-- -VoL2- JANUARY, 1945 ——s“No.l_~

FARLOWIA

A Journat or CryprocamMic Borany —

ANNIVERSARY NUMBER >
COMMEMORATING ©
THE ONE HUNDREDTH BIRTHDAY
OF
WILLIAM GILSON FARLOW
1844-1944

ay PUBLISHED BY
FARLOW LIBRARY AND HERBARIUM OF HARVARD UNIVERSITY
-20 Drvinrry AvE., CAMBRIDGE, Mass.

FARLOWIA
Published semi-annually. Four numbers compose one volume of scotorintitely five
hundred pages.

Subscription prices: Vol. 1, $5.00; Vol. 2, $7.50. Single numbers: Vol. 1, $1.50;
Vol. 2, $2.25.

Subscriptions and remittances should be addressed to FARLOWIA, 20 Divinity Ave.,
Cambridge 38, Mass., U.S. A.

Numbers lost in the mails will be replaced if reported within a reasonable period.

Epiror1AL Boarp
DAVID H. LINDER, Editor

E. V. SEELER, Jr., Managing Editor -
J. R. BARTHOLOMEW R. SIncER
E. B. BARTRAM F, VERDOORN
N.F.Conant W.H. WEston, Jr.
H. F. Harris R. M, WHELDEN

CONTENTS OF VOL. 2, NO. 1

In Honor or Wirt1am Guson Fartow. By David H. Lie Boe Cg a ay

THE CORRESPONDENCE OF WILLIAM G. Fartow purine His StupEnt. Dae AT
SYERABHOURG, Hittnuaacr. degeres 8 o.. Op eg a Rae ee
NoTEs on Fartow’s AGARICALES FROM CHocoruA. By Rolf Singer . . . . 39
Wittiam Gitson Fartow: Promoter or PuycotocicaL RESEARCH IN AMERICA,
ied4-1919.. By Won. ancora Tavior. 6 ea ee BB
FarLow’s INTEREST IN AN INTERNATIONAL ABSTRACTING JOURNAL. By Frans Ver-
PIN RF GO pO we the a 4%, kk en So ee eee e OCE ae
_ Dr. Fartow’s INFLUENCE on Mycotocy. By William H.Weston,Jr.. . . . 85

THE BOLETINEAE OF FLORIDA WITH Notes oN EXTRALIMITAL SPECIES. I, STROBILO-
BEVCETACEAR: DY AGN) SON geR ee ee ee OF

Vol. 1, No. 4, was issued on July 18, 1944.

&

FARLOWIA

A JoURNAL OF CRYPTOGAMIC BOTANY

VOLUME 2
1945-1946

PUBLISHED BY
FARLOW LIBRARY AND HERBARIUM OF HARVARD UNIVERSITY
20 Divinity AvE., CAMBRIDGE 38, Mass.

con

Rhodophyllus Farlowii Singer

L. C.C. Krieger pinxit See page 49

FARLOWIA

A JOURNAL OF CRYPTOGAMIC BOTANY

Vor, 2 January, 1945 No. 1

IN HONOR OF WILLIAM GILSON FARLOW
Davin H. LINDER

Although this journal is but two years or one volume old, it is never-
theless not too young to recognize the one hundredth anniversary of the
birth of the man who, through the founding of the Farlow Reference Li-
brary and Herbarium of Cryptogamic Botany, made possible its existence.
The occasion is of greater importance than the mere fact of the journal’s
existence, for Dr. Farlow was more than a means of making its existence
feasible, he was the first professor of cryptogamic botany in America and
played an important part in the development of this branch of botany. Not
only did he stress the importance of systematic cryptogamic botany and
morphology, but he initiated in this country the teaching of what is now
known as phytopathology and he also began, with Professor W. R. Nichols,
the study of the réle of algae as impurities in drinking water. His versa-
tility and his contributions have been ably and more amply described in
the biographies appended to this paper and for this reason only a brief
sketch is needed on this occasion.

William Gilson Farlow was born in Boston on December 17, 1844 and
there spent the first fourteen years of his life before the family moved to
Newton, Massachusetts, then considerably less urban than it is today.
Perhaps it was the life in the country where he was surrounded on all sides
by Nature’s beauties and curiosities that stimulated his enquiring mind and
influenced him to become a botanist or maybe it was the influence of his
father, John S. Farlow, who although a successful business man, was
equally enthusiastic about horticulture, as is attested by the prizes that he
won at the exhibitions of the Massachusetts Horticultural Society and by
the gifts that he made to that institution. Whether it was the father’s in-
terest in plants or his own discovery of the wild Hepatica, which has been
said to have stimulated him to pursue botany as a life study, there can be
little doubt that William Farlow received the encouragement and backing
of his family, and that his father by example set the pattern that he was
to follow.

While at Harvard College, from which he graduated in 1866, he contin-
ued his interest in natural history, taking the courses offered by Asa Gray
and being elected president of the then active Harvard Natural History

1

2 FartowiA, Vor. 2, 1945

Society. On graduating from college, he spent four years at the Harvard
Medical School where he obtained his M.D. degree. He did this, even
though highly successful as a student, not because he was primarily inter-
ested in medicine, but because in those days botany was the hobby of
amateurs who were for the most part physicians and ministers and there
was little opportunity for the professional botanist. To quote his words
(Science N. 8S. 37: 79. 1913) “When I graduated from college in 1866 and
wished to become a botanist, Professor Gray told me that I ought to study
medicine first because the possibility of gaining a living by botany was so
small that one should always have a regular profession to fall back upon.”
. . . “In 1866, there were very few botanical professorships in this coun-
try, the salaries were very small and the equipment very shabby. Gray was
professor at Harvard, D. C. Eaton at Yale and Porter at Lafayette. Tor-
rey, in spite of his distinction as a botanist, really depended on his position
as a Chemist for his living. The comparatively few positions in government
and state stations offered few attractions and changes were frequent. To a
young man the prospect was not assuring.”

Upon graduating from medical school, Farlow returned to botany, be-
coming Asa Gray’s assistant. He began in earnest to pursue his studies of
cryptogamic botany but after two years, found that if he wished to pro-
gress in this field, it would be necessary for him to go to Europe. Hence, in
1872, he went abroad and spent the two following years, for the most part
in the recently founded laboratories at Strasbourg under the guidance of
Anton De Bary, a brilliant crytogamic botanist whose scientific conclusions
were often in advance of his time and whose teaching appears to have been
most provocative. But despite the fact that he was learning De Bary’s
methods of teaching and research, he nevertheless, on the recommendation
of Dr. Gray, took out time for intensive studies of the lichens with Dr. J.
Mueller and of the algae with Thuret and Bornet. The trips, if we may
judge by his letters, gave him an opportunity to visit many important
herbaria in different parts of Europe and to make the acquaintance of the
distinguished botanists of the period, acquaintances with whom, after his
return to America, he maintained an active correspondence. His stay in
Europe also provided him with an opportunity to obtain rare and important
volumes for his library and equally important specimens for his herbarium
to which he added while still abroad, the most important Curtis collection
which contained parts of the types of species described by Berkeley and
other specimens obtained by exchange with Schweinitz, Ravenel, Sprague,
Fries, and Duby. Thus in spite of being small in stature, a fact about which
Dr. Farlow was very sensitive, he was a giant when measured by his physi-
cal and mental activity.

Although worried about the prospects of a position as a professional
botanist on his return home, when he did come back to America he was
appointed through the friendly efforts of Dr. Gray to an assistant profes-
sorship and five years later, in 1879, became Professor of Cryptogamic
Botany. During the early days he taught not only cryptogamic botany, but

LINDER: WILLIAM GILSON FARLOW 3

at the Bussey Institution where he was stationed the first five years, gave
instruction to special students in the applied side of mycology, probably
the first to be given in that now thriving field known as phytopathology,
to which he was then making contributions.

When he became Professor of Cryptogamic Botany he moved from the
Bussey Institute to Cambridge where he apparently enjoyed greater free-
dom of action and where he continued with zeal to build up his already im-
portant herbarium and library. Although still devoting his attention to
mycology, he devoted more time to the algae, his first love, and published
among other papers, the “Marine Algae of the New England and Adjacent
Coast,” a manual that has been in use up to very recent days. Although
interested in the morphology and taxonomy of the algae, for which his
studies well prepared him, just as with the fungi, he saw the practical im-
portance of the freshwater algae and published three or four papers which
resulted from his study of the water supply problems of the City of Boston.
Again, although primarily a systematic botanist, he was one of the first in
this country to demonstrate the importance of the lower forms of plant life
in the welfare of the people.

Little more need be said here on Dr. Farlow’s reputation as a mycologist
or phycologist since these subjects are more amply discussed in accompany-
ing papers. Less well known were his interest in lichenology and bryology,
perhaps because he published little on these two subjects. However, that
he did much work in these two fields is attested by a perusal of the speci-
mens in his herbarium where can be found a considerable amount of ma-
terial that had been named by him, as well as the critical notes and observa-
tions that were written in his characteristic and occasionally not too legible
hand. His remarkable memory and his very broad knowledge of these
groups enabled him to provide information of great value to students and
specialists alike, and his herbarium was available to those who were prop-
erly qualified. At times he has been criticized for not allowing the herbarium
to be used unless under supervision. This was not because of selfishness on
Farlow’s part but rather the result of abuse by thoughtless visitors who
ruined or even entirely removed some precious specimens from his collec-
tions. Farlow did not bring together the many specimens for immediate
consumption and dispersal, for his endeavors were all directed towards
building a reference collection that would serve future generations of
botanists who were capable of profiting by what he had done.

One cannot but be impressed by the tremendous energy shown by Dr.
Farlow. Although he gave up undergraduate instruction twenty-three years
before his death, he continued to take a keen interest in graduate students.
In addition to his teaching, he gave considerably of his time to answer
correspondence and through it to maintain the friendships that he had
made as he went through life; the administrative needs took no small por-
tion of his efforts, and withal he did a tremendous amount of research in
very diverse groups of non-vascular cryptogams. Yet somehow he found
time to make collecting trips inside and outside of the country, to attend

4 Fartowia, Vo. 2, 1945

meetings of the various societies to which he belonged, and to be present
at the International Botanical Congresses. It is small wonder that he had
little time for publishing the results of his own observations. In addition
to the activities indicated above, he was also interested in founding an
international journal in which could be found the names of new species of
plants that had appeared in the scattered literature during the short inter-
vals between issues, thus serving to lighten the bibliographic tasks of the
systematists and applied botanists as well. As papers were being published
at an increasing rate in widely scattered journals, it was even at that time a
problem to keep abreast of the current literature and only by personally sup-
porting an assistant and bibliographer was he able to maintain an index of
his own covering the fungi that had been described from North America.
This index, in 1919, contained some 300,000 entries and was available to
colleagues for consultation. The reason behind all these efforts seems best
to be explained by his own words (Science N. S. 37: 7. 1913): “I, who was
almost the first American student to seek the benefit of botanical instruc-
tion abroad, have lived to see the time when a very large number of bota-
nists have brought back to America the best that Europe has to offer. There
was a time when our botany might have been said to bear the mark ‘made
in England.’ In more recent years it may be said to have been ‘made in
Germany.’ There are some patriotic souls who hope the time will come, if
it has not already come, when we may say ‘made in America.’ I do not
share their feeling. To me it seems that botany is destined to become world-
wide and it will be enough if we can contribute our proper share to the
general stock.” Dr. Farlow died in 1919 before the intense nationalism that
grew up in the following years exploded into the chaos of our time. Let us
hope that when the present conflagration is over, humanity may have
learned the need of not only the scientific but also the political cooperation
that he hoped to see.

While still a student in Europe, Farlow realized the dangers of the mad
rush to every new fad that comes up in each generation to the neglect of
systematic botany, for he wrote to Gray “I have been to see Prof. Pfitzer,
who is a pupil of Hanstein, the man who doesn’t think it any consequence
to study the forms of common plants, but much better to mope over the
‘Vegetationspunkt.’ Dr. Hillebrand and I agree that, at the rate the Ger-
mans are going on, in twenty years there won’t be a botanist in Germany
who knows anything about the flora of the country.” In many respects his
prophecies were fulfilled, though at a slightly later date. At all events, he
was sufficiently convinced of the importance of systematic botany to bend
all his energies and a goodly portion of his means towards building up the
library and herbarium to make possible the continuance of this line of
study. He bought books, new and old, as they appeared on the market; he
bought exsiccati sets and had one of the best collections of these in the
country; he collected specimens during his trips at home and abroad and
determined many specimens for his colleagues and thus gathered an her-
barium that was very representative of nearly all groups of cryptogamic

LINDER: WILLIAM GILSON FARLOW 5

plants. All of this he gave to Harvard University along with an endowment
which, although not sufficient to maintain an adequate staff was, until the
recent depression, enough to carry on in a very modest way the one that
he had started.

Just as Dr. Farlow was a systematist who saw the importance of other
biological fields and, indeed, helped to initiate them, his successor, Dr.
Roland Thaxter was of much the same stamp and likewise continued to
stress the importance of systematic botany when all young mycologists
were stampeding into the rapidly growing field of phytopathology. A for-
mer student and later a very close friend of Dr. Farlow, Roland Thaxter
continued his predecessors’s policies, buying the important mycological
herbaria of von Hoehnel, Patouillard, Burt, and Bucholtz, the lichenological
herbaria of G. K. Merrill and of Sbarbaro, and the bryological collection
of Fleischer and parts of the Stephani herbarium. To these have since been
added the Sprague and Bartholomew herbaria of fungi and the Verdoorn
herbarium of Hepaticae. These herbaria in addition to the Curtis collection
of fungi, the James, Austin, and Sullivant collections of bryophytes, and
the lichenological herbarium of Tuckerman, are rich in type specimens and
accordingly very important to the systematist. As a result of the accretions
over the years, the Farlow Herbarium is one of the largest, if not the larg-
est, of its kind in America and rivals the few large ones in Europe. Today
the library contains some 36,000 volumes, including periodicals, pamphlets,
and reprints, most of which deal with the field of cryptogamic botany. It
contains many rare and indispensable books of which some are not repre-
sented elsewhere in this country. The herbarium, with its many types or
classical species, today contains 969,610 specimens distributed in the vari-
ous groups as follows: fungi 529,885, lichens 119,315, algae 54,582, mosses
222,579, and hepatics 43,249. Thus the Farlow Reference Library and
Herbarium of Cryptogamic Botany in collaboration with the Laboratories
of Cryptogamic Botany, which have become separated and are now located
in the Biological Laboratories building nearby, is in an excellent position
to continue the tradition established by Farlow.

Since Dr. Farlow placed so much emphasis on systematic botany and
showed his faith in the importance of the subject by devoting the greater
part of his life and means to building up the institution that bears his
family’s name, it seems desirable to examine the standing of this phase of
botanical science as it is at present. There is little doubt that with the
opportunities opened up by plant and forest pathology, and to a lesser
extent medical mycology, systematic mycology has been neglected, much
to the detriment of both mycology and those related fields. The war has
brought to light additional fields in the study of antibiotics (with penicillin
as a shining example) and of deterioration of manufactured materials
which, although a less spectacular field of investigation, is nevertheless im-
portant because it increases the cost of the war and thus effects every
American through that sensitive organ known as the pocketbook. There is
little doubt that these two new lines of endeavor — they cannot be called

6 FarLowiA, VoL. 2, 1945

fads since they are of tremendous immediate importance — are for some
time going to drain young men from mycology, and that systematic studies,
in the eyes of the public and even of scientists themselves, for they are not
beyond the reproach of being shortsighted when it comes to recognizing a
subject outside of their special fields, will fall into oblivion and accordingly
will lose the meagre support that they now receive. We have seen how
German botany, once virile, almost perished without systematic botanists.

How would the related fields of botany, or of chemistry or any other
branch of science that deals with living organisms suffer? We have already
seen in this war that when a person not properly trained is assigned to a
primarily botanical task, it is not until the botanist with systematic training
and with the accompanying techniques is called in that the essentially
botanical problem begins to clarify and to yield results or to clear up in-
consistencies that have previously been baffling. Just as it is nearly impos-
sible for a chemist to step in and solve a mycological problem (and vice
versa) most effectively, so too is it when a chemist, a doctor, a physiologist
or what-have-you tries to do monographic work or determine unusual forms
of plant life. To do this needs the experience that the taxonomist has re-
ceived through training and, even more important, through the experience
of constantly studying a wide range of material and the necessary litera-
ture. The classification of organisms is but one phase of systematic work.
Another phase is the collecting of material. Taxonomists are looked upon
much in the same light as the pack rats, — never satisfied with what they
have. This is rather praise than criticism, for it is only by gathering exten-
sive lots of material from various localities and under different ecological
conditions that it is possible to determine the range of variation of a species.
Furthermore, it has been from the kleptomaniac instinct that our knowl-
edge of the organisms that inhabit this green globe has increased, not in-
frequently to the great benefit in a practical way to the economics of a
people, as is shown by the development of disease resistant strains of wheat.
Whatever of an experimental nature is performed, it is fairly certain that
the experiment cannot be repeated or of general value unless the organism
has been accurately described and properly named.

Much more could be written concerning the importance of systematic
botany and of the necessity of libraries and herbaria, to say nothing of
financial support, for its proper and continued development. However, it
is hoped that the present remarks are sufficient to emphasize that while
immediate practical or economic application of knowledge is of great im-
portance, nevertheless it is impossible to tell when the so-called useless
knowledge of pure science may be called upon. It is therefore desirable
that there be a balance of a sort between applied and pure science. This
seems to be one of the lessons that we can learn from Dr. Farlow’s life.

HaArvarp UNIVERSITY
CAMBRIDGE, Mass.

*Emmons, C. W. — Misuse of the name “Trichophyton rosaceum” for a saprophytic
Fusarium. Jour. Bact. 47: 107-108. 1944.

LINDER: WILLIAM GILSON FARLOW 7

A LIST OF NOTICES AND BIOGRAPHIES

Blakeslee, A. F., R. Thaxter & W. Trelease. William Gilson Farlow. Amer. Jour.
Bot. 7: 173-181. 1920. With portrait and bibliography.

Clinton, G. P. William Gilson Farlow. Phytopath. 10: 1-8. 1920. With portrait.

Jackson, C. L. William Gilson Farlow. Proc. Amer. Acad. Arts & Sci. 57 (18): 484—
491. 1922.

Mangin, L. Notice sur M. William Gilson Farlow. Comptes Rend. Acad. Sci. (Paris)
169 (11): 445-448. 1919. ;

Riddle, L. W. William Gilson Farlow. Rhodora 21: 1-8. 1920. With portrait.

Robinson, B. L. in S. E. Morison. The Development of Harvard University since the
inauguration of President Eliot. Chapter XXII Botany, 1869-1929. pp. 366-370.
1930.

S. H. V. William Gilson Farlow. Ann. Bot. 33 (132): XV-XVI. 1919.

Setchell, W. A. William Gilson Farlow, 1844-1919. Nat. Acad. Sci. Mem. 21: 1-22.
1927. With portrait and bibliography.

Thaxter, R. William Gilson Farlow. Harvard Graduates Magazine. pp. 269-277. 1919.
With portrait.

. William Gilson Farlow. Bot. Gaz. 69 (1): 83-86. 1920. With portrait.

———.. William Gilson Farlow. Amer. Jour. Sci. 49 (290): 87-95. 1920. With por-

trait.
. The Farlow Library and Herbarium of Cryptogamic Botany. Harvard

Alumni Bull. 29 (24): 679-686. Illus. 1927.

—_—_.,, W. J. V. Osterhout & T. W. Richards. Minutes on the life and services of
Prof. William G. Farlow. Harvard Univ. Gaz. 15: 60. 1919.

Wl[akefield], E. M. William Gilson Farlow. Kew Bull. Misc. Inf. 1919 (9) ; 388-390.
1919.

THE CORRESPONDENCE OF WILLIAM G. FARLOW
DURING HIS STUDENT DAYS AT STRASBOURG

Hitpa F. Harris

2(1): 9-37 FARLOWIA January, 1945

THE CORRESPONDENCE OF WILLIAM G. FARLOW DURING
HIS STUDENT DAYS AT STRASBOURG

Hitpa F. Harris

In June, 1872, Asa Gray wrote to his friend Alphonse De Candolle in
Geneva, “My youthful assistant of the past two years goes in a week or
two to Europe, to study in some German university for a year or two; to
Strasburg, I think, unless he first should go to Sweden, and there study
Algae, with Agardh, if he will receive him. He takes a fancy to lower
Cryptogamia. His name is Farlow, an honest, good fellow. He will most
likely be in Switzerland in the summer; and I shall give him a letter of
introduction to you, whom he will wish to know. But take no trouble on
his account, except to introduce him to Dr. Miiller, from whom, as a work-
ing lichenologist, he could learn much.” 4

With this modest introduction, William Gilson Farlow set out to indulge
his “fancy for Cryptogamia” in two profitable years of study and travel in
the outstanding botanical centers of Europe. Farlow’s interest was well
placed, and received the whole-hearted approval of Gray, who was quick
to recognize the abilities of his young assistant, and was eager to have him
return home equipped to introduce the study of the lower plants into the
Harvard curriculum. The story of Farlow’s study in the laboratory of
Anton De Bary is chronicled in an exchange of letters between the two
men — long letters from Farlow, rich in newsy narratives of the activities
at Strasbourg, and briefer ones from Gray, revealing his deep interest in
Farlow’s progress and generous with wise advice as to the most profitable
plan of study. By direct quotation from these letters? it is possible to
create a picture of this interesting period in Farlow’s life in which he re-
ceived the preparation which was to influence so strongly his botanical

1 Letters of Asa Gray edited by Jane Loring Gray. 2: 625. Houghton Mifflin. Bos-
ton. 1893.

? The letters from Gray and De Bary may be found in the Farlow Library; those
from Farlow have been generously lent by the Gray Herbarium, and the writer wishes
to express her thanks to the Director, Prof. Merritt L. Fernald, for permission to quote
from them. The two Christmas letters from Strasbourg have been added recently to the
collection at the Farlow Library by Dr. David H. Linder, whom the writer wishes also
to thank for suggestions made during the preparation of the manuscript. Dr. Rolf
Singer has very kindly contributed the translation of De Bary’s letter from the German.
In a few of the letters quoted, obvious errors in grammar or spelling have been corrected.

Prate I, A portrait of W. G. Farlow, sent home during his European trip. Gray ac-
knowledged its receipt in his letter of March 23, 1873, of which the portion is here
reproduced in which he characteristically remarked, “Thanks for your photo. Very nice!
How your moustache sets you off!”

11

12 FartowlA, Vo. 2, 1945

work in this country, and during which time he acquired the nucleus of his
now famous herbarium and library.

Farlow’s first months in Europe were spent in traveling, but even then
he was anticipating the prospect of his winter’s work, for he wrote on
August 12:

Everyone speaks in glowing terms of De Bary and I hope to be suited with Stras-
burg. Hofmeister has gone to Tiibingen and it will not be far to go to see him if
it seems best . . . I am anxious to get to studying. I enjoy roaming about very
well but I like better to be settled.

It was not until October 6 that he wrote that he was in Strasbourg at last,
and had met De Bary. “His appearance is not striking. He is small and
homely and looks too young to have written as much as he has.”

If this initial meeting with De Bary failed to come up to his expectations,
Farlow’s impressions of Strasbourg were even more disappointing. Stras-
bourg had surrendered to the Germans in 1870 during the Franco-German
war, and Farlow arrived there two years later to find the city still filled
with soldiers, the university enrollment shrunk to a mere 250 students, and
the German language, which he had hoped to master thoroughly during the
course of his studies, spoken poorly and with an ‘“‘Alsatian twang.” Farlow
took an immediate and permanent dislike to the city itself. In his letter of
October 6 he complained:

I do not, now, think I shall be able to endure Strasburg longer than one term. It
is one of the most disagreeable places I have ever been in. The weather has been
horrible. I have been here, in all, ten days and not a single one at all pleasant
and the weather cold and raw. I cannot keep warm. As for society, there is
absolutely none. The university instead of having 1500 students has only 250
and no English or Americans . . . Prof. De Bary thinks Strasburg will be a very
good place to learn German but I do not agree with him. The people don’t speak
German and there is no society here except military men with whom I am not
likely to become acquainted. I am almost sorry that I didn’t go to some northern
city and study the language and let botany alone for some months.

As time passed, only a growing realization of the excellence of De Bary’s
instruction forced him to admit reluctantly to himself that he could not
hope to receive such training in any other place. As he said later (April 23,
1873) after a visit to Heidelberg:

[It] is certainly one of the loveliest places I ever saw and is particularly beauti-
ful in spring when all the trees are in flower. I was glad to come here because I
wanted to find out, for myself, whether it is really a good place to study or not
. . . Accordingly Veni, Vidi, but mot Vici. I find that H. is not to be compared
with Strasburg for botany and I leave tomorrow for the latter city to stay during
May and June with De Bary.

Farlow has described conditions as he found them at the University of
Strasbourg in a short paper which he wrote in October, 1873, intending it

Harris: CORRESPONDENCE OF WILLIAM G. FARLOW 13

for publication in the series of articles which appeared in the American
Naturalist. In it he said:

The University of Strassburg which has, for two years, been under the control
of the Germans, stands, as far as botany is concerned, in the first rank. It is
rather the fashion to say that the University has been created by the Germans.
This, however, is not true, unless we adopt the view that nothing can be a Uni-
versity which is not conducted on the same plan as those of Germany. Before
the Franco-German war, the name of Academy was given to what is now the
University, although the number of students was then much larger than at pres-
ent. Strassburg has for a long time been the residence of distinguished scientific
men. Of the botanists of the old régime, Schimper and Buchinger still remain.
Millardet, the most promising of the younger men, is now in Nancy. Prof.
Schimper, perhaps the most distinguished member of the scientific faculty, still
gives lectures on fossil botany and has a very large and rich collection of fossil
plants.

The active botanical work is performed by Prof. De Bary and Ass. Prof. Solms
von Laubach with Dr. Fritz Sc[h]mitz as assistant in the laboratory. The num-
ber of students attending botanical lectures, if we except the course on Medical
plants, which must properly be included in the medical department, has never
been greater than fifteen and quite frequently not more than five. The attendance
has sometimes been as low as two. I give these numbers simply to show how
different are the conditions from those of our own universities. An American pro-
fessor would hardly think it worth his while to deliver a lecture to five students.
The different proportion of instructors and students affords the clue as to how
the Germans are able to do so much high scientific work. The professors have
very little else to do. It is for this object that they are appointed professors
while in America the professors are only able to do any purely scientific work in
intervals between long courses of elementary instruction which is done in Ger-
many in the gymnasium.

Farlow began his own studies at the laboratory in October, 1872, and he
described his work in two letters to Gray.

Strasburg, Oct. 6.
Dear Sir;

_. . L have now been in Strasburg five days in my rooms, up four flights, in
the Hotel Allemannia. Every day I go to the laboratory where there are three
other students besides myself: a young man, quite young [Rostafinski] who is a
genius. He draws and paints very well, and is now at work on a book on the
Myxomycetes; another, a native of Strasburg [Gilkinet] who is skilful with his
fingers and a third, somewhat older [Suppanetz] from Gratz who is doing some-
thing or other with Pediastrum and some of the Desmids. The first two students
have been with De Bary three years.

Prof. De Bary is a very good teacher. He is very thorough. He put me, at first,
on to Chara and makes me draw everything with a camera. I find it rather slow
work, and am afraid I shall have to stay two or three years with him to learn
much, which is longer than I think I shall be able to. Here every one thinks that
a knowledge of the larger fungi or algae is contemptible. That the only thing
worth living for is to study the development of some of the lower forms. A sys-

14 Fartowia, Vor. 2, 1945

tematic botanist of Fungi or algae is regarded as a shallow person, of course,
without any ability . . . You will be surprised to hear that I have come to the
conclusion that England is, on the whole, the best place to study as they have
not such a mortal dread of anything systematic. I should like your advice about
what I should study.

W. G. Farlow. Hotel Allemannia.

Strasburg, Nov. Ist. 1872.
Dear Sir:

. . . I have now been a month in the laboratory but the lectures did not begin
till this week. The lectures I have decided to attend are, De Bary’s on Vegetable
Anatomy and on the Lower Forms of Life, Graf Solms on Cryptogamia, and
Schimper on fossil botany. Each lecturer lectures three times a week. Graf
Solms has the uncomfortable hour of 6 to 7 in the evening. None of the lectures
begin till a quarter past the hour. When it is light and I am not at lecture, I
work in the laboratory, with the three students that I mentioned in my last letter.
At first, I had the development of Chara and, then, started on vegetable anatomy,
first, with the sieve cells, which are interesting, and, now, I am studying the milk
ducts in Euphorbia, a tedious but not uninteresting work. There is a new mono-
graph on the latter subject by David, a pupil of Cohn. The probability is that I
shall devote six months to anatomy and then take the Myxomycetes. At first,
Prof. De Bary gave me as much attention as I could desire, but, lately, he has
been entirely occupied by his duties as Rector in matriculating the students. He
told Rostafinsky that he should soon be able to devote considerable time to me,
which will please me very well, as I don’t like his assistant Dr. Schmitz very well.

De Bary is an excellent instructor and a very pleasant man. The students all
like him. His German is a little peculiar and it is hard to understand him but he
speaks very good English. He has not much mechanical or artistic skill but he is
business like and doesn’t poke like most of the others. As a lecturer he is poor.
He doesn’t speak distinctly, doesn’t draw well, and is not at all interesting. Here,
there is nothing to learn in the way of addressing an audience. The Professors
have a very careless way. De Bary’s audience consists of five persons. Graf
Solms, as he is called, doesn’t speak English at all. I understand his lectures very
well and they are interesting. He seems to be a very industrious man and very
well trained in microscopic anatomy and the general structure of the lower orders.
I should hardly think that he knew as much about flowering plants as was neces-
sary for such a position as he has. He also lectures on Pharmakognosie but I
don’t attend. His delivery is somewhat better than De Bary’s but still he is not
a model. He has an audience of six .. .

On Wednesday I was matriculated. The imposing ceremony consisted in listen-
ing to a short poorly delivered address from Prof. De Bary, who is rector, then
shaking hands with him, and receiving a ticket of admission. There are, now,
about three hundred students in the University. I have to pay for lectures and
laboratory 120 fr., not including Prof. Schimper’s lectures which have not yet
begun, and for which I have not paid. Schimper is, it appears, a fine zodlogist
and geologist as well as botanist. He is attached to the zodlogical depart-
ment...

Yours, in haste,
W. G. Farlow

HARRIS: CORRESPONDENCE OF WILLIAM G. FARLOW 15

Farlow had not been long at Strasbourg when the opportunity arose for
him to acquire, for his personal herbarium, his first famous collection of
fungi. Before he left America he had discussed with Gray the possibilities
of buying the Curtis Herbarium, and in October he received the following
note from Gray.

Cambridge Oct. 16, 1872
My Dear Farlow

We have much enjoyed your letters — the last from Berlin, & including several
of the Linné photographs. They are most delightful, and I beg you will keep up
the habit of so writing to me. When I can I will write to you in return . . .

When you left you were a good deal disposed to acquire the late Mr. Curtis’s
coll. of Fungi. Mrs. C. is now ready to divide the Fungi from the rest and to
take $1000. I have to-day seen Mr. Sprague of Boston who advised my securing
them, and offerred — long ago — to pay $500 towards it. He tells me that he is
now in different circumstances from what he was, and wishes to forego the pur-
chase — don’t think he can afford to pay $500 at all. And putting in the steam-
warming here has used up the means of the Herbarium for last year & much of
this year.

So, if you are still very fungously inclined, and want to be set up in this de-
partment, I am out of the way, and you can buy the collection if you will. What
say you? If you decline, I shall offer it to the British Museum.

Ever, dear Farlow,
Yours sincerely,
Asa Gray

Farlow replied promptly (November 1):

As to the Fungi, I have concluded to take them. The price seems to me to be
a fair one and, although I know nothing about fungi now, I hope to some time
hereafter. The collection is unique and certainly should not leave America. I
think it will be worth a thousand dollars to me as a recommendation in securing
a position when I return. It is something to possess the finest collection of fungi
in America and I hope, before long, to have a good collection of algae, certainly
the best in America . .

I will write, at once, to my father about paying you for the Fungi and will ask
my brother to call and see you about the payment. What shall be done with the
collection? Had I better have it sent to my father’s house? How can it be for-
warded to Boston? I will ask my brother to settle these questions with you.

Cambridge Nov. 25, 1872
My Dear Dr. Farlow,
I received your letter —saw your Father, same morning — wrote to Mrs.
Curtis — and now have this reply: —
Will attend to all...
Ever yours,
A. Gray

16 FartowlA, VoL. 2, 1945

Gray enclosed in this note the letter from Mrs. Curtis.

Wilmington. Nov. 22nd. /72
Dear Doctor Gray,

I have this morning received your last letter, forwarded to me from Hills-
borough, and hasten to reply, lest you should imagine me doubly neglectful. The
former one, advising us to pack and forward to you, the books relating to Botani-
cal science, left by my dear Husband, was received a few days before I left for a
visit to my friends in this place, and it was scarcely possible to write just then.

I shall return home in two or three weeks, and will try to pack the Fungi, as
you wish, although it is a subject for regret to all of us, that the Collection must
be separated. When we have all arranged to send off, I will communicate with
you again, and let you know in what manner to make the payment.

With the greatest respect and regard, I am, dear Sir, Yours,

M. De R. Curtis.

Farlow acknowledged Gray’s kindness in offering to look after the collec-
tion, and in the same letter expressed his rather dubious opinion of De
Bary’s way of running the laboratory.

Strasburg, Dec. 19th.
Dear Sir;

I received your letter with the accompanying one from Mrs. Curtis a few days
ago and am glad that you have consented to see whether the collection is in
proper order although, I am afraid, that it may take more time than you ought to
give. The collection of M. De Brebisson is for sale in fragments and I have
ordered Cape and S. American algae to the extent of a few dollars. His collection
of diatomes is very good and is supposed to be worth 2000 Thalers. A Dutchman
I believe is going to buy it.

.. , Asa whole, the laboratory looks about as much like a chemical laboratory
as a botanical, since there are any quantity of bottles of reagents, test-tubes,
evaporating dishes and a great hood. There is an immense amount of strong
40 p.c. alcohol used as everything, even algae, are put into it at once and we have
each a small wash bottle full to moisten our sections every few minutes.

In some things they are deficient since there is only one drawing prism and one
Redresseur and, what is abominable, not a single dissecting microscope.

On Monday next, begins a recess of a fortnight and I am going to Frankfort
for a few days to rest, and, then, shall come back to study. I have now been here
three months and can form an opinion of the merits of Strasburg. As far as the
town itself and social advantages are concerned, it is detestably bad. It is a very
bad place to learn any language and there is no redeeming feature about it. As
far as the botany goes, sometimes I am satisfied and sometimes not. From what
I learn from the other students, what I regard as deficiencies in the instruction
are common to all Germany and not peculiar to this place. Everything is slow
to me, but, then, I am in Germany. There is no plan of instruction whatever.
That is, I must know what I want to study and ask for the material myself and
go to Prof. De Bary as I would to a dictionary, for information. I have learnt,
as have all the others, not to ask Dr. Sc[h|mitz questions, as the answer is more
perplexing than previous ignorance and, when he comes round to ask what I am
about, I tell him I have material enough to last for some time. I never let him

Harris: CORRESPONDENCE OF WILLIAM G. FARLOW 17

know that I want something more for, in that case, he picks up the first thing
that comes handy whether it has any reference to my previous studies or not.

About eleven, Prof. De Bary comes into the laboratory for a little while and,
generally, again late in the afternoon. He knows a great deal and is very pleasant
but is very often interested in abstract propositions which no one but a German
cares anything about. A fortnight ago Dr. Schmitz gave me a Lycopodium Selago
to study and, after making sections and preparations of one kind and another, I
thought I knew it pretty well when De Bary asked whether the capsule arose
from the stem or the leaf. After two or three days’ work I managed to get sec-
tions which satisfied him and, of course, myself that it arose from the leaf. Then
came the question how it arose which he didn’t know or, as far as I can find out,
anyone else. After much more labor, I discovered that the epidermal cells begin
to divide parallel to the surface and it grew out in that way. Then came the, to
me, transcendental question what might the subepidermal cells do, and, for a
week, I made section after section of the Vegetationspunkt with no result what-
ever till I used up a great bottle of material which put a stop, fortunately, to
further investigation. I ventured to suggest to my neighbors that I thought, as I
wasn’t to stay in Europe forever, that my time might be better employed than
searching for a needle in a haystack which I could do as well at home. My
Alsatian friend, from whom I learn more than anyone else, said Oh! in a few
months you will be able to write a monograph on the subject. That remark was
a clue to the whole thing; they study for the sake of writing monographs, just
what I don’t wish to do here. I can do that when I get back... |

Remember me to Mrs. Gray, and all the gentlemen in the herbarium.

Yours, respectfully,

W. G. Farlow

Farlow was always on the lookout for books as well as specimens which
would provide him with a good working collection when he returned to
America. He browsed through every available bookshop and catalog, and
gradually accumulated for himself and for Gray an enviable collection of
the best botanical books in Germany. By the summer of 1873 he wrote
that he had a trunkful of books which he would like to send home, and the
following summer he observed with some pride that as an added qualifica-
tion for a position he had “the Curtis Fungi and the finest collection of
algae and algological books in America and a good collection of modern
works on fungi.” Many of his letters to Gray contained reference to books
which he had seen on the market, as well as observations on the usefulness
of some of the newer works.

[December 19, 1872] Books are uncommonly dear, it seems to me, and we get
German books in America very nearly as cheap as here. Among books which
may interest you, are Thomé’s of the Real Schule, Cologne, Lehrbuch der Bo-
tanik. He has also written a Zodlogy and both books are the standards in Ger-
man schools. In the Botany, the plates are very good and he goes into the
anatomy and Cryptogamia much farther than in our own text books, and, I
think, too far for common school scholars. But that is a German failing. The
third edition of Sachs Lehrbuch has just appeared. The principal change is in the
article on Aleuron. The Lehrbuch in Germany is a sort of botanical Bible and
there is no use in telling a German that there is a great deal in it that is stupid

18 FarLowI!A, VoL. 2, 1945

and had better have been omitted particularly as it is borrowed from Hofmeister.
A book which I am going to buy sometime and which would be a great help in
your lectures on Economical Botany is Berg & Schmidt’s Officinelle Pflanzen, in
four moderate sized volumes with very good colored plates. It costs, I believe,
26 Thalers, a pretty high price.

[January 4, 1873] A book which I shall buy for myself and which, certainly,
ought to be in the Cambridge library is Tulasne’s Carpologia Selecta Fungorum,
of which, if I remember rightly, you have only the volume on the Tuberacei. It
is expensive, coming to, at least, fifty dollars, perhaps more. Another work, which
I, possibly, mentioned in my last letter, is Schimper’s Paleontology in three vol-
umes, in French and horribly expensive considering. It is 30 fr. a volume, I was
told. Strasburger’s Coniferae & Gnetaceae is regarded as a good morphological
work, that is, a book which nobody ever finds time to read.

The algae of De Brebisson which I ordered were previously engaged. I think,
before I come home, I shall get Fuckel’s Fungi Rhenani and Rabenhorst’s Fungi
Exsiccati unless they are already in Curtis’s collection. Have you Rabenhorst’s
Characeae? I have been advised to get them. Dr. Schmitz has sent you his
article from the Flora and I asked him also to send Mr. Wright a copy.

I spent Christmas week in Frankfurt stopping at the Hotel du Nord but spend-
ing most of my time at the Consul’s Mr. Webster, father of the only real
American in Strasburg besides myself. I went into the store of Joseph Baer and
spent some hours looking over his books. He has a Jacquin’s Flora Schénbrun-
nensis, in good condition for 68 thalers, a Hooker & Greville which Prof. Eaton
may like. I bought Martius’s Plant. Crypt. Brasil for $18, Mohl Vermischte
Schriften for $1, Boerhaave’s Historia Plantarum Lugduni-Batav. 1731, Dillenius
Cat. Plant. Circa Gissam 1719, for 50 cents, the two last simply as curiosities.
I found a Vegetable Materia Medica of the United States by Barton, Vol. 1 for
$1.50 and bought it. I don’t know whether a second volume was ever published.
Michaux’s Flora Am. Bor. 1803, 2 vols for $3.00. There was a Barton’s Flora of
the United States Ist vol. complete, and second not bound and not certainly
complete for about $10. I couldn’t find out how many plates ought to be in the
second volume. If I had been sure that it was complete, I should have bought it
for you as I heard you say you wanted it. In Mrs. Curtis’ letter she mentions
sending her husband’s cryptogamic books to you. Have you bought them for the
library or is there something which I ought to purchase?

If I felt sure of getting a permanent position at Cambridge, I should limit my
purchases of books to those which, as far as I know, are not in the library at
Cambridge. But, as it is, in consequence of losses by the fire, I suppose the
college will not feel like increasing the number of its instructors for some time
to come, so I must buy books which will be necessary if I go to a place where
the library is not as well stocked as at Cambridge. The same feeling interests me
in collecting. I don’t want to preserve the phaen[er]|ogams which I may collect
but the knowledge that I have a collection of flowering plants will help me in
getting a position in some other place than Cambridge. The fungi and algae, of
course, I save and perhaps I shall the mosses . . .

| March 6, 1873] Last week I saw a catalogue of Moser in Tiibingen with some
excellent things remarkably cheap. I wrote at once but found the best things
gone to my regret. I bought several pamphlets, on algae principally, and an
Endlicher. I saw a copy of Sprengel’s Das Entdeckte Geheimniss, which I know
you wanted, but it had been sold unfortunately. I agree with you that single odd

Harris: CORRESPONDENCE OF WILLIAM G. FARLOW 19

volumes are a delusion and a snare. About De Bary’s work every day something
of his which I have never seen before turns up. As I expect to be here next term
I shall secure more of his pamphlets for myself and next week I shall buy some
of those for sale. Perhaps, when the photographs arrive will be a good time to
suggest that he should send you a complete set of his works. Brefeld’s Penicillium
has just arrived in manuscript with ten plates said to be very interesting. Stras-
burger’s Azolla is like all his works but I don’t believe you will want it. Russow,
a Russian, has written a book in German on the structure of higher cryptogams
which I may possibly buy but I don’t think you will want. Have you Schwen-
dener’s Flechten-Thallus?

[May 12, 1873] I have just ordered and secured! Sprengel’s Entdeckte Ge-
heimniss which I know you wanted. At last, I have got hold of a Postels &
Ruprecht for 32 thalers in Berlin.

[June 30, 1873] A copy of Corda’s Icones Fungorum is for sale and De Bary
has told me that it is an indispensable work and advised me to buy it although
the price is 130 Thl.

[November 15, 1873] Ruprecht’s Tange, I bought before I left Cambridge.
Eaton wants to use my big Postels & Ruprecht until my return and I shall write
to my father to mail it to him. I have lately bought a few books and shall order
more. I expect soon a Tulasne for 60 Thl. Two indispensable and unfortunately
unprocurable books are Corda’s Icones Fungorum and Sowerby’s British Fungi.
A Corda was sold last year for 130 Thl. Is there a Corda in America? Sowerby
is in the Bost. Nat. Hist. I now have a good many pamphlets which have cost a
heap of money. However, most of them are very useful and not to be found in
America. Is the set of Pringsheim’s Jahrbiicher in Cambridge complete? I shall
try to get De Bary to let me have his duplicate set, not quite complete, of
Fuckel’s Fungi.

[June 13, 1874. London] De Bary has bought Duby collection. I have bought
Sowerby’s Fungi and a Harvey for my own use. I can’t make up my mind to
give $200 for Kiitzing’s Tabulae Phycologicae. I feel more like giving $80 for a
Bulliard which is very useful for fungi and not often to be had.

After Farlow found himself more or less oriented to the German way of
research, he began to shape up more definitely his own plan of study. His
language difficulties, coupled with his intense dislike for Strasbourg, brought
up the question of whether to stay on with De Bary, or to move to some
more congenial city. Some of his problems were settled when, early in
January, he received a number of suggestions from Gray who, kept at home
by an attack of influenza, found himself with plenty of time for planning.

Cambridge Jany. 10, 1873
My dear Farlow,

Yesterday came another of your welcome letters — to solace me in illness: for
I suffer with a dreadful influenza — & Goodale is away, at Baltimore.

While lying sleepless I have thought over your studies &c —

We are going to have the Cryptogamic Flora of U.S. done up, pari passu with
the Phanerog. Sullivant is engaged to do the Mosses, in a volume —to which I
hope to have Austin do the Hepaticae.

I want you to come home prepared to do —

20 FartowiA, VoL. 2, 1945

1. The Algae — one vol.

2. The Fungi —

3. — since they are only Algae & Fungi! either dwelling together — or
the lamb inside of the lion—you will probably have to do the
Lichenes! Unless we can get them out of Tuckerman, in an intelligible
form — which is doubtful. He is somewhere among your friends the
Germans.

I should think you are doing pretty well — learning what you ought to know —
getting the ways of working. But tell De Bary that if you stay there another
semester, you must be put at work on & be taught Fungi. Of course, only low
Fungi could be noticed there — or could be worthy of notice by your philosophers,
such as the mould upon a Strassburg cheese or a bad-smelling sausage: Wurst —
I think they call them, which is descriptive when spelled with an O.

If De Bary can’t see that you are taught this special lore, why, I would migrate
in due time...

I doubt if Hofmeister has ever been much of a teacher. Yet I always thought
much of his work. I suppose your best chance would be with Nageli. If they
find him intelligible, there need be no despair anywhere.

Tubingen should be a cheap place to live, and a complete University town.
Hegelmaier who worked up Lemna & other low Pha[e|nogams is a Botanist, and
I should think you might learn much of him, as to ways of research.

Suppose you look in at Tubingen & see a little for yourself.

Munich is cold enough in winter, but an attractive town. Négeli conducts a
well-ordered botanical laboratory. Radelkoffer, his aid is a jolly good-natured
man, who knows plants & flowers somewhat, as well as vegetable anatomy. So, I
should be content with Munich for one trial.

But I have a capital idea for you when your summer vacation comes — Dr. J.
Miiller, Candolle’s curator, at Geneva, is one of the best practical workers at
Lichens &c.— the very man to initiate you into Lichen genera & Lichen work.
When summer or spring arrives go up there, and arrange with him to take you
in hand, & have with him, & De Candolle, & Boissier your respect raised for
Botany. In 3 weeks of work with Miiller you would learn Lichens, and Geneva
is the place of departure & return for the Swiss excursions you will enjoy .. .

They are going over the collection of Fungi, Mrs, Curtis & son, — and in reply
to a letter, I insisted that they were to see that all was thoroughly poisoned or
repoisoned!

Get as many of the Lehrbuch’s as you think worth while . . .

Oh — as to Cohn at Breslau. I think you might do well with him, for a while,
as you say. And then, if Goeppert is not too old you might really learn somewhat
of fossil-plants — which you never would from Schimper’s lectures.

But I have another good idea. Get Schimper in a series of private lessons, as
the days get longer, to put you up to Musci.

He is an old & famous hand at that, — and you might learn all his wrinkles in
a fortnight.

Mrs. Gray is as much broken down as I am — & suffers with, what Dr. Wyman
assures us is Pelliosis rheumatica! —I, with influenza diabolica!

Always Dear Farlow,
Yours truly
Asa Gray

HARRIS: CORRESPONDENCE OF WILLIAM G. FARLOW pal

Farlow replied on January 27.

. .. | am very sorry to hear that you and Mrs. Gray are sick. I had hoped
that, with the better heating of the herbarium, you wouldn’t be liable to attacks
of bronchitis as before. However, I suppose, you are glad that Mrs. Gray is sick,
since misery always likes company.

I was glad to get your advice about my studies. In a week or two I shall have
finished my anatomy and shall then go on to Pilze. So far, my regular study has
been anatomy but I have made occasional excursions into other branches. I learn
a great deal, also, from the studies of the other students who always show, to me
what they have interesting. Stahl is at work making a lichen out of a fungus and
an algae. I have now saved about 70 microscopic preparations which will be of
very great service to me as I know exactly what points each specimen illustrates.
We are taught to make very thin sections, no matter how small or ragged they
look. I think I could stay with De Bary another term to study Fungi and Con-
jugatae and lower algae. Strasburg itself, however, is utterly unendurable and I
find it fearfully hard to learn the language here. I had some thoughts of going to
Bonn to spend the summer semester and study botany only forenoons and the
language afternoons and evenings. I must learn the language at once. If I was
in a private family and could give my whole time to German, I could get it. As
it is, my best energy is spent entirely on botany. Your letter however, changes
my plans. I must decline going to Tubingen or any other small town in South
Germany until I have learnt the language better. I hope to study in Munich
next winter. Your statement of what I am to learn hardly comes within the
bounds of your usual modesty. I think I could stand the algae and, in course of
time, the fungi, but, when you come to add the lichens, I begin to despair. I
should like, as you propose, to study in Geneva and should like a letter of intro-
duction to Mueller if you feel inclined to give me one.

As the term neared its end Farlow was able to estimate the value of the
courses he had studied, and he seemed suddenly to find a surprisingly large
number of reasons for enjoying his work at Strasbourg. On March 6 he
wrote:

. . . For myself, the most interesting thing at present is that the semester will
end next week and we have a recess of about six weeks, till May ist, when
according to all appearances I shall return to Strasburg for the months of May
and June. .=

I like De Bary better and better the more I know about him and the students
in the laboratory are pleasanter than I can again expect to meet in any one
laboratory. Rostafinsky leaves and goes to Munich where I shall meet him again.
Suppane[t]|z also leaves to secure an appointment in Gratz. Dobrach goes to
Bonn, but he was by far the least interesting of all. Gilkinet and Stahl remain
but go with me to Munich next winter. Besides the botanists, I have only one
acquaintance in Strasburg of any account, that is Webster, the American law
student.

As to my studies (in botany not German) I suppose I am as near satisfied, as
I, who am a first class grumbler, can expect. I have been over anatomy practi-
cally and by lectures very much more thoroughly than ever before and there is
but one point which I have omitted which can be better studied in summer, the
development of the embryo. I have about a hundred preparations and some

ANTON DE Bary Ernst STAHL

ALFRED GILKINET JoseFr ROSTAFINSKI

PratTeE II

Harris: CORRESPONDENCE OF WILLIAM G. FARLOW 23

drawings which will be of use to me and what is most important have learnt the
German way of work and managing a laboratory which is radically different from
ours. Lately I have been at work on the Pilze and their development in which
branch De Bary has more practical knowledge than anybody else in the world.
He is very pleasant and takes an interest in his students and gives them a reason-
able amount of time and attention. I have studied the potato disease, Mucor
and the allied forms, Piptecephalis and Chaetocladium, Ascobolus and Eurotium
as types of Sphaeriaceae and Dictyostelium mucoroides. The last was particu-
larly interesting from sowing the spores till the ripe sporangia were produced on
the object glass. De Bary spent an hour showing me the original preparations of
Brefeld’s Mucor and Hallier’s cholera Pilze, the latter very shabby, dirty and
composed principally of striped muscular tissue. Incidentally I have examined
other fungi, Uredineen &c. My neighbors in the laboratory are very obliging
and help me in any way they can.

Next term, De Bary has two courses, a course on general anatomy which I
shall not regularly hear as it is too simple, elementary, and once a week on the
important medical plants. Graf Solms has two courses on Systematic Botany
which I shall attend, if they are good, and on Phaenogamic Parasites. With re-
gard to Schimper I don’t think I care to study the mosses with him; he is learned
but now old and a little tedious. I could learn nothing from him in a short time.
I may instead study lichens with Graf Solms one or two afternoons in the week
as a relief to the Pilze study. With De Bary I shall study only fungi and their
development and leave the algae for Munich or, if I am so lucky, Thuret .. .

I have taken tea twice at De Bary’s and a few Sundays ago all the botanists
were invited by Gilkinet, the Belgian to dine. De Bary was there and we sat
down to the table at one and rose at half past nine, a fearful feed. At the begin-
ning each guest was provided with two enormous bottles of wine, and, when these
were disposed of, more was brought on . . . Next week, I believe, Rostafinsky
is to give a dinner.

Farlow derived a great deal of pleasure and no little amusement from his
contact with De Bary’s assistant, Graf Solms von Laubach. He described
him as “a strange man, good hearted and well meaning but very awkward
and, as a Graf, filled with strangely absurd stories about America.” The
Graf, however, was not the only one to have extraordinary ideas of America,
because Farlow remarked in one letter “I am not unfrequently asked
whether our lectures are in English or German by intelligent people. I have,
also, been asked if it wasn’t very pleasant collecting near Boston because
the Gulf of Mexico must be a very good place for sea-weeds.” At another
time he told of meeting Miiller’s young son, who had recently been reading
Cooper, and was “delighted to see some one who could tell him all about
the Indians of Boston.’”’ Unfortunately Farlow did not elaborate upon the
Graf’s conception of this country, but he has described how he enlivened
the work at Strasbourg. In a letter to Mrs. Gray (October 27, 1873) Far-
low told of a collecting trip to which Graf Solms contributed most of the
entertainment.

Prate II. Contemporaneous portraits of Anton De Bary and three of the laboratory
students with whom Farlow associated at Strasbourg.

24 FartowlA, VoL. 2, 1945

Day before yesterday we all went on an excursion to the other side of the
Rhine to a village called Kork. First, of course, we had to have some dinner
and Graf Solms, who is one of the most awkward and amusing men I ever saw,
began by upsetting his soup all over his clothes which was followed by a volley
of Donnerwetters and wailings over grease spots which never can be taken out,
requiring all the skill of De Bary to quiet him. The rest of us were so con-
vulsed with laughter that we couldn’t say or do anything. We all ran for the
train and had to pay a fine of 50 ct. for coming so late. Kork is a flat plain
interspersed with mud puddles and ditches and here the Kork youth with woollen
night caps watch large herds of swine and geese. We proceeded slowly through
this charming region the only question as to how wet we should get. At length
we came to a very dirty pool full of different species of Elatine and Graf Solms
began digging in the mud and screaming every time he fished up an Elatine to
the great delight of all the swineherds and De Bary who called the plant Elatine
spectaculosa in consequence. Another pool was full of Marsilea and the botanists
all lay down flat in the muddy grass and began digging up the mud with their
fingers and my umbrella, Graf Solms screaming at intervals and jabbering mixed
French and German. When we were all dirty enough to go home, he discovered
that he had not worn his excursion shoes which distressed him exceedingly. His
troubles were drowned in a glass of beer at the depot.

As to his lectures, Farlow said (February 4, 1873):

Graf Solms I don’t like particularly well on the Cryptogamia. He is now on
the algae, of which he has made a perfect botch. There is no classification of
the algae to be allowed because the ‘Entwickelungs-geschichte’ isn’t well enough
known; therefore, by German logic, Graf Solms is not to have any system in his
lectures, in which negative point of view his lectures are a perfect success. Today
a specimen of Gracilaria was passed round to illustrate Gigartina compressa and
Wormskioldia was described as Delesseria. Porphyra was also in Florideae. The
mixture was so bad that I don’t wonder that the students here, not one of whom
has ever seen the sea, think that the algae are in a wretched state.

Later he wrote (March 6, 1873):

Lately he [Graf Solms] has been lecturing on the Pilze and as his lecture room
is next to De Bary’s private laboratory he was afraid De Bary might hear his
mistakes so that he has lectured in a whisper for some days to our great amuse-
ment. His greatest fault is that he has no system about his lectures but perhaps
that is because he is an aus[ser]ordentlich[er] professor.

There was some thought at this time of calling a European botanist to
Harvard on Gray’s retirement, and this possibility made Farlow wonder
about his own prospects after his return to America. Fearing that Harvard
might not have a place for him, he asked Gray about the opportunities in
California, saying that while he preferred to be in Cambridge, he was not
“of the number who believe that all colleges but Harvard are barbarous.”
Gray, in his letter of February 23, 1873, hastened to reassure him. He
wrote:

Take your time — prepare thoroughly. Meanwhile, if you get in a hurry —
and if, as is likely any other place turns up — and places are sure to turn up — I
will see to your interests all round.

Harris: CORRESPONDENCE OF WILLIAM G. FARLOW 25

Do not be at all uneasy. By the time you are ready there will be some good
place for you. Anyway as soon as you come back there will, at the least, be an
assistant-professorship here for you, I cannot doubt.

But your winning card, as it is what is most wanted in U. S. is Cryptogamy —
especially low Cryptogamy. Fungi will be the most telling card. There you will
have an exhaustless and a popular field —in which, well prepared — you can
make a mark here.

Algae — are well, but marine soon exhausted .. .

Then Lichens. Tuckerman is hors de combat, I fear. You must really go into
them! And, indeed, when Sullivant goes off the stage — which must be before
very long — there is no fit moss-man.

Whether you should stay at Strasburg longer I am not competent to advise.
I should say stay if Strassburgh were a pleasanter residence. You are just now
entering upon the field in which De Barry can teach you much, & I hope would
do so. But you must take the universities in their turn.

Perhaps in some you will find good lecturing — which is also to be studied.
Perhaps you must go to France for that. The French ‘expose’ best.

. We were so amused with your account of Solms’ lecture on Algae. If
you stay at Strassburg, you had best be a Privat-docent in Algae. It would tell!

Farlow replied gratefully on March 24:

Dear Sir;

Yours of Feb. 24 was received a few days ago and I was even more glad than
usual to hear from you. Of course, there are times when I feel anxious to know
what I am to do when I get back and that anxiety does not help my studying for
the present moment, but, as long as I feel that there is some one who has an eye
to my interests, I do not feel so much inclined to hurry up my studies here. I
cannot express my obligations for the expressions of interest in me contained in
your last letter.

There are evidently two ways of studying the botany. The first, to catch the
plan of work in different places and then myself to transfer that plan to America.
With such an object I feel as though I had already had enough of Strasburg. I
know De Bary’s method of investigation and could apply it at home —if I had
time. A term with Nageli and a short time with Hofmeister and two or three
months in France would teach me the methods of study adopted by European
botanists and in a year and a half I might come home, not so very badly pre-
pared, at least, to tell what other people are doing.

The second plan involves in addition a continuous work under the different
professors and requires, I am afraid, double the time required by the first men-
tioned course. In pursuance of this plan, the most thorough and the one I should
rather adopt, if I could spare time, I ought to stay longer in Strasburg. Accord-
ingly, when I left Str. a week ago I told De Bary that I should probably return
for the months of May and June. I left my preparations and microscopic affairs
there. Next winter as I have said, I expect to go to Munich .

If you were amused at the account of Solms lectures on the algae you would
also have been amused at those on the lower fungi. He told us that we must
regard the Myxomycetes as animals for he didn’t have time to lecture on them.
If he had had two more lectures he should have made them out to be plants.
He tumbles over his chair puts his fingers in his mouth and goes through con-
tortions that would make a cat laugh. He is goodnatured however. Just before

26 FARLOwWIA, VoL. 2, 1945

I left Strasburg I invited all the botanists, eight in number, to dinner. De Bary
was at one end of the table and I at the other and Graf Solms in the middle
where he kept up his old habit of swigging down red wine. Suppane[t|z the
Austrian was more blooming than ever and went into rhapsodies over some figs
on the table, Siid-europaische friichte, Liebe[r| Gott, classische, colossale, ganz
eminente. He flourished his knife in a way to make me fear for my life. The
supper lasted, for Strasburg, a short time, that is from half-past eight till a little
before twelve. The greater part of the company then retired to the Deutscher
Kaiser to finish up with a few glasses (10 or 12) of beer. The laboratory broke
up last week. Suppane[t]z is ordered by the Austrian government to Prague in-
stead of Gratz where he will probably get a place; Rostafinsky went home, Ddl-
brach to Bonn and I cleared for Frankfurt. At Carlsruhe I ran into Graf Solms
who was characteristically trying to back round a corner. He spent the time till
we reached Heidelberg in repeating to me how ‘komisch’ it was that we had
MEL“. 4 «

You are getting so modest in your demands that in your next I expect to hear
that I must study the ferns, as, in course of time, Eaton will depart this life, and
possibly the flowering plants also. The lichens I should like to study and in
Geneva where I understand one can learn French better than in Paris. However,
I must first learn German, and it will be provoking to find that just as I am
getting so as to be comfortably up in the language I must go home.

Rostafinski’s plans of studying algae with Thuret raised the hopes of
Farlow that he might be as fortunate, for, as he said “There is no one in
Europe I should so much like to study with as Thuret.” On receiving the
letters of introduction which Gray wrote to Thuret and Miller, Farlow lost
no time in sending them off to the two men, with his own request for per-
mission to study with them. On May 12, Farlow wrote:

I had a very polite letter from Thuret saying that he should soon leave
Antibes for the summer and he should be happy to have me meet him this sum-
mer in the vicinity of Nantes or next winter in Antibes. I accepted the latter
alternative with pleasure . . . Thuret wished, in very polite language, to be re-
membered to you and Mrs. Gray.

Of the new semester’s work he said in the same letter:

I omitted to write you, I believe, that, during the vacation, De Bary had a call
to Vienna which he declined. Since then Sachs has also declined. It appears that
the University in Vienna is on the decline in every department except medicine
and there is so much intrigue and chicanery at work there that no distinguished
men will accept positions. It was a disappointment to me that De Bary didn’t
accept, as I should have then spent next winter with him in V. and this summer
have gone to Munich.

. . . I returned here at the beginning of the semester. I do not regret return-
ing. In a social point of view, the laboratory is not so pleasant as last semester
since Rostafinsky and Suppanetz are gone and Stahl is at home most of the time
getting ready for his examination in June. A new Pole is here, a beginner, and
three students who only work three afternoons in the week. But De Bary is
capital. He has a general course from 8 to 9 which I don’t hear and then is in
the laboratory nearly all the morning. He pays me a great deal of attention. At

HARRIS: CORRESPONDENCE OF WILLIAM G. FARLOW ou

present, I am on the Uredineen and I am cultivating rusts and inoculating sound
barberry leaves. I have an ergot Cultur underway but, I am afraid, the material
is poor. Tomorrow I begin on Cystopus. There yet remain the fermentation
pilze which I shall soon take up. We have all been catching flies for Saprolegnia
and Achlya. De Bary began today on Medicinal Plants to a class of 36, a large
number. It was simple and practical, in short, excellent. The same cannot be
said of Graf Solms’ lectures which I attend simply from personal attachment not
because I like the lectures. Systematic Phaenerogams is a hurried humble-jumble
and Parasitic Phaenerogams once a week, considering the materials which he has
spent so much time and money in collecting, are not very interesting. He is soon
going to publish something on the latter subject. He has several ‘junks’ of
Rafflesiae in alcohol . . .

The semester opened a week ago, on Thursday. De Bary, as Rektor, gave an
address, delivery abominable, substance very good. De Bary’s personal appear-
ance is certainly unprepossessing. Everyone says, at first, is that man really
De Bary? In the evening was a torchlight procession in honor of the three pro-
fessors who declined calls to Vienna. The scene was very interesting as the pro-
cession passed by the Cathedral to the Palace where the Professors addressed
the students. Afterwards they all marched to the Kleber Platz and burnt up
their torches in a heap and sang Gaudeamus as at the funeral: of a Chor student.
At ten o’clock began the Commers and lasted till nearly morning. I left at two.
4500 glasses of beer were drunk on this intellectual occasion. The singing was at
first splendid, afterwards slightly confused. There were generals and professors
and students all mixed up together drinking, all dressed as gayly as possible.
Gilkinet and I sat in the gallery where we could see the whole performance.
There were speeches, songs, Salamanders, shouts, howls, and above all drinking.
The scene was new and interesting and I shall not forget it. Several times the
waiter upset our table with several glasses of beer upon it. Fortunately, the beer
ran over onto Gilkinet’s side and he soon got to be tolerably wet. After a while
I found myself about ankle deep in beer and concluded to find another seat. I
am happy to say that I managed to get along without what is here known as a
Katzenjammer or as the students say a Kater. In the course of the evening
Graf Solms appeared miraculously in the gallery with Dr. Schmitz and we drank
the health of the University. What that means I don’t know. Afterwards the
ubiquitous Graf wandered round the hall holding up an empty beer glass until
after a while he lighted upon General Carminsky and the two talked ‘Faderland’
till morning.

A letter from Cambridge in June left no doubt that Gray was looking
forward to seeing Farlow established at Harvard on his return to America.
Gray also suggested that it would be well to begin to work up some special
research which could be published within the coming year, and Farlow im-
mediately responded to the idea.

June 9, 1873
My Dear Farlow
You know how very busy I must have been — and still am, with Goodale gone
since May 1. So you see why I have not promptly responded to yours of March
24, from Frankfurt, April 23, Heidelberg, & May 12, Strasburg — And even now
— late in evening, I can only write a line.

28 FarLowlA, VoL. 2, 1945

You seem to be going on famously well, and doing your best and wisest. I am
glad you are back with De Bary. And I like the way you are laying out your
work for summer and winter. Especially I am glad of Thurets kind response to
you, & that you will get some time with him next winter or early spring . . .

Mrs. Curtis writes that she has sent the Fungi; but I have not yet heard if
your Father has received them.

Goodale has been appointed ‘Assistant Professor of Vegetable Physiology &
Instructor in Botany.’ — his name will go to Overseers, I think to-morrow. The
Corporation begin to be anxious about Bussey School — wants some Professors
to work there —& are determined to have students. Things are shaping as fast
and formally as I could wish,—and when you come home — unless you are
called away by better offers elsewhere, you will certainly find something to do
here. I see they are beginning to want you, and you may fairly count upon
offers when you come home — very likely before. I have said, that you can’t
think of returning to America for a year yet — perhaps not even then — your
time & opportunity in Germany too valuable.

I think it best for you—after consulting De Bary as to what —to get up
some research which you can publish in Germany — or —still better — send
over for me to publish here —in Amer. Academy — or in Amer. Naturalist —
according to circumstances.

Then another thing will be very well.

Send home to me, from time to time, notices of interesting works or researches,
Botanical news — criticisms &c —to be published with your initials among the
miscellanies in Sill. Journal. Or, if you will write some sprightly letters on such
matters that I can print in Amer. Naturalist, it will be well.

But lose no working time. Solid work is better than any display ... All
good things you can send me to print, under your own name, or initials — which
will duly be known — will be advantageous . . .

Ever yours truly
Asa Gray

Strassburg, June 30.
Dear Sir;

Shortly after sending my last letter I received your letter of the 9th and the
notices of the death of Dr. Torrey and Sullivant. I was quite surprised to hear
that the latter was dead and I had no idea that he was so old a man ae

I have spoken to De Bary about some special work. He proposed, The fertili-
zation and fructification of Agarics, ditto of the Oscillariae, ditto of Scenedesmus
and the Entwickelung (cultivation and growth) of Oidium lactis. The last I
selected but, as one may work an indefinite length of time without meeting with
any success or result, I have also taken as nebensache or collateral, the micro-
scopic structure and growth of the Haftorganen i.e. the disks of Ampelopsis, and
the rootlets of Tecoma &c. From one or the other I ought to make out something,

As for writing notices, I should be glad to do that and must try and find time
in the vacations. The following subjects suggest themselves: late German Bo-
tanical Text Books, reviewing Sachs and Thomé, Rostafinsky’s Myxomyceten;
or, possibly, Cornu, Saprolegnieen and Strasburgers, Coniferen, For light letters,
an account of Upsala, and botanizing in the Vosges. I might give a detailed ac--
count of the management of De Bary’s laboratory and cultures but that is too
much like letting the public into the secret . . .

Harris: CORRESPONDENCE OF WILLIAM G. FarRLow 29

I should like, after finishing my studies to write a compendium, something
like Sachs, about Algae and Pilze, giving the latest modes of working and results
so far as known. Such a book is very much needed as Berkeley is thoroughly
bad and there is positively no other book in English. The writing would serve
for me as a review of my studies and could also be available as a text book in
classes where I had to teach the subject. The first thing, however, is for me to
learn the subject myself. Remember me kindly to Mrs. Gray.

Yours, respectfully, but tired,
W. G. Farlow.

Farlow left in August to spend a pleasant six weeks in Geneva with Dr.
Miiller, and returned regretfully to Strasbourg in October, to take the last
of the courses with De Bary which he needed to round out his studies there.

Strassburg, Oct. 27th.

Dear Sir;
I am again installed in Strassburg and, after Geneva, dislike the place more
than ever . . . In comparison with other cities, it seems barbarous . . . I found

my six weeks in Geneva very profitable. Dr. Miller gave me an unlimited
amount of attention and I was surprised to find how much, with his help, I was
able to do in so short a time. I have no means at present, to repay his kindness.
Of course, I thanked him and gave him a book as a memento, but it would give
me pleasure, if you should hereafter, if you have occasion to write to him, men-
tion that I have written you that I feel under obligations to him for his attention
and instruction. While in Geneva, I had an opportunity to make several excur-
sions into the Jura and mountains near the city, in which excursions I succeeded
in collecting a large number of lichens which are not only numerous but valuable,
as I have studied them tolerably carefully, and, also, saved duplicates with an
eye to future exchanges in America. Although, perhaps, having studied lichens
under the most favorable and agreeable circumstances possible, I must confess
that I think they are the stupidest and, at the same time, the most difficult plants
I have ever studied. However, if it is necessary for me to study them in America,
I feel as though I had had a good preparation here. During the summer, I find
that I have collected quite a lot of Erysipheen, Uredineen, Perinosporeen, and
the smaller fungi generally and have laid by a number of duplicates which I
think will be of good service, hereafter, in America.

So far, although I have constantly regretted being obliged to shut myself up
in Strassburg, I feel as though I could no where else and in no other manner
have learnt so much of that part of botany which I needed to know and which
is in America entirely unknown, certainly practically. I have now learnt so much
anatomy, that is histology, that De Bary doesn’t advise my spending any more
time on it here. Then I have studied the orders of minute fungi tolerably thor-
oughly. Of course, to know the subject perfectly is a work of a life time. I have
learned De Bary’s method and, although a knowledge of forms and species is
here discouraged, I have learnt a great many forms and, with the knowledge of
their development and growth which I now possess, the determination of species
comes tolerably easy ...

The remaining botanical subjects which I wish to study are, the development
of the higher cryptogams, i.e. the fertilization of ferns, Marsilea, &c; second, the
algae, marine and fresh water in respect to their development, and physiology;

30 FarLowIA, Vou. 2, 1945

and thirdly, a short study of the cell, in the abstract, as has been done by Hof-
meister in his latest work. Whatever I now study comes a good deal easier than
it did a year ago, showing that the somewhat slow preparation recommended by
De Bary is beginning to bear good results. I don’t feel as though I wanted to
stay much longer with De Bary. I should only be continuing the same work I
have already begun. It is simply now a matter of detail and, if I am to stay here,
till I know all he can teach I shan’t get back to America for a long time. If I
should stay here too long, I should be more likely to turn out a botanist of the
Graf Solms type rather than like De Bary himself . . .

If I can return in September, as I hope, the question will be whether to go to
Pringsheim in May, which will perhaps be decided in the negative if I study fresh
water algae with Thuret, which may be possible, or to go to Wurzburg where
Stahl and Gilkinet are going, and study the cell under Sachs and attend his course
on general Vegetable Physiology without actually going through the experiments
in his laboratory. In this way, I should get all the Physiology necessary for me
as a cryptogamist. Brefeld is also a physician in Wurzburg and is now studying
hefe pilze and, perhaps, could help me some. I hardly think I should find it
profitable to stay more than two months in Wurzburg. Afterwards, there is
nothing which I care specially to do but, perhaps, to visit Rabenhorst, of whom
both De Bary and Prof. E. Fries have a poor opinion. Then a few days on the
coast of France or England and a visit to the leading English cryptogamists . .

I think now I have stated the whole story. I should like to return next fall.
If it is necessary for me to remain a year longer, I must simply spin out longer
the list of studies I have mentioned and, possibly spend a short time with Stras-
burger in Jena, and, although I should at the end of that time know more botany,
I should, perhaps, be less qualified to teach in America. Whether, as far as
America is concerned, the opportunity is good for my returning you know very
much better than I. I think you also know from a former letter of mine that,
although I should like to return to Cambridge, I don’t feel as though my future
success entirely depends on it. As I then wrote, I should like also to go to either
New York or California should there be opportunity in either place. At any rate,
I feel perfectly safe in trusting to you, knowing that you will recognize what is
for my best interest.

Here in the laboratory are Stahl, Gilkinet, and Carminski who remain, and
two new students, Prof. Czerokin of Kasan, Russia, and Dr. Linsted of Berlin
who has written on Saprolegnieen. After Christmas, Dr. Schmitz goes to Halle
as Docent and Rostafinsky, who is now in Petersburg, may take his place. De
Bary repeats his course on Anatomy and soon expects to finish his book. Graf
Solms also repeats his Cryptogams. In accordance with your request I have
written four gossipy letters for the Naturalist. I do not know whether you will
not find them too light . . . I also send the promised critique on Thomé and
Sachs. In a few days, I shall send notices of some recent magazine articles. I
hope to be able to send you soon a critique or summary of the Lichen, algo-
fungus question...

Farlow

These ‘“‘gossipy letters” were edited by Gray, and soon appeared in three
parts in the American Naturalist, vol. 8, 1874, under the title Notes from
the Journal of a Botanist in Europe. Part I. Sweden. Part II. Norway, etc.
Part III, Geneva and the Alps. Gray wrote on November 17:

Harris: CORRESPONDENCE OF WILLIAM G. FARLOW 31

I am arranging for the articles in Am. Naturalist — to begin in Jany no. Nat-
urally enough these articles are not so sprightly as your letters. I wish you would
write only on one side of your paper. I should like to season up the articles by
sandwiching in bits from your letters.

On January 11, 1874, he announced their publication:

The Ist part of your Notes is in Jany Naturalist. It reads nicely. I have had
proofs of the 2d — for Feb. number. I made it up out of your letters!

In the meantime, at the suggestion of De Bary, Farlow was at work on a
new topic for research, which was to culminate in the much discussed paper
on the discovery of apogamy in ferns. He first mentioned it on December 7:

Last week was rather an eventful one. I discovered an entirely new Puccinia
on the common mallows in the botanical garden which germinates at this season.
I presume De Bary will cultivate and describe it. My work has been the fertili-
zation of ferns and a week ago De Bary found a case in Asplenium molle where
the fern is produced directly from the prothallus without the intervention of an
archegonium and fertilization. He gave that to me, as a subject to work up and
I am now working busily away and have made quite a number of drawings. I
shall finish the work as soon as possible for I want to go to Paris with Jackson
about Christmas. However, I cannot leave till my work is completely done. It
will probably be printed in the Botanische Zeitung and I will send you the same
in English for the Am. Journ. if you wish. What shall be done about the figures?
Had they better be copied from the Botanische Zeitung or shall I send you the
drawings?

With regard to the letters I sent, it was difficult to say just how ‘sprightly’ I
ought to make them. What I sent would here have been considered dreadfully
frivolous. You did not say whether they wanted any more or not. You can make
any insertions you please, but I don’t want the editors of the Nat. to change
anything.

With regard to sending criticisms or abstracts of magazine articles, now I am
too busy with my paper to write any and if I am to leave sooner than I expected
for home I ought to spend my whole time in study . . .

You will of course imagine that I am anxious to know what will be done in
Cambridge this summer as my plans and travels may be so much affected thereby.
If the college determines to introduce the study of cryptogamic botany I hope
they wont forget that they must have microscopes. I think you will be disap-
pointed when you again see me in the amount of systematic fungology which I
have learnt but, on the other hand, I have studied development as applied to
some of the minute forms, to a much greater extent than I should have ever done
if I had not come abroad.

The work on the fern prothallus was not finished by Christmas, so instead
of going to Paris, Farlow joined his laboratory friends Rostafinski and
Gilkinet, in celebrating Christmas Eve with the lively De Bary family. On
the following day he wrote his mother and sister his description of the
festivities.

a2 Fartowia, VoL. 2, 1945

Strassburg, Dec. 25.
Dear Mother:

I take advantage of today to wish you all a merry Christmas. As far as I am
concerned the day is Sunday intensified, that is very much quieter than Sunday
is here. Last night we were invited to De Bary’s to see his Christmas tree. It
was all explained to me as though I had never seen a tree and Rostafinsky with
his usual impudence and fondness for giving good advice hoped that I would
introduce the custom into America. After the tree we had supper and at the end
I was presented with a plate of candy on the top of which was a mock sausage
for my benefit as I am so fond of such things. Today the weather is raw and
foggy and I gave up my project of a long walk thinking the less I got of such air
the better. This morning Rostafinsky made me a long call and this afternoon I
called on Dr. Alexander and also took a look at the Vespers in the Cathedral
where the altar was all lighted up. The soldiers had a tree in St. Thomas church
but as there are 18000 soldiers I imagine that if one tree held all the presents a
good many must have gone without. I should have liked to have seen the tree
as it stood directly in front of the celebrated monument to Marshal Saxe which
is very large and of white marble. Today the city has swarmed with soldiers and
many of them being new recruits the uniforms looked clean and fresh.

Last Saturday I went to Frankfurt to see the Websters . . . In the evening a
woman called who is keeping an American boarding house in Frankfurt. She had
her pockets filled with great red apples she had received from America. After
being so long in Strassburg the Frankfurters looked very swelly. The greatest
treat at the Webster’s is an American bed. That is, they don’t have the feather
plumons which the Germans always have. The plumon is a thing too large for a
pillow and not large enough to cover me although I am not very long. Conse-
quently as they don’t have such things as bedquilts and comforters one’s feet and
hands are always out in the cold...

Your affectionate son,
William

Strassburg, Dec. 25.
Dear Mary:

Unless you consider yourself included in ‘our’ family I will wish you and your
family a Merry Christmas. Last night I went to De Bary’s to see his tree. I
wasn’t there at the opening at six o’clock being too busy to go so early. Mrs.
Gilkinet was there and said there was a great running about and trying to look
through the key hole. When I arrived I found the Norway spruce already lighted
up. Instead of the hideous wax doll called the Christ child there was a bunch of
gilded pine cones which Mrs. De Bary had prepared for the occasion. There
were four children Wilhelm the oldest who was very sheepish and didn’t say a
word, August more lively, Marie, a rather too lively young lady six years old
with a pug nose and little black eyes which are generally half strained out of her
head by the two black pig tails on the back of her head, last night however her
hair was flying in all directions, and little Hermann with very red cheeks and
blue eyes. The presents were put on little tables near the tree. Hermann was
perfectly absorbed and could not say a word all the evening. All the boys had a
lot of soldiers. They seemed to have more toys than American children. Marie
had only four dolls and a doll house besides a lot of other stuff and the older
ones had books and games. The candies and cakes were a sight to behold and

Harris: CORRESPONDENCE OF WILLIAM G. FARLOW 33

came from all parts of Germany from Berlin to Strassburg. There was a young
law student named Von Jacoby from Berlin who had a table with presents. He
seemed to be a sort of protégé of Prof. De Bary’s. He had an enormous mouth
out of which he spilt his words. His conversation was principally about how
much beer they could drink in his Chor. On his table were a lot of gloves, and
such gloves. The Germans never wear any which are not ten sizes too large.
There was an indescribable thing which Rostafinsky told me was a Pommern
Gansebrust and a great delicacy. It was a great goose skin sewed up at both ends
and dripping with oil. It was full of uncooked smoked goosemeat and said to be
delicious! The Pommern geese are very celebrated. There were also Erlangen
Lebkuchen and Berlin gingerbread and two cans of condensed milk on his table.
Mrs, De Bary had amongst other things two bottles of arrack punch. At the
table the company appeared much interested in my description of peanuts which
they thought couldn’t be very nice and they couldn’t understand how our You
meant the same as their Du Ihr and Sie. They of course didn’t see the point
when I explained how senseless and inconvenient their Sie was. The other day
De Bary had a great present from America. A box containing a swamp pitcher
plant, some hickory nuts and twelve cranberries. As I said they were good to eat
they cooked them and liked them very well.

Today I dined with Gilkinet. I asked if his little Georgie 1114 months old
was coming home with his Alsatian nurse to spend Christmas. Ever since he was
born he has been farmed out to a peasant near Strassburg. His mother didn’t
know whether he would be at home or not today but had an idea, she wasn’t
quite certain, that he was to spend the day in some village near the Vosges. Such
independence in a child of his age would be remarkable in America. His mother
seemed very calm and philosophical on the subject. So is Prof. De Bary who
after giving Hermann a glass of wine before sending him to bed refused to let
August, aged 8 or 7 yrs., have any more because he brought on a typhoid fever
last year by drinking too much punch New Years.

Will.

A few more days of hard work brought the paper to its conclusion and
on New Year’s Day, 1874, Farlow wrote:

Today I send you the manuscript of an article on which I have been studying
some weeks. I presume you will think it very short for the time spent upon it
but all such work requires an everlasting amount of time. The article seems to
me not at all adapted to the Naturalist but rather to Silliman’s Journal. You are
at liberty to dispose of it as you please provided, wherever it appears, the draw-
ings are as good as those usually found in the Botanische Zeitung. I must make
this a condition because when printed I should feel bound to distribute a few
copies in this region and if the execution was not up to the mark there might be
some very invidious remarks made about American publications here. I wrote
the first draught in German and then afterwards wrote in English, having shown
the German to De Bary. I have made a German abstract which will appear at
once without plates in the Botanische Zeitung . . .

There may be a good many clerical errors in my manuscript but there is no
English speaking person here to whom I can show it. I should like to see the
proof of the figures I send before the article is published.

Gray acknowledged the paper with a brief note on January 11:

34 FaRLowIA, VoL. 2, 1945

Your New Year letter, and the Mss. & drawings have come to hand. I write
a hasty line to tell you so. I think of reading your paper to the Amer. Acad.
And, if they will engrave the drawings to print in the Proceed. — otherwise in
Naturalist unless Sill. Jour. will engrave on stone or copper.

We will see soon. Your paper is very interesting!

On February 15 Gray sent the first proof.

I read your paper to Amer. Academy — it is accepted for Proceedings. Here
is a rough, unread proof. I may keep it back till I can get corrections from you.

That the figures may be photographed for the new process of reproduction, I
have to get the drawings gone over in pen & india ink: pencil will not do.

Farlow returned the proof from Antibes, where he was enjoying at last
his long anticipated visit with Thuret, and he protested that the drawings
were not in pencil, but in neutral tint, and should not be inked over without
his supervision. On March 21 he wrote again from Antibes.

A week ago I returned the proof corrected which you were so kind as to send
me. Last night Rostafinsky and Janchewsky arrived and Rostafinsky said that
the fern prothalli are now grown up into Pteris cretica instead of Aspidium molle.
Today De Bary sent me a copy of the Botanische Zeitung with my short Ger-
man extract in which he had changed Aspidium molle to Pteris cretica through-
out. Will you be kind enough to do the same with the English or if it is too late
for that, to write a little note at the end saying that in consequence of the further
development of the prothalli enabling the species to be accurately determined
it is necessary to change the Aspidium for Pteris cretica throughout. I thought
myself that it was more probably Pteris cretica but De Bary thought the con-
trary. Now it is settled.

Gray replied disconcertingly that the “proof returned came just too late,”
and Farlow wrote again on May 4:

I reached Strassburg Saturday evening and, on my arrival, I found yours of
Apr. 10th which had been forwarded from Antibes. I am quite anxious about
the publication of my article as the preliminary remarks in the Botanische Zei-
tung have been attacked on all sides particularly in Berlin and Wurzburg. I don’t
understand exactly what you mean by my proof arriving too late. I hope you do
not mean that it has been published just as it was. If so, don’t let a copy get
into Germany for there were some horried mistakes in the quotation from Wi-
gand. I hope also that the drawings are well done otherwise the remarks of the
Germans, who don’t believe the thing because it is contrary to their theories,
will be disparaging to the last degree. For De Bary’s sake as well as my own I
must be extremely careful about the form in which the article appears.

The paper, titled Au Asexual Growth from the Prothallus of Pteris ser-
rulata, was published in the Proceedings of the American Academy, vol. 9,
1874, and the correction of the name to Pteris cretica had to be made in a
note at the end. That Farlow was far from satisfied with the appearance
of the article is evidenced in his letter of May 21 when he wrote:

As to the Fern Prothallus, I must confess that I was decidedly disappointed
in the appearance of the plates which fall very far short of those in the Bot. Zeit.

Harris: CORRESPONDENCE OF WILLIAM G. FARLOW 35

and the criticisms have been so unfavorable on the execution of the plates that
I hope you will not distribute any copies in Germany. I may have to translate
and reprint the article with better plates in the Bot. Zeit. As it is, I shall have
to leave any preparations, which I should like to take to America, with De Bary
who will show them to other botanists. I hope you have kept the drawings . . .

As Farlow’s sojourn in Europe drew near its close he became more anxious
about what he was to do on his return to America, and although Gray
hoped and expected to have a place for him at Harvard, there was as yet
no official confirmation of a position. On March 27 Gray wrote:

Presdt Eliot is in England, till May . . . My impression is that he wished in
spring to put you on to the Bussey-foundation, & do some work there — perhaps
start a fungus-laboratory — as well as here. I’ll be responsible that you get the
full of an Assistant-Prof. salary ...& in time more...

As the Bussey develops, I think it likely summer work may be in order there.
But if so, you will have an equivalent in winter. Do not fear about that.

The President is getting a great idea of the importance of research —and will
see that you have a chance. As to Goodale’s course of this summer, —as you
need not begin till 1st August, I advise you to agree to come & take it then —
give a month to it —if you intend to come home this fall at any rate. —If you
think of staying another winter, that is another matter.

I think it will be for your advantage to take hold of this work next summer —
tho’ it wont amount to much. It makes the right beginning.

The last letter from Gray which we have of this long correspondence
expressed once more his confidence in the establishment of a cryptogamic
laboratory.

Cambridge May 4, 1874
My dear Farlow.

Yours of April 11, came while I was in Washington. What I & Goodale had
previously written will by this time tell you all we can say. Presdt E. is not home
yet, — but may be expected to-morrow.

I see you are decided to come home this summer, any how. In that case, I
would, if I were you, get home before the end of July. But I would not hurry to
be here sooner. The summer course this year will not probably amount to much,
but will lay a foundation. The plan of Goodale taking it for July, & turning it
over to you in August is best — You will then work independently. And you can
go on into Sept. if you like. We shall, I doubt not be able to establish a labora-
tory for Cryptogamic & Anatomical Botany, in which you can make name &
fame .. . Your paper has done you great service. More of them, and your
success as teacher will do the rest .. .

I am sadly hurried — will write again soon.

Ever yours cordially

Asa Gray

Farlow sailed for home in June, 1874, and immediately received from
Harvard his appointment as Assistant Professor of Botany. He entered at
once upon the task of creating a laboratory in cryptogamic botany and

36 Fartow!A, Vor. 2, 1945

phytopathology, and his subsequent success at Bussey and later at Cam-
bridge has fully justified the years of preparation that had gone into his
European study. He continued to keep in contact with his friends in
Europe, and his correspondence with De Bary was maintained until the
latter’s death. A letter written by De Bary in the summer of 1877 is of
special interest, as by giving news of the whereabouts of the students in the
laboratory, and of the completion of De Bary’s own book, it brings to a
natural close the narrative of Farlow’s two years at Strasbourg.

Montreux, August 17, 1877
Dear Mr. Farlow:

From the stamp you will see that I have taken refuge in order to get peace, not
in the desert like the ancient prophets, whom I do not emulate in general, but at
that very corner on Lake Leman which you will remember as beautiful and quiet,
especially at this time of year, because now people think it is too hot, while in
winter the English and Berliners abound here. When you imagine this charming
bit of land and water where redwood and giant trees, together with laurel, Lauro-
cerasus, Punica, and Passiflora are being reflected in the blue surface of the lake,
and right now the whole atmosphere is filled with the scent of the flowers of
Ligustrum japonicum — while nearby the Phyllachora Congress is having its in-
ternational meetings, and the wine ripens untouched by the disease these gentle-
men are talking of —if you imagine all this, your heart will probably become
reminiscent too, and you will forgive an old sinner inasmuch as he has retired to
this idyllic spot in order to write you at last.

The narrator may proceed subjectively, so, I start with the news that the
Anatomy is entirely finished four to six weeks ago, 640 plus 20 pages strong. I
have been working desperately on it since last summer, except for a short inter-
ruption which I shall mention later. That’s why you did not hear from me for
so long a time. Two proofs which may have surprised you in April have been
sent to you as a mute sign of life on my part . . . Since the middle of July I
have finished everything excepting the last proofs. I have become human again,
but I had to arrange and clean up so many things, and consider so many other
matters that I could not find time to write a detailed letter.

The cleaning-up-process also includes working up the material that has col-
lected in the last three or four years for your ‘Pteris story.’ I have finished this
during these last days, and I hope to have cleared up this matter now as far as
three species of ferns are concerned. Your side is thoroughly correct. It turns
out — when compared exactly — to be a very instructive special case of a com-
mon phenomenon. With this I have come to answering your letters, for yours of
last October starts with Pteris. As for the complaints which you add to your
remarks, I can answer only with the wish that you may not take to heart what
the American botanophiles and mycophiles say about you and Cooke. If you
continue in the same way as in the last papers you sent me, these gentlemen will
certainly come to reason, if not all, then at least many of them. It’s not different
with me: Berkeley and his set always consider my kind as very wretched fellows,
and still, by now, all intelligent people know what they should think of it. The
only way to achieve something, is not to let yourself be bewildered. I had a lot
more trouble with Hallier, Sollman, Hoffman etc. than anybody has nowadays.

To continue with your mycological questions —I see in Fischer von Wald-
heim’s new survey of the Ustilaginales that he indicates Urocystis colchici on

Harris: CoRRESPONDENCE OF WILLIAM G. FARLOW oy

several monocotyledons, also on Allium (A. rotundum). On another Allium,
A. magicum, he indicates another Urocystis species, U. magica. This is about all
that is known about Urocystis on Allium. Considering the great similarity of the
spore characters of most of the species a decision on the identity of species seems
to be very difficult to me. Why are not all species of Urocystis, or all species on
monocotyledons, a single species? I shall send your fungus to Fischer, so he can
compare it with his materials. But I shall wait till winter because he is probably
absent from Warsaw now...

You know all that’s important about Rostafinski, that is that he is in Krakov,
very industrious, and with good success. I would not know much more myself
without going into details. As for Stahl, you probably don’t know that he is
privatdocent in Wiirzburg. He went there in the summer, and is very satisfied
with everything thus far. I talked with him recently in Strasbourg, and I hope
he will also write me.

You do not have to fear that Gilkinet has forgotten you even if he should not
have written to you in the meantime. I can tell you so with certainty, for we
talked about you very much when I visited him last April in Liittich. Here in
Strasbourg we had the same experience with him as you had — not a word from
him. In April I had a chance to go to the Horticulture Exposition in Amsterdam,
and I took advantage of this trip to visit him in Littich. After a very friendly
reception, I soon found out the good reason for his personal and botanical silence
—he has accepted the position of chargé de cours at the University, that is to
say for pharmaceutical chemistry. He has to manage a chemical laboratory and,
at least for a long time, he was entirely absorbed by the necessity of remodeling
himself for work in this new job. This is understandable and it will justify him
also in regard to you...

There is not much to say about Solms except that he is as you know him —
short active semesters, alternating with long vacation trips. Last fall he was in
Moscow and Stockholm, at Easter in Athens. Otherwise there were many changes
in this laboratory. Dr. Wilhelm, whom you knew, is the only stable man. This
summer it was very lonely, since, because of illness and other reasons, only two
people were working. I shall send you some dissertations which were made dur-
ing the last year...

In order to achieve a symmetrical conclusion to this letter, I shall talk sub-
jectively again. In my house, everyone is rather well, and so am I, especially the
last four to six weeks. I accompanied my family to Switzerland yesterday — they
are in Interlaken again. I left them for a week to go here, and then to Bex
(Rhéne Valley) to the Swiss Meeting of Natural Scientists, and then to Inter-
laken, too. In mid-September we shall be at home again.

Goodby for this time, best regards from my family and myself, and do not.
punish me by too long a silence.

Sincerely yours
A. De Bary
Harvarp UNIVERSITY
CAMBRIDGE, MASSACHUSETTS

2(1): 39-51 FARLOWIA January, 1945

NOTES ON FARLOW’S AGARICALES FROM CHOCORUA
RouF SINGER !

To those who are acquainted with the White Mountains of New Hamp-
shire, the name Chocorua brings to mind the beautiful lake in which is
reflected picturesque Mount Chocorua, but few realize that the region is a
mycologist’s paradise. In this region of beeches, birches, maples, oaks,
pines and hemlock at the lower elevations and of spruce and balsam below
the nearly bald granitic summit of the mountain, there are habitats of
widely varied types and ecological conditions that make possible the oc-
currence of many different species of fungi.

Whether it was beauty of the scenery, the presence of congenial faculty
colleagues, or the presence of these varied and excellent collecting grounds
or some other factor that caused Dr. Farlow to choose Chocorua as his
summer home it is impossible to say. However, it is obvious from the large
number of fungi in his herbarium which were collected in this region, that
when he left his office on the top floor of the Botanical Museum in Cam-
bridge where his herbarium was at that time maintained, he lost few oppor-
tunities for going out into the woods of Chocorua and vicinity to make
collections of fungi, many of which he sent to his colleagues who were
interested in special groups. Among these colleagues was his friend and
former student, Dr. E. A. Burt, who cites many of Farlow’s Chocorua
specimens in his monographic studies of the Thelephoraceae and who in
1909 wrote to Farlow that he was turning up many unusual and interesting
species of Hypochnus (i.e. Tomentella) in that region. While he was mak-
ing these miscellaneous collections, he also had in mind the eventual pub-
lication of an illustrated agaric or mushroom flora and it was at Chocorua
that Joseph Bridgham and L. C. C. Krieger, under his critical supervision,
executed in water color and at natural size a large number of exquisite
illustrations, a small number of which have now appeared in the ‘‘Icones
Farlowianae.” The larger proportion of the paintings that remain unpub-
lished, and unfortunately not available except to those mycologists who
come to the Farlow Herbarium to study specimens and plates, nevertheless
_are extremely valuable as permanent records of the fungi as they appeared
when freshly collected. They are also of value because they illustrate
species which, although Farlow collected and determined many of them,
were in many instances seen and verified by Charles Peck of the State
Museum at Albany, the foremost American agaricologist of the period, as
well as by European specialists in the group. Thus Chocorua was the
source of a large number of specimens, not only for Dr. Farlow but to
many colleagues with whom they were shared, as well as for those who

+ The writer is indebted to Dr. David H. Linder for the introduction to this article.

39

40 FaRLOWIA, VoL. 2, 1945

were fortunate enough to be guests at his summer home and do their own
collecting. However, despite the sharing of specimens and his efforts to
have identified accurately all those fungi that were illustrated by his artists,
there nevertheless remain many that are still to be identified. Also, there
are others which, although named as accurately as the knowledge of the
time permitted, are worthy of additional notes. A few of these are dis-
cussed in the following paragraphs.

I. Surirtus GRANULATUS IN NortH AMERICA AND EUROPE

One of the common species observed near Chocorua is a bolete generally
determined as Boletus granulatus L., and under this name illustrated and
described in the [cones Farlowianae(3). This species, described by Linnaeus
and Fries from Sweden, has previously been studied by the author in
Europe. It was, therefore, interesting for him to compare his notes with
data gathered on abundant living material at Chocorua. We shall at this
point anticipate our eventual conclusion, and stating that the American
form actually differs macroscopically, microscopically, and ecologically
from the European type, name it Suillus 2 granulatus ssp. Snellii * Sing.
ssp. nov.

Suillus granulatus ssp. Snellii ssp. nov.

Notis macroscopicis subspeciem leptopodem fortiter revocantibus in mentem; notis
chemicis haud notabilibus in comparatione cum typo; sporis minoribus (6.8-8.2x3 mu)
a subspeciebus ceteris recedens; habitatio: cum Pinibus strobis (rarissime cum aliis coni-
feris) ad terram plerumque arenosam gregatim in America boreali; collectio typica in
Farlow Herbarium conservata atque a D. H. Linder & R. Singer prope Chocorua, N. H.
collecta est.

Macroscopical characters: There is no need of wasting space by rede-
scribing S. granulatus ssp. Snellii macroscopically. The reader will find
good descriptions of it in most American books and papers dealing with
boletes, e.g. in “Icones Farlowianae,”’ Atkinson, and others. We may only
add that the spore print is ‘Isabella color.” 4

Microscopical characters: Spores pale yellowish or pale melleous, thin-
walled ellipsoid-subfusiform, frequently attenuate toward the upper end
and usually with a subhilar depression, 6.8-8.2 x 3-(3.5) pw; basidia
clavate, 4-spored, hyaline, but at some spots covered with brownish resin-
ous masses emanating from the cystidia, 20-24 x 6-7 p; cystidia frequent,
extremely crowded on the pores where their excretions or incrustations
cover large portions of the hymenium, honey color or hyaline when seen
without the incrustation (after crushing the preparation in an alkaline

* Since it has been proved by D. P. Rogers that S. F. Gray’s “Natural Arrangement”
is post-Friesian, the author thinks that Snell’s (12) proposal of the name Swallus instead
of Ixocomus is justified.

*'W.H. Snell was the first accurately and reliably to indicate one of the main char-
acters which separate the American subspecies, i.e. the small spore size; we therefore feel
that it is most appropriate to dedicate this subspecies to him.

* All colors cited in quotation marks are those of Ridgway unless indicated otherwise.

SINGER: AGARICALES FROM CHOCORUA 41

medium), thin-walled, smooth, clavate-fusoid, more rarely claviform or
fusiform, or ampullaceous, or subcapitate, or a combination of these, 34-38
x 6-10.5 3 trama of the tube walls divergent and with a distinct regular
mediostratum; cuticle with a distinct ixotrichoderm, some of the hyphae
somewhat incrusted; kyphae of the context hyaline, thin-walled, consist-
ently without clamp connections; glands consisting of masses of strongly
incrusted and pigmented or rarely hyaline dermatocystidia similar to the
ones observed on the pore mouths, but more variable in size.

Chemical characters: KOH on the surface of the pileus: lilac, becoming
grayish brown in a few seconds, young stages more intensely and persist-
ently lilac; on the context of the pileus: lilac, becoming grayish brown,
young stages usually not showing any reaction except where the context is
showing the first yellowish spots (near the tubes); on the pores and tubes:
almost black immediately. — NH (prolonged action) and NH,OH on
the surface of the pileus: slowly lilaceous brown, in young stages slowly
gray or grayish brown; on the context of the pileus: lilac (except for young
stages with no appreciable reaction) ; on the pores and tubes: lilac (except
for young stages with no appreciable reaction).

Habitat: On the ground in coniferous and mixed woods, in mycorrhizal
connection with Pinus strobus and P. monticola, rarely, near stands of these
pines, transgressing into pine-free stands of hemlock (Tsuga) or fir (Abies).
Fruiting from June to November.

Distribution: The distribution of this subspecies has not been established
satisfactorily because of the fact that many species have been wrongly de-
termined as “Boletus granulatus” in the past. The geographic area of ssp.
Snellii most probably coincides with that of the five-needle pines with which
it is associated.

COMPARISON WITH CLOSELY RELATED SUBSPECIES AND SPECIES OF SUILLUS

1. Suillus granulatus (L. ex Fr.) Snell ssp. typicus. In the Ameri-
can form the context, tubes and the stipe remain white or pallid for a
longer period than in ssp. typicus; the color of the pileus and the glands of
the stipe reach darker tones than in the latter. In our specimens from
- Chocorua as well as from Massachusetts and New York, the color of the
well-pigmented pilei was about “orange cinnamon,” sometimes near “‘cin-
namon rufous,” or darker than these, and then reaching almost all shades
familiar in Suillus luteus (L. ex Fr.) S. F. Gray, but pale or bleached
forms are not uncommon, the bleaching starting mostly in the center of
the pileus; the glands tend to become purple or blackish long before such
a change is seen in the European form. There are, however, cases where
the glands remain rather inconspicuous in the American form, and others
where they darken early in Europe. The former should be interpreted as
transitions toward Suillus brevipes (Peck) Sing. comb. nov. (Boletus
brevipes Peck) while the latter appear to be transitions toward another
subspecies of S. granulatus which we shall discuss later. The macroscopical
difference between the European type and the American form is easy to see

42 FarLowiA, Vor. 2, 1945

when good colored illustrations of both these forms are compared side by
side, though the comparison of dried material does not show striking
differences.

The microscopical differences between these two forms are restricted to
the spore size. This, however, is the most important distinguishing charac-
ter. The spores of a specimen collected by Buchholtz in the Mikhailovski
Park in the Moscow region were found to be 8.3-9.8 x 3.3-3.5 p, and other
notes on ssp. typicus contain similar measurements which, in comparison
with ssp. Snellii, are decidedly larger. The other microscopical characters
do not seem to differ noticeably in these plants.

The chemical characters of the European form are nearly identical with
the chemical characters given above for ssp. Snellii.

2. Suillus granulatus ssp. leptopus (Pers.) Sing. comb. nov. (Bo-
letus circinans ssp. leptopus Pers.) (8). This is the same as the species
this author indicated under the name /xocomus Bellinii (Inz.) Maire.® Cer-
tain features of the American form, ssp. Snellii, seem to indicate a close
relationship with ssp. leptopus. However, the spores of ssp. leptopus are
not small as in ssp. Snellii but large as in ssp. typicus. The author found
them 8.5—-10 x 3.5-4.5 , and occasionally reaching 16.7x7 p, and the
basidia 27-29.5 x 6.8-9.5 4 in a specimen from Hammam EI Cif, Tunisia,
collected by Bénier and determined by Patouillard.

3. Suillus placidus (Bon.) Sing. comb. nov. (Boletus placidus Bon.).
This is the same as Boletus albus Peck as found abundantly in Massachu-
setts and New York as well as in Europe by the author. Kallenbach thinks
that this is the same as Boletus Bellinii or what we call Suillus granulatus
ssp. leptopus. But this confusion merely results from the fact that Kallen-
bach does not know the latter except from the literature. We have found
that though S. granulatus ssp. Snellii and S. placidus often grow associated
with one single tree, side by side, there is no difficulty distinguishing these
species macroscopically.

The spore size of S. placidus is about intermediate between S. granulatus
ssp. leptopus and ssp. Snellii. This difference even though slight is constant.
It is in accord with European measurements. Snell (11) has also indicated
the slightly larger size of the spores of S. placidus when compared with the
American form of S. granulatus.

While thus the microscopical difference between S. granulatus ssp. lep-
topus and S. placidus is very slight, the chemical characters which are about
the same in all subspecies of S. granulatus including ssp. leptopus, tend to
separate the latter from S. placidus. The ammonia reaction tends more
towards red tones in adult specimens of S. placidus than in S. granulatus
sensu lato. It is remarkable (and should be kept in mind when the useful-
ness of chemical reactions in boletes is discussed) how constant these re-
actions are if specimens of corresponding stages are compared. In the

° Revue de Mycologie 3: 48. 1938. “Et si nous preferons de le classifier comme va-
riété de ce-dernier (J. granulatus), nous pouvons reprendre le nom de Persoon.” Konrad
(6) considered it as a subspecies of S. placidus; Gilbert (4), as an independent species.

SINGER: AGARICALES FROM CHOCORUA 43

writer’s paper cited above (9), this same reddish reaction was indicated
from specimens growing on Pinus cembra on granite soil in Europe, and
later was checked on specimens growing under Pinus strobus on sandy soil
in the Adirondack Mts. of New York, U. S. A. The results were identical.
In order to give the reader the full descriptive facts as drawn from Ameri-
can material of S. placidus for comparison with S. granulatus, American
form, we cite our notes on the chemical characters of the former species:

KOH on the surface of the pileus: lilac to violet, after a while becoming
deep reddish brown to chocolate, in young stages occasionally salmoneous
or purplish salmoneous at first, lilac or violet after a while; on the context
of the pileus: lilac to violet, strongest on the yellowish colored places in
very old specimens, and only on these places (if there are any) in younger
material; on pores and tubes: reddish brown in adult specimens, but no
reaction or else a slow change to pale sordid brown in young stages. NH3
on the surface of the pileus: lilac to violet; same reaction on the context,
but none when young; on pores and tubes: initially salmon color to reddish
tan, but no reaction when young. NH,OH on surface of the pileus:
stronger and deeper than with NH3; on context of the pileus: same re-
action as with NH; on pores and tubes: blue to gray, but no reaction on
young stages; in the very old yellow stage the tubes reach “zinc orange”
(R.) with NH3-vapors, and the surface of the pileus becomes brownish-
red, the context of the pileus salmon color under their influence.

The above data prove that Kallenbach’s opinion (5) is erroneous. Kal-
lenbach thinks that he can divide S. placidus into four “formae”: 1. f. Pini
strobi; 2. f. Pini cembrae; 3 {. Pini halepensis (our ssp. Bellini); 4. f.
americana. As it looks to us, if there are any forms to be distinguished, they
should be f. Pini cembrae and f. americana (in Kallenbach’s terminology).
The European plant on P. strobus is either introduced with the trees, and
therefore the same as f. americana, or it has originated from the European
form on P. cembra (which is much less probable), and then it would be the
same as f. Pini cembrae since there is no evidence of acquired specialization
(inability to grow on the old host, P. cembra), nor any morphological or
chemical differentiation. Kallenbach’s f. Pini halepensis is not a form of
S. placidus as shown above. Consequently, all he says about the geographic
area of his S. placidus is to be understood “cum grano salis.” The African
and Asiatic localities given refer to S. granulatus ssp. leptopus. The Eu-
ropean ones are almost irreparably mixed up.

MYCORRHIZAL CONNECTIONS OBSERVED IN THE RACES OF S. GRANULATUS

The writer has studied the typical European form in Central and Eastern
Europe. It has consistently been found in the neighborhood of Pinus sil-
vestris, in some cases under Pinus montana (in the Alps) and Pinus uncinata
(in the Pyrenees); it has also been collected under Pinus austriaca in the
Wiener Wald. This latter location, however, may not be the correct place .
to look for the typical S. granulatus; at least, my notes seem insufficient for
definite conclusions when seen in the light of more recent developments.

44 FaRLowIA, VoL. 2, 1945

The Mediterranean form, S. granulatus ssp. leptopus, was studied by the
writer in large numbers on the sand dunes of the Mediterranean Coast near
Prat de Llobregat, Catalonia, Spain, in 1934. They were associated with
pine, mostly Pinus pinea, and were fruiting most plentifully in October. In
Africa they reach a maximum in January and February. Other hosts or
symbionts are: Pinus halepensis, P. pinaster, and P. pithyusa. In mixed
stands of these Mediterranean pines and Pinus silvestris some intermediate
forms may be observed, thus the soil also may play an important réle in
this. Besides, old specimens of the Mediterranean race often are hard to
distinguish from S. granulatus typicus.

The American form, in contrast with the European forms, grows with
Pinus strobus, at least preferably so. The Chocorua specimens, observed by
D. H. Linder and the writer, were obviously connected with this five-needle
pine, and occurred as well in pure pine stands as in mixed woods with
scattered pines. However, some of the fruit bodies were too far from any
pines to assume connection between their mycelium and the pine roots.
They grew in a Fagus grandifolia stand with intermixed Tsuga canadensis,
and were obviously connected with the latter. They were not otherwise
different from the pinophilous majority. In Newcomb Co., N. Y., where
the boletes were likewise found to be connected with Pinus strobus, a sim-
ilar ecological anomaly was observed when a form similar to S. granulatus
in a few rare cases grew far from pines in a very open stand of Populus
tremuloides and Salix spec. with numerous intermixed firs (Abies balsamea),
and was clearly connected with the latter. But in this case, the fruit bodies
seemed to be morphologically modified: they were somewhat larger than
the average pinophilous plants and the stipe was pure white as in Suillus
brevipes (Peck) Sing. comb. nov. (Boletus brevipes Peck), and not dotted-
glandulose in its lower third to three-quarters, and only minutely granulose
in its upper portion; it was rather elongate and slightly attenuate or swollen
below; the margin of the pileus was strongly fibrillose-tomentose, probably
from a rudimentary veil as occasionally observed in S. brevipes; the tubes
were strongly decurrent as in a certain form of S. brevipes, growing under
abnormal ecological conditions (in dense mesophytic hammocks) in Florida
(this form was described as Boletus pseudogranulatus Murr.) ; the context
showed reddish discoloration. All this points to the conclusion that the
American form of S. granulatus is not, as the European races probably are,
at least when growing spontaneously in nature, confined to very definite
soil conditions and to a certain limited number of hosts (two-needle pines
only), but able under certain conditions to pass over to other neighboring
conifers, either with slight morphological changes (as in New York with
Abies °), or without any such changes (as in Chocorua). In neither case
did the soil contain a considerable amount of lime. In the light of recent

° At the present stage of my studies, I do not think that it would be wise to insist on
the above determination of the specimens concerned. It is also possible that they were
not closest to the American form of S. granulatus, but rather an extreme form of what
Peck called Boletus albidipes.

SINGER: AGARICALES FROM CHOCORUA 45

publications it seems probable that the western white pine (Pinus monti-
cola) is another symbiont of S. granulatus ssp. Snellii, a relationship which,
considering the taxonomic position of this pine, would not be surprising
at all.

As for the ecological characters of S. placidus it is well known that this
species grows in symbiosis with two five-needle pines, P. strobus and P.
cembra. It thus appears to be closest ecologically to S. granulatus ssp.
Snellii, not ssp. leptopus.

Consequently each of the races of Swillus granulatus evidently has, in
nature, its own requirements as to mycorrhizal hosts. Since these hosts
have very definite geographic areas, the fungi concerned have them also.
All this corroborates the statement that there are myco-ecological as well
as geographic races for which we use the taxonomic term subspecies. S.
placidus, however, not having a host of its own, yet morphologically and
chemically more differentiated than the other forms discussed, must be
treated as a separate species. I am aware of the fact that many taxonomists
prefer to call forms like ssp. Smellii autonomous species rather than sub-
species. More detailed information on Suillus albidipes (Peck) Sing.
comb. nov. (Boletus albidipes Peck) will be needed to decide whether in
this case we have to deal with a group of small species, or rather with a
complex species consisting of several subspecies, possibly including even
S, albidipes and S. brevipes.

II. REDESCRIPTION AND NOTES ON BOLETUS RUBINELLUS PECK

Macroscopical characters: Pileus “light Corinthian red,” “Corinthian
red,” “Etruscan red,” “deep Corinthian red,” and often reaching “Indian
red” in the middle, often more brownish tones mixed in when old, or bleach-
ing to a uniform pale yellow, but normally only the margin, if at all, light
yellow to yellowish white, warty-floccose, occasionally almost squarrulose-
verrucose on the disc, more faintly tomentose on the margin, the flocculae
more or less detersible, subviscid when wet, dry in dry weather, the margin
sterile (about 0.5-2 mm.), broadly conical or convex, eventually often
broadly umbonate, 10-50 mm. broad. — Tubes (bluish) pink to salmo-
neous, or dingy reddish, eventually with a brownish tone, adnate or de-
pressed around the stipe, medium long (about 5 mm.), pores initially
daedaleoid, later angular, and rather small to rather wide (up to 1 mm. in
diameter), sometimes radially elongate near the stipe. Spore print fer-
ruginous-brown. — Stipe subconcolorous, often mixed with yellow or yel-
lowish white, subglabrous to slightly pustulate-subflocculose, dry or almost
so, smooth or almost so, equal or slightly enlarged near the base or with
widened apex, solid, 10-50x 2-10 mm.— Context whitish yellow to
brownish yellow, usually more intensely colored in the stipe, or near the
base with some pink portions in some specimens; odor none; taste mild. —
Habitat: On the ground in woods, always under conifers. July to Septem-
ber. Mostly gregarious. Infrequent to locally moderately frequent.

Microscopical characters: Spores 9.5—13.8 x 34.2 p, long ellipsoid, some

46 FartowiA, Vor. 2, 1945

more fusoid, smooth, thin-walled, brownish to melleous, rather pale; basidia
25-32 x 8-10.5 p, 4-spored; cystidia usually ampullaceous or fusoid with
a bottleneck-shaped apex, but also subulate or fusiform without an elongate
apex, 38-52x 7-10 yw, the “bottleneck” if present about 3-3.8 » thick,
hyaline or more rarely brown, not as conspicuously incrusted as in S. pipe-
ratus,’ rather numerous and occurring on the pores as well as in the tubes;
trama of the tubes well differentiated in a completely hyaline, more gelat-
inized and consequently somewhat loosely arranged and slightly but dis-
tinctly divergent lateral stratum, and a slightly colored, denser, regular
mediostratum, though in mature specimens this differentiation becoming
more and more indistinct, and the hyphae more and more parallel, without
clamps, but with occasional nodules or bridge-like connections between
parallel hyphae; warts of the covering layer consisting of long, cylindric,
hyaline, thin (1.5—8 » in diameter) hyphae with thin walls and without
clamp connections, their wall more or less gelatinized and imbedded in a
gelatinous mass, their free ends obtuse, and sometimes with strange pro-
tuberances.

