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TRANSACTIONS

OF THE

American Microscopical
Society

VOLUME XXXII

1913

iP ok

TRANSACTIONS

OF THE

American Microscopical
Society

ORGANIZED 1878 INCORPORATED I8gI

PUBLISHED QUARTERLY

BY THE SOCIETY
EDITED BY THE SECRETARY

See
VOLUME XXXII LG

NuMBER ONE

Entered as Second-class Matter December 12, 1910, at the Postoffice at Decatur, Iill-
nois, under act of March 3, 1879.

DecaTur, ILL.
Review PRINTING & STATIONERY Co.
1913

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SIV aitabdie WA wien”, as, The ps ne tal hain io a as |

PRR UR ARE? Aine Pui,
“0 Sesame Mee He
<“ Es LE em

. J

OFFICERS.
President: F. CREIGHTON WELLMAN, M. D................ New Orleans, La.
First Vice President: F.C: Wattt, Ph:D...........s.0----- Cleveland, Ohio
Second Vice President: H.E. Jorpan, Ph.D............. Charlottesville, Va.
I ERELATY 20 | Vat NV GeATICOWIAW IE Mtctarerss 20a, aay etal -a\evs 1s 5: o.01 os Sreemenersieraie sete Decatur, Ill.
DFCASUKET Als DELANERANGONP aera 8 5 alt tescye o's's\s s «a's ROMO Ore Charleston, Ill.
Custodian: MAGNUS PReAUMee tre tee ch A oiscw cscs 6 25 mene Meadville, Pa.

ELECTIVE MEMBERS OF THE EXECUTIVE COMMITTEE

Rech. SHANG atte coe cote ne hee Bureau Plant Industry, Washington, D. C.
LR RS Se oe See ee eee ne SP ee ae oO Tee Manhattan, Kans.
GO ABC Trerefil Pe Ie ee Oe eae are eRRI rers Se Granville, Ohio

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.
Apert McCatta, Ph.D., of Chicago, II.
at Chicago, Ill., 1883
T. J. Burritt, 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.
Stmon Henry Gace, B.S., of Ithaca, N. Y.,
at Ithaca, N. Y., 1805 and 1906.
A. Ciirrorp Mercer, M.D., F.R.M.S., of Syracuse, N. Y.,
at Pittsburg, Pa., 1896.
A. M. BueiLe, M.D., of Columbus, Ohio,
at New York City, 1900.
C. H. EicENMANN, Ph.D., of Bloomington, Ind.,
at Denver, Colo., 1got.
Cuartes E. Bessty, LL.D., of Lincoln, Neb.,

at Pittsburg, Pa., 1902.
E. A. Birge, LL.D., of Madison, Wis.,

at Winona Lake, Ind., 1903.
Henry B. Warp, A.M., Ph.D., of Urbana, Ill,
at Sandusky, Ohio, 1905.
Hervert Oszorn, M.S., of Columbus, Ohio,

at Minneapolis, Minn., Igto.
A. E. Herrzier, M.D., of Kansas City, Mo.,
at Washington, D. C., tort.
F. D. Heatp, Ph.D., of Philadelphia, Pa.,
at Cleveland, Ohio, 1912.

The Society does not hold itself responsible for the opinions expressed
by members in its published Transactions unless endorsed by special vote.

TABLE OF CONTENTS

FOR VOLUME XXXII, Number 1, January, 1913

Dissemination of Fungi Causing Disease, by F. D. Heald (Presidential
PGULeSGL EOD TOUS) Geisheels's.a wie» bra, ules Soak & Pacetneseaaen ele te ec © waa

Periodicity of Algae in Illinois, 8 Text Figures, by Edgar Allen Transeau

Nature and Classification of Plant Rusts, 5 Text Figures, by Frank D.
BEOER eRik oh cee sis Cities Seu mide Suehedle susie Bhs LER ee ee eee

Notes and Reviews—Notes on Some Peculiar Sense Organs in Diptera
(illustrated) ; Convenient Dropper for Use in Cutting Celloidin Sec-
tions (illustrated) ; Critical Illumination for the Microscope; Clean-
ing Diatoms; Staining Protozoa; Double-stain Method for Polar
Bodies in Diphtheria Bacilli; New Technic. in Staining Diphtheria
Specimens with Toluidin Blue; Notes from the Meeting of the
Illinois Microscopical Society; Bog Solutions and Plants; Effects of
Cropping on Soil Bacteria; Alternation of Generation in the Phaeo-
phyceae; Experiments on the Germination of Teleutospores; Direc-
tion of Locomotion in Starfish; A Rotifer Parasitic in Egg of Water
Snails; Englenids and Their Affinities; An Ameba with Tentacles;
Some American Rhizopods and Heliozoa; Size of Chromosomes and
Phylogeny; Spermatogenesis in Hybrid Pigeons; Male Germ Cells
in Notonecta; Interstitial Cells of Testis and Secondary Sex .Char-
acters; Microbiology in Relation to Domestic Animals; Beginners’
Guide to the Microscope; Microscopy and Drug Examination.......

TICE cig 0. (9) ay\"" SAL SPO nT ee RN EPC SR SL et
Masiutes ab Cleveland . Meeting... 0). 6d 0086 25 Gis eset sinclar Gee ee
CustodiamsP Report «6. sss c Mh eee A ee cs ee
SEWEASUPED (IS MRETOLE | ochre 6s csics dere cebinw oh ap Henne tose eel at tae

31

41

TRANSACTIONS

OF

American Microscopical Society

(Published in Quarterly Installments)

Vol. XXXII JANUARY, 1913 No. 1
THE ADDRESS OF THE PRESIDENT
FOR 1912

THE DISSEMINATION OF FUNGI CAUSING DISEASE
By F. D. HEALD

INTRODUCTION

A great deal has been written concerning the dissemination of
seeds and this topic constitutes one of the regular subjects for treat-
ment in every text book of elementary botany, but the dissemina-
tion of fungi is rarely mentioned. Even the text-books on fungi
or plant diseases give but an inadequate account of this phase of
mycology. Observations and experiments show that fungi pro-
vide for the dissemination of their offspring and the perpetuation
of the species in many and varied ways and in many cases with
great effectiveness. It is undoubtedly true that the agricultural
and commercial practices of our present civilization have very
materially assisted nature in spreading broadcast numerous para-
sitic forms as well as countless numbers of harmless saprophytes.
That fungus pests are more numerous now in this country than
in former years is not imaginary, but a stern reality. It is true
that with the rapid development of plant pathology during the last
decade we have had diseases of plants brought to our attention
more than ever before. The history of our agriculture shows that
with the intercourse between nations, fungus pests have been fre-

6 F. D. HEALD

quently transported from one country to another. It is my purpose
to consider briefly some of the ways by which fungi, and especially
those causing disease, have been and are being disseminated.

HOW FUNGI ARE CARRIED

In the first place brief reference may be made to the state in
which fungi exist during their transport. They may be carried as
spores, as sclerotia, or as mycelium. Most fungi produce from one
to several different kinds of spores, or specialized reproductive
bodies, which provide for the perpetuation of the species, just as
seeds of our Spermatophytes provide for the production of more
seed plants. Spores may be active, that is, motile or capable of
locomotion, but in most cases they are not endowed with the power
of movement; in the former case their own activity may carry them
in an aqueous medium to points far away from the parent plant
that produced them, but in the latter they must be transported by
some outside agency.

Spores are produced by fungi in enormous numbers. It is un-
doubtedly true that vast numbers of spores perish without ever
finding suitable conditions for the production of new plants. It
has been estimated for some species of mushrooms that only one
spore out of twenty billion ever produce a new plant capable of
spore production.t Many contain a minimum of reserve food and
become exhausted in their first attempt to establish themselves ; some
are not able to withstand adverse conditions, such as desiccation,
low temperatures, etc.; certain types germinate at once without a
resting period and thus frequently fail to reach a suitable sub-
stratum upon which to develop. Conidiospores may germinate at
once and they are generally produced in much greater numbers than
the more resistant ascospores which frequently require a resting
period. Figures give but a slight conception of the enormous num-
bers of spores produced by fungi but they serve to emphasize their
prodigality in spore production. It has been determined by careful
analytic methods that a small “spore horn” or “tendril” of the chest-
nut blight fungus may contain as many as 115,000,000 pycnospores.
Cobb states that a single head of smuted oats may contain as many
as 500,000,000 spores, or a sufficient number to give 1000 per square

DISSEMINATION OF FUNGI ye

foot if they were scattered evenly over an acre of ground. The
marvel is that chestnut blight has not become more widely dis-
seminated or oat smut a greater pest. Buller’ estimates that a single
wheat “berry” affected with bunt or stinking smut may contain
over 12,000,000 spores; also that a single fruit body of Polyporous
squamosus produced 11,000,000,000 spores, while the giant puff
ball, Lycoperdon bodista L. produced the enormous number of
7,000,000,000,000 spores.

The production of sclerotia, or dense aggregates of fungus
tissue by fungi is not uncommon, and these structures vary from
minute masses to organs of appreciable size. Some fungi which
produce spores but rarely, rely upon sclerotia for carrying the
species over unfavorable conditions or for dissemination, while in
other cases as in ergot, the sclerotium is only one stage in a rather
complex life cycle. The origin of sclerotia is perhaps not uniform;
they are probably due, in some cases, to the sterilization of a spore
fruit, a pycnidium or perithecium, and certain fungi with scler-
opycnidia suggest this origin. Sclerotia appear to be very effective
structures in perpetuating the species, if we may judge from the
wide dissemination of certain fungi which are propagated almost
entirely by this method. Why more fungi have not discarded the
wasteful and uncertain process of spore propagation for the more
certain method of sclerotia production can only be conjectured.

Fungi may be transported small or even great distances in the
vegetative or mycelial stage. This mycelium may be included with
some dead organic material which furnishes the substratum for its
development or it may be included within the tissue of a plant or
a plant structure upon which it is parasitic.

The following brief outline will give some of the principal
ways in which the dissemination of fungi is effected:

I. Seed-borne fungi—Seed dissemination.
1. By true seeds or fruits.
2. By vegetative reproductive structures.
Il. Air or wind-borne fungi—Wind dissemination.
1. No explosive apparatus.
2. Provided for by an explosive apparatus.
(a) Forcible ejection of the fungus spores from the fun-~
gus fruit.

8 F. D. HEALD

IlI. Water-borne spores—Water dissemination.
1. Active or motile spores.
2. Passive or non-motile spores.
IV. Insect-borne fungi—Insect. dissemination.
1. Insects as carriers.
2. Insects as hosts.
V. Dissemination by other animals.
VI. Dissemination by agricultural and commercial practices.
1. Transport of soil or manure.
2. Transport of infected seed, nursery stock, or hast: cultural
stock.
3. Transport of various commodities.

SEED-BORNE PLANT DISEASES

Many fungi have certainly solved the problem of dissemina-
tion in a most effective way by a relation to the seed of the host
plant at some point in their life history. The fungus may be car-
ried in either the spore, sclerotium or mycelial stage, upon or with-
in true seeds, fruits or vegetative reproductive structures such as
tubers, roots, bulbs, etc. Many of these seed-borne fungi are
pernicious pests and have attained as wide distribution as the hosts
themselves. Since plant diseases have been more intensively studied,
more and more illustrations of seed-borne fungi have been brought
to our notice. Some seed-borne fungi appear so constantly and
generally upon some of our common crop plants that the attendant
symptoms are not unfrequently interpreted by the untrained, as a
normal accompaniment.

Anthracnose of the bean was one of the first diseases that was
demonstrated to be disseminated by true seeds. This was first
proved by Frank? in 1883, although later investigations have re-
peatedly claimed the honor. The mycelium of the fungus grows
through the pod and into the seed during the period of maturing,
and is there ready to resume its growth when the seed germinates.
Experiments tend to show that this disease is introduced into a
field very largely if not entirely by the ruse of infected seed, and
that the spores are not generally spread from one field to another
by the wind.* The Ascochyta blight of peas is another disease that
behaves in a similar way* and Barre has recently shown that the
widely disseminated anthracnose of cotton bolls is of a like nature.®

DISSEMINATION OF FUNGI 9

Since infected seeds, in so many cases, show no external evidence
of the disease, the fungus is insured a wide dissemination, even
with the most careful practice in the selection of apparently clean
seed. Seed-borne fungi do not appear to be confined to any definite
groups but this method of dissemination may prevail whenever the
ripening ovary is infected. The downy mildew of the Lima bean
and the white rust of the oyster plant constitute two excellent il-
lustrations from the Peronosporales. The seed of the oyster plant
may be so badly infested with the white rust as to entirely destroy
the crop. The black-leg or Phoma wilt of cabbage,®° a disease
known in this country only during the past few years, was un-
doubtedly introduced from Europe with imported seed. Chapman‘
has recently called attention to the seed dissemination of three dii-
ferent onion diseases: smut due to Urocystis cepulae; brown mold,
caused by Macrosporium porri; and downy mildew, referred to
Peronospora schleideniana.

Fruits which function as seeds may also act as carriers of the
parasite. Perhaps the best known and most familiar illustrations
of this class are the seed-borne smuts of cereals, in which case the
fungus is either on the surface of the caryopsis in the spore stage
or has penetrated the pericarp and persists as a dormant mycelium.
The loose smut of oats and the stinking smuts or bunt of wheat
are good illustrations of the former, while the loose smut of wheat
is one of the most notable illustrations of the latter condition. The

‘bunt or stinking smut of wheat is undoubtedly more prevalent in

our cultivated wheat fields than any similar species upon a wild
host, or at least it was until the introduction of fungus “‘steeps” for
its prevention. The ordinary process of harvesting is one that has
very materially assisted in the dissemination of the fungus. When
a wheat plant becomes infected with bunt every head produced by
a single stool is smutted and all the “berries” are destroyed or trans-
formed into “smut berries.”” Smutted plants may be scattered here
and there through the field, and during the threshing process smut-
ted berries are mixed with the normal sound grain; but many of the
smut grains have the thin outer membrane ruptured, thus setting
free the spore mass and the individual spores become scattered over ~
the normal grains, lodging particularly in the “brush” and in the

10 F. D. HEALD

suture.s Wheat may be so badly smutted that the normal grains
are discolored by the large numbers of smut spores adhering to
the berries. Of course nobody would think of using such grain
for seed, but unfortunately smut spores may be present in minute
quantity without giving any indication of their presence. It is pos-
sible, however, for the scientist to determine whether seed wheat
is infected with bunt to even a slight extent. If the seed is washed
or scrubbed in sterile water, the washings centrifuged, and the sedi-
ment examined with the microscope, the presence or absence of smut
spores can be determined. This method was first used in this coun-
try by Bolley.®

In the loose smut of wheat wind dissemination of spores is
combined with seed transport of the dormant mycelium. The inflor-
escence of wheat is completely destroyed and the dry, powdery mass
of spores scattered by the wind. This maturity of the smut coin-
cides nearly with the blossoming time of the normal head and the
scattered spores may be responsible for a blossom infection. The
fungus penetrates the developing ovaries and remains as a dormant
mycelium hidden within the seed and exhibiting no warning of its
presence. It is there, however, ready to resume activity with the
awakening of the seed. Two diseases of beets are known to be car-
ried by the seed (fruits): the Phoma rot of beets which has been
so prevalent in Europe, and our well-known Cercospora leaf spot.
The fungus (Phoma betae) which causes a rotting of the maturing
beets also causes a serious damping-off of young seedlings.1° A re-
cent illustration has come to the writer’s attention of chestnuts
bearing the sclerotia of a fungus upon the surface.

Fungi which cause disease may be disseminated by the use
of infected vegetative reproductive structures such as tubers, roots,
rhizomes, bulbs or corms, for the production of a new crop. All
such structures are gorged with reserve food and their tissues in
a semi-dormant condition, easily invaded by certain types of fungi.
In the majority of these structures the presence of an internal in-
fection is revealed by some discoloration or disorganization of the
storage tissue, or the fungus may be more superficial and exist in
either the mycelial or the sclerotium stage. The Irish potato stands
preeminent among the cultivated plants, for its numerous tuber-

DISSEMINATION OF FUNGI Il

borne diseases.. In recent years many of these tuber-borne dis-
eases have made such headway as to seriously threaten the potato
growing industry in many parts of the country. Mention may be
made of the late blight of the potato with its internal mycelium,
the dry rots which also produce internal discolored areas pervaded
by mycelium, scab with the more superficial corroded areas, potato
wart with its external warty excrescences, and Rhizoctonia or po-
tato rosette with superficial sclerotia. It has been shown by Massee
that wart may also be transported by the use of tubers which show
no external evidence of the disease, the spores being lodged in the
“eyes.” Some of these tuber-borne fungi may so exist in the soil
as to infect perfectly healthy seed, but their original introduction
can be traced in numerous instances to the use of infected seed.

The sweet potato affords several illustrations of diseases which
are frequently introduced by the use of infected seed roots. Among
these may be mentioned the black rot, due to Sphaeronema fimbri-
atum.1+ Several onion diseases may be introduced into new fields,
by the use of infected sets or the use of imported bulbs for growing
seed. Disease of the gladiolus are spread by the planting of corms
already infected, and the same may be said concerning the bulbs
of various greenhouse plants. As specific examples of bulb-borne
diseases mention may be made of the Japanese lily disease, due to
Rhizopus necans, and the anther smut of Scilla latifolia which de-
velops from mycelium present in the bulb.

AIR OR WIND DISSEMINATION

The spores of many fungi are carried away from the parent
plant by means of air currents. In general it may be stated that
spores which are adapted to wind dissemination are liberated from
the fungous fruit as a dry, powdery mass or are born singly or in
loosely connected chains upon the ends of aerial sporophores, from
which they are easily detached. The minute size of fungous spores
makes unnecessary special devices for rendering them buoyant, and
it is always possible to obtain various forms from the dust that
settles to the surface of objects in closed rooms or in the garden
or fields. The majority of the air-borne spores appear to be those
of harmless saprophytic forms, and many of the statements made

I2 F. D. HEALD

concerning the part which wind plays in the dissemination of para-
sitic species are based largely on analogy rather than supported by
direct experimental evidence.

Observational evidence is of little value in determining the part
of air transport of fungus pests which are confined to a single
host, but some heteroecious rusts give us undoubted examples of
wind dissemination which so far as I know have never been defi-
nitely proved experimentally. The cedar rust which alternates be-
tween the cedar tree and the apple tree, and must pass from one
to the other, is an excellent example. Susceptible varieties of apples
standing adjacent to infected cedars will show a high percentage of
infection, sometimes as many as 200-300 distinct spots to each leaf
and the number of infections per leaf will decrease with the dis-
tance until the average is either one or less to each leaf. In addi-
tion, the rapidity and universality of the infection following the
gelatinous stage of the cedar apple can be explained in no other
way than by the wind dissemination of the sporidia, which Coons’?
has shown are forcibly discharged from the promycelia. The part
which wind plays in the spread of our cereal rusts has probably
been greatly overestimated. The old idea that rusts are spread
gradually by the wind from the southern plains country to the
north as the season progresses, has been largely abandoned as our
ideas of their life habits have been modified. The theory of the
wind dissemination of the cereal rusts gives a beautiful example
of the inefficiency of reasoning alone as applied to the processes
of nature.

It is undoubtedly true that many of our present day state-
ments in regard to wind dissemination of spores would need re-
vision if the rigid test of experimental evidence were applied to
them. The application of scientific methods to the determination
of the part which wind plays in the dissemination of parasitic fungi
opens up an interesting field for investigation. It may not be amiss
to mention briefly some of the methods which may be utilized, and
it may be stated at the beginning that. these have already yielded
valuable results in the few cases where tried.

In the employment of the exposure plate in the field under
natural conditions the pathologist is but imitating the bacteriologist.

DISSEMINATION OF FUNGI 13

As far as I have been able to determine this method was first em-
ployed in this country in the field for determining the prevalence
of parasitic fungi by Wolf** at the writer’s laboratory in 1907. It
had, however, previously been used by Saito in Japan. It should
be emphasized in this connection that the exposure plates should
be made under natural conditions if we are to obtain reliable re-
sults. Erroneous conclusions may be drawn from results obtained
under artificial conditions even if they are obtained from field ex-
periments. It is evident that this method is of value only when
the spores of the pathogens under investigation will germinate
upon the medium that is available. There are many parasitic forms
the spores of which will not germinate upon the ordinary culture
media, and in investigating the prevalence of these spores this
method is clearly at fault. Exposure plates may be made in the
field or in the laboratory with the substitution of an artificial air
current. As an illustration of this type, the experiments of Ful-
ton** with the spore horns of the chestnut blight fungus may be
mentioned, although his negative conclusions would have been ob-
tained by a priori reasoning.

It is at once evident that the exposure plate can give no exact
quantitative results, but only the relative abundance of the forms
obtained. The aspiration of the air through a “spore trap” and
the determination of the number of spores per unit quantity of air
can be readily performed by the employment of the ordinary bac-
teriological method. This method was used by Wolf in 1907 and
has recently been employed by Anderson. The poured plates in
this method do not reveal the possible presence of spores which will
not germinate upon our culture media. In order to determine the
presence of these, other methods must be employed. There are
two methods suggested which are somewhat comparable to the two
just outlined. First, if quantitative results are not desired a glass
funnel, the inner surface of which is coated with glycerine, may be
exposed for a certain length of time in the open and the spores
which fall into it washed down with sterile water and the washings
centrifuged, after which the sediment thrown down may be ex-
amined with the microscope. If quantitative results are desired
the aspirator should be used, the contents of the sugar tube dis-

14 F. D. HEALD

solved in a known volume of sterile water, all or a unit quantity
centrifuged and the sediment examined microscopically by the use
of the Leitz-Wetzlar counting apparatus. The results of the micro-
scopic tests may be substantiated by inoculations made by using a
suspension of the spores obtained from the air analyses, and in
certain cases this method can be used to good advantage.

Inoculations by wind-borne spores under controlled conditions
has been used in some cases for determining the part which this
method of transport of spores plays in the life history of the fungus.
As an illustration, the inoculations with chestnut blight fungus made
by Anderson may be cited. The wounds were protected so as to
exclude insects and spores washed down the tree by rains.

A large number of fungi, the spores of which are disseminated
by air currents, produce the spores in such a manner that they are
easily detached and carried away by the wind or in a dry powdery
mass. In such cases there is no explosive apparatus that ejects the
spores into the air and it is entirely the force of air currents that
sweep them away from some exposed position, or they are borne
in such a manner that they rattle out of the fungous fruit, frequently
as a result of agitation of the host plant by the wind.

In many fungi, particularly the ascomycetes and basidiomycetes,
the spores are forcibly ejected into the air' and can then be swept
away from the fruit by the wind. It should not, however, be con-
cluded that all fungous spores that are forcibly ejected, are adapted
for wind dissemination. There are numerous cases in which the
forcibly ejected spores are not adapted for wind dissemination but
after they have come to rest are removed to more distant locations
by other agencies.

The loose smuts of cereals and other grasses which produce
myriads of spores in the deformed or destroyed inflorescences of
their hosts are supposed to be disseminated largely by wind-borne
spores. The powdery condition of the spores, and their elevated
position upon the host certainly suggests this method of transport.
The uredospores and aecidiospores of many rusts and particularly
the sporidia are wind-borne, while the summer spores of the
powdery and downy mildews are produced in such a way as to
suggest this method also. The aecial stage of Gymnosporangium

DISSEMINATION OF FUNGI 15

macropus shows an interesting adaptation.‘* During dry weather
the segments of the pseudoperidium are curved outward in stellate
form but during humid or rainy periods they approach each other
and partially or completely close over the spores. The spores are
thus set free at a time when they are most likely to be carried away,
and this is especially important in heteroecious forms which must
reach the alternate host or fail to develop.

Judging from the results of experiments in the field it seems
that the most prevalent spores belong to species of the imperfect
fungi, and especially to the Hyphomycetes. Perhaps these facts are
to be explained by the omnipresence of certain species of this group
rather than to the fact that they are better adapted for wind dis-
semination. In the work carried out by Wolf only a single
pycnidial form, Phyllosticta, was obtained during a long series of
orchard tests. The work of Burrill and Barrett'* on the wind dis-
semination of Diplodia zeac, the fungus causing dry rot of corn,
may be mentioned in this connection. They found by field tests
that spores of this fungus were carried by the wind and they at-
tribute much oi the infection to wind-borne spores from old stalks.
There appears to be little direct experimental evidence to show to
what extent ascomycetes which forcibly eject their spores from
the asci are disseminated by the wind. In many cases the ejected
spores are surrounded by a sticky material and a priori reasoning
would suggest that in such cases they are not extensively carried
by the wind. It is at least suggestive that the spores of ascomycetes
are obtained so rarely in exposure plates in the open. Further in-
vestigations may show that ascospores are more generally scat-
tered by the wind than present experiments show. It is the idea of
the writer that wind-borne ascospores may be responsible for the
spread of a fungus in the immediate environment, but that long
jumps or wider dissemination are accomplished by other agencies
than the wind.

The puffing of spores as in Peziza, Urnula, and other Dis-
comycetes in which there is a simultaneous discharge of numerous
asci, is undoubtedly an adaptation for wind dissemination. Since
most of these fruits are produced close to the ground, the forcible
expulsion of the spores must materially assist in their being car-

16 F. D. HEALD

ried away by air currents. In the Hymenomycetes the forcible ex-
pulsion of the basidiospores from the sterigmata helps simply in
liberating the spores from the sporophores, while the normal posi-
tion of the hymenium makes the fall of the spores inevitable, and
convection currents assisted by wind carry them away to more
distant points.

Spores that are destined for wind transport may be set free in
a cloud by the explosion of the fruit of the host plant. A beautiful
illustration of this is to be found in the smut infected fruits of
Oxalis, which burst and liberate the spores in much the same way
that the capsules of touch-me-not expel their seeds.

DISSEMINATION BY WATER

Liquid water plays a very important part in the dissemination
of some disease fungi. In certain groups of fungi free water is
necessary for the development of some stage in the life cycle, that
is, in those that produce active or motile spores, swarm spores, or
zoospores. These active spores make their way for a shorter or
greater distance from their point of origin as a result of their own
power of locomotion. In other cases passive or inactive spores
may be washed down from the host plant by rains and carried
away by natural water currents or spread along the course of irri-
gation ditches.

The aquatic habit has been retained to a greater or less extent
by the various species of the pond scum parasites, the water-molds,
the white rusts and the downy mildews. The parasites of the pond
scums are completely dependent upon free water for their dis-
semination and this appears to be equally true of some Chytridiales
parasitic on seed plants. The natural habitat of the cranberry is
particularly favorable for the development of the cranberry gall
due to Synchytrium vaccint. It is also noteworthy that Urophlyctis
alfalfae has made its appearances in this country in a number of
regions in the Pacific coast country where alfalfa is grown under
irrigation.

The Saprolegniales or water-molds include a much larger num-
ber of saprophytes than parasites and in the majority of forms are
strictly aquatic in habit. Some species are parasitic on the eggs

DISSEMINATION OF FUNGI 17

and young of fish and also frequently gain entrance to the bodies
of adults. While countless numbers of young fish and other aquatic
forms annually fall a prey to the ravages of these fish molds, it 1s
under the artificial conditions of the fish hatcheries, that the water
molds are most likely to become epidemic. Not only may the water
become filled with myriads of these motile spores of the water
molds, but the diseased fish may transport the fungus for long dis-
tances and introduce it into entirely new localities.

There are two fungi which belong to the water molds that
cause destructive plant diseases. One of these, Pythiwm de Bary-
anum, is the cause of a damping-off of seedlings, a great variety of
species being attached. This fungus has attained practically a
world-wide distribution. At just the periods that young seedlings
are establishing themselves in the soil, this damping-off fungus
finds conditions favorable for the development and dissemination
of its swarm spores. These motile spores are able to swim actively
in the soil water and are also spread by the spattering of rain and
the meteoric water which flows over or through the surface layers
of the soil. It is this production of enormous quantities of motile
spores at times when the seedlings are young and susceptible that
makes this one of the most destructive of the damping-off fungi.
The other water mold referred to has been known to science only
since 1906,'* but when it first appeared in California it made such
headway as seriously to threaten the citrus industries of that sec-
tion of the country. The fungus in question, Pythiacystis citroph-
thora, causes the disease of lemons known as the brown rot, and
the history of the discovery of the cause of this disease and of its
method of dissemination forms one of the most interesting chap-
ters in modern plant pathology. The natural habitat of the fungus
is the damp soil of the orchard, and irrigation apparently favors its
development. The spores of the fungus are not wind-borne and
only fruit either in contact with the soil or very low down on
the tree becomes infected. The motile spores can easily reach fruits
in contact with the soil by swimming through the soil moisture, and
the spattering of rain is supposed to carry spores to the lowermost
fruits that are free from contact with the soil. If lemons were
handled like apples in preparing them for the market, the brown

18 F. D. HEALD

rot would never have become such a serious pest, but lemons must
be washed or scrubbed, and it is just this process that makes a more
extensive infection possible. Dirt bearing the fungus is transported
to the washer on the surface of the fruits and the fungus finds in
the water of the washer favorable conditions for its development.
The washers thus become infected with the fungus and the water
through which the lemons must pass becomes filled with myriads
of the motile spores, some of which may gain entrance to the fruit
during the washing and scrubbing process. Of course greater care
is now taken in cleaning the washers and the use of fungicides in
the water prevents the development of the fungus.

The Peronosporales, including the downy mildews and white
rusts are not so completely dependent upon moisture for the dis-
semination of the spores, since their conidia (sporangia) are borne
in such a way as to be Carried away by the wind. They do, how-
ever, show a greater dependence upon moisture than many of the
fungi that have abandoned entirely the production of motile spores.
It is an especially noteworthy fact that the late blight of the potato,
caused by Phytophthora infestans, is epidemic during wet seasons
and is limited geographically by rainfall. This partial dependence
upon free water for its development probably explains the reason
the late blight has never been a serious potato disease in the drier
portion of the plains country.

It is undoubtedly true that rain and water currents play a very
important part in the dissemination of fungus spores. In the first
place rain may assist in the further transport of wind-borne spores
that have been lodged upon plant surfaces. In case of wound in-
fections the spores may be finally carried into the wound by rain
washing down over spore laden surfaces, the spores finally coming
to rest in a more favorable position for germination. Many fungus
spores are rarely carried away from the fruits in which they de-
velop except through the agency of rain. This seems to be par-
ticularly true of many forms producing pycnidia surrounded by
a more or less evident mucilaginous secretion which prevents their
release from the fruit or their separation from each other except
in the presence of sufficient water to dissolve the cementing sub-
stances. Such spores may accumulate as sticky or waxy masses

DISSEMINATION OF FUNGI 19

over the acervulus or they may be pushed out through the ostiole
of a pycnidium as the result of growth of others within. In certain
forms the extruded spore mass takes on the form of a long, coiled,
flattened or cylindrical thread, which is designated as a “spore horn”
or tendril. These sticky spores are produced in enormous quantities
during warm, humid periods and the spore masses dry down and
become hard if they are not washed away by rains. Many spores
of this type retain their vitality for a considerable period as long
as they are embedded in their mycelaginous secretion, but soon
succumb to desiccation and other unfavorable factors as soon as
they are separated by the rains.

In this connection several illustrations may be mentioned.
Valsa leucostoma, the fungus causing die-back of peaches, plum
and apricots produces conspicuous, brown or amber-colored ten-
drils which ooze out from the pycnidial stromata embedded in the
bark. These are always abundant during the humid period follow-
ing a rain, but disappear entirely except from especially protected
positions during the first precipitation of any amount. During a
warm rain the spores are being produced in enormous numbers but
they are washed away as rapidly as they are extruded so the ten-
drils do not become visible until the rain has ceased. What has
been said concerning the conidiospores of the die-back fungus ap-
plies equally well to those of the chestnut blight fungus, Endothia
parasitica. It has recently been determined by experiments carried
out under the writer’s direction that the so-called summer spores
are washed down from the blight lesions in enormous numbers,
even during the winter rains when the temperature is but little above
the freezing point.

There seems to be little evidence that the spores of bean
anthracnose are wind-borne. The facts known concerning this dis-
ease indicate that rain and dew are of utmost importance in its
spread after it has once been introduced by the use of infected seed.
The fact that the spores of the Melanconiales are found so infre-
quently in the air lends support to the theory that these fungi are
largely dependent upon rain and other agencies besides wind for
their transport.

There is not much direct evidence to show the part of running

20 F. D. HEALD

water in the transport of non-motile spores. If these spores fall
into streams or irrigation ditches there is every reason to suppose
they will be transported for some distance. In some forms germina-
tion would take place in a few hours, and so the possibility of trans-
port for long distances would be excluded. It is claimed that irri-
gation water plays a very important part in the dissemination of
the late blight of celery in California.** The spores may be washed
away from the pycnidia and carried along the trenches with the
irrigation water.

INSECTS AND DISSEMINATION OF FUNGI

The relation of insects to the spread of plant disease is a sub-
ject to which sufficient attention has not been directed. The nu-
merous insect-borne diseases of man and animals suggests a similar
relation between insects and plant diseases. In most insect-borne
animal diseases the insect acts as an intermediate host, and is not
simply a carrier as is true in the case of the “typhoid fly.” Not a
single instance of an insect acting as an intermediate host for a
fungus causing a plant disease has yet been brought to light, but
the part which insects play in the dissemination of fungi is limited
by their work as carriers and as producers of wounds which make
infection possible.

It seems probable that insects play a very important part in the
dissemination of saprophytic fungi. Fungus fruits are in many
cases rich storehouses of food, and insects have become mycopha-
gists, either utilizing’ the natural growths or becoming cultivators
of fungi as is exemplified by the “ambrosia” beetles, or the ants
with their fungus gardens. In visits to fungus fruits insects
cannot fail to carry away spores, in much the same way that in-
sects carry away pollen (spores) from the flowers which they visit.
In some cases there seems to be a definite adaptation to insect trans-
port of spores, while in others the transport is apparently only acci-
dental.

The carrion fungi, of which Phallus impudicans may be tak-
en as an example, attract flies to their spore-producing surfaces by
their characteristic odor. The greenish slime in which the spores
are embedded also contains three sugars, levulose, dextrose, and

DISSEMINATION OF FUNGI 21

another intermediate between dextrose and gum. These and the
spores are greedily eaten by flies. Fulton? has shown that flies
transport the spores in millions by the adherence of them to their feet
and proboscides, and also that the spores will germinate after they
have passed through the digestive canal of these insects.

The sphacelia stage of the ergot of rye and other grasses gives
a beautiful example of insect dissemination. The ovaries become
infected at flowering time by wind-borne ascospores and the pro-
duction of conidia soon begins. This production of conidia is ac-
companied by the secretion of a sweet substance, the so-called honey
dew, which is eagerly sought by insects. A rapid dissemination
of the fungus is accomplished since the visiting insects carry away
spores and scatter them as they fly from flower to flower. Although
the sooty mold of the orange and other citrous fruits is not a defi-
nite parasite, it becomes a troublesome pest. This fungus is asso-
ciated with and spread by the white fly, or Aleyrodes and other
species of aphid-like insects.2* The secretions of sweetish fluid
constitutes the pabulum which makes possible the development of
the fungus. Some anther-inhabiting fungi are undoubtedly dis-
seminated by insects. This is true of the smut of various species
of the pink family. The affected anthers produce smut spores in-
stead of pollen and these are carried from plant to plant by the
visiting insects, thus assisting in the dispersal of the fungus.

The manner of spread of fire blight of the pear and apple was
for many years more or less of a mystery. The bacteria causing
the disease are set free upon the surface of the diseased parts in
sticky droplets, and are quickly killed by exposure to sunlight and
desiccation. Waite*® first showed that the rapid spread of the dis-
ease during the spring is due to insects and especially to bees. The
bacillus lives over winter in only a small percentage of the affected
branches and is spread from these by insects. The blossoms be-
come infected, the bacteria multiplying in the nectar, and thus the
disease is spread from flower to flower by bees. Insects like leaf
hoppers and others which bite the delicate young shoots are also
important agents in the spread of fire blight.

An intimate relation between certain mites and the bud rot
of carnations was established by the writer.2* The mites are always

22 F. D. HEALD

found in the buds that have been rotted by Sporotrichum poae ; they
find in the mass of rotted petals a most favorable substratum for
their development. The young mites which migrate from diseased
to healthy buds carry spores of the fungus with them and thus in-
oculate the healthy buds, their presence serving to accentuate the
severity of the trouble.

The literature of plant pathology contains not infrequent refer-
ence to the part which insects play in the dissemination of plant
diseases, but these are in many cases generalizations not based on
direct experimental evidence. Murrill and others** have stated
that the spores of the chestnut blight fungus are carried by insects,
but up to the present date there are no published experiments which
really substantiate this statement. It seems probable, however,
that this early claim will be supported by experiments now in pro-
gress.

Massee”’ has pointed out the fact that the rapid spread of apple
canker due to Nectria ditissima in Engiand coincides with the in-
troduction and spread of the “American blight or wooly aphis.’”’ He
makes the following statement: “I think it would be scarcely an
exaggeration to say that if we had no “wooly blight” we should
have no “canker,’’* that is in the sense of an epidemic. It should
be pointed out, however, that this opinion which is not based on ex-
perimental evidence is not entirely acceptable. The whole problem
of the relation of insects to plant diseases is one that merits more
attention than has been given to it, and investigations in this line
may be expected to yield important results.

DISSEMINATION BY OTHER ANIMALS

The prevalent notion in regard to the part which other ani-
mals play in the dissemination of disease-producing fungi is ex-
pressed by the following quotations:

“Insects, birds, snails and slugs are known to be unconscious
agents in the dispersion of spores, whereas dogs, hares, rabbits,
etc., running through a field of corn, potatoes or turnips act after
the fashion of the wind by bringing into contact adjoining
plants.”°

“Mites, flies, birds, mice, etc., carry spores adhering to their

DISSEMINATION OF FUNGI 23

bodies from one place to another; and probably are frequently the
unconscious cause of a new infection or the rapid spread of an epi-
demic due to fungi.”

While the above are generalized statements based on but little
experimental evidence this possibility has been demonstrated in
some cases. Massee reports that snails and slugs are instrumental
in spreading powdery mildews. Slugs allowed to crawl over mil-
dewed leaves and then over healthy leaves left behind spores which
soon caused the appearance of mildew along their pathway.

Birds, especially woodpeckers, have been mentioned by var-
ious writers in connection with the dissemination of the chestnut
blight fungus. While the few tests reported to date (15) were
negative it seems reasonable to believe that bird transport is a
possibility. Woodpeckers frequently visit the chestnut blight lesions
in search of insect larvae, and it will be remarkable if they do not
carry away blight spores upon their feathers, bill or feet. While
the writer is not yet ready to make any positive statement in re-
gard to the part which birds play in the spread of this pernicious
disease of the chestnut, a pertinent fact may be mentioned. By
the employment of careful analytic methods a single hairy wood-
pecker has been found to be carrying as many as twenty different
kinds of fungus spores. Johnson has suggested the possibility that
the bud-rot of cocoanut may be carried by turkey buzzards as well
as by certain insects and reports some experiments which seem to
lend support to his contention.?°

In considering this subject observations on and experiments
with saprophytic fungi may be mentioned. Voglino”? has shown
that slugs eat the sporophores of fleshy agarics, especially the
hymenium, and that the spores begin to germinate in their intes-
tines and afterwards continue to grow in the ground in which
the slugs burrow. The subterranean ascus-bearing tubers of truffles
are sought as food by rodents and the spores of these fungi are sup-
posed to be dispersed by this means.

Herbivorous animals play a very important part in the dis-
persal of certain dung-inhabiting fungi. A considerable number
of these dung inhabiting fungi expel their spores with considerable
force from the fruiting body. If it were not for the grazing ani-

24 F. D. HEALD

mals these spores would in most cases, be carried no farther than
the force of their projection would take them for they are sticky
and adhere to the surface of the objects upon which they light.
Foliage with attached spores is eaten by grazing animals and the
spores being able to pass through the intestines of the animal un-
harmed, find a suitable substratum for their development at some
distant point. Among those forms which have developed this
habit the following may be mentioned: Pleurage, Ascobolus and
other black-knot allies which shoot their spores by the explosion
of the ascus; and certain Hymenomycetes like Coprinus and allies.

DISSEMINATION BY AGRICULTURAL AND COMMERCIAL PRACTICES

Numerous instances of the transport of disease producing or-
ganisms by man as a result of agricultural and commercial prac-
tices are known. With the development of our agriculture and
the intercourse between nations the part of man in the dissemina-
tion of plant diseases has become more pronounced. The possi-
bility of the spread of diseases and insect pests from one region
to another has long been known and states have endeavored to
safeguard the agricultural and horticultural interests of the people
by laws relating to inspection of nursery and other stock. From the
standpoint of fungus diseases this inspection has not been as
effective as might be desired for in the majority of states the in-
spectors have been entomologists, familiar only with the more
evident plant diseases such as crown-gall or black-knot, and not
skilled in the detection and diagnosis of the more obscure troubles.
This statement is not imaginary but is based on facts, for the
writer has, in numerous instances, visited nurseries immediately
after the official inspection and found various fungus diseases
prevalent that were entirely overlooked. The demand for nation-
al legislation making restrictions which might govern the intro-
duction of pests from foreign countries and the spread of troubles
from infected regions to those free from the disease led to the
recent passage of the Plant Quarantine Act.?®

Fungi which are primarily soil dwellers may be carried by
transport of soil. Spores which have not yet germinated may be

DISSEMINATION OF FUNGI 25

incorporated with the soil but in some of the most serious troubles
the fungus is present in the mycelial or in the sclerotial stage. One
of the agricultural practices that should be condemned on this
account is the use of alfalfa soil for the inoculation of a field with
the nitrogen-fixing bacteria. If, for example, the soil selected
contained the mycelium of the alfalfa Rhizoctonia or that of
the southern fungus of cotton root rot, these troubles might be
introduced into new fields. We have reason for believing that the
sterile mycelium of such fungi will endure considerable dessica-
tion without losing its vitality. The “spawn” of mushroom grow-
ers is but mycelium preserved and temporarily dormant in dried
bricks of compost. It is particularly in the cultivation of plants
under glass that we find the introduction of fungi and other pests
with the soil. The drop or Sclerotinia disease of lettuce is one
that persists by the development of sclerotia that may remain in
the soil.*° It seems to be true that root-knot of roses is frequently
introduced into greenhouses by the selection of soil previously in-
fected with eel-worms.

Some parasitic fungi are capable of passing one stage in their
life history in the soil or in compost. This is especially true in
the case of certain smuts, of which corn smut is a most notable
example. It is a common practice on farms to feed corn fodder
or stover to cattle and return the compost to the soil. The smut
spores find in the compost especially favorable conditions for
germination and also for the production of secondary sporidia in
countless numbers by a process of budding. Compost originating
from the use of smut-infected corn may thus contain billions of
sporidia of smut that are returned to the soil of the corn field
where they are ready to produce new infections. The work of
cultivation and the movement of wagons and teams from one field
to another may be responsible for the spread of disease producing
organisms. It is undoubtedly true that the mycelium of the cotton
root rot is extensively spread through the fields during the culti-
vation of the crop. It is claimed by Massee that club root, or the
finger and toe disease of cruciferous plants, may be spread by soil
adhering to cart wheels, tools, shoes, etc. The practice of allow-
ing diseased plants, fruits or other products to fall to the ground

26 F. D. HEALD

and remain there unmolested save for the work of nature’s scav-
engers is a too common practice that favors the spread of disease.

The part which man has played and is playing today in the
dissemination of plant diseases can not be overlooked. This is the
inevitable result of our specialized agriculture and modern com-
mercial practices, but the distribution of diseases by the importation
of infected seed, nursery and horticultural stock, and the transport
of various commodities, can and should be reduced to a minimum
by the employment of all possible safeguards.

Many of our serious plant diseases have been brought to this
country from Europe or other foreign countries, and we can point
in turn to pests which have been transported from America to
Europe and elsewhere. The influence of climatic and edaphic fac-
tors upon the development of disease in epidemic form must be
taken into consideration. It is by no means certain that a fungus
pest which has proved serious in one country will prove equally
serious in another, but the existence of serious diseases in a coun-
try or region should be kept in mind, and importations of sus-
ceptible stock made with extreme care.

Commercial concerns, state agricultural experiment stations
and departments of agriculture, and the United States Department
of Agriculture are all importing seeds and plants from foreign
countries, in the endeavor to find plants valuable for the trade or
better suited to the agriculture of the country. If we reflect upon
the nature of seed-borne fungi it must at once be evident that this
wholesale importation of seed is bound to be a prolific source of
the spread of disease. It is undoubtedly true that the black-leg
of cabbage previously referred to was brought to this country by
infected seed® and that potato wart recently reported from New-
foundland was introduced from England.*® These are illustra-
tions of recent importations and it was the discovery of this latter
disease in this country that gave one of the strong arguments for
the passage of the recent Plant Quarantine Act.

The shipment of nursery stock is frequently responsible for the
appearance of diseases in hitherto uninfected territory. In the
greater percentage of even well managed nurseries plant diseases of
various kinds may be found in profusion, the massing together of

DISSEMINATION OF FUNGI 27

individuals favoring their development, but the neglected nursery
is literally a pest house of plant diseases. The Pennsylvania Chest-
nut Tree Blight Commission has records of spot infections of blight
that were traced to the planting of nursery stock that was diseased
at the time of shipment. The diseases of nursery stock that are
accompanied by easily recognized symptoms, may easily be guarded
against by rigid inspection of all stock offered for shipment, and
fumigation is a reasonable safeguard against the spread of many
insect pests, but unfortunately some of the most serious diseases
can be carried by stock which shows no indication whatever of its
presence. One of the most striking examples of this is to be found
in the case of the seedlings of white pine affected with the so-called
blister rust.4t_ The fungus causing this trouble has a period of in-
cubation in the bark of nearly a year before it causes the character-
istic hypertrophies, and for this reason inspection at the time of
importation is but an imperfect insurance against its introduction.
In such extreme cases the quarantine of infected regions and the
restrictions of shipment of stock that might carry the disease is
entirely justifiable. Diseases like the black-knot, peach leaf curl,
orange rust of raspberries and blackberries, and many others that
produce a perennial mycelium in the host may easily be transported
by infected nursery stock, while there are many opportunities for
the transport of resistant spores on the surface of florists’ green-
house stock or field-grown plants. Besides this, incipient infec-
tions of various fungi may be present in either herbaceous or woody
plants, and entirely escape detection at the time of shipment.

The transport of various commodities such as hay, grain, pack-
ing material, fruits, vegetables, wood, lumber and any crude plant
products must play a part in the spread of plant disease. With our
diversified trade relations with foreign countries and the extensive
trans-continental shipments of plant products from west to east
and from south to north, opportunities for the transport of plant
diseases over wide ranges of territory are greater than ever before
in entire progress of our agriculture.

Zoology Building, Univ. of Pa.

Philadelphia, Pa.

28

12.

13.

14.

15.

F. D. HEALD

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Futton, H. R.

Recent notes on the chestnut bark disease. Penn. Chest. Blight
Conference Rpt. 48-56. 1912.

Anperson, P. J., Erza, W. H. and Bascock, D. C.

Field studies on the dissemination and growth of the chestnut blight
fungus. Bull. Penns. Chestnut Blight Commission. 3:I-, 1913.

16.

17.

18.

19.

20.

2I.

22.

23.

24.

25.

26.

27.

28.

20.

30.

an

DISSEMINATION OF FUNGI 29

Luioyp, F. E. and Rineway, C. S.
Bull. Ala. Agr. Dept. 39:1-I9. IQII.
Burritt, J. T. and Barrett, T. J.
The ear rots of corn. Bull. Illinois Agr. Exp. Sta. 133 :65-109. 1908. .
SmitH, R. E.
The brown rot of the lemon. Bull. Cal. Agr. Exp. Sta: 190:1-70.
1907.
Rocers, C. S.
The late blight of celery. Bull. Cal. Agr. Exp. Sta. 208 :83-115.
Furiton, T. W.
The dispersal of the spores of fungi by the agency of insects with
special reference to the Phalloidei. Ann. Bot, 3 :1899-90.
Weseser, H. J.
Sooty mold of the orange and its treatment. Bull. Div. Veg. Phys.
and Path. U. S. Dept. of Agr. 13:1-34. 1897.
Waite, M. B.
Cause and prevention of pear blight. Yearbook, U. S. Dept. Agr.
1895 :295-300.
HEALp, F. D.
The bud rot of carnations. Bull. Nebr. Agr. Exp. Sta. 103 :1-24.
1908. :
Mourriti, W. A.
A serious chestnut disease. Jour. N. Y. Bot. Garden. 7 :143-153.
1906.
MasseEE, GEo.
Diseases of cultivated plants and trees. 1-602. 1910. Macmillan
& Co.
Jounston, J. R.
The history and cause of the cocoanut bud rot. Bull. Bur. Pl. Ind.
228 :1-175. I912.
VocLino, P.
Richerce intorno all’ azione delle lumache e dei rospi nello sviluppo
di Agaricini. Nuovo Giornale Botanico 27 :181-185. 1895.
Hays, W. M.
Rules and regulations for carrying out the plant quarantine act. U.
S. Dept. Agr. Cir. Office of Sec. 41:1-12. Ig12.
Stevens, F. L.
A serious lettuce disease. Bull. N. C. Agr. Exp. Sta. 217 :1-21. I91I.
Gussow, H. T.
A serious potato disease occurring in Newfoundland. Bull. Can.
Cent. Exp. Farm. 63:1-8. 1900.
SPAULDING, PERLEY.
The blister rust of the white pine. Bull. Bur. of Pl. Ind. 206:1-88.
IQII.

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THE PERIODICITY OF ALGAE IN ILLINOIS
By Epcar NELson TRANSEAU

The following notes on the periodicity of algal occurrence and
reproduction are based on a study of collections made at intervals,
at more than a hundred stations in East-central Illinois. They cover
the period from January, 1908, to January, 1913. About half of
these stations are in the vicinity of Charleston. These have been
visited at frequent intervals, while those at a distance have been
examined at critical times of the year as suggested by conditions at
Charleston.

All collections have been preserved in a solution, a liter of
which contains 100 cc. formalin, 300 cc. alcohol, and 600 cc. of
water. Each collection is labeled and numbered in the field and as
soon as convenient a 4 by 6 inch index card is labeled and num-
bered to correspond. On this is kept (1) a record of the weather
conditions, water conditions, temperatures, relative abundance of
algae in general, and whether floating or attached, etc.; (2) an
analysis of the collection made in the laboratory, showing all algae
present as far as identifiable. In the case of many only the genus
can be given, together with measurements that might aid in de-
termining them from later collections containing the same forms in
a fruiting condition. In summarizing the work it is possible then
to go back to cards or to the collections at any time and correct any
errors or get any further data desired.

“The waters of Eastern Illinois are rich in dissolved mineral
matters derived from the prairie soils. For example, the water of
the Embarras river near Charleston contains on the average .14042g.
of soluble matter per liter. It is consequently not surprising that
the algal flora should be large and varied. Its extent may be roughly
indicated by the fact that the collections contain more than forty-
five species of Oedogonium, and the genus Spirogyra is represented
by at least thirty-five species and varieties. These numbers are
considerably more than have been reported from Massachusetts

32 E. N. TRANSEAU

whose algae are better known probably than those of any state in
the Union.

In order to get at the periodicity the card records have been
listed by months for the five-year period. An examination of the
resulting chart shows that on the basis of their periods of greatest
abundance, the duration of their vegetative cycles, and the times of
their reproduction, the algae may be divided into seven classes.

I. Winter ANNuAtLS. These are species which begin their
vegetative cycle in the autumn, increase up to the time the ponds
are frozen, and last over the winter under the ice. During pro-
tracted winter thaws—which usually occur in January—they may
develop further and even fruit. Their period culminates in March
and April. Sexual reproduction may occur at any time from No-
vember to April. Zodspores are formed from the beginning through
the period of increase. Aplanospores and akinetes develop mostly
during the period of decline (Fig. 1). Some local examples of

NLL

Apr | May |June

Fig. 1. Frequency curve of winter annuals: In this and subsequent figures the
probable occurrence of sexual reproduction is indicated by F, zoospore reproduction by Z,
and the formation of aplanospores or akinetes by A.

algae belonging to this type are Vaucheria geminata, Vaucheria
sessilis, Draparnaldia plumosa, Tetraspora lubrica, and Stigeoclo-
nium lubricum varians.

II. Spring ANNUALS. These are forms in which the veg-
etative period begins in late autumn or early spring, culminates in
May and declines in June. Sexual reproduction occurs in April,
May, and June. Zodspores are formed mostly in early spring, and

PERIODICITY IN ALGAE 33

aplanospores and akinetes during the period of maximum abundance
and decline (Fig. 2). This type includes the largest number of

ie tae Se i eile

Fig. 2. Frequency curve of Spring Annuals

species, among which are Spirogyra varians, Spirogyra Weberi,
Zygnema stellinum, Oedogonium rufescens, and Ulothrix variabilis.

Ill. Summer Annuats. The vegetative period of the algae
of this class begins in early spring and culminates in July and
August. The decline is gradual and extends through the autumn
months. Sexual reproduction occurs in July, August and Sep-
tember. Zodspores, when formed, are most abundant in Spring
and early Summer. Aplanospores develop mostly in August and
September (Fig. 3). Among the local examples of this class are

Fig. 3. Frequency curve of Summer Annuals

Spirogyra decimina, Spirogyra maxima, Schizomeris Leibleinii,
Calothrix stagnalis, and Oedogonium V aucherit.

34 E. N. TRANSEAU

IV. Autumn Annuats. These species begin their vegetative
development in late spring, increase through the summer and have
their period of maximum abundance in the autumn. They may dis-
appear at the time of freezing up of the ponds, or gradually through
the winter. It has been noticed in at least one instance, Spirogyra
setiformis, that when the freezing occured at the time of fruiting,
a large part of the filaments still in a vegetative condition remained
over the winter and completed the fruiting in the early spring. This
is indicated in Fig. 4 by the dotted line. The sexual reproduction

Jan [Fes [ar Apr [may [Jane] July [Any [S000 [O08 [Pow [Dee

Fig. 4. Frequency curve of Autumn Annuals

usually occurs during September, October, and November. Among
the algae of this class are Spirogyra nitida, Rivularia natans, Oedo-
gonium crassum amplum, Spirogyra setiformis, and Oedogonium
obtruncatum.

V. PERENNIALS. This group includes forms in which the
vegetative cycle goes on from year to year without interruption.
The algae may become very scarce during unfavorable periods but
they are capable of at least maintaining themselves without the
production of reproductive bodies. They commonly attain their
greatest development during the summer and early autumn. Sex-
ual organs are mostly produced in late spring or early autumn—
sometimes in both. Zodspores are most abundant in spring and
summer,—not infrequently they are also produced in autumn (Fig.
5). Cladophora glomerata, Rhizoclonium hieroglyphicum, Pitho-
phora oedogonia, Pleurococcus vulgaris, and Ocdogontum grande
belong to this class.

PERIODICITY IN ALGAE 35

eae
22 TPaaeeaneanata| all

Fig. 5. Frequency curve of Perennials

VI. Epuemerats. These species have very short vegetative
cycles, usually best reckoned in days—at most in weeks. Genera-
tions succeed one another rapidly through the periods of favorable
conditions. Also because of varying capacities to respond to en-
vironmental conditions the generations overlap. It is therefore
difficult to represent this group by a curve or collection of curves.
Figure 6 is an attempt to represent it in general. It must be re-

KN)

/

Rh "
BPE aa Tine se Vinay | Jeselealy eg [end oes [oe neal

Fig. 6. Frequency curves of Ephemerals

SSS

membered that the lengths of the curve for a generation will vary
in the different species and under different conditions of temper-
ature, moisture and illumination. The species are mostly soil-
surface and plankton forms. Few of them reproduce sexually.
Zodspores, aplanospores and akinetes are the usual means of in-
crease, dissemination and passing an unfavorable period. The soil

36 E. N. TRANSEAU

forms are favored by wet weather. All the forms may be found in
greater or less abundance in all but the winter months. Among the
commoner local Ephemerals are Botrydium Walrothii, Scenedes-
mus quadricauda, Pediastrum Boryanum, Vaucheria terrestris, and
Ine ffigiata neglecta.

VII. Irrecuvars. In addition to the above six types of per-
iodicity there is a group of forms for which the combinations of
environmental conditions necessary to induce marked vegetative
development or bring about reproduction do not occur with seasonal
regularity. The period between maxima may be of more or less
than a year’s duration. Because of their uncertainty it is difficult
to discover them in a five year collecting period. There is always
the danger that they might have been overlooked during the first
year or two when my collections were not so thoroughly repre-
sentative as during the last three years. I will therefore venture
but a single example: Oscillatoria princeps, for which I have a
fairly satisfactory five-year record. This alga has been exceedingly
abundant at times in one of the ponds from which I have collec-
tions at short intervals. These periods of maximum development
have occurred without seeming reference to season—aside from
absence during the winter.

The diagrams for the several classes of periodicity show their
own relative abundance during the different months of the year.
The classes, however, do not represent equal parts of the algal
vegetation of this region. In Fig. 7 I have attempted to show the
relative importance of these several’classes. For reasons already
stated the Irregulars are not included. One reason why the group
of Spring Annuals is of such great importance is that the variety
of habitats at this season is vastly greater than at any other period
of the year. Habitat diversity and algal variety both have their
minimum during late July and August. Fig. 7 also indicates the
approximate composition of the algal flora at any time of the year.
Thus in October the Perennials and Autumn Annuals constitute the
bulk of the forms, with some remnants of the Summer Annuals,
some Ephemerals, and some early stages of the Winter Annuals.

With regard to periodicity in general it must not be forgotten,
that the classes observed in one locality do not necessarily occur in

PERIODICITY IN ALGAE 37

Jan €6 |Mar |Apr ay une ares Au ept reset Gens ov |Dec
[Yan _| Fe Thay | J Stag Haase fee oe

Fig. 7. Estimated relative importance of the several types of algal periodicity,
and the composition of the algal flora at any time of the year. This leaves out of ac-
count the irregulars.

all others. Indeed, I have good reason to believe that farther north
the number of classes is reduced and that two or more may be-
come merged into one. Judging by the publications of Fritsch, in
England many of the forms which occur here as winter annuals
attain their maximum development there in the summer. Hence
we may expect to find considerable local variations in the floristic
composition of the periodicity classes.

There is a notion prevalent in the text books and laboratories
that algae produce sex organs more abundantly during the low water
stages. This is sometimes expressed by saying that we may look
for sexually reproductive material when the water begins to con-
centrate, or “when the conditions become hard enough,” whatever
that may mean. Or it is said that they remain in a vegetative con-
dition so long as the waters are high. That the opposite of this
statement is true was strongly indicated by an examination of my
records at the end of my first two years of collecting. In the three
years that have since ensued I have watched this particular point
with much care, and there can be no question that in this region at
least (1) the greatest number of species fruit sexually, (2) a par-
ticular species fruits most abundantly, and (3) when a species pro-
duces more than one kind of spore, the greatest variety of spores

38 E. N. TRANSEAU

occur during periods of high water. The spring of 1912 was a
period of heavy rainfall. The remnants of old prairie ponds along
the railroad rights-of-way contained fruiting algae in quantity and
variety beyond anything seen during the preceding four years. The
high water-level was maintained until after the spores were mature.
On examining the corresponding collections for the spring of 1911
I find what are probably the vegetative filaments of many of these
same forms, but only a few produced spores. Again, the autumn
of 1911 was one of exceedingly high water,—the rainfall of Sep-
tember being more than three times normal. Coincidently a num-
ber of species which I had before found fruiting only in the spring,
developed and fruited before the pools froze. These algae fruited
again in the spring in some of the same pools. Whether the fila-
ments developed from the spores of the preceding autumn, or from
spring spores which failed to germinate in the autumn it is impos-
sible to say. But the fact of importance is that the continued high
water of the autumn of 1911 was attended by increased fruiting of
algae.

The origin of the prevailing notion that algae fruit during low
water stages may be connected with the fact that such a large num-
ber of algae fruit in late spring when the rainfall is decreasing and
the water levels are lowering. This is a coincidence of the lower-
ing water level with the time of fruiting and there is no causal re-
lation between the two. If the weather conditions are such that
the water level does not fall at this time of year, but remains con-
stant or rises the fruiting will not only take place but its amount
will be increased.

Fig. 8 shows the number of species known to have fruited ei-
ther sexually or asexually during each month for the years 1911 and
1912. The years are also divided into seasons and the total num-
ber of monthly records per season is indicated in the second line
from the bottom. In the last line is given the water level, as shown
by Weather Bureau records of rainfall, and my own notes. It
should further be noted that the seasons during these two years are
exactly opposite in so far as rainfall and water levels are concerned.
Leaving out of account the temperature conditions which at most
are of secondary importance, we have here a basis for direct com-

——————

PERIODICITY IN ALGAE 39

fet tS | SPS

olen RES sles ol at

hoa 44 ae ee Pie wae mas
eee

Fig. 8. Number of monthly fruiting records (both sexual and asexual) grouped by
seasons and compared with the water level. It will be noted that for each of the seasons
the conditions were opposite in these two years. Note correlation between high water
level and increased fruiting.

parison of wet and dry seasons. The close correlation between
high water level and increased fruiting is too obvious to need further
comment. The only record which shows a marked temperature in-
fluence is the one for the winter of 1912. There was not the usual
thaw in January and February, which would bring this number near-
er the winter mean.

Another notion prevails that sexual reproduction is confined to
the time of maximum vegetative development. While it is prob-
ably true of a majority of algae, I wish to call attention to the fact
that there are many exceptions. When a more detailed report of
these collections can be made, there is reason to believe a list of con-
siderable length will show that many algae may reproduce sexually
at any or all stages of their vegetative cycle. In other words vege-
tative development may follow sexual reproduction or may go along
with it, as well as precede it.

The failure of algae to fruit in streams has been mentioned by
numerous authors. Klebs and Oltmanns speak of the Vaucherias,
and Fritsch speaks of the Spirogyras. For eastern Illinois I can
record the fact that all the species of Vaucheria, Spirogyra and
Oedogonium known to grow in our streams have been collected at

40 E. N. TRANSEAU

one time or another in fruit. I do not, however, doubt the accuracy
of the European observations. Our streams possibly are more slug-
gish and perhaps there is a chemical difference of importance. The
flowing water is generally supposed to retard fruiting by furnish-
ing improved vegetative conditions. Hence we should expect a re-
tarding effect on fruiting which would show clearly in the relative
time of fruiting in ponds and streams. On going over the records
for species which have been collected in fruit in both situations |
find that the stream record is likely to be simultaneous with, or pre-
cede, or follow the pond record. As far as I have studied the
records there is no evidence of a retarding effect of running water.
There is abundant evidence to show that the number of possible
combinations of external factors that will produce sexual re-
production is very much less than the number that will induce
zooOspore production. Zodspores may be formed at short intervals
throughout the life cycle while the sex organs usually develop at a
definite time. The formation of non-motile spores in Pithophora
seemingly occurs under any and all conditions. This represents the
one extreme. At the other is Zygnema pectinatum which produced
aplanospores but once in the five years and then they were common
wherever the Zygnema was found.

It seemed best to the writer to await the completion of the ex-
amination of the collections before attempting to discuss the litera-
ture and the details connected with this problem of periodicity.
The names of the green algae used in this paper correspond to those
found in Collins’ “Green Algae of North America.”

Eastern Illinois Normal School,

Charleston, II.

>) es

SUMMARIES IN MICRO-BIOLOGY

For some months the Secretary has been planning to secure for this Journal and its
Department of Summaries, a series of papers from biologists dealing with the chief groups
of microscopic plants and animals. It has not been the purpose to present a complete
survey of any of the groups. The wish has been rather to bring together in one article
a statement of the following things:—general biology, the method of finding, the methods
of capture and of keeping alive and cultivating in the laboratory; how best to study; the
general technic; the most accessible literature; and a brief outline of the classification,
with keys for the identification of at least the more representative genera and species of
the micro-organisms likely to be found by the beginning students in the United States.

It has been felt that the getting together of such data as this, while not a contribution
to science, would be a contribution especially to isolated workers and to teachers and stu-
dents in the high schools and smaller colleges.

Papers have already appeared treating the aquatic Oligochetes and the Melan-
coniales. The following is the third paper of the series. It is proposed to have such
synopses from time to time until the more common American species of such groups as
the following have been covered: The Blue-green Algae, Conjugating Algae, Diatoms,
other Green Algae, Zygomycetes, Downy Mildews, Yeasts, Powdery Mildews, Hypho-
mycetes, Smuts, Rhizopods, Infusoria, Turbeliaria, Bryozoa, Water Mites, Entomostraca,
etc.—[ Editor. ]

THE NATURE AND CLASSIFICATION OF PLANT RUSTS
By Frank D. Kern

1. Introduction.

The rusts are small, mostly microscopic fungi, parasitic in the
tissues, especially the leaves, of the higher plants. They belong to
the order Uredinales (or Uredineae) which contains without doubt
the largest array of forms of any order of parasitic fungi. There
is an extensive economic interest in the rusts because of the fact
that they do great damage to most of the cultivated crops. Their
varied spore-formation makes them at once of unusual interest to
the general student with the microscope. Many species have spores
of five morphological sorts. In some species these occur in regular
succession upon one sort of host-plant but in many there is a strik-
ing change of hosts (known as heteroecism), a definite part of the
life-cycle being produced quite apart and dissociated from the other
part.

The spores are borne in more or less definite groups called sori
(rarely singly), covered at least at first by overlying host tissue and
set free either by early rupture or by weathering. On account of
the fact that the mycelium is always wholly buried within the host

42 F. D. KERN

tissues it is quite natural to fall into the habit of thinking and speak-
ing of the spore-structures, which manifest themselves upon the
surface, as if they constituted the whole plant instead of representing
only the reproductive portions. Although it is true that this treat-
ment will confine itself chiefly to the spore-structures yet it is well
to get the conception at the start that we must look upon the
mycelium with its resulting spore-forms, in its entirety, if we would
compare these little organisms with other and higher plants.

On account of the vast array of these forms, many of which
have never been investigated by anyone, it is manifestly impossible
to present much in the way of description of specific forms. It is
hoped, however, that a.statement of the main features of morphology
and life-history together with their application to classification may
serve to break down some of the apprehensive feelings which many
now entertain toward the group as a whole. It is with this object
in view that the following discussion is presented. The systematic
account is confined to genera and species found in the United States.

2. Habitat and Distribution.

The rusts are strictly parasitic upon ferns and flowering plants
and are liable to be found anywhere upon these hosts. Although the
spores are microscopic in size, when aggregated into sori they are
often conspicuous even to the naked eye and can usually be recog-
nized easily under a hand lens. The sori may appear upon any part
of the host above ground but the leaves are most commonly affected.
Presence of rust may often be indicated by yellow or discolored
spots upon the leaf-blades, or by swellings and galls upon the petioles
and stems, or by fasciations of the branches known as witches’
brooms.

In consistency the sori may be powdery (pulverulent), from the
falling away of the spores, or they may be compact and firm, or in
some species gelatinous. In shape and size there is great variation.
Often they are roundish or oval, about 0.2-1 mm. across and more
or less cushion-shaped (pulvinate); some are cup-shaped (cupu-
late), 0.1-0.4 mm. in diameter ; others project as cylindrical, filiform,
columnar, or wedge-shaped masses varying in length from 2 or 3
mm. up to 10 or 20 mm. In practically all cases the spore-mass is
elevated to some extent above the surface of the host tissue and by

EEE eee

PLANT RUSTS 43

this means alone it is often possible to distinguish in the field be-
tween true rusts and many spot-fungi which simulate rusts in gen-
eral appearance. This is especially true of grass and sedge rusts.
In color the various shades of yellow and brown predominate, but
some are so pale as to appear almost white, while many are dark
enough to be called black.

Rusts attack plants in all sorts of physical and climatic condi-
tions from the seashore to the summits of the highest mountains,
and from the tropics to the polar regions. There is scarcely a
family? of flowering plants in which some of the members are not
affected by these parasites. In most any region where there is vege-
tation some rusts can be found. In the fields on wheat, oats, and
other cereals; in the orchards on apples, pears, and quinces; in the
gardens on asparagus and beans; in the ornamental plantings on
roses, hawthorns, and cedars; in greenhouses on carnations and
chrysanthemums ; in the forest on pines, spruces, firs, oaks, cotton-
woods, and willows; in low places and swamps on sedges and crow-
foots ; in semi-arid regions on sage-brush and greasewood; in wild
and waste places everywhere on grasses, sunflowers, asters, golden-
rods, dandelions, and hundreds of other weeds and flowers.

Because a rust is known to live upon a certain host it does not
necessarily follow that the rust can be found wherever the host
grows. Wild roses have several species known on them; one of
these is found practically everywhere throughout the region of the
hosts, while the others seem restricted to certain geographic loca-
tions, for example one is in the northeastern states, another in the
prairie region of the central west, another in the Rocky Mountains
and so on. Some rusts which change hosts, as indicated in a fore-
going paragraph, might be expected to be limited to the region which
is common to both hosts, but the fact that many of these have the
capacity to maintain themselves independently on one host upsets
this expectancy. A notable illustration of this is the common stem-
rust of wheat which can have one stage on the barberry if any
bushes are in proximity but which flourishes equally well in regions
where the barberry is unknown.

1. The order Pandanales, of which the cat-tail (Typha) is our representative, and

the Palmales, the palms, are conspicuous examples of large alliances upon which no
Tusts are known.

44 F. D, KERN

It becomes evident from the foregoing discussion that in a study
of these fungi a study of the hosts is also not only important but
necessary. A knowledge of the hosts is essential in classification
and identification. A good way to begin is to examine and become
familiar with all of the forms of rust on some particular host or
closely related groups of hosts. In that way an interesting know-
ledge of flowering plants will be built up as the study proceeds
from one group to another.

3. Collecting.

In collecting one keeps the eye on practically all of the vege-
tation, looking especially for discolored spots and swollen (hyper-
trophied) areas but does not fail to take hold of and examine closely
many a plant which appears perfectly normal. A leaf or shoot
which is more upright than usual is always suspicious. A hand
lens is very useful and may assist greatly in forming a judgment as
to whether a rust is present. Until one becomes familiar with the
gross appearances of the various sori it is well to take home ques-
tionable material for microscopic examination.

It is always best to gather a fair amount of material. The im-
portance of gathering sufficient to give some clue as to the identity
of the host after the specimen has been preserved and packeted can-
not be over emphasized. Flowers or some portion of the infloresence
should be included whenever available, portions of the stem, un-
rusted leaves, basal leaves, etc., are advantageous. Care should al-
ways be taken to make sure that the rusted specimens and the por-
tions included for host identification are from the same plant, or
species, otherwise some very curious results may be obtained. Such
a warning may seem unnecessary but such things have happened
to experienced collectors. Some make it a point to gather separate
phanerogamic specimens for the host determination but such is not
necessary as a rule and is less convenient than the inclusion of
smaller diagnostic port‘ons of the host to be included with the fun-
gous specimens.

4. Care and Cultivation.

If specimens are desired for future study only they do not
require any special treatment but are best preserved by pressing

PLANT RUSTS 45

them in the ordinary method between some sort of absorbent driers.
If it is desired to keep the spores alive so that they may be germi-
nated and studied, or used for inoculating purposes, then certain
precautions are necessary.

In general we may divide the spores into two classes, active
and resting. The active class includes the cluster-cup spores, the
summer or red-rust spores, and certain others such as those of the
common cedar-apples. These spores are ready for germination
upon maturity and will lose the power to grow unless kept in a reas-
onably fresh condition. If it is desired to keep them alive the parts
of the host upon which they are growing should be kept as near
normal as possible. In the case of small herbaceous plants often
the best way is to remove them to pots, taking care to transfer a
sufficient ball of earth so that the shock of transplanting will be
reduced to a minimum. Oftentimes portions of the host-plant may
be kept fresh for a sufficient time by placing the cut ends of stems
or branches in water.

The winter or so-called black-rust condition of grass and sedge
rusts furnish fine examples of the resting class of spores. These
spores are produced in the late summer or fall and normally re-
tain their viability through the winter and germinate in the spring.
Collections made in the fall and kept in a warm dry room during
the winter usually fail to germinate. The freezing temperature of
the outdoor atmosphere is not detrimental. It is necessary to pre-
vent the specimens from thoroughly drying. If put up in cheese
cloth packets and tied to the branches of a shrub close to the ground
the spores will usually winter over well. Resting spores collected
in the field in the early spring usually show good germination. In
the spring the cloth packets should be brought into the laboratory
about the time conditions are favorable for growth outside. The
packets may be sprayed and after a few days of warmth and mois-
ture the spores should show signs of growth.

Germination can be nicely observed in a hanging drop culture.
Ordinary tap water is used for the hanging drop. If care is taken
to make the drop rather shallow it will be possible to focus with
the ordinary high power. The time required for germination de-
pends upon the conditions in which the material has been kept. The

46 F. D, KERN

germ-tubes may begin to show up in an hour or two. A drop cul-
ture which does not show germination in twenty-four hours may
as well be discarded.

If it is desired to make an inoculation indoors some small vig-
orous potted plants must be available. In case it is desired to carry
out such an experiment indoors for demonstration purposes it is
necessary to know the species with which one is dealing in order to
attempt the inoculation upon the right species of host or else the re-
sults would be very uncertain. For example there are about one
hundred species of rusts on grasses in North America. It is certain
that they all produce cluster cup stages on various broad-leaved
plants, but the life-histories of more than half of them are still un-
known. If one is conducting an investigation many trial inocula-
tions are attempted and some of them occasionally meet with suc-
cess, but for demonstration one must select forms which can be
expected to produce success. A few suggestions may not be out of
place here.

The black-rust spores from the stems of wheat will infect the
leaves of the barberry (Berberis). Young barberries may be grown
in pots from seeds. The grayish-black rust from the leaves of oats
will inoculate the buckthorn (Rhamnus), which may also be grown
easily from seeds. The sunflower rust does not change hosts and
the spores from the dark brown sori on wintered over leaves may
be transferred to young sunflower plants and will produce there
the cluster-cup stage. Spores from the common large cedar-apple
on the red cedar will produce abundant infection on the wild crab-
apple or the cultivated apple.

For indoor experiments the spores are removed from the
grasses with a knife or scapel blade and applied to the moistened
leaves of the trial host. In the case of the cedar rust it is not neces-
sary to apply the spores but merely to suspend the cedar-apple over
the plant. A moist surface and a saturated atmosphere are neces-
sary factors for the germination of the spores. In order to insure
these conditions the plant is sprayed with an atomizer before the
spores are sown, the parts which will not dampen being rubbed
with the fingers until water will adhere. After the sowing is made
the plant is placed under a bell-jar and set in a shaded position for

PLANT RUSTS 47

two or three days. The bell-jar is temporarily removed each day
to permit a change of air and is sprayed on the inside with an ato-
mizer before being replaced.

After an inoculation is made an incubation period of about a
week or ten days will elapse before infection will be evident by
the appearance of sori on the areas where the spores were sown.
This period must be taken into account if a teacher desires to have
a demonstration ready at some given time.

5. Methods for Study.

The spores of practically all species of rust make excellent ob-
jects for microscopic study by simply mounting them in a drop of
water on a slide and adding a cover glass, without any treatment
whatsoever. It makes little difference whether the spores are fresh
or whether they are from dried specimens they will as a rule make
a good mount in water. It is even possible to allow a slide to dry
out and then to run water under the cover glass and secure very
good results. Distilled water is preferable to tap water.

Sometimes when spores are quite old and dry they do not wet
up easily or appear somewhat shrunken. A good treatment in
such cases is the addition of a little lactic acid to the drop of water.
This will cause the spores to round out and take on a normal rotund
appearance without producing any appreciable swelling.

