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i D. Hyatt, of New Rochelle, N. Y.,

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Smaon Henry Gace, B.S., of Ithaca, N. Y.,

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Cuartes E. Bessty, LL.D., of Lincoln, Neb.,

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at Sandusky, Ohio, 1905.

The Society does not hold itself responsible for the opinions expressed

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TABLE OF CONTENTS

FOR VOLUME XXVIII

The Annual Address of the President, The Origin and Development of

the Projection Microscope, by Simon Henry Gage, with plates 1 to

WA, and. €OXt-BOUCES. oo... ed we cee a prsleesce + Sisllaie cs egies eye 5

A Preliminary List of Invertebrates, Parasitic or Otherwise Noxious to

Man, collected in Portuguese West Africa: 1904-1906, by F. Creighton

Wellman oi. e705 © akine, octso sik crorsle rscciRle ison hin Severe areats eee hee 61

Observations on the Micro-fauna of an Oregon Pond, by Elda R. Walker,

with: plate (WIeisss:3 soos sd cle toes oo He wren oie te Sree ae eee eee 75

The Arrhenuri of the United States, by Ruth Marshall, with plates vm to

SKOKDD, aie, toycl oy shot a isle se aie ov uevatie iotuneiian aie yeucttesen ote tenetane cof tteeielin cle es seiiete tenets teen ken nate aaa 85

Further Studies in Volvox, with Descriptions of three New Species, by

J. H.. Powers, with plates xx1il 40 XXVI. . .. 2.02. fc ss ontels oe 141

Data for the Determination of Human Entozoa, II, by Henry B. Ward,

With plate XKVIT. ci. secs 665 20s ocsinte ss orcs dee wig ale orale mate ern a eee 177

Permanent Preparations of Tissues and Organs to show Glycogen, by

Summon entry. (Gages 2 66 2s6c sire sine + 1 evel olsl ais tele 203

Necrology—Frank L. James, Ph. D., M. D., with plate........ . 2 522cc0e 207

Rudolph Siemon; ‘with plate... <5... 4. 6S... Uo ens a sintetee setae eee 210

Notes, Apparatus, Reviews, etc. Weigert’s Iron-Haematoxylin Stain,

by James H. Stebbins, Jr.: Watson and Sons’ New Catalogue: The

Microscopy of Technical Products, by T. F. Hanausek, translated

and ‘revised, by “A. 1. Winton.....05 66 608 6. ale doen soe Ose eee 211

Minutes) of the Annual Meeting: 2. 0.55 cc cc ddities « coee «slate eee ene 215

Mreasuner si NeEporty ears ce ceets sein ae “cine Sie 6.0.9 Shes le mi te eee 219

Custodian’s Report ici). s cec cies oes visi eb eile «oes eee siete ee 220

Gonstitution. | sciic'.. sacctewe dius bonacinve ee aues Cocina: tee tan 221

BS GTA WS hag cinco oie eee bee wpeieso oss (etd were @ ale cee a yoie 8s cca sl epee een 222

List: Of Membersiijisiccs css falc os lice @ tics «nie coven a: ctojlele rt siete tet chetetele eet aie 225

MISE OF “SubScribersiees fics < ae ais csi ave cusie arece a 9 Sunterole is pveiene ta Mae teteke aes ketenes 232

1 by lc Ge en ne PrN te aera Aberin oomoh Oooo OL OT Se 233

TRANSACTIONS

The American Microscopical Society

TWENTY-NINTH ANNUAL MEETING, HELD AT ITHACA, NEW

YORK, JUNE 29 AND 30, 1906

THE ANNUAL ADDRESS OF THE PRESIDENT

THE ORIGIN AND DEVELOPMENT OF THE

PROJECTION MICROSCOPE"

By SIMON HENRY GAGE

WITH SIX PLATES AND NINETEEN FIGURES IN THE TEXT,

INTRODUCTION

No more timely subject, it seems to me, can be discussed in a

presidential address before the American Microscopical Society

than the origin and development of the projection microscope’, since

by the universality of the electric light many can make use of this

most potent means of illustration.

When the task was begun it was not thought that the trail would

lead so far back into the past nor be so obscure and difficult to

follow. In following up the trail it becomes increasingly evident

that the records of human achievement are secondary records only;

1The main part of this address was given with demonstrations at the

meeting of the Society in June, 1906, but the final revision was finished only

in January, 1908. It is hoped that during the present year the work may

be carried to completion, adding to the figures in this address, illustrations

of the best modern apparatus with full directions for installation, and

methods of use; and the numerous applications in teaching and in investi-

gation.

*Some of the names by which the projection microscope is known; it

will be seen that the name is frequently derived from the source of light:

6 SIMON HENRY GAGE

the real individuals who discovered the principles and made the

original, fundamental inventions were often lost sight of, while the

mere recorder or encyclopedist is credited with the inventions and

discoveries which his writings popularized. It is hoped that by

the full notes and references given the reader will gain some ade-

quate notion of the steps of progress, and that, where there is no

certain knowledge, he will also perceive the uncertainty.

The author wishes to acknowledge his indebtedness to Andrew

D. White’s Warfare of Science with Theology in Christendom.

This work gives a broad view of the history of science and with

its numerous references puts one in the way of following an his-

torical search in any field of science no matter how special, like

this on the projection microscope. Without this book and the

numerous old Latin folios and other works in the White Historical

Library of Cornell University, it would have been impossible to

trace so far and so fully the development of the subject of this

address.

In nature one can trace more and more, with increasing knowl-

edge, each result to some antecedent; so in human attainment, when

time has given the necessary perspective, every stage seems not

only logical, but as necessary and unavoidable as the steps in a

mathematical demonstration.

It might have been predicted that the genius of man would in

the fullness of time learn to compensate for the congelation of the

once elastic lens of the eye and the consequent ineffectiveness of the

muscle of accommodation. In this age of artificial helps for

defective eyesight, one can but faintly conceive of the cloud that

in earlier times slowly settled upon the aging human being as the

Camera obscura microscope, Ger., Camera-obscura Mikroskop; electric

microscope, Ger. Photoelectrische Mikroskop, Ital. microscopio fotoelet-

trico; lucernal, lamp or lantern microscope, Lat., lucerna megalographica,

Ger., Lampenmikroskop; laterna magica, or magic lantern; oxy-hydrogen,

hydro-oxygen or gas microscope, Fr., microscope au gaz, Ger., Gasmikro-

skop, Hydrooxygenmikroskop, Ital., microscopio a gaz; picture micro-

scope, Ger., Bildmikroskop; projection microscope, Lat., microscopium per

projectionem, Fr., microscope de projection, Ger., Projectionsmikroskop ;

solar microscope, Lat., microscopium solare, Fr., microscope solaire, Ger.,

Sonnenmikroskop, Ital., microscopio solare.

THE PROJECTION MICROSCOPE 7

crystalline lens gradually lost its elasticity, and near objects could

no longer be clearly seen. Cooper makes one feel this keenly in

his descriptions of the young hero in the Deerslayer, named by the

Indians Hawkeye, and of the same hero in The Prairie with the

weight of years upon him. One voice has come down to us from

the time when this cloud was first brushed aside. It dates from

the year 1299, and comes from Florence in Italy: “I find myself

so pressed by age that I can neither read nor write without those

glasses called spectacles, lately invented to the advantage of poor

old men when their eye-sight grows weak.” (Carpenter-Dallinger,

p. 124.)

Figs. 1, 2. Concave, thick-edged, or divergent, and convex, thin-edged,

or convergent lenses. Spectacles for “poor old men when their eyesight

grows weak” are almost always of convex lenses, while shortsighted eyes

are fitted with spectacles with concave lenses.

‘CARPENTER-DALLINGER, p. 124: “The name of microscope, like that of

telescope, originated with the Academy of the Lincei, and it was Giovanni

Faber who invented it, as shown by a letter of his to Cesi, written April 13.

1625, and which is amongst the Lineei letters in the possession of D. B.

Boncompagni. Here is the passage in Faber’s letter: ‘I only wish to

say this more to your Excellency, that is, that you will glance only at

_ what I have written concerning the new inventions of Signor Galileo; if I

have not put in everything, or if anything ought to be left unsaid, do as

best you think. As I also mention his new occhiale to look at small things

and call it microscope, let your Excellency see if you would like to add

8 SIMON HENRY GAGE

It may be asked, what have spectacles to do with projection

microscopes? Everything, for from the invention and knowledge

of spectacle lenses grew directly the possibility of the invention

of all kinds of optical instruments, including the microscope in all

its forms. The spectacle lenses for “poor old men when their eye-

sight grows weak” means here and generally convex lenses, that

is those which will form real images, and on the possibility of

forming real images by lenses rests the whole development of the

telescope, the compound microscope, and the projection microscope.!

that, as the Lyceum gave to the first the name of telescope, so they have

wished to give a convenient name to this also, and rightly so because they

are the first in Rome who had one’.” PoccenporFF, Gesichte der Physik,

p. 197, says: “Ein Mitglied der Accademia dei Lyncei, ein geborener

Grieche Demiscianus, gab den Fernréhren and Vergrdésserungsglisern ihre

jetzt gebrauchlichen Namen Teleskop und Mikroskop, welche bis dahin

Conspicilia, Perspicilia, Occhiali, Occhialini genannt wurden.” Poggendorff

does not give the date when the Greek, Demiscianus, gave the name

telescope and microscope, so it is difficult to reconcile Carpenter-Dallinger’s

statements and his.

*Porta, Natural Magick: The 17th Book; Of Burning-glasses, and

the wonderful sights by them. Chap. XXI, p. 379. How spectacles are

made: “We see that Spectacles were very necessary for the operations

already spoken of, or else lenticular Crystals, and without these no wonders

can be done. It remains now to teach you how Spectacles and Looking-

glasses are made, that every man may provide them for his use. In Ger-

many there are made Glass-balls, whose diameter is a foot long, or there

abouts. The Ball is marked with the Emril-stone round, and is so cut into

many small circles, and they are brought to Venice. Here with a handle

of Wood are they glewed on by Colophonia melted; and if you will make

Convex Spectacles, you must have a hollow iron dish, that is a portion of a

great Sphaere, as you will have your Spectacles more or less Convex; and

the dish must be perfectly polished. But if we seek for Concave Spec-

tacles; let there be an Iron-ball, like to those we shoot with Gun-powder

from the great Brass Canon: the superficies whereof is two, or three foot

about: Upon the Dish, or Ball there is strewed white-sand, that comes

from Vincentia, commonly called Saldame, and with water it is forcibly

rubbed between our hands, and that so long until the superficies of that

circle shall receive the Form of the Dish, namely, a Convex superficies, or

else a Concave superficies upon the superficies of the Ball, that it may fit

the superficies of it exactly. When that is done, heat the handle at a

soft fire, and take off the Spectacle from it, and joyn the other side of it

THE PROJECTION MICROSCOPE 9

Fig. 3 Fig. 4

Fig. 3. A convex lens showing that when used for producing real

images the object is beyond (below in the figure) the principal focus, and

the real image is formed on the opposite side of the lens and is inverted.

The objectives of a compound microscope, and of a projection microscope

act like this simple lens.

Fig. 4. A convex lens used as a simple microscope or magnifier. In

this case the object is within the principal focus (above in the picture),

and the image appears to the observer to be on the same side of the lens

as the object and not inverted.

From the first production of lenses every kind of experiment was

tried to see what could be done with them. In this period of about

four centuries from the introduction of lenses to the invention of

the compound microscope there was a very close alliance between

to the same handle with Colophonia, and work as you did before, that

on both sides it may receive a Concave or Convex superficies; then rubbing

it over again with the powder of Tripolis, that it may be exactly polished;

when it is perfectly polished, you shall make it perspicuous thus. They

fasten a woolen-cloth upon the wood; and upon this they sprinkle water of

Deparr, and powder of Tripolis; and by rubbing it diligently, you shall see

it take a perfect Glass. Thus are your great Lenticulars, and Spectacles

made at Venice.”

10 SIMON HENRY GAGE

science and magic, or rather science had not yet emerged from

magic. The effort was great to produce striking effects and by

secret devices to make them appear supernatural. In the multi-

tude of trials some one, perhaps several quite independently, found

that a convex lens placed in a small opening of the shutter of a

darkened room gave a much more brilliant and distinct image of

the outside world than did a chance hole in the shutter when no

lens was used.

A darkened room or cabinet with a white wall or screen, a

convex lens in the wall or shutter, and a brilliantly lighted object

or landscape outside form a camera obscura or magic lantern

(laterna magica). A projection microscope is simply a magic lan-

tern in which the lens or system of lenses in the wall or shutter is

of relatively short focus and the image on the screen is much larger

than the object.

The magic lantern and its various uses were described by many

writers between 1500 and 1700. L. da Vinci, who died in 1519,

spoke of a Benedictine monk, dom. Panunce, who used a camera

obscura. Porta in his book, Magie naturalis, 1553, described the

camera obscura', and was so successful in its employment that he

‘Porta, Natural Magick: The 17th book; Of Burning glasses and the

wonderful sights by them. Pp. 364-5. How in a Chamber you may see Hunt-

ing, Battles of Enemies, and other delusions: “Now for a conclusion I

will add that, then which nothing can be more pleasant for great men, and

Scholars, and ingenious persons to behold; That in a dark Chamber by

white sheets objected, one may see as clearly and perspicuously, as if they

were before his eyes, Huntings, Banquets, Armies of Enemies, Plays, and all

things else that one desireth. Let there be over against that Chamber,

where you desire to represent these things, some spacious Plain, where the

Sun can freely shine: Upon that you shall set Trees in Order, also Woods,

Mountains, Rivers, and Animals, that are really so, or made by Art, of

Wood, or some other matter. You must frame little children in them,

as we use to bring them in when Comedies are Acted; and you must

counterfeit Stags, Bores, Rhinocerets, Elephants, Lions, and what other

creatures you please; Then by degrees they must appear, as coming out

of their dens, upon the plain: The Hunter, he must come with his hunting

Pole, Nets, Arrows, and other necessaries, that may represent hunting:

Let there be Horns, Cornets, Trumpets sounded; those that are in the

Chamber shall see Trees, Animals, Hunters, Faces, and all the rest so

THE PROJECTION MICROSCOPE 11

‘

got the reputation of being a wizard, a reputation not quite so

safe and complimentary then as now. Cellini in the middle of the

16th century described the phantasmagoric images he saw projected

upon smoke in the Colosseum at Rome. In the second edition of

his work, Ars magna lucis et umbre, Kircher discusses at consid-

erable length the magic lantern; some of his figures are reproduced

in this address. Dechales says in the Mundus mathematicus, that

in 1665 a learned Dane showed him a magic lantern (text-fig. 6).

Zahn in the Oculus artificialis, 1685-6, figures and describes several

forms of magic lantern. From the above it is seen that a knowl-

edge of the magic lantern was widely diffused during the 16th and

17th centuries. No one knows who the first inventor was. Prob-

ably several individuals devised it in various degrees of perfection.

It has been the custom to ascribe its invention to Bacon, to Porta,

plainly, that they cannot tell whether they be true or delusions: Swords

drawn will glister in at the hole, that they will make people almost afraid.

I have often showed this kind of Spectacle to my friends, who much admired

it, and took pleasure to see such a deceit; and I could hardly by natural

reasons, and reasons from the Opticks remove them from their opinion,

when I had discovered the secret. Hence, it may appear to Philosophers,

and those that study Opticks, how vision is made; and the question of

intromission is taken away, that was anciently so discussed; mor can there

be any better way to demonstrate both, than this. The Image is let in

by the pupil, as by the hole of a window; and that part of the Sphere,

that is set in the middle of the eye, stands instead of a Crystal Table. J

know ingenious people will be much delighted in this. It is declared more

at large in our Opticks. From hence may one take his principles of de-

claring anything to one that is confederate with him, that is secret, though

the party be far off, shut up in prison. And no small Arts may be found

out. You shall amend the distance by the magnitude of the Glass. You

have sufficient. Others that undertook to teach this, have uttered nothing

but toyes, and I think none before knew it.”

In this paragraph Porta describes the camera obscura, and: while he

does not here explicitly state that the hole into the darkened room has a

lens in it, one can feel sure that a lens was there, for he compares his camera

with the eye, the pupil corresponding “with the hole in the darkened room.

the sphere set in the middle is the crystalline lens of the eye. Then in the

paragraph on the preparation of spectacles (lenses) he says “without these

glasses no wonders, like those already spoken of, can be done.” Such a

camera obscura involves all the fundamental principles of projection whether

with a single lens in the shutter or the most complex microscopical outfit.

12 SIMON HENRY GAGE

to Dechales, to Kircher and to other writers whose works have

come down to us. It would be just as correct to ascribe the inven-

tion of the telegraph, the discovery of radium and the many other

wonderful things of our period to popular writers in magazines

or to editors of encyclopedias.

In nearly all the old writings in which the magic lantern is dealt

with, the subject is discussed and references are made to their prede-

cessors and to other books as if what they were discussing had

been known for a long time. If one reads Porta’s preface (see the

Bibliography) one could hardly ascribe to him the invention or

discovery of the wonderful things described in his book and he

nowhere claims such distinction. So in the works of Kircher,

Dechales, Zahn, and others one is constantly reminded of modern

encyclopedias, that is of collections and expositions of what is

already known or imagined.

THE PROJECTION MICROSCOPE

To deal now more specifically with the particular form of magic

lantern known as a projection microscope, it is, as stated above,

simply a magic lantern with a short-focus objective serving to show

small objects greatly enlarged. To comprehend the purpose of a

projection microscope it must be remembered that the microscope,

whether a simple magnifier or the most elaborate compound micro-

scope, is an aid to the eye and becomes for the time being a part

of the visual apparatus of the person using it. But the social and

teaching instincts could not be satisfied without being able in some

way to share the pleasure derived from the exquisite forms revealed

by the microscope. Hence there was an effort to do away with

the virtual or individual image seen on looking into a microscope,

and to produce real images on a white screen where all in the room

could see at the same time, and in which the leader could point out

parts as in a large picture while the members of the audience could

ask questions and discuss points, being sure that every one knew

what was under discussion.

In some cases drawings or colored paintings were made on metal

mirrors and properly arranged before a lens in the shutter of a

darkened room. The mirror being set at the proper angle reflected

THE PROJECTION MICROSCOPE 13

the light through the lens into the room and upon the screen where

the drawing on the mirror appeared more or less perfectly (text-

fig. 5; pl. 1). Sometimes a thin coat of honey was spread over

Fig. 5. Diagram showing the method of producing magic lantern pic-

tures by painting the object to be exhibited directly on the mirror. Com-

pare plate 1 from Kircher. S/, rays of the sun; M, mirror; O, object (an

arrow) on the mirror; D, hole in the shutter of a darkened room; Ob,

objective; J, image (inverted on the screen).

the mirror and then living insects placed upon it. The honey

prevented the too rapid locomotion of the insects and increased

their efforts... If one repeats this experiment at the present time

*KIRCHER, p. 794. Caput vu. De scenica, seu historica repraesenta-

tione rerum: “Si vero muscas vivas exhibere desideres; limbus speculi

melle illiniatur, & ecce muscarum per superficiem speculi quaqua versus

gradientium umbrae in murum projectae vivas ibidem, sed insignis magni-

tudinis muscas repraesentabunt. Hoc idem artificium per magentem ex-

hiberi poterit; mam muscae, vel aliae quaevis res acu instructae ductum

magnetis ex posteriori parte speculo applicati, quocumque artifex voluerit

sequentur. Certe hae repraesentationes adeo arcanae sunt, ut nisi modus

expresse spectantes doceretur, vix quispiam Magicae artis suspicionem

evadere posset.” In this place Kircher describes the use of honey on the

mirror to show living insects. He also speaks of the use of a magnet be

hind the mirror and the necessity of an explanation to the spectators to

avoid the suspicion of magic. The text with the figures from Kircher

(Pls. 1, 1 and 11) will give a good idea of his general treatment of the

subject.

14 SIMON HENRY GAGE

he will have no difficulty in appreciating the effect such exhibitions

must have had in lessening the tedium of those old days.*

Dechales says concerning the magic lantern of the learned Dane

(1665) that it is a veritable microscope, and for many purposes far

better than an ordinary microscope. Zahn also, in the Oculus

artificialis, describes the use of the magic lantern for showing small

objects and says it is a kind of miscroscope. While Kircher did

not specifically call the magic lantern a microscope, he used it for

the purpose of showing small insects on the screen.

As early then as 1665 the laterna magica was used for microscopic

projection and it was recognized as a kind of microscope.

Sometime before 1736 Fahrenheit of thermometer fame produced

a projection microscope which was seen by many people in Am-

sterdam. Among these were Lieberktthn the anatomist. Naturally

it appealed greatly to him as a means of demonstrating the fine

structures he was so skillful in preparing. He had a projection

microscope made and took it with him on his visit to England

(1737-8). In England it created great interest, and as this was the

first knowledge the English opticians and scientific men had of

the projection microscope they very naturally attributed its inven-

tion to Lieberkthn. This error has persisted up to the present day

in English and French works on the microscope (see Mayall, p. 41;

Chevalier, p. 65), although as shown above, Deschales and Zahn

had used and recognized the magic lantern as a microscope over

70 years before.* In justice to Lieberktthn it should be said that

nowhere in his writings does he claim to be the inventor.

In the middle of the 18th century the projection microscope with

its essential features was a well known and much used instrument

*In my own work with the projection microscope and in the prepara-

tion of this address, I have repeated all the principal experiments of the

older workers, with, to use the expression of a previous generation, “much

edification and no little enjoyment.”

*"DECHALES, Mundus Mathematicus, T. 01, Dioptricae, Liber 1, p. 680.

Propositio tvu, Theorema. Exigui prototypi, uniea lente convexa ampli-

ficatam imaginem, in pariete exhibere. “Videmus hic Lugduni dioptricam

machinam, sub nomine laternae magicae, é qua radii luminis, per tubum

uno circiter pede longum erumpentes, in pariete 10 aut 12 pedes distante

THE PROJECTION MICROSCOPE 15

both in England and on the Continent, not only for popular exhi-

bitions but for scientific purposes.

The crude forms figured by Dechales, Kircher and Zahn con-

tained all the essential elements: A radiant to illuminate the object,

a convex lens or a combination of lenses to project the image; and

a darkened room or a cabinet with a whitened surface or screen

to receive the projected image.

All the efforts since the fundamental invention of the camera

obscura have been directed towafd improving the method of light-

ing and perfecting the projecting lenses and the mechanical means

for facilitating the working of the instrument.

exigui prototypi imaginem suis coloribus illustrem, & mirum in modum

amplificatam exprimebant.” Dechales shows in this proposition that one

can produce the same and even better projection with a single than with

two convex lenses (see fig.'6) and concludes that these two lenses of the

magic lantern in fig. 6 act like a single lens.

Ibid. Liber 11, p. 696, Propositio xx, Problema. De nocte exigui pro-

totypt ingentem in muro imaginem distinctam exhibire duabus lentibus.

“Jam superiori libro (p. 680) indicavi eruditum Danum, hoc anno 1665

Lugduno transiisse, qui in dioptrica bene versatus inter alia laternam

[magicam] exhibuit, é¢ qua erumpebat tubus unius circiter pedis, -

Tubus duabus lentibus convexis instructus erat [fig. 6]. . . . . Primo

quo murus in quo exprimenda erat imago magis distabit, eo etiam major

erat imago . . . . Tertio imaguncula in laterna inversa erat, ut sui

efigiem erectam in opposito pariete exhiberet, ablata imaguncula solus

apparebat circulus integer lucidus.” In the magic lantern of the learned

Dane with two convex lenses (fig. 6), the large image of the object will

be erect if the small object is inverted, and the more distant the wall or

screen the larger will be the image. If the object is removed there will

be a circle of light on the wall.

Ibid. Liber u1, p. 698, Corollarium [to proposition xx]. ‘“Microscopium

habes in hujusmodi machina [fig. 6] quod tamen ad usum revocare poteris

sine illa. Si enim tubo eadem vitra inseras nempé primum digitorum 5,

secundum digitorum 10. primdoque imponas muscam aut quodctimque ob-

jectum minutum, tum illud ita soli obvertas, ut transmittatur solis radius

in opposito pavimento; habebis illius objecti imaginem. Nam solis radius

idem praestat quod lumen a speculo reflexum.” In this corollary Dechales

states emphatically that in this machine (fig. 6) one has a microscope

which can be put to greater use than an ordinary microscope; further, that

by turning the tube toward the sun the sunlight serves for illumination

instead of light reflected from a mirror.

16 SIMON HENRY GAGE

Fig. 6. Diagram from Dechales (p. 697), showing the laterna magica

of the learned Dane (1665). It is described as a veritable microscope,

the two lenses acting as a single lens to produce the enlarged image

(p. 680). Toy magic lanterns precisely like this are on sale at the present

time. G-H, concave speculum or mirror to concentrate the light of the

lamp (O-N) on the object at (A-B); C-B, E-F; two convex lenses serv-

ing to form the image on the screen (K-L). The two lenses are movable

and the closer the screen the farther apart are the lenses placed. The

farther off the screen the larger the image and the closer are the lenses

brought together. By discarding the lamp and turning the tube directly

toward the sun good images are also produced (p. 698). (See fig. 7.)

LIGHTING

As shown by the quotation from Porta (1589) and by the figures

copied from Dechales and ‘Kircher (figs. 6-8), both sunlight and

lamp light were used from the earliest times. As also seen, mir-

rors, concave and plane, were used to reflect the light upon the

object or into the instrument.

Kircher and Deschales used convex mirrors for lamp light.

Dechales also states that a good light may be obtained by omitting

the lamp and directing the instrument toward the sun.

The apparatus taken to England by Lieberkiithn was without a

mirror and must be directed toward the sun (fig. 7). Cuff, a

London optician, devised a method used to this day whereby the

apparatus remained stationary and the light of the sun was reflected

into the instrument by a movable plane mirror (fig. 8; pl. 1v, A).

THE PROJECTION MICROSCOPE 17

Fig. 7. Diagram of the projection microscope or magic lantern with

the object illuminated by turning it directly toward the sun as described

by Dechales and Lieberkiihn. S/, rays of the sun; D, diaphragm or hole

in the shutter; O, object; Ob, objective; J, inverted image.

Fig. 8. Diagram of the projection microscope or magic lantern in

which the apparatus remains stationary and the sun’s rays are reflected

into it by means of a plane mirror. This shows also the presence of a

condensing lens to concentrate the wide beam of light from the plane mirror

upon the object. SJ, sunlight striking the plane mirror (M) and being re-

flected through the hole in the shutter (D) to the condenser (C) and through

the stage (S) to the object (O). Ob, objective; J, inverted image.

The instrument was set up on the sunny side, on the south if pos-

sible, and the mirror elevated and turned to the right or left to

catch the sun’s rays and reflect them in the desired direction. A

crcular motion was given the mirror to counterbalance the earth’s

18 SIMON HENRY GAGE

rotation by turning the mirror directly or indirectly by a worm-

screw, rack and pinion or some other means inside the shutter.

Instead of a mirror to be moved by hand a heliostat is now fre-

quently employed and the reflection of the sun’s rays is held con-

stant in the desired direction by the clock-work of the heliostat.

There is no doubt about the superiority of sunlight for illumination

with the projection microscope. All other lights appear weak

beside it. As the sunlight is not to be depended upon in many

parts of the world, and of course is not available in the evening

when many lectures must be given, great effort has been made to

find a substitute. Lamps burning various forms of oil have been

and still are used. Such light is naturally vastly inferior to sun-

light. An artificial light approximating sunlight in brightness was

the great desideratum. In 1801 Dr. Robert Hare of Philadelphia

opened the way to the discovery of such a light by the invention

of the oxyhydrogen blow-pipe, by means of which a much greater

heat could be obtained than in any way previously known. Dr.

Hare also pointed out. that in striking contrast with a flame of

oxygen and carbon, the flame of burning oxygen and hydrogen is

not in itself brilliant, and adds in a note: “The inferiority of the

light emitted by the flame of the hydrogen and oxygen gases to

that which irradiates from bodies exposed to its action, adds one

to the many instances in combustion in which the quantity and

color of the light do not seem to be so much dependent on the

quantity of oxygen gas consumed as on the nature of the sub-

stance heated or burned.’

HARE, RosBert, Jk. Memoir on the supply and application of the blow-

pipe. Philosophical Magazine, xiv (1802): 238-245; 298-306, pl. vr. Also

Annal. de Chemie, xiv (1802); 113-138. In this memoir Dr. Hare de-

scribes his invention of the oxy-hydrogen blow-pipe, and discusses the

comparative heat obtained by it and by other means, and says, p. 301, “More

caloric ought to be extricated by this than by any other combustion.” On

p. 303, with reference to the light and heat: “However, it is worthy of

notice that the light and heat of this combustion do not become evident until

some body is exposed to it from which light may be reflected or on which

the effect of the heat may be visible. This is not the case with combustion

supported by oxygen and carbon.”

THE PROJECTION MICROSCOPE 19

Within twenty years after the invention of the oxyhydrogen

blow-pipe by Hare, experiments with it showing the intense heat

and the splendor of light produced by it when the flame was directed

against refractory bodies like lime became a part of the best chem-

ical courses, as for example those of Brande and Faraday given

at the Royal Institute in London.

The real discoverer of the lime light was then Dr. Hare; but I

have not been able to find out who first utilized the light for illu-

mination. According to Andrew Pritchard in the Micrographia

of Goring and Pritchard, the oxyhydrogen or lime light was used

by Birkbeck in 1824 with a large magic lantern to illustrate a lecture

on optical instruments at the London Mechanics Institution.?

J. T. Cooper, an eminent chemist, assisted Mr. Birkbeck at the

lecture. In 1832 this same Dr. Cooper was giving public exhi-

bitions with a projection microscope, using the oxyhydrogen or

lime light as a radiant.

Capt. Drummond, a young Scotch engineer engaged in the trigo-

nometric survey of Great Britain, attended the lectures of Brande

"According to Andrew Pritchard (Goring and Pritchard’s Micro-

graphia, pp. 170-171). “The oxyhydrogen microscope so attractingly ex-

hibited in the present day, and unquestionably meriting all the encourage-

ment that can possibly be bestowed upon it by the promoters of rational

instruction, may be defined to be a mere modification of the solar [micro-

scope] adapted to receive, and employ to the greatest advantage, the rays

of an artificial light diverging from a central point, instead of the parallel

rays from the sun. In the year 1824, Dr. Birkbeck delivered two lectures

on optical instruments at the London Mechanics’ Institution; in one of

which he took occasion to delineate on a screen, by means of a large magic

lantern, representations of magnified objects intensely illuminated by the

light emitted during the combustion of lime by hydrogen and oxygen gases,

and to indicate the practicability of applying successfully this method of

illumination to the microscope. I would not omit, however, to mention,

that, about the same time, Mr. Woodward instituted some experiments with

the phantasmagoria, where the light was obtained in the same way. In

the interval between that and the present time [1824 and 1837], various

amateurs and artists have studiously exercised their talents in perfect-

ing the several parts of the instrument, which, like the solar [microscope]

assumes its name from the source whence the light requisite to its action is

derived [oxyhydrogen gas microscope].” In a note, p. 171, Pritchard says

“Mr. Cooper assisted Dr. Birkbeck in this experiment [with the lime light].”

20 SIMON HENRY GAGE

and Faraday and saw the brilliant light produced by intense heat

on lime. He conceived the idea of using such a light for signaling

in this survey work, and in 1826 published in the Philos. Trans.

Roy. Soc., p. 324, the method of producing a brilliant light by the

use of an alcohol and oxygen flame directed against lime. In 1830

(Philos. Trans. Roy. Soc., p. 383) he abandoned the alcohol and

adopted the oxyhydrogen blow-pipe for heating the lime. This

brilliant light was applied by Drummond to light houses. From

the importance and publicity of this application of the lime light

by Drummond it came to be called the Drummond light, although

he was not the discoverer of the light, and it had been adapted to

magic lantern illumination six years before he applied it to light

houses. On the Continent the lime light was also much used for

the projection microscope.’

*As a further note on the history of the projection microscope in Eng-

land, and especially the development of the oxyhydrogen gas microscope

the following, from the Microscopic Journal and Structural Record, vol. 1

(1841), is appended: “A brief sketch of the rise and progress of micro-

scopic science, and the principal means enumerated which have tended to

its general advancement.—By the Editor, Dr. Daniel Cooper.

“The first and most important attempt to develop to the public gaze

the microscope on a large scale, was made by Mr. Carpenter of Regent

St., who for many years exhibited a solar microscope for the gratification

of the public. The uncertainty, however, of the weather and the state

of the atmosphere generally in this country, and more especially in the

metropolis, was the great obstacle to this exhibition. This difficulty, at

first sight insurmountable, was at length overcome by Mr. J. T. Cooper

[eminent chemist], who had for many years applied for private purposes

the oxyhydrogen gases projected on lime (generally known as the oxy-

hydrogen light) as a means of illustrating in his laboratory and lectures

many of the important facts connected with light.

“At a meeting of a few scientific friends to witness the results of some

experiments with this light, at Mr. Cooper’s laboratory, then at the Alders-

gate street school of medicine (twelve years since, 7. e., in 1830), Mr.

Cooper and John Carey of the Strand, feeling assured of the principle and

stability of the application, proposed to apply this substitute for the solar

rays to the illustration of microscopic power and accordingly arrangements

were made, and a microscope constructed, adapted expressly to the peculiar

nature of the light, which, as is well known, differs in many respects from

that received from the sun. The first microscope (an experimental one)

was opened in the Strand, in the year 1832, nearly opposite the end of Nor-

THE PROJECTION MICROSCOPE 21

The electric are light which, as shown by Davy (Philos. Trans.

Roy. Soc., 1821), could be produced by using carbon terminals

with strong currents from batteries, was also much experimented

with. In 1845 Donné and Foucault constructed a hand-feed electric

lamp to hold the carbons and Chevalier adapted a projection micro-

scope for use with it. Owing to the difficulty of keeping the light

at a uniform intensity with this lamp, efforts were made to construct

one in which the distance between the carbon terminals and conse-

quently the length of the electric arc should remain practically con-

stant. Such so-called automatic arc lamps were constructed in 1848,

1849 and 1850 in both England and France. While the automatic

lamps are very convenient, even at the present day many workers

prefer the hand-feed lamps, as they are not so liable to get out of

order.

In the original arc lamps the carbons were vertical as in ordinary

street lamps. For general lighting this answers very well, but for

folk street; this spot was selected on account of the contiguity to Mr.

Carey’s workshops as a matter of convenience only. When by dint of

much time and experimental application Messrs. Cooper and Carey had

accomplished their labors to their satisfaction, the scientific public, it will

be remembered, were invited to attend at 21 Old Bond street, on the 18th

of February, 1833, to witness the first public exhibition of the kind ever

presented, in which the oxy-hydrogen light was made to perform all that

had been hitherto effected with direct solar light [for the projection micro-

scope]; and it is but justice to those gentlemen to affirm that this exhibi-

tion was considered to be, both by scientific men and the public at large,

not only most creditable to the labors of the projectors, but the most in-

teresting and important that had ever been offered to the public, and which

could not fail to attract the attention of persons in every age, rank and

station in life; but by possessing the noble aim of enlarging the views of

the multitude by drawing their attention to the wonderful and beautiful

adaptations of nature to secure her end. No exhibition was for a period

better attended than was this; others in the course of a short time sprang

up in various parts of the metropolis and the provinces, and two are even

daily exhibited at the galleries of Practical Science in London, forming

the leading attraction, and exciting the general interest and amusement

of those who visit these institutions.

“The application then of the hydro-oxygen light to microscopic pur-

poses by Messrs. Cooper and Carey in place of the very uncertain means

(solar light) by Mr. Carpenter, created at this period a very general taste

for microscopic science.”

29 SIMON HENRY GAGE

Fig. 9. Front and side views of the carbons of an arc light with inclined

carbons; -++ and — indicate the positive and negative poles. Compare

fig. 17 and pl. v.

A is a side view showing the carbons in section at an angle of 30 de-

grees from the vertical and the negative (—) or lower carbon slightly in

front of the positive (+) or upper carbon. The carbons have soft cores.

B is a front view of the carbons as seen projected on the screen with a

42 mm. objective. It is a projection image of the carbons. This figure

shows that the source of light is the crater in the positive (+-) or upper

carbon; it shows also that the lower carbon is slightly below the upper

carbon as well as slightly in front. This avoids a shadow from the lower

carbon.

In the center of the crater is shown a slight shadow. This is due to

the pit formed in the soft core of the carbon.

THE PROJECTION MICROSCOPE 23

the magic lantern or the projection microscope much light is lost.

To overcome this defect concave or parabolic reflectors have been

used as in the early forms with oil lamps (fig. 6; pl. 11). Every

position of the carbons was tried and many careful experiments

made for determining that best suited for the projection microscope.

An angular position was found advantageous by Lewis Wright, who

induced a London maker to construct a lamp for him with the

carbons at an angle of 30 to 40 degrees from the vertical (fig. 9).

Finally Mr. Albert T. Thompson of Boston employed (1894) car-

bons at a right angle in his arc lamps (fig. 10). In lamps with

Fig. 10. Diagram of a simple arrangement for projection. R, right-

angled carbons with the illuminating crater in the upper carbon; C, con-

denser, the first lens being a meniscus; S, stage; O, object; Ob, objective;

I, inverted image. There is no water-bath either with the condenser or the

stage.

the carbons at an angle of 30 to 40 degrees and at right angles the

current should always pass from the upper to the lower carbon,

thus giving the brilliant crater in the upper carbon, that is in a

position whence the light extends most directly to the condenser.

Lamps with right-angled carbons are being used more and more

for projection.”

*The following is the statement of Mr. Albert T. Thompson concerning

the 90° arrangement of the carbons in an arc lamp for the magic lantern:

Boston, Dec. 6, 1907.

“Replying to your valued communication of the 2d, I will state that I

first manufactured the 90° arc lamps in 1894 and a careful search of all arc

lamp and stereopticon catalogs published about that period, fails to show arc

lamps of the 90° construction.

24 SIMON HENRY GAGE

As the crater in the horizontal carbon is the source of light it is

desirable that it should be equally luminous at all times with a given

current, and it should have a constant position. With a given cur-

rent the equal luminosity depends on the length of the arc. In

hand-feed lamps this simply requires the proper amount of atten-

tion on the part of the operator. In the automatic form the device

for striking the arc when the current is turned on should retain

the upper carbon in the horizontal position. A device for auto-

matically striking the arc and retaining always the horizontal posi-

tion of the upper carbon was worked out and applied to his lecture

room lamps of the Thompson pattern, by Prof. Wm. A. Anthony

of the Cooper Union, New York. In the picture of his lamp (pl.

Iv, B), this device is shown by his permission. The arc-striking

device is formed by links giving a hinge or parallel motion.

For microscopic and ordinary lantern projection, and especially

for high power work, it is exceedingly desirable that the crater,

which is the source of light, should remain in a fixed position. With

solid carbons it shifts around the end of the positive carbon more

or less. To avoid this soft-cored carbons were devised. The soft

core seems to serve as a kind of guide, and the crater remains much

more constant in position than with the solid ones.

The modern arc lamp is a source of light which serves for the

projection microscope very well. While not as brilliant as sunlight,

the results are so uniform and the light so ready at hand at all

hours of the day or night that it has practically superseded all other

radiants.

For the magic lantern the alternating current is fairly satisfac-

tory, but more or less noisy. For both the magic lantern and the

projection microscope the constant or direct current is much to be

preferred. For micro-projection if one cannot use sunlight or the

“T did not patent the lamp, for at that time there was no demand for

them, and of course it was difficult to look into the future and realize that

in a few years thousands and thousands would be sold.

“The facts to the best of my knowledge and belief were never published

in any scientific journal.

Yours very truly,

A. T. THompson.”

THE PROJECTION MICROSCOPE 25

constant current electric light it is hardly worth while going to the

trouble and expense of installing the necessary apparatus.

CONDENSERS

In the earliest magic lanterns the light was concentrated upon the

object by a concave mirror or by means of a convex lens called a

condenser. With sunlight a condenser was often omitted, the sun-

light being allowed to fall directly upon the object (fig. 5; pl. 1).

In the more modern forms of apparatus dating from the time of

Fahrenheit, Lieberktthn, Baker, Adams, Goring and Pritchard,

Chevalier, etc., some form of condenser for concentrating the light

upon the object was practically always used, whatever might be

the source of light. In the early form the condenser was usually a

simple convex lens. With the fuller possibility of rendering lenses

achromatic great pains were taken to make the condensers as nearly

achromatic as possible. It was well understood too that in changing

from sunlight to artificial light of any kind a different form of con-

denser was necessary. If a single lens or combination is used for

sunlight with its practically parallel rays, an artificial light with

divergent rays requires at least two lenses, the one next the light

to collect and render the divergent rays from the radiant approxi-

mately parallel and another lens or combination to concentrate the

rays upon the object. Several forms are shown in the accompany-

ing figures. The one composed of two plano-convex lenses with the

Fig. 11. Various forms of condensers. From Lewis Wright, Optical

Projection. The form E is very commonly employed.

convex surfaces facing each other is common (fig. 11). At present

there is much used a condenser of the Zeiss’ pattern consisting of

26 SIMON HENRY GAGE

three lenses: a meniscus with the concavity next the radiarit and

the other two lenses plano-convex with their convex surfaces facing

each other as in the one just mentioned (figs. 12, 17). Such a

system requires a near approach of the meniscus to the radiant (8-10

centimeters) and there is danger of breaking the lens unless it is

mounted with especial care. Pritchard (1837) recommends a plate

of mica between the first lens and the radiant to avoid breakage.

Mica is not perfectly homogeneous and transparent and so lessens

somewhat the brilliancy of the light, but it seems to be effective,

as it gives time for the more gradual heating of the meniscus (fig.

12). Mica is still often used to protect the condenser.

aos ve a

Fig. 12. To show the elements of the illuminating and condensing

apparatus used in fig. 12, pl. v, A, B, and the position in the cone of light

of objects of various sizes. This figure shows the necessity of moving

the stage to accommodate objects of various sizes and ensure their complete

illumination by the entire cone of light from the condenser. 1, focus of

the condenser where objects for high powers are placed; 2, 3, position of

larger objects; 4, front lens of the condenser; 5, condenser water-bath;

6, 7, two lenses of the condenser next the radiant; 8, the dotted line over

the meniscus represents the sheet of mica which serves to prevent the too

rapid heating of the lenses. This is very satisfactorily held in position

by a cap of sheet iron or copper.

In the earlier forms of projection microscopes the object was

put somewhere in the cone of light made by the condenser, depend-

ing of course on the size of the object and the objective used

(fig. 12).

Substage Condenser.—In the fourth edition of George Adams’

Micrographia (1771, pp. vit-v111) he describes as a “principal im-

THE PROJECTION MICROSCOPE 27

provement in the solar microscope” a secondary condenser placed

very near the object. His son in his Essays on the Microscope

(1787) figures this secondary, or as it is now called substage, con-

denser (figs. 14, 18; pl. 1v, A). A substage condenser is recom-

mended also by Goring (1837) and he advocates in strong terms

that it should be achromatic, as do also Brewster and many others.

At the present day the Abbé substage condenser and sometimes

also a special achromatic form are often used. By some a more

elaborate system still is advocated, viz., a plano-concave lens in the

path of the converging cone to make the rays parallel, and then a

substage condenser to concentrate these parallelized rays upon the

object (fig. 13).

Fig. 13. Wright’s Projection Microscope. C, condenser of three lenses;

A, alum or water cell for removing the radiant heat; P, plano-concave of

highly dispersive glass correcting largely the aberrations of the condenser,

and rendering the beam parallel; SC, substage condenser (for low powers

but a single lens is used) ; S, stage; O, object—next the object is the ob-

jective, and at the end of the tube is an amplifier, 4M; R’, rack and pinion

for focusing the condenser; R’*, coarse adjustment of the microscope; F, fine

adjustment of the microscope.

In my own experience, although several forms of substage con-

denser have been thoroughly tested, the projection microscope has

been more successful without a substage condenser. Furthermore

the substage condenser prohibits the use of a proper stage water-

28 SIMON HENRY GAGE

bath for keeping the specimens cool (figs. 16, 17), and greatly

limits the size of objects which can be projected.

Fig. 14. Projection Microscope with substage condenser and ocular

showing that the substage condenser shortens the cone of the main con-

denser, and that the field lens of the ocular acts with the objective to pro-

duce a real image, and that the eye-lens or combination projects this

real image on the screen. As the objective inverts the object, so the eye-

lens inverts the real image and the final screen image is erect like the

object. R, radiant—in this case an arc lamp with right-angled carbons;

C, condenser, consisting of two plano-convex lenses with the convexities

facing each other; Sc, substage condenser. This brings the rays to a focus

sooner than the main condenser would focus them. This shortening of the

condensation cone enables one to shorten the apparatus, as the object can

be brought closer to the main condenser. S, stage; O, object; Ob, objective;

Oc, ocular, with the field-lens (fl) aiding the objective in the formation

of a real image (J), and the eye-lens (e/) serving as a second objective to

project the real image as the final erect screen image (J*).

REMOVAL OF HEAT

The early users of the projection microscope with sunlight dis-

covered that living objects were soon killed in the concentrated

cone of light used to illuminate them, and that delicate objects were

liable to be destroyed. In the works of Baker (1744-1785) and

Adams (1746-1785) their directions for using the solar microscope

caution the operator not to put living and delicate objects in the

focus of the illuminating cone, but to put them in the cone before

the focus is reached.

In 1837 Goring and Pritchard took a very important step in

advance by advocating that the delicate living things, when shown

by the projection microscope, be mounted in plenty of water, and

THE PROJECTION MICROSCOPE 29

that a water-bath be placed in the cone of light from the sun or

the lime light.

They also recommend that the slide containing the delicate living

objects in water be placed just in front of the water-bath, and re-

mark that in this way the heat is greatly reduced. At about this

time Melloni published his important paper “On the free transmis-

sion of radiant heat through different solid and liquid bodies” (Ann.

de Chem. et de Physique, L111:1 and Lv:337). It was shown that

alum absorbs about 90% of the heat. Solutions of alum are also

found to absorb most of the heat. From that time onward solu-

tions of alum were interposed between the radiant and object to

remove the heat. It is interesting to note that the first heat absorb-

ing bath used by Pritchard was water. As the careful thermometric

experiments of E. Nichols have shown (Physical Review, 1:14;

1893), water is a better absorber of radiant heat than solutions of

alum. It is also far less troublesome, and is now almost always used.

With the great impulse to employ the projection microscope

which came in with the electric light there was naturally a much

greater number of workers striving to perfect the instrument and

render it applicable to the needs of a modern department of biology.

One of the difficulties mentioned by the older observers still re-

mained, and delicate and living objects were injured in spite of

the condenser water-bath put in the course of the illuminating cone

(fig. 12). With the perfecting of condensers the light was also

more exactly brought to a focus. This made the use of great mag-

nification possible but at the same time that the light was focused

the heat rays were also greatly concentrated, and so many passed

through the water-bath that delicate and living objects could be

studied and exhibited with high powers for only a very short time.

Fortunately the living things to be studied are mostly mounted in

water and this serves as a special heat absorber. For many prep-

arations, however, there is no such protection and they are soon

ruined, as every one knows who has undertaken this kind of micro-

scopic demonstration.

To overcome the difficulties incident to the presence of much

radiant heat with high powers when the object must be nearly in

the apex of the illuminating cone, Selenka (1887) used a blast of

30 SIMON HENRY GAGE

air from a rubber bag to cool the specimen while it was under

observation. Carbon dioxid has also been proposed, but so far as

I know has never been actually utilized for the purpose.

In 1893 Zoth (Zeit. wiss. Mikr., p. 154) adapted and modified

a Stricker’s warm stage to keep the slide cool by direct conduction

as well as by. absorption, by circulating cold water in the apparatus

instead of warm. This is a realization for modern conditions of

what Pritchard and others secured in their time by mounting the

specimens in water, and is more generally applicable as not every-

thing can be put in water. Leitz, to avoid the necessity of circu-

lating water, used a stage water-bath with a neck-like process ex-

tending through the stage to the level where the slide rests (fig. 15).

Fig. 15. Letiz’s stage water-bath. In A it is shown below the stage;

in B it is shown in section. F, stage; G, water-bath. It is held in place by

springs.

In 1904-5, after using Leitz’ specimen cooler, the writer became

convinced that the cooling of specimens by conduction was of as

great or greater importance than the absorption of the radiant heat

by the stage water-bath. ;

As it is necessary in a biologic laboratory to project objects of

greater length than the usual diameter of the stage opening, a stage

THE PROJECTION MICROSCOPE 31

Fic. 16, A. Sectional view

B of the stage with the stage

water-bath. S, stage in sec-

tion; Sw, stage water-bath;

gsf, glass front for the stage.

It has attached to it the stage

water-bath.

Fic. 16, B. Opening in the

stage 8 centimeters square; C,

stem of the stage in the socket

of the sliding piece (D); E,

sectional view of the lathe bed

with its V’s to fit the grooves

of the sliding pieces supporting

the condenser, stage and mi-

croscope. See pl. v, a and zw

32 SIMON HENRY GAGE

water-bath like that in the accompanying figure (fig. 16) was

devised. It is of sufficient size to fulfill the requirements of modern

projection; viz., from objects near the size of lantern slides down-

ward to the smallest that can be successfully shown (fig. 16, a).

The front is of the full size of the stage. It is made by cementing

with Canada balsam a piece of glass to an oblong museum Jar about

75 mm. wide, 75 mm. high and 37 mm. thick. The museum jar

should of course be polished on both sides. One may have such

a glass stage-front and a water-bath constructed of thin plate glass,

by the fusion method employed in Germany. The slide then always

rests its full length against the cool water-bath and heat is con-

ducted away from the slide. The additional water also serves to

absorb some of the radiant heat from the illuminating cone. Such

a combination of condenser water-bath and stage water-bath answers

for nearly all purposes. For a uniform and more complete control

of the heat in the illuminating cone, Dr. Greenman, Director of the

Wistar Institute of Anatomy and Biology (Anatomical Record,

1907, p. 170), has devised a condenser water-bath with a hollow

jacket around it. Cold water is circulated through the jacket and

serves to keep the water-bath cold. He also uses the stage water-

bath for the more delicate objects and when high powers are used,

especially when the objects must remain a considerable time in the

field as for drawing.

Zeiss, Reichert, Beck, A. T. Thompson & Co., and others have

supplied condenser water-baths in which cold water circulates in

the bath itself. Strange quivering figures appear on the screen

occasionally on account of the difference of density when the cold

and the heated water mix.

The following table is introduced to show the heating of the

condenser and the stage water-baths in actual work with different

currents and in different positions of the water-baths:

THE PROJECTION MICROSCOPE 33

TABLE SHOWING THE RISE IN TEMPERATURE OF THE Two HEAT-ABSORBING

WaATER-BATHS OF THE ProJECTION MICROSCOPE SHOWN IN PLATE vy

(A direct or constant current of 110 volts at the dynamo was used for

all the experiments. The amperage varied as indicated in the first column

of the table.)

Temperature of| Temperature o! v%

Amperes Time condenser wa-| stage water- Position of water-baths

ter-bath (Cw.) | bath (Sw.)

A Start ily (Ge Ge (C Cw. as in ply, B;° Sw:

10 Amp. for low power (pl.

v, A)

30 min. Bie C De ACS

1 hr. 45°C. 285. ACs

1% hr. 53° .€. oh ee

2 hrs. 69m € B57.

B Start 2ary G Sw. for high powers

15 Amp BS (G. (pl. v, B)

45 min. 64°C: 280 +G:

2 hrs. O7caG: S6cnG:

G Start 22° C. 22° C. |Sw. for high powers;

15 Amp Cw. in front of the

: condenser

30 min. se:5°. C: Pineal Bh.

thr. 26006: Aare (Ce

D Start 22° C. 92° ¢. |Water-baths as shown

20 Amp. in pl. v, B

30 min. Ge (Ce age (C.

Ahr: 91° €, 315.

This table shows very clearly that the heating of the water-baths depends

upon the amperage used. It also shows that the stage water-bath is heated

about the same for the high power as for the low power position. The

condenser water-bath is heated very much more when on the metal saddle

between the lenses of the condenser than when in front of the condenser and

unconnected with it.

From the arrangement of the different parts of the apparatus

it is plain that the condenser water-bath must get a great deal of

heat from the metal saddle on which it rests. In the less compact

form of Zeiss for example where the support for the water-bath is

independent and receives no heat by conduction, the water-bath

will become heated only or mainly by the absorption of radiant

heat from the illuminating cone. The great advantage of such a

device as Dr. Greenman’s for circulating cold water around the

34 SIMON HENRY GAGE

condenser water-bath can be seen from the above table, especially

in B and D, where the experiment lasted a considerable time or

where the amperage is high.

It is a fact easily understood that there is great difference in

objects for projection. Some colors transmit freely the radiant

heat while others absorb the heat and soon the specimen is ruined.

Specimens stained with carmin are especially favorable for pro-

jection. Thick sections stained with hematoxylin absorb the heat

very rapidly. Osmic acid specimens are also unfavorable as they

absorb much heat and can be used only for a short time, especially

in high power projection.

PROJECTION OBJECTIVES

The earliest objectives for projection were simple convex lenses.

If one is inclined to underrate the results obtainable in low power

magnification with them let him take any convex lens, say one of

those in the cheap tripod magnifier and he will be astonished at the

excellence. Of course there will appear around the edges the

color fringes seen in all uncorrected lenses.

Naturally the early workers, full of enthusiasm for the pro-

jection microscope, added improvements as they arose. The early

achromatic objectives (1823 and onward) were applied to the pro-

jection microscope with greater or less success, but as the aper-

tures were rather small the loss of brilliancy seemed a greater defect

than the color fringes of the uncorrected lenses. Among the first to

really grasp the principles governing the projection microscope and

to discuss them in a masterly way were Goring and Pritchard (1837)

and Chevalier (1839). It is true that Euler, Brewster, and others

had discussed from the theoretical side the application of achroma-

tism to the projection microscope, but theory rarely works out just

right in practice because so many factors not duly considered by

the theorist, are sure to obtrude themselves in actual work.

In ordinary microscopic observation the curvature of the field

does not greatly trouble the observer for the attention is con-

centrated upon a very small part of the field and the fine adjustment

brings into perfect focus the part to be especially studied at any

given moment. With the projection microscope the image is like a

THE PROJECTION MICROSCOPE 35

great picture and is far more satisfactory if it is distinct over its

whole extent. Then one of the greatest advantages of the projection

microscope is that it shows objects as wholes. For the purposes of

general morphological study as in embryology and in the study of

the large sections of the nervous system for fiber tracts and rela-

tions, the large field shown by the projection microscope is its

greatest advantage.

Among the first compound objectives to give good results were

those made on the principle of the Ramsden oculars (Brewster,

p. 109). Lewis Wright found lenses of the photographic type

made on the Petzval system to work well. For large embryologic

preparations, large sections of the central nervous system and other

large specimens, photographic objectives of 100 to 120 mm. equiva-

lent focus serve a good purpose, the “planars” of Zeiss giving par-

ticularly good results. For such specimens Messrs. Howland

Brown and Morris Earle devised a variable objective called a

mediascope to indicate its function between the ordinary lantern

slide objective and microscopic objectives.

To secure flatness of field and brilliancy of image many opti-

cians have made special objectives for projection. As early as

1874 Zentmayer produced excellent ones. They had the metal

mounting blackened outside as well as inside so that there should

be no confusion arising from internal reflections and no blinding the

eyes of the operator by reflection from polished surfaces on the

outside.

Many opticians now produce excellent projection objectives.

They are sometimes adapted only for the projection microscope, but

more often they are adapted for both screen projection and for

photo-micrography. The projection objectives of higher power than

about 10 mm. equivalent focus have not proved satisfactory in my

hands. For all powers, and especially for the higher ones, objec-

ives for ordinary microscopic work are used. Not all objectives

which answer well for ordinary microscopic use are satisfactory

for projection. One must test many and select those best adapted

for the work. As stated by Lewis Wright, the price of the objective

is no guide to its excellence for projection.

36 SIMON HENRY GAGE

USE OF OCULARS

The objectives prepared for ordinary use are corrected to give

a real image at a distance of 160 or 250 mm., and it could hardly

be expected that they would give equally good real images on a

screen at a distance of 5 to 10 meters. To use these objectives for

projection and preserve the conditions for which they were cor-

rected many workers use oculars. When the ocular is used if it is

Huygenian in form or any form of negative ocular, the field-glass

acts with the objective to form a real image, then the eye-glass

or lens at the upper end of the ocular acts like another objective

to project the real image upon the screen. If the Huygenian ocular

is used the final image on the screen is formed by a simple convex

lens. Whatever the form of ocular the screen image is erect as

the objective inverts the first image and the ocular inverts the

real inverted image, restoring it to its original position. To fully

appreciate this one must have a translucent screen like ground glass

and look at the back of it, otherwise the image will look as it does

when looking on the wrong side of a printed sheet.

For photographic and projection work Zeiss devised a special

projection ocular in which the upper or eye lens is a corrected com-

bination. This gives more perfect screen images than a simple

lens such as is found in an ordinary ocular (fig. 14).

Fig. 17. Diagram of a projection microscope with a triple lens con-

denser (C) with water-bath (W) ; a stage water-bath (Sw), and an

amplifier (4). The dotted lines indicate the course of the outer rays if no

amplifier were used. AR, radiant; S, stage; O, object; Ob, objective.

THE PROJECTION MICROSCOPE 37

The difficulty with all forms of oculars is that the brilliancy of

the image is much lessened, but the greatest defect is the restriction

of the field of view. A large field is one of the greatest benefits

afforded by the projection microscope, but the ocular does away

with that advantage, and the object must be moved to bring in

small fields in succession as with the ordinary microscope.

AMPLIFIER

Among the numberless experiments tried in the early develop-

ment period of the microscope some one hit upon the idea of

employing a concave lens above the objective to diverge the rays

and therefore to increase the magnification (figs. 17, 18), hence

the name “amplifier.”

According to Petri (p. 136), Conradi, 1710, used a double con-

cave lens above the objective. In the achromatic microscope of

Selligue, as made by Chevalier, 1823-24, there are two draw-tubes.

In the upper of these is the ocular and in the lower a biconcave

amplifier. Two such lenses were furnished with the microscope

for different powers.

In the projection microscope made and figured by Chevalier,

1839, an achromatic combination is used as an amplifier (fig. 18).

As shown in the accompanying figure from Wright, he also em-

ployed an amplifier consisting of a plano-concave lens at the end

of the short tube of the projection microscope, no ocular being

used (fig. 13).

For ordinary demonstration purposes the author has found the

amplifiers —5 diopters and —10 diopters a great addition to the

accessories of the projection microscope. With the —5 diopter

amplifier the size is increased about one-half, and with the —10

diopter amplifier about once, making the images one and one-half

and twice the size that they are when only the objective is used.

To get this amplification with objectives would lessen the light

greatly and also cut down the field; the use of oculars would also

reduce the light greatly and cut down the field still more. (See

under the 8 mm. objective in the following table; see atso pl. Va.)

38 SIMON HENRY GAGE -

TABLE SHOWING WHAT CAN BE DONE WITH THE PROJECTION MICROSCOPE SHOWN

IN PL. v, A AND B, WITH A SCREEN DISTANCE OF 8 METERS, A DIRECT

CURRENT OF 110 VoLTs aT THE DyYNAMO, AND THE AMMETER, BY THE

ProJECTION APPARATUS READING 12 TO 15 AMPERES

Screen

Objective Ocular | Amplifier Field Magnifi: image Remarks

105 mm. None None | 50-60 mm. |x 80 4000 mm. ee tube

pl. v, a

75mm. | None | None | 70mm. |X 105 7350 mm. ay

Planar

50mm. | None None | 45mm. |X 160 7200 mm. 4

Planar :

30mm. | None None 13mm. |X 260 3380 mm. eae as in

pl. vw

30mm. | None |—5d. | 13mm. |X 390 5000 mm. i

30 mm. None |—10 d. 13mm. |X 520 6760 mm. so aca

12.5mm. | None None | 45mm. |X 600 2700 mm. to eee

12.5mm.| None |—5.d. | 45mm. |X 900 +| 4050mm. cS

12.5mm.{ None |—10 d. | 45mm. |X 1200+) 5400mm. aes

8mm. None None 26mm. |X 990 2574 mm. eee ase

8mm. |Proj. x2} None 0.6mm. |X 2000 1200 mm. |Narrow tube

8mm. |Proj. x4| None 0.6mm. |X 4000 2400 mm., 1) fe

8mm. | None |— 5 d. 2.6mm. |X 1485 | 3861mm. |Tube as in

oe pl. v, B

8 mm. None |—10 d. 2.6mm. |X 2000 5200 mm. ibe os

2mm. | None | None | 0.5375mm.|X 4000 | 2150mm. «4

Hom. Im.

SCREEN

From the earliest use of the magic lantern it was recognized

that a white screen made of cloth or a whitewashed wall was

desirable (see quotations from Porta). Translucent screens were

also used. These were made of ground glass or of oiled cloth.

Then the lantern could be entirely out of sight of the audience.

This was of course necessary for the striking exhibitions of the

phantasmagoric magic lantern. Even at the present day ground

glass screens are sometimes used for lecture purposes. In such

cases the apparatus is eliminated from the lecture-room.

Baker (1742-1785) recommends a screen made of “the largest

elephant paper” stretched on a frame like that used for fire screens.

Adams (1746-1771) recommends the same kind of screen. Both

recommend for a large screen, one made by pasting several sheets

of the elephant paper on cloth. The screen is then let down from

the ceiling like a roller map.

THE PROJECTION MICROSCOPE 39

Pritchard, 1837, gives in a few words the requisites of a good

screen for projection purposes: “It should be smooth, white and

opaque; it should reflect the greatest quantity of light and absorb

the least.” He advocates among other things screens made of

plaster of Paris or a wall with a thin coat of plaster of Paris spread

upon it. Lewis Wright also speaks well of plaster of Paris screens.

From my own experience a smooth wall with a wash of plaster

of Paris forms the best screen for micro-projection. Such a screen

fulfills the requirements given for a screen by Pritchard. For the

highest powers a small screen made by flowing a thin layer of

plaster of Paris on a perfectly clean plate glass gives the most per-

fect results. In making the screen a frame with wire netting is

placed on the glass and the plaster of Paris poured on until the

wire is covered and the frame filled with the plaster. After the

plaster has set and partly dried there is no difficulty in removing it

from the glass.

Large bristol board sheets with a smooth, lusterless white sur-

face also serve well. The coated roller screens now on the market

are good for general work, but the surface is too rough for the

most satisfactory micro-projection.

Screen Distance.—With lamp light or even with the lime light

the screen distance cannot be very great or the projected images

will be too faint. With sunlight and the electric light, the screen

distance is almost unrestricted. For ordinary projection with a

current of 12 to 15 amperes and objectives up to 6 mm. equivalent

focus, a screen distance of 7 to 10 meters gives good effects. Every-

thing that can be shown well can be made large enough so that each

person in an audience of 200 to 500 can see with satisfaction.

For the higher powers (3 mm., 2144 mm., 2 mm. objectives), a

screen distance of 3 to 5 meters is more satisfactory, and the spec-

tators must be few and close to the screen or they must use opera

glasses. For objects requiring high powers, 2%4 or 2 mm. objec-

tives, it is far more satisfactory to use compound microscopes in

the ordinary way.! The writer has not found projection of details

"For example the oval red blood corpuscles of the camel were 25 mm

long with a 2 mm. homogeneous immersion objective and a screen distance

of 3 meters; but the screen picture was far less satisfactory than the image

of the same corpuscles seen under the microscope.

40 SIMON HENRY GAGE

satisfactory which could not be seen with a compound microscope

supplied with a low ocular and a 16 mm. objective. For such

objects the projection microscope shows them on the screen as satis-

factorily for 500 as the compound microscope shows them to a

single individual.

DARKENING THE ROOM FOR PROJECTION

From the earliest experiments with projection (see note from

Porta), the effect was known to depend largely on the darkness of

the room where the projection took place. It was for this reason,

in part, that the evening was so often selected for lantern exhibi-

tions. Baker (1742-1785) says: ‘When this microscope is em-

ployed the room must be rendered as dark as possible, for on the

darkness of the room and the brightness of the sunshine [or other

light] depend the sharpness and perfection of your image.” Pritch-

ard goes further and would have everything in the projecting

room dark except the screen. This is often done in rooms for draw-

ing with the projection microscope.

With a magic lantern for lantern slides and the powerful modern

electric light the room does not need to be very dark, but for micro-

projection unless it is very dark the screen image will have a gray

appearance and lack in crispness.

MECHANICAL PARTS

With every increase in complexity of the projection microscope

and its work, the mechanical parts must keep pace. Naturally

the earliest forms were very simple (fig. 6; pls. 1 and 11).

The construction and arrangement to meet the increasing require-

ments have been worked out very differently by different ones. The

diagrams (figs. 5, 7, 8, 10, 14 and 17) represent fairly well the

composition and arrangement of the different optical and mechanical

parts in different cases. There is, it seems to me, one fundamental

requisite, viz., that each part should be independent and so mounted

that it may be easily adjusted to work most effectively with the

other parts. That is the radiant, the condenser, the water-bath,

the stage with its water-bath or condenser, and finally the projection

THE PROJECTION MICROSCOPE 41

objective with the focusing arrangement should all be separate and

separately adjustable. They should also be on some kind of lathe

bed or guide so that they may all be moved back and forth in a

longitudinal direction without getting out of line. In a very simple

form (fig. 6), where no great variety of work is demanded all the

parts may be more closely united, but it is very restrictive to have

them so united in modern apparatus. For compactness it is of

great advantage to have the water-bath on the same support as the

condenser lenses, but as shown by the table above, the water-bath

is greatly heated by conduction from the lamp as well as by the

absorption of radiant heat. The apparatus may be more compact

with the stage and tube and focusing device in one piece as with

an ordinary compound microscope, but unless the coarse adjust-

ment has a very long rack for focusing it is impossible to arrange

the object and the objective in the position giving the best results

in the projected image.

In the earlier editions (4th, 1899) of Zeiss’ catalog of projec-

tion apparatus, there is one, the so-called simplified projection

apparatus which fulfills the above requirements, because each piece

is independent and all are arranged on a kind of lathe bed, and one

may easily pass from high power projection to that of objects

25 to 30 mm. in diameter. The one presented in pl. v, A, B,

closely approximates the best construction. The contact of the

condenser water-bath with the metal part of the condenser holder

is objectionable as the water-bath is heated by direct conduction

(see table above).

One of the greatest defects is the small size of the opening in the

stage. Of course no specimen larger than the stage opening can

be projected. This prohibits the use of the large sections of the

central nervous system necessary for understanding its morphology ;

it also prohibits the use of sections of organs and of large embryos,

t. e., objects of 25 to 60 mm.

In early projection microscopes there was no tube extending

beyond the projection objective (figs. 6, 8 and pl. 1v, A), therefore

there was no restriction of the field from that cause, as with the

relatively long and narrow tube of the ordinary compound micro-

scope so commonly used in projection at the present day.

42 SIMON HENRY GAGE

To avoid this restriction of the field by the tube, either no tube

is used with photographic and other low objectives (pl. v, a), or for

ordinary projection a very large and short tube (pl. v, B) is used,

#. e., one 45 to 50 mm. in diameter and 9 to 10 centimeters long.

Harting, in 1839, figured and described such an arrangement.

Selenka, 1887, used no tube, and Zeiss and others recommend large

and short tubes like that of Harting. If projection oculars are to be

used a small tube with an adapter may be attached to this large tube.

BLACKENING THE APPARATUS

As recommended by Goring and Pritchard in the Micrographia

(1837), all the metal and exterior parts of the projection micro-

scope should have a dead-black finish. Sunlight, the lime light,

and the electric light are so brilliant that any polished surface

reflects the light so strongly that the operator is almost blinded

and becomes unable to focus the image on the screen properly.

Much light is also reflected into the room.

DRAWING WITH THE PROJECTION MICROSCOPE

Baker (1742-1785) describes with enthusiasm the advantages of

the projection microscope for drawing’; Adams, father and son

(1746-1787), describe the ease by which drawings can be made on

vertical surfaces, including ground glass. By means of a special

*BAKER, Henry P., Of Microscopes and the discoveries made thereby,

Disco:

“This Microscope [the solar microscope] is the most entertaining

of any; and, perhaps, the most capable of making Discoveries, in Objects

that are not too opake: as it shews them much larger than can be done

any other Way. There are also several Conveniences attending it, which

no other Microscope can have; for the weakest eyes may use it without the

least Straining or Fatigue: Numbers of People may view any Object to-

gether at the same Time, and, by pointing to the particular Parts thereof,

and discoursing on what lies before them, may be able better to under-

stand one another, and more likely to find out the Truth, than when, in

other Microscopes they must peep one after another, and perhaps see the

Object neither in the same light, nor the same Position. Such too as have

no Skill in Drawing, may, by this contrivance, easily sketch out the exact

Figure of an Object, they have a mind to preserve a Picture of; since they

need only fasten a Paper upon the Screen, and trace it out thereon, either

with a Pen or Pencil, as it appears before them.”

THE PROJECTION MICROSCOPE 43

camera obscura arrangement which can be set in a vertical position,

the object being illuminated by sunlight, drawings can be made

in a horizontal position. George Adams, Jr., devised an arrange-

ment for drawing with the lamp or lucernal microscope also. In

1767, Brander devised a camera obscura for drawing with the

solar microscope; the illuminating and projecting part being hori-

zontal, a plane mirror reflected the image downward upon a hori-

zontal surface where it could be drawn.

In the Micrographia of Goring and Pritchard, ground glass is

used for a screen and drawings are made on paper in a vertical

position, either on the front or the back of the screen. Working on

the back of the screen has the advantage that the hands and pencil

cast no shadows.

Dr. Goring describes an accessory to his solar microscope by

which the light is reflected downward by means of a plane mirror

or prism into a drawing box. At the bottom of the drawing box

is a field curved to correspond with the curvature of the field of

the microscope (see figures in the Micrographia and in Brewster On

the Microscope, 1837).

In Chevalier’s work (1839), the advantages of the projection

microscope for drawing are mentioned, and he figures a prism intro-

. 6

4 ‘

=e eel

Lente A ae

TAR L a NM

ae ae SEAS

sb es

ee A —

—— ee at

H “S

Fig. 18. Projection microscope from Chevalier (Planche 2). M, mirror

reflecting the sun’s rays (RR’, rr’) to the condenser (C) ; from the con-

denser they pass to the substage condenser (c) and are condensed upon the

object (0). L, achromatic objective; A, amplifier composed of a plano-

convex and a double concave lens,—this amplifier makes the rays much

more divergent, #. e., BB’ instead of bb’; P, right-angled prism acting as a

45-degree mirror to project the image down upon a horizontal surface for

drawing.

44 SIMON HENRY GAGE

duced in the path of the rays to reflect them downward on a hori-

zontal surface for drawing (fig. 18).

At the present time the projection microscope is much used for

the numerous drawings necessary for making models. Sometimes

the drawings are made on a vertical surface but some laboratories

have also a device for reflecting the image downward upon a hori-

zontal drawing surface. It is tiresome to draw for a considerable

length of time on a vertical surface, but on a horizontal surface

the drawing hand is supported and it is easier to get an exact copy.

For making large diagrams of microscopical specimens one naturally

uses a vertical surface.

C Sw O

= |

; OR \

Det ie

i a fide tL

Z ee daa ng ai a nde ae Be pad

Fig. 19. Projection microscope for drawing on a horizontal surface.

The optical bench carrying the radiant (R), the condenser and condenser

water-bath (CW), the stage and stage water-bath (Su) and the micro-

scope (O), are on one table, Lt, and are perfectly horizontal. The drawing

table, Dt, bears the mirror (M) at an angle of 45 degrees. This insures an

undistorted image on the horizontal surface of the drawing table. This

apparatus, and the furniture for photography are in a special cabinet about

3 meters square atid 2%4 meters high, in a large room. It is painted dead

black on the interior. V7, Light-excluding ventilator.

THE PROJECTION MICROSCOPE 45

In some institutions special rooms are set apart for drawing and

for photography, for example in the Johns Hopkins medical school,

the Wistar Institute for Anatomy and Biology, etc. Such rooms

are usually made dead black to avoid reflections. Crisper pictures

are also easier to produce under such conditions. In fig. 19 is

shown a small drawing room (3x3x2% meters) or cabinet within a

large laboratory. See also the Anatomical Record, November, 1907,

where Dr. Greenman describes the drawing or photographic cabinet

of the Wistar Institute. Either a special room convenient to the

laboratory or some such cabinet in the laboratory is almost a neces-

sity for the work of a modern biologic department.

PROJECTION MICROSCOPE FOR OPAQUE OBJECTS

In the study of modern histology and embryology the majority of

the objects to be studied are lighted by transmitted light and are

therefore seen as transparencies. In the earlier days before the

technique of microscopic work had become developed to any great

extent a large number of the objects studied were viewed as opaque

objects, and all sorts of devices (bull’s eye condensers, concave

specula, etc.) were used to illuminate the objects. Naturally there

was a great desire to show opaque as well as transparent objects

by the projection microscope, and the Adamses (1746-1787) devised

what they called opaque solar and lucernal microscopes. Martin

(1774) is said to have produced a fairly good one. Sometime

before 1756, Lieberkiithn, according to Harting, devised a solar

microscope for opaque objects, and Euler and his followers, Aepinus

and Zeiher (1750-1779), suggested improvements for thts purpose.

Harting also speaks of his countryman, Hendrik Hen (1807), as

having made an excellent microscope which was adapted for opaque

objects and also for use as a solar microscope. In the Micrographia

of Goring and Pritchard (1837) there are given very good figures

and descriptions of a projection microscope for opaque objects

adapted either for sunlight or the lime light.

Briefly stated an opaque projection microscope is one in which

light from the radiant (sun or some form of lamp), instead of

passing through the object and directly to the objective is received

by a plane mirror, or a series of plane mirrors arranged in a circle,

46 SIMON HENRY GAGE

or by a concave mirror and thrown upon the upper side of the

opaque object. The light reflected from the surface of the object

passes to the objective and the image is by it projected upon a screen

as usual.

In the form devised by Adams there was a plane mirror in

line of the beam of light from the radiant. The object and objective

were at a higher level. The light from the plane mirror was

reflected upon the object and illuminated its upper surface. More

often the objective and object were in the line of the illuminating

beam, but a series of small plane mirrors were arranged around the

objective to reflect the light back upon the upper side of the opaque

object, or a concave mirror with a hole in the middle for the objective

was used for the same purpose. The perforated concave mirror is

said by Harting to be due to the ingenuity of Leeuwenhoek, but its

invention is often ascribed to Lieberktihn, and goes by his name.

Opaque projection is now little used with the microscope except

for photographing metals and alloys in micro-metallography. The

fundamental principles of Adams’ apparatus have been closely fol-

lowed, however, in the modern magic lantern devices known as

episcopes or reflectoscopes for projecting pictures in books, various

forms of mechanism, etc.

Proj ECTION AND PHOTOMICROGRAPHY

One of the most important uses of projection is the production

of photographs of microscopic objects. The general principles for

obtaining the real images necessary in photography and for screen

images are the same.

Among those who raised photography with the microscope to a

high degree of perfection (1866-1876) should be mentioned, in

the first place, Col. J. J. Woodward of the United States Army

medical corps. His papers on the subject and the accompanying

photographs form a notable landmark. In the period when the

projection microscope for teaching and public exhibitions had

receded into the background he and others used projection for

photography and worked out with great precision the conditions of

lighting, objectives, etc., upon which success depends. While he

employed oculars and amplifiers for some of his work, he says in

THE PROJECTION MICROSCOPE 4?

his report (1871) on photographing histologic preparations by sun-

light, that he discarded all oculars and amplifiers and got the desired

magnification by the objective at the necessary distance. His photo-

graphic laboratory was a large room and the sensitive plate could

be placed at any desired distance from the objective. Dr. Woodward

speaks very highly of the electric light and compares it favorably

with sunlight. While the interest in photo-micrography helped to

keep alive the knowledge of projection, its influence on projection

for public exhibitions and class-room work has not been altogether

helpful, for while the general principles are the same, the best effects

in photography are only obtained when sharpness with fine detail

are secured, while with projection for teaching and exhibition pur-

poses, brilliance and striking contrast are all important. The fine

detail so necessary in a photograph are lost on a distant screen

image.

Furthermore, in photography only a very limited field can be

shown in a single picture, while in projection for teaching purposes

a very large field should be available so that the relations of the

various parts of the objects can be seen. It seems therefore to the

writer that in striving to realize all of the excellence of a photo-

graphic image for ordinary projection work there has been a retard-

ing effect upon projection, by a too close adherence to the condi-

tions necessary in photography. The substage condenser so de-

sirable in photography limits the size of the stage opening and con-

sequently the size of the object which can be shown. It also pro-

hibits the use of a really effective stage water-bath, and consequently

makes it impossible to use delicate and living objects for a sufficiently

long period. In photomicrography high powers give satisfactory

results, but for projection on a screen for many observers only

moderate powers are really satisfactory (100 mm. to 5 mm. ob-

jectives).

CoNCLUSION

In tracing briefly the course of events since the first production

of lenses it is seen that the enthusiasm aroused by the use of these

glasses was great and enduring. To the visible forms with which

nature is so abundantly endowed they revealed an abundance of

invisible ones, rivalling the visible both in numbers and in beauty.

48 SIMON HENRY GAGE

With the deep social instinct that makes men wish to share with

their fellows, there came about the development of means to show

these marvels of beauty and complexity to many at once, hence has

been more fully perfected the Jaterna magica in its various forms

during the last three hundred years.

In our own generation the interest in minute forms with beautiful

markings and strange ways do not arouse the same enthusiasm as

with the first observers, yet, after the introduction of the doctrine

of organic evolution, they were studied with an intensity never

before known in the hope that nature might perhaps in these simple

forms reveal herself. Furthermore there is a widespread conviction

that some personal and intimate knowledge of nature should be the

heritage of every human being. Hence has come in the period of

“Nature Study,” and the immense improvement of laboratory archi-

tecture, and the betterment of laboratory facilities. In this advance

the improvement and use of the microscope has not lagged behind.

But one can hardly assert the same of the projection microscope.

The simple and general pictures given by it in earlier times no

longer satisfy. Now more frequently it is the definite and specific,

and the minutest detail that contain the points of greatest interest.

As every individual student must repeat the history of the race in

his development the wise teacher does not too rigidly insist on the

minute and the specific at first, but deals with the general, and the

broad relations of things.

This being true there is still a place in which the projection

microscope can supply a need in instruction. This need is some

method of demonstrating the forms and relations of parts in the

specimens themselves. It is the need of getting back to nature in

lectures and demonstrations instead of relying too exclusively on

diagrams and models.

As nearly every institution now has an electric lighting plant and

the ordinary magic lantern for lantern slides is almost universally

used the addition of a projection microscope to the teacher’s outfit is

not prohibitive.

Every person installing such an outfit should know what can be

reasonably expected of it, and the range of its possibilities. The

modern teacher of biology, judging from my own experience, re-

THE PROJECTION MICROSCOPE 49

quires a projection microscope which will begin where the ordinary

magic lantern leaves off, and show specimens from 50 to 60 mm.

in diameter to those of 1 mm. or less, and so enlarge them that they

in all their details can be seen by 500 people as easily as by one

looking into an ordinary microscope.

The aid which the projection microscope is capable of rendering

the investigator in the preparation of drawings and in the study

of the relations of structures is so great that when once fully aware

of this possible help he will not willingly forego it.

Not many projection microscopes meet all the requirements

given above, but that, in part at least, is because maker and user

have become too much separated by modern specialization. In

earlier times the user of optical instruments was also frequently the

maker (Galileo, Newton, Campani, Herschel, Huygens, Hooke,

Leeuwenhoek, etc.), or the optician was the friend and co-worker of

the master or patron for whom the apparatus was made.

With the constant or direct electric current guaranteeing a radiant

at all hours of the day, which is but little inferior to sunlight,

biologists and other users of the microscope are turning again to the

projection microscope and the greatest optical manufacturers in the

world are taking hold of the problem in good earnest. Fortunately

also many men with laboratory training and who therefore know

the actual needs are coming more and more to have positions of

responsibility in these great manufacturing establishments.

As with the laboratories of physics, chemistry and physiology, so

in a less degree of laboratories of biology, much of the apparatus

needed to work out special problems is first produced in the labora-

tory workshop under the direction of the one who knows best the

object to be attained. Enough has been accomplished already by the

laboratories and the opticians in bringing the apparatus abreast of

modern requirements to show that there is a movement on foot to

utilize more and more this most striking of all means of study and

demonstration, and it is confidently predicted that the projection

microscope will soon come to its own again.

50 SIMON HENRY GAGE

ANNOTATED BIBLIOGRAPHY

For the better appreciation of the appearance and character of the older

works consulted the entire title page is given, including the various mottoes

or proverbs appended thereto.

ADAMS, GEORGE.

Micrographia illustrata: or the microscope explained in several new

inventions, particularly of a new variable microscope for examining all

sorts of minute objects and also of a new Camera Obscura Microscope de-

signed for drawing all minute objects either by the light of the sun or by

a lamp in winter evenings to great perfection; with a description of all

the other microscopes now in use. Likewise a natural history of aerial,

terrestrial and aquatic animals, &c., considered as microscopic objects. By

George Adams, mathematical instrument maker to His Majesty. London,

1746, 1771.

The 4th edition of 1771 was used. It is illustrated by 72 copper plates

containing 560 delineations of various microscopic objects. The title page

of this work gives a very good table of contents.

ADAMS, GEORGE.

Essays on the microscope containing a practical description of the most

improved microscopes, a general history of insects, their transformations,

peculiar habits and ceconomy; an account of the various species and singular

properties of the hydrae and vorticellae; a description of 379 animalcula

with a concise catalogue of interesting objects; a view of the organization

of timber and the configuration of salts when under the microscope. Thirty

double copper plates of objects and apparatus. By George Adams, mathe-

matical instrument maker to His Majesty, and optician to his royal high-

ness the Prince of Wales. 4to, London, 1787.

Son of the preceding. Gives a very good account of the projection

microscope, and like the other English writers, ascribes its invention to

Lieberktihn. Jn the preface, p. x, he says: “When I first undertook the

present essays I had confined myself to a re-publication of my father’s

work, entitled, Micrographia Illustrata’ (see above). At the end of the

preface is appended a list of 50 titles of authors consulted in the prepara-

tion of his essays. It is like a latter-day bibliography.

AMERICAN MICROSCOPICAL SOCIETY.

Proceedings and Transactions: 1878 to date. Published by the Secretary.

As the secretary changes frequently there is no fixed place of publication.

AmeErRICAN Montriy Microscoricat JournaL, THe. Iilustrated. Washing-

ton, D. C.: 1880—1902.

THE PROJECTION MICROSCOPE 51

BAKER, HENRY.

Of Microscopes and the discoveries made thereby. Illustrated with

many copper-plates. By Henry Baker, fellow of the Royal Society, and

member of the Society of Antiquaries in London. In two volumes. Vol. 1—

The microscope made easy. Vol. 1—Employment for the microscope.

Vol. 1, A new edition, “Rerum Natura nusquam magis quam in Minimis

tota est.” Plin. Nat. Hist. Lib. xic. 2. London, 1785.

According to the Index Catalog of the Library of the Surgeon General’s

Office, N. S., the first edition was produced in 1742, the second in 1743,

and the “new edition” in 1785. The 1785 edition has been consulted in the

preparation of this address. Baker’s work was translated into French and

into German, and it was well worthy of this distinction. It is probably the

mistake of Baker in attributing the discovery of the projection microscope

to Lieberkiihn that gave the error such wide circulation through the trans-

lations of his work. The feeling of uncertainty concerning the history of

the microscope in the mind of Baker may be inferred by this extract from

his introduction: “Of Microscopes in General: To what accident, to what

Country or to whom, we are obliged for the Invention of Microscopes, is

not in me to determine.” (Pp. 2-3.)

The dedication of this work, 1742, breathes so earnestly and with so

much enthusiasm and good judgment the spirit of the true naturalist, that

it is in large part here reproduced. The discovery of the importance of the

bacteria and other micro-organisms since this dedication was written, now

over 150 years ago, gives special point to what he says comparing the im-

portance of the microscopic forms and those of large size—the “capitals”

and the “little letters,’ as he calls them.

At a meeting of the Royal Society, October 28, 1742. Dedication: “To

Martin Folkes, Esq., President, and to the Council and Fellows of the

Royal Society of London. Gentlemen: An Attempt to excite in Mankind

a general Desire of searching into the Wonders of NATURE, will, I

persuade myself, be accepted favorably by you, whose Endeavours for’ the

Advancement of Natural Knowledge, according to the Purpose of your

Institution, are esteemed by all the World.

“It is something more than an hundred and twenty Years since the

MICROSCOPE was happily invented; and to the valuable Discoveries

made thereby, we stand indebted, as the following Sheets will shew, for a

great Part of our present Philosophy. In such a Length of Time, it is

however probable many more Advantages might have been reaped from

it, had not some Difficulties and Discouragements prevented its general

Use.

“At the Beginning it was confined to very few; who, making a Secret

of it, endeavoured all they could to keep it to themselves; and, when it

became a little more publick, the Price was fixed so high, that the most

Curious and Industrious, who have not always the greatest Share of

52 SIMON HENRY GAGE

Money, could not conveniently get at it. Of late Years, indeed, the Ex-

pense has been much less; but then new Discouragements have started up

from Mistake and Prejudice.

“For many have been frighted from the Use of it, by imagining it

required great Skill in Optics, and Abundance of other Learning, to com-

prehend it to any purpose; whereas nothing is really needful but good

Glasses, good Eyes, a little Practice, and a common Understanding, to

distinguish what is seen; and a Love of Truth, to give a faithful Account

thereof. Others have considered it as a mere Play-thing, a Matter of

Amusement and Fancy only, that raises our Wonder for a Moment, but is

of no farther Service; which Mistake they have fallen into, from being

unacquainted with any Principles whereby to form a right Judgment of

what they see. Many, again, have laid the Microscope aside, after a little

Use, for want of knowing what Objects to examine, where to find, how

to prepare, and in what Manner to apply them. The Trouble of managing

it has also frighted some.

“But we are now so fortunate as to have this Instrument greatly im-

proved amongst ourselves, the Apparatus made much easier as well as

more useful, and the Price considerably reduced. The Solar or Camera

Obscura Microscope, and the Microscope for viewing Objects that have no

Transparency, by throwing a strong reflected light upon them, are also

new, Inventions, from whence great Things may be expected.

“Nothing therefore is now wanting, but a general Inclination to employ

these Instruments, for a farther Discovery of the Minute Wonders of the

Creation; which may not, perhaps, improve our Knowledge less than the

grander Parts thereof. Bears, Tigers, Lions, Crocodiles, and Whales, Oaks,

and Cedars, Seas and Mountains, Comets, Stars, Worlds and Suns, are

the CAPITALS in Nature’s mighty Volume and of them we should not be

ignorant; but whoever would read there with Understanding, must make

himself Master of the little letters likewise, which occur a thousand Times

more frequently, and, if he does not know them, will stop him short at

every Syllable.

“The likeliest Method of discovering Truth is, by the Experiments of

Many upon the same Subject; and the most probable Way of engaging

People in such Experiments, is, by rendering them easy, intelligible, and

pleasant. To effect this, is my Endeavour in the following Treatise.”

BREWSTER, SIR DAVID.

A treatise on the microscope, forming the article under that head in

the seventh edition of the Encyclopaedia Britannica, by Sir David Brewster,

vice president of the royal society of Edinburgh, and corresponding member

of the Institute of France. Edinburgh, 1837.

There is a very good account of the projection microscope with sunlight

or oxyhydrogen light for illumination.

THE PROJECTION MICROSCOPE 53

CARPENTER, WILLIAM B.

The Microscope and its revelations.

This is probably the greatest work on the microscope in the English lan-

guage. Its various editions from the first, in 1856, to the last (8th, 1901),

edited by the Rev. W. H. Dallinger, have been a storehouse of knowledge

and the best that is known in the microscopical world. It does not deal

to any great extent with the projection microscope, but is admirable in the

history of the microscope itself.

CHEVALIER, CHARLES.

Des Microscopes et de leur usage. Description d’appareils et de pro-

cédés nouveaux, suivie d’expériences microscopiques puisées dans les meil-

leurs ouvrages anciens et les notes de M. Le Baillif, et d’un mémoire sur

les diatomées etc. par M. de Brébisson. Paris, 1839. C. Chevalier was an

“Ingénieur-Opticien, membre de la Société d’encouragement pour 1|’industrie

nationale; l’un des lauréats (médaille d’or) a l’exposition de 1834, etc.”

There is a good discussion of the history of the microscope in this

work. Chevalier follows the English in ascribing the discovery of the pro-

jection miscroscope to Lieberkiihn. Chevalier’s figure of the projection micro-

scope (fig. 18 of the text, above) indicates how advanced were the instru-

ments constructed by him.

CoLE, AARON HopGMAN.

Manual of biological projection and anesthesia of animals. Pp. 200,

Chicago, IIll., (1907). Illustrated.

This work is founded on the series of articles on the subject originally

published in the Journal of Applied Microscopy, and contains many good

hints.

DECHALES.

(R. P. Claudii Francisci Milliet Dechales Camberiensis e’ Societate

Jesu.) Cursus, seu Mundus Mathematicus. Tomus Tertius. Complectens

Architecturam Militarem, Hydrostaticam, Tractatus de Fontibus & Fluviis,

de Machinis Hydraulicis, & de Navigatione, Opticam Perspectivam, Catop-

tricam, & Dioptricam. Editio altera ex Manuscriptis Authoris aucta &

emendata, opera & studio R. P. Amati Varcin ejusdem Societatis. Annison

che fiorisce. Lugduni, Apud Anissonios, Joan. Posuel & Claud. Rigaud.

M. DC. LXXXX. Cum Privilegio Regis.

The part relating to thé projection microscope-is contained in: “Trac-

tatus XXIII. Dioptrica, seu de Radio Refracto.” Liber 1, Propositio Lvm.

Theorema (p. 680). “Exigui prototypi, uniea lente convexa amplificatam

imaginem, in pariete exhibere.” Also, Liber 11, Propositio xx, Problema (p.

696). “De nocte exigui prototypi ingentem in muro imaginem distinctam

exhibere daubus lentibus.” See the figure (fig. 7) and quotation above, p. 14.

There are four folio volumes of this work. Each volume contains many

figures. The first edition appeared in 1674.

54 SIMON HENRY GAGE

GAGE, SIMON HENRY.

The Microscope. An introduction to microscopic methods and to his-

tology. Tenth ed., 1908. First edition, 1881. Ithaca, New York.

GorRING, C. R., AND PRITCHARD, ANDREW.

Micrographia, containing practical essays on reflecting, solar, oxy-hydro-

gen gas microscopes; micrometers, eye-pieces, &c, &c. London, 1837.

This is one of the most satisfactory of the older works, and deals in a

modern fashion with the subjects named in the title page. On pp. 170, 171,

Pritchard gives a very detailed account of the early use of the oxy-hydrogen

light for use with a lantern by Birkbeck in lectures at the London Me-

chanics’ Institute, 1824.

HArTING, PIETER.

Das Mikroskop, Theorie, Gebrauch, Geschichte und gegenwartiger

Zustand derselben, von P. Harting, Professor in Utrecht. Deutsche Origi-

nalausgabe vom Verfasser revidirt und vervollstandigt. Herausgebeben von

Dr. Fr. Wilh. Theile, grossherzoglich Sachsischem Medicinalrathe, in drei

Banden. Zweite wesentlich verbesserte und vermehrte Auflage. Mit 466

in den Text eingedruckten Holzstichen und einer Tafel in Farbendruck.

Braunschweig, 1866.

First German edition, 1858. Original Holland edition: Het mikroskoop,

deszelfs gebruik, geschiedenis en tegenwoordigen toestand. Een handboek

voor natur—en genees-kunde. Utrecht, 1848, 1849. Gr. 8°, 3 din, met 5 pl.

Dr. Harting (born 1812, died 1885) was for many years professor in

Utrecht university, teaching, among other things, the microscopic structure

of plants and animals, etc. In his later years he went over to zoology and

comparative anatomy. He was a good type of the scholarly men Holland

has been producing for so many generations. This book on the microscope

is one of the best that has ever been produced, and one refers to it over

and over again with renewed pleasure. In discussing the controverted

points of history in the development of the microscope his step is almost

always sure in the slippery maze of conflicting testimony. His research in

this field was profound and his judgment sound. Considering the state-

ments of Poggendorff and a study of original sources leads me to think

that Dr. Harting was mistaken in ascribing the invention of the projection

microscope to Kircher. (See above, in the address, and Harting, Bd. mm,

p. 279 et seq.)

JourNAL oF APPLIED Microscopy AND LABORATORY MeEtHonps. Rochester,

N. Y., 1898—1903.

This contains a series of articles by A. H. Cole on projection.

JoURNAL OF THE RoyAL Mrcroscoricat Society. Illustrated. London, 1878

to date.

Bibliography of works and papers relating to the microscope, micro-

scopical methods, and histology. It also includes a summary of many of

the papers.

THE PROJECTION MICROSCOPE 55

KircHER, ATHANASIUS.

(Athanasii Kircheri Fuldensis Buchonii € Soc. Jesu Presbyteri). Ars

Magna Lucis et Umbrae. In X. Libros digesta. Quibus Admirandae Lucis

& Umbrae in mundo, atque adeo universa natura, vires effectusque uti nova,

ita varia novorum reconditiorumque speciminum exhibitione, ad varios mor-

talium usus, panduntur. Editio altera priori multo auctior. Sicuti tenebrae

ejus ita & lumen ejus, Psal, 138. Amstelodami, Apud Joannem Janssonium

a Waesberge, & Haeredes Elizaei, Weyerstraet. Anno clo Ioo LxxI, (1671)

Cum Privilegio Sacr. Caesar. Majestatis, & Ord. Holl & Westfr.

The first edition of this work was published in 1646. According to

Poggendorff, p. 436: “Kircher spricht von der Laterna magica in seiner

Ars magna lucis et umbrae, aber nicht in der ersten Auflage von 1646,

sondern in der spatern von 1671.” In the edition of 1671, the projection

microscope is considered in Liber x, p. 768, Problema iv (see pl. 1, II,

below). Also in Cryptologia Nova, pars prima; De Projectione Figurarum

inquamlibet distantiam per Solem, p. 792 (see pl. 1).

This is a wonderful folio volume of over 800 pages, ranging in interest

from the “experimentum mirabile, de imaginatione Gallinae,” p. 112, where,

to illustrate the point, is a hypnotized chicken on a chalk line, to the burning

of the investing Roman fleet at Syracuse by Archimedes with a “Speculum

causticum,” with, on p. 764, a picture of the scene; and finally to the

height of all knowledge in the last chapter, p. 795, “Epilogus sive Meta-

physica Lucis et Umbrae.” No part of knowledge, real or imaginary, was

neglected by the old encyclopedists.

Letss, C.

Die optischen Instrumente der Firma R- Fuess, deren Beschriebung,

Justierung und Anwendung; Mit 233 Holzschnitten im Text und 3 Licht-

drucktafeln. Leipzig, 1899.

This work deals fully with optical projection, including micro-projection.

All sources of illumination are discussed and a very excellent account is

given of the different forms of heliostat.

MAYALL, JoHN, Jr.

Cantor Lectures on the [History of the] Microscope delivered before

the Society for the Encouragement of Arts, Manufactures and Commerce,

Nov., Dec., 1885. London, 1886. This monograph of 97 pages is a model

of clearness, accuracy, and fairness. It is abundantly illustrated. On p. 42

he ascribes the invention of the projection microscope to Lieberkiihn. This

is difficult to understand, as the author is so accurate in most matters.

He gives two figures of the solar microscopes of Adams and Cuff, 1744-

1746, but does not deal adequately with the projection microscope.

Petri, R. J.

Das Mikroshop von seinen Anfangen bis zur jetzigen Vervollkommnung

fiir all Freunde dieses Instruments, von. Regierungsrath Dr. med. R. J.

56 SIMON HENRY GAGE

Petri, ordentl. Mitglied des kaiserlichen Gesundheitsamtes und Vorstand

des bakteriologischen Laboratoriums daselbst. Mit 191 Abbildungen im

Text und 2 Facsimiledrucken. Berlin, 1896.

“Niemals zurtick” is the proverb on the title page. While the projec-

tion microscope is not dealt with in this book the development of the micro-

scope is well done in text and figures. At the end there is a bibliography

of 73 different works or editions, beginning with the Optice thesaurus

Alhazeni (1535, 1572, Latin translations), and ending with the work of

Dr. Henri van Heurck, 1891.

PHILOSOPHICAL TRANSACTIONS OF THE RoyAL Society oF Lonpon. IIlus-

trated. London, 1665 to date.

PoGGENDoRFF, J. C.

Geschichte der Physik Vorlesungen gehalten an der Universitat zu

Berlin. Mit vierzig holzschnitten. Leipzig, 1879.

On the title page is also this motto: “Ex umbra in solem.” The his-

tory given by Poggendorff is divided into three periods: (1) The ancient

or Greek period extending to the fall of Alexandria (460 A. D.); (2) The

Arabic and Middle Ages (460 to the end of the 16th century, including

therefore Kopernicus and Keppler); (3) The period of Galileo and Newton,

17th and part of 18th centuries—about 150 years.

Porta, GIOVANNI BATTISTA DELLA.

