HARVARD UNIVERSITY.

LIB RAR Y

OF THE

MUSEUM OF COMPARATIVE ZOOLOGY.

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THE

KANSAS UNIVERSITY
SCIENCE BULLETIN.

DEVOTED TO THE PUBLICATION OF THE RESULTS OF RESEARCH
BY MEMBERS OF THE UNIVERSITY OF KANSAS.

VOL. V.

PUBLISHED BY THE UNIVERSITY,

LAWRENCE, KANSAS,

191L

I

CONTENTS OF VOLUME V.

No. 1.— The Dakota-Permian Contact in Northern Kansas. . . F. C. Greene.
2. — Further Notes on the Pueblo Ruins of Scott County, H. T. Martin.
3.— The Influence of Magnesium Sulphate on the Motor Cells of

the Cerebral Cortex H. F. Hyndman and W. E. Mitchner.

4. —A Study of the Respiratory and Cardiac Activities of the
Skate following Intravenous Injection of Salt Solution. . .

Ida Hyde.

5. — Nutrition of the Embryo Sac and Embryo in Certain Labiatse,

F. H. Billings.

6. — Mitosis in the Root Tips of Podophyllum peltatum, Aute Richards.

7. — Koeberlinia spinosa Zucc. : An Ecological Study of the

Anatomy of the Stem and some other parts. . Miriam Sheldon.

8. —A Case of Absolute Tone Memory Archibald Hogg.

9. —The Hackberry Psylla, Pachypsylla celtidis-mammae Riley:

A Study in Comparative Insect Morphology H. B. Stowe.

10.— Investigations Regarding the Phloem and Food Conduction

in Plants ; Frank U. G. Agrelius.

11.— Histology of Townsendia exscapa and Lesquerella spathulata,

Lillian Bunion.
12. —Cell Changes in the Alveoli of a Carcinoma of the Mamma,

Earl F. Clark.
13.— The Temnospondylous Amphibia and a New Species of Eryops

from the Permian of Oklahoma Roy L. Moodie.

14. —An Armored Dinosaur from the Cretaceous of Wyoming

Roy L. Moodie.

15. —A Contribution to the Soft Anatomy of Cretaceous Fishes,
and a New Promotive Herring-like fish from the Texas
Cretaceous Roy L. Moodie.

16. —A New Species of Holotrich Nadine Nowlin.

17.— Orthoptera from the Santa Rita Mountains, Arizona, collected

by the University of Kansas Expedition Jas. A. G. Rehn.

18.— Observations on the Gryllidge: III. Notes of the Classifica-
tion and Some Habits of Certain Crickets, W. J. Baumgartner.

19.— Observations on the Gryllid^: IV. Copulation, W. J. Baumgartner.

20.— An Examination of the Chromosomes of Anasa tristis

C. E. McClung and Edith Pinney.

21. —Foundations of Arithmetic Arthur Bolles Frisell.

iitr .C 1910

VOL. XI. No. 7.

BULLETIN

OP THE

UNIVERSITY OF KANSAS.

PUBLISHED MONTHLY BY THE UNIVERSITY OP KANSAS.

AT LAWRENCE.

Science Bulletin,

Vol. V, Nos. 1, 2, 3, 4, 5, 6, 7, 8, 9,
10, and 11.

(Continuation of Kansas University Quarterly.)

^I^RIL, 1910.

LAWRENCE, KANSAS.
Entered at the post office as second-claas matter.

1028

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J. N. VAN DER VRIES.

THE

KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 1 — October, 1909.

(Whole Series, Vol. XV, No. 1.)

CONTENTS:

The Dakota-Permian Contact in Northern Kansas, . F. C. Greene.

PUBLISHED BY THE UNIVERSITY,
LAWRENCE, KAN.

Entered at the post-offlce in Lawrence as second-class matter.

THE KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 1] OCTOBER, 1909. [vo'rxv.'NTi

THE DAKOTA-PERMIAN CONTACT IN NORTHERN

KANSAS.

BY F. C. GREENE.
Plates I to IV.

INTRODUCTION.

TOURING the summer of 1906, while working on the Uni-
'-^ versity Geological Survey of Kansas, under the direction
of Dr. J. W. Beede, the Permian-Cretaceous contact in Kansas,
north of the Smoky Hill river, was mapped. The most of the
northern third of the region was mapped jointly with Doctor
Beede and the remainder mapped individually, in alternate
stretches.

The region covered in this account lies in Washington, Riley,
Clay, Dickinson and Ottawa counties, and is included in the
Washington, Marysville, Clay Center, Abilene, and a comer
of the Minneapolis topographic sheets of the U. S. Geological
Survey. The area lies in the drainage basins of the Little
Blue, Republican, Smoky Hill and Solomon rivers.

The topography is characterized by bold bluffs of the Da-
kota sandstone and smaller benches of the harder Permian
limestones and shales. There is one peculiar feature of the
topography throughout the region: northern slopes of the
divides are generally steep, while the southern slopes are very
gradual, and often without rock exposures. The effect is as
if the land had been tipped to the south, or as if there had been
an interruption of a southeastern drainage which turned the
master streams to the east, thereby causing the streams to
erode their southern banks.

In regions of the Dakota formation, this often results in

2 KANSAS UNIVERSITY SCIENCE BULLETIN.

an extensive series of low hills, composed entirely of loose
sand, such as are seen in the vicinity of Abilene.

In the extreme northern part of the state, the country-rock
is overlain by the glacial drift of the Kansan ice sheet. At
the points where the contact crosses the Kansas-Nebraska
line, the original topography and geology are obscured by
drift, 60 to 100 feet in thickness, as observed from well bor-
ings.

GENERAL SECTION.

Pleistocene. — Glacial drift of Kansas sheet and loess (?) ter-
races.

Cretaceous, — Dakota formation, sandstones and clays.
Permian. — Limestones and shales.

Permian. — No attempt was made to distinguish between the
formations of the Permian, but it was necessary to note such
local aspects as would help in determining the contact line.
There is everywhere an unconformity of considerable relief
between the Permian and Cretaceous systems; which repre-
sents in time the Upper Permian, Triassic, Jurassic and Co-
manche Cretaceous.

The relief of this unconformity amounts, in places, to nearly
100 feet in the distance of a quarter of a mile. In one place
(two miles northeast of Emmons, Washington county) there
is a Permian monadnock which has patches of Dakota lying
here and there on its sides, owing to erosion since Dakota
times. This results in exposures of Permian above the level
of the surrounding Dakota. Similar conditions on a smaller
scale are found elsewhere.

Owing to this unconformity, the section of the Permian dif-
fers in places, the Permian having been eroded before the
deposition of the Dakota to a lower horizon in some places
than in others. The result of this is that the Permian is rep-
resented at the contact by different horizons, none of which are
present over very large areas. At most places a series of thin-
bedded limestones and shales was present at the contact, but
in others a very porous limestone, evenly bedded red and
yellow shales, or blue clay shales, represent the horizon of the
Permian present at the contact. Over the whole area the Per-
mian beds appear to be very evenly stratified.

The following section, taken from a cut on the Burlington
railroad about two miles northwest of Hanover, Washington

GREENE: DAKOTA-PERMIAN CONTACT. 3

county, shows a typical succession of the Permian beds of that

region :

10. Limestone, thin-bedded, and yellowish shales 5 ft.

9. Limestone, massive 6 in.

8. Shales, calcareous olive-g:ray, with limestone about four feet

from bottom 16 ft.

7. Limestone, grading through marly concretions to marly

shales 1 ft.

6. Shales, marly gray 4 ft.

5. Shales, red, evenly bedded, with occasional indurated bands;

in the lower portion are two layers of very impure semi-
crystalline limestone or dolomite, apparently thrown down
by precipitation. There are also four or five very thin lay-
ers of greenish shales 15 ft.

4. Covered 10 ft.

3. Limestone, two layers, separated by bluish shale ; upper lime-

stone hard 3i ft.

2. Shales, olive, indurated 4 ft.

1. Shales, yellow calcareous and argillaceous limestone 2 ft.

Covered slope below.

The character of the Permian from a little different horizon
is shown in a section from the north bank of Mill creek, near
the middle of section 13, Charleston township, Washington
county :

7. Covered.

6. Limestone, light buff, cellular, rather thin-bedded 8 ft.

5. Shales, bluish and yellowish 13 ft.

4. Limestone, blue, laminated, fossiliferous 4 in.

3. Shales, bluish 3 ft.

2. Limestone, hard, buff, fossiliferous 8 in.

1. Shale, blue, and covered slope to water level 17 ft.

Cretaceous-Dakota. — The prominent bluffs of this forma-
tion are composed of a deep-red or brown, rather coarse-
grained, ferruginous sandstone. For this reason the forma-
tion is often spoken of as the Dakota sandstone, but this is
a misnomer, as the bulk of the formation is probably not a
hard sandstone, but clays and shales.^ Other exposures vary
greatly. In fact, it is hard to imagine a formation of a more
diversified character. It gives much evidence of being a shal-
low-water deposit, such as would be formed along shores and
in small estuaries by delta deposits, etc. Both the nature of
the deposits and the fossil content bear out this statement.
The land mass was probably a short distance east of the pres-
ent eastern outcrop, as the Dakota becomes more evenly strat-

1. Gaukl. Ti-^nns. Kan. Acad. Sci., XVII, pp. 122-178. 1901.

4 KANSAS UNIVERSITY SCIENCE BULLETIN.

ified to the west. The fossils consist of the leaves of such
genera as Quercus, Sassafras, Salix, Ficus, Protophyllum,
Platanus, Betulites, etc., all land plants.

Near the contact line the diversified nature is well shown.
Yellow, red, brown and white sandstones are found, both con-
solidated and loose. These colors may be in separate layers
or all in the same stone, in a space six inches square. In places
a conglomerate predominates. In Charleston township, Wash-
ington county, about a quarter of a mile west of the line be-
tween sections 23 and 24, there are twenty-five to thirty feet
of variegated, multi-colored clays, such as are typical of the
Dakota a little west of Brookville, Saline county. The local
name for this is "rainbow" clay, all the colors of the rainbow
being jumbled into a variegated mass. There is no consist-
ency in the structure in the easternmost outcrop of the Da-
kota. Within a quarter of a mile a road may cut through
both a deposit of hard dark-red sandstone and a light-colored
clay. In the sandstone cross-bedding is very much in evi-
dence. In places large nodules of iron pyrites are found, while
iron-oxide concretions of various sizes and shapes interest the
inhabitants throughout the region.

The loose Dakota sands wash far down over the Permian
and often obscure the contact.

As the sandstone is very porous, while the limestones and
shales of the Permian are impervious, springs are found along
the contact line and are helpful in locating it.

Pleistocene. — Glacial drift of the Kansan ice sheet. This
formation overlies the five northern rows of townships in the
region mapped. It is as diversified in character as the forma-
tions underlying it. Besides the sands and boulders of quartz-
ite, greenstone, granite, etc., it is composed of fragments of
Permian and Dakota rocks, the latter generally forming the
bulk of the deposits. This is probably due to the fact that the
Dakota formation extends some distance to the east in the
region north of Kansas, thus lying directly in the path of the
Kansan ice sheet. In the region of Hanover (and several
other places) these materials have been cemented together into
a conglomerate. Variegated clays and other clays, resembling
those of the Dakota, have been deposited in places.

This causes some very perplexing problems in locating the
contact in the drift-covered region. In the first place, it ob-

GREENE: DAKOTA-PERMIAN CONTACT. 5

scures the underlying formations over large areas, and in the
second, it contains deposits of clay, sand and conglomerate
almost identical with those of the Dakota. However, if tnere
are any wells present, the water will be hard in the drift and
soft in the Dakota sandstone.

Other Deposits of the Pleistocene. — In much of the region
along the west side of Mill creek in Washington county tnere
are hfteen feet of very sandy, jointed clays, gray above and
brown beneath, overlying the plain sloping towards the creek.
This deposit may be due to the changes which seem to have
been made in tne direction of Mill creek in glacial times. The
topograpny and geology suggest the possibility that Mill creek
once flowed soutneast from Washington, with a large branch
from the north joining it just east of Washington, while a
small stream used the present outlet of Mill creek into the
Little Blue river. There is a channel beneath Greenleaf , nearly
100 feet deep and filled with glacial debris, as shown by the
wells supplying water to Greenleaf. Along the railroad be-
tween Washington and Greenleaf there are no exposures of
rock in place, but as the region was not carefully studied this
evidence is not conclusive. Glacial damming of this stream
in the region southeast of Washington may have formed a
lake which sought an outlet to the north in the present valley
of Mill creek.

If these suggestions are true, it was probably in this lake
that these deposits were formed.

West of the Republican river at Clay Center the river bluffs
are terraces composed of loess-like material with calcareous
concretions, and native rock below. They rise to a height of
forty to sixty feet at the edge of the river bottom, and are higher
here than farther back. There is, as a rule, a fair back slope
of the terrace to the hills of the Dakota to the west, and the
junction is usually occupied by streams or branches of them
and the drainage is away from the river down the terrace,
which has a maximum width of three miles.

It may be that this deposit is the same material as that
which is known as ''plains marl" in the western part of the
state. In regard to the latter, Professor Haworth^ believes it
to have had the same origin as that of the Tertiary mortar

2. Univ. Geol. Surv. of Kan., vol. II, p. 276.

\

6 KANSAS UNIVERSITY SCIENCE BULLETIN.

beds, and thinks it probable that many of the properties of the
plains marl are largely due to the action of wind.

In the region mapped the loess was found in valleys having
a north-and-south direction. This may be due to the fact that
a wind from the west, carrying the material, would deposit it
in valleys lying north and south, while it would not accumulate
in valleys having an east-and-west direction.

DISCUSSION OF THE MAPPING.

The mapping began at the Kansas-Nebraska line. Here the
contact was concealed by 60 to 100 feet of drift. Just west of
the Little Blue river, opposite Hollenberg, the Permian was
exposed near the 1300-foot contour, while east of this town the
Dakota rocks were exposed near the same level. Thus, in a
general way, the contact is at the 1300-foot level between the
state line and Washington west of the Little Blue. On the
west bank of Mill creek there is a deposit of Pleistocene, cover-
ing the slope. The region is mapped as Cretaceous rock,
though there is little doubt that it is material washed from
the higher hills to the west. A tongue of Dakota extends north
in the area between Mill creek and this river. If the sugges-
tion that Mill creek formerly had its outlet to the south is true,
this tongue would probably be an outlier.

On the east side of the Little Blue are outliers within six
miles north and south of Hanover, but inside the boundaries
of Washington county. South of Washington the contact rises
nearly to the 1400-foot line, but west of that place pitches
down, so that the Permian extends up Mill creek only two
miles. The contact is found in the valley of Beaver creek, at
an elevation of about 1375 feet. At Greenleaf, owing to the
presence of the old channel noted above, neither Permian nor
Dakota is exposed, but the former is only thirty feet below
the level of the town a few miles east and west. Across this
area the contact is dotted.

From Greenleaf to Clifton there is a continuous dip to the
west. There is an outlier of Dakota in the vicinity of Chep-
stow, and it also extends south into the extreme northwest
corner of Riley county, south of Kimeo. West from this point
to Clifton the contact lies mostly in the northern row of town-
ships of Clay county.

In the Republican valley the contact is concealed by a de-

GREENE: DAKOTA-PERMIAN CX)NTACT. 7

posit of loess-like terrace material and alluvium, but a few
miles east of Clifton it is found at the 1280-foot contour, and
probably crosses the river about a mile southeast of Clifton.
West of the Republican river the Dakota is found as a thin
sheet on the terrace, as shown by well borings. Here the con-
tact is at about the 1240-foot mark, but rises steadily to the
south until the divide south of Manchester is reached, where a
height of 1360 feet is reached. There is, however, a constant
westward dip, so that the contact never extends very far west
in the valleys of eastward flowing streams. In the region just
described the contact passes near Idana, Oak Hill, Longford
and Manchester.

East of the Republican river, in Clay county, the Dakota
forms a large outlier about three miles northwest of Green.
Here the contact is nearly up to the 1400-foot contour. From
this outlier west to the Republican river the dip of the Per-
mian is clearly and distinctly shown.

From Manchester to Abilene the Dakota lies on a narrow
divide, but at its southern extension expands into a large
area. In this area it is very thin, and disappears in a large
expansion of sand hills, which may have originated from
poorly lithified sandstone, or from stream or wind sediments.

From Abilene east to within three miles of Chapman, sand
and sand hills occur on the north side of the Smoky Hill.
There is much doubt as to the origin of these deposits. In this
connection, the observations of Hay^ in the vicinity of Junction
City are interesting. He found sand hills near Junction City
at or about the same level as those in the vicinity of Abilene.
He says: "A more recent examination of the high-level sand
dunes previously referred to revealed the fact that they are
residual beds, resting on and abutting against undoubted out-
liers of the Dakota." Two areas of Dakota are shown on his
map. These are only one-half of a mile west of the eastern-
most outcrops in Washington county, although the trend of
the eastern outcrop of the Dakota is southwest throughout the
state.

3. Hay. Geol. of Ft. Riley, etc.. Bull. 137. U. S. G. S.. p. 28.

2-Univ. Sei. Bull., Vol. V. No. 1.

8 KANSAS UNIVERSITY SCIENCE BULLETIN.

CONCLUSIONS.

There is an unconformity of varying, but considerable, re-
lief between the Permian and Cretaceous systems.

The eastern part of the Dakota sediments was deposited in
shallow water along the shore and in estuaries and lagoons, as
shown by the nature of the formation and the fossil content.
Some of them may be of sub-serial origin.

The shore-line of the Cretaceous sea was probably some dis-
tance to the east of the main body of the Dakota outcrop, as
is shown by the presence of outcrops east of Hanover, Wash-
ington county, and in the vicinity of Junction City.

THE

KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 2— October, 1909.

(Whole Series. Vol. XV, No. 2.)

CONTENTS:

Further Notes on the Pueblo Ruins of Scott County, H. T. Martin.

PUBLISHED BY THE UNIVERSITY,
LAWRENCE, KAN.

Entered at the post-oflBce in Lawrence as second-class nuittcr.

THE KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 2] OCTOBER, 1909. [^o^ ^v.^nTs

FURTHER NOTES ON THE PUEBLO RUINS OF SCOTT

COUNTY.

BY H. T. MARTIN.

(Contribution from the Zoological Laboratory, No. 188.)

Plates V to IX, and one text figure.

A T the annual meeting of the Kansas State Historical Society
-^ at Topeka in January, 1899, Dr. S. W. Williston read be-
fore that body a paper entitled "Some Pueblo Ruins in Scott
County, Kansas," in which was fully set forth the results of
the excavation of the ruins now known as Quartelejo, together
with a list of the implements and weapons procured from the
rooms which composed the old buildmg. This paper will be
found in the Kansas Historical Collections, volume 6, 1897-
1900. Unfortunately it was not then possible to publish along
with the paper the photographs and drawings of the ruins
which were made during the excavation. In order to make the
photographs and ground-plan drawing which accompany this
paper as comprehensible as possible, the original description
by Doctor Williston will be given almost verbatim. Since this
account was based upon the field notes of the present writer,
who made the excavations under the directions of Dr. Will-
iston, its repetition here will be somewhat in the nature of a
personal account.

DOCTOR WILLISTON'S ACCOUNT OF THE RUINS.

"The ruins are situated in the valley of Beaver creek
(wrongly called Ladder creek on the map), in the northern
part of Scott county, twelve miles due north from Scott City,
and about ten miles south of the Smoky Hill river, as shown
on the maps, precisely where the township line touches the

(11)

12 KANSAS UNIVERSITY SCIENCE BULLETIN.

most eastern bend of the creek. At this place the valley of the
creek, which here runs nearly north, is less than a mile wide,
surmounted on either side by high bluffs of Tertiary material.
The immediate valley is excavated in the Cretaceous chalk.
The result is that here, as elsewhere in western Kansas where
like geological conditions obtain, the underflow through the
Tertiary sandstones, over the impervious chalk floor, comes
abundantly into the valley, furnishing a considerable stream
of water. Perhaps no stream in the western part of the state
offers more favorable conditions for irrigation than does this
in its lower part. In the dryest years there is always an
abundance of water in the stream, and in the deep pools along
its course there are always many fish. About a half mile above
the site of the present ruins, the Tertiary underflow comes to
the surface along the side of a hill in such perpetual abundance
that it is utilized in the irrigation of a considerable tract of
land.

"These two facts — easy facilities for unfailing and exten-
sive irrigation, and a fish- and beaver-producing, perpetually
flowing stream — are undoubtedly explanatory of the location
of the ruins at this place. The ruins are situated near the
middle of the valley, close to the stream, and away from any
possibility of ambush by hostile savages. They occupy a small
knoll of ground, and, as first seen by us, consisted of a low,
rounded heap of soil and stone, perhaps 75 or 100 feet in diam-
eter, the soil wholly overgrown by buffalo grass. The rocks
are the coarse sandstone of the neighboring hills. A small
excavation had been made in the middle of this mound by pre-
vious explorers, perhaps two feet in depth and of a dozen
square feet in area."

In conclusion. Doctor Williston says: "One fact is estab-
lished from the explorations — the ruins are of Pueblo origin.
Of this there can be no question. The plan of the structure is
only such as the Pueblo Indians could have devised and carried
out. It is not the work of white men, either Spanish or
French, though it is very probable that both the Spanish and
French may have occupied this and other structures at this
locality at later times, or even contemporaneously with the
Pueblos. The finding of an iron ax, of rude and primitive
workmanship, it is true, indicates white men's skill. It is very
evident, also, that the metal instruments were used by the

martin: some pueblo ruins. 13

occupants. Several of the manufactured articles show clearly
the imprints of metal saw teeth.

"The origin of the ruins is of course not positively proven,
yet I believe that concerning this there is scarcely a doubt that
they represent the old fortified place known as Quartelejo,
founded about 1650 by a party of Indians who fled from the
oppression of the Spaniards, from Taos, in New Mexico. The
only information concerning this place that I have so far been
able to obtain is from the works of Hubert Bancroft, volume
17, on Arizona and New Mexico."

The following is the full description of the ruins as published
in the Kansas Historical Collections by the present writer :

"In the excavation of the chief structure referred to in the
cited paper, all possible care was taken to avoid mutilating the
plastering with which the walls were covered, thus permitting
the exact size and shape of each room to be ascertained. As
now excavated, the walls are about two and a half feet in
height. The structure measures fifty by thirty-two feet in
size, and stood as nearly due east and west in its greater meas-
urements as it would be possible to locate it with an ordinary
compass. The outer walls were of heavy stone, from eighteen
inches to two feet in thickness, and were cemented or grouted
together, making the full measurement of the building about
fifty-three by thirty-five feet. The building site, as has been
described by Doctor Williston, was a slightly raised mound,
about seventy-five yards from the bed of Beaver creek, which
here affords an abundance of water for both irrigational and
domestic use. By the side of the building there are two large,
hollowed out places, which had probably been used for the
puddling and mixing of the adobe employed in the construc-
tion of the building. The stone used in the building, all of
which had been brought from the surrounding hills, was con-
siderable in amount, and many single pieces are all that a man
can lift.

"About 100 yards south of the main edifice there is evidence
of several other smaller buildings, all of which must have been
constructed of adobe alone, since no rock remains. These
smaller structures, two of which were examined by us, yielded
no utensils or other relics; nor could their size and shape be
made out with certainty. Both of these buildings, as well as
the large one, present evidence of having been destroyed by fire,
whether as the result of some accident or by Indian foes one

14

KANSAS UNIVERSITY SCIENCE BULLETIN.

martin: some pueblo ruins. 15

cannot say, of course. From the fact that no human bones
were found anywhere about, the probability of design is less-
ened. That the larger structure had been destroyed by fire
there can be no doubt, since the adobe is burnt, and charcoal
is thickly scattered everywhere; the stone and bone imple-
ments also all show evidence of fire. Rooms IV and VII, as
I have designated them, show only slight evidence of the fire,
and it is possible that one or both of these rooms had never
been covered, and hence contained but little subject to de-
struction by fire. Room IV had portions of the rotted posts,
evidently used as a ladder, remaining.

"Charred corn was found in every room except VII, in some
places four or five inches deep. In room V there had been four
or five bushels of this corn in a slightly hollowed out place at
one side.

"About twenty-five yards north of the main structure there
appears to have been three or four small structures, each sepa-
rated a small distance in an east and west line parallel with
the main building. These structures were apparently circular
in outline, and were perhaps teepes.

"The most interesting room in the structure is the one I will
designate as room I. Its dimensions were seventeen feet by
thirteen feet and nine inches. It had a raised dais or platform
on two sides, about six inches high; that on the west side five
feet and three inches wide; that on the north side two feet.
The wider one was doubtless used for sleeping purposes, and
the narrow one as a bench. Very near the center of the room
there is a box-like receptacle, formed of thin stone set edge-
wise. Like the others described further on, the bottom of this
one was about six inches below the level of the floor, and its
size was eighteen by twenty-one inches. It had been plastered
at the bottom, and contained, when examined, a quantity of
clean wood ashes. The receptacle may have been for the grind-
ing and mixing of corn. In the southwest corner of the room
there is a peculiar structure three feet nine inches in length by
two feet and one inch in width, inside measurements, built of
adobe. Its walls are eighteen inches high at the west end and
twelve at the east end, the slope gradual from one end to the
other. The walls, five or six inches in thickness, had been
nicely rounded at the top. In the middle, and joined to the
west end, is a small platform, about sixteen inches in length
by twelve in width, raised about six inches above the bottom

16 KANSAS UNIVERSITY SCIENCE BULLETIN.

of the grooves which surround it. These grooves, shaped some-
what like a U, slope toward the closed end. This part was
filled with ashes, suggesting that the use of the oven was for
the baking of pottery. Near the east end was a large hole,
twelve inches in diameter and eighteen inches in depth, cov-
ered with a flat rock. It contained nothing save fine dust.

"The walls and floors were nicely plastered. The plastering
gave no indications of finger marks, but seemed to have been
smoothed off with some instrument. Stones that might hav^'
answered such uses were found in the rooms. In this room
was found a small pipe, decorated with horizontal markings.
Here also were found a needle or awl for the sewing of hides,
several arrow-heads, fragments of pottery, and bone needles.
The remains of two posts, about eighteen inches apart, were
found in the northeast corner, evidently for the uprights of a
ladder for ingress and egress. Similar holes in like positions
were found in the other rooms. There were no indications of
doors or other openings in any of the rooms. The roof was evi-
dently made of willow poles or brush covered with adobe, as
large quantities of the latter show impressions of twigs.

"Room II was sixteen feet and four inches in width by eigh-
teen feet six inches in length, and had both wall and floor
plastered. The fireplace was two feet by one foot seven inches
in size, and close by it was a hole twelve inches in diameter.
On the east end there was a bench, as in room I, four feet two
inches in width, and on the north side one two feet in width,
while on the other two sides the width was but twelve inches,
but raised to about ten inches in height. Close to the fireplace
was found a grooved stone maul, ribs with marks of a saw
upon them, arrow points, pottery, bone and stone scrapers, and
a small pipe. On the ledge at the east end was found the half
of an iron ax or wedge. The iron of course is much rusted,
and the tool appears to have been split longitudinally and
transversely by some mishap. It had a groove near the head,
instead of an eye, for the attachment of the handle, after the
manner of the stone axes of the aborigines. This room con-
tained more charred corn than did any of the others.

"Room III was fourteen by thirteen feet in size, with plas-
tered walls and floor, the corners rounded at the east end
and square at the west. It had a fireplace eighteen inches by
twenty-four, and a raised dais four feet wide at the west end.
The holes for the posts supporting the roof and for the ladder

martin: some pueblo ruins. 17

were as in the other rooms. The plastering turned up about
the posts showed that this work had been done after the roof
had been placed over the structure. This room furnished
grinders and several bone implements — scrapers and fleshers
— made from the shoulder-blade of the buffalo and deer and
antelope. The wall posts were rotted in the ground, and not
burnt as in the other rooms, nor were the bone implements
partly burned, as was the case with those in the other rooms.
In this room, also, was found part of a musical instrument, a
flute or flageolet, made from the wing bone of a large bird;
also a bone implement with a serrated edge,

"In room IV fewer implements were found than was to be
expected, for few marks of fire were visible. Owing to this,
the room was in better condition for articles to have been pre-
served. It is just possible that the fire started on the south
side, giving the occupants a chance to save most of their be-
longings from this room. Several scrapers, knives, also the
small arrow points on plate I, Nos. 1, 2, 4, 5 and 12, 16, 17, 18,
19, 20, all of which are fine examples of workmanship, were
found.

"Room V was the smallest in the building, being only ten by
fourteen feet in size. It had well-plastered walls and floor, a
fireplace seventeen by twenty-two inches in size, a large quan-
tity of corn, arrow-heads, grinders, scrapers, pottery, etc.
Close by the fireplace there was a hole in the floor covered by a
flat stone that had been undisturbed. At its bottom was found
half of a clam shell, which had been sawed lengthwise with a
toothed saw, the tooth marks being very plainly apparent. In
the northwest corner was a small oven, nine inches in width
and sixteen in depth, excavated from the wall of the room and
plastered throughout. It contained three or four inches of
wood ashes in the bottom. In this were also found three oval
and one square adobe bricks, about ten by fourteen inches in
size, flattened above and rounded below. They may have been
used in the baking of tortillas.

"Room VI was ten feet five inches in width by thirteen feet
and eight inches in length. The level floor had been plastered,
as also the fireplace, which was eighteen by twenty-six inches
in size. There was a narrow partition between this room and
room V, and since no indications of a ladder were found here
it is possible that the two rooms had been connected. Several

3-Univ. Sci. Bull.. Vol. V. No. 2.

18 KANSAS UNIVERSITY SCIENCE BULLETIN.

scrapers, of bone and flint, together with grinders, etc., were
found here.

"Room VII, thirteen feet square, differed from all others in
having no fireplace or plastered walls. Numerous bones of
bison, deer, antelope, coyote, badger, etc., were discovered in
this room. The only relics were bone and flint scrapers. Prob-
ably the room had been used as a sort of storehouse, and not
for human dwelling.

"The pottery found was in part composed of plaster of Paris,
possibly obtained from the crystals of selenite scattered over
the chalk exposures in the vicinity. A number of ribs were
found which had been smoothed at one end into a sort of spat-
ula, and had probably been used in the making of pottery or in
the plastering of the building. Coiled as well as smooth pot-
tery was found, but only a single piece that showed evidence
of decoration. Some of this pottery has been submitted to
Professor Hewitt, of Las Vegas, N. M., who has given much
attention to the work of the Pueblo Indians. He was of the
opinion that all this pottery had been introduced from New
Mexico, and had not been made in the vicinity of the building
or village. Probably this is the furthest east that such pottery
has yet been found. In one of the rooms were found several
squash seeds ; some between two pieces of pottery, in good con-
dition, others much decayed."

In the above account by Doctor Williston, he very graph-
ically described the locality, surroundings and natural re-
sources in the immediate vicinity of the ruins, which he had
visited in 1896, but unfortunately was unable then thoroughly
to investigate. He also gave a number of important quota-
tions from the works of Hubert Bancroft, an eminent writer
on Arizona and New Mexico. These works treat largely of the
early history of the Spanish in America, and contain important
data relative to the position of the pueblo known as Quartelejo,
and the condition under which this isolated post was first built
and peopled. From the above-named works it appears that
about 1650 a band of Taos Indians fled to the plains and for-
tified a place called Quartelejo. In the papers of the Archaeo-
logical Institute of America, American series V, Hemenway
Archaeological Expedition, A. F. Bandelier, page 181, says:
"In 1704 the Indians of the pueblo of the Picuries abandoned
their village in northern New Mexico and fled to the plains,
where they established themselves at a place called El Quar-

martin: some pueblo ruins. 19

telejo. Superstition was, from all indications, the cause of this
hasty movement." In a foot note he adds: ''This flight of the
Indians of the Picuries to the Quartelejo is mentioned in sev-
eral documents. I quote here only the witchcraft trial, entitled
'Causa Criminal co7itra Gevonimo Dirucaca, Indio del Pueblo
de Picuris,' 1713, MS., fol. 16, 17. The same document also
fixes the date. In regard to the return from Quartelejo, it was
effected in 1706 by Sargente Mayor Juan de Uribarri."

For some reason or other there appears to be a conflict as to
the exact date of the settlement of the Indians at Quartelejo,
yet both the above writers agree that the return to Mexico was
effected in 1706. This would indicate clearly that both Ban-
croft and Bandelier were writing about the movements of one
and the same band of Indians. Furthermore, it does not ap-
pear credible that the governor of New Spain would have al-
lowed these discontented Indians to have stayed at Quartelejo
from 1650 until 1704, a period extending over fifty years, with-
out an effort being made to effect their return, and in all prob-
ability Bandelier's date of the founding of this isolated pueblo
in 1704 is the more correct.

He further states that "in all probabilities the place got its
name from the fact that these Picuries Indians made a tempo-
rary stay there." This is also intimated by Fray Silvestre
Valez de Escalante in his letter to Father Agustin Morfi, April
2, 1778. After the evacuation of the pueblo in 1706 at the in-
stance of Uribarri, another band of friendly Indians must have
taken possession of and occupied the place for a number of
years, for from Bandelier we learn that in 1719 Governor
Don Antoni Valverde Cossio, having determined upon a cam-
paign into the north and northeastern plains (a movement
made chiefly against the Yutes and such of the Comanches as
had joined them), penetrated as far as the Quartelejo, and
even beyond. A part of Valverde's object was to secure the
rancherias of the Jacarrilas against their foes. Part of these
friendly Apaches lived at Jacarrila, forty leagues north of
Santa Fe, and another band lived at Quartelejo. Again, we
find that in the following year, 1720, Don Pedro de Villazur
called at this pueblo on his ill-fated trip to the Pawnee viflage
which is supposed to have been located on the Platte river,
when all perished at the hands of the treacherous Indians.
After this I have been unable to find the place mentioned by
any other writers of early southwestern history. And it is

20 KANSAS UNIVERSITY SCIENCE BULLETIN.

more than probable that after the Pawnees had annihilated
Villazur and his soldiers they also, to check the inroads and
settlement of the Spanish and their allies, destroyed the Quar-
telejo, thus effectually clearing the whole country north of the
Arkansas river. This would admirably account for the entire
pueblo and settlement being destroyed by fire, as it undoubt-
edly was. From the position of the utensils and implements
found, a very hurried departure must have been made from
the main structure. Most of the meal-grinding apparatus
found in each room appeared to be left in a group, as though
just laid aside after use. This, and the finding of so much
charred corn in most of the rooms, would surely indicate that
no prearranged plan of evacuation had been decided upon.
That the ruins are those of the Quartelejo mentioned by the old
Spanish writers, to which conclusion Doctor Williston came
after seeing the group of relics obtained, there can be no doubt.

Dr. F. W. Hodge, of the Bureau of Ethnology, says, in the
American Anthropologist, volume 2, October and December,
1900, that he regards the discovery of the ruins as of great in-
terest, that they are of typical pueblo fashion, and, in conclu-
sion, says : ''The former existence of this pueblo has been
known to students of southwestern history and ethnology for a
number of years, but not until the publication of the results of
the investigation had its situation been known, and there is no
reasonable doubt that the identification is correct, for pueblo
architecture is intrusive in Kansas."

If the identification by Doctor Williston as Quartelejo is cor-
rect, then the exact locality of this pueblo, so often referred to
by the old Spanish writers, and to which Don Juan de Archu-
leta journeyed, is assured, and as such should be of great value
in assisting to prove more definitely that the localities desig-
nated by Brower and Hodge as the site of Quivera and Hara-
hey are correct, and thus have some bearing as to the probable
line of march of Coronado, and help to prove that he did cross
the Arkansas river.

Brower says, in his "Harahey," page 27, volume 2 : "When
the final acceptance shall have been established that Coronado
reached and crossed the Arkansas river and followed its course
to the great bend, then the true site of Quivira and Harahey
will have been finally reached and comprehensively under-
stood."

By referring to a quotation given by Doctor Williston in his

MARTIN: SOME PUEBLO RUINS. 21

paper taken from the narrative of Fray Silvestie Velez de
Escalante, April 2, 1778, translated "In the Land of Sunshine,"
we find that Don Archuleta found in the possession of the
Indians of Quartelejo pieces of copper and tin, and that they
said they had procured them from the Quivera pueblos, to
which they had journeyed from Quartelejo.

If these Indians had visited Quivira as they claimed, and
Quivira and Harahey was where Brower and Hodge have lo-
cated it, northeast of Quartelejo, then the presence in the ruins
of the sandstone arrow straighteners can be satisfactorily ex-
plained, and the possession of knives and arrow heads of the
Harahey type and material does tend to prove they had com-
municated in some way with the Quivira settlement. The
arrow straighteners referred to, and which were found in the
ruins, are made of the Dakota Cretaceous sandstone.

The writer has found many arrow straighteners on the west-
em Kansas prairie made by the plains Indians, within a few
miles of the ruins, but these were all of a different and harder
material. Dakota sandstone in any shape is intrusive in the
neighborhood of the ruins, but a journey to Quivira by the
Indians of the Quartelejo would take them directly through
the outcroppings of Dakota sandstone which occur in Ells-
worth, Barton and Rice counties, and enable them to procure
and carry back with them this valuable acquisition.

In a previous paper the writer made mention of a peculiar
U-shaped structure that looked like it might have been used
for the baking of pottery, on the raised platform-like center
piece, a photograph of which is shown on plate VI, fig. 3.
This, Dr. F. W. Hodge, in the American Anthropologist of
October and December, 1900, says is a typical pueblo grinding
trough, and on the raised platform in the center was undoubt-
edly fixed the grinding stone.

The small fireplace and chimney built into the wall of room
V is due to the influence of Spanish associations, for at this
early date chimneys were unknown to the Pueblo Indians
of New Mexico. The chimney leading to the roof was carried
up the center of the wall between rooms V and VI, and was
plastered inside, provision being made to allow for room for
this by a thickening of the wall at this point. It is of course
not known of what material the inner walls were composed,
but from the amount of rock that had been removed from the
ruins for building purposes by settlers the past twenty years,

3-Univ. Sci. Bull., Vol. V. No. 2.

22 KANSAS UNIVERSITY SCIENCE BULLETIN.

and the amount remaining at the time of the excavations, I
think it is safe to presume that the whole structure, with one
exception, was composed of rock. The wall separating rooms

V and VI was entirely too thin to allow of being built of stone,
and was in all probability made of willows woven together and
plastered over. The division wall extended clear across the
room to a height of eighteen inches, and although no opening
for communication showed at this height, it is more than pos-
sible that these two rooms were connected by some opening in
the division wall higher up, considering the fact that in room

VI no post holes remained showing any indications of their
having had a ladder for communication with the roof.

It is much to be regretted that out of the vast amount of
material that has been collected in and around the 100 or more
village sites enumerated in Brower's Harahey, not more than
a dozen specimens have been secured for the University collec-
tion of archaeological material. Yet Brower states that more
than 10,000 archaeological specimens have been gathered, and
are deposited in the museum of the State Historical Society of
Minnesota.

As a means of advancing the study of the earliest history
of the state, and as an inducement to students interested in
archaeology and ethnology, scarcely could there be anything
more valuable and important than a well labeled, systemat-
ically arranged series of archaeological specimens collected
from these historical old village sites of Quivira and Harahey,
visited so long ago by Coronado and his followers.

I here wish to convey my thanks to Dr. C. E. McClung for
his assistance in the preparation of this paper, and for his
kindly advice and corrections.

In the near future a model of these interesting ruins, the
first ever inhabited by white man in the state of Kansas, will
be constructed and placed in the University museum.

BIBLIOGRAPHY.

J. V. Brower. Harahey, vol. 2.

A. F. Bandelier. Archaeological Institute of America, American series
V, Hemenway expedition.

Dr. F. W. Hodge. American Anthropologist, vol. 2, October and De-
cember, 1900.

Dr. S. W. Williston and H. T. Martin. Kansas Historical Collections,
vol. 6, 1897-1900.

THE

KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 3— October, 1909.

(Whole Series, Vol. XV, No. 3.)

CONTENTS:

The Influence of Magnesium Sulphate on the Motor Cells of
THE Cerebral Cortex, . . H. F. Hyndman and W. E. Michener.

PUBLISHED BY THE UNIVERSITY,
LAWRENCE, KAN.

Butered at the post-offlce In Lawrense as second-class matter.

THE KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 3] OCTOBER, 1909. ['^"^xvf nTs

THE INFLUENCE OF MAGNESIUM SULPHATE ON THE
MOTOR CELLS OF THE CEREBRAL CORTEX.

BY H. F. HYNDMAN AND W. E. MICHENER.
(Contribution from the Physiological Laboratory of the University of Kansas.)

'T^'^HE experiments of which the results are here briefly re-
^ corded were undertaken for the purpose of ascertaining
whether magnesium sulphate paralyzed the motor cells of the
cerebral cortex. It was found that 1.5 grams of magnesium
sulphate per kilo, when injected in rabbits subcutaneously, pro-
duced anaesthesia in about fifty minutes that lasted two hours.

Rabbits were employed in all of the tests and three series of
experiments were carried out.

In the first set ether was given, and when anaesthesia had
developed the motor areas were exposed, and a definite, well-
defined small area, which when stimulated with insulated
platinum electrodes with a determined minimal strength of
induced current, caused the hind leg of the side opposite to the
one stimulated to give a definite contraction. The cortex was
then carefully removed and again the minimal strength of cur-
rent determined that would produce the same extent of con-
traction as before in the same leg. The animal was then
allowed to come from under the influence of ether, and then an
anaesthetic dose of magnesium sulphate was injected. The
same area on the opposite side was now stimulated with a de-
termined minimal strength of current that would produce the
same extent of contraction as was secured with ether. The
cortex was then removed and again the minimal strength of
current determined.

The experiment was also modified in that the minimal

(25)

26 KANSAS UNIVERSITY SCIENCE BULLETIN.

strength of current for contraction was determined after cortex
stimulation, and then, when the animal recovered from the
ether effect, magnesium sulphate anaesthesia was secured.
Again the cortex was stimulated with a minimal strength of
induced current.

The results of this set of experiments were that it required
exactly the same strength of current to produce the same extent
of contractions, both with the cortex intact as well as when the
motor cells were removed and the fibers directly stimulated
under ether as under magnesium sulphate anaesthesia. In each
case the strength of current required for the same contraction
was greater when the motor fibers were directly stimulated
after the motor cells were removed than when the contractions
were secured indirectly by direct stimulation of the motor cells.

The second set of experiments differed from the first set in
that the same procedure was carried out under magnesium
sulphate anaesthesia alone.

The third set of experiments consisted in determining the
strength of current when applied to the central ends of the
eighth and seventh cervical dorsal spinal nerves under ether
and magnesium sulphate anaesthesia that would reflexly cause
contraction of certain muscles of the mouth.

A comparison of all the experiments showed that the same
minimal strength of current was required to produce the same
extent of contraction under ether as under magnesium sulphate
anaesthesia, whether applied directly to the motor fibers, indi-
rectly to the fibers through the motor cortical cells, or when
applied to afferent fibers that secured a reflex response; and
that a greater strength of current was required when applied
directly to the motor fibers than when the impulses reached
these through stimulation of the motor cortical cells.

We conclude, therefore, that magnesium sulphate anaesthesia
does not paralyze the motor cells of the cerebral cortex in the
rabbit.

We take this opportunity to thank Doctor Hyde, under whose
supervision these experiments were conducted, for her many
kind suggestions.

THE

KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 4— October, 1909.

(Whole Series, Vol. XV. No. 4.)

CONTENTS:

A Study of the Respiratory and Cardiac Activities and Blood
Pressure in the Skate Following Intravenous Injections
OF Salt Solutions, Ida H. Hyde.

PUBLISHED BY THE UNIVERSITY,
LAWRENCE, KAN.

lOutfifil ;ii I lie [>ost-offlep iu l.n\vren«K' as second-class matter.

THE KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 4] OCTOBER, 1909. [^orxV™:

A STUDY OF THE RESPIRATORY AND CARDIAC AC-
TIVITIES AND BLOOD PRESSURE IN THE SKATE
FOLLOWING INTRAVENOUS INJECTIONS OF SALT
SOLUTIONS.

BY IDA H. HYDE.

Plate X, and 49 figures.

(From the Marine Biological Laboratories of Woods Hole and Leland Stanford' and the
Physiological Laboratory of the University of Kansas. )

' I '^HE experiments that are recorded in this article were con-
■ ducted on the skate, Raia ennacea and Raia binoculata,
during the summers of 1902, 1903, 1906 and 1907.

A host of articles have accumulated since Humboldt first
studied the action of salt solutions upon the rhythmical changes
in the heart's contractions; but with the progress of physical
chemistry new problems and different views presented them-
selves and left many important questions still unsettled. To
aid in solving some of these I undertook the study from a less
general standpoint by employing a method and an animal sel-
dom heretofore adopted in the solution of the questions. A
few experiments were begun on the fresh-water sturgeon and
on the frog in order to compare the results with those obtained
with the skate. These, however, will be extended and varied
before the results are published.

It was hoped that simultaneous records and comparisons of
the three activities might throw some light upon the factors
that influence them and indicate whether their nerve centers
are alike stimulated by the same ions. At first the object was
also to determine whether it was the anion or cathion that

1. I take this opportunity to express my hearty thanks to the directors for the priv-
ilegre of working in these institutions and for the courtesies I receive<l there.

(29)

30 KANSAS UNIVERSITY SCIENCE BULLETIN.

stimulated the different centers, and if there were ions that
simulated one but proved neutral or inhibitory to the other
centers. Moreover, whether valency or solution tension were
influential factors, and if the toxicity produced on either the
respiratory, cardiac or vasomotor centers by certain salts could
be counteracted by definite amounts of other salts that proved
antitoxic. Before I had proceeded far, however, I desired, in
addition to the solution of some of these questions, to study
other physical and chemical effects that were produced in the
body by the salts; e. g., osmotic pressure, the products of chem-
ical reactions, such as oxygen, carbon dioxide, hydrogen and
hydroxyl ions, and the dosage of the solutions,

I realize that in experiments of this character a host of phe-
nomena may exert their influence and that it is difficult with
the data at hand to assign to any special one the cause that
produces the particular result. Some of the events which may
deserve consideration can, however, be controlled and their in-
fluence either accounted for or removed from further consid-
eration. That is especially so in the investigation of the fishes,
where the temperature and supply of the water can be con-
trolled and are kept normal and constant throughout the ex-
periment and where the amount of solution injected bears a
direct proportion to the weight of the animal, and also where
from the method employed mechanical stirnulation, either due
to the insertion of the cannula or brought about by retaining
the fish in a definite position during the experiment, is avoided.

On the other hand, we assume that the addition of an isotonic
salt solution to the blood, the chosen amount of which does not
cause any change in any of the functions under consideration,
will, if hypo- or hyper-tonic, produce various physical and
chemical alterations both in the fluids and tissues of the body.
Introducing an additional salt into the blood assumes changes
in its osmotic pressure and in the contents of the corpuscles,
also alterations in the permeability of the walls of the capilla-
ries, corpuscles and other tissue cells. Or the new composition
in blood salts may make new combinations with colloids or
ferments or proferments, or set free ions that will thus stimu-
late or inhibit functions by acting directly on cardiac or muscle
cells, or indirectly upon special centers. That is, the physio-
logical property of the blood is altered by changing the propor-
tion and character of the electrolytes in it, and this must have

HYDE: EXPERIMENTS ON THE SKATE. 31

its peculiar influence upon the life phenomena of respiration
and heart action and blood pressure. To which of these must
be ascribed the important changes produced in respiration and
cardiac activity and blood pressure is a question with which T
shall not for the present concern myself, since in addition to
the experiments which I conducted it would be necessary, as 1
later saw, to carry on extensive physical and chemical investi
gations for which I lacked the time. I shall content myself,
therefore, by simply stating my method and observations, and
not attempt theoretical considerations of the underlying
causes, trusting that the data obtained from over 200 experi-
ments will aid in the correct interpretation of results secured
by other investigators working upon problems related to those
that I undertook.

According to Garry- and Sumner'^ the percentage of salts in
the skate and selachian's blood does not differ from that in
sea water. The lowering of the freezing point being — 1.8° C.
and the osmotic pressure twenty-two atmospheres, the sea
water is therefore isotonic with the skate's blood, while Bag-
lioni^ estimated it isotonic with an m% NaCl or a 2 per cent
NaCl plus 2,5 per cent urea. Although the osmotic pressure
of the selachian's blood is the same as sea water, its salt con-
tent is less, but the lowering of the freezing point is compen-
sated by the urea in the blood.

Employing Welcker's method, I ascertained that a small
skate weighing from 550 to 700 grams possessed 25 to 30 grams
of blood, or about one-twentieth of the body weight. Harris''
states that the weight of the blood in the skate is only one-
fortieth of the body weight, but he does not describe the method
employed for ascertaining his data.

Although there are wide variations in blood pressure, I
found that the mean pressure for an average-sized skate was
about 20 mm. of mercury.

The photograph on plate X illustrates one of the methods
employed for securing records of the respiratory and cardiac
action and blood pressure. The skate was placed ventral side
up on an inclined board, partly submerged in water, and held
in place by fish netting; a constant stream of fresh sea water

2. Garry, W. E. Biological Bulletin, 1905, vol. VIII, No. 4, p. 257.

3. Sumner, F. B. Bulletin Bureau of Fisheries, 1906, p. 596. Baglioni. Zentralblatt
fiir Physiologic, 1905, vol. XIX, p. 12. Harris. D. F. .Journal of Physiology, 1903,
vol. XXX, Nos. 3 and 4, p. 319.

32 KANSAS UNIVERSITY SCIENCE BULLETIN.

of a definite volume, velocity, and temperature entered the
mouth through a forked glass tube covered with rubber tubing.

As soon as the water enters the mouth and flows out of the
gill slits over the body, thus keeping the skate practically sub-
merged, and the netting comes in contact with the body, the
fish is perfectly quiet and remains so for hours, apparently
contented and comfortable. A cannula, filled with 20 per cent
glucose or glucose and 1 per cent ammonium oxalate, is in-
serted in one of the side branches of the aorta and joined to a
delicate mercury manometer placed quite on a level with the
blood vessels. To the open end of the manometer is attached
a Hurthle manometer, which records the heart action, while
the pressure is read oflF directly from the manometer.

By means of a silk loop one of the gill arches is attached to a
light lever which records the respiratory movements. In con-
trol experiments the tip of the ventricle was exposed and by
means of a silk loop joined to a light lever, so that in addition
to the above records, separate ones were obtained of the rate
and force of the heart's activity.

The salts employed in the experiments were all chemically
pure, either from Kahlbaum or specially prepared in the lab-
oratory, and they were all standardized. The water employed
in making up the solutions was twice distilled in glass.

To avoid mechanical stimulation a cannula, to which was
connected a rubber tube, was inserted either in the median or
lateral caudal vein and tied in place. Subsequently 4 cc. of the
solution per kilo weight of fish was slowly pressed from a
graduated glass tube into the cannula by means of a rubber
bulb attached to its upper end.

During the inspiratory phase the mouth and spiracles are
opened, the floor of the mouth is lowered, the gill clefts or ex-
ternal branchial apertures are closed, and water enters the
mouth and spiracles. In the expiratory phase the mouth and
spiracles close, the floor of the mouth returns to its normal
position or is elevated and the gill clefts open to allow the
water to escape. Occasionally a peculiar spouting takes place
during which water is thrown from the mouth and spiracles.
This is observed in the aquarium also and may be due to irri-
tation, touch, stimulation of the mucous lining of the mouth by
slime particles or vitiated water.

T first investigated the effect of different concentrations of

HYDE: EXPERIMENTS ON THE SKATE. 33

the same solution, seeking the weakest solution that produced
any change in the three functions under consideration; also,
which concentration proved toxic, and whether the injurious
dose could be modified or antagonized by a definite dose of an-
other salt.

Observations were made and curves secured before, during
and after the injection of the salt solutions. From one to five
minutes after the injection a change in one or more of the
functions under consideration usually occurred, and this was
followed by a second change that lasted from five to twenty
minutes, before the records again assumed their normal char-
acter. The first effect may be the reaction of the salt upon the
constituents of the blood and indirectly upon muscle and nerve
cells ; the second is due to osmotic pressure changes. That in-
jections of hypertonic salt solutions in the blood of the dog in-
creases the osmotic pressure, which may become normal within
twenty minutes, was proved by Magnus.^ Moreover, those
salts that set free 0, CO2, H, or OH ions, in their reactions,
exert their influence more often through these ions, and effect
results that differ from those produced by salts having the
same cations, but do not react with the same resulting ion
products; e. g., NaCl is different in its effects from Na2C0:!,
probably because of the OH ions set free by the Na2C03 dis-
sociation.

It is possible that the blood pressure, respiratory and cardiac
activities depend upon the number of molecules and ions in the
blood, and upon the dissociation tension of the salts. The pre-
potent anions or cations may stimulate or inhibit enzyme ac-
tion, and thus aid in producing the inner stimulus that affects
alike the force of cardiac and respiratory activity and directly
or indirectly the blood pressure.

A careful study of very many experiments with the same
salts was made, the average results summed up, and the most
typical of a series of experiments put in tabular form. Thus
table n gives a survey of the different series of experiments and
is a summary of all the experiments conducted on the skate
with NaCl solutions. It shows that m 32, m/16, m/8, NaCl
solutions produce as a rule practically no change in the heart
or respiratory activity or blood pressure. An m% NaCl solu-

4. Magnus, Archiv fiir Experimentelle Pathologie, 1900, vol. 44, p. 99.

34 KANSAS UNIVERSITY SCIENCE BULLETIN.

tion (see curve 1) increases the force and blood pressure but
leaves the rate of the heart and respiration unaltered.

An m/1 solution (curve 2) causes a rise in blood pressure
and force of heart's rhythm at once, which usually continues
for about twenty minutes. But the force and rate of respira-
tion is either unchanged for one to two minutes and then de-
creases ; or may, as does the heart rate, first decrease and then
return to normal.

Following the injection of a 2m NaCl solution (curves 3 and
4) the blood pressure increased for about five minutes, then
became normal. The rate and force of the heart and respira-
tory movements decreased for one to twenty-five minutes ; but
in many cases the force of the heart's activity first decreased
for one to two minutes and then increased, or increased as the
blood pressure rose. In every instance the systole and inspira-
tory phases were prolonged above normal with a 2m NaCl
solution.

A resume of the NaCl experiments shows that injections of
4 cc. per kilo fish of m/64 to m/8 solutions are as a rule prac-
tically indifferent; m% stimulates the force of heart and res-
piratory activity and raises blood pressure for from one to
five minutes, but is usually indifferent to rate; m/1 increases
the force of the heart's action and also blood pressure, but the
rate and force of respiratory activity differs in that they may
for one minute remain normal, then decrease; or may, as does
the heart rate, decrease for one to three minutes before re-
suming again the normal activity ; 2m decreases rate and force
of heart and respiration, but the heart's force is often increased
after one to two minutes' decreased action. The blood pressure
is raised during the first one to five minutes, then falls.

It might be concluded that the blood salts can be slightly in-
creased by 4 cc. per kilo m/64 to m/8 without stimulating or
inhibiting either of the three functions that are considered,
and that equilibrium of the osmotic pressure is rapidly estab-
lished. On the other hand, m% stimulates the force of both
heart and respiration. I infer that its effect is directly upon
the nerve centers for one to five minutes, before the attraction
of water to the blood establishes equilibrium. An increase to
m/1 has a stimulating effect only upon the force of the cardiac
center, thereby increasing also the blood pressure, but

HYDE: EXPERIMENTS ON THE SKATE.

35

strangely enough does not affect the respiratory center in the
same direction but rather depresses its force and rate, as well
as the rate of the heart's activity. But a 2m solution has again
a like depressing influence upon both force and rate of respira-
tory and cardiac rhythm and at first causes a rise in blood pres-
sure.

Curve 1. Following an injection of m% NaCl. Lever attached to
heart and gill arch.

Cnrve 1.

Upper, respiration. Lower, cardiac curve. Injection at X. Force of
heart and respiratory action increased, but not rate.

Curve 2. After NaCl. Lever attached to heart and gill arch. Normal
before X.

Curve e.

Before

After

-L.

Upper, respiration. Lower, cardiac curve. Heart and respiratory
rate unchanged; force of former increased but of latter unchanged or
even less. Time, five seconds.

Data of the mean results in respiration, heart action and
blood pressure of many experiments on the skate. Different

36

KANSAS UNIVERSITY SCIENCE BULLETIN.

concentrations of NaCl were intravenously injected in the pro-
portion of 4 cc. per kilo fish :

Table I. Effect of NaCl solutions.

Salt amt.

Mol.

Heart
rate.

Heart
force.

Respir-
atory
rate.

Respir-
atory
force.

Blood pressure.

Time of
change.

Curves.

Nor-
mal.

Change

NaC12cc.

m/32

Neutral.

Neutral.

Neutral.

Neutral.

24

24

NaC12cc.

m/16

Neutral.

Neutral.

Neutral.

Neutral.

20

20

NaC12cc.

m/8

Neutral.

Neutral.

Neutral.

Neutral.

14

14

NaC12cc.

m%

Neutral.

In-
creased.

Neutral.

In-
creased.

26

28

First 1-5
minutes.

1

NaC12cc.

m/1

Fell 1-3
minutes,

then
normal.

In-
creased
at once.

Normal
or less.

Normal
or less.

17

20

During

20
minutes.

2

NaC12cc.

2m

Fell 36-
30; sys-
tole pro-
longed.

First 1-2
miniites

rose,
then de-
creased.

First 5
min. fell,
inspira-
tion pro-
longed.

De-
creased.

20

23

First 5
minutes.

3, 4

Curve 3. After 2m NaCl. Heart directly attached to lever.

Cvarre 3.

Upper, respiration. Lower, heart. Time, five seconds. Systole and
inspiration much prolonged.

Curve 4. 2m NaCl. Hiirthle mercury manometer,

CUTTO 4.

Normal, 10:35 A. M.
Heart rate, 36.

Respiration rate, 36,

Pressure, 20.

10:40 A.M.

Heart rate, force slightly
increased.

Respiration rate 27,
force slightly de-
creased.

Pressure, 23.

10:45 A.M.

Heart rate, 30, force in-
creased.

Respiration rate 36,
force less.

Pressure, 23,

HYDE: EXPERIMENTS ON THE SKATE.

37

Curve 5. M/32 KCl. Heart and blood pressure were obtained with
Hiirthle manometer. For respiratory records the lever was attached to
the gill arch.

Curve 5.

The upper curve heart, and from abscissa to heart blood-pressure
record.

Below the latter respiratory curves, and the time in five seconds.

Blood pressure remained unchanged, respiratory rate decreased, force
unaltered, expiratory phase prolonged. Heart rate from 24 to 22, pro-
longed diastole and force unchanged.

Curve 6.
arch.

M/8 KCl. Lever attached to the tip of the ventricle and gill

Curre 6.

AAAnAn/WWlJYUl

Upper, respiratory curve. Lower, heart. Time in five seconds.

The first part of the upper curve of respiration, taken before injection,
registered 36; immediately after, force slightly less, but the rate increased
to 42, and in ten minutes was normal again. The first part of the lower
heart curve before injecting was 34 per minute. For about twenty
minutes it was less (27), with prolonged diastole, then gradually in-
creasing during fifteen minutes to normal. Force was unchanged.

Curve 7. Effect of m/8 KCl.

Curra 7.

Upper, heart and pressure recorded with Hiirthle mercury manometer.

Lower, respiratory records from lever attached to a gill arch.

Pressure before injecting m/8 KCl 20 mm.; after injecting for two
minutes 22 mm., then normal. After injecting the solution, the respira-
tion for about twenty seconds decreased in rate with prolonged inspira-
tion, then for about one minute increased, and in fifteen minutes was nor-
mal. Force was unchanged for twenty to sixty seconds, heart rate de-
creased with prolonged diastole, then in five minutes normal; force
unchanged.

38

KANSAS UNIVERSITY SCIENCE BULLETIN.

Curve 8. M% KCl. Curves secured with heart and respiratorj- ievers.

Curve 8.

rv\m\jy

Upper, respiration. Lower, heart cui've.

Within one to two minutes after the m% KCl was injected, the heai't
action ceased. Respiratory force decreased, rate increased, and stopped in
twelve minutes.

Curve 9. m% KCl. Curves secured with lever and Hurthle man-
ometer.

Curve 9.

The upper are cardiac, the lower are respiratory records. Time, five
seconds.

Within one minute blood pressure and heart action were zero. Respira-
tory rhythm decreased and in twenty minutes stopped.

Table IL Mean results in respiration, heart action and blood pressure fol-
lowing injection of different concentrations of KCl solutions.

Salt mol.
4 cc. per
kilo.

Heart
rate.

Time.

Heart
force.

Respira-
tory
rate. '

Time.

Respira-
tory
force.

Blood p

Nor-
mal.

resBure.
Change

Curves.

KCl
m/100

Neutral.

Un-
changed.

Un-
changed.

Un-
changed.

20-60 sec.

Un-
changed.

De-
creased
or un-
changed.

Un-
changed.

18

18

KCl

m 32

Slower.

20-60 sec.

Increase.

then less,

or less

at once.

18
20

18
20

5

KCl

m/8

De-
creased.

20-60 sec.

Un-
changed.

De-
creased.

At once.

20
20

18. 22.

then

20

6
7

KCl

m%

De-
creased,

then
ceased.

5 sec.

In-
creased
for 30 sec.
then less.

In-
creased,
stopped
or de-
creased.

5 sec;
7 min.

De-
creased.

19
or

20

22.

1 min.

0, in

3 min.

8
9

The influence of KCl injections are summarized in table II,
which represents the mean results of many experiments of the
different series of KCl solutions. Curve 5 is a typical curve
secured after injecting the usual amount of an m/32 KCl solu-
tion. From the table and the curve it is observed that an m/32

HYDE: EXPERIMENTS ON THE SKATE. 3d

KCl solution produces a first effect of from twenty seconds to
one minute upon the heart and respiratory activities. More
often during this time the heart rate is slower, due to prolonged
diastole, and the force remains unchanged. The inspiratory
phase is, however, lessened while the expiratory is relatively
prolonged.

Curves 6 and 7 represent records obtained with different
methods after injecting an m/S KCl solution. Although many
other records were made these were chosen because they rep-
resent the more general type. There were exceptions in every
case, but those variations were accounted for either because
of the morbid condition of the skate, because it had been in the
aquarium several days, or had been under observation for sev-
eral hours and was fatigued or more or less irritable. After
the solution was injected for from twenty to sixty seconds the
heart activity greatly decreased and diastole was much pro-
longed. Occasionally the action increased above the normal
rate for one to two minutes, and in ten to fifteen minutes was
normal. The force remained unchanged. The respiratory
rate as a rule decreased for fifteen to thirty seconds, with
prolonged inspiratory phase, then either suddenly increased
above normal or gradually returned to the normal rhythm.
The force was usually unchanged. The blood pressure either
fell or increased slightly, about 2 mm. mercury, for from one
to two minutes.

Following the injection of m% KCl, pressure fell either very
suddenly to zero or in rapid leaps from 20 to 16 in one minute
and zero in two minutes, or often 19 to 22 in one minute then
zero within three minutes. The heart decreased in rate; e. g..
from 30 to 18, with increased force in one and zero in two
minutes. Respiration slowed from 39 to 30 in five seconds to
18 and very shallow and often irregular and spasmodic in ten
minutes, or rapid, 46 to 60, in five seconds, and gradually to
zero in seven minutes, but in some cases continued active as
long as twenty minutes after the heart had ceased contracting.

A resume of the effects of KCl solutions brings to light that
diastole and inspiratory phases are prolonged above normal in
the different strengths of solution. Blood pressure falls in all
but the most dilute solutions. The force of the heart and res-
piratory activity is practically unchanged in all but the toxic
strength, m%, after which it decreases. The rate of the heart's

40 KANSAS UNIVERSITY SCIENCE BULLETIN.

contractions as a rule is at first decreased in all strengths but
the m/100; m% is toxic to heart and respiratory activity,
though usually the respiratory rhythm continues for five to
twenty minutes after the heart has ceased.

A summary of some experiments that were undertaken to
study what salts counteracted the toxic influence of KCl solu-
tions is shown in table 115, and curves 10, 11, 12 and 13 illus-
trate a few of the experiments.

Curve 10 records the antitoxic effect of an m% NaCl solution.
The rate of the heart's action was 36, that of the respiratory
26. At 10:05 m% KCl was injected; within a few seconds the
heart action became irregular and respiration rapid, then ir-
regular, with prolonged expiratory phases. Blood pressure
fell. At 10 :15, an injection of m% NaCl was followed in a few
seconds by regular and strong cardiac and respiratory con-
tractions, which in ten minutes were quite normal, but the
force of the heart-beat was stronger. In other experiments
the toxic effect of an m% KCl was quickly counteracted by an
m/8 NaCl solution.

Curve 11 shows the favorable influence of CaCh with or pre-
ceding a KCl solution; m/4 KCl following m/4 KCl-fm/32
CaCk shows a decrease in force of heart and respiratory ac-
tivity, and a toxic m% KCl following a mixture of m% KCI4-
m/32 CaCk shows, in addition to a decrease in rate, a marked
decrease in force, but in twenty minutes the heart and respira-
tory rhythm is almost normal. The CaCk evidently proved
antitoxic, and with KCl enhances the force of both respiratory
and cardiac action.

Curve 12 also shows that a dose of m/4 CaCk that preceded
an m% KCl and ml KCl counteracted their toxic effects.

Curve 13 : After CaCk mVs, blood pressure stood at 14 mm.,
heart action at 24, respiratory action at 36. When m% KCl
was injected, pressure fell to 2 mm, heart to four contractions
and respiration increased to 42. At the end of two minutes
pressure was again 14, heart and respiratory rate had in-
creased, though shallow. At the end of five minutes, pressure
and action of heart and respiration were quite normal, but
respiratory rate more rapid. This shows that when stronger
solutions of Ca are injected, and the blood is in excess of the
normal, it counteracts the toxic effect of m% solution of KCl
quicker and the toxicity is less marked on respiratory than

HYDE: EXPERIMENTS ON THE SKATE.

41

heart action. In some experiments the toxic effect of KCl
m% was overcome if CaCL was injected soon after the KCl
sokition.

Table 116. Salts that counteracted the toxic effect of m% KCl.

Salt amt.

Mol.

Heart.

Respir-
atory.

Blood
pres-
sure.

Anti-
toxic.

Mol.

Heart.

Respir-
atory.

Blood
pres-
sure.

a

4 cc. KCl.
per kilo.

m%

IrregTi-
lar.
then
spas-
modic.

Rapid,
then
spas-
modic.

Pres-
sure
fell.

NaCU cc.
followed
m% KCl.

m%

Regular

in two

minutes.

Regular

in two

minutes.

Rose.

10

KCl

in/ 4
subse-
quent.

Rate un-
changed,

force

less.

Rate in-
creased,

force

less.

CaCU +

KCl.
Preceded-

m/32

m/4

Rate un-
changed,
force in-
creased.

Rate
less,
force in-
creased.

11

KCl

m%
subse-
quent.

Rate and
force
less.

Rate and
force
less.

CaCH-

KCl.

Preceded.

m/32

m%

Rate and
force in-
creased.

Rate and
force in-
creased.

11

KCl
KCl

m/1

Slight

increase

in rate

and force

Un-
changred.

Rate

slight in-
creased,

force

less.

CaCU.
Preceded.

m,4

Stimu-
lant force
and rate
anti-
toxic.

Stimu-
lant force
and rate
anti-
toxic.

Rose.

12

KCl

in%
subse-
quent.

Almost

stopped;

in five

minutes

quite
normal.

Rate in-
creased,

force
fell, then
normal.

Fell.

CaCU.
Preceded
m% KCl.

mi

24

36

Nor-
mal.

13

Curve 10. Toxic action of KCl m% counteracted by NaCl m%.
attached to ventricle and gill arch.

Levers

nfWmyy^ryrv-^^^^_^^

Time, five seconds. Upper, respiration. Lower, cardiac curves.^

5. Within a few seconds after injecting m% at X, the heart action became irregular
and spasmodic in long diastole. In twenty-five seconds the greatly increased respiratory
movements assumed prolonged expiratory spasms and a few seconds following m% NaCl
injection both respiratory and cardiac action became more regular again. m% NaCl
proved antitoxic for the toxic dose of m% KCl.

42

KANSAS UNIVERSITY SCIENCE BULLETIN.

Curve 11. Toxic effect of KCl m% counteracted by CaCl2 m/32 for the
upper curve. For lower curve, the lever was attached to the ventricle,
and to the gill arch for upper.

Curve 11.

m mmvm]^'''^^^^^^^^^^

■^ '^ " f+!2 Ca Cl

uKU

K ci " r c« «i ^

X K CI m -

Tlonml

Showing the antitoxic and stimulating action of CaCl2 2 cc. m/4 KCI4-
m/32 CaClo. Rate of heart contractions increased. KCl m/4 decreased
the force of heart and respiration, while rate of respiration increased.
The difference between the effect of m% KCl and KCl m% plus
CaCl2 m/32 demonstrates that CaCl2 increases force of heart and respira-
tion.

Curve 12. Toxic effect of KCl m% or m/1 counteracted by a previous
dose of CaCl2 m/4. For lower record the lever was attached to the ven-
tricle, and to the gill arch for the upper curve.

Curve 12.

^C^Cl

tnX KCl

caA/v^

5 KCl

Following 8 cc. of gradually increasing to m/4 doses of CaCl2 the
heart became distended and contracted more forcibly, due to the intra-
cardiac pressure; m% KCl injected increased the rate of cardiac and
respiratory activity and an m/1 KCl injected ten minutes later failed to
produce its usual toxic effect.

Curve 13. KCl m% following m% CaClo. After m% CaCb was in-
jected the pressure stood 14 mm., heart rate 24 and respiratory 36. At
3:50 P. M. KCl m% was injected. In one minute pressure fell to 2 mm.,
heart rate to 4 and respiratory rate increased to 42. At the end of two
minutes pressure stood 14, heai't force increased rate 8, respiratory I'ate
48, force less. At the end of five minutes pressure and heart action were
quite normal while respiration was abnormal in rate but normal in force.

Curv-o 13.

CaCl "> !!'

KCI "X

fXTt\r\

r\^^^^/'^^'^A/^^vr\^\^v^^J\A/^AA^\P\/^^

Upper curve, heart and pressure from abscissa. Lower curves, res-
piration. Time, five seconds. Records with Hiirthle and lever manometer.

HYDE: EXPERIMENTS ON THE SKATE.

43

The results of the experiments with CaCk are briefly tabu-
lated in table III and partly shown in curves. It was observed
that after an injection of the usual amount of m/32 CaCk
solution (curves 15, 16), the blood pressure remained un-
changed. Respiratory activity was usually increased for one
to three minutes, and the cardiac contractions continued un-
changed or did so after about one minute of decreased rate and
increased force. Following an injection of an m/8 CaCk solu-
tion, the rate of cardiac and respiratory activity is usually de-
creased and the force of both increased. The blood pressure
rises for about one minute. Often systole and inspiratory
phases predominate, and furthermore, the solution proved an-
titoxic to many toxic salt solutions. The effect of an m% CaCk
was first a decrease in rate of cardiac and respiratory activity,
often irregular, with prolonged systole and inspiratory phases.
The force of both activities was usually increased, as was also
the blood pressure, for from one to three minutes. If the force
of the heart-beat was not increased the blood pressure did not
rise above normal.

Table III. Mean results in respiration, heart action and blood pressure,
following injections of different concentrations of CaCls solutions.

Salt mol.

Heart
rate.

Heart
force.

Dura-
tion.

Respir-
atory
rate.

Respir-
atory
force.

Dura-
tion.

Pressure.

Curves.

Nor-
mal.

Change

CaCU
m 32

Un-
changed
or de-
creased.

Un-
changed

or in-
creased.

1 min.

Un-
changed

or in-
creased.

Un-
changed
or in-
creased.

3 min.

20

20

15.16

CaCl,
m,8

De-
creased
or un-
changed.

In-
creased
or un-
changed.

1-3 min.

Un-
changed
or de-
creased.

Un-
changed

or in-
creased.

1-3 min.

17

18

17,18

CaCl2

m%

De-
creased

or ir-
regular.

In-
creased.

1-3 min.

De-
creased

or ir-
regular.

In-
creased.

1-3 min.

22

24.22

19,20

Curve 15. Following an injection of m/32 CaCl2.

' Curve 15

2-Univ. Sci. Bull.. Vol. V, No. 4.

44

KANSAS UNIVERSITY SCIENCE BULLETIN.

Rate less and force of heart unchanged; rate of respiratory activity
decreased for one minute, then normal, force increased.

The upper respiratory and the lower cardiac records were secured with
levers directly attached to gill arch and apex of heart.

Curve 16. After m/32 CaClo.

Curve 16.

The upper curve, Hiirthle manometer heart record. Lower curve, gill-
arch lever records of respiratory activity. Rate of heart decreased, force
unchanged. Respiratory activity and pressure unchanged.

Curve 17. Effect of m/8 CaCl2.

Currs 17.

— >

' — ■—*■-■■ ■ , ■ .. ■ -.1. ■ ■ _ _i 1 III ■ *

The upper curve is a record of the cardiac and blood-pressure changes
secured with a Hiirthle manometer. The lower respiratory is a gill-arch
lever record. Rate decreased, force increased for both activities and
blood pressure showed slight rise for a minute or two.

Curve 18. Effect of m/8 CaC^.

Ourre 10.

^"^^^^ f\{\AAAAM_

Upper, cardiac record secured with lever attached to the heart. Lower,
gill-arch record. Respiratory rate unchanged, force increased. Cardiac
rate decreased, force increased.

Curve 19. Effect of m% CaC^.

CUTTB 19.

^'^^^rr^'^^^^fNAf;;^^

HYDE: EXPERIMENTS ON THE SKATE.

45

Hiirthle cardiac records above, gill-arch lever records below. Time,
five seconds. Cardiac and respiratory rate decreased, force and blood
pressure increased for about three minutes, then quite normal for about
thirty minutes. First effect often irregular and spasmodic, especially for
cardiac rhythm.

Curve 20. Effect of m% CaCla.

Curve 20.

Upper curve respiratory, lower cardiac. Levers directly attached to
heart and gill arch. Cardiac and respiratory rates first decreased, force
increased; irregular prolonged diastole and inspiratory phases.

The results following injections of MgS04 solutions are
summarized in table IV, and illustrated in the curves. It was
observed that an m/64 or m/8 MgS04 had a more depressing
influence on respiration than on cardiac activity. The rate was
either unchanged or less, but force and blood pressure usually
sHghtly greater for cardiac, while rate and force of respiratory
activity were generally decreased.

An m% or m/1 solution produces for about two minutes an
increase in blood pressure followed by a decrease. Respiratory
and cardiac rate and force both decrease, but occasionally the
force of the heart's activity is first increased. Sensory re-
flexes are all reduced. With every strength of the solution the
force and rate of the respiratory rhythm and the cardiac rate
were decreased, but blood pressure and cardiac force were in-
creased or decreased after a previous increase. With strong
solutions, e. g., m/1, diastole and expiratory phases predomi-
nated and the activities may become irregular, but an m/8
CaCl2 overcomes the toxic effect and increases the force of the
respiratory phase and blood pressure.

46

KANSAS UNIVERSITY SCIENCE BULLETIN.

Table IV. Mean results in cardiac and respiratory activity and blood pres-
sure following injection of different concentrations of MgS04.

Salt mol.

Heart
rate.

Heart
force.

Dura-
tion.

Respir-
atory
rate.

Respir-
atory
force.

Dura-
tion.

Pres-
sure.

Pressure

change.

Curves.

MgS04
m 64

Less
or un-
changed.

Un-
changed

or in-
creased.

15 min.

Less
or un-
changed.

Less.

15 min.

20

21
20

21

MgS04
m/8

Less
or un-
changed.

Increased

or un-
changed.

6 min.

Less
or un-
changed.

Less.

5 min.

18

19
18

22

m5 8 or
ml

Less
or un-
changed.

De-
creased
after rise.

Rise 2 m.

Less.

Less.

3 min.

22

23

21

23

Curve 21. Effect of MgS04 m/64.

Jiiryo 2'

"lormal

f-\ArvrvYM\r\r\-

Upper curves, cardiac and pressure with Hurthle. Lower, respiratory
gill-arch lever. Time, five seconds. At x, injection. Cardiac rate less,
force and blood pressure slightly greater; respiratory rate less, force
unchanged.

Curve 22. Effect of MgS04 m/8.

Curve 22.

Upper curve, cardiac and blood pressure shows decrease in cardiac
rate, slight increase in force and blood pressure, for two minutes. Lower,
respiratory gill-arch lever curve; decrease in force, not in rate.

Curve 23. Effect of MgS04 m%.

Ourre 23.

Upper curves, cardiac and blood pressure; rate unchanged, force and
blood pressure slightly increase for two minutes, then decrease. Lower,
respiratory curve, decrease in rate and force.

HYDE: EXPERIMENTS ON THE SKATE.

47

The effects of NaiiCO.-; injections are tabulated in table V.
It was observed that Nai^CO.; solutions are most effective in
strengthening the force of the cardiac rhythm and thus rais-
ing the blood pressure. They do not stimulate respiratory
activity except in strong doses, when they may increase the
force. Its pronounced influence upon the heart may be due to
OH ions that are set free by the dissociation.

^ Table V. Resume of Na^COs injections upon cardiac and respiratory

activity and blood pressure.

Salt.

Heart
rate.

Heart
force.

Dura-
tion.

Respir-
atory
rate.

Respir-
atory
force.

Dura-
tion.

Pres-
sure.

Pressure
changes.

Curves.

Na^COa
m/32

Un-
changed.

In-
creased.

15 min.

Un-
changed.

Un-
changed.

22

24

24

m/'8

Un-
changed
or
less.

In-
creased.

15 min.

Un-
changed.

Un-
changed.

20

23

28

25

m%

Un-
changed
or
less.

In-
creased.

2 min.

Un-
changed
or de-
creased.

In-
creased.

26

26

Curve 24. Effect of Na2C03 m/32.

Cuire 24.

Upper curves, cardiac and blood pi'essure with Hiirthle. Rate un-
changed, force and blood pressure increased. Lower, respiratory; force
and rate unchanged. Time, five seconds.

Curve 25. Na2C03 m/8.

Qam ZS.

rNrsiM\r\r\r\r\rMArNrvr\

Upper curves, cardiac and blood pressure. Rate decreased, force
greatly increased, pressure rose. Lower curve, respiratorv; force and rate
unchanged. Time, five seconds.

48

KANSAS UNIVERSITY SCIENCE lU 11 FTIN.

Curve 26. Na2C03 m/2.

Curre 26.

Upper, respiratory gill-arch lever records. Increased force, decreased
rate. Lower, cardiac lever directly attached to heart. Increased force,
decreased rate. Time, five seconds.

Na2HP04 solutions are not so stimulating to cardiac rhythm
as are Na-CO.-: solutions. This is shown in table VI. More-
over, for the same strength they are more toxic. The solutions
act like Na2C0n chiefly upon the force of the cardiac action and
are indifferent in weak solutions on respiratory movements.

Table VI. A summary of experiments of intravenous injections of
Na2HP04 relating to cardiac and respiratory activity and blood pressure.

Mol.
sol.

Heart
rate.

Heart
force.

Dura-
tion.

Respir-
atory
rate.

Respir-
atory
force.

Dura-
tion.

Pres-
sure.

Change.

Curves.

HPO4

m 64

Less or

un-
changed.

In-
creased
or un-
changed.

5 min.

Un-
changed.

De
creased.

5 min.

23

24

27

m 8

Less or

un-
changed.

In-
creased
or un-
changed.

5 min.

In-
creased
or un-
changed.

De-
creased.

5 min.

24

22

28

m/4

Less.

then
rapid or
stopped.

Less.

5 min.

Less,

then
rapid.

De-
creased.

3 min.

23

18

29

Curve 27. Effect of Na2HP04 m/64.

Curre 27.

Upper, cardiac and blood-pressure Hiirthle manometer cui'ves; show
force of heart action and blood pressuie; slight increase, rate decrease.
Lower, respiratory rhythm unaltered. Time, 5 seconds.

HYDE: EXPERIMENTS ON THE SKATE.

49

Curve 28. Effect of Na2HP04 m/8.

Curra 28.

Upper, cardiac and blood-pressure Hiirthle manometer curves. Rate
little less, force slight increase. Lower, respiratory record; shows rate
unchanged, force decreased, pressure decreased.

Curve 29. Na2HP04 m/4.

Cnrre 29.

Upper, cardiac and blood-pressure Hiirthle curves; show fall in blood
pressure and for three minutes almost cessation of cardiac action. Lower,
respiratory, force much decreased.

Curve 296.
tions.

Effect of gradually increasing strength of Na2HP04 solu-

^AMAAAA

Upper, respiratory gill-arch lever curves show that with increasing
strengths from m/64, m/32, m/8, m/4, solutions that the rate increased
and force decreased from the normal and that this solution is more favor-
able to respiration as stimulating rate. Lower, cardiac apex lever curve
shows that the rate remained practically unchanged except in m/4, when
it became rapid and force decreased gradually from the normal.

The influence of NaOH solutions is briefly tabulated in
table VII. It is demonstrated that it acts, augmenting in all
concentrations, on the force of the heart-beat and has not the
same but rather a depressing effect on the respiratory center.
The OH ions here, as in the Na2C03, are the stimulating fac-
tors, but act more powerf ally than in Na^COa solutions. The

50

KANSAS UNIVERSITY SCIENCE BULLETIN.

respiratory force decreases with increased strength of the so-
lution, while the blood pressure rises in all concentrations em-
ployed, showing that increase in blood pressure is not neces-
sarily associated with rise of respiratory activity but rather
with force of heart-beat.

Table VII Showing the effect of NaOH so'utions upon respiratory,
cardiac and blood- pressure changes.

Salt tnol.

Heart
rate

Heart
force.

Dura-
tion.

Respir-
atory
rate.

Respir-
atory
force.

Pres-
sure.

Pres-
sure
change

Curves.

NaOH m 64

Un-
changed
or less.

In-
creased.

5 min.

Un-
changed.

Un-
changed.

25

26

30

m/32

Un-
changed
or less.

Much
increased.

15 min.

Decreased

or un-
changed.

Decreased

or un-
changed.

24

29

31

m,8

Un-
changed.

In-
creased.

10 min.

In-
creased.

De-
creased.

24

27

32

m 4

Less
or un-
changed.

In-
creased.

10 min.

Un-
changed or
increased.

De-
creased.

26

29

33

Curve 30. Effect of m/64 NaOH injections.

curva 30.

Upper, Hiirthle curves of cardiac and blood-pressure changes; decrease
in rate, increase in force and blood pressure for two minutes. Lower,
gill-arch lever curves of respiratory changes unaltered.

Curve 31. Effect of m/32 NaOH solutions.

Cxirve 31.

Upper, Hiirthle curves of cai'diac and blood-pressure changes show a
first effect of two minutes of greatly increased force and slight decrease
in rate of cardiac rhythm. The blood pressure increased for two minutes,
respiratory rhythm decreased very little if at all.

Curve 32. Effect of m/8 NaOH solutions.

Curre 32,

Upper, Hiirthle curves of cardiac and blood-pressure changes show in-
creased blood pressure and rhythm. Lower, respiratory curves, mcieased
rate but decreased force.

HYDE: EXPERIMENTS ON THE SKATE.
Curve 33. Effect of m/4 NaOH solutions.

51

Upper curves, blood pressure and force of cardiac rhythm increased.
Lovi^er, respiratory activity decreased in force, rate increased.

From the series of experiments with different concentra-
tions of urea solutions the conclusion is drawn that injection
of urea solutions of high concentrations, 2m to 4m, increases
force of respiratory and cardiac activity mainly. The rate is
practically unchanged, and weaker solutions are quite indiffer-
ent on blood pressure, cardiac and respiratory activity. It has
been found by Schroder*^^ that selachian's blood normally con-
tains 2.6 per cent of urea, and Baglioni" found that the selach-
ian's heart beats longer in urea than in NaCl. According to
Loeb it is due to the OH ions which are present when the dis-
sociation of the urea occurs. A weak solution of m/20 of
Na2C03, to which a 2m urea solution has been added, greatly
increased the force of cardiac action and blood pressure.

Table VIII. Effect of urea solutions.

Salt mol.

Heart
rate.

Heart
force.

Dura-
tion.

Respir-
atory
rate.

Respir-
atory
force.

Dura-
tion.

Pres-
sure.

Pressure
changes.

Curves.

Urea.
m/50

Un-
changed.

Un-
changed.

15 min.

Un-
altered.

Un-
changed.

15 min.

20

20

ml

Un-
altered.

Un-
altered.

15 min.

Unal-
tered or
less.

Unal-
tered or
increase.

1 min.

21

21

34

2m.

Slight
rise.

Slight
increase.

2 min.

Un-
changed.

Un-
altered.

15 min.

20

20

35

4m.

Un-
changed.

Increase.

Un-
changed.

Increase.

22

21

36

Curve 34. Urea m/1.

I u

6. Schroder. V. z,eitschrift fiir Physiologische Chemie, XIV, S. 596.

7. Baglioni. Zentralblatt fur Physiologie, 1905, XXX, No. 12.

52 KANSAS UNIVERSITY SCIENCE BULLETIN.

Upper, respiratory curve, with lever attached to gill arch, shows first
decrease in rate for about one minute, then rate normal, force slightly in-
creased. Lowei-, cardiac curve, with lever attached to apex. Cardiac
activity unaltered.

Curve 35. Urea 2m.

Carre 35.

Upper, Hiirthle cardiac record shows little increase in force and rate.
Lower, respiratory rate and force unaltered.

Curve 36. Urea 4m.

Curve 36,

Upper, Hiirthle cardiac curve; increased in force and pressure, rate
unchanged. Lower, respiratory cvirve; force slight rise, rate unchanged.

The results obtained with standardized HCl solutions are
tabulated in table IX. The weaker solutions, ranging from
m/100 to m/64, increase for the first minute or two the force
of the cardiac and respiratory and blood-pressure changes, and
decrease or act indifferently on the rate of respiratory and car-
diac activity. A stronger solution, m/32, for the first minute
or two decreases the rate but usually increases the force of
both the respiratory and heart action, and the blood pressure
usually rises for the first minute ; m /8 is toxic, but the respira-
tory rhythm continues several minutes after the heart ceases,
or if it continues it is rapid and faint and the respiratory
movements are continued with less force and rate. It is sup-
posed that the stimulating action is due to the H ions. In all
concentrations between m 100 and m 16 there is an initial
increase in pressure and force, followed by a decrease in all
activities before a return to either normal or less.

HYDE: EXPERIMENTS ON THE SKATE.

53

Table IX. Effect of HCl solutions.

Salt mol.

Heart
rate.

Heart
force.

Dura-
tion.

Respir-
atory
rate.

Respir-
atory
force.

Dura-
tion.

Pres-
sure.

Pressure
changes.

Curves.

HCl m/ 64

De-
creased.

In-
creased.

2 min.

Un-
changed.

In-
creased
or un-
chang:ed.

2 min.

19

20

37

m/32

De-
creased.

In-
creased.

1 min.

De-
creased.

Un-
changed

or in-
creased.

1 min.

24

26

38

m/8

Rapid.

Weak.

1 min..
then de-
creased.

In-
creased.

Un-
changed

or in-
creased.

1 min.,
then de
creased

21

10
0

39

Curve 37. Effect of m/64 HCl on cardiac and respiratory activity and
blood pressure.

Cur»e 37

Upper, gill-arch lever respiratory curve shows for two minutes in-
creased force, rate unchanged. Lower, apex lever cardiac curve for two
minutes increased force, decreased rate.

Curve 38. m/32 HCl injection.

Carre 38.

rV\r'Ar~\r~Y--Ar~'Arv^rY

Upper, cardiac Hiirthle manometer curve and blood pressure shows de-
creased rate and increased force and rise of blood pressure for one
minute.

Lower, gill-arch lever record of respiratory activity shows rate less,
force unchanged for one minute.

54

KANSAS UNIVERSITY SCIENCE BULLETIN.

Curve 39. Effect of m/8 HCl on cardiac, respiratory and blood pres-
sure changes.

Upper, gill-arch lever respiratory curve shows increase at first of rate
and force, then gradual decrease. Lower, apex lever curve shows at once
fall in force and rapid heart-beat that soon becomes irregular and spas-
modic before ceasing.

Injection of different strengths of NH4CI solutions show that
weak solutions are quite indifferent. An m/8 solution increases
blood pressure and force of respiratory movements, but an
m% depresses cardiac and blood pressure activities and greatly
stimulates the expiratory phase of respiration, causing an ir-
regular spasmodic rhythm for fifteen minutes, and this is re-
placed by a weak but regular activity.

Table X. Effect of|NH4Cl!solutions.

Salt,
mol.

Heart
rate.

Heart
force.

Dura-
tion.

Respir-
atory
rate.

Respir-
atory
force.

Dura-
tion.

Pres-
sure.

Pressure
changes.

Curves.

NHXl
m 64

Un-
changed
or less.

Un-
changed.

3 min.

Un-
changed

or in-
creased.

Un-
changed
or less.

5 min.

26

26

m 8

Un-
changed
or less.

Un-
changed.

15 nnin.

Un-
changed.

Expir-
ation in-
creased.

15 min.

26

28

40

m%

Much
less.

Much
less.

15

Long ex-
piratory,
irregu-
lar.

First in-
crease,
then
regular
or less.

15

26

22

41

Curve 40. Influence of NH4CI m/8 on cardiac respiratory and blood-
jressure changes.

CvirTB 40.

rxf-xrNTM-srNrxrMNfxr
, I I ■ - — 1 — ■

Upper, Hiirthle cardiac and blood-pressure curves show that rate and
force are unchanged, blood pressure rose, Lowei', respiratory curve;
rate unchanged, force slight increase.

HYDE: EXPERIMENTS ON THE SKATE.

55

Curve 41. Effect of m% NH4CI.

Cxirvo 41.

Upper, Hiirthle cardiac and blood pressure show a decrease in cardiac
activity and blood pressure. Lower, i-espiratory curve; for fifteen min-
utes prolonged expiratory phase and irregular rhythm, then weak but
regular.

Effects of dehydrated chemically pure Na2S04 solutions are
most pronounced in the strong concentration of m/2 and m/1,
both of which increase the force of cardiac, respiratory and
blood-pressure activities, leaving the rate unchanged. The
effect lasts longer with the weaker solution, and the after-
effects are not, as with the saturated m/1 solution, a weaken-
ing of the rhythm and fall in pressure.

Table XI. Summary of NajS04 effects on respiratory cardiac, and blood-
pressure changes.

Salt mol.

Heart
rate.

Heart
force.

Dura-
tion.

Respira-
tory
rate.

Respira-
tory
force.

Dura-
tion.

Pres-
sure.

Pres-
sure
change

Curves.

NasSO.

m 64

Un-
changred.

Un-
changed.

15 min.

Un-
changed.

Un-
changed.

15 min.

26

26

42

in/2

Un-
changed.

Slight
increase.

15 min.

Un-
changed.

Slight
increase.

15 min.

26

27

43

ml

Un-
changed.

Slight
increase.

5 min.

Un-
changed.

In-
creased.

5 min.

27

29

44

Curve 42. Effect of m/64 Na2S04.

carve 42.

Upper, Hiirthle cardiac and blood-pressure curve, and lower, respira-
tory, all show no change after injection of the solution.

Curve 43. Effect of m/2 Na2S04.

Corra 43.

56

KANSAS UNIVERSITY SCIENCE BULLETIN.

Uppei', Hiirthle cardiac and blood-pressure curves; rate unchanged,
force and pressure slight increase. Lower, respiratory curve; rate un-
changed, force increased.

Curve 44. Effect of m/1 Na2S04.

Curve 44.

Upper, cardiac and blood-pressure curves, show slight increase in force
and pressure, rate unchanged. Lower, respiratory; rate unchanged, force
increased.

The effects of intravenous injections of BaCk solutions are
increased irritability and increased force and blood pressure,
especially with weak solutions. Like most of the other salt
solutions, it does not increase the rate of either heart or re-
spiratory activity. In strong solutions it is toxic, producing at
first irregular spasmodic cardiac and respiratory phases with
strong diastolic and expiratory phases and with rise of blood
pressure, or cessation of activity. The toxic effects can be
avoided if small doses are injected with gradually increasing
strengths, and the toxic effect is overcome by injections of
CaCk solutions.

Table XIL

Effect of BaCli on cardiac, respiratory and blood-
pressure changes.

Salt mol.

Heart
rate.

Heart
force.

dura-
tion.

"""
Respir-
atory
rate.

Respir-
atory
force.

Dura-
tion.

Pres-
sure.

Pres-
sure
change

Curves.

BaCU
m 64

Un-
changed.

In-
creased
diastole.

15 min.

Un-
changed

or
slightly

less.

Expira-
tion in-
creased.

2 min.

18 mm.

20 mm.

45

m 32

Un-
chansred.

Un-
changed

or in-
creased.

1 min.
8 min.

Un-
changed.

creased.

After
8 min.

16

16
18
14

46

m/8

Un-
changed
or slow
or spas-
modic.

In-
creased
irrita-
bility.

2-5 min.

Stops or
rapid
spas-
modic.

10-20
min.

2 min.

23

25

47

in/4

Toxic.

Toxic.

HYDE: EXPERIMENTS ON THE SKATE.

Curve 45. Effect of m/64 BaCl2.

Curve 46.

r\r\PvNMVV\rv

1 ■

Upper, cardiac curve; rate unchanged, force and blood pressure slight
increase. Lower, respiratory curve; rate unchanged, force increased.

Curve 46. Effect of m/32 BaCl2.

(Jarve 46.

Upper, Hiirthle cardiac curves; after one minute, rate, force and pres-
sure unchanged. In five minutes, force and pressure increased. Lower,
respiratory curve, shows rate unchanged, force increased.

Curve 47. Effect of m/8 BaClo.

B half an hour after A. Upper curves of A and B, cardiac and blood-
pressure records, show first spasmodic activity, then strong force and in-
creased pressure. These activities decrease in half an hour but the rate
continues unchanged. Lower, respii'atory curve, shows first spasmodic
action, then this stops for twenty minutes. In half an hour it is weak
and slower.

The effect of distilled water passes off quite rapidly. For the
first two minutes the rate of cardiac and respiratory activity
is lessened and the force of the heart-beat and blood pressure
slightly increased. The blood is rapidly affected and clots
easily; soon the equilibrium of salts is again established and
the normal action resumed.

Table XIII. Showing the effect of distilled water.

Distilled
H,0.

Heart
rate.

Heart
force.

Dura-
tion.

Respir-
atory
rate.

Respir-
atory
force.

Dura-
tion.

Pres-
sure.

Pressure
changes.

Curves.

3-4 cc.

Decrease.

Slight
increase.

2 min.

De-
creased.

Un-
changred.

2 min.

24

26
24

48

58

KANSAS UNIVERSITY SCIENCE BULLETIN.

Curve 48. Effect of distilled water.

Curve 48.

Upper, Hiirthle cardiac and blood-pressure curves, show decrease in
rate, slight rise in force and blood pressure for two minutes, then normal.
Lower, respiratory lever, gill-arch curve, shows decrease in rate, force
unchanged for two minutes.

Acid sodium phosphate solutions cause an increase in force
of respiratory and cardiac activity, and for the weaker solu-
tions at first also an increase in rate. Diastolic and inspiratory
phases are often prolonged and the toxic effect is overcome by
an alkali solution of sodium phosphate or one of CaCk.

Table XIV. Showing mean results of NaH2P04 solutions.

Mol.

solution.

Heart
rate.

Heart
force.

Dura-
tion.

Respir-
atory
rate.

Respir-
atory
force.

Dura-
tion.

Pres-
sure.

Pres-
sure
changre

Curves.

m/64

Rapid to
normal
in two

minutes.

Weak,
increase.

1 min.

2 min.

In-
creased.
Normal.

Less
increase.

1 min.