On account of the ease and satisfaction in making spore mounts
as described in the foregoing paragraphs it is rare that there is any
occasion for attempting permanent mounts of spores. For purposes
of studying the structure of the sori it is often desirable to have
sections and very beautiful results can be obtained by fixing fresh
material, embedding in paraffin and proceeding in the ordinary way,
no special precautions being necessary. It is possible, however,
to secure good preparations without resorting to the cytological
methods. With some practice many will find it possible to cut good
free-hand sections in pith. If the specimens are dry a small portion
containing the sori is soaked in very hot water—if the water comes
to a boil it will do no harm. Pith soaked in alcohol is preferable to
dry pith. The pith cylinder is partially split to allow the insertion
of the material and then, with some water on the razor blade to
float the sections, all is in readiness. The sections can be removed

48 F. D. KERN

with a needle or sharp wooden pick to a slide and are ready at
once for microscopic examination.

6. Characters that may be used in distinguishing the species.

In the classification and identification of the rusts there are
three features which are of importance (1) the microscopic char-
acter of the spores and sori, (2) the life-cycle, i. e. the number of
stages in development, so far as it can be made out, and (3) the
name and systematic position of the host. The first can be learned
from the microscope; the second cannot always be. made out, but
after a little practice helpful inferences may often be drawn; while
the third must depend upon the familiarity with the flowering
plants, the ability to work them out, or to secure competent aid.

AAHNOLO

a b
NI
oe
Sat
Cc d
Fig. 1. A teliospore in process of Fig. 2. Different types of free, stalked

germination. Two of the lower cells have teliospores: (a) 1-celled, the wall smooth

young promycelia, the uppermost cell has (Nigredo Polemonii); (b) 2-celled, the wall

a well advanced promycelium. This one smooth (Dicaeoma Grossulariae); (c) 3-

shows the division into four basidia, three ceed by oblique septa, the wall spinous;

of which are shown forming basidiospores. (d) several-celled by transverse septa, the
wall verrucose, the pedicel swollen (Phrag-
midium subcorticinum).

The rusts usually have more than one spore stage, the differ-

PLANT RUSTS 49

ent stages or phases appearing in a definite sequence and collective-
ly referred to as the life-cycle or life-history. Of the five morpho-
logical sorts of spores mentioned in a foregoing paragraph only four
are borne in sori on the host, the fifth being of a secondary nature
produced upon the germination of one of the other forms (see Fig.
1). This fifth sort, known as a basidiospore because it is produced
on a basidium, is important in indicating the relationship of the
rusts to other fungi but is of no importance in identfication. The
basidia themselves are of importance, especially as regards their
formation whether within or without the spore.

Of the four sorts of spores borne in sori only one is common
to all species, this one (together with the basidiospores) compris-
ing the full life-cycle in some species. This stage which is never
lacking in any life-history is known as the teliwm (plural telia)
and the spores as teliospores, sometimes called also teleutospores,
and represented by the symbol III. The teliospores may be 1-several-
celled (see Fig. 2), the wall may be smooth or rough but is not in
any known species set with prickles (echinulate). Upon germina-
tion the teliospores produce the secondary basidiospores, which
upon successful infection usually produce the pycnial stage, the
sori being known as pycnia or often as spermogonia. The pycnio-
spores are functionless so far as known and do not produce in-
fections, but the presence of this stage is often of value in de-
termining a life-cycle. Depending upon the life-cycle the rusts

Fig. 3. A vertical section through a portion of telium which shows a single layer
of spores compacted laterally. The sorus is subepidermal and the flattened epidermis is
shown extending over the spores. The species represented is Melampsora Medusae on
Salix.
may be divided into two groups, one with a short cycle and the other
with a long cycle. In the short-cycle forms the mycelium from a
basidiospore produces pycnia which are followed at once by telio-
spores or sometimes there is a suppression of the pycnia. The

50 F. D. KERN

pycnia usually appear as honey-yellow specks at first, often becom-
ing blackish with age. The stage is often designated by the
symbol O.

The long-cycle forms have the pycnia and telia and in addition
have between the two either aecia or uredinia or both produced, in
the order named. These two additional stages form important
parts of the life-cycle.

The aecial stage is the so-called cluster-cup stage, deriving that
name from the fact that each aecium, or aecidium, is in many species
provided with a covering (peridium) which later opens out into a
cup-like receptacle enclosing a mass of spores. The edge of this
peridium often becomes toothed or fringed. In some forms the
peridium becomes long and cylindrical while in others it is entirely
lacking. Sometimes the aecia are encircled by clavate or capitate
structures known as paraphyses (see Fig. 4, b). The aeciospores
are usually borne in chains, are always 1-celled, the wall is rough-
ened with more or less evident roundish warts (verrucose), and is
in many species colorless. The symbol for the stage is I.

a b

Fig. 4. (a) Showing the surface sculp-
turing on the side wall (longitudinal radial)
of a peridial cell of Gymnosporangium
globosum. The different species differ in
the character of the markings. When in
place in the peridial tissue other cells are

a b

joined end to end and side by side. (b)
Showing the general nature of a paraphysis.
These structures surround the spore groups
in some species and in others may be inter-
mixed with the spores.

Fig. 5. Two types of urediniospores:
(a) an ellipsoid spore with echinulate walls
and four equatorial germ-pores (Dicaeoma
poculiforme); (b) a globoid spore with ver-
rucose walls and six scattered pores.

The uredinial stage is the one often popularly referred to as

the red rust stage.

In most genera the wrediniospores (see Fig 5),

or uredospores, are borne singly on pedicels in naked sori, but in
some they are in chains and may be surrounded by peridia or by

paraphyses.

The walls of the spores are usually colored and are

PLANT RUSTS 51

always rough, either echinulate or verrucose. The spores afe
single-celled. Functionally these spores are repeating spores, i. e.
they may reproduce themselves over and over indefinitely. The
symbol for this stage is II.

The microscopic spore-characters most used are shape and size,
surface markings, color and thickness of walls. In the case of
teliospores the number of cells is important as is also the shape, size,
and color of the pedicel which often remains attached. With regard
to the urediniospores the number and location of the germ-pores
are often of great value. These pores appear as lighter circular
areas about I-1.54 in diameter and as they are the places through
which the germ-tubes penetrate they are called germ-pores. The
lactic acid treatment will usually assist in bringing them out more
clearly than water alone. In some groups the characters of the
peridial cells must be observed, especially the surface markings
(see Fig. 4,a). The fine and varied character of the surface sculp-
turing on some of these cells almost makes them rank with diatoms
as objects of microscopic interest.

7. Topics for Investigation Suitable to the General Student of
the Group.

The rusts form an interesting group in which much remains
to be done in the United States. One of the most fascinating and
at the same time profitable opportunities for botanical students
everywhere is to institute a careful study of heteroecious forms.
Heteroecious species are divided into two wholly unlike halves
and actual culture (inoculation) experiments are necessary to prove
a relationship. In order that the work of connecting the halves
may go on expeditiously, with as little unprofitable labor as possible,
it is essential that the experimenter be guided by some ideas of
probable relationships. These ideas can be gained in the field. The
finding of aecial and telial stages in close proximity in the field is,
to be sure, not proof of their affinity but is a bit of prima facie
evidence. The closeness of the association, the abundance of the
infection, and the occurrence of known forms must all be taken
into account. Observations can best be begun in the early spring
when new growth is starting. To find a tuft of grass or sedge cov-
ered with wintered over teliospores in contact with some new shoots

52 F. D, KERN

of a broad leaved plant bearing aecia is a strong suggestion of
genetic relationship. Since in North America there are about one
hundred aecial forms whose relations to telial forms are unknown
it will be recognized that there is abundant opportunity for field
observations. The problem may be stated from the other view-
point by saying that there are scores of telial forms whose rela-
tions to aecia must exist but are unknown.

Some observers without greenhouse facilities may like to verify
their clues by means of actual cultures. It is often possible to
obtain very satisfactory results by means of outdoor cultures in a
garden or other protected place. Much valuable work has been
done in this way. For example, plants known to bear aecia can be
_ transplanted to the garden and cared for until they establish them-
selves. During the winter rust on grass suspected of being related
can be placed over the ground so that the young shoots will have
to push up through it. In this way results may be obtained early
before the danger of stray infections is so great. If more than one
such experiment is tried in the same garden much care must be
observed to prevent cross infections which might lead to confusion.

8. Systematic (General).

Teliospores compacted laterally into flattened, cushion-like
masses (see Fig. 3), or filiform, columnar masses (rarely
solitary within the tissues), without stalks.

Walls of teliospores gelatinous, especially at apex, dividing
internally into four basidia. Family 1. Coleosporiaceae.
Walls of teliospores firm, without internal division of con-
tents. Family 2. Uredinaceae.

Teliospores free (see Fig. 2) or united in bundles, stalked, the
walls firm, or with an outer hygroscopic layer.

Family 3. Aecidiaceae.

Some authors include the first two families in one under the

name Melampsoraceae, and use the name Pucciniaceae for the third
instead of Aecidiaceae as given above.

9. Systematic (Special).
FAMILY 1. COLEOSPORIACEAE
This family contains only one genus of importance, Coleospor-

PLANT RUSTS 53

ium, with about 24 species. The genus has all four spore-stages.
All the species are heteroecious, the aecia being the blister rusts on
the leaves (not on the twigs or bark) of pine trees (Pinus). The
uredinia are yellowish and powdery; the telia form waxy cushions ;
the teliospores germinate upon maturity in the fall and the in-
ternal division of the contents into four basidia can generally be
observed with the microscope without difficulty. The following
common species may be mentioned.

Coleosporium Ipomoae (Schw.) Burr. on Ipomoea, urediniospores with
uniformly thin wall 1-1;54; C. Campanulae (Pers.) Lev. on Campanula,
urediniospores with uniformly thick wall, 2-3.54; C. Vernoniae B. & C. on
Vernonia, urediniospores with wall 1-2 at sides, often 2-5u above; C. Soli-
daginis (Schw.) Thiim. on Aster, Euthamia, and Solidago, urediniospores
with uniform wall, about 1-2u. The urediniospores of the different species
do not vary much in size, averaging 14-22x20-30n.

FAMILY 2. UREDINACEAE

This family is represented in North America by seventeen or
eighteen genera and a considerable number of species. In the
United States only seven of these genera are common, the others
being chiefly from tropical regions.

Key To THE PRINCIPAL GENERA
Teliospores in definite and limited sori, usually on the leaf-blades; uredi-
niospores rounded.

Telia conspicuous, raising or breaking through the epidermis, telio-

spores I-celled.
Telia in the form of cushion-like masses; urediniospore-wall

verrucose.
Teliospores in a single layer; urediniospores with inter-
mixed .pataphiyses. gosh dk. dias: Genus Melampsora
Teliospores in chains; uredinia with a delicate peridium
or ‘naked. . 0s wsaverses dees testes Genus Melampsoropsis
Telia extruding as long filiform columns; urediniospore-wall
echintlate. : 25.2252. :20 heen aeeees as Genus Cronartium

Telia inconspicuous, in a layer in the epidermal cells or just below
them, teliospores 2-4-celled.

Teliospore-wall brownish; uredinial peridium opening with a
definite orifice surrounded by longer cells, urediniospores
eehtuilate 204 5:5 /ssd Je seae taken seme Genus Pucciniastrum

Teliospore-wall colorless; uredinial peridium without definite
orifice, the cells longer at the sides and shorter toward apex,
urediniospores verrucose................ Genus Hyalopsora

54 F. D. KERN

Teliospores solitary, or in very loose groups, usually buried within the

parenchymal tissues; urediniospores pointed.......... Genus Uredinopsis
Teliospores forming continuous layers around elongated and thickened stems,
not erumpent; uredinial stage lacking............... Genus Calyptospora

GENUS MELAMPSORA CAST.

A prior name for this genus is Uredo. But as that word has been in general use
as the name of a stage, the one called in this paper the uredinial stage, and its restriction
to a true generic application might lead to confusion, a later and more commonly used
name is here maintained.

The genus contains both heteroecious and autoecious species.
The aecia have no peridium.?. A conspicuous feature of the uredinia
are the numerous, large paraphyses. Both aeciospores and uredinio-
spores have colorless, verrucose walls. There are three common
species.

Species

M. Medusae Thiim. Urediniospores smooth on two sides
which are thickened, I on Larix, II and III on Populus.

M. Bigelowii Thiim. Urediniospores with walls evenly thick
and evenly verrucose, I also on Larix, very similar to the preceding,
II and III on Salix (see Fig. 3).

M. Lint (Schum.) Desm. Autoecious, on Linum.

Genus MELAMpsoropsis (SCHROT.) ARTH. (Sometimes included in
CHRYSOMYXA).

Found in its uredinial and telial stages only on the order
Ericales. The aecia so far as known occur on the leaves or cones
of spruces (Picea). Several of the species are rather rare. M.
Pyrolea (DC.) Arth. on wintergreen (Pyrola) is common. There
are two species on Labrador tea (Ledum) ; M. ledicola (Peck) Arth.
with the II and III on the upper side of the leaves, urediniospores
moderately large, 18-29 x26-36p, wall 2.5-3n thick; and WM. abietina
(A. & S.) Arth. with sori on the under surfaces of the leaves, ure-
diniospores moderately small, 14-22 x20-30p, wall 1.5-2.5 thick.
A uredinial stage on Cassandra calyculata, which is very rarely ac-
companied by telia, is M. Cassandrae (P. & C.) Arth.

GENuS CRONARTIUM FRIES.
A very striking genus in the telial stage on account of the long
(0.5-3 mm.) filiform spore-columns. Cultures have proven that

2. The term caeoma is often applied to such forms, i. e. to aecia in which the
peridium is lacking.

ee

—

.
}
:

PLANT RUSTS 55

the aecial stages are the blister rusts of the twigs, branches and
trunks of pines (Pinus).
Species

C. Comptoniae Arth. A common form along the north At-
lantic coast on the sweet gale (Myrica Gale) and sweet fern (M.
asplenifolia).

C. Quercuus (Brond.) Schrot. is widely distributed on various
species of oak (Quercus). The aecial stage (called Peridermium
cerebrum Peck) on pines forms globoid swellings of the branches
upon which the orange-yellow aecia are arranged in a cerebroid
fashion.

C. ribicola Fisch. de Waldh. is a rather recent importation from
Europe and is a very serious disease of white pine (Pinus strobus)
seedlings. The telial stage on Ribes (currants) is also appearing
in this country.

GENUS PUCCINIASTRUM OTTH.

The characteristic feature of this genus is the hemispherical
or subconical peridium of the uredinial stages with a pore-like
orifice at apex surrounded by elongated cells, which are often
echinulate above. Owing to the fact that the telia remain covered
(indehiscent) they are somewhat difficult to study, and the parti-
tions of the teliospores being vertical are not readily made out. Of
the nine or ten species the following are the more common ones,
others may be found upon Hydrangea, Rubus, Arctostaphylos, and
Vaccimum.

Species

P. Agrimomiae (Schw.) Tranz. Common on Agrimonia from
New England to North Dakota southward to Florida and Mexico.

P. pustulatum (Pers.) Diet. Widely distributed, especially
northward on various species of Epilobium.

P. Pyrolae (Pers.) Diet. on Pyrola and Chimaphila, can be dis-
tinguished from the Melampsoropsis on Pyrola by the nature of the
uredinial peridium and the echinulate markings of the uredinio-
spores.

GENERA Hyatopsora Maen. and UREDINIopsis Macn.

These genera include all of the rusts which are known on ferns in America. The
cycle of development in both genera is not well understood. Both have two spore-forms

56 F. D. KERN

.

known on the fern-hosts aside from the telia. Some authors have looked upon one of
these forms as aecia and the other as uredinia but evidence is lacking to prove the
correctness of this assumption and recent work® indicating the heteroecious character of
certain species of Uredinopsis throws some doubt upon that disposition. For the most
part the two genera occur upon different genera of ferns; Hyalopsora on Phegopteris,
Cystopteris, Polypodium, and Pellaea; Uredinopsis on Osmunda, Onoclea, Pteridium,
Asplenium, and Dryopteris. The two genera can be further separated by the fact that
one of the spore-forms of Uredinopsis has fusiform spores which are acute or beaked
above, with a wall which is smooth except for two longitudinal ridges bearing single rows
of minute projections, while both spore-forms in Hyalopsora have rounded spores with
evenly verrucose walls. H. Aspidiotus (Peck) Magn. is the most widely distributed
of the four species belonging to that genus; U. Osmundae Magn. on Osmunda, U.
mirabilis (Peck) Magn. on Onoclea, and U. Atkinsonii Magn. on Asplenium and Dryop-
teris are the best known of the seven described species of Uredinopsis.

GENus CALytTosporA Kuhn.

Only one species is at present recognized in this genus, C.
columnaris (A. & S.) Kuhn (C. Goeppertiana Kiithn). Uredinia
are lacking; the telia are found on Vaccinium, and the aecia on
the balsam fir (Abies balsameum) in this.country. The telia form
an even, polished, reddish-brown layer around the elongated and
enlarged stems; the teliospores are closely packed in the epidermal
cells, the wall of each spore very thin at the sides 0.5-0.8u, some-
what thicker above I-I.5y.

FAMILY 3. AECIDIACEAE
(Called also PUCCINIACEAE)

In this family belong the largest number of rusts, including
for the most part those that cause serious injury to economic plants.
The number of genera to be dealt with is dependent upon the
scheme of classification which one follows. According to the old
method any species of the group having free teliospores would be-
long to the genus Puccinia if it possessed a single other character,
i. e. two-celled teliospores (see Fig. 2, b). Likewise those forms
would belong to Uromyces, which possess one-celled teliospores
(see Fig. 2,a). Such a scheme, based on only one character, brought
together, as a genus, species of the most diverse forms and varied
affinities. A classification which takes into consideration the nature
of the spore-wall, germ-pores, the origin of the sorus, i. e. whether
under the cuticle or under the epidermis, the life-cycle, whether
one or more stages are lacking, and other important characters
will, of course, segregate the species usually placed under Puccinia

3. Fraser, W. P. Science, N.S. 36:595. 1912.

PLANT RUSTS 57

and Uromyces and increase the number of genera, but it will have
the very great advantage of forming groups which have some
affinities. Following such a system we have among the more com-
mon forms in the United States about twelve genera to consider
in the place of two, but this number might be decreased nearly
one-half by not recognizing the purely artificial character of num-
ber of cells as a basis for generic separation. The present state of
knowledge does not seem sufficient, however, to warrant such a
change.
Key To THE PrINcIPAL GENERA
Teliospores or pedicels, or both, more or less united; uredinia when present
naked but often with intermixed paraphyses.
Teliospores united into a head, or cushion-like body, on a compound

PECL O I eas SSete Sate ito Pe roe ea eS Ur ate Howto a as Genus Ravenelia
Teliospores free but borne in groups of two to eight on a common stalk.
Life-cycle with all spore-forms.............. Genus Tranzschelia
Life-cycle with pycnia and telia................ Genus Polythelis

Teliospores and pedicels both free; uredinia when present without peridium
but sometimes with encircling paraphyses.

Teliospores becoming imbedded in masses of jelly formed by gelatin-
ization of the pedicels, teliospore-pores varying in number and
arrangement; uredinia lacking........... Genus Gymnosporangium

Teliospores in definite sori, not becoming gelatinous.

Pycnia subcuticular, other sori subepidermal; teliospore-pores
when more than one in a cell lateral; uredinia usually with
encircling paraphyses.

Teliospore-wall more or less conspicuously laminate.
Teliospores 2-celled, the wall finely and sparsely
VErrhense Pees tes ess ke. 5 3 Be Genus Uropyzxis
Teliospores 2 to several-celled, more or less coarse-
ly verrucose or even smooth.
Life-cycle with all spore-forms..............
aia hil aia eS OEY cc Sat Genus Phragmidium
Life-cycle with pycnia, aecia and telia........
Be Abe. ee Genus Earlea
Teliospore-wall not noticeably laminate.
Teliospore-wall spinous, teliospores 3-celled by
oblique septa................ Genus Nyssopsora
Teliospore-wall nearly or quite smooth, the spores 2-
or several-celled by transverse septa.
Teliospores 2-celled....... Genus Gymnoconia
Teliospores 3-13-celled...... Genus Kuehneola

58 F. D, KERN

Pycnia and other sori subepidermal; teliospore-pores one in a
cell and apical; uredinia rarely with encircling paraphyses.
Life-cycle with all spore-forms.

Teliospores I-celled ............... Genus Nigredo

Teliospores 2-celled .............. Genus Dicaeoma
Life-cycle with pycnia, aecia and telia.

Teliospores I-celled ........... Genus Uromycopsis

Dehospores: 2-Celled:: .... « » s:0s\s was avon Genus Allodus
Life-cycle with pycnia, uredinia and telia.

Teliospores 1-celled .............. Genus Klebahnia

Deliospores | 2-celled ooo o was ece se Genus Bullaria
Life-cycle with pycnia and telia, or only telia.

Delospores. I-celled> 266. sss aesc Genus Telospora

Teliospores 2-celled .............. Genus Dasyspora

GENUS RAVENELIA BERK.

This genus is especially characterized by the manner in which the teliospores are
fascicled on compound pedicels. The spores form heads which are bordered by hyaline
cysts that swell more or less in water. The urediniospores are often paler below. The
genus occurs, with the exception of one species, upon leguminous hosts included in the
families, Mimosaceae, Caesalpiniaceae, and Fabaceae. The exception is on Phyllanthus
belonging to the Euphorbiaceae. Thirty-eight species have been described in North
America, chiefly from Mexico, Central America, and the West Indies. Several occur
along the southern border of the United States but only one comes into the central
and northern states, R. epiphylla (Schw.) Diet. on Cracca (Tephrosia).

GENUS TRANZSCHELIA ARTH.

A small genus, only two species at present known. The uredin-
iospores have the wall thicker and less echinulate above. The tel-
iospores are 2-celled and a characteristic feature about them, aside
from the manner in which they are borne, is the ease with which
the two cells separate. One species, T. cohaesa (Long) Arth.,
known only from Texas, is autoecious on Anemone decapetala; the
other, 7. punctata (Pers.) Arth., is widespread and heteroecious,
O and I on Anemone, Hepatica, and Thalictrum, II and III on

peaches, cherries, and plums.

GeNuS PoLyTHELIS ARTH.

A small genus which is confined to hosts of the family Ranun-
culaceae. The teliospores are very similar to those of Tranzschelia
but the two genera differ very markedly in the life-cycle. A species
having both cells of the teliospores globoid is P. fusca (Pers.) Arth.
on Anemone quinquefolia common east of the Mississippi; another
with the lower cell considerably elongate, on Pulsatilla hirsutissima,
is P. Pulsatillae (Rostr.) Arth. common from the Mississippi to

—ey-” ~-

PLANT RUSTS 59

Colorado and Montana; and a third with the lower cell somewhat
elongate, on Thalictrum, is P. Thalictri (Chev.) Arth., distributed
throughout the northern United States and Canada.

GENUS GYMNOSPORANGIUM HeEbw. F.

Characterized, with a few exceptions, by a dingy-white, mem-
branous peridium, which elongates into a tubular form and tends to
rupture along the sides; by large peridial cells usually conspicu-
ously sculptured on the inner and side walls (see Fig. 4, a); by
aeciospores with colored walls and evident germ-pores*; and by
teliospores with hyaline pedicels of considerable length, the outer
portions of which swell in moisture and become gelatinized to form
a jelly-like matrix in which the spores appear imbedded. As regards
hosts the genus is restricted in its aecial stage to the family Malaceae
(Pomaceae), with three known exceptions, and in its telial stages
to the Juniperaceae without any known exceptions. About thirty
species have been reported in the United States, of which the fol-
lowing are most likely to be collected.

Species

G. Juniperi-virginianae Schw. (G. macropus Link). The com-
mon “orchard rust” forming globoid galls on the Virginia red cedar
in the telial stage and attacking crabapples and cultivated apples in
the aecial stage. The telia on the galls are cylindrical, the galls die
after producing a crop of telia.

G. globosum Farl. Also forming telia on the red cedar but
chiefly on the genus Crataegus in its aecial stage. The telia are
wedge-shaped and the mycelium in the galls is perennial, produc-
ing new telia between the scars of the sori of previous seasons.

G. germinale (Schw.) Kern (G. clavipes C. & P.). The hem-
ispherical telia in this species do not form galls but long gradual
enlargements of the twigs or branches. The aecia attack the fruits
and often the twigs of Cydonia (quince), Amelanchier, Aronia, and
Crataegus. The peridium is unusually whitish. The telia occur
not only on the red cedar (Juniperus virginiana) but also on the
junipers (Juniperus communis and J. siberica).

Along the Atlantic coast are two conspicuous species on the branches of

4. In most genera germ-pores are apparently wanting or obscure in the aeciospores
but are usually evident in the urediniospores and teliospores.

60 F. D. KERN

the white cedar (Chamaecyparis thyoides); G. Ellisii (Berk.) Farl. with
yellowish filiform telia and G. Botryapites (Schw.) Kern with brownish
pulvinate sori. G. Betheli Kern is a gall form very destructive to the red
cedar (J. scopulorum) in the Rocky Mountains; G. guvenescens Kern in the
same region causes witches’ brooms on the cedars.

GeNuS Uropyxis SCHROT.

A genus usually separable from all others here described by the
laminate wall of the teliospores, the outer layer of which is gelatin-
ous, swelling in water. The species are more common southward
into Mexico. U. sanguinea (Peck) Arth. on Mahomia (Berberis)
is distributed throughout the western mountain region from Wash-
ington and Wyoming south to Guatemala. U. Amorphae (Curt.)
Schrot. on Amorpha is widely distributed over the United States
and especially abundant in the Mississippi valley. In the former
the gelatinous outer layer is relatively inconspicuous, in the latter
1-3u thick at apex and base of spores and 7-15y at the sides.

GeNus PHRAGMIDIUM LINK.

The cycle of development in this genus includes all spore-
forms and all species are autoecious. For hosts it is restricted to
a single family, the Rosaceae. The aecia and uredinia are both
without peridium but usually with encircling paraphyses (see Fig.
4, b). The teliospores are usually more than two-celled by trans-
verse septa (Fig. 2, d). Sixteen species have been described in
North America, four on the tribe Rubeae, eight on the tribe Roseae,
and four on the Potentilleae. P. imitans Arth. on Rubus strigosus
is the most widely distributed of the first group. P. disciflorum
(Tode) James and P. subcorticinum (Schrank) Winter are com-
mon on cultivated roses in many parts of the United States, espe-
cially the northern states east of the Rocky Mountains. The tel-
iospores of the former are 5-9-celled, with walls blackish-brown,
opaque, 5-7p thick, of the latter 5-7-celled, the walls chestnut-brown,
not very opaque, 3-5 thick; in both species the teliospore-walls are
verrucose and the pedicels swell in water. P. Andersom Shear on
Dasiphora fruticosa and P. Potentillae (Pers.) Karst. on various
species of Potentilla are representatives of the third group. In P.
Andersoni the teliospores are furnished with a hyaline papilla at
the apex and the pedicel is much swollen in the lower part, while
in the other the apex has no apiculus and the pedicel is not swollen.

PLANT RUSTS 61

Genus EarLEA ARTH.

This genus resembles Phragmidium but the cycle of develop-
ment includes only pycnia, aecia, and telia. Several species have
been referred here but only one of them is common, that on various
roses. This species differs from the species of Phragmidiuwm com-
mon on roses by the teliospores having smooth walls and pedicels
not swelling in water and also by the fact that the telia are large
and appear always upon the stems, while in that genus they are
small and only upon the leaves.

Genus NyssopsoraA ARTH.

The teliospores differ from those of all other genera (except
Triphragmium) in having the teliospores divided into cells by
oblique partitions in such a way as to make them triangularly 3-
celled (Fig. 2c). They differ from Triphragmium, which is not
discussed in this paper, by the short life-cycle and the spinous char-
acter of the teliospore walls. Only one species, N. clavellosa
(Berk.) Arth. on Aralia nudicaulis is known east of the Rocky
Mountains; another in the western mountainous region is N.
echinata (Lev.) Arth. on Ligusticum and Oenanthe.

GENUS GYMNOCONIA LAGERH.

Here belongs the orange rust of blackberries and raspberries
(Rubus spp.) which is so well known. It is the only species of im-
portance and is best known under the name G. interstitialis
(Schlecht.) Lagerh. The pycnia and aecia are the conspicuous
stages; no uredinial stage exists.

Genus KuEHNEOLA (LINK) ARTH.

Another genus with several species on the Rosaceae but differ-
ing from those already described on that family. The teliospores
are smooth and few- to many-celled by transverse partitions. The
aecial stage is lacking. K. obtusa (Strauss) Arth. with 3-5-celled
teliospores is a common form on Potentilla canadensis; K. uredinis
(Link) Arth. with 5-13-celled (usually 5-6) teliospores is another
rust of Rubus, but is not at all conspicuous and can not be confused
with Gymnoconia. One species, K. Gossypti (Lagerh.) Arth., is a
rust of the cotton plant known from southern Florida and the West
Indies.

62 F. D. KERN

Genus NicrEDo Rouss.

To this and the following seven genera belong most of the
species formerly referred to the old composite genera Uromyces
and Puccinia. By the use of the generic names here adopted the
important information concerning the life-cycle is conveyed in the
name without the necessity of the roundabout method of explaining
the status with a phrase.

The aecia are usualy cupulate, aeciospores borne in chains
with colorless, verrucose walls; the uredinia are without peridium
or encircling paraphyses, urediniospores borne singly on pedicels,
the walls colored, echinulate or verrucose, the pores variously ar-
ranged; the telia are sometimes long covered by the epidermis,
teliospores free, stalked, 1-celled (see Fig. 2a), the wall firm,
colored, smooth or verrucose, with one apical pore. The genus
Nigredo is represented by a large number of species, many of which
are common in the United States. It will be possible to mention
only a few of those most likely to be found.

Species

Host belonging to grass family (Poaceae).
Urediniospore-pores 3 or 4, equatorial, the spores medium-sized (15-19
x18-23u) ; on species of Panicum, chiefly P. virgatum; I unknown.
OER WE eB nin. 235s cared ue Sas kee N. graminicola (Burr.) Arth.

Urediniospore-pores about 8, scattered, the spores large (19-27x25-37p) ;
on species of Spartina; I on Steironema, Polemonium, Phlox, and
SS TE eI NS ous 0.5' 6, senile ok fe, Foe es N. Polemonii (Peck) Arth.

Host belonging to sedge family (Cyperaceae).
Urediniospore-pores 4, equatorial; on Scirpus; I on Cicuta and Sium
OR aac A ETE ROS o's co orc biecehij os Biante sauce N. Scirpi (Cast.) Arth.

Urediniospore-pores 2, above the equator; on Carex; I on Aster and

SOUIEGO cee TRIES <=) o.0.ereidte, came tore tine crete N. perigynia (Hals.) Arth.

Host belonging to’family Araceae; urediniospore-wall thicker above, pores 4;
on Caladim > atitoectottse. .,.. 22250 blo Seeeene N. Caladii (Schw.) Arth.
Host belonging to family Juncaceae; urediniospore-pores 2, equatorial; on
Juncus; I on Ambrosia, Arnica, and Cirsium...... N. Junci (Desm.) Arth.
Host belonging to family Polygonaceae; urediniospore-pores 4, equatorial;
on Polygonum, autoecious.........5.esecceccece N. Polygoni (Pers.) Arth.

Host belonging to family Carophyllaceae; urediniospore-pores 3 or 4, equa-
torial; on Dianthus (carnation); I on Euphorbia (not known in United
BSEALES ) chs Rie AEE ELAR oe Se ee ey ae N. caryophyllina (Schrank) Arth.

—

indie,

kt ee eg

PLANT RUSTS 63

Host belonging to family Fabaceae.
Urediniospore-pores 3-6, scattered; on Trifolium pratense (red clover) ;

DE eeakeiowal cgay. bee cee tes a c's s N. fallens (Desm.) Arth.
Urediniospore-pores 3 or 4, equatorial; on T. repens (white clover;
AULOCCIOUISAS eee Rie ee sate ede lela N. Trifolii (Hedw. f.) Arth.
Urediniospore-pores 2, equatorial; on Strophostyles, Vigna, and Phaseolus
(including “the garden :beafi); atitoecious...... 0's. i006 oe oc ee ences

FRA ony cone eo Once bene Oc re Ore N. appendiculata (Pers.) Arth.

Host belonging to family Asclepiadaceae; urediniospore-pores 4, equatorial ;
on Asclepias; DiGMRROWH +e os ic. wl. ie eae. N. Howei (Peck) Arth.

GENuS DIcAEOMA S. F. GRaAy.

This genus resembles Nigredo in every important character,
differing only in having teliospores with two cells. It is without
doubt the largest of the rust genera. Here belong the bulk of
grass and sedge rusts, including the important cereal rusts.
Dicotyledonous plants of eighty or ninety genera representing about
twenty-five families serve as hosts for species of this genus, but
most of these are not of economic interest or of common occurrence.

Species

Host belonging to grass family (Poaceae).

Telia early naked, blackish, chiefly on the culms and sheaths; uredinio-
spore-pores 4, equatorial; on wheat, oats, rye, timothy and several wild
grasses (Agrostis, Agropyron, Elymus) ; I on barberry...............
PEAS Paige ee ee Sik, SN ee eT SORT Siete el D. poculiforme (Jacq.) Kuntze

(=Puccinia graminis Pers.)

Telia long covered by the epidermis, often grayish-black; chiefly on the
leaf-blades.

Urediniospore-wall brown; the pores about 6, scattered.

Urediniospores with intermixed paraphyses; telia rarely formed
in our region; on species of Poa, common on blue-grass; I on
Uitssulago! Lave: 5. .1srspee eter D. epiphyllum (L.) Kuntze

(=Puccinia poarum Niels.)

Urediniospores without paraphyses; teliospores germinating in
the fall; on rye (Secale cereale) ; I on Lycopsis, not yet found
#n CAmerGas 26 :2.4)s. siete Sages D. Asperifolii (Pers.) Kuntze

(=Puccinia rubigo-vera DC.)
Urediniospore-wall yellow or colorless.

Teliospores with finger-like projections at the apex; on oats and
wild grasses (Cinna, Holcus and others); I on buckthorn
CRHOINNWS eS cia avaie it's atoeie sop tore stats D. Rhamni (Pers.) Kuntze

(=Puccinia coronata Cda.)

64 F. D. KERN

Teliospores with smooth apex; on wheat; I unknown..........
sminstcme MLR: os 4 viele oa'Sb ad D. triticina (Erikss.) Kern.
(=Puccinia triticina Erikss.)
Host belonging to sedge family (Cyperaceae).
Urediniospore-pores 3 (in occasional spores 4), equatorial.
Urediniospores large (18-26x24-39u) ; teliospores large (39-71u long) ;
on species of Carex; I on Urtica..... D. Urticae (Schum.) Kuntze
Urediniospores medium-sized (15-21x19-25u); teliospores medium-
sized) ((37-SSuimlones: on 'CarevsiL ion: (Ribess. «ic: aan eens eee
ryote tote cee elect at eRe he SIS pve Apts bolo he D. Grossulariae (Schum.) Kern.
(=Puccinia Grossulariae (Schum.) Lagerh.
Urediniospore-pores 2, in the upper part of spore.
Urediniospores medium-sized (15-19x19-24u); teliospores medium-
sized (35-50u long); on Carex; I on Aster, Solidago, and Eri-
GEFAG ee eh onG hao c x oe sees D. Erigeronatum (Schw.) Arth.
Urediniospores large (17-21x23-32u) ; teliospores large (42-65u long) ;
on Carex; I on Sambucus.............. D. Sambuci (Schw.) Arth.
Host belonging to composite family, genus Helianthus; autoecious..........
EA RRS Hh TES RSS EAS REPRE PE ae RORY A EAS D. Helianthi (Schw.) Kuntze

GENus Uromycopsis (ScHROT.) ARTH.

The character of the pycnia, aecia, and telia are essentially like
the genus Nigredo, but the uredinial stage is wanting. The telia
often arise within the aecia or about them from the same mycelium.
A good example of the genus is U. Psoraleae (Peck) Arth. on var-
ious species of Psoralea from Minnesota, Illinois and Texas west-
ward to the Pacific coast. The genus is more common westward.

Genus ALLopus ARTH.

This genus bears the same relation to Dicacoma that Uromy-
copsis does to Nigredo. A. Podophylli (Schw.) Arth. is a com-
mon and widely distributed species, occurring on Podophyllum pel-
tatum. The teliospore-walls of this species are especially interest-
ing on account of the straight or curved conspicuous spines with
which they are beset.

GENUS KLEBAHNIA ARTH.

No cupulate aecia are present in this genus, the pycnia being
followed by a stage of the uredinial-type. Only a few species have
been referred here of which the more common one is K. Glycyrr-
hizae (Rabh.) Arth. on Glycyrrhiza. This is found from North
Dakota and Kansas westward.

PLANT RUSTS 65

Genus Butraria DC.

Resembling Klebahnia except for the possession of teliospores
having two cells. A widespread species is on various members of
the family Cichoriaceae, Hieracium, Agroseris, Nothocalais, and
Crepis, for which the oldest name seems to be B. Hieracit (Schum.)
Arth. The teliospore-walls are finely verrucose and uniformly
thick, I-1.54. Another species on false boneset (Kuhnia) is B.
Kuhniae (Schw.) Kern (Puccinia Kuhniae Schw.) with telio-
spore-walls smooth and thicker above, 3-4 at sides, 5-7» above.

GENERA TELOSPORA ARTH. and Dasyspora B. & C.

To these genera belong species with short life-cycles. In some
the teliospores germinate only after a resting period (micro-forms),
in others they germinate at once (lepto-forms). The telia are usual-
ly compact and arranged in circinating or crowded groups. Most
specimens showing teliospores germinating upon maturity can be
placed here with considerable confidence, as they very rarely be-
long to genera with other spore-forms in the life-cycle. The I-
celled forms belong to Telospora, and the 2-celled forms to Dasys-
pora. Only a few species are known in the former genus. Telo-
spora Rudbeckiae (A. & H.) Arth. on Rudbeckia laciniata is the
most likely to be met with. Dasyspora is a large genus. D. Ane-
mones-Virginianae (Schw.) Arth. on Anemone and D. Xanthu
(Schw.) Arth. on Xanthium are of common occurrence.

FORM-GENERA

In addition toethe forms of known life-cycle, which may be re-
ferred to true genera, there are many forms whose life-cycle is too
imperfectly understood to permit them to be placed with confidence
in any of the known genera. Many of these can be recognized
merely as a stage and judging from analogy it is safe to assume that
they cannot be independent but must be associated with other stages.
In order that such forms may have names so that they may be dis-
cussed more easily the practice has grown up of using certain
terms as if they were really generic names, when in fact they rep-
resent only stages. For example aecial forms of the usual cluster-
cup type whose connections are unknown are placed under Aeci-
dium; aecial forms of the blister-type inhabiting the pine family

66 F. D. KERN

(Pinaceae) are treated by most writers under Peridermium; while
aecial forms lacking a peridium are considered under Caeoma.
Uredinial and other similar looking stages are referred to Uredo.
These names which are accorded generic treatment, but which in-
clude only isolated stages, are referred to as form-genera, and as
such they serve a useful purpose in disposing of the residue of im-
perfectly known forms.

Agricultural Experiment Station.
Purdue University.

BIBLIOGRAPHY
ArTHUuR, J. C.

North American Flora, Vol. 7, Part 2 (1907), Part 3 (1912) ; pub. by The
New York Botanical Garden. In this work are given keys, descriptions,
synonomy, hosts and distributions. The genera are treated essentially
in the order given in this paper and the parts published extend through
the genus Nigredo.

Terminology of the Spore-Structures in the Uredinales. Bot. Gaz. 39:
219-222. 1905.

Clues to Relationship among heteroecious Plant Rusts. Bot. Gaz. 33 :62-
66. 1902.

Problems in the Study of Plant Rusts. Bull. Torrey Bot. Cl. 30 :1-18. 1903.

North American Rose Rusts. Torreya g:21-28. 1909.

CarRLETON, M. A. °

Investigations of Rusts. U.S. Dept. Agric., Bureau Plant Industry, Buil.

No. 63 (1904).

Cereal Rusts of the United States. U.S. Dept. Agric., Div. Veg. Phys. &
Path. Bull. No. 16 (1899).

Cuinton, G. P.

Heteroecious Rusts of Connecticut having a Peridermium for their Aecial
Stage. Report of the Botanist for 1907, Conn. Agric. Exper. Station
(1908), pp. 369-396, with several plates.

FREEMAN, E. M. and Jounson, E. C.

The Rusts of Grains in the United States. U. S. Dept. Agric., Bureau of
Plant Industry, Bull. No. 216 (1911).

PLANT RUSTS 67

Jounson, A. G.
The Unattached Aecial forms of Plant Rusts in North America. Proc.
Indiana Acad. Science for 1911, 375-413 (1912). Enumerates 1o1 forms,
with data concerning hosts and localities.

Kern, FRANK D.

Methods Employed in Uredineal Culture Work. Proc. Indiana Acad.
Science for 1905, 127-131 (1906).

The Morphology of the Peridial Cells in the Roesteliae. Bot. Gaz. 49:
445-452, with plates XXI & XXII. toro.

The Rusts of White and Red Clover. Phytopathology Vol. 1, pp. 3-6.
IQII.

A Biologic and Taxonomic Study of the genus Gymnosporangium. Bull.
N. Y. Botanical Garden Vol. 7, No. 26, pp. 391-483 (1011), illustrated.

Orton, C. R.
Correlation between certain species of Puccinia and Uromyces. Mycolo-
gia Vol. 4, pp. 194-204 (1912), illustrated.

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DEPARTMENT OF NOTES, REVIEWS, ETC.

It is the purpose, in this department, to present from time to time brief original
notes, both of methods of work and of results, by members of the Society. All members
are invited to submit such items. In the absence of these there will be given a few brief
abstracts of recent work of more general interest to students and teachers. There will be
no attempt to make these abstracts exhaustive. They will illustrate progress without at-
tempting to define it, and will thus give to the teacher current illustrations, and to the
isolated student suggestions of suitable fields of investigation [Editor.]

NOTES ON SOME PECULIAR SENSE ORGANS FROM DIPTERA

The Diptera are generally conceded to be descended from four-
winged ancestors, the posterior pair of wings having become rudi-
mentary. The rudiments of this posterior pair of wings are called
halteres, and are found as small club-shaped organs just back of
the normal wings.

These organs play an important part in the orientation of the
body during flight. If they are removed or otherwise interfered
with, the flight is disturbed and in some cases prevented.

At the base of the stalks of the halteres are to be found some
highly developed organs, which appear to be sense organs.

In Figs. 1 and 2 which are drawn from an Ortalid, called
Stranzia longipenis, will be seen a dorsal and a lateral view of a
halter. There are apparently two kinds of sense organs depicted
here, one (A) and (C) situated on opposite sides of the stalk, and
one (B) situated on the chitinous sheath which covers the base of
the organ on the dorsal side. (See Plate I).

Fig. 2 shows a side view of the halter. These sense organs
when viewed with a higher power present an appearance something
like Fig. 3. There are ten rows of the oval disks, with as many
rows of rudimentary hairs or spines between them. Organs (A)
and (C) are identical in these particulars. These oval disks have
a swelled or crowning surface, which leaves them distinctly raised
in rows.

In Fig. 4, is a diagram of the disk arrangement of the basal
sheath. Here the disks lie in rows between chitin ridges, those
nearest the center being nearly overgrown with small spines.

70 NOTES, REVIEWS, ETC.

An explanation of the nature and function of these organs is
not certain nor easy. The following is offered as suggestive.

Let us look for a moment at the more primitive type of fly;
we may find here a clue to the course these structures may have
followed in their evolutionary degeneration.

In the Brachycera, we find a type of insect which has very
simple forms of wings, the venation being mostly absent except
for a few parallel ribs which run lengthwise of the wing. The
wings are covered with spines which lie in rows alternating with
each other as in Fig. 7. The halteres still retain their wing shape
and the spines on them preserve the arrangement found on the
anterior wings.

In another family still more highly organized, we find the rows
of spines more definitely gathered and specialized into rows, which
rows of spines are separated by spaces such as are seen in Fig. 5.

Finally, in the elaborate organization of the soaring and pois-
ing flies we find, as described above for Stranzia, the rows of hairs
or spines alternating with the rows of disks. Microtome sections
of these organs show the oval disks as hollow and filled with fluid
during life.

The rows of degenerate spines seem to be connected with the
central nervous system, and are assumed to be sensory.

From a histological standpoint the halter is composed of an
ectodermal layer of cells, which secretes the chitin with its many
sensory spines, and an interior mass composed of trachea, nerves,
and fluids with corpuscles. See Fig. 9 for a very diagrammatic
view of a section of the halter.

Fig. 8 is a much enlarged view of the cells in the sense organ
on the stalk of the halter. The disks are formed from large oval
cells (A), and the sensory cells are at (B).

The writer is unable to say whether the disc cells are also
sensory; altho it is possible that they are. It seems at least prob-
able to him that they may be considered as homologous with the
ordinary smooth membrane interspinal spaces on the normal wing.
It is not clear from the structure of these organs just how they
contribute to equilibrium, unless in some way they control the blood
supply to the vascular terminal bulb.

Sete

AMERICAN MICROSCOPICAL SOCIETY 71

Some experiments conducted to determine what relation these
sense organs on the halteres have to flight and to orientation in
flight may prove interesting to those who have not made special
study of the subject.

In order to determine the relation of the halteres to flight the
writer removed the entire halteres, by cutting, in a number of
specimens of Muscidz. Flies so treated were all incapable of con-
trolling their flight, usually pitching violently downward when at-
tempting to fly.

A similar number of flies was taken, and, without removing
the halteres, a small amount of liquid balsam was introduced
under the sheath and over the sense organs. Specimens treated in
this way could not be induced to undertake flight.

These two experiments show clearly that the halteres play an
important role in equilibrium in flight, and that they can be put
out of commission, as effective organs, without actual removal.
This suggests the existence of certain subordinate parts on which
the functioning of the organ depends.

It now remains to localize, if possible, the responsible portion
of the halter. In doing this, larger flies, as Sarcophagide,
Syrphidz, and Tachinidz, were used. An effort was made in these
flies to injure the structures referred to above as sense organs, and
to confine the injury to these. Cauterization with a hot needle
was attempted; but this was difficult to control, and often resulted
in too extensive a wound. The other method used was the appli-
cation to the so-called sense organs of a small amount of nitric or
sulphuric acid, without allowing it to reach the terminal bulb. The
flies were held for a minute or so to allow the acid to act. Insects
treated in this way pitch headlong in attempted flight much as those
whose halteres were removed. Some forty specimens were so
treated. One of two conclusions seems necessary :—either the acid
penetrates and essentially destroys the whole organ, or there is a
special sensory portion which was destroyed and prevented the
ordinary reaction.

The conclusions which the writer thinks reasonable are:

1. The halteres are necessary to successful balancing in flight
in Diptera.

N
to

NOTES, REVIEWS, ETC.

2. The peculiar and definite organs at the base are sense
organs, and are necessary in giving the halteres functional value.

3. These sense organs are in some way aroused by the changes
in position, and thru them the central nervous system is enabled to
control the process of balancing.

A CONVENIENT DROPPER FOR USE IN CUTTING CELLOIDIN SECTIONS

A very useful aid in cutting celloidin sections is shown in the
accompanying figure (Plate Il). This piece of apparatus was in
stock when the writer assumed charge of this laboratory, and he is
not acquainted with its history. While it is not listed in any of the
dealers’ catalogs that the writer has examined, it may be made at a
very slight cost in any machine shop.

It consists of a glass oil-cup (1) of about 40 cc. capacity, with
a mill-head (2) at the top to regulate the flow of alcohol. The cup
is fastened to a bar (3), which is slotted for about 34 its length to
receive the bolt that extends through the column (4) that holds the
cup a few inches above the knife (5). The head of the bolt men-
tioned above is of the proper shape to fit into the slot in the knife-
carrier, and the thumb-nut (6) on the other end of the bolt tight-
ens at one time both the bar (3) to the column (4) and the column
to the knife carrier. This thumb-nut and its bolt, which, except in
length, are exactly those (7) that hold the knife in position, make it
possible instantly to adjust the cup so that the alcohol will fall on
any desired part of the knife; and since the apparatus is attached
to the carrier it will always be over the same part of the knife even
in microtomes where it is the knife that moves. If all the metal
parts are nickel-plated it will obviate trouble in drying off the alco-
hol to prevent rusting.

A. M. REESE.
Department of Zoology, West Virginia University.

CRITICAL ILLUMINATION FOR THE MICROSCOPE

In a brief paper (J. Queck. Micr. Club. Nov. 1912) Reid gives
some important suggestions for critical illumination, which will cer-
tainly be of value to beginners in the use of the microscope and to
many older users who have not given critical attention to the sub-

474
0,0.00,0
0,0,02°
Fig. 3
Res preccemetees - a
Np Se TOON —
Bese ie =
e—
cdhh4dd44a4
Fig. 5 ae

PLATE |
Sense Organs of Diptera

PLATE I]
Dropper for Celloidin Sectioning

AMERICAN MICROSCOPICAL SOCIETY 73

ject. There is little question that most of us, reared in school labor-
atories, do not get the nice, exact results in the use of microscopes
which are obtained by the thorogoing students of microscopy.

Certain simple precautions leading to good illumination intro-
duce the paper :—Cut out all unnecessary light from the room, so
that no light gets to the eye except thru the microscope; save the
best eye for critical moments by using the other eye for prelim-
inary steps; use color screens complementary to the stains used,
green for red, yellow for blue, etc. The subject of illumination
itself the author discusses under these heads: The most suitable
light; collecting lenses; principles of correct illumination both of
the field and of the object itself ; condensers; distance of lamp from
substage mirror; critical and non-critical illumination; working
aperture; general arrangement of light and apparatus in high,
medium, and low-power work.

For the detailed discussion of these topics the readers must be
referred to the original paper.

CLEANING DIATOMS

Blake (Am. Jour. Sci. Jan. 1913) calls attention to the interest
in cleaning, mounting, and study of diatoms. After recounting the
difficulties attendant on the usual methods he describes a method
originated by himself some twenty years ago.

Instead of the older method of treating with acid, diluting with
water, and repeated decanting, the author devises an organic seive
made by cementing a thin cross-section of some coniferous wood
to a small glass vial whose bottom has been cut off for the purpose.
The wood is cut about one-quarter millimeter in thickness, from a
suitable piece of wood kept until the operation in boiling water.
This is done by means of a sharp, thin-edged chisel.

The operation of cleaning the diatoms consists of placing the
digested diatom material, moderately diluted, in the vial, and by
means of a suitable rubber compression bulb, alternately pressed
and released, of forcing the acids and salts thru the seive, and the
clay and fine sand thru or into its pores. These diatoms which are
longer than the diameter of the pores will remain behind with larg-
er grains of sand which must be removed in some other way.

74 NOTES, REVIEWS, ETC.

It is necessary to see that the strainer does not become choked
This may be prevented by shaking. The strainer should, of course,
be kept in water between uses. When it finally becomes clogged
with sand, a new one must be put on.

By using wood with different sized conducting vessels, a sort-
ing of the diatoms may be affected. By using pine, spruce, white
wood of the red cedar, a graded series of strainers can be had, the
last being much finer than the first.

STAINING PROTOZOA

Darling (Science: Jan. 10, 1913) calls attention to the dearth
of knowledge of the acidophilic substances in the nuclei of protozoa,
owing to the predominant use of basophilic staining substances, and
to the “lack of a satisfactory technic for demonstrating acidophilic
substance in wet fixed films.”

The author suggests careful differentiation of such polychrome
stains as Romanowsky or Hastings-Giemsa, by ammoniated ethyl
alcohol. Under such conditions, studying Entamebae, he found a
definite arrangement of an acidophilic substance (oxychromatin)
within the nucleus, showing a structure quite different from that
shown with the usual basichromatin stains. He believes that care-
ful and critical study will reveal that this oxychromatin may have
important functional relations to the changes that are so well known
in the true chromatin in nuclear activity.

DOUBLE-STAIN METHOD FOR THE POLAR BODIES OF DIPHTHERIA BACILLI

Dr. Marie Raskin (Apoth. Ztg. XX VII, p. 10; Abstr. United
St. Naval Med. Bull. Vol. 6, No. 4, p. 611) proposes a technic for
these bodies, whose distinctive value lies in the fact that only two
operations are necessary, i. e., the application of a stain with both
colors present, then water washing.

Formula for stain :—

Glacial (Acefie Acid.) 2% 22608 BGe:
Dist: Watert19.425. 428, aR gg) *
Ade A G5G6) 2 ivi A a ee 100 “
Old Sat. sol Methylene blue..... 4 “

Ziehl’s phenol fuchsine Sol..... 4

AMERICAN MICROSCOPICAL SOCIETY 75

Drop mixture in a thin layer over the specimens on the cover
glass; heat through the flame. The alcohol ignites and is permit-
ted to burn off, after which the specimen is washed in water and
dried. The entire process takes 20-25 seconds, and the stain re-
mains serviceable for any length of time. Polar bodies appear deep
blue and the bacilli bright red. Even in smears with a preponder-
ance of other bacteria, individual diphtheria bacilli may be readily
and unmistakably identified.

A NEW TECHNIC IN STAINING DIPHTHERIA SPECIMENS WITH
TOLUIDIN BLUE

Dr. Constant Ponder (Lancet, July 6, 1912; Abstr. U. S. Naval
Med. Bull. Oct. 1912, p. 612) recommends the following treatment
for diphtheria bacilli :—

The stain:
Toluidin blue (Grubler)....0.02 gram.
Glacial acetie acide: 20.) 2t EMEC.
ADSHAlOR) SNE OIN Heh are
Distilled Water to make..... 100 ‘“

The film made on cover glass is fixed as usual. Spread stain
on film. The cover glass is then turned over and mounted as a
hang-drop preparation. Typical diphtheria bacilli are said to stain
blue, with red granules. The author gives this as a new method,
and says it is preferable to either Methylene blue or Neisser’s stain.

NOTES FROM MEETING OF THE ILLINOIS MICROSCOPICAL SOCIETY,
Chicago, Oct. Io, 1912

Mr. N. S. Amstutz showed a useful contrivance for keeping
pond life in place. It consisted of a piece of brass about 7/8 in.
square and 5/32 in. thick. A series of seven holes were drilled
thru it so as to imprison that many varieties of pond life at one
time. The plate was placed in a flat bottomed watch glass and
each specimen transferred with a pipette to its proper “cell.” These
could be then studied at will very nicely with a 2/3 objective and
various combinations of oculars. The specimens were confined
laterally so they were unable to move out of the field of view though
having abundant room for vertical movement. With the coarse ad-

76 NOTES, REVIEWS, ETC.

justment the up and down variation could easily be followed. It
proved a great satisfaction to examine water fleas, mosquito larvae,
etc., when fenced in. The holes were arranged 6 in a circle of 1/2
in. diameter and the seventh in the center. Their diameter was de-
termined by measuring the diameter of the field with a stage
micrometer and then selecting the next smaller size of twist drill
by which to do the drilling. To guard against the smallest
animalcule creeping between the brass and the watch glass the
bottom face could be covered with a thin film of balsam, air dried
until quite of proper consistency, and then a cover glass pressed
into intimate contact, so that no balsam would run into the spaces.
Vipa A. LATHAM, Secy.

BOG SOLUTIONS AND PLANTS

Dachnowski (Bot. Gaz. Dec. 1912) writes on the physiological
effects of peat or bog solutions on the plants subjected to them. It
has been clearly established that the nature of these organic solu-
tions and of the bacterial flora maintaining life therein is a very im-
portant factor in limiting the higher life of these regions. The fact
that some plants tolerate these conditions and others do not makes
clear a difference in the plants themselves. The writer is endeavor-
ing to see what it is that makes this difference in plants exposed to
the solutions. The responsibility must rest either upon difference in
diosmotic qualities of the plasmic membranes, or upon differences
in cytoplasmic resistence, or on both. He finds the following facts
which help to localize the solution of the problem: (1) Some plants
may cause the precipitation of the hurtful materials in the solu-
tions in an insoluble form, by enzymic action. This conceivably
may take place outside the membrane, inside the cell ; or in the mem-
brane itself, affecting its permeability ; (2) other plants may possess
the power of assimilating with impunity these organic substances.

It is well known that these solutions have little effect on certain
xerophytic plants, while they totally inhibit agricultural plants. The
value of the work is evident as bearing on the agricultural use of
peat lands, on the nature of xeromorphy itself, as well as on the
successions of vegetation in the bogs.

AMERICAN MICROSCOPICAL SOCIETY 77

EFFECT OF CROPPING ON SOIL BACTERIA

Brown (Centralbl. Bakt. Abt. 2, XXXV, 1912, p. 248) has
studied the effect of different kinds of cropping on the bacterial
content of the soil. He finds that the number of microorganisms
in the soil is much increased by rotation of crops as compared with
continuous cropping. The same is true of the nitrifying and nitro-
gen-fixing powers of the soil. He compares various systems of
alternation of crops in this regard. He also discusses the effect of
turning under clover, as green manure. He claims that the two
year rotation with green manuring is not so effective in increasing
the bacteria and bacterial products as the longer term rotations. It
is shown that the productivity of the soil is closely related to the
bacterial activities within it.

ALTERNATION OF GENERATION IN THE PH/ZO0PHYCEZ

In a beautifully illustrated article (Bot. Gaz. Dec. 1912)
Yamanouchi gives, from the study of the nuclear and experimental
conditions, the grounds for believing that Cutleria multifida is the
gametophytic phase of a species of which Aglaozonia reptans is the
sporophytic stage. The nuclei of both male and female Cutleria
plants contain 24 chromosomes, which is true also of the gametes
themselves. The sporelings resulting from the union of these
gametes contain 48 chromosomes and develop into an Aglaozonia
form similar to A. reptans in nature. On the other hand the
nuclei of Aglaozonia reptans contain 48 chromosomes, which is
reduced in zoospore formation to 24. These zoospores germinate
without conjugation, and produce plants similar to young Cutleria in
nature, and with 24 chromosomes.

EXPERIMENTS ON THE GERMINATION OF TELEUTOSPORES

Dietel (Centralbl. Bakt. IT. 31:95. 1911) reports on the effects
of age and temperature and drying, etc., on the germination of
teleutospores of Melampsora. In the early spring these spores
germinate in about 3 days if brought into favorable conditions of
temperature and moisture. As the spores grow older the time
necessary to germinate decreases. This might be due either to in-
ternal ripening or to the progressive changes in the spring tempera-

78 NOTES, REVIEWS, ETC.

ture. Temporary drying hastened germination; strong light de-
layed it; temporary freezing had no effect. Germination takes place
at 6°-10° C., but is hastened by higher temperature of 15°-20°.

DIRECTION OF LOCOMOTION IN STARFISH

Cole (Jour. Exp. Zool. Jan. 1913) finds that Asterias forbesi in
the absence of directive stimuli, in crawling advances most frequently
with that part of the body forward in which the madreporite occurs.
He found a tendency in these animals to persist in moving with the
same parts foremost in a series of succeeding trials; tho there is
also a tendency to shift or “rotate” this anterior point successively to
other parts. The author thinks the madreporic body may be what
determines anteriorness, and shows that the “physiological an-
terior” of the starfish corresponds in this respect to the anterior
parts of the more bilateral spatangoids.

A ROTIFER PARASITIC IN EGG OF WATER SNAILS

Stevens (Jour. Queck. Micr. Club., Nov. 1912) describes a
rotifer of the genus Proales which is able to bite a small opening in
the tough egg membrane of the snail Limnaea auricularia, and by
squeezing thru this enters the more fluid portion within. The
rotifer feeds on the fluid gelatin of the egg with an occasional attack
on the snail embryo itself. As the result of these attacks the snail
embryo is finally killed.

In the meantime the rotifer lays its eggs, and later leaves this
to enter still other eggs. The larvae hatch and undergo their de-
velopment, devouring the dead snail embryo and other available
substance of the egg. They too later escape and enter other eggs.

This looks somewhat like a parasite in the making. The author
says the rotifers do not seem “at home” in the water while making
their way from egg to egg.

EUGLENIDS AND THEIR AFFINITIES

Alexieff (Arch. Zool. Exp. Notes et Rev., No. 4. 1912) in con-
nection with the discussion of certain euglenoid forms that are partly
or largely parasitic on other animals, makes some interesting sugges-
tions as to the relationships of Protozoa. He thinks the Euglenids

AMERICAN MICROSCOPICAL SOCIETY 79

are near the flagellate source of the Sporozoa, and from thence as a
main stem arise the Trypanosomes, Coccidians, Gregarines,
Haemogregarines. He feels also that the Euglenids may give rise to
lines leading to Cystoflagellates and Ciliates.

AN AMEBA WITH TENTACLES

Collin (Arch. Zool. Exp. N. & R., No. 4. 1912) describes a
new protozoan combining the characters of Ameba and the Suctoria.
The organism has a gelatinous covering whose form is easily
changed, and possesses tentacles by which it attaches itself to objects.
It has the nuclear and pseudopodial structure of the Ameba. It is
a marine form occuring in a culture of seaweed along with other
amebz and Foraminifera.

SOME AMERICAN RHIZOPODS AND HELIOZOA

Wailes (Jour. Linn. Soc. Dec. 17, 1913) reports 161 species and
varieties of Rhizopods and 4 species of Heliozoa from collections
made in 1911 at Augusta, Georgia, in New Jersey, and at various
points in New York. Comment is made upon the small amount of
work done on the American species of these groups since the time
of Leidy.

Of these, 5 species and 10 varieties are new. Forty of them
are recorded for the first time from the United States. About 80%
of the species are similar to those found in Europe. The remainder
are made of species rarely or not at all found in Europe. The
author states that considerable local variation exists in some of
the species.

SIZE OF CHROMOSOMES AND PHYLOGENY

_ Meek (Jour. Linn. Soc. Sept. 24, 1912), thru a study of the
diameters of chromosomes, has reached the conclusion that there
are three diameters of chromosomes found in animals,—.2Ip in
Protozoa, .42 in low Metazoa, and .83u in high Metazoa. He holds
that these measurements are remarkably constant. This arith-
metic progression is believed by him to mean a lateral fusion of these
chromatic elements in phylogeny.

In respect to length, the author finds, by study of spermatogen-

8o NOTES, REVIEWS, ETC.

esis in several species of Stenobothrus that the chromosomes of
the spermatocytes are made up of rods, sometimes 2 and sometimes
4. The length of these rods varies in arithmetic progression. In
each of 4 species studied there are 5 short chromosomes, no two
of which are the same length; altho the 5 short chromosomes in one
species correspond with the 5 short ones of the others. There are
also 3 larger chromosomes in each species, but these long chromo-
somes do not belong in the different species to the same numerical
series. The author believes that the external specific differences
between the species are dependent on the differences in the long
chromosomes, altho he is unable to establish the correlation between
the rod-lengths and the body characteristics.

SPERMATOGENESIS IN HYBRID PIGEONS

Smith (Quart. J. Mic. Sci. 1912, p. 159) reports studies of the
sperm formation and structure in the hybrids formed by mating a
male pigeon and female domestic dove, and compares these with
the condition in pure breeds.

In the first maturation division in the hybrids the chromosomes
do not unite to form 8 bivalent chromosomes but occur quite ir-
regularly about the spindle and are finally distributed to the poles
irregularly.

The second maturation division is practically suppressed. The
secondary spermatocytes proceed at once to form spermatids and
spermatozoa. Many of these are on the average twice the normal
size, altho otherwise apparently normal structurally. In other cases
there were structural abnormalities.

It is known experimentally that hybrids of these stocks are in-
fertile, and it seems that the sterility may be due to the inability of
the specifically different chromosomes to unite in the normal synapse,
with the consequent disturbance in the whole maturation process.

MALE GERM CELLS IN NOTONECTA

Browne (Jour. Exp. Zool. Jan. 1913) discusses the differences
in form and number of the chromosomes in three species of Noto-
necta. She finds that the differences in the chromosome condition
may be explained in these species by the relations of two particular

ee

AMERICAN MICROSCOPICAL SOCIETY 81

chromosomes. In N. undulata the two chromosomes in question
are always separate; in N. irrorata are always united to form a
single body; and in N. insulata they may be separated in the first
spermatocyte division, but are united in the second.

The author traces the origin of the chromosomes from the
karyosphere in the three species, and their behavior in the growth
stages and maturation divisions.

INTERSTITIAL CELLS OF TESTIS AND SECONDARY SEX CHARACTERS

J. des Cilleuls (C. R. Soc. Biol. Paris, 1912, p. 371) finds a strict
coincidence in the development of the interstitial cells of the testis
and the secondary sexual characteristics of the cock. In chickens
the external marks of sex do not begin to appear until about the
thirtieth day. By the time the chicks are 45 days old the pullets
show a greater development of the tail feathers and the cockerel
more color and size of comb. The sex distinctions increase from
this point. The author claims that the secondary sex characters in
the male bird begin to show with the oncoming of the interstitial
cells, and increase as these increase. The author believes that the
secretion of the interstitial cells acts as a hormone in stimulating the
growth of the characteristic male secondary structures.

MICROBIOLOGY IN RELATION TO DOMESTIC ANIMALS

This book, entitled “Principles of Microbiology,” with a sub-
title “A Treatise on Bacteria, Fungi, and Protozoa Pathogenic for
Domesticated Animals,” is written primarily for veterinary stu-
dents beginning the study of microbiology. It consists, in about
equal parts, of matter belonging to general bacteriology and to spe-
cial applications of this to veterinary science. In the very nature of
the case this makes the treatment of general bacteriology somewhat
less satisfactory than may be had from text-books on this subject,
and limits the author somewhat in his treatment of the part of the
subject which is peculiar to the book.

The first twelve chapters are given to such subjects as the
biology, morphology, classification of bacteria; the apparatus, meth-
ods of sterilization, cultivation, staining, and examination of bac-
teria; the relation of bacteria to disease. In the part relating to

82 NOTES, REVIEWS, ETC.

the work of the veterinarian there are, first, two introductory
chapters dealing with the Use of Animals in Bacteriological Exam-
inations and Investigations, and the Bacteriology of Water and Milk.
These are followed by eight chapters dealing with the various prin-
cipal genera and species of microdrganisms that produce diseases in
domestic animals, together with their pathogenesis and, where
known, the treatment. These chapters present very valuable ma-
terial for the general student of biology, as well as for the vet-
erinarian.

In the concluding chapters the author discusses some of the
broader questions of physiology, theory, diagnosis and therapy of
the bacterial diseases under the heads:—Specific Bacterial Pro-
ducts, Tissue Reactions and Immunity ; Serum Diagnosis ; Immunity
and Vaccine Therapy. This resumé is very readable and valuable
to the general student. The mechanical excellence of the book is
all that could be desired.

Principles of Microbiology, by VY. A. Moore. Pages 506; illustrated. Carpenter &
Co., Ithaca, N. Y. Price $3.50.

BEGINNERS GUIDE TO THE MICROSCOPE

This is an elementary handbook designed to aid the untechnical
person to use the microscope for his own pleasure and that of his
friends. The need of such a book seems to the author to lie in the
great complexity of the modern instrument and the wealth of its
accessories, and in the elaborate character of the modern books
about the microscope. In a very simple, gossipy way quite suitable
to his expressed purpose, the author describes the microscope and
its essential parts, the formation of images, illumination; discusses
the principles that should guide in the choice of an instrument ; gives
rules for the use of the instrument and for its care; tells of interest-
ing objects for temporary mounts. There are also sections on the
home aquarium, on collecting objects, on mounting for permanent
display, and on storing slides.

In many ways it is much to be regretted that there are not more
of our modern Americans who turn to such methods of interest
and diversion as are suggested here. The use of the microscope as
a serious instrument of education and research in schools has in-

AMERICAN MICROSCOPICAL SOCIETY 83

creased greatly in this country; but it is remarkable that so few

people use it as a means of recreation, pleasure, and general culture.
The Beginners Guide to the Microscope, by Chas. E. Heath, F. R. M. S. Illustrated;
120 pages. Price 1 shilling. Percival Marshall & Co., London.

MICROSCOPY AND DRUG EXAMINATION

In this little book the author seeks to present in a simple and
condensed form the elements of microscopy and histology demanded
by pharmaceutical students. In Part I, which is given to Micro-
scopy, are discussed briefly,—often too briefly to be satisfactory,—
microscopes, microscopic photography, manipulation and care of
the microscope, reproduction and measurements of microscopic ob-
jects; histology, microchemistry; the preparation and mounting of
microscopic objects; cells; plant and animal tissues; microscopy of
starches, etc. A series of laboratory exercises illustrating certain
part of plant and animal biology follow.

Part II is taken up with suggestions as to the microscopical ex-
amination of some 35 “drugs” in their commercial form. In Ap-
pendix A is a valuable table defining the various elements consti-
tuting and produced by cells, giving their properties and the meth-
od of identifying them by staining or otherwise.

The last 50 pages of the book are given to figures illustrating
lenses, microscopes, drawing apparatus, tissues, organs, drugs.

Mechanically the book is marred by the unnecessarily large
type in which the words desired to be emphasized are printed.

Microscopy and the Microscopical Examination of Drugs, by Charles E. Gabel,
Ph.D., Microscopical Food and Drug Analyst Iowa State Dairy and Food Commission.
Illustrated; 114 pages. Price $1.00, postpaid. Des Moines, Iowa.

NECROLOGY

Announcement of the death of the following members of the
American Microscopical Society has been received since the issue of
the last number:

A. E. Aubert, ’12, New York City.

Geo. C. Crandall, M.D., ’04, St. Louis, Mo.

J. D. Hyatt, Past President and Honorary Member, New Ro-
chelle, N. Y.

AMERICAN MICROSCOPICAL SOCIETY 85

PROCEEDINGS

of the American Microscopical Society
MINUTES OF THE CLEVELAND MEETING

The Society was called to order by President F. D. Heald in the Biologi-
cal Laboratory of the Western Reserve University, Cleveland, Ohio, at 3:30
p. m., Jan. I, 1913.

The reports for 1912 of the Custodian and Treasurer were read and
ordered referred to an auditing committee. This committee was later named
by the President, consisting of Professor Frank Smith and Mr. J. E. Ackert,
both of Urbana, Illinois.

Due to the fact that only one business session was provided for, it was
unanimously agreed to suspend By-laws V and VI, and to nominate officers
from the floor. The following officers were nominated and elected for 1913:

President: Professor F. Creighton Wellman, School of Tropical Medi-
cine, Hygiene and Preventive Medicine; Tulane University, New Orleans, La.

First Vice President: Professor F. C. Waite, Medical Dept., Western
Reserve University, Cleveland, Ohio.

Second Vice President: Professor H. E. Jordan, University of Virginia,
University, Va.

Treasurer (3 years): Professor T. L. Hankinson, Eastern Illinois Nor-
mal School, Charleston, IIl.

Elective Members Executive Committee: Dr. H. L. Shantz, Bureau
Plant Industry, Washington, D. C.; Professor J. W. Scott, Kansas Agricul-
tural College, Manhattan, Kansas; Professor George E. Coghill, Denison
University, Granville, Ohio.

Members of the Council of the A. A. A. S.: Professor F. D. Barker,
University of Nebraska, Lincoln, Nebraska; Professor A. M. Reese, Uni-
versity of West Virginia, Morgantown, W. Va.

An informal discussion, without vote, was had concerning the utilization
of the Spencer-Tolles Fund for research, and other items of general policy.

The verification and publication of the Minutes were left in the hands
of the President and Secretary.

86 AMERICAN MICROSCOPICAL SOCIETY

CUSTODIAN’S REPORT FOR YEAR 1912

SPENCER-TOLLES FUND

Reported at) Wadshineton Mecting oe hi iosc es vcs yes nash dane sehn ies $3,352.16

Dividends teceived Muring: Veat AO lee et kiiotese\er ois sstere bislela eM aie eeite ne 203.28

SNE EMER) ome ioncis We wines © eke Wate ew aie AER oboe inate: ce pis See atures ewer 4.00

$3,559.44

Less ‘dues paid for Lifte-members.: . 2002 eee ed RE SRY 8.00
Total WvVEsted eres: < kM eee eee aa ate tale Melons le saidee aia abeeles $3,551.44
het unchease ‘dariny years. 2.8. kOe ARUP ea $ 199.28
Grand Totals:

All:contributionsetondater: sic. cc cince cuise tee wtoe s cies Oeisaee $ 700.27

Pill Saleeas TOTOCCERINGS. . ose h sno cc vane ce seb cree cewar's 625.73

PATA EIEIO SHINS CP eS ok sein lpictcas vet ee ccs see bea siemet 250.00

Pill COPE AIA GIVIGENGS 7 cose acc heats ces ese teateak pee 2,115.44 $3,601.44
Less:

AIDS, eek as 5 Sie 64nd smiks cok creole Mee $ 100.00

AG Hite-menberchip 1aues: Sols Ss. cad. cae we eter dese wretts 40.00 $ 140.00
INGENDalanlce tales yc at's site BOTS eee ee el cercheet ete els $3,551.44

Life-members and Contributors of $50 and over: John Aspinwall, Rob-
ert Brown, (deceased), J. Stanford Brown, Henry B. Duncanson, A. H.
Elliott, John Hately, Iron City Microscopical Society, and Troy Scientific
Association. :
Macnus Priaum, Custodian.

We the undersigned committee hereby certify that we have carefully
examined the foregoing account, compared it with vouchers and found the
same correct. ,

FRANK SMITH,
J. E. Ackert,
Auditing Committee.

a ays.

To
To
To
To
To
To
To
To
To
To
To
To
To

AMERICAN MICROSCOPICAL SOCIETY 87

ANNUAL REPORT OF TREASURER OF AMERICAN
MICROSCOPICAL SOCIETY

December 21st, 1911 to December 26th, 1912.

RECEIPTS

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Signed, T. L. HANKINsSoN, Treasurer.

Examined and found to correspond with books and vouchers.
FRANK SMITH,
J. E. Acxert,
Auditing Committee.

TRANSACTIONS

OF THE

American Microscopical
Society

ORGANIZED 1878 INCORPORATED I89Q1

PUBLISHED QUARTERLY

BY THE SOCIETY

EDITED BY THE SECRETARY

VOLUME XXXII

NuMBER Two

Entered as Second-class Matter December 12, 1910, at the Postoffice at Decatur, Illi-
nois, under act of March 3, 1879.

Decatur, ILL.
Review PriInTING & STATIONERY Co.
1913

OFFICERS.

President: F. CREIGHTON WELLMAN, M. D................ New Orleans, La.
Mist Vice President: F.C. Waite, PHD. boc. i cos eee sien Cleveland, Ohio
Second Vice President: H.E. Jorpan, Ph.D............. Charlottesville, Va.
pecretarvie Le We GAREOQWAY 2 otic ciejeicnisit nie ciedla de ae etnlalel lee ede Decatur, Ill.
EF CRSUrey 1), La 1 EUANIKUN SONI ae ciclo acitnn 4s ee esis els sieteeeier Charleston, Ill.
Cectoutan.’ “NIAGNOUS: PREATINE loi. .5'40 o's ies Gasica a bid Ode rece Meadville, Pa.