Magiae Naturalis, sive de miraculis rerum naturalium. First edition 1553-

4; 2d, 35 years later, 1589.

Natural Magick by John Baptista Porta, a Neapolitane. In twenty

books : e

1. Of the Causes of Wonderful 11. Of Perfuming.

things. 12. Of Artificial Fires.

2. Of the Generation of Animals. 13. Of Tempering Steel.

3. Of the Production of new Plants. 14. Of Cookery.

4. Of Increasing Household-Stuff. 15. Of Fishing, Fowling, Hunting,

. Of changing Metals. &e.

. Of counterfeiting Gold. 16. Of Invisible Writing.

. Of the Wonders of the Load- 17. Of Strange Glasses.

stone. 18. Of Statick Experiments.

8. Of strange Cures. 19. Of Pneumatick Experiments.

9. Of Beautifying Women. 20. Of the Chaos.

10. Of Distillation.

Wherein are set forth all the Riches and Delights of the Natural

Sciences. London. Printed for Thomas Young, and Samuel Speed; and

are to be sold at the three Pigeons, and at the Angel in St. Paul’s Church-

yard. 1658.

This English edition is from the second Latin edition of 1588-9. One

can but admire the zeal and industry shown by the old writers who were

THE PROJECTION MICROSCOPE 57

“learned in all the knowledge of their time.’ A couple of paragraphs from

the preface will give the point of view more clearly than anything else.

The capitalization and punctuation of the original is preserved so that the

reader may compare the appearance of a printed page in 1658 with one at

the present day:

“If this Work made by me in my Youth, when I was hardly fifteen years

old, was fo generally received and with fo great applau fe, that it was forth-

with tran {lated into many Languages, as Italian, French, Spani{h, Arabick;

and pa{ {ed through the hands of incomparable men: I hope that now com-

ing forth from me that am fifty years old, it 5 hall be more dearly entertained.

For when I jaw the fir {t fruits of my Labours received with fo great Alacrity

of mind, I was moved by the{e good Omens; And therefore have adventured

to § end it once more forth, but with an Equipage more Rich and Noble.

“From the fir {t time it appeared, it is now thirty-five years, And (without

any derogation from my Mode|ty be it i poken), if ever any man labored

earne {tly to disclo{e the fecrets of Nature, it was I: For with all my Minde

and Power, I have turned over the Monuments of our Ance(tors, and

if they writ anything that was Secret and concealed, that I enrolled in my

Catalogue of Rarities. Moreover, as I travelled through France, Italy, and

Spain, I con {ulted with all Libraries, Learned men, and Artificers, that if

they knew anything that was curious, I might under {tand fuch Truths as they

had proved by there long experience. Thofe places and men, I had not the

happine(s to fee, I weit letters too frequently, earne {tly desiring them to

furni{h me with thofe Secrets, which they e{teemed Rare; not failing with

my Entreaties, Gifts, Commutations, Art, and Indu ftry. So that what { oever

was Notable, and to be de{ired through the whole World, for Curio {ities

and Excellent Things, I have abundantly jound out, and therewith Beautified

and Augmented the{e, my Endeavours, in NATURAL MAGICK, wherefore

by moft earne{t Study, and con {tant Experience, I did both night and day

endeavour to know whether what I heard or read, was true or fal{e, that I

might leave nothing una { fayed: for I oft thought of that Sentence of Cicero,

It is fit that they who de{ire for the good of mankinde, to commit to memory

things mo ft profitable, well weighed and approved, Should make tryal of all

things. To do this I have {pared no Pain nor Coft, but have expended my

narrow Fortunes in a large magnificence. . . . .

“Tf I have over-pa{ fed fome Things or not {poken fo properly of them

as I might; I know there is nothing fo Beautiful, but it may be Adorned;

Nor (o full, but it may be Augmented. Jab Bie

Waite, ANprEw Dickson.

A history of the Warfare of Science with Theology in Christendom.

Two vols. New York, 1896.

One cannot read Dr. White’s book without a profound conviction of the

truth of the quotations placed at the beginning: “Thoughts that great hearts

once broke for, we breathe cheaply in the common air.”—Lowell. “Discipu-

58 SIMON HENRY GAGE

lus est prioris posterior dies.”—Publius Syrus. “Truth is the daughter of

time.”—Bacon. “The Truth shall make you free.”—St. John, vit, 32.

Woopwarbd, JoSEPH JANVIER.

On Photo-micrography with the highest powers. Amer. Jour. Sci. and

Arts, 1866.

Report on Photographing histological preparations by Sunlight. 1871.

Magnesium, Oxy-calcium and electric lights applied to photography. 1870.

Application of Photography to micrometry. Phila. Med. Times, 1876.

Dr. Woodward, Med. Department, U. S. Army, produced some of the

best photo-micrographs which have ever been made. He says, in the re-

port on photographing by sunlight, p. 8: “And rejecting all eye-pieces and

amplifiers I have aimed to obtain a magnification of not less than 1,000

diameters by distance alone.” He commends very highly the electric light.

WRIGHT, LEwIs.

Optical projection, a treatise on the use of the lantern in exhibition

and scientific demonstration with 232 illustrations. London, 1891.

This work covers the whole field of projection with the magic lantern,

including the projection microscope. The work is by one who had much

experience, and who gave in this book the benefit of that experience. Ii

one has not a personal instructor this book will be found almost equal

to one.

ZAHN, JOANNES.

Oculus artificialis teledioptricus sive telescopium, ex abditis rerum

naturalium et artificialium principiis protractum nova methodo, eaque solida

explicatum a comprimis e triplici fundamento physico seu naturali, mathe-

matico dioptrico et mechanico, seu practico stabilitum. Folio, 770 pages,

Plates and tables. Herbipoli, 1685-6. Same, second edition, 797 pages,

plates and tables. Norimbergae, 1702.

Both these editions are in the Library of the Surgeon General’s office

at Washington. Zahn, p. 225, says: “Cum in Lucerna megalographica veri

microscopii speciem habeamus.” . . . On comparing the two editions

this expression was found in both. Zahn gives several figures of the magic

lantern which look strikingly like some of the simplest forms of the pres-

ent day. Compare also plate Va.

ZEISS, CARL.

Catalogs for the last 25 years.

Ii one would see in the most striking way the development of the

microscope and its accessories he must compare the successive editions of a

great optical manufacturing house. That of Zeiss has been selected, for it,

by universal consent, would be considered the greatest in our times.

ZEITSCHRIFT FUR WISSENSCHAFTLICHE MIKROSKOPIE UND FUR MIKROSKOPISCHE

TecuNIK. Illustrated. Braunschweig, 1884 to date.

Methods, bibliography and original papers.

PLATE I

Artes Magna Liber X. Magia (atoptrica.

leonismus XXXIV.

Alphaberurn Catoptricum

Apart Aanm AB CDEFGHiKLMNOPQRSTVXYZ

mp oe ~~ p CDELGHI ET WYO b Oth oem

Aiphab: Hebreum NWIPRYAAYOIIONCFII’ONMIINIAAN

imerfien peu OER Rd DA DI] OD) 13. Se ae

Ape je"! ABT REZHOIKAMN = OnPIeYGawe

imerfien in fperslo oy yGESHOIKVWUSOND=SLaweere

ee eran

oF) ris iy

4 o i ne

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set la

PLATE, It

Artis Magna Libtr X. Magia Pars II!.

Problema FIL. lefcopii infpectas nulto negotio exhibeat,

: Remote vero flammam intuentes , ingen-

Lacernam artificiofam conftruere , qua in

tem ignem effe exiftimabunt ; augebuot

remota diflantia {criptalegendaexhibeat. | lumen, fi latera cylindri rmtertora ex fal-

| lucerna , ea, qua hic faétum effe vi- | gido ftanno in ellipfin elaborata tuenn:.

des fi lind : = Sed inventum figura appofita fats decla-

Se nee ES AB rabit. Emanubrium, D feneftram, C m-

fj ? i

peculam concayum umibulum defignat.

uod parabolain

quantum fieri poteft,

Cet , erigatur. '

| Problema IV.

De Lucena Magice fen Thauma-

surge conftrudione.

‘Oe 1n arte Magna Lucis & ame

brz folio 767. hujufmodi Lucerne

mentionem fecerimus & fol. 793. modum

| per folem fimulacrorum in ob{curum lo-

| cum tranfinittendorum , una cum colori-

| bus ad ea depingenda requifitis tradideri-

tucems Iptta hujus {peculi focum applicetur F mus: Quia tamen in citatis locis , inven

Cavpree, flamms candelz , habebifque queficum. ) nonem hanc prorfus fingularem ab aliis

Nam tam inufitato fplendore fu gebit , ut. majortbus inventionibus adornandam re-

podtu etiam ee ae literas ope tes liquimus, accidit , ut multi rei novitate

{(i

rit

Seer

one

2 1 ad eam excolendam animum adje-

Certot, Quos inter primus fuir Thomas

Walgenftemus Danus , haud infime nota

Mathematicus , qui recolens meas in de- | fuo lucro diverfis

{cribendis iis inventiones Lucernam fol,

767¢ a nobis defcriptam , in meliorem

formam reduxit , quam & poftea magno

in Italia principibus ven-

didits

PEATE LET

Magia Catoptrrca.

didit, ut promde jam Rome res poene

valgaris fit. Non eft autem alia inter illam

& a nobis defcriptam Lucerrsim differen-

t12, quam quod complurium :maginum

fpecies in obfcuro cubiculo dictus Wal-

genitenius oftendat fatis nitidé & polité ,

nec non cum fumma fpetantium adinira-

tione. Nos in Collegio noftro 1p ob{curo

cubiculo, 4. novifiima fummo intuenttum

ftupore exhtbere folemus. Eft autem res

vifa digniffima, cum ejus ope, vel inte-

gras fcenas fatyricas, Tragicas theatrales

& fimilia ordine ad vivum exhibere liceat.

Artificium verd Catoptricum , quod

769

que finulacra rerum defdripta fant , re-

| fle€tentibus 1n immobili interiors alicujus

domus aut cubiculi pariete , coloribus ad

vivum cxhibemus ommic ea , que per Lu-

| cernam mobilem monitrari folent; quam-

vis eam eodem inlocomodum fine fo<

lis raduis, per Speculum concayuin aut leg-

tem diaphanam res reprefentandi docea:

mus. Que omnia hic fofius profecums

fum, ut Leétor, unde hac nova arcana

Lucernz (quam pon immeritd Magicam

& Thaumaturgam , 4 mirfica rerum qua-

rumcunque tandem yn ob{curato cubiculo

aut sntempelte nodtis filentio reprafen-

nos fol. 793. Artis Magne Lucis & um-| tatione appellandam duximus ) originem

brz docuimus, non differt ab hac nova fuam traxerint. Quibus premiffis oi! jam

Lucerna , nifi quod illa per mobilem Lu-/rettat, nifut fabricam ejus paucis expo-

namus.

cernani, nos radiis folis in fpeculum, in

Fiat ex ligno receptacolum A. B.C.D.

| planum probe elaboratum ponatur, 18

deinde in L caminus, ut Lucerna peri-| quo coloribus aqueis & diaphanis quid-

‘fum fomam fuum emuttere poflit , Lucer- | quid volnens pingatur ; hoc lee intra

na vero K in medio ponatur vel affixa filo | cubiculnm VTSX tn muro can ido lumen

ferreo vel fupra fulcrum M é regione fo- lucernz vitrum lentculare tranfiens ima-

raminis H, jntra quod tubus palmans) 0 H vitro. plano depictam (que

commuttatur, in fubi verd principio I Jen- anverlo fia in vitro ponitur ) rectam 8c

riculare vitram melioris notz inferatur |in muro grandiorem exhibebit , omnibus

in foramine vera) fei fe Reiclba El vactta| coloribus ad vivumexpreffam, Nota ta-

Pppp 3 mens

THE PROJECTION MICROSCOPE 59

EXPLANATION OF PLATES

Plate I

Reproduction of a part of Kircher, p. 792, showing the use of sun-

light for illumination, and that the objects to be shown in the darkened

room (camera obscura) were painted on plane metallic mirrors (N, V, R).

Below are Roman, Hebrew, and Greek letters showing the usual appear-

ance and also the mirror image which must be painted on the mirrors

in order that the image on the screen will appear in the normal form

after passing the lenses. In fig. 1 at the top of the cut is a kind of

optical bench showing the metallic mirror (4) with letters in reverse

upon it but erect in the screen image at the right. The projection lens is

shown at B.

Plate II

Projection apparatus shown by Kircher, p. 768. The radiant is a lamp

(K) with a concave mirror or speculum (S) for concentrating the light

upon the object. The lamp is in a dark box. The objects are on a long

glass slide as in the present day toy lanterns. The image at G illustrates

one of the amusements of the period.

Plate III

Magic lantern shown by Kircher, p. 769. The apparatus in this case

is separated by a partition from the room in which are the screen and the

spectators.

60 SIMON HENRY GAGE

Plate IV

A. Projection or solar microscope (Adams Essays, 1771, plate 6,

figs. 4, 5, 6, 7; 8). Fig. 4 shows the movable mirror (K-L) placed out-

side the shutter in the sun; O-P, screws in the square plate to fasten the

instrument in the shutter; M-N, thumb screws by which the mirror is

turned to hold the sun’s rays in the right direction. The large tube,

A-C-D, contains the condenser and receives the shorter tube, fig. 5.

Fig. 5 shows the tube into which the objectives are fixed. If for large

objects the lens (fig. 6) is screwed into the end, g, for smaller objects,

the objectives are arranged in a piece (fig. 8) sliding into the opening

at g. Notches along the objective slider indicate when the lens is cen-

tered. The specimen to be examined is inserted at h. For high powers

the substage condenser shown at fig. 7 is put in the tube between d-h.

At b is a rack and pinion for focusing the object.

B. Thompson automatic 90-degree arc lamp with Prof. Wm. A. An-

thony’s parallel movement, arc-striking device. M, electromagnet; A. M, arc-

striking mechanism consisting of an armature of soft iron, and parallel

links or hinges allowing the upper carbon to extend to the right and down-

ward when the current is off. This brings the carbons in contact. When

the current is on the electromagnet pulls up the armature, thus lifting the

upper carbon vertically and to the left. The arrows indicate the direction

of the current.

Plate V

A. Shows the arrangement of parts for projection with a photographic

objective oi! 105 mm. focus. No tube is used and objects up to 60 mm.

in diameter can be projected.

B. Shows the projection microscope in position for high powers. The

tube of the microscope is large (50 mm.) and short (9 to 10 centimeters),

so that the field may not be restricted. Al, automatic arc lamp with An-

thony’s arc-striking device; Cw, condenser and water-bath; Sw, stage and

stage water-bath; Ob, Sc, objectives on a triple nose piece with a 25-centi-

meter screen just behind them; Lb, lathe bed on which the different parts

move independently.

Plate Va

From Zahn’s Oculus Artificialis, 1685 and 1702, showing the use of an

Amplifier.

Fig. 2. Projection with a double convex lens (C-D), and divergence of

the rays, thus giving a large image, by means of a plano-concave lens or

amplifier.

This is the earliest illustration I have found showing the use of an

amplifier in projection. (See p. 37 and figs. 13, 17 and 18 in the text.)

Fig. 3. Projection by means of two convex lenses. The first, (C-D),

gives a real, inverted image at O-K; the second, (E-F), projects this real

image and re-inverts it, thus giving a final erect image (G-H). Compare

the modern projection microscope when an ocular is used (fig. 14 in the text).

PEATE: EV.

London Printed tor Bc Pubtjhid by Geo Adam: "60 Fleet Street Hay 267178 7.

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PLATE Va.

Tubum ex mens Lentibus convexis apparacum, pofluntque oculares Lentes effe duz

vel una rantum habebicque fe ordinatio Tubihujus per modum Telefcopu Aftrono-

mici, quodalias oculo applicacum objecta exhibet everfa , hic aucem debité aptatum

Solem-oftendét erectum,

Cum primo modo immiffioinltituitur, debet Tubus communis five Telefco-

pium Hollandicum paulo magis produci, quam opus eflet ad ordinari¢ objeCa

fivédebet Lens cava pauld amplius 4 Lente convexa

A PRELIMINARY LIST OF INVERTEBRATES, PARA-

SITIC OR OTHERWISE NOXIOUS TO MAN,

COLLECTED IN PORTUGUESE WEST

AFRICA: 1904-1906

By F. CREIGHTON WELLMAN

It has seemed to the writer that a tentative list of the animal

parasites and other allied enemies of man thus far encountered in

what is as yet almost virgin territory to scientists ought to be of

interest to those concerned with microbiology and the geographical

distribution of parasitic disease. I have accordingly gone over

my small collection of the fauna of the Angola highlands (consisting

of about 700 species) and prepared the following list of Protozoa,

Vermes and Arthropoda which are noxious to man—using the term,

sensu generali, to denote any invertebrate animal which may be

inimical during any stage in its life history to the health or comfort

of human beings. Speaking strictly from the academic point of

view some of the worms and arthropods mentioned are not micro-

scopic. However if the worms are not microscopic their ova (upon

the detection of which generally depends the diagnosis of the

adults’ presence) certainly are, and while the insects, etc., are

macroscopic in size (as every entomologist knows) few of them

can be specifically determined with the naked eye alone. So, even

if I did not know of the highly laudable interest taken by this society

in practical parasitology, I should not feel that I were forcing a

personal hobby upon your attention. Writing purely as a micro-

scopist, I have not discussed the symptoms and pathology of the

diseases subtended by the various organisms mentioned, and indeed

space considerations have made it possible for me to transcribe

only the briefest possible extracts from my notes on morphology

and bionomics. I have included an insect and a mite (marked in

62 F. CREIGHTON WELLMAN

the list by asterisks) which are only suspected of possibly playing

a mischievous role in their larval stage. I have considered more

than the occasional mention of related parasites of animals which

are common or of peculiar interest, as lying without the scope of

this report.

PROTOZOA

Rhizopoda

Entamoeba histolytica Sch. What I have referred to this species

is not seldom seen. It is not, however, always found in connection

with dysentery, but can occasionally be demonstrated in the faeces of

apparently healthy natives. The organism is somewhat larger than

is usually stated for this species and measures 0.007—0.01 mm. in

diameter. The closely allied E. coli Lésch can be certainly differ-

entiated from the foregoing only by a study of their pathological

effects. What I suspect to be another and much smaller species than

either of these is more rarely seen. An allied form is also found

in the intestines of pigs. Charcot-Leyden’s crystals are often seen

in mucous containing such organisms.

Mastigophora

Trichomonas sp. Seen in intestinal mucous of a child with slight

diarrhoea; measures about 0.02x0.01 mm. Three flagella.

Cercomonas sp. (? hominis Dav.). These spermatozoa-like or-

ganisms were also from a case of diarrhoea. Body measures about

0.01 mm. and flagellum about 0.02 mm. in length.

Trypanosoma gambiense Dutt. My specimens from cases of

human trypanosomiasis seem to be morphologically identical with

Dutton’s published descriptions. The posterior end is truncated

and the arrangement of the centrosome, nucleus, and flagellum, as

well as the “set” of the organism on the slide correspond with the

same points in gambiense. The protoplasm stains somewhat irregu-

larly with Romanowsky stain, taking on a basophile reaction. The

centrosome stains a dark purple hue and the flagellum (which

stains pink) seems to rise from or near it. The nucleus lies near

the middle of the body and often occupies more than two-thirds

the width of the parasite. It is oval in shape and stains red like

PRELIMINARY LIST OF INVERTEBRATES 63

other chromatin material. Following are measurements of a stained

specimen :

TR caial o's cic ss faccreis he ws ahha nle ovaeale eh ea edie en 2 (about) 24 p

Pepe EES MENG TEE sn er atcpclstaten esis. ciaisy Sos a mre atom cites wlctehwcal ae 3.5

Distance of centrosome from posterior end................ 3 pb

Menethy Of. free MAGE luis: 0.5 opeiscodorate is ase ces d = ewlere's« (about) 9

ieerigin of diatueter Of nucleus... Jo.) «36 <5 5s smysieleiceiedie eae 6 4 op

I have also found several animal trypanosomes which may be

mentioned here.

Trypanosoma avium Dan. Found in blood of a wild dove, Treron

calva. Plump, fish-shaped, with the posterior end rather sharp.

Faint longitudinal striae or myonema lines can be made out in the

protoplasm as well as very fine granules. The nucleus extends

about two-thirds across the body and seems to almost completely

fill the space in which it lies. The flagellum and undulating mem-

brane are wavy in very regular curves. The free flagellum is

comparatively short. Following are the principal measurements

of a stained specimen:

Length Se ciatoneixisl sel al chalelialetaleiclichslnoltalelctstsaiaiet= teistel sisi ai'el oi sin\telstaliel/alelel stele 28 B

RERUEOSE WAGE: c.g titeeie te. s's abiciae Gcleos Mig hwlew died oa ayen- dare rn

Distance of centrosome to posterior end..................+6- 4p

Penetnvoiciree: Hage aay 5. 5.50022 2 > sis xis 2 ioe nls aeynidleleaty ‘(about) 8 p

oneesh tiamicter Of TUCIEUS s « o.5cj<6 nafa0- ode oes oe went m tis warn aie 6p

Trypanosoma lewisi Kent. From rat (Mus decumanus). No

differences in structure between these and specimens secured by me

from sewer rats while I was at the University of London in 1904.

The slender contour and narrow undulating membrane are well

shown. I give measurements of a stained specimen:

ste We nes seq ts 2 SiaciS clot macs avers scree «ac et Ne ets aoa 60

Rec ategt AOE. <<. scte score aise goceite See are eS OEE tive 2 :

Distance from centrosome to posterior end................. ce

enern of tree: flagellum... eben oe a peso nee Sin tals c 10.5 p

ieareest diameter Of nicleuS.%. 3250.41: 5 Shea ee ole (about) 3 op

Trypanosoma mega Dutt. et Todd. My specimens from frogs

(Rana tuberculosa) resemble rather closely the descriptions of this

species. The same parasite or a very similar one has been found

recently on the Congo. The distinguishing points of this trypano-

some are the large size, the longitudinal markings, the great distance

64 F. CREIGHTON WELLMAN

of the centrosome from the posterior end, the ill-defined nucleus

and the tortuous flagellum. Following are the measurements of a

stained specimen:

ee ttg Seer es Maar ae Se tee ce wecetls cL eres Pn (about) 60 y

Greatest width oo 32) ss. Pees se oe tae ci Lees oaks teen ee eree 8 om

Distance of centrosome from posterior end................ ee

Penciwor irce Hacellant: : 2.50552... 5.50- see see ser (about) 18 yp

Bargestidiameter, of nucleus..42,4: cicero toe ie eee eee 3.5 ps

Spirochaeta duttoni Novy et Knapp. In including spirochaetes

here I do not necessarily imply an opinion as to the protozoal nature

of these organisms. The opinions of Schaudinn et alu are well

known. Still should the bodies turn out (as most investigators now

predict) to be schizomycetes, no great harm will be done by record-

ing in a report on animal parasites the occurence of a few species

in West Africa. S. duttont, the cause of “tick fever” in Africa,

is not uncommon. The writer was among the first (if not the first)

to point out that the species can be morphologically distinguished

from S. obermeieri by the difference in the curves of the two

organisms.

Novy and Knapp in differentiating duttoni from obermeiert

adopt this character in the following words: “A further important

characteristic is afforded by the width of the turn of the whole

spiral. In the case of S. obermeieri this width measures quite con-

stantly 1.0 », whereas in S. duttoni it amounts to 2.0 to 2.7». This

fact is referred to by Wellman when he says that the spirillum

studied by him in West Africa ‘dies in rounder and more flowing

curves than are generally described and pictured,’ etc., etc.’

Spirochaeta pertenuis Cast. While my discovery of spirochaetae

in the disease called “yaws” was entirely independent and my pre-

liminary note was on its way to America long before Castellani’s

announcement reached me, yet I have of course explicitly? accorded

him priority in the matter since he was the first to publish his

observations. It is rather remarkable that the same discovery should

*Journal of Infectious Diseases, May 18, 1906, p. 382.

*Vide Journal of Tropical Medicine, Dec. 1, 1905, p. 345; Boston Medical

and Surgical Journal, May 3, 1906, p. 489; Archiv fiir Schiffs-und Tropen-

hygiene, 1906, p. 727, etc.

PRELIMINARY LIST OF INVERTEBRATES 65

be almost simultaneously made in two such widely separated coun-

tries as Ceylon and West Central Africa. There are several types of

organisms found in the lesions. I think my species from the un-

ulcerated papules are rather closer to the pallida type than Castel-

lani’s description of his pertenuts is.

Spirochaeta spp. I have announced the demonstration of spi-

rochaetae in a very peculiar and severe marginal ulceration of the

gums endemic in Africa and South China. The organism can,

I believe, be distinguished from the spirochaetae usually seen in

healthy mouths. My investigations of this subject will be published

in a short time.

Sporozoa

Haemamoeba vivax Gross. et Fel. Comparatively rare. In a

series of 531 blood examinations made in this colony for the Societa

per gli Studi della Malaria only 14 individuals were found with

quartan parasites... This form of malaria among the blacks is an

almost negligible quantity both from its rarity (0.26 per cent) and

from the mildness of the symptoms set up. It rarely causes rise of

temperature in the native, the schizogenesis of the parasite going

serenely on without interfering with the host’s health or happiness.

Haemomonas praecox Lav. Very common. In the series of

blood examinations just mentioned 285 individuals were found to

harbor this species. To this should be added 153 individuals not

showing parasites in the blood but whose large mononuclear niu-

clei formed 20 per cent or more of the whole number. These facts

taken with the results of the spleen index of 513 natives, made at

the same time, have led me to calculate that about 83 per cent of

apparently healthy natives show parasites or other evidence of

malaria in their blood and that moderate invasion is almost uni-

versal and that at least half the population are heavily infected.

Infusoria

Balantidium sp. An infusorian found in faeces. No symptoms.

Measurements: 0.15 mm.x0.06 mm. Peristome large, ectosare

ciliated. Macronucleus round or oval. A similar form is found

here in pigs.

*Individuals who were not taking quinine.

66 F. CREIGHTON WELLMAN

VERMES

Trematoda

Paramphistomidae

None of the Paramphistomidae have been seen in human beings

although I have frequently found an Amphistomum in the stomach

of the lesser reed buck (Cervicapra bohor).

Fasciolidae

Fasciola hepatica L. var. angusta Raill. Several cases seen. This

fluke is also common in oxen and antelope of different sorts. Meas-

urements: Average of formalin specimens, 30.5x7 mm. Eggs:

0.15 x0.85 mm.

Schistosomidae

Schistosomum haematobium Bilh. Fairly common. Infections

all slight. Ova not very plentiful and lesions insignificant. Ova

with side spine from rectum occasionally seen. The differences in

the eggs are very striking and suggest different species.

Cestoda

Bothreocephaloidea

Dibothriocephalus latus L. Not seldom seen. Gotten through

eating raw fish.

Taeniidae

Hymenolepis sp. One incomplete specimen.

Taenia saginata Goeze. Not very common. More often seen in

the interior where beef is more easily obtainable.

Taenia echinococcus v. Lieb. Rare. One case. I have seen the

scolices | bladderworms ?—Ed.] several times in oxen.

Taenia sp. (nov.). I have been informed that this specimen

represents a new species but the description has not yet been pub-

lished.

Nematoda

Anguillulidae

Gen. et sp. incert. This minute entozoal embryo was found in

the blood of a negress and was first mistaken for a filarial embryo.

I referred it to Dr. Low and Sir Patrick Manson on the occasion

PRELIMINARY LIST OF INVERTEBRATES 67

of my last visit to London. They both thought that it probably

belongs to the Anguillulidae and that its presence in the blood was

doubtless accidental.

Angiostomidae

Strongyloides intestinalis Bavay. Seen several times (the rhab-

ditiform larvae) in faeces, once in connection with a severe diar-

rhoea. The remarkable heterogenetic reproduction is difficult to

see, as the parasitic-living generation does not appear spontaneously

in the faeces.

Filariidae

Filaria perstans Mans. Not common. About one native out of

every hundred infected. I have been working for about two years

on the life history. Although my investigation is not complete I

have succeeded in proving certain points of interest. [Some slides

illustrating which were exhibited at the meeting —Ed.] The pres-

ence of the parasite causes no symptoms.

F. nocturna Mans. Very rare in the highlands. Common in the

littoral zone where elephantiasis abounds.

Filaria sp. One solitary embryo in the urine of a Bantu man.

Sheathless and sharp-tailed. Much smaller than F. demarquayi.

Trichotrachelidae

Trichocephalus trichiuris L. Rare; two cases. The characteristic

ova (like an oval tray with handles at the ends) measure about

0.05 x 0.22 mm. No symptoms.

Strongylidae

Strongylus spp. I have not yet found any of these forms in

man but a large number infest animals (e. g., Strongylus contortus

in goats). I have also found an undescribed species of Cylichnosto-

mum frequent in the same host.

Ankylostomum duodenale Dub. Very common in some districts ;

13 infections out of 310 random faeces examinations from different

parts of the country.

Ascaridae

Ascaris lumbricoides L. Found in 158 infections out of 310

routine faeces examinations. Almost universal in children. Some-

times occurs in enormous numbers. J have seen 96 worms passed

68 F. CREIGHTON WELLMAN

by a child two years old the result of a single dose of castor oil and

santonin.

Ascaris lumbricoides var X. In February, 1904, I described

some very peculiar ova from a variety of Ascaris infesting a white

child then resident in this colony. These were bean-shaped, with an

opaque, rugose, albuminous envelope. There were also some anat-

omical peculiarities of the adult worm that led me to suspect that

I was dealing with an instance of variation, or possibly even a new

species. Quite recently I see that O. T. Logan of China states

that he has found the same type of eggs from adults that show some

of the same peculiarities mentioned by me in my original note.

Dr. Logan refers to the correspondence between his observations

_and mine and asks if we are dealing with a new species. I should

like to see the question settled and propose to place some specimens

in Dr. Ward’s hands soon for examination.

Oxyuris vermicularis L. Common in children.

Gordius sp. This is not a human parasite but a very interesting

worm which in its larval stages infects the migratory locust

(Schistocera peregrinatoria). Free living in its adult age. The

blacks believe it invades human beings.

Hirudinei

?Hirudo sp. Found clinging to a native porter after having

crossed a marsh. I found another species in a plain half a mile

from the nearest water. The report on these leeches (which were

sent to Paris for determination) has not yet been received.

ARTHROPODA

Arachnida

Scorpionida, Araneida

Several scorpions and spiders which have not yet been reported

on have poisonous bites. Among the latter are a tarantula and a

large species of Nephila. A huge pedipalp is also feared by the

natives but is probably harmless.

Acarina

Trombidiidae

*Trombidium grandissimum. In native camps. I have found -

a larval mite which bites like Leptus autumnalis. I suspect the

PRELIMINARY LIST OF INVERTEBRATES 69

adult may be the large red mite above named. It is predatory in

its habits.

Ixodidae

Rhipicephalus decoloratus Koch. Found on cattle and other

domestic animals.

Amblyomma varigatum Fabr. Cattle, etc. I have never seen these

ticks listed as parasites of man but they readily bite human beings.

I have personally been victimized by A. variegatum.

Ornithodoros moubata Murr. Usually known as O. savignyi Aud.

var. caeca Neum., but the name I have given is the earliest one

given the tick and Dr. Nuttall of Cambridge writes me that he will

use it in the new monograph of ticks. Very common. Within an

hour’s time I have found as many as a hundred in an old native hut.

Hides by day in thatch, walls, etc., and comes out at night to bite.

Bites man and all domestic animals. Is the disseminator of Sp1-

rochaeta duttont and probably an officient for Filaria perstans.

[Preparations of this tick fed on perstans cases were exhibited at

the meeting.—Ed. |

Sarcoptes scabei L. Common. Also other varieties on animals.

M yriapoda

A number of venomous centipedes are common. Two different

genera of Julidae (millipedes) are remarkable for causing intense

smarting and burning when they crawl on the body. Their poison-

ous secretion is probably from the foramina repugnatoria which are

in the sides of the segments and look like tracheal stigmata. The

real stigmata are on the ventral surface of the segments and so

minute as to be easily overlooked. These millipedes belong to the

genera Spirostreptus and Odontopyge.

INSECTA

Orthoptera

There are several grasshoppers (Phymateus, Pamphagus, etc.)

which defend themselves by means of a stinking secretion which is,

however, nothing more than unpleasant to the nostrils. Three

species of giant earwigs can draw a large drop of blood with their

pincers and one so frequently introduces septic matter in this way,

70 F. CREIGHTON WELLMAN

that the natives greatly fear it and believe it to be venomous.

These species are Apachys chartacea Pal.-Beauv., A. reichards

Karsch, and a third representing a new genus and species which

will soon be described.

Hemiptera

Reduviidae

Several species that bite man have been collected. The most

interesting of these is Phonergates bicoloripes Stal, which also

habitually mulcts Ornithodoros moubata (vide supra) of its ingested

blood.

Acanthiadae

Clinocoris lectularius Wer. Common.

?Cimex sp. A large bug which has not yet been determined.

Pediculidae

Pediculus capitis de Geer. Universal among natives.

P. vestimenti Nitzsch. Very common. Phthirius pubis has not

yet been seen.

Lepidoptera

The larvae of a number of these insects defend themselves by

means of stinging hairs and bristles. The effect of touching some

of these is astonishingly severe, even alarming symptoms being set

up. The worst of them belong to three different families, viz.,

Limacodidae, Arctiidae, and Liparidae (resembles larva of some

Tortricidae).

Diptera

Culicidae

I have made a large collection of these. Only the most common

can be mentioned here. These are: Anopheles welcomet Theob.,

Myzomia funesta Giles, Pyretophorus austentt Theob. (nov.), My-

zorhyncus umbrosus Theob., Cellia squamosa Theob., C. pharoensis

Theob., Danielsia wellmani Theob. (nov.), Culex hersutipalpis

Theob., C viridis Theob., C taenitorhynchoides Giles (nov.), etc.,

etc. So much has been written about mosquitoes that I must refrain

from discussing their bionomics at this time, excepting to say that

M. funesta is the great carrier of malaria and that I have found

PRELIMINARY LIST OF INVERTEBRATES 7a

as high as 13 per cent of this species to be infected with malarial

parasites. Two new points in morphology are the following :Hepta-

phlebomyia simplex Theob., has hitherto been known only from

female specimens, which differ from all other mosquitoes by having

a distinct seventh scaled wing vein, upon which character Mr. Theo-

bald has founded a new genus and a new sub-family. From some

specimens bred by me and sent to the British Museum last year,

however, appears the remarkable fact that the males do not share

this peculiarity. In these there is no true scaled seventh vein, but

the sixth is bent at right angles near the edge of the wing. A suite

of specimens of C. hirsutipalpis Theob., also bred from eggs, which

I sent to the Museum at the same time, also showed sexual di-

morphism (the males having no pole band at the apex of the palpi)

besides remarkable variation in size, some being a third smaller

than the type.

Chironomidae

Ceratopogon sp. Small sand fly. Very vicious.

Simulidae

Simulium sp. Likewise very vicious.

Tabanidae

Tabanus sp. (near rubricundus Walk). Occasionally bites man.

Tabanus sp. (near latipes Macq.). Bites animals and man. These

two species are probably new.

Tabanus socius Walk. Bites man also.

Haematopota spp. Eight or ten new species which will soon be

described. Bite man.

Chrysops spp. Several species. Bite man.

Muscidae

Stomoxys spp. Bite domestic animals and man,

Glossina palpalis wellmani Aust. This is probably the representa-

tive of the species in South Africa. Glossina palpalts palpalts is not

found in this district. Wellmani has also been found in central and

east Africa, and reports show that it takes the place of palpalis

(south of 10° S. lat.) to at least 30° E. long. I have elsewhere

shown that wellmani is the carrier of human trypanosomiasis in my

72 F. CREIGHTON WELLMAN

region. Other species (longipalpis and tachinoides) are found in

the northern part of the colony.

Musca domestica L. Sometimes lays eggs in open wounds. This

species with Homalomyia scalaris and Pycnosonia chloropyga are

probably the worst bacteria carriers among our local diptera, and

are doubtless responsible for much enteric fever and other diseases.

Sarcophagidae

Sarcophaga africa Wied. I have proven this species (and S. albo-

fasciata Macq. as well) to be capable of causing severe destructive

myasis. I also found larvae living under the skin of man. Dr. L. O.

Howard writes as follows concerning a specimen of these latter:

“The larva removed with a pair of forceps from under the skin is

most interesting. It appears not to be a muscid, but its affiliations

are with the Sarcophagidae, the true flesh flies; but I know of no

record of the occurrence of the larva of this family under the skin

of a human being. Still, new things are coming up all the while,

and we may have here something absolutely novel.”

Auchmeromyia luteola Fabr. The larva of this fly sucks human

blood and is one of the commonest pests of the region. I have

elsewhere published a study of its life history.

Anthomyidae

Anthomyia desjardinsu Macq. A case of serious intestinal myasis

caused by this fly. A number of the larvae were passed per rectum.

I am not aware of this species having been previously convicted

of causing myasis.

*Hylemyia fasciata Walk. A case of maggots infesting a wound

is possibly due to this fly. The larvae were not bred out, however,

as they were in my other cases of myasis.

Hippoboscidae

Hippobosca sp. (rufipes Th.). Not very common. Oftenest seen

on donkeys. Sometimes attacks man.

Siphonoptera

Pulex irritans L. Not very common.

Pulex cheopis Roth. Commonest flea on rats, etc. Bites man.

Pulex spp. Several species of fleas sent to London have not yet

been heard from.

PRELIMINARY LIST OF INVERTEBRATES Df

Ceratophyllus fasciatus Bosc. Dogs, etc. Bites man.

Sarcopsylla penetrans L. Universal throughout the region. Lives

in dust, etc., in old kraals and camps. It bites (and the gravid

female burrows into the flesh of) all warm-blooded animals, includ-

ing man. Causes lameness, deformities, etc.

Coleoptera

?Drilus sp. A larva of a beetle belonging to the Malacodermata

and probably to the genus just named; is armed with poisonous

bristles which, when one steps on it with the bare feet, break off

and, working through the thick sole into the flesh cause pain,

inflammation and even sloughing. The presence of the bristles in

the skin can be demonstrated by looking at sections under a low

power of the microscope.

Hymenoptera

A large number of wasps (e. g., Pelopoeus spwifex, Synagris

cornuta, etc.) have painful stings, also ants (Polyrhachis muilitaris

Fabr., etc.) Mutillidae (two common species are Odontomutilla

thymele Pér, and Barymutilla pythia Sm.) and bees. Among the

latter may be mentioned two giant carpenter bees, Mesotrichia mixta

Rad. and Xylocopa tarsata wellmam. Ckll. These latter not only

sting severely, but are also of interest to microscopists in their role

of hosts for the remarkable mites belonging to the genus Greenta

which live in the peculiar pouch which opens on the basal surface

of the first abdominal segment of these bees.

In conclusion it should be understood that the foregoing list does

not pretend to form a complete report on the human parasites of

this district. For I have mentioned only species which I have per-

sonally collected and studied and have excluded as being com-

paratively unimportant nearly as many species as are mentioned in

the list. I am afraid that I have given you little more than a bare

catalogue of names, but fear of making my paper of unwieldy

length has kept me from presenting many interesting details. I

hope, however, that the mention of several species not before shown

to be human parasites and several not before reported from this

part of the world will, at least, be of interest. A list of this kind

is necessarily always incomplete and it is my intention to present

74 F. CREIGHTON WELLMAN

on some future occasion the result of additional observations. Be-

fore closing I must acknowledge my indebtedness to several

zoologists, chief among whom is my friend Dr. Walther Horn of

Berlin, president of the German Entomological Society.

BIBLIOGRAPHY

My original communications on a number of the species above men-

tioned may be seen in the transactions of several European scientific socie-

ties, chiefly the Deutsche Entomologische Gesellschaft (Berlin), the Société

de Médicine et d’ Hygiene Tropicales (Paris), and the Societa per gli Studi

della Malaria (Rome). A large number of notes and articles have also been

published in the Journal of Tropical Medicine (London). A few of the

most important papers in other journals have the following dates: New

York Medical Journal, Aug. 12, 19, 26 and Sept. 2, 1905. Journal of Medical

Research, Jan., 1906. Entomological News, Feb., 1906. Journal of Infectious

Diseases, Nov. 25, 1905. American Jour. of Medical Sciences, May, 1906.

Medicine, July, 1906. Journal of Hygiene (Cambridge, England), July, 1906.

Deutsche Medizinische Wochenschrift (Berlin), No. 27, 1906. Polytechnia

(Lisbon), No. 5, 1906. Annals and Mag Nat. Hist. (London), p. 242 et seq.,

1906. Most of my communications relating to microbiology for several years

past have been summarized from time to time in the Bulletin de 1! Institut

Pasteur and the Archiy fir Schiffs-und Tropenhygiene.

Benguella, West Africa.

March 15, 1907.

OBSERVATIONS ON THE MICRO-FAUNA OF AN

OREGON POND

By ELDA R. WALKER

WITH ONE PLATE

The material under consideration in this paper is from north-

western Oregon, a region almost completely separated from the rest

of the continent by high mountain ranges on the east and south

and bounded on the west by the ocean. This part of Oregon is

conspicuous for the absence of large bodies of standing water.

Only small ponds, such as the one to be described, are found.

The only data on the micro-fauna of this part of the West

concern Seattle, Washington, and Crater Lake, Oregon. There are

a few records also from Spokane, Washington, and from the Flat-

head region of Montana and British Columbia which, although east

of the Cascade Mountains, belong to the Columbia system of drain-

age. These localities are widely separated from each other as well

as from the pond studied, which is near Forest Grove, Oregon.

It is interesting to notice that, although so separated from the

rest of the continent, the species, except a few new or very rare

forms, are those commonly found in similar bodies of water.

I am indebted, for assistance in this study, to Professor Henry

B. Ward, under whose direction the work was carried on, and, for

help in identification of species, to Doctors C. Dwight Marsh,

Charles Fordyce, Robert H. Wolcott, Ruth Marshall, H. S. Jen-

nings, R. W. Sharpe, and Joseph H. Powers.

Todd’s Pond (plate v1, fig. 1) is about a mile in length and a

half a mile in width. It is fed by a stream which arises from moun-

tain springs and is augmented by water from the tile ditches drain-

ing neighboring farming land. This stream flows for some distance

through swampy land covered with willow, maple, fir, and ash

trees. Where it enters the pond the stream is about thirty feet

wide and three feet deep. Its bed is of mud and a thick growth of

pond-lilies and other water plants fills the stream. The water is

76 ELDA R. WALKER

sluggish but clear. The pond itself is shallow, six inches to two

feet in depth. On the south and west the bank rises rapidly to high

pasture and farm lands. On the north and east the pond ends in

a swamp filled with rushes. There is a dense growth of pond-

lilies, cat-tails, willows, rushes, many algae and other water plants,

filling the pond, and its south end is covered with a thick growth

of willow, ash, and, a little distance from the water, fir, maple,

cedar, and oak trees. The vegetation is so dense that during the

summer the water is only visible in small areas. The bed is of

mud, covered with a thick layer of debris, decaying vegetable

matter. Many trees have fallen into the pond where the partly

decayed, moss-covered trunks remain. The water is clear but

slightly yellow. The color is probably due to the wash of fir for-

ests near by. There is no odor from the water except in very dry

seasons when the water is exceptionally low. The pond was much

larger until a few years ago, when the beaver dams at the outlet

were blown out to keep the stream open.

The dense growth of vegetation in the pond, the shallow water

and the forests of large fir trees within a sort distance on all sides

make it impossible for the light breezes that occur during the sum-

mer to have much effect in stirring the water. During the entire

summer the water was never moved enough to prevent its being

clear, although the bed was of such loose light debris that great

care was necessary in collecting to prevent the net from becoming

filled. The elevation is about one hundred feet above the sea level.

Collections were made two or three times a week from July 7 till

August 25, 1905. Each time specimens were taken from a bridge

crossing the inlet (plate v1, fig. 1, B) and from the west side of

the pond (plate v1, fig. 1, W). Collections were also made from

the same places in the summer of 1906. The fauna of the two sta-

tions differed in such minor points that they are considered to-

gether.

MICRO-FAUNA OF AN OREGON POND

The following table gives the

dance on each day. The signs —,

rare, medium, and abundant.

Volvox spermatosphara.......

Volvox perglobator. ......

Eudorina elegans.........

Pandorina morum........

Stentor pyriformis........

Miydra viridis... -..----

EKOUIMOSAS fy ac.2,2 = fs

Dorylaimus attenuatus....

Trilobus pellucidus .......

Monostyla bulla. .........

Monostyla lunaris.........

Anuraea aculeata.........

Anuraea hypelasma.......

Anuraea cochlearis........

Diurella stylata..........

Polyarthra platyptera.....

Furcularia longiseta.......

Cathypna luna. ..:.......

Pterodina (elliptica?)

Dinocharis tetracis........

Cyclops prasinus.........

Cyclops albidus..........

Cyclops serrulatus. .......

Cyclops fimbriatus........

Cyclops viridis. ..........

Pleuroxus denticulatus... .

Alona intermedia.........

Simocephalus vetulus. ....

Eurycercus lamellatus........

Scapholeberis mucronata. ....

Macrothrix laticornis var. ....

Chydorus sphaericus. .....

Cypris ophthalmica.......

Chironomid larva.........

G@ulieid larva: 2.52. ..;2-...

aL ye

Salpina mucronata.. . Sie

Metopidia lepadella.......

Euchlanis deflexa.........

a en i

>| >| >|] >

Beene

= | F> | = | Fs

* *]4+ |+

* i

* * * *

ie ee

| J+ J+

Pal doe

| + j+

eal aaa

Gai he

I+ |+ |+ J+

+ | * [+

+ |+ [+

ea hae be

an +

+ |+ | *]+

ae ete al

+ *

+ [+ | * |+

Saal aed ele

* ee *

— |+ * *

— *

ents

+ |+

+ |+ {+

ae Se” ae

el hil eso

att yttt ++

+ yt++ +44

+++++

ll lal ++

J++ +

ele

Loot lel ea

species and their relative abun-

+, and *, indicate, respectively,

m= le IA

a2l/aj|e

<q j]/—a)<4

+ |+ |—

+ |—

+ |j— j—

— |j+ |—

+ —

+ j|— |—

* | * | *

+ |+ |+

= | t- *

+ |+ |+

78 ELDA R. WALKER

Limnesia sp., juv............ |

Piona reighardi. (6045. 9.5.1 — |\— |

joo] | S| cesahoee

r{=(S (S/S Rleglelelielelz

=\zlz)eleie| 8/88/5818

515 ay fads acs ai oa

May-lyilanvasss sence — Ee tee | + | *| ae

Hydrobatid bug larva........ | | = | | Se

Beetle larva (Gallerucella) ... | | _— |

May-fly nymph (Baetis sp.). . . | | — —_— |-

Coniaid Waive peon was ecieasee | | — =

Dragon-fly larva........... ae | | ==

Hemipterous larva (Corisid) .. | | | tas

Beetle (Cnemidotus sp.)...... | | ==

| | | | | |

| | | |

Mideopsis orbicularis.... . Segoe i= | === ih ay) yl —

Limnesia histrionica.......... = ae at ry |

Limnesia undulata........... { {| j= |j— =e =e

Piona sp., juv..... a ete te ae

Lebertia sp., undet ..........

Unionicola crassipes ......... | — |—

Arrhenurus prominulus....... | | — |4+ |

Arrhenurus sp., juv.......... | \—

Arrhenurus krameri.......... |

Atractides sp., juv...-.2--.-- | | |

OxtN Ap: ANOSC io dish. Bea | —

Diplodontus despiciens....... |

Laminipes sp., undesc........ |

Neumania sp., undet......... | |

| —

: | a

Protozoa were rare in species but abundant in individuals.

Both Volvox spermatosphara and Volvox perglobator were extreme-

ly numerous until the middle of August. Endorina elegans and Pan-

dorina norum were only moderately abundant. All are widely

distributed forms, found in ponds and ditches both large and small.

The two species of Volvox are purely American and only recently

described (Powers, 1907 and 1908). Volvox perglobator is found

in all parts of the United States. Volvox spermatosphara has, as

yet, only been found in the central and western states. I find no

data on the Volvocaceae west of the Rocky Mountains.

Stentor pyrifornus was described by Johnson (1893) as a new

species. He found it only in one body of water, Lake Quinsiga-

mond, Worcester, Massachusetts. The form found on the Pacific

coast agrees with the above very closely in all points which John-

son describes. In my specimens, however, the frontal field is some-

what narrower than the diameter of the body just below (plate v1,

MICRO-FAUNA OF AN OREGON POND 79

figs. 2 and 3). A few individuals were found showing the new

peristome forming as is typical for S$. coeruleus. This was not

described by him. The macronuclei were typically two, but one

specimen was found with only one and another with three. The

macronucleus (pl. v1, fig. 4) varies in diameter from 40u to 45y

and is surrounded by a large but variable number of micronuclei, as

Johnson describes for S. igneus. The body is 255 wide and 357p

long. Stentor pyriformis was very abundant during July and Aug-

ust and especially so during the former month. It was found in

equal abundance in the summer of 1906.

Hydra viridis was present in moderate abundance in a few

collections and isolated specimens were found at other times. No

attempt was made to collect Hydra and it was probably the acci-

dental scraping of the stems of water plants that caused its pres-—

ence at all. This species is widely distributed, but has not been re-

corded before from the northwest.

The worms were of very moderate numbers. Dero limosa is

very widely distributed. It is reported from the Philippines, Eng-

land, Africa and North America in bodies of fresh water. Dory-

laimus attenuatus is found in wet earth and ooze in Germany. No

account of it was found for America. Trilobus pellucidus is found

in mud, slime, and ooze at the bottom of ponds, streams and ditches

in Germany and England. None of these forms have been recorded

from any northwestern locality.

Rotifiers were quite abundant in all collections. Fourteen spe-

cies were found and all were fairly abundant in individuals except

Furcularia longiseta. This was only found in small numbers on

one day. Of the forms found (see table) all, except Anuraea hy-

pelasma, are widely distributed, being found all over the world in

all kinds of bodies of water. They are especially abundant in ponds

of the type under consideration, where there is a great deal of veg-

etation. In addition to the species given in the table there was a

large soit rotifer, probably belonging to the Notommatide, which

was too badly contracted for identification. This was quite abun-

dant. There were also a number of specimens of a form belonging

to the Philodinide, but these were also too badly contracted for

identification.

80 ELDA R. WALKER

The Copepoda were quite abundant in all collections taken, but

were slightly less so in the latter part of August. All of the spe-

cies found are widely distributed ones. Cyclops prasinus was the

abundant form. It is reported elsewhere as widely distributed but

few in numbers. Here it occurred in very great abundance, far out-

numbering all other Copepoda. It is found in ponds, rivers, ditches,

lakes, and even wayside puddles, and distributed over the eastern

and central United States, but is not found in the collections noted

previously from the far west. Its location as well as its abundance

are unusual. Cyclops albidus and Cyclops serrulatus were second in

abundance. Both are reported as very abundant forms everywhere.

They are present in nearly all collections from all parts of America.

They are found in temporary ponds, rivers and even wells. They

are among the most common forms from the mountains of the north-

west, even occurring in Crater Lake, Oregon. This is the nearest

place from which this form is mentioned definitely. Cyclops fimbri-

atus is a rare form found throughout the central states from Man-

itoba to Alabama. Cyclops viridis is one of the most common forms

in temporary ponds and other bodies of water in Illinois, Ontario,

Iowa, and northwestern Colorado, as well as in the mountains of

the northwest. It has been found at Spokane, Washington, which,

although east of the Cascade Mountains, belongs to the Columbia

system.

The Cladocera were not abundant either in species or individ-

uals. All of the forms are widely distributed and have been found

in the southern, eastern, northern and middle states. Pleuroxus

denticulatus was the most abundant form. It has been found in the

southern and middle states, but I find no mention of it farther west

than Nebraska. Simocephalus vetulus is abundant over the north-

ern hemisphere in all the more shallow lakes among vegetation, as

well as in some large lakes. It prefers the more shallow, clear cold

water. This form as well as the three following below are also re-

ported (Forbes, 1891) from the Yellowstone Park and from the

Flathead region, Montana. Eurycercus lamellatus has much the dis-

tribution of Simocephalus vetulus, but it is more common in the

deeper, clear waters. Chydorus sphericus is abundant in smaller

lakes over the whole circumpolar land area. The above four forms,

MICRO-FAUNA OF AN OREGON POND 81

found in the Flathead region of Montana, are of especial interest

here because this belongs to the Columbia system of drainage. Alona

intermedia is found in various parts of the United States. Macro-

thrix laticornis undoubtedly represents a new variety. The species

is widely distributed in Europe and is found in Minnesota and New

Mexico. The specimens found differ from the type as described by

Herrick and Turner (1895), as follows: The body is 450y long.

The antennule (pl. v1, fig 6) is much straighter and the distal end

is much enlarged and bears on the end two long spines with four

about half their length and several short ones at their bases. On

the ventral protuberances are several spines instead of hairs as de-

scribed for the type. At the base of the antennule is one lateral

spine. The dorsal side of the shell (pl. vi, fig. 5) is serrate nearly

to the head, the serrations decreasing in size and becoming farther

apart anteriad. The caudal end is marked by a very blunt protuber-

ance. The lower margin is marked as in Macrothrix tenuicornts.

The ventral spines extend nearly to the head.

Ostracoda were very abundant in numbers at all times except

in the last of August. Only one species was found, Cypris ophthal-

mica Jurine. This is a very widely distributed species and occurs in

all Europe, England, Scotland, Java, Paraguay, Celebes Islands,

Zanzibar and in Minnesota, Georgia and Illinois.

Insect larve were found in fair abundance in all collections.

They were common insects whose larve abound in such ponds

everywhere. Specific determinations could not be made because of

the absence of literature on the subject.

The mites were not very abundant, but one to several species

were found in nearly every collection. Limnesia histrionica (Her-

mann), Limnesia undulata (Miiller) and young speciments of some

species of Limnesia were present. These are generally distributed

forms, found in many parts of Europe and America. They prefer

bodies of water with abundant vegetation. Piona reighardi ( Wolcott)

is an American species and has only been reported previously from

the Mississippi valley, Michigan, Illinois, and Louisiana. It is found

in ponds, lakes and sloughs connected with rivers. Some forms of

this genus were found which were too young for specific identifica-

tion. Unionicola crassipes (Miiller) is also a very widely distrib-

82 ELDA R. WALKER

uted form, found in Europe and America. In the United States it

is reported from Michigan, Wisconsin and Nebraska. It is found

in both large and small lakes and in cut-offs connected with rivers.

It is one of the few plankton forms found in these collections.

Arrhenurus prominulus Marshall is a new species described and

named from my specimens by Doctor Ruth Marshall (Marshall,

1908). Arrhenurus krameri Koenike is also an American species.

It has only been reported once before. That was from the head

waters of the Flathead River, in British Columbia (Koenike, 1895).

This is of special interest here because the Flathead River belongs

to the Columbia system. This is the only mite found which had

been recorded before from the west. Young Arrhenuri were also

found. Mideopsis orbicularis (Miller) is another widely distrib-

uted species. It is found in Europe and America in lakes and ponds

with abundant vegetation. An undetermined species of Lebertia

was found. The genus is quite characteristic of the more northern

and more alpine lakes. Diplodontus despiciens (Miiller), young

specimens of Atractides, an undescribed species of Oxus, and one of

Neumania were also present. These are all widely distributed gen-

era. All of the above forms are not only generally distributed in

lakes and ponds with clear, cool water and much vegetation, they

are also the forms commonly found in collections made in the lati-

tude of Michigan and Wisconsin. They are therefore exactly the

forms to be expected in this pond. Except Arrhenurus kramer,

none of these species have been reported before west of the Rocky

Mountains.

The pond is conspicuous for a large number of individuals rep-

resenting a comparatively small number of species. With few ex-

ceptions, the species are those of wide distribution and are those

characteristic of small bodies of water densely filled with vegeta-

tion. They are also the species common among the vegetation near

the shore of larger lakes. Very few true plankton forms are found.

The pond has been reduced in size by drainage and partly filled with

debris until, in dry seasons, it is little more than a swamp, yet the

shore fauna of a lake remains.

MICRO-FAUNA OF AN OREGON POND 83

LITERATURE CITED

Fores, S. A.

1891. A Preliminary Report on the Aquatic Invertebrate Fauna of the

Yellowstone National Park, Wyoming, and of the Flathead Region

of Montana. Bull. U. S. Fish Comm., x1: 207-258.

Herrick, C. L., anpD TuRNER, C. H.

1895. Synopsis of the Entomostraca of Minnesota. Copepoda, Cladocera,

Ostracoda. Geol. and Nat. Hist. Survey of Minnesota, Zool.,

Series II.

JoHNsoN, HERBERT P.

1893. A Contribution to the Morphology and Biology of the Stentors.

Jour. Morph., vir; 468-562, PLS. XXIII-XXVI.

KoENIKE, F.

1895. Nordamerikanische Hydrachniden. Abhandlungen des Naturwiss.

Vereins Bremen, xt: 172-185.

MARSHALL, RUTH.

1908. Arrhenuri of the United States. Trans. Am. Mic. Soc., xxv:

85-140, PLS. VII-XXII.

Powers, J. H.

1907. New Forms of Volvox. Trans. Am. Mic. Soc., xxvit: 123-150.

1908. Further Studies in Volvox, with a Description of three new

Species. Trans. Am. Mic. Soc., xxviii: 141-175, PLS. XXIII-XXVI.

84 ELDA R. WALKER

EXPLANATION OF PLATE VI

Fic. 1. Map of Todd’s Pond.

The dotted area is a swamp in winter and is covered with rushes. |

Areas covered with trees are represented by “xxxx”.

Points marked “B” and “W” are the places from which collections

were made.

Stentor pyriformis; view of the upper surface.

Stentor pyriformis; lower surface of the same specimen shown in

figure 2, viewed as a transparent object. The two drawings were made with-

out moving the specimen. Both of the above show the macronuclei in their

typical location and the green algal cells with which the body is filled.

Fic. 4. Stentor pyriformis; macronucleus, much enlarged, showing the

micronuclei surrounding it.

BiG:

Pig. 3:

Fic. 5. Macrothrix laticornis var.; showing the arrangement of the ser-

rations on the dorsal side of the shell.

Fic. 6. Macrothrix laticornis var.; one antennule, showing the charac-

teristic arrangement of the spines.

PEATE VI

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THE ARRHENURI OF THE UNITED STATES

By RUTH MARSHALL

WITH SIXTEEN PLATES

The Arrhenuri comprise the largest genus of the Hydrachnide,

and one of the most highly differentiated and widely distributed.

The genus is easily recognized by the following characters. The

thick chitinous covering of the body is pierced by large pores. A

circular furrow on the dorsal side marks off an area which is en-

tirely enclosed in the female, in the male usually open onto the pos-

terior extension of the body. The females are oval in form and

much alike; the males are smaller and vary greatly from each other

and from the females. They are characterized by a posterior pro-

longation of the body of varying length and complexity called the

appendix, at the end of which there is often developed an accessory

sex organ, the petiole. The epimera are in three groups. The ca-

pitulum has the form of a shield with a wedge-shaped notch closed

by a membrane. The genital cleft in the female is flanked by two

large semicircular discs, from each of which extends a wing-shaped

area covered with small acetabula. The male genital area, at the

base of the appendix, has narrow plates forming an elliptical plate

for the cleft, from which extend narrow wing-shaped areas. The

palpi are short and stout, ending in a pincer formed by the fifth

segment and a prolongation from the distal end of the fourth. The

legs are relatively short and much alike throughout the genus; the

last three pairs have swimming-hairs, and the fourth segment of

the fourth leg in the male usually has a spur bearing a bunch of

hairs.

The Arrhenuri vary in size from something more than half a

millimeter to over two millimeters in length. The color is usually

dull blue green, less often dull orange red—colors which harmonize

with the environment of these mites, which are shore and bottom

86 RUTH MARSHALL

forms, found in shallow clear waters where chara and other water

plants are abundant.

The species fall into three, or perhaps four, large groups, ac-

cording to the development of the appendix and its structures in

the male; these groups have been designated as subgenera. There is

first the simple type in which the body form approaches that of the

female, the appendix slightly developed and the petiole absent; this

is the type of the subgenus Truncaturus Thor. From this simple

type there appear to have been three divergent lines of development,

two of which have produced “short-tailed” Arrhenuri, and one

“long-tailed.”” The latter, the subgenus Megalurus Thon, has the

appendix enormously elongated, but comparatively simple in struc-

ture and lacking the petiole, at most having no more than a rudi-

ment of it. The subgenus Micrurus Thon has a short appendix with

a deep median incision, over which lies a petiole and certain hyaline

structures. The third type to be differentiated has an appendix of

varying length with prominent posterior lateral corners and a con-

spicuous petiole and accessory structures. This group, which ap-

pears to be the most highly differentiated as well as the largest in

species, is the subgenus Petiolurus of Thon. Wolcott (1905) has

pointed out, however, that this name should be changed to Arrhen-

urus, as the subgenus contains the type of the genus. The four

subgenera are not sharply marked off from each other, there being

intermediate species difficult to assign to any one group.

One hundred and forty-seven species and varieties of the ge-

nus are already recognized, of which nineteen have been described

from the females and immature males only. Besides this number,

there are over fifty species which are designated by Piersig as

doubtful, the work chiefly of earlier writers.

The greater number of species described are European, as

might be expected. As far as work on the Arrhenuri of the rest

of the world has gone, it reveals the fact that each continent has its

own peculiar fauna; the same subgenera are to be recognized in

each case, but the species are distinct. The only exception to this

is the occurrence of five European species in Asia Minor; but this

region may be regarded as biologically a part of Europe. Three new

species have been described for Asia Minor and eight others from

ARRHENURI OF THE UNITED STATES 87

Ceylon, Java, and Dutch New Guinea; twenty-two have been de-

scribed from Africa; nine for South America, and sixteen for North

America.

It will be seen that the Arrhenuri of the Western Hemisphere

have received but little attention. The first work on any North

American water-mites was done by the eminent German hydrach-

nologist, Dr. F. Koenike. In 1895 he published descriptions of four

new Arrhenuri; the material for the study was collected by Dr. J.

B. Tyrrell in British Columbia. In 1903 the author described ten

species, and in 1904 added two more, the material coming from Wis-

consin and Massachusetts. Of these twelve species, all belonging to

the subgenus Megalurus, half were identified with European forms ;

but it was at once pointed out by Dr. Richard Piersig of Germany, a

recognized authority on hydrachnids, that these also were new spe-

cies and they were consequently so christened by him.

Material for the study of the Arrhenuri of the United States

embodied in this paper has been secured from eighteen states, the

largest part from the Middle West. The author’s own collecting

has been confined to Wisconsin, where over forty bodies of water

were visited. To this material valuable collections from New Hamp-

shire and Vermont were added by Mr. George D. Nourse; from

Oregon by Dr. Elda R. Walker; from Louisiana by Mr. E. Foster;

and from Maine by Mr. A. A. Doolittle. A great part of the ma-

terial, however, came from the large collection of water-mites be-

longing to Dr. Robert H. Wolcott, who generously allowed the au-

thor to work out the Arrhenuri. Dr. Wolcott’s personal collections

were made in Michigan, Missouri, Nebraska, Wisconsin, New Hamp-

shire, Colorado, Indiana, Massachusetts, New York,and Ohio, chiefly

in the three first-named states. Other collections were added to these,

the chief of which were those of the State Biological Station at

Havana, Ill. Dr. Wolcott was also fortunate enough to secure ma-

terial from Dr. Henry B. Ward, from Illinois, Michigan, and Ne-

braska; from Dr. C. Dwight Marsh, from Michigan; from Dr. Chas.

Fordyce, from Nebraska; from Mr. E. Foster, from Louisiana ; from

Miss Caroline E. Stringer, from Nebraska; from Mr. J. C. Craw-

ford, from Nebraska; from Mr. O. D. Noble, from Nebraska; from

Mr. A. S. Pease, from Nebraska and Massachusetts; from Mr. A.

88 RUTH MARSHALL

J. Coats, from Wisconsin ; from Prof. J. G. Needham, from Illinois;

from Mr. J. B. Shearer and Dr. R. H. Ward, from Michigan; from

Mr. R. S. Gray, from California; from Mr. E. W. Berry, from New

Jersey; from Mr. R. H. Johnson, from New York; from Prof. S. A.

Forbes, from Yellowstone National Park; and from Dr. Alfred

Dugés, Guanajuato, Mexico.

In this large amount of material over three thousand individ-

uals of the genus have been identified and described. The four sub-

genera were represented by a total of forty-six species, thirty-six

of which are new. The subgenus Megalurus has the largest num-

ber of species, and a majority of all the individuals; in Europe the

subgenus Arrhenurus is the richest in species. It is significant that

Wisconsin, the state in which the largest number of collections were

made, has twenty-six, or over half the number of species present.

To this number might be added three other species previously de-

scribed by the author from this state, but not recurring in the later

collections. It is reasonable to suppose that more extensive collect-

ing will bring to light other species and a wider range for those al-

ready described. The total number now known for the United

States is about half the number found in Europe. The females are

known in ten species.

DESCRIPTIONS OF THE SPECIES

The species are put in their subgenera, and an attempt is made

to arrange them in natural groups.

SUBGENUS TRUNCATURUS

No species were found as undifferentiated as A. knauthet Koen.,

designated by Thor as the type. The five species here grouped to-

gether have each a small simple appendix, but no petiole or acces-

sory hyaline structures. The end of the appendix has a median

incision of varying depth, and consequently this group stands near

the next subgenus; the dorsal median surface is likewise more or

less depressed. The body is regularly oval, with no large humps,

though slight elevations bearing hairs are found where the appen-

dix joins the body. There is a slight bulging over each eye, the

ARRHENURI OF THE UNITED STATES 89

genital wings are broad, and the dorsal enclosed area is large. The

fourth leg of the male bears no spur on the fourth joint. These

Arrhenuri are relatively small. To this group belong:

A. rotundus n. sp.

A. ovalis n. sp.

A. crenellatus n. sp.

A. bicaudatus n. sp.

A. acutus n. sp.

Arrhenurus rotundus n. sp.

Pl. vu, figs. 1-4; pl. 1x, fig. 128

Arrhenurus rotundus has a form approaching that of the Afri-

can species A. pectinatus Koen, with a narrow, slightly indented ap-

pendix. In the depth of this shallow incision lies a papilla. The

greatest width of the body is found just anterior to the enclosed

dorsal area; the latter region is slightly depressed, the sides of the

furrow turning out to end at the genital wings. These areas project

slightly over the body wall; the anterior borders are indistinct. The

third epimera have very narrow inner borders. At the point where

the appendix joins the body there are two low humps, each with a

fine hair. There are four pairs of long hairs on the end of the ap-

pendix, and shorter ones on the body; the arrangement is best shown

by the figures. The palpus closely resembles that of A. manubria-

tor, the fourth joint being somewhat rectangular in outline. The

fourth leg is very simple in structure.

Female. The body is an ellipse, bulging slightly at the an-

terior end. The dorsal enclosed area is ovate, very narrow in front,

with a concave posterior border. A lateral view shows the body

moderately arched with no humps. The third epimera shows the

same peculiarity as in the male, a very narrow inner edge. The

genital plates are large, the wing-shaped areas wide and short and

directed obliquely back. The males measure 0.83 mm. in length and

0.68 mm. in width; the females are 1.09 mm. long and 0.98 mm.

wide. The color varies with age from dull yellow to orange red.

Twenty-seven males and fifty-eight females occurred in eight

collections, all but one being in the same region. These collections

were made in the fall of 1905 in small pools near Lake Winnebago

90 RUTH MARSHALL

and the city of Appleton, Wisconsin. One collection was also made-

June 2, 1906. The other locality, where was found one male, was

the mill-pond at Big Spring, Adams county, Wisconsin, Aug. 17,

1905.

Arrhenurus ovalis n. sp.

Pl. vu, figs. 5-7

Arrhenurus ovalis male is closely related to A. setiger Koen.,

described from Canada; it is larger, however, and the appendix is

slimmer. The body is nearly elliptical; the genital wings, wide at

their origin, run some distance up on the dorsal side. The area in-

side the dorsal line is elevated ; it extends well over on the appendix,

where it is narrow and completely enclosed by the furrow. The low

appendix has a distinct though small median incision with a small

papilla in the depth; the outline shows three low scallops on each

side, the posterior two belonging to the depressed central region.

There are two pairs of fine hairs on each side. The length is 0.93

mm., the width 0.75 mm. The color is dull yellow green.

Two individuals of this new species are known. One was found

in a small pond near Appleton, Wisconsin, September 28, 1905; the

other was collected by Dr. R. H. Wolcott at High Island Harbor,

Michigan, August 18, 1894.

Arrhenurus crenellatus n. sp.

Pl. viii, figs. 11-13

This new species, like the preceding, is closely related to A.

setiger Koen., from which it differs in several details. There is a

slight indentation between the eyes; the area inside of the dorsal

line is depressed and the genital wings are of nearly the same width

throughout. The appendix has the same general form as in the re-

lated species, but the indentation at the end is broader. The dorsal

depressed surface bears a little peg-like structure (A) placed ob-

liquely ; this was not always present, however, in the material ex-

amined. The dorsal shield is oblong and entirely enclosed, the fur-

row running out on the base of the appendix to form two scallops

which enclose each a little hump. The palpus is unusual; the second

segment, which is very large, bears a dense area of blade-like hairs

ARRHENURI OF THE UNITED STATES 91

on the inner surface. The length of this mite is 0.84 mm.; the width

0.68 mm. The color is dull yellowish green.

Thirty-seven individuals occurred in small numbers in collec-

tions from three states, as follows: Wisconsin—Lake Mason, near

Briggsville, August 16, 1905; Buffalo Lake, at Packwaukee, Sep-

tember 5, 1904; Lake Wingra, Madison, August 29, 1905. Michigan

—Susan Lake, Charlevoix, August 21, 1894 (Dr. R. H. Wolcott);

Les Chenaux Ids., August, 1895 (Mr. J. B. Shearer) ; Grand Rapids,

summers of 1895 and 1897 (Dr. R. H. Wolcott). New Hampshire

—Charlestown, summers and falls of 1905 and 1906 (Mr. G. D.

Nourse.

Arrhenurus bicaudatus n. sp.

Pl. vu, figs. 8-10

The body of the male is obovate, narrow in front of the eyes,

where it is slightly indented. The dorsal shield is large, oval and

elevated, the ends of the furrow bending out to lose themselves on

the base of the appendix. The fourth epimera do not have as pro-

nounced posterior angles as do those of the related species. The

genital areas are unusually wide and extend slightly over toward

the dorsal surface. The narrow appendix is well marked off from

the body and is narrow. There is a central deep incision in which

are seen a few very small hairs; this incision divides the appendix

into two large lobes. Anterior to the incision, where the appendix

is low, is a triangular depression (A) ; the two anterior angles have

little rounded projections (B), and in front of these are two little

points (C) and a pair of fine curved hairs. This species is the

smallest in the collection, the length being 0.75 mm. and the width

0.62 mm. The color is dull orange.

Only four individuals of the species are known, but the range

is wide. The collection of Dr. R. H. Wolcott contained three; one

found in Eagle Lake (Winona Lake), Indiana, July 30, 1903, and

two in Lake St. Clair, Michigan, in the summer of 1893. A fourth

was found in the Calcasieu River, Louisiana, by Mr. E. Foster, Sep-

tember 11, 1906.

92 RUTH MARSHALL

Arrhenurus acutus n. sp.

Pl. vit, figs. 14-16

Arrhenurus acutus is a rare form closely resembling A. bicau-

datus in the form of the appendix, but most readily distinguished

from it by its greater size. The body is elliptical; besides the slight

bulging out over each eye there is a slight protuberance anterior and

internal to this, a character so often found in the next subgenus.

The dorsal enclosed area is large and elevated; the furrow bends

inward at the anterior end and is lost on either side when it bends

out to meet the genital wings. The genital areas are very wide and

scarcely reach the lateral surfaces of the body; the anterior borders

are very indistinct. The appendix is divided into two acute lobes,

by a large, deep rounded incision, in the depth of which are a few

very small hairs. On the dorsal depressed surface in front of this

incision is a large triangular hollow (A), the anterior wall of which

is very sharply defined and has at each end a rounded corner (B).

A little forward of this wall is a tiny point (C) on each side, and a

fine curved hair. Thus the structure of the appendix closely resem-

bles that of the related species. On each side are three long hairs

and two short ones. The length of the animal is 0.99 mm., the

width 0.75 mm. The color is dull brownish yellow in the preserved

material.

Only two individuals of the species are known. One was found

in Mirror Lake, at Delton, Wisconsin, August 21, 1905; the other

in the collection of Dr. R. H. Wolcott in Soft Water Lake, Grand

Rapids, Michigan, August 4, 1896.

SuBGENUS MicRrURUS

This subgenus, for which Thon designated A. forpicatus Neu-

man as the type, has a general oval outline, but it shows a tendency

to produce protuberances over the eyes and humps on the dorsal

surface. The dorsal shield is small and completely enclosed. Over

the median incision of the appendix lies a petiole; in the species

studied this springs from a hollow in the center of the depressed

dorsal surface which appears to be a continuation of the dorsal fur-

row, all of this area presenting a complex structure. The subgenus

ARRHENURI OF THE UNITED STATES 93

is represented in the present collections by five new species, all of

which are rare:

. scutulatus n. sp.

. nfundibularis n. sp.

. lyriger n. sp.

. laticaudatus n. sp.

. montifer n. sp.

mwa A A

Arrhenurus scutulatus n. sp.

Pl, vii, figs. 17-19

The body of this new form is nearly oval in outline, slightly

bulging over each eye and moderately arched in the center. The

dorsal shield, which is very small, has an unusual form; it is broad,

indented at the anterior end and constructed abruptly behind to

broaden out again on the appendix into a somewhat four-sided ele-

vated piece. The genital wings have an unusual outline on the

posterior border, each broadening out strongly half way between

the plates and the outer end; they extend well over toward the

dorsal side. The appendix is large and very broad, the two lobes

formed by the median incision simple in outline. In the center of

the depressed dorsal area there is a hollow (A), inverted heart-

shape; the petiole, which springs from this, is a funnel-shaped hya-

line structure (B), directed outward, the ventral surface strength-

ened by a spoon-shaped piece. Three pairs of fine hairs grow on the

appendix, and another pair at the end of the enclosed dorsal area.

There is no spur on the fourth joint of the fourth leg. The length

of this mite is 0.85 mm. and the width 0.67. The color is dull blue

green.

But four individuals are known; these came from as many dif-

ferent localities, as follows: Wisconsin—Green Lake, September 9,

1905; Buffalo Lake, at Packwaukee, September 5, 1904. Michigan

—Susan Lake, Charlevoix, August 21, 1894 (Dr. R. H. Wolcott) ;

Saginaw Bay, August, 1895 (Mr. J. B. Shearer).

Arrhenurus infundibularis n. sp.

Pl. vu, fig. 20; pl. rx, figs. 21, 22

Arrhenurus infundibularis bears a general resemblance to A.

94 RUTH MARSHALL

scutulatus; the appendix, however, is more sharply differentiated.

The oblong body bulges out strongly over each eye, with two median

protuberances. The small dorsal shield is oval, the posterior end

indented ; the enclosing furrow is wide at the rear and runs over

into the hollow of the appendix. In the middle region of the body

are two low humps, seen best in lateral view, marking the greatest

elevation of the body. The third epimera are slightly wider on the

inner edge than are the fourth. The genital wings are narrow and

extend a short distance toward the dorsal side of the body. The

appendix is well marked off from the body; the median incision is

deep and narrow. The hollow (A) on the depressed dorsal face is

large and circular, and connects with the dorsal furrow as already

noted. From the anterior end of this springs the large hyaline petiole

(B), in shape like a funnel, the rim of which has been flattened down

on the upper side. Two fine hairs lie on the face of this structure.

The fourth leg lacks the spur. The length of the body is 0.9 mm.

and the width 0.68 mm. The color is dull blue green.

Six individuals have been found in as many different collec-

tions ; the range, however, is wide. Wisconsin—Portage, the canal,

August 24, 1905. Michigan—Intermediate Lake, Ellsworth, August

9, 1894 (Dr. C. D. Marsh) ; Lamberton Lake, Grand Rapids, July

22, 1898 (Dr. R. H. Wolcott). Missouri—Rocheport, Roby’s Pond,

July, 1904 (Dr. R. H. Wolcott). Oregon—Forest Grove, Todd’s

Pond, August 7, 1906 (Dr. E. R. Walker).

Arrhenurus lyriger n. sp.

Pl, 1x; fig..26;; pl. ‘x, ‘figs..27,,°28

The complex and unusual development of the dorsal face of the

appendix separates this Arrhenurus clearly from any previously de-

scribed species, although it is closely related to the two preceding.

The body is almost circular in outline, except for the bulging over

each eye and the two median protuberances which are close to-

gether. The body rises to a low cone on each side in the middle

region, as seen in a lateral view. The dorsal enclosed area is very

small, somewhat seven-sided. The furrow which encloses it runs

into the depressed part of the appendix. The third epimera are

slightly broader at the inner ends than are the fourth. The genital

ARRHENURI OF THE UNITED STATES 95

wings are rather wide; they extend up on the dorsal body wall. The

moderately large appendix has a deep median incision, partly cov-

ered by the large petiole. The dorsal depressed area of the appen-

dix is broadly triangular, the apex at the end of the dorsal shield.

Near the apex on either side are two prominent rounded projections

(A), extending over onto it, the posterior bearing a long, fine hair.

In the center of the depression lies a very complex hyaline structure

and petiole. The former is a flat, somewhat triangular sac (B),

cleft on the upper side, and seemingly wrapped around the thick

stem of the petiole (C), whose shape at the end is that of an in-

verted harp. Two fine hairs arise near the end of the cleft of the

hyaline structure, and two more from a pair of small humps placed

one on each side of the end of the petiole. Other fine hairs are best

shown by the drawings. There is no spur on the fourth leg. The

length of the entire body is 1.00 mm.; the width is 0.79 mm. The

color is dull blue green.

Six males of the species were found in five collections from

four different states, as follows: Wisconsin—Green Lake, mill

pond, September 9, 1905. Michigan—Grand Rapids, summers 1895

and 1896 (Dr. R. H. Wolcott). Missouri—Rocheport, Roby’s Pond,

July 24, 1904 (Dr. R. H. Wolcott). New Hampshire—Charlestown,

August 20, 1905 (Mr. G. D. Nourse).

Arrhenurus laticaudatus n. sp.

Pl. 1x, figs. 23-25

This Arrhenurus presents a very unusual form because of the

great width of the appendix and the strong arching of the body.

The anterior region bulges out over each eye, with two median pro-

tuberances besides; back of this the body keeps nearly the same

width to the appendix. The enclosed dorsal area is small, the pos-

terior end greatly elevated at two points (A). The body is highest

just outside of the furrow, where it rises into a large cone on each

side. The genital areas are narrow and scarcely projecting. The

first and second epimera are bluntly pointed; the fourth is unusually

small, its inner border being narrower than that of the third. The

appendix is wider than the body; it rises on each lateral border to

form a triangular platform (B). In the center of the depressed

96 RUTH MARSHALL

dorsal area is a broadly oval hollow (C) in which lies a bladder-like

sac of similar shape with a rounded structure on top of it, the latter

bearing on each side a short hair. These structures represent the

petiole. The hollow is connected by a crease (D) with the broad

dorsal furrow. The median incision of the appendix is small and

rounded with a tiny point in the center. Four pairs of hairs, two

long and two short, are placed near the end of the appendix in the

center. The fourth leg bears a spur. The length of the body is

1.14 mm.; the greatest width, at the appendix, is 0.85 mm. The

color is dull yellow.

Eight individuals of the species have been found in the follow-

ing collections: Green Lake, Wisconsin, September 9, 1905; Fisk’s

Lake, near Grand Rapids, Michigan, September 8, 1894; Roby’s

Pond, Rocheport, Missouri, July 24, 1904. The last two collections

were made by Dr. R. H. Wolcott.

Arrhenurus montifer, n. sp.

Pl. x, figs. 29-31

This rare and unusual form is at once recognized by the wide ~

appendix, greatly elevated dorsal shield and peculiar and complex

petiole. The body, slightly obovate, bulges out over the eyes, with

two median protuberances. The dorsal enclosed area is indistinctly

pentagonal; it is greatly elevated above the rest of the body, rising

into a cone which has a little hump on each side, and falling to the

furrow behind, which is here rather wide. The first and second epim-

era are bluntly pointed; the fourth has a narrow inner end and a

pronounced posterior angle. The genital wings are narrow, and

reach barely to the sides of the body. The appendix flares out from

its base to become as broad as the body. The depressed dorsal area

is long and narrow, making the appendix appear composed of two

layers, each of which ends in lateral scallops, the ventral being

longer and narrow. The median incision is very small. The petiole

is large and stout; it arises from a longitudinal crease in the cen-

ter of the appendix. There is an obliquely placed cylindrical base

(A), with a pair of fine anteriorly directed hairs, on the dorsal

face of which lies a heavy rod-like structure (B), also placed ob-

liquely. From the ventral side of the latter extends a thin irregu-

ARRHENURI OF THE UNITED STATES 97

larly shaped vertical transparent plate (C). The entire length of

the animal to the end of the petiole is 1.12 mm.; the width, 0.76 mm.

The color was lost in preserving.

A. montifer is represented by one individual in the collection

from the Illinois State Biological Station, from Havana, Illinois,

April 3, 1895.

SUBGENUS MEGALURUS

In the Arrhenuri of this group the appendix is at least more

than half as long as the body alone. It is comparatively simple,

however, and bears no petiole, unless certain small structures de-

veloped in a few species be taken as the beginnings of such. The

body is ovate or obovate, but does not usually show very pronounced

elevations. The fourth leg is relatively long and its spur is usually

well developed. A comparison of the described species as shown by

published drawings, together with a careful study of the species in

the present collection, brings out the fact that three types are pres-

ent, though not sharply defined. These three types or series will be

designated for convenience by the first three letters of the alphabet.

In series A the appendix does not attain its greatest length, and the

end is decidedly narrower than the base. Series B shows a tendency

to widen the posterior end of the appendix, while the body may de-

velop small elevations, and the dorsal shield is depressed. In series

C the appendix keeps much the same width throughout, but attains

its greatest length; there is a tendency toward the development of

humps on the dorsal side and an elaboration of the end. The nine-

teen species here represented will be grouped as follows:

Series A SERIES B Series C

. birget Mar.

. mamullanus n. sp.

. solifer n. sp.

. scutuliformis n. sp.

. pseudocylindratus

Piers.

. capillatus n. sp.

. manubriator Mar.

. marshalli Pier. . krameri Koen.

. megalurus Mar. . rectangularis n. sp.

A. pseudocaudatus Piers.

. parallelatus Mar. A. senucircularis Piers.

. prominulus n. sp.

DSA AA

ms AAA AA AA A

expansus Nn. sp. . longicaudatus n. sp.

. cornicularis n. sp.

. apetiolata Piers.

Arrhenurus birget Mar.

1903. A. birgei Marshall, Trans. Wis. Acad., x1v:158-159, pls. 16-

17, fig. 10.

1904. A. birgei Marshall, Trans. Wis. Acad., x1v :520-521.

98 RUTH MARSHALL

A. birgei appears to be the least differentiated member of the

subgenus, the appendix being simple and relatively short. It is like-

wise the smallest member of the group in these collections.

The species was described from individuals found in Wiscon-

sin and Massachusetts; it has now been found in six more states.

In addition to those already reported, collections from the seven dif-

ferent states have given one hundred and fourteen individuals:

Wisconsin—Buffalo Lake, at Endeavor, August 24, 1905; Buffalo

Lake, at Packwaukee, September 5, 1904; Underwood’s Pond, Mon-

tello, September 6, 1904; Green Lake, September 9, 1905; Lake

Wingra, Madison, August 29, 1905; Peaslee’s Pool, Lake Spooner,

August 3, 1906. Illinois—Havana, April and August, 1895 (coll.

from State Biol. Sta.). Michigan—Grand Rapids, summer of 1895

(Dr. R. H. Wolcott); Kawkawlin R., August, 1895 (Mr. J. B.

Shearer). Missouri—Columbia, artificial lakes, summer of 1901

(Dr. R. H. Wolcott) ; Rocheport, ponds, June and July, 1904 (Dr.

R. H. Wolcott). Nebraska—Bellevue, Wiley’s Pond, September 4,

1897 (Dr. H. B. Ward) ; Omaha, summer of 1903 (Dr. R. H. Wol-

cott); South Bend, pond, September 1, 1897 (Dr. H. B. Ward) ;

Springfield, Niobrara River, June 23, 1902 (Mr. J. C. Crawford).

Indiana—Kosciusko county, lakes, July 31, 1903 (Dr. R. H. Wol-

cott). Louisiana—New Orleans, pond in Audubon Park, August

and October, 1901; October, 1903, and October, 1904 (Mr. E. Fos-

ter) ; Lake Charles, September 12, 1906 (Mr. E. Foster) ; Slidell,

October 19, 1901 (Mr. E. Foster).

One male was found colored rusty red instead of the usual

blue green. This was from Peaslee’s Pool, near Lake Spooner, Wis-

consin, where other individuals having the same peculiarity have al-

ready been reported (1904).

Arrhenurus mamillanus n. sp.

Pl x, figs. 32-34); pit ser, fies 35

A, mamillanus has a relatively short appendix, which narrows

to a point at the end, where it bears several small transparent mem-

branes. This character, together with the general form of the body,

places the new species near A. membranator Thor. The anterior

end of the body bows out; the large oval slightly elevated dorsal

ARRHENURI OF THE UNITED STATES 99

shield shows no humps. The enclosing furrow runs far down on

the appendix. The fourth epimera are very broad, and the two

posterior groups are placed very close together. The narrow geni-

tal areas slightly overlap the ventral surface. The appendix is very

broad at the base, cylindrical in the middle, and ends in a three-

lobed platform. On each lateral scallop is a small transparent sac-

like structure (A). The extreme end of the middle scallop has a

blunt point (B), which a lateral view shows to be likewise a small

membrane. Two other tiny membranes (C) are made out by care-

ful search near the low humps on the dorsal surface. There are

four pairs of long, fine hairs on the end of the appendix, and two

pairs of very short ones on the dorsal surface. A. mamillanus 1s

1.03 mm. long and 0.62 mm. wide. The color is yellowish brown in

preserved specimens.

Only one individual of the species has been found. This was

collected by Mr. G. D. Nourse, July 29, 1906, at Charlestown, N. H.

Arrhenurus solifer n. sp.

Pl. x1, figs. 36-38

A. solifer resembles A. morrtsoni Mar. in the form of the body

and appendix. The body is very broad where the slightly elevated

dorsal shield begins; there is a slight bulging over each eye, and

one on each side between the epimera and the genital field. The

dorsal furrow opens ventrally on the base of the appendix. Epimera

and genital field are much the same as in the related species; the

latter projects so as to be seen from the dorsal side. The appendix

is narrow at the base, becomes very wide and arched in the anterior

third, and then much constricted in the posterior third. Here it

rounds out and is slightly scalloped. At the point where the con-

striction begins may be seen a small pair of elevations (A). There

are four pairs of short hairs near the end, and two more pairs on the

dorsal surface. The color in preserved specimens is brownish green.

The entire length is 1.23 mm.; the greatest width, 0.76 mm.

Four individuals of this species were found by Mr. G. D.

Nourse at Charlestown, New Hampshire, one on August 20, 1905,

and three more on October 29 following.

100 RUTH MARSHALL

Arrhenurus scutuliformis n. sp.

Pl. x1, figs. 39-42

A. scutuliformis male resembles A. pseudoconicus Piers., one in-

dividual of which has been found in Wisconsin, and the European

form A. conicus Piers. It differs from the former, to which it is

more closely related, in being larger and relatively less stout. The

body is orbicular, bowed out anteriorly; the dorsal enclosed area

has the same form and is well arched, the end of the furrow being

- on the base of the appendix. The first and second spimera are wide

and have blunt outer ends. The genital wings are short and broad,

slightly over-reaching the ventral surface; a few short hairs grow

on the posterior margin. The appendix is slightly constricted near

the center, and a lateral view shows that the dorsal surface is not

uniformly arched but has two low humps, the anterior one the

higher. The extreme end is much narrowed and low, with a slight

indentation in the center. Just in front of the posterior constriction

is a pair of small elevations with a fine hair in front of each. Four

pairs of longer hairs grow from the depressed end of the appendix.

The body measures 1.4 mm. in length and 0.75 mm. in width. The

color of the preserved material is dull yellowish brown.

A. scutuliformis fem. is ovate, the region between the eyes much

narrowed and slightly bulging; there are slight posterior lateral an-

gles on the body. The anterior epimera resemble those of the male,

being broad and very blunt. The inner borders of the fourth, how-

ever, are much rounded off. The capitulum in both sexes is long

and lacks the usual central wedge-shaped incision; its typical shield-

shaped form has suggested the specific name. The genital wings are

long and oval, much constricted where they join the genital plates,

and placed obliquely. The length of the body is 1.23 mm. ; the width,

islam.

Three individuals of this species are known, all of them from

the collections of Dr. R. H. Wolcott from Michigan. One male and

one female were taken, August 18, 1893, at New Baltimore; and

one male, in the summer of 1895, at Grand Rapids.

ARRHENURI OF THE UNITED STATES 101

Arrhenurus pseudocylindratus Piers.

Pl. xvi, fig. 80

1903. A. cylindratus Marshall. Trans. Wis. Acad., x1v:156-157, pl.

17, fig. 8.

1904. A. pseudocylindratus Piersig. Zool. Cent., x1:210.

A. pseudocylindratus is not closely related to the preceding spe-

cies, but it has an appendix narrowed at the end. The structures

here, however, and the slightly elevated central region, are like those

of Series C. The body is unusually long and the projections over the

eyes are pronounced. The palpi are stout; the second joint bears

several bristles, and the fifth, which is cleft, has a forked hair at its

base.

This is one of the rarer Arrhenuri, but it is now known in six

states. Since first reported, fourteen other individuals have been

found in eleven collections, as follows: Wisconsin—Underwood’s

Pond, Montello, September 6, 1904; Lake Mason, Briggs-

ville, August 16, 1905; Benoit Lake, Burnette county, Au-

gust 15, 1906. Michigan—Grand Rapids, summer of 1895 (Dr.

R. H. Wolcott) ; Susan Lake, Charlevoix, August 21, 1894 (Dr. R.

H. Wolcott). Indiana—Kosciusko county lakes, July 31, 1903 (Dr.

R. H. Wolcott). New Hampshire—Ammonoosuc Lake, August 22,

1900 (Dr. R. H. Wolcott) ; Charlestown, September 17, 1905 (Mr.

G. D. Nourse). Louisiana—Slidell, August 18 and October 19,

1901 (Mr. E. Foster). Wyoming—Yellowstone Park, August 8,

1890 (Prof. S. A. Forbes).

Arrhenurus capillatus n. sp.

Pl. x11, figs. 43-45

A. capillatus, with its relatively short appendix, appears to stand

at the head of a line of development. The body is almost orbicular

except for a moderate bulging out in the anterior region. The dor-

sal furrow runs over on the base of the appendix, where the ends

join. The region outside of the shield is arched. The first and sec-

ond epimera have moderately sharp points; the third has a narrow

inner border. The genital wings are narrow, the borders indistinct,

and they do not form the usual rolls at the sides. A peculiar fea-

102 RUTH MARSHALL

ture, which has suggested the specific name, is the fringe of hairs on

the posterior border of the area. The appendix is one of the short-

est in the subgenus; it is rather stout, slightly broadest in the cen-

ter, where it is moderately arched. A pair of low humps (A), close

together, lie just back of the central arch, each bearing a fine hair.

The posterior end is low and slightly indented; here are found four

pairs of short hairs and a pair of tiny bristles on the dorsal surface.

The color in preserved material is yellowish brown. The average

length is 1.1 mm.; the width, 0.72 mm.

Only two individuals are known; these were collected by Mr.

R. S. Gray in the outlet of Laguna de la Merced, San Francisco,

California, October 15, 1905.

Arrhenurus manubriator Mar.

Pl. x11, figs. 46, 47

1903. A. manubriator Marshall. Trans. Wis. Acad., xv1:151-152,

pls. 15-17, fig. 3.

A. manubriator is closely related to A. capillatus; the appendix,

however, is relatively longer, and both body and appendix show a

slightly greater development of humps.

A. manubriator fem. is now figured and described for the first

time. The body is obovate; the anterior region projects, the pos-

terior end shows slight side corners, between which the body bows

out. The enclosed dorsal area is ovate, the posterior border slightly

bent in. The ventral plates are much like those of the male, the

first and second very wide at the outer ends. The most character-

istic feature is the genital area; the semicircular plates closing the

opening are unusually large, and the genital areas extending from

them short and wide. The body has numerous hairs, which are

mostly fine ones. The palpi are like those of the male. The length

is 0.9 mm.; the width, 0.82 mm.

The species is one of the commonest; the localities where it

has been found in the United States are here given, together with

one collection from Mexico. By far the largest number were taken

in Michigan. It is noticeably absent from Missouri and Nebraska,

although material was examined from several different localities.

ARRHENURI OF THE UNITED STATES 103

Over three hundred individuals of the two sexes were found, in

twenty-two collections. Wisconsin—Lake Winnebago, at Oshkosh,

August, 1897 (Dr. R. H. Wolcott) ; Lake Mason, Briggsville, Aug-

ust 16, 1905; Buffalo Lake, at Endeavor, August 24, 1905; Lake

Wingra, Madison, August 1, 1904; Lake Mendota Bay, Madison,

September, 1905; canal at Portage, August 24, 1905; Lake Spooner,

Washburn county, August 8, 1906. Illinois—Fox Lake, September

17, 1894 (Dr. H. B. Ward). Michigan—Charlevoix, July, 1894 (Dr.

R. H. Wolcott) ; Charlevoix, Susan Lake, August 21, 1894 (Dr. R.

H. Wolcott) ; Black Lake, Emmett county, September 3, 1894 (Dr.

R. H. Ward) ; Grand Rapids, summers of 1895 and 1900 (Dr. R. H.

Wolcott) ; Saginaw Bay, 1895 (Mr. J. B. Shearer) ; Lake St. Clair,

summer of 1893 (Dr. R. H. Wolcott). Indiana—Shoe Lake, Kos-

ciusko county, July 31, 1903 (Dr. R. H. Wolcott). Massachusetts

—Falmouth, Shiverick Pond, August 14, 1900 (Dr. R. H. Wolcott).

Mexico—Guanajuato, 1900 (Mr. Alfred Dugés).

Arrhenurus marshalli Pier.

Pl. xm, figs. 48, 49

1903. A. globator Marshall. Trans. Wis. Acad., x1v:148-150, fig. 1.

1904. A. marshalli Piersig. Zool. Cent, x1:210.

This Arrhenurus shows an elongation of the body and a greater

development of humps upon it than do the preceding species, while

the end of the appendix is high and much broadened out at the end.

Each palpus bears a large bunch of small bristles on the inner sur-

face of the second joint; the fifth claw-like joint is cleft. The palpi

are alike in the two sexes except that the female’s is slightly larger.

This has been found to hold true in all the Arrhenuri examined, and

the principle is of great value in identifying the females.

The female of this species shows some variation in form, as has

already been noted for the male (1903). The posterior lateral humps

are often not strongly developed. It may be distinguished from

other female Arrhenuri which it closely resembles by the general

obovate outline of the body, the region of the eyes being narrowed ;

by the development of the body humps, and by the shape of the

104 RUTH MARSHALL

genital areas, the inner posterior borders of which drop down ab-

ruptly from the genital lips. One individual was orange red with

red legs instead of the usual blue green color. A male was also

found of the same color.

A. marshalli is the commonest and most widely distributed

Arrhenurus in this country. It is now known in ten states, over

eight hundred individuals of the two sexes being found, in some

eighty collections.

Wisconsin—Pools at Appleton; Buffalo Lake, Endeavor; Buf-

falo Lake, and Underwood’s Pond, Montello; Buffalo Lake, Pack-

waukee ; North Park lagoon, Oshkosh; Green Lake and mill-pond at

Green Lake; Lake Wingra and Lake Monona, Madison; pool at

McFarland; pool at Poynette; Mirror Lake, Delton; Lemonweir

River, Mauston; Lake Spooner, Washburn county; Lake Mason,

Briggsville ; and Benoit Lake, Burnette county ; during the months of

the summers and falls of 1904, 1905, and 1906; Lake Winnebago,

Oshkosh, August, 1897 (Dr. R. H. Wolcott). Illinois—Galesburg,

September, 1895 (Prof. J. G. Needham); Havana, summers and

falls of 1894, 1895, and 1896 (coll. from State Biol. Sta.). Michigan

—Lakes near Grand Rapids, summers of 1895, 1896, 1898, and 1899

(Dr. R. H. Wolcott) ; Kawkawlin River, August, 1895 (Mr. J. B.

Shearer) ; Susan Lake, Charlevoix, August 21, 1894 (Dr. R. H.

Wolcott) ; Black Lake, Emmett county, September 3, 1894 (Dr. R.