2 min

49

m 16

Increase.

Increase.

Decrease.

Normal.

1 min.
2d min.
10 min.

Un-
chan^red.

In-
creased.

10 min.

50

m 8

De-
creased.
Normal.

In-
creased.
Normal.

1 min.
5 min.

In-
creased.

In-
creased.

10 min.

51

Curve 49. Effect of m/64 NaH2P04 solutions

Upper records of respiratory activity before, during and following for
two minutes the injection of an m/64 acid phosphate solution. First
minute rate greatly increased, force less; within two minutes force gradu-
ally increased to slightly above normal and rate normal. The lower cai--
diac record .shows first rapid contractions that increase during the second
minute in force, becoming normal in rate and force in two minutes.

HYDE: EXPERIMENTS ON THE SKATE.

5^

Curve 50. Effect of m/16 NaH2P04 solutions.

Curve 60.

Upper, gill-arch lever records of respiratory activity, show increase in
force, rate unchanged. Lower, cardiac lever, records first an increase in
rate and force; second, rapid, shallow; in ten minutes quite normal in
rate and force.

Curve 51. m/8 NaH2Po4 solutions.

CurTe 61.

Upper, gill-arch lever records of respiration show increase in rate, and
after two minutes also in force; inspiration prolonged. Lower, cardiac
apex records. First minute prolonged diastole and increased force, fol-
lowed by slower rate; in five minutes quite nonnal in i-ate and force.

3-Univ. Sci. Bull.. Vol. V, No. 4.

60

KANSAS UNIVERSITY SCIENCE BULLETIN.

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HYDE: EXPERIMENTS ON THE SKATE,

61

Table XVI. Toxic and Antitoxic
Solutions.

Toxic.

Antitoxic.

Curves.

2m NaCl.

CaCU m 32

3

tn% KCl.

m% NaCl

11

m% KCl.

CaClo m 32

12

m*« KCl.

CaCl. m 4

13 .

m 1 NMgrSO,

CaCU m 8

14

NaH2P04 m 4

NaoHP04. or
CaCU m 16

m 1 NajSOi

CaCU m 16

BaCU m 4

CaClo m 8

BaCl.. m 4

NaOH m 8

NaOH m 16

HCl m 100

The following is a resume of the conclusions drawn from
the different experiments :

M% NaCl increases the force of cardiac and respiratory-
rhythm and blood pressure, but is indifferent to the rate during
the first five minutes. Weaker solutions are neutral, and a
stronger, m 1, solution is rather depressing except to heart
force and blood pressure, while 2m is generally depressing,
prolonging systolic and inspiratory phases.

The different strengths of KCl solutions proved depressing
to the activities under consideration. Diastole and inspiratory
changes are prolonged and the toxic effect of an m% solution
is counteracted by CaCl m '32 or NaCl m%.

CaCk solutions are indifferent or depressing to the rate of
both cardiac and respiratory activities, but stimulating to the
force and slightly so to the blood pressure. M% solutions pro-
long systolic and inspiratory phases.

Solutions of MgSOi are depressing to the respiratory mech-
anism and cardiac rate, but at first stimulate heart force and
blood pressure. Diastole and expiratory phases are prolonged
and the toxic effects are neutralized by m/8 CaCh solutions.

NaiCOo solutions stimulate heart force and blood pressure
and are indifferent to the heart rate and the respiratory mech-
anism. Their stimulating effect upon the force of cardiac
activity is enhanced when the solution is combined with urea,
and like the latter exerts a similar influence upon respiration
and heart action.

62 KANSAS UNIVERSITY SCIENCE BULLETIN.

Na-HP04 solutions stimulate as a rule only the force of heart
action. The blood pressure, heart rate, as well as respiratory
activities, are generally decreased or unaltered.

Solutions of NaOH proved more stimulating to cardiac force
and blood pressure than any of the other solutions that were
employed. They were depressing to cardiac rate and respira-
tory activity, except in strong solutions they increased the rate
of respiration. They affect the center that controls the force
of the heart-beat oppositely from what they do the center
which governs the respiratory force. Like other alkaline solu-
tions, they may exert their influence by favoring oxidation.

Urea solutions, except the strongest, are quite indifferent to
the functions under consideration. A 4m solution increases
the force of both the cardiac and respiratory activities; possi-
bly through the OH ions that are set free when dissociation oc-
curs.

Weak HCl solutions produce initial stimulations for a brief
period in cardiac and respiratory force and blood pressure, but
depress the rate, and a stronger, m 8, solution is toxic for
cardiac but less so for respiratory activity.

Weak solutions of NH4CI are indifferent. An m 8 has a
slight augmenting effect on blood pressure and the expiratory
phase, while an m.% solution depresses blood pressure and
cardiac rhythm, but heightens the force of the respiratory
mechanism for a short time.

Following the injection of strong Na-S04 solutions the force
of respiration and heart action and height of blood pressure
is increased, while the rate is unchanged. The addition of a
weak solution of CaCb is favorable.

The effect of a weak solution of BaCk' is increased force of
cardiac and respiratory rhythm and a rise of blood pressure.
Diastolic and expiratory phases are prolonged. The solutions
are more toxic than other salts of the same concentration, and
the toxicity is overcome by CaCk.

Injections of distilled water produce transient stimulating
effects in the force of cardiac contractions and blood pressure,
and depress the rate'.

Weak solutions of NaH-POa accelerate the rate and force of
heart and respiratory action. Diastole and inspiration are
prolonged and toxic solutions are neutralized by Na2HP04.

A consideration of the results tabulated in the foregoing

HYDE: EXPERIMENTS ON THE SKATE. 63

pages discloses the fact that increased force of cardiac activity
is usually accompanied by increased force of respiratory ac-
tion and blood pressure. But NH4CL m/8, urea, and Na2HP04
are exceptions. NH4CI increases blood pressure without in-
creasing cardiac force. Urea produced increased force of re-
spiratory and cardiac activity without raising blood pressure,
and NaiHPOi m/8 increases force of cardiac but decreases
force of respiratory action. Moreover, most of the salt solu-
tions of a definite concentration that augmented the force of
cardiac also increased the force of the respiratory rhythm and
proved either indifferent or depressing in their influence upon
the rate of both of these activities.

The exceptions are MgS04 m/8, which stimulates force of
cardiac but not that of respiratory activities and is depressing
to rate, and NaOH m/8, that stimulates cardiac force and rate
but not the respiratory rhythm.

The toxic effect of many solutions at certain concentrations
was counteracted by definite strengths of other salts. Of these
antagonistic solutions, CaCk m/32 or m/8 proved most ex-
tensively favorable. It was interesting to learn that certain
concentrations of some salt solutions prolong the cardiac sys-
tole and the inspiratory phases, while other solutions favor
diastolic and expiratory phases. M '2 NaCl and also m%
CaCli' prolong systole and inspiration, while KCl m/8, NaHs
PO4 m/8 prolong inspiratory activity and diastole. Moreover,
strong solutions of MgS04, NH4CI and BaCL prolong diastole
and expiratory activity.

The salts that were especially depressing upon the cardiac
and respiratory phases were KCl and MgS04. There are cer-
tain optimum per cents of salts and acid solutions, below or
weaker than which the effect is stimulating or initiates tempo-
rarily increased activity, and above or greater concentration
than which the solution inhibits or decreases activity or proves
toxic. Moreover, increased olood pressure, as a rule, is accom-
panied by a decrease in rate of cardiac and respiratory activity.

4-Univ. Sci. Bull . Vol. V, No. 4.

THE

KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 5— October, 1909.

(Whole Series, Vol. XV, No. 5.)

CONTENTS;

The Nutrition of the Embryo Sac and Embryo in Certain

Labiate, F. H. Billings.

PUBLISHED BY THE UNIVERSITY,
LAWRENCE, KAN.

Entered at the t)ost-offlce iu l.nwreuw :is secondclHss matter.

THE KANSAS UNIVERSITY
SCIENCE BULLETIN.

VoL.V, No. 5] OCTOBER, 1909. [™xv'™l

THE NUTRITION OF THE EMBRYO SAC AND EMBRYO

IN CERTAIN LABIATE.

BY F. H. BILLINGS.
Plates XI to XIV.

^^OON after the publication of a paper on seed development,
' in which various sympetalous families were considered,
Ihe writer began an investigation of embryo-sac nutrition as
found in the Labiatse. Literature upon the subject indicated
the presence of certain nutritive structures that resembled
some of those described in the paper mentioned above. The
structures consisted of outgrowths or specialized portions of
the embryo sac which by enlargement frequently replaced con-
siderable portions of the integument. The position and shape
of the structures are characteristic in some instances of the
family in which they occur. It is therefore one of the prob-
lems of the present investigation to ascertain what degree of
uniformity prevails among a number of genera of the Labiatse,
and thus determine, if possible, whether any special type of
embryonic structure is liable to become characteristic of the
family. Though the number of species studied is but a small
representation of the total number distributed over the earth,
still, as will presently be shown, a great enough similarity ex-
ists among them to warrant the supposition that a like re-
semblance probably exists among the remainder. It would not
be safe to extend such a conclusion to any family taken at
random, inasmuch as some families are aggregates of groups,
such as subfamilies, which may not be closely allied phylo-
genetically but which are associated on account of fulfilling
certain taxonomic requirements.

(67)

68 KANSAS UNIVERSITY SCIENCE BULLETIN.

One of the services that maj' ultimately be performed for
taxonomy is that of testing the homogeneity of families. The
results could be reached satisfactorily only after all the species
of any given family had been examined as to their ontogeny.
Taxonomically speaking, the Labiatse are regarded as a fairly
homogeneous family with well-marked characteristics.

Species whose common ancestors are not remote should show
similarity in embryonic particulars, since these are doubtless
the ones least influenced by environment.

No attempt was made to work out the stages preceding fer-
tilization in every instance. Several species were examined,
however, and the series of events constituting the earlier his-
tory of embryo-sac development were found to be identical.

The ovular primordium gives rise to a single megaspore
mother cell, from which originates an axial row of four poten-
tial megaspores. The most deeply seated of the four develops
into the ordinary type of an eight-celled embryo sac.

The growth of the embryo was followed in detail in Lamium
ample xicaule, and it was found that the various stages in de-
velopment correspond closely to the plan observed in most
dicotyledonous embryos. However, it is not with the develop-
ment of the embryo but with the behavior of the embryo sac
that this paper has most to do.

Many of the writings of early investigators who worked along
embryological lines show that there was little thought of as-
signing physiological significance to peculiar shapes assumed
by the embryo sac. More recently quite a number of observ-
ers have entered the field, and they have shown the wisdom
of correlating morphology and physiology. Bibliographies of
their papers have been published from time to time, so that
only the literature cited in the text, or very recent papers, will
be given here.

The single and unusually thick integument of many Sym-
petalse, including the Labiatse, appear to serve a double func-
tion. The outermost layers become transformed into a seed
coat, while the inner or remainder, wholly or in part, may be-
come a nutritive tissue (Nahrgewebe) which invites, as it
seems, the digestive and absorptive activity of the endosperm.
Sometimes the nutritive tissue is more or less localized, as can
be seen from the deposits of starch in certain regions. In other
instances the deposition of food material is general throughout

billings: embryo nutrition in labiate. 69

the integument, so that no special part would be likely to be-
come a particular point of attack by the endosperm.

When the absorptive activity of the endosperm is largely
centered in some outgrowth or differentiated portion of the
embryo sac, the term "haustorium" is applied to such a por-
tion. In instances where haustoria are not developed, the di-
gestive and absorptive power is more or less equally distrib-
uted over the entire surface of the embryo sac. Where ab-
sorption is going on most rapidly, the cells directly concerned
in the appropriation of the nutritive material are usually
abundantly supplied with plasmatic contents. In case of the
presence of a haustorium the endosperm cells lying adjacent
to the base of the haustorium show the greatest richness in
plasmatic contents.

The following genera and species of Labiatje form the bases
of the detailed description :

Lamium amplexicaule. Mentha cmiadensis.

Stachys palustris. Salvia lanceolata.

Phrysostcgia virginiana. Salvia azurea.

Leonurus cardiaca. Monarda fistulosa.

Brunella vulgaris. Nepeta catana.

Teucri'iim canadefise. Dracoccphalum sp.

Pycnanthemum lanceolatwm. Scidellaria parvula.

Lycopus rubelhis.

Lamium amplexicaule.
This species will be given more detailed consideration than
the others, for it will serve in a measure as a type of the fam-
ily and also as a basis of comparison for the other species.
The embryo sac, at the time fertilization is accomplished, con-
tains the usual form of egg apparatus (fig. 1) . The endosperm
nucleus is large, spherical in form, and contains a vesicular
nucleolus. The nucleus does not lie near the egg apparatus,
as is the case in so many plants, but at the opposite end of the
sac near the disintegrating antipodals. The contour of the sac
shows a constriction which may be used to distinguish a larger
upper, or micropylar portion, in which the egg apparatus lies,
from a smaller, lower, or antipodal portion, in which the endo-
sperm nucleus lies. The endosperm nucleus does not leave the
antipodal region, but immediately after fertilization develops
a tissue in its end of the sac. Double fertilization probably
occurs, though it was not observed.

70 KANSAS UNIVERSITY SCIENCE BULLETIN.

The egg responds to fertilization by elongating greatly, and
pushing down into the forming endosperm tissue (figs. 2,
3). The elongation is at first due to the growth of the en-
tire egg cell. A transverse wall soon separates the suspensor
from the proembiyo. It is thus seen that the cell which is to
develop into the embryo is removed altogether from the mi-
cropylar portion of the sac. For a time this portion contains
no active nuclei, the synergids being in a process of disintegra-
tion. But endosperm nuclei soon migrate into it through the
space lying between the suspensor and the wall of the embryo
sac. The nuclei are but few in number and they do not form,
tissue (fig. 4). From this time on, the two extremities of the
embryo sac are very sharply distinguished. The antipodal end
is soon filled with a compact growing tissue that surrounds
the embryo. Increase in size of the tissue takes place rapidly,
so that the micropylar portion shortly becomes the smaller
(fig. 4). A remnant of nucellus can usually be seen as a dis-
organized mass of cells lying at one side of the micropylar ex-
pansion of the sac. For a considerable length of time the endo-
sperm tissue extends only to the constriction, beyond which lies
the micropylar region Mdth its wandering endosperm nuclei.
An examination of the integument cells surrounding this region
shows that dissolution is in progress — due, of course, to the
predatory activity of this part of the sac. The antipodal por-
tion, with its endosperm tissue, exerts a similar absorptive
influence, but the material absorbed by the latter is retained or
used up in building more tissue, while that absorbed by the
former is not, but is transferred to the endosperm tissue. The
micropylar portion of the embryo sac may therefore be re-
garded as a haustorium. Evidence of the results of haustorial
activity is seen in the unusual richness of plasmatic contents
in the endosperm cells bordering on the haustorium. The pres-
ence of starch in the basal portion points to the same conclu-
sion.

The nuclei of the haustorium do not form tissue, as has al-
ready been stated. In Lamium ample xicaule they take no
definite position. Although they have the same origin as the
nuclei of the endosperm tissue they differ considerably from
them. The chief difference lies in their much greater size.
These may attain a length of over thirty micromillimeters.
which is greater than the average diameter of an endosperm

billings: embryo nutrition in labiate. 71

cell (fig. 5). The haustorial nuclei vary usually from twenty
to thirty micromillimeters in diameter, but the smallest are
much larger than any formed in the endosperm tissue, since
these average only about five micromillimeters in diameter.
Another difference is the much greater size of the nucleoli of
the haustorial nuclei. In addition, the nucleoli are not vesicu-
lar, as are most of those in the endosperm tissue, but are solid.
The large size of the nucleoli would seem to indicate an in-
creased nutritive function on the part of the nucleus, especially
in light of the theory that nucleoli represent reserve food sub-
stance. There is no evidence that the haustorial nuclei undergo
division, either in Lamium or in the coenocytic haustoria of any
other of the Labiatsg investigated. There is some variance in
the explanations of the peculiarity of these nuclei, but dis-
cussion of them will be reserved till the close of the paper.

The suspensor elongates during the growth of the embryo
sac and finally attains an unusual length (fig. 6). Its course
is for some time plainly visible in the haustorium and for a
longer time in endosperm tissue. The portion nearest the em-
bryo persists while the remainder slowly disintegrates. The
appearance of the suspensor is such that at no time does it
seem to serve as a conductor of food substances to the embryo,
at least from any locality other than that of the endosperm
tissue immediately surrounding it. It appears to have fulfilled
its function when it carried the embryo from the haustorium
down into the midst of developing endosperm tissue.

While the micropylar haustorium is transferring materials
to one end of the endosperm tissue the opposite end is nour-
ished by substances conducted through a vascular bundle that
passes down the integument and terminates near the chalaza.
The little depression or pocket in which the antipodals undergo
disintegration is deepened until it forms a short canal, which
extends from the endosperm toward the terminus of the vas-
cular bundle. It does not merit the name of haustorium to the
extent that the micropylar expansion of the sac does, since it
exerts little or no absorptive influence ' upon the integument,
cells bordering it, but rather seems designed to facilitate con-
duction from the bundle. Its increase in length is not accom-
panied by a pronounced increase in diameter nor does it contain
nuclei.

Later stages in ernbryo development show increase of the

72 KANSAS UNIVERSITY SCIENCE BULLETIN.

endosperm tissue and a consequent encroachment upon the
haustoria. Just when the micropylar haustorium ceases to
function is hard to say. Its activity probably does not cease
as soon as it begins to be encroached upon by the endosperm
tissue. The nuclei in fact remain till late stages in seed de-
velopment.

Stachys palutris.

The embryo sac seen in figure 7 presents markedly differen-
tiated portions. The stage shown here is reached in the same
manner as a corresponding one in Lamium. While the endo-
sperm is beginning its formation in the antipodal end of the
sac a number of nuclei migrate into the haustorium. They
distribute themselves over the micropylar end of the hausto-
rium, where the cytoplasm is chiefly located. The opposite end
is distinctly vacuolated. It will be noticed that the constric-
tion is very narrow. Through it passes the suspensor with an
enveloping layer of cytoplasm. The regions adjacent on each
side of the constriction are stored with starch, the result evi-
dently of the absorptive activity of the haustorium. As in
Lamium, the cells of the endosperm tissue nearest the hausto-
rium are richer in plasmatic contents than those at a distance.

The canal that conducts from the terminus of the vascular
bundle is comparatively long in Stachys. It is sharply bent
about midway in its course, one arm running longitudinally to
connect with the end of the vascular bundle.

Stachys and Lamium vary in the structure of their embryo
sacs only in minor details, the main difference being the more
definitely placed haustorial nuclei in Stachys. The embryo is
nourished in both by the endosperm tissue surrounding it, this
in turn being assisted in its growth by the activity of the
haustorium. It is probable that the suspensor does not func-
tion in the conduction of materials to the embryo, on account
of the disorganized condition of its cells. The food materials
gathered by the haustorium are transmitted to the endosperm
tissue through that part of the constriction lying around the
suspensor.

Phrysostegia virginica.

This species has a peculiar appearing embryo sac when the
stage shown in figure 8 is reached. Such difference as may be
noted, when it is compared with Lamium and Stachys, is due
principally to the direction of growth taken by the endosperm

BILLINGS: EMBRYO NUTRITION IN LABIATE. 73

tissue. If the growth had been uniform in all directions the
long axes of the haustorium and endosperm tissue would have
been approximately parallel. In Phrysostegia the endosperm
grows in a one-sided fashion, producing a bend in the embryo
sac of about ninety degrees. The haustorium is a club-shaped
sac containmg a few large nuclei. Its upper or micropylar end
is vacuolated, while its lower end, adjoining the endosperm
tissue, is densely protoplasmic. The suspensor is exceedingly
long.

The position of the antipodal canal is peculiar in this species.
It is located near the haustorium — a situation explained by the
short length of the vascular bundle, as well as the one-sided
growth of the endosperm tissue. The constriction existing be-
tween the haustorium and endosperm tissue is comparatively
wide, with the result that the cytoplasm of the haustorium
comes in contact with the endosperm tissue to a greater extent
than in any other species examined.

Leonurus cardiaca.

Leonurus has a large haustorium with a contour much like
that of Stachys (fig. 9) . The haustorial nuclei have a more or
less fixed position in that portion lying nearest the endosperm
tissue. This is the reverse of the condition in StacJujs, where
the nuclei lie in the micropylar region. Their number varies
from four to six. The definite position of the haustorial nuclei
would suggest the presence of cell walls, but there was no evi-
dence of them.

Leonurus bears a close resemblance to Phrysostegia in the
proximity of the antipodal canal to the haustorium. The canal
does not appear in the section shown in figure 9, so its position
is indicated by the dotted lines. The lower portion of the endo-
sperm tissue and its connection with the vascular bundle
through the canal is shown in figure 10.

BruneUa vulgm'is.

This species possesses a large coenocytic haustorium contain-
ing nuclei without definite position. In this respect it is like
Lamium. The antipodal haustorial canal becomes laterally
placed, but, unlike the two previously described genera, it re-
mains near the base of the endosperm tissue.

The five genera thus far mentioned belong to the tribe
Stachydese. No marked variations occur; in fact, they are

74 KANSAS UNIVERSITY SCIENCE BULLETIN.

• much more nearly alike than they are different. Phrysostegia
appears to be the most divergent, but such divergence is really
trivial. All are characterized by a large micropylar hausto-
rium — a feature, however, which it shares with representa-
tives of other tribes.

Teucrium canadense.

A glance at the diagram shown in figure 11 will show the
relative position and size of the different portions of the em-
bryo sac. The most noticeable difference between this species
and those previously described is the relatively small size of the
haustorium. The failure of the haustorium to enlarge much
is doubtless mainly due to its proxity to the end of the beak-like
extremity of the integument. There is but little room for en-
largement. The haustorium contains a few nuclei that do not
form a tissue. Whether this organ is really functional or not
is difficult to determine with certainty. At most its activity
can be only very slight, otherwise the extremity of the integu-
ment would undergo dissolution. The endosperm nuclei which
are found in it do not have the size nor the altered appearance
seen so frequently in vigorously acting haustoria. It would ap-
pear from this that alteration in the haustorial nuclei is to
some extent at least proportional to the activity of the organ
to which they belong.

Pycnanthemum lanceolatum.

The general configuration of the embryo sac is much like
that of Laminm (fig. 12). The ccenocytic haustorium attains
a large size, with the result that most of the micropylar por-
tion of the integument becomes absorbed. The constriction
between haustorium and endosperm tissue is soon obliterated,
so that the two lie contiguous over a considerable area. The
suspensor is long, as in the previously mentioned species, but
owing to the size of the haustorium most of its length is found
within the limits of this structure.

Lycopus riibellus.

LycopuH ruhelliis, when contrasted with Pycnanthemum,
which belongs to the same tribe, differs in two respects; the
haustorium is small, and the constriction lying between it and
the endosperm tissue is relatively long (fig. 13). The basal
portion of the endosperm tissue extends in a beak-like process
to the end of the vascular bundle. This process occupies the

BILLINGS: EMBRYO NUTRITION IN LABIATE. 75

position of the antipodal canal of previously described species.
Its origin may be explained by a growth of the tissue into the
canal, which it thus displaces. The haustorium contains but
one or two nuclei with no formation of tissue.

Mentha canadensis.
The integument in this species is not extensive, yet there is
a large cosnocytic haustorium present which closely resembles
that of Pijcnayithemiim. Like Pycnanthemiim, there is in
Mentha an early obliteration of the canal connecting hausto-
rium and endosperm tissue. Lycopus, on the other hand, rep-
resents the other extreme, in which the canal persists for a
long time. PycnantheTmim, Lycopus and Mentha belong to the
tribe Satureinese.

Salvia azurea and Salvia lanceolata.

These two species will be considered together, as they pre-
sent many points in common. As compared with the repre-
sentatives of other genera, however, they show marked differ-
ences. In the first place, they are unique in having two haus-
torial outgrowths, one coenocytic and one composed of ordinary
endosperm tissue. In addition, there is a well-developed antip-
odal canal.

In order to follow better the development of the haustoria, it
will be advantageous to examine the definitive embryo sac and
consider some of the phenomena immediately following fertili-
zation. The definitive embryo sac of Salvia lanceolata is shown
in figure 14. It is short, and in comparison with the extensive
integument is relatively insignificant (fig. 15). The shortness
is the more noticeable when the amount of space occupied by
the egg apparatus is taken mto consideration. The embryo
sac is filled with a reticulated cytoplasm, which is distinguished
from that of Lamium amplexicaule and other species examined
by the absence of vacuoles. The antipodals lie in the depression
which marks the beginning of the canal to the vascular bundle.
The small size of the embryo sac necessitates the proximity of
the endosperm nucleus to the egg apparatus.

Fertilization is followed by a very slight elongation of the
egg cell — a marked variation from the condition found in the
species of other genera examined. The explanation for this
difference lies in the fact that none of the space occupied by the
definitive embryo sac is to be included in haustorium formation,

76 KANSAS UNIVERSITY SCIENCE BULLETIN.

since elongation of the egg and suspensor is always for the pur -
pose of removing the embryo from the sphere of haustorial
activity. In Lamium approximately half of the sac becomes
haustorium, and it is in view of this fact that the embryo is
transferred for development to the antipodal region. The
elongation of the suspensor in the majority of Labiatse exam-
ined appears to terminate its function, for disintegration fol-
lows. In Salvia, on the contrary, there is reason to believe that
it plays an important part in embryo nutrition. The cells com-
posing it are short and relatively rich in plasmatic contents.

The first division of the endosperm nucleus results in the
formation of a cell at the extreme antipodal end of the embryo
sac. The wall cutting off this cell lies approximately in a plane
at right angles to the long axis of the embryo and suspensor.
The separating walls of cells produced in succeeding divisions
are at first parallel to the first wall. The embryo is soon sur-
rounded by endosperm tissue (fig. 16). One endosperm cell
takes a position near the end of the suspensor for the purpose
of entering directly into the formation of a haustorium. The
growth of the endosperm tissue surrounding the embryo is
greatest in a direction toward the upper part of the integument
and approximately parallel with the vascular bundle. As a re-
sult, an endosperm process, cylindrical in shape, projects into
the region where lies the most extensive portion of integument
tissue (fig. 17). The outgrowth is evidently designed to reach
the nutritive tissue remote from the main mass of endosperm.
Functionally, it is a haustorium, differing principally from
tissue-containing haustoria of certain other families of plants
in the greater number of cells. Figure 17 shows a young haus-
torium of Salvia lanceolata. In its maturer stages, seen dia-
grammatically in figure 19, it is composed of much elongated
cells. Such differentiation suggests conduction as a function.
The elongated cells lie principally in the axis of the haustorium
with their long axes parallel to that of the haustorium. The
peripheral cells serve as digestive and absorptive cells, while
those of the interior of the haustorium are conductive. The
products of its activity are transferred to the main mass of
endosperm which it thus serves as an absorbing organ.

The other haustorium is coenocytic, and its activities are
directly concerned with embryo nutrition through the suspen-
sor. It is not a structure developed from the suspensor, though

billings: embryo nutrition in labiate. 77

it continues it, in a way, out into the integument. It recalls
some of the haustorial outgrowths in the suspensor cells in
certain Orchidacese- and Rubiacese."' In Salvia lanceolata two
endosperm nuclei take part in haustorium formation; more
than two in Salvia azurea (fig. 18). In both species one end of
the haustorium completes the basal or proximal portion of the
suspensor. In Salvia lanceolata more of the suspensor is sur-
rounded than in Salvia azurea. In the former the suspensor
becomes bent while in the latter it remains straight.

The haustorium grows outward through the integument,
taking a direction nearly at right angles to that taken by the
other haustoria. The purpose of the coenocytic haustorium
is to dissolve the integument cells surrounding it, but it is
unique in communicating the products of its activity to the
suspensor. Attention has already been called to the striking-
differences between the Salvia suspensors and those of the
other Labiatse examined. As the former take part in embryo
nutrition their cells present an appearance betokening activity.
Instead of losing their protoplasmic contents or undergoing
dissolution, as in instances where excessive elongation takes
place, the suspensor cells of Salvia are healthy appearing, re-
sembling the endosperm cells in this respect not a little.

The coenocytic haustorium does not attain the size of the one
composed of tissue (fig. 19). As in similar haustoria previ-
ously described, the nuclei are larger than those of the endo-
sperm tissue.

A canal, bent at nearly right angles, is formed between the
endosperm and the vascular bundle. There appears to be no
nucleus in it, but granular material resembling cytoplasm is
found along the side walls and at the end. As the main body
of endosperm pushes outward at the expense of the integument
cells, the canal does not suflFer for a time (fig. 20). Instead,
the endosperm becomes laid down around its proximal end.
This gives the appearance of the canal piercing the tissue. The
canal forms a direct communication with the vascular bundle,
serving probably as a conduction to the endosperm.

Monarda flstulosa.

This species is patterned after Lamium rather than after
Salvia. There is a large coenocytic micropylar haustorium
which does not diflfer materially from that of Lamium. A few
nuclei are present, though of a relatively smaller size. The

78 KANSAS UNIVERSITY SCIENCE BULLETIN.

embryo sac is narrow and the suspensor is very long. There is
no antipodal canal, but the endosperm tissue bends toward the
terminus of the vascular bundle in the form of a process.

Nepeta cata7'ia.

The haustorium of Nepeta cataria is coenocytic, and contains
four nuclei that occupy a more or less definite position at its
base or near the place where the suspensor passes into the con-
striction (fig. 21). The haustorium is relatively small, con-
taining much less than half the length of the suspensor.

Dracocephalum sp.

This varies from the other species of Labiatse examined in
the character of the outline of the endosperm tissue. In the
other species this tissue enlarges quite uniformly, presenting
a smooth or entire outline when sectioned. The outgrowths
in Dracocephalum have no definite position or form and hence
are not to be regarded as other than integral portions of the
endosperm tissue and not haustoria (fig. 22). The haustorium
is much elongated, as is the endosperm tissue. The embryo lies
in the upper portion of the latter, being attached at the end of
a very long suspensor.

Scutellaria parvula.

The bent embryo sac of Scutellaria, resulting in a bent em-
bryo in the ripe seed, is of course well known to taxonomists.
It conforms to the other species in Labiatse in possessing a
haustorium, but located, as it is, at the extreme tip of the ovule,
its growth is greatly limited, so that it remains small (fig. 23).
The endosperm is elongated and tapers at the base into a hook-
like process whose end lies near the terminus of the vascular
bundle.

DISCUSSION AND CONCLUSIONS.

In a large number of plants, especially those in which two
integuments are present, the embryo sac assumes a somewhat
elliptical or oval form in longitudinal section, with an approxi-
mately entire outline. The nutrition designed for growth of
the endosperm may be absorbed fairly uniformly over the en-
tire surface of the sac, or it may be absorbed more rapidly at
one point than at other points, but in either case the elliptical
or oval outline is maintained. If added organs mean increased
specialization, then embryo sacs of the type just mentioned,

BILLINGS: EMBRYO NUTRITION IN LABIAT^E. 79

without special absorptive structures, may be regarded as a
simpler, less differentiated, perhaps more nearly primitive,
sort.

In some instances the innermost cell layer of integument,
which borders on the nucellus or else on the embryo sac, is
specialized into a digestive layer, whose activity is directed
against the remaining integument for the benefit of the em-
bryo sac. The name "tapetum" has been frequently applied to
this layer, but its use is justified only on account of the nutri-
tive function it performs. GoebeP proposes the term "epithe-
lium" as a substitute. Three characteristics usually pertain to
the epithelium: first, it remains intact for a longer period
than the integumental layers adjacent; second, its cells be-
come differentiated in form or contents from those of the re-
maining integument ; third, it apparently exerts a digestive ac-
tion. The digestive action is doubted by some, among whom is
Schmid," who ascribes a protective function to it. Since the
species of Labiatse herein described do not develop an epithe-
lium, it is without the province of this discussion to do more
than call attention to the fact that Schmid challenges the re-
sults obtained by the "Goebelische Schule," thus showing that
the problem is open for further investigation.

The Labiatse, in common with some other sympetalous fami-
lies, depart from the type described for many plants having
two integuments by differentiating haustoria. The haustorium
implies increased absorptive activity in one portion of the
embryo sac, but not necessarily cessation of the function else-
where. Among the labiates studied, all (with the single ex-
ception of Salvia) have a micropylar haustorium of some sort.
In most of the species it is a well-developed structure, exhibit-
ing manifest activity as a nutritive organ. But in such species
as Scutellaria parvula, Teucnum canadense, and Lycopus ru-
bellus, it is of small size, and its usefulness may with safety
be said to be either greatly reduced or else obsolete. Whether
such a condition really represents actual reduction from a pre-
viously more highly developed state, or whether it is primitive
in its nature, is a problem yet to be solved. The much elon-
gated suspensor is here offered as evidence favoring the former
view, since the purpose of elongation is believed to be the re-
moval of the embryo from the region of the haustorium. Ex-
traordinary length in proportion to the size of the haustorium

80 KANSAS UNIVERSITY SCIENCE BULLETIN.

might therefore mean that shortening of the former did not
accompany reduction in size of the latter. Conditions external
to the embryo sac may have had something to do with hausto-
rium reduction in such a species as Scutellaria parvula, where
narrowing of the integument to a beak left but little tissue to
be exploited.

The nuclei in the coenocytic haustorium merit interest on
account of the extraordinary size and appearance some of them
attain. Such nuclei are found in other families where similar
haustoria exist. They have been described by Balicka-Iwanow-
ska,'' Schmid'' and others. That they are transformed endo-
sperm nuclei is of course easy to observe. Knowing the kind of
work the haustorium is called upon to do, one might expect in-
crease in size, both of entire nucleus and also of nucleolus, on
account of extraordinary nutritive function. The loss of power
to divide accompanies the development of this function. Ac-
cording to Schmid,^ the nuclei should be considered degenerate
structures, whose condition has been produced by excessive
nourishment, as is attested by loss of power to divide. In-
stances of like nature are not limited to embryo sacs, but occur
in mycorrhiza and in animals. There is room for difference of
opinion, however, as to the governing principle underlying such
nuclear change. It is at least open to investigation whether
the increase in size is a direct result of degeneration, due to
unusually favorable conditions regarding food supply, or
whether it is due to the added functional responsibility de-
manded of them by the haustorium. It is questionable whether
either failure to undergo division or exceptional abundance of
nutriment should be used as a test of degeneracy. As evidence
against the latter view, attention is called to the normal ap-
pearance of the nuclei in the cellular haustorium in Salvia and
those in the endosperm cells adjacent to a coenocytic hausto-
rium. Both are in very favorable condition as to food supply,
yet there is no nuclear enlargement. It would appear that
coenocytic structure is conducive to such enlargement, but it
cannot be merely a question of food supply, A difference is to
be noted in the lack of cell walls, which in a tissue may pos-
sibly exert a limiting influence on increase of size in nuclei.
Another difference is seen in the relatively small number of
nuclei in a coenocytic haustorium. It might be difficult to
prove, however, that for this reason the work demanded of each

BILLINGS: EMBRYO NUTRITION IN LABIATE. 81

nucleus would be greater, though of course there is a possi-
bility that it might be true.

Haustoria have either indefinite or more or less definite
positions of their nuclei. Wallc might be expected in instances
where the nuclei are definitely placed. Balicka-Iwanowska''
found a very delicate gelatinous membrane present in cer-
tain instances of this sort. No walls were evident, however,
in any of the micropylar haustoria of the Labiatse.

Indefinite location of nuclei are associated with irregular
form of haustorium, while those having a more nearly fixed
position are found in such as have an approximately symmet-
rical form.

The Labiatse investigated generally have long suspensors,
which are associated with the presence of micropylar haus-
toria. An exception is seen in Salvia, which has a short sus-
pensor and no micropylar haustorium. The direct results of
suspensor elongation are the removal of the embryo from the
haustorium and its transfer to the vicinity of the center of the
growing endosperm tissue. These results are doubtless suffi-
cient to serve as causes for suspensor elongation. It is at least
evident that a position within the haustorium would not be as
favorable as a position without in so far as the products of
haustorial activity are concerned. The endosperm tissue itself
exerts a digestive action on the integument, and it also receives
the nutriment from the haustorium, hence this tissue is more
favorable for embryo development than the haustorium alone.
Although integument cells are destroyed by the haustorium,
there is no reason for believing that those of the embryo would
likewise be, since endosperm tissue bordering on the hausto-
rium is in no way affected by its digestive action. There is,
therefore, no reason for thinking that suspensor elongation is
to remove the embryo from the predatory action of the haus-
torium.

In general, it does not appear that the suspensor functions
as a conductor of nutrition to the embryo. Evidence for this
view is found in the collapsed condition of the suspensor
through nearly its entire length, especially in that part lying
within the haustorium. The greater portion of it is apparently
without cell contents of any sort, and may or may not be con-
tinuous from the micropylar end of the embryo sac to the
embryo. It would be expected of a cell or tissue that is serving

3-Univ.[Sci. Bull., Vol. V, No. 5.

82 KANSAS UNIVERSITY SCIENCE BULLETIN.

actively as a means of translocation of food materials that some
vestige of life would be discernible. In Salvia, for instance,
a healthy, vigorous condition of the suspensor cells accompanies
activity in food transmission. In all the species examined
(except Salvia), the only suspensor cells that appeared to be
living were those immediately adjacent to the embryo. The
nutrition of the embryo is accomplished through absorption
over its entire surface, whereas in Salvia an added source of
nutriment probably lies in the conducting power of the sus-
pensor.

The length of the suspensor is roughly proportionate to the
size of the haustorium through which the embryo has to be
transported. Usually the movement is past a coenocytic struc-
ture to a tissue. An instance is to be noted, however, in My-
oporum^ (Myoporacese) , in which the growth is directly
through one portion of the endosperm to another. The beaked
portion of the endosperm is apparently a remnant of what may
once have been a coenocytic haustorium, but in which the coe-
nocytic condition has been lost.

The integument varies in thickness and extent in the differ-
ent species. Of course, it is in those in which it is most ex-
tensive that the largest haustoria are possible. Through the
integument in each case there runs a vascular bundle which
terminates near the antipodal region. A slight outpocketing
of the embryo sac is found extending into the integument to-
ward the end of the bundle. Later it may develop into a canal
without nuclei, or into a process of the endosperm. Its func-
tion in all cases is evident — to facilitate the passage of nutri-
ment from the vascular bundle to the endosperm.

In summing up the characteristics that are common to the
species described in this paper, with the exception of Salvia,
they have been found to possess three that in the future may
be shown to belong to the family as a whole. They are the
micropylar haustorium, the much-elongated suspensor, and the
antipodal canal or process. Marked variations from a common
type, such as are found in Salvia, suggest a taxonomic re-
arrangement, even though the usual macroscopic characteris-
tics resemble those of the family in which the varying species
is placed. In a previous paper,' for instance, attention was
called to the striking difference in the ovular structure of the
Menyantheae when compared with the coordinate subfamily

BILLINGS: EMBRYO NUTRITION IN LABIATE. 83

Gentianese. The two have since been raised to family rank,
probably for other reasons, yet it is interesting to note that
the structure of the ovule justified the wisdom of the change.
In this way can a study of sporangial, gametophytic and em-
bryonic structures be of service in establishing more natural
relationships and groupings.

LITERATURE CITED.

1. F. H. Billings. Beitrage zur Kenntniss der Samenentwickelung.

Flora Bd., 88 Heft 3, 1901.

2. M. Treub. Notes sur I'embryogenie de quelques Orchidees, Natur-

kund. Verhandl. Konigl. Akad. Deel. 21, 1879.

3. F. E. Lloyd. The comparative embryology of the Rubiaceae. Mem-

oirs of the Torrey Botanical Club, vol. VIII, No. 1, Pt. 2, 1902.

4. K. GoEBEL. Organographie der Pflanzen, vol. II, Jena, 1901.

5. E. SCHMID. Beitrage zur Entwickelungs-geschichte der Scrophulari-

aceae, Zurich, 1906.

6. G. P. Balicka-Iwanowska. Contribution a I'etude du sac embryon-

naire chez certaines Gamopetales, Flora 86, 47-71, 1899.

THE

KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 6— October, 1909.

(Whole Series, Vol. XV, No. 6.)

CONTENTS:

Mitosis in the Root-tip Cells of Podophyllum peltatum, A. Richards.

PUBLISHED BY THE UNIVERSITY,
LAWRENCE, KAN.

Entered at the post office in La\vrenc(> as .second-class matter.

THE KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 6] OCTOBER, 1909. [vJI°xVnT6

MITOSIS IN THE ROOT-TIP CELLS OF PODOPHYLLUM

PELTATUM.

BY A. RICHARDS.

Plates XV and XVI.

(Contribution from the Botanical Laboratory of the University of Kansas.)

''pHE work, the results of which are herein presented, was
*■ undertaken during the summer of 1907, under the direc-
tion of Prof. W. C. Stevens, whom I have to thank for his kind-
ness and much helpful advice. My thanks are also due Mr.
F. A. Hartman for helpful suggestions.

In discussing the root-tip cells of Podophyllum peltatum, 1
shall use the following outline :

I. Methods.
II. Resting cells.

A. Nucleus.

1. Nuclear membrane.

2. Chromatin.

3. Achromatin.

4. Nucleolus.

B. Cytosome.

1. Cell wall.

2. Cytoplasm.

3. Archoplasm.

III. Dividing cells.

A. Prophase.

1. Nucleus.

2. Cytosome.

B. Later phases of division.

1. Chromatin.

2. Achromatin and cytoplasm.

IV. General considerations.

A. Achromatic and archoplasmic structures.

B. Individuality of chromosomes.

(87)

88 KANSAS UNIVERSITY SCIENCE BULLETIN.

METHODS.

The root-tips of Podophyllum peltatum are peculiarly fitted
for the study of mitosis in the vegetative cells of plants. The
cells are larger than those of the onion root-tip, and in properly
stained material the cell organs, especially the chromosomes,
are to be seen with almost diagrammatic clearness.

The material used in this study was collected during the lat-
ter part of March and fixed in Flemming. For staining the
sections three methods were used : Haidenhain's iron-hsema-
toxylin (some sections counterstained with orange G), the
safranin-gentian violet-orange G stain, and a combination of
these two. Satisfactory results were obtained with slides
stained in haematoxylin, both with and without the counter-
stain ; the three-color method also gave fair results. Only a
few sections were successful with the combination method,
but they were promising; without doubt the method will
repay careful experiment. It was followed as given by Schaff-
ner (2).

THE RESTING CELL.

The most typical resting cells are not often to be found at the
tip of the root, for this, of course, is the region of greatest
mitotic activity and the resting stages are very short, but back
from the tip as much as two millimeters. Farther from the
tip the cells are apt to be plasmolyzed in fixing.

The nuclear membrane of the resting cell normally disap-
pears as soon as the cell begins division ; it is, however, quite
constant in its appearance and staining reaction, and varies
only slightly in thickness. It appears to stain very deeply with
chromatin stains, a fact which is probably due to the peripheral
arrangement of the chromatin granules, as seen in figure 1.
These chromatin masses are arranged over the linin network of
the nucleus in large or small patches, depending on the age of
the cell, i. e., nearness to the tip.

In examining the achromatic structures of the cells of P.
peltatum I have had considerable difficulty, especially in the
stages of mitosis. In some cells they can be seen quite clearly ;
in others, which appear to be equally well fixed and have been
treated with exactly the same methods, they cannot be seen at
all. Owing to this irregularity in their staining capacity, I
have not been able to work out their history accurately.

The nucleoli of these cells are very interesting. They vary

RICHARDS: MITOSIS IN PODOPHYLLUM. 89

in number from one to several — rarely more than four — and in
size and shape. In general they are irregular-spherical, but
may be compressed into different shapes. The variation in size,
which is considerable, depends on the age of the cell. As the
time for division approaches they grow smaller, finally disap-
pearing about the same time as the nuclear membrane. This
last fact sustains the conclusion that the nucleolus is simply a
storehouse for food material for the rest of the cell.

In most cells up to the time of division there is noticeable a
peculiar vacuolated condition of the nucleus, and in these vac-
uoles lie the nucleoli. Figure 1 shows two such cells, one of
which has a single large nucleolus surrounded by a vacuole
and the other three. The term "vacuole" for the clear space
surrounding the nucleoli may be misleading. Experiments
seem to show that this colorless space contains something more
than a watery fluid, which is extracted by dehydration. By sub-
jecting the roots of Zea mays, Vicia faba, and other plants to
strong centrifugal force, Mottier ( i ) found that the nucleolus
with its surrounding clear space was thrown out of the nucleus
into the cytoplasm. "The nucleolus was still surrounded by the
space, a fact that seems to show that the colorless substance
has greater specific gravity than the other contents of the
nucleus. The colorless substance may represent the unor-
ganized nucleolar material."

The cell wall of the resting cells stains much like the nuclear
membrane and is only slightly thicker.

The cytoplasm stains fairly dark with iron hsematoxylin,
and shows its fine reticular structure quite plainly. Scattered
uniformly over the threads of the reticulum are large granules.
The cytoplasm of the cell stains slightly denser than the nuclear
sap.

With regard to archoplasmic structures, not the slightest evi-
dence was seen to indicate that such structures as have been
described by Schaffner are ever present : /. e., centrosomes and
centrospheres.

DIVIDING CELLS.

The plane of division in these cells is mostly at right angles to
the axis of the root. This rule, however, is by no means in-
variable, for numerous cases have been found with the division
planes parallel to, or at an angle of forty-five degrees with, the
longitudinal axis (figs. 14, 20).

90 KANSAS UNIVERSITY SCIENCE BULLETIN.

When the nucleus begins division the chromatin masses ar-
range themselves in a spireme (figs. 2, 3) and the linin reticu-
lum can no longer be seen. Perhaps the network has draw^n
tighter, the threads stretching out to form a framework for the
spireme. This stage has been called "close mother skein"
(Schaffner) . In figure 2 the nucleoli are still present, but they
are now without the surrounding clear spaces.

The spireme soon becomes looped with the heads pointing
toward the poles. I have not observed any connection between
this fact and the fact that the chromosomes are J- and U-
shaped. The segmentation of the spireme does not seem to
occur regularly near the equator of the nucleus, as would be
necessary to make the heads of the chromosome loops identical
with those of the spireme loops.

At about this stage the nuclear membrane disappears, as do
the nucleoli. There is, of course, variation about the time of
disappearance, for the membrane is still present in figure 4,
where the granules have split to form a double spireme. This
split is very clearly shown in figures 5 and 6. The spireme
granules here have segmented to form the chromosomes. Im-
mediately after segmentation the chromosome granules con-
dense and the segments shorten, thus giving the apparently
solid character to the chromosomes. In figure 6 the two lower
segments have lagged behind the others in condensation.

At this stage or earlier the milage of the spindle should ap-
pear, for in the next stage observed the chromosomes were
arranged on the bipolar spindle in the metaphase. The origin of
the spindle was not observed, however, nor were any traces of
a multipolar spindle seen. This may be due to any one of the
three causes : the duration of the multipolar spindle may be
short ; the spindle substance at this stage may not take the stain
readily; or the bipolar spindle may have been formed in the
manner usual in the cases of higher plants, but with the omis-
sion of the multipolar stage. Of these three hypotheses for the
absence of the multipolar stage, the third seems, perhaps, the
best from the evidence in hand. The probability of the hy-
pothesis decreases when we consider that the central spindle
fibers stain more deeply than the traction fibers. No centro-
some or centrosphere was seen in these cells, although in one
or two cases a massing of protoplasmic granules at the poles
of the cells might have been taken on a superficial examination
for a centrosphere (figs. 7, 11). It was not difficult to de-

RICHARDS: MITOSIS IN PODOPHYLLUM. 91

termine, however, that such was not the case, and it seems quite
certain that the spindle is formed without the influence of
other archoplasmic structures.

In most cells there is no metaphase for the entire chromo-
some mass, ?'. e., an equatorial plate, in these cells, but each
chromosome passes through the stage without waiting for the
others. Figures 7 and 11 are the only approach to such a con-
dition, and even here there is some lack of uniformity in the
division of the chromosomes. Figures 8, 9 and 10 show the
usual condition of the equatorial plate. Figure 10 shows the
longitudinal split of the undivided chromosomes very clearly.
This cell was crushed, but the crushing served to bring out the
split all the better.

During the metaphase the first signs are seen of the concen-
tration of the cytoplasm at the center of the cell, where the
new wall is to be formed, and the consequent vacuolation of the
cytoplasm at the poles. (This phenomenon is the usual occur-
rence in these cells but there are numerous instances where it
is only slightly marked.) This concentration seems to aid in
the construction of the new cell plate, for the more marked the
concentration the more rapid the cell-plate formation.

In regard to the division of the chromosomes nothing un-
usual was observed. The chromosomes divide and pass to the
poles as in most forms. In passing up to the poles the chromo-
somes usually turn with the short arm of the J toward the cen-
ter of the spindle. One chromosome seemed to stand out from
the rest slightly and was especially marked by its size ; this was
the long chromosome m (figs. 9, 14). Polar views of the ana-
phase present the best opportunities for counting the chromo-
somes, but there is much uncertainty even here, owing to the
diflficulty of getting all of the plate in one section or of tracing
it through neighboring sections. Fourteen chromosomes were
counted in about a dozen cells, but this, of course, is insuffi-
cient evidence on which to base a decision.

Arrived at the poles the chromosomes fuse end to end and
then form a loose daughter "skein." This "skein" becomes a
close one and the chromatin forms a dense mass at the poles.
The remainder of the nuclear reconstruction consists in the
formation of the membrane, the distribution of the chromatin
granules in the nuclei, the appearance of the linin network and
the formation of new nucleoli. It is a general rule that the
nucleoli arise on the side of the cell nearest the new cell wall.

92 KANSAS UNIVERSITY SCIENCE BULLETIN.

While the reconstruction of the nuclei has been in progress,
several changes have occurred in the spindle and the cytoplasm.
As soon as the concentration of the cytoplasm is complete the
spindle broadens out somewhat. Figure 23 shows a maximum
bulging, while figure 22 is perhaps a minimum. Figure 22
shows an unusual phenomenon in the process; the majority of
the central spindle fibers (only the central fibers now remain)
seem to have become free at their polar ends and to have short-
ened towards the center, staining very darkly in this region.
The threads disappear as soon as the cell walls are formed. It
is difficult to determine what the bulging out of the spindle and
the concentration of the threads and cytoplasm signify, but it
seems probable that the threads contain some substance useful
in the construction of the new cell wall. This suggestion is in
line with the work of Strasburger (4) and Haberlandt (5),
who have discussed this phenomenon of the spindle in detail.
The rate at which these processes occur varies. One may
find a nuclear membrane before the cell plate has started (fig.
20), or he may find the cell plate well started while the chro-
matin is still in the loose daughter "skein" (fig. 19).

GENERAL CONSIDERATIONS.

Ach7'omatin and Archoplasm. — In regard to the origin of
the spindle Schaff'ner concludes that, in root-tips of the onion
and of Sagittaria, at least, the multipolar spindle is due
either to pathological causes or to ''improper manipulation
in preparing the sections," and that a bipolar spindle is formed
not by the transformation of a multipolar stage but under the
influence of a centrosome and centrosphere as commonly de-
scribed for animals. As previously stated, I find no evidence
whatever for concluding that such archoplasmic structures are
present.

Although no traces of a multipolar spindle have been seen
here, it is not necessary to assume that a centrosome is present
to guide the formation of the bipolar spindle, as the fibers them-
selves may possess sufficient directive energy to bring about
the bipolar form.

Individuality. — A single bit of evidence was found in support
of the theory of individuality of the chromosomes in vegetative
cells. In several of the cells there is an unusually long chromo-
some, m, in general dividing before the others in the cell. Al-
though this meager bit of evidence is by no means conclusive.

RICHARDS: MITOSIS IN PODOPHYLLUM. 93

it points toward the view that while the cell is the unit of
structure the chromosome is the unit of potentiality, and that
the chromosomes themselves may all differ in structure and, if
in structure, also in potentiality.

An interesting problem concerning these root-tip cells is the
relation of the cells to the entire tissue and the cytological
processes of later stages of embryonic growth. Material for
this study is not at hand at this time, but I hope to be able to
trace out these processes at some subsequent period.

BIBLIOGRAPHY.

1. D. M. MOTTIER. Fecundation in Plants, Carnegie Institute, 1904.

2. J. H. SCHAFFNER. Karyokinesis in Root-tips of Allium cepa, Botanical

Gazette, 36:226.

3. Adolph Hof. Botanisches Centralblat, LXXVI, 65, 113, 166, 221.

4. Eduard Strasburger. Ueber Plasmaverbindungen pflanzlicher Zellen,

Jahrbiicher fiii- wissenschaftliche Botanik, 1900.

5. Haberlandt. Function und Lage des Zellkernes — Jena, 1887.

NOTE.

The drawings of the Podophyllum root-tips were all made with the aid
of a camera lucida (B. & L.) A C. Reichert Mo objective was used and
with it a B. & L. i/^-inch ocular for the larger figures and a %-inch for the
smaller, making a magnification of 2466 and 2000 diameters, respectively.

Figures 9, 13, 14, 15, 16, 18, 20, 22, 23, 24, were made from material
stained with the Flemming three-color method. The others were made
from sections stained with the iron-haematoxylin method.

2-Univ. Sci. Bull.. Vol. V. No. 6.

THE

KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 7— October, 1909.

(Whole Series, Vol. XV, No. V.)

CONTENTS:

KOEBERLINIA SPINOSA ZUCC. : AN ECOLOGICAL STUDY OP THE ANATOMY

OF THE Stem and some other Parts, .... Miriayn Sheldon.

PUBLISHED BY THE UNIVERSITY,
LAWRENCE, KAN.

Entered at the post-oflBoe in Lawrenco as second-class matter.

THE KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 7] OCTOBER, 1909. [^^o'^xv^Nr?

KOEBERLINIA SPINOSA Zucc. :

AN ECOLOGICAL STUDY OF THE ANATOMY OF THE STEM

AND SOME OTHER PARTS.

BY MIRIAM SHELDON.
Plates XVII to XXV.

INTRODUCTION.

SOME plant forms are peculiar and striking. Those of the
desert are particularly individual, many being even gro-
tesque in their chance variations. The choice of one of these
plants for study was suggested by Professor Stevens, of the
Botany Department of the University of Kansas. Accord-
ingly, after sectioning and examining about fifty or sixty
plantsw I selected and studied three more carefully. Of these
the subject of this sketch is perhaps the most interesting.

The material for this study w^as obtained in the summer of
1908. In July of that year collections of about sixty or seventy
diff'erent species were made by Mr. L. M. Peace, of the Uni-
versity of Kansas laboratories. These were taken from the
region around about Tucson, Ariz., representatives of some
species being taken from several habitats, as from mesa,
wash, irrigating ditch or hill top. As the short summer
season of rains had just begun some of the plants were ob-
tainable before the leaves had appeared. Specimens were, of
course, also taken after the leaves came out. The jars with
their preserving fluid (two per cent formalin) were taken into
the field, and the plants put into these as soon as collected. In
many instances the whole plant was secured, one jar containing
the root, stem, leaves and flower or fruit of the same plant. It
will be seen that this collection is particularly valuable because

(97)

98 KANSAS UNIVERSITY SCIENCE BULLETIN.

of the great care taken in obtaining it. Many thanks are due
Mr. Peace on this account, as well as for numerous verbal ex-
planations and descriptions of the desert, the vegetation and
the individual plants.

The literature dealing with the desert plants is, as is well
known, somewhat meager. So far very little more than a de-
scription or a mere mention of KoeherUyiia has been found.
Gray describes it in his Plantse Wrightiana^, and places it
with the Rutacese. Engelmann mentions it twice in his works :
once in his review of the Wislizenus plants and once as erro-
neously numbered by Emory among his Cactacese of the South-
west. Bentham and Hooker describe it under the family
Simarubacese. The Ailanthus, or "Tree of Heaven," is perhaps
the most familiar representative of this family with us north-
ern dwellers among mesophytic forms. John Charles Van Dyke
alludes to the "crucifixion thorn" in "The Desert." He says he
has met with it in several localities although it is reported as
"very scarce." Several travelers in Mexico and the Southwest
have made collections of this plant or have reported it. Prof.
Charles Bessey, of the University of Nebraska, in the "Burro
Thorn" (Holocantha emoryi) finds a plant of great interest, if
not one wholly as attractive morphologically, if one may judge
by the brief account he was able to give for lack of the "oft
promised material."

A description of the plant as a whole has not been found.
We do not know that anyone has ever had the hardihood to dig
up a root. It is said to make an impenetrable barrier, so that a
plant three feet high is of such dense growth that the main
stem is hardly to be seen. Among the earlier collectors of the
plant might be mentioned Chas. Wright, Dr. Josiah Gregg,
Gen. Wm. H. Emory and Dr. N. Wislizenus. The plant is
sometimes called "Junco," It seems as the northern border of
Mexico is approached the individual plants become more
stunted and less numerous than farther south. The plant is
thus described by Sargent:

"Leaves, not more than one inch long. Flowers, appearing
in May and June, about one-fourth inch in diameter. A bushy
tree, rarely twenty to twenty-five feet high, with a short, stout
trunk sometimes six to eight feet long and a foot in diameter ;
more often a low-branching shrub forming impenetrable
thickets, often of considerable extent. Wood very hard, heavy,
close-grained, dark brown somewhat streaked with orange;

SHELDON: KOEBERLINIA SPINOSA. 99

becomes almost black on exposure, with thin yellow or nearly
white sapwood of twelve to fifteen layers of annual growth.
Distribution : Dry gravelly mesas and foothills, valley of the
Lower Rio Grande, Texas, westward to southern through
northern Mexico."

The four or five plants seen by Mr. Peace were of various
sizes, the largest — the one from which the specimens were
taken — being about three feet wide and very compact in
growth. The branches were pale green in color, no leaves, the
flowers whitish to cream color. This was about three miles
southwest of Tucson.

As far as observed every branch is a thorn, some of them a
little blunt at the end and not tipped with the brownish needle-
like point. The thorns vary from 1.5 cm. to 7 cm. in length,
being from 1 mm. to 4 mm. in diameter at the base. The
largest stem obtainable for study was about 4.5 mm. in diam-
eter, and was probably five or six years old, possibly more. To
all appearances the bark is smooth, but microscopically it is
somewhat rough, the roughness due to the dome-shaped pro-
jections which occur over the stomatal chambers. The flowers
are borne along the sides of very small sharp thorns. They
have the appearance of being in a raceme — an umbel-like
raceme, some authors call it. The leaves, said to be scalelike,
are soon dropped.

INVESTIGATION.

The study of the desert plants was taken up in February,
1909, in connection with other graduate work at the University
of Kansas, and continued until the middle of August of that
year.

The methods employed in pursuance of this study were vari-
ous. The chlor-zinc-iodide treatment of sections fresh from the
formalin or paraffin, or softened from the dried state, proved on
the whole the most satisfactory.

All the figures, except a few of the flower and some diagram-
matic ones, were made by the aid of the camera lucida, the pro-
jection lantern or the photographic camera.

Stem. — The stem as a whole shows some interesting features
in cross section, of which the thick epidermis, deeply sunken
stomata, the deep palisade tissue, compact wood and small
water tubes are not the least striking.

The tissues might be taken in their order, beginning with
the innermost one. It has been observed that no generalization

100 KANSAS UNIVERSITY SCIENCE BULLETIN.

can be made in regard to the structure of these desert plants,
each one having its own pecuHar way of getting on.

The pith occupies a sort of triang;ular area. The cells are
comparatively large (see table), are heavily lignified, deeply
pitted, stored with starchy material, and possessed of — in al-
most every case — a small crystal having the form of the cal-
cium oxalate crystal. In the younger flower branch we find
cells with thick walls, also, but retaining their rounded shape,
pitted, and plentifully surrounded with air spaces. No food
material was observed stored in this region. In the older stem
elongated pit cells next the trachial tubes are densely filled
with material that stains blue with c. z. i. This has a fine
granular appearance (fig. 2, plate XX).

The medullary rays are conspicuously broad in cross section,
many being five cells wide. The cells are comparatively large.
They have the ordinary cylindrical shape with thick lignified
and pitted walls. (Figs. 3 and 5, plate XIX.) Intercellular
spaces are frequent. Most of the cells are full of non-nitrog-
enous material, while a few narrow and elongated ones appear
adapted for quick conduction. Those between the phloem
strands show the characteristic proteid reaction with c. z, i.
and have the usual unpitted walls. Crystals are often found in
these cells.

The xylem makes up a great part of the stem. The wood
fibers cannot be said to be very numerous. They have the
usual shape and are rather short. (See table.) The fiber-
tracheids are found quite plentifully and correspond closely to
the wood fiber. The bordered pits, however, are very numer-
ous, there being at least 200 to a tracheid. (Fig. 15, plate
XX.) The pits are small. The spiral tracheids have the single
and double spiral threads. These vessels are decidedly small.
The tracheids are of three kinds and vary in length and diam-
eter. (See table of measurements.) The tracheal tubes have
many small bordered pits and have thickenings in ridges in
the walls. The whole tube is divided at more or less regular
intervals, as from 0.6 to 0.27 mm., by thick rings. Upon in-
vestigation these may sometimes be seen to be in the plane of
union of neighboring tubes, the walls being so completely
fused as to obliterate all trace of joining. It seems that these
rin^s are really the borders of an aperture in the ends of tube
elements, occurring in the earliest laying down of the tissue.
They were observed in the last vessels laid down before col-

SHELDON: KOEBERLINIA SPINOSA. 101

lecting in the midst of the rainy season of 1908. (See figs. 5
and 6, plate XX.) These pitted vessels, often with ragged
inner thickenings (fig. 6), are found from the spiral vessels
next the pith to the latest growth. They are also found in the
young thorn which bears the flowers. The end of this division
of the tracheal tube is often very much elongated and gradu-
ally tapering (figs. 3, 4, 9). Sometimes the long tapering is
omitted and we have a peculiar cell ending as in figures 6a
and 7. These elements sometimes taper at both ends, but fre-
quently and abruptly in horizontal base. (Fig. 3.)

Besides the tube elements just described, there are the more
or less rectangular tracheids that join their neighbors above or
below by horizontal or oblique walls. (Fig. 13.)

No means to be had will give an exact account of the age of
the stem. This is true of most of the perennials of the desert,
we believe. Upon projecting upon the screen by means of the
lantern, a cross section of the older stem showed traces of five
or six seasons' growth, a slight increase in the size and number
of the vessels being noticeable in each successive yearly growth
as well as a somewhat greater compactness of growth in the
wood itself. Probably for the year 1907 a new set of vessels
has been formed. (Fig. 1, plate XVIII.) Outside of these is a
second set of larger and unquestionably new water vessels laid
down at the beginning of the rainy season of 1908. The width
of an annual ring varies somewhat, but on the whole has an
average of about 0.2 mm. The entire radial measurement of
the wood from pith to phloem is about 1.5 mm. This would
make the stem about seven and a half years old.

The Phloem region shows the usual storage of nitrogenous
materials in the sieve and companion cell portion and in the
parenchyma. The elements of the sieve vessels are rather long
(see table) and full of fine granular material. The sieve plates
are heavy, with very minute perforations on the sides. The
phloem parenchyma in my material occurs in a series of five
parts, each being about three cells wide. (Fig. 1, plate XXI.)

The stone cells and bast make a hard, rigid cylinder around
this more delicate portion, the stone cells occurring between the
bundles of bast and extending around them.

The sclerenchymatous or stone cells are of various shapes :
some are squarish, some oblong, and some so elongated as to be
confused with the bast as far as shape goes, but possessing
the deeply pitted characteristic of the stone cell. This zone

102 KANSAS UNIVERSITY SCIENCE BULLETIN.

occupied by the stone cells is broad. Sometimes the cells are
found in the region of the palisade cells, in scattered patches,
or in many cases filling up air cavities below the stomatal
chamber. (Fig. 1, plate XXIII.)

The bast appears to be of twofold character — the ordinary
long, tapering cells, united by a long splicing, then another
kind, shorter and tapering abruptly at one end, while the other
end, rather blunt, fits securely into the socket made by the two
cells below. (Figs. 8, 9, 10, plate XXI.) These fibers occur in
the regular bundles, but they also make a sort of inner bundle,
we think ; at least, in all the cases so far examined they appear
to occupy that position. They may make a sort of inner sup-
porting shaft to the whole group. When separated from the
long fibers a bundle of them looks like a barbed shaft.

The spoyigy parenchyma tissue takes up the inner collen-
chyma region. These are about five or six cells deep radially.
The walls of some of the parenchyma of the innermost portion
are heavily lignified. Neighboring cells have not so changed
materially, but the cellulose wall is thicker in many cases. The
walls in all the sponge are about twice as thick as those of the
palisade region. The spongy cells are spherical, cuboidal or
ovoid in shape, and about twice the diameter of the palisade
cells. The numerous and large intercellular spaces in this re-
gion are conspicuous. There are besides whole rows of open
spaces, seen in either cross- or longi-sections. (Fig. 9, plate
XXII.) This will be taken up in a later paragraph, however.

The palisade cells lie in characteristic position, two or three
deep radially. They are thin walled, cylindrical in shape, with
some, but rather small, intercellular spaces. The chloroplasts
are of goodly size, and show a deep blue in treating with c.
z. i. In this region also are to be found very large aerating
chambers, ovoid in shape, connecting directly with the stomatal
chamber above, and thence with the outer world. These are
not quite so numerous as the smaller cavities mentioned in the
sponge. (Figs. 3, 4, plate XXII.)

The epidermis is remarkable. The cuticle itself is very
heavy, it being 0.015 mm. in thickness in the old stem. The
whole epidermis as found in the older stem measures from
0.075 mm. to 0.105 mm. There is but one row of epidermis
cells. These are elongated radially, and in the younger parts
of the plant are of thin cellulose walls. In the older parts the
walls become very much thickened, so much so that in time only

SHELDON: KOEBERLINIA SPINOSA. 103

a conical space at the bottom represents the cell cavity. (Figs.
4, 5, plate XXII.) At intervals these picket-like cells give way
to large cavities, which they surround and protect by a slight
modification at the outer extremity, a curving upward and in-
ward, thus making a little dome of from eight to a dozen cell
tips. (Figs. 3, 1, plate XXIII.) These, not entirely closing the
cavity, leave an aperture allowing of communication between
the outer world and the stomatal apparatus within and thence
farther beyond to inner tissues. (Fig. 8, plate XXII.) The
details of this arrangement, although seemingly complex, are
actually simple. The stoma lies at the base of the epidermal
air chamber; slight projections of the walls of the guard cells
make a shallow stomatal chamber immediately above, and like
ones make one below. Beyond the lower lies the large ovoid
cavity referred to in the palisade tissue. (Figs. 8, 9, plate
XXII, fig. 11, plate XXIII; and for measurements, see table,
p. 7.) A curious little Phycomycete makes its home in the out-
side stomatal cavities. It occurs in a majority of the stem
stomata. (Fig. 7, plate XXII.) There are about 150 stomata
to a square millimeter. (Fig. 2, plate XXIII.) In the younger
stem a tissue for water storage occurs just outside the peri-
cycle. Sometimes this appears to be represented by a single
cell only. (Figs. 8, 9, 10, plate XXIII.) The region of spongy
parenchyma here stains deep blue with c. z. i., while the pali-
sade cells remain yellowish.

Flower. — The flow^er, cream color to yellow, shows the four-
parted arrangement — four very short triangular sepals, green-
ish in color; four long, blunt, prominently veined petals, en-
closing eight stamens, which show an inequality in length of
filament. The anthers are elongate-ovoid, and open on the
side. They are attached to the filament at about one-fourth
the distance from the base. The filaments are stout and taper-
ing upward. The superior ovary is globose in shape, tapering
above into a pillarlike style, which in turn tapers slightly at the
apex to form the stigma, which is a plain surface surrounding
a circular orifice.

Upon sectioning the ovary longitudinally the numerous
ovules are noticed in longi-section ; they appear hanging down
from the central placenta. In cross section two cells are seen,
and in some parts a suggestion of two side partitions. The
ovary wall is cutinized heavily without and within, insuring

104 KANSAS UNIVERSITY SCIENCE BULLETIN.

protection for the ovules against the intense aridity of the des-
ert atmosphere. The mesophyll of the ovary is well supplied
with chorophyll bodies and contains numerous mucilage cells,
thus assisting in the provision for intensest drought. A resin-
ous gum is found in the seed vessel surrounding the ovules and
the vascular bundles in the placenta. (Figs. 1, 2, 3, 4, plate
XXIV.) The resinous gum is not found elsewhere.

The contents of the cells of the stomata of the inner wall of
the seed vessel stain a deep blue with c. z. i., showing that the
guard cells here are living and filled with non-nitrogenous ma-
terial.

The mucilage cells finally break down, thus forming large
mucilage reservoirs. (Figs. 8, 7, 5, plate XXIV.)

CONCLUSION.

Koeberlinia is what Clements would call a stem xerophyte.
The low stature of the plant, its compact habit of growth and
the heavily cutinized surface everywhere, reduces evaporation
to a minimum. The dead-air spaces furnished by the outer
stomatal chambers also contribute toward this end. A supply
of moisture is insured for the more strongly growing parts by
the storage of water in special reservoirs and by the presence
of mucilage in the seed vessel. The resinous gum aids in con-
serving moisture. Greater strength is secured, perhaps,
through the peculiar splicing of the large tracheids. The stone
cells, with their firm adherence to the bast fibers, also give
strength. The air spaces in epidermis, palisade tissue, sponge,
medullary rays and pith insure a goodly supply of CO2 without
much evaporation. These points might be summed up thus :

Evaporation minimized by:

1. Dispensing with leaves.

2. Heavy cutinization on all exposed surfaces.

3. Dead-airspaces (outer stomatal chambers).

4. Low stature.

5. Compact growth.

6. Resinous gum surrounding tender growths.

Moisture supply provided for by :

1. Special reservoirs for water storage.

2. Mucilage in cells in seed vessel.

3. Tracheal elements.

4. Presence of resinous gum.

SHELDON: KOEBERLINIA SPINOSA. 1U5

Strength attained by :

1. Extreme lignification.

2. Long splicing of bast.

3. Long splicing of large tracheids.

4. Heavy zone of stone cells firmly adhering to bast.

Supply of air provided for by intercellular spaces in :
Epidermis.
Palisade.
Sponge.
Medullary ray.
Pith.

On the whole, this plant presents a wonderful array of
"aids" to getting on in the desert. No other plant examined
showed so thick an epidermis and such an aerating system. A
few were almost leafless, a few had a heavy epidermis, some
were strongly lignified, others showed resinous secretions. Al-
most all had larger water-conducting vessels. With all the
adaptations that this plant had it seemed strange that it was
so rare, but upon investigation we find that by abortion the
many seeds are reduced to one, two or three to a berry. We
have not yet found out whether the berry is eaten by animals
or not.

Tables of measurement, rainfall and evaporation are pre-
sented for reference ; also, a very superficial table giving some
facts in regard to a few other desert plants. I am indebted
to Mr. Peace for the tables on rainfall and evaporation. He
has also kindly furnished the two photographs as well as much
valuable information. Thanks are due to Doctor Trelease.
Doctor Land and Doctor Spalding, for kindness in answering
inquiries. I also wish to thank Doctor Billings for sugges-
tions, and Professor Stevens especially for ever-ready assist-
ance. It is a pleasure here to express gratitude to Doctor Mac-
Dougal for freely granting the facilities of the desert labora-
tory while the materials were being collected.

106

KANSAS UNIVERSITY SCIENCE BULLETIN.

TABLE OF MEASUREMENTS.

1.

2.

3.

4.

5.

6.

7.

8.

9.
10.
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.
21.
22.
23.
24.
25.
26.
27.
28.
29.
30.
31.
32.
33.
34.
35.
36.
37.
38.
39.

Structure. Length,

Mid pith cell 0.03 to

Marginal pith cell 045

Medullary ray cell 05

Wood fiber

Fiber tracheid

Tracheal tubes 06

Tracheids

Sieve cell

Phloem parenchyma

Stone cell

Bast ( long )

Bast ( short )

Spongy parenchyma

Palisade cells

Epidermis, older stem

Cutinized portion epidermis, old stem

Epidermis flower branch

Cutinized portion epidermis, flower stem

Epidermis-ovai-y, outer

Cutinized portion epidermis-ovary, outer

Epidermis-ovary, inner

Cutinizetl portion epidermis-ovary, inner

Epidermis of anther

Cutinized portion epidermis, antner

Epidermis of petal, outer

Cutinized portion epidermis-petal, outer

Epidermis petal, inner

Cutinized portion epidermis-petal, inner

Upper stoma tal space

Lower stomatal space

Outer stomatal aperture

Inner stomatal aperture

Air space in sponge

Mucilage cells

Mucilage reservoirs

Water storage cells

Resinous gum cells

Intercellular space, pith

Intercellular space, sponge

.03

.022

.05

.06

.015

.015

.03

.03
.03

0.045

075

075

24

25

27

114

25

12

12

15

03

03

12
06

045

06
092

Width, mm.

0.03 to 0.075
.01 .015

.012 .014

.01
.015

.015

.01
.075

.03

.01
.02

.025

!06

.03

.015

.007

.003

.015

.03

.025

.015

.015

.06

.013

.007

.02

.012

.105

.075

.06

.013

.065

.015

.045

.01

.02

.0019

.02

.0019

.015

.0019

.04

.03

.008

.015

.032

. 0325

.15

.045

.02

.015

.015

RAINFALL NEAR TUCSON.

Jan.

Feb.

Mar.

3

15

23

30

3

4

7

10

11

13

22

4

19

20

26

2'(

With snow.

0.62 inch.

Apr. 23

.05 "

May 3

trace.

" 23

"

July 5

0.29 inch.

7

.57 "

" 10

ti-ace.

•' 13

0.16 inch.

•' 15

.47* "

" 16

.24t "

•• 17

.15 "

" 19

•' 23

trace.

•• 25

0.38 inch.

•• 27

.03 "

" 28

.01 "

Aug. 2

0.11 inch.
.05 "
.03 "

trace.

0.04 inch.

trace.

0.80 inch.

1.00 ••

2.6

.01 ••

.61 "
1.81 •'
.02 "
.42 "
.42 '•

With sleet and snow.

TEMPERATURE AT THE DESERT LABORATORY.

Lowest. Highent.

July 6 77° F. 99° F.

" 7 77 103

" 8 74 99

" 9 78 90

" 10 78 95

" 11 74 101

" 13 71 100

" 15 67 97

•• 16 69 88

" 17 64 82

Lowest. Highest.

July 18 65° F. 74° F.

•• 19 68 87

" 20 75 95

'• 21 75 99

•' 22 75 94

" 23 62 92

"24 65 90

"25 72 91

"26 62 92

" 27 69 84

SHELDON: KOEBERLINIA SPINOSA.

107

RAINFALL OF TUCSON VICINITY.

January ..
February .
Mai-ch . . .
April. . . .

May

June

July

August . . .
September
October . .
November
December .

1904.

1905.

1906.

1907.

1908.

1.91

0.60

2.47

0.67

4.11

1.26

0.08

1.89

4.27

0.31

0.41

0.52

3.11

0.34

0.00

0.11

i.29?

0.03

0.02

0.45

0.08

0.04

0.24

0.00

0.00

0.00

2.07

1.47

2.07

4.67

5.73

3.00

0.47

1.16

3.66

*0.42

0.61

2.24

0.28

0.28

0.07

0.05

0.01

0.95

. * . <

0.05

4.52

0.66

0.74

1.07

0.90

4.40

0.00

Totals

* To August 2.

8.20

23.31

11.10

13.72

9.42

From July 5 to July 31 there were twelve days upon which rain fell. There was rain
during one day of each of the months of April, May and June. (These statements apply
only to the year 1908.)

WEEKLY AVERAGES OF EVAPORATION,
In cubic centimeters, from free .surfaces at a height of 25 centimeters above the ground.

June.

July.

Tuscon

St. Louis

Lincoln

Chicago

New York

Temperatures, July 5, 1908, to August 2,

August. September.

1908

338

228

212

278

112

157

140

137

96

90

113

144

93

96

109

95

48

63

35

i : Lowest,

62°

F.

; highest.

103°

F.

108

KANSAS UNIVERSITY SCIENCE BULLETIN.

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BIBLIOGRAPHY.

Bbntham et Hooker, 1862. Genera Plantarum, vol. I, pt. I, p. 315.

Bessey, Chas. E., 1904. The Chimney-shaped Stomata of Holocantha
emoriji. Bull. Torr. Bot. Club 31, 523-527.

Bower, 1908. The Origin of a Land Flora, p. 9, 480 to 494.

Cannon, W. A., 1905. Water Conducting Systems of some Desert Plants.
Bot. Gaz. 39, 397-408.

Clements, F. E., 1905. Research Methods in Ecology, Lincoln, Neb.

Coulter, J. M., and Chamberlain, J. C, 1903. Morphology of the Angio-
sperms, pp. 24-26.

Coulter, J. M., 1891. Botany of Western Texas. Contr. from U. S. Nat.
Herb. 11, 55, Wash.

Coville, F. v., and MacDougal, D. T., 1903. The Desert Botanical Labo-
ratory of the Carnegie Institution of Washington. Carnegie Inst, of
Wash. Publication No. 6.

Coville, F. V., 1892. Descriptions of the New Plants from Southwestern
California, Nevada, Utah and Arizona. Proc. Biolog. Soc. Wash.
7, 65.

Coville, F. V., 1893. Botany of the Death Valley Expedition. Contr.
from U. S. Nat. Herb., vol. IV, Wash.

CoWLES, H. C, 1899. The Ecological Relation of Sand Dunes of Lake
Michigan. Bot. Gaz. 27, 195.

Engelmann, George, 1847. Sketch of Doctor Wislezenus's Expedition,
also Cactaceae of Emory's Reconnoissance.

Engelmann, George, 1848. Botanical Works of George Engelmann. Edr
ited by Wm. Trelease and Asa Gray, p. 57; also, 113.

Engler und Prantl, 1895-'97. Die Natiirlichen Pflanzenfamilien, 6, III,
319-321; 4, III, 95, 230.

Gray, Asa, 1850. Plantae Wrightianae, Texano, Neo-Mexicanae. Smith-
sonian Contributions to Knowledge, Wash., vol. Ill, pt. I, p. 30.

Haberlandt, Dr. G., 1904. Physiologische Pflanzenanatomie,

Heller, A. A., 1900. Catalogue of North American Plants, North of
Mexico, p. 138.

Livingston, B. E., 1905. Relation of Soils to Vegetation. Bot. Gaz., 39,
22-41.

Livingston, B. E., 1906. The Relation of Desert Plants to Soil Moisture
and to Evaporation. Carnegie Inst, of Wash. Publication No. 50.

Lloyd, F. E., 1904. Pop. Sci. Monthly — A Botanical Laboratory in the
Desert, 66, 323-342.

Lloyd, F. E., 1906. Palo Verde: The Evergreen Tree of the Desert.
Plant World, July, pp. 165-171.

110 KANSAS UNIVERSITY SCIENCE BULLETIN.

Lloyd, F. E., 1908. Physiology of Stomata. Carnegie Inst, of Wash.
Publication No. 82.

Lloyd, F. E., 1908. Some Features of the Anatomy of Guayule {Par-
theninm argevtatnm Gray). Plant World, August, pp. 172-179.

MacDougal, D. T., 1904. Delta and Desert Vegetation, Bot. Gaz. 38,
44-63.

MacDougal, D. T., 1908. The Vegetation of the Tucson Region, Uni-
versity of Arizona Monthly, May, pp. 1-18.

MacDougal, D. T., 1908. The course of the Vegetative Season in South-
ern Arizona, Plant World, vol. II, pp. 216, 237, 261.

Ramaley, Francis, 1909. Wild Flowers and Trees of Colorado. Plants
and their Distribution, pp. 11-20.

Sargent, C. S., 1905. Manual of the Trees of North America, p. 682.

SCHIMPER, A. F. W., 1903-'04. Plant Geography upon a Physiological
Basis. Translation, Oxford.

Spalding, V. M., 1904. Biological Relation of Certain Desert Plants.
Bot. Gaz. 38, 122-138.

Stevens, W. C, 1907. Plant Anatomy.

Van Dyke, J. C, 1904. The Desert, p. 140.

Vassey, Geo., and Rose, J. N., 1890. List of Plants Collected by Dr.
Edward Palmer in Lower California, in 1889. Contr. from U. S.
Nat. Herb. I, 33, Washington,

THE

KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 8- October, 1909.

(Whole Series, Vol. XV. No. 8.)

CONTENTS:

A Case op Absolute Tone Memory, Archibald Hogg.

PUBLISHED BY THE UNIVERSITY,
LAWRENCE, KAN.

Entered at *he post-offlce In Lawrence as second-class matter.

THE KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 8] OCTOBER, 1909. [^oT xv^no'I

A CASE OF ABSOLUTE TONE MEMORY.

BY ARCHIBALD HOGG.

r?XTRAORDINARY forms of consciousness are necessarily
^ few, otherwise they would not be extraordinary. They
are usually worthy of study and report directly by reason of in-
herent interest and indirectly through the light they throw on
mental process in general. Among the exceptional forms of
memory activity we find in hearing what is called by musicians
"recollection of absolute pitch," by psychologists "absolute tone
memory." A striking instance of the possession of this faculty
led the writer, assisted by a friend' who is a skillful amateur
violinist, to make the two tests herewith reported.

The subject^ is a boy thirteen years of age, a quiet, gentle
lad, imaginative, introspective, fond of his music and of long
walks, and is perhaps less vigorous than the average child of
the same age.

He gave early evidence that he possessed unusual musical
gifts. One night when he was about four years old, and while
lying in bed, he listened to the playing of a wedding march at a
marriage ceremony in a neighbor's house. The next morning
he climbed on the piano stool at home and reproduced the
march — but in a simpler way. At the age of six his musical
education began with lessons on the piano, and it has continued
ever since except when interrupted by the vacations and ill-
nesses incident to childhood. His possession of absolute tone

1. Mr. A. K. Hubbard, assistant professor of civil engineering; in the University at the
time.

2. The boy is the son of a professor in our Univei'sity. Both jjarents are musical. One
older brother plays the cello, another plays violin. Two younger children cannot be char-
acterized with much certainty yet with respect to musical ability.

(118)

114 KANSAS UNIVERSITY SCIENCE BULLETIN.

memory was so strikingly emphasized as to attract attention
one day about three years ago, when he remarked to an older
brother who had just tuned a cello that the pitch of the instru-
ment differed from that of a few days before — an observation
that was verified by appeal to the piano. Since then the boy
has been tested, usually for amusement or out of curiosity,
some five or six times, but without any records being kept.
Composition comes easy to him and he has composed not a few
melodious pieces during the last four or five years.

The notes given in the tests and the observer's judgment are
indicated in accordance with a quite simple plan. The notes
from and including middle C to its octave above are left un-
marked. A tone in the next higher octave is indicated by a
single accent mark above the letter, in the second higher by
two accents placed above, and so on ; similarly with the lower
octaves, except that the accent marks are placed below the let-
ter in such instances. The usual symbols for sharp and flat are
used after the prescribed fashion.

The note or notes struck are placed first in the record and
are followed immediately by the observer's judgment. In both
tests the observer himself was seated with back turned at a
distance of about eight feet from the instrument and was look-
ing away from it. In other and unrecorded tests at home he
has been sent at different times to various places about the
house within hearing distance of the piano. The results of the
unrecorded tests have been practically the same, as conversa-
tion with people present on such occasions has revealed. The
uniformity of testimony regarding these unrecorded tests is the
reason why more tests were not made by the present experi-
menters.

The first test was made on the observer's piano in April of
this year. The results are given in table I.

Several times the observer was asked not to name the notes
but to come to the piano and point them out without striking
the keys. It was doubtless an unnecessary precaution.

A few miscellaneous tests were next made. The observer

was asked to reproduce, either by whistling or singing, certain

notes that were named, and thereupon sang or whistled in

correct pitch, as shown by the piano, the following notes,

f*', g**, c^/, b^, f. h'" a", d.

Following that a simple tune was played on the violin and
the key asked for. It was promptly and correctly given. The

HOGG: ABSOLUTE TONE MEMORY.

115

Table I.

Stimulus.

Judement.

Stimulus.

Judgment.

c

c

D/

D,

E

E

FBi

FBi

K'"

A'"

i B/G

B/G

D"

D"

GG#

A6A

B///

B///

E///G///

G'

G'

B /// F ///

B '" F '"

F.,,

F,,,

G// B// D/ G/

G // B // D / G /

B'"

B'"

EA// G// BA / D/

EA// G// B6/ D/

D#///

D#///

CF#

CF#

G'"

Qin

E,,,G'"

E,,,G'"

Ah'

Kb'

D,C;t

D,C#

experimenters having an idea as to the method of recognition,
parts of three or four tunes were played, care being taken not
to stop on the key note. The recognition was again prompt
and accurate. Being questioned, the observer said his judg-
ment in the first instance was determined by following the
general rule that a simple melody ends with the key note. In
the subsequent selections the judgment was determined by the
sequence of the notes.

Then a high note on the violin between b^" and b" was
struck. When asked the pitch the observer mentioned b", and
upon being told it was not exactly that he named bj^".

Next, several double stops were given on the violin, one note
being a shade off the key in each case. The note that was in
error and the direction in which it should be changed were cor-
rectly given. The observer was now asked to direct the opera-
tion of tuning the A string of the violin until it corresponded
with his own piano. The result, as shown by the piano, proved
that the judgment was slightly in error in the flat direction.
The same test on the D string gave a correct judgment. Prac-
tice with the A string doubtless helped here.

Lastly, the opening notes of a piece were sung. They were
named correctly by the observer, as appeal to the piano demon-
strated. He is ignorant of fingering of the violin.

The second test was made one afternoon about a month later
with the piano used twice a week by the observer when he

116

KANSAS UNIVERSITY SCIENCE BULLETIN.

takes his music lesson. The instrument is tuned to interna-
tional pitch and is lower than his own. He said it sounded
flat. The results are in table II.

TABLE II.

Stimulus.

Judgment.

Stimulus.

Judgment.

c#

c

QUI

G'"

G///

G///

Kb'

G'

A'"

A'"

D.

D,

D"

D"

Kb'

G'

Bb/^j

Bb ,11

Gff'"

G'"

G,

G. ;i

1

FB6

EA

Bb

Bh 1

B6,G

Aj±,F#

W

D'"

CFtt

BF

F^'"

p/"

G// Bi, D/ G/

G// B/^ D; G/

F.,.

F.v

G#
G

' Bit" D#'" Gs'"
"Bb" !>"¥'"

G"B"D"'

g///

B'" '

Fr A"Cr'

D*t//,

D.,

EiGB^D

DF#AC

With the first of the errors it occurred to the experimenters
that the judgments were being determined exclusively by
simple comparison with the image-standards from the home
piano — an hypothesis which would have involved uniformity
in the errors. Notes a half tone lower in pitch than those
really given should have been named.

Later it occurred to us that the observer himself might be
trying to correct the difference between the two pianos. That
this was usually the case became known from answers to ques-
tions asked after the test. The observer arrived at most of his
judgments by traveling the following mental highway: The
tone was first identified and named by means of the images
from his own piano — the instrument on which he daily played
and with which his familiarity was great. Having thus iden-
tified the tone as nearly as possible (exact correspondence in
any instance would be highly improbable), the simple addition
of a half tone brought about the judgment announced.

In a few cases direct recall of the tone-images from the teach-
er's piano rendered any additional mental labor unnecessary
to identification.

HOGG: ABSOLUTE TONE MEMORY. 117

In closing, a few statements are made by way of addition
and partial summary. For our observer the difficulty of iden-
tifying a single tone is scarcely harder in one region of the
piano scale than another. But there is a decided difference in
ease of analysis according as there are high or low complexes
of tones. In the third octave below the mental separation of
two notes was not attempted, the remark being made, ''It's
such a jumble, I can't remember," while in the third above the
identification was made with accuracy and without a noticeably
great amount of concentration. Images from the organs of
utterance seemed to play no part whatever in his recall of
tones. Sound images apparently furnished exclusively the ma-
terial used in all the judgments. Upon being questioned as
to his method of identification and of reproduction the observer
said he "just heard the tones in his head and told in that way."
Of a piece of music recently purchased for him he said he had
played it off "in that way" before touching a finger to the
piano. His introspective competency was greater than was
expected.

THE

KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 9— April, 1910.

(Whole Series, Vol. XV, No. 9.)

CONTENTS:

The Hackberry Psylla, Pachypsylla celtidis-mamm.e Riley.

A Study in Comparative Insect Morphology, . . H. B. Stough.

PUBLISHED BY THE UNIVERSITY
LAWRENCE, KAN.

Entered at the post-offce in Lawrence as second-class matter.

THE KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 9] APRIL, 1910. ['"^Txv'™:

THE HACKBERRY PSYLLA, PACHYPSYLLA
CELTIDIS-MAMM^ RILEY.

A STUDY IN COMPARATIVE MORPHOLOGY.

BY H. B. STOUGH.
Plates XXVI to XXXV, and three figures.
(Contribution from Entomological Laboratory, No. 158.)
Submitted in partial fulfillment of the requirements for the degree of Master of Arts.

OWING to their minute size and obscure life habits, the
members of the homopterous family Psyllidse have re-
ceived comparatively little attention in North America. In
the present paper an endeavor will be made to add to our
knowledge of this family, using as a type Pachypsylla celtidis-
mammse Riley, the same being a study in comparative mor-
phology of mouth parts, thorax, and genitalia, and develop-
ment of wing venation and wing pattern. The specimens upon
which I have based my studies emerged from their galls upon
the hackberry trees during October of 1908, at Lawrence, Kan.
I wish to thank Prof. S. J. Hunter, upon whose suggestion the
problem was taken up, and under whose guidance and counsel
the work was carried on, for his interest in my investigations
and for his assistance rendered.

In search for literature on the Psyllidse of North America,
and especially that dealing with Pachypsylla celtidis-mammai
and its genus, I was impressed with the fact that very little of
a definite and final treatment has ever been given to these in-
sects in this country. A few species have been described, a
few figured, and the life histories of a number worked out,
while keys for the species of only about one genus can be found.

In Europe, on the contrary, the contributions to the knowl-

(121)

122 KANSAS UNIVERSITY SCIENCE BULLETIN.

edge of this family have been numerous and have added greatly
to our information on the subject. The first of these contribu-
tions was made by Forster, in 1848. He was followed by Flor,
Thomson, and Hartig, but not much of an advance was made
upon Forster's work until there appeared in 1879 a treatise,
"Zur Systematik der Psylliden," by Low; this, together with
other contributions by the same author, has long remained
authoritative. The researches of Low in the taxonomy and bi-
ology of the Psyllidse, together with the anatomical work of
Witlaczil, have much simplified the study of this family in
Europe.

James Edwards, in "Hemiptera-Homoptera" (1896), treats
of four genera of the family, including twenty-eight species of
British Psyllidse, with tables for the separation of the genera
and species. He also illustrates these forms in color and gives
the typical wing venation of each genus.

Maskel (1889) lists and describes four species of Psyllidse
from New Zealand, giving keys for separating the subfamilies
represented in that locality, and figures the larval, pupal and
adult stages, and some of their typical anatomical characters.
This paper contains more figures of these characters than any
other I have been able to find, and was quite useful from that
standpoint. This writer also discusses the use and taxonomic
importance of the difi'erent characters of the family.

Froggatt (1903), in a paper on "Australian Psyllidse" de-
scribes eighteen species, mostly of the two genera Aphalaria
and Trioza. He figures the galls made by three species of the
latter genus.

The reasons for the dearth of material in this country may be
found in the fact that the family Psyllidse, like some other little-
known groups, being composed of small and inconspicuous in-
sects, has not been largely collected and studied, two or three
species excepted. One of these, the pear Psylla, Psylla pyricola,
is of great economic importance in some localities, and has long
been known and its life history carefully worked out. The life
histories of these insects are very difficult to determine def-
initely, and since some of them are gall-formers, the biological
side of the work becomes still more complicated. Only about
three specialists have ever devoted much time to their study in
this country, among whom was C. V. Riley. In an article on
"The Psyllidse of the United States" (1883), he gives a resume
of the present knowledge of the family and its principal char-

stough: the hackberry psylla. 123

acters, habits, life history, etc., listing thirteen species. Un-
doubtedly the most important work published so far is that by
the same writer entitled "Notes on North American Psyllidse"
(1884). This paper lists nineteen North American species,
distributed among the following genera: Livia, Aphalara,
Psylla, Pachy psylla, Tnoza, Calophyla, Rhinopsylla, and Cera-
topsylla, the descriptions of three genera and two species being
new. The most important part of this work, from the stand-
point of the present paper, is that dealing with the new genus
Pachypsijlla. This genus is peculiar to this continent, having
no equivalent in Europe, so Riley thinks, and to it belong the
hackberry psyllids. Only three species of these are placed in
his key. Riley also discusses "hackberry galls," in the Cana-
dian Entonfiologist (1883) .

An important contribution to the biology of the subject, es-
pecially of the gall-making phase, is an article on psyllid gall-
making in the fifth report of the United States Entomological
Commission, written by A. S. Packard (1887). This gives
keys to three species ; descriptions of pupa ; key to, and descrip-
tions of, the galls of the following species and varieties of the
genus Pachy jDsylla: venusta, gemma, asteriscus, umbilicus,
mamma, pubescens, globulus, cucurbita, and cucurbita var. To
this is added a detailed description of each gall and a con-
densed general life history of a hackberry psyllid. No mention
is made of the stimuli causing the gall-formation. Ashmead
(1881) describes the life histories and gall-making habits of
these insects, and lists nine species, adding three new ones.

Two papers by Charles W. Mally, entitled, respectively,
"Hackberry Psyllidse Found at Ames, Iowa" (1893), and
"Psyllidse Found at Ames" (1894), give a series of observa-
tions on the winter habits and spring egg-laying of Pachypsylla
celtidis-mammse, with descriptions of the eggs, larvae, pupse,
and galls, and record thirty-one species for North America.
He describes four new species and figures the early stages and
wing venations.

Where mention is made of the galls in the papers that I have
mentioned, only the gross structure of the gall is treated. Cook,
however, in "Galls and Insects Producing Them" (1903-'04),
has made a contribution toward the histology of the psyllid
gall ; he also discusses mouth parts and ovipositors in their re-
lation to gall-formation, but in common with other investiga-

124 KANSAS UNIVERSITY SCIENCE BULLETIN.

tors was not able to add much to our knowledge of the gall-
producing stimuli.

On the mouth parts of Hemiptera, in addition to works of a
general nature, I would cite two of importance in throwing
light upon the homopterous mouth as seen in the Psyllidse.
These are by Charles L. Marlatt (1898) and Walter J. Meek
(1903), and deal with the mouth parts of the cicada. Besides
these, such works as those of Sanborn on the Aphididse
(1905), Berlese on Coccidse (1896), and others, were of use
in a comparative study. The best work upon mouth parts of
Hemiptera is recorded in the German of Leon (1887), Geise
(1883), Wedde (1885), and Heymonds (1899). Dohrn (1865)
contributed to the internal anatomy of the Hemiptera, and
Witlaczil (1885) worked out the brain, eyes and digestive
system.

GENERAL CHARACTERS OF THE FAMILY PSYLLID^.

The Psyllidse were formerly classified with the Aphididse,
to which they are closely related, and are still sometimes
known as "the jumping plant lice." These insects are from
one-eighth to one-fifth of an inch long and in general appear-
ance may be likened to a minute cicada. Their principal char-
acters follow: Hind legs developed for jumping; antennae
nine- or ten-segmented (in life in constant vibration), last seg-
ment bearing two short spines ; tarsi two-segmented ; mouth
parts suctorial ; wing venation comparatively simple. Some
of the species are of economic interest, being injurious through
their feeding habits. A number form galls, notably upon
hackberry. A few species secrete "honey dew."

According to Mally (1894) there are thirty-nine species and
varieties in North America. Only nine species are to be found
in the collection of the University of Kansas, four of these hav-
ing been taken in Kansas. So far, not many fossil insects of
this family are known. Scudder lists in "Tertiary Insects of
America" (1890) only two species, Necropsylla Hgida and
Catopsylla pHma, the latter being of interest, as its wing
venation resembles more closely that of Riley's genus Pachy-
psylla than of any other.