Chemical characters: KOH on the surface of the pileus: blackish, then
becoming brown, eventually brown-red; on pores and tubes: dirty brown;
on context: tawny brown. — NH,OH on the surface of the pileus: yellow,
then becoming olive yellow; on the pores and tubes: deeper pink, eventu-
ally gray. — Aniline oil on the surface of the pileus: characteristically
rapidly deep grayish purple. — Formol: no reaction.

The above description has been drawn from collections made by D. H.
Linder and R. Singer near Chocorua, N. H., and additional fresh material
collected by the writer near Newcomb, N. Y. This material has been com-
pared with the type at Albany which macro- and microscopically proved to
be identical. The type of Boletus Vanderbiltianus Murr., collected in North
Carolina and preserved at the New York Botanical Garden, is the same
species. It is said to grow in thin oak woods, but Dr. H. D. House who
collected it there remembers that there were conifers scattered in the woods
(oral communication). On the other hand, Coker and Beers (2), describe
and figure (p. 38, pl. 3, fig. 2-3, and pl. 62, fig. 13-14) this species in a
way suggesting that some other species must have been mistaken for Boletus
rubinellus. This opinion is in accordance with the fact that these authors
do not recognize Boletus Vanderbiltianus as a synonym of B. rubinellus,
but say, instead, that they have not collected the former. Furthermore, in
a note concerning their conception of Boletus rubinellus, they suggest that
it “may be only a form” of Suillus piperatus. This is probably correct as
far as B. rubinellus sensu Coker & Beers is concerned, but in our opinion
their species is the same as Suillus piperatus var. amarellus (Quél.)
Sing. comb. nov. (Boletus amarellus Quél., Boletus Pierrhuguesii Boudier,
Boletus rubrotubifer Kauffm.), a species which I collected in fresh condi-

‘Suillus piperatus (Bull. ex Fr.) Sing. comb. nov. (Boletus piperatus Bull. ex Fr.,
Ixocomus piperatus Quél., Chalciporus piperatus Bat.)

SINGER: AGARICALES FROM CHOCORUA 47

tion in the Pyrenees, and which is mild and also has the other characters of
the North Carolina fungus. The true B. rubinellus is a northern species,
more frequent in the Adirondacks and the White Mountains, reaching Ken-
tucky in the West, and the mountains of North Carolina and Tennessee in
the South. It does not occur in Europe. As for its affinities, the gelatinized
surface hyphae and the color of the tubes and spores, as well as its habitat
near conifers suggest a close affinity with the section Piperati of the genus
Suillus (Ixocomus sec. Piperati Sing. 1938, genus Chalciporus Bat. 1908).
This relationship is confirmed by the anatomy of the tube walls which is
quite typical for a Swéllus. The chemical reactions also agree with those
indicated by Singer (9) for this group. Consequently, the new combination
Suillus rubinellus (Peck) Sing. is proposed.

III. On LACTARIUS NIGROVIOLASCENS ATK. EX BuRL.

Macroscopical description: Pileus dark brownish gray or dark grayish
brown to almost black, opaque, pruinate-tomentose to strongly velutinous,
dry, smooth, convex, eventually depressed, but usually only in the center,
and, at the same time, showing a small but distinct papilla, especially when
young, 33-73 mm. broad; the margin frequently short-sulcate or coarsely
crenate as in many Leucopaxilli.— Lamellae light ochraceous yellow,
arcuate sickle-shaped, rather narrow: 3-4 mm. broad, acute in front and
behind, not anastomosing, moderately close to subdistant, polydymous,
simple, decurrent or adnate but from the point of attachment down the
apex of the stipe proliferating into very short and low parallel ribs. Color
of the spore print on white paper yellow, “F” (Crawshay).— Stipe con-
colorous, whitish at the extreme base which is buried in moss, versiform but
usually subequal or equal with only the extreme base tapering downward,
faintly tomentose-velutinous, solid, 50-75 x 7-15 mm.— Context white,
becoming dark violet when wounded. Latex white, showing dark violet
spots when exposed to the air in contact with the context. Odor not sig-
nificant. Taste mild. — Habitat: Coniferous wood or mixed wood under
conifers, on moss or sometimes on naked soil.

Microscopical description: Spores 8-13—(15) x 7-11.5—(13) yu, subglo-
bose, with conspicuous 1.5—2 » high ridges forming an ornamentation of
the type II or IIIa, more or less asymmetric; in authentic print from N. Y.,
10.5—11.7 x 8.5-10.5 » with a moderately dense to rather loosely arranged
ornamentation of type IT to IIIa (as above), or IIIb, more rarely IV or V,
the spines and ridges usually connected by an extremely faint network
which is easily overlooked, 1.3-2.3 » high; basidia 27-46 x 9-13 yp, 4-
spored, but very few 2-spored ones intermixed; cystidia none; cheilo-
cystidia on the heteromorphous edge of the lamellae versiform (cylindric,
clavate-balloon-shaped, fusoid, or basidiomorphous), hyaline, 14-52 x 3.3-8
pw; cuticle of the type called Virescens-type in Russula, consisting of a
subcutis of globose cells (about 17-20 x 15-17 4) which mostly proliferate
into a subulate, more rarely clavate or cylindrical hair with narrowed or
broadly rounded apex and 0-2 septa (not counting the wall between the

48 FartowlIA, VoL. 2, 1945

subcuticular cells and the hair); these hairs, 20-120 x 6.5—9 uy, are strictly
erect and form the epicutis; pigment grayish brown, dissolved in the cell-
sap; all hyphae without clamp connections.

OBSERVATIONS ON THE ANATOMY OF THE CUTICLE, THE SPORE WALL
AND THE HISTORY OF THE SPECIES

The above description shows that this interesting and rare species be-
longs in the section Plinthogali Burl., because of the structure of the surface
and the combination of the macroscopical characters. The structure of the
cuticle, strongly reminiscent of what is known as Virescens-structure in the
genus Russula, has never been described correctly in any species of Lac-
tarius where it is characteristic for the whole section Plinthogali. We have
observed this structure in L. fuligineus, L. azonites, L. lignyotus, L. acer,
L. nigroviolascens, a southern species with unchanging latex, and a new
Florida species with yellow watery latex. Unfortunately, Neuhoff in his
work on the ‘“Milchlinge” (7), misinterpreted the structure of the Plintho-
gali by taking the cellular subcutis for the outer layer, and the hairs for
the inner layer. Nothing like this, however, can be observed in any Euro-
pean or American Lactarius. The spores of L. nigroviolascens, as described
above, seemed to be more cristulate than the description given by G. S.
Burlingham (1) would indicate. However, Miss Burlingham was kind
enough to send me some authentic material, probably from the type locality,
or not far away from it, consisting of a good spore print obtained from a
specimen that she herself collected near Ithaca, N. Y. If the extremely faint
connecting veins found here between the warts and ridges are not taken in
consideration, at least a minority of spores of this print shows about the
same marks as two of the three spores figured (1. c., fig. 37). It must, there-
fore, be admitted that the ornamentation of the spores of L. nigroviolascens
Atk. ex Burl. varies to a certain amount not only in different collections
but also occasionally in a single print. The uniformly cristulate spores were
observed in all our New Hampshire material. Farlow had it from Chocorua,
Shelburne, and other places in New Hampshire, and his collections remained
partly undetermined, or were tentatively determined as L. lignyotus Fr., a
closely related but different species, easily distinguishable by the pink to
latericious color of the wounds. Fresh material in excellent condition was
collected by D. H. Linder and R. Singer in the same region where Farlow
collected this species for the first time, i.e. in 1891, and 1910. Farlow had
the essential data on this species, and gave correct notes with the specimens,
indicating how it differs from similar Lactarii. Though realizing it was
different from all species known to him, he did not describe it as a new
species. At that time, many mycologists would have described a species on
far less evidence than Farlow had, but Farlow was much more conscientious
in this regard. Incidentally, Atkinson, who collected his specimens many
years after Farlow, did not describe them either, but gave them the her-
barium name Lactarius nigroviolascens which was later (1932) validly
published by Miss Burlingham.

SINGER: AGARICALES FROM CHOCORUA 49

IV. A NEW SPECIES OF RHODOPHYLLUS: R. FARLOWII
COLOR PLATE

Rhodophyllus Farlowii Sing. spec. nov.

Pileo laete aurantiaco-luteo, non fortiter hygrophano, convexo, umbilicato, ad mar-
ginem pellucide striato, levi, glabro, haud viscido, 24-40 mm. lato; lamellis salmoneis
vel pallide salmoneo-latericiis, subangustis vel moderate latis, adnatis vel dente decur-
rentibus, confertis vel subconfertis; stipite concolori pallidioreve, glabro, levi, sicco, cavo
vel farcto-subcavo vel solido, aequali vel ad apicem subattenuato, 50-76x 2-6 mm.;
carne alba vel subconcolori in adultis, inodora; habitatio ad ligna et humum in silvis
montanis et in paludosis, aestate.

Pileus rich orange-yellow, more yellow between the deeper orange striae,
tending to bleach from the center outward, and somewhat hygrophanous,
convex with a distinct broad umbilicus, the margin transparently striate
about halfway to the center, smooth, glabrous, not viscid, 21-40 mm. in
diameter. — Lamellae salmon-pink to salmon-pale-lateritious, 2.5—5 mm.
broad, broadest in the middle or in the inner third, adnate with decurrent
tooth or subdecurrent to decurrent, close or subclose. Spore print pink. —
Stipe concolorous with the pileus, or slightly paler, the base more whitish,
glabrous, smooth, dry, hollow or solid, or stuffed-subcavous, equal or
slightly tapering upward, 50-76 x 2-6 mm., usually about 4 mm. broad at
the base, and 2-3 mm. at the apex. — Context white, at times somewhat
concolorous when old. Odor insignificant. — Habitat: On wood or on
humus in mountain forests and swamps, under Tsuga and other trees in
mixed stands. July and August.

Microscopical description: Spores 7-10 x 5.2-6.3 yw, pale pink, approxi-
mately hexagonal and the shape of a benzine ring when seen frontally, or
the sides odulose and the spores thus becoming symmetrical-complex, their
hilar portion forming an angle of about 90° when seen frontally; basidia
about 50x 14 y»; cystidia not seen.

This beautiful species is easily mistaken for a Hygrocybe (Hygropho-
raceae) because of its color and habit. It has been found by D. H. Linder
and the writer in July, 1941, near Chocorua, N. H., and was compared with
dry material preserved at the Farlow Herbarium, collected by W. G. Far-
low himself near Chocorua in August 1906 (“‘Kr. 229”). Another collection
was made by Davis at Stow, Mass. (“‘Kr. 280”). Both were painted by
L. C. C. Krieger and are unmistakably the species described above as the
northern form. Then, in 1942, the writer twice collected what seems to be
the same species in Highlands Hammock State Park in Florida. These
southern specimens are sometimes of a pure yellow rather than orange, and
the pileus is perhaps somewhat smaller. Though these differences are slight
and not striking, and the microscopical characters are the same, it seems
wiser to describe them separately.

Pileus “light orange yellow” to “deep chrome,” or more yellow without
an orange tinge: “light cadmium” with the margin “mustard yellow,”

50 FartowlA, Vor. 2, 1945

eventually much more sordid and bleached: “honey color,” margin ‘‘car-
tridge buff,” hygrophanous, but not very strongly so, transparently striate
on the margin, convex with narrowly depressed disk, later infundibuliform,
smooth, glabrous, not viscid, 13-22 mm. in diameter. — Lamellae “pale
orange yellow” or (in the yellow specimens) ‘‘maize yellow,” bleaching to
“cartridge buff” or more pinkish from the spores when old, horizontal,
decurrent, rather narrow, subclose. — Stipe subconcolorous with the pileus
or “lemon chrome” at the apex and gradually paler downward, the base
pallid or whitish, glabrous, smooth, not viscid, subequal, 40-55 x 2-3 mm.
Mycelium white. — Context whitish or somewhat concolorous. Odor in-
significant. — Habitat: On the forest soil in low hammock, especially
where the broad leaved trees are intermixed with Pinus palustris. Septem-
ber. Infrequent.

Farlow did not make an attempt to determine this species, and even the
determination of the genus was very difficult at the time. Thus, one collec-
tion was called “Eccilia or Clitopilus” while the other was marked “Leptonia
spec. vel. Entoloma,” and this in spite of the fact that Farlow was aware of
the specific identity of the collections mentioned. This proves once more
how ridiculous the argument is that Fries’s classification, though not nat-
ural, allegedly has the advantage of being very practical for determination.
Here we have the example that a man of the skill of Farlow put two col-
lections of the same species tentatively in four different genera. In the
modern scheme used by Romagnési (im litt.) and the writer (10), Rkodo-
phyllus Farlowii is easily determined as belonging in the subgenus Leptonia
Fr. sensu Romagnési & Sing. In color, this new species comes close to R.
salmoneus (Peck) Sing. which likewise occurs around Chocorua. Micro-
scopically, however, and also as far as the shape of the pileus is concerned,
the new species belongs in a different taxonomic group. Two colored plates,
executed by Krieger, and illustrating the specimens from Chocorua and
Stow, are preserved at the Farlow Herbarium, and so are the specimens
from these collections. The type collection is the one collected in 1941,
and this is also preserved at the Farlow Herbarium.

This may be a fitting occasion to recommend the genus Rhodophyllus
Quél. as a nomen conservandum to the next International Congress dealing
with the problems of fungus nomenclature. Not only is this name used by
those authors who have proved, by adding valuable new information on the
genus (Kiihner, Lange, Romagnési), that they are competent specialists,
but it also would be historically just to use the name given by the author
who first discovered that the Friesian subgenera and Saccardo’s genera
Entoloma, Clitopilus (excluding the C. prunulus-group), Leptonia, Nolanea,
Eccilia and Claudopus (excluding the C. variabilis-group), are all con-
generic. However, the most important reason for recommending that
Rhodophylius be conserved is that if the rules are to be followed literally
in this case, one of the just mentioned Friesian subgenera must be emended
and all the Rhodophylli would become either Entoloma, or Nolanea, or
Eccilia, etc. In this case, whatever the “legally” chosen genus may be, the

~

SINGER: AGARICALES FROM CHOCORUA 51

number of new combinations necessary to transfer the species listed by
Fries and his followers in one of the remaining genera, plus the species
recently described in Rhodophyllus only, would be much higher than the
number of new combinations still necessary in order to transfer the species
not yet transferred to Rkodophyllus (such as a number of American species
properly belonging in this latter genus in case of its conservation). This
statement may or may not be true if only the number of species and so-
called species in Saccardo’s ‘‘Sylloge Fungorum” are compared, but it is
overwhelmingly true if the number of important, widely distributed and
valid specific names is taken as a base for comparison. If this question is
not taken care of soon, it is to be feared that some McGinty redivivus will
turn out a hundred new combinations a day, making Entoloma rhodopolium
an Eccilia, or Claudopus byssisedus an Entoloma, or Rhodophyllus Farlowit
a Claudopus, depending on which combination of names appeals to him at
the time. The scores of resulting synonyms will have to be carried along
for centuries.

HARVARD UNIVERSITY
CAMBRIDGE, Mass.

LITERATURE CITED

(1) Burlingham, G. S. Two new species of Lactaria. Mycologia 24 (5): 460-463.
1932.

(2) Coker, W. C., and A. H. Beers. The Boletaceae of North Carolina. Chapel
Hill. 1943.

(3) Farlow, W. G. Icones Farlowianae. Cambridge. 1929.

(4) Gilbert, E. J. Les Bolets. Paris. 1931.

(5) Kallenbach, F. Die Pilze Mitteleuropas I. Die RG6hrlinge. Leipzig. 1926-38.

(6) Konrad, P. Notes critiques sur quelques champignons du Jura. Troisiéme série.
Bull. Soc. Myc. Fr. 43 (2): 145-204. 1927.

(7) Neuhoff, W. ix Die Pilze Mitteleuropas II (b) Die Milchlinge. Leipzig 1935-[39].

(8) Persoon, C. H. Mycologia Europaea II. Erlangen. 1825.

(9) Singer, R. Sur les genres Ixocomus, Boletinus, Phylloporus, Gyrodon et Gom-
phidius. Rev. Myc. 3: 36-53, 157-177. Pl. IV. 1938.

(10) —————. ‘Type studies on Agarics. Lloydia 5: 91-135. 1942.

(11) Snell, W. H. Notes on Boletes II and VI. Mycologia 25 (3): 221-232. 1933 and
Mycologia 33 (1): 23-37. 1941. (The latter in collaboration with E. A. Dick).

(12) —————. New proposals relating to the genera of the Boletaceae. Mycologia 34
(4): 403-411. 1942.

2(1): 53-70 FARLOWIA January, 1945

WILLIAM GILSON FARLOW
PROMOTER OF PHYCOLOGICAL RESEARCH IN AMERICA
1844-1919

Wma. RANDOLPH TAYLOR

It is not often that an American botanist has deeply influenced two
diverse fields of his subject, but William Gilson Farlow is recognized as a
dominant figure both in the study of algae and of fungi. His active career
started just as the subject of Botany in this country began to be divided
into specialties, and men began to concentrate their attention somewhat
on a particular field, thus following the example of the Europeans. Farlow
was not without the knowledge of general field botany appropriate to his
day, but he both in teaching and research was a specialist in cryptogamic
botany. He exercised a profound influence in this field. The algae were
his first enthusiasm; he never lost his liking for them and it is the purpose
of this sketch to outline the development of his prestige among phycologists,
and to try to see how he influenced the study in this country. It is not
proposed to attempt to give a general biographical account; that has been
well done by others, particularly Setchell.

Farlow seems to have gone through his college career with some definite-
ness of purpose. Biographical sketches by his students and associates indi-
cate that his scientific taste exhibited itself in the active pursuit of botany
during his undergraduate days, though this was balanced by a marked skill
in music. Probably there was little formal instruction available to him in
his future field, although what the academic customs of the day denied the
infectious enthusiasm of his teacher, Professor Asa Gray, no doubt supplied.
The career of a professional botanist doubtless looked like a rather barren
prospect to his parents; he with his keen mind for business affairs no doubt
had a fair estimate of it. He pursued what we today would consider an
essentially classical training and followed the academic term with two years
at Harvard Medical School, where he did exceedingly well. Thereby he
equipped himself for a remunerative profession; having, thanks to his
family’s comfortable circumstances, no urgent need for exercising it, he
turned to his favorite science of botany and entered Gray’s laboratory as
his assistant.

For an insight into Farlow’s earliest botanical activities we have avail-
able little material. In a day when the keeping of diaries was a common
custom, he seems to have abstained, and the correspondence of his college
days does not seem to have survived. As a result we have nothing before
his adult scientific correspondence to tell us when he became especially
interested in algae, though of later scientific correspondence there is an
abundance.

53

54 FartowlA, Vo. 2, 1945

We do know that when Farlow joined Gray’s Cambridge group he did so
not to follow in his steps as a vascular botanist, but specifically to enter
the practically vacant field of American cryptogamic botanical training.
The study of these plants in this country was the professional concern of
almost no one. A very few scattered persons were doing specialists’ work
on lichens, mosses or hepatics which was excellent, but they did not gen-
erally hold positions in institutions of such prominence and facilities that
their work could be very widely effective. A man in a leading university,
concerning himself with the whole field, and organizing its facilities for
general training in this field, was urgently needed. In many of the floras
of the day a few cryptogams appeared, but in no group had studies in
America begun to give for this country the knowledge equivalent to that
available to Europeans. Gray saw this clearly, and he had begun to assem-
ble materials as occasion favored. It seems that he had a small but notable
collection of algae, the result in the first instance of his friendship with
W. H. Harvey who had held the professorship at Dublin, and J. G. Agardh,
who was professor at Lund. With this as an incentive it is not strange that
Farlow’s first active botanical productivity concerned the algae. While still
in his apprenticeship he studied a small collection of Cuban marine algae
collected by Charles Wright, and published an account of the Chlorophy-
ceae among them in 1871. This, his earliest botanical essay, quaintly illus-
trated with negative line-cuts on a black ground, shows his painstaking
care. He was not yet familiar with the European literature necessary to
make this a critical work, but it does show skill and care. It is representa-
tive of a type of research which will appear frequently during his career,
the report on materials for some exploring expedition.

Meanwhile, his major research work on marine algae was developing.
Farlow had joined a group of biologists who spent the summer of 1871 at
Woods Hole, Massachusetts, laying the foundation for a biological survey
of the area. The United States Commissioner of Fisheries was looking to-
ward the establishment of a research center and hatchery somewhere along
the coast, and Commissioner Spencer F. Baird over several seasons exam-
ined intensively the advantages of several sites which had been suggested
along the New England coast. A small laboratory was set up on the Light-
ship Service wharf on Little Harbor. Just what share the botanical part of
this survey played in the unfolding plans of the Bureau is not clear. In
Farlow’s own development it confirmed his interest in the New England
coast flora. Knowledge of these plants in this country, such as it was, lay
with Professor D. C. Eaton of Yale University. He published very little on
algae, but did have at New Haven a small collection of New England ma-
terial, and some additional material from abroad secured by exchange. The
friendly connection between these two students of Gray was very useful.
Even at first no doubt it greatly fortified Farlow’s beginning algal interests;
later under pressure of his work on ferns, Eaton gave over the algal field
entirely into Farlow’s hands. The first fruits of Farlow’s interest in New

TAYLOR: PHYCOLOGICAL RESEARCH 55

England algae was a short note (1872) on the algae of the Atlantic Coast,
indicating the general character of the flora and some of the striking species.

By now, twenty years after its appearance, the inadequacy of Harvey’s
Nereis Boreali-Americana was evident. Excellently prepared though they
were, and for the times accurate, his three volumes had quite too little field
work behind them. The months of his too brief visit, with considerable
demands on his time apart from collecting of marine algae, were too short
to obtain an idea of the algae of the long American coastlines and so he
relied on the collections submitted by a few amateurs to complete his
catalogue. Local botanists everywhere soon found that Harvey had in-
adequately presented the algal vegetation of their particular coast, and
there began to appear lists aimed at amplifying the record. However,
knowledge of marine algae in America was as yet too ill-advanced to permit
these lists in general to be taxonomically accurate. They did signalize the
general desire for a more complete account of these plants. Farlow, sharing
this desire for more complete studies, brought together his observations in
a list published in 1873. This is the true start toward his major algal work.
Based especially on his 1871 collections at Woods Hole, but including
Olney’s Rhode Island algae collected in 1846-48, of which those of 1846
and 1847 had been checked by Harvey, and also 1870—71 materials from
Greenport and Orient L. I., this paper also contained records which Eaton
contributed of plants from New Haven and Watch Hill, and a Miss Pease
from Edgartown and Vineyard Haven. This gave Farlow a very respectable
list of 103 plants, with eight others considered rare or new to America. It is
a matter of interest that Farlow visited and collected intensively at Woods
Hole localities recognizable and productive today, over seventy years later.
He made this more than a list; though not giving much of descriptive
morphological data regarding the various species, he analyzed the flora in
relation to other shores, recognized its peculiarities and the importance of
Cape Cod as on the line dividing the northern and southern North Atlantic
floras.

The publication of this paper was delayed a year and meanwhile Farlow
had gone abroad on the first of his notable visits to European institutions.
He went with a considerable lot of algae in hand for comparison. His chief
base for the taxonomic study of marine algae in Sweden was at Lund,
where C. A., succeeded by J. G. Agardh had developed resources for the
study of marine algae unapproached elsewhere. Hospitably received there,
he firmly established himself in the good graces of Professor Agardh, who
through his life gave generous help to Farlow on every occasion. These
Agardhian collections are still intact, the specimens numbered in the origi-
nal sequence, the very large specimens such as Ecklonia, Macrocystis and
Durvillaea on great cardboard sheets as Farlow saw and admired them. At
Stockholm he was able to accomplish little on his first visit and went on to
Uppsala. Here he enjoyed meeting both Fries, the elder at seventy-eight
feeble, but assigning Farlow to his son, now the Professor, to be shown the

56 FarLow!IA, VoL. 2, 1945

University. Professor Areschoug, another phycologist, lived nearby, and
Farlow noted the excellence of his extensive microscopic preparations of
algal structures. He also visited the tomb of Linnaeus at the cathedral, a
matter of sentimental duty to all botanists. After about a year with De
Bary at Strasbourg, he went to Switzerland where he collected lichens with
J. Miiller and, in line with his early general botanical training and his
notable aesthetic sense, admired hugely the spread of colors which flower-
ing plants developed on the alpine meadows. The developing facilities for
botanical research at Geneva impressed him. He contrasted the attitude
here with that in Germany where, he writes with scorn, the “Vegetations-
punkt mania . . . affects many of the younger botanists to such an extent
that they are quite unfitted for practical work.” However, he returned to
study under De Bary at Strasbourg and there developed such a respect
for the morphological approach to a fundamental knowledge of algae that
he nearly always insisted that his students develop research from this
angle, urging that they reserve the systematics for a time when, by fuller
experience, they were more thoroughly equipped. He himself here began
a study of Pteris, particularly its apogamy, which resulted in the publica-
tion of three papers. These papers were not really very important ones in
his career, nor did he return afterward to a similar topic for research, but
they signalize his conversion to a particular approach in research for
younger investigators.

Another most interesting algal contact which he made on this trip was
that with J. B. Eduard Bornet in France. Bornet in his collaboration with
Gustav Thuret was in a position of great influence in French botanical
circles; scientific skill and judgment here combined with financial resources
to produce elegantly scientific work of exceptional excellence, and the re-
sult was widely acclaimed. Bornet heartily welcomed Farlow and these
men became close family friends as well as scientific collaborators. At
Bornet’s laboratory he was joined by Rostafinski and Janczewski, Polish
phycologists who worked chiefly on morphology. Farlow developed the
utmost regard for Bornet, and a very extensive correspondence continued
throughout their lives, centering chiefly on comparison of specimens. How-
ever, the profits from this friendship with Bornet were to develop slowly;
the most immediate results from this European trip was to come from his
prolonged residence at Strasbourg.

This visit to biological centers in Europe lasted through two years and
was followed by other trips during which he firmly established and extended
the high regard in which he was held there, resulting, among other advan-
tages, in generous contributions of valuable reference specimens, and ex-
changes with his herbarium. We cannot follow his tour further, but must
consider the later development of his algal interests in America.

He returned from Europe to Harvard, and an appointment which from
1874 was primarily at the Bussey Institution, although he gave instruction
in cryptogamic botany at the Botanical Garden in Cambridge, and in 1879
transferred to Cambridge as Professor of Cryptogamic Botany. However,

TAYLOR: PHYCOLOGICAL RESEARCH a7

the brief contact with applied botany at the Bussey Institution induced
him to attempt some studies and publication on certain disease-producing
fungi, and so began a shift of interest which, though slow to absorb all his
research attention, eventually did so.

During this time Farlow worked on the project of a catalogue of Ameri-
can marine algae foreshadowed by his first brief papers. His collections,
thanks to his correspondence with Dr. C. L. Anderson and others on the
west coast, as well as with all phycologists on the east coast, and his friends
abroad, had grown to the extent that he felt accurately informed about
these plants. Many specimens had been verified during his conference with
Agardh at Lund, and others were sent to him from time to time. In 1873
he published his list of the algae of the south coast of New England, and in
1875 after his return from Europe he expanded this to a list of United
States marine algae, adding especially west coast and Florida records, to-
talling 430 items. This was a list with a short general introduction and a
few notes, a less elaborate form of the same appearing the next year under
the auspices of the U. S. Fish Commissioner with the addition of notes
upon economically useful seaweeds.

A small accumulation, especially of California and Florida algae, was
published with a number of new species in 1877, but he never developed
this general project further. Instead, he took New England records out of
the 1875 list, provided a very useful general introduction, descriptions to
the major categories and the species, and in 1881 published his Marine
Algae of New England and the Adjacent Coast, the work which closed his
major algal productivity. This paper well deserved its long career of use-
fulness; clearly presented and complete for its time, it was the uncontested
reference work for its area for over fifty years. His fame as a phycologist
chiefly rests on this work.

So much for the general main line of Farlow’s algal evolution. What
other lines of productivity did he develop? It would seem that we can
group them into three classes. First in importance was the exsiccata which
he founded and issued with the aid of D. C. Eaton of Yale and C. L. Ander-
son of California. Second was a group of papers of considerable importance
locally, dealing with algal contaminants of Boston’s water supplies. Third,
a group of minor algal taxonomic papers. Almost all his remaining work
falls into one of these three classes.

First we may deal with the Algae Americanae Boreali Exsiccatae. This
appears in his correspondence but little before it was actually produced.
Probably he talked it over with Eaton on some visit, so that little was left
for written arrangement. However, Eaton (March 5, 1876) refers to a dis-
cussion of label size, perhaps for the exsiccata, and in succeeding letters
references to specimens in multiple probably have the same significance,
while Eaton (September 8, 1877) refers to “. . . enough for the 30 sets
with a few I had already,” and a week later refers to “‘. . . specimens too
large for the small fasciculus,” after which this work entered considerably

58 Fartowia, Vor. 2, 1945

into the interchange. The Eaton correspondence was voluminous, that with
Anderson less so. Anderson writes (Sept. 16, 1876) “I like your sugges-
tion in regard to publishing an authentic set of our algae. I shall have con-
siderable time this winter that may be devoted that way. I do not know
that I have enough of our species to make a set of fifty but . . .” This
seems to open up the business of western participation in the series. The
fifth and final issue appeared in July, 1889. The importance of the series
lies in the good quality of the specimens, the accuracy of the determina-
tions, and particularly the fact that this first exsiccata of American algae
now introduced these plants into many important collections at home and
abroad, where before probably only the Harveyan and Agardhian collec-
tions had any large representation. It did very much to increase Farlow’s
prestige abroad. He was largely instrumental in starting the project and
in the end it was practically under his full control.

The series of five papers 1876-80 dealing with algae in the reservoirs of
the Boston water supply and related topics probably called for little critical
research, but they were very important in discharge of Farlow’s civic re-
sponsibilities. These papers at first dealt with a simple problem of unpleas-
ant taste. Farlow found nothing clear-cut in the algal flora that could be
specifically responsible. He indicated a few of the most characteristic
freshwater algae present, and was able to assure the public that these algae
were in no way dangerous to health. Apparently in at least one ‘pond’ the
trouble was due to a Spongilla. Finally in the 1880 paper he wrote a semi-
popular treatise on the general features of the algal floras of reservoirs.

Lastly one turns to what may best be described as Farlow’s shorter algal
works. When one inspects the list it is clear that relatively little bears upon
his own collections. Farlow travelled and collected extensively on the west
coast, in Florida and in Bermuda. The material went into his collections
and presumably was used for exchange, but he published few of the new
records which it contained. As papers covering material which he had col-
lected himself, or which at least came from an area and a flora familiar to
him, we may recognize his Notes on New England Algae in 1882, in 1889
On Some New and Imperfectly Known Algae of the United States I, and
in 1899, Three Undescribed Californian Algae.

The prominent feature of these shorter papers is based upon his com-
manding position in American botany. While the era of great unspecialized
natural history exploration was passing, nevertheless it had not disap-
peared. Parties were still going off and bringing back heterogeneous, often
small, often inexpertly collected lots of material to be parceled out to vari-
ous specialists for study. As always the algae were a minor part, assembled
very clumsily, but they represented a cherished bit of the treasure of the
often arduous journey, and they could only be entrusted to Dr. Farlow.
So, because he greatly liked these plants and felt it his duty to care for
these hard-won little collections, he studied them, occasionally reported
new species, and published an annotated list. The geographical range cov-
ered is very great, though sometimes there were as few as a half-dozen

TAYLOR: PHYCOLOGICAL RESEARCH 59

species included. His Cuban Seaweeds of 1871 was followed by reports on
algae from Kerguelen Island in 1876, Cumberland Sound in 1879, Texas
and northern Mexico in 1883, Alaska in 1885, Ungava Bay in 1886, South
America in 1888, Peary Arctic Expedition in 1895, Azores in 1897 and the
Galapagos Islands in 1902.

Farlow’s last strictly algal paper appeared in 1916, a very scholarly dis-
cussion of the general character and distribution of Pacific Ocean algae,
showing that his interest had persisted through the years. His last recorded
paper was directly connected with algae too, as it was a biographical sketch
of his friend J. B. E. Bornet, published in the same year.

One of the chief circumstances of Farlow’s career and one of great poten-
tial importance for the development of his phycological interests was his
share in the development of the Marine Biological Laboratory at Woods
Hole in Massachusetts. The prospectus for this organization was dated
6 April 1887, and Farlow was the Trustee first named in this document.
He desired to resign the following year, but letters from Minot and others
urged him not to do so, and he served for some years before he severed his
connection. During this period of development under the directorship of
Professor C. O. Whitman of the University of Chicago, the Trustees were
called upon for much financial help. Eventually some came to feel that
they were losing control of the finances of the institution, that its expendi-
tures were exceeding what the Trustees should be called upon to bear, and
that the limitations which they set upon the activities of the Director were
disregarded. Director Whitman clearly had a larger vision than these
Trustees, a vision less trammeled by concern over details of economical
operation and balanced budget. A group of the Trustees resident near
Boston were particularly resentful of the attitude of Whitman and the
business management of the Laboratory. Having organized and financed
the institution, they felt that they should direct its administration. On the
other hand, in the course of ten years the biologists who formed the Cor-
poration, who actually came to work at the Laboratory, now felt that the
institution was peculiarly their own, that they could shape, expand, and
staff it as they chose, and that the Trustees who were commonly not per-
sons working at the Laboratory during the season, should defer to the
general desire of the active group.

The meetings of the Corporation and Trustees had been held in Boston
in the winter and most of the Corporation members could not attend; a
strong desire for greater participation of the biologists from all over the
country led to a demand that meetings be held at Woods Hole during the
summer. This was done in 1894, when a meeting of the Trustees was held
there, but the Annual Meeting at which the major business was transacted
was still held in Boston. The actively dissatisfied group of Trustees, some
seven or eight out of twenty-one, engaged in an exchange of bitter letters
among themselves, looking toward more effective control of the Laboratory.
A special Corporation meeting was called at Boston, 16 Aug. 1897, to pass

60 FartowlA, VoL. 2, 1945

by-laws changing the Annual Meeting to Woods Hole, and the date from
January to August. This meeting was presided over by Farlow as Acting
President; he tried to show that the meeting was illegally called, but failing
of support, withdrew. The meeting proceeded under C. G. Kidder to make
important changes in the by-laws, improving greatly the scheme of organi-
zation. However, seven of the Boston group of Trustees afterward resigned.
Six of them, and one other dissatisfied Trustee, attacked the administration
of the Laboratory in a very severe article in the journal Science, and to this
Farlow was a party. The correspondence in Farlow’s file shows the stages
in the development of this Science article, the anger of the participants, and
Farlow’s share in keeping the text within reasonable bounds. The support-
ers of the Whitman administration replied in a privately printed pamphlet,
subsequent to which, with the withdrawal of the chief dissidents, the whole
matter quieted down.

Thus ended Farlow’s direct part in what has evolved into the most am-
bitious project of codperative biological organization ever attempted. It
remains for us to enquire into his share in the work of the Laboratory dur-
ing his trusteeship, and the persistence of his influence after he resigned.
It does not appear that Farlow was ever active in instruction at the Labora-
tory, or that he was in any large way an investigator there. The first year
there was no staff botanist; in 1889 James E. Humphrey was appointed, to
be followed the next year by W. A. Setchell. Humphrey had been intro-
duced to the algae by the Rev. J. D. King at the Martha’s Vineyard Sum-
mer Institute; he studied the development of the perforations in Agarum
for his B.S. degree thesis at Harvard, though he turned to the Saproleg-
niaceae for his doctorate in 1892, and eventually he deserted mycology for
cytology. Setchell, another student under Farlow, was obviously an ap-
pointment favored by him. Setchell returned in 1892 and subsequently
through 1895, when he departed for the west coast. Obviously Farlow, al-
ready retiring from teaching at Harvard, was content to transmit his algal
enthusiasms through his students. In 1896 a greatly enlarged scheme of
botanical instruction was initiated under the leadership of Professor J. M.
Macfarlane of the University of Pennsylvania, who had been trained at
Edinburgh, but here again Humphrey and one of Farlow’s students, George
T. Moore, guided the work on cryptogamic botany. This brings one to the
year of the great upheaval, but in 1897 B. M. Davis, another student from
Farlow’s laboratory, who had gone to the University of Chicago, took con-
trol and with G. T. Moore again gave the cryptogamic work. The greatly
expanded list of botanical instruction soon was restricted, but Davis re-
mained in charge, and Moore was associated with him until 1905. During
this time various other additional instructors appeared, for algae in par-
ticular J. J. Wolfe from 1902-1906, again trained by Farlow. Moore took
charge of the cryptogamic botany in 1906 and carried it through 1918;
associated with him until 1914 was G. R. Lyman —a Farlow student —
and others, including some Harvard-trained men, but probably none who
were directly trained by Farlow.

TAYLOR: PHYCOLOGICAL RESEARCH 61

Here, then, ends the immediate influence of Farlow and his students in
this venture, which has grown from a handful of investigators in 1888 with
most trifling equipment, to a group averaging over 360 for the five years
before the present war, and about 125 students, with extensive equipment,
large permanent buildings and a considerable maintenance staff. By 1900
it had become clear that the Laboratory was not to collapse; in spite of the
adverse judgment of Farlow and his friends for which there was some jus-
tification, the administration rallied scientific and financial support, and
went its way. Farlow never took further active share in it; neither, how-
ever, did he dissuade his students from doing so and under their guidance
its botanical work prospered for thirty years. While it is extremely doubt-
ful that Farlow actually was diverted from algal study to any great degree
by the Woods Hole dispute, it is clear that for other reasons he was willing
to let others do the research on these plants. By now he had two promising
protégés, his student Setchell and an amateur, F. S. Collins, who was an
accountant and business man of Boston and Malden. For the rest of our
analysis of Farlow’s influence we may go to his list of academic students
and to his friends, their records and their correspondence with him.

A formal list of Farlow’s students does not seem to have been compiled.
From the congratulatory volume of photographs and letters sent to him on
the occasion of his seventieth anniversary we may, however, glean the
names of most of those of his students who specialized in algae at least
for a time.

The first man, apparently, to take a doctorate degree under Farlow was
B. D. Halsted, who in 1878 submitted his successful thesis on American
charophytes. This was the nearest to a taxonomic dissertation that Farlow
ever permitted a student to submit. Halsted became Professor of Botany
at Rutgers College where his interests changed and he mainly concerned
himself with plant diseases and the breeding of crop plants.

However, perhaps an earlier student contact was that with F. W. Hooper,
who took a bachelor’s degree at Harvard in 1875, but his master’s degree
only in 1898, and did not proceed to the doctorate. There is no record of
significant phycological publication on his part, but he contributed Florida
algae to the Algae Exsiccatae Americae Borealis. His interests shifted to
other fields and he became Director of the Brooklyn Institute.

The next student was a very notable one, and for Farlow his appearance
was a fortunate circumstance, because into his hands Farlow consigned
with confidence the teaching of cryptogamic botany when his affairs in-
duced him to drop it in 1897. This was Roland Thaxter, who took his
bachelor’s degree in 1882 and his advanced degrees under Farlow six years
later. Thaxter’s brief excursion into the algal field included especially his
paper on Compsopogon, a morphological study. He then turned his atten-
tion to the fungi and became a very notable figure in that field.

At this period there came under Farlow’s influence Kingo Miyabe, an
able Japanese botanist who had had his early training at Sapporo. He took

62 FarLow1A, Vou. 2, 1945

his doctorate at Harvard in 1889 with his phanerogamic thesis on The Flora
of the Kurile Islands. In later years he published extensively on the marine
algae, and especially the Laminariaceae, of Japan.

While Thaxter was a student, three others were pursuing algal problems
with Farlow. J. E. Humphrey took a bachelor’s degree in 1885 and went
out to teach at Indiana University; he returned to take his doctorate in
1892 with his thesis upon the Saprolegniaceae. He introduced the study of
cryptogamic botany and especially the algae at the Marine Biological
Laboratory, but his interests shifted to cytology in later years. While hold-
ing a post at Johns Hopkins University and in charge of a biological party
in Jamaica, he contracted yellow fever and died in 1897.

R. P. Bigelow, likewise taking a bachelor’s degree at Harvard in 1887,
studied and published upon the structure of the adult growing point of
Champia. He shifted his interest to zodlogy and held a professorship at the
Massachusetts Institute of Technology.

In 1887 Farlow’s friend, Professor Eaton of Yale, sent a very promising
new graduate to study with him. This was W. A. Setchell who, like Miyabe,
continued actively in algal research to the end of his long career. These
were the only students of Farlow who did so. Setchell took a master’s
degree in 1888 and a doctorate in 1890, going eventually to a professorship
at the University of California. His publications on the marine algae of
the west coast are extensive and fundamental, but he had many and varied
auxiliary interests which also led to publication.

G. J. Peirce took a bachelor’s degree at Harvard in 1890 and thus came
under Farlow’s influence, though he went to Leipsic for his doctorate.
Peirce was essentially a physiologist, and eventually Professor of Botany at
Stanford University. He published on the relation of Nostoc to Anthoceros,
gamete extrusion in Fucus, irritability in algae and on brine organisms.

The next year H. M. Richards likewise graduated from Harvard; how-
ever he continued and completed the doctorate in 1895. His early algal
work dealt among other matters with parasitic algae; later his interest
shifted to the fungi, wound reactions and the like. He held a professorship
at Barnard College.

From this time on, all of Farlow’s students held a bachelor’s degree from
another institution, although some added the Harvard one to it. B. M.
Davis came to Harvard from Stanford University. Taking the master’s
and doctor’s degrees in 1894 and 1895, Davis went on to a detailed study
of algal distribution in the Woods Hole area for the U. S. Fish Commission,
which was published in 1913. Although Davis’s interest in the morpho-
logical side of algal studies continued, his actual research interest changed
to plant genetics. He held a professorship at the University of Michi-
gan.

G. T. Moore came to Harvard in 1895 and repeated the bachelor’s de-
gree, taking the master’s the next year and completing the doctorate in
1900. His early interests were particularly in the freshwater algae; later
the burden of administration kept him from activity in the field. He be-

TAYLOR: PHYCOLOGICAL RESEARCH 63

came Professor of Botany at Washington University and Director of the
Missouri Botanical Garden.

In 1897 E. W. Olive took a master’s degree, and in 1902 a doctorate at
Harvard. His early interest was in Myxophyceae, particularly their diffi-
cult cytology. Later his interests shifted to mycology and he served as
curator at the Brooklyn Botanic Garden before leaving academic fields.

J. J. Wolfe came to Harvard, to take his doctorate in 1904, and made a
cytological study of Nemalion which was well considered at the time. He
did not continue with algal research. He held a professorship at Trinity
College, Durham, N. C.

The next student at Harvard with claims to specialization in algae was
R. F. Griggs. He took his doctorate in 1911 and studied the various fea-
tures of the west-coast brown algae before becoming interested more in
general matters of plant distribution. He holds a professorship at George
Washington University.

Finally we may mention F. K. Butters, who took a second bachelor’s
degree at Harvard in 1900, returning to complete his doctorate in 1917.
He studied especially the genus Liagora and was interested in Hawaiian
algae, but later concentrated his attention on ferns and phytogeography,
holding a professorship at the University of Minnesota.

Reviewing this list we see that only two men continued consistently in
algal research. Four, namely B. M. Davis, G. T. Moore, H. M. Richards
and R. F. Griggs continued profitably with algal work for a time, and then
went on to other fields. In teaching no doubt these all transmitted some-
thing of their early enthusiasm; to the teaching at Woods Hole, largely
algal, Humphrey, Setchell, Davis, Wolfe, Moore and their academic de-
scendents, all contributed.

It is hard to judge the worth to science of these men. Some were active
researchers, like Thaxter and Setchell, others administrators like Moore.
Of the fifteen names, eleven carried the star with their listing in American
Men of Science, indicating a considerable recognition by their colleagues;
Humphrey died before the custom was initiated, and Miyabe was a for-
eigner and so not included. In the voting for the first edition, Farlow him-
self ranked first among botanists, Whitman second among the zodlogists,
exceeded only by W. K. Brooks. Seven of Farlow’s students were starred
in this first edition, one being in zodlogy. Almost all of his phycological
students came to hold full professorships in large and well-equipped uni-
versities, many being the departmental heads. That the actual output of
algal research is not great for such a number over so long a period is prob-
ably related to the specialized character of the field and the few openings
favorable to specialists in marine algae. The thread of inspiration was not
broken, however, but carried on into the next generation through Setchell.

The topic of Farlow’s relations with.other botanists is too huge to de-
velop fully in the space remaining for this article. One may well doubt if
any botanist of his generation or since has conducted as extensive a per-

64 Fartowia, Vou. 2, 1945

sonal correspondence. Farlow knew every phycologist of note, and kept
the acquaintances active. The lesser men at home and abroad constantly
referred material to him for advice; he in turn sent material overseas to
persons in a position to give expert help, and thereby kept up the acquaint-
ances established during his trips abroad. Farlow never cared for either an
amanuensis or a typist; he wrote his letters, one may judge from some
remarks, concise ones, with his own hand and seldom kept copies. The
letters reaching him he kept, thousands of them, and in his later years he
had them sorted and bound into scores of volumes. A person who had
grown up with knowledge of Farlow since college days would be in a much
better position than the present writer to extract a connected story of Far-
low’s life and friends from these letters. Only half of the story is there, the
other half must be surmised, for in only such rare cases as those of Gray,
Bornet and Collins has the other side of the exchange been brought into
the Farlow library. The earliest correspondence certainly is missing, and
there is no way of telling how much has been eliminated before what Far-
low thought was important was bound together.

In the first place we must recognize that the bulk of the scientific matter
in these letters consists of lists of identifications and notes on various
plants, discussions of nomenclature and the like. From this, little suitable
to the present presentation can be extracted. Of the general material only
a few glimpses can be offered, to give an idea of the nature of the problems
which were referred to Farlow for advice.

The first group of correspondents are those dating from Farlow’s Bussey
Institute period and his first European trip. Eaton, Anderson and Wolle
represent the main American contacts of these days. Professor D. C. Eaton
of Yale University, also interested in the marine algae, was for many years
his best and most. active correspondent. The exchange opens with a letter
from Eaton describing the results of dredging under Baird’s direction near
Eastport, Maine. He writes that the dredge had brought up Delesseria
sinuosa and Ptilota serrata from at least 400 feet depth. Two years later
Eaton seems to have visited Farlow at Woods Hole for a collecting trip.
The trip must have been strenuous, or at least ended in a rush, for Eaton
accounts for the whereabouts of his own and borrowed attire after what
one suspects to have been a classical attack of professorial absent-minded-
ness: “‘Dear Dr. I remembered the coat before I left Woods Hole,
but too late to reclaim it. Your night shirt I have here, & will presently send
it to your Boston address by the Express Co. . . . I suppose you are hav-
ing a real good time of it now your students have gone.” (20 August, 1875).
The next year begins a very long correspondence about the exsiccata for
which Eaton and Anderson shared the responsibility with Farlow, which
includes their own collecting and that of Edward Palmer and F. W. Hooper
in Florida. In 1879 Eaton was proposing Farlow for a position at Johns
Hopkins University, as he mentions having written to this effect to Mr.
Gilman. Nothing came of it. In earlier letters it is Eaton’s preoccupation
with ferns which calls for apology, but from 1881 it becomes evident that

TAYLOR: PHYCOLOGICAL RESEARCH 65

Farlow’s interest was shifting to the fungi. Setchell’s name is mentioned
first in 1885 at a time when it is evident that these correspondents were.
building up their bryophyte collections; two years later a discussion of
Setchell’s plans appears, and in June Eaton refers with pleasure to his
appointment at Harvard. Apparently Eaton did not think much of or-
ganized scientific groups: in 1893 he writes to Farlow expressing disap-
proval of a botanical congress: “. . . I have the heartiest possible con-
tempt for the whole crowd of priority-worshippers . . . and shall not feel
the slightest obligation to conform to their ‘Laws’,” and regarding an em-
bryo organization in this country “. . . the longer I think of the botanical
society the worse I think.” There seems to have been close social contact
between the families, and Eaton’s daughter kept Farlow closely informed
of her father’s illness and death in 1895.

Dr. C. L. Anderson was the other partner in the Exsiccata enterprise.
He writes to Farlow in 1876 expressing enthusiasm for the prospective
series, and continues with letters regarding the specimens and labels for it.

Francis Wolle, clergyman head of a Moravian girls’ seminary at Bethle-
hem, Penna., was an indefatigable student of freshwater algae, on which he
published largely, his books on freshwater algae, desmids, and diatoms
having a considerable vogue fostered by the numerous, if not very precise,
illustrations and descriptions. The correspondence with Farlow lasted for
about ten years from 1875, mainly with regard to specimens the identity
of which was in question. Apparently Farlow distrusted Wolle’s determina-
tions and for this reason sent some of his specimens to Bornet for redeter-
mination. From a last letter of 1884 we gather that the edition of the
Desmids of the United States was limited to 460 copies, a venture with the
initial success of which Wolle was quite pleased.

The foreign correspondence beginning in this period centers on Farlow’s
first European trip, and continues until when toward the turn of the cen-
tury, these older botanists had passed from the scene. The earliest Agardh-
ian exchange opens when J. G. Agardh (8 January, 1872) welcomes Farlow
as a new correspondent, and indicates his desire for west-coast algae, espe-
cially Laminariaceae. Later that year, correspondence arranging for Far-
low to visit Lund appears and the facilities there are explained. Twenty
years later Agardh again writes to Farlow at Paris inviting him to revisit
Lund, and in August expresses his pleasure in the occasion after Farlow
had left. There was much reference to the notable “Algae Muellerianae”
from Australia, of which he was sending a series to Farlow. Farlow’s mar-
riage in 1900 was greeted with the friendliest expressions and a congratu-
latory poem.

In 1873-74 we have letters from Farlow to Thuret and Bornet arranging
for his visit to Antibes. Farlow was having the traveller’s perennial diffi-
culty of fitting his plans into the vacation plans of the Europeans. The
correspondence with Thuret was necessarily brief, but that with Bornet
was very extensive for many years. In 1874 he wrote to Farlow encourag-
ing him to prepare an introduction to the algae of his region, 7.e., New

66 FartowI!A, Vor. 2, 1945

England, and telling him the gossip regarding the Antibes group of bot-
anists which Farlow had lately left. Of course the correspondence dealt
largely with determinations of Myxophyceae. They exchanged their major
as well as minor works: Bornet in 1878 writes of sending the Etudes
Phycologiques and in 1882 of receiving four copies of Farlow’s New Eng-
land Algae and of Hervey’s Sea Mosses. In 1883 writing in reply to an ac-
count received of Farlow’s trip to Minneapolis, he indicates the part which
Longfellow’s Hiawatha and Cooper’s tales bore in forming his ideas of that
part of America!

Correspondence with Areschoug, Hauck and very many others of lesser
note exists, but lends little to the interest of this account. That with Sir
Joseph Hooker is more significant; Farlow could act as intermediary and
tell him of the waning health of his friend Professor Gray. Fairly early, in
1878, replying to a request for portraits of botanists, Hooker tells what he
can supply and which exsiccatae he has in duplicate for exchange. Finally,
in 1907 he acknowledges formal congratulations, received on attaining his
ninetieth birthday.

With the next age group the importance of the letters shifts to the
American field. Farlow is by now a recognized leader, and while his con-
tact with older Europeans flourishes, he is not particularly impelled to
active correspondence with the younger group. However, we find some new
names, as contact was developing, for instance, with Hariot and Lagerheim.

Among Americans, we have now the period of development of Collins as
a phycologist, Farlow’s encouragement of the Curtiss family in botanical
collecting, and the establishment of his earlier students. Frank S. Collins
was a business man employed by the Boston Rubber Company in charge
of their accounting. He had little leisure, but he employed it in ardent
botanizing. He was quite as much interested in freshwater algae as in
marine plants; from contributing to Dame and Collins’ Middlesex Flora
and the algae of Rand and Redfield’s Flora of Mount Desert, through
many smaller papers, he developed his Green Algae of North America. On
his Green Algae ... , the Algae of Bermuda which he prepared with
Hervey’s cooperation, and on his spectacularly successful Phycotheca
Boreali-Americana, largest exsiccata of algae ever issued, his fame chiefly
rests. His work was for its day very accurate and scholarly, particularly
the smaller papers in which he described the novelties he later incorporated
in his larger works. For much of this Farlow provided the critical back-
ground, the reference collections and encouragement, although ultimately
Collins assembled an excellent herbarium and library of his own. As it was
possible to confer personally, the amount of correspondence was all the
more remarkable. The letters begin in April 1881, but refer to earlier
matters, discussing trips to Nahant and Woods Hole, and the plants col-
lected. In Oct. 1894 appears a sentence, ‘“. . . I want to distribute this
species, either in the Phykotheka Universalis or in the set which Setchell,
Holden and I have been talking of issuing, if we ever get to it; and I don’t
want to give a new name only to have to take it back again.” There is

TAYLOR: PHYCOLOGICAL RESEARCH 67

much interchange at about this time regarding the C. E. Pease and E.
Butler collections of Jamaican algae on which Collins was to publish. The
queries about labels for the Phycotheca, to which Farlow contributed
specimens, begin to appear and Farlow writes to Collins in February 1895
acknowledging the receipt of Fascicle I, and in April Collins writes “. . . I
have sent out about all the first lot I made up of Fascicles I and II, and am
laying out the specimens for another batch of the same.” From letters to
Setchell in 1908 it seems that Farlow was trying to secure money to bring
Collins to the herbarium at Harvard, but that enough was not forthcoming.
As early as Oct. 1901 Collins mentions that five or six years earlier he had
done a good deal of work toward a book on New England marine algae,
and that he now contemplated making it cover the whole east coast; this
manuscript was never modernized and completed, but went on Collins’
death with his collections to the New York Botanical Garden. This inter-
change, too long to follow through, only ceased with the death of Farlow
in 1919, Collins dying the next year.

Another useful interchange, likewise outside the academic sphere, took
place about this time between Farlow and the Curtiss family, Mrs. Floretta
A. Curtiss and her son, A. H. Curtiss. Mrs. Curtiss, living in Florida, issued
three series of Marine Algae of Florida starting in 1895, with many of the
determinations by J. G. Agardh. Much earlier correspondence had been
exchanged between Mrs. Curtiss and Farlow regarding various Florida
algae, starting in 1877. Apparently Mr. Curtiss travelled much for govern-
ment agencies; he also seems to have collected natural history specimens
of all sorts for sale. Mrs. Curtiss especially cared for the algae, which her
son rough-dried for her in the field, and which she soaked out and mounted.
In 1899 he writes of his mother’s partial paralysis and her desire to have her
personal collection of algae, which contained much valuable foreign mate-
rial received by exchange, gotten together for the U. S. National Museum.
Three months later her death is referred to, and the plans for assembling
the Algae Curtissianae into volumes for preservation. As late as 1904 let-
ters were exchanged, but seemingly no more algae were to be expected.

Letters between Farlow and his student J. E. Humphrey began earlier,
but extend into this period. What we have from 1888 tells of Humphrey’s
arrival at Indiana University, the lack of a botanical library, his plans to
build up the department and his hope for separate departmental status.
In 1897 the letters came from Johns Hopkins University, his last post. A
curious exchange here illustrates the lack of centralized authority at the
Marine Biological Laboratory at the time. Humphrey had been teaching
the algae course there; he was preparing for his last Jamaica trip and was
asked to secure a substitute. Humphrey arranged with H. M. Richards to
come, but Director Whitman without consulting Humphrey or Farlow,
engaged B. M. Davis and printed the announcement. Farlow and Hum-
phrey were indignant at having the matter taken out of their hands with-
out warning; Humphrey died on this Jamaica trip, but both Davis and
Moore served on the Woods Hole staff repeatedly. This was no doubt one

68 FarLowiA, VoL. 2, 1945

of the items which collectively induced Farlow to sever his connection with
the laboratory.

Except for the Collins correspondence, that with Setchell is the most
important domestic correspondence relating to American algae. The first
letter is one dated December 1885. Through Eaton, Setchell had sent a
specimen to Farlow for identification which proved of interest, and Setchell
was forwarding more of it. Then follow a series of letters dealing with
Lemanea, Batrachospermum and Tuomeya. There intervenes next appar-
ently a period when Setchell was ill with rheumatism and spent some
months at Sharon Springs, N. Y., and did some field botanizing about
there, but with land plants rather than algae. The work at Woods Hole
comes up for conference in 1891, and various persons as Davis, Brannon
and Rothrock are mentioned as students who later became well known in
the botanical world. A perennial topic of discussion still, the Marine Bio-
logical Laboratory ‘‘mess” is at that time reported as somewhat better
than it had previously been, but Setchell complains of the high room rent:
$1.00 a week each, two in a room in the Gardiner cottage! Each summer
there is more about Woods Hole, and in July 1894 he mentions settling
into the then new Botany building. Next June from Yale he writes to ask
Farlow to recommend him for appointment to a professorship at the Uni-
versity of California at Berkeley; this after testing the disposition of Yale
to meet the competition and getting no encouragement, he reiterates, and
tells that he will receive $3000 as department head. In December he
writes from Berkeley of the pleasure at the welcome he received, says he
has about 500 volumes in the botanical library and tells what other funds
are available to him. M. A. Howe and W. L. Jepson were there as instruc-
tors; but Setchell felt that since Howe’s interests were similar to his own,
that Howe should go elsewhere so that the activities of the department
might be diversified; he recommended him highly to Farlow and asked
help in placing him. The early issues of the Phycotheca came in for discus-
sion and Setchell writes appreciatively of Farlow’s review in the American
Journal. Later in 1896, when the large expansion of the botanical lecture

list at Woods Hole occurred, Setchell wrote with some amusement “. . . I
understand that Macfarlane wants a botanical garden at Woods Hole
&c — &c. . . .” Apparently Setchell doubted the wisdom of such am-

bitious plans. Setchell wrote to Farlow in some detail of his trips abroad,
as to New Zealand and Europe in 1905 and again in 1911, and of the algae
he collected and those he saw at the British Museum, Kew and the
Linnean Society’s rooms.

M. A. Howe opened correspondence from Berkeley with a question on
Fucus evanescens, and asks help on the identification of Pacific Grove
algae. He mentions his paper on California hepatics as his doctorate thesis
and as a collateral interest. In 1894 he wrote inquiring of any opening at
Harvard and in 1898 there were some moves toward bringing Howe to the
Harvard herbarium in charge of bryophyte work. The financial terms
were set forth at some length, Howe being very explicit regarding what he

Taytor: PHYCOLOGICAL RESEARCH 69

felt were the conditions under which he could come. Nothing resulted,
however, and other applications to Missouri and Oberlin which he men-
tions likewise having petered out, he wrote hopefully to Farlow regarding
obtaining the headship of the department at Berkeley, which went, how-
ever, to Setchell. The latter encouraging him to move, in 1897 he writes of
having taken an assistantship at Columbia University. After a break, the
correspondence begins actively again from the New York Botanical Gar-
den, and a great many inquiries regarding various plants occur, as finally
those during 1919 regarding the Bahama Flora. Howe was eventually the
Director of the Garden, and his work on marine algae, while less extensive
than Setchell’s, was at least equal in quality.

It is not possible to follow the correspondence with European botanists
through this period, but one must not fail to notice the friendly interchange
with Alexander Gepp as it covered the period of his marriage to Ethel S.
Barton, another English phycologist, her later very precarious condition
due to tuberculosis, and death. An amusing reference to the compilation
of the British Museum of Natural History catalogue occurs, which Gepp
describes as having been made up by J. Britten, himself, Wiltshear the
attendant, with two words contributed by George Murray. In Gepp’s own
words (5 July, 1905) “We were the scribes, & George Murray was the
Pharisee who contributed but 2 words — viz his own name.”

It is impossible for a person who had no personal acquaintanceship with
Farlow to write as understandingly of his witty, friendly personality as his
own students. One could hardly improve on the charming biographical
account by Setchell 1 to which all should refer for details of his life and the
complete list of his publications.

Without question Farlow dominated algal study in America for a con-
siderable period, and represented it before European scientists. Also un-
questionably this high regard was justified; the problem is to appreciate
truly how this came about. It was not due to the quantity of publication
which, respecting algae, was never great and for the later years very small.
Nor was it due to the importance of the, material studied, for with the
exception of the New England report it was never comprehensive, nor in
general were many new species described. It did not derive from the sump-
tuousness of publication, for his best work appeared in government reports,
nor the elegance of the illustrations, which when from his pen were few,
stiff and labored rather than skillful. So far as his publications go they,
by the care in preparation, clarity and lack of error, confirm rather than
accent his reputation.

One must recognize that the high esteem in which Farlow was held was
due to other factors. In the first place he was skilled in gathering together
collections. He developed a personal herbarium and library of rare excel-
lence, including items scarce and costly even at that day. His collections

*Setchell, W. A. 1927. William Gilson Farlow 1844-1919. Memoirs Nat. Acad. Sci.
21 (4): 1-22. Portrait and complete bibliography.

70 FartowlA, Vor. 2, 1945

were in only secondary degree the result of his own field work. Again ex-
cepting New England, where at a few places he did get a variety of mate-
rial, his collecting was not extensive and did not contribute largely to his
publications. Rather by purchase and exchange did he secure from others
splendid representation of the flora of many parts of the world. These
treasures he was quite willing for others to consult, under strict super-
vision. As a result the more active descriptive phycologists, such as Collins
and Setchell, came constantly to him for consultation, appreciated his
careful assistance, and spread his fame. In the same way his European
travels and active correspondence served abroad.

Here then we have probably the clue to Farlow’s most significant con-
tribution. He developed the first good reference collections for algal study
in this country. He published skilfully, if not abundantly. He encouraged
several young investigators to attempt algal problems, gave them rigorous
training in meticulous research, and the two or three most productive
among these he assisted in every way, so that they in turn dominated algal
studies in America after Farlow’s own interests were largely forced to the
fungi. Professor Farlow was, then, clearly more important as a promoter,
developing resources and trained men, than he was as an investigator him-
self, and as such he fills a distinguished place in American botanical history.

For the hospitality of the Farlow Herbarium and Library, for access to
Professor Farlow’s extensive correspondence, and for patient help in using
these treasures, the writer is greatly indebted to the kindness of the curator,
Dr. D. H. Linder.

UnIvErsITY oF MICHIGAN
Anw Arzpor, MIcHIGAn

2(1): 71-82 FARLOWIA January, 1945

FARLOW’S INTEREST IN AN INTERNATIONAL
ABSTRACTING JOURNAL

FRANS VERDOORN !

In collecting data about the history of international relations and codper-
ation in the plant sciences, I have gathered a large number of letters written
by the late W. G. Farlow. Dr. Farlow, though a charter member of the
Association Internationale des Botanistes, took no great part in its activi-
ties. He has never been a member of the council or of one of its commis-
sions. Yet, during his fruitful life he was in touch with more men abroad
than most of his contemporaries and was particularly active in undertaking
and developing exchange relations. He was greatly interested in various
aspects of abstracting and listing the new botanical literature of the world.
In this field he exercised probably more influence than is realized. The need
of establishing a truly international botanical abstracting journal was felt
and discussed by him at an early date.2, Dr. David Linder has suggested
that it would be useful to publish in this special number of Farlowia a few
of Farlow’s letters on this subject. They are not only of interest for the
student of international relations but throw an interesting light on Farlow’s
attitude towards a number of other problems and persons.

Since 1890 dissatisfaction had been growing, both in the Old and the New
World, about the way the Botanisches Centralblatt, at that time the only
abstracting journal in our field, was being conducted.

In October 1900, Goebel, who had already repeatedly viewed his opinion
about the matter and who had even been planning to establish a second
abstracting journal in Germany, met Lotsy in Ambach and discussed the
situation with him. Lotsy was much interested and subsequently held con-
ferences with R. von Wettstein in Vienna, with M. Raciborski in Lemberg,
and with S. Nawaschin in Kiev. In November 1900 he wrote Goebel about
a definite plan to start a new international botanical abstracting journal, as
well as an international association of botanists which would not only edit
and publish the journal, but would also undertake many other things.

In the United States there was at that time no less dissatisfaction. In an
address (December 27, 1899) to the American Society for Plant Morphology
and Physiology, Dr. Farlow had spoken in detail about the need of either
changing the Botanisches Centralblatt or establishing a new journal. A
committee was then established consisting of Dr. Farlow, Dr. D. T. Mac-
Dougal and Dr. von Schrenk. This issued a first report on June 27, 1900

1 The author is under considerable obligation to F. O. Bower, Sc.D., LL.D., F.RS., of
Glasgow and to William Trelease, Sc.D., LL.D., of Urbana, Ill., for their help in securing
data about the history of international relations work in botany and for their permis-

sion to reproduce the letters reprinted in this article.
2 Dr. J. C. Bay’s earlier efforts in this field are not discussed in this article.

71

rz FartowiA, Vor. 2, 1945

and a definite report on December 27, 1900. The committee, and Dr. W. F.
Ganong, at that time secretary of the society, had been very successful.
They obtained from Dr. O. Uhlworm, the editor of Botanisches Central-
blatt, various promises, described in detail in Dr. Farlow’s letter of Feb-
ruary 4, 1901, reproduced below. As we will see from the following letters,
Dr. Lotsy and his group went on with their plans and established the Asso-
ciation at a meeting in Geneva in August, 1901. The society acquired the
Botanisches Centralblatt. We reproduce now three letters by Dr. Farlow
which throw some interesting light on the events of that time. The first
letter is addressed to Dr. Bower of Glasgow, an old friend of Farlow and
also a friend and supporter of Lotsy and his work. The second letter is
addressed to Dr. Trelease, a former student of Farlow, who was also much
interested in botanical bibliography and who had undertaken to share in
the work of the American editorial committee of the Botanisches Central-
blatt, as referred to in Farlow’s letter of February 4, 1901 to Dr. Bower.
The third letter is again addressed to Dr. Bower. Lotsy’s letter to Farlow,
of which a copy is in the Chronica Botanica Archives, is not so impulsive or
Naive as one may expect after Farlow’s reaction. Lotsy was planning some-
thing much bigger than only a journal, he was dreaming of an international
botanical bureau, as many others after him have done. Farlow was only
interested in the journal and seems to have had little sympathy with Lotsy’s
plans and way of thinking.

Cambridge
Feb. 4, 1901.
Dear Prof. Bower,

I suppose that last June you received a copy of the report of a committee of
the Society of Plant Morphology and Physiology appointed to consider the best
means of securing a good record of current botanical literature. A second and
final report was sent you about a month ago.

Assuming that the reports were received by you it is not necessary for me to
repeat their contents now. The last report was presented by me as chairman of
the committee at the meeting of the Soc. Plant Morph. in Baltimore on Dec. 27.3

After I returned from Baltimore I received a letter from Lotsy which took me
entirely by surprise. I did not know that there was any project for a new journal
of reviews in project. About six months ago, I believe, I had a letter from Goebel
in which he expressed great dissatisfaction with the Centralblatt and said that
he had sometime thought that he would like to start a new journal. But from
what he said I did not understand that there was any intention of doing so now
and the project seemed to me to be all in the air. Up to June of last year the
reply of the editors of the Centralblatt to our suggestions for improving their
journal had not been satisfactory to us but at the meeting of last June our com-
mittee was directed to continue negotiations with the Centralblatt and, if they
would not do what we wanted, to present a plan of a new journal at the meeting
of Dec. 27, 1900.

Early in November the editors replied agreeing to do all that we had asked

° Cf. Farlow in Bot. Gaz. 29: 64. 1900. A further report was issued in 1901 by Farlow,
MacDougal and von Schrenk.

VERDOORN: INTERNATIONAL ABSTRACTING 13

them to do. They decided to put all their worthless articles into the Beihefte and
to put all the reviews and the list of new works into the Centralblatt and also to
allow one to buy the Centralblatt without the Beihefte (which nobody wants) at
the old price. They also asked us to select American editors whose names should
appear on the title page to see that American notices were kept up to the mark.
They furthermore said that they hoped to have similar editors appointed by
botanists in other countries.

Such an arrangement seemed to us at Baltimore to be a good one and a com-
mittee was appointed (of which I am a member) to inform the Centralblatt that
their proposition seemed satisfactory and to nominate American editors. In fact,
we have already selected one editor and have been considering other candidates.

You will imagine my surprise after returning from the Baltimore meeting on
hearing from Lotsy 4 that a new journal was proposed and he wished me to join
his International Association. I should be glad to join the Association were it
not that its main object appears to be the starting [of] a new journal in opposi-
tion to the Centralblatt. Having, in perfectly good faith, gone so far in our
negotiation with the Centralblatt it seems to me that it would not be honest for
me, no matter how brilliant a list of names of associates Lotsy reports, to join
now an association which is directly opposed to the Centralblatt. ...

I have written this to you not knowing to what extent you feel confidence in
Lotsy as a manager but merely wishing to let you know the circumstances which
prevent my accepting Lotsy’s invitation. I feel that we here are in duty bound to
give the Centralblatt a fair trial under the proposed arrangement. It may or may
not succeed but, if it does not succeed now, it will be in part our own fault. If it
turns out that the Centralblatt people won’t do as they should, then all the sub-
scriptions will go to Lotsy’s journal if it is a good one. I take it for granted that
the Centralblatt people mean to act in good faith. Certainly they already have
-dropped original articles from the Centralblatt itself, and, as far as American
editors are concerned there has not yet been time for us to select editors and,
until that is done, one could hardly expect a change in the character of American
reviews. As far as improvement in the quality of the notices is concerned, I
should expect that it would be gradual not sudden and we cannot complain if it
takes at least six months before a perceptible change is seen.

Yours, truly,

. W. G. Farlow.

Cambridge, March 1, 1901.
Dear Trelease,

This noon I received a letter from Lotsy which quite took away my breath.
After receiving my cable and letter saying that it was too late for me to accept
his invitation to join his International Society since we had promised to help the
Centralblatt, Lotsy decided to buy the Centralblatt. They have agreed to sell out
to him and he now wants me to cable to him at once that I will join the new
Society. I shall take the liberty of thinking over the matter until tomorrow
morning. I have consulted Thaxter and he agrees with me in thinking that I can
and should accept. The way things have turned out is, I think the best solution
of the complication.

* For a biographical account of Lotsy vide Genetica 13: 1. 1932.

74 Fartowia, Vor. 2, 1945

Uhlworm 5 is to be kept for five years and the American editors we may select
will be accepted by the managers. It is proposed to capitalize the Centralblatt at
$10,000. Stock is to be placed on the botanical market and we shall have the
opportunity to invest. The assets on hand are something like 100 complete sets
of the Centralblatt &c. The whole thing seems very funny but I think it will
succeed. The main point as far as we are concerned is that our Committee is
practically tied up for the present awaiting the cons[u|mmation of the purchase
by Lotsy. There is one snag in the way; the purchase must be confirmed by the
board of the new Society. What the board is I do not know but I suppose that
Lotsy’s action will be endorsed.

It is too dark to see what I am writing and I must close.

Yours, truly,

[W. G. Farlow. |

Cambridge, April 5 1901.
Dear Prof. Bower,

Since I wrote to you in regard to my correspondence with Dr. Lotsy I have had
another communication from him which surprised me even more than his previous
letters. A week ago I had a letter in answer to my inquiry as to the individual
responsibility of stockholders in which he said but little about the Centralblatt
but instead wrote several pages describing a project of forming a sort of museum
for collections of microscopical slides and type-specimens in general and a bureau
for collecting all sorts of miscellaneous information for botanical travellers.

I presume that you have received a similar letter. The letter seemed so very
visionary and reckless that I could not help coming to the conclusion that Lotsy
is what we call a “crank.” He certainly must have very little commonsense. To
manage the Centralblatt so that it would become a first-class journal will be quite
enough to absorb all his time and energy without launching out on any such wild-
cat sort of speculation as his proposed museum. He stated that he needed an
endowment of $130,000. He added that Goebel despaired of being able to raise
that sum in Europe. I am not surprised that Goebel thinks so.

Lotsy however is confident that some of the numerous American millionaires
will give him all he wants. He said that if I thought that I could introduce him to
a number of our millionaires, he would start for America at once and come to
Cambridge to see me. Unfortunately I do not know any millionaires and if I did,
I most certainly would squeeze all I could out of them for our own botanical
department which needs money very much indeed and I should never think of
asking them to give anything to Lotsy for a museum. I feel sure that all our
botanists who are connected with universities or other institutions would act just
as I have said I would do. There is only one rich botanist in this country and he
certainly would give nothing to Lotsy in any case.

I have not yet answered Lotsy’s letter because he said that he was going to send
me at once the proof of the circular to be issued in regard to his new Society.
After I have seen the proof of his circular I shall be able to write to him some-
thing definite.

° Shortly after Lotsy’s Association acquired the Botanisches Centralblatt he and Uhl-
worm quarreled. After this Uhlworm retired entirely. He may have had some part in
abortive attempts, made in Berlin a little later, to start another abstracting journal.
These plans, however, did not materialize (cf. letter of Thost to von Schrenk, 15 Jan.
1902, in Chron. Bot. Archives).

VERDOORN: INTERNATIONAL ABSTRACTING 75

He will not be able to see me in Cambridge because I intend sailing for Europe .
on April 30. My plan of going to Europe is due to the fact that I have been
asked to attend the celebration of the University of Glasgow in June as a delegate
of the National Academy of Sciences. I am so busy that it will be difficult for
me to leave then but there [are] certain matters of business for me to attend to
in Europe and my wife has two nieces of whom she is guardian now studying in
Paris and she wishes to go to see how they are getting along. Unless something
happens to prevent, we shall start for Cherbourg on the 30 and after attending to
my affairs on the Continent, we shall go to England and then to Scotland so as to
reach Glasgow in time for the meeting on June 15. Therefore there is a chance
that I may have the pleasure of seeing you in June.

Yours, truly,

W. G. Farlow.

Between January 1902 and the last year of the First World War, Lotsy

edited the Botanisches Centralblatt with great care. Except for Schramm’s
Botanical Abstracts we have never had anything in our field that was as
satisfactory as the Botanisches Centralblatt between 1902 and 1919. With-
out doubt it was more satisfactory in its early years than later when Lotsy
became involved in many other activities. During this entire period, Dr.
William Trelease acted as American editor (“Président du Comité National
des Etats Unis”). He gave freely of his time and energy and has done more
than any other North American botanist to report well in the Botanisches
Centralblatt, then still the only abstracting journal in our field, on the dra-
matic progress made in the U.S. A. between 1900 and 1918. He had the
.codperation of a small, rather constant group of American ‘“Rédacteurs
spéciaux.” Of some of these men the names do not often occur in the col-
umns of the Centralblatt. That the journal was not always too satisfactory
from the point of view of the North American editors and their colleagues
we may gather from the following two letters of Dr. Farlow: —

Cambridge, Mass.,
Jan. 27, 1908.
Dear Trelease,

I see no reason why your letter should not be sent to Lotsy except that —
suppose that he lets you go? Then I do not envy you if you feel that you must
go ahead and start an American journal. I understand your reference to the
Americans as “prepared to start an abstract journal of their own when etc” to be
a bluff rather than a statement of fact. My recollection is that when I tried to
suggest a journal for giving abstracts of American literature I was very, very
coldly received. Galloway I remember as particularly “pooh-poohy” and declared
it impossible. In fact, I do not remember that anyone gave me any real encour-
agement. Perhaps you mean an American journal giving abstracts also of Euro-
pean papers. Ughm!

I had no confidence in any real improvement in the Centralblatt when I heard
that Lotsy was to run it . . . He is impractical and given to putting all sorts of
unnecessary irons on the fire. Still I suppose it was a question whether any better
man could be got.

Although far from what one would like, it may perhaps be the fact that, as at

76 FartowiA, VoL. 2, 1945

present conducted, it is as good as such a journal is likely to be. Some want
notices like those in Engler’s Jahrb. but that is impossible unless few things are
to be noticed. The “sympathetic” notices as I have heard them called can only
come from experts who have taken their time to read and use the books noticed
and that takes a long time and experts are not to be depended on in a journal like
the Centralblatt. People complain of Just [Botanischer Jahresbericht]| that it is
too far behindhand. For my part, I think that Just is as near perfection as any-
thing of the sort can be.

I was sorry not to go to Chicago but I was too dreadfully busy and I was
especially sorry that none of our other botanists could go but Goodale was not
well enough and Robinson, Fernald & Thaxter were driven by the printer and
Jeffrey had only that time for going after fossil material. Like us, I suppose that
you have had a remarkable winter. I do not object to the mildness.

W. G. Farlow.

Cambridge, Mass.,
Feb. 10, 1908.
Dear Trelease,

While you are “jogging” the Centralblatt let me suggest one point where it
seems to me that something should be done. I do not like the idea of giving
notices in the Neue Litteratur. Why should not the notices be given all of them
in the regular place? It looks as if the reviewers despairing of getting their
notices into the regular pages smuggled them in in the Litteratur where no one
would look for them. It is unjust to writers so noticed since it looks as if their
papers were not worth noticing.

Generally for my purpose the very short abstracts in the Neue Litteratur are
enough and that brings up the question, Why not give all the notices in this short
form with the titles issued in Neue Litteratur? Who wants (see p. 24) more than
a page of Dittrich & Pax, Herbarium cecidiologicum? The trouble is and prob-
ably always must be that a good many reviews are given to immature students
who have no sense of proportion.

Yours, very truly,

W. G. Farlow

The Botanisches Centralblatt was first published for the Association by
Messrs. Brill of Leyden and printed in Germany at the same press where it
had been printed before it was acquired by the Association. Later it was
printed in Leyden at Sijthoff’s and published by Messrs. Fischer of Jena
for the Association. Though the Association remained a purely international
society till the end (in 1917 its council was formed by D. H. Scott, Wm.
Trelease, and J. P. Lotsy, and its editorial board by Wm. Trelease, C.
Bonaventura, A. D. Cotton, C. Wehmer, and C. H. Ostenfeld) and though
its journal was edited and printed during the First World War in a neutral
country, the fact that it was published in Germany and that much (without
doubt much too much for an international journal) of the contents was in
German caused highly complicated difficulties with American (not British! )
authorities and resulted in only a few copies of the journal reaching Ameri-
can subscribers. This and other things caused considerable ill-will amongst
a number of North American botanists. In Great Britain, where the inter-
national character of the journal was better understood, as Drs. Bower,

VERDOORN: INTERNATIONAL ABSTRACTING 77

Seward and others have assured me, there never was any feeling against the
journal or the society. Feeling dissatisfied, a group of U. S. colleagues,
amongst whom Dr. Livingston was very influential, established at that time
Botanical Abstracts.

This probably happened too recently to allow us to decide if it was wise
to do so, especially as Dr. Lotsy, tired of the difficulties of keeping his jour-
nal going during the war years, offered the North American group the fullest
possible codperation of himself and the Association, a somewhat informal
offer in which he would, without doubt, have had the backing of the Council.
This offer was not accepted. It was hardly considered, which seems a great
pity to the student of international affairs. As a result, we have two not
entirely successful ® abstracting journals from 1919 onwards.

Since this article is not primarily concerned with these problems, it may
be more interesting for us to see what Dr. Farlow, then nearly seventy-five,
had to say about these developments in 1917, 1918 and 1919 in his letters to
Dr. Trelease, the chief American editor of the abstracting journal of the As-
sociation Internationale des Botanistes till its discontinuation.

Cambridge
Nov ctA. 1OL7.
Dear Trelease,

I agree with you entirely about the desirability of improving the Bot. Central-
blatt rather than starting a new and similar journal in this country.

Reddick was here a fortnight ago and asked my opinion about an American
Centralblatt to include everything in this world and apparently everything in the
Solar System. I do not think that he favored it. His account of the genesis of
the idea of an American Centralblatt was not altogether clear. He said that
Stevens had been impressed by the success of the new American journal of
bacterial diseases to which he said there were a good many medical men who
subscribed. The universal success of that journal did not impress me because I
had never seen a copy and had not heard anyone speak about it. If, however, as
Newcombe appears to desire, the new journal is to include not only reviews of
bacterial papers but also all papers on plant diseases and even botanical papers
on non-bacterial or pathological Sue I have an idea that the medical sub-
scribers would soon drop out.

My first answer to Reddick was that it seemed to me to be folly to start a new
journal of any kind at this time for financial reasons. Until the war is ended no
one knows whether he has any money to spend or not and it is now evident that
the war will not end for a long time if it is to end in favor of the Allies and, if
not, Germany will take all the money that can be squeezed out of us.

Reddick said that it was expected (?) that the Brooklyn Bot. Gard. would
furnish the money or most of it. My answer to that was that if Brooklyn fur-
nished the money they would have to do with it what the Brooklyn people wanted
and that the desires of those who did not furnish any money would not count
for much.

® Tt is not the fact that they were commercially not too successful which I deplore, but
the fact that none of them was truly international, while in 1919 we did have the frame-
work of a truly international botanical society! Cf. also my remarks in Nature 154:
595. 1944,

78 FartowlA, VoL. 2, 1945

Furthermore, I told him that I did not desire to see established a new journal
of reviews. As it is, I am loaded down with journals which give reviews of the
same papers over and over again and unless the current journals would agree to
give up reviews or notices of new papers I did not want to be bothered with more
reviews. He said that he believed that Mycologia would agree to give up reviews.
Mycologia, however, is practically the organ of the N. Y. Bot. Gard. and has no
interest in noticing any work not done in connection with the Garden. Other
journals probably would not agree to give up reviews as they help to sell the
journals. Those who take only one journal would be likely to select that one
which gave reviews in addition to original papers. The notices in the Bot. Gaz.
for example certainly draw in subscribers.

Again I have little faith in the ability of our enthusiastic friend to produce a
journal which would replace the Bot. Centralblatt. As you say, at the present
time that journal does not help us as we do not receive the current numbers but
after the war I presume that it will reach us as before. If the Centralblatt gives
out later, then it will be soon enough to think about an American journal.

The Bot. Centralblatt certainly might be made better. I hope that it will. I
have my own ideas of what is desirable. In the first place no author should be
allowed to write a notice of his own papers unless he happens to write in a lan-
guage like Japanese, or Czech perhaps, which is not readable by the ordinary
editors. In the next place the editors should not be allowed to turn over papers
to their students to review unless they themselves revise the reviews themselves.
The younger students are not capable of writing good condensed reviews and,
being paid by the page, there is no inducement to be brief.

My own opinion, not shared by some, is that in a Centralblatt, as far as pos-
sible all papers should be noticed as soon as possible after their issue and that
the reviews should be descriptive not critical. That is, one should have the titles
and a brief statement of contents of as many papers as possible so that everyone
can tell whether a paper on subjects in which he is interested is one which he
ought to look at and read. Really critical reviews are generally too long for a
Centralblatt, they must be written by experts and they cannot be written at
short notice.

Robinson does not agree with me. He thinks that reviews should be “sym-
pathetic.”” That means I suppose in a review of a systematic paper there should
be a list of at least the new species and perhaps others with comments. Those
not interested in the subject would I think have little ‘‘sympathy” with reviews
of that kind.

I really do not believe that there are any botanists to speak of who really want
a new journal but at botanical meetings the crowd is easily persuaded to vote for
anything proposed by a pleasing and plausible speaker. If the question comes up
at Pittsburgh (I shall not be there) it may be voted to start the journal. Some
very foolish things have been voted by the A. A.A.S.

The war news is certainly depressing and people here see no end of the war in
sight unless it is to be very unfavorable. Our family have lately been rather
anxious. My niece we hear has reached France. Her brother, an aviator, is on
his way somewhere, we do not know where. Our available relatives are now all
in Europe except one who is still at the Hydroplane school in Pensacola.

Remember me kindly to Mrs. Trelease.

Yours, very truly,
W. G. Farlow.

VERDOORN: INTERNATIONAL ABSTRACTING 79

Cambridge
Febr. 26, 1918.
Dear Trelease,

Do you remember the Guepinia lutea Bres. which you collected in Alaska? I
have recently received from near Seattle a Guepinia near G. merulina which may
be your plant. We have here what I take to be G. merulina but it is rather rare.
The Seattle fungus has spores which are rather larger and more septate than in
our plant. The basidia are narrow and the tips are very short . . . but I do not
know whether this is constant. I had only a very little of the Guepinia with a
large junk of the host. I have written for more and, if I get it, I will send you
some for comparison if you have any of the G. /utea on hand.

I am just sending in my preliminary ballot of the Nat. Acad. I put on the
names of Atkinson and Forbes. The latter will probably be elected. About Atkin-
son I am more doubtful for there are some who are strongly opposed to him. At
any rate I do not see how I can go to Washington in April to the meeting. It
appears now to be practically impossible to secure a room at any hotel and the
journey is subject to long delays. Prof. Palmer a short time ago was 27 hours on
the road to New York. To be sure he was so imprudent as to take the boat on a
very cold day and was blocked by ice.

Reddick was here two days ago and told me about the Pittsburgh meeting.
O. L. Howard was also here apparently to make arrangements about the Ass.
meeting in Boston next winter. He is down on Richards for advising giving up
the last meeting. If next winter is anything like this and no more coal than now,
the A.A. A.S. will not be welcome guests.

Both Reddick and Howard said that the Pittsburgh meeting was a great suc-
cess. Reddick had become a convert to the new extract journal and he said that
they were going to start work at once and get out the first number in June. It
appears that the Waverly Co. of Baltimore which issues the bacterial extract
journal thinks it can make a success of the new journal. It is supposed that the
larger libraries would be willing to pay a good price if they receive slips during
the year and the whole in book form at the end of the year with the right to buy
other copies at a reduced rate. Something was said of charging $10 a year to
individuals. I thought that not many botanists would want to pay that amount
but Reddick thought they would if the journal was indispensable. The zf seems
to be important but also uncertain. The Brooklyn Garden was said also to be
ready to help. I was told that it was Macdougal who was negotiating with the
Waverly Press.

I wonder what you wrote on your card of the Man Power Census sent by Kern
I presume. Not knowing what else to write I wrote “Mycology” diagonally across
the card but Thaxter says that that will only show what a fool I am for only a
short time ago a prominent Phytopathologist said that he thanked God that he
was not a mycologist. A letter has just come from Whetzel asking me to attend
a meeting in Boston on March Ist to organize work. I can hardly suppose that
they want me to help them do the talking for they are already well equipped in
that department.

Winter seems to be gradually slipping away. That is we now have some com-
fortable days as well as some very uncomfortable and March is still to come.

Yours, very truly,
W. G. Farlow.

80 FartowlA, VoL. 2, 1945

Cambridge,
Dec. 27, 1918.
Dear Trelease,

Things are in a mess. I presume that the fact is that the Centralblatt manage-
ment has become unpopular on account of Lotsy’s . . . domineering and not
conciliating [manners]|.7 Chodat I presume represents the attitude of some Euro-
peans outside Germany and they would be glad to see a rival established in some
other country. Most Americans, especially the young men, seem determined to
cut out the Centralblatt as too decidedly under German control... .

If Botanical Abstracts seems to fill the bill it will have a circulation abroad as
well as here. My question is, will it fill the bill? The first number was not re-
assuring but later numbers may be better. Livingston’s statement that botanists
were much pleased with it would have more weight if I knew who the botanists
were who liked the first number. . . .

At any rate, the war is now on. The Bot. Abstracts committee after Livingston’s
letter do not desire to conceal the fact that they propose to squelch the Central-
blatt if they can. If I were you, I would wait until getting an answer to your
letters of Dec. 24 to Livingston and Williams & Wilkins. They will have to
say whether they intended to have their letters sent to the Ass. Intern. or not.
Very likely you already have their answers. If they did intend to have their
letters, or their contents, sent, then of course they should be sent. If they say no,
all you can do is to tell Lotsy that you have communicated with Williams &
Wilkins and that they have no intention of explaining or apologizing.

As to your own relations to the Ass. Int. Bot., if you have reason to suppose
that its affairs are run by “secret diplomacy” as Chodat says or, if you have
reason later to think so, you had better withdraw. Otherwise you will be in the
position of endorsing measures which you know nothing about and may not
approve at all.

Whether the abstracts will win the day is after all to a great extent a question
of dollars and cents. Just now it is a matter of sentiment but in the end the
question will be can we in America compete with the Germans in cost of printing
and can we control at as small cost as the Germans a large body of reviewers,
revisers, etc. No one knows what the condition of scientific workers is to be in
Germany. It may be that the hitherto large number of trained men willing to
spend their time in writing reviews, bibliography and what may be called dic-
tionary work at a very small price will be much diminished. I am skeptical as to
the reduced condition of Germany. I believe that they are still in good condition
and really better off than France and perhaps as well off as England in equipment
for scientific work. Of course much depends on what sort of a new government
is formed in Berlin. So far at least the German leopard shows no signs of chang-
ing his spots.

Yours, very truly,
W. G. Farlow.

7 Lotsy, his work and his personality, has often been criticized. He was, however, a
great man, and has done more than anyone in the history of our scientia amabilis to
promote international relations, cooperation, and understanding. Dr. Farlow’s impres-
sion, without doubt shared by others at that time in the U.S.A. (but not in Great
Britain, where Lotsy had more friends than anywhere else), of Lotsy’s political sym-
pathies is so erroneous that one can hardly understand it. It teaches us how easily
misunderstandings may arise, during years of mental stress, even amongst the repre-
sentatives of a single race.

VERDOORN: INTERNATIONAL ABSTRACTING 81

Cambridge,
Dec. 31, 1918.
Dear Trelease,

A letter from Mangin dated Dec. 13 contains the following which will interest
you.

“Nous avons eu a discuter la question de l’Association Internationale des Bo-
tanistes. Cette association est virtuellement dissoute et les membres ne (parvien-
dront . . . word not clear but this as far as I can make out) la liquidation. Mais
il serait utile de la reconstituer entre alliés et d’utiliser comme urgane de cette
association les abstraits botaniques qui ont fait leur apparition chez vous.”

If I understand the above, the association is bankrupt which can hardly fail to
be the case since subscriptions must have fallen off very much. I am a loser to
the extent of $500, the amount of bonds I took... .

It seems to me just as well that the Centralblatt should stop. I should prefer
an international reorganization to a purely American conducted journal but Bot.
Abstracts has started in ahead... .

Happy New Year,
W. G. Farlow.

Cambridge
Jan. 26, 1919

Dear Trelease,

It is to be regretted that Mangin’s letter to me and Flahault’s letter to Duggar
did not reach us before the Baltimore meeting. Had that been the case it would
have been possible for you to state the facts contained in those letters to the
representatives of Botanical Abstracts at Baltimore and you might (?) have been
able to convince them of the desirability of trying to make some international
combination which would result in making the Abstracts really an international
organ.

Now, the case seems to be rather hopeless. It looks as if our younger botanists
were cock-sure that our American botanists were better able than any others to
issue an international journal. I hear that a contract has been made with Williams
& Wilkins to run the Abstracts for five years. Of course W. & W. are not well
informed as to the requirements of a real international journal and are quite
satisfied that the Abstracts, as illustrated by the single number issued is just
what is wanted.

The number issued did not strike me favorably considered as a general inter-
national review of botanical literature. It was really nothing but a guide to
American botanical literature and the value of notices of papers in Bot. Mag.,
Torr. Bull., Phytopathology, Am. Journ. Bot. does not seem to me to be great
since all our botanists practically see all those journals for themselves. Further-
more the comparatively large amount of space taken by Shull to review papers
having no relation to botany is looked upon unfavorably by all botanists I know
and, unless he is to be restricted in later numbers, the journal will suffer.

Flahault’s letter is interesting. He speaks of the possibility of the non-germanic
members of the Assoc. Intern. breaking up the agreement with Fischer and ap-
parently is not sure whether it can be done. Of course it can be done for in these
times international pre-war agreements count for nothing.

82 Fartowia, VoL. 2, 1945

It looks to me as if the feeling against the Germans is too great at the present
time to make it possible to form any international union. Opinions may be modi-
fied later but not for some time to come. Meanwhile the Bot. Abstracts is to be
pushed in the hope that it will replace the Centralblatt. If it is to succeed in that
object, future numbers will have to be very different from the first number.

There is no doubt, as you say, that an abstract journal with a German title
would not be taken in the allied countries now. But any journal would have to
give abstracts of German botanical papers as well as others. Such reviews need
not be in the German language although I see no reason why they might not.
There are certainly more botanists here who read German than who read Italian.
It may be said, however, that one can often “guess” what the Italian means.

The meeting 1° in London referred to by Flahault appears to be merely the
general meeting of scientific delegates and it could not have had any special sig-
nificance for botanists. I should be glad to know what Scott’s opinion is.

In short, the Centralblatt is dead and cannot be revived as a world journal
under a German title and issued by a German house. . . .

From the point of view of science itself, I do not think that a union of English,
French, Italians and Americans alone would be for the general advantage. There
therefore remains the question, what are the chances that Botanical Abstracts
will fill the place of the Centralblatt? That cannot be even guessed now.

I hear that the editors of Bot. Abstr. wish to raise a fund of $1000 to pay
reviewers. That would not go very far and the Brooklyn Garden people or others
specially interested ought to be able to get that amount without a general appeal.
If their credit is no greater than that, the future is not promising. . . .

W. G. Farlow.

Though Farlow’s interests were wide, ranging from the lowest to the
highest of the cryptogamic plants, from general botany to taxonomy and
phytopathology, from music to the humanities, though he travelled very
widely and intensively in the interval between 1872 and 1874, and he also
visited Europe thereafter, he was not in the broadest sense an internation-
alist, as were some of his contemporaries. His own collections came first
and these he protected for future botanists by adopting measures, often
criticized by his colleagues, that would as far as possible assure that the
collection of books and specimens be available to future botanists. How-
ever, despite this preference for his own collections, he was much interested
in the personality of his foreign colleagues, as Setchell (Mem. Nat. Acad.
Sci. 21: 14. 1927) has told us, but as a practical and typical New Englander,
was still more interested in the results of their work and especially in the
abstracting and listing of the botanical literature on a truly international
basis. To this extent he was an internationalist, and as his early letters
show, a pioneer in American abstracting work.

CHRONICA BOTANICA
WALTHAM, Mass,

* One of the early meetings of the Conseil International des Recherches (now Inter-
national Council of Scientific Unions).

PLATE I

2(1): 84-95 FARLOWIA January, 1945

DR. FARLOW’S INFLUENCE ON MYCOLOGY
WiLLiAM H. WESTON, JR.

In attempting to evaluate the imprint on a field of science made by an
outstanding worker in that field it soon becomes obvious that lists of publi-
cations often tell but part of the story and that a man’s influence may
make itself felt through various channels less obviously recorded. In Dr.
Farlow’s case, even the partial list of his publications in two of the bio-
graphical memoirs noted in the introduction to this number, reveals that
the scope of his activities was very broad including work on algae, bacteria,
fungi, lichens, liverworts, mosses, ferns, and flowering plants. In addition
he participated actively in botanical affairs for more than forty years, stim-
ulated and promoted the development of several fields which were part of
the rapid progress of botany in the United States during that time and
played an active and constructive part in maintaining effective working
relationships between botanists in this country and in Europe.

Withal it was probably the field of mycology which he influenced most
strongly. On attempting to analyze his effect on this branch of botany, it
becomes clear that his was a manifold influence making itself felt to some
extent through his teaching, to some extent through his published work,
but chiefly through his devoting to the service of mycology his wide knowl-
edge of the fungi and his extensive herbarium, library, and indices on which
this knowledge in part was based. The force which gave especial strength
to all these lines of influence was the power of his dynamic, stimulating,
and catalytic personality. His influence on mycology was thus of the same
pattern as his much wider influence on the whole field of cryptogamic
botany.

To understand the conditions which shaped his early aspirations into
definite objectives that permanently motivated his life’s work it is necessary
to appreciate the status of botany in the United States during the forma-
tive years when Dr. Farlow began his training.

During the 1860s, botany as a science numbered very few professional
devotees in its ranks in this country and was largely the avocation of those,
such as ministers and doctors, earning their living in other professions. As
an indication of conditions at that time it is worthy of note that Asa Gray,
Goodale, Farlow and others in this country, like De Bary and numerous
botanists in Europe, on graduation secured medical degrees because this
was the only available degree involving training and graduate work appli-
cable to botany and also because the profession of medicine would yield a

Doctor and Mrs. Farlow on the porch of their summer home in Chocorua, New
Hampshire. Picture taken a year or two before his death.

85

86 FaARLowIA, Vor. 2, 1945

living permitting work in the unremunerative and precarious field of bot-
any. Also, botany in the United States consisted almost wholly of work on
the flowering plants with emphasis on their taxonomy, nomenclature, flo-
ristics, and gross anatomy. The leading exponent and symbol of this type
of work was Asa Gray who from his contact with the botanists of Europe
was well aware, upon his return from abroad in 1870, that rapid expansion
of botany was beginning and if the United States was to keep pace, other
aspects of the science should be developed. Accordingly, as he was fully
occupied with his own field of work in the Fisher professorship of Natural
History which he had held since 1842, he was instrumental in the selection
and appointment of two promising young men well fitted to develop im-
portant fields in the rapidly expanding realm of botany.

The first, George Lincoln Goodale, by his skill as a teacher and lecturer
gave impetus to the growing field of plant physiology which was rapidly
assuming importance in Europe, and by his efficiency as an organizer and
administrator built up the Botanic Garden, which already had more than
half a century of development and service to its credit, to its final perfection
of usefulness and beauty; he vitalized and enriched the collections of the
Botanical Museum started by Gray around 1860 and brought about the
building of the spacious and well-equipped Botanical Section of the Uni-
versity Museum.

The second, William Gilson Farlow, as the man to devote himself to the
development of cryptogamic botany, was a wholly logical choice, for even
before graduating in 1866 he had shown unusual interest and promise in
that newly burgeoning field while his further preparation, through attain-
ing his medical degree in 1870, was the same as that which Gray and De
Bary, the two who influenced him most strongly, had themselves received.
After being appointed Gray’s assistant in 1870, Dr. Farlow worked with
him for two years increasing the teaching emphasis on the lower plants,
and carrying on research of his own, especially in the algae. Then, realizing
the need for further training which only the laboratories of Europe could
offer at that time, Dr. Farlow spent the two years from the summer of 1872
to the summer of 1874 travelling through England, the Scandinavian
countries, Russia, Germany, France, Italy, and Switzerland and working in
various laboratories, especially at Strasbourg in Anton De Bary’s labora-
tory, at that time the world’s leading center of mycological activity.

These were highly important years in Dr. Farlow’s development. He
met distinguished botanists and visited and worked in their laboratories,
establishing valuable contacts and friendships which he maintained through
correspondence and occasional visits for many years. He became acquainted
with the botanists of the period, both the recognized authorities of the time
and the students working with them, who were to become the leading
mycologists and algologists of their generation. He visited the famous
herbaria and libraries and the convincing evidence of their indispensable
function as working tools essential to fundamental botanical work gained
through his own use of these treasures, started him on the discriminating

WESTON: FARLOW’S INFLUENCE ON MycoLocy 87

purchases of books, periodicals, exsiccati, and specimens that were the
nucleus of the Farlow Herbarium and Library so important in the ensuing
development of cryptogamic botany in this country. Also in these two
years he became aware of the trends and currents in botanical work and
began to appraise what was fundamental and enduring, what held promise
for future development, and what was merely transient in its temporary
popularity.

The evidence seems clear that Dr. Farlow returned from his two years in
Europe with the firm conviction that the development of cryptogamic
botany in the United States would be aided most effectively by three essen-
tial facilities. First, there would be needed a herbarium of authentic ref-
erence specimens to facilitate dependable identification of critical material.
Second, accompanying this reference herbarium an extensive and complete
reference library on the fungi and other lower plants would be an imperative
supplement for adequately identifying material and establishing its rela-
tionships and significance. Third, supporting both these, there would have
to be an index comprising all significant published references to such ma-
terial. One gains the impression that this enduring conviction was not so
much the result of his earlier training with Gray as it was the logical out-
come of his own analysis, during these two years, of the fundamental basis
of botanical supremacy in Europe. There he saw, beneath the fads and
fashionable trends which were gathering those adherents who gravitated
naturally to the most publicized aspects of botany, the sound foundation
on which botanical progress in Europe had depended. He realized the inter-
dependence of such fundamental aspects of the science as taxonomy, mor-
phology, and physiology, and retained his sense of proportion throughout
his work in De Bary’s laboratory where, even in the 1870s, the develop-
ment, reproduction, life histories, parasitism, and other activities of fungi
were stressed and taxonomy was rather looked down upon as old fashioned,
the systematists with their vasculi being significantly and disparagingly
dubbed “Heusammler.”” There he worked on the morphology, development,
and reproduction of fungi and other cryptogams and there he completed |
the work on the apogamous development of Pteris, work which, on publi-
cation in 1874 immediately attracted wide attention, a pioneer contribution
to a field later developed by others to far reaching significance. His brief
but vivid pictures of the atmosphere, activities, and personalities in De
Bary’s laboratories as well as in others, such as Bornet’s, in which he
worked for shorter periods are interesting sidelights on botanical history,
enlivened by his characteristic humor and well worth seeking out in the
bibliographies already noted.

In the biographical sketches which appeared after Dr. Farlow’s death,
there is detectable a tendency to apologize for his not too extensive list of
published contributions. This is hard to understand for certainly his biog-
raphers were well aware that the major part of his activity had been the
accumulation and working over of collections not as an objective in them-
selves but as an essential working tool applied to the service of cryptogamic

88 FARLowIA, VoL. 2, 1945

botany. After Dr. Farlow’s death in 1919 his herbarium, when officially
taken over by Harvard, comprised around 500,000 specimens and the
greater proportion of these had been studied and worked over critically by
Dr. Farlow himself. Many of them bore enlightening notations in his
characteristic handwriting, evidence of their having been compared criti-
cally with exsiccati, type specimens, and other pertinent material and care-
fully studied in relation to the significant and authoritative literature. No
one who has consulted his extensive herbarium can fail to realize that it had
been worked and reworked by Dr. Farlow until it became a fundamental
part of his knowledge, his first hand familiarity with this material and
with the wealth of original work in the extensive library he accumulated
being the basis of his impressively wide knowledge so valuable to many
workers in this country and in Europe.

The list of his publications is ample evidence of the breadth of his interest
and knowledge since it ranges through the cryptogams from the algae to
the ferns, and in the fungi, which have the major emphasis, touches on all
of the chief groups from the bacteria and Myxomycetes up through the
Phycomycetes, Ascomycetes, and Fungi Imperfecti, to the Basidiomycetes.
Here are found the several published lists comprising some of the many
fungi studied and identified by him, fungi representative of many different
localities, some collected by him and by his associates in the vicinity of
Boston, in the White Mountains, and elsewhere in this country, others
brought back from the arctic by the Peary expedition or by other expedi-
tions from such regions as Alaska and Labrador, or the Galapagos and the
Bahamas, or from Bolivia and elsewhere in South America. Here are pio-
neer contributions to the field of plant-pathology, the papers on the Potato
Rot, American Grape Vine Mildew, Black Knot of Cherries, and Diseases
of Orange and Olive Trees in California representing part of his work at
the Bussey Institution under his appointment as Assistant Professor of
Cryptogamic Botany at Harvard from 1874 to 1879. There are also nu-
merous more general papers in this field discussing common diseases
caused by fungi, diseases of fruit-bearing trees, fungous diseases important
in Massachusetts in various years and injurious fungi of California, while
forest pathology shows a few pioneer papers on diseases of forest trees and
diseases of trees following mechanical injury, and there are numerous
studies of rusts, smuts, mildews, blights, and other fungi pathogenic to crop
plants. There is also a brief note, in 1881, on the identity of a fungus found
growing in the human ear, one of the early contributions in this country to
the field of medical mycology, destined for rapid development forty years
later. Papers on the edibility and poisonous quality of mushrooms and on
the diseases of mushrooms with notes on truffles and other rarities are evi-
dence of his interest in these economic aspects of the fleshy fungi as well
as in their identity, classification, and relationship. There are notes also on
some of the more spectacular aspects of mycology such as the identity of
the fungi stored up by squirrels, the fungous diseases of insects, the mim-
icry of fungi by insects, the famous ginger beer plant with its combination

WESTON: FARLOW’s INFLUENCE ON MycoLocy 89

of yeast and bacteria collaborating to produce a liquor more or less potable,
and the nature of the spots of rose leaves thought to be a fungous disease
but in reality merely the adherent sporangia explosively discharged from a
mold growing on the mulch below, a phenomenon that still continues to
mislead botanists of the present as it did those of Europe and the United
States in 1886.

His publications offer convincing evidence that even though his interests
were primarily in systematic botany in its broadest sense, yet he was never
oblivious of the practical importance of various aspects of cryptogamic
botany and when need arose, turned his capabilities to such problems with
telling effect, producing pioneer work of significance in several such fields.
In plant pathology his work stands with that of Burrill and other pioneers
as the first in this country, the beginning of a rapid development that
within two generations gave preéminence in this field to the United States.
Similarly his study of the pollution of drinking water by fresh water algae
was a classic, pioneer investigation paving the way for the later develop-
ment by Moore, Kellerman, and others, of this field of applied algology.

Even more than by his publications, however, he contributed to the
advancement of botany by the services his herbarium, library, and indices,
as well as his own knowledge, stimulus, and counsel rendered to workers in
many areas of that rapidly expanding field of science. Here was available
authentic reference material unequalled by any similar assembly in this
country, a source where under the authoritative and interpretive guidance
of Dr. Farlow, the interested seeker could find the answer to difficult ques-
tions as to the nature, identity, and relationship of the lower plants. Al-
though such service is hard to estimate in quantitative terms, yet it was
probably of the greatest value to mycology for while the library and her-
barium covered the whole field of cryptogamic botany, the herbarium
comprised more fungi than all the other specimens of algae, lichens, and
Bryophytes combined, while the two Indices were devoted to the fungi
alone. These facilities served, in the case of potentially harmful fungi, as
the fingerprint files and criminal records from which the identity and dan-
gerous characteristics of poisonous, injurious, or disease producing fungi
could be determined with certainty, and, in the case of fungi potentially
valuable to science, industry, or agriculture, as the personnel roster from
which their names, locations, and capabilities could be ascertained. Visit-
ing scientists in increasing numbers sought this center, coming from labora-
tories and institutions from all parts of this country and even from Europe,
while by correspondence and exchange or loan of specimens and other ma-
terial the service rendered by Dr. Farlow and the herbarium was extended
even more widely. The numerous published acknowledgments frequently
occurring in the significant contributions to botany over a period of nearly
fifty years eloquently witness the effectiveness of this aspect of his par-
ticipation in the development of mycology.

Dr. Farlow’s influence on cryptogamic botany and on its specialized
branch of mycology began through his teaching in 1874 when on returning

90 FarLow1iA, VoL. 2, 1945

from Europe he was appointed Assistant Professor of Cryptogamic Botany
at Harvard University and commenced work at the Bussey Institution in
Jamaica Plain. There, along with his investigations of plant diseases, he
guided the research of advanced students, the first of whom was Byron D.
Halsted, while two days a week he gave more elementary instruction in
general cryptogamic botany to undergraduates in Cambridge and in the
summer gave similar instruction in summer school. After five years of pro-
ductive activity in this dual role, he relinquished his work at the Bussey
and under his appointment as Professor of Cryptogamic Botany in 1879
devoted all his time to a full program of research, teaching, herbarium
work, and other activities in Cambridge. In 1891 he secured the appoint-
ment of Thaxter, one of his most promising Ph.D.’s then working at the
Connecticut Agricultural Experiment Station, to carry on the undergraduate
teaching while he travelled and worked in Europe and on his return in 1892
resumed only the instruction of graduate students working for higher de-
grees and the guidance of other investigators coming to the laboratories
from the United States and abroad to carry on advanced research under his
direction. In 1896, to devote himself untrammeled to the steadily growing
herbarium, library, and indices that had become ever more important and
widely known, he relinquished to Dr. Thaxter even this advanced instruc-
tion and guidance although his helpful and stimulating influence on gradu-
ate students through occasional contacts continued until shortly before his
death.

Thus through seventeen years of undergraduate courses and twenty-
three of official graduate instruction his influence as a teacher made itself
felt on a long list of men from students gaining chiefly general pre-medical
background to advanced workers who became important figures in various
fields of botany. Of the courses he gave it was the general survey of crypto-
gamic botany variously designated in successive catalogues that most effec-
tively demonstrated his unusual ability as a teacher. This covered the
structure, development, reproduction, relationships, classification, and 1m-
portance of the lower plants, a stimulating and challenging course with
characteristically enlightening and enlivening lectures by Dr. Farlow and
with strong emphasis on the laboratory study of representative types espe-
cially of the algae and fungi. He was at his best in the laboratory where
his practical and psychologically sound approach of leading on the student
by patient but persistent questioning used this plant material as a means
for developing the student’s latent capabilities for accurate observation,
effective interpretation and precise description. His methods of inducing
the student by his own efforts to master the difficulties of interpreting three
dimensional features of complicated solid structures through interpreting
the two dimensional visual images seen through the microscope were highly
effective. A modern teacher might wonder what sections of a block of pine
wood might have to do with cryptogamic botany or with mycology but as
applied by Dr. Farlow, the study of this complicated structure became a
highly challenging and enlightening initiation and apprenticeship to the

WESTON: FARLOW’S INFLUENCE ON MycoLocy 91

student with any spark of initiative or resourcefulness. Some.of Dr. Far-
low’s convictions on the basic philosophy of teaching, the importance of
the student’s seeking and obtaining his answers from the material itself
through his own efforts, are well brought out in his few articles on teaching,
especially in his paper in the Scientific Monthly of 1886. Here the inter-
ested reader will find a discerning portrayal of some of the “modern” meth-
ods of teaching biology with a keen analysis of the types of students he
finds in his own classes today and no teacher, no matter what school of
education may have given him his training, can fail to profit by reading
this paper of fifty years ago.

The facilities for this undergraduate teaching would now be considered
meager. The microscopes were the early type Leitz, the low power focussing
achieved by means of adroitly twisting the barrel in its friction collar, the
high power focussing accomplished by a very simple but effective lever
mechanism. Crude though they seem to the modern student, these little
microscopes are still capable of excellent work in the hands of a discrimi-
nating user and several of them, fifty years later, still in good working con-
dition, have given excellent service on field trips to present members of the
staff. As there was no botany building until 1890, his courses were held in
various buildings, one being Boylston Hall which still survives after years
of service as a chemical laboratory, as a library, and in other capacities
despite occasional arsonous attempts of students to do away with its gray
grimness. From Dr. Farlow’s class records it can be assumed that it was
even more cheerless then than now for entries, during the winter of 1879,
in his characteristic handwriting, record: “January 7: Lecture on Fucus
and Vaucheria. Laboratory too cold for work. January 8: Lecture on the
Peronosporeae. Laboratory too cold for work,” while on January 14 it is
safe to assume that milder weather enabled the class to make up for lost
time as there was no lecture and the laboratory successfully covered Fucus,
Cystopus, and the Saprolegniineae. Also, in these records there is a sug-
gestion of the locker trouble which still afflicts the biology teacher, for in
June, 1878, after the class of ten (six seniors, two juniors, one Divinity
School and one Lawrence Scientific student) had successfully completed
Natural History 9, he notes with relief that the microscopes (borrowed
from the Botanic Garden and the Bussey Institution) are all accounted for
but a new key will have to be made to replace the one missing for locker 16,
although fortunately key #9 can do double duty as with a little coaxing it
can be used for locker 8 in place of its own key, now lost. Incidentally it is
of interest to note that Alfred Worcester, who became a leading and dearly
loved physician in the vicinity, had locker 3 in 1877-78, Wm. Trelease,
later Director of the Missouri Botanic Garden, had locker 1 in 1880-81,
while in 1881-82, Wm. Patten, later widely known for his theory of the
evolution of higher vertebrates from the Ostracoderms had #4 and Roland
Thaxter #1. :

The general course in cryptogamic botany served primarily to give a
foundation knowledge of the lower plants to premedical and other students

92 FaRLowIA, VoL. 2, 1945

concentrating in biology, and it fulfilled this function most ably giving the
students also the training in observation and interpretation already noted
and stimulating them with the enthusiasm and ideals which were essential
ingredients of Dr. Farlow’s teaching. Some, however, such as Thaxter,
found in this course and in the more advanced ones to which it led,
the impetus in mycology as a life work, hence even this undergraduate
teaching contributed in some measure to Dr. Farlow’s influence on this
field.