H. Ward); New Baltimore, Lake St. Clair, August 18, 1893 (Dr.

R. H. Wolcott). Missouri—Columbia, artificial lakes, summers of

1901 and 1904 (Dr. R. H. Wolcott) ; Rocheport, ponds, June and

July, 1904 (Dr. R. H. Wolcott). Nebraska—Cherry county, ponds,

June 9, 1903 (Dr. R. H. Wolcott) ; Ft. Robinson, pond, August 22,

1906 (Dr. R. H. Wolcott) ; Omaha, pond, summer, 1903 (Dr. R. H.

Wolcott) ; Springfield, Niobrara River, June 23, 1902 (Mr. J. C.

Crawford). New Hampshire—Charlestown, ponds, August and

October, 1905 and 1906 (Mr. G. D. Nourse). Indiana—Eagle Lake

and other lakes in Kosciusko county, July 30 and 31, 1903 (Dr. R.

H. Wolcott). Louisiana—New Orleans, Audubon Park, August

and October, 1901 (Mr. E. Foster). New York—La Salle, Little

Niagara River, August 22, 1904 (Dr. R. H. Wolcott). New Jersey

—Passaic, June 19, 1902 (Mr. E. W. Berry).

ARRHENURI OF THE UNITED STATES 105

Arrhenurus megalurus Mar.

Pl. x11, figs. 50-52

1903. A. megalurus Marshall. Trans. Wis. Acad., x1v:150-151,

pls. 14, 15, fig. 2.

A. megalurus male is closely related to A. marshalli; the elon-

gation of the body, the development of the humps, and the widening

and differentiation of the end of the appendix have gone still fur-

ther.

The variations in the form of A. megalurus male, particularly

in the size and sharpness of the humps in front of the eyes, in the

development of the scallops at the posterior end of the appendix,

and in the size and shape of the dorsal hump of the appendix, al-

ready noted (1903) in collection from Wisconsin and Massachu-

setts, are found in the larger amount of material now at hand. This

variation does not appear to be due entirely to the age of the indi-

viduals. In collections from two localities a striking variation in

color from the normal blue green was found—a variation, however,

which has been found in several other species of Arrhenuri. One

male from Charlestown, New Hampshire, had the markings on the

body bright brick red, while all of the species (four males in all)

collected from Dock’s Lake, Washburn county, Wisconsin, and

from a small pool near by, were entirely of this color.

A. megalurus fem., which is now known, has a striking form

which easily distinguishes it from other females of the genus. The

sharp bumps on the body which are so characteristic a feature of

the male are found to a degree unusual in the female. The anterior

part of the body projects slightly and is concave between the eyes,

as in the male; and here there is also the same pair of conical pro-

jections over the eyes, sometimes longer and more abrupt than are

figured. Just outside of the dorsal enclosed area, at the anterior end,

are two large, stout humps, farther back than the corresponding

pair in the male. Two smaller ones lie posterior to these, on the

side. But the most prominent feature of the body is the unusual

development of the posterior end. A pair of large, stout humps

project out from the posterior lateral corners of the body. Behind

these the body is usually strongly bowed out, and here are two more

106 RUTH MARSHALL

pairs of conical humps, one dorsal and one ventral. All of the

humps have hairs. The epimera and genital wings are almost iden-

tical with those of A. marshalli. A. megalurus female measures

0.96 mm. in length and 0.85 mm. in width. The palpi are like those

of the male, and resemble those of A. marshalli.

The statement made earlier by the author (1903) that this fe-

male was hardly to be distinguished from A. marshall female is

thus shown to be inaccurate.

This mite is one of the commonest in the subgenus of the same

name, being represented in fifty collections from ten different states.

In all, one hundred and eighty-three individuals of the two sexes

were found. The localities where collections were made are given.

Wisconsin—Lake Mason, Briggsville; Brown’s Pond, near

Briggsville; Buffalo Lake, at Endeavor; Buffalo Lake, at Pack-

waukee; Buffalo Lake and Underwood’s Pond, Montello; Green

Lake and mill-pond, Green Lake; Lake Wingra, Madison; Mirror

Lake, Delton; Lemonweir River, Mauston; Lake Spooner, Dock’s

Pond and pool near Dock’s Pond, Washburn county; Benoit Lake,

Burnette county; various dates during June, July, August and Sep-

tember of the years 1904, 1905 and 1906. Illinois—Havana, Octo-

ber 10 and 11, 1894 (coll. from State Biol. Sta.) ; Fox Lake, Sep-

tember 17, 1894 (Dr. H. B. Ward). Michigan—West Twin Lake

and Susan Lake, Charlevoix, August, 1894 (Dr. R. H. Wolcott) ;

Intermediate Lake, Ellsworth, August 9, 1894 (Prof. C. D. Marsh) ;

various lakes near Grand Rapids, summers 1895, 1896, 1897, 1898,

and 1900 (Dr. R. H. Wolcott). Missouri—Columbia, artificial

ponds, summer of 1901 (Dr. R. H. Wolcott) ; Rocheport, ponds,

summers of 1901 and 1904 (Dr. R. H. Wolcott). Nebraska—

Omaha, September 1, 1903 (Dr. R. H. Wolcott) ; Pilger Lake, Pil-

ger, August 2, 1900 (Miss C. A. Stringer). Maine—Sebago Lake,

August 1, 1907 (Mr. A. A. Doolittle). Louisiana—New Orleans,

September 28, 1906 (Mr. E. Foster). New Hampshire—Charles-

town, summers and falls of 1905 and 1906 (Mr. G. D. Nourse).

Indiana—Eagle Lake (Winona Lake), and other lakes in Kosciusko

county, July 30 and 31, 1903 (Dr. R. H. Wolcott). New Jersey—

Passaic, April 20, 1902 (Mr. E. W. Berry).

ARRHENURI OF THE UNITED STATES 107

Arrhenurus parallelatus Mar.

1903. <A. parallelatus Marshall. Trans. Wis. Acad., x1v:154-155,

pls. 16-18, fig. 6.

The position of this Arrhenurus is uncertain. The shape of the

body is like that of some members of Series C, but the end of the

appendix is not developed in the same way. A striking feature is

the rounded incision at the end in which lies a tiny stalk, perhaps

representing a petiole, characters which are found in the subgenus

Micrurus,

A. parallelatus is not a common species. Described from in-

dividuals collected in Massachusetts, it is now known from three

other states. Twenty-two individuals were found.

Wisconsin—Underwood’s Pond, Montello, September 6, 1904;

Buffalo Lake, at Packwaukee, September 5, 1904; Buffalo Lake, at

Endeavor, August 24, 1905; Lake Spooner, Washburn county,

August 31, 1906. Michigan—Grand Rapids, summer of 1895 (Dr.

R. H. Wolcott) ; Les Chenaux Islands, August, 1905 (Mr. J. B.

Shearer). Massachusetts—Shiverick Pond, Falmouth, August 14,

1900 (Dr. R. H. Wolcott) ; Cranberry Pond, Woods Hole, August

13, 1900 (Dr. R. H. Wolcott). New Hampshire—Charlestown, Oc-

tober 11, 1906 (Mr. G. D. Nourse).

Arrhenurus expansus n. sp.

Pl. x1, figs. 53-55

The most striking characteristic of this species is the very broad

fan-shaped posterior end of the appendix. The body, which is oval,

projects slightly in the region of the eyes. On the ventral side be-

hind the epimera there is a pair of prominent humps, so prominent

that they are seen from the dorsal side also. The genital area is

very small. The fourth epimera have rather sharp posterior angles.

The ends of the dorsal furrow join ventrally at the base of the

appendix. The body has a low hump on either side outside of the

dorsal shield.

The very long appendix broadens gradually a third of the dis-

tance from the base. Just anterior to the center is a slight enlarge-

ment and a lateral view shows a small hump in this region on the

108 RUTH MARSHALL

dorsal side. The broad end has prominent low rounded corners,

between which it is rounded out without indentations. There are

three pairs of long hairs here, with two shorter pairs on the dorsal

surface and one on the ventral. The length of the body is 1.25

mm.; the width, 0.65 mm. The color is brownish in preserved spe-

cimens.

Three males of the species were obtained by Mr. E. Foster

from collections in Louisiana in 1901; one was made August 11,

in Audubon Park, New Orleans, and the other two at Slidell,

October 19.

Arrhenurus pseudocaudatus Piers.

1904. A. caudatus Marshall. Wis. Acad., x1v:521-523. pl. 40,

fig. 1.

1905. A. pseudocaudatus Piersig. Zool. Cent., x11 :185.

A. pseudocaudatus appears to stand near the beginning of a

line in which the long appendix of the male reaches its highest

development.

This species was described from one male found in the inlet

of Lake Spooner, in northwestern Wisconsin, and later unfortu-

nately lost. No other individuals were ever found in that place,

although several collections have been made. But in material

from High Island Harbor, Lake Michigan, secured by Dr. R. H.

Wolcott, August 18, 1894, five individuals were found wich appear

to be the same species. In all details of structure there is close

agreement with drawings of the type form; but measurements

show that the Michigan forms are uniformly larger.

Arrhenurus prominulus n. sp.

Pl. x111, figs. 56-60

In general shape of body and appendix and in the structures

on the latter the new species appears to be closely related to

A. pseudocaudatus Piers. The body is obovate, bulging out in the

eye region. The dorsal shield follows the outline of the body. The

fourth epimera are broad, somewhat rectangular, and close together.

The genital area do not overlap the ventral surface. The appendix

is widest in the anterior third, and here it is also thickest. Back

ARRHENURI OF THE UNITED STATES 109

of this point it is constricted and bears a large double hump (A).

At the base of this structure are four low humps, much like the

corresponding structures in the related species. The two outer

(B) have on the inner border a semicircular row of small oblong

structures; the other pair, close together, are smaller rounded

humps (C). The end of the appendix widens slightly and ends in

four scallops, the central longest; between the latter is a very

small papilla. Four pairs of short hairs are found on the sides,

and one dorsal pair. The entire length of the body is 1.15 mm.;

the width, 0.65 mm. The color in preserved speciments is bluish

dull brown with darker markings on the dorsal side.

Female. The color and markings are the same as those of the

male; the general outline of the body is obovate, with a very nar-

row convex region in front of the eyes. The dorsal shield, also,

is very large and obovate. The body is rather flat. The epimera

are much like those of the male. The shape of the genital area

differs somewhat from the usual form and affords a means of dis-

tinguishing this female from other species. The semicircular

plates of the genital openings are large, while the wing-shaped

areas are relatively small and extend almost straight outward with

slightly enlarged ends. The total length of the body is 1.13 mm.;

the greatest width, 0.95 mm.

This species occurs only in collections from Forest Grove,

Oregon, made by Dr. E. R. Walker. The collecting ground was

Todd’s Pond, a large shallow body of water filled with water plants.

Several collections were made in the summer of 1905 which yielded

six males and twenty females; and in the following summer there

were found seven males and six females. It is interesting to note

that only two other species of the genus were found, each repre-

sented by one individual only, together with one unidentified female.

Arrhenurus kramert Koen.

1895. A. kramert Koenike. Abh. Ver. Bremen, x111:182, pl. 1

figs. 16-20.

1901. A. krameri Piersig. Das Thierreich: 85.

This species, described by Dr. F. Koenike from one male found

in the Flathead River, British Columbia, near the international

’

110 RUTH MARSHALL

boundary, has not been reported since. It is, therefore, a matter

of great interest to find this Arrhenurus in a new locality. In col-

lections made by Dr. E. R. Walker at Forest Grove, Oregon, in

the summer of 1905, there was found one male, which by a careful

comparison with Koenike’s figures, appears to be A. kramer.

A. prominulus stands very close to it, but does not show as great a

differentiation of the end of the appendix.

Arrhenurus rectangularis n. sp.

Pl. xiv, figs. 61-63

This species is closely related to A. kramert, A. prominulus,

and the following species. The characteristic form of the appendix

is suggested by the specific name which was first employed by Dr.

R. H. Wolcott. The body is obovate, slightly bulged out over each

eye and in the region of the genital area. The dorsal shield is

ovate, the enclosing furrow ending posteriorly far down on the

base of the appendix. The first and second epimera have blunt

anterior ends. The genital wings are rather broad and scarcely

project over the body wall. The hairs on the dorsal side of the

body, marking the openings of skin glands, are very long. The

appendix is widest in the anterior third, where it is slightly elevated

as in the related species. It ends rather abruptly with very pro-

nounced lateral corners which project out farther than the middle

region; the latter is bowed in and has a papilla in its depth. The

last third of the appendix is highest, having a sharp double hump

(A) with a pair of fine hairs. Posterior to it is a pair of humps

(B), each one close to the body margin, with a semicircular row

of oblong structures of unknown significance, similar to those of

A. prominulus. Between these humps is a pair of smaller ones

(C), on whch are the openings of glands. The entire length

of the body is 1.3 mm.; the greatest width, 0.68 mm. The color in

preserved material is dark bronze green.

Three individuals of this species are known, all of them in the

collection of Dr. R. H. Wolcott. One was found in Lake St. Clair,

Michigan, August 10, 1893; the other two in Cranberry Lake,

Woods Hole, Massachusetts, July 28, 1900.

ARRHENURI OF THE UNITED STATES 111

Arrhenurus semicircularis Piers.

Pl. xiv, fig. 64; pl. xv, fig. 129

1903. A. securiformis Marshall. Trans. Wis. Acad., x1v:152-153,

pl. 18, fig. 4.

1904. A. semicircularis Piersig. Zool. Cent., x1:210

This species closely resembles A. krameri Koen. and the pre-

ceding species, but the appendix is relatively slimmer, and the end,

as seen in the lateral view, is the thickest part and bears a more

conspicuous united dorsal hump. The body is decidedly obovate.

The fourth leg is slender, the last two joints being long, as is

characteristic of the males of the subgenus. The third, fourth and

fifth joints are well supplied with swimming hairs, and the spur

on the fourth is conspicuous. The palpus is stout.

A. semicircularis is seldom met with. It was first described

from Massachusetts; it has since been found in three other states.

Twenty-two individuals were found.

Wisconsin—Underwood’s Pond, Montello, September 6, 1904;

Lake Mason, Briggsville, August 16, 1905. Michigan—Grand Rap-

ids, summer of 1895 (Dr. R. H. Wolcott). Massachusetts—Shiver-

ick Pond, Falmouth, August 14, 1900 (Dr. R. H. Wolcott) ; Cran-

berry Pond, Woods Hole, July 28, August 13, 1900 (Dr. R. H. Wol-

cott). New Hampshire—Charlestown, September 17, October 1,

1905 (Mr. G. D. Nourse).

Arrhenurus longicaudatus n. sp.

Pl. xiv, figs. 65-67

The most prominent feature of this new species is the great

length of the appendix, which is relatively longer than in any other

species of the genus. The specific name which it has suggested

was first employed by Dr. R. H. Wolcott. The conspicuously obvate

body is like that of A. semicircularis Piers; but it is more strongly

arched. The fourth epimera are very broad. The genital wings

project beyond the ventral surface and are very broad. The ap-

pendix is narrowest at the base, but does not vary greatly in width

throughout. The center is broadest and greatly arched. Just pos-

terior to this is a prominent hump; at its base is a shallow furrow

112 RUTH MARSHALL

in which lies a peg-like structure (A). This structure did not ap-

pear in two individuals, which were young. The end of the ap-

pendix, which is low, flares out somewhat; in the slightly indented

center lies a very small papilla. Four pairs of short hairs grow

from the sides. The average length of the entire body is 1.24 mm.;

the width, 0.78 mm. The color of the preserved specimens is brown-

ish orange.

Only three males of A. longicaudatus are known, all from New

Hampshire. Two were found by Dr. Wolcott in Ammonoosuc

Lake near Crawford in the White Mountains, August 22, 1900; and

one by Mr. G. D. Nourse at Charlestown, July 29, 1906.

Arrhenurus cornicularis n. sp.

Pl. xiv, figs. 68-70

This species and the next, A. apetiolata, form an interesting

group by themselves whose relationship is not clear. In this group

belong also the Brazilian species A. corniger Koen. and A. ludificator

Koen., and A. uncatus Daday from Paraguay. All are characterized

by the presence of a large pointed hump on the anterior dorsal sur-

face of the appendix like similar structures common near the base of

the appendix in the subgenus Arrhenurus. It has been shown by

Thon (1900) that these humps in forms like A. newman Piers. cover

large accessory genital glands which in the subgenus Megalurus are

accommodated in the prolonged appendix. It would be an inter-

esting subject for future investigation to determine the disposition

of these glands in this group of New World species which have

both the long appendix and the large humps.

A. cornicularis is a small species, the entire length being 0.93

mm. and the greatest width 0.59 mm. The general form of the

body is almost orbicular with slight projections in front of the

eyes. Epimera and genital area are of the usual form; the anterior

plates do not have the sharp points of the related species. The

appendix is broad at the base and of nearly the same width through-

out, being relatively shorter and stouter than in A. apetiolata. The

sharp abrupt projection (A) already referred to is just anterior

to the center, and here the body is slightly widened. The extreme

end has a slight indentation and bears two pairs of little knobs

ARRHENURI OF THE UNITED STATES 113

(B, C) on the dorsal surface; at the base of the former is a little

point (D). The spur on the fourth leg is small. The color of

A. cornicularts in preserved specimens is dul\ bronze green.

Only two individuals are known. These were collected by

Dr. R. H. Wolcott at Grand Rapids, Michigan, in the summer

of 1895.

Arrhenurus apetiolata Piers.

Pi. xv, fig. 11

1903. A. corniger Marshall. Trans. Wis. Acad., x1v:155-156, pl.

15, fig. 7.

1904. A. apetiolata Piersig. Zool. Cent., x1:210.

When the species was first described (1903), the female was

unknown. It has since been found and a description is now pos-

sible.

A. apetiolata fem. very closely resembles A. marshalli fem., so

closely indeed that the two forms are separated only with great

difficulty. The body is ovate, a trifle broader relatively at the pos-

terior end. The small humps above the eyes, at the posterior cor-

ners and near the dorsal line are not as prominent as in the related

form; and the body at the posterior end is simply bowed out, with

no trace of the small elevations which are here often rather well

developed in A. marshalli. The epimera are nearly the same in

form, but the second pair ends in sharper points in A. apettolata.

The point of special difference is in the form of the genital region.

The wing-shaped areas are relatively short ; the upper margin slopes

out and back; and most important of all for purposes of identifica-

tion, it will be noticed that the inner posterior margin runs obliquely

out and back, not straight back, as in A. marshalli. A. apetiolata

fem. measures 0.93 mm. in length and 0.88 mm. in width.

A. apetiolata is fairly abundant and has been found in thirty-

two collections from six states. The total number of individuals

of both sexes found was over three hundred and fifty. The locali-

ties are here given:

Wisconsin—Buffalo Lake at Endeavor, August 24, 1905; Un-

derwood’s Pond, Montello, September 6, 1904; Green Lake and

millsvond, September 9, 1905; Lake Spooner, Washburn county,

August 6, 18, 1906; Benoit Lake, Burnette county, August 15, 1906.

114 RUTE MARSHALL

Illinois—Havana, summers and falls of 1894 and 1895 (coll. from

State Biol. Sta.). Micligan—Grand Rapids, September 30, 1895

(Dr. R. H. Wolcott). Missouri—Columbia, artificial lakes, sum-

mers of 1901 and 1904 (Dr. R. H. Wolcott) ; Rocheport, ponds, sum-

mer of 1904 (Dr, R. H. Wolcott). Nebraska—Omaha, ponds, Sep-

tember 1, 1903 (Dr. R. H. Wolcott) ; Pilger Lake, Pilger, August

2, 1900 (Miss C. E. Stringer) ; Springfield, June 23, 1902 (Mr. J.

C. Crawford). Louisiana—New Orleans, Audubon Park, August

11, 1901 (Mr. E. Foster) ; Slidell, October 19, 1901 (Mr. E. Fos-

ter) ; Lake Charles, September 12, 1906 (Mr. E. Foster).

SUBGENUS ARRHENURUS

In this division of the genus there is found the greatest devel-

opment as seen in the elaborate structures upon the appendix, al-

though the latter is not as long as it is in the preceding subgenus.

The petiole is always a conspicuous feature; it is usually somewhat

club-shaped and grows from the middle of the ventral surface of

the appendix, presenting a great variety of form. The petiole is

believed to be a copulatory organ. At its base on the dorsal side

is usually found a transparent plate of chitin called the “hyaline

appendage.” A pair of curved bristles usually stand on either side

of the appendix. The fourth leg is stouter than in the subgenus

Megalurus and its spur is well developed.

Within the subgenus there are three rather pronounced types.

In one, designated here as Series A, the appendix is very short and

narrow, the posterior lateral angles are small, there are no con-

spicuous humps upon the body, and the hyaline appendage is small

or wanting. In Series B the appendix is well developed with con-

spicuous posterior lateral projections directed outward; near its

base are found a pair of sickle-shaped elevations within the dorsal

furrow. In Series C the appendix tends to elongate, while near

its base lies a pair of large conical bumps.

The new species to be described are grouped as follows:

Series A Series B Series C

A. trifoliatus n. sp. A. pistillatus n. sp. A. amplus n. sp.

A. planus n. sp. A. compactilis, n. sp. A. magnicaudatus n. sp.

A. angustocaudatus n. sp. A. falcicornis n. sp. A. superior n. sp.

A. dentipetiolatus n. sp. A. laticornis n. sp. A. americanus n. sp.

A. reflexus n. sp. A. major n. var.

A. wolcotti n. sp. A, flabellifer n. sp.

A. fissicornis n. sp.

ARRHENURI OF THE UNITED STATES 115

Arrhenurus trifolatus n. sp.

Pl. xv, figs. 72-74

This Arrhenurus does not conform closely to the usual struc-

ture of the subgenus; in some of its characters it shows a more

primitive condition than do any of the following species. The body

is oval, strongly arched in the center, with no pronounced humps.

There is a bulging out in front of each eye and in the posterior lat-

eral region. The dorsal enclosed area is oblong and slightly con-

stricted near the center; the extremities of the furrow end blindly

on the dorsal side of the base of the appendix, reminding one of

the condition found in the subgenus Truncaturus. The last two

groups of epimera are very close together. The appendix is nar-

row, short and low. The posterior lateral regions are simply

rounded. The petiole is very broad at the base, simply a continu-

ation of the appendix; it narrows rapidly to end in a trifoliate piece.

A lateral view shows that it bends sharply to the ventral side. No

hyaline appendage is present. There are an unusual number of

hairs at the end, five pairs in all, and one pair on the dorsal side.

The length of the entire body is 1.02 mm.; the width, 0.74 mm.

The color is dull green.

Five individuals of the species are known from the following

localities: [llinois—Havana, April 3, 1895 (coll. from Ill State Biol.

Sta.). Nebraska—Wayne, September 8, 1899 (Miss C. E. String-

er). Missouri—Rocheport, July 24, 1904 (Dr. R. H. Wolcott).

Louisiana—Audubon Park, New Orleans, August 11, 1901 (Mr. E.

Foster ).

Arrhenurus planus n. sp.

PL xvienios: 79:70

This species is closely related to the European form, A. papil-

lator (Mull.) ; it is smaller, however, and shows clearly some differ-

ences in detail. The appendix and its structure are not well devel-

oped and the body is low and flat; thus it stands among the lowest

of the group. The body is almost orbicular except for the bulging

at the anterior end. The dorsal furrow encloses a very large oval

space, and its ends disappear dorsally at the base of the appendix

116 RUTH MARSHALL

as they do in the preceding species. The genital regions are un-

usually small, the wing-skaped areas ending far short of the edge

of the body. The fourth epimera are broad. The appendix is very

short and low; but the posterior lateral angles are well developed.

The petiole (A) is likewise very small; it is almost hidden by the

overgrowth of the dorsal side of the body. In form it is like the

letter Y inverted ; on either side of it is a stiff hair, but these are not

bent to form “Krummerborsten.” There are also four other pairs

of hairs. No trace of a hyaline appendage was found. The entire

length of the mite is 1.06 mm. and the width is 0.94 mm. The pre-

served specimens were strongly tinged with magenta.

Four males of the species were found by Mr. G. D. Nourse at

Springfield, Vermont, May 21, 1906; and five others at Charlestown,

New Hampshire, in the spring of 1907.

Arrhenurus angustocaudatus n. sp.

Pl. xvi, figs. 77-79

The body is stout and broad with very pronounced outstanding

posterior lateral regions. There are two projections in each eye

_Tegion. The dorsal enclosed area rises back of the center in two

conspicuous cones. The genital wing-shaped areas have an unusual

form; they are broad when they join the genital plates, then bend

rather sharply to run obliquely down and out to the body’s edge

which they overlap. The appendix is short and conspicuously nar-

row. It has well developed posterior lateral angles, each bearing

two hairs. Dorsally near the end are two pairs of little elevations,

the middle ones each with a stiff hair; on the ventral side are two

more pairs, each bearing a hair. The hyaline appendage (A) is

very broad and narrow. The petiole is somewhat peg-shaped; it

has a central thickened piece (B) lying in a trough formed by the

rolled-up sides (C). On either side is a stiff curved hair. The

length of the entire body is 1.3 mm., and the width, 1.1 mm. The

color is dull sage green.

Thirteen males of this species were collected by Professor S.

A. Forbes in a weedy pond near Baronett’s Bridge in Yellowstone

National Park, August 30, 1891.

ARRHENURI OF THE UNITED STATES 117

Arrhenurus dentipetiolatus n. sp.

Pl. xvi, fig. 1; pl. xvii, figs. 82, 83

This rare Arrhenurus has a stout body with very broad out-

standing posterior lateral regions like the preceding species. There

are two projections of the body in the region of each eye, while the

center on each side shows a low cone. The fourth epimera are very

wide. The genital wings are large; this region projects strongly

over the dorsal side. The short stout appendix has rather well

developed posterior lateral angles; each bears two hairs. Between

them the dorsal region grows over the ventral to form a slightly

projecting band (A), on the edge of which is a thin transparent

strip which appears to represent the hyaline appendage (B). The

petiole projects considerably beyond the body; it consists of a stout

cylindrical heavy piece with a transparent conical tooth-like struc-

ture (C) on the end. On each side near the middle of the petiole

there grows out a small curved bristle. There are two pairs of

small humps near the base of the petiole, one dorsal, and a larger

ventral pair, each bearing a hair. The entire length of the body is

1.24 mm.; the width, 1.03 mm. The preserving fluid has destroyed

the color.

The species is known by two individuals collected by Dr. R. H.

Wolcott in Colorado, November 16, 1901, one in a pond on Rush

Creek, east of Laird, the other in a pool by Olive Creek, east of

Wray.

Arrhenurus reflexus n. sp.

Pl. xvi, figs. 84-86

The body is stout, thick, somewhat circular in outline, widest

in the anterior half. There is a slight bulging out in front of each

eye. The posterior part of the body is very well filled out on the

ventral side, so much so that this region with the ends of the nar-

row genital wings (A) shows noticeably in a dorsal view. The

dorsal furrow runs over on the lateral angles of the appendix. In

the posterior part of the enclosed dorsal area are two small but

very abrupt circular humps (B) each with a hair. Back of each

is a small, sharp, curved tooth-like structure (C). The third epi-

mera are narrow, the fourth very wide; near the attachments of

118 RUTH MARSHALL

the fourth legs the latter bear patches of short bristles. The ap-

pendix is very short; it has rounded posterior lateral angles each

bearing two hairs. There is a pair of small elevations on the ven-

tral side, each bearing a long hair. The petiole is very well devel

oped. At its base are two little projections with a pair of long

straight hairs. The stout central part of the petiole curves slightly

upward; toward the end it has a transparent corner piece on each

side (D). Down through the center on the dorsal face runs a

long transparent spoon-shaped structure (E) grooved in the center

and reflexed at the end. Two short forward curving hairs extend

out from the central part near the end. The entire length of the

body is 1.09 mm.; the width, 0.77 mm. The color is blue green.

Six males of this species are known. One occurred in a col-

lection from the mill-pond at the village of Big Spring, Adams

county, Wisconsin, August 17, 1905; the others were collected by

Mr. G. D. Nourse, one at Springfield, Vermont, May 21, 1906,

and four at Charlestown, New Hampshire, July 29 and August

20, 1905.

Arrhenurus wolcotti n. sp.

Pl. xvu1, fig. 87; pl. xv111, figs. 88, 89

This species closely resembles A. berolinensts Protz found in

Germany. It is readily known by its very broad body and very

long, large arid peculiarly formed petiole. The body form is like

that of A. dentipetiolatus, with two projections in each eye region,

very broad rounded-out posterior parts, and an arched central

region. The enclosed dorsal area is elevated; just anterior to the

place where the appendix joins the body there are slight indications

of a pair of humps. The genital wings reach barely to the body

edge. The short broad appendix has rounded projecting lateral

angles. The petiole projects far beyond the appendix. The anterior

half is stout and heavy, ending on each side in a scallop bearing a

curved bristle. Over its dorsal face and extending beyond it is a

thin complex structure ending in two divergent prongs (B).

Through the center runs an elongated bladder-like piece, broad at

the end (A). The two very characteristic structures (B) appear-

ing like prongs in face views are seen to be irregular vertical plate-

like pieces when viewed laterally; they are attached in the ventral

ARRHENURI OF THE UNITED STATES 119

region to a thicker central piece. In these details the new species

differs from A. berolinensis Protz. The hyaline appendage is broad

but short; it is reported as entirely absent in the related form.

There are two small humps anterior to its place of attachment; and

on the ventral side are seen the usual pair of larger humps. In all

there are four pair of hairs extending beyond the body. The entire

length of the body including the petiole is 1.13 mm.; the greatest

width, 0.85 mm. The color is dull green.

Three individuals were found by Dr. R. H. Wolcott, in whose

honor the species is named; two came from a pond on Rush Creek,

east of Laird, Colorado, November 16, 1901; and one from a pond

at Glen, Cherry county, Nebraska, August 25, 1906.

Arrhenurus pistillatus n. sp.

Pl. xvii, figs. 90-92

In this species and the three following there is close resem-

blance in the form of the body and development of the appendix,

a noteworthy feature being the sickle-shaped dorsal humps. The

character of the petiole will most readily separate the species. In

this group A. pistillatus is the simplest, as it is the smallest. The

broadly oval body is slightly bulged out over each eye. The dorsal

enclosed area, constricted at the anterior end, is strongly convex and

bears posteriorly the pair of sickel-shaped elevations, their points

directed forward. Outside of this area, in the center of the body,

there is a slight elevation on each side. Epimera are of the usual

form. The genital fields are narrow, their outer ends showing

like bunches on the side of the dorsal view. The appendix is well

developed but the posterior lateral angles are not conspicuous. Two

pairs of small humps lie at the base of the appendix on the dorsal

side as in the related species; the anterior bear two long hairs, the

posterior are pointed. On the ventral side are two larger, low round-

ed humps, each with two long hairs. The pestle-shaped petiole has

on the end, dorsally, a little rounded bladder (A), very conspicu-

ous in a lateral view; two elongated bladders (B) are attached on

either side of the center ventrally. The curved bristles at the side

of the petiole arise from little bunches. The hyaline appendage is

120 RUTH MARSHALL

large and narrows outward. The length of the entire body is 1.0

mm, and the width is 0.75 mm. The color is dull blue green.

Only one individual of the species is known; this was found

in material collected by Mr. R. S. Gray from a pool in an old quarry

in Alameda county, California, December 10, 1905.

Arrhenurus compactilis n. sp.

Pl. xvitt, figs. 93-95

A. compactilis resembles A. compactus Piers. as well as the

species with which it is here grouped. The body is broadly oval,

slightly projecting over each eye. The enclosed dorsal area is small.

At the point where it narrows into the appendix there stand two

small sickle-shaped humps like those of A. pistillatus. The genital

area has the usual elongated form, and the body here is wider ven-

trally than dorsally. The fourth epimera are broad, and the pos-

terior inner border of each is rounded in. The appendix has well

developed posterior lateral angles which turn strongly outward.

The most characteristic feature of the species, and the one by which

it is most easily recognized, is the petiole. It has the usual pestle

shape, but it bears on the dorsal surface a bladder-like structure

shaped like a leaf (A) ; this is also a prominent structure in a lat-

eral view. The hyaline appendage is wide and narrows to its pos-

terior border. There are the same humps at its base as in A. pis-

tillatus: the ventral pair are large, each with a long hair; the dorsal

ones are small, the anterior pair bearing each a long hair on a

papilla, while the posterior have sharp points directed inward. The

entire length of the body is 1.12 mm.; the width, 0.9 mm. The

color is the usual blue green. A. compactilis is one of the rarer

species; but fifteen males are known, all but two of which were

taken in Wisconsin.

Wisconsin—Oshkosh, inlet of Lake Winnebago, August 1897

(Dr. R. H. Wolcott); Fond du Lac, river shallows, September 7,

1905; Green Lake, mill-pond, September 9, 1905; Lake Spooner,

Washburn county, August, 1906. New Hampshire—Charlestown,

September 17, 1905, October 11, 1906 (Mr. G. D. Nourse).

ARRHENURI OF THE UNITED STATES 121

Arrhenurus falcicornis n. sp.

Pl. x1x, figs. 96-98

This species resembles A. tetracyphus Piers. and the others in

this group. The body is very stout, the general outline oval. It

bulges out near the end where the appendix joins it; and it is still

broad up to the anterior third, when it narrows abruptly to bulge

out over each eye. The ventral part is unusually broad where the

appendix joins the body, so that this region seems to project be-

yond the dorsal side. At the point when the enclosed dorsal area

narrows to run over onto the appendix, there are found a pair of

sickle-shaped humps as in the related species. Except at this point

the body is most elevated near the anterior border of the dorsal

line. The fourth epimera are very broad and the inner posterior

borders are strongly rounded in. The genital area is narrow and

each wing-shaped piece runs out to the body edge. The appendix

has very pronounced posterior lateral angles, directed out. Be-

tween them and extending beyond them is the well developed petiole

with the hyaline appendage (A) at the base. The latter is wide

and its posterior corners are acute. The petiole, of the usual pestle

shape, has three very characteristic bladder-like structures on it.

In the center on its dorsal side is a little appendage somewhat vase-

shaped as seen from above (C), the base having two little points.

On the ventral side of the petiole are two little elongated bladders

(B) attached on either side of the center and projecting beyond it

only at the end. Sometimes these projecting ends appear dorsally

as little points. At the base of the petiole are the small humps

found in other males of this group. The pair on the ventral side

are rather stout and blunt. Two other pairs of tiny elevations are

situated on the dorsal side; the posterior pair have sharp points

directed inward, the anterior bear small hairs.

A. falcicornis is a large mite: the entire length is 1.33 mm.;

the width, 1.02 mm. The color is the usual dull blue green. It is

not common, only twelve individuals being known, all but two of

which occurred in collections from central Wisconsin.

Wisconsin—Goose Pond, near Jorden Lake, Adams county,

summer of 1894; Mirror Lake, Delton, August 21, 1905; Green

123 RUTH MARSHALL

Lake, and mill-pond, September 9, 1905; Lemonweir River, shal-

lows, Mauston, September 3, 1906. Indiana—Eagle Lake (Winona

Lake), July 30, 1903 (Dr. R. H. Wolcott).

Arrhenurus laticornis n. sp.

Pl. x1x, figs. 99-101

A. laticornis closely resembles A. falcicornis but is smaller.

The body is more nearly oval and both body and appendix are rela-

tively longer, while the structure of the appendix, particularly the

petiole, differs in the two species. The body inside of the dorsal

line is arched. The sickle-shaped humps, so conspicuous a feature

of the lateral aspect in all males of the group, are not as well de-

veloped as in A. falcicornis. The fourth epimera are broad but not

as much rounded in on the posterior border as in the related species.

The region of the genital area is broader on the ventral side than on

the dorsal, causing a bulging out over the side walls as in the re-

lated species. There is the same development of humps and hairs

on the appendix, but the ventral pair of humps at the base of the

petiole are greatly enlarged in A. laticornis. The lateral posterior

processes of the appendix are rather more conspicuous. The peti-

ole, pestle-shaped, has a little bladder-like structure (A) on its

dorsal face which is uniformly broad and deeply notched. The

entire length of the body is 1.06 mm.; the width, 0.8 mm. The

color is the usual blue green.

Sixty-four males of this species were found as follows: Wis-

consin—Mirror Lake, Delton, August 21, 1905. Illinois—Ha-

vana, August, September, 1894 and 1895 (coll. from Ill. State Biol.

Sta.) ; Galesburg, fall of 1896 (Dr. R. H. Wolcott). Missouri

—Columbia, artificial ponds, summer of 1901 (Dr. R. H. Wolcott) ;

Rocheport, Roby’s pond, summer of 1904 (Dr. R. H. Wolcott).

Arrhenurus amplus n. sp.

Pl. xx, figs. 102-105

This mite, although very large, has a somewhat simpler struc-

ture than the following species in the same group. In all of them

the posterior lateral angles of the appendix are well developed and

the petioles are of elaborate structure. In outline 4. amplus is

ARRHENURI OF THE UNITED STATES 123

ovate with a slight double bulge over each eye. The dorsal enclosed

area on the body is nearly circular; a lateral view shows this region

arched prominently but there are no humps where the appendix

joins. The ventral side of the body in the region of the genital

area projects strongly beyond the dorsal; the genital wings are nar-

row and reach barely to the body edge. The appendix has nearly

the same width throughout; the well developed posterior corners

flare out slightly. The hyaline appendage (A) is very small,

merely a scallop over the center of the base of the petiole. The

petiole is very large; it has a heavy, somewhat Y-shaped median

ventral piece. Thin side pieces roll up dorsally toward the center.

Ventrally on either side of the base of the petiole are two well

developed humps; dorsally there is a small pair, all provided with

hairs. A large curved hair stands on each side of the petiole. The

palpi are stout; the fourth segment is long, and the second bears

several bristles. The spur on the last leg has a bunch of very long

hairs. The color of this mite is blue green tinged with brick red.

The entire length is 1.5 mm.; the width, 1.1 mm.

Female—The outline is ovate with slight posterior corners.

The dorsal enclosed area is nearly circular. The epimera resemble

those of the male, the first and second having rather blunt points,

the fourth being moderately wide. The size and form of the genital

wings will distinguish this species; they are moderately long, wide,

slightly enlarged at the outer ends; the inner borders are wide and

rounded out to fit around the genital plates, which, however, they

do not appear to touch. The inner posterior borders drop con-

siderably below the plates. The entire length of this female is 1.83

mm. ; the width, 1.57 mm.

This species is known in thirty-one individuals, fourteen of

which are males. All of them were found in five collections made

at Charlestown, New Hampshire, by Mr. G. D. Nourse, from July

to October in 1905 and 1906.

Arrhenurus magnicaudatus n. sp.

Pl. xx, figs. 106-108

This species is easily recognized by its great size, the large

broad appendix, and the form of the petiole. The body has nearly

124 RUTH MARSHALL

the same width throughout, bulging out over each eye. The dorsal

enclosed area is small upon the body; the furrow runs along the

sides of the appendix dorsally to end finally on the ventral side

of the projecting angles. Within this area at the base of the ap-

pendix is a big elevation rising abruptly to a broad blunt apex and

sloping gradually back to the end of the appendix, forming the

greatest elevation of the body. Near the anterior end of the en-

closed dorsal area the body rises on each side to a low cone. The

genital areas are small, not reaching the edge of the body. The

fourth epimera are large and the two posterior groups are close

together. The appendix takes up about one-third of the length of

the body and it likewise is of almost uniform width. At the pos-

terior end it broadens out a little and here are two well developed

posterior lateral angles. The petiole has a heavy wrench-shaped

piece on the ventral side; thinner side pieces roll in dorsally to

enclose a space at the bottom of which lies a little thickened ridge.

A slightly curved bristle lies on either side of the petiole. The

hyaline appendage (A) is small; it is merely a narrow border to

a scallop at the base of the petiole. There are two pairs of small

humps just in front of the hyaline appendage, and a large pair on

the ventral side in the same region bearing each a long hair. The

average length of the body is 1.64 mm.; the width, 0.10 mm. The

color is the usual dull blue green.

Seventeen individuals were present in collections from three

different states as follows: Wisconsin—Goose Pond, near Jorden

Lake, Adams county, summer of 1894; Green Lake and mill-

pond, September 9, 1905; Lake Mason, near Briggsville, August

16, 1905 ; Lake Spooner, Washburn county, August 18, 1906. Michi-

gan—Susan Lake, Charlevoix, August 21, 1894 (Dr. R. H. Wol-

cott); Grand Rapids, July 8, 1899 (Dr. R. H. Wolcott). New

Hampshire—Charlestown, September 17, October 1, 1905 (Mr.

G. D. Nourse).

Arrhenurus superior n. sp.

Pl. xx, figs. 109-111

This Arrhenurus, the largest member of the genus thus far

reported, bears a close resemblance to A. magnicaudatus, from

ARRHENURI OF THE UNITED STATES 125

which it differs most conspicuously in the shape of the big dorsal

hump and in the structure of the petiole. The form of the body

is nearly identical; the lateral view shows, however, some differ-

ences. The body is arched on each side near the beginning of the

dorsal enclosed area, but not as abruptly as in the related form. The

large hump at the base of the appendix is uniformly rounded. The

narrow genital area reaches just to the body edge. The posterior

lateral angles of the appendix are rather more pronounced, while

the three pairs of small humps near the base of the petiole, two

dorsal and one ventral, are not as conspicuous. The petioles are

distinctly different in side view. Surface views show that the ven-

tral heavy wrench-shaped piece has attached to it two large thin

extensions along its length. The hyaline appendage (A) is the

same shape in both forms, but is larger in A. superior. This mite

measures about 2.06 mm. in length and 1.2 mm. in width, with some

variation. The blue green color is strongly tinged with brick red.

Six individuals were found in three collections, in two of which

A. magmicaudatus also occurred.

Wisconsin—Lake Mason, near Briggsville, August 16, 1905.

Michigan—Susan Lake, Charlevoix, August 21, 1894 (Dr. R. H.

Wolcott) ; Saginaw Bay, August, 1895 (Mr. J. B. Shearer).

Arrhenurus flabellifer n. sp.

Pl. xx11, figs. 122-124

In general form A. flabellifer resembles the two following

species but is readily distinguished from them by the large size and

unusual form of the petiole. The oval body with a projection over

each eye is of the usual type in this group of the subgenus. The

dorsal enclosed area is depressed, the body on either side rising

to a low hump. The fourth epimera are very wide, the posterior

angles slight. The genital wings project over on the sides of the

body and the posterior border bears several hairs. At the point

where the body and appendix join is a pair of well developed conical

humps as in related species. The moderately long appendix has

well developed outstanding side corners, each with the customary

two hairs. There are the usual three pairs of small humps at the

base of the petiole; the ventral pair are rather large. The hyaline

126 RUTH MARSHALL

appendage is very narrow and can easily escape attention. The

petiole is very broad at the end; dorsally the sides appear to roll in

toward the center; through the middle is developed a thicker piece

(A), ventral and slightly projecting. The entire length of the body

is 1.05 mm.; the width, 0.73 mm. The color is the usual dull blue

green.

Seventeen individuals were found in collections from three

states as follows: Wisconsin—Buffalo Lake at Packwaukee, Sep-

tember 5, 1904; Buffalo Lake at Endeavor, August 24, 1905; Green

Lake and mill-pond, September 9, 1905. Illinois—Havana, August

14, 1894 (coll. from Ill. State Biol. Sta.).. Missouri—Columbia,

summer of 1901 (Dr. R. H. Wolcott).

Arrhenurus americanus n. sp.

Pl. xxi, figs. 112-117

A. americanus resembles several European members of the

genus, particularly A. cuspidifer Piers. The ovate body is slightly

bulged out over each eye. The dorsal enclosed area is depressed,

the sides of the body on either side rising to a low cone. The furrow

runs far along on the dorsal side to end on the corners of the ap-

pendix. The epimera are of the usual form, the fourth being much

the widest, its posterior inner border much rounded in. The gen-

ital wings are narrow and run up on the sides of the body. At the

base of the appendix lies a pair of large pointed humps directed

forward, each with a hair; these structures are slightly smaller

than in A. flabellifer. The long appendix is widest at the end where

it has very pronounced lateral angles. On the dorsal side near the

end are the two pairs of small humps so often found in the Ar-

rhenuri; the anterior pair are rounded and bear each a hair, while the

posterior have points directed inward. On the ventral side of the

appendix is likewise the customary larger pair of humps each with

a long hair. The petiole is pestle-shaped and rather simple. The

hyaline appendage is large and almost rectangular. A curved bristle

lies on each side of the petiole with a long straight hair outside.

The entire length of the body is 0.92 mm.; the width, 0.64 mm.

The color is greenish, usually strongly tinged with dull red.

ARRHENURI OF THE UNITED STATES 127

The palpi are stout. The second joint has several large bristles ;

the claw-like fifth is cleft. The fourth leg is rather stout, as is true

of other species of the short-tailed Arrhenuri. The fourth joint

much exceeds the others in length and its spur is highly developed ;

the last two joints are conspicuously short.

Female. The body is broadly ovate and shows rather pro-

nounced posterior lateral angles. It is distinguished from other

females of the genus chiefly by the shape of the fourth epimera

and of the genital area. The former are very narrow on the inner

border; and the genital wings, which are close to the last plates,

are very long, of almost uniform width throughout, and slope

obliquely out from the genital plates. The length of the body is

1.12 mm.; the greatest width, 1.0 mm.

This is the second most widely distributed and abundant spe-

cies present in the collections; over four hundred and fifty individ-

uals were found in some fifty collections from six different states.

By far the largest single collection was from North Park Lagoon,

a small pond connected with Lake Winnebago at Oshkosh, Wiscon-

sin, where ninety-two males and one hundred and one females were

found, this number comprising the entire collection of adults except

one. No representatives of the species were found in the numerous

collections from Missouri.

Wisconsin—Buffalo Lake at Montello; Buffalo Lake at Pack-

waukee; Buffalo Lake at Endeavor; North Park Lagoon, Oshkosh ;

Fox River shallows, Appleton; Fox River shallows, Kimberly;

Green Lake and mill-pond at Green Lake; Lake Wingra and Ya-

hara River, Madison; pool near McFarland; Lemonweir River,

Mauston; Lake Spooner, Washburn county; Mirror Lake, Delton;

summers and falls, 1904, 1905, and 1906. Illinois—Havana, Aug-

ust, September, and October, 1894 and 1895 (coll. from Ill. State

Biol. Sta.). Michigan—Grand Rapids, summer of 1895 (Dr. R. H.

Wolcott) ; Intermediate Lake, Ellsworth, August 9, 1894 (Dr. C.

D. Marsh) ; Kawkawlin River, August, 1895 (Mr. J. B. Shearer) ;

26 Lake, Charlevoix, August 6, 1894 (Dr. R. H. Wolcott) ; Susan

Lake, Charlevoix, August 21, 1894 (Dr. R. H. Wolcott) ; Saginaw

Bay, August, 1895 (Mr. J. B. Shearer) ; Les Chenaux Islands, Aug-

ust, 1895 (Mr. J. B. Shearer). Nebraska—Omaha, September 1,

128 RUTH MARSHALL

1903 (Dr. R. H. Wolcott) ; Omaha, Child’s Point, June 6, 1903

(Mr. A. S. Pearse) ; Hackberry Lake, Cherry county, June, 1902

(Dr. R. H. Wolcott) ; Pilger Lake, Pilger, August 2, 1900 (Miss C.

E. Stringer) ; Linwood, August, 1898 (Mr. O. D. Noble) ; Wayne,

August 2, 1899 (Miss C. E. Stringer); St. Michael, October 28,

1899 (Dr. R. H. Wolcott) ; South Bend, September 1, 1897 (Dr.

H. B. Ward) ; South Bend, November 3, 1894 (Dr. R. H. Wolcott) ;

Niobrara River, Springfield, June 29, 1902 (Mr. J. C. Crawford). -

New Hampshire—Charlestown, August 12, 1906 (Mr. G. D.

Nourse). New York—Little Niagara River, La Salle, August 22,

1904 (Dr. R. H. Wolcott).

Arrhenurus americanus var. major n. var.

Pl. xx1, figs. 118-120; pl. xx11, fig. 121

In collections containing A. americanus male there were usu-

ally found also a smaller number of individuals very closely re-

sembling them in structure but conspicuously larger. Ninety-seven

of such males were found. In a few cases more of the larger forms

occurred. than of the smaller; and in some cases, even, the larger

form predominated, while in the Missouri collections only the

larger form was found. These larger individuals measure 1.12

mm. in length and 0.75 mm. in width, and they are blue green with-

out the reddish tinge.

A careful examination of the two forms shows several differ-

ences in structure. The middle region of the body is not quite so

elevated; the double hump at the base of the appendix is lower

and blunter than in the larger form. On the ventral side of the

appendix the small rounded humps near the base of the petiole

are not present. But the most striking differences, aside from size,

are in the hyaline appendage and the petiole. The former is con-

spicuously narrower on the posterior border than at the base; the

petiole has the same shape in both forms, but the larger has on

the dorsal face a bladder-like structure cleft at the top, projecting

above and beyond the end. (Compare figs. 113 and 114 with figs.

119 and 120.)

Examination of over two hundred males of A. americanus

shows a very great uniformity of structure. But about one-third

ARRHENURI OF THE UNITED STATES 129

of the larger form showed a hyaline appendage like that of the

smaller form, that is, sharply rectangular; while a few individuals

had not only this peculiarity but also a corresponding petiole, with-

out the bladder-like ending. Moreover, eleven individuals differed

from the rest in that the lateral projecting corners at the end of

the appendix were less developed and more strongly directed out-

ward. And besides, the hyaline appendage of these individuals was

like that of A. americanus, while the petiole was like that of the

larger form. (See fig. 118.)

It would seem, therefore, that we have here a variable and yet

a distinct type from that of A. americanus. But as it agrees in so

many ways with the latter, and is found so often associated with it,

it has seemed proper to regard it as a variety of the smaller, com-

moner, and less variable species.

Wisconsin—Underwood’s Pond at Montello; Buffalo Lake at

Endeavor; Fox River shallows at Appleton; Fox River shallows

at Kimberly; Green Lake; Lake Mendota, Madison; pool near Mc-

Farland; Lemonweir River at Mauston; Lake Spooner, Washburn

county; Goose Pond near Jorden Lake, Adams county; August,

September, and October, 1894, 1904, 1905, and 1906; Lake Winne-

bago, Oshkosh, August 31, 1897 (Dr. R. H. Wolcott). T[llinois—

Havana, August and September, 1894 and 1895 (coll. from State

Biol. Sta.). Michigan—Grand Rapids, summer of 1895 (Dr. R. H.

Wolcott) ; Grand River at Grand Rapids, July 27, 1898 (Dr. R. H.

Wolcott) ; Kawkawlin River, August, 1895 (Mr. J. B. Shearer) ;

Reed’s Lake, Grand Rapids, June 28, 1899 (Dr. R. H. Wolcott) ;

Saginaw Bay, August, 1895 (Mr. J. B. Shearer). Missouri—Co-

lumbia, artificial lakes, summers of 1901 and 1904 (Dr. R. H. Wol-

cott) ; Rocheport, ponds, summer of 1904 (Dr. R. H. Wolcott). Ne-

braska—Omaha, September 3, 1903 (Dr. R. H. Wolcott) ; Wayne,

August 22, 1899 (Miss C. E. Stringer) ; South Bend, Fish Hatch-

ery, September 1, 1897 (Dr. H. B. Ward) ; Bellevue, Wiley’s Pond,

September 5, 1897 (Dr. H. B. Ward) ; Niobrara River, Springfield,

June 23, 1902 (Mr. J. C. Crawford). New Hampshire—Charles-

town, summers and falls, 1905 and 1906 (Mr. G. D. Nourse). In-

diana—Eagle Lake (Lake Winona), July 30, 1903 (Dr. R. H. Wol-

cott). Louisiana—Slidell, October 19, 1901 (Mr. E. Foster).

130 RUTH MARSHALL

Arrhenurus fissicornis n. sp.

Pl. xx1t, figs. 125-127

A. fissicornis is a large and rare form whose relationship is

uncertain. It is easily recognizable by the big, elaborately developed

petiole. The body is almost circular in outline, with only a slight

projection over each eye. At the point where the appendix joins

it, there is a large conical hump on each side having a double point

(A). The fourth epimera are very wide. The genital wing-shaped

areas are rather narrow but long, and curve up onto the projecting

ventral side in an unusual manner (B). The appendix is stout

and the end, where projecting lateral angles are well developed, al-

most as wide as the body. The hyaline appendage (C) is very

small, not even covering the base of the petiole. The petiole is

large and stout, somewhat resembling that of A. reflexus. The

anterior and middle portions are covered with a heavy integument

with pore-openings as is the body; a small curved hair on either

side marks the limit. The end has two lateral rounded transparent

pieces (D). Through the center dorsally extends a spoon-shaped

piece (E) with four tiny bristles at the end. At the base of the

petiole on the ventral side is a pair of rather large humps, each

with a long bristle. Another stout bristle stands on either side of the

petiole, but these are scarcely bent, as are the hairs usually found in

the same position in other species of the subgenus. The spur of the

fourth segment of the last leg is well developed and the distal end

of the fifth is produced into two chitinous prongs. The length of

the body is 1.27 mm.; the width, 0.86 mm. The color, as usual, is

dull blue green.

But two individuals of the species are known, both from the

collections of Dr. R. H. Wolcott: one came from Reed’s Lake,

Grand Rapids, Michigan, July 23, 1898; the other from Eagle Lake

(Winona Lake), Indiana, July 30, 1903.

Ill

ACKNOWLEDGMENTS

To Dr. Robert H. Wolcott, the American hydrachnologist, the

author is deeply indebted for the use of his private library and col-

ARRHENURI OF THE UNITED STATES 131

lections and for continued advice and assistance. Doctor Henry B.

Ward, head of the Department of Zoology of the University of

Nebraska, by his interest and aid, has made this study of the Ameri-

can Arrhenuri possible. The author wishes to express most sincere

thanks to both.

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ARRHENURI OF THE UNITED STATES 133

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134 RUTH MARSHALL

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eth ; ©) tay

Barer eas 2

are es

Vi pete

Fic.

Fic. 10.

CHAD MP wp

A, triangular depression on the appendix; B, rounded projections;

point.

1: (coe be

Fic. 12.

Fic. 13.

ARRHENURI OF THE UNITED STATES

EXPLANATION OF PLATES

Plate VII

Arrhenurus rotundus, lateral view.

Arrhenurus rotundus, dorsal view.

Arrhenurus rotundus, ventral view.

Arrhenurus rotundus, fem., ventral view.

Arrhenurus ovalis, dorsal view.

Arrhenurus ovalis, ventral view.

Arrhenurus ovalis, lateral view.

Arrhenurus bicaudatus, lateral view.

Arrhenurus bicaudatus, dorsal view.

Arrhenurus bicaudatus, ventral view.

Plate VIII

Arrhenurus crenellatus, lateral view.

Arrhenurus crenellatus, dorsal view.

Arrhenurus crenellatus, ventral view.

A, peg-like structure on the appendix.

Fic. 14.

Fic. 15.

Fic, 15.

Arrhenurus acutus, ventral view.

Arrhenurus acutus, dorsal view.

Arrhenurus acutus, lateral view.

135

A, triangular depression on the appendix; B, rounded projection;

ie. 17.

Fic. 18.

Fic. 19.

C, point.

Arrhenurus scutulatus, dorsal view.

Arrhenurus scutulatus, ventral view.

Arrhenurus scutulatus, lateral view.

A, heart-shaped depression; B, petiole.

Fic. 20.

Arrhenurus infundibularis, dorsal view.

A, depression; B, petiole.

RUTH MARSHALL

Plate IX

Fic. 21. Arrhenurus infundibularis, lateral view.

Fic. 22. Arrhenurus infundibularis, ventral view.

B, petiole.

Fic. 23. Arrhenurus laticaudatus, lateral view.

Fic. 24. Arrhenurus laticaudatus, dorsal view.

Fic. 25. Arrhenurus laticaudatus, ventral view (the specimen tipped

forward).

A, humps on the dorsal shield; B, elevated corners of the appendix;

C, hollow in which lies the petiole; D, crease.

Fic. 26. Arrhenurus lyriger, dorsal view.

A, rounded projections; B, hyaline structure; C, petiole.

Fic. 128. Arrhenurus rotundus, fourth leg.

Plate X

Fic. 27. Arrhenurus lyriger, ventral view.

Fic. 28. Arrhenurus lyriger, lateral view.

C, petiole.

Fic. 29. Arrhenurus montifer, lateral view.

Fic. 30. Arrhenurus montifer, ventral view.

Fic. 31. Arrhenurus montifer, dorsal view.

A, base of petiole; B, obliquely placed rod; C, vertical plate.

Fic. 32. Arrhenurus mamillanus, dorsal view.

Fic. 33. Arrhenurus mamillanus, end of the appendix.

Fic. 34. Arrhenurus mamillanus, lateral view.

A, B, GC, transparent structures.

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ARRHENURI OF THE UNITED STATES 137

Plate XI

Fic. 35. Arrhenurus mamillanus, ventral view.

Fic. 36. Arrhenurus solifer, lateral view.

Fic. 37. Arrhenurus solifer, dorsal view.

Fic. 38. Arrhenurus solifer, epimera and genital field.

A, small elevations.

Fic. 39. Arrhenurus scutuliformis, dorsal view.

Fic. 40. Arrhenurus scutuliformis, lateral view.

Fic. 41. Arrhenurus scutuliformis, mas., epimera and genital field.

Fic. 42. Arrhenurus scutuliformis, fem., epimera and genital field.

Plate XII

Fic. 43. Arrhenurus capillatus, dorsal view.

Fic. 44. Arrhenurus capillatus, ventral view.

Fic. 45. Arrhenurus capillatus, lateral view.

A, humps.

Fic. 46. Arrhenurus manubriator, fem., ventral view.

Fic. 47. Arrhenurus manubriator, fem., right palpus.

Fic. 48. Arrhenurus marshalli, mas., right palpus.

Fic. 49. Arrhenurus marshalli, fem., right palpus.

Fic. 50. Arrhenurus megalurus, fem., ventral view.

Fic. 51. Arrhenurus megalurus, fem., dorsal view.

Fic. 52. Arrhenurus megalurus, fem., right palpus.

Plate XIII

Fic. 53. Arrhenurus expansus, dorsal view.

Fic. 54. Arrhenurus expansus, ventral view.

Fic. 55. Arrhenurus expansus, lateral view.

Fic. 56. Arrhenurus prominulus, ventral view.

Fic. 57. Arrhenurus prominulus, dorsal view.

Fic. 58. Arrhenurus prominulus, lateral view.

Fic. 59. Arrhenurus prominulus, palpus.

Fic. 60. Arrhenurus prominulus, fem., ventral view.

A, B, C, humps.

138 RUTH MARSHALL

Plate XIV

Fic. 61. Arrhenurus rectangularis, ventral view.

Fic. 62. Arrhenurus rectangularis, dorsal view.

Fic. 63. Arrhenurus rectangularis, lateral view.

A, B, C, humps.

Fic. 64. Arrhenurus semicircularis, fourth leg.

Fic. 65. Arrhenurus longicaudatus, dorsal view.

Fic. 66. Arrhenurus longicaudatus, lateral view.

Fic. 66. Arrhenurus longicaudatus, epimera and genital field.

A, peg.

Fic. 68. Arrhenurus cornicularis, dorsal view.

Fic. 69. Arrhenurus cornicularis, lateral view.

Fic. 70. Arrhenurus cornicularis, epimera and genital field.

A, conical projections; B, C, knobs; D, point.

Plate XV

Fic. 71. Arrhenurus apetiolata, fem., ventral view.

Fic. 72. Arrhenurus trifoliatus, dorsal view.

Fic. 73. Arrhenurus trifoliatus, ventral view.

Fic. 74. Arrhenurus trifoliatus, lateral view.

Fic. 75. Arrhenurus planus, ventral view.

Fic. 76. Arrhenurus planus, dorsal view.

A, petiole.

Plate XVI

Fic. 77. Arrhenurus angustocaudatus, dorsal view.

Fic. 78. Arrhenurus angustocaudatus, ventral view.

Fic. 79. Arrhenurus angustocaudatus, lateral view.

A, hyaline appendage; B, thickened portion of the petiole; C, rolled-

up outer portion.

Fic. 80. Arrhenurus pseudocylindratus, palpus.

Fic. 81. Arrhenurus dentipetiolatus, dorsal view.

A, outgrowth from the dorsal surface; B, hyaline appendage; C,

tooth-like structure of the petiole.

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ARRHENURI OF THE UNITED STATES 139

Plate XVII

Fic. 82. Arrhenurus dentipetiolatus, lateral view.

Fic. 83. Arrhenurus dentipetiolatus, ventral view.

A, outgrowth from the dorsal surface; B, hyaline appendage; C,

tooth-like structure of the petiole.

Fic. 84. Arrhenurus reflexus, dorsal view.

Fic. 85. Arrhenurus reflexus, ventral view.

Fic. 86. Arrhenurus reflexus, lateral view.

A, genital wing; B, circular hump; C, tooth-like structure; D,

transparent corner of petiole; E, reflexed central part of the

petiole.

Fic. 87. Arrhenurus wolcotti, ventral view.

Fic. 129. Arrhenurus semicircularis, palpus.

Plate XVIII

Fic. 88. Arrhenurus wolcotti, dorsal view.

Fic. 89. Arrhenurus wolcotti, lateral view.

A, bladder-like structure; B, prongs of the petiole.

Fic. 90. Arrhenurus pistillatus, lateral view.

Fic. 91. Arrhenurus pistillatus, dorsal view.

Fic. 92. Arrhenurus pistillatus, ventral view of the appendix.

A, rounded bladder-like structure; B, elongated bladder-like piece.

Fic. 93. Arrhenurus compactilis, ventral view.

Fic. 94. Arrhenurus compactilis, dorsal view.

Fic. 95. Arrhenurus compactilis, lateral view.

A, leaf-like structure.

Plate XIX

Fic. 96. Arrhenurus falcicornis, dorsal view.

Fic. 97. Arrhenurus falcicornis, lateral view.

Fic. 98. Arrhenurus falcicornis, ventral view.

A, hyaline appendage; B, elongated bladder-like structure; C, vase-

shaped structure.

Fic. 99. Arrhenus laticornis, dorsal view.

Fic. 100. Arrhenurus laticornis, ventral view of the appendix.

Fic. 101. Arrhenurus laticornis, lateral view.

A, bladder-like structure; B, hyaline appendage.

140

Fic. 102.

Fic. 103.

Fic. 104.

Fic. 105.

Arrhenurus

RUTH MARSHALL

Plate XX

amplus, fem., ventral view.

amplus, lateral view..

amplus, ventral view.

amplus, dorsal view.

A, hyaline appendage.

Fic. 106. Arrhenurus magnicaudatus, dorsal view.

Fic. 107. Arrhenurus magnicaudatus, ventral view of the end of the

appendix.

Fic. 108.

Arrhenurus

magnicaudatus, lateral view.

A, hyaline appendage.

Fic. 109.

Arrhenurus

superior, ventral view of the appendix.

Fic. 110. Arrhenurus superior, lateral view.

Fic. 111.

rrhenurus

superior, dorsal view of the end of the appendix.

A, hyaline appendage.

Fic. 132:

Fic. 113.

Fic. 114.

Fic. 115.

Fic. 116.

rotated).

Fic. 117.

Fic. 118.

mediate form.

Fic. 119.

Fic. 120.

Fic. 121.

Fic. 122.

Fic. 123.

Fic. 124.

Arrhenurus

Plate XXI

americanus, ventral view.

americanus, dorsal view.

americanus, lateral view.

americanus, palpus view.

americanus, fourth leg (the last three joints are

americanus, fem., ventral view.

major, dorsal view of the appendix of the inter-

major, dorsal view.

major, lateral view.

Plate XXII

major, ventral view.

Arrhenurus flabellifer, lateral view.

Arrhenurus

flabellifer, dorsal view.

flabellifer, ventral view.

A, thickened central part of the petiole.

Fic. 125.

Arrhenurus

fissicornis, dorsal view.

Fic. 126. Arrhenurus fissicornis, lateral view.

Fic. 127. Arrhenurus fissicornis, ventral view.

A, conical cleft hump; B, genital wing; C, hyaline appendage; D,

transparent corner of the petiole; E, central spoon-shaped piece.

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FURTHER STUDIES IN VOLVOX, WITH DESCRIPTIONS

OF THREE NEW SPECIES

By J. H. POWERS

WITH FOUR PLATES

In the last volume of the TRANsacTIONS the writer published an

article on “New Forms of Volvox,” in which two hitherto unrecog-

nized types were described. Each differed radically from all pre-

viously known species. But as the description of each rested upon

but a single collection of material and as their nearest ally—V olvox

aureus—is known to be a highly variable species, it was deemed

unwarranted to bestow upon them new specific names until further

investigations of material from wider areas had been made to

throw light on possible connecting forms. Such further studies

have now been made. A number of valuable collections of Volvox

have been secured, from points, in some instances, as _ widely

separated as Maine, Washington and Louisiana; while continued

effort in the writer’s own state (Nebraska) resulted during the

past summer in finding the organism in nearly a score of bodies of

water. Many of these were insignificant in size and duration. But,

in the full blaze of Nebraska sunlight, Volvox is able to appear,

multiply, and riot in sexual reproduction in pools of rain water of

scarcely a fortnight’s duration. Indeed, the ecology of the organism

deviates here decidedly from that which has usually been assigned

to it. Upon another occasion the writer may describe this more in

detail. Suffice to say now that the writer’s repeated failure to find

Volvox in previous years has probably been due to seeking for it in

too large bodies of water, e. g., ponds one to two feet deep. Prob-

ably a hundred of these have again been examined during the past

summer without success in a single instance.

The amount of material secured from all sources has been very

considerable. Nearly a hundred bottles stand on my shelves, many

of them containing many cubic centimeters of Volvox alone. A

brief indication of the results thus far obtained may be given in

142 J. H. POWERS

tew words. First, the hope that a form had been discovered in the

process of extensive variation has not been substantiated. Second,

on the contrary, the “first form” described by me in the previous

paper proves to be a new and definite species. Third, as to the

“second form” described, nothing resembling it has again been

found. As above stated, no Volvox has again been met with in

deeper ponds resembling the one from which this form was taken.

Fourth, the surprising fact has developed that at least two other

distinct species exist in our fauna, displaying varied and interesting

characters which separate them distinctly from the accepted Euro-

pean species as well as from those which I have described. Fifth,

no less surprising is the almost total absence, from the material I

have examined, of the European types. In three instances only

have isolated colonies been met with showing feeble sexual repro-

duction of the European type of V. aureus. Not a trace of the

true V. globator in sexual reproduction has yet been found. Buta

beautiful form resembling V. globator in its somatic cells I possess

in copious quantities. It has undoubtedly been mistaken for V.

globator in repeated instances. But measurements and, above all,

the study of its sexual phases at once reveal it as distinct. Indeed,

it is not only a separate species but, again, a surprisingly aberrant

form, extending in yet a new respect the limits of generic characters.

It will be seen at the close of this paper that the genus Volvox

includes diversities, both on the sexual and somatic side, that would

better become a family than a genus. Yet its division into separate

genera is plainly out of the question because of the contradictory

nature of these two classes of equally fundamental characters.

In the remainder of this paper I wish, first, to name, refigure and

complete the description of the “first form” of Volvox described in

my preceding publication; second, to figure and describe in detail a

second interesting species, discussing in connection with it a remark-

able process of invagination and final inversion of young colonies

which it displays more frequently than any other species. Third, I

will briefly describe, without figures, the third new species; adding,

finally, a few words of theoretical discussion.

VOLVOX SPERMATOSPHARA

The first form of Volvox described by me in the above mentioned

FURTHER STUDIES IN VOLVOX 143

paper I now name Volvox spermatosphara, the allusion being to

the “sperm spheres” which constitute its most obvious and inter-

esting character. It is not necessary to re-describe this species in

detail as this has been sufficiently done in the previous paper. The

figures there given also represent many aspects of the species cor-

rectly, although the general views of entire coenobia were so im-

perfect, owing to the inadequate material at my disposal, that I have

bettered and extended them by the figures on plate xx111; while

figures 14 to 22 and also figure 26 on plate xxiv show more

abberrant coenobia and certain special details in regard to repro-

duction. It remains at present to give evidence for the specific

distinctness, to modify one or two partial errors, to confirm several

doubtful points, and to add such new details as the study of the

living material has disclosed.

As to the independence of the species, it proves, in most respects,

to be quite constant. By persistent search, I was able to collect a

large amount of the material during the months of July, August,

and September, from quite a number of distinct pools. I observed

the species both in its appearance and disappearance, although

none of the locations allowed me to follow it for many weeks at a

time. But all phases of its life history were observed time and

again, without noting other deviations than those in size, etc., to be

mentioned later. The same uniformity was noted in collections of

material submitted to me from two or three distant points. The

first of these, for which I am much indebted to Dr. Elda R. Walker,

whose article on “Observations on the Micro-Fauna of an Oregon

Pond” appears in this volume of the TRANSACTIONS, was from

Washington. Although sexual phases were not abundant, sufficient

were present to completely prove its identity with the form here

described. The second was an invaluable and copious collection

of Volvox kindly submitted to me by Prof. R. H. Wolcott, of the

*This remarkable collection contained not only examples of the entire

three species here described, besides two instances of decidedly variant coeno-

bia which may very possibly belong to a fourth species, but it also contained,

in lesser quantity, a wide range of species from other genera of the Volvo-

cine, Among these latter were numerous individuals, in all stages of growth

and vegetative reproduction, of the beautiful organism Pleodorina californica

Shaw. T have also seen this organism in the collections of E. Foster of New

144 J. H. POWERS

University of Nebraska. It was taken in a broad but shallow pond

near Rocheport, Missouri, on July 24, 1904. The water of the pond

was said to be decidedly warm to the touch when the collection was

made. V. spermatosphara comprised more than half the collection,

showing every phase of the organism’s development in countless

numbers. It is from this collection that the additional figures above

mentioned were taken. As above stated the evidence from this

locality also showed the species to be a fairly constant type. Volvox

colonies found in plankton collections kindly loaned by E. Foster of

New Orleans showed the probable presence of the species there,

though the sexual forms were not found. It will thus be seen that

the localities for the species, thus far, are all from western America.

Yet it can scarcely be restricted to this area.

The first point of my previous description that requires modifica-

tion is the matter of size. I there gave the limits of 500 to 1,000 »

for the diameter of the colonies. Although the best estimates I

could make, and certainly not far from correct for the material

then at my disposal, I find they greatly exaggerated the size of the

species as I have since found it. All, indeed, of the material since

examined by me has averaged small. This may no doubt be due

to the circumstances of its growth, viz., little water and much

crowding, but the first and only material seen by me when I wrote

the original description was doubtless quite as exceptional or more

exceptional than the larger quantity examined since. The species

averages small, perhaps smaller than any other in the genus. My

more recent measurements show mature reproductive coenobia of

vegetative and mixed content reaching a common maximum of 600

to 650 ». Purely female colonies reach but about 500 p, while

the smallest showing mature sperm spheres are at times as small

as 150 » or even smaller. The most frequent size of mature

coenobia in the above mentioned larger collection from Missouri

Orleans. Hitherto this Pleodorina has to my knowledge, been recorded only

from California and Illinois. I may also here take occasion to correct an

oversight touching Pleodorina in my last article. I there spoke of observing

transition types between Pleodorina and Eudorina. I should have said Pleo-

dorina illinoiensis. This species and this species only is a variety of Eudorina

elegans.

FURTHER STUDIES IN VOLVOX 145

was not far from 350 » in average diameter. These dimensions

illustrate the extreme variability in size of the species, paralleling

in this respect . aureus which, according to Klein, varies from

about 170 to 850 ». The colonies shown on plate 1 will display

these dimensions approximately to the eye. All are photographed

under the same magnification (about 79 diameters) but it should

be borne in mind that the flattening necessary for photography has

not always been in equal degree, while in any case, some coenobia

crush rather than expand in the process. The relations are therefore

approximate, not exact. All the figures on plate xxl except

figure 3, represent nearly or quite adult colonies, and all are taken

from the same collection of material. The minimum size of free

young colonies also falls far below that found in the material first

described. They frequently escape from the parental coenobia at

100 » or less (fig. 3).

Observation of living material emphasizes another minor though

constant character, viz., the uniformly oval shape of the coenobia

at all ages. In no other species of the genus, if we may trust

descriptions and figures, is this character so strongly developed.

Even before birth it is strongly marked (figs. 1 and 2), while in

V. aureus the young are said to be spherical. This character is

no doubt correlated with the unusually active habit and extraor-

dinary development of polarity which this species manifests. To

these I will return later.

As to the general nature of the various reproductive contents of

the colonies, most of the facts remain as previously described. The

decreased average size of the colonies reduces somewhat the average

number of reproductive bodies occurring, though by no means pro-

portionately. This is especially noticeable in the numerous smaller

female colonies (pl. xxi, figs. 9 and 10). These show a marked

tendency to contain exactly eight ova; but frequently this number

is increased to eleven, twelve, or more; while large female colonies,

like figure 4, although they now and then contain many more,

usually do not do so. The ova or oosperms in tiny colonies like

figure 9 often well nigh fill the entire interior.

Similar to this last phenomenon, but much more striking, is the

frequent overcrowding of parent colonies, both rather large as

146 J. H. POWERS

well as very small ones, with asexual daughter colonies or with

these and sperm spheres (pl. xxiv, figs. 20, 23, and 26). No

such hypertrophied reproductive development has, to my knowl-

edge, been hitherto recorded for the genus. All the figures I have

seen have shown the daughter colonies as fairly roomily situated

within the parent. When first found, I was inclined to attribute the

extraordinary appearance to some unwonted shrinking action of

the preservative. But I have since verified the exact appearances

on living material. As will be seen from the figures, the parent

colony is often stretched by the growing young until it becomes a

me.e taut film assuming any form which the chance aggregation

of the progeny requires. Not only is the gelatinous wall of the

parent colony, normally of considerable thickness in this species,

reduced to attenuation, but even the individual cells of the parent

colony, where the pressure of the growing young is greatest, are

flattened out until their longer, longitudinal axis has become much

shorter than the transverse. An interesting result of this over-

crowding is the frequent crushing of the more mature sperm

spheres when these are present (pl. xxiv, fig. 26). The ordinary

daughter colonies prove very resistent, showing but now and then

a little distortion; but the spheres of sperm platelets succumb at

once to the pressure, another significant proof of their non-inde-

pendent character. As they mature, they are, indeed, most fre-

quently flattened against the inner surface of the colony producing

them. And this, again, I find to be not an artefact, but present in

living material.

Touching this same matter of the independence or dependence

of these cell aggregates which resolve themselves wholly into sperms

within the mother colony, I have made some further observations

upon the living material. It is obvious that the sperm spheres

when made up of mature sperm platelets cannot be independent

locomotive bodies. The cilia of the separate sperm bundles do not

extend freely into the surrounding medium but are quite confined

within the thick, if fragile, envelope of the entire sphere. Somatic

cells are not present whose flagella might serve for common locomo-

tion. However as sexually mature Volvox colonies, even when free,

usually tend to quiescence, we might still suppose these spheres

FURTHER STUDIES IN VOLVOX 147

capable of a free life and of locomotion in their earlier stages. More-

over, I have again verified upon living material the somewhat un-

certain observation first made on the original slides to the effect that

the sperm spheres possess rudimentary cilia in the spermogonia

stage. I was, however, unable to find a single free sperm sphere

among living material rich in sexual colonies. 1 therefore tried the

experiment of freeing them by artificial rupture of the parent. I

chose coenobia carrying both sperm spheres and the ordinary daugh-

ters, as these latter would furnish an obvious control in the results.

In every instance, when thus freed, the ordinary daughter colonies

showed the utmost celerity in locomotion. They seemed to me the

most rapidly moving Volvox aggregates I had ever seen. But

the sperm spheres were powerless to change their location. Those

slightly less mature were quite inert, while a few, evidently in about

the last stages of development preceding the final divisions, showed

an interesting rudimentary power of movement. Without changing

their location they rotated, with a slow continuous movement, about

an axis vertical to the substratum. The movement was so slow as to

be but just perceptible under low magnification. It was continued,

however, as long as I watched them, for perhaps a quarter of an

hour. Unfortunately time did not permit me to experiment upon

the possible development of these artificially freed groups of purely

sexual cells. Such experiments could not fail to be of interest.

The above observations are sufficient, however, to establish the

thoroughly dependent nature of these aggregates. If they are

reduced Volvox colonies, they are, as I have stated in my earlier

article, devoid of the majority of the characters of such. They stand

to the parental colony wholly on the plane of a spermary or anthe-

ridium. I shall return to a few words of a theoretical nature touch-

ing these at the close of the article, showing that quite a different

hypothesis may be made with regard to their origin. I may remark

here, however, that a few new observations have been made upon

the size of these sperm spheres. In my preceding description I

gave the numbers of sperm platelets as 64, 128, or 256, allowance

being made for irregularities due to the occasional degeneration of

cells. I find, however, in the very small coenobia, frequent in many

of my later collections, that the number of platelets in mature sperm

148 J. H. POWERS

spheres is often but 32 (pls. xx111 and xxiv, figs. 11, 12, 13, and 14).

If, then, the final interpretation put upon these bodies makes them

true colonies, we must begin our definition of the genus, not with,

“cells numerous, usually above a thousand, etc.” but “number of cells

various, from 32 to at least 22,000.” However, a word later in this

matter, after the next species has been described, for it, again, has a

related, though less extreme, mode of sperm production, which will

cast considerable light upon the matter of interpretation.

As to the number and size of the individual sperms, there is more

variation in the latter than I at first noted, but they are fairly

constant and I have not made further measurements. The number

in the platelet appears, too, more constant than in any other species

of the genus; but among many thousands observed I have seen

perhaps half a score of cases where the number was 64 instead of

the usual 32. Such is the case with the sperm bundles in the

mature and flattened sperm sphere shown in pl. xxiv, fig. 16. All

the platelets evidently preserve a uniform number in these rare

deviations, just as they invariably do in the ordinary type.

In connection with the formation of sperms, I have sought assidu-

ously for transitional variations between ’. spermatosphara and

V. aureus or V. tertius. These would naturally appear, either as

adult colonies developing at least a few sperm bundles in the

ordinary isolated or scattered fashion; or, second, as young pre-

cocious male colonies midway between sperm spheres and the pre-

cocious progeny which Klein described in the rare variant of V.

aureus and which Meyer found in V. tertius. As to the former, not

an adult colony of VY. spermatosphara has shown so much as one

sperm bundle in the position occupied by them in the two European

species, more especially in V. aureus. It is true that I collected a

few colonies of a small Volvox from a spring-fed pool in the

sand-hill district of western Nebraska, among which I at last dis-

covered one or two sexually mature male coenobia that conformed

wholly to the type of V. aureus. But as none of them showed

definite characters, so far as my investigation could be carried, of

*The maximum is given by Klein for V. globator. It is quite possible

that the last species to be described in this paper, which far exceeds V.

globator in size, may also exceed it in number of cells, etc.

FURTHER STUDIES IN VOLVOX 149

V. spermatosphara, I conclude that these colonies probably were

the true European species. It is the only certain instance of it

that has thus far fallen under my observation.

As to the second possible line of connection, a search of thousands

of coenobia has shown me two, which may be doubtfully regarded

as such connecting types. They were from the Missouri collection

which was so exceedingly rich in varied forms of Volvox and related

genera. But in each instance it appeared rather more probable that

they were pathological colonies than either true variants, or yet

examples of another special species. They were small coenobia,

each showing injury and rather weak somatic cells. But the point

in question was that each bore within it, daughter colonies made up

largely of sperm platelets, but with a few undivided and therefore

possible somatic cells. Figure 24 on plate xxiv represents one of

these daughter colonies, under a magnification of 217 diameters.

The sperms were mature, but will be seen to be exceedingly small,

much smaller than any that I have met with in typical V. spermato-

sphara; and, besides this, there were in every case, but sixteen in a

bundle, a number common in typical V. aureus, but never found

by me in the special species under consideration. These two isolated

instances of possible yet improbable connecting links demand no

more attention until their nature has been demonstrated by the

discovery of more material of a similar type.

There remains to be discussed several points of interest con-

cerning the structure of the individual cells of the coenobia. In

the American Naturalist for 1889, John A. Ryder described a so-

called ‘anterior sense organ” in a form of Volvox identified by him

as Volvox aureus. The fact discovered—as interesting as the desig-

nation was unhappy—related to the development of the stigmas.

Those at the anterior pole proved to be “eight or ten times” as

strongly developed as those at the posterior pole. It is curious

that this interesting discovery has not, to my knowledge, been

made, or even repeated by other observers, European or American.

Possibly the same species may or may not display the character ac-

cording to the environment (character of water ; amount of sunlight)

which it is developed. This was my thought when, upon examining

my first V. spermatosphara, taken from a decidedly muddy pool,

150 J. H. POWERS

I found its stigmas powerfully developed in the cells of the forward

pole of each coenobium, yet regularly decreasing in size, until, a

little posterior to the equator of the colony, they were no longer

present, or at least no longer present as colored bodies. I was

unable to decide whether at the posterior pole, they were really

quite absent, or whether one of a group of small colorless granules

really was the homologue of a pigment body. However, I could

find no coenobium which showed recognizable stigmas at the pos-

terior pole, even when well viewed under immersion lenses. Even

in daughter colonies before birth the differentiation is strongly

marked. The stigmas are visible, even under low magnification, at _

the anterior poles, but in each instance, I failed to follow them,

despite most careful search, beyond a point a little past the equa-

torial plane. These facts point to the pculiarity as most probably

a constant specific character, rather than a local adaptation only, for

the young colonies are by no means regularly so placed in the

parental coenobium that the anterior pole only is illuminated. It is

probable that this extreme differentiation is correlated with the

unusual degree of polarity in this species, with its habit as a very

active swimmer, and with its ability to withstand decidedly muddy

water. Further study must determine whether this peculiarity is

present in other species as well. Unfortunately conclusions cannot

be drawn from preserved material, as the stigmas are not conserved.

Finally, a word concerning the important feature of the cyto-

plasmic strands or processes supposed to extend from cell to cell of

the Volvox colony. Much debated for V. globator and finally

definitely established by Arthur Meyer, they seem also invariably

present in VY. aureus, though delicate, straight and as fine as the

cilia throughout their entire length. In V. tertius, too, Meyer

found them in the younger colonies, but, despite his long research

and elaborate technic upon this particular point, they quite escaped

his observation in the older colonies. Still he doubted their absence.

I have carefully repeated his technic, using the same Zeiss lenses

which he employed. I have also used other and newer reagents

which seemed indicated for the purpose in hand. But in this species

of Volvox I detect not the smallest trace of connectives in colonies

of any age whatever. If this finding is borne out by the observa-

FURTHER STUDIES IN VOLVOX 151

tions of others, it seems gratuitous to assume the presence of such

connection between cells as a generic character of Volvox. I may

add that the reagents employed brought out with much plainness

the minutest features hitherto observed in other species. The dem-

onstration of the finest fibrils which Meyer showed to be the true

connectives uniting the larger processes, cell with cell, in V. globa-

tor, gave me no difficulty. Cilia were plain, even in living daughter

colonies, viewed through the thick matrix of the parent. I think

I should have discovered the connectives easily had they been

present at one-fifth the diameter of the cilia.

As to the structure of the gelatinous sheaths surrounding the

individual cells, | have, following Meyer, made it out fully, using

his reagents on fresh material and also by very prolonged staining

of preserved material in carmalum. I regret that lack of time

prevents my preparing figures of this feature for present publica-

tion. My preparations, however, show the present species closest

in this respect to V. tertius, though distinct from it. Most notice-

ably so in the marked polarity of the sheath. The thickness of the

matrix about the posterior cells is several times that of those at

the anterior pole.

Summing together the more important characters for the species

Volvox spermatosphara, we get the following definition:

Coenobia strongly oval, the sexually mature members from 150

to nearly 1,000 » in diameter, more commonly between 250 and

600 », containing approximately 1,000 to 3,000 cells. Polarity and

activity of the coenobia strongly developed, the single cells show-

ing stigmas decreasing regularly from the anterior toward the

posterior pole. Form of individual cells spherical to oval in side

view, without connecting strands of cytoplasm. Numbers of pri-

mary sex cells approximately the same, whether becoming par-

thenogenetic daughter colonies, ova, or aggregates of male cells

(spermogonia)—from 1 to 25, or perhaps more. Ripe sperms

in bundles of 32, situated, not in the matrix of independent colonies,

but in aggregates within the parental colony—sperm spheres—

each of which contains from 32 to approximately 256 bundles.

152 J. H. POWERS

VOLVOX WEISMANNIA

While studying the beautiful collection of material collected by

Professor Wolcott at Rocheport, Missouri, I met with numerous

coenobia of a Volvox quite unknown to me which at first glance I

thought might be a variant of V. spermatosphara or possibly a link

between it and some other type. A little study, however, showed

it to be wholly distinct from V. spermatosphara and from all other

species hitherto described. It is indeed one of the most unique

species of the genus. Not only does it display several very excep-

tional and yet unvarying characters, but it shows its specific dis-

tinctness still more conclusively by the large number of constant

minor characters which separate it from other described species.

In order to facilitate to the utmost recognition of the species, which

I am especially desirous should be restudied in other developmental

stages and in other localities, I will describe these characters in the

order of their significance and of their obviousness, rather than in

the conventional series.

Perhaps the most striking and certainly the most fundamental

of these characters is the early period at which the primary repro-

ductive cells are set apart as distinct from those which are to become

the vegetative or somatic members of the adult aggregate. The

ten smaller figures in the upper half of plate xxvi make this fairly

plain. All are photographed with a magnification of 217 diameters

and all are from decidedly young, though not equally young,

coenobia. Several are taken just before the time of closure (figs.

50 to 54), while others (figs. 45, 47, 48, 49, and 56) in stages just

subsequent to this event. Perhaps most characteristic and instructive

of all are figures 51 and 52. These are taken from the two opposite

poles of two similar daughters in the same coenobium. Young

coenobia in exactly this stage and showing exactly this appearance

were present by hundreds in the material. It will be seen that the

young colony is still quite unclosed, yet the reproductive cells are

not only distinct and well developed but they have already quvite

lost their connection with the periphery of the colony, lying merely

in delicate contact with the inner ends of the somatic cells which

are now evenly distributed above them. Figure 53 shows, poorly,

an earlier stage, while figures 50 and 54 were intended to show

FURTHER STUDIES IN VOLVOX 153

yet earlier stages, the number of cells still being small, although no

less certainly separable into somatic and reproductive individuals

than in the somewhat later stages. At just what stage in embryonic

development (if this expression is permissible) the separation first

takes place I have not been fully able to ascertain. Despite the

number of coenobia at my disposal I found none showing the exact

stage desired. Evidently, however, the differentiation of the re-

productive cells must begin at about the 64-celled stage; if not,

certainly at the next step. This brings us directly to the conclu-

sion that we have in this simple Volvox aggregate a perfect exam-

ple of the continuity of germ cells. It is in emphasis of this that

I have named the species after the celebrated advocate of this con-

ception.

Correlated, no doubt, with this early setting apart of reproductive

cells is another interesting fact of their almost mathematically

regular arrangement. Despite the fact that their numbers vary,

occasionally running to high limits, yet even most of the variations

are reducible to regular deficiencies of certain definite cells, on

the one hand, or to equally regular additions or multiplications on

the other. The phenomena remind one strikingly of the regularity

of cell arrangement resulting from the segmentation of the ovum in

higher organisms. The commonest numbers of primary sex cells

are eight and ten, the latter predominating. The arrangement of -

the eight cells is as simple as possible (figs. 45 and 48), four, dis-

tributed at equal distances from each other, lie in one plane, wel!

toward the posterior pole of the colony. The final closure of the

young colony is always at this pole between these four cells. Situ-

ated at a considerable distance above these four cells lies the other

four. They are invariably on a plane considerably above the equator

of the colony and are so placed as to alternate with the lower four

(note again figs. 45 and 48), cell for cell. {f do not find an in-

stance where the cells in the upper plane stand directly above those

below, although they soon come to be so far above them that no

reason is discernible for the special alternating arrangement. The

ten-celled grouping repeats exactly that of the eight-celled, with the

addition, however, of two more reproductive cells, in a pair, placed

still nearer the vegetative pole. These cells are very peculiar.

154 J. H. POWERS

To begin with, in the younger colony at least, they seem never to

reach the size of the other cells; they are uniformly smaller, a

difference which is frequently, though not always, maintained

throughout their entire subsequent development. I am unaware of

any other species of Volvox showing thus a regular discrepancy

in the size of its sex cells. Again these two smaller reproductive

cells are arranged in a definite manner with regard to those in the

series just above them; they are not in a position of perfect sym-

metry, or only assume this position later. They lie almost touching

two of the cells in the series just above them. The assumption of

this position is again due, it would seem, to the position of the

quadrangular or diamond-shaped opening of the unclosed colony;

it is so large in nearly all cases as to thrust the smaller pair of

cells out of their natural situation (see fig. 51). These eight and

ten-celled groupings constitute by far the major number in the

material which I have studied, and they also constitute the basis

for all of the deviations. To begin with, the most frequent of these

are due to the omission of one of the reproductive cells in any

ane of the series. Most frequently of all, one of the two smaller

is wanting. But it is surprising to note that neither its mate nor

any of the other cells are thereby shifted in their positions or

altered in number or character. A single member is simply dropped

out. Other deviations consist in interpolations and duplications or

divisions. I have never found a single cell added on an old plane,

making five in one circle, although nothing is commoner in other

species. When ceils are added they are added in pairs. Thus the

two lower cells may rarely become four, all of the same small size

which characterizes the lower two; much more frequently another

pair is introduced, of full size, on a new level, as between the two

main series. Rarely the cells of the lower four are shifted out

of one plane, so that all, except the upper circle, appear plainly

in pairs (fig. 39). Not only may we have these interpolations, but

not infrequently each single cell in all the series has divided, as

shown by the fact that in the young colonies these cells are but

half the size of those in sister coenobia with ordinary counts (fig.

47); the two or four smaller cells at the base of the colony, how-

ever, may or may not divide with the rest. In very rare instances

FURTHER STUDIES IN VOLVOX 155

both interpolation of series and division of members has taken

place, giving surprisingly high numbers, which, so far as my obser-

vation goes, then always become ova.

This introduces another point which interested me much, viz.,

the study of the destination of the different groups of cells. Such

study showed that the definite arrangement applied equally to all.

It is maintained with singular beauty in colonies which develop ova

only, as figures 36 and 38 will show to a considerable extent, al-

though the difficulties of flattening these colonies sufficiently for

photography without distorting the symmetry of arrangement in

their reproductive bodies is discouragingly great. A special ten-

dency is plainly marked for the two or four smaller, posterior cells

to develop into ova when the others do not (figs. 38, 42, and 43),

while the opposite case I have never seen. Yet this latter is the

only invariable rule that may be laid down. Otherwise every pos-

sible combination occurs. Any reproductive cell in any part of the

colony may become either an ovum, a parthenogenetic colony of the

ordinary type, or one of the dwarf male colonies later to be described.

As above stated the regular arrangement tends strongly to be pre-

served throughout development, although it may finally become par-

tially lost and is much less noticeable when the smaller reproductive

bodies develop into large daughter coenobia. Yet figure 57, de-

spite some displacement in the vigorous flattening necessary for

photography, shows a beautifully typical arrangement, the two

minor cells having developed into correspondingly small vegetative

daughters. Rarely is the discrepancy in size so great, though sel-

dom wholly absent. All in all, this definiteness in arrangement,

yet almost entire indeterminateness in development, shows in this,

as in no other species, the almost perfect parity of the different

classes of primary reproductive cells. In no other species is this

parity so great, although as I have suggested in my previous study,

it is strongly indicated in V. spermatosphara. With these species

before us it becomes, I think, impossible to deny the perfect homol-

ogy of all the primary reproductive cells of the genus. And, if so,

Klein’s elaborate argument to the contrary; his refusal to admit

the applicability of the conception of parthenogenesis, which

Biitschli had applied to the asexually developing progeny ; his desire

156 J. H. POWERS

to homologize these asexually reproductive cells with the zoospores

of the algae,—all these contentions fall to the ground. This will

appear still plainer as we proceed, and especially at the close.

Touching next on the ova, we find them marked, in this species,

by considerably greater size than in any other. Unfortunately, in

the entire mass of material, none seemed to have been fertilized

and I cannot state the nature of the oosperm wall. The unfertil-

ized eggs, however, will be seen (figs. 36 and 38), although their

maximum development may stili not have been reached, to be larger

than the zygotes in large female colonies of V. spermatosphara

(fig. 4). I have not measured the very largest observed, but turn-

ing again to my slides, I readily find examples over 90 y» in their

greater diameter. A further peculiarity lies in the fact that the

Ova are not spherical when seen from the side, so to speak. They

are slightly flattened against the wall of the colony (fig. 36) just

as are, in this species, the young colonies. This may be due to the

action of the preservative (formalin), but it probably is not. Un-

fertilized ova of other species are not thus flattened by the reagent.

Finally, the mode of sperm-cell production in V. weismanmia is

also unique, though showing relationship to several other types.

It is by means of dwarf male colonies which ripen their sperms

before they are liberated, and indeed, probably lead but a brief

and ineffectual motile life after they are freed from the parental

organism. The species was but just entering upon a full period

of sexual reproduction at the time the collection was made. An

unlimited amount of material was therefore not at my disposal.

Nevertheless, I examined, in one series, 131 parental coenobia

bearing young male colonies in some easily recognizable stage of

development (plate tv, figs. 46, 55, 57, 58, 59, 60). In all these the

process bore evidence of striking uniformity even in nunor details,

the only exception being in the number of such dwarf colonies formed

in one parental coenobium. This varies from one to ten, as the

figures will show, the difference being largely due to the fact that

usually a part of the primary reproductive cells develop into vege-

tative daughters instead. Figure 57 shows the only instance found

by me in which all of the reproductive cells had given rise to male

progeny. Among the oldest coenobia I did not find the male colo-

FURTHER STUDIES IN VOLVOX 157

nies side by side with ova, but they were so in the somewhat younger

ones. Indeed, as I have already indicated, nearly all combinations

occur. Thus on plate m1, figure 39 shows a colony in which all

the progeny are ordinary vegetative daughters save one of the

lower, the only one present of the lower pair, which is a young male.

In figure 35 one of the lower pair and one of the first four are

dwarf male colonies; in figure 41 one of the upper four only, and in

figure 42 one of the lower four only. I shall return to this matter

of the distribution of reproductive content, giving a series of con-

crete cases, as soon as | have described other phenomena connected

therewith.

The presence of these dwarf male colonies, with accelerated

sperm production, at once suggests a resemblance, if not a rela-

tionship, between this species and the alleged variety of V. aureus

described by Klein,’ as well as to V. tertius. Both of these forms

showed the development of ripe sperms in daughter colonies before

their birth. The resemblance is thus a real one, and a less close

one exists to V. spermatosphara even, the sperm spheres, without

all somatic cells, being, perhaps, comparable to these dwarf male

colonies with but few somatic cells. The differences in detail are,

however, very marked and very constant. Thus, in this species

the dwarf male colonies are not paralleled, as in the European

forms, by dwarf female colonies. Although the egg cells are set

apart never so early in V. wetsmannia, they grow slowly and mature

only in colonies that have long been free. The structure, too, of

these precocious male colonies is unique. Thus the number of

somatic cells, in proportion to the spermogonia, is surprisingly few,

though it by no means varies to especial closeness to the sperm

spheres of V. spermatosphara (see especially pl. rv, fig. 60). The

number of sperms in the bundles is again quite beyond that in any

other Volvox save the unrelated forms of V. globator and its

American ally, viz., 64 and 128. In every instance observed by

me these two sizes occur side by side, and, curiously enough, in

about the same proportions, the larger size always much the most

numerous. Even in the younger colonies with undivided spermo-

*Ludwig Klein: Neue Beitrage zur Kenntniss der Gattung Volvox.

Berichte der Deutschen Botanischen Gesellschaft, vir: 42, etc.

158 J. H. POWERS

gonia, of which I observed a great many, these latter cells were

always of the same two distinct sizes and the larger always much the

more numerous. These facts can, with care, be made out in fig-

ures 58 and 59. Both the position and the shape of the sperm

bundles is also peculiar. They lie well within the colony, leaving

the mere smattering of somatic cells to form the entire peripheral

contents (fig. 60). Even the spermogonia take up this position

some time before their final divisions. The process is just begin-

ning in figure 59 and is carried further in figure 58. In all other

species heretofore described the bundles of sperms are represented

as quite flat in their final form; here they are all of them slightly

concave (fig. 61). The sperms, too, while showing the terminal

cilia and general form of the awreus-type, are more slender than in

any other species except VY. globator.

The above may be considered a fair resumé of the most important

characters of this species so far as they are peculiar and so far

as they could be ascertained from the material at my disposal. One

other character, however, was very evident in the material and was

so new and strange that I at first thought it peculiar. I have, how-

ever, since discovered it occasionally in other species. And in all

it remains the most interesting phenomenon which I have thus far

met with in the genus. I refer to the invagination and, finally,

the complete inversion, or turning inside out, of many of the young

colonies. The process begins at various periods before the closing

of the young colony and is finally completed by a reclosure of the

colony, the surfaces of which are now reversed, at a somewhat

later period. I have seen no allusion to similar observations by

other students of the genus. This fact, together with the bizarre

nature of the phenomenon itself, led me to be very doubtful as to

its cause. The possibility suggested itself that the whole process

might be due to some unwonted effect of the five per cent formalin

used as a preservative. Yet this usually conserves well the gen-

eral shape of colonies, both young and old. In any case I deemed

the process peculiar to V. weismannia, and that if not due to the

action of the formalin, it might well be a pathological result of

growth at too high a temperature. The volvox collection in ques-

tion had been taken in water that “felt warm to the hand.” One

FURTHER STUDIES IN VOLVOX 159

after another, however, each of these conjectures was disproven.