ELECTIVE MEMBERS OF THE EXECUTIVE COMMITTEE

| WALES 0 6 Ae Bureau Plant Industry, Washington, D. C.
MENA SENET 49) rcfa Merah 2 abe cence Mer daieiabinae |” w cea cto etal a Leigh ols Manhattan, Kans.
Seta MINI Ea ce ects aia, ois eA eal wa tialne se G.8 sie,s oes omak led Granville, Ohio

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
Apert McCa tra, Ph.D., of Chicago, II.
at Chicago, Ill. 1883
T. J. Burrmt, Ph.D., of Urbana, III.,
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., r89o.
Stmon Henry Gace, B.S., of Ithaca, N. Y.,
at Ithaca, N. Y., 1895 and 1906
A. Ciirrorp Mercer, M.D., F.R.M.S., of Syracuse, N. Y.,
at Pittsburg, Pa., 1896.
A. M. Bree, M.D., of Columbus, Ohio,
at New York City, 1900.
C. H. E1icENMANN, Ph.D., of Bloomington, Ind.,
at Denver, Colo., rgor.
Cuartes E. Bessey, LL.D., of Lincoln, Neb.,

E. A. Birce, LL.D., of Madison, Wis.,
Henry B. Warp, A.M., Ph.D., of Urbana, IIL,

at Pittsburg, Pa., 1902.
at Winona Lake, Ind., 1903.

at Sandusky, Ohio, 1905.
Hersert Oszorn, M.S., of Columbus, Ohio,

at Minneapolis, Minn., 1910.
A. E. Herrzzer, M.D., of Kansas City, Mo.,
at Washington, D. C., 1911.
F. D. Heatp, Ph.D., of Philadelphia, Pa.,
at Cleveland, Ohio, 1912.

The Society does not hold itself responsible for the opinions expressed
by members in its published Transactions unless endorsed by special vote.

TABLE OF CONTENTS

FOR VOLUME XXXII, Number 2, April, 1913

Notes on Japanese Protozoa, with Plate III, by C. H. Edmondson and

TER MEU SINRNTST SRT LIMP CY wie’ s nba Sic cise cinta Pemiah ie alee mer eet Aloha. bieasbeatcty elk een 93
Experimental Amitosis in Onion Root Tip, by H. E. Jordan............. 103
Summaries in Micro-biology: The Black Moulds (Mucoracee), with

I JSS Wasa CAN AVR vac Payee santo Grace sducic icky lo vere iaccle ett eee bin stale seta ales Siebel meherntete 113

Notes and Reviews: Summary of the Elements in the Reproductive
Process (Plate VI), by T. W. Galloway; The Growth of the Com-
pound Eye (Plates VII and VIII), by E. W. Roberts; Fresh-water
Hydroids, by Edward Potts; Mutation in Micro-organisms; Differ-
entiation in Chromosomes; Bud-formation in Syllids; Cork Oak in
Portugal; A Simple Method to Remove Paraffin Sections Which are
Stuck to a Sheet of Paper or to the Hand; To Kill Mosquitos or
Other Insects; To Keep Slides at Constant Temperature; Section-
cutting in Gelatin by Freezing; Household Bacteriology............ 127

PEG OUAEETA [AE TS Ptae ee e ie ron ais wl din Jo nos a a sees stew we Ome Rat ialele ys Ot ie 159

TRANSACTIONS

OF

American Microscopical Society

(Published in Quarterly Installments)

Vol. XXXII : APRIL, 1913 No. 2

NOTES ON JAPANESE PROTOZOA
WITH FIGURES AND DESCRIPTIONS OF NEW AND RARE SPECIES

C. H. EpMoNDSON AND R. H. KinGMAN

The fresh-waters of Japan afford a wonderful opportunity for
the enthusiastic microscopist. Conditions under which simple or-
ganisms thrive are not wanting anywhere in that country. Flooded
rice fields of the lowlands, cool mountain streams and innumerable
lakes, large and small, are teeming with low plant and animal forms.

To what extent systematic study of the microscopic fauna and
flora of the waters of Japan has progressed, under the direction of
the eminent biologists of that country, the writers of this article are
not able to state.

With a view of determining the species of Protozoa characteris-
tic of Japan and comparing them with the American forms, micro-
scopic studies were carried on by C. H. Edmondson during July and
August, I912, in various parts of the main island. Beginning with
Kobe, observations were made through the central and eastern sec-
tions of the country and as far north as Lake Chuzenji.

Material was gathered from rice fields, small pools, streams and
lakes. Collections were made from the following large lakes: Lake
Biwi, altitude above sea level 328 ft.; Lake Hakone, altitude 2,378
ft.; Lake Chuzenji, altitude 4,375 ft. Since the survey covered a
wide territory with considerable variation in local conditions as well
as in altitude, the list of species embodied in this brief report may
well represent the characteristic unicellular fauna of the entire
country. The portion of this article concerned with Rhizopoda is
largely a result of the work of R. H. Kingman, a student of zoology,

O4 EDMONDSON AND KINGMAN

who identified and studied many forms from preserved material.
By comparing the list which follows with numerous local records of
observers in America and other parts of the world one sees some
added evidence of the wide distribution of many species of Protozoa.

The accompanying figures, prepared by Mr. Kingman from
permanent mounts, represent new, or rare species of Rhibopods or
forms showing considerable variation.

Phylum PROTOZOA: Subphylum SARCODINA:
Class RHIZOPODA Subclass AMOEBEA

Order GyMNOMOEBIDA
Family Amoebidae
Amoeba Ehrenberg. <A. proteus Leidy; A. guttula Duj.; A.
Sphaeronucleus Greef; A. striata Penard; A. radiosa Ehr.; A.
saphrina Penard.

The species of this genus were not common in any locality.
Material from Myoho-in Temple grounds, Kyoto, furnished the
best examples. Large individuals of A. radiosa were taken from
Lake Hakone.

Hyalodiscus Hertwig and Lesser. H. rubicundus H. and L.

3ut one individual was observed. A very typical form, red-
dish-brown in color. From a rice field, Kyoto.

Arcella Ehrenberg. A. vulgaris Ehr.; A. discoides Ehr.; A. costata
Ehr. ; A. arenaria Greef.

Of the above species A. vulgaris is the more widely distributed
in Japan. Lake Chuzenji and the region of Kyoto furnished the
best material. |

Centropyxis Stein. C. aculeata Stein.

Found in all localities. Very abundant in Lake Hakone. Great
variation in size occurs in this species and some very large forms
were observed.

Pixidicula Ehrenberg. P. cymbalum Penard.

A species rarely observed. Found in material from Lake
Hakone.

Lecquereusia Schlumberger. L. spiralis Ehr.; L. modesta Rhumbler.

These two speices are widely distributed in Japan, the former
being much more abundant. In the typical L. spiralis the aperture

JAPANESE PROTOZOA 95

is usualy directed obliquely toward one side with a prominent
hump at the outer base of the neck. In the common form in
Japan the aperture is directed almost straight forward, in very
rare cases there being a slight prominence at the base of the neck.
Common in Lake Hakone. Typical examples of L. modesta were
found in lakes on Mt. Rokkozan.

L. epistomium Penard, a common species of the high lakes of
Colorado, was not observed in Japan.

Difflugia Leclerc. D. pyrifornus Perty; D. lobostoma Leidy; D.
constricta Leidy; D. acuminata Ehr.; D. tuberculata Wallich;
D. lebes Penard; D. bacillariarum Perty; D. elegans Penard.

Of the species of Difflugia in Japan, D. elegans is apparently
the most common. It is widely distributed and shows a great
range of variation. D.Jebes, not uncommon in some of the lakes
of Colorado, was observed but onee, in material from the bottom
of Lake Hakone.

Pontigulasia Rhumbler. P. spectabilis Penard.

But one individual observed: From Lake Hakone. A very
typical form.

Quadrulella Cockerell. Q. symmetrica Schultze; Q. symmetrica
var. curvata Wailes.

Very typical forms of the species were taken from shallow lakes
on Mt. Rokkozan. The variety, observed but once, was found in
Lake Hakone.

Nebela Leidy. N. collaris Leidy; N. crenulata Penard; N. hippo-
crepis Leidy; N. triangulata Lang.

In the material collected in Japan species of Nebela were very
rare.

There can be no reason to believe, however, that the genus is
not well represented in that country. One individual of the rare
species, N. hippocrepis, was found in material from Mt. Rokko-
zan. In the ooze from the rocks along the shore of Lake Hakone
and from the border of a shallow lake on Mt. Rokkozan was
found a species which is here listed under the name JN. triangulata
Lang.

The Japan species resembles, in some particulars, Nebela bipes
Carter, as described in Clare Island Survey, Part 65, by Wailes

96 EDMONDSON AND KINGMAN

and Penard, and may represent an intermediate form between
N. triangulata and N. bipes.

In the Japan form the shell is very transparent, compressed,
irregular in outline with the fundus region inflated in an asymmet-
rical manner. The aperture is slightly oval.

Great variation exists in the form of the shell and in the
arrangement of the plates. In some the plates are circular or
oval, distinctly separated from each other with the ground sub-
stance of the shell intervening. In others the plates are closely
crowded together and very irregular in outline, while in some
the plates are regular in outline but distinctly overlap each other.

The irregular inflation of the fundus is a characteristic feature.
Usually the posterior lateral borders are expanded into lobes
of variable size. In some these prolongations are pointed as in
N. bipes, but more often they are blunt or rounded. Occasionally
the fundus is truncated posteriorly, sometimes it is strongly con-
cave. The extensions of the fundus are seldom uniform on the
two sides of the shell and are never the same in two individuals.
Usually the narrow view of the shell presents an irregular outline.
The compression of the shell is seldom uniform, but is always
stronger at the fundus border.

The size of the Japanese form ranges from 80 to 100m in
length, including the prolongations of the fundus; from 60 to 80p
in breadth of fundus and from 28 to 60n in the long diameter of
the aperture.

No living individuals were observed.

Heleopera Leidy. H. picta Leidy.

Material from Mt. Rokkozan furnished the only species of the
genus observed. Under high power the plates are seen to be
circular, slightly overlapping. Little foreign material is attached
to the shell.

Phryganella Penard. P. hemisphaerica Penard.
Frequently observed in many localities.
Campascus Leidy. C. dentatus, sp. nov.

In 1877 Leidy discovered Campascus cornutus in China Lake,
Wyoming, at an altitude of 10,000 feet. Apparently the species
has not been observed since that time.

~~

i ee ee

ai. a!

JAPANESE PROTOZOA 97

More recently Penard described two species of the genus,
Campascus triqueter and Campascus minutus, from the deep
lakes of Switzerland. In both species described by Penard the
fundus is without the horn-like prolongations of the form ob-
served by Leidy. Campascus minutus was reported by Wailes in
1912 from the New York water-supply drawn from Croton Lake
Reservoir.

The form under consideration, which is apparently a new
species, was found in the ooze taken from the rocks along the
shore of Lake Hakone, Japan, in August, 1912.

The description follows: Shell of yellowish, chitinoid material
similar in general outline to Campascus cornutus. Under high
power the shell has the appearance of being distinctly punctate.
In some individuals the punctae are arranged in a regular diagonal
manner, in others there is no regularity about the arrangement.
In no specimens examined can outlines of plates be detected even
with the oil immersion lens.

The neck is short and sharply bent, nearly at right angles to
the long axis of the shell. The circular aperture is bordered by
a thin delicate membrane of approximately 4» in breadth.

A number of short, blunt, tooth-like prolongations are present
on the posterior border of the fundus. From three to seven of
these processes are usually present. They vary in size and when
numerous give an irregular, crenulated appearance to the posterior
edge of the fundus, when the broad side of the shell is viewed.

In Leidy’s species the two horns are directed laterally and pos-
teriorly, their tips not projecting beyond the posterior border,
giving the fundus a rounded outline when the narrow side of
the shell is observed. In this species the teeth-like points are
directed backward and project beyond the border, giving the
fundus the appearance of terminating in a spine when the narrow
side of the shell is seen.

Leidy records the size of Campascus cornutus as ranging from
0.112 mm. to 0.14 mm. long by 0.18 mm. broad.

This species of Japan is much smaller. The length of the
shell, including the spines and the collar about the aperture,
ranges from 60 to 80n. Breadth of fundus from 50 to G6y.

98 EDMONDSON AND KINGMAN

Greatest thickness, narrow view, 284. Aperture 12 in diameter.

The living organism was not observed.
Paulinella Lauterborn. P. chromatophora Lauterborn.

Empty shells of this very minute form were found in material
from the bottom of Lake Hakone and also from shallow lakes on
Mt. Rokkozan. The shell is composed of five longitudinal rows
of plates and possesses a short neck. The Japan form is very
typical.

Cyphoderia Schlumberger. C. ampulla Ehr.; C. ampulla var. papil-
lata Wailes.

The species is very common in Lake Hakone and was found in
other localities. Considerable variation in size and also in the
arrangement of plates occurs. The plates are usually placed in
diagonal rows, but this regularity is not always maintained.

The variety was observed but once and that in material from
Lake Hakone.

Sphenoderia Schlumberger. S. lenta Schiumb.

Very widely distributed and also very common in Japan. The
only species of the genus to be determined.

Euglypha Dujardin. £. alveolata Duj.; E. brachiata Leidy;
E. filifera Penard; E. laevis Perty; E. ciliata Ehr.; E. armata
Wailes.

A few species of this genus are very abundant in Lake Hakone
as well as in other localities. Two species, E. filifera and E.
ciliata, were rarely observed, the others mentioned are common.

Assulina Ehrenberg. A. seminulum Ehr.

Observed in material from Kyoto. A very typical form, choco-
late-brown in color.

Plagiopyxis Penard. P. callida Penard.

Indentified in material from Kyoto. Not common.

Trinema Dujardin. T. enchelys Ehr.; T. lineare Penard; T. cam-

planatum Penard.

The genus represented by T. enchelys is very common in many
localities. The other two species were rarely observed.

.
——— ———< =.

JAPANESE PROTOZOA 99

Class ACTINOPODA Subclass HELIOZOA
Order APHROTHORACIDA

Actinophrys Ehrenberg. A. sol Ehr.

Observed in great abundance at Kyoto; rarely seen in other
localities.

The following list is a record of the species of Mastigophora
and Infusoria identified in material taken from the fresh waters
of Japan. Flagellates and ciliates are very abundant in that
country, as elsewhere, and the small number of species here listed
indicates brevity of observation rather than any dearth in pro-
tozoan fauna. The remarkable thing to be noticed is the identity
of the Japanese forms with our common American species.

Subphylum MASTIGOPHORA:
Class ZOOMASTIGOPHORA

Order HETEROMASTIGOPHORA

Notosolenus Stokes. A’. orbicularis Stokes.
Anisonema Dujardin. A. acinus Duj.
Order Monapipa
Anthophysa Bory d. St. Vincent. A. vegetans Mill.
Order EUGLENIDA
Euglena Ehrenberg. F. viridis Ehr.; E. deses Ehr.; E. acus Ehr.
Phacus Dujardin. P. pleuronectes Mill.; P. longicaudus Ehr.
Trachelomonas Ehrenberg. T. hispida Stein; T. volvocina Ehr.;
T. armata Stein.
Astasia Ehrenberg. A. trichophora Ehr.
Distigma Ehrenberg. D. proteus Ehr.

Subphylum INFUSORIA:
Class CILIATA
Order HoLorricHIDA
Coleps Ehrenberg. C. hirtus Ehr.
Lacrymaria Ehrenberg. L. olor Mill.

Lionotus Wrzesniowski. L. fasciola Ehr.
Dileptus Dujardin. D. gigas C. and L.

100 EDMONDSON AND KINGMAN

Chilodon Ehrenberg. C. cucullulus Mill.

Nassula Ehrenberg. N. oronata Ehr.

Loxocephalus Ehrenberg. L. granulosus Kent.
Cinetochilum Perty. C. margaritaceum Ehr.

Frontonia Ehrenberg. F. leucas Ehr.

Paramaecium Miiller. P. caudatum Ehr.; P. bursaria Ehr.
Cyclidium Ehrenberg. C. glaucoma Ehr.

Pleuronema Dujardin. P. sp. (undetermined).

Order HETEROTRICHIDA

Spirostomum Ehrenberg. S. ambiguum Ehr.
Stentor Oken. S. caeruleus Ehr.; S. polymorphus Ehr.
Gyrocoris Stein. G. oxyura Stein.

Order HypotricHIDA

Oxytricha Ehrenberg. O. pellionella Mill.
Stylonychia Ehrenberg. S. notophora Stokes.
Euplotes Ehrenberg. E. charon Mull.
Aspidisca Ehrenberg. A. costata Duj.

Order PERITRICHIDA

Vorticella Linnaeus. V. sps.
A number of undetermined species were observed.
Cothurnia Ehrenberg. C. sp. (undetermined).

Class SUCTORIA

Sphaerophrya Claperéde and Lachmann. S. magna Maupas.
Washburn College, Topeka, Kansas.

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102 EDMONDSON AND KINGMAN

EXPLANATION OF FIGURES

Pirate Il

Fig. 1, Lecquereusia spiralis Ehrenberg; X 272. From Lake Hakone.
Fig. 2, Lecquereusia spiralis Ehrenberg; X 257. From Lake Hakone.
Fig. 3, Lecquereusia spiralis Ehrenberg; 272. From Lake Hakone.
Variations of the species common in Japan.
The aperature is directed almost straight.
Fig. 4, Lequereusia modesta Rhumbler; X 225. From Lake Chuzenji.
Fig. 5, Difflugia bacillariarum Perty; X 225. From Lake Hakone.
Fig. 6, Difflugia elegans Penard; X 195.
Very common. Individuals observed ranged from 60-194” in length.
Fig. 7, Quadrulella symmetrica var. curvata Wailes; X 427.
Near the aperture the plates become small and irregular. Rarely ob-
served. From Mt. Rokkozan.
Fig. 8, Nebela hippocrepis Leidy; X 108.
Broad view of a shell. From Mt. Rokkozan.
Fig. 9, Nebela hippocrepis Leidy; X 198. Narrow view of same.
Fig. 10, Nebela triangulata Lang; X 325.
Broad view of a shell. From Lake Hakone.
Fig. 11, Nebela triangulata Lang; X 378. From Lake Hakone.
Fig. 12, Nebela triangulata Lang; X 354. From Lake Hakone.
Fig. 13, Nebela triangulata Lang; X 315. From Lake Hakone.
Fig. 14, Nebela triangulata Lang; X 325. Narrow view of a shell. From
Lake Hakone.
Variation in the shape of the fundus and in the arrangement of the plates
shown in these figures.
Fig. 15, Campascus dentatus, sp. nov.; X 370.
Broad view of a shell with the posterior border of the fundus provided
with numerous teeth-like prolongations. From Lake Hakone.
Fig. 16, Campascus dentatus, sp. nov.; X 390. Broad view of another
shell. From Lake Hakone.
Fig. 17, Campascus dentatus, sp. nov.; X 390. Broad view of another
shell. From Lake Hakone.
Fig. 18, Campascus dentatus, sp. nov.; X 390. Narrow view of same.
From Lake Hakone.

2
2

Fig. 19, Paulinella chromatophora Lauterborn; X 1050. From Lake Hakone.

apanese Protozoa

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Pirate IIT.

102

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EXPERIMENTAL AMITOSIS IN ONION ROOT TIP
H. E. Jorpan

INTRODUCTION

My recent study of amitosis in the lining epithelium of the
epididymis of the mouse,’ and in certain other ciliated epithelia,
led me to the tentative conclusion that the fundamental causative
factor here involved is the partition and consequent destruction of
the centrosome in the formation of “basal granules” from which
the cilia grow. The suggestion was made that direct cell division
is proximally related to factors inducing loss of integrity or im-
pairment of function of the centrosome. Such factors are vari-
ous, including those originally and recently sugested, namely, in-
tense secretory activity, and disturbed metabolic conditions charac-
terized as “degenerative” (Flemming,? Ziegler? vom Rath,* et
cetera), lack of adequate nutriment (Child,® Patterson®), insufh-
cient supply of oxygen (Wieman’).

To test experimentally this hypothesis a series of experiments
were made by growing onion roots in water to which ether was
added. Two assumptions, apparently legitimate, are involved:
1) the onion cell has an analogue (indiscernible) of the centrosome
of animal cells, similarly concerned in the formation of the mitotic
spindle; 2) ether, and other anesthetics, may be expected to produce
a “stupefying’ effect upon the “dynamic center” of the cell thus
enforcing a direct method of actual multiplication.

The sole result claimed for the investigation is the fact that
onion roots continue to grow vigorously in water to which small
quantities of ether are added, and that many of the cells are in
various phases of undoubted direct division. No originality is
claimed either for the hypothesis or for the experimental procedure.
I believe, however, that the facts adduced from the study of the
epididymis of the mouse offer the strongest histologic evidence yet
reported in support of the hypothesis. The experiments were under-

104 H. E. JORDAN

taken simply to test this hypothesis. The same idea must have
suggested the experiments of Pfeffer and Nathansohm ® with spiro-
gyta grown in water with ether, and of Wasielewski® with Vicia
faba roots grown in chloral hydrate solution.

Nathansohm (1900) claims to have induced amitosis in spiro-
gyra and the desmid closterium by transferring karyokinetically
dividing filaments to a 1% solution of ether in water. Roots of
Phaseolus, Lupinus, Phalaria and Marsilia were similarly treated,
but without success in changing cell-division from the indirect to
the direct method. Treatment of growing roots of Vicia faba with
a 0.7% chloral hydrate solution for various periods is claimed by
Wasielewski (1902-1904) to change the majority of cell divisions
from mitotic to amitotic. Némec?® (1904), however, interprets
the cell pictures. (in roots of Vicia faba, Allium cepa and Pisum
sativum, treated with 0.75% chloral hydrate solution) in terms of
nuclear fusions, and disputes their amitotic significance. My ex-
periments with onion root tip grown in ether solution, on the con-
trary, show unmistakable amitosis. However, in view of the
fact that the roots grew vigorously, and that mitoses are always
exceedingly rare, and in several instances totally lacking, indisputable
healthy amitotic divisions (i. e., apparenty non-degenerative) are
unexpectedly relatively rare. 7

DESCRIPTION OF THE EXPERIMENTS

The procedure was simply to place an onion, with stem and
roots either absent or just appearing, into the mouth of a small
jar so selected for size that the root pole of the bulb was immersed
in the solution to a depth of about half an inch. The experiments
with controls consisted of three series, with summarized results
as follows:

First Series

a) Bulb in 4% alcohol solution—Neither stem nor roots appeared.

b) Bulb in water unchanged for four days.—Both stem and roots
appeared. Of 5 roots examined, two were disintegrating; the
remainder showed occasional mitoses in the plerome cells and
several doubtful instances of degenerative amitoses in the
pleriblem cells.

Ae

a)

EXPERIMENTAL AMITOSIS 105

Bulb in water to which a few drops of ether were added thrice
daily for two days.—Stem barely developed, many vigorously
growing roots appeared. Of 6 roots examined only two showed
mitotic figures, limited to the plerome; and all showed occa-
sional direct divisions among the periblem cells.

Second Series (three to eight tips each)
Bulb with roots just showing; in water from 10 o’clock A. M.
to 4.30 P. M.—Normal; many mitoses both in plerome and
periblem; also a few in dermatogen.
Bulb in water, unchanged for 4 days.—Mitoses still frequent ;
a few tips distintegrating.
Bulb in water changed twice daily for 4 days——Normal; nu-
merous mitoses.
Bulb in moist cotton 1 day.—Tips of young development;
neither mitoses nor amitoses.
Bulb in moist cotton 2 days.—Distintegrated.
Bulb in water unchanged for one week.—Normal; many
mitotic figures.
Bulb in water + ether, changed twice daily for 1 day.—Neither
mitoses nor amitoses.
Bulb in water + ether, changed twice daily for 3 days—No
mitoses; many amitoses.
Bulb grown in cotton moistened with water and ether for 4
days.—No growth of either stem or roots.

Third Experiment
Bulb in water to which a small amount of ether was added
three times daily for 5 days, and allowed to remain in the un-
changed water for 2 days longer.—Normal; numerous mitotic
figures.
The roots were fixed in Flemming’s strong solution.
The sections were cut at from 5 to 10 microns, and stained in

iron hematoxylin, with and without eosin counterstain.

Description of Normal Root
A brief description of normal conditions seems desirable before

proceeding to an analysis of the experimental results of the several

106 H. E. JORDAN

series. The description is based upon an excellent preparation
previously made and used for demonstrating mitotic figures in class-
room work.

The root-tip contains characteristic cells in the several regions:
dermatogen, periblem, plerome, and root-cap. The dermatogen con-
sists of from 2 to 5 layers of very long rectangular cells, with the
longest axis parallel to the long axis of the root. The nucleus is
pale and finely granular, with one or several nucleoli. The cyto
plasm contains large vacuoles and numerous dcep-staining spherical
granules of various sizes (mitochondria). Some of the cells are in
process of indirect division.

The periblem consists of from 7 to Io rows of small rectangular
cells surrounding the central plerome. The longest axis of these
cells is generally perpendicular to that of the dermatogen cells. The
nuclei are pale staining, finely granular or delicately reticular,
mostly with two nucleoli frequently surrounded by a clear halo. The
cytoplasm is considerably vacuolated and contains granules similar
to those described for the dermatogen. The cells of the layers, more
particularly from the third inward and from just above the upper

- :

Fig. 1. Sketch of periblem cell from normal root showing early process in

constriction of nucleolus. Abundant earlier and later phases also ap-
pear. The vacuolated cytoplasm contains deeply staining granules (mito-

chondria). The nucleus is pale-staining, finely granular, or delicately
reticular.

Fig. 2. Typical rectangular bi-nucleolate cell of periblem of normal root.
One of the nucleoli is surrounded by a clear halo, probably a fixation
artefact.

Figs. 3 and 4. Periblem cells from root undergoing early degenerative
changes. The cytoplasm contains huge vacuoles, and lacks mitochondria.
The nuclear wall is irregular and apparently suffering an equatorial con-
striction closely simulating, if not actually, a direct division. Excessive
number of nucleoli, i. e., more than two is a common condition in the
cells of these tips.

EXPERIMENTAL AMITOSIS 107

limit of the root-cap, show large numbers of nucleoli at all phases of
constriction and separation in the process of the formation of
binucleolate nuclei (Figs. 1 and 2). The appearance is exactly that
of Remak’s classic illustration of the initial step in amitosis. How-
ever, the further steps, viz., nuclear and cytoplasmic fission, no-
where appear; and mitoses at all phases are very abundant.

The plerome consists of a variable number of layers. The
cells are long rectangular, relatively huge rectangular, and approxi-
mately square, in shape. The nuclei are large oval or spherical,
mostly with deep-staining coarse reticulum, and with one or two
chromatic nucleoli, sometimes appearing achromatic. The cytoplasm
contains smaller vacuoles than in the periblem and dermatogen
cells and relatively fewer mitochondria. Mitoses are most abundant
in this portion. Two chromatic nucleoli are frequently present even
at the segmenting spireme stage, of irregular oval shape but with
sharp contour.

The root-cap contains larger and smaller polyhedral cells with
enormous vacuoles in the cytoplasm and very sparse mitochondria,
and pale granular nuclei with one or generaly two nucleoli. These
cells are only rarely seen in mitosis in this preparation.

The points of special significance in normal root tip for this
study are: 1) absence of amitoses; 2) great abundance of mitoses,
especialy in plerome; 3) nucleolar constriction in, and binucleolate
character of, the periblem cells ; 4) presence of mitochondrial gran-
ules (an index of virility); and 5) general pale-staining, finely
granular, character of the periblem nuclei.

ANALYSIS OF THE First SERIES OF EXPERIMENTS

The non-appearance of roots or stem on the bulb in the 4%
solution of alcohol is interpreted to mean that this solution was
too strong to permit development. Since the experiment was not
repeated with weaker solutions the interpretation must be tentative ;
but since this is the only complete failure of development in the
several series (save one with bulb in cotton moistened with ether
solution) it is very plausible. The reason for not further experi-
menting with alcoholic solutions at this time was the fact that posi-
tive results were obtained with the ether solutions.

108 H. E, JORDAN

The roots of the bulb grown in water unchanged for four days
differed somewhat from normal, and several were in the later stages
of disintegration. Mitoses are numerous in the plerome cells, but
practically absent in the periblem cells. Mitochondrial granules
are relatively sparse and pale-staining. Relatively more cells have
two nucleoli; a number have three (Fig. 3) and even four nucleoli.
The periblem nuclei are frequently irregular in outline, with numer-
ous blunt processes; a few may possibly be in process of amitosis
(Figs. 3 and 4). On this bulb the stem grew very vigorously.
Cytologic appearances must probably be interpreted in terms of
early degeneration, involving possibly slight amitotic division.

Roots of bulbs grown in a 3% aqueous solution of ether grew
very vigorously, while the stem developed but little. This result
seemed to indicate a stimulating effect to root formation, causing
indirectly a retardation of stem growth. The jar with bulb was
covered under a larger jar to prevent excessive volatilization of
the ether. When uncovered as in the later experiments this differ-
ential growth of roots and stem was not so evident. The roots
have a perfectly healthy appearance, in section in all respects like
the normal except that mitoses are almost, and in some tips entirely,
lacking even in the plerome cells. Amitoses are abundant in the
periblem cells (Figs. 5 and 6), usually appearing in groups of four
or eight. Many of these nuclei have an irregular contour with short
pseudopodial processes like in the foregoing set of roots. Only
rarely can the final process of the formation of a wall between the

Fig. 5. Periblem cell from root grown in 3% ether solution (first set of ex-
periments), showing early stage in amitotic division.

Fig. 6. Similar cell, showing a later phase of direct division, including the
formation of a new cell membrane between the separating daughter nuclei.
The cytoplasm contains large vacuoles and lacks mitochondria, in which
respects it resembles a degenerating cell.

Fig. 7. Diagram of frequent, type of nucleus showing character of initial
phase of nuclear direct division.

EXPERIMENTAL AMITOSIS 10g

amitotic moieties be seen (Fig. 6). Many nuclei show a sharp cut
at one point on the surface (diagram, Fig. 7), the initial step in
the nuclear constriction. This appearance is practically lacking in
the previous set.

The number of bi-nucleolate cells does not seem relatively
larger than in the normal or degenerating tips. However, multi-
nucleolate cells are absent. No significance can be attached to the
bi-nucleolate nuclei, from the standpoint of amitosis, since these
are almost equaly abundantly present both in normal and degenerat-
ing tips. Still, when followed by nuclear fission, as occasionally in
tips grown in ether solution, the nucleolar fission represents the
initial step in amitosis.

There remains no question, I believe, that true amitosis occurs
in these tips, and as the result of ether in the water; but whether
this result is direct (i. e., specific, the ether acting anzsthetically
upon the astral system) or indirect (1. e., due to abnormal environ-
mental conditions produced by the ether), must remain uncertain.
However, the fact that undoubted amitoses are relatively much more
abundant in the tips grown in water with ether than the doubtful
ones in the unchanged water speaks in favor of a direct retarding
influence to centrosomal activity. This conclusion is further
strengthened by the fact that the roots grew vigorously; and that
no cytologic indications of degeneration, beyond a number of irreg-
ular nuclei, appear. The result is disappointing, however, in that
the amitoses are unexpectedly rare in view of the rapid growth and
the absence of mitoses. It cannot be asserted with full surety that
growth was entirely by amitosis, since a few mitoses appear in the
plerome cells; and, moreover, the possibility remains that the tips
were cut at the interval between mitotic waves. But if this were
the true explanation of the absence of mitoses, not all of the roots
would be expected to show substantially the same condition. It
seems then that amitosis plays a considerable part in the growth
of root tips in water with ether, and that it does not necessarily
signify degeneration since the tips after two days are still vigorous
and normal. That the complete amitotic process is consummated,
though the final step is difficult to demonstrate, is further indicated
by the complete absence of bi-nucleolate cells.

110 H. E. JORDAN

It seemed possible that the degeneration and incidental prob-
able amitosis of the roots grown in unchanged water might be due
to the presence of the extract of onion held by the water; also,
this method of sprouting bulbs is so far from normal that only
degeneration phenomena (including amitosis) could perhaps be
expected. To check these uncertainties, and to further test the
influence of ether on mitotic proliferation, the second set of experi-
ments was devised.

SECOND SERIES OF EXPERIMENTS

Growth in moist cotton gave wholly negative results as the
above summary shows. This evidently makes a most unfavorable
growth medium.

The roots grown in water changed twice daily were normal.

Those grown in unchanged water, even after an entire week,
were still vigorous and essentially normal. Thus extract of onion
appears to have no inhibitory or deleterious effect, at any rate if
allowed to volatilize freely. In the previous experiment the bulb
fitted more closely into the mouth of the jar. The cause of the
degeneration in several roots in the first series of experiments
remains obscure.

,9and 10. Periblem cells from roots grown in ether solution (second
set of experiments), showing three stages in the nuclear constriction;
in Fig. ro a new cell membrane is forming and has progressed into the
depth of the constriction on the right.

Figs. 11 and 12. Periblem cells from the same set showing initial and final
stages in direct division.

Figs.

The bulb grown in the ether solution again sprouted roots,
showing the identical condition described in the first series (Figs.
eto! 12):

EXPERIMENTAL AMITOSIS Itt

THIRD EXPERIMENT

The third experiment was planned for the purpose of testing
whether tips induced to presumable amitotic growth by ether could
again revert to mitotic growth on resumption of normal conditions.
With this end in view a bulb was placed as above in a 3% solution
of ether (ether being added thrice daily) for 5 days, and then al-
lowed to continue development for two more days’ without further
addition of ether. Six roots were cut and fixed at the end of the sev-
enth day. All except two which were distintegrating were appar-
ently normal and the majority contained abundant though frequently
atypical mitotic figures. The result of this experiment indicates
that the direct cell division in the roots developing in the ether
solution does not necessarily lead to degeneration. Significantly,
too, several of the roots showed a few bi-nucleate cells; as if the
amitotic process had been halted short of consummation at the dis-
appearance of the ether from the water.

CoNCLUSION

Onion root-tips grown in appropriate solutions of ether
(approximately 3% in water) show an almost complete absence of
mitotic figures, and a considerable number of periblem nuclei in
process of direct division. Amitosis is not present in roots grown
in pure water, while mitoses are abundant. Ether would appear
to have a “stupefying” effect upon the astral apparatus, preventing
karyokinetic cell division. Certain appearances (e. g., pale periblem
nuclei of irregular contour) suggest degeneration ; but roots similarly
treated with ether for several days and then allowed to develop for
several more without further addition of ether, proceed to grow
normally and show abundant mitoses. Thus the amitosis seems
capable of reversion to mitotic type. These results confirm those
of Pfeffer and Nathansohm with spirogyra and closterium, and
those of Wasielewski with roots of Vicia faba grown in chloral
hydrate solution. Wasielewski, however, could only report a ma-
jority of direct, as compared with indirect, divisions in Vicia faba;
and my own results are disappointing in that, in view of the almost
total absence of mitoses, undoubted amitoses are rare. Concerning a
certain small number, however, there can be no doubt. It must

112 H, E. JORDAN

still remain uncertain whether these represent early indications of
a reversible degenerative process due to unfavorable environment,
or the result of a direct influence on the part of the ether upon the
achromatic spindle complex. The apparent selective action upon
the periblem cells also remains obscure. Whether degenerative in
significance, as seems more probable in view of certain cytologic cor-
respondences above detailed, or otherwise, the amitosis present in
etherized roots seems clearly the result of a superior sensitiveness
on the part of the centrosome and the astral associates to untoward
environmental influence.

LITERATURE

i. Jorpan, H. E.

Amitosis in the epididymis of the mouse. Anat. Anz., Bd. 44. 1913.
FLEMMING W.

Neue Beitrage Zur Kentniss der Zelle. Archiv. mikr. Anat., Bd. 37.

1801.

ZIEGLER, H. E.

Die biologische Bedeutung der amitotischen Kerntheilung in Tier-

reiche. Biol. Centrbl., Bd. 11. 1801.
4. Vom Ratu, O.

Ueber die Bedeutung der amitotischen Kernteilung im Hoden. Zool.
Anz., Bd. 14. 1891.

rs)

sd

5. Crip, C. M.
Amitosis as a factor in normal and regulatory growth. Anat. Anz.,
Bd. 30. 1907.

6. Patterson, J. T.
Amitosis in the pigeon’s egg. Anat. Anz., Bd. 32. 1908.
7. Wireman, H. L.
A study in the germ cells of leptinotarsa signaticollis. Journ. Morph.,
vol. 21, No. 2. rgto.
8. NATHANSOHN, A. .
Physiologische Untersuchungen tiber amitotische Kernteilung. Jahr.
Wiss. Bot., 35:1. 1900.
9. WASIELEWSKI, WALDEMAR VON.
Theoretische und experimentelle Beitrage Zur Kentniss der Amitose.
Jahrb. Wiss. Bot. 1 Abschnitt, 1902, Bd. 38: Heft. 3. II Abschnitt,
1904, Bd. 39: Heft. 4.
10. NEmEc, B.
Ueber die Einwirkung des Chloralhydrats auf die Kern-und-Zell-
theilung. Jahr. Wiss. Bot. 39:4. 1904.

Anatomical Laboratory,
University of Virginia.

—,

SUMMARIES IN MICRO-BIOLOGY

For some months the Secretary has been planning to secure for this Journal and its
Department of Summaries, a series of papers from biologists dealing with the chief groups
of microscopic plants and animals. It has not been the purpose to present a complete
survey of any of the groups. The wish has been rather to bring together in one article
a statement of the following things:—-general biology, the method of finding, the methods
of capture and of keeping alive and cultivating in the laboratory; how best to study; the
general technic; the most accessible literature; and a brief outline of the classification,
with keys for the identification of at least the more representative genera and species of
the micro-organisms likely to be found by the beginning students in the United States.

It has been felt that the getting together of such data as this, while not a contribution
to science, would be a contribution especially to isolated workers and to teachers and stu-
dents in the high schools and smaller colleges.

Papers have already appeared treating the aquatic Oligochetes, the Melanconiales
and the Rusts. The following is the fourth paper of the series. It is proposed to have
such synopses from time to time until the more common American species of such groups
as the following have been covered: The Blue-green Algae, Conjugating Algae, Diatoms,
other Green Algae, Downy Mildews, Yeasts, Powdery Mildews, Hyphomycetes, Smuts,
Rhizopods, Infusoria, Turbelaria, Bryzoa, Water Mites, Entomostraca, etc.—[Editor.]

THE BLACK MOULDS (Mucoraceae)
Leva B. WALKER

The Mucoraceae receive the name “Black Moulds’ from the
fact that in a number of the most conspicuous genera the fruiting
bodies and older hyphae are dark colored or “black.” A young
growth of any of these fungi is, however, always colorless. They
are largely saprophytic and are found abundantly on decaying or-
ganic matter, especially on dung, and in the soil as common soil
fungi. The parasitic forms live upon other mucors or upon basidi-
omycetes. Almost all are easily cultivated on bread, on dung or
on ordinary culture media (either directly or by culturing the host
and parasite together) if bateriological apparatus is at hand.

1. Plant Body

The Mucoraceae form a well-defined group. The plant body,
as is true for almost all phycomycetous fungi, is siphonaceous, cross
walls being few or none, always coenocytic. The protoplasm in
young growing filaments shows characteristic movement in a “gla-
cier-like” fashion. The mycelium may be superficial, developing
rhizoids that penetrate the substratum, or it may develop almost
entirely within the substratum.

Il4 LEVA B. WALKER

2. Sexual Reproduction

The most uniform character in the Mucoraceae is the produc-
tion of zygospores. The zygospores are developed by the union of
cells from the same filament or from other filaments. First very
much enlarged branches (progametes) branch off from two adjacent
filaments (Plate IV, Fig. 1, a), a cell is cut off from the end of each
progamete, these end cells becoming the gametes (Fig. 1, b (x)),
the enlarged supporting cell being called the suspensor (Fig. 1, b
(y)). The gametes fuse (Fig. 1, c) by the absorption of the wall
between the gametes, and a thick wall is built up around the fertil-
ized cell, which is then called the zygospore. While zygospores are
known for most of the species of the Mucoraceae, they are rarely
found. When they do appear in cultures they appear in great abun-
dance. The conditions necessary for the production of zygospores
has been the subject of many researches, the most extensive of
which are those of Blakeslee, who has shown definitely that the
Mucors are separable into two distinct types which he terms hhomo-
thallic and heterothallic. 1n homothallic forms the gametes are pro-
duced from branches of the same filament and zygospores are
usually abundant in nature wherever the fungus is found. In the
heterothallic forms the gametes must be produced from separate
strains (termed for convenience + and —) of the fungus which
often differ so greatly in appearance that they might easily be taken
for separate species or at least varieties. Extensive experiments
made by planting + and — strains of many Mucors together show
that when a + and a — of different species are planted together the
progametes will form (but will not form zygospores), but that when
a + and a + or a — and a — are planted together no such proga-
metes are formed. The + strain is usually a little more vigorous
than the —, and in some cases the gametes are larger, so we can well
think of the + strain as female and the — as male. In heterothallic
forms the zygospores are rarely found, as either the + or the — 1s
rare. The germination of the zygospores has been observed in
relatively few cases, but where observed takes place much as shown
in Fig. 3, 1. Zygospores germinate only after a resting period.

3. Asexual Reproduction

SPORANGIOSPORES—The sporangiospores (usually referred to

THE BLACK MOULDS Its

simply as spores) are typically formed in sporangia which are pro-
duced on erect well-differentiated hyphae known as sporangiophores.
The sporangia are separated from the sporangiophores by a cross
partition which usually rounds up into the sporangium, forming a
columella (Fig. 1, {). The outer wall of the sporangium is usually
delicate, dissolving in water, or fracturing easily so that in exam-
ining material in water mounts under the microscope the spores
are often seen surrounding the columella, the only remnants of the
outer wall being seen at the point of its attachment to the columella.
The sporangia are in most cases evident and many spored, but in
one tribe they are reduced seemingly to one spored sporangia and
the one celled sporangia are usualy called “conidia.” In another
tribe the spores are arranged in a single row in a sporangium which
breaks down so readily that it is rarely distinguishable and gives the
appearance of a chain of “conidia.”

The asexual reproduction as above described is always the
most abundant form of reproduction and some forms are only known
by their asexual reproduction. For this reason the keys which will
follow are based upon the sporangial reproduction. The spores
are capable of germination at once (Fig. 3, above i), but will usually
remain viable for an indefinite period.

Azycospores—These are often found in cultures of Mucors,
they usually appear much like the zygospores, but are formed with-
out the union of gametes.

CHLAMypbosporEsS—Thick-walled spores, known as chlamydo-
spores, are often found in the main vegetative filament, or on the
ends of filaments (Fig. 3, 1, k). They may germinate at once or
after a resting period.

4. Systematic

The Mucoraceae comprise a group of about forty described
genera and over three hundred described species. Probably most
of these are present in our flora. Only a few of the most typical
and abundant genera and species will be mentioned here.

116 LEVA B. WALKER

KEY TO THE TRIBES OF MUCORACEAE.

A. Reproduction asexually by spores contained in evident sporangia.
I. Columella present.
a. Sporangia many-spored, generally of one kind or if two kinds
the smailer irregularly disposed on the main sporangiophore.
1. Membrane of sporangium easily dissolved or fractured

a AY, Os, iceeme eral afc Aue AS Wen tl, sees Pst ees ae eee Mucoreae (I)
2. Membrane of sporangium usually solid, persistent...........
Pe eee ee ae Tee eee Piloboleae (II)

b. Sporangia of two kinds, the larger many spored, the smaller
few spored and formed on the ends of regularly branched
SUGFan CIO PNOLeS es ciao ete e s </<8' se eeree Thamnidiae (III)

II. Sporangia without a columella.................. Mortierelleae (IV)

B. Reproduction asexually by so-called “conidia” produced either solitary
or in chains.

I. “Conidia” solitary, produced upon the ends of usually much branched

SEOGMNGNROLES Eats . ck o tees See een vee ee Ue Chaetocladiae (V)
II. “Conidia” in chains, produced in head-like masses upon sterigmata
at the ends of “conidiophores”............ Cephalideae (VI)

I. TRIBE MUCOREAE

Sporangia generally of one kind with a columella and a mem-
brane that dissolves or fractures easily. Smaller sporangia (sporan-
gioles) with persistent membranes occur very rarely and in such
cases are disposed without order upon the main sporangiophore.
Zygospores naked or surrounded by appendages.

Key To GENERA.

I. Sporangiophores fasciculate on a rhizoidiferous stolon.

1. Sporangia globose; zygospores unprotected................- Rhizopus

2. Sporangia pyriform; zygospores protected by flexuous unbranched

outgrowths from the suspensors.............. Absidia

II. Sporangiophores emerging solitary from the mycelium, no rhizoidiferous
stolons.

1. Sporangiophores unbranched, or not dichotomously branched.
a. Mycelium of one kind.
1. Zygospores unprotected.
a. Gametes about equal (heterothallic or produced from

widely separated parts of the mycelium)....... Mucor
b. Gametes unequal (produced from closely related branches
Of thesame lament) icpacesok se eens cake Zygorrhynchus

c. Zygospores protected by spiny branched outgrowths from
the: suSpensorss 82 Ses Saiotae es eases Phycomyces

THE BLACK MOULDS 117

b. Mycelium of two kinds, the one colorless in the substratum, the
other aerial, brown and spiny, producing sporangiophores

BG -ZVSOPHOTES: Vaan cdtiaaw ace tw aialnnen da cten centet Spinellus

2. Sporangiophores repeatedly dichotomously branched. Zygospores pro-
duced between dichotomously branched hyphae......

pinta ibe Shahn anes wha ar eAbaVa 6 6 toil otnha shar kie Stabata(aiene a eieite eletart Sporodinia

Ruizopus (=<Ascophora)—Saprophytic fungi, the hyphae
non-septate and much branched, forming long stolons and rhizoids.
Sporangiophores clustered at the nodes (Fig. 1, e) above the
rhizoids. Sporangia spherical containing many spores. Zygospores
spherical or nearly so with a thick, warty dark-brown wall:
heterothallic.

R. nigricans Ehrb. The stolons far spreading, often 1-4 cm.
long, covering the substratum with a cobwebby growth which is at
first colorless, but finally brown. Rhizoids much branched, colorless
at first, finally becoming brown. Sporangiophores mostly in clusters
of 3-5 (seldom single), unbranched 0.5-4 mm. high. Sporangium
and columella (Fig. 1, £). Sporangia 100-3504 in diameter. Spores
irregularly globose or broad oval 6-17 with longitudinal ridges,
light gray (Fig. 1, g). Zygospores 160-220 in diameter, brown-
black, opaque, warty (Fig. 1, d).

On all kinds of organic matter. It is our most common black
mould and will usually appear in a few days upon moist bread or
any organic substance when placed in a moist chamber. Cultures
can always be easily obtained by breaking open a sweet potato that
has rotted with a soft rot and placing it in a moist chamber for a
few days. The rot is caused by the fungus. It also causes large
proportion of the rot of strawberries. In growth upon these hosts
the stoloniferous habit is often not seen.

Agsip1A—Saprophytic fungi. Mycelium stoloniferous; spo-
rangiophores in groups produced only on the tips of the arched
internodes (Fig. 2,a). Sporangia pyriform (Fig. 2,c). Columella
blue-black. Zygospores protected by flexuous circinate outgrowths
from one or both suspensors (Fig. 2, b). Both heterothallic and
homothallic forms known.

A. caerulea Bainier. Vegetative hyphae blue-violet, sporangio-
phores up to 25 mm. in length. Sporangia pale-violet to brown bear-
ing many pale-violet spores 4-74. Columella hemispherical or ab-

118 LEVA B. WALKER

conical, often surmounted by a nipple (Fig. 2, d). Zygospores 60pn
in diameter with suspensors provided with 10-20 long slender
circinate appendages ; heterothallic.

On dung, in humous soil, etc.

Mucor. Saprophytic fungi. Mycelium of one kind, largely
penetrating the substratum, without rhizoidiferous stolons. Spo-
rangiophores produced singly (Fig. 3, a-d). Zygospores unpro-
tected. Gametes about equal, heterothallic or at least produced from
widely separated parts of the mycelium. (Over one-third of the
described Mucoraeceae belong to this genus.)

Key to Species

I. Sporangiophores not branched.

a. Sporangiophores not exceeding 1 cm. in length.........../ M. hiemalis
b: Sporangiephores 2-15 ‘em! long... 05.2 ose A M. mucedo
II. Sporangiophores branching indefinitely (Fig. 3, b)......../ M. racemosus

IIfl. Sporangiophores branched in sympodial cymes.

a. Sporangiophores non-erect ending in a large sporangium and produc-
ing a short distance below more or less closely clustered branches
which "bear sporangia’ (Rig SNe )eie make clnepete cies M. botryoides

. Sporangiophores circinate (Fig. 3, d)...............1 M. circinelloides

c. Sporangiophores straight, not circinate columella spinescent (Fig. 3, £)
Mae ened <n lh Bhs ipods vos ign! x tats he SiG ahr atasouliia pis meas M. plumbeus

M. hiemalis Wehmer. Mycelium a bright gray, sporangia

green to yellow-black 50-80 in diameter. Columella globose when
young becoming somewhat elongated. Spores regularly ellipsoidal
4-10xX2.5-5u. Chlamydospores numerous in the substratum, irreg-
ular pyriform, barrel shaped, etc. (Fig. 3, k). Zygospores globose
70-100p black warty; heterothallic.

Common in soil and on organic matter.

M. mucedo (Linne) Brefeld. Sporangiophores erect, rigid
simple 2-15 cm. high becoming brown when old. Sporangia spher-
ical (Fig. 3, e) 100-200n becoming brown when old, covered with
slender crystals. Columella high-cylindrical to globose (Fig. 3, g).
Spores twice as long as broad 6-12 by 3-6, smooth weak yellow to
colorless. Zygospores globose go-250u in diameter, black, hetero-
thallic.

Common in soil or on organic matter.

THE BLACK MOULDS ; 119

M. racemosus Fresenius. Sporangiophores 5-40 mm. long,
rigid, irregularly branched (Fig. 3, b) in mass a dirty light yellow-
ish color. Sporangia spherical 20-70n in diameter, dirty yellow to
brownish, the sporangial wall smooth, not dissolving in water, but
breaking open easily. Spores hyaline to dirty yellow ellipsoidal to
globose 4-8x4-10n smooth. Zygospores globose 70-85» in diameter,
brown warty, heterothallic. Chlamydospores abundant even on the
sporangiophores ; globose to oblong (Fig. 3, i) 10-20x25-30p» wall
smooth, contents containing oil drops.

Common in soil and on organic matter.

M. botryoides Lendner. Sporangiophores as shown in Fig. 3, c,
1.5 cm. in height. Sporangia globose, clear gray, the walls diffluent
in water. Terminal sporangium 8o0p in diameter; columella globose
(Fig. 3,h). Spores globose 4-10u in diameter, uneven. Chlamydo-
spores lemon shaped 16-22x10-16p.

Fairly common in soil.

M. circinelloides Van Tieghem. Sporangiophores sympodially
branched (Fig. 3, d), branches 5-6 often appearing as sessile,
brownish. Sporangia globose variable in size, those of the larger
having diffluent walls, while those of the smaller are persistent.
Spores 3-4x5-6» smooth, weak gray in mass. Zygospores globose
red-brown with long thorn-like pointed warts that are streaked
longitudinally ; heterothallic.

Fairly common in soil and on organic matter.

M. plumbeus Bonorden. Sporangiophores rigid, erect, straight,
up to 1 cm. high, branched, all branches ending in sporangia.
Sporangia globose, small, up to 10op in diameter, dark brown to
black at maturity, finely spiny, membrane dissolving leaving a basal
collar. Columella long cylindrical to pyriform with one or more
spines on the summit (Fig. 3, £). Spores globose, yellowish brown
with irregular folds. Chlamydospores as in M. racemosus.

Fairly common in soil and on organic matter.

ZYGORRHYNCHUsS—Separated from mucor by the gametes, which
are unequal and arise comparatively close together, almost invariably
originating from a single aerial hypha (Homothallic).

Z. vuilleminii Namyslowski. Sporangiophores flexible, septate
variable in length. Sporangia globose 30-70» in diameter covered

120 LEVA B. WALKER

with needles of calcium oxalate, membrane diffluent only when old.
Columella broader than high (Fig. 4, b). Spores 2x4u (Fig. 4, d).
Chlamydospores 40x14p, usually in chains. Zygospores brownish,
covered with tubercles 2-34 high (Fig. 4, a and c). Azygospores
frequent, smaller than zygospores.

Z. moelleri Vuillimin. Differs only in size of spores. Spores
3X5-7h-

Both species are found in soil and on organic matter.

Puycomyces—Saprophytic fungi; sporangiophores simple
arising singly metallic-green or olive, terminated by a large
sporangium; sporangia many spored, the membrane dissolving;
columella pear shaped (Fig. 5, c). Zygospores dark brown, pro-
tected by dichotomously branched spiny outgrowths from the sus-
pensors.

P. nitens. Kunze. Sporangiophores 7-30 cm. high (Fig. 5, a).
Sporangia about I mm. in diameter; spores ellipsoid 16-30x8-15u
(Fig. 5, b). Zygospores round, 300p thick (Fig. 5, d), typically
heterothallic.

On oily or decaying organic matter, especially on old bones or
other oily matter.

SPINELLUS—Differs from Mucor only in the mycelium being
of two kinds. (It is often included in the genus Mucor.) The
mycelium in the substratum is flexuous, smooth and colorless, that
in the air brown, and covered with spines. Sporangiophores and
zygospores are both produced only on the aerial mycelia (Fig. 6).

S. fusiger (Lk.) Van Tieghem. Sporangiophores single, un-
branched, rigid below bulbous inflated (Fig. 6, e), blue-gray to
chocolate-brown at maturity, 0.1-6 cm. high; sporangia black at
maturity 180-300 in diameter, with a sub-conical columella (Fig.
6, b). Spores spindle shaped, brown 30-40x9-I2n (Fig. 6, d).
Zygospores dark brown 180-400p thick; homothallic (Fig. 6, c).

Common, parasitic on agarics.

SPORODINIA (Syzygites)—Vegetative filaments delicate, pene-
trating the substratum. Aerial fllaments dichotomously branched
producing sporangia and zygospores. Sporangia spherical with
hemispherical columella. Zygospores spherical, smooth, homothallic
(Fig. 7; Plate V).

THE BLACK MOULDS 32!

S. grandis Lk. Sporangiophores repeatedly dichotomous, sep-
tate (Fig. 7, a and b). Sporangia pale red or orange when young,
at maturity brownish or blackish brown. Spores round or ellipsoid
I1-40n. Zygospore mycelium brown, the ends tapering (Fig. 7, c).

Common in nature upon decaying Boleti and other large, fleshy
fungi, but can readily be grown upon bread or other organic media.

II. TRIBE PILOBOLEAE

Sporangia of one kind only with membrane for the major part
solid, persistent, of a very dark blackish color, or swelling only
toward the base. Sometimes the sporangium dissolves, leaving the
columella, while more often it is forcibly thrown off with the colu-
mella and opens only after swelling of the membrane. Zygospores
naked (Fig. 8, e).

PitopoLus (==Hydrogera)—This is the only genus found in
our flora. In this genus the sporangiophore is swollen above and
the sporangium thrown off.

Key to Species

1. Swelling at top of sporangiophore ovoid.

a. Sporangiophore slender, spores oval..............+---- P. crystalines
b. Sporangiophores short and thick, spores globose.......... P. oedipus
2. Swelling at top of sporangiophore almost spherical ............ P. roridus

P. crystalinus Tode. Sporangiophores 5-10 mm. long (Fig. 8,
a), columella conical (Fig. 8, b). Spores 5-10x3-6p, colorless
(greenish yellow in mass).

On dung (usually appears on horse dung left for a few days
under a bell jar).

P. roridus Persoon. Sporangiophores 1-2 cm. high (Fig. 8, d),
columella rounded, short. Spores 8-6x3-4p, colorless (pale yellow
in mass).

On dung.

P. oedipus Mont. Sporangiophores 1-3 mm. high (Fig. 8, c),
contents orange red, columella conical, reaching almost to the sum-
mit of the sporangium. Spores round, 10-I4p, orange. 2

On excrement of animals, on mud, on decaying algae.

[22 LEVA B. WALKER

Ill. TRIBE THAMNIDIAE

Sporangia as in the Mucoreae, but of two kinds: the one
many spored, with membrane that dissolves, leaving only a naked
columella; the other (sporangioles) few spored with a persistent
membrane, often without columella. The sporangioles are produced
onthe ends of branched sporangiophores, which are formed at reg-
ular intervals on the principal sporangiophores. Zygospores as for
Mucoreae.

Key TO GENERA
rt Primary sporangia with, sporangioles without, columella... 7hamnidium
2. Both kinds of sporangia with columella.................. Dicranophora

_. THAMNIDIUM—Sporangiophores erect, principal sporangia
terminal on the main branches, with columella; sporangioles on side
branches, without columella.

T. elegans Link. Sporangiophores (Fig. 9, a) 0.5-3 cm., occa-
sionally 6 cm. high, the branching very variable. Sporangia 100-
200u in diameter, white, with large columella, many spored (Fig.
9, d). Sporangioles globose, small, white 8-16» in diameter, mostly
4 spored (Fig. 9, b). Spores 8-10x6-8% smooth, weak gray brown.
Zygospores globose, black, warty (Fig. 9, e).

On dung, in soil, on decaying plant parts, etc.

T. amoenum (Preuss) Schroet. Differs from T. elegans in
that the sporangioles are produced on the coiled tips of lateral
branches (Fig. 9, c). Sporangia are brownish, with a large egg-
shaped columella. Spores 6-8x4-6p.

On decaying wood, dung, etc.

DicraNoPpHora (Fig. 10)—This genus is rarely found, but is
mentioned because of its having gametes entirely unequal, homo-
thallic (Fig. 10, a), and because of its peculiar sporangiophores
(Hig. 10, a-b).

IV. TRIBE MORTIERELLEAE

Sporangia without a columella (Fig. 11, a-b), membrane dis-
solving readily. The zygospores surrounded by a densely interwoven
mass of hyphae which grow from the suspensors, and from the
branches from which they arise (exterior Fig. 11, d, section Fig.
It, e).

THE BLACK MOULDS 123

Only one genus and one species will be mentioned, Mortierella
polycephala, Coemans. Mycelium much branched and stolon-like,
fusing with neighboring hyphae to form a network, septate when
old. Sporangiophores erect 250» high, in groups of 5-20, swollen
at the base, tapering to the top, terminating in a large sporangium,
and on the upper portion bearing 2-10 short branches terminating
in small sporangia. Sporangia round, white 4-20 spored, spores
10-12y, with a large, glistening oil drop.

On dung, decaying fungi, etc.

V. -TRIBE CHAETOCLADIAE

Sporangia and “conidia” both produced or only “conidia” (the
conidia are to be regarded as reduced one celled sporangia).
“Conidia” formed singly (not in chains) upon the ends of the
usually much-branched conidiophores. Zygospores naked. Chlamy-
dospores, round intercalary.

CHAETOCLADIUM—This is the only genus commonly met with.
Usually parasitic upon other Mucoraceae, occasionaly saprophytic.
Mycelium thin, colorless, forming clusters of short, thick haustoria
at point of attachment with the hyphae of the host (Fig. 12, d).
Conidiophores creeping, verticillately branched. Conidia produced
on the swollen middle portion of the branches, the ends of which
are sterile (Fig. 12, b, c).

C. jonesii (B. and B.) Fresenius. Conidia round 6.5-10p,
singly, colorless, but blue in mass.

On Mucoraceae (partially saprophytic).

_ C. brefeldii Van Tiegh and Le Mon. Conidia globose or
globose elliptical, smooth, colorless 2-5y.

Parasitic on Mucor mucedo and Rhizopus nigricans.

VI. TRIBE CEPHALIDEAE

“Conidia” seemingly produced in chains on the ends of simple
or branched conidiophores. (Really produced in a row in an
elongated sporangium which soon disappears.) Zygospores naked.

Key To GENERA.

¥.. |“ Conidiophores” not swollen at. tips) 2.2. 42:2 Jo.2.0 eG Piptocephalis
2 “Canidiophares” (Swolletr at tipy. Si ces088 sect 25 Ve eels Syncephalis

124 LEVA B. WALKER

PrprocEPHALIS—Parasitic on other Mucoraceae by means of
filiform haustoria (Fig. 13, b). “Conidiophores” repeatedly
dichotomously branched (Fig. 13, a), erect, septate, brownish with
age, not swollen at tip. ‘“Conidia’” cylindrical or spherical in radial
chains clustered on the ends of the branches. Zygospores spherical,
naked (Fig. 13, c).

P. tieghamiana Matruchot. Conidia spindle-shaped to cylindri-
cal, 4-5X2-2.5p.

Parasitic on Rhizopus nigricans (rare).

SyNCEPHALIS—Parasitic on other Mucoraceae (or saprophytic).
Mycelium of very slender branching and anastomosing filaments,
producing numerous clusters of rhizoids which penetrate the host
(Fig. 14, i). “Conidiophores,” stout, erect, mostly unbranched,
enlarged above; “conidia” cylindrical to fusiform, in many radiating
chains clustered on the enlarged summit of the conidiophore.
Zygospore spherical, naked, rarely produced as a lateral outgrowth
from the fertilized cell, Many species are apt to be found, but
none very common, so only a key to species will be given.

Key to Species
I. “Conidiophores” erect (Fig. 14, a).
1. “Conidia” produced directly upon the enlarged end of the
“Conidiophore” (Fig. 14, g).
a. Spores 8-9x3-4u

“Conidiophores” 420-720H high.............seeeee- '..S. sphaerica
b. Spores 20-27x7-11% “Conidiophores”
Vemraslender AG0-A75iuniGhy sen «oe cake cts aioe nets sheets S. tenuts

2. “Conidia” produced upon short branches of the enlarged end of the
conidiophore (Fig. 14, d, e, f).
a. Spores cylindric 60-80x5-64

“Comidiannores’” 2-3mm, ‘high. vse ess oseknGess sebeakue S. cordata
b. Spores rectangular 13-16x7-8u

“Conidiophores” , 300-350. high .e.s 6 sscay «on bice's oles S. pycnosperma
c. Spores cylindric 8-1ox6u

“Conidiophores” 120-150 high. .....ccsssccccesescccns S. nodosa
d. Spores cylindric 5-6x3u

“Gonidianores . ‘O:Smh WISI, .i vebisk ee btiiadc oee a ct S. depressa

II. Conidiophores incurved (Fig. 14, b and c).
a) Spores 10-12x4-5"
“Conidiophores” 170-200# high (Fig. 14, c)..........5. S. cornu
b) Spores 7-8x3-4u
“Conidiophores” 100-120" high (Fig. 14, b)............ S. reflexa

THE BLACK MOULDS 125

. Sphaerica Van Tieghem—On horse dung with Mucoraceae.
. tenuis Thaxter—On sphagnum.

. cordata Van Tieghem—On dung.

. pycnosperma Thaxter—On dung of mice and sheep.

. nodosa Van Tieghem—Parasitic on Mucoraceae.

. depressa Van Tieghem—On horse dung.

. cornu Van Tieghem and Le Mon.—Parasitic on Mucoraceae.
. reflexa Van Tieghem—On dung.

NNnNNnNnNNNMNMNMN

5. Literature

Aside from the general treatment of the Mucoraceae in such
publications as Engler and Prantl’s “Die Natiirlichen Pflanzenfam-
ilien,’ Saccardo’s “Sylloge Fungorum,” Rabenhorst’s “Kryptogamen
Flora,” the literature is very greatly scattered. Only a few of the
more easily obtained publications in English will be listed.

ANDREWS, F. M.
Protoplasmic streaming in Mucor. Bull. Torr. Bot. Club 39:455-510.
1912.
BEsseEy, C. E.
The structure and classification of the Phycomycetes. Trans-Amer. Mic.
Soc. 24:27-54. 1902.
BLAkESLEE, A. F.
Zygospore germinations in the Mucorineae. Annals Mycologici IV,
No. I. 1906
BLAKESLEE, A. F.
Sexual reproduction in the Mucorineae. Proc. of Amer. Acad. Arts &
Sci. 40, No. 4. 1904.
BLAKESLEE, A. F.
Differentiation of sex in thallus gametophyte and sporophyte. Bot Gaz.
42 :161-178. 1906.
BLAKESLEE, A. F.
Zygospores and sexual strains in the common bread mould, Rhizopus
nigricans. Sci. II 24:118-122. 1906.
Grove, W. B.
Monograph of the Pilobolidae. Midland Naturalist. 1884.
Jensen, C. N.
Fungous flora of the soil, Cornell Agr. Exp. Sta. Bull 315. 1912.
PounpD, ROSCOE.
A revision of the Mucoraceae with especial reference to species reported
from North America. Minn. Bot. Studies Bull. 9, 1894 :87-104.

126 LEVA B. WALKER

Swincte, D. B.
Formation of spores in the Sporangia of Rhizopus nigricans and Phy-
comyces nitens. U. S. Dept. Agr. Bureau Plant Industry Bull. 37.
1903.
THAXTER, ROLAND.
New or Peculiar Zygomycetes Bot. Gaz. 24:1-15. 1897.

University of Nebraska.

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DEPARTMENT OF NOTES, REVIEWS, ETC.

It is the purpose, in this department, to present from time to time brief original
notes, both of methods of work and of results, by members of the Society. All members
are invited to submit such items. In the absence of these there will be given a few brief
abstracts of recent work of more general interest to students and teachers. There will be
no attempt to make these abstracts exhaustive. They will illustrate progress without at-
tempting to define it, and will thus give to the teacher current illustrations, and to the
isolated student suggestions of suitable fields of investigation.—[Editor.]

SUMMARY OF THE ELEMENTS IN THE REPRODUCTIVE PROCESS

Teachers of Biology often have occasion to devise special meth-
ods to put before students, in a single view, the various elements
in a complex biological concept. Such a device, if successful, is
in reality a summary of the essential factors or parts of t’ 2 process
or structure, together with an expression of the relations between
the factors, so formulated that the pupil may successfully visualize
them in their proper proportions and be conscious of them in his
complete synthesis of the concept. Such a device is peculiarly
essential with certain classes of pupils who, after an analysis of
such a complex, refuse to re-synthesize all the elements—but allow
one or two major factors to usurp the meaning of the whole. The
need of this sort of aid is apparent especially in the study of the
evolutionary processes where the comparative element enters
largely.

Reproduction is an illustration of such a complex and sig-
nificant phenomenon in biology. It is so varied in the different
groups of plants and animals, and includes or has associated with it
so many different elements at the different levels of life that it is
almost impossible to bring even the most important of these into a
scheme in such a way that the student may grip their interrelations,
and it is not possible in any scheme of reasonable complexity to in-
clude all the related facts.

The device presented herewith is submitted purely as a peda-
gogical scheme summarizing the main features of the process of re-
production in plants and animals, and showing at least a portion of
the evolutionary tendencies connected with the fundamental fune-

128 T. W. GALLOWAY

tion. It is offered to assist teachers and not as a contribution to
knowledge.

The following brief discussion of the major items connected
with reproduction, as exemplified by the two series of organisms,
will serve to explain and elaborate the heads included in the table
(Plate VI).

1. The essential nature of reproduction, no matter how simple
nor what additional processes and complications may be introduced,
is division. The divisions may be equal or unequal, may be of
single-celled organisms or of complex; but without any exception
reproduction always involves the division of parental material to
make offspring. New individuals are thus formed at the expense
of the old. It is always in contrast with assimilation and growth,
and means a sacrifice of the individual structure that has been
builded up by these processes. This reproduction or division re-
sulting from growth is due apparently to some stimulus which is
normally brought on by the accumulation of internal materials and
the near culmination of internal processes and powers which we
describe by the term maturity. The exact nature of these stimuli

we do not know. (See first column of chart.)

2. Primary Kinds of Division (Reproduction). There are
two main types of parental division, depending on the complexity
of the organisms (second column) :—

(a). In unicellular organisms reproduction is a matter of
nuclear and cell division merely. Such daughter cells separate and
lead individual lives. The division of one cell into two or more
cells is at the same time the division of an individual one-celled
parent organism into two or more individual offspring. The cell
being the body of the organism, reproduction is at once a bodily
division and a nuclear or cell division. This is reproduction at its
simplest. Clearly in such a case as this the daughter organism,
which is a small cell, reaches maturity by a simple process of
growth into a large cell, whereupon reproduction may occur again,
when the stimulus of maturity becomes operative.

(b). By far the greater number of our plants and animals
are, however, multicellular in their adult stage. They have reached
this condition by a long series of nuclear and cell divisions in which

SUMMARY OF REPRODUCTION 129

the cells did not separate into distinct individuals, but remained to-
gether as parts of a more complex individual. Division of such an
organism as this is a somewhat different thing from what was de-
scribed in (a). The many-celled organism may divide bodily into
two or more smaller, but many-celled, offspring. This is analogous
to the division in (a) in that it is a mass division. It differs from
(a) in that it is not at all a cell division.

Again the multicellular organisms may reproduce by freeing,
through division, single-celled offspring, which are, of course, the
immediate product of cell and nuclear divisions taking place in a
certain line of cells in the body of the multicellular parent. These
divisions are homologous with the reproduction described in (a)
in that these reproductive cells are formed by nuclear and cell di-
vision. The process differs from (a) only in the fact that the
dividing cells that form the offspring are harbored in a many-celled
body. We may say then that reproduction in (a), with its double
aspect of cell division and bodily division in one operation, has
become specialized in (b) into two separate possible operations,
one of the multicellular body and the other of the unicellular germ
cells, contained in that body.

3. The Ratio in Size of the Parental Body to That of the Off-
spring. In its simplest form the division of the parent, whether
one or many-celled, into new individuals may be considered as pro-
ducing two practically equivalent offspring, each of which has one-
half the endowment of original parental material. So far as this
ratio is concerned the condition is much the same in the unicellular
organisms and in those multicellular ones,—as flatworms, naidiform
oligochetes, and the like-—in which there is division into essentially
equivalent daughters. In both cases we may fairly say that the
parental organism is completely destroyed in the act of reproduc-
tion. Strictly speaking, there is no parent left, and the two off-
spring formed are mutually at the highest advantage possible to
offspring in that they only have to double their material to be at
complete maturity of individuality. (See third column of chart.)

4. Depreciation in the Volume of the Reproductive Units
(Offspring). The method just described is at a maximum of sac-
rifice on the part of the parent with maximum biological advan-

130 T. W. GALLOWAY

tage to a limited number of offspring. It leads to well endowed off-
spring, brief periods of immaturity, numerous reproductions, and
impermanent individuality on the part of the parent. From this
simple condition as a starting point there are, in practice, two de-
partures found in organisms whereby a certain form of increase of
effectiveness is obtained: (1) there may be a division of the en-
tire parent into many offspring, by fragmentation, sporulation, etc.,
which continues the complete destruction of the parent as in the
former case, but reduces the endowment of each offspring. The
compensating fact is the increased number of offspring. This de-
vice increases the period of immaturity and in consequence pushes
apart the successive reproductive generations; (2) the second case
is illustrated by a budding process such as is seen in the yeast cell
or in hydra,—or by the condition to be described later for still
higher forms. In these the parent is not completely destroyed,
and the one or many offspring are reduced correspondingly in
volume and in initial vital efficiency. This diminution in volume
may be such that the offspring of a multicellular organism may con-
sist of a single cell. This is clearly a much more economical mode
of reproduction with very much less drain on the parent; it allows
a more continuous parental existence with an increased emphasis
on permanency of individuality; it makes possible frequent or even
continuous reproductive activity, with more numerous offspring.

5. Depreciation in Value Accompanying Depreciation in
Volume. It will be realized in connection with the above that the
economy and gain through safeguarding the parent and in the capa-
bility of producing many offspring has been purchased at the ex-
pense of the amount of the original parental material going to
each offspring. As the result of this reduction in volume the off-
spring necessarily has a longer, more difficult course to run in
coming to reproductive maturity. This means that the decreased
volume entails a decreased present biological value in the repro-
ductive units. The more the offspring differs from the parent
the more extended and difficult is its task in reaching maturity. The
reduction in volume and in value is greatest in our higher, more
complex multicellular animals and plants in whose case the new
offspring is merely a single cell at the outset. This condition is il-

SUMMARY OF REPRODUCTION I3I

lustrated both by the spores and gametes. In the gametes, which
are reproductive units that normally unite before development, and
in the spores, at least of the higher plants, there is another most
interesting reduction to which further reference is made in Sec-
tion 9. Here the offspring is not merely reduced to one cell; the
number of chromosomes in the nucleus of that cell is reduced to
one-half the number characteristic of the individual body cells of
the species. One further step in the depreciation in volume is seen
in the sperm (or male) cells in most animals and plants. In ad-
dition to the loss of one-half its chromosomes, the male cell is much
reduced in the amount of cytoplasm.

6. The Necessity of Restoration. The depreciation of the one-
celled offspring of the higher animals and plants described in the
preceding section is shown further by the fact that these new off-
spring are frequently unable to enter, as soon as they are formed,
upon the series of cell divisions and other changes which have been
described as necessary to maturity. Their further development
seems to be checked in part by the excessive reduction they have
suffered. This is very often true of spores, which may not under-
take to germinate for some time after their formation, even though
external conditions seem favorable. The gametes, eggs and sperm,
are normally even more incapable of entering at once on the stages
that lead to maturity, without external help. It is even more true
of the sperm than of the egg. The sperm, it will be recalled, is
like the egg in having only one-half endowment of chromosomes;
it has also lost the major part of its protoplasm, of which the egg
has an abundance. The spores and the eggs of many species may
develop rather promptly, but so far as we know the sperm is so
reduced in the typical higher plants and animals as to be wholly
incapable, in itsef, of developing into the adult. (See column six
of chart.)

It would seem, therefore, that the tendency to economize paren-
tal expenditure and to increase the number of offspring had over-
reached its mark,—defeating in large degree its prime result of
economic reproduction. At this point in nature we find a series of
recuperative processes by which these reduced reproductive bodies
are given the power to resume their activities and are restored ap-

132 T. W. GALLOWAY

parently to a vigor which enables the single cell to develop, in
proper order, all the essential characteristics of the species to which
it belongs. Because of these restorative processes these single-celled
reproductive units cease to be merely single cells; they are poten-
tial adults.

7. Methods of Restoration. This need of restoration, sug-
gested by the great reduction in the parental substance—the cyto-
plasm and chromatin—that goes to each of the unicellular offspring
and reinforced by the fact that they are often unable at once to
develop, is met in nature in two or three principal ways.

In the first place, and most simply, it frequently happens that
the offspring may regain this power of developing to the mature
state by a mere season of rest or quiescence, without any other
change of condition. This is quite commonly true of the spores.
It is not always true that the spores need such a rest period, how-
ever; they may often develop directly. (See column seven of
chart.)

Secondly, in addition to the mere lapse of time the spores,
after leaving the parental tissues, are undoubtedly subjected to de-
cided changes in respect to moisture, temperature, and other im-
portant environmental conditions. These changes may themselves,
rather than the mere rest, be the stimulating and restoring agency.
We know that decided changes in environmental conditions do
stimulate renewed activity in vital processes. This may be anal-
ogous to the artificial stimuli that arouse unfertilized eggs into
action as described below.

In the third place, the gametes usually demand more than rest.
In this type of reproductive units, where chromosome reduction is
well nigh universal, it is ordinarily essential that two gametes (off-
spring )—each containing one-half the specific number of chromo-
somes—shall unite and bring the contents of both the cells into one.
Such a united cell, which is no longer a single reproductive body
or offspring, but rather a fusion of two offspring, has the power
of development, which did not reside in either of the original cells.
This is called conjugation or fertilization. It frequently happens
that a period of rest is also needed in connection with these unions,
though this is not the rule.

,

SUMMARY OF REPRODUCTION 133

Finally, it has been discovered by experiments that this restora-
tion and immediate power of development of gametes (the egg par-
ticularly) can be secured in some organisms in various artificial
ways. These consist for the most part in applying to the gametes
certain conditions,—changed mechanically or chemically from the
normal. This is known as artificial parthenogenesis,—and means
that certain eggs which would not ordinarily begin development
without union with another cell may be caused to develop by chem-
ical and mechanical stimuli. It will be seen that this is analogous
to the condition described in the second type of restoration above,
by which spores may come to the point of developing through
changed external conditions. The same power is sometimes nor-
mally possessed by the larger of the two gametes,—the ovum (par-
thenogeneses: section 9).