The species taken for study belongs to this last genus,
Pachypsylla. The namer, C. V. Riley, says of it : "There is no
genus among those characterized by Dr. Fr. Low with which

stough: the hackberry psylla. 125

Pachypsylla can properly be compared. In the convexity of
the body it greatly surpasses Psylla, from which it is at once
distinguished by the vertical and rugosepunctate head, the
quadrate vertex, the short frontal cones, the less filiform and
stout antennae." This species was first listed for Kansas by
Tucker in 1907.

The coloration of Pachypsylla has been well described by
Riley. The general color is dark brown, with the abdomen
showing pinkish shades along the edges of each segment. The
wing pattern is dark brown, and the ocelli are red.

This insect is quite short bodied ; the sexes are easily distin-
guished, as in all representatives of the family, the end of the
abdomen being pointed in the female but blunt and appendage-
bearing in the male. The total length of body averages, for the
male, 3.2 mm., and for the female, 3.6 mm.

HEAD.

The head is carried in such a position relative to the re-
mainder of the body that the epicranium is in an almost ver-
tical position (3J^-4)* the frontal cones pointing downward,
with the outer face of the frons in a horizontal position, its
distal end projecting caudad. Viewed from the front, the head
corresponds roughly, in general shape, to a right triangle,
truncate at the vertex, the posterior edge of the epicranium
lying on the base of this triangle, and a point somewhat be-
yond the frontal cones representing the vertex (28-4). The
whole head is rather flat, thin, rectangular, and boxlike, the
open side of the box being the caudal side (28-1), against and
into which opening fit the anterior part of the prostemum, the
procoxse and the mouth parts.

The following parts and regions of the head, excluding the
frons and clypeus, which will be described under "mouth
parts," are to be found : epicranium, gense, frontal cones, an-
tennal sockets, compound eyes, and ocelli.

The epicranium forms the front wall of the head and is ver-
tical in position. Its posterior margin is slightly concave, and
curves caudad and ventrad to the opposite side of the head,
forming a roll-like edge. This roll-like portion is the dorsal
part of the head when the latter is in normal position. This
region of the head (28-4, ep) is rectangular, about 0.45 mm.

* Read plate XXXIV, fig. 4.

126 KANSAS UNIVERSITY SCIENCE BULLETIN.

wide and 0.62 mm. long. Its posterior edge is concave, as are
also the lateral edges which border on the compound eyes,
while the anterior margin is quite convex and has a somewhat
more than semicircular excision at its median point, in which
region is located the frontal ocellus, seen in figures 4 and 5, fo,
and in 2 in longitudinal section, pi. 28. The epicranium is
seen to be flat in surface when the head is viewed in profile.
The other two ocelli (25-4, o) are placed at the two posterior
angles of the epicranium, near its margin and slightly posterior
to the compound eyes. They are about 0.07 mm. in diameter.
The epicranium is divided along its median line by a very
plainly cut suture into two equal sclerites. This suture is deep,
and in making dissections of the head these two sclerites were
often separated readily along this line. The anterior, convex
suture between the epicranium and the rest of the head was
not so apparent in most specimens that I examined, though
usually those boiled thoroughly in potassium hydrate and then
carefully cleared in xylol showed it more plainly. The surface
of the epicranium is rugose-punctate, slightly pubescent.

The frontal cones, erroneously called "cones of the clypeus"
by Slingerland and "face lobes" by Scott,' are found, one on
each side of the median line, below the epicranium {28A, fc).
As seen from the front (28-5) they diverge from the vertex
just beyond the front ocellus. They are separated from the
cheeks or gense by a curved suture, which appears somewhat
ovate when the cones are viewed from below. This suture
could be made out much more easily on the cephalic than caudal
surface of the head.

The gense (28-1, 4:,5,g) extend from the cephalic edge of the
epicranium down on each side under the compound eyes and
fuse with the caudal, folded portion of the epicranium. Be-
tween the cones and the compound eyes is the antennal socket
(fig. 5, as), which is quadrate in shape, with angles much
rounded, and with a blunt projection, p, breaking into it on its
caudal edge, which is concerned with the articulation of the
basal segment of the antenna. The aperture filled by the
prosternum, procoxse, frons and mouth parts is similar in
shape to that of the epicranium but narrower. On the edge
of the gena, midway between the vertex and the compound
eye on each side, is situated a sharp projection (figs. 1 and

1, Insect Life, vol, V, pp. 229-230,

STOUGH : THE HACKBERRY PSYLLA. 127

5, ij) , forming the attachment for a membrane which runs
caudad between the front coxse, and which will be mentioned
more fully later. The outline of the compound eye is somewhat
more angular than rounded, viewed from either the front or
the rear.

In my study of the sclerites of the head I have followed Com-
stock and Kochi, and my conclusions for Pacliijpsylla agree
largely with the facts recorded by them.

ANTENNA.

The antennae are filiform, rather stout (a characteristic of
the genus Pachypsylla) , and average 0.83 mm. long. They are
10-segmented (28-3). The relative lengths of the ten an-
tennal segments on the basis of the length of the first segment
may be represented thus: 1, 1, 2, 1.2, 1.1, 1.2, 0.8, 1, 0.7, 0.4;
the second segment is wider distally than proximally and is
distally notched; the fourth, fifth, sixth, seventh and eighth
segments are constricted proximally ; the tenth segment is the
smallest segment and bears at its distal end two stout, blunt,
diverging spines (fig. 12), which are about as long as the seg-
ment and appear to be hollow. The last few segments are
compressed dorsally.

The cuticula of the first two segments is very deeply rugose,
forming scales, which on the second segment become longer
at the distal end (^5-3). The surface of the remaining seg-
ments is covered with deep, transverse corrugations (28-6),
which give to each segment a sharply serrate lateral edge, the
teeth pointing distad. A fringe of sensory hairs is found on the
distal ends of the segments, and a few scattered along their
surfaces, the tenth segment being without conspicuous hairs.
Figure 9, plate XXXIV, shows the last four segments, with
their corrugations and the terminal spines, while figure 6,
plate XXVIII, is of the articulation of the third and fourth
segment, showing the sensory hairs and the cuticular struc-
ture.

MOUTH PARTS.

The suctorial mouth as found in the homoptera, and as de-
scribed by Meek (1903) and others, consists of labrum, epi-
pharynx, labium, hypopharynx, and setse. These parts are all
represented in Pachyvsylla. Closely associated with them are
the frons, clypeus, mandibular and maxillary sclerites, and
tentorium, corresponding in position to those in the cicada.

128 KANSAS UNIVERSITY SCIENCE BULLETIN.

These parts are so closely associated with the mouth parts
proper (labium excepted), forming as they do a distinct re-
gion of the head, which is torn loose in dissection as a single
complex of sclerites (32-1), that they will be described in
this connection.

On account of the vertical position of the epicranium, the
mouth parts assume an orientation by which the frons, clypeus,
and labrum are almost horizontal, their distal ends projecting
caudad (28-6). The basal portion of the labium has a similar
orientation, but distad of its bend (29-2) it observes a general
ventral direction, though possessing some latitude of move-
ment cephalad and caudad.

Frons. — The frons is the largest and heaviest sclerite in this
region. Viewed from in front (looking dorsad) it is seen to
be obovate in outline (^.9-3). Viewed from the side (fig. 1)
its ventral outline is strongly rounded basally, with a dent at
about the beginning of the distal third. Its basal edge is
thickened (figs. 5 and 1) and continued laterally into proc-
esses (figs. 5, 3 and 1) which join the lateral ends of the
rod-shaped tentorium, t, and the chitinized rods, cmx. At the
place where the rods xijz (fig. 5) above mentioned leave the
frons at x, same figure, there springs a ligamentary process
(28-h, Ip) , which runs cephalad and dorsad of frons, forming
a small plate above the latter, and this plate in turn is con-
nected to the vertex by two small ligaments. These are for the
attachment of the frons to the head, still leaving some oppor-
tunity for motion. The frons is rugose-punctate of surface.

Clypeus. — The clypeus (29-1, 3, c) is about one-third as
long as the frons. It is more or less rectangular as viewed
from the front, broader at proximal end, and its lateral edge is
bent dorsad, thus partly enveloping the stylets or setse. The
slightly curved suture between this sclerite and the frons was
clearly marked in most specimens examined, though in a few it
was not readily discerned.

Mandibular and Maxillary Sclerites. — Between the lateral
edge of the frons and the rod cmx (29-1), and bounded proxi-
mally by the arm of the frons which joins cmx, is a more or
less circular foramen through which the bases of the setse may
be seen (figs. 3 and 1, for). This foramen is bounded dis-
tally by two sclerites; the proximal is crescent-shaped and is
known as the mandibular sclerite (fig. 1, mds) ; the distal is

stough: the hackberry psylla. 129

triangular and extends forward about as far as the end of the
labrum and is called the maxillary sclerite (fig. 1, mxs) . The
end of this sclerite has a rather slender, curved portion at its
apex, called the "maxillary process." These two sclerites are
shown in figure 3 in ventral aspect, and in figure 1 in lateral
aspect, where the clypeus, labrum, setae, etc., have been bent
ventrad in order to show the parts free and not overlapping.
The edge of the maxillary sclerite and maxillary process are
also shown in figure 5 looking ventrad. According to Smith
(1892) these two sclerites represent the true mandibles and
maxillae, the setge not being considered as such.

Mouth Parts Proper. Labrum. — The labrum {29-S and 1,
lab) is only about half as long as the clypeus and differs in
shape from the labrum of the cicada in that instead of being a
narrow, pointed process, it is, as viewed from in front, rec-
tangular in general shape, slightly longer than broad, with
distal corners rounded. It is not so flat as in the cicada, but
its lateral edges are bent dorsad in the same manner as those of
the clypeus, so as to partly surround the setse and the pharyn-
geal canal. At the proximal end of the labrum is found a
slightly curved suture between it and the clypeus. This suture
was very hard to make out, and the two sclerites in some speci-
mens appeared almost as if fused. Viewed from one side, the
ventral outline of the labrum is seen to be slightly convex, with
the distal portion bent somewhat ventrad. Marlatt's nomen-
clature makes the frons the clypeus and the clypeus and la-
brum a two-piece labrum.

Epipharynx. — Situated under the labrum (29-2, 3 and 1,
cp), and shorter than the same, is an indistinct organ, the
epipharynx. In the psyllid this organ is a delicate, fleshy,
tongue-like structure, more or less triangular in shape, with a
blunt tip, extending as a sort of outgrowth from, and exten-
sion of, the labrum. It appears as if it were a prolongation of
the labrum, and it was difficult in most cases to determine
where the one ended and the other began. The epipharynx is
pressed tightly upon the upper surfaces of the setse, surround-
ing them somewhat on the sides, and thus supplements the la-
brum in keeping them together to form a tube.

Labium. — The labium is attached below the hypopharynx, on
the lower side of the head (29-1, labi, and 3J^-6). This at-
tachment is by means of a delicate membrane made fast to the

130 KANSAS UNIVERSITY SCIENCE BULLETIN.

edge of the foramen next to the frons, which is easily torn
away in dissection. At first the labium is merely a mem-
branous band with only very slightly raised edges, which in
this region of the organ are somewhat chitinized. These edges
gradually become more raised and lose their chitinization until
finally a little before the bend in the labium these edges be-
come broad flaps which come together, and from this point to
the tip there is left only a slit (29-4:). The epipharynx lies
naturally against the floor of the labium, but it can be lifted
clear, together with the remainder of the parts, for some dis-
tance from this position. The first or basal segment of the
labium is membranous, not chitinized, since it is so well pro-
tected between the procoxse in a membranous fold. Each
procoxa is placed very close to and almost fits the foramen
laterad and dorsad of the frons {29-1, for). The first seg-
ment of the labium lies between the front coxae, embedded in a
membrane, and these coxae almost meet over (ventrad of) it
(29-2 and 31-1).

The attachment of the labium to the prothorax, in this case
to the region between prothorax and mesothorax, typical of
the Homoptera, is very strong as compared with that between
labium and head. In fact, in most cases when the head was
removed from the rest of the body the labium remained fas-
tened to the thorax but tore loose at its base from the head.
Meek mentions such a condition as present in the cicada, call-
ing the structure on the thorax to which the labium is attached
a "collar," describing it in the following words: "A heavy
membrane attaches the labium to a chitinized collar of the
thorax." In Pachypsijlla this collar is a chitinized process
(27-4) embedded in the membranes between the prothorax
and mesothorax (29-2, p). This process consists of two
heavily chitinized structures, flattened and concave at their
distal ends and large and swollen at their proximal ends, which
are connected by a curved, chitinous rod as a sort of yoke,
which is immediately under or dorsad of, and partly encircles,
the labium just cephalad of its bend (29-4, b) . The bases of
this process are embedded in the membrane dorsad and some-
what caudad of the cephalic margin of the mesothorax, from
whence the process projects ventrad and cephalad. The distal,
parallel ends of these are roughened on their edges and hol-
lowed out to form a groove, which, assisted by membranous

STOUGH: THE HACKBERRY PSYLLA. 131

attachment, holds that portion of the labium just beyond the
bend. Connected to the distal ends of these processes are mem-
branes which run cephalad between the bases of the pro-
thoracic coxae, surrounding the labium, to form a sort of fold
or groove for the same. This arrangement may be seen in
figure 2, plate XXIX, in diagram, m representing the mem-
brane surrounding the labium, p the process holding labium to
thorax ; the position of the labium and setse may also be seen.
Figure 1, plate XXXI, a cross section through the base of the
mouth parts and procoxse, corresponds roughly to Meek's
"Fig. 13" for Cicada, and shows the great similarity between
the two forms. The bases of the membranes surrounding the
labium are here illustrated.

The labium is slitted throughout its entire length. At its
base, where it is almost flat, it has only short, low, lateral
edges or flaps, and hence is broadly open, and the upper parts
of the mouth fit into this (29-2, 4). The labium, epipharynx
and setse all lie on the floor of the labium. Just at, or a little
beyond, the end of the epipharynx the lateral flaps become
larger and converge, their wavy margins fitting rather closely
together. From here on to the end of the labium these flaps do
not diverge again. Figure 6, plate XXXI, represents a cross
section of the first segment of the labium just beyond its bend.
Here the slit is seen to be held together by means of grooves
and ridges. There is also a depression on its caudal wall. The
lining of the labial canal is somewhat chitinized. On each side
of the canal are found two longitudinal muscles, m of the same
figure. The first labial segment is much roughened distally;
the second is quite short, chitinized, and bears sensory hairs
at its distal end ; the third segment is heavily chitinized, with
a pointed tip ending in two lateral, tonguelike processes. The
lateral outline of these is seen in figure 2, plate XXXI, and in
cephalic aspect in figure 3. The most notable feature of these
processes is the three large sense organs on each — large spines
on pegs. These same three sense organs are found on the
labium of Aleurodidse. This segment also bears several large
sensory hairs, two of which are much longer than the segment
itself.

Pharynx. — This is tube-shaped, chitinized, spreading some-
what at its cephalic end (29-5, p) . Its ventral surface is
supplied with eight pairs of muscles and tendons, which run

132 KANSAS UNIVERSITY SCIENCE BULLETIN.

to the inner surface of the frons. At its distal end the pharynx
fuses with the hypopharynx and hypopharyngeal lamellae. The
organ is heavily chitinized and thick-walled for some distance,
but soon becomes narrower and, losing its chitinization, pro-
ceeds caudad until about the middle of the back part of the
head, where it turns abruptly caudad and enters the thorax
to become the oesophagus.

Mandibles and Maxlllim. — In dissecting the mouth parts the
labium is generally torn loose from its connection below the
hypopharynx and other parts and left connected to the thorax,
as already mentioned. The setae, however, remain fixed firmly
to the upper parts of the mouth {32-1) and can be sepa-
rated and torn loose from these parts only with difficulty. They
are held together by the labrum and epipharynx above, the
maxillary sclerites and maxillary processes on the sides, and
by the hypopharynx below. From this point the bases of the
mandibles and maxillae diverge cephalad and laterad in a man-
ner which has aptly been compared to the form of the letter
"Y." The orientation of these parts was best seen in either a
lateral view of the frons, clypeus and other parts removed
from the head, or by examining the parts in such a position
as to look down on the bases of the setae in the direction of
their distal ends. In such a position this divergence, which
takes place immediately cephalad of the hypopharynx and
other closely enveloping parts, can be plainly seen. The bases
of these setae (^^-8, 9) are hollow, funnel-shaped, heavily
chitinized, and attached to the other parts by tendons and
muscles. One pair of these setae, from their cephalic posi-
tion, were recognized as the mandibles, and when dissected
out were found to surround the other pair of stylets through-
out their entire length, their slightly concave inner surfaces
fitting closely against them. This latter, inner pair represents
the maxillae. On dissection they were found to be held firmly
between the mandibles on each side and were difficult to sep-
arate. The mandibles are thicker than the maxillae, but not so
wide. Their relative shapes can be seen in cross section in
31-^. As in the cicada, the setae shortly after leaving the
head begin to curve so that the mandibles are above and below
the maxillae instead of laterad of them. This curving was seen
in sections distad of the bend of the labium and had begun in
the section from which figure 5 was drawn. In this figure the

STOUGH: THE HACKBERRY PSYLLA. 133

lumina in each mandible may be seen. These may also be seen
running to the tip when the mandibles are viewed extended, as
they are generally filled with air, which renders them visible.
No indication of lumina could be found in the maxillae. They,
however, by their concave surfaces fitting together, form a
canal through which the plant juices are sucked. The tip of a
mandible is very slender, sharp, and sickle-shaped, the convex
side bearing blunt serrations, the teeth pointing proximad
{26-1). The tip of a maxilla is of similar shape but with a
sharp projection on the convex side some distance from the
apex (fig. 6). The apices of these setai are so transparent
that the details of their structure were difficult to discern
clearly.

The homologies of the hemipterous mouth have been much
discussed in the past, and the opinions held have been various.
1 have followed the nomenclature used by Meek for Cicada,
which seems to be in harmony with late investigations along
this line. In their place I have compared the parts of the mouth
as seen in Pachypsylla with the corresponding parts in Cicada.
I also found in my work that comparison with other homop-
terous forms was helpful, showing the close relationship of
the Psyllidse to the other families.

The labium of the Aphididse is quite similar in general struc-
ture to that of Pachypsylla except in relative length and num-
ber of segments and in the fact that it does not possess the
bend characteristic of the latter insect.

The mouth parts, exclusive of the labium, are almost iden-
tical with, though larger than, those of the Aleurodidse which
I have examined. The same sclerites are present, arranged
similarly, though somewhat differently shaped. The labrum is
much shorter and smaller proportionately than in this family,
and the setae seem to be relatively shorter. The end of the
labium has the same lateral processes, but these are relatively
smaller. The Aleurodid labium has a bend similar to that of
Pachypsylla.

The coccid mouth parts also correspond in many ways. Here
the labium is very short and the setae are retracted into a
special pouch when not in use. The parts exclusive of the
labium are similar in general structure to the same parts in
Pachypsylla. Berlese's nomenclature and figures in his work
on Italian Coccidse were very useful to me. According to this.

134 KANSAS UNIVERSITY SCIENCE BULLETIN.

the clypeus or ipostoma (epistoma) corresponds to the frons of
Pachijpsijlla, the two terms "ipostoma" and "clypeus" being
generally used interchangeably by him. Cmx of my figures is
the ipostoma; xyz is the 7od-shaped apophyse of the clypeus;
the thickened, basal edge of the frons, z, is the transverse bar of
the ipostoma; the opening, for (29-1), is the foramen of the
ipostoma. The structure which Meek calls the tentorium is
designated by Berlese the transverse branch of the ipostoma.

THORAX.

The thorax as described by Riley: "Pronotum moderately
short, of equal width, slightly emarginate behind, steeply as-
cending posteriorly ; lateral impressions sculptured and colored
as the head ; well marked. Dorsulum well developed, thrice as
long as the pronotum, and about twice as wide as long; poster-
ior lobe distinctly longer than the anterior ; hind margin sinuate
each side and truncate at middle; surface finely alutaceous;
color, light brownish yellow with a large brown apical spot
divided by a yellow median line. Mesonotum convex, wider
than head, sculptured as dorsulum, with four vittse (longi-
tudinal) of brown or greenish-brown color, the outer ones
usually wider than the inner ones, all bordered and divided
transversely by lines of brighter yellow."

Prothorax. — Viewed from above the thorax is seen to be
composed of a number of sclerites, as seen in 27-S, A and
B, which upon closer examination and dissection may be shown
to make up the three typical regions of the thorax. In the
prothorax there is only one sclerite composing the tergum —
the pronotum (27-3, A and 31, pn) . This is collarlike, steeply
ascending, as seen in the longitudinal section (fig. 3 B), some-
what wider at the middle than laterad, where an obscure
suture divides it from the pleural sclerites. This is shown in
plate XXVII, fig. 1, and plate XXXV, fig. 12. The proepi-
sternum and sternum are very closely fused and the suture is
scarcely evident. The proepimeron (27-1, em i) is a rather
wedge-shaped piece, rounded on its ventral end, lying back of
the ventral portion of the pronotum and the dorsal portion of
the proepisternum. Situated in the pleural membrane beneath
the proepimeron is the prothoracic spiracle, sp. As seen in
35-6, the prosternum is only slightly developed, its cephalic
edge on each side being produced entad into a slender portion.
The prothoracic coxae are large and occupy, together with the

stough: the hackberry psylla. 135

labium, which is imbedded in a membranous fold between the
former, most of the prosternal region, leaving the prosternum
to develop mostly on the sides. The "collar," already described
as a chitinized process holding the labium to the thorax, may
be of the nature of the apophyse as seen in the endoskeleton of
the mesothorax.

Mesothorax. — The mesothorax is next in complexity of
structure, naturally being larger and more complex than the
prothorax, as it bears, besides one pair of legs, the fore wings.
The tergum is composed of three sclerites (34-2, and 27-3, A,
B), Edwards thus describes the mesotergum of the Psyllidse:
"Mesonotum large, generally suborbicular ; a portion of its
area in front in the shape of a broadly truncate triangle (the
dorsulum) , separated from the remainder by a distinct suture,
and a small crescent-shaped piece projecting from the middle
of its hind margin (the scutellum) , also marked off by a
suture."

The dorsulum {27-S A and 1) is in shape an equilateral
triangle, truncate at its angles, with the edges of the lateral
truncations notched. The posterior margin is not straight, but
produced on each side into a small projection midway between
the median line and the edge of the sclerite. The anterior third
of the dorsulum is covered by the posterior edge of the prono-
tum (fig. 3 A).

The mesonotum (27-S A and 1, mm) is the largest sclerite
of the thorax, ovate, convex, and about twice as wide
as long, with anterior margin almost straight and posterior
margin deeply convex. The sclerite is laterally narrowed to a
blunt, notched point, which is best seen in lateral view (fig. 1).

The scutellum (figs. 3 A and B, sen, and 1) is, viewed from
above, ovate, twice as wide as long, lateral edges rounded, an-
terior margin slightly concave and posterior convex and hol-
lowed out at the median line. The structure of the pleurum
of the mesothorax is exceedingly complicated and the sutures
are all indefinite. The structure of the same is shown (partly
diagrammatic) in figure 1. The suture between the meso-
sternum and the sclerite just dorsad could generally be made
out. This latter sclerite I take to be the mesoepisternum, es 2.
The mesoepimeron, es 2, is a very irregular, much folded scle-
rite intimately connected with the inwardly extending endoskel-
eton (fig. 2, es 2 and eytd) . These pleural sclerites are so folded

2-Univ._Sci. Bull.. Vol. V. No. 9.

136 KANSAS UNIVERSITY SCIENCE BULLETIN.

and fused one upon the other that practically only the region ot
each could be indicated roughly in figure 1. The origin of the
anterior wing may be seen in figure 1, bp. Just cephalad of
its base are two fleshy processes — the cephalic, larger and some-
what renifoi*m in shape; the articulatory epidemes, artep.
Cross sections show the larger epideme to be almost spherical,
with a broad, short base or pettiole.

The mesosternum is rectangular, wider than long, anterior
margin somewhat convex, with corners rounded. The coxal
cavities or acetabula are large, ovate, and are situated on the
posterior margin (35-11).

When the mesothorax is dissected free from the other seg-
ments, and viewed from behind, the endoskeleton is plainly
seen. I shall not attempt to give the orientation and detailed
description of its different parts, which may be seen in 27-2
(partly diagrammatic). This internal skeleton is a complex
of plates and rods, braced in all directions, and fused with the
outer sclerites of the segment. Three different internal pro-
jections may be seen, dorsal, lateral, and ventral. The dorsal
piece, the phragma, phrg, is a symmetrical, transverse plate
projecting ventrad. A similar structure in the mesothorax of
a stag beetle is figured by Packard (Textbook of Entomology)
and called "Diaphragm for the attachment of the tergal mus-
cles of the metasternum." On each side there is a simple pro-
jection which seems to correspond to the apodeme, apod. The
lower projection or apophyse ("medifurca," according to Pack-
ard) is seen at apoph, and in 55-11, and is shaped like a
letter "Y" ; its dorsal edge is hollowed out, this hollow together
with a similar one on the lower edge of the phragma above
giving passage to the digestive system, etc. (;?7-33, c.)

The mesothorax is built for great strength, the sclerites be-
ing much fused and the internal structures all large and heavy.
Although this segment carries the anterior wings, which are
used mainly in flight, I have no evidence that this insect is a
very strong flyer ; on one occasion a specimen was captured on
the top of a high building, but probably it was carried to such
a height independent of its own flying powers, by a strong wind
blowing at the time.

Metathorax. — This segment is the strongest in structure of
the thoracic segments, and the endoskeleton and external scle-
rites are so strongly fused that the homologies of this region
present a great many perplexing problems. I shall proceed to

stough: the hackberry psylla. 137

describe the different parts as I have found them, together with
my interpretation as to their homologies. The structure of
the notum is not so difficult to determine. Three sclerites are
plainly made out (27-3 A). The anterior of these is crescent-
shaped, and in life is concealed by the scutellum of the meso-
tergum. The dorsal portion of the second sclerite is a small
ovate plate, while the third and posterior sclerite is rather
broad, its posterior margin somewhat indented and its lateral
margins concave.

The metasternum (27-5', 30-8; 31-1, sir) is best developed
anteriorally where it is connected with a very light, narrow,
median portion between the acetabula, which in turn is con-
nected with and almost seems to be a part of the metafurca, mf,
of the endoskeleton. Closely connected to the internal pieces
of the latter, and separated from the sternum by an obscure
suture, is a sclerite whose dorsal end fuses with the other parts.
This probably represents the reg-'on of the metaepisternum
(31-1, es 3). The portion (x) in figure 7 which runs ven-
trad and entad to join a part of the metafurca has notches on
its ventral edge at x, and with these notches articulates the end
of a complex of sclerites extending down the side and across
the ventral surface of the segment (27-5 and 31-8). Much
difficulty was found in homologizing these sclerites, authorities
differing as to the names of some of the parts. Upon tearing
loose this complex carrying the metathoracic leg from its at-
tachment at X and y (31-8) and examining it under a high
power of the microscope, its structure could be made out. It is
composed of five sclerites. Entad (somewhat caudad and
dorsad) is a more or less crescent-shaped sclerite (fig. 8, str).
This I believe to be a part of the sternum. The next sclerite,
mer, extad and cephalad, is of a similar shape but bears a blunt
projection or tubercle. Authorities differ as to what is the
homology of this part. Westwood says : "Beneath, the epimera
of the metathorax are singularly produced behind the place of
insertion of the hind legs (which are pushed forwards), and
terminated by two strong spurs." Edwards states : "Meso-
sternum produced behind into two large, sharp spines." Riley
states: "Metasternal processes cylindrical, hardly narrower
toward the tip, which is obtuse, not pointed." I find a similar
structure in all the Cicadidse which I have examined. In "Les
Cicadines D'Europe," translated from the German of Fieber
by Reiber (1875), in the discussion of the general structure of

138 KANSAS UNIVERSITY SCIENCE BULLETIN.

the Cicadidse, under legs, this sclerite is called the "meracan-
thus" ("A la base des hanches et specialement aux posterieures,
nait, visible chez les Fulgorides et remarquablement developpee
chez Cicada, une epine subulee, au lobe triangulaire, lanceolee,
cornee, qui est encore assez distincte aux hanches intermedi-
aires, Le Dr. Hagen [Die europ. Cicaden, Stett. ent. zeit. 1886]
appelle trochanter [Klappe] cet appendice corne qui doit etre
designe sons le nom vrai de Meracanthus, pi. 10, fig. 9, b, car
le trochanter n'a pas d'apendice").

Focusing upward with the high power, the next sclerite, also
crescent-shaped, with parallel edges, which is seen to be the
next extad, comes into view (fig. 8, em 3). It is this sclerite
which articulates at x and y (figs. 7, 8). This I believe to
represent the metaepimeron. Cephalad and laterad of these
sclerites is a platelike piece, the cephalic end of which bends
dorsad and entad, and forms a ring. This structure is seen in
figure 8, cox. On this ring, the trochanter articulates (also
upon the end of the episternum), and since this sclerite is the
only one which in any degree approaches the tj^ical cylindrical
coxal form, being in this case a sort of ring at its end, I take it
to be the coxa. Entad (dorsad) of this, and slightly cephalad,
is an accessory sclerite (55-8 and 9, scl) , a rod whose caudal
end is flattened out into a plate to which are attached muscles
that arise from the inner surface of the meracanthus, or epi-
meron (which of these two, I am as yet unable to state). The
cephalic end is connected to the cephalic edge of the trochanter
and serves to pull it towards the body. Its position is shown in
figure 8, plate XXXV.

Turning now to the internal structure of the metathorax,
which is best seen from the front, when this segment is severed
from the others the metafurca (50-8; 31-1] 27-5, mf) are
the most prominent features. The two large forks which pro-
ceed dorsad and caudad are connected with the lower surface
of the tergum. The whole endoskeleton is so closely fused with
the sclerites of the metathorax that it cannot always be sepa-
rated from them. This internal structure is large and heavy,
and by bracing all parts makes the segment exceedingly strong
and rigid.

The structure of the pleural sclerites in a species of Aleuro-
did which I examined was also found to be very complex, there
being a large number of sclerites homologous to those found in

stough: the hackberry psylla. 139

Pachypsylla, and a careful comparison of the two forms would
throw much light on the question. •

The metathorax is not so heavily braced as in Pachypsylla,
but the coxal region is still well developed. A study of the
psyllid pupa would probably show much of interest in this con-
nection, as these parts above mentioned are all represented
there but somewhat simplified. The folding and fusion of parts
in the adult, and the presence of a heavy, braced endoskeleton
and the complex articulation of the trochanter of the meta-
thoracic leg, are all specializations to render possible and in-
crease its jumping power.

My studies on the thorax of the Psyllidae agree with the
recent articles by Snodgrass (1909) , in which the old theory of
Audouin making each tergal segment to be composed of four
primitive sclerites is discredited; I have been able to find no
more than three sclerites composing the mesoterga and meta-
terga.

WINGS.

I. VENATION.

The wing venation of the Psyllidse seems never to have been
reduced to any uniform system such as that of Comstock and
Needham (1898-'99). I find the venation figured by Mally,
Edwards, Slingerland (1896) and others, but the nomencla-
ture is not uniform and is of a special nature. I have at-
tempted to reduce the venation as seen in the insect under dis-
cussion to the system of Comstock and Needham, above
mentioned. In this study my first conclusions as to the homolo-
gies of the veins were arrived at through the study of adult
wings alone, but later I have made use of nymphal wings and
wings of newly emerged adults.

On March 6 a large number of pupae of Pachypsylla celtidis-
gemma were procured from their galls on the stems (lateral
buds) of the hackberry, some seven or eight being found in
each large-sized gall. Their wing pads, after soaking for about
a day in dilute formalin, were removed, mounted in dilute
glycerine, and studied with special reference to their tracheae.
As soon as I had begun this study I found that my conclusions
arrived at earlier were borne out in almost every detail by the
number, position, character, etc., of the trachese of the
nymphal wings, these tracheae corresponding to the veins of
the adult wing.

140 KANSAS UNIVERSITY SCIENCE BULLETIN.

Venation of Nymphal Wing Pads and Wings of Newly Emerged Adults.

Fore Wing (26-3 and 32-S). — The costa, C, rises from a
large tracheal trunk from the thorax, proximad of the radius
and subcosta (fig. 3). It follows the anterior margin, but
drops down at the region of the stigma and fuses with the
subcosta.

The subcosta, Sc, and radius, R, are fused throughout the
basal third of the wing, and the free portion of the first-named
vein is comparatively short, as it soon fuses with the costa.

The radius is represented by a single trachea, and at its
base, where it is fused with the base of the subcosta, it rises
from the large tracheal trunk, distad of the origin of the
costa (fig. 3).

Media and cubitus, M, and Cu. The media rises from the
main trunk just cephalad of the base of the cubitus and distad
of the base of the subcosta (fig. 3). It is two-branched and
shows its double structure throughout the distal half of the
pad, the two branches being close together and parallel for
some distance. The three tracheae, the subcosta plus radius,
the media and the cubitus run parallel throughout the basal
third of the wing. In normal position, these three tracheae are
very close together, as in figure 8, but in the diagram (fig. 3)
they are represented more widely separated to show their con-
nections with the main tracheal trunk from the thorax. The
occurrence and relative positions of these three tracheae are of
the greatest importance in correctly determining the homolo-
gies of the adult veins, as the upper of these branched veins,
formerly called the "upper branch of the cubitus," arises from
a trachea distinct from the true cubital trachea, and therefore
cannot be a part of the cubitus, while the trachea representing
the subcosta plus the radius fused is distinct from those of
media and cubitus, though all four veins appear as a single
vem in the basal portion of the adult wing. Both media and
cubitus are two-branched, their distal ends bending so as to
run parallel with the costa for some distance before fusing
with it, forming at some places as many as three parallel or
more or less intertwining lines of tracheae (fig. 3).

The anal fold or "claval suture" (fig. 3, af) is present as a
distinct trachea which fuses at its distal end with the second
branch of the cubitus and the costa near the posterior margin.
Only one anal vein (fig. 3, A) could be made out, but as this

STOUGH : THE HACKBERRY PSYLLA. 141

trachea was torn more or less when the fore wing pad and the
hind wing pad, which are grown together at this point, were
separated, such evidence in itself is not conclusive. However,
from my studies I. have every reason to believe that normally
only one anal vein is represented. Likewise, the basal connec-
tions of the costa of the posterior margin and the trachea of
the anal fold to the main trunk from the thorax could not be
accurately determined.

For the further study of the development of the venation, a
number of Pachypsylla celtidis-mammse which had emerged
during the fall of 1908 were thrown into weak formalin im-
mediately after emergence, before the chitinized veins had
been laid down (26-4; 32-9, 10 and 11). In one specimen
(26-4 and 32-d) the wing was still delicate and transparent,
with its edges crumpled and folded, and the veins still prac-
tically in their tracheal condition, while in other specimens the
wing membranes had hardened, the pattern gained its pig-
ment, though still light, and the veins fully formed but with
the tracheae in some cases visible within them (-5^-10 and 11),
In the youngest specimen mentioned above the three parallel
trachese of the subcosta plus radius, media and cubitus were
strictly separate and corresponded in position to those of the
nymph of P. c.-gemma. The distal ends of the veins, becoming
lost in the folds of the margin of the wing, could not be traced.
The position and direction of these trachese here were more
closely like the veins found in the adult than were those of the
nymphal wing pads.

These studies aided me much in studying the development
of the trachese into veins, and the coalescence of two or more
of the former into one vein, as seen in the case of the sub-
costa plus radius, media and cubitus. In some specimens
these could be seen to be still single and parallel until they
reached the internal trunk (26-4 and 32-9). The radius and
subcosta fused (32-10) was on the cephalic margin of the com-
pound vein in process of formation, the cubitus (fig. 11) being
on the caudal margin.

The vein in the anal fold could still be plainly seen (26-4),
but its basal connection was not made out. Since there is no
thickening of the wing here, as soon as the air leaves this
trachea the only suggestion of a vein that is left is the trans-
parent "fold,"

142 KANSAS UNIVERSITY SCIENCE BULLETIN.

Since there is so little chitinization and thickening of the
membrane around the trachese of the hind wing, these can be
studied best in specimens such as the above, where air in the
tracheae renders them visible. This is especially true of the
costa, subcosta and vein in the anal fold. The subcosta cannot
be made out in the adult hind wing, and has not been figured
for it, but shows plainly when filled with air. By placing a
piece of black paper under the condenser of the microscope
the veins appear a glistening white, and under the low powers
are much more easily studied in this manner.

An interesting point I observed in the hind wing was the
presence — in one specimen — of what should probably be re-
garded as a "sport" vein (26-2, sv) . It is connected with the
anal vein and also fuses with the costa. Since I have found in
one hind wing of two specimens, both male and female, the
radius forked, as is the case with the normal cubitus, which
branch, though a "sport," probably points back to a former
two-branched condition of that vein as seen in the fore wing,
this vein above mentioned may show the same condition for
the anal wing, and represent the second anal vein. I find
under "Psyllidse" in the "Zoological Record" for 1867 the fol-
lowing: "From the occurrence of irregularities in the vena-
tion of the wings of this species, [Amsosiro/j/m ficus Linn.] he
[Frauenfeld] takes occasion to remark upon the frequency of
such irregularities among the Psyllidse ; the variation is almost
always confined to one wing."

Adult Venation.

Fore Wing (27-6). — The costa can be easily traced around
the entire margin of the wing and is very thick at the basal
portion of the anterior edge. All the rest of the veins except
the anal arise from a single main trunk, which is the main
support for the wing. This trunk is composed of the three
tracheae mentioned above, fused — the subcosta plus radius,
media and cubitus. The subcosta and radius are fused as far
as the stigma, s, where the subcosta runs up toward the costal
margin, forms the lower margin of the stigma, and meets the
costa somewhat proximad of the apex of the wing. The radius
is composed of a single vein, probably representing R 4 -f 5.
At about the middle of the cell, /, a vein leaves the main trunk
which divides into two branches, each of these again dividing
once. The two upper branches represent M 1 -|- 2, and M

STOUGH : THE HACKBERRY PSYLLA. 143

3 -f- 4. In one specimen of Psylla coryli I found that on one
wing the media was single instead of two-branched, but never
found such a case in the present species. The two lower
branches above mentioned are Cu 1 and Cu 2. Below this is
what is called the "claval suture," corresponding to the anal
fold (fig. 46, af). The region posterior to the anal fold or
claval suture is generally called the "clavus," from the corre-
sponding region in heteropterous wings, cl. One anal vein is
present; this, by a dorsal and cephalic folding over of the
distal half of the margin of the clavus, takes the place of the
costa at this point (that is, forms the posterior edge of the
wing here), the two veins appearing upon first observation
to have crossed. This condition was also observed in the
nymphal wing pads and in the wings of newly emerged indi-
viduals.

Hind Wing (27-7). — This is more simple in venation than
the fore wing. The costa soon becomes weak and indefinite.
The subcosta is not visible except in newly emerged individ-
uals where the tracheae are filled with air, and the radius is
seen as in the fore wing, as a single vein. The media is also
a single vein, though in two specimens I found it two-branched
in one wing after the fashion of that vein in the fore wing.
The cubitus has two short branches. The claval suture and
anal vein are practically the same as those in the fore wing,
only there is no folding over of the posterior margin of the
clavus.

The adult wings as above described are specialized by re-
duction, the coalescence of veins being outward.

According to Mally's nomenclature, the common base of the
media and cubitus which joins the subcosta plus radius is
known as the "Petiolus cubite," Pc; the media and cubitus be-
fore branching are known as the "first cubitus" and the "sec-
ond cubitus," and their branches the "first, second, third and
fourth furcal veins." The presence of a media is not recog-
nized by him.

Edwards calls the media and cubitus the "upper and lower
branches of the cubitus," and the "Petiolus cubite" of Mally
the "Stalk of the cubitus." He mentions no media.

Woodworth (1906) adheres to a nomenclature somewhat
different. According to this, the media and cubitus are called
the "independents" which "appear as a twice-forked branch

144 KANSAS UNIVERSITY SCIENCE BULLETIN.

from the primary," the fact that these veins are distinct,
arising from separate tracheae — as I have shown — not being
recognized.

The Psyllidae are the highest of the group Phytophthires of
the Homoptera. These all have simple wing venations, that of
the Coccidse being the simplest, Woodworth says of these :
"The venation of the Phj^ophthires has a nomenclature of its
own, not relating it to any other groups, the only investigator
who has attempted to homologize the veins being Redtenbacher.
This author admits to only two branches as belonging to vein
V (corresponding to my independents) [vein m] in the Psyl-
lidse, and sees none at all in the Aphididse."

II. PATTERN.

Fore Wing. — I quote Riley's original description of the wing
markings of this species : "Front wings narrower and, there-
fore, apparently longer than in vemusta, very little wider at
terminal third than at basal third; costal margin at base but
slightly subhyaline, more or less densely covered with small,
brownish, scalelike specks, as follows: an oblique and gradu-
ally narrowing band (extending from the tip of the ptero-
stigma to near tip of radius), which runs across the wing to
the tip of the first furcal, is usually freer from these dots than
the rest of the wing, while its edges are usually* more crowded
with the dots, so as to bring the pale band into greater relief ;
quite frequently there is an intensified patch of brown about
discoidal part of subcostal vein and at all vein furcations ; also,
the terminal space outside of the pale oblique band is uniformly
dark, but with three marginal pale spots (one on each side of
the cubital, second marginal and discoidal cells), always more
or less evenly speckled; exceptionally, both inside and outside
the oblique band, there is a border of variable extent, of a uni-
form dark brown color, not composed of small dots."

There is great sexual difference in color pattern and in in-
tensity of markings, as seen in figures. The dark border of the
light band mentioned in Riley's description is generally very
weak in the female, sometimes almost obsolete, except where
it crosses the veins, as in figure 6, plate XXXIII. In general,
the wing of the male is darker and more heavily marked than
that of the female. Also, the spring forms of both sexes are
said to be darker than those that emerge in the fall and hiber-
nate over winter.^

2. Insect Life, vol. V, p. 227.

stough: the hackberry psylla. 145

In the males there is generally a large dark spot between
Cu 1 and Cu 2, and on and below the radius proximad of
its fusion with the subcosta, this spot tending to spread
across cell I. These spots, proximad from the stigma, tend in
the female to be reduced to dusky portions only a little darker
than the surrounding parts, the former being almost indistin-
guishable and the latter confined to a border to the vein. The
three marginal spots mentioned by Riley are generally more
distinct in the male. There is a great deal of individual varia-
tion in the wing markings as to the number and arrangement
of the small spots composing the pattern, no two individuals
showing precisely the same combinations. Figures 1 and 5,
plate XXXIII, show how this pattern is produced, and inci-
dentally the difference in shape of the apices of the male and
female wings. Figure 4, plate XXXIII, is another view of the
pattern at the branching of the median veins, more highly
magnified than the two preceding.

The upper surface of the wing is entirely covered with disk-
like cuticular outgrowths about 0.005 mm. in diameter, best
seen in cross section through the wing {30-2, a and h) . These
are more or less circular in outline though often very irregular.
Some of these, h, are filled with dark brown pigment, the same
diffusing slightly into the surrounding cuticula. These pig-
mented disks are arranged in groups of 2, 3, 4, 5, 6, 7, 8, and
so on, to form polygons and other figures. These groups form
the "spots" which make up the pattern, and are either distinct
or banked up together to form a dark spot or band, in which
case the polygonal structure is not so apparent. The following
elements make up the pattern: (1) The individual pigmented
cuticular processes, which I shall designate as "disks"; (2)
polygonal and other combinations of these disks — "groups" ;
(3) arrangements of these groups making up the large and
small spots, bands, etc. — "markings."

There are a great many different forms of groups produced
by these disks, the principal types of which are seen on pago
146. Polygonal groups predominate, though chainlike, broken,
and irregular forms are also found. Groups of "two's" and
"three's" arranged lengthwise with the wing are found pre-
dominating in the basal cells, while more complex polygonal
figures are the rule in the distal regions of the wing. Groups
consisting of only one disk were not found.

Any difference between the composition of the groups or the

146 KANSAS UNIVERSITY SCIENCE BULLETIN.

frequency of certain group types in the male and the female
was not found. The polygonal gi'oups are oftener without the
central disk dark than with. When this was not pigmented,
the space was still clouded the same as the cuticula surround-
ing the pigmented disks.

I

tt

^

^. A

^ :• 'i

^ :: •'

•

Fig. 61. — Diagram showing principal forms of
groups found in the pattern.

The disks were found to be smaller proximad of the base of
the wing and near the veins and the anal fold, and consequently
the groups there were either of uniformly small disks or of part
normal and part small-sized disks. There were no groups on
the stigma ; here the disks became reduced to mere low, conical
projections. The anal fold was smooth and transparent, with
clear-cut edges.

Cross section (30-9, c) shows on the lower side of the wing,
!»pposite each disk, a small, blunt outgrowth, generally unpig-
mented.

Hind Wing, — Here the disk structure of the membrane is
barely visible under favorable conditions of illumination of the
microscope. Only those disks on the clavus are pigmented.

stough: the hackberry psylla. 147

The number of groups averages thirty to thirty-five. They
tend to be irregular and not polygonal. Groups of "four's"
are perhaps the most common, being generally found singly,
though sometimes two or three groups are associated together
to form a figure, generally a chain.

III. STRUCTURE.

Fore Wing. — Of the sixty wings that I examined, the fore
wings averaged 2.94 mm. long by 1.19 mm. at greatest width
in the male, and 3.27 mm. by 1.37 mm. in the female. Varia-
tions in outline of the wings and in the shape of the cells were
very slight. There was, however, variation noticed in the
length and breadth of the wing between rights and lefts in
both sexes.

In the membrane at the base of the fore wings are found two
small sclerites, called by Audouin "articulatory epidemes,"
already mentioned {27-1, artep). The cephalic of these is
seen in cross section to be more or less spherical in outline.

On each side of the wing veins is a row of rather stiff, curved
hairs in circular pits, seen in 33-1 and 5. On the anterior
portion of the costa there are three rows of hairs, the an-
terior of which is separated from the others by a central,
dorsally projecting ridge of the costa. From a point near the
proximal edge of the light band, already referred to under
"pattern," these hairs are found in a single row until the place
is reached where the anal vein and the posterior costa seem to
cross. From here on the costa bears no hairs, but in their
place a row of very minute, sharp teeth on its outer edge.
These teeth are also found on the edge of the anterior costa
below the smooth-edged cuticular margin of the wing ; they are
somewhat larger here, but become obsolete a little before the
apex of the wing is reached.

Upper and lower surfaces of the wing are essentially the
same, as regards number and position of hairs. The structure
of the membrane has already been treated. It is submem-
branaceous in texture.

Hind Wing. — The hind wing averages 2.5 mm. long by 0.97
mm. wide. The apical portion is wider than the basal and is
well rounded. The anterior margin is fairly straight, though
somewhat undulating in outline. No hairs are found on the
veins of the hind wing except two or three on the fused portion
of the subcosta, radius, media and cubitus, and on the basal

148 KANSAS UNIVERSITY SCIENCE BULLETIN.

portion of the anterior costa, where fifteen or more — the num-
ber being- variable — long, stiff hairs project cephalad, curving
upward {S3-S). The hind wings are very weak and delicate,
and these hairs are of use in coordinating the movements of
the hind wing with those of the fore wing, resembling some-
what the "hamulus" of the Hymenoptera.

The anterior margin of the costa is only slightly roughened
—covered with low, knob-like processes, distad of the hairs—
the basal two-thirds of its length, but from here on as far as
the beginning of the clavus these knobs become lengthened out
into spine-like scales pointing distad. The margin of the
clavus is comparatively smooth except where roughened at the
humeral angle.

The membranes of this wing are transparent and thin, and
were often separated from each other to form a sac, when
boiled too long in caustic potash. This is in harmony with the
studies of Weismann, Kiinkel d'Herculais, Dewitz, Van Rees
and Pratt, which established the fact that insect wings are of
hypodermal origin, being evaginations of the integument, and
the separation of these membranes through the action of
caustic potash shows plainly that the wing membrane is made
up of two laminae, an upper and a lower, which are fused to
one another between the veins. The veins, except the costa,
are almost without chitinization, and the subcosta could be
made out only when it was filled with air as in newly emerged
specimens.

The fore wing is quite flat, except the clavus, which is bent
somewhat ventrad, and that portion of the clavus between the
anal vein and the costa is bent quite sharply ventrad, so that
it is almost horizontal in position when the wings are folded
along the side of the body. When the insect is at rest and the
wings are in this position the anal angles of the fore wings fit
up closely against the edges of the scutellum of the meso-
thorax, and their edges are contiguous from the distal end of
the clavus to the tip of the wing. Through the gap between
the claval portions of the wings can be seen the posterior mar-
gins of the hind wings, which are contiguous throughout the
claval region, their anal angles fitting against the small oval
sclerite of the metathorax (the second tergal sclerite).

stough: the hackberry psylla. l4d

LEGS.

The prothoracic and mesothoracic legs are essentially the
same in structure, while the metathoracic leg is greatly spe-
cialized. The coxae of the former are not free and movable,
as in many insects, but immovably fixed to the thorax, and,
according to Witlaczil, coalescing with it. They are greatly
produced laterally, this portion being flattened and longer in
the mesocoxa than in the procoxa.

The trochanters are short, cylindrical sclerites, somewhat
constricted proximally. All three pairs are similar.

The two anterior pairs of femora are swollen, the outlines
of their dorsal surfaces somewhat convex, and the cuticula of
ventral surfaces much roughened. The femora of the protho-
racic legs are somewhat longer than those of the mesothoracic
pair. The distal ends are on their ventral faces deeply gouged
out, into which groove, bounded by strong lateral edges, the
tibise fit when the leg is flexed. The femora of the metatho-
racic legs are similar in structure to those of the other pairs,
but longer and relatively more slender. There was very little
variation of this part, as may be seen by the following table of
measurements :

AVERAGE LENGTH.

Male. Female.

Right. Left. Right. Left.

0.656 mm. 0.654 mm. 0.686 mm. 0.691 mm.

The articulation between femur and trochanter of meso-
thoracic leg is shown in figure 78, page 150. This is similar in
the other pairs.

The structure of the tibise is of great interest. The tibise of
the prothoracic and mesothoracic legs are about as long as the
femora, but those of the metathoracic legs are slightly longer
than the femora. The distal ends of the first and second pairs
of tibise bear a fringe of long, sharp spines which point distad,
those of the prothoracic pair being slightly the weaker. The
distal ends of the metatibise are swollen and bear nine short,
thick, blunt, black spines (figs 73, 74 and 75). These spines
are a part of the scheme of specialization for jumping, pre-
venting the foot from slipping in the same way as do the
spines on the grasshopper's tibise.

The tarsi are two-segmented. In the first two pairs of legs
the second segment is larger, but the first is the larger in the
metathoracic pair. In all three pairs the segments are much

150

KANSAS UNIVERSITY SCIENCE BULLETIN.

Pig. 71. > 1H7.

Pig. 79. Rloo

Pig. 77. » at"*

Fig. 72. — First metatarsal segment, c, claw; p, pad.

Figs. 73 and 74. — End of metatibia showing spines.

Fig. 75.— End of mesotibia.

Fig. 77. — Metatrochanter. sh, sensory hairs ; sp, sensory pits.

Fig. 78. — Articulation between coxa and trochanter mesothoracic leg.

cox, coxa; troc, trochanter; m, membrane.
P'ig. 79. — Prothoracic foot, c, claw; s, str, tendons moving claws.

constricted proximally. The first segment articulates with the
tibia by a joint much like that between femur and trochanter.
On the ventral surface of the first tarsal segment of the meta-
thoracic leg is a soft, membranous pad, projecting distad be-
tween the two stout, laterally placed claws (fig. 72) . This pad
is best developed in the metathoracic tarsi and these claws are
found only here.

The second segment bears the claws, which are similar in all
three pairs of legs. These are short, heavy, and rather blunt.

stough: the hackberry psylla. 161

with several small sclerites at their bases embedded in the
membrane. These are illustrated in figure 79, which shows
the tendons controlling their movement, and the two-lobed
empodium, one lobe being under each claw. This empodium
is functional, as in many insects, and is used in holding the in-
sect to a smooth surface, as it can cling readily to a glass sur-
face.

Femora, tibiae and tarsi of all three pairs of legs are cov-
ered with hairs, many of which are probably tactile. Both
coxae and trochanters (except of metathoracic legs) bear hairs,
the latter with one or two hairs on their distal ends longer
than the segments themselves. On the trochanters is a row
of oval sensory pits (fig. 77, sp).

ABDOMEN.

The relative shapes of the abdomen in the sexes can be
seen in 35-lS and 14 (figure inverted), also in 34.-4:. The
abdomen of the female is larger than that of the male and
seems much longer in proportion, as the ninth segment of the
male bearing the genitalia is generally carried bent dorsad
and cephalad so that the supra-anal plate fits up tightly against
the seventh segment, thus making the abdomen appear quite
short. In both sexes it is slightly compressed laterally. Ac-
cording to Edwards, there are visible five segments above and
six below, not counting the genital segment.

The first two segments are smaller than those caudad. "The
first abdominal somite of both sexes is added to the meta-
thorax to enlarge the springing-gear ; the second abdominal
somite forms a short stalk for the abdomen. . . ." —
(Macloskie.)

According to Witlaczil, in the male the supra-anal plate rep-
resents the tenth somite, the ninth is not represented above,
and below forms the subgenital plate; the eighth is scarcely
at all developed above ; in the female, the supra-anal and sub-
genital plates or valves represent the tenth segment.

In the pleural membrane are found seven spiracles, each
surrounded by a wide, chitinized ring. The cuticula of the
abdomen is punctate, not so pubescent as that of the thorax,
and the membrane between the chitinized plates of each seg-
ment is covered with great numbers of minute sensillia stylo-
conica. On the posterior edge of each segment is a row of

3-Univ. Sci. Bull., Vol. V, No. 9.

152 KANSAS UNIVERSITY SCIENCE BULLETIN.

sensory hairs, in prominent pits, much more evident, however,
in the male than in the female.

GENITALIA.

Male Genitalia. — The genital segment of the male of
Pachypsylla celtidis-mammse is described by Riley as follows :
"Male genital segment a little longer than the preceding ven-
tral segment, brown, shining; plate as high as the length of
the segment, lateral lobe barely indicated, anterior margin
straight, posterior margin very slightly oblique, i. e., the plate
gradually increases in width towards the tip, which is truncate
and not arctuate, as in the preceding species. Forceps as in
venusta [two-thirds as high as plate, front margin straight,
hind margin slightly sinuate at basal half, tip rounded, outer
face smooth and very shining]."

The genital segment of the male consists of the following
parts: Supra-anal plate or "genital plate," subgenital plate,
forceps or "posterior processes," and copulatory organ or
"sedeagus."

The supra-anal plate is a conspicuous fleshy process attached
to the anterior edge of the subgenital plate. It is generally
carried at about right angles to this plate, and is shown in
55-10 as drawn forward out of normal position. It averages
about 0.48 mm. in length (measured on cephalic surface) and
its greatest width is 0,19 mm. Looking cephalad at this plate
it is seen to taper at top and bottom, and to be convex with
rather wide flaps on its lateral edges (30-6, sppfi). At its
distal end these flaps recede to the edge, where the organ
ceases to be an open trough, and becomes a short, rounded,
truncate, cone-shaped structure (30-6 and 7, x) , connected
by its cephalic wall to the basal wall of the trough below. This
structure ends in a short, flaring, membranous ring (figs. 6
and 7, a), which is the external opening of the anus. The
concave face of the plate is lined with a delicate, folded mem-
brane, between which and the cephalic wall is situated a tube,
the rectum. Almost the entire surface of the plate carries
numerous long, stiff hairs.

The lower extremity of the supra-anal plate consists of a
chitinized ring which expands into a rather broad, two-lobed
plate, Ip, on each side. These plates drop ventrad at their inner
edges much after the fashion of a funnel, and form the base of
the copulatory organ.

stough: the hackberry psylla, 15^

Extending ventrad, with its dorsal edges articulating with
the lower side of each of the above-mentioned lobed plates, is
a chitinized band or loop (figs. 6 and 7, 0 extending down-
ward almost to touch the inner surface of the subgenital plate.
Between the arms of this loop the proximal portion of the
copulatory organ swing, almost entirely concealed within the
subgenital plate when the genital segment is telescoped into
the next segment cephalad and the supra-anal plate is forced
caudad. At emergence, however, the genital organs are ex-
tended and not telescoped, and at time of copulation the copula-
tory organ is protruded from the cavity of the subgenital plate.

Extending ventrad from the lateral plates at the base of the
supra-anal plate is the chitinized couplatory organ or extremity
of and sheath for the penis — "the sedeagus" of some writers.
This organ (figs. 6 and 7, co, and 35-1), soon after it leaves
the base of the supra-anal plate, forms a sharp bend, reversing
its direction to dorsad. After reaching a position about oppo-
site the middle of the supra-anal plate there occurs a conspicu-
ous swelling, and beyond that a joint or geniculation {30-
Q, g). The proximal portion of the second segment of the
organ is also swollen. This segment in normal position drops
ventrad. The distal end bends caudad and is swollen greatly,
becoming club-shaped (fig. 6, c), convex cephalad, and con-
cave caudad, this segment having a vertical orientation. The
convex, cephalic face of the club is gouged out to form a deep
longitudinal groove running from base to tip.

The proximal portion of the first segment of the copulatory
organ is not tubular, but slitted on its convex surface, and
through this slit the penis enters the sheath, p. The concave
surface of this portion is coarsely notched or corrugated trans-
versely for some distance beyond the bend. The remainder of
the organ is tubular, the canal running through the inner half
of the sheath — that is, through the cephalic half of the first
segment and through the caudal half of the second segment.
This canal runs into the center of the club, makes a sharp turn,
and, proceeding back upon itself, curves to the convex side of
the club, where it issues at the base of the club in the bottom of
the groove above mentioned.

The copulatory organ is more strongly chitinized at its distal
end than it is in the curved portion of the proximal segment,
and along its outer surface, than on its inner. The first seg-
ment measures (from base to geniculation, the distance along

154 KANSAS UNIVERSITY SCIENCE BULLETIN.

curved side being much greater) 0.81 mm., the distal segment
0.38 mm., and the club 0.12 mm.

The subgenital plate has already been mentioned in the quo-
tation from Riley. Its measurements are as follows : Depth,
0.37 mm. ; length, 0.40 mm. ; greatest width, 0.26 mm.

Situated at the posterior end of the subgenital plate are the
forceps (fig. 10, /", and 35-16). These two pieces are some-
what swollen at their bases, seem to be articulated one against
the other, and are connected to the subgenital plate by means of
a long, thin, tendonous strip, which reaches ventrad to the
inner surface of the plate, where it is fastened. The forceps
are about 0.29 mm. in length, spread at widest 0.29 mm., and
project above the subgenital plate 0.20 mm.

Female Genitalia. — The genital segment and ovipositor of
the female are of great morphological interest. The genital
plates have been described by Riley as follows: "Female
genital segment a little shorter than the three preceding ven-
tral segments together; . . . upper plate a little longer
than the lower one, gradually tapering toward the tip, which
is straight; . . . lower plate also simple." This is in
substance all that I have found in the literature to which I have
had access concerning the female genitalia of this family. The
genital valves or plates, being of taxonomic value, have gen-
erally been included in descriptions of species, but the oviposi-
tor remains unmentioned, except in a few cases where it is
spoken of as "ending in an acute point" (Edwards). West-
wood (1839) mentions it as a "pleurivalve conical ovipositor."
I have been unable to find anywhere a treatment of its struc-
ture.

I shall first treat of the two plates mentioned above, and then
consider the structure of the ovipositor proper.

Of these plates, the supra-anal is of the greater interest. It
is wedge-shaped, with its apex blunt and its i)roximal end three-
sided. It averages in length 0.98 mm. and in width 0.42 mm.
Its shape is well shown in 55-4, where the ovate openings
at its basal angles can be seen. These latter may be of use in
attaching the plate to the abdomen. The plate is somewhat
convex but not so much so as the subgenital plate. It bears on
its upper surface sensory hairs, probably tactile, and is (luite
heavily chitinized except for a large oval space, //, about 0.26
mm. long surrounding the anal opening, where the plate is

STOUGII : THE HACKBERRY PSYLLA. 155

almost transparent. Along the median line in this space is a
slit, into the cephalic end of which the anus opens ; this could
be seen in a longitudinal-vertical section, where the rectum
was found to run just under the dorsal surface of the posterior
end of the abdomen and end at the cephalic end of this slit.
About midway between this slit and the edges of the clear
space are situated the curiously ornamental openings of the
circumanal wax glands, outside of which are rather broken
and irregular rows of short, heavy, curved, sensory hairs in pits.
The dorsal aspect of this region of the plate is represented
in 31-4:. This encircling band of wax glands is double;
their openings are ovate to rectangular, with borders measur-
ing 0.015 mm. in width, and are seen in the greatly magnified
portion shown in 26-5 B. Longitudinal-vertical sections
showed the glands themselves. These are of large size, flask-
shaped, with long necks, and are closely packed in a double row
under the dorsal openings. They are illustrated in section in
figure 5 A, irg. Their nuclei, n, could in most cases be made
out quite plainly. These glands are of hypodermal origin, and
are homologous to the circumgenital glands or spinnerets found
in the Coccidse. They are functional in the nymphal stages of
most Psyllidse, and in those forming galls the wax enters into
the composition of the gall, making an impervious lining for
the cavity, while it is found as a stringy, exuvial covering to
some of the nongall-formers. These glands are functional in
the nymph of Pachypsylla, but I have no evidence that they are
so in the adult.

The subgenital plate is much broader and more concave than
the supra-anal plate, though not so long as the latter. Its
shape when flattened out is shown in 27-S. The anterior
margin is almost straight, except laterad, where it turns caudad
and joins the lateral margin, forming an acute angle. The
apex is blunt, as in the supra-anal plate. The approximate
measurements of the plate are : Width at base, 0.73 mm. ;
median length, 0.63 mm. ; length of lateral margin, 0.67 mm.
The plate is covered with rather stiff" hairs, probably tactile,
more numerous at the posterior end and wanting at the anterior
end; it is almost as heavily chitinized as the supra-anal plate.

Ovipositor. — The insects of the suborder Homoptera possess
a well-developed ovipositor. This is well preserved and of
quite complicated structure in the family Psyllidse.

156 KANSAS UNIVERSITY SCIENCE BULLETIN.

The typical structure of an ovipositor is described in the fol-
lowing quotation from Packard : "Morphologically, the ovi-
positor is composed of three pairs of unjointed styles (rhab-
clites of Lacaze-Duthiers, gonapophyses of Huxley), which are
closely appressed to or sheathed within each other, the eggs
passing out from the end of the oviduct, which lies, as Dewitz
states, between the two styles of the lowest or innermost pair
and under the cross-bars or at the base of the stylets men-
tioned ; the styles or blades spreading to allow of the passage
of the es^.''

The structure of this organ I find to be second in interest
only to the mouth parts. It is more complicated in structure
than those parts in the cicada, but the homologies between the
two can be worked out in part, as will be seen later. It is very
similar in most points to the ovipositor in the Aleurodidae.

Lack of material has confined my work to postembryonic
studies. There has been much investigation on the segmental
origin of the ovipositor parts, the researches of Wheeler and
Heymons being of chief importance. Wheeler (189^), who
studied orthopteran embryology, holds for the "direct con-
tinuity of the embryonic appendages with the gonapophyses,"
while Heymons (1899), from studies in Heteroptera and
Homoptera, considers these gonapophyses as hypodermal out-
growths. However, it is not my purpose to discuss these ques-
tions, but to treat of the anatomy and homologies of the parts.

A specimen of Pachypsylla female, mounted in balsam after
having been thoroughly boiled in caustic potash and cleared,
showed, when viewed from the side, the relative position of
the ovipositor as regards supra-anal and subgenital plates, and
the posterior abdominal segments (30-4). The ovipositor is
seen to be an elongate organ with dorsal and ventral outlines
about parallel. It measures in length about 0.82 mm., in
greatest width about 0.19 mm., and in thickness less than half
that of width. The caudal extremity is acutely pointed, this
portion bending dorsad, then caudad, with its apex extending
caudad even with the tip of the subgenital plate, while the
supra-anal plate extends caudad from this apex 0.11 mm. in the
specimen described. The proximal or cephalic end of the ovi-
positor is composed of several sclerites of irregular form, to be
described later, the ends of which project 0.16 mm. beyond a
line joining the bases of the enclosing external plates to a

stough: the hackberry psylla. 157

position about opposite and above the last spiracle, well into
the dorsal portion of the last abdominal segment. Its position
relative to the two plates is shown in figure 1 B, plate XXVI,

Seven longitudinal rods or valves are to be found running
throughout the organ. Six of these are paired, two pairs form-
ing the lateral edges of the ventral face of the organ, and a
third pair lateral and well towards the dorsal face ; the seventh
and unpaired rod is dorsal and median.

Each upper and each lower valve of the two ventral pairs are
very closely connected, being held together throughout their
entire length by means of a tongue on the upper fitting into a
groove in the lower, which may be seen in the cross section in
26-1 A, tg. The tips of these valves are very sharp and
finely pointed, and come together at their tips, where they are
enclosed in a sheath. The cephalic ends of each ventral and
inner stylet is connected with a more or less crescent-shaped
sclerite (26-1, blv) . The cephalic end of each ventral and
outer stylet connects with a sclerite (fig, 1, buv) which runs
dorsad and caudad. When this reaches about the dorsal face
of the ovipositor it turns caudad and becomes a thin rod, Ir,
and at about the middle of the ovipositor becomes compressed
laterally to form a wide, thin plate, which gradually loses its
chitinization, and is lost in the fleshy tongues on each side of
the ovipositor sheath (figs. 1, 2 and 3, stp) .

Situated medially and a little dorsad of the above-mentioned
rods is a rather thick, cylindrical rod bent at its cephalic end
slightly dorsad, then ventrad (fig. 1, mr) . This rod does not
project quite as far cephalad as the stylets below. It is greatly
swollen at its caudal extremity, with a square end, and its ven-
tral portion is continued into a slender process, in shape much
like a human femur, procv. This last-named process works
against a ventrad-projecting outgrowth of the sheath sur-
rounding the stylet tips, which serves to close the oviduct (fig,
1, v) . Lying above this slender sclerite are two pairs of chit-
inized sclerites (figs, 1 2 and 3, sc 1 and sc 2), which are
connected with and probably serve to bring about lateral ad-
justment of the lateral tongues above mentioned. Viewed
laterally, these sclerites appear as one pair (fig. 1), but when
viewed from above, and pressure was applied to the dissec-
tion so as to spread the sides apart, they were seen as two
distinct pairs of sclerites. Viewed in this manner, the sclerites

158 KANSAS UNIVERSITY SCIENCE BULLETIN.

of the cephalic pair, sc 1, are pointed, with convex inner sur-
faces. Their tips have a connection with the lateral plates
(figs. 1, 2 and 3, pi), the latter probably serving to spread
these sclerites apart. The caudal pair, sc 2, reach almost to
the base of the sheath surrounding the stylet tips, are rectan-
gular in shape, with rounded ends, and with their inner sur-
faces corrugated, the corrugations corresponding to the
striations on the surfaces of the lateral tongues of the ovi-
positor.

The sheath already mentioned (figs. 1, 2 and 3, sh) is
slitted on its dorsal surface from near its cephalic edge to the
apex of the ovipositor, to allow of the spreading of the stylets
and the passage of the egg during oviposition. The cephalic
portion of its dorsal face carries the small, valve-like, ventrad-
projecting process (fig. 1, v).

Fig. 1, pi. 26, shows a cross section about midway through
the organ. In this the fleshy structures that were destroyed in
boiling in caustic potash can be made out. The grooved and
toothed valves, uv and Iv, are seen fitted together, and the
chitinized portion, x, which is grooved and forms the sides of
the oviduct. Above the oviduct, and forming its roof, is the
structure, y, whose ventral surface is covered with minute,
soft, hair-like outgrowths, and whose dorsal portion is chitin-
ized. The part marked z probably represents the lateral
plates, Ipt, of 30-1, 2 and 3. The duct of the cement (?)
gland is seen at cmgd in this figure and appears in all sec-
tions cephalad of this, and in those caudad up to the point
where it turns ventrad and enters the oviduct. The entire
course of this duct was best seen in longitudinal-vertical sec-
tions, but could be traced well in longitudinal-horizontal sec-
tions, where its union with the oviduct was plainly seen, and
in the next succeeding sections ventrad the oviduct was laid
bare. The cement (?) gland itself, a small sac, was found
somewhat cephalad of the base of the ovipositor in most of
my dissections. Figure 6, plate XXXI, is of a cross section
near the base of the ovipositor sheath. Here the shape of the
supra-anal and subgenital plates can be seen, also the position
of the oviduct. The lateral tongues or sting-palpi are also
represented.

I have pointed out the structure of these parts and shall
now give my theories as to the homologies. The typical struc-

STOUGH: THE HACKBERRY PSYLLA. 159

ture of an ovipositor as being composed of several pairs of
stylets has already been mentioned.

To homologize the structure of the ovipositor of Pachypsylla
with the structure of the typical ovipositor is more difficult
than the comparison of it with that as seen in the different
families of the Homoptera. I find that it is more specialized
than a typical ovipositor such as found in the Orthoptera,
though it is possible to show the homologies; but since it re-
sembles more closely that found among the Hymenoptera —
where it is specialized to form a sting — these homologies can
be traced through the latter. Taking the sting of the honey-
bee as a type of the hymenopterous ovipositor, since this has
been so carefully worked out by Cheshire and others, it can be
seen that the lower valves of Pachypsylla (30-1 and 3, Iv)
correspond to the lower and outer pair of darts in the honey-
bee. The upper valves of Pachypsylla correspond to the inner
pair in the bee, where they are fused into one piece to form the
sheath. It should be noted here that the sheath does not corre-
spond to that part of the ovipositor of Pachypsylla which I
have designated by that name. In the honeybee there are two
fleshy processes, one on each side of the sting, which are
thought to represent the upper pair of stylets and are called
the "sting-palpi" or "feelers." These are evidently the same
as the lateral tongues found in Pachypsylla.

The homologies up to this point are comparatively clear,
but as to the remainder of the parts I can only theorize, as
their structure is complicated and there are present more
pieces than in the typical organ. The lateral tongues evidently
are the "sting-palpi" and together with the lateral rods and
lateral plates form the upper pair of stylets; but this expla-
nation does not account for the median rod with its processes
and appendages. There may be three possible explanations
for these structures:

1. This median rod might be regarded as a fusion of the
dorsal pair of stylets, and the rods and lateral plates only as
tendons causing the spreading of the sc 1 and sc 2 and conse-
quently the sting-palpi.

2. The rods and lateral plates might be considered as com-
posing the dorsal pair of stylets and the median rod as an
accessory structure which, moving backward and forward,
would spread and close the sting-palpi, for these latter are

160 KANSAS UNIVERSITY SCIENCE BULLETIN.

evidently in as intimate a relation to the median rod as the
lateral rods. Hence, there is possible a third theory:

3. According to Lacaze-Duthier's view, there are "two
pieces forming the outer pair of rhabdites" (after Packard),
as seen in the figure illustrating his conception of the ideal
ovipositor. In Pachypsylla, one of these "pieces" might be
the rod and the lateral plate which connects basally with the
lever supporting the base of the upper valve, buv, and distally
with the tongues or sting-palpi. The median rod, then, might
represent the second of the "pieces," its two parts fused into
one. Thus rods and lateral plates, together with the sting-
palpi, would form one portion of the upper pair of valves
while the median rods represent the other paired portion,
fused. The sheath may also belong to the second piece of the
upper valves, as do the two pairs of small sclerites, sc 1 and
sc 2 (30-2). The levers supporting the bases of the lower
valves, blv, find their homologues in similar structures in the
honeybee, termed there simply "levers," which are the point
of attachment of muscles moving the sting. They have a sim-
ilar function here, serving to spread the styles.

The ovipositor of the cicada has been described and figured
by Marlatt (1895). A rather superficial study of this form
from my own dissections has rendered the following facts
and hypotheses as to the homologies between this form and
Pachypsylla: There are found two pairs of blades, the dorsal
pair grown together, and the two pairs sliding on one another
by means of tongues and grooves much as I have shown to be
the case with Pachypsylla. The tips of the ventral pair of
blades are serrated for cutting purposes. These two pairs of
blades evidently correspond to the upper and lower valves or
styles ot Pachypsylla, though in the latter there is no need for
their tips to be modified for cutting, as the eggs are not de-
posited within plant tissue. In Cicada there is a very large
sheath surrounding the entire ovipositor, except for a slit be-
neath, which probably corresponds to the supra-anal plate,
Marlatt says, "The ovipositor is protected and covered when
at rest by two valves, which form a sort of sheath or scab-
bard." These second valves mentioned by Marlatt, which are
dorsad of the blades, proximad are broad, and made up of
chitinized and more or less membranous portions, and distad
are composed of two distinct and distally separable scabbards

stough: the hackberry psylla. 161

covering the tips of the blades dorsally and laterally, and open
entad and ventrad. The basal portion is evidently a complex
made up of parts corresponding perhaps to the median rod,
lateral rods and lateral plates, with their processes, append-
ages, etc., of Pachypsylla, while the two scabbards may be
homologous to the sheath. The sting-palpi are not repre-
sented in Cicada.

The ovipositor of one species of Aleurodidse which I have
examined showed features most similar to those of Pachy-
psylla of any other insect, the parts being much more easily
compared with those of this insect. The upper and lower pairs
of converging blades or styles were found, together with a
sheath enclosing their apices, while their bases bore irregu-
lar supports as in Pachypsylla. Sting-palpi and sc 1 and sc 2
were also present.

GENERAL SUMMARY.

1. The head is made up of the typical sclerites.

2. The mouth parts conform in general to those of other
homopterous forms, such as those of the Cicadidse, Aphididse,
Aleurodidse, and most closely to those of the Aleurodidse, differ-
ing most from those of the Coccidse.

3. The mesoterga and metaterga show three sclerites only.
The pleural sclerites are much folded, with their sutures almost
obsolete. The internal skeleton of mesothorax and metathorax
is large and heavy, making the segments very rigid.

4. The metathoracic legs are greatly specialized for jump-
ing, the tibise being provided with heavy spines on their distal
ends, and the coxa, epimeron, meracanthus, a part of the
sternum, and an accessory sclerite compose a single complex
of sclerites which encases powerful jumping muscles.

5. A study of the tracheation of nymphal wing pads and
wings of recently emerged adults has shown that there are
present in the adult wing the following veins : one costa ; one
subcosta ; one radius ; one two-branched media ; one two-
branched cubitus ; one anal vein ; and a vein occupying the
anal fold, seen only in nymphal wings, being represented only
as a fold in the adult wing. Previous writers have not indi-
cated the media and cubitus as separate veins, but I have shown
them to be represented by separate parallel trachese in the wing
pads. The subcosta of the hind wing, previously unfigured,
was found in recently emerged specimens. A sport vein, pos-

162 KANSAS UNIVERSITY SCIENCE BULLETIN.

sibly representing the second anal vein, was found in one
specimen.

6. The color pattern was found to consist of polygonal or
irregular groups of pigmented, disklike outgrowths of the
cuticula. The manner of producing the pattern is various.

7. The male genitalia were found to consist of supra-anal
plate, subgenital plate, couplatory organ and forceps.

8. The female genitalia consist of supra-anal plate (orna-
mented with a double row of wax glands, flask-shaped in longi-
tudinal section), subgenital plate and ovipositor. The ovi-
positor was found to be homologous in many points to a
hymenopterous ovipositor as seen in the honeybee (a sting).
Comparison with other homopterous forms showed it to be
homologous in most points with that of the cicada, but almost
identical to that of the Aleurodidse, showing, with the buccal
appendages, the close structural relationship of the Psyllidse
to that family.

TECHNIQUE.

My studies have been based upon dissections and serial sec-
tions.

Dissections were made in three ways: (1) A dissecting
stand fitted with an 0.8-inch lens was used in some cases; (2)
some of my work was done under a compound microscope (one-
inch eye-piece and one-inch objective) on the lower end of
whose draw-tube was placed a three-inch objective, thus pro-
ducing an erect image. This combination was useful but its
disadvantage was that the eye was too far from the specimen
being dissected on the microscope stage; (o) the greater part
of my dissecting, however, was done under a "Pfeiffer's Erect-
vision Dissecting Microscope" using objectives three inch, one
inch and two-thirds inch. This was found to be the most con-
venient, especially in the more delicate dissections, such as of
mouth parts and genitalia. By the use of very fine, sharp-
pointed needles, these minute organs could be torn apart with
ease.

Most of my material was boiled for a short time in caustic
potash, and then treated with alcohol and xylol previous to
dissections, no fresh specimens being available at the time
when this work was done. Xylol was of great use in clearing
the parts, but at the same time it rendered them brittle and
easily broken.

Dissections were made under water, alcohol, xylol and glyc-

stough: the iiackberry psylla. 163

erine. Xylol was a very clear medium, having the advantage
that uncleared specimens could be dissected in it, soon becoming
clear, but it evaporated very quickly, and, lacking viscosity, al-
lowed too much movement of the parts being dissected. Glyc-
erine was found to possess the necessary viscosity and was the
most satisfactory medium in most cases.

The nymphal wing pads were removed with fine scissors,
holding the nymph the while between the finger-tips ; the pads
were mounted in dilute glycerine after being kept in formalin
for about twenty-four hours.

Serial sections were made through the mouth parts and geni-
talia, and cross, longitudinal-horizontal and longitudinal-ver-
tical sections through the entire insect.

Killing and Fixing. — Live insects were killed by plunging
into a boiling saturated solution of corrosive sublimate con-
taining about one per cent of acetic acid, and after being
punctured with a piece of glass drawn out into a fine point
were left in the solution to fix for twenty-four hours.

Dehydrating and CleaHng. — The insects were placed in
small vials with ends covered with cheesecloth, and run through
alcohols of percentages 70, 85 and 95, twenty-four hours in
each. They were removed to absolute alcohol for about twelve
hours and then run into xylol for about twelve hours.

hifiltration and Embedding. — Leaving the specimens still in
the vials they were infiltrated in a paraffin oven with paraffin
of melting-point 50-55" C. for forty-eight hours. They were
then embedded in paraflfin in small paper dishes, being oriented
properly as the paraffin hardened when the dish was floated on
cold water.

Sections. — Serial sections 10-20 microns thick were cut on a
sliding microtome. The ribbons were floated onto Mayer's
albumen fixative on a clean slide, the excess of fixative drained
off" with filter paper, and the slide dried on top of the paraffin
oven for twelve or more hours. The slide was then left for
ten to fifteen minutes in xylol to remove the paraffin, when it
was run through the following grades of alcohol, five to ten
minutes in each : 95 per cent, 85 per cent, 70 per cent, 50 per
cent, 35 per cent, 25 per cent, and then into water, after which
the sections were stained in Mayer's Carmalum, ten to fifteen
minutes, washed in water and run back through the alcohols,
beginning with twenty-five per cent, into absolute alcohol, five

164 KANSAS UNIVERSITY SCIENCE BULLETIN.

to ten minutes in each. They were then cleared in xylol and
mounted in balsam.

In running the sections into and out of the stain (aqueous)
the lower grades of alcohol, 25 per cent and 35 per cent, and
then water, were necessary, as without their use the transfer
into the stain from 70 per cent alcohol was found to be too
sudden, causing currents which tore the sections loose from
the slide. Any dirt or grease on the slides made this more cer-
tain. However, cleaning the latter in 95 per cent alcohol con-
taining a few drops of hydrochloric acid obviated this.

BIBLIOGRAPHY.

AsHMEAD, W. A., 1881. On the Aphididae of Florida. Can. Ent., vol.
XIII, 1881, pp. 220-225.

Berlese, a., 1896. Le Cocciniglie Italiane Viventi Sugli Agrumi. Parte
II, plate VIII, fig. 1; parte III, figs. 180, 181.

COMSTOCK, J. H., and KocHi, C, 1902. The Skeleton of the Head of
Insects. Amer. Nat., vol. XXXVI, No. 421.

CoMSTOCK, J. H., and Needham, J. G., 1898-'99. The Wings of Insects.
Amer. Nat., vols. XXXII and XXXIII.

Cook, M. T., 1903-'04. Galls and Insects Producing Them. Parts III,
IV and V, Ohio Nat., vol. Ill, pp. 419-436; parts VI-IX, Ohio Nat.,
vol. IV, pp. 125-147.

DoHRN, A., 1865. Quaedam de anatomia hemipterorum. Uratislaviae.

Edwards, J., 1896. Hemiptera-Homoptera, pp. 233-261.

Froggatt, W. W., 1903. Australian Psyllidae, part III. Proc. Linn. Soc.
N. S. W., 1903, part 2.

Geise, O., 1883. Die Mundtheile der Rhynchoten, Bonn.

Heymons, R., 1899. Beitrage zur Morphologie und Entwicklungsge-
schichte der Rhynchoten, Halle.

Leon, N., 1887. Beitrage zur Kenntnis der Mundtheile der Hemipteren,
Jena.

Low, Fr., 1879. Zur Systematic der Psylliden. K. K. Zool.-bot. Gesell-
schaft in Wien, 1882.

Macloskie, G., 1886. Witlaczil on Psyllidae. Amer. Nat., vol. XX,
March, 1886.

Mally, C. W., 1894. Hackberry Psyllidas found at Ames, Iowa. Proc.
la. Acad. Sci., 1893, vol. I, pt. IV, pp. 131-138.

STOUGH: THE HACKBERRY PSYLLA. 165

Mally, C. W., 1895. Psyllidas found at Ames. Proc. la. Acad. Sci., 1894,
vol. II, pp. 152-171.

Marlatt, C. L., 1895. The Hemipterous Mouth. Proc. Ent. Soc. Wash.,
vol. Ill, pp. 241-249.

Maskel, W. M., 1889. On Some Species of Psyllidae in New Zealand.
Trans. N. Z. Inst., vol. XXII, 1889, art. XVII, pis. X-XII, pp. 157-170.

Meek, W. J., 1903. On the Mouth Parts of the Hemiptera. Kan. Univ.
Sci. Bull., vol. II, No. 9.

Packard, A. S., 1887. Hackberry Psyllidae. Fifth Rept. U. S. Ent. Com.,
pp. 614-622.

Reiber, F., 1875. Translation of Dr. Franz-Xavier Fieber's "Les Cica-
dines D'Europe."

Riley, C. V., 1893. The Pear Tree Psylla. Insect Life, vol. V, pp. 226-
230.

Riley, C. V., 1889. Note on work done by Low and his predecessors on
the Psyllidae. Insect Life, vol. II, No. 6, p. 196.

Riley, C. V., 1883. Hackberry Psyllid Galls. Can. Ent., 1883, vol. 15,
pp. 157-159.

Riley, C. V., 1883. The Psyllidae of the United States. Science, 1883,
vol. 2, p. 337.

Riley, C. V., 1884. Notes on North American Psyllidae. Proc. Biol. Soc.
Wash., 1884, vol. II, pp. 67-79.

Sanborn, C. E., 1906. Kansas Aphididae. Kan. Univ. Sci. Bull., vol. Ill,
No. 1.

ScuDDER, S. H., 1890. Tertiary Insects of North America. Plate 12, figs.
11 and 21; pp. 276-278.

Slingerland, M. v., 1896. The Pear Tree Psylla. Bull. 44, Cornell
Univ. Agr. Exp. Sta.

Smith, J. B., 1892. Structure of the Hemipterous Mouth. Science, vol.
19, No. 478.

Snodgrass, R. E., 1909. The Thoracic Tergum of Insects. Ent. News,
vol. XV, No. 3, pp. 97-104.

Snodgrass, R. E., 1907. A Comparative Study of the Thorax in Orthop-
tera, Euplexoptera, and Coleoptera. Proc. Ent. Soc. Wash., vol. IX,
Nos. 1-4.

Tucker, E. S., 1907. Some Results of Desultory Collecting of Insects in
Kansas and Colorado. Kan. Univ. Sci. Bull., vol. IV, No. 2.

Wedde, H., 1885. Beitrage zur Kenntniss des Rhynchotenriissels, Berlin.

Wheeler, W. M., 1893. A Contribution on Insect Embryology. Journal
of Morphology, vol. VIII, No. 1.

WiTLACZiL, E., 1885. Die Anatomie der Psylliden. Zeitschrift fiir
Wissenschaftlichen Zoologie, vol. XLII, pp. 560-658.

WooDWORTH, C. W., 1906. The Wing Veins of Insects. Univ. of Cal.
Publications, vol. I, No. 1, pp. 122-125.

Zoological Record, 1886. Frauenfeld on Anisostropha ficus Linn, in
Verh. zool.-bot. Ges. in Wien, XVII, pp. 801-803.

THE

KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 10 — April, 1910.

(Whole Series, Vol. XV, No. 10.)

CONTENTS:

Investigations Regarding the Phloem and Pood-conduction

IN Plants, Frank U. G. Agrelius.

PUBLISHED BY THE UNIVERSITY
LAWRENCE, KAN.

Entered at the post-office in Lawrence as second-class matter.

THE KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 10] APRIL, 1910. [r.'xvSTo

INVESTIGATIONS REGARDING THE PHLOEM AND
FOOD-CONDUCTION IN PLANTS.

BY FRANK U. G. AGRELIUS.
Plates XXXVI-XXXVII.

THIS work is an inquiry into the special devices for the
lateral transfer of food materials in the phloem of vascu-
lar plants; to see whether sieve tubes are always present in
the higher plants ; and to make some comparative studies of the
phloem in stems, petioles and fruitstalks.

At first material in the winter condition was tried but did
not prove satisfactory, on account of the difficulty of identify-
ing the sieve tubes in their winter condition. Material was
again collected, this time including stems, petioles and fruit-
stalks, from the spring and summer growth, where the sieve
tubes were functioning. Vines were selected at first, from the
supposition that the conducting elements would be better de-
veloped and therefore plainer. This material was preserved in
two-per-cent formaldehyde. As shrinkage of the protoplasm
would really assist in making the cell walls more distinct, but
little attempt was made to prevent it in the various processes
of preparation.

The celloidin method of imbedding was tried at first but it
did not serve the purpose well. It was rather essential in the
hard material first collected, but the paraffin method worked
better in most cases. By the celloidin method slides having
the required large number of sections upon them were difficult
to make. If the celloidin were left in the sections it stained
too much like the cellulose walls. If dissolved out the cuttings
were liable to cupping or to injury in the region of the cam-
bium. These facts led to the choice of the paraffin method,

(169)

170 KANSAS UNIVERSITY SCIENCE BULLETIN.

which was used where possible and which proved quite satis-
factory in the majority of cases. The oldest material possible
was commonly used.

Stains adapted to the work in hand were not found at first.
Iodine green and erythrosin were tried, but this method left the
phloem red or pink — a poor color for the purpose. Finally
fuchsin for the lignified walls and methyl blue for the cellulose
ones were found to be quite satisfactory. The latter gave a
fairly dark wall and slime body — the one better for making the
drawings, the other rendering the identification of the sieve
tubes less diflficult.

The sections of such varied kind as to subject and age of
material stained quite differently under the same treatment, so
that each one usually required some variation. The slides were
left in the fuchsin from two hours to several days. The excess
stain was removed with acid-alcohol, and the slides were then
placed in methyl blue for from twenty seconds to three or four
minutes.

As a rule, cross, radial and tangential sections, with many
mounts on each slide, were prepared for each subject — in all,
more than 300 slides. These included specimens from young
and from somewhat older material. The tangential sections
proved the best for demonstrating sieve tubes. The cross sec-
tions were best for the measurements of cell walls and cell
cavities, and to show the locality of the various tissues, etc.
All were used to determine, if possible, the presence of pits, or
of any other facts bearing on the problem.

PLANTS EXAMINED.

The following is a list of the plants examined, arranged ac-
cording to the classification of Gray's revised manual :

Pinaceae:

Pinus laricio Port.
Alismaceae:

Sagittaria latifolia Willd.
Gramineae:

Zea Mays Linn.
Cyperaceae:

Scirpus validus (Vahl.) I
Bromeliaceae:

Tillandsia usneoides L.
Liliaceae:

Yucca glauca Nutt.

Asparagus officinalis L.

Smilax rotundifolia L.

Salicaceae:

Salix alba, var. vitellina (L.)
Koch.

Populus alba L.

Populus deltoides Marsh.
Juglandaceae :

Juglans nigra L.
Urticaceae:

Ulmus fulva Mx.

Ulmus americana L.

Celtis occidentalis L.

Cannabis sativa L.

Humulus Lupulus L.

Morus rubra L.

AGRELIUS: FOOD-CONDUCTION IN PLANTS.

171

Aristolochiaceae:

Aristolochia macrophylla Lam.

Aristolochia tomentosa Sims.
Ranunculaceae:

Clematis Pitcheri T. & G.
/inonaceae:

Asimina triloba Dunal.
Menispermaceae :

Menispermum canadense L.
Saxifragacese :

Deutzia Fortunei Hors.
Rosaceas :

Spiraea japonica L. (?)

Pyrus Malus Linn.

Fragaria virginiana Deuchesne.

Rubus occidentalis L.

Rubus cuneifolius Pursh.

Rosa pratincola Greene.

Prunus americana Marshall.
Leguminosge :

Gymnocladus dioica (L.) Koch.

Gleditsia triacanthos L.

Melilotus alba Desr.

Amorpha fruticosa L.

Desmodium canescens (L.) DC.
Rutacese :

Zanthoxylum americanum Mill.
Simarubacese:

Ailanthus glandulosa Desf.
Anacardiaceae:

Rhus glabra L.

Rhus toxicodendron L.

Rhus cotinus L.
Celastraceae:

Celastrus scandens L.
Aceraceae :

Acer saccharinum L.

Acer Negundo L.
Rhamnaceae:

Rhamnus lanceolata Pursh.
Vitaceae :

Psedera quinquefolia (L.)
Greene.

Vitis asstivalis Mx.

Tiliaceae:

Tilia americana L.
Malvaceae :

Abutilon Theophrasti Medic.
Cactaceae :

Opuntia Rafinesquii Engelm.
Haloragidaceae :

Myriophyllum heterophyllum Mx.
Cornaceae :

Cornus asperifolia Mx.
Oleacese :

Fraxinus viridis Mx. (?).

Syringa vulgaris L.
Asclepiadaceae:

Asclepiodora viridis (Walt.)
Gray.

Asclepias syriaca L.
Convolvulaceae:

Cuscuta glomerata Choisy.
Labiatae :

Monarda fistulosa L.
Solanaceae :

Lycium halimifolium Mill.
Scrophulariaceae :

Verbascum Thapsus L.
Bignoniaceae:

Tecoma radicans (L.) Juss.

Catalpa speciosa Warder.
Plantaginacese :

Plantago virginica L.
Caprifoliaceae:

Symphoricarpos orbiculatus
Moench.

Triosteum perfoliatum L.

Sambucus canadensis L.
Cucurbitaceae:

Cucurbita pepo L.
Compositae :

Brauneria pallida (Nutt.) Brit-
ton.

Gaillardia grandifloi'a ?

Arctium Lappa L.

Cirsium discolor (Muhl.) Spring.

Lepachys pinnata (Vent.)

These range from the Pinacese to the Compositse. They in-
clude one parasite, one epiphyte, three hydrophytes and two
xerophytes. The list embraces twenty-three herbaceous plants
(not including vines), five herbaceous vines, ten shrubby vines,
thirteen other shrubs and twenty-one trees. These are em-

172 KANSAS UNIVERSITY SCIENCE BULLETIN.

braced in thirty-nine families, sixty-five genera, and seventy-
two species.

DRAWINGS.

In general, camera-lucida drawings have been made to show
the pits in the phloem when found, to show sieve tubes and
some of the adjacent tissues from three points of view. A
Bausch and Lomb 1/12 objective was used with but few ex-
ceptions, and for measuring a Bausch and Lomb micrometer
eyepiece — divisions 1/10 mm. In all about 175 drawings have
been prepared for use in evidence.

FINDINGS.

Problem I. — What special devices occur for the lateral
transfer of food materials in the phloem ?

After considerable search I have found pits in the lateral
walls of the phloem of fourteen plants, as shown, for example,
in figures 1 and 2. In figure 1 they are evidently between the
medullary ray and the phloem ; in figure 2, between the sieve
tubes and the adjoining cells.

Frank* seems to imply that there are no pits between the
parenchyma cells of the phloem. Figure 1 shows some as they
appeared between the parenchyma and the parenchyma, and
between the parenchyma and medullary-ray cells of the stem
of Popultis alba. He also states that the sieve tubes are the
widest phloem elements.- This is hardly true in quite a num-
ber of instances, as in Amorpha fruticosa (fig. 6), where the
parenchyma is the wider. Quoting from Briosi,^ he states that
the slime bodies of the sieve tubes sometimes contain starch
granules. Some are shown in figure 7 in Populus alba. They
were noted also in Rhus toxicodendron. Frank states also^
that the lateral walls of the sieve tubes are never strength-
ened. Figures 4 and 5 show thickenings on the lateral walls
of the sieve tubes in the roots of Populus deltoides. It would
seem that these are for strengthening the wall. They occur
also in the end plates of the sieve tubes of the same plant
(figs. 8 and 9) and in Populus alba.

Strasburger'^ finds pits between the phloem parenchyma and
the medullary rays. Figures 1, 2 and 3 show such. He also
states® that the cortical rays are specially adapted by means

1. Frank, A. B. : Lehrbuch der Botanik, 1892, vol. I, p. 184.

2. Loc. cit., p. 183.

3. Loc. cit, p. 183.

4. Loc. cit., p. 183.

5. Strasburprer, Ed., Textbook of Botany, 1897, p. 135. (Trans, by Porter.)

6. Loc. cit., p. 186.

AGRELIUS: POOD-CONDUCTION IN PLANTS. 173

of these pits and their contact with the companion cells for
taking up and transferring the food materials laterally, espe-
cially to the cambium. Figure 6 shows a very common ar-
rangement of the medullary rays which bears out the rest of
his statement. It may be added to by saying that the medul-
lary rays are often close to each other tangentially, and the
rays and ray-like cells of the pericycle often sweep around the
narrow portions of the phloem, as seen in figure 6 of Amoy^pha
fi-uticosa, as though for the special purpose stated above, and
in a way especially fitting them for that function. Some re-
cent research^ has further verified Strasburger's statements
in certain plants.^

One thing to be noted in botanical literature is that work on
sieve tubes and sieve plates is usually done with plants from
the Vitacese, Cucurbitacese, Tiliacese, and a few others, where,
as will be seen by reference to table I, the walls of the phloem
are unusually thick. Presumably this is because here the
things to be seen are more pronounced, and hence more easily
found and studied. Table II shows comparative data collected
from the plants having pits in the lateral walls of the phloem
as observed in this work and from those in which such pits
were not observed. In Equisetum pits are general in the
phloem elements and in the rest of the cortical tissues. In
others this is not true. In the tables mentioned it will be seen
that pits are present in those plants having thicker walls than
those in which pits were not observed, and, as a rule, the sec-
tions were from actively functioning tissue. It is seen that
the phloem walls are as thin in some cases as 0.25 micron, and
in many cases 0,35 micron. In fact, in the phloem in many
instances there was but little evidence of appreciable addition
in thickness to the original vertical wall formed by the cam-
bium or procambium. As a result of this, in drawing with the
high magnification of 1200 diameters, it was not feasible to
use a double line for representing the thickness of the walls,
and the attempt to do so was soon abandoned and choice made
of the measurement and tabulation method as shown by the
tables. It is quite plain that such thin walls could neither
have nor require pits. In the xylem I found pits invariably

7. Hill, A. W., Histology of the Sieve Tubes of Angiosperms. Annals of Botany,
vol. 22, No. 86, pp. 246-290.

8. Sykes, M. G., Anatomy and Histology of Macrocystis pyrifera and Laminaria sac-
charina. Annals of Botany, vol. 22, No. 86, pp. 291-326.