It is for his guidance and training of more advanced workers that Dr.
Farlow was best known and therein lay the major influence of his teaching
on mycology. Stimulating in his leadership, exacting in his requirements,
unfailing in his extensive knowledge, he contributed notably to the develop-
ment of the men who carried on advanced work under his direction and
guidance. It is significant that the problems he suggested for these men
were almost without exception in the fields of morphology, histology,
cytology, development, or parasitism rather than taxonomy. This was be-
cause, from the breadth of his own experience, he realized that such work
would afford them better training than premature attempts at taxonomic
monographs of various groups. Also he realized that systematic botany in
its broad requirements needed all the evidence available from the rapidly
developing fields of morphology and physiology as these, in turn, were
dependent on taxonomy. In the testimonial volume of letters and photo-
graphs given him on his seventieth birthday, there is an impressive list of
some fifty-eight workers who had studied with him or benefited from his
guidance in the Cryptogamic Laboratory between 1874 and 1914. Among
them are some who had become outstanding figures in algology, others had
contributed notably to zodlogy, medicine, bacteriology, or even more spe-
cialized fields while the greater number were those who had played an im-
portant part in developing mycology and plant pathology in this country.
The names of these men are eloquent testimony to the influence of Dr.
Farlow’s teaching; here are the algologists Setchell and others, discussed
more fully elsewhere in this number; here are the outstanding teachers
such as Lyman, Butters, and Grover; the famous men of medicine such as
Kelly; the bacteriologists such as Barber; the botanists who contributed
notably to physiology, genetics, cytology, taxonomy or other branches,
men such as Blakeslee, Duggar, Davis, Robinson, Peirce, and Oakes Ames;
the plant pathologists and mycologists such as Clinton, Rorer, Johnston,
Faull, Schrenk, Fink, Riddle, Burt, Jackson, and Thaxter.

All the aspects of Dr. Farlow’s influence on mycology and indeed on the
whole field of cryptogamic botany whether through his teaching, research,
or herbarium activities derived power and enhanced effectiveness because
they were quickened and vitalized by his personality. In him there was no
dullness nor stolidity, rather he was dynamic, highly active, quick of mind
and motion, congenial, witty, and stimulating. Any discussion in which he
participated was activated through his participation, any meeting he at-
tended was enlivened and enlightened by his presence, any scientific move-

WESTON: FARLOW’s INFLUENCE ON MycoLocy ' 93

ment in which he took part was given impetus. Even as a student he had
shown the qualities of leadership and as his prestige grew through years of
constructive activity it was the stimulus of his personality as well as the
soundness of his judgment and the scope and accuracy of his knowledge
that made him one of the leading figures in American botany.

In the biographical sketches which appeared after his death, especially
in the excellent memoir by Setchell, are many revealing glimpses of Dr.
Farlow’s personality, while the spontaneity of his wit and the deftness of
his humor enliven the pages of his contributions. Some of the many stories
concerning him have already been written, numerous others are treasured
in legend. A couple of these, apocryphal perhaps, are yet so characteristic -
that they may not be out of place here. For example, it is said that his
method of informing Dr. Thaxter of his engagement to Lilian Horsford was
to remark at the end of one of their daily discussions of laboratory affairs,
“By the way, did you know that Miss Horsford is engaged?” and on
Thaxter’s astonished reply “No, to whom?” his answer was “To me, of
course.” Also, as an instance of his ability to deflate pomposity there is the
story that once, when he was crouched, digging in a patch of shrubbery by
the side of a road near Chocorua, a dowager driving by with all the pomp
and circumstance of footman, driver, and prancing team of horses, im-
periously stopped the equipage and called out “Little boy, who are you and
what are you doing here,” at which Dr. Farlow straightened up and with
impressive dignity answered, “Madam, I am Dr. William Gilson Farlow,
Professor of Cryptogamic Botany at Harvard University and I am collect-
ing hypogaeous fungi,’ at which disconcerting and unexpected reply the
woman is reported to have driven rapidly away.

Those of us who were fortunate enough to have had even a slight asso-
ciation with him as graduate students looked upon him with awe because
of his attainments, and delight because of his whimsical humour. While I
was in charge of the teaching laboratory as Dr. Thaxter’s assistant, Dr.
Farlow liked to use the large tables on free days for sorting specimens and
his comments, while rapidly shuffling packets from one herbarium folder to
another and critically looking over new accessions, were worth straining
the ear to hear and such straining was necessary, for in his rapid sorting,
both hands would be busy and he would frequently hold one or two packets
between his teeth. Yet even when thus occupied he would carry on a run-
ning fire of comment on the specimens, with anecdotes of those who had
collected them and reminiscences of his own collecting trips. It was on one
such occasion that he spoke of collecting algae on the West Coast and dry-
ing his specimens in his hotel room using the iron bed, which he had turned
upside down and festooned with cords, as an improvised drying rack. His
imitation of the Chinese houseboy opening the door, taking in the situation
with one glance, uttering a wailing ‘““A-wah, A-wah” and retiring for a more
favorable opportunity to tidy the room, was unforgettable.

In some of the biographical sketches his quick, light step is noted. This
step was familiar to all of us in the laboratories, as we noted also his char-

94 FarLow1A, VoL. 2, 1945

acteristic habit of taking a couple of shorter, quicker steps just before com-
ing to a threshold, then hurdling the threshold as it were and taking a few
more quick steps before resuming his regular pace. For this characteristic
entry I used to listen, for he had a way of trying out my reactions on ma-
terial he was studying, seeking, no doubt, the opinion of the general public
of less experience and lower I.Q. in addition to the expert opinion of Dr.
Thaxter. On one occasion he brought in two slides, asked me to compare
them and after watching me examine them carefully under high power
asked my opinion. My reply, noting their general similarity but pointing
out certain differences, seemed to please him highly and he rapidly departed
with the slides remarking ‘‘Very interesting, some people seem to have
thought they were the same.’’ When his note in Science in 1912 appeared
shortly after, I realized he had been showing me some of the critical ma-
terial on which his transfer of the chestnut blight fungus from Diaporthe
to Endothia was based.

It was typical of his geniality that he would invite all of us, graduate
students and investigators in the laboratory, to his house for occasional din-
ners. Those were memorable evenings. Mrs. Farlow, gracious and kindly,
presided at the table over a dinner excellent and sumptuous, in striking
contrast to the meager Foxcroft Hall menu which the limited finances of
most of us permitted. Then followed examination of priceless volumes in
the small fire-proof room which projected toward what is now the Faculty
Club; perhaps some music in which he showed himself a versatile and ac-
complished pianist; and certainly reminiscences and anecdotes of his own
experience and of well-known mycologists and algologists here and abroad,
unforgettable to us who were just beginning our mycological work. On one
occasion there was discussion of ant lions with which, of all of us, only
Owen Burger was familiar from his work in Florida and I remembered well
Dr. Farlow’s vivid demonstration, on the deep piled Oriental rug, of the
proper posture to assume while calling ‘““Doodle, doodle” to the ant lion in
the right pitch to jar him with the vibrations so that he would emerge from
his hiding place in the center of his effective sand trap. (Incidentally, some
years later in the Philippines I found this technique successful with the
ant lions under the eaves of my house at Los Bajos even though the magic
words are certainly not Tagalog! )

All of us working in the laboratory at that time received some measure
of help in our research and large measure of inspiration in our lives from
our contacts with Dr. Farlow and this was in 1911 to 1915, only a few
years before his death and long after he had, officially, relinquished all
direct contact with teaching. Still treasured in my collections are specimens
he gave me to facilitate identification of algae I had gathered at Woods
Hole and around Halifax, Nova Scotia; but one of many instances of his
helpfulness toward all of us.

That his aid to the younger generations of botanists was not only scien-
tific but in many cases financial is less well known, largely because such aid
was very unobtrusively and often vicariously and anonymously achieved.

WESTON: FARLOW’s INFLUENCE ON MycoLocy 95

Setchell mentions one or two such instances and Mangin gratefully acknowl-
edges other cases in which French botanists were benefited while many
others have received only private, though equally grateful, recognition.

To the beginner as well as to the expert, the assistance of Dr. Farlow’s
incomparably broad and expert knowledge was ever available if sought,
and as his prestige grew he occupied a position of esteem highly regarded
in this country and abroad as the final authority in difficult questions in
the taxonomy of the lower plants. Because of his wide contacts with the
work going on in this country and Europe, his familiarity with a wider
range of workers and laboratories, his grasp of important developments in
various fields, his opinion was sought not only in taxonomic problems but
also on questions of general policy and purpose. It was the combination of
his accomplishments and his personality that made him the spokesman and
representative of mycology for many years.

Finally, as an adjunct to all the other channels through which he influ-
enced cryptogamic botany and especially mycology, the part he played in
fostering and promoting publications should be noted. He was the first of
the American editors of the influential British periodical The Annals of
Botany and, through his efforts over several years, it was strengthened by
notable contributions from workers in the United States. In 1883, as the
volume of work accomplished by investigators in his laboratory began to
increase, he started the well-known series of “Contributions from the Cryp-
togamic Laboratories of Harvard University” and in 1896 when he relin-
quished the guidance of research workers to devote himself to his other
activities, the numbered publications in this series, together with a few
similar contributions for one reason or another not included, had reached
over forty. Each of these papers by students or associates was within its
limits a significant contribution and the whole series has materially ad-
vanced the development of botany. This very periodical, Farlowia, itself
is not only a tribute in name to Dr. Farlow, but concrete evidence of the
ways in which his influence on mycology and other branches of cryptogamic
botany still makes itself felt.

HARVARD UNIVERSITY
CAMBRIDGE, Mass.

4

2(1): 97-141 FARLOWIA January, 1945

THE BOLETINEAE OF FLORIDA WITH NOTES ON
EXTRALIMITAL SPECIES

I. THE STROBILOMYCETACEAE
-RoLFr SINGER

The present paper is intended to be the first of a series of monographic
studies devoted to the basidiomycetous flora of the State of Florida. For
the first time an attempt will be made to treat taxonomically the Boletineae
of a large region with due consideration of the anatomical and chemical
characters which, according to the author’s view, are of no less importance
in the taxonomy of this group than are adequate macroscopical descriptions,
Considering the many inevitable changes that will be noticed when this
paper is compared with the traditional American and European system of
classification, the author has thought it necessary not to confine himself to
the discussion of the species occurring in Florida. While giving complete
descriptions only of Floridan species, notes on extralimital species have
been added in order to avoid the mistake bound to impair the work of the
locally minded taxonomist. At the same time, this procedure seems to offer
the only means of providing all the evidence for the solutions proposed in
this paper. This seemed especially important in some thus far controversial
groups such as the Strobilomycetaceae, Jugasporaceae, and the genera
Pulveroboletus and Phylloporus (Boletaceae). It will also illustrate the
specific representation of Florida fungi in the world flora of Boletineae.
Another factor determining the present arrangement was the author’s wish
to coordinate European and American nomenclature and classification. As
a matter of fact, many so-called American species have been known previ-
ously to European mycologists under other names, and many European
species have turned out to be later synonyms of species published by Amer-
ican authors. In the case of Boletellus ananas and Strobilomyces pallescens
a similar situation has developed between this country and Australia.

The mycoflora of Florida is especially fascinating as a part of a transi-
tional vegetation, changing from the North American temperate flora into
the neotropic flora of the West Indies and South America. This transition
is a very gradual one, the northern and the southern elements overlapping
each other’s areas of distribution until, in a comparatively narrow zone
along the south tip of the peninsula it becomes more truly tropical. Never-
theless, while not all of the representatives of tropical South and Central
America reach South Florida, there are still some more northern elements
following the species of Quercus all through the peninsula. The extra-
tropical part of what is usually called South Florida, and Central Florida
up to a line running somewhere through North Florida is characterized by
a progressive (towards the north) increase of the absolute number of species

97

98 FARLOwIA, VoL. 2, 1945

of Boletineae observed, and also by the progressive increase of the percent-
age of northern elements, and a progressive division of the fungus flora into
a thermophilous summer flora, and a more hardy winter flora. While all
tropical elements reaching as far north as the zone where this bi-seasonal
fruiting period becomes most distinct naturally belong to the summer crop,
and all the truly northern elements belong to the flora appearing during the
winter rainy period, it nevertheless is true that some unexpected fruiting
periods are observed for certain species, e.g. Boletellus Russellii with the
center of its area far to the north, still appears among the summer-fruiting
species. The bi-seasonal division rather seems to follow generic lines: with
very few exceptions the genera Strobilomyces, Porphyrellus, Boletellus (i.e.
all Strobilomycetaceae), all Boletaceae except Suillus, and the genus Clito-
pilus belong to the summer-fruiting group, while Gomphidius, Suillus, Pax-
illus, and also some single exceptional species of the Boletaceae belong to
the winter-fruiting group. These seasons are strictly parallel with the main
periods of precipitation in this part of the state, ie. June to August, and
depending on yearly fluctuations, frequently also in May (even April) and
September (October) and again in December and January (sometimes in-
cluding November and February, or even a part of March). The periods
between these rainy seasons are not altogether devoid of fungus growth,
but only very exceptionally will any representatives of the Boletineae be
found during the dry spells between the main seasons, and never will a
typical summer species be collected in winter or vice versa. It will be re-
membered that a somewhat similar situation has been reported from Cuba
by Berkeley and Curtis though their statement regards only the montane
zone. A very interesting fact, noteworthy in this connection, is the discovery
by the writer of true seasonal dimorphism in boletineous species in Florida.
Two species, Suillus hirtellus (Peck) Snell and Suillus australis Sing. ined.,
show morphological and chemical differences according to the period of
appearance and these are constant in different regions and analogous in
both cases cited. They will be discussed in the second part of this paper.
As for the association of the mycorrhizal fungi with certain hosts, we
have some interesting examples of specialization in Florida. While it is
generally assumed that there is little evidence of a mycorrhizal association
with forest trees in the tropics, the occurrence of several species of pines
down to the south tip of the peninsula and in the Caribbean area, and also
the presence of many species of Quercus, explain the fact that Florida, even
the tropical zone of the State, makes an exception of this rule. It is remark-
able that two closely allied species, or, in the writer’s opinion, subspecies of
the genus Pinus, i.e. Pinus palustris and Pinus caribaea, clearly differ bio-
logically if their symbionts are considered. P. palustris has many more spe-
cies forming mycorrhiza obligatorily confined to Pinus, and most stands
and a fair number of individual trees if not practically all in certain regions
have either Boletinus-, Suillus-, or Gomphidius-mycorrhiza, or at least a
mycorrhizal flora consisting of such Agaricales that do not occur on or with
Pinus caribaea. This latter pine is comparatively rarely found to be asso-

SINGER: FLORIDA BOLETINEAE. I oo

ciated with mycorrhiza-forming Agaricales, and if it is, the species are dif-
ferent from these observed with Pinus palustris. Pinus caribaea is associated
with Boletus griseus, Boletus rubricitrinus, Boletus rubellus or in other
words, fungi belonging to a genus where the formation of mycorrhiza with
conifers is mot a generic character, and not even a specific character as far
as the species just cited are concerned. This strict differentiation may best
be explained by the fact that to the slight morphological difference between
these two pines there is added a somewhat stronger climatic, and a very
strong ecologic factor. Most of the area of Pinus caribaea is situated in the
tropical zone of South Florida, and the soil usually is hot and dry lime rock.
Whatever the explanation, this interesting relationship tends to furnish one
more argument in favor of the results of the recent investigations by De
Vall.1 For example, in Highlands Hammock State Park, Pinus caribaea
was indicated by some authors. The analysis of the accompanying fungus
flora shows that it is typical for Pinus palustris, not P. caribaea. Actually
De Vall and West were able to state that the pines in question were Pinus
palustris.

As for broad leaf trees the main mycorrhiza-bearing genus is Quercus.
Here again I was surprised to learn how little some of the mycorrhizal fungi
seem to be influenced by the variety of vegetation types in which they find
themselves growing, as long as there is a suitable mycorrhiza-host. In
Florida all species of Leccinum (group of Boletus scaber) are dryophilous,
i.e. confined to oak; yet, Leccinum chalybaeum Sing. ined. was found in a
flatwood under Pinus palustris. It was not until I found a patch of two
inch high Quercus minima that I felt satisfied to explain the occurrence of a
Leccinum in pine land. This little oak also grows in symbiosis with Por-
phyrellus subflavidus. Only a few feet from there, I observed several car-
pophores of T'wlopilus conicus, an indubitable pine-symbiont. This shows
how carefully field observations on questions of mycorrhiza must be made
in order to provide information usable as a basis of future experimental
studies or as a basis for taxonomic purposes, instead of being a source of
perpetual confusion.

Among the vegetation types of Florida in which species of Boletineae
participate, are all types of hammocks (low, mesophytic and high ham-
mocks as well as tropical hammocks), pinelands (high pine and flatwood,
and even the lime rock flats of the south) and scrub areas on white sand
in sand dune regions. This enumeration is given in the order of relative
importance. In open grassland (lawns, grassy places in open woods and
gardens, so-called prairies and marshes, possibly some parts of the ever-
glades), Boletineae are found exceptionally: Clitopilus Giovanellae and
C. scyphoides var. floridanus being the only species growing outside the
wooded areas, entirely independent of mycorrhiza, or as it seems even of
the cooling shadow of the trees. In the scrub, under scrub oak and Pinus

*De Vall, Wilbur B. The taxonomic status of Pinus caribaea Mor. Proc. Fla. Acad.
Sci. for 1940, 5: 121-132. 1941.

100 FARLowIA, VoL. 2, 1945

clausa, I have observed two species of boletes, Boletus luridellus and Xero-
comus subilludens, but both these species are not exclusively inhabitants of
the scrub.

A phenomenon of interest for those who search for new and more con-
vincing facts about the problem of the phylogeny of the Boletineae, has
been observed in Florida: the occurrence of gastroid conditions of two now
well-defined species of Boletineae, Boletus rubellus ssp. caribaeus and Bole-
tinus decipiens. In the former they were sterile while in the latter they
produced spores of the same kind as were produced on the pseudoangio-
carpus (i.e. normal) carpophores. These parallel forms of fructification
will be discussed at their proper place.

The number of species of Boletineae now known to occur in Florida is at
least sixty-five. This is a remarkably high figure if the fact is kept in mind
that several genera of prevalently or exclusively northern distribution such
as Gomphidius, Paxillus, Hygrophoropsis, and Boletinus are represented by
only one or two species each, and still the total is higher than in France,
and probably higher than in the entire Caribbean area.

Some special arrangements and abbreviations to be used on the following
pages need explanation. In the key, the Florida species are numbered, the
numbers referring to the figures preceding the name and description of the
species in this paper. The unnumbered species appearing in the keys are
those studied by the author and discussed in this paper under the heading
“Extralimital species.” For these species the synonymy has been kept down
to the most essential binomials, while for Florida species the full synonymy
has been established. The keys are thought to enable the experienced reader
to determine all sufficiently known species, and it is hoped that this arrange-
ment will make the paper useful beyond the state limits of Florida, and also
abroad. Under the heading “Material studied” we indicate all material that
has influenced the description given. The herbaria in which the material is
preserved are indicated by the abbreviations recommended by J. Lanjouw
(Chron. Botan. 5 (2/3): 143-150. 1939). A special effort has been made
to secure type material of the species treated, and all material collected in
Florida, though the latter is scarce with the only exception of the collections
preserved at the Agricultural Experiment Station of the University of
Florida at Gainesville. All the descriptions and notes including the syno-
nyms are entirely based on the author’s own studies; in the very few cases
where citations from other sources have been necessary, these sources have
been indicated. A complete list of literature and an index of the genera and
species will be given at the end of the last part of this paper which will
be published at a later date. The color names in quotation marks given in
the descriptions and keys are those of R. Ridgway. Color Standards and
Color Nomenclature, Washington, D. C., 1912. Colors that cannot ade-
quately be determined in Ridgway are cited from A. Maerz & M. Rea Paul,
Dictionary of Color, New York, 1930, and then ““M. & P.” is added after the
quotation marks.

SINGER: FLoRIDA BOLETINEAE. I 101

ACKNOWLEDGMENTS

A fellowship of the Guggenheim Memorial Foundation has made this
study possible. It enabled the writer to get a first-hand knowledge of the
fungi of Florida in the field, and also to carry through a careful examination
of the types of species described by W. A. Murrill at Gainesville, Florida.
The author wishes to thank Dr. D. H. Linder for loan of scientific appa-
ratus during the trip, for reading the manuscript and for the drawings on
plate I of this paper. He also wishes to acknowledge his gratitude for per-
mission to study material from other institutions than the Farlow Her-
barium, namely to Dr. John N. Couch and Dr. Alma Holland Beers of the
University of North Carolina, Dr. Homer D. House of New York State
Museum, Dr. Fred J. Seaver, New York Botanical Garden, Dr. Walter H.
Snell, Brown University, Mr. E. West and Dr. W. A. Murrill, University of
Florida. It is a pleasure to acknowledge the kind and efficient help the
author had from all botanists he met in Florida. He wants to extend his
thanks for innumerable courtesies to Erdman West, University of Florida,
for laboratory space and valuable advice to Dr. Williams, University of
Miami, for office and transportation facilities to Mr. Altvater, Highlands
Hammock State Park, now in the Armed Services, for determination of
hosts to Dr. Buswell, University of Miami, and Miss Lillian Arnold, Uni-
versity of Florida. The writer is also thankful for the help in planning the
trip offered by Dr. E. D. Merrill, Harvard University, and Dr. D. Fairchild,
Coconut Grove, Fla. Their competence in everything connected with plant
life in Florida made it possible for the author to be at the right time at the
right place, probably the most important thing for anyone who wants to
collect in Florida. It is impossible to mention all the names of those who
have turned in specimens of fungi, but their contributions are highly ap-
preciated.

SUBORDER BOLETINEAE
(Opat. 1836 ut fam.) Pilat 1926, em. Sing. 1936.

KEY TO THE FAMILIES

A. Spores never pale pink, mostly ornamented, or nearly black to black, or “warm
sepia” to “Sudan brown” or “hazel” in print, or extremely long (most of the spores
of a print over 20 uw in length), or with thick (thicker than 1 «) wall applanate or
provided with a germinative pore at the apex; hymenophore ordinarily (in Florida
always) tubulose; clamp connections none. Strobilomycetaceae

A. Spores either pale pink, or smooth (constantly and persistently so), if nearly or dis-
tinctly black, and if larger than 20 w, the hymenophore is lamellate; if thick walled,
the hymenophore usually lamellate and/or clamps present; germinative pore none;
hymenophore either lamellate or tubulose, also merulioid, or showing some configu-
ration intermediate between lamellae and tubes; clamp connections absent or present.
B. Hymenophore tubulose, rarely merulioid or (in one species) Jamellate, and then

the surface of the pileus turning deep blue or greenish blue with ammonia.
Boletaceae

102 Fartowla, Vo. 2, 1945

B. Hymenophore lamellate. Pileus not turning deep blue nor greenish blue with
ammonia.

C. Spores evenly rounded, not angular when seen from one end, nor furrowed;
carpophore usually well pigmented, mostly strongly reacting with NHs, KOH,
FeSQ,, and other reagents; lamellae sometimes very thick and distant; clamp
connections absent or present; cystidia absent or present.

D. Clamp connections absent; spore print black or nearly so; spores clearly
elongate. Gom phidiaceae

D. Clamp connections present; spore print brownish to “chamois,” or creamy
white; spores not elongate. Paxillaceae

C. Spores not evenly rounded but angular when seen from one end, and often
furrowed longitudinally; pigment none, or grey, not reacting strongly with
NH:, KOH, FeSO,; lamellae not thick, not distant; clamp connections none;
cystidia none. Jugasporaceae

STROBILOMYCETACEAE Gilb. 1931 (as Strobilomyceteae)

Characters of the family: pileus squamose to squarrose, or squamulose
to glabrous, viscid, or not viscid, small to large, the margin frequently pro-
jecting; hymenophore tubulose, rarely lamellate, tubes usually compara-
tively long halfway between the margin and the stipe, strongly convex be-
neath, depressed to adnate, pores medium wide to rather wide when mature,
discolorous or more frequently concolorous with the tubes: whitish to gray,
or whitish to pale grayish cream color, becoming vinaceous pink to sordid
purplish, or yellow, golden yellow, the pores sometimes orange to red, at
last olivaceous or yellowish brown, sometimes becoming black or red or blue
on pressure; spore print black or nearly so, or, if not black, a very deep
brown with an olive tinge, or deep porphyry brown to reddish brown
(‘warm sepia,” “hazel,” “Sudan brown’’), rarely (in some olive brown col-
ors) approaching spore colors observed in Boletaceae, and then the spores
differing microscopically from the latter; spores richly colored when quite
mature, fuscous to deep succineous-melleous with darker ornamentations,
more rarely persistently smooth, and then either remarkable for their unique
size (20-30 u), or for their comparatively thick walls (more than 1 »), the
endosporium usually interrupted at the apex forming an incomplete germi-
native pore, or the apex applanate as in Ganoderma, or even more acumi-
nate than it is in the Boletaceae, outline globose, subglobose, short-ellipsoid,
fusoid, rarely ovoid-fusoid; basidia and cystidia comparatively voluminous,
mostly more so than in species with equally large spores in the Boletaceae;
trama of the tube walls pronouncedly bilateral in youth, hyphae without
clamp connections; stipe usually approximately cylindrical and compara-
tively long and slender, smooth or ornamented with woolly or fibrillose
scales, or with shallow to conspicuously projecting alveolate networks
which in the most typical cases are waxy-slippery and bearing a hymenium;
veil often present and then woolly or membranaceous, sometimes viscid,
either fugacious, or persistent as an annulus on the stipe or as an appendicu-
late margin on the pileus, or both, never entirely glutinous, not yellow-
pulverulent; stipe always destitute of glandulae. Habitat: on earth, more

SINGER: FLORIDA BOLETINEAE. I 103

rarely on the base of trees, or on decaying wood, most species preferring
warmer climates, only few species reaching Canada, and only two species
known from Europe and adjacent regions.

When any local flora is analyzed, and the representatives of the Strobilo-
mycetaceae compared with each other and with the Boletaceae, it usually
so happens that they are not easily recognized as a homogeneous group. In
Europe, the spores of the two species found there are very different, espe-
cially when studied superficially, and the classification of the boletes accord-
ing to the macroscopical characters of the pileus and the stipe that had
been usual in both the American and European literature up to very
recently, deepened the impression that the North American Strobilomy-
cetaceae were widely different macroscopically, and only the “incidental”
characters of the spores simulated an allegedly non-existing relationship.
Even in South Carolina with its comparatively rich flora, Coker and Beers
(The Boletaceae of North Carolina, p. 74. 1943) offer the opinion that “‘to
attempt to segregate them as a unified group would seem to us unadvisable.”
Strangely enough, the strobilomycetaceous species of Boletus sensu lato in
the same book are arranged much like a unified group. They are treated
in the following order: B. chrysenteron sensu Coker & Beers, B. sordidus,
B. chrysenteroides, B. ananas, B. Russellii, B. betula; — missing only B.
gracilis (with the spores erroneously described as smooth), B. projectellus
(with actually smooth spores), and S. floccopus (already transferred to
Strobilomyces). If all characters and all species known (including those
from Asia and Africa) are considered, there is no doubt that the Strobilo-
mycetaceae constitute a natural group though related to the Boletaceae,
but not showing clear kinship with any genus of the latter, differing not
only in the finer structural characters of the spores, but also in their color.
They also show a combination of macroscopical characters that makes it
possible for the experienced collector to foretell the spore characters in a
general way. The relatively large size of the basidia and cystidia in most
species is an additional criterium. The astonishing similarity of the chem-
ical characters in Porphyrellus pseudoscaber (Secr.) Sing. and Strobilomyces
floccopus convinced the author still more that the two species and genera
are more closely related than formerly has been suspected. Nevertheless
the gaps between some of the species known to belong to the Strobilomyce-
taceae may seem rather wide, and some striking types may, rightly or
wrongly, cause the taxonomist to erect several separate genera. That is
why, indeed, a number of genera already have been proposed for them:
Strobilomyces Berk. for S. fleccopus and allied species, Porphyrellus for B.
pseudoscaber, Boletogaster for a form near B. jalapensis, Frostiella for B.
Russellii and B. betula, and Boletellus for B. ananas.

However, if all the groups and species from all over the world are taken
into consideration, the gaps between them do not appear to be so striking,
and one has difficulty in distributing the species in genera or sections rather
than in finding connecting links between them.

In Gilbert’s scheme the Strobilomycetaceae are divided in two genera

104 FARLOwIA, VoL. 2, 1945

according to the color of the pores which are whitish to gray in Strobilo-
myces and yellow in Boletellus. In Leccinum where a similar situation
exists, this character has certainly no more than sectional value, and neither
in Leccinum nor in the Strobilomycetaceae is it accompanied by any one
correlated character. As a matter of fact, the squamose condition of the
pileus in Strobilomyces is found in the type species of Boletellus as well as
in some other species and the reticulate short spores known in S. floccopus
are also found in Boletellus retisporus, a species characterized by yellow
pores. Thus, the characters on which Berkeley based his genus Strobilo-
myces are not confined to this genus. On the other hand two African species,
S. costatisporus and S. pterosporus, have all the essential characters of S.
floccopus but short ellipsoid spores with the same longitudinal wing-like
ridges as known in Boletellus singaporensis or B. Linderi where the pores
are yellow or red and the veil of a quite different structure.

Likewise the color of the spore print, often useful in the classification of
the Boletaceae, cannot be used to the same extent in the Strobilomycetaceae
because of the uncorrelated occurrence of some shades and tinges. The
spores of Strobilomyces floccopus and Boletellus ananas are practically
black in print though very different in ornamentation and shape. Porphy-
rosporus subflavidus and Boletellus betula, otherwise similar, have the
hymenophore, including the pores, and the spore print differently colored.
Porphyrellus subflavidus and P. pseudoscaber though having the same color
of the spore print, are chemically different. The species, intermediate be-
tween these, Porphyrellus gracilis, has the spore print of a shade differing
from both P. subflavidus and P. pseudoscaber.

The situation is not even as simple as it seemed at first, since further
studies revealed that there are not merely white, gray, and yellow-pored
species, as Gilbert thought, but there also are species with initially pallid
cream-colored pores (e.g. Porphyrellus tristis), species with white pores
that become vinaceous pink (Porphyrellus gracilis, P. malaccensis, P. sub-
flavidus) and species with discolorous pores that are tinged orange to red
(Boletellus Linderi, B. pictiformis).

The spores are not only of two types, as was assumed formerly, but
there is a third type, usually described as punctate or warty; in this case
cylindric spines are imbedded in the exosporium, protrude slightly or
scarcely, and when they are seen from above, they are described as punctate
(Boletellus betula), but when they are seen in an optical section showing
the outline, the spores are mostly described as smooth (Porphyrellus gra-
cilis, P. subflavidus), or warty-rough (Porphyrellus malaccensis). In some
cases species with the essential macro- and microcharacters of a related
rough-spored strobilomycetaceous species, have not seemingly but actually
smooth spores. These smooth spores are easily recognized as strobilomy-
cetaceous by their size, color, or thickness of the walls, or by the shape of
the apex of the spore, and the combined remaining characters put these
forms in the neighborhood of some indubitable representative of the Stro-
bilomycetaceae rather than of the Boletaceae where most smooth-spored

SINGER: FLorRIpA BOLETINEAE. I 105

species belong. If it could be doubted hitherto that there are any smooth-
spored species actually related to some species of Strobilomycetaceae, there
is no longer room for doubt in view of the fact that both smooth and per-
forated spores are found in a single preparation of some specimens of Por-
phyrellus pseudoscaber and Strobilomyces velutipes. In both cases the
majority of the spores is smooth even when mature, and the minority is
ornamented.

The above-cited facts show that the Strobilomycetaceae are a homo-
geneous and probably natural family within the Boletineae, and have to be
treated as such. The difficulty in achieving a satisfactory division of the
family into genera could be avoided by recognizing only one genus for all
tube-bearing Strobilomycetaceae. However, this would mean only a
delay in solving the problem, for poor genera will never constitute
good sub-genera, and a single genus for all species involved would not
match well with the generic conception now generally adopted in the
Boletineae.

As the author thinks now, the difficulty will be solved most naturally by
adopting a third genus for the tube-bearing Strobilomycetaceae, besides
Strobilomyces and Boletellus. This genus has to take care of the species
with elongate, smooth or perforated spores that are somehow reddish brown
without an olivaceous tinge. Since it would therefore, among others, em-
brace the Porphyrellus pseudoscaber-complex on which the genus Porphy-
rellus has been based, it would not be necessary to propose a new generic
name, but merely transfer Porphyrellus from the Boletaceae to the Strobilo-
mycetaceae. Gilbert, not satisfied with the naming of a genus to replace
Bataille’s homonym Phaeoporus, transferred several species to Porphyrellus,
American species which he did not know personally. Here started a con-
fusion which could only be straightened out by studies in America using
fresh spore prints and good fresh and authentic dried material. It turned
out that Boletus alboater Schw. and Boletus nigrellus Peck, synonyms by
the way, are not related to Porphyrosporus pseudoscaber, as claimed by
Gilbert, but belong in a section of Tylopilus Karst. (Boletaceae) , not repre-
sented in Europe. This explains why Gilbert, not knowing any species of
this section, and not having recognized the genus T ylopilus at all, misplaced
the two American species in his Porphyrellus. Porphyrellus differs from
the above-mentioned section of Tylopilus in having more strongly pig-
mented spores with a different color in print, and with thicker walls and
either acuminate or truncate apex. It also differs in having the pores much
wider in age than they are in Tylopilus, and finally, their hymenial elements
are decidedly more voluminous, especially in proportion to the spore size.
The chemical reactions also support this distinction. Though both in 7'y-
lopilus and in Porphyrellus there are species with a pileus covered by a
trichodermium, the structure of this layer on the stipe is not quite identical,
and the trama of the dissepiments of the hymenophore becomes subregular
in Porphyrellus in a much earlier stage than in Tylopilus. The occurrence
of spores with imbedded warts in some forms of Porphyrellus pseudoscaber

106 FartowlA, Vor. 2, 1945

emphasizes the difference between the type species of Porphyrellus and the
genus Tylopilus with its constantly smooth spores.

Porphyrellus pseudoscaber is closer to Strobilomyces than it is to Tylo-
pilus. This is shown by the occasional presence of spores with imbedded
warts as in Strobilomyces velutipes, by the above-mentioned similarity of
the chemical characters of P. pseudoscaber and S. floccopus, the intensely
colored spore wall in both Porphyrellus and Strobilomyces and the rather
large size of the basidia when compared with spore size in both these genera.
It is therefore surprising to see that Fries, who did not know any of these
characters, by mere intuition treated Boletus porphyrosporus (i.e. Porphy-
rellus pseudoscaber) and Boletus strobilaceus (i.e. Strobilomyces floccopus)
within one and the same subgroup, the section Favosi Fr. Bataille had
Eriocorys strobilaceus (S. floccopus) and Phaeoporus porphyrosporus (P.
pseudoscaber) in the same series Porphyrospori Bat.

There are now about forty-five species known to belong or probably be-
longing in the Strobilomycetaceae, of which roughly two thirds are suffi-
ciently studied. Only about one quarter of the total is not prevalently or
exclusively tropical or subtropical. Eight species and varieties have been
observed in Florida. Three of these are endemic and have not thus far been
found except in Florida. This puts Florida among the regions richest in
strobilomycetaceous forms: the southern part of North America, the West
African tropics, and the Austral-Asiatic tropics.

KEY TO THE GENERA AND SECTIONS

A. Hymenophore, at least in its larger part, distinctly tubulose.

B. Spores globose to short-ellipsoid; hymenophore white to gray at first, becoming
darker in age; stipe not waxy-costate-lacunose; pileus and stipe either warty, or
woolly, or spinose-squarrose. Gen. Strobilomyces Berk.

C. Spores smooth, at least a considerable percentage of them, or beset with warts
and short ribs, or either of these, imbedded in the exosporium, or with more
or less imbedded to subfree reticulate ridges, forming a complete network on
the spores. Sect. Genuini Sing.

C. Spores with longitudinal wing-like thin ridges running from the hilar end to-
ward the very apex of the spore where they abruptly stop.
Sect. Pterospori Sing.

B. Fungi not combining the characters outlined above, ie. spores either distinctly
elongate (Q larger than 1.5), or hymenophore not white, nor gray at first; if the
pileus is adorned by projecting warts, floccons, or erect spines, the hymenophore
is yellow to red, and the stipe is not strongly floccose, or a veil is entirely want-
ing, or both.

D. Tubes and pores of the young carpophores white to pale grayish cream color,
becoming light pinkish vinaceous or sordid gray to porphyry brown with a
grayish tinge when quite mature; spores with short cylindric warts or with
slender spines perforating the exosporium, making the spores appear punctate
when their upper surface is focussed upon under an oil immersion lens; in
some cases the vast majority, or even all spores are virtually smooth; spore
print reddish brown (varying from “warm sepia” to “Sudan brown” or
“hazel”) without any olivaceous tinge (at least in the species studied in this
regard). Gen. Porphyrellus Gilb. em.

SINGER: FLoRIDA BOLETINEAE. I 107

E. Pileus sordid or at least very dull-colored, gray or umber. Context
strongly reacting with KOH, NH.OH, FeSO, etc.; stipe not reticulate, its
trichodermium usually forming a distinct velutinous layer.

Sect. Tristes Sing.

E. Pileus brighter-colored: yellowish brown, light yellow to occasionally
white, chestnut, or some other shade of reddish brown; context either
weakly or not at all reacting with KOH, NH.OH, FeSO,, etc.; stipe at
least finely or even obscurely reticulate, or partly so, or deeply waxy-
costate-lacunose. Sect. Graciles Sing.

D. Tubes yellow in youth, becoming melleous or brownish melleous, or oliva-
ceous with age; spores with an ornamentation as described above, or smooth,
or—most frequently —longitudinally winged to striately ribbed, in one
species reticulate as in Strobilomyces floccopus; spore print deep brown with
an olivaceous tinge to blackish, not reddish when fresh.

Gen. Boletellus Murr. em.

F. Pileus pink, purple, or carmine, but frequently at least partly expallent,
not viscid; spores ornamented with longitudinal wing-like ridges or striate
with narrow ribs or veins from the hilar end to the tip, never smooth, nor
otherwise ornamented; length of the majority of the spores when fully
mature more than 17 wu. Sect. Ananae Sing.

F. Pileus rarely showing the above colors, and then either viscid, or (and)
spores smooth, or differently ornamented, or shorter.

G. Spores either smooth and very large (length 20 » and above), or short
(around 12 ) and reticulate; apex of the stipe with a rather indistinct
to very distinct reticulation, or most of the stipe with longitudinal
veins that anastomose, but never entirely smooth, nor with waxy-
costate-lacunose surface in the species known.

H. Spores strongly elongate, smooth; margin decidedly projecting.
Sect. Mirabiles Sing.

H. Spores short, reticulate; margin not distinctly projecting.
Sect. Retispori Sing.

G. Spores either smaller than 19.5 « if smooth, or, if they are not smooth,
never reticulate; stipe either waxy-costate-lacunose, or smooth, or
woolly-shaggy-subreticulate.

I. Neither pileus nor stipe viscid when wet, and veil, if present, not
viscid. Sect. Chrysenteroidei Sing.

I. Either pileus or stipe or veil (if present) viscid, or all three viscid.
K. Spores with longitudinal wing-like ridges.
L. Stipe smooth or almost so, sometimes annulate; pileus not
velutinous-tomentose. Sect. Ixocephali Sing.

L. Stipe waxy-costate-lacunose, the thin lamellose ribs very high,
limiting deep honeycombs where rain-water remains stored;
pileus tomentose (at least in the American species), and not
distinctly (if at all) viscid. Sect. Dictyopodes Sing.

K. Spores with short cylindric warts which for the most part are
imbedded in the exosporium, and when the upper surface of the
spores is focussed upon, it appears punctate.

Sect. Allos pori Sing.

A. Hymenophore lamellate. Gen. Phyllobolites Sing.

108 Fartowia, VoL. 2, 1945

Strobilomyces Berk., Decades of Fungi 32-33. Hook. Journ. Bot. 3: 77.
1851.
Eriocorys Quél., Ench. Fung., p. 163. 1886.

Characters of the genus: see key (p. 106). Type species Strobilomyces
strobilaceus (Scop. ex Fr.) Berk., a synonym of S. floccopus (Vahl in FI.
Dan. ex Fr.) Sacc.

Sect. Genuini Sing. sect. nov.
Sporis levibus, perforato-verrucosis, cristulatis, reticulatisve.

Characters of the section: see key (p. 106) and the above Latin diag-
nosis.

KEY TO THE SPECIES

A. Spores smooth or with short perforating warts in the episporium, making the spores
appear punctate when seen from above. S. velutipes

A. Spores not smooth (only a small minority occasionally smooth, and many immature
spores smooth, but then the ornamented ones not as above), and not punctate when
seen from above.

B. Spores with short ridges and occasional isolated warts, incompletely connected
with each other to form a fragmentary network; pileus with a covering of rigid,
very acute, erect, thin spines or pyramids which often are cristate, or fimbriate,
or fasciculate, the single ones frequently applanate in one direction, and never
with a base wider than 1 mm. along its smallest diameter, and 2 mm. along its
largest diameter in dried condition. 1. S. confusus

B. Spores with a complete network; pileus with less rigid and more woolly scales
with a larger basal diameter when dried, or with the same kind of warts as de-
scribed above (the scales rather smaller than those), but at the same time with
merely squamulose marginal half.

C. Warts or spines or scales of the pileus larger than 2 x 1 mm. at their base, and

not very rigid. 2. S. floccopus
C. Warts acute, and their base smaller than 2 x 1 mm., rigid, confined to the
disk; the marginal half merely squamulose. S. nigricans

DESCRIPTIONS OF THE SPECIES OBSERVED IN FLORIDA

1. Strobilomyces confusus Sing. spec. nov.
Pratt I, Fic. 1

Pileo atro, spinulis erectis, rigidis, acutissimis, subinde fimbriatis vel cristatis
obtecto, spinulis densioribus in centro, margine velo appendiculato, sed haud distincte
nec persistenter lanato, sicco, 40-85 mm. lato; tubulis porisque albido-griseolis vel
griseis, siccando nigrescentibus, cinnamomeo-maculatis ubi tacti sunt, at maculis eis
gradatim nigrescentibus, 10 mm. longis vel longioribus, plerumque subdepressis ad
stipitem; sporis in cumulo nigris, cristulatis et incomplete reticulatis, 10.5-12.5 x 9.7—
10.2 uw; basidiis 37-48 x 11-13.5 uw; cystidiis clavatis, dein fusoideo-mucronatis vel
vesiculosis apice ampullaceo, fuscescentibus, (22)-34-61 x 7.5-26.5 mw; stipite griseo, ad
apicem reticulato, lanato-lacerato infra zonam reticulatam, plerumque attenuato basin
versus, ad ipsam basin saepe incrassato, solido, 42-78 x 10-20 mm.; velo pro ratione
sparso; carne pilei alba, celerrime rubescente, in stipite alba, autoxydatione et rubente
et nigrescente; odore haud notabili. Habitatio: in pratulis umbrosis, in hortis, in
pinetis mixtis prope Quercus minimas, in dumetis frondosis sub Quercubus laurifoliis
et Q. virginianis. Secundum litus Atlanticum Septentrionale occidentemque versus
usque ad Ohio et Tennessee.

SINGER: FLoRIDA BOLETINEAE. I 109

Pileus between fuscous black and black, with thin but rigid, not woolly,
very acute, erect spines which are denser toward the center, and often
cristate, fimbriate, or fasciculate, also not rarely compressed, the basal
diameter not over 2 x 1 mm. in dried material, occasionally the spines oblit-
erated or evanescent, and then the pileus appearing areolate-squamose and
not much different from similar forms of S. floccopus, never viscid, not
woolly, or at least not strongly and persistently so because of the greater
coherence and scarcity of the veil which often hangs in small patches from
the margin, surface convex, the center flattened when old, 30-85 mm.
broad. — Hymenophore whitish-gray to gray, becoming blackish even if
carefully dried, becoming spotted with “cinnamon” where touched in fresh
condition, these cinnamon places then changing to deep chocolate and
finally black; tubes 10-18 mm. long, very slightly depressed around the
stipe, but also adnate-subdecurrent in smaller, and decidedly depressed in
larger individuals; pores medium wide, about as in S. floccopus; spore print
black. — Stipe gray, reticulate at the apex above the annulus (or remnants
of the veil), shaggy-woolly below it, solid, tapering downward or more
rarely equal, the base often somewhat thickened, 42—78 x 10-20 mm.,; veil
not abundant, and annulus not persistent. — Context whitish, quickly red-
dening in the pileus reaching a deep carrot color except for a narrow zone
under the cuticle, the context of the stipe white, reddening and blackening
simultaneously on exposure; odor not remarkable.

Spores short-ellipsoid to globose, fuscous, with very fragmentary dense
network, or merely echinate, cristulate with occasionally connected spines,
the ornamentation 0.3-1.7 » high but usually partly or almost entirely im-
bedded in the much paler exosporium, wall rather thick with an incomplete
or complete germinative pore, 10.5-12.5 x 9.7-10.2 yw; basidia (3)-4-
spored, 37-48 x 11-13.5 »; cystidia clavate, then becoming fusoid-mucro-
nate and brown, also sometimes clavate-mucronate, or vesiculose with am-
pullaceous apex (neck), (22)—34-61 x 7.5-26.5 «; étrama of the dissepi-
ments of the hymenophore with a colored, denser mediostratum and a
paler lateral stratum of divergent hyphae, hyphae without clamp-connec-
tions; cuticle (tips of the warts) of strands of parallel hyphae which are
brown, 17.5—63 x 3-20 ,, with transverse outgrowths, horns, buttons etc.,
the terminal members sometimes clavate.

Chemical reactions: not separately studied (hardly much different from
those of S. floccopus).

Habitat: On shaded grass lawns in gardens, in mixed pine woods (e.g.
woods of Pinus palustris and Quercus minima), and also in high and meso-
phytic hammocks under hardwood trees (e.g. Quercus laurifolia and Q.
virginiana). Summer.

Distribution: From Massachusetts west to Ohio and Tennessee south to
central Florida. In Florida not rare though not quite so common as S. floc-
copus which seems to go farther north as well as farther south.

Material studied: Fla.: Eustis, Lake Co., Oct. 1897, Thaxter 6177 (immature) (FH) ;
Flatwood east of Gainesville, Alachua Co., July 1943, Singer F 2782, type (FH) ; other

110 FartowlA, Vo. 2, 1945

collections in and near Gainesville, Singer F 2766 I, F 2531, F 2647, co-types (FH).
Mass. Wood’s Hole, Farlow (FH) ; Plymouth, Aug. 1889, Farlow (FH).— Pa.: Martic
Forge, Marticville, on humus in chestnut orchard, Aug. 1910, J. F. Collins (FH).—
Ohio: Symmes, Hamilton Co., July 1935, W. B. Cooke 41452. (FLAS, FH).— Tenn.:
Cades Cove, Blount Co., July 1938, J. P. Porter, mixed collection (FH).

This species, so strikingly different microscopically, has never been dis-
tinguished from S. floccopus (S. strobilaceus) in this country. Both these
species apparently require about the same conditions for their develop-
ment since many collections in the herbaria are mixed. Once the distin-
guishing features are understood, they are so pronounced that it is possible
to foretell what type of spores a certain dried specimen will have, and vice
versa. Allowance, however, should be made for specimens which under
certain conditions, do not develop or preserve the characteristic acute
spines on the pileus. These forms are, fortunately, rather rare. On the
other hand, the ornamentation of the spores is quite constant. Among pub-
lished photographs Krieger, Popular Guide ... , p. 444, fig. 125, phot.
Cummins, may well belong here, while Atkinson, Mushrooms, fig. 172-174
certainly is S. floccopus.

2. Strobilomyces floccopus (Vahl in Fl. Dan. ex Fr.) Karst., Bidr. Finl. Nat. Folk
37: 16. 1882.

Boletus floccopus Vahl in Fl. Dan. ex Fr., Syst. Myc. 1: 292. 1821; Persoon,
Myc. Eur. 2: 144. 1825. non Rostkov.

Boletus gossypinus Pers., Myc. Eur. 2: 144. 1825.

Boletus squarrosus Pers., Myc. Eur. 2: 145. 1825,

Boletus cinereus Pers., Myc. Eur. 2: 144. 1825.

Boletus strobilaceus Scop. ex Fr., Elench. 1: 127. 1828.

Boletus stygius Wallr., Comp. Fl. Germ. Sect. II, 4: 608. 1833.

Boletus strobiloides Krombh., Nat. Abb. 10: 21. pl. 74, fig. 12-13. 1846.

Strobilomyces strobilaceus Berk., Dec. Fung. 32-33, Hook. Journ. Bot. 3: 77.
1851; Outl. Brit. Fungol., p. 236. 1860.

Boletus lepiota Vent., Micet. Agr. Bresc., p. 37. pl. 43. 1863.

Boletus hydriensis (Hacquet ut Clathrus ex) W. Voss, Oesterr. Bot. Zeitschr.
32: 40. 1882.

Eriocorys strobilacea Quél., Enchir., p. 163. 1886.

Eriocorys strobilacea var. floccopus Bat. Bolets, p. 11. 1908.

Prater I, Fic. 2.

Pileus between “mummy brown” and black, or about “sepia,” or more
gray all over or particularly near the margin, the rimosely diffracted por-
tions of the pileus between the scales either much paler, almost white, or
even darker (black) if the discoloration of the context has reached the
surface, paler on the margin because of the abundance there of the woolly
gray or white or sordid veil that hangs in soft patches from the margin and
initially continues into the corresponding layer of the stipe, the scales often
with blackened tips, softer than in S. confusus, entirely covering the surface
of the pileus, not viscid, large (larger than 2 x 1 mm. at their base in dried
specimens), erect and pyramidal, often curved, or imbricate and more or less
appressed, hardly denser toward the center of the pileus, surface pulvinate,
finally flattened, 44-155 mm. broad. — Hymenophore gray, or white, be-

SINGER: FLorIDA BOLETINEAE. I 111

coming gray, the pores concolorous but reddening or blackening on pressure,
or reddening and subsequently blackening, angular, usually lamellose
around the stipe when mature, medium to wide, tubes 12—14 mm. long or
occasionally longer, adnate to somewhat decurrent, or slightly depressed
around the stipe; spore print black. — Stipe gray to concolorous with the
pileus, and with gray or white or sordid abundant veil at least in the middle
portion of the stipe where it forms a thick woolly covering, sometimes
several subparallel belts up to a usually amorphous annulus, the apex sub-
glabrous, striate with longitudinal veins running down from the walls of
the tubes, the base tomentose, solid, subcylindric, or tapering upward, or
downward, 50-125 x 12-21 mm.; veil sometimes consisting of two distinct
layers, the outer one more colored. — Context white or whitish in the pileus,
either merely blackening, or reddening while blackening, or reddening and
then blackening, these discolorations sometimes blending into a dirty pink
or fuliginous-lilac shades, the same colors and color changes taking place in
the stipe though the discolorations there quite frequently in contrast with
the context of the pileus, rather dry, especially in the stipe when compared
with some other species of Strobilomycetaceae and especially with Boleta-
ceae, and slightly tougher than these; odor not remarkable; taste mild, nutty.

Spores 9.7-15 x 8.5-12 , covered entirely by a constantly complete net-
work that raises 0.3-1.7 » high, but frequently almost entirely imbedded,
the wall rather thick, with a more or less distinct germinative pore, asym-
metrical, buff brown with deeper brown ornamentation; basidia 30-58 x
(7.5)—-10-18 yp, 4-spored; cystidia numerous, 17—90 x 8-26 yp, clavate, very
frequently mucronate-apiculate, hyaline, occasionally containing a brown
drop when very young, later brown throughout, few cystidia more lanceolate
or fusoid, but never really acute above; trama of the dissepiments of the
tubes consisting of a denser, brownish mediostratum, and a looser, hyaline
divergent lateral stratum, hyphae of the mediostratum cylindric (few ve-
siculose), parallel-subinterwoven, clampless; spines of the pileus made up
of strands of parallel hyphae with clampless septa at a distance of (20)—
40-88 yp, cylindric or slightly narrowed at the septa, brown, the terminal
members often slightly attenuate toward the apex, but always rounded,
7-17.5 p thick.

Chemical reactions. KOH on surface of pileus brown; on context red
then usually becoming brown or brownish, the red reaction usually bright
and distinct, rarely dull because of the beginning blackening due to autoxy-
dation; NH,OH on context ochraceous, or (in Dade Co. specimens) nil;
FeSO, bluish gray, bluish-greenish-gray, or glaucous; HNO, on context
yellow.

Habitat: In mixed woods, more frequently under frondose trees than
under conifers, in Florida frequently on well-shaded lawns, in gardens, also
in mesophytic, high, and tropical hammocks, usually under oaks. Summer
and fall.

Distribution: With certainty from Spain to the Caucasus, and from
Sweden to North Africa in the Old World, and from tropical Florida to

112 FartowiA, VoL. 2, 1945

Canada, apparently west to the Pacific Coast, also observed in eastern Asia
(e.g. Japan); its occurrence near the Equator and southward is not quite
certain. In Florida probably in all counties.

Material studied: Fla.: Dade Co., Fairchild Tropical Garden under Quercus vir gini-
ana Sept. 1942, Singer F 750 (FH); Brickell’s Hammock under Q. virginiana, November
1942, Singer F 931 (FH); Alachua Co., numerous collections in 1943, Singer F 2766,
2647a, 2194, 2752, 2020a, 2020b, 2606, 2899, 2532, 2736/II (all FH), some Murvill
(FLAS).— Canada: Petawawa Forest Experiment Sta. C. G. Ridley 9531 (FH).—
Me.: Kittery Point, Aug. 1897, Thaxter (FH) and Aug. 1895, Thaxter 1559 (FH).—
Mass.: Milton, Aug. 1896, H. Webster (FH); Plymouth, Farlow (FH); Hornblower,
det. D. H. Linder (FH); Wakefield, Singer (FH); Harvard, Singer (FH).— Pa.:
Center Co. State College, Sept. 1933, W. L. White 1284 (FH).—Ga.: vicinity of
Tallulah Falls, July-Sept. 1901, A. B. Seymour, det. W. G. Farlow (FH).— Iowa:
Iowa City, G. W. Newton, det. A. P. Morgan (FH).— Mich.: Whitmore Lake, July
1921, L. E. Wehmeyer (FH).— Tenn.: Cades Cove, Blount Co., July 1938, mixed with
S. confusus, J. B. Porter (FH).— Austria: Rekawinkel, Niederoesterreich, Hoehnel
(FH) ; — Germany: Chiemsee, southern Bavaria, Aug. 1922, Singer; — France: sev-
eral collections of fresh material; — Spain: Catalonia, La Sellera, Codina, det. Singer,
Aug. 1934 (BC) ;— Caucasian Mts.: Swanetia, Kha, under Corylus ?colurna, Aug.
1929, Singer (W) ; Filimonov Range, under Fagus orientalis and Abies Nordmaniana on
decayed wood, Aug. 1935, Vasilieva (LE, KAZ.) ; also other Caucasian localities, once
on rhizomata of Struthiopteris germanica, etc.

There is no doubt that this species can easily be separated from all other
species of this group. Unfortunately, the well-introduced and generally
accepted name S. strobilaceus is illegal since Fries in his Systema Mycolo-
gicum used a name ex Flora Danica, but later has both species side by side,
with the unexpectedly jovial comment (translated): “in my sense not
enough different, but I figure, I have to concede a bit to others also.” Had
it been possible to emend this and similar situations by replacing Fries’s
Systema by the Epicrisis as the nomenclatorial starting point, while there
was still time (see Revue de Mycologie 3: 39. 1938), the usual name could
have been saved. Since, mostly because of the war, no steps have been
taken, I do not want to make my own rules in monographic papers, but I
think it is the general desire of all taxonomists that now, since it is too late
for drastic changes, no belated halfway measures may punish those who
have followed the rules strictly. Many important monographic papers have
appeared since my suggestion appeared, and it is to be hoped that their
nomenclature, not the nomenclature of older and now less important papers
be preserved. As for S. strobilaceus, however, there may still be a faint
chance to preserve this name for a valid species, if future careful studies
should prove that S. floccopus and S. strobilaceus represent two independ-
ent species. I have tried to distinguish between a form with reddening flesh
and another one with blackening context, and at first it seemed that a slight
difference in the KOH reaction runs parallel with this differentiation; it
also appeared that there was a correlation between these characters and the
more sepia or more gray color of the pileus, and the shape of the scales;
but on the other hand, there seemed to be at least one more form with a
different combination of characters (the form from Dade Co.). Intermedi-

SINGER: FLoRIDA BOLETINEAE. I 113

ate forms, in the long run, made me abandon my initial scheme. As I see
now, these intermediate forms may at least partly be explained by the fact
that many of them eventually turned out to be S. confusus, but even after
their elimination, the distinction between the typical S. floccopus and S.
strobilaceus is not quite clear, and it would be unwise to identify these forms
as observed by us with European species described as poorly as Boletus
floccopus in Fries’s Systema. Unless further accurate data on a large num-
ber of specimens in Europe as well as in America prove the opposite to be
true, I treat S. floccopus as a variable but homogeneous species.

S. floccopus is considered, by mycophagists, as edible though mediocre
in quality.

EXTRALIMITAL SPECIES

Strobilomyces velutipes Cooke & Mass., Grevillea 18: 5. 1889.
PLATE I, FIc. 6.

Described from Australia, it is said to be ‘“‘resembling S. strobilaceus and
S. nigricans, but entirely different in the spores.”’ I have studied a fragment
of the type (NY). This species actually differs from 5. floccopus and S.
nigricans in having spores which are either smooth, or differently orna-
mented, and smaller than in the other species. I found them short-ellipsoid
to subglobose, fuscous, thick-walled, more often without any ornamentation
than with it, and, in the latter case faintly rough in the outline of the main
wall which is often surrounded by an indistinct hyaline perisporium, 8.5—
10.5 x 7—7.7 y»; the ornamentation consists of warts which frequently are so
dense as to form an interrupted partial reticulation, the reticulation at least
in its lower part imbedded in the main wall much as in Porphyrellus gracilis
but less conspicuously so because of the low ornamentation and the slight
difference in color between the warts and the remaining portions of the wall
(ornamentation rarely reaching 0.7, in height) ; basidia 24-30 x 10-10.5 p,
clavate-subfusiform and short, 2-3—4-spored, often aborted and cystidia-
like; cystidia fusoid and usually apiculate at the apex; hyphae without
clamp connections. S. indicus Lloyd (Mycol. Notes 7: 1331. 1924) is, ac-
cording to the data given by its author, the same species.

S. velutipes demonstrates the transition from smooth to ornamented
spores. It does not demand too much imagination to draw a connecting line
between the smooth short spores of this species and the elongate smooth
spores of Porphyrellus, both with a varying percentage of perforately warted
spores intermixed in a single preparation, with the same chemical characters
(if it can be supposed that the reactions of S. velutipes are essentially the
same as in S. floccopus), the same dull colors, the same habit, etc. The
elongate spores, naturally, have the wall slightly thinner and lighter col-
ored. For those who are used to phylogenetic thinking, there remains little
doubt that the genus Porphyrellus has derived from such forms as S. velu-
tipes.

114 Fartowia, Vor. 2, 1945

Strobilomyces nigricans Berk., Dec. Fung., Lond. Journ. Bot. 4: 139. pl. 6. 1852.

Described from Khassya, British India, it was rediscovered by Patouillard
in one of the collections sent to him from Indo-China. In his Herbarium,
now preserved at the Farlow Herbarium, the dried specimens described by
him (Bull. Soc. Myc. Fr. 39: 54. 1923) are found in good condition though
still labeled with an herbarium name since Patouillard initially thought
this species was new. The acute warts of the center of the pileus remind
one of S. confusus, but S. nigricans is smaller, at least the Indo-Chinese
collection, and the warts are denser. When compared side by side with
either of the American species, it seemed to be different. The spores were
found to be identical with these of S. floccopus: dark fuscous, thick-walled,
short-ellipsoid to subglobose or globose, with a complete network of arcuate
ridges and veins which are up to 0.5 » high, 9-11.5 x 8.5-10.3 p.

The existence of this species shows that S. floccopus, usually characterized
as a very isolated form, is only one out of a whole series of closely related
species.

SPECIES IMPERFECTLY KNOWN

Strobilomyces polypyramis Hook. fil. apud Berk., Dec. Fung., Hook. Journ. Bot.
3: 78. 1851.

The type comes from India. I have not seen any specimens. This is
probably another species of this section.

Strobilomyces montosus Berk., Dec. Fung., Hook. Journ. Bot. 3: 78. 1851.

The type comes from India. I have not seen any specimens. This may
be another species of the Floccopus-complex.

Strobilomyces echinatus Beeli, Bull. Soc. Roy. Bot. Belg. 58: 212. pl. 15, fig. 11.
1926.

The type comes from the Belgian Congo. I have not seen any specimens,
but this species appears to be remarkable because of the acuminate disc,
and the punctate spores (according to Gilbert, Bull. Soc. Mycol. Fr. 52:
258. 1936). It probably belongs in this section.

Sect. Pterospori Sing. sect. nov.
Sporis de apice usque ad hilum alato-costatis.

Characters of the section: see key (p. 106) and the above Latin diagnosis.

Strobilomyces pterosporus Sing. spec. nov.
Pratt I, Fic. 8.

Pileo umbrino-fuligineo, inter squamas subpallidiore, squamis umbrino-atris, ap-
pressis maculato, plus minusve 40 mm. lato, convexo; tubulis griseo-brunneolis in
adultis, longissimis (radio pilei longioribus), circum stipitem depressis, poris concolo-
ribus, irregularibus, compositis, angulatis, amplis, superficiem hymenophoralem ventri-
cosam efformantibus; sporis fuscis vel melleo-fuscis sub microscopio, atris in cumulo,
longitudinaliter (6)-12-costatis vel -alatis, ornamentatione 1.6-2 u alta, ad polum api-
calem abrupte levibus, 16.5-21 x 14-14.5 4; basidiis tetrasporis, circa 25x17 ; cystidiis

SINGER: FLORIDA BOLETINEAE. I 115

versiformibus; hyphis tramalibus haud fibuligeris; stipite concolori, centrali, plus du-
plico longiore quam diameter pilei, e velo ipsum apicem stipitis approximante fibroso-
tomentoso, cylindrico, solido; carne pallida, mutabili, tenui (1-3 mm.) in pileo. Habi-
tatio: Ad terram in silvulis intersecatis apricis Augusto mense. In Liberia Africae
Tropicalis Occidentalis, Firestone # 3, Du River. D. H. Linder 312 (FH).

Strobilomyces costatisporus (Beeli) Gilb., Bull. Soc. Mycol. Fr. 52: 260. 1936.
Boletus costatisporus Beeli, Bull. Soc. Roy. Bot. Belg. 59: 162. 1927.

Described from the Belgian Congo, this species differs from S. pterosporus
in being larger, but having shorter tubes and smaller pores, and also defi-
nitely smaller spores. I have not seen any specimens. S. lepidellus Gilb.
(Bull. Soc. Myc. Fr. 52: 259. 1936) nom. subnud., has, as far as I know,
not been validly described since, and therefore cannot be compared with
either S. pterosporus or S. costatisporus.

Porphyrellus Gilb., Bolets, p. 99. 1931.
Phaeoporus Bat., Bolets, p. 11. 1908, mon Schroeter, Krypt. Fl. Schles.
(1888).
Boletus subgen. Porphyrosporus Smotlacha, Monog. Cesk. h. Hribov.
Dr oil hott
Characters of the genus: see key (p. 106). Type species Boletus por-
phyrosporus Fr., i.e. Porphyrellus pseudoscaber (Secr.) Sing.

Sect. Tristes Sing. sect. nov.
Pileo haud laete colorato; sporis plerumque levibus; carne fortiter colorata reagen-
tium alcalinorum ope.

Characters of the section: see key (p. 107), and the above Latin diag-
nosis. Type species Boletus tristis Pat. & Baker.

DESCRIPTION OF THE SPECIES OBSERVED IN FLORIDA

Porphyrellus pseudoscaber (Secr.) Sing. comb. nov. ssp. typicus.
Boletus pseudoscaber Secr., Mycogr. Suisse 3: 13. 1833.
Boletus porphyrosporus Fr. in Hok, Boleti, p. 13. 1835.
Krombholzia porphyrospora Karst., Hattsw., p. 17. 1882.
Gyroporus porphyrosporus Quél., Enchir., p. 162. 1886.
Boletus nebulosus Peck, 51st Rep. N. Y. St. Mus., p. 292. 1898.
Boletus umbrosus Atk., Journ. Myc. 8: 112. 1902. sec. Snell.
Phaeoporus porphyrosporus Bat., Bolets, p. 11. 1908.
Tubiporus porphyrosporus Ricken, Vademecum, p. 205. 1918.
Porphyrellus porphyrosporus Gilb., Bolets, p. 99. 1931.

There is no need for a detailed description of the type subspecies, since
Kallenbach has given a fairly complete account on the European species,
and Snell has described the American collections (see the following illus-
trations: Die Pilze Mitteleuropas, pl. 26, fig. 1-13, and Mycologia 24: 336.
1932). I found the spores (average of several measurements on various
materials from Austria) 13.5-19—(20.4) x 6.3-8.5-(10.2) p», and among

? Porphyrosporus Smotlacha has been incorrectly cited as a genus by both Gilbert and
Kallenbach.

116 FartowlA, VoL. 2, 1945

them some smaller spores, several of which (among several hundreds of
normal spores) have warts imbedded in the pallid exosporium, 8—9 x 5.4-6
pw; basidia 34-41 x 13.5-15.3 p, 4-spored; cystidia fusoid-mucronate with
the tip rather acute or quite acute, hyaline or more often brown, especially
in old specimens; trama of the hymenophore and structure of the cuticle
exactly as in ssp. cyaneocinctus Sing. (p. 117), the hyphae of the tricho-
dermium are illustrated in Kallenbach, l.c., pl. 33, fig. 47 7%%14°. Old
herbarium material from Europe (Hoehnel, FH) did, however, not show a
true trichodermium. This latter was preserved in but a few fragments, in
others the repent thin cylindric hyphae of the surface (up to 6.5 » in diam-
eter) were either the remaining base of the trichodermium after it had been
rubbed or washed off, or such a trichodermium is not constant in European
plants. The context of the European form is also variable. Sometimes there
is no bluing at all, and these forms are mostly found in coniferous woods
(see Singer, Pilze aus dem Kaukasus, Beih. Botan. Centralbl., Abt. II, 48:
519. 1931); they also were described as a variety, Boletus porphyrosporus
var. minor Bat. & Crawshay (Bull. Soc. Myc. Fr. 39: 267. 1924) but the
decision whether these variations are indications of the presence of a col-
lective species in Europe, or merely fluctuations within an homogeneous
species, must be left for European mycologists. W. H. Snell’s American
collections have, notwithstanding statements to the contrary by the original
authors of evidently identical species, all more or less bluing flesh, and seem
to represent a single form, certainly identical with at least a part of the
European P. pseudoscaber ssp. typicus. In the type of Boletus nebulosus,
the larger spores are thin-walled, and the smaller ones thick-walled, fusoid
or ellipsoid, or constricted; the dimensions of the spores are extremely vari-
able as far as the length is concerned (variation of 100%), but little
variable in breadth: 11—20.5 x 5.8—7.5 ys, mostly 12.5-15.5 x 6—7 yp; basidia
large, 4-spored; cystidia brown often clavate, often gigantic; trama truly
bilateral as in ssp. cyaneocinctus; epicutis of erect to repent clampless
hyphae, some fascicles of hyphae forming fragmentary trichodermia, termi-
nal members of the chains 5—11 p» in diameter, subfusoid, or clavate, not
subulate. When trying to distinguish the typical form of Europe and of
the northern part of the Atlantic Coast States from ssp. cyaneocinctus, one
should keep in mind that most boletes tend to become rimose-tessellate in
age, and unless several observations on many specimens corroborate the
first observations, the positive-finding should not necessarily mean that the
form studied is ssp. cyaneocinctus. It is also important to note that the
characteristic color ring of the latter subspecies does not serve to distinguish
dried specimens, since it disappears on drying.

3. Porphyrellus pseudoscaber (Secr.) Sing. ssp. cyaneocinctus Sing. nom. nov.
Boletus fumosipes Peck, Ann. Rep. N. Y. St. Mus. 50: 108. 1897, pro parte.
Ceriomyces fumosipes Murr., Mycologia 1: 154. 1909.

Pileus “snuff brown” to ‘‘Saccardo’s umber” with paler portions, veluti-
nous or subvelutinous, cracking readily into areoles so that all adult speci-

SINGER: FLoriIpA BoLeTINEAE. I 17

mens are conspicuously rimose-tessellate, the crevasses whitish and running
in all directions, not viscid, convex, 25-115 mm. broad. — Hymenophore
between “cartridge buff” and “cream buff,” becoming grayer, and eventu-
ally with a tinge from the spore color, greenish blue on injury, tubes up to
21 mm. long, deeply depressed around the stipe, leaving no sterile margin
outward, pores forming a very convex outer-lower surface, concolorous
with the tubes, and discoloring on injury the same way as these, the dis-
colored patches finally becoming chestnut brown, diameter up to 2 mm. Ve:
rather wide when mature; spore print “warm sepia” in a good print. —
Stipe pale yellowish at the very apex with lamelloid proliferations of the
walls of the tubes which run down to a characteristic though sometimes
indistinct greenish blue to green (‘deep lichen green” to “Rejane green’)
color ring below which the stipe is “wood brown,” rarely without an apical
coloring, extremely faintly fibrillose or velutinous on pale yellowish ground,
later becoming ‘“‘Saccardo’s umber,” subequal usually with thickened base,
solid, 46-70 x 12-16 mm. — Context white, on exposure staining “porce-
lain blue” to “gobelin blue” near the tubes, but almost entirely bluing when
young and fresh specimens are squeezed, the expressed sap coloring white
paper blue, slightly dirty lilaceous under the cuticle, mostly more or less
bluish in the apex of the stipe; odor none; taste somewhat sour.

Spores variable in shape, usually more or less fusoid, also ellipsoid-oblong,
10.5-14.3-(18) x (4.5)—6—(7.7) , with suprahilar depression, rarely with-
out it, usually with a wall consisting of a brownish inner layer and a hyaline
or pallid, thin exosporium which looks like an optical halo when well de-
veloped and seen under oil immersion, the exosporium usually optically
empty, and then the spores definitely smooth, or in some rare cases per-
forated with thin cylindric spines which do not emerge above the surface of
the exosporium but make the spore appear punctate when the upper surface
is in focus, the walls occasionally seemingly thick but this is caused by the
invisibility, in some instances, of the inner surface of the main wall so that
the latter seems to extend from the surface of the spore to the surface of
the oil drop, but occasionally actually thick, especially in the apical, rarely
in the basal portion; basidia 4-spored, 30-36 x 9.5-14.5 mw; cystidia fusoid,
or broadly ventricose-mucronate, but obtusate, hyaline to brownish, 35—
58 x 10-20 ,; trama of the young tube walls truly bilateral, but very soon
becoming xerocomoid, i.e. bilateral-subregular, the mediostratum initially
darker and denser, soon all hyphae equally colored and equally dense, lat-
eral stratum initially divergent, soon divergent but in a narrow zone where
the trama turns into the subhymenium; cwticle a trichodermium, consisting
of a hymeniform epicutis with dermatocystidioid terminal members of
erect hyphae, arising from a subcuticular layer of more or less erect hyphae
which frequently are to a certain extent broken down into chains of almost
cubical to globose elements, and have a larger diameter than the basal
septum of the cystidioid elements of the epicutis, the latter (dermatocys-
tidia) subulate, a smaller number of them fusoid, the very apex rounded,
26-42 x 7-14 p, hyaline, smooth.

118 FartowlA, Vor. 2, 1945

Chemical reactions. KOH on surface of pileus purplish to reddish but
not as bright as with NH,OH; on context “Etruscan red,” becoming “‘vina-
ceous tawny.” — NH,OH on the surface of the pileus deep purple (‘dark
livid brown’’), the center of the reacting spot eventually ‘English red”’; on
context at some places reddening then browning, at others sordid (little
reaction). — HNO; on context brownish. — FeSO, “deep glaucous gray”
or “dark glaucous gray.” — Formol salmon color. — Methylparamido-
phenol negative.

Habitat: Under oaks on the soil, gregarious. May to August. Rather
frequent though not abundant anywhere.

Distribution: New York to North Florida, and “west to Kentucky” ac-
cording to Murrill. In Florida known only from Gainesville, but certainly
of quite general occurrence in North Florida.

Material studied: Fla., Alachua Co. Gainesville, under Evergreen oak (Quercus lauri-
folia), Aug. 1937, W. A. Murvrill (Boletus fumosipes) (FLAS, co-type) ; also other col-
lections in high hammocks under oaks, June 1—July 25, 1943, Singer F 2193, 2607, 2825
(FH, co-types).— N. Y.: Type, also type of Boletus fumosipes, p.p. (NYS).

As pointed out by Snell, a portion of the type of B. fumosipes Peck has
striate spores and belongs to Boletellus chrysenteroides Snell, while another
portion has smooth spores and this seems to be the basis of Peck’s descrip-
tion. However, the fact remains that the type of B. fumosipes is a mixed
sample, and no type specimen is designated. The author therefore pre-
ferred to avoid Peck’s name by choosing a new name for the subspecies
described above which has been called Boletus fumosipes by Murrill, and
B. sordidus by Coker and Beers.

EXTRALIMITAL SPECIES

Porphyrellus tristis (Pat. & Baker) Sing. comb. nov.
Boletus tristis Pat. & Baker, Journ. Straits Branch R. Asiatic Soc. No. 78: 70. 1918.

I have studied the type in the Patouillard Herbarium (FH). This is an
interesting species. The stipe is subvelutinous, somewhat thickened at the
base, the tubes are subadnate at maturity, wide in age, 3.5 mm. long when
fresh. The spores are fusoid to boat-shaped, porphyry-umbrinous, darker
than all other elongate spores known in this family and in the Boletaceae,
reminding one of Psathyreila (Psathyra), transparent, smooth, some at-
tenuate almost from the base up, wall thin though not extremely so, the
outer layer in some cases slightly obscurely cloudy-punctulate with paler
and darker brown zones, 14—-17—(19) x (3)-4-5.5 w, with or without a
suprahilar depression; basidia 4-spored, 9.5-12.5 y. broad. The cystidia
are fusoid with rounded tips, or subulate, variable in size, hyaline, crowded
at the pores. The trama of the tube walls is truly bilateral with a melleous
to mahogany brown rather narrow mediostratum, and a hyaline, strongly
divergent, somewhat looser, lateral stratum. The cuticle of the pileus is
characterized by erect dermatocystidioid terminal hyphae which form an

SINGER: FLORIDA BOLETINEAE. I 119

epicuticular trichodermium-palisade; they are brown, fusoid-ventricose
and rounded above, or fusoid-mucronate, 70-100 x 14-28 p. The velvet of
the stipe is formed by thin brown erect hyphae with a capitate-balloon-
shaped or clavate terminal hypha, 15-28 x 9-10 » (very characteristic).
All hyphae have clampless septa.

The flesh of this species is said to be white and unchanging, but the same
statement has been made on other species of this group. The color and the
whole appearance strongly suggest P. pseudoscaber which is the nearest
species we know. The strange heterogeneity of the exosporium of many
spores is indicative for the origin of the ornamentation of the spores in the
Strobilomycetaceae by alternating condensations in the exosporium.

This species has been abundantly collected in Singapore but not else-
where.

SPECIES IMPERFECTLY KNOWN

Boletus sordidus Frost, Bull. Buff. Soc. Nat. Sci. 2: 105. 1874.
Ceriomyces sordidus Murr. N. Am. FI. 9 (3): 149. 1910.

Boletus sordidus reminds one of both the type subspecies and the more
southern subspecies cyaneocinctus Sing. of Porphyrellus pseudoscaber. The
spores are 10—15—(17.5) x (4)—5.5—7.5-(9.7) ,» much the same as in P.
pseudoscaber if the measurements are expressed in range limits as above,
but actually differing in being enormously variable as well in length as in
breadth, some spores being as short as 10-11.5 x 6—7 yp, approaching the
shape of the most elongate spores of Strobilomyces velutipes. The walls of
the broadest spores of B. sordidus are very thick and have more or less in-
complete germ pores at the apex which may also be applanate as in Gano-
derma, the walls reaching 1.7 » in diameter at the apical end.

The above data have been taken from authentic material received at
Farlow Herbarium from the Boston Society of Natural History where some
of the types of Frost’s were preserved; it is very probable that this speci-
men actually represents a portion of the type collection. The type has been
described from Vermont, but fresh material has thus far been re-collected
neither there nor elsewhere in New England, and it is to be feared that B.
sordidus is merely a form with abnormal spores of one of the two estab-
lished subspecies of Porphyrellus pseudoscaber, either as Snell first thought
(1.c.) ssp. typicus, or as he later says (Mycologia 28: 475. 1936) Boletus
fumosipes Peck, i.e. Porphyrellus pseudoscaber ssp. cyaneocinctus Sing. As
it seems to me, Boletus sordidus has to remain among the dubious species
of Porphyrellus, sect. Tristes until further evidence shows its correct po-
sition.

Sect. Graciles Sing. sect. nov.

Pileo laete colorato; sporis perforato-punctatis; stipite subreticulato vel lacunoso-
favoso; actione acidorum et alkalium haud intensa.

Characters of the section: see key (p. 107) and the above Latin diagnosis.

120 Fartow!A, Vor. 2, 1945
KEY TO THE SPECIES

A. Stipe merely finely and sometimes obscurely, reticulate, not lacunose. P. gracilis.

A. Stipe deeply anastomosing costate, and lacunose because of these anastomoses, the
ribs lamellose in shape and waxy in consistency.
B. Spores 14.5-18 x 6.5-8.2 ww. Pileus 47-107 mm. in diameter.
4. P. subflavidus

B. Spores 10-14 x 5-6.8 uw. Pileus 40 mm. broad. P. malaccensis

4. Porphyrellus subflavidus (Murr.) Sing. comb. nov.
Tylopilus subflavidus Murr., Mycologia 30 (5): 520. 1938.
Boletellus subflavidus Snell, Mycologia 33 (4): 422. 1941.

Prate I, Fic. 4.

Pileus yellowish to almost argillaceous, or whitish, or with pallid center,
most frequently about ‘‘Massicot yellow” and eventually often ‘clay color’’
when the weather is dry, occasionally with a slight “pale ochraceous buff’
or “pale ochraceous salmon” tinge, flocculose-velutinous, usually becom-
ing rimose-areolate, eventually frequently fibrose-squamose, not viscid,
convex, at length slightly depressed, 47-107 mm. broad. — Hymenophore
pallid grayish with a vinaceous tint, eventually almost as in Tylopilus coni-
cus, very convex or forming a wall near the depression which surrounds the
stipe, tubes long (11-21 mm.), pores concolorous, rather wide; spore print
on white paper ‘‘warm sepia,” in thinner layer “Army brown” or ‘Verona
brown.” — Stipe yellowish white, pale yellow, or light yellow, lamellose to
alveolate-lacunose, solid, occasionally becoming hollow in age, cylindric, or
tapering down, the base usually acuminate, more rarely bulbose, 45-145 x
7-30 mm. — Context pure white, in the base of the stipe usually rich yel-
low, unchanging in all parts; odor slight, fruity, or almost none; taste
bitterish.

Spores melleous with brown markings, somewhat less rich-colored than
in Boletellus betula, with subsmooth to crenulate outline, the exosporium
with imbedded, thin, cylindric spines which when seen from above make
the spore appear punctate, fusoid, with a slight suprahilar depression, 14.5—
18—(20.5) x (6)—6.5-8.3-(8.8) ; basidia 4-spored, 33-38 x 11.5-15.5 p;
cystidia not as striking as in most species of this family, versiform, i.e.
clavate-fusoid, irregularly fusoid, elongate-claviform, fusoid-ampullaceous,
etc., moderately numerous, 27—54x 5.5-10.2 yw, hyaline, not incrusted;
trama of the tube walls truly bilateral, the mediostratum thin, very pale
melleous, denser than the strongly diverging, loose, pure hyaline lateral
stratum, hyphae without clamp connections, somewhat wavy in the medio-
stratum, and somewhat gelatinizing in the lateral stratum, multi-septate,
especially in the mediostratum and, as usual, in the subhymenium; cuticle
of the pileus consisting of a trichodermium, the erect hyphae of the outer-
most layer with ascendant or irregular basal hyphae, more or less fascicu-
late, loose, and not parallel, with cylindric, cylindric-subcapitate, or clavate
terminal hyphae which are neither colored nor incrusted, and have a not
extremely thin wall, broadly rounded above, 30-78 x 3.5-11.5 1; lamellae
of the stipe formed by rather parallel, thin, tender, thin-walled, cylindric,

SINGER: FLORIDA BOLETINEAE. I 121

multi-septate, clampless hyphae of 3—3.5 » diameter, the surface formed by
somewhat inflated, hyaline dermatocystidia e.g. 38 x 11.3-11.5 yp.

Chemical reactions: KOH on pileus cinnamon buff; on context negative;
NH; on pileus avellaneous; NH,OH on pileus cinnamon buff; on context
negative; HNO; on pileus yellow; on context negative; FeSO, on context
negative.

Habitat: In high hammock vegetation, under Quercus virginiana and
O. laurifolia, and in flatwoods under Pinus palustris with Quercus minima,
on the sandy soil, usually gregarious. June to October.

Distribution: Northern Florida.

Material studied: Alachua Co., Murrill (FLAS, type); authentic material (FH) ;
Singer F 2200, F 2539, F 2673, F 2842 (FH).

A well-defined species among the Strobilomycetaceae of this country, but
closely allied to P. malaccensis and P. lacunosus.

EXTRALIMITAL SPECIES

Porphyrellus gracilis (Peck) Sing. comb. nov.

Boletus gracilis Peck, Ann. Rep. N. Y. St. Mus. 24: 78. 1872.
Tylopilus gracilis Henn. in Engl. & Pr., Nat. Pfl. 1 1**: 190. 1897.

Puate I, Fic. 5.

I have studied authentic material sent to Patouillard by Peck (from
Ballston, N. Y.). This material seems to be identical with the Albany types
(NYS). It is remarkable that no author has as yet seen the characteristic
structure of the spore wall which is perforated, as in Ganoderma, by brown
cylinders, scarcely projecting above the surface of the exosporium, but
making the spore clearly appear punctate when the upper surface is focussed
upon; they are 11—-15.3-(17.5) yw long and (5.5)—6.5—60.7 p», broad, melleous,
basidia four-spored, 24—30 x 9.8-11.8 ». The cystidia are versiform, similar
to these of the preceding species. Macroscopically, this species resembles
a new species of Tylopilus, related to T. fellews which will be described in
the second part of this paper. Old spore preparations are of the same color
as in P. pseudoscaber, not as in Tylopilus. Coker and Beers indicate the
spores as being “hazel’’ but a four-year-old good spore print from a speci-
men of Coker’s was found to be “Sudan brown” by the writer. P. gracilis
occurs in New York, New England, and North Carolina, and is said to be
distributed “from Nova Scotia to Georgia.” It has, as Tylopilus gracilis
(Peck) Henn., been indicated by Murrill (Florida Boletes, Mim. Contrib.
Herb. Univ. Fla. Agr. Exp. Sta., p. 2, 1942) as occurring in northern Flor-
ida. However, I was unable to find any evidence in the Herbaria consulted.

Porphyrellus malaccensis (Pat. & Baker) Sing. comb. nov.
Phylloporus malaccensis Pat. & Baker, Journ. Straits Branch R. Asiatic Soc. No. 78:
71. 1918.
It is hard to understand why this species has been published as a Phyl-
loporus. It has nothing in common with that genus, and belongs in the

122 FartowIA, VoL. 2, 1945

closest neighborhood of P. subflavidus. The type from Singapore, which is
deposited in the Patouillard Herbarium (FH), is somewhat smaller than
the Florida species, and has smaller spores, 10—14—(17.5) x 5-6.8 ». No
other difference between these two species can be discovered. Nevertheless
we consider them as independent species.

SPECIES IMPERFECTLY KNOWN

Porphyrellus Cookei (Sacc. and Syd.) Sing. comb. nov.
Boletus lacunosus Cooke & Mass., Grevillea 18: 5. 1889.

The type, which I have not seen, comes from Australia. This species evi-
dently belongs in the P. malaccensis-subflavidus-group. If the spores are
described correctly, this species would be distinguished easily by the spore
measurements which are given as “15x10 »’’ (Cooke & Massee). If it
were not for this one character, I would be unable to point out how it could
be distinguished from P. subflavidus unless “‘pileo subviscoso” indicates an
actual difference in the structure and consistency of the cuticle.

SPECIES EXCLUDENDAE

Porphyrellus alboater (Schw.) Gilbert, Bolets, p. 99. 1931. This is a Tylopilus
(Boletaceae).

Porphyrellus nigrellus (Peck) Gilbert, Bolets, p. 99. 1931. This is the same as T'ylo-
pilus alboater (Schw.) Murr.

Porphyrellus indecisus (Peck) Gilbert, Bolets, p. 99. 1931. This is a Tylopilus (Bo-
letaceae) .

Boletellus Murr., Mycologia 1: 10. 1909, em. Gilbert, Bolets, p. 106. 1931.
Boletogaster Lohwag, Beih. Botan. Centralbl. 42 (2): 274. 1931.
Frostiella Murr., Mim. Contrib. Herb. Univ. Fla. Agr. Exp. Sta. Jan.

5, p. 6. 1942; Snell, Mycologia 34: 409. 1942 (nomen nudum).

Characters of the genus: see key (p. 107). Type species Boletellus
ananas (Curt.) Murr. (Boletus ananas Curt.).

Section Ananae Sing. sect. nov.

Pileo roseolo, purpurascente, rubello, rubido-vaccineo, saepe fortiter squamoso vel
squamuloso-furfuraceo, haud viscido, margine velo appendiculato vel subnudo; sporis
elongatis, longitudinaliter striatis vel vittatis, magnis (i.e. magnitudine majoritatis
sporarum maturarum 17 yw excedente).

KEY TO THE SPECIES

A. The longitudinal ridges of the spores high (projecting about 1 « above the surface
of the spore) ; stipe sometimes red, but not red-tomentose above, nor squamulose,
nor furfuraceous; pileus imbricate-squarrose, bearing large scales (see B1, B2, B3).
B1. Stipe umber brown; carpophore small. B. porphyrius
B2. Stipe pallid or white, sometimes becoming red at the apex or gray at the base,

but never prevalently umber brown; carpophore medium to large.
5. B. ananas
B3. Stipe yellow above, red below; carpophore medium to large. B. emodensis

SINGER: FLORIDA BOLETINEAE. I 123

A. The longitudinal ridges of the spores usually low (projecting much less than 1 4, or
occasionally just reaching 1 « height), never transversely veined-striate; stipe either
red furfuraceous to tomentose above, or squamulose or furfuraceous; pileus mi-
nutely squamulose to glabrous, or tomentose-scrobiculate.

C. Stipe longitudinally striped red, white tomentose at the base, red tomentose-
furfuraceous above, 100 x 12-13 mm.; pileus uneven, scrobiculate and tomentose,
not scaly, not glabrous, 60-70 mm. broad. B. obscurecoccineus

C. Stipe not striped with red, not red tomentose-furfuraceous above, but distinctly
furfuraceous or transversely squamulose, 30-50x4-9 mm.; pileus minutely
squamulose or glabrous, 20-40 mm. broad. B. cubensis

B. lignatilis
B. guadelupensis

DESCRIPTION OF THE SPECIES OBSERVED IN FLORIDA

5. Boletellus ananas (Curt.) Murr., Mycologia 1: 10. 1909.

Boletus ananas Curt., Amer. Journ. Sci. II 6: 251. 1848; Berk., Hook. Journ.
Botan. 1: 101. 1849.

Strobilomyces pallescens Cooke & Mass., Grevillea 18: 5. 1889.

Boletus isabellinus Peck, Bull. Torr. Club 24: 146. 1897.

Boletellus pallescens Gilbert, Bolets, p. 107. 1931.

?Boletus coccineus Fr., Epicr., p. 423. 1838.

?Strobilomyces coccineus Sacc., Syll. 6: 50. 1888.

?Boletus ananaeceps Berk., Linn. Soc. Lond. Journ. (Bot.) 13: 161. 1873.

?Strobilomyces ananaeceps Sacc., Syll. 6: 50. 1888.

?Secotium excavatum Kalchbr., Ertek Termész. Kéreb Kiadja a Magy. Tud. Ak.
13 (8): 7. pl. 3. 1884.

?Strobilomyces excavatus Henn., Hedwigia 43: 187. 1904.

Prate I, Fic. 7.

Pileus with “deep vinaceous,” or “Pompeian red” and “Isabella color”
fibers, a general color of about “russet vinaceous” resulting, bleaching when
old, fascicles of the fibres forming large woolly-squarrose or subimbricate
scales which are the more erect the nearer they are to the center, non-viscid,
subglobose at first, becoming convex-expanded, 45-95 mm. in diameter;
margin extended into a thick membranaceous veil which is white or nearly
so, and initially runs down the apex of the stipe in two indistinct layers,
the outer one thin and fragmentary, later breaking off the stipe to leave a
sleeve just above the surface of the stipe, finally becoming ragged to form
a sordid and pale stramineous, irregularly crenulate-denticulate appendicu-
lation on the margin; these remainders of the appendiculate veil disappear
at or after maturity. — Hymenophore yellow, often tinged with reddish
brown, staining blue where injured, eventually becoming yellowish brown
or reddish brown all over; tubes depressed around the stipe, 11-16 mm.
long; pores concolorous, large, irregular, angular, forming a distinctly con-
vex lower surface; spore print black or nearly so. — Stipe white, or pallid,
often with a red belt on the apex (‘‘Pompeian red” or lighter, corresponding
to the blue or green belt of Porphyrellus pseudoscaber ssp. cyaneocinctus),
described as grayish below (Cleland for Australian specimens), but only
becoming pinkish sordid or sordid when old in American specimens, sub-
fibrillose-subglabrous to fibrous, but rarely with an actual annular remnant

124 FartowiA, Vor. 2, 1945

of the veil on the stipe, not scaly-floccose nor shaggy, smooth, tapering up-
wards when young, later rarely with subbulbous base, or sometimes taper-
ing downwards, or more frequently subequal, solid, 60-140 x 8-15 mm. —
Context yellowish white or whitish yellow in the pileus, white in the stipe
or only in the base of the stipe, readily and strongly bluing when cut, at
least in the pileus, occasionally reddening in the stipe; odor none; taste
mild.

Spores 13.8-25.5x 7-11 py, mostly 17.5-22.5 x 7.5-9.2 » in American
specimens, with strongly projecting longitudinal, slightly spiralling, lamel-
losely-thin wing-like ridges which abruptly are cut off before they reach
the indistinct germinative pore, leaving a very narrow hole at the very apex
of the spore, the sides of the ridges and the bottom of the furrows between
them transversely venose-striate, the ridges numbering from 12 to 14, and
measuring from 0.6—1.2 », mostly 1 » in height; basidia 22.5—50 x 9.5-17.5
pt, 2- to 4-spored; cystidia fusoid, mucronate, or ampullaceous to ampulla-
ceous-capitate, hyaline, brittle, 24-50 x 5-15 yu; trama of the tube-walls
truly bilateral with more colored mediostratum and looser, divergent lateral
stratum, the hyphae without clamp connections; hyphae of the outer layer
of the vezi hyaline, semiopaque, long-cylindric, irregularly interwoven, thin-
walled, 4—7 p» thick, the septa without clamp connections, the terminal
members of the hyphae long, with attenuate though rounded, or suddenly
broadly rounded, or subcapitate tips.

Chemical reactions: KOH on context yellow, becoming brown; on the
surface of the pileus olive-yellow, becoming maroon; NH,OH on the sur-
face of the pileus olive-yellow; FeSO, on the context green; aniline nega-
tive; chlorovanilline on context becoming yellow where the context had
been white (probably only the acid — HCl — reacting).

Habitat: In various kinds of woods, from open pine-oak woods to dense
tropical hammocks, frequently on the bases of trees (Pinus palustris, P.
caribaea, Quercus virginiana, Q. Catesbaea, Nectandra coriacea), or on
shaded stone walls, climbing at times as high as three feet above the ground,
but also on sandy soil under oaks or pines, solitary to gregarious. From
May to November in the Northern Hemisphere, December to May in South
Australia (according to Cleland).

Distribution: Southern States of the U.S. A., from South Carolina to the
south tip of Florida and west to Alabama and Mississippi, probably reach-
ing Mexico and the South Pacific Islands but known with certainty only
from New Caledonia, also in Queensland and South Australia. In Florida
through all zones.

Material studied: Fla.: Dade Co. Matheson Hammock and Fairchild Tropical Gar-
den, September to November 1942, Singer F 709, F 709a, F 709b, F 1391 (FH); Lake
Co., Eustis, Oct. 1897, Thaxter 2444; Alachua Co., Gainesville, August 1937, Murrill
(FLAS, FH); Sugarfoot Hammock, July 1943, Singer F 2541, F 2541a (FH); New-
nan’s Lake, July 1943, Murrill (FLAS); Madison Co., Cherry Lake, July, Murrill
(FLAS): Marion Co., Goldhead Park, May, Murrill (FLAS).—S. C.: Santee River,

type of Boletus ananas Curt. (Curtis Herbarium, FH).—Ga.: North-east Georgia,
vicinity of the Tallulah Falls (Tallulah Falls, Bad Branch, Sunset Rock, Rock Moun-

SINGER: FLORIDA BOLETINEAE. I 125

tain), from August to September, A. B. Seymour & W. L. Moss, det. W. G. Farlow
(FH).— Ala.: Earle (FH).—Miss.: Type of Boletus isabellinus (NYS).— New
Caledonia (Nouvelle Calédonie): on iron-rich soil, Le Rat, 1808, det. Patouillard
(S. pallescens), (FH).— Queensland, Australia: Type of Strobilomyces pallescens
(NY).

Coker and Beers, and also Murrill, Earle, and others emphasized the
growth of the carpophores on the base of trees, exclusively pines as it was
understood by these authors. Earle supposed parasitism, and Murrill based
the genus Boletellus, among other characters, on the epixylous growth of the
carpophores. This, however, is not in full agreement with the facts. I have
in numerous careful observations never been able to find any injury on trees
that could be attributed to this fungus, nor is it true that this species is
always epixylous, or confined to pines. At some stations, there were abso-
lutely no pines in the vicinity, and at others, the carpophores definitely
preferred the bases of oak trees to those of pines in a mixed stand. The fact
that many specimens grew, of all places, on scanty humus and débris on
stone walls, the mycelium appressed to the rocks, seems to show that
Boletellus ananas merely prefers to grow along hard surfaces, but most cer-
tainly it is not a parasite unless, of course, the mycelium is connected with
the roots, and has loose mycorrhizal relationship with oaks and pines, with
possible disturbances of the mutual character of the symbiotic relations in
favor of the fungus, as it happens under certain conditions with other
boletes. However, the lack of specialization on the part of B. ananas, as
well as the growth on stone walls instead of close to the roots of the trees,
or on the ground under trees, makes it highly dubious whether it is a my-
corrhizal fungus at all. Without experimental studies proving the fact of
actual formation of mycorrhiza, field observations are not sufficient to
answer this particular question. At any event, the idea that B. ananas is an
exclusive parasite of pine trunks must now be abandoned. Another interest-
ing fact about B. ananas is the geographic area occupied by this species. If
it is interrupted, this would by itself constitute a rare and interesting phe-
nomenon, but it may well be that the many little explored regions and small
islands between the known areas, after more exploration, will furnish a
bridge between the American and the Australo-Oceanic centers of distri-
bution.

The spores seem to average a trifle larger in the Australian plant, but we
obtained two practically identical measurements from the New Caledonian
material in Patouillard’s Herbarium and from our Dade Co. (Fla., U.S. A.)
collections: 19—22.5 x 8-9.5-(11) » in New Caledonia, and 18—22.5 x 8.8-
9.2 » in South Florida. There is also Cleland’s indication that the stipe is
gray below, but Cooke (Queensland-type) says it is pallid. It seems there-
fore that these spurious differences cannot be used for the substantiation of
geographic races. Lloyd’s Strobilomyces pallescens from this country is, no
doubt, plain B. ananas.

Krieger (The Mushroom Handbook, p. 441. 1936) uses the name Stro-
bilomyces coccineus (Fr.) Sacc. for our Boletellus ananas. As for the ge-

126 FartowlA, VoL. 2, 1945

neric name, I refer to the observations in the present paper (p. 105); the
specific name has the advantage of being undoubtedly the oldest possible
name having priority over Boletus ananas. When Fries described this spe-
cies, he based his data on Plumier’s laconic characterization and somewhat
fantastic picture in black and white. Notwithstanding the very inadequate
description and the poor drawing, it is hard to imagine what other species
it could possibly represent. On the other hand, it has to be remembered
that Plumier gives no locality, and all we know is that it was found some-
where in America. Nobody has ever heard of there being any specimens
preserved. This is, in my opinion, not enough evidence to unsettle an
established name like B. ananas, and Murrill is right in referring to Boletus
coccineus in the list of doubtful species. Murrill also was right in referring
Boletus isabellinus Peck to Boletellus ananas asa synonym. The type speci-
mens at Albany do not leave any doubt as to their identity. If there is the
slightest similarity between plate 3 of Kalchbrenner (l.c.) and the fungus
painted, this latter, Secotium excavatum, cannot be a Boletellus, and by no
means B. ananas. But Hennings who has seen the “originals” flatly de-
clares Boletus pallescens and Secotium excavatum to be synonyms. Re-
examination of the types of the latter will decide if he is right.

EXTRALIMITAL SPECIES

Boletellus porphyrius (Pat. & Baker) Gilbert, Bolets, p. 107. 1931.
Strobilomyces porphyrius Pat. & Baker, Journ. Straits Branch R. Asiatic Soc. No.
78: 72. 1918.

Described from Singapore, this species has been transferred to Boletellus
by Gilbert because of the indication in the original description ‘Spores
longitudinally striate” and because of the sordid yellowish hymenium. The
examination of the type reveals that Boletellus porphyrius actually belongs
in this genus, and that it can be characterized as a smaller B. ananas with
umber brown stipe. The spores are the same in both these species. I found
them with exactly the same double markings as in B. ananas and of the
same size: 17.5—21 x 7.5—9.5 ». Basidia 32-46 x 12-18 »; hyphae without
clamp connections.

Boletellus emodensis (Berk.) Sing., Ann. Mycol. 40: 18. 1942.
Boletus emodensis Berk., Hook. Journ. Bot. 3: 48. 1851.
Strobilomyces annamiticus Pat., Bull. Soc. Mycol. Fr. 25: 6, pl. 1, fig. 1. 1909.
Boletellus annamiticus Gilbert, Bolets, p. 107. 1931.

The characters of this beautiful Asiatic species are plainly those of the
section Ananae, even the peculiar veil of B. ananas being well represented.
The originals of Hooker’s paintings of Sikkim fungi are preserved at Far-
low Herbarium, and they show clearly enough that B. emodensis is very
similar to B. ananas. We have not studied the types which are in England,
but we have had material from China, province of Kwangsi, and the types
of Strobilomyces annamiticus Pat. in the Patouillard Herbarium. The lat-
ter is doubtless a mere form of B. emodensis if not a plain synonym. The

SINGER: FLoRIDA BOLETINEAE. I 127

spores of B. emodensis are 18-22.5 x 8.3-11 mw with the wings transversely
striate, and 1-1.5 » high. The spores as indicated by Patouillard come from
immature material. Immature carpophores of the Chinese collection have
the same spores as indicated by Patouillard (though there is a printing
error in his description: 10 instead of 20 »). The basidia are 39-68 x 12—
14.3 u. The cystidia we have seen are smaller than the basidia or equal the
basidia and they are smaller than indicated by Patouillard. The lateral
stratum of the hymenophoral trama is distinctly divergent. The mycelium
of all specimens is very strongly developed, forming a large, sometimes
yellow tomentum. This species differs from B. ananas in the color of the
stipe, and possibly the deeper red of the pileus and of the hymenophore.
It also differs from B. porphyrius in the colors and the size. Nevertheless,
these three species form a closely related group (a stirps) within the sec-
tion. This stirps is characterized by the marginal veil, large spores with
double ornamentation, and scaly pileus. Boletus squamatus Berk. is prob-
ably another species of this group, if it is not a synonym of B. emodensis
with somewhat faded colors.

Boletellus obscure-coccineus (H6hn.) Sing. comb. nov.
Boletus obscure-coccineus Hohnel, Sitz.-ber. Akad. Wiss. Wien 123 (1): 88. 1914.

Hohnel, who collected and described this species in Java, mistakenly
thought it was especially close to Boletus versicolor i.e. B. rubellus Krombh.
The type specimens preserved in the Herbarium Hohnel (FH) show that
it is a Boletellus with the hymenophore convex beneath, 8-9 mm. thick,
and the spores 15.5—21 x 7.5—8 y, melleous when mature (not brown) with
comparatively thin walls but with distinct, fine, venose, longitudinal ridges.
Basidia 30—42 x 8.5-12.5 yu, 4-spored. Cystidia mucronate, about 100 p»
long and 14—24 » broad. Hyphae of the tube-trama hyaline, cylindrical,
smooth, without clamp connections. This species reminds one macroscopi-
cally of Boletellus emodensis, and microscopically of Boletellus cubensis
and its allies.

Boletellus cubensis (B. & C.) Sing. comb. nov.
Boletus cubensis Berk. & Curt., Journ. Linn. Soc. 10: 304. 1868.

The spores of the fragment of the Kew type (NY) are 20-22.5 x 7.5-10 p,
and striped with longitudinal low ridges (less than 1 » high), deep mel-
leous, fusoid, somewhat truncate at the apex, the ridges slightly anastomos-
ing but not transversely veined as in Boletellus ananas, with suprahilar
depression. This shows that it belongs in the neighborhood of Boletellus
obscure-coccineus, It may well be that it will be discovered in South Florida
or on the Keys since many Cuban species have been found there by the
writer.

Boletus lignatilis Berk. & Curt., Journ. Linn. Soc. 10: 303. 1868.

This species also comes from Cuba, and the spores of the fragment of
the Kew type (NY) are longitudinally ridged (ridges up to 0.8 » high)
and the ridges connected by occasional to frequent anastomoses (but not

128 FartowI1A, VoL. 2, 1945

transversely venose as in Boletellus ananas or B. chrysenteroides). They
are deep melleous and depressed above the hilar end, 18-21.5 x 7.5-9 up.
This is doubtlessly a Boletellus of this section, but no new combination is
proposed because of the probability that this and the preceding species are
identical.

Boletus guadelupensis Pat., Bull. Soc. Myc. Fr. 16: 177. 1900.

Collected on Guadeloupe, West Indies, this is possibly another synonym
(a younger stage) of Boletellus cubensis (B. & C.) Sing. The spores of the
type (FH) are 14~-19.5 x 6.5-8.5 yw, and longitudinally striate from the
hilar end to the tip where the striae suddenly end, very few mature (col-
ored) spores remaining smooth or almost so, melleous, the walls browner,
with a slight suprahilar depression or applanation; I have not seen the
yellow prominent cystidia. According to the original description, this spe-
cies differs from B. cubensis in chestnut brown pileus and appendiculate
margin. The former difference may be due to the fact that the specimen
dried to that color, but was reddish when quite fresh, as is evident in the
dried specimen. This may be explained by the hypothesis that this species
represents a younger stage of B. cubensis which would also account for the
smaller size of the whole carpophore as well as of the spores. There are
also minor differences in the description of the surface of the stipe. Never-
theless, I think it is necessary to refrain from transferring B. guadelupensis
to Boletellus as an independent species, as long as no data on the fresh
plants are available. Unfortunately the Cuban plants as well as B. guade-
lupensis seem to be rare.

SPECIES IMPERFECTLY KNOWN

a

Boletus squamatus Berk., Hook. Journ. Bot. 4: 137. 1852.

I think that this Indian species is another member of the section Ananae,
but since nothing is known about the spores, further studies are necessary,
before a new combination can be justified. Berkeley thinks that his species
is allied to Boletus subtomentosus and B. chrysenteron, but Hooker’s origi-
nal painting reminds one of Boletellus porphyrius and B. emodensis rather
than of any Xerocomus.

Sect. Mirabiles Sing. sect. nov.
Pileo minute tomentoso vel squarroso, margine secedente sterili; stipite plus minusve
reticulato; sporis giganteis, levibus.

Characters of the section: see key (p. 107), and Latin diagnosis above.
This is a remarkable section because of the enormously large, smooth spores,
and the projecting margin. It has been treated by Singer conditionally in
Xerocomus, but it actually is close to the remaining sections of Boletellus.
The type species is Ceriomyces mirabilis Murr. There are no representa-
tives of this section known to occur in Florida.

SINGER: FLORIDA BOLETINEAE. I 129

Boletellus mirabilis (Murr.) Sing. comb. nov.
Ceriomyces mirabilis Murr., Mycologia 4: 98. 1912.
Xerocomus mirabilis Sing., Rev. Myc. 5: 6. 1940.

The type of this western species has been studied by the writer and com-
pared with material obtained by Alexander H. Smith. As I said in 1942,
“The exact structure of the trama could not be stated. If this would be
extremely bilateral, generic separation from Xerocomus should be taken in
consideration.”’ Actually, the trama of the tube walls zs truly bilateral. The
spores are 18—26.3 x 7-10.5 pu, smooth, ellipsoid-oblong to subfusiform or
fusiform, yellowish brown (color in mass unknown to me), wall 1-1.8 p»
thick, but also thinner in many cases, slightly obtusate at the apex in many
cases, with suprahilar applanation; basidia 33.5-39 x 10-14.5 yu, 4-spored;
cystidia 60-80x 11-16 yw, ventricose or ampullaceous or more or less
strongly capitate above, or plainly fusoid; hyphae of the truly bilateral
trama without clamp connections and divergent in the lateral stratum;
papillae of the cuticle consisting of chains of thin (about 8 ,»), or thick
(10-18 ») clampless hyphae, in the latter case attenuated toward the septa,
brown, about 20—60 p» long between septa, the terminal member occasionally
shortened, and rarely spherocystoid. Kauffman (Pap. Mich. Acad. Sci.
Arts & Lett. 5 (1925): 115. 1926) redescribed this species from Mt. Hood,
Ore. (Wehkmeyer 28130, MICH). Snell had it from Manitoba, Slip and
Snell indicate it from Idaho, the type came from Washington, and Overholts
indicates it in Pennsylvania. This latter extension of the area of distribu-
tion is surprising, if true.

Boletellus projectellus (Murr.) Sing. comb. nov.
Ceriomyces projectellus Murr., Mycologia 30: 524. 1938.
Boletus projectellus Murr., Mycologia 30: 525. 1938; Coker & Beers, Boletaceae,
p. 51. 1943.

This species has been found in Virginia and North Carolina, U.S. A. For
a good macroscopical description and a colored picture see Coker and Beers,
l.c., pl. 1, fig. 3-4. The type specimens (Murrill F 9296, FLAS), authentic
material (J. N. Couch F 16045, FLAS) and other North Carolina material
(J. N. Couch 13193, FH), were compared by the writer and found to be
identical. Couch’s collections are evidently the basis of the description and
plate of Coker and Beers. The anatomical characters of these plants have
been found to be as follows: spores (20)—22—30—(32) x (5.8)—-8.5-11-(12)
p», smooth, melleous to deep melleous, the walls 1-1.5 » thick when mature,
the upper portion attenuated from the lower third, or the outline more
evenly elliptic-oblong to boat-shaped in frontal view, with a slight depres-
sion in profile, always more or less acuminate though obtuse at both ends,
the inner wall interrupted and the germ pore incomplete since the exospo-
rium is continuous; basidia 23-36 x (10)—12-15 yp, 4-spored, few 2-spored;
cystidia fusoid with ampullaceous apex, or lanceolate with obtuse tip,
hyaline, (33)—35—70 x 10-17 yw; trama truly bilateral, the mediostratum
slightly denser and dilutely colored, the lateral stratum somewhat diverg-

130 FarLowI1A, Vor. 2, 1945

ing (in nearly adult specimens), clamp connections none. Boletellus pro-
jectellus is one of the representatives of the Agaricales with the longest
spores, and belongs to the species with the most voluminous spores of the
whole order. It certainly has the largest spores of all boletes.

Sect. Retispori Sing. sect. nov.

Sporis breviusculis, reticulatis.

Characters of the section: see key (p. 107) and the Latin diagnosis
_above. The section is unique in combining spores of the type of Strobilo-
myces floccopus with entirely different macroscopical features. The only
species is Boletellus retisporus which has not been found in Florida.

Boletellus retisporus (Pat. & Baker) Gilbert, Bolets, p. 108. 1931.
Boletus retisporus Pat. & Baker, Journ. Straits Branch R. Asiatic Soc. No. 78: 72.
1918.

This Singapore species is extremely interesting, and easily identified by
the combination of yellow hymenophore and short, reticulate spores. I
have studied the type specimen in the Patouillard Herbarium (FH). The
spores are asymmetrical, brown, the ridges forming the complete and con-
spicuous network are 1.3—2 » high, but often half or entirely imbedded in
a colorless outer wall. My measurements are: 12—12.5 x 8-9.5 w. Basidia
28x11 yw. Hyphae 5-12 » thick in the trama, without clamp connections.
The mycelium is white, and the pileus, though this is not mentioned in the
original description, is distinctly appressed-squamulose.

Sect. Chrysenteroidei Sing. sect. nov.

Pileo haud roseo nec purpurascente nec carmineo, at flavo, fusco, castaneo; sporis
longitudinaliter vittatis vel striatis, aut levibus, elongatis, aut latissimis (10.5-11 ») aut
moderate voluminosis (circa 12.5-17 « longis) ; velo spurio vel nullo; pileo stipiteque
haud viscidis; stipite haud ceraceo-lacunoso.

Characters of the section: see key (p. 107), and the above Latin diag-
nosis. The type species is Boletellus chrysenteroides (Snell) Snell.

KEY TO THE SPECIES

A. Spores consistently smooth. Boletellus turbinatus
A. Spores, at least their overwhelming majority, ridged because of longitudinal lamel-
lose wings. .
B. Wings of the spores striped transversely by low veins, or, at least, the bases of
the wings connected by such transversal veins, giving the spores an appearance

reminiscent of the ornamentation of the spores in Boletellus ananas.
Boletellus chrysenteroides

B. Wings not connected nor striped by regular transversal veins.

C. Pores discolorous, not yellow to melleous as the tubes, but red.
D. Spores 15.5-17 x 10.5-11 yu, with the lamellose wings projecting 1-1.7 uw.
Boletellus Linderi

D. Spores 12.5-15.2 x 6.5—7 w; wings lower than 1 yu.
6. Boletellus pictiformis

SINGER: FLORIDA BOLETINEAE. I 131

C. Pores concolorous, yellow or melleous.

E. Spores (12.2)—-13-13.8-(15.3) x (5.5)-6-(6.8) w. Boletellus pictiformis var.
fallax

E. Spores 14—22.5 x 6.5-10 » (compare B. chrysenteroides, if collected on the
North American Continent, and B. guadelupensis if collected in the West
Indies) .

DESCRIPTION OF THE SPECIES OCCURRING IN FLORIDA

6. Boletellus pictiformis (Murr.) Sing. comb. nov. var. typicus.
Suillellus pictiformis Murr., Lloydia 6 (3): 226. 1943.

Pileus castaneous, densely shaggy-tomentose, convex, 40-60 mm. broad.
— Hymenophore yellow or melleous with red pores, changing at once to
cyaneous when wounded, plane pores 1 mm. in diameter. — Stipe lateri-
tious above, castaneous below, very shaggy, sulphureous at the apex, en-
larged upward, 60-80 x 10-13 mm.— Context citrinous, yellow, turning
blue at once; taste mild or acid. (Macroscopical description drawn from
W. A. Murrill’s notes.)

Spores 12.5-15.2 x 6.5—7 yp, intensely melleous, subfusoid, with longi-
tudinal ridges from the tip downward to the hilar tube, the ridges occa-
sionally anastomosing or forking, but their sides and the space between
them not veined transversely; basidia 27-34 x 11.5-14.2 p, 4-spored; cys-
tidia clavate-mucronate, or fusoid-ventricose with a cylindric ampullaceous
neck which sometimes is curved, or merely fusoid with obtuse apex, hyaline,
or with a melleous belt-like incrustation; terminal hyphae of the tomentum
of the pileus melleous, clavate, 82 x 10.8 », the lower hyphae 4-14 p» in
diameter, without clamp connections.

Habitat: On banks under trees. July to August.

Distribution: North Florida.

Material studied: Kelley’s Hammock, Alachua Co., Murrill (FLAS, type); also
authentic material from Sanchez and Planera Hammock, Alachua Co., Murrill F 17945,
F 17862 (FLAS).