8. Conjugation (Isogamy). In that form of restoration of
power to the offspring, in which gametes unite and, by fusing two
half-sets of chromosomes, regain the full complement of chromo-
somes, the most simple form has uniting gametes in which no dif-
ference is distinguishable. So far as we can see, each of the off-
spring entering into the fusion is exactly equivalent to the other.
Each has suffered equal reduction as compared with the original
parent, both in protoplasmic substance and in chromosomes. We
cannot properly use here the terms male and female, nor fertiliza-
tion, nor sex. The fusion is mutual and there seems no differen-
tiation of function in the gametes. This is known as isogamy or
conjugation. In some cases the conjugation is facultative; that
is, the gametes usually conjugate, but if this does not occur, they
may after a period of rest develop without it.

9. Differentiation Among the Gametic Offspring (Heter-
ogamy). In many species of organisms the same individual may
produce two different kinds of offspring or gametes. In such cases
the gametes may be nearly alike, differing only slightly in size. In
other species there may be great difference in size and behavior.
At the extreme of this differentiation we have the two types of
offspring known as ova and sperms. Characteristically, the eggs are
unicellular offspring, which are large, spherical, well nourished, and
sluggish cells, with much protoplasm, but only one-half the specific

13-4 T. W. GALLOWAY

number of chromosomes at maturity. The sperms, on the other
hand, are usually actively motile cells, of very various shape, though
not spherical like the egg. They are much reduced in protoplasm,
but they have the same number of chromosomes as the ova,—that
is, one-half the specific number. (See column four of chart).

This condition introduces ser. The ova are known as female
and the sperm as male. The union of the ova and sperm, in pairs,
which is usually necessary to restore the power of development, is
called fertilization. In some species the ova may develop without
union with the male cell. This is known as parthenogenesis, and
it is possible that this ability represents the action of some con-
dition of the parental body, analogous to hormone action, which sub-
stitutes some other stimulant for that of the male cell, which is the
usual or normal one.

It is necessary to insist that the eggs and the sperm are the
real offspring, and that the act of reproduction in the female is
the production of eggs; that reproduction in the male is formation
of sperm; and that fertilization, which is union, is not reproduc-
tion at all, but the direct opposite of it. It is in part at least a
restorative process following upon the reduction and depreciation
of the gametes in reproduction, and instituting a series of cell di-
visions which provides both for the new parental body and the new
generation of reproductive cells (offspring) which are derived from
it.

10. Differentiation of Organs of Sex. In the simplest state of
the formation of the reproductive units, especially before these units
themselves are sharply different, the organs or structures in plants
and animals which produce the two kinds of offspring are not only
in the same individual ( hermaphroditism), but they are much alike,
- except for the fact that the gametes they produce differ more or
less. There is, however, a perfectly clear tendency for the organs
and structures connected with the production and distribution of
eggs (female offspring) and the sperm (male offspring) to become
different,—and often very markedly different, even when they oc-
cur in the same individual,—quite as diverse indeed as are the off-
spring which they produce. This is a quite common kind of her-
maphroditism, in which a single plant or animal develops two dif-

SUMMARY OF REPRODUCTION 135

ferent, and often very complex, sets of sex structures to produce
and take care of the two kinds of offspring so as to insure their
fertilization and development. It is illustrated in earthworms, snails,
and many other organisms. Ordinarily this union is supposed to
be not between the offspring of the same parent, but of different
parents (cross-fertilization). Doubtless self-fertilization is also fre-
quent.

11. Diversification of Parents (Sex-dimorphism). The prob-
lem and function of producing and caring for ova is so different
from that of producing and caring for sperm that such a differen-
tiation as we have seen in the last section takes places in the organs
that do the work. This differentiation does not in the majority of
animals and plants stop here. The differentiation of sex, first seen
in the offspring (eggs and sperm) and then in the organs producing
them (ovaries and spermaries), comes to show in the individuals
which bear the organs; and male individuals come to be differen-
tiated from female individuals as much as ovaries differ from testes
or ova from sperm. (See column five of chart).

Among animals and plants can be found all degrees of these
sex differences. We have individuals that produce ova and have
female organs and characteristics at one period of life, and later
are males and produce sperm. In others, permanently male or fe-
male, the external differences are so slight that dissection of the
organs of reproduction alone can determine the sex of the parem.
In still others there are such striking differences between the sexes
that they would not be regarded as belonging to the same species
of organisms from structure alone. This is the condition in most
of the higher organisms, and is particularly striking in all the
higher animals. The differences between the male and female in
man, in birds, in many insects and spiders are matters of common
observation.

12. Parental Care of Offspring. The depreciation of offspring
in the interest of economy to the parent and of increased number of
offspring, coupled with the resulting need of restoration by fertiliza-
tion or some other device, has led to other most important biolog-
ical results. The longer course over which the offspring must go
to reach maturity, the diversification of the gametes, and the con-

136 T. W. GALLOWAY

sequent diversification of the male and female parents furnish the
biological groundwork for very interesting adaptations for mating,
home-making, parental care. These adaptations may be structural,
instinctive, emotional, or intellectual. These are among the highest
and most stimulative of the biological phenomena,—upon both
parents and offspring. Because of such conditions the care-taking,
sympathetic, social aspects of parents are emphasized, the period
of filial dependence is made longer and more fruitful through asso-
ciation with parents, and the motives for social life and development
are introduced. It is not easy for the student to exaggerate the
evolutionary importance of this group of phenomena correlated with
reproduction, and common in many animals of the higher groups.

13. Combination of the Reproductive Methods in One Indi
vidual. In what has been said the process has been treated as though
a given species of organisms had only one mode of reproducing.
As a matter of fact we often find that a plant or animal will for a
period show some form of mass division (vegetative reproduction)

EXPLANATION OF PLATE VI

The scheme represents an effort to put in tabular form some of the
more important relations suggested in the discussion. The vertical columns
indicate certain of the classes of facts connected with reproduction. The
horizontal subdivisions express some of the important variations in respect
to the particular phenomena named at the head of the columns. In a
general way the category nearer the bottom of the page at each step is
looked upon as the lower, and as giving rise to that above it. For example,
the following free translation will illustrate what is conceived as the
simplest condition from which others have arisen: “Direct mass-division
of unicellular parents into (2) offspring, equal to one-half the parent and
equal to each other, which demand no special restoration to enable them to
begin the new growth, is exemplified by fission among the Bacteria.” The
signs + and — in column six indicate “requiring” and “not requiring”
restoration respectively.

It is not intended always to imply that the subdivisions in a brace are
strictly logical subdivisions of the category preceding. For example:
“Many O.” near the bottom of the fourth column manifestly cannot be a
subdivision of “O = % parent”; it is rather thought of as a derivative
and extension of the fission process, whereby “2 equal O.” are produced
from the one-celled parent.

ia METHOD: TYPE OF
NN
£ LESL REPRODUCTION
| imorphic Sexualitg.

io ' Rarthenogenesis- BEES, 2
| UNION \Hermaphrodile E.WORMS
| |Spore Formation
LS | Tabers

| | BUdAING; EG. MYOROZOA.
| Flat Worms (FISSION)
|

|

|

|

|

|

|

|

|

|

|

UNION \Eadorine.

Te

72 | UNION '‘Vorticella.
es. UNION ‘Some Heliozoa
fea!) AEOF Sores

a) eas, f ete,

9 UNION \|Llasmodium.

(of Two | Chlamgdomonas
UNION | __THE SHALL ZOOSPORES ___
hg O*MANY ee”

/ MYXOMYCETES

er keST eee

COR ! __ SSeS FOES ce
PERMT \Desmids

fea y\ Paramecium
REST Zaglena__
| \Bacleria (FISSION)

¥

FARENTS
eal 6. BODILY ne REPRODUCTIVEUNITS | PARENTS ee NL. REST | REPRODUCTION

ESSENCE a. CELLULAR ee BETWEEN | DI VERSIFICATION
aS )

DIVERS'N ‘neo of Power and| METHOD: TYPE. OF

DEPRECIATION ees ; 2 DIVERSE wi kestovati ZOF?2 UNION \Dimorphic Sexaallg.
piece } jo PARENTS| —Feestoration \ Rarthenogenesis: Cm
Parent | +Restoration | UNION _\Hermaphrodile £.worms

Malfeetiatar MUCHEARGER
YOUnicellular | 7Pypen7r q
Ss Oregeliatar | (PARENT | +0r~Kestoration Spore Formation

parents \liregaal |
|
|
|
ee |
Opspring (eter gy a +Restoration | REST | Tubers
— UC Egecead oraneguay -Restoration | | Budding; £6. AroROZOA.
oe RECT O.=22 PARENT \QEqual : 2 Restoration | ' Flat Worms (FISSION)
ASS” ie y ee OV ! (Qunegua i DBE +Restoration | UNION ‘\Eadorina.
PA 5 - ata PARENTS | ee SOE +Restoration | UNION ‘Vorticella.
MEL NGS:
peal | «Restoration | UNION Some cintiozoe
Unicellular Opepring, | Q. \OLgaal emer Teer) ae | REST i Seieees
Bodies | -Restoration | est ie
Unegual | | +Restoration | UNION Plasmodium. .

ALL ZOOSPORES

of Two | Char MGI AOTRONAS
re) any Aetinospheriam

MYXOMYCETES

ee Foaal- +kestoration a
| @Ponuaren) REST | Gnsceses

|
|
!
a / |
O.= 22 PARENT | ror-FRestoration | Mecist | Ye
|
|
I
|
|
|
|

PERM'T \Desmé as
union}

TEMP’Y | | Paramecium
REST iL aglena
-kestoration | ' Bacteria (FSSION)

+ftestoration

a Liga al Oission)

Piate VI

SUMMARY OF REPRODUCTION L37

or spore formation, and will then, owing to some change of con-
ditions internal or external, enter upon reproduction by gametes.
The same individual, thus, at different periods of life or with dif-
ferent stimuli in the way of temperature or nutrition and the like,
may combine the advantages outlined in the preceding sections in
the various special methods of securing the reproductive act. The
possession of these several methods of reproducing of course makes
possible a much more equable and adequate response to the varying
environmental pressure.

14. Alternation of Generation —a Combination of Reproduc-
tive Methods in Different Individuals Constituting a Single Life-
Cycle. Another form of combination of reproductive methods than
that described in section 13 is found in most of the complex plants
and in many types of animals. It consists essentially in, we may say,
a reproduction by gametes, in which the egg is fertilized by the
sperm in the usual way resulting in a type of individual which we
shall call “A”; individuals of the type ‘““A” become mature and do
not have the power to produce gametes at all, but, on the contrary,
they may bud, or divide vegetatively, or form spores; when these
new individuals come to their reproductive maturity they are dif-
ferent, often very different, in appearance from individuals “A”,
and may be called “B”. This second type of organisms in its turn
cannot reproduce as “A” reproduced, but reproduces by gametes,
and the embryo resulting from their union matures into a type like
the original “A”. We have thus come round to the same point
in the “cycle” at which we Started, and in doing so we have had
two succeeding types of individuals in the same species and two
different methods of reproduction regularly alternating. “A” pro-
duces non-sexually individuals “B”, and “B” reproduces sexually
individuals “A”. In coelenterates “A” is the tubular type, multi-
plies by budding and ultimately produces the medusoid individuals
“B”. These latter reproduce by eggs and sperm, from the union
of which the tubular hydroid forms are again produced.

In plants, “A” is known as the sporophyte generation, which
produces spores. When the spores germinate they produce the new
and very different gametophyte generation, which reproduces by the

138 T. W. GALLOWAY

formation of gametes, male and female. The union of these initiates
the sporophyte generation again. (See text Fig. 1.)

Aniferidéum.
x.
Zoteg ni iN
xX.
Mature Gam elophyte. QAntherozotd. x.
(Thelivs). Kr Ovurm
Spore. x. Fectilized Ovum.
Begins 2x-gene sation.
Molter -cell
of Spoves. Fuaitue e
Begins x-gene cation 5 i igelye e

\e

pec Sovangi Um. 2X.

Fig. 1.—A diagram to illustrate the life-cycle, the alternation of generation including
the relation of the x and 2x condition of the chromosomes, in one of the higher plants
(e. g. a fern). The fertilized ovum is the beginning of the sporophyte generation. The
spore is the beginning of the gametophyte generation.

15. Relation of the Chromosome Reduction to the Alternation
of Generation. The two kinds of gametes possess a number of
chromosomes characteristic of the species. This number is spoken
of as x. When they unite the chromosomes of male and female
gametes do not lose their integrity, but the resulting embryo has
the double number of chromosomes, or 2x. As cell division oc-
curs and the resulting plant becomes multicellular, the number of
chromosomes is not reduced, but all the body cells of the new
plant (sporophyte) have 2x chromosomes. This remains true
until the nuclear divisions immediately preceding the formation of
the spores, or non-sexual reproductive bodies. At this time there
is a reduction to one-half the number, and thus the spores them-
selves contain nuclei with only the x number of chromosomes.

SUMMARY OF REPRODUCTION 139

As the spore germinates and the new multicellular generation
(gametophyte) is produced this condition is unchanged; and since
the gametes are formed from the gametophyte they have likewise
the x chromosomes, as we saw at the beginning of this section.
In the union of the gametes the double number is again restored.

The condition is pictured in the accompanying diagram (Fig.
1). The relative length and importance of these two generations
is very varied in the different groups of plants, but the alternation
of generation, and the distinctive reduction and restoration of
chromosomes by which the generations, are marked are very con-
stant.

Immature Polae Bodces
Ovum. 2x. Foemed.x.
spermatocyte.
Spetrn Kit’ 4 en
Mother Cell. 2%.
Witte Mature
e pod Ovurt.x.
Marore
Male Body.2zx.
Mature J
Female Babe
ey Begins 5 cb
ae Cle avage
Differentiation Stages. 2x.

of Sexes. 2x } ‘ee :
f Sutra of
Tis sues. 2x.
Fig. 2.—A diagram to illustrate the life-cycle and the x and 2x state of the chromo-
somes in higher animals. It will be seen that the x stage is very brief.

A condition somewhat analogous is seen in diagram, Fig. 2,
which represents the condition in higher animals. Whether it is
more than analogy we are not now prepared to say. The ordinary
body of the animal is made up of cells with 2x chromosomes. This
continues up to the time of the formation of the gametes. In their
formation and maturation the number is normally reduced to x,
immediately to be doubled upon the union of the gametes in fer-

140 NOTES, REVIEWS, ETC.

tilization. This reduction and doubling of the chromosomes is
strongly believed to be closely connected with the hereditary trans-
mission of characteristics from two parents, and seems to have to do
with the changes in the germ plasm that produces variation within
the offspring of the same pair of parents.

T. W. GALLoway.

THE GROWTH OF A CoMPpouND EYE

As is well known to students of insect life, there is a period be-
tween the larval and adult stages of development when important
structural changes take place. For instance, take one particular
type of organ, such as the eye. In a moth larva such as the Tus-
sock, Notolophus leucostigma, the larva has several single eyes
grouped on each side of the head. Somewhere between the time
of its entering upon the pupal stage and its emergence therefrom
as an adult it exchanges these simple larval eyes for an elaborate
pair of compound eyes. How it does this, and what becomes of the
old larval eyes, is a process so well hidden from view in the pupal
case that only cytological work can reveal the secrets.

When ready to pupate the larvae of most moths seek various
sheltered positions in which they undergo their final molt or shed-
ding of the larval cuticula. This leaves the hypodermal (epider-
mal) cells of the creature raw, under which conditions they exude
a fluid which hardens and forms the pupal case.

As most individuals of a brood undergo their stages at similar
intervals one may by collecting a number of pupae at this stage
get material for studying not only the stages of the eye, but of
other structures as well. By selecting individuals, at say two-day
intervals from the first day of pupation onward, we will get a series
showing the progressive development of the parts.

The pupa so selected should be opened in the back by a sharp
instrument to allow the rapid penetration of the killing and fixative
fluids. If any particular part of the tissue is wanted, it is neces-
sary to be very careful, in making openings, not to injure or de-
form the tissues in the immediate vicinity.

During the pupal life, two very important and entirely oppo-
site processes are necessarily very active. One is the breaking

GROWTH OF COMPOUND EYE I4I

down and absorption of the useless larval structures. This de-
structive process is carried on by two different factors :—by phag-
ocytes, a special form of cell which gives out certain products
which break down and disintegrate passive adult cells into blood
pabulum; and by natural breaking down processes of cells which
have lost their functional places in the changing economy. The
other or opposing process is one of rapid cell division and growth
of new types of cells which are to become differentiated into new
adult organs.

In the first class of structures which disintegrate, we may in-
clude the larval types of eyes. These, following all the previous
larval molts, have grown a new cuticula and have retained their
function ; but now it is no longer so. With this last molting process
they immediately collapse and shrink toward the brain to which
they are attached. Here they will be recognized in the photos of
subsequent stages as the dark pigmented masses at the posterior
lobe of the brain. Finally through growth of the brain they come
to lie well down on the stalk of the lobes.

Let us now turn toward another aspect of the head of the
creature. During larval life the cheeks of the larva were plump
with masses of muscle which were absolutely essential in working
the cutting jaws during the active voracious life of the caterpillar.
From now on these muscles are useless, as the biting jaws will
never be used again. These rounded cheeks are destined to be-
come the seat of a new activity; they are to become the site of the
enormous compound eyes of the adult insect, containing many
hundreds or thousands of eyes, as the case may be. The phago-
cytes have been active for several days and there is a large cavity
filled with body fluid in each cheek. This blood fluid contains
numerous phagocytes, many broken down tissues, and many fat
cells which enter through the wide neck opening from the thorax.
In the meantime, the ectodermal layer of cells has proliferated to
quite an extent so that there is a seemingly chaotic mass several
cells deep.

Fig. 1 (Plate VII) represents a frontal section of the head in
the region described above, passing through both brain lobes and
through the regions to be occupied by the compound eyes. The

142 E. W. ROBERTS

larva is in about a four-day stage of pupation. This photograph
may serve as a topographic chart to which may be referred all the
special organs mentioned in later figures.

Fig. 2 is an enlarged view of the optical elements of the same
4-day stage. Practically all the cells which are to form the future
eye tissue are now present and there is to be no increase in num-
ber henceforth. Later change is due to growt® any! differentiation
of these cells. That is, the cytoplasmic vegetative systems of these
cells are to grow into their hereditary forms, and the individual
cells are to adjust themselves into their peculiar group relations.
What looks at this stage as a hopeless jumble of cells without order
or form, is the foundation of an order which will develop itself
with remarkable speed.

If we now turn to the next or 5-day stage, shown in Fig. 3, we
shall see that the cells have undergone a remarkable arrangement
into definite groups, each of which is an exact duplicate of its
neighbor. Also within these groups, they are beginning to show
definite shapes and relations to each other. Their vegetative sys-
tems also grow to show more of their future structural characters.
Order has appeared out of seeming chaos.

By examination of the 6-day stage, Fig. 4, we will see that the
mass of optical elements has doubled in thickness by the growth of
the cell systems.

Fig. 5 is a view of the optic lobe of the brain at this same 6-day
stage. There has grown a bewildering complexity of elements
here. The large dark pigmented mass is the rudiment of the
larval eyes. ’

We will now turn to the 8-day stage well shown in Fig. 6.
Here we find the cells again doubled in bulk as compared with the
6-day stage. The pigment cells and nerve cells are greatly elongated
and now reach backward nearly to the brain itself.

A cross section of these groups of optical elements is shown
in Fig. 7 (Plate VIII). Here the elements are seen to be grouped
into hexagons, an effect produced by pressure of the opposing
groups of cells. Toward the margin of the section the elements
are, of course, cut obliquely. A surface view of the cornea, as

GROWTH ,OF COMPOUND EYE 143

Fig. 8, shows the beautiful regularity of the hexagonal visual ele-
ments.

As to the cytological structure of these compound eyes, there
are several views. Two of these taken from, Lang’s “Comparative
Anatomy,” pages 470 to 471, are here illustrated in diagrams (text
figures) A and B. These authors both consider the hypodermal
layer of cells as distinctly a layer by itself. We can hardly agree
with this view, however. In all the types of eyes examined we find
that in the early stages there is: but a single layer of cells in the
hypodermis. Later some of the cells draw inward and_ by
division give rise to other cells which form. the ommatidium group.
So it appears to us that the whole group of cells is strictly hypo-
dermal in origin. These cells now elongate, forming spindle shaped
cells which extend more or less the entire depth of the ommatidium.
Such elongated undifferentiated, cells are seen in the eye of the

The structure of an ommatidium (single eye) of the compound eye :—A, ac-
cording to Patten’s view; B, according to Grenacher’s view. cl, cuticular corneal
lens; hy, hypodermis cells; r, retinophorae-crystal cells; mr, nuclei of the same;
k, crystalline cone; p, pigment cells; ret, retinulae; rh, rhabdome; n, nerve.

According to Patten (A) the ommatidium is apart from the corneal hypo-
dermis, of one layer, all its elements passing by means of thin processes through
its whole thickness from the base of the, corneal lens; according to Grenacher the
ommatidium, apart from the corneal lens, consists of two layers. (Taken from
' Lang’s Comparative Anatomy, page 471.)

€, Section through the ocellus of a young Dytiscus larva (after Grenacher.) :
ct, chitenous cuticle ;l, cuticular lens; gh, cells of the vitreous body; hy, hypo-
dermis; st, rods; re, retinal cells; no, optic nerve,

144 E. W. ROBERTS

male Ephemera, Fig. 9, (1). The nuclei of these cells gradually
migrate as they grow to the localities where they are found in the
adult eye.

The corneal hypodermis consists of a varying number of cells
arranged in hexagons following the shape of the compressed om-
matidium below. When the crystal cells begin their enlargement
these corneal cells are displaced by their outward growth and fin-
ally come to lie around the base of the cone group of cells. This
is well shown in Fig. 9, (2) of the Ephemera eye. They are thus
seen to be the last of the hypodermal cells to assume the elongated
spindle shape lying vertical with the rest of the ommatidium group.
See also Fig. 10. So we consider there is really but one layer in
the eye when they have all reached adult stages. In both the dia-
grams the rhabdome should have been shown as a nucleated cell,
being the central cell of the group of retinular cells. The nucleus
of the rhabdome cell finally comes to lie at the inner margin of
the eye. We believe, therefore, that the usual method of dia-
gramming of the optic nerve is wrong, the central retinular cell

DESCRIPTION OF FIGURES
Pirate VII
Fig. 1. Tussock Moth, 4-day pupal stage. Frontal section thru brain
and compound eye.
Fig. 2. Tussock Moth, same stage. Portion of developing eye much
enlarged. :

Fig. 3. Same, 5-day pupal stage. Region similar to Fig. 2.

Fig. 4. Same, 6-day pupal stage. Similar region.

Fig. 5. Same, 6-day pupal stage. Optic lobe of brain.

Fig. 6. Same, 8-day pupal stage. Region similar to that of Figs. 2-4.

Pirate VIII

Fig. 7. Tussock Moth. Eye elements cut in cross section.

Fig. 8. The corneal lenses of the compound eyes of Tabanus astrata.

Fig. 9. Eye of male Ephemerid: 1, spindle cells; 2, hypodermal cells;
3, thabdome cell.

Fig. 10. Eye of a female: Rhamphomyia—a Dipteran.

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GROWTH OF COMPOUND EYE 145

being the nerve cell, which extends outward among the crystal
cells and inward where it connected with the ganglion cells of the
brain, as seen in the Tussock photos.

The compound type of eye is generally considered to be an
elaboration of the ocellus or single eye type of larval forms. If
we compare the diagrams of the ommatidium by Patten and Gren-
acher with the ocellus of a larval form (see text Fig. C, after
Grenacher) we can see the probable homology of the various cell
structures. We may consider the rods (st) on the retinal cells
(re) as the remains of spines on the hypodermal cells which form
the optic invagination. This same spine structure also forms the
olfactory nerve endings on the antennae of the Diptera. In the
ommatidium of the compound eye, of the seven retinular cells,
all but the central one has lost its rod. This rod we consider to
terminate at the base of the crystal cells as seen in the Ephemera
material. This termination corresponds with the position of the
rods in the ocellus. See Fig. 9 (3).

The cells which close in from the sides in both the ocellus
and ommatidium forming the crystal cells may be considered as
homologous. They may be considered as full length cells much
compressed in a lateral direction.

The space in which the crystal cones of the ommatidium are
deposited is homologous with the space between the rods and the
vitreous body cells of the ocellus type.

These observations do not lead us to believe that the ideas of
either Patten or Grenacher are entirely correct in regard to the
structure of ommatidium.

E. W. Roperts.
Battle Creek, Mich.

146 EDWARD POTTS

FRESHWATER HYDROIDS

The now familiar Hydra seems to have been first noticed in
the year 1703. About 40 years later the observations or experi-
ments of Abraham Trembly of Geneva excited a widely extended
interest in it, not so much because of the peculiarities of its life
history, as from its apparent indifference to the many seemingly
fatal wounds given it to produce its death. On account of the
interest taken in these experiments, this creature was often re-
ferred to as the Zoophyte of Trembly, but this had long since
lapsed into the common, as well as generic, name of Hydra. Three
species are generally recognized in this genus, as Hydra vulgaris,
Hydra viridis, and Hydra fusca, on account of a variation in the
number of their tentacles, a difference in their color, or an increase
in their extensibility and consequent contractility. This is most
conspicuous in the case of Hydra fusca, where we may first see
it after disturbance as a “ball of greenish jelly,” lengthening a
moment later into a slender thread or stem, from near the distal,
or outer, end of which six or more tentacles are budded out rap-
idly, lengthening into a drooping mass or fringe of extremely
delicate filaments knotted along their whole length with what are
known as lasso, or poison, cells. The water-flea or other Protozoan
which accidentally touches one or more of these, is liable to be
paralyzed by some of these poisonous darts, when the fortunate
tentacle shrinks down to the mouth of the Hydra, quickly opened
to receive it, and—“facilis decensus averni.” So much for the
common Hydra, which will easily be recognized.

The next Hydroid to join the list of freshwater forms was
Cordylophora lacustris, much more complicated in its structure,
wherein it nearly resembles several of the marine forms, and, while
it does not throw off any free-swimming medusz, it does, at certain
seasons of the year, give birth to so-called hydranths that take
root and grow up directly into a new generation of hydroids.

Following this come the Medusae, Limnocodium sowerbu and
Limnocnida tanganyike, found respectively, the first in Regents
Park Gardens in London in 1880, the other in 1883 in Lake
Tanganyika, Central Africa. The parent hydroid of neither of
these, or from which presumptively it must have arisen, has not,

FRESH WATER HYDROIDS 147

as yet, been positively ascertained, although Mr. Bourne and others
did find in other tanks and in the Kew Gardens, also in London,
a hydroid greatly resembling the next to be described form, from
which they assumed it may have descended. This connection the
writer thinks has not, for reasons that will be given, been fully
proved.

In the spring of 1885 the writer of the present paper, while
studying the life history of a new Polyzoan, which he had named
Paludicella erecta, found upon the surface of some stones collected
during the previous autumn in the neighborhood of Philadelphia,
Pa., and kept over winter in his home, some novel forms that
he soon convinced himself were of hydroid character, though
entirely destitute of tentacles, or of other organs of prehension or
locomotion. These, with the consent of his friend, Dr. John A. -°
Ryder, of the University of Pennsylvania, form the new genus
Microhydra ryderii. They are about one-half a millimeter in length
when single, or when branching near the base, which will be called
the pedal disc, the total length is about one millimeter from head
to head. The diameter of the cylindrical body is about one-tenth
millimeter. A few lasso cells are scattered along it, but a great
many, say 40 or 50, are collected upon each capitulum or head.
Here the mouth is placed, but, except when the lips are everted
while feeding, it is with difficulty recognized. As no means have
yet been discovered by which it may remove or re-attach itself
upon its pedal disc after removal, and having no grasping organs,
our perplexity may be easily pardoned, when we strive to under-
stand how this animal can.catch others of better motive powers,
and feed itself by killing them when caught. Many observations
looking toward a determination of that point are narrated in a
paper entitled “Microhydra during 1907” and published in the
Proceedings of the Delaware Co. Institute of Science, Vol. III,
No. 3, issued May 15, 1908. The space allotted me in the present
publication will not allow of many of these, except to say that,
lying as it were perdue upon the surface of these stones, under
the protection of a crowded growth of Polyzoa and other localized
animals, they are very likely to be crawled over by small annelids,

148 EDWARD POTTS

and many Protozoa still smaller, whom they can paralyze with
their darts, and twist their mouths around so as to secure.

Although a stock of these interesting creatures was rarely,
perhaps never, absent from the jars upon my study table, it will
be noticed that it was not until 1897, or twelve years after their
first discovery, that medusa buds were seen to be formed upon the
hydroid stems, a millimeter in length, nor was an opportunity
found until a lapse of ten years more for a more particular study,
as will be presently described.

To supplement this, which is known as the sexual process of
its development, nature, or, more properly its Author, has provided
another, an a-sexual method of reproduction that deserves to be
at least briefly described:

We may see quite frequently, but better when Microhydra is
located upon the edge of a stone and stretches out so as to be
brought into clear view by transmitted light, depressions at both
ends of the middle-third of one side of its body. That, at the distal
end, deepens more rapidly than the other, and, by a novel method
of longitudinal fission, gradually approaches the other, the cellular
structures of both parent and larva healing up and rounding out
as the separation progresses. Finally the larva is nearly liberated
and hangs by an invisible thread, until in our jars it is by any
motion of the water waited against the glass, where it temporarily
adheres, an organism without organs, no capitulum, no pedal disc,
no apparent mouth, no means of catching prey, or feeding upon it
when caught. Plainly it is an inert, helpless body that we may |
safely call a larva, until we find, a week or ten days later, that
a capitulum has been formed, a pedal disc prepared upon which
it now stands upon the surface on which it may have been lying,
and is now prepared to sustain life on its own account. I have
watched this whole operation perhaps a score of times, and have
found the process of segmentation to take about eight hours, I
have said that during the larval period it is without organs of
locomotion, yet we have been always ready to admit that it does
move, probably by some amezboid action of its surface cells, that
my eyes have not been quick enough to catch, even with the
microscope.

FRESH WATER HYDROIDS 149

It is recognized that any suggestions to students to collect and

study this interesting group must be ineffective if they do not
include information from my experience as to their favorite living

places. Tacony Creek, although it furnished the first specimen,
is not ideal, while Flat Rock Dam certainly is such. Most suc-
cessful collectors of the plant-like fauna of our fresh waters early
learn that these prefer to grow where rapid currents bring them
a constant supply of food, and, at the same time, prevent silt from
gathering over and smothering them. The factories along the
Schuylkill Canal require much more water than is furnished by the
infrequent opening of the canal gates to let boats through. For
this reason a number of tunnels have been built to pass water from
above the dam into the canal below the first set of gates, and at

the point where they enter, at a depth of 6 or 8 feet, it becomes
the really “raging canawl’ of our derision. A dredging net of

suitable length and strength is almost practically certain to bring
up stones, large and small, covered with all varieties of the fauna
I have already mentioned. These stones are placed in glass jars
of from ¥% to 2 gallons content, and, occupying places upon my
table in a moderately warm room, rarely fail to supply me with a
healthy stock for several months.

It is possible that the general students of zoology may be in-
terested in a quotation from the paper referred to above, “Micro-
hydra ryderii during 1907,” describing the discovery of medusoid
buds and their formation:

“Tt is possible that a few medusae were seen after their first
discovery in 1897, say in 1898 or 1899, but no opportunity was
found for such observation as was above suggested until May 16th
of last year. On that morning a bud was doubtfully suspected,
watched during that day and the next, and by g p. m. of the 17th
the evidence of a coming medusa became convincing. Yet its posi-
tion on the side of a jar, and in relation to the other members of
the group, was not such as made possible the determination of the
two points named above. The ‘microscopical observatory’ had not,
at that date, been devised, and the best we could do was to stand the
jar upon the side of which the budding medusa had been detected
on a pile of books before a Welsbach gas light and examine it with

150 EDWARD POTTS

a Coddington lens or, later, through the tube removed from a com-
pound microscope, and laid across another pile of books.

“This was the situation when, at 9.30 p. m., five of us deter-
mined not to lose sight of it during the night; wherefore one or
more were continuously on the watch until 6.30 a. m. of May 18th.
The first differentiation of parts had appeared about 9.30 p. m.,
May 17th, and all hands took part, though without artistic skill
or scientific training in recording what we saw, by drawings, the
most characteristic features of which I have here preserved. An
examination of them will show the first recognizable feature to
have been the manubrium at the proximal extremity of the bud—
bearing upon its summit a circular or spherical form more or less
complete in every figure, though variable in size; whose meaning
must be left to elucidation through other specimens. Above or
beyond this there was always a light-cavity of varying size and
shape; and, almost from the beginning, transverse lines were to
be seen at the distal end of the bud, suggestive of two membranes ;
and still more faintly longitudinal lines that ultimately resulted in
becoming the radial canals. From 12.45 a. m. of May 18th and
persistently thereafter, the innermost of the transverse lines men-
tioned gave convincing proof that it was to be the velum, including
the marginal canal and circular aperture; and a few minutes later
every observer noticed more or less distinctly, upon the outer mem-
brane or surface, radial lines diverging from the apex or crown
toward the position of the marginal canal, adjoining the velum.
From 2.15 a. m. the approximately circular outline of the meduse
changed to a pear shape, widening, with nearly straight lines, from
the proximal to the distal end; and the faint lines of the radial
canals became more marked. About 4.30 a. m. pulsation or throb-
bing of the velum was observed; at first a pair, one, two; then, say
a half minute later, one; a pause, then, one, two—one, two, and so
on, very irregularly ; and thus continued, perhaps increasing in force
until 5.30, when the velum with its aperture could easily be seen,
distended, pressing up against and separating, at 6.20, the seg-
mented tentacles as shown in two excellent drawings by Miss C. W.
Beekley, as she saw them, parted, as when an orange is peeled

ee

AMERICAN MICROSCOPICAL SOCIETY ISI

from any central point down to an equatorial line, and then
forced upward by internal pressure.

“T know not what other observers may have written as to the
formation of the earliest tentacles in marine medusa; but all our
night watchers unhesitatingly agreed that my impressions of ten
years ago had been proved correct, in relation to this species of
freshwater forms. Of course, my theory assumes that the wider
portions of these wedge-shaped segments contract, or, as it were,
roll up upon themselves so as to form the nearly cylindrical tenta-
cles as we know them. I place great weight upon the simultaneous
appearance of the whole eight, without the slightest suggestion of
longitudinal growth.

“The throbbings of the velum continued irregularly after the
last drawing was made, finally liberating the medusa about g a. m.
of the same day (May 18th). Two days had passed since the first
determination of the bud, and the liberated medusa lived but two
days longer, so that this specimen did not secure us any better sight
of possible sense organs than had those seen ten years before.”

Epwarp Ports.

MUTATION IN MICRO-ORGANISMS

Dobell (Jour. Genetics, Nov. 1912 and Feb. 1913) gives a
valuable review of the literature and a summary of the conclusions
of investigators concerning mutation in micro-organisms.

In Trypanosomes (Nov., 1912,) it appears that definite struc-
tural changes may be produced by use of certain dyes, by cultiva-
tion in cold blooded vertebrates and certain invertebrates, which
changes persist through subsequent divisions and apparently do
not impair the power of division. In case of those treated with the
dyes the kineto-nucleus is destroyed. The loss of this organ seems
to decrease the virulence of the action of the Trypanosomes on the
host. Virulence is changed also by the passage of the organism
through the blood of certain animals. Resistance is developed by
them also to certain drugs which. are gradually administered. This
increased resistance is transmitted in breeding.

In respect to the Bacteria, the author summarizes his digest
in these words: “First it seems. established that the Bacteria are

152 NOTES, REVIEWS, ETC.

subject to mutation—that is to say, in a given race individuals
may occur which differ from their fellows in their genetic con-
stitution. Individuals frequently occur which possess new struc-
tural or functional features; and these features, though often the
transient peculiarities of the individual only, are in some cases
transmitted to the offspring for many successive generations. There
is reason to suppose that this phenomenon occurs in nature as well
as in laboratory cultures. The progeny of an organism which varies
may thus constitute a new race, in which every individual possesses
the new character.”

The author defines mutation as a permanent change, however
small it may be, which takes place in a micro-organism and is trans-
mitted to subsequent generations. These mutations are classed as
structural and physiological,—the latter comprising those in which
the power of producing pigments, ferments, etc., is seen.

In some instances the mutations seem to be caused by chem-
ical or other conditions of the medium; in others, in which effort
was made to secure uniformity of medium, changes still occurred
where it seems necessary to assume that the conditions of the
changes were primarily internal.

DIFFERENTIATION IN CHROMOSOMES

Agar (Q. J. M. S. Dec. 1912) reports studies of chromosomes
in Lepidosiren in which he shows that there is a widespread ten-
dency for chromosomes to be constricted or to segment trans-
versely. This is especially noticeable when the chromosomes are
short in comparison with their length. The point at which this con-
striction takes place in a given chromosome is constant for that
chromosome, and is the point at which it most readily tends to
form the angle of the V when that form is taken. The author be-
lieves that the constancy of this position denotes a constant dif-
ferentiation of the chromosomes in the long axis. The presence
of the constrictions is not, however, necessarily to be considered as
evidence of bivalency or of a future division in that plane.

AMERICAN MICROSCOPICAL SOCIETY 153

BUD-FORMATION IN SYLLIDS

Potts (Q. J. M. S. Jan. 1913) proposes the following classifi-
cation of budding found in these worms :—

1. Linear budding (terminal). Stolons produced at the end
of the stock, and arranged end to end in chains, e. g. Autolytus.

2. Lateral budding. Stolons produced singly as lateral out-
growths from the stock. Syllis ramosa.

3. Collateral budding (ventro-terminal). Stolons produced
from a ventro-terminal proliferating cushion on the stock, and ar-
ranged side by side in rows. Trypanosyllis gemmipara.

In this latter genus the author has made a study of the forma-
tion of the buds. The process is as follows:—The leucocytes col-
lect in the mesoblast of the posterior segments where the buds are
to appear; the epiblast gives rise to cell proliferations which be-
gin the stolons; the mesoblast from the stock invades these young
stolons; the mesoblast proliferates and shows its first signs of
segmentation in the form of incipient septa; two bundles of muscle
fibres and a single ventral nerve cord grow directly from the cor-
responding structures of the stock into the stolon; the epiblast of
the stolon segments and forms its appropriate segmental structures.

Those who have studied the formation of new segments in the
segment-forming zone of worms will be impressed with the simi-
larity between the processes.

CORK OAK IN PORTUGAL

. Klein (Naturwiss. Zeitschr. Forst- und Landwirtsch. Nov.

1902) discusses the cork oak and its products in Portugal in a
very interesting article. As is well known this oak, Quercus suber,
is native to Southern Europe. In Portugal there are some 550,000
acres of this oak. During the first 20 to 25 years its growth and
cork production are rapid, and at the end of this period a crop
may be gathered. The cork oak forests are mostly private prop-
erty and are rented for periods of 20 to 40 years, or worked by the
owner. The most of the old forests are natural; but of recent years
new forests are being planted of acorns selected from known pro-
lific trees. In such plantations the first crop may be had in 10
years.

154 NOTES, REVIEWS, ETC.

MISTLETOE OF THE INCENSE CEDAR

Meinicke (Proc. Soc. Am. Foresters. Mch. 1912) has a very
interesting study of the California mistletoe, which is parasitic on
the Incense Cedar. This mistletoe is a small hanging shrub pro-
ducing barrel-shaped swellings on the trunks of such trees as have
been long infected. Account is given of the examination of some
of these swellings as old as 350 years and 45 inches in diameter.

The living “sinkers” of the mistletoe were found 7% inch long,
extending into the sap-wood and going through 19 rings. The dead
“sinkers” were also to be seen persistent in the heart wood. In
one tree at a point where there was no infection during the first 37
years, as shown by the inner rings, the last 219 years show con-
tinuous infection.

The parasite begins on the young tree as a semi-parasite with
green leaves. The enormous development of the bark gradually
eliminates both the green shoots and the aerial haustoria, and leaves
the plant with a widespread root system extending into the tissues
of the host, apparently without serious injury to it.

A SIMPLE METHOD TO REMOVE PARAFFIN SECTIONS WHICH ARE
STUCK TO A SHEET OF PAPER OR TO THE HAND

Ribbons of paraffin sections temporarily set aside on a sheet
of paper frequently adhere so firmly to the paper as to be unde-
tachable without special means.

A simple method to remove such sections I have found to be
as follows :—

Cut out a piece of the paper together with such a length of the
ribbon as is desirable for mounting ‘on one slide. Drop 50-75%
alcohol on the piece so that the alcohol may diffuse under the paraf-
fin sections. As soon as the paper underlying the ribbon is soaked
with alcohol immerse the piece gradually in water. The ribbon
will float off and may be drawn up on a wet slide and mounted in
the usual way.

In cases where sections accidentally adhere to the hand, drop
some alcohol so that it may diffuse under the sections. They may

then be easily removed. Ropert CHAMBERS, JR.,

Biological Department, University of Cincinnati.

AMERICAN MICROSCOPICAL SOCIETY 155

TO KILL MOSQUITOES OR OTHER INSECTS

Mix equal parts of 90% alcohol and a 1:500 aqueous solu-
tion of HgCl2. Gently boil the insect in this for a minute or two
to expel the air in the trachee. As the solution cools it is drawn
through the stigmata into the body of the insect to all the tissues.
Leave for a few hours, then pass at proper intervals through 90%
alcohol, absolute, oil of turpentine, and paraffin.

R. Ross says this is particularly good for salivary glands of in-
fected mosquitoes, as the Sporozoa are well preserved.

In case it is desired to mount whole a larva or small adult
insect, after killing as above, use Farrant’s Medium. Ring with
Hollis’ glue.

Abstracted by V. A. Latham.

TO KEEP SLIDES AT CONSTANT TEMPERATURE

Use a sheet of copper 15 inches long, 3 inches broad, and 1-12
inch thick. Support on 2 or more suitable feet and place a small
lamp beneath. In this way graduation of temperature can be had
by varying the height, and at different distances from the heated
point. Boa i
V. A. LatHam.

SECTION CUTTING IN GELATIN BY FREEZING

Gaskell (J. Path. and Bact. July 1912) recommends cutting
certain materials by freezing in gelatin rather than by the usual
processes. It is claimed that it avoids distortion such as occurs in
use of fluids like alcohol and xylol, and also the vacuolation found
in paraffin preparations. The fats are of course preserved. It is
valuable in examining small objects, and objects with loose tissues
like lung and tissues liable to disintegration. It is especially use-
ful in examination of lung in broncho-pneumonia, as the contents
are preserved im situ. Many other similar occasions of usefulness
are cited. It prevents the disintegration often resulting from ordi-
nary methods in pancreas, liver and spleen and the like.

The most important item is to get the proper consistency of
the gelatin.

156 NOTES, REVIEWS, ETC.

Fix in some formal mixture, as 10% formalin in Millers
fluid; wash thoroughly (over night or equivalent time), as formol
will act on the gelatin and prevent penetration.

Tear up and soak Gold Label gelatin in water from 1 to 4
minutes, depending on room temperature. Squeeze the gelatin by
hand, place in beaker, cover and melt in paraffin oven till viscid.

Transfer to ordinary incubator at 37° C., take tissue from

water, dry or blotting paper, drop into gelatin, and leave for 2
hours.

Imbed the tissue in a paper box in some of the gelatin in
which it has soaked. Let the mass set at room temperature. Harden
for 3 or 4 days in vapor of formol, supporting above the fluid in
suitable way. It may be left in this state indefinitely, or be stored
in 5% formol. Return to the vapor for a few days before cutting.

When ready to cut pare the block and place in water I-10
minute, depending on hardness, before freezing. The freezing
microtome recommended is one by Aschoff made by Sartorius of
Gottingen.

Various methods of successful staining are also given.

Abstracted by V. A. L.

HOUSEHOLD BACTERIOLOOGY

This book is the outgrowth of courses in the subject given in
the department of Home Economics at the Iowa State College of
Agriculture. It is somewhat more than its name implies. It is
rather a study of micro-organisms from the point of view, first, of
the general student, and, second, of the student of the economic
and sanitary applications.

The treatment is compact, and the authors succeed admirably
in securing an intelligible discussion of an enormous number of
phases of the subject. It appears to the reviewer as one of the
most teachable of the books on the subject of economic micro-
biology.

After an overbrief opening chapter introducing the subject
in a historic way, Sections I-III, consisting of 19 chapters, follow,
dealing with the general considerations (1) of Morphology and
Classification; (II) of the Technic of Culture, Sterilization, and

AMERICAN MICROSCOPICAL SOCIETY 157

observation of Micro-organisms, and (III) of their Physiology
and Ecology. In addition to the chapters on classification, which
are remarkably clear and helpful to the beginner, an illustrated key
of some 35 pages is given for the identification of the families and
genera of the common molds. This without a doubt will add greatly
to the usefulness of the book to the college student and to the in-
dependent worker. The authors have done for these household
micro-organisms exactly what the American Microscopical Society
is trying to do for the common genera and species of microscopic
and near-microscopic plants and animals, on behalf of its members.
They deserve the thanks of teachers for this contribution to the
teaching of this interesting subject.

Section IV deals with Fermentation both from the scientific
and the economic viewpoint. It treats the general phenomenon of
fermentation, enzymes of micro-organisms, relation of these to
food preservation, the changes in organic substances as sugars,
milk, celluloses, gums, fats and nitrogenous compounds through the
action of micro-organisms.

Section V treats of Micro-organisms and Health. Here are
included the expected subjects: Theories of disease, resistance of
the body to disease, organisms normal to the body, classification
of disease-producing organisms, the various special types producing
special disturbances in the body. In addition there are chapters on
water contamination, examination and purification; contamination
and examination of air; contamination and examination of milk and
of other foods.

Household Bacteriology; Buchanan. Cloth, 8 vo., 536 pages and index. Illustrated.
The Macmillan Co., New York. 1913. Price $2.25 riet.

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NECROLOGY

GroRGE C. CRANDALL, M. D.

Dr. George C. Crandall of St. Louis, Mo., who became a
member of the American Microscopical Society in 1904, died re-
cently at the age of 47 years, from Bright’s disease, after an illness
of two months.

He was born at Elgin, Ill., received his education at the Uni-
versity of Michigan,-and began the practice of medicine at St.
Louis in 1895. At the time of his death he was Professor of
General Medicine in the Medical Department of St. Louis Univer-
sity, was president of the consulting staff of the City Hospital,
medical director of the St. Louis Society for the Relief and Pre-
vention of Tuberculosis, a member of the St. Louis Medical
Society, Missouri State Medical Association, American Medical
Association, Medico-psychological Association, Academy of Science,
Civic League, Citizens’ Industrial Association, and various social
and business clubs.

He is survived by Mrs. Crandall, who was Miss Nellie Merry
of Syracuse, N. Y., and one son, 16 years of age.

A. B. AUBERT

Professor Alfred Bellamy Aubert was born in New York
City in 1854. He was educated in the Columbia grammar school,
in the Lycee of Strasburg, France, and received his B. S. in chem-
istry at Cornell University in 1875. Upon graduating from college
he was appointed to the Chair of Chemistry at the University of
Maine, which position he held for thirty-five years, retiring June,
1910, on account of failing health. :

On assuming his position at the University of Maine, chem-
istry was taught only as a culture study, but through his efforts

160 NECROLOGY

a strong chemical course was developed, from which a large
number of well-known chemists have been graduated.

As a teacher he had the ability to inspire in his students the
utmost confidence. He was a member of several chemical and
microscopical societies, to whose journals he contributed many
interesting and valuable articles. As a man he was modest, retiring
and unassuming, diffident almost to bashfulness, respected and
beloved by all who knew him.

His death occurred November 12, 1912.

BonNER N. McCraAvEN

Mr. Bonner McCraven of Houston, Texas, who joined the
society in 1904, died during August, 1912.

TRANSACTIONS

OF THE

American Microscopical
Society

ORGANIZED 1878 INCORPORATED I89QI

PUBLISHED QUARTERLY

BY THE SOCIETY

EDITED BY THE SECRETARY

VOLUME XXXII

NuMBER THREE

Entered as Second-class Matter December 12, 1910, at the Postoffice at Decatur, Illi-
nois, under act of March 3, 1879.

Decatur, ILL.
Review Printinc & STATIONERY Co.
1913

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OFFICERS.

President: F. CREIGHTON WELLMAN, M. D................ New Orleans, La.
Fagst Vice President: EB. Gp WaAttE, PRDiie cs. cecccnces ss Cleveland, Ohio
Second Vice President: H.E. Jorpan, Ph.D............. Charlottesville, Va.
SECEELOLY = Us: We GAETOWAY: coats csrstetccis ols's so Santee baivic dame eaes Decatur, Ill.
ixeastrers| Ula EAN KINSON tt. \t5 a2 ace cals oe owaoea oes ee Charleston, Ill.
Gusistase- “MAGNUS. PLATE. 020. oe coc led asei ok oe wha ea we Meadville, Pa.

ELECTIVE MEMBERS OF THE EXECUTIVE COMMITTEE

EES SS ae ee ee ae Bureau Plant Industry, Washington, D. C.
Ha ANGEARS RO 0G ts CAR AE ying) Stee ae arctan a and ae mae a Ea Re aad Laramie, Wyo.
LEO ep By COG ENTE etc ei a cia ee cai SSIS A we aes ha Lawrence, Kans.

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

Apert McCatta, Ph.D., of Chicago, IIl.
at Chicago, IIl., 1883

T. J. Burr, 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.
Srmon Henry Gace, B.S., of Ithaca, N. Y.,
at Ithaca, N. Y., 1895 and 1906.

A. CuiirForp Mercer, M.D., F.R.M.S., of Syracuse, N. Y.,
at Pittsburg, Pa., 1806.

A. M. Buiette, M.D., of Columbus, Ohio,
at New York City, 1900.

C. H. E1icenMANN, Ph.D., of Bloomington, Ind.,

at Denver, Colo., 1901.
CuHartes E. Bessty, LL.D., of Lincoln, Neb.,
E. A. Brirce, LL.D., of Madison, Wis.,

Henry B. Warp, A.M., Ph.D., of Urbana, IIL,

at Pittsburg, Pa., 1902.
at Winona Lake, Ind., 1903.

at Sandusky, Ohio, 1905.
Hergert Oszorn, M.S., of Columbus, Ohio,

at Minneapolis, Minn., 1910.
A. E. Herrzzer, M.D., of Kansas City, Mo.,
at Washington, D. C., 191.
F. D. Heap, Ph.D., of Philadelphia, Pa.,
at Cleveland, Ohio, 1912.

The Society does not hold itself responsible for the opinions expressed
by members in its published Transactions unless endorsed by special vote.

TABLE OF CONTENTS

FOR VOLUME XXXII, Number 3
The Spencer-Tolles Fund: Grants to be Made From it, by the Com-

BIMGECE «55 SF he, Seah ee LG a Sa simr voters tests eae ik nena este dares aie ee ae 165
Notes on the Trematode Genus Clinostomum, with Plate IX, by William
Walter Ont. a oicke bere TN COG ni ban en aete oilers Cre ae 169

Notes, Reviews, Etc—1. Notes on Rhizopods from Michigan; 2. Specula-
tions on the Nature of the Olfactory Organs; 3. Vaginicola, an Inter-
esting Protozoan; 4. An Inorganic Cell, by E. W. Roberts (with
Plates X-XII). Polyembryony in the Nine-banded Armadillo;
Persistence of Bacillus abortivus in Tissues; Persistence of Tubercle
Bacilli in Cultures; Cambium Growth in American Larch; Cell-
division in the Sex Cells of Taenia; Demonstration of Brownian
Movement; Fresh-water Diatoms and Their Preparation; Mount-
ing Fresh-water Algae; Mounting Volvox; Counting Leucocytes in
Cerebro-spinal Fluid; Toxic Secretions of Infusoria; Relations of
Cell Size and Nuclear Size in Oxytricha; Influence of Mating in
Paramecia; Amitotic Division in Ciliated Cells; Spermatogenesis in
Silkworms; Education of Infusoria in Ingestion of Food; Spirostyle
in Spermatozoa; Nerve Fibrils in Dentine; Succession in Fungi;
Rusts and their Host Tissues; Physiological Effects of Bordeaux
Mixture; Hermaphroditism in Amphioxus; Resistance in Hiber-
nating Animals; Microscopic Measurements by Camera Lucida;
Micro-radiography; Circulation by Convection Currents in Labor-
atory Aquaria; Simple Histological Methods; Reducing Stock Solu-
tions; Laboratory Guide in Parasitology; Prevention and Control
GE WUISEASEY Bot oe ok ois sdk os ew eb pee eS <UL S Ue Ce nee eee 183

TRANSACTIONS

OF

American Microscopical Society

(Published in Quarterly Installments)

Vol. XXXII JULY, 1913 No. 3

THE SPENCER-TOLLES FUND

The Spencer-Tolles fund was established by the American
Microscopical Society for the encouragement and furtherance of
research, especially among its members and in lines of study in
which the microscope is the principal instrument of research. While
the total has not yet reached the limit desired to give an income
effective for the purposes which the founders had in mind, yet the
committee to whose charge the Spencer-Tolles fund was entrusted
feel that the amount of the fund is now sufficient to warrant the
society in appropriating regularly some part of the income for the
encouragement of research while the remainder will still be added
to the principal until the fund is completed. In order that the
Society may be familiar with the situation and with the conditions
laid down for securing appropriations from the fund the commit-
tee has formulated the following rules for its guidance in making
grants. These received the approval of the Executive Committee
at the annual meeting last December. They are intended to guide
the committee in its action until experience demonstrates the advisa-
bility of changes; but they will not be modified without due notice
to the Society and adequate opportunity for discussing from every
standpoint any proposed alterations. The present financial status of
the Spencer-Tolles Fund may be ascertained by consulting the report
of the Custodian (Trans., v. 32, p. 86), and its growth is readily seen
by comparing these reports for a series of years. They are published
annually and give a full account of the Fund to date of the meeting.

It is not the intention of the committee to hamper the investi-
gator by limiting the use of grants to very precise purposes or to
control narrowly their expenditure, but to allow the fullest freedom
for the exercise of individual judgment in particular cases. As

166 SPENCER-TOLLES FUND

appropriate purposes in definite cases are recognized the collection
or preparation of material for work, the construction or purchase of
special apparatus, provision for field expenses, payment of services
for control, observations or experiments, and the preparation of
illustrations or other special expense incident to the publication of
the research in proper form. Since the amount now available is
small the committee is inclined to favor those definite purposes
which may be served by the appropriation of smaller sums and is
especially favorable to making such a grant in connection with funds
available from college, university, or private sources where it will
enable the investigator to complete a task already begun or to pub-
lish work already finished but delayed by the expense of printing.

In this connection attention is directed to grants already made
from this Fund. No one of them was large enough alone to attain
the results actually achieved, but in each case the completion of the
work would not have been possible without such assistance as the
Fund gave. In view of the large sums dispensed by other scientific
organizations for experimental purposes in research the commit-
tee believes that aid in this direction is less needed, and is inclined
to emphasize the aid to publication which it is in position to offer and
which is not often given by other agencies. For the present, the
committee will not limit its grants exclusively to those given to aid
in the publication of research papers, even though it feels the need
of such aid and the great advantage which will accrue to the entire
Society as well as to the individual member by adopting such a
policy.

REGULATIONS GOVERNING GRANTS FROM THE SPENCER-TOLLES EUND

1. The Committee will receive formal application for grants
from the Spencer-Tolles Fund at present only from members of the
American Miscropical Society.

2. Under ordinary circumstances not more than $100 will be
voted in any one year to research purposes. Under the Constitution
of the Society no money can be granted for any other purpose from
the income of this Fund.

3. Applications for grants shall be filed with the chairman of
the committee, who shall at once communicate all the facts to other

SPENCER-TOLLES FUND 167

members and after their discussion and action shall inform the appli-
cant of the result.

4. Each applicant shall submit to the committee a record of
his professional or academic training and of any other research
work already done. He shall name three persons qualified to speak
through personal knowledge of these facts, especially concerning
his ability to carry on investigation successfully.

5. Each applicant shall outline the topic on which he seeks
the assistance of the Fund and shall indicate the manner in which
he proposes to expend the grant asked, the reasons for seeking aid,
and the results he expects to attain by this aid.

6. On completion of the work each recipient of a grant shall
give the committee a report of the use to which the grant allowed
has been put, preferably in the form of a paper ready for publica-
tion and embodying the results of the work in connection with
which the grant was used.

7. Every grant is made upon the express condition that all
results obtained by its aid shall be offered to the American Micro-
scopical Society for publication in advance of their announcement
elsewhere. In case the Society or its properly constituted authority
is unable or unwilling to undertake the publication of the complete
work, then permission will be granted to publish elsewhere; but
wherever published publications including the results of this work
shall contain the distinct statement that the work contained in the
paper was done with the aid of a grant from the Spencer-Tolles
Fund of the American Microscopical Society.

8. Payment of a grant shall be made by the Custodian on
certification of the chairman that the committee has approved the
grant and in accord with the specifications made by the committee
for this particular grant. Payment of any sum will ordinarily be
made, one half when the grant is approved and one half when the
report is received and accepted by the committee. But the special
circumstances associated with an individual grant may lead the
committee to modify this rule and provide some other manner of
payment in that instance.

g. The expenditure of the money shall be entirely in the con-
trol of the person receiving the grant and he shall not be asked to

168 SPENCER-TOLLES FUND

secure or furnish any vouchers covering the expenditure in detail.
On completion of the work he shall file with his report a statement
that such a sum, mentioning the amount, has been expended and
the results of the work are contained in the accompanying report.
Any unexpended balance retained by the custodian in making the
final payment, or if paid out of the grant not covered by this state-
ment shall be returned to him and shall be again placed in the Fund.

(Signed) Henry B. Warp, Chairman
S. H. GAGE
Macnus PFLAM
H. R. HOWLAND
A. M. BLEILE

NOTES ON THE TREMATODE GENUS CLINOSTOMUM

By WILLIAM WALTER CortT*

During the fall of 1911 I found encysted in the mesentaries and
under the peritoneum of several specimens of the leopard frog
(Rana pipiens) a number of larval distomes belonging to the genus
Clinostomum Leidy. Recently more cysts were found in a frog of
the same species from North Judson, Indiana. Two specimens of
this genus from cysts in the black bass (Micropterus sp.) near
Washington, D. C., were sent me by Dr. B. H. Ransom, and some
of the same kind of material was given me by Dr. George R. La
Rue, which he had found in perch (Perca flavescens) from Douglas
Lake, Michigan. Additional material of the larval stages of this
genus was turned over to me by Professor Henry B. Ward from
his private collection. This material was from the following hosts
and localities: From cysts in frog, Oshkosh, Wisconsin; from
perch (Perca flavescens) and blue gill (Lepomis pallidus), Bass
Lake, Michigan; from rock bass (Ambloplites sp.), Alma, Michi-
gan; from perch, Lake Spooner, Wisconsin; from black bass (Mic-
ropterus sp.), Sebago Lake, Maine. Also two adults were collected
from the mouth of a black-crowned night heron (Nycticorax nycti-
corax naevius) which was taken alive near Urbana, Illinois, and
given me by Professor Frank Smith.

I wish to express my appreciation of the helpful suggestions
and criticisms given me by Professor Henry B. Ward during the
preparation of this paper.

During the past year I have been engaged in a comparative study
of the above material. Two recent papers by Osborn on the North
American representatives of the genus Clinostomum have covered
much the same ground as my studies. The first of these (Osborn,
1911), considers the distribution and behavior of this type of tre-
matodes, and goes over the literature thoroughly. The second

*Contributions from the Zoological Laboratory of the University of Illinois, under
the direction of Henry B. Ward, No. 25.

170 W. W. CORT

(Osborn, 1912), takes up their anatomy in so much detail that fur-
ther morphological description seems unnecessary at the present
time. Although in the main my observations confirm Osborn’s, I
must dissent from his conclusions in regard to the forms from the
frog. Osborn finds so little difference in form and proportions of
the body between the late immature stages from cysts in the fish and
frog and the mature worms from the heron, that he considers all the
flukes of this genus so far collected from North America to belong
to the same species, Clinostomum marginatum Rudolphi. I find
that the specimens from the frog show so many differences that I
am forced to consider them as a distinct species. The Clinostomum
from the bittern (Wright, 1879) also shows several important points
of difference from the adults from the heron, and is probably the
adult stage of the frog species.

Distomum reticulatum described by Looss (1885), a larval
Clinostomum from Costa Rica (not Porto Rica, as stated by Os-
born and Linton), has given rise to considerable discussion. This
form is much larger than any of those from North America, has a
lobed ovary, and the cirrus sac extends to a point posterior to the pos-
terior testis. Osborn (1912: 219-220) doubts whether these differ-
ences are great enough for specific distinction, and states that too
much importance should not be attached to differences in shape of an
organ like the uterus or its parts. He further states that writers from
Leuckart down have considered Distomum reticulatum Looss to be
identical with the North American representatives of the genus
Clinostomum. This statement is not correct since Braun (1900:
44-45) states that Distomum reticulatum is distinct from the North
American species, and relates the former closely to Clinostomum
sorbens, a South American species which he described. I agree with
Braun’s conclusion. Since Looss’ name for this form was preoccu-
pied Monticelli renamed it Mesogonimus dictyotus. But he over-
looked Leidy’s earlier genus Clinostomum; therefore Looss’ Costa
Rican trematode should be known as Clinostomum dictyotum.

In the study of the North American representatives of Clinos-
tomum very little attention has been given to those from frog hosts.
MacCallum (1899) reports them, and Osborn (1911) describes
their position in the cyst. He also includes them in his later paper

GENUS CLINOSTOMUM 171

on the structure of Clinostomum marginatum, although apparentiy
all his anatomical descriptions and figures represent specimens from
the black bass and heron. Both the above authors have referred
this form to Clinostomum marginatum. I propose to show that
the Clinostomum form from the frog belongs to a distinct species,
which may be named Clinostomum attenuatum, since it is the most
slender of all the species so far described in this genus.

The late larval stages of Clinostomum attenuatum have been
found encysted only in frogs, and of Clinostomum marginatum only
in fish. The cysts in the frogs are scattered in the mesentaries, em-
bedded under the peritoneum of the body cavity and in the lymph
spaces between the skin and muscles of various parts. They are
never found within the muscle tissue surrounded by its fibers. The
cysts from the fish in my experience and Osborn’s are always found
in the midst of muscle bundles.

Osborn (1911) notes the position of the worm in the cyst as
the only difference between the forms from the fish and frog. I
have found this condition to be dependent upon the location of the
cyst. In those cysts embedded in the muscles of the fish where there
is considerable pressure from all sides, the worm is folded three
times with the acetabulum to the outside and is very tightly enclosed
by its cyst. Where the cysts are loosely held in the mesentaries of
the frog and subjected to little if any external pressure, the worm
is very loosely enclosed in its cyst, and is usually folded but once.
But in those cysts which are located in the frog where they are
subjected to pressure as between the large muscles of the upper
thigh, the worm is folded three times and compressed tightly in its
cyst, offering then much the same appearance as the fish cysts.

Differences are to be noted between the two species in size and
shape. In length the range of variation within each species is so
great that no constant difference is found. The specimens from the
frog ranged from 3.9 mm. to 5.52 mm. in length. Among those from
the fish the smallest individual measured 3.5 mm. and the largest
6.6 mm. in length. In body shape the differences are very distinct
and constant. The frog type is slender and of almost uniform width
and thickness throughout its length (Fig. 1), while the fish type
has a broad rather flat post-acetabular region (Fig. 2). In Clinosto-

172 Wim W. CORT

mum attentuatum the pre-acetabular region is oval, measuring on
the average about 0.67 mm. in width by 0.41 mm. in thickness, while
the whole body back of the acetabulum has a thickness greater than
half its width. The average measurement for the region of the
ovary are about 0.68 mm. in width by 0.37 mm. in thickness. In
Clinostomum marginatum from the fish cysts the pre-acetabular
region is more nearly cylindrical, the average width for my speci-
mens being 1.05 mm. and the average thickness 0.85 mm. Just back
of the acetabulum the ratio of the width to the thickness is about
three to two, the average width being 1.35 mm. and the average
thickness 0.91 mm. At the ovary, however, this worm is wider
and more flattened than the other species, having an average width
of 1.55 mm. and a thickness of 0.71 mm. The difference between
these two species in relative width of the different body regions is
even more pronounced. In the frog forms the width of the pre-
acetabular region is equal to or only a little less than that of the
post-acetabular, while in the fish forms it is only a half or two-
thirds as great. In the first species the post-acetabular region has
about uniform width throughout its length, while in the second it
is considerably wider at the ovary than just back of the acetabulum,
and becomes narrower toward the posterior end. These relations are
illustrated fully in the measurements given in the table.

Braun (1900), in his diagnosis of the species of Clinostomum
lays considerable emphasis on the structure of the anterior tip.
This region is truncated obliquely ventrad so that the dorsal sur-
face extends somewhat further forward than the ventral. Braun
calls this surface the oral field. In the center of this field on the
oral cone is the oral sucker. Surrounding the oral cone in both
Clinostomum attenuatum and Clinostomum marginatum is the fur-
row and the projecting margin described by Braun in some of the
species of this genus. Braun’s suggestion that the whole oral field
is used as a sucker has been confirmed by Osborn (1911: 368), and
is fully discussed later in this paper. In Clinostomum attenuatum
the oral cone fills a greater portion of the oral field than in Clinosto-
mum marginatum, whereas in the latter the furrow is deeper and
wider and the projecting margin higher. Measurements of four
individuals of like contraction express the first difference. In two

—s

———

GENUS CLINOSTOMUM 173

specimens from the frog the oral cone measured 0.45 mm. and 0.52
mm. in diameter at its base, and the oral field had a width of 0.56
mm. and of 0.65 mm. making the ratio very nearly 4:5. But in two
specimens from the fish the oral cone measured 0.60 mm. and 0.67
mm., and the oral field 0.93 mm. and 0.97 mm. in diameter, the
ratio in this case being about 2:3.

Further differences between these species are found in the
relation of the length of the pre-acetabular region to the total length
of the body, and in the position of the genital glands. In Clinosto-
mum attenuatum the region in front of the ventral sucker is short,
being only about one-sixth or one-seventh the total body length,
while in Clinostomum marginatum it is one-fourth or one-fifth. To
determine the position of the genital gland field the distance from
the posterior edge of the acetabulum to the ovary was measured. In
the frog type the ovary is always back of the middle of the post-
acetabular region, and in some cases the whole genital gland field is
back of that point. In the fish type this condition is almost always re-
versed, the ovary in my specimens being at about the middle or in
front of the middle of the post-acetabular region. Reference to the
table will make these differences more clear.

In the genus Clinostomum the uterus empties into a sac which
extends longitudinally from the genital pore anteriad toward the
acetabulum (u s Fig. 1 and 2). In the larval stage this sac is very
narrow and contracted. In both the species under consideration the
uterine sac has about the same length, but its position in respect to
the other organs is different. In Clinostomum attenuatum the dis-
tance from the posterior edge of the acetabulum to the anterior tip
of the uterine sac is quite great, being almost equal to and in some
cases exceeding the total length of the sac. But in Clinostomum
marginatum the anterior tip of the uterine sac comes very close to
the acetabulum and in the contracted specimens almost touches it.

Depending upon the great variation in size of the animals, the
diameter of the suckers varies rather markedly in these two species.
In spite of this variation distinct differences can be traced. In
Clinostomum attenuatum the oral sucker measured on the average
0.19 mm. in transverse diameter and in Clinostomum marginatum
0.30 mm. The acetabulum in the first species measured on the

174 W. W. CORT

average 0.56 mm. and in the second species 0.73 mm. In the ex-
treme individuals, however, the sizes overlap. The difference be-
tween these two species in the ratio of the size of the suckers is
constant. In Clinostomum attenuatum the acetabulum is about three
times as large as the oral sucker, while in the Clinostomum margi-
natum the ratio is about 2.4 to I.

The most clear cut specific difference between these two forms
is found in the difference in the structure of the cuticular spines.
In Clinostomum attenuatum the spines range from 0.013 mm, to
0.016 mm. in length and from 0.005 mm. to 0.009 mm. in thickness
at their bases, and in Clinostomum marginatum they vary in length
from 0.007 mm. to 0.011 mm. and in thickness from 0.0015 mm. to
0.002 mm. In both species they are about equally numerous in a
given area but on account of their greater width they appear more
thickly set in the flukes from the frog. Figures 4 and 5 show this
difference in size more clearly than any description.

The differences given above between the advanced larval stages
from the frog and from the fish seem to me to be so distinct and
so far reaching, that it is impossible to consider these two forms
as belonging to the same species.

MacCallum (1899) and Osborn (1912) have worked out the
anatomy of the adult Clinostomum marginatum from North Amer-
ica in considerable detail, so that I shall give only a sufficient de-
scription of the specimens collected from the black-crowned night
heron to determine their relationship to the larval forms. The fol-
lowing measurements were taken from the worm shown in figure 3:

Waa: a et 5 et ee Mee ORES La EE EE 3.4 mm.
Wadth: at tlic: atetror end .6 2056.5 . ie eee 067°"
Width half way from the anterior end to acetabulum.... 0.63 “
Width At “arctan... . «. ico ean ttgupategasace eee O56
Width half way from acetabulum to ovary .............. O82) is
MPA OR SOCEM RESE | 0s 5 a cunt a neine Eee cee O21) as
Width half way from ovary to posterior end ............ ORF. 7
Length’ of pre-acetabular region 2.146.500; aca AGA e077 455
Length of post-acetabular region «......<.)-cccseersee000~ 2A ye
Distance from acetabulum to ovary ...........ss.eeeee- 0.075
Distance from ovary to posterior end ................... 1.4 4
Eegoth OT MEE TSE . ices arcane cate lt ee EER O45 ee
Distance from anterior tip of uterine sac to acetabulum... 0.07. “
Length ‘of *genttal ailand: field) 500. es Oe eee 0.76 = “

GENUS CLINOSTOMUM 175

Mransverse diameter of Joralustickety. Got c. snacsseeas ove ONT hs
diransvetse diameter of, acetapiuluineas enh access eins ase rome (0 a
Ratio of oral sucker to acetabulum ................. 1tO123 oe
Weneths ob eggs (average) secss cere coca tee oe prsice aials 0.091 “
Wadthcoimesos: (Caveragein sa titan rsie tera niels ate sleclore stor OOSka

These measurements agree so closely with those given above
for the advanced larval stages from the fish that both forms must be
considered as different developmental stages of the same species.
This adult was very small and in length and size of suckers falls
slightly below the smallest of the larval forms. The anterior end is
more attenuated in the adult than in any of the larval forms which
I examined, but this is due to difference in contraction. On the other
hand it can be seen that the larval stage from the fish and the adult
from the heron are almost exactly alike in the general shape of the
body, the proportions of the various regions, the position and con-
figuration of the genital organs, and the ratio in size of the suckers.

Osborn’s description of Clinostomum marginatum from the
bass and heron agrees in all points with my forms from similar
hosts. Since he observed more individuals he notes a greater range
of variation than was found in my material. His worms vary in
length from 3 mm. to 8.2 mm. and in greatest width from 0.7 mm. to
2.2mm. It is of interest to note that Osborn (1912: 191) also finds
his smallest individuals among the adults, indicating a still greater
variation than he has found for the larval stages.

That the fluke described by MacCallum (1899) as Clinosto-
mum heterostomum is really Clinostomum marginatum cannot be
doubted. He notes individuals up to 10 mm. in length and in his
drawing of a toto preparation the ovary is back of the middle of
the post-acetabular region. In all other respects his description
of this worm agrees with Osborn’s and my own for Clinostomum
marginatum. These observations and comparisons seem to show
that all the representatives of the genus Clinostomum which have |
been found up to date in North American fish and herons belong
to the one species, Clinostomum marginatum.

Clinostomum marginatum was originally found in Brazil. Braun
(1900) compared Rudolphi’s type specimens of Clinostomum margi-
natum from Brazil with material gathered by Natterer in the same
country from several different localities and hosts. He states that

176 W. W. CORT

all this material corresponded so closely to Rudolphi’s type speci-
mens that it must be considered as belonging to Clinostomum mar-
ginatum. After a careful comparison with Braun’s descriptions, I
can find no constant differences between the North American forms
just considered and these Brazilian forms except in the greater size
of the eggs of the latter. In the specimens from South America
the eggs vary in length from 0.104 mm. to 0.140 mm. and in width
from 0.055 mm. to 0.073 mm. The largest measurements recorded
for the eggs of North American forms is 0.099 mm. in length by
0.066 mm. in width. It will be seen that in width this falls well
within the range of variation of the South American flukes, and is
only slightly less in length. Braun’s material from Brazil agrees
so exactly with Osborn’s and my specimens from North America,
that his figures 8, 9, or 20 (Braun 1900), might be used to illustrate
our descriptions. |

Clinostomum marginatum is thus a species of very wide distri-
bution, having been reported from Brazil in South America, and
from North America very widely. It has also a wide range of hosts.
The advanced larval stages have been reported from cysts in the
pike, the perch, the blue gill, the black bass, the rock bass, the sunfish
and the trout. The adult has been found in three different genera
of water birds: From three true herons, Ardea sp. (Brazil), Ardea
cocot, and Ardea herodias; from one species of stork, Mycteria
americana, and from the black crowned night heron, Nycticorax
nycticorax. The presence of Clinostomum marginatum in both
North and South America is easily explained by the great range of
of the adult host. Ardea herodias and Nycticorax nycticorax nae-
vius for example both range over North America at large, and
central and northern South America.

There remains for consideration the Clinostomum described by
Wright (1879) from the American bittern. Wright’s description
and drawing of this form differs in several respects from Clinosto-
mum marginatum. In his drawing the genital field is shown behind
the middle of the post-acetabular region, the uterine sac reaches
only about half way from the genital pore to the acetabulum, the
pre-acetabular region is only about one-sixth of the total body length,
and the worm is rather long and slender with fairly uniform width

GENUS CLINOSTOMUM 177

throughout. All these points agree with Clinostomum attenuatum
rather than Clinostomum marginatum. Since frogs form an irh-
portant part of the food of the American bittern, Wright’s Clinos-
tomum from this host might well be the aduit Clinostomum at-
tenuatum.*

While collecting two adult Clinostomum marginatum from the
heron, I was able to make some observations on the activity and re-
lation to its host of the living parasite. The two flukes were below
the average in size for this species and had only a few eggs in
their uteri. The activities of the parasite were studied both while
in the mouth of the heron, and after removal into normal saline
solution. The heron came into my hands still alive and not more
than five minutes elapsed between the killing of the bird and the
finding of the flukes.

In the bird’s mouth the worms were very much contracted, and
adhered so firmly to the mucous membrane that it was very difficult
to loosen them. Their position was well suited to resist the friction
of food taken into the mouth of the heron. Not only was the ace-
tabulum firmly attached but also the oral field functioned as a
sucker. The pre-acetabular region was bent over so that the oral
field was almost in contact with the acetabulum. The anterior end
was given very firm attachment by the sucking action of both these
structures. The post-acetabular region was much contracted longi-
tudinally and arched so that it was quite convex. The edges were
pressed closely into the mucous membrane, and evidently by the
drawing up of its central part this whole region also acted as a
sucker. In fact the posterior end was so firmly attached that it was
almost as difficult to loosen as the anterior. Such a sucking activity
of the post-acetabular region accounts for the great development of

*Since the completion of the above observations I have received from Professor
Henry B. Ward material of the genus Clinostomum collected by A.“L, Cooper from the
vicinity of Go-Home Bay, Zoronto, Canada. This material, which was both larval and
adult, was collected from two fish hosts—Perca flavesens and Micropterus dolomieu,—
one frog host—Rana catesbiana,—and one bird host—the American herring gull (Larus
argentatus). A careful examination, using for species determination the points brought
out above, showed that the specimens from the fish and bird belonged to the species
Clinostomum marginatum, and the form from the bull frog to Clinostomum attenuatum.
This gives additional data to support the hypothesis, that Clinostomum marginatum in
its advanced larval stages is limited to fish hosts and Clinostomum attenuatum to frogs.
Also this collection adds a new genus to the bird hosts of Clinostomum marginatum, and
extends the list of hosts of Clinostomum attenuatum to two frog species.