174 KANSAS UNIVERSITY SCIENCE BULLETIN.

present, but with the cell walls always much thicker. Even
here there are no actual perforations.

Plants often adopt the device of obliquely placed end plates,
possibly for the purpose of furnishing greater surface for the
pits. This is noticeably true in many instances, especially in
the Salicacese. As a result it seems necessary to strengthen
these walls, and cross banding with thickenings in the walls
are seen in figures 2, 8 and 9. The best illustrations (figs. 8
and 9) are from a root of Populus deltoides, taken about eight
feet below the surface of the ground. Here there is a much
increased sieve-plate surface, with strengthening bands at
regular intervals, as seen from two directions.

Problem II. — Are sieve tubes always present in the phloem?

As to this problem, my data, many of which were obtained
only after long search, are quite positive. Plants differ greatly
in arrangement, location, number, size, staining qualities, etc.,
of their sieve tubes in the various specimens. However, one
can be reasonably sure of their presence by noting the phloem
in cross section. The companion cells and the sieve tubes ad-
joining form units in outline, which, together with the proto-
plast of the companion cell and the slime body of the sieve
tube, assist in the discovery (fig. 6). Sieve plates are not so
easily found, unless it may be in those plants where sieve
tubes are numerous, or where one happens to get a section
through an unusually favorable place. For still better evi-
dence the slime body is good, but the perforated sieve plate is
best. I was usually not satisfied until all of these requirements
were met, and the drawings were then made.

Perforations were not found in the end plates in Pinus
laricio, Sagittaria, and Aspa)-agus. They were quite numer-
ous, although very small, in Opnntia. In Myriophyllum, a
hydrophyte, they were evident, as in Cuscuta, a parasite. In
this last were found cells of large dimensions, apparently
serving especially as conducting cells. One was 0.1 mm. in
tangential diameter by 1.62 mm. in axial length. We could not
tell how much longer, because of the limits of the section.
Sieve tubes are also present in Tillamlfiia, an epiphite. Here
there is a cylinder of sclerenchyma inclosing the vascular bun-
dles, and probably permitting of but little lateral transfer of
foods.

Data concerning the sieve tubes are contained in tables I-IV.

AGRELIUS: FOOD-CONDUCTION IN PLANTS. 175

It will be seen that the walls are commonly quite thin — those
between them and the companion cells being thinnest. Thick-
ness of their walls ranges from 0.25 micron to 4.17 microns.

Problem III. — The facts observed concerning the third part
of the problem are tabulated in table III. The tendency is
shown to be toward a reduction of the diameter of the vessels
in the fruitstalk and petioles, with an increase in thickness
of the walls.

RESUME.

First. — There are special devices for the better lateral trans-
fer of foods present in the phloem and cortex of the plants
studied. These are (a) pitted walls; {h) the arrangement of
the phloem in narrow wedges with pericycle cells sweeping
around these and connecting with the medullary rays, and
(c) radially elongated medullary-ray cells adapted readily to
conduct the food as stated. It is probable that in many plants
the extreme thinness and large area of the phloem walls per-
mits a considerable lateral movement of materials without
resort to pits, and that no pits are to be found in some (per-
haps many) cases.

Second. — Sieve tubes with perforated sieve plates were dem-
onstrated in all but three of the plants examined, and here I
was satisfied of their presence. They often increase the area
of their sieve plates by placing them obliquely, and these may
require special thickenings to support them. The number of
the sieve tubes in various species varies considerably. They
are present in some plants when probably almost functionless.

Third. — Relatively, the phloem and xylem do not vary
greatly in amount in the various parts of the plants, such as
the stems, petioles, etc., but the main differences in this respect
are in the other tissues.

2-Univ. ScL Bull., Vol. V, No. 10.

176

KANSAS UNIVERSITY SCIENCE BULLETIN.

Table I. Thickness of cell walls in stems (measurements in microns).

6S

2.!!

re 3

Between sieve
tubes and com-
panion cells

Between sieve
tubes and med-
ullary rays

Between tracheal
tissues and xyl.
parenchyma.. . .

Between tracheal
tissues and med-
ulls^ry rays

Pinus laricio Bort

Saggittaria latif olia Willd

0.65
0,35
0.50
0.30
0.30
1.00
0.58
0.35
0.40
0.35
1.08
0.50
0.35
0.40
0.35
0.35
0.30
0.35
0.40
1.28
0.45
0.42
0.35
0.50
0.45
1.65
0.60
0.40
0.65
0.40
0.45
0.40
0.55
0".50
0.75
0.40
0.55
0.50
0.68
1.00
0.50
0.70
2.33
0.55
0.65
0.50
0.50
0.45
1.15
0.60
0.42
0.60
0.55
0.45
0.50
0.60
0.60
0.65
0.50

0.32
0.40
0.25

0.65

2.58
1.50
1.65
1.50
1.30
1.65
1.78
2.79
1.85
2.73
2.73
2.80
3.53
3.83
2.31
2.60
3.86
2.75
3.08
3.72
2.48
3.72
1.58
3.11
2.19
3.65
2.43
3.94
2.94
3.93
2.80
2.70
3.37
4.48
2.64
1.85
2.35
3.08
2.35
2.60
2.50
2.20
3.83
2.48
3.25
1.66
3.10
3.50
4.47
2.85
3.40
3.80
3.02
2.46
5.56
4.44
2.78
2.68
3.25

1.60

Zea mays L

Scirpus validus L

Tillandsia usneoides L

Smilax rotundifolia L

6.60
0.40
0.30
0.35
0.30
0.53
0.35
0.30
0.35
0.25
0.25
0.30
0.30
0.35
0.92
0.40
0.40
0.25
0.40
0.45
1.71
0.40
0.30
0.30
0.25
0.35
0.33
0.35
0.35
0.55
0.35
0.55
0.38
0.55
0.45
0.35
0.50
1.72
0.50
0.45
0.38
0.30
0.25
0.75
0.40
0.35
0.35
0.40
0.25
0.30
0.40
0.40
0.35
0.45

"6'.40'
0.35
1.25
0.50
0.35
0.35
0.40
0.35

"0.35
0.55

"o.si"

0.25
0.55
0.35
3.30
0.55
0.45
0.58
0.40
0.40
0.40
0.50
0.45
0.75
0.50
0.90
0.50
1.00
0.85
0.70
0.50
4.17
1.00
0.80

"OAO
1.30
0.80

"6!45"
0.30
0.55
0.50
0.50

Yucca e:lauca Nutt

Asparagus officinalis L

Salix alba, var. vitelHna (L.) Koch

1 28

J uglans nigra L

1 65

Populus alba L

1 65

Populus deltoides Marsh

1 85

Ulmus Mx f ulva

2.61

Ulmus americana L

1 67

Celtis occiden talis L

1 56

Cannabis sativa L

1.72

Humulus lupulus L

Morus rubra L

1 60

Asimina triloba Dunal

Clematis pitcheri T. & G

3 50

Menispermum canadense L.

Deutzia fortunei Hort

2 78

Spira?a japonica Li.? . . .

Pyrus malus L

1 72

Rubus occidentalis L

1 75

Rubus cuneifolius Pursh

1 85

Prunus americana Marshall

1.92

Gymnocladus dioica (L.) Koch

3.16

Gleditsia triacanthos L

2.46

Melilotus alba Desr

1 85

Amorpha f ruticosa L

1 65

Desmodium canescens (L.) DC

2.08

Zanthoxylum americanum Mill

3.25

Ailanthus glandulosa Desf

3.61

2 20

Rhus toxicodendron L

1.35

Rhus cotinus Nutt

1.50

Celastrus scandens L

1.85

Acer saccharinum L

2 00

Acer negundo L

1.87

Rhamnus lanceolata Pursh

1.80

Psedera quinquefolia (L.) Greene

1.8S

Vitis aestivalis Mx

1.77

Tilia americana L

Abutilon theophrasti Medic

Opuntia rafinesquii Engelm

Myriophyllum heterophyllum Mx

Cornus asperifolia Mx

2.08

Fraxinus viridis Mx.?

2.27

Syringa vulgaris L

2.02

Cuscuta glomerata Choisy

Lycium halimifolium Mill

Tecoma radicans Juss

2.13

Catalpa speciosa Warder

2.15

Sambucus canadensis L

2.3C

Triosteum perfoliatum L

Symphoricarpos orbiculatus Moench

1.92

Brauneria pallida ( Nutt) Britton

Gaillardia grandiflora

Minimum thickness of wall

0.300
2.330

0.250
1.720

0.250
4.170

1.300
5.660

1.280

Maximum thickness of wall

3.610

Average thickness

0.598

0.430

0.715

2.866

2.095

AGRELIUS: FOOD-CONDUCTION IN PLANTS.

177

TABLE II.

Pinus laricio Bort

Yucca g-lauca Nutt

Salix alba, var. vitellina (L.) Koch

Populus alba L

Populus deltoides Marsh, (stem)

Populus deltoides Marsh, (root)

Menispermum canadense L

Rubus occidentalis L

Gymnocladus dioica (L.) Koch

Celastrus scandens L

Psedera quinquefolia (L.) Greene

Vitis aestivalis Mx

Abutilon theophrasti Medic

Symphoricarpos orbiculatus Moench

Average for the above plants with pitted
walls

Average from plants where pits were not
observed

Difference (+) or (— )

Thickness in microns of walls between—

S. t. and

s. t.

0.65
0.58
0.40
1.08
0.50
1.39
0.45
0.45
0.40
0.50
0.70
2.3S
0.65
0.60

0.763
0.549

S. t. and
comp.
cells.

C.40
0.35
0.53
0.35
1.00
0.40
0.45
0.30
0.38
0.50
1.72
0.45
0.40

0.556
0.394

S. t. and
m. ray.

0.65

0.40
1 25
0.50
1.22

+0.214 +0.162

0.35

0.45

0.50

0.50

4.17?

0.80

0.50

0.941
0.633

+0.308

Tracheal
tissues

and
xyl. par.

2 58
1.78
1.85
2.73
2 80
4.88
2.48
2.19
3.94
3.08
2.20
3.83
a. 25
2.78

2.884
2.922

-0.038

Tra-
cheal

tissues

and
m. ray.

1.60

1.28
1.65
1.85
3.24

1.75
3.16
1.85
1.85
1.77

1.92

1.993
2.133

—0.140

CO

<;

H
W
H

«

Q
Z
<!

m

pj
O

I— t

H

W c
US (u

s

:^^

<!

o

^;
o

•«!

S
o
o

H

•^1

Fruitstalks.. . .

1

1

Petioles

;

43 9

Stems

»

to

si

.2 2

Fruitstalks . . .

CO

d

£3
§3

1-H

Petioles

eo

1—1

Stems

co'
CO

o

C CO

.J

Fruitstalks . . .

ei m'

Petioles

CO

CO
CO

CO

Stems.

C-

1-1

g2
1-J

§3

Walls between tr.
and xyl. parench.

Fruitstalks....

\

8

O

f-H

CO

g

CO

Petioles

CO

s

co'

o
o

Stems

CO

CO

3

C4

CO

g

OS

1

Fruitstalks.. . .

I>

1^

Petioles

•<*

00

00

oi

T— 1

Stems

CO
00

CO

00
CO

00

04

CO
CO

d

9

CO
CO

SI

00

Fruitstalks . . .

;

d

CO

d

d d

Petioles

CO

".

d

o

1-1

Stems

CO

U3

CO

d

d

o
d

d

d

c1

OS rt

a'

Fruitstalks....

:

g
d

CO

d

o

CO

d

o
d

Petioles

CO

cm'

r-l

d

Stems.

d

d

eo

CO

d

d

CO

d

Walls between
s. t. and s. t.

Fruitstalks . . .

g
d

d

d

d

Petioles

S SI

i-i d

Stems

S3

1 =-•

CO

CO

d

d

o
d

d

o
d

.2
>

I
■I

>

1

'S

3
u

■

3

cm

00

3
1

1

0

1

.>

'■a
CO

CO

3

u

s

3

E
S
Q

d
Q

n

5)
o
m

B

s

s

3

CI

J

>

n
3
tn

&

5

1
B

3
B

1

3

5

1

AGRELIUS: FOOD-CONDUCTION IN PLANTS.

179

TABLE IV. Resume.
(Measurements in microns.)

Walls
between
sieve
tubes
and
sieve

tubes.

Walls
between
sieve
tubes
and
comp.
cella.

Walls
between
sieve
tubes
and
med.
rays.

Diam-
eter of
sieve
tubes.

Walls
between

tra-

cheals

and

xyl.

parench.

Walls

between

tra-

cheals

and

med.

rays.

Diameter

of
tracheals.

Lengrth

of

sieve

tubes.

Stems, only.
Petioles ' '

0.598«'
1.317*

0.504"
1.390'

0.430''*
1.093'

0.364'
1.000

0.715"

1.470*

0.528'
1.220

12.193°
13.166=

8.302'°
41.640

2.866«'
3.526=

2.884"
4.880

2.095
2.997'

2.229'
3.240

33.633"
30.612''

15.830"
206.55

183.608*

Fruits talks,
only

Roots, only.

182.180

Minimum.. .
Maximum . .

0.300
2.330

0.250
1.720

0.250
4.170(?)

3.500
41.640

1.300
5.560

1.280
3.610

9.950
206.55

97.600
256.780

Average

0.622

0.471

0.746

12.009

2.936

2.166

32.845

183.608

8-Univ. Sci. Bull., Vol. V. No. 10.

THE

KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 11— April, 1910.

(Whole Series. Vol. XV. No. 11.)

CONTENTS:

Histology of Townsendia exscapa and Lesquerella spathulata,

Lilian Biinton.

PUBLISHED BY THE UNIVERSITY
LAWRENCE, KAN.

Enteieil at the post-office in Lawrence as second-class matter.

THE KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. II] APRIL, 1910.

Whole Skkies
Vol. XV, No. 11

HISTOLOGY OF TOWNSENDIA EXSCAPA AND
LESQUERELLA SPATHULATA.

bY LILIAN BUNTON.
Plates XXXVIII to XLV.

''pWO excellent representatives of xerophytic vegetation,
^ Toumsendia exscapa and LesqueveUa spatludata, were col-
lected in Gove county, Kansas, at an elevation of 2700 feet.
These plants are not common throughout this district and are
limited to certain localities. Toivnsimdia was found growing in
draws in the immediate vicinity of the town of Gove and also
on the flat, open prairie around "Castle Rock," about twenty-
five miles southeast of Gove, and in the vicinity of a group of
rocks known as the "Pyramids," thirty miles southwest.

A general idea of the appearance of the vegetation of this
district may be derived from figure 1. My examples of Toivn-
sendia, therefore, were growing on the flat, open prairie as well
as in the draws, occurring not in clumps or patches, but as
isolated individuals within three or four feet of one another.
The plants on the open prairie show a more luxuriant growth
than those in the draws.

The specimens of LesqueveUa were collected about twelve
miles southwest of the town of Gove on a rocky, clay bluff".
This plant, unlike Toiimsendia, was found growing not as iso-
lated individuals but in rather large colonies.

Of all the ecological factors influencing the structure, habit
and distribution of plants the most potent is the water supply.
In Gove county the total precipitation for the year 1904 was
18.10 inches ; the greatest monthly precipitation was 4.13 inches,
in May ; the least monthly precipitation was 0 inches in Febru-
ary and November, The total snowfall for the year was 5.5

(183)

184

KANSAS UNIVERSITY SCIENCE BULLETIN.

inches and the number of rainy days 39*. In the following
table is given the monthly and annual precipitation in Gove
county for the year 1904, with departures from the normal :

January.

Precipitation 0.01

Departure — 0.57

February.

Precipitation 0.00

Departure — 0 . 72

March.

Precipitation 1 . 00

Departure +0.32

April.

Precipitation 1 .79

Departure — 0 . 27

May.

Precipitation 4.13

Departure +0.89

June.

Precipitation 2 . 62

Departure — 1 . 25

July.

Precipitation 2 . 18

Departure — 1 . 47

August.

Precipitation 2.57

Departure +1.11

Septembei'.

Precipitation 1.26

Departure — 1 . 08

October.

Precipitation 2 . 00

Departure +0.98

November.

Precipitation 0 . 00

Departure — 0 . 51

December.

Precipitation 0. 55

Departure • +0.03

Annual.

Precipitation 18.10

Departure —2.27

From comparisons with other places in Kansas where
weather records have been kept, it is seen that the amount of
precipitation of this habitat is remarkably small; in fact, be-
ing the least, excepting that of Oberlin, Lakin, and Dodge City;
^A'here the annual precipitation was 15.72 inches.

One may acquire a general idea of the temperature of this
locality from the following tables :

Monthly and annual mean temperature for year 1904, with departure

from normal, Fahrenheit.

January.

Temperature 29 . 0

Departure +0.9

February.

Temperature 30 . 6

Departure +1.9

March.

Temperature 43 . 8

Departure +6.4

April.

Temperature 48 . 6

Departure 0.0

May.

Temperature 57.8

Departure — 4 . 6

June.

Temperature 67.4

I'eparture — 4.9

July.

Temperature 74.8

Departure — 3 . 8

August.

Temperature 72 . 6

Departure — 4 . 0

September.

Temperature 68 . 3

Departure — 0 . 1

October.

Temperatuie 55.3

Departure +1.5

November.

Temperature . 44.6

Departure — 0 . 9

December.

Temperatuie 30.2

Departure — 2.4

Annual.

Temperature 51.9

Departure — 0 . 5

BUNTON: XEROPHYTIC VEGETATION.

185

Monthly maximum temperatures for the year 1904, with dates.

January 18 65°

February 9 72°

March 2 82°

April 10 79°

May 24 93°

June 23 99°

July 29 102°

August 24 95°

September 9 103°

October 3 87°

November 18 73°

December 31 68°

Monthly minimum temperatures for year 1904, with dates.

January 25 — 2°

February 19 3°

March 3 6°

April 17 25°

May 14 34°

June 11 50°

July 26 54°

August 22 50°

September 14 41°

October 25 25°

November 11 11°

December 26 4°

Another ecological factor which has much to do in shaping
plants is the wind. On the prairies the wind has full sweep ; it
blows most of the time at a rapid rate and tends to dry up what-
ever vegetation there may be in its path. No definite statement
can be made as to the velocity of the wind of this special local-
ity. In the following table, however, are shown some of the
results of observations made by Stevenson, of Edinburgh, upon
the increase in velocity of the wind with the height above the
ground, as given in Schimper's Plant Geography :

Height of instrument
al>oveprroun(l, in feet.

V2 ....

Velocity of wind
in miles per hour.

6.83

2% 8.73

41/3 9.77

91/2 10.45

14 10.54

25 11.54

50 12.21

Height of instrument
aboveground, in feet.

V2
31/2

9
14
25
51

Velocity of wind
in miles per hour.

.... 9.8

12.4

. ... 13.8

14.3

15.0

16.3

Height of instrument
aboveground, in feet.

IVs ....

31/2 ....

9

14

25

51

Velocity of wind
in miles per hour.

.... 22.2

.... 25.6

.... 31.9

33.7

37.1

42 . 7

From the above tabular statement it is quite clear that plants
that are only slightly raised aboveground experience the effects
of the wind much less than tall plants, and especially trees.

186 KANSAS UNIVERSITY SCIENCE BULLETIN.

The action of the wind on plants is partly direct by tensile
stresses and partly indirect by increasing transpiration ; both
these actions are the more energetic the taller the plants or the
higher their location.

Where my Townse^idias occurred the soil is a loose, cal-
careous deposit, allowing the water quickly to percolate
through it.

When the character of its habitat is considered, it would be
expected that the anatomy of Townsendia must be that of a
xerophyte.

TOWNSENDIA EXSCAPA.

ToivTisendia exscapa is perennial from a woody taproot
which averages ten centimeters long and three to four milli-
meters in diameter at the thickest portion. The stem grows
from one to four inches high. The fact that this plant is low
helps it to endure the strong winds prevailing in its locality.

The character of the stem varies in different plants. In
some, while young, it is quite simple, bearing only a few heads
of flowers ; in others it is several-branched, bearing six or more
heads. As a rule, however, it gradually becomes more and
more branched with age until the whole plant may attain a
breadth of ten to twelve inches, bearing about 100 heads of
flowers. The leaves are all clustered at the base, narrowly
linear or slightly spatulate, and one to two inches long. The
flowers are white and are borne in sessile heads one to one and
one-half inches broad. In figure 2 is shown a photograph of
Townsendia. When the flowers of the larger plants have gone
to seed the plants have the appearance seen in figure 3.

In the discussion of the histology of Townsendia the stem
will be first treated, then the root, and last the leaf, and the
diff"erent tissue systems under each will be examined.

The stem.

Tegiimentary Tissues. — The protection of the mature stem
is provided for by a cutinized epidermis and cork tissue. All
the walls of the epidermis are cutinized. In figure 4 is shown
a small portion of the cutinized epidermis. In the following
table is given the average thickness of the walls of the epi-
dermal cells and the average length and width of the cavity of

BUNTON: TOWNSENDIA EXSCAPA.

187

the epidermal cells of Townsemlia and a few mesophytes and
xerophytes :

Outer
wall.

Inner
wall.

Radial
wall.

Vertical
length.

Radial
width.

Eupatorium
altissimum.

0.005 mm.

0.001mm.

0.001mm.

0.022 mm.

0.015 mm.

Mesophytes.

Salvia
azurea.

0.006 mm.
Cutinized.
portion,
0.005 mm.
Cuticle,
0.001 mm.

0.006 mm.

0.003 mm.

0.02 mm.

0.01mm.

Hydrophtes.

Sagittaria.

0.003 mm.

0.001mm,

0.001mm.

0.02 mm.

0.01 mm.

Nelumbo-
lutia.

0.006 mm.

0.003 mm

0.003 mm.

0.04 mm.

0.02 mm.

TOWNSENDIA.

0.05 mm.

0.02 mm.

0.018 mm.

0.09 mm.

0.068 mm.

In the following table is given the average thickness of the
layer of cork, the average thickness of the walls of the cork
cells, and the average length and width of the cell cavity in
Townse7idia:

Thickness of cork 4-7 cells.

Average thickness of wall 0.004 mm.

Average length of cavity 0.14 mm.

Average width of cavity 0.087 mm.

In figure 5 is shown a small portion of cork tissue in cross
section.

When compared with the mesophytes and hydrophytes
above mentioned, it is seen that Toivnsendia has a well devel-
oped protective system, one of the primary requisites of a
plant growing in a dry, windy habitat.

The Streyigthenmg Tissues. — The strengthening tissues
found in ToumseyuHa consist exclusively of collenchyma and
short sclerenchyma cells. From figures 6 and 7, collenchyma
cells as seen in cross and longitudinal sections, it may be seen
that the cells of the collenchyma tissue have remarkably thick
walls. The walls of the short sclerenchyma cells also are much
thickened, as shown in figures 8 and 9. No true bast fibers are
found ; the tissue that in cross section appeared to be bast
when examined longitudinally proved not to be bast, but a
tissue of short sclerenchyma cells. Wood fibers are also

188 KANSAS UNIVERSITY SCIENCE BULLETIN.

wholly lacking. This deficiency, however, is in a measure com-
pensated by the modification of other tissues, the walls of the
wood parenchyma, cortex, pith and medullary-ray cells being
remarkably thickened. The walls of the cortex and pith cells
are, on the average, 0.03 mm. thick. In figures 10 and 11 are
shown a few cortex cells as seen in cross and longitudinal
sections, respectively. In figures 12 and 13 are shown a few
pith cells as seen in cross and longitudinal sections, respec-
tively. As shown in figure 14, the tr icheids have thick, ligni-
fied walls and constitute the smaller portion of the xylem area ;
these, together with the wood parenchyma cells, tend to
strengthen the plant. A longitudinal section was mounted in
a drop of concentrated solution of chromic acid ; in a few min-
utes everything had disappeared but the tl^acheids. The
chromic acid dissolved the wood parenchyma cells, leaving the
tracheids isolated as shown in figure 15. The average length
of a tracheid is 0.243 mm. ; the average width, 0.016. mm.

Tracheal Tissues. — In the stem the conduction of water is
provided for by tracheids varying in length, and in longi-
tudinal section having the appearance of typical tracheal
tubes. When, however, a longitudinal section was macerated
under a cover glass in chromic acid, the tracheids were isolated
from the other tissues and stood out prominently as spiral and
scalariform tracheids. By computing the number of tracheids
in one xylem wedge and multiplying by the number of such
wedges, where the cross section of the stem is 19.635 sq. mm.
in area, there were found to be 1313 tracheids. The average
diameter of these stem tracheids is 0.016 mm. The total
water-conducting area of the stem, cross section, is 0.266
sq. mm., which is 1.3 per cent of the whole stem. Of the xylem
portion of the stem, 6.6 per cent is given up to tracheids.

When compared with the water-conducting tissue of several
mesophytic herbaceous plants, that of Toivnsendia seems quite
small. In Saponaria officinalis 22.7 per cent of the whole stem
is devoted to the conduction of water; of the xylem portion
29 per cent, the average diameter of a tracheal tube being
0.026 mm. Thus it is seen that 17 times as much of the stem
of S. officinalis as in Toivnsendia is devoted to the conduction
of water.

In Verbena stricta there was devoted to the conduction of
water 9 per cent of the stem cross section, where the area was

BUNTON: TOWNSENDIA EXSCAPA. 189

found to be 3.801 sq, mm. Of the xylem portion, 62 per cent
was devoted to the conduction of water. In Verbena, then,
there was found devoted to the conduction of water seven times
as much of the stem as in Toivnsendia. In Mentzelia oligo-
sperma there was devoted to the conduction of water 9 per
cent of the stem cross section, where the area was 9.07 sq. mm.
Of the xylem poition, 33 per cent was devoted to the conduc-
tion of water. In Mentzelia also there was found to be de-
voted to the conduction of water nearly seven times as much of
the stem as in Townse^idia. In Ambrosia artemisifolia there
was found devoted to the conduction of water 21 per cent
of the stem cross section, where the area was 13.86 sq. mm.
Of the xylem portion 52 per cent was devoted to the conduction
of water. In Ambrosia, then, there was devoted to the conduc-
tion of water about sixteen times as much of the stem as
in Toivnsendia. In Abutilon avicennx there was devoted to the
conduction of water 2 per cent of the stem cross section, where
the area was 38.48 sq. mm. Of the xylem portion, 6 per cent
was devoted to the conduction of water. In Abutilon, accord-
ingly, there was devoted to the conduction of water approxi-
mately twice as much of the stem as in Toivyisetulia.

When compared with the water-conducting tissues of desert
plants, which have have been discussed by W. A. Cannon^ in
his paper on the ''Water-conducting Systems of Desert Plants,"
Toumsendia has a relatively large water-conducting area.

In the following table is given the number of tracheal ele-
ments in 1 sq. mm. of stem, cross section, of Townsendia, and
of several mesophytes and xerophytes; the latter cited from
Cannon :

Townsendia 66

Mesophytes :

AbutUon avicennse 33

VerbeiM stricta 734

Mentzelia oligosperma 77

Ambrosia artemisifolia 124

Xerophytes :

Celtis pallida 1

Covillea tridentia 2

Fouquieria splendens 7

•

Tissues for Food Conduction and Storage. — The medullary
ray is composed of two kinds of cells, some with rather thick

1. Bot. Gazette, June, 1905, pp. 396-408.
•J-Univ. Sci. BulK. Vol. V. No. 11.

190 KANSAS UNIVERSITY SCIENCE BULLETIN.

cellulose walls which measure 0.001 mm. in thickness. The
thick-walled medullary-ray cells constitute approximately one-
third of the whole area of a medullary ray. An explanation
of there being both thick- and thin-walled medullary-ray cells
may possibly be that the thick-walled cells have sacrificed
their function of conduction and have assumed that of
strengthening. The average vertical length of a medullary-
ray cell is 0.01 mm. ; the average width 0.008 mm. In figure 16
are shown a few medullary cells as seen in cross section ; in
figure 17 as seen in longitudinal section.

Very little special provision for the passage of food longi-
tudinally seems to have been made by Toivnsendia. The ordi-
nary collateral vascular bundles are found. A comparatively
small portion of the vascular bundle, however, is devoted to
phloem — about 12 per cent; in Verbena stricta 25 per cent of
the vascular bundle is devoted to phloem. In Mentzelia oligo-
speTTiia there is the same proportion of the vascular bundle de-
voted to phloem as in Toivnsendia — 12 per cent. Neither sieve
tubes nor sieve parenchyma cells are found in Townsendia.
The whole work of conduction of food longitudinally is ap-
parently assumed by the cambiform cells and the undivided
mother cells of the sieve tubes. Presumably the sieve tubes,
when present in a plant, would carry the food more rapidly.
The most elaborated sieve tubes are always found in plants of
considerable length, especially in trailing, clambering and
climbing plants. Toivnsendia, however, is so low that it seems
that the food can be conducted longitudinally rapidly enough
and in sufficient quantities by the cambiform and undivided
ipother cells of the sieve tubes without the aid of sieve tubes.

In figure 18 are shown phloem cells as seen in cross section ;
in figure 19, as seen in longitudinal section : a, cambiform cell ;
h, undivided mother cell of the sieve tubes. The average
length of these cells is 0.18 mm; the average width, 0.01 mm.
The average thickness of their cell wall is 0.0006 mm.

In Toivnsendia, the xylem parenchyma cells and the medul-
lary ray, cortex and pith cells afford sufficient means for the
storage of food. Upon examination of sections of the stem of
the plant collected at the time of flowering and fruiting, there
appeared stored up in the tissues of the stem no food what-
ever, thus showing that the food must at this time be used

BUNTON: TOWNSENDIA EXSCAPA. 191

by Totvnsendia as fast as it is made. No statement, however,
can be made of the storage of food in the plant later in the
season.

The Root.

Tracheal Tissues. — As in the stem, the conduction of water
in the root is provided for by spiral and scalariform tracheids.
By computing- the number of tracheids in a certain portion of
a cross section, and multiplying by the number of such por-
tions, there were found to be 1271 tracheids in the cross sec-
tion of the root. In figure 20 may be seen how these tracheids
are distributed throughout the section. The average diame-
ter of these tracheids is 0.018 mm. The total water-conduct-
ing area in a root, where the cross section area is 7.496 sq.
mm., is 0.325 sq. mm., or 4.2 per cent of the whole root area.

In structure, the other tissues of the root do not differ es-
sentially from those of the stem; hence the discussion of them
as found in the stem will suffice for the root.

The Leaf.

The Tegumentary Tissues. — The outer wall of the epidermis
of the leaf is not wholly cutinized ; the inner portion is of cel-
lulose and is 0.0108 mm. thick, and the outer portion of the wall
is cutinized and is 0.0072 mm. thick. The inner tangential
wall is of cellulose and is 0.0036 mm. thick. The radial walls
are likewise of cellulose and 0.0036 mm. thick. The average
tangential length of the cells of the epidermis is 0.0396 mm. ;
the average radial length, 0.025 mm. No difference was found
between the upper and lower epidermis, hence the figures given
above will suffice for both. In figure 24 is shown a portion of
the cutinized epidermis as seen in cross section.

Tracheal Tissues. — After a leaf boiled in alcohol had re-
mained in a mixture of chloral hydrate and hydrogen peroxide
for several days, it had become so well bleached that its vena-
tion could be seen easily with the naked eye. There appeared
three prominent longitudinal veins, one central and two lat-
eral, from which extended other veins branching in every
direction and terminating in water-storage tracheids, as
shown in figure 21o. In the cross section of the midrib
thirough the center of the leaf were found ninety-five tracheids,
the average diameter of which was 0.0075 mm., making the

192 KANSAS UNIVERSITY SCIENCE BULLETIN.

total water-conducting area there 0.00004 sq. mm. The cross-
section area of the midrib was 0.01207 sq. mm., 32 per cent of
which was devoted to conduction of water. In figure 21b is
shown the midrib as seen in cross section. In the cross sec-
tion of each of the lateral veins were found twenty-eight
tracheids, the average diameter of which was 0.0059 mm.,
making the total water-conducting area in each vein 0.00073
sq. mm. The total cross-section area of each vein was 0.003
sq. mm., 24 per cent of which was devoted to the conduction of
water. The average distance between the ultimate branches
of the veins — a and b, figure 22 — is 0.031 mm.

In the leaf, the phloem consists only of cambiform cells and
undivided mother cells of the sieve tubes ; the xylem is made up
of wood parenchyma cells and spiral and scalariform tracheids.
The average diameter of the cavity of the tracheids is 0.006
mm. The walls of the parenchyma cells, however, remain
quite thin. In figure 23 are shown the phloem cells as seen in
longitudinal section of the midrib : o, cambiform cell ; b, undi-
vided mother cell of the sieve tubes. The average length of
these cells is 0.14 mm., the average width 0.01 mm., the aver-
age thickness of the cell wall 0.0006 mm.

The Photosiinthetic Tismie^. — On both sides of the leaf are
found palisade cells ; the sportgy parenchyma cells are not very
prominent and toM^ards the center of the leaf, in some places
in the leaf not really being differentiated from the palisade
cells. The explanation of the palisade cells on both sides of
Townscndia may be that out on the open prairie the reflected
light from below is sufficiently intense to stimulate the forma-
tion of palisade tissue on the under side of the leaf.

On each side of the leaf there are from two to three layers
of palisade cells, the average length of which is 0.14 mm., the
average width 0.05 mm. The average number of chloroplasts
in a single cell is twenty-nine and the average diameter of a
chloroplast is 0.0008 mm. In figure 25 are shown the palisade
cells as they appear in surface view; in figure 26 spongy pa-
renchyma cells as they appear in surface view.

In the following table is shown the number of chloroplast^

BUNTON: TOWNSENDIA EXSCAPA. 193

found in a single mesophyll cell of a number of plants, as given
in Haberlant's "Pflanzen-Anatomie" :

Spongy
Palisade cells, parench. cells.

Fragaria elatior 86 14

Pulmonaria officinalis 85 15

Ricinus communis 82 18

Brassica rapa 80 20

Galeopsis tetrahit 79 21

Ft'opaeolum majus 77 23

Helianthus anmms 73 27

Phaseolus vinltiflorus 69 31

Bellis perennis 67 33

When compared with the plants above mentioned, it is seen
that in the leaf of Toivnsendia the chloroplastic surface is
relatively small. On both sides of the leaf the number of
chloroplasts in a single mesophyll cell is approximately
twenty-nine, thus giving to both sides of the leaf the same
chlorophyll-producing volume. The small number of chloro-
plasts found in the mesophyll cells of the upper side allows
the sunlight to penetrate to the lower side of the leaf, thus
giving to the cells there the power to carry on photosynthesis
more effectively.

The Water-storage Tissues. — In the leaf of Townsendia
there is an admirable arrangement for all the mesophyll cells
to have immediate access to water. Throughout the full
length of the leaf on both sides are found large cells which
are used primarily for the storage of water. These water-
storage cells, however, seem to assume other functions; they
evidently take part in the conduction of water and aid to a
slight degree in photosynthesis. They have very thin walls,
which are lined by a few chloroplasts much smaller than those
found in the other mesophyll cells. Bordering on these water-
storage cells are mesophyll cells which draw upon them for
water when there is need.

In order to secure the greatest access to water the palisade
cells occur in various relations : bordering on veins, on water-
storage cells, on spongy parenchyma cells. In the following
table is shown the relative frequency of palisade cells bor-
dering respectively on water-storage cells, veins and spongy
parenchyma cells :

Boi'dering on water-storage cells 222

Bordering on veins 81

Bordering on spongy parenchyma cells 42

194 KANSAS UNIVERSITY SCIENCE BULLETIN.

From the above figures it is seen that the palisade cells occur
most frequently bordering on the water-storage cells, there be-
ing nearly three times as many bordering on the water-storage
cells as on the veins and twice as many bordering on the veins
as on the spongy parenchyma cells. Figure 27 is a diagram-
matic drawing, showing the distribution of water-storage cells,
b, as seen in a cross section of a leaf. In figure 28 are shown
the palisade cells, B, bordering on water-storage cells, A. In
figure 29 are shown the palisade cells. A, bordering on the
veins, C. Some places in the leaf the palisade cells, D, have
still greater access to water, bordering as they do on both
water-storage cells. A, and storage tracheids, C. This relation
generally occurs at the ends of the leaf. In figure 30 are
shown the palisade cells as found in this relation. Sometimes
the palisade cells are found bordering on water-storage cells,
A, which in turn border on veins, C, as shown in figure 31.
Such an arrangement as this is an excellent one; for as fast
as the water-storage cells are deprived of the supply of water
on hand they can immediately draw on the veins and thereby
continue to supply the palisade cells with water as it is needed.
In figure 32 are shown palisade cells, A, bordering on spongy
parenchyma cells, B.

Another means for storage of water is the water-storage
tracheids found at the termination of the veins. Palisade
cells bordering on these tracheids can obtain water when there
is need. In figure 33 are shown these tracheids as they ap-
pear at the ultimate ends of the veins ; in figure 34 the water-
storage tracheids as they appear in other places in the leaf.

The Aerating Spaces. — In Townsendia there were found
devoted to aeration only 20 sq. mm. of the surface of a leaf
which measured 100 sq. mm. ; or, in other words, the volume
of the intercellular spaces and the volume of the whole leaf,
roughly estimated, stand in the relation to ' each other as
8:1000. When compared with Pistia texensis, an hydrophyte,
whose aerating volume and the volume of the whole leaf
stand in the relation to each other as 760 : 1000, the amount
of volume devoted to aeration in Townsendia seems quite
small.

There were found on either side of the leaf an approximate
average of 344 stomata to 1 sq. mm., the average number
usually found in plants. There was no difference in the char-

BUNTON: TOWNSENDIA EXSCAPA. 195

acter of the stomata on either side of the leaf. It seems rather
surprising to find in the leaf of a xerophyte the usual number
of stomata found in plants and to find them at the surface of
the leaf instead of being sunken beneath the epidermis; for,
as a rule, in xerophytic plants there are fewer stomata and
they are often sunken beneath the epidermis.

In figure 35 are shown the stomata as they appear in the
surface view of the leaf of Toivnsendia. In figure 36 is shown
a stoma as it appears in cross section of the leaf.

SUMMARY.

1. Size and Habit of Toivnsendia. — Townsendia is a low
plant, growing from one to four inches high; it grows on the
flat, open prairie as well as in the draws, and occurs not in
clumps but as isolated individuals. The stem is at first rela-
tively simple, but, as a rule, it becomes more and more
branched with age until the whole plant may attain a breadth
of ten to twelve inches.

2. Tegumentary Tissues. — In the mature stem of Town-
sendia the cutinized epidermis, whose outer walls measure
0.05 mm., and the cork tissue several layers thick, constitute
the tegumentary tissues.

In the leaf of Toivnsendia there is found an epidermis
whose outer wall measures 0.017 mm. in thickness. The outer
portion of the wall is cutinized and measures 0.0072 mm. in
thickness ; the inner portion is of cellulose and measures
0.0108 mm. in thickness.

3. Strengthening Tissues. — In Toivnsendia the strengthen-
ing tissues of the stem consist of collenchyma and short
sclerenchyma cells. True bast fibers and wood fibers are lack-
ing. The remarkably thick cell walls, not only of the strength-
ening tissues but of nearly all the tissues, make the stem of
Toivnsendia strong.

4. Tracheal Tissues. — The area devoted to conduction of
water in Townsendia is comparatively small, there being only
1.3 per cent of the stem devoted to that function. Growing as
it does in a location where the water supply is poor, Toivn-
sendia would not require very extensive water-conducting
tissues.

5. Food-conducting Tissues. — Longitudinally there seems
to have been made very little provision for the conduction of

196 KANSAS UNIVERSITY SCIENCE BULLETIN.

food in the stem. The phloem portion is very little differen-
tiated; it consists only of cambiform cells and undivided
mother cells of the sieve tubes, there being found neither sieve
tubes nor sieve parenchyma cells.

6. Storage Tissues. — Since Toivnsendia grows in a xero-
phytic habitat it is not surprising to find in it some provision
for the storage of water. In the stem, the cortex, medullary
ray and pith cells are well adapted to the storing of water.
In the leaf, however, special provision is made: throughout
the whole length of the leaf are found large water-storage
cells which not only reduce transpiration but also rapidly fill
whenever the water supply is increased, and yield their con-
tents to the assimilating cells as the supply of water there is
reduced.

7. Aerating Spaces. — As would be expected to be found in
a xerophyte, the amount of space devoted to aeration is com-
paratively small. Only one one hundred and tw^enty-fifth of
the volume of the leaf of Toivnsendia is devoted to aeration ;
While in Pistia about two-thirds of the volume of the leaf is
thus employed.

LESQUERELLA SPATULATA.

Lesquerella spatulata is perennial by a much branched
caudex which varies from one to two inches in length. It has
a taproot which measures from one foot to a foot and a half in
length and one-fifth of an inch in diameter at its thickest por-
tion. The leaves are separated by short internodes and ap-
pear tufted. They measure from one-half to one inch in
length and one-fifth to one-seventh of an inch in width. They
are entire and vary in form from narrowly elliptic to ovate-
lanceolate, oblanceolate and spatulate, and they, as well as the
pedicels, are beset with stellate hairs. The flowers are yellow
and are borne on pubescent scapes which measure from one-
half inch to six inches in length.

The Stem.

The Tegumentary Tissues. — The protection of the mature
stem is provided for by a cutinized epidermis and cork. There
is, in addition to these tissues, in older portions of the stem, a
borke. All of the walls of the epidermis are cutinized. Figure
37 represents a small portion of the epidermis as seen in cross

BUNTON: LESQUERELLA SPATHULATA. 197

section ; figure 38a, a small portion of the epidermis as seen in

longitudinal section.

In the following table is given the average thickness of the

cell walls and the average length and width of the epidermal

cells :

Outer tangential wall 0 . 0U49 mm.

Inner tangential wall 0 . 0049

Radial wall 0.003

Vertical diameter of cavity 0.075

Radial diameter of cavity 0.026

Tangential diameter of cavity 0.04

In the following table is given the average radial breadth of
the cork zone, the average thickness of the walls of the cork
cells, and the vertical, radial, and tangential dimensions of the
cell cavity :

Average breadth cork zone 15-20 cells.

Average thickness of wall 0 . 001 mm.

Vertical diameter of cavity 0.025

Radial diameter of cavity 0 . 026

Tangential diameter of cavity 0.036

Figure 386 represents a few cork cells as seen in cross sec-
tion.

In addition to this large zone of cork there are found in the
borke rather large stone cells, which, as shown in figure 39,
have relatively thick and pitted walls.

The tegumentary tissues are remarkably well developed for
such a low, herbaceus plant as Lesquerella. When, however,
its xerophytic habitat is taken into consideration, the necessity
for this provision is readily seen.

The Strengthening Tissues. — The strengthening tissues in
the stem consist of collenchyma, bast fibers, stone cells and
wood fibers.

The radial breadth of the collenchyma zone is 0.093 mm.
The average thickness of the walls of the collenchyma cells is
0.006 mm. ; the average tangential breadth of the cell cavity is
0.036 mm. ; the average radial thickness is 0.01 mm. In figure
40 are shown a few collenchyma cells as seen in cross section ;
in figure 41 as seen in longitudinal section.

The sclerenchyma tissue is made up of cells which vary from
stone cells to short bast fibers. As shown in figure 42, the stone
cells found here are smaller than those found in the borke.
Figure 43 represents the stone cells as they appear in longi-
tudinal section.

3-Univ. Sci. Bull. Vol. V. No. 11.

198 KANSAS UNIVERSITY SCIENCE BULLETIN.

The bast fibers are 0.11 mm. long and 0.016 mm. wide, the
average thickness of the cell wall being 0.006 mm. As may be
seen in figure 44, the ends of the fibers overlap but little. In
figure 45 are shown a few bast fibers as seen in cross section.

The most striking characteristic of the strengthening tissues
of the stem is the presence of large zones of thick-walled wood
fibers, which constitute 26 per cent of the area of the stem and
78 per cent of the area of the xylem. These wood fibers, as
shown in figure 46, occur among spiral tracheal tissues. They
average 0.11 mm. in length and 0.01 mm. in width. The walls
are remarkably thick — 0.006 mm. ; the cell cavity is exceed-
ingly small, in some places being almost entirely obliterated.
Figure 47 represents a wood fiber as seen in longitudinal sec-
tion.

The strengthening system in the stem of Lesquerella is more
extensively developed than that in the stem of Toivnsendia, the
only strengthening tissues found in the stem of Toivnsendia
being collenchyma and short sclerenchyma cells, bast fibers and
wood fibers being absent. This deficiency in Townsendia, how-
ever, is compensated by remarkably thick cell walls, not only
of the strengthening tissues but of nearly all the tissues. The
strengthening tissues in the stem of Lesquerella are so well de-
veloped that there is no need of the other tissues being modi-
fied, as was the case in the stem of Toivnsendia.

The Tracheal Tissues. — The tracheal system in the stem of
Lesquerella consists of spiral and scalariform tracheal tubes
and spiral and scalariform tracheids alone. There was found
devoted to the conduction of water 2 per cent of the stem cross
section, where the area was 7 sq. mm. ; 4 per cent of the xylem,
where the cross section area was 3.46 sq. mm., was devoted to
this purpose.

In the following table is given the average thickness of the
cell wall, and the average tangential and radial dimensions of
the cell cavities of the various tracheal elements found in the
stem of Lesquerella :

Average thickness Radial diametev TanRential di-
of cell wall. of cavities. ameter of cavities.

Largest tracheal tubes 0.002 mm. 0.016 mm

Smallest tracheal tubes 0.003 0.002

Tracheids 0 . 002 0.016 0 . 04 mm.

As shown in figure 46 — a small portion of the xylem as seen
in cross section — the tracheal elements occur among alter-
nating, definite zones of wood fibers, A, and wood parenchyma

BUNTON: LESQUERELLA SPATHULATA. 199

cells, B, the largest ones being among the wood fibers and the
smallest among wood parenchyma cells. It is generally con-
ceded that spiral vessels are formed by the procambium ex-
clusively, and wood fibers by the cambium. As the tracheal
elements found interspersed among the wood fibers are spiral
and scalariform, the activity here must be due either to the
cambium or procambium. From the fact that these spiral
vessels are found somewhat remote from the pith, there fol-
lows a probable conclusion that the cambium, contrary to the
general rule, has formed spiral vessels.

The tracheal elements found in the stem at the nodes were
different from those in the internodes. At the nodes the
tracheal tissues consist exclusively of short and peculiarly
shaped tracheids. In figure 48, A, are shown the scalariform
type of tracheids; B, the spiral type of tracheids as seen in
longitudinal section.

In the internodes the tracheal elements consist exclusively of
spiral and pitted tracheal tubes, with the exception of a
few tracheids interspersed here and there. Figure 49, A, illus-
trates the spiral tracheal tubes found among the wood fibers,
as seen in longitudinal section; figure 49, B, spiral tracheal
tube found among the wood parenchyma cells ; figure 49, C,
pitted tracheal tubes as seen in longitudinal section.

Tissues for the Conduction and Storage of Food. — The med-
ullary rays found in the stem of LcsqnereUa present an inter-
esting characteristic. Throughout the entire length of the
medullary ray are found leaf traces containing spiral tracheal
tubes as shown in figure 51, a. These tracheal tubes are much
smaller than those found in the xylem area of the stem, the
average diameter of the cavity being 0.007 mm. and the aver-
age thickness of the cell wall 0.002 mm. Figure 50, a repre-
sents a leaf trace. The internodes are so short that it is im-
possible to cut a section without having leaf traces traversing
the medullary rays. These leaf traces run horizontally into the
primary xylem, and when the cambium begins its activity it
builds up the medullary rays around them. The average
width of a medullary ray is 0.06 mm., and the average vertical
length of a medullary-ray cell is 0.03 mm., and its averags
width is 0.01 mm. Figure 51, b represents a small portion of
the medullary ray as seen in cross section.

In the stem, the ordinary collateral vascular bundles are

200 KANSAS UNIVERSITY SCIENCE BULLETIN.

found ; 20 per cent of the vascular bundle is devoted to phloem,
as in Toivnsendia. Neither seive tubes nor seive parenchyma
cells are found. The whole work of conduction is assumed by
the undivided mother cells of the sieve tubes. Lesquerclla is
a very low plant and it seems that the food can be conducted
longitudinally fast enough and in sufficient quantities without
the aid of the other phloem elements. In figure 51, c, are
shown a few phloem cells as seen in cross section ; in figure 52,
as seen in longitudinal section ; at the angles of these cells
occur small intercellular spaces. A,

In the stem of LesquereUa the storage of food is provided for
by means of the thin-walled parenchyma of the cortex and the
xylem parenchyma and the medullary rays. Figure 53 repre-
sents a few cells of the thin-walled parenchyma of the cortex
as seen in cross section. Figure 54 represents a few cells of the
xylem parenchyma as seen in longitudinal section. In all of
these tissues starch and proteid were found in small quantities,
but neither sugar nor oil.

Figure 50 is a diagrammatic drawing of the cross section of
the stem traced as seen when projected upon a screen. From
this may be derived a general idea of the various tissues of the
stem, their relation to one another, and the proportion of the
stem devoted to each: A, borke; />, collenchyma; C, scleren-
chyma; D, phloem; E, xylem; 1, tracheal tissues; 2, wood
fibers; 3, xylem parenchyma; F, medullary ray; a, leaf-trace.

The Root.

The Tegumeyitary Tissues. — The tegumentary tissues of the
mature root consist of a cutinized epidermis and cork tissue
which are similar to those tissues found in the stem, both in
structure and proportion.

The Strengtheni7ig Tissues. — The strengthening tissues
found in the root of LesquereUa consist of collenchyma, stone
cells and wood fibers. These tissues show no difl'erence what-
ever from those found in the stem.

The Tracheal Tissues. — The tracheal tissues in the root con-
sist entirely of scalariform and spiral tracheal tubes. In a
root where the cross section area was 18.75 sq. mm., there was
found devoted to the conduction of water 4 per cent. Here the
cross-section area of the xylem is 6.7 per cent of the entire
root.

In the following table is given the average diameter and the

BUNTON: LESQUERELLA SPATHULATA. 201

average thickness of the cell walls of the tracheal elements in
the root:

Average diameter of the larger tracheal tubes 0.03 mm.

Average diameter of the smaller tracheal tubes 0.007

Average thickness of cell wall 0 . 005

As in the stem the tracheal elements of the root are inter-
spersed among definite, alternating layers of wood fibers and
wood parenchj'ma cells. Figure 55 represents a few wood
fibers and tracheal tubes as found in the outermost layer of
xylem (fig. 56a, E) as seen in cross section. Here the tracheal
tubes are relatively large and the wood fibers have compara-
tively large cavities. The next succeeding layer of xylem
(fig. 56a, F) consists of very small, thick-walled tracheal tubes,
A, and wood parenchyma cells, B, shown with greater magnifi-
cation in figure 57. In figure 58 are shown a few wood paren-
chyma cells as seen in longitudinal section. The average length
of these cells is 0.07 mm. and the average width 0.015 mm.
Figure 59 represents a few wood fibers and tracheal tubes
found in the next succeeding layer of xylem (fig, 56a, G) . The
average diameter of the tracheal tubes in this layer of xylem is
0.023 mm. ; here the wood fibers have remarkably thickened
walls and a smaller cavity than those found in the outermost
layer of xylem (fig. 56a, E).

Figure 56a, a segment of the cross section of the root, shows
the constitution of the xylem: A, wood fibers; B, tracheal ele-
ments ; C, wood parenchyma.

In figure 56& are shown the tracheal elements found in the
root: A and B, scalariform tracheal tubes, C, spiral tracheal
tube.

Tissues for the Conduction a7id Storage of Food. — In the
root are found only secondary medullary rays, which are nar-
rower than those found in the stem. In these medullary rays is
found no tracheal system as in those of the stem. As in the
stem, there seems to have been made very little provision for
the passage of food longitudinally. Yet, however, a larger pro-
portion (25 per cent) of the vascular bundle of the root is de-
voted to phloem than in the stem. The phloem consists ex-
clusively of undivided mother cells of the sieve tubes.

Starch and proteids in small quantities were found in the
medullary rays and phloem-parenchyma cells, but neither glu-
cose nor oil. In the root of Toiunsendia there was found
glucose, but neither starch, proteid nor oil.

202 KANSAS UNIVERSITY SCIENCE BULLETIN.

In figure 60 — a diagrammatic drawing of the cross section
of the root traced as seen when projected upon a screen — are
shown the different zones of tissues of which the root con-
sists and their relation to one another: A, cork; B, collen-
chyma ; C, stone cells ; D, thin- walled parenchyma ; E, phloem ;
F, wood fibers; G, wood parenchyma; H, tracheal elements;
/, medullary ray.

The Tegumentary Tissues. — The tegumentary tissues con-
sist of an epidermis with trichomes. All the walls of the epi-
dermis are cellulose except the outer tangential wall, which has
a thin film of cutin on the outer portion. The outer and inner
tangential wall each measures in thickness 0.006 mm., and the
radial wall 0.003 mm. The average radial diameter of the cell
cavity is 0.02 mm., the average tangential diameter 0.016 mm.
On both sides of the leaf the epidermis is of the same charac-
ter. Figure 61 represents a small portion of the epidermis:
a, cutinized portion; b, cellulose portion. The trichomes are
thickly distributed over both sides of the leaf, and must be
quite effective in giving protection. The discussion of these
trichomes will be taken up in connection with the water-
storage system.

The Tracheal Tissues. — For the study of venation, a leaf was
used which had been bleached by standing four or five hours
in dilute hydrochloric acid and then two or three days in
chloral hydrate. As shown in figure 62, there could be seen
in this leaf one large central vein, A, running longitudinally
throughout the length of the leaf and sending off near the
center of the leaf two large lateral branches, B, and below the
center the large branch, G. On either side of the midrib there
are two prominent longitudinal veins, C. These primary veins
send out other veins which branch in every direction and end
abruptly, as shown at D. The average distance between the
ultimate branches of the veins, E and F, is 0.16 mm., there
being included within this area, on the average, twenty-five
mesophyll cells. The phloem consists only of undivided mother
cells of the sieve tubes, whose average length is 0.16 mm. and
width 0.006 mm. The xylem consists exclusively of spiral
tracheary tracheids. The average diameter of the tracheids
in the midrib is 0.006 mm., and the average diameter of those
in the smaller veins is 0.003 mm. In the midrib, where the
cross-section area is 0.015 sq. mm., 20 per cent is devoted to

bunton: lesquerella spathulata. 203

the conduction of water. Figure 63 represents a small portion
of the midrib as seen in longitudinal section: A, spiral tra-
cheary tracheid ; B, undivided mother cell of the sieve tubes.

Tlie Photoi^ynthetic Tissues. — The leaf of Draba is bifacial,
that is, palisade cells occur on both sides of the leaf, which is
from twelve to fifteen layers of cells thick. The average radial
length of a palisade cell is 0.027 mm., the average cross diame-
ter 0.02 mm. Figure 64 represents a few palisade cells as
seen in surface view.

The average number of chloroplasts in a single mesophyll
cell on either side of the leaf is twenty-seven. The small num-
ber of chloroplasts found in the mesophyll cells of the upper
side of the leaf allows the sunlight to penetrate to the lower
side, thus giving to the cells there the power to carry on photo-
synthesis effectively.

The Water-storage Tissues. — In the leaf of Lesquerella there
are no water-storage cells as are found in that of Townseyidia.
The only structures in the leaf that might serve principally
as water-storage elements are the large trichomes which occur
on both sides of the leaf. These trichomes, as shown in figure
65 (surface view) are quite frequent, and are stellate-peltate
in form. These, as a rule, are found above a vein, and, since
they are alive even on old leaves, they presumably have some
active function to perform. Their basal portion is slightly
depressed beneath the lower level of the epidermis, and the
wall here is cellulose. The remaining portion of the wall of
the trichomes is cellulose, except the very outer portion, which
is cutinized. The fact that these trichomes occur above the
veins and that their basal portion is cellulose-walled seems
to point to the conclusion that when there is plenty of water
furnished by the veins they absorb water, store it up, and
yield it to the mesophyll cells as it is needed. Figure 66 rep-
resents a trichome as seen in a leaf cross section : A, cutinized
portion of a wall of trichome; B, cellulose portion of wall of
trichome ; C, mesophyll cells ; D, tracheids.

The Aerating Spaces of the Leaf. — Mounting the bleached
leaf so that it could be looked through from surface to surface,
I estimate that approximately 2.7 per cent of the volume of the
leaf is devoted to intercellular spaces ; in other words, the vol-
ume of the intercellular spaces and the volume of the whole
leaf, roughly estimated, stand in relation to each other as

204 KANSAS UNIVERSITY SCIENCE BULLETIN.

27 : 1000, On either side of the leaf are found 640 stomata to
1 sq. mm., a much larger number than was found in the leaf
of Townseiulia. The stomata are at the surface of the leaf in-
stead of being sunken, as is often the case with xerophytes.
In figure 67 are shown a few stomata as seen in surface view.
In figure 68 is shown a stoma as seen in cross section.

Figure 69 represents a small portion of a leaf as seen in
cross section: A, upper epidermis; B, lower epidermis; C,
palisade tissue; D, vascular bundle.

SUMMARY.

1. Size and Habit of Lesquerella spatJiulata. — Lesquerella
spatliulata, a perennial from a branched caudex, is a low plant
growing from one to two inches high. It was found growing
on a rocky clay bluff, occurring not as isolated individuals, but
in colonies.

2. Tegnmentary Tissues. — In the mature stem of Lesque-
rella the tegumentary tissues consist of a cutinized epidermis,
cork, and borke. In the mature root the tegumentary tissues
consist of a cutinized epidermis and cork.

The tegumentary tissues of the leaf consist of an epidermis,
all the walls of which are cellulose except a thin film of cutin
on the outer tangential wall. There is, in addition to the ordi-
nary epidermal cells, stellate trichomes on both sides of the
leaf, which would be effective in protection against too intense
illumination and in the reduction of transpiration, as well as
in the storage of water.

3. The Strengthening Tissues. — The strengthening tissues
consist of collenchyma, bast fibers, stone cells and wood fibers.
The wood fibers constitute a very important part of the
strengthening system. These have remarkably thickened
walls, the cavities in some cases being almost obliterated.
Contrary to the general rule, the wood fibers are found among
spiral tracheal tubes, both wood fibers and spiral tracheal
tubes having been formed by the cambium.

4. Tracheal Tissues. — The tissues devoted to the conduction
of water constitute 2 per cent of the stem and 4 per cent of the
root. This seems a comparatively small area devoted to this
purpose, but growing, as Lesquerella does, in a location where
the water supply is poor, it would not require very extensive

BUNTON: LESQUERELLA SPATHULATA. 205

water-conducting tissues. The tracheal tissues consist of
spiral and scalariform tracheal tubes and tracheids.

5. Photosynthetic Tissues. — The photosynthetic tissues of
the leaf consist exclusively of palisade cells, and there is found
in a single palisade cell on either side of the leaf an average of
twenty-seven chloroplasts.

6. Food-conducting Tissues. — As in Toivnsendia, the
phloem is but little differentiated. It consists exclusively of
undivided mother cells of the sieve tubes; sieve tubes and
sieve parenchyma cells are wholly lacking.

7. Storage Tissues. — In the stem the phloem cells and
medullary-ray cells afford provision for the storage of food.
In these tissues starch and proteids were stored up in small
quantities. The only storage tissues found in the leaf are the
large stellate hairs which, possibly, rapidly fill whenever the
water supply is increased, and, being cellulose-walled at their
bases, yield their contents to the mesophyll cells as the supply
of water is reduced.

8. Aerating Spaces. — As would be expected to be found in
a xerophyte, the amount of space devoted to aeration is com-
paratively small, there being only 2.7 per cent of the volume of
the leaf devoted to this purpose.

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THE

KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 12— March, 1910.

(Whole Series, Vol. XV, No. 12.)

CONTENTS:

Cell Changes in the Alveoli of a Carcinoma of the Mamma,

Earl F. Clark.

PUBLISHED BY THE UNIVERSITY,
LAWRENCE, KAN.

October, 1911.

Entered at the post-ofRce in Lawrence as second-class matter.

3310

THE KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 12] MARCH, 1910. [vo'^xv^noTb

CELL CHANGES IN THE ALVEOLI OF A CARCINOMA

OF THE MAMMA.

BY EARL F. CLARK.

(Contribution from the Zoological Laboratory, No. 188.)

Submitted in partial fulfillment of the requirements for the degree of Master of Arts.

Plates XLVI to XLVIII.

THE literature upon the pathological new growths of the
human body scarcely mentions amitosis as a method of
cell increase. It occurred to me to treat in a comparative
way the two methods of cell division, mitosis and amitosis, as
they are found in neoplastic material.

It is not intended to make this paper conclusive, but to make
it the first of a series of case studies from the results of which
a general conclusion may be deduced as to the relative im-
portance of indirect and direct cell division in these new
growths.

I am indebted to Dr. M. T. Sudler for the following history :
"In the first week of October, 1907, she fell, bumping the
breast against a hard object. No further notice was taken of
the incident, however, until the latter part of November, when
she noticed a hard, thin, round lump in the right breast about
the size of a silver dollar. It was then freely movable. About
the 25th of December she noticed a tingling in the right arm,
and on January 1, 1908, the arm began to ache. The original
mass increased rapidly in size until the whole breast was in-
volved. Its rate of growth seemed intermittent, some weeks
being much more rapid than others. Upon examination the
entire breast was found to be occupied by a hard, firm mass,
which was intimately incorporated in the connective tissue of

(209)

210 KANSAS UNIVERSITY SCIENCE BULLETIN.

the breast. The nipple was somewhat retracted but not com-
pletely. The mass was movable over the muscle. The axillary
nodes were enlarged and hard. On February 5 the breast, to-
gether with the axillary lymph nodes, was removed. On March
13 a recurrence in the left breast was discovered, which was
removed on March 20. Twelve days later metastatic nodules in
the neck were removed. The patient died soon afterwards
from an apparently general carcinomatous involvement of the
body. From the foregoing history it is plain that this was an
extreme type of the most rapidly growing and malignant of
tumors."

This material was not prepared especially for fine cytological
work because the discovery of its excellence for investigation
was not made until after its fixation. It was fixed in ten-per-
cent formalin and remained in a weaker solution until used.
I found the cell nests somewhat shrunken away from the sur-
rounding stroma and the chromatin either so swollen or so
poorly preserved that I was unable to make out details in re-
gard to the chromosomes. The fixation otherwise was good.
The tissue was mounted in paraffin and cut in sections eight
micra thick. Sections were stained in hsematoxylin for twelve
hours after being in a mordant of iron-alum for twelve hours.
They were then decolorized in the iron-alum solution to the
desired extent.

The cells studied appeared in variously shaped pockets or
alveoli surrounded by dense white fibrous connective tissue.
This connective tissue stroma was penetrated more or less by
columns of tumor cells, one to three cells in diameter. In
short, this specimen in parts conformed to the scirrhus while
in others it was more of the medullary type of carcinoma. In
this paper I will confine my study to the cell nests or alveoli.

In the first four figures I chose a typical field from widely
separated parts of the tumor for each drawing, and drew all
the structures appearing in that field. I did this to show the
similarity of the cell content of the tumor mass throughout.
The normal resting cell, with its variation in size, shape and
chromatin content, composed the greater number of cells of
each group. It was considered the normal resting cell because
of its uniform staining quality, its maintenance of cell and
nuclear wall, its content of regularly outlined chromatin nu-
cleoli within a single nucleus. Marked variation in any of these

CLARK: CYTOLOGY OF CARCINOMA. 211

characteristics was considered a cell change having as its
ultimate end either the production of new cells or degeneration.

Among the resting cells were others which could be easily
identified as being in the process of indirect division. Al-
though no detail of the chromosomes could be made out, yet the
increased amount of chromatin over the resting cell, its affinity
for the stain, the irregular outline of the chromosomes, the ab-
sence of the nuclear membrane and the shape and position of
the chromatin masses identical with those of the prophase,
metaphase, anaphase, and telophase leave no doubt as to the
identity of such cells. Equally distributed among the resting
cells were other cells which were clearly abnormal, owing to
their unusual size and their content of two or more nuclei or
clefts of greater or less depth which either separated or tended
to separate the single nucleus into two or more distinct seg-
ments. These separated segments had all the characteristics
of the nucleus of the resting cell. I have considered such cells
as in the process of direct cell division, although I have been
unable to find a single cell showing distinct separation of the
cytoplasm. Adami thus describes amitosis : "The nucleus
divides without any apparent preliminary rearrangement of
its structure. It becomes elongated, then dumb-bell shape (the
process can be followed in the amceba), the connecting neci<:
becomes broken across and the two daughter nuclei pass apart,
their separation being in some cases followed by division of the
cell body while in others this further division is wanting and
the binucleate cell is produced." Patterson, in "Amitosis in
the Pigeon's Egg," says : "In most cases the nucleus elongates
in the direction transverse to the plane of the future division,
after which a constriction in the nuclear membrane appears
about the entire circumference of the nucleus. This con-
striction continues to deepen until a complete division is ef-
fected. A modified form of this type is found in those cases
in which the constriction proceeds from one side."

Although the process in this material differs somewhat, I
think, from that described by Adami and Patterson for other
material, the result is the same. Therefore I feel safe in as-
suming that cells with the nucleus wholly or in various degrees
of separation are in the process of amitosis. Ziegler and
Vom Rath consider such cells as already in the process of de-
generation. If so, it will not be out of place to consider along
with these division figures structures that appear throughout

212 KANSAS UNIVERSITY SCIENCE BULLETIN.

this material and which are plainly cells in varying degrees of
degeneration, as evidenced by changes in the chromatin, cy-
toplasm and cell wall, and note if there is any similarity be-
tween them.

Figures 5, 6, 7 and 8 are used to further illustrate these three
cell changes as they appear in various parts of the tumor. In
these drawings I omitted the greater number of the resting
cells.

I will supplement this preliminary account by short de-
scriptions of individual cells in the several plates with inter-
pretations of their variation from the normal.

I have used the term "cell nests" to mean the group of cells
within the alveoli ; the term "chromatic cells" to mean those
cells in that condition in the degeneration process just before
the chromatin loses its staining power and when it has swollen
so much and become so diffuse as to fill the entire nucleus as
a more or less densely staining mass.

OBSERVATIONS.
The Cells of the Nests Change by Mitosis, Amitosis and Degeneration.

In showing the cell changes of these nests I will simply use
a number of figures, describing each one with some amount of
detail and inserting comments as observations suggest.

Figure 1 is drawn from cell nest 2. It includes three cells
dividing directly, one dividing indirectly, two cells in the
process of degeneration, and a number of resting cells.

The resting cell is smaller than the dividing cell, is of poly-
hedral shape, has a limiting membrane, and a large, well-
marked nucleus. Its cytoplasm is denser than the karyoplasm
and is of a reticular appearance. The nucleus has a well-
marked nuclear wall and usually contains from one to three
regularly round, deeply-stained chromatic bodies which I will
call chromatin nucleoli. A directly dividing cell is of much
larger size than the resting cell and usually a bit clearer, a
condition suggesting that it has included a large amount of
fluid preparatory to its division. The nucleus divides by the
formation of a groove or cleft on one side. I have not seen a
cell where the cleft appeared on the two sides. The cleft
deepens and the nuclear substance pulls apart to form the
regular outline of the daughter nuclei. The nuclear membrane
seems to follow the deepening cleft so that as soon as the seg-
ments have separated the karyoplasm is enclosed by its wall.

CLARK: CYTOLOGY OF CARCINOMA. 213

In the cells shown I have drawn them as they appeared in out-
line in one plane, but included the chromatin through the depth
of the cell. Each well-defined segment of the nucleus usually
contains a chromatin nucleolus. In cell A the lower nuclear seg-
ment has a second cleft appearing upon its inner border. Cell
B has two distinct nuclear segments, each one of which has a
secondary groove appearing upon its upper surface. The
upper segment shows no chromatin at all. Cell C shows two
nuclear segments, each containing a chromatin nucleolus. Cell
D shows a central constriction of its nucleus with the mother
chromatic mass in the plane of constriction, but I do not say it
is in the process of division.

Cell E is a representation of the typical indirect division
figures found in this material. It is in the early anaphase.
As I have already said, this material, being fixed in formalin
with no particular care regarding its cytological preservation,
shows no detail characteristics of the chromosomes. I have
used in my counts only the well-marked division figures, from
the close spireme stage on. In these indirect divisions the
cells are larger than the resting cells, the nuclear membrane
had disappeared, and about the chromatic figure the cytoplasm
was clear. I was unable to make out the spindle or any of the
details farther than shown.

Cells F F are two fragments of degenerated cells in each of
which the chromatin has swollen and coalesced into one homo-
geneous, deeply staining mass.

Figure 2. Among the cells of this group there is not the
noticeable difference in size usually seen between the dividing
cells and the resting ones. Cell A shows an indirect division
plainly. Cell B shows the nucleus being divided by a cleft
passing in from its upper surface. Each segment contains a
chromatin nucleolus. Cell C, an otherwise ordinary resting
cell, contains in its cytoplasm, half way from the cell border
to the nuclear membrane, a clear vesicle, regular in outline
and with a definite limiting border indicating that it is filled
with a substance differing from the cytoplasm.

Figure 3 shows no new structures, but attention is called to
the regularity of the chromatin nucleoli. Cells A and B show
the nuclei entirely segmented and each segment containing
regular chromatin nucleoli, the lower segment of cell B con-
taining two.

214 KANSAS UNIVERSITY SCIENCE BULLETIN.

Figure 4 is a group of twelve tumor cells drawn from cell
nest 1, a small nest of seventy-five cells containing six indirect
and no direct divisions. In this group are shown four of the
six indirect divisions. Cells A and B are noticeable for their
large size, while cells C and D are of the same size as the rest-
ing cells. Cell E is a common-sized resting cell, containing one
chromatin nucleolus but with a hollow clear vesicle bulging
into the wall of its nucleus. Cell F is a common resting cell ex-
cept that it has its chromatin diffused through the karyoplasm.

Figure 5. In this group and in those following I have not
included all the cells that appeared in the field, but simply in-
structive ones and enough resting cells to show a comparison.
The group was drawn from cell nest 3, of 253 cells, which in-
cluded six indirect and five direct divisions. Cell C was in the
fourth tier from the edge and on the peripheral side of the
group. It was large, being .0125 mm. in its greatest diameter
and showing a nucleus in two segments, each segment con-
taining a regularly round chromatin nucleolus. The cell was
noticeably clearer than the others. Cell B also has a biseg-
mented nucleus, but its chromatin was scattered in irregular
masses through the karyoplasm. It was also large, being .01
mm. in its greatest diameter. The nucleus of cell A was in
three segments, each part having a regularly round chromatin
nucleolus, and beside this a certain amount of chromatin scat-
tered in clumps through the karyoplasm. It was .0125 mm. in
its greatest diameter. In all my cell measurements I used a
micrometer scale which I have found by measurement to give
the following dimensions : With a drawing board on a level
with the object and at the same inclination as the microscope
stage, using ^2 oil-immersion objective and a 8X Huyghenian
ocular, an object that fills the square of the scale is .0125 mm.
in diameter; with the M; objective it is .03 mm. in diameter, and
with a % objective it is .15 mm. in diameter. Three-fourths
of the square equals .0093 mm. ; one-fourth of the square is
equal to .0031 mm. ; one-half of the square is equal to .0062
mm. Cell D is in the indirect division process. Cell £7 is a
structure that is common among the tumor cells and I have
concluded that it is a degeneration result, although it may
occur anywhere in the cell nest among the otherwise vigorous
cells. It is characterized by a large amount of deeply staining
chromatic material, homogeneous and closely massed, having

CLARK: CYTOLOGY OF CARCINOMA. 215

no definite form but with a more or less regular outline. This
mass evidently represents the nucleus, which has become filled
with a diffused, swollen mass of chromatin. Its cell membrane
has broken down, allowing the cytoplasm to flow out into the
surrounding intercellular spaces. Notice the regular form of
the chromatin nucleoli in this group of cells.

Figure 6. This group of eleven cells was drawn from the
same cell nest as the previous group. Cell A is a large cell,
.0093 mm. in its greatest diameter. In the cytoplasm about the
nucleus are four vesicles, similar to those described under
figures 2 and 4. The chromatin of its nucleus is of regular
outline but is unusual on account of its size, being a trifle over
one-half the diameter of the nucleus itself, at least twice the
amount found in the ordinary resting cell. I think it is a cell
just beginning the degeneration process shown by the swelling
of its chromatin. Cell C is undoubtedly in the process of de-
generation, for its cell membrane has disappeared, allowing
the cytoplasm to flow out uncontrolled. It is a cell almost as
large as cell A. Its chromatin, which exists as a deeply stain-
ing mass somewhat irregular in outline, was larger in diameter
than that of cell A. The nuclear wall could not be made out.
Cell B is one farther along in the process of breaking up than
cell C, for most of its cytoplasm has gone. The diameter of
its chromatic element is not so much larger than that of cell
C but it is more condensed and of more regular outline. These
three cells, I think, are respectively in advancing stages of dis-
integration. Cell Z) is a cell dividing indirectly, having a chro-
matic mass larger than that of cell C but of a looser arrange-
ment, indicating individual chromosomes. It also maintains its
cell borders perfectly.

Figure 7. This group of eleven cells was drawn from cell
nest 6 in the region of a necrotic area. Cells A and B are in
the fifth and sixth tiers respectively from the free edge of the
tumor on the peripheral side of the group. I will use this
group and the one following it, figure 8, to show changes in the
cells passing inward from the free edge of the cell nest to a
necrotic area. On the side peripheral to cell A there is a small
necrotic spot two or three cells in diameter, which accounts
for cells A and B on that side of the group. Cell A is a good
specimen of such cells as C and B of figure 6 or cell E of figure
5. It shows the swelling of the chromatin and the disin-

216 KANSAS UNIVERSITY SCIENCE BULLETIN.

tegration of the cytoplasm well. While there has been a
swelling of the chromatin in each of these cells there has been
an equally great shrinkage of the nucleus, so that at this stage
there has been a real loss of material to the nucleus. Cell B is
peculiar in that its nucleus, while practically the same size as
normal, stains very dark; not solid, as in A, but granular.
There is also a faint indication of its chromatin nucleolus re-
maining, not indicating so much that it has become diffused
through the karyoplasm as that the karyoplasm itself assumes
a new staining power. The cell wall of cell B is broken down
in one place, showing signs of disintegration. Cell C, a directly
dividing cell, is .0093 mm. in its greatest diameter, while cell
D is just a trifle less than .0125 mm. The average diameter of
the resting cells of this group is near .007 mm. So I think
that a cell preparing for direct division increases in size to a
marked extent.

Figure 8. This group of sixteen cells is taken from cell nest
6, the same nest that group 7 is taken from, and drawn from a
zone of cells to the inner side of group 7, that is, between
group 7 and a necrotic core which lies in the central part of
the nest. These two figures show well the transition from the
more peripheral nondegenerating cells to the entirely necrotic
mass of cell debris which fills the central portion of so many
of the cell nests. No dividing cells were shown. Cell A, the
most peripheral of the group, is the only one showing a chro-
matin nucleolus. It is an ordinary resting cell. In cells B
and B', a type of many cells found at this zone about the ne-
crotic area, and for that reason described as characteristic,
no chromatin nucleolus could be made out through the depth
of the cell, but in each of these the chromatin was scattered
through the karyoplasm in the form of a loose network, stain-
ing lightly. The cell and nuclear membranes were preserved
perfectly. Passing inward toward the center of the nest is a
field of cells which I have called ''chromatic cells," and which
are perfectly characteristic of a zone of cells that immediately
surrounds the necrotic core of the cell nest. They are a
transition stage between the normal cells and the wholly de-
generated ones. They resemble closely the disintegrating cells
described in figures 5, 6 and 7, but seem to have another history
of formation. As in the other cells, the nucleus is filled with a
homogeneous chromatic mass, the outer layers staining darker

CLARK: CYTOLOGY OF CARCINOMA. 217

than those farther toward the center. In this field the stain
was not as intense as that in the cells of the previous figures but
was of a grayish shade, and passing toward the necrotic area
the light in the gray increases until in cells D and D' there is
just enough black remaining to outline the nucleus well. From
this stage it gradually fades out until the cells are entirely ne-
crotic and do not stain at all. The cell membrane in this
group could not be decided upon satisfactorily. In cells A,
B and B' it was perfectly intact, while in cells E it seemed to
be entirely gone. In other cells, as C, C, the cell outline was
definitely preserved, while yet again in cells D and D' the cell
boundary was irregular and indefinite. But I am safe in say-
ing that when the cell has so far degenerated that the color
has disappeared from the nucleus the cell membrane has also
disappeared and disintegration immediately follows. But in
many of these cells the cell membrane has at least partially
gone while the chromatin stains deeply. The nucleus of cell
B' is 0.004 mm. in diameter while the chromatin mass of cell
C is near 0.003 mm. in diameter, showing an appreciable
shrinkage of this element in the degeneration process.

The Mitotic Divisions Occur in General Upon the Periphery of the
Alveolus, the Amitotic Nearer the Center.

I will use camera lucida drawings, together with tables, to
prove this. I counted the number of cells, the number of di-
rect divisions and the number of indirect, and also counted the
tier of cells from the periphery of the cell nest in which each
division occurred. I hoped by this method to arrive at some
conclusion as to the rapidity of growth of the tumor, in order
to compare it later with the growth of the adult and em-
bryonic mammary tissue. I also wished to decide as to the
relative frequency in occurrence of the direct and indirect di-
visions and to note, if possible, whether there was any com-
parison between the relative proportions of these and the sort
of nests in which they occurred. I also wished to decide if
there was a well-founded relation between either of these
methods of cell division and the necrotic areas indicating that
one or the other was related to degeneration, or that they
occurred equally in the same region showing that they were
equally vital methods of cell growth.

Cell Nest 2 was the non-necrotic limb of a necrotic body.
It was counted up to the beginning of the necrotic area. I can-

218 KANSAS UNIVERSITY SCIENCE BULLETIN.

not give the tiers of cells in which the various divisions oc-
curred in this nest.

No. of cells. Direct divisions. Indirect divisions.

563 9 16

Percentage of cells in divisions 4.42

" of cells in direct divisions 1.50

" of cells in indirect divisions 2.90

" of divisions which ai'e direct 36

" of divisions which are indirect 64

Cell Nest 3. This nest apparently is a small nest in di-
ameter but it has four "chromatic cells," which indicates that
it is somewhat degenerating.

Tier 123^56789 10 11 Av. Total

Direct D 2 1 ... 2 ... . 6.2 5

Indirect D 3 1.11 2.3 6

No. of cells. Direct divisions. Indirect divisions.

253 5 6

Percentage of cells in divisions 4.34

" of cells in direct divisions 1.90

" of cells in indirect divisions 2.40

" of divisions which are direct 45.40

" of divisions which are indirect 54.50

Cell Nest 4. This was a nest with a necrotic center .125
mm. to .1125 mm. in diameter, while around this on the edge
of the nest was a zone of cells varying in width from .05 mm.
to .037 mm. which was undegenerate and which constituted
the number of cells counted, so that the total number of cells
counted does not show the real size of the nest in point of num-
ber of cells, one-half of the area of the nest being degenerate.

Tier 123ji.56789 10 11 Av. Total

Direct D 1.312111. 1 1 5.5 12

Indirect D 12 11.11. 1 . 5.2 8

No. of cells. Direct divisions. Indirect divisions.

515 12 8

Percentage of cells in divisions 3.88

" of cells in direct divisions 2.30

" of cells in indirect divisions 1.50

" of divisions which are direct 60

" of divisions which are indirect 40

Cell Nest 5. This was one of the smaller sized nests and
was quite vigorous, showing no degenerating cells at all.

Tier 123^56789 10 11 Av. Total

Direct D 00 00

Indirect D 3 . . 1 1.7 4

CLARK: CYTOLOGY OF CARCINOMA. 219

No. of cells. Direct divisions. Indirect divisions.

106 00 4

Pei'centage of cells in divisions 3.77

of cells in direct divisions 0

of cells in indirect divisions 3.77

of divisions which are direct 0

of divisions vi^hich are indirect 100

a
II
It
a

Cell Nest 6. This was a cross section of a large alveolus,
being .2875 mm. in length and .1758 mm. in width. It was
regularly oval in outline. There were two degenerate spots
in it at opposite sides, the size of six to eight cells. With these
exceptions the cells were all counted.

Tier 123^56789 10 11 Av. Total

Direct D 14 1.3.21.. . . 5.0 12

Indirect D 1 1 .... 2 5.2 4

No. of cells. Direct divisions. Indirect divisions.

676 12 4

Percentage of cells in divisions 2.36

" of cells in direct divisions 1.70

" of cells in indirect divisions 0.60

" of divisions which are direct 75

" of divisions which are indirect 25

Cell Nest 7. This was a small, oval, non-necrotic nest.

Tier 123^56789 10 11 Av. Total

Direct D 00 00

Indirect D 14 1.8 5

No. of cells. Direct divisions. Indirect divisions.

123 00 5

Percentage of cells in divisions 4.06

" of cells in direct divisions 0

" of cells in indirect divisions 4.06

" of divisions which are direct 0

" of divisions which are indirect 100

Cell Nest 9. This was a large necrotic nest, .2125 mm. in
diameter, with a necrotic center .1125 mm. in diameter, which
had a border of vigorous cells .05 mm. wide.

Tier 123^56789 10 11 Av. Total

Direct D 1 . 4 . 1 1 . . . 1 . . 4.2 8

Indirect D 1.1 2.0 2

No. of cells. Direct divisions. Indirect divisions.

299 8 2

Percentage of cells in divisions 3.34

" of cells in direct divisions 2.60

" of cells in indirect divisions 0.70

" of divisions which are direct 80

" of divisions which are indirect 20

220

KANSAS UNIVERSITY SCIENCE BULLETIN,

In the following cell nests I have taken the serial sections of
the same nest and counted them in the same way as I did the
single sections of the previously counted nests. I hope in this
way to in a manner corroborate my results and also to study
the history of the cell nest farther.

Cell Nest 10. This nest was .4125 mm. in length, .0875
mm. in width. It was a nest that showed occasional spots of
six to eight cells that had broken down, an average of two of
these spots in each nest. The direct divisions occurred largely
in the upper, wider end, while the indirect occurred largely in
the lower, narrower end, but I have not illustrated this by
sketches. It was therefore taken as a nest, non-necrotic, but
near the stage when it would rapidly have become so, indicated
by the degenerate spots.

Cell Nest 10, A.
Tier. 1 2 8 ^ 5 6 7 8 9 10 11 Av. D.D. I.D. Cells.

Direct D. . . 1 2 1 2 1 . . 1 . . 5.6 8

Indirect D. 1 6 2 3 . 2 3.0 . . 14 532

Cell Nest 10, B.

Direct D. 1 1 2 5 1 2 4.2 12

Indirect D. 2 3 3 3 2.6 .. 11 600

Cell Nest 10, C.

Direct D. . 3 1 . . 3 3.8 7

Indirect D. 2 2 . 3 2 3.1 . . 9 557

Cell Nest 10, D.

Direct D. . 1 1 3 2 2 1 4.6 10

Indirect D. 2 3 2 2 2.4 . . 9 655

Cell Nest 10, E.

Direct D. . . 3 2 2 1 1 4.4 9

Indirect D. 1 1 2 1 1 1 3.4 . . 7 654

Cell Nest 10, F.

Direct D. . . 4 3 2 2 4.1 11

Indirect D. 2 2 3 4 . 2 1 3.5 . . 14 650

Cell Nest 10, G.

Direct D. . . 1 . . 4 2 5.8 7

Indirect D. 3 7 2 1 . 1 1 2.6 . . 15 564

No. of cells. Direct divisions. Indirect divisions.

4212 64 79

Percentage of cells in divisions 3.39

" of cells in direct divisions 1.50

" of cells in indirect divisions 1.80

" of divisions which are direct 44

" of divisions which are indirect 56

The direct divisions averaged 1.68 rows nearer the center
than the indirect divisions.

CLARK: CYTOLOGY OF CARCINOMA. 22l

Cell Nest 11. This nest was probably one-half of a half
necrotic alveolus. A sketch of it in cell nest 11 A is given in
figure 9. The cross-hatched area shows that part of the nest
that is necrotic, while the white border about it is the area of
the cells counted.

Cell Nest 11, A.

Tie7\ 1 2 3 4 5 a 7 8 9 10 11 Av. D.D. I.D. Cells.

Direct D. . 2 . 1 2.6 3

Indirect D. . 1 . 2 3.3 . . 3 334

Cell Nest 11, B.

Direct D. . 3 . 1 . 3 . 1 . 1 . . 5.1 9

Indirect D. 2 . 1 1.6 . . 3 305

Cell Nest 11, C.

Direct D. ... 1 1 3 . 1 5.8 6

Indirect D. 1 4 . 1 . 1 2.7 . . 7 279

Cell Nest 11, D.

Direct D. 1 . 2 1 1 2 1 4.3 8

Indirect D. 1 2 1 2.0 . . 4 296

Cell Nest 11, E.

Direct D. . 1 1 . 2 . . . 1 ... 4.8 5

Indirect D. 1 1 . 1 1 3.0 . . 4 279

Cell Nest 11, F.

Direct D. . . 1 . 1 . 1 . . 1 . . 6.2 4

Indirect D. 2 . 2 . . 3 3.7 . . 7 271

Cell Nest 11, G.

Direct D. . 2 1 1 3 . . 1 1 1 . . 5.3 10

Indirect D. 3 1 . . 1 2.0 . . 5 404

Cell Nest 11, H.

Direct D. . 2 2 1 . . 1 . 1 . . . . 4.2 7

Indirect D. . 3 1 1 1 3.0 . . 6 391

Cell Nest 11, I.

Direct D. . . 2 . 2 2 . . 1 1 . . 5.8 8

Indirect D. . 3 1 2 2 . 1 3.6 . . 9 393

Cell Nest 11, J.

Direct D. . 1 .... 2 5.3 3

Indirect D. 1 . 1 3 1 3.5 . . 6 306

No. of cells. Direct divisions. Indirect divisions.

3258 63 54

Percentage of cells in divisions 3.59

" of cells in direct divisions 1.90

" of cells in indirect divisions 1.60

" of divisions which are direct 53.80

" of divisions which are indirect 46.10

The direct divisions in this nest averaged 2.1 rows farther
away from the free edge of the nest than the indirect, or 2.1
rows nearer the center of the nest.

222 KANSAS UNIVERSITY SCIENCE BULLETIN.

Cell Nest 8. The work upon this nest was done with
greater care than that upon the previous nests. I first made a
camera lucida sketch of the nest in outline, then went over the
nest carefully with the high power 8x compensating ocular
and marked each division I found upon the outlined sketch. I
then used an 8x Huyghenian ocular, upon the diaphragm of
which I had placed a micromillimeter scale, and went over the
field again, carefully counting the cells by the aid of the mi-
crometer scale, and verified the previous locations of the divid-
ing cells and located any new ones that I may have missed in
the first survey. I thus corroborated my work and made it
doubly sure by locating each dividing cell upon an outline
sketch. I also used these sketches in the hope that together
with the tables I might farther establish the apparent relation
of directly dividing cells to the degenerating areas. Where
this relation has been especially noticeable I have made high-
power camera lucida drawings of the particular field to show
some detail about the cells I counted, simply drawing the more
important structures in relation to them. Using these large
drawings and marking their position upon the smaller sketches
one cannot avoid showing in the same way the slide does the real
relation of the various dividing cells to these necrotic areas.
In these outline sketches I only outlined the nest, tracing into
any irregularities among the cells along the edge, so that, un-
less otherwise marked, rifts in the nests will mean spaces left
where the cells have apparently been pulled apart by external
means. The areas that are entirely necrotic will not be dis-
tinguished from those that show definite degenerating changes ;
cells that are in the "chromatic cell" stage I have counted as ne-
crotic. These areas are shown by cross-hatched work. The in-
direct divisions are shown by a dot or "X" while the direct
divisions are shown by a circle. Starting with figure 10 the
sections are in serial arrangement. This cell nest 8 is only a
portion of the entire alveolus, showing at the right side of the
sketch just the end of a necrotic plug, and leaving the greater
part shown in the vigorous condition. I used this nest to con-
trast, if possible, the two divisions in these two areas.

CLARK: CYTOLOGY OF CARCINOMA. 223

Fig, 10. Cell Nest 8, A.
Tier. 1 2 3 A 5 6 7 8 9 10 11 Av. D.D. I.D. Cells.

Direct D. . 1 . 1 3 . . 2 1 . . . . 5.4 8

Indirect D. 2 4 4 1 2 2 3.2 . . 15 534

Fig. 11. Cell Nest 8, B.

Direct D. . 1 2 3 2 . 1 4.1 9

Indirect D. 1 4 4 3 1 2 3.3 . . 15 511

Fig. 12. Cell Nest 8, C.

Direct D. . 1 1 1 5 1 3 2 1 .... 5.7 15

Indirect D. 7 3 3 3 2 1 4 1 3.5 . . 24 554

Fig. 14. Cell Nest 8, C.

Direct D. . 1 2 2 2 2 . 1 4.6 10

Indirect D. 2 2 2 5 2 1 1 1 4.0 . . 16 555

Cell Nest 8, D.

Direct D. 1 2 . 1 2 3 2 5.1 11

Indirect D. 3 1 3 3 4 1 1 3.6 . . 16 626

SUBNEST 8, D.

Direct D. ... 1 1 4.5 2

Indirect D. 3 5 1 1.7 .. 9 172

Cell Nest 8, E.

Direct D. . 1 2 2 3 2 1 4.9 11

Indirect D. 4 3 7 4 4 3.0 . . 22 869

Cell Nest 8, F.

Direct D. . . 3 2 6 2 3 5.0 16

Indirect D. 3 5 3 2 2 2 ........ 3.0 . . 17 745

Fig. 16. Cell Nest 8, G.

Direct D. . 2 3 1 3 4 1 1 4.7 15

Indirect D. 2 6 2 1 . 1 1 2.8 . . 13 916

Fig. 18. Cell Nest 8, H.

Direct D. 1 2 3 1 . 3 3 4.3 13 ... ...

Indirect D. 3 1 7 1 .. 2 . 1 .... 3.5 . . 15 907

Fig. 19. Cell Nest 8, I.

Direct D. 1 . 2 1 2 . 1 3 . . . 1 5.7 11

Indirect D. 1 1 7 2 4 1 1 3.8 . . 17 912

No. of cells. Direct divisions. Indirect divisions.

7301 121 179

Percentage of cells in division 4.01

" of cells in direct division 1.60

" of cells in indirect division 2.40

" of divisions which are direct 40.33

" of divisions which are indirect 59.66

The direct divisions of this nest averaged 1.78 rows nearer
the center than the indirect.

2-Univ. Sci. Bull., Vol. V, No. 12.

224 KANSAS UNIVERSITY SCIENCE BULLETIN.

DESCRIPTION OF OUTLINE SKETCHES AND DRAWINGS.

Cell Nest 8, "A" (fig. 10) . The amount of necrotic material
in this figure is greater than in the following ones, the necrotic
plug showing less and less in each consecutive section. There
are no divisions shown in this figure near the necrotic plug,
but the nearest two are direct. The greater part of the divid-
ing cells are in the limb of the nest.

Cell Nest 8, "B" (fig. 11). The number of divisions pres-
ent in the distal part of the limb is noticeable. There are three
direct divisions immediately adjacent to the necrotic plug and
one nearer than the nearest indirect division.

Cell Nest 8, "C" (fig. 12). In this nest there was an un-
usually large number of dividing cells. At the point of the
arrow in figure 12 is the center of the group of cells drawn on
a large scale in figure 13. About the arrow point are the four
direct divisions and the two indirect divisions shown in the en-
larged group. This group of nineteen cells includes a larger
area under the lens than any previously drawn. I drew all the
cells of importance under the field of the camera lucida, in-
cluding a field .05 mm. in diameter, while the previous groups
drawn have included an area only .0375 mm. in diameter. It is
located on the edge of the necrotic area and includes the whole
width of cells lying between this area and the free edge of the
cell nest, cell F being in the first row of the nest, while cell B is
among the necrotic cells. The direct divisions of this section
are deep in the center of the nest, while the indirect are
largely very close to the edge, seven cells out of twenty-five
dividing indirectly lying in the first row and no directly di-
viding cells nearer than the second row, and then only one.
A general survey of these sketches together with the tables
strongly indicates that the cells on the edge of the tumor
increase almost, if not wholly, by indirect divisions, while those
deeper away from the edge increase largely by direct divisions
but occasionally by indirect. I want to emphasize by the large
drawings the intimate association of the directly dividing cells
with the degenerate ones. Cells H, H', H" are the "chromatic
cells" that I have described previously under figure 13. Sur-
rounding these cells closely are cells A, B, C, D and E, plainly
in the process of direct division. Cell B is peculiar in that it
shows no chromatin nucleolus in either segment of the
nucleus. This same occurrence was noticed in numbers

CLARK: CYTOLOGY OF CARCINOMA. 225

of resting cells, B, B' , figure 8, etc., just outside the zone
of "chromatic cells" H, H', H". Its chromatin had wholly
or largely disappeared, and if any remained it was loosely
diff'used through the karyoplasm. Cell E is another cell show-
ing no chromatin in any of the segments of its nucleus. It
evidently was in decay for it was very translucent. The frag-
ments of its nucleus showed no chromatin at all. It may be that
a cell such as B, which has vigor enough to cause its nucleus
to segment but not to cytoplasm, gradually loses its staining
ability and its definiteness of outline and later becomes merely
a mass of debris through a stage represented in cell E. Cells
F and G are two cells in the indirect division process. The di-
rectly dividing cells are much larger than the resting cells or
those dividing indirectly. Cell A is .0093 mm. in its greatest
diameter, cell B is .007 mm., cell C is .0093 mm., cell G is .0062
mm., cell F is .0062 mm., cell E is .0093 mm., cell D is .0085
mm., while cell /, an average resting cell, is .0062 mm. in its
greatest diameter. This cell shows the breaking up of the
chromatin and its diffusion through the karyoplasm, possibly
preceding cells H, H', H". Its nucleus is .005 mm. in diameter,
while the diameter of the "chromatic mass" in cells H, H', H"
is .0031 mm., showing evidently a shrinkage in the latter.

Figure 14, In this section two indirect divisions are shown
located on the very edge of the necrotic plug, and two more
near. There are also close to the edge of the nest three direct
divisions, still showing a greater proportion of direct divisions
near the necrotic area than indirect, when compared with the
total number of divisions of each in the section. But this
shows that indirect division may occur on the very edge of a
degenerating area.

At the lower extremity of the limb of the regular section
following is found a circular group of cells, which in the next
section shows a connection with the larger body. This is the
first appearance of this group, but in the previous sections
there is a circular space indicating where it has been but from
which it has in some way been lost. There is a cleft entirely
through the regular section, but this was where the cells had
pulled apart and not where they had degenerated. The direct
divisions are intimately associated with the necrotic spots. In
the middle segment there are two indirect divisions on the edge

226 KANSAS UNIVERSITY SCIENCE BULLETIN.

of the necrotic area, but usually they are near the free edge of
the nest.

In the succeeding section the subnest has joined the limb of
the main body and it was all counted together. There are
three necrotic areas in this section. In the lower one the two
directly dividing cells of that region are on the edge of the
degenerated area.

A later section shows the nest in two segments, broken apart
from each other. A pluripolar cell division shows at one place.