The type variety is evidently very rare even in the large hammock re-
gion northwest of Gainesville where it has been found exclusively. It is so
rare indeed that W. A. Murrill has collected it only four times since 1938,
and I was not able to gather specimens during my stay in Gainesville. It is
remarkable because of its relation with an African species, B. Linderi.
These species are very similar in many regards and are no doubt closely
related. It is interesting to find two species with striking affinities at two
isolated points on opposite sides of the Atlantic. B. pictiformis may well
be derived from forms with African ancestry, but it would be dangerous to
generalize concerning this probability for all Strobilomycetaceae of this
hemisphere, since B. ananas and B. porphyrius, Porphyrellus subflavidus,
P. malaccensis and P. lacunosus, and also Boletellus jalapensis show quite
different geographic relationships, extending rather across the Pacific than
across the Atlantic.

i3z FartowiA, VoL. 2, 1945

Boletellus pictiformis (Murr.) Sing. var. fallax Sing. var. nov.
A typo differt poris concoloribus cystidiisque paucis incrustatis.

Pileus brown because of the “‘Prout’s brown,” “mummy brown,” or “bis-

ter” appressed fibrose scales with the pale yellow flesh showing through
between them, not viscid, convex, without or with only slightly projecting
margin, 42-53 mm. broad. — Hymenophore “yellowish citrine” to “olive
lake” when mature, with concolorous pores which are angular and number
5-10 per 5 mm. transversely, and individually 0.3—1.5 mm. in diameter, i.e.
rather large when quite mature, the tubes somewhat depressed around the
stipe, moderately long (5-7 mm.), turning blue on injury. — Stipe below
“mummy brown,” in the middle ‘“‘madder brown” to “Vandyke brown,” or
paler and with yellow shining through, and at the apex “Pinard yellow,”
fibrous-scaly-shaggy in the color of the pileus, the scales often arranged in
a coarsely reticulate manner, solid, equal, with acuminate-tapering base,
60 x 12-13 mm. — Context of the pileus yellow, richer yellow at the apex
of the stipe (“‘baryta yellow” to “empire yellow”), between ‘“‘bay” and
blackish brown at the base, strongly and readily turning blue in the pileus
when cut, rather thick in the pileus; odor and taste not remarkable.

Spores (12.2)—13-13.7-(15.3) x (5.5)—6-(6.8) , deep melleous, with
low to medium high longitudinal lamellose wings which are not connected
by regular transverse veins, but occasionally by oblique anastomoses of the
wings which are often forked, with suprahilar depression or applanation,
ellipsoid-fusoid; basidia 4-spored, 25-39x 12-13.7 yw; cystidia numerous
near pores, mostly ampullaceous, pale melleous to hyaline, not incrusted or
occasionally somewhat incrusted, (23)—38—60 x 10-14.2 »; also some ex-
tremely versiform cheilocystidia are present at the very edge of the tube
walls; cuticle of the pileus made up of repent or obliquely ascending strands
of parallel, cylindric hyphae with the second-last member always elongate
and 4-10 ,» thick, the terminal member cylindric with rounded ends and as
long as the other hyphae, or shorter, with some rare spherocysts on the
very surface.

Chemical reactions: KOH on the surface of the pileus eventually deep
reddish brown; on tubes and context ochraceous brown. — NH,OH on the
surface of the pileus black; on the tubes and on the context negative, or
removing the blue stain when applied after autoxydation has taken place.
— HNO; on the surface of the pileus rufous; on the context pale reddish-
brown. — FeSO, on the surface of the pileus negative; on the context
negative, or removing the blue stain if applied after the autoxydation has
taken place. —— Formol on the context provoking more blue coloration.

Habitat; Under mixed hardwoods, in a mesophytic hammock. July.

Distribution: North Florida.

Material studied: Kelley’s Hammock, N. W. of Gainesville, Alachua Co., July 14,
1943, Singer F 2716 (FH).

This variety lacks the most characteristic feature of the type, and Mur-
rill did not recognize it as his species when he saw it in fresh condition.

SINGER: FLORIDA BOLETINEAE. I 133

However, the complete identity of the anatomical characters and the fact
that the type of var. typicus and the type of var. fallax were found in the
same hammock, and also the agreement of the important macroscopical
characters, makes it impossible to consider the variety as a separate species.
I have called it var. fallax because it suggests other species rather than B.
pictiformis, e.g. Boletellus chrysenteroides (Snell) Snell which, however,
has not a floccose-fibrillose-squamose stipe, but only shows furfuraceous
punctations. Its spores usually have distinct transverse, regular anastomos-
ing veins between the wings, reminding one of the ornamentation of B.
ananas. Nothing like this is seen in spores of B. pictiformis var. fallax. The
cuticle of B. chrysenteroides and B. pictiformis var. fallax also are very
different in structure.

EXTRALIMITAL SPECIES

Boletellus chrysenteroides (Snell) Snell, Mycologia 33: 422. 1941.
Boletus chrysenteroides Snell, Mycologia 28: 468. 1936.

This has been described from New York, but additional material has been
found in the Farlow Herbarium from Glenbrook Ravine, northeast Georgia,
A. B. Seymour, det. W. H. Snell (B. fumosipes), and there is material in
E. V. Seeler’s Herbarium, collected by him on Nantucket, Mass., det. D. H.
Linder & R. Singer. W. H. Snell was kind enough to send me his entire
material, including specimens from North Carolina, West Virginia, Ten-
nessee, Missouri, and Pennsylvania, all the same species with the covering
of the pileus like that of Porphyrellus pseudoscaber, and actually with the
same type of structure, but of a different brown. The spores are somewhat
variable from specimen to specimen, (9.5)—11—17.3—(18.2) x 5.8-9.8 yp,
with longitudinal lamellose wings of 0.6—-0.8 » height, melleous, the wings
connected by regular, transverse veins recalling those of Boletellus ananas
but not always running all through and up the sides of the wings reaching
their edges; basidia 4-spored, 25-39 x 11-19 p»; cystidia fusoid to ampul-
laceous, hyaline or pale melleous, not incrusted, numerous on and near the
pores, 34-62 x 9.5-18.5 »; trama initially truly bilateral (Boletus-type),
later more regular; epicutis formed by a palisade of erect, septate hyphae,
with the last members conic-fusoid or cylindric, the second-last ones conic-
obtuse or like the lower ones, i.e. almost as broad as long and rather volu-
minous, fulvous brown; they measure: last member 17-18 x 5.8-9.8 p,
second 17-18 x 13-17 » below, and 7-11 » above, third and following mem-
bers 17-18 x 13-17 »; hyphae without clamp connections.

One of the specimens, a co-type of Snell’s, collected at Burnet, Quebec,
Canada, differs from all other specimens in having the longitudinal wings
of the spores more frequently anastomosing, or even with transverse ridges,
but not always showing the characteristic thin, low veins found in the type;
these spores are a trifle larger, 13-18.7—-(20) x 7.3-9.2-(10.2) mw, and thus
are intermediate between B. chrysenteroides and B. guadelupensis; their
wings are 0.8-0.9 » high, some lower. The external appearance of the fruit

134 FartowiA, Vow. 2, 1945

bodies is nearly the same as in the type, but they are larger in all parts.
Since two small characters are coupled in this case, distinguishing the
northernmost collection from all the rest, one may suspect that the Ca-
nadian form is a geographic race. Further observations on this subject are
needed. B. chrysenteroides is apparently the northern equivalent of the
Floridian B. pictiformis.

Boletellus Linderi Sing. spec. nov.
Pirate I, Fic. 10.

Pileo flavo (“Naples yellow” Ridgway), fibrillulis squamulisque badiocastaneis (“ma-
hogany red” Ridgway) dense obtecto, (25)—60 mm. lato; hymenophoro poris primum
rubris et aequalibus, deinde aurantiacis et irregularibus instructo, e tubulis longis con-
sistente; sporis ellipsoideis, brunneis, 17—23—alatis, ornamentatione 0.9-1.7 yw alta, polo
apicali levi, 15.5-17 x 10.5-11 w metientibus; basidiis 34x 14.7 yu, clavatis, tetrasporis;
cystidiis circa 63x22 mu, ventricoso-ampullaceis; stipite ad apicem luteo (“light cad-
mium” Ridgway) et atro-squamoso, fibroso, elongato; carne fracta caerulescente.
Habitatio: Ad folia delapsa ad basin arboris Augusto mense. In Liberia Africae Occi-
dentalis, prope Miamu. D. H. Linder 408 (FH).

Boletellus turbinatus (Snell) Sing. comb. nov.
Pirate I, Fic. 3.

The type from Missouri (FH) shows the spores to be deep honey color
with a fulvous tinge in the episporium, smooth but thick walled, the apical
portion attenuate and mostly subacute, often with an incomplete germina-
tive pore. These are typical Boletellus-spores in spite of their being smooth.
I found them 13.5—20 x 5.5—9.5 yw; Snell finds them up to 21 » long. I sus-
pect that the turbinate habit is merely incidental.

SPECIES IMPERFECTLY KNOWN

Boletus chrysenteron Bull. sensu Coker & Beers, The Boletaceae, p. 70, pl. 44 (be-
low), pl. 64, fig. 8. 1943 (non Fr.).

Discovered in North Carolina, this species agrees with the preceding one
in many ways, but differs in the more truncate apex of the spores and the
spore measurements as given by Coker and Beers. Material from Chapel
Hill, however, has the spores larger than given by these authors, viz.
(11.5)-13-17—(26) x 4.8-6.5 yw. This cannot be Bulliard’s plant, since
species with spore characters like those described and figured by Coker and
Beers do not occur in the environs of Paris. It is also not Fries’s Boletus
chrysenteron which equals Boletus communis Bull. sensu Coker & Beers
(d.c., p. 62). This latter name can not be accepted according to the rules of
nomenclature. If Boletus chrysenteron sensu Coker & Beers should turn
out to be different from Boletellus turbinatus, it must be renamed. How-
ever, we are inclined to think that the turbinate shape of the latter is an
accidental character of no taxonomic value and little if any constancy;
consequently these species may be synonymous.

SINGER: FLORIDA BOLETINEAE. I 135

Sect. Ixocephali Sing. sect. nov.
Superficiebus sterilibus carpophororum plus minusve viscidis; sporis longitudinaliter
vittatis; stipite haud subceraceo-lacunoso.

Characters of the section: see key (p. 107) and Latin description above.
The type, Boletellus singaporensis (Pat. & Baker) Gilbert (Boletopstis
singaporensis Pat. & Bak.), has a distinct annulus, but I think that evelate
forms like Boletellus jalapensis are close enough to be placed in this same
section. The section /xocephali is not represented in Florida.

Boletellus singaporensis (Pat. & Baker) Gilbert, Bolets, p. 107. 1931.
Boletopsis singaporensis Pat. & Baker, Journ. Straits Branch R. Asiatic Soc. No. 78:
69. 1918.

The species has a beautiful, distinct, tardily lacerate annulus which is
said to be viscid in fresh condition. The type (FH) shows a pretty species
with long stipe and thick hymenophore, well agreeing with the original de-
scription. The spores are comparatively short, 14-15 x 10.5-12.5 p, with
about 13 longitudinal wing-like lamellose ridges which are brown on mel-
leous ground, and 1.2-1.6 » high, and suddenly cut off at the apex, with a
globose oil-drop and asymmetrically inserted hilar tube, some of the ridges
not reaching the opposite end, not transversely veined, wall 0.8-1.1 » thick;
basidia 23-27 x 15-17 ,; cystidia subulate-fusoid with rounded tips, 45x
10.5 ». This species, under a temporary herbarium name, is represented in
Patouillard’s Herbarium (#4992).

Boletellus jalapensis (Murr.) Gilbert, Bolets, p. 107. 1931.
Ceriomyces jalapensis Murr., Mycologia 2: 248. 1910.

The type (NY) from Mexico has 16-18-(25) x 10-11-(13.5) u large,
deep melleous spores with 10-14 longitudinal ribs with scarce anastomoses
but usually once or twice forked ribs among them, the ribs 1-1.5 » high;
basidia 38—42 x 14-15.5 , 4-spored; cystidia not found but probably pres-
ent; hyphae without clamp connections; structure of the cuticle of the
pileus not clear, hyphae clampless, elongate, some claviform, some very
thin, yellowish-subhyaline, of variable diameter.

I have not seen the material determined as Boletogaster jalapensis by
Lohwag but judging from the picture and the description, it may be a long-
stemmed form of this species. This would extend the area of B. jalapensis
to South China, another indication of the close relationship between the
boletineous flora of North and Central America and Eastern Asia. It may,
however, turn out that Lohwag’s determination was only approximate, and
that the Chinese fungus is a third species of this section characterized by
its stipe which is twice as long as in typical B. jalapensis. As for the de-
scription of the ornamentation of the spores, I am not sure that Lohwag’s
observations are quite exact. If they are, it would indicate an additional
difference between this and the Chinese species. The spores of the Mexican
type have the projecting angles (corresponding to the ribs) when seen from
above, not semicircular but lamellose-rectangular, thinner than the distance

136 Fartowla, Vor. 2, 1945

they project from the surface of the wall proper, and their number, when
counted at one of the poles never reaches twenty.

Sect. Dictyopodes Sing. sect. nov.
Pileo tomentoso, haud viscido; sporis longitudinaliter costato-vittatis, elongatis;
stipite subceraceo-lacunoso et ex ea re reticulato vel Jongitudinaliter lamellato.

Characters of the section: see key (p. 107) and the above Latin diagnosis.
The type species is Boletellus Russellii (Frost) Gilbert (Boletus Russellii
Frost).

7. Boletellus Russellii (Frost) Gilbert, Bolets, p. 107. 1931.
Boletus Russellii Frost, Bull. Buffalo Soc. Nat. Hist. 2: 104. 1874.
Ceriomyces Russellii Murr., Mycologia 1: 144. 1909.
Boletogaster Russellit Lohwag in Handel-Mazzetti, Symb. Sin. 2: 56. 1937.
Frostiella Russellii Murr., Mimeogr. Contr. Herb. Univ. Fla. Agr. Exp. Sta.
Gainesville, p. 6. 1942.

Pirate I, Fic. 9.

Pileus “Sayal brown” and “wood brown” mixed, later about “avella-
neous,” strongly tomentose, becoming strongly furfuraceous to felty-
squamulose, dry or scarcely subviscid after prolonged rains, with smooth
margin which strongly projects beyond the hymenophore, appearing lobate
or crenate from above, pulvinate, 35-80 mm. broad. — Hymenophore
“naphthaline yellow,” becoming “barium yellow” to “strontian yellow,”
pores concolorous but with a slight pinkish hue when young, surrounded
by the sterile projecting margin which is pallid and reddening, tubes me-
dium long, 5-9 mm., adnate, becoming depressed around the stipe when
mature, pores rather small to medium wide, widest in the marginal row,
3 per mm., then becoming wider, most frequently 0.8 mm. wide, but some-
times up to 1 mm. in diameter; spore print “dark olive.” — Stipe “cacao
brown” to “walnut brown,” at least above, the ribs and mostly also the
entire base “vinaceous pink” to “buff pink,” with very high (about 2-3
mm.) lamellose longitudinal ribs which strongly anastomose, making the
surface appear conspicuously lacunose-favose, the single honeycomb of
varying length (1-10 mm.), the base usually smooth but often covered
with white to whitish yellow mycelial felt, solid, becoming slightly hollow,
slightly to strongly viscid outside, especially below, tapering at the base
into the ground, and also tapering upward in the upper one or two thirds,
or thickened at the base and gradually tapering to the apex, the basal por-
tion usually covered by rotten leaves, 63-80 x 14-20 mm. — Context whit-
ish to creamy pallid, stramineous-pallid, becoming eventually melleous in
the base of the stipe, and reddish under the cuticle, more so when wounded,
eventually almost ‘walnut brown” in the upper portion of the pileus, almost
unchanging on injury, rather hard in the stipe but the consistency of the
ribs almost waxy, soft-fleshy in the pileus; taste mild; odor almost none,
eventually slightly mawkish.

Spores 16-19.5 x 8-11 », melleous, ornamentation and wall brown, with
longitudinal, wing-like, Jamellose ridges which run from the apical pole to

SINGER: FLORIDA BOLETINEAE. I ey

the hilar tube, suddenly stopping at the apex, without the transverse veins
of Boletellus ananas, occasionally some ridges connected by a few scattered
anastomosing ridges, the longitudinal ridges numbering about 13 at the
poles, and projecting 1.8—2.1 »; basidia 25-35 x 13-16 p, 4-spored; cystidia
mostly fusoid-ampullaceous, often compressed, 45-70 x (6.5)—9.5-15 4p,
occasionally with a colored incrustation but wall and contents always
hyaline; ¢rama of the hymenophore truly bilateral, mediostratum slightly
darker and denser than the strongly diverging hyaline lateral stratum;
cuticle of the pileus consisting of multi-septate, erect chains of hyphae with
clampless septa, most members of these chains rather short to almost sub-
globose, but the terminal cell elongate (about 50 » long), cylindric or subu-
late; surface layer of the stipe consisting of a palisade of dermatocystidia
and abundant dermatopseudoparaphyses, also some basidia; the dermato-
cystidia of the young specimens hyaline to brown, ampullaceous, up to
80-100 x 10.8 »; dermatopseudoparaphyses up to 11 p» broad; basidia
4-spored, about 46x 13.2 »; hyphae of the context without clamp con-
nections.

Chemical reactions. KOH on the surface of the pileus chestnut or ful-
vous; on the context of the upper portion of the pileus brown; on the con-
text of the stipe slowly brownish; NH,OH on the surface of the pileus
somewhat deeper and more brick colored; on the context of the upper por-
tion of the pileus red; on the context of the stipe slowly reddish; FeSO,
on the surface of the pileus deep olive; on the context of the pileus and
stipe negative; formol negative; phenol negative.

Habitat: Under oaks, in Florida under Quercus laurifolia in high ham-
mock formations on the soil amongst leaves. Summer.

Distribution: All along the Atlantic Coast from New England to Central
Florida and west to Wisconsin (according to Murrill) and Mississippi. In
Florida not reaching the tropical part of the peninsula.

Material studied: Fla.: Alachua Co., in and near Gainesville, R. Singer F 2500,
F 2500a (FH); Lake Co., Eustis, on wood of Sabal, R. Thaxter (FH) ; — Vt.: type of
Boletus Russellii, Brattleboro, in damp woods, 1865, C. C. Frost (FH).—N. H.: Shel-
burne, Aug. 1891, Farlow (“form with scaly top”) (FH); Chocorua, September 1917,
Farlow (FH); Intervale, August 1901, Thaxter 3473 (FH).—Mass.: Arlington,
‘HH. M. N. (FH): Middlesex Fells, August 1897, Farlow (FH); Canton, August 1925,
D. H. Linder (FH).—N. C.: Glassy Mts., Flat Rock, August 1934, A. S. Rhoads (det.
W. A. Murrill) F 9385 (FLAS).

B. Russellii is unmistakable for anyone who has once collected it. Its
history shows clearly that there was doubt about its specific taxonomy only
once when Lloyd claimed its identity with B. betula, an error subsequently
corrected by W. G. Farlow. There was, however, considerable fluctuation
in its generic position. I cannot see any reason for splitting the genus
Boletellus into several genera, but if it is done (which would make it nec-
essary to go much farther in breaking up the genus than we do now in
dividing the boletaceous genera and the agarics) it should be done accord-
ing to the sections outlined in the key p. 107. However, Frostiella is not a

138 FarLowIiA, Vor. 2, 1945

valid name since it has been published without a Latin diagnosis, and in
the sense of Murrill covers several sections of Boletellus; thus it becomes a
synonym of the older name Boletogaster Lohwag. There is a fine colored
picture in Jcones Farlowianae, pl. 80, and a good photograph has been pub-
lished in Coker and Beers, The Boletaceae, pl. 48. Macllvaine pl. 118, fig.
2 is likewise correct. Elrod and Blanchard (Mycologia 31: 701, fig. 2, E.
1939) published a photomicrograph showing the structure of the trama.
The difference of the picture obtained as compared with our results is due
to insufficient consistency on the part of these authors in their choice of
corresponding and comparable stages of development. In young hymeno-
phores, the trama of practically all Boletineae is more or less bilateral, and
all Strobilomycetaceae have the truly bilateral trama of the Boletus-type.

SPECIES NOT KNOWN TO THE AUTHOR

Boletellus costatus (Rostrup) Sing., Ann. Myc. 40: 18. 1942.
Boletus costatus Rostrup, Botan. Tidskr. 24: 357. 1902.

Described from Thailand, tropical Asia, this probably belongs in this
section.

Sect. Allospori Sing. sect. nov.
Pileo viscido; sporis spinis exosporio immersis ornatis; stipite subceraceo-lacunoso
et ex ea re reticulato vel longitudinaliter lamellato.

Characters of the section: see key p. 107, and Latin diagnosis above.
The type species, B. betula (Schw.) Gilbert, is so far the only species of
this section. It has not been found in Florida.

Boletellus betula (Schw.) Gilbert, Bolets, p. 108. 1931.

A North American species, reported from many eastern and middle
western states, has also been called Ceriomyces betula Murr., Frostiella
betula Murr., and Boletus Morgani Peck. It was initially described as
Boletus betula by Schweinitz, Schr. Naturf. Ges. Leipzig 1: 90. 1822. I
have studied specimens from several localities, and the results of my micro-
scopical analyses follow:

Spores 16—21.5 x 8.5-10 p, deep ochraceous-melleous, with suprahilar
applanation, ellipsoid-fusoid, with isolated punctiform warts if the upper
surface of the spore is focussed upon, but actually the wall perforated by
cylindric thin spines which are darker than the wall and hardly pass be-
yond its outer surface; basidia 25-40 x11.6-16.5 p, 4-spored; cystidia
present; the pellicle consists of a palisade of dermatocystidia (100 x 8 y)
and more basidium-like bodies; the surface of the stipe consists of a pali-
sade of sterile elements.

SPECIES INCERTAE SEDIS

Strobilomyces paradoxus Mass., Bull. Misc. Inf. Bot. Gard. Kew 1909 (5): 209. 1909.
Boletellus paradoxus Gilbert, Bolets, p. 107. 1931.

SINGER: FLoRIDA BOLETINEAE. I 139

This species has been described from Singapore. I have seen no speci-
mens.

Boletus Rostrupii Syd., Ann. Myc. 1: 177. 1903.
Boletus lacunosus Rostr., Bot. Tidskrift, 24: 357. 1902 (non Cooke).

This species has been described from Thailand. I have seen no speci-
mens but it seems reasonable to assume that it belongs in the Strobilomyce-
taceae.

Strobilomyces fasciculatus Cooke, Grevillea 20: 4. 1891.

This. species from Australia may or may not belong in the Strobilomyce-
taceae. No specimens have been examined by the writer.

Strobilomyces ligulatus Cooke, Grevillea 20: 4. 1891.
Boletellus ligulatus Sing., Ann. Myc. 40: 18. 1942.

Described from Australia, this seems to be a Boletellus, provided the
description is correct. However, the spore ornamentation (if there is any)
has not been mentioned by Cooke.

Strobilomyces rufescens Cooke & Mass., Grevillea 18: 5. 1889.

The description of this Australian species reads much like that of Stro-
bilomyces pallescens which is a synonym of Boletellus ananas.

Boletellus pustulatus (Beeli) Gilbert, Bolets, p. 107. 1931.
Boletus pustulatus Beeli, Bull. Soc. Roy. Bot. Belg. 58: 211. 1926.

Described from Belgian Congo, this species has been transferred by Gil-
bert to Boletellus, and, judging from the original description, correctly so.
If the spores actually are as small as indicated by Beeli, one would expect
this species to belong in the section Chrysenteroidei, though its color is
more suggestive of section Ananae.

Phyllobolites Sing., Ann. Myc. 40: 59, 1942.

No representatives occur in Florida.

Harvarp UNIVERSITY
CAMBRIDGE, Mass.

140 FARLOWIA, VoL. 2, 1945

EXPLANATION OF PLATE I

All drawings are made with the aid of a camera lucida from dried material mounted
in KOH-phloxine (basidia and cystidia) or either ammonium hydroxide or Melzer’s
reagent (spores) under oil immersion lens. Magnification approximately x 1050.

1. Strobilomyces confusus Sing. Spores, basidium; from the type specimen.

2. Strobilomyces floccopus (Vahl ex Fr.) Sacc. Spores, basidium; specimen from
Harvard, Mass.

Boletellus turbinatus (Snell) Sing. Spores, basidium; from the type specimen.

Porphyrellus subflavidus (Murr.) Sing. Spores; from authentic material.

Porphyrellus gracilis (Peck) Sing. Spores, basidium; from authentic material.

Strobilomyces velutipes Cooke. Spores, basidium; from the type material.

Boletellus ananas (Curt.) Murr. Spores, basidium; specimen from North Florida.

Strobilomyces pterosporus Sing. Spores, basidium; from the type specimen.

Boletellus Russellii (Frost) Gilb. Spores, basidia and cystidia.

Boletellus Linderi Sing. Spores, basidium; from the type specimen.

© 60 GT OF

a

141

SINGER: FLorIpA BOLETINEAE. I

PLATE I

Be

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‘SEYMOUR, A. B. Host. Index of the Fungi of North America.
Te are, Clot oe 18

-THAXTER, R. Contribution towards a Monograph of the Laboul-
beniaceae. (Mem. Amer. Acad. Arts & Sci.) Unbound.

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TOG ee ee ee
go Se ee ee ee

TM Ae ae ee ee ee
pe ee

THAXTER, R. Entomophthoreae of the United States. 1888. |
(Mem. Bost. Soc. Nat. Hist.) Unbound . ... .. . 3.00

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FARLOWIA
A JourNAL oF CryptoGAMic BOTANY | -

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CONTENTS OF VOL. 2, NO. 2

A Taxonomic anp Ecoxocica, Stupy or Somr DiaToMs FRoM THE Pocono PLa-
TEAU AND ADJACENT REcions. By Ruth Patrick. . . . . ©.) . 143°

THE BOLETINEAE OF. FLORIDA wiTH NoTES oN EXTRALIMITAL Spectres. II. THE
BOLETACEAE (GYROPOROIDEAE). By Rolf Singer. . . . . . . . 223

Vol. 2, No. 1, was issued March 22, 1945.

AVLVIALSITATTV ahd
peor plnoys JYIALSATATIVAKd ‘42439 | ‘1E8t ‘duo

‘snuadqns WY NICIHLNVNHOF
pear plnoys snusdqns OTH ENV NHOF Suipeey ‘21 ‘| “991 ‘d uO

“SPOT ‘Gur “(Z) 2 vimopsny “SUOYEI] OUDD0g : yoy 4“
VLVUUF

FARLOWIA

A JOURNAL OF CRYPTOGAMIC BOTANY

Noles Jury, 1945 Now Z

A TAXONOMIC AND ECOLOGICAL STUDY OF SOME
DIATOMS FROM THE POCONO PLATEAU
AND ADJACENT REGIONS

RutH PATRICK

The Pocono plateau, which is located in the northeastern section of Penn-
sylvania, is part of the general Appalachian system. Geologically it is mostly
sandstone of the Pocono and Catskill formation which was laid down in the
late Devonian and early Carboniferous times. Here and there are present
local limestone outcrops.

In this study the diatoms were obtained from regions adjacent to as well
as in the Pocono plateau. Various types of bodies of water in Carbon,
Lackawanna, Luzerne, Monroe, Northampton, Pike, and Wayne County
were included. Thus localities on the Martensburg shale, Helderberg, and
Chemung formations were also studied.

In this region there are many so-called lakes. From the limnological
standpoint these are not lakes but ponds, for they have no thermocline.
They are formed in one of two ways; either they are of glacial origin or are
dammed up streams. Immediately after the Civil War there was much lum-
bering in this region. The dams were constructed in the streams to form
storage basins for the logs until the spring thaws. At that time they would
be floated down stream. After these lumbering operations had ceased these
ponds were used as a source of ice. During the winter, ice would be cut and
stored in ice houses which were built on the shores of the ponds. With the
advent of artificial ice, this business ceased and summer resorts were built
about the ponds. :

A description of the various localities from which the diatoms were col-
lected is given below. The ecological information concerning these habitats
is to be found in the ecological discussion.

LACKAWANNA COUNTY
2044 Indian Lake, from squeezings of surface vegetation, Hodge, 7 July 1941
LUZERNE COUNTY
2042 Harvey Lake, from squeezings of the bottom vegetation, in quiet shallow
water, Hodge, 7 July 1941
2043 Harvey Lake, from squeezings of vegetation in running water at the outlet,
Hodge, 7 July 1941

143

144

FArLowiA, VoL. 2, 1945

MONROE COUNTY

262

263

2010
2011
2012
2013
2014
2015
2016
2017
2018
2019
2020
2025
2031

2032

2033
2034

2035°

2036

2037
2038
2039
2040
2045
2046
2047
2049
2051
2056

2065

Pocono Lake, a composite collection made by Mr. F. J. Myers during the
summer of 1940.

Jaggie’s Bog, Shulze

Jaggie’s Bog, Shulze

South Pocono Lake, in a small bay one foot deep, on rocks and on moss
and weeds growing on bottom, Patrick, 8 July 1941

South Pocono Lake, squeezings from Sphagnum in 18 inches of water,
Patrick, 8 July 1941

A small stream from springs, running under road to Davy’s Run, shallow,
filled with Sphagnum, Patrick, 8 July 1941

Shallow ditch along road to Davy’s Run, very cold, in shade, Patrick,
9 July 1941

Davy’s Run, beaver dam, surface mud and scum around weeds, growing in
3-4 inches of water, Patrick, 9 July 1941

Davy’s Run, in running water below beaver dam, attached or enmeshed in
Fontinalis antipyretica, Patrick, 9 July 1941

Davy’s Run, surface ooze in 3-4 inches of water, about 50 feet above beaver
dam in shade, Patrick, 9 July 1941

Davy’s Run, open sun, running water, on or enmeshed in weeds, Patrick,
9 July 1941

Davy’s Run, deep shade, surface of mud, still, shallow water, Patrick,
9 July 1941

Davy’s Run, deep shade, rapid water, squeezings from Hypnum and Fonti-
nalis, Patrick, 9 July 1941

Lost Lake, squeezings from Sphagnum, A. Turner, 5 July 1941
Raymondskill Falls, surface scum on rocks in pool in falls, Patrick, 2 July 1941
Mud Run, squeezings from moss in spillway, Hodge, July 1941

Tobyhanna Creek, shallow water near flat rocks on side of creek known as
“Lover’s Leap”, Patrick, 8 July 1941

Tobyhanna Creek, near bank of creek, Patrick, 8 July 1941

Pocono Lake, on stones in 1 foot of water at Emlen dock, Patrick, 30 June 1941
Pocono Lake, from squeezings of Elodea, Dicranium, and Hypnum growing
in 1 foot of water at Emlen dock, Patrick, 30 June 1941

Pocono Lake, from squeezings of Myriophyllum in 2 feet of water at Emlen
dock, Patrick, 30 June 1941

Spring run near Emlen cottage, squeezings from weeds, C. Myers, 6 July 1941
Pocono Lake, plankton taken by tow net, Patrick and Hodge, 7 July 1941
Lake Tamaque, surface scum, Hodge, 7 July 1941

Lake Tamaque, scrapings from log, Hodge, 7 July 1941

Popomoning Lake, from squeezings of Elodea, Hodge, July 1941

Wier Lake, squeezings from surface vegetation, Hodge, July 1941

Wier Lake, squeezings from bottom vegetation, Hodge, July 1941

Trout Lake, squeezings from Spirogyra, Hodge, July 1941

Mountain Lake, from squeezings of surface vegetation, Hodge, July 1941
Davy’s Run, full sun, in running water, enmeshed in Fontinalis antipyretica,
Patrick, 8 July 1941

Pocono Lake, scrapings from logs supporting Emlen’s dock, Patrick, 10 July
1941

NORTHAMPTON COUNTY

2050

Green Pond, from squeezings of vegetation, Hodge, July 1941

PIKE COUNTY

1

Z
3

Greeley, squeezings from Fontinalis in brook running through swamp,
J. Burke, 4 July 1941

Greeley, squeezings from Sphagnum in swamp, J. Burke, 4 July 1941
Shohola Falls, from algae at top of falls, J. Burke, 4 July 1941

PATRICK: Pocono DIATOMS 145

4 Shohola Falls, from moss at top of falls, in swift water, J. Burke, 6 July 1941
5 Lake Wallenpaupek tributary passing under Route 507, J. Burke, 5 July 1941
7-9 Shohola Falls, scrapings from dripping rock face, side wall of gorge, above
concrete bridge crossing road below falls, J. Burke, 5 July 1941
10 Shohola Falls, from algae in brook at head of falls near pond, J. Burke,
6 July 1941
11 Shohola Falls, from moss in brook above falls, near pond, J. Burke, 6 July
1941
12 Shohola Falls, edge of pond above falls, J. Burke, 6 July 1941
13 Shohola Falls, scrapings from dripping wall in small glen at head of falls,
J. Burke, 6 July 1941
14 Shohola Falls, scrapings from face of rock at side of falls, J. Burke, July 1941
15 Shohola Falls, gelatinous algae on rock wall facing falls, J. Burke, 6 July 1941
127-130 J. Shulze (no other information except Pike County given)
1621-1623 Bushkill Creek, in spring opposite head of island, J. Shulze. (I cannot be
sure whether this locality is in Pike or Monroe county.)
2188-90 J. Shulze (no other information except Pike County given)
WAYNE COUNTY
2021 Angels, Mill Creek, in cool, shallow, swift water on rocks in middle of creek,
J. Burke and R. Patrick, 5 July 1941
2022 Angels, small tributary of Mill Creek, deep shade, slow running, 2-5 inches
deep, J. Burke and R. Patrick, 5 July 1941
2023 Angels, Mill Creek tributary, deep shade, from surface of leaves in spring
bubbling through sand, J. Burke and R. Patrick, 5 July 1941
2024 Angels, scraping from surface of leaves in a very shallow pool or puddle,
J. Burke and R. Patrick, 5 July 1941
2026 Angels, Mill Creek, full sun, growing on stones in swift, shallow water,
R. Patrick and J. Burke, 5 July 1941

In this study it was thought wise to consider all obtainable collections
that had ever been made in this region. Some-of these collections are quite
old and no information is available concerning. the ecological conditions
which existed when the collections were made. Because of this most of the
ecological studies are based on a few of the most diversified ponds which
were selected for this purpose. Much of the ecological data was obtained
by Mr. Frank J. Myers who studied the rotifers present in these bodies of
water (Proc. Acad. Nat. Sci. Phila. 94: 251-286. 1942). The information
concerning the calcium, bicarbonates, and color of the water was furnished
by Dr. Thomas Edmondson, Woods Hole Oceanographic Laboratories.

In the study of all collections, hyrax mounts were made and the species
identified were ringed by means of a diamond marker. In the taxonomic
discussion under each species the localities are given in which the species
were found and the number in parenthesis indicates the slide in the her-
barium of the Academy of Natural Sciences of Philadelphia on which the
identification was made.

The materials prepared for the ecological studies were made from a quan-
titative standpoint. The original amount of material collected in each case
was as nearly as possible the same. If collections were made for epiphytic
diatoms, one-half pint (236 c.c.) of weeds or debris was collected and
squeezed. The drippings were retained and concentrated, and one half of
this concentrate was cleaned. The cleaned material in each case was diluted

146 FARLowIA, VOL. 2, 1945

to a common dilution factor. This solution was then well shaken and im-
mediately one tenth of a cubic centimeter was withdrawn and deposited on
tonic and encrusted forms.

In this study five terms have been used to denote the relative abundance
of the species found. The slides, with the aid of a mechanical stage, were
methodically studied. In most cases a 3 mm. objective was used in making
these counts. However, when large forms were being studied a 16 mm.
objective was used. “Rare” indicates that only a very few specimens were
found on the slides studied. “Frequent” indicates that one or two specimens
were present in about fifty per cent of the fields of the microscope. ‘Very
frequent” indicates that an average of two or three specimens were found in
each field. ‘“‘Common or abundant” indicates that several specimens were
found in each field.

The author is particularly indebted to Mr. Frank J. Myers who furnished
much valuable information which he had obtained concerning these bodies
of water, and who also made available the equipment of his summer labora-
tory at Pocono Lake Preserve. The author also wishes to express her appre-
ciation to Dr. Thomas S. Stewart who made the photographs included in
the plates; to Dr. Thomas Edmondson who made the chemical analysis
of the water; to Miss Jane Stenz who made the slides for this study; to
Mr. Joseph Burke who made available his collections from Pike County; to
Dr. Francis W. Pennell for his help with Latin descriptions; and to Miss
Hilda F. Harris of the Farlow Herbarium, to Mrs. Gertrude Hess of the li-
brary of the American Philosophical Society, and to Mr. Frank Patrick who
made available additional literature necessary for this study.

MELOSIRA AcarpH

A. Sulcus absent, in girdle view frustule very finely striated ... M. varians
A. Sulcus present
B. Sulcus a distinct concave throat or channel
C. Lives in running water on rocks or in moss; in valve view, puncta ar-

ranged in definitely radiating rows ........... M. roeseana and var.
C. Typically a pelagic form .................0.. M. ambigua
B. Sulcus a narrow groove ......... 0c cece eee eens M. italica

Melosira ambigua Miiller, 1905, Bot. Jahrb. Syst. Pflanzengesch. u. Pflanzengeog.
34: 283. pl. 4, figs. 9, 10.

For a discussion of this species see my paper in Proceedings of the Acad-
emy of Natural Sciences of Philadelphia 92: 195. 1941.

This species was very frequent in a gathering made by Shulze in Pike
County. Unfortunately we have no habitat data concerning this collection.
Krieger (1929) states it is found in waters with a pH of 5.5-7.8.

Monroe County: Pocono Lake Preserve, Pocono Lake, 2036 Patrick
(A-2148), 2038 Patrick (A-2152); Tobyhanna Creek, 2033 Patrick (A-
2142); Trout Lake, 2047 Hodge (A-2172). Pike County: 127-128 Shulze
(A-2215-17).

Patrick: Pocono DIATOMS 147

Melosira italica (Ehr.) Kiitz., 1844, Bacill., p. 55. pl. 2, fig. 6.
Gaillonella italica Ehr., 1836, Ber. K. Akad. Wiss. Berlin 1836: 53.

This species was common in the pond above Shohola Falls.

Monroe County: Raymondskill Falls, 2025 Patrick (A-2131); Jaggie’s
Bog, 262 Shulze (A-2223). Pike County: 130 Shkulze (A-2221); Shohola
Falls, 10-11 Burke (A-2201-02), 12 Burke (A-2204), 14 Burke (A-2209).

Melosira roeseana Rabh., 1852, Algarum Europaea no. 382, 1853, Sussw. Diat., p. 13.
pl. 10, suppl. fig. 5.

It is hard for me to be sure of the difference between M. roeseana and
M. roeseana v. epidendron. I have checked my specimens with those of
Rabenhorst’s on Rabh. Alg. Sach. Resp. Mitteleurop. no. 383 and with
Van Heurck’s Type Slide no. 465. I find that M. roeseana on these slides
has the ridges, perhaps spines, on the margins of the valve arranged at
regular intervals, usually between each stria. In Melosira roeseana v. ept-
dendron according to Van Heurck’s figure (Syn. Diat. Belgique, fl. 89,
figs. 17-18) and in Van Heurck’s Type Slide no. 467 the spines are irregu-
larly arranged. Also the structure of variety epidendron seems to be
somewhat coarser.

This species was present only in the locality given below.

Pike County: Shohola Falls, 15 Burke (A-2211).

Melosira roeseana v. epidendron (Ehr.) Grun. apud Van Heurck, 1882, Syn. Diat.
Belgique, pl. 89, figs. 17, 18.
Sphenosira epidendron Ehr., 1848, Ber. K. Akad. Wiss. Berlin 1848: 219.

In collection no. 7 of Burke were found specimens which differ in number
of striae in 10 » from the figure in Van Heurck’s Synopsis Diatomées Bel-
gique. In girdle view the striae are 17—18 in 10 » instead of 14 in 10 p.

This variety was of frequent occurrence in the collections cited.

Pike County: Shohola Falls, 7-9 Burke (A-2196-98).

Melosira varians Ag., 1827, Flora (Bot. Zeit.) 10 (2): 628.

This species though present in two habitats was not of frequent occur-
rence in either.

Monroe County: Raymondskill Falls, 2025 Patrick (A-2131). Pike
County: Shohola Falls, 4 Burke (A-2192).

CYCLOTELLA Ktrz1nc

A. Valves circular, center of valve marked by radiating lines producing a star-shaped
DAtLERN etter ce. cet hte unia eerie yeas aes C. stelligera

A. Center of valve otherwise marked or clear

B. Marginal striae interspersed with thickenings, middle field of valve filled with
radiating punctae’ ...28 o.4... eos C. comta v. radiosa

B. Marginal striae without such thickenings C. Meneghiniana v. rectangulata

Cyclotella comta v. radiosa Grun. apud Van Heurck, 1882, Syn. Diat. Belgique, i.
92, fig. 23, text (1885) p. 214.

148 FARLOwWIA, VOL. 2, 1945

Though only found in one habitat it was common there. Hustedt states
that it is especially found in sub-alpine lakes. It seems to be found most
commonly in this country in habitats north of or above the frost line.

Luzerne County: Harvey Lake, 2042-43 Hodge (A-2160-63).

Cyclotella Meneghiniana y. rectangulata Grun. apud Van Heurck, 1882, Syn. Diat.
Belgique, pl. 94, figs. 17-19.

Pike County: Shohola Falls, 4 Burke (A-2192), 10-11 Burke (A-2200-
02).
Cyclotella stelligera (Cl. and Grun.) Van Heurck, 1882, Syn. Diat. Belgique, pl. 94,
figs. 22-27.

Cyclotella Meneghiniana v. ? stelligera Cl. and Grun. apud Cl., 1881, K. Sv. Vet.
Akad. Handl. 18 (5): 22.

This species was found only once and then it was rare.
Monroe County: Trout Lake, 2049 Hodge (A-2174).

ODONTIDIUM Ktrzinc emenp. BESSEY
Odontidium anceps Ehr., 1843, Abhandl. K. Akad. Wiss. Berlin 1841: 127.

This species was rare in several localities. It was common in very shal-
low water, on the surface of leaves at Angels, Wayne County. The tempera-
ture of the water was between 50-60° F.

Pike County: 2188-90 Shulze (A-2234-36), 128 Shulze (A-2217); Lake
Wallenpaupek tributary, 5 Burke (A-2195). Wayne County: Angels, 2021
Burke and Patrick (A-2123), 2023-24 Burke and Patrick (A-2127-29).

Odontidium hiemale v. mesodon (Ehr.) Grun., 1862, Verh. Zool. Bot. Ges. Wien
12: 357.

Fragilaria mesodon Ehr., 1839, Abhandl. K. Akad. Wiss. Berlin 1838: 57. pl. 2, fig. 9.

This variety was very frequent in shallow running water at Angels, and
in squeezings from the bottom vegetation of shallow water in Wier Lake.

Monroe County: Raymondskill Falls, 2025 Patrick (A-2131); Wier
Lake, 2046-47 Hodge (A-2169-70). Pike County: 127 Shulze (A-2215);
Lake Wallenpaupek Tributary, 5 Burke (A-2195); Shohola Falls, 15 Burke
(A-2211). Wayne County: Angels, 2021-24 Burke and Patrick (A-2123-
29), 2026 Burke and Patrick (A-2134).

TABELLARIA EHRENBERG

Tabellaria fenestrata (Lyngb.) Kiitz., 1844, Bacill., p. 127. pl. 17, fig. 22; pl. 18, fig.
2; pl. 30, fig. 73.

Diatoma fenestratum Lyngbye, 1819, Tent. Hydrophyt. Danicae, p. 180. pl. 61, fig. 3.

From this study it would seem that this species preferred shallow water.
It is of very frequent or common occurrence in Mud Run, Pocono Lake,
Tobyhanna Creek and the pond above Shohola Falls, in scrapings from
dripping rocks.

PATRICK: Pocono DIATOMS 149

Lackawanna County: Indian Lake, 2044 Hodge (A-2164). Luzerne
County: Harvey Lake, 2042 Hodge (A-2161). Monroe County: Pocono
Lake Preserve, Mud Run, 2031 Hodge (A-2138); Pocono Lake, 2034-36
Patrick (A-2144—48), 2038 Patrick (A-2152), 2065 Patrick (A-2182),
F. Myers (A-2213); South Pocono Lake, 2010-11 Patrick (A-2101-04) ;
Tobyhanna Creek, 2032-33 Patrick (A-2140-42); Lake Tamaque, 2039
Hodge (A-2154); Wier Lake, 2046 Hodge (A-2168). Pike County: 2188-
90 Shulze (A-2233-36), 127-30 Shulze (A-2215-21); Greeley, 1 Burke
(A-2184); Shohola Falls, 10-14 Burke (A-2201-10).

Tabellaria floceulosa (Roth) Kiitz., 1844, Bacill., p. 127. pl. 17, fig. 21.

Conferva flocculosa Roth, 1797, Catalecta Botanica Fasc. 1, p. 192. pl. 4, fig. 4; pl.
5, fig. 6. (This reference was checked for me by Miss Harris of the Farlow Library.)

This species was very frequent or common in Pocono Lake, Mud Run,
Tobyhanna Creek, Davy’s Run and Shohola Falls. It seems to prefer some-
what more acid conditions than the preceding species and is often found
associated with moss. Krieger (1929) states it prefers pH 3.5—-5.5.

Monroe County: Davy’s Run, 2014-19 Patrick (A-2109-19), 2056 Pat-
rick (A-2180); Jaggie’s Bog, 262 Skulze (A-2223); Pocono Lake Preserve,
Mud Run, 2031 Hodge (A-2138); Pocono Lake, 2034-36 Patrick (A-2144—
48), 2038 Patrick (A-2152), 2065 Patrick (A-2182), F. Myers (A-2213);
Tobyhanna Creek, 2032-33 Patrick (A-2140-42); South Pocono Lake,
2010 Patrick (A-2102); Popomoning Lake, 2045 Hodge (A-2166); Ray-
mondskill Falls, 2025 Patrick (A-2131); Trout Lake, 2049 Hodge (A-
2174); Wier Lake, 2047 Hodge (A-2170). Pike County: 2188-89 Shulze
(A-2233-35); Bushkill Creek, 1621-23 Shulze (A-2227-31); Greeley, 1—2
Burke (A-2184—-88); Shohola Falls, 3-4 Burke (A-2190-92), 10-14 Burke
(A-2200-10). Wayne County: Angels, 2021 Burke and Patrick (A-2124).

MERIDION AcaArpH

Meridion circulare (Grev.) Ag., 1831, Consp. Crit. Diat., p. 40.
Echinella circulare Grev., 1822, Mem. Werner. Nat. Hist. Soc. 4 (1): 213-15. pl. 8,
fig. 2.

The specimens most commonly found in this study differ from the type
in having the apex or head end of the valve acute instead of rounded. This
species reached its best development in shallow, flowing, somewhat acid
waters; — ina tributary of Mill Creek, in a brook running through a swamp
at Greeley, at Jaggie’s Bog, and at Shohola Falls.

Luzerne County: Harvey Lake, 2042 Hodge (A-2161). Monroe County:
Davy’s Run, 2014-15 Patrick (A-2109-11), 2018-19 Patrick (A-2117-
19); Jaggie’s Bog, 262 Shulze (A-2223); Pocono Lake Preserve, Mud Run,
2031 Hodge (A-2138); Tobyhanna Creek, 2033 Patrick (A-2142); Trout
Lake, 2049 Hodge (A-2174); Wier Lake, 2046-47 Hodge (A-2169-70).
Pike County: 128 Shulze (A-2217), 130 Shulze (A-2221); Bushkill Creek,
1621-23 Shulze (A-2227-31); Greeley, 1 Burke (A-2184); Shohola Falls,

150 FARLowIA, VoL. 2, 1945

3-4 Burke (A-2190-92), 10 Burke (A-2200), 14 Burke (A-2208). Wayne
County: Angels, 2021-22 Burke and Patrick (A-2123-25), 2026 Burke and
Patrick (A-2134).

Meridion circulare v. constricta (Ralfs) Van Heurck, 1881, Syn. Diat. Belgique, pl.
51, figs. 14, 15, text (1885) p. 161.

Meridion constrictum Ralfs, 1843, Ann. Mag. Nat. Hist. 12: 458. pl. 18, fig. 2.

This variety was found most often in Wayne County. As contrasted with
the species the only habitat in which it was very frequent was from scrap-
ings of dead leaves in a small, very shallow puddle at Angels.

Monroe County: Davy’s Run, 2016 Patrick (A-2113); Raymondskill
Falls, 2025 Patrick (A-2131). Pike County: 127 Shulze (A-2215); Gree-
ley, 2 Burke (A-2188); Shohola Falls, 11 Burke (A-2203), 12 Burke
(A-2204). Wayne County: Angels, 2021-24 Burke (A-2124-29),

FRAGILARIA LyNncBYE

A. Valves united to each other at middle portion of valve, thus having the ends of the
valve separate and free ... 2.2.0. eee cece ee euee F. crotonensis
A. Valves forming bands by other methods
B. Axial area linear and narrow, sometimes lacking or indistinct, central area
lacking. See B’ and B”
C. Valve constricted at center: ..éiccc4esss0xcieeens F. undata v. lobata
C. Valve not constricted

D. Valve elliptic or linear with rounded ends, striae about 11 in
LO Wh gsi a ane ew oiob ee bea be eee cones F. pinnata

D. Valve linear or lanceolate in shape, striae more numerous
E. Valve capitate
F. Some of the striae not opposite each other on
either side of the pseudoraphe thus producing an
irregular effect ............. F. bicapitata
F. Striae regular, not as above ..F. virescens var.
E. Valve not capitate ..... acerca ae: F. virescens
B’. Axial area linear and narrow, central area present
G. Central area bilateral, valve constricted in the middle
F. capucina var.
G. Central area unilateral, valve swollen on one side F. Vaucheriae
B”. Axial area more or less lanceolate, wider at center than at ends of valve
Hi, Striae very coarse; 6-8 In 10 @ ..ccncvavesseeave F. leptostauron
H. Striae much finer, 14-17 in 10 uw
I. Striae marginal
J. Valve elliptical ................... F. elliptica
J. Valve lanceolate with rostrate ends .F. brevistriata
I. Striae longer, valves linear or lanceolate ...F. construens

Fragilaria bicapitata A. Mayer, 1917, Denks. K. Bayer. Bot. Ges. 13: ns. 7: 21. pl. 1,
fig. 26.

This species was only once of very frequent occurrence and that was in

PATRICK: Pocono DIATOMS Tha

Wier Lake in association with bottom vegetation in shallow water. The
pH was 6.6.
Monroe County: Wier Lake, 2046-47 Hodge (A-2169-70).

Fragilaria brevistriata Grun. apud Van Heurck, 1881, Syn. Diat. Belgique, pl. 45,
fig. 32, text (1885) p. 157.
This species was rare in the two localities in which it was found.
Monroe County: Wier Lake, 2046 Hodge (A-2169).

Fragilaria capucina v. mesolepta Rabh., 1864, Flora Europaea Algarum, p. 118.

This variety was rare in the localities given below.

Monroe County: Popomoning Lake, 2045 Hodge (A-2166); Mountain
Lake, 2051 Hodge (A-2178). Pike County: 2189 Shulze (A-2235), 128
Shulze (A-2217).

Fragilaria construens (Ehr.) Grun., 1862, Verh. Zool. Bot. Ges. Wien 12: 371. pl. 7,
fig. 10.
Staurosira construens Ehr., 1843, Abhandl. K. Akad. Wiss. Berlin, 1841: 424.

This species was found only in Pocono Lake, and then it was rare.
Monroe County: Pocono Lake Preserve, Pocono Lake, 2034 Patrick
(A-2144), 2035 Patrick (A-2146), F. Myers (A-2213).

Fragilaria crotonensis Kitton, 1869, Science Gossip 1869: 110. fig. 81.

This species was very frequent in one locality. It was probably washed
into the brook from the lake.
Pike County: Shohola Falls, 10 Burke (A-2200).

Fragilaria elliptica Schuman, 1867, Schrift. Phys. Okon. Ges. Konigsburg 8: 52. pl. 1,
fig. 5.

This species was common in one collection: squeezings from littoral bot-
tom vegetation in Lake Wier.

Monroe County: Wier Lake, 2047 Hodge (A-2170). Pike County: tribu-
tary of Lake Wallenpaupek, 5 Burke (A-2195).

Fragilaria leptostauron (Ehr.) Hustedt, 1931, in Rabenhorst’s Krypt.-Flora, 7(2):
153. fig. 668a-f.
Biblarium leptostauron Ehr., 1854, Mikrogeologie, pl. 5, fig. II, 24, pl. 12, figs. 35, 36.

This species was rare in the three habitats.
Pike County: Lake Wallenpaupek tributary, 5 Burke (A-2195). Wayne
County: Angels, 2021-22 Burke and Patrick (A-2124-25).

Fragilaria pinnata Ehr., 1843, Abhandl. K. Akad. Wiss. Berlin 1841: 415.

This species was rare in occurrence.
Pike County: Lake Wallenpaupek tributary, 5 Burke (A-2195).

Fragilaria undata v. lobata var. nov.

Valvae constrictae ad medium, longae 20-54 mu, apicibus porrectis, pseudoraphe nulla,
striis transversis tenuibus 16-17 in 10 uw.

152 FaRLowlA, Vor. 2, 1945

William Smith (Ann. Mag. Nat. Hist. 2d ser. 15: 7. pl. 1, fig. 7) first
refers to this variety as F. undata v. y. What may be this variety, though
we cannot be sure as only the outline is shown and Ehrenberg questions it
being in the genus Fragilaria, is given as F. constricta Ehr., 1843, Abhandl.
K. Akad. Wiss. Berlin 1841: 415. pl. 1, fig. I, 21; pl. 3, fig. VI, 10. Hustedt
has placed in synonymy the specimens of W. Smith under Ehrenberg’s
name, Fragilaria constricta, and has clearly stated in his description that
there are horns or spines on the margins of the valve. He has cited as
reference Tempere and Peragallo, 2d Ed. Diatomées du Monde Entier nos.
831, 868, 907, 939. I have checked these slides and also one from Christiana
(Febiger 3437) which may be one of W. Smith’s collections. In none of
these specimens do I find any evidence of spines. It is true that on two
specimens from Pocono Lake I saw evidence of spines, but this condition
seems to be the exception rather than the rule.

Since Ehrenberg clearly indicates that he is doubtful about placing his
species in the genus Fragilaria and also since one cannot be sure from his
short description just what he had, I believe it best to make this form a
variety of W. Smith’s F. wndata. The specimens of W. Smith which he con-
sidered typical for F. wndata are very different from the figures of Ehren-
berg for F. constricta.

Monroe County: Pocono Lake Preserve, Pocono Lake, F. Myers (A-
2213). Pike County: Shohola Falls, 13-14 Burke (A-2206-09).

Fragilaria Vaucheriae (Kiitz.) Boye, 1938, Bot. Not. 1938(1-3): 164-71.
Exilaria Vaucheriae Kiitz., 1833, Linnaea 8: 560. pl. 15, fig. 38; Kiitz. Dec. no. 24.

Though found in several habitats this species was frequent in none.

Luzerne County: Harvey Lake, 2042 Hodge (A-2160). Monroe County:
Davy’s Run, 2018 Patrick (A-2118); Trout Lake, 2049 Hodge (A-2174);
Wier Lake, 2047 Hodge (A-2170). Pike County: Shohola Falls, 3 Burke
(A-2190), 11 Burke (A-2202). Wayne County: Angels, 2026 Burke and
Patrick (A-2134).

Fragilaria virescens Ralfs, 1843, Ann. Mag. Nat. Hist. 12: 110. pl. 2, fig. 6.

This species was very frequent in squeezings from H ypnum and Fonti-
nalis in Davy’s Run, from the swamp at Greeley, from surface of dead
leaves in shallow water at Angels, and from a locality of Shulze which was
merely designated as Pike County.

Monroe County: Davy’s Run, 2014 Patrick (A-2110), 2016-19 Patrick
(A-2113-19), 2056 Patrick (A-2180); Pocono Lake Preserve, Mud Run,
2031 Hodge (A-2138); Pocono Lake, 2037 Patrick (A-2150), F. Myers
(A-2213); Tobyhanna Creek, 2032-33 Patrick (A-2140-42). Pike County:
2188-90 Shulze (A-2233-36); 127-130 Shulze (A-2215-21); Bushkill
Creek, 1621 Shulze (A-2227); Greeley, 1 Burke (A-2184); Shohola Falls,
14 Burke (A-2208-10). Wayne County: Angels, 2024 Burke and Patrick
(A-2129), 2026 Burke and Patrick (A-2134).

Fragilaria virescens v. capitata Oestrup, 1910, Danske Diat., p. 193. pl. 5, fig. 125.

Patrick: Pocono DIATOMS 153

In Jaggie’s Bog this variety was very frequent.

Monroe County: Jaggie’s Bog, 262 Shulze (A-2223); Pocono Lake Pre-
serve, Pocono Lake, F. Myers (A-2213). Pike County: 128 Skulze (A-
2217). Wayne County: Angels, 2024 Burke and Patrick (A-2129).

ASTERIONELLA HaAssAtyi

Asterionella formosa Hassall, 1855, Microscopical Examination of Water, p. 10. (As
stated in previous papers I have been unable to verify this reference.)

This species reaches its best development in eutrophic water, though, as
seen from this study, it can grow in slightly distrophic water but does not
flourish so abundantly.

Luzerne County: Harvey Lake, 2042-43 Hodge (A-2161-62). Monroe
County: South Pocono Lake, 2010 Patrick (A-2102); Tobyhanna Creek,
2033 Patrick (A-2142); Pocono Lake, 2034-36 Patrick (A-2144-49), 2038
Patrick (A-2152); Trout Lake, 2049 Hodge (A-2175).

Asterionella Ralfsii W. Sm., 1856, Brit. Diat., 2: 81.

This species, unlike the one above, seems from this study to have its
best development in strongly distrophic, acid water. It reaches its best
development in Lake Tamaque with a pH of 3.8, and is frequent in Lost
Lake.

Monroe County: Lost Lake, 2020 Turner (A-2121); Lake Tamaque,
2039 Hodge (A-2154). Pike County: Shohola Falls, 14 Burke (A-2210).

SYNEDRA EHRENBERG

A. Striae typically 15 or more in 10 », often small diatoms with rather delicate structure

B. Ratio of breadth to length 1:11 or greater ....... .,..S. minuscula
B. Ratio of breadth to length less than above, central area distinct, often the
valve slightly swollen at the central area ............ S. rumpens and vars.

A. Striae less than 15 in 10 p
C. Central area usually broader than long, striae 12 or less in 10
S. ulna and vars.
C. Central area longer than broad, striae 12-14 in 10 », sometimes appearing
(oma horhio me Mynona noo cabnuqodotecguoe oo unuomdn S. acus and vars.

Synedra acus Kiitz., 1844, Bacill., p. 68. pl. 15, fig. 7.

This species was frequent among the moss growing at the top of Shohola
Falls in swift water. This is evidently not the most desirable habitat for
this species.

Pike County: Shohola Falls, 4 Burke (A-2192), 10 Burke (A-2201).

Synedra acus v. delicatissima (W. Sm.) Grun. apud Van Heurck, 1885, Syn. Diat.
Belgique, p. 151.
Synedra delicatissima W. Sm., 1853, Brit. Diat., 1: 72.

In W. Smith’s original description he states that this diatom may vary
from 130 to 220 p (he gives it in terms of inches).

154 FartowiA, VoL. 2, 1945

This variety was frequent in the surface water of Mountain Lake, pH 7.3.
Monroe County: Popomoning Lake, 2045 Hodge (A-2166) ; Mountain
Lake, 2051 Hodge (A-2178).

Synedra minuscula Grun. apud Van Heurck, 1881, Syn. Diat. Belgique, pl. 39, fig. 13.

Though found in several habitats this species was of frequent occurrence
in none.

Monroe County: Davy’s Run, 2014 Patrick (A-2110), 2017 Patrick
(A-2115), 2019 Patrick (A-2119), 2056 Patrick (A-2180); Pocono Lake
Preserve, Tobyhanna Creek, 2032 Patrick (A-2140); Raymondskill F alls,
2025 Patrick (A-2131). Pike County: Lake Wallenpaupek tributary, 5
Burke (A-2195); Shohola Falls, 4 Burke (A-2192), 10-11 Burke (A-2200—
02), 13-14 Burke (A-2206-08). Wayne County: Angels, 2022 Burke and
Patrick (A-2125), 2026 Burke and Patrick (A-2134).

Synedra rumpens Kiitz., 1844, Bacill., p. 69. pl. 16, fig. VI, 4, 5.

Though found in several habitats this species was always rare.

Monroe County: Davy’s Run, 2019 Patrick (A-2119); Mountain Lake,
2051 Hodge (A-2178) ; Raymondskill Falls, 2025 Patrick (A-2131); Trout
Lake, 2049 Hodge (A-2175). Pike County: Shohola Falls, 3-4 Burke
(A-2190-92), 10 Burke (A-2200). Wayne County: Angels, 2021 Burke
and Patrick (A-2124).

Synedra rumpens vy. familiaris (Kiitz.) Grun. apud Van Heurck, 1881, Syn. Diat.
Belgique, pl. 40, figs. 15, 16. (Grunow questions whether this should be a variety.)
Synedra familiaris Kiitz., 1844, Bacill., p. 68, pl. 15, fig. 12.

Monroe County: Wier Lake, 2046-47 H odge (A-2169-70). Wayne
County: Angels, 2026 Burke and Patrick (A-2134).

Synedra rumpens vy. scotica Grun. apud Van Heurck, 1881, Syn. Diat. Belgique,
pl. 40, fig. 11.

Pike County: Lake Wallenpaupek tributary, 5 Burke (A-2195). Wayne
County: Angels, 2021 Burke and Patrick (A-2123).

Synedra ulna (Nitzsch) Ehr., 1832, Abhandl. K. Akad. Wiss. Berlin 1831: 87.
Bacillaria ulna Nitzsch, 1817, N. Schrift. Naturf. Ges. Halle 3(1): 99.

In these collections were found specimens which agree with S. una in
regard to number of striae and the ratio between length and breadth but
which had a central area which was longer than broad, as found in S. acus.

This species was common in the squeezings from surface vegetation of
Wier Lake.

Monroe County: Mountain Lake, 2051 Hodge (A-217 8); Raymondskill
Falls, 2025 Patrick (A-2131); Wier Lake, 2046 Hodge (A-2169). Pike
County: 127 Skulze (A-2215); Bushkill Creek, 1623 Shulze (A-2231);
Lake Wallenpaupek tributary, 5 Burke (A-2195). Wayne County: Angels,
2022 Burke and Patrick (A-2125), 2026 Burke and Patrick (A-2134).

Patrick: Pocono D1aTomMs 155

Synedra ulna v. danica (Kiitz.) Van Heurck, 1881, Syn. Diat. Belgique, pl. 38, fig.
14a, text (1885) p. 151.
Synedra danica Kiitz., 1844, Bacill., p. 66. pl. 14, fig. 13.

This variety was very frequent in Mountain Lake, a shallow tributary of
Mill Creek, and at Shohola Falls. It is often a plankton form and it is un-
usual to find it so well developed in running water.

Monroe County: Mountain Lake, 2051 Hodge (A-2178); Popomoning
Lake, 2045 Hodge (A-2166). Pike County: 2188-89 Shulze (A-2233-35) ;
Bushkill Creek, 1621-23 Shulze (A-2227-31); Shohola Falls, 3-4 Burke
(A-2190-92), 10-14 Burke (A-2200-10). Wayne County: Angels, 2022
Burke and Patrick (A-2125).

Synedra ulna v. oxyrhynchus (Kiitz.) Van Heurck, 1885, Syn. Diat. Belgique, p. 151.
Synedra oxyrhynchus Kiitz., 1844, Bacill., p. 66. pl. 14, fig. 8, 2, IX—-XI.

This species was found only twice and then it was rare.
Monroe County: Raymondskill Falls, 2025 Patrick (A-2131). Wayne
County: Angels, 2026 Burke and Patrick (A-2134).

Synedra ulna v. Ramesii (Herib. and Perag.) Hustedt apud Pascher, 1930, Sussw.
Flora Mitteleurop. 10: 152. fig. 163.

Synedra Ramesii Herib. and Perag., 1903, Diat. Foss. d’Auvergne, p. 80. i. 11, fig.
28. (This reference was checked through the kindness of Miss Harris of the Farlow
Library.)

This variety was only found once and then it was rare.
Wayne County: Angels, 2021 Burke and Patrick (A-2124).

EUNOTIA EHRENBERG

A. Dorsal and ventral margins of valve parallel or nearly so
B. Valve with raphe extending backward from terminal nodules, a jelly pore
present at one end! of valve. ..-. 22.2.5... ---.---- E. flexuosa
B. Valve not as above
C. Valve strongly arcuate, spanning about 120° ..E. elegans
C. Valves not so strongly arcuate
D. Relatively large forms
E. Ends of valve distinctly wider than rest of valve, some-
what wedge-shaped

F. Valves slightly swollen in middle
E. formica

F. Valves not swollen in the middle
E. tautonensis

E. Ends of valve not distinctly wider than rest of valve,
but capitate, striae coarse, 8-10 in 10 pw
E. monodon v. major
D. Relatively narrow forms
G. Ends of valve distinctly capitate on dorsal margin, ven-
tral margin flat. See G’ and G”.
H. Valve arcuate

156 FartowiA, Vor. 2, 1945

I. Striae 14-17 in 10 yw, terminal nodules small
E. lunaris v. capitata
I. Striae 10-13 in 10 uw, terminal nodules large
E. gracilis
I. Striae 12-14 in 10 y, terminal nodules dis-
tinct but not large .E. arcus v. curta
H. Valve not arcuate, sometimes slightly curved,
striae 12-15 in 10xu..... E. fallax
G’. Ends of valve not capitate
J. Ends rounded, valve lunate in shape

E. lunaris
J. Ends rounded, cell wall thick, valve only slightly
arched ..................E. valida

G”. Ends of valve narrowed and attenuate
K. Valve very narrow, striae 15-20 in 10 u
E. Naegelii
K. Breadth and length of valve variable, sometimes
swollen at ends of valve, valves frequently occur-
ring in long chains ....... E. pectinalis and vars.
A. Dorsal margin smooth, convex, ventral margin straight or somewhat concave
L. Ends of valve distinctly set off from rest of valve
M. Typically large forms with coarse striae, not more than 12 in 10 u
N. Valves with ventral margin straight or only slightly concave,
ends of valve formed by a distinct narrowing of the valve, ends
TUUDCAEG~ 4.24 i Gieaueealeceie eau ate es E. praerupta and vars.
N. Ends of valve somewhat arcuate
O. Ends slightly swollen to form rounded apices
E. monodon
O. Ends distinctly capitate ..... ...E. indica
M. Typically small forms, striae more than 15 in 10 uw
P. Dorsal margin only slightly convex, ends strongly capitate,
usually as wide if not wider than main part of valve
E. exigua
P. Dorsal margin strongly convex
Q. Ends of valve more or less capitate
R. Striae about 21 in 10 u ...E. Meisteri
R. Striae 15-19 inlOw..... E. septentrionalis
Q. Ends of valve not capitate ..... E. rostellata
L. Ends of valve not distinctly set off from rest of valve
S. Striae 16 or more in 10 u
T. Dorsal margin of valve somewhat pointed in the middle, thus
giving the margin a slightly wedged appearance
E. trinacria
T. Dorsal margin of the valve smoothly convex, not as above
E. tenella
S. Striae usually 15 or less in 10 w
U. Ends of valve distinct from rest of valve, ends attenuate,
HOUMGER. 6 pcs bes He eG de lewinwavemue ree G E. sudetica
U. Ends of valve not distinct from rest of valve

Patrick: Pocono DIATOMS Ey

V. Ends of valve acute, giving a sharp pointed appearance
to the ends of the valves, sometimes slightly rounded
E. veneris

V. Ends of valve broadly rounded. .E. faba
A. Valve with dorsal margin with definite undulations

W. Valves small, two to five undulation on dorsal margin, striae about 18
sire) hase Gok ere were bs ote ee arene semen oes E. microcephala

W. Valves more robust, striae 15 or less in 10 wu

X. Ends of valve distinctly attenuate, undulations serrate, breadth of
valvei/—bl uw. sce ow and, Xe mle +s E. septena

X’. Ventral margin almost straight, undulations on dorsal margin crenate,
two in number

Y. Ends of valve distinctly truncate

Z. Undulations well developed ....E. suecica
Z. Undulations formed by a slight constriction in the dorsal
MN ALON Se ater, cose. Lace ee Cee E. praerupta v. bidens

Y. Ends of valve not truncate, but rounded E. diodon

X”. Ventral margin more or less concave, undulations crenate, four or
TTY O TG iy See ee rae Caap teense ehelene cpstelors E. robusta

Eunotia arcus v. curta (Grun.) Schonf., 1906, p. 116.
Himantidium arcus v. curtum Grun., 1862, Verh. Zool. Bot. Ges. Wien 12: 339. pl. 6,
fig. 16.

The identity of these specimens has been checked with Van Heurck’s
Type Slide No. 268. My specimens vary in length, some of them being 30 »
long. The striae are 12 in 10 p.

Monroe County: Jaggie’s Bog, 262 Shulze (A-2223). Pike County: 127
Shulze (A-2215).

Eunotia diodon Ehr., 1837, Ber. K. Akad. Wiss. Berlin 1837: 45.

This species, though found in several localities, was never represented by
more than a few specimens. Hustedt states that it is typically found on
bare rocks in Northern Europe, and also in damp moss. This corresponds
to the type of habitat where it was found in this study.

Monroe County: Davy’s Run, 2015 Patrick (A-2111), 2019 Patrick
(A-2119). Pike County: 127-128 Shkulze (A-2215-17); Bushkill Creek,
1621 Skulze (A-2227); Shohola Falls, 14 Burke (A-2208).

Eunotia elegans Oestrup, 1910, Danske Diat. p. 172. pl. 5, fig. 105.

I believe this is the first time this species has been recorded from North
America. The length of the valve measuring along its curvature was 48.7 pn,
the breadth was two p. The striae were 24-25 in 10 p. This species was
found in one collection.

Pike County: Greeley, 2 Burke (A-2188).

Eunotia exigua (Breb.) Rabh., 1864, Flora Eur. Algarum, p. 73.
Himantidium exiguum Breb. ex Kiitz., 1849, Spec. Algarum, p. 8.

This species was rare in all habitats. Krieger (1929) says it is found in
pH 3.5-5.5, especially with Sphagnum.

158 FarLow!A, VoL. 2, 1945

Monroe County: Davy’s Run, 2019 Patrick (A-2119). Pike County:
2188 Shulze (A-2234); Bushkill Creek, 1623 Shulze (A-2231).

Eunotia faba Ehr. emend. Van Heurck, 1881, Syn. Diat. Belgique, p. 143. pl. 34, fig. 34.
Eunotia faba Ehr., 1837, Ber. K. Akad. Wiss. Berlin, 1837: 45.

In all three habitats, this species occurred in running water associated
with Fontinalis or other plants living in water. It was rare in all habitats.
Monroe County: Davy’s Run, 2017 Patrick (A-2115); Pocono Lake
Preserve, Spring Run, 2037 C. Myers (A-2150). Pike County: Greeley, 1
Burke (A-2184).
Eunotia fallax A. Cl., 1895, Bihang K. Sv. Vet. Akad. Handl. 21 (3) no. 2: 33. pl. 1,
fig. 35.

Monroe County: Tobyhanna Creek, 2032 Patrick (A-2140).

Eunotia flexuosa Breb. ex Kiitz., 1849, Spec. Algarum, p. 6.

The forms of this species found during this study were shorter and nar-
rower than in typical specimens of the species. The ends were not inflated.
I have carefully studied these forms and checked them with the isotype
specimens in H. L. Smith, Spec. Typicae No. 555. These specimens differ
a great deal from those which I found in my study of Brazilian diatoms,
which have distinctly capitate ends.

The specimens were found in distinctly acid water. This species was of
common occurrence in Bushkill Creek and at the edge of the pond at
Shohola Falls.

Monroe County: Mountain Lake, 2051 Hodge (A-2178); Pocono Lake
Preserve, Pocono Lake, 2035-36 Patrick (A-2146-49), 2039 Patrick (A-
2154); Wier Lake, 2046 Hodge (A-2169); Lake Tamaque, 2039 Hodge
(A-2154). Pike County: 130 Skulze (A-2221), 12 Burke (A-2204), 14
Burke (A-2210).

Eunotia formica Ehr., 1843, Abhandl. K. Akad. Wiss. Berlin 1841: 414.

This species was rare and may be a contamination.
Monroe County: Mountain Lake, 2051 Hodge (A-2178).

Eunotia gracilis (Ehr.) Rabh., 1864, Flora Europaea Algarum, p. 72.
Himantidium gracile Ehr. 1843, Abhandl. K. Akad. Wiss. Berlin 1841: 417. pl. 2,
fig. 1, 9; pl. 3, fig. I, 41.

This species was found only once and then in small numbers.
Pike County: Greeley, 2 Burke (A-2188).

Eunotia indica Grun. apud Rabh., 1865, Beitrage Naheren Kennt. und Verbreit. des
Algen, 2: 5. pl. 1, fig. 7.

This species though found only twice was of frequent occurrence at the
spring on Bushkill Creek.

Monroe County: Pocono Lake Preserve, Pocono Lake, 2065 Patrick
(A-2182). Pike County: Bushkill Creek, 1621 Shudze (A-2227).

PaTRIcK: Pocono DIATOMS 159

Eunotia lunaris (Ehr.) Grun. apvd Van Heurck, 1881, Syn. Diat. Belgique, pl. 35,
figs. 3, 4, 6.
Synedra lunaris Ehr., 1832, Abhandl. K. Akad. Wiss. Berlin 1831: 37.

This species is widely distributed. Its most favorable habitats are found
in Pike County, particularly Bushkill Creek and Shohola Falls.

Luzerne County: Harvey Lake, 2043 Hodge (A-2162). Monroe County:
Davy’s Run, 2010 Patrick (A-2102), 2012-14 Patrick (A-2105-09), 2016
Patrick (A-2113), 2019 Patrick (A-2119); Jaggie’s Bog, 262 Shulze (A-
2223); Mountain Lake, 2051 Hodge (A-2178); Lake Tamaque, 2040
Hodge (A-2156); Wier Lake, 2046 Hodge (A-2168). Pike County: 127-
128 Shulze (A-2215-17); 2188-90 Shulze (A-2233-36); Bushkill Creek,
1621-23 Shulze (A-2227-31); Greeley, 1-2 Burke (A-2184-88) ; Shohola
Falls, 3-4 Burke (A-2190-92), 7 Burke (A-2197), 10-14 Burke (A-2200—
08).

Eunotia lunaris v. capitata (Grun.) Hustedt apud Pascher, 1930, Siissw. Flora Mittel-
europ., Heft 10: 185. fig. 250.
Synedra lunaris v. capitata Grun., 1862, Verh. Zool. Bot. Ges. Wien 12: 389.

Pike County: Greeley, 2 Burke (A-2188); Bushkill Creek, 1621 Shulze
(A-2227).

Eunotia Meisteri Hustedt apud Pascher, 1930, Siissw. Flora Mitteleurop. Heft 10: 179.
fig. 230.

Though occurring in several different habitats, this species was only once
very frequent and that was in association with Hypnum and Fontinalis, in
Davy’s Run.

Monroe County: Davy’s Run, 2014-19 Patrick (A-2109-19), 2056 Pat-
rick (A-2180); Pocono Lake Preserve, Mud Run, 2031 Hodge (A-2138) ;
Tobyhanna Creek, 2032-33 Patrick (A-2140—42) ; Wier Lake, 2047 Hodge
(A-2170). Pike County: Shohola Falls, 3 Burke (A-2190), 12-13 Burke
(A-2205-06).

Eunotia microcephala Krasske, 1932, Hedwigia 72: 102. pl. 2, fig. 7.

This is the same as Van Heurck’s concept of Eunotia tridentula and
E. tridentula v. perminuta. 1 have critically examined Van Heurck’s Type
Slides Nos. 309, 347, and his figures in Synopsis des Diatomées Belgique,
pl. 34, fig. 29, 30. Eunotia tridentula was first described by Ehrenberg
(Abhandl. K. Akad. Wiss. Berlin 1841: 414. pl. 2, fig. J, 14). From Ehren-
berg’s figure it is clear that it is a larger diatom than the one which Van
Heurck had. Therefore Van Heurck’s specimens should be considered a
misidentification and Krasske’s name should be adopted. This species was
common in Spring Run.

Monroe County: Davy’s Run, 2056 Patrick (A-2180); Pocono Lake
Preserve, Tobyhanna Creek, 2033 Patrick (A-2142); Spring Run, 2037
C. Myers (A-2150).

160 FartowiA, VoL. 2, 1945
Eunotia monodon Ehr., 1854, Mikrogeologie, pl. 2, fig. IJ, 26 a. b.

This species was only once of frequent occurrence and then in a spring
in Bushkill Creek.

Monroe County: Wier Lake, 2046 Hodge (A-2169); Jaggie’s Bog, 263
Shulze (A-2225). Pike County: 128 Shulze (A-2217); Bushkill Creek,
1621 Shulze (A-2227).

Eunotia monodon v. major (W. Sm.) Hustedt apud Pascher, 1930, Siissw. Flora
Mitteleurop., Heft 10: 186.

Himantidium majus W. Sm., 1856, Brit. Diat., 2: 14. pl. 33, fig. 286.

This species was present in several habitats, but was very frequent or
common in only two, Wier Lake and Mountain Lake. In both instances it
was found in association with surface littoral vegetation.

Monroe County: Davy’s Run, 2056 Patrick (A-2180); Pocono Lake
Preserve, Pocono Lake, 2035 Patrick (A-2146); Wier Lake, 2046 Hodge
(A-2168); Mountain Lake, 2051 Hodge (A-2178). Pike County: 2188-90
Shulze (A-2234-36); Shohola Falls, 4 Burke (A-2192), 11-12 Burke
(A-2202—-04), 14 Burke (A-2209). Wayne County: Angels, 2023 Burke
(A-2127).

Eunotia Naegelii Migula, 1907, Kryptogamen Flora, 2: 203.

This species is sometimes called Eunotia alpina (Naeg.) Hustedt. The
name alpina in the genus Eunotia is preoccupied by a species of Kutzing,
and therefore cannot be used for this species which when first named was
in the genus Synedra.

From this study, it seems that this species prefers definitely distrophic,
acid water. It was of very frequent occurrence in Lake Tamaque and
Lost Lake.

Monroe County: Lost Lake, 2020 Turner (A-2121); Lake Tamaque,
2039-40 Hodge (A-2154-56). Pike County: Shohola Falls, 12 Burke
(A-2205).

Eunotia pectinalis (Dillw.) Rabh., 1864, Flora Europea Algarum, p. 73.
Conferva pectinalis Dillw., 1809, Brit. Conf., p. 43. pl. 24.

From this study it seems that this species prefers moving water. It was
very frequent in Davy’s Run, at the spring in Bushkill Creek, and on drip-
ping rocks at Shohola Falls.

Monroe County: Davy’s Run, 2019 Patrick (A-2119), 2056 Patrick
(A-2180) ; Pocono Lake Preserve, Pocono Lake, F. Myers (A-2213); Toby-
hanna Creek, 2032 Patrick (A-2140); Wier Lake, 2046 Hodge (A-2169).
Pike County: 2188-90 Shulze (A-2233-36); Bushkill Creek, 1621 Shulze
(A-2227); Shohola Falls, 3-4 Burke (A-2190-92), 14 Burke (A-2208).

Eunotia pectinalis v. minor (Kiitz.) Rabh., 1864, Flora Europaea Algarum, p. 74.
Himantidium minus Kiitz., 1844, Bacill. p. 39. pl. 16, fig. 10, 1-4.

Though found in several localities this species was very frequent only in
Davy’s Run.

PATRICK: Pocono DIATOMS 161

Monroe County: Davy’s Run, 2018-19 Patrick (A-2118-19), 2056 Pat-
rick (A-2180); Lost Lake, 2025 Turner (A-2131); Pocono Lake Preserve,
Tobyhanna Creek, 2032-33 Patrick (A-2140-42); Wier Lake, 2046 Hodge
(A-2168); Mountain Lake, 2051 Hodge (A-2178). Pike County: 127
Shulze (A-2215); Bushkill Creek, 1623 Shulze (A-2231); Shohola Falls,
4 Burke (A-2192).

Eunotia pectinalis v. minor f. impressa (Ehr.) Hustedt apud Pascher, 1930, Stissw.
Flora Mitteleurop., Heft 10: 182.

Eunotia impressa Ehr., 1854, Mikrogeologie, pl. 3, fig. IV, 20; pl. 14, fig. 66.

This form was very frequent in one locality designated as Pike County.
Krieger (1929) states it is found in pH 4.5-S.5.

Monroe County: Pocono Lake Preserve, Spring Run, 2037 Myers (A-
2150); Pocono Lake, 2038 Patrick (A-2152), 2065 Patrick (A-2182). Pike
County: 2188-90 Shulze (A-2233-36); Bushkill Creek, 1623 Shulze (A-
2231); Greeley, 1-2 Burke (A-2184-88); Lake Wallenpaupek tributary,
5 Burke (A-2195); Shohola Falls, 13 Burke (A-2206).

Eunotia pectinalis v. recta Rabh. ex Mayer, 1917, Denks. K. Bayer. Bot. Ges. 13
(ns. 7): pl. 1, figs. 37, 38.

Mayer gives no description and I have not been able to find the descrip-
tion of Rabenhorst. Therefore I am supplying the following Latin de-
scription.

Valvae lenes arcuatae 19-20 » longae, 3 wu latae, apicibus linibus rotundis; striis
15-16 in 10 ph.

This variety was rare and found in only one locality.
Monroe County: Davy’s Run, 2016 Patrick (A-2113).

Eunotia pectinalis v. undulata (Ralfs) Rabh., 1864, Flora Europaea Algarum, p. 74.
Fragilaria pectinalis v. undulata Ralfs, 1843, Ann. Mag. Nat. Hist. 12: 108. pl. 2,
fig. 3d.

Ralfs refers to Pritchard, 1841, A History of the Infusoria, p. 220. fig.
171. I have looked up this reference and find that the species being con-
sidered is listed under the name Fragilaria grandis.

This species though found in several habitats was common only in Bush-
kill Creek and at Shohola Falls from moss in swift water at the top and in
scrapings from dripping rocks at Shohola Falls.

Monroe County: Davy’s Run, 2016 Patrick (A-2113), 2019 Patrick
(A-2119); Mountain Lake, 2051 Hodge (A-2178); Pocono Lake Preserve,
Mud Run, 2031 Hodge (A-2138) ; Pocono Lake, 2034-36 Patrick (A-2144—
48), 2038 Patrick (A-2152), 2065 Patrick (A-2182), F. Myers (A-2213);
South Pocono Lake, 2010 Patrick (A-2101); Tobyhanna Creek, 2032 Pat-
rick (A-2140). Pike County: 262 Shulze (A-2223); Bushkill Creek, 1621
Shulze (A-2227); Shohola Falls, 3 Burke (A-2190), 4 Burke (A-2193), 10
Burke (A-2200), 12 Burke (A-2204), 14 Burke (A-2210).

162 FARLOWIA, VoL. 2, 1945
Eunotia praerupta Ehr., 1841, Ber. K. Akad. Wiss. Berlin 1841: 143, 204.

This species was found only once and then it was rare.
Pike County: Greeley, 2 Burke (A-2188).

Eunotia praerupta v. bidens (Ehr.) Grun., 1880, K. Sv. Vet. Akad. Handl. 17 (2):
109.

Eunotia bidens Ehr., 1843, Abhandl. K. Akad. Wiss. Berlin 1841: 413.

Hustedt states that Ehrenberg’s identification is doubtful, so he uses
W. Smith’s Himantidium bidens as the basis for the name bidens. W. Smith,
however, gives Ehrenberg as the authority for the name Himantidium
bidens, and refers to Kiitzing’s “Species Algarum,” p. 9. Kiitzing refers to
Ehrenberg in Abhandl. K. Akad. Wiss. Berlin, 1841: 413 where he states
that Himantidium bidens equals Eunotia bidens, which is described in the
reference given above.

This species was found only in Pike County. In the Greeley habitats it
was associated with either Fontinalis or Sphagnum. In one collection, 2188
Shulze, specimens were found with spines on the dorsal margin of the valve.

Pike County: 129 Shulze (A-2219), 2188 Shulze (A-2234): Bushkill
Creek, 1621 Shulze (A-2227); Greeley, 1-2 Burke (A-2184-88).

Eunotia praerupta v. curta Grun. apud Van Heurck, 1881, Syn. Diat. Belgique,
pl. 34, fig. 24. (in the text, 1885, Van Heurck refers to this as forma curta).

This variety is rare in each of the localities.
Pike County: Greeley, 2 Burke (A-2188); Shohola Falls, 12 Burke
(A-2204).

Eunotia praerupta v. inflata Grun. apvd Van Heurck, 1881, Syn. Diat. Belgique,
pl. 34, fig. 17, text (1885) p. 143.

Pike County: Greeley, 1 Burke (A-2184).

Eunotia robusta Ralfs apud Pritchard, 1861, A History of the Infusoria, p. 763.

Unless, as I have previously suggested, De Toni’s “Sylloge Algarum”’ is
taken as a starting point for diatoms, this name, according to the Inter-
national Rules of Botanical Nomenclature section 10, article 56, will have
to be discarded.

In this study, the species was typically found in acid water. It reached
its best development in Lost Lake which is strongly distrophic, and in
South Pocono Lake in association with Sphagnum.

Monroe County: Lost Lake, 2020 Turner ( A-2121); Pocono Lake Pre-
serve, South Pocono Lake, 2010-11 Patrick (A-2101-03); Tobyhanna
Creek, 2032-33 Patrick (A-2140-42); Pocono Lake, 2034-35 Patrick
(A-2144-47), 2065 Patrick (A-2182), F. Myers (A-2213). Pike County:
128-29 Shulze (A-2217-19), 2188-90 Shulze (A-2234-36); Bushkill
Creek, 1621 Shulze (A-2227); Greeley, 1 Burke (A-2184); Shohola Falls,
12 Burke (A-2204).

PATRICK: Pocono DIATOMS 163

Eunotia rostellata Hustedt. This species was invalidly described by Hustedt, 1913, in
A. Schmidt’s Atlas der Diatomaceen-Kunde, pl. 289, figs. 3, 4. I have therefore in-
cluded a Latin description.

Valvae arcuatae 46 u longae, 4-5 u latae, dorsali et ventrali lateribus prope parallelis;
striis 14-17 in 10 pw.

I have checked my specimens with those of H. L. Smith, Spec. Typicae
no. 160 which are from the type locality, and they seem to be the same.
The striae usually vary from 14-15 in 10 » at the center of the valve to 17
in 10 » at the ends.

Pike County: 2189 Shulze (A-2235).

Eunotia septena Ehr., 1843, Abhandl. K. Akad. Wiss. Berlin 1841: 414. pl. 4, fig. I, 13.

It would perhaps be wise to unite several of these species of Ehrenberg
as Ralfs and Grunow have done. If this is done, however, according to the
International Rules of Botanical Nomenclature, one of Ehrenberg’s names
must be retained, as any additional name would be superfluous.

This species was rare in one locality.

Pike County: 127-30 Shulze (A-2215-21).

Eunotia septentrionalis Oestrup, 1897, Meddl. om Gronld. 15: 274. pl. 1, fig. 10.

This species occurred typically in acid water. It reached its best develop-
ment in association with moss in running water and was very frequent in
Sphagnum in the swamp at Greeley, and of frequent occurrence in Davy’s
Run and Mud Run.

Monroe County: Davy’s Run, 2014 Patrick (A-2109), 2019 Patrick
(A-2119); Pocono Lake Preserve, Mud Run, 2031 Hodge (A-2138). Pike
County: 2188 Shulze (A-2233), 128 Shulze (A-2217); Greeley, 1-2 Burke
(A-2184-88); Shohola Falls, 3 Burke (A-2190), 14 Burke (A-2208).

Eunotia sudetica O. Miill., 1898, Forschungsber. Biol. Stat. Plén 6: 59. pl. 3, figs. 25, 26.
Monroe County: Wier Lake, 2046 Hodge (A-2168).

Eunotia suecica A. Cleve, 1895, Bihang K. Sv. Vet. Akad. Handl. 21 Afd. 3 (2): 29,
pl. 1, figs. 31, 32.

This species occurred only once and then it was rare.
Pike County: Shohola Falls, 9 Burke (A-2198).

Eunotia tautonensis Hustedt.

This species, lacking any kind of description, is invalidly described by
Hustedt, 1913, in A. Schmidt’s Atlas Diatomaceen-Kunde, pl. 291, figs. 1-3.

Valvae arcuatae elongatae, 94-257 uw longae, 5-10 u latae, lateribus parallelis, apicibus
cuneatis; striis 12-17 in 10 uw.

As in Eunotia flexuosa the terminal nodules are distinct, and at one end
of the valve there is a jelly pore. The striae vary from 12-14 in 10 p» in the
center of the valve to 17 in 10 » at the end of the valve. Some of the speci-
mens from Lost Lake, where the species was very frequent, had small spines

164 FArRLowIA, VoL. 2, 1945

on the dorsal margin. However, this was not typically the case. No doubt
this species is near to the genus Desmogonium.

Monroe County: Lost Lake, 2020 Turner (A-2121); Pocono Lake Pre-
serve, Pocono Lake, F. Myers (A-2213).

Eunotia tenella (Grun.) Hustedt apud Pascher, 1930, Siissw. Flora Mitteleurop., Heft
10: 175. fig. 220.

Eunotia arcus v. tenella Grun. apud Van Heurck, 1881, Syn. Diat. Belgique, pl. 34,
figs. 5, 6.

Pirate I, ric. 1

This species was first invalidly described by Hustedt, 1913, A. Schmidt’s
Atlas Diatomaceen-Kunde, pl. 287, figs. 20-25.

A small form of this species which corresponds to Hustedt’s illustration
in A. Schmidt’s Atlas Diatomaceen-Kunde, pi. 287, fig. 25, was also found.
It is 9.3 » long, 3.5 » wide, and the striae are 17 in 10 ». As it is quite
different in shape from the usual F. tenella, I have illustrated it.

This species was very frequent in the swamp at Greeley.

Monroe County: Davy’s Run, 2016 Patrick (A-2113), 2018-19 Patrick
(A-2118-19); Pocono Lake Preserve, Mud Run, 2031 Hodge (A-2138).
Pike County: 2188 Shulze (A-2234); Bushkill Creek, 1623 Shulze (A-
2231); Greeley, 2 Burke (A-2188).

Eunotia trinacria Krasske, 1929, Bot. Arch. 27: 349. fig. 1.

This species was present in only two habitats.
Monroe County: Davy’s Run, 2017 Patrick (A-2115). Pike County:
Greeley, 2 Burke (A-2188).

Eunotia valida Hustedt apud Pascher, 1930, Siissw. Flora Mitteleuropas. Heft 10: 178.
fig. 229.

This species was rare in one habitat where it might have been a contami-
nation from some other habitat.
Pike County: 130 Skulze (A-2221).

Eunotia veneris (Kiitz.) De Toni, 1891, Sylloge Algarum, 2: 794.
Himantidium veneris Kiitz., 1844, Bacill. p. 40. pl. 30, fig. 7.

In 1854 Gregory (Quart. Jour. Micros. Sci. 2: 96) describes a diatom
which he states W. Smith has named Eunotia incisa. He states that he has
sent some of these specimens to Kiitzing who replied that they were dif-
ferent from Himantidium veneris. Gregory further states that his species is
very similar in shape to H. veneris, but that H. veneris does not have the
notches close to the terminal puncta nor does it have striae as Eunotia incisa
does. Obviously, if it belongs to the genus Himantidium, it has striae. As
for the notches to which Gregory refers, they are the terminal nodules. I
find that by careful focusing, this notch can be shown to appear and dis-
appear. Unfortunately I have not seen any authenticated specimens of
either Kutzing or Gregory.

PaTRICK: Pocono DIATOMS 165

In some cases specimens were found with obtuse ends as Gregory shows
as variety @. The striae in some of my specimens were coarser than is
usually found in E. veneris. They were in some cases as few as 12 in 10 p.
Perhaps these forms should be separated as varieties.

This species was very frequent at Shohola Falls in a collection made from
a dripping wall and in a collection from algae and moss in the brook. It
was common in squeezings from plants (Elodea, Myriophyllum, etc.) in
shallow water in Pocono Lake and Lake Popomoning, and in Spring Run.

It is interesting to note that this species reached such good development
in two habitats which were quite diverse in some respects. Pocono Lake has
an average pH of 6.4. Analysis of the water shows calcium 2.37 p.p.m., and
bicarbonates 15.6 p.p.m. Popomoning Lake has a pH of 8-8.1, calcium
8.82 p.p.m., and bicarbonates 33.6-34.2 p.p.m. In Pocono Lake the speci-
mens had rounded ends, and striae about 14 in 10 p. In Lake Popomoning
the specimens had acute apices and striae 17-20 in 10 w. Krieger (1929)
states that this species is found in pH 4.5—5.5, which is lower than either of
these two habitats.

- Luzerne County: Harvey Lake, 2042 Hodge (A-2160). Monroe County:
Davy’s Run, 2015-16 Patrick (A-2111-13), 2018 Patrick (A-2118); Po-
cono Lake Preserve, Mud Run, 2031 Hodge (A-2138); Pocono Lake, 2034—
37 Patrick (A-2144—-50), 2065 Patrick (A-2182); Tobyhanna Creek, 2032—
33 Patrick (A-2140—-42); Popomoning Lake, 2045 Hodge (A-2166). Pike
County: 2188 Shulze (A-2233); Bushkill Creek, 1623 Shulze (A-2231),
Greeley, 2 Burke (A-2188); Shohola Falls, 3 Burke (A-2190), 10-14 Burke
(A-2200-09). Wayne County: Angels, 2021 Burke and Patrick (A-2123).

COCCONEIS (Eur.) CLEVE

Cocconeis placentula Ehr., 1838, Die Infusionsthierchen als volkommene Organismen,
p. 194.

This species was of common or very frequent occurrence at Raymonds-
kill Falls, Harvey Lake, Popomoning Lake, and Mountain Lake.

Luzerne County: Harvey Lake, 2042 Hodge (A-2160). Monroe County:
Raymondskill Falls, 2025 Patrick (A-2131); Popomoning Lake, 2045
Hodge (A-2166); Mountain Lake, 2051 Hodge (A-2178). Northampton
County: Green Pond, 2050 Hodge (A-2177). Pike County: Shohola Falls,
10-11 Burke (A-2201-02); Lake Wallenpaupek tributary, 5 Burke (A-
2195); Bushkill Creek, 1623 Shulze (A-2231). Wayne County: Angels,
2021-22 Burke and Patrick (A-2124-25).

Cocconeis placentula v. lineata (Ehr.) Cl., 1895, K. Sv. Vet. Akad. Handl. 27 (3):
169.

Cocconeis lineata Ehr. is first mentioned by Ehrenberg, Abhandl. K.
Akad. Wiss., 1841, p. 369, but is neither described nor illustrated. It is
first illustrated, I believe, in his Passat-Staub und Blut-Regen, Abhandl.
K. Akad. Wiss. Berlin 1847, and published separately in 1849, where it
appears on pi. 5, J, fig. 10.

166 FaRLowiA, VoL. 2, 1945

This variety was very frequent at Shohola Falls and at Angels. In both
these cases it was found attached to stones in shallow swift water.

Monroe County: Popomoning Lake, 2045 Hodge (A-2166); Wier Lake,
2047 Hodge (A-2170). Pike County: Shohola Falls, 4 Burke (A-2192).
Wayne County: Angels, 2026 Burke and Patrick (A-2134).

ACHNANTHES Bory

Cleve has divided this genus into several genera. Hustedt prefers to use
Cleve’s names to indicate subgenera or groups rather than to split the
genus into several genera. Since Hustedt’s arrangement is the one most
generally used today, and until I have studied the genus more critically, I
shall follow it. On this basis three groups are recognized in this study.

A. Transverse rows composed of very coarse punctae, the pseudoraphe usually excentric.
y p ] d

mee AY and AM. i. be acceaheas ap aeulee aaauniat ave rmeakddawews Achnanthidium
A’. Transverse rows composed of very fine punctae, the pseudoraphe central.
Microneis
A”. Transverse rows composed of a double row of punctae......... Achnanthes

ACHNANTHES subgenus

Achnanthes coarctata (W. Sm.) Grun. apud Cl. and Grun., 1880, K. Sv. Vet. Akad.
Handl. 17 (2): 20.
Achnanthidium coarctatum W. Sm., 1855, Ann. Mag. Nat. Hist. 2nd Ser. 15: 8,
fig. 10.

W. Smith cites Brebisson as the authority and gives as reference Kiitz.
Spec. Algarum, p. 54. This is not a correct reference and I cannot find that
Brebisson ever published such a name.

The only habitat in which this species was found, was on the surfaces of
dripping rocks which were outcrops of limestone.

Pike County: Shohola Falls, 15 Burke (A-2211), 9 Burke (A-2198).

MicroNEIS subgenus

A. Rapheless valve with a horseshoe-shaped area well-developed on one side of the
middle of the valve. See A’ and A”.
B. Valve with raphe with the striae irregularly longer and shorter in the central
area, More numerous on valve with raphe than on rapheless valve
C. Valve elliptic with attenuate rostrate ends ....A. Peragalli
C. Valve elliptic but ends usually rounded, not as above
A. Oestrupii var.
B. Not as above, striae about the same number on both valves
A. lanceolata and vars.
A’. Rapheless valve with striae broken so as to form a longitudinal line on each side of
the pseudoraphe ....... 0... ccc cece cece cece ecceenanene A. Lewisiana
A”. Rapheless valve without horseshoe-shaped area or longitudinal lines
D. Valve with raphe with a well-developed stauros reaching to the edges of the
valve

Patrick: PocoNo DIATOMS 167

E. Stauros on one side of raphe having an area which appears to be
raised, thus giving the stauros on that side a bifurcate appearance
A. hungarica

E. Stauros having the same appearance on both sides of the central nodule
A. exigua —

D. Valve with raphe with or without a stauros, if present not reaching to edges
of valve, striae fine, over 20 in 10 u
Pee Valvie line ate sais tke. cs) duns tlatsas S Oe ea stekoteparus A. linearis

F. Valve with capitate-rostrate ends, striae usually slightly more distantly
spaced at center of valve around central nodule A. microcephala

F. Hard to distinguish from the above, but ends usually rounded or some-
what rostrate, not capitate, striae not more distant at central nodule
A. minutissima

Achnanthes exigua Grun. apud Cl. and Grun., 1880, K. Sv. Vet. Akad. Handl. 17 (2):
ale
This species was found only once and then it was rare.
Monroe County: Popomoning Lake, 2045 Hodge (A-2166).

Achnanthes hungarica Grun. apud Cl. and Grun., 1880, K. Sv. Vet. Akad. Handl.
17 (2): 20.
Achnanthidium hungaricum Grun., 1863, Verh. Zool. Bot. Ges. Wien, 13: 146.
This species was only found once and then it was rare. It is characterized
by the fact that the stauros appears to be bifurcate on one side of the
central nodule, but close examination shows that this is not true.
Monroe County: Mountain Lake, 2051 Hodge (A-2178).

Achnanthes lanceolata (Bréb.) Grun. apud Cl. and Grun., 1880, K. Sv. Vet. Akad.
Handl. 17 (2): 23.
Achnanthidium lanceolatum Bréb. ex Kiitzing, 1849, Spec. Algarum, p. 54.

This species was very frequent in the shallow water of a small tributary
of Mill Creek, from the surface of leaves of a very small puddle near Mill
Creek, enmeshed in moss in the swift water of Mill Creek, and in the sur-
face scum on rocks in Raymondskill Falls.

Monroe County: Raymondskill Falls, 2025 Patrick (A-2131); Mountain
Lake, 2051 Hodge (A-2178). Northampton County: Green Pond, 2050
Hodge (A-2177). Pike County: Shohola Falls, 3 Burke (A-2190), 15
Burke (A-2211), 5 Burke (A-2195). Wayne County: Angels, 2021-24
Burke and Patrick (A-2124—29).

Achnanthes lanceolata var. apiculata var. nov.
PrateE I, Fics. 4, 5

Valvae elliptico-lanceolatae, 28 « longae, 10-11 4 latae, apicibus apiculatis. Valva
superior ad instar soleae equinae effigiem in medio ostendens, pseudoraphe anguste
lanceolata; striis radiantibus, 10-12 in 10 w. Valva inferior aream axialem angustam
et aream centralem rotundatam exhibens; striis validis radiantibus, 10 in 10 yu. Type:
Shohola Falls, 3 Burke (A-2190).

A noteworthy feature of this diatom is that the valve with a raphe has
much more strongly radiate striae than the valve without a raphe. It dif-

168 FARLow!IA, VOL. 2, 1945

fers from A. lanceolata v. robusta in that the ends are more apiculate than
in robusta and the axial area is narrower and not granular.

This diatom was only found in one geographical habitat, Shohola Falls,
but it was well distributed in the various ecological habitats here.

Pike County: Shohola Falls; 3 Burke (A-2190), 10-14 Burke (A-2201-
10).

Achnanthes lanceolata v. rostrata Hust. apud Pascher, 1930, Siissw. Flora Mittel-
europ., Heft 10: 208. fig. 306b.

In this study this variety was only found in running water. It was very
frequent in the surface scum on rocks in little pools in Raymondskill Falls.

Monroe County: Raymondskill Falls, 2025 Patrick (A-2131). Pike
County: Shohola Falls, 10-11 Burke (A-2200-02); Lake Wallenpaupek
tributary, 5 Burke (A-2195). Wayne County: Angels, 2026 Burke and
Patrick (A-2134).

Achnanthes Lewisiana sp. nov.
PLATE I, Fics. 2, 3

Valvae lineari-ellipticae, 15 » longae, 4.6 uw latae. Valva superior pseudoraphem dis-
tinctam exhibens; striis 15-17 in 10 ys; utrimque per lineam longitudinalem inter-
ruptis. Valva inferior aream axialem ad valvae ipsae '4-™% latitudinem prope nodulum
centralem exhibens; area centrale nulla; striis parum radiantibus, 15-17 in 10 uw. Type:
Shohola Falls, 14 Burke (A-2210).

This diatom is named in honor of Dean Ivey F. Lewis with whom I did
my graduate work at the University of Virginia.

This species is particularly characterized by the interrupted striae on the
rapheless valve. The most closely related fresh water species is A. Such-
landti Hustedt. The resemblance of my specimen to this species is mainly
in the interrupted striae of the rapheless valve, for it differs from it in shape
of axial area, number of striae, and size.

Both valves were seen and drawn from the same frustule.

Pike County: Greeley, 2 Burke (A-2188); Shohola Falls, 14 Burke
(A-2208, 2210).

Achnanthes linearis (W. Sm.) Grun. apud Cl. and Grun., 1880, K. Sv. Vet. Akad.
Handl. 17 (2): 23.
Achnanthidium lineare W. Sm., 1855, Ann. Mag. Nat. Hist. 2d ser. 15: 8. pl. 1, fig. 9.

Though found in a great many habitats, this species was common or very
frequent only in Davy’s Run. It was found in association with moss, par-
ticularly Fontinalis, in running water.

Monroe County: Davy’s Run, 2017 Patrick (A-2115), 2019 Patrick
(A-2119), 2056 Patrick (A-2180); Pocono Lake Preserve, 2031 Hodge
(A-2138), 2032 Patrick (A-2140), 2065 Pétrick (A-2182). Luzerne
County: Harvey Lake, 2042 Hodge (A-2160). Pike County: Shohola Falls,
10-14 Burke (A-2200-09).

PATRICK: Pocono DIATOMS 169

Achnanthes microcephala (Kiitz.) Grun. apud Cl. and Grun., 1880, K. Sv. Vet. Akad.
Handl. 17 (2): 22.

Achnanthidium microcephalum Kiitz., 1844, Bacill., p. 75. pl. 3, figs. 13, 19.

Luzerne County: Harvey Lake, 2042 Hodge (A-2160). Monroe County:
Trout Lake, 2049 Hodge (A-2174).

Achnanthes minutissima Kiitz., 1844, Bacill., p. 75. pl. 13, fig. 2c.

This species was found only once and then it was rare.
Pike County: Lake Wallenpaupek tributary, 5 Burke (A-2195).

Achnanthes minutissima v. cryptocephala Grun. apud Van Heurck, 1880, Syn.
Diat. Belgique, pl. 27, figs. 41-44.

Monroe County: Pocono Lake Preserve, Tobyhanna Creek, 2032-33
Patrick (A-2140-42).

Achnanthes Oestrupii v. parvula var. nov.

Prate I, Fics. 6, 7

Valvae ellipticae apicibus parum rostratis, 12 « longae, 6 m latae. Valva superiot
spatium hyalinum et in latere uno effigiem ad instar soleae equinae ostendens; striis
parum radiantibus 15-16 in 10 w. Valva inferior aream axialem angustam et aream
centralem transversam exhibens; striis 28 in 10 «, radiantibus, longis inaequaliter et ad
nodulum centralem crassis. Type: Shohola Falls, 14 Burke (A-2209).

This variety differs from the species in that it is consistantly smaller in
size. It also differs from the original description of the species in the num-
ber of striae of the rapheless valve. This species was first noted by Oestrup
and considered a variety of A. lanceolata. A. Cleve (Acta Forst. Fenn. 22:
53) first named it A. lanceolata v. Oestrupii. The illustrations of both
authors indicate the striae number of the rapheless valve as 10—12 in 10 p.
Hustedt raised the variety to specific rank, and states the striae number of
the rapheless valve is “10-14 (measuring on the edge of the short striae
forms) or 14-18 (measuring on the pseudoraphe of the longer striated
forms).” My specimens have striae 15 in 10 » whether measured at margin
or inner edge of striae.

Both valves were seen on the same frustule.

Pike County: Shohola Falls, 10, 14 Burke (A-2200, 2209).

Achnanthes Peragallii Brun and Herib. apud Herib., 1893, Diat. Auvergne, p. 50.
pl. 1, fig. 4.

This species was rare in the two localities where it was found. In both
cases it was in running water.

Pike County: Shohola Falls, 10 Burke (A-2200); Lake Wallenpaupek
tributary, 5 Burke (A-2195).

ACHNANTHES subgenus

Achnanthes Stewartii sp. nov.

170 FarRtowlA, VoL. 2, 1945
Puate IT, rics. 1-3

Valvae ellipticae, 16 « longae, 7.5 « latae, area axiali angusta, costis tenuibus 14 in
10 u, serie duplici punctorum parvorum interpositu. Type: Pike County: Lake Wallen-
paupek tributary, 5 Burke (A-2194).

This species is named in honor of Dr. Thomas S. Stewart, Curator of the
Microscopic Department of the Academy of Natural Sciences of Philadel-
phia, who has kindly made most of the photographs of diatoms which I
have published.

It is with some hesitation that I put this species in this subgenus. The
costae are not very pronounced, but the presence of a double row of
punctae between each two costae places it here.

This species was only found in one locality, but as it is a very distinct
species and of frequent occurrence I have described it.

Pike County: Lake Wallenpaupek tributary, 5 Burke (A-2194-95).

AMPHIPLEURA Kirzinc

Amphipleura pellucida Kiitzing, 1844, Bacill., p. 103. pl. 3, fig. 52.
Frustulia pellucida Kiitz., 1833, Linnaea, 1833: 543. pl. 13, fig. 11.

This species seems to prefer circum-neutral waters. It was found only
once in this study.
Luzerne County: Harvey Lake, 2043 Hodge (A-2162).

VANHEURCKIA Bresisson

The name Frustulia, sometimes applied to this genus, is invalid according
to the International Rules of Botanical Nomenclature.

Vanheurckia rhomboides (Ehr.) Bréb., 1869, Ann. Soc. Phy. Mic. Belge 1: 204.

Navicula rhomboides Ehr., 1843, Abhandl. K. Akad. Wiss. Berlin 1841: 419. pl. 3,
fig. I, 15.

This species was very frequent in Lost Lake which is a strongly distrophic
lake which contains a great deal of Sphagnum. It is found only in acid
water, and usually in association with moss.

Lackawanna County: Indian Lake, 2044 Hodge (A-2164). Monroe
County: Lost Lake, 2020 Turner (A-2121) ; Pocono Lake Preserve, Pocono
Lake, F. Myers (A-2213); South Pocono Lake, 2010-11 Patrick (A-2101-
03). Pike County: Shohola Falls, 11 Burke (A-2202).

Vanheurckia rhomboides v. amphipleuroides (Grunow)
Navicula rhomboides v. amphipleuroides Grun. apud Cl. and Grun., 1880, K. Sv.
Vet. Akad. Handl. 17 (2): 47. pl. 3, fig. 59.
Grunow places this species in the group Vanheurckia under the genus
Navicula.
This variety was found in shallow running water and was very frequent
in a shallow tributary of Mill Creek. It was not found in the standing,
strongly distrophic waters of some of the habitats studied.

Patrick: Pocono DIATOMS 171

Monroe County: Raymondskill Falls, 2025 Patrick (A-2131). Pike
County: Bushkill Creek, 1623 Shkulze (A-2231); Lake Wallenpaupek trib-
utary, 5 Burke (A-2195). Wayne County: Angels, 2021-22 Burke and
Patrick (A-2124-25), 2026 Burke and Patrick (A-2134).

Vanheurckia rhomboides v. crassinervia (Bréb.) Van Heurck, 1881, Syn. Diat.
Belgique, pl. 17, figs. 4-5, text (1885) p. 112.

Navicula crassinervia Bréb. apud W. Sm., 1853, Brit. Diat., 1: 47.

Brébisson (Ann. Soc. Phy. Mic. Belge 1:204) later transferred this spe-
cies to the genus Vanheurckia.

This variety is the same as the invalidly published Frustulia rhomboides
v. saxonica (Rabh.) De Toni.

This variety was well distributed in the more acid waters of this region
and was:of common occurrence in Davy’s Run, Lost Lake, Pocono Lake,
and Lake Tamaque.

Lackawanna County: Indian Lake, 2044 Hodge (A-2165). Monroe
County: Davy’s Run, 2013-15 Patrick (A-2107-11), 2018-19 Patrick
(A-2117-19), 2056 Patrick (A-2180); Lost Lake, 2020 Turner (A-2121);
Pocono Lake Preserve, Pocono Lake, 2034-38 Patrick (A-2144—-52), 2065
Patrick (A-2182); Mud Run, 2031 Hodge (A-2138); South Pocono Lake,
2011-12 Patrick (A-2104—-05); Tobyhanna Creek, 2032-33 Patrick (A-
2140-42); Lake Tamaque, 2039-40 Hodge (A-2154-56). Pike County:
2188-90 Shulze (A-2234-36); Greeley, 2 Burke (A-2188); Shohola Falls,
4 Burke (A-2193), 10 Burke (A-2200), 12 Burke (A-2204), 14 Burke
(A-2210).

Vanheurckia rhomboides v. crassinervia f. capitata comb. nov.

This form is the same as the invalidly published Frustulia rhomboides v.
saxonica {. capitata (Mayer) Hustedt.

Valvae elliptico-lanceolatae, ca. 45 « longae; apicibus rostrato-capitatis; striis sub-
tilissimis, 34-36. in 10 w.

This form is characterized by its capitate-rostrate ends. It is usually
about 45 » long.

This form was common in Pocono Lake from squeezings of Eleocharis,
Dicranum, and Hypnum growing in one foot of water and was very frequent
in scrapings of piles supporting the Emlen Dock in Pocono Lake. These
two habitats were within fifty yards of each other.

Monroe County: Davy’s Run, 2016 Patrick (A-2113); Mountain Lake,
2051 Hodge (A-2178); Pocono Lake Preserve, Pocono Lake, 2035-36
Patrick (A-2147-48), 2065 Patrick (A-2182); Tobyhanna Creek, 2032-
33 Patrick (A-2140-42); South Pocono Lake, 2010 Patrick (A-2101).
Pike County: Bushkill Creek, 1623 Skulze (A-2231); Shohola Falls, 3—4
Burke (A-2190-92), 11 Burke (A-2202), 14 Burke (A-2208).

Vanheurckia vulgaris (Thw.) Van Heurck, 1880, Syn. Diat. Belgique, pl. 17, fig. 6,
text (1885) p. 112.

Colletonema vulgaris Thw., 1848, Ann. Mag. Nat. Hist. 2d ser. 1, pl. 12.