I at last found young colonies of V. spermatosphara which were

also undergoing the process at the time of their preservation.

While, even in my earliest collections of living material, in July, I

succeeded in detecting a few instances, not one of which showed

any other sign of abnormality. Later in the season I met with

them, never in great numbers, but from time to time, in each of

the species which I collected. They occurred, too, in freshly col-

lected material brought from near-by sources with the utmost care.

Last of all, I found the same thing, in at least one instance, in a

colony of the large globator-like species next to be described, which

I took from relatively cold spring water. I judge the phenomenon,

then, to be not wholly abnormal. I regret that I have in no case

been able to retain and watch the development of the inverted colo-

nies. I have only watched them through a few stages of the

process.

An examination of figures 30, 31, and 32 will show how conspicu-

ous are the young colonies of V. weismannia which result from

this inversion. A superficial glance at even unstained material

revealed the astounding presence of the primary sex cells on the

outside of young colonies. Sometimes one, sometimes several,

and, now and then, all of the young daughters in the half-grown

coenobia were in this condition. Unfortunately the oldest coenobia

with still larger daughters did not contain them, and I was accord-

ingly unable to trace their development beyond about the time of

the reclosure of the inverted progeny. I was not able to photograph

the earlier stages of the transformation very clearly, but figure 22,

from V. spermatosphara, and figures 25, 27, 28, 29, and 33, from

V. weismannia, suffice to make apparent most of its stages. The

young colonies, at the beginning of the process, lie with the unclosed

pole directed toward the center of the coenobium. The closed or

anterior pole, lying against the wall of the parent, folds inward,

exactly as in a true case of gastrula formation. When, however,

the invagination is completed (fig. 22), the process does not pause,

the opening at the posterior pole yields to the pressure of the oppo-

site infolding layer; it is stretched, though apparently not torn

asunder, and the anterior pole of the young colony presses quite

160 J. H. POWERS

through the open posterior pole. An intermediate stage in the

process at which it seems to slacken somewhat, and is therefore

quite frequently found, is shown in figures 25 and 32. It looks ex-

actly like a rolled up child’s cap. Soon, however, the crown is

pressed quite through the rim and a form is reached resembling a

wide vase or an old time kettle with flaring rim (fig. 27). This

then rapidly closes, the whole assuming the original form of a

young colony save for indentations or pockets which are soon

formed under the reproductive cells (figs. 30, 31, and 32). The

ova which show but indistinctly in the inverting colonies as fig-

ured, are, in reality, very conspicuous in shifting foci. So loosely

are they held to their points of anchorage during the inversion, it

seemed at first probable that they might frequently be quite dis-

lodged, I was even inclined to interpret certain loosely scattered

cells of appropriate size found in the matrix of the parental coeno-

bium as possibly of this origin. Further observations did not bear

this out, however. The reproductive cells are not loosened during

the inversion; and the scattered cells in the old colony proved to

be chlorophyl-containing flagellate parasites, which penetrate the

matrix from without and there undergo a further development.

The reproductive cells of the young inverted colonies meanwhile,

loosely held at first, rapidly sink into little depressions, well shown

in figure 32. I have seen none that were sunk deeper or gave

signs of re-entering the colony; but, in the material at hand, hun-

dreds of young colonies were present, with the eight and ten repro-

ductive cells respectively, looking, according as seen from one

pole or the other, exactly like figures 31 and 32.

Seeing the abundance of these forms, and noting considerable

variation in the developmental stage at which the process begins, I

sought for instances of invagination in young but completely closed

colonies. I half feared I should find such, and thus unwittingly give

occasion for some younger Haeckelian philosopher to load our deli-

cate Volvox with a yet more ponderous weight of phylogenetic

fancy. Fortunately, however, no such “true gastrulae’ could be

found. Slightly cupped individuals were common enough, but none

deeply invaginated, so we need not on account of this new phenom-

enon dream yet further of ’olvox as the primal metazoon. The real

FURTHER STUDIES IN VOLVOX 161

interest in the process lies in its possibilities as, so to speak, a

natural experiment. The reproductive cells, which have originally

three distinct modes of development, are, in mid-career, suddenly

placed in a new yet evidently not fatal environment. Will this in-

troduce into the already complex life-cycle of this species yet other

and varied links? The single though copious collection of material,

which is all I have of this species, does not even suggest an answer.

I should be much pleased if others should be so fortunate as to

find the complementary phenomena.

Having described the reproductive phenomena of this species,

it may be worth while to indicate briefly the actual contents of sev-

eral coenobia taken almost at random from the slides. This will

substantiate, to some extent, the foregoing statements as well as

supplement the meager representation in the figures.

Number 1. Coenobium with content of 10 bodies; upper row

of four daughters, each with eight reproductive cells, two inverted,

two typical; second row, one typical daughter with eight reproduc-

tive cells, two with high numbers (18 or 20) and one in process of

inversion with eight or ten reproductive cells; two lower members

of the content remaining as true ova.

Number 2. Large coenobium with content of ten daughters ;

the first and second series, with four in each, all typical daughters

with eight reproductive cells each; the two lower members also

daughter coenobia but smaller, one with eight, the other with six-

teen reproductive cells of half the size. No inversion.

Number 3. Smaller coenobium with content of eight bodies ;

all of upper series and two of lower are very young daughters, seem-

ingly all with eight reproductive cells ; remaining two in lower series,

large ova.

Number 4. Content of ten bodies. Upper series: two typical

daughters with 9 and 10 reproductive cells, respectively ; remaining

two, in process of inversion, show 8 or 10 reproductive cells. Second

series: one typical daughter with 18 or 20 reproductive cells; one

with but 6, and two completely inverted with each 8 reproductive

cells. Lower pair of bodies remain as large ova.

Number 5. Coenobium with content of nine daughters. Upper

series: four typical young colonies with 8 reproductive cells each.

162 J. H. POWERS

Second series shows absence of one member, leaving but 3, each of

which bears 8 reproductive cells; one completely inverted. Lower

two members small daughters, both completely inverted, with 8 re-

productive cells each.

Number 6. Coenobium with content of ten daughters. Each

member in the two upper series a typical young colony with 10 re-

productive cells. The lower two, much smaller, with 8 reproductive

cells each, one completely reversed and one in process.

Number 7. Coenobium with content of ten bodies, all ova ex-

cept one of the upper series, which is a young dwarf male colony.

Number 8. Coenobium with content of eight bodies. Upper

series: 2 large ova, one typical and one inverted daughter, each with

& reproductive cells. Lower series: three large ova and one young

dwarf male colony.

Number 9. Coenobium with content of ten bodies. Upper se-

ries: all young daughters with 8 reproductive cells, of which two are

completely inverted, one in mid process and one just beginning.

Second series: likewise. all daughters with 8 reproductive cells each,

of which one is typical and 3 are completely inverted. Lower two

members: one an inverted daughter with 8 reproductive cells, the

other a young dwarf male colony.

Number 10. Coenobium with content of ten bodies. First and

second series each contain 4 typical daughters with 8 reproductive

cells each. Lower two members both ova.

Number 11. Coenobium with content of ten bodies. First and

second series composed of typical daughters with 8 or 10 reproduc-

tive cells each, save one member in upper series which has a much

higher number too difficult to make out. Two lower members are

very small inverted daughters, each with 8 reproductive cells.

Number 12. Coenobium with content of ten bodies, all typical

daughters with 8 reproductive cells each, except the lower pair, one

of which is an ovum and one a young dwarf male colony.

VOLVOX PERGLOBATOR

I little thought to hit upon yet another distinct species of the

genus. I had indeed several years ago seen a few colonies of a

large Volvox closely resembling V. globator save that a very few

FURTHER STUDIES IN VOLVOX 163

female colonies showed a wholly unwonted number of ova or zyg-

otes. This suggested at least marked variation from the European

type. Hardly had I found my first Volvox pool this season, how-

ever, than I hit upon this type, the first time I had ever seen it alive

or in any quantity. It proved to be a most rampant grower in the

writer’s locality (Lincoln, Neb.). After a heavy rain in mid-August

ten days sufficed to produce, multiply and bring to copious reproduc-

tive phases an amount of material which almost thickened the water

in tiny depressions about the border of a weedy pool on low, soggy

ground. When sexual reproduction was at its height the reproduc-

tive colonies, especially the females, lay literally in handfuls at the

bottom of such cavities. Even a cursory examination of such ma-

terial showed it to be specifically distinct from the European V.

globator as this has been described and figured. Moreover, this lat-

ter species has not been considered of a highly variable nature. As

I have not had time to prepare plates of this species, I will in the

present article but note its more obvious specific differences.

To begin with, it grows in this locality to just about double the

size ascribed to V. globator. The maxima given for this latter

species have been from 500 to 800 p» for the vegetative colonies, the

sexual colonies being usually, though it appears not always, smaller.

V. perglobator always exceeds these dimensions in this locality, even

when the conditions of excessively rapid multiplication and over-

crowding should tend to reduce its size as much as possible. All of

my collections showed vegetative colonies beyond 1 mm. in diam-

eter, while 1200 to 1400 » colonies were readily obtainable, and from

a slightly larger pool I measured one old furrowed colony of nearly

1600 » diameter. Even the female colonies from this source were,

many of them, just about 1 mm. in diameter, sometimes exceeding

this in the longer dimension.

The second prominent character separating this species from

V. globator is its dioecious sexuality. Overton says of V. globator

that all observers agree upon its monecious (hemaphrodite) char-

acter. Klein, it is true, in his monograph published simultaneously

with Overton’s, throws some doubt upon this assertion, deeming that

V. globator “may yet be shown to possess more complicated sexual

relations.” This has not, to the writer’s knowledge, yet been shown;

164 J. H. POWERS

but, in any case the classic species must be chiefly hermaphrodite.

Its American congener, on the other hand, in the large masses of

sexually reproductive material that I have examined, shows uni-

form separation of the sexes. Among thousands of coenobia I did,

indeed, find one enormous female colony, loaded with zygotes, which

showed, on one side, a single well developed group of sperms, while

near it a vacant space showed evidence of the escape of another.

But a single instance among thousands indicates an abnormality only.

The sperm masses in this species, seem, further, to always escape

outward, into the surrounding water, never into the interior of the

coenobium. Among material rich in male colonies these groups of

free sperms may readily be found, swimming with vigorous inde-

pendence among the various coenobia. Self-fertilization, therefore,

which is asserted for VY. globator, is obviously precluded here.

Not only is this species dioecious, but even the combination of

sexual cells with parthenogenetic reproduction, spoken of as if not

rare in V. globator, is here only an anomaly. I have seen but-three

such colonies.

Much more striking to the eye than any of the above charac-

ters is the number of reproductive cells which this superb species

presents. Even in the most poorly developed material collected in

the writer’s locality, the number of ova, or oosperms, often exceeds

one hundred, while in the finest material yet found it is not infre-

quently between three and four hundred, the large oval female

coenobia being thickly studded with crenate zygotes, leaving only a

small area of purely somatic cells at the anterior pole. When final

counts are made, I think considerably higher numbers will be

found. V. globator, on the other hand, shows as maxima, accord-

ing to different observers, from thirty to fifty female cells.

The male colonies differ much in size, number, and manner of

development. Sometimes they develop all their sperms nearly _si-

multaneously; at others their development is spread over a very

considerable period, nearly all stages being present at one time. Old

colonies may be found almost devoid of sperms or anlagen of them,

all or nearly all having been shed. Excepting such colonies as these,

however, the numbers of sperm groups is always very high. In

FURTHER STUDIES IN VOLVOX 165

place of the four or five of V. globator, we have, in the main, from

50 to 150 sperm masses or their anlagen in one colony.

Finally I will mention the most unique and most interesting

character of the species, viz., the manner of development and the final

form of the sperm bundle or sperm group; for “bundle” it is not and

still less a “platelet.” I introduce the term “sperm globoid,” for the

single aggregate of sperms here, instead of being flat and plate-like

is, in its formative period, a hollow, globular body, strangely simu-

lating a minute Volvox colony. As the spermocytes become differ-

entiated into perfect sperms this globular body becomes much flat-

tened, retaining, however, its cavity and showing sperms with out-

wardly directed flagella on all sides. These latter do not project

similarly from all the sperms; one side, even while the globoid is

still within its parental lodgement, always shows the flagella parted,

or, rather, radiating from a center; as we approach the periphery

they bend around the sides and extend straight backward, as do all

those on what we may call the posterior side. The whole arrange-

ment is obviously an adaptation to a special mode of locomotion,

a rapid spiral rotary movement with one pole habitually directed

foremost.

I have not as yet sufficiently studied the form of the single

sperms. They are obviously shorter and smaller than any I have

seen before, and are very numerous. I think a single globoid prob-

ably contains twice the maximum number hitherto given for the

single sperm platelet in any species. I find that these sperm globoids

are subject to some variation, which I hope to study in detail later.

I will but mention now that in material grown in much less sunny

climates, as Maine and Michigan, although undoubtedly of this spe-

cies, the globoids do not become always complete. The form is quite

the same. But a small, round opening remains on one side. This

suggests the possibility of a variation so considerable in amount as

to link this species with the typical VY. globator. There is little pros-

pect, I think, however, of such extensive variation, more especially

as V. globator has shown itself a rather non-variable species.

As to the minor characters of the vegetative cells., etc., they

plainly coincide, in the main, closely with those of V. globator. The

excessively numerous, highly stellate cells often quite duplicate the

166 J. H. POWERS

appearances of the figures of Overton, Meyer, etc. In the oldest

vegetative coenobia the appearance is a great exaggeration of any-

thing hitherto described. The single cells, becoming far removed

from each other, extend their cytoplasmic processes in irregular,

bent lines, until the cell body is hardly noticeable, and the appear-

ance, under a moderate magnification, is that of a sponge-like retic-

ulum. As mentioned earlier in this paper, I was able to verify per-

fectly the extremely delicate connecting fibrils which Meyer showed

were the real connection of the cytoplasmic processes one with an-

other. But in a few cases I observed, in old colonies, isolated in-

stances of a very different type of union. The heavy cytoplasmic

processes had themselves quite penetrated the bounding surfaces of

the gelatinous envelopes and united pairs of cells in a wholly differ-

ent manner. The connection in rare cases had been strengthened

until two cells had entered into close plastogamic union, quite resem-

bling that seen among rhizopods; the diameter of the connecting

bridge being little less than that of the individual cell bodies. Such

cases resembled binucleate or dividing cells,.save that the age of the

colony and the size of the cells precluded such interpretation.

In distribution, Volvox perglobator is probably little less than

universal in the United States. The most frequent species in my

own locality, I have also found it in the arid sand-hill region of

western Nebraska. Although not quite certainly established, veg-

etative colonies probably belonging to this species are found in ma-

terial from Washington. It occurred sparingly in the collection

from Rocheport, Missouri, and was well demonstrated and abun-

dant in plankton from ponds near New Orleans. I have seen the

same species from St. Louis, Missouri; from the vicinity of Ann

Arbor, Michigan; probably from Massachusetts; and, finally, from

the neighborhood of Sebago Lake, Maine.

Undoubtedly this is the species most frequently called V. glo-

bator in America. But Volvox would seem to have been more cas-

ually studied than well nigh any other organism of interest. Figures

have been published and labeled lV. globator that bore no resemblance

to the description or figures of European writers; even vegetative

coenobia bearing young colonies of thousands of cells each have

been mistaken and figured for colonies containing ova. All of

FURTHER STUDIES IN VOLVOX 167

which shows the need of more careful and extended work. Whether

the European V. globator really occurs with us at all or not, I have

as yet no exact evidence to determine. Very probably it does,

though less abundantly.

GENERAL REMARKS

In conclusion it is worth pointing out that these new species of

Volvox raise or throw light on several interesting problems with

regard to the genus. Without attempting full treatment, a little space

may be devoted to each.

The first of these is the nature of the Volvox aggregate as such:

what value has it over and above the unicellular condition, or that

of the much simpler aggregates in the same family. The answer has

been given that it is an ““Ernahrungsgenossenschaft,” a nutritive so-

ciety. The evidence for this has lain mainly in the fact of the

cytoplasmic connectives uniting cell with cell, and especially their

increased number in V. aureus at the points of union between re-

productive and somatic cells. The further fact has been cited that

division and growth of the somatic cells is not continued after the

development of the roproductive cells has begun. Now a rapid sur-

vey of the facts recorded for V. spermatosphara and V. weismannia

will show, I think, that these positions are not well taken. The first

species possesses no connectives between the cells whatever, if my

observations are correct. Its reproductive cells slip below the sur-

face at a very early stage in their development and seem to rest only

in the slightest contact with the somatic cells. In V. weismanmia I

am less certain about the general fact of connectives, but they are

probably absent, and the case of the reproductive cells is much more

convincing. Here their advent does not coincide with the cessation

in division of the somatics, and the development of both is for a

long time simultaneous. Especially instructive in this connection is

also the development of the sperm spheres and the dwarf male col-

onies of the two species respectively. In the case of the former a

large number of germ cells develop rapidly without the immediate

proximity of any somatic cells whatever. In the case of the latter

the somatic cells are small and utterly insignificant in number, while

the germ cells are rapidly developed and numerous, producing sperm

bundles of unusual size. Such instances seem an absolute disproof

168 J. H. POWERS

of the value commonly assigned to the cytoplasmic connection as

found in V. aureus and V. globator. I may add the observation

which I have more than once made that in instances where young

colonies within the parent have been almost completely destroyed by

rotifers, leaving a part of their reproductive cells with little or no

somatic support, these cells have none the less continued to develop

and even exceeded in size the reproductive cells of sister coenobia

within the same parent which had not been injured. If, then, we are

to regard the Volvox aggregate as a communistic nutritive society,

the medium of nutritive transfer must be, not the cytoplasmic pro-

cesses, but the gelatinous matrix itself and this alone. Even this,

however, is rendered improbable by such facts of reproduction as I

have cited in my previous article in regard to the “second form” of

Volvox there described. The life of the cells in Volvox is probably

nearly independent, that of the reproductive cells with the rest; they

doubtless grow mainly by their own nutrition, their chlorophyl be-

ing always abundant as long as growth continues. Protection, and

locomotion, leading to suitable illumination, are doubtless their only

advantages.

Upon another general matter of still greater interest I will, for

the present, touch with but a word. The determination of sex is a

matter of great present interest. Complex facts and complex inter-

pretations seem tending to supplant the simple nutritive hypothesis

that for a time was deemed of universal application. Sex in Volvox

is almost at its beginning, yet it is universally conceded. The rela-

tions, moreover, are anything but simple; parthenogenetic egg cells

developing into colonies of like content; or into male colonies, or

female colonies; or into colonies of various mixed types. Even in a

single species the phenomena are involved. Yet just herein lies, per-

haps, the possibility of tracing cause and effect and arriving at a

general conclusion. The different species, moreover, offer alto-

gether varying degrees of advantage as subjects for analysis. With-

out attempting, at the present time, a restatement of the facts with

this end in view, the writer will but remark that the detailed phe-

nomena of reproduction in V. spermatosphara and still more in

V. weismannia seems to yield strong support to the hypothesis that

nutritive causes, manifesting themselves especially in the size of the

FURTHER STUDIES IN VOLVOX 169

cells, are an important factor in determining the sex of the repro-

ductive cells or their products in these incipient multicellular or-

ganisims.

Next as to the general interpretation to be placed upon the pe-

culiar mode of sperm formation in V. spermatosphara and V. wets-

mannia. I have said of the sperm spheres that they may be inter-

preted either as reduced colonies in which somatic cells have been

completely replaced by reproductive members, or as mere groups

of multiplied spermogonia which, coincident with the general habit

of all growing cell aggregates of the genus, have assumed a spherical

form. The dwarf male colonies of V. weismanma might well tend to

support the first hypothesis of simple, reduced, precociously repro-

ductive coenobia. But again the ontogeny of these dwarf male

colonies reduces this probability. For these dwarf male progeny

are distinct from almost their earliest segmentation, every stage of

their development being peculiar to themselves. While seeking out

these facts it suddenly occurred to me that another hypothesis was

possible, and, indeed, far more probable than either of those men-

tioned, viz., that both the sperm sphere and the dwarf male colony

represent, not secondary acquisitions of any type, but earlier phylo-

genetic stages in the evolution of the genus, preserved in these spe-

cies on the male side, so to speak, only. In my former article I

spoke of the striking resemblance between a sperm sphere in the

spermogonia stage and a Eudorina colony. And since the finding

of numerous sperm spheres with but thirty-two members the paral-

lelism becomes still closer. We might even, without violence, as-

sume the original development of Volvox to have taken place di-

rectly from Eudorina, the sperm sphere serving as an almost perfect

connection between the two. Colonies of Eudorina have been re-

ported in which every cell became a platelet of sperms. The only

change to the sperm sphere would be the loss of independence, a

possible increase in size, reduction of cilia, etc., all minor changes.

The dwarf male colonies would evidently represent a slightly higher

stage in the evolution toward the Volvox aggregate, viz., the split-

ting up of a few cells, in the early time of segmentation into somatic

members, while the majority develop as before. On the female and

ordinary parthenogenetic sides the species have obviously reached a

much higher plane.

170 Jj. Bs ROWERS

If the foregoing interpretation is correct it suggests at once an

arrangement of the known species of the genus in a more or less

perfect series. V. spermatosphara stands lowest, not only because

of its Eudorina-like sperm spheres, but because of its lack of cyto-

plasmic cell connections, its small size, etc. Just above it stands V.

weismannia; above this undoubtedly V. tertius and the peculiar form

(very probably a distinct species) described by Klein as a variation

of V. aureus. Possibly, indeed, this form, when fully studied, may

rank yet lower in the series. Considerably above these, at least in

its ordinary types, stands V. aureus (minor); while yet further re-

moved stands . globator, with its late maturity and reproductive

cells which seem to be derived directly from somatic cells which

have already reached a high degree of differentiation. Highest of

all stands V. perglobator, which seems to intensify or carry further

nearly every feature of its better known relative. The sperm glob-

oids of this species might seem possibly open to an opposite inter-

pretation, as primitive sperm aggregates that retained the form of

a complete colony. The form of the ordinary sperm platelet is,

however, as long since pointed out, a still more primitive phylo-

genetic form, and the sperm globoid of V. perglobator undoubtedly

represents a later hypertrophy, so to speak, of the simpler and more

common type.

Be these interpretations, however, as they may, the little genus

Volvox is evidently rich in interesting surprises, and this paper will

quite have served its purpose if it stimulates their discovery as well

as aiding, in some degree, the general study of our American forms.

In conclusion I wish to thank the following persons, from whom

I have received valuable collections of material, greatly aiding me in

the present study: Prof. R. H. Wolcott, Lincoln, Neb.; Dean H. B.

Ward, Lincoln, Neb.; Dr. Elda R. Walker, Lincoln, Neb.; E. Foster,

New Orleans, La.; Bertram G. Smith, Syracuse, N. Y.

FURTHER STUDIES IN VOLVOX 171

EXPLANATION OF PLATES

Plate XXIII

Volvox spermatosphara

The figures on this plate give a typical representation of the forms,

sizes, and content of the different colonies of the species. All magnified

about 79 diameters.

Fic. 1. Medium to large coenobium containing 14 typical daughters.

Fic. 2. Medium to large coenobium containing 11 rather large daughters

and one sperm sphere composed of platelets of developed sperms.

Fic. 3. A free coenobium showing approximately the smallest size at

which they escape from the parent.

Fic. 4. Large female coenobium containing 11 oosperms.

Fic. 5. Medium to large coenobium contains six daughter coenobia and

six mature sperm spheres. Several of the daughters are somewhat distorted.

Fic. 6. Coenobium containing one daughter, one young oosperm and

one sperm sphere disintegrating into its component sperm bundles in the

manner invariable in this species.

Fic. 7. Coenobium containing 10 nearly mature sperm spheres and one

daughter coenobium.

Fic. 8. Coenobium containing seven nearly mature sperm spheres, one

young oosperm and one daughter coenobium.

Fics. 9 AND 10. Small female coenobia (very characteristic) containing

eight ova each.

Fic. 11. Small coenobium containing three immature (two in spermo-

gonia stage) sperm spheres and two ova as young oosperms. This com-

bination is by far less common than any of the others on the plate with

the possible exception of the balanced coenobia in figures 5 and 12.

Fic. 12. Small coenobium containing four daughter coenobia and four

immature sperm spheres.

Fe. 13. Small coenobium with two daughters, one immature sperm

sphere and three ova.

172 J. H. POWERS

Plate XXIV

The figures in the upper portion of plate show more eccentric coenobia of

V. spermatosphara; the smaller ones toward the lower end show the phe-

nomena of inversion in young coenobia of V. weismannia.

Fic. 14. V. spermatosphara, small, containing one sperm sphere of 32

sperm bundles, and one ordinary daughter. X 79.

Fic. 15. Small sterile coenobium of same species. 79.

Fic. 16. Coenobium with six daughters and one sperm sphere, the

platelets of which contain 64 sperms each. Very rare. X 79.

Fic. 17. Small coenobium with one relatively large daughter and one

ovum or young oosperm. The latter is mot in the daughter, but is below it

pressed against its wall. 79.

Fic. 18. Small coenobium containing mature sperm sphere only. X 79.

Fic. 19. Medium to large coenobium containing but single daughter.

SOT

Fic. 20. Coenobium showing maximum amount of crowding by daugh-

ter coenobia, of which there are 10, only seven or eight showing in the single

focus. x 79.

Fic. 21. Smaller coenobium showing single daughter of relatively large

size. X 79.

Fic. 22. Detail from one side of a young coenobium of V. spermato-

sphara. The young daughter coenobia still show no sex cells. One of them

is still unclosed and displays well the process of invagination, the first stage

in inversion. The daughter coenobium beside it, still not quite closed, is

perhaps beginning the same process, being flattened on the side toward the

parental wall. 217.

Fic. 23. Coenobium of V. spermatosphara with four daughters, show-

ing moderate degree of crowding. 79.

Fic. 24. Detail from one of two coenobia found representing possible

connecting types between Il’. spermatosphara and some other species. The

figure shows, mainly, a sphere or young colony made up almost entirely of

sperm platelets of 16 sperms each, a few undivided and therefore possibly

somatic cells being among them. It is hard to separate these, in the figure,

from the cells of the parental coenobium, but this may be done near the

periphery. 217.

Fic. 25. Young daughter coenobium of ’. weismannia in process of in-

version. Three of the sex cells are dimly visible. > 319.

Fic. 26. Coenobium of V. spermatosphara with four daughters and one

mature sperm sphere; shows crowding and crushing of the less resistent

sperm sphere. X 79. -

Fic. 27. Nearly inverted daughter of V. weismannia in the “kettle-

shaped stage,” the loosely attached reproductive cells show but poorly at the

sides. X 217.

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FURTHER STUDIES IN VOLVOX 173

Fic. 28. Inversion of very small and young coenobium of V. weismannia,

at the earliest stage found; the reproductive cells are present but hard to

make out in figure. 217.

Fic. 29. Another instance of inversion in same species; young colony

removed from parent. X 217.

Fic. 30. Completely inverted daughter—same species—about to reclose,

as shown by the x-shaped cleft in the middle. The reproductive cells are

here numerous, probably 18 or 20, and are plainly visible. They have but

just begun to sink into the depressions which form beneath them.

Fics 31 AND 32. Small daughter coenobia of V. weismannia, seen from

opposite poles, one removed from parent before photographing. Each has

eight reproductive cells, those in figure 32 having sunken well into the pits

that receive them.

Fic. 33. Another, rather early, stage in the same process and same

species.

Plate XXV

V olvox weismannia

All magnified about 79 diameters.

Fic. 34. Coenobium with nine daughters typically arranged. The lower

one lacks a mate that would have made the typical number 10. The lower

daughter is typically smaller; it is completely inverted, showing reproductive

cells on outside. The other daughters are still unclosed, showing the quad-

rangular openings on one side in several instances.

Fic. 35. Typical coenobium with 10 progeny, all of upper series with

10 reproductive cells each. Of second series, three are typical, while one is

a young male colony. Lower two show one small inverted colony and one

young male colony.

Fic. 36. A typical female coenobium with eight ova. The regular ar-

rangement is fairly well preserved; note flattened form of ova when seen

from side.

Fic. 37. Medium-sized, nearly mature coenobium with 10 daughters,

the regular arrangement being nearly preserved. The reduction in size of

the two lower members is somewhat greater than usual.

Fic. 38. Young coenobium with 10 reproductive bodies showing typical

arrangement. One of the two lower is an ovum. The one of the lower four

toward the left is in the process of inversion.

Fic. 39. Female coenobium with 10 ova; regular arrangement well pre-

served. The arrangement is, however, not typical, in that only the upper

four ova are in one plane.

Fic. 40. Coenobium with 10 reproductive members, of which one in

upper series (left hand) is becoming a young male. In the second plane

174 J Ve POWERS

the two toward the right are both inverted, as is also the farther of the two

lower ones.

Fic. 41. Female coenobium with nine ova, one in the upper series want-

ing. The ova are considerably displaced in the flattening of the whole colony

but the arrangement can still be discerned, especially by the two lower and

smaller members.

Fic. 42. Coenobium with 10 reproductive members, somewhat displaced.

The two lower are remaining as ova, and one of the lower four is a young

male colony.

Fic. 43. Coenobium containing the less frequent number of 12 repro-

ductive members. Unfortunately the originally regular arrangement is much

disturbed, but can be made out. The four lower members are undisturbed

in position and are all ova.

Fic. 44. Coenobium with eight progeny, the arrangement considerably

disturbed. All are typical save the left member in the lower series, which

has just inverted, showing the “kettle stage.”

Plate XXVI

Volvox weismannia

Fic. 45. Young coenobium (taken shortly after closure) seen from side,

showing the typical oblate form peculiar to young of this species. The eight

ova occupy typical positions. X 217.

Fic. 46. Nearly mature coenobium with but seven progeny. Note, how-

ever, preservation of the typical positions, one member being represented by

a noticeable vacancy in the lower series. One of the upper four is a dwarf

male colony with spermogonia nearly ready for division. X 79.

Fic. 47. Young daughter colony slightly older than figures 45, 48 and 49.

It shows a less frequent instance, double the number of reproductive cells.

Their position in pairs may be partially noted; also their size reduced by one-

half. 217.

Fic. 48. Typical colony, with eight reproductive cells, seen from one

pole. Compare with figure 45. XX 217.

Fic. 49. Typical young colony with 10 reproductive cells. The two

slightly smaller cells, removed a little farther from the periphery, are the

lower pair. X 217.

Fics. 50 AND 51. Still younger, unclosed, coenobia seen from opposite

poles. Each has 10 reproductive cells. Note, in figure 50, the determina-

tion of the position of the smaller pair by the fold-like borders of the open-

ing. X 217.

Fic. 52. Early differentiation of reproductive and somatic cells.

Fic. 53. Like figure 50, but a stage younger.

FURTHER STUDIES IN VOLVOX 175

Fic. 54. Very young dwarf male colony showing (dimly) the very

early differentiation of reproductive cells (spermogonia) and somatic cells.

Fic. 55. Nearly mature coenobium showing four dwarf male colonies,

their spermogonia being just ready to divide. This coenobium was included

in the plate partly as an example of irregular number in general content,

i.e., seven instead of eight young. Re-examination of the slide, however,

shows fragments of another dwarf male colony which had been nearly de-

stroyed by a rotifer, thus making the typical number eight. The small size

of one of the lower four is, nevertheless, not usual in an assemblage of eight

young. X 79.

Fic. 56. Young coenobium much like that in figure 49; the lower pair

of cells are dimly visible. X 217.

Fic. 57. Coenobium, nearly mature, bearing ten young colonies, all

dwarf males. Rather rare. X 79.

Fic. 58. Immature male colony in the stage at which most of the

spermogonia have taken up their final position within the colony, leaving

the few somatic cells alone to constitute the peripheral series. 217.

Fic. 59. Like the preceding save at a somewhat earlier age, some of

the spermogonia being still at the periphery of the sphere while others are

receding below the somatics. X 217.

Fic. 60. Mature dwarf male colony from within the parent, showing

mature sperm bundles. The focus is at the periphery of the colony to show

the few small somatic cells at the periphery while the mature sperm bundles

(of two sizes) are well within, forming as it were a second layer. X 217.

Fic. 61. Single sperm bundle of approximately 128 sperms (larger size)

seen from side and showing the slightly concave form characteristic of this

species.

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DATA FOR THE DETERMINATION OF HUMAN

ENTOZOA—II

By HENRY B. WARD

WITH ONE PLATE

Four years ago I contributed an article on this subject for the

Transactions. That paper dealt with then known facts regarding

human parasites as the means for their determination. The latter

problem is essentially a microscopical subject and the constant and

increasing use of this instrument in clinical work since then has

resulted in considerable additions to our knowledge. It is all im-

portant for the accurate identification of these forms that precise

data be at hand with reference to the individual species. As I

pointed out previously, the data in our possession are fragmentary,

and at times conflicting and inaccurate, so that such a positive de-

termination is often a matter of considerable difficulty. In this

paper I wish to add to the previous article a brief statement regard-

ing the important discoveries of recent years concerning the eggs

of human entozoa. It will be noted accordingly that this paper is

limited to a consideration of only a single diagnostic feature, the

eggs. While it is my intention subsequently to consider other feat-

ures noted in my earlier paper, yet, without that, the emphasis on

this one element would be justified by the paramount importance in

fecal examinations of accurate knowledge concerning the ova.

There has been much discussion also concerning the meaning of

supposed variations in these structures and some have gone so far

as to proclaim the unreliability of evidence adduced from them. In

the subsequent pages I hope to show that such opinions are due to

a misconception of the facts, and that research has been confirming

the accuracy of differential diagnosis based upon the character of

the ova.

At the outset it is necessary to dwell once more upon the fact

that in common with all similar structures, the ova are not abso-

lutely uniform in size. Some modern investigators have attempted

to make and publish measurements of these structures so carefully

178 HENRY B. WARD

expressed as to extend the values to tenths of a micron. Such accur-

acy is largely imaginary and is apt to be misleading, for the range of

variation between individual specimens, while small, is nevertheless

far beyond such insignificant limits. The average size may readily

be obtained by taking ten or a dozen normal specimens. Directly in

connection with this matter, it is important to call attention to the

error which is introduced by the habit of some investigators in re-

porting as the limits of size the absolute range which can be found

among the eggs in any preparation. This is likely to give a false

idea concerning the characteristic type of egg in the species under

question and is distinctly misleading when the figures are taken

up by a subsequent observer as a means of determining the nature

of a questionable species. Where the limits are unreasonably large,

such an observer is very likely to think that the eggs he has under

examination may properly be included in the species in question,

when, as a matter of fact, they fall within the limits of abnormal

eggs only and do not conform to the measurements of normal ova.

In this matter I agree entirely with Looss, who says (1907 :149,

footnote) : There exist, of course, among the immense number of

ova in an individual worm always some which are either larger or

smaller than the rest, or even evidently misshapen. In my opinion

it is of no use to record carefully the measurements of these eggs

also. For the description and definition of a species it is much

more important to select for measurement those ova which appear

to be normal and to present the size and shape typical for the spe-

cies. It may be added in passing that young worms with few ova

in their uteri usually do not afford normally-shaped and normally-

sized ova.

Constant differences in size between specimens measured and

the measurements given for a species under consideration create a

prejudice at once in favor of the view that the two species are dis-

tinct. Disregard of this almost axiomatic statement has brought

much confusion. More than once in the past, some of the sup-

posed wide variations in size among the eggs of a certain species

have been found to be due to the confusion of two or more closely

related forms under a single specific name. This matter was em-

phasized in the previous article; and, in at least one instance, dis-

cussed there, eggs which had been assigned to a certain species, but

DETERMINATION OF HUMAN ENTOZOA 179

which yet appeared to be radically smaller than the ordinary meas-

urements of ova in that species, have since been shown to belong

to a different species. Further mention of this item will be found

later under the discussion of Paragonimus Westermanit.

The egg of Schistosoma haematobium figured in my previous

article (Ward, 1903a, pl. 1x, fig. 10) appears to have been some-

what exaggerated in size through inadvertence either in copying

or reproducing the figure. Accordingly I have deemed it wise to

present a new figure of the egg in this species. It will be noted by

comparison that the one shown this time is somewhat narrower in

proportion and that the embryo fills a larger part of the shell. The

latter feature is one which to some extent at least is dependent

upon the stage of contraction of the embryo and will vary consid-

erably in different eggs. No other errors in text or figures of the

first paper have been called to my attention.

In agreement with the plan followed in the previous article all

figures of ova reproduced on the accompanying plates are repre-

sented at the uniform magnification of 500 diameters. Much labor

has been expended in the effort to secure for each species the best

illustration available. In some cases no figures of the egg exist and

in order to secure a representation for ready comparison with other

species I was forced to make use of geometric lines drawn as ellipses

on the dimensions given for the desired ova. While necessary, this

method is very undesirable, since few eggs are a mere ellipse. The

method disregards entirely the taper towards the poles of the egg,

the size and character of the lid when present, the color, thickness,

transparency and other features of the shell, and the nature of the

contents. All of these items furnish distinctive features of import-

ance in certain cases, and should be more generally and fully re-

corded.

The egg shells are the only immobile factors in trematode

structure; they are also the portions most usually found and most

readily used in the clinical diagnosis of infection with helminthes.

Evidently, then, more accurate data concerning them will prove of

great value to the clinician.

The idea seems to be prevalent that these structures form an

unreliable basis for diagnosis on account of the large variation to

which they are subject. I am convinced that this is the most serious

180 HENRY B. WARD

sort of an error. All my own observations tend to show great uni-

formity in size under similar conditions. This uniformity is

strongly evidenced in the work of Looss among Trematoda. Whole

genera, if not sub-families, among the groups which he is handling

in admirable fashion, manifest such close agreement in this respect

that the general description of the group includes only a narrow

range of size among the ova in the entire series of forms within

its limits. These data are far more valuable than has yet been

clearly appreciated, and the very existence of marked variation in

the records of a single form is presumptive evidence that in the

absence of errors of observation, two or more species are confused

under the single appellation.

The most extensive additions to our knowledge in this field

which have been made within the last four years fall under the

Trematoda. Consequently, the major portion of this paper is de-

voted to that group. No important additions along this line have

been made to our knowledge of cestodes parasitic in man. Among

the nematodes, however, several species have been discovered and

some valuable data can also be presented with reference to older

torms. For general data to be utilized in the differentiation of ova

and in the diagnosis of parasites by other structures available for

clinical purposes, the reader is referred to the preceding paper

(Ward, 1903a).

Of Cladorchis Watsoni, which has been found but once, para-

sitic in a negro of German West Africa, Shipley (1905:7) says that

the eggs are encased in a shell and contain many deeply staining

yolk-granules. He gives measurements for the egg of 122 » by 80 p,

but Conyngham records the size as 130 by 75 ». The egg has not

yet been illustrated. Only a single case of parasitism by this spe-

cies has been recorded as yet, so that its pathological significance

is uncertain.

Since the publication of my first paper Fasciola gigantica, a

form previously confused with Fasciola hepatica or at most re-

garded as a variety of that parasite, has been clearly differentiated

as an independent species. One case of its occurrence in the hu-

man lung is on record. A second probable case is noted by Mus-

grave (1907:17) for the Philippine Islands. According to Looss

DETERMINATION OF HUMAN ENTOZOA 181

(1902:783) the eggs of Fasciola gigantica, with rare exceptions,

vary between 145 and 152 p» in length by 80 » in breadth. In the

same species described by Railliet under the name of F. angusta the

size of the eggs is given as 145 to 150 pu by 82 to 88 u.

The eggs of Fasciolopsis Busku, which in my previous paper

were described after Odhner and which then had not been delin-

eated, have been studied and illustrated by Loos (1905-6). I repro-

duce a copy of his figure in the accompanying plate (pl. xxvu1, fig. 3).

He gives the range of size among the ova as 120 to 130 p» in length

and 70 to 80 » in breadth. While he as well as Odhner note that

these eggs are in all respects like those of Fasciola hepatica, yet the

figure of Looss shows them to taper more toward both. poles, to

have smaller operculum and a thinner shell than the ova of F.

hepatica. All of these points are clear on comparison of the figures

for the two species. The contents of these ova as they appear in

the feces show plainly the single germ cell and mass of yolk cells.

Development has not yet begun. Manson (1907:738) says that

these ova are closed by a very delicate operculum.

A single specimen of F. Buskit was found at the necropsy of a

Laskar sailor in Galveston, Texas, by Moore and Terrill (1905).

These authors give the average size of the ova as 150 by 75 pw. If

the magnification assigned to their microphotographs is correct not

even the largest reaches this length and the actual size agrees close-

ly with the measurements of Looss cited above.

Goddard has recently (1907 :195) reported a case of this para-

site from China; the material was referred to a committee and that

part of the report of the Investigation Committee which deals with

the eggs reads as follows: Eggs (possibly immature). Size not

measured, about a half that of Ascaris lumb. Shell, very thin

walled. Contents clear, small and granular, well marked nucleus in

center. Nearly spherical. No operculum observed.

An error is certainly present here, for the size of these ova

would equal only 40 to 45 » by 25 to 30 » on the basis they give in

their estimate. This is far too small for Fasciolopsis Buskii. If one

infers that the proportions relative to Ascaris lumbricoides are by

accident reversed in their statement these ova would approximate

130 to 150 » by 100 to 116 » and this again fails to agree with the

182 HENRY B. WARD

species F. Buskii to which they assign these specimens. In fact, the

designation “nearly spherical” which they use to characterize the

ova can not by any means be applied to the figure which Looss

gives of ova in F. Buskit. Renewed investigations alone can tell

whether the authors are in error as regards their measurements and

descriptions or with respect to their identification of the species.

Fasciolopsis Rathouisi was originally described by Poirier in

1887 and since that time no case of its occurrence has been record-

ed before the present year. Many authors, including Leuckart, who

examined one of the original specimens, have considered it iden-

tical with Ff. Busku; but in common with others I have constantly

maintained its specific integrity, although in a recent paper (Ward,

1903 :867) assigning it to the genus Fasciolopsis, originally created

for F. Buskit.

In the paper just cited Goddard (1907:195) has recorded one

case of simple infection supposedly with this species and one case

of mixed infection with it and F. Buskii. The original description of

Poirier is inaccessible to me; according to various authors it states

that the eggs are ovoid, measure 150 by 80 yp, and show great simi-

larity to those of Fasciola hepatica. The account of Goddard is as

follows: The eggs of the D. rath. are about two-fifths of the micro-

scopic field under a one-sixth-inch objection and one-inch ocular,

have a thin shell and appear to possess a hyaline body moderately

well filled with coarse granules of a greenish yellow tint [ ? in fresh

feces]. These eggs were present in both cases reported [one was

a case of mixed infection with F. Buskii. |

The report of the Investigation Committee to whom these spec-

imens were submitted is given in the same paper (Goddard 1907:

198). The description of the ova reads thus: Egg, oval, size not

measured, about one-third larger than Asc. lumbric. Thin walled

and smooth with very small operculum. Contents appear to con-

sist of large granules.

From this account the size of the ova may be estimated as ap-

proximately 100-115 » by 65-75 ». Such an ovum would be very

much smaller than that described by Poirer for his Distomum

Rathouisi. The Investigation Committee notes in its report some

doubt as to the correctness of the identification, and the doubt is

DETERMINATION OF HUMAN ENTOZOA 183

emphasized by this discrepancy in the size of the ova. Further

study of these forms is needed to establish their true character.

In the previous paper (Ward, 1903a:118) I called attention to

the great discrepancies in published records regarding the ova of

Paragonimus Westermanti, and put together there the measurements

of various authors in an outline sketch (pl. vin, fig. 1). In the text

attention was especially called to the smallest measurements, namely,

those recorded by Yamagiwa and the suggestion made that an er-

ror had been made or another species was concerned. Since then

the discovery of Schistosoma japonicum has demonstrated the truth

of the latter alternative and the cases of Yamagiwa, who gave

measurements of ova from the brain and lung, are now believed to

concern this new species. Even after the elimination of these and

of the earlier, probably incorrect records of Baelz, there still re-

main marked differences between various records. These differ-

ences are not reconciled by more recent investigations, but on the

contrary rather emphasized, as the following record of such inves-

tigations will show.

The only case of parasitic hemoptysis, due to infection with

Paragonimus Westermant, yet observed, in man in this country

(Mackenzie, 1904:1134) was diagnosed by means of microscopic

examination and demonstration of the ova in the sputum of the pa-

tient, a Japanese fisherman, working on the Columbia River. Thirty-

one ova were measured and found to vary from 85.5 » to 99.7 p» in

length and from 48 to 69 » in breadth, with an average size of 91.3 »

by 55.2 ». The measurements given by Katsurada (1900) vary from

87.5 » to 102.5 w in length by 52.5 » to 66.3 » in breadth, with aver-

age values of 93.5 by 57 w. These two series agree very closely

with each other and with records of Baelz, Leuckart and Manson.

It should be noted that all were made on parasites from hosts of

the same nationality and undoubtedly represent the same species of

parasite.

Of importance is the observation of Looss on this species. He

says (1905 :283, note 1): The size of the ova in Paragonimus Wes-

termanti is given rather differently by previous authors: 80 to 100 »

in length by 50 to 70 » in thickness. Since I have found them in

mature intact worms only 77 to 81 » long and 43 to 50 » thick, the

184 HENRY B. WARD

larger measurements cited above speak for confusion with other

eggs. The eggs of P. Westermanii possess, moreover, a relatively

thick yellowish to reddish-brown, plainly operculate shell and at the

time of elimination from the body of the host always contain only a

simple germ cell surrounded by yolk cells.

These measurements of Looss do not agree with any others

from man, though nearest those of Musgrave (1907) and markedly

different from those of Katsurda and Mackenzie. The source of

the material used by Looss is not given. The accuracy of Katsur-

ada’s work can not be doubted and his material came from the

classic field of infection with the Asiatic lung distome; hence the

eggs of this species were before him when preparing his record of

the size and character of the ova. Nevertheless, the average meas-

urements of Loos agree exceedingly closely with those of Ker-

bert, based on the original material of P. Westermanu obtained

from the tiger.

Paragonimus Westermanti has been most carefully studied by

Musgrave (1907) in the Philippines. He says of the ova: The

eggs are oval in shape and vary in color from a reddish-brown to

light-yellow, the color depending somewhat upon the stage of their

development and the source from which they are obtained and per-

haps, also, upon other conditions which are not fully understood.

Again, their size varies considerably, and a glance at Table No. 3

will show the marked differences in the measurement given by

some of the principal observers. I have carefully measured several

specimens with the Zeiss photomicrographic apparatus, with the fol-

lowing results:

Length, 0.0062 to 0.0098; average, 0.0074.

Breadth, 0.0045 to 0.0063; average, 0.0057.*

Eggs in sections of tissues and those preserved in any other

manner contract somewhat and these are not considered in the

measurements given above. A well-marked operculum is probably

present in all mature specimens, but in many instances this is in-

distinct and at times it can not be made out at all. This operculum

has quite a distinctive appearance, which is noticeably different from

**These figures are evidently far too small and I conclude that the decimal

point is incorrectly placed and they should read 0.062 mm., etc., or 62 y, ete.

DETERMINATION OF HUMAN ENTOZOA 185

that of some of the other fluke eggs. The operculum in the ova

under consideration appears to fit into an opening very much in the

manner of a cork into a bottle, whereas in some of the other trema-

todes, particularly Opisthorchis, the opercula appear to be placed

over openings more in the fashion of a cap.

The ovic cell in many specimens is quite distinct, and several

yolk cells are then also present, but in others, even if they are taken

from the same abscess, no such structure can be made out.

The record of Musgrave agrees in breadth of the ova exactly

with the determination by Katsurada (1900:508). But the latter

records the average length of eggs as 93.5 » as against Musgrave’s

average measurement of the length at 74 ». This adds another dif-

ficulty to a series of measurements already manifesting greater va-

riations than are known to occur among the eggs of any other tre-

matode. I have been for some time of the opinion that the Amer-

ican form reported from the cat, dog and hog, constitutes a dif-

ferent species from that found in man in the East. Yet the adop-

tion of this view does not clear up the question satisfactorily. Either

one must acknowledge an unusual variation in size among the ova

of this species, or an exceptional number of errors in observing or

recording their measurements, or be forced to conclude that several

species of closely similar appearance and structure have been

bunched under the one name. Musgrave’s cases apparently all con-

cern natives of the Philippine Islands.

These measurements and those given in my previous paper

may be most readily compared in the form of a table which I ap-

pend here.

—RANGE OF SIZE— AVERAGE

AUTHOR DATE LENGTH BREADTH SIZE HOST COUNTRY

Kerbert 1881 80 x45 Tiger Holland

Leuckart 1889 80 -100 - 56 90* x56 Man Japan

Ward 1894 96 -118 48 -55 102 x53 Cat WreSeA:

Stiles and

Hassall 1900 78 - 96 48 -60 85.6x53.2 Hog Wi-S:"A:

Katsurada 1900 87.5-102.5 52.5-66.3 93.5x57 Man Japan

Mackenzie 1904 985.5- 99.7 48 -69 91.3x55.2 Man (Jap.) U.S-A.

Looss 1905°- 77 —-81 43 -50 79* x46* Man? ?

Manson 1907 80 -100 40 -60 ?90* x50* Man ?

Musgrave 1907 62 - 98 45 -63 74 x57 Man Philippines

All measurements in microns.

* Estimated.

186 HENRY B. WARD

The entire matter may be summed up in the following para-

graph, in which regard is had not only to the ova, but to other

structural elements as well. After extended study I am convinced

that the American form which I described originally and identified

as Paragonimus Westermani is undoubtedly a distinct though close

related species to which I now wish to give the name of Paragomni-

mus Kellicotti. It is well known through the original description

and the subsequent complete accounts of Stiles; yet it is my inten-

tion to publish shortly a study of this species, giving more complete

justification of its specific distinctness from the Japanese form.

I am endeavoring to secure material from the Philippines also

to compare with each of these two forms as well as with material

from the tiger. The great similarity in the various measurements

of Japanese material made by different observers at widely sepa-

rated times and places, namely: Baelz, Leuckart, Manson, Katsura-

da, and Mackenzie, whose averages vary only a few microns, indi-

cate that this form may be after all a distinct species from Kerbert’s

tiger parasite with which the Japanese measurements do not agree.

The Japanese form will, however, not need a new name, but prob-

ably will go back properly to take one of the earlier names used

by investigators of the human parasite in Japan. To determine cor-

rectly the facts in this case demands most careful study and hasty

action will only add to the confusion already existing.

In a recent most valuable paper, Looss (1907) has differen-

tiated the old species Opisthorchis sinensis into two distinct forms,

Clonorchis sinensis, the Distomum innocuum of Baelz, and Clonor-

chis endemicus, the Distomum endemicum of Baelz; the latter prac-

tically takes the place of the true Clon. sinensis in the literature pub-

lished after 1883 and is at present the one usually described in the

text books, etc., as “Opisthorchis sinensis.”

Regarding the ova of Clonorchis sinensis Looss says (1907:

149): The average dimensions of the eggs are 0.029 mm. in

length and 0.016 mm. in width; the limits of the former being

0.026 and 0.03 mm., and of the latter 0.015 and 0.017 mm. In many

specimens of the species the eggs show a distinct narrowing to-

wards the anterior extremity, and their rather high lid is marked off

by a sharply projecting brim. I have, however, also seen specimens

in whose ova these peculiarities were but little pronounced.

DETERMINATION OF HUMAN ENTOZOA 187

The description of the ova of Clonorchis endemicus given by

Looss (1907:151) is as follows: The eggs have about the same

_ length as those of the preceding species (0.026 mm.), but their width

is decidedly somewhat smaller, amounting on an average to 0.015

mm., with the lower limit at 0.013, the upper at 0.016 mm. The nar-

rowing towards the anterior end is in the main not so marked and

the margin of the rather flat lid not so sharply projecting as in Clon.

sinensis ; but these differences are on the whole very slight and not

recognizable in every specimen.

I have reproduced here (pl. xxv, figs. 1 and 2) figures of the

two species given by Looss to illustrate his descriptions. Evidently

the exact determination of the species on the basis of the ova alone

will be a difficult matter even under favorable conditions. It should

be noted that Looss does not find the blunt process at the pole of the

egg opposite the lid which was illustrated after both Ijima and Kat-

surada in my previous paper (Ward, 1903a:137, pl. 8, fig. 7).

Of the greatest importance for the proper recognition and ac-

curate identification and differentiation of the human parasites are

careful studies of the ova under different conditions and at various

phases of development. A most valuable work of this type has re-

cently been published by Saito (1906:133) on “Distomum spathu-

latum.”’ The species on which he worked was evidently the Japa-

nese form, which Looss more recently has designated Clonorchis

endemicus. Saito presents a brief record of the data regarding

these eggs as given by others, especially the Japanese investigators,

and then states his own results, from which the following is taken:

At its tapering end the egg shell is provided with a helmet-shaped

lid; the line of union is marked by a projecting rim. The embry-

onic development occurs in the uterus and eggs taken from its te

minal coils contain completely developed embryos. Fresh eggs in

the uterine coils occasionally open of themselves, naturally much

more easily those from feces. The so-called “Stabchenkorper,”

which up to the present time has been regarded as a yolk remnant,

must be considered as a part of the embryo.

The appearance of the egg at different stages is both described

and figured, as also the appearance and structure of the embryo. Un-

fortunately the author does not give a series of careful measure-

188 HENRY B. WARD

ments and his figures are not accompanied by a record of the exact

magnifications. It is worthy of note that he represents fresh ova

both with the short, blunt process at the posterior end and without

this structure.

During the past few years our knowledge of the schistosomes,

or blood flukes of man, has been greatly extended. To the well

known Egyptian blood-fluke, Schistosoma haematobium, two new

species have been added which also occur in the human host. Of

the original species, S. haematobium, a recent valuable publication

by Looss (1905b:101) records the following important data: When

in the uterus of the female worm and immediately after deposition

these ova measure on the average 80 by 30 » and contain an unseg-

mented egg cell surrounded by a mass of yolk cells. During their

stay in the tissues where cleavage and development of the embryo

takes place, the eggs increase so markedly in size that at the close

of this period they usually measure 110 to 120 p» in length by

46 to 50 » in breadth. The thin, yellowish shell carries at one end

a short pointed spine of variable length. Among these one finds

often another form which is characterized by a larger, lateral spine.

Regarding the latter type of ovum, which is said to occur in

feces only, more will be said later on in this paper. Certain it is

that among eggs taken from the urine there is found more than one

type which departs noticeably from the normal average form of the

egg in S. haematobium. The form with a lateral spine has been

taken as the basis of a new species, S. Mansont, which is considered

below ; but the same method can hardly be followed in the case even

of all recorded variations which are of an extreme type. Promi-

nent among such let me call attention to the observations of Chris-

tophers and Stephens (1905:259). In an important note on a pe-

culiar schistosome egg, these authors record that these eggs were

met with in the urine of a native of Madras suffering from haema-

turia. The egg differed from that of the Schistosoma haematobium,

inasmuch as it was of a long spindle shape, and from the thick end

of the egg a long, snout-like process projected. The egg measured

205.2 by 53.2 p, being therefore much longer than other schisto-

some eggs. In the urine there occurred also eggs indistinguishable

from those of S. haematobium, so that the significance of the spindle-

shaped variety was uncertain.

DETERMINATION OF HUMAN ENTOZOA 189

When one considers that the ova observed in this case are only

slightly broader than the normal egg and yet 75 to 80 per cent

longer, it will be acknowledged that such a difference can not pos-

sibly be accounted for by any normal variations in size. What were

the abnormal conditions producing such ova or whether a new spe-

cies is concerned, must await further investigations. The possibil-

ity of a mixed infection with two species of Schistosoma can not be

dismissed without consideration. Further observations in the same

region would throw light upon the question; but so long as only one

case is on record, these ova must be considered as merely abnormal

variations of a radical type.

A new species of Schistosoma, designated the Asiatic blood-

fluke (S. japonicum) was first described and named by Katsurada

(1904). His account of the eggs of this parasite may be abstracted

as follows: Eggs from human feces; usually oval or elliptical, thin-

shelled, yellow or light brownish yellow, without lid, and without

spine. In fresh material they measure 75 to 90 » by 52.5 to 72.5 p,

with an average size of 83.5 » by 62.5 ». The egg from feces con-

tains a well developed, ciliated embryo. Ova in human liver tissue

measured, on the average of 25 specimens, 72 by 49 uy, being thus

somewhat smaller than in feces, owing to the influence of the pre-

servative.*In the liver of a cat the ova measured 64 yp, or in a sec-

*Compare in this connection the more probable explanation of

Looss, cited above (p. 188) for the increase in size of the ova of S. haem-

atobium between the time of deposit by the female worm and of exit

from the tissues of the host. So large a contraction as indicated by Kat-

surada’s figures, 7. e., one-sixth or more, could not in my opinion be

brought about by the action of preserving fluids. I have measured ova

in several species while fresh and after preservation in various fluids and

have never been able to demonstrate any positive constant difference.

Irregular shrinkage does of course occur, so that many ova can not be

measured at all, but uniform contraction appears to be confined at least

within narrow limits. It is important that this matter be re-examined

to determine whether the changes in size from liver to feces are con-

stant. Katsurada indicates that the transition is accompanied by the

growth of the embryo which is not formed in eggs from the liver, but

present in eggs taken from feces. I see no reason why eggs in intestinal

tissue or in feces should contract less than those in liver tissue under

methods of preservation probably similar. It is clear, however, how the

growth of an embryo might gradually enlarge the shell.

190 HENRY B. WARD

ond host 70 by 74 yn, and in the intestinal wall 67 by 50 ». In the

uterus of the female worm the average length of a series of ova was

58.4 pw, the average breadth 42.5 »; thus the uterine egg is the

smallest of those described. The hosts were all native to Japan.

Almost simultaneously with Katsurada, Catto described the

same species. He gives the measurements of the ova as 60 to 90 »

by 30 to 50 w, with an average of 70 by 40 ». He speaks of the shell

as stout, but otherwise agrees with the description cited above. The

host came from China.

Recently a case of this parasite has been reported from the

Philippine Islands by Woolley. In the description of the parasite

this author says (1906: 87): The following comparative measure-

ments of the ova were furnished to me by Dr. Shiga, after he had

examined my specimens and compared them with those of Fujinami

and Manson:

Manson Fujinami Woolley

| VS TY eG 9 «Re ie, Nae eg SR A 72.8 p 66.2 p 62.4 p

BOAGEMY Oo Stes eek eC ms 48 43.6 43.6

This case was in a native Filipino who had never been out of

the islands.

In general one may say that the eggs of Schistosoma japonicum

are smaller than those of S. haematobium, have blunter ends and no

spines. There are also considerable differences between the meas-

urements given by the various observers, some of which are

explained on the basis of Katsurada’s observations concerning the

stage of development of the ova. These observations are cited

above.

The early history of Catto’s material is worthy of note as indi-

cating the difficulties at times in the way of an accurate micro-

scopic diagnosis. At a necropsy certain lesions were observed and

tissue preserved and later sectioned from the liver, mesenteric lymph

glands and intestine. These sections were examined by at least

three pathologists and showed numerous small oval bodies having a

smooth, stout capsule. Opinions differed as to their character, but

the case was originally published as one of coccidiosis in man. This

is not the first case in which trematode eggs have been diagnosed

and published as coccidia (Ward, 1903a:115). Specimens were also

DETERMINATION OF HUMAN ENTOZOA 191

shown at the Medical Research Club, London, but no definite con-

clusion was reached regarding the nature of the oval bodies. An

eminent German authority to whom pieces of the tissues were sent

stated “that the oval bodies were neither coccidia nor the eggs of a

trematode, but those of a nematode of unrecognizable species, etc.”

At the British Medical Association in July, 1904, the case was

reported to the section on tropical diseases, and the abstract of the

proceedings (J. Trop. Med., Aug. 15, 1904; London Lancet, Aug.

27, 1904) includes but one interpretation of these structures, viz., as

eggs of an oviparous nematode new to pathology. Stiles (1905)

errs in stating that the author claimed he had a new species of

trematode for man, apparently also in giving the date of this pub-

lication as September 7, 1904.

I was at the International Zoological Congress at Berne,

Switzerland, in August, 1904, when Sir Patrick Manson sent some

of Catto’s slides to Professor Blanchard, who at once called into

consultation Looss, Monticelli, Stiles, and myself. The view that

these questionable bodies were nematode eggs was reported to us

and promptly discarded in favor of the opinion advanced first by

Looss that they were the ova of a new blood-fluke, evidently related

to S. haematobium, but readily distinguishable from that species.

Later in the same summer both Stiles and I had the opportunity of

examining Catto’s material in London and of confirming the pre-

liminary diagnosis. By most careful work Catto dissected adult

worms out from the infected tissues and published a full account of

the case and its parasite as Cragg’s Prize Essay for 1904 from the

London School of Tropical Medicine.

The eggs of this species are by no means new in pathology.

Many cases from Japan in which eggs have been reported previously

in nodules from the omentum, mesentery, peritoneum, rectum, and

from a cirrhotic liver, are interpreted by Looss (1905 :282) as prop-

erly belonging to this new Schistosoma japonicum. Katsurada in-

cludes in the same category the cases of Yamagiwa in which eggs

were found in the cortex cerebri and were regarded as the cause

of Jacksonian epilepsy. I pointed out in my previous paper (1903a:

118) that they could hardly belong to the species Paragonimus Wes-

termanti, to which the Japanese author had assigned them, and sug-

192 HENRY B. WARD

gested that if the records were correct, a new species was probably

concerned. The drawing | gave agrees well with the measurements

assigned to ova of S. japonicum by both Catto and Katsurada. Looss

very justly calls attention to the necessity of distinguishing sharply

between the two species in cases where ova occur in nodules in

human tissues and suggests that. this will not be difficult in view of

the different structure of the shell and of its contents. The data for

the ova of Paragonimus are given above.

Booth (1907 :201) is the first to give a detailed descrption of the

eggs and embryos of S. japonicum from fresh material. At the

Hankow (China) Hospital, he examined ova from fresh feces. They

were a little larger than those of Ascaris lumbricoides in the same

material, oval in shape, transparent, thick-shelled, and occupied by

a well developed embryo. The shell had neither spine nor opercu-

lum. The embryo was oval and smaller than the shell, while within

the shell on each side of the embryo lay two oval yolk bodies. The

embryonic cilia were in motion and two hours’ incubation of feces

at 37° C. resulted in finding many empty shells. In every case the

shell ruptured at the side and the last part to leave was the anterior

end of the embryo.

Very recently Sambon has described a third species of Schisto-

soma from man and bases the description entirely on the ova. He

lists for them the following characteristic features: the ova are

large, with rounded ends and a lateral spine, which is heavy, promi-

nent and sharply pointed. According to Holcomb, who has been able

to examine a large number of these eggs, they measure from 112 to

162 » in long diameter, and from 60 to 70 » in breadth. So far as

known, these lateral-spined ova are voided exclusively with the feces

and endemic haematuria is totally unknown in the West Indies

where, according to the latest reports, this form is extensively prev-

alent. Sir Patrick Manson discovered such ova in 1903 and sug-

gested their specificity. Similar ova had indeed been seen before

and according to Bilharz and Mantey they occur singly in isolated

cases.

On the basis of personal experience Looss (1905b:102) rejects

this view regarding them as abnormal structures which with regu-

larity are produced at the outset of sexual activity and suggests that

DETERMINATION OF HUMAN ENTOZOA 198

possibly mature but unimpregnated females produce such ova. He

represents the mode of formation of such an egg and also illustrates

a transition form between the normal terminal-spined egg and the

lateral-spined form which he regards as abnormal. I| reproduce here

(fig. 7) from his work a lateral-spined ovum to show the type he

has in mind. He calls especial attention to the extreme rarity of

this type, which is by far most frequently found in the liver where

they not rarely occur unmixed with normal eggs. The figure of

such an ovum copied from Looss seems to confirm his view since in

the place of the usual well formed embryo this egg contains a gran-

ular mass indefinite in character save for numerous oil (?) globules

(fig. 7). Such an appearance indicates a disintegrating mass, and

accords with the theory that the ovum was not fertilized. On the

other hand, the constant differences in size, place of deposition and

geographical distribution point very strongly to the specific inde-

pendence of this form. It may be that in reality there are two

forms: an abnormal variety of S. haematobium, rarely found, which

contains a degenerating mass of cellular debris and a normal, lat-

eral-spined ovum, belonging to an independent species. The adinis-

sion of Looss that the lateral-spined ova are by far most frequent

in the liver accords with their attainment of the intestinal wall and

ultimate escape from the body by the feces rather than by the urine

as in the case of S. haematobium. Most careful study should be de-

voted to the cases in which lateral-spined ova are found. In case

the somewhat larger size, noted by Sambon, is confirmed on further

measurements the specific identity of the worm will be establishicd

even before the adult is seen.

The lateral-spined ovum, copied after Looss, is much smailer

than the minimum figures for the ova from the West Indies, which

Sambon regards as belonging to S. Mansoni. This difference in

size as well as a distinct variation in furm are evident from a com-

parison of Looss’ egg (fig. 7) with an outline giving the dimen-

sions assigned to ova of S. Mansoni by Sambon. This lends color to

the view that the type which Looss has illustrated is not the true

S. Mansoni but some abnormal form. I incline to the opinion that

the view of Manson, worked out more recently by Sambon, wi!!

prove to be the correct one.

194 HENRY B. WARD

Since writing the above discussion, I have secured a copy of a

paper by Holcomb (1907) which gives a most exhaustive discussion

of the subject and leaves no doubt of the specific validity of S. Man-

sont. He shows that there is throughout the West Indies an exten-

sive schistosome infection which is characterized by lateral-spined

ova voided in the feces of the host. This same form is also most

frequent in Central Africa, while the variety producing terminal-

spined ova exists in its purest form in Cape Colony and a more or

less mixed infection occurs in Egypt. Holcomb believes that the

females described by Looss and others were possibly not abnormal

types, since such an ootype as described by Fritsch must have pro-

duced invariably the lateral spine. The mere fact that the spine was

lateral would hardly account for the fact that these females usually

selected the intestinal tract in which to deposit their eggs, ror would

it explain why this particular worm should find the conditions to

perpetuate its type when transplanted to the West Indies. The West

Indian infection proves that they are not the eggs of unfertilized

females, and some of his cases, which were under observation for

one year or more, show only too well that the persistence of this

type of egg can not be attributed merely to young females. In the

light of these facts, Holcomb believes that these other observers have

described not abnormal forms, but really the essential difference

between the female Schistosoma haematobium and the Schistosoma

Mansoni.

It is not impossible that the Schistosoma Mansoni of the West

Indies is a lineal descendant of the parasite now known as existing

in Central Africa and that it was carried to these localities by the

traffic in slaves.

This somewhat lengthy discussion is justified by the confusion

which has existed and the very recent date of the elucidation which

the subject has received. In the light of Holcomb’s careful report

there remains only the single doubt created by the great dissimilar-

ity between his figure of the ovum of S. Manson (fig. 8) and that

for the so-called abnormal ovum of S. haematobium, according to

Looss (fig. 7). The exactness of the latter observer suggests

again the view I advanced above that the type he illustrates was after

all not S. Mansoni. It is impossible to believe that the same species

DETERMINATION OF HUMAN ENTOZOA 195

could have ova varying so greatly as these figured by Looss and by

Holcomb.

In his complete and detailed description of the new species, Hol-

comb (1907 :66) says of the eggs that they are found without diff-

culty in the feces when present.* * * They do not occur in the

urine. * * * They are oval in shape and of a pale straw color,

and so transparent that the inclosed miracidium is readily made out.

My measurements of a number of eggs have shown them from 112

to 162 » in the long diameter. In breadth they measure from 60 to

70 p. They are provided with a spine of from 15 to 17 ,» in length,

and which is sharply pointed. This spine, although I have previous-

ly reported it at the juncture of the posterior and middle third of

the shell, is, from further measurements, more correctly located at

the juncture of the last and third quarter of the egg. The spine is

directed backward. When the shell is broken by the miracidium

the rent is usually in the direction of the short diameter and curving

toward the long diameter as it approaches it.

Lining the inside of the shell is the vitelline membrane. This

membrane is usually made out without much difficulty and is clear-

est in the pole of the egg, at which the cephalic extremity of the

miracidium presents. Here there is usually a clear space between

the membrane and the shell.

Within the vitelline membrane is inclosed a miracidium—a

brownish membrane of granules in a high state of molecular vibra-

tion—and several clear, slightly opalescent bodies. The brownish

membrane consists of a thin layer of granules which surrounds the

whole miracidium. They are usually found in a high state of mole-

cular activity in the matured eggs. The opalescent bodies are more

frequently two in number, but I have found as many as five. They

are of a somewhat hyaline substance and though of oblong dimen-

sions, usually saddle the miracidium so that the side view presents,

giving them the appearance of oval bodies jammed between the

miracidium and the shell. These bodies appear to be of the nature

of food for the embryo, as in cases where the feces have been mixed

with sand and kept for one or more days they are usually absent.

The miracidium can be readily seen through the shell.

No important data regarding the ova of Cestoda have been se-

196 HENRY B. WARD

cured since the publication of my previous paper. Moreover, for

the clinical determination of tapeworms the eggs are of subordinate

value. The alimentary canal being the seat of adult cestodes in

man and a diagnosis being regularly made by the discovery of pro-

glottides in the feces, the problem is much simpler than with tre-

matodes or nematodes.

Among the round worms or nematodes half of the species or

more are viviparous, and consequently do not enter into considera-

tion here. Among the others there are a few species concerning

which important new data are recorded.

On the basis of a careful study of an extensive series of speci-

mens and of type material, Looss (1905a) has been able to distin-

guish three separate species among the nematodes formerly diag-

nosed as Strongylus subtilis and to establish the identity of the true

species subtilis with that earlier designated as instabilis by Railliet.

The forms belong properly to a distinct generic group named Tri-

chostrongylus by Looss. Of the three species which occur in man,

Tr. mstabilis is the most common; it is also the only one as yet

found in the human host outside of Egypt, having been reported

from Japan by Ijima. In Egypt Looss records that he has found

the parasite frequently, but always in insignificant numbers, not ex-

ceeding 30 to 40 individuals in a single host. Its infrequence, small

size, and lack of hooks, bristles, and buccal armature, indicate that

its presence is not accompanied by mechanical disturbance of the

alimentary lining and Looss is unable to detect any pathological sig-

nificance in its parasitism. In Japan, however, it occurs occasionally

in man in great numbers and under these circumstances may be of

pathological importance.

The ova are thin-shelled, colorless and in form and appearance

much like those of the European hookworm (Ancylostoma duode-

nale), so that on superficial examination of fecal material from man

it may be confused with them. In the uterus of the young female

parasite they have not begun development, but in older worms or

when taken from feces they show 8 to 32 cleavage cells already

formed. The egg shell measures 73 » to 76 p, or rarely 80 mw in

length by 40 » to 43 p» in breadth, being thus much larger than those

of the European hookworm, which measure 56 p» to 61 » in length

DETERMINATION OF HUMAN ENTOZOA 197

by 34 » to 38 w in breadth, and somewhat larger than those of the

American hookworm (Necator americanus), which are 64 p to 72 u

long by 36 » thick. According to Looss, the latter eggs are espe-

cially characterized by a marked tapering toward one pole, a feat-

ure not found in either of the other species.

In Trichostrongylus probolurus, a closely related species, the

eggs are similar in appearance and nearly like in size, measuring

76 » to 80 » in length by 43 » to 46 » in breadth. This species has

been reported hitherto only in Egypt as a human parasite and even

there it appears to be infrequent in the human host.

Still a third species, Tr. vitrinus is also recorded as a rare hu-

man parasite in Egypt. Its eggs are somewhat larger, measuring

84 to 90 » in length by 46 to 50 » in breadth. The eggs of these

three species are so much alike that on the basis of present knowl-

edge nothing more accurate than a diagnosis of the genus could

probably be made from an examination of the fecal matter contain-

ing them.

A new sclerostomid, parasitic in man, has been described by

Railliet and Henry (1905) as Triodontophorus deminutus. It was

collected in Africa at the autopsy of a native. The ova are ellip-

soidal; measured through the body wall, they vary from 60 to 65 p

in length and from 38 to 40 » in breadth. The same authors have

also described (Railliet and Henry, 1905a) another new sclerosto-

mid, Oesophagostoma Brumpti, from a negro in Africa, but in this

case no eggs were found in the worm.

Only a single record is at hand concerning Physaloptera cau-

casica. Of the ova of this species its describer, von Linstow, says

that they are very thick-shelled and measure 57 » long by 39 yp

broad. This species was noted in my earlier paper as uncertain in

importance and data regarding its eggs were not cited. On account

of its possible recurrence in the Orient, if at all, these facts are given

briefly here to make them available for comparison.

A single record concerns a new nematode from man, reported

from Texas and described as Ascaris texana. It appeared doubtful

that the form should retain its position in this genus, and this view

is strengthened by the re-examination of the material by Stiles. The

question must remain undecided until the male is found and de-

198 HENRY B. WARD

scribed. According to Smith and Goeth (1904) the ova of this para-

site were numerous in both specimens. The description concerns, of

course, the uterine egg which is reported as well advanced in cleav-

age, colorless, provided with a thick wall, without visible external

marking or envelope as met in the eggs of certain of the Ascarides.

The ova measure 60 by 40 p, with a variation of 4 to 5 » in different

specimens measured. The interior appeared granular, no nuclet

being appreciable in the specimens at hand. Two drawings of ova

are given; they do not agree well in form, appearance or internal

structure. Further data are essential if this species is to be recog-

nized from fecal examinations ; meantime, it cannot be averred that

the species is more than purely casual as a human parasite.

DETERMINATION OF HUMAN ENTOZOA 199

BIBLIOGRAPHY

Boorg, R. T.

1907. Notes on Ova and Embryos of Schistosomum Cattoi vel japonicum.

Jour. Trop. Med., x: 201-203.

CurisToPHERrS, S. R., AND STEPHENS, J. W. W.

1905. Note on a Peculiar Schistosomum Egg. Jour. Trop. Med., vim:

259.

GopparRD, F. W.

1907. Two Rare Fasciolidae. China Med. Jour., xx1: 195-198.

Hotcoms, R. C.

1907. The West Indian Bilharziosis in Its Relation to the Schistosomum

Mansoni (Sambon, 1907), with memoranda in ten cases. LU eres)

Naval Med. Bull, 1: 55-80.

KaTSURADA, F.

1900. Beitrag zur Kenntniss des Distomum Westermanni. Beitr. pathol.

Anat. u. allg. Pathol., xxvii: 506-522.

1904. Schistosomum japonicum ein neuer menschlicher Parasit, durch

welchen eine endemische Krankheit in verscheidenen Gegenden Jap-

ans verursacht wird. Annot. zool. japon., v: 147-160. 1 PI.

Linstow, O. von.

1902. Zwei neue Parasiten des Menschen. Centralbl. Bakt. u. Par., Orig.,

XXXI: 768-71. 4 Figs.

Looss, A.

1902. Ueber neue und bekannte Trematoden aus Seeschildkroten. Zool.

Jahrb., Syst., xvr: 411-894. 11 Pls.

1905. Schistosomum japonicum Katsurada, eine neue asiatische Bilhar-

zia des Menschen. Centralb]. Bakt. u. Par., Orig., xxx1Ix: 280-285.

1905a. Notizen zur Helminthologie Aegyptens VI. Das Genus Tricho-

strongylus n. g. mit zwei neuen gelegentlichen Parasiten des Men-

schen. Centralbl. Bakt. u. Par., Orig., xxxrx: 409-422. 2 Pls.

1905b. Von Wiirmen und Arthropoden hervorgerufene Erkrankungen.

Mense’s Handb. der Tropenkrankh., 1: 77-209. 2 Pls.

1907. Some Parasites in the Museum of the School of Tropical Medi-

cine, Liverpool. Annals Trop. Med. and Par., 1: 123-152.

Mackenzie, A. D.

1904. A case of parasitic hemoptysis or infection with the Distoma Wes-

termantit. Jour. Am. Med. Assn., XLIt: 1133-1135.

MANSON, Sir P.

1907. Tropical Diseases, a Manual of the Diseases of Warm Climates

4th Edit., 853 pp.

200 HENRY B. WARD

Moors, J. T., and TerRIL1, J. J.

1905. Fasciolopsis Buskii (Distomum Buskii): the first reported case in

the United States; found in a patient dying of typhoid fever. Jour.

Am. Med. Assn., xL_v: 1002-3. 1 PI.

Muscrave, W. E.

1907. Paragonimiasis in the Philippine Islands. Philippine Jour. Sci.,

m Be 15-38:

RAILLIET, A., AND HEnry, A.

1905. Un nouveau sclerostomien (Triodontophorus deminutus nov. sp.)

parasite de l’homme. Compt. rend. Soc. Biol., Paris, Lvut: 569-571.

1905a. Encore un nouveau sclerostomien (Oesophagostomum Brumpti

nov. sp.) parasite de homme. Compt. rend. Soc. Biol., Paris, tym:

643-645.

Sarto, S.

1906. Ueber den Ejiinhalt des Distomum spathulatum und die morpho-

logische Beschaffenheit eines Embryos. Centralbl. Bakt. u. Par.,

Orig., XLi1: 133-138. 10 Figs.

SAMBoN, L. W.

1907. Remarks on Schistosomum Mansoni. Jour. Trop. Mod., x: 303-

304.

SHIPLEY, A. E.

1905. Cladorchis Watsoni (Conyngham), a human parasite from Africa.

Thompson, Yates & Johnston Lab. Rept., Liverpool, vr: 129-135.

iL were

Situ, A. J., AND GoETH, R. A.

1904. Ascaris Texana; a note on a hitherto undescribed Ascaris parasitic

in the human intestine. Jour. Am. Med. Assn., XLiit: 542-544.

STILEs, C. W.

1905. The new Asiatic bloodfluke (Schistosoma japonicum, 1904, Schist-

osoma Cattoi, 1905) of man and cats. Amer. Med., 1x: 821-823.

Warp, Henry B.

1903. Trematoda. Wood's Ref. Handb. Med. Sci., Rev. Ed., vir: 859-874.

1903a. Data for the Determination of Human Entozoa. (Studies from

the Zoological Laboratory of the University of Nebraska, No. 55).

Trans. Amer. Micr. Soc., xxiv; 103-138. 4 Pls.

WooLLey, P. G.

1906. The Occurrence of Schistosoma japonicum vel Cattoi in the Phil-

ippine Islands. Philippine Jour. Sci., 1: 83-89.

DETERMINATION OF HUMAN ENTOZOA 201

EXPLANATION OF PLATE XXVII

All of the figures have been brought to the same scale in copying and the

reproduction has made them of the uniform magnification of 500 diameters.

Unless otherwise stated they represent normal and average ova for the species

indicated.

Fic. 1. Clonorchis sinensis. The chief forms presented by the eggs,

each egg taken from a separate specimen of the worm. After Looss, 1907,

pl. rx, fig. 7.

Fic. 2. Clonorchis endemicus. Eggs figured in the same manner and

under exactly the same enlargement as in the preceding drawing so as to

allow of comparison. After Looss, 1907, pl. 1x, fig. 8.

Fic. 3. Fasciolopsis Buskii; egg from uterus of full grown worm. After

Looss, 1905, pl. 1x, fig. 4. é

Fic. 4. Schistosoma haematobium; egg from uterus of female worm,

occasional only in feces and then generally calcified. After Looss, 1905, pl.

Ix, fig. 8.

Fic. 5. Schistosoma haematobium; normal egg from urine. After Looss,

1905, pl. 1x, fig. 7.

Fic. 6. Schistosoma japonicum; egg from fresh feces. After Katsurada,

1904, pl. vu, fig. 1.

Fic. 7. Schistosoma haematobium; abnormal (?) egg from feces. After

Looss, 1905, pl. 1x, fig. 9.

Fic. 8. Schistosoma Mansoni; showing the lateral-spined egg with the

miracidium. The different structures are shown diagramatically. After

Holcomb, 1907, p. 66, fig. 3.

Fic. 9. Trichostrongylus instabilis; egg from feces. After Looss, 1905,

pl. rx, fig. 21.

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PERMANENT PREPARATIONS OF TISSUES AND OR-

GANS TO SHOW GLYCOGEN

By SIMON HENRY GAGE

From the time of Claude Bernard’s discovery of glycogen (1850-

1860) it has been known that strong alcohol retains it in the tissues,

and therefore alcohol has been used whenever one wished to deter-

mine the presence of this substance by microscopic examination. Ab-

solute alcohol is usually recommended for the fixation, but as gly-

cogen does not dissolve in alcohol above 60% it seems unnecessary

to use absolute. 95% alcohol has given perfectly satisfactory results.

Some authors recommend formalin and other fixers. It is true

that some glycogen remains and can be demonstrated after the use

of formalin and some other fixers; but all are agreed that alcohol

is the most satisfactory fixer when glycogen is to be demonstrated.

As alcohol penetrates slowly one can get the best fixation by using

only small or thin pieces suspended in the alcohol. Bernard himself

recommended the use of a mixture of equal parts of a saturated al-

coholic solution of iodin and glacial acetic acid prepared at the time

of use. This mixture is not so satisfactory as alcohol alone. A

modification of Bernard’s method has been found excellent, and it

has the advantage of not shriveling the tissues so much as pure al-

cohol. The formula is 95% alcohol, 100 cc.; 10% iodin in 95%

alcohol, 2.5 cc.; glacial acetic acid, 1 cc.

For sectioning, the paraffin method is the most satisfactory. The

sections are spread on slides, using one of the following iodin solu-

tion instead of water: Water, 500 cc.; sodium chlorid, 1.5 grams ;

potassium iodid, 3 grams; iodin crystals, 1.5 grams (or 15 cc. of a

10% solution of iodin in 95% alcohol). If one has to do with very

soluble forms of glycogen an alcoholic solution for spreading the

sections is preferable. Formula: Water, 250 cc.; 95% alcohol,

250 cc.; sodium chlorid, 1.5 grams; potassium iodid, 3 grams; iodin

crystals, 1.5 grams.

When the sections are spread care must be taken not to over-

heat the slides, for the paraffin should in no case be melted. While the

204 SIMON HENRY GAGE

sections are being spread the iodin stains the glycogen, and one can

examine the sections with low powers without further treatment.

The stain remains in the paraffin sections for an indefinite period.

This gave a clue, for permanent preparations of iodin-stained glyco-

gen. The ordinary hard paraffin shows crystals too plainly. It was

thought that if a liquid or semi-liquid paraffin could be used it might,

like the solid paraffin, preserve the iodin stain in the glycogen. In

sections so mounted high powers of the microscope could be used.

Many forms of liquid paraffin were tried, but none of them

were satisfactory. White vaseline was then used at the suggestion

of Dr. Mall, and it answered fairly well. The best results were,

however, obtained by the use of the ordinary yellow vaseline sold by

druggists. After many experiments the final mounting was accom-

plished as follows: If the paraffin sections had faded or were not

deeply enough stained the slide bearing the sections was immersed

in one of the staining solutions given above for 5 or 10 minutes or

longer. The glycogen will be deeply stained if the paraffin has not

been melted in spreading the sections. The slide is then allowed

to dry thoroughly—half an hour in a drying oven or an hour or

more (preferably over night) in the laboratory. The yellow vase-

line is melted over a water-bath, and then the slide with the sections

is put into xylene until the paraffin is dissolved out of the sections

—2 to 5 minutes. After the paraffin is removed the sections are

mounted in the yellow vaseline as one mounts preparations in bal-

sam. The slide may be slightly warmed so that the vaseline can be

mostly pressed out; the excess of vaseline is wiped away and the

cover sealed with balsam or shellac, so that it will not be easily

moved. Preparations so made have retained the glycogen stain of

iodin for three years.

Another method was tried with success, viz., that of mounting

in balsam without a cover-glass as with Golgi preparations. Prep-

arations so made have retained the stain for over six months. The

balsam preparations are somewhat more satisfactory for study with

the highest powers.

If homogeneous immersion objectives are to be used on these

uncovered preparations it is better to revert to the original homo-

geneous immersion liquid, viz., thin Canada balsam (See Cox, Proc.

PREPARATION OF TISSUES 205

Amer. Micr. Soc., 1884; Mayall, p. 96). Of course, the ordinary

cedar oil may be used on the covered preparations mounted in vase-

line. It will be found in most, if not in all, of the tissues which were

quickly and thoroughly fixed that the glycogen is by no means in

granules, but appears as a homogeneous substance in the cells.

Many attempts have been made to use other stains than iodin

for the glycogen, that is, stains which would be permanent and en-

able the investigator to mount the sections in balsam. None of the

methods so far proposed give so differential a stain as iodin; and

they are difficult of application. They are of course desirable to use

in connection with the iodin stain if one is making a critical study.

The two methods most often employed are the gentian violet

method of Lubarsch and the carmin method of Best. Both these

methods are quite fully and satisfactorily dealt with in Ehrlich’s En-

cyclopedia of Mikroskopie, under Glycogen.

In working upon embryonic and some other material it is some-

times desirable to decalcify the contained bone. This can be safely

done, as follows: Fix thoroughly in the absolute or 95% alcohol or

the iodin alcohol given above. Decalcify in 67% alcohol containing

3% nitric acid (Gage, Proc. Amer. Micr. Soc., 1892, p. 121). The

sections are made by the paraffin method as before. Of course, one

may use the collodion method of sectioning, but then the only way

known to the writer for making permanent preparations stained

with iodin is to allow the collodion sections to dry. They may then

be mounted in vaseline or in balsam as directed above.

If one attempts to dehydrate the sections the iodin will be dis-

solved out. The secret of success is to stain the sections before the

removal of the paraffin; to use no alcoholic solutions after the sec-

tions are stained with iodin, but to mount directly in vaseline and

cover, or in balsam without a cover-glass, as directed above.

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NECROLOGY

FRANK L. JAMES, Ph. D., M. D.

Dr. Frank L. James, past president of the American Micro-

scopical Society and for eighteen years editor of the scientific de-

partments of the National Druggist, of St. Louis, Mo., died in

that city May 19, 1907. He was taken sick May 5, grew steadily

worse, and erysipelas developed in its most violent form. It is

supposed the infection took place from a sore caused by the friction

of a celluloid shield worn over a defective eye, the result of an

accident about ten years before.

Dr. James was a man of varied and unusual attainments and

had quite an eventful career. He was born in Mobile, Alabama,

August 27, 1841. A New England ancestry was back of him, on

his mother’s side; paternally, he came of English yeoman stock,

which settled in the “tidewater” country of Virginia in early Colo-

nial days. His was a rare intellect, and his facility in the acquire-

ment of languages was little short of marvelous. There is hardly

a language that has a literature with which he was not more or less

familiar, and most of them he could read and translate with ease.

At sixteen he was prepared to enter the Polytechnique at Carlsruhe,

Germany, whither he was sent to study civil engineering. His

passion, however, was for chemistry, and by rare good fortune, at

eighteen, he became a member of the household and a worker in the

laboratory of Baron Justus von Liebig.

Leaving the companionship of Liebig, he graduated in medi-

cine in Paris, just as the coming on of the Civil War sent young

Americans home to espouse one side or the other. Dr. James’ sym-

pathies, naturally, were with the people of his native section, and he

determined to come back to his native country, and to throw his

fortunes with the Southern side. The passing of the Federal lines

presented a difficulty, since the blockade had already been estab-

lished, but his wit brought him through. A fluent speaker of Ger-

man and somewhat of German physique, he made his way under

208 NECROLOGY

the guise of a military officer to Havre, where friends procured

him a passport. Before leaving Paris, he had received from Con-

federate Commissioner Slidell dispatches for President Davis.

Passing over to London, he undertook the same commission from

Mr. Mason. A few days found him in New York, “a German inter-

ested in the war between the States.”” There he met an old friend,

a former business partner of his father, who supplied him with $250

in gold. He went to Washington, and thence, by invitation, to

Alexandria, where he was the guest of Federal officers. All the

while, the dispatches were concealed in the holsters of the pistols

he carried. He then gradually made his way toward the Confed-

erate lines, and, finally managing safely to slip into them, was soon

with his family in Mobile. The dispatches were in due time deliv-

ered to President Davis, who, becoming acquainted with Dr. James’

thorough knowledge of European languages, and being impressed

with his youthful daring and discretion, appointed him to a place

in the secret service. Here his technical knowledge and skill found

use in the making of the first torpedoes used in the memorable strug-

gle. He was personally engaged in planting the explosives in Mobile

Bay which blew up a Federal war vessel and caused the loss of

many lives. As a secret service man he was at one time directing

the catching and salting of fish for the army, using opportunities

thus afforded him for communicating with the outside.

The war over, he went into the temporary retirement of country

life on a farm in Mississippi, where his sister, Mrs. Mosby, had

her home.

In 1866 he became connected with the topographical survey

of the Mississippi River under Col. Folsom. While thus engaged

he became so much interested in the Indian mounds in Osceola

county, Arkansas, that he began and conducted a systematic explora-

tion of them, the results of which are recorded in the records of

the Smithsonian Institution. In the early seventies, he was con-

spicuous in reconstruction history as associate editor of the Mem-

phis Appeal, but gave up journalism in 1877 to enter on the prac-

tice of medicine in St. Louis. A born writer, however, he continued

to make frequent contributions to the secular as well as to the

scientific press, and in May, 1884, he was selected as editor of the

NECROLOGY 209

therapeutic department of the National Druggist. He assumed the

editorial management at the beginning of 1888, and from that time

until his death, with only a brief interval, he held this position.

Possessed of a many-sided genius, his life abounded in contra-

dictions. Capable of tender affection, he never married; profoundly

learned in medicine, he was personally careless of many conditions

physicians insist upon; a fluent and graceful writer, he had no am-

bition for publication, his only published work, so far as is known,

outside of his contributions to journalism, being a manual for stu-

dents, entitled “Elementary Microscopical Technology.” With an

intellect of great breadth, he was content to shut himself in; with

vast acquirements in fields that have made other men famous, he

pursued the even tenor of his way; lavish in the expenditure for

books, he spent little otherwise.

As age advanced, his friends and associates gradually grew

fewer and fewer, and his last days were days of loneliness and soli-

tude. But in his library he always found solace and entertainment,

and his books were to the last his closest and dearest friends.

Dr. James was a skillful microscopist, and his researches with

this instrument were to have been given in full to the scientific

world when the accident already referred to deprived him of the

sight of one eye, and defeated his cherished purpose. He was an

inventor and maker of both chemical and microscopical appliances

for his own use, and, until the last, maintained his great skill in

the handling of tools. It was characteristic of him that he never

took out a patent, though there were many evidences of his handi-

work stored in his laboratory.

Dr. James became a member of this Society in 1882 and served

once as president, presiding at the Washington meeting in 1891,

and though in latter years compelled to forego the pleasures which

the microscope afforded him, he never lost interest in the Society.

210 NECROLOGY

RUDOLPH SIEMON

Rudolph Siemon, the subject of this sketch, was essentially a

self-made man, born March 25, 1837, at Ziesar, Prussia. He ar-

rived in the United States in June, 1856, and came directly to Fort

Wayne, Indiana, where he had a brother and two sisters living.

Shortly after his arrival, he, together with his brother, established

the book store of Siemon & Brother, which they succeeded in mak-

ing one of the leading establishments of its kind in the state of

Indiana.

In the year 1886, Mr. Siemon sold his interest in the book

business to his brother and spent some time in travelling. Upon his

return home he established the Siemon Wall Paper Company, which

he conducted until shortly prior to his death.

Mr. Siemon was an active and successful business man; he

was fond of study and was held in high esteem by all who knew him.

He was a bachelor and died at the age of seventy years.

In 1890, Mr. Siemon joined Sections A and F of the American

Association for the Advancement of Science, and a year later he

became a member of the American Microscopical Society. He took

great interest in the proceedings of both associations, but of late

years was unable to attend the meetings, on account of his declining

health. F. W. KuEHNE.

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NOTES, APPARATUS, REVIEWS, ETC.

WEIGERT’S [IRON-HAEMATOXYLIN STAIN

BY JAMES H. STEBBINS, jR.

In the Van Gieson muscle and fibrous connective tissue stain,

it is customary to stain the tissue first with one of the haematoxylin

stains, and then try to counterstain with picric acid-acid fuchsin.

Those familiar with the process will have noticed that in order to

obtain satisfactory final results, that is, to have the haematoxylin

show up well, it is necessary to greatly over-stain with the latter,

as the subsequent treatment with the picric acid-acid fuchsin exerts

a strong bleaching action upon the haematoxylin, and greatly re-

duces its strength. To therefore overcome this defect, Weigert has

worked out the following stain, which has given me the most satis-

factory results. Tissue stained by this process shows the connective

tissue stained a bright red to carmine, and the muscle fibres of a

bright yellow color, while the cell nuclei stand out sharply in

either blue-black or black. Those interested in photo-micrography

will find this stain of great help in obtaining sharp, crisp pictures.

The iron-haematoxylin stain is prepared as follows:

PEPIN ALARVLEN trp hr E Wry tes a ale dena ¥ a Gackt 1 grm.

Salpraléohiolt! si shes Eleet Stn evcag kc Mean Aneel Oke ra! 100 c.c.

Perric chloride: solution: (109%))i a. eee cee 4 c.c.

Eivarochioric acid* (259) towed. fas see ere

mseIicd Watel s/s Sede AU Ee it aes OO Sate 95 «exe:

For use mix equal parts of 1 and 2 and stain tissues fixed in

alcohol, or formalin, to the desired shade; wash well in one or two

changes of water, and then counterstain in:

Saturated ag. ‘solution of picric acid: ..! 4 jus uae 100 c.c.

_ somo of acid fuchsin i. 1a. Ske 1O<ee;

212 NOTES, APPARATUS, REVIEWS, ETC.

Rinse in water, dehydrate in alcohol, and mount in balsam.

The iron-haematoxylin solution may be kept for several days in

good condition, or until it begins to smell strongly of ether.

Watson & Sons’ NEw CATALOGUE

There is a tendency on the part of microscopists to underesti-

mate the importance of the condenser or illuminating apparatus

which is used in all high-power microscopical work. A large num-

ber of well-known houses do not even quote anything beyond the

Abbé illuminator. But it has been pointed out time and again that

the limitations in working with a condenser of this description are

very great, and its continued use is probably due to the failure of

workers to appreciate these limitations, and because its very want

of aplanatism renders it easier to work with than a more accurately

made system. Briefly stated, the Abbé illuminator consists of two

lenses only, and it is neither aplanatic nor achromatic. The maxi-

mum solid cone which it is capable of transmitting is .50 N. A.,

with the result that, however large an aperture the objective may

have, its effective working under critical conditions is reduced by

the condenser to .50 N. A.—that is, if an investigator is using a

1-12-inch oil immersion of 1.30 N. A. and Abbé illuminator, he is

only actually employing about half of the effective aperture of the

lens.

We are reminded of this by a perusal of a new edition of the

catalogue of the well-known English house, W. Watson & Sons,

313 High Holborn, London, in which, as is common with the lead-

ing English houses, a great feature is made of substage condensers

and principally those of large aplanatic aperture. They range from

an oil immersion system having a full aperture of 1.35 and an

aplanatic aperture of 1.25, through other systems having respect-

ively apertures of 1.0 N. A. with an aplanatic aperture of .9, and

another of .5 N. A. with an aplanatic aperture of .48, to the macro

illuminator, which is designed especially for producing a uniform

illumination of large objects under low powers.

On this account alone this catalogue is worth the considera-

tion of microscopists; beyond this there is a wealth of information

in it concerning not only the typical English stands which are associ-

NOTES, APPARATUS, REVIEWS, ETC. 213

ated with this firm’s name, but probably every accessory which

microscopists can wish for.

Special interest attaches to the introduction by them of two

new objectives. One is a 1-6-inch of their parachromatic series,

which has been made by them on a newly computed formula of their

expert, Mr. Conrady. The unique feature of this lens lies in the

tact that it possesses a working distance of 1 millimetre. This

means that the bacteriologist can use it on “hanging drop” cultures

and employ it freely with his haemacytometer covers and thickly

covered objects without any risk whatever of damage. In addi-

tion to this, apart from the claims of the makers as to its fine per-

formance, there is the testimony given at the October meeting of

the Royal Microscopical Society of London in a paper communi-

cated by Mr. E. M. Nelson, the well-known authority, that “ew

lenses, apart from apochromats, have shown as well as this new

1-6 inch, a balsam-mounted Pleurosigma formosum—the most se-

vere test to which any dry lens such as this can be subjected”; and

further statements to the same effect were given by Dr. E. J. Spitta,

the well-known author of “Microscopy,” and Dr. Eyre, bacteriologist

to Guy’s Hospital, London.

The other lens is a 2 mm. of Watson’s holoscopic series, which

is now made with a guaranteed numerical aperture of 1.37.

Many pages in the 176-page catalogue are devoted to photo-

micrographic apparatus, and we can heartily commend an exam-

ination of this list to microscopists. It will well repay the writing

of a request to forward it, which Messrs. Watson inform us they

will have much pleasure in complying with, free of charge.

THe Microscopy oF TECHNICAL PRopUCTS

A companion volume to Dr. Winton’s Microscopy of Vegetable

Foods, and equally attractive in make-up and contents is the Mi-

croscopy of Technical Products, by Dr. T. F. Hanausek, revised

by the author and translated by Dr. Winton. Like the former, this

volume is published by John Wiley & Sons, of New York City.