178 W. W. CORT

the dorso-ventral parenchymous muscles which has been noted for
this species.

The only reference in literature to the position of the Clinosto-
mum in its bird host is by Osborn (1911: 363). He found speci-
mens of Clinostomum marginatum adhering to the heron’s throat
only by means of the anterior end. As the bird had been dead for
a day or two, the worms were probably beginning to loosen their
hold. In his later paper on this form Osborn (1912: 193) writes as
if attachment in this species was effected by the anterior end alone.
He says that the reason for the large size of the ventral sucker had
not been indicated by the behavior of the worm, and that although
the structure of this sucker suggests full functional power he had
noticed no activities for it. My observations show that the ace-
tabulum is not only fully functional in this species, but that it plays
a very important part in holding the parasite in position in the
mouth of the host.

After the position of the worms had been noted they were re-
moved and observed for some time in normal saline solution. The
living fluke was a semi-transparent whitish cream color, with the
testis and ovary showing as opaque pure white areas. The intes-
tinal ceca were dark brown, and the region surrounding the uterine
sac was light pink. The animal manifested considerable activity
(Figs. 6 and 7), the whole body expanding and contracting rythmi-
cally. In its most contracted state the total length of the worm
was about 2.5 mm. with a width at the region of the ovary of 1.56
mm. When most extended it reached a length of 4.96 mm. with a
width at the ovary of 0.92 mm.

A study of the movements of the worm after removal from its
host confirmed the observations made on its sucking activities while
in position, and suggested a possible method of locomotion. This
fluke has the long distance from the stomach to the mouth of the
bird to travel, after the larval forms have been freed from their
cysts in the fish by the digestive juices. In the most contracted
position the anterior end was much shortened and turned ventrad,
the oral field with the oral cone at its center being in the same plane
and very close to the acetabulum. The edge of the oral field was
very mobile and strong sucking movements were noted. This posi-

GENUS CLINOSTOMUM 179

tion of the anterior end suggests that which was noted when the
animal was attached to the mucous membrane of the heron’s mouth.
In the contracted position the post-acetabular region was very short
and broad and so arched up as to be very thin. The edges showed
considerable movement, curling in and then extending. The position
and movements of this region suggested that if its edges were in
contact with a soft surface considerable sucking power would be
developed, just as in a small boy’s leather sucker. This position of
greatest contraction I shall call the sucking position. It was the
assumption of this sucking position which made the removal of the
worms from the mucous membrane of their host so difficult. The
need of such a strong sucking reaction on the part of the parasite is
apparent, when it is considered that whole fish and other very hard
particles of food are taken into the mouth of the heron.

In the series of rythmical contractions made by this fluke as it
lay free in the salt solution was suggested a possible method of
locomotion. The cycle of expansion and contraction was as fol-
lows: From the sucking position the pre-acetabular region would
stretch out and the oral field go through sucking movements. At
the same time the post-acetabular region would be extended, reach-
ing its greatest length just as the pre-acetabular region was begin-
ning to contract. The worm would then contract again into the
sucking position and a new cycle of movement would be initiated by
the extension of the anterior end. Sucking movements of the aceta-
bulum were noted during this process.

When the larval trematodes incysted in a fish eaten by the
heron are liberated in its stomach by the action of the digestive
juices, they turn toward the esophagus. The series of rythmical
movements described above is begun, and when the pre-acetabular re-
gion is extended and turned ventrad, the oral field comes in contact
with the mucous membrane. The sucking movement causes it to
take hold and on the contraction of the pre-acetabular region, the
worm is pulled forward until the acetabulum comes into close con-
tact with the oral field. The acetabulum then takes hold in its turn
and on the next extension of the anterior end holds the ground
gained. By a laborious repetition of these movements the worm
could make its way up to its final position. If any food coming

180 W. W. CORT

down the esophagus should strike the parasite it would con-
tract strongly, assume the sucking position, and hold on with its
whole body until the way was clear again. Having once gained its
final position, the adaption of the whole body for sucking would en-
able it to hold its position against the friction of the heron’s food.

TABLE OF MEASUREMENTS

tnostomum inostomum
attenuatum marginatum

co atl hee 2 So ee

1 Na Hs

>
—)
-}

1 FT Oe Re Be SARS ROCCO ae ei
Width wat vanteriana (Oto. cas came aie einer
Width half way from anterior end to

BCCTAUITITIN on ais ae nie aie s hiele bicelo eels
Wadth: at acetalmltims sc cccciicislssjeu sax cme
Width half way from acetabulum to ovary
WAGE Cab (OVANY.s ao ccraje'w eins ties olc/eretotaveo in ara
Width half way from ovary to posterior end
Length of pre-acetabular region..........
Length of post-acetabular region.........
Distance from acetabulum to ovary......
Distance from ovary to posterior end.....
Length of uterine Sac.......-.cccrcccees
Distance from anterior end uterine sac

EG WACELAUIN II | 1c iolcicie's ely a's)o'e.s/ainiete'= pa
Length of genital gland field.............
Transverse diameter of oral sucker......
Transverse diameter of acetabulum......

So
©
w
cS
Se RR eh eles
awn

ON FURR SINTER =D 00
>

Shoo sor ewooseocs SS
AEUDA YORBNADRAAAD Wo

PENS SSO So (SM
BND BKANWERE DRO PS

SrdfPOSCSSSSO
neo DH wWUNom mbt into
NOR OCOW He LHhOL
esss
Ome OC RFORBANNADAN Ue
eess Serressssss Sf
DRA VYROEURDUADD Ww
ONO FRADE UROUOF NM
eeeor Sdy-Foooses SoM
DAWUS BWwEUNYODLDO Ao
Diwiow NOOHERANHN OD

SS Sree roes
ND NUWOOR NR OwO
Hh RNOMORUNIOR

OORA MAD Woo
Se PS PSE Ne Soees Sa

ANwoh NWO HhROWODH
eae Si ale a ne
OOH CWOANANMNAW OND
NNAOD DAWOUDWeRNHrRK WOO Ne
ROMO FRPHAOMSMN NOM)
UNAKH DHWOwnwowwwn
OfOW Chee COM ROO

mre COM tO 00t1wWOO

Sees
NNO

The above table gives measurements from ten representative
toto mounts, five of Clinostomum attenuatum and five of Clin-
ostomum marginatum.

GENUS CLINOSTOMUM 181

LITERATURE CITED

Braun, M.
1900. Die Arten der Gattung Clinostomum Leidy. Zool. Jahrb.,
Syst., 14 :1-48.
Looss, A.

1885. Beitrage zur Kenntniss der Trematoden. Distomum palliatum
n. sp. und D. reticulatum n. sp. Zeit. f. wiss Zool., 41 :390-446.

MacCatium, W. G.
1899. On the species Clinostomum heterostomum. Jour. Morph.,
15 697-710.

Oszorn, H. L.
1911. On the Distribution and Mode of Occurrence in the United
States and Canada of Clinostomum marginatum, a Trematode
Parasitic in Fish, Frogs and Birds. Biol. Bull., 20 :350-366.
1912. On the Structure of Clinostomum marginatum, a Trematode
Parasite of the Frog, Bass, and Heron. Jour. Morph., 23 :189-223.

Wricut, R. R.
1879. Contributions to American Helminthology. No. I. Proc.
Canad. Inst., 1 :54-75.

182 W. W. CORT

EXPLANATION OF PLATE IX

Figures 1-5 were drawn with a camera lucida.

Fic. 1. Clinostomum attenuatum from Rana pipiens. Larval speci-
men. X 57.

Fic 2. Clinostomum marginatum from Perca flavescens. Larval speci-
men. X 57.

Fic 3. Clinostomum marginatum from Nycticorax nycticorax naevius.
Adult. X 57.

Fic. 4. Cuticula and spines of Clinostomum attenuatum. X about 1000.

Fic. 5. Cuticula and spines of Clinostomum marginatum. X about 1000.

Fics. 6 and 7. Free hand drawings of a living specimen of an adult
Clinostomum marginatum expanded and contracted. X about 33.

ABBREVIATIONS USED IN PLATE

a, acetabulum
i, intestine

om, oral mound
0, Ovary

os, oral sucker
t, testis

u, uterus

us, uterine sac
v, vitellaria

PLATE IX

DEPARTMENT OF NOTES, REVIEWS, ETC.

It is the purpose, in this department, to present from time to time brief original
notes, both of methods of work and of results, by members of the Society. All members
are invited to submit such items. In the absence of these there will be given a few brief
abstracts of recent work of more general interest to students and teachers. There will be
no attempt to make these abstracts exhaustive. They will illustrate progress without at-
tempting to define it, and will thus give to the teacher current illustrations, and to the
isolated student suggestions of suitable fields of investigation.—[Editor.]

AMATEUR MICROSCOPISTS

The following group of four notes, by Mr. Roberts, represent
work done by an enthusiastic photographer, who is greatly inter-
ested in studying and photographing histological and cytological con-
ditions. They represent a type of worker and of work which the
American Microscopical Society wishes to encourage. So much
expert work is done with the microscope in our great laboratories
that too many students come to feel that good work cannot be done
away from them. It is the hope that amateur workers will come
more and more to use this department, and make it helpful to other
amateurs.

I. NOTES ON RHIZOPODS FROM MICHIGAN

During the summer of 1912 monthly collections at the same
locality were made. These collections were killed and fixed at
early morning hours, stained in iron-hematoxylin, dehydrated and
carried into xylol. Some of the material was mounted whole in
balsam and other portions imbedded and sectioned.

Many varieties were studied, but the common Arcella vulgaris
was the easiest material for observation, both on account of its
shape, ease of identification and number of individuals found.

The Arcella is very polymorphic in form, and has a number
of different life phases which are not altogether understood.

The life cycle is a year in length, and the majority of individuals
arrive at sexual maturity in May and June. From then onward
only straggling mature individuals are found.

The ordinary method of propagation is by gametes. These are
furnished with a pair of flagella by which they move about. See

184 NOTES, REVIEWS, ETC.

Fig. 1-4 and 2-A. These gametes are macro- or female, and micro-
or male. Shey conjugate and form a new individual which, after
a resting stage, develops into a mature Arcella. Along in October
the young individuals begin to form the chromidial net from which
the future generations are cut out.

1-A. Arcella—male gamete.

2-A. Arcella—female gamete.

3-A. Arcella—conjugation of gametes.

4-A. Arcella—resting stage.

5-A. Arcella—early vegetating stage.

6-A. Arcella—later vegetating stage.

7-A. Arcella—showing details of chromidial net formation.
There are 12 chromosomes which bud out chromidia into the cytoplasm, these
subdivide and form the net.

8-A. Arcella—showing the formation of the flagella fibrils from the centrosomes.

9-B. Clathrulina in asexual series.

10-B. Clathrulina in resting spore stage—asexual.

This stage was observed as shown in Fig. 7-A in the diagrams.
Certain chromidial bodies are budded out from the nucleus into
the cytoplasm, where they undergo repeated subdivisions until their
progeny form a dense network of deeply staining granules con-
nected by threads which follow the divisions. This process con-
tinues slowly until the warmth of Spring hastens feeding, which
increases nourishment and brings the metabolism of the organism
to a climax.

When the net is fully formed it undergoes constrictions in the
night, forming at intervals masses of the chromidial granules. See

AMERICAN MICROSCOPICAL SOCIETY 185

Figs. 1 and 2, Plate X. The inclosed granules now bud out a certain
number of granules which unite and form a nucleus for the mass.

The formation of the flagella by repeated divisions of centro-
some bodies was observed. This agrees in all essential details with
the later observations of the origin of motile organs on various
type of plant and animal sperms. See text figure 8-A.

This end, bearing the flagella, is the mouth end of the adult
rhizopods, so that in all forms, both naked and shelled, there is
always a definite polarity or relation of the nucleus and cytoplasm.
See Fig. 5-A.

The Rhizopods are now considered to be degenerate flagellates.
The flagellate gamete stage gives the clue to their line of descent.

These sexual gametes escape when mature, leaving temporary
scars in the matrix in which they were bedded, as is shown in Fig.
3, Plate X.

There was seen occasionally an alternate sexual generation, or
process of schizogony. This stage or form is produced by the
constriction of the chromidial net into masses. These masses have
at this stage no nuclei, each mass being inclosed by a network of
regularly placed granules connected by threads. These granules,
bud inward, forming granules which group and form the new nuc-
leus, while they themselves form the cytoplastids, or vegetative
systems. See Fig. 4, Plate X.

Thus we see the cell formation processes reversed from the
common method of budding from the nucleus outward. This seems
to the writer to show that the chromidial elements of the cyto-
plasm are of the same rank as the nuclear chromidia, being capable
of reversing their generations in either direction.

This seems to represent a degenerate function which is only
repeated at rare intervals, as but very few are to be found.

Then there is a process of blastogamy, in which an individual
deserts its shell and unites with another individual in its shell, the
two forming a joint network. This chromidial mass later breaks
up into swarm spores.

The peculiar markings on the chitinous shell are produced by
cyto-somes, which take nuclear stains, and the membrane on which

186 NOTES, REVIEWS, ETC.

the chitin is deposited is easily observed in many specimens where
the individual is shrunken away from the wall by the reagents.

Many fine specimens of our only form of fresh water Poly-
cistinee, the Clathrulina, were found, and several interesting phases
of their life history studied.

They form a fenestrated silicious shell on a long stalk, as shown
in Fig. 5, Plate X. An interesting phase of their asexual repro-
duction was observed. The body divides, part escaping from the
shell. This part then forms a new shell for itself. The stalk of
the new individual forms in attachment to the old shell, and by
growth gradually elongates until adult size is reached when the pro-
cess is again repeated. As many as eight individuals were found
thus formed a series. See text figure 9-B.

They also reproduce sexually by motile macro- and micro-
gametes, which escape and conjugate and after a resting stage fol-
low the usual route to maturity.

An interesting asexual stage was found, the body breaking
down into three stellate resting spores, much resembling those of
Desmids. See Fig. 6, Plate X. The cell wall is dissolved and the
spores escape and are found in abundance in the mud.

As late as October they still remain in this stage, very likely
spending most of the winter thus. Early Spring collections ought
to show good stages of their development.

2. SPECULATIONS ON THE NATURE OF THE OLFACTORY ORGANS

In the vast families of insects and among other nearly related
animals are found certain organs called antenne. These are situ-
ated on the head and are generally conceded to be modified limbs,
of which each segment of these forms of bodies once possessed
a pair.

Such a modified pair of legs from a Black Syrphus Fly is
shown in Fig. 1, Plate XI. Here, at least three segments of the
limb remain. The second segment is greatly expanded laterally,
forming a bulb much flattened, to which is attached the third seg-
ment in the form of a long whip-like filament.

AMERICAN MICROSCOPICAL SOCIETY 187

This modified limb is supposed to be the seat of several senses ;
indeed, in some insects each segment is credited with a different
function by some writers.

These limbs, originally used for walking, were doubtless pro-
vided with various sensory cells adapted to the necessities of their
peculiar functions. The sensory cells are usually in the form of
hypodermal spines of various shapes, covered with a chitinous ex-
terior to give them greater firmness.

All the living cells of animal bodies are supposed by many
students to be connected by minute filaments with the nervous sys-
tem. Thus all cells are potential sensory cells, both those on the
interior and exterior of the body.

By looking at the enlarged lobes of the antennz of the fly in
Fig. I, it will be seen to be covered with small spines. Also there
will be seen near the base a dark circular pit. On many insects
these are very numerous and are considered as olfactory in func-
tion. These olfactory pits are invaginations of the hypoderm, and
the spines on the exterior are sunk into the pits.

If we look at Fig. 2, which is a section of these pits on the
antenne of a fly, called Sarcophaga, we will see the pointed spines
which are the sensory structures.

The sensory cells being thus sunk are much protected so that
their chitin envelop may be but feebly developed, leaving them
nearly naked and therefore more sensitive.

So we have here two kinds of sensory cells, the external exposed
cells, and the sunken or protected cells.

It is apparently these same hypodermal spines which form
the rhabdome rods in the optic invaginations of insect eyes.

If we now look at another form of limb called a palpus (See
Fig. 3), from a moth, Pieris raphae, we will see the end of the organ
is invaginated into a pit. Into this pit will be seen projecting the
sensory scales, while below is the connecting nerve cord.

In Fig. 4 we get a view of the elaborate development of these
pits on the antenna of a honey bee. Here the same spines are seen
in the pits, also the elaborate nerve connections.

Figure 5 is a cross section of the olfactory pits on the antenna
of a wasp, Vespa. The outer openings of these pits are closed by

188 NOTES, REVIEWS, ETC.

elongated lips of chitin, while the pits proper are more round in
shape. The naked tips of the sensory spine cells are seen in cross
sections in great numbers. While these pits are regarded as olfac-
tory, it may well be that the olfactory sense is not confined to the
pit spines alone, but is only more sensitive here because of the thin-
ness of the chitin. It may occur in some degree over the exterior.

On many Lepidoptera the males have these external sensory
scales so well developed that they are enabled to pick up the trail
of a female of their species by the scent she leaves in passing through
the air. In such cases the antenne are largely developed, while the
shoulders and fore limbs also are covered with a special growth of
scales which seem actively to function as olfactory organs.

A moth called Chyfolisa morbidalis has in the male a brush of
enormous scales on each front leg, which exceeds the entire com-
bined size of the head and antennz. In this case the whole front of
the body becomes a quivering agency of sex determination.

It seems reasonable that this is a case of animal tropism, the
pathway of chemical particles left in the air by the female acting on
the sensitive spines much as light acts on the eyes.

It is possible that this chemical sensitiveness to particles in the
air is partly responsible for the irregular flight of such animals.

3. VAGINICOLA; AN INTERESTING PROTOZOAN

The form described herein is apparently related to the Vorti-
cellidee, a family of infusorians remarkable for beauty and variety.
From their shape these animals are often called Bell-animalcules.
They are attached, either temporarily or permanently, and often
have a distinct stalk.

They are usually marked histologically by a long ribbon-shaped
nucleus, a circle of vibratile cilia around the oral end, and by a
lengthwise binary division as one of the methods of its multipli-
ction.

The form described here was found June, 1912, in collections
from Goguac Lake, near Battle Creek. It is free, has a capsule,
seemingly of a chitinous or horny nature possessed of an oval aper-
ture. In all the specimens studied there is a pair of individuals in
each capsule.

AMERICAN MICROSCOPICAL SOCIETY 189

Figures 1 and 2, Plate XII., are photographs of longitudinal
and cross-sectional views, respectively, of the animal. Figures 3
and 4, Plate XII, are diagrams simplifying and interpreting the for-
mer figures. From these the main details of the anatomy can be
made out.

The twin individuals arise apparently by binary division of the
parent. Further multiplication is by motile gametes, which bud
from the adult. These arise in a string and suggest ova in higher
animals.

The oval cilia are not in a wreath form, but line the gullet into
which they are retracted when not in use.

The body wall contains fibrils similar to those found in the
Vorticellidz.

4. PROCYTOS VULGARIS; AN INORGANIC CELL

There is a question which often comes into the mind of stud-
ents of cytology: ‘““Where does the cell form come from? What was
its origin and what relation does it bear to organic and inorganic
nature?”

There is a growing number of students of natural phenomena
who are diligently striving to show the relations of the organic
world to the inorganic. To such the following study may be of
interest.

A look at the photo (Plate XII., Fig. 5), which I have called
Procytos vulgaris, shows a cell structure which bears a striking re-
semblance to many animal and plant tissues.

Here are to be found wholly inorganic formations which sug-
gest cells with cell walls, nuclei, nucleoli, filaments and cyto-plasm
bodies. These structures are purely inorganic and can be produced
in the liquid form of any material, by observing certain conditions
of temperature.

The material used in the present experiment is ozokerite, a
refuse product from oil refineries in the form of wax. If this is
melted and poured on a hot plate and allowed to cool slowly we get
the effect pictured above.

The Giant’s Causeway in Ireland is a somewhat similar produc-
tion on a large scale, in an ancient outpoor of lava or melted rock.

190 NOTES, REVIEWS, ETC.

The cells are centers of boiling by which heat is conducted from
the base to the air, where the material is cooled and then moves
downward again.

Diagram of Circulation in the
Inorganic Cell

By consulting the diagram there will be seen two streams of
fluid of opposite polarity. One stream of positive heat units goes
upward in the center of the cell while the cooled units descend at
the margin. Particles of various kind which may be in the fluid
are carried along by the movements of the streams.

This boiling is easily seen in the fluid preparation as long as
the material stays within a certain range of temperature.

The cells should in theory be correct hexagons, but some cells
are more vigorous in action than others, resulting in the encroach-
ment on the domain of other cells, thus deforming them.

The descending streams of cooled units deposit particles of
various materials in the cell walls where they are cemented up by
congelation into substantial walls.

The nucleus-like spots are caused by introducing bodies that
do not readily mix with the wax—in this case air bubbles. These

AMERICAN MICROSCOPICAL SOCIETY I9g!t

become the center of a secondary activity. When the cooling has
progressed far enough they collapse, forming the crater-like circles
in the cells. The nuceoli are secondary eruptions of the air.

The markings within, which suggest cytoplasm, are caused by
crystals of some fat which solidifies at a temperature different from
the mass with which it is mixed.

Now, let us compare this structure with the similar characters
of living cells and see wherein they both show dependence on natural
conditions. First, we have here in both cases a definite enclsoure
of certain activities. In both cases one form of this activity is heat;
only within certain bounds of temperature are the activities of either
possible. With the lowering of the temperature the components
congeal, with the raising they disintegrate.

Second, the cells are rudely hexagonal in form. This follows
from the association of semi-fluid bodies, their mutual pressure de-
termining their shape.

Third, the formation in both cases, by precipitation, due to con-
gelation or other process, of cell walls at the boundaries of the cell.

Fourth, a definite circulation of the cell contents which lasts as
long as the components are in their requisite relations.

This may be plainly seen in plant cells where the protoplasm
streams outward from the nuclei, bathing the cell wall on the side
toward the source of heat and returning to the nuclei on the cooled
wall.

Fifth, the formation of nuclei of substances of a nature differ-
ent from that of the surrounding material, which results in definite
secondary activities of a complicated kind.

Sixth, the precipitation and coagulation of various substances
in the cytoplasm when they reach critical temperature points.

We thus see that there are structures and functions which
occur in both the organic and inorganic world in quite similar ways.
It is quite impossible to escape the conviction that forces that act
in the one are also real causes in the other.

It is not pretended that the processes in this inorganic material
are identical with those in the living cell, nor that temperature is the
only factor in organic activity.

192

NOTES, REVIEWS, ETC.

There are many types of fluid-formed crystals now known;
some 300 or more. These are molecular arrangements of inorganic

substances.

Modern theories of crystal formation point to a fluid

pre-crystal stage in which the components are adjusted into the rela-
tions in which they congeal.

When more definite studies of pre-crystal stages are to be had,
we do not doubt that conditions similar to these we have described
in this inorganic cell will be found.

Battle Creek, Mich. E. W. Roserts.

Fig.
Fic.
Fic.
Fic.
Fie.
Fie.

Fic.
Fic.
Fic.
Fic.
Fic.

Fic.
Fie.

An hw Pw

AEA spre eam

1
2.

EXPLANATION OF PLATES

PLATE X—Rhizopods of Michigan
Arcella vulgaris; constrictions of chromidial net into gametes.
Arcella vulgaris; network matured into gametes.
Arcella vulgaris; escaping gametes have left scars.
Arcella vulgaris; asexual individual.
Clathrulina; adult asexual form.
Clathrulina; resting spore stage, also asexual.

Piate XI—Nature of Olfactory Organs
Antenne of Black Syrphus Fly.
Section of sensory pit in Sarcophaga.
Section of palpus of Pieris raphae.
Section of antenna of Honey Bee.
Cross-section of olfactory pits on antenna of Vespa.

PLaTe XII

Vaginicola; photograph of longitudinal section.
Vaginicola; photograph of cross-section.

Fic. 3. Diagram of details in Fig. 1: A, sheath; B, aperture of same;
C, the individual animal; D, nuclei (macro and micro) ; E, gametes; F, fibrils

in ectoderm.

Fic. 4. Diagram of details in Fig. 2. Lettering as in Fig. 3.

Fic. 5.

Photograph of ozokerite cooled in such a way as to suggest cells.

PLATE X

PuatTe XI]

PLATE XII

aA
= ies?

~~
8
a

ik

AMERICAN MICROSCOPICAL SOCIETY 193

POLYEMBRYONY IN THE NINE-BANDED ARMADILLO.

Newman (Am. Nat., Sept. 1913) brings together many inter-
esting facts concerning the biology of the Texas Armadillo, gath-
ered from numerous papers which he has published previously.
Since the paper is itself a summary, it is impossible here to sum-
marize it.

Some of the most interesting facts relate to polyembryony.
There are four embryos, which are enclosed in a common chorion.
These four are always of the same sex. They are formed from a
single egg fertilized by a single spermatozoon. The cleavage gives
rise to an inner cell mass and blastodermic vesicle similar in all es-
senital respects to the rodents. It is in connection with mesoderm
formation that the writer finds first two, and then four, buds with-
in the general vesicle. The writer believes that this breaking up
of embryonic activity, which might naturally be expected to con-
tinue as a unit. may be caused by lowered vitality, due to an egg
parasite, coupled with an external pressure on the developing em-
bryo exerted by a groove in the uterine wall. The latter tends
though pressure to isolate the two halves of the embryo, and be-
cause of the low vitality it is not able to unify these bilatcral growth
activities.

Many interesting questions concerning heredity, sex-determi-
nation, and the like, offer themselves in connection with this very
favorable material. For example, it is clear that these four embryos
are much closer in kinship than is true of members of the same
litter in mammals generally. Furthermore they occur in pairs which
are mutually more alike than they are like the other pairs.

Each embryo ultimately develops its own independent connec-
tion with the mother, and they often differ signally in their nourish-
ment, as shown by the rate of development. Since the quadruplets
are invariably of the same sex, irrespective of their size, it is clear
that the sex is determined before the isolation of the four centers
of growth in the vesicle.

The female diploid number of chromosomes is 32, reducing to
16. In the male the diploid is 31, reducing to 15 and 16. This
duplicates the conditions described in other vertebrates.

194 NOTES, REVIEWS, ETC.

The same author (Jour. Exp. Zool., Aug., 1913) discusses in a
much more extended way the many interesting questions of heredity
suggested in the more general article.

PERSISTENCE OF BACILLUS ABORTIVUS IN TISSUES.

Fabyan (Jour. Med. Research, May, 1913) presents facts to
show that B. abortivus has a quite prolonged life in the tissues of
apparently healthy laboratory animals—as guinea pig, rabbit, mouse,
rat, pigeon, etc. In one instance they were harbored without any
external signs of ill effects for 67 weeks. Two additional conclu-
sions seem warranted from the experiments: First, that there
seems to be at least a slight temporary multiplication of the germs
after inoculation ; and, second, that the animals are not without the
power slowly to destroy the bacilli.

The study is interesting as bearing on possible periods of en-
durance and latency of pathogenic bacteria after the disappearance
of the symptoms of the disease.

PERSISTENCE OF TUBERCLE BACILLI IN CULTURES

Smith (Jour. Med. Res., May, 1913) tests the current view
that tubercle bacilli lose their vitality in cultures in periods of I to
6 months. He found that cultures which completely ceased to mul-
tiply on the artificial media under wholly favorable conditions were
still infectious to guinea-pigs for from 7-19 months. This was true
both of human and bovine strains; though of the two types when
reared side by side the bovine is the more resistant. It is true that
the number of bacilli surviving in such cultures is very small. The
series of biological facts is suggestive: Tubercle bacilli (bovine),
which on removal from the diseased animal do not at first multiply
on glycerine agar, may in time become partially saprophytized and
grow luxuriantly on such culture media; gradually this culture
medium fails to serve their purpose, and most of them die; as long
as vitality lasts the fresh tissues of the guinea-pig furnish an ade-
quate medium for their restoration.

CAMBIUM GROWTH IN AMERICAN LARCH

Knudson (Bul. Tor. Bot. Club, June, 1913) presents a study
of the American larch in respect to place and time of beginning of

AMERICAN MICROSCOPICAL SOCIETY 195

cambial activity, the relation of xylem and phloem formation, and
the like. As the result of two years’ studies he concludes, for this
species, that the development of the phloem precedes that of the
xylem; that the first increase in diameter of xylem begins a few
meters below the apex; that the development of the xylem begins
about a month later than the leaf formation, altho there seems, ac-
cording to other investigation a diameter increase about the time
the leaves appear. This, the author thinks, is due mainly to a
swelling of the tissues, of the turgor type.

It is suggested that temperature of the soil, moisture of the
air, the thickness and color of the bark, as well as other unknown
factors may determine the place and date of diameter increase.

CELL-DIVISION IN THE SEX CELLS OF TAENIA

Harmon (J. of Morph., June, 1913) presents evidence that the
division of the spermatogonial cells and the two spermatocyte divi-
sions in Tznia are mitotic. In the ova mitosis is frequent and there
is no evidence of amitosis in the oogonial division; the maturation
divisions are mitotic, and mitotic divisions occur both in early and
late cleavages. The author believes that there is no reason to be-
lieve that amitotic division occurs in this animal—contrary to the
conclusions of earlier studies. She believes that the close contact
of nuclei and other items that have been interpreted as meaning
amitosis are due merely to special conditions of mitosis—as the
nuclei dividing more rapidly than cytoplasm, shortness of cleavage
spindle, swift reconstruction of nucleus after splitting of chromo-
somes, rapid growth of daughter nuclei, etc.

METAMORPHOSIS OF FILARIA LOA

Dr. Leiper (Lond. Sch. Trop. Med., Jan., 1913) telegraphs from
Calabar that the Metamorphosis of Filaria loa has been proved to
take place in the salivary glands of a fly belonging to the genus
Clorysops.

DEMONSTRATION OF BROWNIAN MOVEMENT
Mr. Travis of the Queckett Club describes a satisfactory method

of demonstrating striking Brownian movements. Rub a_ small
amount of gamboge for a few moments on an ordinary microscope

196 NOTES, REVIEWS, ETC.

slide. Place a drop or so of water where the gamboge has been
rubbed. Gently push the edge of a cover glass up to the gamboge;
with suitable illumination the whole field is seen in very brisk
motion.

FRESH WATER DIATOMS AND THEIR PREPARATION

Groom (Eng. Mech., March, 21, 1913) invites the microscopist
to the study of the Diatoms. He quotes: “Contemplation of the
netted beauty of some small part of nature may bring man, as he
grows older, the admiration of a vision of the whole,” and thus
advises: Go to a pond, collect a quantity of Pond weed. Wash this
by shaking thoroughly in, say a wide-mouthed 2-pound jar nearly
filled with water. Pick out all of the weeds possible, shaking to
free them of the diatoms. Allow to settle for two hours and pour
off all the water possible. Put the sediment into a saucer with a
fair quantity of water and place it in a sunny window. Next morn-
ing take a fine piece of muslin big enough to cover the saucer; soak
it, wring it out, place it over saucer; press it down so that it
rests lightly on the surface of the sediment. After a time the free
diatoms will make their way through the meshes of the muslin and
discolor it. The diatoms are thus collected on the upper surface of
the muslin free from the sandy debris. They can be mopped from
the muslin into clear water with a camel’s hair brush.

MOUNTING FRESH-WATER ALGAE

English Mechanic (Nov., 1912) makes the following sugges-
tions for beginners. For unicellular alge (diatoms, desmids, etc,).
remove as much water as possible from them with a pipette, trans-
fer to a watch glass of filtered rain water or clear water of the kind
they live in. Carefully stir up the sediment, allow to settle, remove
the water and the very surface of the sediment with a pipette. Re-
peat this process until a supply has been obtained freed from the
sand and mud. Remove water and add a fluid made up of alcohol, 3
parts, water 2 parts, and glycerine I part. Stir the specimens well
and leave it in an open watch-glass for 10 days or more until the
alcohol and water are evaporated—nearly covering the vessel to
keep out foreign dirt. Do not use heat to hasten evaporation. Mount
in warm, not hot, glycerine jelly.

AMERICAN MICROSCOPICAL SOCIETY 197

MOUNTING VOLVOX

English Mechanic (Feb. 14, 1913) offers the following method
for preparing this interesting organism. [If in sufficient quantity
pour through a fine cloth funnel. After a couple of minutes place
the cloth in tube or dish of 2% per cent formol. Fix for about 5
minutes. Remove the muslin containing the material, which will
now be a gelatinous mass of volvox and other small organisms that
may have been present.

To remove volvox from muslin, place some 2% formal in a
slightly sloping white plate. Open out the muslin and place the
volvox in contact with the fluid. A sable brush and a hand lens
should be used to push each volvox off the cloth and up the inclined
plate out of the formol. If there is much dirt pour off the old fluid,
wipe the plate and replace with clean formol.

Transfer to ringed cell with pipette, and mount. Care is
necessary to prevent drying out and the formation of air bubbles.

COUNTING LEUCOCYTES IN CEREBRO-SPINAL FLUID

Chauvet (Le Monde Med.) refers to the necessity of technical
exactness in this process for diagnostic and other reasons. The
small amount of the fluid available is a complicating fact. He de-
scribes at some length the two methods in use: Centrifuging, and
Nageotte’s Cell Method.

The centrifugation method involves centrifuging, the clotting
of the cellular elements, dropping clot on slide, drying it in open air,
fixing with absolute alcohol or ether, staining in aqueous eosin
(1 per cent) or with hematoxylin, mounting in cedar oil. Under
normal conditions one finds not more than 2 or 3 mono-nuclear
cells on an average in a I-12 immersion field. It would imply a patho-
logical lymphocytosis if 7 or 8 lymphocytes were to be found in
that space. It is quite clear that the method is open to such pos-
sible errors that enumeration by means of it must be uncertain in
diagnostic value.

Nageotte’s Cell Method involves the use of a cell of known
dimensions. Its bottom is ruled into minute parallelograms. This
cell is completely filled with cerebro-spinal fluid to which a minute
amount of Unna’s blue has been added for staining. The filled cell

198 NOTES, REVIEWS, ETC.

is placed on the movable stage and allowed to rest for five minutes
until the lymph cells settle. The cells in a few adjacent rectangles
can be readily counted by means of Leitz No. 5 or 6 objective. The
number per unit of volume can be computed from the known depth
of the cell and the size of the parallelograms.

The latter method is much more exact and convenient.

TOXIC SECRETIONS OF INFUSORIA

Woodruff (Jour. Exp. Zool., May, 1913) reaches the con-
clusion that Parameciam and some of the hypotrichous protozoa
excrete substances that are toxic to, and tend to inhibit the rate of
reproduction in their own species. These products are specific in
their action since their presence does not uniformly influence the
rate in a species other than that producing it.

It is apparent that this factor is necessarily one of importance
in determining the continuance of cultures and the succession of
organisms in them.

A few pedigreed specimens of Paramecium were placed in cul-
ture media, which differed only in that one had contained a rich cul-
ture of paramecia for several days, another had contained a similar
culture of hypotrichs, and another had no protozoa. All the media
had the same bactrial flora.

RELATIONS OF CELL SIZE AND NUCLEAR SIZE IN OXYTRICHA

Woodruff (Jour. Exp. Zool., July, 1913) finds that there
is great variation both in the actual size of nuclei and cells and in
the size of these in relation to one another in all the periods of the
life of the race. That is to say there is no absolute or relative size
which is characteristic of any age. The average size of the nucleus,
and of the cell itself is smallest, and the proportion of nuclear to
cytoplasmic matter is highest during the period of greatest reproduc-
tive activity. The author believes that the mass relations of nucleus
and cytoplasm are not the determining features of reproduction.

INFLUENCE OF MATING IN PARAMECIA

Jennings and Lashley (Jour. Exp. Zool., April and Aug., 1913)
find that the conjugation of paramecia has a distinct influence upon

AMERICAN MICROSCOPICAL SOCIETY 199

the character of the descendants of both conjugants. These two
lines of offspring descended from a pair of conjugants are more alike,
both in their rates of fission and in the length of their bodies. There
is a correlation, as has been previously seen, in the body length of
parents, owing to assortative mating. But the correlation between
the offspring of parents that have mated is 48 per cent greater than
that between the parents themselves, which shows inheritance in the
offspring from both parents.

AMITOTIC DIVISION IN CILIATED CELLS

Jordan (Anat. Anzeig. XIII, 1913, p. 598) contributes to the
study of the behavior of ciliated cells, a report of the epithelial cells
in the epididymis of the white mouse. He finds that division here
is exclusively amitotic. Not a single mitotic figure was seen; but
all stages of direct nuclear division are found. The prevalence of
amitotic division has been shown in the epididymis of other animals,
in the ciliated cells of the trachea, and in the ciliated cells of the
gills of the clam. Jordan believes that the loss of power of mi-
totic division in these ciliated cells is due to the fact that the cen-
trosome, whose activity institutes indirect nuclear division, is used
up in the formation of the basal granules from which cilia are de-
veloped. In a way the power of mitotic division is the price they
pay for cilia.

SPERMATOGENESIS IN SILKWORMS

Yatsu (Annot. Zool. Japan., Vol. VIII., Pt. IL, July, 1913)
undertakes to find whether there are any chromosomal differences
between the various races of silkworms that are correlated with the
morphological differences. He studied in all some seventeen do-
mestic varieties of Bombyx—Japanese, Corean, Chinese, Turkish
and European. His results were negative; that is to say, he found
no differences of shape, size or number in the chromosomes of the
morphologically different races of domestic worms. The haploid
number he finds to be 28; the unreduced number is therefore 56.

The wild silk worm, Theophila mandriana, however, has 27 as
the haploid number. If therefore, as some writers think, the wild
form is the ancestor of the domesticated races, the latter have

200 NOTES, REVIEWS, ETC.

acquired two chromosomes (in the unreduced nucleus) in the course
of domestication.

EDUCATION OF INFUSORIA IN INGESTION OF FOOD.

Metalnikow (C. R. Soc. Biol., Paris, 1913, pp. 701-704) states
that infusoria may be brought to use more selection in the taking of
substances. By using substances only slightly injurious or even
substances with no nutritive qualities, he found such substances
would be taken indiscriminately at first; but after a period of hours
or days they cease to take them in. Such substances, at first taken
freely and later refused, were aluminium in emulsion, sudan red,
phosphorus, sepia, and carmine. In some instances the presence
of another substance would induce them to swallow particles which
they had learned to refuse. For example, they would take a mix-
ture of sepia and carmine when they refused carmine alone.

SPIROSTYLE IN SPERMATOZOA

Champy (C. R. Soc. Biol., Paris, 1913, pp. 663-4) makes a com-
parative study and an interpretation of the spiral, rod-like body found
in many spermatozoa. He suggests axostyle and spirostyle as its
name. He finds it in several amphibians ; it has also been described in
some reptiles, birds and mammals. He traces the development in am-
phibian from a simple axial rod in the nuclei of the spermatids to
a twisted spiral one in the early stages of sperm formation, and
finally to its partial or total disappearance in mature sperm. Its
twisting in development involves both the nucleus and the cytsplasm,
and thus may give a definite torsion to the whole spermatozoan.
The result in the motion of the sperm is to produce a spiral course
such as we see in many of the protozoa.

NERVE FIBRILS IN DENTINE

Contrary to the usual interpretation, Mummery (Proc. Roy
Soc., Ser. B., 1912, p. 79) holds that the dentine of the teeth is
innervated clear to its outer edge by nerve fibrils from the pulp
cavity. There is a plexus over the outer surface of the pulp, and
from this the neurofibrils, usually two to each tubule, enter the

AMERICAN MICROSCOPICAL SOCIETY 201

dentine tubules and run their whole length to the point where the
enamel or cement joins. This enables us better to understand the
power of the dentist over us.

SUCCESSION IN FUNGI

Brown and Graff (Philip. Jour. Sci., VIII, Sec. C. I; 1913, p.
21) report studies on the succession of fungi growing on dung. This
is a class of studies always of value to directors of laboratories,
and more of such should be made. The authors’ record that the
moulds, as the Mucors, first appeared, followed by Oospora. These
disappeared in about 10 days. Next appeared the sporophores of
species of Coprinus, which persist for a long time. The authors
believe the order of appearance is due to different periods of latency
and rates of development of the spores of the species; and that the
poor persistence of the early types were due to hurtful micro-organ-
isms or to toxins formed in the dung about the fungi. Experiment
showed that the Mucors were not short-lived on sterilized materials.

RUSTS AND THEIR HOST TISSUES

Tischler (Flora 104, 1911; Bot. Gaz., Aug., 1913) describes
the relation between Uromyces Pisi and Euphorbia Cyparissias, its
host in the zcidial stage. The rust winters in the buds of the sub-
terranean shoots, and as these grow it tends to keep pace with them.
If the parasite thrives the host is deformed in a characteristic way.
The author investigated the following among other questions.
Under what conditions do the shoots of the host outgrow, and es-
cape as it were, the ill effects of the parasite? Along what routes do
the hyphae of the rust run in keeping pace with the new growth?
Just at what time in the cell history do the hyphz change the cell
so as to produce the deformed growths?

It was found that the shoots might grow away from the fungus
by furnishing high temperature and other conditions which would
force the growth of the host. Also when the rust approaches its
fruiting stage the host may outgrow it. If kept in the dark so that
zcidia do not form, the buds cannot grow away from the rust.

The hyphz of the rust do not succeed in sending haustoria into
the meristematic cells, and hence the deformation is not due to

202 NOTES, REVIEWS, ETC.

effects at this point, whether at the tip or in the cambium. But as
soon as the cells cease to be meristematic, or embryonic, and begin
to form vacuoles the haustoria are formed and modification of the
cells begins.

The hyphz keep up with the growth of the shoots by following
the trache, from which they penetrate the surrounding tissues.

PHYSIOLOGICAL EFFECTS OF BORDEAUX MIXTURE

It has been claimed that Bordeaux mixture, in addition to its
fungicidal effects, augments the assimilative activity or plants on
which it is sprayed. Ewart (Zeitschr. Pflanzenkrank. XXII, p.
257: Bot. Gaz., June, 1913) finds by experimenting with potatoes,
radishes and beans that the yield was always decreased by cover-
ing the leaves with the mixture, and in proportion to the strength
of the mixture. He also found that the sugar content of currants
was increased by spraying the fruit with the mixture, and decreased
by spraying the leaves alone.

HERMAPHRODITISM IN AMPHIOXUS

Goodrich (Anat. Anz., 1913, p. 318) describes an interesting
abnormality in this animal. A male specimen with 25 testes on one
side had one of the 25 gonads on the other side a perfectly developed
ovary with numbers of large ova. All the 49 testes were perfect
and full of sperm.

RESISTANCE IN HIBERNATING ANIMALS

Bertarelli (Centr. Bakt., 1te Abt. Orig. XVIII, 1913, p. 566)
finds that marmots are not more resistent to rabies, anthrax, tetanus,
and diphtheria during hibernation than at other times. Blanchard
had previously reported these animals to have increased resistance,
during hibernation, to cobra venom, diphtheria, tetanus, trypano-
somes and trichina.

MICROSCOPIC MEASUREMENT BY CAMERA LUCIDA

Joly (Sci. Proc. Roy. Dub. Soc., XIII, 1913, p. 441) suggests
a simple method for measuring microscopic objects by means of the
camera-lucida. Draw two fine lines, diverging from a point, on a

AMERICAN MICROSCOPICAL SOCIETY 203

piece of white paper. The angle of divergence will be determined
by the size of the object to be masured. The image of the object
to be measured is projected on the sheet of paper. The paper is
moved until the object just fills the space between the lines, and a
mark is made across the lines at this point.

A stage micrometer scale is then substituted for the object and
is moved along the diverging lines until a number of the divisions
exactly cover the space between the lines. This point is marked as
before by a cross line. The distance from the intersection of the
lines to each of the cross lines is measured, and one has two similar
triangles from which a single proportion can be derived in which
the size of the object is the one unknown quantity—diameter of
object: micrometer divisions:: distance from intersection to
object: distance from intersection to micrometer.

MICRO-RADIOGRAPHY

Goby (Comp. Rend. CLVI, pp. 686-8: Trans. in J. R. M. S.,
Aug., 1913) reports the application of the X-ray to making visible
the internal structure of opaque microscopic objects. “It replaces
the method of section cutting, which is often slow and costly, and
always indirect and destructive of the object, by a method which,
whilst rapid and preserving the object itself, reveals sufficient detail
to make it only necessary to enlarge the minute radiogram directly
obtained, in order to be able to study it with the naked eye with
the same facility as an ordinary macro-radiogram.”

The difficulty of doing this has arisen in getting the necessary
clearness of detail by means of Roéntgen rays. This is overcome
by an ingenious contrivance which suppresses the secondary or
superfluous rays, and insures that the incident rays shall be normal.
For details of the apparatus the reader must refer to the citations
above. Figures are given which are enlarged (x 19-25) reproduc-
tions of micro-radiograms of Foraminifera and of the limbs of a
small three-toed lizard. The results are remarkable. |

CIRCULATION BY CONVECTION CURRENTS IN LABORATORY AQUARIA

Gemmill (J. R. M. S., June, 1913) describes a simple method
for getting a gentle circulation and zration in single or serial small

204 NOTES, REVIEWS, ETC.

laboratory aquarium jars. It depends in principle upon a water
current some degrees lower than that of the aquaria in the lab-
oratory.

The author recommends tall beakers with about 9 inches of
water in them. The current of colder water is carried through a U
tube of glass, which is connected with the tap and the sink by rub-
ber tubing. The U tube is of %-inch tubing, and dips some 4%
inches into the beaker of water. This leaves an equal distance of
water in the flask below the U tube.

The cool current flowing through the U tube cools the water
in immediate contact with its surface. A downward convection cur-
rent is thus caused in the middle of the jar. The water at the wall
of the jar, exposed to the higher temperature of the laboratory will
supply an upward convection current. Enough of the surface
water is carried downward in the descending current to insure
oxygenation of the whole volume.

For delicate floating larve, such as Asterias, the author shows
that this method is much more satisfactory than streams of air
bubbles. The danger of mechanical injury is eliminated, and it is
possible to isolate the vessels so as to prevent infection even from
the atmosphere. Manifestly a series of U tubes can be used so as
to make the same stream serve a whole battery of vessels.

SIMPLE HISTOLOGICAL METHODS

Salkind (J. R. M. S., Aug., 1913, p. 426) has brought to-
gether some simplifications of histological methods:

1. Sublimate fixation—Instead of removing the salts of mer-
cury by the usual method, the iodine treatment may be carried on
during the removal of the paraffin by placing the mounted paraffin
sections in xylol saturated with iodine. In order to do this, after
fixing in Zenker’s or Helly’s fluid, the objects are placed in a solu-
tion containing 3 per cent potassium bichromate and I or 2 per cent
hydrochloric acid. If acid solutions are not suitable, use the follow-
ing instead: Water, 100 c.c.; corrosive sublimate, 4 grms. ; potassium
bichromate, 2.5 grms.; chloral hydrate, 4 grms.

2. Aceton-Ether Method of Paraffin Embedding—Remove
tissue from water or weak alcohol and place in a fluid composed as

et ie

AMERICAN MICROSCOPICAL SOCIETY 205

follows: Acetone, 2 parts; ether, I part; water, I part. Keep in
this at least one hour for each millimeter of thickness of the tissue.
Transfer to a mixture of equal parts of acetone and ether saturated
with paraffin. Transfer to paraffin.

3. Simultaneous Polychrome Stain—Saturated watery toluid-
in-blue with 3 per cent formol, 12 parts; alcohol, 90 per cent, 8
parts; acetone, 4 parts; saturated naphthol-yellow in go per cent
alcohol, 2 parts; saturated erythrosin pur., in go per cent alcohol, 3
parts. Mix in above order. Add 5 to Io parts of distilled water.
Let stand. No precipitate should appear. The fluid should be a
dark blue, with a violet shade in a few minutes.

4. Adhesions of Sections to Slide—When the paraffin sec-
tions are floating in warm water, add one drop of cedarwood oil.
This spreads as a thin film over the surface of the water. Sections
mounted direct from this fluid will adhere firmly.

REDUCING STOCK SOLUTIONS

Lowe (Zeits. wiss. Mikr., XXIX, p. 545) suggests a simple
method for reducing concentrated stock solutions of reagents to the
dilute form in which they are to be used. Pour into the graduate
a quantity of the stock solution, whose cubic centimeters equal in
number the percentage strength desired in the dilute solution. Add
to this enough of the diluting fluid to make a total number of cubic
centimeters equal to the percentage strength of the original stock so-
lution. If, for example, one wishes to make a 2 per cent solution from
a 15 per cent stock solution, put 2 c.c. of the stock solution into the
graduate and then fill until it totals 15 c.c.

PARASITOLOGY ; LABORATORY GUIDE

This laboratory manual for the study of parasites will be of
great value to zoology teachers who are not themselves experts in
parasitology. The exercises included in the book are based on
courses in the University of California on Human Parasitology and
Veterinary Parasitology, each of one half year.

The introduction deals briefly with the biology of parasitism.
The body of the book is divided into three parts, as follows: L.,
Medical Etomology; II., Helminthology ; — III., Life History
Studies on Living Parasites.

206 NOTES, REVIEWS, ETC.

Part I. opens with a brief discussion of insects and diseases.
Exercises follow, among others, on the mouth parts of insects; the
internal structure of insects; biting lice; sucking lice; bed bugs;
mosquitos ; buffalo gnats ; horse flies ; house flies ; fleas ; ticks ; mites ;
venomous spiders; ameba; trypanosomes; malarial parasites.

Part II. includes exercises on the round worms, hookworms.
lungworms, trichina, filariae; leeches; liver fluke and other trema-
todes, cestodes.

Part III. provides for the study of the life history of the com-
mon house fly, the mosquito, and the flea, and gives suggestions of
general procedure in the investigation.

There are useful exercises on parasiticides and anthelmintics,
and their value. .

In most groups both morphological and systematic studies are
outlined.

The book could have been made more valuable for the gen-
eral zoologist without a great increase in its size, by the addition
of a few devices for identification at least of some of the less com-
monly known parasites, with suggestions for finding, preparing, and
displaying them. The exercises pre-suppose ready-made prepara-
tions, except in the three life-histories in Part III. The exercises
do not make as much use of the suggestive question and the re-
search spirit on the part of the student as the reviewer feels is wise;
but rather follows the method of indicating what is to be found and
expecting the student to verify and identify the findings.

A Laboratory Guide to the Study of Parasitology, by W. B. Herms, The Macmillan
Company, New York. 72 pages. Price 80 cents net.

PREVENTION AND CONTROL OF DISEASE

This book is designed to bring the remarkable work of research
students of recent years, in respect to the prevention and control
of diseases, within reach of the general public. It is felt that such
an increased audience which understands something of the steps
necessary to control disease will advance the work in two ways.
It will make the general public more sympathetic with the investi-
gations that are necessary to get the facts, and more willing to sup-
port the legislators and the health officials who must apply them.

AMERICAN MICROSCOPICAL SOCIETY 207

In style and content the treatise is admirably adapted to the
needs of those for whom it is written—the intelligent general reader
and the undergraduate student in college classes.

An enumeration of some of the principal chapter headings will
enable the reader to appreciate its scope. Chapter I. is an introduc-
tion dealing briefly with some elementary biological matter. Suc-
ceeding chapters are Death-rate and Disease-prevention; Various
Types of Disease and Hygienic Considerations; the Germ Theory
of Disease and other Theories; The Life of Micro-organisms;
Plant and Animal Parasites; Micro-organisms in Air, Water, and
Food; Infection and the Spread of Disease; Disinfection and Dis-
infectants; Susceptibility and Resistance; Immunity; Specifics in
the Treatment of Disease; Colds and Their Like; Filth Diseases,
Typical and Special; Smallpox and Vaccination ; Wound Infections ;
Diphtheria and Pneumonia; Contagious Diseases of Childhood;
Tuberculosis, Its Manifestations and Causes, Its Prevention and
Control, Its Cure; Yellow-fever and Malaria; Cancers; Diseases of
the Second Half of Life.

The general order of discussion in each group of diseases is:
(1) The general nature of the disease; (2) the history and geo-
graphical distribution; (3) cause and manner of infection; (4)
prevention and control.

In our recent agitation against tuberculosis and typhoid we
have been inclined to forget that there are scores of other less
common or less fatal diseases, that are largely preventable, which
diminish human efficiency very greatly. Without removing any of
the necessary emphasis on these more important diseases this book
will give the general reader a less hysterical or unbalanced attitude
toward the whole question of preventable diseases.

The matter of the volume is presented in clear attractive style
which will make it welcome for the purposes for which it was
written.

Prevention and Control of Disease, by Frances Ramaley and Clay E. Giffin. 386

pages. For sale by the University Store, Boulder, Colorado. Price $3.00. Special rates
where employed as a text book.

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TRANSACTIONS

OF THE

American Microscopical
Society

ORGANIZED 1878 INCORPORATED I8QI

PUBLISHED QUARTERLY

BY THE SOCIETY

EDITED BY THE SECRETARY

VOLUME XXXII

NuMBER Four

Entered as Second-class Matter December 12, 1910, at the Post-office at Decatur, IIli-
nois, under act of March 3, 1879.

Decatur, ILL.
Review Printinc & STATIONERY Co.
1913

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OFFICERS.

President: F.CreicgHToN WELLMAN, M. D................ New Orleans, La.
Furst Vice President: F.C. Watre, Ph.D... 2.0.5. c cece Cleveland, Ohio
Second Vice President: H.E. Jorpan, Ph.D............. Charlottesville, Va.
SeEeelary Wet GAILOWAY cle oe inc ad's 6 mgisinjs-oim'e a bins no's ine bien ote Decatur, Ill.
Tie CUSUER 12 lists EUANIKENSON & 6 255 5:c.cve cisiviolote otis ster oe baal eroratane Charleston, Ill.
OS TOMLA sd WNLAGN US RTA TI Mo fetes ev saier ol ceto lec taleveta'e sn iota > 2/01 obese Meadville, Pa.

ELECTIVE MEMBERS OF THE EXECUTIVE COMMITTEE

Dea NG eh SVAINIEZ: 5 6.5; 55 evsveyateratay ibis? avalos Bureau Plant Industry, Washington, D. C.
SEINE, J.) cnt. Rie aie aa stil potaclstaiey sae a cd bala eeaiaisrs da'bla es Laramie, Wyo.
Se DM COGEDIETS rete Ae ee ee ae cin eee Raatoete ese oeeee Lawrence, Kans.

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
Apert McCatra, Ph.D., of Chicago, Ll.
at Chicago, Ill., 1883
T. J. Burritt, Ph.D., of Urbana, Ill,
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.
Stmon Henry Gace, B.S., of Ithaca, N. Y.,
at Ithaca, N. Y., 1895 and 1906
A. CiiFForp Mercer, M.D., F.R.M.S., of Syracuse, N. Y.,
at Pittsburg, Pa., 1896.
A. M. Bueite, M.D., of Columbus, Ohio,
at New York City, 1900.
C. H. E1iceENMANN, Ph.D., of Bloomington, Ind.,
at Denver, Colo., 1901.
Cuartes 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 Urbana, Ill,
at Sandusky, Ohio, 1905.
Herpert Osporn, M.S., of Columbus, Ohio,
at Minneapolis, Minn., 1910.
A. E. Herrzier, M.D., of Kansas City, Mo.,
at Washington, D. C., Io1T.
F. D. Heatp, Ph.D., of Philadelphia, Pa.,
at Cleveland, Ohio, 1912.

The Society does not hold itself responsible for the opinions expressed
by members in its published Transactions unless endorsed by special vote.

TABLE OF CONTENTS

FOR VOLUME XXXII, Number 4, October, 1913

Report of the Secretary and Editor, by T. W. Galloway................. 213
The Powdery Mildews,—Erysiphacex, with Plates XIII-XVI, by George
Md Reeders once Ser oe rere eee eae eenteeoske iets aie ae ie ee 219
A Descriptive List of Cephaline Gregarines of the New World, with
Plates: 2CV.LIE eX by Max. Mes BINS oon shere cisiotlemicine ae eee eee 259°
Notes and Reviews—A Paraffin Ribbon Carrier, by R. T. Hance, with
Plates XXI-XXII; Insects and Disease; Adaptation in the Gall
Midges; Biology of the May-flies; Hibernation of the House-fly;

Parasite of the Chinch Bug; Grasshoppers..............ceeeeceeees 297
BCASE DOE WeHIDETG. (So sos eihts arse iel icine ater ee bee DRM ne chee Cee oh eee 303
EASPaDE | SUDSCEIDCES - pie as «+ Oe oe ee nee on eeiakoas dues caves noe eee 310
UR GER yen se Secs aie SORE Dea Hoe OI ee in Ee Cla eee 313

NOTICE TO MEMBERS

Members are hereby notified that the regular annual meeting
of the American Microscopical Society will occur in Atlanta,
Georgia, at 3:30 p. m., on Wednesday, December 31st, in connec-
tion with the American Association for the Advancement of Science.
The place of meeting will be announced in the official program of
the A. A. A. S. The meeting of the Executive Committee will be
held at Georgian Terrace at 12:30 noon on December 30th, at which
time the committee members who are present will lunch together.

TRANSACTIONS

OF

American Microscopical Society

(Published in Quarterly Installments)

Vol. XXXII OCTOBER, 1913 No. 4

REPORT OF THE SECRETARY AND EDITOR
T. W. GALLOWAY

The issuance of this number closes the period for which the
present Secretary was elected. It has not been the custom for secre-
taries of the American Microscopical Society to issue an address
to its members thru the pages of the Transactions; nor is it the
usage for editors of scientific magazines to do so. There are cer-
tain reasons however why it appears to the writer appropriate for
him, in the combined capacity of Secretary and Editor, to bring be-
fore the Society in this form a statement of the work done during
this triennium, and to indicate what seem to be the deductions fairly
to be made as the result of the developments of the period.

This has been a time of transition for the Society. Histo-
tically, there was a long pericd in the early history of the Society
when there was a successful annual meeting supported by a limited
number of enthusiastic microscopists from the country at large and
by a considerable number of actual microscopists and well-wishers
worked up at or near the place of meeting.

With the inevitable extension of the use of the microscope in
all institutions of learning and the specialization of the people
using it, there came a period in which the annual meetings dwindled,
and could scarcely be called successful from the point of view of
even the most optimistic friend of the Society. Even where the
attendance was fair, the cleavage of interest between the amateur
worker and the professional work became so pronounced as to make
the scientific program something of a trial to both.

When the present Secretary assumed the work it soon became
apparent that a choice of emphasis must be made. On account of
the great distances in our country, because of the diversity in the

214 T. W. GALLOWAY

specialization of our membership, and because of the great develop-
ment of the special scientific societies for the promotion of particu-
lar realms of work it seemed wise frankly to give up hope of having
successful programs of the American Microscopical Society, and to
confine the annual meeting to such business routine as is absolutely
essential to preserve the integrity of the corporation.

The necessary corollary of this elimination of the scientific
meeting, which was formerly one of the chief forms of service ren-
dered by the Society to these members who could attend, was to
bring to all members such timely papers, on subjects suited to the
general membership of the Society, as the income would allow. All
the energy and funds put into holding a meeting were to be put into
the publications. |

To effect this it was decided to change the form of the Trans-
actions from an annual to a quarterly issue. While the expense of
such issue is greater, it was felt that the timeliness would more than
compensate for this. It was felt furthermore that a general journal
devoted to somewhat broad aspects of microscopic biology, pub-
lished in the middle west would, for a period at least, serve a real
end in science, and might increase the appeal of the Society to the
new generation of investigators and teachers. It would furnish a
new avenue of periodical publication, and might be made to meet
the needs of such groups of biological workers as are least well sup-
plied.

It was urged by some friends of the Society that the Quarterly
Transactions undertake to fill some very special field,—as cytology
—which is not at the present time occupied by any American publi-
cation. Such a course presents many elements of attractiveness.
Partly because such narrow specialization would be a complete
break with all the traditions of the Society and partly because the
Secretary did not feel competent to pilot such a venture, it seemed
wiser, however, merely to find a fairly permanent clientage in need
of certain things and to undertake to supply this need.

With this task in mind and because of his own experience as a
teacher of biology in small colleges, the Secretary was convinced
that there would be the least departure demanded from the tradi-
tion of the Society to undertake to supply certain features that

REPORT OF THE SECRETARY 215

would be of practical value to the teacher or student with inade-
quate journal facilities. Such teachers usually find it necessary to
be reasonably familiar with a rather wide range of special interests,
and yet do not have access to the publications necessary to secure
this familiarity.

In pursuance of this idea it appeared that there are at least
three types of contribution that would be of conspicuous value to
general students. These are:

First—Notes, reviews, abstracts of such special articles as are
most valuable from the point of view of the teacher. The purpose
is not at all to make these abstracts cover the whole field of biology,
but rather to indicate some main lines of progress and to furnish
the teacher with illustrations of the progress. Arrangements are
now being made to get the help of several biologists to make ab-
stracts of the most suggestive work done in their respective fields.

Second—Occasional digests by experts of recent progress in
some restricted departments. It was contemplated that these would
represent the work of specialists who would bring to the general
reader the discussion of the main conclusions and tendencies and
prospects within such fields. Several such have appeared. Others
are in preparation.

Third—It was felt that synopses of some of the more important
groups of microscopic and near-microscopic plants and animals,
would be of great value to teachers. The thought again is not sci-
entific completeness ; but such a treatment of the species more likely
to be met by a teacher or his students, together with keys, figures,
and descriptions, as would enable identification by the student. It is
a fact that we are beginning to recognize that our schools and col-
leges are neglecting to teach the students to identify and name at
least a few of the forms they meet in their work. It is important
that every biologist, no matter how narrowly he is specializing,
should have some mastery over the classification of the more com-
mon animals and plants. From the point of view of the general
teacher of biology there is most important pedagogical value in a
certain amount of systematic work. These simple descriptions,
keys, and figures of the more common species of microscopic or-
ganisms are designed to aid the general student and teacher, rather

216 T. W. GALLOWAY

than the specialist in the department. Quite a number of these have
already appeared, as follows: Aquatic Oligochetes, Melanconiales,
Rusts, Black Moulds, Powdery Mildews, and Cephaline Gregarines.

Others are arranged for and will appear from time to time un-
til all the more common American forms have been touched upon.
The Editor desires to express his appreciation of the help of the
scientists who are consenting to do this task. Their prompt and
cordial response convinces him that they believe the enterprise
worthy.

It has been the purpose to maintain the space given to re-
search articles over as wide a field of microscopic biology as condi-
tions will allow. The flow of manuscripts has not been sufficient to
require or enable the editor to have any plan other than to produce
an issue of a reasonable size. This has meant that thus far those
writers who have availed themselves of our pages have had very
prompt publication. As the permanency of the Quarterly becomes
manifest this supply will doubtless increase, with a corresponding
delay in publication.

By vote of the Executive Committee the Spencer-Tolles Fund
now becomes operative in encouraging and aiding investigators. The
details of this arrangement may be had by application to Dr. H.
B. Ward, Urbana, IIl., who is Chairman of the Committee having the
awarding of this grant in hand. Results of investigation conducted
under this grant will be published in the Transactions and will add
to the value of the journal to members.

During these three years we may fairly say that the Society
has prospered in respect to finances‘and in membership. In the
latest list available to the Secretary when he took up the work,
there were 226 names of members, and 33 of subscribers. There
have been added in these three years 188 members and 36 subscrib-
ers. This old list with which we started had not been revised for
some years, and thru death and resignations the original list has
lost 115 names. Some of these names have been carried for these
three years on the rolls with the hope that the members might re-
new their standing. Some have done so. This year about 65 of
the old members’ names are finally dropped from the list. During
this year 45 members and 25 subscribers have been added. As this

REPORT OF THE SECRETARY 217

is written we are printing a list of 293 members and of 68 sub-
scribers—a total of 361. This list is almost absolutely a net list of
members who are promptly keeping alive their membership. One
more year of growth such as we have been having will place the
membership at 400. This number will give the Society complete self
support and enable the issue each quarter of a magazine of 80 to
100 pages of scientific material.

The Secretary would also call the attention of the Society to
the gradual change in the character of the membership. We have
lost the memberships, formerly numerous, of amateurs and well
wishers brought into the Society thru the special efforts of the mem-
bers living in and near a place of meeting. Most such members
were transient ; but among them were many amateur microscopists
who meant very much to the life of the Society. This type of mem-
ber we have not found a way to enlist as formerly. Our great gain
has been among teachers. These have come in, one by one, on the
recommendation of other teachers. Without doubt they will give us
a more stable membership than was true formerly. While we have
many physicians as members we have not been able to get hold of
physicians thru our physician members as we have in the case of the
teachers. This is doubtless due in part to the excellent character of
some of the publications now available to physicians in the realm of
medico-biological research.

The Secretary feels that the Society owes something still to
amateur microscopists in America. Without forgetting any of the
difficulties in the way, it would seem possible to conduct a depart-
ment in which the matters most needed by isolated and incompletely
trained students might be developed and dwelt upon. The various
methods, devices, short cuts, and so forth, in connection with the dis-
covery, collection, culture, preservation, mounting, and study of
microscopic materials might well be brought out. Possibly a brief
department of questions and answers might be established for ama-
teur workers. The Secretary has been seeking some one to edit
such a department, but as yet without success.

In conclusion, the writer wishes to renew a suggestion made

last year in connection with the giving up of the annual scientific
program. li successful meetings of general microscopic workers

218 T. W. GALLOWAY

are to be held, it must be under the auspices of societies of limited
area and not by a national body. “It may be pertinent to suggest
that, in the opinion of the Secretary, the Microscopical Societies of
more local character should be able to carry on successful meetings.
These may be state or city societies. There is no reason why really
live, valuable meetings for practical discussions and demonstrations
cannot be had under these circumstances. The number of people
who use the microscope and are interested in its application is
greatly increased. There are more indeed in a single city than were
found in the whole nation when this society was organized. The
A. M. S. stands ready to serve such local societies in any way pos-
sible. Indeed it seems as tho an effective division of labor would
be: (1) the national society to furnish a magazine of microscopy
and microscopic research; and (2) state and city societies to fur-
nish the personal contacts and stimulus thru meetings. Some such
broad affiliation of national and local societies ought to be possible
and mutually supportive.”

SUMMARIES IN MICRO-BIOLOGY

For some months the Secretary has been seeking to secure fer this Journal and its
Department of Summaries, a series of papers from biologists dealing with the chief groups
of microscopic plants and animals. It has not been the purpose to present a complete
survey of any of the groups. The wish has been rather to bring together in one article
a statement of the following things:—-general biology, the method of finding, the methods
of capture and of keeping alive and cultivating in the laboratory; how best to study; the
general technic; the most accessible literature; and a brief outline of the classification,
with keys for the identification of at least the more representative genera and species of
the micro-organisms likely to be found by the beginning students in the United States.

It has been felt that the getting together of such data as this, while not a contribution
to science, would be a contribution especially to isolated workers and to teachers and stu-
dents in the high schools and smaller colleges.

Papers have already appeared treating the aquatic Oligocketes, the Melanconiales,
the Rusts, and Black Moulds. The following is the fifth paper of the series. It is pro-
posed to have such synopses from time to time until the more common American species of
such groups as the following have been covered: The Blue-green Algae, Conjugating
Algae, Diatoms, other Green Algae, Downy Mildews, Yeasts, Hyphomycetes, Smuts,
Rhizopods, Infusoria, Turbellaria, Bryozoa, Water Mites, Entomostraca, etc.—[Editor.]

THE POWDERY MILDEWS—ERYSIPHACE/E
By GeorcE M. REED

Introduction. The Erysiphacez constitute a well defined
group of the Ascomycetes. Various common names are applied to

these fungi, as mildew, powdery mildew, white mildew, blight. They
have long been interesting forms for study, partly on account of the
microscopical interest in observing the fruiting bodies with their
characteristic and frequently ornate appendages, but more especially
on account of their parasitic nature, in many cases being of consider-
able economic importance due to the injury caused to various seed
plants.

The powdery mildews are obligate parasites, attacking various
Angiosperms. In fact these parasites are mainly limited to the
Dicotyledons, only one species being reported as common upon
Monocotyledons—Erysiphe graminis on members of the grass family.

The group is characterized by the possession of two distinct
fruiting stages. One, the conidial stage, develops during the sum-
mer and results in the formation of a very large number of white,

220 GEORGE M. REED

more or less ellipsoidal spores which are easily distributed by air
currents. This stage has been referred to a distinct genus, Oidium,
belonging to the Fungi Imperfecti. It is now described as the Oidial,
or better still, the conidial stage. Later in the season the ascus
stage of the fungus is produced. The asci, one or more, are borne
within the fruiting bodies or ascocarps. The latter are small, more
or less globular, dark brown or black structures without different-
iated openings for the escape of the asci or ascospores. These
ascocarps, or perithecia, are provided with peculiar appendages which
are outgrowths of the outer layer of cells of the perithecial wall.
They are sometimes ornately branched and their characteristics are
of considerable value in classification. The asci are sac-like struc-
tures, more or less oval in shape, and at maturity contain two to
eight ascospores depending upon the species. In general the conidial
stage serves to spread the fungus rapidly and widely during the
growing period of the host while the ascocarp stage serves to tide
the fungus over the winter season.

Mycelium and Conidial Stage. With few exceptions the
mycelium of the powdery mildews is external to the host tissues.
It generally develops on either the upper or lower surfaces of the
leaves, but frequently is found on the stems, flowers and fruits. The
mycelium is generally hyaline, well developed and much branched
and the hyphae which compose it are septate. Thus it is composed
of many cells, each cell containing regularly one nucleus.

At numerous places on the mycelium a special hypha grows
towards the host tissue, penetrates a cell of the host, and forms a
haustorium. In many cases, these haustoria arise from a flattened
expansion, called an appressorium, of the superficial mycelium. The
haustoria function as absorbing organs and obtain the necessary ma-
terials for the mycelium from the host protoplast.

Generally these haustoria are confined to the epidermal cells.
In a few cases they are formed in the deeper lying cells. For ex-
ample, in Uncinula salicis, according to Smith (62), some hyphe
from the surface mycelium grow through the epidermal cells and
form the haustorial expansions in the cells of the mesophyll tissue.
(Pl. XIII, Fig. 2). Even in this mildew, however, most of the
haustoria develop in the epidermal cells. In many cases, notably the

THE POWDERY MILDEWS 221

' grass mildew, several of these absorbing organs may be found in a
single host cell.

Smith (62) has quite fully described the haustoria of the
Erysiphacez and their relation to the host cells. In most cases they
are more or less globular swellings at the ends of the special pene-
trating branches (Pl. XIII, Figs. 1, 2). In the grass mildew, Erysiphe
graminis, the haustoria seem to be highly specialized structures (PI.
XIII, Fig. 3). They consist of an ellipsoidal central portion with long
finger-like processes at one or both ends. They would appear to be
efficient absorbing organs and may be correlated with the vigorous
development of mycelium and abundant production of conidia char-
acteristic of this mildew. The haustoria of E. galeopsidis are also
somewhat lobed. In fact the lobing of the haustoria of this species
is the principal basis for separating it from E. cichoracearum.

The haustoria of all the mildews regularly contain a single
nucleus; very rarely are two nuclei found in one haustorium.

As already indicated the mycelium of some mildews, in part at
least, is endophytic. The mycelium of the genus Phyllactinia grows
in the intercellular spaces of the mesophyll of the host leaves. The
haustoria develop as side branches from the intercellular mycelium
and penetrate the various host cells in the interior of the leaf.

Salmon (59) has described Erysiphe taurica as being even more
endophytic in its habit than Phyllactinia. The mycelium is at first
wholly endophytic ; from this arise conidiophores which pass through
the stomata to the exterior. The perithecia are developed on a
superficial mycelium which originates from that in the interior of
the leaf. Globular haustoria are found in the mesophyll cells of the
host. Salmon has recently separated this species, on the basis of
these characters, and placed it in a distinct genus—Oidiopsis.

As the mycelium develops special branches arise which grow
away from the surface of the leaf. These hyphae form the conidio-
phores (Pl. XIII, Fig. 4), one of the fruiting stages of the fungus.
At the end of each conidiophore a more or less ellipsoidal conidium is
cut off. The hypha elongates again and a second conidium is cut
off. This process of basipetal abstriction continues, finally re-
sulting in the formation of a chain-like row of oval conidia, the
oldest being at the apical end. These easily separate as they mature

222 GEORGE M. REED

and are readily distributed by air currents. The young cell which
forms the conidiophore contains a single nucleus. Nuclear division
precedes the cell division and thus each conidium comes to have a
single nucleus.

The conidia germinate readily in water or moist air by pushing
out one or more germ tubes. (Pl. XIII, Fig. 5). Unless applied to the
proper host, however, but little growth occurs. If the conidia are
placed on the proper host, mycelia with their haustoria are quickly
developed which soon give rise to other crops of conidia. In the
grass mildew 48-72 hours are sufficient for this cycle of development.
Consequently during a single season a succession of crops of conidia
may be formed and many millions of spores may be produced.

Development of the Perithecium. Although many students
of the mildews had examined carefully the structure of the mature
perithecium and had worked out systems of classification based upon
its characteristics, De Bary (5) in 1863, was the first to adequately
trace the origin of this structure. He studied the development of
the perithecium of Sphacrotheca castagnei on Taraxacum. De Bary
(6) later (1871) studied other forms and found their development
essentially similar to that of Sphaerotheca.

De Bary describes the ascocarp or perithecium as arising at the
point of contact er crossing of two branches of the mycelium. These
branches push out protuberances at the same time which rise erect
from the epidermis of the host. They are soon cut off by cross
walls and the one from the lower hypha grows and takes the form
of an elongated ellipsoidal cell. This is the archicarp or oogonium.
The other branch, the antheridial branch, remains cylindrical and
is closely applied to the oogonium, its upper end bending over and
covering the apex of the latter. A cross wall cuts off a short nearly
isodiametric cell, the antheridium, which is borne on the basal cell
or stalk. The oogonium now develops into the sporocarp, usually
being divided into two cells, the upper one forming the solitary ascus
of this mildew while the other remains as a stalk cell to the ascus.
The ascus subsequently produces eight ascospores. As the young
ascus develops the envelope apparatus is formed by the outgrowth of
seven to nine tubular hyphz from the base of the oogonium and
antheridium. These hyphae elongate, remaining in close contact

THE POWDERY MILDEWS 223

with each other and also with the oogonium and antheridium, finally
meeting above the apex of the antheridium. Each hypha then divides
by one or two transverse walls and the young sporocarp is sur-
rounded by a single layer of cells. From the inner surfaces of these
hyphae secondary branches arise which ramify and develop into a
dense parenchyma-like weft formed of two or more layers of cells.
From the outer layer of cells, those first formed, the appendages
arise. The walls of these outer cells also thicken and assume a dark
brown color.

In Erysiphe the main difference lies in the fact that the archicarp
or oogonium grows into a curved tube and divides by transverse
walls into a row of several cells. From these cells a number of club-
shaped, erect asci are formed by each cell growing out into an ascus
or putting out a few short branches which finally terminate in asci.