172 FarLowla, VoL. 2, 1945

This species was found in several localities but was rare in each case. It
was usually found in running water.

Monroe County: Davy’s Run, 2015 Patrick (A-2111); Wier Lake, 2047
Hodge (A-2171); Raymondskill Falls, 2025 Patrick (A-2131). Pike
County: 2189 Shulze (A-2235); Shohola Falls, 4 Burke (A-2192), 14
Burke (A-2208). Wayne County: Angels, 2021 Burke and Patrick (A-
2124), 2026 Burke and Patrick (A-2134).

CALONEIS CLeve

Caloneis Lewisii sp. nov.

Pirate II, Fic. 4

Valvae bis constrictae longae 27 w ad medium 8.7 uw apicibus cuneatae; striis trans-
versis 19-20 in 10 w; area axiali angusto, nodulo centrali majusculo subquadrangularis
signis lunatis duobus crasso. Type: Illinois, Du Page County, 2423 F. Drouet, P. C.
Standley and J. A. Steyermark (A-2018).

It is regrettable that the often used name, Caloneis trinodis, cannot be
retained for this species. Caloneis trinodis is based upon the specimens of
Lewis. Lewis misidentified his specimens as Navicula trinodis W. Sm. I
have conferred with Dr. E. D. Merrill and Dr. Francis W. Pennell about
using the name Caloneis trinodis, but they assure me that since this species
is based upon the misidentification of Lewis, this cannot be done. In 1878
Grunow, apparently unaware of Lewis’ specimens, called this species Navi-
cula Trochus var.? biconstricta. Unfortunately the name biconstricta is
preoccupied in the genus Caloneis. Thus it seems that the best thing to do
is to name the species after Lewis who found the first specimens. I do not
believe it correct to consider it a variety of Caloneis Schumanniana as Cleve
(K. Sv. Vet. Akad. Hand]. 26 (2): 53) and Reichelt (Archiv Hydrobiol.
Planktonk. 1: 231-32) have done.

In my recent paper (Proc. Acad. Nat. Sci. 95: 71) I have given Boyer
the credit for the name Caloneis trinodis. This was a mistake as Meister
first made the combination in 1912. However, as shown above, this name
is invalid.

Northampton County: Green Pond, 2050 Hodge (A-2176).

Caloneis silicula v. alpina Cl., 1893, K. Sv. Vet. Akad. Handl. 26 (2): 51.

It is very interesting to find this variety in the Pocono Plateau. The
identification of the variety has been checked with Cleve and Moller type
no. 273. Hustedt stated that this variety is only found in the high moun-
tains of Europe.

In this study, the diatom was found only once, and then it was rare.
From this I would infer that Davy’s Run is not the ideal habitat, and the
specimens which I found might be contaminations brought in by birds, etc.
from some other habitat.

Monroe County: Davy’s Run, 2014 Patrick (A-2110).

PATRICK: Pocono DIATOMS 173

NEIDIUM PFITZER

A. Striae more than 20 in 10 yu
B. Ends of valve rounded, not at all constricted
-C. Sides of valve parallel ............... N. bisulcata
C. Sides of valve undulate .............. N. bisulcata v. undulata
B. Ends of valve more or less constricted, from slightly rostrate to protracted
rostrate ends
D. Ends of valve only slightly constricted N. affine

D. Ends of valve distinctly protracted and rostrate
N. affine v. amphirhynchus

A. Striae less than 20 in 10 »
E. Valves linear

PPP serOuUnGe dine 38.0 5 ti access wich we ae ee N. iridis f. vernalis

Bo buds; wedee shaped .22. 6. i500 0s caseen N. iridis v. amphigomphus
E. Valves linear-lanceolate

GUebnds rounded 3.6... see ese Nath sea ies N. iridis ©

G@. Ends. subrostrate ccs nk be eee to N. iridis v. ampliata

Neidium affine (Ehr.) Cl., 1894, K. Sv. Vet. Akad. Handl. 26 (2): 69.
Navicula affinis Ehr., 1843, Abhandl. K. Akad. Wiss. Berlin 1841: 417. pl. 2, fig. I, 7.

This species was rare in the two localities. In both cases it occurred in
running water.

Monroe County: Davy’s Run, 2013 Patrick (A-2107), 2019 Patrick
(A-2119). Pike County: 1621 Shulze (A-2227).

Neidium affine v. amphirhynchus (Ehr.) Cl., 1894, K. Sv. Vet. Akad. Handl. 26 (2):
68.
Navicula amphirhynchus Ehr., 1843, Abhandl. K. Akad. Wiss. Berlin 1841: 417.
pl. 3, fig. I, 10.
This variety was rare in two habitats.
Monroe County: Davy’s Run, 2014 Patrick (A-2110). Wayne County:
Angels, 2021 Burke and Patrick (A-2123).

Neidium bisulecatum (Lagst.) Cl., 1894, K. Sv. Vet. Akad. Handl. 26 (2): 68.
Navicula bisulcata Lagst., 1873, Bihang K. Sv. Vet. Akad. Handl. 1 (14): 31.
pl. 1, fig. 8.
This species was rare in the two localities where it was collected.
Pike County: 127 Shulze (A-2215), 2188 Shulze (A-2234).

Neidium bisulcatum v. undulatum O. Miill., 1898, Forschungsber. Biol. Stat. Plon 6:
62-63.

This variety was rare in occurrence.
Pike County: Greeley, 2 Burke (A-2188).

Neidium iridis (Ehr.) Cl., 1894, K. Sv. Vet. Akad. Handl. 26 (2): 69.
Navicula iridis Ehr., 1843, Abhandl. K. Akad. Wiss. Berlin 1841: 418. pl. 4, figs. 1, 2.

Though found in several localities, it was of frequent occurrence in none.
Luzerne County: Harvey Lake, 2042 Hodge (A-2160). Monroe County:

174 FarLowIA, VoL. 2, 1945

Davy’s Run, 2014 Patrick (A-2109), 2019 Patrick (A-2119); Raymonds-
kill Falls, 2025 Patrick (A-2131); Wier Lake, 2046 Hodge (A-2169).
Northampton County: Green Pond, 2050 Hodge (A-2177). Pike County:
2188-90 Shulze (A-2234-36), 128-29 Shulze (A-2217-19); Greeley, 1
Burke (A-2184); Shohola Falls, 4 Burke (A-2192), 12 Burke (A-2204).

Neidium iridis v. amphigomphus (Ehr.) Mayer, 1917, Denks. K. Bayer. Bot. Ges.
13: n.s. 7: 30.
Navicula amphigomphus Ehr., 1843, Abhandl. K. Akad. Wiss. Berlin 1841: 417.
pl. 3, fig. I, 8.
This variety was frequent in a locality simply designated by Shulze as
Pike County.
Monroe County: Jaggie’s Bog, 262 Shulze (A-2223). Pike County: 130
Shulze (A-2221).

Neidium iridis v. ampliatum (Ehr.) Cl., 1894, K. Sv. Vet. Akad. Handl. 26 (2): 69.
Navicula ampliata Ehr., 1842, Ber. K. Akad. Wiss. Berlin, 1842: 337.

These specimens differ from the typical forms of this variety in that the
sides of the valve are parallel and not convex.
Pike County: 2190 Shulze (A-2236).

Neidium iridis v. vernale Reichelt, 1897, Sitzungsber. Naturforsch. Ges. Leipsig 24:
8-9.

This variety was only found once and then it was rare.
Pike County: 128 Shulze (A-2217).

DIPLONEIS EHRENBERG

This genus was rarely encountered in this study. The two species listed
below were only found twice and then they were rare.

Diploneis ovalis (Hilse) Cl., 1891, Acta Soc. pro Fauna et Flora Fennica 8 (2): 44.
pl. 2, fig. 13.
Pinnularia ovalis Hilse in Rabenhorst, 1861, Algen Europa, no. 1025.

Luzerne County: Harvey Lake, 2042 Hodge (A-2160). Monroe County:
Raymondskill Falls, 2025 Patrick (A-2131).

Diploneis Smithii (Bréb. ex W. Sm.) Cl., 1894, K. Sv. Vet. Akad. Handl. 26 (2): 96.
Navicula Smithit Bréb. ex W. 5m., 1856, Brit. Diat. 2: 92.

Monroe County: Raymondskill Falls, 2025 Patrick (A-2131).

STAURONEIS EHRENBERG

A. Ends of valve with septum, valves slightly biconstricted ...... S. Smithii

A. Ends of valve without septum
B. Raphe broad, striae usually less than 20in 10 u........ S. phoenicenteron
B. Raphe linear, striae 20 or more in 10 u.............0.- S. anceps and vars.

B. Raphe linear, striae about 30 in 10 mw, small forms usually less than 25 » long
S. Kriegeri

Patrick: Pocono DIATOMS 175

Stauroneis anceps Ehr., 1843, Abhandl. K. Akad. Wiss. Berlin 1841: 306, 422. pl. 2,
fig. I, 18.

This species was found in several localities but was frequent in none.

Lackawanna County: Indian Lake, 2044 Hodge (A-2165). Monroe
County: Davy’s Run, 2014-15 Patrick (A-2110-11), 2018-19 Patrick
(A-2118-19); Mountain Lake, 2048 Hodge (A-2172); Pocono Lake Pre-
serve, South Pocono Lake, 2010 Patrick (A-2102); Trout Lake, 2049
Hodge (A-2174); Wier Lake, 2046-47 Hodge (A-2169-70). Pike County:
2188 Shulze (A-2234), 128-30 Shulze (A-2217-21); Shohola Falls, 12
Burke (A-2204).

Stauroneis anceps v. amphicephala (Kiitz.) Cl., 1894, K. Sv. Vet. Akad. Handl.
26 (2): 148.
Stauroneis amphicephala Kiitz., 1844, Bacill. p. 105. pl. 30, fig. 25.

This variety is characterized by having the striae easily resolved into
punctae. It was only found in one locality and then it was rare.
Pike County: 127 Shulze (A-2215).

Stauroneis anceps v. gracilis (Ehr.) Brun, 1880, Diat. Alpes et Jura et Région Suisse
et Francaise des Environs de Geneve, p. 89. pl. 9, fig. 2.
Stauroneis gracilis Ehr., 1843, Abhandl. K. Wiss. Berlin 1841: 423. pl. 1, fig. II, 14.

This variety though found in several localities was frequent in none.

Monroe County: Davy’s Run, 2015 Patrick (A-2111); Pocono Lake
Preserve, Mud Run, 2031 Hodge (A-2138); Pocono Lake, 2036 Patrick
(A-2148). Wayne County: Angels, 2022 Burke and Patrick (A-2125).

Stauroneis anceps v. linearis (Ehr.) Van Heurck, 1880, Syn. Diat. Belgique, p. 69.
pl. 4, figs. 7, 8.
Stauroneis linearis Ehr., 1843, Abhandl. K. Wiss. Berlin 1841: 423. pl. 1, fig. I, 11.

This variety was found only in one locality and then it was rare.
Monroe County: Pocono Lake Preserve, Pocono Lake, 2036 Patrick
(A-2148).

Stauroneis Kriegeri nomen nova
Valvae lineares 21 mu longae, 4 mw latae, apicibus constrictis capito-rostratis; striis
radiantibus, 26 in 10 u, area axiali angusta; stauro lineari.

This species was first observed by Krieger, who named it Stauroneis
pygmaea (Beitrage zur Naturdenkmalpflege 13: 272. pl. 2, fig. 28 (1929)).
However, this name is preoccupied by a species described by Castracana in
1886 (Voyage of H.M.S. Challenger, Botany 2: 25. pl. 29, fig. 7).

As Krieger points out, this species may vary between 20-23 » in length
and 4—5 » in breadth. He states the striae are 30 in 10 », but in my speci-
mens they are 26 in 10 ». The stauros may appear broader on one side of
the central nodule than on the other.

This species is named in honor of Mr. Krieger.

Monroe County: Davy’s Run, 2017 Patrick (A-2115). Wayne County:
Angels, 2023 Burke and Patrick (A-2127).

176 FaRLowIA, VoL. 2, 1945

Stauroneis phoenicenteron (Nitzsch.) Ehr., 1843, Abhandl. K. Akad. Wiss. Berlin
1841: pl. 2, fig. V, 7.
Bacillaria phoenicenteron Nitzsch. 1817, N. Schrift. Naturf. Ges. Halle 3: 92.

This species was common in the squeezings from surface vegetation in
shallow water of Wier Lake where the pH of the water was 6.6. It is also
common in some collections by Shulze from Pike County.

Luzerne County: Harvey Lake, 2042 Hodge (A-2160). Monroe County:
Jaggie’s Bog, 262-63 Shulze (A-2223-25); Pocono Lake Preserve, South
Pocono Lake, 2010 Patrick (A-2101); Raymondskill Falls, 2025 Patrick
(A-2131); Wier Lake, 2046 Hodge (A-2168). Northampton County:
Green Pond, 2050 Hodge (A-2176). Pike County: 127-29 Shulze (A-
2215-19), 2190 Shkulze (A-2236); Bushkill Creek, 1621 Shulze (A-2227).
Wayne County: Angels, 2021 Burke and Patrick (A-2124).

Stauroneis Smithii Grun., 1860, Verh. Zool. Bot. Ges. Wien 10: 464. pl. 4 (Grunow’s
pl. 6), fig. 16.
This species, occurring only in flowing water, was rare in the localities
given below.
Monroe County: Davy’s Run, 2018 Patrick (A-2117); Pike County:
Shohola Falls, 3-4 Burke (A-2190-92), 11 Burke (A-2202).

NAVICULA
A. Punctae of striae forming longitudinal and transverse rows perpendicular to each
OM ei iin wae ace ne aaeead Mane cnne ketene aciaa eases Naviculae Orthostichae
A. Punctae of striae forming decussating rows ........... Naviculae Decussatae

A. Punctae of striae not as above
B. Striae transversely lineate ..........00eeeceeeas Naviculae Lineolatae
B. Striae not transversely lineate
C. Raphe enclosed in silicious rib, terminal nodule thickened
Naviculae Bacillares
C. Raphe not enclosed
D. Central pores somewhat distant, axial area narrow or uniting
with central area to form a lanceolate space
Naviculae Entoleiae
D. Central pores not distant
E. Central area not distinct — small forms less than 20 pu
Naviculae Minusculae
E. Central area variously formed
F. Striae radiate throughout
G. Striae clearly alternately long and short at
central area, valves more or less elliptic in
shape ......... Naviculae Heterostichae
G. Striae may be uneven but not definitely
longer and shorter at central area — central
area quadrate or forming a transverse fascia
Naviculae Mesoleiae
F. Striae not radiate throughout, median striae dis-
tinctly more distant from each other than those in
other parts of the valve Naviculae Decipientes

PATRICK: Pocono DIATOMS eit

NAVICULAE ORTHOSTICHAE Cleve

Valves usually elongate, lanceolate to linear. The median line with
small or elongate central nodule, and with small or indistinct terminal
fissures.

Navicula cuspidata Kiitz., 1844, Bacill., p. 94. pl. 3, fig. 24.
Frustulia cuspidata Kiitz., 1833, Linnaea 8: 549. pl. 14, fig. 26.

This species was present only in one locality.
Luzerne County: Harvey Lake, 2042 Hodge (A-2161).

NAVICULAE DECUSSATAE Cleve

Valves elliptical with attenuate rostrate ends. The punctae of the striae
are arranged in obliquely decussating rows. There is only one species in
this region.

Navicula placenta Ehr., 1854, Mikrogeologie, p. 33. pl. 12, fig. 23.

This species, though found in several habitats, was very frequent in only
one, where it occurred on a wall dripping with water to the side of Shohola
Falls.

Pike County: 2188-89 Shulze (A-2234-35); Bushkill Creek, 1623 Shulze
(A-2231); Greeley, 1 Burke (A-2184), 2 Burke (A-2188); Shohola Falls,
14 Burke (A-2208-10). Wayne County: Angels, 2021 Burke and Patrick
(A-2123).

NAVICULAE LINEOLATAE Cleve

A. Terminal striae parallel or convergent
B. Striae usually 15 or more in 10 yw; valves usually less than 50 u long

C. Linear or slightly lanceolate with rounded ends, the median stria on
each side of the central nodule often decidedly longer than the other

striae about the central nodule ........ N. cincta and var.
C. Central area more or less rounded
D. Valve rhombic lanceolate ...... N. cryptocephala v. pumila

D. Valve lanceolate with attenuate, rostrate ends
N. cryptocephala

D. Valve lanceolate with acute ends N. radiosa v. tenella
B. Striae less than 15 in 10 u
E. Valves 30 u or less in length

F. Valves with strongly capitate ends, striae 8-10 in 10 u
N. hungarica v. capitata

F. Valves with slightly rostrate ends, striae 14 in 10 uw
N. cryptocephala v. veneta
E. Valves longer than 30 u
G. Terminal nodule of valve expanded to produce a hyaline space
N. globulifera
G. Terminal nodule not so expanded

178 FartowiA, VoL. 2, 1945

H. Ends of valve variously produced

I. Ends of valve attenuate-rostrate and slightly capi-
tate; valve lanceolate
N. rhynchocephala

I. Ends of valve cuneate; valve linear
N. viridula v. linearis
H. Ends of valve not produced; valve lanceolate
N. radiosa
A. Terminal striae radiate throughout
J. Striae at central area equally shortened
K. Central area more or less rounded

L. Valve lanceolate, striae at central area more distantly spaced
N. lanceolata

L. Valve ovate lanceolate with attenuate, rostrate ends
N. anglica

K. Central area transverse, central striae not more distantly spaced
N. dicephala
J. Striae at central area irregular in length
M. Striae at central area alternately longer and shorter and distinctly

more distantly spaced than in the rest of the valve
N. quadripartita

M. Striae not distinctly more distantly spaced about the central area
N. Striae 16-18 in 10 w; valve elliptical-lanceolate with distinctly

attenuate rostrate ends ........ N. decussis
N. Striae coarser; valve ovate-lanceolate, ends distinctly attenuate
POSETSUO essa e5 ec aaSadakanes aor N. exigua v. capitata n. var.

Navicula anglica Ralfs apud Pritchard, 1861, Syst. Hist. of the Infusoria, p. 900.

This species was rare in two collections.
Monroe County: Raymondskill Falls, 2025 Patrick (A-2131). Pike
County: 127 Shulze (A-2215).

Navicula cincta (Ehr.) Ralfs apud Pritchard, 1861, Syst. Hist. of the Infusoria, p. 901.
Pinnularia cincta Ehrenberg, 1854, Mikrogeologie, pl. 10, fig. I, 6.

For further discussion concerning the authorities for this name see Proc.
Acad. Nat. Sci. Phila. 95: 72.

This species was found only in one habitat and then it was rare.

Monroe County: Popomoning Lake, 2045 Hodge (A-2166).

Navicula cincta v. angusta (Grun.) Cl., 1893, K. Sv. Vet. Akad. Handl. 27 (3): 17.
Navicula angusta Grun., 1860, Verh. Bot. Zool. Ges. Wien 10: 528. pl. 3, fig. 19.
This variety was rare in both localities. The specimens in Tobyhanna
Creek were shorter than usual, being only 40 y» long.
Monroe County: Pocono Lake Preserve, Tobyhanna Creek, 2033 Patrick
(A-2142). Pike County: Shohola Falls, 14 Burke (A-2210).

Navicula cryptocephala Kiitz., 1844, Bacill., p. 95. pl. 3, fig. 26.

Monroe County: Trout Lake, 2049 Hodge (A-2175); Wier Lake, 2047
Hodge (A-2170); Raymondskill Falls, 2025 Patrick (A-2131). Northamp-

PatrRIcK: Pocono DIATOMS 179

ton County: Green Pond, 2050 Hodge (A-2176). Pike County: Lake
Wallenpaupek tributary, 5 Burke (A-2195); Shohola Falls, 3 Burke (A-
2190), 4 Burke (A-2192). Wayne County: Angels, 2022 Burke and Patrick
(A-2125), 2026 Burke and Patrick (A-2134).

Navicula cryptocephala v. pumila Grun. apud Van Heurck, 1880, Syn. Diat. Bel-
gique, pl. 8, fig. 6, 7.
Grunow suggests this is a variety but questions it. However, later writers
such as Cleve and Hustedt have given him credit for this combination.
Pike County: Lake Wallenpaupek tributary, 5 Burke (A-2095). Wayne
County: Angels, 2026 Burke and Patrick (A-2134).

Navicula cryptocephala v. veneta (Kiitz.) Rabh., 1864, Flora Europaea Algarum,
p. 198.
Navicula veneta Kiitz., 1844, Bacill., p. 95. pl. 30, fig. 76.

This variety was rare in the one locality.
Northampton County: Green Pond, 2050 Hodge (A-2176).

Navicula dicephala Ehr., 1838, Die Infusionsthierchen als volkommene Organismen,
p. 185.
This species was rare in the various localities in which it was encountered.
Monroe County: Raymondskill Falls, 2025 Patrick (A-2131). Pike
County: 2188 Shulze (A-2234); Bushkill Creek, 1623 Shulze (A-2231);
Shohola Falls, 12 Burke (A-2205).

Navicula decussis Oestrup, 1910, Danske Diat., p. 77. pl. 2, fig. 50.

My specimens are as Oestrup’s illustration except that the striae appear
to be somewhat curved. Hustedt in A. Schmidt’s Atlas Diatomaceen-
Kunde, pl. 398, figs. 36, 37, illustrates a diatom with curved striae and a
punctum in the central nodule which he assigns to this species.

I cannot be sure, from the descriptions and original illustrations, of the
difference between NV. decussis and N. Thingvallae. I have not seen the
types so will not try to make any critical evaluation of the two species.

This species was not of frequent occurrence in any of the habitats.

Monroe County: Raymondskill Falls, 2025 Patrick (A-2131). Pike
County: Lake Wallenpaupek tributary, 5 Burke (A-2195). Wayne County:
Angels, 2021 Burke and Patrick (A-2123-24).

Navicula exigua v. capitata var. nov.

Valvae elliptico-lanceolatae, apicibus rostrato-capitatis, 29 mw longae, 10 yu latae;
striis 14-15 in 10 yw, radiantibus, ad nodulum centralem longis inaequaliter; area axiali
angusta; area centrali transversa. Type: Shohola Falls, 10 Burke (A-2200).

The specimens vary in length from 22-30 y, in breadth from 10-11 u.
Hustedt has illustrated in A. Schmidt’s Atlas der Diatomaceen-Kunde,
pl. 398, figs. 24-26, specimens which seem to be the same as mine. He has
placed them under the species name. However, if one examines the original
figure and description of Gregory, it is clear that Hustedt’s specimens are
not the same as the type. Gregory (Quart. Jour. Micros. Sci. 2: 98-99.

180 FARLOWIA, VoL. 2, 1945

pl. 4, fig. 14) states, “its form is broadly elliptical with acute but not pro-
duced apices — the striae are distinct and radiate.”

Grunow in Van Heurck’s Synopsis des Diatomées Belgique (pl. 8, fig. 32)
and in Cleve and Moeller Type Slide no. 261 indicates that this species has
a transverse central area formed by the irregular shortening of the central
striae. Otherwise his specimens are as Gregory’s illustration. The question
as to whether Grunow’s and Gregory’s specimens are the same will have to
await the study of type or isotype material since Gregory makes no state-
ment concerning the central area.

Pike County: 2188 Shkulze (A-2234); Lake Wallenpaupek tributary, 5
Burke (A-2195); Shohola Falls, 10-11 Burke (A-2200-02), 14 Burke
(A-2210).

Navicula globulifera Hustedt, 1927, Archiv fiir Hydrobiol. 18: 164. pl. 5, fig. 7.

I believe this is the first record of this species in North America. My
specimens are 37-39 » long, 5.8—7 » broad, and the striae are 12 in 10 p.

Monroe County: Trout Lake, 2049 Hodge (A-2175). Pike County:
Greeley, 2 Burke (A-2188); Lake Wallenpaupek tributary, 5 Burke (A-
2195). Wayne County: Angels, 2021 Burke and Patrick (A-2123).

Navicula hungarica v. capitata (Ehr.) Cl., 1895, K. Sv. Vet. Akad. Handl. 27 (3): 16.
Pinnularia capitata Ehr., 1854, Mikrogeologie, pl. 37, fig. HIT, 9.

In 1848, Ehrenberg (Ber. K. Akad. Wiss. Berlin 1848: 18) just mentions
this species. In 1838 Ehrenberg (Infusionsthierchen als volkommene Or-
ganismen, p. 185, pl. 13, fig. 20) describes a Navicula capitata which is
quite different from the species in question.

Monroe County: Raymondskill Falls, 2025 Patrick (A-2131). Pike
County: Shohola Falls, 4 Burke (A-2192).

Navicula lanceolata (Ag.) Kiitz., 1844, Bacill., p. 94. pl. 30, fig. 48; pl. 28, fig. 38.
Frustulia lanceolata Ag., 1827, Flora (Bot. Zeitung) 10: 626.

Northampton County: Green Pond, 2050 Hodge (A-2177).

Navicula quadripartita Hustedt, 1937, Archiv fiir Hydrobiol. und Planktonk. Suppl.
15: 263.

This species is first published by Hustedt in 1934 in A. Schmidt’s Atlas
Diatomaceen-Kunde, p/. 400, figs. 12-15. According to the International
Rules of Botanical Nomenclature this is not a valid publication so the above
later record is given.

Hustedt states that this species is usually associated with moss.

Pike County: Bushkill Creek, 1623 Shulze (A-2231); Greeley, 2 Burke
(A-2188); Lake Wallenpaupek tributary, 5 Burke (A-2195).

Navicula radiosa Kiitz., 1844, Bacill., p. 91. pl. 4, fig. 23.

This species was very frequent from the squeezings of the surface vegeta-
tion of Mountain Lake.
Luzerne County: Harvey Lake, 2042-43 Hodge (A-2160-62). Monroe

PATRICK: Pocono DIATOMS 181

County: Mountain Lake, 2051 Hodge (A-2178); Popomoning Lake, 2045
Hodge (A-2166). Pike County: Lake Wallenpaupek tributary, 5 Burke
(A-2195); Shohola Falls, 10-11 Burke (A-2200-02).

Navicula radiosa v. tenella (Bréb.) Cl., 1895, K. Sv. Vet. Akad. Handl. 27 (3): 17.
Navicula tenella Bréb. ex Kiitz., 1849, Spec. Algarum, p. 74.

This species was of very frequent occurrence in certain habitats of Po-
cono Lake, Trout Lake, and Shohola Falls. It seems to prefer shallow
water and is associated with algae or moss.

Luzerne County: Harvey Lake, 2042-43 Hodge (A-2160-62). Monroe
County: Davy’s Run, 2015 Patrick (A-2111), 2019 Patrick (A-2119) ;
Mountain Lake, 2051 Hodge (A-2178); Pocono Lake Preserve, Pocono
Lake, 2035-36 Patrick (A-2146-48), 2065 Patrick (A-2182); Tobyhanna
Creek, 2032-33 Patrick (A-2140-42); South Pocono Lake, 2010 Patrick
(A-2101); Trout Lake, 2049 Hodge (A-2174). Pike County: Greeley, 1
Burke (A-2184); Shohola Falls, 3-4 Burke (A-2190-92), 10-11 Burke
(A-2200-02), 13 Burke (A-2206). Wayne County: Angels, 2021 Burke
and Patrick (A-2123).

Navicula rhynchocephala Kiitz., 1844, Bacill., pl. 30, fig. 35.

This species, though found in several localities was rare in all.

Luzerne County: Harvey Lake, 2042-43 Hodge (A-2160-62). Monroe
County: Raymondskill Falls, 2025 Patrick (A-2131). Pike County: Lake
Wallenpaupek tributary, 5 Burke (A-2195); Shohola Falls, 10 Burke
(A-2201). Wayne County: Angels, 2026 Burke and Patrick (A-2134).

Navicula viridula v. linearis Hustedt

This variety was first published in 1936 in A. Schmidt’s Atlas Diato-
maceen-Kunde #1. 405, figs. 13, 14. No description was present. Later,
1937, Archiv fiir Hydrobiologie, Supplement 15: 264 a description in Ger-
man was given. To date no Latin description has been furnished so I am
including one in order to validate the name.

Valvae lineares, 65-100 u longae, 12 u latae, apicibus cuneatis; striis 8—9 in 10 p, inter-
mediis radiantibus, terminalibus convergentibus, distincte transverse lineatis.

This species though rare in several localities was frequent in none.

Luzerne County: Harvey Lake, 2042 Hodge (A-2160). Monroe County:
Raymondskill Falls, 2025 Patrick (A-2131). Pike County: Shohola Falls,
10 Burke (A-2201). Wayne County: Angels, 2021 Burke and Patrick
(A-2123), 2026 Burke and Patrick (A-2134).

NAVICULAE BACILLARES Cleve

Navicula americana Ebr., 1843, Abhandl. K. Akad. Wiss. Berlin 1841: 417.

At Shohola Falls a form only 38 » long and 7 » broad was found. In no
case was this diatom of frequent occurrence.

182 FarLowiA, Vor, 2, 1945

Pike County: 127 Shulze (A-2215), 130 Shulze (A-2221); Shohola Falls,
10 Burke (A-2201).

Navicula bacillum Ehr., 1839, Abhandl. K. Akad. Wiss. Berlin 1838: 130.

This species was found in only two localities and was frequent in neither.
Luzerne County: Harvey Lake, 2043 Hodge (A-2162). Wayne County:
Angels, 2021 Burke and Patrick (A-2123).

NAVICULAE ENTOLEIAE Cleve

A. Frustules in filaments
B. Valves lanceolate, striae marginal .................. N. Keeleyi

B. Valve concave, striae not marginal.................. N. contenta v. biceps
A. Frustules not in filaments.
C. Valves concave, length of valve 20 uw or more, ends narrowed, rostrate
N. concava
C. Valves not concave
D. Striae more than 20 in 10 u
E. Valves swollen at center, about 15 «long . N. perpusilla

E. Valves linear-lanceolate, not swollen at center of valve over
20 Ge MOUS spe etoendottie es dcouala ete neead bak N. poconoensis

D. Striae less than 20 in 10 uw, ends of valve distinctly capitate
N. hassiaca

Navicula concava sp. nov.

Prate II, Fic. 6

Valvae utrimque concavae, 21.7 » longae, 5 uw ad nodulum centralem latae; apicibus
rostratis, area axiali late lanceolata; area centrali inaequali; striis 27-28 in 10 i, prope
medium radiantibus in apicibus parallelis. Type: Pike County; Greeley, squeezings
from Sphagnum in swamp, 2 Burke (A-2188).

This diatom is typical of the members of the Entoleiae except the striae
are parallel or only very slightly radiate at the ends. Cleve in describing
the group states that the striae are radiate at the ends. However, as the
species fits this group in all other respects, I think it well to include it until
a more thorough study is made.

This species occurred in only one locality, but was of frequent occurrence.

Pike County: Greeley, 2 Burke (A-2188-89).

Navicula contenta v. biceps (Arnott) Van Heurck, 1885, Syn. Diat. Belgique, p. 109.

Diadesmis biceps Arn. ex Cl. and MOll. no. 175. The determinations of these speci-
mens were made by Grunow.

This variety is rare in the two localities.
Pike County: Lake Wallenpaupek tributary, 5 Burke (A-2195); Shohola
Falls, 9 Burke (A-2198), 14 Burke (A-2208).
Navicula hassiaca Krasske, 1925, Abhandl. u. Ber. Ver. fiir Naturk. 56: 47. pl. 2,
fig. 26.

Pike County: Greeley, 1 Burke (A-2184); Lake Wallenpaupek tributary,
14 Burke (A-2210).

PATRICK: Pocono DIATOMS 183
Navicula Keeleyi sp. nov.

Prate II, Fic. 7

Valvae lanceolatae 19-30 u longae, 4-5 u latae, in fibris; apicibus capitato-rostratis,
area axiali et area centrali late lanceolata; striis marginibus, 30 in 10 uw. Type: Pike
County, Shohola Falls, scrapings from dripping rock face, side wall of gorge, above
concrete bridge crossing road below falls, 7 Burke (A-2197).

This species is named in honor of Mr. Frank J. Keeley who first called
my attention to this diatom. He found it in wet moss at Flat Rock, Mont-
gomery County, Pennsylvania. Mr. Keeley has also helped me in describ-
ing the species.

This species is very common in these collections. Its manner of growth
in filaments immediately calls it to one’s attention. It is distinguished from
the other filamentous forms of this group by its broad axial and central
areas which form a lanceolate space.

Pike County: Shohola Falls, 7 Burke (A-2196-97), 9 Burke (A-2198-
99).

Navicula perpusilla Grun., 1860, Verh. Zool. Bot. Ges. Wien 10: 552. pl. 4 (Grunow
pl. 2), fig. 7.

Hustedt states that this species is found especially in mountainous re-
gions. This species was very frequent on a dripping rock face at Shohola
Falls.

Pike County: Shohola Falls, 7 Burke (A-2196), 9 Burke (A-2198).

Navicula poconoensis sp. nov.

Pirate II, ric. 5

Valvae linearis-lanceolatae 26.4 u longae, 4.6 uw latae, apicibus late rostratis; area
axiali lanceolata, area centrali rotundata; striis 31 in 10 yw, in apicibus parallelis. Type:
Pike County, Greeley, squeezings from Sphagnum in swamp, 2 Burke (A-2188).

This species, as V. concava, differs from other members of the Entoleiae
in that the terminal striae are parallel instead of radiate, but it agrees in
all other respects with members of this group.

The species is most closely related to N. Flotowdi but differs from it in
the size of the valve, the shape of the apices, and the fact that it does not
form filaments.

Pike County: Greeley, 2 Burke (A-2188-89).

NAVICULAE MINUSCULAE Cleve

A. Valve elliptical in outline, striae about 30 in 10 wu
B. Striae parallel or slightly radiate 1.1.1... 0... cece eee eee eters N. muralis
B. Striae strongly radiate <2 sine. ert siete: oe ee cee N. atomus

A. Valve linear-elliptic with rostrate ends, striae about 30 in 10 w. As seen through
the microscope the striae at margin of valve are on a different level of focus than

At raphe cs cu oss sacs Oe ey ocr oe hs N. festiva

184 FARLowIA, VOL. 2, 1945

Navicula atomus (Kiitz.) Grun., 1860, Verh. Zool. Bot. Ges. Wien 10: 552. pl. 2, fig. 6.
Amphora atomus Kitz., 1844, Bacill., p. 108. pl. 30, fig. 70.

This species was only found in one locality.
Pike County: Shohola Falls, 11 Burke (A-2202).

Navicula festiva Krasske, 1925, Abhandl. u. Bericht. Ver. fiir Naturk. Cassel 56: 47.
pl. 1, fig. 16.

The name Navicula vitrea (Oestrup) Hustedt cannot be applied to this
species as it is preoccupied by N. vitrea Cleve, 1893, (K. Sv. Vet. Akad.
Handl. 26 (2): 111).

This species was of very frequent occurrence at Greeley, where it was
found in association with Sphagnum.

Pike County: Greeley, 2 Burke (A-2188).

Navicula muralis Grun. apud Van Heurck, 1880, Syn. Diat. Belgique, pl. 14, fig. 27.

This species was rare in the one locality.
Monroe County: Trout Lake, 2049 Hodge (A-2174).

NAVICULAE HETEROSTICHAE Cleve

A. Valve elliptical in shape, central area large and rounded... .N. scutiformis
A. Valve lanceolate to rhombic-elliptical, central area indistinct .N. cocconeiformis
A. Valve orbicular, central area indistinct, less than 20 u long. .N. pseudoscutiformis

Navicula cocconeiformis Greg., 1856, Quart. Jour. Micros. Sci. 4: 6. pl. 1, fig. 22.

This species was only found in running water, but it was not of frequent
occurrence in any habitat wherein it was found.

Monroe County: Davy’s Run, 2014 Patrick (A-2110), 2016-18 Patrick
(A-2113-18), 2056 Patrick (A-2180); Pocono Lake Preserve, Tobyhanna
Creek, 2033 Patrick (A-2142). Pike County: Shohola Falls, 10-11 Burke
(A-2201-02), 13 Burke (A-2206).

Navicula pseudoscutiformis Hustedt apud Pascher, 1930, Die Siissw. Flora Mittel-
europ., Heft 10: 291. fig. 495.

This species was rare in the localities where it was found.

Monroe County: Davy’s Run, 2019 Patrick (A-2119); Wier Lake, 2047
Hodge (A-2170). Pike County: Shohola Falls, 11 Burke (A-2202), 14
Burke (A-2210).

Navicula scutiformis Grun., 1881, in A. Schmidt’s Atlas Diat., pl. 70, fig. 62.
This species was found in only one locality.
Monroe County: Davy’s Run, 2019 Patrick (A-2119).
NAVICULAE MESOLEIAE Cleve

The valve is symmetrical, linear to elliptical with usually obtuse or
rostrate ends. The axial area is narrow or indistinct. The central area is

Patrick: Pocono DIATOMS 185

large, quadrate or a transverse fascia. The striae are fine, punctate and
radiate throughout. The connecting zone is not complex.

A. Valves with isolated stigma in the central area; valve often somewhat lanceolate in
shape and may have rostrate capitate ends...............-.-- N. mutica

A. Valves without an isolated stigma in the central area, ends of valve broadly rounded.

B. Terminal nodules laterally expanded to form a clear area . N. pupula and vars.

B. Terminal nodules not so expanded
C. Terminal striae radiate and curved.............. N. bacilliformis
C. Terminal striae not so formed
D. Median striae at the central area approximately the same length
so that the central area appears clear-cut but small
N. minima var.

D. Median striae of central area very irregular in length

E. Very small forms, almost orbicular in shape
N. subatomoides

E. Valves over 12 u long, elliptical in shape

F. Central area broad, somewhat rhomboidal in shape
N. variostriata

F. Central area narrowly rectangular in shape
N. Rotaeana
Navicula bacilliformis Grun., 1880, K. Sv. Vet. Akad. Handl. 17 (2): 44. pl. 2, fig. 51.

This species was rare in the two localities cited below.
Monroe County: Popomoning Lake, 2045 Hodge (A-2166); Wier Lake,
2047 Hodge (A-2170). ;
Navicula minima v. atomoides (Grun.) Cl., 1894, K. Sv. Vet. Akad. Handl. 26 (2):
128.
Navicula atomoides Grun. apud Van Heurck, 1880, Syn. Diat. Belgique pl. 14,
fig. 12, text (1885) p. 107. Van Heurck Type Slide no. 219.
This species was very frequent in the bottom vegetation in shallow water
in Wier Lake and Davy’s Run and in scrapings from rocks at Shohola Falls.
Monroe County: Davy’s Run, 2016-17 Patrick (A-2113-15); Pocono
Lake Preserve, Tobyhanna Creek, 2032 Patrick (A-2140); Pocono Lake,
2037 Patrick (A-2150); Wier Lake, 2047 Hodge (A-2171). Pike County:
Shohola Falls, 14 Burke (A-2208).

Navicula mutica Kiitz., 1844, Bacill., p. 93. pl. 3, fig. 32.

This species was rare.
Pike County: Shohola Falls, 10 Burke (A-2200).

Navicula pupula Kiitz., 1844, Bacill., p. 93. pl. 30, fig. 40.

Though found in several localities this species was only frequent in the
squeezings from moss collected in the spillway of Mud Run.

Monroe County: Davy’s Run, 2014 Patrick (A-2109), 2019 Patrick
(A-2119); Raymondskill Falls, 2025 Patrick (A-2131); Pocono Lake
Preserve, Mud Run, 2031 Hodge (A-2138); Pocono Lake, 2036 Patrick
(A-2149). Northampton County: Green Pond, 2050 Hodge (A-2176).
Pike County: 127 Shulze (A-2215).

186 Fartowia, Vor. 2, 1945

Navicula pupula v. bacillaroides Grun. apud Cl. and Grun., 1880, K. Sy. Vet. Akad.
Handl. 17 (2): 45.

I do not believe as Hustedt does that this variety and var. rectangularis
of Grunow should be united. I have seen Cleve and Moeller’s Type Slide
No. 86 which has on it var. rectangularis. These specimens are quite distinct
from those of var. bacillaroides. The sides of the valve are linear or some-
what concave while those of var. bacillaroides are somewhat convex. Also
var. bacillaroides seems to have a more robust structure than var. rectan-
gularts.

This variety was only encountered once in this study.

Northampton County: Green Pond, 2050 Hodge (A-2176).

Navicula pupula v. capitata Hustedt apud Pascher, 1930, Siissw. Flora Mitteleurop.,
Heft 10: 281. fig. 467c.

Though found in several localities this variety was always rare.

Monroe County: Pocono Lake Preserve, Pocono Lake, 2065 Patrick
(A-2182); Raymondskill Falls, 2025 Patrick (A-2131). Pike County:
Shohola Falls, 11-12 Burke (A-2202-05). Wayne County: Angels, 2021
Burke and Patrick (A-2123).

Navicula Rotaeana (Rabh.) Van Heurck, 1880, Syn. Diat. Belgique, pl. 14, fig. 17.
Stauroneis Rotaeana Rabh., 1852, Hedwigia 1: 103. pl. 13, fig. 7.

Though found in many habitats this species was common only in Davy’s
Run. Here it was often found associated with the mosses Hypnum and
Fontinalis,

Luzerne County: Harvey Lake, 2042 Hodge (A-2160). Monroe County:
Davy’s Run, 2014-19 Patrick (A-2110-19); Pocono Lake Preserve, Toby-
hanna Creek, 2032-33 Patrick (A-2140-42); Pocono Lake, 2035 Patrick
(A-2147); Spring Run, 2037 C. Myers (A-2150); Trout Lake, 2049 Hodge
(A-2174). Pike County: Lake Wallenpaupek tributary, 5 Burke (A-2195) :
Shohola Falls, 3-4 Burke (A-2190-92), 13 Burke (A-2206). Wayne
County: Angels, 2021 Burke and Patrick (A-2123).

Navicula subatomoides Hustedt, 1936, in A. Schmidt’s Atlas der Diatomaceen-Kunde,
pl. 404, figs. 33-35.

As this species has not been validly described I am supplying the fol-
lowing description.

Valvae ellipticae 6-7 w longae, 4-5 mw latae, area axiali anguste, area centrali lata
stauroneiformi; striis 44 in 10 yu, radiantibus.

This species was very common in the squeezings from Hypnum and
Fontinalis in Davy’s Run.

Monroe County: Davy’s Run, 2019 Patrick (A-2119). Pike County:
Lake Wallenpaupek tributary, 5 Burke (A-2195).

Navicula variostriata Krasske, 1923, Bot. Archiv. 3: 197. fig. 12.

This species was rare in the localities in which it was found. Hustedt

Patrick: Pocono D1aToms 187

states that it is especially found associated with Sphagnum. This did not
seem to be necessarily the case in the habitats in which it was found in
this study.

Pike County: Bushkill Creek, 1623 Skulze (A-2231); Greeley, 2 Burke
(A-2188); Shohola Falls, 14 Burke (A-2209-10).

NAVICULAE DECIPIENTES Cleve
Navicula halophila v. minor Kolbe, 1927, Pflanzenforschung 7: 67. pl. 1, fig. 4.

Kolbe states that this variety is a “mesohalobien.” If I am correct in
believing that my specimens belong to this variety, then it should be noted
that this variety also is found in somewhat acid, shallow, fresh water. I
have not seen any authenticated specimens of var. minor, but a careful
analysis of Kolbe’s figure would indicate that my specimens are the same.
My specimens are about 17 u long, 6 » broad, and the striae are 19 in 10 p.

Monroe County: Pocono Lake Preserve, South Pocono Lake, 2010 Pat-
rick (A-2101); Tobyhanna Creek, 2032 Patrick (A-2140). Pike County:
Shohola Falls, 4 Burke (A-2192), 11 Burke (A-2202).

Navicula semen Ebr., 1843, Abhandl. K. Akad. Wiss. Berlin, 1841: 419. pl. 1, fig. IT,
17-a?; pl. 4, fig. I, 8.

This species was very frequent in two localities or collections of Shulze
which were simply indicated as Pike County.

Monroe County: Jaggie’s Bog, 262-63 Shulze (A-2223-25). Pike
County: 128-129 Shulze (A-2217-19).

PINNULARIA EHRENBERG

A. Striae distinctly separated from each other.................000-- Distantes
A. Striae close to each other
B. Striae short, perpendicular or almost so to the raphe
C. Large forms, striae typically less than 15 in 10 uw... .Brevistriatae
C. Small forms, striae more than 15 in 10 yw............ Parallelestriae
B. Striae not perpendicular to the raphe nor typically short
C. Raphe linear

D. Valves with capitate or rostrate ends which are narrower than
the rest of the valve, usually small forms..... Capitatae

D. Valves with ends not distinctly narrower than rest of valve
E. Striae somewhat radiant in the middle and convergent at
the ends, larger forms, usually over 40 » long
Tabellariae
E. Striae strongly radiant in the middle and convergent at
the ends, raphe at terminal nodules bayonet-shaped
Divergentes
C. Raphe broad
Bs Simple sence seer ee sien tave cles aos ete eteects Maiores

Fi Twisted aces gene tains earths ons 6 crane esis Complexae

188 FarRLowIA, VoL. 2, 1945
DISTANTES Cleve subgenus

Pinnularia borealis Ehr., 1843, Abhandl. K. Akad. Wiss. Berlin 1841: 420. pl. 1, fig. IJ,
6; pl. 4, fig. I, 5; pl. 4, fig. V, 4.
This species was frequent on dripping rocks at Shohola Falls. Pike
County: 128 Shulze (A-2217); Shohola Falls, 9 Burke (A-2198).

BREVISTRIATAE subgenus

A. Valve triundulate, ends rostrate-capitate, axial area granular
P. nodosa

A. Valves not triundulate
B. Valves more or less swollen at center, ends capitate, axial area granular
P. acrosphaeria and var.

B. Valve linear, ends rounded, axial area clear....... P. brevicostata
Pinnularia acrosphaeria W. Smith, 1853, Brit. Diat., 1: 58. pl. 19, fig. 183.

This species, though found in several localities in Pike County, was never
of frequent occurrence.

Pike County: 2190 Shulze (A-2236), 128-130 Shulze (A-2217-21);
Bushkill Creek, 1621 Shulze (A-2227); Greeley, 2 Shulze (A-2188);
Shohola Falls, 3 Burke (A-2190), 13 Burke (A-2206), 14 Burke (A-2210).

Pinnularia acrosphaeria var. turgidula Grun. apud Cleve, 1895, K. Sv. Vet. Akad.
Hand]. 27 (3): 86.
This very distinctive species was never of frequent occurrence.
Monroe County: Pocono Lake Preserve, Pocono Lake, 2035 Patrick
(A-2146); Raymondskill Falls, 2025 Patrick (A-2131). Pike County:
Shohola Falls, 3 Burke (A-2190).

Pinnularia brevicostata Cleve, 1891, Acta Soc. pro Fauna et Flora Fennica 8 (2): 25.
pl. 1, fig. 5.
This species was only found once and then it was rare. It may have been
a contamination.
Pike County: 2190 Shulze (A-2236).

Pinnularia nodosa (Ehr.) W. Smith, 1856, Brit. Diat., 2: 96.
Navicula nodoso Ehr., 1838, Die Infusionsthierchen als volkommene Organismen,
p. 179. pl. 13, fig. 9.

This species was of common occurrence in a locality given by Shulze
only as Pike County.

Monroe County: Jaggie’s Bog, 262 Shulze (A-2223). Pike County:
127-128 Shulze (A-2215-17); 2188 Shulze (A-2233); Bushkill Creek,
1621-23 Shulze (A-2227-31); Greeley, 1 Burke (A-2184); Shohola Falls,
12 Burke (A-2205).

PARALLELISTRIATAE Cleve subgenus

Pinnularia fasciata (Lagst.) Hustedt apud Pascher, 1930, Siissw. Flora Mitteleurop.,
Heft 10: 316. fig. 569.

Patrick: Pocono DIATOMS 189

Navicula fasciata Lagst., 1873, Bihang K. Sv. Vet. Akad. Handl. 1 (14): 34. pl. 2,
fig. 11.

This species was rare in one locality.
Pike County: Shohola Falls, 14 Burke (A-2210).

Pinnularia sublinearis (Grun.) Cl., 1895, K. Sv. Vet. Akad. Handl. 27 (3): 74.

Navicula sublinearis Grun. apud Van Heurck, 1880, Syn. Diat. Belgique, pl. 6,
figs. 25, 26, text (1885) p. 76.

This species was rare in the one locality given below.
Wayne County: Angels, 2017 Burke and Patrick (A-2115).

CAPITATAE Cleve subgenus

A. Margins of valve triundulate with central area forming a stauros
P. mesolepta v. stauroneiformis

A. Margins of valve not triundulate
B. Valves lanceolate, axial area broad

C. Valve 30 or more yu long, striae 11-12 in 10 uw
P. Braunii v. amphicephala

C. Valve 20 u long, striae 17 in 10 » P. Burkei
B. Valves linear, ends of valve distinctly capitate
D. Striae almost parallel, slightly radiate at middle and convergent at ends
E. Central area a broad fascia P. Hilseana

E. Central area only slightly broader than the axial area ~
P. subcapitata

D. Striae distinctly radiate at center of valve, convergent at ends, central
area rhomboidal sometimes forming a stauros
P. interrupta
B. Valves linear, ends subcapitate

F. Striae gradually shortened to form a very broad fascia
P. subcapitata v. paucistriata

F. Striae usually abruptly shortened to form a transverse fascia
P. subcapitata v. stauroneiformis

Pinnularia Braunii v. amphicephala (Mayer) Hustedt apud Pascher, 1930, Siissw.-
Flora Mitteleurop., Heft 10: 319. fig. 578.
Pinnularia amphicephala Mayer, 1917, Denks. K. Bayern Bot. Ges. in Regensburg
13: ns. 7: 136. pl. 2, figs. 15, 16.

This species was only found in one locality and then it was rare.
Monroe County: Pocono Lake Preserve, Davy’s Run, 2014 Patrick
(A-2109).

Pinnularia Burkei sp. nov.
Pirate III, Fic. 1
Valvae lanceolatae apicibus rostrato-capitatis, 19 « longae, 3 uw latae, striis 17 in 10 p,
radiantibus, apice versus convergentibus. Type: Pike County: Greeley, 2 Burke
(A-2189).
I place this species in the Capitatae with some hesitation. It has the
characteristics of the species typically included in this group except for the

190 FaRLowlA, VoL. 2, 1945

axial area. Typically, this group has a narrow axial area. However, if we
include Pinnularia Brauni v. amphicephala within it, I believe we are
justified in including this species.

This species is named in honor of Mr. Joseph F. Burke of the New York
Botanical Gardens who has given me many diatom collections from Pike
County.

This species was only found in two localities.

Pike County: Greeley, 2 Burke (A-2188-89); Shohola Falls, 14 Burke
(A-2210).

Pinnularia Hilseana Janisch apud Hilse, 1860, Jahres-Ber. Schles. Ges. Vaterl. Cult.
36: 82.

The identification of these specimens has been checked with Rabenhorst’s
Algen. Sachs. resp. Mitteleurop. No. 953. This species was found only in
association with Sphagnum. Because it has this very distinctive habitat as
well as definite taxonomic characters, I do not believe it wise to consider it
a variety of P. subcapitata, but think it should be kept as a separate species.

This species was of very frequent occurrence in Lake Tamaque.

Monroe County: Pocono Lake Preserve, South Pocono Lake, 2011-12
Patrick (A-2103-05); shallow ditch on way to Davy’s Run, 2013 Patrick
(A-2108); Lake Tamaque, 2040 Hodge (A-2156).

Pinnularia interrupta W. Sm., 1853, Brit. Diat., 1: 59. pl. 19, fig. 184.

Though found in several localities this species was rare.

Lackawanna County: Indian Lake, 2044 Hodge (A-2165). Monroe
County: Pocono Lake Preserve, Davy’s Run, 2014 Patrick (A-2110), 2019
Patrick (A-2119), South Pocono Lake, 2010 Patrick (A-2102); Wier Lake,
2046 Hodge (A-2169). Northampton County: Green Pond, 2050 Hodge
(A-2176). Pike County: Greeley, 1-2 Burke (A-2184-88); Shohola Falls,
10 Burke (A-2201), 12 Burke (A-2204).

Pinnularia mesolepta v. stauroneiformis (Grun.) Cl., 1895, K. Sv. Vet. Akad.
Handl. 27 (3): 76.

Navicula mesolepta v. stauroneiformis Grun., 1860, Verh. Zool. Bot. Ges. Wien
10: 520. pl. 4 (Grunow il. 2) fig. 22b.

This variety was rare in one locality.
Pike County: 127 Shulze (A-2215).

Pinnularia subcapitata Greg., 1856, Quart. Jour. Micros. Sci., 4: 9. pl. 1, fig. 30.

This species, as figured by Gregory, has distinctly capitate ends and an
indistinct central nodule. Certainly it does not form a stauros. Some
workers have confused this typical species with its variety stauroneiformis.
They have called specimens with a stauros and subcapitate ends by the
specific name. A study of Gregory’s original plate will show that the
diatom he was describing is distinctly different from the stauroid variety.

Monroe County: Lake Tamaque, 2039 Hodge (A-2154).

Patrick: Pocono DIATOMS 191

Pinnularia subcapitata v. paucistriata Grun. apud Van Heurck, 1880, Syn. Diat.
Belgique, pl. 6, fig. 23.

This diatom was only found once, but then it was of frequent occurrence.
Pike County: Shohola Falls, 3 Burke (A-2190).

Pinnularia subcapitata v. stauroneiformis Van Heurck, 1880, Syn. Diat. Belgique,
pl. 6, fig. 22.

This variety was found in several habitats but was frequent in none.

Monroe County: Pocono Lake Preserve, Davy’s Run, 2056 Patrick
(A-2180), Spring Run, 2037 C. Myers (A-2150). Pike County: Greeley,
1-2 Burke (A-2186-88). Wayne County: 2021 Burke and Patrick (A-
2123), 2024 Burke and Patrick (A-2129).

TABELLARIAE Cleve subgenus

In many instances it is very difficult to distinguish this group from the
Divergentes. As I have not studied these groups critically, I shall follow
the classification of Cleve and Hustedt.

A. Valves with lunate markings on each side of the central nodule P. stomatophora
A. Valves without such markings

B. Valves linear, apices rounded, striae strongly radiate at center of valve,
gradually shortened to form a stauros............... P. substomatophora

B. Valves linear or linear-lanceolate with somewhat rostrate-capitate ends
P. gibba and var.

Pinnularia gibba Ehr., 1843, Abhandl. K. Akad. Wiss. Berlin 1841: pl. 1, fig. II, 8;
pl. 3, fig. I, 4.

Pirate III, ric. 2-5

I have not been able to find the original reference to Pinnularia gibba.
In the above cited article, Ehrenberg gives four illustrations of P. gibba.
One illustration, Pl. 7, fig. 17, 3, Ehrenberg questions as being this species.
The other three illustrations seem to refer to two distinctly different
diatoms. Two illustrations, Pl. 1, fig. 17, 8, and Pl. 3, fig. I, 4, are more as
the present concept of P. gibba. The third illustration, Pl. 2, fig. I, 24, is
hard to identify as being any form recognized to-day. Ehrenberg also
describes in the above reference, p. 423, Pl. 1, fig. II, 3, Stauroptera gibba.
He states that this is in some ways like P. gibba, but is a doubtful form.
I have included in my plates reproductions of Ehrenberg’s figures. Ehren-
berg states the magnification is 300 times.

As to my specimens, two diatoms were found which belong to the P.
gibba series. One was like the specimens on H. L. Smith Spec. Typ. No.
275 which is probably the typical form. The specimens which I found
were always in association with Sphagnum.

This species was very frequent in a composite collection made by Mr.
Frank Myers from Pocono Lake.

Monroe County: Lost Lake, 2020 Turner (A-2121); Pocono Lake Pre-

192 FarLow1A, VoL. 2, 1945

serve, Pocono Lake, F. Myers (A-2213), South Pocono Lake, 2011 Patrick
(A-2103).

Pinnularia gibba v. rostrata var. nov.

Prate III, Fic. 6

Valvae lineares, 69 « longae, 10 u latae, apicibus rostratis; area axiali lata; striis
9 in 10 », intermediis parum radiantibus, terminalibus convergentibus. Type: Monroe
County: Pocono Lake, F. Myers (A-2213).

Monroe County: Mountain Lake, 2051 Hodge (A-2178); Pocono Lake
Preserve, Pocono Lake, 2034 Patrick (A-2144), F. Myers (A-2213); Wier
Lake, 2046 Hodge (A-2169).

Pinnularia stomatophora (Grun.) Cleve, 1895, K. Sv. Vet. Akad. Handl. 27 (3): 83.

Navicula stomatophora Grun., 1876, in A. Schmidt’s Atlas Diatomaceen-Kunde,
pl. 44, figs. 24, 26.

Though found in several localities this species was rare in all.
Pike County: 128 Shulze (A-2217), 2190 Shulze (A-2236); Shohola
Falls, 14 Burke (A-2208).

Pinnularia substomatophora Hustedt, 1935, Archiv fiir Hydrobiol. und Planktonk.
Suppl. 14: 160. pl. 2, fig. 14.

This species is present in several localities but frequent in none.

Monroe County: Pocono Lake Preserve, Pocono Lake, F. Myers (A-
2213); Wier Lake, 2046-47 Hodge (A-2169-70). Pike County: Greeley,
2 Burke (A-2188); Shohola Falls, 4 Burke (A-2192), 13 Burke (A-2206).

DIVERGENTES Cleve subgenus

This group seems to be closely allied to the Tabellariae and in some
cases it is very hard to decide in which group a species belongs. As I have
not made a critical study of these groups, I have followed the classification
of Cleve.

A. Central area with a horseshoe-shaped thickening on each side of the central nodule
P. divergens and vars.
A. Central nodule without such thickenings

B,. Edge of valve triundulate ..5:...dsdviwnsdesaveous P. legumen
B. Valve linear, narrowed toward the ends
C. Length less than 25 wi... eee eee eee P. obscura
C. Length over 40m cc cisisseesaresicccasavaes P. Brebissonii

Pinnularia Brebissonii (Kiitz.) Rabenhorst, 1864, Flora Europea Algarum, p. 222.
Navicula Brebissonii Kiitz., 1844, Bacill., p. 93. pl. 3, fig. 49; pl. 30, fig. 39.

Kiutzing says that this species is the same as Frustulia bipunctata Breb.
Cons, Diat., p. 18 (sec. specim.). Miss Hilda F. Harris has verified this
reference for me, and states that no description or illustration is given.
Therefore even if it is the same as Frustulia bipunctata, the name Pinnu-
laria Brebissonii is valid.

PaTRIcK: Pocono DIATOMS 193

Monroe County: Pocono Lake Preserve, Pocono Lake, F. Myers (A-
2213). Pike County: 128 Shulze (A-2217), 2188 Shulze (A-2234).

Pinnularia divergens W. Sm., 1853, Brit. Diat., 1: 57. pl. 18, fig. 177.

This species though found in several localities is frequent in none.

Monroe County: Davy’s Run, 2015 Patrick (A-2111), 2017-18 Patrick
(A-2115-18); Pocono Lake Preserve, Pocono Lake, F. Myers (A-2213),
2065 Patrick (A-2182); South Pocono Lake, 2010 Patrick (A-2102);
Tobyhanna Creek, 2032 Patrick (A-2140). Pike County: 2190 Shulze
(A-2236); Greeley, 1 Burke (A-2184).

Pinnularia divergens v. parallela (Brun) Freng., 1924, Anales Soc. Cient. Argen-
tina 97: 102.

Pinnularia parallella (divergens W. Sm. var.) Brun, 1895, Le Diatomiste 2: pl. 14,
fig. 7.

This species was found twice but was not frequent in either locality.
Pike County: 129 Shulze (A-2219), 2190 Shulze (A-2236).

Pinnularia legumen Ehbr., 1843, Abhandl. K. Akad. Wiss. Berlin, 1841: pl. 4, fig. I, 7.

This species is also mentioned in the Ber. K. Akad. Wiss. Berlin 1843:
255, but is not described. There may be an earlier description which I
have not found. I have checked this identification with Van Heurck’s Type
Slide no. 69.

This species was found only in one locality and then it was rare.

Pike County: Shohola Falls, 12 Burke (A-2205).

Pinnularia obscura Krasske, 1932, Hedwigia 72: 117. pl. 3, fig. 22.

In his description Krasske states that the striae are 10 in 10 ». In my
specimen they are 12 in 10 yp, otherwise it is the same as Krasske’s descrip-
tion. This species was found in only one locality but there it was very
frequent.

Pike County: Greeley, 2 Burke (A-2188).

Mayrores Cleve subgenus

Pinnularia Clevii sp. nov.
Pirate III, Fics. 7-10

Valvae bis constrictae, 112 « longae, ad medium 16 yp latae; apicibus leviter rostratis ;
costis 9 in 10 u, centro aliquid radiantibus, in apicibus convergentibus, area axiali lata,
raphe simplice lata. Type: Monroe County: Jaggie’s Bog, 263 Shulze (A-2225).

This species was first observed by P. T. Cleve and for that reason I am
naming it after him. In 1891 (Acta Soc. pro Fauna et Flora Fennica
8 (2): 24. pl. 1, fig. 3) Cleve referred this species to Pinnularia essox
Ehrenberg. An examination of Ehrenberg’s original plate (Abhandl. K.
Akad. Wiss. Berlin 1841: pl. J, fig. JJ, 4) shows clearly that these two spe-
cies are not the same. Ehrenberg’s Pinnularia essox was no doubt the same
as Navicula essox on H. L. Smith’s Species Typicae No. 272. This species

194 FARLowI1A, Vor. 2, 1945

really belongs to the genus Caloneis. I have included a copy of Ehrenberg’s
and Cleve’s original figures. Also I have included a photograph of one of
the specimens of H. L. Smith’s Species Typicae No. 272.

This species was only found in one locality.

Monroe County: Jaggie’s Bog, 262-63 Shulze (A-2223-2225).

Pinnularia maior (Kiitz.) W. Sm., 1853, Brit. Diat., 1: 54. pl. 18, fig. 162.
Frustulia maior Kiitz., 1833, Syn. Diat., p. 547. pl. 14, fig. 25.

This species was found in several localities but was frequent in none.

Monroe County: Davy’s Run, 2016 Patrick (A-2113); Pocono Lake
Preserve, Pocono Lake, F. Myers (A-2213). Pike County: 2188-90 Shulze
(A-2234-36); Greeley, 1 Burke (A-2184).

Pinnularia maior v. transversa (A. Schmidt) Cl., 1895, K. Sv. Vet. Akad. Handl.
27 (3): 90.
Navicula transversa A. Schmidt, 1876, Atlas Diat., pl. 43, figs. 5, 6.

This variety was rare in the localities where it was found.
Monroe County: Pocono Lake Preserve, Pocono Lake, F. Myers (A-
2213). Pike County: 129 Shulze (A-2219), 2190 Shulze (A-2236).

CoMPLEXAE Cleve subgenus

A. Striae usually 5 or less in 10 u

B. Valves broader at the middle and ends, thus producing a swollen appearance
P. nobilis

B. Valves linear with ends slightly narrower, but broadly rounded
P. streptoraphe

B. Striae usually more than 6 in 10 u

C. Valves linear, with rounded ends; ends as broad if not broader than the rest
OF THE Valve 5p cicaigegaew dosed teas ems a paeateegeeekie’ P. gentilis

C. Valves linear or very slightly lanceolate, gradually diminishing in width to-
wards the ends of the valve, thus producing in some cases a slightly elliptical
effect, the band across the striae very variable as to its appearance

P. viridis and vars.
Pinnularia gentilis (Donk.) Cl., 1895, K. Sv. Vet. Akad. Handl. 27 (3): 92.
Navicula gentilis Donk., 1870-73, Nat. Hist. Brit. Diat. p. 69. pl. 12, fig. 1.

This species was very frequent in Jaggie’s Bog, Monroe County.

Monroe County: Jaggie’s Bog, 262-63 Shulze (A-2223-25).
Pinnularia nobilis Ehr., 1840, Ber. K. Akad. Wiss. Berlin 1840: 214.

This species was very frequent in a collection made by Shulze which was
simply designated as Pike County. It was also common in a collection
from Jaggie’s Bog.

Monroe County: Jaggie’s Bog, 263 Shulze (A-2225). Pike County:
128-29 Shulze (A-2217-19).

Pinnularia streptoraphe Cl., 1891, Acta Soc. pro Fauna et Flora Fennica 8 (2): 23.

This species was common in a collection made by Shulze which was
simply designated as Pike County.

Patrick: Pocono DIATOMS 195

Monroe County: Pocono Lake Preserve, Pocono Lake, F. Myers (A-
2213). Pike County: 128-29 Shulze (A-2217-19); Greeley, 1 Burke
(A-2184).

Pinnularia viridis (Nitzsch) Ehr., 1843, Abhandl. K. Akad. Wiss. Berlin 1841: pl. 1,
aa ye rhe a Ub (cio fl see
Bacillaria viridis Nitzsch, 1817, N. Schrift. Naturf. Ges. Halle 3 (1): 99. pl. 6,
figs. 1-3. F

This species was very frequent in a composite sample of collections
made by Myers from Pocono Lake. Actually it may not be so frequent in
any single habitat.

Monroe County: Davy’s Run, 2013-14 Patrick (A-2107-10); Jaggie’s
Bog, 263 Skulze (A-2225); Pocono Lake Preserve, Pocono Lake, F. Myers
(A-2213); Wier Lake, 2046 Hodge (A-2169). Pike County: 127-29
Shulze (A-2215-19), 2188-90 Shulze (A-2234-36); Bushkill Creek, 1621
Shulze (A-2227); Shohola Falls, 3 Burke (A-2190), 12 Burke (A-2204).

Pinnularia viridis v. elliptica Meister, 1912, Beitrage zur Krypt.-Flora Schw., 4 (1):
150. pl. 23, fig. 4.
This variety occurred only in one locality and then it was rare.
Monroe County: Jaggie’s Bog, 263 Shulze (A-2225).

Pinnularia viridis v. fallax Cl., 1895, K. Sv. Vet. Akad. Handl. 27 (3): 91.

This variety occurred only once and then it was rare.
Monroe County: Davy’s Run, 2017 Patrick (A-2115).

Pinnularia viridis v. sudetica (Hilse) Hustedt apud Pascher, 1930, Siissw. Flora
Mitteleurop. Heft 10: 335.
Pinnularia sudetica Hilse, 1860, Jahres-Berichte Schles. Ges. Vaterl. Cult. 36: 82.

As this name is older than P. commutata for this variety it is the one
that should be used. It is doubtful to me that this form should be con-
sidered a variety of viridis because the raphe is simple instead of being
complex as is characteristic for this group. However Grunow, Cleve, and
Hustedt considered it to be. I have not as yet arrived at any definite
conclusions.

This variety was very frequent in Lost Lake which is a strongly dis-
trophic Sphagnum bog.

Monroe County: Lost Lake, 2020 Turner (A-2121). Pike County: 129
Shulze (A-2219).

AMPHORA CLEVE

Amphora ovalis Kiitz., 1844, Bacill., p. 107. pl. 5, fig. 35.
Frustulia ovalis Kiitz., 1833, Linnaea 8: 539. pl. 13, fig. 5.

Pike County: Shohola Falls, 10 Burke (A-2201).
Amphora ovalis v. Pediculus (Kiitz.) Van Heurck, 1880, Syn. Diat. Belgique, pl. 1,

figs. 6, 7, text (1885) p. 59.
Cymbella pediculus Kiitz., 1844, Bacill., p. 80. pl. 5, fig. 8.

196 Fartow1A, VoL. 2, 1945

This variety, like the species, was not often found in this study. It was
rare and found in only two localities.

Monroe County: Wier Lake, 2046 Hodge (A-2168). Pike County: Lake
Wallenpaupek tributary, 5 Burke (A-2195).

Amphora perpusilla Grun., 1884, Denk. Akad. Wiss. Wien 48: 50. pl. 1, fig. 6.

This species was rare in one locality.
Wayne County: Angels, 2021 Burke and Patrick (A-2123).

CYMBELLA AcarpH

A. Central nodule with an isolated stigma or the median striae ending in an isolated
punctum at the central nodule

B. Stigma present
C. Stigma on dorsal side, valve more or less naviculoid with the ventral
margin usually swollen at the central area....... C. sinuata
C. Stigma on ventral side of central nodule, valve cymbelloid
C. tumida
B. Stigma not present, ventral striae ending in isolated puncti, in valve view
distinctly cymbelloid in shape..................2.0000- C. cistula
A. Central nodule without an isolated stigma, ventral striae at central nodule not end-
ing in isolated puncti
D. Valve view distinctly cymbelloid, raphe excentric
E. Raphe forming a straight line usually quite close to the edge of the
valve
F, Striae distinctly lineolate, ventral edge of valve straight or
somewhat convex at central nodule.......C. turgida
F. Striae not distinctly lineolate
G. Terminal nodules distant from the ends of the valves,
the fissure of the terminal nodule prolonged at a slight
angle to the general direction of the raphe
C. scotica
G. Terminal nodules near the ends of the valve, fissure not

prolonged as above................ C. ventricosa
E. Raphe somewhat curved or arched, more distant from the ventral
MOSTEI Laeieae ese ewan cava bade boa eoseaesciwls C. aspera

D. Valve more naviculoid in shape, raphe usually only slightly excentric
H. Small forms with indistinct central area
I. Striae 24 or more in 10 yu................. C. microcephala
I. Striae typically less than 20 in 10 uw, raphe more excentric
C. amphicephala
H. Larger forms with distinct central area
J. Striae less than 10 in 10 yp, valve elliptically lanceolate
C. Ehrenbergii
J. Striae more numerous
K. Ends of valve produced, apiculate. .C. cuspidata
K. Ends of valve produced, capitate to rostrate
C. naviculiformis
K. Ends only slightly produced........ C. Hauckii

Patrick: Pocono D1aToMs 197
Cymbella amphicephala Naeg. ex Kiitz., 1849, Spec. Algarum, p. 860.

This species was rare in the various habitats listed.

Monroe County: Davy’s Run, 2015 Patrick (A-2111), 2056 Patrick
(A-2180); Tobyhanna Creek, 2032 Patrick (A-2140), 2033 Patrick (A-
2142); Wier Lake, 2046 Hodge (A-2169).

Cymbella aspera (Ehr.) Herib., 1893, Diat. Auvergne, p. 69. pl. 3, fig. 10.
Cocconeis aspera Ehr., 1839, Ber. K. Akad. Wiss. Berlin 1839: 30.

Though found in two of the acid lakes, this species was very frequent
only in Mountain Lake in which the pH is 7.3.

Monroe County: Mountain Lake, 2051 Hodge (A-2178); Pocono Lake,
F. Myers (A-2213); Wier Lake, 2046 Hodge (A-2169).

Cymbella cistula (Ehr.) Kirchn., 1878, in F. Cohn’s Kryptogamen-Flora von Schles.,
2: 189.

Cocconema cistula Ehr. apud Hempr. and Ehr., 1831, Symbolae Physicae: Ani-
malia Evertebrata Exclusis Insectis — First Series, no page number, pl. 2, fig. 10.

Luzerne County: Harvey Lake, 2042 Hodge (A-2160). Monroe County:
Popomoning Lake, 2045 Hodge (A-2167); Trout Lake, 2049 Hodge
(A-2175).

Cymbella cuspidata Kiitz., 1844, Bacill., p. 79. pl. 3, fig. 40.

Though found in many habitats, this species was always rare.

Monroe County: Raymondskill Falls, 2025 Patrick (A-2131); Pocono
Lake Preserve, South Pocono Lake, 2010 Patrick (A-2101); Tobyhanna
Creek, 2032. Patrick (A-2140); Pocono Lake, 2036 Patrick (A-2149);
Wier Lake, 2046 Hodge (A-2169). Pike County: Shohola Falls, 10 Burke
(A-2201), 12 Burke (A-2205); 130 Shulze (A-2221); Bushkill Creek,
1621 Shulze (A-2227).

Cymbella Ehrenbergii Kiitz., 1844, Bacill., p. 79. pl. 6, fig. 11.

Monroe County: Davy’s Run, 2019 Patrick (A-2119); Jaggie’s Bog,
263 Shulze (A-2225).

Cymbella Hauckii Van Heurck in Hauck and Richter Phykotheka Universalis No. 147.
(Reference checked for me by Dr. David H. Linder)

Pike County: Shohola Falls, 12 Burke (A-2204), 127 Shulze (A-2215),
128-130 Shulze (A-2217-21).

Cymbella microcephala Grun. apud Van Heurck 1880, pl. 8, figs. 36-39, text (1885)
p. 63.

The identification of this species was checked with Van Heurck Type
Slide 211. In both habitats in which this species was found, the water was
approximately pH 7.

Luzerne County: Harvey Lake, 2042 Hodge (A-2160). Monroe County:
Trout Lake, 2049 Hodge (A-2174).

198 FARLOWIA, VoL. 2, 1945
Cymbella naviculiformis Auersw., 1861, in Rabenhorst Algen Europea, no. 1065.

This species though found in several habitats was always rare.

Monroe County: Davy’s Run, 2014 Patrick (A-2110). Pike County:
Shohola Falls, 12 Burke (A-2205), 127 Shulze (A-2215). Wayne County:
Angels, 2021 Burke (A-2124).

Cymbella scotica W. Sm., 1853, Brit. Diat., 1: 18. pl. 2, fig. 25.

This species is one of the most widely distributed of those found in this
study. Its distribution in no way seems to be correlated with the acidity or
alkalinity of the water. It was found in shallow water. It is of common
occurrence only in Lake Pocono, and very frequent in Trout Lake.

Lackawanna County: Indian Lake, 2044 Hodge (A-2165). Luzerne
County: Harvey Lake, 2042-43 Hodge (A-2160-62). Monroe County:
Davy’s Run, 2014 Patrick (A-2110), 2018 Patrick (A-2118), 2056 Patrick
(A-2180); Pocono Lake Preserve, South Pocono Lake, 2010 Patrick (A-
2101); Tobyhanna Creek, 2032-33 Patrick (A-2140-42); Pocono Lake,
2034-36 Patrick (A-2144-48); 2038 Patrick (A-2152), 2065 Patrick
(A-2182). Pike County: 2188-89 Shulze (A-2234-35); Shohola Falls, 3
Burke (A-2190), 4 Burke (A-2192), 10-12 Burke (A-2201-05), 14 Burke
(A-2208).

Cymbella sinuata Greg., 1856, Quart. Jour. Micros. Sci. 4: 4. pl. 1, fig. 17.

Monroe County: Raymondskill Falls, 2025 Patrick (A-2131). Wayne
County: Angels, 2021-22 Burke and Patrick (A-2123-25).

Cymbella tumida (Bréb.) Van Heurck, 1880, Syn. Diat. Belgique, pl. 2, fig. 10, text
(1885) p. 64.
Cocconema tumidum Bréb. ex Kiitz., 1849, Spec. Algarum, p. 60.

The identification of this species has been checked with Van Heurck
Type Slide no. 21. It was rare in the two localities given below.

Monroe County: Raymondskill Falls, 2025 Patrick (A-2131). Pike
County: Lake Wallenpaupek tributary, 5 Burke (A-2195).

Cymbella turgida Greg., 1856, Quart. Jour. Micros. Sci. 4: 5. pl. 1, fig. 18.

This species was rare in the various localities.
Monroe County: Trout Lake, 2049 Hodge (A-2174). Pike County:
2188-90 Shulze (A-2233, 2236).

Cymbella ventricosa Ag., 1830, Consp. Crit. Diat., part 1, p. 9.

This species, though found in many different habitats in this study, was
only twice of very frequent occurrence, in Mountain Lake and in Shohola
Falls.

Luzerne County: Harvey Lake, 2042 Hodge (A-2160). Monroe County:
Davy’s Run, 2014 Patrick (A-2110), 2016 Patrick (A-2113), 2018 Patrick
(A-2117), 2056 Patrick (A-2180); Mountain Lake, 2051 Hodge (A-2178) ;
Pocono Lake Preserve, Tobyhanna Creek, 2033 Patrick (A-2142); Mud

PATRICK: PocoNo DIATOMS 199

Run, 2031 Hodge (A-2138); Pocono Lake, 2034—36 Patrick (A-2144—48),
2065 Patrick (A-2182); Popomoning Lake, 2045 Hodge (A-2166); Ray-
mondskill Falls, 2025 Patrick (A-2021); Trout Lake, 2049 Hodge (A-
2174); Wier Lake, 2046-47 Hodge (A-2168-70). Pike County: 127
Shulze (A-2215), 130 Shulze (A-2221), 2190 Shulze (A-2236); Bushkill
Creek, 1621 Shulze (A-2227); Shohola Falls, 3-4 Burke (A-2190-92), 10
Burke (A-2200), 11 Burke (A-2202), 14 Burke (A-2209), 15 Burke (A-
2211), 2190 Shulze (A-2236), 127 Shulze (A-2215), 130 Shulze (A-2221);
Bushkill Creek, 1621 Shulze (A-2227).

GOMPHONEMA AcArDH

All of the species of this genus which were found in this study belong
to that section of the genus which is characterized by having a punctum
ending one of the median striae or present in the central area.

A. Valve distinctly asymmetrical to the transverse axis of the valve, apices distinctly
different
B. Striae at center of valve alternately longer and shorter, thus producing a
Taciate venecke amet sieepeiws se suck oa a fsa anda tus teapot G. constrictum

B. Striae not alternately longer and shorter at center of valve
C. Valve broadest at apex of valve

D. Valve distinctly constricted just below apex
G. acuminatum

D. Valve gradually tapering to opposite apex G. augur
C. Valve broadest below apex of valve
E. Apex wedge-shaped, somewhat apiculate . G. turris
E. Apex distinctly capitate................ G. sphaerophorum
A. Valve, though asymmetrical to the transverse axis, with apices not so distinctly dif-
ferent as above
F. Apices of valve acute, the valve narrow and somewhat delicate in appearance

G. gracile
Es Apicessof valve obtusesss.% ss. secrete cee oe G. angustatum var.
F. Apices of valve capitate, valve not over 30 uw long..... G. parvulum

Gomphonema acuminatum Ebr., 1832, Abhandl. K. Wiss. Berlin 1831: 88.

This species was rare in all localities cited.

Luzerne County: Harvey Lake, 2042 Hodge (A-2161). Monroe County:
Popomoning Lake, 2045 Hodge (A-2166); Wier Lake, 2046 Hodge (A-
2168). Pike County: 127 Shulze (A-2215); Bushkill Creek, 1623 Shulze
(A-2231); Lake Wallenpaupek tributary, 5 Burke (A-2195).

Gomphonema acuminatum v. coronatum (Ehr.) Rabh., 1864, Flora Europaea
Algarum, p. 290.
Gomphonema coronatum Ehrenberg, 1840, Ber. K. Akad. Wiss. Berlin 1840: 211.

This species was only found in one locality and there it was rare.
Pike County: Bushkill Creek, 1621-23 Shulze (A-2227-31).

200 FarLowl!A, Vou. 2, 1945

Gomphonema acuminatum v. pusillum Grun. apud Van Heurck, 1880, Syn. Diat.
Belgique, pl. 23, fig. 19.

Monroe County: Popomoning Lake, 2045 Hodge (A-2167); Mountain
Lake, 2051 Hodge (A-2178). Pike County: 130 Shulze (A-2221); Shohola
Falls, 14 Burke (A-2209). Wayne County: Angels, 2021 Burke and Pat-
rick (A-2123).

Gomphonema acuminatum v. laticeps (Ehr.) Grun. apud Van Heurck, 1880, Syn.
Diat. Belgique, pl. 23, fig. 17.
Gomphonema laticeps Ehr., 1843, Abhandl. K. Akad. Wiss. Berlin 1841: 416.

This species was found in one habitat and there it was rare. It may well
be a contamination from some other locality.
Monroe County: Mountain Lake, 2051 Hodge (A-2178).

Gomphonema angustatum v. obtusatum (Kiitz.) Grun. apud Van Heurck, 1880
Syn. Diat. Belgique, pl. 24, figs. 43-45.

Sphenella obtusata Kiitz., 1844, Bacill., p. 83. pl. 9, fig. 1.
Pike County: Lake Wallenpaupek tributary, 5 Burke (A-2195); Shohola
Falls, 10 Burke (A-2201). Wayne County: Angels, 2021 Burke and Pat-
rick (A-2123).

Gomphonema augur Ehr., 1840, Ber. K. Akad. Wiss. Berlin 1840: 211,

’

This species was very frequent in the surface vegetation of Mountain
Lake. The pH was 7.3.

Monroe County: Mountain Lake, 2051 Hodge (A-2178). Pike County:
Shohola Falls, 12 Burke (A-2204).

Gomphonema constrictum Ehr., 1832, Abhandl. K. Akad. Wiss. Berlin 1830: 63.

Luzerne County: Harvey Lake, 2042 Hodge (A-2160). Monroe County:
Mountain Lake, 2051 Hodge (A-2178); Popomoning Lake, 2045 Hodge
(A-2166).

Gomphonema constrictum v. capitatum (Ehr.) Van Heurck, 1880, Syn. Diat.
Belgique, pl. 23, fig. 7, text (1885) p. 123.

Gomphonema capitatum Ehr., 1838, Die Infusionsthierchen als volkommene Or-
ganismen, p. 217. pl. 18, fig. 2.

Monroe County: Davy’s Run, 2010 Patrick (A-2102), 2015 Patrick
(A-2111), 2056 Patrick (A-2180); Mountain Lake, 2051 Hodge (A-2178):
Pocono Lake Preserve, Pocono Lake, 2065 Patrick (A-2182); Tobyhanna
Creek, 2032-33 Patrick (A-2140-42); Popomoning Lake, 2045 Hodge
(A-2166); Trout Lake, 2049 Hodge (A-2174); Wier Lake, 2046 Hodge
(A-2169). Pike County: 2188-90 Shulze (A-2233-36): Bushkill Creek,
1621 Shulze (A-2227); Shohola Falls, 3-4 Burke (A-2190-92), 10-12
Burke (A-2200-04).

Gomphonema gracile Ehr., 1838, Die Infusionsthierchen als volkommene Organis-
men, p. 217. pl. 18, fig. 3.

Though this species was found in several habitats, it was of frequent

PATRICK: Pocono DIATOMS 201

occurrence only in Bushkill and Tobyhanna Creeks. My specimens cor-
respond to those illustrated by Van Heurck in the Synopsis Diatomées
Belgique, but they vary from the specimens on Cl. and Moll. Type Slide
no. 274 in that the central areas of my specimens are more distinctly
unilateral.

Northampton County: Green Pond, 2050 Hodge (A-2176). Monroe
County: Tobyhanna Creek, 2033 Patrick (A-2142); Wier Lake, 2046-47
Hodge (A-2169-70). Pike County: Bushkill Creek, 1621-23 Shulze
(A-2227-31); Shohola Falls, 14 Burke (A-2209).

Gomphonema gracile v. auritum (A. Braun) Grun. apud Van Heurck, 1880, Syn.
Diat. Belgique, pl. 24, fig. 15, text (1885) p. 125.

Gomphonema auritum A. Braun ex Kiitz., 1849, Spec. Algarum, p. 68.

I have checked these identifications against Van Heurck’s Type Slide
no. 212 and find that my specimens agree with this variety except in the
structure of the central area. In my specimens the central area is more as
v. lanceolata, but in other respects they seem to be v. auritum. No doubt
they are an intermediate form. I refer to the specimens from Tobyhanna
Creek. In the specimens from Bushkill Creek the striae are 9 in 10 p.
Those on Van Heurck’s slide have 14 striae in 10 ». However, Cleve (K.
Sy. Vet, Akad. Hand]. 26 (2): 182) states that this variation is possible.

Monroe County: Tobyhanna Creek, 2032 Patrick (A-2140). Pike
County: Bushkill Creek, 1623 Shulze (A-2231).

Gomphonema gracile v. naviculoides (W. Sm.) Grun. apud Van Heurck, 1880, Syn.
Diat. Belgique, pl. 24, fig. 13.

Gomphonema naviculoides W. Sm., 1856, Brit. Diat., 2: 98.

This variety was very frequent once and then in very shallow water on
the surface of leaves in a puddle at Angels near Mill Creek. The tempera-
ture of the water was between 50-60° F.

Pike County: Shohola Falls, 10 Burke (A-2201). Wayne County:
Angels, 2024 Burke and Patrick (A-2129).

Gomphonema parvulum Kiitz., 1849, Spec. Algarum, p. 65.
Sphenella ? parvula Kiitz., 1844, Bacill., p. 83. pl. 30, fig. 63.

Hustedt states that this species prefers standing water. In this study it
seemed to reach its best development and to be of very frequent occurrence
in shallow, cool, swift, flowing water of Davy’s Run and Mill Creek.

Monroe County: Davy’s Run, 2015-16 Patrick (A-2111-13), 2019 Pat-
rick (A-2119), 2056 Patrick (A-2180); Pocono Lake Preserve, Pocono
Lake, Mud Run, 2031 Hodge (A-2138); Pocono Lake, 2036 Patrick (A-
2148), 2065 Patrick (A-2182); Tobyhanna Creek, 2033 Patrick (A-2142) ;
Trout Lake, 2049 Hodge (A-2175); Wier Lake, 2047 Hodge (A-2170).
Pike County: 127 Skulze (A-2215); Bushkill Creek, 1623 Shulze (A-2231) ;
Lake Wallenpaupek tributary, 5 Burke (A-2195); Shohola Falls, 3 Burke
(A-2190), 10-11 Burke (A-2201-02). Wayne County: Angels, 2021 Burke
and Patrick (A-2123), 2026 Burke and Patrick (A-2134).

202 FaRLowlA, Vor. 2, 1945

Gomphonema parvulum v. micropus (Kiitz.) Cl., 1894, K. Sv. Vet. Akad. Hand.
26 (2): 180.
Gomphonema micropus Kiitz., 1844, Bacill., p. 84. pl. 8, fig. 12.

This variety was very frequent in only one locality, in Wayne County.
The habitat was in very shallow water on the surface of leaves. The tem-
perature of the water was between 50—60° F.

Monroe County: Raymondskill Falls, 2025 Patrick (A-2131). Wayne
County: Angels, 2022-24 Burke and Patrick (A-2125-29).

Gomphonema sphaerophorum Ehr., 1845, Ber. K. Akad. Wiss. Berlin 1845: 78.

This species was only found once and then it was rare.
Monroe County: Raymondskill Falls, 2025 Patrick (A-2131).

Gomphonema turris Ehr., 1843, Abhandl. K. Akad. Wiss. Berlin, 1841: 416.

Monroe County: Mountain Lake, 2051 Hodge (A-2178). Pike County:
Shohola Falls, 10 Burke (A-2201).

CYSTOPLEURA Bresisson

Cystopleura has to be used instead of Epithemia for the latter is pre-
occupied by a genus of flowering plants. Species of this genus were rarely
found in this study and those noted below were rare in each of the
localities.

Cystopleura turgida (Ehr.) Kuntze, 1891, Revis. Gen, Plant., 2: 891.
Navicula turgida Ehr., 1832, Abhandl. K. Akad. Wiss. Berlin, 1830: 64.

Monroe County: Mountain Lake, 2051 Hodge (A-2178).

Cystopleura Zebra v. proboscidea (Kiitz.) De Toni, 1892, Sylloge Algarum (Bacill.)
2: 784.

Epithemia proboscidea Kiitz., 1844, Bacill., p. 35. pl. 15, fig. 13.

This identification has been checked with Van Heurck’s Type Slide no.
254. My specimens are like the specimens on this slide except the ventral
margin of the valve is straight instead of being slightly arched in the center.

Monroe County: Mountain Lake, 2051 Hodge (A-2178).

Cystopleura Zebra v. saxonica (Kiitz.) De Toni, 1892, Sylloge Algarum (Bacill.)
2: 784.

Epithemia saxonica Kiitz., 1844, Bacill., p. 35. pl. 5, fig. 15.
Monroe County: Popomoning Lake, 2045 Hodge (A-2166).

RHOPALODIA O. MULLER

Rhopalodia gibba (Ehr.) Miill., 1897, Engler’s Bot. Jahrb. 22: 65. pl. 1, figs. 15-17.
Navicula gibba Ehr., 1832, Abhandl. K. Akad. Wiss. Berlin 1831: 80.

This species, though found in several localities, was always rare.
Monroe County: Mountain Lake, 2051 Hodge (A-2178); Pocono Lake

PATRICK: PocoNo DIATOMS 203

Preserve, Pocono Lake, 2035 Patrick (A-2146). Northampton County:
Green Pond, 2050 Hodge (A-2176). ‘

HANTZSCHIA Grunow

Hantzschia amphioxys (Ehr.) Grun. apud Cl. and Grun., 1880, K. Sv. Vet. Akad.
Handl. 17 (2): 103.

Eunotia amphioxys Ehr., 1843, Abhandl. K. Akad. Wiss. Berlin, 1841: 413. pl. 1,
fig. I, 26.

This species was only found once and then it was rare.
Pike County: 128 Shulze (A-2217).

Hantzschia elongata (Hantz.) Grun. apwd Cl. and Grun., 1880, K. Sv. Vet. Akad.
Handl. 17: 104.

Nitzschia vivax v. elongata Hantz., 1860, Hedwigia 2(6): 35-36. pl. 6, fig. 5.

In the text the citation is as given above. In the explanation of the plate
it is cited as Nitzschia elongata.

This species was very frequent in Jaggie’s Bog.

Monroe County: Jaggie’s Bog, 263 Skulze (A-2225). Pike County: 128
Shulze (A-2217); Shohola Falls, 12 Burke (A-2204).

NITZSCHIA Hassati

A. Frustule with middle part of valve spindle-shaped, ends long attenuate
Nitzschiellae
A. Frustule not so formed

B. Keel punctae prolonged into ribs extending over the surface of the valve

Grunowiae
B. Keel punctae not forming ribs, valve usually not constricted in the middle,
valve usually lanceolate ............. Dace lacdishs cnt. s Lanceolatae

GRUNOWIA4E (Rabenhorst) Grunow

Nitzschia sinuata v. Tabellaria Grun. afud Van Heurck, 1881, Syn. Diat. Belgique,
pl. 60, figs. 12, 13, text (1885) p. 176.

Denticula Tabellaria Grun., 1862, Verh. Zool. Bot. Ges. Wien 12: 548. pl. 18 (Gru-
now’s plate 12) fig. 206.

I have checked this determination with Grunow’s determination on Van
Heurck’s Type Slide no. 386.

This variety was frequent in a collection which was made by squeezing
out plants of Elodea found in Popomoning Lake.

Monroe County: Popomoning Lake, 2045 Hodge (A-2166).

LANCEOLATAE Grunow

A. Striae less than 20 in 10 yu, distinctly punctate......... N. amphibia
A. Striae more than 24 in 10 », keel punctae 7-10 in 10 u... N. Hantzschiana and vars.

A. Striae usually more than 30 in 10 yw, keel punctae 11 or more in 10 u
N. palea

204 Fartowia, VoL. 2, 1945
Nitzschia amphibia Grun., 1862, Verh. Zool. Bot. Ges. Wien 12: 574. pl. 12, fig. 23.

This species was found only once and then it was very rare.
Northampton County: Green Pond, 2050 Hodge (A-2176).

Nitzschia Hantzschiana Rabh., 1860, Hedwigia 2: 40. pl. 6, fig. 6.

Rabenhorst refers one to his Algen Sachs. resp. Mitteleurop. no. 943,
but no description is furnished. He simply compares this species to others.
This species was found only once but in that habitat it was common.

Pike County: Shohola Falls, 15 Burke (A-2211).

Nitzschia Hantzschiana f. subserians Grun. apud Van Heurck, 1881, Syn. Diat.
Belgique, pl. 69, fig. 2.
This form was found in only one locality, but it was common there.
Pike County: Shohola Falls, 15 Burke (A-2211).

Nitzschia palea (Kiitz.) W. Smith, 1856, Brit. Diat. 2: 89.
Synedra palea Kitz., 1844, Bacill., p. 63. pl. 3, fig. 27; pl. 4, fig. 2.

This species was found in several habitats, but was common in only one
locality, Green Pond, a body of water which has a higher mineral content
than the other lakes or bodies of water that were studied.

Monroe County: Raymondskill Falls, 2025 Patrick (A-2131). North-
ampton County: Green Pond, 2050 Hodge (A-2176). Pike County: Sho-
hola Falls, 3 Burke (A-2190), 10 Burke (A-2200).

Nitzschia palea v. debilis (Kiitz.) Grun. apud Cl. and Grun., 1880, K. Sv. Vet. Akad.
Handl. 17 (2): 96.
Synedra debilis Kiitz., 1865, Bacill., 2d ed. p. 65. pl. 3, fig. 45.

This variety was found only once but there it was common. It was found
associated with the species.

Northampton County: Green*Pond, 2050 Hodge (A-2176).

Nitzschia palea v. tenuirostris Grun. apud Van Heurck, 1881, Syn. Diat. Belgique
pl. 69, fig. 31, text (1885) p. 183.

This variety was found only twice and then it was rare.
Pike County: Shohola Falls, 4 Burke (A-2192), 11 Burke (A-2202).

NITZSCHIELLAE Rabenhorst
Nitzschia acicularis (Kiitz.) W. Sm., 1853, Brit. Diat. I: 43. pl. 15, fig. 122.
Synedra acicularis Kiitz., 1844, Bacill., p. 63. pl. 4, fig. 3.
The specimens found were smaller than usual, being 45 » long and 2.3 p

wide. This species was only found in one locality, and then it was rare.
Pike County: Shohola Falls, 10 Burke (A-2200).

STENOPTEROBIA Brespisson

Stenopterobia intermedia (Lewis) Van Heurck, 1896, Treatise on the Diatomaceae,
p. 374.

Surirella intermedia Lewis, 1863, Proc. Acad. Nat. Sci. Phila. 15: 338. pl. 1, fig. 2.

PATRICK: Pocono D1aToms 205

This species was rare in one collection which was designated by Shulze
as Pike County.
Pike County: 127 Shulze (A-2215).

SURIRELLA Turpin

A. Valves isopolar

B. Wings clearly evident in valve view, caniculi 20 or less in 100 uw, valves linear
HATICCOLALC) sire ayers pe eee Si hs cris Sei -cis: nee a taieenaey eran te S. biseriata

B. Wings clearly evident in valve view, caniculi 30 in 100 xu, valves elliptical
S. Moelleriana

B. Wings not clearly evident in valve view
C. Caniculi 50-60 in 100 yz, ends of valve rounded...S. gracilis

C. Caniculi more than 60 in 100 g, ends of valve somewhat wedge-shaped
S. angusta

A. Valves heteropolar

D. Wings not clearly developed in valve view, apparently marginal
S. guatemalensis

D. Wings clearly developed in valve view
E. Frustule distinctly wedge-shaped in girdle view. ..S. splendida

E. Frustule linear only slightly wedge-shaped in girdle view
S. tenera and vars.

Surirella angusta Kiitz., 1844, Bacill., p. 61. fl. 30, fig. 52.

Though found in several localities it was frequent in only one which was
from squeezings of moss in Mud Run. From this study it would seem that
this species preferred running water.

Monroe County: Davy’s Run, 2015 Patrick (A-2111), 2017 Patrick
(A-2115), 2056 Patrick (A-2180); Pocono Lake Preserve, Mud Run, 2031
Hodge (A-2138); Tobyhanna Creek, 2033 Patrick (A-2142); Raymonds-
kill Falls, 2025 Patrick (A-2131). Pike County: Shohola Falls, 10-11
Burke (A-2201-03), 14 Burke (A-2210). Wayne County: Angels, 2022
Burke and Patrick (A-2125), 2026 Burke and Patrick (A-2134).
Surirella biseriata Bréb., 1836, Mem. Soc. Acad. Sci. Artes Belles Lettres de Falaise,

1835: 53. pl. 7.

This species was found only once and then it was rare.
Monroe County: Raymondskill Falls, 2025 Patrick (A-2131).

Surirella gracilis (W. Sm.) Grun., 1862, Verh. Zool. Bot. Ges. Wien 12: 450. pl. 7
(Grunow’s plate 10) fig. 11.

Tryblionella gracilis Smith, 1856, Brit. Diat., 2: 35. pl. 10, fig. 75. (Grunow ques-
tions that this is the same.)

This species was found only once and then it was rare.
Monroe County: Raymondskill Falls, 2025 Patrick (A-2131).

Surirella guatemalensis Ehr., 1854, Mikrogeologie, pl. 33, fig. VI, 7.

This species has previously been reported from Massachusetts and Con-
necticut. It was only found once or twice in the collection. However, since

206 FARLowIA, VoL. 2, 1945

it is such a distinct species and one not to be confused with any other, I
have decided to include it.
Monroe County: Raymondskill Falls, 2025 Patrick (A-2131).

Surirella Moelleriana Grun., 1870, Catalogue and “400” Type Plate of Moeller,
2-3-10.

As indicated above this species is recorded as a new species in the cata-
logue which accompanied Moeller’s “400” Type Plate. It is stated that
these identifications are according to the classification of Grunow. It has
been generally assumed that the identifications were made by Grunow.

In some ways this species seems to be very similar to Surirella Amphioxys
W. Smith, but until William Smith’s slide is seen, this problem cannot be
settled. Smith does not give a figure and his description is not complete
enough to be definite.

This species was only found in one locality, but then it was of frequent
occurrence,

Monroe County: Raymondskill Falls, 2025 Patrick (A-2131-33).

Surirella splendida Ehr., 1854, Mikrogeologie, pl. 2, fig. IIT, 16, fig. XIV, 35, pl. 35,
figs. A-1, F-6.

This species was found only once and then it was rare.
Monroe County: Davy’s Run, 2056 Patrick (A-2180).

Surirella tenera Greg., 1856, Quart. Jour. Micros. Sci. 4: 11. pl. 1, fig. 38.

This species was found only once and then it was rare.
Monroe County: Davy’s Run, 2019 Patrick (A-2119).

Surirella tenera v. nervosa A. Schmidt, 1875, Atlas Diatomaceen-Kunde, pl. 23,
figs. 15-17.
This species was rare in the two localities cited below.
Monroe County: Raymondskill Falls, 2025 Patrick (A-2131). Pike
County: Lake Wallenpaupek tributary, 5 Burke (A-2195).

Surirella tenera v. Palmeri (Boyer) Hustedt, 1927, in A. Schmidt’s Atlas der Diato-
maceen-Kunde, pl. 366, fig. 4, 5.

Surirella Palmeri Boyer, 1920, Bull. Torrey Bot. Club 47: 70. pl. 2, figs. 14, 15.

This variety was fairly frequent in the one locality where it was found.
Monroe County: Raymondskill Falls, 2025 Patrick (A-2131).

ECOLOGICAL DISCUSSION

In the consideration of the ecological relations of the species studied, I
have thought it wise to consider in detail only those species which were
very frequent or common in a given habitat. It is the purpose of such an
ecological discussion as this one to correlate the presence of certain species
to given ecological conditions. Therefore if a species or variety is not well
established in a given habitat, it cannot be typically indicative of the con-
ditions present. It is in the taxonomic consideration that the detailed

PatTRIcK: Pocono DIATOMS 207

distribution of each species is given. The pH range given in parenthesis is
that which Krieger ! has indicated for the species in question. A study of
the various habitats shows the following relationships.

Lackawanna County: Indian Lake, altitude 1960 ft., pH 5—5.2, bi-
carbonates 8.5-11.6 p.p.m., calcium 1.54 p.p.m., color U.S.G.S. scale 10,
length 0.7 kilometer. The flora was not very plentiful as to specimens or
species in the collections made from this lake.

Luzerne County: Harvey Lake, Catskill formation, altitude 1230 ft.,
pH 7, bicarbonates 21.4-32.9 p.p.m., calcium 7.64 p.p.m., color U.S.G.S.
scale 10. Length about 4 kilometers. This is the largest natural lake in
Pennsylvania. It is a deep body of water of irregular outline formed by
glacial action and is surrounded by wooded moraine and farming country.
The principal aquatics are Ceratophyllum demersum, Elodea canadensis,
Fontinalis, disticha, Megalodonta Beckii, Najas flexilis, Potamogeton per-
foliatus, Potamogeton epihydrus, Potamogeton crispus, Ranunculus tricho-
phyllus. In the collections made from the bottom vegetation in shallow,
still water, Cyclotella compta v. radiosa and Cocconeis placentula were the
dominant forms. The collections made from the vegetation in running
water at the outlet showed a more meager diatom flora.

Monroe County: Davy’s Run, Pocono formation, is a mountain torrent
arising on the Pocono Lake Preserve and emptying into Tobyhanna Creek.
Its average pH is 5.8. The principal aquatic is Fontinalis dalecartica. The
collections were made from several different habitats. In squeezings from
Hypnum and Fontinalis growing in rapid water in deep shade, Achnanthes
linearis, Eunotia Meisteri, Fragilaria virescens (pH 6.8-7.8), Gomphonema
parvulum (pH 6.8-7.8), Navicula Rotaeana, N. subatomoides and Tabel-
laria flocculosa (pH 3.5-5.6) were very frequent in occurrence. Squeezings
from Fontinalis growing in full sun in running water yielded most frequently
Achnanthes linearis, Eunotia pectinalis (pH 6-7.8), E. pectinalis v. minor,
Navicula minima v. atomoides, N. Rotaeana and Vanheurckia rhomboides
v. crassinervia (pH 4—-5.5). Though the observed average pH was 5.8, the
species of most frequent occurrence had a minimum pH range of 3.5-6.8
and a maximum pH range of 5.5-7.8.

In a spring fed ditch six inches deep which runs along the path in the
woods which leads to Davy’s Run, Vanheurckia rhomboides v. crassinervia
(pH 4—5.5) was commonly found. Sphagnum was also growing abundantly
in the ditch.

Lost Lake, Pocono formation, is a strongly distrophic Sphagnum bog.
The temperature of the water in July was between 50-60° F. The well-
developed diatom flora consisted of relatively few species. Those of very
frequent or common occurrence were Eunotia robusta (pH 4.5-5.5), E.
tautonensis, Pinnularia viridis v. sudetica, Vanheurckia rhomboides and
V. rhomboides v. crassinervia (pH 4—-5.5). Noteworthy also was the fre-
quent occurrence of Asterionella Ralfsii and the very frequent occurrence

1 Krieger, W. 1929. Beitrage zur Naturdenkmalpflege 13 (2): 233-300.

208 FARLOWIA, VOL. 2, 1945

of Eunotia Naegelii. The flora of this bog most nearly resembles that of
Lake Tamaque which is also strongly distrophic.

Mountain Lake, Catskill formation, altitude 600 ft., length about 0.5
kilometer, pH 7—7.3, bicarbonates 41.5—68.3 p.p.m., calcium 11.71 p.p.m.,
color U.S.G.S. scale 25. This is a narrow pond formed by damming Pond
Creek and is surrounded by open farm country. The principal aquatics
are Ceratophyllum demersum, Chara vulgaris, Elodea canadensis, Najas
flexilis, Lemna minor, and Potamogeton epihydrus. In the collections which
were made from the surface vegetation, a well-diversified flora was present.
Those species which were very frequent or abundant were Cymbella aspera
(pH 6.8-7.8), C. ventricosa (pH 6.8-7.8), Cocconeis placentula (pH 6.8—
7.8), Eunotia monodon v. major, Gomphonema augur, Navicula radiosa
(pH 5-7.8) and Synedra ulna v. danica.

Pocono Lake Preserve is in the Pocono formation at an altitude of about
1640 ft. This is a large area in the Pocono Plateau which has been set aside
by the state in order to preserve the native flora and fauna.

Mud Run is a small run on the preserve near Tobyhanna Creek. Squeez-
ings from the moss growing in the spillway contained mainly Tabellaria
fenestrata and T. flocculosa (pH 3.5-5.5). The pH at the time the collec-
tions were made was pH 5.2.

Pocono Lake, length about 5 kilometers, average pH 6.4, bicarbonates
15.9 p.p.m., calcium 2.37 p.p.m., color U.S.G.S. scale 40. Pocono Lake,
surrounded by wooded country, is a long pond of irregular outline formed
by damming Tobyhanna Creek. The principal aquatics are Drepanocladus
exannulatus in large patches, Eleocharis acicularis lining areas of the shore,
Fontinalis disticha, Fontinalis novae-angliae, Myriophyllum humile in
large patches, Najas flexilis, Nitella flexilis in large patches, Potamogeton
epihydrus adhering to submerged stumps and rocks, and Sphagnum subse-
cundum. Several collections were made from the shallow water in the vi-
cinity of the Emlen dock. The piles supporting the dock were also scraped.
The diatoms of very frequent or common occurrence were Cymbella scotica
(pH 4.5-5.5), Eunotia veneris (pH 4.5-5.5), Navicula radiosa vy. tenella,
Tabellaria fenestrata, and T. flecculosa (pH 3.5-5.5), Vanheurckia rhom-
boides v. crassinervia (pH 4-5.5), and V. rhomboides v. crassinervia f. ca-
pitata. These findings according to the pH ranges of the species given by
Krieger would mean a pH 3.5—5.5.

Collections were also made from that portion of the lake known as
South Pocono Lake. The principal aquatic plant is Sphagnum. The pH
varies from pH 5.2-5.9 depending on the direction of the wind. The main
diatoms associated with the Sphagnum were Eunotia robusta (pH 4.5-5.5)
and Vankeurckia rhomboides v. crassinervia (pH 4—5.5). The two diatoms
were also commonly associated with Sphagnum in Lost Lake.

In Spring Run which empties into Pocono Lake, Eunotia microcephala
and FE. veneris (pH 4.5—5.5) were commonly found. The water in this run

is very shallow.
Tobyhanna Creek is a rapid mountain stream from the damming of

PATRICK: Pocono DIATOMS 209

which Pocono Lake is formed. Below the dam the stream continues and
empties into the Lehigh River at Stoddartsville, Luzerne County. Collec-
tions were made from the shallow water near the banks a few hundred feet
below the dam. The pH of the water was 6.4. The most common species
were Tabellaria fenestrata, and T. flocculosa (pH 3.5—5.5). However a
fairly diverse flora is present. The common occurrence of Tabellaria floc-
culosa at this pH is somewhat higher than typical for the species. Perhaps
there is some other more important factor than pH for the successful
development of this species.

Popomoning Lake (Sailor’s Lake) is in Chemung formation at an altitude
of 660 ft., and is about 0.7 kilometer long, pH 8-8.1, bicarbonates 33.6—
34.2 p.p.m., calcium 8.82 p.p.m., color U.S.G.S. scale 15. This is a deep
body of water, ovoid in shape, formed by glaciation and is surrounded by
hilly wooded moraines. The principal aquatics are Elodea canadensis,
Najas flexilis, and Potamogeton foliosus. Squeezings from the aquatics,
particularly Elodea, yielded a fairly diverse flora with Eunotia veneris (pH
4.5-5.5) and Cocconeis placentula (pH 6.8-7.8) the principal species.
Noteworthy was the frequent occurrence of Nitzschia sinuata v. Tabellaria.
This was the only locality where this species was found in this study. The
finding of Eunotia veneris in this lake was very contrary to expectations
as far as pH is concerned.

Lake Tamaque is in the Pocono formation at an altitude of 1800 ft.,
length about 0.4 kilometer, pH 3.8—4.2, bicarbonates 6.1 p.p.m., calcium
1.21 p.p.m., color U.S.G.S. scale 170. This round pond, surrounded by hills
and low forest, is formed by the damming of Beaver Creek. It is strongly
distrophic. The principal aquatics are Cladopodiella fluitans, Fontinalis
disticha, Fontinalis novae-angliae, Myriophyllum humile, Sphagnum cuspt-
datum, and Utricularia vulgaris. The flora is limited to a very few species.
The most common is Vanheurckia rhomboides v. crassinervia, and of very
frequent occurrence is Pinnularia Hilseana. In the collections taken from
surface scum, Asterionella Ralfsti was also of common occurrence. Notable
was the very frequent occurrence of Eunotia Naegelii. From this study it
would appear that Asterionella Ralfsii and Eunotia Naegeli prefer strongly
distrophic water.

Trout Lake, Pocono formation, altitude 960 ft., length about 1 kilometer,
pH 6.8, bicarbonates 12.2-15.8 p.p.m., calcium 2.57 p.p.m., color U.S.G.S.
scale 10. Trout Lake is an oblong pond formed by the damming of Moun-
tain Spring Creek and is surrounded by open farming land. The principal
aquatics are Elodea canadensis, Najas flexilis, and Potamogeton epihydrus.
The diatom flora was relatively richer in Navicula than that of other lakes
studied. The flora was fairly well diversified. The most frequent species
encountered were Cymbella scotica (pH 4.5-5.5) and Navicula radiosa v.
tenella. The finding of Cymbella scotica at this pH is contrary to the find-
ings of Krieger who lists this species under the name Cymbella gracilis,
which I have previously shown to be an invalid name for this species.

Wier Lake, Catskill formation, altitude 700 ft., length about 0.2 kilo-

210 FarLow!A, VoL. 2, 1945

meter, pH 6.6—6.8, bicarbonates 17.1-19.5 p.p.m., calcium 3 p.p.m., color
U.S.G.S. scale 10. This is a roundish pond formed by damming Wier Creek
which is surrounded by fairly level farming country. The principal aquatics
are Elodea canadensis, Fontinalis antipyretica, and Fontinalis disticha.
The flora is well developed as to species and specimens. In squeezings from
the surface vegetation, Eunotia monodon v. major, Stauroneis phoeni-
centeron (pH 6.8-7.8), and Synedra ulna (pH 6.8—7.8) were of common
occurrence. In collections taken from the bottom vegetation in shallow
water, Fragilaria bicapitata, F. elliptica, Navicula minima v. atomoides,
and Odontidium hiemale v. mesodon were very frequent or common.

Northampton County: Green Pond, Martensburg shale formation, alti-
tude 400 ft., length about 0.1 kilometer, pH 8—8.1, bicarbonates 117.1-
137.3 p.p.m., calcium 35.8 p.p.m., color U.S.G.S. scale 50. This is a small
roundish body of water which is formed from old wells and springs in the
center. It is surrounded by open farming country. The principal aquatic
is Najas flexilis. As seen by the water analysis, this pond has a higher
mineral content than any of the other bodies of water which were analyzed.
As might be expected, the diatom flora has a decidedly different aspect
than that of the other bodies of water studied. The flora is less diversified
and the dominant species is Nitzschia palea and its variety, debilis.

Pike County: Bushkill Creek is located on the Catskill formation. As
Bushkill Creek runs through both Pike and Monroe Counties it is doubtful
in which county these collections were made. Mr. Shulze is dead and no
one seems to know just where he collected. The collections were made in a
spring on Bushkill Creek opposite the head of an island. A well-developed
diatom flora is present. The genus Eunotia is the dominant genus, and the
species of common or very frequent occurrence are Eunotia flexuosa, Euno-
tia lunaris (pH 4-7.8), Eunotia pectinalis (pH 6—-7.8), and Eunotia pecti-
nalis v. undulata. Thus it would seem from the diatoms that the pH was
probably between 6-—7.8.

The swamp at Greeley is on the Catskill formation. Collections from
Fontinalis in the brook running through the swamp contained a flora fairly
rich in Eunotia and Pinnularia. However, the only two species of very
frequent occurrence were Fragilaria virescens (pH 6.8-7.8) and Meridion
circulare (pH 6.8-7.8). Collections from Sphagnum were also rich in
Eunotia. The most frequent species were Eunotia septentrionalis, FE. tenella,
Navicula festiva, and Pinnularia obscura,

The collections made at Shohola Falls, which is on the Catskill forma-
tion, were from a variety of habitats offered by a pond, a brook, and a fall.
Also just below the falls is a gorge in a limestone outcrop. The flora in the
gorge is distinctly different from that found in the other habitats of this
region. In collections taken from the edge of the pond above the falls, the
following species were common or very frequent, Eunotia flexuosa, E. lunaris
(pH 4-7.8), Melosira italica (pH 6.5-7.8), Tabellaria fenestrata, and
Tabellaria flocculosa (pH 3.5-5.5). From the moss and algae in the brook

Patrick: Pocono DIATOMS 211

the following species were very frequent or common: Cymbella ventricosa
(pH 6.8-7.8), Eunotia lunaris (pH 4--7.8), Eunotia veneris (pH 4.6—-5.5),
Fragilaria crotonensis, and Synedra ulna v. danica, From the moss in swift
water at the top of the falls, the species of very frequent occurrence were
Cocconeis placentula v. lineata (pH 6.8-7.8), Eunotia pectinalis v. undu-
lata, Navicula radiosa v. tenella, and Synedra uina v. danica. In the algae
at the top of the falls, the species of very frequent occurrence were Eunotia
veneris (pH 4.5-5.5), Meridion circulare (pH 6.8-7.8), and Navicula
radiosa v. tenella. In the scrapings from dripping rocks to one side of the .
falls the species of very frequent occurrence were Eunotia lunaris (pH
4-7.8), E. pectinalis, E. pectinalis v. undulata, E, veneris (pH 4.6-5.5),
Meridion circulare (pH 6.8-7.8), Navicula minima v. atomoides, Navicula
placentula, Tabellaria fenestrata, and T. flocculosa (pH 3.5-5.5).