Dr. Hanausek is distinguished as an investigator, a teacher,

and a technical expert, and his work is distinguished by scientific

accuracy, clearness, and utility as a guide in diagnosis. In the

214 NOTES, APPARATUS, REVIEWS, ETC.

revision of the book, previously recognized as authority on the sub-

ject, much new matter has been added to the chapters on textile

fibres and the number of practical examples largely increased. The

number of cuts is also increased.

The volume, as it stands, is an octavo, of XII + 471 pages, with

276 illustrations. It is designed to serve both as a reliable scientific

guide to the student entering the field of technical microscopy, and

also as an aid to the technical worker in the solution of purely prac-

tical problems. The volume is unique in that it teaches the micro-

scopic identification of technical products and at the same time the

fundamental principles of vegetable histology and the histology of

certain animal materials.

Part I deals with Apparatus and Methods; Part II with the

Microscopy of the Most Important Types of Technical Raw Ma-

terials. Under the latter head are chapters on starches, in which

are discussed the general character, formation, and chemical char-

acters of starch, and the source, methods of preparation, and micro-

scopical characters of the different kinds; and chapters on animal,

vegetable, and mineral fibres, including a treatment of the exam-

ination of paper and textile fabrics. A chapter on stems and roots

includes also a discussion of barks and contains an analytical key

for the identification of the most important economic woods, which

has been revised to include all the important American woods; 32

pages are devoted to this key and 147 species of woods are included

and described. A chapter is devoted to leaves, and, under the

head of flowers and parts of flowers, another to insect powders.

Still another chapter on fruits and seeds includes a discussion of

various oil cakes, and one is devoted to teeth, bone, horn, etc. The

book closes with a chapter on micro-chemical analysis.

The volume is generously illustrated, very carefully written,

in an unusually lucid style, and cannot help but be of great value

to all microscopists who deal especially with technical products, as

well as of interest to all workers with the microscope who meet with

such material, incidentally, in their more general work.

PROCEED!NGS

The American Microscopical Society

MINUTES OF THE ANNUAL MEETING

HELD AT

ITHACA, NEW YORK, JUNE 29 AND 30, 1906

The twenty-ninth annual meeting of the American Microscop-

ical Society was held in conjunction with that of Section F of the

American Association for the Advancement of Science, in Stimson

Hall, Cornell University, Ithaca, New York, and was presided over

by President S. H. Gage.

FIRST SESSION

The first session, called to order at 4:30 Pp. M., June 29, was

occupied by the annual address of the President on the subject,

“The Origin and Development of the Projection Microscope.” The

address was accompanied by demonstrations; it proved a most at-

tractive topic and Prof. Gage’s able presentation of it was enjoyed

by a large audience.

At the conclusion of the address a brief recess was taken,

after which the Society again came to order for a business session.

The Secretary presented an informal report, which was, on motion,

received and accepted. The Custodian’s report was read and, on

motion, accepted and referred to an auditing committee consisting

of Drs. A. T. Kerr and M. D. Ewell. The Treasurer’s report was

also presented, accepted, and referred to an auditing committee

consisting of Profs. H. B. Ward and Chas. Fordyce.

A nominating committee was elected consisting of Dr. R. H.

Ward, Prof. B. F. Kingsbury, Dr. W. C. Krauss, Prof. W. H.

Watson, and Dr. M. D. Ewell.

The Abbé Fund Committee presented a report of progress;

on motion the committee was continued and was instructed, on the

216 PROCEEDINGS

accumulation of a suitable sum, to close the subscriptions and turn

the amount over to the secretary of the committee, who should for-

ward it to the secretary of the Denkmal committee at Jena.

The matter of a monument to Herbert R. Spencer was brought

up by Dr. Ewell, who described a visit made to his grave and stated

that he found it unmarked in any way. This was accompanied by

a recommendation by the Executive Committee that the Society

consider favorably some method of securing funds for such a monu-

ment. On discussion it appeared that there was no way, according

to the Constitution, by which the proceeds of the Spencer-Tolles

Fund, being a memorial fund, to be used for the encouragement of

research only, could be used for this purpose. Thereupon Dr.

Ewell gave notice of an amendment to Article VII of the Consti-

tution changing the clause referring to the use of the income of

this Fund by adding the words, “or for the erection of any appro-

priate memorial to the persons in whose honor the fund is named,”

and the Secretary was instructed to secure from the members of

the Society an expression of opinion as to the desirability of so

modifying the Constitution.* In view of the fact that a year would

have to ensue before this amendment could be acted upon, the Ex-

ecutive Committee recommended the appointment of a committee

authorized to solicit funds for the purpose proposed, which recom-

mendation was favorably received, and the President appointed

Drs. M. D. Ewell and W. C. Krauss.

Mr. J. D. Hyatt, of New Rochelle, New York, was elected an

honorary member, in recognition of long and faithful service in the

Society.

On motion, the Society voted to loan Dr. Ewell “Centimeter

A” for the purpose of investigation, the Custodian being authorized

to take his receipt therefor.

Adjourned at 7 P. M.

SECOND SESSION

At the second session, called to order at 8 Pp. M. of the same

day, the following papers were read:

*Correspondence showed a practically unanimous opposition to the change,

so the matter was dropped.

PROCEEDINGS 217

“The Ecological Relations of a New Carnation Mite,” by Dr.

R. H. Wolcott.

“The Identification of Blood Stains,” by Prof. W. F. Watson.

“The Pietzsch Microtome,” by Mr. Edward P. Dolby.

“Two New Pieces of Apparatus,’ by Mr. A. B. Porter, pre-

sented by Dr. M. D. Ewell.

The following papers were read by title:

Variation in the Vitellaria and Ducts of Three Species of the Genus Opis-

thorchis, by Prof. F. D. Barker.

Vertical Distribution of Plankton in Flathead Lake, by Prof. M. J. Elrod.

On the Cladocera of Nebraska, by Prof. Charles Fordyce.

A biological study of the lakes of the Pike’s Peak region, by Dr. H. L. Shantz.

On the occurrence of Trypanosoma in the blood of Rana clamata, by Mr.

James H. Stebbins, Jr.

A new Linguatulid from some of our Gulls, by Prof. Henry B. Ward.

The need of a trained microscopist in every community, by Dr. William C.

Krauss.

Permanent preparations of tissues and organs to show glycogen, by Prof.

S. H. Gage.

Adjourned.

THIRD SESSION

The third session was called to order at 8:45 a. M., June 30.

The auditing committee on the Custodian’s report reported favorably

and the report was adopted. The auditing committee on the Treas-

urer’s report was continued and instructed to present the report,

when ready, to the Secretary for publication.

The nominating committee recommended the following persons

for election to the offices named:

President, Dr. Marshall D. Ewell, Chicago, Il.

Vice-Presidents, Prof. Herbert Osborn, Columbus, Ohio; Prof.

William H. Watson, Greenville, S. C.

Custodian, Mr. Magnus Pflaum, Pittsburgh, Pa.

Executive Committee, Dr. A. T. Kerr, Ithaca, N. Y.; Mr. F.

W. Kuehne, Fort Wayne, Ind.; Prof. B. L. Seawell, Warrens-

burg, Mo.

The report was adopted and the Secretary instructed to cast

the vote of the Society for the persons named.

The usual allowance was voted for the expenses of the Secre-

tary.

918 PROCEEDINGS

The Society then adjourned sine die. However, the members

joined in the meeting of Section F of the American Association for

the Advancement of Science, held in the afternoon of the same day.

During the meeting the members participated in all the social

entertainments tendered the other scientific bodies and enjoyed to

the full the very generous hospitality extended by Cornell Uni-

versity.

Rosert H. Wo tcutt, Secretary.

TREASURER’S REPORT

FROM JUNE 18, 1906, TO JANUARY 1, 1907

DR.

eee aticestcoral last, RepOck-co:. aca tne Meteone ten. ss $319 25

Mt MCHIDETSHIP IES, TIDAL... icsiccedsaersiqievsisialsvceyast weiss es nae $ 800

Mom Vembexship Guess 1905 < hac cei agsinolete ods. s cca c cvns oa e-s 26 00

Momvlenibership: dies: 1906: 4.2 con ccc Gases be cwec css ot 188 00

meeembership: dues, W907) oie soles oes ba less td 22 00

MomMembership:, diuess 190805. ile. . She boss hem Jae 400 248 00

“dh JaCL SUERY TCT RS 22) Se a 21 00

DO SUIISEEIBEES SW. OV eee o.oo io os eveseseiovsv ores ote ate 38 00

Memsumscripers, VOL MM VIT.. ol. oo. cc cece ece echoes ORE 6 00 44 00

SMEAR ACNCEETEES SOLE 5 i) cosy 5 oad ns secoale eee CR Py ds

memadvertisers, Vol, XVI... . 24 Mee eee Sas es 80 00

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mM as AIONy |. he inv eae ek Fn OL, eres}. 20 00

CR. $738 50

By Postage, Stationery and Printing, Secretary.......... $ 34 26

Eeapesiices hreasuier. 3 ft are eos ce te ee ates 5 00 $ 39 26

ieee XMLESSAGe, SCCLELARY:. oc. 0s oh vehagree « bec avaule eb een 999

Pe pLessaee. “lb reasuien: oo}. 2's | ecu need mane oe 150 11 49

iy hypewritnic Secretary. <2 w Se. ove 2c ee eee as 14 30

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By Fare to Ithaca Meeting, Secretary...................- 25 00

Rageimtine, Vol. xox Vil... 2.4 do eee os hee 508 30

By Bank Charges for Collecting Checks, Treasurer...... 200

Seta ECE SORE TANG 55 2 os. og ce Se Sarak ata ee ee 129 45

$738 50

J. C. Smitu, Treasurer

We hereby certify that we have examined the Treasurer’s books and

vouchers and found the foregoing account correct.

( Signed)

Henry B. Warp,

CHARLES ForDYCE,

Auditing Committee

CUSTODIAN’S REPORT FOR YEAR ENDING JUNE

20, 1906

SPENCER-TOLLES FUND

Reported ‘at. Sandusky, Mectiag: 9: 02s se. ss an oe oe old aes ae ee $2,081 09

Dividends: Retaivedy seus eee te tei he: RS ee 149 44

Hite Memberships chasse brs Chew isd Sa Seen we SOSA ee ee 100 00

Contribution inden ccc. coSa kis Gacrourswn's aincls Geis AOGR te ee ee 50

$2,331.03

Ress Duesitom Life Membership: .....22cce-> nee nee $6 00

1Se55 = AP EHSESites hoses Re ee eee Re ee eee eee 25 6 25

$2,324 78

MotalMlinvested: .4c.ceey ce wGoevieios sateen hee $2,321 72

GaShy somes etic jaoeeeroaya Syesc Senn oa Chere he RO 306 $2,324 78

Net incredse< diinitte. “yea x6 ou jn.0c eae oes chon eee $ 243 69

GRAND TOTALS

otal Contributions: to. ‘date. .\,. <0... 0 24505 o-cet oe ke oe $ 69407

otalstromesalerof Proceedings tol date,.....--.-4 4-45. oe eee 570 73

Potal Wnterest and! Dividends: to date. 2... 4 2.2-.<caee +) eee 1,025 15

Totaleleife Memberships) to dates. 2... 40.802 = pele oe sone CO ee 150 00

$2,439 95

leesssmGrantaeNon le eee cece ieee iene $50 00

(Grant cI. (25 sorta cho ko hc Sheen ei 25 00

GrantyaNO: 137. ocean cee ne tic se eee 2500 $100 00

Dues on) Lite Memberships... 5-5-0 ee 6 00

Expenses: ..oc:scee ote teeny eee atone 665 coaeenan 917 $11517

$2,324 78

LIFE MEMBERS AND CONTRIBUTORS OF $50 AND OVER.

John Aspinwall A. H. Elliott

Robert Brown John C. Hately

J. Stanford Brown Troy Scientific Association

Henry B. Duncanson Macnus Priaum,

Custodian

IrHaca, N. Y., June 30, 1906

We, the undersigned committee, hereby certify that we have carefully

examined the account of the custodian as given in the foregoing report,

compared the same with the vouchers, and found the same to correspond and

to be correct. ABRAM T. KERR,

MarSHALL D. EWELL,

Auditing Commuttee

CONSTITUTION

ARTICLE [

This Association shall be called the AMERICAN MICROSCOPICAL

Society. Its object shall be the encouragement of microscopical

research.

ArticLe II

Any person interested in microscopical science may become a

member of the Society upon written application and recommenda-

tion by two members and election by the Executive Committee.

Honorary members may also be elected by the Society on nomina-

tion by the Executive Committee.

ArtIcceE III

The officers of this Society shall consist of a President and two

Vice-Presidents, who shall hold their office for one year, and shall

be ineligible for re-election for two years after the expiration of

their terms of office, together with a Secretary, a Treasurer, and

a Custodian, who shall each be elected for three years, be eligible

for re-election, and whose terms of office shall not be coterminous.

ARTICLE IV

The duties of the officers shall be the same as are usual in simi-

lar organizations; in addition to which it shall be the duty of the

President to deliver an address during the meeting at which he pre-

sides; of the Custodian to receive and manage the property and

permanent funds of the Society under the direction of the Executive

Committee and in conjunction with a permanent committee to be

called the Spencer-Tolles Fund Committee, and to make a full and

specific annual report of the condition of all the property, funds,

and effects in his charge; and of the Secretary to edit and publish

the Transactions of the Society.

ARTICLE V

There shall be an Executive Committee, consisting of the offi-

cers of the Society, three members elected by the Society, and the

past Presidents of the Society and of the American Society of Micro-

scopists who still retain membership in this Society.

222 CONSTITUTION AND BY-LAWS

ARTICLE VI

It shall be the duty of the Executive Committee to fix the time

and place of meeting and manage the general affairs of the Society.

ArtTIcLe VII

The initiation fee shall be $3, and the dues shall be $2 annually,

payable in advance. But any person duly elected may upon payment

of $50 at one time, or in installments within the same year, become

a life member entitled to all the privileges of membership, but ex-

empt from further dues and fees. All life membership fees shall

become part of the Spencer-Tolles Fund, but during the life of such

member his dues shall be paid out of the income of said fund. A

list of all life members and of all persons or bodies who have made

donations to the Spencer-Tolles Fund in sums of $50 or over, shall

be printed in every issue of the Transactions. The income of said

fund shall be used exclusively for the encouragement and support of

original investigations within the scope and purpose of this Society.

The principal of the fund shall be kept inviolate.

ArticLe VIII

The election of officers shall be by ballot.

ARTICLE IX

Amendments to the Constitution may be made by a two-thirds

vote of all members present at any annual meeting, after having

been proposed at the preceding annual meeting.

BY-LAWS

ARTICLE I

The Executive Committee shall, before the close of the annual

meeting for which they are elected, examine the papers presented

and decide upon their publication or otherwise dispose of them.

All papers accepted for publication must be completed by the

authors and placed in the hands of the Secretary by October Ist

succeeding the meeting.

CONSTITUTION AND BY-LAWS 293

ArTIcLeE II

The Secretary shall edit and publish the papers accepted, with

the necessary illustrations.

ArTIcLE III

The number of copies of Transactions of any meeting shall be

decided at that meeting. But if not decided, the Secretary shall,

unless otherwise ordered by the Executive Committee, print the

same number as for the preceding year.

ARTICLE IV

No applicant shall be considered a member until he has paid his

dues. Any member failing to pay his dues for two consecutive

years, and after two written notifications from the Treasurer, shall

be dropped from the roll, with the privilege of reinstatement at any

time on payment of all arrears. The Transactions shall not be sent

to any member whose dues are unpaid.

ARTICLE V

The election of officers shall be held on the morning of the last

day of the annual meeting. Their terms of office shall commence at

the close of the meeting at which they are elected, and shall con-

tinue until their successors are elected and qualified.

ARTICLE VI

Candidates for office shall be nominated by a committee of five

members of the Society. This committee shall be elected by a

plurality vote, by ballot, after free nomination, on the second day

of the annual meeting.

ARTICLE VII

All motions or resolutions relating to the business of the Society

shall be referred for consideration to the Executive Committee before

discussion and final action by the Society.

ArticLe VIII

Members of this Society shall have the privilege of enrolling

members of their families (except men over twenty-one years of

age) for any meeting upon payment of one-half the annual sub-

scription ($1).

224 CONSTITUTION AND BY-LAWS

ARTICLE IX

There shall be a standing committee known as the Spencer-

Tolles Fund Committee to take general charge of the fund and to

recommend annually what part of the income shall be expended for

the encouragement of research, but the apportionment of the sum

thus set apart shall be made by the Executive Committee.

The Spencer-Tolles Fund Committee shall also have general

charge of the expenditure of such money as may be apportioned,

under the conditions laid down by the Society for its use.

The Custodian shall be an ex-officio member of this committee.

ARTICLE X

The Executive Committee shall have the power annually to

appoint two members to represent the Society on the Council of the

American Association for the Advancement of Science, in accord-

ance with the regulations of the latter organization.

Revised by the Society, July, 1903.

LIST OF MEMBERS

(Corrected to January 1, 1908)

HONORARY MEMBERS

Crisp, FRANK, LL.B., B.A., F.R.M.S., .

5 Landsdowne Road, Notting Hill, London, England

DALLINGER, REv. W. H., F.R.S., F.R.M.S.,

Ingleside Lee, S. E., London, England

LISTE) [a0 Detain en an I 69 Burling Lane, New Rochelle, N. Y.

Warp, R. Hatstep, A.M., M.D., F.R.M.S........ 53) Fourthe St.) hroyse Newye

LIFE MEMBERS

ROMANCE OTANEORD ::), c's cm oclee ross scree 489 Fifth Ave, New York City

ROWING ROBERT )o)c)5-< alsvese oie sna sone cuene Observatory Place, New Haven, Conn.

IDUNGANSON, ROK, HENRY, BE A. Me... 2... State Normal, Peru, Neb.

ISERIORT weROF ARTHUR El. cos cieereee cee one 4 Irving Place, New York City

EVATET Vm] OPIN Cc -ceers ci elas syeicio ta ee reese Chicago Beach Hotel, Chicago, III.

MEMBERS

The figures denote the year of the member’s election, except ’78, which

marks an original member. The TRANSACTIONS are not sent to members in

arrears, and two years’ arrearage forfeits membership. (See Article IV of

By-Laws.)

MEMBERS ELECTED DURING THE YEAR 1906-7

(For address see regular list )

Duncanson, Henry B., A.M. Moe tter, H., M.D.

(Life Member) SHaw, Wo. C.

GREGER, Pror. DARLING K. Soar, C. D., F.R.M.S.

Kriss, H. G., A.B., B.D. WaLKER, Epa R., Ph.D.

LitTERER, Wm., A.M., M.D. WASTENEYS, HarpoLPeH

MarSHALL, RutH, Ph.D. WILLIAMSON, Wo.

ALLEGER, WALTER W., M.D., ’94........ 143 U St., N. W., Washington, D. C.

EEE, NVYNERED EB. A.M:, 04.46. .02 0000055 809 W. 24th St., Kearney, Neb.

AR NOPD MOET JEL. COG= so) secesyc' sic ee 155 W. Concord St., Boston, Mass.

INSEIN WATT SOHN! MicAe) MEE: OOS! «5 <cocesco ss oo Lose eee Newburgh, N. Y.

Atwoop, E. S., ’79....Water Witch Club, Highlands, Monmouth Co., N. J.

PREM OOD ETE PB. Jo sissies a elodien aban 16 Seneca Parkway, Rochester, N. Y.

BARCLAY OUTS! P.. M.D.5 05% :e0ievenl eo ok ss 537 St. Paul St., Rochester, N. Y.

Pasnre) Arpert S.,. 97). .6.45...; 1033 Witherspoon Bldg., Philadelphia, Pa.

226 LIST OF MEMBERS

BARKER, FRANKLIN D., A.M.,.’03...... University of Nebraska, Lincoln, Neb.

BAUSCH. EDWARD: 7 USte aaa one 179 N. St. Paul St., Rochester, N. Y.

BAUSCH. EIENRYS- SOs cm irotaaeie es moet 735 N. St. Paul St., Rochester, N. Y.

BAUSCH, IWALL TAME SSH ttae ses! votveniesd os St. Paul St., Rochester, N. Y.

Brar,. PROP JAMES SHARTEEM. 796 «0025. bec acs nse scene Scio College, Scio, Ohio

BEARDSLEY; PRORSe AGEs MOM sea eee: cris eicrorcielc 1412 Tenth St., Greeley, Colo.

Bewi, Avert T., B.S., A.M., ’03..Neb. Wesleyan Univ., University Place, Neb.

Bernt, CuARR ci Siem, Ser. oss vis crtcarte Sees 39 Broadway, New York City

BENNETT, HENRY C., ’93.... Hotel Longacre, 157 W. 47th St., New York City

BensLey, B. A., Ph.D. (Col.), ’05..Biol. Dept. Uni. of Toronto, Toronto, Can.

BERING sO DWAR OO racine ieee siete eee 421 W. William St., Decatur, Ill.

BESSEY, (PROF. CHAREES EDWIN, ED, 110.) 98.0 none seneeee Lincoln, Neb.

BEYER} RORMGEOS Ein OUmia eniemenicnes cre 4422 Coliseum St., New Orleans, La.

BIRGE EP ROR ea AS SCD le De Oe ae 744 Langdon St., Madison, Wis.

BISCOE, PROF HOMAS 3 Diy Oi. t:. a5 .mcmeiuaios 404 Front St., Marietta, Ohio

BiErmes Ae Me AVA), Sil. ee Ohio State University, Columbus, Ohio

BonvINE, Pror. DoNALDSON, ’96........... 4 Mills Place, Crawfordsville, Ind.

Bootu, Mary A., F.R.ML.S., ’82........60 Dartmouth St., Springfield, Mass.

Hoeven; C39. Aci oe ees fee UE ease 3223 Clifford St., Philadelphia, Pa.

IBREDINGH GEOte See O Bertani re aeae 104 Powelton Ave., Lansdowne, Pa.

BROMLEY, ROBERT Unis, M1). > "9S 2.nos silico acts << se ciokieies cae Sonora, Cal.

BrooxoveEr, «Gas, AvBi MiSs 0b. 60). vet aces. Buchtel Coll., Akron, Ohio

Brown, NE; HOWEAND; (Oley ¢ sooe8% wk eens’ 918 Chestnut St., Philadelphia, Pa.

Browpace, A. He, M-D., 94.2.6... aeis's 1073 Bushwick Ave., Brooklyn, N. Y.

BUBEEL VE eb lol Obese 804 Bergen Ave., Jersey City Heights, N. J.

BULL. PAMESUEDGAR GE SOn 02 Geek oor piscine eels 141 Broadway, New York City

IBURRILE, “PROR: hs Jin or nae 278 5 oo. ssc sw ever hones ere eee ee Urbana, III.

BYERS SD) OR Aulus ee eye ee ie 114 W. Second St., Oil City, Pa.

CAID WEED: SOTIS Vem Heh Oat ets ets State Normal, Charleston, II.

CARPENTER. DHOS) BS MaDe 990 te0 esse 533 Franklin St., Buffalo, N. Y.

CARTER, 2) OHIN p> -SOn tee eee 5356 Knox St., Germantown, Philadelphia, Pa.

CrARK sGAVEORD <b) Via 9b) eee oe 619 W. Genesee St. Syracuse, N. Y.

GrARK, «GEORGES PDW:, oVUD= 96 ees eee Genesee St., Skaneateles, N. Y.

CLEMENTS, Freperic E., A.M., Ph:D., "98: >..2..2.0.5.0 ob eee

SI OAR atcte che Slo Geke Sielecearh LONG Univ. of Minnesota, Minneapolis, Minn.

CLEMENTS, Mrs. Epitua Scuwartz, A.M., Ph.D., ’03...... Minneapolis, Minn.

Cocks, Pror. REGINALD S., ’99....McDonogh High School, New Orleans, La.

CoLTon AS Ty MED). 704-5 322.8ssbacc8s8- Ferry and Otis Place, Buffalo, N. Y.

Crospy;’ Cyrus REVAC Bs 08S es okies tome sche 43 East Ave., Ithaca, N. Y.

Coucn,. PRanwers: Gi, “660. heh. ev 3481 Broadway, New York City

Cox, Cras, PER SM ese: Grand Central Station, New York City

CRAIG, "THOMAS; '95.0 85.5 000.654; 597 Sherbrocke St., W., Montreal, Canada

CRANDALL, Geo. C., B.S., M.D., 04........... 4287 Olive St., St. Louis, Mo.

LIST OF MEMBERS 227

as. © 1, MD 99n.. ices ons 2s es ees 209 Locust St., Evansville, Ind.

Dissrow, WituiaM S., M.D., Ph.G., ’01...... 151 Orchard St., Newark, N. J.

Doiy, Enwann P., 06.:.......->.-------25. 948 N. 43d St., Philadelphia, Pa.

RS Oi? Seer ean 293 Hampshire St., Buffalo, N. Y.

Rages tionary, Ph.G., '95...:...---. 22-6: 907 Seventh St., Buffalo, N. Y.

Darescuer, W. E., ’87....... Care Bausch & Lomb Opt. Co., Rochester, N. Y.

Hegeyeeeta, Patio, MD., 02... -...<. 200-00: Hospital, Limon, Costa Rica

Bucay bs O., B.S. 04... 0.22.0. see e ec ee eens 125 E. 25th St., Chicago, Il.

EiGenMANN, Pror. C. H., ’95........... 630 Atwater Ave., Bloomington, Ind.

epidies EATER S., 98.0 662-. 00 vesccrtacee ves 17 Birr St., Rochester, N. Y.

Bema non MORTON 9.9 MLA., MiS., “9B. owe sio.c ema! o)aim vicie o'shoye's rom wine

SS nae: dela re oy ne University of Montana, Missoula, Mont.

Pr oareR JOMN, IMD... (885 vos asian. occede'e« 1014 Fourteenth St., Denver, Colo.

LED Re ie Ld re ee 16 Pearl St., Council Bluffs, Iowa

Bwert, MarsHaut D., M.D., LL.D., ’85........... 59 Clark St., Chicago, IIl.

eee dors Ws EA MDS Me. RS 5700 ips ayaa apes elect eid nse.

A OSRGRE Fee on Pec cee eee Guy’s Hospital, London, E. C., England

BREE CADOEPH,. MCD, "81... <oii50:- ac s ernlericnntace 520 E. Main St., Columbus, Ohio

Bere, Gro. F., M.D.,-F.R.M.S., °78....... 5. 30 Woodlawn Ave., Buffalo, N. Y.

Fettows, Cuas. S., F.R.M.S., ’83..912 Cham. of Comm., Minneapolis, Minn.

BEVGuSON a VIEADE MUS: "PhDs G02. 3% caecccs saeceeees esc Blacksburg, Va.

mec Me Jits WED. FOR cc hs oe ws wea 2 Union Place, Troy, N. Y.

Psat OT SOL Be Dy 22 78 8 ee: Em ea 3212 Pine St., St. Louis, Mo.

PIER INTRO ns aaa akice jane ences Se sie e 644 Broadway, Milwaukee, Wis

iscuer. .CHas, i. Mi, M.D, P°R.M-S., 703°... 52 477 Rice St., Chicago, IIl.

(USES ES aR: eae en Penner Zeiss Optical Works, Jena, Germany

Deere: AMES Mi M.D. 91... cc. a8 Stoneleigh Court, Washington, D. C.

EGE ES G's DET Sa eee ae 202 S. Thirty-first Ave., Omaha, Neb.

Forpyce, Cuarves, B.S., A.M., Ph.D., ’98...Univ. of Nebraska, Lincoln, Neb.

ROSTER EDWARD, - 99 025.355 cnesieic1aser ee The Daily Picayune, New Orleans, La.

BOULTON, EIARRY BR, AGB. (04.5...0.4:-- La. Agric. Exp. Sta., Baton Rouge, La.

Prawiss, HH. W., M.D., PhD. 05... .. U. S. Consulate, Port au Prince, Haiti

icReEEE ERRORS SIMON EH: Db. 95, 82 seecese cadena. 4 South Ave., Ithaca, N. Y.

Gace, Mrs. SUSANNA PHELPS, “87. ........-.:.cce. 4 South Ave., Ithaca, N. Y.

Perea. TRor D. W., Oly... oso ases nes 1332 West Wood, Decatur, II.

ESE R! OR. os c.n clare os oily oe evicia a le oe Te ee Chevey Chase, Md.

Seeeeer Orn WED). 02. 22 foc nse Sonera a ahr feOR Sparta, Kent Co., Mich.

GiLttmorE, Miss GERTRUDE A., B.A., ’03...... 27 Charlotte Ave., Detroit, Mich.

POSS Te RRS pig 1S aS a ee 424 E. Fourteenth St., Oakland, Cal.

GrossxkopF, Ernest C., M.D., ’99. ......187 Thirty-sixth St., Milwaukee, Wis.

GreGER, ProF. Dariine K., ’07............. Westminster College, Fulton, Mo.

228 LIST OF MEMBERS

Haac, D. E., M.D., ’86......--+-+2++-2000e- Liberty Center, Henry Co., Ohio

Hazis Victor S:, OM. ..-:-cpavces<- <2 1911 Webster St., San Francisco, Cal.

HANAMAN, GC. F.PURsMES 79. o. 2s eer State and Second Sts., Troy, N. Y.

EL IN SONG do cs Wad cre ote oie etare cle solo ho eine ae Napier ente ee Charleston, IIl.

HATFIELD, JOHN J. B., ’82.......... 2 eee rece cece cece ees Malott Park, Ind.

Heap, PF. D; Ph.D; "06. Ae. ic. <= University of Nebraska, Lincoln, Neb.

Hertzier, ArTHUR E., M.D., ’96......... 402 Argyle Bldg., Kansas City, Mo.

Hertzoc, MAXIMILIAN, M.D., ’01..Room 800, 103 E. Randolph St., Chicago, IIl.

Hitt, Hersert M., Ph.D., ’87.........----0 +20 24 High St., Buffalo, N. Y.

Hitton, Davin Crarx, A.M., M.D., ’01....-......--- 1240 O St., Lincoln, Neb.

PI GHGE (Be Me Ld tases ce hye le oases eee tee eee eee Jefferson City, Mo.

Horrman,. Jos. H., M.D., ’96....2.-:.-- «++ 111 Steuben St., Pittsburgh, Pa.

Howits, FrepertcK S., Ph.D., ’99....Yale Medical School, New Haven, Conn.

Hormes, A. M:, M.D., ’98.......---+2-+s2008 205 Jackson Block, Denver, Colo.

Hoskins, WM., "79.....-----2e see ceeee Room 54, 81 S. Clark St., Chicago, IIl.

How .anp, Henry R., A.M., ’98........---+++- 367 Seventh St., Buffalo, N. Y.

Humpurey, Pror. O. D., Ph.D., 95..... State Normal School, Jamaica, N. Y.

Ives, FrepErRIC E., ’02......... Woodcliff-on-Hudson, Weehawken P. O., N. J.

Jackson, Dante Dana, B.S., '99.......--. 941 President St., Brooklyn, N. Y.

JoHNSON, FRANK SM.D.,. BORNE Ss, "83. 066 = 2521 Prairie Ave., Chicago, II.

Jones, Mrs. Mary A: Dixon, M.D., F:R-M-S., 98-2... 5-220

cia ta AOR ROR C IDR OD Ee 249 E. Eighty-sixth St., New York City

Jupay, CHANCEY, 700.......06-.0 seen cree eee 610 Lake St., Madison, Wis.

KELLNER, HERMANN, Ph.D., ’04.......-..-+: Spencer Lens Co., Buffalo, N. Y.

RECLOGG: Jose eV, 18. oa ce ree minis 202 Manchester St., Battle Creek, Mich.

Kerr, ABRAM TUCKER, JR, M.D., ’95...........-. Stimson Hall, Ithaca, N. Y.

Kororp, CHartes A., Ph.D., ’99........University of California, Berkeley, Cal.

KOEZ, Ao day Eg Snes cae meinen einisin = lesen ao 32 S. Fourth St., Easton, Pa.

Kraere’ WHEISM, 95.0... 2emeeninss os 411 W. Fifty-ninth St., New York City

KEauss, Wit. G5, MI G0. os oan we 479 Delaware Ave., Buffalo, N. Y.

Kriss, H. G., A.B., B.D., 07. .300 Highland Ave., Chestnut Hill, Philadelphia, Pa.

FIDE SoU og LO eicrete cielo nase © ietheveye eae sins 723 Court St., Fort Wayne, Ind.

DANDACER. Feith, Dita Ole des <ake om. Ohio State University, Columbus, Ohio

Teaiee MCH AEE, 20: UG. so eines Saicuara aetaaoetan 451 Jackson Blvd., Chicago, Ill.

LATHAM, Miss V.-A., M.D:; D-D.S., F.RUM-:S. “882.225-eeee eee

AREA ea VIAL Kine AO ae ea 808 Morse Ave., Rogers Park, Chicago, IIl.

TAWION, AP DWARD. Ey OSs. c otic pc ideaiets ots tases anlar Sheldon Ave., Troy, N. Y.

TGRTPBE:( Vos, EV ARRVA AG thi sscn cet alo a niaiala erin 5 ial epee 336 Pine St., Reading, Pa.

Lewis, Mrs. KATHARINE B., ’89...“Elmstone,” 656 Seventh St., Buffalo, N. Y.

LEWIS; IRA Wee St criion oe hetas ctec, Os tere ciara 406 Galena St., Dixon, IIL.

LIST OF MEMBERS 229

Err, J) Fowarpe DD Ss 782. -... 2.2... 1104 Granite Bldg., Rochester, N. Y.

Paerure line ye Wee, OG. |S ose ies oe cis aw eens ace ves Nashville, Tenn.

eer NEEM ase cele cir.e ol cta)= Svs ro aiaea wi vie)elas Sele een ee = Haverhill, N. H.

Pama Anoery, 92) 22)... ioe See ewes 48 Cumberland St., Rochester, N. Y.

Meee NRY,? B40. 605.2. ween dese boas 48 Cumberland St., Rochester, N. Y.

Boars CHANDLER HST 022022506 eee ess 316 Seneca St., Manlius, N. Y.

Looss, ARTHUR, Ph.D. (Lips.), ’05....Govt. School of Medicine, Cairo, Egypt

OES EC ne) Ding CR Weegee 80 E. Fifty-fifth St., New York City

Evan, Re A., A.M; M:D., 701.......: University of Nebraska, Lincoln, Neb.

vow, Howarp N., M:D., ’84....'.)..... 5% 828 N. Wheaton Ave., Wheaton, IIl.

RS Soha &, (eal gene.) 2 B OMe | 9 Do ee aE Ca ee 1828 Fifth Ave., Troy, N. Y.

MarsHAltt, CoLtins, M.D., ’96........... 2507 Penn. Ave., Washington, D. C.

MAaARSHATHeRUTHG Ph Ds 0%e2.. se: University of Nebraska, Lincoln, Neb.

MASTERMAN, ELMER E., ’97...............- F. R. D. No. 2, New London, Ohio

Riecree. ES NOD Pie DD: (02.05 «0. oa as 5 80 Park Place, East, Detroit, Mich.

MAYWALD, FREDERICK J., ’02........ 1028 Seventy-second St., Brooklyn, N. Y.

MoctartaA. Arpert Ph. D)-"80. 2. 5.0%.8 aes 162 Twenty-fifth St., Chicago, III.

McCraven, Bonner N., ’04.............. 1203 McKinney Ave., Houston, Tex.

pear MOSER IS R490 oe Sa) os Foo SoS Sens 259 Eighth St., Troy, N. Y.

MGM REVS EVASERTIS 85). salva cciste-e! 9 W. Forty-eighth St., New York City

TLD Gn pe): Sie a 319 Fifth Ave., Pittsburgh, Pa.

Mercer, A. CuirForD, M.D., F.R.M.S.,’82..324 Montgomery St., Syracuse, N. Y.

Mercer, FrepericK W., M.D., F.R.MLS., ’83....2540 Prairie Ave., Chicago, IIl.

1 SEED OME DR 23 8 i 1 ee 200 E. State St., Athens, Ohio

MicHENor, Ava, M.D., ’02....... State Training School for Girls, Geneva, III.

Miter, Joun A., Ph.D., F.R.M.S., ’89......... 44 Lewis Block, Buffalo, N. Y.

DS OS oe re Burr Block, Lincoln, Neb.

Moetter, H., M.D., ’07.............341 W. Fifty-seventh St., New York City

Marny nosreT © M.D:, O75 34 << os 2scesideiar 1733 Bonté Ave., Berkeley, Cal.

Lu Se 5S Gag | oe Se ne ae ee ee Woodlawn Ave., Winona, Minn.

Norpianner, A. G. E., M.D., Sc.D., Ph.D., LL.D., ’05

Ci RS DE ATT SD Se Re Rs ea ee cae ee Earl Block, Boulder, Colo.

ROUSE, “Ga UIINGE: TD OS a8 oo rela tn gan Steet ak ee Charlestown, N. H.

mem, Ricnarp J., M:D., ’83; 0. 2.6% j<. <2: 5 York St., East Savannah, Ga.

Beater fers WI), 025, is.s.eees Medical Dept. Univ. of Ga., Augusta, Ga.

Mere SED. ORs 5 aU hoe a al eed 18 Locust St., Portland, Me.

Oszorn, Pror. HERBERT, M.S., ’05..... Ohio State University, Columbus, Ohio

rr Hanvey NA! M.. 03:00 22" .. ..Spencer Lens Co., Buffalo, N. Y.

PARK ROSWELL, A. Ma NMED. 79455) ..:..2. 510 Delaware Ave., Buffalo, N. Y.

ParKER, Horatio N., ’99..... Be ro U. S. Geol. Survey, Washington, D. C.

Parmicn Pea Nic PhD 98 oe he 802 Buchanan St., Topeka, Kan.

230 LIST OF MEMBERS

Pease, PRE Ni; SVs. cece swiss b seen evens ses P. O. Box 503, Altoona, Pa.

Peanoeg. as 019-3256 sence eet ee 3609 Woodlawn Ave., Philadelphia, Pa.

Priaum, Maenus, EsQ., '91...25.. 0500000 440 Diamond St., Pittsburg, Pa.

Porter, ALBERT B,, 06... 0.205. 5-00- eee eeee ees 324 Dearborn St., Chicago, IIl.

Pounn; Restor, Adj PED 708.52 2.2. ces ine enct as sens oem emilee

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INDEX

Acarina, of an Oregon pond, 81;

of the genus Arrhenurus of the

United States, 85; parasitic, from

Portuguese West Africa, 68

Africa, Portuguese West, parasitic

and noxious invertebrates from, 61

Amplifier, for projection microscope,

37.

Apparatus, 211

Arachnida, noxious, from Portu-

guese West Africa, 68

Arrhenuri of the United States, The,

by Ruth Marshall, 85

Arrhenurus, characters of, 85; de-

scriptions of North American

species, 88; distribution of, 86;

divisions of, 86

Arrhenurus acutus, 92; americanus,

126; americanus var. major, 128;

amplus, 122; angustocaudatus, 116;

apetiolata, 113; bicaudatus, 91;

birgei, 97; capillatus, 101; com-

pactilis, 120; cornicularis, 112; cre-

nellatus, 90; dentipetiolatus, 117;

expansus, 107; falcicornis, 121;

fissicornis, 130; flabellifer, 125; in-

fundibularis, 93; krameri, 109;

laticaudatus, 95; laticornis, 122;

longicaudatus, 111; lyriger, 94;

magnicaudatus, 123; major (var.),

128; mamillanus, 98; manubriator,

102; marshalli, 103; megalurus,

105; montifer, 96; ovalis, 90;

parallelatus, 107; pistillatus, 119;

planus, 115; prominulus, 108;

pseudocaudatus, 108; pseudocylin-

dratus, 101; rectangularis, 110; re-

flexus, 117; rotundus, 89; scutu-

latus, 93; scutuliformis, 100; semi-

circularis, 111; solifer, 99; supe-

rior, 124; trifoliatus, 115; wol-

cotti, 118

By-laws, 222

Camera obscura, 10

Cestoda, from Portuguese West

Africa, 66

Cestodes, eggs of, 196

Cladocera, of an Oregon pond, 80

Cladorchis watsoni, egg of, 180

Clonorchis endemicus, egg of, 187

Clonorchis sinensis, egg of, 186

Coleoptera, noxious, from Portu-

guese West Africa, 73

Colonies of Volvox, destination of

groups of cells in, 155; dwarf,

male, 156; fine structure of, 150,

165; inversion of young, 158;

nature of, 167; nature of repro-

ductive contents of, 145

Condenser, substage, 26

Condensers, for projection micro-

scope, 25

Constitution, 221

Copepoda, of an Oregon pond, 80

Custodian’s report, 220

Data for the Determination of Hu-

man Entozoa, II, by Henry B.

Ward, 177

Determination of sex, in Volvox, 168

Diptera, parasitic and noxious, from

Portuguese West Africa, 70

Eggs, of human entozoa, 177

Entozoa, human, data for the de-

termination of, 177; eggs of, 177

Eudorina elegans, Pleodorina illi-

noiensis a variety of, 144

234

Fasciola gigantea, egg of, 180

Fasciolopsis buskit, egg of, 181

Fasciolopsis rathouisi, egg of, 182

Fauna, microscopic, of an Oregon

pond, 75

Further Studies in Volvox, with De-

scriptions of three New Species,

by J. H. Powers, 141

Gage, Simon Henry, Permanent

Preparation of Tissues and Or-

gans to show Glycogen, 203;

President’s Address, The Origin

and Development of the Projec-

tion Microscope, 5

Glycogen, preparation of tissues and

organs to show, 203

Hanausek, T. F., and Winton, A. L.,

Microscopy of Technical Products,

The (review), 213

Heat, removal of, in projection mi-

croscope, 28

Hemiptera, noxious, from Portu-

guese West Africa, 70

Hydra, from an Oregon pond, 79

Hydrachnids, of an Oregon pond, 82;

of the genus Arrhenurus of the

United States, 85

Hymenoptera, noxious, from Portu-

guese West Africa, 73

Insect larvae, of an Oregon pond, 81

Insecta, noxious, from Portuguese

West Africa, 69

Invagination, in Volvox, 158

Invertebrates, parasitic or moxious to

man, collected in Portuguese West

Africa, 61

James, Frank L., sketch of, 207

Kuehne, F. W., sketch of Rudolph

Siemon, 210

INDEX

Lenses, invention and development

of, 8

Lepidoptera, noxious, from Portu-

guese West Africa, 70

Light, arc, 21; Drummond, 20;

lime, 18

Lighting, for projection microscope,

Macrothrix laticornis, a new variety

of, 81

Magic lantern, 10

Marshall, Ruth, The Arrhenuri of

the United States, 85

Members, list of, 225

Microscope, origin of name, 7; pro-

jection, 5

Microscopy of Technical Products,

The, by Hanausek and Winton,

review, 213

Minutes, of the twenty-ninth annual

meeting, 215

Missouri, Volvox and other forms

from pond at Rocheport, 143

Myriapoda, noxious, from Portu-

guese West Africa, 69

Necrology, 207

Nematoda, from Portuguese West

Africa, 66

Nematodes, eggs of, 196

Notes, 211

Objectives, from projection micro-

scope, 34

Observations on the Micro-fauna of

an Oregon Pond, by Elda R.

Walker, 75

Oculars, for projection microscope,

Officers, list of, 3

Oregon. Micro-fauna, of

Pond, Forest Grove, 75

Organs, Preparations of, to show

glycogen, 203

Todd’s

INDEX

Origin and Development of the Pro-

jection Microscope, The, Presi-

dent’s Address, by Simon Henry

Gage, 5

Orthoptera, noxious, from Portu-

guese West Africa, 69

Ostracoda, of an Oregon pond, 81

Paragonimus kellicotti, egg of, 186

Paragonimus westermanii, egg of,

183

Parasites, from Portuguese West

Africa, 61

Permanent Preparations of Tissues

and Organs to show Glycogen, by

Simon Henry Gage, 203

Pflaum, Magnus, custodian’s report,

220

Photomicrography,

microscope, 46

Physaloptera caucasica, egg of, 197

Pleodorina californica, from Mis-

souri, 143

Pleodorina illinoiensis, a variety of

Eudorina elegans, 144

Pond fauna, of Todd’s Pond, Forest

Grove, Oregon, 75

Powers, J. H., Further Studies in

Volvox, with Descriptions of three

New Species, 141

Preliminary List of Invertebrates,

Parasitic or otherwise Noxious to

Man, collected in Portuguese West

Africa; 1904-1906, A, by F.

Creighton Wellman, 61

President’s Address, The Origin and

Development of the Projection Mi-

croscope, by Simon Henry Gage, 5

Projection, and photomicrography,

46; darkening room in, 40

Projection microscope, amplifier for,

37; blackening, 42; character of,

12; condensers for, 25; darkening

room for, 40; drawin.; with, 42;

for opaque objects, 45; lighting,

with projection

235

16; mechanical parts for, 40;

names for 5; objectives for, 34;

origin and development of, 5; re-

moval of heat of, 28; screen for,

38; use of oculars, 35; water-bath

for, 29

Protozoa, of an Oregon pond, 78;

parasitic, from Portuguese West

Africa, 62

Reproductive cells, in Volvox, ar-

rangement of, 153

Review, 213

Rotifers, of an Oregon pond, 79

Schistosoma haematobium, egg of,

179, 188

Schistosoma japonicum, egg of, 189

Schistosoma mansoni, egg of, 192

Screen, for projection microscope, 38

Screen-distance, in projection micro-

scope, 39

Sex-determination, in Volvox, 168

Siemon, Rudolph, sketch of, 210

Smith, J. C., treasurer’s report, 219

Spencer-Tolles Fund, report of, 220

Sperm-formation, in Volvox, 156

Stain, to show glycogen in tissues

and organs, 203; Weigert’s iron-

haematoxylin, 211

Stebbins, James H., Jr., Weigert’s

Iron-Haematoxylin Stain, 211

Stentor pyriformis, form from Ore-

gon, 78

Subscribers, list of, 232

Temperature, rise in, in water-baths

in projection microscope, 33

Tissues, preparations of, to show

glycogen, 203

Treasurer’s report, 219

Trematoda, from Portuguese West

Africa, 66

Trematodes, eggs of, 180

236

Trichostrongylus instabilis, egg of,

196

Trichostrongylus probolurus, egg of,

197

Trichostrongylus vitrinus,

197

Triodontophorus deminutus, egg of,

197

egg of,

United States, Arrhenuri of, 85

Volvox, arrangement of species of,

170; destination of groups of cells

in, 155; determination of sex in,

168; differentiation of primary re-

productive cells in, 152; hypertro-

phied reproductive development in,

146; independence of sperm

spheres in, 146; invagination in, 158;

inversion of young colonies in,

158; mode of sperm-formation in,

156; nature of, 167; sperm-glo-

boid of, 165; stigmas of, 149

INDEX

Volvox aureus, 148; globator, 166;

perglobator, 162; spermatosphara,

142; weismannia, 152

Walker, Elda R., Observations on the

Micro-fauna of an Oregon Pond,

Ward, Henry B., Data for the De-

termination of Human Entozoa—

LUE alar¢

Water-bath, for projection micro-

scope, 29; stage, 30

Watson & Son’s New Catalogue, 212

Weigert’s Iron-Haematoxylin Stain,

by J. H. Stebbins, Jr., 211

Wellman, F. Creighton, A Prelimi-

nary List of Invertebrates, Para-

sitic or otherwise Noxious to

Man, collected in Portuguese West

Africa: 1904-1906, 61

Worms, of an Oregon pond, 79;

from Portuguese West Africa, 66

TRANSACTIONS

OF THE

American Microscopical

Society

. ORGANIZED 1878 INCORPORATED 1891

PUBLISHED QUARTERLY

BY THE SOCIETY

EDITED BY THE SECRETARY

VOLUME XXIX

1909-10 -

Decatur, ILL.

Review PrInTING & STATIONERY Co.

IQIO

THE AMERICAN MICROSCOPICAL SOCIETY

TRANSACTIONS

AMERICAN MICROSCOPICAL SOCIETY

TWENTY NINE VOLUMES ISSUED

AS A TEACHER, do YOU realize

That you can get systematic papers, including keys, which will

be of great value to your students in the study of the follow-

ing and other groups of organisms?

Nae genera of Water-mites, by Professor Wolcott and

others.

Various genera of Protozoa, of Rotifers, and Polyzoa.

Genera of Diatoms, Desmids, Conjugatae, Protophytes,

Protococcoideae, Phycomycetes, and other Classes, by

Professor Bessey. (This is the best systematic account of

the lower plants available to microscopists.)

Determination of Human Entozoa, by Professor Ward.

AS A PHYSICIAN, do YOU realize

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on medico-biological subjects of great interest to you? Some

of these are: medical and medico-legal microscopy; miscroscope

in diagnosis; parasitology; bacteriology; trypanosomes; blood

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AS A MICROSCOPIST, do YOU realize

That nowhere else in American Biological literature is there

so complete a record of the development of the microscope as

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AS A GENERAL STUDENT OF BIOLOGY do YOU realize

That the Transactions contain many biological papers of the

greatest general value and interest? Among many others:

Professor Eigenmann’s studies on the Eyes of Cave Ver-

tebrates.

Professor Bessey’s address on Evolution in Microscopic

Plants.

Dr. Ewell’s paper on the Use of the Microscope and

Camera in Detection of Forgery.

Dr. Gage’s address on the History of the Projection Mi1-

croscope.

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plasm.

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Address the Secretary,

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OFFICERS.

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ABS SHERTZEMR VED) sy cette, shots nein on: Kansas City, Mo.

Secretary: FRED C. ZAPFFE, M.D......... 3431 Lexington St., Chicago, Ill.

iecusirers Davin: ZOOK ...- 19: ee << Ashland Block, Chicago, Ill.

PUShGUtON- NUAGNUS) PFLAUM . so oclsc senile cee tise aaenaiere Media, Pa.

ELECTIVE MEMBERS OF THE EXECUTIVE COMMITTEE

AO MBE DAUNAUMTAIN sc) Se. «oi cle nants a aie el eeueetats erie sade ele yee RTOs Ne Ne

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eee MOOKEET 050 oss eink ere Beas Chere oid oes incense Lincoln, Neb

EX-OFFICIO MEMBERS OF THE EXECUTIVE COMMITTEE

Past Presidents still retaining membership in the Society

R. H. Warp, M.D., F.R.M.S., of Troy, N. Y.,

at Indianapolis, Ind., 1878, and at Buffalo, N. Y., 1879.

J. D. Hyatt, of New Rochelle, N. Y.,

at Columbus, Ohio, 1881.

ALBERT McCAtta, Ph.D., of Chicago, Ill.,

at Chicago, I1l., 1883.

T. J. Burritt, Ph.D., of Urbana, I11.,

at Chautauqua, N. Y., 1886, and at Buffalo, N. Y., 1904.

Gro. E. FELL, M.D., F.R.M.S., of Buffalo, N. Y.,

at Detriot, Mich., 1890.

MARSHALL D. EWELL, M.D., of Chicago, I11.,

at Rochester, N. Y., 1892, and at Boston, Mass., 1907.

Simon Henry GAGE, B.S., of Ithaca, N.Y.,

at Ithaca, N. Y., 1895 and 1906.

A. CLIFFORD MERCER, M.D., F.R.M.S., of Syracuse, N. Y.,

at Pittsburg, Pa., 1896.

W. C. Krauss, M.D., of Buffalo, N. Y.,

at Columbus, Ohio, 1899.

A. M. BieILe, M.D., of Columbus, Ohio.,

at New York City, 1900.

C. H. EIGENMANN, Ph.D., of Bloomington, Ind.,

at Denver, Colo., 1901.

CHARLES E. Bessey, LL.D., of Lincoln, Neb.,

at Pittsburg, Pa., 1902.

E. A. BrrcE, LL.D., of Madison, Wis.,

at Winona Lake, Ind., 1903.

Henry B. Warp, A.M., Ph.D., of Lincoln, Neb.,

at Sandusky, Ohio, 1905.

The Society does not hold itself responsible for the opinions expressed

by members in its published Transactions unless endorsed by a special vote.

CONTENTS

FASCIOLOPSIS BuSKII, F. RATHOUISI AND RELATED SPECIES IN CHINA.

TEN RY) UB WARD cre sia Se csie seals einte ei evaiersl eh eualeUatete pale eee rr 5

ON THE MORPHOLOGY AND DEVELOPMENT OF A NEW CESTODE OF THE

GENUS PROTOCEPHALUS WEINLAND. GEORGE R. LARUE......... 17

A New SPECIES OF THE TREMATODE GENUS ULLOCREADIUM. IVAN E.

WVUATETETING = Sr Ue ey ee es Sie es eA a ate eM oe ee 50

A Pies ror SyMPoSIUM WoRK. VIDA A. LATHAM.................. 67

REPORT OF COMMITTEE ON ERNEST ABBE DENKMAL.................. 71

In) MEMorRTAM. (C5 'C.) Mrnrog) MM. PEVAUM 27.5 2.) 2) 12 sien 73

FOREWORD

It is now some time ago since it was decided by the Executive

Committee to discontinue the publication of the annual volume of

transactions and to issue in its stead a quarterly. The needs for a

journal of this sort have long been apparent, and there can be no

question that this venture, with such a sponsor as the American

Microscopical Society, will prove successful—not from a financial

standpoint, because that is not considered primarily as essential—

but from a scientific standpoint. It will furnish an excellent means

for the publication of scientific papers recording the researches made

in a branch of science which is always a live and an interesting

one. The fact that this publication will appear regularly every

quarter—January, April, July and October—will be sufficient

assurance for the speedy appearance of contributions.

It is desired to publish only high-class matter, which must be

submitted to and approved by the Executive Committee for pub-

lication. The membership and others are urged to send in their

contributions, which, if accepted, will be published in the order as

received. The rules which governed publication of papers in the

annual volume also apply to the quarterly, and the editor will appre-

ciate a strict observance of them as promoting harmonious colab-

oration.

THe EpItor.

TRANSACTIONS

American Microscopical Society

(Published in Quarterly Installments)

VoL. XXIX DECEMBER 1909 No. 1

FASCIOLOPSIS BUSKII, F. RATHOUISI, AND RELATED

SPECIES IN CHINA

HENRY B. WARD

[WITH TWO PLATES. ]

In 1843 Dr Busk discovered a parasite which he took from the

duodenum of a Lascar sailor who died in the Seamen’s Hospital in

London. Lankester (1857) published the original description of

this parasite, which he named Distoma Buskii. It appears that

the discoverer objected to this appellation and in consequence, when

the specimen was later described by Cobbold (1860), he gave it

the name of Distoma crassum, a name which had been preoccupied

by von Siebold in 1837 for another species of distome. Moreover,

preference has no influence in scientific nomenclature, and the

original specific designation must stand. Of the original specimens

the majority have been lost. One which was given to Leuckart is

figured in his work (1863:586). A second is preserved in the

museum of the Middlesex Hospital, London, and a third in the

museum of the Royal College of Surgeons. Two others are in the

museum at King’s College, making in all only five type specimens

in existence.

The second and third cases of the occurrence of this parasite

were recorded in 1873 by Leidy, to whom the material was sent

by Dr. J. G. Kerr, a missionary physician in Canton. Two others,

under the date of 1875, are reported by Odhner (1902) ; the sixth

case under date of 1878 by Cobbold (1879), and the seventh case

under date of 1890 by Odhner (1902). This last author made a

* Presented at the Sixth Annual Meeting of the American Society of

Tropical Medicine, held at the U. S. Naval School, Washington, D. C.,

April 10, 1909.

6 HENRY B. WARD

careful study of the structure of the species, which meanwhile had

been assigned to the new genus Fasciolopsis by Looss (1899), and to

him we owe our first accurate data with regard to its morphology.

I have found references to cases since that date in India, of which

the record is inaccessible to me, and in China, reported by Heanley

(1908). In addition to these Looss (1907) has examined about a

dozen specimens from Hongkong which are now in the museum of

the Liverpool School of Tropical Medicine. According to Heanley

these came from the pig. Finally Goddard (1907: 196) reports

two cases from China.

From correspondence with Dr. W. H. Jefferys of Shanghai,

China, who cites the opinions of others as well as his own experi-

ments in China, it seems clear that this species is far more abun-

dant and of much greater importance than has hitherto been be-

lieved. I have myself received specimens from Dr. Jefferys and can

confirm the reports of Odhner and Looss with regard to its mor-

phology. The characters of the genus and of the species may be

outlined as follows:

FASCIOLOPSIS Looss, 1899.

Fasciolinae without anterior region clearly set off from rest of

body. Cuticula smooth. Acetabulum powerfully developed, with

cavity extended posteriad as sacculate invagination, and much

larger than oral sucker. Intestinal crura simple, slender, wavy, but

without evaginations. ‘Testes dendritic, with branches growing

smaller toward distal ends. Cirrus pouch very long, cylindrical,

containing spiral tubular seminal vesicle with peculiar caecal appen-

dage. Cirrus closely covered with fine spines. In alimentary canal

of mammals. Type species F. Buskii.

FASCIOLOPSIS BUSKII STILES, 1901.

Syn: Distoma Buskii Lank., 1857; D. crassum Cobbold, 1860,

nec v. Sieb. 1837.

Length 24 to 45 mm., or even 75 mm. (Busk), usually about 30

mm.; breadth, 6 to 16 mm., usually 10 to 12 mm.; maximum

thickness, 1.5 to 4 mm. Body moderately elongated, nearly regu-

larly oval, ventral surface flattened, skin without spines.t Oral

1. Odhner and most others say the cuticula is without any sort of arm-

ature, but Heanley states most positively that it has spines, whether taken

from man or pig, although they are very difficult to find in some mounted

specimens.

FASCIOLOPSIS 7

sucker 0.5 mm. in diameter, completely ventral; acetabulum separ-

ated from anterior end by about its diameter, 1.6 to 2 mm., with

deep triangular lumen extending caudad. Pharynx large, powerful;

prepharynx present. Esophagus very short, intestinal crura slen-

der, extending to posterior end, with two characteristic curves

toward the median line, one at center of body, the other between

the testes. Genital pore immediatly anterior to acetabulum. Cirrus

sac cylindrical, median, prominent, extending from acetabulum

about half way to shell gland. Testes, dichotomously dendritic,

posterior to transverse yolk duct and in median field, one behind

the other. Germarium small, dendritic, just anterior to transverse

yolk duct on the right side. Laurer’s canal present, but receptac-

ulum seminis wanting. Uterus in irregular open coils anterior to

ovary. Vitellaria well developed, with minute acini, extending

from acetabulum to posterior end where they merge, although the

band is nearly interrupted at the posterior end in the median line.

Tiggs very thin shelled, 120-130 microns long by 75 to 80 microns

broad, with minute operculum and finely granular contents. Devel-

opment unknown.

Parasitic in the duodenum of man in India, Siam, China, Assam,

and Sumatra, according to various authors. Very common in South

China pigs (Heanley).

The presence of this parasite is said to be accompanied with high

temperature (to 106° F.), bloody diarrhea, emaciation, tympanic

abdomen, edema, in general, typhoid symptoms. Several of the

cases have terminated fatally, owing, in part, at least, to the late

date at which treatment was sought.

Calomel, thymol with castor oil or salts, and eucalyptus oil have

all been used with success to expel these parasites.

It is important to note here the various records concerning the

occurrence of this species in the United States which are to be

found in the literature. The first of these is given on the authority

of Leidy (1891), where the parasite is listed under the name of

Distomum crassum. In fact, as early as 1873, he reported the

specimens which in this later paper he places in the species named.

The material, as already noted, came from Dr. J. G. Kerr of Can-

ton, China, and was vomited by a Chinese boy. A girl of English

parentage, in Canton, is also recorded as having passed the same

parasite from the bowels. While first recorded in our literature,

8 HENRY B. WARD

it should be noted that the parasites were not obtained on this

continent, but were sent here for determination merely. These are

the only specimens which Leidy reported from the human host.

In the same paper, and immediately in connection with these

specimens of D. crassum from man, Leidy also records that he had

received specimens of this parasite from New York, Arkansas and

Texas, where they were found in the liver of a doe (N. Y.) and of

cattle (Ark. and Tex.). Although Leidy says these specimens pre-

served in alcohol “appear to be identical with D. crassum,”’ and,

though they have frequently been recorded under this name, yet

they do not actually belong to the species under consideration. The

forms in question were in reality Fasciola magna, a species which

is now well recognized as a parasite of American herbivora.

On the other hand, the probable introduction of Ff. Buskiu from

the East, which I predicted in 1903, has been demonstrated in the

discovery of a case by Moore and Terrill (1905). The parasite

was found at the necropsy of a Laskar sailor from an English

steamer, who died at Galveston, Texas, of typhoid fever. This is

the only record of the species for this continent, so far as I can

ascertain. With regard to their record of the parasite, I wish to

call attention to one error which may be the cause of some confu-

sion if not corrected. These authors give the average size of the ova

as 150x75 microns. If the magnification assigned to their micro-

photographs is correct not even the largest reaches this length and

the actual size agrees closely with the measurements of Looss cited

above. A similar error has been made in the case which Goddard

has recently (1907: 195) reported from China; the material was

referred to a committee, and that part of the report of the investi-

gation committee which deals with the eggs reads as follows: Eggs

(possibly immature), size not measured, about one-half that of

Ascaris lumbricoides. Shell, very thin walled. Contents clear,

small and granular; well marked nucleus in center. Nearly spher-

ical. No operculum observed.

An error is certainly present here, for the size of these ova would

equal only 40 to 45 microns by 25 to 30 microns on the basis they

give in their estimate. This is far too small for 7. Buskw. If one

infers that the proportions relative to Ascaris lumbricoides are by

accident reversed in their statement, these ova would approximate

130 to 150 microns by 100 to 116 microns; and this again fails to

FASCIOLOPSIS 9

agree with the species F. Buskii to which they assign these speci-

mens. In fact, the designation “nearly spherical” which they use

to characterize the ova can not by any means be applied to the

figure which Looss gives of ova in F’. Buskw. Renewed investiga-

tions alone can tell whether the authors are in error as regards

their measurements and descriptions or with respect to their iden-

tification of the species.

Recently Barrois and Noc (1908) reported this species as fre-

quent in Cochin China. Among 133 Annamite prisoners, coming

from various provinces, 16 were found on systematic treatment

with thymol to be carriers of F. Buskii. In 36 autopsies, however,

not a single case of infection with this species was discovered.

Among the 16 cases infected the flukes were solitary in 5 instances,

3 parasites were present in each of 3 cases, 4, 5, 6, and 24 parasites

were found in one case each, and finally even 36 flukes in one case.

It is worthy of note that Barrois and Noc think this parasite of

little or no consequence unless present in large numbers. In this

opinion they are at variance with other observers whose opinions are

already cited.

These authors also record the size of the parasite as 25 to 70 mm.

in length by 6 to 12 mm. in breadth and 1.5 mm. in thickness. It

is longer, they continue, than Fasciola hepatica, of which it has not

the leaf-like form, but may be most easily distinguished at a glance

by its considerable thickness. As regards anatomic details, they

merely confirm, in general, the description of Odhner (1902) with

the difference that the ovary and shell gland, in place of being sit-

uated at the middle of the body, as he indicates, are located about

at the union of the anterior and middle third of the body.

In 1887 Poirier described as Distoma Rathouisi a new human

parasite from a single specimen sent him from China by the Rev.

Father Rathouis. According to the record the fluke was passed

by a Chinese woman, 35 years of age, at the mission Zi-ka-wei. The

woman had suffered long from hepatic pains which were not amen-

able to treatment. Hence, Poirier concludes that the specimen

came from the biliary ducts. Poirier gives an extended description

of the anatomy, which he compares in detail with D. hepaticum, the

well known liver fluke, common throughout Europe in sheep and

occasionally also found in man.

10 HENRY B. WARD

For twenty years after its discovery and description Poirier’s

species was not reported again. Many authors, among whom

Leuckart only need be noted, inclined to the view that it was iden-

tical with F. Buskwi, originally reported from India. When the

latter species was carefuliy studied by Odhner his work showed

that the two forms were in all probability not identical. A care-

ful examination of the records also indicated to me their evident

relationship. Accordingly, in a revision of human trematode para-

sites, I (1903) included Poirier’s form as a distinct species in the

genus Fasciolopsis. Of Odhner’s most recent paper (1909) I shall

speak later.

Recently Goddard (1907) reported from Shaohsing, China, two

cases of its occurrence. In the first case, a woman, 42 years of age,

there was a mixed infection with F. Busku. After the administra-

tion of eucalyptus oil, chloroform and castor oil, specimens of both

parasites were passed on three successive days. The patient was

greatly emaciated, and treatment was delayed so long that she died.

An autopsy was impossible. In the second case, a boy, 6 years old,

the eggs were found in the feces and administration of eucalyptus

oil, as before, brought away many parasites at intervals of two or

three days. ‘Two weeks later two specimens were vomited. The

patient is still under observation.

According to Goddard, the parasite is common in that region and

is thought usually to cause death. The cardinal symptoms are en-

larged abdomen, diarrhea, wasting, and occasionally jaundice. The

stools are usually light yellow in color and of a peculiarly offensive

odor. Goddard observed under the microscope groups of bile

stained cells resembling liver cells, sometimes with no definite out-

line, sometimes three or four lobules held together by the enclosing

network of fibrous tissue. Yet he observed no symptoms of liver

involvement. Other cases of the disease are under observation.

The specimens from these cases were submitted to Drs. J. L.

Maxwell and W. H. Jefferys of The Research Committee, who say

that the specimen appears to be D. Rathouisi, though the deserip-

tion of this worm on record differs in some particulars from their

observations. They had at the same time specimens of F. Buskit

from the other case, and note emphatically that they “dissent alto-

gether from the statements of Scheube that these two distomata are

varieties of the same worm. In our specimen their form, size and

FASCIOLOPSIS 11

consistency differ in many particulars.” The ova are important

factors in determining a parasite fluke. Goddard’s account of these

structures is as follows: The eggs of the D. Rathouisi are about

two-fifths of the microscopic field under a one-sixth inch objective

and one inch ocular, have a thin shell and appear to possess a hya-

line body moderately well filled with coarse granules of a greenish

yellow tint in fresh feces. These eggs were present in both cases

reported. One was a case of mixed infection with F. Buskw and

the figures given in this report show a difference in size of the ova.

The report of the Investigation Committee to whom these speci-

mens were submitted is given in the same paper (Goddard 1907:

198). The description of the ova reads thus: Eggs, oval, size not

measured, about one-third larger than Ascaris lumbricoides. Thin

walled and smooth with very small operculum. Contents appear

to consist of large granules.

I have discussed the records of this species in a recent paper

(1908) thus: “From Goddard’s account the size of the ova may be

estimated as approximately 100-115 microns by 65-75 microns.

Such an ovum would be very much smaller than that described by

Poirier for his D. Rathouisi. The Investigation Committee notes

in its report some doubt as to the correctness of the identification,

and the doubt is emphasized by this discrepancy in the size of the

ova. Further study of these forms is needed to establish their true

character.”

Since writing the above account, through the great kindness of

Dr. Jefferys, I have received three specimens of this parasite of

which he writes: “The other worm, of which I can spare you two

specimens, is less certain. Maxwell (Tainan, Formosa) believed it

to be D. Rath., and I think it possibly is so, though I felt doubtful,

as he, too, does now. If the testicles of D. Rathowisi are right and

left, then this is not that worm. Otherwise it corresponds to such

descriptions as given in Braun, for instance. Anatomically it is

much like F. Buskii and appears in the same patients at times.

There are these few differences—it is shorter and thicker in propor-

tion and this factor is constant; that is, if the stool has both kinds

of worms, it is easy and simple to place each worm present in one

or the other group. It is more regularly oval and does not have

wavy edges or lean to one side (See also Goddard’s report). As

you know, there has always been a doubt as to whether or not

12 HENRY B. WARD

these two worms are the same or different. Personally I am inclined

to believe they are the same and that the original description of

D. Rathouisi was faulty with regard to the position of the testicles.

Possibly a difference in date of infection might account for the dif-

ference in size, but why can we not have a large breed of worms

and a small, as a large breed of horses and a small.”

Again, in October, 1908, Dr. Jefferys sent me another lot of the

same species; there were nine specimens, which, he writes, were ob-

tained by Goddard, but gives no further data. At the same time,

he sent a batch of five flukes labeled “probably rathowisi also, passed

with last. Seem different to naked eye. Have just acquired and

not cleared any yet.”

From this material I have been able to make an extended study

of this species. The original description of Poirier is in error at

many points, but this is not strange, in view of the fact that he only

had a single specimen at his disposal. The results of my work may

be summed up in the following specific diagnosis.

FASCIOLOPSIS RATHOUISI WARD, 1903.

Syn. Distomum Rathouisi Poirier, 1887.

Length 15 to 19 mm., breadth 8.5 to 10.5 mm., thickness up to

3 mm.; shape bluntly oval or elliptical, with short cephalic cone,

sharply marked off from body in profile aspect only and usually

bent ventrad and even slightly posteriad. Alcoholic specimens light

erayish brown, sometimes darker. Usually flexed with dorsum con-

cave and edges only very slightly crenate, if at all. Oral sucker

small, subterminal, 0.25 to 0.29 mm. broad by 0.2 mm. in antero-

posterior diameter, with cavity looking ventrad, separated from

ventral sucker by about twice its own diameter. Acetahulum 1.32

2. It is interesting to note how the observer, after having noted and re-

corded clearly the differences in external appearance between the two species,

is led at the close of his report to question their specific identity on the

basis of an argument often advanced and very specious, yet thoroughly

fallacious. The fallacies of this argument are so well exposed by Looss in

various papers (1899, 1907, et alia) that it would not be profitable to go

into them here. I have myself recently (1908) discussed the question with

especial reference to the ova and have shown some of the errors which have

followed in the past from the assumption of a position such as suggested

by Dr. Jefferys. Closely related species of flukes are continually inter-

changed, and the errors in determination resulting therefrom have led in

the past to great confusion.

FASCIOLOPSIS 13

to 1.38 mm. broad by 0.68 to 0.7 mm. in antero-posterior diameter.

Esophagus almost lacking. Internal organs much as in F. Busku,

except in detail. Only prominent differences noted here. Intes-

tinal crura somewhat more irregular and probably with more and

more pronounced curves. Cirrus sac not conspicuous as in /’. Bus-

kii. Testes more compactly branched, broader and denser, than in

F. Buskii, posterior to transverse yolk duct and in median field.

Contrary to the report of Poirier they lie one behind the other.

Germarium on right side, small, coarsely branched. Uterus in

broad, heavy, closely grouped coils, anterior to ovary. Vitellaria

outside of intestinal crura from about level of acetabulum to pos-

terior end where they merge without indication of interruption.

Masses of acini also drift over intestinal crura toward center of

body at several places. Eggs oval, thin shelled, with delicate oper-

culum.

Concerning /’. Rathowisi in China and its effect on the host God-

dard (1907) writes as follows: “This parasite, known to the Chi-

nese as , is quite common here and is thought usually to

cause death, because the Chinese remedy is so obnoxious to the pa-

tient that the parents (for patients are usually children) are too

sympathetic to make them take it. The cardinal symptoms are

enlarged abdomen, diarrhea, wasting, and occasionally jaundice.

The appetite is usually preserved. The cause is supposed to be

the excessive eating of aromatic foods, such as peanuts, or, more

especially, the eating and drinking of all kinds of things at all

times, thus preventing digestion. This produces the worm, so say

the Chinese. I am inclined to think the egg may be carried on

uncooked vegetables or raw fruit, but thus far have failed to

find it.

“The relation of this disease to the liver is interesting. The

stools are usually light yellow in color and of a peculiarly offen-

sive odor. Under the microscope I have found groups of bile-

stained cells resembling liver cells; sometimes with no definite

outline, sometimes three or four lobules held together by the enclos-

ing network of fibrous tissue. Clinically I have observed no symp-

toms of liver involvement.

“Several other cases of this disease are now under observation

and will be reported later on.”

14 HENRY B. WARD

To the original case reported by Poirier may be added the two

noted by Goddard, and one, sent me by Jefferys in 1908, as noted

above, which also comes from Goddard. Two of these are mixed

infections; the one of Goddard (1907) being complicated by F.

Buskw and the last case noted having, as Jefferys writes, another

form, of which more later.

Within a very short time I have seen a paper by Odhner (1909)

in which, after an examination of Poirier’s original specimens

from the Paris museum, he endeavors to demonstrate that D.

Rathouisi represents only a contracted specimen of Ff. Busku. He

rightly points out some errors in the original description of F.

Rathouist which include some items given in the above synopsis,

and yet I can not possibly share his conclusions. Several good

observers in China, among them Jefferys, Goddard and Maxwell

(v.a.), note the differences in the appearance of the living speci-

mens. ‘The worms are equally easily distinguished in the alcoholic

material. The form differs radically and is constant. The surface

is not wrinkled as it would be if noticeably contracted; there is a

sort of cephalic cone here which is absent in F. Busku. The uter-

ine coils are more frequent and more filled with ova. The acini

of the yolk gland are more numerous and somewhat differently dis-

tributed. The suckers are not precisely alike in size, though more

nearly so than indicated by Poirier. In short, the differences, while

slight, are sufficient to constitute the two forms of different spe-

cific rank, and F’. Rathouisi is undoubtedly a good species.

Among the specimens forwarded to me by Dr. Jefferys in Octo-

ber, 1908, were five which he noted as being different to the eye

from F. Rathouisi, although they were passed with the batch of

that species sent me at the same time. They were somewhat larger

than F’. Rathouist, measuring 21 to 22 mm. in length by 9 mm. in

width. The general aspect is of a larger and slenderer worm and

the light yellowish gray color of the alcoholic specimen enables one

to pick them out at a glance. At first thought they appeared as

transitional forms between F’. Rathouisi and F. Buskwi and likely

hence to establish the identity of those two forms which has been

maintained, but further examination revealed the improbability

of this view. The oral sucker is somewhat smaller, the testes differ-

ent in form and the uterus much more closely coiled. Even if one

may suppose that all of these differences are produced by varying

FASCIOLOPSIS 15

contraction, there yet remains a conspicuous difference in the size

of the yolk gland acini which is very striking and apparently beyond

modification by contraction or other change in form. It may be

that this form represents merely an older specimen of F’. Rathouisi,

but reference to Odhner’s figures will show at once that it can

hardly be a younger or less developed form of F. Buskii since it is

in every way fully developed. I am inclined to consider it a new

species to which the name F’. Goddardi may be given. There is

hardly opportunity here to discuss the matter in detail. I expect

te consider it most extensively in a larger paper on these forms now

under preparation. Both figures are drawn from alcoholic speci-

mens and to the same scale.

BIBLIOGRAPHY

Bagrois, Tu., Et Noc, F.

1908. Sur la fréquence du Fasciolopsis Buskii (Lank., 1857) en

Cochinchine. Bull. soc. path. exot., Paris, i, 216-221.

Cosson, T. S.

1860. Synopsis of the Distomide. Jour. Proc. Linnean Soc., Lon-

don, i, 1.

1879. Parasites: A Treatise on the Entozoa of Man and Animals.

London.

GoppARpD, F. W.

1907. Two Rare Fasciolide. China Med. Jour., xxi, 195-198.

HEANLEY, C. M.

1908. A Large Fluke of Man Probably Not Hitherto Described. Fas-

ciolopsis Buskii as a Parasite of Man in Hong Kong; its usual

host probably the pig. Jour. Trop. Med., xi, 122-3.

LANKESTER, E. R.

1857. Appendix B in Kiichenmeister, Manual of Parasites, p. 437.

(Cited after Cobbold, 1879.)

Lerpy, J.

1873. On Distoma hepaticum. Proc. Acad. Nat. Sci., Philadelphia,

1873, 364.

1891. Notice of Some Entozoa. Proc. Acad. Nat. Sci., Philadelphia,

1891, 234-6.

LEucKAktT, R.

1863. Die Menschlichen Parasiten und die von Ihnen Herriihrenden

Krankheiten. Leipzig.

Looss, A.

1899. Weitere Beitriige zur Kenntniss der Trematoden-Fauna Aegypt-

ens, zugleich Versuch einer natiirlichen Gliederung des Genus

Distomum Retzius. Zool. Jahrb., Syst., xii, 521-784.

1907. Some Parasites in the Museum of the School of Tropical Medi-

eine, Liverpool. Annals Trop. Med. and Par., i, 123-152.

Moore, J. T., AND TERRILL, J. J.

1905. Fasciolopsis Busku (Distomum Buskii) ; the first reported case

in the United States; found in a patient dying of typhoid fever.

Jour. Am. Med. Assn., xlv, 1902-3. 1 pl.

16 HENRY B. WARD

ODHNER, TH.

1902. Fasciolopsis Buski (Lank.) (== Distomum crassum, Cobb.) ,

ein bisher wenig bekannter Parasit des Menschen in Ostasien.

Centralbl. Bakt. u. Par., Orig., xxxi, 573-781.

1909. Was ist Distomum Rathowisi? Arch. Parasitol., xii, 467-471.

POIRIER, J.

1887. Note sur une nouvelle espéce de Distome, parasite de ’homme,

le Distomum Rathouisi. Arch. zool. expér. géner., series 2, v, 203.

1 tab.

Warp, H. B.

1903. Trematoda. Wood’s Ref. Handb. Med. Sci., revised edition, vii,

857-874.

1908. Data for the Determination of Human Entozoa II. (Studies

from the Zoological Laboratory, The University of Nebraska, No.

86.) Trans. Amer. Micro. Soc., xxviii, 177-202; 1 pl.

EXPLANATION OF PLATES.

PLATE I.

PLATE II.

(Both figures are drawn from alcoholic specimens and to the same scale.)

PLATE I

Plate 1—Adult of

Vasciolopsis Rathouisi.

—'%

ut,

—

PLATE II

Plate 2—Aduit of Fasciolopsis Goddardi,

ON THE MORPHOLOGY AND DEVELOPMENT OF A NEW

CESTODE OF THE GENUS PROTEOCEPHALUS.*

WEINLAND

[WITH FOUR PLATES. |

GEORGE R. LARUE.

Since Fuhrmann’s work, “Die Taenien der Amphibien,” a consid-

erable number of species have been added to the Genus Proteocepha-

lus, the addition being made from the cestodes of snakes and of

fishes. No further Proteocephalus species have been recorded from

the group of Amphibia.

It is my purpose to describe in this paper the cestode Proteo-

cephalus filaroides, n. sp. from Amblystoma tigrinum Baird. Dr.

J. H. Powers of the University of Nebraska, in carrying on a series

of investigations on Amblystoma tigrinum, found a cestode parasite

occurring in considerable abundance. Some fragments were turned

ever to Dr. Henry B. Ward, who later interested the writer in this

form. Besides the fragments collected by Dr. Powers and placed

at my disposal through the kindness of Dr. Ward, specimens have

been collected from Amblystoma tigrinum in Cherry County,

Nebraska, at Crete, Neb., and at Belleville, Kan. The host is quite

abundant in the region west of the Missouri river, and, since infec-

tion is almost general, the accumulation of a relatively large

amount of material has been fairly easy.

Liihe (1899) gives certain reasons for retaining the name

Ichthyotaenia suggested by Lénnberg (1894) for this group. How-

ever, since the name Proteocephalus Weinland (1858) has the right

of priority, it should be used to denominate the genus.

Material has been killed in formol, corrosive-acetic, picro-subli-

mate-formol, Kleinenberg’s picro-sulfuric with an equal part of 5

per cent. solution of corrosive sublimate to which 5 per cent. acetic

acid has been added, and in other killing and fixing agents. Kill-

ing fluids were generally used hot, and, since the species is small

and not muscular, but little contraction resulted.

* Studies from the Zoological Laboratory, The University of Nebraska,

under the direction of Henry B. Ward, No. 95.

18 GEO. R. LA RUE

GENERAL APPEARANCE. When removed from the small intes-

tine of the host the cestodes are very active, the strobila undergoing

constant changes in length and breadth. The scolex passes through

changes of form with great rapidity. In this respect it fully bears

out its generic name. Before killing, the cestode is white, at times

somewhat transparent. Pieces which have lain for a time in the

rectum of the host frequently become stained yellow or even brown.

In external appearance preserved specimens are long and ex-

tremely attenuated. One notes the long, slender neck region, the

long chain of proglottids which are more or less distinctly set off

from one another, and the extreme length of the mature and ripe

proglottids. A typical specimen yields the following measurements :

Length of body, 80 mm.; number of proglottids, 112; breadth of

scolex, .46 mm.; suckers in antero-posterior diameter, 0.180 mm. ;

length of neck, 34 mm. The following table gives some relative

measurements.

Number of cm. Proglottids

from scolex. in em. Breadth.

1 (includes neck) 39 0.36 mm.

2 26 0.46 mm.

3 14 0.50 mm.

4 10 0.53 mm.

5 6 0.63 mm.

6 5 0.68 mm.

i 6 0.65 mm.

8 6 0.80 mm.

While these figures may be taken as typical, considerable varia-

tion occurs. The length may run up to 110 mm. and possibly

more. ‘The scolex measurements show the least variation, while

the length and breadth of the proglottids show the greatest. Dur-

ing the course of some experiments the writer has taken from the

aquarium jars containing Amblystoma many long pieces of stro-

bila, consisting of ripe proglottids. One piece in particular meas-

ured 78 mm. in length and contained 28 empty proglottids. Another

piece 50 mm. long contained 16 proglottids, of which some single

proglottids measured 4 mm. by 0.75 mm.

The cestode strobila from which the stages of uterine develop-

ment were taken contained about 40 proglottids, in the ripest of

which were a few eggs. The posterior part of the chain was broken

A NEW CESTODE 19

off. A portion of a second strobila which was broken off at either

end contained 90 proglottids, in which the sex organs had attained

a fair stage of development, yet no proglottids contained ripe eggs.

This explains why at times 20-30 ripe proglottids may be passed in

a single string. In this piece of strobila the first proglottid meas-

ured 1.012x0.478 mm., the forty-first measured 0.810x0.626 mm.,

and the eighty-eighth 1.012x0.478 mm.

The scolex (Fig. 9) is small; just distinguishable to the naked

eye. Three scolices give the following measurements of width:

0.366 mm., 0.386 mm., 0.46 mm. Frequently no difference can be,

' detected in the lateral and dorso-ventral diameter, but transverse!

isections show the lateral diameter to be greater. Grooves or fur-

rows may or may not be seen between the suckers, according to the

state of contraction. Grooves have not been seen at the tip. The

apex is conical, without depression and without a fifth sucker.

Neither hooks nor rostellum are present. The four suckers are quite

symmetrically arranged on the broadest zone of the head. ‘The

antero-posterior diameter of the suckers varies from 0.165-0.184

mm., the lateral diameter being slightly less.

The transition of scolex to neck is gradual. The neck, which is

0.30-0.36 mm. in breadth, varies in length from 3.40 mm. to

4.04 mm.

The transition from the neck to the first proglottids is almost

imperceptible. The youngest proglottids are broader than long,

measuring 0.30-0.36 mm. in breadth by 0.10-0.17 mm. in length.

With the growth of the proglottid the relations between the length

and breadth change gradually until in sexually mature or in ripe

proglottids the length much exceeds the breadth. An extreme

length of 4.0 mm. by 0.75 mm. has been noted in empty ripe pro-

glottids, while a length of 1.6 mm. x 0.8 mm. is not uncommon.

An end-proglottid of characteristic shape is found in young and

in maturing cestodes. The posterior end is more or less pointed,

with usually a small indentation at the tip. The end proglottid is

lost with the first shedding of the ripe proglottids, and thereafter

the end of the strobila may round off or may present a more or less

ragged or broken appearance.

The surfaces of well extended proglottids are rarely thrown

into longitudinal or transverse folds, which fact facilitates the de-

termination of the segmentation.

20 GEO. R. LA RUE

HistoLtocy AND MorpHo tocy. In their main features the cuti-

cula and subcuticular structures (Fig. 37) do not vary much from

the current descriptions. The cuticula measures about 8.6 microns

in thickness, but this measurement is not constant over the entire

body. Its surface in well-preserved specimens is never ragged or

broken, but may show minute serrations. Minute pores are present.

The subcuticular muscles consist, first, of a layer of circular

muscle fibers lying next to the cuticula, then a layer of longi-

tudinal fibers. Together these muscles form a layer about 8.5

microns thick. Within the muscular layer are the pyriform sub-

cuticular cells, whose bodies le 5-20 microns from the muscular

layer.

The parenchyma (Fig. 37) is characterized by the occurrence of

large spheroidal spaces, which may reach a length of 30-45 microns.

These large spaces occur within the parenchyma, in both the medul-

lary and cortical regions, the larger spaces in mature proglottids

being grouped near the vitelline glands.

Fresh material treated with a solution of Sudan III in 70 per

cent. alcohol reveals the presence of oil globules whose size and

arrangement agree with the spaces seen in sectioned material.

Osmic acid used as a check yields the characteristic test for fat

globules.

An entire strobila treated with Sudan III shows the arrange-

ment of the oil globules. In the scolex itself no oil globules are

found, or, if present, are extremely small and widely scattered.

In the mid-region of the neck (Fig. 23), about 0.47 mm. from the

scolex tip, hes a large mass of oil globules measuring up to 20

microns in diameter. From the posterior part of the mass, which

in a somewhat flattened specimen is about 0.17 mm. broad, smaller

fat globules extend posteriad in two parallel bands. These bands

broaden out toward the periphery of the strobila, but remain dis-

tinct for several millimeters, after which they fuse in the midfield.

In this anterior region the small oil globules are not densely massed,

but as the proglottid becomes older and approaches the stage of

sexual maturity the globules, which are here much larger, become

closely massed. This massing is apparently due more to an in-

crease in the size of the individual globules than to an increase in

number. In this region the fields about the vitellaria show denser

masses of globules than the middle proglottid area. The outer-

A NEW CESTODE 21

most globules are quite small. These grade over into the larger

globules about the vitellaria and these in turn grade over into

smaller globules in the mid-region of the proglottid. These last

globules, however, are not as small as are those lying outermost.

With the development of the uterus and the uterine outpocket-

‘ings and the production of eggs the larger part of the fat globules

_in the medullary parenchyma disappears. Apparently they are used

up in the rapid metabolic processes incident to egg production.

‘However, not the whole supply of fat is used up in this process, for

even those proglottids which have shed their eggs show some large

oil globules in the region of the vitellaria.

In sections of Proteocephalus Lonnbergi Fuhr. from Necturus

maculatus Raf. determined and studied by the writer similar large

globular spaces within the parenchyma were found whose distribu-

tion agreed with that of the globular spaces within P. filaroides.

_ Sections of an undescribed Proteocephalus species from Micropte-

_ rus salmoides, to which I shall refer in this article to P. sp., do not

show such large spaces as do either P. filaroides or P. Lonnbergu

Fuhr. Since neither fresh nor formol material is at hand no tests

can be made.

In looking over the literature on the subject I find only two ref-

erences to oil droplets in cestodes. R. Leuckart (1879-86: 552)

barely mentions the presence of oil droplets in the lateral fields

of the proglottids of a human cestode, but he does not further de-

scribe their distribution, nor does he point out their significance

in the nutrition of cestode tissues. Schiefferdecker (1874) describes

oil droplets in Dipylidiwm caninum and in Taenia solium. He

describes with some accuracy their distribution in the parenchyma

of the proglottid, but he does not attempt to describe their distri-

bution in the strobila. Minute fat droplets were found in the

cuticula and in the subcuticular cells. The fat droplets of the

parenchyma were larger and were scattered throughout the medul-

lary and cortical parenchyma.

In a rough way I have used fresh material (Dipylidium cani-

num?) as a check upon P. filaroides, and I have demonstrated the

presence of oil globules therein, but I have not attempted to work

out their distribution accurately in the strobila or in the proglot-

1. This form may prove to be Proteocephalus ambloplitis Leidy, which

it very much resembles.

22 GEO. R. LA RUE

tid. Thus far in all my work I have not found oil droplets in

the cuticula and seldom in the subcuticula, except when they were

in the act of being forced out of the proglottid by pressure.

Sections of Taenia serrata (?) show infrequent globular spaces

10-14 microns in diameter in both medullary and cortical paren-

chyma. These spaces, which were undoubtedly left vacant by the

dissolved oil globules, are most numerous in the cortical paren-

chyma of the lateral fields.

So far as I have been able to learn no one else has described

oil droplets in cestodes, this in spite of the many investigations on

their finer anatomy. No one hitherto has found oil droplets in the

Proteocephalidae, nor has anyone hitherto shown the distribution

of oil droplets in the strobila of any cestode, so far as I know. Von

Linstow (1891) has described large parenchyma spaces in Proteo-

cephalus longicollis Rud., but he does not attempt to interpret

them ; they are probably the spaces left vacant by these oil globules.

Blochmann (1896) and Young (1908) describe the subcuticula

and parenchyma and show their intimate relationships with the

cuticula. Neither investigator notes the presence of oil droplets or

the spaces where they occur.

From our knowledge of these structures we must consider the

cuticula to be a membrane through which the foods in solution

pass. ‘Then the subcuticular cells and the processes of the paren-

chyma which extend out to the cuticula absorb these foods (Young

1908). No attempt need now be. made to explain the nature of

absorption. Here synthesis takes place and these products are

then stored in the parenchyma spaces throughout the proglottid.

I shall not undertake an extended discussion of Schiefferdecker’s

theory of fat absorption from the intestinal contents of the host.

His theory supposes the physical capture of food particles by sub-

cuticular cell processes thrust out through pores in the cuticula.

He found his confirmation in the now discredited theory of Than-

») hoffer.2 Studies on assimilation in cestodes are now under way and

in that connection I shall further consider Schiefferdecker’s theory.

Chalk bodies occur in the parenchyma which agree in general

with those described from other Proteocephalus species.

2. Thanhoffer, L. von: “Beitriige zur Fettresorption und histologischen

Struktur der Diinndarmzotten,” in Pfliiger’s Archiv. f. d. ges. Physiol., Vol.

VIII; 391.

A NEW CESTODE 23

MUscULATURE OF SCOLEX AND SrrospitaA. The musculature of

the scolex is fairly complex. As seen in a series of transverse

sections the muscles are arranged as follows: At a depth of about

20 microns from the tip, general oblique muscles (Figs. 13 and 14)

appear, which course from the right and left lateral surfaces to the

dorsal and ventral surfaces respectively forming a rhomboid of

muscle fibers. This rhomboid of fibers is encountered in the

deeper sections until their course is interrupted or diverted by

the increasing size and prominence of the muscular walls of

the suckers which begin to appear at different levels from 60-100

microns from the apex of the scolex. At places the fibers (Fig. 15)

appear to be attached to the sucker wall. The tangentially cut

ends of these fibers may be seen even down to a depth of 200 mi-

crons from the scolex tip.

Beginning at a level of about 30 microns from the apex of the

scolex and persisting at successive levels to the posterior edge of

the suckers may be found a system of dorso-ventral and of trans-

verse (lateral) muscle fibers. Thus a muscle cross (Fig. 14) is

formed within the rhomboid of fibers. The fibers of the muscle

cross are most numerous between the levels of 40-90 microns from

the apex; thereafter their number diminishes, but the individua!

fibers are stronger.

With the appearance of the suckers in the series we find a diag-

onal muscle cross (Fig. 16), whose fibers connect opposing struc-

tures. Each muscle bundle is closely compacted in the middle, from

which point it begins to flare out broadly toward either end, giv-

ing a broad attachment on the sucker wall.

To summarize the musculature of the scolex as seen in the trans-

verse sections, we have, in order of appearance from the tip, first,

the rhomboid of muscles; second, the transverse muscle cross; third,

the diagonal muscle cross, the last two forming the so-called muscle

star (“Muskelsterne,” Riggenbach, 1896).

Liihe (1894) found no rhomboid of muscle fibers in the scolex

of Anoplocephala perfoliata. The muscle relationships in the an-

terior part of the scolex were not similar to those in P. filaroides.

In the region of the suckers he found the transverse and diagonal

muscle crosses as we have it in P. filaroides, but much more strongly

developed. The muscle relationships posterior to the suckers are

again unlike those in P. filaroides.

24 GEO. R. LA RUE

Passing beyond the suckers the muscles immediately take on the

arrangement of the muscles of the neck and of the younger pro-

glottids. In this respect this cestode differs from P. fossata Rig-

genbach in that it does not possess a muscle cross posterior to the

suckers. Fuhrmann (1895) diagrams the scolex musculature of

Nematotaenia dispar Goeze, but the diagram does not agree in all

respects with the musculature of this form. This furnishes addi-

tional evidence that Proteocephalus and Nematotaenia belong to

distinct groups of cestodes.

A study of longitudinal sections through the scolex confirms the

interpretation of the diagonal muscle-cross. Moreover, such sec-

tions give the best idea of the longitudinal muscles, which in gen-

eral are heavier than the transverse fibers. Sections through the

middle areas of the scolex show long fibers (Fig. 17) extending

almost to the tip and terminating at either side of the median line.

A vertical muscle-cross connecting the anterior part of one sucker

wall with the posterior part of the wall of the adjacent sucker, and

vice versa, may be made out (Fig. 18). This structure is fairly

well developed. Strong, heavy, longitudinal muscle fibers contin-

uing the longitudinal muscle system of the body pass from the neck

into the base of the scolex as isolated fibers or, more infrequently, as

small bundles (Figs. 17, 18). Here they find their attachment at

various levels of the lower half of the suckers. A few of these

fibers are attached anterior to the middle region of the sucker. The

arrangement of the longitudinal muscles of the scolex agrees quite

well with Riggenbach’s description of P. fossata Rigg.

In marked contrast to the heavy musculature of the strobila in

the fleshy Proteocephalus species, such as P. ambloplites and P.

torulosa, is the extremely weak development of strobilar muscula-

ture in P. filaroides. Dorso-ventral and lateral muscles are pres-

ent, but are weakly developed. In place of strong bands or plates

of longitudinal muscles, delimiting the cortical from the medul-

lary parenchyma, there are quite isolated fibers of comparatively

small size. Even in contracted proglottids the longitudinal mus-

cles do not present the appearance of being strong and closely set.

ORGANS OF THE Scotex. Reverting now to the structure of the

organs of the scolex we find that the unarmed suckers, four in num-

ber, do vot differ in their histological detail from those of other

Proteocephalus species. There is no terminal sucker or depression.

A NEW CESTODE 25

However, just below the cuticula at this point lies an organ (Figs.

13 and 17), the structure of which has not yet been worked out.

The organ has a length of about 63 microns and a breadth of about

34 microns, a matter of some individual variation. That this is not

a thickening of the tissues of this point is attested by the fact that

it lies within a narrow, clear zone which separates it effectually

from the surrounding parenchyma. Apparently it is not a low-

lying sucker or an invaginated rostellum, for no exterior opening

could be observed. The histology of this organ has not yet been

worked out, but will be undertaken in connection with the mor-

phology, histology, and histogenesis of the plerocercoid of this ces-

‘tode, a study which is now under way.

In this connection it might be said that in the plerocercoid (Figs.

4, 5,6) this organ may be seen with great distinctness, being much

larger than in the adult. In the plerocercoid occurring in the black

bass Micropterus dolomieu Lac.) of Pelican Lake, Minnesota,

this organ finds a very strong development. Unfortunately no adult

specimens of this form are at hand, so no comparison of this organ

in the plercercus and in the adult could be made. The plero-

cercoid is apparently the young form of a Proteocephalus species.

Riggenbach (1896) found a plerocercoid with a large terminal

organ embedded in the scolex of Corallobothrium lobosum. From

its appearance the scolex, I think, belongs to a Proteocephalus spe-

cies. Of this Riggenbach writes: “Sein Scheitel besteht aus einem

langsovalen Korper der aus homogener oder fein granulirter Sub-

stanz zu besetzen scheint. Vielleicht haben wir in diesem gebilde

die Anlage eines Rostellums zu sehen.”

Thus far this end organ has been found only in those species

which are devoid of a terminal sucker and of a rostellum. It is

found in P. ambloplitis Leidy, where it is figured by Benedict

(1900). P. sp. shows it also. P. Lénnbergw studied by the writer

shows a similar structure, much smaller, however, than in P. am-

bloplitis or in P. sp. Riggenbach figures no such organ for P. fos-

sata Rigg., which has a terminal depression, but no terminal sucker,

nor does he figure it for P. abscisa Rigg., which is without terminal

depression or sucking disk. A new species which I shall describe

in detail later has been studied. It has a fifth sucker, but this organ

is not present. With the discovery in snakes of a series of new

Proteocephalus species having either a rudimentary or functional

26 GEO. R. LARUE

rostellum, with or without hooks, comes an added interest in the

scolex relationships. Unfortunately, the writer has not been able

to examine slides of these interesting forms and must rely on the

figures and descriptions of others. That such an organ, espe-

cially if it be small, may be overlooked is attested by the fact that

it has been overlooked in P. Lénnbergiit Fuhr. until discovered

by the writer. A careful survey of the members of the entire

genus would doubtless yield some extremely interesting facts that

perhaps would throw some light upon the relationships of this

genus which is so widely distributed both geographically and in

regard to hosts.

Furthermore, it would be of interest to find out whether we

have here to do with a cestode structure common to many unarmed

species in different genera or whether it be a structure found only

in this one genus and in members which are found only in North

America. Apparently it has never been described in species occur-

ring outside of North America.

Thus far no attempt has been made to assign a function or an

origin to this organ. A solution of the question of origin would

probably throw light upon the question of function. After all we

may be compelled to adopt the dictum of Lithe (1894) that the

fifth sucker is but a modified rostellum; and we may find that we

have to do in this organ with a modified fifth sucker or a much

modified rostellum.