Although De Bary maintained that the perithecium originated as
the result of a fusion of sex cells he was unable to determine the
actual fusion of the protoplasts of the oogonium and antheridium.
Harper (28, 29, 30), using modern cytological methods, has been
able to verify De Bary’s conclusion and has added many additional
facts to the history of the development of the perithecium and the
following account is based on his studies of Sphaerotheca, Erysiphe
and Phyllactinia.

Both antheridium and oogonium arise as side branches of neigh-
boring hyphae. The development of the oogonium generally pre-
ceeds that of the antheridium and it soon forms a short oval branch
which can easily be distinguished from vegetative branches by be-
ing vertical to the leaf surface and also by containing denser pro-
toplasm. After the oogonial branch has elongated until it is two to
three times as high as wide, with a transverse diameter twice that
of a vegetative hypha, it is separated from the latter by a cross wall.
(Pl. XIV, Fig. 9). The oogonium contains a single nucleus which is
hardly distinguishable from that of a vegetative cell. During this
process the young antheridial branch bends up and grows close to
the side of the young oogonium. After a time a cross wall is formed
cutting off a single nucleated cell. After the complete formation of
the oogonium the antheridial branch elongates and its nucleus di-
vides (Pl. XIV, Fig. 10), followed by cell division which cuts off a

224 GEORGE M. REED

small terminal antheridium and a stalk cell (Pl. XIV, Figs. 11, 12).
The antheridium is carried upward by growth, becomes closely ap-
pressed to the oogonium, and appears as a cap on the latter. Next
the cell wall between the oogonium and antheridium is dissolved and
the antheridial nucleus migrates through the opening and approaches
the egg necleus which lies near the center of the oogonium (PI. XIV,
Fig. 13). The nuclei now fuse and soon the opening between the
antheridium and oogonium is closed (Pl. XIV, Fig. 14).

At the same time with the entrance of the male nucleus into the
oogonium the development of the perithecial wall begins (Pl. XIV,
Figs. 15-20). Hyphe arise at the base of the oogonium and grow up
around the antheridium and oogonium. These become multi-cellular
by nuclear and cell division. The antheridium relaxes and collapses
but persists among the wall cells for some time. Next, due to the
enlargement of the stalk of the oogonium, the first series of wall
cells are bent out and other hyphae grow up inside them and, by
branching and dividing, form a sphere of cells about the oogonium.
The wall of the perithecium then contains several layers of cells,
the outer for protection, the inner for nourishment. From the
former the appendages arise when the perithecium is about half
grown (PI. XIII, Fig. 8). These soon become thick walled and lose
their protoplasmic contents on the further ripening of the perithe-

cium.
The fusion of the two nuclei in the oogonium takes place before

the completion of the first wall layer. As further development pro-
ceeds, the oogonium grows into the ascogonium. The fusion nucleus
in the oogonium first divides, following by cell division (Pl. XIV,
Fig. 17) ; the lower cell of these two develops no further. The nucleus
of the upper cell again divides and this is followed by cell division ;
this process is repeated until a series of five to six cells is formed
(Pl. XIV, Fig. 19). Each of these cells has regularly one nucleus ex-
cept the next to the last which invariably has two. This penultimate
cell develops into the ascus (Pl. XIV, Fig. 20).

In the early stages the development of the mildews with several
asci is similar to that of Sphaerotheca. Harper (29, 30) has care-
fully traced the development of sex organs and established the nu-
clear fusion in the young oogonium of both Erysiphe and Phyllac-
tinia.

THE POWDERY MILDEWS 225

The oogonium, after the fusion of the antheridial and oogonial
nuclei, develops into the ascogonium (PI. XIII, Fig. 6). This is ac-
complished by the elongation of the oogonium which becomes curved
in a very irregular fashion. Nuclear division occurs but this is not
followed at once by cell division. Instead further nuclear divisons
occur. Soon, however, cell division takes place and there is formed
a row of three to five cells. The end cell of the fully developed
ascogonium regularly contains one nucleus while the next to the
last or penultimate cell always contains more than one nucleus.

Following the formation of the ascogonium, the ascogenous
hyphae arise as lateral branches of the former (Pl. XIII, Figs. 6, 7).
Most, if not all, of these hyphae arise from the penultimate cell. The
number of cells in each hypha varies but one cell in each becomes
an ascus. These cells always contain two nuclei while the other
cells of ascogonium and ascogenous hyphae are almost without ex-
ception uninucleated.

During these processes the perithecial wall is formed by the up-
growth of hyphae from the stalks of both antheridium and oogon-
ium. These, by continued division and branching, form finally the
many celled wall of the perithecium (Pl. XIII, Fig. 8). When the
perithecial wall is fully differentiated the further development of the
asci begins.

As already noted each young ascus always contains two nuclei
(Pl. XIV, Figs. 19, 21). Harper (30) has traced with exceptional
completeness the nuclear phenomena which occur in the further de-
velopment of the asci (Pl. XIV, Figs. 21-24, Pl. XV, Figs. 25-33).
The asci rapidly increase in size and consist of an upper enlarged
portion, in which the nuclei lie, and a lower stalk-like portion.

When the ascus has attained about half of its mature size the
two nuclei fuse to form the primary ascus nucleus. (Pl. XIV, Figs.
22-24). The union of the two resting nuclei results in the formation
of a single spherical nucleus which increases in size with the further
growth of the ascus (Pl. XV, Fig. 25). This primary ascus nucleus
next undergoes division resulting in the formation of two nuclei;
each of these divide giving rise to four which in turn divide forming
eight nuclei in the ascus (Pl. XV, Figs. 26-30). This triple divi-
sion of the primary ascus nucleus is characteristic not only of the

226 GEORGE M. REED

mildews but of by far the larger number of other Ascomycetes as
well.

During all the nuclear processes from the first development of
the young oogonium Harper has been able to demonstrate the occur-
rence of a central body and to follow its behavior. This central
body is located on the nuclear membrane. “It constitutes through-
out a point of attachment for the elements of the nucleus, and in all
the various modifications which it and they undergo in the processes
of division and fusion this relation is maintained in the most
definite fashion. The central body by its position determines in
an important sense a definite polar organization on the part of
the chromatin, and thus of the nucleus as a whole.” “In every
stage the chromatin is definitely attached to either one or two
central bodies on the periphery of the nucleus. The nucleus is hence
strictly unipolar throughout its so-called resting stages, becoming
bipolar by division of the center for the formation of the two daugh-
ter nuclei.” Harper has given a continuous account “of the exist-
ence of the central body and the maintainance of its connection with
the material of the chromosomes through two nuclear fusions in the
oogonium and in the young ascus, through a series of divisions in
the ascogenous hyphae, and the triple division in the ascus, and
finally through the formation of the ascospores by free cell forma-
tion.” “The central bodies are thus seen to be permanent structures
of the cell during both the dividing and resting stages of nuclear
development.”

During the process of all three nuclear divisions there is a well
developed system of astral rays which extend from the central body
into the cytoplasm. This polar aster persists after the third division
and functions in the process of spore formation.

In Phyllactinia regularly only two ascospores are formed, each
with a single nucleus. The other six nuclei formed in the triple di-
vision of the primary ascus nucleus are regarded as supernumerary
nuclei which soon disintegrate in the cytoplasm. In some mildews,
however, as in Sphaerotheca, all eight nuclei function and the ma-
ture ascus contains eight ascospores. In the different mildews the
number of ascospores varies from two to eight.

THE POWDERY MILDEWS 227

Harper has described ascospore formation as a process of “free
cell formation.” Following the completion of the third nuclear di-
vision, a beak is developed on the functional nuclei which are usually
more or less pear-shaped.

The central body is located at the apex of the beak. In general
the beaked nucleus and aster lie free in the cytoplasm but sometimes
lie quite close to the ascus wall. The astral rays are folded over to
form the plasma membrane of the spore. (Pl. XV, Figs. 31, 32).
“The rays become elongated during the process by growth which
apparently proceeds from the central body outward, and at the same
time they fold over and combine side by side to form a continuous
broad, umbrella-shaped membrane. Sometimes the rays on one side
seem to be in advance of those on the other in the process of enclos-
ing the spore mass. If, in folding over and elongating, the rays of
one center come in contact with those of another, they tend to fuse,
at least temporarily. Later, however, they must separate again,
since one almost never finds spores with two nuclei.” “The broad
umbrella-shaped membrane gradually closes in to form, by further
marginal growth, the ellipsoidal plasma membrane of the spore.
The whole spore body is cut out of the previously undifferentiated
cytoplasm of the ascus by the formation of a new plasma membrane
derived from the fibers of the polar aster and without the deposition
of a cellulose wall.”

Following the enclosure of the spore plasm by the new plasma
membrane, the central body breaks away from the membrane, the
nucleus regains its spherical or oval shape, with the central body
lying on the surface of the nuclear membrane. Finally a wall is
formed about the spores and the development of the perithecium
with its ascospores is complete (PI. XV, Fig. 33) and there follows
the resting condition.

In the formation of the ascospores only a part of the cytoplasm
is used up. The delimitation of the spores leaves a considerable
portion of unused material. The spores thus are imbedded in the re-
maining part of the ascus cytoplasm. This material is called
epiplasm or periplasm and its presence is a constant feature in the
ascus of the Ascomycetes.

228 GEORGE M. REED

In at least two species of Erysiphe—E. graminis and E. gale-
opsidis—spore formation does not take place on the living host.
When mature perithecia of these forms are found an examination
will show the absence of the ascospores. Instead these are formed
the following spring. To just what stage the nuclear phenomena
proceed in the fall in these forms has not been worked out.

The ascospores escape from the perithecium in the spring.
Galloway (25) states that the perithecium may suddenly burst and
forcibly eject the asci. The cells of the inner wall of the perithe-
cium, which retain their protoplasmic contents, may produce a sub-
stance capable of swelling in water and so cause the rupture of the
perithecium as suggested by Harper.

The ascospores, when placed in a damp atmosphere or in water,
germinate by the formation of germ tubes. If the ascospores are
sown on the epidermis of a suitable host the germ tube penetrates
and forms a haustorium in the host cell. The superficial mycelium
also develops from this tube and, by growth and branching, soon
spreads over a considerable area of host surface, giving use to
numerous conidiophores.

On some hosts of the mildews perithecia are rarely formed.
One noted illustration of this is the mildew on the cultivated grape
in Europe. Berkeley (7) in 1847 described the conidial stage of this
fungus as Oidium Tuckeri and, although this mildew was recognized
as a serious disease, the perithecia were not observed until 1893
when Coudere (15) found these fruiting bodies and determined
finally the identity of this mildew with the American Uncinula ne-
cator. The perithecia are quite common _on grapes in this country.

In 1907, 1908 and 1909 a serious disease of various oaks in
Europe, caused by a mildew, was observed by a large number of
workers. The conidial stage was very common and quite destruct-
ive to the foliage of young oaks. While perithecia have since been
found, they are of rare occurrence on these hosts in Europe.

It is well known that the conidial stage of Erysiphe graminis
is very common on blue grass and other grasses but perithecia are
quite infrequent. In the vicinity of Columbia Evonymus atropur-
pureus is severely attacked by a mildew but, while a large number
of plants have been carefully examined by the writer, the perithecia

THE POWDERY MILDEWS 229

have not been found. Various species of Xanthium are also at-
tacked by the conidial stage but perithecia are rarely produced.

As already noted characteristic appendages are found on the
mature perithecia of the powdery mildews. These are outgrowths
of the outer layer of cells of the perithecial wall. They develop
from different parts of the wall in different species and show char-
acteristic differences in their form. Their function is uncertain. In
many cases, perhaps all, they serve to set the perithecium free from
the surface of the leaf. In Phyllactinia for example, Neger (40)
has observed the bending down of the appendages and their straight-
ening up as a result of the alternately moistening and drying of the
perithecia. In fact, in the case of the Phyllactinia, it is quite usual to
find the perithecium in the mature condition turned over on the
surface of the leaf so that they rarely are found in the normal po-
sition.

In addition to the characteristic appendages which are found in
all the powdery mildews, certain penicillate cells develop upon the
apical portions of the perithecium of Phyllactinia. These arise as
an outgrowth of the cells of the outer wall of the perithecium, then
grow up more or less vertically, branching repeatedly. After they
have attained their full size the wails begin to swell and become
gelatinous. They fuse together laterally and form a slimy mass
crowning the perithecium. They serve to attach the perithecia in
an inverted position to various objects, more commonly to the living
epidermis of the host after the perithecia have been loosened from
their place of development by the hygroscopic movements of the
appendages. The perithecia may fall upon Fomes, as has been re-
corded, and be re-attached by these penicillate cells. Perhaps the
unusual host distribution of this species may be, in part at least, ex-
plained by the accidental re-attachment of the perithecia.

There are several questions of great importance raised in con-
nection with the facts of cytological study of the mildews. One
question is that of the significance of the double nuclear fusion
which occurs in the life history of these plants. One nuclear fusion
occurs in the oogonium, when the nucleus from the antheridium
migrates through the opening in the cell wall and fuses with the

230 GEORGE M. REED

egg nucleus. The second fusion occurs in the young ascus, this
nuclear fusion resulting in the formation of the primary ascus nu-
cleus.

Some investigators of this group deny that there is any fusion
of nuclei in the oogonium and that the nuclear fusion which occurs
in the ascus is the real fertilization process in these forms. The
ascus, by these workers, is interpreted as an egg. Harper, however,
has given very convincing evidence of the sexual nature of the
hyphe first described by De Bary as oogonium and antheridium and
leaves no doubt as to the nuclear fusion in the oogonium. His work
has also been confirmed by Blackman and Fraser (8) although de-
nied by Dangeard (16) and more recently by Winge (72). If the
nuclear fusion in the oogonium is a sexual process, as seems certain,
then the subsequent nuclear fusion in the ascus cannot be considered
a sexual process and the ascus cannot be interpreted as an egg.

Harper (30) has attempted to explain this second fusion on the
basis of the nucleo-cytoplasmic relation. He believes that this
fusion is “correlated in some way with the vegetative development
of the relatively gigantic size of the ascus as compared with other
cells of the fungus.” Large cells in general have large or numerous
nuclei and small cells have small or few nuclei. Nuclear and cyto-
plasmic masses are in equilibrium when there is a certain proportion
between them. “Any increase in the mass of either tends toward
producing a corresponding increase in the other; a reduction in one
necessitates a reduction in the other, in order that the nucleo-cyto-
plasmic equilibrium may be maintained.”

“The ascus is to be developed as a relatively large cell to serve
as a storehouse, with an abundant supply of material for the forma-
tion of ascospores; and in order that the nucleo-cytoplasmic equili-
brium may be maintained, it must be provided with an excess of
nuclear material as compared with the other cells of the ascogenous
hyphae and the ascogonium. There are several stages in this differ-
entiation of the ascus as to its nuclear content. It is binucleated
from the first, while the other cells mentioned are uninucleated ; and,
further, its two nuclei fuse with the union of all their corresponding
parts to form a single larger nucleus, which in turn grows with the
further growth of the ascus.”

THE POWDERY MILDEWS 231

“Tn the process of spore formation we have again a most strik-
ing example of the controlling influence of the so-called nucleo-
cytoplasmic relation. The nucleus of the ascus divides to form two
daughter nuclei, and these in turn divide successively to form eight
nuclei; but in thus passing from the uninucleated to the multi-
nucleated condition the nucleo-cytoplasmic equilibrium is main-
tained. The two daughter nuclei are proportionally smaller than
the mother nucleus, and the four and eight nuclei in the end bear
approximately the same relation to their cytoplasmic masses as did
the primary nucleus of the ascus to the cytoplasm of the entire ascus.
The two nuclei which become the centers for the formation of spores
grow to a somewhat larger size than the remaining six, and accord-
ingly the mass of cytoplasm included in the two spores is more than
one-fourth of that of the entire ascus.”

Another fact to be explained is the universal triple division
which occurs in the ascus of the powdery mildews as well as in
that of all the higher Ascomycetes. The fusion nucleus regularly
divides three times in rapid succession forming eight nuclei. It is
very common for eight ascospores to be formed, although frequently
the number is less, due to the non-functioning of one or more.

It is well known that in all the higher plants there is a double
division of the spore mother cell during which the number of
chromosomes is reduced. The doubling of the chromosomes occurs
upon the fusion of the nuclei of the two sex cells. Since this double
division is necessitated by the single fusion, Harper believes that the
double nuclear fusion in the mildews, one in the oogonium and one
in the ascus, necessitates a triple division in order to bring about
the corresponding reduction in the number of the chromosomes.
Harper also interprets the ascus as a spore mother cell.

Host Distribution. Some morphological species of the
powdery mildews are limited to a single host. For example,

Podosphaera biuncinata occurs only on Hamamelis virginiana, and
Uncinula geniculata on Morus rubra. Others are confined to the
species of a single genus of host plant, for example, Uncinula cir-
cinata on Acer, Erysiphe aggregata on Alnus. Again some occur
on a number of different genera all of which belong to the same
family, for example, Erysiphe graminis on various grasses. Several

232 GEORGE M. REED

of the mildews, however, have a very wide range of hosts. Perhaps
the most striking cases are Erysiphe polygoni on 355 hosts belonging
to 42 families, E. cichoracearum on 280 hosts belonging to 27 fami-
lies and Phyllactinia corylea on 144 hosts belonging to 36 families.

Several cases are known where two or more different species
of mildews occur on the same host. These may have the same or
a different geographical distribution. For example, Microsphaera
alni and Phyllactinia corylea both occur on the common hazelnut,
Corylus americana; Erysiphe polygoni and Microsphaera alni occur
on Lathyrus venosus; and three different mildews, Sphaerotheca
mors-uvae, Microsphaera grossulariae and Phyllactinia corylea have
been reported on Ribes Grossularia.

Biologic Specialization. In recent years a great deal of ex-
perimental work has been done to determine whether a mildew
growing on one host can produce infection on another plant which
is known to be infected by the same morphological species. Neger
(39) was the first to show that some of the morphological species
of mildews are broken up into biologic forms, limited to definite host
plants. Marchal (35-36), Reed (45-49), Salmon (52-57), Steiner
(63), and Voglino (68), have very greatly extended our knowledge
regarding the biological specialization of different mildews.

Thus far one or more species of five genera-Uncinula,
Erysiphe, Microsphaera, Phyllactinia and Sphaerotheca—have been
tested for host specialization, no results as yet having been recorded
for Podosphaera. Most of the data obtained, however, are very in-
complete except possibly in the case of three species of Erysiphe,
namely E. Polygoni, E. cichoracearum and E. graminis, and one
species of Sphaerotheca, Sph. humulh. -

The grass mildew, Erysiphe granunis, has proven to be excep-
tionally favorable for studies on biologic specialization. The result
has been the discovery of a large number of facts bearing upon
this problem. As already noted this morphological species occurs
on approximately sixty different grasses. So far, however, the work
indicates that in practically every case the biologic forms are re-
stricted to the species of a single genus. For ’example, conidia
from wheat will not infect barley, oats nor rye; conidia from barley

THE POWDERY MILDEWS 233

will not infect wheat, oats nor rye. In other words, in every case
the mildew on one cereal is unable to pass over onto the species of
other cereals.

Not all species of a particular genus, however, may be sus-
ceptible to the mildew. For example, the mildew on barley will infect
the common barley and also Hordeum decipiens, H. hexastichon, H.
intermedium, H. bulbosum, H. distichum, H. maritinum and H.
zeocriton, but will not pass over onto H. jubatum, H. murinum, H.
secalinum nor H. sylvaticum. Young plants of Hordeum nodosum
are easily infected with the barley mildew but older plants are im-
mune. A similar case has been found by Salmon (56) among the
brome grasses. As a result of his work upon the mildews of the
brome grasses, Salmon believes that there are four or perhaps five
biologic forms within this one genus alone.

The wheat mildew, while able to pass over onto one or more
varieties of every species of Triticum tested, is not capable of infect-
ing all of the varieties of wheats. Ina recent paper (49) I published
the results of tests with seventy-eight varieties distributed among
nine different species of this genus. Of these seventy-eight varie-
ties four proved to be immune, two belonging to T. dicoccum and
two to T. vulgare. Ina few other cases the percentage of infection
was rather low but in a great majority of cases the percentage of
infection approached 100.

In some unpublished results I have been able to transfer the
wheat mildew to different species of Aegilops. Several species were
tested and nearly all of them proved highly susceptible. If Aegilops
is to be regarded as a distinct genus as some systematists believe, we
have a case of one biological form of grass mildew occurring on the
species of two different genera. A similar relation is found in the
case of the oat mildew, for I have been able to confirm Marchal’s
statement that the conidia from oats are able to infect seedlings of
Arrhenatherum elatius, although the percentage of infection is not
very high.

Well developed biologic forms have been found on other
grasses. The mildew on species of Agropyron are confined to the
hosts of this genus. Similarly the orchard grass mildew is confined
to Dactylis glomerata and the blue grass mildew to species of Poa.

234 GEORGE M. REED

In addition to the grass mildew very full results have been ob-
tained with Erysiphe cichoracearum. As already noted, this mildew
occurs upon 280 hosts belonging to 27 families. It is therefore a
very cosmopolitan species. The results indicate, however, that it
is broken up into biologic forms. I (47) have found that distinct
forms occur on cucurbits, asters and golden rods. I have also found
that the cucurbit mildew occurs on at least eleven species of cucur-
bits belonging to seven genera. Six other species were also slightly
infected. In fact only three species of cucurbits tested remained
entirely resistant to the mildew. I further found that plants of
Plantago rugelii and of Helianthus annuus could be infected by the
mildew growing upon the cucurbits. In this case therefore this
biologic form not only occurs on several genera of a single family
but even passes out beyond the limits of the family.

Salmon (56) in his work on the mildew of the brome grasses,
and Steiner (63) more recently in his work with the mildew of
Alchemilla, have described what they call “bridging species.” Sal-
mon found that the mildew on Bromus racemosus failed to infect
B. commutatus (twelve trials), while it infected B. hordeaceus in
one hundred per cent of the cases (thirty-four trials). Further-
more, conidia from B. commutatus failed to infect B. racemosus
(thirty-six trials), while the mildew occurring in nature on B.
hordeaceus infected B. commutatus (forty out of forty-nine trials).
From these data, Salmon concludes that B. hordeaceus may act as a
“bridge” for the mildews on B. racemosus and B. commutatus.
Salmon tested this in one case by infecting B. hordeaceus with
conidia from B. racemosus. The conidia produced on the former
were then used to infect B. commutatus. Similarly Steiner regards
Alchemilla pastoralis as a “bridge” for Spherotheca humuli to pass
from A. connivens to A. micans.

The question of the degree of biologic specialization in other
mildews is one of great interest and one which can be attacked with
relative ease. There are many cases of peculiar host distribution
which should be investigated from the physiological standpoint. For
example, Microsphaera diffusa is reported by Salmon in his mono-
graph as occurring on twelve herbaceous legumes and three species
of Symphoricarpos, a shrubby plant belonging to the Caprifoliacez.

THE POWDERY MILDEWS 235

To be sure a distinct species, based on morphological characters, is
recognized by some as occurring on Symphoricarpos. Infection ex-
periments would determine whether the mildew on the legumes is
transferable to Symphoricarpos or not and might indicate whether
the minor morphological differences are of sufficient importance to
regard the form on Symphoricarpos as a distinct species.

It is further of interest that Phyllactinia corylea occurs on such
a wide range of hosts and that the one species of the genus has a
world wide distribution. While certain workers have described
other species of the genus yet the great morphological similarity of
this mildew on its various hosts is very striking. Voglino (68) has
found some evidence for biologic specilization in this mildew.

Economic Importance. The mildews are obligate parasites
and as such frequently cause a great deal of injury to their hosts.
While as a general rule these fungi are not considered as destructive
as the rusts and smuts, still there are many plants of economic im-
portance that are seriously attacked by some species of powdery
mildew.

The powdery mildew of the grape, Uncinula necator, has for
a long time been recognized as a serious menace to the culture of
grapes in various parts of Europe. In 1847 Berkeley (7) reported
that “the grapes in the neighborhood of Margate (England) have
for the past two years been attacked by a peculiar mildew of a most
destructive character.” Very soon after this the disease was re-
ported from the vineyards of southern France and other European
countries, causing destruction to the grape harvest.

The American gooseberry mildew, Sphaerotheca mors-uvae,
has proven very destructive to the English varieties of gooseberries,
so that their cultivation in this country is attended with great loss.
The fungus attacks especially the fruit, covering the berries with
a close felt-like mycelium. The mycelium becomes dark colored
and thick-walled with age and the perithecia are imbedded in it. In
recent years (21, 58) the mildew has occurred in epidemic form in
England and other European countries. Its appearance everywhere
has occasioned great loss to growers of gooseberries, not only by de-
stroying the year’s crop of fruit but also by weakening the bushes
themselves through injury to twigs and leaves. It is interesting to

236 GEORGE M. REED

note that the European gooseberry mildew, Microsphaera grossu-
larie, is not ordinarily troublesome although injury has been re-
ported, the affected leaves becoming shrivelled and falling prema-
turely.

The rose mildew, Sphaerotheca pannosa, is often very destruct-
ive to roses. While especially likely to attack roses cultivated in
the greenhouse, the fungus also attacks certain varieties in the gar-
den. The crimson rambler is notably susceptible, for the fungus
develops on the young leaves causing these to curl and arch; the
young stems are also attacked and more or less deformed. It also
attacks the flower buds forming a white mealy growth and blasting
the flowers. The conidia are formed in great abundance and the
disease spreads very rapidly.

The hop, in many regions, suffers serious injury from the
spread of the hop mildew, Sphaerotheca humuli. The worst damage
is done when the disease attacks the cones causing them to shrivel
up. In some parts of England this disease is much dreaded by the
hop growers. Comparatively recently the hop mildew has become
serious in the hop yards of New York (9g). In some counties the
entire hop crop has been ruined by the ravages of this fungus while
in others the yield has been decreased and the quality of the crop
much impaired.

Among other plants that are sometimes seriously attacked by
the mildews the following may be mentioned: Grasses are some-
times seriously injured by the spread of Erysiphe graminis. Ander-
son (2) reports that this mildew in the northwestern states “affects
chiefly the Poas and is especially damaging to P. tenuiflora, one of
our most valued forage grasses.” Stewart (64) has reported the
occurrence of the mildew on wheat in the spring but, although the
disease spreads very rapidly, no serious permanent injury to the
wheat crop results. Cucumbers when forced in the greenhouse are
sometimes severely attacked by Erysiphe cichoracearum. Nursery
stock, as apples, peaches and cherries, are frequently damaged by
powdery mildews.

As a general rule it is not difficult to control the outbreaks of
diseases due to powdery mildews. Repeated experience has shown
that flowers of sulphur, dusted on the infected plants, is a satisfac-

i

THE POWDERY MILDEWS 237

tory remedy. Most of the mildews seem to be controlled by this
treatment, especially when prompt measures are taken at the first
indication of an outbreak. A relatively strong solution of potassium
sulphide—one ounce in two gallons of water—is also effective in
the treatment of some diseases caused by mildew.

Pathological Effects. As a general thing the mildews do
not cause any striking pathological changes in the cells of the host.
Ordinarily no appreciable malformation occurs or hypertrophy.
However, an enlargement of the twigs of Physocarpus, similar to
Witches’ Brooms, are produced by the development of Sphaerotheca
humauli.

One effect of some of these mildews is very striking. When
Maple leaves infected with Uncinula circinata are collected in the
late fall they will be found to show characteristic yellow and green
areas. Examination shows that the green areas are infected with
the mildew. The infected host cells are stimulated to retain their
green color longer than the other cells of the leaf. A similar effect
may be seen on grass leaves when badly covered with the mildew.
The results may be made more striking by placing leaves of barley,
for example, in a moist glass chamber and inoculating with conidia
from barley. In three to four days numerous infected areas will
be distinctly visible and possess a deep green color while the other
portions of the leaves will be a pale yellow.

Collection, Preservation and Cultivation of the Powdery
Mildews. The powdery mildews are usually conspicuous ob-
jects on leaves and other parts of living plants. Sometimes careful
search is needed to locate the perithecia. When hunting for mil-
dews one needs to examine both surfaces of the leaves for many
forms occur mainly or entirely on the under surface of leaves. This
is particularly true of Phyllactinia corylea.

Special care, of course, must be taken to properly identify the
host plant. In fact it is generally a good plan to collect flowering
or fruiting parts of the host and keep these with the infected leaves.

For the systematic study of the powdery mildews dry material
is very satisfactory. Infected parts of plants are collected and

238 GEORGE M. REED

placed between driers for a few days. The material is then trans-
ferred to suitable envelopes, properly labeled, and laid aside for fu-
ture study.

Many of the mildews are easily cultivated in greenhouses and
afford exceptionally fine material for the study of various problems.
In general a succession of crops of young hosts must be provided
so that the mildew will be able to spread over on to the young plants.
The grass mildew on various hosts, when once secured, may easily
be propagated by planting additional seed of the proper hosts at in-
tervals of one to four weeks. Various other mildews are also read-
ily propagated in a similar manner. The writer has successfully cul-
tured for long periods the mildews on cucurbits, goldenrod, aster,
Erigeron, apple, sunflower, various cereals and grasses, etc. As
pointed out by Melhus: (37) the sunflower mildew is particularly
favorable for class use because of the readiness with which peri-
thecia appear upon the host.

The writer has experienced a great deal of difficulty in propa-
gating the mildews during the summer months. The high tempera-
ture of the greenhouse seems to be the controlling factor. The mil-
dew, however, is especially infested by thrips, a small insect which
seems to spread very rapidly during the summer season.

Classification. The powdery mildews have long been an
object of study from the systematic standpoint. Practically all of

the earlier mycological workers have devoted more or less attention
to the group. Probably the first mention made of one of these
fungi was by Linnaeus in Species Plantarum (1753) under the
name of Mucor Erysiphe. Following Linnaeus, Persoon (42, 43),
Rebentisch (44), (who published the first illustration), de Candolle
(13), Fries (24), and others made additions to our knowledge of
mildews. The most important papers dealing with the systematic
arrangement of these fungi are those by Wallroth (69, 70) (1819)
who insisted upon the distinction of species on the basis of morpho-
logical characters and not by the host plant upon which they grew;
by Schweinitz (60) (1834), who described several North American
forms; by Léveillé (33) (1851), who divided the group into six
genera, based upon the number of asci in the perithecium and the
characters of the appendages, the names and limits of these genera

THE POWDERY MILDEWS 239

being retained to the present day; by the Tulasne brothers (66)
(1861), who described and very fully illustrated by exceptionally
fine copper plates sixteen species, although they placed them all un-
der one genus—Erysiphe ; by Cooke & Peck (14) (1872) who listed
the mildews of the United States; by Burrill (12) (1892) who de-
scribed the North American forms then known; by Salmon (50)
(1900) who has monographed the group with exceptional complete-
ness and whose work is the standard at the present time.

Many lists from various states have also been published. These
give us much desired information regarding the occurrence and dis-
tribution of the mildews in the United States. Among the more
important of these state lists the following may be mentioned: An-
derson (Iowa) (1), Anderson (Montana) (2, 3), Atkinson (Caro-
lina and Alabama) (4), Brannon (Indiana) (10), Burrill and Earle
(Illinois) (11), Davis (Wisconsin) (17, 18, 19, 20), Farlow
(Massachusetts) (22), Freeman (Minnesota) (23), Griffiths
(northwestern states) (26), Harkness and Moore (California)
(27), Lawrence (Washington) (32), Millspaugh and Nuttall (West
Virginia) (38), Selby (Ohio) (61), Tracy and Earle (Mississippi)
(65), Underwood and Earle (Alabama) (67), Walters (Kansas)
(71).

In the following classification of the Erysiphacee Salmon
(50, 51) is largely followed. Most of the facts stated are taken
from his monograph and supplementary notes.

Salmon in his monograph (50) recognizes six genera, forty-nine
species and eleven varieties.

Saccardo on the contrary recognizes a much larger number of
species. Of Salmon’s species and varieties, thirty-one species and
seven varieties are found in North America. Of these thirteen
species and five varieties have been reported only from this con-
tinent.

Some interesting cases of distribution may be noted. The ma-
ple mildew of North America is Uncinula circinata, while a different
species, 4. aceris is reported on the maples of Europe. Sphaerotheca
morse-uvae in North America specially attacks species of Ribes; in
Europe this same form is reported on species of Euphorbia. Very
recently, as already noted, this mildew has also appeared on Ribes

240 GEORGE M. REED

in Europe, but it is thought to have been introduced from America.
Microsphaera alni is very common on the lilac, Syringa vulgaris, in
this country, but according to Magnus (34) is not found on this
host in Europe, although the fungus on other hosts is quite common.

Following Léveillé, Salmon divides the Erysiphacee into six
genera’ and uses the same generic names as proposed by Léveillé.
The following keys, which are based on Salmon’s monograph and
which are quite generally used in this or in slightly different form,
may serve to differentiate the genera and species.

Key To GENERA

A. Perithecium containing a single ascus B.
B. Appendages unbranched, more or less flexuous, arising from the base

Ofa thee pertthectttiles cc coca ee ee Meeks eo lette eee Spherotheca.
B. Appendages one to several times dichotomously branched at the
ADO Cree Sete rete tee Shade Ty ahe ua aeeR Uae ace Sere rae eer ee Podosphera.

A. Perithecium containing several asci C.
C. Appendages without bulbous enlargement at the base D.
D. Appendages generally straight, dichotomously branched at the

UPON Peeters: coe ak ect er ra teereei ote sree Siete cenc rae Microsphera.
D. Appendages simple, uncinate or spirally inrolled at the apex.
1A Ee ETE DIOR NG a CUE Sine Grane Uncinula.

D. Appendages simple or irregularly branched, more or less flex-
uous, usually somewhat similar to the mycelial hyphe, not
dichotomously branched nor uncinate at the apex....Erysiphe.

C. Appendages simple, straight, rigid, with a bulbous enlargement at
Ges BaGS ie he Sk te Lec a kad teens ae ee ee ee Phyllactinia.

SPHAEROTHECA Léveillé. (Plate XVI, Figs. 38, 38a)

The perithecia are subglobose, containing a single ascus which
is regularly 8-spored. The appendages are flexuous, ‘brown or
colorless, spreading horizontally and often interwoven with the
mycelium which they frequently resemble; they are simple or
rarely branched, sometimes lacking.

Sphzrotheca is represented by five species and one variety,
all of which have been reported from the United States.

1. Salmon (59) has since placed Erysiphe taurica in a new genus by itselfi—
Oidiopsis—making in all seven genera.

THE POWDERY MILDEWS 241

Key To SPECIES

A. Mycelium persistent, thick, pannose, forming dense patches composed
of special hyphz, in which the perithecia are more or less immersed B.

B.

Persistent mycelium usually satiny and shining, white, sometimes
becoming gray or pale brown............ S. pannosa (Wallr.) Lév.

B. Persistent mycelium dark brown C-.

C. Inner wall of the perithecium separating from the outer; hyphe
of persistent mycelium very tortuous....... S. lanestris Harkn.

C. Inner wall not separating, hyphae straighter...............
ROO RAR IPSEC BE OSS ONT RET S. mors-uvae (Schw.) B. & C.

A Mycelium without these characters D.
D. Perithecia 60-784 in diameter, ascus 60-75x42-50u; inner wall of

perithecium separating from the outer..S. phytoptophila K. & S.

D. Perithecia 50-1204 in diameter, ascus 45-90x50-72u; inner wall scarce-

Hosts:

ly separating from the outer E.
E. Cells of outer wall of perithecium 10-20% wide, averaging I5u
Bes ea cl SOAP rere ho aces Eee S. humuli (DC.) Burrill.
E. Cells of outer wall of perithecium 20-30 (rarely 40) wide,
averaging 25u...... S. humuli var. fuliginea (Schlecht.) Salm.

Spherotheca pannosa; Rosa (various species).

S. lanestris: Quercus agrifolia, Q. alba, Q. macrocarpa.

S. mors-uve: Ribes (various species).

S. phytoptophila: Celtis occidentalis.

S.humuli: Agrimonia striata, Geranium maculatum, Geum cana-
dense, Physocarpus opulifolius.

S.humuli var. fuliginea: Bidens chrysanthemoides, B. frondosa,
Erigeron annuus, Taraxacum officinale.

PoposPH#RA Kunze. (Plate XVI, Figs. 34, 34a).

Perithecia globose, or globose-depressed, containing a single
ascus which is 8-spored. The appendages are equatorial or apical,
dark brown or colorless, dichotomously branched at the apex,
ultimate branches simple and straight or swollen and knob-shaped.

The genus contains four species and one variety, all, except
one species, being represented in the United States.

Key to Species

A. Basal appendages present in addition to the apical, the latter usually
unbranched. sm .encucoe ts och as P. leucotricha (Ell. & Everh.) Salm.
A. No basal appendages present B.

242 GEORGE M. REED

B. Appendages arising from near the apex of the perithecium, some-
what erect and fasciculate, one to eight times the diameter of
the perithecium, dark brown for more than half their length
atid Raised hacen Ebest tare P. oxyacanthe var. tridactyla (Wallr.) Salm.

B. Appendages equatorially inserted and more or less spreading C.

C. Appendages colorless, or faintly tinged with brown at the base,
branched apex not swollen............. P. biuncinata C. & P.

C. Appendages dark brown for more than half their length, ultimate
branches of. the apex. knob-shaped ..20\55..<> sc> ke onamh eee

S hs Shel erate Aieiare ee Re deed he RES P. oxyacanthe (DC.) De Bary.

P. leucotricha: Pyrus malus.

P. biuncinata: Hamamelis virginiana.

P. oxyacanthe: Prunus (various species).

P. oxyacanthe var. tridactyla: Spirea Douglasii.

UNcINULA Léveillé. (Plate XVI, Figs. 37, 37a).

This genus is easily distinguished by the uncinate apex
of the appendages. The perithecia are globose or globose-depressed
and contain several asci, two to eight spored. The appendages are
simple in all American forms.

There are eighteen species and two varieties. Of these ten
species occur in the United States.

Key to Species

A. Appendages colored for half their length or more..................--
Ss ncaa rk lc A EEN (oven DS eee ONY A U. necator (Schw.) Burrill.
A. Appendages colorless B.
B. Asci containing 2-3 spores C.
C. Perithecia very large 215-3202 in diameter; more than 30 asci

in the perithecium............. U. polychaeta (B. & C.) Ellis
C. Perithecia averaging 1304; asci 8-20 in the perithecium....
at 4h). 5 EEE Mae RT Ete cL AE RRO U. macrospora Peck.

B. Asci containing 4-8 spores D.
D. Appendages delicate, narrow, 3-4u wide; asci 4-7 spored E.
E. Perithecia 150-2004 in diameter; asci about 25..............
MI, Ody a ene ee eres «Aha cea U. confusa Massee.
E. Perithecia 86-122u in diameter; asci 5-8 G.
G. Appendages 50-160, 14-34 times the diameter of perithe-
CROMES £54 caus oe ies em nee oe oe U. parvula C. & P.

1. Reported only from Washington.

THE POWDERY MILDEWS

243

G. Appendages 24-46, 114-2 times the diameter of perithecium,

Ofien> geniculate... oo sa. eee woe cos U. geniculata Gerard.

D. Appendages stouter, wider, or if narrow with asci 8-spored F.
F. Appendages abruptly flexuose, about equalling the diameter of
perithecium, spores usually 8............... U. flexuosa Peck.

F. Appendages all straight H.

H. Appendages thick walled, refractive or rough at base, per-
ithecia 64-146 in diameter............ U. clintonii Peck.

H. Appendages thin walled throughout I.

I. Perithecia 90-1754 in diameter, averaging 135”; asci con-

taining 4-6 spores.......... U. salicis (DC.) Winter.
I. Perithecia 160-2254 in diameter, averaging 1I90u; asci
containing 7-8 spores........... U. circinata C. & P.

Hosts:

Uncinula necator: Psedera quinquefolia, Vitis (various species).

polychaeta’: Celtis occidentalis.
macrospora: Ulmus americana, U. fulva.
confusa’: Celtis occidentalis.

parvula: Celtis occidentalis.

geniculata: Morus rubra.

flexuosa: Aesculus glabra.

clintonti: Tilia americana.

her Uy cere

circinata: Acer saccharinum.

ErysipHE Hepw. (Plate XVI. Figs. 36, 36a).

salicis: Populus (various species), Salix (various species).

Perithecia are generally globose and contain several asci, 2-8
spored. Appendages are floccose, simple or irregularly branched,
never with a definite apical branching, sometimes obsolete, usually

more or less similar to the mycelium and interwoven with it.

The genus contains eight species and one variety of which

seven species and the variety occur in the United States.

Key to Species

A. Perithecia large, 135-2804 in diameter, averaging 200u B.

B. Perithecia immersed in a lanuginose persistent mycelium; asci not
containing spores on the living host.............. E. graminis DC
B. Perithecia not immersed in a lanuginose persistent mycelium; asci

containing spores on the living host C.

1. Reported only from S. Carolina, Alabama and Mississippi.
2. Never found but once and then associated with U. parvula.

244 GEORGE M. REED

C. Asci containing eight spores, rarely six or seven; spores some-

what spherical, 16-20x10-15u.....E. aggregata (Peck) Farlow.
C. Asci containing four to six spores; spores 20-22XI0-I2u......
Perkasie teks E. polygoni var. sepulta (Ell. & Everh.) Salm.
C. Asci containing two spores; spores 28-40x18-224; perithecia at
maturity becoming cup-shaped................. E. taurica Lév.

A. Perithecia smaller, 65-180, not immersed in a lanuginose mycelium D.

D. Asci not containing spores on the living host; haustoria lobed....

Sittchsors a aly Tee ve ies eal dal todd Seema eee E. galeopsidis DC

D. Asci generally containing spores on the living host; haustoria not
lobed E.

E. Perithecia containing 4-25 asci, usually 10-15; asci 58-90x30-50n;

spores generally two, 20-28x1I2-20m...... E. cichoracearum DC.

E. Perithecia containing few asci, 2-8, rarely as many as 22; asci

46-72x30-45u; spores 3-8, rarely 2, 19-25x9-14u.E. polygoni DC.

A. Perithecia small, 52-6ou in diameter, containing usually three asci; asci

48-50x28-364; asci two, rarely three, spored...... E. trina Harkn.

Hosts:

Erysiphe graminis: Agropyron repens, Avena sativa, Dactylis
glomerata, Hordeum vulgare, Poa pratensis, Secale cereale, Trit-
icum vulgare.

E. aggregata: Alnus incana (catkins).

E. galeopsidis: Scutellaria latifolia, Stachys tenuifolia.

E. polygoni: Astragalus canadensis, Clematis virginiana, Lathy-
rus venosus, Polygonum aviculare, Ranunculus abortivus.

E. polygoni var. sepulta’ Bigelovia graveolens.

E. cichoracearum: Actinomeris alternifolia, Ambrosia trifida, As-
ter cordifolius, A. laevis, A. sagittifolius, Eupatorium purpureum,
Helianthus annuus, Cucumis sativus, Cucurbita maxima, Solidago
canadensis.

E. trina®: Quercus agrifolia.

E. taurica®: Heliopsis scabra.

MicrospHarA Léveillé. (Plate XVI. Figs. 39, 39a).

The perithecia are globose to globose-depressed and contain
several asci, 2-8 spored. The appendages are not interwoven with
the mycelium; they are divided several times in a dichotomous
manner at the apex.

1. Reported from Rocky Mountain States.

2. Reported only from California.

3. Reported by Salmon as an old world species but recorded by Anderson (1) from
Towa.

THE POWDERY MILDEWS 245

The genus is represented by thirteen species and six varieties
of which five species and four varieties occur in North America.

BI Key to Species

A. Appendages 214-7 times the diameter of the perithecia, usually much
contorted and angularly bent; apical branching of appendages very
irregular and lax, with the branches flexuous and more or less
PNRICR eins Seat tae de 4k SEs eo aeons sole M. euphorbie (Peck) B. & C.

A. Appendages long or short without above characters B.

B. Tips of some or all of the ultimate branches of the appendages

recurved C.
C. Appendages long and flaccid D.

D. Apex of appendages much branched, ornate, more or less

GlOSESP SPOLES 1 22—2OWIA=TWS tarsiacts oo cn esiaeere a sea nee A sicle rs ste

Be a8 Ah Ae sor M. alni var. extensa (C. & P.) Salm.

D. Apex of appendages less branched, more or less widely

forked, or branching close and simple; spores 18-23x9-13u

ALA Rb Snares M., alni var. vaccinii (Schw.) Salm.

C. Appendages short, not exceeding 2% times the diameter of the

perithecium E.

E. Appendages more or less contorted, apical branching very

laxeandiisregilate ny. . er one M. alni var. ludens Salm.

E. Appendages not contorted, apical branching closer and reg-
ular; tips regular, recurved F.

F. Axis of some appendages not dividing dichotomously
at the apex, but bearing sets of opposite branches....
aA eee M. alni var calocladophora (Atk.) Salm.

F. Appendages regularly dichotomous at apex............
Re POP RAN oe) are peri hh M. alni (Wallr.) Salm.

B. Tips of appendages not recurved G.

G. Appendages 3-7 times diameter of perithecia; colored, nearly to
APOK) . Valle dees be sae Saat eet ee M. Russellii Clinton.

G. Appendages colorless H.
H. Branching of appendages lax, irregular; ultimate branches
long, forming a narrow fork.......... M. diffusa C. & P.
H. Branching closer and more regular; apex of appendages
with very short primary and secondary branches, more
or less digitate.............M. grossularie (Wallr.) Lev.

Hosts:

M. euphorbiae: Euphorbia corallata.

M. alni: Gleditsia triacanthos, Lathyrus odoratus, Platanus occi-
dentalis, Syringa vulgaris, Viburnum Lentago.

M. alni var. vaccinii: Vaccinium (various species).

240 GEORGE M. REED

M. alni var. ludens’: Vicia americana.

M. alni var. extensa: Quercus (various species).

M. alni var. calocladophora*: Quercus aquatica.

M. Russellii: Ovxalis stricta.

M. diffusa: Desmodium canadense, Lespedeza capitata, Symphori-
carpos orbiculatus.

M. grossularie: Ribes nigrum, Sambucus canadensis.

PHYLLACTINIA Léveillé. (Plate XVI. Figs. 35, 35a).

This genus is easily recognized by the large (140-270 in
diam.) globose-depressed perithecia with the equatorial rigid, color-
less, acicular appendages with a bulbous swelling at the base. The
perithecia contain many asci which are regularly 2-spored, rarely
3-spored. The apex of the perithecium is provided with numerous
crowded, branched penicillate cells which arise from the outer cells
of the perithecial wall.

The mycelium of this genus is characteristically internal, de-
veloping in the intercellular spaces of the leaf. Special branches,
haustoria, penetrate into the cells of the spongy parenchyma.

This genus is widely distributed and is represented by a single
species—Phyllactinia corylea (Pers.) Karst.

Hosts:

Betula alba, Celastrus scandens, Cornus stolonifera, Corylus Amer-
icana, Osirya virginiana.

Host Index. The following list includes about one hundred
and seventy-five hosts of the Erysiphaceze. The list is by no means

intended as a complete one of the powdery mildews of the United
States, but it does include the common hosts, most of which have
a wide distribution and upon which the various species of mildews
may be found. Collections of nearly all of these are found in the
herbarium of the writer. A few are taken from the records of
different state lists. In naming the hosts, Gray’s New Manual of
Botany, revised by Robinson and Fernald, is strictly followed.

1. Reported only from S. Dakota and Montana.
2. Reported only from S. Carolina, Alabama and Mississippi.

THE POWDERY

NCEE AE UD Gtttile [rey yaya s\n) adele airs

Acer saccharum Marsh
Mices saccharinium Us. save cuewaes «
Actinomeris alternifolia (L.) DC....
MeScHilisy glabray Willdzu.. aaa cor.
Agrimonia striata Michx
Agropyron repens (L.) Beauv
Alnus incana (L.) Moench
Ambrosia artemisifolia L
Pmanrosia: thitidas Li. as oxen ds ween
Amphicarpa monoica (L.) Ell
Apios tuberosa Moench..............
Aquilegia canadensis L
Aquilegia coerulea James
Aster cordifolius L
Aster laevis L
Aster paniculatus Lam
Aster puniceus L
Aster sagittifolius Wedemeyer
INStet hcadescaniti lx -2i faces derenlaivion
Astragalus canadensis L
| AIRES Tri 7 0 0 ee
Betula alba L. var. papyrifera (Marsh)

‘S [DEVE SS eC SOR RE ER eae Ra
Bidens cernua L

ey .

eee ewww
© a me see se @

Sea ole eve e.

re ey
wa) oe 61a 6, oe \e)'s © 600) 616.0 0,6 8 oe

ey

{oy bce) 6a) a! .6, 6 oe! UO, a 4, 6.0 6x8 0. 8 are

Bidens levis (L.) BSP

BAdense dt tondosSaw yarns so acl6 ase eiete

Bigelovia graveolens A. Gray........
Carpinus caroliniana Walt...........
Castanea dentata (Marsh.) Borkh...
Catalpa spectosa \Watdeti. ave5os2 se
Ceanothus americanus L
Celastrus scandens L
Celtis occidentalis L
Celtis occidentalis L
Celtis occidentalis L
Chelone glabra L
Cirsium lanceolatum (L.) Hill
Cirsium muticum Michx

O) 0 MG S's, # wie OM el tls) mS

ee

Cie) @ ete) e Ole of eae, @ aeeiele

ee

MILDEWS

Funcus
Uncinula circinata C. & P.
Uncinula circinata C. & P.
Uncinula circinata C. & P.
Erysiphe cichoracearum DC.
Uncinula flexuosa Peck.
Spherotheca humuli (DC.) Burr.
Erysiphe graminis DC.
Erysiphe aggregata (Peck) Farl.
Erysiphe cichoracearum DC.
Erysiphe cichoracearum DC.
Erysiphe polygoni DC.
Microsphera diffusa C. & P.
Erysiphe polygoni DC.
Erysiphe polygoni DC.
Erysiphe cichoracearum
Erysiphe cichoracearum
Erysiphe cichoracearum
Erysiphe cichoracearum
Erysiphe cichoracearum
Erysiphe cichoracearum
Erysiphe polygoni DC.
Erysiphe graminis DC.

DE
DC.
DC.
DC.
DC.
DC.

Phyllactinia corylea (Pers.) Karst.
Spherotheca humuli (DC.) Burr. var.
fuliginea (Schlecht.) Salm.
Spherotheca humuli (DC.) Burr. var.
fuliginea (Schlecht.) Salm.
Spherotheca humuli (DC.) Burr. var.
fuliginea (Schlecht.) Salm.
Erysiphe polygoni DC. var. sepulta (E.
& E.) Salm.
Microsphera alni (Wallr.) Salm.
Microsphera alni (Wallr.) Salm.
Microsphera alni (Wallr.) Salm. var.
vaccinii (Schw.) Salm.
Microsphera alni (Wallr.) Salm.
Phyllactinia corylea (Pers.) Karst.
Spherotheca phytoptophila K. & S.
Uncinula polycheta (B. & C.) Ellis
Uncinula parvula C. & P.
Erysiphe galeopsidis DC.
Erysiphe cichoracearum DC.
Erysiphe cichoracearum DC.

248 GEORGE M. REED

Clematis’ vitginiana TWaicc. es te csiccaets Erysiphe polygoni DC.

Cornus alternifolia’ bats ee eevee Microsphera alni (Wallr.) Salm.

Cornus stolonifera Michx............. Phyllactinia corylea (Pers.) Karst.

Corylus americana Walt.<'.... 50. ..... Microsphera alni (Wallr.) Salm.

Corylus americana Walt.............. Phyllactinia corylea (Pers.) Karst.

Cratecus Crasswalli. Goose as bo ieaiee Podosphera oxyacanthe (DC) de B.

Cratees -coccitied lic. : coatee dace eh Podosphera oxyacanthe (DC.) de B.

Crepis acuminata “INGER. ss os Erysiphe cichoracearum DC.

Gucumispeatnvirs 1s, enter che kere Erysiphe cichoracearum DC,

Cucurbita maxima Duchesne......... Erysiphe cichoracearum DC.

Cucurbita sPepo las icwiht caierees ei cist Erysiphe cichoracearum DC.

Dactylis’ slomerata (Lior ice saree Erysiphe graminis DC.

Desmodium canescens (L.) DC....... Microsphera diffusa C. & P.

Desmodium canadense (L.) DC....... Microsphera diffusa C. & P.

Desmodium paniculatum (L.) DC.....Microsphera diffusa C. & P.

Elymus. striatus Willd.) ic. 82.0004 Erysiphe graminis DC.

Erigeron annuus (L.) Pers:......... Spherotheca humuli (DC.) Burr. var.
fuliginea (Schlecht.) Salm.

Frigeron canadensis: o70i20) e862 Spherotheca humuli (DC.) Burr. var.

fuliginea (Schlecht.) Salm.
Eupatorium urticefolium Reichard... Erysiphe cichoracearum DC.

Eupatorium perfoliatum L............ Erysiphe cichoracearum DC.
Eupatorium purpureum L............. Erysiphe cichoracearum DC.
Euphorbia corollata clock creme ere oH Microsphera euphorbie (Peck) B.
& C.
Euphorbia marginata Pursh.......... Microsphera euphorbie (Peck) B.
& C.
Braxintiss amoenicanay lane ne acne ae Phyllactinia corylea (Pers.) Karst.
Fraxinus Pennsylvanica Marsh....... Phyllactinia corylea (Pers.) Karst.
Galtum -Aparine Wid ayatotenmete ee Erysiphe cichoracearum DC.
Geranium carolinianum L............ Spherotheca humuli (DC.) Burr.
Gerantdnr mactlatum ib..2-3.0 8 ees: Spherotheca humuli (DC.) Burr.
Gerardia grandiflora Benth........... Spherotheca humuli (DC.) Burr. var.
fuliginea (Schlecht.) Salm.
Genmremadence acd. 40822 i.e tees Sphzrotheca humuli (DC.) Burr.
Gilia micrantha (Kell.) A. Nels...... Spherotheca humuli (DC.) Burr.
Gleditsta triacamthos iis csc00e 0s Microsphera alni (Wallr.) Salm.
Grindelia squarrosa (Pursh.) Duval..Erysiphe cichoracearum DC.
Hamamielis witemana’ Ly.\0. odes we «s Podospheera biuncinata C. & P.
Heleniumimautaummnate Wiis) Erysiphe cichoracearum DC.
Helianthiswanniunsele.aene eee ae Erysiphe cichoracearum DC.
Helianthus decapetalus L............. Erysiphe cichoracearum DC.
Helianthus-‘strumosus sl. 22) 0 ih Erysiphe cichoracearum DC.
Helianthus tuberosas is 2.52 85. Erysiphe cichoracearum DC.

HeltopsisscabravDuvaleaeis eee Erysiphe taurica Lév.

THE POWDERY MILDEWS

Hordeum vulgare L
Humulus Lupulus L
Hydrophyllum appendiculatum Michx.
Hydrophyllum macrophyllum Nutt...
Inula Helenium L

Lathyrus ochroleucus Hook..........
athyrissodonatis, Ds.) o's acleceis00.«' a6
Pehyras palisttis: Te... ssi. ianieincss =
Lathyrus venosus Muhl...............
Lespedeza capitata Michx............
Ligustrum vulgare L
Lonicera sempervirens -L.............
Ponicera tattarien ie ached. bescdieas,
Lupinus perennis L
Morus rubra L
Oenothera biennis L
Ostrya virginiana (Mill.) K. Koch...
WM VawVIGPINI ANAS css ciccsl aces stl 8
Meas weerietey Tonks Rise cts oad oi
Parietaria Pennsylvanica Muhl
Parnassia caroliniana Michx
Philag paniediata, Tbs. tc dcitse es.
Physocarpus opulifolius (L.) Maxim.
Pisum sativum L
Plantago major L
Plantago Rugellii Decne
Elatanus., occidentalis: Ib.352)..5.<.55052.
Poa pratensis L
Polygonum aviculare L

ee ee ee ee

sim (eta @ (cio, 0] o *\ Ble «8, 21a 0) eve) 6:0
ee)

Cas wa © oo Osu sie

Polygonum erectum L
Polygonum exsertum Small
Polygonum scandens L
Populus deltoides Marsh
Populus grandidentata Michx
Populus tremuloides Michx
Prunus Besseyi Bailey
Prunus (cultivated cherry & plum)...
Psedera quinguefolia (L.) Greene.
Pyrus malus L

PO ee a a a a
eee eee eeeee
5 718 © (Swe slp ew ew ee
nw ee © ee, wee ale
ee
ee ce ees eee

Mirercus: alba; Ma./s.0 eee tei elon =
Quercus alba L
Quercus agrifolia Née
Quercus agrifolia Née

re ae a ee Oe

ee ee ee

249

Erysiphe graminis DC.

Spherotheca humuli (DC.) Burr.

Erysiphe cichoracearum DC.

Erysiphe cichoracearum DC.

Erysiphe cichoracearum DC.

Microsphera alni (Wallr.) Salm.

Microsphera alni (Wallr.) Salm.

Microsphera alni (Wallr.) Salm.

Erysiphe polygoni DC.

Microsphera diffusa C. & P.

Microsphera alni (Wallr.) Salm.

Microsphera alni (Wallr.) Salm.

Microsphera alni (Wallr.) Salm.

Erysiphe polygoni DC.

Uncinula geniculata Gerard.

Erysiphe polygoni DC.

Microsphera alni (Wallr.) Salm.

Phyllactinia corylea (Pers.) Karst.

Microsphera Russellii Clinton.

Erysiphe cichoracearum DC.

Erysiphe polygoni DC.

Erysiphe cichoracearum DC.

Spherotheca humuli (DC.) Burr.

Erysiphe polygoni DC.

Erysiphe cichoracearum DC.

Erysiphe cichoracearum DC.

Microsphera alni (Wallr.) Salm.

Erysiphe graminis DC.

Erysiphe polygoni DC.

Erysiphe polygoni DC.

Erysiphe polygoni DC.

Erysiphe polygoni DC.

Uncinula salicis (DC.) Winter.

Uncinula salicis (DC.) Winter.

Uncinula salicis (DC.) Winter.

Podosphera oxyacanthe (DC.) de B.

Podosphera oxyacanthe (DC.) de B.

Uncinula necator (Schw.) Burr.

Podosphera leucotricha (E. & C.)
Salm.

Microsphera alni (Wallr.) Salm.

Spherotheca lanestris Harkn.

Erysiphe trina Harkn.

Spherotheca lanestris Harkn.

250 GEORGE M. REED

Otiercus: aquatica Walt: os... «> «eee Microsphera alni (Wallr.) Salm. var.
calocladophora (Atk.) Salm.

Quercus macrocarpa Michx.......... Sphzrotheca lanestris Harkn.

Quercus macrocarpa Michx.......... Microsphera alni (Wallr.) Salm.

Quercus Muhlenbergii Engelm........ Microsphera alni (Wallr.) Salm.

Quercus pedunculata Ehrh. var. fasti-

Siattan (Geran plan ce ek wii bie Sarena Microsphera alni (Wallr.) Salm.

Oiuerctigvermustlageee cca osc cc. eee Microsphera alni (Wallr.) Salm. var.
extensa (C. & P.) Salm.

Oiereuseribra: los acre ce eek ee eee Microsphera alni (Wallr.) Salm. var.
extensa (C. & P.) (Salm.)

Ouerciis wwelutina an. 2. seach ese Microsphera alni (Wallr.) Salm. var.
extensa (C. & P.) (Salm.)

Ranvnewlusabortivis do-= 2 asca sew ee Erysiphe polygoni DC.

PeatiiinCulas aAcCrisnli. seceiakicnn serene Erysiphe polygoni DC.

Bhs Sabra -Lajt asieteee ses cae eneees Spherotheca humuli (DC.) Burr.

Rds: typhina Lio. ice sks ifeeoanea ee Spherotheca humuli (DC.) Burr.

RibeswmGrossulariawlicscer. cctee eee ac Spherotheca mors-uvae (Schw.) B.
& C.

Rabes niece tines ae eh ee oe ee eee Microsphera grossularie (Wallr.) Lév.

Rosa (cultivated species, Crimson

RAMIER VC toe eine eee Somme ke Spherotheca pannosa (Wallr.) Lév.

Rubus ‘iseitus? 1.650553 wk secede we Spherotheca humuli (DC.) Burr.

Salix<cordatasMGhysenanke seine Uncinula salicis (DC.) Winter.

Saltx discolor: Wuhl ox. cee Uncinula salicis (DC.) Winter.

Salix: ‘itera eWalsileteo nso. soe eee Uncinula salicis (DC.) Winter.

Satmbicis canadeusts is. 328 fhe Microsphera grossularie (Wallr.)

Lev.

scatellaria’ laterifiora Lig. see ee Erysiphe galeopsidis DC.

Shepherdia argentea (L.) Nutt....... Spherotheca humuli (DC.) Burr.

Sicyos anetlatus diol ic< jis Dac ede Erysiphe cichoracearum DC.

Solanum carolinense- L... 2.3 0362.56 Erysiphe cichoracearum DC.

Spirea’ Douglasiv Hook.<. sa ccainen<- Podosphera oxyacanthe (DC.) de B.
var. tridactyla (Wallr.) Salm.

Stachys palustris L.................. Erysiphe galeopsidis DC.

Stachys tenusfolia Willd... ........%6.%. Erysiphe galeopsidis DC.

Symphoricarpos orbiculatus Moench... Microsphera diffusa C. & P.

Syrinea valears aera: kniele as tvs dec Microsphera alni (Wallr.) Salm.

Taraxacum dumetorum Greene....... Spherotheca humuli (DC.) Burr.

Taraxacum officinale Weber.......... Spherotheca humuli (DC.) Burr. var.
fuliginea (Schlecht.) Salm.

Teucrium, canadense! Ib.i 25200005520 Erysiphe galeopsidis DC.

Tilia-amencanadserieeoe wee ene ee Uncinula clintonii Peck.

Trfoham pratetiseLs, 22c37 cate Erysiphe polygoni DC.

Triticum ‘yulwaré “Villas as eeeekt Erysiphe graminis DC.

THE POWDERY MILDEWS 251

Troximon parviflorum Nutt.......... Spherotheca humuli (DC.) Burr. var.
fuliginea (Schlecht.) Salm.

Ulmus fulva Michx..................Microsphera alni (Wallr.) Salm.

Neilimatrsichtt hve Witchy ei2)< chet syeie cccre)s- «ioe Uncinula macrospora Peck.

Vaccinium pennsylvanicum Lam.......Microsphera alni (Wallr.) Salm. var.
Vaccinii (Schw.) Salm.

WialerianawedtlishsN ltt. tassels asecesie eee Erysiphe cichoracearum DC.

WreeietsD ASEAEAS Pac ...o2 cosine aic)s so oles oe Erysiphe cichoracearum DC.

Metnena stricta: Vents, sii 6.53 jens « Erysiphe cichoracearum DC.

Merhena, utticefolia Le... 2... cee. eee Erysiphe cichoracearum DC.

Vernonia Baldwini Torrs............. Erysiphe cichoracearum DC.

Mibuniuimelentaco ile. .tccniceemoer Microsphera alni (Wallr.) Salm.

Wiciasamericana MithlSaanco5,. aoc -- Microsphera alni (Wallr.) Salm. var.
ludens Salm.

Witis (cultivated serape)) inn. ssn -eiece o<: Uncinula necator (Schw.) Burr.

Witt Saconditoltam Vinchaenenc ace acres artiel es Uncinula necator (Schw.) Burr.

Matic Oliipinad clos. co.css. consis cise dates. Uncinula necator (Schw.) Burr.

Zanthoxylum americanum Mill....... Phyllactinia corylea (Pers.) Karst.

University of Missouri
Columbia, Mo.

BIBLIOGRAPHY.

1. ANDERSON, J. P., 1907

Iowa Erysiphacee. Proc. Iowa Acad. Sci. 14 :1-34.
2. ANDERSON, F. W., 1889

Brief notes on a few common fungi of Montana. Journ. Mycol. 5:31.
3. ——_____, 1889

Preliminary list of the Erysiphee of Montana. Journ. Mycol. 5 :188-194.
4. ATKINSON, G. F., 1891

Some Erysiphee from Carolina and Alabama.

Journ. Elisha Mitch. Sci. Soc. 7 :61-73.
5. De Bary, A., 1863

Uber die Fruchtentwicklung der Ascomyceten. Leipzig.
6. ——__—__, 1871

Beitrage zur Morphologie und Physiologie der Pilze, XIII, XIV, pp.

23-75-, 91-85.
Abhandl, d. Senckenb. Naturf. Gesell. 7.

10.

Va ie

12.

13.

14.

15.

16.

18.

10.

20.

GEORGE M. REED

BERKELEY, M. J., 1847.
Gardener’s Chron. 7 :779.
BLACKMAN, V. H. & Fraser, H. C. I., 1905

Fertilization in Spherotheca. Ann. Bot. 19 :567-560.
Biopcett, F. M., 1913

Hop Mildew. Cornell Agr. Exp. Sta. Bult. 328.
Brannon, M. A., 1808

Some Indiana Mildews. Proc. Ind. Acad. Sci. pp 291-295.

Burritt, T. J. & Earze F. S., 1887
Parasitic Fungi of Illinois—Erysiphee.
Bult. Ill. State Lab. Nat. Hist. 2: pt. 2, 387-432.
Burriti, T. J., 1892
Erysiphee. Ellis & Everhart’s North American Pyrenomycetes,
pp 2-30, and 728.
De CaANnpoL_e, A. P., 1805-1815
Flore Francaise 2:272-275; 6:104-109.
Goone) Ms iGe"GPrex ©, He 1872
The Erysiphei of the United States. Journ. of Bot. (1872) :11-14,
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Couperc, G., 1893

Sur les périthéces de l’Uncinula spiralis en France et identification
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Comptes Rendus 116:210, 211.

DANGEARD, P. A., 1904
Recherches sur le développement du périthéce chez les Ascomycétes.
Le Botantiste, Ser. IX.
Davis, J. J., 1803
Supplementary List of Parasitic Fungi of Wisconsin.
Trans. Wis. Acad. Sci. Arts and Letters 9:156-158, 164, 165.
, 1897
Second Supplementary List of Parasitic Fungi of Wisconsin.
Trans. Wis. Acad. Sci. Arts and Letters 11 :165-178
, 1902
Third Supplementary List of Parasitic Fungi of Wisconsin.
Trans. Wis. Acad. Sci. Arts and Letters 14 :83-106.
, Sept. 1909
Fourth Supplementary List of Parasitic Fungi of Wisconsin.
Trans. Wis. Acad. Sciences, Arts and Letters 16 :739-772.

21.

22

23.

24.

25.

26.

27.

28.

20.

30.

ai;

22.

33.

34.

THE POWDERY MILDEWS 253

Eriksson, J., 1906

Der amerikanische Stachelbeermehltau in Europa, seine jetzige Ver-
breitung und der Kampf gegen ihn. Zeitsch. f. Pflanzenkr. 16 :83-90.

Fartow, W. G., 1874-76

List of Fungi found in the vicinity of Boston. Bull. Bussey Inst. 1 :437.

FREEMAN, E. M., Igoo.

A preliminary list of Minnesota Erysiphee. Minn. Bot. Stud. 2:423-430.

Fries, E., 1820.

Systema mycologium 3 :234-247.

GaLtoway, B. T., 1895

Observations on the development of Uncinula spiralis.
Bot. Gaz. 20 :486-401.

GriFFITHS, D., 1899

Some Northwestern Erysiphacee. Bull. Torr. Bot. Club 26:138-144.

Harkness, H. W. & Moore, J. P., 1880

Catalogue of the Pacific Coast Fungi. Calif. Acad Sci. :32, 33.

Harper, R. A., 1895

Die Entwickelung des Peritheciums bei Spherotheca Castagnei.
Berichte d. d. Bot. Gesell. 13 :475-481.

, 1806
Uber das Verhalten der Kerne bei der Fruchtentwickelung einiger
Ascomyceten. Jahr. f. w. Botanik 29 :656-686.
, 1905

Sexual Reproduction and the Organization of the Nucleus in Certain
Mildews. Carnegie Instit. of Washington, Publication 37 :1-104.

Keusey, F. D., 1889

Study of Montana Erysiphee. Bot. Gaz. 14 :285-288.

Lawrence, W. H., 1905

The powdery mildews of Washington. Wash. Agr. Exp. Sta. Bult. 7o.

LeveiL_e, J. H., 1851

Organisation et disposition methodique des espéces qui composent le
genre Erysiphe. Ann. Sci. Nat. III, 15:109-179, 381.

Macnus, P., 1808.

Der Mehlthau auf Syringa vulgaris in Nordamerika.
Bericht. d. d. Botan. Gesell. 16 :63-70.

254 GEORGE M. REED

35. MarcHat, E., 1902
De la specialization du parasitisme chez l’Erysiphe graminis.
Comptes Rendus 135 :210-212.
36. Marcuat, E., 1903
De la specialization du parasitisme chez l’Erysiphe graminis.
Comptes Rendus 136:1280, 1281.
37. Metuus, I. E., 1912
Culturing of parasitic fungi on the living host.
Phytopathology 2 :197-203.
38. MuritspaueH, C. F. & Nurratt, L. W., 1806
Flora of West Virginia.
Field Columbian Museum Publication 9, Vol. 1, nr. 2, 130, 131.
39. Necer, F. W., 1902
Beitrage zur Biologie der Erysipheen. Flora 90 :221-272.
LOS,
Neue Beobachtungen iiber das spontane Freiwerden der Erysipheen-

frucht korper.
Centrb. f. Bakt. Parasit. u. Infekt. Abt. 2, 10:570-573.

41. PAMMELL, L., 1909

Some plant diseases of 1908. Iowa Agr. Exp. Sta. Bult. 104.
42. Persoon, C. H., 1796.

Observationes mycologice I :13.

43. ——_——_—, 1801
Synopsis Methodica Fungorum 1 :124.
44. REBENTISH, J. F., 1804
Prodromus Flore Neomarchice, 360, 361.

45. Reep, G. M., 1905

Infection Experiments with Erysiphe graminis DC .
Trans. Wis. Acad. Sci. Arts and Letters 15 :135-162.

46. ———______., 1907

Infection experiments with the mildew on cucurbits, Erysiphe
cichoracearum DC.

Trans. Wis. Acad. Sci., Arts and Letters 15 :527-547.

47. ————,, 1908

Infection experiments with Erysiphe cichoracearum DC.
Bull. of the University of Wisconsin, Science Series 3 :337-416.

48.

49.

50.

5I.

52.

57:

58.

59.

61.

THE POWDERY MILDEWS 255

EEE Tee
The mildews of the cereals. Bull. Torr. Bot. Club 36:353-388.
, 1912
Infection experiments with the powdery mildew of wheat.
Phytopathology 2 :81-87.
Satmon, E. S., 1900
A Monograph of the Erysiphacee. Mem. Torr. Bot. Club. 9.
, 1902
Supplementary notes on the Erysiphaceze.
Bull. Torr. Bot. Club 29:1, 83, 181, 302.
s 1903
On specialization of parasitism in the Erysiphacee.
Beihefte Zum. Bot. Centralblatt 14 :261-315.
, 1903
Infection powers of ascospores in Erysiphacee. Journ. of Bot. 41:1!
, 1904
Cultural experiments with biologic forms of the Erysiphacee.

Phil. Trans. Royal Society of London Series B, 197 :107-122.

» 1904
Cultural experiments with the barley mildew, Erysiphe graminis DC.
Annal. Mycol. 2:70-99.
s 1904
On Erysiphe graminis DC., and its adaptive parasitism within the
genus Bromus. Annal. Mycol. 2:255-267, 307-343.
» 1905 .
Further cultural experiments with ‘Biologic Forms’ of the Erysiphacee.
Ann. of Bot. 19:125-148.
» 1905
On the present aspect of the epidemic of the American gooseberry
mildew in Europe. Journ Roy. Hort. Soc. 29 :102-I10.
, 1906
On Ojidiopsis taurica Lév., an endophytic member of the Erysiphacez.
Ann. Bot. 20:187-200.
pE ScHweEINiTz, L. D., 1834

Synopsis Fungorum in America Boreali.
Trans. Amer. Phil. Soc. 4:269, 270.
Seitpy, A. D., 1893

The Ohio Erysiphee. Ohio Agr. Exp. Sta. Bult. 3 :213-224.

256 GEORGE M. REED

62. SMITH, GRANT, 1900
The haustoria of the Erysiphez.
Bot. Gaz. 29 :153-184.
63. STEINER, J. A., 1908
Die Specialisation der Alchemillenbewohnenden Sph@rotheca Humuli
(DC.) Burr.
Centrallblatt fiir Bakt., Parasitenkunde, und Infextious Krankheiten,
Abt. II, 21 :677-736.

$4. Stewart, F. W., 1910
Notes on New York Plant Diseases I. N. Y. Agr. Exp. Sta. Bult. 328

4&5. Tracy, S. M. & Earte, F. S., 1895
Mississippi Fungi. Miss. Agr. Exp. Sta. Bult. 34 :95-07.

66. Tu asne, L. R. & C., 1861
Selecta Fungorum carpologia I :191-216.
67. Unperwoop, L. M. & Earte, F. S., 1897
Preliminary list of Alabama Fungi.
Ala. Agr. Exp. Sta. Bult. 80:176-181.
68. VocLtno, P., 1905
Contribuzione allo studio della Phyllactinia corylea.
Nuovo Giornale Botanico Italiano 12:313-327.
69. WatirotH, F. W., 1819
Naturgeschichte des Mucor Erysiphe L.
Berl. Ges. Nat. Freunde Verhandl. 1 :6-45.
70. ————————__, 1819
De mucore Erysiphe Linnaei observationes.
Annal. Wett. Gesell. 4:226-247.
71. Watters, L. A., 1806
Erysiphee of Riley County, Kansas.
Trans. Kansas Acad. Sci. 14:200-206.
72. WInNGE, O., IQII
Encore le Spherotheca Castagnei Lév.
Bull. Soc. Mycol. de France 27 :211-2109.

EXPLANATION OF FIGURES

Figures 1-3 are copied from Smith. Figures 6, 7, 9-33 are copied from
Harper. The others are from drawings made by G. T. Kline.

Prate Derr:

Fig. 1. Haustorium of Erysiphe polygoni DC. on Geranium maculatum.

Fig. 2. Haustoria of Uncinula salicis (DC.) Winter on Salix discolor.
One haustorium is located in the epidermal cell, while the other
is in a subepidermal cell.

Fig.
Fig.

Fig.

Fig.

Fig.

Fig.

THE POWDERY MILDEWS 257

Haustorium of Erysiphe graminis DC. on Poa.

Three conidiophores of Erysiphe graminis DC. The characteristic

enlargement at the base of the conidiophores is well shown.
(Magn. 228 times).

Three germinating conidia of Erysiphe graminis DC. (Magn. 228

times).

Section through a young perithecium of Phyllactinia corylea

(Pers.) Karst. The ascogonium of five cells is shown; the penul-

timate cell is budding out in ascogenous hyphe.

Median section of an older perithecium of Phyllactinia corylea.

A portion of the ascogonium and sections of the multinucleated

ascogenous hyphe are shown.

Median section of a perithecium of Uncinula salicis. The section

shows three asci, each with the primary ascus nucleus; surround-

ing the asci is the inner wall composed of cells filled with dense

protoplasm; outside of these the outer wall whose cells are largely

devoid of protoplasmic contents and have thick walls. Four young

appendages are shown.

PLATE XIV.

Figs. 9-20 Spherotheca castagnei Lév.
Fig. 9. Oogonium and antheridial branch each containing one nucleus.

Fig.

Figs.

Fig.

Fig.
Fig.

Fig.
Fig.

Fig.

Fig.

Fig. 20.

10.

Antheridial branch cut off and nucleus divided into two.

11-12 Antheridium cut off from the stalk cell.

rz

14.

ry.

16.
17:

18.

19.