Below Shohola Falls there is a limestone outcrop. The diatom flora found
here is different from that of other habitats and clearly reflects the differ-
ence of its geological and ecological environment. Collections made from
the dripping rocks facing the falls contained mainly Nitzschia Hantzschiana,
and Nitzschia Hantzschiana f. subserians. Other collections made from
dripping rocks of the side wall of the gorge further down stream but above
the concrete bridge, contained as the dominant species: Eunotia lunaris
(pH 4-7.8), Melosira roeseana v. epidendron, Navicula Keeleyi, and N avi-
cula perpusilla (pH 6.8-7.8).

Wayne County: Mill Creek, altitude 1800 ft. Mill Creek is a rapid
mountain stream located in the Catskill formation, and empties into the
Paupack Creek. Enmeshed in the moss growing in the cool, shallow, swift
water in the middle of the creek, Achknanthes lanceolata (pH 6.8-7.8),
Gomphonema parvulum (pH 6.8-7.8) and Meridion circulare (pH 6.8—7.8)
were very frequent. Growing on the stones in the swift water in the middle
of the creek was Cocconeis placentula v. lineatus which was of very frequent
occurrence. In the slow running water of a shallow tributary of Mill Creek,
Achnanthes lanceolata (pH 6.8-7.8), Meridion circulare (pH 6.8-7.8),
Odontidium hiemale v. mesodon, Synedra ulna v. danica and Vanheurckia
rhomboides v. amphipleuroides were of very frequent occurrence. From the
surface of dead leaves in very shallow water, Achnanthes lanceolata (pH
6.8-7.8), Fragilaria virescens (pH 6.8-7.8), Gomphonema gracile v. navi-
culoides (pH 4.5-5.6), Gomphonema parvulum v. micropus, Meridion
circulare v. constricta and Odontidium anceps were very frequent. The
temperature of the water was between 50-60° F.

It is interesting to note that though geographically these habitats are
relatively close together, the dominant species are so variable. This is but
another indication of how dependent the luxurious growth of a diatom is
upon its immediate ecological habitat. Thus the more accurate information
we have as to the requirements of each species the more valuable the diatom
will be as an indicator of water conditions.

21z FARLOWIA, VoL. 2, 1945

As pointed out by Hustedt,* certain diatoms seem to be found typically
associated with moss. In this study Eunotia Meisteri, E. microcephala,
Fragilaria virescens, Navicula Rotaeana and Navicula subatomoides were
most often associated with Hypnum and Fontinalis. Those species which
reached their best development in association with Sphagnum were Eunotia
robusta, Vanheurckia rhomboides v. crassinervia, Navicula festiva, N. quad-
ripartita and Pinnularia obscura. ‘This study supports the conclusions of
Hustedt that they are typically aerophils as usually the moss was in a par-
» tially submerged habitat. In this group of aerophils is Fragilaria bicapitata,
but in this study it was not found typically associated with moss. Other
diatoms found in this study which seem to flourish in moss on dripping
rocks or in partially submerged habitats are Melosira roeseana v. epi-
dendron, Navicula Keeleyi and Tabellaria flocculosa.

The genus Vanheurckia is interesting as an acidophil group. Vanheurckia
rhomboides and V. rhomboides v. crassinervia, and V. rhomboides v. crassi-
nervia f. capitata are found in various habitats. V. rhomboides v. crassinervia
is one of the few diatoms which seems to thrive in strongly acid or distrophic
water. V. rhomboides v. amphipleuroides on the other hand reaches its best
development in much less acid water.

Three other diatoms which seem to reach their best development in
strongly distrophic conditions are Pinnularia Hilseana, Asterionella Ralfsii
and Eunotia Naegeli.

On the other hand Nitzschia palea and Nitzschia palea v. debilis were
only found under alkaline conditions. Likewise Nitzschia Hantzschiana v.
subserians was only found in association with a limestone outcrop. It might
appear that Melosira roeseana v. epidendron also prefers water of a high
mineral content. However, Mr. F. J. Keeley tells me that he has found
Melosira roeseana v. epidendron growing on crystalline rocks. He believes
that it is the habitat of dripping rocks in association with moss that is the
crucial point.

Certain diatoms considered in this study were most typically found grow-
ing on rocks in shallow water. Achnanthes lanceolatum, Cocconeis placen-
tula v. lineata, and Gomphonema parvulum seemed to prefer rocks in swift
water, while Navicula per pusilla reached its best development on dripping
rocks.

Interesting also was the very frequent occurrence of Synedra ulna v. danica
in the running water of brooks. This variety has most often been reported
in the flora of lakes or ponds.

It will be noted that in some cases the water was studied from a chemical
standpoint. These were the bodies of water which Myers (Proc. Acad. Nat.
Sci. Phila. 94: 251-287. 1942) studied particularly in his work on the
rotifers of this region. The chemical analyses were made to determine the
pH, the amount of bicarbonates, and calcium. The richest diatom flora as
to number of species present was between the pH ranges of 6.4—7.3. Lake

*Hustedt, F., 1934. Abhandlungen und Vortrage herausgegeben von der Bremer
Wissenschaftlichen Gesellschaft. 8/9: (1934) 362-403.

Patrick: Pocono DIATOMS 213

Tamaque with a pH of 3.8-4.2, on the other hand, had a well-developed
flora consisting of only a few species. The same tendency as to fewness
of species was also noted in Green Pond with a pH of 8-8.1.

It is interesting to compare the floras of Green Pond and Lake Popomon-
ing. Both bodies of water have a pH of 8-8.1, but the mineral content of
the water of Green Pond, as seen by the bicarbonate and calcium deter-
minations, is much greater. The diatom floras are also distinctly different.
As might be expected this would seem to indicate that the type of diatom
flora is more closely correlated to mineral content than to pH.

As the waters considered in this study are mostly acid, it is interesting to
compare them with the acid water floras of New Jersey. Though similar in
many respects, certain apparent lacks are of interest. For instance the
genus Anomoeoneis, so commonly found in New Jersey, was not found in
this study. Also certain species of Surireila described by Lewis from the
White Mountains and frequently found in New Jersey were scarcely present
in this study. Also the diatom Tzbiella punctata, which is characteristically
found in New Jersey was only found once, in Pocono Lake. This record
was not included, as I believe it might have been contamination since the
collection was made with apparatus often used in New Jersey. However, I
must add that there is no reason so far as I know why it should not be found
in the Pocono region.

Interesting also to note is the occurrence of Eunotia tautonensis which
was originally found in New England and to date has not been reported
from New Jersey.

Generally speaking the diatom flora is that of a cool temperate region.
It contains more species common to northern or mountainous regions than
to the more moderate temperate regions.

SUMMARY

In this study two hundred forty-eight nomenclatorial entities were con-
sidered and studied. Fifteen of these are new.

The names of three species and one variety were validated by supplying
an adequate description.

The ecological factors which seem to be beneficial to the development of
the various species have been considered in the ecological discussion.

As this study is mostly concerned with the study of the diatom floras in
moss and more or less acid water, the genera Eunotia, Tabellaria, Van-
heurckia and certain sections of the genus Navicula are most abundant
from a quantitative and qualitative standpoint. The genus Pinnularia
reaches its best development in collections from Pike County, particularly
in those collections made by Shulze. However, two Monroe County habi-
tats — Pocono Lake and Jaggie’s Bog — have a fairly rich Pinnularia flora.
Several genera such as Achnanthes, Cymbella, Gomphonema, Neidium and
Surirella are fairly well represented as to number of species present, but
seldom are these species of very frequent or common occurrence in the flora.

As might be expected, the genera Amphora, Diploneis and Nitzschia are

214 FartowiA, Vor. 2, 1945

not well represented. Species of the genus Nitzschia are common only in
Green Pond and in moss at the limestone outcrop below Shohola Falls,
which have a relatively high mineral content.

It is interesting to note the complete absence of the genus Anomoencis,
Tibiella, and some of the species of Surirella which are often found in acid
water floras. Likewise some of the species of the genera Cystopleura and
Rhopalodia, which are considered by some to be aerophils, are poorly
represented.

ACADEMY OF NATURAL SCIENCES
PHILADELPHIA, PENNSYLVANIA

Vy
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216

FARLowIA, VoL. 2, 1945

PLATE I
1. Eunotia tenella (Grun.) Hustedt ....................... x2000
2,3. Achnanthes Lewisiana sp. nov. ...................... X2633
4,5. Achnanthes lanceolata v. apiculata var. nov. .......... x2500
6,7. Achnanthes Oestrupii v. parvula var. nov. ............ x2617

8. Navicula exigua v. capitata 2.0.00. 0... cee cece ee ees x2100

217

PATRICK: Pocono DIATOMS

Cm,

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PLateE I

218

STO PW hh

FARLow1A, VoL. 2, 1945

PLATE II
Achnanthes Stewartii sp. nov. ......... 000 ccceceeeceees x3031
Achnanthes Stewartii sp. nov. ....... pleats eee aie eos 4 x1719
Achnanthes Stewartii sp. nov. .............0. eee eeaee x2200
Calonels Lewiell spe M00). jo cpreiic suse eee eater uedeents x2100
Navicula poconoensis sp. NOV. ...........0000 eee eee eee x2537
Navicula concava sp. NOV. 2.0... elec eee e eee x2036

Waviciis Kreleyi ep. 00. 2. cpccended. dss idevisawiains x2160

Pocono DIaToMs

PATRICK

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PuraTeE II

220

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6
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9
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FARLOWIA, VoL. 2, 1945

PLATE III
Polaris Buck 2p. ROG. ciisexvniesinisedecaneiwues x2526
3,5. Pinnularia gibba, Ehrenberg’s illustration ........... x300
. Stauroptera gibba, Ehrenberg’s illustration .............. x300
. Pinnularia gibba v. rostrata var. NOV. ........... 000. eae x1500
Pinnularia essox as on H. L. Smith’s slide ............... x800
. Pinnularia essox, Ehrenberg’s illustration ............... x300
. Pinnularia Clevei sp. nov. ...... 0.00. c eee cece ees x598

. Pinnularia essox as illustrated by Cleve ................. x500

7rd

Pocono DiaToms

PATRICK

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Prate III

2(2):223-303 FARLOWIA Jury, 1945

THE BOLETINEAE OF FLORIDA WITH NOTES ON
EXTRALIMITAL SPECIES

Il. THE BOLETACEAE (GYROPOROIDEAE)

R. SINGER

The Boletaceae are the largest family of the suborder Boletineae. Much
attention has been paid to them since Linné’s day because of their edible
qualities. In recent years the practical aspect has shifted to another prob-
lem, their physiological relationships with forest trees, and the economic
significance of mycorrhizal symbiosis in forestry. It is obvious that no
actual progress toward practical evaluation of the complicated biology of
the mycorrhiza will be made without a profound knowledge of the taxon-
omy of the Boletaceae and extensive, accurate field observations concerning
their ecological characters.

The present paper is an attempt to contribute to the knowledge of the
Boletaceae of the state of Florida. Being a continuation of the first part
treating the Strobilomycetaceae of Florida,’ this article is based upon the
same concept as the foregoing part, and the reader is referred to the in-
troduction to that paper.

BOLETACEAE R. Maire (1901)

Characters of the family: Pileus scaly, fibrillose, mealy, tomentose,
granulose, velutinous, or glabrous, often becoming tessellate-rimose, viscid
or dry, small to large, the margin sometimes projecting; hymenophore
tubulose, rarely lamellate (in Phylloporus); tubes short to long, decurrent
or adnate to depressed around the stipe, or free; pores very small or closed
when quite young to very wide and open from the beginning, sometimes
lamellately arranged (“boletinoid”), and then not so easily separated from
the context of the pileus, usually very easily separable from the context, in
one genus (Ixechinus), all the single tubes free from each other, discolorous,
or more frequently concolorous with the pores, whitish to sordid, or whitish
to yellowish, or yellow to golden yellow, more rarely orange or red, the
pores sometimes orange to red or reddish brown, at last often olivaceous
or yellowish brown, sometimes becoming dirty livid or brownish or blue to
green on pressure; spore print olivaceous to deep olivaceous, cinnamon, or
fawn color to vinaceous pink, or ochraceous, or lemon yellow; spores
usually not very richly colored under the microscope even when quite ma-
ture (except in Xanthoconium where they are bright golden), most fre-

1 Singer, R. The Boletineae of Florida with notes on extralimital species. I. Farlowia
2 (1): 97-141. 1945.

229

224 FARLow1IA, VoL. 2, 1945

quently pale melleous to melleous or brownish melleous, or pale yellowish,
even hyaline, always smooth with homogeneous walls which rarely reach
1 » in diameter, their length rarely over 20 yw, usually well below 20 y,
without germinative pore, with globose, subglobose, short-ellipsoid, fusoid-
subcylindric, fusoid-ellipsoid, ovoid-fusoid, or cylindric outline; basidia
and cystidia usually comparatively small, more rarely large: trama of the
tube walls more or less bilateral in youth, in some genera (Phyloporus,
Xerocomus) subregular, without an interlaced hymenopodium; hyphae
with or without clamp connections; stipe cylindric, attenuate or thickened
toward the base, or ventricose to bulbous, smooth, and glabrous or orna-
mented with glandulae, with furfuraceous floccons, with scabrous squa-
mulae, or with reticulate lines, solid to hollow; veil often present, and then
membranaceous, or membranaceous-floccose, or glutinous, or pulverulent
(and then mostly yellow), either fugacious or persistent as an annulus on
the stipe; habitat: on earth, more rarely on decayed wood in wooded areas,
one species (Xerocomus parasiticus) parasitic on Scleroderma, the majority
symbiotic with forest trees, forming mycorrhiza. Type genus Boletus Dill.
ex Fr.

It has already become a commonplace to state the desirability of a parti-
tion of the once all-embracing genus Boletus into smaller units. Yet the
existing schemes proposed by W. A. Murrill and E. Gilbert respectively
(the remaining classifications are either stubbornly Friesian, or they have
not been accepted by the mycologists, or they are merely variations of
Gilbert's arrangement) were satisfactory only in a very limited way. Mur-
rill’s classification was satisfactory at a time when a system had to be mainly
workable, i.e. an easy road leading to a generic name. While Murrill’s
classification was mostly original, Gilbert’s scheme is merely an emendation
and modernization of Queélet’s classification (1886-1888). Although Gil-
bert expressly detests the evolutionary approach, he is more alluring to the
phylogenetically minded modern mycologist because, at the expense of
clear-cut divisions, he reunites mostly natural groups, or, at least, seems to
have the tendency to do so. The real test came when additional anatomical
and also chemical * characters were used to check on its principles and on
its definition of genera, and when more facts on the richer boletaceous flora
of North America and other extra-European regions became available to
test its value. It cannot be denied that after careful examination of the
resulting data, most of Gilbert’s definitions and some of his genera have
become untenable; others are complex or collective, and many of the species
cited by him will have to be transferred to other groups than those desig-
nated for them by that author. As a whole, Gilbert’s classification, quite in
contrast to its author’s claim, is a locally limited or, more accurately, a
European system. When checked merely with European material, and by
using only the limited number of characters admitted by Gilbert, it does

* Since chemical characters are not available for all species of the Boletaceae, they do

not appear much in the keys of this paper. I expect them, however, gradually to take
an important place in scientific keys to genera as well as to the sections and species.

SINGER: FLORIDA BOLETINEAE II. 225

not reveal many weak points, however, and this is the reason why it has
been adopted with some modifications by most French authors, by this
writer in his “System der Agaricales,” and, in the absence of anything
better, even by the most competent American specialist, W. H. Snell
(Mycologia 33: 415. 1941).

The first step toward a more universal solution, based not only on macro-
scopical and spore characters but on all anatomical, chemical and physio-
logical characters available, was made by this writer in his essay on a group
of genera he had studied in Siberia and compared with European and
North American species: Suillus (Ixocomus ), Boletinus, Phylloporus, Gyro-
don. Gyroporus is an easily defined genus. Consequently there was left
mainly the genus-complex around Boletus sensu stricto, i.e. the genera
Xerocomus, Pulveroboletus, Boletus, Tylopilus, Leccinum, and_ besides
some genera that had more recently been described from the tropics and
subtropics: Gastroboletus, Phlebopus, Ixechinus, Boletochaete, Phaeogyro-
porus, and Xanthoconium. As for the genera Phlebopus, Phaeogyroporus,
and Xanthoconium, the author was able to secure all data necessary to
insert these genera in any new classification; the genera Gastroboletus,
Ixechinus, and Boletochaete, however, are in need of one or two descriptive
details in order to receive their place in the system. We have attempted to
reorganize the species belonging to all the above-mentioned genera in order
to make these genera more natural and, at the same time, provide a more
precise, delimitation between them enabling the student actually to deter-
mine the genus prior to determining the species. These determinations are
not easy, for in order to get down to the name, it is necessary to provide
many more data on a given specimen than was hitherto required. But,
provided the data are available, no guesswork will be necessary, and the
keys should lead smoothly to the correct genus and species. The delimita-
tion of the genera will be more fully discussed in its proper place. In the
end, no verbal justification will make a classification acceptable. The
actual test will be in the laboratories and herbaria. In order not to com-
plicate unnecessarily the determination of the genera of Florida boletes by
providing one key to all known genera, we thought it wise to add a second
key for use in Florida only. This latter key is partly artificial, and takes
into consideration only the characters of species treated as occurring in
Florida in this paper.

SURVEY OF THE SUBFAMILIES AND GENERA OF THE BOLETACEAE
A. Group of genera imperfectly known but admitted because of the presence of some
important character or correlated characters. (See B1, B2, B3)

B 1. Setae present in the hymenium. Africa and Asia. Boletochaete
B 2. Tubes separated from each other, free. Madagascar. Ixechinus

’ Singer, R. Les genres Ixocomus, Boletinus, Phylloporus, Gyrodon et Gomphidius.
Rev. de Mycol. 3: 35-53, 157-177. pl. 4. 1938.

226 FArRLow1A, VoL. 2, 1945

B3. At least part of the surface layer of the pileus and the stipe connected and
including the young hymenophore like a peridium. South-China.
Gastroboletus
A. Genera completely known anatomically and chemically, not showing any of the
characters indicated under B1, B2, or B3.

C. Clamp connections constantly present except in occasional parthenogenetic
forms, and easy to observe on at least a fair percentage of the septa * (Species
with elongate spores and large incrusted cystidia, forming mycorrhiza with
conifers, and macroscopically characterized by having boletinoid hymeno-
phores, are not included in this subfamily.) Subfam. Gyrodontoideae

D. Hymenophore depressed or subfree around the apex of the stipe, not
arcuate-decurrent.

E. Spore print yellow. 1. Gyroporus
E. Spore print olive brown. 2. Phaeogyroporus
D. Hymenophore more or less arcuate-decurrent.
F. Veil present; cystidia rather distinct. 3. Paragyrodon
F. Veil none; cystidia usually very rare and inconspicuous.
4. Gyrodon

C. Clamp connections either not constant in a genus (Boletinus) or very sparse
(not more than 1-10 clamps per 100 septa), or persistently lacking.

G. Hymenophore either boletinoid (lamellately arranged, radiately elon-
gate pores, adnato-decurrent or deeply decurrent), or covered by a
viscid-membranaceous or glutinous veil, or stipe with glandulae, or at
least pileus (not stipe except, possibly, for the veil) strongly viscid
(this latter character has to be combined with small, pale melleous,
elongate spores, large, incrusted cystidia and cylindric or subcylindric
stipe; —if not showing these additional characters, the species is not
included in this subfamily). All species exclusively connected with
coniferous trees by mycorrhiza (species forming mycorrhiza with fron-
dose trees, or not forming mycorrhiza with Gymnospermae only, are
not included in this subfamily). Subfam. Suilloideae

H. Veil none; stipe somewhat hollow; hymenophore more lamel-
lately arranged near the margin than near the stipe; pileus not
viscid. 5. Psiloboletinus

H. Veil present or absent; stipe hollow or solid; hymenophore, if
arranged lamellately at all, more so toward the stipe than at
the margin of the pileus; pileus viscid or dry; fungi not com-
bining the characters indicated for Psiloboletinus.

I. Veil present; pileus dry, or rarely becoming viscid during
its development, starting from the margin, fibrillose or
scaly or floccose, these ornamentations or coverings being
part of the cuticle proper, not the veil; stipe always with-
out glandulae; hymenophore usually very strongly boleti-
noid, never composed of fine (diameter 0.5 mm. and less)
pores; clamp connections present in many specics.

6. Boletinus

I. Veil present or absent; if there is a veil, the pileus is
never scaly or squamulose, or/and the hymenophore is
composed of fine pores; pileus always viscid; glandulae
frequently present on the stipe, and sometimes on the

“Use the mycelioid base of the stipe, or one of the cortical layers for examination.
For further technical hints see Singer, Lloydia 5: 98. 1942.

SINGER: FLORIDA BOLETINEAE II. i

pores; hymenophore boletinoid or not boletinoid; clamp-
connections none. 7. Suillus

G. Hymenophore not boletinoid; veil, if present at all, either glutinous
or pulverulent and then yellow (yellow-green, reddish-yellow) and
dry; stipe without glandulae; pileus viscid or not; species forming
mycorrhiza with conifers or frondose trees; or possibly not forming
mycorrhiza.

K. Lateral stratum of the hymenophoral trama of the Phylloporus-
type (its hyphae only slightly divergent, rather close, and only
little paler, if at all so, than the mediostratum, often becoming
irregular in age); hymenophore made up of lamellae, or of
tubes, the tubes usually wide (about or above 1 mm. in diam-
eter) ; spore print olive brown, never pinkish, nor ferruginous-
ochraceous, nor avellaneous-ochraceous, nor cinnamon-isabel-

line. Subfam. Xerocomoideae
L. Hymenophore lamellate. 8. Phylloporus
L. Hymenophore tubulose. 9. Xerocomus

K. Trama truly bilateral-divergent (Boletus-type) when young,
(lateral stratum loosely arranged, strongly divergent, much
paler than the mediostratum, usually hyaline, but its hyphae
becoming stretched from rapid longitudinal growth of the tubes
shortly before full maturity, and then becoming parallel with
the mediostratum) ; hymenophore always made up of tubes
with either wide or small pores; spore print not always olive
brown. Subfam. Boletoideae

M. Stipe mot combining the following characters: tapering
from the base or more rarely from the middle to a
slender apex, of fibrous-hard context, with scabrous
squamulae or furfuraceous dots which usually are darker
than the remaining surface of the stipe, or becoming so
in age; pores small or wide, red or concolorous, free or
adnate; mycorrhiza with frondose trees or with conifer-
ous trees, or possibly no mycorrhiza formed.

N. Spore print consistently brownish with an oliva-
ceous tinge.

O. Veil present or absent; if the veil is present,
it consists of a yellow (yellowish-green,
reddish-yellow) pulverulence, or it is en-
tirely glutinous; if the veil is absent, the
stipe is viscid in fresh, young specimens, or
the pileus is covered by a yellow pulveru-
lence; rarely, in avelate forms, there is no
pulverulence and the stipe is not viscid; in
this case the cuticle of the pileus is viscid;
in old or dried specimens the hymenophore
often assumes a very deep golden yellow,
or golden olive, or reddish orange color.

P. Hymenophore a thin to medium thick
layer of decurrent tubes, arcuate at
least when young; stipe frequently
swollen ventricose and sometimes
furrowed at the base where it is con-
tinued into a dense mycelial growth
which often takes the shape of a

228 FartowlaA, VoL. 2, 1945

pseudomycelium; context frequently
turning blue on exposure; veil none;
size of the carpophores often re-
markable. 10. Phlebopus

P. Hymenophore consisting of rather
long tubes, not decurrent, either ad-
nate or depressed around the stipe,
either applanate or convex beneath;
stipe usually not swollen-ventricose,
mostly more or less cylindric, or
slightly thickened downwards; pseu-
dosclerotium none; context rarely
turning blue on exposure, and, if it
does, there is a yellow pulverulent
veil present. 11. Pulveroboletus

O. Veil none; stipe not viscid; pileus not cov-
ered with yellow pulverulence; if the pileus
is viscid, or the base of the stipe covered
with abundant yellow mycelium, the cuticle
is made up of a trichodermium-pallisade or
epithelium. 12. Boletus

N. Spore print not showing a trace of an olivaceous
tinge even if it is quite fresh.”

Q. Spore print rusty-yellow; spores bright
golden under the microscope, long-cylindric,
bacilliform, and narrow; stipe smooth and
glabrous; context white, unchanging, mild.

13. Xanthoconinm

Q. Spore print deeper ferruginous, or wood
brown, or fawn color, or pinkish vinaceous,
not golden under the microscope, not ex-
actly cylindric in their majority, or rarely
so; stipe rarely smooth and glabrous as
well; context rarely white and unchanging,
and mild as well. 14. Tylo pilus

M. Stipe (though somewhat variable in shape) mostly
tapering upwards from the base or more rarely from the
middle, beset with darker (or becoming darker) furfura-
ceous or squamulose scabrosities, rather fibrous-hard in
the larger lower part but tender-fleshy in the apex, the
scabrosities consisting of dermatobasidia, dermatopseu-
doparaphyses, and dermatocystidia, forming a terminal
hymenium on a fascicle of parallel hyphae; pores very
small, never red; tubes long, especially in the middle
between the stipe and the margin of the pileus, deeply
depressed-free around the stipe; mycorrhiza almost
constantly with trees of the orders Fagales and Salicales.

15. Leccinum

* * *

° The spore print should be examined immediately after a thick layer of spores on
white paper — without the hymenophore of the specimen touching it — has been ob-
tained, and the correct equivalent in Ridgway must be noted. Old spore prints change
their color.

SINGER: FLORIDA BOLETINEAE II. 229

KEY FOR DETERMINATION OF THE GENERA OF BOLETACEAE
OCCURRING IN FLORIDA

A. Clamp-connections present (choose this alternative even if they are very scarce).

B. Hymenophore depressed around the stipe; spore print yellow.
1. Gyroporus

B. Hymenophore decurrent; spore print brownish.
C. Hymenophore lamellate. 8. Phylloporus
C. Hymenophore tubulose. 4. Gyrodon
A. Clamp-connections none.
D. Veil present.
E. Veil yellow, pulverulent. 11. Pulveroboletus
E. Veil not yellow, and not pulverulent.

F. Veil viscid in wet weather; pores not radiately arranged (i.e. not boletinoid) ;
pileus viscid when wet; stipe with glandulae, rarely without them (and then

the veil mostly rudimentary, only marginal). 7. Suillus
F. Veil dry, even in wet weather; pores radiately arranged (boletinoid) ; pileus
non-viscid; stipe without glandulae. 6. Boletinus
D. Veil none.
G. Hymenophore lamellate. 8. Phylloporus

G. Hymenophore tubulose.
H. Spore print brown with an olive tinge, or decidedly olive.

I. Cuticle of the pileus with a distinct epithelium, consisting of several layers
of spherocysts which are usually arranged in erect chains, later becoming
free, and eventually often washed off; hymenophore depressed around the
stipe, pores small, yellow; stipe more or less furfuraceous or squamulose-
scabrous. 15. Leccinum

I. Pileus without a distinct epithelium.

J. Stipe fibrous-fleshy and rather hard, gray to black squamulose-scabrous
on white to pale grayish ground; context white, changing to sordid lilac
on exposure. 15. Leccinum

J. Stipe not.scabrous and usually soft-spongy-fleshy; context white or
colored, changing or unchanging.

K. Pileus with a yellow pulverulence, or stipe with glandulae, or stipe
viscid, or hymenophore arcuate-decurrent when young with the tubes
rather short to barely medium long.

L. Glandulae present on the surface of the stipe; mycorrhiza with

Pinus. 7. Suillus

L. Glandulae none.

M. Hymenophore arcuate-decurrent in young specimens shortly be-
fore full maturity, tubes rather short to barely medium long; spores
small (5.5—-11 x 2.7-4.8 «) ; growing on pine trunks.

10. Phlebopus

M. Hymenophore arcuate-decurrent but in primordial stage; tubes
medium to long in mature specimens; spores usually larger than
indicated above; not or only accidentally growing on pine trunks.

11. Pulveroboletus

K. Pileus without yellow pulverulence in any stage; stipe neither beset
with glandulae, nor viscid in any stage; hymenophore arcuate-decurrent
but in primordial stage, tubes medium to long.

230 FartowiA, VoL. 2, 1945

N. Pores wider than 1 mm. in diameter.

O. Pileus viscid and tomentose at the same time; carpophore small
(pileus 10-20 mm.) ; spores small (6—9.3 x 2.8-4.2 uw); trama of the
Phylloporus-type. 9. Neroconus

O. Pileus rarely viscid and tomentose at the same time; carpophore
mostly larger than 20 mm. broad; spores larger than 9.3 pu, or at
least some of the spores of a print larger; trama of the Phylloporus-,
or of the Boletus-type; fungi not combining the characters indicated
above.

P. Surface of the pileus (if fresh and young) strongly reacting
with NH: and NH,OH, becoming vivid blue, to bright bluish
green; trama of the PAylloporus-type. 9. Xerocomus

P. Surface of the pileus not reacting as indicated above; trama of
the tubes of the Boletus-type.

Q. Pores vivid golden yellow, remaining so on drying; stipe not
very thick, subglabrous and smooth; context unchanging;
pileus viscid when wet. 11. Pulveroboletus

Q. Not combining the above characters. 12. Boletus

N. Pores decidedly smaller than 1 mm. (if in doubt, or if pores around
1 mm. wide, see “O’’). 12. Boletus

(but compare Pulveroboletus retipes)
H. Spore print without an olivaceous tinge (‘Isabella color,’ some kind of
vinaceous pink to wood brown, fawn color, etc., or a rusty brownish yellow).
R. Spore print rusty brownish yellow; spores under the microscope bright
and intensely golden yellow; context combining white (unchanging) color
and mild taste; stipe combining smooth and glabrous surface; pileus not

viscid, not scrobiculate. 13. Yanthoconium
R. Spore print not rusty brownish yellow, at least not when fresh; spores
under the microscope brownish melleous, vellowish melleous, pale melleous
or subhyaline to hyaline; context not combining white (unchanging) color
and mild taste unless the pileus is viscid or scrobiculate; stipe not combining

smooth and glabrous surface unless the pileus is viscid or scrobiculate.

S. Stipe with distinct or indistinct glandulae, usually cylindric and not very
thick, rarely swollen; mycorrhiza with pines; spore print “ochraceous
tawny” to “Isabella color”; hymenophore adnate-decurrent; pileus glu-
tinous. 7. Suillus

S. Stipe always completely devoid of glandulae (lens), usually cylindric and
then very thick, or thickened at some part; mycorrhiza with pines or with
other trees, never exclusively with pines; spore print not “Isabella color”
(the nearest it may occasionally come to “Isabella color” is ‘colonial
buff”) ; pileus viscid or dry. 14. Tylopilus

Subfam. Gyrodontoideae Sing. subfam. nov.
Hyphis constanter fibuligeris; sporis ellipsoideis vel globosis, brunneolis vel flavis.

Characters of the subfamily: see key (p. 226) and Latin diagnosis. The
type genus is Gyrodon Opat.

The genera Gyrodon, Paragyrodon (tribus Gyrodonteae Sing.) on one
hand, and Gyroporus (tribus Leucoporelleae (Gilbert) Sing.) on the other
hand did not seem to be easily combined into one subfamily or tribe, not-
withstanding their obvious similarities, that is not until the tropical genus

SINGER: FLORIDA BOLETINEAE II. 231

Phaeogyroporus was discovered. The latter represents a perfect connecting
link between the two former tribes Gyrodonteae and Leucoporelleae which
now become superfluous.

1. Gyroporus Quél., Enchir., p. 161. 1886, emend. Pat., Ess. tax., p. 124.
1900.
Suillus Karst., Bidr. Finl. Nat. Folk 37:v. 1882, non S, F. Gray
(1821).
Coelopus Bat., Bolets, p. 12. 1908.
Leucobolites G. Beck, Zeitschr. Pilzk. 2:146. 1923.
Leucoconius (Reichenb.) G. Beck, Zeitschr. Pilzk. 2:146. 1923.

Characters of the genus: Pileus non-viscid, glabrous to coarsely fibrous-
subsquamose; cuticle made up of repent or subascendant elongate but
sometimes rather short and thick hyphae with thin or thick walls; hymeno-
phore tubulose with concolorous, small to more often medium to large pores,
depressed around the stipe, rather long, white, pink, or pallid-stramineous;
spore print yellow; spores stramineous-subhyaline to yellow, ellipsoid,
rarely more elongate than twice as long as broad, of medium size but rather
variable in this regard; cystidia present; trama of the hymenophore-walls
truly bilateral; stipe hollow or solid, its surface glabrous or fibrous, not
reticulate; context white or whitish, unchanging, or becoming bright blue
on exposure; hyphae (except for rare parthenogenetic aberrations) con-
stantly with clamp connections; the usual inorganic reagents causing very
slight or no color reactions; said to contain no boletol. Growing on the
ground in temperate as well as in tropical regions, possibly occasionally
forming mycorrhiza but sometimes fruiting without mycorrhizal connec-
tions. The type species is Gyroporus cyanescens (Bull. ex Fr.) Quél. This
is also the type species of the genus Leucoconius Beck, and it is assumed
that the logical type species of Suillus Karst. is the same species. The genus
Leucobolites Beck is obviously framed around the characters of Gyroporus
castaneus, the first species cited, and since this belongs in Gyroporus, all
three genera are synonyms of Gyroporus. In the original sense of Quélet,
Gyroporus contained several species which are now excluded from this
genus, but Patouillard’s emendation leaves no doubt about the sense in
which Gyroporus has to be restricted. The genus Gyroporus has been al-
most unanimously accepted by mycologists at present though it usually is
incorrectly (spore print “white’’) or insufficiently (no mention of clamps)
characterized.

KEY TO THE SPECIES

A. Pileus white, whitish cream color, stramineous, greenish, clay color, pinkish or umber.
B. Context unchanging on exposure.

C. Pileus “tawny olive,” “Isabella color,” “Saccardo’s umber” when fresh,

strongly and coarsely fibrous and usually subsquamose, assuming a

characteristic green color on drying if the specimens have been old,

watersoaked and exposed for several hours before drying.
2. G. umbrinisquamosus

232 FarLowlA, VoL. 2, 1945

C. Pileus not as dusky colored as above, more white to pale stramineous,
or pinkish, less strongly fibrous to completely glabrous, not assuming
a green color on drying. 1. G. subalbellus
B. Context strongly bluing on exposure. 3. G. cyanescens
A. Pileus fulvous, fulvous-ferruginous, castaneous-fulvous, tawny, cinnamon to whitish
cinnamon, or deep red, purplish carmine, or deep purplish violet.

D. Pileus deep purplish violet. G. atroviolaceus
D. Pileus not colored as above.
E. Pileus deep red, purplish carmine. 4. G. purpurinus
E. Pileus not colored as above. 5. G. castaneus

1. Gyroporus subalbellus Murr., N. Am. FI. 9:134. 1910.

Suillus subalbellus Sacc. & Trott., Syl]. 21:252. 1912.
Gyroporus roseialbus Murr., Mycologia 30:520. 1938.

Pileus with a basic white color but sometimes flushed with “sea shell
pink,” or “apricot buff,” the disc often “cinnamon buff,” and besides fre-
quently with cinnamon or pale cinnamon dots, rarely pale yellow instead of
white, frequently partly or entirely covered with a coarse but thin ‘colonial
buff” fibrillosity reminiscent of G. cyanescens and then parts of the pileus
or the entire surface shaded with this color, the disc, if fibrillose, even
reaching ‘‘chamois,” the glabrous surface sometimes minutely tessellate,
not viscid, the margin sterile and then incurved when young, or fertile,
convex, frequently with more or less depressed center, (25)—35-100 mm.
— Hymenophore white or whitish, becoming “cream color” to “mustard
yellow” with age, about 4 mm. deep, depressed or rarely adnate to the stipe,
unchanging on injury; tubes rather long, (3)—5-8 mm. long; pores con-
colorous, rather small (2.5 per mm.) to medium wide (0.9 per mm.) ; spore
print “colonial buff” with a hue of ‘straw yellow.” — Stipe white or some-
what concolorous with the pileus, very frequently flushed with pink, “sea
shell pink,” “shell pink,” “buff pink,” “light salmon orange,” frequently
stained with ‘‘cinnamon buff” like the pileus, or with brownish, olivaceous
or umbrinous spots near the base, the apex usually more white than the rest
of the stipe, glabrous or somewhat furfuraceous to slightly tomentose-
fibrillose, smooth, equal, or with enlarged base, rather frequently swollen
and distorted, solid, becoming stuffed and finally with cavities or hollow,
(25)—33-80 x (6)—-11-30 mm. — Context white, sometimes salmoneous in
the cortex of the stipe, absolutely unchanging; odor none, or agreeable,
reminding one of cinnamon; taste mild.

Spores 7.5-13.8 x 4-5.5-(6) ym, short-cylindric-ellipsoid, ellipsoid, or
ovoid, hyaline with a slight or bright yellowish tint, with suprahilar de-
pression or applanation, smooth; basidia 28-33 x 8-11.5 pw, 4-spored; cys-
tidia versiform, mostly fusoid; trama of the tube-walls truly bilateral as in
G. castaneus; fibrillosity of the pileus and the stipe of strands of parallel
hyphae, or of irregularly interwoven hyphae with often thick, yellowish
walls, occasionally rather short members between septa, 3-18 yp. thick, the
terminal ones usually clavate, more rarely subulate and more or less acumi-

SINGER: FLORIDA BOLETINEAE II. 233

nate; all kyphae with clamp connections, or at least a large percentage of
them with clamp connections, in some rare depauperate forms all hyphae
clampless.

Chemical reactions: KOH negative (faintly sordid). —NH,OH and
NH; negative. —HNOs; negative (faintly yellowish).— FeSO, faintly
grayish-pallid, stuffing of old stipes pale green (weak reaction). — Methyl-
paramidophenol in the fibrillose form in Highlands County negative, in
the glabrous form in Alachua County distinctly and quickly positive (deep
violet). — Phenol very slowly becoming more or less deeply chocolate. —
Formol negative or almost so (rarely very pale grayish red). — Aniline
negative.

Habitat: Under oaks, and under pines in mixed and pure stands, on lime
rock, gravel and sandy soil, outside the woods and in dense high, mesophytic
and tropical hammocks, solitary, or more commonly in groups, fruiting
from May to October.

Distribution: From North Carolina to Florida and west to Mississippi.
In Florida common in all zones.

Material studied: Fla. Alachua Co., Gainesville, under oaks, August
1937, type of G. roseialbus (FLAS) ; also co-type and authentic material
(FH), F 9292 (FLAS); F 19576 (FLAS); in and near Gainesville,
R. Singer, F 2560a (FH); F 2021 (FH); F 2105 (FH); F 2147 and
2147a (FH); Sugarfoot Hammock, under oak, R. Singer, F 2570 (FH);
Newnan’s Lake, Sept. 1941, under scrub oak, G. Weber, (det. Murrill)
F 18577 (this is a forma depauperatus) (FLAS); Highlands Co., High-
lands Hammock State Park, sandy roadside under Pinus palustris, August
1942, R. Singer, F 389 (FH); Marion Co. near Altoona, October 1941,
W. A. Murrill, F 21541 (FLAS); Dade Co., Matheson Hammock, R.
Singer, F 707 and F 707 a (FH); Brickell’s Hammock, R. Singer, F 1071
(FH).® — Miss., type of G. subalbellus (N.Y.)

Murrill says that his G. roseialbus differs from G. subalbellus “in its rosy
tints, glabrous surface, and solid stipe.” I did not find these characters
correlated in any way. The type of G. subalbellus is partly fibrillose, but
I have found more distinctly fibrillose specimens with pink stipe, and per-
fectly glabrous specimens without a suggestion of pink. The solid stipe
later becomes hollow, a phenomenon that can be observed at varying stages
of development exactly as in G. castaneus. My number F 2570 has slightly
furfuraceous pileus and is consequently intermediate between the glabrous
and the fibrous form. Somebody may yet find it worth his while to erect
formae for all these variations and combinations of unimportant characters.
At present, I think we can do without them. The only character that gives
some weight to the distinction between glabrous and fibrous forms is the
reaction with methylparamidophenol; further tests will probably show that
the reaction is variable in both forms.

® There is also an indication of this species from Seminole Co., Altamonte Springs
(fide Beardslee).

234 FarLowlA, Vor. 2, 1945

Coker and Beers (Boletaceae, p. 14. 1943) mention a “form with un-
changing flesh” of G. cyvanescens, There is no doubt but that this is G.
subalbellus,

Murrell’s number F 18577 is interesting because it represents an example
of a depauperate (much smaller in all its parts), probably parthenogenetic
form without clamp connections; the spores are normal, and the basidia
4-spored as usual. It belongs to the glabrous form. Murrill’s co-types of
G. roseialbus (F 9887 and F 9288) are not cited here because they belong,
at least partly, to Xanthoconium stramineum. The latter species and G.
subalbellus are sometimes macroscopically similar but the shape of the
spores, their color both under the microscope and in print, and, as a rule,
the presence or absence of clamp connections help to identify them.

G. subalbellus, like other species of this genus, does not seem to show any
preference for specific forest trees. I have found it growing exclusively
under Pinus australis at one locality, under Pinus palustris at another, and
many times under various species of Quercus without any conifers nearby.

2. Gyroporus umbrinisquamosus Murr., Bull. Torr. Bot. Club, 66:33. 1939.

Pileus “tawny-olive,” “Isabella color,” “Saccardo’s olive,” strongly fi-
brous-tomentose-subsquamulose all over, more so on the margin, not viscid,
convex, 55-75 mm. broad. — Hymenophore whitish becoming yellowish,
deeply depressed around the stipe, tubes moderately long (about 6 mm.),
pores concolorous, small (about 2.5 per mm.), spore print as in the preced-
ing species. — Stipe concolorous with the pileus and covered with the same
felt, more so toward the base, solid, soon becoming hollow, enlarged below,
50-85 x 12-25 mm. — Context white, unchanging; odor none, taste mild.

Spores 9.5-16 x 4.5—-7.5 p, ellipsoid to rarely somewhat allantoid, with a
suprahilar applanation or depression, smooth, yellowish-hyaline; basidia
25-28 x 9-14 yu, 4-spored; cystidia 35-38 x 6.3-10 », numerous, especially
near the pores, hyaline, more or less fusoid, the thickest part in the middle
or nearer the apex which is rounded or acute; trama not studied; the
fibrous coating of the pileus and the stipe consisting of strands of parallel,
cylindric, greenish-melleous hyphae, with slightly thickened walls, the ter-
minal members cylindric, fusoid, or clavate, 7-15 ,» thick; Ayphae with
clamp connections.

Chemical reactions: Watersoaked, old material turns a characteristic
green when dried slowly, this green color disappears later in the herbarium;
reactives tested gave the same results as obtained with G. subalbellus.

Habitat: In turkey oak woods (Quercus Catesbaea) and in high ham-
mocks (with Quercus laurifolia and Q. virginiana), on sandy, humous soil,
fruiting from July until October. Solitary and rare.

Distribution: Known only from Alachua Co., Florida, U.S.A.

Material studied: Fla. Gainesville, Sept. 1938, type (FLAS) ; July 1943,
R. Singer, F 2776 (FH); West of Micanopy, Oct. 1939, W. A. Murrill
(FLAS); Newnan’s Lake, July 1943, R. Singer, F 2911 (FH).

This species is very closely related to G. subalbellus, and though the

SINGER: FLORIDA BOLETINEAE II. 235

color change of dried specimens which I have never observed in the latter
species, as well as possibly the higher upper limit of the spore measure-
ments, would seem to favor its distinction as a separate species, I have
hesitated a long time to admit it as such. One of W. A. Murrill’s collec-
tions (F 18461) looks exactly like G. umbrinisquamosus in dried condition,
and at the same time, the color of the pileus was indicated as pale yellow,
and it was determined by Murrill as G. roseialbus. One may be inclined
to consider this as a transitional form, yet it is possible that a line between
this latter form and the typical G. umbrinisquamosus will prove to be con-
stant enough to maintain the latter as an autonomous species.

3. Gyroporus cyanescens (Bull. ex Fr.) Quél., Enchir., p. 161. 1886.

Boletus cyanescens Bull. ex Fr., Syst. Mycol. 1:395. 1821.

Leccinum constrictum (Pers. ex) S. F. Gray, Nat. Arr. Brit. Pl. 1:647. 1821.
Boletus lacteus Lév., Ann. Sci. Nat. sér. III 9:124, pl. 9, fig. 1-2. 1848.
Suillus cyanescens Poir. in Lam. ex Karst., Bidr. Finl. Nat. Folk 37:1. 1882.
Gyroporus lacteus Quél., Enchir., p. 161. 1886.

Gyroporus cyanescens var. lacteus Quél., Fl. Mycol., p. 425. 1888.

Pileus stramineous-yellowish, subalutaceous, sordid or pale sulphureous,
pale straw color, greenish yellow, or nearly whitish pallid, eventually often
becoming a pale brownish umber, covered with a coarse fibrous felt, not
viscid, pulvinate, usually becoming subplane or irregular, 35-120 mm.
broad. — Hymenophore white, then yellowish, bluing on injury, depressed
around the stipe, often almost free, tubes rather long (2.5-8 mm.), pores
concolorous, irregularly polygonal or subcircular, moderately wide; spore
print between “colonial buff” and ‘“‘straw yellow.” — Stipe cremeous-sordid,
yellow, or otherwise similar to the pileus in color, paler (to white), or gray-
ish white at the apex, rarely subglabrous, mostly covered with the same
kind of felt as the pileus, smooth, solid or stuffed, then with several small
cavities, or hollow, tapering upward, more rarely subequal, 30-120 x 10-30
mm.— Context of pileus white, strongly turning cyaneous (the blue of
bachelor’s buttons) when wounded, sometimes somewhat reddish brown in
the base of the stipe and palest yellow—stramineous in the rest of the stipe,
becoming greenish blue on injury; odor none; taste mild.

Spores 8-16 x 4-8 uy, ellipsoid to ovoid, most frequently 10-11 x 5-6 up,
smooth, stramineous-subhyaline; basidia 25-40 x 8.5-12 p; cystidia 35-
52 x 8-10.5 p, cylindric-clavate, or cylindric-fusoid; trama truly bilateral;
fibrosity of the pileus consisting of (5)—9-20-(25) y thick, cylindric, stiff,
yellowish hyphae, the thin ones lemon yellow, most septa with clamp con-
nections.

Habitat: In woods and even on meadows of all kinds, most frequent on
sandy soil and on sandy loam, also on clay soil; fruiting from July until
November.

Distribution: Europe, Caucasus, North America. In Florida very rare,
only in the northern part.

Material studied: Fla. Alachua Co., Gainesville, on the ground under
pines, W. A. Murrill (FLAS). — Canada, N.B., July 1902, W. G. Farlow

236 FARLowIA, VOL. 2, 1945

(FH). —N.H., White Mts., Crystal Cascade, on ground near stump, Sept.
1891, W. G. Farlow (FH); Chocorua, Aug. 1904, W. G. Farlow (FH). —
Mass. Newton, Sept. 1892, W. G. Farlow (FH); Waverley, Aug. 1897,
W. G. Farlow (FH); Magnolia 1907, &. B. Blackford (FH).—N.Y.
Ebor, Nov., C. #. Peck, (as Boletus variegatus) (217) (NYS).— Tenn.
Great Smoky Mts., Chimney Trail, birch woods, Aug. 1939, D. H. Linder
(det. Snell) (FH); Greenbrier, A. J. Sharp (det. L. R. Hesler) (FH). —
Numerous fresh collections and specimens in European herbaria. — Cauca-
sus, Abkhazia, Saken R., under roots of Fagus orientalis Aug. 1929, R.
Singer (W); Malaya Laba R., Aug. 1935, L. NV. Vasilieva (KAZ).

This species cannot be distinguished with certainty from the two preced-
ing species unless studied in fresh condition. I have to rely on W. A. Mur-
rill’s determination of the fresh material as far as the occurrence of this
species in Florida is concerned. G. cvanescens is edible. It does not seem
regularly to form mycorrhiza, at least no preference of any forest tree is
shown, and sometimes fruiting bodies are formed far from any tree at all.
The above description has been filed from notes on European and Cauca-
sian collections of this author, except for the characterization of the trama
which resulted from New England material, and strongly disagrees with
the findings of Elrod and Blanchard who found it decidedly not bilateral.
I can only repeat that it is essential to study young stages, just when the
hymenium begins to sporulate, in order to be sure that comparable material
of the species examined is at hand. If this is done, there is not the slightest
difference in the structure of the trama of the various species of Gyroporus.

4. Gyroporus purpurinus (Snell) Sing. comb. nov.
Boletus castaneus f. purpurinus Snell, Mycologia 28(5) :465. 1936.
Prater I, ric. 1-4.

Pileus “neutral red,’ “mineral red,” almost “bordeaux,” or “garnet
brown” and “maroon,” or ‘‘deep vinaceous,” velutinous or punctate-subto-
mentose, then somewhat furfuraceous, pulvinate-convex, then plano-convex,
often with more or less depressed center, with obtuse margin, not viscid,
11-50 mm. broad. — Hymenophore white, eventually becoming ‘‘naphtha-
lene yellow,” adnate-subdepressed, becoming deeply depressed around the
stipe, convex beneath, pores small, then wide (1 to 3 per mm.), concolorous;
spore print between ‘amber yellow” and “citron yellow,” i.e. brighter vel-
low than in the G. cvanescens group. — Stipe concolorous with the pileus
and occasionally slightly inclining to brownish, apex frequently white,
entirely velutinous-subfurfuraceous, stuffed and with cavities, or plainly
hollow, equal or tapering upward, 30-40 x 2.5-8 mm.— Context pure
white, unchanging; odor none; taste mild.

Spores 8.2-11.5 x 5.5-0.8 pw; basidia 21-27x 11.5-12.2 yp, 4-spored;
cystidia at the pore mouths characteristically narrowly subulate, the apex
acuminate-obtusate or subacute, hyaline, thin-walled, numerous, 23-43 x
3.3-4.7 w; trama of the tube walls truly bilateral, the mediostratum scarcely

SINGER: FLORIDA BOLETINEAE II. . 237

hyaline; cuticle consisting of irregularly arranged, unequal hyphae, the
terminal members of which are mostly erect to form (with 1-2 of their
basal hyphae) a trichodermium-palisade, the terminal members cystidioid,
subulate, ampullaceous or fusoid, rarely vesiculose, 40-100 x 5.3-20 yp,
thin- or moderately thick-walled, filled with a purple cell-sap; a similar
covering on the stipe; Ayphae with clamp-connections, but the clamps may
be rather scattered.

Chemical reactions: NH;, NH,OH, KOH and aniline-oil on cuticle
and on context almost negative.

Habitat: Shaded places under various trees, in tropical and low ham-
mocks, and also in forest of a more northern type outside of Florida, in
general growing under the same conditions as G. castaneus, but never as far
from forest trees as the latter. Solitary to gregarious, fruiting from July
until October.

Distribution: North America from New York to Florida, and west to
Michigan. In Florida widely distributed, moderately frequent in all zones.

Material studied: Fla. Dade Co. Matheson Hammock, Sept.—Oct. 1942,
under Bursera, Ficus, Nectandra, Quercus, R. Singer, F 878, F 1174 (FH);
Oct. 1942, under Pinus caribaea, Casuarina spec., Quercus virginiana, R.
Singer, F 1174. (FH); Alachua Co., Gainesville, July 1943, under Liquz-
dambar, Ostrya, Carpinus, Prunus, Quercus spec., R. Singer, F 2623 (FH);
Planera Hammock, July 1943, R. Singer, F 2885 (FH); July 1938, W. A.
Murrill & Erdman West, F 7917 (FLAS).

G. purpurinus is, in the author’s opinion, not a form or variety of G.
castaneus but a good species. Aside from the constant and persistent dif-
ference in color, this species is of slightly smaller average size than G.
castaneus, and its surfaces have a much more pronounced tendency to
become furfuraceous than in the latter species. This is by no means a color
variant within the ordinary variation of G. castaneus since it is absent in
parts of the north and west of the United States and in Europe, and it is
also not a geographic race because it grows together with G. castaneus in
some states of this country. This most beautiful species of the Gyrodon-
toideae has first been validly described by W. H. Snell but it has also been
collected by W. A. Murrill who filed it under an unpublished varietal name.

5. Gyroporus castaneus (Bull. ex Fr.) Quél., Enchir., p. 161. 1886.
Boletus castaneus Bull. ex Fr., Syst. Mycol. 1:392. 1821.
Boletus cyanescens B fulvidus Fr., Syst. Mycol. 1:395. 1821.
Boletus fulvidus Fr. ex Fr., Epicr., p. 426. 1838.
Boletus testaceus Pers., Mycol. Eur., 2:137. 1825.
Suillus castaneus Poir. in Lam. ex Karst., Bidr. Finl. Nat. Folk 37:1. 1882.
Gyroporus castaneus var. fulvidus Quél., Enchir., p. 161. 1886.
Boletus rufocastaneus Ell. & Ev., N. Am. Fungi, 2nd ser., no. 2302. 1890, nomen
nudum,

Pileus “cinnamon rufous” to ‘‘Kaiser brown,” or “tawny” to “hazel,” or
“‘Sanford’s brown” to ‘xanthine orange,” or “pecan brown” to “Rood’s
brown,” rarely paler (‘‘cinnamon buff,” “pinkish buff,” or even “pale pink-

238 Fartowla, VoL. 2, 1945

ish buff”), usually lighter colored on the margin, pubescent-subvelutinous
to finely tomentose, rarely subglabrous, not viscid, convex, then irregularly
applanate, or with depressed center, or rarely entirely concave, (20)—30—80
mm. broad. — Hymenophore white or whitish at first, then becoming cream
color to light yellow or light lemon yellow from the spores, sometimes stain-
ing brown on injury, depressed around the stipe, or becoming so, tubes up
to 8 mm. long; pores small to medium (1-3 per mm.); spore print as in
G. purpurinus. — Stipe concolorous with the pileus, the apex usually paler,
slightly tomentose-subvelutinous, not viscid, sometimes irregularly fur-
rowed, tapering upward, more rarely subcylindric, solid, then stuffed, and
eventually hollow, the hollow stage beginning at different stages depending
on the individual, rarely stuffed even when quite old, 35-80 x 8-22 mm.
— Context white or whitish and unchanging on exposure, the cortical layer
of the stipe and sometimes under the cuticle of the stipe concolorous with
the surface, firm and fleshy, later soft and brittle; odor none; taste mild.

Spores 7-11 x 4.5—6 p, ellipsoid with suprahilar applanation, stramineous-
hyaline to stramineous, smooth, many of them with an ellipsoid to sub-
globose central oil-drop, with thin to rather thick, simple wall, non-amyloid;
basidia 27-40 x 8-11.2 yw, 4-spored; cystidia on the pore mouths subulate,
gradually tapering from the 4—6 » thick base toward the acuminate but
rarely acute apex, 20-55 y» long, thin-walled, hyaline, numerous; on the
side of the tube walls the cystidia are scattered, more or less fusoid, 38—
46x 7-11 p; cuticle of the pileus and the stipe consisting of rows of fila-
mentous hyphae which are rarely shortened and irregularly vesiculose, their
diameter 5—17.5 y, their septa with clamp connections (some clampless),
with rather thick wall, with fulvous, yellow, or hyaline cell-sap, terminal
hyphae often upright with rounded apex; trama of the hymenophore truly
bilateral with distinctly divergent, looser lateral stratum in young speci-
mens; the majority of the thin-walled hyphae with clamp connections.

Chemical reactions: KOH and NH,OH very slight or no reaction;
methylparamidophenol positive on context of very fresh material, reach-
ing ‘“‘vinaceous-purple,” in less fresh material merely staining brown;
phenol chocolate.

Habitat: In mixed forests, coniferous woods, frondose woods, open places,
low hammocks, mesophytic hammocks, and high hammocks, on lawns under
oaks, on fields and pastures, on various kinds of soil, however rare or want-
ing on strongly calcareous soil, very rarely on very decayed wood, fruiting
from May till October.

Distribution: With certainty known from Europe and North Africa,
and, what appears to be the same species, in temperate North America,
common in both dry-warm and damp-warm climates, also indicated in
Australia but unconfirmed from there; in Florida common but not reach-
ing the tropical zone.

Material studied: Fla. Alachua Co. Gainesville, det. W. A. Murrill
(FLAS); R. Singer, F 2754 (FH); F 2145 (FH); F 2145a (FH); High-
lands Co., Highlands Hammock State Park, R. Singer, F 51, F 51a, F 516

SINGER: FLORIDA BOLETINEAE II. 239

(FH). —N.H. Shelburne, W. G. Farlow (FH).— Mass.: Bedford, W. G.
Farlow (FH); Cambridge, W. G. Farlow (FH); R. Singer (FH); G. D.
Darker (FH); Jamaica Plain, G. D. Darker, 5223 (FH); 6141 (FH);
6153 (FH); Wellesley, D. H. Linder (FH); Newton, W. G. Farlow (FH);
G. D. Darker, 5364 (FH); Canton, July 1933, D. H. Linder (FH); Aug.
1933, id. (FH); Purgatory swamp, D. H. Linder & R. Singer (FH); Wake-
field, D. H. Linder & R. Singer (FH); Nantucket Co., E. V. Seeler, 108
(FH). —N.J. Newfield, 1886, Ellis, Herb. Farlow ex Herb. Ellis (B. rufo-
castaneus Ell., authentic) (FH); 1891 (FH); Ellis and Everhart, N. Am.
Fungi, l.c. (type of B. rufocastaneus) (FH).— Spain, Catalonia, Aigua-
moix Valley, Aug. 28, 1934, R. Singer (BA), and numerous collections of
fresh material from France to the Caucasus; also abundant material in
various herbaria.

EXTRALIMITAL SPECIES

Gyroporus atroviolaceus (Hoehn.) Gilbert, Bolets, p. 102. 1931.
Suillus atroviolaceus Hoehn., Sitz.-ber. Akad. Wiss. Wien 123:87; Fragm. 16
(834) :39. 1914.

This species has been described from Java. The type (FH) shows ellip-
soid hyaline-citrinous spores with a slight suprahilar depression, or applana-
tion, 8.8-9.5 x 6.3-6.7 3; basidia 26.5-31.5 x 12-14 yp, 4-spored; cystidia
fusoid, brownish, 41-75 x 7-16 p; cuticle consisting of elongate hyphae
with cylindric, clavate, or broadly fusoid terminal members, the apices of
which are frequently thickened, 25-90 x 9.5-21 », many septa with clamp
connections; they are now brown, but Hoehnel, on fresher material, found
them violet, and up to 140 x25 up.

SPECIES INCOMPLETELY KNOWN

Gyroporus Earlei Murr., Mycologia 13:60. 1921.
Boletus Earlei Overh. in Seaver & Chardon, Sci. Surv. Porto Rico 8:156. 1926.

This species, described from the West Indies, is preserved but not in a
condition that would permit any conclusions.

Gyroporus caespitosus Clel., Toadst. & Mushr. 2:186. 1935.

This species, described from South Australia, cannot be considered as
certainly belonging in this genus as long as no specimens are available.

SPECIES EXCLUDENDAE

Gyroporus rufus (Schaeff. ex) Quél. belongs in Leccinum.

Gyroporus scaber (Bull. ex Fr.) Quél. belongs in Leccinum.

Gyroporus porphyrosporus (Fr.) Quél. is Porphyrellus pseudoscaber (Secr.) Sing.
Gyroporus asprellus (Fr.) Quél. is a doubtful species, hardly a Gyroporus.

Gyroporus deflexus Murr. (Leucogyroporus deflexus Snell) is a young stage of some
bolete but most certainly not a Gyroporus.

240 FarLowlA, VoL. 2, 1945

Gyroporus praeanisatus Murr. belongs in Tylopilus.

Gyroporus pisciodorus Murr. is Tylopilus tabacinus (Peck) Sing.
Gyroporus fumosiceps Murr. is Boletus fumosiceps Murr.

Gyroporus Rhoadsiae Murr. is Tylopilus Rhoadsiae (Murr.) Sing.
Gyroporus stramineus Murr. is Yanthoconium stramineum (Murr.) Sing.

Gyroporus jamaicensis Murr. (Suillus jamaicensis Sacc. & Trott.)

This species, described from the West Indies, shows some interesting char-
acters. The spores of the type (NY) are ellipsoid, smooth, non-amyloid, about
twice as long as broad, with suprahilar depression or applanation, thin-
walled, subhyaline, with hyaline walls and faintly yellowish contents, asym-
metric, 9.5-11 x (4)—-4.8-5.2 ; basidia 20-25 x 10-12 w; there are some
hyaline cystidia with swollen base and ampullaceous apical neck, 20-30 x
11-12 p, the neck e.g. 10 x 2.5-3 w; trama of the hymenophore decidedly
truly bilateral with a melleous, denser, more parallel-subinterwoven axillar
mediostratum and a loosely arranged, strongly gelatinized hyaline lateral
stratum with distinctly divergent hyphae; all septa are clampless; cuticle of
the pileus consisting of hyphae which are little more than 1 ,» thick but
toward the surface become broadened up to 12 » in diameter, and clavate;
some clavate bodies of 6-8 » diameter were observed at the apex of the
stipe. As long as nothing is known about the variabilities of the characters
and the exact color of the spore print and the chemical reactions, I hesitate
to transfer this species to any other genus. However, it is obvious that this
is not a Gyroporus, and my guess is that it belongs in Tylopilus.

Gyroporus albisulphureus Murr., Lloydia 7:325. 1944.

I have seen only a small fragment of this species, and cannot say what
it is. If it is not an abnormally colored form of some known species, it will
probably be sought for among Pulveroboletus (if viscid) or Boletus where
it would stand out because of its milk-white pileus. The microscopical
characters which I could ascertain did not bear out the opinion of Murrill,
who links it with Gyroporus. The spores are too elongate, not right in color,
and the hyphae are without clamp connections.

Gyroporus bisporus Murr., l.c.

A small fragment of this species (FH), kindly sent me by its author, is
insufficient to dispose of it. However, it can be said with certainty that it
is not a Gyroporus.

2. Phaeogyroporus Sing., Mycologia 36:360. 1944.

Characters of the genus: see key, p. 226, and the original description, /.c.
The type species is Phaeogyroporus Braunii (Bres.) Sing, There are no
representatives of this genus known to occur in Florida. However, in view
of the taxonomic importance of this genus, and the possibility that at least
one of its species may yet be discovered in the state, we add the descriptions
of two extralimital species.

SINGER: FLORIDA BOLETINEAE II. 241

Phaeogyroporus Braunii (Bres.) Sing., Mycologia 36:360. 1944.

Boletus Brauni Bres., Bull. Soc. Myc. Fr. 6: XX XVII. 1890.
Suillus Braunii Kuntze, Rev. Gen. Plant. 3°:535. 1898.

Pileus olive becoming dirty brown, smooth, glabrous, convex, becoming
partly concave, 80-150 mm. broad. — Hymenophore yellow, turning brown,
attenuate-adnexed or emarginate-depressed around the tubes, applanate
beneath, but concave near the margin, not decurrent except in youth, tubes
8-11 mm. long, pores small (about 2 to a mm.); spore print olive brown.
— Stipe yellow turning dirty brown, subglabrous, smooth, solid, tapering
upward or downward, or thinnest in the middle, 55—63—(100, according to
Bresadola) x 20-42 mm.; mycelium dingy melleous. — Context cream color
in both pileus and stipe, moderately juicy, firm in the stipe; odor and taste
not recorded, probably not distinctive.

Spores 7-8 x 5.3-6.3 yp, mostly 8x6 p, smooth, pale brown, ellipsoid;
basidia 21-25 x 9-9.5 yw; cystidia 40-50 x 6-18.5 p, vesiculose, claviform,
or subfusoid;-tvama of the hymenophore consisting of 3.5—9 » thick, often
somewhat wavy-subinterwoven (in mature material) hyphae, and some
laticiferous vessels of about 4 p» diameter; Ayphae of the cuticle of the
pileus somewhat interwoven, with up to 7.7 » thick terminal hyphae which
are clavate, hyaline, and usually thick-walled (about 1 » thick), with nu-
merous large clamp connections.

Habitat: Solitary on the ground, fruiting during the early rains.

Distribution: Tropical West Africa, Camerouns to Liberia.

Material studied: Liberia, Ganta, April 1939, G. W. Harley (det. Singer)
40 (FH).

I have not seen the type collection. However, the few insignificant dis-
crepancies between Bresadola’s description and our own, which is entirely
drawn from Liberian material, do not seem to indicate that there is a
chance of a mistake. The spores are considerably smaller in Bresadola’s
diagnosis but young spores in our specimens are 5x4 ». The smaller size
of Bresadola’s specimens and the shorter, subdecurrent tubes clearly indi-
cate that the type was a rather young specimen, probably immature.
Therefore Bresadola did not find basidia. What he thought were basidia,
are probably cystidia according to the measurements given. Boletus suda-
nicus Har. & Pat. seems to be close to this species, possibly identical; cf.
Heim, R. Rev. de Mycol. 1:13. 1936.

Phaeogyroporus tropicus (Rick aud Rehm & Rick) Sing., Mycologia 36:360. 1944.
Boletus tropicus Rick apud Rehm & Rick, Broteria 5:223. 1905.

Pileus yellowish brown, velutinous to minutely subtomentose-squamulose,
sometimes areolate, usually smooth, semiglobose, later depressed in the
center and more or less wavy at the margin, 30-100 mm. broad. — Hy-
meno phore ochraceous, then with an olivaceous tinge, the lower surface not
or not much concave under the margin of the pileus, the rest applanate to
convex and deeply emarginate around the stipe. — Stzpe brownish yellow,

242 Fartowia, Vou. 2, 1945

smooth, glabrous, solid, cylindric, or tapering downwards or upwards,
25—40 x 8-24 mm. — Context whitish; taste and odor not recorded.

Spores (6.3)-6.8-9.5-(11) x (4.9)-5.3-6.8 y, short-ellipsoid, smooth,
light brown, without suprahilar depression; basidia 9.3 » broad, 4-spored;
cystidia not seen but probably present; cuticular tomentum consisting of
rich-melleous to golden-melleous, unequal or cylindrical, filamentous, often
waved and nodose hyphae which often look like laticiferous vessels, 2.8-12
# in diameter, scarcely intermixed with much thinner and less opaque
hyphae; most of the septa with clamp connections.

Habitat: On the ground, the mycelium forming mycorrhiza with trees.

Distribution: Brazil, south to Argentina, also in Liberia, Africa.

Material studied; Brazil 1906, J. Rick, 31 (Boletus crassus but recog-
nized as authentic material of Boletus tropicus by Rick in litt.) (FH). —
Liberia, Ganta, G. W. Harley (FH).

The above materials are identical with each other. P. tropicus differs
from P. Braunii in being somewhat smaller on an average (though large
specimens of the former may be as large or larger than small specimens of
the latter) and also in the shape of the hymenophore and the size of the
pores. P, tropicus may yet turn up in the lists of Central American or even
Floridan collectors. P. Braunii is more accurately known at the time being,
but the congenerity and the specific autonomy of both these species cannot
be doubted.

P. tropicus has been shown to form a mycelium-crust closely connected
with the roots of Citrus spp. in Brazil covering the colonies of Pseudococcus
Comstocki which attack the roots of these citrus after they have been car-
ried there by an ant (Solenopsis saevissima var. moelleri Forel); these
mycelial crusts of the Phaeogyroporus are called criptas by the Brazilian
writers. The Pseudococcus living in symbiosis with the fungus is thought to
be the immediate reason for the subsequent dying of the trees affected,
though the action of an endophytic mycorrhizal fungus weakens the plants
sufficiently prior to becoming subject to a fatal attack by the complex of
fungus and Pseudococcus. This extremely complicated and _ interesting
cooperation of symbiosis, epibiosis and parasitism has been described and
illustrated, discussed and compared with an analogous case in Africa, v7z.
the phthiriosis of the coffee tree in Cameroun, by C. R. Goncalves (Ob-
servacgoes sobre Pseudococcus Comstocki (Kuw. 1902) atacando Citrus na
Baixada Fluminense. Rodriguesia 4:179-198. fig. 1-39. 1940) and F. R.
Milanez (Observagoes sobre uma estranha doenca das laranjeiras. Rodri-
guesia 4:199-263, pl. 1-18. 1940). Since we have been able to state that
P. tropicus occurs in Liberia, it would be interesting to find out whether or
not it has anything to do with phthiriosis in Cameroun, or what precisely
its biological rdle in Africa is.

3. Paragyrodon (Sing.) Sing., Ann. Mycol. 40:25. 1942.
Gyrodon subg. Paragyrodon Sing., Rev. de Mycol. 5:7. 1940.
Characters of the genus: see key, p. 226. Type species P. sphaerosporus

SINGER: FLORIDA BOLETINEAE II. 243

(Peck) Sing. (Boletus sphaerosporus Peck). Paragyrodon is not repre-
» sented in the flora of Florida.

4. Gyrodon Opat., Comm. Hist.-Nat. Fam. Fung. Bolet., Wiegmann’s
Areh, 2(1) 25,1836.

Uloporus Quél., Enchir., p. 162. 1886 (proposed lecto-type U. livi-
dus)
Boletinellus Murr., Mycologia 1:9. 1909 (type B. merulioides )

Characters of the genus: Pileus not quite glabrous and inclining to be-
come viscid in wet weather; hymenophore of irregularly arranged (gyrose)
or boletinoid, or almost lamellate tubes or honey-combs, arcuate-decurrent;
spore print brown or olive brown; stipe not reticulate, not hollow, not
glandulose, avelate; context either changing color or unchanging on ex-
posure; spores short-ellipsoid to subglobose, more rarely subcylindric-
subreniform (but very short), smooth, brownish; cystidia not differentiated
excepting cheilocystidia which are rather inconspicuous; hyphae with
clamp connections. Habitat on wood or on the ground, and in the latter
case often forming mycorrhiza with Alnus, Fraxinus. The genus is almost
cosmopolitan. Of the six or seven species known, one occurs in Florida.

The type species of Gyrodon is G. sistotremoides (Fr.) Opat., a synonym
of Boletus sistotrema Fr., Syst. Mycol. 1:389. 1821, which would appear to
be the legal name of the plant. It has been argued (Kallenbach, Pilze Mit-
teleuropas 1(15):113. 1935) that since Opatowski leaves Boletus lividus,
the well-known European species of Gyrodon in the genus Boletus, and
does not know Gyrodon sistotremoides himself, this latter may not be
congeneric with G. lividus, and may rather be a rare deformation of some
species with other affinities. It seems indeed that Karsten’s G. sistotrema
is nothing but Suilus bovinus since he indicates the spores as elongate.
Despite the fact that Fries cites this description of Karsten’s in his Hy-
menomycetes Europaei, this does not necessarily mean that Fries’ species
is likewise a Suillus. In fact, Fries cites at the same time “B. brachyporus
Rostk.” as synonym, and this figure in Sturm’s Flora is considered to be
a genuine Gyroporus lividus (see Gilbert, Bolets, p. 216. 1931). Konrad
and Maublanc and other authors think that G. sistotrema and G. sisto-
tremoides are synonyms of G. lividus, and judging from the description
which is the only document we have, we admit that there is no good reason
for eliminating this species from Gyrodon since it may well pass as an
extreme form, G. lividus. It would thus become a form with more elongated
stipe and “whitish” context, and the configuration of the hymenophore
described by Fries would find its logical explanation when considered to be
of the type observed in old stages of Gyrodon lividus rather than by look-
ing for rare deformations in other genera. The secret of Gyrodon sistotrema
will never be lifted in an absolutely conclusive way, and, as it seems to us,
it would be best to let it go at that, and not cause nomenclatoral trouble
by refusing “endorsement” of this species as a Gyrodon sensu stricto. The

244 FARLowIA, VoL. 2, 1945

general consensus appears to be rather to model Gyrodon along the char-
acters of Gyrodon lividus even at the risk of having to conserve the generic
name Gyrodon in this sense.

KEY TO THE SPECIES

A. Spores medium to large (6)-6.8-11 x 4.7-9 w; pores wide, later becoming very wide,
boletinoid or forming a hymenophore similar to that of Phylloporus rhodoxanthus.
The American forms not combining bluing context with russet-staining stipe. Not
connected with Alnus.

B. Pileus “russet”; hymenophore nearly of the Phylloporus rhodoxanthus type
G. intermedius
B. Pileus olivaceous, sepia, or tan-clay-color, not reddish; hymenophore more
or less boletinoid.
C. Context and tubes not bluing on injury. 6. G. proximus
C. Context and tubes bluing on injury.

D. Stipe usually slightly eccentric to central; carpophores growing
neither on fern-trunks, nor in association with Fraxinus ameri-
cana. South American species. G. Rompelii

D. Stipe usually strongly eccentric or lateral; carpophores growing
either on fern trunks, or in association with Fraxinus americana.
North America and Asia.

E. Carpophores rather robust, stipe comparatively thick;
growing on very decayed wood or on earth in (probably
mycorrhizal) association with Fraxinus americana. North
American species. G. merulioides

E. Carpophores thin and small; stipe slender, growing on
fern trunks. Javanese species. G. spec.

A. Spores small: 4-8 x 3-5 ; pores rather small but elongate, later becoming irregu-
larly elongate, of varying level, and therefore appearing as if toothed; context
strongly bluing; stipe becoming russet-spotted, always associated (probably by
mycorrhiza) with Alnus nigra, A. viridis, A. incana. G. lividus

6. Gyrodon proximus Sing. spec. nov.

Pileo obscure fusco, dein biso vel umbrino, saepe pallescente usque ad alutaceum,
tomentoso in centro, subviscido, margine incurvo, dein attenuato, convexo, denique
irregulariter applanato, 20-115 mm. lato. Hymenophoro submerulioideo, dein boleti-
noideo, poris tubulis concoloribus, immutabilibus, inaequalibus, flavis, subviridulis in
vetustis ; sporis 7-9.5—(9.8) x 4.8-6.2—(6.8) qu, melleo-brunneis; hyphis fibuligeris. Stipite
pileo concolori, ut minimum ad basin, gradatim vel subabrupte hymenophoro apicem
versus concolori, solido, subglabro, sublevi, 20-63 x 9-15 mm. Carne superficiebus pal-
lidiore, plus minusve alutacea, aeri exposita immutabili; odore subtili vel nullo. Habi-
tatio: Ad terram et ad ligna in silvis humidis Americae subtropicalis tropicalisque.

Pileus initially deep brown (‘mummy brown”) becoming “bister” or
“Saccardo’s umber,” or “buckthorn brown,” in many specimens bleaching
to alutaceous (“pinkish buff,” “cinnamon buff”) or argillaceous (“clay
color”), frequently with still paler portions (‘cartridge buff”), tomentose,
especially in the center, and often shining because of a tendency toward
viscidity during heavy rains, with the margin incurved when young, later
progressively thinning out and narrowly sterile in most specimens, convex,
eventually irregularly flattened, 20-115 mm. broad. — Hymenophore “am-

SINGER: FLorIDA BOLETINEAE II. 245

ber yellow” to “wax yellow,” or “aniline yellow,” assuming a greenish tinge
- when old, deeply decurrent, tubes rather short, pores concolorous, un-
changing, submerulioid, becoming more boletinoid, 0.5-1.3 mm. wide, on
unequal level; spore print olive brown. — Stipe concolorous with the pileus,
in the lower part, or at least at the base, gradually or somewhat abruptly
paler and yellower toward the apex, usually reaching the color of the
hymenophore just below the pores, both the stipe and the spores, or either
of them exuding a chestnut colored watery liquid in some specimens,
straight, or curved, eccentric to central, more rarely lateral, solid, 20-63 x
9-15 mm. — Context paler than the surface, more or less alutaceous, not
changing on injury; odor slight, agreeable, or none.

Spores 7—-9.5—(9.8) x 4.8-6.2-(6.8) p, short-ellipsoid, smooth, honey-
brown, occasionally subreniform, with rather thin or medium thick walls,
with a short-ellipsoid central oil-drop, or without it; basidia 21-43 x 9-9.5
p, 4-spored; cystidia on the pore-mouths (cheilocystidia) fusoid, bottle-
shaped, or subulate, 15-17 x 4 m, neck, if present, 2 p in diameter; trama
of the hymenophore truly bilateral, mediostratum slightly denser than the
diverging lateral strata; Ayphae with clamp connections.

Chemical reactions: KOH on surface of pileus, deep chestnut, or brown,
usually fading to ochraceous but with a dark ring; on context brown, —
NH; on surface of pileus, lilac to deep brownish lilac. — NH,OH on sur-
face of pileus, deep chestnut or blackish to lilaceous, with a brownish-
lilaceous or lilac ring around the wetted spot; on the context and on the
tubes, blue. — HNO3; everywhere light reddish brown, reddish brown. —
H.SO, on surface of pileus, deep chestnut; on context, ochraceous brown,
or negative; on tubes orange brown. —Formol on context, greenish or
bluish. — FeSO, on tubes, green to blue-green.

Habitat: In moist lime sink hammocks and near creeks in swampy or
low hammock vegetation, also said to occur in high hammocks and oak
woods, on decaying wood and on the ground, fruiting from February
(British Guiana) and June (Florida) to September.

Distribution: Known to occur in north Florida and British Guiana,
probably also on Jamaica, apparently all over the Caribbean area.

Material studied: Fla. Alachua Co., Gainesville, R. Singer, F 2649
(FH); W. A. Murrill, F 18174 (FLAS); Devil’s Millhopper, R. Singer,
F 2425 (type), F 2425 a (FH).— British Guiana, Plantation Vryheid,
D. H. Linder, 1005 (FH)."

This species differs from G. merulioides in having unchanging flesh and
tubes, being somewhat more variable in its habit (even more often sub-
centrally stipitate than lateral), and showing somewhat different chemical
reactions. It also differs slightly in the shades and distribution of the colors
but this character is not quite reliable. In flat frondose woods, on the
ground, it usually assumes the habit of G. Rompelit (from which it differs

7Specimens from Jamaica without notes cannot be listed as more than probably
belonging here: Blue Mts., Morca’s Gap, Sept. 21-24, 1908, E. G. Britton (det. W. A.
Murrill) (as Boletinus porosus ?) (NY). .

246 FarLowIA, VoL. 2, 1945

in having unchanging context and hymenophore) whereas, on steep land-
sides, banks, vertical surfaces of logs, etc., it tends rather to assume the
general appearance of G. merulioides. The spores are definitely different
from these of G. merulioides and G. Rompelii as well. However, the meas-
urements and the indication of the shape would not clearly express the
difference. The spores of all Gyrodon are variable in their proportions
(Q=length : breadth), but in the bluing American species (G. merulioides,
G. Rompelii), the spores with Q=1.35 and less are more numerous than in
the unchanging species (G. proximus) where the spores with Q = 1.45-1.60
are very common. Besides, the bluing species of this continent have spores
which frequently, though not always, reach larger figures in length and
especially in breadth, than G. proximus.

It is not clear what trees this species prefers, if any, and whether or not
there is a mycorrhizal relationship between forest trees and the mycelium.
There was in my original notes no comment made on a possible association
with Fraxinus. The type locality has been made the subject of a second
floristic analysis,* and the species of trees recorded then were: Liguidambar
styraciflua (the decaying wood on which some of the specimens grew was
likewise identified as belonging to sweet gum), Ostrya virginiana, Car pinus
caroliniana, Quercus virginiana, also Evonymus and Toxicodendron were
present. Ulmus alata, Magnolia grandifolia, Tilia heterophylla, Fraxinus
americana, F. pauciflora were found farther away. Judging from the ex-
periences with Gyrodon merulioides in the northern states, one would be
inclined to think that Fraxinus is, here too, the mycorrhiza-tree of the
Gyrodon. But, on the other hand, Fraxinus was definitely absent at least
from one place where the Gyrodon was collected. Aside from this, Fraxinus
does not occur in British Guiana, and is also not known on Jamaica. Since
the specimens from British Guiana can hardly be anything but G, proximus,
the association with Fraxinus, if ever existent farther north, must be flexible
enough to allow replacement of this tree by some other species of the
Oleaceae, or else the occurrence of the fungus is not necessarily restricted
to the simultaneous presence of a mycorrhizal symbiont. In this regard,
G. proximus seems to differ from G. merulioides.

EXTRALIMITAL SPECIES

Gyrodon intermedius (Pat.) Sing., Rev. de Mycol. 3:172. 1938.
Phylloporus intermedius Pat., Bull. Soc. Myc. Fr. 11:86. 1895.
Suillus intermedius Kuntze, Rev. Gen. Plant. 3°:535. 1898.
Boletus intermedius Sacc., Syll. 14:165. 1899.

Boletinellus intermedius Gilbert, Bolets, p. 105. 1931.

The type (FH) from Madagascar has the spores 7.8-9.2 x 5-6.8 , and

* A floristic paper (Lillian E. Arnold. A Field Trip to the Devil's Millhopper. Jour.
E. Mitch. Sci. Soc. 52(1):77-90. pl. 7-10. 1936) dealing with this lime sink vegetation
was of great help. The author divides the sink-slopes in four zones. In the higher two
zones where my number F 2425 a (co-type) was collected, Fraxinus has not been in-
dicated.

SINGER: FLORIDA BOLETINEAE II. 247

the hyphae with clamp connections. No mention of the bluing of the con-
text is made in the description. We also have West African material
(Liberia, Firestone # 3, July 25, 1926, D. H. Linder (det. R. Singer)
(FH) that belongs to this species because of the color of the pileus and
the shape of the hymenophore, but the spores are slightly smaller (6-9.3 x
4.7-5.5 p, mostly 6.8-7.8x4.8-5 pw); basidia 26-28 x 7.5-11 »; hyphae
with clamp connections. The specimens show larger and more central stipes
like G. Rompelii, and the flesh turns blue on injury (when fresh) as in the
latter. The pileus is “russet” and the hymenophore “lemon chrome” in
fresh condition. For the time being, we do not think it wise to separate
these forms specifically.

Gyrodon Rompelii (Pat. & Rick) Sing., Rev. de Mycol. 3:172. 1938.
Phylloporus Rompelii Pat. & Rick, Broteria 6:81. 1907.

The type and authentic material (FH) has spores exactly as G. meru-
lioides, (7.5-10.5 x 6-7.5 »), and the authors indicate that the fresh context
turns blue when broken. The habit is the only visible difference between
this and G. merulioides. There may be a difference in the chemical char-
acters, and there is one in ecology, since Fraxinus americana, the constant
accompanying tree of G. merulioides, does not occur in Brazil. Yet it may
well be that, as indicated by this author in previous papers, G. Rompeli is
not specifically different from G. merulioides. In fact, the author has seen
specimens from the southern hemisphere with lateral stipe. The absence
of Fraxinus at its habitats and the fact that, between the geographic areas
of G. merulioides and G. Rompelii, a gap is noticeable with G. proximus
replacing the northern and the southern species in the tropical belt, prevent
me from reducing G. Rompelii to synonymy.

Gyrodon merulioides (Schw.) Sing., Rev. de Mycol. 3:172. 1938.

Daedalea merulioides Schwein., Trans. Amer. Phil. Soc. II 4:160. 1832.
Paxillus porosus Berk. in Lea, Cat. Plants Cincinnati, Ohio, p. 54. 1839.
Boletus lateralis Bundy in Chamberlain, Geology Wisconsin 1:398. 1883.
Boletinus porosus Peck, Bull. N.Y. State Mus. 8:79. 1889, incl. var. opacus.
Boletinellus merulioides Murr., Mycologia 1:7. 1909.

This species is so similar to G. proximus of Florida that a complete de-
scription of our collections appears to be necessary for the comparison of
these plants.

Pileus olive brown or brown (“Isabella color” and “light brownish olive”
in various combinations), often with darker portions or spots (“mummy
brown” or deeper), also darker colored where handled, circular or elongate
in circumference, dry, finely subflocculose or glabrous and like kid, or
tomentose, or partly so, convex, eventually flattened or concave, 40-110: -
mm. broad. — Hymenophore “old gold” with a flush of “Isabella color,”
bluing where injured, becoming more brown or green when old or dry,
boletinoid to submerulioid, some of the perpendicular walls not reaching
the level of the radial (lamellose) walls, decurrent and forming an incom-
plete or complete wide network of veins on the very apex of the stipe, 4-14

248 FARLowI1A, VoL. 2, 1945

mm. thick, pores irregular, angular, more radially elongate near the margin
than farther inside, more than 1 mm. wide in middle zone (seemingly much
wider than 1 mm. but the lower connecting walls inconspicuous, and there-
fore often not taken in consideration) ; spore print slightly deeper colored
than ‘Isabella color” when quite fresh, and slightly lighter than this in an
old print. — Stipe “buckthorn brown” and “yellow ocher” at apex, abruptly
concolorous with the pileus in the lower portion, and there eventually al-
most black (“citrine drab” and ‘clove brown”) in some specimens, the
very base again paler in some cases, subglabrous, almost smooth, almost
constantly either lateral or eccentric, rarely only slightly eccentric-sub-
central, curved or straight, solid, 10-34 x 8-18 mm.— Context “baryta
yellow” to “pinard yellow” in the pileus, ‘‘fawn color” to almost “army
brown” in the base, often pale brown almost throughout the stipe, becoming
dirtier here, and slowly bluing in the pileus when exposed; odor none, or
of raw potatoes, or slightly spermatic.

Spores (6.8)—7-11 x 4.8-9.2 p, mostly 7.5—-10.5 x 5.8-9 u, short-ellip-
soid, with well-colored, somewhat thickened walls, smooth: basidia about
25 x 10-10.8 », 4-spored, with 3.5-4.2 long sterigmata; cystidia rare, not
very conspicuous, cylindric, up to 6.5 » broad; trama of the hymenophore
truly bilateral; cuticle of the pileus of filamentous, 2-3.5 u thick hyphae
which are incrusted by the pigment, 3—4 ,» thick in the cuticle of the stipe,
some hyphae thickened up to 15 » in diameter, and filled with a yellowish
sap; most Ayphae with clamp connections.

Chemical reactions: KOH on the surface of the pileus, a dusky mahog-
any brown; on context, light brown to very deep brown. — NH: on surface
of pileus, “Rood’s brown.” — NH,OH on surface of pileus, very deep
brown where it had been brown, more chestnut to paler tan where it had
been more olivaceous; on context of pileus provoking the same color as
the natural color of the context of the base of the stipe, or becoming dirty
salmon color, or dirty gray; on tubes, “Mars brown” to “Prout’s brown”
(somewhat more sordid than these).— H»SO, on surface of pileus, as
with KOH; on context and tubes, subferruginous. — Phenol on context,
more blue than by mere autoxydation, eventually becoming brick red. —
FeSO, on context, olivaceous-green.

Habitat: In open woods and shaded lawns under Fraxinus americana, or
in its neighborhood, on the soil, or on decayed wood of various trees,
usually gregarious, fruiting from June to October.

Distribution: Common in most northeastern states, also in eastern Can-
ada and south to Georgia and Alabama, west to Wisconsin.

G. merulioides has been studied by the author in numerous dried and
fresh specimens. D. H. Linder (in schedulis) first brought the Fraxinus-
relation to the author’s attention, and then his own observations as well
as W. H. Snell’s independent experiences (orally to the author) confirmed
this relationship. However, in artificial conditions of the laboratory, it is
possible that this species may form mycorrhiza with other species than the
above mentioned. In fact, it has been shown recently that a fungus de-

SINGER: FLORIDA BOLETINEAE II. 249

termined as Boletinus porosus forms mycorrhiza with pines. This, however,
if it happens in the laboratory, is not likely to happen in nature.

Gyrodon spec.

One of the undetermined boletes in the Hoehnel Herbarium (FH), col-
lected near Tjibodas, Java, in 1909 on a fern-trunk, F. v. Hoehnel B.n. 1187
was examined by this writer, and found to belong in Gyrodon, more pre-
cisely in the G. merulioides-Rompelii-group, differing from both G. meru-
lioides and G. Rompelii in the slender, eccentric stipe, and the much thinner
pileus. The spores are 7.8-10.2 x 5.5-6.8 y, short-ellipsoid, brownish,
smooth. It is possible that the habitat is only incidental, and that there is
Fraxinus near the place where it was collected (Fraximus does occur on
Java). It may also be that the thin pileus and slender stipe do not normally
characterize the Javanese form. Therefore, I do not think it wise to publish
it under a new specific name but merely wish to call the attention of other
mycologists to the subject.

Gyrodon lividus (Bull. ex Fr.) Sacc., Syl]. 6:52. 1888.

Boletus lividus Bull. ex Fr., Syst. Mycol. 1:389. 1821.

Boletus sistotrema Fr., Syst. Mycol. 1:389. 1821, non Rostk. in Sturm.
Boletus brachyporus Pers., Mycol. Eur. 2:128. 1825.

Gyrodon sistotremoides Opat., Comm. Hist.-Nat. Fam. Fung. Bolet., Wiegmann’s

Arch. 2(1):5. 1836.

Boletus rubescens Trog, Flora 22°:449, 1839.

Boletus lividus ssf. alneti Lindgr. ex Fr.. Hymen. Eur, p. 519. 1874.
Boletus lividus ssp. labyrinthicus Fr., Hymen. Eur., p. 519. 1874.
Boletus lividus var. rubescens Quél., Ass. Fr. Avance. Sci., 1885, p. 7. 1885.
Uloporus lividus Quél., Enchir., p. 162. 1886.

Uloporus lividus var. rubescens Quél., Enchir., p. 162. 1886.

Gyrodon lividus ssp. alneti Sacc., Syll. 6:52. 1888.

Gyrodon lividus ssp. labyrinthicus Sacc., Syll. 6:52. 1888.

Gyrodon rubrescens Sacc., Syll. 6:52. 1888.

?Boletus gyrosus Pers., Mycol. Eur. 2:138. 1825.
?Boletus caespitosus Mass., Brit. Fung. Flor. 1:297. 1892.
?Gyrodon caespitosus Sacc., Syll. 11:81. 1895.
?Gyrodon Mikhnoi Karst. (descr. desid.) ex Gilbert, Bolets, p. 104. 1931.

This is the only species known thus far from Europe but it has not been
observed in America. For a description and plate see Kallenbach, Pilze
Mitteleuropas 1:109. pl. 10, fig. 4-7; pl. 39, fig. 1-22; pl. 44, fig. 70-72,
and for the microscopical characters R. Singer, Rev. de Mycol. 3:172. 1938.

SPECIES INCOMPLETELY KNOWN

Gyrodon purpureus (Beeli) Sing., Rev. de Mycol. 3:171. 1938.
Favolus purpureus Beeli, Bull. Soc. Roy. Bot. Belg. 58:213. 1926.
Boletinellus purpureus Gilbert, Bolets, p. 105. 1931.

This species from the Belgian Congo has been admitted in Gyrodon by
this author conditionally (“Si ce que M. Gilbert affirme relativement au
Favolus purpureus est exact...”), on the authority of Gilbert who has
seen the type.

250 FAarRLow!A, VoL. 2, 1945
Gyrodon Ledermannii Kallenbach, Zeitschr. Pilzk. 8:96. 1929, ad int.

Described from New Guinea, this species is too incompletely charac-
terized.

SPECIES EXCLUDENDAE

Gyrodon capensis Sacc. is Phlebopus capensis (Sacc.) Sing.

Gyrodon castanellus (Peck) Sing. see under Xerocomus squarrosoides (Snell) Sing.

Gyrodon Emilei (Barbier) Killermann, Denkschr. Bayer. Bot. Ges. Regensb. 16:22.
1925 is Boletus spec., probably B. aff. rubellus Krombh.

Gyrodon filiae (Gill.) Sacc. & Cub. is Boletus edulis Bull. ex Fr. (a monstrosity)

Gyrodon fusipes (Heufl. in Rab. Exs.) Sacc. is Suillus placidus (Bon.) Sing.

Gyrodon immutabilis (Britz.) Sacc. & Trott. is a Boletus spec., possibly B. appen-
diculatus; according to Killermann, it is Phlebopus sulphureus (Fr.) Sing.

Gyrodon miramar (Roll.) Sacc. & Trott. is Suillus granulatus (L. ex Fr.) Kuntze
(sec. Konrad & Maublanc)

Gyrodon Mougeotii (Quél.) Sacc. is Boletus pulverulentus Opat. (sec. Konrad &
Maublanc)

Gyrodon Oudemansii (Harts. in Flor. Bat.) Sacc. is Suillus placidus (Bon.) Sing.

Gyrodon placidus (Bon.) Sacc. is Suillus placidus (Bon.) Sing.

Gyrodon rubellus McWeeney fide Rea is not a Gvrodon.

Gyrodon volvatus (Pers.) Opat. is a parasitized or anomalous Amanita spec.

Subfam. Suilloideae Sing. subfam. nov.

Hyphis fibuligeris vel haud fibuligeris; hymenophoris boletinoideis (parietibus lamel-
lose i.e. radialiter dispositis) aut velatis aut glandulosis (glandulis numerosioribus ad
stipitem) aut — si characteribus his carent — pellicula viscosissima praesente et tunc
sporis constanter minoribus, pallide melleis elongatis, cystidiisque plerumque incrustatis
et majusculis, stipite plerumque cylindrico vel subcylindrico, hymenophoro plerumque
haud distincte depresso circum stipitis apicem et tramate distincte bilaterali praedito;
si autem his characteribus haud clare gaudet, tubuli ex integro rubidi sunt. Habitatio:
Cum arboribus coniferis mycorrhizam formant.

Characters of the subfamily: see key, p. 226, and Latin diagnosis.

The genera Boletinus, Psiloboletinus, and Suillus have very decidedly a
close and natural affinity notwithstanding the difficulties one experiences in
briefly outlining the characters they have in common, as may be seen in
the above Latin description of this subfamily. When working on these
fungi one cannot fail to notice that there is a much closer relationship
between all these genera than between any one of them and any other
boletaceous genus.

5. Psiloboletinus Sing. gen. nov.

A genere Boletino veli absentia et hymenophoro magis lamelloideo marginem pilei
versus differt.

Characters of the genus: see key p. 226 and the above Latin diagnosis.
The type species is P. lariceti (Sing.) Sing. comb. nov. (PAviloporus
lariceti Sing.)

At the time of the discovery it seemed wiser to refrain from the erection
of a new genus for what we then called Phylloporus lariceti. A more pro-

SINGER: FLoRIDA BOLETINEAE II. 251

found study of the affinities of this species, however, as well as of Phyl-
loporus rhodoxanthus has become possible since then and, as a result, we
now propose to base a new genus of Suilloideae on the former while the
latter enters the subfamily Xerocomoideae as the type species of Phyl-
loporus. The ecology and the chemical characters of P. lariceti and P. rho-
doxanthus make this disposition necessary. Psiloboletinus does not occur
in Florida.

6. Boletinus Kalchbr., Bot. Zeit. 25:182. 1867.

Euryporus Quél., Enchir. p. 163. 1886. Type, E. cavipes (Opat.)
Quél.

Characters of the genus: Pileus fibrillose to squamose, or even squarrose,
the fibrils or scales not superimposed upon a viscid layer and therefore not
easily detersible as fragments of a veil, the surface decidedly dry even in wet
weather except for Boletinus spectabilis when the pileus becomes viscid
starting from the margin and gelatinizing inwards; spores comparatively
small to medium (up to 13.2 » in our measurements) pale melleous with
occasional darker spores, cystidia voluminous and mostly covered by a
resinous colored incrustation; clamp connections potentially present in
most species, i.e. at least some populations or carpophores of a majority of
species have at least some scattered clamps while in some species a ma-
jority of specimens does not show clamps at all, and in some species,
clamp-bearing specimens have not yet been found; hymenophore more
(in the majority of cases) or less (especially in B. amabilis and B. pictus)
boletinoid, i.e. pores radiately arranged and rather long, the radial walls
lamella-like, especially toward the stipe; stipe usually cylindric or sub-
cylindric and constantly without glandulae (some glandulae might how-
ever be seen on the pores if a lens is used); veil always present though not
always persistent and not regularly forming an annulus, either simple or
double; mycelium connected with certain conifers (Pinus, Larix, Picea,
Pseudotsuga) with which all Boletini are associated, forming mycorrhiza.
The type species is Boletinus cavipes (Opat.) Kalchbr. (Boletus cavipes
Opat.).

KEY TO THE SUBGENERA AND SECTIONS

A. Stipe hollow, or, if solid, the pores comparatively extremely wide; mycorrhiza al-

ways with Larix; clamps constant. Subgenus Eu-Boletinus
B. Stipe hollow. Sect. Cavipedes
B. Stipe solid. Sect. Palustres

A. Stipe solid; pores medium-wide to wide, not as wide as in the sect. Palustres if the
size of the pileus is considered; clamps either not constant, or very scattered, or
even entirely absent in all the material studied thus far. Subgenus A porpiellus

C. Veil duplex, the interior veil gelatinizing and forming a glutinous inner layer
of the annulus and a viscid marginal zone in older specimens.
Sect. Spectabiles
C. Veil simple or double but neither layer gelatinizing with age.
Sect. Solidipedes

252 FaRLowI14, Vor. 2, 1945

Subgen. Eu-Boletinus Sing. subg. nov.
Hyphis fibulis numerosis constanter instructis; stipite interdum cavo; mycorrhiza
cum larice.

Characters of the subgenus: see key and Latin diagnosis above. Type
species Boletinus cavipes (Opat.) Kalchbr.

Sect. Cavipedes Sing., Rev. de Mycol. 3:163. 1938.

Characters of the section: Stipe hollow; pileus squamose; hymenophore
strongly boletinoid, pores wide but not as extremely wide as in the Palus-
tres; veil not gray; gelatinizing surfaces absent; spores often small (smaller
than 10 » in a majority of cases); clamp connections numerous and con-
stant; context unchanging. The type species is B. cavipes (Opat.) Kalchbr.
Another species is B. asiaticus Sing. None of these has been observed in
Florida.

Sect. Palustres Sing., Rev. de Mycol. 3:162. 1938.

Characters of the section: Stipe solid; pileus squamose; hymenophore
strongly boletinoid, pores extremely wide in comparison with the size of
the pileus; spores small (smaller than 10 »); clamp connections numerous
and constant; hyphae of the cuticle with a wall of about | diameter,
grayish-fuscous in ammoniacal medium; context unchanging on the air
but somewhat more readily reacting with chemical reagents, especially
FeSO,.? Type species B. paluster (Peck) Peck. No representatives are
known in Florida.

Subgen. Aporpiellus Sing. subg. nov.

Fibulis haud praesentibus in omnibus carpophoris (i.e. inconstantibus) et si adsunt,
haud praesentibus in omnibus septis (ic. infrequentibus) ; stipite nunquam cavo;
mycorrhiza cum larice et aliis arboribus coniferis.

Characters of the subgenus: See key and Latin diagnosis above. Type
species B. pictus (Peck) Peck.

Sect. Spectabiles Sing., Rev. de Mycol. 3:157. 1938.

Characters of the section: Veil duplex, i.e. interior veil membranaceous
and gelatinizing in the annulus and on the margin of the pileus: exterior
layer squamulose; hyphae of the pileus thin-walled; stipe solid; clamp
connections none, or in some specimens present though very rare. The
only species known is B. spectabilis Peck, forming mycorrhiza with larch,
and therefore not occurring in Florida.

"In B. paluster, the ““carmine” color of the scales becomes more vivid with formol
which also provokes at times a very slight reddening of the context; KOH on surface
of pileus black for about 1 second, then pale brownish; on context red for about one
second, then lilac, then sordid; NH,OH on surface of pileus, darker and then quickly
bleached; on context, darker yellow; FeSO, on stipe, greenish gray. The whitish vel-
low to lemon yellow context of the pileus, and the pale orange context of the stipe, the
more reddish context of the base and the “Isabella color” pores and tubes are not
subject to autoxydation.

SINGER: FLORIDA BOLETINEAE II. 253
Sect. Solidipedes Sing., Rev. de Mycol. 3:158. 1938.

Characters of the section: Veil double or simple but not showing different
consistency in age, the whole plant remaining dry, never viscid, or scarcely
so; stipe solid, eventually sometimes slightly excavated; pores wide (about |
as wide as in the sect. Cavipedes, or smaller), in two species (B. amabilis,
B. pictus) not quite as distinctly boletinoid as usual in the other sections;
veil frequently tending to be or to become gray. Type species B. pictus
(Peck) Peck.

This section is the one richest in species, and also shows the greatest
variation in distribution and in ecologic requirements. The author formerly
(Rev. de Mycol. 5:6. 1940) transferred Boletus Lakei to Ixocomus (i.e.
Suillus) but further studies on this species and allied forms have shown
that it belongs in Boletinus rather than Suillus. It grows with Pseudotsuga.
B. Benoisti, B. grisellus, most forms of B. oxydabilis and B. ochraceoroseus
grow with larch and B. amabilis has been indicated with Picea. Naturally,
these species do not occur in Florida. Of the species associated with pine,
B. pictus does not occur so far south, thus only B. decipiens is found in the
pine woods of this state.

KEY TO THE SPECIES

A. Context distinctly and strongly (in young fresh specimens) changing color when
bruised; annulus often gray or tending to become so; mycorrhiza with Pinus, or
with Larix sibirica or L. dahurica; pileus ‘““Vandyke red” to “acajou red” when
young, if growing with pine.

B. Mycorrhiza with Pinus. B. pictus
B. Mycorrhiza with Larix.
C. Pileus deep raspberry-red to bright brick red; mycorrhiza with Larix
dahurica. B. Benoisii
C. Pileus merely with an occasional purplish brick color shade; mycor-
rhiza with Larix sibirica. B. oxydabilis

A. Context slightly changing color over very limited areas when broken, or indistinctly
and inconstantly discolored after rather long exposure to the air, or not changing
at all; annulus not gray; pileus never entirely ‘““Vandyke red” or “acajou red”;
fungi forming mycorrhiza with Pinus, Larix (but not LZ. dahurica nor L. sibirica),
Pseudotsuga, Picea.

D. Mycorrhiza with Larix occidentalis and other species of Larix.
E. Spores around 4-5 uw broad; pileus never pinkish. B. grisellus
E. Spores around 3-3.5 w broad; pileus inclining to pink.
B. ochraceoroseus
D. Mycorrhiza with other conifers than Larix.
F. Mycorrhiza with Pseudotsuga or Picea; western species.
G. Spores around 4.2 w broad and (9.5)—11-(12.3) mw long.
B. amabilis
G. Spores around 3.5 w broad and (7)-8.5—(10.7) yu long.
B. Lakei

F. Mycorrhiza with Pinus; eastern species. 7. B. decipiens

254 Fartowia, Vor. 2, 1945

DESCRIPTION OF THE FLORIDA SPECIES

7. Boletinus decipiens (Berk. & Curt.) Peck, Bull. N.Y. State Mus. 8:78. 1889.

Boletus decipiens Berk. & Curt., Ann. Mag. Nat. Hist. II, 12:430. 1853.
Suillus decipiens Kuntze, Rev. Gen. Plant. 3°:535. 1898.

Boletinus Berkeleyi Murr., Mycologia 1:6. 1909.

Boletinus floridanus Murr., Lloydia 6:224. 1943.

Prater I, ric. 5-7.

Pileus “maize yellow” or paler, with “cinnamon buff” or “clay color”
scales, sometimes becoming somewhat “Verona brown” after rains, not
viscid at any time, the scales irregular, rather small (1-2 mm. in diameter),
appressed, the surface convex, not or indistinctly umbonate, 35-70 mm.
broad. — Hymenophore “deep colonial buff,” nearing “honey yellow” when
mature, typically boletinoid and very compound, adnato-decurrent, the
decurrent tooth running down over about one third to one half of the stipe’s
length, and this decurrent portion usually entirely covered by the veil,
tubes moderately long, pores concolorous. — Stipe “cinnamon buff,” or,
especially below, “pinkish cinnamon,” frequently with some “Sudan brown”
below, with an incomplete and not quite constant network immediately
below the fertile zone of the apex, solid, tapering downwards, 30-36 x
10-12 mm.; veil “pale pinkish buff” to “tilleul buff,” apical, dry, fibrillose,
appressed, rarely forming a well-developed annulus; mycelium sordid. —
Context of the pileus and the apex of the stipe ‘“‘pinkish buff,” of the
middle of the stipe “straw yellow,” of the base “orange cinnamon,” most
of the context unchanging but at places becoming ‘“‘vinaceous tawny”
(reddening) in the pileus on prolonged exposure; odor agreeable but
different from the agreeable odor of the Suilli, not strong; taste mild.

Spores 7-10.5 x 3-4.2 p, mostly 8.7-9.5 x 3.5-3.7 , pale brownish or
pale melleous, smooth, more or less cylindric to subfusoid or subellipsoid,
decidedly asymmetric, with rather thin wall; basidia 24-30 x 6-8.5 up,
4-spored, the spores formed on sterigmata of the usual shape (half-sickle-
shaped), 7 » long; cystidia up to 50x 13 p», usually clavate to subfusoid,
incrusted by a ferruginous-fulvous, rarely hyaline incrustation; tramal
hyphae without clamp connections, truly bilaterally arranged; hyphae of
veil clampless, 1.5-8 » in diameter; Ayvphae of the cuticle in interwoven
strands of parallel filaments, hyaline, with hyaline or brown punctate in-
crustations or entirely golden-ferruginous at places, the terminal members
often clavate, many with a mucilaginous incrustation, the walls very thin
to moderately thick, the diameter 3-11 p», the septa without clamps.

Chemical reactions: KOH on surface of pileus, green-black; on context
of stipe, green-black, almost black; NH on surface of pileus, pink and on
context of stipe, “cinnamon russet”’; NH ,OH on surface of pileus, rusty-
brown or after a short time becoming sordid blackish lilac; on context of
stipe, “cinnamon russet,” then slowly almost black; HNO, on surface of
pileus, negative; on context of stipe, “cinnamon rufous”; FeSQ, in all
parts, slowly black.

SINGER: FLORIDA BOLETINEAE II. 255

Gastroid condition: Some of the smaller carpophores of the same my-
celium do not, at some localities, open up at all and remain indefinitely in
a Gasteromycete-like condition; they are piriform with columella (stipe)
and labyrinthiform loculi (hymenophore) confined to the underside of a
pileate portion of the peridium (pileus), producing spores of the same size
(7-9.5 x 3-3.8 »), shape, and color, on basidia of the same type as in the
pseudoangiocarpous form.

Habitat: On the ground in flatwoods and in sphagnose swamps under
Pinus palustris, P. caribaea and possibly other species.

Distribution: Southeastern states of the U.S.A. north to New Jersey and
west to Kentucky (according to Murrill); occurring throughout Florida
but rarely collected.

Material studied: Fla. Alachua Co. east of Gainesville, July 20, 1943,
R. Singer, F 2840 (FH); Lake Co., Eustis, Oct. 1897, R. Thaxter (det.
Singer) (FH); Dade Co., Addison Hammock, February 2, 1906) Je4K.
Small (det. W. A. Murrill) (NY). S.C. type and authentic material from
Society Hill and Santee Canal (1312) and Ravenel (401) (FH). Ala.
Auburn, F. S. Earle (FH) ; other specimens in the herbaria consulted are not
in good condition, and have therefore not been studied carefully, or they
obviously belong to other species such as Xerocomus squarrosoides (Snell)
Sing. or Gyrodon merulioides, etc.

Peck correctly transferred this species to Boletinus. Whether his own
collections belong to B. decipiens is not quite certain, but inconsequential.
Specimens in the Curtis Herbarium, collected by Peck, are not too con-
vincing, and may belong to B. pictus. Material from Maine may belong to
B. cavipes. All Massachusetts material known to me is Xerocomus squar-
rosotdes.

The existence of gastroid forms is considered by this author as an addi-
tional evidence of the affinity of the Boletineae and certain Gasteromycetes.
These forms are fructifications with incomplete individual development,
not reaching the second gymnocarpous stage characterizing Boletinus in
general. If the Boletineae as a whole are, as this writer thinks, derived
from certain Gasteromycetes such as Truncocolumella and related forms, it
would follow that the gastroid forms of Boletinus decipiens are atavistic
aberrations whose formation may be favored by climatic factors, e.g. sudden
decrease in humidity during the fruiting period. Specimens were sent to
Dr. S. M. Zeller of Corvallis, Oregon with the pointed question as to simi-
larity with gasteromycetous genera. Dr. Zeller replied that “the gastero-
mycetoid stage of Boletinus decipiens was received in good condition. It
comes closer to Truncocolumella than any other described genus of Gastero-
mycetes, I believe.” It ought to be noted well that this comes from: the
author of the genus Truncocolumella, an undoubtedly gasteromycetous genus
with the spores borne acrogenously on the sterigmata. It is worth remember-
ing that this genus had been described, shortly afterward, as Dodgea by
Malencon, and that Dodgea was the main subject of discussions on boleti-
noid affinities by Malencon and this author. With these facts at hand, it

256 FARLOwIA, VOL. 2, 1945

would certainly not be a scientific approach to shut the eyes and talk of
convergence.

EXTRALIMITAL SPECIES

Boletinus pictus (Peck) Peck, Bull. N.Y. State Mus. 8:77. 1889.

Boletus pictus Peck, Ann. Rep. N.Y. State Cab. 23:128. 1872.
Boletus Spraguei Berk. & Curt., Grevillea 1:35. 1872.
Boletus Murraii Berk. & Curt., Grevillea 1:36. 1872.

The pileus of fresh material is ‘“Vandyke red,’ margin “acajou red,”
scales ‘‘burnt lake,” later ‘‘Pompeian red” with the scales “brick red,” and
the yellow of the flesh showing through; pores somewhat paler and yel-
lower than “‘old gold”; stipe “‘acajou red” above or ‘‘acajou red” to “brick
red,” or with the base ‘brick red” to “Hessian brown” (actually this is the
color of almost reticulate fibers which are appressed to the “antimony yel-
low” to “light orange yellow” ground); mycelium dirty whitish; context
of the pileus ‘straw yellow,” soon becoming clouded with pinkish to gray-
ish, context of the stipe “mustard yellow,” of the base “ochraceous tawny,”
soon becoming more “tawny” because of autoxydation; veil densely corti-
noid-fibrillose, initially “light pinkish cinnamon” becoming “tilleul buff”
and eventually gray in the outermost layer.— KOH on flesh of pileus,
deep olive, becoming blackish olive to black; FeSO, on flesh of pileus, deep
sordid olive; formol on flesh of pileus, incarnate; NH,OH as with KOH
but reaction weaker.

Boletinus Benoisii (Lebedeva ex) Sing., Rev. de Mycol. 3:159. 1938.

Occurring under Larix instead of under pines, this seems to be specifically
different from B. pictus,

Boletinus oxydabilis Sing., Rev. de Mycol. 3:160. 1938.

For the complete data on this species and its variety see the original
description (d.c.) of the Siberian type.
Boletinus grisellus Peck, Mem. N.Y. State Mus. 3:169. 1900.

Described from Massachusetts, but rather profusely distributed, this
species has been described by Peck (/.c.), Murrill (N. Am. Fl. 9(3):159.

1910), Singer (/.c. p. 162), and others. Type specimens are preserved
(NYS, FH).

Boletinus ochraceoroseus Snell, Mycologia 33:35. 1941.

Described from Idaho, this species has later been proved to be associated
with Larix (see Slipp & Snell, Lloydia 7:18. 1944). The type (FH) ap-
pears to be different from B. Lakei and B. amabilis.

Boletinus amabilis (Peck) Snell, Lloydia 7:17. 1944.
Boletus amabilis Peck, Bull. Torr. Bot. Club 27:612. 1900.

Described from spruce woods in Colorado, part of the type is preserved

SINGER: FLORIDA BOLETINEAE II. Zo7

at FH, and the spores were found to be decidedly larger than in B. Laket.
Judging from what material I had, I found it difficult to believe that B.
amabilis as described by Slipp & Snell (/.c.) is the same as the type of this
species. The description fits much better in B. Lakei which actually is as-
sociated with Pseudotsuga. Whether or not this and the following species
are races or forms, cannot be decided with certainty.

Boletinus Lakei (Murr.) Sing. comb. nov.

Boletus Lakei Murr., Mycologia 4:97. 1912.
Ixocomus Lakei Sing., Rev. de Mycol. 5:6. 1940.

Though the specimens received by this author in 1939 from Dr. Alex-
ander H. Smith were identical with the type specimens (NY), they do not
seem to belong in Suillus now that the related species, B. amabilis and B,
ochraceoroseus, have been studied and found to belong in Boletinus. The
section Pseudotsugini of Ixocomus is therefore to be abandoned as a
synonym of the section Solidipedes of Boletinus.