Nervous System. The nerve ring (Fig. 15) occurs at a depth

of about 70-80 microns from the tip of the scolex. In longitudinal

sections this ring may be seen at about the level of the anterior

sucker edge (Fig. 17). The structure is somewhat octagonal in

shape, measuring about 35-40 microns in breadth by 50-55 microns

in length. At its corners nerve processes extend out toward the

suckers and in some cases appear to reach them.

In the strobila the main pair of lateral nerve trunks may be

observed in either longitudinal or transverse sections (Fig. 37) or

in toto mounts (Fig. 2). No accessory nerve trunks have been

made out. The main nerve trunk passes dorsal to the cirrus and

vagina (Fig. 11).

Excretory SysT—eM. The excretory system in the scolex shows

a complex system of anastomosing and coiling ducts. Thence two

pairs of lateral excretory ducts, remaining always within the medul-

A NEW CESTODE 27

lary parenchyma, take a slightly sinuous course posteriad well

within the paired vitellaria (Fig. 2). In the region of the cirrus-

sheath (Fig. 11) the ventral duct passes below and the dorsal

duct above the cirrus and vagina, and in the ovarial region (Fig.

19) they pass dorsal and ventral to the ovaries. Here the ventral

vessels pass between the ovaries and the paired vitelline ducts. In

this region also may be seen the ventral transverse anastomosis.

It seems that a dorsal transverse anastomosis may also exist.

No anastomoses connecting the dorsal and ventral vessels have

yet been demonstrated. The ventral vessel is usually more expanded

than the dorsal, hence its wall appears thinner. Histologically they

seem to be alike.

At regular intervals branching or simple anastomoses (Fig. 10)

arise from either ventral or dorsal vessels, being more numerous

from the ventral vessel in the posterior part of the proglottid.

These anastomoses proceed to the ventral and dorsal surfaces. At

times the anastomosis expands into a little bladder in the region

of the subcuticular structures, then its lumen contracts sharply

and it passes to the exterior as a small canal in the cuticula. Bene-

dict (1900) has shown very similar structures in P. ambloplitis.

I have seen them in both of my new forms and in P. Lénnbergi.

From the fact that these anastomoses to the exterior are second-

ary excretory openings they are called “foramina secundaria.”

However, there exists a wide variation in the types of structures

which are grouped under this name. The “foramina secundaria”

described by Kraemer (1892) for P. ocellata and P. torulosa, and

by Riggenbach (1896) for P. fossata, P. abscisa and Coralloboth-

rium lobosum, are muscular pulsatile vesicles which open at the

posterior lateral margin of each proglottid. Benedict (1900) finds

no such vesicles for either P. ambloplitis or for P. ocellata, nor do

any of the secondary excretory openings seen by me come under

the type described by Kraemer and Riggenbach.

The flame cells may be easily made out in the region contiguous

to the dorsal and ventral excretory vessels. These may form little

fan-shaped groups whose collecting tubules come together to form

larger tubules, which later fuse with other tubules, and these in

turn empty into the main ducts. Many small groups of flame

cells empty directly into the main vessels by means of a common

duct.

28 GEO. R. LA RUE

A fairly careful study has shown almost no flame cells in the

cortical parenchyma, nor are they found in the midfield of the

proglottid. They are relatively less frequent in the posterior part

of the proglottid about the ovaries and they are not present in the

inter-ovarial space. A sketch (Fig. 10) indicates approximately

the area of their most frequent occurrence.

On the distribution of flame cells Braun (1894-1900) yields the

following data: Schiefferdecker (1874) found them thickest be-

tween the transverse musculature of the proglottid; Hamann

(1885) in Taenia litterata almost always only within the trans-

verse musculature, very seldom in the peripheral parenchyma. On

the contrary, Pintner (1881) and Zschokke (1888) reported that

the flame cells lie thickest right here—i.e., in the peripheral pa-

renchyma—and only a few penetrated into the tissue lying within;

they especially accompanied the fibrils of the larger muscle fibers.

In living animals they may be seen most numerously in the head

and in the posterior proglottid, but are here less numerous. In

Bothriocephalus punctatus Fraipont (1880, 1881) found them in

like manner superficial; in Taenia echinococcus especially at the

posterior end and in the scolex of Tetrarhynchus tenuis in the head,

as well as in the posterior end irregularly arranged, while, on the

contrary, in the neck, in the summit of the proboscis sheath and

in the probosces there was a regular arrangement. Zernecke

(1895) saw the majority of flame cells in Jagula in the region

of the outer excretory plexus, also between the subcuticula and the

longitudinal musculature; a number are also found between the

last, also singly even in the medullary layer.

Proteocephalus Lonnbergu Fuhrmann shows the same regional

relationships of the flame cells as does P. filaroides. P. sp. shows

a like arrangement. No other Proteocephalus species have been

investigated on this point.

GENITAL SysTEM. In a general way the genitalia of P. fila-

roides (Figs. 1, 2, 3) agree with the type of genital system for the

genus. Cirrus and vagina have a common lateral genital sinus.

The uterus is a median ventral tube in its early stages; later it has

25-35 lateral out-pocketings on either side. The vitellaria are in

the lateral fields and run the length of the proglottid. The ovary

is bilobed, and is located at the posterior end of the proglottid.

Behind it is the shell gland and odtype. An odcapt (Schluck-ap-

A NEW CESTODE 29

parat) is present. The testes are numerous, lying in two fields

between the vitellaria.

In very young proglottids no sign of genitalia is visible, but in

the region of the second centimeter the anlage of the genitalia may

be seen in toto mounts as a longitudinal median darkly staining

mass of nuclei which extends from the posterior end of the pro-

glotiid anteriad. Near the anterior end of the proglottid this dark

streak turns toward the lateral surface. Riggenbach (1896) gives

a good description of the early appearance of the developing gen-

italia.

The common lateral genital sinus alternates irregularly from

right to left in the strobila. It is situated within the anterior fifth

of the proglottid, and is not marked by a papilla. The vagina

opens anterior to the cirrus.

Mae Orcans. The testes, which number from 70-114, meas-

ure up to 50-60 microns in diameter. In nearly mature proglottids

they are almost spherical, but with their further development they

come to be polygonal on account of pressure of one on another.

They lie between the vitellaria on either side of a free median zone

extending back to the ovarial space. At the extreme anterior end

there is a tendency for the two groups of testes to fuse (Fig. 2).

In ripe proglottids they are pressed clesely against the vitellaria by

reason of the swollen condition of the uterus and its lateral

pouches.

Each testis is covered with a delicate nucleated membrane, the

tunica propria, which is continuous with the walls of a minute vas

deferens of essentially the same structure. The vasa efferentia lie

in a plane dorsal to the testes. These ducts fuse to form larger

vessels (Figs. 1, 21), which are irregularly connected by cross

anastomoses. The larger vessels finally converge to join the larger

vas deferens. The vas deferens arises at a point on the inner dorsal

surface of the dermo-muscular sac near the midfield and at about

the end of the anterior third of the proglottid. Thence the vas

deferens (Fig. 1) makes a few coils in its course anterial to the

cirrus sheath. In mature proglottids the lumen of the vas defer-

ens is small. Its walls are made up of a thick nucleated membrane

surrounded by epithelial cells. A little later the testes begin to dis-

charge their spermatozoa and the vasa efferentia are distended.

These pour their contents into the vas deferens, whose walls dilate

30 GEO. R. LARUE

greatly, and become extremely thin (Fig. 20). The nuclei of both

the inner membrane and the epithelial layer become widely separ-

ated. Only the upper end of the vas deferens retains approximately

its original size and structure (Fig. 22).

In one specimen observed the vas deferens near the cirrus was

greatly enlarged before any spermatozoa were present. This fact

is difficult to account for, but it does not seem to point to a spe-

cific difference. It may have been an abnormality.

Entering the cirrus sheath the vas deferens enlarges somewhat

and now may be called the ductus ejaculatorwus. The ductus (Fig.

2) extends from a fourth to a half the length of the cirrus sheath,

then it doubles back on itself almost to the proximal end of the

sheath, where it is gradually transformed into the cirrus. The

course of the cirrus is then direct to its distal end where its tissues

pass over into the muscular wall of the cirrus sheath. Hence when

protrusion occurs the cirrus is evaginated.

The cirrus (Fig. 25) is lined by cuticula, which is continuous

with that covering the proglottid. Lying next to the cuticula is a

heavy layer of circular muscle fibers, and outside of the circular

muscles is a layer of more weakly developed longitudinal muscles.

Then an epithelial layer follows. Prostate gland cells are numer-

ous on the ductus ejaculatorius and on the cirrus. No retractor

muscles could be demonstrated. The space between the cirrus and

cirrus sheath is filled with a fibrous parenchyma.

The protruding cirrus (Fig. 35) may be frequently seen in ripe

proglottids as a cylindrical fleshy organ, blunt at the tip. There is

no evagination of the cirrus pouch as observed by Schwarz (1908)

in P. Marenzellert Barrois.

The cirrus sheath (Fig. 25) is a thin walled muscular sac, whose

tissues are continuous with those of the cirrus at the distal end.

Its length is about 0.22 mm., its breadth about 0.11 mm., being

broadest in the posterior or proximal half. The musculature con-

sists of weak circular and longitudinal fibers, the circular fibers

lying within the longitudinal. A thin epithelial membrane forms

the outermost layer. At infrequent intervals are gland cells.

FEMALE OrGANS. The vagina (Fig. 2) lies anterior to the

cirrus. From its opening into the common genital sinus it de-

scribes a gentle curve to the midfield of the proglottid, then it

passes directly back to the interovarial space. It lies dorsal to the

A NEW CESTODE 31

median uterus. The size of the vagina varies greatly in different

proglottids, being much larger when distended with semen. This

distension may extend along the whole course of the vagina, or it

may be confined more to the upper and the lower stretches, the

middle part being more nearly normal in size. The broadened pos-

terior part may be called a receptaculum seminis (Fig. 3). It lies

just within the inter-ovarial space. Its posterior end is closely

constricted to form a sort of valve.

The cuticula-like lining of the vagina, which seems to be con-

tinuous with the external cuticula, extends the full length of the

organ to the end of the receptaculum seminis. It is by no means

certain that this lining is a cuticula; it may be a delicate epithe-

lum. In P. Ambloplitis, P. sp. and P. ocellata it bears cilia.

Surrounding this lining is a layer of epithelial cells with occa-

sional gland cells, which are greatly increased in number and in

size in the upper dilated portion of the vagina (Fig. 25). In this

region is a thin layer of circular muscles. Near the opening into

the genital atrium is a weakly developed sphincter. The middle

part of the vagina is lined by a cuticula-like layer, about which

are muscular elements, whose nature could not be determined. The

outermost layer is composed of epithelium and gland cells. The

structure of the lower vagina will be taken up later.

THE ORGANS OF THE INTER-OVARIAL Space. The organs of the

inter-ovarial space (Fig. 3) are arranged on the general plan of

the members of the genus. The ovaries are connected by a mid-

piece which discharges the ova into the oviduct through the odcapt.

The oviduct passes backward in a sinuous course to the post-ovarial

space, then it turns anteriad and passes over to the odtype. About

midway in its course the oviduct receives the vagina, which enters

the inter-ovarial space from the dorsal side. Lying ventral to the

ovaries and at about the level of the odeapt are the paired vitelline

ducts. These unite to form a single vitelline duct, which may be-

come distended to form a vitelline reservoir. The vitelline reser-

voir passes posteriad to the odtype, which it enters at the beginning

of the shell gland. The odtype is surrounded by the shell gland,

the individual cells of which empty into it by means of little tu-

bules. The odtype leads directly into an uterine passage, which

passes more or less directly anteriad to a point some little dis-

32 GEO. R. LA RUE

tance anterior to the posterior end of the uterus. It now discharges

into the dorsal side of the uterus.

This uterine passage persists as a narrow passage and does not

dilate with ova, nor does it form out-pocketings along its course.

Hence it certainly should be distinguished from the uterus proper

and it is manifestly wrong to call the passage an oviduct. The

name oviduct should be retained for the passage between the odcapt

and the odtype. Furthermore the vagina discharges into the ovi-

duct, and not the reverse, as Kraemer (1892) has it. This point

is determined by the histological structure of the organs.

While in general the relationships of the organs of the inter-

ovarial space are as described and as delineated (Fig. 3), some

variations in this plan occur. The vitelline reservoir may lie on

either the right or left side of this space. The courses of the ovi-

duct and of the vagina are not rigidly followed. The state of con-

traction of the proglottid has much to do with the relaxed or con-

tracted condition of the coils of these organs.

Taking up now the histology of the organs in the inter-ovarial

region: That part of the vagina (Figs. 26, 39) extending from

the valve of the receptaculum seminis to the oviduct is composed

of a layer of cubical epithelial cells, surrounding which is a layer

of muscles, the nature of which could not be determined. Then

follows a layer of epithelial cells which may have a glandular

nature.

The odcapt (Figs. 24, 38) is a muscular organ lined with a thin

epithelium and richly supplied with gland cells, whose cell proc-

esses can be traced well into the muscular walls. No boundary

epithelium could be discovered.

The oviduct (Fig. 24) is lined with ciliated cubical epithelium,

surrounding which is a muscular coat, apparently composed of both

circular and longitudinal fibers. Then a layer of epithelium with

scattered gland cells envelops the whole.

The odtype (Fig. 40) seems to be a development of a portion

of the oviduct, for its structure is essentially the same. So far as

could be ascertained, its cubical epithelium bore no cilia. Its mus-

cular walls are more heavily developed than the walls of the ovi-

duct. It possesses a continuation of the oviduct’s external epithe-

lium. The new element is the large mass of single club-shaped

gland cells composing the shell gland. The long slender cell proc-

A NEW CESTODE 33

esses of these cells can be traced well into the muscular layers. The

shell gland covers an area about 63 microns in diameter.

The uterine passage is composed of a layer of epithelial cells,

whose nuclei are quite numerous. A muscular coat can not be

made out. Enveloping the entire organ is a layer of superficial

epithelium.

The paired vitelline ducts (Fig. 36) are composed of a single

layer of flattened epithelium cells, whose nuclei appear as scat-

tered bodies on the surface. It seems that these cells have con-

tractile elements, for the wall has a fibrous appearance. The fibers

run lengthwise of the ducts. At the junction of the vitelline ducts

a new layer or lining of cubical epithelium is found extending

throughout the course of the vitelline reservoir. The presence of

muscle fibers could not be demonstrated.

THs VITELLARIA AND OvaRIES. The vitellaria themselves (Fig.

2) are long, lateral-lying masses of follicles, grouped about an

axial duct, of the same structure as the paired ducts with which

they are continuous. The vitelline cells are large, deeply staining,

spherical or ovoidal cells, with prominent nuclei and nucleoli. In

them oil globules abound.

The bilobed ovaries are long wing-like bodies connected at the

middle by a mid-piece. They are made up of branching and anas-

tomosing swollen tube-like bodies. The margins of the ovaries are

entire. The external covering is a delicate nucleated epithelium.

The ova are large ovoidal or somewhat polygonal cells with prom-

inent nuclei and nucleoli. They stain deeply.

THE UTERUS AND ITS DEVELOPMENT. The early development of

the uterus, and especially the development of the outpocketings

in this genus as preformed organs, was first correctly described by

Schneider (1904) for P. percae Miiller. His account is extremely

brief, but his conception is correct. He says: “Der Langskanal

des Uterus ist bereits angelegt und hat ein weites Lumen, lange

bevor die Production der ersten reifen Hier eintritt. Er. verlauft

an der ventralen Seite als ein Kanal von kreisformigem Querschnitt,

der bereits auf beiden Seiten Auslaiufer (Uterusiste) auszusenden

beginnt.” This view was not held by the investigators on this sub-

ject before Schneider. They considered that the outpocketings were

caused by the breaking down of the uterine walls on account of the

pressure of the contained eggs.

34 GEO. R. LARUE

My own observations bear out Schneider’s statement completely,

and, since the process deserves more complete treatment than he

gives, I shall outline the process of development of the uterus (Figs.

30, 31, 32, 33, 34, 35) and its outpocketings as it was observed

in eleven consecutive proglottids. The drawings are camera out:

lines made from a toto preparation mounted in cedar oil. Later

these drawings were carefully compared with frontal and trans-

verse sections to insure correctness of detail. For convenience I

have numbered the proglottids serially (1-11), beginning with the

one a part of whose uterus is delineated in Figure 30.

As the proglottid approaches maturity the uterus may be ob-

served as a small organ traversing almost the length of the proglot-

tid. It is a slender tube of epithelial cells, whose nuclei are closely

massed and whose cell bodies stain more deeply than do those of

the surrounding parenchyma. At a stage in the proglottid history,

when a few sperms have accumulated in the vas deferens, but none

of which have yet been passed into the vagina, the uterus begins to

take on the appearance shown in Figure 30. It will be noticed that

at irregular intervals the epithelial cells of the uterine wall are

massed together. While leading out into the parenchyma from these

masses are strands of nuclei closely packed in a single row. This

effect can also be seen in the preceding progiottids, but not so mar-

kedly. In them the strands of nuclei are not as long as they are

here. Proglottid 2 shows but little advance in this structure; if

anything the nuclei in the strands are a little more numerous, the

strands a little longer. |

Proglottid 3 shows sperms in the vas deferens. The vagina is

dilating, but there are no sperms in it. The lumen of the uterus

(Fig. 31) is of about the same size as in Figure 30. We now

see that the nuclei of the strands have increased greatly in num-

bers, though at the tips of the strands the nuclei form a single

line. Passing over proglottids 4 and 5 as showing gradual advances

in development, the following one, 6, shows some marked changes

over proglottid 3. Here the lumen of the uterus (Fig. 36) has

increased in size. At the bases of the strands of nuclei the cells

have moved apart and a lumen appears. The outer ends of

the strand remain solid and taper out to single strings of nuclei.

Branches appear on these strands. We have here the first definite

appearance of the lateral out-pocketings of the uterus. In this

A NEW CESTODE 35

proglottid no eggs were present, indeed no sperms had yet entered

the dilated vagina. Proglottid 8 shows the vas deferens full of

sperms, but the much dilated vagina empty. The lumen of the

uterus (Fig. 33) is much larger; the lateral out-pocketings are

longer and have a greater volume. From their tips the double and

single strands of nuclei may be traced well into the parenchyma.

Proglottid 9 shows the vagina containing sperms throughout its

entire length, while many sperms remain in the vas deferens. A

very few eggs are to be seen in the median uterus tube. Plainly,

then, the out-pocketings of the uterus are not due to the pressure

of the contained ova, rather they are preformed organs.

Proglottid 10 shows a few more eggs in the anterior part of the

uterus. Figure 34 shows the extreme anterior end of the uterus

of proglottid 11. Here a number of eggs are seen in the lateral

pockets. They are not yet sufficiently numerous to cause dilation

of the uterus. In this figure the nuclei of the uterine wall, as wel!

as those of the lateral strands, have been omitted. The latter could

still be distinctly traced.

Unfortunately the strobila was broken off here and the posterior

part was missing; hence the drawing of the ripe proglottid (Fig.

35) has been made from a different worm. It will be noticed here

that the lateral pouches have been so dilated that their walls lie

upon each other until we have the appearance of septa. This ex-

treme dilation of the uterine pouches, and, in fact, the expansion

of the uterine pouches in all stages, is apparently due to the absorp-

tion, the breaking down, of the parenchyma and the food material

stored within the parenchyma as fats. These materials are probably

used up in the metabolic processes incident to egg formation.

The method of discharging the eggs has received some attention

by investigators of Proteocephalus species, who, with the exception

of Schneider, described the process as a thinning of the body wall

in the mid-ventral line. Into this space the eggs are crowded until.

with the further growth of the uterus and with development of the

eggs, the body wall becomes thinner and thinner, until it breaks

apart and a cleft is formed. Pressure upon the wall of the uterus

at this point causes a rupture and the eggs escape. Thus we have

a locus minoris resistentiae developed.

However, the process is not so simple as the above description

indicates. The locus minoris resistentiae, or, rather, the series of

36 GEO. R. LA RUE

loci minoris resistentiae, are formed by a process very similar to,

and simultaneous with, the development of the lateral outpocket-

ings. At various levels in the uterus occur outpocketings, which are

directed ventrally between certain pairs of longitudinal muscles.

Such an outpocketing is shown in Figure 29. Here a sharp point is

seen to be directed toward the cuticula. A more frequent appear-

ance of the ventral outpocketing is shown in Figure 27, while Figure

28 delineates a condition of the uterus in the section adjoining the

one from which Figure 27 was drawn. All three figures were taken

from the uterus in the anterior part of the same proglottid. At

least 10-12 separate regions similar to Figure 27 may be found in

the same proglottid, while regions similar to Figure 29 occur more

rarely.

In this condition the ventral diverticula remain for some time,

then as the uterus fills the eggs are crowded out into the ventral

diverticula, as well as into the lateral ones. With increasing pres-

sure of contained eggs in the ripe proglottid, and probably with

a further absorption of tissue materials in the region of the ventral

diverticula, the points on the ventral wall, already undermined by

the ventral outpocketings, give way, and the ventral wall now splits

throughout its length in a definite line, much as a perforated piece

cf paper, when torn, follows the line of perforations. Thus the

egos are discharged.

It should be noted here that, since P. filaroides is not a fleshy

form, the ventral diverticula are necessarily small. In P. sp. the

ventral diverticula are much better developed. Indeed they pass

out between the median pair of strong longitudinal muscles and

there, in the cortical parenchyma, they expand into spheroidal

spaces, which finally come to hold eggs. In this case not so many

ventral outpocketings are formed, there being three to five. But

since the proglottid is much shorter than in P. filaroides a large

number of ventral diverticula are not necessary to produce the

weakened wall. Again in the salamander tapeworm we have 25-35

lateral outpocketings on each side and in the other species but

10-16. There seems to be a correlation between the number of

ventral and lateral outpocketings.

It is a singular fact that no outpocketings proceed toward the

dorsal wall in either species observed by the writer. It is true that

in Benedict’s (1900) description the splitting of the uterus occurs

A NEW CESTODE 37

on the dorsal surface, but an examination of his figures and

descriptions shows an error in orientation and that the species

P. ambloplitis and P. ocellata described by him follow the rule of

the genus. That no dorsal diverticula occur bespeaks a directive

force which is difficult, if not impossible, to explain. Schneider

(1904) attempts an explanaticn by making the single ventral out-

pocketing or “Spitze” of P. percae homologous with the preformed

ventral uterus opening of Bothriotaenia, which is held to be homol-

ogous with that of Bothriocephalus. Whether any such homology

between the uterus openings in Proteocephalus and Bothriocepha-

lus exists is difficult of determination and cannot now be settled.

A further comparative study of the species of Bothriocephalus, the

Bothriotaenia, and the Proteocephalus must be made before the

homology can be pressed. .

Judging, however, from Schneider’s drawings of Bothriotaenia

rugosa it is thinkable that its ventral uterus opening arises in a

manner similar to that in Proteocephalus. The fact that a single

ventral uterus outpocketing is found in P. percae, while several

occur in some of the members of the genus, does not argue against

an homology, for a serial homology is possible.

In P. filaroides the discharge of the eggs may be accomplished

while the strobila is yet intact. More frequently, however, a num-

ber of ripe proglottids break loose from the chain and pass to the

exterior per anum. The majority of the proglottids have begun

the act of discharging the eggs before they reach the exterior. In

this way, when experimental hosts have been well fed on liver,

large pieces of cestodes, consisting of 10-28 empty or nearly empty

proglottids and measuring up to 78 mm. in length, have been

passed. The feces of such hosts frequently contain large quanti-

ties of free eggs.

THE Eac. When discharged from the uterus the egg (Fig. 12)

is covered with three membranes. The outer mucilaginous mem-

brane is very thin and transparent. In diameter it measures from

35-100 microns, every possible gradation between these two meas-

urements being represented. If all Proteocephalus species show

similar variations in the measurements of the outer egg membranes

this measurement is of little value. The second membrane or egg

shell is hyaline, but heavier than the first. It measures from 30-31

microns in diameter, a fairly constant measurement. The third

38 GEO. R. LA RUE

membrane is a delicate hyaline, flexible membrane, just enclos-

ing the six hooked embryo or oncosphere which is developed in the

uterus. Between the second and third membranes is a granular

mass.

The oncosphere, which may be removed from the egg by careful

pressure manipulation on the cover-glass, measures about 21 mi-

crons in diameter. This measurement shows no great variation. It

is still surrounded by the closely investing third membrane. The

three pairs of hooks which were somewhat obscured before now

show up distinctly. In oncospheres which are turned at the proper

angle the three pairs of hooks are seen to be placed in such a way

that two pairs oppose each other at an angle of 180°, while the

cther pair is set midway between them on the quadrant. The

hooks are about 9 microns in length.

If one has some degree of success in making his preparations

some oncospheres will begin to show signs of life. The hooks begin

to go through their characteristic motions. So far as I have been

able to observe, the opposing hooks usually act in unison, while the

third pair alternates with them. More seldom all three pairs of

hooks act simultaneously. In action the opposing pairs bring their

distal ends into proximity, while their proximal ends approach the

periphery. Then the proximal ends move toward each other, while

the distal ends are thus pressed outward from the center against

the investing membrane and downward from the pole. When this

movement is well under way the points of the third pair are

brought to a polar position, and then, about the time of the relaxa-

tion of the opposing pairs, their points are moved toward the

equator. While this alternation of movements is not absolutely

regular, there is a certain rhythm about it which is striking.

With the equatorial movement of the two pairs of hooks the

oncosphere undergoes a change of form from the somewhat spher-

ical to a broadly pear-like form, the anterior pole being the broader.

At the same time the polar diameter is considerably shortened by

an active forward movement of the posterior part of the body.

It has been impossible to diagram the phases of these motions

satisfactorily. But since the writer has also been successful in

watching the movements in the eggs of Hymenolepis nana it seems

that anyone interested in this point could easily watch the process

to his own satisfaction.

A NEW CESTODE 39

INFECTION EXPERIMENTS. A number of infection experiments

have been undertaken using Chironomous larvae, a favorite food of

the secondary host, Daphnia, Cyclops, Notonecta, some larve of

the Dytiscidae, tadpoles of Rana catesbiana, besides the salamander

itself. Thus far all these infection experiments have failed.

That the experiments should fail on the salamander is peculiar,

because, as will be shown later, the plerocercus may develop in it.

Unfortunately the experiments were carried on under most adverse

conditions, when the animals were feeding either not at all or very

poorly at best and when but few eggs could be obtained. The

physiological conditions of the resting animal are probably inimical

to infection.

Schneider (1904) attempted feeding experiments with Gam-

marus locusta, Idotea entomon, Phoxinus laevis, and Gasterosteus

spinachia with no results. The Gammarus died soon after eating

of the eggs (possibly from too heavy infection). The Jdotea and

the fish yielded no results. Since, apparently, he examined only

livers of the fish, he might have overlooked an infection.

Leuckart (1879-86: 464) reports a plerocercus found by Gruber

in Cyclops serratulus which he believed to be the larva of Proteo-

cephalus torulosa Batsch. From a comparison of his figure, which

is drawn to scale, with measurements of P. torulosa it seems that

there is some foundation for this view.

The plerocercus of P. filaroides is even smaller than the one fig-

ured by Leuckart; hence it is not beyond a possibility that Cyclops,

or Daphnia, or Chironomous larve might be intermediate hosts.

Salamanders living under certain conditions are almost always

infected with plerocercoids. The first condition for infection in

the wild salamander is that he must live in an infected pond—i. e.,

the tapeworm must be present in some of the salamanders. During

the summer of 1908 one pond was found whose salamanders were

not infected with either cestodes or plerocercoids. Further data

shows that in infected ponds density of salamander population con-

ditions the amount of plerocercoid infection. The heaviest infec-

tion also exists in salamanders which have remained larval for a

year or more. ‘Thus the element of time increases the chance of

infection.

During the summer of 1908 some larval salamanders of the first

year were obtained from a Kansas pond. Every salamander opened

40 GEO. R. LARUE

at that time possessed a moderate infection of cestodes, but no cysts

containing plerocercoids were noted; they may have been over-

looked. The remaining larve were kept in a large aquarium, where

they underwent metamorphosis. Some of these were put in small

aquarium-jars from time to time, while others were allowed to

remain in the large aquarium. They were trained to feed on liver

and after a short period of feeding ripe proglottids were passed.

When well fed the number of proglottids passed, and hence the

number of eggs discharged, was quite large. In the limited quan-

tity of water in the jars a great opportunity for autoinfection

through the mouth was offered. Yet, whether the infection with

plerocercoids resulted from this period or from an earlier period

could not be ascertained with certainty, because no control experi-

ments could be undertaken at that time.

After a period of 4-6 weeks, during which it was possible for

infection to occur, the salamanders began to die from some cause not

yet ascertained, though a species of Opalina was abundant in many

that died. These dead animals were found to have a moderate infec-

tion with cestodes. Plerocercoids were encysted in all the abdominal

viscera and in the muscles of the abdominal wall. The extent of

the plerocercoid infection in the first few animals examined was

not fully ascertained because of lack of experience in collecting

such material and because the significance of the infection was not

realized. Later animals had a heavy infection. Cysts have seldom

been found in the liver, the organ of heaviest (?) infection accord-

ing to the other investigators. Cysts were most frequent in the

abdominal muscles, less numerous in the submucous layer of the

alimentary tract. They have been removed from the spleen, the

fat bodies, the mesentery, the ovaries, the testes, the pericardium,

and the kidneys. Some plerocercoids have been removed from the

abdominal cavity, where they were moving about freely. Further-

more, some plerocercoids have been removed from the lumen of

the intestinal tract itself when the host had been in captivity

and had fed on a strictly liver diet for three to five months.

Plainly no invertebrate host figured in the development of these

plerocercoids and in their transfer to the alimentary tract.

The plerocercoids (Fig. 6) from the lumen of the intestine are

quite invariably longer than those from cysts and from the abdomi-

nal cavity. There is, moreover, a change in the peculiar end-organ,

A NEW CESTODE 4]

which is much reduced in size in these specimens. Compare Fig-

ures 4, 5, 6, 7, 8, which show a series of five specimens, Figure 4

from a cyst, Figure 5 from the abdominal cavity, and Figures 6, 7,

8 from the intestinal tract. A marked reduction in the size of the

end-organ is noted in Figures 6 and 7, while in Figure 8 no end-

ergan is visible. The end-organ in the adult scolex is small (Fig.

23) and may not always be seen in toto mounts.

The finding of these larve free in the intestinal tract and free

in the abdominal cavity in spite of all precautions to guard against

error and after a period of from three to five months of liver feed-

ing, during which the intervention of invertebrate hosts was impos-

_ sible, hints toward a period of wandering after the plerocercus has |

_ gained his development. That a plerocercus can penetrate the sub-

mucous and mucous layers of the salamander intestine is not beyond

comprehension. Young (1908: 186) considers that the young

larvee of Taenia serrata, after they have lost their hooks, may leave

the cysts to wander out of the liver. In every host well infected

with plerocercoids some cysts must necessarily lie very close to the

inner mucous layer of the intestine. In these cases the possibility

of the plerocercus reaching the intestinal lumen is not too remote to

account for an infrequent infection. Although in one host one

plerocercus was found moving freely in a bloody area under the

peritoneum and in another host one plerocercus was found in a

bloody area in the muscles of the stomach wall, I am not ready to

say that an active migration actually occurs. For the most part

it appears, until more data is at hand, that the majority of the

- encysted larvae are doomed to fail to develop into cestodes. This

would be true except in cases of cannibalism, which cases occur

only rarely among salamanders. My data seem to hint rather

strongly toward direct development, but the possibility and the

probability of invertebrate hosts playing a réle in the infection must

not be overlooked. Indeed, even if direct development be eventually

proven, the invertebrates are not thereby disproven to act as in-

fective agents.

The discrepancy in size between the end-organ of the plerocercoid

from cysts and the end-organ of adult cestodes was so great that it

seemed best to determine by further experiments whether this plero-

cercoid was the larva of our salamander cestodes. Consequently on

Oct. 17, 1908, two salamanders known to be uninfected were anes-

42 GEO. R. LA RUE

thetized, and each was injected per mouth with 16 cysts. Later

one of these animals was put into earth to hibernate, while the other

was trained to eat liver. This last one was fed November 13 with

31 plerocercoids embedded in a piece of kidney. On November 29,

42 days after the first infection and 16 days after the second infec-

tion, the first ripe proglottids were passed. At this time there were

four pieces 10-20 mm. long and 10-12 pieces 2-5 mm. long. The

proglottids and eggs were typical for the salamander cestode. Since

this time a large number of proglottids have been passed, not all of

them being ripe. The host is yet alive, so no idea of the extent

of the infection can be given.

At the time of the second infection, November 13, two other sala-

manders, believed to have a light infection, were fed cysts in pieces

of kidney, the larger of the two receiving 30 cysts, the smaller 24

cysts. Since this time these two salamanders have yielded many

pieces of unripe cestodes and a few pieces containing ripe proglot-

tids. On Jan. 21, 1909, 69 days after the infection, the smaller

specimen died. When examined many plerocercoids were removed

from the body, none from the bony cavity, none from the intestine ;

32 cestodes were taken from the pyloric end of the intestine. These

cestodes were in all particulars like those removed from wild sala-

manders. This is by far the heaviest infection in salamanders

which has come under my observation.

Two days later another salamander died which had come from

the same pond and which had been kept in an adjoining aquarium-

jar under similar conditions except that it had not been fed plero-

cercoids. Thus it served as a control animal. This animal had

no adult cestodes, but three plerocercoids were removed from the

body cavity and three larval cestodes from the lumen of the intes-

tine, besides, of course, the many plerocercoids from the other parts

of the body.

These few experiments with feeding plerocercoids furnish con-

clusive evidence, first, that the encysted plerocercoid is the larval

form of the cestode found in the same host; second, that the period

of development from the plerocercus after ingestion. is short.

The experimental part of the work is necessarily incomplete, but

it is the intention of the writer to continue the experiments in the

hope that the life history of this genus of cestodes may be fully

elucidated.

A NEW CESTODE 43

RELATIONSHIPS OF THE SPECIES. The nearest described species to

Proteocephalus filaroides is Proteocephalus Lonnbergu Fuhrmann,

from Necturus maculatus Raf., a North American perennibranchi-

ate amphibian. In nearly all its measurements P. Lénnbergu

exceeds P. filaroides, but in their general characteristics they are

much alike. Both have the peculiar end-organ, the unsegmented

neck, the relatively thin and flat strobila, young proglottids longer

than broad. In both cestodes the musculature is relatively weak.

The excretory ducts are very similar in their relations to the other

organs. ‘The numerous testes are arranged on either side of a free

median zone. The organs of the inter-ovarial space have similar

relationships. The ovaries of P. Lonnbergw are a little more

branched than in P. filaroides.

These two species are widely separated from Nematotaenia dis-

par Goeze, the only other well-described cestode of amphibia.*

Thus the cestodes of amphibia divide themselves into two groups,

3. Leidy (1851) reported a new species of cestode from Bufo Americanus

which he named Taenia pulchella Leidy. His description, which is inade-

quate for diagnostic purposes, follows:

Taenia pulchella, n. sp.

“White, without any admixture of any other color, variable, usually

broadest anteriorly. Head quadrilateral, subclavate, obtusely rounded,

broader than the neck. Acetabula circular, cupshaped, lateral and op-

posite, sessile, protractile. Neck very long, cylindroid. Articuli containing

several colorless globules; anteriorly subglobular or transversely oval; pos-

terior moniliform, longitudinally oval, or cylindroid and centrally incras-

sate.

Measurements.—Longest, 9 inches (228.5 mm.). Articuli commencing

to be distinctly separate 4 inches (101.5 mm.) from the head. Breadth

anteriorly, 1/4 line (.52 mm.); posteriorly, 1/6 line (.35 mm.) ; anterior

articula, 1/6 line (.35 mm.) long; posterior, 1/4 line (.52 mm.). Acetabula,

1/166 inch (.153 mm.) in diameter. Smallest, 2 inches (50.8 mm.). Head,

1/75 inch (.338 mm.) broad. Articuli commencing distinctly separate 3/4

inch (19.05 mm.) from the head. Broadest part of neck, 1/90 inch (.282

mm.); short distance posterior to the head, 1/125 inch (.203 mm.). An-

terior articuli, 1/100 inch (.253 mm.) diameter; posterior, 1/144 inch

(.176 mm.) long; 1/200 (.127 mm.) broad at extremities, 1/133 inch

(.191 mm.) broad at middle. Acetabula, 1/200 inch (.127 mm.) diameter.

Habitation—Small intestine of Bufo Americanus.

Remarks.—Closely resembles the Taenia dispar. Goeze, found in the Bufo

viridis, ete.,.but it is relatively longer and is never colored.”

It is not possible now to determine whether this be a Proteocephalus

species or whether it belongs with the Nematotaenia or is, in fact, Nema-

totaenia dispar. The latter species has been reported by Stiles and Has-

sall (1894), and by Wright (1879), from North American species of Bufo.

44 GEO. R. LARUE

the one, represented by Nematotaenia dispar, allies itself with the

Taeniadae in M. Braun’s classification, the other group, represented

by P. Lonnbergii and P. filaroides, allies itself with the family Pro-

tcocephalidae as true Proteocephalus species.

The study of this cestode is necessarily incomplete, yet it is hoped

that the facts here presented may prove to be an addition to our

knowledge of the genus Proteocephalus Weinland, and in particu-

lar to our knowledge of those members of the genus inhabiting

amphibia.

The work on this form was carried out in the Zoological Lab-

oratory of the University of Nebraska under the direction of Dr.

Henry B. Ward. To him my sincere thanks are due for the use of his

extensive private library, for the use of material for comparisons,

and for his interest and cooperation. I am also indebted to Dr. J.

H. Powers for assistance in securing preserved and living material

and for his advice in the care of experimental animals.

Benepict, H. M. BIBLIOGRAPHY

1900. On the Structure of Two Fish Tapeworms from the Genus

Proteocephalus Weinland, 1858. Jour. Morph., xvi, 337-368; 1 pl.

BLOCHMANN, F.

1896. Die Epithelfrage bei Cestoden und Trematoden. 12 pp., 2 pl.

Hamburg.

Brawn, M.

1894-1900. Vermes, Abt. 1 b. Cestodes, in Dr. H. G. Bronn’s Klassen

und Ordnungen des Thier-reichs. Leipzig.

Buecr, G.

1902. Zur Kenntnis des Excretionsgefiisssystems der Cestoden und

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FRAIPONT, J.

1880-1881. Recherches sur lVappareil execréteur des Trématodes et

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FUHRMANN, O.

1895. Die Tiinien der Amphibien. Zool. Jahrb. Anat., ix, 207-26; 1 pl.

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thyoténias. Arch. se. phys. nat. Genéve (4), xvi, 335-337; also in

Bull. Soe. Se. nat. Neuchatel, xxxi, 386-88.

GRUBER, A.

1878. Ein neuer Cestoden-Wirth. Zool. Anz., i, 74-75.

HAMANN, O.

1885. Taenia lineata Goeze, eine Taenie mit fliichenstiindigen Gesch-

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KRAEMER, A.

1892. Beitriige zur Anatomie und Histologie der Cestoden der Siiss-

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LEIDY, JOSEPH.

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A NEW CESTODE 45

LEUCKART, R.

1879-86. Die Parasiten des Menschens. 2 Auufl., 827 pp. Leipzig u.

Heidelberg.

Linstow, O. VON.

1891. Ueber den Bau und die Entwickelung von Taenia longicollis

Rud. Ein Beitrag zur Kenntnis der Fischtiinien. Jen. Zeitschr.

fiir Naturwiss., xxv, 565-74; 1 pl.

LONNBERG, E.

1894. Ueber eine neue Tetrabothrium-species und die Verwandt-

schaftsverhiiltnisse der Ichthyotiinien. Centralbl. Bakt. u. Par.,

xv, 801-3.

LuUue, M.

1894. Zur Morphologie des Taenien-scolex. 133 pp. Kénigsberg.

_ 1899. Zur Kenntnis einiger Distomen. Zool. Anz., xxii, 524-26.

PINTNER, TH.

1881. Untersuchungen iiber den Bau des Bandwurmkérpers mit

besonderer Beriicksichtigung der Tetrabothrien und Tetrarhyn-

chen. Arb. a. d. zool. Inst. d. Univ. Wien., iii, 163-242; 5 pls.

RIGGENBACH, E.

1896. Das Genus Ichthyotaenia. Revue suisse de zool., iv, 166-275;

3 pls.

SCHIEFFERDECKER, P.

1874. Beitriige zur Kenntnis des feinern Baues der Taenien. Jena.

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SCHNEIDER, G.

1904. Beitriige zur Kenntnis der Helminthenfauna des Finnischen

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1-34; 1 pl.

ScHWARZ, R.

1908. Die Ichthyotaenien der Reptilien und Beitrige zur Kenntnis

der Bothriocephalen. 50 pp.; 7 pls. Basel.

Stites, C. W.

1893. Ueber die topographische Anatomie des Gefiiss-systems in der

Familie Taenidae. Centralbl. Bakt. u. Par., xiii, 457-65.

Strimzs, C. W., AND HASSALL, A.

1894. A Preliminary Catalogue of the Parasites Contained in the

Collections of the U. S. Bureau of Animal Industry, U. S. Army

Med. Museum, Biological Department of the University of Penn-

sylvania (Coll. Leidy) and in the Coll. Stiles and Coll. Hassall.

Vet. Mag., 1894, 245-53, 331-54.

WEINLAND, D. F.

1858. An essay on the Tapeworms of Man, etc. 93 pp.; 12 figs. |

Cambridge, Mass.

WricHT, R. RAMSEY.

1879. Contributions to American Helminthology I. Proc. Canad.

Inst., i, 3-23; 2 pl.

Youne, R. T.

1908. The Histogenesis of Cysticercus pisiformis. Zool. Jahrb. Anat.,

xxXvi, 183-246; 4 pls.

ZERNECEE, BH.

1895. Untersuchungen iiber den feinern Bau der Cestoden. Zool.

Jahrb. Anat., ix, 92-151; 8 pl.

ZSCHOKKE, F.

1888. Recherches sur la structure anatomique et histologique des

cestodes. Mém. de l'Institut nation. Genévois, xvii, 396 pp; 9 pl.

46 GEO. R. LARUE

EXPLANATION OF PLATES.

All drawings, unless otherwise stated, were drawn with the aid of a

camera lucida, and details were added with the same magnification. The

magnifications given are approximate.

ABBREVIATIONS.

Ci. Cirrus. od. Oviduct.

Cip. Cirrus pouch. oocp. Obdcapt.

cm. Circular muscles. oot. Odtype.

cut. Cuticula. Ov. Ovary.

cvid. Common vitelline duct. rs. Receptaculum seminis.

dex. Dorsal excretory duct. sh. Shell-gland cell.

e0. End-organ. sm. Subecuticular muscles.

ep. Epithelium. t. Testis.

fd. Fat globules. up. Uterine passage.

gle. Gland cells. ut. Uterus.

Im. Longitudinal muscles. va. Vagina.

lop. Lateral uterine outpocket- vd. Vas deferens.

ings. vef. Vas efferens.

mrh. Fibers of muscle rhomboid. cer. Ventral excretory duct.

mst. Muscle star. vi. Vitellaria.

nN. Longitudinal nerve. vid. Vitelline duct.

nr. Nerve ring. vop. Ventral uterine out-pocket-

nst. Strands of nuclei. ings.

PLATE I.

Fig. 1—Schematic diagram of male genital system from the dorsal side.

Reconstruction from camera outlines. X 40.

Fig. 2.—A sexually mature proglottid seen from the ventral surface.

From a toto mount. X 55.

Fig. 3.—Diagram of organs of inter-ovarial space. Reconstruction from

camera outlines, somewhat simplified by displacing the organs. Ventral

side uppermost. XX 250.

Fig. 4—Plerocercoid from a cyst, optical section. The suckers, bo,

drawn within the body. Note large end organ, eo. X 40.

Fig. 5.—Plerocercoid from the body cavity, optical section. X 40.

Fig. 6.—Plerocercoid from intestine, optical section. x 40.

Fig. 7.—Scolex of plerocercoid from intestine, optical section. Note the

reduction in size of the end organ. X 40.

Fig. 8.—Scolex of larva from intestine, optical section. No end-organ

visible. XX 40.

Fig. 9.—Scolex of adult cestode, optical section. X 40.

Fig. 10.—Schematic diagram of the lateral field of proglottid. Redrawn

from a camera outline. The dotted circle indicates the area in which the

flame cells are most numerous. Anastomosis of the ventral excretory duct,

vexa. X 130.

Fig. 11—Diagram similar to Figure 10, showing relation of cirrus,

nerve-cord, and excretory vessels. The cirrus passes between the dorsal and

ventral excretory ducts. X 130.

PLATE I

eof

BeQ,

PLATE II

29 ARN

har td :

A NEW CESTODE 47

PLATE II.

Fig. 12—Egg showing three membranes, m/, m2 and m3; also the onco-

sphere, onc, with its six hooks. X 325.

Fig. 13.—Scolex of adult cestode; cross-section showing rhomboid of

muscle-fibers, mrh, the subcuticular muscles, sm, and the end-organ, €o.

Nuclei are figured in figures 13, 14, 15, 16, 17. X 315.

Fig. 14.—Scolex of adult cestode; cross-section 30-40 microns below tip

showing rhomboid of muscle fibers, also weak transverse and dorso-ventral

muscle fibers. X 315.

Fig. 15.—Scolex of adult cestode; cross-section 60-70 microns below tip

showing nerve-ring, the persisting fibers of the muscle rhomboid, also weak

dorso-ventral and transverse muscle fibers. X 195.

Fig. 16.—Scolex of adult cestode; cross-section 220-230 microns below

tip, showing muscle star, composed of the horizontal muscle cross and the

dorso-ventral and lateral muscle fibers. X 195.

Fig. 17.—Scolex of adult cestode; longitudinal section through oppos-

ing suckers. X 195.

Fig. 18.—Scolex of adult cestode; longitudinal section showing vertical

muscle cross, vme, between adjacent suckers. Two excretory vessels are

seen at ex. X 195.

48 GEO. E. LA RUE

PLATE III.

Fig. 19.—Cross-section of proglottid through the region of the odcapt.

Note the vitelline ducts passing ventral to the ventral excretory duct.

x 80.

Fig. 20.—Cross-section of vas deferens after distention by spermatozoa.

xX 315.

Fig. 21.—Vasa deferentia and their anastomoses, showing frequency of

nuclei on the delicate membrane. X 315.

Fig. 22.—Vas deferens, proximal end in cross-section. Note the two

layers of epithelia. X 315.

Fig. 23.—Head and neck of adult cestode, optical section. Stippled area

indicates area of fat globules. Head flattened. X 50.

Fig. 24.—Odécapt and oviduct. Cubical epithelium bearing cilia, cep.

Longitudinal section. X 630.

Fig. 25.—Cirrus and vagina from a frontal section of a proglottid. Com-

mon genital sinus or atrium, gsi; sphincter muscle of vagina, sphm; ductus

ejaculatorius, dej; prostate gland cells, prgl.. X 315.

Fig. 26—Vagina, musculature somewhat idealized. Cross-section.

630.

Fig. 27.—Uterus and a ventrally directed out-pocketing. From a cross-

section. X 315.

Fig. 28.—Uterus from section adjacent to that figured in Figure 27. No

ventral out-pocketing. X 315.

Fig. 29.—Uterus and ventral out-pocketing. Note the way in which the

ventral wall of the uterus apparently separates the median uterus from the

ventral diverticulum. Figures 27, 28 and 29 from the same proglottid

delineate about the same stage of uterine development as is shown in

Figure 33. X 315.

PLATE III

Shan

i a0 3

PEST Eh a

ope f

bd, Aly Csthey

PLATE IV

0 - —

eA OwoP

P 3

(

A NEW CESTODE 49

PLATE IV.

Figs. 30, 31, 32, 33, 34. Stages in the development of the lateral uterine

out-pocketings, from a toto mount. Note in Figure 30 the simple strands

of nuclei arising from little heaps of cells; in Figure 31 the multiplication

of nuclei and the lengthening of the strands; in Figure 32 the appearance

of a lumen in the strands, and the strands branching. Note in Figure 33

the broader median uterus and the increased volume of the out-pocketings.

Note in Figure 34 the few eggs, o, in the uterus, a septum forming at sep

by the crowding together of the walls of the neighboring diverticula.

Nuclei omitted. X 230.

Fig. 35.—Ripe proglottid. Septa, sep, between the large lateral diver-

ticula of the uterus. Cirrus evaginated. From a frontal section. X 30.

Fig. 36.—Paired vitelline ducts and the beginning of the common vitel-

line duct. X 315.

Fig. 37.—Lateral field of a proglottid, cross-section. Note the large oval

spaces where the fat globules are located; circular and longitudinal sub-

cuticular muscles, sem and sm; subcuticular cells, sct. X 315.

Fig. 38.—Odcapt in cross-section. Note the gland cells and the absence

of a limiting epithelium. XX 630.

Fig. 39.—Lower vagina. Receptaculum seminis, epithelium, cubical

epithelium, cep. X 315.

Fig. 40.—Oétype and shell-gland with the beginning of the uterine pas-

sage in outline; cross-section. XX 630.

A NEW SPECIES OF THE TREMATODE GENUS

ALLOCREADIUM

WITH A REVISION OF THE GENUS AND A KEY TO THE SUB-FAMILY

ALLOCREADIINAE*

IVAN E. WALLIN

[WITH TWO PLATES. ]

Looss (1899), in his work on the trematode-fauna of Egypt and

a natural organization of the genus Distomum Retzius, established

a genus Creadium with the following diagnosis: Worms of sub-

medium size, with a thick, in a contracted condition, almost cylin-

drical, body that diminishes anteriorly into a thin and very flexible

neck part. The posterior end is rounded; suckers greatly devel-

oped; cuticula smooth. Intestine with well developed pharynx,

long esophagus (in a contracted neck it is bent into an 8 shape,

and on this account appears short) ; intestinal coeca long. Gen-

ital aperture near the forking of the esophagus. Cirrus pouch

large, bag-shaped, with well developed cirrus. ‘Testes large and

spherical, median, one back of the other, in the posterior part

of the body. Ovary also large, anterior to the testes and to the

side. Receptaculum seminis and Laurer’s canal present, the for-

mer voluminous, pear-shaped. Yolk glands very greatly developed,

back of the testes they come together and are continuous and com-

pletely fill the body. Uterus very short, between the anterior testes

and the acetabulum, describing a few convolutions. Eggs rela-

tively large (60 to 90 microns), with weakly colored shells, lying

in a row in the uterus, somewhat ventral. The first stage in the

development appears to be completed while in the mother. Inhabit

(freshwater?) fish. Type: Creadium isoporum Lss.

Looss placed Distomwm angusticolle Hausm. in the genus and

pointed out the similarity of a number of distomes described by

Stossich to O. isoporum. He suggested that they might belong to

* Studies from the Zoological Laboratory, The University of Nebraska,

under the direction of Henry B. Ward No. 96.

A NEW TREMATODE 51

different closely related genera, and would then be grouped under a

new sub-family Creadiinae.

The name Creadion, given to a genus of birds by Viellot in 1816,

being very similar to Creadiwm, and according to the usual view

precluding the use of the latter term, Looss (1900) changed the

name to Allocreadium.

Stossich (1900) placed the following species in the genus Allo-

creadium: A. pegorchis Stoss., A. obovatum (Molin), and A. asym-

phyloporum Stoss.

Odhner (1901) revised the genus Allocreadium, placing the fol-

lowing species in it: A. fasciatum (Rud.), A. labri (Stoss.) [later,

1902, found it to be synonymous with A. pulchella (Rud.)], A.

sinuatum (Rud.), A. atomon (Rud.), A. transversale (Rud.), A.

tumidulum (Rud.), A. labracis (Duj.), A. commune (Olss.), and

A, genu (Rud.). Many of the forms that Rudolphi named are diffi-

cult to identify on account of the very meager descriptions and the

poor illustrations, when given; as a result a number of forms de-

scribed by later investigators are synonymous. Odhner examined

some of the original specimens described by Rudolphi, and with

the aid of identical forms that he collected in Sweden he gives com-

plete descriptions and figures of some of the heretofore obscure

species.

In a later paper (1902) Odhner placed A. fasciatum, A. pul-

chella, and A. sinuatum in a new genus, Helicometra. These forms

differ essentially from the species of Allocreadiwm proper in havy-

ing eggs with long unipolar filaments. Helicometra pulchella

(Rud.) was given as the type of the genus.

Stossich (1902) described a new Allocreadiinian form, charac-

terized by eggs with long unipolar filaments, naming it Loborchis

mutabilis; it is synonymous with Helicometra mutabilis. He men-

tions in the introduction of his description that Liihe had intimated

in a personal communication that he was preparing a description

of a new genus, Loborchis, for the forms in Allocreadium, charac-

terized by eggs with long unipolar filaments. Stossich’s form

agreeing with the diagnosis that Liihe was preparing, he placed it

in Loborchis. I have been unable to find this description of Lobor-

chis, or any other references to it, and take it for granted that it

was never published. Stossich (1903) placed H. flava in the genus;

52 IVAN E. WALLIN

in a later paper (1904>) he includes H. gobui and gives a compara-

tive table of all the described species in the genus Helicometra.

In an earlier paper in the same year (1904*) Stossich established

a new genus in the sub-family Allocreadiinae, naming it Lepocrea-

dium. The distinguishing characteristics of the genus follow: Cu-

ticula covered with scales, easily coming off; suckers small and

equal in size, oral sucker terminal; prepharynx present; pharynx

elongated and well developed; genital aperture at anterior edge of

acetabulum. Type: ZL. album Stoss. He also places L. pegorchis

Stoss. in the genus.

There are very few references to Allocreadiinian forms in Amer-

ican literature. MacCallum (1895) describes Distomum isoporum

Lss. var. armatum from Lake Erie and the Grand River of Ontario,

occurring in the small intestine of Aplodinotus grunniens, and in

the intestines of Lepomis gibbosus and Acipenser rubicundus. ‘This

form does not belong in the sub-family Allocreadiinae, as was

pointed out by Looss (1902: 785). Linton (1898) described Dis-

tomum simplex Rud. [synonymous with Allocreadiuwm atomon

(Rud.) |, from Woods Holl, Massachusetts. A single specimen was

taken from the intestine of Microgadus tomcod. He also describes

Distomum pallens Rud. [synonymous with Allocreadium pallens

(Rud.) |], from the same locality in the intestine of Alutera schoepfi.

In the same paper he gives a description of a Distomum, of which

the species was not determined. A single specimen was collected

from Woods Holl, Mass., in the intestine of Lagocephalus laeviga-

tus. The description given places the form in the sub-family Allo-

creadiinae, but it is far too incomplete to establish the exact posi-

tion of the worm. Stafford (1904) catalogues Allocreadium. iso-

porum Lss. from the intestine of Semotilus bullaris. He gives the

dimensions 2.28 by 0.68 mm., which are quite small for A. tsoporum

(5 by 0.75 mm.).

The trematode that is about to be described is one of a collection

made by Prof. Henry B. Ward at Sebago Lake, Maine, in the sum-

mer of 1907. It differs specifically from all heretofore described

trematodes, and I therefore suggest for it the name:

Allocreadium. lobatum, u.sp.

Habitat: Stomach of Semotilus bullaris.

The length of Allocreadium lobatum varies from 3 to 6.7 mm.

and the breadth from 0.72 to 1.5 mm. The general shape of the

A NEW TREMATODE 53

body is somewhat cylindrical, with the posterior part slightly flat-

tened dorso-ventrally and the extremity rounded. From the acet-

abulum the body diminishes slightly in diameter to the anterior end,

forming an ill-defined “neck” region. In a greatly contracted indi-

vidual the anterior end may be similar in shape to the posterior

end (Fig. 1).

The cuticula covering the body is a smooth homogeneous structure

without spines or scales and about 7 microns in thickness. In some

cross sections the cuticula showed fine lines dividing it into sections

of unequal size (Fig. 12). This could not be seen in all sections,

so is undoubtedly the result of fixation or of sectioning. Lying

deep in the parenchyma are the elongated epithelial cells which se-

crete the cuticula; v. Gronkowski (1902) investigated the struc-

ture of these cells beneath the cuticula in Distomum hepaticum,

Amphistomum conicum, Distomum variegatum, and Distomum

lanceolatum and concludes that they are genuine epithelial cells.

The two suckers are of about equal size, the variation in size in

the same individual is, at most, the same as for A. isoporum—

namely, as 13:14 (Looss, 1894). The oral sucker is circular in

outline and is situated sub-terminally; it varies in diameter from

0.455 to 0.473 mm. It has a well-developed musculature, the muscle

fibers radiating outward from the central part (Fig. 2). A few

large oval cells are present at irregular intervals between the muscle

fibers. On the inside it is lined by a thick cuticula, the continua-

tion of the body cuticula. Beneath the cuticula is a thinner mem-

brane, which is continuous around the whole sucker and forms an

attachment for the muscles. The acetabulum, in most individuals,

is slightly oval in outline, its longer axis lying at right angles to

the longer axis of the body. It varies in diameter from 0.455 to

0.509 mm. In structure it is similar to the oral sucker. Its posi-

tion in the body is at the posterior part of the first fourth of the

body.

The digestive system begins directly back of the oral sucker with

a well developed pharynx. There is no prepharynx present. The

pharynx is slightly oblong in shape, varying from 0.236 to 0.298

mm. in length and about 0.222 mm. in breadth. It has a well

developed musculature, which in general appearance is similar to

that of the suckers. The heavy cuticula of the oral sucker is con-

tinuous through the pharynx and esophagus, terminating at the

54 IVAN E. WALLIN

forking of the intestine. Following the pharynx is a long esopha-

gus, which, in the majority of individuals examined, forks at

about the anterior margin of the acetabulum. In one individual

the forking occurred at the posterior margin of the acetabulum.

Embedded in the parenchyma about the esophagus are a few deeply

staining epithelial cells, in appearance like those described by

Braun (1879-1893: 671), Looss (1902: 648), and others. These

cells secrete the thick cuticula that lines the esophagus. Cells of

similar structure are also found about the metraterm, ductus ejac-

ulatorius and beneath the surface of the body cuticula. Surround-

ing the cuticula of the esophagus is a circular layer of muscular

fibers and outside of this is a longitudinal layer (Fig. 5). In nor-

mal conditions the esophagus is about 0.1 mm. in diameter. The

two intestinal coeca lie laterally and near the dorsal surface. They

are about 0.185 mm. in diameter. In some individuals one of the

intestinal coeca has a smaller caliber than the other. There is no

indication that this condition is the result of contraction. The

intestinal coeca are lined by a single layer of cells which are more

or less irregular in outline. Large, somewhat oval, nuclei lie near

the bases of the cells (Fig. 8). Some of the cells in this layer

appear like gland cells, drops of secretion lying on the periphery;

clear oval spaces may also be seen at intervals, reminding one of

goblet cells in higher animals. Next to the epithelial layer is a

single layer of circular muscle fibers, then follow a few scattered

longitudinal fibers. The intestinal coeca end for the most part a

short distance from the end of the body, in some individuals about

midway between the posterior testis and the end of the body.

The excretory system is composed of a single median vesicle

reaching from the posterior end of the body to the posterior testis;

at this point two fine branches, of much smaller caliber, are given

off. The branches lie close together in the dorsal part of the body

and pursue a straight course to about the region of the ovary, where

they separate and run to the sides. From this point they pursue a

very tortuous course and it was impossible to definitely follow the

many coils that occupy a great part of the anterior region of the

body. The median vesicle is surrounded, especially in the posterior

part, by a great number of cells, which have the appearance of

gland cells, small ducts leading away from each cell to the vesicle.

The vesicle (Fig. 6) is lined with a heavy cuticula, which is more

A NEW TREMATODE 55

or less ragged in places and gives the appearance of cilia. At its

anterior end the vesicle has a smaller caliber and the cuticula is

not as heavy here as it is in the posterior part. The “gland” cells

lying about the vesicle are similar in appearance to the epithelial

cells mentioned in this paper and should be classed with them.

The testes in Allocreadium lobatum differ from all other species

in the genus in that they are distinctly lobed. They occupy a posi-

tion in the body that is characteristic for the genus, one iying back

of the other in the anterior part of the posterior half of the body.

There is quite a variation in the dimensions of the testes in differ-

ent individuals; the length varies from 0.374 to 0.546 mm. and

the breadth from 0.400 to 0.728 mm. The most common type of

form is broader than long, but this is reversed in some specimens.

From each testis a vas deferens is given off, which pursues an

almost straight course forward and each testis opens separately

into the seminal vesicle. The vasa deferentia are easily recognized

in section on account of their size and the darkly stained spermato-

zoa that are generally present in the lumen. The lumen of the

tube measures from 0.028 to 0.044 mm. in diameter.

The genital pore lies in the median line from 0.14 to 0.29 mm.

anterior to the acetabulum. Directly beneath the pore is the com-

mon genital atrium, which is lined by a thick cuticula that has a

rough appearance and is surrounded by a great number of mus-

cular fibers.

The cirrus pouch (Fig. 7) extends from the center of the acet-

abulum to the genital pore; it curves down over the acetabulum

so that in a sagittal section it would appear bent into a crescent

shape. It measures about 0.15 mm. in breadth at its posterior end

and diminishes in size anteriorly. The wall of the cirrus pouch is

made up of a single layer of muscle fibers enclosed by limiting

membranes. Darkly staining nuclei were observed, at intervals, in

the outer membrane. In the posterior part of the cirrus pouch is

a large seminal vesicle containing a vast number of spermatozoa.

The vesicle, which is bent into a half-circle, communicates anteri-

orly with a large pars prostatica, which is also curved. The pars

prostatica and seminal vesicle together form a figure S. The wall

of the pars prostatica is made up of circular and longitudinal mus-

cle fibers enclosed by thin membranes. The two muscular layers

and the arrangement of their fibers can best be seen in a tangential

56 IVAN E. WALLIN

section, such as shown in Figure 10 at x. In the lumen of the tube

can be seen spherical masses of unequal size, which at first sight

look like cells, but a closer study shows them to be structureless.

These spherical masses are drops of secretion that are the product

of the many prostate glands embedded in the parenchyma about the

pars prostatica. The ducts of the prostate glands can easily be

traced into the wall of the prostatica. The pars prostatica com-

municates anteriorly with a long ductus ejaculatorius which meas-

ures about 0.9 mm. in length and from 0.039 to 0.065 mm. in

thickness. Lying outside of the wall of the ductus and attached to

it are a great number of longitudinal muscle fibers (Fig. 9) which

undoubtedly serve to draw the ductus back into the cirrus pouch

after it has been evaginated. In a cross section of the ductus

ejaculatorius these muscle bands have the appearance of a thick

fringe about the tube. Beneath this longitudinal layer is a layer

of circular muscle fibers. The ductus ejaculatorius is lined by a

heavy cuticula, which, in the anterior half of the tube, is thrown

into folds and has a ragged appearance. Epithelial cells lie about

the whole length of the ductus ejaculatorius and undoubtedly

secrete the heavy cuticula lining the organ.

The ovary lies in the median line directly posterior to the acet-

abulum. It is spherical in shape and measures from 0.303 to 0.327

mm. in diameter. The ovary is completely filled with mature and

developing ova, the mature ova having a large circular nucleus in

which the chromatin is in a spireme and surrounded by cytoplasm,

which is variously shaped to accommodate the space in which it

is found. The mature ova measure about 0.015 mm. in diameter.

From the ventral side of the ovary the oviduct leads away: it meas-

ures from 0.015 to 0.020 mm. in width. There is no muscular tis-

sue in the walls of the ovary and oviduct; a thick membrane limits

the ovary and is continuous into the oviduct. Large nuclei are

found at intervals in the membrane. The oviduct has the same

structure and caliber throughout its length, there being no indica-

tion of an odcapt (Fig. 3) ; it measures about 0.106 mm. in length.

Between the ovary and the testes lies the receptaculum seminis,

a little to the right of the median line. It is a very large oblong

sack measuring about 0.418 mm. in length and 0.236 mm. in width.

The anterior end is slightly drawn out and communicates with

Laurer’s canal, which pursues a straight course to the right dorsal

A NEW TREMATODE 57

surface. A swelling occurs in the tube a short distance from the

receptacle; at this point a branch is given off which enters the ovi-

duct (Fig. 11).

The yolk glands in Allocreadium lobatum are profusely devel-

oped and are characteristic of the genus. The follicles are large

spherical masses filling the lateral areas of the body and are contin-

uous posterior to the testes. Anteriorly the yolk glands extend to

the level of the ovary. Two main ducts collect the yolk cells from

the two sides (Fig. 11) ; they come together in the median line and

form a yolk reservoir, which lies ventral and posterior to the ovary.

The yolk reservoir is drawn out anteriorly and communicates with

a tube whose caliber is large enough to contain only a single row of

yolk cells. This tube runs forward a short distance and then turns

back. At the point of the bending the oviduct enters it. The cal-

iber continues the same for a short distance, then widens out to

form the ootype. In this part of the tube—that is, from the point

that the oviduct enters—the lining is very thick and contains nu-

clei. Embedded in the parenchyma about the ootype are great num-

bers of gland cells, the shell glands, with their ducts, leading into

the wall of the ootype. The wall of the ootype is composed of quite

a thick membrane containing nuclei, outside of which is muscular

tissue.

The ootype is followed by a short tube of very much smaller cali-

ber, which is the beginning of the uterus. The uterus makes many

bends and fills up a great part of the space between the anterior

testis and the acetabulum. It was impossible to follow the uterus

in its complete length on account of the tearing produced when

the knife passed through the eggs. The uterus has a large caliber,

about two or three times the width of an egg. A little posterior

to the cirrus pouch the metraterm unites with the uterus; it pur-

sues quite a straight course to the genital atrium. The metraterm

is lined by a thick cuticula, which has a very ragged surface.

Beneath the cuticula is a layer of circular muscle fibers, then follow

the longitudinal fibers arranged like those about the ductus ejacu-

latorius. The lumen of the tube is much smaller than the width

of an egg, and when an egg is passing through a swelling is pro-

duced at the place where it may be located. Darkly staining epithe-

lial cells are also present in the parenchyma about the metraterm

(Fig. 7).

58 IVAN E. WALLIN

The eggs in Allocreadium lobatum are very numerous. They are

quite large, measuring from 0.067 to 0.085 mm. in length and

from 0.046 to 0.057 mm. in breadth. A small cap is found at

one pole of the egg. This could only be recognized when an oil

immersion objective was used (Fig. 4). The shell is colored a very

light yellow.

The body musculature is not greatly developed; a few scattered

fibers may be seen in a cross section. The fibers are slightly more

numerous in the anterior part of the body.

The parenchyma of the body does not differ from that usually

found in trematodes.

In reading the literature on trematodes I have met with a num-

ber of forms described, notably, by Stossich, which bear a striking

resemblance, both in illustration and description, to the forms in

the genus Allocreadium. Looss (1899: 577) points out the close

relationship of these forms to Creadiwm (Allocreadium) and sug-

gests that they might belong to closely related genera. While they

do not fully agree with the diagnosis of Allocreadwm, the varia-

tions, in my estimation, are not of sufficient importance to place

them in new genera. I have ventured, relying on the original

descriptions and illustrations of Stossich, to place the following

forms in the genus Allocreadium: “Distomum fasciatum Rud.” of

Stossich, D. mormyrt Stoss., and D. pallens Rud.

Allocreadium colligatum n. sp.

1885. “Distomum fasciatum Rud.,” Stossich p. 5.

1898. “Distomum fasciatum. Rud.,” Stossich p. 46.

This form described by Stossich is certainly not identical with

Helicometra fasciata (Rud.), as described by Odhner (1901).

Monticelli (1893: 192) gives an illustration of D. fasciatum Rud.

or Hel. fasciata (Rud.), and points out that it differs essentially

from Stossich’s D. fasciatum Rud. He also makes mention of eggs

with long unipolar filaments which are not mentioned by Stossich.

Braun (1879-1893, pl. xxii, Fig. 8, A. and B.) gives a reproduction

of Stossich’s drawing of D. fasciatwm Rud., and beside it he repro-

duces Willemoes-Suhm’s drawing of an egg from D. fasciatum

Rud. which has a long unipolar filament. This is undoubtedly a

confusion of the two distinct species, the egg belonging to Helico-

metra fasciata. Comparing the description and illustration of

Stossich’s D. fasciatum with Odhner’s description and illustration

A NEW TREMATODE 59

of Helicometra fasciata, the following differences are notable:

In Stossich’s D. fasciatum the ovary is globular, testes spherical,

and the vitellaria extend to the oral sucker; in Helicometra fasciata

the ovary is lobed, testes lobed, and the vitellaria extend to the an-

terior part of the acetabulum. These differences clearly show that

the two forms are not identical, but distinct species, and with the

difference in eggs they belong in separate genera, Stossich’s D. fas-

ciatum falling in the genus Allocreadium.

Stossich (1885) gives the following description for this form:

Collected from the intestine of Labrus miztus.

Body elongated, elliptical, somewhat depressed, unarmed, uncol-

ored; posterior extremity rounded. Oral sucker terminal, robust,

and united directly with a large pharynx. Acetabulum very large,

prominent, without pedicle, in form somewhat circular. Testes two,

large, sub-spherical, and situated one on the other in the middle

of the body. Cirrus very long, cylindrical, bends into a spiral and

terminates in an enlarged knob. Vitellaria very greatly developed ;

they occupy not only the whole posterior part of the body, but some

of the follicles are found almost at the oral sucker.

Length 2 mm.; breadth 0.3 to 0.5 mm.

Allocreadium mormyri (Stoss.).

1885. Distomum mormyri Stoss., Stossich p. 5.

1898. Distomum mormyri Stoss., Stossich p. 36.

This form is very much like Allocreadium obovatum Molin; it

differs from it, however, in the size of the body and in the shape

and size of the pharynx. The esophagus in mormyri, as well as in

obovatum, is very short and in this particular does not agree with

the diagnosis of the genus. A comparative study of the Allocrea-

diinae forms shows a gradation in the length of the esophagus from

a long to a very short one. It would be hard to draw a line between

forms which should be excluded from the genus and those that

would satisfy the diagnosis on this one character. In all other par-

ticulars the form agrees with the diagnosis of Allocreadiwm. Fol-

lowing is the description of D. mormyri given by Stossich:

Body unarmed, depressed, subovate, rounded at the two extremi-

ties. Oral sucker sub-terminal, circular, and is bordered by a mus-

cular girdle. Acetabulum much larger than oral sucker, elliptical-

transverse, somewhat prominent, aperture transverse. Genital aper-

ture is situated under the pharynx. Cirrus somewhat cylindrical,

60 IVAN E. WALLIN

unarmed, somewhat sharpened at the extremity; cirrus pouch bent

into a semi-circle extending beyond the posterior margin of the

acetabulum. Testes two, large, ovate, situated one behind the other

longitudinally in the body. Vitellaria greatly developed, follicles

small spheres; they occupy the whole posterior part of the body and

extend to the pharynx. Uterus short; eggs relatively large, ellip-

tical, colored intensely yellowish-brown.

Length 2 to 2.5 mm.; breadth 0.75 to 1 mm.

Allocreadium pallens (Rud.).

1819. Distomum pallens Rud., Rudolphi p. 408.

1887. Distomum pallens Rud., Stossich p. 4.

1898. Distomum pallens Rud., Stossich p. 47.

1898. Distomum pallens Rud., Linton p. 526, pl. xlvii, figures

8 and 9.

I have been unable to find a good illustration of this form, so

have had to rely for generic determination on the accuracy of the

description given by Stossich. Linton (1898) figures a D. pallens

Rud., but the drawing is so poorly executed that it is not service-

able. The form fully agrees with the diagnosis of Allocreadium

with the exception of the esophagus, which is short. The following

description is compiled from Stossick (1887 and 1898) :

From the intestine of Chrysophrys surata.

Body elongated, unarmed, posterior and rounded. Acetabulum

very prominent, situated at the base of the neck, twice as large as

the oral sucker; aperture elliptical-transverse.

Oral sucker sub-terminal, globular, and provided with longi-

tudinal aperture; from the center a short canal (prepharynx) leads

back and communicates with a large bulbus esophagus (pharynx),

which has a quadrangular shape; this departs into a short esopha-

gus, which divides into two intestinal coeca. The intestinal coeca

extend to the extreme posterior end of the body.

Testes two, posterior, large, elliptical; cirrus pouch in the vicin-

ity of the acetabulum, contains the seminal vesicle. Ovary globu-

lar, next to the testes. The yolk glands are numerous, composed of

small globular follicles, extend from the extreme caudal end to the

ovary. Seminal receptacle large and situated to the right side of

the ovary. Eggs elliptical, limited in number. Genital aperture sit-

uated below the forking of the esophagus.

Length 5 to 6.5 mm.; breadth 0.5 to 1 mm.

A NEW TREMATODE 61

The following key comprises all the species described in the sub-

family Allocreadiinae with the exception of Helicometra sinuata

(Rud.) and Allocreadium asymphyloporum Stoss. H. sinuata has

not been definitely established as a distinct species, so it has been

emitted on this account; the description of A. asymphyloporum is

so incomplete that it was not possible to give it a definite place in

the key.

Worms from small to medium size; cuticula without spines; long intes-

tinal coeca; genital aperture near forking of esophagus; seminal receptacle

large; yolk glands profusely developed, coming together and are continuous

in the posterior body; uterus short between anterior testis and acetabulum.

In fish. SUB-FAM. ALLOCREADUNAE Looss.

A.—Eggs with long, unipolar filaments; esophagus of moderate length; geni-

tal aperture in median line; ovary lobed. In sea-fish.

GeNus Helicometra Odhner.

I. Testes lobed.

1. Acetabulum little larger than oral sucker? pharynx globular or

elongated, prepharynx short.

a. Ovary 6-8 lobed; vitellaria extend to anterior margin of

acetabulum; pharynx globular; length 2 to 3 mm.

Helicometra fasciata (Rud.)

b. Ovary 3 lobed; vitellaria extend to genital aperature; cirrus

pouch extends to anterior margin of acetabulum; pharynx

globular; length 2 to 3 mm. Helicometra flava (Stoss.)

2. Acetabulum almost double the size of the oral sucker; pharynx

squarish in outline; prepharynx short or long; cirrus pouch

extends to anterior margin of acetabulum; ovary 4-5 lobed.

a. Vitellaria extend to oral sucker; prepharynx short; pharynx

sub-quadrate; length 4 to 5 mm.

Helicometra mutabilis (Stoss.)

b. Vitellaria comparatively sparse; extend to the genital aper-

ture; prepharynx long; pharynx broader than long; length

1 to 2.5 mm. Helicometra gobii (Stoss.)

II. Testes rounded; acetabulum larger than oral sucker; pharynx

globular.

1. Vitellaria extend to the bifurcation of the esophagus; prepharynx

short; intestinal coeca extend to extreme posterior end;

ovary 3 lobed; length, 2 mm.; breadth, 0.5 to 0.75 mm.

Helicometra pulchella (Rud.)

2. Vitellaria extend to oral sucker; prepharynx absent; intestinal

coeca extend to short distance from posterior end; ovary

multilobed; length, 2.3 mm.; breadth, 0.9 to 1 mm.

Helicometra sinuata (Rud.)

B.—Eggs without filament.

I. Suckers well developed; cuticula smooth; esophagus long; genital

aperture near forking of intestine. In (fresh-water?) fish.

GENUS Allocreadium Looss.

IVAN E. WALLIN

Ovary lobed; prepharynx present; pharynx globular; forking of

esophagus about midway between suckers; eggs numerous.

a. Vitellaria extend to acetabulum; cirrus pouch slender, extends

to a little beyond acetabulum; length, 1.5 to 3.4 mm.

Allocreadium atomon (Rud.)

b. Vitellaria extend to pharynx; cirrus pouch club-shaped, ex-

tends to posterior border of acetabulum; length, 2 to 6 mm.

Allocreadium labracis (Duj.)

. Ovary spherical.

a. Testes lobed; vitellaria do not extend anterior to acetabulum.

aa. Suckers of equal size; no prepharynx; cirrus pouch ex-

tends to ovary; seminal receptacle between ovary and

testes; eggs very numerous, 0.067 to 0.085 by 0.046 to

0.057 mm.; length, 4 to 6.7 mm.; breadth, 1 to 1.5 mm.

Allocreadium lobatwm Wallin.

b. Testes rounded; vitellaria extend to acetabulum or beyond.

aa. Prepharynx present.

aaa. Vitellaria extend to ovary; acetabulum twice as large

as oral sucker, and pharynx quadrangular; length, 5

to 6.5 mm.; breadth, 0.5 to 1 mm.

Allocreadium pallens ( Rud.)

bbb. Vitellaria extend to acetabulum.

x. Acetabulum equal in size to oral sucker; length,

5 mm.

Allocreadium isoporum Looss.

y. Acetabulum 114 times as large as oral sucker.

Allocreadium transversale (Rud.)

ece. Vitellaria extend to pharynx.

x. Cirrus pouch long and slender, extends to ovary;

ovary to right of median line; genital aperature

in median line; length, 2 to 2.5 mm.; breadth, 0.5

to 6 mm. Allocreadium genu (Rud.)

y. Cirrus pouch quite large, extends a little back of

acetabulum; ovary to left of median line; genital

aperture to right of median line; length, 1.5 mm.;

breadth, 0.3 mm.

Allocreadium angusticolle {(Hausm.)

z. Cirrus pouch curves into a crescent shape, extend-

ing a little back of acetabulum; ovary to right of

median line; genital aperture to left of median

line; esophagus very short.

xx. Pharynx almost equal in size to oral sucker,

globular; ovary sub-spherical; length, 2.5 to

4.5 mm.; breadth, 0.8 to 1 mm.

Allocreadium obovatum (Molin.)

yy. Pharynx elongated; ovary spherical; length,

2 to 2.5 mm.; breadth, 0.75 to 1 mm.

Allocreadium mormyri (Stoss.)

bb. Prepharynx absent.

aaa. Pharynx long and cylindrical; esophagus short (0.06

mm.), genital aperture in median line; cirrus pouch

to 1 mm. Allocreadium commune (Olss.)

extends to ovary; length, 1.4 to 3 mm.; breadth, 0.4

A NEW TREMATODE 63

bbb. Pharynx globular; esophagus short; cirrus pouch ex-

tending to middle of ovary; length, 2 mm.; breadth,

0.3 to 0.6 mm. Allocreadium fasciatum (Stoss.)

ece. Pharynx globular; esophagus long (0.1 mm.) ; genital

aperture left of posterior edge of pharynx; cirrus

pouch extends to anterior border of acetabulum;

length, 1.5 to 2.25 mm.; breadth, 0.35 to 0.6 mm.

Allocreadium tumidulum (Rud.)

II. Suckers small, of equal size; cuticula with scales easily coming off;

esophagus short; genital aperture anterior and left of acet-

abulum. Jn sea-fish. GENus Lepocreadium Stossich.

1. Long prepharynx; testes at extreme posterior end of body; semi-

nal receptacle spherical; penis unarmed; vitellaria extending to

pharynx; length, 2 mm.; breadth, 1.75 mm.

Lepocreadium pegorchis Stoss.

. Short prepharynx; testes at anterior second half; seminal re-

ceptacle bottle-shaped; penis armed; vitellaria extending to acet-

abulum; length, 1 to 2.5 mm.; breadth, 0.3 to 0.6 mm.

Lepocreadium album Stoss.

This study was carried out in the Zoological Laboratory of the

University of Nebraska during the academic year 1907-1908. I

wish to take this opportunity to express my sincere thanks to Prof.

Henry B. Ward for his guidance in the work, for the preserved

materials, and for the use of his extensive private library.

BIBLIOGRAPHY

Brawn, M.

1879-1893. Vermes-Trematodes. Bronn’s Klassen und Ordungen des

Thier-reichs, Leipzig, iv, 1.

Linton, E.

1898. Notes on trematode parasites of fish. Proceed. U. S. Nat. Mus.,

xx, 507-548.

Looss, A.

1894. Die Distomen unserer Fische und Frésche. Biblioth. Zool., xvi,

1-293.

1899. Weitere Beitrage zur Kenntniss der Trematoden-fauna Aegypt-

ens. Zool. Jahrb., Syst., xii, 521-784.

1900. Nachtriigliche Bemerkungen zu den Namen der von mir

vorgeschlagenen Distomidengattungen. Zool. Anz., xxiii, 601-608.

1902. Ueber neue und bekennte Trematoden aus Seeschildkroten.

Zool. Jahrb., Syst., xvi, 411-894.

MacCattum, W. G.

1895. On the Anatomy of two distome parasites of fresh-water fish.

Veterinary Mag., ii, 1-12.

MOonrTICELL, FR.

1893. Studii sui Trematodi endoparassiti. Zool. Jahrb., Supp. 3.

ODHNER, TH.

1901. Revision einiger Arten der Distomen-gattung Allocreadium Lss.

Zool. Jahrb., Syst., xiv, 483-520.

1902. Mitteilungen zur Kenntniss der Distomen. Centralbl. f. Bakt.

u. Parasit., xxxi, 152-162.

64 IVAN E. WALLIN

RUDOLPHI, C. A.

1819. Entozoorum synopsis, Berol., 811 pp.

STAFFORD, J.

1904. Trematodes from Canadian Fish. Zool. Anz., xxvii, 481-495.

Stossicu, M.

1885. Brani di elmintologica tergestina. Boll. Soc. Adriat. se. nat.

Trieste, Series 2, ix, 1-9.

1887. Brani di elmintologica tergestina. Boll. Soc. Adriat. se. nat.

Trieste, Series 4, ix, 1-7.

1898. Saggio di una fauna elmintologica di Trieste e provincie con-

termini. Progr. civ. scuola reale sup.; 162 pp.

1900. Osservationi elmintologiche. Boll. soc. Adriat. sc. nat. Trieste,

xx, 89-103.

1902. Sopra una nuova specie delle Allocreadiinae. Arch. Parasit.,

v, 578-582.

1903. Una nuova specie di Helicometra Odhner. Arch. Parasit., vii,

373-376.

1904. Note distomologiche. Boll. soc. Adriat. se. nat. Trieste, xxi,

193-201.

1904a. Alcuni Distomi della collezione elmintologica del Museo Zoolog-

ico di Napoli. Annario Mus. Zool. Univ. Napoli, n. s., i, 1-14.

GRONOWSEI, C. VON.

1902. Zum feineren Bau der Trematoden. Polnischer Archiv. f. biol.

und med. wissensch., i, 1-29.

A NEW TREMATODE 65

EXPLANATION OF FIGURES

All figures were drawn with the camera lucida. The following abbrevia-

tions hold for all figures:

A. Acetabulum. OD. Oviduct.

C. Cuticula. Ov. Ovary.

CM. Cireular Muscle. PG. Prostate Glands.

CP. Cirrus Pouch. PP. Pars Prostatica.

DE. Ductus Ejaculatorius. 8. Secretions.

EC. Epithelial Cells. SG. Shell Gland.

ED. Excretory duct. SR. Seminal Receptacle.

LM. Longitudinal Muscles. Re Testis.

LC. Laurer’s Canal. U. Uterus.

M. Metraterm. V. Vitellaria.

Mu. Muscle fibres. VD. Vas Deferens.

Oo. Oédtype. YR. Yolk Reservoir.

RupotPH, C. A.

PLATE I.

Fig. 1—Allocreadium lobatum ; toto, the uterus is not represented in the

figure. X 25.

Fig. 2.—Tangential section of anterior end. X 250.

Fig. 3.—Section through ovary and oviduct. X 400.

Fig. 4.—Egg. X 400.

Fig. 5.—Cross-section of esophagus. X 400.

Fig. 6.—Portion of a cross-section of excretory vesicle. XX 400.

66 IVAN E. WALLIN

PLATE II.

Fig. 7.—Optical section of cirrus pouch and metraterm. X 80.

Fig. 8.—Cross-section of intestinal coecum. X 400.

Fig. 9.—Section of ductus ejaculatorius. X 250.

Fig. 10.—Section of pars prostatica. X 400.

Fig. 11.—A reconstruction of the egg-forming organs. X 80.

Fig. 12.—Part of a cross-section showing the cuticula. X 400.

A PLEA FOR SYMPOSIUM WORK.

PONDLIFE AND NEW METHODS OF NARCOTIZING POLYZOA ROTIFERA.

VIDA A. LATHAM, D.D.S., M.D.

CHICAGO

In view of a tendency in this Society to have its volumes fill up a

gap in scientific study, it was thought the subject of “Plankton”

and its results might furnish a good field for research, at the same

time providing a special corner which can fill out a want both from

microscopic, biologic and economic reasons.

In an age of books it is strange that the rambler beside ponds and

ditches should have been left without a pocket companion until the

little manual by the late M. C. Cooke appeared in the series entitled

“Natural History Rambles” called “Ponds and Ditches,” and the

one by C. O. Groom Napier on “Lakes and Rivers” popularized the

subject so much loved by the late H. Stack in his “Marvels of

Pondlife.” As a suggestion I would ask the consideration of form-

ing a group or groups along some line of research and endeavor to

work in one or other line of study, viz.: “Pondlife,” to include (a)

fauna; (b) methods of collecting; (c) preserving and mounting;

(d) staining or the value and use of reagents; (e) results of

studies in varieties and their morphology, etc.

This subject can be used to exchange material with those engaged

in other lines of work; so that members can have material named

by those who know what the organisms are and to use them to iden-

tify other forms and to familiarize themselves with an unknown

subject, for the general complaint is “I do not know protozoa when

I see them and so pass them by.” Unfortunately, our workers are

scattered and many members wish for help in their rambles, and

we cannot do better than make a record in our Transactions by the

successful “fielders” to notify of good places to study and gather

material in the vicinity of our cities. When members go to Atlantic

City, New York, Boston, etc., who is there who would not be glad

to hear of a good nearby collecting spot that can be studied with

ordinary apparatus, and so obtain material, fresh and preserved, of

marine forms unknown to those of us living in the middle west.

Some of us who visit California, Catalina and Puget Sound might

be glad to know of localities, material and those willing to assist the

visitors when there, and exchanges could be had and study work

provided for winter months and rainy days.

68 Vv. A. LATHAM

It is a great disappointment to find after one has left Florida,

New Orleans, or some other point, of the fine hunting naturalists’

paradise we have passed by because we did not know of it. Some

may say if we publish a locality where some fine colonies of frederi-

cilli are to be found, we shall have the pond spoiled. It is hoped

no member of our society is so deficient in humane collecting and

a true scientific spirit as to wantonly destroy or take more than is

reasonable for study and an exchange.

The difficulty still remains unsolved as to the best manner of

preserving specimens of desmids, and such like small alge, for

future reference. The same is true of rotifers, though Mr. Rousse-

let’s formula with cocain or eucain has been a valuable help. Poly-

zoa and many of the other varieties all need special methods to

preserve them in a natural way. Some forms kill and expand

easily under a 14 per cent. solution, where again nothing but a 10

per cent. will affect another species. A society like this can do a

valuable amount of work in just this experimental study. Some

of us who are expert technicians can mount in deep cells with some

ringing cement known to be permanent if used for formal, cocain

or chloretone solutions. Let them give us their ideas, for exchange

is no robbery and only fair, unless a patent is held for his process.

To those interested in this line of work it will be of interest to

take up the group of chemicals known as anodynes, among which

we have cocain, eucain, beta-cain, stovain, kélene, chloretone,

hydroxyline, phenol, camphor, thymol, etc. Various combinations

of these give excellent results for special things. There is no excuse

for lack of material, for many forms can be obtained in January,

even under the ice and all the year round. Preparations to show the

urticating bodies or thread cells have so far been unsuccessful.

Who has ever made serial sections of some of the polyzoa and

studied their anatomic details? Some papers on the preparation of

hydroids, mostly marine forms, by Mr. Harris of London, and pub-

lished in the Annals of Microscopy, will be worthy of noting by any

one interested in methods of killing these beautiful animals.

A method recently brought to my notice, through the kindness of

Mr. H. E. Hurrell of England and communicated to him by Mr.

Bradley of Adelaide for narcotizing, has certainly yielded most

excellent results. The animals, say plumatella or other polyzoa, if

in 2-6 ounces of pond water, are carefully washed in several changes

of fresh spring or tap water by very gentle shaking in a suitable

vessel. Care must be taken not to detach them from the plant if it

can possibly be avoided. At any rate, care has to be exercised that

they are not broken up by too vigorous action (reversing the tubes

SYMPOSIUM WORK 69

or bottle should be sufficient if done several times). Now allow the

polyzoa to stand in pure tap water or twice filtered river water

until they have recovered from shock and extended freely. I use

wide tubes of about 4 or 5 inches in length with not too many ani-

mals in each tube so as to give one a perfect view of the progress

toward narcotization. Next have a special lot of stovain or eucain

at hand and take up one drop only of the drug (15 grains dissolved

in 100 c.c. of water), but before applying it fill up the pipette with

fresh water so that it may dilute the dose. Then, with all care not

to disturb the protozoons, squirt the mixture down the side of the

tube in which they are, continuing the pipetting some 6 or 8 times

so as to ensure that the agent thoroughly combines with the water.

Repeat the one-drop dose at intervals of 10 minutes for 3 or 4

times, then go on with 2 drops for every 5 minutes, for a quarter of

an hour; then proceed with about 4 drops if by this time the ani-

mals be not too sleepy. Great care and absolute attention is

required at a certain psychologic moment, viz.: when there is the

slightest tendency for the tentacles to curl over outward. At this

_ moment one of the animals should be touched carefully with the

pipette, and if it remains without flinching the process is nearly

complete. It should then be given a dose of about 6 drops and

allowed to stand for a short time, and if you have a sufficient

material a colony should be placed by itself in a small watchglass

with a sufficient quantity of the narcotized water; then a pipette

full of formalin (10 per cent. of the 40 per cent. commercial forma-

lin) discharged over it, and if sufficiently narcotized this will

remain extended and be killed and fixed at the same time. (N. B.

The pipettes must on no account be mixed or used for any but the

one reagent and it is well to color or mark them so no confusion

can be made.) ‘The stovain is used in the same strength that the

eucain is for the narcotizing of the polyzoa, rotifera and ento-

mostraca (Hurrell).

Chloretone is a most excellent solution, used in 1-5-10 per cent.

strength solutions and is worthy of a trial.

If desired for class study, the animals can be rendered insensible

and thoroughly examined, and when done with add more fresh water

and they recover and seem no worse for their treatment. As noted

by Prof. A. H. Cole of Chicago, it makes an excellent method for

projecting animals on the screen when all can study their anatomy

and many peculiarities. It affords both study and a pleasant sub-

ject for a lecture.

Those of the members who frequent the inland lakes of Wiscon-

sin, Michigan, Indiana, Montana, Minnesota, New York and

70 Vv. A. LATHAM

Florida remember to collect a few vials for others interested and

exchange for insects, diatoms, etc., and receive the gratitude of your

less fortunate members. Remember the rotifera are prolific in bog-

mosses or turf-mosses, as the Germans call them. The biologist

confesses to the presence of infusoria within the cells of the leaves,

but who can enumerate the creatures which sport outside, the water-

fleas, wheel bearers and small snails; the confervoid alge?

The sphagna suck up the atmospheric moisture and convey it to

the earth. So they also contribute to it by pumping up to the sur-

face of the tuft formed by them the standing water which was their

cradle, diminish it by promoting evaporation, and later become

material for fuel.

As a favorite locality for stephanoceros note the foliage of the

water milfoil (myriophyllum) and on the slender roots of the wil-

lows which run into the water. Floscularia ornata or cornuta is met

in the leaves of submerged plants as the water crowfoot. Take

some of the slender-leaved water plants and examine a few branches

at a time in a vial with a pocket lens. A small square vial is best

made with two pieces of glass separated by three small pieces care-

fully joined together with marine glue or Ward’s brown cement or

the gelatin bichromate insoluble cement. It should be wide enough

to easily admit the stems of water plants, so as not to squeeze them

in putting them in. Two or three of these will be a great help, as

one can stand quietly and allow the animal to recover from shock.

The first ghmpse reveals an egg-shaped object of a brownish tint,

stretching itself upon a stalk, and many show signs of hairs or

cilia at its head. Now carefully manage the light obliquely, and

the dirty brown hue disappears and is replaced by brilliant colors ;

the cilia become very long and resemble spun glass.

In regard to carrying collections of animals in bottles with water,

much difference of opinion exists as to whether the jars should have

an air vacuum left or not. Davis’ Practical Microscopy, p. 134,

says: “Do not have any portion of the bottle or tube filled with air

if they are to be exposed to shaking or concussion.” Dr. A. C.

Stokes fills his bottles. I find a good way is to have a quill passed

through each cork and cut off close at the outside, and this will

prevent the water coming out and admit air at the same time.

If this method is of any use we should be glad to hear from

others who might give their experiences in the forms of sympo-

siums and a good discussion.

REPORT OF THE COMMITTEE ON THE ERNST ABBE

DENKMAL.

At the Sandusky meeting of the American Microscopical Society,

July 6, 1905, the President presented an appeal for cooperation of

the American Microscopical Society in the erection of a monument

at Jena, Germany, to the memory of the late Professor Ernst Abbé.

The peculiar propriety of our participation was urged by all who

participated in the discussion and the undersigned were appointed

a special committee to select funds and forward them in the name

of the Society (see Transactions, xxvil, 162). Some money was

also subscribed by those present and more was secured during the

fall by private solicitation.

During the following winter the committee sent a circular appeal

to every member of the society, and also prosecuted its efforts to

secure personal subscriptions. The announcement was made that

exact amounts subscribed would not be made public, but that all

money received would be forwarded in a lump sum under the name

of the donors as well as of the society.

Believing that its work is now finished, the committee has for-

warded recently the sum collected with the following communica-

tion to Dr. Gustav Fischer, Jena, treasurer of the fund, with the

following memorandum :

The American Microscopical Society, recognizing the great services of

the late Prof. Dr. Ernst Abbé to the cause of scientific investigation, espe-

cially as connected with the use of the microscope, desires to contribute to

the Ernest Abbé Denkmal and encloses herewith draft for 450 marks* for

this purpose.

On the part of the society by

Allen, W. E. Haag, D. E. Seawell, B. L.

Barelay, L. P. Hollis, F. S. Shultz, C. S.

Bessey, C. E. Holmes, A. M. Siemon, R.

Birge, E. A. Krauss, W. C. Slocum, Chas. EF.

Bleile, A. M. Kuehne, F. W. Smith, J. C.

Brown, Robert McCraven, B. N. Ward, Henry B.

Buffet, E. P. Moody, R. O. Ward, R. Halsted

Flint, Jas. M. Pennock, E. Whelpley, H. M.

Gage, S. H. Pflaum, M. Wolcott, R. H.

Gifford, H. Ransom, B. H.

In response the committee received the following letter from the

treasurer of the fund:

* The sum originally sent was 425 marks, but 25 marks was received

and forwarded later.

12 ABBE DENKMAL

JENA, 22 Juli, 1907.

Herr Henry B. Ward, President of American Microscopical Society,

Lincoln:

Hochgeehiter Herr! Ich erheilt heute Ihren Check in Betrage von M.

425 als Beitrag der Amer. Microscopical Society fiir das Abbé Denkmal.

Ich spreche Ihnen meinen verbindlichsten Dank fiir die liebenswiirduge

Sendung aus und bitte Sie auch den Herren, welche so freundlich waren

sich mit Beitriigen zu beteiligen, im Namen des Comité zur Errichtung

eines Abbé-Denkmals den wiirmsten Dank zu iibermitteln.

Mit grésster Hochachtung.

[Signed] Gustav FISHER,

Kassenfiihrer des Comités zur Errichtung eines Ernst Abbé Denkmals.

In all the committee has received $107.50 and has paid out for

drafts of 425 marks and 25 marks the sums of $101.40 and $6.06

and for postage 14 cents.

The committee begs that, having submitted herewith its final

report, it may be honorably discharged.

(Signed) Henry B. WARD,

Macnvus PFLAUM,

Rosert H. WoLcort.

Su Memoriam

Cc. C. MELLOR.

C. C. Mellor, Pittsburg, Pa., treasurer of this society from 1889

to 1894, died, after a lingering illness, on April 2, 1909, at the age

of 73 years. He departed too soon for those who knew him; and

few were better known and more highly and deservedly esteemed

than he was in the city of his birth and manhood.

He was an exceptional man. Absorbed in practical affairs, own-

ing and managing a large business, yet he found time for and

delighted in whatever tended to culture, refinement and learning.

It is not too much to say that to a great extent Pittsburg owes to

him her present position as a center of music, art and education.

To mention the organizations for these purposes which he founded

and to those to which he gave his active, hearty and unstinted

financial support would occupy too much space. Whatever and

whoever interested him made of him a warm friend ; and his friend-

ship and assistance meant success, for he was born to succeed. In

his varied relations he met many minds, yet such was his well-

poised tact and judgment that his presence alone caused harmony

and silenced controversy. He was eminently a man of peace;

because liberal and generous in his views and careful in his con-

clusions, he respected the opinions, prejudices, and rights of others.

He was one of the truest, most simple-hearted and whole-souled

of men. ‘To meet him was to know him, to know him compelled

trust and admiration, and his friendship was a delight and inspira-

tion.