The group of cells drawn under high power in figure 15 is
from this section. Figure 15 shows a group of five directly and
one indirectly dividing cells. In this drawing and those made
hereafter I shall indicate the segments of the nucleus as they
appear through the depth of the cell, demonstrating their over-
lapping. The border line used was drawn at the edge of the
cells which presented no marked degeneration changes, but it
is not a hard, fast line, a few vigorous cells jutting out farther
than the line and a few chromatic cells being behind it. Cells
A, B, C, D and E show well-marked direct divisions. Cell A
is on the necrotic border. Cell B, a very small cell for a di-
rectly dividing one, is a half cell width from the necrotic
border. Cell C is a trifle closer than B to the border. Cell D
is in the third row from the border. It is peculiar in that its
chromatin nucleoli in each segment are not staining deeply and
the chromatin is diffused through the karyoplasm. Cell E is in
the fourth row from the necrotic edge. Cell F is an indirectly
dividing cell and is in the same row of cells as cells C and B.
Cells G and G' are the "chromatic cells" described under
figure 13. They mark the edge of the necrotic plug and are in
various stages of degeneration, as indicated by the disintegra-
tion of their cytoplasm and the loss of the staining power of
their chromatin. This sketch together with the smaller one
shows well this collecting of large numbers of the directly di-
viding cells about these necrotic areas. There are mitotic cells
here, but not in such proportion in comparison to their total
number as the amitotic.

Figure 16. In this section there are more direct divisions
than indirect. There is a noticeable change in the number of
indirect divisions in the lower part of the nest, in what was
termed a "subnest" in a preceding section. In one section
there are nine indirect divisions, two direct. In the next there

CLARK: CYTOLOGY OF CARCINOMA. 227

are eight indirect divisions, two direct. In the next there are
five indirect divisions, three direct. In a later (fig. 16), there
are five indirect divisions, four direct. In another (fig. 19),
there are four indirect divisions, three direct. This shows
consistently in this case that as the center of the nest is ap-
proached in the sections the number of direct divisions increase
in proportion while the indirect decrease in proportion. In the
region of the arrow point of figure 16 an enlarged drawing was
made (figure 17). I used this to show more detail about the
cells shown in figure 16. Cell A was on the edge of the necrotic
area. Its chromatin was small in amount and gathered in two
clumps in one segment and three in the other. Cell C is an
ordinary dividing cell. Cell D is in the second row from the
necrotic edge. The cross-hatched work shows the necrotic
area, this area being entirely necrotic and showing no transi-
tion cells along the edge.

Figure 18. There are but two dividing cells in the upper
segment of this nest, while in an earlier there are fourteen, the
number decreasing with each section until this one. They
have increased steadily in the middle part of the limb, however,
from eleven to nearly twenty in the section shown in figure 18.
This carefully worked out will demonstrate that at the same
'time the cells in one part of the nest are rapidly dividing those
in another part are not dividing at all. There is either an
unequal stimulus to growth in the nest or an unequal negative
cause.

Figure 19. This nest is the same one as that in the preced-
ing figure, but in this section the two parts have been sepa-
rated. At the point of the arrow is the location of figure 20,
and here are shown the three dividing cells drawn in figure 20.
In figure 20, cell A is plainly among the degenerating cells in
the necrotic plug, and is surrounded by "chromatic cells," no
vigorous cell being near. It is a large clear cell with a biseg-
mented nucleus and showing but very little chromatin. Its
appearance indicates that it is a degenerating cell. Cell B
lies just outside the border of vigorous cells. Its nucleus is bi-
segmented and it stains much deeper than the cytoplasm. The
chromatin nucleoli stain very faintly. Cell C is a large clear
cell lying in the third row of cells from the necrotic edge. It
stains lighter than the other cells about it and shows a small
amount of chromatin and that stained faintly. Cells A and B

228 KANSAS UNIVERSITY SCIENCE BULLETIN.

are noticeable because they have broken off from the vigorous
growth and lie among cells that are distinctly necrotic.

Out of the 294 direct divisions in all the nests only 7 occurred
in the first row on the edge of the nest.

Out of the 357 indirectly dividing cells 63 occurred in the
first row on the edge of the nest.

The direct divisions in all the nests averaged 1.68 rows
nearer the center than the indirect divisions.

Cell Nest 12. This nest was sketched and counted in
forty-two serial sections. It began as three or four groups
of cells which were the cross sections of limbs which later
fused into a common trunk. It was non-necrotic throughout.

Cell Nest 12, I.

Tier 12 3 4 5 6

Direct D 1 . . .

Indirect D 2 1 4 . . .

Cell Nest 12, II.

Direct D

Indirect D 4 . i i ! '. 2.8 '. 6 140

Direct D

Indirect D 1 2.0 . 1 78

Av.

D.D.

I.D.

Cells

3.0

1

2.2

,

7

166

Cell Nest 12, III.

.2

2.0

. 1

2.0

Cell Nest 12, IV.

. . 1 . . . .

3.0

2 . . 1 . . .

2.0

Direct D

Indirect D 2 . . 1 . . . 2.0 . 3 77

Cell Nest 12, V.

Direct D

Indirect D 2 3 1 .

Cell Nest 12

Direct D 1 .

Indirect D 1 1

Cell Nest 12

Direct D 1 1 .

Indirect D 2 1

Cell Nest 12

Direct D

Indirect D 1 3 . 3

Cell Nest 12

Direct D 1 1

Indirect D 1 1 1

VI.

VII.

VIII.

IX.

1.8 . 6 180

3.0 1

2.5 . 2 81

2.5 2

2.3 . 4 85

2.7 . 7 83

3.5 2

2.0 . 3 86

Cell Nest 12, X.

Direct D 2 . . . 4.0 2

Indirect D 2 2 1.4 , 4 169

CLARK: CYTOLOGY OF CARCINOMA. 229

Cell Nest 12, XI.

Direct D

Indirect D 3 3 2 . . . . 1.9 . 8 206

Cell Nest 12, XII.

Direct D •

Indirect D 2 2 2 . . . . 2.0 . 6 227

Cell Nest 12, XIII.

Direct D 1 1 . . 3 . . 3.6 5

Indirect D 6 1 1.1 . 7 235

Cell Nest 12, XIV.

Direct D 4 . . . 4.0 1

Indirect D 5.2.... 1.5 . 7 360

Cell Nest 12, XV.

Direct D 1 . . 5.0 1

Indirect D 3.2.11. 2.8 . 7 328

Cell Nest 12, XVI.

Direct D 1 . . . 4.0 1

Indirect D 3.1121. 3.2 . 8 335 ■

Cell Nest 12, XVII.

Direct D 1 1 . . . 3.5 2

Indirect D 4 2 11. . . 1.9 . 8 365

Cell Nest 12, XVIII.

Direct D 1 . . . 4.0 1

Indirect D 9 . 2 . 1 . . 1.6 . 12 362

Cell Nest 12, XIX.

Direct D 1 1 . 5.5 2

Indirect D 5 . . . . 1 . 1.6 6 . . 309

Cell Nest 12, XX.

Direct D 1 . . 1 . 4.5 2

Indirect D 13.1.1. 2.5 . 6 332

Cell Nest 12, XXI.

Direct D 2 2.0 1

Indirect D 2 1 1.3 . 3 215

Cell Nest 12, XXII.

Direct D .

Indirect D 14 11. . . 2.4 . 7 209

Cell Nest 12, XXIII.

Direct D 1 1 . 1 1 . . 3.0 4

Indirect D 2 . 2 . . . . 2.0 . 4 254

Cell Nest 12, XXIV.

Direct D . 1 2.0 1

Indirect D 6 1 2 . . . . 1.5 . 9 288

Cell Nest 12, XXV.

Direct D 1 . . 5.0 1

Indirect D 2 3 1.7 . 5 449

230

KANSAS UNIVERSITY SCIENCE BULLETIN.

Cell Nest 12, XXVI.

Direct D 1 1 . 1 .

Indirect D 5 4

Cell Nest 12, XXVII.

Direct D 1 2 . . 1 .

Indirect D 6 3 2 . . . .

Cell Nest 12, XXVIII.

Direct D 1 1 2 . .

Indirect D 3 2 1 1 1 .

Direct D. ,
Indirect D.

Cell Nest 12, XXIX

. 1 . . . .
3.22. .

Cell Nest 12, XXX.

Direct D 2 . . .

Indirect D 5 2 1 . . .

Cell Nest 12, XXXI

Direct D 1

Indirect D 3 2 1 . . .

Cell Nest 12, XXXII

Direct D 1 . . . 1

Indirect D 1 2 1 . . .

Cell Nest 12, XXXIII.

Direct D 1 . . . 1 1 .

Indirect D 5 1 1 . . . .

Cell Nest 12, XXXIV.

Direct D 1 1 . . 1 . .

Indirect D 2 . 1 . 1 . .

Cell Nest 12, XXXV.

Direct D 1 . . . 2 .

Indirect D 5 2 . . 1 . .

Cell Nest 12, XXXVI.

Direct D 1 1 .

Indirect D 4 3 1 2 . . .

Cell Nest 12, XXXVII.

Direct D 3 . 1 . . .

Indirect D 3 6 1 1 . . .

Cell Nest 12, XXXVIII.

Direct D 1 . . . 1 .

Indirect D 6 2 1 1 . . .

Cell Nest 12, XXXIX.

Direct D 1 . . 3 . 1 1

Indirect D 5 4 1 1 . . .

Cell Nest 12, XL.

Direct D 2 , .

Indirect D 6 2 2 . . . .

4.3
1.4

3

'9

sie

3.5
1.6

4

ii

230

3.2

2.4

4

'8

334

2.0
2.4

1

'7

293

3.0
1.5

2

"s

307

1.0
1.6

1

6

3'34

4.0
2.0

2

'4

28i

4.0
1.4

3

'7

435

2.6
2.5

3

*4

38i

4.6
1.7

3

"s

43i

5.5
2.1

2

io

479

2.5
2.0

4

ii

537

4.0
1.7

2

io

55i

4.3
1.8

6

ii

498

5.0
1.6

2

io

4i6

CLARK: CYTOLOGY OF CARCINOMA. 231

Cell Nest 12, XLL

Direct D 2 2.0 2

Indirect D 2 3 4.6 . 5 371

Cell Nest 12, XLII.

Direct D 1 1 . . 2 . 4.2 4

Indirect D. 4 2 3. . 1 . 2.3 . 10 421

No. of cells. Direct divisions. Indirect divisions.

12,228 81 285

Percentage of cells in divisions 2.99

" of cells in direct divisions 6

" of cells in indirect divisions 2.3

" of divisions which are direct 22.1

" of divisions which are indirect 77.9

Table of all the nests counted, showing percentage of cells
in division, percentage of cells in direct division, and percent-
age of cells in indirect division :

Percentage of

Percentage

of

Per

•centage of

Cell nest.

cells in

cells in

cells in

division.

direct divisio7i.

indirect division

2

4.42

1.5

2.9

3

4.34

1.9

2.4

4

3.88

2.3

1.5

5

3.77

0.0

3.77

6

2.36

1.7

.6

7

4.06

0.0

4.06

8

4.01

1.6

2.4

9

3.34

2.6

.7

10

3.39

1.5

1.8

11

3.59

1.9

1.6

12

2.99

.6

2.3

The writer of this paper wishes here to acknowledge his in-
debtedness to Doctor McClung, whose kind suggestions made
this work possible, and to Doctor M. T. Sudler for the material
and clinical data upon which this study is based.

CONCLUSIONS.

1. The percentage of cells in division shows no relation to
the percentage of amitoses or mitoses.

2. The percentage of mitoses is high in small non-necrotic
nests.

3. The percentage of amitoses is high in large necrotic nests,
indicating that it is the division associated with degeneration.

4. Amitosis is the form of division occurring in the necrotic
area.

5. The rate of division in the various nests is inconstant.

6. The directly dividing cells lie nearer the center of the cell

3-Univ. Sci. Bull.. Vol. V. No. 12.

232 KANSAS UNIVERSITY SCIENCE BULLETIN.

nests and thus nearer the necrotic areas than the indirectly
dividing cells.

7. The indirectly dividing cells lie nearer the periphery of
the nest than the directly dividing cells.

8. From these observations it may therefore reasonably be
concluded that amitosis represents the final effort of the cell at
reproduction, and that shortly after its occurrence degenera-
tion ensues.

9. Amitosis, mitosis and disintegration comprise the cell
changes in the alveoli.

10. Cells in direct and indirect division are larger than the
resting cells.

11. In one stage of degeneration of the cells the chromatin
becomes diffused through the karyoplasm and so swells that it
fills the nucleus as a deeply staining, homogeneous m^ass. In
another process of degeneration the chromatin loses its staining
power and the cell becomes clear.

12. There is a shrinkage of the nuclear bulk in degeneration.

13. A border of chromatic cells lies between the vigorous
cells and the cell debris of the necrotic area and marks the
limit of degeneration; such cells are themselves degeneration
results.

BIBLIOGRAPHY.

C. M. Child. Studies on the Relation Between Amitosis and Mitosis.
Biological Bulletin, vol. XII, No. 2, January, 1907.

J. George Adami. General Pathology.

J. E. S. Moore and C. E. Walker. First Report on the Cytological In-
vestigation of Cancer, 1906.

Alfred Stengel. General Pathology.

J. T. Patterson. Amitosis in the Pigeon's Egg. Anatomischer Anzeiger,
Bd. 32, 1908.

THE

KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 13 — March, 1910.

(Whole Series, Vol. XV. No. l."}.)

CONTENTS:

The Temnospondylous Amphibia and a New Species op Eryops

FROM THE Permian of Oklahoma, . . . Roy L. Moodie.

PUBLISHED BY THE UNIVERSITY,
LAWRENCE, KAN.

October, 1911.

Entered at the post-office in Lawrence as second-class matter.

3310

THE KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. .3.] MARCH, 1910. [vorxv.'™!

THE TEMNOSPONDYLOUS AMPHIBIA AND A NEW
SPECIES OF ERYOPS FROM THE PERMIAN .
OF OKLAHOMA.

BY ROY L. MOODIE.

(Contribution from the Zoological Laboratory, No. 189.)

Plates XLIX to LIV.

OUR knowledge of the fossil Amphibia begins with Dr. J. J.
Scheuczer's discovery of two skeletons in the Miocene of
Oeningen, which he referred to as "Homo diluvii testis
et Theoskopos," and which later Hermann von Meyer desig-
nated "Andrias scheuczeri," thus keeping the same idea in the
generic name as Scheuczer had expressed in his letters and
descriptions. The discovery was made about 1725, so far as I
can ascertain. His letter to Sir Hans Sloane in 1726, which
was translated in the republished volume of the early trans-
actions of the Royal Society of London, is the first definite
mention I find of this discovery in the literature which is ac-
cessible to me.

Exactly one hundred years elapsed from the time of Scheuc-
zer's discovery of the giant salamander skeletons to the time
when Dr. Georg Friedrick Jaeger discovered in the Triassic of
Germany some labyrinthodont remains which he figured and
described in his folio published in 1824 at Stuttgart with the
title "De Ichthyosauri sive Proteosauri Fossilis Specimenibus
in Agro Bollensi in Wurtembergia." He figured the occipital
portion of the skull and teeth of what he, in 1828, called "Mas-
todonsaurus." In 1833 Doctor Jaeger pointed out the syn-
onymy of Salamandroides with Mastodonsaurus. Previous to

(235)

236 KANSAS UNIVERSITY SCIENCE BULLETIN.

Doctor Jaeger's discovery, however, was that of the form
Pygopterus, the history of which is given below.

The literature on fossil Amphibia for the next twenty years
following Jaeger's last mentioned paper deals almost entirely
with the Tertiary Amphibia, although in 1844 and 1845 Her-
mann von Meyer published some interesting papers on the more
ancient amphibians. It is interesting to note that in von
Meyer's "System der fossilen Saurier," 1845, he does not men-
tion a single temnospondylous amphibian.

Our knowledge of the Temnospondylia dates from the year
1847, in which year Dr. August Goldfuss described Archego-
saurus, the first known and for many years the only representa-
tive of this order of amphibians. Doctor Goldfuss gave as the
title of the paper in which he described Archegosanrus "Ueber
das selteste der mit Bestimmtheit erkannten Reptilien." It will
thus be noticed from the literature that the early writers re-
garded the labyrinthodonts and their allies as reptiles. It was
Quenstedt who in 1850 made the first contention that "Die
Mastodonsaurier im gruenen Keuper Sandstein Wurtemburgs
sind Batrachier," and from that time dates the development of
our knowledge of these extinct forms as Amphibia. Three
years after Quenstedt's paper appeared, Vogt upheld the same
contention in his essay "Archegosaurus und all Labyrintho-
donten sind Amphibien nicht Reptilien."

In 1850, after the discovery of the Archegosaurus and its
identification. Doctor Jaeger published an interesting essay en-
titled "Ueber die Uebereinstimmung des Pygopterus liichis Ag.
mit dem Archegosaurus dedienii Goldf." In this essay Jaeger
brings out the interesting facts of the early discovery of the
Archegosaurus. I give the following translation of the first
paragraph of Jaeger's essay: "In the collection of the late
Professor Storr, of Tubingen, which was received in 1817 at the
Imperial Museum of Natural History at Stuttgart, there was
found a skull enclosed in a spherosiderite nodule. This skull
was listed in 1777 as No. 192, in the printed catalogue of the
Pasquay collection (which was the foundation of the Storr col-
lection), without giving the locality, as a fish skull which had
been discovered in a non-fissile gray shale. Agassiz, at whose
disposal the entire collection of fossil fishes was soon after
placed, designated this specimen on a label in his own hand-
writing at first as Aspidorhynchus nov. sp. On a label written

MOODIE: PERMIAN ANPHIBIA. 237

by him later it is designated as Pygopterus lucius. In the first
part of the second volume of the Fossil Fishes (Poissons fos-
siles) , page 10, Agassiz placed this skull in the family Sauroidse
as the second species of the genus Pygopterus, with the short
description : 'Pygojjtei^us lucius Ag., une tete seulement, dont
la machoire superieure est plus allongee. Houille de Saar-
briick.' In the same work, part II, page 78, this specimen is
designated as the fifth species of Pygopterus, with the descrip-
tion, 'une tete avec des dents tres acerees, de la houille de Saar-
briick. L'original se trouve au Musee de Stuttgart.' He re-
gards the same specimen as the third species on page 162 in the
synoptic table of the family Sauroidse arranged in the order of
the formations of the Carboniferous which contained Sauroidse.
It is also given this place in the general table of fossil fishes
which appeared in 1844, page xxxvi."

Jaeger affirmed the supposition of Agassiz that the speci-
men was from Saarbriick, from the similarity of the nodule to
those found at that place. He then proceeded to show that the
skull described by Agassiz as Pygopterus lucius really shows
reptilian characters which Jaeger regarded as intermediate
between those of a crocodile and an Iguana. He then compared
it with the skull of a young crocodile and also with the skulls of
various lizards. He found that the measurements and propor-
tions of the skull compare favorably with those of the crocodile,
and that the teeth have the form and size of those of a newly
born crocodile. The position of the orbit is also identical with
that of the crocodile. He made the important observation that
the nostrils have the lateral position of the lizards. He then
compared the specimen with other known fossil forms, i. e.,
Zygosaurus, RJmiosaurus and the labyrinthodonts, and found
considerable differences. He then made the following interest-
ing observations regarding Archegosaurus: "At the meeting of
naturalists in Aachen in 1847 Goldfuss described the remains in
one of the many spherosiderite nodules, which contained a rep-
tile skull, to which he gave the name ArcJiegosam-us. He estab-
lished three species for the genus, which he described more
fully in a later publication. Only one species of this genus,
Archegosaurus clecheni, was described and figured in the
Neues Jahrbuch fiir Mineralogie, 1847, Tab. VI, with a line
drawing." Jaeger then pointed out in a conclusive manner the
identity of Pygopterus lucius with A)-cJiegosau)-us. He re-

238 KANSAS UNIVERSITY SCIENCE BULLETIN.

garded Archegosauriis as allied to the Crocodilia but having
characters of the Batrachia as were exhibited in the labyrintho-
donts, and on account of the geological position of the fossils
he supposed that Archegosaurus would really belong with the
labyrinthodonts.

Huxley in 1862 described Pholidogaste7' pisciformis from the
Carboniferous of Scotland. In the next year he distributed the
ancient Amphibia into the two groups, the Archegosauria and
the Masto(^onsauria of von Meyer. In the former group he
placed Pholidogaster, to which it manifestly does not belong.
Huxley was himself doubtful as to its location, for he says :
"Archegosaurus, of course, takes its place among the Archego-
sauria ; and Pholidogaster, I suspect, must go with it, though its
vertebra are far better ossified" ; and in a footnote he makes the
interesting statement : "It seems to me probable that the verte-
bral centra of Archegosaurus may really have been osseous
rings, such as are found in embryo frogs and salamanders,"
which shows that he was on the right track but did not quite
comprehend the nature of the vertebrae of Archegosaurus.

Our knowledge for the next few years of our history is
advanced but little, and that only by sundry discussions on
Archegosaurus. In 1866 Owen placed Archegosaurus in his
subclass "Dipnoa," under the order "Ganocephala." In the
same year Haekel retained Owen's classification, except that
he displaced Owen's "Dipnoa" with 'Thractamphibia." Cope,
in 1868, retained essentially the same classification as proposed
by Owen. In 1874 the committee of the British Association
for the Advancement of Science reported on the extinct Am-
phibia, and placed Archegosaurus by itself in group VI, the
Archegosauria.

No further additions were made to our knowledge of the
Temnospondylia until 1875 and 1878, when Cope described
Cricotus from the Carboniferous of Illinois and T rimer orhachis
from the Permian of Texas. In the next year Gaudry described
Actinodon from the Permian of France. In 1882 a decided ad-
vance was made in the separation by Professor Cope of the
rhachitomous forms such as Eryops, Actinodon, Zatrachys, Tri-
merorhachis and other genera, which have subsequently proven
to be synonyms of the genera mentioned, into a distinct group.
Cope proposed the term Rhachitomi for this subordinal group,
and designated two families, the Eryopidse and the Trimeror-

MOODIE: PERMIAN ANPHIBIA. 239

hachidae as its members. Later in the same year Cope de-
scribed Acheloma from Texas. In 1885 Lydekker added other
genera and families to the group, notably Gondwanosaurus,
which he placed in the family Archegosauridse. In 1884 Cope,
and in the following year Fritsch, contributed further to the
knowledge of the Temnospondylia by the description of the
forms Dijjlospondylus and Cricotus from the Permian of Bo-
hemia and Texas. In 1888 Zittel proposed to place all temno-
spondylus amphibians under the suborder Temnospondyli in
two groups — A, those with rhachitomous vertebrse; B, those
with embolomerous vertebrae — for which two groups Cope had
previously, 1885, proposed the terms Rhachitomi and Embo-
lomeri, but had regarded them as coordinate with the group
Stegocephali.

Case, Broili, and others have in recent years given additional
information on the forms previously described and have de-
scribed some new forms. In 1909 Dr. S. W. Williston, in his
essay on ''Trematops milleri from the Permian of Texas," was
enabled to give a restoration of the skeleton of that form. In
the present bulletin the writer will describe an interesting new
embolomerous amphibian under the name Spondylerpeton
spinatum, from the Coal Measures of Mazon Creek, Illinois.
This is the oldest of the known Temnospondylia.

We know nothing of the ancestry of the Temnospondylia.
They occur in the Carboniferous and Permian, and if the Ster-
eospondylia are highly specialized Temnospondylia, as seems
probable, they also occur in the Triassic, but this is yet to be
proven.

The geographical distribution of the Temnospondylia is as
follows : In the Carboniferous and Permian of North America ;
Europe; and Asia, in the Permocarboniferous of India, Gond-
wanosaurus. So far as the writer is aware they occur nowhere
else unless some of the little known forms from South Africa
will, on further search, reveal temnospondylous characters. It
is entirely too soon to arrive at any conclusions as to the paths
of migration of this group. They followed the way which was
open to other land animals, whatever that may have been.

Nor can we say in what region the first of the Temnospondy-
lia lived. Judging from the remains so far discovered their
place of origin must be located in North America. The an-
nouncement of the discovery of the eryopoid form from Pitts-

240 KANSAS UNIVERSITY SCIENCE BULLETIN.

burg, Pa., by Case, and the discovery by the writer of the
cricotoid form in the Mazon Creek fauna, as well as the pres-
ence of Carboniferous Cricotidse in Illinois and Kansas, all
point to an origin of the temnospondylous group at an earlier
date in North America than in any other region of the globe.
The embolomerous forms are by far the older.

The species of fossil Amphibia from the Carboniferous,
Permian and Triassic are so little known that it is difficult to
give a definite list of the species of the Temnospondylia. The
list given below is believed to be approximately accurate. It
does not differ in essential outlines from that given by Zittel
in 1887. The species are distributed in the various groups to
which, at the present time, they seem to belong :

CLASS— AMPHIBIA, Linne, 1758.

Order — Temnospondylia, Zittel, 1887.
Suborder — Rhachitomi, Cope, 1885.

Family — Eryopid^e Cope, 1882.

Eryops megacephalus Cope, 1877. Permian of Texas.
" erythrolithicus Cope, 1878. Permian of Texas.
" ferricolus Cope, 1878. Permian of Texas.
" reticulatus Cope, 1881. Permian of Texas.
" ? platypus Cope, 1877. Carboniferous of Ohio.
" latus Case, 1905. Permian of Texas.
" willistoni Moodie, 1911. Permian of Oklahoma.
Trematops milleri Williston, 1909. Permian of Texas.
Acheloma cumminsi Cope, 1882. Permian of Texas.
Ichthycanthus ohiensis Cope, 1877. Carboniferous of Ohio.
Gondwanosaurus bijoriensis Lydekker. Bijori, India.
Zatrachys apicalis Cope, 1881. Permian of Texas.
" conchigerus Cope, 1896. Permian of Texas.

" crucifer Case, 1905. Permian of Texas.

" micropthalmus Cope, 1896. Permian of Texas.

" serratus Cope, 1878. Pemiian of Texas.

Actinodon frossardi Gaudry, 1880. Permian of France.
" latirostris Jordan. Permian of France.

* (?) Anisodexis enchodus Cope, 1897. Carboniferous of Ohio.

* " imbrecarius Cope, 1882, Permian of Texas.
*Rhytidosteus capensis Owen, 1884. Karoo Beds, South Africa.
(?) *Euchirosaurus rochei Gaudry. Permian of France.

Family — Trimerorhachid^ Cope, 1882.

Trimerorhachis bilobatus Cope, 1883. Permian of Texas.

" conangulus Cope, 1896. Permian of Texas.

" insignis Cope, 1878. Permian of Texas.

" mesops Cope, 1896. Permian of Texas.

" leptorhynchus Case, 1902. Permian of Oklahoma.

Cricotillus brachydens Case. Permian of Oklahoma.
Archegosaurus austricus Makowsky, 1876.

" decheni Goldfuss, 1847. Permian of Germany.

" latifrons Gein. & Deich. Permian of Saxony.

" ornatus Woodward, 1905. Permian of India,

MOODIE: PERMIAN ANPHIBIA. 241

Platypodosaurus ricardi Twelvetrees, 1880. Permian of Russia.
" stiickenbergi Trautschold. Zechstein of Russia.

Cochleosaurus bohemicus Fritpch, 1876. Permian of Bohemia.
Gaudrya latistoma Fritsch, 1885. Permian of Bohemia.
Chelyosaurus vranyi Fritsch, 1878. Permian of Bohemia.
*Sphenosaurus sternbergi Fitzinger, 1840. Permian of Bohemia.
Sparagmites lacertinus Fritsch, 1885. Permian of Bohemia.
*Sclerocephalus hauseri Goldfuss. Permian of Germany.
*Sclerocephalus credneri Fritsch. Permian of Bohemia.
*Sclerosaurus bavaricus Branco. Rothl. Ohmbach.
*Melosaurus uralensis H. von Meyer. Kupfersandstein.
*Osteophorus roemeri H. von Meyer. Rothl. Schliessen.
*Zygosaurus lucius Eichwald. Permian of Russia.

Gonioglyptus huxleyi Lydekker from the Gondwana beds of
India also possibly belongs in this family.

So many genera and species of extinct Amphibia are based
on fragments of crania or limbs that an exact diagnosis of the
forms is impossible. The species marked with a star (*) are
uncertain as to position. Many of the species listed under the
Trimerorhachidse are placed in this family on the authority of
Lydekker, who included them in his family Archegosauridse,
which was preoccupied by Trimerorhachidse. Doctor Williston
regards Trematops milleri Will, and Acheloma cumminsi Cope
as representatives of a family distinct from the Eryopidse for
which in 1910 he proposed the name Trematopsidse. He also
regards the species of Zatrachys as entitled to family rank in
the Zatrachydidse, Williston, 1910.

Euchirosaurus of Gaudry is probably a reptile.

Suborder — Embolomeri, Cope, 1881.
Family — Cricotid^ Cope, 1884.

Cricotus gibsoni Cope, 1877. Carboniferous of Illinois.

" crassidiscus Cope, 1884. Permian of Texas.

" heteroclitus Cope, 1875. Carboniferous of Illinois.

" hypantricus Cope, 1884. Permian of Texas.

" sp. indet. Case. Carboniferous of Illinois.
Diplospondylus punctatum Fritsch. Permian of Bohemia.
Spondylerpeton spinatum Moodie, 1910. Carboniferous of Illinois.
? *Discosaurus permianus Credner. Permian of Saxony.

Family — Dissorophid^ Williston, 1910.

Dissorophus multicinctus Cope, 1895. Permian of Texas.

" mimeticus Cope, 1896. Permian of Texas.

" testudineus Cope, 1896. Permian of Texas.

Cacops aspidephorus Will. 1910. Permian of Texas.
Aspidosaurus chiton Broili, 1904. Permian of Texas.

From the foregoing diagnosis is will be seen that about 55
species of the 350 which have been assigned to the fossil
Amphibia of the world can be located in the order Temno-

242 KANSAS UNIVERSITY SCIENCE BULLETIN.

spondylia. Without doubt some of these species will prove in-
valid and there will be others already described which will in
time prove to belong to this group, but the above statements
represent an approximation of our present knowledge.

Since the present essay is to deal especially with a new mem-
ber of the genus E^ijojJs, only those groups will be here de-
fined to which Eryops and its allies pertain.

Order — Temnospondylia, Zittel, 1887.

Terrestrial or semiaquatic vertebrata ; skull bones pitted and
grooved; lateral line canals present; pineal foramen some-
times absent; sclerotic plates present; vertebrae rachitomous
or embolomerous ; notochord usually persistent ; one or two
sacral vertebra; tail present, short or long; limbs and girdles
all well developed ; pectoral and pelvic girdles composed of the
usual stegocephalan elements; an osseous pubis present;
cleithrum present on the scapula in some forms; limb bones
well developed and bones of forearm and leg separate; carpus
and tarsus osseous, 12 elements present in tarsus of one form,
11 carpals; five digits in hand and foot; phalangeal formula
2, 3, 3, 4, 2 (?) for the hand and 2-3-3-3-2 (?) for the foot;
venter covered with an armature of osseous scutes, sometimes
overlapping; skin of back bare or armored; ribs heavy, double-
headed, curved, and either moderately long or short. Body
short and heavy. As compared to skull, about two to one.
Temporal foramina present in a few forms.

Range. Pennsylvanian to (Triassic) Upper Permian.

Distribution. North America — Illinois, Kansas, Oklahoma
and Texas. Europe — Germany, Bohemia, France. Asia —
India.

The suborder Rhachitomi is characterized solely by the na-
ture of the vertebral column, which is cut into triangular seg-
ments.

Family — Eryopid^ Cope, 1882.

Large terrestrial or amphibious vertebrata; skull bones
deeply marked with pits and grooves which at times take the
form of lateral line canals ; carpus and tarsus osseous ; pubis
osseous ; pentadactyl fore and hind limbs ; orbits in the typical
genus located far back on the skull and near the median line;
cleithrum present on the scapula ; vertebrae rachitomous, the
intercentrum supporting the arch in the dorsal region; para-

MOODIE: PERMIAN ANPHIBIA. 243

sphenoid well developed or reduced ; teeth on pterygoids, pala-
tines, prevomers and parasphenoid.

Range. Upper Pennsylvanian to Permian.

Distribution. America, Europe, Asia,

Genus — Eryops Cope, 1877.

The following definition of the genus is given by Doctor
Branson : "Skull long, comparatively narrow ; proportion of
length to breadth about nine to seven. Roof bones coarsely
sculptured posteriorly, finely sculptured anteriorly. Nasals
and premaxillse very large; f rentals excluded from orbits by
junction of pre- and post-frontals. Pterygoids not meeting in
the median hne; parasphenoid dagger-shaped, tapering grad-
ually to a point just in front of the palatine foramina; pre-
vomers large. Orbits subcircular, situated in the posterior half
of the skull ; nares subovate, remote, at a considerable distance
from the tip of the skull. Many minute denticles on pterygoids,
palatines, prevomers and parasphenoid. Teeth circular in
cross section, strongly ribbed near base, dentine strongly in-
folded. Three large teeth on each palatine. Mandible without
postcotyloid process. Vertebrae rachitomous. Ribs double-
headed. Pelvic bones coalesced."

There have been up to the present five species of Eryops de-
scribed, with a doubtful sixth, ? Eryops platypus Cope, from
the Carboniferous of Ohio.

In 1889 Lydekker described a species of Eryops, E. africanus,
from the Karoo beds of South Africa. Broom has subse-
quently shown that this species does not belong in this genus
and has placed that species with two others in his genus
Rhinesuchus, in which he includes R. (Macromerion) gumbeli
(v. Ammon), R. (Eryops) africanus (Lydekker), and R.
whaitsi Broom.

An additional species of Eryops is represented by remains
in the collection of the University of Kansas and it is here
described as new.

Eryops willistoni, new species.

Dr. S. W. Williston in 1899 described and partially figured
m the Kansas University Quarterly portions of a skeleton of
a species of Eryops, which he tentatively referred to the
species Eryops megacephalus Cope. He has several times
expressed to me the opinion that the form is distinct from
E' megacephalus, and a recent careful study of the ma-

244 KANSAS UNIVERSITY SCIENCE BULLETIN.

terial has led me to regard the remains as a new species,
which may be very appropriately named Enjops ivillistoni as
a partial expression of gratitude to my honored preceptor in
paleontology. Since, also, several interesting points in
anatomy and morphology have come up for consideration the
following discussion is presented in the hope of adding some
knowledge on this peculiar Permian genus.

The material which represents the present species comprises
the following portions of the skeleton : Portions of the skull ;
several teeth and a nearly entire right mandible; the scapula-
coracoid of both sides, incomplete ; the right clavicle ; portion
of the interclavicle ; right humerus, radius, ulna and a
phalange ; about a dozen vertebrae and the left sacral rib.

There are various fragments of the skull preserved. On one
of these fragments is observed a portion of the right orbit and
a part of the prefontal and frontal bones clearly marked off
by sutures. The present form differs in this respect from the
known specimens of Erijops megacephahis, in which the sutures
remained unknown until they were discovered by Doctor
Branson, who was able to trace them with the aid of a lens.
On the present specimen the sutures stand out with almost
startling distinctness. They do not have the sharply zigzag
form assumed in E. megacephalus and other of the larger
Amphibia, but the sutures occur in bold curves. I detect a
portion of the supraorbital lateral line canal on the 'fragment
containing the orbit. The sculpture of the cranial bones in the
present form is quite different from that of E. megacephalus.
It is much more rugose and the rugosities have a larger form.
This holds true for the mandible also (plate XLIX, fig. 1), and
it is taken as one of the specific characters.

There are preserved in the collection also a single complete
tooth and fragments of others. The complete tooth (plate
XLIX, fig. 2) is of the typical labyrinthodont form. It is oval
in cross section with sides flattened. It arises from a tumid
base of cancellated bone as in the mosasaurs. It is recurved
and sharply pointed.

In 1899 Ludwig Strickler (Paleontographica Bd. XLVI, p.
85, plates XI, XII) investigated the minute structure of the
teeth of Eryops megacephalus.

The mandible of the present species is preserved in a much
broken condition, but its form is nearly complete. The form
and ornamentation of the posterior portion were well shown in

MOODIE: PERMIAN ANPHIBIA. 245

Williston's figure (Kan. Univ. Quart., Ser. A, vol. VIII, pi.
XXX, fig. 1), which is reproduced here with slight changes.
The sharp angulation and the posterior position of the angle is
a specific character which serves to distinguish this species
from Eryops megacephalus Cope, to which the present form is
most nearly allied.

The jaw (plate XLIX, fig. 1), is heavy and elongate. The
portion posterior to the coronoid angle is an acute triangle
which ends in the articular. Only a portion of the suture
bounding the articular can be distinguished. This is indicated
in the figure. The coronoid suture is evident for its entire
course. The suture starts from the articular suture, and run-
ning directly forwards turns upward just before reaching the
teeth, which do not occur on the coronoid. This element is a
thin bone superiorly but more thickened below. The sutures
separating the angular, surangular and dentary cannot be de-
termined. The angular and surangular contain, on their exter-
nal surface, the peculiar nodose ornamentations which help to
characterize the present species.

The sculpture on these elements consists of scrobiculate
knobs some 5 to 7 mm. in diameter. This ornamentation ends
apparently at the dentary, since the tooth-bearing element,
which is quite large, is ornamented only with coarse elevations
and pits.

The operculo-mandibular lateral line canal can be traced as a
depression beginning on the articular and running forward as
far as the bone is clearly preserved. It occurs as a shallow
groove interrupted by the nodose and scrobiculate sculpturing
of the mandibular elements. The internal surface of the man-
dible is not preserved in sufficient detail for description.

Complete teeth are lacking from the mandible in its present
condition. Doctor Williston figured tvv^o of them as complete,
and it is quite probable that they have been broken off since he
studied the specimen, because the material is excessively
fragile. The hollow character holds for all of the tooth bases
which are preserved on the mandible. An additional portion
of the mandible is shown in figure 5, plate XLIX. Its char-
acters do not differ from those described above.

In regard to the hollow nature of the teeth. Tomes has the
following explanation : "The laminse of pulp, with their several
systems of dentinal tubes, instead of passing out in straight

246 KANSAS UNIVERSITY SCIENCE BULLETIN.

lines like the spokes of a wheel, pursue a tortuous course as
they run from the central small pulp chamber toward the sur-
face. Not only do they undulate, but they give off lateral proc-
esses, and at their terminations near the surface of the tooth
the thin laminae of pulp (so thin that the radiating pulp cham-
bers are mere fissures) become dilated; so that on section cir-
cular canals are seen at these points, as is also the case at the
points where subsidiary processes branch off." (Dental An-
atomy, 5th edition, 1898, p. 65.)

Measurements of the mandible of Eryops willistoni:

Greatest length 48 cm.

Length of posterior portion (fig. 1) 33

Greatest width 10

Width of dentary midway from coronoid process 6.5

Width across articular 2.6

Length of tooth 2 ,

Thickness of same at base 5

The right arm of this species is preserved in an incomplete
condition, as may be seen by reference to plate LII. The form,
of the humerus is well shown in the figures, plate LIII, fig. 7,
and plate LII. It suggests to a certain extent the humerus of
a mosasaur. The present element does not have the bizarre
appearance which exists with the specimen of that element of
Eryops megacephalus figured by Professor Cope in 1890
(Trans. Amer. Philos. Soc, vol. XVI, p. 367, fig. 3). The large
projections are wanting entirely from one side and on the other
they are not so highly developed, and yet the form of the bone
is such that it can be nothing else save a humerus. It is too
rugged for a femur.

The specimen is abraded to some extent, so that the exact
form of the element cannot be determined. It may have had
the form suggested by the broken lines in the figures. The
humerus is that of the right side. Viewed from in front the
element is seen to have a peculiar form (plate LIII, fig. 7).
The lower portion of the inner surface is distinctly concave.
The inner side of the bone is comparatively smooth. There is
a low ridge which runs obliquely across the antero-superior
angle of the bone. There is a distinct twist in the inner sur-
face, which is continued to such an extent that the inner sur-
face is carried around almost parallel with a portion of the
outer surface. It then takes another and inward curve and
ends in a projection.

One of the projections from the outer surface of the humerus

MOODIE: PERMIAN ANPHIBIA. 247

is in the form of a hook and strongly suggests the hooklike
process which is developed in the quadrate of the mosasaurs.
This hook and the other elevations on the edge of the bone
without doubt served as points of attachment of the deltoid and
pectoralis muscles, which were undoubtedly well developed in
the species of Eryops.

The outer surface of the bone is exceedingly rough and there
are no less than six distinct elevations on this surface. One
of these elevations is a ridge which runs at somewhat of an
angle across the posterior edge. There is an indication of a
foramen opposite the pit which occurs on one side of the bone,
which at first was thought to represent the entepicondylar for-
amen. This was such an important matter that the bone was
cut through at this point by Mr. Martin and fully demonstrated
the total absence of this foramen in the element although there
is quite a distinct pit at this point. Cope described such a fora-
men in Acheloma, but Case, who has recently examined the
type, says the foramen is an accidental opening. Gaudry like-
wise described such an opening in Actinodo7i.

Measurements of the humerus :

Length of specimen as preserved 96 mm.

Estimated length along same line 110

Width at middle of shaft 50

Width of top of humerus 90

Width at lower end 72

Thickness of head as preserved 23

Diameter of entepicondylar pit 3

The element which is here regarded as a radius is very sim-
ilar in form to that described by Cope for Eryops mega-
cephalus. Its form is well shown in the figure. The lower end
is somewhat larger than the upper, which is abraded and
broken. Viewed from the lower surface a cross section of the
bone shows a semicircular form. It is hour-glass shaped in its
entire length, with a shallow groove near the lower end. Both
ends of the bone near the articular surfaces are roughened by
vascular pits for muscular attachment or for the union of the
cartilaginous tips.

Measurement of radius:

Actual length of specimen 91 mm.

Estimated length of specimen 95

Width of upper end (estimated) 40

Width at middle of shaft 24

Width at lower end 60

Thickness of base 32

248 KANSAS UNIVERSITY SCIENCE BULLETIN.

The element here figured as an ulna (plate LII, U) is doubt-
fully referred to the arm, since the bones were found disasso-
ciated. It may belong to the leg. Its form is specifically difl"er-
ent from that which Cope figured as the ulna of Eryops mega-
cephalus, although its form is, in general, the same. It does
not depart widely, in point of form, from that of the radius,
which we should expect to be the case in a generalized verte-
brate. It is smaller, somewhat shorter, and the groove on the
ventral ( ?) surface is lacking. The radius and ulna are both
peculiarly amphibian in the way they have weathered. The
ends have assumed the concave form so characteristic of the
Branchiosauria and the modern Amphibia, indicating a higher
development of the perichondrium than of the endochondrium.

Measurements of the ulna :

Actual length of specimen 80 mm.

Estimated length of specimen 93

Width of shaft at middle 22

Width of upper end (estimated) 47

Width of lower end (estimated) 40

Associated with the limb bones there is a portion of a pha-
lange of the hand ( ?) of the specimen. There are no carpals
preserved. The element figured (plate LIII, figs. 4 and 5) is
probably a metacarpal. Its form is very closely similar to that
figured by Cope for the metacarpal element of Eryops mega-
cephalus. Its upper end is very broad, with recurving edges
and with rugosities for muscular attachment. Its form is
crescent shaped and concave. The shaft narrows abruptly
from the upper end to the middle and then widens for the lower
end, which is lost.

Measurements of the metacarpal :

Actual length of specimen 23 mm.

Estimated length of bone 27

Width of upper end 22

Thickness of upper end v 8

The following description of the coraco-scapula (plates
L, LI) is taken from Doctor Williston's paper. The descrip-
tion of the cleithrum is appended :

"The bone is elongated, and nowhere very thick or massive.
The distal part of the scapula is much thinned and considerably
expanded; the immediate margin here, however, is wanting,
so that the precise outlines cannot be given.

"The position of the two bones must have been very oblique,
as is evident from the position of the glenoid cavity. The pos-

MOODIE: PERMIAN ANPHIBIA. 249

tero-inferior border is moderately thickened, and rounded;
gently concave along the proximal part and convex distally.
The antero-superior border is much thinner than the opposite
one, and is concave throughout the extent of the shaft, except
distally, where it is coossified with the procoracoid. The union
of these two bones is very close in this region, the sutural line
being distinguished with difficulty if at all. The procoriicoid
is narrower and more thickened below, reaching to the lower
part of the conjoined bone, and lying in close apposition though
not suturally united. Both the lateral surfaces of the scapula
are nearly flat. A little proximad to the narrowest part of the
shaft the bone is much thickened by a stout ridge on the inner
side, which includes between it and the remainder of the bone,
just back of or above the cotylus, a large, elongated foramen,
both of whose orifices can be seen from the inner side of the
bone only.

"The glenoid surface is elongated and deeply concave in its
long diameter. The scapular portion is much smaller than the
coracoid, and is partly separated from it by a constriction ;
this surface is nearly flat and is placed at right angles to the
plane of the coracoid surface, looking directly downward when
the bone is lying horizontally. The rest of the cotylus is mod-
erately concave and looks outward and somewhat backward
and upward with the bone in its former position. Just how
much of the conjoined bone is formed of each element it is im-
possible to say since the union is so close that no trace of a
junction is perceptible. Evidently, however, the coracoid
forms only a small part of the whole bone. Its anterior or in-
ferior border is gently convex, the posterior one slightly
concave.

"The anterior end of the superior border of the scapula is
gently convex, as is also the broad external surface. On the
inner side the foramen described above opens into a deep,
elongated, boat-shaped cavity at its distal end, the cavity
formed by the upper border of the ridge described. In the
lower or anterior end of the cavity there are apparently two
smaller foramina, the external orifices of which are just within
the anterior margin of the scapular face of the glenoid surface.
The proximal surface of the scapula is concave; that of the
coracoid, if the bone is limited by the ridge spoken of, is for the
most part gently convex and lies in a more mesial plane."

a-Univ Sci. Bull.. Vol. V, No. 13.

250 KANSAS UNIVERSITY SCIENCE BULLETIN.

The scapula is evidently that of the right side, the same as
the humerus described below.

On the antero-superior corner of the scapula is a part of a
peculiar element — the cleithrum. Its occurrence in the genus
Eryops has been noted by Case in two species. In E. latus the
cleithrum takes a most peculiar form. The form assumed by
the element in the present species is impossible to determine,
since the greater part of it has weathered away.

The cleithrum is, apparently, simply laid on the upper edge
of the scapula, the union being hardly a sutural one. In the
present specimen it is represented only by the lower part, but
it could not have had the immense expanse superiorly as Case
and Williston have figured for the cleithrum in Eryops latus
Case. It was probably quite slender, as it is in the dicynodont
reptiles. The cleithrum is a thin plate of bone with a ridge
running along the interior border. The edge of the scapula
is depressed to receive the element.

The characters of the scapula which are at variance with the
scapulse of the known species are the narrowness of the shaft,
the location and size of the cleithrum, and the position of the
cotylus.

In regard to the three foramina seen and described in the
scapula Williston has recently proposed the terms supra-, post-
and infra-glenoid or supra-coracoid to designate these open-
ings. The foramina occur in all known Temnospondylia, as
well as in the Cotylosauria (2) and in the Pelycosauria (2).
The writer has recently described a branchiosaurian from the
Pennsylvanian of Mazon Creek in which three foramina are
visible in the scapula. This was thought to be of interest in
connection with the Temnospondylia, but investigation showed
that in the temnospondyles the foramina are in no case to be
regarded as occurring strictly in the scapula, but rather as be-
longing to the coracoid which is united with the scapula.

An interesting question of homologies, or it may be parallel
development, arises in comparing the present scapula-coracoid
with the clavicular apparatus of Portheus.

Measurements of scapula-coracoid :

Actual length of specimen 33 . 7 cm.

Estimated length of element 36

Width of upper end of scapula 15.5

Thickness at upper end 3 mm.

Width at middle of shaft 6.6 cm.

Width across coracoid portion 17

MOODIE: PERMIAN ANPHIBIA. 251

Length of cotylus 7.2

Width of cotylus 3.8

Diametei" of infraglenoid foramen 5 mm.

Thickness of bone at lower edge 7

Thickness of bone through cotylus 4.3 cm.

Length of portion of cleithrum preserved 63 mm.

Greatest width of cleithrum 22

The clavicle of the right side is preserved almost completely
(plate LIII, figs. 1 and 2). The element was figured by
Doctor Williston but not described. It is strongly curved and
spatulate. Its outer surface is ornamented with longitudinal
grooves, which arise near the center of the bone and run each
way, thus indicating its ossific center. The inner surface of
the bone is smooth excepting for two vascular pits near the
center. A prominent ridge runs along the dorsal inner edge
of the bone.

Measurements of the clavicle :

Length along curve 24.7 cm.

Length across arc 20

Least width 1.3

Greatest width 4.6

Greatest thickness 1

Least thickness 2 mm.

There is a fragment of what I regard as the posterior
projection of the interclavicle. If it is such it is either a
pathological specimen or there were two projections from the
posterior edge of the interclavicle, for the fragment is highly
asymmetric. The spine, for such it is, has an irregular nodose
surface as if for the attachment of cartilage. It is long, slender
and somewhat flattened. (Plate LIII, fig. 3.)

Measurements of spine of interclavicle :

Length of specimen 13.7 cm.

Greatest width 2.2

Least width 75

There are remains of about a dozen vertebrae preserved.
Portions of three of these are figured in plate LIV, figures 1
and 2. They all have the characteristic rachitomous form as-
sumed by the other species of Eryops. The divisions between
the pleurocentra, hypocentra and neurocentra are well marked.
The zygapophyses are well formed. The upper part of the
neural spines is much thickened, as is the case often in speci-
mens of Eryops megacephalus Cope. According to Broili the
spines may also be bifurcate, as Cope has described for E.
ey^ythrolithicus. The enlarged spine was undoubtedly for the
attachment of muscles to sustain the enormous head, and

252 KANSAS UNIVERSITY SCIENCE BULLETIN.

spines showing different characters might very easily occur

in various portions of the vertebral column.

Measurements of vertebrse :

Length of centrum 2.5 cm.

W^idth of centrum 3.2

Height of vertebras to tip of spine 13.4

Thickness of top of spine 3

Broili (Monatsb. d. Deutsch, Geol. Gesell., Bd. 60, Texttafel
p. 236, fig. 2) has figured in Eryops megacephalus a peculiar
element which he has identified as the sacral rib, having found
it in place with a sacral vertebra. There is no doubt that the
present element (plate LIV, figs. 3 and 4) is the same as he
has figured. Its interpretation as a sacral rib is somewhat
surprising on account of the extreme thinness of the bone. It
thins out to less than a millimeter on one edge. Its location
any other place in the skeleton is, however, doubtful.

The element has a broad base as if for attachment suturally
with the vertebral transverse process. The base of the speci-
men has been broken transversely and only a portion of the
suture remains. The bone is flat immediately from the base
and is greatly expanded and thinned distally. It is, for the
most part, smooth, though there is a small roughened place on
the dorsal (?) surface. The ventral (?) surface is marked by
a depression which runs the entire length of the bone. The
distal end is lost.

Measurements of sacral rib :

Actual length of specimen 90 mm.

Width at distal end 62

V^idth at middle of blade 32

Width at inner end 55

Eryops ivillistoni differs from E. ferricolus Cope in the
absence of a raised rim to the orbits. From E. erythrolithicus
Cope it differs in the non-bifurcation of the neural spines.
This last is not a well-known species and may be synonymous
with E. megacephalus. E. reticulatus is a small species, from
which E. willistoni differs in the character of the cranial sculp-
ture. It is distinct from E. latus Case on account of the di-
mensions of the scapula and the cleithrum. E. megacephalus
also shows characters which are divergent from those of the
present species. These are the shape and structure of the
mandible and its ornamentation, the form of the humerus and

MOODIE: PERMIAN ANPHIBIA. 253

the shape and structure of the scapula-coracoid, as well as
other structures which are characteristic.

The material on which the species Eryops willistoni is based
is from the Permian of Oklahoma. It was collected by Prof.
C. N. Gould in 1896, in the red sandstone southwest of Black-
well, Okla. The specimen is No. 348 of the Kansas University
Museum.

I am indebted to Mr. G. Dallas Hanna for all of the draw-
ings illustrating the present essay, except that on plate XLIX
figure 1, which is from the pen of Mr. Sydney Prentice, at
present with the Carnegie Museum.

3-Univ. Sci. Bull.. Vol. V, No. 13.

THE

KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 14 — March, 1910.

(Whole Series, Vol. XV. No. 14.)

CONTENTS:

An Armored Dinosaur from the Upper Cretaceous of Wyoming,

Roy L. Moodie.

'PUBLISHED BY THE UNIVERSITY,
LAWRENCE, KAN.

October, 1911.

Entered at the post-office in Lawrence as second-class matter.

3310

THE KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 14 ] MARCH, 1910. [vo"!xv,'.v" u

AN ARMORED DINOSAUR FROM THE CRETACEOUS

OF WYOMING.

BY ROY L. MOODIE.

(Contribution from the Zoological Laboratory, No. 190.)

Plates LV to LIX, and one text figure.

IN Science for October 20, 1905, Dr. S. W. Williston (1) pub-
lished a brief account of a remarkable armored dinosaur
from the upper Cretaceous of Wyoming. He likewise gave
a short sketch of the horizon from which the specimen was
obtained, naming the horizon Hailey shales, and suggesting
the name Stegopelta landerensis for the dinosaur.

During the summer of 1906, while searching these beds for
plesiosaur remains for the Carnegie Museum {2) the writer
followed the horizon eastward and northward to the southern
end of the Big Horn mountains. The beds increase in thick-
ness towards the east and the lithological character becomes
different, changing from a soft fine shale in the west to a hard
sandy rock in the Big Horn region. The Hailey shales lie just
above and comformable on the Mowry beds, which Darton {3)
describes as follows :

"The stratigraphic succession of the formation [Colorado]
is uniform throughout. The lowest beds are several hundred
feet of dark-colored shale, usually containing toward the base
a deposit of sandstone from 2 to 20 feet thick. This is capped
by the Mowry beds, consisting of from 100 to 150 feet of hard
shales and thin-bedded, fine-grained sandstone of a dark gray
color, which on exposure weathers to a characteristic light
gray. These rocks are especially characterized by large num-
bers of fish scales in nearly every bed. They are often so

(257)

258

KANSAS UNIVERSITY SCIENCE BULLETIN.

abundant as to number from five to ten on a surface of six
inches square. Owing to their hardness the beds give rise to
round-topped ridges of considerable prominence."

Fig. 1. A typical exposure of the Hailey shales on Connant creek, Wyo-
ming. The Mowry beds may be seen on the left in the background.

In those parts of Wyoming where I worked, the Mowry beds
stand out as prominent ridges of hard whitish shales which
can be readily recognized from a long distance, and on that
account serve as excellent landmarks for the identification of a
region where the Hailey shales may occur. In thickness the
Hailey shales vary from a few feet north of Lander, Wyo., to
something over 200 feet in the Big Horn region. "Near Soap
Creek the dark shales overlying the Mowry beds are about 200
feet thick," etc. (6). In most of their extent the Hailey beds
have numerous brown sandy clay concretions which very often
contain vertebrate remains. These concretions vary in size
from a few inches to many feet in diameter. In the Big Horn
region the concretions are stratified into distinct layers, but
such is not the case in other places where the horizon was
examined.

So far as is at present known the Hailey shales outcrop along
the eastern flank of the Wind River mountains, in the Shoshone
anticline; in the anticline along Beaver creek near Hailey,
Wyo., the type locality; in an anticline north and west of the

moodie: armored cretaceous dinosaur. 259

Granite mountains, on Connant creek ; in a small anticline west
of the Rattlesnake mountains ; along the northern and eastern
flanks of the Rattlesnake mountains. There are extensive ex-
posures on the eastern slope of the Big Horns, and I do not
doubt that they occur also to the west of these mountains. In
text-figure 1 is given a view of a typical exposure of the
Hailey shales, on Connant creek, north of the Granite moun-
tains, Wyoming. On the left of the photograph the Mowry
beds stand out in the characteristic ridge.

The vertebrate fauna of the Hailey shales still remains to be
accurately determined, but so far as known it is represented
bj^ large and small fishes, amphicoelian crocodiles with hollow
limb bones, turtles with completely ossified carapaces and
plastra like Toxochelys and plesiosaurs of two types. The re-
mains of plesiosaurs are in places exceedingly abundant, evi-
dences of no less than fifty-five individuals having been dis-
covered during one summer. Occurring in these deposits, also,
have been discovered the remains of three dinosaurs. The re-
mains described below represent one of these animals. The
other two are known in one case from a single worn dorsal
vertebra found in one of the massive shell-concretions con-
sisting of myriads of gastropod and lamellibranch shells to-
gether with plesiosaur teeth and vertebrae, about ten miles
north of where the Stegopelta remains were exhumed ; and in
the other case from three caudal vertebrae discovered in the
same beds nearly a hundred miles to the east, near the western
base of the Rattlesnake mountains.

The Stegopelta remains herein described were once encased
in one of the brown sandy clay concretions so abundant in this
formation, but many years ago it was exposed and by the
weathering processes of a long space of time had become re-
duced to hundreds of fragments. These fragments had washed
and rolled down the slope and had been trodden in the mud
by cattle so that when first discovered it was discarded as a
worthless plesiosaur specimen. When Doctor Williston, how-
ever, in cleaning up some of the fragments came across some
typical stegosaurian dinosaur teeth, the whole aspect of the
matter was changed and every fragment was carefully col-
lected. As boxed the specimen weighed only 450 pounds, of
which about two-thirds was bone. It has been a long and
tedious task to fit together the scattered and disassociated
fragments, but through the combined eff'orts of Doctor Willis-

260 KANSAS UNIVERSITY SCIENCE BULLETIN.

ton and myself a number of elements of the skeleton have been
reconstructed.

The specimen as it now stands consists of the following
parts: Fragments of the upper and lower jaws with a dozen
or more teeth, most of which are fragmentary; seven frag-
mentary dorsal vertebrae; portions of the sacrum; two caudal
veitebrse; many fragments of the ribs; a complete ulna be-
longing to the left side ; portions of the right ulna ; portions of
the radii ; the left ilium nearly entire ; a part of the right ilium,
showing the acetabulum ; the right pubis nearly entire ; a por-
tion of the left pubis; the lower portion of the right tibia; the
upper and lower ends of the right fibula ; many large fragments
of the metatarsals ; nearly a hundred small bony scutes ; several
large bony plates, girdles and bony spines representing va-
rious portions of the dermal armor; together with a great
quantity of waterworn fragments which it is impossible to
determine.

The remains described as Stegopelta landerensis indicate a
stegosaurian dinosaur about two-thirds the size of Stegosauiiis
ungidatus Marsh. The head was small, as is indicated by the
size of the teeth. The fore limbs were not relatively so large
as in Stegosaurus. The entire pelvic region was covered with a
dorsal shield of rather thick, scrobiculate plates with elevated
centers which are asuturally united over the pelvis. On each
plate there is found, either on one side or in the central emi-
nence, a pit, which possibly supported a more or less elongate
horny spine. Anterior to the pelvic shield there were many
dermal plates and scutes of various sizes scattered over the
dorsal region of the body. The anterior portion of the animal
appears not to have been armed. Back of the pelvis there were
massive dermal plates, huge bony spines and a bony girdle
probably encircling the tail in some such manner as in the
glyptodonts. The neural spines of the caudal vertebrae were
probably each surmounted by an elongate bony plate which
quite possibly bore a horny excrescence of some kind. While
closely similar in many respects to Polacanthus, Stcgopelta
shows wide structural differences from that form, although it
must have had much the .same appearance as Nopsca (5) has
figured in Polacanthus. The present dinosaur, in life, may
have stood nearly seven feet high at the hips and attained a
length of sixteen to eighteen feet. Armed as it was with horns,

moodie: armored cretaceous dinosaur. 261

spines and excrescences it must have been a bizarre creature,
even in those Cretaceous days.

The teeth (plate LVI, figs. 4 and 5) of Stegopelta are typi-
cally stegosaurian. There are a dozen of them in more or less
perfect state of preservation. They are all of practically the
same size and character, showing but little variation among
themselves. The crowns of the teeth, which measure 7 mm. in
breadth, are subcompressed and narrowly palmate with eight
radiating costse, all of which end at the tip in sharp points.
The roots of the teeth measure 15 mm. from the base of the
crown to the tip of the fang. In point of size and number of
costse the present teeth are so closely similar to those described
by Leidy (5) as a supposed lacertilian form, Paleoscincus,
that had the teeth only been found they would undoubtedly
have been referred to that genus. It is possible that Stegopelta
may eventually prove to be a Paleoscincus form. This, how-
ever, is hardly probable, on account of the great interval of
time separating the strata which have yielded the two speci-
mens, PaleoscincKS occurring as it does near the top of the Fort
Pierre Cretaceous and Stegopelta near the top of the Fort Ben-
ton. Were the form one of a stable group this might not be
sound reasoning, but in a group of animals which evolved so
rapidly as did the dinosaurs one would hardly expect a genus to
continue for so long a time. At all events, the name Stegopelta
may be retained until more evidence is forthcoming concerning
Paleoscincus. If, as Hatcher believed (23), Lambe's Stereo-
cephalus is a Paleoscincus form, then Stegopelta is a good
genus, since it shows wide structural differences from the form
which Lambe has described. The tooth which Lambe (7) has
figured on page 57 certainly resembles those of Paleoscincus,
and if the tooth really is to be associated with the remains
described as Stereo cephalus a question might arise as to
whether these remains were not in reality portions of the skele-
ton of Paleoscincus as Hatcher intimated. Lambe {8) says
the tooth provisionally associated with the Stereocephalus
remains "differs from those of the Red Deer river district, re-
ferred to the two species of Paleoscincus, and is about twice as
large as those of P. costatus." Just how it differs he does not
say, and one is at a loss in attempting to separate the forms
from his figures. Size alone is usually no safe criterion for the
separation of animal forms, and more particularly genera. I

262 KANSAS UNIVERSITY SCIENCE BULLETIN.

can but agree with Hatcher that the tooth figured as Stereo-
cephalus resembles the teeth of Paleoscincus.

The vertebral cohimn is represented by posterior dorsal ver-
tebrae, portions of the sacrum and two caudal vertebrae. The
remains are more or less fragmentary, but enough is preserved
to give a fairly accurate idea of this portion of the animal's
anatomy, although it is impossible to determine the vertebral
formula.

The dorsal vertehrse (plate LVIII, figs. 5 and 6) obtained are
twelve in number, for the most part imperfectly preserved.
They are cylindroid in form, very slightly amphicoelous, nearly
amphiplatyan, and the arches are elevated as in Stegosaurus
and Polacanthus. The centra are slightly excavated below the
base of the neural arch as in Scelidosaurus. The neural canal
is small and is wider in the ventral than in the dorsal portion.
The centra measure 60 mm. in width by 70 mm. in length.

The sacrum (plate LIX, figs. 6 and 7) as preserved consists
of two sections. One includes three firmly coossified vertebrae
of the posterior end and the other a part of the anterior end
representing two vertebrae. The anterior section measures
120 mm. in length by 70 mm. in width at the widest part.
The fragment is very smooth and of a dense bony texture.
The neural canal is broad posteriorly but narrows somewhat
anteriorly. In plate LIX, figure 2, is represented a cross
section through this fragment to show the sacral enlargement
at the posterior end. The portion representing the posterior
end of the sacrum (plate LIX, figure 7) measures 203 mm. in
length by 64 mm. in width at the caudal end. The texture of
this part is very fine and compact externally, but internally
there is a tendency to a cancellated structure. An interesting
characteristic of this element of the skeleton is the high degree
of coossification of the various elements composing it. There
is not the slightest indication of a suture where the different
vertebrae have become joined so that the sacral centra are con-
stituted in a solid bony rod tapering somewhat at the anterior
end, as do the sacra of Polacanthus (9) and Hyl%osauru>^
(10). The attachments for the sacral ribs are large, and are
placed about midway dorso-ventrally of the centra of the sa-
crum. While the sacrum of Stegopelta resembles in a great
measure that of Polacanthus, yet it varies from it in that the
elements composing it are more highly coossified. This condi-

moodie: armored cretaceous dinosaur. 263

tion cannot be entirely attributed to the age of the individual
but is expressive rather of the higher specialization of the pres-
ent form.

Attached to the neural spines of the sacral elements was a
long, narrow dermal plate (plate LVI, fig. 3), evidently in firm
union with them. This plate occupied the mid-dorsal line of
the great pelvic shield and lay between the iliac armor of the
two sides. The plate, like all the rest of the dermal skeleton, is
scrobiculate and somewhat of the texture of woven cloth, as
Marsh has described in Nodosauriis textilis (11). It becomes
slightly expanded posteriorly, where it measures 30 mm. in
width.

The caudal vertehrse (plate LIX, fig. 1, and plate LVIII,
fig. 3) vary to a remarkable degree from those of the dorsal
series. They are greatly compressed dorso-ventrally ; are very
short, with the sides much rounded and the centra plano-con-
vex. The vertebrae have strong transverse processes. In
neither specimen is the neural spine preserved, so it is im-
possible to say as to its extent. This, however, may be inferred
to have been low and stout, if one may judge from its broken
bases. The neural canal is not large, measuring only 14 mm. in
diameter. The centra measure 82 mm. in width by 38 mm. in
length by 41 mm. in height. It is thus seen that the caudal
vertebrae are more than twice as wide as long. This condition
is not a unique one, since Hulke {12) has described the same
condition in Polacanthus.

The rihs are all so fragmentary that no discussion of them
can be given. Many of the fragments are triangular in cross
section, agreeing in this respect with other armored dinosaurs.
Attached to some of the rib fragments are found small bony
scutes. The fact is of service in determining the position of
these dermal scutes on the body of the animal.

The idna (plate LIX, figs. 8 and 9) of the left side is pre-
served complete. It shows considerable variation from any-
thing hitherto described for any other armored dinosaur. The
shaft of the bone is flattened from side to side and is twisted
from right to left. It ends in a rather pronounced expansion
distally. The coronoid process is quite prominent and is ex-
cavated to form a large sigmoid cavity. The olecranon is long,
measuring 70 mm. in length from the base of the coronoid
process to the proximal end of the bone. It is flattened from

264 KANSAS UNIVERSITY SCIENCE BULLETIN.

side to side and ends rather broadly, not in a point as does the
ulna of Stegosaurus. Viewed from in front the ulna curves
strongly outward, especially toward the distal end. It is not
so straight as in Stegosaurus. The articular surface for the
humerus is small. The radial articulations are, however, well
marked, with the upper one forming a distinct fossa below the
coronoid process. The lower articular surface of the ulna is
well rounded and is oval in outline. The ulna measures 385
mm. in total length, 106 mm. antero-posteriorly at the widest
part, 45 mm. near the middle of the bone and 60 mm. at the
proximal end.

The radius is represented by fragments only. The head of
one of the elements is nearly circular in outline and measures
60 mm. in cross section. The articular surface is so abraded
that it is impossible to determine its form.

The ilia (plate LV) are both represented in the collection.
The left one is preserved nearly entire. Enough is present to
determine the total length and breadth of the bone, and the
missing parts have been restored, as is evident from the photo-
graph. The portion of the right ilium preserved shows the
acetabulum. The description of the ilium will, however, be
confined to that of the left side. The bone is elongate and
slightly convex upwards ; the convexity becoming strongly
pronounced on the outer side. It is 50 mm. thick at the acetab-
ulum, but thins to 5 mm. on the inner side and at the anterior
end. The thinning is gradual in these two directions. On
the outer edge, however, the bone remains thick for over
three-fourths of its length, when it gradually thins, but not so
much as on the inner edge. The entire dorsal surface of the
ilium is covered by a mosaic of thick, bony, scrobiculate plates
firmly united to it. These plates are more or less pentagonal
in shape and are not all of the same size. On nearly all there
occurs, on some in the center, on others near the edge of the
plate, a pit, which in life may have supported a more or less
elongate horny spine. When the pit is central it is located in
the top of the central eminence which occurs on all of the plates.
The plates were once, apparently, suturally connected, but
there is now no indication of a suture. They form one con-
tinuous bony mass over the entire dorsal surface of the ilium.
The diameter of the individual plates varies from 56 mm. to 105

moodie: armored cretaceous dinosaur. 265

mm., and they are all from 5 mm. to 15 mm. in thickness. A
single plate is shown natural size in plate LVI, figure 8.

The lower surface of the ilium is nearly smooth. The only
irregularities are the acetabulum and small ridges for muscular
attachments. It is gently concave, the concavity becoming
stronger towards the outer edge. The acetabulum is in the
shape of an elongate oval and is defined by a high ridge of
bone. The acetabular cavity is rather shallow and is deeper
anteriorly than posteriorly. Its length is 190 mm. and the
width, as preserved, is 82 mm. It may have had a somewhat
greater width, but not to exceed 90 mm. at the most. The
entire ilium measures 48I/2 inches in total length, IOI/2 inches
at the widest part, and 8I/2 inches near the middle of the bone.

The ilium must have had a horizontal position during life,
for the position of the acetabulum is such that the femur could
not have articulated in it had the bone been in any other
position. This condition is not exceptional among dinosaurs,
since the ilia of Polacayithiis had a similar position and the
ilia of Triceratops are horizontal to a much less extent. The
articular surfaces for the pubes and ischia have been broken
away and lost, so it is impossible to determine the relations
of these elements. However, they must have had much the
same relations as Seeley (13), Lydekker (17) and Hulke (li^)
have described for Polacanthus.

The pubes (plate LVIII, figures 1 and 2) are both repre-
sented, the right one nearly complete but lacking the entire
post-pubic process. This element resembles more nearly that
of Claosauriis (15) than it does that of its nearest allies,
Polacanthus (13) and Stegosau7'us (16). In form the bone is
elongate, flattened and spatulate. It curves gently inwards
and is only slightly twisted. In thickness the pubis is nearly
uniform, becoming thickened proximally to support the pec-
tineal process (ectopubis of Baur). The pubis measures 300
mm. exclusive of the pectineal process, which is 57 mm. in
length. In width the bone measures 112 mm. at the widest
part and 86 mm. at the distal end. The external surface of
the pubis is crossed by a diagonal ridge for muscular attach-
ments. The post-pubis is not represented in the collection
save by a fragment which may represent the disal end. The
place where the process has been broken from the pubis is
large, so the post-pubis may have been of some extent.

266 KANSAS UNIVERSITY SCIENCE BULLETIN.

The tibia (plate LVIII, figure 4) is represented bj^ the lower
end of the bone belonging to the right side. Firmly coossified
with it is the astragalus, which rises into a rather prominent
ascending process on the anterior surface. The union of the
astragalus with the tibia is so complete that were one to view
the specimen from the posterior side its presence would never
be suspected. On turning the bone over, however, the astraga-
lus is distinctly seen, as is shown in the figure. The spine of
the astragalus is broken off, but judging from the groove for
its reception it must have been nearly 80 mm. in length. It
measures 50 mm. in width at the base. The tibia measures
145 mm. in width distally. The articular surface is broad and
pulley-shaped. There is a broad, shallow, trochlear groove
which is bounded by two rounded projections. The groove
runs up prominently on the posterior surface of the bone but
is not so prominent on the anterior surface. From the nar-
rowness of the broken surface of the bone, only 50 mm. in
diameter, one would judge that the shaft of the tibia was very
slender as described by Hulke (18) in Hylseosaunis and in
Polacmitlnis (19). It resembles these forms also in that the
shaft of the tibia is triangular in cross section, forming in
this case a nearly isosceles triangle. The section shows a
marked difference between the exterior of the bone, which is
smooth and compact, and the interior, which is coarsely can-
cellated.

The fibula (plate LIX, figures 4 and 5) is represented by the
proximal and distal portions of the bone belonging to the right
side. The proximal end, which is subtriangular in cross sec-
tion, measures 85 mm. in width and is considerably expanded
as compared to the shaft. The upper end is so weathered that
no traces of the rugose surface for the attachment to the tibia
are left. However, one side is flattened as if for such a union.
The external side of the head is marked by a rather prominent
condyle, which projects some 5 mm. from the surface. The
shaft of the bone is very slender. At a distance of five inches
from its upper end the fibula measures 20 mm. by 30 mm. in
diameter being thicker antero-posteriorly than from side to
side. In structure the fibula is much like that of the tibia,
already described. The distal end, also, triangular in cross
section, is of the same proportions as the proximal end.

The metatarsaU (plate LVIII, figure 7) are represented by

moodie: armored cretaceous dinosaur. 267

several fragments. The proximal and distal ends of one of
them are preserved so that a reconstruction is possible. These
fragments represent a rather elongate, slender element with
the upper surface rounded and the lower surface somewhat flat-
tened. The proximal articular surface is widely concave and
the distal end is pulley-shaped. It is thus evident that the meta-
tarsal is not different from that of other herbivorous dinosaurs.
The length of the bone as restored is 150 mm., the breadth of
the proximal end is 80 mm., and the height 60 mm.

The dermal armor consists of small bony scutes, large dermal
spines and heavy plates, all of various shapes and sizes.
Mingled in with the other bones of the skeleton were more
than four score small bony scutes (plate LVII), varying in
diameter from 15 to 75 mm. and of various thicknesses from
4 to 20 mm. The scutes are all scrobiculate and are for the
most part rounded, though some are elongated, one scute be-
fore me measuring 60 mm. in length by 30 mm. in breadth.
The majority of them possess a ridge asymmetrically placed,
which runs nearly the entire length of the scute. Others run
up to a point in the center ; others are flat.

In structure all of the scutes are of a coarsely cancellated
character, such as Hulke has described for the scutes occur-
ring in Polacanthus. Associated with them were many frag-
ments of heavy triangular ribs with which some of the scutes
were firmly united. From this fact it would appear probable
that the scutes were scattered along the back of the animal an-
terior to the great pelvic shield. The elongated scutes re-
ferred to above may possibly have been situated on the neural
spines of the vertebrae, as in Scelidosaurus (20). The larger
scutes, some of which are as large as the palm of one's hand,
were very probably located near and anterior to the great
pelvic shield.

Scutes very similar to these described for Stegopelta are
found in a number of different armored dinosaurs as well as
among the crocodiles. There are really no structural differ-
ences between the scutes of Stegopelta herein described and
those found in the common Mississippi alligators. The pittings
on the Stegopelta scutes are, however, smaller than in the latter.
Hulke (21) has described a few scutes belonging to Polacan-
thus which resemble the present specimens very much. Lambe
(22) has figured three scutes associated with the form Stereo-

268 KANSAS UNIVERSITY SCIENCE BULLETIN.

cephalus which are almost identical with the scutes of Stego-
pelta. Hatcher {23) was certainly in error in criticizing Lambe's
association of these scutes with dinosaurian remains and in
claiming that they were of a crocodilian nature, since in the
present specimen there can be no doubt as to their dinosaurian
character. Nopsca (^4) has also fallen into the same error
as did Hatcher when he says that crocodilian remains were
associated with the Polacanthus specimen. Seeley {25) has
described and figured scutes similar to the above in Crateomus.
Huxley {26) has described and figured identical scutes in Acan-
thopholis. Marsh {27) has figured some small bony scutes in
Triceratops. Owen has figured scutes of a like nature in
Scelidosatirus {20). Possibly other armored dinosaurs pos-
sessed such scutes.

The question of the morphological value of these scutes is
an interesting one. Judging from the scutes in Stegopelta, one
would incline to the opinion that they represent the rudiments
of a complete bony armor which in time, had the dinosaurs
survived, might have covered the whole dorsum of Stegopelta'. "i
descendants. Such a suggestion might apply equally well to
our modern crocodiles. It is pretty certain that the scutes in
the crocodiles are not vestigial, since it is not now conceded
that the modern Crocodilia are descended from the ancient
armored teleosaurs {28).

The dermal spines (plate LVI, fig. 1, plate LIX, fig. 3) are
all of a dense bony texture with the external surface pitted.
A portion of a small dermal spine is represented, natural size,
in plate LIX, fig. 3. Its base is broken away so it is impossible
to determine its extent, but this was, in all probability, not
considerable. One of the most peculiar portions of the dermal
armor is that represented in plate LVI, fig. 1, which is called
a large dermal spine, and which I have provisionally located
near the base of the tail. The spine is asymmetrical and is
lateral in position, having had a mate on the opposite side of
which nothing remains.

The spine preserved is much broken and the distal tip is
lost. As reconstructed it is of massive proportions in com-
parison to the rest of the exoskeleton. In cross section it is
for the most part triangular, with two sides of the triangle
concave and one convex. It is rounded near the distal end.
The spine, unlike most elements of a similar nature, is bifid,
as is represented in the figure (plate LVI, fig. 1). The short

moodie: armored cretaceous dinosaur. 269

spine measures 80 mm. from the point of its inception to the
tip, and the main spine measures 16 inches from tip to tip.
At its widest part it is 100 mm. in breadth. The surface of the
bone is pitted with irregularly placed cavities as though it were
encased with horny substance.

The exact position of this large spine on the body of the
animal is, of course, largely a matter of conjecture, since it was
found broken and disassociated and mingled with various
other parts of the skeleton, but it seems best to locate it back
of the pelvic shield, since the caudal vertebrae are much stouter
than the dorsal ones and since there is no evidence of any armor
excepting the small bony scutes anterior to the pelvis. The
spine is totally unlike anything described for any other stego-
saurian dinosaur.

The dermal plates, represented in plate LVI, figs. 2 and 9,
are massive and of a compact texture. Their position on the
skeleton is a puzzle. The plate represented in plate LVI, fig.
9, is not entire. It is asymmetrical, one side being crossed by a
rather high ridge. Its upper surface has pittings and grooves.
The under surface is smooth and rounded. It is possible that
this is a portion of another girdle similar to the one described
below. That it is not a part of that girdle is evident from its
proportions. The plate measures 105 mm. in diameter. The
other dermal plate (plate LVI, fig. 2) is apparently complete,
with a broad surface for articulation with some element which
is now lost. This plate is concavo-convex. It measures
140 mm. across in the longest diameter. It is without vascular
markings of any kind, but is smooth as though it were entirely
embedded in the flesh of the animal.

The part of the girdle represented in plate LVI, figs. 6 and 7,
is like the last described plate in that it is of a compact texture.
In the girdle the upper surface is pitted, and crossed at regular
intervals by rather high ridges the sides of which are slightly
concave. This bone at once reminds one of the similar ele-
ment figured and described by Lambe (29) as occurring in
the Stereocephalus form. Lambe (29) however, located the
girdle back of the head, and if his interpretation of the large
fragment which he calls the skull is correct such a location
would be quite plausible. The present girdle differs in a great
degree from that described by Lambe, since it is asymmetrical
and had a mate on the opposite side, of which there are frag-
ments in the collection. It differs further in being a solid mass

270 KANSAS UNIVERSITY SCIENCE BULLETIN.

of bone instead of a bony ring to which the scutes are united.
The girdle as preserved is in the form of an arch and is com-
posed of three plates, only two of which are ridged. The under
surface is smooth and longitudinally convex. The part of the
girdle preserved measures 320 mm. in length and 110 mm. in
width.

Stegopelta finds its nearest allies in Polacanthus, from the
Wealden of the Isle of Wight; in Paleoscincus, which must, I
think, include the Stereocephalus of Lambe, from the Belly
river deposits ; and Stegosaurus, from the Lower Cretaceous of
Wyoming and Colorado. The characters which separate Steg-
opelta from Stegosaurus and Paleoscincus have already been
considered. From Polacanthus the present form differs in the
whole surface of the ilium being covered in the former animal
with a firm mosaic of very small, somewhat rounded, plates,
which are scarcely at all sculptured, as I learn from Doctor
Williston, who has examined the type specimen. These plates
in Stegopelta are relatively quite large and strongly sculptured.
The present form differs from Polacanthus also in the diver-
gence of the points of the ilium anteriorly. In Polacanthus
the iliac region was entirely covered by a bony shield, but in
Stegopelta this shield was not continuous, but there was a space
between the anterior ends of the ilia which was filled, probably,
with small bony scutes. There have been many stegosaurian
dinosaurs described from various parts of the world, notably
North America and Europe. The group seems to have had a
very wide distribution geographically.

In conclusion, I wish to express my gratitude to Dr. S. W.
Williston, not only for his kindness in turning over the material
to me for description but for his kindly interest and help during
the entire investigation.

The specimen is in the Walker Museum, University of Chi-
cago.

Since the above was written there have appeared two papers
describing two new types of dinosaurian reptiles closely allied
to the above-described form. One of these papers,* from the
American Museum of Natural History, is of especial interest,
since the material supplements in an excellent way that which
has just been described. Mr. Brown's restoration of the Ankij-
losaurus magniventris, while based in large part on Marsh's

* Brown, Barnum, 1908. Bulletin Amer. Mus. of Nat. Hist., vol. XXIV, p. 187.

moodie: armored cretaceous dinosaur. 271

drawing of Stegosaurus ungulatus, is not accurate as to the
form of the animal. The pelvic region was undoubtedly cov-
ered over with a heavy shield as in Stegopelta. Indeed,
Doctor Willistonf says, "Ankylosaurus is either very closely
allied to or identical with Stegopelta Williston, a genus over-
looked by Mr. Brown."

The remains described by Mr. Brown consisted of a skull,
some small dermal scutes, a few vertebrse, a scapula, a few
ribs and other parts, most of which were lacking in the above-
described animal.

Dr. O. Abel has given (Verhandl. d. k. k. Zool.-bot. Gesell. in
Wien, Bd. LVIII, p. 215, fig. 3) a restoration of Ankylosaurus
magniventris Brown. It is to be doubted if he has added to our
knowledge of dinosaurian attitudes, since the reader is at once
struck with the grotesque and impossible attitude which he has
caused the animal to assume. It is unfortunate that Brown's
ideas of the arrangement of the scutes on the back should have
been given by Doctor Abel, since there is not the slightest evi-
dence that they assumed anything like the arrangement he has
represented.

The other dinosaur is from the Niobrara Cretaceous of Kan-
sas and was collected by Mr. Charles Sternberg in the Hack-
berry creek region.

Doctor Wieland$ describes these remains as Hierosaurus
sternbergii, new genus and species. This form is also doubt-
fully distinct from Stegopelta, unless it be that the skeletal
characters were widely at variance to the characters of the der-
mal plates. A comparison of figure 9, plate LVI, of the present
essay, with Doctor Wieland's figure 6, will show how very
closely the dermal elements of the animals resemble each other.
Comparisons also of the figures on plate LVII of the present
paper with Doctor Wieland's figures 2, 4 and 5 will reveal the
close similarity of the small dermal elements. It would, how-
ever, be too previous to say certainly that Hierosaurus and
Stegopelta are identical. The geological horizon of the two
specimens is certainly not far apart, or perhaps identical, since
the Hailey shales occupy relatively the same position in Wyom-
ing as the Niobrara chalk does in Kansas.

t American Naturalist, vol. XLII, No. 501, p. 629.

+ Wieland, G. R., 1909. Amer. Jour. Sci., March, p. 250.

a-Univ. Sci. Bull . Vol. V, No. 14.

272 KANSAS UNIVERSITY SCIENCE BULLETIN.

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16. Ibid, fig. 6.

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19. HULKB. Phil. Trans., 1881, p. 657.

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22. Lambe. 1902. Contribution to Canadian Paleontology, vol. Ill, p. 56,

pi. XXI, figs. 6, 7, 8.

MOODIE: ARMORED CRETACEOUS DINOSAUR. 273

23. Hatcher, J. B. 1905. Vertebrate Fauna of the Judith River Beds.

Bull. U. S. G. S., pp. 67-103.

24. NOPSCA. 1905. Geol. Mag., N. S., Dec. 5, vol. II, p. 241.

25. Seeley. 1881. Q. J. G. S., p. 653, pi. 31, fig. 3.

26. Huxley. 1867. Geol. Mag., vol. IV, p. 66, pi. V.

27. Marsh. 1896. Dinosaurs of North America, pi. LXX.

28. WiLLiSTON, S. W. American Amphicoelian Crocodiles, Jour. Geol.,

vol. XIV, No. 1, p. 1, 1906.

29. Lambe. Contribution to Canadian Paleontology, vol. Ill, p. 56,

pi. XII, figs. 3, 4, 1902.

THE

KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 15— March, 1910.

(Whole Series, Vol. XV, No. 15.)

CONTENTS:

A Contribution to the Soft Anatomy of Cretaceous Fishes
AND A New Primitive Herring-like Fish from the Texas
Cretaceous, Roy L. Moodie.

PUBLISHED BY THE UNIVERSITV,
LAWRENCE, KAN.

October, 1911.

Entered at the post-office in L.iwi-ence as second-class matter.

3310

THE KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 15.] MARCH, 1910. [yjL'xVNo!"

A CONTRIBUTION TO THE SOFT ANATOMY OF CRE-
TACEOUS FISHES AND A NEW PRIMITIVE
HERRING-LIKE FISH FROM THE
TEXAS CRETACEOUS.

BY ROY L. MOODIE.

(Contribution from the Zoological Laboratory, No. 191.)

Plates LX to LXII.

''f'^HERE have been, during the past few years, several ad-
1 ditions to our knowledge of the soft parts of extinct
animals. This knowledge has to do, in large part, with
the firmer tissues, such as the cartilaginous portions of the
skeleton, the skin, the muscles, but in some cases the kidneys,
oviducts, nervous tissues, blood vessels and alimentary canal
are clearly preserved. Dean (1) has been especially fortunate
in the discovery of some of these structures in the sharks of
the Cleveland shales of Ohio. He has described very fully the
preservation of the kidneys, muscles, skin, the cartilaginous
elements of the fins and arches and portions of undigested food.
So perfectly are the remains preserved that the tissues, in
some cases, admit of histological differentiations into the
component elements. Eastman (2) has described the preser-
vation of the outline of some acanthodians from Mazon Creek.
Woodward (3) has contributed to our knowledge of the soft
anatomy of fossil fishes in many ways and has added interest-
ing information on the anatomy of the lateral line system of
Cretaceous selachians. Jaekel (J^), Dean (1, p. 267) and Gill
(5) have discussed the anatomy and the significance of Juras-
sic and Cretaceous chimseroid egg cases. Otto Reis (6) has
written much on the soft anatomy of various fossil fishes, more

(277j

278 KANSAS UNIVERSITY SCIENCE BULLETIN.

especially the Coelacanthidse, in which he has described the
form of the muscle fibers, the swim bladder and other struc-
tures. Eastman and Parker (21) have described the preserva-
tion of the brain, the internal ear and arterial vessels in
Rhadinichthys deani Eastman from the base of the Waverly
shales, Kentucky. Dean (20) has mentioned the preservation
of the lateral line sensor^' canals of the head, the auditory
organ and the rim of the nasal capsule in Acanthodes bronni
from the Pe/mian of Lebach, preserved in the Berlin Museum.
Fritsch (7) has described very accurately the outlines of the
body and fin membranes of Pleur acanthus. Traquair (5),
Dean {1) and Sollas {9) have added to our knowledge of the
anatomy of Paleospondijlus. Patten (iO), Eastman, Traquair
and others have written on various structures of the Ostra-
cophores. Other authors have contributed from time to time
on the subject, until we have, in some instances, e. g., Bothrio-
lepis, Paleospondylus, Cladoselache, a fairly definite idea as
to the outward form and internal structure of the creature.
Among higher animals something has been done on the soft
anatomy of the extinct Amphibia, Ichthyosauria, and Dino-
sauria.

It is with some degree of pleasure that the w^riter is able to
add to the knowledge of the soft anatomy of extinct forms by
the discussion of the alimentary canal of two Cretaceous fishes.
One is a species of Empo, probably E. nepaholica Cope, from
the Niobrara Cretaceous of western Kansas, and the other is
a new species of clupeoid fish from the Cretaceous of Texas.

The specimen which probably belongs to Empo nepaholica
Cope consists of the cast of a large stomach which, in all
probability, represents a fish of some ten or twelve feet in
length. It is No. 347 of the University of Kansas Paleontologi-
cal Collection. The remains were discovered in 1897 by Mr.
H. T. Martin in the Niobrara chalk four miles northwest of
Elkader, Kan. The specimen has recently been presented to
the Museum of the University by Mr. Martin.

The specimen consists of a cast of the larger portion of the
alimentary canal of a large species of fish. Attached to the
matrix of the cast on one side is the major portion of the right
pectoral fin, which is described and figured below. So far as
the writer is aware, the present specimen is the most perfect
example of the pectoral limb of an Empo which has been de-

moodie: cretaceous fishes. 279

scribed. Cope (ISa) figured an incomplete one and Hay men-
tioned another. Hay (11) has figured a portion of the caudal
fin of an Empo showing the extreme character of segmenta-
tion of the rays. The same character is shown in the pectoral
fin.

The stomach is rounded, somewhat laterally compressed,
and elongate in a shghtly U-shaped curve. There are eight
muscular constrictions on the ventral surface and four on the
dorsal. The constrictions, on the ventral surface, occur in
groups. Anteriorly there are two close together. This group
is separated by a space of an inch and a quarter from the next
group, in which there are three, which occur a little over one-
half inch apart. The last group, also of three, is separated from
the second group by one inch. The surface of the stomach
cast is covered with a dark, apparently carbonaceous, ma-
terial which may be carbonized muscle, together with a few
large scales of the typical Empo form. Running the entire
length of the specimen are longitudinal ridges and grooves
showing the cast of the muscular walls of the stomach.

The interior of the stomach, in cross section, shows no food
material, but only chalk. It is possible that the fish, like some
of its modern relatives, may have been a bottom feeder and its
stomach may have been partly filled with Niobrara mud at the
time of its death. There must, however, have been some sedi-
ment enter the stomach after death, for the full form of the
organ is preserved as though the entire stomach cavity had
been packed with mud. Furthermore, the form of the stomach
is that of a carnivorous fish, and recalls very strongly the
stomach of a mountain trout or of the sunfishes of our inland
streams, all carnivorous in habit.

The portion of the alimentary canal preserved is in two
lobes. The first lobe is undoubtedly the stomach proper, and
the constriction between the lobes is the pyloric region. The
other lobe is unlike anything among modern fishes with which
I am at present acquainted. It is undoubtedly an enlargement
of the intestine and possibly served as a secondary stomach.
It lacks the muscular constriction and the longitudinal plicse.
The plicse are, however, continued well across the pyloric
region to the beginning of this second enlargement.

The pectoral fin, as preserved, is well characterized in the
photograph (plate LXII, figure 2). It is somewhat turned in-

280 KANSAS UNIVERSITY SCIENCE BULLETIN.

ward and bent, during interment, back against the stomach.
There are eleven rays preserved. The anterior rays are cross
segmented with long divisions, which measure 7 mm. in
length in the second ray. The square notches mentioned by
Hay {11, p. 87) as occurring on the specimen of Em'po nepaho-
lica Cope in the United States National Museum, are entirely
lacking from that portion of the anterior ray which is pre-
served in the present specimen. The teeth on the edge are
also absent, nor do I find that they are evident in Cope's figure
referred to by Hay. The figure is very indistinct, and if the
notches were present they could not, in the nature of the case,
be normal, but would represent places where the segments had
dropped out. The first ray is not a spine. In other respects
the present specimen agrees well with that figured by Cope
on plate LU, figure 1 (Cretaceous Vertebrata). The fifth and
succeeding rays are segmented like the anterior ones, but the
segments are smaller and measure, on the average, only about
2 mm. The seventh ray is especially broad, equaling in its
proportions two and one-half of the other rays. All of the
rays are split distally. The seventh divides into four second-
ary rays and the divisions ascend more and more to the base of
the fin posteriorly. The fin supports are obscured by scales
and matrix so that their nature cannot be determined. On
the opposite side of the specimen, below the pectoral fin, there
are large scales and fragments of ribs.

The second intestinal enlargement is interesting, entomo-
logically, as showing ttie borings of some fossorial hymenop-
teron ; possibly some one of the smaller species of the
Andrenidse. There are fragments of pupa cases in the burrows,
so there is no doubt as to the recent origin of the holes.

The present specimen is so far the only remains known of
the soft anatomy of the Kansas Cretaceous fishes, and, so far
as I can learn, the first indication of the alimentary canal of
Cretaceous bony fishes of any region. Whether the stomach
and intestines in their various forms will ever be of any help
in determining the relationship of the various osseous fishes
remains to be determined. It is to be feared, however, thai
the fishes have been so diversified according to food habits that
these structures will not be of any great phylogenetic value.
The remains are interesting, however, as indicating, in a
measure, the habits of life of at least one of the Cretaceous
fishes.

MOODIE: CRETACEOUS FISHES. 281

Measurements of the specimen of Empo nepaholica Cope :

Entire length of the alimentary canal as preserved . . 53 . 2 cm.

Greatest diameter at anterior end 10

Least diameter, across pyloric region 3

Greatest diameter of posterior enlargement 7.4

Length of pectoral fin as preserved 9

Greatest width of fin 3.7

Length of first ray. 8

Width of first ray 2 mm.

Diameter of large scale 15

Thrissopater intestinalis new species.

A species of clupeoid fish is represented in the University
Museum by the remains here described as a new species. The
form is located in Thrissopater of Giinther, described from the
Gault of Folkstone in 1872 (12). My thanks are due Dr. A.
Smith Woodward for the suggestion of a comparison of the
present form with that of Thrissopater. It was thought for a
time that the present form represented a genus distinct from
Thrissopater. The distinguishing character was thought to be
found in the position of the pelvic fins, which has served as a
generic character in other fishes. In Thrissopater salmoneus
the pelvic fin is opposite the dorsal and in the present form it is
distinctly posterior to it. There is, however, a great range of
variation in the position of the pelvic fin, especially among the
lower osseus fishes. My thanks are due Prof. E. C. Starks for
aid in reference to the characters of the modern bony fishes.
During the summer of 1909 the writer spent some weeks study-
ing with him the fishes of Puget Sound. He first called the
writer's attention to the wide variation of the location of the
pelvic fin in the clupeoid fishes. This variation is easily un-
derstood when it is remembered that the pelvic fin lies free
from any firm attachment and hence its variation in location
would not mean as much as though it were attached to the
scapular arch. Further aid was rendered the writer in de-
termining the characters of the clupeoid fishes by Dr. W. G.
Ridewood, of London. An examination of the essays of this
gentleman has been of great service.

The absence of material for direct comparison with the
species of Thnssopater makes it best to locate the present
form temporarily in that genus. The systematic position of
Thrissopater has been the subject of a wide variance of
opinion. Dr. Giinther regarded Thrissopater as closely allied
to the modern Clupeidse and located it {13) in that family, in
which he also included such forms as Spaniodon, Albula, Elops

282 KANSAS UNIVERSITY SCIENCE BULLETIN.

and Engraulis, all of which have been regarded by different
authors as types of distinct families. Boulenger {H) regards
Thnssopater as a member of the subfamily Thrissopatrinae,
which is one of his four subfamilies of the Clupeidse. Dr.
Jordan (15) located the form in the family Spaniodontidse,
which is closely related to the Elopidse, between which and the
Clupeidse Boulenger regards Thrissojyater as being interme-
diate (I.e., p. 564). Professor Starks writes me that Doctor
Jordan now regards the family Spaniodontidse as untenable.
Dr. A. Smith Woodward (16) regards Thrissopater as a mem-
ber of the Elopidse, which differ from the Clupeidse in the pos-
session of a single supramaxillary, the degree of union of the
parietals and the gape of the mouth and the presence of a
gular plate. The present form presents the characters of the
Elopidse in so far as they are preserved. In recent forms, the
presence of a gular plate in the Elopidse serves as a convenient
landmark for the distinction of the families of the Elopidse and
Clupeidse. As a matter of fact, the families are so closely allied
that the characters used for their separation must in time be
broken down by the discovery of new material.

Herrings and herring-like fishes are not at all rare in the
Cretaceous deposits of the world. Davis {17) has described
many forms of clupeoid fishes from Mount Lebanon. Before
him Agassiz advanced the knowledge of these forms, and lat-
terly Woodward has described several interesting clupeoids.
Cope described several clupeoids from the Eocene of Green
River, Wyoming {18), and Jordan has cited the interesting
relations of these forms to forms now living in the rivers of
Australia and Chili. At the present time herrings form an
important item in the economic history of the world. Huxley
has dwelt {19) especially on the anatomy and relations of the
herring in this connection. The present form adds yet another
mite to our knowledge of these interesting fishes. It is believed
to be as early as or perhaps somewhat older than many of the
described clupeoids. The specimen comes from the Austin shales
or limestone, which is a probable equivalent of the Niobrara
Cretaceous. It is from near Baylor, Tex., and is No. 300 of
the University of Kansas Museum.