SPECIES INCOMPLETELY KNOWN

Boletinus appendiculatus Peck, Bull. Torr. Bot. Club 23:418. 1896, from Washing-
ton, D. C.

Boletinus solidipes Peck, Bull. N.Y. State Mus. 167:38. 1912, from Maine.

Boletinus subgrisellus Sing. ad int., Ann. Mycol. 40:27. 1942 nom. subnud.

Boletinus glandulosus Peck, Bull. N.Y. State Mus. 131:34. 1909, from Nova Scotia.

SPECIES EXCLUDENDAE

Boletinus castanellus Peck.

This is hardly a Boletinus. No type is preserved and the species, as
understood by some is Gyrodon merulioides, or as understood by others,
e.g. Coker and Beers, is Xerocomus squarrosoides (Snell & Dick) Sing.

Boletinus flavoluteus Snell apud Snell & Dick is a Suillus.

Boletinus merulioides (Schw.) Coker & Beers, is a Gyrodon, G. merulioides.
Boletinus porosus (Berk. im Lea) Peck is also G. merulioides.

Boletinus punctatipes Snell & Dick is Suillus punctatipes.

Boletinus squarrosoides Snell & Dick is not a Boletinus. It can be considered as
either a Phylloporus or a Xerocomus, X. squarrosoides (Snell & Dick) Sing.

7. Suillus Micheli ex S. F. Gray, Nat. Arr. Brit. Pl. 1:646. 1821 emend.
Snell, Mycologia 34:406. 1942.

Pinuzza Micheli ex S. F. Gray, l.c. (type Pinuzza flava (Bolt.) S. F.
Gray)

Rostkovites Karst., Rev. Mycol. 3:16. 1881 (proposed lecto-type
R. granulatus (L.) Karst.)

Cricunopus Karst., l.c. (proposed lecto-type C. luteus (L.) Karst.)

Boletus Dill. ex Fr. sensu Karst., l.c. p. 17, non S. F. Gray.

258 FARLOWIA, VoL, 2, 1945

Viscipellis Quél., Enchir. p. 155. 1886 (proposed lecto-type V. lutea
(L.) Quel)

Versipellis Quél,, l.c. p. 157 1°

Ixocomus Queél., Flore Mycol. p. 411. 1888 (proposed lecto-type
I. luteus (L.) Quél.)

Characters of the genus: Hyphae of the pellicle strongly gelatinized;
spores usually small; cystidia usually rather large, incrusted; pileus more
or less viscid to glutinous; pores small and round to very wide and boleti-
noid; tubes adnate and rarely slightly depressed around the stipe, more
often subdecurrent or decurrent; trama of the hymenophoral walls truly
bilateral, of the Boletus- or Boletinus-type, not of the Phylloporus type;
stipe usually more or less cylindric, with or without glandulae (consisting
of fascicles of dermatocystidia, or consisting of hyphae terminating in
dermatocystidia of two types, dermatopseudoparaphyses and dermato-
basidia); spore print either between “ochraceous tawny” and “Isabella
color” or a rather deep olivaceous; veil either present or absent. either en-
tirely glutinous-gelatinous, or membranaceous and viscid when wet; mycor-
rhiza with conifers only. The type species is S. /uteus (L. ex Fr.) S. F. Gray
(Boletus luteus Linné ex Fr.).

From viscid species of Pulveroboletus, this genus differs in several re-
gards, either by the characters of the veil, indicated above, or by the more
boletinoid hymenophore if the pores are wide, or by forming mycorrhiza
with coniferous trees instead of frondose trees, or by a combination of
characters like these. The presence of a viscid veil, or glands, or red hy-
menophore, or boletinoid hymenophore on a species which is constantly
associated with conifers suffices to consider it as a Suillus, if it is not a
Boletinus (see key, p. 226). The only Pulveroboletus known to have a
glutinous veil, P. corrugatus (Pat. & Baker)Sing. (Boletopsis corrugata
Pat. & Baker) is a tropical Asiatic species, not associated with conifers.
Similar characters or alternative characters or combinations of characters
delimit Suillus from viscid species of other genera (Xerocomus badius,
X. brasiliensis, Leccinum rubropunctum, L. chalybaeum, Boletus Frostii,
Tylopilus plumbeoviolaceus, and others) of the Boletaceae, and Boletellus
of the Strobilomycetaceae.

The author has formerly divided the present genus into four sections,
and though he has studied numerous additional species since then, he still
sees no reason to change this very workable and natural arrangement. The
largest section, Granulati, in fact the only one that occurs in Florida, has
been subdivided by this author in two subsections. In the light of recent
studies it appears that a third subsection should be added in order to take
care of the species with weak alkali-reactions and deeper olive spore print,

*Tt is here proposed to accept Murrill’s proposal to choose the first species of the
genus Viscipellis, V. variegata, as type of the genus. Versipellis also contains elements
of Xerocomus and Boletus, Phlebopus, etc. The same approach was also thought to be
most practical in the case of the genus Rostkovites Karst.

SINGER: FLORIDA BOLETINEAE II. 259

ie. S. hirtellus, S. punctipes, 5. cembrae. The sections and subsections of
Suillus represent themselves, thus, in the following way:

KEY TO THE SECTIONS AND SUBSECTIONS

A. Stipe with a veil, without glandulae; mycorrhiza with Larix. Sect. Larigni
B. Pores usually grayish or cinnamon-orange, rarely yellowish, more than 1 mm.

in diameter, rarely about 1 mm. in diameter. Subsect. Megaporini
B. Pores yellow to ochraceous-olive, less than 1 mm, in diameter, reaching 1 mm.
only in old, large fruiting bodies. Subsect. Leptoporini

A. Stipe either without a veil, or with glandulae, or, at the same time, without a veil
and with glandulae; mycorrhiza with Pinus, Picea, Abies, Pseudotsuga, Tsuga, and
possibly with other conifers, not with Larix.

C. Glandulae present (except sometimes in S. brevipes where they may be so
indistinct as to appear lacking) ; if they are seemingly absent, the stipe re-
mains white for a long time. Sect. Granulati

D. Spore print between “ochraceous-tawny” and “Isabella color”; pores.
small to wide; context of the pileus with a distinct reaction with NHs
(red or lilac) ; pileus without innate fibrils and squamulae.

E.

Pores more than 1 mm. in diameter or about 1 mm. wide; veil
present but often either gelatinized and entirely glutinous, or
reduced to a marginal veil that is rather fugacious.

Subsect. Latiporini

Pores small, decidedly below 1 mm. in diameter (some pores
may become 1 mm. in diameter when old in very large speci-
mens) ; veil well developed, or indistinct, or absent.

Subsect. Angustiporini

D. Spore print deeper olive; pores around 1 mm. in diameter, occa-
sionally either less or more; context of the pileus with a markedly
weak reaction with NHs (almost or quite negative at first, after very
long exposure becoming sordid to gray or violet) ; pileus with an in-
nate fibrillosity at least at first, or fibrillose-squamulose-hirtose.

Subsect. Hirtellini

C. Glandulae entirely lacking; stipe colored from the beginning.
F. Pores and tubes not reddish in fresh condition. Sect. Bovini

GC

G.

Pores large (larger than 1 mm. in diameter); pileus not
squamulose; context with NHs dark rose, then red, and eventu-
ally brownish vinaceous. Subsect. Euryporini

Pores small to medium (1 mm. or slightly less in diameter) ;
pileus squamulose; context reacting indistinctly with NHs (red-
dish-lilac) Subsect. Stenoporini

F. Pores and at least a portion of the tubes colored a dull red or rose-red
in fresh condition. Sect. Piperati

Sect. Larigni Sing., Rev. de Mycol. 3:38. 1938.

Characters of the section: see key. Type species: Suillus Grevillei (K1.)
Sing. (Boletus Grevillei K1.), also type of subsect. Leptoporini Sing. /.c.

Subsect. Megaporini Sing., Rev. de Mycol. 3:41. 1938.

Characters of the subsection: see key. Type species: Suillus aerugi-
nascens (Secr.) Snell (Boletus aeruginascens Secr.). Other species: Suillus

260 FARLowIA, VOL. 2, 1945

flavus (With. ex Fr. sensu Bres., Nuesch) Sing. comb. nov. (Boletus flavus
With. ex Fr. emend. Bres., Zxocomus flavus (sensu) Sing.); S. tridentinus
(Bres.) Sing. comb. nov. (Boletus tridentinus Bres.). None of these have
been observed in Florida.

Sect. Granulati Sing., Rev. de Mycol. 3:38. 1938.
Characters of the section: see key. Type species: same as type of genus.
Subsect. Latiporini Sing., Rev. de Mycol. 3:45. 1938.

Characters of the subsection: see key above. Type species: Suillus
flavidus (Fr. ex Fr.) Sing., comb. nov. (Boletus flavidus Fr.). Other spe-
cies: S. sibiricus (Sing.) Sing. comb. nov. (Jxocomus sibiricus Sing.) ;
S. americanus (Peck) Snell; S. subaureus (Peck) Snell; S. flavoluteus
(Snell) Sing. comb. nov. (Boletinus flavoluteus Snell), and some more,
none of them occurring in Florida.

Subsect. Angustiporini Sing., Rev. de Mycol. 3:45. 1938.
Characters of the subsection: see key above. Type species: S. granulatus
(L. ex Fr.) 8. F,. Gray.

KEY TO THE SPECIES”

A. Glandulae manifest, in occasional forms with indistinct glandulae and then stipe
elongate (longer than the diameter of the pileus) and mycorrhiza not formed with
2-needle pines (at least in nature).

B. Veil present, forming a distinct and constant though not always persistent
annulus.

C. Stipe about six times longer than thick, or more, with numerous
glandulae almost to the base; annulus small to medium sized, not very
distant, and not broadly sheathing, pallid and frequently appressed,
sliding down in some instances; mycorrhiza with 5-needle pines (Pinus
strobus and P. monticola). S. subluteus

C. Stipe and annulus not as described above; mycorrhiza with 2-needle
pines (in nature).

D. Context a warm yellow; annulus very strikingly sheathing and
not distant in its free part; mycorrhiza with Pinus taeda, P.
palustris, P. australis. 8. S. cothurnatus

D. Context white to partially lemon yellow; annulus sheathing
but widely distant in its free portion when normally developed
in not too old specimens; mycorrhiza in nature with Pinus
pinea, P. silvestris, P. nigra, P. montana, P. resinosa.

S. luteus
B. Veil absent or indistinct, never forming annulus.

E. Pileus white or white for the most part and for a very long period;
stipe elongate; NHs-reaction red in mature specimens; mycorrhiza
with 5-needle pines. S. placidus

™ This key, as usual, takes into account all species known to the author, and will be
useful to determine, in this particular case, those species occurring in the eastern United
States, eastern Canada, and Europe.

SINGER: FLORIDA BOLETINEAE II. 261

E. Pileus yellow to brown, more rarely initially white but soon with
brown and eventually often with yellow shades, or pallescent from the
center; stipe short or elongate; NHs-reaction not distinctly reddish in
mature (though not too yellow) context; mycorrhiza with various
sections of Pinus.

F. Spores larger than 8.2 «. European and African races of
S. granulatus.

G. Central and North European form with elongate stipe,
yellow to yellowish brown pileus, and concolorous
glandulae which become darker only in some specimens
and then very late; preferring calcareous soil; associated
with Pinus silvestris and P. montana.

S. granulatus ssp. typicus

G. Mediterranean form with short stipe, white to yellow-
ish brown or fuscous pileus, strongly colored grandulae,
and preference for sandy soil, not growing with Pinus
silvestris in nature. S. granulatus ssp. leptopus

F. Spores smaller than 8.2 4; associated with Pinus strobus (and
P. monticola ?), rarely with other conifers such as Abies.

H. Glandulae conspicuous, not limited to the apex of the
stipe; tubes adnate; veil rarely present.
S. granulatus ssp. Snellii
H. Glandulae minute, present only at apex of stipe; tubes
decurrent; veil marginal, usually rather distinct.
S. albidipes
A. Glandulae very indistinct and small, pallid to brownish, confined to the upper third
of the frequently very short stipe, or more often entirely lacking, at least to the
unaided eye; mycorrhiza with 2-needle pines. 9. S. brevipes

DESCRIPTION OF THE FLORIDA SPECIES

8. Suillus cothurnatus Sing. spec. nov.

Puate I, ric. 8-10.

Pileo approximate coloribus S. lutei gaudente, viscoso, glabro, levi, 16-63 mm. lato;
tubulis ochraceo-alutaceis, olivascentibus in vetustis, poris subconcoloribus, exiguis;
sporis 7.5-9.8x 2.7-3.5 , levibus, pallide melleis; stipite albido, griseolo, pallide
aurantiaco-flavello, saepe fuscescente, glandulis pallidis fulvescentibus vel nigrescentibus
adornato, solido, annulato, 23-40-(60) x 4-11-(13) mm.; annulo pallido, olivascente
vel grisente, ample cothurnante, membranaceo, viscido; mycelio salmoneo; carne
marmorata, aurantiaco-alutacea vel aurantiaca atque carneo-alutaceo-flammata, vel
pallide alutaceo-maculosa. Habitatio: Prope pinus (Pinus australis, P. palustris, P.
taeda) in Florida Americae Borealis. Distinguntur duae subspecies, ssp. aestivalis Sing.
(subspecies typica) stipite elongato, et ssp. hiemalis Sing., stipite breviusculo.

Ssp. aestivalis Sing.. (type subspecies), ssp. nov.

Pileus “ochraceous buff” and “ochraceous tawny,” or “cinnamon buff”
and “clay color,” or just “cinnamon buff,” viscid, glabrous, smooth, naked,
sometimes marbled in these colors, subconic or convex-subumbonate, or
eventually with flattened center, 16-40 mm. broad. — Hymenophore
“ochraceous buff” to “colonial buff” towards the margin, more rarely all
over, with “capucine orange” spots, or between “orange buff” and “light
orange yellow” tending to “yellow ocher” when maturing, or near “honey

262 FARLoOwIA, VoL. 2, 1945

yellow” when mature, adnate with a more or less pronounced decurrent
tooth; tubes about 4 mm. long; pores concolorous, irregular, some radially
elongate and 0.5-1.0 mm. wide in radial direction but never wider than
0.5 mm. transversely. — Stipe brownish white to white, then sordid grayish
white, or assuming yellowish or pale orange yellow hues, finally sometimes
darkening to “hair brown,” glandulose, with the dots being all equally large
or larger above the annulus (and then more scattered than below), brown-
ish becoming badious, tawny, or vinaceous brown, inside solid, equal or
tapering downward, 23-40 x 4-7 mm.; mycelium salmon color; veil con-
necting the margin with the apex of the stipe and running down the stipe,
initially white near the stipe and more glutinous-hyaline near the margin,
disrupted in this latter zone, the inner, lower portion forming an annulus
which sheathes the stipe and has a strong tendency to become inversely
infundibuliform (wider below), becoming sordid-white and then gray on
maturing, more rarely with an olivaceous shade, viscid when wet (the cov-
ering gluten colorless), as high as broad or slightly reflexed-horizontal. —
Context marbled “orange buff” and “light buff,” unchanging, soft and
somewhat watery; odor as in S. punctipes (agreeable), or none; taste not
significant.

Spores 7.8-9.8 x 2.7-3.2 p, most frequently 8.3-8.5 x 2.8-3 p, pale mel-
leous, smooth, ellipsoid-oblong to subcylindric; basidia 19-22 x 5.8-6.8 By
4-spored; cystidia 40-68 x 6.5-10.3 pw, strongly to moderately strongly in-
crusted, mostly elongate-clavate to broadly clavate, more rarely more
fusoid or cylindric, or varying frequency even on the pores; trama truly
bilateral; pedlicle an “ixotrichodermium”; dots of the surface of the stipe
consisting not only of the ordinary dermatocystidia of the Suéllus-type but
also of hyaline structures recalling the scales of Leccinum (Krombholzia)

stipe “dull Indian purple” or “deep purplish vinaceous”; on glandulae dark
lilac. — NH; on surface and context of pileus and on pores little or slow
reaction, especially in young specimens; on cortex of stipe ‘“vinaceous
rufous” when quite mature. —NH,OH on surface of pileus ‘cinnamon
drab”; on context of pileus “cinnamon drab” or indeterminably darkening;
on pores and tubes lilaceous pink, becoming drab, old specimens “vinaceous
rufous’? becoming ‘“‘testaceous”; on context of stipe lilaceous pink, in age
slowly to “dark vinaceous brown,” middle portion more date brown. —
FeSO, very deep olive to black.

Habitat: On the naked or mossy ground and on mossy trunks in ham-
mocks (low and high as well), more rarely in flatwoods and scrub, always
in the neighborhood of either Pinus palustris or P. taeda though occasionally
as far as 11 m. from the nearest pine tree, scattered or in small groups,

SINGER: FLORIDA BOLETINEAE II. 263

fruiting from May to September in north Florida, in central Florida until
October.

Distribution: North and central Florida.

Material studied: Fla. Highlands Co., Highlands Hammock State Park,
R. Singer, F 478, F 478 a, etc. (the type is F 478) (FH); Lake Co., Eustis,
R. Thaxter (det. Singer) (FH); Levy Co., Gulf Hammock, W. A. Murrill,
F 18751 (FLAS); Alachua Co., Gainesville, R. Singer, F 2164 (FH);
near Dayville, R. Singer, F 3035 (FH); Tung oil-mill on Newberry Road,
W. A. Murrill, F 17882 (FLAS); there were notes by W. A. Murrill at
FLAS indicating the occurrence of what must be this form in Columbia Co.

Ssp. hiemalis Sing. ssp. nov.

Pileus between “wood brown” and ’tawny olive” or “sayal brown” when
young, later “ochraceous orange” and “pinkish buff” or “clay color” mixed
with “sayal brown” and “bister,” glabrous, smooth, naked, pulvinate, then
convex with slightly depressed center, 50-63 mm. broad. — Hymenophore
“antimony yellow” to “yellow ocher,” or “primuline yellow” to “aniline
yellow” with a shade of “honey yellow” to “olive ocher” when old, adnate,
decurrent or more or less shallowly sinuate with a slight decurrent tooth,
or plainly adnate; tubes 7-8 mm. long; pores concolorous or with some
glands of the same color as these of the stipe, small (up to 0.5 mm. in diam-
eter) with some larger ones intermixed (up to 1 mm.), not quite round;
spore print between “ochraceous tawny” and “Isabella color.” — Stipe
pallid or whitish, becoming “light cinnamon drab,” on the base “light
salmon orange,” eventually often with sordid shades or yellow portions,
glandulose because of scattered to crowded glandular dots above and below
the annulus which are concolorous with the pileus or paler, later becoming
tawny or black (more often black), equal, solid, 30-35—(60) x 9-1 1-(13)
mm.; mycelium salmon color; veil pallid, later becoming “citrin drab” to
“deep olive” or gray, or some of these colors combined, or concolorous with
the pileus, ruptured along a circular line near the apical surface of the stipe,
and thus leaving an annular sheath on the stipe; annulus having the shape
of a broad sheathing belt on the upper half of the stipe, tending to separate
from the surface of the stipe at its lower white marginal rim, the outer
(marginal) portion of the veil less persistent, eventually even the annulus
which is fibrillose-membranaceous, and viscid to glutinous on its surface,
collapsing or sliding downward, becoming yellow after freezing. — Context
marbled “zinc orange” and “pinkish buff,” or “buff yellow” near the sur-
face of the pileus and sometimes also near the tubes, unchanging; odor like
that of S. punctipes (agreeable), or none; taste mild or very slightly acid.

Spores 7.5-9.8-(12.3) x 2.7-3.5-(5) y, ellipsoid-oblong to cylindric,
smooth, pale melleous; basidia 20-26 x 6-7 p, 4-spored; cystidia 34-90 x
4.7-11 », with castaneous or melleous incrustations, elongately clavate, not
very numerous except at certain spots; trama and pellicle as in ssp. aestivalts.

Chemical reactions: KOH on surface of pileus plumbeous-gray; same
reaction on tubes and context. — NH; on surface of pileus almost negative;

264 FARLowIA, VoL. 2, 1945

)

on pores quickly “carrot red,” then ‘‘vinaceous rufous,’ sometimes even
“Hay’s russet”; on context “carrot red” to “vinaceous rufous,”’ sometimes
even ‘‘Hay’s russet,”’ sometimes bright and lasting pinkish red in the base.
— NH,OH on surface of pileus slowly as with KOH; on pores as with
NHs, then slowly as with KOH; on context as on pores. — H2SO, on con-
text deep reddish brown, subferruginous. — FeSO, on context very deep
olive to black (a very sensitive reaction).

Habitat: On soil under Pinus palustris, P. taeda, and P. australis, fruit-
ing in Florida from October (north Florida) or November to January, farther
north apparently during the fall months.

Distribution: From North Carolina to Florida. In Florida rather fre-
quent in the north and probably also occurring in central Florida.

Material studied: Fla. Alachua Co., Gainesville, December and January
1942-1943, R. Singer, F 1666 (type), F 1666 a (FH); Sugarfoot Ham-
mock, January 1943 R. Singer, F 16666 (FH); Orange Heights, Novem-
ber 1938, W. A. Murrill, F 19400 (FLAS); Gainesville, W. A. Murrill,
8778; from schedulae (with the specimens removed) at FLAS by W. A.
Murrill, it would seem that ssp. Aiemalis also occurs in Marion Co.

The chemical differences of these two subspecies are similar to those
between winter (below freezing) collections and summer collections of Suilli
in general. The carrot red with ammonia is a reaction due to low tempera-
tures, but there is, besides, a more purple reaction with alkali in the summer
form. This and a slight difference in the color of the pileus are, in addition
to the relative length of the stipe, the main distinguishing characters in all
three cases of genuine or spurious seasonal dimorphism in Suillus. Despite
careful and prolonged observation during two growing seasons, we were
unable to ever find any alternation of summer and winter fruiting bodies
at the same station, and, moreover, such an alternation could not even be
considered possible during the time of our observation since the fruiting
bodies of the summer form failed to show up even in the neighborhood of
the places where the winter form had been collected. It must therefore be
assumed that there exist, in this species, two parallel races, each adapted
to one of the two different growing seasons of northern and central Florida.
This biformity may have developed up to this point from a start similar to
that found in Flammulina velutipes (Curt. ex Fr.) Karst. [Myxocollybia
velutipes (Curtis ex Fr.) Sing.| whose seasonally dimorphous forms may be
a more primitive stage of differentiation than what we have observed in
S. cothurnatus. This is the first time that anything like the classical sea-
sonally dimorphous races of R. Wettstein has been discovered in fungi.

It seems that Atkinson’s Boletus luteus belongs partly to S. subluteus,
and partly (fig. 169) to what in North Carolina takes the place of the winter
form of S. cothurnatus. It may also be suspected that S. cothurnatus, to-
gether with similar forms, is hiding in Coker and Beers’ description of
Boletus luteus. Parts of the description and likewise their pl. 5, fig. 2-3
suggest S. cothurnatus. I do not know whether in the Carolinas, so much
farther north, with the biseasonal aspect of the mycoflora apparently in-

SINGER: FLORIDA BOLETINEAE IT. 265

comparable with what is observed in Florida, or at least showing it in a
much lesser degree, the distinction between ssp. aestivalis and ssp. hiemalis
holds. It is quite possible that in another region another race occurs that
shows less seasonal adaptation. It would be interesting to investigate fur-
ther these things now that the distinction of S. cothurnatus from S. luteus
has ceased to be a problem.

S. cothurnatus differs from S. luteus in having the whole context of an
entirely different color. The difference is striking for anyone who knows
the true S. luteus. Seasonal dimorphism has not been observed and obvi-
ously does not occur in S. Juteus. Some smaller differences may be added
but the easiest way to identify S. cothurnatus in the field is by the warm
yellow context. I suspect that the forms associated with the series Australes
of Pinus are always S. cothurnatus, and those associated with the pines of
the series Lariciones are always S. luteus.

As for S. subluteus, commonly found under pines of the section Cembra,
it is decidedly different from both S. cothurnatus and S. luteus. It is true
that the glandulae of S. Juteus are mostly supra-annular though occasionally
also showing below the annulus, while S. subluteus and S. cothurnatus al-
ways have them both above and below the annulus, and more distinct at
that. However, there is no need to stress this point since the main differ-
ences between all these species lie elsewhere. S. subluteus differs from S.
luteus much more evidently in its more elongate stipe, its less distant veil,
and its different colors. S. subluteus differs from S. cothurnatus in the shape
of the annulus and the colors.

S. cothurnatus is edible, and as good as S. Juteus though rarely abundant
enough to be of much use.

9. Suillus brevipes (Peck) Kuntze, Rev. Gen. Plant. 3°:535. 1898. ‘

Boletus brevipes Peck, Ann. Rep. N.Y. State Mus. 38:110. 1885.
. Boletus viscosus Frost, Bull. Buffalo Soc. Nat. Sci. 2:101. 1874.
Boletus pseudogranulatus Murr., Bull. Torr. Bot. Club 67:63. 1940 (a variety).

Var. typicus.

Pileus in all colors occurring in S. luteus, viz. “fawn color,” “Mikado
brown,” after cold rainy weather between ‘“‘Sanford’s brown” and ‘Mars
orange” in some specimens, exposed portions soon bleaching to “cartridge
buff,” “cream buff,” “chamois,” “honey yellow,” sometimes discolored to
an indescribable drab (near “olive buff” or ‘deep olive buff”), often mar-
bled in all these colors, or some of them, occasionally yellow showing
through in frost-bitten specimens, with entire, rarely rimosely cracking,
smooth, viscid pellicle which is easily entirely removable, glutinous in wet
weather, naked and glabrous except for occasional appressed or fimbriate
fibrils of the veil on the margin, often with narrowly sterile margin, with a
dirty whitish covering all over in some of the veiled specimens, especially
as long as they are young and grow in dry weather, convex, often somewhat
depressed in the center, 40-105 mm. broad. — Hymenophore “baryta yel-
low,” “‘straw yellow,” “lemon yellow,” eventually somewhat duller, adnate

266 FarRLowIA, VoL, 2, 1945

or mostly very slightly sinuate near the stipe; tubes 4-6 mm. long, i.e.
rather short; pores concolorous, finally becoming “deep colonial buff,”
small but of varying diameter, in average (2)—3 pores toa mm.; spore print
between “ochraceous tawny” and “Isabella color.” — Stipe white, becom-
ing yellowish at apex and brownish at base when old, but always pure white
all over when young, without glandulae, or often with very small concolor-
ous or pallid to pale brown glandulae which are not readily visible to the
naked eye and are limited to the upper third of the stipe, or sometimes to
the middle portion, better visible on well-dried stipes, non-viscid, solid,
cylindric or slightly ventricose, remarkably short, sometimes even shorter
than thick, 8-30 x 10-24 mm.; veil inconstant and not persistent, never
forming a true annulus and only rarely forming a narrow belt, but often
found on the margin of the pileus, membranaceous-fibrillose, sordid drab to
white or with these colors mixed, more distinctly visible in dry and warm
weather. — Context pure and uniformly white in both pileus and stipe, later
yellowish above the tubes, and in very old, especially frozen material,
sometimes yellowish in most parts (reaching ‘‘yellow chrome’’), soft and
watery as in S. luteus; odor agreeable as in S. /uteus, or none; taste mild
and slightly acidulous.

Spores (6.7)—-7.5-8.8—(11) x 2.7-(3) p, pale melleous, subellipsoid-sub-
fusoid to cylindric (more so from the profile), smooth, thin-walled; basidia
20-21 x 6-7 p, 4-spored; cystidia 55-83 x 8-10.3 p, numerous, strongly in-
crusted, only a few of them non-incrusted or little incrusted, clavate with
a capitate apex, rarely subfusoid-subcylindric; trama truly bilateral; pel-
licle of the pileus consisting of an “ixotrichodermium”; stipe without der-
matocystidia or with scattered ones but in those specimens with more or less
distinct glandulae in dried condition, the latter are composed of fascicles of
hymenium consisting of strongly incrusted clavate-capitate, or subfusoid,
or clavate dermatocystidia, 27-67 x 8-14 p.

Chemical reactions: KOH on surface of pileus, dark gray; on tubes,
sordid gray to nearly ‘bone brown”; on white context, dirty lilac becoming
more grayish and in older drier material lilac for a very long time and then
drab; in the yellow context of the stipe, dirty lilac, becoming more grayish.
— NH,OH on the paler portions of the pellicle causing them to become
darker, then ‘‘purplish vinaceous,” darker portions slowly “purplish vina-
ceous”’; on context of older drier material, lilac then blue. — NH on sur-
face of pileus reacting as with NH,OH; on white context, “alizarin pink”
for a moment, then vanishing, in older and drier pilei “salmon pink”’; on
yellow portions of the context of the pileus, between ‘cadmium orange”
and “orange rufous,” becoming bluish-caesious; on tubes, ‘ochraceous
salmon,” in older drier material reacting as with the yellow context. —
H.SO, on the pellicle nil; on the pores, rusty ochraceous. — Methyl-
paramidophenol very pale grayish hyaline (almost negative), except for
the zone above the tubes and across the apex of the stipe where a slow
vellow reaction takes place, not becoming violet anywhere.

Habitat: In pine woods, under Pinus taeda, P. australis (in Florida; in

SINGER: FLORIDA BOLETINEAE II. 267

other regions with related species of pines), also in mixed woods under
2-needle-pines, gregarious, usually in large groups or rows, fruiting in
Florida from October to May, farther north in fall.

Distribution: North America, most common in the southeastern states.
In Florida mainly in the northern part, not observed in south Florida.

Material studied: Fla. Alachua Co., Gainesville, Dec—Jan. 1942-43,
R. Singer, F 1646, F 1646 b, F 2044 (FH); W. A. Murrill, F 18440 (Bo-
letus granulatus ) (FLAS).— N.Y. Karner (authentic material of Boletus
viscosus Frost) (NYS); also several collections of Peck (authentic ma-
terial of Boletus brevipes) (NYS).

This species is very closely related to S. granulatus ssp. Snellii, and since
material of this latter was the main source of information on S. granulatus
in general available to Murrill, it is not surprising that he considered it as
a synonym in North American Flora. However, when studied in Florida
where S. granulatus ssp. Snellii and similar fungi except S. brevipes do not
occur at all, the species seems to be quite constant in its characteristic
features except for such definite varieties with longer or swollen stipe as
shall be described below. These varieties show a limited though undeniable
convergence toward S. albidipes. If it were not for some minor characters
and the mycorrhizal association, these uncommon atypical forms of S.
brevipes would seem to be indistinguishable from S. albidipes. This may
lead to confusion in places where S. brevipes, S. albidipes, and S. granulatus
ssp. Snellii occur simultaneously, and where mixed stands of conifers make
the separation difficult. It is very probable that these seemingly convergent
forms actually mark the evolutional starting point of the American group
of species gravitating around S. granulatus. S. brevipes, then, would mark
the stage of complete adaptation to 2-needle pines. Since S. brevipes and
its varieties in Florida form a very definite group within the subsection
Angustiporini, I do not combine it with S. granulatus as a subspecies, inas-
much as others may feel that not only S. albidipes and S. brevipes but also
S. granulatus ssp. Snellii and ssp. leptopus (S. Bellinit) should all be treated
as autonomous species, rather than as subspecies of S. granulatus. While
no argument pro or contra is offered here, the author thinks it necessary to
call the attention of the specialists to this very interesting though compli-
cated problem in order to get more new facts from different regions. The
necessity to do so will arise still more urgently when western collections
are compared with material gathered in the East and South.

S. brevipes is, as also has been stated by Peck and others, a good edible
species, providing large quantities of palatable food during the winter
months for the few people who know it.

Var. aestivalis Sing. var. nov.
A typo differt stipite elongato et statura paulum minore. Sub Pinibus taedis in
Florida Americae Borealis Junio-Augusto mensibus.

Differs by a longer stipe than the type variety, and thus approaches
S. albidipes but easily distinguished by the smaller size, the almost non-

268 FarRLowlA, VoL, 2, 1945

decurrent tubes, the white interior that becomes yellow with age (nowhere
brownish red), and the mycorrhizal association with Pinus taeda. It is
known only from the type locality in Gainesville, Alachua Co. Fla. where
it grows infrequently in small groups. The microscopical and chemical
characters are the same as in the type variety. Because of the longer stipe
it corresponds to the seasonal dimorphous subspecies of S. cothurnatus and
S. Airtellus but the difference seems to be less pronounced and less constant.
We are still not entirely convinced that this is a summer race rather than
a mere summer-adaptation alternating, on the same mycelium, with the
winter fruiting bodies. As long as no more observations can be had, we
shall have to term this a varietas in our sense (as a temporary nomencla-
toral status).

Var. pseudogranulatus (Murr.) Sing. comb. nov.
Boletus pseudogranulatus Murr., Bull. Torr. Bot. Club 67:63. 1940.

Pileus “avellaneous” with a slight drab or cinnamon tinge, or rather
tending toward clay color or umber, with paler places, viscid when wet,
moderately shining when dry, smooth and glabrous, but sometimes with
easily separable inconspicuous sordid patches of a thin membranaceous
deep dirty-gray veil on the margin which is very acute, strongly convex and
usually remaining so in the middle while the marginal part becomes flat-
tened, or becoming subplane all over, thick even for a bolete, 60-110 mm.
broad. — Hymenophore between “empire yellow” and “picric yellow,”
somewhat irregular but not gyrose-merulioid, not quite readily separable
from the context, adnate-decurrent, or decurrent and mostly continuing on
the apex of the stipe as an indistinct narrow reticulated belt; tubes 4.5 mm.
Jong in quite adult specimens, i.e. rather short; pores concolorous, small
(about 2 to a mm.).— Stipe initially white but soon becoming “lemon
chrome” and “light cadmium” at places and eventually entirely yellow,
smooth or rarely very indistinctly rugulose near the base, glabrous or with
very small, not very distinct pallid to blackish glandular dots as in S. bre-
vipes var. typicus but often, at least on one side of the stipe, connected by
very fine lines, and thus forming a network that may extend over a large
portion of the stipe, or be confined to the apex, avelate or with a narrow,
indistinct, soon disappearing annular belt which is deep dirty-gray, with
swollen ventricose lower half which may again be tapering toward the very
base, 30-60 x 26-40 mm. (near the middle or base), 17—24 mm. at the apex,
solid; mycelium yellowish or yellowish white. — Context white, vellow in
the peripheral parts of the stipe and in the base, often also toward the
tubes, very thick and solid-fleshy in the pileus except for the swollen part
of the stipe where it may be soft and spongy; odor as in S, luteus; taste
mild or slightly acidulous.

Spores (6.7)—7.5-9.5—(11) x 2.7-(3.3) jy, in shape and color similar to
those of var. typicus; basidia as in var. typicus; cystidia 30-65 x 7.5-10.5
#, Numerous, strongly fulvous-castaneous or melleous from a resinous in-

SINGER: FLORIDA BOLETINEAE II. 269

crustation, oval or clavate above, more rarely fusoid or ampullaceous, with
thin lower part.

Chemical reactions: KOH on context and surface of pileus, drab; on
context of stipe, grayish green becoming black; on pores, deep grayish
brown. — NH; and NH,OH on the surface of the pileus, slate gray; on
context of pileus, rosy pinkish then becoming slate gray; on pores and
tubes, “‘vinaceous rufous,”’ then becoming grayish green, and same colors
seen in the cortex of the stipe. — H»SOx, on the tubes a similar reaction as
with NH;. — Aniline negative. — Formol negative. Methylparamido-
phenol negative. Note: these reactions were obtained with quite mature
fruiting bodies in cold weather.

Habitat: In dense hammocks, under or near Pinus palustris, P. taeda,
or P. australis, solitary, fruiting from October to April.

Distribution: Known only from Alachua Co., Fla.

Material studied: Fla. Type (FLAS); Gainesville, R. Singer, F 1704
(FH); F 1877 (FH).

This species has not a distinctly elongate but rather a ventricose or in-
versely claviform stipe. It is much more variable than may be assumed
from Murrill’s description. The thicker stipe with decurrent tubes distin-
guishes it from the type. The reticulation stressed by Murrill is an incon-
stant character. In only one carpophore out of all I have seen, do I find a
reticulation worth recording. The veil cannot serve as a distinguishing
character since the same kind of veil is observed also in a certain percentage
of fruiting bodies of the type variety, but these fruiting bodies do not differ
from the avelate ones in any other way. Var. pseudogranulatus seems to be
an aberration due to peculiarities of the habitat. It occurs in moister soil
of hammocks where the carpophores are less exposed to the rain. Whether
this is just a forma, or a constant race, cannot be decided immediately.

EXTRALIMITAL SPECIES

Suillus subluteus (Peck) Snell apud Slipp & Snell, Lloydia 7:34. 1944.
Boletus subluteus Peck, Bull. N.Y. State Mus. 1:62. 1887.

Since this species has been confused with S. luteus as well as with the
Floridan S. cothurnatus, I think it is necessary to give a complete descrip-
tion of S. subluteus, based on collections made by this writer in the Hunt-
ington Wild Life Forest, Newcomb, N. Y. (FH, NYS) which were com-
pared with the type of S. subluteus (NYS).

Pileus more or less “chamois” (colored somewhat like the European S.
flavidus), but sometimes more reddish than “chamois,” sometimes more
olive, glutinous when wet, often slightly squamulose with indistinct, ap-
pressed squamulae, convex, 57-63 mm. broad. — Hymenophore various
shades of yellow with an olivaceous shade in age, adnate to slightly de-
current; tubes medium long; pores decidedly smaller than 1 mm. in diam-
eter, only in large, old specimens reaching 1 mm., somewhat angular, be-

270 Fartowra, Vor. 2, 1945

coming somewhat sordid on pressure, somewhat glandular-dotted; spore
print between “ochraceous tawny” and ‘Isabella color.’ — Stipe concol-
orous with the pores, more pallid near the base, beset with light brown,
later, especially below, often with dark brown to almost black glandulae
above and below the annulus, with watery, colorless, or watery-yellow
droplets distilled from the glandular dots on the pores next to the stipe and
on the apex of the stipe, annulate, solid, equal or slightly thickened down-
ward, slender as compared with the usual shape of the stipes of S. luteus,
about 60x 9 mm.; mycelium sordid-grayish; annulus pallid, appressed or
somewhat distant in young specimens, not conspicuously broad-sheathing,
and soon collapsing into a mere slimy belt, apical but sometimes sliding
down to the middle, membranaceous but very glutinous when wet and even-
tually sordid or concolorous with the remaining surface of the stipe. —
Context white, very slightly and slowly turning sordid when exposed, in the
stipe more yellowish when bruised; odor like that of S. punctipes but
weaker; taste mild, slightly acidulous.

Spores 9-10.7—(11) x 3.3-3.8 4, mostly 9.5 x 3.5 w, rather pale melleous,
smooth; cystidia 38-120 x 5.5-7.5 yp, cylindric, obtuse at the apex, or
clavate, almost constantly incrusted with a brown, resinous crust or gran-
ules, glandular dots consisting exclusively of (20)—70-125 x (3.5)-7-9.5 p
large, narrowly clavate, rarely fusoid and then often 6.5 » broad, incrusted
dermatocystidia; Ayphae without clamp connections.

Chemical reactions: KOH on surface of pileus, dark sordid gray; on
pores, dark reddish brown, — NH; and NH,OH on surface of pileus, red-
dish brown; NH; on the context of the pileus, salmon pink; NH,OH on
the context of the pileus, blue; on the pores, dark gray. — Methylpara-
midophenol negative.

Habitat: With Pinus strobus (and probably with other 5-needle pines)
in woods, often together with S. granulatus ssp. Snellii.

Distribution: Eastern States of the U.S.A., not reaching Florida in the
south; the western limits of the species have not been established.

The spores which are larger than those of S. cothurnatus and S. luteus,
provide a good character to distinguish poor specimens or dried material
of S. subluteus from S. luteus and S. cothurnatus. The chemical characters
also seem to be different in these species (see also observations under
S. cothurnatus).

Suillus luteus (L. ex Fr.) S. F. Gray, Nat. Arr. Brit. Pl. 1:646. 1821.

Boletus luteus Linné ex Fr., Syst. Mycol. 1:386. 1821.
Ixocomus luteus Quél., Flor. Mycol. p. 414. 1888.
Cricunopus luteus Karst., Rev. Mycol. 3:16. 1881.

The description and figures given by Kallenbach, Pilze Mitteleuropas
1(8):45, pl. 19. 1928, should be compared with the foregoing descriptions
of S. subluteus and S. cothurnatus. In order to make this comparison com-
plete, I want to add some data on S. /uteus, either not contained in Kallen-
bach’s account or mentioned there in a different manner:

SINGER: FLORIDA BOLETINEAE IIT. 271

Spores in a very fresh print “ochraceous tawny,” under the microscope
(6.8)—7.5-9-(10) x 2.8-3.3 », pale melleous, smooth, ellipsoid-subfusiform
to cylindric. — KOH on surface and context of pileus, “‘slate olive” to dark
dirty-gray; on the context, occasionally reddish at first; on the pores, “bone
brown,” or reddish to brownish red and eventually becoming “bone brown,”
at times passing through a lilac phase before becoming brown. — NH; on
surface of pileus, negative to very pale pink; on context of pileus (where
yellow), ‘salmon buff,” eventually with lilac shades; on pores and tubes,
pale reddish to brownish pink or orange-tawny. — NH,OH on surface of
pileus, little reaction, or slowly dark gray; on context of pileus, “slate
olive” (where it had been yellow). — H2SO, on the yellow context of the
pileus, “primulin yellow’; on pores, ‘ochraceous orange”; on surface of
pileus, negative.

Habitat: Under Pinus silvestris, P. montana, P. nigra, P. resinosa (also
reported with P. pinaster by Kallenbach but this seems doubtful; in experi-
ments, forming mycorrhiza even with Picea and Larix), i.e. in nature always
with pines of the section Lariciones.

Due to the erroneous identification of S. subluteus and S. cothurnatus as
S. luteus, the author was for a while doubtful as to whether the true S. luteus
occurs in America at all. However, a large collection of excellent fresh
material growing under Pinus resinosa (coll. P. Gast, September 1944 in the
Harvard Forest, Petersham, Mass.) proved the fact that a species entirely
identical with the European S. /uteus occurs in this country. It is now diffi-
cult to say whether this was originally imported with Pinus silvestris and
then spread, or — more probably — if it is native in North America. It
does not reach Florida.

Suillus placidus (Bon.) Sing., Farlowia 2:42-43. 1945.

Boletus placidus Bon., Mohl’s Botan. Zeitschr. 19:204. 1861.
Boletus albus Peck, Rep. N.Y. State Cab. 23:130. 1873.”

As for the discussion of this species see Singer, /.c. 1938, and l.c. 1945.
The types of Peck’s have been compared with material collected by this
writer in New York State. B. albus and B. placidus are most certainly
identical.

I do not think that S. placidus occurs in Florida. I have not collected
white Suzlli in Florida at all. W. A. Murrill collected a specimen of what
he calls Boletus granulatus (in his sense a collective species), adding, on
the label (F 16407, FLAS) “white form.” It was more slender-stemmed
than the usual S. brevipes but this would not mean much since it was col-
lected in summer when the slender-stemmed summer form of S. brevipes is
found. The glandulae are not too clearly visible. Schematically, this speci-
men would have to be determined as S. placidus but it may also be a white
form of S. brevipes var. aestivalis, or an exannulate form of S. acidus (Peck)
comb. nov. (Boletus acidus Peck).

* For a complete list of synonyms see Singer, Rev. de Mycol. 3:48. 1938.

272 Fartow!A, Vor. 2, 1945
Suillus granulatus (L. ex Fr.) Kuntze, Rev. Gen. Plant. 3°:535. 1898.

The European type (ssp. typicus), the American (ssp. Snellii), and the
Mediterranean (ssp. leptopus) race have been discussed in Farlowia 2:41-
45. 1945. When studying the western collections of Swili, the author came
across an abundant collection of what is easily recognized as S. granulatus,
but the determination of the subspecies causes difficulties. I cannot, with-
out extensive study of fresh material, indicate whether this Idaho subspecies
is identical with any known subspecies or represents an allied subspecies,
but it probably is not ssp. Swellii. This partly explains the fact that Slipp
and Snell find slight modifications in size and veil of Idaho specimens de-
pending on whether the plants occur with Pinus monticola (sect. Cembrae)
or P. contorta. The specimen shown in the illustration (Lloydia 7:41, pl. 4,
fig. A. 1944) would suggest S. brevipes by its habit but it probably is the
Pinus contorta form of S. granulatus. Good separate descriptions of these
collections may help solve this problem.

Suillus albidipes (Peck) Sing., Farlowia 2:45. 1945.
Boletus granulatus var. albidipes Peck, Rep. N.Y. State Mus. 54(1):168. 1901.
Boletus albidipes Peck, Rep. N.Y. State Mus. 65:22. 1912.

This little known species is about intermediate between S. brevipes and
S. granulatus ssp. Snellii. Since the summer form of S. brevipes or S. bre-
vipes var. pseudogranulatus may be confused with S. albidipes, and also
because of the lack of adequate descriptions of S. albidipes, we include here
a description of it, obtained from material collected by this writer near
Newcomb, N. Y. (FH) and compared with the type (NYS).

Pileus “cinnamon rufous,” somewhat deeper colored when young, slightly
tending toward ‘‘zinc orange,” with the yellow of the flesh showing through
the pellicle of very old and watery specimens, darker and paler spots some-
times tessellately alternating, usually slightly rivulose under the viscose
surface, or areolate when old, pallescent from the middle outward in many
specimens, the margin in very many of the young specimens with a white
or pallid velutinous or fibrillose-tomentose band from an indistinct veil,
convex, obtuse, 40-105 mm. broad. — Hymenophore flesh-whitish, then
yellow as in S. granulatus ssp. Snellii, adnate-decurrent to strongly decur-
rent; tubes as long and the concolorous pores as wide as in S. granulatus
ssp. Snellii; spore print between ‘‘ochraceous tawny” and “Isabella color”
(nearer to the latter). — Stipe pure white becoming somewhat lemon-
yellow in the upper fourth to two thirds when old or in cold weather,
usually stained with tan color in the middle, yellowish fibrillose at the base,
indistinctly glandulose all over or with small scattered white, later pale
brown glandulae at the apex (upper fourth to half), otherwise glabrous and
smooth, non-viscid, solid, often becoming stuffed, cylindric or clavate
(tapering upward), or tapering downward, or ventricose, straight or curved,
50-115 x 8-28 mm.; mycelium white. — Context white, later reddish or
brownish (reminding one of Amanita rubescens in this regard) in the stipe,
especially in the middle of the stipe, in the apex of the stipe rather becoming

SINGER: FLoriIpA BOLETINEAE II. 273

bright lemon yellow than reddish, and also somewhat lemon yellow toward
the tubes; odor as in S. luteus, but very slight; taste mild to slightly
acidulous.

Spores 6.8-9.2 x 2.7-3.2 p, mostly 7.8-8.5x3 p, pale melleous, smooth,
cylindric, fusoid-cylindric, or fusoid-ellipsoid; basidia about 20x 5 yp, 4-
spored; cystidia either hyaline and scattered, or more or less incrusted and
in fascicles or scattered, cylindric-clavate or broadly clavate, their wall
slightly thicker than the wall of the basidia, 42-72 x 6.5-11.5 »; trama and
pellicle as well as glandulae as in S. brevipes.

Chemical reactions: KOH on surface and context of pileus, dirty grayish,
not violet; on pcres, deep dirty fuscous, becoming almost black, blackish
fuscous. — NH; and NH,OH on surface of pileus, more grayish; on con-
text of pileus, pink then lilac (‘‘pale brownish vinaceous”); on context of
stipe, negative, only lemon-yellow places becoming deep orange-brown-
rufous; on tubes, orange-ferruginous.

Habitat; In open coniferous and mixed woods under or near Pinus strobus
(sect. Cembrae), but occasionally, in regions where white pine is common,
also growing with Abies balsamea.

Distribution: From Massachusetts to New York, south apparently to
North Carolina, but western limits unknown.

This differs from S. brevipes in the elongate stipe, the reddish tones in
the context, and the habitat; from S. granulatus in the reduced glandulosity
of the stipe, a slight difference in the chemical reactions, and the reddish
tones in the context. It may eventually be reduced to the status of a sub-
species of the latter.

Subsect. Hirtellini Sing. subsect. nov.

Sporis in cumulo saturate brunneo-olivaceis ; poris circa 1 mm. amplis, interdum
paulum amplioribus vel angustioribus; carne pilei ammoniaci ope vix vel tardive
reagente (grisente vel violascente) ; pileo innate fibrilloso in juventute vel fibrilloso-
squamuloso-hirtello.

Characters of the subsection: see key (p. 259) and the above Latin diag-
nosis. Type species Suillus hirtellus (Peck) Kuntze (Boletus hirtellus
Peck).

KEY TO THE SPECIES

A. Context unchanging or at least not becoming blue on exposure; stipe only very
slightly and gradually tapering toward the apex or almost perfectly equal.

B. Stipe strongly thickened toward the base; pores brownish to brownish-olive,
never clear yellow; glandulae crowded, often running into each other and
making the stipe sticky; numerous non-incrusted fusoid cystidia present on
the pores; spores (7.8)-8.8-11.5 x 3-3.7 mw; color of the pileus often about
“Mars yellow”; mycelium white; ammonia (also vapors) on the surface of
the pileus definitely negative. S. punctipes

B. Stipe slightly if at all thickened toward the base; pores from the early im-
mature to the final overmature stage showing a clear yellow, or more pinkish-
ocher in youth and more yellowish-olive in age; glandulae though distinct
and numerous not really crowded and not forming sticky patches; non-

274 FarLow1a, Vor. 2, 1945

incrusted cystidia infrequent, fusoid cystidia exceptional, or wanting; spores

(6.8)-7.5-10.2 x 3-3.5 «; color of the pileus never ‘““Mars yellow”; mycelium

not always white; ammonia (vapors) on the center of the pileus after long

exposure “Persian lilac,” on the margin “acajou red”; ammonia water lilac,

then carmine, then amethyst color around an eventually “cinnamon buff” spot.
10. S. hirtellus

A. Context more or less bluing; stipe more or less thickened toward the base.

C. Pileus and stipe golden yellow, then sulphur yellow to yellowish cream;
mycelium white, “in seral association in the Thuja-Tsuga-zone” in Idaho.
S. hirtellus var. mutans

C. Pileus ocher brownish to orange brownish with darker squamulae; mycelium
pink; in Larix-Pinus-Picea-stands of the northern slopes of the Altai Mts.
associated with Pinus sibirica. S. cembrae

10. Suillus hirtellus (Peck) Kuntze, Rev. Gen. Plant. 3°:535. 1898.

Boletus hirtellus Peck, Bull. N.Y. State Mus. 2(8) :94. 1889.
Ixocomus hirtellus Sing., Ann. Mycol. 40:30. 1942.

Rostkovites hirtellus Murr., Mycologia 1:14. 1909.

Boletus hirtellus var. siccipes Coker & Beers, Boletaceae, p. 83. 1943.

Ssp. cheimonophilus Sing. ssp. nov.

Mycelio plerumque manifeste albido, carpophoro majore, stipite breviore latitudine
pilei, tubulis circum stipitem raro distincte depressis; sporis saepe elongatis, tenuibus,
plus minusve cylindricis; forma hiemalis.

Pileus ‘‘barium yellow” to “pinard yellow” with subconcolorous or some-
what darker innate fibrils which form subappressed but hirtose squamulae
on the disk, dry in dry weather, decidedly viscid in wet weather, occa-
sionally showing a grayish band around the extreme margin (veil), convex,
becoming subplane, 50-90 mm. broad. — Hymenophore “wax yellow” to
“olive lake,” adnate to slightly depressed, not decurrent; tubes moderately
long; pores concolorous but often with some brownish glandulae, medium
wide (around | mm. in diameter), not exuding drops unless diseased; spore
print rather deep olive. — Stipe yellow, pallid toward the base, beset with
numerous pale brown, then testaceous-chocolate, eventually blackish glan-
dulae over its entire length, solid, equal or at least equal in its upper two
thirds, and then the base slightly thickened and again acuminate below,
40-70 x 5-20 mm., mostly 45-60 x 5-20 mm.; veil none on the stipe;
mycelium mostly whitish. — Context pale yellow, unchanging; odor none;
taste mild.

Spores 8-13.5 x (2.5)—3-3.3-(3.7) 2, melleous, smooth, cylindric or sub-
fusoid-cylindric, rarely subellipsoid with suprahilar depression, or without
depression; basidia and cystidia as in ssp. thermophilus; trama and glan-
dulae not studied.

Chemical reactions: KOH on surface of pileus and stipe and on the pores
and context, “‘bay” but somewhat deeper. — NH. and NH,OH on surface
of pileus, negative for a long time, then assuming a vinaceous hue, then
“cinnamon buff”; on pores, more sordid grayish olive, then becoming deep
olive gray with a cinnamon ring; on context, an undefinable grayish tinge,
then becoming pallid sordid brown.

SINGER: FLORIDA BOLETINEAE II. 205

Habitat: On sandy soil under pines, in Florida under Pinus palustris,
P. australis, and P. taeda, usually solitary, or in small groups, fruiting in
Florida in December and January.

Distribution: Florida (if the northern form is included, it is distributed
from New York to Florida with the western limits unknown).

Material studied: Fla. Alachua Co., Newnan’s Lake, Jan. 30, 1943,
R. Singer, F 1772 (FH); Magnesia Springs, Jan. 5, 1941, G. F. Weber,
(indet.) F 19826 (FLAS).

Ssp. thermophilus Sing. ssp. nov. TEXT-FIGURE 1.

Mycelio plerumque manifeste roseolo; carpophoro minore; stipite longiore latitudine
pilei; tubulis circum stipitem plerumque distincte depressis; sporis forma sua haud
distinctis; forma aestivalis thermophila.

Pileus initially cinnamon, soon becoming “baryta yellow,” somewhat
“maize yellow” on the disc, then “Martius yellow” or “sulphur yellow,” or
“light Chalcedony yellow,” subviscid, innately fibrillose and minutely hir-
tose, the margin initially covered with cobweb-like to woolly remnants of
the veil which in the primordia continues into an indistinct, glutinous,
hyaline veil that disappears before full maturity, convex, soon flat around
the umbo, but sometimes without an umbo (subumbonate or obtuse),
38-58 mm. broad. — Hymenophore between “deep colonial buff” and “olive
ocher,” depressed around the stipe; tubes 5-8 mm. long; pores initially
“capucine buff,” then “ochraceous buff,” then “barium yellow,” then “light
ochraceous buff,” and eventually between ‘“‘deep colonial buff” and ‘“‘dark
olive buff,” or ‘‘sulphine yellow,” small (2 toa mm.) to medium (7 to 5 mm.
tangentially, or 4 to 5 mm. radially), up to 1 mm. in diameter; spore print
deep olivaceous. — Stipe concolorous, the brightest portions about “primu-
line yellow,” beset with glandular dots which are sparse to very numerous,
initially white or cinnamon, later concolorous with the remaining surface of
the stipe, or whitish, then becoming carmine red, brownish red or indefi-
nitely remaining concolorous, the upper part of the stipe and also sometimes
the pores guttulate (droplets consisting of a whitish, rarely avellaneous
latex), entirely solid, cylindric (after having been thinner at the apex in
very young stages) to subequal, or with a root-like tapering base, 50-60 x
6-9 mm.; mycelium mostly salmon color; veil none. — Context yellow,
unchanging; odor none, or very slight, like that of S. /uteus; taste mild.

Spores (6.8)—7.5-10.2 x 3-3.2 y, brownish melleous, some pale melleous,
smooth, fusoid to ellipsoid, in profile cylindric to fusoid; basidia 26 x7 yn,
4-spored; cystidia 37—71 x 5.3-16 », with about 0.5 » thick walls (the often
strongly developed resinous incrustation not included), most of them colored
because of the incrustation, few hyaline, elongate-clavate, in fascicles;
dermatocystidia of the glandulae of the stipe of one type only, viz. the
incrusted, clavate (Suillus-) type, 17-77 x 6—-9.5 »; hyphae without clamp
connections.

Chemical reactions: KOH on surface of pileus, deep brownish yellow or
lilaceous brown; on pores, dirty brown; on context, various shades of

276 FarLow!A, VoL. 2, 1945

brown. — NH, little reaction immediately but if exposed long enough, the
disc of the pileus becomes “Persian lilac,” and the margin “acajou red”; on
pores, slowly drab; on context, negative or similar to the reaction on the
surface of the pileus. —NH,OH on surface of pileus, lilac, then carmine,
then amethyst color, then fading; on the context of the stipe, almost nega-
tive, or slowly sordid; in the base of the stipe, rapidly drab. — FeSO, on
the context, slowly olivaceous yellow.

Habitat; In mixed stands of low hammock and in flatwoods under Pinus

TEXT-FIGURE 1. Suillus hirtellus (Peck) Kuntze ssp. thermophilus Singer.
Fresh specimens from Highlands Hammock State Park, Florida after pro-
longed exposure to ammonia vapors to bring out the hirtose character of
the pileus. x 34.

palustris in small groups, fruiting from June to September, or perhaps
October.

Distribution: From Georgia to south Florida (not in the tropical zone).

Material studied: Fla. Highlands Co., Highlands Hammock State Park,
Aug.—Sept. 1942, R. Singer, F 543 (type) (FH); F 543 a, F 5436 (FH);
Alachua Co., Gainesville, W. A. Murrill, (as B. granulatus) F 16526
(FLAS).— Ga. Vicinity of Talullah Falls, north bank of river above
bridge, Sept. 1901, A. B. Seymour & W. L. Moss (det. W. H. Snell as
B. hirtellus) (FH); Fern Valley, Sept. 1901 (FH); Fern Brook and Bad
Branch, Sept. 1901 (FH).

I am at present not in a position to decide whether the northern form,
z.e. the type form, belongs to any of these subspecies or constitutes a sub-
species by itself. From the specimens I have seen, and from the best de-
scriptions available, I would guess that it belongs rather to ssp. cheimo-

SINGER: FLoRIDA BOLETINEAE II. Agi

nophilus. More phenological observations, however, will have to prove this
point. It is possible that the northern form is unadapted to any special
season and shows some distinguishing character or characters that would
make it advisable to separate it as ssp. typicus.

We have examined authentic material of Boletus hirtellus var. siccipes
Coker & Beers, J.c. The differences between this and the Albany specimens
are so slight as to be considered negligible.

EXTRALIMITAL SPECIES

Suillus hirtellus var. mutans (Peck ex Snell) Snell apud Slipp & Snell, Lloydia 7:23.
1944.

Boletus hirtellus var. mutans Peck ex Snell, Mycologia 33:26. 1941.

Boletus tomentosus Kauffman, Pap. Mich. Acad. Sci., Arts & Lett. 1:117. 1921,
non Krombh. Nat. Abb. Schw. 5, fl. 36, fig. 19-20. 1836 (year uncertain),
nom. nudum; nec Boletus tomentosus Raddi ex Sacc. Syll. 23:346. 1925.

I have seen only Henderson’s specimens (NYS) of Peck’s unpublished
variety which may well be an autonomous species or a geographic race
(subspecies) of S. Airtellus. If recognized as a species, it should be known
under the specific epithet used by Kauffman, since the figures in Krombholz
do not provide essential characters and do not give analyses as would be
necessary to really understand this species. Art. 44 (3) of the International
Rules is therefore not applicable in this case, and is, for that matter, very
rarely applicable at all in groups where the color of the spores and their
shape, the presence or absence of clamp-connections, the structure of the
trama, the viscidity of the pileus or stipe etc. are essential characters while
the characters shown in these early pictures are “essential” (if the meaning
of the word is stressed) only if the painting is executed and reproduced so
well as to leave no doubt about the identity of the fungus. This is not the
case with B. tomentosus Krombh. The description given by that author
later under another name, combines two tentative ‘new species” and is,
consequently, useless as a descriptive basis for B. tomentosus. B. tomen-
tosus Raddi is pre-Friesian, and was validated by Saccardo four years after
Kauffman. The only way of ruling out Kauffman’s species is by proving it
to be a synonym (of S. Hirtellus or a subdivision of it, or perhaps of
Rostkovites californica), not a homonym.

Suillus cembrae (Sing.) Sing. comb, nov.
Ixocomus cembrae Sing., Rev. de Mycol. 3:49. 1938.

Described from the Altai Mts. in Siberia, this is an independent species,
close to S. punctipes (Peck) Sing.

Suillus punctipes (Peck) Sing. comb. nov.

Boletus punctipes Ann. Rep. N.Y. State Mus. 32:32. 1880.
Ixocomus punctipes Sing., Ann. Mycol. 40:30, 1942.

The data obtained through study of fresh and dried material in New
York State show that this species, observed under Pinus strobus by this

278 FARLOwIA, VoL. 2, 1945

author, has white mycelium (not pink as S. cembrae), and that the context
remains unchanged when broken. This, coupled with the difference in the
mycorrhiza-host, indicates that these specimens are different. The chemical
reactions of the New York material were: KOH on surface of pileus,
brown; on context of pileus, immediately sordid lilac, then lilaceous brown:
NH; on surface of context of all parts, negative; NH,OH on surface of
pileus, negative; on context of pileus, slowly pale and sordid lilaceous;
FeSO, on context of pileus, pale gray.— The spores are subfusoid-sub-
elliptical in frontal view, subfusoid in profile, melleous, smooth, (7.8)—8.8—
11.5 x 3-3.7 w; basidia 20-28 x 6.5-7 y, 4-spored; cystidia broadly clavate
to mostly elongate-clavate, or fusoid, either non-incrusted or more or less
strongly incrusted, very numerous, especially near the pores, the walls
moderately (0.5 ») thick, 30-70 x 4.5-10.5 , the incrusted ones more often
clavate than fusoid and the non-incrusted ones more often fusoid. — This
species is distributed in the northeastern States reaching south as far as
North Carolina with the western limits unknown.

Sect. Bovini Sing., Rev. de Mycol. 3:38. 1938.

Characters of the section: see key, p. 259. Type species: S. bovinus
(L. ex Fr.) Kuntze, Rev. Gen. Plant. 32:535. 1898 belonging in subsect.
Euryporini Sing. l.c., p. 53. Another species, S. variegatus (Sw. ex Fr.)
Kuntze, /.c. is the type of subsection Stenosporini Sing. l.c. None of these
occurs in Florida, and even their occurrence in North America is, though
claimed, not quite certain.

Sect. Piperati Sing., Rev. de Mycol. 3:38. 1938.

This differs from the preceding section mainly in having dull red or pink
tubes. The viscidity of the pileus is not always as strongly developed as in
the other sections, but the hyphae of the cuticle are remarkably loosely
arranged and imbedded in a gelatinous mass. While in other sections the
color of the mycelium usually ranges from white to dirty drab or from
white to pink, it tends to yellow in the Piperati. Two species are known,
the type species S. piperatus (Bull. ex Fr.) Kuntze, Rev. Gen. Plant. 32:535.
1898 with its variety amarellus (Quél.) Sing. and S. rubinellus (Peck) Sing.
None of these has ever been found in Florida.

SPECIES INCOMPLETELY KNOWN

Rostkovites californica Murr., Boletinus flavoluteus Snell, B. glandulosus
Peck are some species apparently belonging in Swidlus but not well enough
known to this author. They have no bearing on Florida species, nor, like
any other species that may enter into this category, do they have outstand-
ing taxonomic significance.

SPECIES EXCLUDENDAE

Suillus castaneus (Bull. ex Fr.) Poir. in Lam. ex Karst.
Suillus cyanescens (Bull. ex Fr.) Poir. in Lam. ex Karst.

SINGER: FLORIDA BOLETINEAE IIT. 279

Suillus subalbellus (Murr.) Sacc. & Trott.

Suillus atroviolaceus Hoehn.
All these species belong in Gyroporus, not in Suillus S. F. Gray.
Suillus jamaicensis (Murr.) Sacc. & Trott. is not a Suillus in our sense, nor is it a
Gyroporus, cf. p. 240.
Suillus hygrophanus Rostr. can be almost anything. No specimens seen.
Suillus cantharelloides (Jacobasch) Sacc. & Syd. is not a bolete.

Suillus Maxonii (Murr.) Sacc. & Syd.

This is a polypore, and, as the type (NY) shows, belongs in the genus
Polyporus sensu str. The spores are hyaline, thin-walled, cylindric or
cylindric-fusoid, 11-12 x 5 »; hyphae of the tube-trama interwoven-sub-
parallel, trama not bilateral but subregular, walls thin, clamp connections
numerous. It is described from Costa Rica, and should be known as
Polyporus Maxonii (Murr.) Sing. comb. nov.

Kuntze, Revisio Generum Plantarum 3°:535-536. 1898, amused him-
self by transferring practically all known species of Boletus which he copied
from Saccardo, to Suillus because of priority or what he used to call so.
Personally, he has never seen any of them except perhaps Boletus edulis
in his kitchen.

Subfam. Xerocomoideae Sing. subfam. nov.

Hyphis haud fibuligeris vel raro paucas fibulas gerentibus; sporis fusoideo-subcylin-
draceis, olivaceo-brunneis in cumulo (numquam roseolis nec flavis nec ferrugineis) ;
pileo subtomentoso vel tomentoso jove sicco, saepe viscidulo jove pluvio; hymenophoro
lamelloso interstitiis venosis vel tubuloso et tunc poris plerumque amplis, irregularibus,
compositis praedito, sed nunquam typice boletinoideo, adnato vel decurrente; cystidiis
hyalinis vel melleo-incrustatis vel melleo-tinctis; tramate hymenophorali subbilaterali-
subregulari (typi “phylloporoidei”) strato laterali vix laxiore quam mediostratum
leniterque tantum divergente; stipite solido, cylindraceo vel subcylindraceo, rarius
ventricoso-subbulboso, glandulis veloque carente. Habitatio: Ad terram et ad lignum
sub variis arboribus.

Characters of the subfamily: see key p. 227 and the above Latin diag-
nosis. Type genus: Xerocomus Quél.

Phylloporus Quél., Flor. Mycol., p. 409. 1888.

This genus has all the characters of the first section of the following
genus of this subfamily except for the different configuration of the hy-
menophore. This is the only genus of the Boletaceae with gills instead of
tubes. It can, however, be distinguished from all gill fungi by its char-
acteristic bright and deep blue to green-blue reaction with ammonia on the
surface of the pileus and the structure of the young gill trama. The hyphae
are usually clampless but in some specimens some rare clamps can be ob-
served if carefully and patiently searched for. The type species is P. Pel-
letieri (Lév. apud Cr.) Quél. which in its turn is the same as P. rhodoxanthus
(Schw.) Bres. ssp. europaeus Sing. Another subspecies of this species oc-
curs in Florida.

280 FARLowIA, Vor. 2, 1945
11. Phylloporus rhodoxanthus (Schw.) Bres., Fung. Trid. 2:95. 1900.

Paxillus flavidus Berk., Lond. Jour. Bot. 6:315. 1847.

Paxillus pinguis Hook. fil. apud Berk., Hook. Journ. Bot. 3:41. 1851.

Agaricus Pelletieri Lév. apud Crouan, Flor. d. Finistére, p. 81. 1867. (=ssp. euro-

paeus Sing.)

Agaricus paradoxus Kalchbr., Icon. Fung., p. 27. 1873 (=ssp. europaeus Sing.)

Paxillus paradoxus Cooke, Grevillea 5:6. 1876 (=ssp. europaeus Sing.).

Clitocybe Pelletieri Gill., Champ. Fr., p. 170. 1878 (=ssp. europaeus Sing.)

Gomphidius rhodoxanthus Sacc., Syll. 511139. 1887.

Flammula rhodoxanthus (sic) Lloyd, Myc. Not. 1(no. 3) :17. 1899.

Paxillus sulcatus Pat., Bull. Soc. Myc. Fr. 5:7. 1909 (=ssp. americanus Sing. ?).

Paxillus rhodoxanthus Ricken, Blatterp. 1:95. 1911 (formally ssp. americanus)

Phylloporus bogoriensis Hoehn., Fragm. Mycol. 16(838) :41. 1914 (=ssp. bogori-

ensis Sing.)

Paxillus Pelletieri Velen., Cesk. Houby 2:356. 1920 (=ssp. europaeus Sing.)

Phylloporus sulcatus Gilbert, Bolets, p. 88. 1931 (=ssp. americanus Sing.?)

Phylloporus paradoxus Cleland, Toadst. & Mushr. 1:178. 1934 (formally = ssp.

europaeus)

Gomphidius foliiporus Murr., Mycologia 35:432. 1943 (=ssp. foliiporus Sing.)
?Agaricus Tammii Fr., Ofvers. K. Vetensk.-Akad. Forh. 18(1):24. 1861.
?Gymnocybe Tammi Karst., Hattsw., p. 412. 1879.

?Paxillus Tammii Pat., Tab. Anal. Fasc. 4:161. 1885 (descr. et fig. excl. quae=P.
rhodoxanthus ssp. europaeus)
?Flammula Tammii Sacc., Syll. 5:810. 1887.

PrateE I, ric. 11-12.
Ssp. foliiporus Murr. l.c.

Pileus when fresh between “maroon” and “claret brown,” or between
“russet” and “hazel,” or “russet,” or “ochraceous tawny,” or “buckthorn
brown,” when old (sooner in sunny weather) fading to between “buckthorn
brown” and “Isabella color” or to between “Isabella color” and “chamois,”
non-viscid, subtomentose or tomentose, in many specimens with soft wart-
like projections, glabrescent with age, and often becoming rimulose or
rimose (mostly concentrically), with the margin forming an obtuse angle
(about 90°) in old specimens, convex, soon with flat or even slightly de-
pressed center, margin eventually also flattening, 18-145 mm. broad. —
Hymenophore “strontian yellow” or between this and “lemon yellow”,
turning blue in fresh material where wounded, eventually more sordid-
brownish, decurrent, lamellae broad (7-14 mm. broad), their edges never
forming pores and only the bases (near the line of attachment to the con-
text of the pileus) up to one third of the breadth of the lamellae, rarely
higher up, connected by transversely or obtusely forking veins, the sides of
the lamellae often transversely veined, the edges entire and somewhat
darkening in age; spore print brownish olivaceous, becoming “chipmunk”
(M. & P.) in older herbarium preparations. — Stipe reddish brown above
(colors much like those of the pileus), at least the furfuraceous squamulae
or farinaceous pruinosity of the apex, and also the thin ribs continuing the
lamellae, besides with a belt of reddish brown below the apex in most
specimens, otherwise more olivaceous below, or yellow (concolorous with

SINGER: FLORIDA BOLETINEAE II. 281

the hymenophore) above, the reddish-brown colors eventually gradually dis-
appearing, grayish greenish pallid below from a tomentose-flocculose cover-
ing, or, if glabrous, merely pallid there, not viscid, solid, equal or tapering
downward, 16-55 x 2-40 mm.; mycelium either yellowish white or rich
yellow; veil none. — Context yellowish white, pale yellow, or yellow, bluing
when quite fresh and young; odor faint as in Xerocomus subtomentosus,
X, illudens etc.; taste mild.

Spores 11-15 x 4.5—5.8 », melleous, smooth, oblong-ellipsoid to subfusoid,
often the upper half only tapering to the apex; basidia 29-42 x 8-14.5 yp,
mostly 4-spored, only occasional 3-spored ones mixed in, which measure
31-36 x 7.5-8.5 , and very rare 2-spored ones exceptionally found (e.g.
43x 6.5 w); cystidia 51-95 x 7-20 p», fusoid, more rarely clavate, rarely
cylindric, apiculate but obtuse at the tip, hyaline or with a melleous apex,
rarely entirely melleous, and very slightly so in most cases, numerous,
especially near the edges, the larger ones mostly found near the edges;
trama consisting of a mediostratum of subparallel-subinterwoven hyphae
that are more irregularly interwoven near the base of the lamella, and a
lateral stratum of slightly though distinctly divergent (parallel with each
other) hyphae which are not or not much more loosely arranged than the
hyphae of the mediostratum and consequently also not or not much paler
than these; kyphae in most carpophores without clamp connections, but
occasional carpophores with a few clamped septa are observed.

Chemical reactions: KOH on surface of pileus, deep rich blue; on la-
mellae, brown; on context brownish.— NH, and NH,OH on not yet
discolored surface of pileus, quickly bright but intensely pure blue to
green-blue, eventually reaching ‘““Hay’s brown” as everywhere else in the
carpophore. — Aniline negative. — Formol negative. — Methylparami-
dophenol negative.

Habitat: On the ground in hammocks as well as in open woods, also on
trunks of trees (Sabal, Quercus, Pinus) in frondose woods, mostly near
Quercus spec., and also in coniferous woods, mostly near Pinus australis,
solitary or gregarious, fruiting from May to November.

Distribution: Florida.

Material studied: Fla., Highlands Co., Highlands Hammock State Park,
R. Singer, F 223/I and F 223/I a (FH), F 632 (FH); Sebring, R. Singer,
F 439 (FH); Polk Co., Lake Alfred, A. S. Rhoads (det. Murrill) (FLAS) ;
Alachua Co., Gainesville, R. Singer, F 2183 (FH); F 2629 (FH); W. A.
Murrill, (authentic) F 17990 (FLAS); Sugarfoot Hammock, E. West,
L. Arnold, W. A. Murrill, F 17747 (type) (FLAS, FH); F 17774 (co-
type) (FLAS); F 15728 (FLAS); F 18157 a young stage, authentic
F 18157 (FLAS); Magnesia Springs, £. West, L. Arnold, W. A. Murrill,
(authentic) F 17991 (FLAS); Columbia Co., High Springs, W. A. Mur-
rill, (authentic) F 15599 (FLAS); Dade Co., Brickell’s Hammock, Miami,
R. Singer, F 940 (FH).

P. rhodoxanthus ssp. foliiporus (Murr.) Sing. is found in two forms, one
with yellow mycelium, and another one with whitish mycelium. It is pos-

282 FARLOWIA, VoL. 2, 1945

sible that these are hereditary forms which however would be of negligible
taxonomic importance unless coupled with some other character. We have
not been able, thus far, to ascertain any correlation of this kind.

When reading the diagnosis of Gomphidius foliiporus Murr., one may be
impressed by the allegedly pallid lamellae and the dark spores. The speci-
mens, however, including the type, do not show any appreciable differences,
either macroscopical or microscopical, from the specimens determined
by Murrill as PAylloporus rhodoxanthus. In fact, the latter as well as
G. folitporus are identical. The common Phylloporus in and around Gaines-
ville, Fla., always had yellow lamellae when young and fresh, never pallid
ones, and the spores were of the ordinary color in the Xerocomoideae in
print as well as under the microscope. Fries indicates spores of a different
color for his Agaricus Tammii, but this should not have influenced the
descriptions of P. rhodoxanthus because Fries’ indication is either an error,
or A. Tammi is not a PAhylloporus but rather a Gymnocybe.

This does not mean that G. foliiporus is a plain synonym of the northern
P. rhodoxanthus. It is rather a geographic race of the latter. The difference
between this and the other races of P. rhodoxanthus does not consist in the
characters emphasized by Murrill but in the characters opposed to each
other in the following key.

KEY TO THE SUBSPECIES OF PHYLLOPORUS RHODOXANTHUS

A. Context changing to blue (green), or “black” by autoxydation when wounded, at
least lamellae bluing or “blackening” when quite young and fresh; tropical and
subtropical races.

B. Spores 11-15 x 4.5-5.8 «; American race. ssp. foliiporus
B. Spores 7.5-11 x (3.5)-4-5 uw; East Indian race. ssp. bogoriensis

A. Context unchanging or changing to reddish or brownish red when wounded; lamel-

lae never bluing or blackening even when young and fresh. Temperate races.

C. Anastomoses of the lamellae constant and broad (high), reaching half the
breadth of the lamellae, or more, more rarely less; decurrent ribs at the apex
of the stipe forming a proliferation of the lamellae, either absent, or rather
indistinct, even if mycelium yellow. ssp. europaeus

C. Anastomoses of the lamellae mostly vein-like, rarely higher and then only
toward the margin of the pileus, or near the stipe, and even there not sur-
passing half the breadth of the lamellae, sometimes entirely lacking; from
the point of attachment of each lamella at the stipe, a longitudinal vein or
rib runs down the apex of the stipe in all forms with yellow mycelium;
American race. ssp. americanus

Ssp. bogoriensis (Hoehn. l.c.) Sing. comb. nov.

The examination of the type material (FH) makes it evident that this
species is another race of P. rkodoxanthus. Aside from the somewhat
problematic difference between bluing and blackening, we find the spores
slightly smaller in all three collections from Java than they are in our
Florida material; the basidia are about 35x 10.5 ,, 4-spored; cystidia
either hyaline or brownish, cylindric, clavate, or fusoid, with 0.8 » thick

SINGER: FLORIDA BOLETINEAE II. 283

walls, bluntly rounded at apex, numerous, especially near the edge,
44-105 x 11-20 p; tramal structure as in ssp. foliporus; no clamps seen.

Assuming as wide a variation of characters as is found in the Florida
subspecies, it may be concluded that all three collections of Hoehnel’s are
the same thing. Even the material marked “Tjibodas” and “Zwergexem-
plare” is hardly different though it shows some similarities with material
collected in Indo-China and China which rather belongs in ssp. americanus.
Specimens without notes on color changes from the Singapore Botanic
Garden were collected by Baker and remained undetermined in Patouil-
lard’s Herbarium (FH). I think they belong to ssp. bogoriensis.

Ssp. americanus Sing., Rev. de Mycol. 3:171. 1938.

Lloyd has first mentioned the difference in the hymenophore of the
American form, the type form of P. rhodoxanthus, and the European form.
The specimens we have collected in this country and the specimens pre-
served in the herbaria, as far as they have been found in North America,
can be divided in two basic forms, one with whitish mycelium, and wanting
ribs on the apex of the stipe, and one, usually larger, with yellow mycelium,
and very distinct longitudinal veins or ribs on the apex of the stipe. The
form with whitish mycelium has been found in New Brunswick, Canada,
and in northern New England, while the form with yellow mycelium is
known from southern New England to the Carolinas and perhaps Georgia.
The type of P. rhodoxanthus ssp. americanus Sing. is, of course, the speci-
men on which Schweinitz based his Agaricus rhodoxanthus, and this is
obviously the form with yellow mycelium. It so appears that the form with
white mycelium does not belong in the same subspecies with the typical
ssp. americanus. However, more data are needed in order to make sure
that this actually is another geographical race, and besides, it will have to
be proved that it is not identical with any European form.

The spores are rather variable in both American forms, in the white-
mycelioid form varying from 10-11 x 3.3-4 » in New Hampshire material
to 11.5-13.5 x 44.8 » in New Brunswick material; in the yellow-mycelioid
form they usually are 10.5-12.5—(14.3) x 4—5.5 ; basidia 20-30 x 6—8.3 yp,
with four 5 » long sterigmata; cystidia hyaline or in old herbarium speci-
mens with a slight yellowish tinge, cylindric or cylindric-fusoid, also some-
times clavate with 0.5—-0.8 » thick wall, rather numerous to very numerous,
50-111 x 6-19 y»; trama as in the other subspecies; no clamps found.

Material from eastern Asia seems to belong here. The description given
by Imai for Hokkaido material fits well. We have also examined Chinese
collections with the spores 9.2-15 x 3.7-5.5 ys; basidia 29-32 x 7-9 yp, 4-
spored; cystidia hyaline to very pale yellowish with 0.7 » thick wall, clavate
or fusoid, also sometimes cylindric, 40-110 x 7-19 »; trama as in the other
subspecies; clamps none; color of the mycelium yellow; ribs at apex of
stipe distinct. Patouillard’s type (FH) of his Paxdllus sulcatus from Indo-
China has 11-12.8 x 4.5-5—(6) » large spores (a few foreign spores from
Boletellus, probably B. emodensis, have fallen onto the hymenophore of the

284 FARLOWIA, VOL. 2, 1945

type specimen); basidia and cystidia as in the yellow-mycelioid form of
ssp. americanus.

Lebedeva claims that she had P. rhodoxanthus from the Far-Eastern
Province of the U.S.S.R. Thus this race probably occurs all through East-
ern Asia from there through Japan and Eastern China to French-Indo-
China.

Ssp. europaeus Sing., Rev. de Mycol. 3:171. 1938.

Good European material from St. Johann in Tyrol, v. Hoehnel, 1908
(FH) has recently been consulted by this author and the mycelium has
been found to be yellow while the apical ribs were absent or very indistinct;
the spores were 11-14 x (3.3)-4.2-5 1; basidia 35-44x 8.5 pw, 4-spored;
cystidia yellowish-brownish to hyaline, 45-70 x (5.5)—10 y, fusoid; trama
not studied, clamps none.

SPECIES INCOMPLETELY KNOWN

Phylloporus infundibuliformis (Cleland) Sing. comb. nov,
Paxillus infundibuliformis Cleland, Toadst. & Mushr. 1:175. 1934.

This is evidently a Phylloporus but not conspecific with P. rhodoxanthus,
as far as can be concluded from the description. What is described as
P. paradoxus by Cleland is a subspecies of P. rhodoxanthus, probably an
undescribed one, or else ssp. americanus or ssp. bogoriensis.

SPECIES EXCLUDENDAE

Phylloporus malaccensis Pat. & Baker is Porphyrellus malaccensis (Pat. & Bak.) Sing.
Phylloporus intermedius Pat. is Gyrodon intermedius (Pat.) Sing.
Phylloporus Rompelii Pat. & Rick.

This species is Gyrodon Rompelii (Pat. & Rick) Sing., and perhaps a
variety of G. merulioides (Schw.) Sing.

Phylloporus lariceti Sing.
This species is here treated as the type of the new genus Psiloboletinus.
Phylloporus squarrosoides (Snell & Dick) Sing.

This species may very well be treated as a Phylloporus, but in the present
arrangement it is transferred to Xerocomus.

Xerocomus Queél., Flor. Mycol., p. 417. 1888.
Xerocomopsis Reichert, Palest. Journ. Bot., Reh. Ser., 3:229. 1940.
Pileus more or less tomentose or subtomentose, frequently with at least
fragmentary trichodermium-pallisade; hymenophore not lamellate but oc-
casionally subboletinoid, most frequently with rather wide and angular
pores, adnate, often with a decurrent tooth, or arcuate-decurrent, more
rarely becoming depressed around the stipe and then usually with the
radial walls of the tubes forming a sublamellate ring around the apex of

SINGER: FLorIDA BOLETINEAE II. 285

the stipe, not free, but sometimes separating in age; stipe usually cylindric
or subequal and rather thin, more rarely some other shape, and very rarely
ventricose-bulbous as in Boletus, sometimes with an ocher brown or yellow
coarse network at the apex, never finely reticulate as in Boletus; veil none;
yellow pulverulence none; spores variable in size but constant in shape, 7.e.
always subcylindric to subfusoid, or oblong-ellipsoid to ellipsoid-subclavate,
always olivaceous-brown in print (never yellow, ferruginous, pinkish-
vinaceous etc.) ; trama of the hymenophore subbilateral of the Phylloporus-
type, i.e. the lateral stratum scarcely (if at all) lighter colored, and scarcely
(if at all) looser than the mediostratum, its hyphae though distinctly di-
verging in youth, not strongly bent outward, and not distinctly imbedded
in a mucilaginous mass, parallel with each other; hyphae without clamp
connections; cystidia medium-sized to rather large, not strikingly incrusted
or not incrusted at all, not in fascicles; setae none; mycelium white or
yellow, not constantly forming mycorrhiza with conifers (except a few
species which do not occur in Florida). — The type species is Xerocomus
subtomentosus (L. ex Fr.) Queél.!®

The genus Xerocomus is the one genus among the Boletineae that is most
difficult to delimit, and yet it appears to be indispensable from a practical
as well as theoretical point of view. It undoubtedly represents itself as a
link between Phylloporus on one side and Boletus sensu str. on the other.
As for its limits with PAylloporus, I have been tempted to redefine Phyl-
loporus by introducing into that genus species with definite pores such as
Xerocomus illudens and the related species showing the characteristic vivid
blue ammonia-reaction on the surface of the pileus known also in Phyllo-
porus. In fact these species are so closely related to Phylloporus that they
almost form a continuous line, and the dividing line between PAylloporus
and Xerocomus could as well have been drawn along chemical as along
macro-morphological characters. Unfortunately these convenient and rea-
sonably sharp characters are not entirely parallel, since the blue ammonia-
reaction is found on some porous species. I finally chose the macro-mor-
phological criterion for the distinction of the two genera for two practical
reasons. First, this character has in its favor tradition as well as the greater
convenience of immediate recognition in old, dry, or dried material. Second,
a modification of the vivid blue reaction which is found in other species of
Xerocomus either as a flash of black-blue or a livid-caesious ring around
the stained spot, or a violet shade of blue, may cause confusion notwith-
standing their obvious difference from the coloration observed in Phyllo-
porus. This more conservative solution, leaving X. illudens and allied forms

18 Tt would mean to reduce Xerocomus to the status of a synonym of Boletus, if the
first species mentioned by Quélet, Xerocomus impolitus were to be the type of the genus,
as has been proposed by others. This latter proposal, however, was made as a conse-
quence of a misinterpretation of the rules which do not prescribe the choice of the first-
described species. Xerocomus, as conceived by its author, is the group centering around
X. subtomentosus, and therefore this author in 1936 proposed this latter species as
lecto-type.

286 FartowlA, VoL. 2, 1945

in Xerocomus, has also the advantage of keeping within a single genus such
species as X. illudens, X. spadiceus sensu Quél.=X. coniferarum and X.
subtomentosus which were in the past even considered to be varieties of one
and the same species. I may add, however, that there is not enough evidence
to prove that X. illudens and X. coniferarum actually are more closely re-
lated to the rest of the Xerocomi than with Phylloporus.

On the other extreme of the genus Xerocomus are forms which apparently
gradually approach the genus Boletus sensu str. In these, the pores are
somewhat smaller than in the others, and the structure of the lateral
stratum of the hymenophoral trama becomes more truly bilateral (Boletus-
type). In fact, even macroscopically, no sharp line between these two
genera can be drawn. If, however, the exact structure of the trama and the
affinities of every single intermediate species are checked and weighed, a
line between these genera will result that is somewhat hard to define but
evidently leaves related forms together and makes both genera as natural
as possible.

Two important papers on the anatomy of the hymenophoral trama in
the boletes have appeared recently. The first authors to mention the dif-
ference between what we call the Phylloporus-type and the Boletus-type,
were Lohwag and Peringer (Ann. Mycol, 35:295-338. 1937). Their results
were checked by this author on the limited amount of material in 1938
and found to be correct. I may add that in my key to the genera of the
Agaricales (Schweizer. Zeitschr. Pilzk. 17:7. 1939) I have for the first time
used the two types of bilaterality for taxonomic purposes. In a later paper,
Elrod and Blanchard (Mycologia 31:704. 1939) also published on the
structure of the trama of the Boletaceae but did not arrive at the same
results as their predecessors. In fact, they concluded that though there was
plenty of difference in different species, this difference did not coincide with
any classification based on other characters and can therefore not be used
for taxonomic purposes. I have regretted that these authors did not publish
precisely what structure they found in what species and at what stage. If
they had, it would probably become more apparent that the differences they
found cannot, in many cases, be used for taxonomic purposes because they
do not exist between species but between stages. I have now checked on
young and good material of most of the specimens available for this purpose
(see descriptions and notes on extralimital material), and rechecked on my
own findings in long and tedious investigations. If care is taken to use for
these investigations nothing but comparatively recent collections and car-
pophores just beginning to sporulate, it will be found that there is a bi-
lateral trama in all Boletaceae with a single modification, that is the
existence of the Phyiloporus- and Boletus-type of bilaterality. As soon as
the tubes begin to elongate, the divergence of the hyphae of the lateral
stratum diminishes slowly until it finally reaches zero i.e. the hyphae show
a generally axillar direction, parallel with each other and with the medio-
stratum. On the other hand, there is no bolete among the scores of speci-
mens I have sectioned that does not show distinctly (though to a different

SINGER: FLoRIDA BOLETINEAE II. 287

degree) divergent hyphae in the lateral stratum at a certain age. Even the
difference between the Phylloporus- and Boletus-type is rather a quantita-
tive than a qualitative one. It is only natural that, under these circum-
stances, intermediate structures occasionally may be observed. An example
is section Pseudoboleti with thus far one species, X. badius. Since the dis-
posal of the species with intermediate structure has been taken care of in
an arbitrary way, with the idea of keeping related forms together, the keys
to the species of Boletus (now in preparation) take into account both
species here considered as belonging to Xerocomus, and species considered
as Boletus (sensu str.).

KEY TO THE SECTIONS

A. Spores short (Q=2, or less), or reaction of the surface of the pileus with ammonia
an intense, vivid pure blue or green-blue (about “porcelain blue” or “dusky green
blue”), not blackish blue, nor violet, livid, or caesious. Sect. 1. Pseudophyllopori

A. Spores elongate and reaction with ammonia not as described above.

B. Pileus not or scarcely viscid in wet weather; species growing entirely as
saprophytes, or with northern forest trees, or parasitic on fungi.
C. Not parasitic on fungi; spore production of the hymenophore abun-
dant; context potentially bluing, or at least fresh tubes sometimes
bluing. Sect. 2. Subtomentosi

C. Parasitic on fungi (Sclerodermatineae) ; spore production not as active
as in other boletes; context not or scarcely bluing, not even in the
tubes. Sect. 3. Parasitici

B. Pileus distinctly viscid in wet weather; either forming mycorrhiza with
northern conifers exclusively, or connected with southern hardwoods, some-
times in open places.

D. Carpophores small; spores smaller than 10 «; context and pores un-
changing; species never connected with conifer mycorrhiza.
Sect. 4. Brasilienses

D. Carpophores medium to rather large with medium (longer than 10 y)
to rather large spores; context bluing; northern species, always con-
nected with conifer mycorrhiza. Sect. 5. Pseudoboleti

Sect. Pseudophyllopori Sing. sect. nov.

Sporis breviusculis (quotiente e longitudine et latitudine vix superante 2) aut reac-
tione NH; et NH,OH in superficie pilei provocata intense azurea vel viridulo-azurea.

Characters of the section: Pileus not viscid; surface of pileus in the
species studied turning immediately to an intense, vivid pure blue or green-
blue (about ‘“‘porcelain blue” or “dusky green blue”), at least when young
and fresh, neither water-soaked nor too dry; pores wide; spores either short
(Q=2, or less), or of normal elongated shape; context bluing or not bluing;
habitat on the ground or on wood in frondose and coniferous woods, ham-
mocks, and in gardens near trees, never parasitic on fungi.’* — The type
species is X. illudens (Peck) Sing. (Boletus illudens Peck).

14 Tt is not known whether these species form mycorrhiza. If they do, they evidently
prefer frondose trees, especially oaks, hornbeam, and perhaps sweet gum and others.
However, one species grows with conifers (X. coniferarum).

288

Fartowia, VoL. 2, 1945

KEY TO THE SPECIES

A. Spores with Q averaging around 2 or somewhat smaller.

B. Pores large, subboletinoid, angular; spores 9-12 x 4.5-6 uw. X. squarrosoides

B. Pores medium-sized, circular to oblong; spores 7.7-8.5 x 5 yu.

X. Housei

A. Spores with Q decidedly above 2.

C. Stipe without a network in all specimens of a given group, 7.e. constantly
smooth; spores 4-5 » broad or broader; growing with oaks (in Florida).

D. Base of stipe distinctly bright yellow; mycelium yellow; carpophores

small and thin. 12. X. hypoxanthus

D. Base of stipe not distinctly vellow; mycelium yellowish white to

white; carpophore large and thick (in Florida).
E. Floridan species; carpophores large, thick, growing in scrub
and in low oak woods; hymenophore a rich yellow
13. X. pseudoboletinus
E. African species; carpophores rather small and rather thin, grow-
ing at the base of termite nests; hymenophore pinkish cinnamon.
X. Linderi

C. Stipe with a network, at least at the apex in most specimens of a group
(sometimes the network is incomplete, and replaced by a coarse brown
punctation on yellowish ground, but the majority of carpophores shows them
connected enough to form a well-distinguishable network); spores either
about 4-5 w broad, or narrower.

F. Pileus usually tapering down into the thin, frequently curved stipe,

and thus the hymenophore distinctly arcuate-decurrent; tubes very
gradually transient into the coarse network (ridges or punctation) of
the stipe; mycelium white or yellowish white; spores 4-5 w broad;
habitat around trunks in dense hammocks, more rarely farther away
from the trunks on humus. 14. X. hemixanthus

. Pileus not shaped as described above, and consequently the hymeno-

phore either flatly adnate (though sometimes plainly decurrent on one
side because of irregularity of growth), and only slightly decurrent
with a tooth, the walls proliferating into the network of the apex of
the stipe but not very gradually so; mycelium yellowish white or yel-
low; spores 3.5—4.2 uw broad in eastern species, often broader in others;
growing in small groups not arranged around the trunks of trees,
usually in rather open stands of northern woods, or in partly cultivated
hammock land, in gardens etc.

G. In beech woods (Fagus) and in coniferous woods, in Europe,
perhaps in North America; spores (3.5)—4-5-(6) w broad,
most frequent sizes 11.5-13 x 4.8 4; mycelium yellow and then
carpophores large and the stipe not tapering downwards, or
mycelium white and then carpophores medium-sized to rather
small with the apex of the stipe deeply reticulate.

H. Stipe brown, very strongly reticulated, pallid near the
base, equal or tapering downward; carpophores medium-
sized, or perhaps rather small; in beech woods (accord-
ing to Rostkovius). X. lanatus

H. Stipe yellow with more or less deeply decurrent brown
network; the base and the mycelium yellow, stipe sub-
equal or tapering upward; large plant in coniferous
woods. X. coniferarum

SINGER: FLORIDA BOLETINEAE II]. 289

G. In mixed oak woods and on lawns and in gardens under fron-
dose trees other than beech, mostly oak in the eastern part
of North America; spores 3.5-4.5 « broad, most frequently
10-11.7 x 3.5-4.1 yw; mycelium yellow and then carpophores
medium-sized and the stipe distinctly tapering downward in
most cases, or mycelium white or yellowish white, and then
carpophores large with the stipe yellow and brown-reticulate.

15. X.illudens

I. Mycelium white to yellowish white; carpophores large
(pileus 63-123 mm. broad; stipe 40-60x 25-35 mm.) ;
the brown network of the stipe usually reaching deep to-
ward the base. Northern plant. ssp. typicus

I. Mycelium yellow; medium-sized carpophores (pileus 43-
87 mm. broad; stipe 35-70x 8-18 mm.); the brown
network of the stipe usually confined to the upper portion
of the stipe. Known only from Florida.

ssp. xanthomycelinus

12. Xerocomus hypoxanthus Sing. spec. nov.

Pileo dilute brunneolo, ochraceo-brunneo-punctatulo (velutino-punctatulo vel fibril-
loso-granulosulo) vel tomentoso, sicco, convexo, 26-66 mm. lato, ammonici ope vivide
azureo; tubulis flavis, deinde subolivaceis, saepe tactu subvirescentibus, adnatis vel
subdepressis; sporis 11-14 x 4.2-5.2 wu, brunneo-olivascentibus in cumulo recente; stipite
flavo, deorsum pallido vel pallide flavo, sed brunneolo-granuloso-subsquamuloso,
squamulis minutis subsparsis obtecto, aequali, mycelio flavo, 30-50x 4-10 mm.; carne
alba vel albida, flavidula vel flava, laesa subtiliter caerulescente in pileo, inodora, miti.
Habitatio: In dumetis subtropicis (“low hammock”) et silvis arenosis (“scrub”) prope
quercus et arbores alias frondosas aestate in Florida Americae Borealis.

Pileus pale brownish but appearing much more vivid brown because of
a dense fibrillose-granular velvet or a tomentum that is ‘mahogany red,”
“Brussels brown,” “russet,” “Mars brown,” or “ochraceous tawny,” and
by side view (seen at an acute angle) usually rather “buckthorn brown,”
non-viscid, only subviscid when wetted thoroughly, not reddening in cracks,
26-28 mm. broad but occasionally reaching 66 mm. in diameter. — Hy-
menophore “strontian yellow” with a tinge of “oil yellow” or “pyrite yel-
low,” adnate-subdecurrent or planely adnate, or slightly depressed around
the stipe, or lamellately stretched near the stipe, and finally separating
from it, pores concolorous and greening on pressure or unchanging, wide;
spore print not obtained in thick enough layer but of about the same color
as in the following species. — Stipe pallid to pale yellow or pale tawny and
mostly more vivid yellow at the apex (“light cadmium” to “lemon chrome”’),
and usually likewise yellow at the very base where the finely fibrillose
bright yellow mycelium is attached, the yellow of the surface of the middle
portion usually obscured by the “Argus brown” granular punctation or
furfuraceous pustulation which becomes denser downwards but is usually
sparse and the particles minute farther above, though occasionally it forms
an annular belt of denser granulae (not a veil), non-viscid, solid, thin,
subequal or equal, 20-32 x 2-4.5 mm. but sometimes reaching 50 x 10 mm.
— Context white or with a very faint yellowish tinge in the pileus, reaching
“Martius yellow” to “strontian yellow,” pallid to light yellow in the stipe

290 FartowlA, VoL. 2, 1945

but often lemon yellow in the apex of the stipe and often marbled, the
context of the pileus never becoming reddish in wounds or cracks, unchang-
ing when bruised, or sometimes slightly bluing; taste mild; odor none or
very slight, like that of Boletus edulis; taste mild.

Spores (8.2)-11—14 x 4.2—5.2 , smooth, melleous, fusoid with suprahilar
depression; basidia 26-41.5 x 8.5-11 p, most frequently 26-36 x 9.5-10 p,
4-spored, very rarely some few basidia 2-spored; pseudoparaphyses clavate
with subcapitate apex, or basidiomorphous, e.g. 33 x (5)-8-11.5 p, not
always present; cystidia 34-70 x 6.8-12.2 p, fusoid with the thickest por-
tion in the middle or in the upper third, with the apex obtuse and often
ampullaceous protracted, hyaline, rather numerous; trama of the tube-walls
of the same general type as in X, illudens; all hyphae without clamp con-
nections.

Chemical reactions: KOH on pileus, dark and intense blue, in older and
drier material “amber brown” to “Argus brown’; on the tubes, light brown
to brown; on the squamulae of the stipe, deeper brown but ground color
unchanged. — NH,OH on the surface of the pileus, deep and rich blue to
green-blue when young and fresh, slightly bluish to negative when old or
dry; on context, blue to bluish when fresh.

Habitat: On trunks of palmetto between old petioles, on very decayed
frondose wood, on the base of frondose trees, or on humus and débris in
low and mesophytic hammocks, and on dry sand in scrub areas, from July
to September.

Distribution: From Georgia (and perhaps Virginia) south to Florida
(except in the tropical part of the state).

Material studied: Fla. Highlands Co., Highlands Hammock State Park,
R. Singer, F 626 (type, FH); F 37 (co-type, FH); F 357 (co-type, FH) ;
F 579/I (co-type, a large form of the scrub, FH); Alachua Co., Sugarfoot
Hammock, R. Singer, F 2473 6 (FH); F 2590/1 (FH).—Ga. Tallulah
Falls, A. B. Seymour (FH).

The combination of characters in this usually small bolete is constant in
all collections, and it does not seem to have been described before.

13. Xerocomus pseudoboletinus (Murr.) Sing. comb. nov.
Ceriomyces pseudoboletinus Murr., Lloydia 7:324. 1944.

Pileus “mahogany brown,” “Verona brown,” “russet,”’ later “sayal
brown,” “Mikado brown,” ‘ochraceous tawny,” or with some ‘“Sanford’s
brown,” or (because of the unequal development or partial lack of the
cuticle) as pale as “honey yellow” in the center and “chamois” to “cream
buff” on the margin, occasionally with some “pinkish buff” to “pinkish
cinnamon” mixed in, sometimes more “cameo brown” at places, tomentose
to subvelutinous, more tomentose toward the margin, partially and un-
equally glabrescent and often areolate-squamose, the squamulae appressed,
becoming more distinctly rimulose when old, the yellow of the flesh show-
ing through the crevasses or squamulae, dry, rugulose to smooth, margin
incurved and slightly projecting when young, the projecting portion pallid

SINGER: FLORIDA BOLETINEAE II. 291

underneath, pulvinate, then convex with an occasional depression in the
center, 40-170 mm. broad. — Hymenophore “lemon chrome” or between
‘“Jemon chrome” and “lemon yellow,” or “sulphine yellow,” becoming “‘sul-
phine yellow” or “lemon yellow” when older, then “citrine” with more
“lemon yellow” pores, adnate or shallowly depressed; tubes 10-17 mm.
long, the radial walls of the tubes often short-decurrent (1-6 mm.) and
occasionally with anastomosing lines; pores on unequal level, radially
elongate, at least near the stipe, 0.5-2.5 mm. wide, 3-5 pores to 5 mm.
radially, and 5—6 pores to 5 mm. transversely; spore print brownish olive.
— Stipe “naphthalene yellow,” “primulin yellow,” “strontian yellow,” or
‘ivory yellow” with “colonial buff” punctations, often at places almost
white, often concolorous with the pileus near the base, or near the base or
in the middle “onion skin pink,” “pecan brown,” or “walnut brown,” dry,
not reticulate, without furfuraceous or reticulate brown ornamentations,
even at the apex, but with occasional short longitudinal coarse ribs which
are subconcolorous with the pileus and rarely show a slight anastomosis,
otherwise quite smooth, but never really glabrous because of a minute,
mostly yellowish or pale tan punctation all over its surface, though often
seemingly subglabrous because the punctation does not show except when
observed with a lens, subequal or tapering downward and frequently with
a strongly enlarged apex, solid, 35-50 x 8-18 mm., the apex sometimes up
to 40 mm. wide. — Context “pale lemon yellow,” “lemon yellow,” or
“barium yellow” near the tubes and yellowish white to “naphthalene yel-
low” or almost white with yellow stripes in the stipe, otherwise yellowish
white to nearly white, often somewhat pinkish under the surface, either
entirely unchanging in any part, or more rarely slowly and weakly greening
in the deeper yellow portions of the context; odor of soap, or none; taste
mild, submild, or scarcely bitterish.

Spores 10.2-13.7 x 4-5.5 w, subfusoid or ellipsoid-oblong with suprahilar
depression, more rarely without it, smooth, melleous; basidia 22.5-43 x
8.8-11.7 (larger in the dry scrub areas than in moist hammocks), hyaline
to pale melleous, 4-spored; cystidia on pores and inside the tubes narrowly
fusoid or broadly fusoid with long cylindric neck which rarely is subacute,
and usually obtuse, hyaline to more rarely pale melleous, 37-56 x 4.5—-8 yp;
cheilocystidia sometimes differentiated, versiform; trama of the hymeno-
phore decidedly of the Phylloporus-type; hyphae without clamp connec-
tions, hyaline.

Chemical reactions: KOH on surface of pileus, “amber brown,” then
“Argus brown” or still deeper brown, later becoming paler; on context of
pileus, brownish; on context of stipe, negative or almost so; on tubes,
brownish; on surface of stipe, negative. — NH,OH (and also NH3 vapors)
on surface of pileus (if fresh enough), becoming “porcelain blue” to “dusky
green-blue,” less vivid on older caps, or in the old ones becoming “dull
Indian purple” or duller, or deep brown on old, dry pilei, then becoming
paler, the vapors usually reacting similarly’*; the portions treated with

* In strongly cracked or squamose pilei, the reaction is likely to be indistinct.

292 FartowiA, VoL. 2, 1945

FeSO, previously, and subsequently exposed to the action of ammonia
show the same blue reaction as fresh young material, even if the material
be rather old and dry (catalysis?) ; on context of stipe, negative except in
some cases where the deep yellow portions of the flesh change to bluish
green with ammonia. — FeSO, on surface of pileus, see under NH,OH;
on context of pileus, slowly dirty greenish. — H»SOy, on surface of pileus,
reddish fulvous; on tubes, rusty-ochraceous; on context of pileus, nega-
tive then sometimes blue. — HNO, on surface of pileus, reddish fulvous;
on the entire context, slightly reddish brown, generally causing a richer
and brighter color.— Formol on context of pileus, negative and later
somewhat blue.

Habitat: Under scrub oaks (Quercus Chapmanii and other species)
turkey oak (Q. laevis), water oak (Q. nigra), laurel oak (Q. laurifolia),
live oak (Q. virginiana), with Q. minima, in the scrub, low open places, on
lawns, and even in flatwoods, on dry sandy as well as on wet acid soil.
Fruiting from July to October.

Distribution: Florida (excepting the tropical part).

Material studied: Fla. Highlands Co., Highlands Hammock State Park,
in the scrub areas, R. Singer, F579 (FH); Alachua Co., near Dayville,
southwest of Gainesville, near the pond, R. Singer, F 2540, (FH): F 3029
(FH); between Gainesville and Newnan’s Lake in flatwoods, R. Singer,
F 2841 (FH); W. A. Murrill, F 19589, co-type, (FLAS, FH); Gainesville,
R. Singer, F 2764 (FH); also less well-preserved material without notes,
determined as B. illudens by W. A. Murrill (FLAS).

This species has obviously been considered as B. illudens by Murrill and
later has been described as a new species by him. It was already given as
a new species in this manuscript when a sample of C. pseudoboletinus
reached me.

14. Xerocomus hemixanthus Sing. spec. nov.

Pileo brunneo vel brunneo-umbrino, castaneo vel ferrugineo in junioribus, fulvo-
rubido vel ferruginascente in vetustis, tomentoso, pulvinato, dein applanato, 27-75 mm.
lato; ammoniaci ope caerulescente; tubulis flavis vel flavo-melleis, in vetustus discoloratis,
obtuse adnatis vel decurrentibus, plus minusve arcuatis, 5-11 mm. longis, poris con-
coloribus, amplis at moderate tantum radialiter elongatis, 1.5-2 mm. in diametro in
maturis; sporis 10.3-15.3x4-5 q; tramate typi Phyllopororum; stipite cremeo, ad
apicem vel omnino castaneo-reticulato, aequali vel subaequali, saepe curvato, 30-60 x
5-12 mm.; mycelio albo vel lutescente-albido; carne aequaliter flava vel pallide flava,
fracta frequenter caerulescente. Habitatio: In silvis densis frondosis (hammocks) circa
et ad bases truncorum arborum, praecipue Ostryae aut rarius ad humum, Julio mense
in Florida Septentrionali Americae Borealis.

Pileus between “snuff brown” and ‘“Saccardo’s umber,” or “Dresden
brown” when young, between “russet” and “auburn” when old, or similar
in color through all stages and then between “chestnut” and “auburn,”
tomentose, becoming faintly granulose or rimulose-tessellate, non-viscid,
pulvinate, then flat, 27-75 mm. broad. — Hymenophore “amber yellow,”
“honey yellow,” “Isabella color,” sometimes becoming as deeply colored as
“Saccardo’s umber,” at last discolored golden brownish with age, broadly

SINGER: FLORIDA BOLETINEAE II. 293

adnate (with or without a decurrent tooth) or decurrent, more or less
arcuate, tubes 5-11 mm. long, pores 1.5—2 mm. wide when mature, 7.e. wide
but only moderately radially elongate. — Stipe “cartridge buff” to “cream
buff,’ apex or entire surface adorned with anastomosing ribs or a perfect
network, the ribs coarse and “chestnut brown,” rarely broken up into
minute squamulae of the same color, solid, terete or compressed, equal or
subequal, straight or more often curved, 30—60 x 5—12 mm.; mycelium white
or yellowish white. — Context yellow or pale yellow in both pileus and
stipe, occasionally slightly bluing, yellow in dried specimens; taste slightly
acidulous or mild; odor none, in old specimens often recalling Roquefort
cheese.

Spores 10.3-15.3x4-5 yp, fusoid, melleous, smooth; basidia 25-31 x
7.5-11 pw, 4-spored; cystidia 27-38 x 6-17 p, rarely longer, always hyaline
in well-preserved material, versiform, on pores fasciculate; hymenophoral
trama of the Phylloporus-type; the lateral stratum little though undoubt-
edly divergent and no differentiation in color or density between it and the
mediostratum visible; terminal members of the cuticular hyphae upright
and often rather short to even subglobose (13-34 yw in diameter), broadly
rounded above, not forming an actual epithelium, the rich ferruginous-
chestnut-brown pigment squeezed out and dissolved in ammoniacal prep-
arations; hyphae without clamp connections.

Chemical reactions: KOH on surface of pileus, deep castaneous; on
tubes, brown; on context, ochraceous tan. — NH on surface of pileus and
hymenophore, blue. — NH,OH on surface of pileus, tan; on tubes, blue;
on context, bluish. — HNO on surface of pileus, orange then reddish. —
H.SO, on surface of pileus, orange then cinnamon; on tubes, orange-
ferruginous.

Habitat: On the bases of the trunks of Ostrya, more rarely of Liquéd-
ambar, Acer, or Carpinus, usually on the trunks of living trees, rarely on
dead trunks, sometimes on humous ground of mesophytic hammocks near
Ostrya or Liquidambar; fruiting in July, gregarious.

Distribution: North Florida.

Material studied: Fla. Alachua Co., in and around Gainesville in meso-
phytic hammocks, R. Singer, F 2473, F 2473 a, F 2473 b, etc. (FH).

‘This species is unmistakable when found typically developed. In its
external appearance, it is much like X. kypoxanthus but differs in the color
of the mycelium and the chestnut brown raised lines on the apex of the
stipe; the spores are slightly narrower on an average but there are collec-
tions where the spores are equally broad in both species. In X. squarrosoides
which, at times, is also similar in stature, the spores are distinctly broader.

15. Xerocomus Uludens (Peck) Sing. comb. nov.
Boletus illudens Peck, Ann. Rep. N.Y. State Mus. 50:108. 1897.
Ceriomyces illudens Murr., N. Am. FI. 9:145. 1910.
Ceriomyces alabamensis Murr., N. Am. Fl. 9:146. 1910 (a subspecies ?).
Boletus alabamensis Sacc. & Trott., Syll. 21:242. 1912.
Ceriomyces flavimarginatus Murr., Mycologia 31:110. 1939. (a subspecies), p.p.

294 Fartowisa, Vor, 2, 1945
Ssp. typicus.

Pileus “tawny” to “cinnamon rufous” when fresh, middle portion with
“ochraceous tawny” and “cinnamon rufous” or “russet’’ stains on generally
“buckthorn brown” ground, margin becoming slightly more grayish fuscous
than “ecru-olive,” center subtomentose, margin tomentose, non-viscid, con-
vex, center sometimes depressed, margin thick and fleshy, 63-123 mm.
broad. — Hymenophore mixed “strontian yellow” and “citron yellow,”
becoming “pyrite yellow” at last, adnate-subdecurrent, tubes 9-15 mm.
long; pores concolorous, sometimes slightly bluing on pressure, 0.8—3.0 mm.
wide. — Stipe between ‘primrose yellow” and ‘‘marguerite yellow,” or
between “ivory yellow” and ‘colonial buff,” with either concolorous or
tawny brown (color of the pileus) very coarse, wide network of strongly
raised, strongly anastomosing lines, most distinct at the apex but reaching
down to the very base, with a concolorous or “light pinkish cinnamon” faint
fibrillosity at places, or almost all over its surface, solid, decidedly tapering
downward, 40-60 x 25-35 mm. at apex; mycelium yellowish white. —
Context yellowish white with pale brownish pink portions to nearly “mas-
sicot yellow,” somewhat brownish inside the stipe; odor none, or slightly
like Boletus edulis but weaker and less aromatic; taste mild.

Spores (8.2)-9.5-12.7-(13.6) x 3.5-4.5 ys, most frequently 10-11.7 x
about 4.0 », melleous, smooth, ellipsoid-subfusoid to strongly fusoid;
basidia 28-35 x 7.5-9.2 uw, (2)—4-spored; cystidia 42-56 x 6-11 pn, hyaline,
non-incrusted, distinctly fusoid, thin- to moderately thick-walled; trama
of the hymenophore not observed in proper condition, but of the general
structure of the related forms; Ayphae without clamp-connections.

Chemical reactions: KOH on surface of pileus, deep chestnut becoming
“ochraceous orange” (but reaction weak and slow). — NH); on surface of
pileus, deep green-blue. — FeSQ, on tubes, more greenish and more gray;
on context, almost negative.

Habitat: In mixed woods on the ground, from August to September.

Distribution: From New England and upper New York State south to
the Carolinas and probably Alabama.

The above description is the result of the study of several good fresh
specimens from Massachusetts, dried material from North Carolina (Alma
Holland Beers 12523) and the type collection at Albany (NYS). Compare
also Coker and Beers’ description (/.c. p. 55). We included this description
in order to make the comparison with the Florida subspecies possible.