The mourning for a departed friend is in this case much relieved

from its sadness by a view of the beauty of his life. He worked in

broad daylight, and the sun sent his approving rays upon his many

beneficent activities. He offended none and enjoyed rather to praise

than to blame. Animated by high, pure aims, he succeeded in

their realization by rare sagacity and discretion. His private,

social, business and public relations he conceived as duties in which

no one did excel him in joyous, modest, unassuming, and conscien-

tious performance.

Thus he lived, and was a practical sermon to his friends and the

community at large. Macnus PFriaum.

(ys

pany

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PLAN ally

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Coalnhinol

OFFICERS.

iBrossdent: TAERBERT OSBRORNG cies ae sia diets oe wis vieieiw Wiel oto! oye Columbus, Ohio

Vice President: A. E; Hertztre, M.D....... 02.2665. 605s os Kansas, City, Mo.

BCRCLOTY 2) Leas MALLU A eee chic cele cpecisln b:s-o)3 6 sein warwipe orate aha Decatur, III.

MPC OSUT ey asl wle, SVAN KUN SONsIa sitstelae cine e iss aaa sistelousiole sree Charleston, III.

Cusiouan: NIRGNUS PREAUM 0 o.0 cos ooo nse wales ain crete olayaisiace lel Meadville, Pa.

ELECTIVE MEMBERS OF THE EXECUTIVE COMMITTEE

MERITS EDATNUAMEAIN 153s dh cae ace Pe ieee ave ISTE ASTOR oT RIOR leah ars as Troy, N: Yi

HERE Se RACY, Sco hk cs pce tu roped ots eee Tae TER aoa Oe Loses om ace he ete Tate) caer Oakland, Cal.

EX-OFFICIO MEMBERS OF THE EXECUTIVE COMMITTEE

Past Presidents still retaining membership in the Society

R. H. Warp, M.D., F.R.M.S., of Troy, N. Y.,

at Indianapolis, Ind., 1878, and at Buffalo, N. Y., 1870.

J. D. Hyatt, of New Rochelle, N. Y.,

at Columbus, Ohio, 1881.

Apert McCatra, Ph.D., of Chicago, III.

at Chicago, Ill., 1883.

T. J. Burrmt, Ph.D., of Urbana, IIl.,

at Chautauqua, N. Y., 1886, and at Buffalo, N. Y., 1904.

Gro. E. Fett, M.D., F.R.M.S., of Buffalo, N. Y.,

at Detroit, Mich., 1890.

MarsHALL D. Ewett, M.D., of Chicago, IIl.,

at Rochester, N. Y., 1892, and at Boston, Mass., 1907.

Simon Henry Gace, B.S., of Ithaca, N. Y.,

at Ithaca, N. Y., 1895 and 1906.

A. CiirForp Mercer, M.D., F.R.M.S., of Syracuse, N. Y.,

at Pittsburg, Pa., 1896.

A. M. Buette, M.D., of Columbus, Ohio,

at New York City, 1900.

C. H. E1iceEnmann, Ph.D., of Bloomington, Ind.,

at Denver, Colo., Igor.

Cuares E. Bessey, LL.D., of Lincoln, Neb.,

at Pittsburg, Pa., 1902.

E. A. Birce, LL.D., of Madison, Wis.,

at Winona Lake, Ind., 1903.

Henry B. Warp, A.M., Ph.D., of Lincoln, Neb.,

at Sandusky, Ohio, 1905.

The Society does not hold itself responsible for the opinions expressed

by members in its published Transactions unless endorsed by a special vote.

TABLE OF CONTENTS

FOR VOLUME XXIX, NUMBER 2.

The Future Activity of the American Microscopical Society, by T. W.

Galloway. 2525 seo cae ae od ot bea Ce bee koe vars Cae er 79

The Phyla, Classes, and Orders of Plants, by Charles E. Bessey..... Serene.

New Studies of the Arrhenuri, by Ruth Marshall, with plates I to III.... 97

The Lakes of the Glacier National Park: 1. Avalanche Lake, by Morton

J-Elrod<with plates:lV- to. Villu ss soa Soncccw sence os Ree te eee III

Recent Progress in Parasitology, by Henry B. Ward.................... II9

Recent Progress in the Pedagogy of Biology, by John G. Coulter........ 159

The Spencer-Tolles Fund; by Magnus Pilaum.<. 5/0. 00. ewes ec eee 167

Notes, Reviews, etc. A Method of Making Absolute Alcohol; A Con-

venient Reagent Case; Cultures of Saprolegnia; Cultivation of Fresh-

water Algae; Distribution of Rotifers; Vitality in Paramecium;

Technic of Tubercle Bacilli; Bacterial Infection by Endoparasites;

Longevity in Trichina; Symbiosis and Parasitism; The Use of the

Microscope:in Elementary Biology... j0...s....sscstestecse nee 17I

Necrology.

William Henry Dallinger, D.Sc., D.C.L., with Portrait, by A. E. Shipley 183

William Christopher Krauss, M.D., with Portrait, by Henry R. Howland 187

Richard S.-Nuanns, M.D,, by H. W.. Witcovero:« os...35.0<5 . cae so eee 189

Goenstitution ‘and. By-Lawsy si 2 s.cud< os coins see 4 opie ts 0 cee ee I9I

MEIStHOL SMIEMDETS Hes, oe oe eins oe ace OOo e se Clete in ane ee 195

Peists GE. SSUNSCEMDERS «3s Vac ove sc.cc 2 c0s Oeh «Fe mwegeeae 8c i 201

1 Bote (o> cat ed tk ie ie ge cara ae iy Oe De REE Oe Te Pe EM ene LIS eee ee et 203

Wxivertisements: ..2cs 2 be cen vaseissta tes Sande check eee meee eure ae I-VI

OFFICIAL NOTICE OF ANNUAL MEETING

MINNEAPOLIS, MINN., DEC. 27-28, 1910

IN CONNECTION WITH THE A. A. A. S.

The Secretary has already received the titles of enough papers

to make an interesting meeting.

Business of unusual importance to the members will be trans-

acted. There should be a good attendance.

Please send to the secretary at once the title of your paper, and

the names of any whom you wish to propose for membership.

PLACE OF MEETING: UNIVERSITY OF MINN.}; PILLSBURY HALL, ROOM 7.

TRANSACTIONS

American Microscopical Society

(Published in Quarterly Installments)

Vol. XXIX OCTOBER 1910 No. 2

Tne PULURE ACTIVELY OF Tits SOCIETY

By THE SECRETARY

As was announced in the last number of the Transactions, the

publications of this Society are to be issued quarterly, instead of an-

nually as heretofore. Unfortunately the numbers have not been

published regularly during the current year. This number, of extra

size, concludes Vol. XXIX. Beginning with January, 1911, the

issues will occur regularly four times per year.

This change at this time from the annual volume to a quarterly

journal marks a new emphasis in our activities. Our prime work

now becomes openly and frankly, as it should be, a work of publica-

tion. The quarterly form means more timely publication of scien-

tific materials, and allows an adaptedness and flexibility denied to

an annual volume. Our organization, our membership, and our

meetings, from this time, come to be confessedly for the end of

making this publication effective for our own use and for that of

all lovers of micro-biology. To this the Secretary pledges his best

efforts.

The fact that our society is named the “AmMeERIcAN MiIcro-

SCOPICAL Society’ need not give any uneasiness to anyone. It has

a splendid record both of research and publication; its growing

emphasis on the results of the microscope, rather than on the mere

instrument itself, is both natural and creditable and exactly dupli-

cates the history of similar societies in Europe. It can find its final

function in the upbuilding of American science just as well with

this name as with another, even though it may not be fully descrip-

tive of its changing field.

80 T. W. GALLOWAY

As the Secretary sees the situation, we, as a society, are just

now in need of some degree of conscious readjustment to the needs

of our times. There appear to him two very distinct ways along

which satisfactory adjustment and usefulness may be had :—(1) to

enter upon some specialized field of microscopic research, now un-

provided with adequate means of publication in this country, and

to make this journal the best exponent of this especial interest; or

(2) to enter confidently and with open mind into the fuller service

of those lovers of micro-biology who, from necessity or choice, wish

and need to have a general and synthetic view of the field, rather

than a highly specialized and technical journal.

The first alternative, while presenting some remarkably at-

tractive features, would involve a complete reversal of our tradi-

tional policy; and while this in itself is not prohibitive, it does de-

mand that we shall see whether some other sound solution may not

be in better harmony with the whole course of our evolution. Such

a policy furthermore would produce an almost complete change in

our membership. Probably less than 25 per cent of our members

would be in need of so narrowly technical a journal as this policy

would necessitate.

If, on the contrary, the present development of biological

science in this country demands a journal which seeks to deal ac-

curately and yet broadly with general micro-biology, this journal is

in an excellent position to fill this need.

It is somewhat true that the specialized research student can

not always resist the temptation to be a snob, and is liable to be in-

tolerant of any publication which undertakes to deal with so large

a field of interest as ours has been. And undoubtedly such treat-

ment does have its pitfalls. The writer does not believe, however,

that this lack of sympathy for the more general biology is sound,

even for the investigator; and he is confident that the very best

and sanest specialists in our society are not out of sympathy with

this point of view, and will not be lost to the society. At any rate

there are a great many students, and their number is increasing,

who do feel the need of just this synthetic view of biology; and

their needs are quite worthy of attention on the part of scientific

men.

Indeed these more general students are needed to stand inter-

mediate between the narrow expert and the unscientific public.

FUTURE ACTIVITY OF THIS SOCIETY 81

Their number and efficiency ought to be increased, for the good

equally of the expert and of the public. We are all familiar with

the miserable inefficiency of the passage of results from the re-

search laboratory to the plain man. The larger and more catholic

this intermediate group of scientists, the more hopeful will be the

future of science, and the more quickly and accurately will the

results of research become a part of the assets of the nation.

As a matter of fact the writer believes that the weakest place

in our whole scheme of biological publications is just at this point

of bringing together the results of advanced research, and of mak-

ing them intelligible—quickly and accurately intelligible—to the

non-professional or general student of biology. If this Society can

contribute in some degree to this service, it will supply a real need,

and worthily fulfill the prophecy of its past.

For these reasons the writer holds strongly and unhesitatingly

to the view that there is a need for this journal; that this need lies

most decidedly in the direction of continuing our evolution of the

past years, rather than in changing to the narrower policy; and that

our best services to the community may be rendered by undertak-

ing really and intelligently to broaden our efforts rather than to

narrow them. In this view many of those, who have done much

for the Society in the past, heartily concur.

It may be allowable to say in this connection, that there is no

purpose to use the Transactions to exploit a weak, dilute, and

superficially popular type of biology. The research articles will be

just as exact and as numerous as in the past; but there will be an

effort to make, in addition to these, such a series of summaries and

digests as will give the members of the Society, who may not see

large numbers of biological journals, a fair idea of the activities

in the whole field.

Taking our present published membership, only about 25 per

cent have been producers of papers for the Transactions; and this

number includes many who have contributed only brief notes.

These contributors are almost equally divided among physicians,

teachers, and the independent and isolated students. Thus some

three-fourths of our membership maintain their connection with

the Society for the sake of the publications, or thru a desire to

encourage biology. As nearly as I can make out, about ten per

cent of the membership is connected with the larger Universities

82 T. W. GALLOWAY

where they have access to numerous technical publications;

eighteen per cent are physicians in large cities, where presumably

they have similar advantages; twenty per cent are unclassified

members who live near large libraries. The remaining fifty per

cent are physicians, teachers of biology in colleges and high schools

(about fifteen per cent of the whole membership), and isolated

amateurs; few of these have the use of many journals. It is fair

to assume that such a publication as the Transactions have been,

broadened as indicated above, will continue to appeal strongly and

to be highly serviceable to this latter half of our membership. This

suggests that we should seek greater growth among these classes

of students, as well as among our more favored specialists.

An examination of the articles of the last ten years shows

that of the major papers, ten per cent deal primarily with medical

subjects; thirty per cent are papers on subjects closely related to

the work of the physician, as parasitology, bacteriology, impurities

of the water supply, and the like; twenty per cent are concerned

with the microscope as an instrument, and its related helps and

their technic; forty per cent deal with fairly technical phases of

micro—or general—biology, either structural or systematic, appeal-

ing to the better class of research students—amateur and profes-

sional—and to the general teacher of biology. Thirty per cent of

the whole body of articles may fairly be classed as general articles

which would be reasonably interesting to any intelligent person,

tho not a specialist in anything.

This analysis shows that our publication, if continued in the

spirit of the past, may be expected to appeal quite largely to teach-

ers of general biology in colleges and high schools; to physicians

who are still interested in the biology foundational to medicine and

in the general questions of the science; to investigators, amateur

and professional, in any of the various fields of micro-biology to

which we have given attention; to an increasing class of people who

are desirous of showing their general interest and sympathy with

the progress of biology, even tho they may not themselves have the

time or disposition to work as experts in this field.

It is therefore among these four classes of people that our

work should be done in enlarging our membership, and our Trans-

actions should be edited with the real needs of these groups in mind.

The editor, therefore, has this proposition to make to the member-

FUTURE ACTIVITY OF THIS SOCIETY 83

ship:—Without weakening the appeal of the TRANSACTIONS to the

research student, he will undertake to increase its value to the gen-

eral student and teacher, the scientific physician, and the intelligent

layman who wishes to keep in touch with the progress being made

im biology. This can readily be done, and can be done only on

condition that the present members of the Society will assist in the

task of bringing the claims of the quarterly to the notice of those

who ought to be interested—namely, the most up-to-date physicians

of your community, the teacher and advanced students of biology in

the high school or college or university, the isolated person who

works with the microscope for mere love of it, the public or medical

library, and the well-to-do public spirited people who would be

willing to contribute the small amount of our annual dues to aid in

edvancing scientific work.

As one of the steps in redeeming his part of this agreement, the

editor proposes to introduce in each issue a brief review or sum-

mary of the most important items of recent progress in one or more

of the various subdivisions of biology. In this way we can have a

resume of the most important fields every two or three years.

These reviews will not be mere minute abstracts of all the special

papers that appear, such as a bibliographer would wish—altho spe-

cific references will be given to the most important papers; they

will rather be digests of the most important results, with appro-

priate discussion of the conclusions reached and of the tendencies

manifest in the special fields. For this work it is purposed to en-

list some of the best biologists of America, both within and without

the Society.

Among the reviews planned at present are those of the follow-

ing departments of interest :—parasitology ; bacteriology; plant and

animal experimentation; behavior of microscopic animals; re-

searches in the microscopic aspects of heredity; biological technic;

cell studies; plankton, and water purification; comparative physiol-

ogy of the cell and of cell-products ; other phases of medical biology ;

pedagogy of biology. Still other topics may be added as need arises.

Some of these topics are only indirectly related to microscopy,

to be sure; but there is none of them to which the microscope has

not largely contributed. And it is not unfitting that a society orig-

inally organized to exploit the microscope should father an effort

to have a family reunion of the legitimate descendants of the mic-

84 T. W. GALLOWAY

roscope, and to synthesize the special interests.

In conclusion, the incoming secretary feels that he has a right

to invite and expect the full cooperation of every real friend of

the Society. This cooperation should take two forms :—the pre-

senting of some of your very best, suitable papers to the secretary

for use in the Quarterly; and (2) the seeking for memberships

among your friends who may in any way be interested in the work

we are doing. T. W. GALLoway.

bHE.PAYLA, CEASSES, AND-ORDERS OF PLANTS

By CHARLES, E, BESSEY, Ph.D.

The Plant World is here regarded as readily divisible into four-

teen phyla, thirty-three classes, and many more orders. Elsewhere*

I have set forth somewhat fully the principles underlying the tax-

onomy of plants, and have given in considerable detail such a result-

ing classification. In this paper there appears for the first time the

key to the phyla of plants which I have used in some of my Univer-

sity classes in systematic botany. The synoptical view of the phyla,

classes, and orders which follows this key will, I hope, prove help-

ful to teachers and students, as well as others who are interested in

the broader aspects of plant classification.

REY TO THE. PHYRA OF PLANTS

A, Cells typically with poorly developed nuclei and chromatophores; re-

producing by fission and spores; mostly blue-green, brown-green or -

fuliginous (or colorless), never chlorophyll green.

I. Unicellular to filamentous plants. Phylum 1. Myxophyceae.

B. Cells typically with well-developed nuclei and chromatophores; reproduc-

ing by fission and spores, and mostly by gametes also; chlorophyll-

green, sometimes hidden by other coloring matter (or colorless).

I, Plants of but one obvious generation, typically aquatic.

a. The fertilized egg developing into a zygote only.

1. Unicellular, to filamentous many-celled plants (rarely a plate

of cells) : isogamic to heterogamic.

Phylum 2. Protophyceae.

2. Filamentous many-celled plants, mostly breaking up early into

single cells; isogamic. Phylum 3. Zygophyceae.

3. Tubular filamentous (or saccate) coenocytic plants, usually at-

tached basally by rhizoids; isogamic to heterogamic.

Phylum 4. Siphonophyceae.

4. Cellular filamentous to massive plants, attached basally by

rhizoids (or roots): isogamic to heterogamic; the green

color hidden by a brownish pigment.

Phylum 5. Phaeophyceae.

*A Synopsis of Plant Phyla. University of Nebraska Studies. Vol. VII. October, 1907.

The Phyletic Idea in Taxonomy. Science, Vol. XXIX. January, 1909.

Outlines of Plant Phyla. University of Nebraska, Department of Botany. September,

1909.

CHARLES E. BESSEY

b. The fertilized egg developing into a spore-fruit.

1. Cellular filamentous to massive holophytic plants, attached

basally by rhizoids (or roots); heterogamic: the green

color mostly hidden by a red or purple pigment.

Phylum 6. Carpophyceae.

2. Cellular filamentous hysterophytic plants, often much degen-

erated, without chlorophyll; heterogamic.

Phylum 7. Carpomyceteae.

II. Plants of two obvious, alternating generations, typically terrestrial.

a. Gametophyte generation larger, and longer-lived than the depend-

ent sporophyte generation.

1. Gametophytes from prostrate and thalloid to erect leafy

shoots; sporophytes from globose to cylindrical or stalked,

neither expanded nor rooted. Phylum 8. Bryophyta.

b. Gametophyte generation smaller and shorter-lived than the inde-

pendent sporophyte generation.

1. Both generations more or less holophytic and independent.

(a) Gametophytes typically flat and thalloid, normally at-

tached by rhizoids, mostly monoecious; sporophytes

consisting of large-leaved, solid stems, which are

rooted below. Phylum 9. Pteridophyta.

(b) Gametophytes typically flat and thalloid, normally at-

tached by rhizoids, mostly monoecious: sporophytes

consisting ef mostly solid, cylindrical, jointed and

fluted stems, bearing small, whorled leaves at the

nodes, and rooted below. Phylum 10. Calamophyta.

rhizoids or none, often dioecious; sporophytes con-

sisting of solid, cylindrical, continuous (not jointed)

and not fluted stems, bearing small spirally arranged

(or opposite) leaves, and rooted below.

Phylum 11. Lepidophyta.

2. Gametophytes hysterophytic, dependent upon and nourished by

the sporophyte.

(a) Sporophylls open, ovules and seeds naked (gymnosper-

mous).

(1) Gametophytes dioecious: sperms ciliated and

motile; sporophytes producing micfospores

and megaspores in spiral or whorled sporo-

phylls, or these aggregated into cones.

Phylum 12. Cycadophyta.

(2) Gametophytes dioecious: sperms not ciliated, not

motile; sporophytes with sporophylls in

cones. Phylum 13. Strobilophyta.

(b) Sporophylls closed, ovules and seeds covered (angio-

spermous).

THE PHYLA, CLASSES, AND ORDERS OF PLANTS 87

(1) Gametophytes dioecious sperms not ciliated, not

motile; sporophytes with sporophylls in

flowers. Phylum 14. Anthophyta.

SYSTEMATIC ARRANGEMENT

Phylum I. MYXOPHYCEAE

The Slime Algae

Usually blue-green, poorly developed cells, or filaments.

Class 1. ARCHIPLASTIDEAE. (Cyanophyceae) Without nuclear

membrane. (Species about 2000.)

Order Coccogonales. Unicellular.

Order Hormogonales. Filamentous.

Class 2, Ho LopLasTIDEAE. With nuclear membrane. (Species

about 20.)

Order Glaucocystales. Dividing in one plane.

Phylum II. PROTOPHYCEAE.

The Simple Algae

Normally chlorophyll-green, with well developed single

cells, or filaments.

Class 3. ProrococcomeEar. Green Slimes. Unicellular. (Species

about 450.)

Order Palmellales. Cells not in colonies.

Order Coenobiales. Cells in colonies.

Class 4. CONFERVOIDEAE. Filamentous Algae. Filamentous, or a

plane. (Species about 640.)

Order Microsporales. Unbranched.

Order Schizogoniales. Unbranched.

Order Ulvales. Plant a plane or tube.

Order Chaetophorales. Usually branched. Zoospores and

ciliated gametes.

Order Coleochaetales. Branched, fusing into discs.

Phylum III. ZYGOPHYCEAE

The Conjugate Algae

Chlorophyll-green, sluggish filaments, often fragmenting

into single cells.

Class 5. ConyucaTar. Typically filamentous, green plants, with

Class

CHARLES E. BESSEY

cellulose walls. (Species about 1300.)

Order Zygnematales. Pond Scums. Filamentous.

Order Desmidiales. The Desmids. Filaments usually early

fragmenting into single cells.

6. BACCILLARIOIDEAE. The Diatoms. Brownish-green plants,

with silicified walls. (Species about 5700.)

Order Eupodiscales. The Round Diatoms. Filaments com-

monly cylindrical, usually fragmented into single cells.

Order Naviculales. The Long Diatoms. Filaments flattened,

usually fragmented into single cells.

Phylum IV. SIPHONOPHYCEAE

The Tube Algae

Normally chlorophyll-green filaments composed of one or

more coenocytes.

7. VAUCHERIOIDEAE. The Vaucherioid Plants. Filamen-

tous, septate or tubular. (Species about 800.)

Order Cladophorales. The Cladophoras. Septate, the seg-

ments coenocytic.

Order Siphonales. Green Felts. Tubular, irregularly branched,

chlorophyllose.

Order Siphonomycetales. (Phycomyceteae) Filaments tubu-

lar, irregularly branched, chlorophyll-less.

8. BryopsipoIpEAE. The Bryopsidoid Plants. Globular to

stipitate or dendroid, septate or continuous. (Species

about 300. )

Order Valoniales. Globular coenocytes to compound septate

plants. |

Order Dasycladales. Regularly branched, non-septate, mar-

ine plants.

Phylum V. PHAEOPHYCEAE

The Brown Algae

Brown-green filamentous to large, massive plants, marine.

9g. PHAEOSPOREAE. The Kelps. Reproductive organs ex-

ternal, isogamic to heterogamic. (Species about 550.)

Order Ectocarpales. Zoospores and isogametes similar and

motile.

THE PHYLA, CLASSES, AND ORDERS OF PLANTS 89

Order Tilopteridales. Zoospores and heterogametes dissimil-

ar, eggs non-motile.

Order Cutleriales. Zoospores and heterogametes dissimilar

and motile.

Class 10. DicryoTINEAE. Reproductive organs external, hetero-

gamic. (Species about 130.)

Order Dictyotales. Plants erect, flat, leaf-like, zoospores and

gametes non-ciliated.

Class 11. CycLospoREAE. The Rockweeds. Reproductive organs

in sunken conceptacles, heterogamic. (Species about

350.)

Order Fucales. Usually flattish, branched.

Phylum VI. CARPOPHYCEAE

The Higher Algae

Typically red to purple filamentous to massive plants;

mostly marine.

Class 12. BanciomEArE. Antherids and oogones developed from

ordinary cells of plant body; propagation by mono-

spores. Red or purple plants. (Species about 50,

doubtfully belonging here.)

Order Bangiales. One chromatophore in each cell.

Order Rhodochaetales. Several to many chromatophores in

each cell.

Class 13. Florideae. The Red Seaweeds. Antherids and oogones

specially developed ; propagation by tetraspores. Red

or purple plants. (Species about 3000.)

Order Nemalionales. Mostly filamentous plants. Sporo-

phores produced directly from fertilized egg.

Order Gigartinales. Parenchymatous plants; sporophores

produced by nearby auxiliary cells branching in the

surrounding tissues.

Order Rhodymeniales. Filiform, cylindrical, to foliaceous

plants ; sporophores produced by nearby auxiliary cells

growing outward in plant body.

Order Cryptonemiales. Filiform, branched, often complan-

ate; sporophores produced by remote auxiliary cells.

Class 14. CHAROIDEAE. The Stoneworts. Antherids and oogones

Class

CHARLES E. BESSEY

specially developed; no tetraspores. Green plants.

(Species about 160.)

Order Charales. Erect, with whorled branches.

Phylum VII. CARPOMYCETEAE

The Higher Fungi

Terrestrial, chlorophyll-less, filamentous, parasites and

saprophytes, producing spore-fruits.

15. ASCOSPOREAE. The Sac Fungi. Spore-fruits containing

one or more asci with ascospores. (Species about

29,000. )

Order Laboulbeniales. The Beetle Fungi. Erect, minute,

few celled, bearing simple ascigerous fruits.

Order Perisporiales. Primitive Sac Fungi. Filamentous,

with simple, mostly spherical spore-fruits.

Order Pyrenomycetales. Black Fungi. Filamentous, with

mostly compound closed spore-fruits.

Order Pyrenolichenes. The Lower Lichens. Lichen-form-

ing fungi, allied to the preceding families.

Order Hysteriales. The Slit Fungi. True fungi; sapro-

phytic; apothecia opening by a slit.

Order Graphidales. Black Lichens. Lichen-forming fungi,

allied to the preceding families.

Order Phacidiales. The Little Cup Fungi. True Fungi,

spore-fruits open (apothecia).

Order Caliciales. True fungi, and lichen-forming fungi;

apothecia spheroidal, pulverulent.

Order Pezizales. Cup Fungi. True fungi; apothecia at length

cup-shaped, fleshy or leathery.

Order Discolichenes. The Higher Lichens. Lichen-forming

fungi allied to the preceding families.

Order Helvellales. The Helvellas. True fungi; apothecia

open from the first, fleshy or gelatinous.

Order Aspergillales. The Little Tubers. True fungi; spore-

fruits minute, mostly not subterranean. (Related to

Perisporiales. )

Order Tuberales. The Tubers.° True fungi; spore-fruits

large, tuberous, subterranean, fleshy, internally ascig-

erous.

Class

THE PHYLA, CLASSES, AND ORDERS OF PLANTS Ol

Order Exoascales. Pocket Fungi. True fungi; apothecia

much reduced and simplified.

Order Hemiascales. Reduced Sac Fungi. True fungi; no

apothecia, asci single, scattered.

16. TELIOSPOREAE. Brand Fungi. Parasitic, much reduced

plants producing erumpent sori (but no definite spore-

fruits) consisting of telioasci and teliospores. (Species

about 4200.)

Order Uredinales. The Rusts. Typically with sporidia,

pycniospores, aeciospores, urediniospores and _telios-

pores.

Order Ustilaginales. The Smuts. Typically with sporidia

and teliospores.

17. BASIDIOSPOREAE. Basidium Fungi. Spore-fruits con-

taining one or more basidia with basidiospores. (Spe-

cies about 14,000.)

Order Hymenogastrales. The False Tubers. Spore-fruits

large, tuberous, subterranean, fleshy, with internal

hymenium.

Order Phallales. The Stink Horns. Spore-fruits large,

fleshy, at first tuberous and subterranean, later stalked

and emerging.

Order Sclerodermatales. The Hard Puff-balls. Spore-fruits

small to large, roundish, eventually pulverulent.

Order Nidulariales. Bird-nest Fungi. Spore-fruits small,

spherical or top-shaped, leathery, containing peridioles.

Order Lycoperdales. The Puffballs. Spore-fruits large,

fleshy, at first subterranean, later emerging.

Order Hymenomycetales. Toadstools, ete. Spore-fruits

large, umbrella-shaped, bracket-shaped or variously

branched ; hymenium eventually external.

Order Exobasidiales. Reduced and degraded plants related

to the preceding families; basidia undivided.

Order Auriculariales. Ear Fungi. Reduced and degraded

plants related to the preceding families ; basidia divided

transversely.

Order Tremellales. Jelly Fungi. Reduced and degraded

plants related to the preceding families; basidia di-

vided vertically.

g2 CHARLES E. BESSEY

Func Imperrecti. The “Imperfect Fungi.” Including

16,000 to 17,000 species with regard to which our

knowledge is quite imperfect. They are generally re-

regarded as conidial states of Ascosporeae. The

classification here given is merely provisional.

Order Sphaeropsidales. The Spot Fungi. Conidia developed

in pycnidia.

Order Melanconiales. The Black-dot Fungi. Conidia de-

veloped on a stroma.

Order Hyphomycetales. The Moulds. Conidia developed up-

on separate conidiophores which do not form a stroma.

Phylum VIII. BRYOPHYTA

The Mossworts

Chlorophyll-green, small, massive, sexual plants (game-

tophytes), producing a small, spore-bearing generation

(sporophyte).

Class 18. Hepaticae. Liverworts. Gametophytes mostly bilater-

al, often thalloid, creeping; sporophytes usually split-

ting and containing elaters. (Species about 4,000.)

Order Ricciales. The Riccias. Sporophyte globose, sessile,

without columella or elaters.

Order Anthocerotales. Hornworts. Sporophyte elongated,

with a columella and elaters, two-valved.

Order Marchantiales. Liverworts proper. Sporophyte round-

ed, usually short stalked, without columella, inde-

hiscent, containing elaters.

Order Jungermanniales. Scale Mosses. Sporophyte stalked,

four-valved; with elaters.

Class 19. Musct. Mosses. Gametophytes multilateral, usually

erect; sporophytes mostly dehiscent by a circular lid,

and without elaters. (Species about 12,600.)

Order Andreaeales. Black Mosses. Sporophyte — short-

stalked, opening by four to six longitudinal slits.

Order Sphagnales. Peat Mosses. Sporophyte short-stalked,

opening by a circular lid.

Order Bryales. True Mosses. Sporophytes mostly long-

stalked, generally opening by a circular lid, usually

with a peristome.

THE PHYLA, CLASSES, AND ORDERS OF PLANTS 93

Suborder Acrocarpi. Sporophytes terminal on the

main axis of the gametophyte.

Suborder Pleurocarpi. Sporophytes terminal on short

lateral axis of the gametophyte.

Phylum IX. PTERIDOPHYTA

The Ferns

Chlorophyll-green, small, sexual plants (gametophytes),

producing a large-leaved, rooted generation (sporo-

phyte). (Here restricted to the ferns alone, and in-

cluding abotit 2500 species. ) /

Class 20. EusporANGIATAE. Old-fashioned Ferns. Sporangia de-

veloped from internal cells.

Order Ophioglossales. Gametophyte tuberous, subterranean ;

sporophyte with large leaves, some parts sporogenous.

Order Marattiales. Gametophyte flat, green, superficial;

sporophyte with large compound leaves; sporangia

hypophyllous.

Order Isoetales. Gametophytes dioecious, rounded; sporo-

phyte with erect, crowded, narrow leaves; sporangia

epiphyllous, basal.

Class 21. LepTOSPORANGIATAE. Modern Ferns. Sporangia de-

veloped from superficial cells.

Order Filicales. Land Ferns. Spores of one kind; gameto-

phytes foliose, monoecious.

Order Hydropteridales. Water Ferns. Spores of two kinds;

gametophytes dioecious, rounded.

Phylum X. CALAMOPHYTA

The Calamites

Minute sexual plants (gametophytes), producing cylin-

drical, jointed and rooted sporophytes. (Species liv-

ing about 20, but very many extinct. )

Class 22. SPHENOPHYLLINEAE. The Wedge-leaved Calamites.

Palaeozoic trees with solid, jointed stems, long extinct.

Order Sphenophyllales. With the characters of the class.

Class 23. EQuiseTINEAE. The Horsetails. Palaeozoic to recent

plants with hollow, jointed stems.

Class

CHARLES E. BESSEY

Order Equisetales. With the characters of the class.

24. CALAMARINEAE. Old Calamites. Palaeozoic plants,

often trees, with hollow stems, long extinct.

Order Calamariales. With the characters of the class.

Phylum XI. LEPIDOPHYTA

The Lycopods

Minute gametophytes, producing branching, small-leaved,

rooted sporophytes. (Species living about goo, but

very many extinct.)

25. ExicuLtate. Lower Lycopods. Isosporous; leaves with-

out ligules.

Order Lycopodiales. Gametophyte much larger than the spore.

26. LicuLaTarE. Higher Lycopods. Heterosporous; leaves

with ligules.

Order Selaginellales. Small plants; stems not thickening.

Order Lepidodendrales. Palaeozoic and Mesozoic trees, long

extinct.

Phylum X11. “CYCADOPHRY TA

The Cycads

Minute gametophytes developed in naked seeds produced

by the large, leafy-stemmed and rooted sporophytes;

sperms motile. (Species living about 140, but very

many extinct. )

27. PTERIDOSPERMEAE. The Seed Ferns. Palaeozoic, fern-

like plants, long extinct.

Order Pteridospermales. With the characters of the class.

28. CycADINEAE. The Common Cycads. Mesozoic to

present plants with pinnate leaves.

Order Cycadales. With the characters of the class.

29. BENNETTITINEAE. The Flowering-Plant Ancestors.

Mesozoic plants with pinnate leaves, long extinct.

Order Bennettitales. With the characters of the class.

30. CoRDAITINEAE. The Conifer Ancestors. Palaeozoic

to present, trees and shrubs with typically parallel-

veined leaves, mostly long extinct.

THE PHYLA, CLASSES, AND ORDERS OF PLANTS 95

Order Cordaitales. Branching trees with elongated, parallel-

veined leaves. (Extinct.)

Order Ginkgoales. The Maidenhair Trees. Branching trees

with fan-shaped, parallel-veined leaves. (All extinct

but one species. )

Order Gnetales. The Joint-Firs. Anomalous woody plants

of doubtful relationship.

Phylum XIII. STROBILOPHYTA

The Conifers

Minute gametophytes developed in naked seeds produced

by the large, leafy-stemmed and rooted sporophytes ;

sperms not motile. (Species about 450.)

Class 31. Pinomear. Mostly trees with increasing stems and

small mostly persistent leaves; sporophylls mostly in

cones.

Order Coniferales. Conifers proper. Microsporophylls and

megasporophylls in cones.

Order Taxales. The Yews. Microsporophylls in cones,

megasporophylls in very small cones or solitary.

Phylum XIV. ANTHOPHYTA

The Flowering Plants

Minute gametophytes developed in seeds enclosed in

pistils in flowers, produced by the large, leafy-stemmed

and rooted sporophytes ; sperms not motile.

Class 32. MonocotyLepoNEAE. The Monocotyledons. Leaves of

sporophyte alternate from the first, usually parallel

veined ; fibrovascular bundles of stem scattered. (Spe-

cies somewhat more than 20,000. )

Subclass MonocoTyLEDONEAE-HypoGyNnak. Perianth and sta-

mens arising below the carpels: (carpels superior).

Order Alistamatales. Carpels separate, superior to all other

parts of the flower.

Order Liliales. Carpels (usually 3) united forming a com-

pound pistil, superior; perianth in two whorls (of 3

each), corolla-like.

Order Arales. Compound pistil mostly tricarpellary, super-

ior; ovules solitary.

96 CHARLES E. BESSEY

Order Palmales. Compound pistil mostly tricarpellary, su-

perior; ovules usually 1; perianth reduced to rigid

scales.

Order Graminales. Compound pistil reduced to 2 or 3 car-

pels; ovule solitary; perianth reduced to small scales,

or wanting.

Subclass MonocoTYLEDONEAE-EPIGYNAE. Perianth and sta-

mens arising above the carpels: (carpels inferior).

Order Hydrales. Aquatics with an inferior ovary.

Order Iridales. Compound tricarpellary pistil inferior ;

whorls of perianth mostly alike and regular.

Order Orchidales. Compound tricarpellary pistil inferior;

perianth irregular.

Class 33. DicotyLEDONEAE. The Dicotyledons. Leaves of young

sporophyte opposite, sometimes continuing so, usually

reticulate veined; fibrovascular bundles of stem in one

or more rings. (Species about 90,000.)

Subclass DicoTyLEDONEAE-THALAMIFLORAE. Axis of the flow-

er (thalamus) normally cylindrical, hemispherical or

flattened, bearing on its surface the hypogynous peri-

anth, stamens and pistils (or the stamens may be at-

tached to the corolla).

Super-Order Thalamiflorae-Apopetalae-Polycarpellatate. Car-

pels typically many, separate or united; petals

separate.

Super-Order Thalamiflorae - Gamopetalae - Polycarpellatae.

Carpels typically many, united; petals united.

Super-Order Thalamiflorae-Apopetalae-Polycarpellatae. Car-

pels typically two, united ; petals united.

Subclass DicoTyLEDONEAE-CALYCIFLORAE. Axis of the flower

normally expanded into a disk or cup, bearing on its

margin the perianth and stamens, (or the latter may

be attached to the corolla).

Super-Order Calyciflorae-Apopetalae. Petals separate. Car-

pels many to few, separate to united, superior to

inferior.

Super-Order Calyciflorae-Gamopetalae. Petals united. Car-

pels few, united, inferior.

NEW STUDIES OF THE ARRHENURI

By RUTH MARSHALL

Since the publication of “The Arrhenuri of the United States,”

in 1908, the author has been fortunate enough to secure enough

more material to justify the publication of a supplementary paper

on this large genus of water-mites. This material consists of col-

lections from new localities in the United States, and of additional

material from old collecting grounds, together with a few specimens

from other countries. The author’s own collections were made in

Wisconsin and Illinois. A few new places were visited, as the lakes

about Milwaukee ; and more extensive collections were made in pre-

viously known bodies of water, as Lake Wingra, Mirror Lake, Lake

Spooner, Wisconsin, and the Illinois River at Havana. As a result,

several hundred more Arrhenuri have been studied, representing

twenty-eight species; the habitats of eleven species have been ex-

tended beyond state lines, and three new species have been added to

the genus, while four more females have been identified.

Through the courtesy of Dr. E. A. Birge and Mr. Chauncey

Juday, the author was allowed to look over the collections at the

University of Wisconsin, including some material from the United

States Fish Commission, as well as private collections. The author

is also indebted to Miss E. J. Rigdon and to Mr. H. S. Pratt

(through Dr. R. H. Wolcott) for other water-mites. The species

will be taken up in the sequence established in the former paper.

Arrhenurus rotundus Mar.

ELL ties

1908. A. rotundus Marshall. Trans. Am. Mic. Soc., XXVIII:

89-90, pl. VII, figs. 1-4; pl. IX, fig. 128. :

One individual of this small and rare Arrhenurus was found in

a small pool between Madison and Lake Waubesa, Wisconsin, June

24, 1909. It had been found in but two other localities, both in Wis-

consin. . .

Arrhenurus setiger Koen.

1895. A. setiger Koenike. Abh. Ver. Bremen, XIII: 178, pl. 1,

figs. 11-13.

98 RUTH MARSHALL

1901. A. setiger Piersig. Das Tierreich: 113-114.

Much interest attaches to the finding of this species, which was

known before only from material collected in Alberta. One indi-

vidual was found with A. rotundus, which it resembles, in a pool

near Madison, Wisconsin, June 24, 1909. In structure and size it

was found to agree with Dr. Koenike’s description; in color, how-

ever, it was not yellow green, but deep red. :

Arrhenurus bicaudatus Mar.

1908. A. bicaudatus Marshall. Trans. Am. Mic. Soc., XXVIII:

gi, pl. VII, figs. 8-10.

This mite was found for the first time in Wisconsin at Neshota,

September 26, 1908; and later was found in Mirror Lake, Delton,

August 17, 1910. Only one individual was secured in each place.

Arrhenurus infundibularis Mar.

1908. <A. infundibularis Marshall. Trans. Am. Mic. Soc.,

XXVIII: 93-94, pl. VIII, fig. 20; pl. TX, figs. 21, 22.

Three individuals were found in early summer in two collec-

tions from Lake Wingra, at Madison, Wisconsin, the first time for

this locality.

Arrhenurus lyriger Mar.

1908. A. lyriger Marshall. Trans. Am. Mic. Soc., XXVIIT:

94-95, pl. IX, fig. 26; pl. X, figs. 27, 28.

This unusual mite was found in single individuals in three dif-

ferent collections from Mirror Lake, Delton, Wisconsin, the first

specimens from this place.

Arrhenurus scapulatus n. sp.

Pl. I, figs. 2-6

This interesting new species has an unusual form which places

it midway between the subgenus Micrurus and the subgenus Megal-

urus. The oval body has pronounced elevations over the eyes, and

a small, completely closed dorsal shield. At the point where the body

and the appendix join are two large, outstanding humps. The ap-

pendix, short and narrow, but distinctly marked off from the body,

NEW STUDIES OF THE ARRHENURI 99

seems at first to relate the mite to the “long-tailed” Arrhenuri. But

the dorsal surface of the end is depressed, and in it lies a tiny blad-

der-like structure (P, fig. 3), which seems to represent the petiole.

The fourth leg lacks the spur on the fourth segment. These charac-

ters place the species in the subgenus Micrurus. The fourth pair of

epimera have indistinct posterior borders. The palpi have a group

of several blunt, blade-like bristles on the inner side of the second

joint, a character which has suggested the specific name. Outside

of this bunch is found a larger group of slimmer, pointed hairs. The

color in the preserved specimen is deep blue. The length of the body

is I.I mm; the width, 0.8 mm.

Only one individual of this species is known. This was found

in the Richard collection, at the University of Wisconsin, taken at

Mayumba, in the Congo, Africa, June 20, 1890. Its nearest known

relative appears to be A. pectinatus Koenike, another African spe-

cies, which was found in Zanzibar and Madagascar. A. scapulatus

has a longer appendix; but in the form of the body, the position of

the humps, the closed dorsal area, and the blades on the palpi, the

two species show their relationship.

Arrhenurus laticaudatus Mar.

1908. A. laticaudatus Marshall. Trans. Am. Mic. Soc.,

XXVIII: 95-96, pl. IX, figs. 23-25.

One specimen of this rare Arrhenurus was found with A.

lyriger in Mirror Lake, Delton, Wisconsin, Aug. 27, 1910, for the

first time.

Arrhenurus birget Mar.

1903. A. birgei Marshall. Trans. Wis. Acad., XIV: 158-159, pls.

aoa 7,) fig: TOs.

1908. A. birget Marshall. Trans. Am. Mic. Soc., XXVIII:

97-98.

This widely distributed mite was found in three new localities

in eastern Wisconsin—Mukwanago, Neshota and Pewaukee lakes—

in the fall of 1908. It was also found again in Mirror Lake, Delton,

Wisconsin; and in collections made by Dr. E. A. Birge at New

Orleans and Slidell, Louisiana. In all, twenty-five individuals were

obtained.

100 RUTH MARSHALL

Arrhenurus solifer Mar.

1908. A. solifer Marshall. Trans. Am. Mic. Soc., XXVIII:

99-100, pl. XI, figs. 36-38.

One more specimen of this rare species has been found; it was

collected by Mr. H. S. Pratt at Cold Springs Harbor, Long Island,

New York. It was known before only from New Hampshire.

Arrhenurus scutuliformis Mar.

1908. <A. scutuliformis Marshall. Trans. Am. Mic. Soc.,

XXVIII: 100, pl. XI, figs. 39-42.

This is another rare Arrhenurus whose range is now extended.

It had been known only in collections from two localities in Michi-

gan. One more individual was found September 3, 1907, in Bass

Lake, near Spooner, Wisconsin.

Arrhenurus pseudocylindratus Piers.

1903. A. cylindratus Marshall. Trans. Wis. Acad., XIV: 156-

157, pln 17, figs s:

1904. A. pseudocylindratus Piersig. Zool. Cent., XI: 210.

1908. A. pseudocylindratus Marshall. Trans. Am. Mic. Soc.,

XXVIII: 101, pl. XVI, fig. 80.

This large mite occurs again in two collections from Mirror

Lake, Delton, Wisconsin, in late August, 1910. While never found

in large numbers, it appears to be widely distributed in the United

States.

Arrhenurus tahoci n. sp.

Pl. I, fig 7; pl. IT, figs. 11-14; pl. ITI, fig. 31.

This new species of the subgenus Megalurus has a large oval

body and a relatively short and simple appendix. The body is regu-

lar in outline, with a large oval dorsal area, the ends of the furrow

running over onto the appendix. Within the dorsal area is a large

unpaired conical hump, an unusual feature in the “long-tailed”

Arrhenuri. The appendix is broad at the base, and is not sharply

marked off from the body. It narrows in the center, and flares out

at the end in well pronounced side corners, on each of which is a

small hump. A short distance in front of these is a pair of smaller

humps. The genital wings are rather small, but well defined. The

NEW STUDIES OF THE ARRHENURI IOI

fourth epimera are only slightly wider than the third. Palpi and

legs have no especially distinctive characters. The color in the pre-

served material is greenish yellow. The length of the body is 1.28

mm; the width, 0.82 mm.

Only a single male of this new species is known. This is espe-

cially interesting, however, as it was found in a collection from Lake

Tahoe, California, made by Mr. Chauncey Juday, on June 29, 1904.

The nearest related species appears to be A. capillatus Mar., which

has been found in but one locality, a pool near San Francisco. The

two males are somewhat alike in the form of the end of the ap-

pendix ; but this is much more sharply marked off in A. capillatus.

One other Arrhenurus was found with A. tahoei. This was a

female, which in size and form suggested the same species. An ex-

amination of the palpi showed a close similarity of structure, and

this may be taken as proof of its identity. A dorsal view is given

of this specimen. The length is 1.45 mm; the width, 1.23 mm.

Arrhenurus manubriator Mar.

1903. A. manubriator Marshall. Trans. Wis. Acad., XIV:

151-152, pls. 15-17, fig. 3.

1908. A. manubriator Marshall. Trans. Am. Mic. Soc.,

XXVIII: 102-103, pl. XII, figs. 46, 47.

The range of this species has been extended to Ohio, where it

was collected by the United States Fish Commission at Put-in-Bay,

July, 1899, forty-six females, but only one male, being found. It

was also found in Lake Neshota and Lake Pewaukee, near Milwau-

kee, September, 1907, and found again in one collection from Lake

Spooner, Wisconsin, July 16, 1909, in each case in but a single speci-

men of each sex.

Arrhenurus marshalli Piers.

1903. A. globator Marshall. Trans. Wis. Acad., XIV: 148-

£50) pl. 14, fig 1.

1904. A. marshalli Piersig. Zool. Cent., XI: 210.

1908. A. marshalli Marshall. Trans. Am. Mic. Soc., XXVIII:

103-104, pl. XII, figs. 48, 49.

This species continues to lead all others in number of individ-

uals; nearly four hundred were found in these collections, and they

came from seventeen localities. It is also the most widely distributed

102 RUTH MARSHALL

Arrhenurus. It has been found in fifteen states and Canada—the

latter together with Ohio, Arkansas, Texas and Pennsylvania being

now added to the list. Material from the United States Fish Com-

mission gave collections from Long Point, Canada; from Bass

Island, Ohio; and from Erie, Pennsylvania (1899). Collections

made by Dr. E. A. Birge (1903) yielded material from San Marcos,

Texas; from Bantig, Arkansas; and from New Orleans and

Shreveport, Louisiana. This was the first material received from

the first two named states. Two females were found by Mr. H. S.

Pratt at Cold Springs Harbor, Long Island, New York, October 8,

1907. The author’s own collections have extended the range of the

species to eastern Wisconsin, where it was found in Lake Neshota

and Lake Pewaukee, September, 1908. It was also found again in

collections from Lake Spooner, Mirror Lake (Delton), and the

Madison lakes. It appeared in numerous collections from the Ili-

nois River at Havana, during June and July, 1910. New collections

were made in pools at Hennepin, Illinois, August 4, 1910; here the

enormous number of one hundred and twelve were secured.

~ Arrhenurus megalurus Mar.

Plat fesr5-1 7.

1903. A. megalurus Marshall. Trans. Wis. Acad., XIV: 150-

T51, pis; 14,15, fie--2:

1908. A. megalurus Marshall. Trans. Am. Mic. Soc., XXVIII:

105-106, figs. 50-52.

Next to A. marshalli, this species now leads in numbers and in

wide distribution. About two hundred new individuals are here rep-

resented. New York is added to its range, as thirteen individuals

were found at Cold Springs Harbor, Long Island, October 8, 1907,

by Mr. H. S. Pratt. Twenty-two were found in collections from

Slidell, Louisiana, by Dr. E. A. Birge, June, 1904, a new locality for

this state. In the author’s collecting in Wisconsin, it was found in

one new locality, Lake Pewaukee (1908) ; and found again in Mir-

ror Lake (Delton), and in Lake Spooner.

A. megalurus has always been found to be a variable form, in

strong contrast to nearly all other species of the genus. The degree

of development of the humps of the body in both sexes, and the

amount of indentation of the end of the appendix in the male are

NEW STUDIES OF THE ARRHENURI 103

variable characters. The individuals from Slidell all showed the

most extreme development of body humps yet found in the species,

both sexes taking on a very bizarre appearance (figs. 15, 17).

Moreover, some of these individuals were young ones. In strong

contrast to these were two females found in a collection from Lake

Wingra, Madison, Wisconsin, May 28, 1909, which showed scarcely

a trace of the humps, although the normal number of hairs was

present (fig. 16). Judging from the crust of the body, they were

rather young; but they were mature, for they were found in copula

with males.

Arrhenurus parallelatus Mar.

1903. A. parallelatus Marshall. Trans. Wis. Acad., XIV: 154-

155, pls. 16, 17, fig. 6.

1908. <A. parallelatus Marshall. Trans. Am. Mic. Soc.,

XXVIII: 107.

Three new individuals of this species were found again in a

single collection from Lake Spooner, Wisconsin, August 30, 1907.

Arrhenurus expansus Mar.

1908. A. expansus Marshall. Trans. Am. Mic. Soc., XXVIII:

107-108, pl. XIII, figs. 53-55.

This species has been found only in Louisiana. One individual

was found in a new locality, Schrievesport, by Dr. E. A. Birge, in

November, 1903.

Arrhenurus pseudocaudatus Piers.

1904. A. caudatus Marshall. Trans. Wis. Acad., XIV: 521-

523, pl. 40, fig. r.

1905. A. pseudocaudatus Piersig. Zool. Cent., XII: 185.

1908. A. pseudocaudatus Marshall. Trans. Am. Mic. Soc.,

XXVIII: 108.

The single individual from which this species was described was

lost. It is a matter of some interest, therefore, to record that an-

other specimen was found, after much search, in the original col-

lecting ground, the inlet of Lake Spooner (Wisconsin), July 16,

1909. It is noticeable for its color, orange red and deep blue green

replacing the dull green usually found in the individuals of this

sub-genus.

104 RUTH MARSHALL

Arrhenurus semicircularis Piers.

1903. A. securiformis Marshall. Trans. Wis. Acad., XIV:

152-153, pl. 18, fig. 4.

1904. A. semicircularis Piersig. Zool. Cent., XI: 210.

1908. <A. semicircularis Marshall. Trans. Am. Mic. Soc.,

RAV ET: 11a, pl AVG, fe: G43 plo XVII, figoazo:

One more individual of this species was found, May 21, 1909.

This was in a new locality, Lake Wingra, Madison, Wisconsin.

Arrhenurus apetiolata Piers.

Pl. II, figs. 18-20.

1903. A. corniger Marshall. Trans. Wis. Acad., XIV: 155-

EEO. sie ih te. 7.

1904. A. apetiolata Piersig. Zool. Cent., XI: 210.

1908. A. apetiolata Marshall. Trans. Am. Mic. Soc., XVIII:

113-114, pl. XV, fig. 71.

This species occurs abundantly, over one hundred individuals

of the two sexes occurring in eighteen collections, scattered over five

states. Its range has been extended to three new states—Pennsyl-

vania, Ohio, and New York. One male occurred in one of the col-

lections made by the United States Fish Commission at Erie, Penn-

sylvania; and another at Put-in-Bay, Ohio (1899). Three females

were found in material collected by Mr. H. S. Pratt, October 8, 1907,

at Cold Springs Harbor, Long Island. It was found abundantly in

material collected by Dr. E. A. Birge from New Orleans and Slidell,

Louisiana (June, 1903; October, 1904). The species was found

rather commonly (especially the females) in collections at various

points about Havana, Illinois, in July, rgto.

The female previously described (1908) as Arrhenurus apetio-

lata is now known to belong to another species, as yet unidentified,

The attention of the author was called to the error by Mr. G. D.

Nourse. Since that time the true form has been established. Males

of this species have several times been found copulating with fe-

males of other species.

The body of the true A. apetiolata is elliptical. The anterior

border is almost straight ; the posterior end is bowed out with prom-

inent side corners. The enclosed dorsal area is large and slightly

constricted in the anterior part. The three groups of epimera are

NEW STUDIES OF THE ARRHENURI 105

close together; the third pair is broad. The wing-shaped genital

areas on either side are large and slant obliquely out and back. The

length of the body is 0.75 mm; the greatest width, 0.65 mm. The

small size, the elliptical form, the small space between the two pos-

terior pairs of epimera, and the large size and oblique direction of

the genital areas are characters which readily distinguish A. petiola-

ta from the females of other species.

Arrhenurus trifohatus Mar.

1908. A. trifoliatus Marshall. Trans. Am. Mic. Soc., XXVIII:

115, pl. XV, figs. 72-74. _

Two individuals were collected in pools at Hennepin, Illinois,

and one at Havana, in the summer of 1910, the former a new local-

ity for the species.

Arrhenurus reflexus Mar.

1908. A. reflexus Marshall. Trans. Am. Mic. Soc., XXVIII:

117-118, pl. XVII, figs. 84-86.

The range of this rare species has been extended to Ohio, where

one individual was found in a collection made by the United States

Fish Commission, July 18, 1899. One new specimen was also found

in a new locality in Wisconsin—Mirror Lake, Delton, August 27,

IQIO.

Arrhenurus falcicornis Mar.

Pl Ti ie? 2o:

1908. A. falcicornis Marshall. Trans. Am. Mic. Soc., XXVIII:

121-122, pl. XIX, figs. 96-98.

Five individuals of this species were found in the present col-

lections. It is known now for the first time from New York,

through the collection at Cold Spring Harbor, Long Island, made by

Mr. H. S. Pratt, October 8, 1907. It was found in one new locality

in Wisconsin—Lake Mukwanago, October 17, 1908.

Arrhenurus laticornis Mar.

Pl. III, figs. 21-24

1908. A. laticornis Marshall. Trans. Am. Mic. Soc., XXVIII:

122, pl. XIX, figs. 99-101.

100 RUTH MARSHALL

In collections made at Havana, in the Illinois River, in July of

1910, A. laticornis was almost always present in considerable num-

bers, outnumbering the other species, and exceeding the numbers

found in any collections of former years. With the exception of

this species, the Arrhenuri were not found to be as abundant in this

region, either in species or individuals, as they were before the

opening of the Chicago Drainage Canal, judging by the comparison

of the 1910 collections with those made in earlier years by the Illinois

State Biological Station. In August of the same year, the author

found this species also occurring, in smaller numbers, in nearly all

of the collections made at Mirror Lake, Delton, Wisconsin. It had

been found but once before in this body of water. These were the

only two localities where this species has been found since the spe-

cies has been established.

A. laticornis fem. is now known; it occurred in almost equal

numbers with the male in the present collections. The body is nar-

row in the region of the eyes, broad and truncated at the posterior

end. The palpi (like those of the male, but larger) are stout, and

characterized by the great length of the saber-like bristle near the

end of the claw. The second segment has four bristles on the inner

face.

Arrhenurus pollictus n. sp.

Pl. I, figs. 9, 10; pl. III, figs. 25-28.

This new and rare species belongs with the most highly differ-

entiated Arrhenuri, the subgenus Arrhenurus; within this subgenus

it is found in a small but well defined group, characterized by the

presence of a pair of sickle-shaped humps placed dorsally near the

base of the appendix. It resembles the European species A. com-

pactus Piers., and the American species A. falcicornis Mar., from

both of which it is at once distinguished by the form of the petiole,

as well as by its smaller size. The body is oval, the enclosed dorsal

area moderately elevated, its furrow running out to lose itself on

the lateral projections of the appendix. The genital area is narrow

and long, with an indistinct anterior border. It forms conspicuous

rolls on the sides of the body. The epimera have the usual form;

the first pair have sharp anterior points, and the fourth are moder-

ately wide and placed close together. The appendix is short and

NEW STUDIES OF THE ARRHENURI 107

broad with strongly developed posterior angles. On the median

dorsal surface the two small humps are unusually large (fig. 25, H),

as are also the pair on the ventral side (G). The petiole, which is

always so characteristic a feature of these Arrhenuri, has a heavy

dorsally curved central piece (A); around it is wrapped a thin-

ner piece (B), open on the dorsal side. From the latter extend

little bladder-like pieces (D). The hyaline appendage (Hy) is

small and very narrow on the posterior border. The number and

position of the hairs on the appendix is typical of the group.

The fourth leg is short and stout in the first four segments.

The fourth segment, the longest, as usual, has its process uncom-

monly well developed, the end being bent back, like a thumb. The

swimming hairs and bristles of this appendage are quite typical.

The palpi are stout, and present no striking characteristics, unless it

be the rather large number of bristles on the inner side of the second

joint.

The entire length of the body of this mite is 0.78 mm; the width,

0.58 mm. The color is the usual blue green. Only one male is

known, a young but fully formed one. This was found in a small

pool near Kilbourn, Wisconsin, August 7, 1907. The pool fluc-

tuates considerably in size, and has yielded very few Arrhenuri. A

female found at the same time and agreeing with the male in the

structure of the palpi, was identified as belonging also to this species.

Arrhenurus magnicaudatus Mar.

1908. A. magnicaudatus Marshall. Trans. Am. Mic. Soc.,

XXVIII: 123-124, pl. XX, figs. 106-8.

One individual of this large species was found in the present

collections, from a new locality. It was taken from Mirror Lake

(Delton), Wisconsin, August 27, 1910.

Arrhenurus americanus Mar.

Pie Et fig: 20:

1908. A. americanus Marshall. Trans. Am. Mic. Soc.,

XXVIII: 126-127, pl. XXI, figs. 112-117.

Arrhenurus americanus, one of the commonest of the American

species, was found in a large number of the collections. Collections

of the United States Fish Commission, July and August, 1899, have

108 RUTH MARSHALL

extended its range to Canada (Long Point), and to Ohio (Put-in-

Bay). It was found again at Madison, Wisconsin, by Miss E. J.

Rigdon, June, 1909. It was collected in two new localities in east-

ern Wisconsin—Lake Neshota and Lake Pewaukee—in the fall of

1908. It was found again in large numbers in Lake Spooner, Wis-

consin, in 1907 and 1909; and in smaller numbers at Havana, IIli-

nois. It did not appear, however, in the numerous collections made

in Mirror Lake, Delton, Wisconsin, in August, 1910, although it had

been found in less extensive collections in three former years.

Arrhenurus americanus var. major Mar.

Pl, fig: 3

1908. A. major Marshall. Trans. Am. Mic. Soc., XXVIII:

128-129, pl. XX1I, figs. 118-120; pl. XXII, fig. rar.

Although never found in large numbers, the range of this

variety is as great as that of A. americanus. It was found again in

Lake Spooner and in two other new localities in Wisconsin: Lake

Mukwanago (1908), and Mirror Lake, Delton (1909).

Rockford, Ill., Nov. 1, rgto.

CoO AN AWD

NEW STUDIES OF THE ARRHENURI

EXPLANATION OF THE PLATES

Plate I

. Arrhenurus major, palpus.

. Arrhenurus scapulatus, fourth leg.

. Arrhenurus scapulatus, lateral view.

. Arrhenurus scapulatus, ventral view.

. Arrhenurus scapulatus, dorsal view.

. Arrhenurus scapulatus, palpus.

. Arrhenurus pollictus, fourth leg.

. Arrhenurus pollictus, ventral view of the appendix.

IIo

ile

12.

ley

14.

15.

16.

17:

18.

10.

20.

aie

22.

23)

24.

25.

26.

27.

28.

20.

30.

an.

Arrhenurus

RUTH MARSHALL

EXPLANATION OF THE PLATES

Plate II

tahoei fem., dorsal view.

tahoet mas., dorsal view.

tahoei mas., lateral view.

tahoet mas., ventral view.

megalurus mas., extreme development of the humps.

megalurus fem., undeveloped body humps.

magalurus fem., highly developed body humps.

apetiolata fem.,, ventral view.

apetiolata fem., dorsal view.

apetiolata fem., palpus.

Plate III

laticornis fem., epimera and genital plates.

laticornis fem., dorsal view.

laticornis mas., palpus.

laticornis mas., fourth leg.

pollictus mas., lateral view.

pollictus mas., dorsal view.

pollictus fem., epimera and genital plates.

pollictus fem., palpi.

americanus fem., dorsal view.

falcornis, palpus.

Arrhenurus tahoei, fourth leg.

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THE LAKES OF THE GLACIER NATIONAL PARK

By Morton J. ELrop

This and subsequent papers by Professor Elrod, to appear during next year, will

introduce our members to new biotic conditions,;—and to new facts of the distribution

of micro-organisms.—[Ed. ]

AVALANCHE LAKE

Glacier National Park is in northwestern Montana, on the line

of heavy travel. As yet it is visited by a small number of people

only. This Park is a wonderland of mountain crags, dizzy cliffs,

dashing waterfalls, clear lakes, eternal snow and ice, primeval

forest, wild game, blue sky and brilliant sunshine. Here dozens of

sharp mountain summits pierce the azure sky, making a ragged saw

edge on the horizon in every direction. Here the perfectly glorious

heaven’s blue is reflected in the unknown depth of glacial ice, or

in the mirror surface of placid lakes nestling at the base of frown-

ing cliffs of awful height. Here the works of nature have not been

marred by the hand of man. Here is the place where clouds are

made; where are seen the largest remnants of that great ice field

that in past ages covered all northern America; where ice melts

into whispering rivulets which form dashing cascades, these flowing

into lakes of wondrous beauty; where exquisitely beautiful land-

scapes of never ending variety meet the wondering gaze from every

direction; where mountains and sky meet and the freshly formed

dew glistens in shimmering icicle on the bare rocks or form into tiny

droplets which bring life and vigor to the struggling alpine plants.

This, indeed, is a fairy land, where dreams of fantastic things come

true, and where interest and wonder never cease.

The continental divide in the Park makes a very irregular line

as it passes through the quadrangle, alternately pointing to every

direction of the compass. This tortuous path is broken by numer-

ous mountain peaks, in part forming the divide, in larger part

lying a short distance to one side or the other of the actual water-

shed. Extending shoulders from the high mountains enclose pro-

II2 MORTON J. ELROD

tected places. Into these the snows of countless ages have piled by

the aid of drifting winds, forming the large ice masses which give

rise to the name Glacier Park. Some of the peaks are so pre-

cipitous that little snow accumulates, and in late summer are com-

pletely bare. This is generally true of isolated mountains.

The mountains of the Park create profound admiration. A

few rise above 10,000 feet. Many are between 9,000 and 10,000

feet elevation. Many are below 9,000. Their beauty and im-

pressiveness appears in their abruptness. Their crests seem to

have been pushed up out of the earth by sudden upheaval, without

the usual long and sloping ridges such as are frequently seen in

mountain chains. Precipices from 2,000 to 3,000 feet high are not

uncommon, nor can one get out of sight of the frowning cliffs.

Geologically they are extremely interesting, and their clean surfaces

and clear fractures invite study.

The glaciers of the Park are numerous and easily reached. Of

these, large and small, there are about sixty. The size varies from

those covering only a few acres up to the largest, Blackfoot Glacier,

covering some eight or ten square miles. The glaciers of most

interest and importance are Blackfoot, Harrison, Sperry, Pumpelly,

Red Eagle, Grinnell, Chaney, Vulture, Rainbow, Kintla and Agassiz.

These lie high up on the mountain shoulders, and consist of solid ice,

for the most part broken and crevassed, especially noticeable in late

summer. The color of the ice is a deep blue, increasing in intensity

with the depth. Some of the glaciers are easily reached, others only

with extreme difficulty. It is possible to ride on horseback to the

ice of either Sperry or Blackfoot. Hundreds of people visit the

former annually, as it is the glacier nearest the hotel at Lake Mc-

Donald. In fact, it is easily possible to leave the hotel in the morn-

ing, spend a couple of hours on the ice at Sperry, and return to the

hotel in the evening. To reach Blackfoot glacier requires more

time, as it is more remote. But the journey is full of interest every

rod of the way, and a visit to Blackfoot means to revel in some of

the finest scenery of the Park. Except for the traveler with

pack horses the glaciers and other features in the northern end of

the Park are not as yet readily accessible.

There are some 250 lakes, large and small, ranging in size from

a few hundred feet to 12 miles in length, and of varying width.

All the larger lakes are full of fish, and many of the smaller ones

THE LAKES OF THE GLACIER NATIONAL PARK 113

would support fish life, could the fish be placed in their waters.

Falls between lakes and in the streams below prevent the ascent of

fish, hence their absence from many lakes. The lakes are dis-

tributed throughout the Park. From any high summit one may

see a number, even as many as 18 or 20 from one point of view.

It is the purpose of the writer to present in this and succeeding

articles some of the interesting features, and to give the results of

study of the life of some of these lakes. It makes little difference

where we begin, hence as a matter of convenience we shall take

Avalanche Lake.*

Avalanche Lake is reached by trail from Glacier Hotel at Lake

McDonald. The distance from Lake McDonald is about nine miles.

There is no wagon road, and but this one trail, to the lake. The

trail is through dense woods, winding over ridges, crossing rivulets,

leading through long lanes of waving ferns, always up and up, until

the border of the lake is reached.

Our outfit for study consisted of the following articles: a

canvas boat; a small dredge made of a rectangular piece of iron

12x8 inches and provided with a net of large mesh next to the iron

frame, the closed end of the bag of fine bolting cloth; a thermome-

ter in a brass tube; a set of maximum and minimum thermometers ;

a set of wet and dry bulb thermometers for humidity determina-

tion; two hair hygrometers; cameras and plates; chloroform, alco-

hol, formaldehyde, bottles, insect boxes, pins, and nets, and dozens

of other necessary articles.

Our material, provisions and beds were transported on the

backs of seven pack-horses of gentle natures and extensive knowl-

edge of the woods. In addition were four saddle horses, while for

lack of other animals two of the party had to walk. The line of

eleven horses and seven men winding along the trail through the

forest was a fine sight. The bell of the leader gave indication to

those in the rear and often out of sight of the progress of the train.

Owing to the difficulties of transportation, the time consumed

in making and breaking camp and in attending to other duties, and

the difficulties attending visits to lakes with difficult approaches and

of caring for and transporting the collections, only a preliminary

*The reader is recommended to make purchase of the topographic map, Chief Moun-

tain Quadrangle, which will clearly present the location of this and other lakes. Price

five cents in currency. Address the U. S. Geol. Survey, Washington, D. C.

I1t4 MORTON J. ELROD

study could be made, and this under trying circumstances.

The visit was made to Avalanche Lake August 4, Igo, the

party remaining over two days, spending three nights at the lake.

The party consisted of Marcus E. Jones, of Salt Lake City, study-

ing the botany of the region; J. E. Kirkwood, Professor of Forestry

in the University of Montana; Walter Lehman, of Lewiston, Photo-

grapher; Austin Warr, Lehman’s young nephew; the writer, who

was making zoological studies, and the guide, T. H. Scott. Each

person attended to his own line of study, all the collections being

transported by the pack train.

Avalanche Lake lies cradled in the mountains. The scenery

on all sides surpasses portrayal by pen, brush or camera. On the

north and east an unnamed lofty mountain rises abruptly from the

water’s edge. On the opposite side the slopes of Mt. Edwards,

densely wooded and covered with underbrush, rise with very steep

grade to the cliffs near the summit. The rim of Avalanche Basin,

on whose top lies Sperry Glacier, forms the remainder of the moun-

tain boundary. Over the precipices numerous cascades and water-

falls come tumbling down to the lake level, bringing the dissolved

glacial mud and the disintegrating rock. Practically all of the

water of the lake comes from the glacier above. The drainage area

is therefore quite small.

The lake is less than a mile in length, elliptical in outline, with

densely wooded shores. At the upper end avalanches have swept a

portion of the ground clear of trees, for the most part covered with

impassable brush. The lake has been filled at the upper end, the

meandering streams and comparatively level floor giving ample

evidence of this. The old lake bed, together with the material

brought down by the avalanche, cover a mile or more of space

from the lake to the cliffs.

On this composite soil a composite flora of great interest is

found, representing the temperate plants coming up from the lower

valleys, as also the alpine vegetation brought down by the snow

and ice, some of which is making a desperate struggle for an ex-

istence.

The lake is shallow, for a mountain lake, and will doubtless fill

in rapidly. A log jam at the outlet effectually prevents the loss of

any but the soluble matter from the lake, thus aiding in the de-

position of material. Nearly half of the lower portion of the lake

THE LAKES OF THE GLACIER NATIONAL PARK 115

has a depth of only a few feet, from four to ten. At the upper end

the lead showed the greatest depth to be 63 feet.

The soundings and dredgings were made from the canvas boat,

the first boat of any kind to be floated on the lake. The microscopic

life was secured by the dredge. Atg a.m. of August 4, on the way

up the lake, very little microscopic life was found. On the return

down the lake at 2 p. m. life was very abundant. The smaller

species were at or near the surface in the upper lake, the larger

species where the lake was shallow. No attempt was made to gather

material from varying depths, as to do so would be impossible. A

small haul from the bottom was made where the lake was shallow.

While no deductions can be safely drawn from the observa-

tions of one day, yet it seems fairly well established that the

microscopic life comes to the surface during the light of the day,

descending at night. It is possible they seek warmth as well as

light. In Flathead Lake I have seen entomostraca so abundant at

the surface on a hot afternoon that they occurred by the dozen in

a cup of water dipped up at random.

The color of the water was pea-green, rather murky looking

from the sediment coming from the glacier. At 3 p. m. the tem-

perature of the water was 57 degree F; of the air, 70. The per-

centage of saturation of the air in the woods on the lake shore at

2:45 p. m. was 43, at 3:45 it had risen to 62, and at 6:00 to 83. At

night it rose to saturation, and in the morning a light rain was fall-

ing. This, however, was local.

The stones at the lake shore in still and shallow water were

covered with a light green alga, forming long slender threads, sey-

eral inches in length. This is a species of Tetraspora, found grow-

ing also in Lake McDonald. It adheres firmly to the stones, requir-

ing considerable force to separate it from the rock. The plant was

most abundant on the smaller pebbles, from the size of a marble to

those the size of an. egg, and grew most luxuriantly in shallow

water of the lake near the inlets.

The widely distributed snail, Pyramidula strigosa, so abundant

throughout Flathead Lake country, was found sparingly in moist

places. It is doubtless abundant in the wet season. There were no

snakes. A single frog was taken. One monster toad was observed

on the log jam, but slid into the water with a big splash as the net

was moved toward it. Two stone-flies were abundant—a gray

116 MORTON J. ELROD

species along the creeks, a slate black one over the lake. The

former were depositing eggs in the rushing water. Every now

and then one was caught by the splashing water, as the abdomen

was dipped into the stream, and was carried away—sad ending to

to an instinctive effort. This was a day flier. The black one over

the lake flew only in the dusk of the evening.

The lake is a great place for fish. To use the popular expres-

sion, it is full of fish. The most inexperienced fisherman can catch

a mess of trout almost any time. In spring and summer the streams

carry into the lake an abundance of food. The lake is a small speck

in a wide forest area, and great quantities of flying insects fall into

it. The myriads of stone-flies make juicy meals, and dozens were

taken before our eyes while depositing eggs. A stone-fly falling on

the water rarely had more than a few minutes of life after wetting

its wings, before it was taken by a fish. The insects for the adult

fish, and abundance of entomostracan life for minnows, makes it

an ideal place for fish to live. The lake does not have a high eleva-

tion, opens rather early in spring, and does not have as long a period

devoid of insect food for fish as one would imagine. With the in-

dications of abundance of microscopic life as shown by our dredge

it seems possible to keep the number of fish large by continued

stocking, even though tourists may come in much greater numbers

than at the present. The streams afford abundant protection and

shelter for minnows in overflowed and shallow places; and in the

inlet streams is the best fishing, especially when insects are flying.

The annual rise of the lake, as indicated by the shore lines, is

about three feet. The beach is small and narrow, with fine pebbles

for the most part.

To build a road for vehicles from Lake McDonald to Avalanche

Lake will require no great engineering skill, and now that the Gov-

ernment has created the Park we may expect roads to be built.

This will doubtless be one of the first pieces of road, as it opens up

one of the nearest and finest bits of scenery in the Park. It will

also open up a fine collecting field for the botanist and entomologist,

as well as for the student of general zoology. The wooded moun-

tains are full of rare treasures, now exceedingly difficult to procure

because of the trouble in caring for them and getting them safely

out of the woods. With our mode of travel most of the time was

spent in travel and fixing camps. If we spent a day now and then

: 4 :

S ° ; -

7 ~ : i

< ~

Sones ' ?

neers ' d =

* 3 rf A : 4 4

y ‘

+ oe

“

T+

y aia a

_ '

THE LAKES OF THE GLACIER NATIONAL PARK LE.

at a lake collecting microscopic life it could be nothing more than

superficial study of qualitative nature, to determine what the coun-

try affords rather than to work out a problem.

AVALANCHE LAKE STORY ILLUSTRATIONS

Plate IV. Avalanche Lake and Basin from the lower end, showing

cliffs above which lies Sperry glacier from which comes the water of the lake.

—Photo by O. H. Barnhill.

Plate V. Log jam at lower end of Avalanche lake. —Photo by Elrod.

Gorge in Avalanche Creek, outlet of Avalanche Lake.

Plate VI, Fig. 1.

—Photo by N. A. Forsythe.

Plate VI, Fig. 2. Sperry Glacier, from which comes the water of

—Photo by Elrod.

Avalanche Lake.

DEPARTMENT OF SUMMARIES

TO BE DEVOTED TO DIGESTS OF PROGRESS

IN BIOLOGY

While the Transactions will continue to be primarily a Journal of research in micro-

biology, it is recognized that the field has become so broad as to preclude the possibility

of frequent articles in any one of the departments of special interest. Because of this

it will be the policy to present, from time to time, supplementary digests of the progress

being made in the various fields of micro-biology. It is also proposed to introduce similar

summaries of the progress made in some departments not represented in our articles of

research. This is done with the feeling that such reviews will increase the permanent

value of the Transactions to all who may not have access to a large list of technical

biological journals, nor the time to make the survey for themselves. The first of the

reviews in this issue covers a territory finding large application for the physicians and for

students of parasites; the second, for those who teach.—[Editor. ]

RECENT PROGRESS IN PARASITOLOGY

By Henry B. Warp*

In an address delivered some years ago before this Society, I

expressed the view that the problems of medical zoology would ac-

quire increasing interest in the immediate future and this view of

the situation has been abundantly justified by events in recent years.

So much so that the task which has been laid upon me by the Sec-

retary is a difficult one in view of the mass of material which has

been produced. In dealing with recent progress in this field it is

possible, within the limits assigned, to include only the major items,

and even at that some of those must be passed by with hardly more

than a mention. Many investigations of marked value will have to

be omitted entirely, although others may appear to have been given

an amount of space out of keeping with their real value. But it

seems necessary to note particularly those investigations which open

up new portions of the field, or which change our point of view or

accepted interpretation of an individual subject, rather than those

which contribute equally valuable results in the better known por-

tion of the territory. While from one standpoint those papers must

be regarded as most valuable which present results obtained from

a study of the human species, either directly or in its relations with

some other forms of animal life, yet it is evident on a moment’s

thought that comparative studies are as indispensable for a thorough

understanding of the field of medical zoology as they are in other

*Contributions from the Zoological Laboratory, University of Illinois, under the

direction of Henry B. Ward, No. 7.

120 HENRY B. WARD

portions of the biological field. For commercial reasons those rela-

tions which affect the domestic animals have commanded greater

attention than those which affect animals of less value to man.

Consequently the bulk of the literature concerns itself with a

description of the parasites of man and of the domestic animals;

nevertheless the smaller number of papers that deal with other

forms often contain results of equal or greater value.

The importance of studies in medical zoology has been empha-

sized in the scientific mind, during very recent years by the estab-

lishment of schools of tropical medicine following hard on those

which originated very recently also in Liverpool and London.

Furthermore, courses in tropical medicine have been introduced

into medical colleges at home and abroad. Chairs in parasitology,

in protozoology, and in medical entomology have been endowed

abroad and have been established under university influences even

in our own country. Finally the medical journals are devoting in-

creased attention to the animal parasites and animal carriers of

diseases which, only a very few years ago, received but scant at-

tention in their pages or were entirely overlooked.

Some part of this movement is undoubtedly due to the recog-

nition of the scientific importance of problems which previously

had been entirely overlooked. But the major part of it is un-

doubtedly due to the increased importance, in a commercial way,

of tropical countries and peoples. It is doubtful whether sleeping

sickness would have demanded as wide-spread attention or as stren-

uous efforts for its suppression if it had not been a grave commer-

cial danger to the nations developing extensive colonial possessions

in Africa. It is certainly true that our own increased contact with

the Orient has resulted in forcing upon us an interest in the dis-

eases which are primarily characteristic of that territory and

which, by virtue of more frequent intercourse, are constantly being

brought to our shores and endangering our territory.

The extent to which such diseases affect tropical regions may

be readily inferred from the medical reports of those regions. A

sample of this evidence is furnished by the records of our insular

possessions. The Philippine Bureau of Health reports from Bilbid

Prison hospital, during less than two years, 1537 cases of hook

worm, 551 cases of amebic disentery, and 174 cases of malaria,

out of 2,800 cases, or, about 81% due to animal parasites. They

RECENT PROGRESS IN PARASITOLOGY I2I

estimate 80% of the entire population to be infected, a higher

percentage of infection than has as yet been definitely reported

from any other people. The results are seen in the general physical

impoverishment of the people and the high rate of morbidity and

mortality charged to other diseases.

Among the general items which demand consideration at the

outset one may place first of all those organizations or foundations

for the study of the diseases caused by animal parasites or con-

veyed through their medium. These enterprises have taken various

forms, being organized under government control, as parts of uni-

versities or educational institutions, or as separate foundations.

Several European governments have organized bureaus or

commissions and have sent parties into the field for the investiga-

tion of sleeping sickness within the limits of their own territories.

Indeed, it is fair to say that no other disease has attracted as much

governmental attention as this. Unfortunately the experiments have

not as yet yielded very positive results with reference to the pos-

sibility of effecting a cure and the problem of prevention is natur-

ally an exceedingly difficult one when applied to peoples who are

unaccustomed to exercise any sanitary precautions and are un-

able to understand the reason for the simplest regulations which

are promulgated.

The French Commission for the study of Sleeping Sickness

in the French Congo published in 1909, an extended and careful

report on the disease and its transmitting agents in that region.

The report abounds in valuable data, far too extensive to be ab-

stracted here. The German Commission which was sent to Ger-

man East Africa published its report in the same year. The work

is characterized by genuine German thoroughness and is magnificent-

ly illustrated. The Sudan Sleeping Sickness Commission also pub-

lished its final report during the same year and this again con-

tains valuable contributions to the knowledge of the disease-pro-

ducing and the disease-transmitting organisms. Prominent men-

tion should also be made of the Sleeping Sickness Commission of

the Royal Society of London, England, to the work of which ref-

erence is made in detail later.

A most important movement in connection with the suppres-

sion of this disease, produced and transmitted by animal organ-

isms, was the organization of the Sleeping Sickness Bureau in Lon-

122 HENRY B. WARD

don under the direction of the most eminent English investigators

of tropical diseases and of leading Colonial officials of the Imper-

ial government. The Bureau began in 1908 the publication of a

Bulletin giving prompt and accurate reports of field work and of

laboratory experimentation on the trypanosomes which produce

this and similar diseases, the flies which carry them, the chemical

substances employed in therapeutic experiments, and the means

of all sorts for attacking, limiting, or eradicating the disease in

man. These splendid reports have been supplemented by advisory

pamphlets for travelers and residents in tropical Africa, by maps

showing the known distribution of the disease and its carriers, and

by an exhaustive Bibliography of Trypanosomiasis.

The permanent Commission for the Suppression of Uncin-

ariasis in Porto Rico closed its work in 1909 since the legislature

failed to appropriate the necessary funds for its continued activity.

This agency reached 81,375 persons in 1908 and as a result of the

treatment given 25,757 of them were entirely freed of hookworms.

The full meaning of the work done by the Commission may be

grasped when one compares the health reports of Porto Rico and

finds a drop in deaths due to anemia from 11,875 during 1900-01

to 1,785 during 1907-08.

The foundation in our own country of a commission for the

suppression of hookworm disease deserves special mention since

it is the first instance in which large private funds have been given

to an organized movement for the suppression of disease caused

by an animal parasite. The stimulus which has been given to work

throughout the infected territory, and the results already attained

with reference to the prevalence of the disease, the conditions

which favor its spread and those which limit it, and the prophylactic

measures which should be inaugurated and can be promulgated,

are abundant justification of the wisdom which prompted the gift

and the skill which has accompanied the organization of the work

under the direction of the commission.

While the government has detailed a part of the time of one

of its officers in connection with the work of this commission, it

is noteworthy that the work represents otherwise the results of

private activity and as such is unique in the scope of its opera-

tions and in the magnitude of the results it seeks to achieve.

RECENT PROGRESS IN PARASITOLOGY 123

The organization in London of a Society for the Destruction

of Vermin, and the demand for information concerning such pests

as shown by articles in newspapers and popular magazines, both

indicate the growth of a serious public movement to reduce the

numbers of these undesirable elements of the fauna which play a

prominent part in the transmission of disease.

From general literature evidences of the movement may be cited

in the establishment in 1908 of a section of the Journal of Hygiene

which was named Parasitology and devoted exclusively to problems

connected with animal parasites and the diseases with which they are

related.

The foundation in 1907 of the Annals of Tropical Medicine

and Parasitology by the Liverpool School of Tropical Medicine has

furnished another medium for the adequate publication of memoirs

in the field under consideration. Most of the articles published

in it present the results of investigations in medical zoology and

their high character together with the effective appearance of the

publication renders it a most important aid to research in this field.

The foundation by the Instituto Oswaldo Cruz in Rio de Janeiro

of a series of Memorias also deserves especial mention. The mono-

graphs deal largely with medical zoology. They are uniformly

important research contributions in this field and are printed and

illustrated in a manner worthy of their character.

The Wellcome Research Laboratories founded several years

ago at the Gordon Memorial College in Khartoum, had as one

of its primary functions the investigation of problems in para-

sitology. The third annual report (Balfour, 1908) is a series of

splendid articles on protozoology, helminthology, and medical ento-

mology, and contributes in a most valuable manner to both general

and local parasitology. The supplement to this report includes a

review of recent advances in tropical medicine which in admirably

clear and succinct form presents inter alia the results of investi-

gations in medical zoology. This review covers recent work in

parasitology about up to the time at which the present paper begins ;

in that, however, the material is grouped under the head of the

various diseases produced and emphasis is laid on the clinical and

pathological aspects rather than on the structural and biological

features which alone are considered here. One may hope that a

124 HENRY B. WARD

publication so valuable and so satisfactorily printed, may receive

the support necessary for its perpetuation.

Undoubtedly the greatest bibliographic work yet undertaken

in this field is the publication by the U. S. Department of Agricul-

ture of the Index Catalog of Medical and Veterinary Zoology

edited by Stiles and Hassall. The printing of this work, begun in

1902, has involved to date about 2500 pages for the author lists,

which even yet are not finished. Of the subject index one splen-

did number on Trematoda and Trematode Diseases has already

appeared. With the completion of this work, which may be looked

for soon as it is progressing with commendable rapidity, the inves-

tigator will have at command not only a record of the literature

of the past but also a complete subject index admirably arranged

and rich in cross references. This will lighten the burden of trac-

ing down the work of his predecessors and at the same time place

it at his disposal far more completely than individual effort has

been able to cover the field in the past.

Of general literature published on this subject within the past

two years the new edition of Braun’s Human Parasites easily de-

serves first place. Besides the addition of a new section on clinical

features of parasitic infections it was so thoroly revised other-

wise as to be practically a new work and by its translation and

publication in English has become accessible to a much wider circle.

The laboratory manual on Parasitology by Braun and Lihe also

deserves especial mention here. Of. practical value for the pre-

cise determination of various species of human helminthes is a

paper published in these Transactions (Ward, 1908a) in which

a critical abstract was given of the work during the preceding five

years, the discrepancies in the records of different workers were

pointed out, and new data furnished for the precise determination

of the several species. Another general paper (Ward, 1909) in-

cluded a brief discussion of the relation of human progress to

diseases caused by animal parasites.

PROTOZOA

The last two years have been signalized by the appearance of

two admirable general works on the protozoa, which devote especial

attention to a consideration of the pathogenic species as well as to

the relations of parasitic forms to disease. There are the splendid

RECENT PROGRESS IN PARASITOLOGY 125 ;

text on Protozoology, by G. N. Calkins and the magnificent Lehr-

buch der Protozoenkunde by F. Doflein. As a briefer treatise of

almost equal importance may be mentioned a chapter by Minchin

in Allbutt’s System of Medicine, while an article by the author of

the present paper (Ward, 1908) gives a brief outline of the present |

state of knowledge concerning the Protozoa.

So far as parasitic amoebae are concerned the advance in our

knowledge has been very great. Many authors have added to the

list of known forms new species from man or other animals, but the

records have been for the most part so scanty as to afford little

positive ground for the acceptance of the views or the identifica-

tion and determination of the species. Sufficient attention has not

been given to extended studies on the life history, and the work of

Schaudinn from a previous period has not been materially expanded,

although his conclusions have been questioned apparently without

sufficient ground by many later workers. It seems probable that

the diversity of the amoeban parasitic fauna will be shown to be

greater even than the present incomplete records indicate; but

without any means of discrimination any efforts to establish a

system of classification or to differentiate new forms from the

old species will lead to little positive results. Among the many

papers on these forms it is difficult to make a selection. To the

few noted as showing present tendencies, many others might well

be added.

As a result of his investigations Craig (1908) confirms the

views of Schandinn as to the presence in man of at least two

amoebae, one pathogenic and the other a harmless commensal. The

paper includes extensive data on the morphology, reproduction and

habits of these forms and on their differential diagnosis.

Craig (1910a) has also published more extended studies on the

species Paramoeba hominis, which he originally described in 1906

as a parasitic amoeba from the human intestine. He has observed

some features in the life history and describes the process of en-

cystment and certain flagellate stages in development. The occur-

rence of the amoeboid, the encysted, and the flagellate stages at the

same time in fecal material renders it easy to differentiate this

from other intestinal amoebas. The flagellate stage is readily con-

fused with Trichomonas hominis but may be told by the absence of

126 HENRY B. WARD

an undulating membrane, the presence of one flagellum and the

spherical form in Paramoeba hominis.

Elmassian (1909), Hartmann (1908), and Koidzumi (1909)

are among others who have made more precise studies of new in-

testinal amoebae from man.

In one direction considerable advance has been made. Vari-

ous efforts towards the cultivation of amoebae under laboratory

conditions have resulted in partial success, and the time is at hand

when this group of organisms can be handled like bacteria and

flagellates under conditions of laboratory experimentation. Prom-

inent among the workers in this line have been Musgrave, Nagler

and Walker. In commenting on these experiments it should be added

that they have given ground for confusion since they deal only

with the most uniform and least easily differentiated stage of exist-

ence in the life cycle of the organisms. Futhermore, it still remains

to be shown that under artificial conditions these organisms mani-

fest the form and display the activities which they possess under

natural conditions. Several authors have suggested that the ap-

pearance and habits of the organisms are radically modified by

cultural conditions, and that great caution must be exercised in

drawing conclusions with reference to their structure and habits in

the normal host.

Investigations concerning various trypanosomes have multi-

plied enormously during the last two years. A mere mention of the

most important contributions in this field would exhaust the pos-

sible limits of this paper, and the bibliography on the subject for

recent years constitutes a substantial volume which has already

been mentioned under the publications of the Sleeping Sickness

Bureau. Of particular importance may be mentioned the experi-

ments of Nuttall (1908), Breinl and Hindle (1910), and others

who have succeeded in transmitting rat trypanosomes through the

medium of rat fleas and lice. The early view that the transmission

of the trypanosome of sleeping sickness was purely a mechanical

transfer has given way under the demonstration that the fly be-

comes infective only after a longer period and remains infected

through a considerable interval of time. Reference is made to

details later in this paper.

The cultivation of Trypanosomes in artificial media was first

successfully carried out by MacNeal and Novy in 1903, and in

RECENT PROGRESS IN PARASITOLOGY 127

subsequent years these same authors have published many details

on the technic and have extended the method from the trypanosomes

of the rat to other mammalian species and to those of birds. Other

investigators have repeated these experiments with varying suc-

ceess but least satisfactorily with the pathogenic species. It has

not yet been shown that cultures aid in the differentiation of spe-

cies or strains, or that it is possible to achieve in any way immunity

with the organisms thus grown. Nicolle has also used the method

to cultivate the protozoon parasite of infantile splenic anemia, and

several authors have recently succeeded in cultivating Treponema

pallidum in an artificial medium.

Of theoretical interest in considering the origin of trypanosomes

is the view of Minchin (1908) whose extensive studies on the group

deserve prominent mention. He is of the opinion that their ances-

tors were flagellates parasitic in the gut of vertebrates and the

cysts were taken in with food. Through the alimentary wall they

reached the circulatory system. Then ingested by blood sucking

insects, they encysted in the gut and with the insect feces came

again into the gut of the final host. Or they accommodated them-

selves to life in the insect gut and wandered into the proboscis,

only to be transferred by the bite of the host so that encystment

became unnecessary and dropped out. In other than blood sucking

insects the flagellates are perhaps neotenic larvae.

Kleine (1909) was the first to demonstrate the infective powers

of tsetse flies for trypanosomes after long intervals, and confirmed

these findings by repeated experiments. It was impossible to escape

the conclusion that Trypanosoma gambiense undergoes a develop-

mental cycle in its transmitting agent, Glossina palpalis.

Kleine’s results were confirmed almost immediately by Bruce

who found flies infective after 16, 19 and 22 days. Bruce showed

that monkeys of the genus Cercopithecus may be used as test ani-

mals for human trypanosomiasis. Cercopithecus ruber is most suit-

able and has the same relation to sleeping sickness as the guinea-pig

to tuberculosis. But “inoculation into test animals is an uncertain

diagnostic method at the best and the incubation period is long;

it is rarely needed in human trypanosomiasis.” As a test of the

permanence of cure or to indicate promptly any relapse, inocula-

tion into this animal would be most useful. ;

,128 HENRY B. WARD

Kleine infected flies hatched from pupae by feeding them on

sick monkeys and then conveying sleeping sickness to healthy mon-

keys. From his work it appears clear that Glossina morsitans is

also a transmitter of Trypanosoma gambiense. The large number

of unsuccessful experiments on hereditary transmission renders its

occurrence rather unlikely. Mechanical transmission does not occur

after an interval of 18 hours.

On the life history of the trypanosomes, Minchin and Thomp-

son (1910) have published the results of studies in the transmission

of Trypanosoma Lewisi by the rat flea. They show that the rat flea

can transmit the flagellate from infected to non-infected rats, that

the transmission takes place by the cyclical and not, so far as present

evidence goes, by the direct method. The incubation period for the

developmental cycle of the trypanosome covers at least 6 or 7 days.

The Bulletin of the Sleeping Sickness Bureau gives (No. 15,

page 89) the following table showing the facts at present known

concerning the cyclical development of trypanosomes.

METHODS OF TRYPANOSOME TRANSMISSION SO FAR ASCERTAINED

Species of Insect. Duration of Place of Duration of Observer.

trypanosome. non-infective development. infectivity.

period.

T. Brucei (?) G., palpalis. 20 days at least 83 Kleine.

P ays.

T. gambiense G. palpalis. 16, 19 and Intestine. At least 75 Bruce,

and a trypan- 22 days. days. Hamerton,

osome of di- Bateman and

morphon type. : d Mackie.

T. gambiense. G. palpalis. 18 days. Intestine. Kleine.

T. cazalboui. G. palpalis. 7 days. Probocis. At least 2% Bouffard.

Tae: months. Minchin and

T. lewisi. Ceratophyllus 6 days. Begins in At least 6 Thomson.

fasciatus. rectum. weeks.

“In the cyclical method” says Minchin, “the invertebrate is

more than a suitable instrument in the transmission; it acts as a

host in which the parasite establishes itself and maintains the exist-

ence of its species.” The evidence cited above makes it probable

that this method is the usual one in the transmission of trypano-

somes and demonstrates the great importance of the msect trans-

mutters.

Prowazek (1909) defined carefully the often confused genera

Herpetomonas, Crithidia and Trypanosoma, and discussed the mor-

phology and development of the last named genus. The paper con-

tains a wealth of important structural details on these forms.

RECENT PROGRESS IN PARASITOLOGY 129

The work of a number of investigators has shown that trypa-

nosomes acquire a specific chemo-resistance to a certain drug which

persists unchanged through long periods. Straines with double

or triple resistance may be produced experimentally; even though

mixed all such strains remain separate and capable of isolation after

repeated passages through infected animals.

Chagas (1909) has discovered a new flagellate which he names

Schizotrypanum Cruzi, in blood of children of Minas Geraes, Bra-

zil. It is transmitted by a bloodsucking bug, common in the inhab-

ited houses of the region. This bug, Conorrhinus (megistus?), is a

true host of the flagellate which requires at least eight days to com-

plete its developmental cycle in the insect. The parasite was trans-

mitted experimentally, and in some cases at least by bite of the

infected bug, to rabbits, dogs, guinea-pigs and monkeys.

Minchin and Woodcock (1910) have worked on the blood para-

sites of fishes and by comparison of the minute structure and

staining reactions are able to demonstrate essential differences be-

tween the nuclei of gregarines and trypanosomes. The nucleus of

Halteridium is clearly different from that of a gregarine and on

the other hand remarkably like that of the trophonucleus of a trypa-

nosome in being of the karyosomatic type. The immediate conclu-

sions from these results are of primary importance in indicating the

closer and more distant relationships of the groups cited. They also

speak distinctly against the proposal to remove the group of Hae-

mosporidia from the Sporozoa and place it with the trypanosomes

under the Flagellata.

Much work has been done recently on the interesting forms

known as spirilla or spirochaetes. Many new species have been

described, especially from bivalves where they occur in the crystalline

style (Fantham, 1908). Authors still differ widely regarding the

structure and relationship of these forms. Some new details re-

garding the spirochaete parasites of higher form have been deter-

mined by the use of dark field illumination, etc., but no agreement

is reached as to their systematic position or their methods of repro-

duction. The number of papers and notes which have appeared on

this group is so enormous that it is impossible to give, within the

limits at the command of this article, any adequate idea of the

work done, but the net results do not include many items of striking

general importance, while even among the most able investigators

130 HENRY B. WARD

one finds diametrically opposed views regarding some simple fea-

tures of structure as well as concerning the existence or non-exist-

ence of supposed stages and forms in the life history.

Concerning the spirochaete of syphilis described by Schaudinn

the most extended study has been made by Krzysztalowicz and Sied-

lecki (1908).

They maintain that no flagella are present and an undulating

membrane is doubtful; the body is not stiff, though little moveable.

Chromatin is distributed through the entire body. Multiplication is

by longitudinal division. The so-called microgametes originate by

cutting off the ends of elongate individuals, i. e., by transverse

division or budding; the role of these microgametes, as also of

the forms called macrogametes is left unsettled. The spirochaetes

belong to the Flagellata, constituting a special group of Spiroflag-

ellata.

The discovery that relapsing fever is due to an organism of the

type of a spirillum or spirochaete falls before the period covered by

this paper as also the demonstration that several species of the

organism exist which are related individually to fevers of different

regions. Much evidence has been accumulated during the past two

years on the organisms of this group and their differentiation from

each other. Darling (1909) has studied carefully the species which

is the cause of the relapsing fever of Panama. He shows that it is

distinct from the species Spirochaeta Obermeieri, Sp. Duttoni and

Sp. Cartert though belonging to the same group. The morphology

of this species shows considerable variation and no positive identi-

fication can be made on morphological grounds. The question of

the genetic relationship of these forms is still swb judice. Darling

considers Spirochaeta as more closely related to the bacteria than

to the protozoa.

Among the important studies on haematozoa of various sorts

conducted by Balfour (1908) especial mention should be made of

his investigations on spirochaetes in fowls. He was able to correct

many erroneous and partial observations of previous investigators

and to demonstrate an intracorpuscular stage of the spirochaete which

in all probability may be regarded as a phase in the life history of

the organism. Contemporary and subsequent investigations tend to

establish the occurrence of such intracorpuscular stages in the life

cycle of all spirochaetes and to regard them as stages leading to the

RECENT PROGRESS IN PARASITOLOGY 131

formation of the very minute bodies out of which a new generation

of spirochaetes is evolved after transfer to a new host.

Our knowledge of the distribution and life history as well as

of the mutual relationships of the Sporozoa has been greatly ex-

tended by the work of Léger. His work deals largely with the

lower forms and is too extended to permit of detailed citation in

this article.

Patton has published valuable papers on the flagellate and

sporozoan parasites of various animals. His experiments at inocu-

lating dogs with Leishmania Donovani; the parasite of Kala Azar in

man, have demonstrated the non-susceptibility of that host. He

favors including this parasite with numerous others in the common

genus, Herpetomonas, a procedure which can not find favor with

zoologists. In a most lucid manner he discusses recently (1909) the

life history of the Kala Azar parasite and its relations to other

organisms.