The remains preserved consist of the nearly complete fish,
as may be determined from an examination of the plate. The
caudal portion is, unfortunately, lacking. The outer surface
of the skull was badly broken and Mr. Martin very kindly ex-

MOODIE: CRETACEOUS FISHES. 283

tricated the skull for me from the matrix. It was a difficult
task and the results were hardly worth the efforts, for the
embedded portion was but little better preserved than the
outer. Enough of the skull is preserved, however, to show
many of the important characters. The head is naked; the
body compressed, but whether the ventral edge was drawn out
into a keel or not cannot be determined from the specimen.
The mandible is fully as long as the skull. The relations of the
articulation to the orbit cannot be determined, nor can the
position of these openings be definitely located. The parietal
bones are, apparently, small. Certainly the supraoccipital
projects forward as in Thrissopater magnus. Maxilla is
slender, with a single supramaxillary. The margin of the
jaws is provided with a single row of small, recurved, sharply
pointed teeth of uniform size throughout the length of the en-
tire mandible and maxilla. The quadrate is broadly V-shaped,
with a prominent articular surface. Nasal, ethmoid and pre-
maxillary bones ornamented with numerous small pits. The
same character occurs on the anterior end of the maxilla of
the right side. A single, squarish, punctate, thick, pharyngeal
bone is present. A very few branchiostegal rays are pre-
served; not over ten. From the relationships of the form we
would judge there were many in the complete fish. The oper-
cular apparatus is smooth. Posterior suborbital plate radiately
furrowed; its extent exceeding one-third of the length of the
skull; remaining elements indistinct. Greatest depth of the
body is slightly greater than the length of the skull from
premaxilla to supraoccipital. The length of the body is
possibly equal to four times the length of the skull. The fins
are relatively small. Dorsal fin median in position. The
pectoral fin has sixteen rays, which are cross-segmented but
are not divided longitudinally. The rays are supported by
five baseosts. The distance between the origins of the pectoral
and pelvic fins is equal to nearly four times the length of the
pectoral fin. The pelvic fins have nine rays, none of which
are cross-segmented. The pelvic bone is large and spatulate.

The body scales are small, cycloid, deeply imbricated and
marked with fine concentric lines. There is a large, elongated
and elegantly sculptured scale at the base of the pectoral fin,
as in Thrissojmter salTnoneus; though in the present instance
the scale is less than one-half the length of the pectoral fin
rays. The vertebrse are preserved to the number of thirty-

?-Univ. Sci. Bull.. Vol. V, No. 15.

284 KANSAS UNIVERSITY SCIENCE BULLETIN.

four. There may have been as many as forty-five or fifty in
the complete fish. They are fully ossified, slightly constricted
and marked with small longitudinal ridges. The length is
slightly greater than the depth. The neural spines are long
and interlock with the interneurals. Supernumerary ribs
present. Six of them occupy the space of a single vertebral
centrum.

The specimen as preserved is well characterized by the fig-
ures. The fish lacks the posterior end of the body back of the
anus. It is chiefly remarkable on account of the extraordinary
preservation of the casts of the rectum and intestine, of which
there are six coils or loops preserved. The remains are em-
bedded on the right side in a calcareous, arenaceous, shaley
limestone, which also contains remains of some species of
Inocey^amus, small fish teeth and the base of a moderately
large shark's tooth.

Perhaps the most interesting portion of the entire specimen
is the intestinal canal, from the presence of which is derived
the specific name. In general features the alimentary canal as
preserved recalls that of the common fresh-water buffalo fish,
Ictiobus bubalus Raf (plate LXII, figure 1). The similarity
in form is undoubtedly indicative of similarity of habit, and
since we know that the buffalo fishes are bottom feeders we
can easily predicate that our ancient Cretaceous fish had similar
habits and at the time of its death the alimentary canal was
filled with mud mixed with some organic substances ; for the
fossil shows a different texture for the cast of the alimentary
canal from the matrix, indicating different materials. The
intestine as preserved consists of six coils or loops of the very
small intestine which immediately precedes the rectum, which
is likewise preserved. The rectum is elongate but no more so
than is the same structure in the buffalo fish. The essential
characters are shown in the illustrations.

The distinction of this species from the other three which
have been assigned to Thrissopater is to be found, first of all,
in the posterior location of the pelvic fin. Its base lies at a
distance posterior to the back edge of the dorsal fin, which is
equal to its own length. So far as I am aware the large axil-
lary scale in other species of Thrissopater is larger and un-
ornamented. From T. salmoneus Giinther the present form is
to be distinguished by the relative proportions of the head and

MOODIE: CRETACEOUS FISHES. 285

body. The head and opercular apparatus is contained only
about twice or at most two and one-half times in the body.
From T. magnus it is to be differentiated by the relative pro-
portions of the opercular and the skull. The former is con-
tained twice in the latter in the present form. In T. magnus
it is contained three times. The present species is indeed very
closely allied to Thrissopater magnus Woodward from the
Lower Chalk of Hollingbourn, Kent. It is to be distinguished
by the relative dimensions of the vertebrae as well as by the
proportions existing between skull and opercular. The verte-
brae in T. magnus are higher than long while in T. intestinalis
they are slightly longer than high, and the ends are occupied
by distinct rims, such as do not occur, apparently, in the
English form. The characters which the two species have in
common are striking. They both have the same notch in the
anterior end of the mandible ; the same finely punctate ethmoid
and nasals; the same form and dimensions of mandible and
maxilla; the same divided posterior suborbital; and the same
relative shape of skull. Many of these are generic characters.

The present species can be distinguished from Thrissopater
(?) megalops Woodward by the proportions of the head. In
T. ( ?) megalops the height of skull from cotylus to supraoc-
cipital is equal to the length of the mandible, while in T. ??-
testinalis the mandible exceeds the height of the skull from
cotylus to supraoccipital by 15 millimeters. It may be further
distinguished by the relative proportions of the pectoral arch
and skull as well as by the absence of the radiately furrowed
suborbital and the notch in the anterior end of the mandible
in T. (?) megalops.

Measurements of Thrissopater intestinalis Moodie :

Length of specimen 29 cm.

Greatest depth 9

Length of skull (with opercular apparatus) 9.7

Depth of skull at quadrate 4.8

Length of mandible 6

Depth of mandible at cotylus 1

Diameter of pharyngeal plate 9 mm.

Length of tooth 2.5

Width of opercular apparatus 3 cm.

Length of clavicle 2.5

Length of pectoral fin 2.5

Width of pecoral fin 1.2

Length of pelvic fin 2.4

Width of pelvic fin 1.2

Length of actinost 3 mm.

Length of dorsal fin as preserved. 2.1 cm.

286 KANSAS UNIVERSITY SCIENCE BULLETIN.

Width of dorsal fin as preserved 1.8

Length of caudal haemapophyses 9 mm.

Length of vertebrae 6

Depth of vertebrae 5

Width of small intestine 3

Length of rectum 12 cm.

Width of rectum 1.3

Length of interneural 1

BIBLIOGRAPHY.

1. Dean, Bashford, 1909. Studies on Fossil Fishes (Sharks, Chima-

roids and Arthrodires) . Memoirs of the American Museum of Nat-
ural History, vol. IX, part V, pp. 211-287, with 65 text-figures and
plates XXVI-XLI.
la. HussAKOF, Louis, has given a very full bibliography of the forms
here discussed. Studies on Arthrodira, Memoirs of the American
Museum, 1906, vol. IX, part III.

2. Eastman, Charles R., 1902. The Carboniferous Fish Fauna of

Mazon Creek, Journal of Geology, vol. X, p. 536.

3. Woodward, A. S., 1888, Paleontological Contributions to Selachian

Morphology, I : On the Lateral Line of Cretaceous Species of
Scyllidae. Proc. Zool. Society of London, p. 126.

1898. Preliminary Note on a New Specimen of Squatina

from the Lithographic Stone of Nusplingen, Wurtemberg. Geo-
logical Magazine, December, IV, vol. V, No. 409.
3a. COLLINE, W. E., 1895. The Morphology of the Sensory Canal System
in Some Fossil Fishes. Proc. Birmingham Phil. Soc, vol. IX.

4. Jaekel, Otto, 1901. Ueber jurassiche Zahne und Eier von Chimaer-

iden. Beilage Band Neues Jahrbuch f. Mineralogie, etc., Bd. 14,
pp. 540-564, fig. 3, Tafeln XXII-XXIII.

5. Gill, Theodore, 1905. An Interesting Cretaceous Chimaeroid Egg-

case. Science, N. S., vol. XXII, No. 567, p. 601, November 10.

6. Rets, Otto, 1888. Die Ccelacanthiden. Paleontographica, Bd. XXXV,

pp. 1-96, Tafeln I-V.
Reis, Otto, 1889. Ueber ein Art Fossilization der Muskulatur.

Gesellschaft f. Morphologic u. Physiologie in Muenchen, pp. 1-6.
Reis, Otto, 1894. Ueber Phosphoritisirung von Cutis der Testikel

und des Ruckenmarks bei fossilen Fischen. Archiv f. Mikro.

Anatomie, Bd. 44, pp. 87-119.
Reis, Otto, 1898. Neues ueber petrificirte Muskulatur. Archiv f.

Mikros. Anatomie Bd. 52, pp. 262-268.

7. Fritsch, Anton, 1895. Fauna der Gaskohle und der Kalksteine der

Permformation Bohmens, vol. Ill, p. 1.

8. Traquair, R. H. See bibliography in reference No. 9.

9. SOLLAS, W. J. and I. B. J., 1904. An Account of the Devonian Fish,

Palieosjwndyhis gunni Traquair. Phil. Trans. Royal Society,
London, vol. 196, B, pp. 267-294, plates 16-17.

10. Eastman, Chas. The results of Patten, Traquair, Dean, Eastman

and others are very adequately discussed in Eastman's two

memoirs :
Devonic Fishes of the New York Formations, Memoir No. 10 of

the New York State Museum, pp. 24-65;
Devonian Fishes of Iowa. Annual Report Iowa Geological Survey,

vol. XVIII, pp. 31-291; 41 text-figures, plates I-XVI. 1907.

11. Hay, O. p., 1903. On Certain Genera and Species of North American

Cretaceous Actinopterous Fishes. Bull. Amer. Mus. Nat. History,
vol. XIX, p. 88, plate I, fig. 4.

MOODIE: CRETACEOUS FISHES. 287

12. GuNTHER, A., 1872. Figures and Descriptions of British Organic Re-

mains. December. XIII, Mem. Geol. Survey No. 1, plate I.

13. GuNTHER, A., 1880. The Study of Fishes, p. 656.

14. Cambridge Natural History, vol. 7. Boulenger, Systematic Ac-

count of the Teleostei, pp. 562-564.

15. Jordan, D. S. Guide to the Study of Fishes, vol. II, p. 43.

16. Woodward, A. S., 1901. Catalogue of Fossil Fishes of the British

Museum, part IV., page 32, plate V, fig. 1, VII, fig. 4.

17. Davis, J. W., 1887. The Fossil Fishes of the Chalk of Mount Leb-

anon, in Syria. The Scientific Trans. Roy. Dublin Society, vol.
Ill (ser. II), art. XII, p. 567, plates XXXI-XXXIV.

18. Cope, E. D., 1884. Tertiary Vertebrata, book I, p. 77, plate IX.
18a. Cope, E. D., Cretaceous Vertebrata, plate LII, fig. 1.

19. Huxley, T. H., 1881. The Herring, Scientific Memoirs, IV, pp.

473-492.

20. Dean, Bashford, 1907. American Jour. Anatomy, VII, p. 218.

21. Eastman, C. R., 1908. The Devonian Fishes of Iowa, vol. XVIII,

Iowa Geol. Survey, pp. 264-272. Article by Dr. G. H. Parker on
the "Auditory Organs and Other Soft Farts," on p. 272.

3-Univ. Sci. Bull.. Vol. V. No. 15.

THE

KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 16 — March, 1910.

(Whole Series, Vol. XV, No. 16.)

CONTENTS:

A New Species of Holotrich, By Nadine Nowlin.

PUBLISHED BY THE UNlVERSITv,

LAWRENCE, KAN.

October, 1911.

Entered at the post-office in Lawrence as second-class matter.

3310

THE KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 16.] MARCH, 1910. [^xv.ToTe

A NEW SPECIES OF HOLOTRICH.

BY NADINE NOWLIN.

(Contribution from the Zoological Laboratory, No. 193.)

Two text figures.

IN April of 1909 there was brought to my room at the
Naples Station a jar of the small barnacle Lepas
'pectiJiata, and in examining microscopically one of the
appendages I observed a minute rotifer-like animal creeping
along its edge. This animal, after being studied for some
time, was found to be an infusorian — a Holotrich, resembling
in a general way Huxley's Dysteria armata. Upon careful
study, however, it differed in so many ways that I concluded
it to be a distinct form.

Symbiosis is such an old and well known condition in the
organic world that any discussion of it for its own sake is un-
necessary here. As in other cases, these two animals live
together, deriving mutual benefit from the combination ; the
ciliate finding a shelter in the appendages and the barnacle, no
doubt, getting a tasty morsel when it succeeds in dislodging
the little animal long enough to sweep it into the gullet. It is a
question how the protozoon manages to thrive under these con-
ditions. We know that the water currents set up by the host
are swift and frequent, and life under such tempestuous condi-
tions would seem worse even than the chances in the open sea.
The structure of the smaller organism partially explains this.
It has first of all an armature, a tough siliceous skeleton, cover-
ing the body, except a narrow strip on the ventral side. With
this protection it is not easily crushed. Then caudally there
protrudes a hook-like tail, which not only helps the animal in

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292 KANSAS UNIVERSITY SCIENCE BULLETIN.

creeping over the rough surface of the appendage, but also
serves as an efficient means of attachment. Near the oral
opening protrudes a long flagellum which lashes rapidly to and
fro and secures food for the animal.

The small visitor thus established on the appendage of a
barnacle obtains its food from the currents of water which are
constantly coming in to the host. When the dangers come and
the appendages are enclosed by the calcareous plates of the
barnacle, the ciliate still has moisture and some food. One
might think that in casting its lot with a sedentary animal the
free-swimming protozoon had limited its opportunities. This
is not wholly true. Lepas pectinata grows on masses of lava,
and during storm these masses become broken and widely
scattered. It is only occasionally that the Naples collectors are
able to find this barnacle — its appearance in the bay depend-
ing upon, first, a storm at sea, and then a strong south wind to
sweep it in. During April and part of May I was able to ob-
tain this material but twice. This was unfortunate as the
nucleus is an interesting one, and promises well for cytological
work. The barnacles live less than a week indoors and the
ciliates perish with them. Moreover, the ciliates are not nu-
merous. As a last resort for material^ I took barnacles from
the storeroom, preserved in formalin, and succeeded in find-
ing the specimens, but they did not prove satisfactory for
nuclear study.

DESCRIPTION OF SPECIMEN.

The animal is oval in form, being slightly wider cephalically.
It is covered with siliceous skeleton, giving the appearance of
a bivalve shell, and possesses a caudal appendage. So differ-
ent is it from the usual infusor that it is little wonder most
zoologists at first glance pronounce it a rotifer, or a mollusk, or
a low crustacean. The animal is flattened dorso-ventrally, the
dorsal shell being more extensive and bending over the right
side. The dorsal is slightly convex ; the ventral is concave and
attaches to the dorsal by a deep groove on the left. The right
edge of the dorsal shell and the left edge of the ventral do not
meet, thus leaving exposed a very narrow strip of protoplasm.
This uncovered protoplasm is ciliated and it is by means of
these cilia that the animal swims. The surface of both shells
is smooth.

The crevice between the ventral and dorsal plate lies then

nowlin: a new holotrich.

293

on the right ventral area of the animal, and because the animal
moves often on the edge we are apt to think of this ciliated
region as purely ventral. Lankeste in describing the family
Dysterina, to which I believe this specimen belongs, calls this
the ventral side. The right ventral groove then extends from
the posterior end of the animal completely along the side and
rounds across the cephalic end. At its caudal extremity is an
appendage, and at the cephalic, a long flagellum.

EXPLANATION OF TEXT FIGURES.

The figure to the left is a view of the right dorsal side of the animal,
showing the flagellum anteriorly and the ventral cilia; the calcareous
supporting rods of the mouth, the two contractile vacuoles and the
nucleus. The caudal appendage is shown by a continuous line and its
range of motion by dotted lines.

The middle figure is a view of the animal on edge, showing a convex
dorsal side, concave ventral, left dorsal groove and caudal appendage.

The right figure is an imaginary cross-section through the nucleus.

The tail, like the body, is covered by the silicious skeleton
and appears hollow, at least part of the way. Whether the
anus lies at its distal end, as is claimed for Onychodactylus, I
am unable to say. Its range of movement is nearly 180 degrees
in the plane of the right ventral groove. In movement the ani-
mal may use this organ as a propeller, shoving itself forward
with long strides by placing the end against a solid structure.
It is this motion which resembles strongly that of a rotifer. In
swimming the tail is usually folded into the groove like a knife-
blade closed in its handle, and the animal is carried forward
by the vibration of its cilia. The animal when creeping over a
solid surface stands on the ciliated edge, much as a mussel when
moving through the sand. In swimming it lies slightly on its
ventral side and has a swinging motion through the water.

The flagellum is located in a deep depression of the cephalic
protoplasm. On superficial examination this depression might

294 KANSAS UNIVERSITY SCIENCE BULLETIN.

be mistaken for the mouth, but the mouth is just ventral to it
and convenient to the currents set up by the flagellum.

The gullet is a long, narrow funnel running obliquely up-
wards and backwards until it nearly reaches the opposite side
of the body. Like other protozoa of this group, it is supported
by calcareous rods. These are plainly visible when picro-
sulphuric acid is first placed on the specimen, but very soon the
acid destroys them and the gullet shows only as a clear place
in the protoplasm.

Two contractile vacuoles are present, and a very large nu-
cleus. The nucleus is especially interesting in these forms be-
cause the meganucleus is heterogeneous, the anterior half re-
maining almost clear when stained with picro carmine, the
posterior half staining densely. The micronucleus may lie
quite far to the posterior end of the meganucleus. The fact
that the two halves of the nucleus react differently to the
same stain suggests a segregation of functions. Since in di-
vision the meganucleus breaks transversely, the resulting ani-
mals are dimorphic as to meganuclear protoplasm.

CLASSIFICATION.

According to Lankester's classification the specimen de-
scribed above belongs to the sub-order Gymnostomata, that
division of the Holotrichs in which the mouth is closed in the
intervals between the acts of ingesting food. It belongs to the
family Dysterinse (Clap. & Lach.), which corresponds to Calk-
ins's sub-family Ervilinse : Cilia confined to ventral surface
or a portion of it; caudal end invariably possesses a movable
style arising from caudo-ventral surface.

Of the various genera grouped under this family, or sub-
family, our specimen corresponds most closely to the last of the
following list :

^gyria (Clap. & Lach.), 1858.

Onychodactylus, Entz, 1884.

Trochilia, Duj., 1841.

Dysteria, Huxley, 1857.

Dysteropsis, Roux, 1902.

The only species of this genus, so far as I can learn, is the
small one found in the lakes of Geneva and called minuta by
Roux, because it measures only 28 micra long and 16 micra
wide. The animal described in this paper is marine; the av-

nowlin: a new holotrich. 295

erage specimen is 83 micra long and 50 wide. Its habitat in
the tentacles of Lepas pectinata suggests the name of Dyster-
opsis pectinata for the new specimen.

BIBLIOGRAPHY.

Bronn, H. G. 1880. Klassen und Ordnungen des Thier Reichs, vols. I,

II, III.
Claparede and Lachmann. 1850-'60. Etudes sur les Infusoires et les

Rhizopodes. Mem. Inst. Genevoise, 5, 6, 7.

Calkins, G. N. 1902. Marine Protozoa from Woods Hole. Bull. U. S.
Fish Comm. 21, pp. 415-468.

Calkins, G. N. 1909. Protozoology. Lea & Febriger.

Huxley, Thos. 1857. Mic. Trans. Jour., vol. 5.

Lankester, Eray. Protozoology, part 1, fascicle 2.

Roux, Jean. 1899. Infusoires cilies des environs de Geneve. Rev.
Suisse, Zool. 6.

Roux, Jean. 1901. Faunne Infusoirenne des eaux stagnents des en-
virons de Geneve. Mem. cour. de I'Universite de Geneve.

Roux, Jean. 1901. Faunne Infusoirienne des environs de Geneve. Vehr.
Zool. Congr. 5 (Berlin).

THE

KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 17 — March, 1910.

(Whole Series, Vol. XV, No. 17.)

CONTENTS:

Orthoptera from the Santa Rita Mountains, Arizona, Col-
lected BY the University of Kansas Expedition, James A. G. Rehn.

PUBLISHED BY THE UNIVERSITY,
LAWRENCE, KAN.

October, 1911.

Entered at the post-office in Lawrence as second-class matter.

3310

THE KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 17.] MARCH, 1910. R^rxv.'No'n

ORTHOPTERA FROM THE SANTA RITA MOUNTAINS,

ARIZONA, COLLECTED BY THE UNIVERSITY

OF KANSAS EXPEDITION.

BY JAMES A. G. REHN.

Academy of Natural Sciences of Philadelphia.

Plate LXIII.

THE collection forming the basis of this paper was made in
the Santa Rita range during the summer of 1907 by
the University Museum expedition under the late Prof.
F. H. Snow.

FORFICULID^.

SpongopJiora apicidentata Caudell.

Santa Rita mountains, 5000 to 8000 feet, July. F. H. Snow.
One female.

This species has previously been recorded from localities in
Arizona ranging from near a hundred feet to about five
thousand feet above sea level.

Blattid^.

Ischnoptera notha Rehn and Hebard.

Santa Rita mountains, 5000 to 8000 feet, July. F. H. Snow.
One female.

This specimen agrees with an individual of the same sex
from the Huachuca mountains previously examined by the
author.*

The measurements of the Santa Rita specimen are as follows :

Length of body 14.0 mm.

Length of pronotum 4.5

Greatest width of pronotum 5.9

Greatest width of tegmen 4.5

Length of tegmen 10.0

* Proc. Acad. Nat. Sci. Phila., 1907, p. 25 (Uhleriana) ; Rehn and Hebard, Ibid. 1910,
p. 443.

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300 KANSAS UNIVERSITY SCIENCE BULLETIN.

In Arizona this species appears to be restricted to certain
elevations on mountains of the southern part of the territory,
viz., Santa Rita, Huachuca and Patagonia ranges.

Blattella dilatafa (Saussure).

Santa Rita mountains, 5000 to 8000 feet, July. F. H. Snow.
One female, one male.

These specimens agree with those recorded by the author*
from the Huachuca mountains.

Mantid^.
Bactromantis virga Scudder.

Santa Rita mountains, 5000 to 8000 feet, July. F. H, Snow.
One female.

This specimen is slightly larger than a Huachuca mountains
individual of the same sex.

Phasmid^.
Pseudoserniyle truncata Caudell.

Santa Rita mountains, 5000 to 8000 feet, July. F. H. Snow.
One female.

Parabacilhis coloradus Scudder.

Santa Rita mountains, 5000 to 8000 feet, June. F. H. Snow.
Two males, one female.

ACRIDIDiE.

Telmafettix aztecns Saussure.

Santa Rita mountains, 5000 to 8000 feet, June. F. H. Snow.
One male.

Mermiria texano Bruner.

Santa Rita mountains, 5000 to 8000 feet, July. F. H. Snow.
One female.

Prorocorypha,^ n. gen.

Allied to Paropomala Scudder, but differing chiefly in the
great development of the cephalic portion of the head, the
greatly elongate supra-anal plate of both sexes and the sub-
genital plate of the male.

Body sub-bacilliform. Head greatly elongate; vertex and
pre-ocular region strongly produced into a rostrate process,
fastigium lanceolate, tectate, carinate; antennse, at least of
male, somewhat triquetrous, subensiform. Cephalic and
median limbs very small. Caudal limbs very slender. Pro-

* Proc. Acad. Nat. Sci. Phila., 1907, p. 26.

' TTpiMi, prow; Kopvfpr), head.

REHN: SANTA RITA ORTHOPTERA. 301

sternum with a median compressed protuberance ; mesosternal
lobes contiguous. Supra-anal plate aculeate in both sexes ; sub-
genital plate of male strongly produced, knife-like.
Type: P. snowi, n. sp.

Prorocorypha snowi n. sp. (Plate 63.)

Types : ^ and 5 (not mature) ; Santa Rita mountains,
Pima and Santa Cruz counties, Arizona, 5000 to 8000 feet, June
i ^), July ( 2 ), 1907. F. H. Snow. (University of Kansas.)
Size medium; form very elongate. Head one and one-half
times the dorsal length of the pronotum, occiput with a hardly
perceptible longitudinal arcuation ; interocular region but little
narrower than the greatest width of the fastigium; rostrum
very long and deep, occupying nearly half the length of the
head and the greater part of the depth ; outline when seen from
the side with fastigium slightly ascending from the interocular
region, face nearly horizontal, inter-antennal portion rounded
to the acute angulate fronto-fastigial angle ; fastigium lanceo-
late, the greatest width contained twice ( ^j ) or more than
twice ( $ ) in the length, apex very narrowly rounded, lat-
eral margins distinctly carinate, surface tectate, median
carina as distinct as the lateral and extending to the
inter-antennal region; lateral foveolse very elongate, lanceo-
late; frontal costa. narrow, very slightly but regularly
expanding ventrad in the male, very narrow dorsad, expanding
somewhat in the inter-antennal region and subequal ventrad in
the female, in both sexes moderately sulcate and becoming ob-
solete before reaching the clypeal margin ; eyes flattened,
elongate ovate, not at all prominent; antennae, in the male at
least, slightly exceeding the head and entire thorax in length,
thick, three-sided, one narrower than the others, ensiform.
Pronotum about twice as long as its width, cephalic margin of
dorsum truncato-rotundate, caudal margin truncate, median
carina distinct ; lateral lobes with the ventral margin truncate.
Wings and tegmina undeveloped. Prosternum with a very dis-
tinct longitudinal rounded tubercle ; mesosternal foramina con-
siderably impressed. Abdomen with the segments considerably
elongate; supra-anal plate in both sexes strongly produced,
needle-like, its length being but very slightly less than the
dorsal length of the pronotum; cerci somewhat compressed,
simple, sub-sty] if orm; subgenital plate of male produced into
a compressed, slightly arcuate knife-like structure, slightly
more than half again as long as the supra-anal plate; ovi-

302 KANSAS UNIVERSITY SCIENCE BULLETIN.

positor jaws of the female placed well under the projecting
supra-anal plate. Cephalic and median limbs very small;
caudal femora reaching to the fifth abdominal segment, very
slender, tapering; caudal tibiae equal to the femora in length,
armed on the external margin with fourteen to sixteen small
spines.

General colors of female burnt carmine and apple green,
the venter of the body and head ochraceous, the carmine
strongest on the dorsum, the green only on the limbs; caudal
femora yellowish proximad. Color of male destroyed by im-
mersion in liquid preservative.

Measurements.

Male. Female.
Length of body (including subgenital

and supra-anal plates) 31.0 mm. 44.0 mm.

Length of head 5.5 7.2

Length of f astigium 2.7 3.8

Length of pronotum 3.2 5.0

Length of caudal femur 10.5 13.6

Length of supra-anal plate 3.4 4.4

Length of subgenital plate 5.0

A single pair have been examined. Doctor McClung, of the
University of Kansas, who has studied the spermatogenesis
of this species and whose assistants collected specimens under
Professor Snow, informs me that "specimens such as those you
have were constantly caught in copulo, and a microscopical ex-
amination of the testes shows the spermatozoa well developed."
The specimens in hand have the tegmina and wing pads in the
reversed nymphal condition.

Eritettix variabilis (Bruner).

Santa Rita mountains, 5000 to 8000 feet, June and July.
1907. F. H. Snow. One male, four females.

One of the females in this series is in a greenish phase of
coloration, while two of the remaining females have a pattern
of chiefly solid contrasting colors. The green phase has the
greater part of the head, dorsal half of the lateral lobes of the
pronotum, the pale stripe on the tegmina and the dorsal face
of the caudal femora apple green, while the strongly contrasted
phase has broad seal brown postocular bars extending over the
head and pronotum and regularly expanding in width, the
dorsal aspect much lighter, either ochraceous or rust red, and
the usual dark portions of the tegmina also seal brown. The
supplementary carinas of the pronotum are distinctly but not
strongly indicated in two of the females, represented by the

REHN: SANTA RITA ORTHOPTERA. 303

merest traces in the female and completely absent in the two
contrasting colored females.

The only previous Arizona record of this species is from
Douglas, Cochise county.

Amphitornus ornatus McNeill.

Santa Rita mountains, 5000 to 8000 feet, June and July,
1907. F. H. Snow. One male, one female.

Cordillacris pima Rehn.

Santa Rita mountains, 5000 to 8000 feet, July, 1907. F. H.
Show. One male.

This species was known only from the Baboquivari moun-
tains, Pima county, Arizona.

Psolcessa texana Scudder.

Santa Rita mountains, 5000 to 8000 feet, July, 1907. F. H.
Snow. One female.

This specimen is in the blackish phase of coloration — the
typical texayia form.

Aulocara fenioratuni Scudder.

Santa Rita mountains, 5000 to 8000 feet, July, 1907. F. H.
Snow. One male.

This species is now known in Arizona from the Huachuca
and Santa Rita mountains, and from Phoenix.

Arphia aberrans Bruner.

Santa Rita mountains, 5000 to 8000 feet, July, 1907. F. H.
Snow. One male, one female.

This species is now known from localities extending from
the Huachuca range to Nogales, Ariz.

Hippisciis coralUpes (Haldeman).

Santa Rita mountains, 5000 to 8000 feet, June and July,
1907. F. H. Snow. One male, one female.

Scirtetica ritensis, n. sp. (Plate 63.)

Type: $ ; Santa Rita mountains, Pima and Santa Cruz
counties, Arizona. Elevation, 5000 to 8000 feet. July, 1907.
F. H. Snow. (University of Kansas.)

A very beautiful species of the genus, differing from its
allies in the absence of a median band on the wings, the dark
color being on the periphery, in the color of the disk and in the
heavy, robust build.

Size medium; form robust. Head with the extreme width
very slightly greater than the greatest width of the cephalic

304 KANSAS UNIVERSITY SCIENCE BULLETIN.

margin of the pronotum ; occiput slightly arcuate longitu-
dinally, the caudal section of vertex with a pair of distinctly
impressed subpyriform areas; interocular region slightly wider
than the fastigium, broadly and deeply sulcate ; fastigium very
greatly declivent, sulcate as in the interocular space, the sul-
cate portion, however, being slightly broader than between the
eyes; frontal costa somewhat narrower than the fastigium
dorsad, very gradually expanding ventrad to very near the cly-
peal suture, where the margins curve strongly laterad, sulcate
ventrad to immediately dorsad of the ocellus, where the sulcus
terminates abruptly, and from the ocellus ventrad it is present
shallower and sub-obsolete as the clypeal suture is approached ;
lateral foveolse lanceolate, trigonal, distinctly impressed ; eyes
broad, somewhat depressed ovoid in outline, about equal in
length to the infra-ocular groove, moderately prominent when
viewed from the dorsum; antennae about equal to the dorsal
length of the head and pronotum ; ventral portion of head dis-
tinctly broader than the dorsal. Pronotum with the length of
the disk about half again the dorsal length of the head, and
with the greatest width of the disk but slightly shorter than the
length; cephalic dorsal margin very slightly obtuse-angulate,
caudal margin decidedly obtuse-angulate ; lateral angles of the
disk very distinct on the metazona and the cephalic portion of
the prozona, broken and sub-obsolete mesad; median carina
distinct, moderately elevated, straight, cut by the transverse
sulcus very slightly cephalad of the middle; lateral lobes dis-
tinctly deeper than long, the cephalic and caudal margins
obliquely emarginato-sinuate, ventro-caudal angle broadly
rounded. Tegmina exceeding the tips of the caudal femora by
slightly more than the length of the pronotum, moderately
broad, slightly narrowing distad, the apex obliquely rotundato-
truncate; intercalary vein prominent, through the greater
part of its length closer to the median than to the ulnar
vein. Wing with the greatest width contained one and
one-thirds times in the length, apex sub-rectangulate with the
distal portion of the costal margin arcuate. Mesosternal lobes
with the interspace very strongly transverse; interspace be-
tween the metasternal lobes transverse. Caudal femora robust,
the ventro-lateral carina distinctly lamellate, reaching its
greatest expansion at two-fifths the distance from the apex,
pattern of the paginse distinct; caudal tibiae slightly shorter
than the femora, armed laterad with seven spines.

General colors vandyke brown and pea green, marbled and

REHN: SANTA RITA ORTHOPTERA. 305

blotched one over the other, the tegmina, dorsal face of the
caudal femora and disk of the pronotum with the green pre-
dominating; ventral surface Isabella color. Dorsum of the
head and pronotum with a regular figure of green which is
margined laterad with clove brown, this figure being con-
stricted at the caudal margin of the head and almost severed
mesad on the pronotum; antennae clove brown annulate with
pea green, their insertion bordered dorsad and ventrad by a
pair of transverse lines of clove brown ; eye walnut brown.
Wings with the disk orange rufous, the distal and caudal
margins with a moderately broad band of vandyke brown which
fails to reach the proximal margin, ulnar tsenia extending
about half way to the base of the wing ; apex with the smaller
areas and a few of the short veins creamy white. Caudal
femora with the dorsal face crossed by a median and a disto-
median bar of clove brown, while the apex is touched with the
same color, ventral sulcus washed with pale blue; caudal tibise
turquoise blue with the lateral face of the proximal third green-
ish white and the extreme distal portion of the internal face
blackish, leaving a pregenicular pale annulus, the apex dark
blue, and a dark blue annulus placed next to the pale one;
tarsal joints very pale ochraceous.

Measurements.

Length of body 23 . 8 mm.

Length of pronotum 4.5

Length of caudal femur 12 . 0

Length of tegmen 21.0

The type specimen is the only one seen by the author.

Tonionotus aztectis (Saussure).

Santa Rita mountains, 5000 to 8000 feet, July, 1907. F. H.
Snow. One male.

Mestohregma rubripeiine (Bruner).

Santa Rita mountains, 5000 to 8000 feet, July, 1907. F. H.
Snow. One male, one female.

Conozoa carinata Rehn.

Santa Rita mountains, 5000 to 8000 feet, July, 1907. F. H.
Snow. One female.

Trimerotropis allicieyis (Scudder).

Santa Rita mountains, 5000 to 8000 feet, June and July,
1907. F. H. Snow. Three males.

These specimens are somewhat smaller than a female from
the Huachuca mountains.

2-Univ Sci. Bull., Vol. V, Nq. 17.

306 KANSAS UNIVERSITY SCIENCE BULLETIN.

TrimerofropiFi laficiucta (Saussure).

Santa Rita mountains, 5000 to 8000 feet, July, 1907. F. H.
Snow. One male, one female.

This species has previously been recorded from Nogales,
Douglas and the Huachuca mountains in southern Arizona,
and Flagstaff in northern Arizona.

Tmmerotropis cyaneipennis (Bruner)-

Santa Rita mountains, 5000 to 8000 feet, July, 1907. F. H.
Snow. One female.

Heliastus benjamini Caudell.

Santa Rita mountains, 5000 to 8000 feet, July, 1907. F. H.
Snow. One female.

This specimen has pale interrupted decussate markings on
the pronotum.

Previous records of this species are from the Huachuca moun-
tains, the Baboquivari mountains, and Nogales, Ariz.

Heliastus aridus (Brunei") •

Santa Rita mountains, 5000 to 8000 feet, July, 1907. F. H.
Snow. One male.

Dactylotum variegatum (Scudder).

Santa Rita mountains, 5000 to 8000 feet, July, 1907. F. H.

Snow. Two females.

Tettigonid^.
Arethsea sellata Rehn.

Santa Rita mountains, 5000 to 8000 feet, July, 1907. F. H.
Snow. One male.

This is the first record of the species outside of the Huachuca
mountains.

Scudderia furcifera ( Scudder) .

Santa Rita mountains, 5000 to 8000 feet, July, 1907. F. H.
Snow. One male.

GRYLLIDiE.

Gryllus alogus Rehn.

Santa Rita mountains, 5000 to 8000 feet, July, 1907. F. H.
Snow. Two females.

These specimens measure as follows :

Length of Pronotum.

Tegmen.

Caudal femur.

Ovipositor.

4.0 mm.

9.0 mm.

12 mm.

14.5 mm

3.3

5.5

11

13.8

This species has been recorded from the Huachuca mountains
and Phoenix, in addition to Albuquerque, the type locality.

THE

KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 18— March, 1910.

(Whole Series. Vol. XV. No. 18.)

CONTENTS:

Observations on the Gryllid^: III. Notes on the Classification
AND on Some Habits of Certain Crickets, W. J. Baumgartner.

PUBLISHED BY THE UNIVERSITY,
LAWRENCE, KAN.

October, 1911.

Entered at the post-ofHce in Lawrence as second-class matter.

3310

THE KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 18.J MARCH, 1910. [^xv.'no's

OBSERVATIONS ON THE GRYLLID^ : III. NOTES ON

THE CLASSIFICATION AND ON SOME HABITS

OF CERTAIN CRICKETS.

BY W. J. BAUMGARTNER.

(Contribution from the Zoological Laboratory, No. 195.)

Classification.

IN the course of collecting material for some cytological
studies on the crickets, I was early confronted with the
question of what are the true species of Gryllus. The first
specimens collected I {2) called Gryllus assimilis, after care-
fully comparing them with the labeled specimens in the Uni-
versity of Kansas collection, and after reading such descriptions
as were then available in the library. Later I collected about
Chicago, 111., and Woods Hole, Mass. My attempts at classify
ing these specimens led me to the conclusion that the species
of Gryllus, the common larger field crickets, are not fixed but
grade into each other. I found that in all of these places there
were two groups with diflPerent breeding seasons — one that
passed the winter in the nymph stage, and another that passed
it in the egg. The former matures and breeds around Law-
rance, Kan., during June and early July, and the other during
the latter part of August and September.

This question of the true species in Gryllus was frequently
discussed with my fellow student. Dr. F. E. Lutz, now of Cold
Spring Harbor, N. Y., during our study at the University of
Chicago. I am glad to confirm his recent publication (i^) in
which he says that the species of Gryllus as now named do not
differ in characters, "but merely in the degree of common
characters." My study has not been especially along the line

(309)

310 KANSAS UNIVERSITY SCIENCE BULLETIN.

of taxonomic characters ; but my attempts at classifying the
specimens collected in Douglas and Harvey counties, Kansas,
in Chicago, 111., in Woods Hole, Mass., in the Santa Rita moun-
tains, Arizona, and in Tarpon, Tex., trying to follow the keys
given by Scudder, Blatchley, De Saussure and others, have led
me to believe that Lutz is right when he says : "Either we
simply name stages in a great continuous mass of variation and
call them species or there is but one species of Gryllns in east-
ern United States, and the names we give are not the names
of species at all, but simply inaccurate, shorthand expressions
for recording the approximate size, proportions and color of
individuals found." This applies to our common field crickets ;
and I do not think that it should be limited to the eastern Uni-
ted States, but should include the central portion as well.

On a collecting trip to Tarpon, Tex., last summer, I found
a color variation which confirms this opinion that all these so-
called species grade into each other. On the low sandy islands
I found that the crickets were straw yellow. Most of them were
in the last or next to the last nymph stage. This was about
June 12. At first I thought they were the imported Gryllus
domesticus; but later collecting disclosed some with a few, and
some with many dark markings. A few of the adults taken
subsequently were quite black. These were under the same
boards or stones with the straw-colored ones, and were mating
with them ; and they probably came from the same mother. A
number of nymphs were brought to the laboratory at Law-
rence and raised to maturity. All of them turned much darker
and some became jet black. As far as I could see these black
ones could not be distinguished from our native species.

One peculiarity of these crickets on the islands at Tarpon
offered an additional reason for thinking they were G. domes-
ticus, or a closely allied species; i. e., the young nymphs varied
much in their stages of development, a peculiarity I had noticed
in the domestic species in the greenhouses in Chicago. This
fact must be due to the climatic conditions — both forms de-
veloping where there is a long-continued breeding season, with
even temperature.

In the laboratory I found that these Texas forms mated
very readily with our Kansas forms, both the spring-maturing
and autumn-maturing broods. Some of the adults I brought
with me paired with some tardy spring forms and some of the

BAUMGARTNER: OBSERVATIONS ON THE GRYLLID^. 311

smallest nymphs were not matured before autumn adults
appeared.

In trying to classify these southern forms I finally concluded
that they were our common black field crickets, which had lost
a little, very much, or nearly all of their pigment. Although
Blatchley, Scudder, and De Saussure use the color difference as
one of the prominent characters separating species, I do not
believe that it can properly be so used. "Black color" and
"straw color" do not stand for different species in the crickets
of the Texas coast. The black gradually shades off into the
straw color ; and a black one and a light one may have the same
mother.

An examination of the germ cells reveals no differences in
cell structure between the southern light-colored specimens
and our native black ones. But both differ markedly from
that found in Grylhis domesticus, as has been and will be shown
by the papers dealing with spermatogenesis.

All of the collections made in the various localities show
dimorphism as to wing length. The short-winged forms are
very much more numerous in all places, but the long-winged
forms vary greatly in frequency in the several localities, as
Lutz {IJf) has found.

Blatchley {6) is correct when he suggests that the failure
of past monographers of this genus is in part due to the fact
that they have neglected the study of the animals in the field.
By using this method he has added some very useful hints on
the habits and structure, as they bear on the classification. He
has plainly shown that there are in many localities really two
broods, one maturing early and the other later. He considers
them as belonging to different species. Lutz denies this. In
whatever region I have observed the two broods, the autumn
specimens are larger and more robust than the earlier ones.
They also differ in proportions and color enough to represent
two species according to ordinary criteria for species ; but the
intergrading of the forms from different localities would re-
move all distinctive characteristics. So while Blatchley is ap-
parently right, I feel confident that extensive careful collecting
will show that Lutz is correct.

The earlier brood lives "in burrows singly or in pairs,"
while the later ones "are more sociable," and there is not much
"forsaking of burrows," as Blatchley {6) thinks. My ob-
servations have led me to the following conclusions: Of the

312 KANSAS UNIVERSITY SCIENCE BULLETIN.

spring brood each individual has a separate well-made burrow,
early in the season, usually under some stone or board. The
male keeps his as long as he lives, or well through the breeding
period at least ; while the female abandons hers when she be-
comes an adult, or even before. Thereafter she may be found
with the male in his burrow or in any convenient hiding place.

The young individuals of the autumn brood never make much
of a burrow, but live under bunches of loose, dry grass or old
rags, or whatever they find. I have frequently found more
than a dozen in an old newspaper in the grass. The adult
males sometimes have a sort of burrow, particularly late in the
season, but most of the time I find them in any kind of a hiding
place. I am quite sure, however, that these creatures, young
and old, especially the males, have a selected spot in the grass
or paper, which serves as their home, and so the diff'erence
between the spring and autumn forms is really this : the former
dig a burrow for a home, while the latter simply select some
convenient place to stay.

Judging from my study of the germ cells of Gryllus domes-
ticus, given in a former paper (3) , this species is quite different
from the other forms. The diff'erence of chromosome number
and shape are such that I should expect the domestic species to
be very different in taxonomic characters; but such is not the
case.

In other genera of the family Gryllidse the species are more
distinct and limited. In CEccmthns, following Hart (10), we
classify the species largely by the color markings on the basal
joints of the antennae, and this seems to be quite constant. I
have found that I can separate the nymphs quite readily by
means of these markings. Nemobius shows more variation,
and probably after large, widespread collecting the species may
prove to intergrade. In Gryllotalpa the species are quite
distinct.

FOOD HABITS.

Very many observers have written of the food habits. It
is known that the common black field crickets may eat almost
anything. In captivity they will sometimes devour each other,
the stronger ones feasting on the weaker ones even before they
are dead. I have seen a female chew the wing of a male, and
I have found a crippled female with her abdomen partly eaten
away. In their free life I think this rarely or never occurs.

BAUMGARTNER: OBSERVATIONS ON THE GRYLLID^. 313

The females will eat the empty spermatophores whenever they
find them.

Among the mole crickets I observed this peculiarity between
two species. In a box of specimens of Scapteriscus sent me
from Porto Rico I never discovered any partially devoured
ones among the few dead specimens; but while collecting our
own species Grijllotalpa borealis, in northern Indiana, I placed
one female adult and six nymphs in a bottle full of sand in
the field. When I returned to the laboratory I found but two
nymphs, the others having been devoured by the adult. This,
with some later experiences, led me to believe that adults will
eat the nymphs whenever they find them in their burrowings.
However, this cannot be true for the very young nymphs, as the
eggs are laid in a mass in a much frequented part of the
burrow, and the mother, no doubt, cares for the eggs and
young for a while.

EGG-LAYING.

Blatchley (6) says: "The eggs of most crickets are laid
singly in the ground." My observations confirm this as far
as Gryllus and Ncmobius are concerned. The large black field
cricket selects usually a somewhat barren spot in a grassy
field, where she lays her eggs. She will force her ovipositor
into the ground and deposit a single egg, then removing the
ovipositor partly will put it down at a different angle and
plant another egg, and repeating the process will leave a third.
On no occasion did I see more than four eggs laid without the
ovipositor being completely removed and pushed into the
grou'nd at a new place. Nemobius lays its eggs in a similar
manner. Two or three, rarely four, eggs are laid almost side
by side, and then the next batch are placed a quarter of an
inch or more away.

In but one instance did I find eggs laid by the mole cricket.
They were "in a heap on the floor in the enlarged part of a side
gallery," just as Barrett (1) has described.

CHIRPING.

A peculiar habit of the mole crickets, of which I made brief
mention in an abstract (4), is the chirping of the female. A
hurried examination of the tegmina of the females will show
that the nerves are modified into a rasping and sounding organ,
which is not as large or as well developed as that of the male,
but well enough to have made thoughtful observers of the past

314 KANSAS UNIVERSITY SCIENCE BULLETIN.

suspect that it might function, and that female Gryllotalpa
might chirp. As far as I have been able to read the literature
no one has observed that they actually do so. Most of our
books say that only the males stridulate. LaCordiare (12) says,
"The chirping organs of the crickets are simple and limited to
the males." Scudder, speaking of the crickets, says, "his
egotistic love song." Comstock (7) writes, "the males of the
crickets have musical organs." Lang (13) says: "In the Lo-
custidse and Gryllidas only the males stridulate, by rubbing the
rough basal portions of their wing cases against each other."
Packard (15), after speaking of the organs in the males, says :
"The females are not invariably dumb, both sexes of the Eu-
ropean Ephippigera being able to faintly stridulate." Henne-
guy (11), in speaking of the musical organs, writes: "Where
they are found they are well developed in the males only; in
the females they are more or less rudimentary. Such is the
case in the Gryllidse." Barrett (1) describes the stridulating
organ in the male "Changa" or Porto Rican mole cricket,
Scapteriscus didactylus; but he has completely overlooked the
same but less well developed organ in the female.

The female mole cricket has quite a loud and distinct chirp.
It usually consists of a single note; but there may be several
at short intervals. This note is less shrill than the ordinary
call of the male. However, the male has a note very similar to
that of the female which it uses for the same purposes, namely,
as a means of recognition in the dark burrows. The call is al-
ways given when one individual is approaching another, es-
pecially when digging a new tunnel. Both genera, Gryllotalpa
and Scapteriscus, have the stridulating organ on the female
elytra, and so both must be able to chirp. I never isolated a
Porto Rican female to hear its chirp, but after hearing the call
of our native cricket I feel sure that I have heard the insular
female's chirp also.

This unusual ability possessed by the female is an adaptation
to life in underground burrows. It enables the individuals to
recognize others which are approaching under conditions where
sight cannot be used. Thus enemies and friends can be dis-
tinguished; while if the female were dumb, as she is in all
other crickets as far as I know, they might often attack even
their mates. "^

BAUMGARTNER: OBSERVATIONS ON THE GRYLLID^. 315

PROTECTIVE GLANDS IN THE MOLE CRICKETS.

As indicated in an abstract (4) , I have found that the correct
interpretation of the function of the anal gland, which has so
long puzzled investigators, is protective. Leon Dufour (8), a
careful French investigator, first described in the mole crickets
in both sexes a pair of azure or skim-milk colored glands con-
nected with the rectum. Their secretion he compares in con-
sistency with the vitreous humor of the human eye. To this
secretion is added some excrement from the rectum, and when
this mJxture is expelled it forms a brown liquid of nauseating
fetidity. He calls the gland "an organ of excremental se-
cretion."

Berlese (5) describes the same structure and thinks it is a
prostatic gland analogous to that found in the locustids. Al-
though he found it in the female also, he does not seem to try
to explain it there.

Fenard quotes both of the above descriptions and adds a good
many observations of his own. He describes the gland from
sections and gives the action of certain fixatives and stains
upon the tissues of the gland and its contents. He states in
detail the macroscopic and microscopic structure. He concludes
as follows : "Judging from the position of this organ, from the
consistency of the liquid which it contains, and from its points
of similarity with the prostatic glands of the locustids, I think
that it ought to be considered also as a gland furnishing a
mucus destined to lubricate the copulating apparatus. This
organ exists in the female, it is true, but in this case it fur-
nishes without doubt still a lubricant for the vagina, or a
liquid to form the nest of these insects." After describing the
details of this gland in the female he says : "I think that these
organs can only be some secreting agent of a mucus destined
to lubricate the genital organs ; or perhaps they glue together
and hold the spermatophores ; or perhaps again they secrete
the substance used to form the nests in which are found, as
we all know, two to three hundred eggs all massed together
and more or less united." It is evident from this uncertainty
that Fenard did not know the function of the glands in ques-
tion, yet he was inclined to follow Berlese and called them
"prostatic glands."

Packard, in his work on Entomology, places tke anal odor-
iferous glands described by Dufour among the repugnatorial

2-Univ. Sci. Bull , Vol. V, No. 18.

316 KANSAS UNIVERSITY SCIENCE BULLETIN.

glands, apparently because he has concluded that all fetid and
anal glands are repulsive.

As far as the position, size and structure of these glands are
concerned, these earlier observers are on the whole correct.
They agree, too, in the main points, Dufour has the glands
attached to the rectum, while Fenard has them attached to the
genital duct. The explanation of this difference of observation
is partially suggested by Fenard, when he says: ''En somme
ces organes paraissent d'eboucher dans une sorte de cloaque ou
arrive I'oviducte." The mole crickets have but a single opening
at the posterior end of the abdomen ; and a short common duct
carries the genital and excrementary products. This should
very properly be called a "cloaca." Into this cavity the short
ducts of the anal glands empty.

Fenard gives as the sizes of the glands "about six millimeters
in length and three millimeters in thickness." I found none
as large as that, but the size would depend in part upon the
amount of secretion in the gland. Both Dufour and Fenard
speak of two lobes and a median constriction. There is some
tendency for such a constriction to show, but it is not constant.
The shape and position of the organ would depend somewhat
on the amount of extension of the abdomen and the full-
ness of the rectum. The two lobes when present do not differ
in histological structure, and not in function, as Fenard has
shown by his careful work by means of sections. The
walls are resistant, the cavity large, and the contents appear
homogeneous, granular, and they coagulate as a result of fix-
ation, and color strongly whenever stained. All these facts
Fenard has correctly described.

But Fenard must have worked with preserved specimens
only, or he would not have made the error concerning the func-
tion of the gland. Although he quotes Dufour, he cannot have
followed his suggestion when the latter says : "If one seizes
a mole cricket of either sex, it squirts from the anus a brown
liquid of nauseating fetidity. This liquid is formed in part by
excrement from the rectum and is in part the product of a
special secretion."

I have studied Scapte^iscus didactijlus from Porto Rico and
Gryllotalpa borealis taken in northern Indiana and in eastern
Kansas. My observations and experiments show that the above
quotation is correct in most parts. If the insect is held or irri-
tated in the region of the head or thorax, there is no discharge.

BAUMGARTNER: OBSERVATIONS ON THE GRYLLID^. 317

But if held or pinched or pricked or chemically irritated on
any side of the posterior part of the abdomen, or on the hind
legs, there is always ejected from the anus a bluish-white
liquid with some excrement. The discharge is directed as
nearly as possible to the point of attack, be it above, below,
behind, or on either side of the abdomen. It is driven with
considerable force, enough in some instances to carry it across
an aquarium eight inches in diameter. After several ejections
there is less excrement in the liquid, which becomes almost
colorless, losing its milkiness.

The ejected mass has a very fetid odor and is very sticky,
so sticky that a half-grown nymph can readily be suspended
by lightly touching a needle to some of the secretion and then
to its abdomen. An adult female, in spite of her strong legs,
was held for nearly a minute as a result of touching her be-
smeared body against the side of the jar.

In some breeding experiments reported elsewhere I was able
to study the effect of this ejection and the conditions under
which it is made. There was no discharge when the male was
carefully introduced into the jar with the female, but on one
occasion it happened that the male became excited and rushed
upon the female in his attempt to get away. He received a
discharge upon his head and into his face. He stood for a long
time trying to clean this off. He apparently could remove but
little of it, and died on the second day thereafter. At another
time a female received a lesser discharge from a male. She,
too, spent hours trying to scrape off the sticky stuff, but failed,
and died on the third day. The other pair lived for many weeks
longer. Perfectly calm individuals, when put into a jar in
which there had been a discharge a day or so before, became
very much agitated and tried hard to escape from the en-
closure. This behavior suggests that when these insects get
this odor it warns them that an enemy has been or is near, and
they try to escape. I repeated this test several times with the
same result. I introduced some affected sand into a jar con-
taining a calm individual. He became agitated. In every in-
stance the crickets became excited when they perceived the
odor.

The fetidity of the liquid must repel very ardent pursuers,
and the stickiness must retard them should they become en-
tangled in a discharge. It is, no doubt, for the purpose of
so entangling the enemy that the cricket directs its discharge
toward the point of attack.

318 KANSAS UNIVERSITY SCIENCE BULLETIN.

This defensive organ probably explains the fact that mole
crickets have so few natural enemies, as reported by Bar-
rett (1).

Since the Gryllotalpidae move most of the time in under-
ground burrows the discharge from the anus would protect
against attacks from the rear. Hence there is no discharge
when the irritation is on the anterior half of the body. The
head and thorax, besides being very hard, are further pro-
tected by the powerful fore legs. The abdomen is compara-
tively soft and without other protection than that described
above.

My observations and experiments prove conclusively that the
secretion of the anal glands, or "prostatic glands" of Berlese
and Fenard, is preeminently protective, as any one who will
take the trouble to secure a live specimen and repeat these tests
can see for himself. Neither Berlese nor Fenard can have
handled live individuals, or they should have seen the use of the
anal secretion.

As far as we know no other orthopteran has these protective
glands, nor has it the same peculiar habits. The mole crickets
running along the narrow underground tunnels have the soft
abdomens constantly exposed to the attacks of enemies which
they cannot see or perceive, so they have developed a special
organ which can instantly repel or retard a pursuer.

SUMMARY.

1. The species of Gryllus in eastern and central United
States are not distinct, but form one large intergrading series,
as Lutz has shown. This is true also for the supposed dis-
tinguishing st7'aiv and dark colors as shown by the specimens
collected in Texas.

2. The female mole cricket has a partially developed chirp-
ing organ on its elytra. With this instrument it produces a
single note used as means of recognition in the dark tunnels.

3. The anal gland of Dufour, the prostatic gland of Berlese
and Fenard, is protective in function. The secretion operates
as a repellant by its fetidity, and as a retardant by its
stickiness.

4. Both the female musical organ and the protective gland
are adaptations to life in underground tunnels.

University of Kansas,
May 10, 1909.

BAUMGARTNER: OBSERVATIONS ON THE GRYLLID^. 319

BIBLIOGRAPHY.

1. Barrett, O. W., 1902. The Changa or Mole Cricket in Porto Rico.

Bulletin No. 2, Porto Rico Agricultural Experiment Station.

2. BAUMGARTNER, W. J., 1902. The Spermatid Transformation in

Gryllus assimilis with Special Reference to the Nebenkern. Kan.
Univ. Sci. Bui., vol. 1, No. 1.

3. BAUMGARTNER, W. J., 1904. Some New Evidences for the Indvidual-

ity of the Chromosomes. Biol. Bui., vol. VIII, No. 1.

4. BAUMGARTNER, W. J., 1905. Some Peculiar Habits of the Mole

Crickets. Science, N. S., vol. XXI, No. 544.

5. Berlese, a., 1882. Ricerche sugli organi genitali degli Ortotteri.

Atti della R. Accad. dei Lincei. Ser. Ill, vol. XI.

6. Blatchley, W. S., 1902. The Orthoptera of Indiana. Twenty-

seventh Annual Report of the Department of Geology and Natural
Resources of Indiana.

7. Comstock, J. H. and A. B., 1895. A Manual for the Study of the

Insects. Ithaca, N. Y.

8. DUFOUR, Leon, 1841. Recherches anatomiques et physiologiques

sur Orthopteres, les Hymenopteres et les Neuropteres. Memoire
des Savants etrangers, Paris.

9. Fenard, a., 1897. Recherches sur les organes complementaires in-

ternes de I'appariel genital des Orthopteres. Bull. Sc. France-
Belg., vol. XXIX.

10. Hart, Chas. A., 1892. On the Species of CEcanthus. Ent. News,

vol. III.

11. Henneguy, L. Felix, 1904. Les Insectes, Morphologie, Reproduction,

Embryogenie. Paris.

12. Lacordiare, M. Th., 1834. Introduction a L' entomologie. Paris.

13. Lang, A. Textbook of Comparative Anatomy. (Translated by H. M.

and M. Bernard.) London.

14. Lutz, F. E., 1908. The Variation and Correlations of Certain Tax-

onomic Characters of Gryllus. Carnegie Institution of Washing-
ton, Publication No. 101.

15. Packard, A. S., 1893. Textbook of Entomology. London.

16. Saussure, Henri de, 1872-'78. Melanges Orthopterologiques. Geneva.

17. SCUDDER, S. H. Notes on the Stridulation of Some New England

Orthoptera.

18. ScUDDER, S. H., 1901. The species of Gryllus on the Pacific Coast.

Psyche, vol. IX.

19. ScuDDER, S. H., 1901. The Species of Gryllus Found in the United

States East of the Sierra Nevadas. Psyche, vol. IX.

a-Univ. Sci. Bull., Vol. V, No. 18.

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THE

KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 19 — March, 1910.

(Whole Series. Vol. XV. No. 19.)

CONTENTS:

Cbservations on the Gryllid^: IV. Copulation, W. J. Baumgartner.

PUBLISHED BY THE UNIVERSITv,
LAWRENCE, KAN.

October. 1911.

Entered at the post-office in Lawrence as second-class matter.

3310

THE KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 19.] MARCH, 1910. [vo'^xVNrif

OBSERVATIONS ON THE GRYLLID^: IV.
COPULATION.

BY W. J. BAUMGARTNER.

(Contril)ution from the Zoitlogical Lal)oiatory, No. 196.)
Plate LXIV.

Historical Note.

'' I"* HE first suggestion of the correct method of the transfer
I of the spermatozoa from the male to the female in the
Gryllidse is given by Siebold (20), who described the
glands annexed to the ejaculatory duct and noted that then-
secretion coagulated very readily when exposed to the air, and
that it probably served to fill and distend the copulatory pouch
just as it filled and distended the penis. The latter organ sur-
rounded a portion of the sperm and thus formed the spermato-
phore. Siebold believed with the older naturalists that the
spermatophore is the end of the penis broken off and often re-
formed.

Stein (22) by his careful investigations gave the correct idea
of the spermatophore as a vesicle formed by the secretion of
the annexed glands and used to carry the sperm. Siebold
(21) accepted this correction and later defended it. But he
studied mostly locustids, and so we do not know that this part
of his notes concerned the gryllids ; however, some of his work
was done on Gnjllus, and so the work must be mentioned in a
review of the subject.

We do have a comparatively detailed description of the act
or acts of copulation in the crickets as early as 1855 by a
Frenchman, M. Charles Lespes (11). But the work was little
known and then questioned by Milne-Edwards (15) ; so that

(323)

324 KANSAS UNIVERSITY SCIENCE BULLETIN.

textbooks of to-day still disregard it, or quote it with a ques-
tion mark. Since I shall use Lespes's work very freely in my
own description, I shall not review it farther here. Suffice it
to say that the observations are strikingly keen and correct
for having been made so early, and they should long since
have been inserted in our texts.

Leydig (13) in his fine work, dealing more with the histo-
logical side of the subject, says that in many invertebrates the
accessory sex glands form a spermatophore.

Milne-Edwards (15), in 1870, said, speaking of the sper-
matophore mentioned by Stein and Siebold : "It is possible
that it may exist in certain cases, but in other cases it is evi-
dent to me that the appendage in question was certainly a
part of the penis." Of Lespes's statements he writes: "It
does not seem to me sufficiently demonstrated that the so-
called spermatophore is not the terminal part of the penis
which in copulation is broken off and remains inserted in the
female apparatus, as it does so often in the case of other
insects."

Milne-Edwards's prominent position, no doubt, gave his
work the preponderance, and so we find Lespes's work hardly
ever quoted. To show how it was disregarded, let me quote
from Girard, who wrote three large volumes on entomology.
In 1873 (7) he said of the glands joined to the vas deferens,
"They pour in a liquid to delay or modify the sperm." In the
third volume, in 1885 (8), he repeated the above statement
about delaying the sperm, and then adds, "or perhaps to give
more activity to the sperm, which are almost immotile in the
testis."

Berlese (-4) studied the external and internal genital organs
of various orthopteran species. In the body of his work he
does not seem to realize that the Gryllidse differ from the other
Orthoptera, and so he describes in Gry litis a penis and all the
other accessory parts, without any thought of their correct
function. In an appended "Physiological Note" he comes
nearer the truth. He says, concerning copulation, that fertili-
zation is internal and that copulation in the Acrididae and per-
haps in the Mantidse is by means of a penis, the act lasting a
relatively long time; while in the Locustidse and Gryllidse,
"the male mounts on the back of the female, the penis being
inserted in the vagina and the sperm injected in a short time."

BAUMGARTNER: COPULATION IN GRYLLID^. 325

He supposes that the mucus secretion of the annexed glands
serves to lubricate the organs. He notes the fact that in
Gryllus and Gryllotalpa the mucous secretion flows out through
the same duct with the sperm. He adds further: "In Gryllus,
I think, fecundation is more complex, occurring by means of a
special organ which is left in the vagina of the female and
which can be reformed up to three times a day, as I myself
have seen. This singular organ holds the sperm in its interior
and is a true spermatophore." The only way I can explain
how Berlese came to overlook the discrepancy between his
description and his "Physiological Note" is to think that in
the body of the paper he was concerned with the homology
of the parts in the various groups of Orthoptera, and so did not
think of the function.

Graber (9) shows that in many Orthoptera there is no true
copulation, but that the sperm are transferred by spermato-
phores. Palmen (17) studied by means of dissection and sec-
tions several species of Orthoptera, and he confirms Graber,
showing the absence of a penis and the correct method of
spermatozoon transfer.

Peytoureau (19) in his long thesis on the "Morphology of
the Genital Armature of Insects," does not even mention
Lespes's work, again showing how little this early, accurate
investigation is known ; or if known, how little it is appreciated.
Peytoureau worked on the mole crickets, some long-horned
and some short-horned grasshoppers. His special study was
the hard parts, and he showed that the penis is absent, ex-
cept in the Acrididse, the Forficulidse and the Ephemeridse. But
his study was centered about the homology of the male and fe-
male hard parts. Questions of the number of abdominal seg-
ments, the relative position of the ovipositor and the penis and
to what segments these belonged, were the ones which he con-
sidered; and so the only observation of importance bearing
upon our topic is the absence of the penis in the Gryllidse.

Fenard (6) in 1896 published a long paper on "Les Organes
Complimentaries Internes de L'appariel Genital des Orthop-
teres." He studied both sections and dissections of the an-
nexed glands in the two sexes of several genera of most of the
families of Orthoptera. In his observations he is correct on
the whole. But in his interpretations he is not always so
fortunate. The errors made are due largely to the fact, I

326 KANSAS UNIVERSITY SCIENCE BULLETIN.

think, that he worked with preserved specimens only, and so
missed the function of parts as proven by the habits of the ani-
mals. The suggestion that the anal gland in the mole crickets
secretes a lubricant for the genital organs is entirely wrong, as
I have shown elsewhere (2) . Concerning the glands annexed to
the ejaculatory duct he reasons correctly, as follows: Gryllids
and locustids form a spermatophore, as shown by Siebold
{21), Lespes {11) and Berlese (4). But these families are
the ones that have this mass of annexed glands; hence these
glands secrete the substance for the spermatophore. He
thinks there must be a spermatophore formed in Gryllotalpa,
although it has never been seen because of the habits of these
insects. In this surmising he was fortunate, as I show below.
But he is mistaken in the function of what he calls the ear-
shaped gland. He thinks that it is analogous to the anal
gland of Gryllotalpa. This is surely wrong. The position of
the pair of glands and its relation to the excretory duct is such
that it can have no such function as I have found for the anal
glands of the mole crickets. My many dissections and obser-
vations lead me to believe that there is no gland which secretes
a lubricant for the sexual organs.

Although the method of copulation by spermatophore and
the function of the annexed glands have been so well described,
it is rather disappointing to see our textbooks still quoting
Carus's figures showing that the male generative apparatus of
Gryllus is among the simplest found in the insects. See
Henneguy {10) et al. Although this Frenchman cites Fen-
ard's paper in his bibliography, he makes no reference to his
findings, but says of the annexed glands : "Files ont ete peu
etudies jusqu' ice." But under the topic of "Copulation" he
gives the process in Gryllus in detail, quoting Lespes's descrip-
tion and some of his figures of the spermatophore. In this
case he does better than our own Packard {18), who, in speak-
ing of the spermatophore, says : "In Locusta, and perhaps
also in Gryllus, the sperm is enveloped by the secretion of the
accessory glands of the seminal duct." This "perhaps" is
very striking when one considers how easy it is for any one to
observe copulation in our common field cricket. One need
only catch an individual of each sex during the height of the
breeding season, as indicated by their great chorus of chirping,
and place them with a little grass over night or longer into

BAUMGARTNER: COPULATION IN GRYLLID^. 327

separate jars. Then slowly turn the glass with the male over
that with the female. In a comparatively short time the male
will begin his chirping and one can see the whole series of
acts of courting and copulation. After observing the process
one can hardly prevent a smile when he reads Loeb's {H)
quotation of Yersin's experiment of having a pair of decapi-
tated crickets copulate, and then see the admonition : "Of
course it was necessary to place the male 07i the female." One
would hope that the observations in the experiment itself, and
any conclusions drawn therefrom, may be more to the point
than is the self-evident suggestion added at the end.

DESCRIPTION.

A free translation of Lespes's paper of 1855 would describe
fairly correctly and with a good deal of detail the various steps
in the process of copulation. He notes at the beginning that
there is no true copulation, but that the male simply deposits
a spermatophore into the posterior end of the abdomen of the
female ; and that this process may be repeated many times.

The Courting.

Lespes (11) studied first the field cricket. Catching speci-
mens of both sexes, he got them accustomed to their new sur-
roundings and then brought the two together. "The male
soon began to chirp, and to move around the female. Ap-
proaching nearer and continuing his chirping he turned his
abdomen, carried very low, toward the head of the female.
She remained quiet for ten minutes, not appearing to notice
the maneuvers of her mate; then she moved forward a little
and began to caress his abdomen with her mouth parts. Pres-
ently she mounted partly on his back; and he, stopping the
chirping, glided back under her. The female vulva now rested
above the end of the male abdomen."

This description of the courting is fairly exact. The caress-
ing by the female is not essential or even usual. Sometimes the
female hardly stops feeding during the whole process, simply
allowing the male to slide under her. When the male ap-
proaches the female, or when she approaches him, his chirp
becomes much softer and less shrill. On some occasions if the
female did not yield after a certain amount of chirping the
male became angry and fought his mate.

328 KANSAS UNIVERSITY SCIENCE BULLETIN.

The Transfer of the Spermatophore.

Lespes says : "The plate, the tenth tergite, which covers the
opening of the anus and the pieces of the genital armature,
was raised, and a hook of this apparatus penetrated into the
vulva. Immediately after this the end of a small brown body
directed by the hook was introduced into this opening. The
female then left the male, carrying the small brown body of
which I have spoken." Lespes then removed the small appa-
ratus from the female quite easily, as only its front end was in-
serted in the vulva. This apparatus he found composed of a
small hard ampulla and a thin, transparent plate curiously
twisted. Upon examining a jar in which he kept a male over
night he discovered that there was a spermatophore lying on
the bottom. He later proved that this was not exceptional, but
that males deprived of females rid themselves of their sper-
matophores.

The male which had copulated with the female seemed ex-
hausted and did not renew his courting for several hours, al-
though he was again put in with the female. Lespes observed
the transfer of the spermatophore very often, and each time
he got a spermatophore from the female, which he studied
later.

Taking up this transfer in detail, the Frenchman describes
the position of the spermatophore in the male. "It lies in the
posterior end of the abdomen covered by the supra-anal plate.
To see it in situ, gently raise the anal plate which covers the
anus; below this there is a horny plate with three hooks pro-
jecting backwards and upwards, and below this there is the
spermatophore lying in the concavity of the eighth sternite.
The ampulla extends backwards and is supported on either
side by fleshy pallets (plate is used by Lespes) . The plate and
the thread of the apparatus are still in the organs which formed
them. At the moment of copulation the supra-anal plate is
raised and the hooks are introduced into the vulva, holding the
two individuals firmly together. The end of the spermato-
phore plate from which projects the thread glides in a kind of
hollow groove in behind the hook in a manner analogous to
that which surgeons use when they introduce a bistoury upon
a groove-director. By a rapid movement from before back-
wards the plate is fastened and the two pallets which hold the
ampulla move aside, and so the spermatophore is carried by the

BAUMGARTNER: COPULATION IN GRYLLID^. 329

female, held by the plate only. Ten minutes after copulation
some males had formed a new spermatophore, but it was soft
and white, and attained its hardness and color only after an
hour. The spermatophore can be quite readily removed from
the male by means of a forceps. To form a new spermato-
phore the male goes through some motions similar to those
used in defecation. The female carries the spermatophore for
some time, in one case six hours, and it was not dropped until
she mounted a male again."

My own observations show that this account is quite cor-
rect in most of the details. Our common black field cricket
differs in the following points : The female if undisturbed
will not leave the male immediately after the placing of the
spermatophore. She may remain perfectly quiet for several
minutes.

A third element of this sperm-bearer, which Lespes seems to
consider of little importance, is very essential, as I shall prove
below. It is the thread which projects beyond the plate. (See
fig. I.) This thread slides in the groove when the apparatus
is being introduced. The mechanical process can be better
understood by examining figure 5. The ampulla lies between the
two pallets, which are then extended backward very far. The
plate lies in the anterior part of the mold and the thread lies
in the groove below the hooks. In transferring the spermato-
phore the male turns the hooks upward by swinging them, or
rather the whole mold, at its attachment to the tergite as a
pivot. By this turning the hooks first extend upward and then
forward and are thus inserted into the vulva of the female,
holding her firmly. Then by means of the muscular mass be-
hind, the thread and plate of the spermatophore are pushed far
into the vulva. The hooks of the plate hold the apparatus and
the thread carries the sperm into the spermatheca. The
thread and the anterior end of the plate are quite flexible, and
so they are bent and directed by the "independent piece with
the short stylet" which guides the thread into the duct leading
to the spermatheca. The "forward and backwards" movements
are concerned with the proper placing of the thread.

The time suggested for a male to copulate a second time is
much too long. I have had a second pairing occur in fifteen
minutes. The spermatophore in this case was whiter and
softer than usual, but as far as I could see it was a normal

330 KANSAS UNIVERSITY SCIENCE BULLETIN.

copulation. This intervening time was unusually short,
but I am sure that it does not take an hour for a spermato-
phore to harden after it is formed.

I am sure that the males of ou)- field crickets rid themselves
of the spermatophore, if deprived of the females, only ex-
ceptionally. In all my many observations I have seen the thing
occur but three times, twice in Gryllus and once in Nemobius.
In every case it was rubbed ofl[' after an unsuccessful attempt
to introduce it into the female. In these trials the plate was,
no doubt, partially removed from its normal position in the
forming organs, because in most cases the males made no
effort to remove the spermatophore, although they had tried
to copulate.

I have not seen that the males make any movements while
forming a spermatophore, but I would not positively deny that
they do so. I have frequently kept them under observation
for a few hours after copulation ; some would form a new
sperm bearer and others not. One male copulated a second
time after a very short interval, but I did not watch him mean-
while. Some other specimens did not have the spermatophore
formed after hours of watching.

The female carries the vesicle very frequently until she is
about to mate again. If this comes soon after a previous copu-
lation, she will remove the vesicle. She does this with her
mouth parts, bending the abdomen ventrally. She may place
herself partly on her back and holding the abdomen against
the ground force her mouth parts back so as to reach the
ampulla of the spermatophore. If the abdomen is too much
distended by eggs, she frequently rubs it off by dragging the
abdomen on the ground. The longer a spermatophore has been
carried by the female the easier it is removed. In only one
instance did I see a female mount upon a male with the sperm
bearer still in place. After a good many efforts the male suc-
ceeded in pushing the old spermatophore partially out of the
way and placing a new one. The female then carried both,
the old one apparently hanging on by one hook.

The Spermatophore.

Lespes says : "The spermatophore of the field cricket is com-
posed of a vesicle almost round, of a brown, more or less dark,
color. It terminates at one extremity by a whitish papilla
and is continued on the other end by a transparent plate, al-

BAUMGARTNER: COPULATION IN GRYLLID^. 331

most quadrilateral, formed by a thin membrane stretched over
three small cartilaginous pieces. One of these is median and
connects directly with the vesicle. It is tubular and contains
a horny thread which continues well beyond the plate. The
other two, situated to the right and left, are arch shaped, and
their two ends covered by the membrane, form on either side
two tooth-like hooks which serve to fix the apparatus in the
vagina. The size of the spermatophore varies a little but
ordinarily it attains almost 4 mm. from the papilla to the end
of the plate. The thread appears longer or shorter according
as it projects more or less from its tube. The spermatophore
is of different consistency in different parts. The vesicle is
extremely solid and its walls are very thick, while the plate
and its cartilaginous thread are soft at the moment of
copulation ; besides, it is covered by a white, thick liquid which
presents to me all the characteristics of a sperm fluid. The
thick, hard vesicle, composed apparently of double walls, to-
ward the extremities is hollowed out at the center. The round-
ish cavity is full of sperm. At one end a blind tube projects
from the cavity into the ampulla. At the other end the cavity
continues as a straight canal through the whole length of the
plate. This canal contains the horny thread.

"The white liquid which covers the plate when first removed
from the female shows, when examined with a high magnifier,
a large number of small, thread-like zoosperms. They are
about .04 mm. long and .002 mm. thick. These same bodies in
great number fill the cavity of the vesicle. They were never
united into feather-like bundles, nor did they ever show any
movement. The fluid taken from the testis of the male or the
copulating pouch of the female contains many very similar
zoosperms. These never exhibited movement, however
treated. Put into water, some of them twisted themselves
into knots.

"When the female lets the spermatophore fall the walls of
the vesicle are slightly ridged, but it still contains some sperm.
The horny thread is no longer found in the tube of the plate."

I have quoted the above description at length because it
gives many facts as I find them in our American species.

The vesicle and the plate are similar to those described
above (see figs. 1 and 2). The thread is not contained in the
tube, but it is the tube itself continued, and it is much longer,
and bent back so as to run parallel with the plane of the plate

332 KANSAS UNIVERSITY SCIENCE BULLETIN.

and vesicle (see fig. 1) . It is a hollow tube for its whole length
and the sperm comes out at the end of the thread, and not at
the end of the plate, as Lespes supposed. The thread varies
in length not because "it projects more or less from the tube"
but because it is broken off in removing it from the insect.
Many of those that I secured at first had more or less of the end
broken off. It is easier to get a spermatophore with a complete
thread from the male than from the female, especially if the
latter has carried the apparatus for some time. This no doubt
accounts for the fact that the apparatus can be more easily re-
moved after it has been carried for a while. When the insect
drops it normally the thread has been broken off, or, more
probably, partly dissolved away; but it is not entirely gone,
as Lespes states.

The vesicle is apparently double-walled as the Frenchman
observes (figs. 1, 2, and 8) . The inner one is thicker and more
granular, while the outer one is harder and quite transparent
(figs. 1 and 2). It is the inner wall that continues forward
and forms the thread, although this substance is less granular
than that around the vesicle. The substance forming the
plate is somewhat granular, and in some cases took the stain
just like the inner coat of the vesicle. In the lateral parts of
the plates I saw in several of my preparations many vacuoles.
The thread in many of the specimens fixed and mounted in
balsam shows a tendency to break away from the plate (fig. 8) .
That the sperm pass out at the end of the thread was nicely
shown by several specimens which I placed into a normal salt
solution. The sperm flowed out very regularly and moved
about for a time. In one instance it took about fifteen minutes
for a vesicle to empty itself through the end of the thread.
After the sperm were all out a little fluid with small granules
flowed out. I do not agree with Lespes when he says that
some sperm remain in the vesicle normally; nor do I think
that the wrinkling is a characteristic of the empty vesicle. I
could not see any difference between an empty spermatophore
and a dark brown one which had been carried by a male for
some time. The wrinkling, which Lespes correctly describes,
is due to the longer hardening of the vesicle and does not seem
to affect the cavity on the inside.

The shape of the cavity differs from that described and
figured by Lespes in the total absence of the blind tube extend-
ing into the papilla. What he describes is only the second layer

baumgartner: copulation in gryllid^. 333

of the wall of the vesicle as shown in figures 1 to 3. The dark
stain here shows the outer limits of the inner lining, and the
cavity is on the inside, as shown in the outline drawings. The
cavity is easily visible in the preparations but the photos do
not bring it out. At the posterior end the cavity is flattened
and at the other end it gradually tapers down to the size of the
thread; so that the shape of the opening is much like a flat-
bottomed flask. In this cavity all the sperm have their heads
directed toward the openings into the thread and their tails
toward the papilla. Figure 11 is a small group of sperm
photographed from a section of a spermatophore. In passing
out of the thread the sperm manifested some movement of
their own, but were not very active. Several may pass out
together but only two or three heads have room side by side.
The sperm are larger than Lespes thought, as can be seen by
comparing with my earlier published measurements (1).

The Spermatophore-forming Organ.

Since many of our textbooks on entomology — see Packard
{18) and Henneguy {10) — still cite Gryllus as showing one of
the simpler conditions for the excreting ducts of the testes
among the insects, I shall give a more detailed account of the
male generative organs, their ducts and the accessory glands,
than I should otherwise do.

The first description of these parts was given by Dufour (5) ,
in Gryllus campestiis, then by Lespes {11), in G. domesticus,
later by Berlese {^), again in G. campestris, and then by Fen-
ard {6), in both of the above species.

The testes lie on either side of the abdomen above the ali-
mentary canal and extend from the second to the sixth or
seventh segments. The shape is much like that of an elongated
strawberry. On the outside is a thin membrane which encloses
the hundreds of straight or slightly curved tubules which
make up the organ (figs. 6 and 7). Their blind ends are
nearly all directed more toward the posterior end of the body of
the animal. At the center of the testes is a large tube, or
rather an irregular sinus, which receives the openings of the
hundreds of tubules (fig. 8). The small vas deferens leaves
this sinus on the anterior ventral side and comes to the ventral
side of the testes a little back of the middle. It passes back-
ward over the tubules, and leaving them at the outer posterior
edge passes back along the body wall muscles till it reaches the

334 KANSAS UNIVERSITY SCIENCE BULLETIN.

ninth segment, where it passes around the cereal nerve and a
few strands of muscle and ligament (figs. 7 and 8). Making
a sharp bend it goes forward and ventralward. It is very
much coiled upon itself from the bend on (figs. 7 and 8) , The
loops of the coils may lie in planes passing dorso-ventrally or
laterally. The two vasa deferentia join and immediately empty
into the posterior ventral angle of the sinus, which receives
the openings of the annexed glands (fig. 9). Beginning at
the cereal nerve and sometimes a little in front of the bend
the duct is swollen so as to be 0.5 to 0.75 mm. in diameter
(figs. 7 and 8). Fenard says there are from four to five coils
and that the total length is 8 mm. In our common field cricket
I found no regularity about the position or number of coils
and the total length is more than 10 mm.

This swollen part of the vas deferens is always in the mature
adult filled with a mass of sperm, all of them with their heads
turned away from the testis.

Fenard (^) , as well as the older investigators, says that "the
union of the two vasa deferentia forms the ejaculatory duct."
This is the statement generally made of the insects higher in
the scale of evolution ; but it is true only in a certain sense — in
the sense that it serves as a common duct to carry out the
secretions of the testes. First, it is not a true ejaculatory
duct in the sense that it "during coition conducts the sperm
into the copulatory pouch of the female" — Packard {18).
This is true only in those insects with a penis; but in the
Gryllidse, where there is a spermatophore formed there can be
no such function. Second, nor is it true in the sense that it is
formed by a growing together of the two vasa deferentia. It
is formed by an invagination from the outside, as was first
shown by Palmen {17) from the anatomical standpoint and by
Nusbaum {16) from the embryological. This invagination
projects quite far beyond the point at which the two vasa def-
erentia join it (fig. 7). It extends dorsalward and forward,
where it receives the openings of the hundreds and hundreds of
tubules of the annexed glands (fig. 9). The lumen is very
much flattened laterally and widened vertically towards the
blind end. The more muscular tube leaves the sinus at the
posterior dorsal angle just above the point of entrance of the
vasa. This is well shown in figure 9. The openings of the vasa
cannot be recognized in this figure, but the serial sections leave
no doubt as to its location.

BAUMGARTNER: COPULATION IN GRYLLID^. 335

The hundreds of glandular tubules which empty into the up-
per end of the common duct vary greatly in length and di-
ameter, apparently depending in part upon the room they find
for development (figs. 6, 7 and 9) . They take up all the avail-
able space, completely surrounding the seminal vesicles, ex-
tending far backward beside the rectum and the spermato-
phore pouch, or crowding forward and upward around the
intestinal tract. Many of them are short while others are long.
The whole group of tubules forms a large mass which takes
up quite a space (fig. 6).

The "prostatic gland," first noticed by Berlese and later
described by Fenard, I found "only after much difficulty," as
the latter says. From my dissections I had concluded that the
part so described was a part of the irregular pouch which
molds the plate of the spermatophore. But a careful study of
serial sections reveals the presence of a pair of oval glands or
pockets attached on either side to the common duct. They
have very short connecting tubes and they lie in the muscular
and connective tissue surrounding that part of the common
duct between the ganglion and the mold. The glands are not
surrounded by "yellowish fat," as Fenard suggested; but they
have "thin walls and are filled with a clear fluid." This fluid
seems to harden into a clear yellowish chitin-like substance
when treated with fixatives. In this it resembles the secretion
from the annexed glands; but their secretion is darker and
more granular. Judging from the behavior of the secretion
and the position and connections of these glands I think they
do not furnish a lubricant, but probably have something to do
with the formation of the more flexible thread and the plate.

The common duct, after passing close to the floor of the ab-
domen, turns dorsalward and then empties into the cavity of
the spermatophore mold (fig. 9). This apparatus was care-
fully studied by Lespes {11). Many of his observations are
keen and to the point, but his description in some points is
not very clear. Berlese (^) studied this structure too, but he
could not get away from the idea that he was dealing with the
ordinary structures, so he applies the name "penis," etc., to
the parts and so makes his description misleading. Fenard
{6) pays no attention to the hard structures and so he does
not make out the relations and functions of parts in this region.
Peytoureau {19) studied mostly only the morphological rela-

336 KANSAS UNIVERSITY SCIENCE BULLETIN.

tions of the hard parts and so he does not explain the functions
of the parts of this organ.

Lespes describes this apparatus "charged with the formation
of the spermatophore" as follows :

"It is situated at the posterior part of the ejaculatory duct
and may be considered as a part of the genital armature. In
the field cricket this armature is formed, first, by a curved plate
composed of many pieces extending to the right and left and
united by a tough membrane ; and second, by a very remarkable
independent inferior part. The plate itself is composed of six
solid pieces. Above is a kind of shield-like piece which is
easily divided along the median line. Posteriorly it ends in
three hooks, of which the middle one plays an important part
in copulation. To the right and left are long slender rods upon
which are inserted the fleshy pallets which support the vesicle
of the spermatophore. Below this shield are two others, thin
and curved, with an irregular contour. Finally, on the inferior
face of the dorsal shield are two small, slender plates. During
copulation the median hook only enters the vagina and serves
to conduct the plate of the spermatophore. The independent
piece is formed in front by a sort of short stylet which is ap-
plied to the inferior face of the plate of which I have just
spoken. At the base this stylet enlarges, changes its nature
and forms a white plate with cross striations. This plate is
bent along its length, forming an irregular circle. It is swollen
into a very thin vesicle situated under the genital armature
and in front of the two pallets which support the ampulla of
the spermatophore. This pocket is largely open in front and
under the genital armature. The ejaculatory duct, after
having passed around the lower part of the spermatophore
pouch, ends near this opening.

"This white plate is the organ for the production of the
spermatophore. On its convex surface is a transparent line
in which is formed the tube which holds the thread of the
spermatophore. The size of the plate is greater at the two
ends. At the terminal part of the enlarged inferior portion
ends the ejaculatory duct."

Lespes claims to have seen the spermatophore while it was
forming. "It had a very thin membrane. When completely
formed the vesicle was removed out to the place between the
two fleshy pallets, and the plate and the thread remained in

BAUMGARTNER: COPULATION IN GRYLLID^. 337

the forming organ, that is, in the stylet of the armature, until
given off to the female."

This description is somewhat indefinite because of the un-
certain meaning of the phrase "genital armature." Besides,
the application of the words "in front" and "behind" is not
certain. I do not believe that the differences between Lespes's
description and my observations can be accounted for by sup-
posing such differences exist in the different species studied.
Both of us used Grylhis domesticus some, and as shown below
we found only small minor differences between the house
cricket and our own species of field crickets.

Before discussing the formation of this peculiar spermato-
phore farther let me state as clearly as possible the mechanical
problem involved. There is a mold made up of a system of
bracing rods of chitinous material and a thin fibro-muscular
membrane. Into this mold leads a single tube, which has to
bring the sperm and the secreted fluid for the covering. The
ampullar part of the finished spermatophore has apparently
a double wall, and so there may possibly be two kinds of secre-
tions, although a careful study of the gland shown in figure 10
makes me doubt it. The question is, then : How are the sperm
kept separate, and especially how are they held in the center
while the soft liquid is poured around them and hardens? If
there is a second layer in the wall the incomplete ampulla would
have to be held in the center while the second inflow and hard-
ening occurs. To place a semifluid substance within another
fluid substance and hold the former at the center while the
latter hardens would be a difficult feat for any mechanic.
I do not see how the apparatus provided is sufficient to ac-
complish the result. Yet I have seen so many well-formed
spermatophores that there can be no doubt of the end product.

The way to solve the problem readily suggests itself. One
needs to get a lot of specimens showing the various stages in
the formation of the vesicle. Such a series of stages I have
been hoping to get, but have thus far not succeeded. In all
my many dissections I have never found a partially formed
vesicle. It is hardly feasible to follow the process in a living
specimen, as the attempts to make observations would interfere
with the normal formation.

Although I am not able to give a description of the exact
method of formation, I shall nevertheless make some sugges-

2-Univ. Sci. Bull., Vol. V, No. 19.

338 KANSAS UNIVERSITY SCIENCE BULLETIN.

tions. Lespes says that the ampulla is formed and then moved
out to its position between the two pallets. I think that this
cannot be correct, as it would mean that the plate and thread
would have to be joined onto the ampulla. In all the specimens
I have seen the inside cavity and the walls are just as smooth
as they can be. I cannot imagine that if the apparatus is
formed in two parts they could be joined together so exactly
and smoothly. I would rather believe that the structure is
formed as a whole in situ. The thing that gave the idea that
it moved back is the fact that when the wall has partly hard-
ened the pallets retract a little, exposing the ampulla.

When there is no spermatophore in a male cricket the hooks
are drawn far down, the pallets are much contracted and
folded, and a part of the loose tissue on the floor of the cavity
is rolled up and fits into the mold as a sort of plug. The pallets
and the plug have deeply grooved and ridged surfaces which
line the small lumen within.

Things occur about as follows when a spermatophore is
formed : The inner lumen becomes a little larger and assumes
the shape of the cavity of the vesicle. Then the sperm pass
down, filling it up. It is possible that the lining cells secrete a
small amount of substance to form a thin membrane, and
Lespes may have found an example in this stage. If a mem-
brane is secreted it cannot be discerned later. Now some secre-
tion comes down the common duct. This flows in and fills up
the grooves. The ridges withdraw, forming new grooves to
be filled up. The process continues till enough is laid down
to form the inner layer of the wall. Then the flow ceases for
a while, and the secretion hardens somewhat. After a longer
or shorter time a second inflow of secretion occurs, and it takes
the place of the withdrawn ridges till the surface of the lining
membrane becomes smooth, thus forming the outer wall of
the ampulla. Then the whole hardens into the completed sper-
matophore. I think the whole takes place in a short time rel-
atively. There may be movements accompanying the process,
although I have not observed them.

Since there is but one duct, the only other assumption which
could be made is that the secretion and the sperm come down
all together instead of in succession. If this be correct, then the
long sperm would have to wiggle their way to the center. This
is hardly possible. Besides, it would be difficult to explain the
second layer in the wall of the ampulla by this hypothesis.

BAUMGARTNER: COPULATION IN GRYLLID^. 339

Another mechanical problem appears in the study of the
spermatophore, namely, What causes the sperm fluid to flow
out of the ampulla? Are they forced by contraction of the
walls, or is there some substance that swells and forces them
out? Does air penetrate the wall in some way and thus prevent
a vacuum?

I do not believe that there is any contraction ; at least not
much. The walls do not collapse. Farther than this I could
find nothing that would throw any light on the problem.

Differences Found in the Domestic Cricket.

Lespes describes the spermatophore and the spermatophore-
forming organ of this species, but says that he has not seen the
act of copulation. I can readily believe the latter statement,
as the insects are much more shy and will not mate readily
when watched. Nevertheless I witnessed the process several
times. The behavior during courting and the movements dur-
ing the transfer of the spermatophore are very similar to
those of the field crickets. The differences are of no conse-
quence.

The spermatophore is lighter in color, as Lespes says. But
this is in accord with the general lack of color in this species.
It is a little shorter and stouter, as the Frenchman indicated.
The plate is also narrower and more bent. But the cavity does
not extend back into the papilla, as I have already explained
about our field crickets.

There is one point to which I should like to call attention.
Lespes does not speak of it, yet shows a difference in the draw-
ings. In the house cricket the thread is a direct continuation
of the canal. This is correct for this species, and is precisely
what I find in our field cricket. I have not seen the spermato-
phore in G. campestHs; but judging from our domestic and field
crickets, and the conditions I find in Nemobius, I do not be-
lieve that Lespes is correct when he indicates a break between
the thread and the plate, as he shows in his drawings, figure 2.
The canal could not be continuous if his drawing is correct. But
Lespes thought that the thread was the mass of sperm, and
so not thinking of the thread as a hollow conducting tube he
might have a break in it.

The mold differs from that of our field cricket about as it
differs from Gryllus campestris, judging from Lespes's descrip-
tion. The hooks on the superior plate are rudimentary or

340 KANSAS UNIVERSITY SCIENCE BULLETIN.

absent, while the stylet is well developed and projects far
backward. The cavity is smaller, making it necessary that
the plate of the spermatophore be bent into a smaller curve.
There are other minor differences in size and relations.
Whether these differences are great enough to prevent inter-
crossing of species is not positively settled. I tried some ex-
periments in cross-breeding, in which I was unable to get any
cross matings; but the tests were not extensive enough to be
very conclusive.

IN NEMOBIUS.

This genus resembles rather closely the form which Lespes
describes under the name Gryllus Sylvestris. It is much smaller
than our Gryllus and, like the European form, it likes the
woods or shady places ; in the grass under the trees of a park
I have found the most specimens. In the months of August
and September they are very numerous.

In its courting it is, like the domestic cricket, much more
shy than the larger form, and it is rather tedious to watch till
one catches a pair copulating. In the main the whole series
of processes is quite similar to that in Gryllus. In the courting
the male is more active while the female less readily yields to
the courting. What Lespes (12) says of his forms holds true
for our two genera : "Except some details of form and size
all is similar. The spermatophore is smaller and more fragile
but is similarly composed. The genital armature presents the
same pieces, but they differ much in form." Lespes again in
this species figures and describes the thread as continuous with
the canal of the plate. This strengthens my belief that there
is some mistake in the description of the spermatophore of
Gryllus campestris.

I have succeeded in getting a good photograph (fig. 4), that
shows the structure of the spermatophore of Nemobius fas-
ciatus, our most common species, very well. The ampulla is
almost a perfect sphere. Its wall is thick and its central cavity
rather small and nearly spherical. As in Gryllus sijlvestiis,
the plate is small and far from the ampulla, but the thread and
plate are much more curved in our species (fig. 4). There is a
fine, hollow, tube-like opening leading from the central cavity
to the tip of the thread. There can be no doubt that the sperm
fluid flows out through it.

BAUM GARTNER: COPULATION IN GRYLLID^. 341
IN GRYLLOTALPA.

Fenard (6) cites the fact that the Locustidse and the Gryl-
lidse produce spermatophores and that those two famihes have
the large mass of annexed glands. Continuing, he says : "Then
wherever we find the glandular tubes similar to those in the
above-mentioned groups we have a right to conclude, until
there is proof to the contrary, that they secrete a substance
destined for the formation of spermatophores more or less
complex." He suggests that in Gryllotalpa, whose copulation
has never been seen, fertilization occurs by means of a sperma-
tophore, because they have the annexed glands similar to
those of Gryllus.

In the abstract {2) I made the statement that in the mole
crickets the sperm are transferred by means of a spermato-
phore. In order to observe copulation I kept four mole crickets,
two pairs, for several weeks each in a separate aquarium partly
filled with wet sand, grass roots and slices of potatoes. For
five nights I placed them by pairs into battery jars which con-
tained just enough sand to allow the animals to make a burrow
around the edge. Thus I could watch them continuously. After
some twenty hours I finally observed copulation. As a result
of these vigils I am able to give the facts concerning chirping
and the protective gland published elsewhere {3), as well as
the following concerning copulation.

The courting is somewhat similar to that in Gryllus. The
male calls the female with loud, long chirps. As she ap-
proaches the chirps become short and much softer. He then
frequently turns his abdomen towards her. As the pair get
ready to copulate the position assumed is quite different from
that of any other animals of which I know. They turn poster-
ior end to posterior end, and ventral side to ventral side, so
that the cloacal openings are just opposite each other. The
female stands erect with her abdomen slightly raised, while
the male lies on his back. The abdomens are tightly held to-
gether by hooks, described with great detail by Peytoureau
{19). The sperm were carried to the female by a spermato-
phore. The time it takes for the transfer is not over a minute ;
but the pair kept their relative position, the abdomens simply
touching each other, for more than ten minutes. After dis-
turbance the male followed the female and again assumed this

342 KANSAS UNIVERSITY SCIENCE BULLETIN.

relative position, but no further transfer of a spermatophore
occurred.

Right after the first separation of the couple the female be-
gan to chew at a part of the sperm vesicle, and only desisted
when disturbed by the movements of the male. She was also
perfectly quiet as soon as her abdomen was touched as in the
position of copulation. One observation is not enough to estab-
lish the purpose of an act; yet, judging from the behavior of
the individuals of this pair, especially from the anxiety of the
male to touch the female with his abdomen, one could easily
conclude that the long lying in the position of copulation was
to prevent the female from chewing at the spermatophore too
soon and thus preventing the proper injection of the sperm.
After the pair had remained in the above-described position
for ten minutes or more the female left the male and mani-
fested no further desire to chew the spermatophore. She
carried it for about a half an hour and then dropped it.