Cell wall between antheridium and oogonium dissolved and the
antheridial nucleus lying adjacent to the oogonial nucleus in the
oogonium.

Fusion of oogonial and antheridial nuclei; wall re-formed between
antheridium and oogonium.

Oogonium with fusion nucleus and surrounded by the first layer of
wall cells.

Oogonium surrounded by two layers of wall cells.

Development of oogonium to form ascogonium. The latter now
consists of two uninucleated cells.

Young ascogonium with three nuclei, cell division not having oc-
curred.

Completely developed ascogonium with the cells of the inner layer
of the perithecial wall shown. The penultimate cell of the ascogo-
nium is binucleated and is the young ascus.

Young ascus with the primary ascus nucleus and two ascogonium
cells.

Figs. 21-24. Phyllactinia corylea (Pers.) Karst.

Fig. 21.

Ascus with two nuclei; chromatin strands distinctly oriented with
reference to the central body.

258

GEORGE M. REED

Figs. 22-24. Stages in the fusion of the two nuclei to form the primary ascus

nucleus.

PLATE XV.

Figs. 25-30. Phyllactinia corylea (Pers.) Karst.

Fig.
Fig.
Fig.

Fig.
Fig.
Fig.
Figs.

Fig.

Fig.
Fig.

25.
26.
27.

28.

20.
30.

Ascus with primary ascus nucleus.

Division of primary ascus nucleus; early stage in synapsis.
Division of primary ascus nucleus; loosening of chromatin fol-
lowing synapsis.

Division of primary ascus nucleus; equatorial plate stage showing
eight chromosomes, and the central bodies with the well developed
asters.

Binucleated ascus following completion of the first division.

Second division; late anaphase stages with eighi chromosomes on
each half of each spindle.

31-32 Erysiphe cichoracearum DC.

31.

a2.

30a.

Spore formation; plasma membrane being formed by bending
back of astral rays and their fusion to form an umbrella-shaped
membrane.

Plasma membrane of spore about complete.

Phyllactinia corylea (Pers.) Karst. Ascus with two spores.

PLATE XVI.

Perithecium of Podosphera oxyacanthe (DC.) De Bary, on cul-
tivated cherry. (Magn. 50 times).

. Ascus of P oxyacanthe. (Magn. 275 times).

Perithecium of Phyllactinia corylea (Pers.) Karst. on Betula alba.
(Magn. 50 times).

. Ascus of P. corylea (magn. 275 times).

Perithtecium of Erysiphe polygoni DC. on Polygonum aviculare.
(Magn. 50 times).

. Ascus of E. polygoni. (Magn. 275 times).

Perithecium of Uncinula salicis (DC.) Winter on Salix discolor.
(Magn. 50 times).

. Ascus of U. salicis. (Magn. 275 times).

Perithecium of Spherotheca Humuli (DC.) Burr., var. fuliginea
(Schlecht.) Salm. on Taraxacum officinale. (Magn. 50 times).

. Ascus of S. Humuli var. fuliginea. (Magn. 275 times).

Perithecium of Microsphera alni (Wallr.) Salm. on Syringa vul-
garis. (Magn. 50 times).
Ascus of M. alni. (Magn.'275 times).

PRATE ally

PLate XIV

PLATE XV

iV

PLATE XVI

A DESCRIPTIVE LIST OF THE CEPHALINE GREGAR-
INES OF THE NEW WORLD
By Max M. ELtis.

Introduction.

Aside from Central Europe little is known of the Cephaline
gregarine fauna of the world. Like many other groups of animals
which have no large economic importance at present, cephaline
gregarines have been neglected. Particularly is this true of the
New World fauna. Gregarines are however deserving of more
attention if only as objects of scientific interest, since they are
easily obtainable and offer excellent material for both class work
and experimental research. It is the object of the present paper
to bring together the references to the new world species and
some information regarding the group Cephalina Delage. Short
descriptions and measurements have been added to aid in the
determination of specimens.

History.

Although gregarines had been seen and reported by several
writers before 1828, the first formal description of a genus and
species of cephaline gregarines was made in that year by Leon
Dufour (1828). He established the genus Gregarina and defined
G. ovata from Forficula auricularia L., an earwig. He had how-
ever discussed gregarines in an earlier paper (1826). Dufour
considered gregarines as peculiar worms and this idea dominated
the work of several subsequent authors. Gregarines were va-
riously regarded as parasitic worms, both Nematode and Trema-
tode, and were even assigned to the plant kingdom by some. Koel-
liker (1848), was one of the first zoologists to maintain that gre-
garines are one-celled animals. The taxonomic knowledge of
cephaline gregarines has been advanced particularly through the
works of A. Schneider (1875 et. seq.), and of L. Leger (1892 et.
seq. ).

The first paper dealing with new world gregarines was pub-
lished by Joseph Leidy in 1849. This included a short diagnosis
of the genus Gregarina and the description of a new species, G.

260 MAX M. ELLIS

larvata from Julus marginatus. In his discussion Leidy states that
“in the state in which Gregarina is found, it would probably hold
rank between the J7vematoda and Trichina, the lowest of the
Nematoidea”. Subsequently Leidy published several other notes
and descriptions of gregarines. No further attention was given
the new world forms until Frenzel (1892), gave the descriptions
and the results of some experimental work upon five new species
of gregarines from Argentine Republic. Crawley (1903a,b; 1907),
in a series of three articles offered the first connected account of
the cephaline gregarines of the United States, describing new spe-
cies from Leidy’s unpublished manuscript and several from his
own investigations.

At the present time 56 species, some of which are incom-
pletely described, are known from North and South America. This
undoubtedly is but an introduction to our fauna.

Anatomy and Life Cycle.

Cephaline gregarines are protozoan parasites, usually found
in the alimentary canal of arthropods. They are most commonly
taken in the free adult stage, the sporont, from the posterior por-
tion of the alimentary canal of the host. A sporont (fig. 13),
typically consists of two parts separated by a septum, so that it
superficially appears to be made up of two cells. The anterior of
these units is the protomerite, and the posterior, containing the
nucleus, the deutomerite. The septum between the protomerite
and the deutomerite is usually well developed but in some genera,
as Gamocystis A. Schneider, the septum is wanting, the protomerite
being represented by but a constriction. The other extreme is
found in the peculiar gregarine Taeniocystis mira Leger from
Ceratopogon solstitialis Winn., in which the deutomerite is divided
by several granular septa. The outside coat of a sporont, the
epicyte, is quite thin and rather firm as contrasted with the semi-
fluid material, the endocyte, which fills it. Often just below the
epicyte a thin clear zone, the sarcocyte, may be seen. The endocyte
is usually quite dense and homogeneous although it may be almost
clear, and may contain large and small granules and oil drops. The
nucleus is suspended in this endocyte unattached as may be demon-
strated by crushing a sporont, with pressure from a cover glass,

————— Oe

<

CEPHALINE GREGARINES OF THE NEW WORLD 261

without crushing the nucleus, when the nucleus will be forced out
of the epicyte. Often the nucleus is not visible but it may always
be made so by staining with a weak solution of Iodine in Potassium
Iodid (the usual Grams solution diluted one-half with water
answers very well for this).

Since the life cycle of all cephaline gregarines is much the
same, although those of the various species differ in detail, Gre-
garina blattarum Siebold from cockroach will be used as an ex-
ample. The fusion of two sporonts in the alimentary canal of the
first host produces a cyst. This is a prolate spheroid covered with
a gelatinous envelope. After being discharged from the alimentary
canal of the first host the cyst, if it be kept moist, passes through
a series of internal changes, which result in the formation of sporo-
cysts, commonly called spores. These are discharged from the
cyst through long tubes, the sporoducts. The period during which
the cyst produces sporocysts, up to the time when they are dis-
charged, is known as the maturation period. The maturation
changes, although begun before the cyst has left the body of the
first host, rarely if ever are completed in the first host.* Each
sporocyst, after it has been subjected to the proper conditions of
moisture, discharges several sporozoites. Infection of the second
host may take place either as the result of the ingestion of sporo-
zoites, or of the sporocysts from which the sporozoites may be
discharged. In either event the intracellular phase of the life
cycle is begun by the sporozoite. This enters a cell of the alimen-
tary canal of the new host and after a time develops into a minute
gregarine, composed of three parts, a deutomerite, a protomerite
and an epimerite, the latter being in front of the protomerite and
joined to it. Later the young gregarine withdraws from the
cell of the host so that only a portion of the epimerite remains
within the cell, the remainder of the body being in the alimentary
canal of the host. When the gregarine has attained a certain
growth it leaves the cell entirely and becomes a free parasite in
the canal, although still bearing the epimerite. This stage is the

*Crawley, (1903b, p. 641) has suggested, from the advanced stage of maturation in
which he found the cysts of G. achetaeabbreviatae in the host, that sporocysts my occasion-
ally be discharged in the host. Leger et Duboscq (1902, p. 412), report the sporocysts of
Pyxnia mobuszi as occuring in the excrement of the host.

262 MAX M. ELLIS

cephalont. The loss of the epimerite constitutes the change to the
sporont stage and the cycle is completed.* The various stages
differ somewhat with the several families and will be discussed in
the diagnoses of the families.

Technique.

Gregarines are best studied while living. The alimentary canal
may be withdrawn from a recently decapitated arthropod and
teased in normal salt solution. Many species of gregarines will
live for hours in either normal salt solution or Ringer’s solution.
When placed in water the osmotic tension usually causes them to
swell up and burst. This difficulty may be overcome by the addi-
tion of a little white of egg. Permanent mounts are usually made
with considerable difficulty if attempt be made to handle the in-
dividual animals. Balsam mounts may be made however by kill-
ing and staining portions of the alimentary canal of the host and
teasing them when in balsam. Sections of the alimentary canal
are also good, showing the intracellular stages as well as the free
forms. The cysts are to be collected from the faeces of the host.
By isolating several individuals of the host species in clean test tubes
plugged with cotton, the faeces may be collected free from debris.
The faeces should be examined with a low power glass after soak-
ing in water. The cysts when removed from the faeces should be
placed in a damp-cell on a slide. Care must be used to protect
the cysts from mold.

Taxonomy.*

Since the sporonts of many species are quite similar the
taxonomic characters are drawn for the most part from the epi-
merite, the cyst and the sporocyst. Often all of the stages were
not at hand, so that many species are incompletely described, in
the original diagnoses. In the descriptions given here the letters
“P” and “D” refer to protomerite and deutomerite respectively,
and the measurements are for average sporonts or spores.

*For a detailed account of the intracellular stages and their development see Leger
and Duboscq, 1904.

*In making up the brief descriptions of the species given, data from specimens
seen by the writer were used as far as possible; these wanting, the descriptions were
composed from the figures given with the original diagnoses, which in the case of gre-

garines must correspond in general to the types of larger animals. The figures given
here are from drawings by the writer unless otherwise credited.

CEPHALINE GREGARINES OF THE NEW WORLD 263

CEPHALINA DELAGE

Sporozoans reproducing by sporulation only, which usually follows the
permanent fusion of two adult individuals; gametes similar or dissim-
ilar; young stages always intracellular; epimerite present in the first
extracellular stage or at least represented in the last intracellular stages;
adult generally divided by a septum into a deutomerite and a protomerite.
This group includes the Polycystid Gregarines of authors.

Key To THE FAMILIES OF CEPHALINA
A, With a free cephalont stage.
B. Sporonts forming associations ;* epimerite simple.
C. Septum of the satellite disappearing; dehiscence by simple rup-

TEURES, sichaikh sens ctairetomis se ayeiiaia as aaa ae Cae es Didymophide

CC. Septum of the satellite present.
D. Dehiscence of cyst by simple rupture........ Hyalosporide
DD. Dehiscence of cyst by sporoducts.............. Gregarinide

BB. Sporonts not forming associations; epimerite usually not simple.
E. Septum present in the sporont.
F. Dehiscence of cyst by simple rupture.
G. Sporocysts without spines...... eens Actinocephalide
GG, sSporocyst with sspies:. 5.62. 4oSiec8.: « Acanthosporide
FF, Dehiscence of cyst by a pseudocyst, either central or lateral.
H. Epimerite symmetrical and symmetrically attached to

Pe Se OLGERETALE.. 6554. \ataetlas «uno Stylocephalide

HH. Epimerite asymetrical or asymmetrically attached to the
PROLOMERILE Ris oct cecre ek, Re eh eee Dactylophoride

2S, | Septum wanting in. the sporontin;.. i. os. ne sidewos od os Doliocystide

AA. Epimerite represented only in the last intracellular stages.Stenophoride

*Two or more sporonts joined in tandem, see fig. 5; the first of these sporonts is
termed the primite and the posterior individuals the satellites.

DiIpDYMOPHIDZ

A family of one genus, the species of which are known only

from Europe.
Didymophes F. Stein, 1848, s. 186.
Type—D. gigantea F. Stein, 1848, s. 186, t. 9, f. 40; from Oryctes
nasicornis (L.) larvae,—Coleoptera.

HYALOSPORID

This family is in part the Gregarinidae of authors. As here
defined it includes only those Gregarinids whose cysts dehisce by
simple rupture. Seven genera, one provisionally, are referred to

264 MAX M. ELLIS

this family. One new world species is known, although doubtless
several occur.

a. Septum present, protomerite and deutomerite distinct.
b. Sporocysts ellipsoidal to spindle-shaped; ends somewhat pointed.
c. Sporocyst with distinct equatorial swelling........ Frenzelina
cc. Sporocyst without equatorial swelling............ Hyalospora
bb. Sporocysts not ellipsoidal; ends broadly rounded.
d. Sporocysts not spherical.

e. Sporocyst ovoid or cylindrical............. Eirmocystis
ee. Sporocyst prismatic, polygonal in outline...... Euspora
dd. Sporocyst spherical or subspherical........... Uradiophora
aa. Septum wanting.
fA POSDPFORE SIODOSE. 255.5) 2 eta iieio edo ee oe RO Spherocystis
tic spotont *clongatests a. sce eh een ea ee eee Ganymedes

Frenzelina Leger et Duboscq, 1907, p. 773-774.
Type—F. conformis (Diesing)=Gregarina conformis Diesing, 1851, II,
p. 15; from Pachygrapsus marmoratus (F.),—Crustacea.
Hyalospora A. Schneider, 1875, 4, p. 583.
Type—H. roscoviana A. Schneider, 1875, 4, p. 584, t.16,f.41-42; from
Petrobius maritimus—Thysanura.
Euspora A. Schneider, 1875, 4, p. 582.

Type—E. fallax A. Schneider, 1875, 4, p. 583, t.18,f.14-17; from Rhizo-

trogus @stivus,—Coleoptera.

Euspora lucani Crawley. Fig. 1.

Euspora lucani Crawley, r903a, p. 50-51, pl. I11,f.38; Swarthmore, Penn-

sylvania, from Lucanus dama Thunb.—Coleoptera.

Epimerite undescribed; elongate and cylindrical, protomerite and deu-
tomerite both broadly rounded; size as given by Crawley, lc., primite 520ux
128u, satellite 360ux108u; cysts unknown. This species is referred to the
genus Euspora because of the shape of the sporont and the coleopteran host,
making the generic determination very uncertain.

Eirmocystis Leger, 1892, p.110.
Type—E. ventricosa Leger,1892, p. 111, t.6,f.1-4; from Tipula oleracea
and Tipula pratensis larve,—Diptera.
Uradiophora Mercier, 1912, p. 108.
Type—U. cuenoti (Mercier)=—Cephaloidophora cuenoti Mercier, IgII, p.
51; from Atyephyra desmaresti Millet,—Crustacea.
Ganymedes J. Huxley, 1910, p. 160.

Type—G. anaspidis J. Huxley, roro, p. 155-175, pl. 11, f. 1-19; from

Anaspides tasmanie (Thompson) ,—Crustacea.

This genus is placed here provisionally because of the similarity of
Ganymedes anaspidis and Uradiophora cuenoti in several features of morph-
ology and in type of host. Since the complete life cycle of Ganymedes has
not been worked out this arrangement cannot be verified at present.

CEPHALINE GREGARINES OF THE NEW WORLD 265

Spherocystis Leger, 1892, p. 115.
Type—S. simplex Leger, 1892, p. 115, t. 6, f. 11-13; from Cyphon pallidus
larve,—Coleoptera.

GREGARINID

Cysts spherical or ovoid, covered by a gelatinous envelope
which is often double; one or more sporoducts forming during
maturation, through which the sporocysts are discharged; sporocysts
often in chains. As here defined this family includes but part of
the species of Gregarinidac of authors, the other genera being re-
ferred to Hyalosporidac. Two of the three genera are represented
in the new world fauna.

a. Septum present; protomerite and deutomerite distinct.

DP SPOEGGHCES, SQUCE AM crak cles 2 aie a Siar creca entail 5 sacee ole Gregarina
Hip SPOMOGUCH TONGL BALM ce oor a Selassie Glcias toa mace Gigaductus
ieee SCPE ats WAIEINGS een re ce 3 Paws yeep an siaye sisieln Kote ote Gamocystis

Gregarina Dufour, 1828, p. 366.

Type—G. ovata Dufour, 1828, p. 366; from Forficula auricularia L.,—

Orthoptera (Euplexoptera).

Of the twenty species from the new world assigned to the this genus the
generic determination of but six is absolute, since the dehiscence of the cysts
has not been described for the other fourteen. It has been the custom of
authors to refer any gregarine found in association to this genus when the
data were insufficient for complete determination; hence a large number of
species have been placed here tentatively.

Gregarina blattarum Siebold. Figs. 20-22.

Gregarina blattarum Siebold, 1839, s.57,t.3: Crawley, 1903a, p. 44; from

Periplaneta orientalis and Ectobia germanica: Hall, 1907, p. 1; Lincoln,

Nebraska, from Periplaneta americana: Ellis, r9r3c, p. 83; Douglas

Lake, Michigan from Ischnoptera pennsylvanica.

Gregarina blatte-orientalis Leidy, 1853, 230, pl. 11, f.11-12; from Blatta

orientalis.

Clepsidrina blattarum, Magalhes, 1900, p. 38-44; Brazil, from Periplaneta

americana 2nd Periplaneta orientalis.

Epimerite short, digitiform to subglobose, about one-half the length of
the protomerite of the cephalont; sporont short and broad, both pro-
tomerite and deutomerite broadly rounded; average P. 1ooux120p, D. 130ux
400“; cysts prolate spheroids, average 450ux900n with gelatinous envelope ;
sporoducts 10 or more, reaching the length of 200”; sporocysts barrel-shape,
4ux8u.

Gregarina panchlore Frenzel. Fig. 9.

Gregarina panchlore Frenzel, 1892, s .209, f. 20; Cordoba, Argentine Re-

public, from Panclilora exoleta Klug.

266 MAX M. ELLIS

Epimerite undescribed; sporont cylindrical, length 180u, width 30-35;
cysts and sporozoites unknown. In associations the protomerite of the
satellite is deeply concave at the anterior end to receive the posterior end of
the deutomerite of the primite. From the measurements given by Frenzel,
a completion of his figure 20 would make the deutomerite about six times
as long as the protomerite. This gregarine may be a synonym of the fol-
lowing species. Specimens of Panchiora sp. from Quirigua, Guatemala, and
from bananas shipped into Boulder, Colorado, were examined by the writer
in 1912 but no gregarines found.

Gregarina blabere Frenelzel. Fig. 2.
Gregarina blabere Frenzel, 1892, s. 300-314, f.21-33; Cordoba, Argentine
Republic, from Blabera claraziana and related forms.

Epimerite long, tapering, shaped like a spear-head, enlarged at the base,
about twice the length of the protomerite of a large cephalont; sporont
elongate, protomerite and deutomerite both broadly rounded; adult sporont
I50ux500u, protomerite about one-fourth the length of the deutomerite;
cysts and sporozoites unknown. A specimen of Blabera sp. from Gualan,
Guatemala contained no gregarines.

Gregarina serpentula Magalhes. Fig. 3.

Gregarina serpentula Magalhes, 1900, p. 40, {.4; Brazil, from Periplaneta

americana.

Epimerite unknown; sporont elongate, 180u4x1200u, protomerite 50u in
length; cysts undescribed. From the figures given by Magalhzs and Frenzel
it seems quite probable that G. serpentula is a synonym of G. blabere, leaving
but two species of gregarines known from the roaches of the world at pres-
ent.

Gregarina achete-abbreviate Leidy. Fig. 5.

Gregarina achete-abbreviate Leidy, in part, 1853, p. 238, pl.11,f.34: Craw-

ley, 1903a, p. 45, pl.III, f. 35; Beach Haven, New Jersey, from Acheta

abbreviata: idem, 1903b, p. 639-641; idem, 1907, p. 220, pl. XVIII, f. 1;

Beach Haven, New Jersey, and Wyncote, Pennsylvania, from Gryllus

abbreviatus.

Epimerite undescribed; sporont short and broad, protomerite almost
spherical, deutomerite rounded posteriorly; average P. 200ux150u, D. 225ux
3004; cysts spherical, about 2504, with a gelatinous envelope; sporoducts 2
to 5, elongate, reaching the length 1ooou, (Crawley, 1907, lc.), sporocysts
cylindrical, tapering slightly at each end, ends broadly rounded, 4.5ux2.3u.
Taken by the writer at Douglas Lake, Michigan, July, 1913, from Gryllus
americanus.

Gregarina longiducta Ellis. Figs. 26-29.

Gregarina longiducta Ellis, 1913c, p. 78-82, f. 1-8; Douglas Lake, Mich-

igan, from Ceuthophilus latens and Ceuthophilus maculatus.

Epimerite short and digitiform, about equalling the protomerite of a
cephalont in length; sporont short and broad; average P. 200ux170n, D. 200ux

CEPHALINE GREGARINES OF THE NEW WORLD 207

230u; cysts spherical with a gelatinous envelope, 200u to 300"; sporoducts
four or rarely five, at one pole, length when everted enormous, reaching
35004; sporocysts barrel-shaped, hexagonal in profile, with rounded edges,
3ux6.5u. A species much like G. achete-abbreviate from which it differs in
the size of the sporocysts, the enormously long sporoducts, and the polar
arrangement of the sporoducts.

Gregarina consobrina sp.. nov. Figs. 23-25.

Epimerite short, simple and digitiform, its length about one-third that
of the protomerite of the cephalont; sporont short and globose; protomerite
hemispherical, not as wide as the deutomerite; length of the protomerite
amout one-half of its width and one-fifth of the total length; deutomerite
broadly oval in outline, its maximum width about equalling its length; cysts
spherical, with a thick, outer, gelatinous envelope and a thin, dense, inner en-
velope, average cysts 250u to 300“; sporoducts four to six in number, all in
one hemisphere, very long, averaging 900u to 12004 in length; sporocysts
cylindrical, slightly rounded at each end, in chains when first discharged,
3.2ux8u; maturation period in water at room temperature during October, six
days or more; average sporonts 600u in length, P. 130ux300u, D.
470ux450u; host, Ceuthophilus valgus Scudder, (det. Prof. T. D. A. Cock-
erell), collected in Boulder Canon, near Boulder, Colorado, at an altitude of
about 6,500 feet, October 5, 1913.

This species, G. achete-abbreviate Leidy, and G. longiducta Ellis are to
be regarded as a species group since they are so closely related yet each
presents a different combination of characters. G. consobrina Ellis differs
from G. longiducta Ellis in the position of the sporoducts, these being all in
one hemisphere although not closely grouped about the pole; in the length of
the sporoducts, which are about one-third as long as those of the latter; and
in the shape of the sporont, this being much more globose and the protomerite
less distinct. From G. achete-abbreviate Leidy G. consobrina differs in
shape of sporont, size of sporocyst, lack of orange color in cyst, as well as
type of host.

Gregarina rigida (Hall). Fig. 13.

Hirmocystis rigida Hall, 1907, p.1-26, f. 1-11, 21; Lincoln, Nebraska, from

Melanoplus differentialis, M. femur-rubrum, M. atlantis, Canon City,

Colorado, from M. bivittatus, M. differentialis, M. angustipennis: Hall

I9QI2, p. 337; Canon City, Colorado, from M. coloradensis, Boulder, Colo-

rado, Colorado Springs, Colorado, Bethesda, Maryland: Ellis, ror3a,

p. 464; Boulder, Colorado, from Brachystola magna.

Gregarina melanopli Crawley, 1907, p. 223, pl. XVIII, f. 6-90; Wyncote,

Pennsylvania from Melanoplus femoratus: Ellis, 1913c, p. 82-83; Douglas

Lake, Michigan, from Melanoplus luridus, M. femur-rubrum; M. bivit-

tatus.

Epimerite short and digitiform; sporont short and broad, both pro-
tomerite and deutomerite rounded, average P. 130uxrIsou, D. 140ux
570u; cysts spherical, 300% to 400u, covered by a gelatinous envelope,

268 MAX M. ELLIS

20u to 2004, usually orange in color; sporoducts 10 or more, exceeding the
gelatinous envelope but a short distance; sporocysts in chains when first
discharged, hyaline, barrel-shaped, rather hexagonal in outline, 5ux8u; both
cysts and sporonts are usually yellow or even orange in color and the sporo-
duct-buds a brilliant orange just before the sporoducts are everted. This is
the common gregarine of North American grasshoppers. Some little con-
fusion concerning the name of this species has arisen as the result of the
almost simultaneous publication of the descriptions of Hirmocystis rigida
Hall and Gregarina melanopi Crawley, here considered as synomymous. The
original diagnoses of both species were without descriptions of the cysts and
their dehiscence. Hall pointed out (1912, p. 337) that the two species were
to be regarded as synonyms. The writer, (1913c) described the cysts and
their dehiscence for G. melanopi Crawley from material collected at Doug-
las Lake, Michigan, and since returning to Colorado has found the cysts of
Hirmocystis rigida Hall to dehisce by sporoducts in the same manner; hence
the name must stand Gregarina rigida (Hall).

Gregarina locuste-caroline Leidy. Fig. to.
Gregarina locuste-caroline Leidy, in part, 1853, p. 230, pl. 11, f. 35-38;
Locusta carolina L.: Crawley, 1907, p. 225, pl. XVIII, f. 13; from
Dissosteira carolina (L.), Wyncote, Pennsylvania.
Stephanophora locuste-caroline, Crawley, 1903a, p. 54, in part.

Epimerite globose, about half the length of the protomerite of the
cephalont; sporont short and rounded, protomerite subglobose, deutomerite
oval; largest individual seen (Crawley, 1907, p. 225), 350"; cysts and dehis-
cence undescribed.

Gregarina passalicornuti Leidy. Figs. 12 and 16.

Gregarina passalicornuti Leidy, 1853, p. 238, pl. 11, Fig. 30-31: Ellis,
1913b; New Orleans, Louisiana, from Passalus cornutus Fab.

Epimerite undescribed; sporont distinctly longer than broad, rather
cylindrical in outline, protomerite hemispherical, deutomerite cylindrical,
usually narrowed near the middle; average P. 60uxson, D. 60ux
150u; cysts and sporocysts unknown.

Gregarina guatemalensis Ellis. Fig. 15.

Gregarina guatemalensis Ellis, 1912c, p. 687, Fig. 6; Quirigua, Guate-

mala, from Nelus interstitialis

Epimerite undescribed; sporont short and broad, especially in the pos-
terior portion of the deutomerite; protomerite subglobose; deutomerite
cylindrical, widening rather abruptly near its posterior end; average P. 70ux
Sou, D. 1604x180“; cysts and dehiscence undescribed.

Gregarina xylopini Crawley. Fig. 17.

Gregarina xylopini Crawley, in part, 1903a, p. 47, pl. III, f. 30; from

Xylopinus saperdioides.

Epimerite undescribed; sporont somewhat elongate; protomerite elon-
gate, distinctly narrowed near the middle, its length twice its width; pro-

—

CEPHALINE GREGARINES OF THE NEW WORLD 269

tomerite cylindrical, its width about one-third its length; cysts and dehiscence
undescribed; size not given in original diagnosis.
Gregarina grisea Ellis. Fig. 18.

Gregarina grisea Ellis, 1913b, p. 200, f. 1; New Orleans, Louisiana, from
Tenebrio castaneus Knoch.

Epimerite undescribed; sporont short and ovoid; protomerite hemispher-
ical, narrower than the deutomerite; deutomerite oval, its posterior margin
broadly rounded; average P. 60uxs5ou, D. 1ooux 370.

Gregarina microcephala Leidy.

Gregarina microcephala Leidy, r889, p. 11, 1 Fig.; from Hoplocephala

bicornis.

This is known only from the original diagnosis. Since its position is
very uncertain Leidy’s description is copied here.

“In some little green beetles, Hoplocephala bicornis, one of the Tene-

brionide, I found a number of gregarines remarkable for the small size

of the head and hence the species may be named Gregarina microcephala.

The body is clavate; the head like a watch crystal with a little ball at

the summit. Length 0.35 mm. by 0.1 wide; head 0.012 long by 0.04 wide.

It bears a close resembldpce to Echinocephalus hispidus of Schneider,

found in Lithobius forcipatus, but in the one described I at no time found

digitiform appendages to the head.”

The host of this species is now known as Arrhenoplita bicornis (Olivier).

Gregarina scarabeirelicti Leidy.

Gregarina sp. Leidy, 1851a, p. 208; from the larve of a large lamellicorn

insect.

Gregarina scarabeirelicti Leidy, 1851b, p. 287; from larve of Scarabeus

relictus.

This species and the following one, G. melalonthebrunnee Leidy, are
known only from the original diagnoses, which are incomplete and without
figures. Until these species are redescribed their position and validity are
doubtful. Leidy’s diagnoses are copied here. :

“Body white, cylindro-fusiform. Superior division presenting four sides

of a hexagon, subacute. Nuclear body of inferior division transparent,

globular or elliptical, containing several coarse granules. Length from

1-66th to 1.25 lines; head 1-400th inch to 1-133d inch long by 1-285th inch

to I-111th inch broad. Anterior portion of inferior division 1-200th inch

to 1-86th inch broad; posterior portion 1-666th to 1-250th inch broad. —”.
Gregarina melalonthebrunnee Leidy.

Gregarina melalonthebrunnee Leidy, 1856, p. 47;-from Melalontha-

brunnea.

“Body oblong oval; head oblate spheroidal, slightly elevated at the sum-

mit. Single and in pairs. Length of body .405 mm., breadth .252 mm.;

length of head .108 mm., breadth .144 mm.”

Gregarina statire Frenzel. Fig. 14.
Gregarina statire Frenzel, 1892, p. 234-286, t. VIII, f. 1-15; Cordoba, Ar-

270 MAX M. ELLIS

gentine Republic, from Statira unicolor Blanch.

Epimerite short, simple, conic; cephalonts and free sporonts ovoid;
sporonts in association globose; protomerite hemispherical to subglobose, its
length about one-fourth of the total length, width of the protomerite less
than that of the deutomerite, in large sporonts about one-half the width of
the deutomerite; protomerite of satellite quite compressed; cysts and sporo-
cysts unknown; large sporonts 3004x350u.

Gregarina bergi Frenzel. Figs. 38-39.

Gregarina bergi Frenzel, 1892, p. 286-208, f. 16-19; Cordoba, Argentine

Republic, from Corynetes ruficollis.

Epimerite simple, styliform, enlarged near the base so that it is arrow-
head-shaped in profile, its length greater than that of the protomerite of the
cephalont, its greatest width about one-half that of the protomerite; sporonts
ovoid; protomerite hemispherical, almost as wide as the deutomerite, length
of the protomerite about one-fourth of the total length; posterior margin
of the deutomerite broadly rounded; cysts and sporocysts unknown; aver-
age individuals 90ux300u. This gregarine has been taken by Wellmer, 1912,
in Prussia from Corynetes violaceus L. He reports it as forming associa-

tions. :
Gregarina elatere Crawley. Fig. 10-11.

Gregarina elatere Crawley, 1903a, p. 46, pl. I, f.11; Wyncote, Pennsyl-

vania, from Elater sp. larve.

Hirmocystis ovalis Crawley, 1903a, p. 50, pl. I, f. 5-6; from larve of

beetles, doubtfully identified as Cucujide. :

Epimerite globose to ovoid, almost equalling the length of the protomer-
ite of the cephalont in diameter; cephalont ovoid; sporont rather cylindrical,
both protomerite and deutomerite broadly rounded; protomerite hemispherical
about one-fourth as long as the deutomerite; deutomerite cylindrical, a little
broader at its junction with the protomerite than the protomerite; cysts and
sporocysts undescribed; no associations observed; maximum length as given
by Crawley, 7ou.

Gregarina termitis Leidy. Fig. 6.

Gregarina termitis Leidy, 1881, p. 441, pl. 52, f. 27: Porter, 1897, p. 65,

pl. 6, f. 73-76; Cambridge, Mass. from Termes flavipes.

Epimerite undescribed; sporont short, distinctly longer than broad, pro-
tomerite oval to subglobose, deutomerite ovoid to cylindrical; average P.
25ux170u, D. 30ux 400u, cysts and sporocysts unknown. The writer has
taken this species at Boulder, Colorado from Termes lucifugus during 1912
and 10913.

Gregarina calverti Crawley. Fig. 4.

Gregarina calverti Crawley, 1903a, p. 48, pl. Il, f. 19-21; Wyncote, Penn-

sylvania, from Lysiopetalum lactarium; idem. 1903b, p. 638, pl. XXX, f.15.

Epimerite undescribed; sporont elongate and cylindrical, protomerite
short, oval in outline, about one-twentieth as long as the deutomerite, some-
what more globose in young sporonts equalling about one-sixth of the length

CEPHALINE GREGARINES OF THE NEW WORLD 271

of the deutomerite, deutomerite elongate and cylindrical, tapering posteriorly
in young sporonts; cysts spherical, about 3004 in diameter; sporocysts barrel-
shaped, 5uxI3u; average sporonts 1000.

Gregarina sp.

Gregarina sp. Ritter, Proc., Cal. Acad. Sci., ser. 2, 4, p. 39-85, 1893. This
description was not seen by the writer.

Gigaductus Crawley, 1903a, p. 633.

Type—G. parvus Crawley, 1903a, p. 633, pl. XXX, f. 10-13; from Har-

palus caliginosus Fab.,—Coleoptera.

Gigaductus parvus Crawley. Fig. 8.

Gigaductus parvus Crawley, 1903a, p. 633, pl. XXX, f. 10-13; Wyncote,

Pennsylvania fram Harpalus caliginosus Fab.: Ellis, 1913a, p. 465; Vin-

cennes, Indiana, from Harpalus pennsylvanicus Dej.

Epimerite undescribed; sporont longer than wide though not greatly
elongate, oval in outline with a distinct constriction at the junction of the
protomerite and deutomerite; protomerite subglobose; deutomerite ovoid,
tapering noticeably toward the posterior end; average P. 7oux45u, D.
S8oux160u; cysts spherical, about 200u in diameter, dehiscence by one, large,
short sporoduct; sporocysts cylindrical, 25uxI2y.

Gigaductus kingi (Crawley). Fig. 7.

Gregarina kingi Crawley, 1907, p. 221, pl. XVIII, f. 10-12; from Gryllus

abbreviatus Serv.

Epimerite undescribed; sporont longer than wide; protomerite of primite
knob-shaped, widest in its anterior half, deeply constricted near the middle;
protomerite of the satellite subglobose; deutomerite oval in outline; average
P. 6oux4ou, D 60ux120u; cysts spherical or oval, about 100” in
diameter, dehiscence by one large, rather long sporoduct; sporocysts barrel-
shaped, 3ux5u.

Gamocystis A. Schneider, 1875, p. 587.

Type—G. tenax A. Schneider, 1875, p. 587, t. 19, f. 10-13, t.21,f.6; from

Ectobia lapponica (L.),—Orthoptera.

This genus is without a known representative in our fauna at present.

ACTINOCEPHALID

Dehiscence of cysts by simple rupture; sporocysts biconic or
navicular to crescentic; epimerite variable; sporonts not forming
associations. As here defined this family includes both the Actino-
cephalide and Menosporide of Leger. The epimerite becomes high-
ly specialized in some species of this family, yet the entire gamut
of possibilities is run from the simple to the extremely elaborate.
Three types are represented if typical species be chosen: (1) epi-
merite simple and styliform—Stylocystis; (2) styliform with a cir-
cular, elevated and divided, basal portion—Py-inia; (3) epimerite

272

MAX M. ELLIS

consisting of a circular elevated and divided portion, with a cen-
tral concavity, suggestive of the disappearance of the styliform por-
tion of the other two types—Menospora. An effort to divide the
family according to these three types of epimerite is unsatisfac-
tory, however, since the various combinations of these epimerite
characters, as regards presence and absence, and degree, intergrade.
A key to the genera, although perhaps somewhat artificial, is pos-
sible on epimerite characters.

a.

Protomerite regular, not divided.
b. Epimerite simple.

c. Epimerite rounded, -hemispherical; protomerite of the
cephalont much compressed and elevated around the base
of the epimerite like a collar............... Amphoroides

cc. Epimerite styliform.
d. Epimerite at first short and styliform, but becoming
rounded and button-shaped as the cephalont develops
Sharh acaabte eats Khoi aie Be rates co is PION? hol Ra Steinina
dd. Epimerite not becoming button-shaped.
e. Epimerite simple styliform, often curved. Stylocystis
ee. Epimerite conical, arrowhead-shaped in profile....
Biers Bich nt Mee eh Seek SN, bees at Pileocephalus
bb. Epimerite not simple.
f. Carried by a much produced portion of the protomerite.
g. With retrose spine-like processes; styliform to subglo-
HOSEMGi lsc cick eG eae ee eee Geniorhynchus
gg. Without retrose spine-like processes.
h. Apical portion with digitiform processes.
i. Apex concave, with a marginal row of re-

curved processes.................-Menospora
iii Apex convex, with six to eight marginal
digitiform processes...... Hoplorhynchus

hh. Without apical digitiform processes; a rounded
marginal portion in the center of which is an
elevated cup-shaped portion with a scalloped edge;
central portion evertible............... Phialoides

ff. Anterior portion of the protomerite of the cephalont slight-
ly if at all produced.

j. Septum wanting; epimerite disk-shaped to subglobose,
its margin scalloped deeply............... Schneideria

jj. Septum present, protomerite and deutomerite distinct.
k. Epimerite consisting of a central elevated portion
surrounded at its base by a marginal elevated or

divided portion.

a

se

CEPHALINE GREGARINES OF THE NEW WORLD 273

1. Central portion rounded, hemispherical, mar-
ginal portion rounded and undivided......

Hin Gea e? DU reL Re pr ae Discorhynchus

ll. Central portion pointed and _ styliform.
m. Basal portion scalloped........... Pyxinia
mm. Basal portion subglobose, produced into
horizontal or slightly recurved teeth....
Pe ee ee ter eens Beloides
kk. Epimerite without a central elevated portion.
n. Deutomerite not divided by septa.
o. Epimerite short, with a series of long
hair-like filaments............ Bothriopsis
Een BLS eee eth ees eee Coleorhynchus
ee rates tees See seis ena sane Legeria
oo. Epimerite without long filaments but with
short digitiform processes.

p. Basal portion of the epimerite longer
than the digitiform processes, cylin-
drical’to- flask-shaped. Jo2.'.30-2 ses
Bese craset a Sates = Amphorcephalus

pp. Basal portion equal to or shorter than
the digitiform processes.

q. Digitiform processes free and
well separated ...Actinocephalus
qq. Digitiform processes placed close
together, more or less united at
the hase 2 Stephanophora
ooo. Epimerite without long filaments, consist-
ing of button-shaped or subglobose mass
deeply fluted.
r. Basal portion of the lobes rounded..

SE Sas 5 fase cers ee Anthorhynchus
rr. Basal portion of the lobes pointed and
LECURVEd Bete we ae a Stictospora

nn. Deutomerite divided by several septa; epimerite
subglobose with recurved hooks............
SRS A pai Bo gat ih ge a Teniocystis
aa. Protomerite produced and divided equatorially so that the whole
has somewhat the appearamce of half-raised umbrella; epimerite
consisting of a circular series of short digitiform processes car-
ried on a narrowed portion of the protomerite....Sciadiophora
Amphoroides Labbe, 1899, p. 20.
Amphorella Leger, 1892, p. 132. Preoccupied.

274 MAX M. ELLIS

Type—A. polydesmi (Leger)=Amphorella polydesmi Leger, 1892, p.
132, t.10,f.9-14; from Polydesmus complanatus (L.),—Diplopoda.
Amphoroides polydesmivirginiensis (Leidy). Fig. 36.

Gregarina polydesmivirginiensis Leidy, 1853, p. 238, pl. 10, £.23-29;

Amphoroides polydesmivirginiensis, Crawley, 1903a, p. 45,pl.II,f.25;

Wyncote, Pennsylvania and Raleigh, North Carolina, from Polydesmus

virginiensts.

Epimerite undescribed; protomerite button-shaped to subglobose, small,
and narrower than the deutomerite, greatest length of the protomerite not
exceeding one-tenth of the length of the deutomerite; deutomerite elon-
gate, rounded posteriorly, widened in the anterior half; epicyte thick; cysts
and sporocysts unknown; average sporonts 400.

Amphoroides fontarie Crawley. Fig. 37.

Amphoroides fontarie Crawley, 1903a, p. 53, pl. I, f£.12-14; Wyncote,

Pennsylvania, and Raleigh, North Carolina, from Polydesmus sp. and

Fontaria sp.

Epimerite undescribed; sporont somewhat ovoid in shape, protomerite
subglobose, its maximum width less than that of the deutomerite, its length
one-fourth or less of the length of the deutomerite; deutomerite oval in
outline, often widened in its anterior half; average sporonts about 170;
sporocysts and cysts unknown. The writer has taken this species from
specimens of Polydesmus sp. collected by Mr. S. A. Rohwer at East Falls
Church, Virginia, in May, 1913.

Steinina Leger et Duboscq, 1904, p. 352.

Type—S. ovalis (Stein)=Stylorhynchus ovalis Stein, 1848, p. 182-223;

from Tenebrio molitor L. larve—Coleoptera.

This species, S. ovalis (Stein), or others of the same genus, should be
looked for in North America since the host and other closely related species
are found in our fauna.

Stylocystis Leger, 1899, p. 526.
Type—S. precox Leger, 1899, p. 526-533; from Tanypus sp. larve—
Coleoptera.
Stylocystis ensiferus (Ellis). Fig. 34.

Stylocephalus ensiferus Ellis, 1912c, p. 686, £.5; Quirigua, Guatemala,

from Leptochirus edax Sharp.

Epimerite simple and styliform, its length about equal to that of the
protomerite of the cephalont; sporont ovoid, the deutomerite broadly
rounded posteriorly, protomerite subglobose, deutomerite cylindrical; aver-
age sporonts 50“; cysts and sporocysts unknown.

Pileocephalus A. Schneider, 1875, p. 591.
Type—P. chinensis A. Schneider, 1875, p. 592, t.16,f.21-24; from Mysta-
cides sp. larve—Trichoptera.

Geniorhynchus A. Schneider, 1875, p. 594.
Type—G. monnieri A. Schneider, 1875, p. 595, t.20,f.21-27; from
Libellula sp. nymphs—Odonata.

CEPHALINE GREGARINES OF THE NEW WORLD 275

Geniorhynchus .eshne Crawley. Fig. 41.

Geniorhynchus eshne Crawley, 1907, p. 227, pl. XVIII, f.4; Southeastern

Pennsylvania, from nymphs of Aeshna constricta Say.

Epimerite subglobose, carried by an elongated portion of the protomer-
its, with numerous short, spine-like processes directed posteriorly; both
protomerite and deutomerite resembling truncated cones with their bases
together; deutomerite according to Crawley often constricted posteriorly;
size given as 4204; cysts and sporocysts not known.

Menospora Leger, 1892, p. I5I.
Type—M. polyacantha Leger, 1892, p. 151, t.19,f.1-5; from Agrion puella
(L.) nymphs—QOdonata.
Hoplorhynchus Carus, 1863, p. 570.
Type—H. oligacanthus (Siebold)=Gregarina oligacantha Siebold, 1839,
t.3; from Calopteryx virgo (L.)—Odonata.
Phialoides Labbe, 1899, p. 24.

Phialis Leger, 1892, p. 135. Preoccupied.

Type—P. ornata (Leger)=Phialis ornata Leger, 1892, p. 135,t.13,f.4-12;

from Hydrophilus piceus (L.) larve—Coleoptera.

Schneideria Leger, 1892, p. 153.
Type—S. mucronata Leger, 1892, p. 153, t.2,{.7-13; from Bibio marci
(L.) larve—Diptera.

Discorhynchus Labbe, 1899, p. 20.

Discocephalus Leger, 1892, p. 134. Preoccupied.

Type—D. truncatus (Leger)=Discocephalus truncatus Leger, 1892, p.

134, t.15,f.10-12; from Sericostoma sp. larve—Trichoptera.

Pyxinia Hammerschmidt, 1838 p. 35.

Asterophora Leger, 1892, p. 129.

Type—P. rubecula Hammerschmidt, 1838, p. 357, t.4,f. a-g; from Der-

mestes lardarius L.—Coleoptera.

Pyxinia crystalligera Frenzel. Fig. 43-44.

Pysxinia crysatlligera Frenzel, 1892, p. 314-332, f. 34-50; Cordoba, Argen-

tine Republic, from Dermestes vulpinus Fab. and Dermestes peruvianus

Castelnau, and larve of the latter.

Epimerite consisting of a circular basal portion with a fluted margin and
a central styliform portion, the length of the styliform portion exceeding
one-half the length of the protomerite of the cephalont; sporont somewhat
elongate, protomerite globose, narrower than the widest portion of the deu-
tomerite, deutomerite broad just posterior to the protomerite; average
sporonts QO0ux250p.

Beloides Labbe, 1899, p. 26.

Xiphorhynchus Leger, 1892, p. 137. Preoccupied.

Type—B. firmus (Leger)=Xiphorhynchus firmus Leger, 1892, p. 138, t.17,

f.1-4; from Dermestes lardarius L.—Coleoptera.

Bothriopsis A. Schneider, 1875, p. 596.

276 MAX M. ELLIS

Type—B. histrio A. Schneider, 1875, p. 596, pl. XXI, £.8-13; from Hydati-

cus cinereus,—Coleoptera.

Bothriopsis histrio A. Schneider. Fig. 42.

Bothriopsis histrio Schneider, 1875, p. 506, pl. XXI, £.8-13; Crawley,
19034, p. 54-55, pl. II, f.15-18; Wyncote, Pennsylvania, from Hydaticus ciner-

eus larve, Colymbetes fuscus and Acilius sulcatus.

Epimerite consisting of a short button-shaped portion from the margin
of which are six or more long hair-like filaments; protomerite of the cepha-
lont subglobose anteriorly and cylindrical posteriorly; deutomerite of cepha-
lont ovoid; sporont variable and very active changing shape readily, in ex-
panded individuals the protomerite is subglobose with a cup-shaped depres-
sion posteriorly into which the conical anterior end of the deutomerite fits,
deutomerite, aside from the portion included by the protomerite, elongate and
conical; sporonts reach the length of 500”; cysts spherical, about 400m in
diameter, dehiscing by simple rupture; sporocysts biconic, 54x7u.

Coleorhynchus Labbe, 1899, p. 23.

Coleophora A. Schneider, 1885, p. 94. Preoccupied.

Type—C. heros (A. Schneider)=Coleophora heros A Schneider, 1885,

p. 95, t.25; from Nepa cinerea L.—Hemiptera.

Although the type of epimerite for this genus has not been described it
is placed in the key with Bothriopsis because of the aquatic host.

Legeria Labbe, 1899, p. 24.

Dufouria A. Schneider, 1875, p. 595. Preoccupied.

Type—L. agilis (A. Schneider)=Dufouria agilis A. Schneider, 1875, p.

595, t.22, f.1-6; from Colymbetes sp. larve—Coleoptera.

Legeria terpsichorella sp. nov. Fig. 30.

Epimerite not seen; sporonts extremely active constantly changing the
shape of the anterior three-fifths of the body and proceeding rather rapidly
in a serpentine path as a result, the protomerite often being bent almost
forty-five degrees from the main axis of the body; expanded individual with
a protomerite equal to or longer than the deutomerite, the anterior fourth of
the protomerite hemispherical to subglobose, below which is an elevated
flange-like portion, remaining two-thirds cylindrical, the posterior portion
with a cup-shaped depression some 60u deep into which the anterior conical
portion of the deutomerite fits; deutomerite excepting the portion included
by the protomerite ovoid, rather sharply rounded posteriorly; average
sporonts about 720u in length; length of the deutomerite to the external
junction with protomerite 320u, of the anterior conical portion of the deu-
tomerite 96u, of the protomerite to the flange portion 3204, from flange to
anterior end Sou; width of deutomerite 1454, of the flange portion of the
protomerite 1754; epicyte thin and flexible; sarcocyte scarcely visible; nu-
cleus seen only with the use of reagents; endocyte dense and homogeneous,
of a light brown color; cysts and sporocysts not seen.

Host, Hydrophilus sp., Douglas Lake, Michigan, July, 1913.

Amphorellus Ellis, r913a, p. 462.

CEPHALINE GREGARINES OF THE NEW WORLD 277

Type—A. amphorellus Ellis, 1913a, p. 463, f. 1-2; from Scolopendra heros
Girard—Chilopoda.

Amphorocephalus amphorellus Ellis. Figs. 51-52
Amphorocephalus amphorellus Ellis, 1913a, p. 463, £. 1-2; Boulder, Colo-
rado, from Scolopendra heros Girard.

Epimerite flask-shaped with a marginal row of small digitiform processes
at its anterior end, its length greater than that of the protomerite of the
cephalont; protomerite with a constriction near the middle; deutomerite of
the cephalont elongate and conical, broadest near its anterior end, where
its maximum width is twice that of the protomerite; deutomerite of the
sporont elongate and cylindrical, rather sharply and abruptly pointed at
its posterior end; sporonts reaching the length of 1,000”; P. 6o0uxs5op, D.
60ux950u; cysts unknown.

Amphorocephalus actinotus (Leidy). Fig. 53.

Gregarina actinota Leidy, 1889, p. 10, f.1; from Scolopocryptops

Sexspinosus.

Hoplorhynchus actinotus, Crawley, 1903a, p. 55, pl. III, £.36-37; Wyncote,

Pennsylvania, Raleigh, North Carolina, and Wallingford, Pennsylvania,

from Scolopocryptops sp.

Hoplorhynchus scolopendras Crawley, 1903b, p. 636, pl. XXX, f.19; Ral-

eigh, N. C. from Scolopendra woodi Meiner.

Epimerite elongate, flask-shaped, bearing at its anterior end a series of
small digitiform processes carried by four horizontal lobes, length of the
epimerite equal to from one-half to one fourth of the total length of the
cephalont; protomerite hemispherical to subglobose; deutomerite elongate,
conical and pointed posteriorly, its maximum width about one-third of its
length; size, as given by Leidy 600u for the cephalont, by Crawley, 485u for
the sporont; cysts unknown.

Actinocephalus F. Stein, 1848, p. 106.

Stephanophora Leger, 1892, p. 127. .

Type—A. lucani F. Stein, 1848, t. 9, f. 33; from Lucanid beetle.

It is to be noted that Stephanophora Leger, was invalid since it in-
cluded the single species Actinocephalus lucani Stein (redescribed by Leger
as Stephanophora radiosa Leger), the type of Stein’s genus Actinocephalus.
Leger, 1892, recognized the synonomy of Actinocephalus lucani Stein with
his species Stephanophora radiosa, but by its removal from the genus of
Stein (wrongly ascribed to Schneider by Leger, 1892, p. 141), the genus
Actinocephalus Stein was left without a species included in its original des-
cription. This situation renders the name Actinocephalus as restricted by
Leger, 1. c., and Labbe, (1899, p. 25), invalid. In restoring the name Ac-
tinocephalus to its type A.‘lucani Stein, Actinocephalus of authors stands
without a name. Since Stephanophora Leger and Actinocephalus of authors
are so closely related it seems best to consider them synonomous, avoiding

278 MAX M. ELLIS

the confusion attendant to the substitution of a new name. Both epimerite
and sporocyst characters of Stephanophora and Actinocephalus intergrade.
Actinocephalus pachydermus (Crawley). Figs. 54-55.

Stephanophora pachyderma Crawley, 1907, p. 226, pl. XVIII, f. 2-3;

Wyncote, Pennsylvania, from Dissosteira carolina (L.)

Gregarina locuste-caroling Leidy, in part, 1853, p. 230, pl. 11, f. 37-38.

Stephanophora locuste-caroline, Crawley, in part, 1903a, p. 54.

Epimerite subglobose, bearing at its apex a marginal row of digitiform
processes; protomerite somewhat hemispherical with the epicyte slightly
produced to receive the epimerite; deutomerite of the cephalont elongate but
rather broad; sporont oval in outline, protomerite short and hemispherical,
its length about one-fourth of the total length; epicyte very thick in both
cephalont and sporont; sporonts reaching the length of 500%; cysts unknown.
It seems quite probable that the undescribed gregarine figured by Hall,
(1907, f. 13) from Chimarocephalus viridifasciata taken at Lincoln, Ne-
braska, was a sporont of this species.

Actinocephalus zophus (Ellis) Fig. 49.

Stephanophora zopha Ellis, 1913b, p. 201, f. 2; New Orleans, Louisiana,

from Nyctobates barbata Knoch.

Gregarina xylopini Crawley, in part, 1903a, p. 47, f. 29; from Xylopinus

saperdioides.

Epimerite short and subcylindrical, with an apical row of marginal
digitiform processes; protomerite subglobose, its diameter equal to or a
little greater than the length of the epimerite; deutomerite elongate, cylin-
drical, pointed posteriorly; sporont elongate and cylindrical, pointed pos-
teriorly, length of. the protomerite 8 to 12 in the length of the deutomer-
ite; sporonts reaching length of 1600”; cysts unknown. The writer has
taken this gregarine from specimens of Alobates pennsylvanicus deGeer
collected at East Falls Church, Virginia in May, 1913 by Mr. S. A. Rohwer.

Actinocephalus crassus (Ellis). Fig. 4o.

Stephanophora crassa Ellis, rorac, p. 688, f. 7; Quirigua, Guatemala,

from Leptochirus edax Sharp.

Known only from the sporont; general shape ovoid with the posterior
portion of the deutomerite narrowed and conical; protomerite hemispheri-
cal, its length equal to about one-third of the total length; deutomerite
broad in the anterior half, narrowed rather abruptly in the posterior half to
a rounded cone; epicyte thick.

Actinocephalus harpali (Crawley). Fig. 46.

Gregarina harpali Crawley, 1903a, p. 49-50, pl. I, f. 1-4; Wyncote, Penn-

sylvania from Harpalus caliginosus.

Actinocephalus harpali, Crawley, 1903b, p. 637-638, pl. XXX, f. 14.

Epimerite undescribed; sporont ovoid; protomerite hemispherical its
length about one-sixth of the total length and about one-half of its own
width; deutomerite ovoid; sporonts reaching the length of 12004. Cysts
about 500u in diameter, covered with a thick gelatinous envelope, dehiscing

CEPHALINE GREGARINES OF THE NEW WORLD 279

by simple rupture; sporocysts 7.54 x Qu, described by Crawley as, “diamond
shaped in longitudinal and hexagonal in transverse section.”
Actinocephalus disceli (Crawley). Fig. 50.

Gregarina disceli Crawley, 1903a, p. 47, pl. I, f. 7-10; Wyncote, Penn-

sylvania, from Discelus ovalis.

Epimerite undescribed; sporont elongate, posterior end of the deutomer-
ite tapering and pointed; protomerite pentagonal in outline, as wide or slight-
ly wider than the deutomerite, its length about one-thirteenth of the total
length in large sporonts; cysts and sporocysts unknown. This gregarine is
placed in the genus Actinocephalus because of the general shape of the
sporont and the coleopteran host; it was removed from the genus Gregarina
since the sporonts do not form associations. The grouping of large numbers
of sporonts with the posterior ends of their deutomerites touching can not be
considered an association in the sense of a Gregarinid association, and has
also been observed for species of Stylocephalus and Actinocephalus.

Actinocephalus dujardini A. Schneider. Fig. 45.

Actinocephalus dujardini A. Schneider, 1875, p. 580, pl. 16, f. 9-20;

Crawley, 1903a, p. 55; from Lithobius forcipatus.

Epimerite subglobose with a short neck, bearing a marginal row of
about twenty short, rigid, recurved, tooth-like processes at its anterior end;
protomerite subglobose to cuboidal; its length equal to half or more of the
length of the deutomerite; deutomerite rather broad, conical; size small.

Actinocephalus americanus Crawley, Fig. 56.

Actinocephalus americanus Crawley, 1903b, p. 636, pl. XXX, f. 22;

Wyncote, Pennsylvania, from Galerita bicolor Drury.

This gregarine was described from a single specimen, 2004 x 454, pro-
tomerite 354 long. Crawley states that it is probably “only sporadically
present in Galerita, and that its usual host is some other animal.” There
exists but this single record of this species.

Actinocephalus brachydactylus sp. nov. Figs. 31-33.

Epimerite very short, composed of a circular row of eight short digiti-
form processes united basally; protomerite globose to dome-shaped, in the
cephalonts slightly broader than the deutomerite; deutomerite subcylindrical,
tapering gradually towards the posterior end which is broadly rounded; av-
erage cephalont 3204 in length, protomerite S80u x S8ou, deutomerite
75u X 2404; sporonts reaching the length of 500u; cysts not seen. Host,
nymphs of Aeshna sp., Douglas Lake, Michigan. Taken July, 1913.

Anthorhynchus Labbe, 1899, p. 19.

Anthocephalus A. Schneider, 1887, p. 69. Preoccupied.

Type—A. sophie (A. Schneider) = Anthocephalus sophie A. Schnei-

der, 1887, p. 69, t. 10, f. 11-17; from Phalangium opilio L—Phalangidea.

Anthorhynchus cratoparis (Crawley). Fig. 47.
Asterophora cratoparis Crawley, 1903a, p. 54, pl. II, f. 23; Swarthmore,
Pennsylvania, from Cratoparis lunatus.

280 MAX M. ELLIS

Epimerite spherical, deeply fluted, borne by a short elevation from the
anterior portion of the protomerite; protomerite subglobose; deutomerite in
the form of a truncated cone, somewhat elongate and narrowed at the an-
terior end where it joins the protomerite; length given as 540u; cysts undes-
cribed.

Anthorhynchus philicus (Leidy). Fig. 48.

Gregarina philica Leidy, 1889, p. 9, 1f.; from Nyctobates pennsylvanica.

Asterophora philica, Crawley, in part, r903a, p. 53, pl. III, f. 31-32.

Epimerite spherical, deeply fluted; protomerite subglobose to cuboidal,
about one-ninth the length of the deutomerite; deutomerite elongate, sub-
cylindrical, tapering to a point at the posterior end; length as given by
Crawley, 300“; cysts undescribed.

Anthorhynchus boletophagi (Crawley). Fig. 57.

Gregarina boletophagi Crawley, 1903a, p. 47, pl. I, f. 26-28; Swarth-

more, Pennsylvania, from Boletophagus cornutus.

Epimerite undescribed; sporont subcylindrical, protomerite oval in out-
line with a short dome-shaped portion at the antefior end, length of the pro-
tomerite a little more than one-fourth of the total length, deutomerite regu-
larly cylindrical excepting the extreme posterior end which tapers rather
abruptly so as to form a truncated cone. This species has been transferred
to this genus from Gregarina although neither cysts nor epimerite are
known, because it is not found in association, and because the anterior por-
tion of the protomerite is suggestive of the slightly produced protomerites
of other species of the genus Anthorhynchus, which bear the epimerites. It
is to be regarded as a provisional determination only.

Stictospora Leger, 1893, p. 117.
Type—S. provincialis Leger, 1893, p. 129-131; from Melolontha sp. larve
—Coleoptera.

Teniocystis Leger, 1906, p. 307.
Type— T. mira Leger, 1906, p. 307-329; from Ceratopogon solstitialis
Winn., larve—Diptera.

Sciadiophora Labbe, 1899, p. 17.

Lycosella Leger, 1896, p. 36. Preoccupied.

Type—S. phalangii (Leger) = Lycosella phalangii Leger, 1896, p. 36,

t. 3, f. 1-15; from Phalangium crassum Duf.—Phalangidea.

The writer has opened the alimentary canal of perhaps two hundred
Phalangidea from Douglas Lake, Michigan, and from Boulder, Colorado,
without finding any gregarine infection, although S. phalangii (Leger) and
related species are reported as very abundant in the Phalangidea of Europe.

ACANTHOSPORIDE

Dehiscence of cysts by simple rupture; sporocysts with spines ;
sporonts always solitary. No species referable to this family have

CEPHALINE GREGARINES OF THE NEW WORLD 281

been taken as yet in the new world. The genera may be separated
by the following key:

a. Epimerite without lateral recurved processes or long filaments;
spines at both the equator and the poles of the sporocysts. Acan-
thospora.

aa. Epimerite with either lateral recurved processes or long filaments.
b. Epimerite with lateral recurved processes.

c. Sporocysts with spines only at the poles........... Corycella

cc. Sporocysts with spines at both the equator and poles. An-
cyrophora

bb. Epimerite with long filaments; sporocysts with spines both at

the poles and above and below the equator....... Cometoides

Acanthospora Leger, 1892, p. 145.
Type—A. pileata Leger, 1892, p. 145, t. 15, f. 1-5; from Omoplus sp.
larve—Coleoptera.
Corycella Leger, 1892, p. 144.
Type—C. armata Leger, 1892, p. 144, t. 16, f.7-12; from Gyrinus natator
—Coleoptera.
Ancyrophora Leger, 1892, p. 146.
Type—A. gracilis Leger, 1892, p. 146, t. 19, f. 11-13; from Carabus
auratus L.,—Coleoptera.
Cometoides Labbe, 1899, p. 29.
Pogonites Leger, 1892, p. 148. Preoccupied.
Type—C. crinitus (Leger) = Pogonites crinitus Leger, 1892, p. 149, t.
18; from Hydrobius sp. larve—Coleoptera.

STYLOCEPHALID

Dehiscence of cysts by simple rupture with a pseudocyst;
sporonts solitary; sporocysts subspherical but asymmetrical, united
in chains usually black or dark brown; sporulation distinctly.
anisogamic. The species of this family are known only from
Tenebrionid beetles*

a. Epimerite cup-shaped, composed of a row of short digitiform pro-

cesses surrounding a membranous portion........ Lophocephalus
aa. Epimerite without digitiform processes.
b. Epimerite large and conical, carried by a short neck............

ere a nee a ecig) ae eee eee Cystocephalus

bb. Epimerite small, carried by a long base.
c. Epimerite spherical or ovoid.............-. Spherorhynchus
cc. Epimerite cylindrical and pointed, with a bulbous basal
Pee 3 ee, ee i a ge a Ee Stylocephalus

Lophocephalus Labbe, 1899, p. 31.

rhe species Stylocephalus caudatus (Réssler) is probably referable to the genus
Stictospora of the Actinocephalidae. This species is from a Phalangid host.

282 MAX M. ELLIS

Lophorhynchus A. Schneider, 1882, p. 435. Preoccupied.

Type—L. insignis (A. Schneider) = Lophorhynchus insignis A. Schnei-

der, 1882, p. 435, t. 13, f. 1-3, 5, 12, 13, 48, 50; from Helops striatus

Fourc—Coleoptera.

Cystocephalus A. Schneider, 1886, p. 99.

Oocephalus A. Schneider, 1886, p. 101.

Type—C. algerianus A. Schneider, 1886, p. 100, t. 27, from Pimelia sp.—

Coleoptera.

Spherorhynchus Labbe, 1899, p. 32.

Spherocephalus A. Schneider, 1886, p. 100, Preoccupied.

Type—S. ophioides (A. Schneider) = Spherocephalus ophioides A.

Schneider, 7886, p. 100, t. 28; from Acis sp.—Coleoptera.

Stylocephalus Ellis, 1912, p. 25.

Stylorhynchus Stein, 1848, p. 195. Preoccupied.

Type—S. giganteus Ellis, 1912, p. 25-27, f. 1-2; from Eleodes sp.—Cole-

optera.

Stylocephalus giganteus Ellis. Figs. 58-59.

Stylocephalus giganteus Ellis, 1912a, p. 25-27, f. 1-2; Boulder, Colorado,

from Eleodes sp.: Hall, 1912, p. 337-338; Amo, Colorado, from Eleodes

hispilabris and Eleodes sp.

Epimerite rather styliform, basal bulbous portion less than half as long
as the distal cylindrical portion; collar joining the epimerite to the pro-
tomerite almost, if not quite as long, as the epimerite proper and exceeding
it in diameter; epimerite and collar exceeding the length of the protomerite
of the cephalont; sporont greatly elongate, sub-cylindrical, pointed poster-
iorly; sporonts exceeding 2,000u. To the diagnosis of this species as or-
iginally given may be added the description of the cysts and sporocysts,
which have recently been secured.

Cysts spherical, average diameter 450u, the entire surface irregular cov-
ered with small elevations and depressions, cyst proper covered with a very
thin gelatinous envelope (entirely wanting in some cysts), white when first
discharged from the body of the host, but becoming lead gray and finally
black as maturation progresses; maturation period for cysts obtained in
September, 1913, and kept in water at room temperature, at least ten days;
dehiscence simple rupture, with an irregularly spherical central pseudocyst;
sporocysts discharged in long chains; each sporocyst subspherical but asym-
metrical, one side being distinctly larger and with a greater curvature than
the other; when in the chains the sporocysts alternate so that the large
side of a sporocyst is always turned away from the large sides of the two
adjoining sporocysts; covering of the sporocyst thick, expanded at each end
to join with that of the next sporocyst in forming the chains; endosporal
mass arranged around a polygonal, hyaline, central spot containing a few
granules; sporocysts black or dark brown in color, measuring 74 xX IIp;
sporozoites differentiating in a few days from the endosporal mass, leaving a
central hyaline space with numerous granules.

CEPHALINE GREGARINES OF THE NEW WORLD 283

The writer has taken this gregarine from Eleodes sp, and Asida sp., col-
lected at Denver, Colorado, also from Asida opaca Say and Eusattus sp. at
Boulder, Colorado.

DACTYLOPHORIDE

Epimerite asymmetrical, or asymmetrically placed on the
protomerite; cysts dehiscing by simple rupture, usually splitting
along the equator, with a pseudocyst; sporocysts cylindrical.

a. Protomerite represented only by a constricted portion of the body;

MRR SU AMAB Se otra cig we ck bys aa Se te eee Rhopalonia

aa. Protomerite distinct from the deutomerite; septum present.
b. Deutomerite not divided by septa.
c. Sporocysts cylindrical, usually in chains.

d. Sporont short and ovoid; epimerite asymmetrical, con-
sisting of a conical pointed lateral portion and a mar-
ginal row of filamentous processes; the conical por-
tion persisting in the sporont stage...... Echinomera

dd. Sporont elongate; epimerite conical, short and lateral;
protomerite broad, upturned on one side, that bearing
the epimerite; protomerite with digitiform processes.

e. Protomerite bifid on the side away from the epi-
BIGGIE £2 565 hea cid & 24 He take mA « Pterocephalus
ec. - Protomerite, not . bifid: .:.....5.: 2.502035 Dactylophorus
cc. Sporocysts more or less ellipsoidal; not in chains.

f. Sporocysts not pointed; epimerite short, coni-
cal and lateral, borne by a much produced por-
tion of the protomerite...... Trichorhynchus

fi. SDOLOCYStS <- pointed... + s-u- se eeke eee Acutispora

bb. Deutomerite divided by one or more granular septa. ..Metamera
Rhopalonia Leger, 1893, p. 1285.
Type—R. geophili Leger, 1893, p. 1285-1288; from Geophilus sp.—Chilo-
poda.
Echinomera Labbe, 1899, p. 16.

Echinocephalus A. Schneider, 1875, p. 593. | Preoccupied.

Type—E. hispida (A. Schneider) = Echinocephalus hispidus A. Schnei-

der, 1875, p. 503, t. 16, f. 36-40; from Lithobius forticatus L—Chilopoda.

Echinomera hispida (A. Schneider). Fig. 60.

Echinocephalus hispidus A. Schneider, 1875, p. 593, t. 16, £. 36-40.

Echinomera hispida. Crawley, 1903a, p. 52; Wyncote, Pennsylvania,

Raleigh, North Carolina, and Cambridge, Mass., from Lithobius forti-

catus: Ellis, 1973a, p. 465; Boulder, Colorado, from Lithobius coloraden-

sis (Cockerell).

Epimerite asymmetrical, consisting of a pointed conical, lateral portion,
and a series of more or less filamentous digitiform processes, the whole be-

284 MAX M. ELLIS

ing carried by a short base equalling the protomerite in width; the pro-
cesses of the epimerite disappearing shortly after the animal frees itself
from the intestinal wall of the host, but the conical portion of the epimerite
persists in the sporont stage giving an asymmetrical margin to the front of
the protomerite; sporont ovoid, length of the protomerite from one-seventh
to one-eleventh of the length of the deutomerite; cysts spherical, sporocysts
cylindrical; average sporonts Sou x 180u.
Pterocephalus A. Schneider, 1887, p. 67.

Nina Grebnicki, 1873. Preoccupied.

Type—P. scolopendre (Kolliker) = Gregarina scolopendre Kolliker,

1848; from Scolopendra sp.—Chilopoda.

Dactylophorus Balbiani, 1889, p. 41.

Dactylophora Leger, 1892, p. 124. Preoccupied.
Type—D. robustus (Leger), 1892, p. 124, t. 9; from Cryptops hortensis
Leach—Chilopoda.
Trichorhynchus A. Schneider, 1882, p. 438.

Type—T. pulcher A. Schneider, 1882, p. 438; from Scutigera sp.—Chilo-
poda.

Trichorhynchus pulcher A. Schneider. Fig. 61.
Trichorhynchus pulcher A. Schneider, 1882, p. 438; Crawley, 1903a, p. 52.
Gregarina megacephala Leidy, 188, p. 11, 1f.; from Cermatia forceps.

Epimerite short and conical, borne by a much produced portion of the
protomerite; sporont elongate, reaching the length of 8004; cysts ovoid;
sporocysts cylindrical.

Trichorhynchus lithobii Crawley. Fig. 62.
Trichorhynchus lithobii Crawley, 1903b, p. 637, pl. XXX, f. 18; Raleigh,
North Carolina, from Lithobius sp.

Since the determination of this gregarine remains quite uncertain until
it is more fully described, a portion of Crawley’s original diagnosis is
copied here to accompany his figure: “An epimerite was not seen. The
protomerite was subcordiform, and displayed in front a differentiation the
exact nature of which could not be determined. The deutomerite varied con-
siderably in shape, the animal being quite polymorphic.—The largest indi-
vidual seen was 195 microns long.”

Acutispora Crawley, 1903b, p. 632.

Type—A. macrocephala Crawley, 1903b, p. 632-633, pl. XXX, f. 1-6;
from Lithobius forficatus L.—Chilopoda.

Acutispora macrocephala Crawley. Figs. 63-64.
Acutispora macrocephala Crawley, 1903b, p. 632-633, pl. XXX, f. 1-6;
Raleigh, North Carolina, from Lithobius forficatus.

Epimerite uncertain; sporont rather elongate, tapering posteriorly, the
posterior end of the deutomerite broadly rounded; protomerite constricted
near its posterior third, narrower than the deutomerite; width of the pro-

CEPHALINE GREGARINES OF THE NEW WORLD 285

tomerite about one-half of its length, which is a little less than one-third of
the total length of the animal; sporocysts ellipsoidal, narrow and pointed,
about 4u x 19u; cysts spherical, with a large lateral pseudocyst.

Metamera Duke, 1910, p. 261.

Type—M. schubergi Duke, rgro, p. 261-286, pl. 15-16; from Glosso-

siphonia complanata and Hemiclepsis marginata—Hirudinea.

This genus at present contains but a single species, M. schubergi Duke,
known from England and Germany. In his description of this species Duke
(1910, p. 262), states that it is “identical with a species briefly mentioned by
Bolsius in 1895, and the subject of a more detailed but still fragmentary
paper in 1896.” On the same page Duke calls attention to the fact that Castle
(1900), “mentions having observed the gregarine seen by Bolsius in about
half the specimens of Clepsine elongata which he examined.” In this
roundabout way there exists a North American record of a gregarine prob-
ably referable to the genus Metamera. This gregarine, listed as Gregarina
complanata by Castle (1900, p. 60) from Glossiphonia elongata, is deserving
of study when material is obtained.

STENOPHORID

Dehiscence of cysts by simple rupture; sporocysts ovoid, not
in chains; epimerite present only in the intracellular stage; anterior
portion of the protomerite with a thin central area in the epicyte
so that the protomerite when seen in optical section appears to have
a central canal in its anterior end.

Stenophora Labbe, 1899, p. 15.

Stenocephalus A. Schneider, 1875, p. 584. Preoccupied.

Type—S. juli (Frantzius) = Gregarina juli Frantzius, 1848, p. 191-194;

from Julus sp—Diplopoda.

Cnemidospora A. Schneider, 1882, p. 446.

The species of this genus are parasites of Diplopods, although two
species, S. erratica Crawley and S. gimbeli Ellis, have been recorded from in-
sects. These two species, as suggested by Crawley (1907) regarding his
species S. erratica, may be accidental and atypical forms of some of the
regular Diplopod-infesting Stenophore, resulting from the introduction of
the sporocysts into the wrong host.

Stenophora robusta Ellis. Fig. 72.

Stenophora robusta Ellis, ror2b, p. 8-11, f.1 a-b; from Parajulus venus-

tus Wood and Orthomorpha gracilis (Koch), Boulder, Colorado.

Short and ovoid, posterior margin of the deutomerite broadly rounded;
protomerite narrower than the deutomerite, subconic, its length about one-
eighth of the total length; size under 250u. The writer has taken this gre-
garine from specimens of Orthomorpha sp., collected at Gold Hill, Colorado,
at an altitude of 8400 feet, in November, 1912, by Miss Rosamond Patton.

Stenophora gimbeli Ellis. Fig. 71.

286 MAX M. ELLIS

Stenophora gimbeli Ellis, r9r3a, p. 464, f. 3-4; Vincennes, Indiana, from

Harpalus pennsylvanicus Dej.

Short and ovoid, posterior margin of the deutomerite broadly rounded;
protomerite almost as wide as deutomerite, hemispherical, wider than long,
its length about one-sixth of the total length; average specimens, 500n.

Stenophora erratica Crawley. Fig. 609.

Stenophora erratica Crawley, 1907, p. 221, pl. XVIII, f. 5; from Gryllus

abbreviatus.

Slightly elongate, posterior margin of the deutomerite broadly rounded;
protomerite equalling the deutomerite in width, subconical, its length about
one-fourth of the total length; reaching the length of 5oou.

Stenophora julipusilli (Leidy). Fig. 65.

Gregarina julipusilii Leidy, 1853, p. 238, pl. 10, f. 21-22; from Julus

pusillus.

Stenophora julipusilli, Crawley, 1903b, p. 634, pl. XXX, f. 16-17; from

Julus sp. and Parajulus sp.: Hall, 1907, p. 149; Lincoln, Nebraska.

Somewhat elongate, (young specimens ovoid), posterior margin of the
deutomerite rounded; protomerite conical to almost biconic, anterior end
rather distinctly pointed; length of the protomerite in adult specimens about
one-tenth of the total length.

Stenophora larvata (Leidy). Fig. 7o.

Gregarina larvata (Leidy) 1849, p. 232; from Julus marginatus.

Gregarina julimarginati Leidy, 1853, p. 237, pl. 10, f. 1-20; from Julus

marginatus.

Stenophora juli, Crawley, 1903a, p. 51; from Julus sp. and Parajulus sp.

Elongate, posterior margin of the deutomerite narrowly rounded to al-
most pointed; protomerite hardly as wide as the widest portion of the deu-
tomerite, hemispherical to subglobose; length of the protomerite about one-
twentieth of the total length of adult specimens.