The group of organisms represented by the parasite of Kala

Azar has received an important addition in the discovery of a new

form by Nicolle (1909).

This species which he names Leishmania infantum, is the cause

of spenomegaly in children, in Tunis and Southern Italy. It is

found in the spleen, liver, bone marrow, but rarely in peripheral

blood, either free or in uninuclear leucocytes. In cultures it develops

into forms similar to Herpetomonas, which multiply actively by

cross fission and form rosettes by agglutination at the flagellar

ends. It can be transmitted to apes and dogs, and in his opinion

is originally a parasite of the latter.

The organism of Kala Azar has also been studied in culture

following the original suggestions of Patton and Christopher. New

experiments by Row show that active multiplication in blood-serum

cultures at 25-28° C. produces colonies of spindle shaped individ-

uals in 48 hours which become free after formation of a flagellum

and manifest two forms, thick and thin. Cultures from old cases

of oriental sore develop parasites of a spindle shaped, non-flagel-

late stage only, and after unequal division produce 4 long and 4

short individuals of which the latter are short-lived. He also dis-

cusses the difference between Helcosoma tropicum and Leishmania

Donovan.

132 HENRY B. WARD

Cummins (in Balfour, 1908) discusses the presence of this

parasite in the Sudan, its probable introduction, distribution, and

significance.

In 1908 Darling reported a new parasitic protozoon encoun-

tered in the Canal Zone. The micro-organism, which he names

Histoplasma capsulatum, gives rise to a fatal infectious disease, his-

toplasmosis, among natives of tropical America. The parasite is

described in detail but no facts were elucidated concerning its life

history or the mode of infection. In a later paper (1909a) Darling

gives further data on, the morphology of the organism which he

describes as small, round, or oval, 1 to 4 microns in diameter,

possessing a polymorphus, chromatic nucleus, basophilic cytoplasm

and achromatic spaces all enclosed within an achromatic refractile

capsule. The form and arrangement of the nuclear chromatin and

the lack of a chromatic rod differentiate it from the Leishman

Donovan body. It occurs in the endothelial cells of smaller lymph

and blood vessels in enormous numbers and causes necroses of liver

and lymph nodes, splenomegaly, and pseudo-gramulomata of the

lungs and intestines, with ulceration of the latter. The organism

may prove to be identical with Helcosoma Donovani and in any

event appears to be closely related to it.

Cole, Hadley and Kirkpatrick (1910) have published an exten-

sive memoir on blackhead in turkeys, in which they discuss broadly

the general topic of avian coccidiosis. Their work is full of import-

ant observations on the organisms and their relation to disease.

They discuss at length the disease, the gross pathological changes

involved and the microscopical findings. They interpret the organ-

ism, Amoeba meleagridis, originally discovered and investigated by

Theobald Smith, as merely a stage in the development of the coc-

cidium to which they attribute the disease. They record the pres-

ence of the same coccidium in a large series of avian hosts and

also in mice, rats and rabbits (“probably”). Finally they outline

the developmental cycle and record a series of experiments in trans-

mission. Such radical modifications in accepted views will require

some corroborative evidence before they can be finally accepted.

Smith has pointed out very recently that important questions must

be answered before the conclusions of Cole and his associates can

be accepted.

RECENT PROGRESS IN PARASITOLOGY E33

The sexual forms of the malarial plasmodia from human blood

are discussed by Craig (1910) who reports also the effect of

quinin upon these sexual stages and finds that their disappearance

from the peripheral blood does not indicate their destruction. It

seems probable that they are merely driven to the spleen and bone

marrow. In any event quinin produces no changes whatever in

morphology of the fully developed gametes.

Koch showed in 1908 that in Babesia (Piroplasma) bigemina

one stage occurs in the eggs of the tick and is transmitted thereby to

the second generation of ticks. Carter found that Spirochaete Dut-

toni multiplied in the ova of ticks and thus infected the second gene-

ration. In flies this may be the usual or the only method of trans-

mission from one generation of the host to the next.

Extensive experimental researches on the drug treatment of

Piroplasma have come from Nuttall’s laboratory at Cambridge,

England. Other contributions from the same source deal with the

development of Piroplasma canis, P. bovis, and other species, and

with Theileria parva.

Hepatozoon perniciosum (n. g., n. sp.) a hemogregamine patho-

genic for white rats has been studied by Miller (1908) who finds

the sexual cycle in a mite (Lelaps echidninus).

The great variety of records concerning the occurrence of Sar-

cosporidia in man of which only two or three cases are positively

known, lends special interest to the report of Darling (1909b)

concerning a new case. The author regards it as a chance infection

of a species different from those hitherto reported from man, but

not definitely determined. The sporozoa disappeared from the

muscle fibers of the subject and this probably took place on a date

when the sporozoites were observed escaping from their capsule.

He concludes that such infection gives rise to little or no discom-

fort. The case was complicated by intercurrent typhoid fever.

Renewed investigations by Bensen (1909) demonstrate the

specific distinctness of Trichomonas intestinalis and Tr. vaginalis.

A study of the life history shows that Trichomonas intestinalis

throws off its flagella and becomes amoeboid. After encystment

a new generation is formed. Tr. vaginalis encysts in the flagellate

stage and after that becomes amoeboid. The same author (1908)

has monographed the genus Lamblia.

134 HENRY B. WARD

Among the ciliates there is to report a careful study of the

genus Opalina by Metcalf (1909) who gives a synopsis of the

species, a description of the structural features, cysts, sexual forms,

and biological characteristics of each species. Infection is direct

and easy with all species of Batrachia.

HELMINTHES—GENERAL

Shipley has carried out extensive investigations on the internal

parasites of the grouse and related birds. The work was under-

taken with a view to ascertaining the cause of grouse disease of

the British Isles. Several papers give information concerning the

abundance and varieties of the grouse entozoa together with data on

their structure. Thus far the life history has not been elucidated.

The relations which these parasites bear toward disease 1s the sub-

ject of a special paper (Shipley 1908) in which the author brings

together also the results of earlier workers on similar relations

between other entozoa and other hosts, including man. On the

basis of the evidence collected concerning man he concludes that

“the passage of bacteria which set up intestinal disease is immensely

aided by any agent which causes a lesion in the mucosa. Such

lesions are normally caused in man—by entozoa.”

Pratt (1909) has published a careful review of our present

knowledge regarding the cuticula and subcuticula of Trematoda and

Cestoda. He concludes that the cuticula in these worms is not

homologous to the similarly designated layer in segmented worms

and arthropods but is the peripheral portion of the parenchyma and

is formed by secretions of the latter tissue. He regards the sub-

cuticula as a genetic portion of the same tissue and not as epithelial in

origin. The subcuticular cells are regularly absent in the earliest

larval stages and often in adults (monogenetic trematodes) ; they

probably form an indifferent embryonic tissue to be specialized in

the later growth of the worm.

Our knowledge concerning the geographic distribution of hel-

minthes has been notably widened during the past two years. In

addition to articles on separate groups by authors quoted in the

special sections of this survey, mention should also be made of work

on South African species by L. H. Gough, on Australian species

by Georgianna Sweet, on South American species by Daday, on

RECENT PROGRESS IN PARASITOLOGY 135

species from Roumania by N. Léon, on species from Bermuda and

Florida by E. Linton and H. S. Pratt, on forms from Central

Africa by Balfour, Leiper and Wenyon.

TREMATODA

No doubt the most striking contribution to the structure of

the flukes which has been made within recent years is the publica-

tion by Goldschmidt (1909) of studies which modify radically our

interpretation of various organs in these forms.

As a result of careful and extensive investigation the author

claims it is necessary to abandon the old view that the shell gland

forms the egg shell. The droplets in the yolk cells are shell-

secretion, and the shell material is produced by the so-called vitel-

laria, while the shell gland secretes a watery substance in which

the eggs are suspended in the uterus. The so-called yolk cells play

no important part in the nourishment of the young embryo.

Stiles and Goldberger (1910) publish the results of an anatom-

ical study of Cladorchis Watson: from man, renamed Watsons

Watsoni, and of allied mammalian species all belonging to the new

superfamily Paramphistomoidea, which corresponds to the family

Paramphistomidae as previously conceived. Many new points in

the morphology of these forms are discussed including a large peri-

suctorial cavity of varied structure which to these authors is sug-

gestive of a rudimentary body cavity absent by definition in the

group of Plathelminthes. The authors use the projection method

in demonstrating the organology of these thick-bodied and not easily

studied species. In this paper Stiles proposes a new and exceed-

ingly important plan for designating the topography of trematodes

under a new terminology which may be outlined in the author’s

words as follows:

“In brief, longitudinal and transverse straight lines are drawn

at the periphery of the various organs; the longitudinal lines bound

fields the transverse lines bound zones. Portions of the body bound-

ed by other than straight lines (as that portion bounded by the

intestinal ceca) are termed “areas.” Organs are then located with

reference to these fields, zones, and areas. Thus, the testicular

zones may coincide, overlap, abut, or be separate ; the testicular fields

may coincide, overlap, abut, or be separate. An ovary may

be described as in the pretesticular, testicular, or posttesticular zone,

136 HENRY B. WARD

or in the extratesticular, testicular, or intertesticular field; a given

organ may be in the prebifurcal zone, preacetabular zone, postace-

tabular, postovarial, postuterine zone, etc. The body is also divided

into five transverse zones, each representing 20 per cent of the body

length; these zones, beginning at the oral pole, are called the first,

second, third, fourth and fifth. It is believed that by aid of this

system, descriptions may be made more exact than they frequently

are at present, and that, especially in the case of tabular keys, the

system will be found useful. A key to the figured species of dis-

tomes is now being formed on this principle; a preliminary study,

based upon about 150 illustrations, has thus far been found to be

very satisfactory.”

Lihe (1909) has brought together all the species of trematodes

found in the fresh-water fauna of Germany into an admirable

brief systematic manual which will be of great service elsewhere

also.

Among taxonomic and distributional studies of trematodes

from hosts other than man especial mention must be made of the

work of Braun and his coadjutors who have added greatly to our

knowledge of the structure and relationships within this group.

Similar valuable contributions in this field have been made by

Odhner on African species, Luhe on numerous families and genera

chiefly of European forms, Lebour and Nicoll on the trematodes of

the British Isles and adjacent waters, Dietz on the Echinostomidae,

etc. Nicoll (1909) has contributed prominently to our knowledge

of the systematic arrangement of the digenetic trematodes. Ssinitzin

(1909) has discussed the origin of the trematodes and their methods

of reproduction on the basis of his investigations in this field.

Much work has been done in determining the diagnostic fea-

tures of various known human parasites from this group. This has

resulted in marked improvement in the description of known forms

and also in the differentiation of new and closely related species.

Thus Looss (1907) has demonstrated that under the familiar name

Opisthorchis sinensis two human parasites have been confused by

recent authors. Baelz in 1883 separated them correctly and named

them Distomum innocuum and D. endemicum. Looss now makes

a new genus Clonorchis to hold both species.

Another instance of advance in knowledge by careful investi-

gation of new material concerns the species Fasciolopsis Buskti and

RECENT PROGRESS IN PARASITOLOGY 137

F. Rathouisi. The actual existence of the latter has recently been

questioned by Odhner (1909) who was able to subject the original

material to re-examination. About the same time, however, with

the aid of material collected by Drs. Goddard and Jeffreys in China,

Ward established (1909a) the validity of Poirier’s species Distoma

Rathouisi not reported since the original record; errors in the

original description were corrected and the differential characters

between this and related species were determined. Important data

concerning the distribution, frequence, and relation of these species

to human disease were quoted from the records of the physicians

named. It seems likely that the new species reported by Roden-

waldt (1909) Fasciolopsis Fiillebornii, is actually identical with the

form originally described by Poirier. In fact the author himself

calls attention to the similarity between his form and the generic

diagnosis given by Odhner for the genus Fasciolopsis. Rodenwaldt

reports that in the Marine Hospital at Hamburg an Hindoo evacu-

ated among other worms three to which Odhner’s generic diagnosis

fits except the coiled form of the much stronger cirrus sac and the

lack of coecal sac in the sperm vesicle. The almost round cephalic

tip is not plainly set off from the body. A broad flat main excretary

vessel extends from the posterior end to the shell gland, dorsal to

testes. At regular intervals it gives off paired cross branches, a

strong pair in the region of the anterior testes with numerous lateral

twigs which go up into the anterior region. At the shell gland the

main vessel splits into two branches extending anteriad. The con-

dition of the cirrus sac which in size and volume far surpass that

of all known trematodes, supports the view of various authors

against Odhner that D. Rathouisi cannot simply be rejected.

The life history of various flukes has also received attention.

A most important contribution in this field has been made by

Ssinitzin (1909a) on the aberrant Gastrostomidae. Among human

parasites Garrison and Leynes (1909) have studied the miracidium

of Paragonimus Westermanu under various physical conditions.

In their experiments when eggs are taken fresh from sputum,

the cleavage is not finished; in 15 days after evacuation the mira-

cidia swarm out; the temperature optimum is 25-28° C. Bad water

and high temperature delay development and ordinarily 25 to 45

days are necessary for hatching, though this is very variable. Mira-

cidia in egg shells remain alive up to 160 days. High temperature

138 HENRY B. WARD

is very dangerous (37°) but over 15° C. is necessary for develop-

ment; below 10°C. the movement stops, but the embryo remains

alive. Direct sunlight is dangerous and any light is unnecessary

for development. Weak saline solution is entirely innocuous but

dessication proves rapidly fatal.

Garrison (1908) has also described a new fluke parasitic in

man. Ortman (1908) has studied the early development of the

fluke embryo and followed in detail the origin of the embryonic

membranes and of the various organs.

The discovery of a new human blood fluke in China and Japan

which just antedated the limits of our summary has led to extended

studies on this organism with a view to determining its distribution

and hygienic importance. Prominent among these investigations

may be mentioned the work of Tsuchiya (1908) who has published

a detailed discussion of Schistosoma japonicum, especially in its

clinical and pathological aspects. Infection takes place through

drinking water and is more frequent in men than in women, and

most of all in children of the lower classes. Enlargement of the

liver and spleen and swelling of the body due to ascites, are the

symptoms. In chronic cases intestinal hemorrhage and general

weakness (marasmus) lead to a fatal termination. The parasite is

very common in cats and dogs of infected districts. The eggs, de-

posited in the stomach or intestinal wall by parasites are set free

directly into vessels and by rupture of these reach the tissues. The

splenic enlargement is caused by waste products of the parasites.

The eggs, 85.7x64.3 microns, are smaller than in S. haematobium,

smooth, never having a spine. The male has no papillae or warts

on body. The length of body and size of suckers is much greater

than in S. haematobium, but variable according to the host. Vitel-

laria and unpaired crus intestinale of female less extended, uterus

and paired crura more so than in S. haematobium. The disease is

wide-spread in China and the Philippines. Prophylaxis consists

in avoidance of impure drinking water. The work of Katsurada and

Hashegawa, noted later, throws grave doubts on this last conclusion.

A new species of human blood fluke, Schistosoma Mansoni, was

established by Sambon in 1907 on the basis primarily of the struc-

ture of the ova which possess a lateral spine and also differ in size

from the long-known Egyptian species (S. haematobium) and the

more recently discovered species of the Orient (S. japonicum). In

RECENT PROGRESS IN PARASITOLOGY 139

an extended critique Looss (1908) reached the conclusion that the

evidence is absolutely insufficient to point out the existence of a

distinct species in the West Indies and certain parts of Africa as

claimed by Sambon. The latter author (1909) replied in detail

to the criticisms of Looss and adduced further evidence to show

the correctness of his views regarding the existence of a new spe-

cies in the regions noted and the distinctness of this species, S.

Mansoni, from forms described previously.

By the work of Piraja da Silva (1908) many details of struc-

ture in the adult worms of Sch. Mansoni are reported with such

clearness and apparent accuracy that it is difficult to reject his

conclusions regarding the existence of this as a distinct species. In

20 cases, all that were observed, only lateral spined eggs were found.

Precise and full measurements and descriptions are given for male,

female, egg and miracidium and the lateral spined ova were observed

in the uterus of the female worm.

In a brief article Looss (1909) after analyzing the views and

observations of other investigators regarding the method of infec-

tion in Egypt by the blood fluke comes to the following important

conclusions :

“The theory of the infection taking place by the mouth( along

with food and drink) must be refuted because it is irreconcilable (a)

with certain biological peculiarities of the miracidium, (b) with the

general distribution of the disease among the population of Egypt.

“The theory of infection by miracidium entering the urethra or

the anus is (a) utterly improbable for general parasitological rea-

sons; (b) in contradiction with a number of biological and anatomo-

pathological facts (for example the incapability on the part of the

miracidium to resist the action of acids, even if very diluted; the

part played in the infection by the liver, etc.).

“The theory of infection by the skin is in accordance with all

the facts thus far known (a) of the biology of the parasite, (b) of

the distribution of the disease among the population (native and

foreign, town and rural) of Egypt. It shows (c) how the chief

sufferers—the children in town, the adult males in the country—

live under conditions which, from the epidemiological point of

view, are essentially the same ,and give the miracidia (d) the op-

portunity of passing, within the short time of their life, from man

to water and from water back to man.”

140 HENRY B. WARD

This view which Looss has been brought to adopt after long

study with many experiments and extended observation on the

field has received startling confirmation in recent experiments con-

ducted by Katsurada and Hashegawa (1910). They took young

cats and dogs from non-infected regions and exposed them for an

hour and a half in the ditch water of infected territory under pre-

cautions which rendered infection per os impossible. Then followed

prompt removal from the infected region. About a month later

several thousand blood flukes were present in the bodies of these

experimental animals. Two conclusions seem unavoidable: first,

the young form of Schistosoma japonicum penetrates through the

skin into the human as well as the animal body and attains sexual

maturity within the limits of a month. Second, the miracidium also

develops to a sporocyst and to the adult in the one host and from

a few cysts countless young worms are produced. According to a

brief supplemental note by these authors, Matsuura has demon-

strated eggs in human feces when the infected individual had only

waded in the water of infected regions and Fujinami has been able

to demonstrate skin infection in cattle also.

CESTODA

Kofoid and Watson (1910) have published a most important

preliminary note on the orientation of the adult cestode based on

their studies on Gyrocotyle. The morphology of the nervous sys-

tem leads them to regard the tapeworm scolex as posterior and the

free end of the chain as anterior. “It is noteworthy that this orien-

tation of the cestode brings the growing zone in the so-called neck

of the strobila into a position homologous with that of the ante-

penult segment of the amelid worm, also the zone of growth.”

The importance of this paper consists in the care with which

the view ts worked out by the detailed study of the nervous system

especially. So far as the theory itself is concerned one should call

attention to the fact that these views concerning the orientation of

the cestode body are in agreement with those advanced somewhat

earlier by Cohn (1907). He based his argument which was pre-

sented in the brief form of a preliminary paper on the structure of

the adult Cestodaria, on a comparative study of the morphology and

development of the onchosphere in different groups of tapeworms,

and on a comparison of growth regions in cestodes and annelids.

RECENT PROGRESS IN PARASITOLOGY I4!

Young (1908) has investigated with great precision the develop-

ment of the individual tissues and organs of the common dog tape-

worm, Taenia serrats, from its bladder worm, Cysticercus pisi-

formis. The origin and multiplication of cells in the early stages of

the bladder worm force Young to conclude that the small groups of

chromatin granules (chromidia?) assemble to form nuclei at various

points. Later these nuclei acquire control of protoplasmic areas

which become delimited from surrounding areas as new cells. Nu-

cleoplasm is fundamentally the same as cytoplasm; and in the man-

ner indicated cells may in his opinion originate de novo.

Glaser (1909) described the development of the bladder worm

of Taenia crassipes in great detail. In the main it resembles the

well-known development of Echinococeus. La Rue (1909) has

worked out some important factors in the development of Proteo-

cephalus and gives the first account of fat tissue in a cestode.

The highly interesting primitive cestode, Archigetes has been

studied by Mrazek (1908) who describes a new species and adds

comparative data on the structure of those already known. Plehn

(1908) has added to these primitive forms, frequently grouped to-

gether as Cestodaria, a new genus, Sanguinicola, which is parasitic

in the blood stream of the carp.

Gasse (1910) has investigated the local reaction produced in

the body of the animal by invasion of the bladder worm. He finds

that the host animal envelopes the inwandering Echinococcus or

bladder worm with a variously constructed cyst. Fertile Echinococci

are almost always surrounded by fibrillar connective tissue. Sterile

Echinococci have a three-fold capsule: inside young connective

tissue cells, then round cells, outside fibrillar connective tissue. The

bladderworm cyst consists of the same tissue elements as the Echino-

coccus capsule. Giant cells are present only in cysts of sterile

Fchinococci.

Little has been done on human cestode parasites within the

past two years save the publication of brief notes or preliminary

descriptions of new species.

Stiles (1908) reported from Florida a proliferating cestode

larva found in man which resembles closely the similar human para-

site known from Japan. Its most striking characteristic is the

production by budding of supernumerary heads which may become

independent and wander through the subcutaneous tissue in which

142 HENRY B. WARD

the parasite is found. Nothing is known of the life history, method

of infection, or adult form of the parasite.

Stephens (1908) has described Dibothriocephalus parvus n. sp.

from a Syrian in Tasmania; the parasite is regarded as “possibly

a Levantine product after all and not Australian in origin.” He

also described in the same paper Taecnia Bremneri n. sp. from a

native woman in Northern Nigeria where the parasite was said to be

common.

Léon (1908) has described a new human tapeworm from Rou-

mania under the name of Braunia jassyensis. Another new form of

human cestode, Diplogonoporus Brauni, has also been described by

Léon (1910) who reports two cases which, like the first species,

come from Roumania.

The cestodes of other hosts than man have received consider-

able attention during the last two years. Among the most important

contributions are those of Fuhrmann and Ransom.

Fuhrmann (1908) published a most extensive and valuable

monograph on the cestodes of birds including 500 species from 200

host species. The species of cestodes found in birds of different

zoogeographical areas are often very distinct. To this he has also

added other papers dealing with more recent acquisitions and broad-

ening notably our knowledge of the structure and distribution of

these forms.

Ransom (1909) contributed a monograph on bird cestodes of

this continent and has thus given the first connected account of these

parasites in a region less known in that respect than any other grand

division of the earth’s surface. He has thus supplied an important

gap in our knowledge of the avian parasitic fauna.

Luhe (1909) has published a valuable synopsis of cestodes

found in German fresh water hosts.

NEMATODA

The group of Nematoda has always occupied an isolated posi-

iton in the animal series and during the past few years numerous

attempts have been made by careful microscopic study of the struc-

tural details and of developmental history to demonstrate the prob-

able relationships of the group to other branches and also to the

aberrant groups which have often been associated with it in the

phylum Nemathelminthes. The habits, life history and pathological

RECENT PROGRESS IN PARASITOLOGY 143

significance of the parasitic species have also been investigated with

the results of importance for the comprehension of the relations in

which these species stand to the causation of disease and for the

formulation of rational measures against the spread of these para-

sites.

Rauther (1909) has brought together the results of his pre-

vious studies on the morphology and relationships of the Nematodes

with results which are at variance with previous views. The author

states at the outset that the structural features, developmental

stages, and larval forms of Nematoda furnish no basis for attaching

the group to other types of worms but afford everywhere hints of

a connection with the Arthropoda. He would associate Gordins and

Nectonema with Solenogastridae, though distantly. In the fore gut

of Nematoda he finds many points of similarity to the Echinoderi-

dae, the Gastrotricha, the Tardigrada and Pentastomidae, even to

the Diptera; and in the opinion of the author all these groups may

be traced back phylogenetically to the highly organized Arthropoda.

On the structure of the Nematoda, and of other groups often

classed with them as Nemathelminthes, many studies have been pub-

lished. Of first rank must be mentioned the work of Goldschmidt on

the nervous system of Ascaris.

Goldschmidt (1g09a) found the nervous system of Ascaris so

simple in form, constancy, and symmetry of parts that it could be

described in full and every individual cell indicated by a fixed num-

ber. Thus for the first time it became possible to depict the com-

plete anatomical foundation for a comprehension of all reflex pro-

cesses. In these species he was able to demonstrate continuity of

protoplasm in the system. These species of nematodes grow not by

cell division but by cell enlargement and the adult has in most organs

just as many cells as the completed embryo. It is then clear that

cell lineage has a mathematical relation to adult structure not pre-

viously suspected.

Martini (1909) has finished an important and extensive study

on the comparative histology of the subcuticula and lateral lines of

nematodes. The embryological history of these structures is care-

fully followed out and on the basis of this evidence the author draws

the following conclusions concerning the classification: the group of

Strongylidae is an unnatural one; the terms polymyaria and mero-

144 HENRY B. WARD

myaria are of true systematic value; but coelomyaria and platy-

myaria are not.

Valuable systematic studies on Nematoda have been made by

Railliet et Henry (1909) on the very large and variable family, the

Strongylidae; from which, as formerly held, they exclude Eustron-

gylus, Hystrichis, and Physaloptera as belonging to the Filariidae.

The remaining forms can be grouped into Metastrongylinae and

Ankylostominae with numerous sub-groups. On the basis of ex-

tended studies Jagerskidld (1909) comes independently to the same

conclusion: that the Eustrongylidae constitute a well limited family

which has no near relation to the Strongylidae proper.

Glaue published a brief report on the difference between the

small dog and cat nematodes (Ascaris canis and A. felis) also

known as occasional parasites of man. His work in distinguishing

these forms as separate species has been anticipated in part by

Leiper,* whose original note seems not to have been followed as yet

by the more complete discussion promised.

Considerable attention has been paid to the structure and de-

velopment of the minute blood-inhabiting embryos, known as

Microfilariae, which are frequent in certain regions and are as-

signed a prominent role among human parasites.

Rodenwaldt (1908) has studied the microscopic filarial embryos

in the blood. No mutation could be found among the canine fil-

ariae. The embryo worms are most common in lungs both in men

with mutation and in dogs without it and wander everywhere in

blood vessels, even in terminal capillaries but are lacking in lymph

vessels. They prefer capillaries of larger vessels and hence are

much more frequent in most organs than in the blood stream. It

seems that diwrna is torn away by day in the swift flowing capillary

region of the pulmonary circulation whereas at night it withstands

the current in this circulation. Nocturna is carried on both day and

night in the swift flowing capillary stream of the pulmonary circula-

tion, by day also in that of the systemic circulation, whereas it can

maintain its place by night in the slower moving capillary blood of

the systemic circulation. In the second part are given many topo-

graphic details of external body form and internal structure of

blood filariae, mainly useful for differential diagnosis of species.

*British Medical Journal, June 1, 1907, p. 1296.

RECENT PROGRESS IN PARASITOLOGY 145

Filleborn (1908b) gives diagnostic differences between F.

diurna and nocturna, length of life of worms, anatomical details,

measurements, and infection of mosquitos.

The same author (1908a) also records experimental work on

dogs with F. immitis. He further discusses length of life of micro-

filariae, problems connected with diurnal and nocturnal appearance,

whether the mosquito takes up more microfilariae than proportion-

ate to the blood sucked, the development of the microfilariae in

various mosquitos, at different temperatures, etc., the migration of

the larvae into the proboscis, the stay of the larvae in the mosquito

when no blood is taken, the length of life of larvae ripened in the

mosquito and kept in various media and finally the penetration of

the skin by the larvae.

Noé has demonstrated that for Filaria Grassii the intermediate -

host is Rhipicephalus sanguineus. The nymphs suck up the lymph

of the dog. The infrequent larvae taken with it pass through the

intestinal wall into the lacunom and go through stages of develop-

ment in any organ or tissue. Even the male tick is a parasite car-

rier and important for the species on account of its greater mobil-

ity. The gravid female gets larvae at the start of sucking when

lymph is taken; when blood reaches the stomach the filariae atrophy,

losing their power of movement. Then follows a description of

the larvae that is very complete.

Filleborn (1908) has published a detailed description of

Filaria volvulus, which is common in Kamerun. Several indivi-

duals, male and female, lie in a simple subcutaneous tumor. Free

larvae are found in the capsule of an older tumor, but have not. yet

been found in blood. They are easily told from other known blood

filariae.

Phalen and Nichols (1909) record the facts concerning filaria-

sis in man and other animals in the Philippines and conclude that

since the carrier of the embryo filaria (Culex fatigans) is univer-

sally distributed, the narrow localization of human infection can

only be explained on the mutual exclusion of malarial infection.

Unterberger (1908) shows in one unquestioned case that

Oxyuris vermicularis does bore into the normal mucosa with its

head. In other cases ulceration may have been primary and en-

trance of the worm secondary.

146 HENRY B. WARD

Wenyon (in Balfour, 1908) conducted some experiments with

embryos of the guinea worm, confirming the observations of Leiper

that after the embryo has completed its metamorphosis in the

cyclops, the addition of very dilute hydrochloric acid kills the ento-

mostracan but causes the embryo to desert actively its dead host.

These conditions simulate gastric digestion. After metamorphosis

the long pointed tail is replaced by a short, blunt, bilobed tail.

Important contributions have been made to the geographical

distribution and systematic arrangement of Nematoda through ex-

tensive papers by Jagerskiold, Shipley and von Linstow.

Stephens (1909) has reported a new nematode, Strongylus

Gibsoni, as a human parasite from Chinese in Hong Kong. He

also furnishes final evidence for excluding Filaria immitis from

the list of human parasites, printing a letter from Bowlby, the sup-

posed authority for such occurrence which states that the use of

this name resulted from the error of a reviewer. :

Lithe (1909a) has published a resume of our knowledge con-

cerning the Acanthocephala which from its complete and critical

character is certain to form the basis for future work on this group.

He reviews the history of investigations in this field from the

earliest references to the work of Westrumb in 1821 and then lists

all the species included under the genus Echinorhynchus with

critical citation of complete data for each species; later he cites

those forms originally included under other generic names and

finally discusses the genera of Acanthocephala, both those that are

deserving of present acceptance and those that are to be rejected

as synonyms or on other grounds.

Our scanty knowledge concerning the life history of the

Acanthocephala renders the work of Riquier (1909) especially note-

worthy ; in the pike he produced experimentally the development of

Pomphorhynchus laevis Zoega (Echinorhynchus proteus West.)

from Tinea vulgaris.

INSECTS

The agency of blood-sucking insects in the transmission of

disease has led to extended studies on the determination of species,

and on the structure and the life history of such forms. Briefer

studies and more extended papers on various species and genera

are exceedingly numerous, and monographs on the different groups

of mosquitos, flies and ticks are among the most important con-

RECENT PROGRESS IN PARASITOLOGY 147

tributions of recent years. In a recent volume on blood-sucking

flies, Austen (1909) completes the British Museum series of mono-

graphs on blood-sucking insects which was planned by Lankester.

While the record is confessedly incomplete, and the museum ma-

terial available from the different regions of very unequal value,

nevertheless Uganda, which has particular interest in the problem

of disease transmission by blood sucking insects, is well treated

and the description of binomic features will be of particular value

to investigators in the field.

King (in Balfour, 1908) reports on the flies and other blood

sucking insects in the Sudan.

The Sleeping Sickness Commission of the Royal Society found

as the result of its early investigations that Glossina palpalis on the

uninhabited shores of Victoria Nyanza can retain its infectivity

for a period of at least two years after the native population has

been removed. On the basis of the evidence they give, one is

forced to conclude either that the fly is long lived or that there is

a local reservoir of the disease-producing organism. The view

that the crocodile, on the blood of which Glossina feeds, constitutes

such a reservoir is not supported by the facts thus far discovered.

Experiments in transmission of animal trypanosomes, as well

as Tr. gambiense, were made by Bruce (1910) first on lake-shore

flies, and later on flies bred in the laboratory. In wild flies the

shortest time before one became infected with Tr. gambiense was

18 days, the longest 45 days, and the average 32 days. In labora-

tory bred flies the shortest time was 27 days, the longest 53 and

the average 36 days. It was very difficult to infect such flies at

all. A wild fly may remain infective at least 75 days.

It appears now probable that sleeping sickness may be con-

veyed by some other agent than Glossina palpalis, some few posi-

tive cases being reported from regions, particularly Lake Nyassa,

where this fly has never been found. It is hardly likely that the

species has been overlooked.

Bruce and others have also shown that mechanical transmis-

sion of sleeping sickness by means of Glossina palpalis can take

place if the transference of the flies from an infected to a healthy

animal is instantaneous,—i. e., by interrupted feeding. But this

mechanical transmission does not occur if any time interval comes

148 HENRY B. WARD

between the feedings, and it plays a much smaller part, if any, in

the spread of sleeping sickness than has been heretofore supposed.

The same authors have proved experimentally that cattle may

act as a reservoir of the virus of sleeping sickness, and that healthy

animals may be infected from them by means of Glossina palpalis.

Furthermore, in the fly area cattle do harbor naturally Trypano-

soma gambiense and may keep up indefinitely the infectivity of the

fly, though proof is lacking that in nature such does actually take

place.

Complete records have been published of 50 cases of sleeping

sickness in Europeans. Of these 30 are known to be dead, 11 sur-

vive, and the fate of 9 is uncertain. At least one well known case

seems to have made a positive recovery, and there are grounds for

like hope in 4 others or more. Their recovery appears to be due

to the resisting power of the human organism, at least as much

as to the treatment.

Among diseases transmitted by blood sucking insects, sleep-

ing sickness has received primary attention during the last few

years, but bubonic plague has taken almost equal prominence in

scientific study. Much work has been done under the direction

of the United States Marine Hospital Service on our western coast

with reference to the occurrence of the fleas and their relation to

rats and ground squirrels in which the disease now seems to be

endemic. Thus it has been shown that rat fleas, caught in San

Francisco, will bite man under experimental conditions, and that

squirrel fleas feed readily on human blood. Furthermore, fleas

from rodents adapt themselves quickly to hosts of a different spe-

cies and pass from rat to squirrel and vice versa, even in the pres-

ence of their proper hosts. The plague bacilli have been demon-

strated in the common squirrel flea and in the lice found regularly

on the same host. It is interesting to note that the average number

of fleas from the squirrel is larger than from the rat or from any

other host yet observed. On the transportation of fleas between

distant points by means of ships, rats and mice, one investigation

made at Hamburg covering a period of three months, showed that

6% of the rodents examined carried fleas in considerable numbers,

and that all but 6% of the fleas were of the species Pulex cheopis,

a form primarily responsible for the transfer of the plague bacilli.

This observation furnished positive evidence that living fleas are

RECENT PROGRESS IN PARASITOLOGY 149

imported from Oriental ports into western harbors; and even in

the absence of living rats, or under circumstances when the immi-

gration of rodents from the ship is absolutely prohibited, they may

transfer infection and be distributed to various points on shore

through personal effects or various articles of freight and baggage.*

In an extended critique and summary of present knowledge,

Nuttall (1908a) deals with the importance of various species of

ticks, especially in the transmission of protozoal diseases. Among

the latter, those due to hematozoal parasites (Piroplasma) take

first rank in the severe effects produced among domestic animals.

Relapsing fevers in man and similar mammal and fowl diseases

due to Spirochaeta, are also tick transmitted. The propagation of

Rocky Mountain or Spotted Fever is due to a tick and mention has

already been made in this paper of the demonstration that Filaria

embryos may undergo their development in a tick which also serves

as a means of conveying the infection from one human host to

another.

The final report of Ricketts (1909) contains a summary of

his work on the spotted fever of Montana and its transmission

from which the following items are excerpted as of importance here.

The wood tick is the natural means by which man is infected with

this disease, to which also some of the small native animals are

susceptible. The latter serve as a reservoir for the disease, keeping

it alive from year to year. The disease is hereditary in the tick,

though this fact alone cannot be responsible for the maintenance

of the disease; hereditary transmission occurs probably in less than

50% of the cases under natural conditions. Experiments showed

that the rock squirrel, chipmunk, the wood-chuck, and the mountain

rat are all also adapted to the maintenance of spotted fever in

nature.

Nuttall and his collaborators have published a long series of

papers on various factors in the structure and biology of the para-

sitic ticks. Among them are articles on the presence of an anti-

coagulin in the salivary glands and intestines of Argus perscius, on

the structure of tick spiracles, of Haller’s organ, on the behavior

of spirochaetes in the bed bug (Acanthia lectularia) and on the

structure and biology of Haemophysalis punctata.

*For details and_references to literature consult my review on this topic in the

American Naturalist, July, 1910, p. 439. ;

150 HENRY B. WARD

Neumann has continued the well known series of papers on

the ticks and has begun also a new series treating of the compara-

tive morphology of the Pediculidae.

Rohr (1909) has contributed an extensive memoir on the

Ixodidae of Brazil, including valuable data on their internal micro-

scopic structure and on the biology of the group.

King (in Balfour, 1908) has made an extensive report on the

ticks of Central Africa with data concerning hosts, life history

and relation to disease.

One of the most important studies in microscopic anatomy pub-

lished recently is the paper by Stiles (1910) on the microscopic

structure of the stigmal plates in Dermacentor. He demonstrates

the high systematic value of these plates which afford a simple

means for the differentiation of the otherwise variable, often easily

confused species.

MAMMALS

The most striking investigations of recent years with refer-

ence to the bubonic plague and its relation to the human species

have not been those which dealt with the transmitting insects, but

rather those which concerned the natural and acquired hosts of

the disease among the lower animals. As is well known the

bubonic plague is probably primarily a rodent disease and the

rodent host is believed by some to be a Mongolian marmot, from

which the disease is transmitted through house rats to the human

species. In California the disease has become endemic among the

ground squirrels which some years ago suffered from a wide-

spread epidemic. It is established on the best of evidence that the

disease has been transmitted from the ground squirrel to the human

species, and that plague infected rodents still exist on the western

coast, although infrequent at present. From the ground squirrel

it has spread in exceptional cases to the brush rat and perhaps to

other native species. Transmission to the human race can occur

usually through the medium of the house rat rather than directly

from the native species, although it has been suggested that trans-

fer from the squirrel to man may be brought about through range

cattle which acquire the fleas in and about squirrel villages and

then through contact with man give opportunity for transfer of

the infected insects. It has been suggested also that the booby

owl, which occupies the same burrows with the ground squirrel,

RECENT PROGRESS IN PARASITOLOGY I5I

may readily carry infected fleas over long distances, infect new

squirrel villages and new regions, and thus complicate greatly

the problem of the reduction of the disease. The habits of the

wood rat militate against the probability that it can play any par-

ticular part in infecting the human species.

A most important contribution in the field of medical zoology

which is also unique in having been published by a committee of

business men, is the record of Plague Eradication in San Francisco

(Citizens Health Committee, 1909). Apart from its medical and

social importance it contains chapters on the role of the rat and

of the flea which present in non-technical form the biological data

concerning these species, especially such items as. were determined

in the course of the work at San Francisco and are related to plague

transmission. The report includes full details concerning the means

taken for the extermination of the rat and for rat-proofing buildings.

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RECENT PROGRESS IN PARASITOLOGY 155

MILLeEr, W. W.

1908. Hepatozoon perniciosum (n. g., n. sp), a Haemogregarine Patho-

genic for White Rats. Hygienic Lab. Bull. 46.

MincHIN, E. A.

1908. Investigations on the Development of Trypanosomes in Tsetse

Flies and Other Diptera. Quar. Jour. Mic. Sci., 52:159-260. 6 PI.

MincuHin, E. A. anp THomson, J. D.

1910. The Transmission of Trypanosoma Lewisi by the Rat-flea (Cerato-

phyllus fasciatus). Proc. Roy. Soc., London, B 555, pp. 273-285.

Mincuin, E. A. anp Woopcock, H. M.

1910. Observations on Certain Blood Parasites of Fishes Occurring at

Rovigno. Quar. Jour. Mic. Sci., 55 :113-154; 3 Pl.

Moors, J. E. S. anp BrEINL, A.

1909. Life History of Trypanosoma equiperdum. Proc. Roy. Soc., Lon-

don, B 80:288-298. 1 PI.

Nagler, K.

1909. Entwicklungsgeschtliche Studien ttber Amoeben. Arch. Protistenk.,

15-53. 6 PL.

Mrazek, Al.

1908. Ueber eine neue Art der Gattung Archigetes. C. B. Bakt. u. Par.,

Orig. 46 :719-723.

NIcoLL, Wo.

1909. Studies on the Structure and Classification of the Digenetic Trema-

todes. Quar. Jour. Mic. Sci., 53 :391-487. 2 PI.

NICOLLE, CH.

1909. Le Kala Azar infantile. Ann. Inst. Pasteur, 23:362-401, 441-471.

2 Pk

Noe, G.

1908. I1 ciclo evolutio della Filaria grassii mihi, 1907. Atti Accad. Lin-

cei, (5) 17:282-93.

NuTTaLt, G. H.

1908. The Transmission of Trypanosoma Lewisi by Fleas and Lice.

Parasitology, I :297-301.

1908a. The Ixodidea or Ticks, Spirochaetosis in Man and Animals, Piro-

plasmosis. Jour. Roy. Inst. Pub. Health, London. 51 pp.

OpDHNER, TH.

1909. Was ist Distomum Rathouisi? Arch. Parasitol., 12:467-471.

OrTMANN, W.

1908. Zur Embryonalentwickung des Leberegels (Fasciola hepatica L ).

Zool. Jahrb., Anat., 26:255-292. 3 Pl.

156 HENRY B. WARD

Patton, W. S.

1909. The Parasite of Kala Azar and Allied Organisms. London Lancet,

Jan. 30, 1909. (Reprinted in 7 pp. with 2 Pl.).

PHALEN, J. M. anv Nicuots, H. J.

1909. The Distribution of Filaria in the Philippine Islands. Phil. Jour.

Sci., B 4:127-139. I map.

PLeuN, M.

1908. Ein monozoischer Cestode als Blutparasit (Sanguinicola armata

und inermis Plehn). Zool. Anz., 33 :427-440.

PRATT Eas:

1909. The Cuticula and Subcuticula of Trematodes and Cestodes. Amer.

Nat., 43 :705-720.

PrRowWAZEK, S.

1909. Kritische Bemerkungen zum Trypanosomenproblem. Arch. Schiffs.-

Tropenhyg., 13 :301-8.

Ratwuiet, A. et Henry, A.

1909. Sur la classification des Strongylidae: 1. Metastrongylinae. 2.

Ankylostominae. C. R. Soc. Biol., Paris, 66:85-88, 168-171.

Ransom, B. H.

1909. The Taenioid Cestodes of North American Birds. U. S. Nat.

Museum, Bull. 60. 141 pp. 42 figs.

RavuTHer, M.

1909. Morphologie und Verwandschaftsbeziehungen der Nematoden und

eininger ihnen nahe gestellter Vermalien. Ergeb. Fortschr. Zool., 1

491-596.

Ricketts, H. T.

1909. Investigation of the Cause and Means of Prevention of Rocky

Mountain Spotted Fever, Carried on during 1907 and 1908. Mont.

St. Bd. Health. Fourth Biennial Report, pp. 77-191.

Rigurer, J. K.

1909. Die Larva von Pomphorhynchus laevis Zoega (—Echinorhynchus

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RECENT PROGRESS IN PARASITOLOGY 157

Ronr, C. J.

1909. Estudos sobre Ixodidas do Brazil. Trabalho do Institute Oswaldo

(Cruz 220) pps 5 eek

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1909. The Development of the Parasite of Oriental Sore in Cultures.

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and Hyg., 12:I-IT.

SHIPLEY, R. E.

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Other Vertebrates. Parasitology, 1 :263-2709.

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1909. A New Human Nematode, Strongylus gibsoni, n. sp. Ann Trop.

Med. and Par., 2:315-316. 2 PI.

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Trop. Med. and Par., 2:317-320.

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72 pp. 43 Pi.

Stites, C. W. AND GOLDBERGER, J.

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and of nineteen allied species of Mammalian Trematode Worms of

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1908. Ueber eine neue parasitare Krankheit (Schistosomiasis japonica)

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Gegenden Japans. Arch. Path. Anat., 193 :323-369. 1 PI.

158 HENRY B. WARD

UNTERBERGER, FR.

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1909. The Influence of Civilization on Human Zoo-parasitic Diseases.

Proc. Seventh Int. Zool. Congress, Boston. (Reprint 7 pp.).

1909a. Fasciolopsis Buskii. F. Rathouisi and Related Species in China.

China Med. Jour.; Also Trans. Amer. Mic. Soc., 29 :5-16.

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Zool. Jahrb., Anat.,

RECENT TENDENCIES IN THE PEDAGOGY OF BIOLOGY

By JOHN G. COULTER

It appears fair to say that more attention than before is being

given to the problem of the teaching of biology. It appears fair to

say that problems connected with the uses of biological materials in

education are receiving better recognition than heretofore as re-

search problems of real value to science as well as to education.

Laboratory methods are being carried over into the investigation of

educational problems; and the biologist, for example, who con-

fesses that his research-interest lies in the study of the uses of

plants and animals as means in education rather than as ends in

themselves needs no longer fear that he will lose caste. Con-

structive research upon the educational uses of biological materials

is finding equal recognition with research in other biological fields.

Theses based upon such work may even find equal consideration

for the doctor’s degree with theses based upon what we somewhat

vaguely differentiate as ‘“‘pure science.”

There is a question of distinction here, however. Professor

Ganong in his recent revision of The Teaching Botanist (p. 58)

puts it as follows:

“The temper and temperament required for investigation and for gen-

eral teaching are not simply different, but are even somewhat antagonistic.

* * * * * This applies to the university type of abstract research, the

kind which is on the forefront of advancing knowledge; on the other hand

it does not apply to some other types of investigation, which are closely and

logically connected with the teaching. Of this kind is investigation into the

pressing educational problems of the science, a field as difficult and service-

able as anything which the abstract phases of the science have to offer the

teacher.”

All of which appears to indicate that the day of a science of

education approaches. The teachers themselves are looking more

to results attained by science-methods, and listening less to the per-

suasive voice of authority unsupported by evidence which will en-

dure the criteria of science. The need for thoroughly trained in-

vestigators in education is enlarging; not so much because the re-

sults of their work will be more accurate now than formerly, as

160 JOHN G. COULTER

because they may obtain now a larger hearing than fomerly. The

results of their researches will have real effects upon procedure.

Whereas, even a decade ago, they would have had slight notice, so

dominant in the educational world was mere opinion; so ascendant

was authority based on little more than the prestige which position

and vigorous assertion give. Such dominance and such ascendancy

are passing. (An excellent presentation of this point is found in

Bagley’s ‘““The Scientific Method in Educational Research,’ Nature

Study Review, Sept. 1910.)

The natural divisions of the uses of biological materials in edu-

cation are :—those of the elementary school, of the secondary school,

and of the college or university. Of the problems lying in the first

of these divisions, Professor Ganong writes that they are “as much

psychological as scientific.” (The Teaching Botanist, p. 3.) We

may leave it to the psychologist to resent any invidiousness in this

distinction, and merely accept the thought behind as reason for the

omission of “nature-study” problems from this review, so far, at

least, as the elementary school is concerned. The omission of col-

lege and university problems is warranted, partly because they are

less than those of the high school, partly because opinion about

them is less accessible, partly by the page limits of the review, and

mainly because the reviewer finds himself even less ready to discuss

them. It seems safe to say, however, that post-high-school biology

teaching is in far more satisfactory condition than that done in the

high school.

Only in a negligible minority of high schools do “botany” or

“zoology” appear as required subjects later than the second year.

Hence the problem in this field narrows itself to youths of from

14 to 16 years, and the ‘psychological’ element is obvious. To

the writer a present basic question appears to be whether children

of this age are, or are not, ready to profit by the study of science;

is it desirable or is it foolish to attempt at this age the inculcation

of the scientific spirit? Upon the answer to this psychological

query appears to rest the whole matter of method in the use of

biological materials in the high schools, at least in the years in

which they are now almost exclusively used. Let it be clear, how-

ever, that this answer will furnish a basis for method alone. It

would furnish no basis at all for the exclusion of plant and animal

materials from the program, though it might furnish ample warrant

RECENT TENDENCIES IN THE PEDAGOGY OF BIOLOGY 161

for discontinuing to name such uses either “botany,” or “zoology,”

or even “general biology.”’ For the thing sought all of these may

prove misnomers, if they are not already misnomers for things

urged and attempted in their name.

We find at present urged at least three types of high school

courses in these materials. These may be roughly differentiated as

“economic,” “natural history,” and “pure science.” The limits of

this review make it almost imperative to assume that these terms are

self explanatory.

Inquiries which the writer has made in the Middle West are

supported by numerous published opinions in warranting the asser-

tion that college and university men—the professional biologists—

while showing considerable variation of opinion in minor matters,

hold practically together in the opinion that “pure science” should

dominate in the use of biological materials in the secondary schools.

The school masters, on the other hand, hold with equal unanimity,

for “economic” or “natural history’ dominance. The position of

the scientists is succinctly put in the following sentence: “To be-

gin with economics and to work back to the scientific basis thereof

seems to me justified neither by theory nor experience; while be-

ginning with scientific study and working thence to economics

seems to me in accord with both.” (Ganong, The Teaching Botan-

ist, p. 46.)

A similarly definite statement of the position of the school men

is not at hand, but it appears to the writer to be somewhat as fol-

lows: Educational values are evident only as conduct is affected.

We are thoroughly sceptical of the conduct-effects of pure science

teaching in the first and second years of the high schools, at least

so far as its effects may be judged from the kind of teaching we

are now able to find and employ. Its values do not appeal to

adolescents of that period. The “human interests’ should domin-

ate in the use of biological materials in these years, whereby we may

directly affect conduct through precept and example, dealing with

familiar and economically important plants and animals. Content

deserves consideration over method at this stage. We grant the

theoretical values of pure science methods, but we have not had

adequate evidence of their benefits in this place in education. We

know that we can teach a boy to make a better garden and that he

will largely do the things we teach him how to do. They appeal to

P90 16h

162 JOHN G. COULTER

him. We do not know that we can make him a better thinker, or

a better citizen, through pure science at this stage, and we prefer

the certain, tangible values to these uncertain mental ones we have

scant proof for. We can teach him how to do at this stage much

better than we can teach him how to think.

It appears to be true that this divergence of opinion is more

apparent than real, at least to the extent that what many school

men are seeking as “botany” and “zoology” are not such in any

strict manner of speaking. If they find fault that the teachers they

obtain from universities, as teachers of botany or zoology, are not

trained in agriculture, horticulture, or forestry, bits of which they

ask to be included as botany, they should not find that fault with

the university which makes no claim to include these divisions in

in its science departments, but rather with conditions which provide

as yet no adequate facilities for the training of such teachers as they

desire.

Dewey (Science as Subject Matter and as Method; Science,

Jan. 28, 1910) makes a case against the educational value in science

as it has been taught, but remains true to the university ideal as

to how it should be taught. He does not pretend, however, to limit

himself to the under years of the high school as to pupils, nor to

the actual limitations as to teachers. His conclusions, however,

point to radical alteration as to content even of the pure science

course. To achieve the training in method he advocates, limitations

of time would require extensive elimination of matter at present

required by universities for entrance.

The infinitely extensive character of natural facts and the universal

character of the laws formulated about them is sometimes claimed to give

science an advantage over literature. But viewed from the standpoint of

education, this presumed superiority turns out a defect; that is to say, so

long as we confine ourselves to the point of view of subject-matter. Just

because the facts of nature are multitudinous, inexhaustible, they begin no-

where and end nowhere in particular, and hence are not, just as facts, the

best material for the education of those whose lives are centered in quite

local situations and whose careers are irretrievably partial and specific. If

we turn from multiplicity of detail to general laws, we find indeed that the

laws of science are universal, but we also find that for educational purposes

their universality means abstractness and remoteness. The conditions, the

interests, the ends of conduct are irredeemably concrete and specific. We do

not live in a medium of universal principles, but by means of adaptations,

through concessions and compromises, struggling as best we may to enlarge

RECENT TENDENCIES IN THE PEDAGOGY OF BIOLOGY 163

the range of a concrete here-and-now. So far as acquaintance is concerned,

it is the individualized and the humanly limited that helps, not the bare uni-

versal and the inexhaustibly multifarious. * * * * *

Something of the current flippancy of belief and quasi-scepticism must

also be charged to the state of science teaching. The man of even ordinary

culture is aware of the rapid changes of subject-matter, and taught so that

he believes subject-matter, not method, constitutes science, he remarks to

himself that if this is science, then science is in constant change, and there is

no certainty anywhere. If the emphasis had been put upon method of attack

and mastery, from this change he would have learned the lesson of curiosity,

flexibility and patient search; as it is, the result too often is a blasé satiety.

I do not mean that our schools should be expected to send forth their

students equipped as judges of truth and falsity in specialized scientific mat-

ters. But that the great majority of those who leave school should have

some idea of the kind of evidence required to substantiate given types of

belief does not seem unreasonable. Nor is it absurd to expect that they

should go forth with a lively interest in the ways in which knowledge is im-

proved, and a marked distaste for all conclusions reached in disharmony

with the methods of scientific inquiry. It would be absurd, for example, to

expect any large number to master the technical methods of determining

distance, direction and position in the arctic regions; it would perhaps be

possible to develop a state of mind with American people in general in which

the supposedly keen American sense of humor would react, when it is

proposed to settle the question of reaching the pole by aldermanic resolu-

tions and straw votes in railway trains or even newspaper editorials. * * *

Mankind so far has been ruled by things and by words, not by thought;

for, till the last few moments of history, humanity has not been in posses-

sion of the conditions of secure and effective thinking. Without ignoring in

the least the consolation that has come to men from their literary education,

I would even go so far as to say that only the gradual replacing of a literary

by a scientific education can assure to man the progressive amelioration of

his lot. Unless we master things, we shall continue to be mastered by them;

the magic that words cast upon things may indeed disguise our subjection

or render us less dissatisfied with it, but after all science, not words, casts

the only compelling spell upon things. * * * * *

The modern warship seems symbolic of the present position of science

in life and education. The warship could not exist were it not for science:

mathematics, mechanics, chemistry, electricity supply the technique of its

construction and management. But the aims, the ideals in whose service

this marvelous technique is displayed are survivals of a pre-scientific age,

that is, of barbarism. Science has as yet had next to nothing to do with

forming the social and moral ideals for the sake of which she is used. Even

where science has received its most attentive recognition, it has remained a

servant of ends imposed from alien traditions. If ever we are to be gov-

erned by intelligence, not by things and by words, science must have some-

thing to say about what we do, and not merely about how we may do it most

164 JOHN G. COULTER

easily and economically. And if this consummation is achieved, the trans-

formation must occur through education, by bringing home to men’s habitual

inclination and attitude the significance of genuine knowledge and the full

import of the conditions requisite for its attainment. Actively to participate

in the making of knowledge is the highest prerogative of man and the only

warrant of his freedom. When our schools truly become laboratories of

knowledge-making, not mills fitted out with information-hoppers, there will

no longer be need to discuss the place of science in education.

Ganong (Some Reflections upon Botanical Education in Ameri-

ca, Science, March 7, 1910) makes a similar point which, to ac-

complish, would also require vigorous pruning of the course under

present high school conditions.

Another phase of our treason to the genius of science is found in the

belief and practise of some teachers that broad generalizations are the true

aim of elementary teaching. I know a recent elementary textbook in which

the author laments that “some teachers do not yet understand the import-

ance of imparting to beginners a general rather than a special view point.”

And I could cite many passages to show a belief of this and some other

teachers that subject matter, accuracy in details, and other fundamental veri-

ties of science, are not important in comparison with “view-points” and

“outlooks on life’ and that sort of thing. In my opinion there can be no

greater educational error. There is no training which American youth needs

more than that in a power to acquire knowledge accurately and to work de-

tails well. Disregard for particulars and a tendency to easy generalities are

fundamental faults in American character, and need no cultivation, but, in-

stead, a rigorous correction. ;

In whichever direction we look radical change appears to be

more or less imminent. It is frequently suggested that such change

may lead to distinct differentiation between city and country condi-

tions, the thought being that city schools must, from the very con-

ditions of their environment, hold to “pure science” ideals, while

the country, with wealth of materials at hand, may more reasonably

make environment the basis, at least for materials.

Another argument for the “natural history’’ method is based

upon the fact that chemistry and physics are firmly entrenched in

the upper high school years, and that much of the experimental

work in biology, urged by “pure science” and “scientific habit of

thought” advocates, is pointless without a knowledge of these. But

the strongest and the simplest argument brought forward by the

school men is, perhaps, that teachers qualified to train boys and

girls in the scientific spirit, admittedly a teaching calling for high

RECENT TENDENCIES IN THE PEDAGOGY OF BIOLOGY 165

qualifications, are not usually obtainable with the present status of

salaries. Society, if it wants better thinkers turned out by its

schools, must pay a higher school tax. Effort has been made,

notably in New York State, to solve the pedagogical problem by a

first year course in General Biology; while important schools, as

the high school at Springfield, Mass., and the University High

School in Chicago, have early courses in General Science. In the

former case the attempt appears to involve no important reduction

in the amount of material used, but rather to increase correlation

and synthesis, with a large emphasis on “human interest.” In the

latter cases there is frank concession to the “natural history” idea,

the course being required of all students. Later electives in botany

and zoology are also offered. To such a plan there are serious

physical obstacles so far as the average high school is concerned.

The following references may be added to those already given,

as having special value in indicating modern tendencies in respect to

the teaching of Biology:

CaLpweLL, O. W.—‘“The Principles that Should Determine the Courses in

Biology in Secondary Schools.” School Sci. and Math.,

Mch. 1909. ‘

GaLLtoway, T. W.—“An Appreciation of the Pedagogical Possibilities of the

Biological Laboratory.” School Sci. and Math., Feb. 1908.

GaLLtoway, T. W.—“Elementary Zoology.” 1910, P. Blakiston’s Son & Co.,

Philadelphia. In the writer’s opinion this book repre-

sents the most distinct advance which has been made in

the way of a text book toward the ideals of Dewey and

toward some of those of Ganong.

GRUENBERG, Beny. C.—“Some By-Products of Biology Teaching.” School

Sci. and Math., Apr. 1908.

THE SPENCER-TOLLES FUND

To the Members:

Although the Society, by reason of circumstances unnecessary

to explain, has rested for a few years, this fund continued its

activity without interruption. At the last meeting in Boston in

1907, it amounted to $2,530.57; its increase since is $620.58.

Its origin, purpose, growth, management and service (the lat-

ter yet in its infancy) are well known to the older members. It may

interest the newer members, as well as those expected to join the

society, to be informed upon these subjects.

At the Rochester, N. Y., meeting in 1884, it was determined

to commemorate the labors of Chas. A. Spencer and Robert B.

Tolles, two American opticians, who should be remembered and

honored as long as the microscope shall endure. *This was to be

accomplished by establishing the ‘“Spencer-Tolles Fund” from

which the income was designed to support a fellowship.

It happened that Rev. W. H. Dallinger, then President of the

Royal Microscopical Society, on a visit to this country, attended

this meeting. Through him that society made a contribution of

five pound sterling. This money, received on December 17, 1884,

was the beginning of the fund.

But mere hopes and expectations are unproductive. In ten

years the fund had reached only about $350.00. While waiting for

better results the society, by the efforts of some members, antici-

pated the expected benefits from the fund by offering prizes for

original microscopical work. In the year 1893 the sum of $130.00

was so distributed. For the following year $125.00 was promised

for the same purpose, but the promise was broken, and, thereby, the

giving of prizes ended. However, the assistance rendered by these.

small sums to investigators, and the results obtained served as a

*“Memoir of Chas. A Spencer” by Hamilton L. Smith, LL.D., F.R.M.S., Transactions,

1882, Vol. IV, pp. 49-74. Portrait.

“Robert B. Tolles and the Angular Apperture Question,’ Presidential Address of

Jacob D. Cox, LL.D., F.R.M.S., Transactions, 1884, Vol. VI, pp. 5-39. Portrait.

“Memoir of Robert B. Tolles” by Geo. E. Blackham, M.D., F.R.M.S. Ibid. pp. 41-46.

“The Debt of American Microscopy to Spencer and Tolles” by William C. Krauss,

B.S., M.D., Transactions, 1901, Vol. XXIII, pp. 19-29. Four portraits.

168 MAGNUS PFLAUM

stimulus for renewed efforts in behalf of the fund. Up to the year

1895, the interest received was under 6%. Then the investment

was changed, and with considerably higher rate of interest, it

reached in 1900 the sum of $756.00. This was an encouragement.

In that year the office of Custodian was separated from that of

Treasurer; thereby better attention could be paid to the growth of

the fund; all proceeds from sales of Transactions were permanent-

ly assigned to it; and an appeal made to the membership for further

contributions. This action brought the fund during the following

year up to $1,200.00. In 1902 life memberships were created. Since

that year gratifying progress has been made. The following is the

condition to date:

Total Conttib ution -reectved’s sos .co es cing eee ee $ 604.33

Total. Proceed sales of Dransacttons. o25. 2500050 3-2-8 625.73

Motaln(S) Lites MenthershipSesmecseiceestios eee eee 250.00

Total Interest and Dividend received.................-- 1,720.58

$3,290.64

esse LURE RS LANES (eee eee cote Loree $100.00

Less dues on Life Membership................ 22.00

Less 7 years’ expense of Custodian’s office*.... 17.49 139.49

Net fundnvested: CSept.wlOL0) e-aro ens oe wcieaes eee $3,151.15

*This expense relates to the care and management of the property of the Society.

The fund is managed wholly free of any charge or expense.

The income now is a little over 6%, compounding semi-an-

nually,

Such a fund, almost sacred in its character, should be invested

in the safest possible manner, but yielding the highest attainable

income commensurate with safety. This was found possible by

taking shares in a reliable well managed Building and Loan Asso-

ciation. A committee, appointed in 1895 for this purpose, selected

The Keystone Building & Loan Association of Pittsburg (office,

Frick Annex Building, Pittsburg, Pa.). This association is now in

its nineteenth year, has over $4,000,000.00 of stock in force, with

$1,722,000.00 assets, and is perhaps one of the best managed insti-

tutions of its kind in the State of Pennsylvania, in which all such

corporations are under supervision of the State Banking Commis-

sioner in the same manner as State banks.

What service has this fund so far rendered?

When in the year 1900 the so long coveted sum of $1,000.00

THE SPENCER-TOLLES FUND 169

was actually in sight the Society created a special committee for

the management of the fund and making grants from its income.

It was deemed wise to spend but a part of the income so as not to

hinder the further growth of the principal. The grants to investi-

gators was therefore, for the time being, limited to $50.00 per year.

So far but $100.00 had been awarded.* If so much was ac-

complished with such a small sum we may easily conceive what

greater means might produce. In a short time the annual income

will be $200.00. This sum will result in some good, but seems a

puny amount considering the important service it is to render, and

that it expresses the gratitude, and represents a monument, to the

men for whom the fund is named, by a national body of the dignity

of this Society.

A greater degree of usefulness of the fund rests with the mem-

bers of the Society and the friends, at large, of microscopical re-

search. Such labor is seldom undertaken for the sake of com-

pensation, but on the contrary is usually attended with expense

which some investigators are unable, and in justice to the cause

and in furtherance of science should not be permitted, to bear.

The purpose of the fund is neither charity, nor generosity, nor

compensation—it is either broad-spirited encouragement of scien-

tific research, with all its possibilities of benefit to mankind, or is

not worth a moment’s care or attention.

The fund at present should be regarded as a healthy, promis-

ing infant, well worthy of generous nourishment to bring it to

beneficient maturity.

Will you assist, dear Reader ?

MAGNUS PFLAUM, Custodian,

Meadville, Pa.

*Grant No. 1. “The Early Morphogenesis and Histogenesis of the Liver in Sus

Scrofa Domesticus,” by David C. Hilton; four plates, Transactions 1902, Vol. 24, pp.

55-58. $50.

Grant No. 2. “The Relation of Leaf Structure to Physical Factors,” by Edith

Schwartz Clements; nine plates, Transactions 1904, Vol. 26, pp. 19-102. $25.

Grant No. 3. This was made “to aid Prof. Fred E. Clements and Mrs. Edith S.

Clements in completing an investigation on the modification of leaves.’”’ Ibid. Minutes,

p. 287. $25.

NOTES, REVIEWS, ETC.

A METHOD FOR MAKING ABSOLUTE ALCOHOL

The necessity of absolute alcohol to the histologist makes it one

of the important considerations in a small laboratory. Since it is

expensive to purchase it as such, and since an incorporated insti-

tution of learning is able to obtain commercial alcohol without the

payment of the large internal revenue tax, most of us have manu-

factured more or less absolute alcohol for use in the laboratory.

The use of unslaked lime to remove water is satisfactory as far

as the resulting absolute alcohol is concerned, but is a “messy” pro-

cess which occupies a great deal of room while going on and involves

distilling apparatus not always to be found in a small place. It

should also be done by an experienced person, and so the instructor

himself must take a couple of days from his other work to go

through the process.

The result in our laboratory was that we bought absolute alco-

hol very often rather than prepare it.

At the suggestion of the head of the chemical department, cal-

cium carbide was used to dehydrate the commercial alcohol and the

process is so simple that I give it in detail.

The calcium carbide used is the finely granular sort prepared

for portable lamps.

Into a round-bottomed glass flask holding 1500cc., put an ex-

cess of carbide—300 to 350 grams—and 1000cc. commercial alco-

hol (94 per cent). This, with a good sized condenser set vertically

in the stopper, should be placed in a chemical hood or by an open

window and allowed to react. After the action slows down, in an

hour or thereabouts, heat is gently applied, preferably with a water-

bath, and the boiling continued for two hours longer. Allow to

cool sufficiently to handle, remove the condenser, and connect it to

the flask in the regular distilling position by a Hopkins (Kjeldahl)

distillation bulb. The other end of the condenser can be connected

to a receiving vessel by an adapter and this vessel should have a

CaCl drying tube in its cork. See that all the joints are tight and

distil over rapidly. If the distillation is stopped just before the

flask is boiled dry the flask will not break.

172 NOTES, REVIEWS, ETC.

The excess of carbide, when water is added after cooling, will

loosen the solid mass in the bottom of the flask so that the flask can

be easily cleaned.

8oocc. out of a liter has been recovered as absolute alcohol.

This will mix with the usual xylol without milkiness. It has a

rather offensive odor of acetylene, but is far cheaper than the abso-

lute alcohol from supply houses and quicker than making absolute

from lime.

S. R. WitiiaMs, Oxford, Ohio.

A CONVENIENT REAGENT CASE

A sectional bookcase has been found most useful for keeping

the reagents and appliances used in microscopy free from dust.

The case stands at the side of my work table in a corner. The

third (13 in.) section from the floor, at the level of the table, is

the one which holds the dehydration series and staining jars. In

the sections above and below this are slide boxes, metric apparatus,

drawing materials and a dissecting microscope.

The short distance needed by the doors for swinging allows

me to sit at the work table and reach into most of the sections.

There is also less danger of upsetting jars when they are not

on the table in front of you.

The reagent shelf of the bookcase should have all varnish re-

moved as alcohol running down the sides of the bottles softens the

varnish and causes the bottles to stick. S. R. WILLIAMS.

CULTURES OF SAPROLEGNIA

A very satisfactory method of cultivating and handling the

various species of Saprolegnia, Achlya, and other of the water-

molds, particularly suitable for allowing individual study in large

classes, is as follows:

I. Secure bottom-ooze from a number of ponds or lakes, etc.,

if no infested materials are at hand. Place in these cultures a few

flies, that have been immersed in alcohol for disinfection and then

2. When any of these begin to show signs of the saprophytes,

as a whitish halo, remove the flies to watch glasses half full of water

for observation, one fly to each vessel.

AMERICAN MICROSCOPICAL SOCIETY 173

3. As zoospores are developed, place recently killed mosquitos,

as many as you want preparations, in a watch glass with the in-

fected fly, until they become infected. This requires only a few

moments.

4. Place each mosquito in a hanging-drop culture, or other

slide culture, in a moist chamber, for individual use.

There is just about enough nutrition in the mosquito to bring

the Saprolegnia through the life-cycle, and not enough to give

trouble with bacteria or infusoria. Since the slides can be trans-

ferred to the microscope at any time, without disturbing the culture,

the life history of the fungus can be easily followed by even an

elementary class. It offers a good laboratory exercise for noting

the effects on life and development of changing conditions of tem-

perature and the like.

CULTIVATION OF FRESH-WATER ALGA®

Professor J. A. Nieuwland of Notre Dame University, has done

a valuable service for teachers, and other workers with fresh-water

alge, by bringing together in the Midland Naturalist a series of

suggestions relative to the conditions of successful culture and man-

ipulation of these plants. Among them are the following:

1. Best use small aquaria (one to two gallons) for most alge.

Larger aquaria encourage the growth of Cladophora, and entomos-

traca, worms, insects, etc., which are not helpful to the small alge.

2. Do not put much material in the jars. Often a very small

amount (1 cu. in. to the gallon of water) is best.

3. Remove crustacea and insects. Utricularia placed in the

jars will aid in this removal of the smaller crustacea.

4. In transplanting, retain as nearly as possible the conditions

under which the plants were growing in nature. Use, so far as

possible, the very water in which they are found growing. Never

allow complete change of water. If necessary to add water, use

only a small percent of the total; and if necessary to take it from

tap, allow it to run 5 or 10 minutes before letting it pass into the

jar. If water is too hard, Chara will help to correct it.

5. Cover the bottom of the jars with an inch of well-washed

sand. This, as well as the vessels, should be thoroly disinfected

with formaldehyde if they have had Oscillatoria growing upon

174 NOTES, REVIEWS, ETC.

them. This precaution should always be taken if the higher alge

are wanted.

6. Even if the vegetative portions should disappear, it is bet-

ter not to throw away the material, unless it is foul with the small,

dark blue Oscillatoria. Often after a period of rest, many valuable

types will renew themselves; and, being better adjusted than at the

beginning, may make remarkable success as cultures, and pass

satisfactorily into the reproductive stages. Several types are re-

ported by Professor Nieuwland as appearing year after year in the

same jar.

7. In collecting. the best materials are usually to be had from

small streams or ponds, in which the water almost or entirely dries

up in summer or fall. It is good practise, even in the dry period,

and particularly in winter when the vegetative stages of alge may

not be apparent, to collect some of the sticks, mud or soil, etc., from

places in which desirable plants have been seen during the appro-

priate season. If this material is placed in laboratory jars, excel-

lent laboratory cultures will often be developed in a few weeks.

This indoor “forcing” of winter collections of alge is a most de-

cided aid to the teacher. It furnishes also a promising field of re-

search for students in our laboratories.

8. On the other hand, by putting late-fruiting species into jars,

in the fall, and placing them in diffuse light in a cool place, they

may be kept for weeks or even months with little change.

SUCCESSION OF MICRO-ORGANISMS IN FRESH WATER

Considerable interest has been shown recently by students, in

the normal succession of micro-organisms in water, both in nature

and in the somewhat artificial conditions of the laboratory. This

is one of the phases of plankton work likely to be especially useful

to teachers and to students. An especially well conceived and ex-

ecuted study of this sort has recently been reported and published

by F. E. Fritch and F. Rich in the Proceedings of the Bristol

Naturalists’s Society for 1909. It extends over a period of five

years and is from an inland body of water near Bristol.

The observers found that the dominant forms of algz in the

pond were Cladophora, Spirogyra, and numerous diatoms, both free-

living and epiphytic. The chief subsidiary forms were Edogonium,

Mougeotia, and various Cyanophycee. Among these forms the fol-

AMERICAN MICROSCOPICAL SOCIETY 175

lowing periodic phases were noted, as forming an annual cycle:

(1) A winter phase with free diatoms in abundance; (2) a spring

phase with Spirogyra dominant; (3) a summer phase with Clado-

phora dominant and epiphytes abundant; and (4) an autumn phase

in which many of the earlier forms become active again after a

latent period. These include Spirogyra, Edogonium, or other forms.

The authors classify and discuss the factors that operate to produce

these changes as seasonal, irregular, and correlated. This kind of

continued work with small bodies of water is needed in this coun-

try; and our own members can make valuable contributions to the

ecology of our micro-organisms by such investigations in the neigh-

borhood of their homes.

DISTRIBUTION OF ROTIFERS

In the Journal of the Queckett Microscopical Club, C. F.

Rousselet offers a discussion of the distribution of Rotifers. He

calls attention to the fact that most species have quite a cosmopolitan

range; and that no continent or climatic zone can really claim a pe-

culiar rotatorian fauna. Even the rarer species are reported from

widely separated regions of the earth. These facts are of course

to be coupled with the ease with which Rotifers are transported.

Some of them, as is known, are even capable of being dried out

and of resuming life when conditions become favorable. Even in

species in which this is not true there are resting eggs that resist

both cold and drouth. The fact that the habitat of many Rotifers

is such that they or their eggs are liable to be dried up once or

oftener each year, and thus to be committed to the winds, is likewise

an important factor.

RELATION OF VITALITY OF PARAMECIUM TO CONSTANCY OF

SURROUNDINGS

Mr. L. L. Woodruff states, in the Biological Bulletin, that Par-

amecia kept for generations in a reasonably constant culture me-

dium undergo cyclical changes in protoplasmic vitality ; and finally

die from internal causes. If, however, the culture medium is kept

changing in such a way as to disturb this cycle, the lowering of

vitality may be prevented and the protoplasmic life may be contin-

ued without any apparent decrease of vitality. Possibly the cycle

leading to senility may even be eliminated altogether in this way.

176 NOTES, REVIEWS, ETC.

He has cultivated Paramecium thru more than 1230 generations,

which occurred at an average rate of more than 3 divisions in two

days. These observations make it seem probable that in nature

Paramecium does not undergo this lowering of vitality, because of

the stimulus of changing external conditions.

NOTES ON THE TECHNIC OF TUBERCLE BACILLI

1. Gassi:

Make smear preparations ;

Stain in warm eosin solution for one or two minutes. (To 5c.c.

of I per cent eosin solution add a crystal of sublimate the

size of a lentil.

Wash in water.

Treat with a mixture (0.5 vol. NaHo, 1 vol. potassium iodide,

and 100 vols. of 50 per cent alcohol) until preparation as-

sumes a pale green color.

Rinse carefully with alcohol, and wash with water.

Stain for two or three seconds in a solution:—methylen blue

1 vol.; abs. alcohol 10 vols.; 0.5¢.c. hydrochloric acid and

goc.c. distilled water.

Wash thoroly and mount.

The tubercle bacilli are red, the rest blue.

2. Bernhardt’s method of examining sputum:

Place 5c.c. of sputum and 20c.c. of a 20% solution of com-

mercial antiformin in a stoppered bottle.

When homogeneous, pour in ligroin until a layer 3-5 mm.

thick is formed.

Shake until well mixed, and allow to stand at temperature of

room for % hour.

Take out loop-fulls of the layer immediately beneath the lig-

roin. Fix, stain, and store the films in the usual way.

3. Hammerl’s method of examining sputum:

Five parts of a solution, consisting of 99% ammonia and 1%

caustic potash, is mixed with 1 part of sputum and vigor-

ously shaken until homogeneous.

To 15c.c. of this mixture add 5c.c. acetone.

Centrifuge for half an hour.

Make films from the deposit and stain in the usual way.

AMERICAN MICROSCOPICAL SOCIETY WIG

4. Coppin’s method:

Make thin smear on glass. Fix by heating.

Stain by covering with Ziehl’s carbol-fuchsin, diluted with %

its volume of dist. water.

Heat specimen until it steams well, and put aside for 2-5

minutes.

Wash excess of carbol-fuchsin with water.

Place in bath of 10% sulphuric acid until smear fades to an

almost invisible gray or pink.

Wash well until it appears merely cloudy when held to light.

Cover smear with 1% solution (aq.) of picric acid for 10-50

seconds. Wash and mount as usual.

This method shows the tubercle bacilli as rose-red beaded rods

upon a pale yellow field.

BACTERIAL INFECTION BY ENDO-PARASITES

A. E. Shipley, in the Proceedings of the Royal Society, Vic-

toria, cites a case of the bacterial infection of the swim-bladder of

a trout through the migration of a nematode worm from the di-

gestive tract to that point. This, taken with other known similar

cases, leads him to suspect that in an analogous way human entozoa,

passing from the digestive tract with its numerous microbes, may

be the means of infecting distant organs in man, just as really as

in the case of the ecto-parasites,—even tho less frequently.

Mehlhose has recently cited numerous cases in which bacteria

have been found in the bladder-forms of tapeworms.

LONGEVITY OF TRICHINA

In an examination, made in Posen, of nearly 100 bodies of

persons more than 60 years of age, it was found that almost 20%

showed cysted Trichina. From facts brought out in connection with

the examination, there is strong ground for concluding that Tri-

china may live more than 40 years without losing its power to de-

velop.

SYMBIOSIS AND PARASITISM

A most interesting quadruple mixture of symbiosis and para-

sitism is reported by a French observer, in which three species of

micro-organisms figure. A ciliate infusorian—Trichodina paradoxa

178 NOTES, REVIEWS, ETC.

—lives in the intestine of a mollusk, Cyclostoma elegans. The sur-

face of the infusorian is covered by an ectoparasitic species of

Spirillum. In the pharynx of the infusorian is a cluster of bacteria,

believed to be living symbiotically with the Trichodina. This asso-

ciation is quite constant.

The observations of Bab, Sauvage, and McIntosh seem to show

that it is certainly possible for Spirochaeta pallida to pass from a

syphilitic woman to her children directly thru the ova. McIntosh

recently cites cases in which the ovaries and other tissues—even the

ova—of a congenitally syphilized child showed great numbers of

the organisms.

Schereschewsky has been able to cultivate Spirochaeta pallida

on horse-serum. These cultivated specimens could net be distin-

guished from the usual form supposed to be the cause of syphilis,

except in that they do not produce the disease in animals. This

suggests that they have*either undergone an attenuation in culture,

or that syphilis is not really caused by S. pallida.

Mano Truffi claims to have produced complete immunity of

rabbits to syphilis by cutaneous infection.

The list of lactic acid-producing bacilli is reinforced by a new

Streptothrix isolated by G. C. Chatterje, from ‘‘Dadhi’—a sour-

milk preparation of India. It coagulates the casein, produces much

lactic acid, and rapidly destroys pathogenic bacteria introduced into

milk cultures with it.

K. Kominami (Tokyo, 1909) reports finding the sexual phases

of Mucor racemosus.

Members of this Society who are interested in the protozoa or

in Mitosis cannot fail to get great pleasure and profit from an ex-

amination of the remarkably perfect slides for sale by Mr. V. S.

Powers, Station A, Lincoln, Nebraska: The Secretary is frank to

confess that he has seen nothing to compare with them in domestic

or imported slides, and recommends that members who work with

these subjects individually or with classes send for the list of offer-

ings.

AMERICAN MICROSCOPICAL SOCIETY 179

THE USE OF THE MICROSCOPE IN ELEMENTARY BIOLOGY

In recent years there has been considerable criticism of courses

in Biology, arranged for high school students, which commence

with the cell and the simpler organisms and pass upward along the

ascending scale of animals or plants.

The nub of this criticism has included two main points: (1)

the time honored doctrine that teaching should pass from the known

to the unknown, rather than the reverse; and (2) the microscope

is itself such a difficult and complex instrument, and the interpre-

tation of its story so uncertain and puzzling to youth, that it ought

to come, if at all, much later in the course.

There is of course something in each of these contentions, and

as scientists and citizens, whether we are teachers or not, we are

all interested in the question. Many good teachers have wholly

given up the strong advantage that comes from following the evolu-

tionary order, and begin with the more complex but better known

organisms ; and some cut out the use of the microscope almost alto-

gether.

The writer is glad, however, to be able to offer here, out of his

own experience, at least one suggestion, which may serve to mod-

erate this attack on the microscope as unpedagogical and undesir-

able for use as a high school implement.

In a series of classes of high school age, a number of records

have been made, unknown to the pupils, of tests as to the attitude

of the pupils toward laboratory work that involves the use of the

microscope in comparison with that which does not. It is unneces-

sary here to give a detailed statement of the points on which the

record was made. Various things that indicate to the teacher an

interest on the part of the pupil were noted :—degree of concentra-

tion; the sustaining of effort completely to the end of the period;

facial and bodily attitudes indicative of attention; the voluntary ex-

pressions of the pupils, etc.

The following conclusions are based on these records:

1. The classes show every evidence of being as much interested

in the study of the microscope itself, as an instrument, through the

whole of a two-hour period given to its examination and to the

discussion of its parts and their uses, as in any thing else in the

whole course in Biology. The microscope, as a man-made instru-

180 NOTES, REVIEWS, ETC.

ment of precision, is well worth the student’s time; there is no place

more appropriate for its study than in a course in Biology, to which

it has made its most notable contributions; and the student is inter-

ested in it. These facts make a good pedagogical combination.

2. All the special technic which the high school student must

have at the outset can be mastered in one two-hour exercise; and

there need not be a single dull moment in that time. The utmost

skill that the secondary pupil will need, in order to have the micro-

scope a very valuable aid thru all his course, can be picked up day

by day incidental to the actual use of it.

3. In my own experience, 95% of these student show by my

tests, and avow for themselves, a more intense interest in the aspects

of biology calling for the use of the microscope than in the non-

microscopic work of the course. The microscopic work compares

favorably with the very best field-work that I have ever been able

to organize, in its attraction for even the elementary students.

It is not the desire of the writer to conclude from these obser-

vations that the microscopic work should bulk large in a beginning

course; but rather to insist that the greater interest of it does at

least mitigate the unpedagogic leap into the unknown territory of

the cell and unicellular organisms. If there is any thing in the

doctrine of interest in education, this increased interest may even

justify the teacher in beginning thus low down, merely in order to

get the inspiration which this new point of view may yield. This

may enable the pupil to approach the organisms, which he thinks

he knows, in a more open spirit; and this is a result quite worth

seeking.

Indeed, we have found in our laboratory that the privilege

of applying the microscope to some part of the larger animals, eith-

er in extemporized preparations or in permanent mounts, is often of

great pedagogical value in holding the pupil’s interest to the gross

studies. The microscopic “demonstrations” often used for general

purposes are much more meaningful to students who have had a

first-hand experience with the instrument in their own work.

For these reasons the writer feels that there is need of a re-

action in favor of a wiser, and probably a more extended, use of this

most splendid instrument in even elementary courses. Most of our

pupils do not get beyond the high school; and it is not fair that the

recent vogue of the utilitarian and out-door phases of biology

AMERICAN MICROSCOPICAL SOCIETY 181

should rob the coming generation of a first-hand chance to appre-

ciate a little of the actual world of the minute, and of the marvel-

ous instrument which makes its study possible.

Mr. Edward Pennock of Philadelphia is featuring the newly

designed Pietzsch Microtomes for general laboratory use. It is

claimed that these instruments are of very perfect workmanship

and do exceptionally accurate work.

As is well recognized, the glass covers used so much by

microscopists vary greatly in thickness even tho sold under stand-

ard numbers. The following table gives the actual result of meas-

urement of the contents of two %4-ounce boxes (taken at random)

graded as No. 1, of a certain make which apparently are largely

imported and sold in this country. The thicknesses given are in

thousandths of an inch:

Y%Z-OUNCE, 92 COVER-GLASSES

(Sold as No. 1, which should not exceed limits of 5 to 7 thousandths)

TiS) 9 0037 1n: 13 are .005 in. 4 are .007 in.

5 are .0035 in. 13 are .0055 in. 5 are .0075 in.

Q are .004 in. 8 are .006 in. I is .008 in.

22 are .0045 in. 10 are .0065 in. I is .0085 in.

%Z-OUNCE, 82 COVER-GLASSES

(Sold as No. 1, which should not exceed limits of 5 to 7 thousandths)

Toi1She O03) in: II are .0055 in. 6 are .0075 in.

3 are .0035 in. 5 are .000 in. 4 are .008 in.

7 are .004 in. IO are .0065 in. 3 are .0085 in.

I5 are .0045 in. 7 are .007 in. I is .0095 in.

Q are .0O5 in.

In each case, it will be seen, 48 PER CENT measure outside the lim-

its of No. I.

Mr. Pennock recommends his —— covers as closely true to

their numbering. He will be glad to furnish estimates.

EPS FES ST

NECROLOGY

WILLIAM HENRY DALLINGER, 1842—1909

William Henry Dallinger,* Honorary member of this Society,

was the son of an artist and engraver, and was born at Devonport

on July 5, 1842. As a boy he had leanings towards natural science

and at one time had thoughts of becoming a medical student; but

the deep piety of his nature prevailed and, after a brief training at

Richmond Theological College, he entered the Wesleyan ministry

in 1861, and for the next twenty years “traveled in the circuits” of

Faversham, Cardiff, Bristol, and Liverpool. At the last named city

he remained for twelve years. During these years he not only

kept up his interest in Natural Science, but taught himself German,

Greek, and Hebrew.

In 1880 Dallinger was appointed Governor and Principal of

Wesley College, Sheffield, where he did much to develop the mod-

ern side of the school. There he remained until 1888, when the

Wesleyan Conference, in recognition of the great interest and value

of his scientific work, allowed him to retain his status and privileges

of a minister without pastoral charge, only retaining his position as a

member of the “Legal Hundred.” On leaving Sheffield, Mr. and Mrs.

Dallinger—he married Emma, daughter of David Goldsmith, of

Bury St. Edmunds, and had one son—were presented with some

plate and a handsome sum of money, which he characteristically.

spent in microscopes and other scientific instruments.

During his tenure of the Principalship of Wesley College,

Dallinger was four times elected President of the Royal Microscopi-

cal Society, in 1884-5-6 and 7. Although living 160 miles from

London, he was constant in his attendance at meetings both of the

Council and of the Society, and in order in no way to allow these

meetings to interfere with his work at Sheffield he was in the habit

of returning by the early newspaper train the morning after the

Meeting. His devotion to the Society and the tact he showed in

the Chair were warmly commented on by Dr. Glaisher, Prof.

*The American Microscopical Society is indebted to Mr. A. E. Shipley of Christ’s

College, Cambridge, for permission to reprint this memorial notice; and to the Royal

Microscopical Society for permission to reprint the portrait accompanying.

184 WILLIAM HENRY DALLINGER

Jeffrey Bell, and Mr. Crisp on his retirement in 1888. So great

was his interest in the Society that shortly after resigning the

Presidency he allowed himself to be nominated Joint Secretary, and

for some years he continued to labour whole-heartedly for its wel-

fare. In 1890, 1891, and 1892 he was President of the Quekett

Microscopical Club.

After leaving Sheffield, Dallinger devoted much of his time to

lecturing. He was for many years senior Lecturer of the Gilchrist

Educational Trust. He first lectured for the Trust in 1879 and con-

tinued without a break until the spring of 1909. During the thirty

years he gave about 450 lectures in different towns in the country

for the Gilchrist Trustees. The titles of some of his most popular

lectures were as follows:

“An Hour with the Modern Microscope,’ “Carnivorous

Plants,’ “Contrasts in Nature—the Infinitely Great and the In-

finitely Small,’ “Evolution as Illustrated in the Minutest Forms of

Life,’ “Spiders: Their Work and Their Wisdom,” “Ants,”

“Wasps,” “The Pond and its Minute Inhabitants.” In 1887 he was

chosen to deliver the seventeenth Fernley Lecture, the Lectureship

being a Wesleyan foundation. The lectures are delivered at the

annual Conference. Dallinger took as his subject “The Creator, and

what we may know of the Method of Creation.” As a lecturer

Dallinger was very successful: he had a vivid descriptive style and

a remarkable ability in illustrating both verbally and by drawings

his subject matter. He spared no pains to make his matter at-

tractive and even painted his own lantern slides; in this his re-

markable artistic gifts were apparent.

Some of his more important scientific articles are mentioned

below. The great service he did to students by editing and partly

re-writing Carpenter’s book on “The Microscope” is worthy of

record.

Dallinger was elected a Fellow of the Royal Society in 1880,

and received the degrees of LL.D. from Victoria University, Toron-

to, in 1884; D.Sc. from Dublin in 1892; and D.C.L. from Durham

in 1896. A striking photograph of him is published in the ‘Journal

of the Royal Microscopical Society,’ 1909, and is herewith repro-

duced.

The original contributions made to Science by Dr. Dallinger

were almost entirely confined to the investigation of certain flagel-

WILLIAM HENRY DALLINGER 185

lates. In conjunction with his friend, the late Dr. J. Drysdale, of

Liverpool, he succeeded in working out the life-history of many of

the minuter forms and of making considerable advances in our

knowledge of the processes of decomposition of organic matter,

and of the degree to which unicellular animals can survive great

changes in the temperature of their environment. The dominant

feature of the work of Dallinger and his colleague Dr. Drysdale

was untiring patience and unwearied application combined with a

profound knowledge and mastery over all the technique of the micro-

scope. By using a binocular, one individual flagellate could pass

from the vision of one observer to that of the other and thus on

one occasion a motionless zygote was continuously watched for

thirty-six hours until it burst into a cloud of swarm pores; on an-

other occasion Dallinger watched the same protozoon for a con-

tinuous period of nine hours. By such persistent observation the

life-histories of several of the flagellates hitherto most incompletely

known were worked out. The papers in which the joint authors

recorded the observations are characterised by a singular modesty

and simplicity of language. The simple organisms investigated were

not overwhelmed with long classical designations. Dallingeria

drysdalei (S. Kent) was to them “the hooked monad,”’ Polytoma

uvella (Ehrb.) “the acorn monad,” Tetramitus rostratus (Pertz)

“the calycine monad,” and so on.

Dallinger and Drysdale contributed important facts bearing on

the theory of “abiogenesis.’”” With great manipulative skill and

under the most rigorous conditions they were able to show that

though the temperature of boiling water is fatal to flagellates in an

active state, the spores of these animals can resist a much higher

temperature without suffering harm. For these minute spores can

sustain a heat up to 268° F. in water, and even up to 300° F. or

higher if in a dry state. The demonstration of these remarkable

powers of resistance showed that organic solutions, which had been

thought to be sterile because they had been boiled, often contained

living spores which had survived the heat and were capable of

starting fresh colonies of flagellates. In connection with this part

of his scientific researches was the remarkable series of experiments

by means of which he was able to habituate successive generations

of Dallingeria drysdalei and other forms to gradually increasing

higher temperatures. From a temperature of 60° F. at which these

186 WILLIAM HENRY DALLINGER

flagellates normally live, he gradually raised the solution to that of

158° F. At this heat, which is at once fatal to the normal animals

not habituated to high temperatures, the animals lived and multi-

plied, differing from the original stock chiefly in the marked vacuo-

lation of the protoplasm. He felt that these experiments weighed

against the position Weismann had taken up on the non-inheritance

of acquired characters, and argument did not shake him. He was

intensely preoccupied in his work and gave his contemporaries the

impression of profound earnestness in all he undertook, combined

with a little absence of business method. Before leaving this short

record of Dallinger’s contributions to the advancement of learning

it is well to recall the honest and the truly scientific spirit which

animated both his researches and his writings. This is admirably

expressed in his own words:—‘Let truth come from whence it

may, and point never so grimly to where it may, he would be re-

creant to science who would for one moment hesitate to receive it.

But no less false is it to the foundation principles of true science,

to accept as true, what must constitute the roots of vast generalisa-

tions, except on evidence which no future scrutiny or analysis can

shake.’’*

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*‘Jour. of the Roy. Mic. Soc.,’ vol. 3, pt. 1, p. 16 (1880).

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WILLIAM CHRISTOPHER KRAUSS, M.D.

In the death of Dr. William C. Krauss which occurred at New

York City September 21, 1909, the American Microscopical Society

has lost one of its most active members who was its President in

1899 after having served it faithfully as its Secretary during the

three preceding years. It is therefore most fitting that some mem-

orial of him should find a place in this volume of our transactions.

Writing of him from the standpoint of one whose relations

with Dr. Krauss were those of warm personal friendship contin-

ued through many years, the bare enumeration of the many activi-

ties in which he was engaged, of the many honors which came to

him as the result of his brilliant qualities and his useful life, seems

but a cold and inadequate expression of that which made up the

man, endearing him to his friends and winning the admiring respect

of his fellows in his own profession.

Born in Attica, New York, October 15, 1863, and entering col-

lege at the age of seventeen, he took the medical preparatory course

and that of science and letters at Cornell University, graduating

with the degree of B.S. in 1884 and from Bellevue Hospital Medical

College in 1886, also from the University of Berlin, magna cum

laude, with the degree of M.D. in 1888. He pursued his medical

studies further in Munich and Paris and returning to America be-

gan his professional career as a specialist in nervous and mental

‘diseases in the City of Buffalo, N. Y., which was his home through-

out his life. He attained a well earned eminence as a neurologist

and was recognized as an expert in all matters relating to insanity

and nervous diseases; was widely sought as a consultant and as an

expert witness was always lucid and positive, commanding marked

attention. He was a lecturer at Cornell University; Professor of

Pathology at Niagara University 1890-94, and of Nervous Dis-

eases 1894-99.

He was a tireless worker, designing many instruments for use

in the study of nervous diseases ; was Associate Editor of the Buf-

falo Medical Journal, also of the Neurologisches Centralblatt, Ber-

lin, Germany, and of the Journal of Nervous and Mental Diseases.

He was a prolific author, having published more than one hundred

188 WILLIAM CHRISTOPHER KRAUSS

papers on medical subjects especially relating to the nervous sys-

tem; he translated Professor Mendel’s Text book of Psychiatry

from the German and had completed just before his death a work

upon Tumors of the Spinal Cord. His literary style was clear and

forcible and his deductions were always fortified by strong and

convincing arguments. He was the first to study the effects of

high voltage upon the brain. He was a neurologist to nine important

hospitals, was President of the Erie County Medical Society and

of the Medical Association of Central New York, an organizer

and Secretary of the Buffalo Academy of Medicine, Medical Super-

intendent of the Providence Retreat for Insane, on the board of

managers of the Buffalo State Hospital for the Insane, and a mem-

ber of seventeen learned societies in America and in Europe.

Such briefly were some of the endless activities that absorbed

time and thought and energy, and in which he found the happiness

that follows successfully accomplished endeavors. Among his pro-

fessional associates he was a leading spirit, always honored and

esteemed. With his friends he was genial and kindly, appreciative

of all things good and beautiful and true, finding his way to men’s

hearts, gaining and holding fast their warm affection. In his later

years he became a sufferer from endocarditis and in July, 1909, he

visited Europe in order to consult eminent physicians there. Re-

turning to America, after a stormy ocean voyage which aggravated

his disease, he reached New York September 20, 1909, and died

there on the following day. He is survived by his wife and three

children

Henry R. Howcanp.

RICHARD J. NUNN, M.D.

Dr. R. J. Nunn, of Savannah, Ga., who has been a faithful

member of this Society since 1883, died at his home June 2gth, 1gIo.

He was born in County Wexford, Ireland, Dec. 13, 1831.

He received his preliminary instruction in the Royal College

of Surgery of London and Apothecary Hall, Dublin.

He came to America in early life and graduated from the Sa-

vannah Medical College in 1854, and he practiced medicine in

Savannah, Ga., until about ten years ago, when he retired from

active practice.

He was a captain of Artillery in the Confederate Army during

the war between the States, but later on account of ill health he re-

tired from command, tho he remained on duty in the hospitals.

In the early part of 1876, his health having become impaired,

he made a trip abroad, but yellow fever breaking out in Savannah

during that year he returned and assisted in the fight against that

malady; and as a result of his devotion and services the Georgia

Medical Society drew up a testimonial to his self-sacrifice and devo-

tion, which testimonial was broadly published in 1876.

During his early years, Dr. Nunn held the Chair of Practice in

the old Savannah Medical College, and later held the same position

in the Oglethorpe College.

Dr. Nunn took a deep interest in all scientific and literary mat-

ters, and was actively connected with a great many institutions in

the United States, as well as in Europe.

Locally, he was very prominent ; being a Curator of the Georgia

Historical Society, member of the Board of Trustees of the Telfair

Academy of Arts & Sciences, member of the Board of Managers

of the Savannah Public Library, and a director in several financial

institutions.

He was deeply interested in Free Masonry, and after retiring

from practice gave the majority of his time to that Order, having

held the highest offices in the state, and at the time of his death was

Inspector General of the Scottish Rite of Free Masonry for the

States of Georgia and South Carolina, and an active member of the

Supreme Council 33.

190 RICHARD J. NUNN

He was a man of broad scientific and philosophic culture, he

possessed an unusual wit and excelled as a controversialist, was a

man of exceedingly tender heart and gentle nature, possessing loving

traits of character. It is literally true to say that he was loved by

all who knew him, his broad charity and gentleness attracting to him

people in all the fields of life.

His funeral was one of the most largely attended ever held in

this city.

H. W. WITcOvER.

In addition to the above, the following members of the Society

have died since the last printing of the list of members:

Henry Bausch Frederick W. Mercer

Clark, Gaylord P. Ava Michenor

Frederick W. Kuhne Albert B. Porter

Lomb, Henry George C. Taylor

The Secretary is unable to get information of the following.

Any member knowing whether they are living will confer a favor

by giving the information to the Secretary:

Victor S. Hall, San Francisco.

Mrs. Mary A. D. Jones, New York City.

Wm. F. Sheridan, U. S. Cruiser “Denver.”

CONSTITUTION

ARTICLE [

This Association shall be called the AMERICAN MICROSCOPICAL

Society. Its object shall be the encouragement of microscopical

research.

ARTICLE II

Any person interested in microscopical science may become a

member of the Society upon written application and recommenda-

tion by two members and election by the Executive Committee.

Honorary members may also be elected by the Society on nomina-

tion by the Executive Committee.

ARTICLE III