As the vesicle was being transferred, or just after it had
been put in place, there was an outflow of some transparent
fluid on either side of the vesicle. This soon hardened. It
is this part of the apparatus that the female was chewing. The
spermatophore was found to consist of an oval ampulla which
contained the sperm in the cavity at the center. At one end
of the ampulla there is a projection by which the apparatus is
held in the vagina, and through which the sperm are carried
into the spermatheca. On either side of this projection is an
irregularly shaped mass formed by the above-mentioned out-
flowing fluid during the transfer. The two sides are unlike, as
part of one side was pulled and eaten away by the female, and
the other side was pressed out of shape by some falling sand
before it had time to harden. Having but this one imperfect
specimen of the spermatophore of these crickets I cannot give
as accurate a description of its detail as I should like. Suffice
it to say that it has the essential parts — a hollow ampulla to
contain the sperm and a projecting part to fasten it to the fe-
male and to carry in the sperm. The structure of the latter is
obscured by the irregular mass.

The outflow of this fluid recalls an observation which I made
repeatedly in the Chicago greenhouses. In the copulation of
a locustid the same phenomenon occurs, but in a more marked
way. The species, an exotic one introduced from Japan, is

BAUMGARTNER: COPULATION IN GRYLLID^. 343

very abundant in the greenhouses. The male has no chirping-
organs and so the courting is all done by means of the long
antennae. The female mounts on the back of the male, when
he hooks an almost spherical ampulla full of sperm into her
vagina. During the latter act a viscid fluid flows out on either
side and forms two somewhat irregular roundish masses larger
than the original ampulla, which now lies between the two.
Sometimes the female began immediately after copulation to
eat this substance, doing it more readily if the couple was dis-
turbed.

As to a use or meaning for this outflow of fluid and its sub-
sequent hardening it is difficult to suggest a satisfactory one.
Of its regular occurrence there can be no doubt, as I observed
it many times in Diastremmena and once in Gryllotalpa. It
may function as an additional fastener to hold the apparatus in
the vagina. Yet this seems hardly necessary, as the spermato-
phore apparently remains in place before it is hardened. It
may have something to do with the emptying of the vesicle,
contracting around the ampulla, thus forcing out the sperm.
Or it may be concerned with the removal of the spermato-
phore from the vagina, acting as a bait for the female to re-
move it.

SUMMARY.

1. The reproductive apparatus of Gryllus is not of the
simplest type, as was reported by the earlier investigators and
as is still suggested by our textbooks. These organs are rather
of the most complex kind found among the insects. They were
studied and described with a good deal of detail, and the cor-
rect method of the transfer of the spermatozoa was indicated
by Lespes in 1855 ; but the work was discredited and neglected.

2. The male carries the sperm to the female in a special
structure, the spermatophore, formed by the secretion of the
glands annexed to the common duct.

3. The essentials of the spermatophore, as shown by the
study of several genera of Gryllidse, are a hollow vesicle to act
as a retainer, and a hollow thread-like tube which carries the
sperm into the spermatheca of the female. To the tube are
added some enlargements which serve to hold the structure in
the vagina.

4. The so-called genital armature of the male crickets con-
sists of a mold for forming the spermatophore and an appa-
ratus to transfer the same to the female.

344 KANSAS UNIVERSITY SCIENCE BULLETIN.

5. The differences between the field and domestic crickets
is probably great enough to prevent intermating, yet this is not
at all proven.

6. Gryllus has no gland which secretes a lubricant for the
copulatory organs. The so-called "prostatic gland" has to do
with the formation of the thread and plate of the spermato-
phore.

7. Mole crickets form a spermatophore. A transparent
viscid fluid flows out during the transfer. The male and the
female assume a very peculiar position relative to one another
during copulation.

University of Kansas,
March 12, 1910.

BIBLIOGRAPHY.

1. Baumgartner, W. J., 1902. Spermatid Transformations in Gryllus

assimilis. Kan. Univ. Sci. Bull., vol. I, No. 2.

2. Baumgartner, W. J., 1905. Observations on Some Peculiar Habits

of the Mole Crickets. (Abstract.) Science, N. S., vol. XXI, No. 544.

3. Baumgartner, W. J., 1910. Observations on the Gryllidae; III.

Notes on the Classification and Some Habits of the Crickets. Kan.
Univ. Sci. Bull., vol. V, No. 18.

4. Berlese, Antonia, 1882. Ricerche sugli organi genitali degli Ortot-

terri. Atti della Accademia dei Lincei, 1882.

5. DuFOUR, Leon, 1841. Recherches anatomiques et physiologiques sur

les Oi"thopteres, les Hymenopteres et les Neuropteres. Mem. d.
Savants etrangers, VII, Paris.

6. Fenard, a., 1896. Recherches sur les organes complementaires in-

ternes de I'appareil genital des Orthopteres. Theses presentees a
la Faculte des Sciences de Paris. Lille.

7. Girard, Maurice, 1873. Traite elementaire d'entomologie, vol. I,

Librairie J. B. Baillere et fils, Paris.

8. Girard, Maurice, 1885. Traite elementairie d'entomologie, vol. Ill,

Librairie J. B. Baillere et fils, Paris.

9. Graber, 1877-'79. Die Insecten, vol. I-II, Mtinchen.

10. Henneguy, L. F., 1904. Les Insectes, Masson et Cie, Paris.

11. Lespes, Chas., 1855. Memoire sur les Spermatophores des Grillons.

Ann. Sc. Nat., T. III.

12. Lespes, Chas., 1855. Deuxieme Note sur les Spermatophores du

Gryllus sylvestris. Ann. Sc. Nat., T. IV.

13. Leydig, Franz, 1867. Der Eierstock und die Samentasche der In-

sekten, zugleich einen Beitrag zur Lehre der Befruchtung. Nova
Acta Acad. Leop. Car., T. XXXIII, Dresden.

14. Loeb, J., 1907. Comparative Physiology of the Brain and Compara-

tive Psychology, New^ Yoi*k.

15. Milne-Edw^ards, H., 1870. Lecons sur la Physiologie et Anatomie

comparee de I'homme et des animaux, T. IX.

BAUM GARTNER: COPULATION ON GRYLLID^. 345

16. NusBAUM, 1882. Zur Entwickelungsgeschichte der Ausfiihrungs-

gange der sexualdriisen bei den Insecten. Zool. Anz., Bd. V.

17. Palmen, J. A., 1884. Uber paarige Ausfiihrungsgange des gesch-

lechtsorgane bei den Insekten. Eine morphologische Untersuch-
ung. Helsingfors.

18. Packard, A. S., 1903. A Textbook of Entomology. The Macmillan

Company, New York,

19. Peytoureau, a., 1895. Morphologie de I'Armure des Insectes, Paris.

20. SlEBOLD, C. T., 1837. Fernere Beobachtungen iiber die Spermato-

zoen bei wirbellosen Thieren. Mueller's Archiv.

21. SlEBOLD, C. T., 1845. Uber die Spermatozoiden der Locustinen. Nova

Acta Acad., T. XXI.

22. Stein, F., 1847. Vergleichende Anatomie und Physiologie der In-

section. I. Die Weiblichen Geschlechtsorgane der Kafer, Berlin.

3-Univ. Sci. Bull.. Vol. V. No. 19.

THE

KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 20— March, 1910.

(Whole Series, Vol. XV, No. 20.)

CONTENTS:

An Examination of the Chromosomes of Anasa Tristis:

C. E. McCIung and Edith Pinney.

PUBLISHED BY THE UNIVERSITY,
LAWRENCE, KAN.

March, 1912.

Entered at the post-office in Lawrence as second-class matter.

3-3310

THE KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 20.] • MARCH, 1910. [^oTxv.TT,

AN EXAMINATION OF THE CHROMOSOMES OF

ANASA TRISTIS.

BY C. E. M'CLUNG and EDITH PINNEY.

I. Introductory statement.

II. The chromosomes during the spermatogenesis of Anasa, an inde-
pendent study: By the junior author.

III. A comparison of the results of studies upon the chromosomes of
Anasa ; By the senior author.

1. Methods.

2. What is the number of spermatogonia! chromosomes, and of

oogonial chromosomes?

3. What is the number of first spermatocyte chromosomes?

4. What is the behavior of the accessory chromosome and of the

plasmasome in the first spermatocyte prophase?

5. What is the behavior of the accessory chromosome in the first

spermatocyte mitosis?

6. What is the behavior of the chromosomes in interkinesis?

7. What is the behavior of the accessory chromosome in the second

spermatocyte mitosis?

8. What is the final chromosome constitution of the spermatozoa?

I. INTRODUCTORY STATEMENT.
The extensive studies now in progress upon the chromo-
somes of numerous species of plants and animals have led to
the formulation of several working hypotheses of considerable
general importance. In order that these may be of value and
serve as an aid to further progress, it is necessary that the
phenomena from which they take their origin be extensively
recognized and uniformly interpreted. Any conception of the
relations existing between germ-cell organization, as exhib-
ited by the chromosomes, and the structural characters of the
body is now unfortunately delayed by certain disagreements

(349)

350 KANSAS UNIVERSITY SCIENCE BULLETIN.

upon the actuality of the phenomena upon which such inter-
pretations are based.

Fundamentally, these disagreements concern more or less
intimately that view of chromosome organization which is
called the "theory of chromosome individuality." Modem
cytologists are either supporters or opponents of this theory,
and its adequacy or inadequacy will be determined by the
weight of evidence now accumulating. It is, of course, im-
possible to conceive that it will stand unmodified, for it is but
the expression of our present limited knowledge; but it must
now be determined whether these structures of the cell have
persistent individualities or whether they are merely incidental
and inconstant expressions of more fundamental phenomena.
Observed facts must support one or the other of these alterna-
tives, although in doing so they may readily enough modify
the established order.

A considerable body of evidence has accumulated within re-
cent years to support the theory of chromosome individuality
and to associate the development of sexual characters with
particular chromosomes. To this has been added the support
of experimental work on sex determination and the parallel-
ism between the segregation of Mendelian characters and the
behavior of the chromosomes in the maturation divisions of
the germ cells. Much of this cytological work has been done
upon the insects, particularly in the orders Orthoptera and
Hemiptera, and, in the large, there is a strong consensus of
opinion. More recently confirmatory evidence has been fur-
nished from studies upon nematodes, including Ascaris, by
Boveri, upon echinoderms by Baltzer, and upon birds and man
by Guyer. Here and there, however, there are workers who
question the fundamental facts and deny the correctness of the
theories founded upon them.

It is necessary, in order to proceed further with our inter-
pretations, that we reach an agreement upon the observable
facts in our science, and no honest question should remain
without answer.

At present one of J:he most serious diversities of opinion
exists regarding the character and the behavior of the acces-
sory chromosome in the bug Anasa tristis. Because of the ex-
tensive and painstaking studies of hemipteran germ cells by
Wilson, the contradictions and inaccuracies in the studies of

CHROMOSOMES OF ANASA TRISTIS. 351

Paulmier and of Montgomery seemed to be cleared up and the
phenomena in this order brought into agreement with those
of the other insects. To this was added the confirmatory
work of Stevens upon several orders of insects, the study
of Lefevre and McGill on Anasa, and more recently that of
Morgan on aphids and phylloxerans. Directly opposed to all
this are the observations of Foot and Strobell upon A7iasa
tristis, the form studied in detail by Paulmier, Montgomery
and Wilson. Their contention, in brief, is that the number
of spermatogonial chromosomes is even instead of odd, that
there is no dimorphism of the spermatozoa due to the presence
in one-half their number of the accessory chromosome, and
that there is no evidence to show that the theory of chromo-
some individuality receives any support from observed mor-
phological continuity of any of the chromosomes. Aside from
these major questions there are others, such as the history
and fate of the plasmasome, upon which these authors are in
disagreement with other investigators.

The issues here defined are so clear-cut and definite and
their settlement of so much importance that the senior author
of the present paper was led to offer his services for a reex-
amination of the whole question. This offer met a hearty
response from Professor Wilson, who sent not only live mate-
rial, but his entire series of mounted slides, including those
used by Paulmier. Professor Lefevre has been kind enough,
also, to permit the use of his slides. Miss Strobell, writing
for Miss Foot, expressed a lively interest in the proposed set-
tlement of this problem, but was unable to send the prepara-
tions used in their studies, because these had all been de-
stroyed on account of limited storage room. It is to be
regretted that this material is not available for comparison,
since it is upon it that the only divergent observations have
been made; but since specimens from the same locality, pre-
pared by all methods, including the special ones employed
by the Misses Foot and Strobell, are at hand, there can be no
reasonable doubt that it is representative.

To reduce the possibility of the influence of preconception
to the lowest possible degree, the junior author has worked
through the entire problem without knowledge of the work
done by others, and without promptings from the senior
author except such as were given by a series of questions for

352 ' KANSAS UNIVERSITY SCIENCE BULLETIN.

which answers were desired. Independently, the senior au-
thor went over the same ground, first upon material prepared
in this laboratory, and later upon the slides of Paulmier, Wil-
son and Lefevre. Whenever possible, photomicrographs were
made, and representations of them accompany the paper. The
two authors reached concordant results throughout, and their
common experience is embodied in what follows.

Since the matters of dispute are purely of facts and inter-
pretations, it will be necessary to take a definite position on
each of the points involved, but so far as possible the material
evidence for the decisions will be presented fully. It is hoped
that this may be complete enough so that others interested
may be able to form their own opinions directly from the ap-
pearances themselves. There will be presented first the per-
sonal work of the junior author, giving her independent
results, and this will be followed by a general discussion of the
problem by the senior author.

II. THE CHROMOSOMES DURING THE SPERMATOGENESIS OF

ANASA TRISTIS.

BY EDITH PINNEY.

Preparatory to an attempted settlement of various disputed
questions in the development of the germ cells of Anasa tristis
an independent reexamination of the entire process was under-
taken. In order to avoid unconscious bias of opinion the pre-
vious reports on the spermatogenesis of this species were not
read. Without referring, therefore, to the accounts of other
observers for comparison or correlation it is purposed to record
the results of these observations.

Material. — Preparation and general description.

Material was taken from specimens of Anasa collected in
Lawrence, Kan., and in Woods Hole, Mass. The latter was
obtained through the courtesy of Dr. E. B. Wilson, of Columbia
University. All material was fixed in Flemming's fluid, the
period of fixation varying from one-half to two hours. The
longer fixation was found to give the better results. Paraffin
sections were cut five micra in thickness and stained by
Heidenhain's iron-hsematoxylin, Flemming's tricolor and
Auerbach's methods. Slides prepared by the author from
nineteen individuals were studied. Most of the drawings
were made at a magnification of 3625 diameters. This is re-
duced one-half in reproduction.

pinney: chromosomes of anasa tristis. 353

The testes of Anasa tristis are paired organs in the form of
two small fig-shaped bodies which lie on either side of the
median line in the cephalo-ventral part of the abdominal cav-
ity. They are composed of a number of long, slender follicles
lying parallel to each other and extending the entire
length of the testis. The follicles themselves taper slightly
from the broader base toward the sperm-duct. The follicu-
lar organization into cysts of germ cells of varying stages
of maturation is similar to that in the Orthoptera, and the
same method which was used to determine the sequence of
the observed changes in the Orthopteran species is applicable
here. Practically all of the cells of a cyst are in the same
phase, although exceptions to this regularity are frequent in
the older cysts, particularly those of the first and second sper-
matocyte generations. These variations are, however, to be
expected, and may reasonably be accounted for if one considers
the unavoidable differences which must occur in the rate of
metabolism in cells situated differently in the same cyst.

The longitudinal section of a follicle presents some noticeable
features of a structural nature caused by characteristic dif-
ferences in the cells of various regions. The spermatogonial
cells occupy the distal portion of the follicle, the extent of the
spermatogonial area being determined by the age of the in-
dividual. These cells are small, and in the older cysts the nu-
cleus is accompanied by little cytoplasm. Hence the peculiar
staining quality of the nuclear material causes this region to
appear more intensely stained than the proximal portion of the
follicle. The cells, too, are crowded closely together. In con-
trast to this is the lightly stained spermatocyte region, due to
the fact that here the cells contain a greater amount of cyto-
plasm in proportion to the amount of nuclear material. The
cells have increased so greatly in size in the later generations
that the cysts are larger. Usually a characteristic barrier ex-
ists between the two main regions of divisions. This is formed
by cysts of cells undergoing the process which has been desig-
nated as synizesis, a detailed description of which will come
later. In almost every instance, too, the line of separation is
emphasized by the presence of many cysts of degenerating cells
with their spherical masses of chromatin.

354 KANSAS UNIVERSITY SCIENCE BULLETIN.

OBSERVATIONS ON THE SPERMATOGONIA.

From a study of polar views of the equatorial plate the num-
ber of spermatogonial chromosomes was determined to be
twenty-one, and in no case was there observed a departure
from this characteristic number. The chromosomes in young
cells are larger than those in older cells and do not exhibit the
tendency to coalesce so readily. Consequently cells for study-
ing the number, form and behavior of the chromosomes were
comparatively few, although enough were found to leave no
doubt in the observer's mind as to the correctness of these
observations and their interpretation. Figures 1, 2, 3, 4, 5, 6,
7, 8, 9, 10, show polar views of the equatorial plate from
spermatogonial cells. Certain elements of the complex exhibit
constant and striking characteristics in size relationships. The
largest three chromosomes in each group are easily identified,
as are also the smallest two. The remaining members of the
group more nearly approximate each other in size, so no at-
tempt was made to homologize these in the several cells.

The form of the chromosomes varies from spherical and el-
lipsoidal to kidney-shaped with variations in size and position.
The smallest two elements show the most striking differences
in these respects from the other members of the group. There
seems to be no fixed arrangement of the individual elements,
except that there is an evident pairing of twin chromosomes.
Sutton reported the presence of similar pairs in Brachystola
magna^- and his observations have been verified by other work-
ers in this laboratory on other species of the Orthoptera.f One
element of such a pair is considered to be of maternal origin
and the other of paternal origin. The tendency for equal ele-
ments to get together is very marked even in young cells, and
makes conspicuous the presence of an unpaired element, one of
the largest three chromosomes. Should its mate exist in the
cell it could not possibly be overlooked when even the smallest
elements are so prominent. Owing to uniformity in size, pre-
cisely which of the three is the unpaired element may be de-
cided upon only when two of them are unmistakably paired.
The smallest two elements show this tendency but rarely, their
behavior at this stage being consistent with subsequent irregii-
larities.

* "Spermatogonial divisions of Brnchystola iiKK/na." Kansas University Quarterly,
vol. 9, No. 2. 1900.

t The Chromo.some complex of SyrhuUi admirubilis. W. R. B. Robertson. Kansas Uni-
versity Science Bulletin, vol. IV, No. 13, 1908,

pinney: chromosomes of anasa tristis. 355

All of the chromosomes lie with their long axis in the plane
of the equatorial plate, and from a study of lateral views of
this stage it is evident that the plane of the split in the chromo-
somes coincides with the same plane. Ideal conditions for equal
division of chromatin are therefore present. Division is simul-
taneous throughout the group. The separated halves move
toward opposite poles, maintaining their relative positions un-
til the pole is nearly reached. Thus the shape of the spindle
during anaphase is altered to accommodate the varying condi-
tions. See figures 12, 13 and 14. There is no evidence as to
the method of spindle-fiber attachment, and it is doubtful
whether the matter can be decided in this material. Figure 15
is a drawing of a section through an entire cyst. Most of the
cells are in metaphase. A few are in anaphase. The polarity
exhibited in this section is typical of all cells of this genera-
tion.

In lateral views of anaphases in young cells the chromosomes
are distinct. In the older, more crowded cysts the tendency of
the individual elements to coalesce produces the appearance of
a solid band of chromatin in the equatorial plate. Later stages
of the same show two separate bands with spindle fibers be-
tween. The plate arrangement is lost when the pole is reached.
Figure 17 shows a polar view of an early telophase. The
chromosomes have not yet lost their identity, although they
have apparently begun to disintegrate. Between the telophase
and the succeeding metaphase there is an increase in the amount
of chromatin. The formation of a nuclear membrane and the
further dissolution of the chromosomes follow. A typical telo-
phase from a young cyst is shown in figure 18. The daughter
nuclei are flattened, but assume a spherical outline as the diffu-
sion of chromatin proceeds. Figures 19 and 20 show the cul-
mination of this process.

The inauguration of the next mitotic cycle is marked by the
appearance of an exceedingly long and much coiled spireme,
whether continuous or not it is impossible to say. At any rate,
after the thread has thickened and shortened considerably it is
plainly seen to exist in segments which finally condense to form
the homogeneous bodies of the chromosomes. Figures 21 to
28, inclusive, show the successive steps in this transformation.
The shortening segments show various twists, bends and con-
tortions. See figures 22, 23, 24 and 25. Not until very late in
the prophase does the longitudinal split in the threads become

356 KANSAS UNIVERSITY SCIENCE BULLETIN.

visible. (Figs. 26 and 28.) At this stage pairing of equal
elements is very common throughout the spermatogonial gen-
erations. Note the marked pairs in figures 1, 2 and 4, plate
LXV. It would seem from this that throughout the history of
the spermatogonial cells there is a consistent provision for the
final processes which mark the transition from spermatogonia
to spermatocyte. This process is called synapsis. The cells of
a mature spermatogonial cyst have not been counted, but judg-
ing from the number of cells in a cyst of developing sperma-
tids, eight mitoses complete the activity of the germ cells as
spermatogonia, that is, that after the last division there are
256 cells in a cyst. In such cysts the cells are poor in cyto-
plasm.

After the final division the typical telophase is replaced by
synizesis. During this process the chromatin is contracted
into a deeply staining granular mass whose position within
the nucleus is either central or slightly eccentric, leaving the
peripheral area clear. It is quite certain that the chrome tin
mass is composed of intimately associated granular threads,
for occasionally ends protrude from the knot, although entire
threads can not be traced. (Figs. 30, 33.) It is true that the
chromatin enters synizesis in the form of individual threads,
and the presynizetic stage is characterized by the same ele-
ments. It is reasonable to suppose that the threads continue
as such during the period of synizesis. That some change i&
taking place within the knot is shown later, but there is no
evidence to show that this change involves the disorganization
of the chromatin threads and its reorganization into new ele-
ments. Nothing can be said in regard to the duration of
synizesis. That an end to end conjugation of the paired ele-
ments of the spermatogonial nucleus occurs during this period
seems established beyond a doubt.

At one side of the conjugating threads lies a definite chro-
matin body of homogeneous consistency. Occasionally this
body shows an uneven outline suggesting a slight disintegra-
tion of the constituent chromatin. Its position is constant and
it is invariably present. From a study of subsequent matura-
tion processes it is shown to be the accessory chromosome
(McClung, Orthoptera) and can be identified as the homo-
logue of the unpaired element appearing in the spermatogonial
metaphases. In studying the history of this odd member of
the complex, drawings of it in that stage were made, as well as

pinney: chromosomes of anasa tristis. 357

in numerous subsequent stages, for size comparisons. Fig. 14,
plate LXVI, shows several drawings from different individuals.
The spermatogonial cells from which drawings of the accessory
were made vary in age, and as there is undoubtedly a decrease
in the chromatin content of these nuclei as the cells multiply we
would naturally expect to find homologous chromosomes vary-
ing in size in the same individual. The position of the chromo-
somes with respect to the equatorial plane varies previous to
division, that is to say, in the formation of the equatorial plate
the chromosomes may lie obliquely to the plane of the equator,
and this would cause apparent differences in the size of camera
drawings. Conceding that these possible causes of size varia-
tion are sufficient to account for the slight variations in size
of the accessory as shown in the drawings, we are forced to con-
clude that the size of this element in the spermatogonia is
strikingly constant. A similar study of the homogeneous body
observed in the nucleus during synizesis shows a like con-
stancy in size. Although these two elements which appear at
different times in the nucleus are so unlike in form we cannot
but consider them as equal chromatin masses. Moreover, if
synapsis is a conjugation of twin chromosomes, obviously the
unpaired element of the earlier stages would take no part in
this process. This evidence is sufficient to incline us to believe
that the two masses are identical. Further evidence will be
given to strengthen this conclusion. An account of the presyn-
izetic stages or growth period will be given in the observations
on tlie first spermatocyte generation.

The First Spermatocyte.

The earliest condition of the spermatocyte cell is the result
of the disentanglement of the chromatin knot formed during
synizesis. Even before the chromatin has spread to the nuclear
membrane, and while the peripheral area is comparatively free
from it, it is easy to determine that the chromatin exists in the
form of granular threads or rods. See figures 1, 2, 3 and 4,
plate LXVI. This diffusion of the chromatin threads makes
possible the detection of a second body similar in size and
staining qualities to the chromatin body described as present
during synizesis. This body is not present before synizesis, is
not apparent during the synizetic period, and is present after-
ward in the interior of the nucleus. The natural conclusion
then is that this is a product of synizesis. Its staining qual-

358 KANSAS UNIVERSITY SCIENCE BULLETIN.

ities indicate that it is composed of chromatin. The distinction
between this and the accessory is only possible when the ac-
cessory lies close to the nuclear membrane, which is its usual
position. In this respect it resembles the accessory chromo-
some of the first spermatocyte prophase of the Orthoptera.

The history of the three distinct nuclear components, active
chromatin, accessory chromosome and plasmasome, will be fol-
lowed separately up to the spermatocyte metaphase, beginning
with the chromatin. The reorganization of the nuclear chro-
matin into typical spermatocyte chromosomes after diffusion is
completed gives no hint of a reversal of the mitotic cycle.
That is, the appearances which characterize chromosome re-
construction do not suggest a reversal of the changes which
were observed to take place in the diffusing chromatin. The
straight threads which typify the early prophase (figs. 1-4,
plate LX VI) gradually elongate and finally are disposed into a
fine reticulum at the intersections of which are very slight ac-
cumulations of chromatin. See figure 4, plate LXVI, and
figure 4, plate LXVII. This might be considered as the climax
of diffusion, for in this stage the chromatin is most evenly dis-
tributed throughout the nucleus. Subsequently there are more
extensive accumulations of chromatin at certain points within
the nucleus. These amorphous aggregations of granular
chromatin gradually assume the form of loosely organized
tetrads resembling a few of the types of tetrads found in the
Orthoptera. Figure 16, plate LXVI, shows four types of tet-
rads taken from one cyst all of the cells of which are in
the same stage of development; (a) shows the largest form.
By analysis it is seen to be composed of four equal chromatids
and represents the pair of the largest spermatogonial chromo-
somes conjugated. The disposition of maternal and paternal
chromatids is indeterminable in this as well as in the forms
shown by (b) and (c), which are different types of the cross.
In each case two opposite arms of the cross are composed of
portions of maternal and paternal threads lying parallel to
each other. The ends of these arms are the synaptic ends of
the chromatids, but whether the synaptic end of a chromatid
represents the polar end, as is certainly the case in the
Orthoptera,* it is impossible to determine. One of the other
pair of opposite arms is composed of parallel maternal threads
and the remaining arm consists of parallel paternal threads ;

* Pinney, Edith, OrKanization of the Chromosomes in Phrynotettix magnus, Kansas
University Science Bulletin, vol. IV, No. 14, 1908.

pinney: chromosomes of anasa tristis. 359

(a) shows a rod form in which the division at right angles to
the long axis may or may not separate elements of opposite
parental origin. It is impossible to ascertain whether these rod
forms have passed through the cross stage, since some of the
earliest recognizable tetrads have this form and keep it as they
condense.

The manner in which these forms are evolved from two con-
jugated chromosomes of opposite sex, each split longitudinally,
is similar to that described for the Orthoptera by McClung;"^
(d) is an unusual condition at this stage of the smallest sper-
matogonia! chromosomes. Even in stages as late as those
shown in figures 22 and 24, plate LXVI, the separate halves
of this pair often lie on opposite sides of the nucleus. Just
previous to the metaphase they approach end to end. The
constriction between the two halves is never obliterated by a
close union. Evidently synapsis has had no effect on the
smallest two members of the spermatogonia! complex, al-
though they undoubtedly entered into the synaptic knot. Sub-
sequent changes consist in the condensation of these filiform
tetrads to form the homogeneous elements of the metaphase.
As the chromosomes approximate the homogeneous condition
they arrange themselves at the periphery of the nucleus. This
tendency may account for the characteristic position of the
accessory during synizesis, the growth period and the sper-
matocyte prophases. The same nuclear membrane which is
formed previous to synizesis continues to function up to the
time of a late spermatocyte prophase. At the latter stage
there are present within the nucleus thirteen distinct bodies.
Figures 10 and 11, plate LXVI, show two parts of the same
nucleus appearing in adjacent sections. The thirteen bodies
are drawn. These must represent the twenty-one sperma-
togonia! chromosomes plus the plasmasome created during
synizesis.

Before stating the homologies between these spermatocyte
structures and those of the previous generation the history of
the accessory will be followed, beginning with synapsis. In
this connection the behavior of the new spermatocyte element,
the plasmasome, will be reported. The time of appearance, the
nature and source of these two bodies, have already been spoken
of. It has also been mentioned that immediately after synizesis
the two bodies are practically equal in size. In the succeeding

* McClung, C. E., '00, The Spermatocyte Divisions of the Acrididae, Kansas University
Science Bulletin, IX, 1, '02; The Spermatocyte Divisions of the Locustidae, ibid., XI, 8.

360 KANSAS UNIVERSITY SCIENCE BULLETIN.

diffusion stages the plasmasome increases in size and the dis-
tinction between it and the accessory becomes very marked.
In these stages the accessory remains closely appressed to the
nuclear wall while the plasmasome is located internally. See
figures 3, 5 and 6, plate LXVI. This period during the dif-
fusion of chromatin and the growth of the plasmasome can
not be considered as belonging to the prophase of the first sper-
matocyte. The nucleus increases steadily in size, reaching its
maximum when diff'usion is complete, so we may consistently
call this interval after synizesis the growth period. Its cul-
mination marks the installation of important changes, one of
which, the re-formation of the chromosomes, has already been
described. The accessory meanwhile maintains its former
position, size and staining qualities. During the prophase the
plasmasome gradually diminishes in size, and finally disappears
just before the metaphase. During this time, too, it does not
stain so readily with chromatin stains. In the Heidenhain's
iron-hsematoxylin preparations the staining quality of the
plasmasome varied with the amount of extraction in the iron-
alum solution. Occasionally the plasmasome presents a vacuo-
lated appearance. This may occur at any time during its
history but is more characteristic of the beginning of its dis-
solution. Occasionally in the prophase there occur two small
homogeneous bodies staining like the plasmasome. As the
typical plasmasome is absent and the accessory present these
are considered as fragments of the plasmasome. There is no
regularity in their occurrence. The cysts in this condition are
not frequent and the majority of their cells contain typical
plasmasomes.

In later stages the plasmasome is identified by its shape, be-
coming spherical as it decreases in size. Figure 7, plate LXVI,
shows a nucleus containing the plasmasome and the accessory.
The difl'erence in staining qualities is apparent. The accessory
remains in its former position, its form to the observer depend-
ing on his point of view. Plate LXVII consists of a series of
drawings showing the size relations of the accessory and the
plasmasome at the time of synizesis, the growth period and the
succeeding prophases. The drawing of nuclei at the top of the
page show the condition of the active chromatin. They are
numbered in the order of the changes which they present.
For instance, figure 1 shows a nucleus during synizesis. The
perpendicular column below figure 1, designated la, shows a

PINNEY: CHROMOSOMES OF ANASA TRISTIS. 361

number of drawings of the accessory taken from cells in the
same stage as figure 1. Figure 2 shows a nucleus of the stage
immediately following synizesis. The perpendicular column,
2a, below figure 2 shows a number of drawings of the accessory
taken from nuclei in the same stage as figure 2. The perpen-
dicular column, 2b, beneath figure 2 shows a number of draw-
ings of the plasmasome taken from cells in the same stage as
that shown in figure 2. In the same way columns 3a and 36,
4a and 4b, etc., are drawings of the accessory and plasmasome
from nuclei such as are shown in figures 3, 4, etc., respectively.
The horizontal columns A, B, C and D indicate the different in-
dividuals from which the drawings were made. The differ-
ences in size which are shown in these monoplane drawings
depend upon the point of view of the observer. That the ac-
cessory is constant in size is easily determined by different
focusings. From these drawings there is also found to be a
regularity in the relation between the size of the plasmasome
and the condition of the active chromatin of the nucleus. From
the foregoing observations it is now clear that the thirteen
nuclear structures present in the late spermatocyte prophase
comprise one accessory, one diminished plasmasome, two small
chromosomal elements of opposite parental origin, and nine
ordinary tetrads.

Figures 1 to 10, inclusive, plate LXVIII, show typical polar
views of the first spermatocyte metaphase. All of the groups
shown are from different individuals, with the exception of
figures 2, 3, 4, plate LXVIII, which were in the same section of
the same cyst. As many as six of these views, all typical, have
been observed in one section in one cyst. There are eleven
chromosomes present, each of which, with the exception of the
accessory, is composed of four chromatids. The plasmasome
has disappeared, also the nuclear membrane. The chromo-
somes lie in a clear area surrounded by the cytoplasm. Nine
of the complex, including the largest, form a rather compact
ring, in the center of which appears the smallest member of
the group. The relative positions of the chromosomes forming
the ring vary. One chromosome lies outside of the ring. Fig-
ure 11, plate LXVIII, shows six of these outer chromosomes at
this stage taken from the same individual from which figure 6
was made. So, also, figures 12, 13, 14 and 15 are drawings of
the similarly located chromosome found in the individuals from
which figures 7, 8, 9 and 10, respectively, were made. This

362 KANSAS UNIVERSITY SCIENCE BULLETIN.

eccentric member shows remarkable constancy in size. Its
size and its position suggest the accessory during synapsis.
See figures 29, 30, 33, plate LXV. Figure 5, plate LXVIII,
shows a polar view of an atypical metaphase. This condition
was observed only twice in this material, both times occurring
in different individuals. There appear to be twelve chromo-
somes present. In addition to the small chromosome within
the ring, there are two large equal elements which are not
large enough to correspond to any elements present in the
typical metaphase. Only eight chromosomes form the ring. It
must be that the missing one of the nine which usually form
the ring is represented by the two small and unusual bodies
which lie within the ring, one on either side of the small ele-
ment, and that this is merely an accidental case in which
synapsis did not occur. In connection with this supposition we
recall the fact that synapsis did not occur in the small chromo-
some at the usual time. This similarity of action during the
synaptic period may be significantly related to their similarity
of position.

In a lateral view of a spermatocyte anaphase the same sur-
face of the chromosomal elements is presented to the observer
as is seen in the equatorial plate of the spermatogonial cells.
By referring to figures 1 to 10, plate LXV, it will be seen that
the chromatids have become slightly shorter and correspond-
ingly thicker. Here again the smallest chromosome asserts its
independence. In the spermatogonial cells these small chromo-
somes present spherical outlines. In the first spermatocyte
generation, instead of shortening, as their fellows, they have
elongated. Only in this element and in the accessory was the
plane of division actually determined. Since the two halves
of the smallest chromosome are never so closely united that we
lose sight of their identity, it can be stated positively that
division in this case is transverse and qualitative, causing the
separation of maternal and paternal elements. Owing prob-
ably to the slight attraction exhibited between the halves of
this chromosome division is effected while the other chromo-
somes are still intact. Lateral views of early anaphases in-
variably show this condition. Similar views of later stages
show its separated halves preceding the larger diads to the
poles. In the ordinary tetrads the formation of the crosses
might be interpreted to have the same significance as is at-
tributed to the like form in the Qrthoptera, in which McClung

PINNEY: CHROMOSOMES OF ANASA TRISTIS. 363

has shown conclusively that the first division plane is longi-
tudinal. However, in this case there is no positive evidence to
offer. The chromosomes enter the equatorial plate with their
longitudinal axes parallel to the spindle fibers. Figures 16 and
17, plate LXVIII, show that the polar ends of the chromosomes
converge toward the poles of the spindle. That both methods
of division may be employed by different members of the group
is shown by a study of the accessory at this stage. This ele-
ment, which is composed of two chromatids, was last observed
in the metaphase lying outside of the ring. The element (x)
in figure 16, plate LXVIII, is undoubtedly the accessory chro-
mosome. This drawing is not limited to one plane. The small
chromosome is shown dividing in its usual manner. This ele-
ment, it will be remembered, is located at the center of the
ring and no element of the ring could lie so far from this center
as the one marked (x) . This, then, must lie on the outside of
the circle. From the arrangement found in polar views of the
metaphase one would expect in the lateral views to find sec-
tions which would show four chromosomes in the same plane.
These would include the small central member, two chromo-
somes of the ring, one on either side, and, at one side, the
accessory. Such a section is shown in figure 17, plate LXVIII.
The small chromosome, it will be seen, occupies the axis of
the spindle, which in this view appears asymmetrical, the
asymmetry being due to the extra divergence of the fibers at-
tached to the accessory.

In these lateral views of anaphases the structure of the ac-
cessory is easily determined. It is composed of two chroma-
tids, which are shown in figures 19, 20 and 21, plate LXVIII.
All of the other chromosomes contain four chromatids,
although in some only two of the four appear in the same
plane. The accessory is easily identified by its form in these
stages, where it is shown to divide tardily, division not be-
ginning until the ordinary chromosomes have entirely divided.
In their journey to the poles the accessory chromatids lag be-
hind the others. Figures 17 to 26, plate LXVIII, show this
behavior at different stages. All of the chromosomes of the
complex keep their relative positions until the pole is reached.
Figures 1, 2, 3 and 9, plate LXIX, show polar views of the
dividing groups with the inner and outer members in their
characteristic positions. As soon as the pole is reached the

2-Univ. Sci. Bull., Vol. V, No. 20.

364 KANSAS UNIVERSITY SCIENCE BULLETIN.

chromosomes crowd together and, on account of the lagging of
the accessory, lateral views of this stage show the condition re-
produced in figures 25 and 26, plate LXVIII; (x) is the ac-
cessory. It is easily identified in both daughter groups. Fig-
ure 24 shows the only case observed which might be inter-
preted as the failure of the accessory to divide. The spherical
form of the structure (x) may indicate a degenerate form of
the accessory or an unusually persistent plasmasome. The
crowding together of the diads at the poles completes the cycle
of changes occurring during the first spermatocyte generation.
No telophase involving the disintegration of the chromosomes
and the appearance of a new nuclear membrane occurs.

The Second Spermatocyte.

The second spermatocyte generation begins with the forma-
tion of an equatorial plate by the members of the new sperma-
tocyte nucleus. These second spermatocyte chromosomes con-
sist of ten diads and one monad or single chromatid, the acces-
sory. In figures 1, 2, plate LXIX, of typical equatorial plates
of this generation will be seen a tendency toward the same
chromosome arrangement which prevailed during the preced-
ing mitotic cycle. From the size relations shown in figure 4,
plate LXIX, which is a group of accessory chromosomes from
cells of the same stage and animal as figure 3, plate LXIX, re-
spectively, we are justified in our conclusions as to the identity
of this element. Figures 5, 6 and 7, plate LXIX, are polar
views of beginning anaphases. The chromosomes are viewed
here in cross-section. Figures 10, 11 and 12, plate LXIX, show
lateral views of the beginning of this ultimate division. Figure
10 suggests what transpired between the stage shown in figures
1, 2 and 3 and that shown in figures 5, 6 and 7. The transi-
.tion is extremely brief, for evidence of it is comparatively in-
frequent. The chromatids which, during the first spermatocyte
division, lay parallel to each other have separated at one ex-
tremity and have swung around until they lie end to end, the
separate ends pointing toward opposite poles. The chromatids
maintain this position during a relatively long period of
time, for this condition and the following telophase are most
commonly noted in cysts of second spermatocytes. A typical
spindle is formed and the fibers are attached at the separated
ends of the chromatids. Separation of chromatids occurs
simultaneously, the smallest member here acting in unison

pinney: chromosomes of anasa tristis. 365

with the others. This may be regarded as due to the fact that
the same conditions are present in all of the chromosomes pre-
vious to division ; that is, the degree of union between the chro-
matids is the same in each diad, while it will be remembered
that this relation between the diads which composed the tetrads
of the first spermatocyte varied.

When the chromosomes separate sufficiently to expose a clear
zone of fibers in the region occupied by the equatorial plate
we find the accessory chromatid still in its place, and it keeps
this position until the remaining chromatids have almost
reached the opposite poles. Figures 13 to 18, inclusive, plate
LXIX, show the successive steps in this division. Well coa-
lesced daughter groups of this division are observed at oppo-
site ends of the spindle, while the accessory lingers half way be-
tween. The accessory finally approaches one of the groups and
almost immediately the dividing cell wall is formed. This
leaves the accessory near the newly formed cell wall, entirely
apart from the group to which it evidently belongs. Figure 19,
plate LXIX;, is a group of such bodies drawn from cells in this
stage. The size of this lagging body was thus found to be
constant. The spindle fibers between the two groups persist
until after the nuclear membranes of the daughter cells are
formed. Before this occurs, however, the changes shown in fig-
ures 20 to 25, inclusive, plate LXIX, take place. This consists
in the withdrawal of the accessory from its position near the
dividing wall into the now shapeless chromatin mass of the new
nucleus. In studying these processes it was very easy to obtain
sections which showed the two daughter cells in the same plane.
The connecting band of spindle fibers leaves no doubt as to
their relation even after the new cell wall has been formed.
The accessory occurs in only one of these cells. A further
significant fact is that in every such case where both daughter
cells can be identified beyond doubt one always shows the ac-
cessory at some stage of its characteristic activity, while in the
other no corresponding body is to be found.

As soon as the membrane forms around the spermatid nu-
cleus which contains the accessory this element becomes de-
tached from the mass of which it forms a part and moves again
to the nuclear wall. Its individuality again becomes prominent.
Figures 26 to 31, plate LXIX, show this condition. Here also
the same precautions in determining its identity were observed.

366 KANSAS UNIVERSITY SCIENCE BULLETIN.

Care was taken to study both daughter groups, in one of which
it invariably occurred while the other offered no evidence of
the presence of a similar element. The constancy in size of
the accessory and the relation of its size here to the size of the
accessory as determined by figures 4 and 19, plate LXIX,
help to establish its identity. This condition of the chromatin
in the spermatid nucleus lasts for only a short time. The
large chromatin mass soon begins to break up into smaller
masses which eventually distribute themselves against the
nuclear wall. In an early stage of this disintegration the ac-
cessory is still easily recognized, as it lies on one side of the
nucleus apart from the rest of the chromatin. It still keeps
the elongated form it displayed in the second spermatocyte
anaphase.

A whole cyst of young spermatid nuclei was counted in order
to determine in just what proportion of the spermatids of one
cyst the accessory occurred. The importance of this observation
is obvious, although it seems that the result could be predicted
safely from the evidence obtained from previous observations.
An account of the method used in counting may help to verify
the results. Serial sections of an entire cyst were obtained.
The cyst chosen for counting ran through twenty-four sections.
The sections were five micra in thickness. The exact condition
of the chromatin was found to vary in almost every cyst of this
generation. In the cyst counted were found the stages shown
in figures 28, 29 and 31, plate LXIX. From the conditions ex-
isting in such nuclei we would expect to be able to identify the
accessory from its shape and its position with respect to the
rest of the chromatin in the nucleus. Evidently the number of
nuclei in such a cyst and the number of spermatids formed
would be equal. All whole nuclei or almost whole nuclei and
fragments containing the accessory were counted. It was im-
possible to avoid the error of counting one nucleus as two where
a nucleus happened to be cut exactly in half; one half appearing
in two adjacent sections. The number of nuclei containing the
accessory was 484. The number of fragments of nuclei which
contained the accessory was 40. Thus the entire number of
spermatids in which the accessory occurred was 524. The
number of nuclei counted which contained no accessory was
577. These, however, would include the nuclei from which
small fragments containing the accessory had been severed.

m'clung: chromosomes of anasa tristis. 337

That would make the number of nuclei Math no accessory 537,
which is slightly larger than the number with the accessory. It
has been pointed out that the nuclei Which were cut in two
equal parts might be counted as two. This would add to the
number of cells which did not contain an accessory. Consider-
ing this source of error the number of cells of each kind may
practically be considered as equal. There can be no doubt
as to the occurrence of two kinds of spermatids which develop
into spermatozoa, one containing one more chromatid, the ac-
cessory, than the other; and from the evidence presented we
feel justified in concluding that the two kinds of spermatozoa
occur in equal numbers.

III. A COMPARISON OF THE RESULTS OF STUDIES UPON

CHROMOSOMES OF ANASA.

BY C. E. m'clung!

1. Methods.

The methods of an investigator are a very important factor
in arriving at conclusions, and it is quite possible to secure
diametrically opposite results upon the same material by
varying the technical processes. It will, therefore, be neces-
sary to consider the methods of preparation and study em-
ployed upon Anasa. Paulmier, Montgomery, Wilson, Lefevre,
McGill and Pinney fixed their material with Flemming's fluid,
corrosive-acetic mixtures, Bouin's fluid and various other
cytological reagents. This material was then sectioned in
paraffin, and stained, principally with the iron alum — hsema-
toxylin stain of Heidenhain. The slides thus prepared by
Paulmier, Wilson, Lefevre and Pinney are beautiful examples
of cytological technic and leave little to be desired in delicacy
and precision. A considerable number of the slides sent me
by Professor Wilson and Professor Lefevre are prepared ac-
cording to the methods of Foot and Strobell, so that I have
been able to study the material under their conditions, al-
though, unfortunately, their own preparations have not been
accessible to me. Since the entire case of these investigators,
as opposed to all their fellow workers, rests upon their own
peculiar methods, it seems desirable to consider in detail the
technic upon which they rely.

The usual methods of cytologists are dismissed with prac-
tically no consideration, and instead of fixing the material, it

368 KANSAS UNIVERSITY SCIENCE BULLETIN.

is spread upon glass slips and allowed to dry. (Foot and
Strobell, '07, p. 282.) In this condition it is stained with
Bismark brown and mounted in balsam. Entire reliance is
placed in this method, and little attempt is made to correlate
the results thus obtained with the appearances presented by
other systems of technic. That the position of these authors
on this point may be clearly understood, I quote their state-
ment of the case.

"Professor Wilson, in his recent paper in Science, February,
'07, replying to our preliminary note, says that he thinks the
contradiction in our results is probably due to the difference
of method employed, we having placed our faith in smear
preparations, while he has relied on sections. We are glad
of an opportunity to emphasize this faith, believing that for
demonstration of the structure and count of chromosomes our
modified smear preparations are more reliable than sections ;
and it is for this reason we have abandoned the use of sections
in studying chromosomes, except for comparative work and
for studying the topographical relation of the cells. In cells
fixed and sectioned nearly all the delicate details shown in
the chromosomes of our smear preparations are completely
lost, and it ought to be too obvious to mention that a method
which presents clearly each individual chromosome in its in-
tegrity offers decided advantages when the question of accu-
rate counting assumes the importance and develops the con-
tradictions familiar in recent literature."

The authors state that they have used sections for com-
parison and topographic work, but none of their photographs
are made from sections, and their entire argument, as pub-
lished, is based upon material prepared according to their
own method.

It is my judgment that this method, used alone, is entirely
inadequate for accurate results, and in this particular case is
responsible for the discrepancy between these investigators
and Paulmier, Wilson and Montgomery. From the work of
Miss Pinney it will be clear, I think, that in the prophases
figured by Foot and Strobell there are two darkly staining
bodies, the accessory chromosome and the plasmasome, instead
of the one shown in their photomicrographs. Owing to the
technic employed by them the plasmasome is practically de-
stroyed, and instead of using other methods to determine the

m'clung: chromosomes of anasa tristis. 369

nature of the nuclear body, they assume that it is a plasma-
some. "As the presence of a plasmasome in the nucleus at
these stages is the typical phenomenon familiar in all known
forms (its absence being most exceptional), we feel justified
in interpreting the structure we find in the resting nucleus as
a true plasmasome, and not an odd persisting spermatogonial
chromosome, i. e., chromosome nucleolus." (Foot and Strobell,
'07, p. 284.) Further, the smear method does not give precise
and accurate pictures of the chromosomes. The variation in
size, owing to the different thicknesses of the smear and amount
of spreading, is very extensive, as may be seen by a com-
parison of the photomicrographs of Foot and Strobell. Com-
pare, also, figures 21-26, plate LXX, which represent smears
of cells in the same stage. Not only may the entire groups of
chromosomes in similar cells vary as three or four to one, but
the members of one cell may suffer unlike expansion or con-
traction to a similar extent, if the conditions of drying are
variable, due to the thickness of the smear film. There is but
one advantage of the smear method over sections, and that is
that it presents all the elements of the cell in one plane, so that
their entire outline is visible for studying and photographing.
It is a method that should be used in connection with sections,
and from the beginning of my work I have utilized it con-
sistently and with great profit in this way. I therefore speak
of it as a warm friend and advocate, but at the same time I
realize its limitations and am convinced that it should not be
used to the exclusion of all other methods.

Foot and Strobell have employed photography alone as a
means of presenting illustrations of their material, and it is
assumed by them that if a thing can be photographed it must
necessarily be a true picture of normal conditions. This I
consider to be a decided fallacy. A photograph is an inter-
pretation by the observer, just as is a drawing. The personal
factor is no more absent from one method of illustration than
it is from the other. Photographs may present with greater
fidelity the details of structure in an object, but the choice of
the object and the nature of details are at the command of
the photographer. It is possible, especially in smear prepara-
tions, to select cells that will illustrate almost any condition
and to photograph them. The highly interpretative character
of the photograph is illustrated admirably by the present con-

370 KANSAS UNIVERSITY SCIENCE BULLETIN.

troversy over the chromosomes of Anasa. Foot and Strobell
publish photographs that seem to show that the accessory
chromosome divides in the second spermatocyte division. This
is their interpretation of the behavior of this element, and
they illustrate it by photography. Not a picture which they
publish would indicate that there is ever a case where the ac-
cessory chromosome positively fails to divide at this time,
and yet literally thousands of unquestionable cases of this
occur in every good preparation. See the group of cells
shown in figure 15, plate LXXI. By selecting a series of early
anaphases, and a very few obscure and distorted individual
cells, they are able to secure pictures which appear to bear
out their contention. In speaking thus of the methods of
Misses Foot and Strobell, I wish it clearly to be understood
that I am not reflecting upon their motives. It is my desire
only to show that photographs are interpretations of observed
phenomena and have no claim to infallibility. Again, as in
the case of the smear method, I would state that I speak as a
friend of the method and not as one who deprecates it. I
believe that scientific papers, particularly cytological, should,
whenever possible, be illustrated by photographs, and in my
own publications I have followed this plan so far as the
circumstances would permit.

2. What is the number of spermato gonial chromosomes and

of oogonial chromosomes?

Confessedly, the accurate estimation of the numbers of chro-
mosomes in these generations of cells is a difficult matter, and
if it were the only means of determining the chromosomal
differences of the sexes there might be occasion for doubt with
regard to theories founded upon such enumerations alone.
Fortunately, it is only one of several criteria of sexual differ-
entiation in the germ cells, and so is not of commanding
importance. Accurate counts of the chromosomes, however,
are entirely possible with care, and concordant results are ob-
tainable by independent observers. In the case of Anasa this
is quite possible, and both Miss Pinney and myself have found
the number twenty-one uniformly present in the sperma-
togonia. Not a single instance of twenty-two chromosomes
was found in any spermatogonium, and we must therefore con-
clude with Wilson that thirty-one is the normal spermatogonia!

m'clung: chromosomes of anasa tristis. 371

number. In plate LXX, figures 1-3, may be seen sperma-
togonia! complexes showing this number clearly.

It is claimed by Foot and Strobell that the number is twenty-
two, and they publish four photographs in support of this
view. Only one of these represents a stage where the chromo-
somes are compact and homogeneous, and this shows twenty-
one chromosomes, although in the explanation of the figure
the authors state that one of the apparent chromosomes is
really two joined at right angles. Since there is no case in the
figure where one chromosome is bent at almost right angles,
the explanation is not very convincing. The remaining three
are of earlier stages, in which the chromosomes have a loose
structure that might readily be disturbed in the process of
smearing, thus separating one of the larger chromosomes. In
two of the photographs such an interpretation would seem to be
entirely justified. Photo 49 is the only one in which there
would seem to be any certainty with regard to the number
twenty-two. Here it appears as if conditions were normal,
and it might be recorded as an instance of twenty-two chro-
mosomes in the spermatogonia.

The conditions regarding the spermatogonia! number may be
summarized thus : Paulmier, Montgomery at the time agreeing
with him, reported twenty-two spermatogonia! chromosomes.
Later, Wilson, working upon Paulmier's original preparations
in part, finds twenty-one spermatogonia! chromosomes, and
Montgomery, restudying his own material, agrees with this
enumeration. Lefevre and McGill report twenty-one sperma-
togonia! chromosomes from their own preparations. Foot and
Strobe!!, worl^ing upon smear preparations, consider twenty-
two to be the normal spermatogonia! number. Miss Pinney,
upon examining material from Kansas, Massachusetts and
Pennsylvania, and worl<;ing without a knowledge of others' re-
sults, finds twenty-one to be the spermatogonia! number.
Finally, from a study of the original material of Paulmier,
Wilson, Lefevre and McGill, and Pinney, I myself find in all
clear cases, where there is no doubt of the presence of all the
chromosomes, that the number is twenty-one. Foot and Stro-
bell object to the testimony of Montgomery, and of Lefevre and
McGill, on the ground that they reported at one time an even
number of spermatogonia! chromosomes and at another an odd
number. It must of course be admitted that this weakens their

372 KANSAS UNIVERSITY SCIENCE BULLETIN.

testimony, for it is an evidence either of careless work or of
the influence of preconceptions. I can recall with what diflfi-
culty I persuaded myself, against the current conceptions of
chromosome numbers, that the spermatogonial number of
chromosomes is odd in the Orthoptera, and I can easily under-
stand the temptation, in an early stage of an investigation, to
seek concordance with accepted opinions. The investigators
who reported even numbers for their material did so when all
the evidence from other forms seemed to demonstrate the uni-
versality of this phenomenon. In a sense their testimony is
more valuable after a restudy of their material, because it in-
dicates the desire to arrive at facts regardless of consequences.
It is a very unusual man, or a very stubborn one, who finds no
occasion to change his opinions.

This is the present status of the question concerning the
spermatogonial number of chromosomes in Anasa. If it is im-
possible for the interested cytologist to come to a conclusion re-
garding the facts of the case from the studies so far made, he is
quite at liberty to undertake an investigation for himself and
I have no doubt but that all the material that has been avail-
able for my investigation will be placed at his disposal.

There appears to be no dispute regarding the oogonial num-
ber, although the counts in these cells have been less numerous
than in those of the male. So far as the opportunity has
offered I have gone over the female complex of chromosomes,
and I have no reason to doubt that the number is twenty-two,
as determined by Wilson.

3. What is the number of first spermatocyte chromosomes?

Regarding the number of first spermatocyte chromosomes,
there seems to be no difference of opinion, since every observer
has recorded eleven as typical for this generation. The clear-
ness with which the chromosomes of the first maturation mito-
sis appear in the equatorial plate almost precludes the possi-
bility of error. Here also may be found a close adherence to a
type of arrangement which presents the small chromosome in
the center, surrounded by a more or less regular ring of nine
chromosomes upon the outside of which is placed the accessory
chromosome. This is not an invariable arrangement, for it is
a culmination of the movements of the chromosomes in the pro-
phase where the elements bear somewhat similar relations to
each other, but it appears in a large proportion of the cells.

m'clung: chromosomes of anasa tristis. 373

The position of the accessory chromosome upon the outside of
the group has led Foot and Strobell to coin another new name
for it, and in their terminology it is the "eccentric chromo-
some."

4. What is the behavior of the accessory chromosome and of
the plasmasome in the first spermatocyte prophase f

While there is no dispute regarding the number of chromo-
somes in the first spermatocyte, there is serious divergence of
opinion with reference to the structure of the accessory chro-
mosome. Upon its nature depends the number of spermato-
gonia! chromosomes and the character of the four spermatids
derived from each first spermatocyte, so its determination is
most important. If it be a tetrad then the spermatogonia!
number must be even and the spermatids all alike in the pos-
session of eleven chromatids ; but if, on the contrary, it be a
diad the spermatogonia! number of chromosomes is necessarily
odd and the spermatids of two types, one of which possesses
eleven chromosomes and the other ten.

With the exception of Foot and Strobell a!! observers are
agreed that there are but two chromatids in the accessory
chromosome, but these observers claim that it possesses four
as do the other chromosomes. As proof for this contention,
they publish a series of chromosome groups from the late pro-
phase in which the various elements are clearly shown. The
identification of the accessory chromosome is correctly made in
each case, I believe, but it is entirely clear to my mind that
there is nothing in the appearance of these photographs to
justify the interpretation of the accessory chromosome as a
tetrad. There is in each case a characteristic difference be-
tween the tetrads and the accessory chromosome, for in the
former there always appears a diamond-shaped opening, the
angles of which point to the two lines of cleavage, while in the
case of the accessory chromosome this is lacking. Every ac-
cessory chromosome, on the contrary, shows a clear, straight
line of division along its entire length. In but a single instance
is there an exception, and that is in their figure 4, plate II,
where the plane of longitudinal cleavage is interrupted by
transverse markings, A careful examination of this chromo-
some will demonstrate, however, that there is not only one of
these apparent cross divisions, but two of them, I am con-
vinced that the excellent photomicrographs of Foot and Stro-

374 KANSAS UNIVERSITY SCIENCE BULLETIN.

bell of this stage are sufficiently clear in their evidence of the
nature of the accessory chromosome, and I have not included
any number of these stages in the photomicrographs accom-
panying this paper. Moreover, in none of the material that I
have examined is there any indication whatever of a tetrad
structure in the accessory chromosome, and I am thoroughly
convinced that it is a univalent chromosome.

If the earlier prophase of the first spermatocyte be exam-
ined, the chief error of Foot and Strobell becomes evident. It
is claimed by these investigators that the accessory chromo-
some behaves during the prophase as do the other chromo-
somes and that there is but a single nucleolus-like body present
— a plasmasome. The work of Miss Pinney showing the tran-
sition in form and size of the accessory chromosome and of the
plasmasome, both of which appear simultaneously in the nu-
cleus, makes it entirely clear that Foot and Strobell were mis-
taken on this point. As I have elsewhere pointed out, the
technic of these investigators is responsible for their failure
to find the plasmasome, for it is this structure, and not the
accessry chromosome, that is missing from the smear prepa-
rations.

These investigators have pointed out the large difference
between the appearance of the accessory chromosome in draw-
ings made by Wilson and the photographs illustrating their
own paper. Such variations in the form of this element will
receive explanation in part from the series of drawings made
by Miss Pinney to illustrate the transition in form of the pro-
phase elements. Both the stage of development and the method
of preparation influence the form of this accessory chromo-
some. As a rule, sections show this element as a spheroidal,
apparently homogeneous body during most of the prophase,
but occasionally, just succeeding synizesis, it appears as a short,
thick thread, with a tendency to be bent at the middle. This
is the nearest approach to a spireme condition that it attains,
and in .this corresponds to similar conditions which I have de-
scribed for certain Locustids. So far as my experience goes it
would seem to be the rule that the accessory chromosome, to a
certain degree, undergoes a granular diffusion of the chromatin
similar to that of the other chromosomes, but that it is less ex-
tensive and is frequently marked by the close approximation of
divisions of the thread.

It will be observed from plate LXX, figures 4-6 and 11, that

m'clung: chromosomes of anasa tristis. 375

the accessory chromosome in this spireme condition is accom-
panied by a large distinct plasmasome which lies toward the
center of the nucleus. Such a structure is absent from the
photographs presented by Foot and Strobell when the accessory
chromosome (their plasmasome) is elongated. As will be noted
in Miss Pinney's description, the plasmasome does not appear
until after synizesis is established. Some of the nuclei figured
by Foot and Strobell would seem to be presynizetic in develop-
ment and could not therefore contain plasmasomes.

I would conclude, therefore, that after synizesis there are
present in the first spermatocyte nuclei two nucleolus-like
bodies, one the accessory chromosome and the other a plasma-
some. Further, that the accessory chromosome exists through-
out the whole period, for a time, as a short, heavy thread, at
other times concentrated into a mass, and finally in the late
prophases as a straight longitudinally split rod. The plasma-
some, on the other hand, is an inconstant structure, absent be-
fore synizesis, and again in the late prophase, and is not to be
mistaken for the accessory chromosome.

5. What is the behavior of the accessory chromosome in the

first spermatocyte mitosis'?

Reference has already been made to the condition of the ac-
cessory chromosome in the late prophase and to' its position in
the equatorial plate. It appears in the metaphase as a uni-
valent, longitudinally split rod lying outside the ring of ordi-
nary chromosomes. In division its halves separate and during
the anaphase move to the two poles of the spindle as do those
of the other chromosomes. Examples of this may be seen in
figures 2, 3, 4, 5, 6, 7, 8, 9, 10, plate LXXI. Both series of
daughter chromosomes may be seen in plate LXXI, figure 3,
which shows a mid-anaphase with the halves of the accessory
chomosome accompanying each group. There appears to be no
question on the part of anyone regarding the movements of the
chromosomes in the first spermatocyte, so that an extended dis-
cussion on this point is not called for.

From the fact, however, that Foot and Strobell have used the
tardy division of the accessory chromosome in the first sperma-
tocyte as an argument to prove its division in the second
spermatocyte, because here also it lingers for some time near
the equatorial plate after the other chromosomes have moved
toward the poles of the spindle, it will be necessary to call at-

376 KANSAS UNIVERSITY SCIENCE BULLETIN.

tention to figures 6, 7, 8, 9, 10, plate LXXL These show that
while the time of division is later than that of the other chro-
mosomes the actual separation occurs and the halves of the
divided accessory chromosome may, in each case, be seen in the
daughter cells. No instance could be observed where there
could be doubt of this fact. The clearness of these phenomena
is due to the fact that the accessory chromosome remains, as in
the prophase, distinct from the other chromosomes, so that it
can be followed even into the telophase. As will be seen later,
the same aloofness on the part of the accessory chromosome
is encountered in the second spermatocyte and is so marked
that there is no difficulty in tracing its behavior.

6. What is the behavior of the chromosomes in interkinesis?

The interval between the two spermatocyte divisions is com-
paratively brief and the changes in the relative positions of
the chromosomes slight. While the chromosomes mass together,
the outlines of the individual elements may be seen clearly in
favorable preparations of the telophase. Since the whole
daughter chromosome complex travels to the pole at about the
same rate, its members at the end of the movement lie at
approximately the same level. When therefore the second
spermatocyte spindle is formed there is little or no change in the
positions of the chromosomes, and the equatorial plate of the
metaphase corresponds in general with that of the first sperma-
tocyte, from which it was derived. While this is true, the slight
movements of all the chromosomes during their poleward mi-
gration usually results in a destruction of the typical ring
arrangement so characteristic of the first spermatocyte. Be-
cause of its isolated position and somewhat greater variation
of movement in the anaphases, the accessory shows more
marked diff'erences in position in the equatorial plate than the
other chromosomes. If it has moved toward the pole at the
same rate as the rest of the complex and maintains its position
during the telophase it will then be found upon the periphery
of the equatorial plate in the second spermatocyte. If, on the
contrary, as often happens, the movement is slightly slower
than that of the other chromosomes and it swings around un-
der the mass during the anaphase movements it will later ap-
pear within the group of chromosomes in the resulting equa-
torial plate. It is not possible, therefore, to recognize the ac-

m'clung: chromosomes of anasa tristis. 377

cessory in the second spermatocyte by its position, as might
frequently be done in the first spermatocyte. I am of the
opinion accordingly that any identification based upon position
alone is not warranted in the case of the accessory chromosome
in the second spermatocyte. By this I do not mean that there
may not be similar arrangements of the chromosomes in both
spermatocyte metaphases, but rather that the opportunities for
movements of the chromosomes during the first spermatocyte
anaphase and telophase are so great that no rule may be laid
down to mark a type of second spermatocyte. (Plate LXXI,
figures 11, 12.)

7. What is the behavior of the accessory chromosome in the
second spermatocyte mitosis?

At this point in the history of the accessory chromosome of
Anasa there occurs the wide diversity of opinion between Foot
and Strobell and the other investigators who have studied it.
The issue is clearly drawn. Foot and Strobell claim that the
accessory chromosome is divided in the second spermatocyte
as in the first spermatocyte, but Wilson and others are just
as positive that it passes undivided into but one of the pair
of spermatids formed by the division of each second sperma-
tocyte. Issues are joined here on a question of fact, and it be-
comes necessary to determine, therefore, whether the accessory
chromosome divides in the second spermatocyte metaphase or
whether it passes undivided into one of the two daughter cells.

I must confess that it is a matter of no little astonishment
to me that there should be any difference of opinion on this
point, because the conditions are so clear and unmistakable as
almost to preclude error. A number of photomicrographs of
the second spermatocyte anaphase have been prepared from
both sections and smears and are reproduced in plate LXXI, fig-
ures 13-27. In figure 15 there are shown ten pairs of spermatids
as they appeared spread upon the glass slip. This slide was
prepared according to the method of Foot and Strobell and
shows the cells practically separated so that there can be no
question of a later division of the accessory chromosome. In
each one of these ten pairs it is clearly seen that one member
contains an accessory chromosome while its mate lacks it. The
same statement holds true of all similar cells drawn and photo-
graphed, and of the thousands of others examined. In no case

378 KANSAS UNIVERSITY SCIENCE BULLETIN.

was a divided accessory chromosome found in any second
spermatocyte. For a short time the accessory chromosome
lies against the mass of chromosomes, but when the spermatids
are separated and the nuclei re-form it moves away again and
lies at one side. Such conditions are shown in plate LXXI, fig-
ures 20-21, 25-27. No clearer or more precise demonstration of
a cytological fact could be asked than is here afforded by the
unilateral movement of the accessory chromosome.

The question naturally arises, why, if this be true, should
two observers decide that the accessory chromosome does here
divide? It is of course, impossible to explain just why this has
occurred, but it would seem to me that Foot and Strobell were
impressed with the necessity of showing every chromosome in
both daughter cells and that they chose stages which exhibited
such arrangements. Since it is impossible to find these con-
ditions after a mid-anaphase, owing to the massing together of
the chromosomes, all their pictures were made in the early
anaphase while the accessory chromosome lies in or near the
equatorial plate. Not a single pliotograph of an unmistakably
completely divided second spermatocyte is shown among all
their figures. Since it is only at this time that the accessory
chromosome may definitely be demonstrated to occur in only
one daughter cell their failure to figure it may be understood.

Since Foot and Strobell base their case upon the evidence
afforded by their photographs, it is proper that these should
be considered. They show in plate III twenty-three figures of
second spermatocyte anaphases and telophases. In all but
figure 45 the accessory chromosome lies undivided in the equa-
torial plate, and in this two independent nuclei with no clearly
marked accessory chromosomes are advanced as evidence that
the accessory chromosome has divided. Figure 46 shows al?o
two nuclei with no indication that they are daughter deriva-
tives. Figures 29, 30, 31 and 44 are introduced to show the
actual division of the accessory chromosome. Figure 29 shows
the accessory chromosome of a mid-anaphase with a slight
constriction at the center, figure 30 a similar cell with two
distorted and ill-defined chromosome masses, figure 31 two
other similar daughter groups with an apparently divided lag-
ging chromosome, while figure 44 exhibits a mid-anaphase with
the daughter ( ?) groups widely separated and turned by spread-

m'clung: chromosomes of anasa tristis. 379

ing and a broken accessory chromosome midway between them.
In all of these cells the chromosomes are thin and of indefinite
outline and have apparently suffered much in the process of
preparation. There is not a clear case of a divided accessory
chromosome in the series, although individual cells, doubtless the
best that could be found, are selected and photographed. That
later stages show conclusively the undivided accessory chromo-
some, not alone in selected cases but in every cell in a micro-
scopical field, is demonstrated in plate LXXI, figure 15, of this
paper.

The objection may be raised that such a stage does not show
all the chromosomes; but it is not a valid objection because it
is impossible to photograph all the chromosomes at this stage
on account of their apparent fusion into a mass. Moreover,
the essential facts are admitted by all. It is agreed that all
the chromosomes, with the exception of the accessory chromo-
some, divide so that each group contains ten ordinary chromo-
somes; it is agreed that the lagging chromosome is the acces-
sory chromosome. The question therefore to be answered
concerns the behavior of the lagging chromosome. Are its
derivatives found in each of the separated daughter cells of the
second spermatocyte; or is it, undivided, included in only one
of the two ? The evidence on both sides has been presented in
statement, drawing, and photograph and the judgment of those
interested must, in the absence of personal study, be based
upon the presentations. The crux of the situation lies here,
for if the accessory chromosome divides there is no dimorphism
of the spermatozoa and no visible sexual differentiation of the
paternal germ cells, but if it remains undivided then there is a
dimorphism of the spermatozoa and a consequent visible chro-
mosome differential between two numerically equivalent
classes.

It is pertinent to consider also whether in evaluating the
evidence in this case, any considerable number of genuine
instances of unusual conditions can be held to invalidate the
common conclusions reached by numerous independent stu-
dents of these phenomena. There can be no question of the
occurrence of abnormal conditions in the development of the
germ cells. Entire cysts, or even testes, become degenerate
from imperfect coordination in development ; certain daughter

3-Univ. Sci. Bull.. Vol. V. No. 20.

380 KANSAS UNIVERSITY SCIENCE BULLETIN.

groups of cells fail to separate and thus produce spermatids
with two or four tails and as many simplex groups of chromo-
somes; minor aberrations occur throughout all stages. Can
these be held to prove that there is no constancy in the processes
and results of spermatogenesis? Can deviations from the
normal or average be regarded as an argument against a type?
If so there can be no general laws in biology.

THE

KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 21 — March, 1911,

(Whole Series, Vol. XV, No. 21.)

CONTENTS:

Foundations of Arithmetic Arthur Bowes Frizell.

PUBLISHED BY THE UNIVERSITY,
LAWRENCE, KAN.

March. 1912.

Entered at the post-office in Lawrence as second-class matter.

3-3310

THE KANSAS UNIVERSITY
SCIENCE BULLETIN.

Vol. V, No. 21] MARCH, 1911. [voTxv'™i

THE FOUNDATIONS OF ARITHMETIC.

DISSERTATION FOR THE DEGREE OF DOCTOR OF PHILOSOPHY,

SUBMITTED TO THE FACULTY OF THE GRADUATE

SCHOOL OF THE UNIVERSITY OF KANSAS.

By Arthur Bowes Frizell, of Boston.

This thesis seeks a foundation for arithmetic in the
ideas underlying Cantor's formulation of his system of
ordinal types.

It proceeds by postulating, but follows D. Hilbert
and G. Peano rather than E. V. Huntington. The
search is for postulates possessing heuristic and didac-
tic, not merely subsumptive value.

A motif is found in the notion of an abstract group,
which secures the development step by step of all num-
ber systems so far studied without further postulates
than those needed for the transfinite ordinals.

As axioms are to be avoided, it is necessary to state
carefully the definitions and theorems used even when
they are well known to the mathematicians, but in this
case the proofs are omitted.

1. Definition. A set of symbols a, b, . . . will be
said to form a K-class if we possess a test which enables
us to assert in every case either that a = 6 or that a is
not = b subject only to the restrictions that

(383)

384 KANSAS UNIVERSITY SCIENCE BULLETIN.

a ) the result of comparison be uniquely determined

b ) the statements b = a and a = b shall be interchange-

able

c ) from a = b and b = c must follow a = c

d ) a = a if K contains no other symbol equal to a.

2. Definition. An infinite set of symbols is one
which contains parts that can be put into one corre-
spondence with the whole. A finite set is one that is
not infinite.

3. Definition. An ordered set is one in which we
are always able to say either that a precedes b (a < 6 )
or that a follows b {a>b) subject only to the restric-
tions that

a ) the result of comparison is uniquely determined

b ) a < 6 shall exclude a = b but involve b>a

c ) it a<b and b<c, then a<c

d) if a' — a and b' — b, then a' < b' or a' > b' according

as a < 6 or a > 6.

4. Definition. An ordered set is said to be well
ordered when it has a first element and every subset
beginning with the first element has an immediate suc-
cessor in the given set.

5. Definition. A procedure whereby to every a, b
of a K-class is assigned a definite symbol c= aob will
be called a C-rule or rule of combination provided that
by this assignment equals with equals give equals.

6. Definition. A K-class is said to possess the
fundamental group property with respect to a C-rule
when a o 6 also belongs to K.

7. Definition. Modulus of a K-class with regard to
a C-rule is a symbol uotK such that aou = a = uoa
for every a in K.

8. Proposition I. An ordered set defined by the
requirements : a ) it shall contain a given symbol e ;
b ) it shall possess the group property for every combi-

FRIZELL: FOUNDATIONS OF ARITHMETIC. 385

nation eoa where a denotes a previously defined mem-
ber of the set is also well ordered, infinite and forms a
K-class with respect to the given rule. Proof : By
hypothesis the set is to contain eoe = e', eoe' = e",
eoe" = e'", . . . Since it is to be an ordered set no two
of its members can be equal .".to every element of the
whole set can be assigned some one of the subset e',
e'", e", . . . That is, the set is infinite. And byb)
every element has an immediate successor. Finally we
have a test for equality which satisfies the require-
ments of § 1.

9. Scholium. A set defined as in § 8 contains no
modulus. In what follows it will be referred to as an
e-sei.

10. PosUdates. We postulate an e-set for a lower
rule of combination which shall contain a lower symbol
u and e-sets for both this and a higher rule which shall
both contain a higher symbol w, the definitions of the
rules of combination to be completed in §§ 17, 26,
28, 30.

11. The different sets K[uo], K[wo], K[^6'^] are
to be so ordered that the lower shall precede the higher.
Thus they form together a well ordered set

u, uou = u', uou' = u", uou" = n"', ....

w, wow = wu', wowu' = wu", wowu" = wu'", . . .

wntv = w''\ wnw''' = w''", waW" ^W'", . . .

12. Postulates. We postulate 6-sets for the lower
rule which shall contain respectively each of the sym-
bols w' , w" , iv^" , ... in succession. Thus
corresponding to every a of the set K[i^n] we shall
have an e-set K[ao]:a, aoa ^ au' , aoau' = au" ,
. . . Now taking e. g. a = w'" , by the principle of
§ 11 every member of K[ii'""'o] precedes ww'" = w^''
and therefore falls between iv'"' and w^\ That is,
all these new e-sets are interpolated between successive

386 KANSAS UNIVERSITY SCIENCE BULLETIN.

elements of K[?('n]. Hence the totality of symbols
forms a well ordered set.

13. Proposition 11. A rule of combination is asso-
ciative throughout a given K-class if

ao(6oc)= aoboc
for every a, b, c in K.

14. Definition. A K-class is said to form a semi-
group with regard to a C-rule associative throughout K
if by this rule unequals with equals always give une-
quals or if from the relation a'oh = aob, resp. aob'=
aob we can always infer a ' = a resp. h' = b.

15. Proposition III. No semigroup contains more
than one modulus for its defining C-rule. For if u and
u' not = u were both moduli, we should have aou' =
a = aou contrary to i? 14.

16. Definition. An abelian K-class with reference
to a C-rule is one for which the rule is without excep-
tion commutative.

17. Proposition IV. Sufficient conditions of an
abelian class are the relations

ao(6oc) = ao6oc and boa = aob

for every a, b, c in the class.

18. Proposition V. A C-rule is distributive over
another in a given K-class if the relations

(aob)c = acobc and a(boc) = aboac

are satisfied by every a, b, c in K.

19. Proposition VI. A necessary and suflficient con-
dition of the generating rule of an e-set being associ-
ative for the set is the inductive formula of definition.

eoaoh = eo{aoh) .

Proof. If the relation of ao(hoc) = aoh oc has been
established for a certain a and every /> , c of the set,
then by hypothesis eoao{hoc) = eo\ao(b oc)\ —
eo\aohoc\ = eo{aoh)oc — eoaohoc.

FRIZELL: FOUNDATIONS OF ARITHMETIC. 387

But eohoc = eo{boc) by definition.
Therefore e'oboc = e'o(boc) and so on.
Hence the theorem by strict induction.

20. Corollary. The rule is also commutative. For
if eoa = aoe, then by the associative law eoaoe =
eo{aoe) = eo(eoa) . But eoe' = e'oe by § 19, there-
fore eoe'' = e"oe, and so on. And if /^oa = ao6for
a certain a and every h then ho{eoa) = bo{aoe) =
i>oaoe = aoboe = eo{aob) = eoaob.

Hence the proposition follows by strict induction.

21. Proposition VII . The formula of § 19 is also a
sufficient condition of the e-set constituting an abelian
semigroup for its generating C-rule. Proof. If aob
belongs to the given e-set, then eoaob — eo {aob)
also belongs to it. Thus the first group property is es-
tablished. We have proved the associative law and by
definition no two numbers of the set are equal. There-
fore a' o 6 is not =aob when a' is not = a.

22. Postulates. Using capital letters M, N, . . . .
to denote elements of the lowest class K[tfo], and
italic letters a, /^ . . . for those of the class K[t^n]
where a shall precede />, we postulate for every MB =
BM 2i set MB o Na for every Na, then using letters
a, b, . . . . where a shall precede b to denote the re-
sulting symbols we postulate inductively new sets boa
where a = Na and b denotes successively the higher
''polynomial" or composite symbols. Every b-set is to
be simply ordered and possess the first group property
for every boa.

23. Proposition VIII. Every b-set is well ordered,
infinite and forms a K-class for the rule denoted by o .
For the b-set consists of the combinations b, b o a ,
b o a li', b o a u", boa u'", .... and the reasoning
of Prop. I holds good.

24. Proposition IX. The b-sets belonging to a

388 KANSAS UNIVERSITY SCIENCE BULLETIN.

given Mb form together an infinite, well ordered
K-class as regards the rule o . For the set of b's :

Mb, Mhoiv^^\ ,Mbow^^"''\ ....
forms a well ordered K-class between consecutive mem-
bers of which the b-sets are interpolated.

25. Proposition X. The totality of the b-sets forms
an infinite, well ordered K-class as regards the rule o .
For the set of Mb belonging to a given -6 is a well or-
dered K-class and so is the whole set of 6's, the latter
being simply K[u'n].

26. New combinations of symbols already defined
are defined inductively in accordance with the formulas

b o a = a o b and
a o (b o c) = a o b o c.

This secures the group property for the whole set and
the reasom'figof F/and F// establishes

Proposition XL The set of symbols generated ac-
cording to two rules, a higher and a lower, from a single
arbitrary symbol iv by aid of the postulate of order,
the restricted form of the group property and the above
inducted formulas of definition, is a well ordered, infi-
nite set constituting an abelian semigroup with regard
to the lower rule.

27. Scholium. The higher rule remains unre-
stricted and has not been defined beyond the set
K [ w; n ] , therefore is not yet a C-rule for the K-class
of § 26.

28. Definition. The higher rule shall be defined
inductively throughout K [ 'w; n ] by the formula

w ab = w a ab,

where a, b are any members of K [ t<; n ] for which a b
has already been defined.

29. Proposition XII. The class K [ w u ] forms an
abelian semigroup with regard to its generating rule.
Proof by Propositions VI and VII,

FRIZELL: FOUNDATIONS OF ARITHMETIC. 389

30. Definition. The higher rule shall be defined
inductively for the remaining symbols of our abelian
semigroup on the lower rule according to the formulas

{aob)c = acobc and a{boc) = aboac,
where the combinations ab, ac, be shall have been
already defined.

31. Proposition XIII. The higher rule is distribu-
tive over the lower throughout the class of symbols
defined in § 22. Proof. First let a = b = c = l where
I is any symbol oi K[iva]. Then II is defined by § 28
and l{lol) =lloll = {lol)l by § 30. Now suppose
that the formulas of § 30 have been established for all
symbols of K [ ^ o ] up to and including a certain a and
for every b and c.

Then {loaob)c ^ \ lo{aob) fcby associative law,

= lco(aob)c by hypothesis,
= lco{acobc) by hypothesis,
= lcoacobc by associative law,
= (loa)cobc.
Similarly in every other case when a, h, c are replaced
by^oa, lob, loc respectively. But by definition
^(^oNa) = /^oanNa and so on. Therefore by strict
induction the formulas hold for every a, b, c belonging
to the same K[^o]. Hence by Prop. V the theorem
is true in this case. If h, k, I denote different mem-
bers of K [ -w; D ] the definition gives

{h o k)l = hi okl Bind h(kol) —hkohl.
Then the proof is completed inductively by the same
reasoning as above.

32. Corollary 1. The higher rule is a C-rule and
by it unequals with equals give unequals. For since
our symbols form a well ordered set according to the
lower rule, we have a = b o x for any two unequal ele-
ments a , b and therefore ac=^bcoxc .". not = b c .

33. Corollary 2. The higher rule is associative.

390 KANSAS UNIVERSITY SCIENCE BULLETIN.

For this property has been established for the ele-
ments a, 6, .... as members of K [ ^c n ] . Therefore
by the distributive law it holds generally.

34. Corollary 3. The higher rule is commutative.
For this also has been proved in the class K [ ii? n ] .

35. Corollary U- The set of symbols defined in ij 22
forms an abelian semigroup with reference to the
higher rule.

36. Scholium. Beginning with any a> iv we have
abelian semigroups with regard to both rules, and so
beginning with iv for one or the other rule, but no part
beginning with iv forms a semigroup on both rules.

37. Proposition XIV. Necessary and suflficient con-
ditions that a set M constitute an abelian semigroup
with respect to each of two rules of combination, one
distributive over the other, that M contain an arbitrary
w and that M be as generated an ordered set, are the
rules of combination thus far defined.

38. Corollary. The properties in question may be
expressed by the following postulates:

A. There shall be a higher and a lower rule of com-
bination.

B. Combinations of symbols shall be so ordered that
those made by the lower rule precede combinations of
the same symbols by the higher.

C. The higher rule shall be distributive over the
lower.

D. There shall be a set M containing the symbol lu .

E. M shall form an abelian semigroup for the lower
rule.

F. M shall form an abelian semigroup for the higher
rule.

G. Every successive set generated according to re-
quirements shall be an ordered set.

FRIZELL: FOUNDATIONS OF ARITHMETIC. 391

39. Scholium. M is a well ordered set with no
modulus.

40. We postulated two rules of combination for the
symbol w , but only one for the lower symbol u . If
we should postulate a higher rule for u by the same
requirements as for iv the resulting set of symbols
would not differ as regards any group property from
the ^{;-set ; the new abelian semigroups would be re-
spectively holoedrically isomorphic with the former.
If we postulate no higher rule the set K[iio] still
forms an abelian semigroup holoedrically isomorphic
with K[i(;o].

41. Instead let us set up postulates for the u class
of symbols differing from those of § 38 only in adding
another postulate H. The symbol u shall be a modulus
for the higher rule, with the restriction on A-G that
they shall not conflict with H. Then the class K [^i^]
reduces to the single element u , the abelian semigroup
on the two rules reduces to the set K[^to] and postu-
late B drops out, since every combination by the higher
rule is found among those made according to the lower
rule. This new set of postulates may be replaced by
the equivalent set.

42. a. There shall be a higher rule defined from a
lower by the inductive formulas {uoa)h = uhoah and
a(uoh) = auoah.

b. There shall be a lower rule defined inductively by
the formula uoaob = uo{aoh).

c. There shall be a set M containing the symbol u .

d. The set M shall possess the group property for
every combination uoa.

e. M shall be an ordered set.

f . There shall be a set M ^ built up by postulates c, d
and e on the symbol uu .

g. The sets M' and M shall be identical.

392 KANSAS UNIVERSITY SCIENCE BULLETIN.

43. Proposition XV. The set of symbols defined
by the above seven postulates constitutes an abelian
semigroup for each rule and has a modulus for the
higher rule, which is distributive over the lower.

Pivof. The abelian semigroup on the lower rule is
established by Prop. VII, the distributive law follows
from XIII and the other semigroup by § 35. It re-
mains to prove the existence of a modulus.

Let e = uu. By postulate g, e must occur in the set
K[?io] : II, u' , u" , . . .

Suppose e = u" . Then every member of K [eo] will
be found in K[?/^o] but beyond n' . That is, u and u'
are not in K[eo], contrary to g. Therefore the only
possibility of satisfying g is e=^u. Since, then,
un = u, it follows by the distributive law that u is a
modulus.

44. Postulates a . . . g define the natural num-
bers as ordinal symbols and by XV contain all the laws
of their arithmetic. The cardinal numbers may be de-
fined in a manner now familiar as names of classes,
e. g., ''five" is the name given to the class of all well
ordered sets which are ordinally similar to the set u ,
u', u" , u'" , u^\

45. Definition. A group is a semigroup which con-
tains, corresponding to every a, l>, in it, symbols p
and q such that aop = h = qoa.

46. Proposition XVI. Every group contains a
modulus with respect to its defining C-rule.

47. Definition. If a class which has a mod-
ulus u for its defining C-rule contains symbols a , a
such that aoa = u = aoa then a , a are said to be
each the inverse of the other.

48. Proposition XVII. No member of a semigroup
can have more than one inverse in the semigroup. For
otherwise equals with unequals would give equals.

FRIZELL: FOUNDATIONS OF ARITHMETIC. 393

49. Proposition XVIII. Every group contains the
inverse of every one of its members.

50. Proposition XIX. A semigroup with modulus
is a group if it also contains the inverse of every one
of its members.

Proof, a o (a o h) — a o a o h = u o h = J)^ and
(!) oa) o a = h o (a o a) = h o u = h , that is, § 45 is sat-
isfied by p = a o /' and q = hoa.

51. Proposition XX. Given an abelian semigroup
G with reference to a rule denoted by o , if in the class
[ ( m, n ) ] = C of pairs of elements of G we declare
[m,q) = (n,p) when and only when mop = noq and
set up a rule defined by the relation {m,q)Q{n,r) =
{mon,qor), then 1 ) C will be a K-class, 2 ) o a C-rule,
3 ) C an abelian group as regards the rule denoted by
the sign o.

Proof. 1) Either mop = nog' or mop is not =noq
by hypothesis and §1. Hence either im,q) = (n,p)
or(m, g) is not =(n,p). Obviously the assertions
{n,p) = {m,q) and {m,q) = {n,p) are identical in
meaning, since this is so for Tto ^ and mop. It re-
mains to verify the euclidean postulate.

Suppose that (m,q) = (l,r) and (^, r) = (?i,p).

Then mor = lo q and lop = nor.

Hence (mor) o (lop) = (nor) o (loq).
But since G is an abelian semigroup (mor)o(^op) =
mor o lop = (mop) o (lor) and likewise {nor)o
(loq) = (noq)o(lor). Therefore (mop)o(^or) =
{noq)o{lor). Whence mop = noq by definition
of semigroup and finally i'm,q) = (n,p) by definition
of equality.

2) Let (m', q') = (m, q) and (?t', r') = (n, r). There-
fore m' oq — moq' and n' or = no r'. Whence as
under 1) {m' on') o {qor) = {mon) o {q' or') ^Xidi
hence (m' on', q' or' = {mon,qor). That is {m',q')Q

394 KANSAS UNIVERSITY SCIENCE BULLETIN.

(n'yr') = (m,q)Q{n,r), or equals by equals give equals
and © denotes a C-rule.

3) The fundamental group property is secured by
the definition. To prove the associative law we have
(m,n)Q \ {p,q)o(r,s) \ = (mopor,noqos) = (m,n)&
(p, g)0(r, s). Suppose that {m', q')o{n, r) = (m, g)©
( 7^, r ) . Then (m' on,q' or') = (mon,qor) by defi-
nition. Hence {m' on) o {qor) = {mon) o {q' or)
and {m' oq)o{nor) = {moq')o{nor). Therefore
m' oq = moq' hy %\4:. That is (m', g') = (m, g') and
similarly for the other form of this property. Thus C
is a semigroup. But the element ( m, m ) is a modulus
and to every {n,q) corresponds an inverse {q,n).
Therefore by XIX, C is a group, which we will denote
byG.

Finally {n,r)Q(m,q) = (nom,roq)
= {mon,qor) — {m, q) o {n, r) .

Therefore by § 16, G is an abelian group.

52. The semigroup C and group G are connected by
Proposition XXL If we declare (m o q, q) = m then

(mo 7, <j)o(n or, r) = mo n. For (m oq, q)o{nor,r)
= {moq onor, qor) = monoqor,qor)= mon.

53. Scholium. The group G contains a semigroup
K holoedrically isomorphic with C in such manner that
the rule 0 becomes identical with the rule o for K.

54. Obviously Prop. XX may be applied to the set
of natural numbers in two different ways according as
we use the semigroup on addition or that on multipli-
cation. An essential distinction between these two
procedures is furnished by

55. Proposition XXII. Given two rules of combi-
nation of which one is distributive over the other, no
set of symbols can form a group with respect to both
rules. For such a set would contain a modulus v for
the lower rule (XVI). Then by the distributive law

FRIZELL: FOUNDATIONS OF ARITHMETIC. 395

a = bx =^ b {x o v) = bx o bv = aobv for every a. That
is hv is also a modulus. But there can not be more than
one modulus (///). Therefore hv = v for every h so
that when h' is not = h we have h'v = v = hv contrary
to definition.

56. Corollary. It is riot possible to define a set of
symbols constituting a semigroup with modulus as re-
gards both rules of § 55.

57. If Prop. XX is applied to the set of natural
numbers and the rule of addition, we lose the semi-
group on multiplication and can not recover it. This
semigroup, in fact, is destroyed if we only annex a
modulus for addition. Consequently we give the pref-
erence to the semigroup on multiplication and proceed
with the group G obtained by applying XX to it.

58. Proposition XXIII. Given a higher rule dis-
tributive over a lower, and a set of symbols forming an
abelian semigroup with respect to each rule, if we build
the group G of § 52 with reference to the higher rule,
then a necessary and suflftcient condition of having a
lower rule over which the higher shall be distributive
and with regard to which G shall constitute an abelian
semigroup is the formula of definition (m, q) o{n, r)
= {rrir onq, qr).

Proof. The necessity of this relation is obvious. To
show that it is sufficient we first let {m' , q') = (m, q)
SLYid {n^, r') = (n, r) .'. (m'q = mq') and n'r = nr'.
Therefore m'r'qr = m'qrr' = mrq'r' and n'q'qr =
nqq'r'. Hence (m'r' on' q)qr ^ (mr onq) q'r' by
distributive law and .'. by definition {m' r' on' q' q'r')
= {mr o nq, qr) or equals with equals give equals and
we have a C-rule. Similarly equals with unequals give
unequals. For the associative law {k, l)o\{m, q)o
{n, r) \ = {k, I) o {mr o nq, qr) = {kqr o Imr o Inq, l(/r)
= (kqo Im, lq)o{n, r) = {k, l)o(m, q) o{n, r).

396 KANSAS UNIVERSITY SCIENCE BULLETIN.

Obviously (??, r)o(m, q) = (m, q)o (n, r) . Thus
we have an abelian semigroup and the distributive
principle follows by taking equals with equals.

59. Definition. The symbols of G shall be ordered
according to the convention that ( m , q ) shall precede
or follow {n, q) according as m precedes or follows n
and by ^ 3.

60. Scholium. If a, h denote two natural numbers
that have no common factor other than unity, the set
of symbols (a, />) is simply ordered by the above defi-
nition. This set, which we will denote by R, may be
taken as representative of G since every element of G
is equal to some member of R.

61. Definition. The symbols constituting the set R
are called absolute, rational numbers.

62. Proposition XXIV. Every two unequal ele-
ments of G are connected by a relation h = g ox where
h, g denote the given elements, g being that which
precedes.

For g = (m, q) = (mr, qr) and h = (n, r) = (nq, qr)
and mr precedes nq hy hypothesis. Therefore h = gox
is satisfied by a? == (d, qr) where nq = mr od.

63. Corollary. The set R is simply ordered accord-
ing to the lower rule denoted by o .

64. Proposition XXV. If we exclude the modulus
u the remaining symbols of R form a set which is sim-
ply ordered according to the higher rule. For let x
denote any element of R which precedes and y any one
which follows u .

Then gx precedes and gy follows g'. That is, the
trios u,g,gx and u,g,gy are ordered whether g
precedes or follows u .

65. Corollary. The modulus u separates the re-
maining symbols of R into two simply ordered classes,
neither of which contains either a first or last element.

frizell: foundations of arithmetic. 397

66. It is possible to divide R into two ordered
classes so that every element of R belongs either to the
one or to the other, but neither class has either a first
or a last member. For example, the class Ri of all ele-
ments whose squares precede 2 and the class Rg of all
whose squares follow 2 together exhaust the set R,
while neither Ri nor R2 has either a first or a last ele-
ment. If we take 4 instead of 2, the element 2 is not
included either in R or in Rg ; it separates them. We
usually make this separation in practice by selecting a
well ordered set, e. g. according to the decimal scale.
We take first the highest integer in R^, then the high-
est number of tenths, hundredths and so on. Similarly
we pick out first the lowest integer in R2 , then the
lowest set of tenths, hundredths, etc.

67. Definition. A well ordered set which has no
last element is called a series.

68. Definition. The series of all symbols of the
well ordered set 1 , 2 , . . . w, w -^1, . . .
2w, . . . Sw, . . . w^, . . . which pre-
cede the element a taken in the above order, is said to
be a series of type a .

69. Definition. An ordered set which has no first
nor last element will be called an unbounded set.

70. Given an unbounded set S ordered according to
a rule with respect to which S constitutes an abelian
semigroup, suppose that a series K [f ^J of type w has
been selected from among the elements of S and let
Wi denote the set of all symbols of S which follow, Vg
all which precede every fj, let Vi denote the set of all
symbols s which precede Wj and Wg the set of all that
follow V2 .

Then one of the sets W^ , V^ must be unbounded and
both may be. So also of V2 and W2 .

3-Univ. Sci. Bull. Vol. V. No. 21.

398 KANSAS UNIVERSITY SCIENCE BULLETIN.

71. In every one of the four cases when

Wi has a first element or V^ has a last
V2 has a last element or W2 a first

this element is called the limit of the set to which it

belongs.

72. In case W^ and V^ or V2 and W2 are both un-
bounded we will introduce a new symbol f ^ which we
will call the limit of K [f ,J .

73. Lemma. If Wi and Vi are both unbounded for
the series i^[f„] the same is true for the series
K[aof„] where a is any symbol in S. For if either
the W or V set corresponding to i^ [a o f „] had a limit
I we could write 1 = aox and x would be a limit for the
same W or V belonging to K [f ,J contrary to hy-
pothesis.

74. Theorem. If the series K[fi\, K[gj] both
divide S into two unbounded sets the same is true of
the set K [f ^ o gj .

Proof. Let hi = f 1 o gi , hg = f 2 o gg , . . . .

h^ + i = f2 0gi, h^ + s^faOga, ...

Then by the Lemma each of the series K[hj],
K[h^+,], K[h2^+,], . . . K[h,,,+J, . . . divides S
into two unbounded sets. Hence the same is true of
tne set n^ , n^_).2) n2,^+3, . . . n^^.^^^.!.

75. On the basis of the preceding theorem and defi-
nitions is built up, by laying down the usual definitions
of equality, order and two rules of combination, the
set X of limits of series K [ f „ ] selected out of R. The
set X forms an abelian group with reference to the
higher rule of combination and an abelian semigroup
on the lower rule. The higher rule is distributive over
the lower and possesses the modulus u. X may be
simply ordered according to the lower rule and is di-
vided by the modulus u into two unbounded sets each

FRIZELL: FOUNDATIONS OF ARITHMETIC. 399

simply ordered according to the higher rule. X is itself
an unbounded set and contains a sub set holoedrically
isomorphic with R as regards each rule of combination.
Thus the whole set of absolute numbers is deduced
from the set of absolute rational numbers by using only
principles of order.