Stenophora spiroboli Crawley. Fig. 66.

Stenophora spiroboli Crawley, 1903a, p. 51-52, pl. II, f. 22; Raleigh, North

Carolina, from Spirobolus sp.

Cnemidospora spiroboli Crawley, 1903b, p. 638-630, pl. XXX, f. 7-9.

Elongate, rather pointed posteriorly, protomerite narrower than the deu-
tomerite; length of the protomerite about one-thirty-second of the total
length; cysts spherical; sporocysts spindle-shaped, with heavy epispore, size
12.54 X 7.54. This species may be a synonym of S. larvata (Leidy).

Stenophora cockerelle Ellis. Fig. 67.

Stenophora cockerelle Ellis, ror2c, p. 681-685, f. 1-3; Quirigua, Guate-

mala, from Parajulus sp.

Elongate, posterior margin of the deutomerite broadly rounded to al-
most square; protomerite subglobose with a distinct papilla at its anterior
end; width of the protomerite about one half that of the deutomerite, length
of the protomerite about one-sixteenth of the total length.

CEPHALINE GREGARINES OF THE NEW WORLD 287

Stenophora elongata Ellis. Fig. 68.

Stenophora elongata Ellis, ror2zc, p. 685-686, f. 4; Quirigua, Guatemala,

from Orthomorpha coarctata (Saussure).

Extremely elongate, posterior margin of the deutomerite rounded; pro-
tomerite as wide or slightly wider than the deutomerite, pentagonal in out-
line, pointed anteriorly; length of the protomerite about one-twenty-fourth
of the total length .

DOLIOCYSTID

Septum wanting, protomerite continuous with the deutomerite ;
epimerite simple and digitiform; sporocysts oval with an enlarge-
ment at anterior ends; habitat, intestine of marine annelids.

Doliocystis Leger, 1893, p. 204-200.
Type—D. pellucida (K6lliker) = Gregarina pellucida Kolliker, 1848, p.
35, t. 3, f. 31, from Nereis sp.—Polycheta.:

Doliocystis rhynchoboli Crawley.
Doliocystis rhynchobli Crawley, 1903a, p. 56; nomen nullum, Porter,
1897b, p. 8, pl. 3, f. 37; from Rhynchobolus americanus Verrill.

Host List
Host Gregarine
ANNELIDA

Rhyncobolus americanus Verrill Doliocystis rhyncobli Crawley
HIRUDINEA

Glossiphonia nepheloidea (Graf)
Glossiphonia elongata Castle... Metamera sp. ?

DIPLOPODA
OWLOTAG, ISD: svc ticielsteieieaite ans syste dmphoroides fontarie Crawley
MAUS ASD sostsc todo, echehivereriaaierslavaroareste Stenophora julipusilli (Leidy)
Julus minutus Brandt
EDS PMSAPIIS, (o.oo oils ae amon a ays eae ienophora julipusilli (Leidy)
Lysiopetalum lacterium (Say)....Gregarina calverti Crawley

Stenophora julipusilli (Leidy)
Orthomorpha coarctata (Saussure) Stenophora elongata Ellis
Orthomorpha gracilis (Knoch) ..Stenophora robusta Ellis

Ovihomorpha sp: 3.5. s0aaceonaeae Stenophora robusta Ellis
Parajulus venustus Wood........ Stenophora robusta Ellis
IPGFOIUIUS? SD: wdieccia tele eee eee Stenophora cockerelle Ellis
Parajuiis Spe, Jee ciecion tot ool se Stenophora julipusilli (Leidy)
Polydesmus virginiensis .......... Amphoroides polydesmivirginiensis
(Leidy)

Polydesmusssp-ae ssa en es Amphoroides fontarie Crawley
Spirobolus marginatus (Say) ....

Julus marginatus Say..........é Stenophora larvata (Leidy)

SPIFOUOIUNS: SD seco eels ais aie.2 fa oars Stenophora larvata (Leidy)

288 MAX M. ELLIS

CHILOPODA
Lithobius coloradensis (Cockerell) Echinomera hispida (A. Schneider)
Lithobius forficatus (Linn.) ...... Acutispora macrocephala Crawley
Actinocephalus dujardini A. Schneider
Echinomera hispida (A. Schneider)
ERROR BD Oo5 be. ostlean ook Trichorhynchus lithobii Crawley
Scolopendra heros Girard......... Amphorocephalus amphorellus Ellis
Scolopendra woodi Meinert ...... Aniphorocephalus actinotus (Leidy)
Scolopocryptops sexspinosus
Et: gy bahia Catt eng bape a tes Pr aa sar Amphorocephalus actinotus (Leidy)
scolopocrypiops sp. tic OTs Amphorocephalus actinotus (Leidy)
Scutigera forceps (Rafinesque) ...... Trichorhynchus pulcher A. Schneider
Cermatia forceps’: sce. os sce Trichorhynchus pulcher A. Schneider
ORTHOPTERA
Blaptica dubia (Serv.)
Blabera claraziana Saussure....Gregarina blabere Frenzel
Blatta orientalis Linn. ............ Gregarina blattarum Siebold
Periplaneta orientalis .......... Gregarina blattarum Siebold
Blaitella germanica (Linn.)
Ectobia germanica ............ Gregarina blattarum Siebold
Brachystola magna Girard........ Gregarina rigida (Hall)
Ceuthopilus latens Scudder ...... Gregarina longiducta Ellis
Ceuthophilus maculatus (Harris) Gregarina longiducta Ellis
Ceuthophilus valgus Scudder.......... Gregarina consobrina Ellis
Chortophaga viridifasciaia (De
Geer )
Chimarocephalus viridifasciata.. Actinocephalus pachydermus (Crawl.)
Dissosteira carolina (Linn.) ...... Actinocephalus pachydermus (Crawl.)
Locusta carolina Linn. ........ Gregarina locustecaroline Leidy
Gryllus abbreviatus Serv. ........ Stenophora erratica Crawley
Acheta abbreviata ............ Gregarina achete-abbreviate Leidy
Gigaductus kingi (Crawley)
Gryllus americanus Blatchley ....Gregarina achete-abbreviate Leidy
Ischnoptera pennsylvanica (De
Geer a aetee see ek eee Gregarina blattarum Siebold
Melanoplus angustipennis (Dodge) Gregarina rigida (Hall)
Melanoplus atlanis (Riley) ...... Gregarina rigida (Hall)
Melanoplus bivittatus (Say)...... Gregarina rigida (Hall)
Melanoplus coloradus Caudell
Melanoplus coloradensis ....... Gregarina rigida (Hall)

Melanoplus differentialis (Uhler).Gregarina rigida (Hall)
Melanoplus femoratus (Burmeis-

CET) Se eee ean Gregarina rigida (Hall)
Melanoplus femur-rubrum (De-
Geer) <2. SF eee EReers eaten Gregarina rigida (Hall)

CEPHALINE GREGARINES OF THE NEW WORLD 289

Melanoplus luridus (Dodge) ..... Gregarina rigida (Hall)

Panchlora exoleta Burmeister ....Gregarina panchlore Frenzel

Periplaneta americana (Linn.)....Gregarina blattarum Siebold

Gregarina serpentula Magalhes

ISOPTERA

Termes flavipes Kollar .......... Gregarina termitis Leidy

Termes lucifugus Rossi .......... Gregarina termitis Leidy
ODONATA

Aeshna constricta Say, nymph....Geniorhynchus eshne Crawley

Aeshna Sp: lymph ts... sie ucise se Actinocephalus brachydactylus Ellis
COLEOPTERA

Acilius sp.

Acilius sulcatus (European)....Bothriopsis histrio A. Schneider
Alobates pennsylvanicus (DeGeer) Actinocephalus zophus (Ellis)

Nyctobates pennsylvanicus ..... Anthorhynchus philicus (Leidy)
Nyctobates pennsylvanicus bar-

Dear CUSOCH)® 225 cave de oes se Actinocephalus zophus (Ellis)
IE DPOLE SAY 5.2522 di ese 0s Stylocephalus giganteus Ellis
Boletophagus sp.

Boletophagus cornutus ......... Anthorhynchus boletophagi (Crawl.)
Calpmberes: spr S22 e.2. dolls ese
Colymbetes fuscus Linn. (Eu-

CREED eens Pey ferale Sarath ='s,5 Sielons Bothriopsis histrio A. Schneider
Cratoparis lunatus (Fab.)........ Anthorhynchus cratoparis (Crawl.)
CM CUIIE LAKUCO oo sicss ae oot Hoje ne ose Gregarina elatere Crawley
Dermestes peruvianus Castelnau..Pyxinia crystalligera Frenzel
Dermestes vulpinus Fab. ......... Pyxinia erystalligera Frenzel
Dicelus ovalis LeConte .......... Actinocephalus disceli (Crawl.)
WER SDs NAT V inal s ovscloe a aeiaiatots Gregarina elatere Crawley
Eleodes hispilabris (Say) ........ Stylocephalus giganteus Ellis
LE] ret aos Cs SIS Ie SOURED AO DOr nc Stylocephalus giganteus Ellis
[DAG ET SAC ae a Reo E IS e enre ws tylocephalus giganteus Ellis
Galerita bicolor Drury............ Actinocephalus americanus Crawl.
Harpalus caliginosus Fab. ........ Gigaductus parvus Crawley

Actinocephalus harpali (Crawl.)
Harpalus pennsylvanicus DeGeer..Gigaductus parvus Crawley
Stenophora gimbeli Ellis
Holocephala bicornis Olivier...... Gregarina microcephala Leidy
Hydaticus sp.
Hydaticus cinereus, larve (Euro-

pean) “sco navoe nas ee es em = 2 Bothriopsis histrio A. Schneider
Eiyaropiilas Sin as sstek vena gee Legeria terpsichorella Ellis
Leptochirus edax Sharp...........: 4ctinocephalus crassus (Ellis)

Siylocystis ensiferus (Ellis)

290 MAX M. ELLIS

Ligyrus relictus (Say)

Scarabeus relictus, larve...... Gregarina scarabeirelecti Leidy

Lucanus dama Thunb............ Euspora lucani Crawley
Necrobia ruficollis Fab.

Corynetes ruficollis ...........- Gregarina bergi Frenzel
Neleus interstitialis Esch......... Gregarina guatemalensis Ellis
Passalus cornutus Fab............ Gregarina passalicornuti Leidy
Serica brunnea Linn.

Melalontha brunnea ........... Gregarina melalonthe-brunnee (Leidy)
Statira unicolor Blanchard....... Gregarina statire Frenzel
Tenebrio castaneus Knoch........ Gregarina grisea Ellis

Xylopinus saperdioides (Olivier). Gregarina xylopini Crawley
‘Actinocephalus zophus (Ellis)
TUNICATA i
Perophora annectens Ritter...... Gregarina sp.
Acknowledgments.

The writer wishes to express his thanks to Dr. Paul S. Welch
who determined the hosts taken at Douglas Lake, Michigan; to
Prof. T. D. A. Cockerell who revised the list of host names;
and to Mr. Arthur G. Vestal for reading the manuscript.

University of Colorado, October 15, 1913.

CEPHALINE GREGARINES OF THE NEW WORLD 291

BIBLIOGRAPHY

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1912a,A new species of Polycystid Gregarine from the United States,
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292 MAX M. ELLIS

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CEPHALINE GREGARINES OF THE NEW WORLD 293

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294 MAX M. ELLIS
EXPLANATION OF FIGURES
Piate XVII
FIGURE

T;

io GON ON

Euspora lucani Crawley. Association. (After Crawley, 1903a, pl.-III, f.
38).

Gregarina blabere Frenzel. Sporont. (After Frenzel, 1892, f. 22).

Gregarina serpentula Magalhes. Association. (After Magalhzs, 1900,
tA).

Gregarina calverti Crawley. Sporont. (After Crawley, 1903a, pl. II,
f. 19).

Gregarina achete-abbreviate Leidy. Association. Douglas Lake, Mich-
igan.

Gregarina termitis Leidy. Sporont. Boulder, Colorado.

Gigaductus kingi (Crawley). Association. (After Crawley, 1907, f. 10).

Gigaductus parvius Crawley. Association. Vincennes, Indiana.

Gregarina panchlore Frenzel. Anterior portion of satellite. (After
Frenzel, 1892, f. 20).

Gregarina elatere Crawley. Cephalont. (After Crawley, 19o03a, pl. I, f.
1T)2

Gregarina elatere Crawley. Sporont. (After Crawley, 1903a, pl. I, f. 5).

Gregarina passalicornuti Leidy. Association. (After Leidy, 1853, pl.

Ti, f.° 30):

Gregarina rigida (Hall). Sporont. Boulder, Colorado.

Gregarina statire Frenzel. Association. (After Frenzel, 1892, f. 1).

Gregarina guatemalensis Ellis. Association. Quirigua, Guatemala.

Gregarina passalicornuti Leidy. Sporont. New Orleans, Louisiana.

Gregarina xylopini Crawley. Association. (After Crawley, 1903a, pl.
MS yoy) ie

Gregarina grisea Ellis. Association. New Orleans, Louisiana.

Gregarina locustecaroline Leidy. Cephalont. (After Crawley, 1907,

£) 53).

Pirate XVIII

Gregarina blattarum Siebold. Association. Douglas Lake, Michigan.

Gregarina blattarum Siebold. Sporocyst. Douglas Lake, Michigan.

Gregarina blattarum Siebold. Cyst with developing sporoduct-buds.
Michigan.

Gregarina consobrina Ellis. Cephalont. Boulder, Colorado.

Gregarina consobrina Ellis. Association. Boulder, Colorado.

Gregarina consobrina Ellis. Dehiscing cyst. Boulder, Colorado.

CEPHALINE GREGARINES OF THE NEW WORLD 2905

Gregarina longiducta Ellis. Dehiscing cyst. Douglas Lake, Michigan.

Gregarina longiducta Ellis. Cephalont. Douglas Lake, Michigan.

Gregarina longiducta Ellis. Sporocyst. Douglas Lake, Michigan.

Gregarina longiducta Ellis. Association. Douglas Lake, Michigan.

Legeria terpsichorella Ellis. Sporont. Douglas Lake, Michigan.

Actinocephalus brachydactylus Ellis. Anterior portion of cephalont.
Michigan.

Actinocephalus brachydactylus Ellis. Cephalont. Douglas Lake, Mich-
igan.

Actinocephalus brachydactylus Ellis. Sporont. Douglas Lake, Michigan.

PiLatTeE XIX

Stylocystis ensiferus (Ellis). Cephalont. Quirigua, Guatemala.

Stylocystis ensiferus (Ellis). Sporont. Quirigua, Guatemala.

Amphoroides polydesmivirginiensis (Leidy). Sporont. (After Craw-
ley, 1903a, f. 25).

Amphoroides fontarie Crawley. Sporont. East Falls Church, Vir-
ginia.

Gregarina bergi Frenzel. Cephalont. (After Frenzel, 1892, f. 16).

Gregarina bergi Frenzel. Sporont. (After Frenzel, 1892, f. 17).

Actinocephalus crassus (Ellis). Sporont. Quirigua, Guatemala.

Geniorhynchus @shne Crawley. Cephalont. (After Crawley, 1907,
Be oA}

Bothriopsis histrio A. Schneider. Cephalont. (After Leger, 1892, pl.
MLL, ft. ¥);

Pyxinia crystalligera Frenzel. Cephalont. (After Frenzel, 1892, f. 35).

Pyxinia crystalligera Frenzel. Sporont. (After Frenzel, 1802, f. 39).

Actinocephalus dujardini A. Schneider. Cephalont. (After Schneider,
1675, pl. XVI, £..0:)

Actinocephalus harpali (Crawley). Sporont. (After Crawley, 19038,

2 Oe Te ea

Anthorhynchus cratoparis (Crawley). Cephalont. (After Crawley,
TO032a-).p)... Ulf: 23°

Anthorhynchus philicus (Leidy). Cephalont. (After Crawley, 1903a,
plelilt: Sr):

Actinocephalus zophus (Ellis). Cephalont. New Orleans, Louisiana.

Actinocephalus disceli (Crawley). Sporont. (After Crawley, 19038,
plate f.2)8

Amphorocephalus amphorellus Ellis. Sporont. Boulder, Colorado.

Amphorocephalus amphorellus Ellis. Cephalont. Boulder, Colorado.

Amphorocephalus actinotus (Leidy). Cephalont. (After Crawley,
1e03a, pl EU tes hi

Actinocephalus pachydermus (Crawley). Sporont. (After Crawley,
zoe7, 1.3):

58.

59.

MAX M. ELLIS

Actinocephalus pachydermus (Crawley). Cephalont. (After Leidy,

1853, pl. II, f. 37).

Actinocephalus americanus Crawley. Sporont. (After Crawley, 1903b,
hie).

Anthorhynchus boletophagi (Crawley). Sporont. (After Crawley,
19038, pl. LI, £. 26).

PLATE XX

Stylocephalus giganteus Ellis. Anterior portion of a cephalont. Boul-
der, Colo.
Stylocephalus giganteus Ellis. Sporont. Boulder, Colorado.

50s. Stylocephalus giganteus Ellis. Sporocysts. Boulder, Colorado.

60.
61.

62.

&

Vas snd

Echinomera hispida (A. Schneider). Cephalont. Boulder, Colorado.

Trichorhynchus pulcher A. Schneider. Anterior portion of a cephalont.
(After Schneider, 1882, f. 4).

Trichorhynchus lithobii Crawley. Sporont. (After Crawley, 1903b, f.
18).

Acutispora macrocephala Crawley. Sporont. (After Crawley, 1903b,
rE De

Acutispora macrocephala Crawley. Sporocyst. (After Crawley, 1903b,
ot):

Stenophora julipusilli (Leidy). Young sporont. After Leidy, 1853,
PL c104 f' 21);

Stenophora spiroboli Crawley. Sporont. (After Crawley, 1903a, pl.
Ei, £22).

Stenophora cockerelle Ellis. Sporont. Quirigua, Guatemala.

Stenophora elongata Ellis. Sporont. Quirigua, Guatemala.

Stenophora erratica Crawley. Sporont. After Crawley, 1907, f. 5).

Stenophora larvata (Leidy). Sporont. (After Leidy, 1853, pl. 10, f. 1).

Stenophora gimbeli Ellis. Sporont. Vincennes, Indiana.

Stenophora robusta Ellis. Sporont. Boulder, Colorado.

SS ee

PLaTE XVII

PLaTE XVIII

PEATE xox

PLATE XX

DEPARTMENT OF NOTES, REVIEWS, ETC.

It is the purpose, in this department, to present from time to time brief original
notes, both of methods of work and of results, by members of the Society. All members
are invited to submit such items. In the absence of these there will be given a few brief
abstracts of recent work of more gencral interest to students and teachers. There will be
no attempt to make these abstracts exhaustive. They will illustrate progress without at-
tempting to define it, and will thus give to the teacher current illustrations, and to the
isolated student suggestions of suitable fields of investigation.—[Editor.]

A PARAFFIN RIBBON CARRIER

The Carrier described below was designed to handle the par-
affin ribbon as it comes from the microtome in such a way as to
preserve a perfect series and to eliminate some of the difficulties
encountered with the usual method. Not only is the old method of
cutting long serial sections into short pieces and laying them upon
a sheet of paper tedious but the danger of losing a part of the sec-
tions in a sudden draft or of having them hopelessly mixed is great.
Without careful shielding, ribbons placed upon paper may not be
allowed to lie for any length of time before mounting. With the
use of the carrier long unbroken series may be wound on the drum
and allowed to remain until used. The writer has allowed a ribbon
to remain on the carrier for three days exposed to all the drafts
common in the average room and at the end of the time was able to
mount a perfect series with no difficulty. The inclined plane shown
in the photograph greatly facilitates mounting. The ribbon is un-
wound from the drum onto the plane where it is cut to the desired
lengths. The continuous ribbon does away with the bother of piec-
ing bits together as is frequently necessary when mounting from
short strips laid on paper.

Directions for Making
Material.

All the materials necessary for making the machine are easily
obtained and at slight expense. The wood used was poplar and
yellow pine taken from an old packing box. A one pound coffee
tin was made to serve as the cylinder or drum.

298 NOTES, REVIEWS, ETC.

Dimensions.

Figure 1. A.

1. The Base. 9x6x¥ inches. A slot 24%4x% inches (Fig. 2-
10) is cut in the center to accommodate the winged nut (9) which
fastens the uprights (3) to the base.

2. The Base to which the uprights are fastened. 7x3x%4
inches. A quarter inch hole is bored in the center of this base to
pass the screw of the winged nut through.

3. The Uprights. 9x3x™% inches. The width narrows four
inches from the bottom to 1% inches. Slots 1x¥% inches are cut in
the tops of the uprights (Fig. 1D) to accommodate the axles of
the drum. The uprights were made particularly for the Leitz Base
Sledge microtome and, though the height to which they raise the
drum works very well with the rotaries, two inches might be cut
from their length with advantage.

4. These are pegs that are inserted four and three quarter
inches from the base to carry the inclined plane shown in Fig. 2-11.

5. The Axle Support. This is a disc of wood 2x% inches
through the center of which a quarter inch walnut axle is thrust.
The end of a maple spool is glued to the top of the disc to serve as
a bearing (Fig. 1, B-3).

6. The Drum. The lid is soldered onto a one pound coffee
tin, the measurements of which are 6x4 inches. The can is covered
with blotting paper. This drum will carry about fourteen feet of
one-half inch ribbon.

7. The Drum Pulley. 2x™%4 inches and grooved as shown in
the diagram. The belt runs from this pulley to one made by fasten-
ing the ends of a small spool together and which is secured to the
inner face of the driving wheel.

8. The Driving Wheel. 3x%4 inches. A handle is inserted
in one side as shown in the diagram though very little use is found
for it.

9. Winged Nut. % inch in diameter.

Figure 2.

10. Slot 21%4x% inches in which the bolt of the winged nut

Ie LVI]
a

IIXxX wv Id

a Dri

G

AMERICAN MICROSCOPICAL SOCIETY 299

slides. This allows the adjustment of the drum to meet require-
ments.

11. Inclined Plane. 12x634x% inches. At one end two pegs
are placed to engage those shown in Fig. 1,-A4.

General Directions

The only difficulty that will be encountered in the making of
the carrier will be to fasten the axle supports squarely in the center
of the drum.

Centering the axles may be easily accomplished by drawing a
circle the exact size of the drum on a board and then, after de-
termining the center, drill a hole the size of the axle (%4 inch)
through it. Insert the axle into this hole (Fig. 1, E). Drive long
brads or nails at the periphery of the circle as shown in this figure
so as to hold the drum firmly in place when it is lowered. Coat the
axle support with glue and press the drum tightly against it. The
nails will hold the drum in place and the axle will be in the exact
center of the cylinder.

Glue may serve to fasten the axle permanently to the drum but
I find that it does not take a very firm hold of the tin and soon breaks
away. This may be overcome by first placing small brads or screw
eyes (Fig. 1,-B 1) in the sides of the axle supports and then
gluing the discs on as directed above. After the glue has set firmly
enough to hold the discs in place solder is run in under the screw
eyes and they are thus firmly fastened to the tin.

Directions for Using

Cut a ribbon from eight to ten inches in length and press one
end lightly against the blotting paper covering the drum. After
this the ribbon is wound on the cylinder as it comes from the micro-
tome (shown in Fig. 2) by thumbing the edge of the driving wheel
which revolves the drum very slowly (the drum revolves once to
every two and a half revolutions of the driving wheel). The winged
nut allows the cyliner to be adjusted to the demands of the particu-
lar microtome in use. The ribbon is wound spirally upon the drum
by sliding the carrier parallel to the knife. When ready to mount
the ribbon is unwound onto the inclined plane which is covered
with blotting paper and cut to the desired lengths.

Univ. of Cincinnati. Rosert THEODORE HANCE.

300 NOTES, REVIEWS, ETC.

DIVISION OF LABOR IN THIS DEPARTMENT

It is the hope of the Editor to secure the aid of special workers
in the preparation of these notes. Dr. Paul S. Welch has agreed to
furnish abstracts of such entomological papers as may seem suitable
for the purposes of this department. The following notes are fur-
nished by him. Dr. George R. La Rue will furnish abstracts in
histology and microtechnic.

INSECTS AND DISEASE

Doane (Journ. Econ. Ent., 6:366-385, 1913) contributes an im-
portant and useful paper entitled: “An Annotated List of the
Literature on Insects and Disease for the year 1912.” Brief men-
tion is made of the work of Brues and Sheppard, Rosenau, and An-
derson and Frost on the transmission of infantile paralysis to mon-
keys by the common stable fly, Stomo.rys calcitrans. Whether this
is the usual method of transmission among human beings remains
to be determined. The Simuliide have continued to receive attention
on account of their possible relation as carriers of pelagra. The
work of Forbes, Garman, and Hunter is referred to as presenting
very important circumstantial evidence but it remains to be proven
that these flies really carry the virus which causes the disease.
Other recent work on malaria and mosquitoes, the house fly and
typhoid, and trypanosomes and sleeping sickness is briefly men-
tioned. The chief value of the article lies in the extensive biblio-
graphy which contains nearly three hundred references to works on
medical entomology issued during 1912.

ADAPTATION IN THE GALL MIDGES

Felt (Canadian Entomol., 45:371-379, 1913) discusses “Adap-

tation in the Gall Midges.” Forms of adaptation are grouped under
three heads: (1) Strength, aggressive and defensive, (2) Prolifi-
cacy, and (3) Evasive adaptations. Bud galls, leaf galls, stem galls,
and root galls are discussed with reference to these classes. In
spite of the fact that this group of insects, because of its general
similarity in habit, might be thought to exhibit slight variations in
structure a number of interesting significant structural modifica-
tions are found in the antenne, palpi, wings, and generative organs.
It is shown that the gall midges can not be counted as particularly
strong or prolific forms but they have been able to maintain them-

AMERICAN MICROSCOPICAL SOCIETY 301

selves largely by evasive adaptations which secure protection for
them at the expense of the host plant. This group of insects pre-
sents many interesting biological and morphological problems which
are unsolved and according to Dr. Felt there is perhaps no insect
family better suited in many ways for the study of adaptation.

BIOLOGY OF MAY-FLIES

Morgan (Ann. Ent. Soc. Amer., 6:371-413, 1913) in an article
entitled “A Contribution to the Biology of May-flies” gives inter-
esting and valuable data on the different stages of the life history
and modifications of the structures of the nymph and adult. The
amount of detail makes a short summary impossible. Aside from
the considerable amount of new data which is presented, the fea-
ture of the paper which is of particular value to teachers and in-
vestigators is the complete bibliography on May-flies at the end of
the paper which contains approximately 300 titles of foreign and
American literature.

HIBERNATION OF THE HOUSE-FLY
Skinner (Ent. News, 24 :303-304, 1913) in discussing the often
repeated question as to what becomes of the common house-fly dur-
ing the winter opposes the views held by Howard and Hewitt who
claim that the fly hibernates as an adult. His observations lead him
to believe that the house-fly hibernates as a pupa and not as an adult.

A PARASITE OF THE CHINCH BUG

McColloch (Can. Ent., 45 :342-343, 1913) gives a preliminary
report of the discovery of a hymenopterous parasite on the eggs of
the chinch bug which promises to be of considerable economic inter-
est. Mr. A. B. Gehen, Entomological Assistant of the Bureau of
Entomology, U. S. Dept. of Agric., to whom the adult parasite was
sent for identification, determined it as a member of the family
Proctotrypide and states that preliminary examination indicated that
it is both a new species and a new genus. The parasite was found
in every wheat and corn field examined around Manhattan, Kansas.
The average percentage of parasitism has been found to be about
20.8. The length of the life cycle was found to vary from Io to
18 days and as many as six generations were noted between May
19 and August 10. A complete account is to appear later.

302 NOTES, REVIEWS, ETC.

GRASSHOPPERS

Vestal (Biol. Bull., 25 :141-180, 1913) contributes an ecological
paper on “Local Distribution of Grasshoppers in Relation to Plant
Associations”. He finds that the distribution of grasshopper spe-
cies within the region studied bears evident relation to the plant
associations and the extent of the latter marks the areas of the
different habitats. Grasshoppers select habitats or associations in
which favorable conditions are to be found, irrespective of past
history, extensiveness, geographical or successional relationships of
the vegetation. Only rarely is there any direct relation between
grasshopper species and species composition of the plant associa-
tions, as few grasshoppers are selective feeders. The following
points are also brought out: Grasshopper species can be arranged
according to gradients of environmental factors. Grasshopper suc-
cession is incidental to the development of vegetation; the change 1s
not only one of species but one of habits as well. Grasshopper spe-
cies have in general the geographic range of the types of associations
which include the necessary physical and vegetational conditions.
Species of least definite local distribution are widespread geograph-
ically. Seasonal differences in time of activity of grasshopper
species, probably in part due to the antagonistic influence of other
animals, is marked. Seasonal and local distribution are interrelated.
Species of indefinite local distribution have also least definite time
distribution.

AMERICAN MICROSCOPICAL SOCIETY 303

LIST OF MEMBERS

HONORARY MEMBERS
Crisp, Frank, LL.B., B.A., F-R.M.S.,
5 Landsdowne Road, Notting Hill, London, England

|S SE Tea [id a ar 69 Burling Lane, New Rochelle, N. Y.
Warp, R. Hatstep, A.M., M.D., F.R.M.S.......... 53 Fourth St., Troy, N. Y.
LIFE MEMBERS
OS RS 489 Fifth Ave., New York City
DuNcANSON, Pror. Henry B., A.M...............: State Normal, Peru, Neb.
Eee Peon Amrnaunt EE oo... . 6s .a ois dea 52 E. gist. St. New York City.
SES ea Oe ge 28 ogee ee ee Chicago Beach Hotel, Chicago, Ill.
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 ADMITTED SINCE THE LAST PUBLISHED LIST

Arnold, Frank
Baldwin, Herbert B.
Bass, > C.
Bennehoff, J. D.,
Brode, Howard S.

Buckingham, Edwin W., Jr.

Garison; ‘C::O:
Chambers, Robert J.
Daugherty, Lewis S:
Dole, J. Wilbur
Eggleston, H. R.
Gabel, Charles E
Goldsmith, G. W.
Gregory, Emily R.
Griffin, Lawrence E.
Hance, Robert T.
Harman, Mary T.
King, Williard V.
Kroeck, Louis
Lewis, L. L.

MacGillivray, Alexander D.

Magath, T. B.
McCreery, Geo. L.

Mullenix, R. C.

Myers, Frank J.

Noll, William C.
Quillian, M. C.

Rankin, Walter M.
Raymond, Rev. R. I.
Rice, William F.

Sawyer, William Hayes, Jr.
Scott, George Filmore
Shira, Austin Flint
Smith, Bertram G.
Spaulding, M. H.
Spurgeon, C. H.
Stumpmeyer, Geo. A.
Stunkard, Horace W.
Tsou, Ying-Hsuan Hsuwen
Turner, Clair E.
Varrelman, Ferdinand A.
Walker, Leva Belle
Wieman, Harry L.
Wright, Stanley H.
Tyrrell, E. G.

304 LIST OF MEMBERS

ACKERT, JAMES EpwarbD, ’II............ Kas. State Ag. Col., Manhattan, Kas.
Reson, Appa: Wi 88 eck 7555s kaos oe Rees 3935 Pine St., Philadelphia, Pa.
ALLEN, Wynrrep E., A.M., ’04........ 1345 N. Harrison St., Stockton, Cal.
IS 0 EEA en, a lens en eee 540 S. Main St., Manchester, III.
PD A IER NE soo wteinemtahas tel 408 House Building, Pittsburg, Pa.
ATHERTON, Pror. L. G., A.B., M.S., ’12..State Normal School, Madison, S. D.
AyWoen, ie. Fie eee a rss plea cee 16 Seneca Parkway, Rochester, N. Y.
Bani, Gesce JEAN Oa. B,, Ws. ie ks se caees 5605 Madison Ave., Chicago, III.
BARN WIN? EIPRBERT: BD. 2 E39). dsc ste keene ae 927 Broad Street, Newark, N. J.
BAER) Re SBR De aA sce a nae 324 North 53rd Ave., Chicago, Ill.
BANKER, Pror. Howarp J., Ph.D., ’11...DePauw University, Greencastle, Ind.
per at a Da Ss 7 ear ee ea A 120 West 46 St., New York, N. Y.
BARKERS ArBERT SA ¥O7 055 So eae eo so saa oe he ee ee Moylan, Pa.
BARKER, FRANKLIN D., Ph.D., ’03..... University of Nebraska, Lincoln, Neb
Payee FW. PB Se MAL ass. nevcks Sek tieutiawsst Clemson College, S. C.
BASS, 2G GeO UD Png irs) fice ee 741 Carondelet Street, New Orleans, La.
BAUSCH 2 EDWARD? 27657005 siaividers sick oe 179 N. St. Paul St., Rochester, N. Y.
BAGSCH AAW IELIAMES (GB. bin doskwses ose. Pos St. Paul St., Rochester, N. Y.
Pocdapetery, Pars AS Begs 2) 2c ks ibe Iie 1412 Tenth St., Greeley, Colo.
Beit, Arsrar T., B:S.,A.M., ’o3......4.... La. State Univ., Baton Rouge, La.
IDPNIE HOREMN Re ESV MSE Ie aiken ace be 9s oe Alfred College, Alfred, N. Y.
BENNETT, Henry C., ’93....Hotel Longacre, 157 W. 47th St., New York City
Bessey, Pror. CHARLES Epwin, Ph.D., LL.D., ’98.............-. Lincoln, Neb.
BETES; SPOH NE eT 1s Seige ee ee eae ele cbs 111 Market St., Camden, N. J.
Bunce, Pao EA ScD, ELD ees: 32. .'. 744 Langdon St., Madison, Wis.
adc. 3 ee MD, ad. tn ee Beek eas 735 Wilson Bldg., Dallas, Texas
Beem .As Me MD aeRy sie a. tape ce Ohio State University, Columbus, Ohio
BoprnE, Pror. DoNALDSON, ’096............- 4 Mills Place, Crawfordsville, Ind.
Booty, Mary A., F.R.M.S., ’82.......... 60 Dartmouth St., Springfield, Mass.
Bors,’ PaorsM Ws Fiery 3 52 nde Municipal Laboratory, Oskaloosa, Ia.
Bove; 62S AM Ve2ieee soe 6140 Columbia Ave., Philadelphia, Pa.
Brope, Howarp S., Ph.D., ’13....... 433 E. Alder Street, Walla Walla, Wash.
Baounoves, Cras. AJB. MiSs 66: S6ci ews nen wn Buchtel Coll., Akron, Ohio
BROWN AMOS: E.. Pie D:, V9T 20 ee ese Fae 20 E. Penn St., Germantown, Pa.
BROw Nica hh: (AL Be) 2s ei. eck William Nast College, Kiukiang, China
BrowNincG, SipNeEY Howarp, ’11..Royal London Ophthalmic Hospital, London
Bryant, Pror. Eart R., A.M. ’Io...... Muskingum College, New Concord, O.
BUCKINGHAM: POWIN Ws Vite. oo o.0 ain op noes cnces woe Box 54 Lexington, Va.
Bont, Samus Wee, FS 702. 2.5. cclce eae eee 141 Broadway, New York City
Beier: (Paam Fro. we BAS OFS oes oe Le ee Re Chapel Hill, N. C.
BURRInSPROR We pee ey 70 toatetatrere Aa oe tos een a came ee ee eee Urbana, IIl.
Cattson (670. Ae es re eines} bx oie oan Doane College, Crete, Nebr.
CARTER; PROF. (GHAREES Dee cos ote cs eaie oss Parsons College, Fairfield, Ia.
Carter, JoHN E., ’86.......... 5356 Knox St., Germantown, Philadelphia, Pa.

Canven, Puor. G. Li Asia: sors bess 526 W. 123rd St., New York, N. Y.

AMERICAN MICROSCOPICAL SOCIETY 305

CHAMBERS, Ropert J., Ph.D., ’13....... 345 Thrall Avenue, Cincinnati, Ohio.
(Grane, Greorce Enw., M.D.,.’06...........20¢ Genessee St., Skaneateles, N. Y.
aU MEIWART WY 2AM TB. os: occa « da ere een ob ihe eelen one Fairport, Iowa
Grements, Mrs. F: E., Ph.D, ’03.....,.. 800 4th St., S. E., Minneapolis, Minn.
Gocautcruor. Grorce BE. Ph.D:, 711... . o.c6 sce: Kas. Univ., Lawrence, Kas.
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Courant, Ricnarp Bi M.D), "2. vs. 2 No. Broadway, Tarrytown, N. Y.
a GEORGE AW) REE ers ih ty had oe dks College View, Nebr.
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Darna cee Leasore Mere, PhiD),: MD), nr... es 6.86. oleh tS hak
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Dupceon, WINTIELD, B.S., ’11....Allabahad Christian Coll., Allabahad, India
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Becreston, ob. R., M.As 713505..0...... Buena Vista, Col., Storm Lake, Ia.
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PemonaLnom SVlontom Io MEAS M.S... "98... « ...< cise aotsies a sperctare ochehele oxen
SRS CS a Soe ST CR a Ce Oe University of Montana, Missoula, Mont.
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LENT ASE, Ue \ 291 DF Rito 0 eee ne Cae ee a 16 Pearl St., Council Bluffs, Iowa
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RE Ie Renee TaN LG nines ieee! cis ars Guy’s Hospital, London, E. C., England
HEART OWE UIEROFVAW. «Gr CDT) osc ists sid score orate iene 24 Quincy St., Cambridge, Mass.
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Bere Grae MDE RMS "98.00 ek so. Morgan Bldg., Buffalo, N. Y.
Fettows, Cuas. S., F.R.M.S., ’83...111 Cham. of Comm., Minneapolis, Minn.
IDERGUSON, MARGARET CLAY, 11.00.26. 020 cscs 46 Dover St., Wellesley, Mass.
Pe eER RM NE OM a ars ci wd diatin « ajale.w sien os 86 62-50% Box 1608, Milwaukee, Wis.
HEnZawANDODEH RAYMOND) Bis ERIM. Se ODT. 2 co. occ ws als lee ch detaichelete ciate
PRES sySeahiciharcisvaina: scaronecove State Laboratory of Hygiene, Trenton, N. J.
Prant, James. MM.D.; ol. ... 0.6.05... Stoneleigh Court, Washington, D. C.
1 ES CRS Se oo 202 S. Thirty-first Ave., Omaha, Neb.
Futon, Harry R., A.B., ’04........ A. & M. College, West Raleigh, N. C.

Purniss; H: W.;M.D!, PhiDs, 70522... ; U. S. Consulate, Port au Prince, Haiti

3006 LIST OF MEMBERS

GaBeL, CuHares E., Ph.D., '13....1340 E. Walnut Street, Des Moines, Iowa

GAcE, Pior. Sriow H., "BS. "Ga eas oa ce iv ovens 4 South Ave., Ithaca, N. Y.
Gace, Mrs. SUSANNA PHELPS,’ 87.............. 4 South Ave., Ithaca, N. Y.
Gattoway, Pror. T. W., A.M., Ph.D., ’or...... 1332 West Wood, Decatur, IIL.
GARRETSON: “UGENE, 21203); Vee .n Sa eee 428 Fargo Ave., Buffalo, N. Y.
GESNER, BROWER CLAIR, ’II........ 110 Steadman St., Moncton, N. B., Canada
Gompsmrrn, GW BIA 13.6025. eB siee S. La. Indus. Inst., Lafayette, La.
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GEA YS Oe ees bret ereds 3535 Telegraph Ave., Oakland, Cal.
GrrconyS Harry Ro Pe ras oe ee bie ds cuales Buchtel Col., Akron, O.
GRIFFIN, LAWRENCE E., ’13....... University of the Philippines, Manila, P. I.
GUTBERLET, JOHN E., A.M., ’11.......... Nat. Hist. Bldg. U. of I., Urbana, Ill.
Guyer, Micwaet F., Ph.D., ’11....... University of Wisconsin, Madison, Wis.
PIAGEER, EE MED. Cana ee cao eos gti eae Hagler Building, Springfield, Ill.
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France; ‘Ropesr T., BiAS 71300 0200.% 723 Freeman Avenue, Cincinnati, Ohio.
PACE SON Ese MIS in rs eS ee bain Saas cae eee Charleston, Ill.
Harman, Mary T., ’13....... Kansas State Agr. College, Manhattan, Kansas.
HEArp be Dee Ph.Ds 66) Zool. Lab. U. of Pa., Philadelphia, Pa.
FPRENDERSON, "WILETAM, 710../0056 5 e'oie oia's oles 1083 E. Eldorado St., Decatur, Ill.
Heney, EpWite CoM. Dit oe ea 554 S. 35th Ave., Omaha, Neb.
HeErtz_er, ArTHUR E., M.D., ’096.......... 402 Argyle Bldg., Kansas City, Mo
Hertzoc, MAXMILIAN, M.D., ’OI........... 06 1604 Mallers Bldg., Chicago, Il.
PRC POL RRE CEP, shot Shite Ses ek oracles 178 Union Ave., Long Branch, N. J.
Phyorray iwpvie Cs 125 ike. Meadowdale, Snohomish County, Washington
Hottts, Freperick S., Ph.D., ’99...... Indiana Med. School, Indianapolis, Ind.
Tlosiknns! Wa 796i. 1285 tt eae sk Shae near 49 6th St., LaGrange, III.
How arp; 'Guorce) £252. ls Sitwell Vale, Moorgate, Rotherham, Eng.
Howarp, Rosert NEssit, ’12..Ookiep, Namaqualand, Cape Province, S. Africa
How.anp, Henry R., A.M., ’08...........--- 217 Summer St., Buffalo, N. Y.
Ives, Freperic E., ’02......... Woodcliff-on-Hudson, Weehawken P. O., N. J.
Jacxson, Dante, Dana, B.S., ’99....... _..930 President St., Brooklyn, N. Y.
SPARENS OVV EE NED: Ono toe he ak eee 1231 Locust St., Philadelphia, Pa.
Pane ee bene PTS ke ca tae ewer 603 S. Fern Ave., Wichita, Kas.
pci GAR ET, RA i rr i ge a Science Hall, Indianola, Ia
JERVIS, SPOWACE Ber Ea TR as sie lg ovec wees voee 8 Charles St., Houlton, Maine
WPEDROSONS HN Mia re aha aie ales we wie ew ssnatstoiede eit oe Joplin, Mo., R. F. D. 4-147
Jounson, Frank S., M.D., F.R.M.S., ’93...... 2521 Prairie Ave., Chicago, Ill.
JORDAN; PROFESS) ataceias ess University Place, Charlottesville, Va.
JUDAYi“CRANCEY: “GON tee Soren ene cee dae 610 Lake St., Madison, Wis.
Kiser an Pie OCP ae Ts cs eiciorwoang oder Jefferson City, Tenn.
Kerroee, J. F., MDS ye eee he 202 Manchester St., Battle Creek, Mich.
Kespen; Prom J. Rs aa Beeches aes wis Baylor University, Waco, Texas

Krncarp, Trevor, A.M., ’12........ University of Washington, Seattle, Wash.

AMERICAN MICROSCOPICAL SOCIETY 307

Keene) WAEEARN GV. Egan cite wis Bea eee eels P. O. Box 261, New Orleans, La.
[AG 4 LS et | oe 5 1o15 Blondeau St., Keokuk, Ia.
Koro, CHartes A., Ph.D., ’99....... University of California, Berkeley, Cal.
apne ees er O Re on lack ocek sino Sota s 32 S. Fourth St., Easton, Pa.
Kriss, H. G., A.B., B.D., ’07....Zool. Lab. Univ. of Pa., Philadelphia, Pa.
ee 29 oa aS Se i 520 Elm Street, San Jose, Calif.
ET NCES BRAY SE Ut Ree Se ee ae ee EY oe oe

tee Bank of New South Wales, Warwick, Queensland, Australia
Exnmacer, Fy 1, B:A;, ’03............ Ohio State University, Columbus, Ohio
ARE Groner. Re 71s... 5)... .).- University of Michigan, Ann Arbor, Mich.
MATT AMAGVITSS. VELAY NE DE DDS: FARMS. 288... e3 so. tec stot taeber

ete eect ays russ ai alors. je feeis, 1644 Morse Ave., Rogers Park, Chicago, Ill.
Latimer, Homer B., M.A., ’11..Neb. Wesleyan Univ., University Place, Neb.
LEHENBAUER, PuHutip, A.M., ’11........Nat. Hist. Bldg., U. of I., Urbana, II.
EEN EMRE aSHEODORE-W Mi" isis. voo<ks eas ouirnee oe Bitter Root Inn, Montana
Lewis, Mrs. KATHERINE B., ’89...“Elmstone,” 656 Seventh St., Buffalo, N. Y.
OAS BRP ABN VERIO Jorn oo oiaio's oS D 2-6 / halo ckeca 5;v wisle ob) 5hs 406 Galena St., Dixon, Il
Ln SDSS Tia i Oe Okla. Ag. Exp. Sta., Stillwater, Okla.
Partarenesvuare cA MT). 200% occ accoh8 cn Sale se cceile daths Nashville, Tenn.
REE NGED ET © O20 225 hi suies cities A aioe 48 Cumberland St., Rochester, N. Y.
LoncFELLoW, Ropert Capes, M.S., M.D., ’11........ 1611 22nd St., Toledo, O.
oOo SE Re 01: Saat 6 ae eo North Gorham, Maine
Dyvonstlowsarp N., MiD.784.20..6.......- 828 N. Wheaton Ave., Wheaton, III.
MacGIitvrAy, ALEXANDER D., ’12....603 W. Michigan Avenue, Urbana, III.
Won BEC UE [Bye A ee Se ee ee ee Decatur, Illinois
Marr, GeorGe Henry, M.E., ’11............. 94 Silver St., Waterville, Maine
NUR SIEATE > COREING. MD). QO ., o:c00c.0;«.0 2:0 2507 Penn. Ave., Washington, D. C.
NPARSITATE RUTH PHD. O75... cs. noc ees Rockford College, Rockford, II1.
WISE SHATT AO LS) en JT 2.6 oo lcti be 139 E. Gilman St., Madison, Wis.
ESSERE ERDR VA. bs ea Sei’ oto ccm owas eS Clemson College, S. C.
WeAsree eee MDL Phe O2 <0 -.2cx28 46 Warren Ave. East, Detroit, Mich.
MAYWALD, FREDERICK J., ’02........ 1028 Seventy-second St., Brooklyn, N. Y.
MeGarra sAveern PhiD:, ’80. 2s5:..2006 55 2316 Calumet Ave., Chicago, III.
MinGerrgre tory Pe P eho HS ats hs Univ. of Nevada, Reno, Nevada.
MI aNADD pent eTocs ore AS eee ORs Pe fee Glen Rose, Texas
ETA Ve OS EPH OA ba tegag tr hips cided Vane 259 Eighth St., Troy, N. Y.
McKeever, Frep L., F.R.M.S., ’06........... 429 Pender St., Vancouver, B. C.
Seemnriere Oy teeon ME FORMS 2s cece wre cue csr ewes vacessteess

= se hc Oe Oo e 324 Montgomery St., Syracuse, N. Y.
LU IRVERTES ijl Seal Ed & OL DB 0 6 eine sea 200 E. State St., Athens, Ohio
Metcatr, Pror. ZENO P., B.A., ’12........ A. & M. College, W. Raleigh, N. C.
PPE MME IONUA RED OG os vs aciosocesscececses ene’ 18 W. 27th St., New York City
Miter, Cuartes H, ’11...... Med. School, John Hopkins U., Baltimore, Md.
Mitter, Joun A.,Ph.D., F.R.M.S., ’89........ 44 Lewis Block, Buffalo, N. Y.

Marr en amor CPG: 3 MED) bie sa. cscs a sc: 403 Ray St., Seattle, Wash.

308 LIST OF MEMBERS

MINEHART, ProF. VELEAR Leroy, A.B., ’11..2070 Rosedale Ave., Oakland, Cal.

Mokewmrnst Ha Say lO! foo oa ewtews Ges 620 First Natl. Bk., Lincoln, Neb.
MornrEn HMDS 7072562 sec desu 341 W. Fifty-seventh St., New York City
Moopy, Rosert O., M.D., ’07...... Hearst Anat. Lab. U. of Cal., Berkeley, Cal.
Morris \GAPEG 712) ..2.)6 <8 Leafield, Gibsons Hill, Norwood, London, S. E.
NSULTENTS OUR AG, PID aia ee a Lawrence College, Appleton, Wisc.
MurreeE, Pror. E. H., A.M., LL.D., ’12...... Brenau College, Gainesville, Ga.
MiERS MRAM Et ate 1s)Asciocs sictiehse pose ee 331 Market Street, Bethlehem, Pa.
Noni Wimrrane C25 BY Bd f71ghtce Bak. so ata teens McCool Junction, Nebr.
INORRBIS: APROF EVARRY WALDO} TTS Sek Sb s0n 816 East St., Grinnell, Iowa
Norton: -GHarrrs. Eve), 21ae sohes bhaces 118 Lisbon St., Lewiston, Maine
INGURSEAMGEORGE DY 205 tea vee dc ceo aera ein bee cise coer Charlestown, N. H.
OGEEVEENG 25) 1B'S.6 Sc Diiss oe. ee ree 1006 N. Union St., Lincoln, Ill.
QR AWEI OU. oc citeia a stasiel ca eee mmaieroesiee 216 Spring St., Portland, Me.
Osh ea ving poe eed BY Se (2) ON cee ee ae oe tee e Eo Granville, O.
OQROETA MD OMINGO WE STS se weeks. 2 Seo el tetra eee oe Gijon (Asturias), Spain
Ossorn, Pror. Herpert, M.S., ’05..... Ohio State University, Columbus, Ohio
One EH ARVER tN ALM Oaths: ices SSRs Spencer Lens Co., Buffalo, N. Y.
IBATMER S HOMAS: CEPAD REVS Eh) Se Ut soe ss. foals sce Media, Pa., R. F. D.
ParKErR, Horatio N., ’99........ College of Agriculture U. of I, Urbana, Ill.
PARsSErE sy HIENRY VivAc; *AGMG, 29RD. fesise cn 122 West 8&1 St., New York, N. Y.
PérRick) PRANK) PhD) Ol. Go. sate ans 603 Dwight Bldg., Kansas City, Mo.
PEASEORREDUN 5) Q7 aera ee eee ete eoies sates eee P. O. Box 503, Altoona, Pa.
PEASTEE? (PRoE LEON (DS PhDs. 71. 252k... Public Museum, Milwaukee, Wis.
BENNOGK | EE BWARD #270. 52264502 scsi 3609 Woodland Ave., Philadelphia, Pa.
PETERSON, NIELS FREDERICK, ’II..........2.000008 La. S. U., Baton Rouge, La.
PEEAGMAUNDAGNUS$:ESO4 7 OTA sey hak ccs ck Cone a eee Meadville, Pa.
PEEUPSS ESSIEN SECTS: Oe so Sascnenel ene 16 N. Summit, Ypsilanti, Mich.
Bais, Epweay; 20 SSB 8 eC Sea oso ok Sao nis hee

...Madeley House, Bulstrode Way, Gerrard’s Cross, Bucks, England
Potvarp, Pror. J. W. H., M.D., 712. Washington and Lee Univ., Lexington, Va.

Pounp, Roscor, A.M., Ph.D., ’908..... Harvard Law School, Cambridge, Mass.
BGWEnSr Eh wb A Birra Beek he ete wilieeiceseutires Waxahachie, Texas.
BarenepeRor sOrro as.) MEDIV 6 Tr.ceceene 5 and 6 Fedl. Bldg., Laramie, Wyo.
BEINGS SOME RED O9e fees ccc 8 ok eos Beas 510 E. Clark St., Champaign, II.
IPYBURN GEORGE IMEI 86>. 5. ccc ce ccseesecceet torr H St., Sacramento, Cal.
Qomiman, MaAgvin ©, “AJM. "13-0003... clas: Wesleyan Col., Macon, Ga.
RAwern: WiAETER@ Vie E35. ck. s.0c 5 = Princeton University, Princeton, N. J.
TRUANSTER: Jbt Bic a Onn et nied | 5 tote ce es 203 Seneca St., Manlius, N. Y.
ReAwsott: -Beavron Eis QME 6 $2 toa fos eon os swans Abt < Jeni eee ee
Set Lee U. S. Bureau of Animal Industry, Washington, D. C.
RASorren. 0.2 Me ae ak seals Lbs dS weed) st eee Clinton, Conn.
Ray, Benya Hse s 335c5.:k castes Pine Grove, Schuylkill Co., Pa.

Raymonp, Rev. R. I. M.A., 713-........ Univ. of the South, Sewanee, Tenn.

AMERICAN MICROSCOPICAL SOCIETY 309

Rector, FRANK Les.iz, M.D., ’11.......... 36 Forty-first St., Brooklyn, N. Y.
FEES SV EATTRECE MET os iether wit Sob te a eM i ne eek kh) em & Tarkio, Mo.
Reese Paar oALeems Mt PD) ((Hop.); 7052. 65 2 6dieies eae es ei ee ees

ay NAMEN EE ones Od ale) Dake eis Sits W. Va. Univ., Morgantown, W. Va.
RGD PPA BRED See ete, reece bere Kuala Lumpur, Selangor, Fed. Malay States
IRGeEA VW EEErAaiEt. AMS Taio oactic oid gor College Avenue, Wheaton, II.
RismaRnS ee Aime. Mehr ose tan oe gee lue University Station, Austin, Tex.
IRATGUARD So EUTTAIS QO cid/e-e dad sic'e.cieieresevercreisiere 1114 Floyd St., Lynchburg, Va.
RIcHARDS, FREDERICK WM., ’I1....212 Notre Dame St. West, Montreal, Can.
ROBERTSMMMONVAT TTS? SET dsc) dete lcls-e:<aiepeicie eigcslacre 65 Rose St., Battle Creek, Mich.
Peer ese oars TD e wie eb 8 bviea. encase 345 West Michigan St., Chicago, III.
Rocers, WALTER E., ’11..... 24 Columbia Ave., Univ. Heights, St. Louis, Mo.
Ross, LurHER SHERMAN, S.M., ’11............ 1308 27 St., Des Moines, Iowa.
NA Te ane VA Mis AERC rE Es Sy 2.0 ig aida oii eed BAIS Hudson, Ohio
SAWYER, WILLIAM Hayes, JR., ’13.......... 18 Arch Avenue, Lewiston, Me.
Scott, GrorGE Fitmore, A.M., ’13.College City of New York, New York, N. Y.
PUREE RIPON yg Uk erie S28 Sia, ai ahan piv n.d nee aie eg eae Univ. of Wyo., Laramie, Wyo.
SUSI Sf He Ll Dig) 2 0 8) D ead oe ae Bureau Plant Industry, Washington, D. C.
SEDARER PRORORS Plot WVVis OS). 5 a oholsis cierto 5 wos aus 168 Parkside, Brooklyn, NY.
PMR UME RE ee 2 is. (a ess ciS wide Gis ieravelbe. anisicea ws 809 Adams St., Bay City, Mich.
SHERIDAN, Wo. F., ’05....U. S. S. Flusses, (care Postmaster, New York City
STEER ANU SENG ETN uve (AT) ch saloalacabotememeteenes Homer, Minnesota
Sirs Urea Crs UNS tS ba) eee Seventh St. Docks, Hoboken, N. J.
Siniyiane, TE, TREN rnin ge Son a ae ae 902 Pine St., Philadelphia, Pa.
Srocuse (CHas. E> PhDs M:Di 778) oe... oe ee oe 218 13th St., Toledo, Ohio
SMUATMMLIOWIARD! 9120.5). 5 o.cia cscs oloteie esos 203 West Avenue, Jenkintown, Pa.
Smirn BreeTpaAm G. Ph.D,, 713.0... 936 Forest Avenue Ypsilanti, Mich.
Situ, Pror. Frank, A.M., ’12.......... 913 W. California Ave., Urbana, III.
RRM tS) OOo as isis os ca kdes deiddies 131 Carondelet St., New Orleans, La.
SOR enn a ER NISG 8 G75 sts iia San ok en kee Jae Up Re Ma eee Mn

a ee ae 37 Dryburgh Road, Putney, London, S. W., England
Spautpinc, M. H., A.M., 713...... 508 W. College Avenue, Bozeman, Mont.
SPuRGEON, CuHar_es H., A.M., ’13..... 1330 Washington Ave, Springfield, Mo.
Sepp, te Ee PhD, M.D 708.323. 65.252 50 E. gist St., New York City
STEVENS mBO Reb sollte eee Siar 2) eine A ee aed Seren nee NY

A SAAR ce Sone ae earn Agricultural Experiment Station, Gainesville, Fla.
STEWART, MAupE GERTRUDE, B.L., ’12...... 410 Winona St., Northfield, Minn.
OOP EME: GrgNy. S25.) BB sd os osels bees das dade delsicle 3 widies Monroe, Mich.
StuNKARD, Horace W., B.S., ’13........ Nat. Hist. Bld. U. of I., Urbana, Iil.
STURDEVANT, LAZzELLE B., A.B., B.S., ’03..... Univ. of Nebraska, Lincoln, Neb.
RMT See ROR Wel teh a AO) eS Als 6 = oi vo oma nig: 6 haehnteeoee eR eee eee Ames, Iowa
SVEMMENPRN MGRORGE? 120) os 1 a Sclgsieles cus asic: P. O. Box 1110, Montreal, Quebec
SwiInc_e, Pror. Leroy D. ,’06.......... Univ. of Utah, Salt Lake City, Utah
Tuomas, ArtHurR H., ’gg........ Twelfth and Walnut Sts., Philadelphia, Pa.

WAMOMIN S» GEORGE; O0i. 02. Mee deal nee owt s 1410 E. Genesee St., Syracuse, N. Y.

310 LIST OF MEMBERS

‘Fons; jase C., Bik Os Fee ois cedcestl Sees eee. Boulder, Colo.
TRENNER, SIMEON, ’I2..... 6328 Jefferson St., Germantown, Philadelphia, Pa.
Tsou, Yinc-Hsuan Hsuwen, M.S., ’13..P. O. Box 78, Univ. Sta., Urbana, III.
‘Thorne: Guar Gs; Avo .2 50S ei 173 Wood Street, Lewiston, Me.
Durrmx? Pros: Az FMD: PRS Pia. so Fossa has cd iwsne cake se Seaeoeeee
Neg Tes oes eee ate University of Virginia, Charlottesville, Va.
‘Termes: 2.5G: Tamer, 71965 kee Park Rynie, Natal So. Africa
Varearein? Pinenrer ©) rao siivicevinksenaase 141 Milk St., Boston, Mass.
Vite CEBAve: IARDEY (Js). "PE0E Gs es See ak 310 N. H. Bldg., Urbana, Ill.
VARRELMAN, FERDINAND A., 7I13......-00.00000: P. O. Box 134, Columbia, Mo.
Wiatre: UPRepenrcr (Ges iPh Do: Tirso d. ade csdencse sleo oa ee ve ae ee eee
dibs Medical Department, Western Reserve Univ., Cleveland, Ohio
Waren Bena (R: PRAD 07. Seccck cn University of Nebraska, Lincoln, Neb.
WALKER SPRvAy BRUER Uitved. vis. Siseaw as aoe Sea Station A., Lincoln, Nebr.
Wa ker, THappEus, M.D., ’I1.............- 33 East High St., Detroit, Mich.
Warton: hervB ¢) Aev: PD 270532 oh ed eek Kenyon Coll., Gambier, Ohio
WER Is IR See cdesine seca eves e aaa 306 E. 43rd St., Chicago, IIl.
Warp, Henry B., A.M.. Ph.D., ’87......... University of Illinois, Urbana, III.
NGATSON WORMCEYDE SoMIL TT Ses )..c bob aca bu 6 saidchwe See Te Kingwood, W. Va.
Wee ci) an OO Ps SGT 5 Oa ss Sa saw Se Box 416, Fergus Falls, Minn.
Wier PARE PSR eens 28k. Pao) savas. Kas. St. Ag. Col., Manhattan, Kas.
WELUMAN: Prom i. CariGuTon, M:D:. 605. 60s saw ee eee eee
Betas dc tutem chinks Med. Dept. Tulane University, New Orleans, La.
Witness) 82°49. PHD 5 ees iss coe sa chic 79 Chapel St., Albany, N. Y.
WHe -p tey, H. M., M.D., Ph.G., F.R.M.S., ’09..2342 Albion Pl., St. Louis, Mo
Wire Cs AS EE OSes os See Center Sandwich, N. H
Wuirtney, Geo. ANSEL, B.M.E., ’I11........200+ 235 Main St., Lewiston, Maine
Wieman, Harry L., Ph.D., 713...... University of Cincinnati, Cincinnati, O.
Wrrirdwison, Wa, FBS BS 7 i i 95) oR ee
Ui Nabicivin Ae hee 9 Plewlands Terrace, Edinburgh, Scotland
Wotcott, Ropert Henry, A.M., M.D., ’98...Univ. of Nebraska, Lincoln, Neb.
ra eae W; 07 ic Bet Arce cat ee aes (Westmount) Johnstown, Pa.
Weicet, Srantey‘'H. C. E., 13... 112’N. Broad Street, Philadelphia, Pa.
ZAPFFE, FREDERICK C., M.D., ’05...........-. 3431 Lexington St., Chicago, III.
Zmss, Gass, (care Dr; M. von. Rohr) .< 3.6.5 200. cae ee ees Jena, Germany
ZOO: QORVID INT BLO Oly toes wok eee odie Dodo 59 Clark St., Chicago, Il.
SUBSCRIBERS
ACADEMY OF NATURAL SCIENCES............. Logan Square, Philadelphia, Pa.
AmeERIcAN Musrum or NatoraL History,....:.......:...02 500s soe
Uns. JSS ee CaS Re eee eet 77th St. and Central Park, New York, N. Y.
Bascocx .Screwrerie Linkane S. Siali evi <4 eae Plainfield, N. J.
BIBLIOTHECA DE FACULTAD DE MEDICINA...........---- Montevideo, Uraguay

~~

AMERICAN MICROSCOPICAL SOCIETY 311

BECKFORD: DIOLOGICAT:ISTBBARYS 0.5.65 ck atac eens dt Bates Col., Lewiston, Me.
OST NE BING TR ote rs 0c FSeuele hid iad PRR Aes ORE Boston, Mass.
Boston Socrety oF NATURAL History............ Berkeley St., Boston, Mass.
BRooKLYN INSTITUTE OF ARTS AND ScIENCES..Lafayette Ave., Brooklyn, N. Y.
BUREAU ORASCIENCE) IATBRARY . 5 o.0's bs dete o's «eds van doicwieas cama e oo Manila, P. I.
EUR GESSAMUN Ee ere reed mrad are loleveicicraiain'e eee oa’ 126 Summer St., Boston, Mass.
Cageemay pi A es enol e ss Casilla de Correo-69, Montevideo, Uraguay
annrcrrsERrMeTRRARN ssc ialseecs< ssc 6d ceies sce cieeasbaeuaoerd Allegheny, Pa.
NEI A PURER oes chy eon isles al che. soleil aiaiene-a ofsie din vb wd aledee Pittsburg, Pa.
Grrenses: Grp BRARY «2 oc0.cvec0. 0: ecere; sire 52 East 41st Street, New York City.
mmreA Cee UNIVERSITY, LIBRARY... <.00ep «ois 04's ose san ve shedbesawcine Chicago, Ill.
CopuRN Lrprary OF CoLoRADO COLLEGE......... Colorado Springs, Colorado.
Earn VaR GOMPEGE IGTRRARY)-notc emis tis ay ssuclevencieise ais.5 6 sles eels @clsretoe Waterville, Me.
CoLLEGE OF PHYSICIANS LIBRARY......... 19 S. 22nd. St., Philadelphia, Pa.
Gere ime Stare NORMAL: LIBRARY... «.....s)cicccncelesiosecseeseeccws Greeley, Colo.
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PARP ERE ROS AO). Scr. pee fo cto aos alias oe ahs P. O. Box 174, Capetown, S. Africa
DecaTuR TEACHERS’ PEDAGOGICAL LIBRARY........ Public Schools, Decatur, II}.
Merete Rte ORTH TBRARY. 92 xic.c)cc baie © s wieveimwis'e Hortite sls oe ee baies we Detroit, Mich.
Ana UUINIVERSTR Ye ILTBRARY: 2 2:s.0005 a sie owen as doae ere maine Des Moines, Iowa
ES Ee 37 Soho Square, London, England
Reece ee DA Te MITISELIME 5. 5:5:0:0)210.0 sc0:ei0 ie 001 Pideld ple eoulale’s oe Chicago, Ili.
FRANKLIN & MARSHALL COLLEGE LIBRARY.............000000 Lancaster, Pa.
LETTS TL UU Ge ee ye ee 159 East 61st St., New York
2 ES a a ear ee Westboro Insane Hospital, Westboro, Mass.
(ELAR Wi VA ne 4575 Forest Park Blvd., St. Louis, Mo.
iE Aare MINTG MPA BO) BANE ORSV a. ahercintn oar oe anole 'e one os ola wis)s eee te a Burlington, Vt.
ILLtino1s STATE LABORATORY OF NATURAL History LrIpBRARY........ Urbana, Ill.
LOWALSOTATE (UEACHERS. GOLLEGE. LIBRARY... 20.42. 2 eh dee Cedar Falls, Ia.
Towa STATE CoLLece Liprary (Pror. PAMMEL) ....Station A. Ames, Iowa.
jameeseMirrrkin WNIVERSITY JLIBRARY .5< sds cisccce se Seceicle © ocraye Decatur, IIl.
epee EDA ITER AUN 0 oy. w bce. wv praia Sersio eo nda AR a ee eee Chicago, Ill.
WetNGHHOPKINS, UNIV: LIBRARY...c 65.3225 0ccc asec ees cane wees Baltimore, Md.
IGAWSAGS STATE AGRa COLLEGE. LIBRARY... 0.:.20- +b be nine es Manhattan, Kas.
[GARORATORY, OF FHIISTOLOGY AND EMBRYOEOGY. . 32.6... 00% coeccccces sees

TR SE University of Minnesota, Minneapolis, Minn.
Paeene OTANFORD, Jr, Univ. LImmaRy. 23.5252 022 50s cee ce nes Stanford, Cal.
L’InstirutTo OswaLpo Cruz (CHEZ Mr. A. SCHLACHTER)...........20055

BET baton Rae Bows cnet 20 Rue des grandes Augustius, Paris, France
PSOE E DOT ANICAE. GARDEN. cose 0.00% ses eos aclesvasvccases St. Louis, Mo.
Mourners HonvOKr) GOLPEGESLIBRARY: 3... s sss [ss cea t v's 0s South Hadley, Mass.
Museum CoMPARATIVE ZooLoGy (HARVARD).........---+-- Cambridge, Mass.
MUSKINGUM (\COLUEGH: ISTBRARYs -s\isiias hive bets cscs eens New Concord, Ohio
INE WeLLAMPSHIRE ‘OTATE UCTRRARY <..2 nox «\.siacis's oc eesiee ceieen Concord, N. H.

312 LIST OF MEMBERS

New: Yorr Pupric Limrary 5.553.000 ...476 Fifth Ave., New York City
NS Yow STATE LIBRARY: 4. oi 'oc eeeuied dcaves Serial Section, Albany, N. Y.
Ome Sranenrvensery (Lime ary., svc. Nee Se aoe ake 2k Columbus, Ohio
Oni1o WESLEYAN UNIVERSITY LIBRARY..............--00000- Delaware, Ohio.
QurENS CUNTVERSITY JETRRARY 31)<).sA2t civraniceion Meee De eins Kingston, Ontario
SOuen rivie LApeaRY os 350 5 2 8 Se ek ..U. S. Patent Office, Washington, D. C.
SmitH COoLLece LIBRARY........ bE Sad We kao oacniee eRinee Northampton, Mass.
SVRAGUSE shUBIIG (LABRARY s, 6 decd clsiesissd oer chiele etek 3B ya oats: Syracuse, N. Y,
U. S. MepicaL MusEuM AND eee Bou. Gen.’s Office, Washington, D. C.
UNIVERSITATS BIBLIOTHEEK...... etapet rae eats ete neo cere Amsterdam, Holland.
WININERSTEY > OF WARIZONA ST LTBRARVE" cs.ic. aataite,srohs net Me then ee ee Tuscon, Ariz.
Unirvensity or Carrronwea)-Lippary, obec Sage cle. ones Berkeley, Cal.
University oF Missourt LIBRARY.............. poe s eae Columbia, Mo.
UINTVERSITY LOR MVIONTAN AS STBRARYE os s/cie cicte ce sainiee contac Missoula, Mort.
UINIVERSTEY OR SINEVADAOICIBRARY 2.525sc:ctos oa Sete ice ee oa aoanae Reno, Nev.
UnIversity oF NEBRASKA LIBRARY..... SASS ee ee Lincoln, Neb.
UNIVERSITY OF TENNESSEE Lrprary..College of Agriculture, Knoxville, Tenn.
UNIVERSITY OF VIRGINIA LIBRARY.........0:0.-00000: Charlottesville, Virginia
VANDERBILT UNIVERSITY LIBRARY..... S Sata slider eee .....Nashville, Tenn.
WASSART COLLEGE MILIBRARYS <cicnc 5 66 tens se Saini vee Poughkeepsie, N. Y.
WASHINGTON STATE COLLEGE LIBRARY....... a paieenee Pullman, Washington.
WESLEYAN UNIVERSITY LIBRARY....... eo kiki Dialga Middletown, Conn.

INDEX

Algz, Periodicity in Illinois, 31
Alternation of Generation, 137

Ameba with Tentacles, 79

American Rhizopods and Heliozoa, 79
Amutosis in Onion Root Tip, 103
Amitotic Division in Ciliated Cells, 199
Aubert, A. B. Memorial of 84, (159)

Beginners Guide to the Microscope

(Book), 82
Black Moulds—Lena B. Walker, 113
Bog Selutions and Plants, 7
Book Reviews, 81, 156, 205

Cephaline Gregarines of the New World,
259

Chinch bug parasite, 301

‘fhromesomes, Size and Phylogeny of,
79

Cilleuls, J., Intersitital Cells of Testis
and Secondary Sex Characters, 81

Classification of Plant Rusts, 41; of
Powdery Mildews, 238; of Cephaline
Gregarines, 262

Collecting Plant Rusts, 44

Collin, An Ameba with Tentacles, 79

Compound Eye, Growth of, 140

Cort, Wm Walter, Notes on the Trema-
tode Genus Clinostomum, 169

Crandall, Dr. Geo. C., Memorial of, 84
(159)

Custodian’s Report of Year 1912, 86

Darling, Staining Protozoa, 74

Diatoms, How to Clean, 73

Differentiation of Organs of Sex, 134

Diptera, Peculiar Sense Organs of, 69

Disease Producing Fungi, Dissemination
ro) a

Dissemination of Fungi
sease, I*. D. Heald, 5

Causing Di-

Diversification of Parents, 135

Edmonson, C. H. & Kingman, R. H.,
Notes on Japanese Protozoa, 93

Ellis, Max M., List of Cephaline Gre-
garines of the New World, 259

Entomological Notes, 300

Erysiphacez, 219

Experimental Amitosis in Onion Root
Tip, H. E. Jordan, 103

l'reshwater Hydroids, 146
Fund, Spencer—Tolles, 165
Fungi, Dissemination of, F. D. Heald, 5

Gall Midges, adaptations in, 300

Galloway, T. W., Summary of Ele-
ments in the Reproductive Process,
127; Report of the Secretary, 213.

Grasshoppers, 302

Gregarines of New World, 259

Growth of the Compound Eye, 140

Habitat and Distribution of Plant Rusts,

A2
Hance, R. T., A Paraffin Ribbon Car-
rier, 207

Heald, F. D., Presidential Address, Dis-
semination of Fungi Causing Disease,

~

FS
Heliozoa and Rhizopods of America, 79
Household Bacteriology (Book) 156
House-fly, Hibernation of, 301
How to Mount Rust Spores, 47
Hyatt, J. D., Memorial of, 84
Hybrid Pigeons, Spermatogenesis in 80
Hydroids, Freshwater, 146

Illinois Microscopical Society, Notes
from Meeting, 75

Index, 313

insects and Disease, 300

Insects and Dissemination of Fungi, 20

Intersitital Cells of Testis and secondary

Sex Characters, 81

314

Japanese Protozoa, Classification of, 94

Japanese Protozoa, Notes on, 93

Jordan, H. E., Experimental Amitosis
in Onion Root Tip, 103

Kern, Frank D., Nature and Classifica-
tion of Plant Rusts, 41

Kingman, R. H. and Edmonson, C. H.
—Notes on Japanese Protozoa, 93

Latham, V. A., (Sec’y.) Notes from
Meeting of Illinois Microscopical So-
ciety, 75

May-flies, biology of, 301

McCraven, Bonner N., Memorial of, 160

Meek, Size of Chromosomes
Phylogeny, 79

and

Meeting of the Illinois Microscopical
Society, Notes from, 75

Microbiology in Relation to Domestic
Animals (Book) 81

Microscopy and Drug
(Book) 83

Mildews, Powdery, 238

Minutes of the Cleveland Meeting, 85.

Moulds, Black, 113

Nature and Classification of Plant Rusts,
41

Necrology, 84, 159

New Technic in Staining Diphtheria
Specimens with Toluidin Blue, 75

Notes from the Illinois Microscopical
Society Meeting, 75

Notes of Japanese Protozoa, Edmonson
& Kingman, 93

Notes on the Trematode Genus Clinos-
tomum, W. W. Cort, 169

Notes, Reviews, etc., 69, 127, 183

Examination

Onion Root Tip, Experimental Amitosis
in, 103

Paraffin ribbon carrier, 297

Parasites of Chinch bug, 301

Parasitic Rotifer in Egg of Water Snail,
78
7

INDEX

Parasitology ; Laboratory Guide (Book)
205

Periodicity of Alge in Illinois, E. N.
Trauseau, 31

Phylogeny and Size of Chromosomes, 79

Physiological Effects of Peat or Bog So-
lutions on Plants, 76

Plant Rusts, Nature and Classification
of, 41

Powdery Mildews, George M. Reed, 238

Presidential Address, 5

Prevention and Control
(Book) 206

Potts, Edward, Freshwater Hydroids,
146

Proceedings of the A. M. S., 85

Production of Sex Organs in Algez, 37

Protozoa, How to Stain, 74

Protozoa, Japanese, 93

of Disease

Reed, George M., Powdery Mildews, 238

Report of Secretary and Editor, 213

Reproduction Among Fungi, 6

Reproduction of Black Moulds, 114

Reproductive Process, Summary of the
Elements in, 129

Rhizopods and Heliozoa, American, 79

Roberts, E. W., The Growth of the
Compound Eye, 140; Four Notes, 183

Rotifer Parasitic in Egg of Water
Snail, 78

Secretary's Report, 213

Sense Organs of Diptera, 60

Smith, Spermatogenesis in
Pigeons, 80

Spencer—Tolles Fund—by the Commit-
tee, 165

Spermatogenesis in Hybrid Pigeons, 80

Summary of the Elements in the Repro-
ductive Process, 127

Synopses: Plant Rusts, 41; Powdery
Mildews, 238; Cephaline Gregarines,
259

Hybrid

AMERICAN MICROSCOPICAL SOCIETY 315

Table of Measurements of Clinostomum, Wailes, Some American Rhizopods and

180 Heliozoa, 79
Technic, 75, 204 Walker, Lena B., Black Moulds, 113
Toxic Secretions of Infusoria, 198 Water Snails, a Rotifer Parasitic in Egg
Transeau, Edgar Nelson, Periodicity of of, 78

Algz in Illinois, 31 Welch, Paul S., Entomological Notes,
Treasurer’s Report for 1912, 87 300

Trematode Genus Clinostomum, Notes

on, 169

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