76. The preceding development does not take ac-
count of the possibility that K[f„] may not really di-
vide S. There may be no symbol which follows every
f,-. Then Wi is an empty class and V^ is taken to coin-
cide with S. Thus if K[f„l and K | g„ | are both
series of this kind, they both have the same W and V,
whence if new symbols t,„ , g,, were to be introduced
the principle of definition used in s^ 75 would lead us to.
declare f,^ = g,^. Therefore we assign to the totality of
the sets K | f „ | for which Wi is an empty class a single
symbol Z which is to follow every element of X. Then
following ^75, Z o g^ is to be defined as h^ where h„ =
f n o gn and f ,„ = Z . But for K [ h^ ] , Wi is empty,
therefore Zox — Zog^ = h,„ = Z . That is, Z-\-x =
Z = x-\-Z, 7jX = Z^xZ, and similarly Z + Z = Z =
ZZ = Z^ = Z^. The associative, commutative and dis-
tributive laws still hold as well as the first group prop-
erty, but the semigroup is destroyed for both rules by
violating the law of equals with unequals.

77. It is also possible that no symbol in S precedes
every f^. Then Vg is an empty class and Wg coincides
with S. On the same principle as in § 76 we assign to
the totality of series of this kind a single symbol v to
precede every x and define v o ic = h^ where h„ = f„ o g„,
g^ = X, f^ = v. And now the two rules must be dis-
tinguished. Since no x precedes every f, the sets
K|f„ + g„] and K[g„] separate S into the same V
and W. Therefore v + a; = h^ = g,„ = a; + v. And for
the same reason no s can precede every f„ g„

.". vo; = (fg)^ = V = xv

400 KANSAS UNIVERSITY SCIENCE BULLETIN.

Also in like manner v + v = v = vv .

In the set K|h, v| the associative, commutative
and distributive lav^s and the first group property are
all preserved. The law of equals with unequals is
violated for multiplication but still holds for addition,
for which v is a modulus.

78. Since the introduction of infinity would destroy
both semigroups while zero destroys only that on mul-
tiplication, it seems preferable to admit zero to our
arithmetic and exclude infinity. But this removes the
only objection to enlarging the addition semigroup into
a group. This is effected at a stroke by applying Prop.
XX to the set X and thus building an abelian group
with reference to addition. In this group we then de-
fine a higher rule by the formulas

aob — ah = a&T> and aeT) = al>.

This completes the system of real numbers, forming
an abelian group on addition and, when we exclude its
modulus, an abelian group on multiplication, which is
distributive over addition. The system of real numbers
can be simply ordered according to the lower rule, but
the abelian group on the higher rule can not be simi-
larly treated.

79. We might have proceeded by first applying XX
to the natural numbers as a semigroup on addition.
This yields the whole set of integers, positive, negative
and zero. Omitting zero and defining by the law of
signs we should have an abelian semigroup on multi-
plication. Applying XX to it, there results an abelian
group composed of all rational numbers except zero.
Then the introduction of the limits would supply the
whole set of irrational numbers and close by reintro-
ducing zero.

80. As long as we postulate two rules of combina-
tion, one distributive over the other, and demand semi-

FRIZELL: FOUNDATIONS OF ARITHMETIC. 401

groups with reference to both, we are led inevitably to
the transfinite types in case we impose no restriction
on the multiplication table and, if we require a mod-
ulus, to the natural numbers, whence the system of
real numbers results from the attempt to build groups.
The two rules of combination connected by the distrib-
utive property may be regarded as defining arithmetic
up to date ; it has not yet been found profitable to pos-
tulate in any other way. Then transfinite arithmetic
is distinguished by postulating the semigroups and
finite arithmetic by the postulate of the modulus.

81. The real numbers form the most general finite
system with a single unit and must enter into every
system with more than one principal unit, one unit
must always be the modulus. Systems with two or
more principal units have been studied exhaustively by
Weierstrass. Here the abelian group on the lower rule
is postulated universally.

With two principal units, if we exclude the modulus
of the lower rule, the necessary and sufficient condition
of an abelian semigroup on the higher rule is the exist-
ence in the system of a symbol i such that ii = u.
Thus the common complex numbers form the most
general system with two principal units satisfying the
postulates for real members.

82. Within the system of common algebra are dis-
tinguished different number bodies each built on a root
of a given algebraic equation as a unit. The algebraic
numbers of a given body form an abelian group on ad-
dition and, excluding zero, an abelian group on multi-
plication, and must contain a given symbol, the root of
the given algebraic equation. For example the system
of common complex numbers is the number body which
contains a root of the quadratic x^ -|- 1 = o ,

402 KANSAS UNIVERSITY SCIENCE BULLETIN.

83. An integral algebraic number satisfies an equa-
tion

where every a^ is an integer. A good illustration of
the serviceableness of the group theory process for
arithmetic is the theorem : The whole numbers of a
quadratic number body form an abelian group on addi-
tion and, if we exclude zero, an abelian semigroup with
regard to multiplication.

84. A more striking illustration is furnished by
Dedekinds' ''Ideals." An ideal is defined as a combi-
nation of the whole numbers of a number body posses-
sing the first group property for both addition and
multiplication. The ideals of a given body form an
abelian semigroup with respect to multiplication.
There is no addition of ideals.

85. Similar applications are found in the expression
of an ideal by aid of its basis and in the cognate formu-
lation of the transformations of a collineation in space
of N dimensions. The latter question resolves itself
into that of complex numbers with N principal units.
Here it is no longer possible to preserve even the re-
stricted abelian semigroup on the higher rule.

Not only is the abelian character lost, as in quater-
nions, but the semigroup property may be violated on
account of the possibility of a combination by the
higher rule being zero when none of the factors is zero.

86. It is now possible to describe more concisely the
relation of the transfinite arithmetic to common arith-
metic. The symbols defined in ^ 22 form a set of num-
bers with infinitely many principal units w, w^, w^, . .
i. e., the principal units form a series of type w. In
finite arithmetic, it is true, the symbols ^t^ u^, ... all
belong to the set generated from u as principal unit,
but if we allow this analogy in the transfinite system
there will be still more new symbols.

FRIZELL: FOUNDATIONS OF ARITHMETIC. 403

For just as the removal of restrictions on the higher
rule carried our series of symbols beyond the finite set
of type w so the introduction of a still higher rule, cor-
responding to involution, will extend it beyond the set
defined in § 22 if we impose no restrictions on the new
rule except that of order and the special form of the
group property.

87. Let us postulate a rule expressed by a^ higher
than the two preceding, that is, the combination a b
shall precede a*'. To distinguish we will now call ab
the lower and a o b the lowest rule.

The lower rule has been shown to be associative and
commutative for the set K [ -m; n J , which forms an
abelian semigroup with regard to it. A set of symbols
shall be generated from w by the higher rule in accord-
ance with the postulate of order and the restricted
form of the group property for iv^, where a is a pre-
viously defined element. By Prop. I this set forms a
series w,^^ = w', iV"' = w" , w'^"w"', . . . and now w'
must follow every w^ (N == 2, 3, . . . ) [^(;^ is not a
combination by rule, the N is a mere index ] . Then
by § 22 we obtain a series holoedrically isomorphic with
the set there considered if to the lowest and lower rules
respectively we make the lower and higher correspond.

88. The articulation of the two lower rules in ^^ ^^ 28-35
leaves room for much freedom in definition. There we
defined the lowest rule first and made the lower de-
pend upon it. Here we have already defined the lower
rule to the extent implied in the application of § 22.
This, however, involves no further restriction than the
inductive associative formula of § 19. But the defini-
tion from the lowest rule carries this property with it.

It is therefore permissible to define the lowest rule
so that the lower shall be distributive over it. Then
by the process of § 22 new combinations are made by
the higher and lowest rule together, from each series

404 KANSAS UNIVERSITY SCIENCE BULLETIN.

of type w in the present set of symbols a new set of
the same type as that in ):^ 22. And all new series
being fitted in by the nature of the process between
consecutive elements of the series previously defined
we have at every stage a totality of symbols forming a
series.

89. The group properties of the two lowest rules
are preserved throughout the preceding process. On
every new symbol not obtained from the preceding
by postulating them is built up a set forming abelian
semigroups on both the lower and lowest rules, the new
symbol playing the part of a unit precisely like w in
the series of ^ 22, and in this set the lower rule is dis-
tributive over the lowest. This process is to be con-
tinued as long as new symbols can be obtained by it.

90. There remains one more possibility of building
new symbols by introducing a combination defined one
way of the higher symbols with those of the series
K [2^ o] according to the lowest rule. Thus to every a
in the higher series will be assigned a ^-series a , slou,
Siou', 2ion" , Siou'", . . .

This series fits in between a and its next following
element in the former set so that we still have a to-
tality forming a series.

91. For completeness it is convenient also to define
combinations one way of certain symbols with those of
K['Zio] according to the other two rules, viz.: those
expressions in which the symbols of Kfiio] have
already been used as marks or indices, e. g. :

u'w, u"w, u"'tv, . . . w' , W"^" , W'" , . . .
u'w' , u"w' , u"'w' , . . . w'""' w'""" , w'""'" , . . .

Expressions uoa, u'oa, . . . u''" , u"'", . . .
u'""', u"'^' , . . . are not defined, neither are there
any combinations according to the higher rule except
the set w where a belongs to this set. The higher

frizell: foundations of arithmetic. 405

rule has not been made a rule of combination for any
whole set. This would involve assigning properties to
it and completing the arithmetic of the symbols we
have defined, which is beyond the present purpose. It
is enough to have them in series ; that forms the foun-
dation of their arithmetic-

92. The mere existence of the series of symbols de-
veloped in the preceding H is enough to solve a great
variety of problems arising in analysis, the nature of
which will be illustrated by comparing this series with
series consisting of absolute numbers. The natural
numbers form a series of type iv if they are arranged
in the order of their genesis. It is, however, possible
to arrange them in series of higher types for e. g. the
odd numbers alone form a series of type w. If to this
we add on the even numbers successively we obtain
series of types

'w-{-l, w-\-2, . . . u'H-N, . . .
and thus the whole set is ordered in type 2w .
If we order the set R of ^ 60 as follows :

(1,1), (2,1), (3,\). (4,1), . . .

(1,2), (3,2), (5,2), (7,2), . . .

(1,3), (2,3), (5,3), (7,3), . . .
we have a series which can be put ordinally into
one to one correspondence with the series 1,2, . . .
w-\-l, iv-\-2, . . . 2w , 2w-\-l, . . . i.e.,
is of type ^^^

The same set of numbers (the common fractions)
can, however, be arranged in a series of higher type.
For every finite, simple, continued fraction is equal to
some common fraction and conversely. Now let us
order the finite continued fractions according to the
values of their successive quotients

Thus the continued fractions containing each a single
quotient form a series of type w . Then from each

406 KANSAS UNIVERSITY SCIENCE BULLETIN.

member of this series by adding a second quotient
results again type lu . Therefore the fractions of the

form — ^ -~ may be arranged in series ordinally sim-
ilar to 1, 2, . . . w-]-l, . . , 2w, . . . 3iv, . . .
i. e. of type w^ . Annexing a third quotient replaces
each element of this set by a series of type w so that
we include types iv'^-\-l, w^-\-2, . . . 2iv^y . . .

Sw^, . . . i. e. the class of fractions -^ -^ 4-

is ordered in type iv^.

Therefore the whole set of simple continued frac-
tions, comprehending types 'w , . . . w^, . . . tv^, . . .
w^, . . . constitutes, as ordered, a series of type

w"' = w'.

93. Now it is easy to arrange the natural numbers
also in a series of type iv^ as follows. We know that
the class of prime numbers can be put into one to one
correspondence with the whole set of natural numbers
(the number of primes is infinite).

Therefore we can set up a one to one correspondence
ordinally between the prime numbers and the simple
continued fractions with a single quotient. By the same
reasoning the continued fractions with two quotients
are shown to be ordinally similar to the class of prod-
uct of two primes, i. e., if to every quotient q^ we
assign that prime pi for which qi is the ordinal number
in sequence (so that e. g. to the quotient 7 we assign
17) and likewise for a second prime pa and quotient qz,
and so on.

Proceeding in this way the class of all products of N
primes is exhibited as ordinally similar to the class of

continued fractions -^ -^ ' ' ' —- But the class of

all products of primes is the whole set of natural num-
bers. Therefore by § 92 the natural numbers may be
arranged in series w' . Q. E. D.

94. In other words we have here a method whereby

frizell: foundations of arithmetic. 407

the elements of any ii;-series may be rearranged so as
to produce a series of type w "' . Applying this method
successively to the ^(;-series in the preceding we obtain
in place of ?<; , 2w, . . . w', 2w' . . . that is, instead
of 2(;^ wegeta type?^i(;'. Thenw^^iv, iv^-\-2tv, . . .
are replaced by !(; it'' + ^(;', ivw' -{-2w' , . . . making
in all type 2ivw' replacing the former 2w^.

Then come in succession types 3ww' , iww' , . . .
so that the original iv^ expands into a type w^w' . It
is clear that in this way we retrace precisely the steps
of the process of § 22, building up on each new symbol
the abelian semigroups according to the lower and low-
est rules.

Therefore by successive applications of the method
it will be possible to rearrange the natural numbers in
series of types as high as any of the set hitherto de-
fined. This result may also be stated. The transfinite
ordinal series so far defined may each be put into one
to one correspondence with the set of natural numbers.

95. The symbols w, tv-\-l, tv-\-2, . . . i. e., the
transfinite symbols, are said to form the second ordinal
class, the finite symbols constituting the first class.
The latter class was said to be of type w, where w is
the symbol following next after all the finite symbols
of K[^to]. Likewise we shall say that the first and
second ordinal classes together form a series of type
W , introducing this new symbol without assigning to
it any properties except that it shall follow next after
all symbols of the second ordinal class, i. e., after all
the setK['M;''] : w, w', w" , . . .

96. We see that the ordinal symbols play a double
role. They were defined as symbols forming a well
ordered set. But to each symbol which has no imme-
diate predecessor corresponds a series of which it is the
type, viz.: the series of all its predecessors or any
ordinally similar series. The arrangements of the nat-

408 KANSAS UNIVERSITY SCIENCE BULLETIN.

ural numbers in § 94 are only a part of these series.
But they are also only a part of the permutations of
the whole set of natural numbers. Other permutations
are obtained by the following Lemma. From any tv
series of symbols may be obtained a set of permuta-
tions of the symbols forming a series of type tv' = iv"\

Proof. Letai, ag, . . . aN, • . . denote the given
symbols in the given order. Without changing the
order of the higher a's put ai successively in every sub-
sequent place. This set of permutations is obviously of
type w .

From every one of them by repeating the process on
a 2 we obtain again a w;-series .'. in all a series of per-
mutations of type td Repeating the process with as,
a4, . . . . successively we have a series of permuta-
tions whose type is w'. Q. E. D.

97. Thus the arrangements of the natural numbers
furnish a series of type not lower than W. For by § 96
we first deduce from the series 1, 2, 3, ... . i. e. , the
normal order, a set of permutations forming a series of
type w'. That is, to every ordinal symbol preceding iv'
is assigned a permutation, and vice versa. Then by >i
94 arrange the natural numbers in series iv' which ob-
viously is a different permutation from any of the pre-
ceding •.' it can not be obtained by the process of ^ 96.
Now repeating the method of ^:J 96 on each ?('-series of
this permutation we use up all ordinal symbols between
w' and that which results from it by a second applica-
tion of § 94. This holds step by step as long as the
latter process can be carried on. Thus to every ordinal
symbol in succession preceding W is assigned a new
permutation. That is, we have a set of permutations
of all the natural numbers forming a series whose ordi-
nal type is W.

98. The process just described for making permu-
tations of all the natural numbers can not yield a series

frizell: foundations of arithmetic. 409

of type higher than W since, as we have seen, it gen-
erates precisely the series W of ordinal symbols. That
there are types higher than W is obvious, for we can
proceed with W just as with tv to generate new e-sets
and new abelian semigroups, and there is no limit to
the possibilities in the way of still higher symbols and
rules. Now it is quite conceivable that there may be
further permutations of natural numbers not obtainable
by the above process. As a first step toward investi-
gating this question let us consider the simpler one
whether the natural numbers themselves can be ar-
ranged in a series of ordinal type higher than W. For
this purpose we will establish the following

Lemma. Permutations of the natural numbers can
be made by the process described and ordered so as to
form a series of type higher than any series of all the
natural numbers, however arranged. For by the proc-
ess in question we can always form a ne w permutation
which differs from the first permutation in at least its
first element, from the second in at least its second

element from the ( i^ + 1 ) st in at least its

( w; + 1 ) st element, and so on, therefore is not included
in any set of permutations ordinally similar to any
possible arrangement of all the natural numbers.
From this Lemma we readily obtain the

Theorem. Every possible arrangement of the natural
numbers is a series of the second class. For by the
Lemma to every such arrangement in series can be as-
signed a set of permutations forming a series of higher
ordinal type. But by § 97 the process by which this is
eflFected yields a set of permutations forming a series of
type W. Therefore every possible arrangement of nat-
ural numbers in series is of type lower than W, there-
fore belongs to the second ordinal class ( being ipso facto
>w). q. E. D.

410 KANSAS UNIVERSITY SCIENCE BULLETIN.

99. The arrangement of the natural numbers in
series of type higher than w finds appHcation in the study
of infinite continued fractions. By aid of the euclidean
algorithm for greatest common divisor every absolute
irrational number less than unity can be expressed as
an infinite continued fraction

_i i_ i_

qi+ q2+ • • • • q^+ • • • •

and conversely. The class of infinite simple continued
fractions may therefore be taken as the representative
of the class of irrational numbers between zero and 1.
An infinite continued fraction can not be obtained
from a finite one merely by annexing quotients; it can
only be described by assigning a law which determines q^
for every value of N. An infinite continued fraction
may be formed, e. g., by the law that every quotient
shall be 2. It is sufficient, however, to consider the class
in which the quotients are all different; this can be put
into one to one correspondence with the whole class.
Accordingly we are concerned with the class of all
possible permutations of all the natural numbers.
These permutations may be examined in the same way
as the permutations of a finite set by imagining a frame-
work of places to be filled, but the number of places is
infinite. Moreover, we must provide for the possibility
of filling the places in a series of order higher than w.
Thus an infinite continued fraction can be formed by
filling the even places successively with the odd num-
bers in their natural order and the odd numbered places
with even numbers in the same way, i, e.,
J 1 1 1^ 1 1

2+ 1+ 4+ 3+ 6+5+

or by filling the odd numbered places with primes and
the even places with composite numbers. Or we can
select first the places whose indices are primes, then
the indices which are products of two primes, three

FRIZELL: FOUNDATIONS OF ARITHMETIC. 411

primes, and so on, and fill these sets with the correspond-
ing sets of numbers permuted in any way we please.

100. We have seen that every possible arrangement
of all the natural numbers in a series is of type lower
than W. Therefore the quotient places of our frame-
work m the order in which we propose to fill them (or
rather state how they shall be filled ) constitute an or-
dinal type of the second class. That is, the numbered
places in a permutation of all the natural numbers, ar-
ranged, however, in the order in which they are to be
filled, form a series obtainable by the process of ^ 97.*
Hence there can be no permutation of all the natural
numbers not obtainable by this process, since such a
permutation would be eo ipso different from all of
those hitherto obtained and well ordered in some defi-
nite numbered places. In other words, this would mean
that the series of the places as they are filled, or the
series of numbers with which each place is to be filled,
was not obtainable by the process of § 97. Therefore
all possible permutations of the natural numbers form
a series closely related to the series of type W.

BIOGRAPHY.

I, Arthur Bowes Frizell, member of the Protestant Episcopal Church,
was born in Boston, July 14, 1865. My parents were Joseph Palmer Fes-
senden Frizell, civil and hydraulic engineer, and Julia Anna (Bowes) Frizelh
My early education was chiefly at home and in a private school in Dorches-
ter, Mass. I graduated from the high school in St. Paul, Minn., and spent
three years at the Massachusetts Institute of Technology, where I after-
wards served as assistant instructor in mathematics, 1888-'91. I received
the degree of Bachelor of Arts from Harvard College in 1893 and that
of Master of Arts from Harvard University in 1900. I served as instructor
in mathematics at New York University 1895-'96, and at Harvard 1897-1906,
when I resigned to study abroad. After three semesters at Gottingen, I
returned to America, and was appointed, 1908, Professor of Mathematics
in Midland College, Atchison, which position I resigned, 1909, to accept an
instructorship in the University of Kansas.

*And the preceding statements apply verbatim to every series of values
admissible in a given numbered place.

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The Green Bug.

PLATE I.

A. Dakota sandstone resting on Permian shales. Man's head in line w th
contact. Spring creek, six miles north of Washington.

B. Unconformity of Pleistocene on Permian. Ravine running from It ft
to right in Permian shales with thin limestone forming the top cros es
a buried gully at right angles to it filled with Pleistocene mater al,
which is etching backward. Note the tipping of the limestone i>»to
the old gully (undisturbed by recent erosion). About three miles east
of Washington.

.'Science Bulletin, University of Kansas.
Vol. V, No. 1.

PLATE I.

PLATE 11.

.4. Unconformity of Dakota on Permian sandstone at contact projects.

Brickyard, east side of Smoky Hill river, Salina.
B. Same. South end of same hill bv old mill.

Science Bulletin, Univeksity of Kansas.
Vol. V, No. 1.

PLATE II.

^nS-'i -

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B

PLATE III.

-4. Cliff of Dakota sandstone. East side of ]Mill cieek, northeast of
Washington.

B. Slumping of soft sediments, clays apparently of Dakota age. Three
miles east of Washington.

Science Bulletin, University of Kansas.
Vol. V, No. 1.

PLATE III.

PLATE IV.

A. Slumped soft Dakota clays suriounded by impure sandstone, on Per-
mian floor. Three miles east of Washington.

B. Alfalfa field on the wide terrace of the Republican liver, west of
Clay Center.

Science Bulletin, Univerkitv of Kansas.
Vol. V, No. 1.

PLATE IV.

PLATE V.

Fig. 1. — View looking southwest from ruins, showing heavy ledges of
Loup Fork rock in background, from which the pueblo was built. In
the foreground two of the party are shown excavating one of the
circular pits which occur about twenty yards north of the main
building.

Fig. 2. — Ground plan of ruins, showing remains of the division walls,
looking west. Remains of the mealing trough shown in the extreme
corner of room I, to the left of the observer. To the right, in room V,
the small fireplace built into the wall can be seen.

Science Bulletin, University of Kansas.
Vol. V, No. 2.

PLATE V.

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PLATE VI.

Fig. 3. — Typical Pueblo grinding trough, three feet nine inches long by
two feet one inch wide and eighteen inches deep.

Figs. 4, 5, 6, 7. — Awls or drills, used principally for perforating skins
and boring holes into bones, etc. Figure 7 is of a reddish flint, beau-
tifully colored, and of fine workmanship.

Figs. 8, 9, 10, 11, 12. — Bone implements, probably used to enlarge holes in
skins. Have been much used.

Figs. 13 to 18, inclusive. — Pipes and pipe stems; all are made of native
clay. Figure 15 is well made and is decorated with the design shown
on plate VIII, figure 62; mouthpiece broken off. Pipe measures two
and one-half inches long and seven-eighths inch in diameter at the
center.

Science Bulletin, University of Kansas.
Vol. V, No. 2.

PLATE VI.

PLATE VII.

Figs. 19, 20, 21, 22, 23, 24.— Arrow shaft straighteners. These are made
of the Dakota sandstone, and probably came from somewhere in the
vicinity of Quivira, for no outcroppings of this material are exposed
nearer to Quartelejo than those which occur on the western boundary
of Brower's Quivira, about 130 miles east of the Quartelejo pueblo.

Fig. 25. — Bone piccolo, made from the wing bone of a large bird, prob-
ably a pelican.

Fig. 26. — Bone implement, possibly a hoe blade, from shoulder blade of
buffalo.

Fig. 27. — Tallying implement, made from rib of buflFalo.

Fig. 28. — Bone tool with a toothed edge, which may have been used for
fleshing skins.

Fig. 29. — Bone tool, made from rib of buffalo, worn smooth and flat,
spatula-like.

Fig. 30. — Bone implement of undetermined use, much worn, with toothed
edge. Made from shoulder blade.

Figs. 43 to 56, inclusive. — Fleshers or scrapers, of flint. Figures 43, 45,
46, 47, 51, 52, are beautifully made. Figures 46 and 51 show cross
sections of Fusilina, a fossil typical of the Carboniferous. The ma-
terial of these two, at least, is not found in the vicinity of the ruins,
and was probably procured from the eastern part of the state.

Science Bulletin, University of Kansas
Vol. V, No. 2.

PLATE VII.

^ ^

PLATE VIII.

Figs. 57, 58. — Knife blades of yellow chert.

Fig. 59. — One of the most interesting implements found, consisting of the
half of an iron ax head, five and one-half inches long by one and a
quarter wide at its greatest width, and five-eighths of an inch thick.
This, by some mischance, had been split in two longitudinally, and
was arranged with a groove around it instead of an eye like our
modern axes have.

Fig. 60. — A fine grinder of conglomerate, eleven inches long. Shows
much usage.

Fig. 61. — The half of a clam shell, plainly showing saw-tooth marks
where it was divided. This was found at the bottom of the twelve-
inch hole in room V.

Fig. 62. — Diagram of pattern on pipe bowl shown on plate VI, figure 15.
Impression was made on clay and photographed.

Fig. 63. — Base of deer antler, which has evidently been used as some im-
plement.

Figs. 64 to 69, inclusive. — Specimens of charred corncobs. Figure 59
shows some well-developed kernels on it.

Fig. 70. — Bone implement, well worn.

Fig. 71. — Portion of 'dobe plastering from walls, showing impressions of
finger tips.

Fig. 72. — Baked clay marble, evidently used in some game.

Figs. 73, 74, 75. — Chert scrapers.

Fig. 76. — Hoe blade made from shoulder blade of buffalo.

Science Bulletin, University of Kansas
Vol. V, No. 2.

PLATE VIII.

PLATE IX.

Fig. 77. — Small metate or grinder, of sandstone.

Fig. 78. — Grinder made of clay mortar and coarse sand, and baked as
hard as rock itself.

Fig. 79. — Hammer worked from a glacial boulder; well made and much
worn.

Fig. 80. — Grinder, of very fine granite.

Fig. 81. — Grinder, of material same as figure 78.

Fig. 82. — Grinder, made from glacial boulder.

Figs. 83, 85. — Grinders, of fine sandstone.

Fig. 84.— Knife blade of chert.

Science Bulletin, University of Kansas
Vol. V, No. 2.

PLATE IX-

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Science Bulletin, University of Kansas.
Vol. V, No. 4.

PLATE X.

SciKNCE Bulletin, Univeksity of Kansas.
Vol. V, No. 5.

PLATE XI.

Science Bulletin, Umveksity of Kansa«
Vol. V, No. 5. n-AiNj,*..

PLATE XII.

Science Bulletin, University of ICansas.
Vol. V, No. 5.

PLATE Xllt.

SciENcR Bulletin, University op Kansas

PLATE XIV.

PLATE XV.

Fig. 1. Two resting cells, showing chromatin granules, and nucleoli with
the surrounding clear space.

Fig. 2. Cell beginning mitosis. Close spireme. Nucleoli without the
clear space.

Fig. 3. A granular spireme. The nuclear membrane has disappeared
but the cytoplasm can still be distinguished from the nuclear
sap.

Fig. 4. The chromatin granules are here arranged in rows. This cell has
its nuclear membrane yet and the nucleoli have not disap-
peared.

Fig. 5. Polar view; spireme segmented to form the chromosomes, has a
longitudinal split, and is uniformly granular. Nuclear wall
just disappeared. Chromosomes looped to form U and J
shapes.

Fig. 6. A little later than figure 5. The longitudinal split is very clearly
seen.

Fig. 7. Late metaphase. Cytoplasm granular in places, may have been
slight plasmolysis in this cell. The spindle fibers are stained
very lightly here.

Fig. 8. Metaphase. Spindle shows marked polarity; chromosomes some-
what indefinite.

Fig. 9. Metaphase. Cytoplasm at the ends becoming vacuolated and at
the center slightly denser. An especially long chromosome
{m) is to be noted.

Fig. 10. The cytoplasm of this cell was crushed; however, the chromo-
somes, which were undivided yet, show the longitudinal split
clearly.

Fig. 11. Anaphase. Cytoplasm very granular in places (homogeneous
portion not shown). The spindle is well stained in this cell,
and the chromosomes have their characteristic bent shape.

Fig. 12. Anaphase. Spindle not clear in this figure. The amount of
cytoplasm is small compared to the amount of chromatin here.

Science Bulletin, Univekshy of Kans/s
Vol. V, No. 6.

PLATE XV.

\"\

PLATE XVI.

Fig. 13. Anaphase. Massing of the cytoplasm at the center of the cell.
Fig. 14. Anaphase. The long chromosomes (m) are to be seen in this
figure.

Fig. 15. Anaphase. Shows relation of chromosomes to spindle during
this period.

Fig. 16. Anaphase. Spindle fibers do not show here; reticular cyto-
plasm (drawn from a slide loaned by Mr. Alban Stewart) .
Fig. 17. Anaphase. Polar view; 14 chromosomes.
Fig. 18. Same as 17.

Fig. 19. Telophase. Shows the loosening up of the chromosomes. Cell
plate beginning to form.

Fig. 20. Chromatin has diffused throughout the nucleus ahead of the
cell-plate formation. A nuclear membrane has also been
formed.

Fig. 21. Chromatin a dense mass, very indefinite at the ends of the cell,
not shown ; cytoplasm much contracted from the cell wall and
concentrated about the spindle.

Fig. 22. A very peculiar spindle; cytoplasm much condensed; a second
dense spindle formed about the center of the cell where the
cell-plate is appearing. Chromatin not shown.

Fig. 23. Concentrated cytoplasm at the spindle center; a very broad
spindle and the cell plate extends entirely across it. Cytoplasm
vacuolated; the daughter nuclei, not shown in this figure, are
nearly reconstructed.

Fig. 24. Cell plate nearly formed; nuclei reconstructed; cytoplasm con-
centrated at the center. (From slide belonging to Mr. Stewart.)

Science Bulletin, University of Kansas
Vol. V. No. 6.

PLATE XVI.

T^Z

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PLATE XVII.

Koeberlinia spinosa.

Fig. 1. — Branch of Koeberlinia spinosa. x about %.

Fig. 2. — Aspect of vegetation on mesa near Tucson in the neighborhood
from which the specimen was taken.
Photographs by L. M. Peace.

Science Bulletin, University of Kansas.
Vol. V, No. 7.

PLATE XVII.

PLATE XVIII.

Koeberlinia spinosa.

Fig. 1. Cross section of stem. P, pith; M, medullary ray; W, wood;
Tr, tracheal tubes of year '07 (?) ; Ph, phloem; Sa, spongy pa-
renchyma air space; Pa, palisade air space; Ea, epidermal air space;
B, bast; Scl, sclerenchymatous tissue. Semi-diagrammatic, x 25.

Fig. 2. (a) Section from an older stem, showing traces of annual rings,
'07, '05. (b) Portion of same stem, opposite side, showing '06 trace.
Semi-diagrammatic, x 25.

Fig. 3. Cross section of flower branch, showing region of wa, water
storage cells, x 283.

Fig. 4. (a) Pith from flower branch, showing air spaces, A. Cross,
X 283. (b) Single cell from flower-branch pith, showing pits, P;
air spaces, A. Cross, x 400.

Fig. 5. Pith of old stem, showing pitted conditions and starch bodies.
Cross, x 283.

Fig. 6. Crystals in old pith cells; longitudinal section, x 283.

Fig. 7. Diagrammatic representation of regions of older stem. E, epi-
dermis; P, palisade; Sp, sponge; B, bast; S, stone cells; Ph, phloem;
W, wood. Enlarged.

Science Bulletin, University uk Kansas
Vol. V, No. 7.

PLATE XVIII.

P S p B S

7 ^^

PLATE XIX.

Koeherlinia spinosa.

Fig. 1. Medullary rays, showing longitudinal extension, longi-tangential
section. Semi-diagrammatic, x 55.

Fig. 2. Portion of (1) in detail, showing thick walls, some pits, and nu-
mei'ous aerating places. Tangential, x 283.

Fig. 3. Single medullary ray cell from (2), showing food bodies F, B;
pits, P; aerating spaces, A. Longi-tangential. x 636.

Fig. 4. Medullary ray cells, showing one elongated and narrow cell e. c,
and some more, box-like, for storage, St. Longi-radial, x 283.

Fig. 5. Single storage cell from (4), showing food bodies. Longi-radial,
X 636.

Fig. 6. Portion of medullary ray extension in phloem region, showing-
thin, unpitted walls and cells filled with nitrogenous material. Longi-
radial, X 283.

Fig. 7. Poi-tion of older wood, showing tracheids, Tr; fiber tracheids,
Ft, and wood fibers, if/. Cross, x 283.

Fig. 8. Detail of (7), showing pitted condition of wood, x 636.

Fig. 9. Water vessels bordering pith. Ti-, tracheal tubes; Td, tracheid;
P, pith. X 636.

Fig. 10. Spiral tracheids, near pith. Semi-tang., X 283.

Fig. 11. Portion of wood, showing scattered tracheids, Tr. Cross, x 283.

Science Bulletin, University of Kansas
Vol. V. No. 7.

PLATB XIX.

PLATE XX.

Koeberlinia spinofin.

Fig. 1. Tracheal tube bordering- elong^ated pith cell, on margin of older
part of bundle. P, pith cell; Tr, tracheal tube. Longi-radial, x 686.

Fig. 2. Elongated pith cells den.sely filled with finely granular food ma-
terial. Pits in walls omitted. P, pith; W, wood. Longi-section, X
283.

Figs. 3, 4, 5, 6. Different views of tracheal tubes, all showing the cir-
cular opening at the end. 6 (a) Third partition in tube 6, showing
section of ring bordering aperture. These have become so completely
welded as to give the appearance of a continuous tube, as in 6.
Longi-section, X 400. Fig. 3 is the cavity outlined.

Figs. 7, 8, 9. Sketches showing joining of ends of tracheids. Longi-
radial, X 400.

Figs. 10, 11, 12. Various odd or malformed tracheids. Longi-radial,
X 283.

Fig. 13. Regular rectangular tracheids. Longi-radial, X 283.

Fig. 14. Single tracheid. Longi-radial, X 283.

Fig. 15. Fiber-tracheid with small but numerous pits. X 283.

Fig. 16. Wood fiber. Longi-section, x 283.

Science Bulletin, University of Kansas
Vol. V, No. 7.

PLATE XX.

PLATE XXI.

Koeberlinia spinosa.

Fig. 1. Five rows of phloem parenchyma cells p. p., as seen in cross
section; m, medullary-ray cells, x 283.

Fig. 2. Sketch of portion of phloem, Ph, region bordering stone cells, S,
showing various sizes of cells. Longi-section, x 283.

Fig. 3. Portion of (2) in detail, sieve tubes, St., and sieve plates, S, and
parenchyma cells alongside, p. Longi-section, x 400.

Fig. 4. Stone cells bordering spongy parenchyma, showing some cells
unlignified, and some but partly lignified, parenchyma cells; s, .stone
cells; pc, partly lignified; P, unlignified. Cross .section, x 283.

Figs. 5, 6. Different sizes and shapes of stone cells. Longi-section,
X 283.

Fig. 7. (a) Tapering bast fiber, (b) Portion of long bast fiber, showing
tapering part and projections of wall to support neighboring cells,
(c) Projections with adjoining cellulose walls in detail, (d) Pits in
long fiber. All x 283.

Figs. 8, 9, lOo, x 55, 283, 636, respectively. Short bast fibers as they
occur securely fastened together. 106, sketch showing pits in short
bast fiber, x 636.

Science Bulletin. University of Kansas.
Vol. V. No. 7.

PLATE XXI.

PLATE XXII.

Koeberlinia spinosa.

Fig. 1. Portion of palisade P, and spongy parenchyma, Sp, tissue, from
stone cells to epidermis. Numerous intercellular spaces /. S., are
found. Cross section, x 283.

Fig. 2. End view of palisade cells, showing number and size of chlo-
rophyll bodies. Tangential section, x 283.

Fig. 3. Epidermis of young stem, showing large cellular space, cs, and
thin walls; this also shows frequency of deeply sunken stomata, ss.
Cross, X 283.

Fig. 4. Portions of older epidermis, showing reduction of cell space by
intense cutinization ; cell space, cs; cutinized portion, cp. Cross,
X 283.

Fig. 5. Tangential view of older epidermal cells, showing thickening of
walls. X 283.

Fig. 6. Single stoma from young stem, showing large guard cells, Gc.
Also, small stomatal chamber proper, above and below, sc. Cross,
X 283.

Fig. 7. Cuter stomatal chamber invaded by phycomycete, Ph. Cross,
X 283.

Fig. 8. Stomatal apparatus. OA, outermost aperture; E, epidermal cell
space; osc, outer stomatal air chamber; isc, inner stomatal air cham-
ber; Pr, projection of thickening, making stomatal spaces propei',
ssp; P, palisade cells; AS, air space in sponge. Cross, X 283.

Fig. 9. Outline of regions of aerating spaces, showing frequency of
spaces. E, epidermal region; p, palisade region; s, sponge region.
Enlarged.

Science Bulletin, University of Kansas.
Vol. V, No. 7.

PLATE XXll.

AS

PLATE XXIII.

Koeberlinia spinosa.

Fig. 1. S, stone cells filling up lower stomatal chamber; Ph, remains of
Phycomycete in outer aperture; d, portion of "dome." Cross section,
X 283.

Fig. 2. Surface view of outer stomatal aperture and epidermal cells.
SA, stomatal aperture; EC, epidermal cells. Tangential section,
X 283. Compare with fig. 3, plate XXII.

Fig. 3. View toward outer stomatal aperture from within, showing
tips of cells making the "dome." Tangential section, x 283.

Fig. 4. Section across mid-chamber. Tangential, x 283.

Fig. 5. View looking toward stoma, showing shelf, s, made by projecting
walls. Tangential, X 283.

Fig. 6. Stoma proper, showing guard cells. Tangential, X 283. Com-
pare with fig. 6, plate XXII.

Fig. 7. Stoma proper, invaded by growth of Phycomycete, Ph. Tan-
gential section.

Fig. 8. Water-storage cells, W, in flower branch, and aerating space, A.
Cross, X 283.

Fig. 9. Portion of older stem, showing empty cells, W, probably for water
storage. Cross, x 283.

Fig. 10. Water-storage cells, W, and aerating space. A, in flower
branch. Cross, x 283.

Fig. 11. Aerating space, A, in spongy parenchyma. Longi-section, x
283.

Science Bulletin, University of Kansas.
Vol. V. No. 7.

PLATE XXIII.

PLATE XXIV.

Fig. 1. Resinous-gum cells, RD, surrounding vascular bundles in elon-
gated portion of receptacle. Cross, X 283.

Fig. 2. Resinous-gum cells in elongated portion of receptacle, RD. Longi-
section, x 283.

Fig. 3. Resinous-gum cells, RD, surrounding ovule. X 283.

Fig. 4. Section of ovary. X, position of structures shown in figs. 1 and
2. Longi-section, x 5.

Fig. 5. Section showing position of mucilage reservoirs and other tis-
sues in ovary. MR, mucilage reservoirs; MC, mucilage cells; E, epi-
dermis; p, palisade tissue; M, mesophyll; vb, vascular bundle. Longi-
section, X 55.

Fig. 6. Ovary. Reference letters same ais (5). Cross, x 55.

Fig. 7. Portions of (6) more in detail. OV, ovule; S, stoma; for the
other letters, see (5). Cross, X 283.

Fig. 8. Diagrammatic representation of regions in wall of ovary. Ref-
erence figures as above. Cross section enlai'ged.

Science Bulletin, University of Kansas.
Vol. V, No. 7.

PLATE XXIV.

PLATE XXV.

Fig. 1. Portion of ovary wall in detail. OE, outer epidermis; M, me-
sophyll; M?% mucilage reservoirs; IE, inner epidermis; IS, inner
stoma; VB, vascular bundles. Cross, x 283.

Fig. 2. Stoma from inner epideraiis of ovary wall. Gross, x 283.

Fig. 3. Outline of portion of ovary wall, showing frequency of inner
stomata, S. Cross, enlarged.

Fig. 4. Outer stoma, enlarged. Cross, x 283.

Fig. 5. Inflorescence, showing fruit. X 1.

Fig. 6. Single flower stalk. Sepals and petals dropped. Stamen L
longer than stamen S. X 5.

Fig. 7. Single flower, x 5.

Fig. 8. Section of anther, showing number of locules and thin-walled
epidermis. Cross, X 283.

Fig. 8a. Pollen grain, x 636.

Fig. 9. Sunken stomata on outer epidermis of petals, x 283.

Fig. 10. Portion of petal, showinq: thin-walled epidermis. Cross, X 283.

Science Bulletin, University of Kansas
Vol. V, No. 7.

PLATE XXV

PLATE XXVI.

Fig. 1. — Cross section through ovipositor, about midway: A. cmgd,
cement gland duct; ovid, oviduct; uv, upper valve; Iv, lower valve;
tg, tongue and groove; spg, subgenital plate. B. Diagram showing
position of ovipositor relative to plates.

Fig. 2. — Sport anal (?) vein of adult: A, anal vein; C, costa; sv, sport

anal (?) vein.
Fig. 3. — Diagram of tracheation of nymphal wing pad of P. celtidis-

gemma fore wing: C, costa; Sc, subcosta; R, radius; M, media;

Cu, cubitus; af, anal fold; A, anal vein.
Fig. 4. — Diagram of tracheation of wing of recently emerged adult of P.

celtidis-mammse, fore wing.
Fig. 5. — Longitudinal-vertical section through wax glands: A. wg, wax

glands; 7i, nucleus; cu, cuticula; sh, sense organs; tvgd, wax gland

ducts; fb, fat body. B. wgo, openings of wax glands, dorsal aspect.

Fig. 6.— Tip of mandible.

Fig. 7. — Tip of maxilla.

Fig. 8. — Base of mandible: te, tendon.

Fig. 9. — Base of maxilla.

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P"'lG. 1. — Lateral aspect of prothorax and mesothorax, diagrammatic:
pn, pronotum; es 1, proepisternum; es i, px'oepimeron ; pstr, pro-
sternum; pcox, procoxa; up, spiracle; dnr, dorsulum; m?», mesonotum;
scu, scutellum; artep, articulatory epidemes; bp, base of primary;
ew 2, mesoepimeron ; es 2, mesoepisternum; mstr, mesosternum; end,
portion of endoskeleton.

Fig. 2. — Caudal aspect of mesothorax, diagrammatic : • ter, tergum ;
str, mesosternum; phr, phragma; apoph, apophyse; c, canal; apod,
apodeme; es '2, mesoepisternum; em 2, mesoepimeron; end, portion of
endoskeleton corresponding to end in fig. 1, plate XXVII.

Fig. ."J. — Sclerites forming tergum of thorax: A. pn, pronotum of pro-
thorax; dor, dorsulum of mesothorax; mm, mesonotum of mesothorax;
scu, scutellum of mesothorax; mett, metathorax. B. median, longi-
tudinal section through tergum.

Fig. 4. — Chitinized process holding labium to thorax (cephalic aspect) :
A. a, lateral arm of process; h, yoke-like piece connecting lateral
arms, caudad and dorsad of labium. B, tip of a lateral arm.

Fig. 5. — Ventral aspect of metathorax, diagrammatic: sir, meta-
stei'num; cox, coxa; em 3, metaepimeron ; mer, meracanthus; troc,
trochanter; mf, ventral edge of metafurca.

Fig. 6. — Venation of adult of P. celtidis-iimnnnx, fore wing: r, costa;
Sc + R, subcostal plus radius; s, stigma; Sc, subcosta; R, radius;
M 1 2, media; M 3 If, media; Cu 1, cubitus; Cv, 2, cubitus; af, anal
fold or "claval suture"; cl, clavus; A, anal vein; Pr, petiolus cubite.

Fig. 7. — Venation of adult of P. celtidis-mammse, hind wing.
Fig. 8. — Ventral aspect of subgenital plate of female: h, base of plate;
a, apex of plate; sli, sensory hairs.

Science Bulletin, University of Kansas.
Vol. V, No. 9.

PLATE XXVII.

PLATE XXVIII.

Fig. 1. — Caudal aspect of head: ep, epicranium; e, compound eye; fo,
front ocellus; g, gena; fc, frontal cone; a, antennal segments; y, pro-
jection from gena; x, portion of membrane forming fold around
labium between procoxae.

Fig. 2. — Longitudinal section of front ocellus: cu, cuticula; hyp, hypo-
dermis; I, lens; ip, iris pigment; v, vitreous body; r, retinal cells; n,
nerve.

Fig. 3. — Antenna.

Fig. 4. — Cephalic aspect of head: ep, epicranium; e, compound eye; o,
ocellus; fo, front ocellus; g, gena; fc, frontal cone; a, antennal seg-
ments; as, antennal socket.

Fig. 5. — Ventral aspect of head: ep, epicranium; e, compound eye; fo,
front ocellus; g, gena; fc, frontal cone; as, antennal socket; p, pro-
jection from edge of antennal socket; y, projection from gena; f,
frons; Ip, ligamentary process of frons.

Fig. 6. — Articulation between third and fourth antennal segments: a,
third segment; b, fourth segment; cor, coiTUgations; sh, sensory
hairs.

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PLATE XXIX.

Fig. 1. — Lateral aspect of mouth parts, labium excepted: /, frons; c,
clypeus; lab, labrum; ep, epipharynx; md, mandible; tux, maxilla;
mds, mandibular sclerite; mxs, maxillary sclerite; cmx, chitinized
plate for muscular attachment; cp, hypopharyngeal lamella; procx,
maxillary process; lahi, labium (base); h, hypopharynx; for, fora-
men.

Fig. 2. — Lateral aspect of mouth parts (diagrammatic) : /, frons; ep,
epipharynx; s, setse; labi 1, lahi 2, lahi 3, three segments of labium;
mestr, mesosternum; p, process of thorax for holding labium; m, mem-
brane forming fold between procoxae for labium.

Fig. 3. — Ventral aspect of mouth parts, labium excepted: /, frons, c,
clypeus; lab, labrum; ep, epipharynx; md, mandible; ms, maxillas;
cmx, chitinized plate for muscular attachment; cp, hypopharyngeal
lamella; mds, mandibular sclerite; mxs, maxillary sclerite; procx,
maxillary process.

Fig. 4. — Ventral aspect of labium: /, frons; c, clypeus; lab, labrum; ep,
epipharynx; labi 1, labi 2, labi 3, three segments of labium; b, point
where bend occurs.

Fig. 5. — Dorsal aspect of mouth parts, labium excepted: f, frons; lab,
labrum; ep, epipharynx; md, mandible; vix, maxillae; cmx, chitinized
plate for muscular attachment; cp, hypopharyngeal lamella; procx,
maxillary process; h, hypopharynx; t, tentorium; z, proximal edge of
frons; p, pharynx.

Science Bulletin, University of Kansas.
Vol. V, No. y.

PLATE XXIX.

f P I

PLATE XXX.

Figs. 1, 2 and 3. — Lateral, dorsal and ventral aspects of ovipositor,
diagrammatic: iiv, upper valves; Iv, lower valves; vs, upper and
lower valves (fig. 2) ; buv, base or support for uv; blv, base or sup-
port for Iv; sh, sheath; v, valve closing oviduct; procv, process of rod
for closing v; sc, laterally diverging pairs of sclerites cephalad of
sheath (fig. 1) ; sc 1, proximal pair of sclerites; sc 2, distal pair of
sclerites; sip, sting-palpi; mr, median rod; Ir, lateral tendenous rod
flattening out into lightly chitinized lateral plates, Zpf.

Fig. 4. — Diagram showing position of ovipositor: spp, supra-anal plate;
a, opening of anus; sgy, subgenital plate; v, valves of ovipositor; sh,
sheath of ovipositor tip.

Fig. 5. — Lateral aspect of metafurca.

Fig. 6.— Caudal aspect of supra-anal plate: s-pp, supra-anal plate; sppfi,
inner edges or flaps of supra-anal plate; x, distal portion of plate; a,
anal opening; Ip, lateral plate; co, base of copulatory organ; /, loop
below copulatory organ.

Fig. 7. — Lateral aspect of supra-anal plate and copulatory organ : spp,
supra-anal plate; sppfi, inner edges or flaps of supra-anal plate; x,
distal portion of plate; a, anal opening; Ip, lateral plate; co, copu-
latory organ; g, geniculation; c, club; p, penis; /, loop below copu-
latory organ; y, slit in base of copulatory organ.

Fig. 8. — Cephalic aspect of metathorax, diagrammatic: str, meta-

sternum; nif, metafurca; la, lateral projection of endoskeleton ; mett,

metatergum.
Fig. 9. — Section through fore wing: wm, wing membrane; b, pigmented

disc; a, unpigmented disc; c, cuticular processes on ventral surface of

wing.

SciEa*cB Bulletin, University of Kansas.
Vol. V, No. 9.

Pig. 94.

PLATE XXXI.

Fig. 1. — Cross section through procoxae and mouth parts: cox, coxa; c,
clypeus; p and pc, parts of pharyngeal canal; mds, mandibular
sclerite; mxs, maxillary sclerite; .s, setae; rp, hypopharyngeal lamella;
m, membrane forming fold between procoxae to hold labium.

Fig. 2. — Lateral aspect of tip of labium: t, lateral tongue-like processes;
sh, sense organs.

Fig. 3. — Cephalic aspect of tip of labium: /, lateral tongue-like processes;
ah, sense organs; si, slit of labium.

Fig. 4. — Base of supra-anal plate of female showing wax glands: a, anal
opening; ivgo, openings of ducts of wax glands; .s/?, sense organs; .r,
rhitinized portion of plate; y, slightly chitinized, translucent portion
of plate.

Fig. 5. — Cross section of lain 1 : nicl, mandible; mx, maxilla; »i, longi-
tudinal muscles; si, slit of labium; /, lumen in mandible.

Fig. 6. — Cross section of ovipositor through base of .sheath : spp, .supra-
anal plate; sgp, .subgenital plate; nvid, oviduct; sh, sheath; sfp, sting-
palpi.

Fig. 7. — Lateral aspect of metathorax, part of metasternum, epimeron,
meracrnthus, co>a. <:t?.., veriiov(.d. (li?G:vannmatic : Dicff, meta-
tergum; vif, metafurca; bs, region of base of secondary; es 3, region
of metaepisternum ; str, part of metasternum; end, a rod connected
to metafurca; x, point of articulation of em 3.

Fig. 8. — Lateral aspect of metathorax, part of metasternum, episternum
and metafuica removed, diagrammatic: mett, metatergum; mf,
metafurca; str, part of metasternum; mer, meracanthus; em 3, meta-
epimeron; cox, coxa; scl, an accessory sclerite; x and ?/, articulation
of em 3; troc, trochanter; fern, femur.

Science Bulletin, University of Kansas.

Vol. V, No. 9.

PLATE XXXI.

Fig. 36. %n\

PLATE XXXII.

Fig. 1. — Frons and mouth parts, labium excepted, removed from head.
(Photo.)

Figs. 2, 3 and 4, — Wings of male. (Photos.)

Figs. 5, 6 and 7. — Wings of female. (Photos.)

Fig. 8. — Tracheation of nymphal wing pad of P. celtidis-gemma, fore
wing. (Photo.) C, costa; Sc, subcosta; R, radius; M, M, media;
Cu, Cu, cubitus.

Fig. 9. — Tracheation of wing of recently emerged adult of P. celtidis-
mammse, fore wing. (Photo.) C, costa; Sc + R, radius plus sub-
costa; R, radius; M, media; Cu, cubitus.

Fig. 10. — Branching of media and cubitus from radius plus subcosta in
fore wing of recently emerged P. celtidis-mammse : Sc + Rr radius
plus subcosta; M, media; Cii, cubitus.

Fig. 11. — Branching of subcosta and radius in fore wing of recently
emerged P. celtidis-mammse: Sc, subcosta; R, radius; C, costa.

Science Bulletin, University ok Kansas.
Vol. V, No. 9.

PLATE XXXII.

PLATE XXXIII.

Fig. 1. — Apex of wing of male, showing pattern. (Photo.)

Fig. 2.— Hind wing. (Photo.)

Figs. 3 and 6. — Wings of females. (Photos.)

Fig. 4. — Highly magnified portion of pattern. (Photo.)

Fig. 5. — Apex of wing of female, showing pattern. (Photo./

Fig. 7. — Wing of male. (Photo.)

Fig. 8. — Base of hind wing, showing pattern and hairs on costa. (Photo.)

Science Bulletin, TTnivrrsity of Kansas.
Vol. V, No. 9.

PLATE XXXTIT.

PLATE XXXIV.

Fig. 1. Dorsal aspect of male. (Photo.)

Fig. 2. Dorsal aspect of male. (Photo.)

Fig. 3. — Ventral aspect of male. (Photo.)

Fig. 4. — Lateral aspect of female. (Photo.)

Fig. 5. — Pachypsylla on bark of hackberry, showing protective markings.

(Photo.)
Fig. 6. — Head, antennae, and mouth parts. (Photo.)
Fig. 7. — Photograph of head and antennae.

Fig. 8. — Lateral aspect of mouth parts, removed from head. (Photo.)
Fig. 9. — Last four joints of antenna.

Science Bulletin, University of Kansas.
Vol. V, No. 9.

PLAtE XXXIV.

PLATE XXXV.

Fig. 1. — Metathoracic tarsi. (Photo.)

Fig. 2. — Prothoracic tarsi. (Photo.)

Fig. 3. — Mesothoracic tarsi. (Photo.)

Fig. 4. — Dorsal aspect of supra-anal plate of female. (Photo.)

Fig. 5.— Mesothoracic leg. (Photo.)

Fig. 6. — Prothoracic leg. (Photo.)

Fig. 7. — Lateral aspect of copulatory organ. (Photo.)

Fig. 8. — Metathoracic leg. (Photo.) scl, accessory sclerite.

Fig. 9. — Lateral aspect of ovipositor. (Photo.)

Fig. 10. — Lateral aspect of male genitalia. (Photo.) spp, supra-anal
plate; sgp, subgenital plate; co, copulatory organ; f, forceps.

Fig. 11. — Mesothoracic legs with mesosternum. (Photo.) apoph,
apophyse.

Fig. 12. — Prothoracic legs, pronotum, etc. (Photo.) pn, pronotum;
es 1, proepisternum; em 1, proepimeron; cox, coxas.

Fig. 13. — Lateral aspect of female abdomen. (Photo.)

Fig. 14. — Lateral aspect of male abdomen. (Photo.) (Figure inverted.)

Fig. 15. — ^Cephalic aspect of forceps. (Photo.)

Science Bulletin, University of Kansas.
Vol. V, No. 9.

PLATE XXXV.

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PLATE XXXVI.

Fig. 1. Tangential section of Poptilus alba with pits in wall between
medullary ray and phloem parenchyma and between two pa-
renchyma cells.

Fig. 2. Side view of obliquely placed end sieve plate from Populus alba
and with pits between the sieve tubes and the other phloem
cells. Tangential section.

Fig. 3. A pit in a vertical wall between a medullary ray and the phloem
of Popvhis alba.

Fig. 4, Transverse or reticulate strengthening bands on the wall of a
sieve tube in a root of Populus deltoides. Tangential section.

Fig. 5. Transverse bands on the wall of a sieve tube in the root of
Populus delfoides, as seen in section.

Fig. 6. Transverse section in Amorpha fruticosa, showing the radially
elongated medullary-ray cells and others similar to these
sweeping around the narrow strip of phloem and arranged
to conduct food from these radially, especially to the cambium.

DESCEIPTION OF FIGURES.

Plates XXXVI and XXXVII.

Explanatoi-y notes: Magnification is 600, except in figure 6, where it is

250 approximately.
Lettering: st = sieve tube; m = medullary ray; p = parenchyma;

camb = cambium; stch = starch; cr = crystal; pt = pit; n =

nucleus.
Symbols: -^ ) and — > ] = direction toward nearest point of circun^-
ference or in the epidermis,

Science Bulletin, University of Kansas.
Vol. XV. No. 10.

PLATE XXXVI.

PLATE XXXVII.

Fig. 7. Tangential section of Populus alba, showing the "beaded" end
plate of a sieve tube and starch granules in the slime body.

Fig. 8. Tangential section in the root of Populus deltoides, showing one
of the remarkable obliquely placed end plates of a sieve tube in
cross section. The strengthening bands here appear as larger
beads and the perforation of the wall as spaces between
smaller beads. This long plate evidently gives more surface
for the distribution of the pits and possibly better communi-
cation between the two cells of the sieve tube.

Figs. 9 and 10 are from tangential sections of the root of Populus del-
toides, and show the reticulated markings caused by the pit-
ting and banding of the obliquely placed end plates in the
sieve tubes. In figure 10 the sieve tube adjoins a medullary-
ray cell.

Science Bulletin, University of Kansas.
Vol. XV. No. 10.

PLATE XXXVII.

V^h}i

10

PLATE XXXVIII.

Townsendia exscapa.

Fig. 1. View from "Castle Rock Bluffs," Gove county, Kansas, showing
the habitat of Townsendia in the foreground.

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PLATE XXXIX.

Tuivm^endia exscapa.

FlO. 2. Photofjraph nf TDirx^oiiJid in flower, showinj; its habit while

young.
Vw,. o. Photognipii of 'r()iriif<cii(li(i ill fruit, to]) view.

Science Bulletin, University of Kansas.
Vol. V, No. 11.

PLATE XXXIX.

"rW^ ..-Jar ,*«-rf^.,^V5-i .-^3;-, '>4i>Si

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PLATE XL.

Townsendia exscapa.

Fig. 4. Cutinized epidermis of stem, cross section. R, outer tangential

wall; S, inner tangential wall. X 650.
Fig. 5. Coi'k cells, cross section, x 650.
Fig. 6. Collenchyma cells, cross section, x 650.
Fig. 7. Collenchyma cells, longitudinal section, x 650.
Fig. 8. Short sclerenchyma cells, cross section, x 650.
Fig. 9. Short sclerenchyma cells, longitudinal section. X 650.
Fig. 10. Cortex cells, cross section, x 650.
Fig. 11. Cortex cells, longitudinal section. X 650.
Fig. 12. Pith cells, cross section, x 650.
Fig. 13. Pith cells, longitudinal section. X 650.
Fig. 14. Small portion of xylem, cross section, showing: a, tracheids;

b, wood parenchyma cells, x 650.
Fig. 15. Tracheids, as seen in a longitudinal section, when isolated from

the wood parenchyma cells by means of chromic acid. X 650.

Science Bulletin, University of Kansas
Vol. V. No. 11.

^xH

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PLATE XL.

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PLATE XLI.

Townsendia exscapa.

Fig. 16. Medullary-ray cells, cross section. X 650.
Fig. 17. Medullaiy-ray cells, longitudinal section, x 650.
Fig. 18. Phloem cells, cross section. X 650.

Fig. 19. Phloem cells, longitudinal section: a, cambiform cell; b, undi-
vided mother cell of the sieve tubes, x 650.
Fig. 20. Cross section of root, showing the distribution of tracheids.

X 115.
Fig. 21a. A bleached leaf, showing its method of venation, x 121/4.
Fig. 21b. Midrib of leaf, cross section, showing: a, phloem cells; b,

tracheids; c, wood parenchyma cells. X 215.
Fig. 22. Portion of a bleached leaf, showing the details of venation.

X 115.
Fig. 2.3. Phloem cells as seen in a longitudinal section of the midrib of

a leaf: a, cambiform cell; b, undivided mother cell of the

sieve tubes. X 650.
Fig. 24. Cutinized epidermis of leaf, cross section: a, cuticle; b, cutin-

ized portion of wall. X 650.
Fig. 25. Palisade cells, surface view: c, intercellular spaces; d, palisade

cell. X 650.
Fig. 26. Spongy parenchyma cells, surface view. X 650.

SCENCE^BULLEXIN, Univ.«sitv o. KaX..s.

PLATE XLI.

2/ a

PLATE XLII.

Townsendia exscapa.

Fig. 27. Diagrammatic drawing showing the distribution of water-
storage cells as seen in cross section of a leaf: a, veins; b,
water-storage cells.

Fig. 28. Palisade cells bordering on water-storage cells: A, water-
storage cells; B, palisade cells; a, chloroplasts found in water-
storage cells. X 650.

Fig. 29. Palisade cells bordering on the veins: A, palisade cells; B,
border parenchyma cells; C, tracheids. X 650.

Fig. 30. Palisade cells bordering on water-storage cells and storage
tracheids: A, water-storage cells; B, chloroplasts found in
water-storage cells; C, water-storage tracheids; D, palisade
cells. X 650.

Fig. 31. Palisade cells bordering on water-storage cells, which in turn
border on veins: A, water-stoi'age cell; B, chloroplasts; C,
tracheids; D, border parenchyma cells. X 650.

Fig. 32. Palisade cells bordering on spongy parenchyma cells: A, pali-
sade cell; B, spongy parenchyma cell; C, intercellular spaces.
X 650.

Fig. 33. Water-storage tracheids as they appear at the ultimate ends of
the veins: A, veins; B, water-storage tracheids. X 650.

Fig. 34. Water-storage tracheids as they appear in other places in the
leaf: .4, ordinary tiacheids; /?, water-storage tracheids.
X 650.

Fig. 35. Stomata, suiface view, x 650.

Fig. 36. Stoma, as seen in cross section of the leaf, x 650.

Science Bulletin, University of Kansas
Vol. V, No. 11.

PLATE XLII.

i\

34

PLATE XLIII.

Lesquerella spathulata.

Fig. 37. Cutinized epidermis of stem, cross section : S, outer tangential
wall ; T, inner tangential wall, x 650.

Fig. 38a. Cutinized epidermis of stem, longitudinal section : S, outer
tangential wall; T, inner tangential wall, x 650.
Cork cells as seen in cross section, x 650.
Stone cells found in the boi'ke, cross section. X 650.
Collenchyma cells, cross section, x 650.
Collenchyma cells, longitudinal section. X 650.
Stone cells found in sclerenchyma tissue, cross section. X 650.
Stone cells found in sclerenchyma tissue, longitudinal section.
X 650.

Bast fibers, longitudinal section, x 650.
Bast fibers, cross section. X 650.

Small portion of xylem, cro.ss section: A, wood fibers; B, wood
parenchyma cells; C, tracheal elements.

Wood fiber, longitudinal section, x 650.

Tracheal elements found in the stem at the nodes as seen in
longitudinal section: A, scalariform type of tracheids; B,
spiral type of tracheids. X 650.

Fig. 49. Trachea] elements found in the internodes of the stem, as seen
in longitudinal section: A, spiral tracheal tubes found among
the wood fibers; B, spiral tracheal tubes found among the
wood parenchyma cells; C, scalariform tracheal tubes. X 650.

Fig. 50. Diagrammatic drawing of the cross section of the stem: A,
borke; B, collenchyma; C, sclerenchyma; D, phloem; E,
xylem; 1, tracheal tissues; 2, wood fibers; 3, xylem pa-
renchyma; F, medullary ray; a, leaf trace. X 87%,

Fig.

386

Fig.

39.

Fig.

40.

Fig.

41.

Fig.

42.

Fig.

43.

Fig.

44.

Fig.

45.

Fig.

46.

Fig.

47.

Fig.

48.

Science Bulletin, University of Kansas
Vol. V, No. 11.

PLATE XLIII.

m

PLATE XLIV.

Lesquerella spathulata.

Fig. 51a. Spiral tracheal tubes found in the leaf trace. X 650.
Fig. 516. Small portion of the medullary ray, as seen in cross section.
X 650.

Fig. 51c. Phloem cells, as seen in cross section: A, intercellular spaces.

X 650.
Fig. 52. Phloem cells, as seen in longitudinal section: A, intercellular

spaces. X 650.

Fig. 53. Thin-walled parenchyma cells of the cortex, cross section.
X 650.

Fig. 54. Xylem parenchyma cells, as .seen in longitudinal section, x 650.
Fig. 55. Wood fibers and ti-acheal tubes found in the outermost layer of
xylem, cross section. X 650.

Fig. 56(1. Segment of the cross section of the root, showing the consti-
tution of the xylem: A, wood fibers; /?, tracheal elements;
C, wood parenchyma. X 53.

Fig. 566. Tracheal elements found in the root: A and B, scalariform
tracheal tubes; C, .spiral tracheal tube. X 650.

Fig. 57. Tracheal tubes and wood parenchyma cells found in the layer
of xylem (fig. 56a., F) : A, tracheal tubes; B, wood pa-
renchyma cells. X 650.

Fig. 58. Wood parenchyma cells, as seen in longitudinal section, x 650.

Fig. 59. Wood fibers and tracheal tubes found in the layer of xylem
(fig. 56a, G). X 650.

Fig. 60. Diagrammatic drawing of the cross section of the root, show-
ing: A, cork; B, collenchyma ; C, .stone cells; D, thin-walled
parenchyma; E, phloem; h\ wood fibers; G, wood paren-
chyma; H, ti-acheal elements; /, medullary ray. X 65.

Fig. 61. Cutinized epidermis of leaf: .1, outer tangential wall; o, cutin-
ized portion; b, cellulose portion; B, inner tangential wall.
X 650.

Fig. 62. A bleached leaf, showing the method of venation: A, central
vein; B, lateral branches sent off from the midrib near the
center; C, two prominent longitudinal veins found on either
side of the midrib; D, E, F, terminations of veins; G, a large
lateral vein sent out from below the center of th^ midrib,
X 27,

Science Bollktin. University of Kansas.

Vol. V, No. 11.

PLATE XLIV.

PLATE XLV.

Lesquerella spathulata.

Fig. 63. Small portion of midrib of leaf, as seen in longitudinal sec-
tion: A, spiral tracheary tracheid; B, undivided mother cell
of the sieve tubes. X 650.

Fig. 64. Palisade cells, surface view, x 650.

Fig. 65. Trichomes, as seen in surface view of the leaf, x 115.

Fig. 66. Trichome, as seen in leaf, cross section: A, cutmized portion of
wall of trichome; B, cellulose portion of wall of ti-ichome;
C, mesophyll cells; D, tracheids. X 250.

Fig. 67. Stomata, as seen in surface view. X 650.
Fig. 68. Stoma, as seen in cross section, x 650.

Fig. 69. Leaf, as seen in cross section: A, upper epidermis; B, lower
epidermis; C, palisade tissue; D, vascular bundle, x 250.

SciENcK Bulletin, University of Kansas
Vol. V, No. 11.

PLATE 3JLV.

SCIENCE BULLETIN, UNIVERSITY OF KANSAS
VOL. V, No. I 2

PLATE 46

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SCIENCE BULLETIN, UNIVERSITY OF KANSAS
VOL. V, No. I 2

PLATE 47

SCIENCE BULLETIN, UNIVERSITY OF KANSAS
VOL. V, No. I 2

PLATE 48

D

20

. B

y

PLATE XLIX.

Fig. 1. External view of the right mandible of Eryops willistoni. X V2-
Drawing by S. Prentice, a = articular; c = coronoid.

Fig. 2. Maxillary tooth of Eryops willistoni. X V2.

Fig. 3. Fragment of cranium, including portion of right orbit and show-
ing very distinctly the suture bounding the anterior border of the right
frontal. A portion of the supraorbital lateral line canal may be also
detected. X %• O = orbit.

Fig. 4. Cross section of mandibular tooth, showing pulp cavity and dental

tubules. X 2.
Fig. 5. External view of anterior portion of the right mandible, x *^-

SCIENCE BULLETIN, UNIVERSITY OF KANSAS.
Vol. V, No. 13.

PLATE XLIX.

PLATE L.

External view of the right coraco-scapula of Eryops willistoni. x Vz-
C = coracoid portion of the coraco-scapula; / = foramen for artery;
L = cleithrum ; 0 = humeral cotylus.

SCIENCE BULLETIN, UNIVERSITY OF KANSAS.
Vol. V, No. 13.

PLATE L.

PLATE LI.

Internal view of coraco-scapula. C — coracoid portion; / = foramen;
L = cleithrum. x %•

SCIENCE BULLETIN, UNIVERSITY OF KANSAS.
Vol. V, No. 13.

PLATE LI.

PLATE LII.

Arm bones of Eryops tvUUstoiii. X %• H — humerus; R = radius;
U = ulna. Cross sections of the various elements are shown at the
sides.

SCIENCE BULLETIN. UNIVERSITY OF KANSAS.
Vol. V, No. 13.

PLATE Lll.

PLATE LIIL

Fig. 1. Posterior view of right clavicle of Eryops willistoni. X %■

Fig, 2. Ventral view of right clavicle, x %.

Fig. 3. Ventral view of the posterior projection of the interclavicle of

Eryops willistoni. X %•
Fig. 4. Dorsal view of phalange (metacarpal ?). X %•
Fig. 5. End view of same. X %•

Fig. 6. Cross section of posterior projection of interclavicle.
Fig. 7. Anterior view of the right humerus. X %• / = supracondylar

arterial foramen.
Fig. 8. Cross section of the clavicle.

SCIENCE BULLETIN, UNIVERSITY OF KANSAS.
Vol. V, No. 13.

PLATE LIII.

PLATE LIV.

Fig. 1. Posterior view of a dorsal vertebral spine, showing apical en-
largement. X %.
Fig. 2. Lateral view of three consecutive dorsal vertebrae, x %•
Fig. 3. Ventral view of left sacral rib. x %•
Fig. 4. Dorsal view of the same.

SCIENCE BULLETIN, UNIVERSITY OF KANSAS.
Vol. V, No. 13.

PLATE LIV.

PLATE LV.

The dorsal surface of the left ilium of Stegopelta, showing the dermal
plates. X Vi.

SCIENCE BULLETIN, UNIVERSITY OF KANSAS
VOL. V, No. 14

PLATE 65

PLATE LVI.

Fig. 1. A large dermal spine which was possibly associated with the

heavy bony girdles near the base of the tail. X ^^•
Fig. 2. One of the massive dermal plates of Stegopelta, which may have

formed part of another girdle like the elem.ents in figs. 6 and 7. X Vz-
Fig. 3. The median dorsal scute between the ilia. X Vi-
Figs. 4 and 5. The teeth of Stegopelta. x IVz.
Figs. 6 and 7. Dorsal and posterior views of large bony girdle, which

probably encircled the tail near the base, x Vi-
Fig. 8. Detail of the surface of the left ilium; one of the dermal scutes,

one-half natural size.
Fig. 9. A heavy dermal plate, which may possibly have been associated

with the element shown in figure 2 to form another girdle. (See

Wieland, 1909, Amer. Jour. Sci., March, p. 251, fig. 6.) X Vi-

SCIENCE BULLETIN, UNIVERSITY OF KANSAS
VOL. V, No. 14

PLATE 56

PLATE LVII.

Dorsal view of thirteen small dermal plates of Stegopelta, nearly natural
size, showing the various shapes, sizes, and manner of ornamentation

SCIENCE BULLETIN, UNIVERSITY OF KANSAS
VOL. V, No. 14

PLATE 57

PLATE LVIIL

Fig. 1. The external surface of the right pubis, x ^A.

Fig. 2. The right pubis seen from the edge, x ^/4.

Fig. 3. The posterior view of an anterior caudal or a posterior sacral

vertebra. X Vi.
Fig. 4. The lower portion of the right tibia, showing the firm union of

the astragalus. One-half natural size.

Fig. 5. Lateral view of one of the dorsal vertebrje. One-half natural

size.

Fig. 6. Posterior surface of a dorsal vertebra. X Vz-

Fig. 7. A metatarsal of Stegopelta seen from above, x V2-

SCIENCE BULLETIN, UNIVERSITY OF KANSAS.
Vol. V, No. 14.

PLATE LVin.

PLATE LIX.

Fig. 1. Dorsal view of an anterior caudal vertebra. X ^/4.

Fig. 2. A cross section of the sacrum to show the sacral enlargement.

X Vs.
Fig. 3. A small dermal spine. X Vz-

Figs. 4 and 5. Upper and lower portions of right fibula. X Vi-
Figs. 6 and 7. Dorsal surface of posterior and anterior segments of the

sacrum.
Fig. 8. The left ulna seen from behind. One-tenth natural size.
Fig. 9. The left ulna seen from the side. One-tenth natural size.

SCIENCE BULLETIN. UNIVERSITY OF KANSAS.
Vol. V, No. 14.

PLATE LIX.

I

Plate LX.

The stomach of Einpo nepaholica Cope from the Niobrara Cretaceous.
The longitudinal muscular plicae are indicated by heavy lines. xVz-

SCIENCE BULLETIN. UNIVERSITY OF KANSAS
Vol. V, No. 15.

PLATE LX.

Plate LXI.

Tho specimen of Tlni.^fiopater ivtestinalis Moodio. >' %.

SCIENCE BULLETIN, UNIVERSITY OF KANSAS
VOL. V, No. 15

PLATE 6 1

IMS^f

Plate LXIL

Fig. 1. Photograph of the alimentaiy canal of the Buffalo fish, Icfiobds
biibalus Raf., from the Kaw river, showing the similarity in the
form of the intestines to that preserved in the fossil Thrissopater
intestinalis Moodie. X V2.

Fig. 2. Photograph of the right pectoral fin of Empo nepaholica Cope.
X%.

SCIENCE BULLETIN, UNIVERSITY OF KANSAS
VOL. V, No. I 5

PLATE 62

2

PLATE LXIIL

Upper figure: Lateral view of female of Prorocorypha snowl, new gen.
and sp. X 2.

Second figure: Lateral view of male of Prorocorypha snowi. x 2.

Left of three middle figures: Lateral view of apex of male abdomen of
Prorocorypha snowi. X 4. (The supra-anal plate of the specimen
from which the artist made the drawing is broken. It is long, as
in the female.)

Middle of three middle figures: Dorsal view of head and pronotum of
female of Prorocorypha snowi. X 2.

Right of three middle figures: Lateral view of apex of female abdomen
of Prorocorypha snowi. X 4.

Next to bottom row, left figure: Dorsal view of head and pronotum of
Scirtetica ritensis, new species, x 2.

Next to bottom row, right figure: Lateral view of type specimen of
Scirtetica ritensis. X 2.

Bottom row, left figure: Wing of Scirtetica ritensis. X 2.

Bottom row, middle figure: Lateral view of apex of female abdomen of
Scirtetica ritensis. X 2.

Bottom row, right figure: Lateral view of male abdomen of Scirtetica
ritensis. X 2. (This figure was drawn by the artist from a speci-
men not studied by the author.)

VOL. V, No. I 7

PLATE 63

SCitr'"-'

^ x^, , . Spernatophore

Side view DorsalNJvievf Side view ^^ jjemobius

Spermatophores of Gryllus

Spermatophore
Mold

Three Dorsal Dissect ions of the Abdomen of Gryllus
Inters! ine 140

Rectam

Median section of abiomen ^fiio>. Section of sinus and

<^ secreting tubules

Sperm from
spermatophoTe

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SCIENCE BULLETIN, UNIVERSITY OF KANSAS
VOL. V, No. 19

PLATE 64

PLATE LXV.

Figs. 1 to 10, inclusive. — Polar views of the equatorial plate from
spermatogonial cells, each showing the entire number of chromosomes.
In figs. 1, 2 and 4 the possible pairing of equal elements is shown. There
are ten pairs. Nos. 1-1 are the largest two spermatogonial chromo-
somes, which form the largest chromosome of the spermatocyte group.
No. 11 is unpaired, and is the accessory chromosome. In figs. 5 to 10, in-
clusive, the connecting bands of linin are not shown.

Fig. 11. — Lateral view of the spindle in metaphase from a young
spermatogonial cyst.

Fig. 12. — Same from an older spermatogonial cyst.

Fig. 13. — Beginning anaphase from an older spermatogonial cyst,
showing the usual confluent condition of the chromatin.

Fig. 14. — Late spermatogonial anaphase.

Fig. 15.— a section of an entire spermatogonial cyst, showing the cells
in different stages of division. The polarity shown is typical.

Fig. 16.— Early spermatogonial telophase.

Fig. 17. — Polar view of the same, in which all twenty-one members of
the complex can be distinguished, (x) Accessory chromosome.

Fig. 18. — Late telophase, from a young spermatogonial cyst.

Figs. 19 and 20. — Nuclei of spermatogonial cells, showing the chro-
matin at the point of greatest diffusion. The reticular structure is still
evident.

Fig. 21. — A typical spermatogonial nucleus, showing the beginnings of
the reorganization of the nuclear chromatin.

Figs. 22 to 28, inclusive. — Successive stages in the re-formation of the
spermatogonial chromosomes.

Figs. 29 to 33, inclusive. — Nuclei in the process of synizesis. (x) Ac-
cessory chromosome, (t) Nuclear membrane, (c) Chromatin mass.

UNIVERSITY OF KANSAS SCIENCE BULLETIN
VOL. V, No. 20

PLATE 65

^^'V

PLATE LXVI.

Fig. 1. — An early stage of the spermatocyte nucleus, succeeding syni-
zesis. (x) Accessory chromosome, (p) Plasmasome.
Figs. 2 and 3. — Same as fig. 1.

Fig. 4. — Slightly later stage of the spermatocyte nucleus than is shown
in figs. 1, 2 and 3. The chromatin is more diffuse and the accessory
chromosome (x) is elongated, granular, and stains more deeply than the
other chromatin.

Fig. 5. — A spermatocyte nucleus at the period of greatest diffusion.
The chromatin is reticular, (p) Plasmasome. (x) Accessory chromo-
some more condensed than the fonn shown in fig. 4.

Fig. 6. — Same as fig. 5. The accessory is a homogeneous chromatin
rod and the plasmasome reaches its maximum size in this stage.

Fig. 7. — Spermatocyte nucleus in early prophase, showing the first
sig-ns of chromosome reorganization. (p) Vacuolated plasmasome.
{x) Accessory chromosome. The difference in the consistency of these
two bodies is apparent in iron-haematoxylin preparations which were
stained for a shorter time, comparatively, or which were subjected to a
longer extraction process.

Fig. 8. — Spermatocyte nucleus, showing the beginnings of tetrad
formation and the accompanying phenomenon, the decrease in size of the
plasmasome.

Fig. 9. — Later spermatocyte prophase. The plasmasome becomes
spherical as it disappears.

Figs. 10 and 11. — Two parts of the same nucleus drawn from adjacent
sections. There are thirteen bodies, all of which are shown. The plas-
masome has diminished greatly in size, (m) The univalent halves of the
small chromosome.

Figs. 12 and 13. — Spermatocyte cells in early prophase. The nuclear
wall in fig. 12 is not shown. The plasmasome is very lightly stained, in
marked contrast to the accessory, which holds the stain much longer.

Fig. 14. — Accessory chromosomes drawn from spermatogonial cells in
metaphase. For the method of identification see text.

Fig. 15. — Accessory chromosomes from cells of the same stage and
preparation as that of fig. 12, where there is no trouble in identifying
this element.

Fig. 16. — Tetrads from the same stage and preparations as figs. 12
and 13. (a) Largest form. (6) and (c) Typical forms of the cross.
(d) The small chromosome, the halves of which are not usually so close
together at this time.

Fig. 17. — Portion of a spermatocyte nucleus in late prophase, showing
the diminishing plasmasome (p) and the halves of the small chromo-
some (m).

Figs. 18 and 19. — Portions of the same spermatocyte nucleus appear-
ing in adjacent sections, showing thirteen elements — nine tetrads, two
ru-chromosome diads, the accessory chromosome, and the plasmasome.
The plasmasome has almost entirely disappeared.
Figs. 20 and 21. — Same as figs. 18 and 19.
Figs. 22 and 23. — Same as the two previous figures.
Figs. 24 and 25. — Same as above. The nuclear wall in fig. 25 is not
shown.

UNIVERSITY OF KANSAS SCIENCE BULLETIN
VOL. V, No. 20

PLATE 66

12

13

s

U

PLATE LXVII.

Fig. 1. — Nucleus in the process of synizesis. (x) Accessory chromo-
some, la. — Drawings of accessory chromosomes from nuclei similar to
fig 1. A, B, C and D indicate the four individuals from which these
comparative drawings were made.

FiG; 2. — Spermatocyte nucleus, immediately after synizesis, showing
both accessory chromosomes and the plasmasome. 2a. — Accessory chro-
mosomes from the same stage as is shown in fig. 2. 26. — Plasmasomes
from the same nuclei as the accessory chromosomes in 2a. The opposite
elements in columns 2a and 26 are from the same nucleus. The same
holds true for columns 3a and 36, 4a and 46, 5a and 56.

Fig. 3.— Spermatocyte nucleus, showing a later stage than fig. 2.
3a. — Accessory chromosomes from the same stage as fig. 3. 36. — Plas-
masomes from the same stage as fig. 3. See explanation of fig. 2, same
plate.

Fig. 4. — Spermatocyte nucleus at the period of maximum diff'usion
and largest plasmasome growth, deeply stained. 4o. — Accessory chro-
mosomes from same state as fig. 4. 46. — Plasmasomes from same stage
as fig. 4. See explanation of fig. 2, same plate.

Fig. 5. — Spermatocyte nucleus, showing the beginning of chromosome
reorganization. 5a. — Accessory chromosomes from the same stage as
fig. 5. 56. — Plasmasomes from the same stage as fig. 5. See explana-
tion of fig. 2, same plate.

UNIVERSITY OF KANSAS SCIENCE BULLETIN
VOL. V, No. 20

PLATE 67

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PLATE LXVIII.

Figs. 1 to 10, inclusive. — The chromosome complex of the first sperma-
tocyte in metaphase. (x) Accessory chi"omosome.

Fig. 5. — An unusual condition of the first spermatocyte metaphase,
showing twelve elements, a-a. — Probably the disjoined halves of one of
the larger chromosomes which normally occurs in the ring.

Fig. 11. — -Drawings of the accessory chromosome from cells of the
cyst from which fig. 6 was made, showing the constancy in size in this
element within a single cyst.

Fig. 12. — Drawings of the accessory chi-omosome from cells of the
cyst from which fig. 7 was made.

Fig. 13. — Drawings of the accessory chromosome from cells of the
cyst from which fig. 8 was made.

Fig. 14. — Drawings of the accessoiy chromosome from cells of the
cyst from which fig. 9 was made.

Fig. 15. — Drawings of the accessory chromosome from cells of the
cyst from which fig. 10 was made.

Fig. 16. — Lateral view of first spermatocyte in beginning anaphase,
(x) Accessory chromosome. The cell membrane is shown.

Fig. 17. — First spermatocyte spindle, in longitudinal section through
the accessory chromosome {x) . (a) and {b) Chromosomes of the ring
at equal distances from (?h), the central small chromosome. The acces-
sory chromosome lies outside of the ring, causing the asymmetry of the
spindle, as drawn.

Fig. 18. — Same view of a slightly later stage, showing the typical
irregularity in the movements of the accessory chromosome {x) and the
central small chromosome {m) with respect to those of the ring (a)
and {b) .

Fig. 19. — Later anaphase, showing the difference in organization be-
tween the accessory chromosome (.r) and the ordinary chromosomes.

Figs. 20, 21 and 22. — Later successive stages of the first spermatocyte
anaphase, (x) Accessory chromosome.

Figs. 23, 25 and 26. — First spermatocyte telophases, in which the acces-
sory chromosome (x) may still plainly be identified.

Fig. 24. — Telophase of first spermatocyte. (x) Unusual element,
which is probably an undivided accessory chromosome or a persistent
plasmasome.

UNIVERSITY OF KANSAS SCIENCE BULLETIN
VOL. V, No. 20

PLATE 68

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PLATE LXIX.

Figs. 1, 2 and 3. — Polar views of the single daughter groups of chro-
mosomes resulting from the first spermatocyte division. This stage is of
very brief duration following immediately that in figure 25, plate LXVIII.

Fig. 4. — A group of drawings of the accessory chromosome from stages
such as are shown in figures 1, 2 and 3.

Figs. 5, 6, 7 and 8. — Polar views of the second spermatocyte complex.
Note diflTerence in arrangement of the chromosomes compared with those
of first spermatocytes.

Fig. 9. — Oblique view of a late first spermatocyte telophase. One whole
daughter group is shown and part of the other.

Figs. 10, 11 and 12. — Lateral view of second spermatocyte spindle in
metaphase. (x) Accessory chromosome, (m) Small central chromosome.

Fig. 13. — Lateral view of early anaphase, second spermatocyte.

Fig. 14. — Later stage of the same, (x) Accessory chromosome.

Fig. 15. — Same as figures 13 and 14, showing all of the chromosomes.
(x) Accessory chromosome.

Fig. 16. — Same as figure 14.

Fig. 17. — Slightly later stage, showing the beginnings of the formation
of a dividing cell membrane. Note that the daughter gj'oups of chro-
mosomes are entirely coalesced. This is typical of these later stages.

Fig. 18. — Same as figure 14.

Fig. 19. — Group of accessory chromosomes drawn from cells such as
are shown in figure 18, showing the constancy in size of the lagging ele-
ment (x).

Figs. 20, 21, 22, 23, 24 and 25.— Later stages in the second spermatocyte
division, showing the behavior of the accessory chromosome (x) . Note
the dividing cell wall and persistent spindle fibers which identify beyond
a doubt the two cells shown in each figure as daughter cells of the same
first spermatocyte.

Fics. 26 and 28. — Later stage of the same after the formation of the
nuclear membranes, (x) Accessory chromosome.

Fig. 27. — Later stage of the same in which the spindle fibers are
fading.

Fig. 29. — Still later stage in which the two chromatin masses are
breaking up and the ordinary chromatin appears granular, (x) Acces-
sory chromosome.

Fig. 30. — A group of drawings, showing the typical appearance of the
daughter nuclei of the second spermatocyte which have received the ac-
cessory chromosome in the second maturation division.

Fig. 31. — A group of drawings showing the early development of the
spermatid which received the accessory chromosome in the second mat-
uration division.

UNIVERSITY OF KANSAS SCIENCE BULLETIN
VOL. V, No. 20

PLATE 69

®(«^«

PLATE LXX.

These photomicrographs were made with a Zeiss 2mm., 1.40 N. A.
apochromatie objective and No. 4 projection ocular, the camera bellows
being extended so as to place the sensitive plate at a distance of thirty-
one inches above the object. Illumination was furnished by Welsbach
mantle, the light passing through a Watson Parachromatic oil immersion
condenser with the diaphragm set at N. A. 1.0. The original magnification
is 1500 diameters which, in reproduction, has been reduced to 1000
diameters. Considerable detail has been lost in reproduction, especially
in figui-es 3, 7 and 8.

Fig. 1. — Polar view of spermatogonial metaphase, showing 21 chro-
mosomes.

Fig. 2. — Same as figure 1.

Fig. 3. — Same as figure 1.

Fig. 4. — First spermatocyte prophase, in which the accessory chromo-
some is bent in the middle. This is the nearest approach to the spireme
condition found in Anasa.

Fig. 5. — Later prophase with the accessory extended and showing the
longitudinal split.

Fig. 6. — Same stage as in figure 5.

Fig. 7. — -A portion of three cysts appears in this photograph. In the
lower one the last generations of spermatogonial chromosomes are seen
in several cells. At the upper right-hand corner are four cells in
synizesis.

Fig. 8. — Parts of three cysts are included in this picture. The cells in
each are in some stage of synizesis. The presence of the accessory chro-
mosome on the periphery of the nucleus and of the plasmasome in the
synizetic knot is demonstrated in several cells. The stages shown in
figures 7 and 8 precede the ones represented in figures 4, 5 and 6.

Fig. 9.— Post-synizetic stage, in which the peripheral accessory and the
more central plasmasome are shown in several cells. This may be com-
pared with a similar condition in cells prepared according to the method
of Foot and Strobell, as shown in their figures 11 and 12, in order to
judge of the correctness of their belief that the smear method preserves
delicate detail better than sections. It may be pointed out also that there
is little liability to confuse either of the nucleolar bodies with karyosomes
resulting from concentrations of the chromatin elsewhere. Wilson's slide
9496. Photos 1 to 9 from sections.

Fig. 10. — Chromosomes of the first spermatocyte metaphase, smear
preparation. The lowermost chromosome is the accessory. It is seen to
have one plane of division, while the others have more or less distinct
second planes indicated. In this cell the five lower chromosomes have
dried thinner and spread more than the upper six and show wrinkles and
other distortions. Wilson's Woods Hole "x" slide.

Figs. 11-15. — Photographs from sections on Wilson's slide 9496, show-
ing the accessory chromosome and plasmasome in the first spermatocyte
prophase at about the same stage as that of figure 9. It is difficult to

get several nuclei close together with the two elements in focus at the
same time, but most of these show at least two cells. Figure 11 is in
focus at the level of the upper part of the nucleus and shows the ac-
cessory extended as in figures 5 and 6.

Figs. 16-20. — Photographs from smears on Lefevre's slide 30, showing
the late prohases of the first spermatocyte. The accessory is distinguished,
as Foot and Strobell point out, by having two halves lying side by side
without lateral extensions. There is no indication of a second plane of
division, as may easily be seen in figures 16 and 20. In figure 20 there
appear thirteen elements.

Figs. 21-26. These are from smears, and show the chromosomes of the
first spermatocyte in metaphase. The great difference in the size of Lhe
chromosomes, due to the variation in the extent of spreading, upon drying
may be judged by comparing figure 22 with figures 23 and 10. In figure 23
there are twelve elements, one much lighter than the others, which may be
interpreted as an unsually resistant plasmasome which has persisted much
longer than common. In all of these the accessory may be identified both
by its position and by the single plane of cleavage indicated. It has been
injured in the cell represented in figure 23, where it is seen next to the
plasmasome with one corner cut sharply off.

Figs. 27 and 28. — ^Photographs of the first spermatocyte metaphase,
polar view, showing the typical arrangement of the chromosomes. The
accessoiy lies without the ring and the ?H-chromosome in the center.
These prints are from the same negative and show how it is possible to
vary the apparent size of structures even by printing. Wilson's section.

Figs. 29-33. — Polar views of first spermatocyte metaphases from sec-
tions. In order to get several cells to show together the focus was shifted
during exposure in making photograph 31 so that the outlines are not
sharp. In figure 29 the accessory does not show, being just out of focus
in the lower cell. From these illustrations it may be seen that while
there is a general agreement in the arrangement of the chromosomes in
the equatorial plate, it may be modified in details. Wilson's sections.

SCIENCE BULLETIN, UNIVERSITY OF KANSAS
VOL. V, NO. 20

PLATE 70

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30

V

32

33

McCluno. photo

COCKAYNE-BOSTON

PLATE LXXI.

Fig. 1. — Lateral view of two second spermatocytes in early anaphase.
It is apparent in both of these that the accessory chromosome is already
in movement to one of the poles, in the upper cell to the right and in the
lower to the left. The accessory in the upper cell already has its longer
axis parallel to that of the spindle. Smear by Wilson.

Fig. 2. — Lateral view of first spermatocyte in metaphase or early ana-
phase. The accessory is the lowermost chromosome. All the elements are
clearly seen to be divided. This is a slightly earlier stage than the ones
of the second spermatocytes in figure 1 and should be compared with
them in order to see the ditference in the behavior of the accessory in
the two divisions. Smear by Lefevre.

Fig. 3. — Mid-anaphase of the first spermatocyte, showing the full com-
plex of chromosomes in each daughter cell. The accessory is the lower-
most one in each group and is slightly removed from the others. Smear
by Wilson.

Fig. 4. — Late anaphase of the first spermatocyte mitosis. The acces-
sory lies to the right of each daughter group at the same level. Smear
by Lefevre.

Fig. 5. — Mid-anaphase of the first spermatocyte, oblique view. The
accessory to the right of each daughter group.

Fig. 6. — Mid-anaphase of the first spermatocyte anaphase,, lateral view.
The accessory at the lower end of each daughter group.

Fig. 7. — Very late anaphase of the first spermatocyte division, lateral
view, with the accessory chromosome of each group proximal to the
equatorial plate. Smear by Wilson.

Fig. 8. — Late anaphase of the first spermatocyte mitosis, showing the
divided accessory accompanying each daughter group of chromosomes.
Smear by Lefevre.

Fig. 9. — Like figure 8. Smear by Lefevre.

Fig. 10. Similar to figures 8 and 9. Smear by Lefevre. This series
of anaphases of the first spermatocyte indicate, I think, that the accessory
is characterized by an independence of movement and position in the
group of chromosomes rather than by a "lagging" tendency. It does
not in any case, apparently, anticipate the poleward movement of the
other chromosomes, but takes part in the general division. On account
of its peripheral position and slighter relation to the plate of chromosomes
it is crowded back and therefore lies below the daughter mass of chro-
matin. Its position in the metaphase of the second spermatocyte depends
upon the degree to which it is crowded aside in the telophase of the first.
If it maintains its peripheral position at the level of the other chromo-
somes it is then an eccentric element in the metaphase of the second
supeimatocyte. If, however, it lies beneath the other chromosomes in
the telophase it may become included within the ring of chromosomes in
the next metaphase.

Fig. 11. — Polar view of second spermatocyte metaphase. In the lower
cell three chromosomes are shown in lateral view. The difference in the
arrangement of the chromosomes, as compared with the first spermatocyte,
is striking and apparent. Section by Wilson.

Fig. 12. — Similar to figure 11. Compared with the cell shown in the
preceding figure it is seen that the arrangement of the chromosomes is
diffei-ent and that both are unlike that of the first spermatocyte where
the 7?i-chromosome practically always lies within a ring of larger chromo-
somes. The upper cell in figure 11 shows how close the resemblance to
the first spermatocyte may be, on the contrary. Section by Wilson.

Figs. 13-27. — Photomicrographs of smears, mostly from slides by Wil-
son, showing the behavior of the accessory in the second spermatocyte. It
will appear from these, I think, that while the accessory may lie near
the equatorial plate in the early and mid anaphases, when actual division
of the cell has occurred the position of the accessory in only one of the
two daughter cells may be determined. Of particular importance in this
connection is figure 15 where numerous cells in telophase are in focus at
one time. Each one of these demonstrates beyond question the unilateral
movement of the accessoiy. This area is but a small part of a much
largei one on the slide, where more than a hundred similar cells show,
without exception, the same undivided condition of the unpaired element.
The lagging chromosome is recognized by all to be the accessory. It is
seen here in all the divided cells to pass without division into but one of
the two daughter derivatives. In comparing these figures with those of
Foot and Strobell it should be borne in mind that these cells are without
question completely divided, that numbers of them are shown together,
and that the whole cell body is represented, so that there can be no ques-
tion of the relation between the members of the pairs of chromosome
groups. Their figures, on the contraiy, are of single cells, mostly in early
or mid anaphases, in which the cell body does not appear. The con-
sistent behavior of the accessory chromosome in the different cell gen-
erations is worthy of attention also. Wherever it is found it tends to
isolate itself somewhat from the rest of the chromosomes and to behave
with some independence. The argument of Foot and Sti'obell is based
on conformity to one type on the part of the accessory as well as on that
of the ordinary chromosomes. The figures on plate LXXI will demonstrate,
I believe, that the accessory does not conform to the processes of the
other chromosomes. Its isolated position in one of the two daughter
spei-matids shown in figures 26 and 27 is just as apparent as it is in
each of the two daughter spermatocytes shown in figures 3-10. This
characteristic may also be seen in figures 20, 21 and 25. Certain cells
in the stages represented in figures 14, 17, 22, 23 may suggest that the
accessory chromosome might later divide, and it is upon such as these
that Foot and Strobell base much of their argument, but later and more
decisive stages demonstrate that it does not do so and this positive evi-
dence has much more value than the presumptive. It is my belief from
a careful study of the material and the photographs of Foot and Strobell
that not one instance which they figure is an indubitable case of a divide:!
accessory in the second spermatocyte. I do not believe that the in-
dentations in outline, or the light places in the middle of the accessory,
have any value as an indication of probable division, for such effects must
inevitably occur to delicate structures like the chromosomes in smearing.
I would submit that if such evidence is to be used,, the chromosomes of the
upper group in the anaphase of the second spermatocyte represented in
figure 37, plate III of Foot and Strobell, are much more certainly divided
than are any of the accessories for which they claim divisions in figures
26-46 of the same plate,

SCIENCE BULLETIN, UNIVERSITY OF KANSAS
VOL. V, NO. 20

PLATE 71

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Date Due