Ssp. xanthomycelinus Sing. ssp. nov.

A subspecie typica mycelio luteo nec non statura paulum minore differt. Sub Quer-
cubus in Florida Americae Borealis.

Pileus reddish brown to fulvous castaneous (“Mikado brown,” “auburn,”
“chestnut’’), tomentose, or subrimulose-tomentose when mature, non-viscid,
pulvinate to nearly applanate, 43-87 mm. broad. — Hymenophore about
“strontian yellow” or somewhat paler, becoming more olive yellow with
age, adnate or adnate-subdecurrent and continued in the reticulation of the

SINGER: FLoripA BoLeTINEAE II. 295

apex of the stipe; tubes medium long; pores concolorous, unchanging, wide
to very wide. — Stipe varying from pallid to yellow but if pallid it is cov-
ered with numerous dense minute flocculae which make the whole surface
appear yellow except for the brownish (testaceous-cinnamon to reddish
brown) ridges or ribs that form, on the upper third or two thirds of the
stipe, a coarse network from which longitudinal lines frequently run down-
wards for some distance, solid, tapering downward from the very apex or
from about the middle of the stipe, or only with a tapering base and other-
wise subequal, 35-70 x 8-12—(18) mm.; mycelium light yellow to rich yel-
low. — Context white with yellow portions above the tubes, or light yellow
throughout the pileus, white or sordid pale yellow in the stipe, not changing
anywhere, slightly paler than the surface immediately under the cuticle of
the pileus; taste mild; odor almost none.

Spores 9.5-14,2 x 3.5-4.5 w, most frequently 10-11.5 x 3.5—4 y, melleous,
subfusoid-subellipsoid, smooth; basidia 24-30x 7-9 p, 4-spored, in few
carpophores some 2-spored ones intermixed; cystidia 28-63 x 5.5—-11.5 p,
fusoid or subfusoid, larger and more scattered in the tubes, smaller and
more numerous on the pores, with rounded apex, hyaline, occasionally with
long slender ampullaceous neck; ¢vama of the hymenophore of the Phyllo-
porus-type; hyphae without clamp-connections.

Chemical reactions: KOH on surface of pileus showing a nearly black
ring around the stain. — NH; and NH,OH on surface of pileus, strongly
and intensely greenish blue or blue, then fading to deep chocolate.

Habitat: On lawns near oaks (Quercus virginiana and Q. laurifolia) and
shrubs in small groups, successively but not abundantly fruiting during the
summer (from May to July and probably longer).

Distribution: Northern Florida.

Material studied: Fla. Alachua Co., Gainesville, May 1943, R. Singer,
F 2167 (type, FH); July 1943, R. Singer, F 2530, F 2612, F 2620 (co-
types, FH).

This subspecies replaced the type form entirely in Gainesville while I
was collecting there. However, Murrill has determined a more yellowish
specimen with thicker stipe from Alachua County as B. dludens (FLAS),
and it would not be surprising if this species were neither the type nor the
ssp. xanthomycelinus. Murrill’s Ceriomyces alabamensis belongs most prob-
ably to one of the subspecies of X. iludens but I was unable to find out
to which. The spores are not as small as indicated by Murrill but rather
9.5-11.5 x 3.7-4.2 » in the type specimens.

EXTRALIMITAL SPECIES

Xerocomus squarrosoides (Snell & Dick) Sing. comb. nov.

Boletinus squarrosoides Snell & Dick, Mycologia 28:468. 1936.
Phylloporus squarrosoides Sing., Rev. de Mycol. 3:170. 1938.

The excellent description of this species given by its author makes it
possible to dispose of this species taxonomically without knawing the

296 FaRLowIA, VOL. 2, 1945

ammonia reaction. There is abundant material at the Farlow Herbarium,
authentic material as well as Farlow’s material from Providence, R. I.
(Boletinus decipiens) and Stow, Mass. (Boletinus castanellus), and D. H.
Linder’s collection (B. subtomentosus) from Canton, Mass., and besides a
fine unpublished color plate (Br. 372). It usually has more boletinoid hy-
menophore than other species of this genus and its color is less yellow or
less uniformly yellow than in the others except X. Linderi. The spores are
melleous-brownish, smooth, asymmetric, ovoid with tapering apex or
oblong-ellipsoid, without a distinct suprahilar depression, 9-12 x 4.56 p,
mostly about 9.8-10 x 4.8-5 yx; basidia 28-37 x (5.5)—7.7-9 y, 4-spored;
cystidia closely resembling the cystidia of Phylloporus rhodoxanthus, either
hyaline or melleous, fusoid with mostly broadly rounded tip, more rarely
ampullaceous or clavate, 40-73 x 7.5-12.2 w; trama of the Phylloporus-
type; hyphae without clamp connections. In 1938, the author placed this
species in Phylloporus. In the present arrangement we take it with all other
porous forms to Xerecomus.

Xerocomus Housei (Murr.) Sing. comb. nov.
Ceriomyces Houset Murr., N. Am. FI. 9:145. 1910.

The type (NY, NYS) has the spores still shorter than Y. squarrosoides,
ellipsoid-ovoid, smooth, melleous, 7.7—8.5 x 5 1; basidia 28 x 9.7 p, 4-spored;
cystidia fusoid, narrowly clavate, or fusoid-ampullaceous, rather numerous
35—40 x (5)—9-9.5 ; cuticle with an interrupted trichodermium-palisade,
the terminal hyphae fusoid or subulate, or capitate, 17-42 x 7-22 »; hyphae
clamp-less. It seems that this North Carolina bolete, rarely collected and
chemically not studied, belongs in section Pseudophyllopori.

Xerocomus Linderi Sing. spec. nov.

Pileo areolato, areolis fulvidis (“tawny 14”), subapplanato, 40 mm. lato, cuticula e
trichodermio palisadiformi brevi-hyphoso nec non e catenulis suberectis sphaerocystarum
consistente; tubulis carneo-cinnamomeis (“pinkish-cinnamon” R. in vivis), fractis
caerulescentibus, amplissimis, angulatis; sporis subfusoideo-ellipsoideis, brunneolo-
melleis, levibus, tenui-tunicatis, depressione supra-hilari praeditis, 11-12 x 4.7-5.2 yw;
basidiis 30-38x 9-11 yw, tetrasporis; cystidiis versiformibus, saepius clavatis; stipite
argillaceo-cinnamomeo (“clay color”) subtus, at ad apicem flocculis fuscidulis (“snuff
brown”) furfuraceis obsito, solido, aequali, 45x 5 mm.; mycelio albo, ex hyphis 2.8—3.5
mM Crassis cylindraceis consistente; carne sordide ochracea in siccis, tenuiore, odore
saporeque haud notatis. Habitatio: Ad basin nidi termitum prope Banga, Liberia
Africae, Octobri mense D. H. Linder, 1211 (type, FH).

This is apparently a Xerocomus but the section is somewhat difficult to
determine. It is in so many ways similar to X. hemixanthus and X. squar-
rosoides that we consider it as belonging in section Pseudophyllopori.

Xerocomus lanatus (Rostkov.) Sing. comb. nov.
Boletus lanatus Rostkov. in Sturm, Deutschl. Fl., Pilze 3:77. 1844.
Boletus hieroglyphicus Rostkov. l.c. 3:93. 1844.
Boletus Leguei Boud., Bull. Soc. Mycol. Fr. 10:62. 1894.
Suillus lanatus Kuntze, Rev. Gen. Plant. 37:535. 1898.
Boletus subtomentosus var. Leguei Bat., Bolets, p. 19. 1908.

SINGER: FLORIDA BOLETINEAE II. 297

Xerocomus subtomentosus var. Leguei R. Maire, Publ. Inst. Botan. Barcelona,
3(4) :43. 1935.

Boletus subtomentosus ssp. spadiceus var. lanatus Konr. & Maubl. Icon. Sel. Fung.
6:463. 1938.

The author has seen material in Boudier’s herbarium at Paris. Accord-
ing to Rostkovius it is found in beech woods, and according to Boudier,
the spores are very broad, much broader than in X. dlludens.

Xerocomus coniferarum Sing. nom. nov.
Boletus ferrugineus Bres., Iconogr. 19, pl. 915. 1931, non Schaeff. nec Frost.

This is also the B. spadiceus “Fr.” sensu Quél., Champ. Jura, p. 262.
1872, X. spadiceus (Fr. sensu Quél.) Quél., B. subtomentosus ssp. spadiceus
(Fr. sensu Quél.) Konr. & Maubl. It occurs under fir in the Vosges Mts. in
France, also in the Trentino, and the Caucasus.

Sect. Subtomentosi Fr., Epicrisis, p. 412. 1838 (sect. gen. Boleti).

Characters of the section: see key p. 287. Type species: X. subto-
mentosus (L. ex Fr.) Quél. Other species: X. chrysenteron (Bull. ex Fr.)
Quél., X. Zelleri (Murr.) Snell apud Slipp & Snell. It is possible that some
tropical species belong in this section. One of the species which the author
tentatively places here, is:

Xerocomus Junghuhnii (Hoehn.) Sing. comb. nov.
Boletus Junghuhnii Hoebn., Sitz.-ber. k. Akad. Wiss. Wien, Math. Nat. KI. 123
(I) :87. 1914.

The type (FH) from Java which we have examined in order to be able
to compare it with X. brasiliensis, shows very large pores in a very small
carpophore. Hoehnel states that the pileus is dry. In one of the transparent
bags containing type material, specimens of another species (a minute
Boletus) are intermixed, but the descriptive notes seem to entirely cor-
respond to the species which we take for the real X. Junghuhni. Even
there, we find the spores very variable in size but not so much in shape.
They are pale melleous (“pale yellowish in masses” says Hoehnel but he
had no good spore print) under the microscope, smooth, and varying be-
tween 9-10x3.8—-4.2 » in one mature specimen to (9.2)—10.2-13.3 x
(3.8)—4.2-5.3 y» in another; basidia 23.8-25.3 x 6.8-8.8 4; cystidia versi-
form, not always conspicuous, about 35-42 x 8.5-12.5 »; hyphae of cuticle
with yellow cell sap; clamps none; structure of the trama not with cer-
tainty decipherable.

Sect. Parasitici Sing., Ann. Mycol. 40:43. 1942.

Characters of the section: see key, p. 287. The only species known is the
circumpolar X. parasiticus (Bull. ex Fr.) Quél. Massachusetts material
(D. H. Linder & R. Singer, Greenwood, Sept. 1943), and Paris material
(R. Singer, Bois de Meudon, Aug. 1935) show the following chemical char-
acters: KOH and NHs on surface of pileus and on the yellow context

298 FaRLowiA, Vou, 2, 1945

castaneous; NH,OH on surface of pileus castaneous, eventually paler but
with darker ring; HNOs on surface of pileus castaneous, eventually with
a bright orange ring; FeSQ, negative. The trama of alcohol-material from
Middlesex Fells, Mass., shows an undifferentiated mediostratum, the outer
larger part of the trama diverging but only slightly so, the non-diverging
portion rather more hyaline than the diverging one, the latter of rather
densely arranged hyphae without clamp connections; both strata continu-
ing around the bottom of the tube. The inverse coloration of the tramal
strata as compared with the Boletus-type may be due to the preserving
medium. In the same material some 1.5 mm. broad and 4.5 mm. high
primordia have been sectioned by the author since they represented the
youngest stage in which the beginning hymenophore could still be demon-
strated. These primordia showed the presence of a distinct ring-hole which
is in apparent contradiction to previous statements in the literature. I am
at a loss to explain these facts unless the ring-hole in our preparations is
also due to an alternating influence on the tissue caused by the preserving
fluid. X. parasiticus has not yet been found in Florida, notwithstanding
the abundance of its hosts, Scleroderma spp., in the state.

Sect. Brasilienses Sing. sect. nov.

Pileo tomentoso et viscido; poris initio parvulis, deinde amplis vel pro ratione
amplissimis; sporis exiguis (6-9.3 x 2.8-4.2 wu); trama typi Phyllopororum; in tropicis
et subtropicis.

Characters of the section: see key, p. 287 and the above Latin diagnosis.

16. Xerocomus brasiliensis (Rick) Sing. comb. nov.
Boletus brasiliensis Rick, Ann. Mycol. 3:235. 1905.

Pileus pale or light yellow, felty tomentose and at the same time viscid,
sometimes with narrow sterile margin, hemispheric to convex, 10-20 mm.
broad, rarely broader. — Hymenophore greenish yellow, slightly depressed
around the stipe, tubes of medium length, pores concolorous, wide when
mature, inequal, irregular; spore print not obtained but judging from small
deposits on the carpophore itself, probably olive brown. — Stipe pale mel-
leous to yellow, smooth or almost so, sometimes mealy on top, solid, cylin-
dric, about 21 x 3-5 mm., rarely larger. — Context subconcolorous with
the surface, unchanging, odor none.

Spores 6-9.3-(10) x 2,8-4.2 p, mostly around 8.5 x 3.5 p» with 0.4 u thick
wall, smooth, melleous, subellipsoid-cylindric or somewhat variable in
shape; basidia (13)—19-28 x (4)—6.5-8.8 », 4-spored with 4.5 long sterig-
mata; cystidia inconspicuous and infrequent, hyaline, not or very little
incrusted, fusoid-ampullaceous, e.g. 17.5 x 5.5 wu: trama very homogeneous,
rather hyaline, of almost equal density and without an appreciably darker
colored zone in the middle, of the Phylloporus-type; cuticle of the dried
material available not showing any particular structure; laticiferous vessels
present, melleous, all Avphae without clamp connections.

Chemical reactions: KOH, NH,OH, NH, everywhere brown.

SINGER: FLORIDA BOLETINEAE IT. 299

Habitat: On the ground in open places and in dense subtropical woods
(low hammock type) on humus and débris, even up to two feet on the living
trunks of trees, never near conifers, solitary or in small groups, in Florida
fruiting in August.

Distribution: Florida (rare), Brazil (frequent). Exact limits unknown.

Material studied: Fla. Highlands Co., Highlands Hammock State Park,
R. Singer, F 641 (FH); Brazil: Near Sao Leopoldo, numerous specimens
including parts of the type and authentic material in Rick: Fungi Aus-
troamericani, 1909 (FH).

The above description proves that this is a Xerocomus, and not related
to Boletus fulvidus (i.e. Gyroporus castaneus) as suspected by Rick. This
small and inconspicuous bolete has not been found, and, for that matter,
not been looked for in the West Indies and northern South America.

Sect. Pseudoboleti Sing. sect. nov.

Pileo viscidulo at subtomentoso; tramate structura intermedia inter typum Phyllo-
pori et typum Boleti gaudente; vaporum ammoniacalium ope brevissime caerulescente-
nigrescit; in silvis coniferis borealibus.

This is the section for the species with intermediate tramal structure
that cannot easily be linked with any group within the genus Boletus, sensu
str. In the type and only species, X. badius (Fr.) Kiihn. ex Gilbert (Boletus
badius Fr.) the subhyaline lateral stratum is slightly looser than the medio-
stratum and its hyphae moderately but distinctly divergent, touching each
other and somewhat broader than in the mediostratum where the hyphae
are regular-subinterwoven and somewhat (very pale) colored. The spores
are in some specimens within the usual limits in Xerocomus, in others
larger. Coker and Beers suggest that the American X. badius has the
spores smaller than the European type. However, material collected by me
near Lynnfield, Mass. (FH) has the spores 11.5—-18.5—(24) x 4-5 yp, and
material from Austria (FH) has them 12.5-16.5 x 5—5.7 ». Some authors
consider X. badius as a Suillus but aside from the habitat, and the subviscid
pileus there are no other characters to support this view. The blackish-blue
flush with ammonia on the surface of the pileus would rather support
Gilbert’s proposal to consider it as a Xerocomus. It has not been observed
in Florida.

SPECIES INCOMPLETELY KNOWN
Xerocomus Boudieri Sing., Ann. Mycol. 40:43. 1942.

This species has not been studied thoroughly enough to decide whether
it belongs in Xerocomus as redefined in this paper. The same is true for
Boletus Roxanae and B. innixus Frost (at least authentic material at FH)
which evidently are very close to X. Boudieri if not identical. Neither of
these three species has been observed in Florida.

Xerocomus lignicola (Kallenbach) Sing., Ann. Mycol. 40:43. 1942.
Boletus lignicola Kallenbach, Pilze Mitteleur. 1:57. 1929.

300 FARLOWIA, VoL. 2, 1945

With the data at hand, I am unable to decide whether this is a Xerocomus
spec. or a form of Phlebopus sulphureus (Fr.) Sing. (Boletus sulphureus
Fr.).

Xerocomus tumidus (Fr.) Gilbert, Bolets, p. 145. 1931.
Boletus tumidus Fr., Hym. Eur., p. 501. 1874.

Fries’ species is dubious as its author states himself. Gilbert bases his
treatment on Peltereau’s emendation but this also is insufficient for generic
determination.

SPECIES EXCLUDENDAE

Xerocomus amarellus Quél. is Suillus piperatus var. amarellus (Quél.) Sing.
Xerocomus gentilis (Quél.) Sing.

This is treated as Pulveroboletus gentilis. The same disposal is made of
Xerocomus auriporus (Peck) Sing. and X. flaviporus (Earle) Sing. In fact,
the entire section Auripori has been transferred to Pulveroboletus.

Xerocomus griseus (Frost) Sing.= Boletus griseus Frost.
The section Grisei has been reunited with Boletus.

Xerocomus impolitus (Fr.) Quél. is Boletus impolitus Fr.
Xerocomus mirabilis (Murr.) Sing. is Boletellus mirabilis (Murr.) Sing.

The section Mirabiles has been transferred to Boletellus.
Xerocomus pulverulentus (Opat.) Gilbert.

This species is here retransferred to Boletus. Xerocomus radicans (Pers.
ex Fr.) sensu Quél. (non Kallenbach) is the same.

Xerocomus rubellus (Krombh.) Quél. is here treated as Boletus.

Xerocomus chrysenteron var. versicolor Quél., X. armeniacus Quél., X.
pruinatus (Fr.) Quél., and X. versicolor (Rostkov. non S. F. Gray) Gilbert
are forms of the same species. In this country, Boletus bicolor Peck and
B. fraternus Peck belong in the same complex, but they have never been
considered as Xerocomi.

Xerocomus sulfureus (Fr.) Queél. is Phlebopus sulphureus (Fr.) Sing.
Xerocomus spec. Heim, Bol. Soc. Broter. 13(2) :53. 1938 is Boletochaete spec.
Xerocomus spec. Snell apud Slipp & Snell, Lloydia 7:50. 1944 is Boletus spec.

302 FaRLowlA, VoL. 2, 1945

EXPLANATION OF PLATE I

All drawings of spores are at a magnification of 2000; all other drawings of micro-
scopic structures at 1000; figs. 5-7 shown at natural size.

Fig. 1-4. Gyroporus purpurinus (Snell) Singer
1. Structure of the cuticle of the pileus.
2. Cheilocystidia of the hymenophore.
3. Basidium.
4. Spore.
Fig. 5-7. Boletinus decipiens (B. & C.) Peck
5. Gasteromycetous conditions of the carpophore sketched at natural size.
6. Longitudinal section of an immature carpophore (natural size). The veil closely
applied to the surface of the hymenophore.
7. Longitudinal section of a mature carpophore (natural size), the remnant of the
veil remaining as a relatively closely attached annulus.
Fig. 8-10. Suillus cothurnatus ssp. thermophilus Singer
8. Part of the hymeniform fragments on the surface of the stipe that are found in
and around the glandular dots.
9. Large dermatocystidia, another element of the glandular dots of the stipe.
10. Spore.
Fig. 11-12. Phylloporus rhodoxanthus ssp. foliiporus (Murr.) Singer
11. Spores.
12. Cystidia of the hymenophore.

SINGER: FLORIDA BOLETINEAE II.

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THAXTER, R. Entomophthoreae of the United States, 1888.

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Vol. 2 JANUARY, 1946 No. 3

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CONTENTS OF VOL. 2, NO. 3

Note on ARGENTINE CHarAsS. By Roy M. Whelden. . . . .
New Mosses FRoM Tierra DEL Furco. By Edwin B. Bartram. .
Tue Genus CUNNINGHAMELLA (Mucorares). By Victor M. Cutter, Jr...

A ComPaARATIVE STUDY OF TORULOPSIS PULCHERRIMA AND TAPHRINA DEFORMANS
mn CuLTuRE. By Catherine Roberis .

THE CYPERICOLOUS AND JUNCICOLOUS SPECIES OF SCLEROTINIA. By H. H. Whetzel

Vol. 2, No. 2 was issued on September 17, 1945.

305
309
321

345
385

FARLOWIA

A JOURNAL OF CRYPTOGAMIC BOTANY

Vous2 JANUARY, 1946 No. 3

NOTE ON ARGENTINE CHARAS
Roy M. WHELDEN

It may be conceded that the Charophyte-flora, even in those countries
most thoroughly explored botanically, is none too well known. In countries
such as Argentina, large in area and varied in climate, our knowledge is
very much less, and is based almost entirely on incidental collections of the
plants made ey those who were primarily interested in gathering quite
different material.

The scarcity of Charophyte-material is clearly indicated by the paucity
of papers mentioning any such plants from Argentina. Alexander Braun
(1882) mentioned Chara foetida A. Br. as occurring in that country, and
referred to a form of Chara Martiana from there also. Spegazzini (1883)
listed, as occurring there, ten species of these plants (6 Nitella, 1 Lampro-
thamnus, and 3 Chara, with several varieties and forms in addition), three
being described as new species. It may be added that a few papers record
the occurrence of additional species from the several countries bordering
Argentina. All these suggest that despite the very small number of refer-
ences to these plants they may be found to be of rather frequent occurrence
there.

The present note results from the study of a small collection of five
folders of the genus Czara, collected in October and November 1938, and
January 1939, by W. J. Eyerdam, A. A. Beetle, and E. Grondona. As is not
unusual with this group of plants, much of the material was received in a
badly fragmented condition; however, species determination of all speci-
mens was easily possible.

Three of the plants proved to be Chara foetida A. Br., noted above as
occurring there. The condition of the specimens was such that determina-
tion of varieties or forms seemed inadvisable. Indeed the question might
be raised as to whether the present state of knowledge of Charas is suffi-
ciently ample to justify the establishment of too many varieties. Certainly
casual examination of the three plants might lead to a conclusion that one
had before him at least two species and perhaps a variety of one, making
allowances for the immature condition of one of them. Yet in every par-
ticular — stem cortication, stipular crown, nature of branches, and fruit
details — they closely vesemble each other, varying only in the amount of
incrustation, in the nature of the bracts, and more noticeably in the size
of the sess

305

306 Fartowla, Vor. 2, 1946

One of the two remaining folders was rather confusing on first examina-
tion, owing to the presence of two distinct species, Chara Martiana A. Br. and
fragments of what seemed to be an atypical form of Chara foetida A. Br.
The Chara Martiana was a male specimen.

The fifth folder contained a plant which proved to be a real puzzle. The
plants were collected from “a still pond, in soft clay mud,” and were noted
to have ‘fruit very malodorous.” The specimens (Fig. 1) were all small, the
largest just over 10 cm. tall, of quite delicate appearance, and each of
several slender spreading branches. The incrustation, present throughout
the plant, was thin and inconspicuous, giving it a rather brownish-green
color (dried specimens).

The stems are rather irregularly tripli-corticate (Fig. 3b), and 0.5—0.62
mm. in diameter, the primary cells usually slightly more prominent than
the secondary. No spine cells occur, but only small internodal cells. The
stipular crown (Fig. 3a) comprises a single row of rather irregularly ar-
ranged cells, twice the number of leaves at the node, each cell rounded or
slightly elongate. Leaves 8-10 in a whorl, up to 1.3 cm. long, comprising
6—9 segments, the last of which is a very short, conical cell. Many leaves
are completely ecorticate; in others, one, two, or the three lowest nodes
may be corticated. The nature of the cortication (Fig. 2b) is in no way
correlated with the node in which the leaf occurs, there occurring every
conceivable variation from whorls of leaves all entirely ecorticate to those
in which the lowest segments of every leaf are corticated. The bracteoles
are likewise quite variable in number and in length, but occur only at the
lower internodes, usually only on those bearing sporangia. Those sub-
tending the sporangia are all cylindrical, with abruptly tapering ends, and
may considerably exceed the sporangia in length. Those opposite the spo-
rangia are much reduced, often to inconspicuous short conical cells. The
antheridia, found only in the youngest leaves near the stem apex (Fig. 2c),
are 280-320 » in diameter; they are soon deciduous. The sporangia occur
on the lower two to three internodes, and are 820-860(—900) » long and
410-425 » in diameter. The odspores are dark brown, have 14-15 spiral
ridges on the surface and are 600 » long and 325 p» in diameter. The wall
between the ridges is quite plane.

This plant has many points in common with Chara Curtisit T. F. Allen,
Ch, delicatula Ag., Ch. hydropitys A. Br., and Ch. leptopityvs A. Br. From
all these it is set off by the absence of spine cells, by its poorly developed
stipular crown of a single row of scarcely elongated cells, and by the nature
of the cortication of the leaves. From other described Charas it is much
more completely separate. It is therefore now described as a new species.

Chara leiopytis sp. nov.

Monoica, tenera et flexilis, leniter incrustata 10 cm. longa; caule 0.5-0.62 mm. in
diam., triplo-corticatis, aculeis nullis sed cellulis verruciformibus; corona stipulari du-
plici foliorum numero, ex unica cellularum paullo elongatarum vel rotundatarum serie
inaequaliter disposita; verticillis foliorum e foliis 8-10 compositis, 1.3 cm. longis,
geniculis 6-9, articulis omnibus ecorticatis vel infimis 1-3 corticatis, corticatis ecortica-

WHELDEN: ARGENTINE CHARAS 307

tisque saepe in eodem verticillo; geniculis infimis bracteiferis, bracteis inaequalibus,
longis cylindricis, apicibus conicis vel breviter conicis; antheridiis 280-320 mw in diam.
mox caducis; sporangiis solitariis, oblongis, 820-860(—900) yu longis, 410-425 uw in diam.;
oosporis 600 w longis, 325 w in diam., 14-15-gyratis, inter gyros planis.

Argentina: In stagnis ad argillam teneram, in Prov, Jujuy, Depto. San Pedro.
Harvarp UNIVERSITY

CAMBRIDGE, Mass.
LITERATURE CITED

1882. Braun, A.. Fragmente einer Monographie der Characeen. Edited by O. Nord-
stedt. Abhandl. K. Akad. Wiss. Berlin 1882. 211 pp. 7 pl.
1883. Spegazzini, C. Characeae Platensis. Anal. Soc. Cient. Arg. 15: 1-14.

308 Fartowia, VoL. 2, 1946

s
SOO}

Chara leiopytis Whelden n. sp.

Fig. 1. Habit of plant. Fig. 2a. Two illustrations of successive whorls, showing
irregularities in branch cortications. Fig. 2b. Three branches showing irregularity of
cortication, of node length, and of bracteoles. Fig. 2c. Two branches near apex of stem
bearing antheridia. Fig. 3a. Two stem nodes showing stipular crown. Fig. 3b. Details
of stem cortication. Fig. 4. Sporangium and odgonium. Fig. 5. Detail of wall of
o6gonium.

The magnification of each figure is given by an appropriately numbered scale.

2(3):309-319 FARLOWIA January, 1946

NEW MOSSES FROM TIERRA DEL FUEGO
EpWIN B. BarTRAM

About ten years ago I had the privilege of studying an extensive collec-
tion of mosses made by Dr. H. Roivainen in his capacity as botanist with
the scientific expedition of the Finnish Geographical Society to Tierra del
Fuego in 1928-29. All of the material was returned to Finland and sub-
sequently I received a complete series of specimens including the types of
many new species which resulted from the study, all of which are now in
my herbarium.

In 1937 Dr. Roivainen published the results of his bryological investiga-
tions relating to Sphagnaceae, Andreaeaceae, Fissidentaceae, Ditrichaceae,
Seligeriaceae and Dicranaceae.1 This was to be followed by treatments of
the succeeding groups but since then the unsettled conditions in Europe
and especially in Finland have not only prevented further publications but
have made it highly uncertain when, if ever, we may expect to have the
remaining results of Dr. Roivainen’s personal observations and studies.
Meanwhile it seems advisable to record the descriptions of the new species,
many of which are of the utmost interest.

DICRANACEAE

Chorisodontium sericeum Bartr. sp. nov.

Caespites densi, aureo-lutescentes, inferne fusci. Caulis ad 2 cm. altus, haud tomen-
tosus, ramosus. Folia caulina circa 5 mm. longa, superne falcato-secunda, ovato-
lanceolata, concava, sensim in acumen artistatum planum apice denticulatum attenuata;
costa pallida, ad basin 50 w lata, excurrente; cellulis alaribus distinctis, sequentibus
linearibus, sinuosis, superioribus subquadratis vel oblongis. Folia perichaetialia e basi
longe convoluta, subulata; seta 2.5 cm. longa, rubra, theca erecta, cylindrica; peristo-
mium destructum. Caetera ignota.

Chile: Prov. de Magellanes, Isle Clarence, Puerto Beaubasin, _ad saxa,
No. 2297 type.

While similar to C. lanigerum (Besch.) Broth., this species seems to. be
constantly distinct in the flat, sharply toothed leaf tip, the narrower costa,
and the stems almost devoid of either radicles or tomentum.

POTTIACEAE

Molendoa fuegiana Bartr. sp. nov.

Sat robusta, dense caespitosa, olivaceo-viridis, inferne fuscescens. Caulis circa 1.5 cm.
altus, ramosus, inferne tomentosus, dense foliosus. Folia sicca contorta, madida arcuato-

* Roivainen, H. Bryological Investigations in Tierra del Fuego. Ann. Bot. Soc. Zool.-
Bot. Fenn. Vanamo, 9(2) :v—x; 1-58. 1937.

309

310 FARLowIA, VoL. 2, 1946

patula, flexuosa, comalia stellata, ad 2.5 mm. longa, lineari-lanceolata, canaliculata,
acuminata; marginibus erectis, undulatis, papilloso-crenulatis; costa valida, fusca, ad
basin 100 wu lata, excurrente, mucronata, apice papillosa; cellulis superioribus rotundato-
quadratis vel transverse dilatatis, chlorophyllosis, obscuris, dense papillosis, marginali-
bus hic illic bistratosis, inferioribus linearibus, rectangularibus. Caetera ignota.

Fuegia media: Lago Blanco, Puesto de los Indios, ad rupes, Nos. 162a,
163 type, 173.

As compared with the description of M. boliviana Broth. and M. Herzogit
Broth. the leaves of MM. fuegiana are appreciably longer while in M. andina
Mitt. the leaves are narrower below and the upper leaf cells smaller and not
transversely elongate.

Didymodon ampliretis Bartr. sp. nov.

Caespites compacti, virides, inferne fuscescentes. Caulis 1-1.5 cm. altus, pluries
divisus, gracilis. Folia sicca parce contorta, humida erecto-patentia, ovato-lanceolata,
canaliculata, sensim angustata, apice subobtusa; marginibus minute papilloso-crenulatis,
medium versus anguste reflexis; costa valida, infra summum apice evanida; cellulis
superioribus rotundato-quadratis, dense chlorophyllosis, obscuris, papillosis, marginalibus
bistratosis, inferioribus rectangulis, pellucidis. Caetera ignota.

Fuegia media: Lago Fagnano, punta Pizzaro, ad rupes, No. 1217 type;
Fuegia bor.: Estancia Esperanza, No. 427.

By comparison with D. diaphanobasis Card. of Mexico, the upper leaf
cells of D. ampliretis are much more chlorophyllose and obscure while the
basal cells are shorter with more firm, pellucid lateral walls.

Tortula robusta Hook. & Grev. var. laxa Bartr. var. nov.
Cellulis laminalibus laxioribus, 15-20 yu latis, grosse papillesis.

Fuegia media: Lago Linch, in silva paludosa, No. 189; Lago Linch, Cerro
Chico, in reg. alp., No. 1357; Estancia Cameron, Puesto Medio, in prato
humido, No. 1895 type; Fuegia occidentalis: Fjordo de Agostini, Bahia
Groth Hansen, No. 254; Fjordo Martinez, ad rupes sericiticas, No. 583.

The collections referred here differ from typical 7. robusta, as I under-
stand it, in the large, lax, thin-walled cells of the upper lamina which are
coarsely papillose with C-shaped papillae. The costa is smooth to papillose
on the back. The leaves are narrower than those of T. rivularis Dus. and
not as strongly serrate, while the cells are appreciably larger than in
T. pseudorobusta Dus.

GRIMMIACEAE

Grimmia scabrines Bartr. sp. nov.

Autoica, caespitosa, sat robusta, fusco-viridis. Caulis prostratus, 3-4 cm. longus,
ramosus. Folia caulina conferta, sicca appressa, humida erecto-patentia, ovato-lanceo-
lata, concava, sensim in pilum hyalinum spinulosum attenuata, 2-3 mm. longa, 1 mm.
lata; costa valida, in pilum producta, dorso papillosa; marginibus integris, toto fere
longitudine late recurvis; cellulis superioribus rotundatis, altissime papillosis, hic illic
bistratosis, margines versus 3-4 stratosis, inferioribus rectangularis, parietibus maximus
incrassatis sinuosisque. Folia perichaetialia erecta, majora, costa angustiore; capsula
immersa, breviter pedicellata, oblonga; peristomium aurantiacum, dentibus lanceolatis,
integris; sporae lutescentes, laeves, diam. 8-10 wu.

BartTRAM: MossEs FROM TIERRA DEL FUEGO Sl

Chile: Prov. de Nuble, Recinto, Las Trancas, ad truncos Nothofagi
antarcticae, alt. 1150 m., No. 1003 type.

Readily distinguished from any of the forms of G. apocarpa and, in fact,
from any of the species of the section Schistidium by the coarsely papillose
leaf cells. A cross section of the leaf shows the papillae to the best ad-
vantage but they may be clearly seen in profile on the roll of the leaf
without sectioning.

Rhacomitrium limbatum Bartr. sp. nov.

Robustum, dense caespitosum, nigrescens. Caulis erectus, ad 4 cm. altus, irregulariter
divisus. Folia conferta, sicca erecta, parce flexuosa, humida patentia, 3.5-4 mm. longa,
ovato-lanceolata, sensim acuminata, superne carinata, toto ambitu limbo lato incrassato
circumducta; marginibus integris, inferne reflexis, superne planis; costa valida, excur-
rente; cellulis inferioribus linearibus, parietibus incrassatis, valde sinuosis, superioribus
brevioribus, oblongis, costa versus bistratosis, margines versus 6—9 seriatis 3—4 stratosis.
Seta 5-6 mm. longa; peristomii dentes rufescentes, ad basin divisi, papillosi; sporae
papillosae, diam. 15-22 u.

Fuegia occid.: Fjordo Finlandia, Brazo Aina, ad saxa, No. 502; Fjordo
Martinez, ad rupes uliginosas, No. 594; Fjordo de Agostini, Bahia Groth
Hansen, No. 2268; Chile: Prov. de Magellanes, Fuegia occid., Monte Buck-
land, alt. 500 m., Nos. 2084a, 2644; Isla Clarence, Puerto Beaubasin, ad
saxa, Nos. 2313, 2316 type.

This exceedingly fine and interesting species is immediately distinguished
from any of the forms of R. crispulum, and also from any member of the
genus with which I am familiar, by the broad, fleshy leaf border extending
from the apex almost to the insertion.

Rhacomitrium scabrifolium Bartr. sp. nov.

Robustum, aureo-lutescens. Caulis ad 8 cm. longus, irregulariter divisus, ramis
breviter ramulosis. Folia 3.5 mm. longa, sicca erecta, humida erecto-patentia, ovato-
lanceolata, plicata, superne canaliculata, sensim in pilum hyalinum denticulatum pro-
ducta; marginibus papilloso-crenulatis, reflexis; costa depressa, angusta, ad medium
folii evanida; cellulis omnibus elongatis, linearibus, papillosis, parietibus incrassatis,
lutescentibus, valde sinuosis. Caetera ignota.

Chile: Prov. de Magellanes, Fuegia occid., Monte Buckland, alt. 500—700
m., No. 241; Puerto Yarton, Pico Nariz, reg. alp. No. 820; Fjordo Fin-
landia, Brazo Aino, No. 1970; Fjordo de Agostini, No. 2068 type; Bahia
Sarmento, No. 2465.

The combination of plicate leaves and papillose leaf cells will separate
this species from any of its Fuegian associates. In R. ptychophyllum Mitt.
of New Zealand, the leaves are plicate but the short lateral branchlets are
lacking and the leaf cells are smooth.

FUNARIACEAE

Funaria (Entosthodon) pungens Bartr. sp. nov.

Autoica, pusilla, gregaria, lutescente-viridis. Caulis 3-5 mm. altus, inferne radiculosus.
Folia ovato-oblonga, acuminata, concava, comalia 2.5-3 mm. longa; marginibus integris,
erectis, saepe ad basin uno latere inflexis; costa tenui, longe ab apice folii evanida;

312 FarLowlA, VoL. 2, 1946

cellulis superioribus oblongo-hexagonis, ad 50 wu latis, inferioribus rectangularibus. Seta
erecta, rubella, ad 18 mm. longa; theca erecta, pyriformis; peristomium simplex, dentes
rufi, papillosi; sporae fuscescentes, diam. 25-28 wu.

Fuegia occidentalis: Fiordo Martinez, Bahia Sarmento, No. 2358 type.

This species will be distinguished from F. chiloensis Mitt. by the sharply
acuminate leaves and the broader and laxer areolation of the upper part
of the leaf. :

SPLACHNACEAE

Tayloria magellanica (Brid.) Mitt. var. gigantea Bartr. var. nov.
Caules 18-20 cm. alti, dense compacti.

This seems to be only an extremely robust form with densely tufted
parallel stems, up to 2 dm. high, showing very clearly the zones of growth.
The development is unusual and in this condition the plant is an excep-
tionally fine one.

Fuegia occidentalis: Fjordo Finlandia, prope Montem Nylandiam, No.
1745 type.

BRYACEAE

Mielichhoferia multiflora Bartr. sp. nov.

Synoica; flores plures stipitati. Caespites compacti, superne lutescente-virides, nitidi,
inferne dense fusco-radiculosi. Caulis erectus, ad 2.5 cm. altus, dense julaceo-foliosus.
Folia erecta, imbricata, ovato-lanceolata, acuminata, carinato-concava; marginibus
erectis, superne denticulatis; costa valida, infra apicem folii evanida; cellulis anguste
rhomboideis, basilaribus rectangularibus. Seta 8-10 mm. longa, lutescente-rubra; theca
inclinata, asymmetrica; annulus latus; endostomium pallidum, corona basilaris humilis,
processus angusti, linea divisurali distincti; operculum depresse conicum, apiculatum;
sporae lutescentes, diam. ad 20 uw.

Chile: Prov. de Nuble, Cordillera de Chillan, alt. 2200 m., No. 1699
type.

In some respects the details of this species closely approximate the de-
scription of M. leptoclada C. M., but in M. multiflora the leaves are erect
and closely imbricated and the segments of the endostome show a median
line.

The numerous stalked flower buds are conspicuous and may be a good
diagnostic character. While the fertile flowers are synoicous, the remaining
undeveloped buds are often composed entirely of antheridia. Either the
archegonia in these buds are abortive or the inflorescence is heteroicous.

Bryum (Cernuiformia) Roivaineni Bartr. sp. nov.

Synoicum; caespitosum, caespitibus densiusculis, sordide viridibus. Caulis ad 5 mm.
altus, basi fusco-tomentosus, innovando-ramosus. Folia ad 3 mm. longa, ovata, acuta,
haud decurrentia, concava; marginibus anguste recurvis, integris; costa crassa, fusca,
infra apicem folii evanida vel percurrente; cellulis oblongo-hexagonis, firmis, basilaribus
rectangularibus, concoloribus, marginibus elongatis, limbum biseriatum, fuscescentem
efformantibus. Seta circa 2 cm. longa, rubra; theca nutans, oblongo-piriformis, sicca
deoperculata, haud constricta; exostomii dentes fusco-lutei, endostomum sordide
luteum, papillosum, processus angusti, exostomio adhaerens; sporae 18-22 mw in diam.

BaRTRAM: MossEsS FROM TIERRA DEL FUEGO ais

Chile: Prov. de Magellanes, Fuegia occid., Fjordo Finlandia, Brazo Aino,
No. 2110 type.

This species seems to be distinct from both B. austro-chilense and B.
dicarpum in the broader, less sharply pointed leaves and the thick, strong
costa.

Bryum (Inclinatiformia) fuegianum Bartr. sp. nov.

Synoicum; gracile, caespitosum, caespitibus mollibus, pallide viridibus, inferne
fuscescentibus. Caulis erectus, 2 cm. altus, gracilis, innovando-ramosus, innovationibus
erectis, circa 2 cm. longis, base radiculosis. Folia sicca erecta, contorta, humida erecto-
patentia, spathulata vel obovata, breviter acuminata, usque ad 3 mm. longa, 1.5 mm.
lata; marginibus inferne reflexis, integris; costa infra apicem folii evanida vel percur-
rente; cellulis laxis, teneris, oblongo-hexagonis, basilaribus rectangularibus, rubris,
marginalibus elongatis, limbum 3-4 seriatum efformantibus. Seta tenuissima, rubra,
5-6 cm. longa; theca nutans; exostomii dentes fusco-lutei; endostomium hyalinum,
papillosum, processus angusti, superne dentibus adhaerens, cilia 0, sporae 25 mw in diam.

Fuegia media: Puesto Medio de la Estancia Cameron, in palude, No. 8
type. |

Bryum (Cernuiformia) austro-chilense Bartr. sp. nov.

Synoicum; caespitosum} viridibus, inferne fuscescentibus. Caulis ad 1 cm. altus, basi
fusco-tomentosus, innovando-ramosus. Folia sicca erecta, contorta, late ovata, breviter
acuminata, carinato-concava, circa 1.5 mm. longa; marginibus anguste reflexis; costa
fusca, percurrente; cellulis laxis, oblongo-hexagonis, basilaribus rectangularibus, con-
coloribus, marginalibus elongatus, limbum biseriatum efformantibus. Seta ad 2 cm. alta,
rubra; theca nutans, pyriformis, 3 mm. longa; exostomii dentes fusco-lutei, apice
hyalini, endostomium pallide luteum, processus angusti, superne dentibus adhaerens,
cilia 0; sporae papillosae, 25 uw in diam.

Chile: Prov. de Nuble, Cordillera de Chillan, alt. 2000-2200 m., Nos.
1042, 1043, 1693 type.

Perhaps near B. obscurum Card. & Broth. but in comparison with the
description of this species, the leaves of B. austro-chilense are smaller, the
costa percurrent and the leaves of the innovations not comose at the tips.
Bryum (Cernuiformia) dicarpum Bartr. sp. nov.

Species precedenti forsan affinis sed costa infra apicem folii evanida, sporis majoribus
et theca pendula, microstoma.

Synoicous; similar in habit and vegetative characters to B. austro-chilense
but the leaves slightly larger, up to 2.5 mm. long with the costa ending just
below the apex; setae constantly two from the same perichaetium; capsules
pendulous, small-mouthed; spores 32-35 yw in diam.

Fuegia bor.: Hotel Bouqueron, in loco paludoso, No. 317 type.

Bryum (Inclinatiformia) macrosporum Bartr. sp. nov.

Dioicum; caespitosum, caespitibus densis, viridibus. Caulis erectus, 3-5 mm. altus,
superne comoso-foliosus, inferne fusco-tomentosus. Folia comalia late oblongo-ovata,
acuminata, carinato-concava, sicca erecta; marginibus integris, anguste reflexis; costa
rubra, in aristam usque 300 u longam, levem excedente; cellulis rhomboideis, basilaribus
rectangularibus, rubris. Seta circa 2 cm. alta, rubra; theca pendula, 1 mm. longa,
oblonga; exostomii dentes lutei, apice hyalini, endostomium hyalinum, processus

314 FartowlA, VoL. 2, 1946

angusti, fenestrati, ciliis interpositis 3, parum brevioribus, haud appendiculatis; sporae
lutescentes, papillosae, 30 » in diam.

Chile: Prov. de Magellanes, Fuegia media, Lago Fagnano, Punta Piz-
zaro, No. 1211 type.

By the dioicous inflorescence this species is allied to B. vernicosum Dus.
and B. Hatcheri Dus. but is distinct from both in the well-developed cilia
of the inner peristome and the larger spores.

Bryum (Pseudotriquetra) brachycarpum Bartr. sp. nov.

Dioicum; caespitibus densis, pallide viridibus, intus fuscescentibus. Caulis erectus,
innovando-ramosus. Folia ovata, breviter acuminata, limbata, ad 2.5 mm. longa;
marginibus inferne anguste revolutis, summo apice minute denticulatis; costa inferne
rubra, percurrente vel breviter excedente; cellulis rhomboideis, basilaribus oblongis,
rubris, marginalibus elongatis, limbum 3—4-seriatum efformantibus. Seta 5—6 cm. longa,
tenuis, rubra; theca pendula, 2.5 mm. longa, piriformis; cilia appendiculata; operculum
convexum, apiculatum; sporae lutescentes, diam. 18-20 x.

Fuegia media, Estancia la Marina, in palude, No. 677 type.

This species is apparently near B. zeballosicum Card. & Broth, but is
clearly distinguished from this and any of the allied group by the much
longer setae and the short, pyriform capsules with the neck only about half
as long as the sporangium.

Bryum (Caespitibryum) pauperculum Bartr. sp. nov.

Dioicum; caespitosum, caespitibus densis, superne lutescenti-viridibus, intus fusces-
centibus. Caulis vix 1 cm. longus, inferne fusco-radiculosus. Folia sicca appressa,
humida erecto-patentia, triangulari-lanceolata, ad 2.5 mm. longa, sensim acuminata;
marginibus fere ad apicem revolutis, integris; costa valida, in aristam laevem excedente;
cellulis oblongo-hexagonis, basilaribus rectangularibus, rubris, alarabis laxis, marginali-
bus elongatis, limbum indistinctum efformantibus. Seta circa 2 cm. longa; exostomii
dentes pallide lutescentes, endostomium hyalinum, papillosum, cilia appendiculata ;
sporae minutissime papillosae, fuscae, diam. circa 30 p.

Chile: Prov. de Magellanes, Fuegia occid., Cabo San Pablo, Nos. 658
type, 670.

This species evidently stands in close proximity to B. caespiticum but
the leaf outline is different, the margins and the arista are entire and the
spores twice as large.

Bryum (Rosulata) pseudothyridium Bartr. sp. nov.

Dioicum; robustum, caespitosum, caespites densi, pallide lutescente-virides, nitidius-
culi. Caulis erectus, dichotome innovans, dense rufo-tomentosus, ad 4 cm. altus. Folia
ad apicem innovationem congesta, sicca erecta, flexuosa, spiraliter contorta, humida
erecto-patentia, oblongo-obovata, apiculata, late limbata, ad 5 mm. longa; marginibus
fere ad apicem revolutis, apicem versus remote denticulatis; costa crassa, basi rubella,
in apiculum laevem excedente; cellulis ovato-hexagonis, basilaribus rectangularibus,
infimis rubris, marginalibus linearibus, multiseriatis, limbum latum distinctum effor-
mantibus. Seta ad 2.5 cm. longa, rubella, flexuosa; theca nutans vel pendula, oblongo-
cylindrica, brevicollis, circa 3 mm. longa; operculum conicum, apiculatum; exostomil
dentes fusco-lutei, apice hyalini, 0.6 mm. alti, endostomium pallide luteum, papillosum,
corona basilaris alta, processus angusti, rimosi, cilia 3, valde, appendiculata; sporae
minute papillosae, diam. 12 wu.

BARTRAM: MOSSES FROM TIERRA DEL FUEGO S15

Chile: Prov. de Magellanes, Fuegia occid., Puesto Yarton, loco uliginoso,
No. 893 type; Fuegia media, Rio Bueno, Nos. 386, 1089.

An unusually handsome moss which is distinct from B. Lechleri C. M.
and B. andicola Hook. in the more elongate and robust stems and the broad
leaf border, up to 150 » wide in the upper half of the leaf and composed of
8-15 rows of narrow linear cells.

BARTRAMIACEAE

Philonotis luteola Bartr. sp. nov.

Caespites compacti, pallide lutescente-virides. Caulis 5-6 cm. altus, inferne parce
radiculosus, ramosus, ramis gracilis, laxifolius. Folia erecta, oblongo-ovata, breviter
acuminata, haud plicata 2-2.5 mm. longa; marginibus planis, superne serrulatis; costa
lata, male limitata, infra apicem dissoluta, dorso superne papillata; cellulis omnibus
laevibus, laxiusculis, superioribus rhomboidalibus, inferioribus longioribus, marginalibus
haud distinctis. Caetera ignota.

Fuegia media: Estancia Vicuna, Puesto 20, Cerro Fuentes, reg. alp.,
No. 734 type.

The unbordered leaves, broader costa and absence of any differentiated
marginal cells at the basal angles will at once separate this species from any
of the forms of P. vagans. It is distinguished from P. acicularis (C. M.) by
the plane margined leaves and from both P. varians Card. and P. parallela
Dus. by the costa ending below the apex.

Breutelia angustiretis Bartr. sp. nov.

Caespites densi, nitidi, superne lutescente-virides, inferne fusci, dense rufo-tomentosi.
Caulis 4-5 cm. altus, irregulariter ramosus. Folia erecto-patentia, subflexuosa, e basi
ovata, plicatula sensim longissime cuspidata, ad 4.5 mm. longa; marginibus integris, e
basi anguste recurvis, superne planis; costa angusta, percurrente vel excurrente ; cellulis
omnibus anguste linearibus, laevibus, parietibus incrassatis, lutescentibus, infimis bre-
vioribus, rubellis, parietibus sinuosis. Fructus ignotus.

Fuegia occidentalis: Fjordo Martinez, Bahia Pluschow, Nos. 592 type,
1136; Chile: Prov. de Magellanes, Isla Dawson, Puerto Valdez, in silva
humida, Nos. 1326, 2563.

This species a B. chrysura (C. M.) Broth. are closely allied to B.
integrifolia (Tayl.) Jaeg. “A critical study of this group may show that the
specific characters are too unstable to afford any practical basis for classi-
fication.

ORTHOTRICHACEAE

Macromitrium (Teichodontium) coriaceum Bartr. sp. nov.

Laxe caespitosum, lutescente-viride, intus fuscescens. Caulis repens, dense ramosus,
ramis suberectis, usque ad 1 cm. longis, ramulosis. Folia sat densa, sicca erecta, contorta,
humida erecto-patentia, lanceolata, plicata, breviter acuminata, superne carinata, 1.5—2
mm. longa; marginibus superne denticulatis; costa sub apice evanida ; cellulis superiori-
bus ovalis, incrassatis, laevibus, basilaribus anguste f fanrulbribus, valde incrassatis.
Seta erecta, laevissima, 7-8 mm. longa; theca laevissima, oblongo-cylindrica, exothecio
coriaceo; exostomium circa 30 m altum, dentibus inter se concretis, papillosis, endos-
tomium exostomio majus, processus 32, lati, obtusi, dense papillosi; calyptra nuda,
laciniata; sporae papillosae, diam. 50-60 pu.

316 FarLow1a, VoL. 2, 1946

Chile: Prov. de Chiloe, Puerto Barroso de la Penins, Tres Montes, Nos.
1563, 1570 type, 1588, 1628.

Clearly distinguished from M. Rusbyanum E. G. Britt. and M. macro-
sporum Herz. by the longer segments of the inner peristome and the naked
calyptrae.

LEMBOPHYLLACEAE

Camptochaete arbusculans Broth. ex Bartr.

Dioica; caespitosa, lutescente-viridis. Caulis sublignosus, bipinnate ramosus, ramis
teretibus, saepe flagelliformiter attenuatis. Folia conferta, concava, imbricata, latissime
ovata vel obovata, minute apiculata, 1.5 mm. longa, 1 mm. lata; marginibus erectis,
superne denticulatis; costa variabilis, nunc simplex et fere ad medium folii producta,
nunc brevis et binis vel nulla; cellulae anguste rhomboideae, laevissimae, apicales
breviores, alares sat numerosae.

Chile: Prov. de Nuble, Recinto, Las Trancas, alt. 1200 m., No. 1600;
Chile australis in montibus Cordillera de la Costa supra Angol oppidam in
truncis arborum, Dusen No. 338, Nov. 11, 1896, type.

Roivainen’s No. 1600 is exactly similar to Dusen’s No. 338, issued as
Cam ptochaete arbusculans Broth. which has been made available for com-
parison through the kindness of M. Theriot. The species is included by
Brotherus in Ed. 2 of the Pflanzenfamilien but as the description seems

never to have been published it may be validated by the above brief
diagnosis.

The stem and branch leaves are not clearly differentiated. The leaves
are very broadly ovate or obovate, minutely apiculate, alar cells rather
numerous, upper cells small, parenchymatous, rhomboidal, median and
lower cells narrower; margins erect, denticulate above; costa variable, short
and double or even lacking, often single and extending nearly to mid-leaf.
The variable costa is an anomalous character in Camptochaete but the
plants are otherwise characteristic so I believe they may be included here
under Rigodiella sect. nov. characterized as follows:

Folia caulina et ramea vix diversa; costa maxime variabilis, nunc simplex et bene
evoluta, nunc binis et brevis vel nulla.

HOOKERIACEAE

Sauloma tenella (H. f. & W.) Mitt.

Fuegia media: Rio Bueno, in silva Nothofagi, No. 2008.

New to South America. The broad distribution of so many austral types
which are common to New Zealand and the tip of the South American con-
tinent is again emphasized by the occurrence of this species in Fuegia.

AMBLYSTEGIACEAE

Hygroamblystegium: I think Reimers (Hedwigia 66:71. 1926) is entirely
justified in transferring the following species to H ygroamblystegium. They
are certainly better placed here than in Pseudoleskea.

BARTRAM: MOosSES FROM TIERRA DEL FUEGO 317

The group is a very intricate one and the distinctions so slight and
inconstant that I doubt if they can be resolved into definite specific identi-
ties. The large series collected by Dr. Roivainen seems to be separable into
two fairly distinct types as outlined in the tentative key presented below.
Leaves closely appressed

EGosely (usecase Plants © arier nae ek Maina cues cnet H. filum
Densely tufted plants, stems parallel ............- H. filum var. compactum
Leaves spreading or secund
Habit rigid, leaves strongly secund, uncinate at tips H. fuegianum var. secundum
Habit lax
Costa short excurrent

Were SICHOCTADIABUSi tee 85 Grants sas te ge H. fuegianum var. gracilis
More robust plants ‘
Leaves erect-spreading or secund ........ H. fuegianum
Leaves widely spreading ...............- H. fuegianum var. Skottsbergii
(Gusta done sexCOITente Ae eRe aah ONS sche ce oe H. fuegianum var. excurrens

Hygroamblystegium filum (C. M.) Reim.

Hygroamblystegium filum var. compactum Bartr. var. nov.

Caules elongati, ad 10 cm. longi, dense compacti, cellulae superiores breviter hexa-
gonae.

Chile: Prov. de Magellanes, Fuegia media, Lago Linch, Cerro Chico, reg.
alp., No. 1380 type.

An unusual form with long, parallel, densely ied stems and short
upper leaf cells.

Hygroamblystegium fuegianum (Besch.) Reim.

The following sporophyte characters are taken from No. 1016 which is
in good fruit.

Seta slender, reddish, smooth, about 2 cm. long, twisted to the right when
dry. Capsule cylindric, curved, constricted below the mouth when dry, urn
2 mm. long; peristome large, pale yellow; lid not seen; calyptra pale, early
deciduous; spores smooth, 10-12 » in diam.

Hygroamblystegium fuegianum var. Skottsbergii (Card.) Bartr. comb. nov.
Pseudoleskea fuegiana (Besch.) Card. var. Skottsbergit Card.

Hygroamblystegium fuegianum var. excurrens (Card. & Broth.) Bartr. comb. nov.
Amblystegium excurrens Card. & Broth.

Hygroamblystegium fuegianum var. gracilis (Card. & Broth.) Bartr. comb. nov.
Pseudoleskea fuegiana (Besch.) Card. var. gracilis Card. & Broth.

Hygroamblystegium fuegianum var. secundum Bartr. var. nov.

Caules rigidi. Folia saepe fortiter falcato-secunda.

Chile: Prov. de Magellanes, Fuegia media, Lago Fagnano, Punta Piz-
zaro, No. 1213 type.

Very different from any of the allied forms in the rigid, rather robust
stems, uncinate at the tips and with the leaves clearly falcate-secund.

Calliergidium austro-stramineum (C. M.) Bartr. comb. nov.
Hypnum austro-stramineum C. M. t

318 FaRLowIA, VoL. 2, 1946

Dr. Grout (Moss Fl. N, Amer. 3(2): 100. 1931) has advanced what seem
to be adequate reasons for separating this group from Drepanocladus.

Calliergon sarmentosum (Wahlenb.) Kindb.

Very similar to the plants of the northern hemisphere in every way except
that the apical leaves of the young branches and stems are more closely
imbricated to form a terete tip ending in a cuspidate point.

BRACHYTHECIACEAE

Brachythecium longidecurrens Bartr. sp. nov.

Autoicum; lutescente-viride, nitidulum, Caulis irregulariter ramosus, ramis arcuatis.
Folia dense conferta, scariosa, sicca imbricata, humida erecto-patentia, caulina e basi
longe decurrente latissime cordato-ovata, in acumen breviusculum constricta, 1.5-1.7
mm. longa; marginibus inferne reflexis, superne planis, fere e base minute denticulatis;
costa valida, ad 4/5 folii evanida; cellulis linearibus, inferioribus brevioribus et latio-
ribus, alaribus rectangularibus, pellucidis, Folia ramea angustiora, sensim acuminata.
Folia perichaetialia ecostata, in acumen patulum subito constricta; seta rubella, 2 cm.
longa, superne scaberula; theca horizontalis, sicca arcuata, 1.5 mm. longa, fusca.

Chile: Prov. de Magellanes, Fuegia media, Rio Bueno, in silva, Nos. 2018
type, 2025, 2063.

These plants have something of the habit of B. rivulare Bry. Eur. but are
immediately distinguished by the acuminate leaves, the longer costa, and
the setae which are smooth below.

Brachythecium (Julacea) micro-collinum Bartr. sp. nov.

Autoicum; tenellum, pallide viride. Caules intertexti, depressi, irregulariter pinnati,
ramis curvato-patulis. Folia minuta, sicca imbricata, humida erecto-patentia, concava,
ovato-lanceolata, acuminata, haud decurrentia, circa 0.5 mm. longa; marginibus toto
ambitu denticulatis; costa valida, ad 4/5 folii evanida, extremitate dentiformi; cellulis
linearibus vel lineari-rhomboideis, basilaribus laxioribus, rectangulis, infimis ovalibus
vel breviter rectangulis. Folia perichaetialia enervia, e base subvaginante in acumen
abrupte constricta, marginibus ad_ basin integris, superne argute denticulatis; seta
rubella, 6-7 mm. longa, laevissima; theca horizontalis, 1 mm. longa.

Chile: Prov. de Nuble, Recinto, in silva umbrosa, alt. 750 m., Nos. 954,
955 type.

Near B. collinum (Schleich.) but much smaller in every way and witha
stouter, longer costa ending in a spine on the back. B. arenarium Card. &
Broth. is described as having the stem leaves ending in a long, subulate
acumen. This character, if constant, should preclude any confusion with
B. micro-collinum.

PLAGIOTHECIACEAE

Plagiothecium nitidum (Hook. f, & Wils.) Bartr. comb. nov.
Leskea nitida Hook. f. & Wils.

I have outlined elsewhere (Manual of Hawaiian Mosses, p. 224. 1933)
my reasons for thinking that Catagonium is a poorly defined genus. The
above species does not appear to be generically distinct from Plagiothecium.

BARTRAM: MOSSES FROM TIERRA DEL FUEGO 319

Calliergonella complanata Card. & Broth.

To judge from a fragmentary specimen of the type collection consisting
of two stems and a few detached leaves which I have seen through the
kindness of Dr. Linkola, is apparently one of the forms of Eucatagonium
politum (Hook. f. & Wils.) with the leaves not clearly distichous.

SEMATOPHYLLACEAE

Sematophyllum aureo-nitidum Bartr. sp. nov.

Dioicum? Caespites depressi, sat densi, lutescente-virides, nitidi. Caulis ad 3 cm.
_ longus, irregulariter ramosus. Folia conferta, falcata, e basi ovato-oblonga abrupte longe
acuminata, ecostata, circa 2 mm. longa; marginibus planis, inferne integris, superne
denticulatis; cellulis anguste linearibus, incrassatis, infimis lutescentibus, admodum
porosis, alaribus 3-4, oblongis, vesiculosis, hyalinis vel lutescentibus. Folia perichaetialia
ovato-oblonga, breviter acuminata; marginibus superne irregulariter denticulatis; seta
rubella, laevissima, circa 12 mm. longa. Caetera ignota.

Chile: Prov. de Magellanes, Fuegia occid., Fjordo Martinez, ad rupes,
No. 579; Fjordo Martinez, Bahia Sarmiento, No. 2351 type.

More robust than S. callidum (Mont.) Mitt. from which it also differs in
the more glossy leaves hooked at the tips of the branches. The description
of Hypnum leucocytus C. M. suggests something similar but Dixon remarks
(Studies in the Bryology of New Zealand, Pt. VI, p. 309. 1929) that this
species is not glossy while S. aureo-nitidum is characterized by a lustrous,
almost metallic sheen.

POLYTRICHACEAE

Dendroligotrichum squamosum (Hook. f. & Wils.) Broth.

Numerous collections of this species show the end cells of the lamellae in
cross section are either two-parted or single and the lamellae 4—5 cells high.
The end cells have two lumens when two-parted and are scarcely suggestive
of Polytrichum commune to which Dixon (Studies in the Bryology of New
Zealand, Pt. IV, p. 235. 1926) indicates that it may have some affinity.

BuUSHKILL
PIKE Co., Pa.

2(3):321-343 FARLOWIA ‘January, 1946

THE GENUS CUNNINGHAMELLA (MUCORALES)
Victor M. CutTTER, JR.

The opportunity has recently arisen to examine a number of isolates of
Cunninghamella from various localities in North America. The confused
synonomy and overlapping specific descriptions available in the literature
have proved most troublesome in attempting to identify these isolates. It is
the author’s opinion that at the present time the accurate identification of
species of Cunninghamella is practically impossible. This situation arises
in large part from the failure of earlier workers to employ a standard re-
‘producible culture medium in their studies, and in part from the fact that
the known species have not yet been studied comparatively. Our expanding
knowledge of the physiology and genetics of the fungi makes it evident
that in such a plastic group as the Mucorales, specific concepts can be
formulated only from isolates grown on rigidly standardized media under
carefully controlled conditions. Descriptions based upon field collections
and gross cultures are of little value to the mycologist interested in accurate
determinations. Unfortunately for the advance of mycological science, the
precision of technique employed by physiologists in their studies of growth
and variation in fungi has too rarely been carried over into taxonomic
investigations.

METHODS

In common with most fungi, Cunninghamella species are very sensitive
to the most minute variations in the culture medium, and the same isolate
grown upon different media may present a completely different aspect.
This plasticity is no less evident at changing levels of temperature and
humidity. Any key to this genus can be of practical value only when the
material to be identified is handled under exactly comparable conditions.
Therefore the standardization of cultural techniques is imperative, and
towards this end the procedure described below is suggested as a standard
to be followed in the identification of species of Mucorales. It is not claimed
that the medium employed represents the best possible choice for cultural
studies or maintenance of mucoraceous fungi. Rather the medium is de-
signed to be easily and inexpensively reproduced and, when formulated
with CP chemicals, to be as uniform as possible. To date approximately
150 species of Mucorales have been grown upon this medium with good
results in all cases.

In connection with this procedure it is understood that all cultures should
originate from single spores. This is easily accomplished, in the case of
Cunninghamelia, by any of the usual techniques. The medium used in this
study is designated as M.D.A. agar and is made up according to the follow-
ing formula, and adjusted to pH 4.2 with 0.1 N HCl.

yal

322 FaRLow!A, Vor. 2, 1946

Mexirase: (4c. 4a02 5 40.00 gr. thiamine chloride . . . 0.5 mgr.
asparagine ....... 2.00 gr. agar flakes >. 0... o.:; 15.00 gr.
ROO hie eee ot a DF, distilled water . . . . 1000.00 cc.
PIgSOg 5 fs 'é oes 0.25 gr.

All reagents used are of CP grade. The thiamine used is Merck’s synthetic,
the asparagine is Bacto-Difco, and the agar is Bacto-agar flakes prepared
by the Difco laboratories.

In the routine of identification, isolates are grown upon M.D.A. agar at
25° C. and 40° C. Specific descriptions are based upon material cultured
at 25° C. At the 40° C. level, cultures are recorded merely as growing or
not growing. Smaller intervals of temperature than these have not proved
practical for the sharp delimitation of species. No attempt is made to
record degrees of growth since quantitative methods are rarely practical in
a taxonomic treatment, and do not lend themselves to rapid identification.

Material for microscopic examination is mounted in a solution of equal
parts of phenol crystals, lactic acid, glycerin and distilled water to which
a detergent, ‘‘Tergitol’’ Penetrant 7, is added in the ratio of 3 drops per
50 cc. of solution. To this mixture 5 percent Sudan III is added, which
furnishes an easy method of distinguishing chlamydospores and hyphal
bodies from conidia since the former rapidly accumulate the stain whereas
the latter remain unstained. The color of conidia and other spores is de-
termined by mounting them in water.

In connection with conidial measurements in this genus a word of cau-
tion is necessary. The shape and dimensions of conidia can not be accu-
rately determined until they are mature and have fallen from the vesicles.
In situ their shape is frequently obscured or distorted by crowding, in most
mounts only immature conidia remain attached to the vesicles, the mature
conidia being readily deciduous. Many of the aberrant measurements found
in the literature as well as the statements concerning supposed differences
in size and shape of the conidia of the terminal and lateral vesicles are
directly attributable to the failure to recognize this fact. The term “vesicle”
is used here in reference to the capitate structure terminating the conidio-
phores after the conidia are detached. The age of cultures determines to
some degree the type of branching displayed by the conidiophores and since
cultures are rarely mature until six days after inoculation, all observations
are made at this time.

Under the discussion of each species the original description is repro-
duced, followed by an emended description of the species based upon
monospore cultures grown at 25° C. on M.D.A. agar six days after inocula-
tion. The key to species is compiled from these emended descriptions.
Technical color terms cited in quotation marks are those of Ridgway (26).
The accession numbers given under the heading ‘‘material examined” refer
to cultures and permanent mounts filed in the author’s herbarium at Cornell
University.

CuTTER: THE GENUS CUNNINGHAMELLA . aco

HISTORY OF THE GENUS

The genus Cunninghamella was established by Matruchot (18) in 1903
upon C. africana collected in the French Sudan. At this time he failed to
provide a generic description which was later supplied by Saccardo (30).
Thaxter (36) shortly pointed out that in reality C. africana only repre-
sented a redescription of his Oedocephalum echinulatum, and that the spe-
cies should therefore be known as C. echinulata. Matruchot had postulated
that in spite of its purely conidial reproduction in the asexual phase, Cun-
ninghamella should be placed in the Mucorales, and supported his claim
with a series of ingenious experiments showing that Cunninghamella will
serve as a host for species of Piptocephalis which are parasitic exclusively
upon the Mucorales. He further indicated that the usually aseptate my-
celium favored the inclusion of Cunninghamella in this group. His opinion
was subsequently confirmed by Blakeslee (5) who discovered the zygo-
spores of C. echinulata in 1904. In the following year Saito (31) described
the genus Actinocephalum based upon A. japonicum. Fitzpatrick (11) has
pointed out that this genus is undoubtedly synonymous with Cunningham-
ella. In 1907 Lendner (14) described C. elegans in Swiss soil, and later, in
1927, he (16) described C. Blakesleeana from North American material
sent to him by Blakeslee. Stadel (32) in 1911 founded C. Bertholletiae
upon a strain from decaying Brazil nuts, and discussed certain aspects of
its growth and development upon various substrates. Shortly thereafter
Bainier & Sartory (2), apparently unaware of the existence of Cunning-
hamella, erected the genus Muratella upon M. elegans collected in France.
Torrey (37) showed that this species was synonymous with Cunningham-
ella echinulata Thaxter, and investigated the development and morphology
of the conidia in some detail. Naumov (20), however, on the basis of its
consistently spherical conidia, considered Muratella elegans distinct from
Cunninghamella echinulata and erected the new combination C. Bainieri to
include it and several of his own isolates. Ling-Young (17), on the other
hand, considered Muratella with its curious giant conidia to be distinct
from Cunninghamella and maintained it as a valid genus.

Paine (22) in 1927 described C. verticillata from the United States. This
species has been reported several times subsequently, but appears to be
synonymous with C. Blakesleeana Lendner, which had been described and
published three months before. Pispek (23) in 1929 discussed a series of
isolates from Yugoslav soils and erected four new species, C. ramosa, C.
dalmatica, C. polymorpha and C. echinata. None of these species has been
subsequently reported and all of them seem conspecific with previously
described forms, although PiSpek’s meager descriptions make positive
identification uncertain. In 1930, Fitzpatrick (11) briefly summarized the
work on Cunninghamella to that date. Zycha (38) in 1935 presented the
first comprehensive survey of the genus and the first key to the species. He
retained the six following species: C. microspora, C. albida, C. elegans, C.

324 FarLowIA, VoL. 2, 1946

ramosa, C. Blakesleeana and C. echinulata, and presented descriptions of
these compiled from the literature. All other described species were reduced
by him to synonomy. This treatment has served as a standard for most
subsequent work. Alcorn and Yeager (1) in 1938 published a monograph
of the genus with emended specific descriptions of species collected by them
in Idaho. This work contained a key to the species based in part upon the
literature and in part upon their own observations, as well as numerous
inaccuracies in the translations of original descriptions. While their mono-
graph purports to cover the entire literature on Cunninghamella, in reality
a number of species are not treated, in particular those of PiSpek and of
Zycha. The latest treatment of the genus is that of Naumov (20) in 1939.
He presents a descriptive key to all the known species compiled from the
original descriptions and his own observations. He retains all the pre-
viously described species and erects the new combination C. Bainieri
to include Muratella elegans. He has stated that this key is only tenta-
tive and the future studies will show that many of these species are not
valid.

Our knowledge of the sexuality of this genus and of the zygosporic stages
is largely confined to the works of Blakeslee (6), Burger (9) and Blakeslee,
Cartledge and Welch (7). The last named authors have shown that C. ele-
gans, C. Bertholletiae, C. Blakesleeana and C. echinulata are heterothallic
forms and are sexually dimorphic, a conclusion contrary to that advanced
by Burger on the basis of experiments with C. Bertholletiae. Blakeslee (6)
had previously described and figured the zygospores of C. echinulata, and
Lendner (16) supplied the same information for C. Blakesleeana. The
sexual stages of other described species are as yet unknown. All these
authors have indicated that zygospore formation is usually initiated at
temperatures above 25° C,

The systematic position of Cunninghamella has been treated in various
ways by different authors. Matruchot (18) considered it to belong in the
tribe Choanephoree in company with Choanephora. Lendner (15) places
it in the family Chaetocladiaceae in the suborder Conidiophorae. Fitz-
patrick (11), Zycha (38), Christenberry (10) and Ou (21) place it in the
family Choanephoraceae, with Blakeslea and Choanephora. However, Nau-
mov (20) creates the family Cunninghamellaceae to include Cunningham-
ella, Thamnocephalis and Sigmoideomyces. At the present state of our
knowledge the latter disposition seems most natural since this group of
genera forms a coherent unit well separated from Choanephora and Blakes-
lea by the complete absence of sporangia and the presence of echinulate
globose conidia. Furthermore, the dark colored rough-walled zygospores
set Cunninghamella apart from Blakeslea and Choanephora, which have
thin-walled, smooth, more or less hyaline zygospores.

It must be pointed out in connection with the validity of the name Cun-
ninghamella that this genus was founded upon the conidial or imperfect
stage of C. africana by Matruchot in 1903. The perfect stage (i.e. the
zygospores) was not discovered until 1904 by Blakeslee who described the

CuTTER: THE GENUS CUNNINGHAMELLA 625

zygospores in C. echinulata. These two species are synonymous. If the
strictest interpretation of the International Rules of Botanical Nomencla-
ture is followed (i.e. Art. 57) Cunninghamella must be attributed to Blakes-
lee, who, however, failed to give a generic description with his report of the
perfect stage of this fungus. On the other hand the International Rules fail
to define exactly what constitutes the perfect stage of a Phycomycete of the
mucoraceous type. Under proper conditions in homothallic species, and in
the case of heterothallic species such as Cunninghamella, in the presence of
two compatible strains, the gametangia and zygotes are borne upon the
same mycelium which gives rise to the asexual reproductive structures.
Since the perfect and imperfect stages are thus inextricably mingled it will
be seen that while these fungi are pleomorphic in regard to their life cycle,
there is no certain way of separating the two stages for taxonomic purposes.
This situation is somewhat different from the case of certain heterothallic
Ascomycetes and Basidiomycetes having pleomorphic life cycles where the
perfect and imperfect stages are clearly distinguishable. Bisby (4) has
pointed out other instances where the strict application of Article 57 to the
Phycomycetes creates nomenclatural problems of this nature. In the Mu-
corales dogmatic adherence to this rule would necessitate a large number of
changes in the author citations, and redescription of genera and species,
because the great majority of forms have been described in their imperfect
stages. Consequently in this and subsequent work on the Mucorales, Bisby’s
suggestions will be followed, and the first valid name applied to genus or
species will stand irrespective of whether it was applied in cases where the
zygotic stage was not known or described.

Cunninghamella Matruchot, Ann. Mycol. 1: 47-50. 1903; — Saccardo,
Syll. Fung. 17: 508. 1905.
Actinocephalum Saito, Bot. Mag. Tokyo 19: 1. 1904.
Saitomyces Ricker, Journ. Mycol. 12: 61. 1906.
Muratella Bainier & Sartory, Bull. Soc. Mycol. France 29: 129. 1913.

Fungi with erect or ascending conidiophores, abundant submerged sterile
mycelium, aseptate when young, becoming septate in age, hyaline or light
colored. Conidiophores variously branched or sometimes simple, each
branch terminating in a capitate vesicle covered with minute sterigmata
bearing conidia. Conidia unicellular, globose to ellipsoid, echinulate,
asperulate, or smooth, deciduous. Sporangia absent. Chlamydospores oc-
casionally present, intercalary, spherical to ellipsoid with granular or oily
contents. Rhizoids present in some species at base of conidiophores or
indiscriminately arranged on aerial parts. Gametangia isogamous. Zygo-
spores globose or doliform, dark-walled, tuberculate or roughened. The
known species are heterothallic and saprophytic on a wide variety of
substrates.

The type species is Cunninghamella echinulata (Thaxter) Thaxter.

326 FaARLOwIA, VoL. 2, 1946

KEY TO THE SPECIES OF CUNNINGHAMELLA
(i eeeMen Uma ACs Ce ci eee ek te Rae Sots Buh ais gins eg met a ta or aed bs
2. Colonies “Smoke Grey”; branches of the conidiophores sympodial, cymose or
irregular, not verticillate; conidia predominantly spherical with long echinula-
tions; giant conidia absent pepe ett BMPs irs =e Mean eee .1. C. echinulata.
2. Colonies “Gull Grey,” branches of the conidiophores verticillate; conidia pre-
dominantly ovoid to ellipsoid with short echinulations. ...4. C. Bertholletiae.
Be GT een at a” Ae ete ts Vg 0 dad eure ee ee eae ce
3. Giant conidia present; colonies “Smoke Grey” to “Pale Olive Buf”; conidia
predominantly spherical with long echinulations or sometimes smooth.
2. C. Bainieri.
3. Giant conidia absent; conidia predominantly ovoid to ellipsoid with short
re AVE PE LEW | te gee RSE AL ge a Re eR ROT Ai re Joh rare ane ee 4,
4. Colonies “White”; branches of the seueliantaes dichotomous, sympodial or
verticillate ; shteoidal tufts apparent macroscopically; conidia up to 35x 25 pw.
3. C. Blakesleeana.

4. Colonies “Gull Grey”; branches of the conidiophores always verticillate;

conidia up to 16x 12 w; rhizoidal tufts not apparent macroscopically. ....... 5.
5. Chlamydospores usually present; mycelium containing much oil; habitat
| ig Tap LT dp ee ree ie Oe SC ae On EEN 4. C. Bertholletiae.

5. Chlamydospores absent; mycelium mostly lacking oil; habitat soil and seeds.
5. C. elegans.

1. Cunninghamella echinulata (Thaxter) Thaxter, Rhodora 5: 98. 1903.
Ocedocephalum echinulatum Thaxter, Bot. Gaz. 16: 17. 1891.
Cunninghamella africana Matruchot, Ann. Mycol. 1: 47-50. 1903.

Prate I, A-I

“White becoming slightly yellowish. Sterile and fertile hyphae not clearly differen-
tiated, the latter more or less irregularly and indefinitely branched. The fertile heads
nearly spherical to obovoid, very variable in size, more or less distinctly areolate:
maximum 45-65 uw: average 28-35 uw. Spores oval to elliptical, finely echinulate: average
10x 12 #; maximum 18-25 uw.” — Thaxter (1891).

On M.D.A. agar:

Colonies up to 1.5 cm. high, spreading rapidly, “Smoke Grey,” powdery
in appearance; growing at 40° C. Sterile mycelium smooth-walled, much-
branched, mostly aseptate, up to 20 » in diam.; rhizoids occasional; chla-
mydospores occasional upon submerged portions, irregular in shape and
size. Conidiophores dichotomously or cymosely, never verticillately,
branched; each branch terminated in a swollen vesicle, broadly clavate to
subspherical, up to 60 p» in diam., average 25-40 » in diam.; smooth or
faintly areolate, no conspicuous difference in shape of lateral and terminal
vesicles. Conidia globose to ovoid up to 20 x 18 », average 11-14 yw in diam.,
thickly beset with very fine spines up to 4 » in length, the immature conidia
sometimes smooth. Zygospores described by Blakeslee (6) and not sub-
sequently reported. Gametangia isogamous or heterogamous; zygospores
nearly opaque, 46-80 x 40-63 p, average 70 x 58 yu, thickly beset with long
heavy tubercles. Zygospore formation taking place only at temperatures
above 25° C.

TYPE LOCALITY: On cheese and cheesy paper, Massachusetts. Thaxter.

CuTTER: THE GENUS CUNNINGHAMELLA PAT

DISTRIBUTION: Camel dung, French Sudan, 1903, Matruchot; dried
flowers, Venezuela, 1905, and Puerto Rico, 1905, horse dung, Philippines,
1905, Blakeslee; Brazil nuts and soil, (18 races), New York, 1921, Blakes-
lee, Cartledge & Welch; soil, Dalmatia, 1932, Johann; soil, Bastrop Co.,
Texas, 1932, Morrow; soil, Cuba, South Carolina, North Carolina, Louisi-
ana, 1940, Christenberry; decaying flowers of Cucurbita pepo, Szechuan,
China, 1940, Ou.

MATERIAL EXAMINED: Panama, C. Z., date and substrate unknown, Weston (Acc. no,
110) ; locality unknown, via Harvard Cult. Coll. (Acc. no. 127); Ontario, peas, 1941,
Groves, (Acc. no. 152); Arnprior, Ont., peas, 1943, Groves, (Acc. no. 294); Ithaca,
N. Y., contaminant in culture, 1943, Cutter, (Acc. no. 345); locality unknown, from
Brazil nuts, Blakeslee, (Acc. nos. 374, 375); locality, date and substrate unknown,
Blakeslee, (Acc. nos. 376, 377, 378, 379).

The type of branching is the most important character separating C.
echinulata from the other members of this genus. The descriptions and
figures of Thaxter and Matruchot indicate that they were not dealing with
a form having verticillately branched conidiophores. Plate I, A, B, D, and
E shows the typical cymose branching encountered in vigorous cultures. In
old cultures, pseudoverticillate conidiophores may be developed through
the production of secondary conidiophores on the primary conidial head.
A similar situation may occur if the conidiophores are submersed, as in the
example shown in Plate I, G. This is, however, an abnormality. The author
does not follow Zycha (38) in considering Muratella elegans Bainier &
Sartory, and C. verticillata Paine as synonyms of C. echinulata inasmuch
as these two forms are described and figured with verticillately branched
conidiophores while a study of the present material of C. echinulata indi-
cates that it is remarkably constant in its cymose branching.

Some slight discrepancies exist between Matruchot’s description of C.
africana and Thaxter’s description of Oedocephalum echinulatum, chiefly
in the dimensions of the conidial heads, which in C. africana are said to
reach 100 » in diameter. However according to Thaxter (36) the two
forms are synonymous and in general the descriptions coincide well. Nau-
mov (20) retains C. africana and makes C. echinulata a synonym of it,
which is an untenable disposition when Thaxter’s opinion that C. africana
merely constituted a redescription of his O. echinulatum is considered.

Alcorn & Yeager’s description of C. echinulata contains the statement
that the vesicles are areolate, an opinion which probably arises from a liberal
interpretation of Thaxter’s more cautious statement that the heads are
sometimes faintly areolate. In the cultures studied here the areolate con-
dition could sometimes be discerned but it certainly was not present in the
majority of isolates and it was usually characteristic only of immature
vesicles. ie

C. echinulata is the only species of Cunninghamella which in this study
consistently showed growth at temperatures above 40° C. This character-
istic, and the absence of the peculiar giant conidia described below, distin-
guish it clearly from the following species.

328 FaRLOWIA, VOL. 2, 1946

2. Cunninghamella Bainieri N. Naumov, Clés des Mucorinées, Paris, 1939.
Muratella elegans Bainier & Sartory, Bull. Soc. Mycol. France 29: 9. 1913.
Muratella sp. Ling-Young, Rev. Gen. Bot. 42: 744. 1930.

Prate I, J-O

“Conidia all spherical 14-17 » in diam. armed with bristles 2 w in length; sterigmata
0.5-1.4 « long; colonies dirty white, 1 cm. high or more; verticils of 3-8 elements,
accompanied as well by longer branches of the conidiophores; swellings of 28-58 u in
diam.” Naumov.

On M.D.A, agar:

Colonies 0.5—-3 cm. high, ‘Pale Olive Buff” to “Pale Smoke Grey,” pow-
dery in appearance, not growing at 40° C. Sterile mycelium much-branched,
up to 30 » in diam., aseptate when young, in the submerged portions fre-
quently swollen into vesicular shapes; true chlamydospores absent. Co-
nidiophores erect, usually verticillately branched below the terminal head,
sometimes with several whorls of branches, lateral branches usually less
than 50 » long. Vesicles up to 60 » in diam., average 40 p, lateral vesicles
usually smaller than the terminals. Vesicles bearing normal conidia gen-
erally smooth, sometimes verrucose when young, vesicles bearing giant
conidia markedly verrucose. Conidia of two sorts, predominantly spheri-
cal, sometimes broad ovoid; normal conidia hyaline 5—15 p in diam. with
slender sharp echinulations up to 3.5 » long, giant conidia yellow to brown-
ish, thick-walled, 15-27 » in diam. with stout sharp echinulations up to
2 w long or sometimes smooth, giant conidia usually borne on verrucose
subterminal vesicles developing in older portions of the colony. Zygospores
unknown.

TYPE LOCALITY: Dead wood, France, Bainier & Sartory, 1913.

DistriBuTion: Soil, France, 1930, Ling-Young.

MATERIAL EXAMINED: Hibiscus flowers, Ann Arbor, Mich., 1940, Cutter (Acc. no.
29) ; locality and substrate unknown, 1942, via American Type Cult. Coll. (Acc. nos.
348, 352, 354); locality, date and substrate unknown, C. B. S. via Blakeslee (Acc.
nos. 386, 387).

This species, which is closely related to C. echinulata, may be easily
distinguished by the presence of numerous giant conidia in older cultures,
as well as by its failure to grow at 40° C. Bainier & Sartory (2) give its
optimum temperature range as 25—27° C, Ling-Young (17) has discussed
the morphology of the “conidies tardives” or giant conidia, of this species,
and states that they are always borne upon specialized lateral heads. This
contention is not strictly borne out in the material studied here where the
giant conidia are frequently interspersed with smaller conidia on both ter-
minal and lateral vesicles. However, Plate I, N shows the usual condition
where these structures are confined to certain lateral vesicles. A study of
Ling-Young’s illustrations show that they were based upon immature
conidiophores which may account for the peculiar distribution of his
“conidies tardives.” These structures have not been encountered in other
species of Cunninghamella.,

CuTTER: THE GENUS CUNNINGHAMELLA 329

3. Cunninghamella Blakesleeana Lendner, Bull. Soc. Bot. Geneva 19: 234. May,
1927.
Cunninghamella “A,” Blakeslee, Cartledge & Welch, Bot. Gaz. 72: 185. 1921.
Cunninghamella verticillata Paine, Mycologia 19: 253. Sept. 1927.
Cunninghamella echinata Pigpek, Acta Bot. Inst. Univ. Zagreb. 4: 101. 1929.

Prate ITI, A-D

“Conidiophores branched, terminated by swollen heads; heads minutely verrucose,
30 w long by 28 w wide, bearing ellipsoid conidia with finely echinulate walls. Branches
of the conidiophore verticillate or solitary, lateral branches bearing spherical, hyaline to
yellow conidia; zygospores spherical to doliform, dusky black, 40 « in diam. up to
48 uw long by 49 w wide.” Lendner.

On M.D.A. agar:

Colonies 2 to 3.5 cm. high, spreading rapidly, “White,” at maturity be-
coming tinged with yellow and compactly matted, not growing at 40° C.
Sterile mycelium at first dichotomous, later irregularly branched up to 20 p
in diam., aseptate, bearing prominent tufts of rhizoids or outgrowths visible
macroscopically upon the aerial portions, occasionally forming vesicular
swellings containing much yellow oily material upon the submerged por-
tions; true chlamydospores lacking. Conidiophores erect, sometimes di-
chotomously or sympodially but usually verticillately branched. Lateral
branches of the conidiophores variable in length and number, usually less
than 50 » long. Vesicles variable, subglobose to broadly clavate up to 65 p
in diam., average 30-40 y» in diam., lateral vesicles usually smaller than
terminal vesicles, verrucose to smooth. Conidia hyaline, predominantly
ovoid to ellipsoid with smaller subspherical conidia interspersed, up to 35 p
long x 25 » wide, average 5-18 » long x 4-12 p» wide, with very short widely
spaced echinulations or sometimes smooth. Zygospores not seen.

TYPE LOCALITY: Unknown.

DISTRIBUTION: Brazil nuts purchased at many localities in eastern
U.S.A. (all nuts presumably from S. America), 53 races, 1921, Blakeslee,
Cartledge & Welch; soil, Iowa City, Iowa, 1927, Paine; soil, Colorado,
1930, LeClerg; soil, Grobnik, Krk Island in Adriatic Sea, 1929, PiSpek.

MATERIAL EXAMINED: Brazil nuts bought in eastern U. S. A. (all nuts presumably
from S. America), 1919 (7 races) Blakeslee (Acc. nos. 360, 361, 362, 363, 364, 365, 366).

Cunninghamella Blakesleeana was described by Lendner (16) in 1927
with no data given concerning the substrate or locality from which it was
collected beyond a brief note to the effect that the material had been for-
warded to him by Blakeslee. Previously Blakeslee, Cartledge & Welch (7)
had investigated and discussed the sexual condition in an undescribed
species which they designated as Cunninghamella “A.” At this time they
pointed out that in many respects Cunninghamella “A” was intermediate
between C. echinulata and C. Bertholletiae, the appearance of the colony
recalling C. echinulata, but the aspect of the spores and conidiophores more
closely approaching C. Bertholletiae. Lendner’s description of C. Blakes-
leeana emphasized this situation, and at the start of this study it was as-
sumed that Cunninghamella “A” and C. Blakesleeana were synonymous.

330 FaRLOWIA, VoL. 2, 1946

Recently in a personal communication Professor Blakeslee has informed
the author that his material of Cunninghamella “A” did indeed constitute
the type material of C. Blakesleeana.

The cultures studied here which were obtained from the Blakeslee collec-
tion agree well with Lendner’s description except that the conidia of these
isolates reach somewhat larger dimensions than those given by Lendner.
It is not unusual to encounter outsize conidia up to 35 » long. Furthermore,
in this material certain outgrowths of rhizoids or sterile mycelium upon the
aerial hyphae were very apparent, a feature not mentioned in Lendner’s
description. The presence of these rhizoidal tufts upon the mycelium in the
aerial portions of the colony imparts a very characteristic appearance dur-
ing the early stages of growth, and in conjunction with the pure white color
of the colonies provides an easy macroscopic method of distinguishing this
species from C. elegans and C. Bertholletiae.

The frequently irregular or pseudoverticillate branching of the conidio-
phores and the more heavily echinulate conidia are further differences
between C, Blakesleeana and the above-mentioned species. Conidial meas-
urements and shapes of these forms tend to overlap and do not seem re-
liable criteria for their separation. The data presented by Blakeslee, Cart-
ledge & Welch based upon a study of the sexual conditions in this form and
in C. elegans and C. Bertholletiae indicate that it is physiologically distinct
from the latter species.

The zygospores are described by Lendner and are said to differ from
those of C. echinulata and C. elegans in the shape of the segments of the
epispore which are somewhat depressed or hollowed in the center, rather
than flat as in C. elegans or tuberculate as in C. echinulata.

Cunninghamella verticillata Paine (22), described three months after
C. Blakesleeana, corresponds so closely with it that it must be incorporated
here. The description of C. echinata Pispek fits C. Blakesleeana so well that
it seems probable the two species are synonymous although Pi§pek’s de-
scription leaves much to be desired. Since C. echinata has never been re-
ported subsequently and since there are no significant differences between
the description of it and of C. Blakesleeana, PiSpek’s species is here reduced
to synonymy. :

In common with C. Bertholletiae, this species occurs predominantly upon
substrates with high oil or fat content, as is indicated by its repeated isola-
tion from Bravil nuts. A detailed study of a large series of isolates from
this and other substrates may reveal strains intermediate in character
between this species and C. Bertholletiae and C. elegans which will necessi-
tate the combining of this complex into one inclusive species. On the basis
of our present knowledge it pRESa desirable to maintain them as distinct
entities.

4, Cunninghamella Bertholletiae Stadel, Dissertation, Kiel, pp. 1-35. 1911.
PrLateE I], E-G

“Colonies white, yellow to light olive, dense with interwoven mycelium, up to 2 cm.
high. Conidiophores verticillately branched, the laterals sometimes showing secondary

CuTTER: THE Genus CUNNINGHAMELLA shat

branching. Conidiophores 9-19.2 w thick, with heads up to 70 w in diam., average
40-50 «; swellings 5x4 u up to 30-55 uw in diam. Conidia oval or having the form of
a cucumber seed, echinulate, 16-25 x 10-12 yu, or spherical, more nearly smooth 8-9 wu
in diam. Sterigmata up to 2.75 x 0.8 uw. Chlamydospores present, up to 30-40 uw in diam.
Oily material usually present in mycelium, conidiophores and conidia. Species showing
an affinity for oily substrates.” Stadel.

Zygospores have been reported by Blakeslee, Cartledge & Welch (7) but
never described.

On M.D.A. agar:

Colonies up to 2.5 cm. high, spreading rapidly, “Gull Giey? somewhat
compacted, sometimes growing at 40° C.; sterile mycelium much branched
with some spherical to ovoid chlamydospores up to 35 » in diam. containing
much oily or granular material; conidiophores erect, verticillately branched
at some distance below the terminal heads, with as many as five whorls of
lateral branches; laterals usually more than 50 p» long; vesicles subglobose
to clavate, when young tuberculate or verrucose, in age becoming smooth,
up to 65 » in diam., at times filled with large oil globules; conidia character-
istically lacrymoid, sometimes subspherical or ellipsoid, with prominent
apiculus, up to 16x12 yw, average 9x7 yp», hyaline, echinulate with very
short spines or asperulate: zygospores not seen.

TYPE Locality: Nuts of Bertholletia excelsa, Brazil, 1911. Stadel.

DISTRIBUTION: Brazil nuts, Massachusetts, 1919, Burger; New York
and localities in eastern U.S.A., 81 races (all nuts presumably collected in
S. America) 1921, Blakeslee, Cartledge & Welch; soil, Moscow, Idaho,
1937, Alcorn & Yeager.

MATERIAL EXAMINED: Brazil nuts, locality unknown via American Type Cult. Coll.,
1943, (Acc. nos. 95, 96, 353); via Blakeslee coll., 1943, (Acc. nos. 367, 368, 369, 370,
371, 372, 373).

Lendner (16) calls attention to the fact that this species is very close to
C. elegans, and considers them synonymous. Zycha (38) follows this treat-
ment, whereas Naumov (20) and Alcorn & Yeager (1) retain C. Berthol-
letiae as a specific entity. In this connection it should be recalled that
Blakeslee, Cartledge & Welch (7) in their extensive work on the ‘sexuality
of Cunninghamella apparently experienced no difficulty in separating the
two species and obtained no evidence that they were conspecific. Further-
more with the exception of Alcorn & Yeager’s record, C. Bertholletiae has
been reported only from decaying Brazil nuts while C. elegans is predomi-
nantly a soil inhabiting form. In the case of the race of C. Bertholletiae
collected from Idaho soil by Alcorn & Yeager, their emended description
does not coincide at all well with the original description. Furthermore
their translation of Stadel’s description contains certain inaccuracies which
may arise from the fact that they cite as the source of this diagnosis not

. Stadel’s own work, but only a review of it prepared by Leeke (13). There-
fore the only valid evidence that the Idaho form is actually C. Bertholletiae
is the fact that it was reputed to form zygospores when contrasted with a

332 FARLowIA, VoL. 2, 1946

minus race of this species in Blakeslee’s laboratory. Under the circum-
stances this record must be viewed with some scepticism.

When cultured upon M.D.A. agar there are certain minor differences
between the two species. C. Bertholletiae usually develops chlamydospores,
whereas these structures are lacking in C. elegans. In all the material ex-
amined here the mycelium of C. Bertholletiae was consistently filled with
granular materials or oil globules, a condition not seen in C. elegans. The
conidia of the former are more consistently apiculate than are those of the
latter, and the lateral branches of the conidiophores of C. Bertholletiae are
on the average somewhat shorter and stouter. It must be emphasized, how-
ever, that these are not reliable criteria for the separation of the two species
and are useful only in connection with a large series of isolates.

Perhaps the most significant difference between the two species is the
ability of certain strains of C. Bertholletiae to grow at 40° C. No strains
of C. elegans were encountered which could endure this temperature. Four
of the ten cultures of C. Bertholletiae studied here exhibited this peculi-
arity. This variability between morphologically identical strains cannot be
satisfactorily accounted for at the present time. However, all the cultures
studied had been grown upon artificial media for a number of years before
they were tested, and it is possible that during this time certain of them lost
the ability to grow at higher temperature levels. While this speculation is
in no sense conclusive, the temperature growth relationships of these species
may prove very significant in effecting a satisfactory separation, and should
be investigated in greater detail with freshly collected cultures. The only
other species of Cunninghamella consistently able to grow at temperatures
of 40° C. or higher is C. echinulata. In the key presented above C. Berthol-
letiae has been so placed that it may be identified in either temperature
category.

In the light of the results of Blakeslee, Cartledge & Welch, and the fail-
ure in the present study to obtain zygote formation between oppositely
sexed races of these two species contrasted under favorable conditions of
temperature and nutrition, it seems reasonable to conclude that they are at
least physiologically distinct. Consequently C. Bertholletiae is retained
until more cogent proof of its conspecificity with C. elegans is presented.

5. Cunninghamella elegans Lendner, Bull. Herb. Boissier (II) 7: 250-251. 1907;
Les Mucorinées de la Suisse. Berne, 1908.

PLATE II, H

“Colonies white to light ashy, mycelium differentiated into two regions, the sub-
merged region very compact, in age becoming pseudoparenchymatous and cartilaginous,

dichotomously several times, aseptate, verticillately branched at a distance below the
terminal head. Verticillate branches variable in number, terminated in heads. Terminal
heads up to 60 w in diam., lateral heads 18-20 uw in diam. Conidia of the terminal head
ovoid, apiculate at point of insertion, 16x 12-14 w up to 14x22 mw, armed with very
short spines. The lateral conidia smaller and more nearly spherical, 8-10 « in diam.
The conidia impart a pale ashy blue color to the colony. After dissemination of the

o

CuTTER: THE GENUS CUNNINGHAMELLA 335

‘conidia the surface of the heads is echinulate with short tubercles, very evenly
distributed, marking the points of insertion of the conidia.” Lendner (1908).

Zygospores reported by Blakeslee, Cartledge & Welch (7) but never
described.

On M.D.A. agar:

Colonies up to 3.5 cm. high, spreading rapidly, “Gull Grey,” not growing
at 40° C., in age becoming compacted. Sterile mycelium much branched,
up to 20 » in diam., aseptate when young, gradually becoming septate in
the submerged portions, frequently forming oidia and swellings, true chla-
mydospores not observed. Conidiophores erect, verticillately branched at
some distance below the terminal head; lateral branches arranged in a
number of successive whorls about the primary conidiophore, laterals usu-
ally more than 50 » long and comparatively slender. Terminal vesicles
subglobose to broad clavate, up to 60 p» in diam., lateral vesicles pyriform,
subglobose or clavate, up to 40 » in diam.; vesicles for the most part very
closely verrucose or tuberculate, sometimes smooth. Conidia hyaline, pre-
dominantly ovoid to ellipsoid, occasionally lacrymoid, up to 14-16 p» long
x 10-12 p» broad, echinulate with very short spines, asperulate, or smooth.
Zygospores not seen.

Type Locality: Garden soil, near Geneva and Vuache, Switzerland,
Lendner.

DistTRIBUTION: Soil, Cold Spring Harbor, N. Y., (42 races) 1921, Blakes-
lee, Cartledge & Welch; Germany, 1929, Janke; South Slavia 1929,
PiSpek; Evanston, Illinois, 1929, Swift; France, 1930, Ling-Young; Bas-
trop Co., Texas, 1932, Morrow; Germany, 1935, Zycha; Ann Arbor, Michi-
gan, 1914, 1917, Povah; Moscow, Idaho, 1937, Alcorn & Yeager; Canada,
1938, Bisby; North Carolina (several localities), 1940, Christenberry.

MATERIAL EXAMINED: Peas, Wellington, Ont., Arnprior, Ont., Brittania Bay, Ont.,
Canada, 1943, Groves (Acc. nos. 141, 312, 316) ; flaxseed, Richmond, Ont., Saskatoon,
Sask., Winkler, Man., Canada, 1943, Groves (Acc. nos. 209, 224, 296); locality and
substrate unknown, via American Type Cult. Coll. and S. A. Waksman, 1943 (Acc. nos.
349, 355, 357) ; locality and substrate unknown via Blakeslee Coll., 1943 (Acc. nos. 380,
381, 382, 383, 384, 385); parsley seed, New Haven, Conn., and Gilroy, Calif., 1942,
Groves (Acc. nos. 388, 390) ; lettuce seed, Philadelphia, Pa., and San Francisco, Calif.,
1942, Groves (Acc. nos. 389, 398) ; spinach seed, Toledo, Ohio, and Milford, Conn., 1942,
Groves (Acc. nos. 395, 396); pepper seed, Toledo, Ohio, 1942, Groves (Acc. no. 391) ;
pumpkin seed, Milford, Conn., 1942, Groves (Acc. no. 392) ; celery seed, Detroit, Mich.,
1942, Groves (Acc. no. 393) ; muskmelon seed, New Haven, Conn., 1942, Groves (Acc.
no. 394); carrot seed, Minneapolis, Minn., 1942, Groves (Acc. no. 397).

This species is common in seeds and soil throughout the temperate re-
gions of the world. As yet no tropical records are available. The small,
asperulate or shortly echinulate conidia and consistently verticillate branch-
ing of the conidiophores, with long, slender, Jateral branches, make Ce
elegans easily distinguishable from all other described species except C.
Bertholletiaec. The distinguishing features of these two species have been

eh.

334 FartowiA, Vor. 2, 1946

previously outlined. Lendner has described the peculiar vesicular swellings
formed in liquid media by this species. The same phenomenon is exhibited
by all isolates of Cunninghamella under unfavorable conditions. It must be
recognized that these structures are not true chlamydospores, which in the
material examined by the author occur only in C. echinulata and C. Berthol-
letiae. These chlamydospores are thick-walled resistant bodies, intercalary
in the mycelium, while the vesicles merely represent thin-walled swollen
portions of the submerged mycelium,

DOUBTFUL SPECIES

Cunninghamella albida (Saccardo) Matruchot, Ann. Mycol. 1: 56. 1903.
Oedocephalum albidum Saccardo, Michelia 2: 288, 1881. —
Haplotrichium albidum Saccardo, Michelia 2: 288. 1881 and Fungi Italici: fig. 805.
1881.

“Sterile mycelium repent, much branched, aseptate, hyaline. Conidiophores erect or
decumbent similar to the sterile mycelium, aseptate and hyaline, simple or occasionally
giving off lateral conidiophores terminated by finely muriculate vesicles. Conidia arising
from the tubercles of the vesicles globose to globose elliptical (not concatenate) 2-10 uw
in diam., hyaline white to pale vellow.” Saccardo.

In dead and decaying roots of Citrus Limonii Salo, Italy, 1874.

As far as is known this species has never been collected since the original
report. There seems little doubt that Saccardo was dealing with a species
of Cunninghamella but the figure and fragmentary description given offer
little possibility of future identification. Zycha and Naumov have inter-
preted C. albida as having unbranched conidiophores and use this character
to separate it from other species of Cunninghamella. This interpretation
arises from a liberal translation of the original Latin description. On the
other hand, Saccardo’s illustration shows a form with verticillate and
dichotomously branched conidiophores, as well as several young unbranched
conidiophores. Were it not for the smooth-walled conidia the species would
conform well with C, Blakesleeana. However, in the absence of further in-
formation concerning the essential dimensions of this species, any specula-
tion in regard to its synonymy appears unwarranted. It is therefore retained
as a provisional species but not included in the key.

Actinocephalum japonicum Saito, Bot. Mag. Tokyo 19: 1. 1904.
Saitomyces japonicus (Saito) Ricker, Journ. Mycol. 12: 61. 1906.

“Turf grayish, mycelium irregularly branched covering the substrate; sporangiferous
hyphae erect, lacking basal rhizoids, in general verticillately branched, branches termi-
nally inflated, vesicles 25-55 w in diam.; conidia globose or sometimes oval, 20 by
18-21 yw, inserted individually on sterigmata, hyaline, echinulate, zygospores and
chlamydospores unknown.” Saito.

TYPE Locaity: Air borne, Musashi Province, Japan.

This species placed in Cunninghamella by Fitzpatrick (11) and subse-
quently by Zycha (38) and Naumov (20), appears identical with C, Bai-
niert. Zycha and Naumov consider it synonymous with C. elegans Lendner,
but the short verticillate lateral branches shown in Saito’s illustrations and

Cutter: THE GENUS CUNNINGHAMELLA a5

the large finely echinulate subspherical conidia are not consistent with the
concept of C. elegans. In the absence of the original material it will prob-
ably never be possible to determine its specific affinity.

Cunninghamella dalmatica PiSpek, Acta Bot. Inst. Univ. Zagreb. 4: 100. 1929.

“Colonies brownish white, 1 cm. high, conidiophores mostly verticillately branched
immediately below the apex, terminal heads 28-52.5-56 mw in diam., lightly verrucose;
lateral heads 14-24.5 w diam.; conidia globose 7-8 « diam. to opal 7-14 wu long by
7-10.5 « wide.” PiSpek.

TYPE LOCALITY: Cultivated soil near Jesenice, Dalmatia.

MATERIAL EXAMINED: Locality, date, and substrate unknown, C.B.S. via Blakeslee
(Acc. no. 386).

The incomplete description given by PiSpek will not allow this species to
be positively identified. The culture received under this designation from
Professor Blakeslee’s collection is believed to represent a subculture of
Pigpek’s type. Upon examination it proved identical with C. Bainieri. This
cannot be accepted as conclusive evidence that these two forms are synony-
mous because of the possibility of contamination during transfer of the
cultures. On this basis, and because of the incomplete diagnosis furnished
by PiSpek, there is no justification for maintaining C. dalmatica as a valid
species.

Cunninghamella ramosa PiSpek, Acta Bot. Inst. Univ. Zagreb. 4: 102. 1929.

“Colonies yellow-brown, 2 cm. high, externally resembling certain Absideae. Conidio-
phores in the middle of the culture for the most part simply branched, frequently
inflated at the base, those on the outer periphery of the culture abundantly sympodially
or irregularly branched; terminal heads for the most part verrucose 42-59-77 yw in
diam.; lateral heads 14-38 « wide; spores mostly globose, very large, yellowish, granu-
lar within, not echinulate, 10.5—24.5-35 mw in diam. or globose and pyramidal, 10.5-14
u long by 7-9-10.5 «4 wide; chlamydospores not seen, zygospores unknown.” PiSpek.

TYPE LocaLity: Rich pasture soil near Kraljevo (Serbia occid.).

On the basis of PiSpek’s description this species appears closely related
to C. Bainieri. The only significant differences are the irregular branching
of C. ramosa and its predominantly smooth-walled conidia. However,
Plate I, M shows an irregularly branched conidiophore of C. Bainieri and
smooth-walled giant conidia are not infrequently found in this species. The
large dimensions of the conidia of PiSpek’s isolate certainly indicate a close
connection with C. Bainieri. This species has never been reported subse-
quently, and until more information can be accumulated its validity must
be questioned.

Cunninghamella polymorpha Pispek, Acta Bot. Inst. Univ. Zagreb. 4: 103. 1929.

“Colonies brownish white, dense, 2 cm. high, conidiophores very diversely branched;
terminal heads verrucose, 32-49-55 yw in diam.; lateral heads 10.5-17.5-25 mw in diam.;
spores globose 7-14—(17.5) mw in diam., or ovoid to ellipsoidal 7-10.5—(14) « wide by
7-10-(17) yw long, hyaline or yellowish; chlamydospores not seen, zygospores un-
known.” PiSpek.

336 FarLowlé, VoL. 2, 1946

TYPE LOCALITY: Soil between rocks on Mt. Puvarov, elevation 1300
metres, Croatia.

MATERIAL EXAMINED: Locality, date, and substrate unknown. C.B.S. via Blakeslee
(Acc. no. 387).

The isolate of this species received from Professor Blakeslee’s collection
was identical with isolates of C. Bainieri. The description furnished by
PiSpek indicates that the measurements of his type isolate overlap those of
C. dalmatica, and he presents no significant data which will permit the
separation of these two species. On the basis of his description positive
identification of C. polymorpha is impossible.

EXCLUDED SPECIES

Cunninghamella microspora (Riv.) Matruchot, Ann. Mycol. 1: 56. 1903.

Gonatobotrys microspora Rivolta, Dei Parassiti Vegetali, p. 490. Torino, 1873.
Prachtflorella microspora (Riv.) Matruchot, Ann. Mycol. 1: 56. 1903.

This species, placed in Cunninghamella by Matruchot, has been incor-
porated in the genus by all subsequent authors, although as far as can be
determined it has been reported only by Rivolta in 1873 and by Berlese in
1889. Rivolta’s description consists of the single line “La Gonatob. micro-
spora. — Ha le spore piccolissime.”, and the single figure given is too frag-
mentary to give any real concept of the species. Knowledge of the form is
therefore confined to the descriptions and figures presented by Berlese from
material encountered on dead and decaying bark of Morus alba in Northern

from any of the forms discussed above. The restricted nature of the colony,
the extremely small dimensions of the mycelium (2-3 » in diam.), and
particularly the presence of unbranched conidiophores with intercalary
spore-bearing nodes are characters not associated with the original concept
of Cunninghamella as outlined by Thaxter and Matruchot. Furthermore,
the elongate, basally apiculate, smooth conidia do not correspond well with
the usual conidial type in Cunninghamella. Matruchot’s comments on the
relationships of C. microspora and his proposal of the alternate name
Prachtflorella microspora indicate that his disposition of it was provisional,
and subsequent studies have not yet confirmed his opinion. In the absence
of any knowledge of the sexual stage of this species it seems advisable to
exclude it from Cunninghamella.

Cunninghamella mandshurica Saito & Naganishi, Bot. Mag. Tokyo 39: 285. 1913.

This species has been shown to belong in Choanephora by Fitzpatrick
(11). The longitudinally striate conidia exclude it from Cunninghamella,
although the type of branching is in some respects intermediate between
the two genera. Zycha (38) considers the species synonymous with
Choanephora cucurbitarum.

CuTTER: THE GENUS CUNNINGHAMELLA oon

ACKNOWLEDGMENTS

The author wishes to extend special thanks to Dr. J. Walton Groves of
the Canadian Department of Agriculture for the contribution of a large
series of cultures which made possible this study. Acknowledgment is also
made to Dr. D. H. Linder and Dr. W. L. White of the Farlow Herbarium,
Harvard University, and Dr. S. A. Waksman who have contributed cultures
and made many helpful suggestions. The curators of the American Type
Culture Collection have most generously cooperated by making available
all cultures of Cunninghamella in their collection. Dr. Kenneth Raper has
extended valuable aid by donating many cultures from the Blakeslee Col-
lection. Finally the author wishes to acknowledge the aid of Miss Shirley
Weinstein, who assisted in the cultural studies of this material.

DEPARTMENT OF BOTANY
CoRNELL UNIVERSITY
Irmaca, N. Y.
BIBLIOGRAPHY

1. Alcorn, G. D., & C. C. Yeager. A monograph of the genus Cunninghamella, with
additional descriptions of several common species. Mycologia 30: 653-658. 1938.

2. Bainier, G., & A. Sartory. Etudes morphologique et biologique du Muratella
elegans n. sp. Bull. Soc. Mycol. France 29: 129-136. 1913.

3. Berlese, A. N. Fungi Moricolae. Padova, 1889.
4. Bisby, G. R. Who is the author? Trans. British Mycol. Soc. 25: 434-435. 1942.

5. Blakeslee, A. F. Sexual reproduction in the Mucorineae. Proc. Amer. Acad. Arts
and Sci. 40: 205-319. 1904.

6. ——————. Two conidia-bearing fungi. Cunninghamella and Thamnocephaiis,
n. gen. Bot. Gaz. 40: 161-170. 1905. :

7. Blakeslee, A. F.. J. L. Cartledge & D. S. Welch. Sexual dimioxphist in Cun-
ninghamella. Bot. Gaz. 72: 185-219. 1921.

8. Briquet, J. International Rules of Botanical Nomenclature. Jena, 1935.

9. Burger, O. F. Sexuality in Cunninghamella. Bot. Gaz. 68: 134-146. 1919.

10. Christenberry, G. A. A taxonomic study of the Mucorales in the southeastern
United States. Journ. Elisha Mitchell Sci. Soc. 56: 333-366. 1940,

11. Fitzpatrick, H. M. The Lower Fungi; Phycomycetes. New York, 1930.
12. LeClerg, E. L. Cultural studies of some soil fungi. Mycologia 22: 186-210. 1930.

13. Leeke, N. Review of, Stadel, O. Uber einen Neuen Pilz Cunninghamella Berthol-
letiae. Dissert. Kiel, 1911. Mycol. Centralbl. 1: 218-219, 1912.

14. Lendner, A. Sur quelques Mucorinées. Bull. Herb. Boissier. ser. II, 7: 250-251.
1907.

15. ——————. Les Mucorinées de la Suisse. Berne, 1908.

16. ——————. Une nouvelle espéce de Mucorinée, le Cunninghamella Blakesleeana

_ Lendner. Bull. Soc. Bot. Geneva 19: 234-238. 1927.

17. Ling-Young, M. Etude biologique des phénoménes de la sexualité chez les
Mucorinées. Rev. Gen. Bot. 42: 722-752. 1930.

18. Matruchot, L. Une Mucorinée purement conidienne, Cunninghamella africana.
Ann. Mycol. 1: 45-60. 1903.

19. Morrow, M. B. The soil flora of a pine forest. Mycologia 24: 398-402. 1932.

20. Naumov, N. A. Clés des Mucorinées. Paris, 1939.

ya
ogee

FaRLOwWIA, VOL. 2, 1946

. Ou, S. K. Phycomycetes of China I. Sinensia 2: 427-449. 1941.
. Paine, F. S. Studies of the fungous flora of virgin soils. Mycologia 19: 248-267.

1927.

. PiSpek, P. A. Edafske mukorinje Jugoslavije. Acta Bot. Inst. Bot. Univ. Zagreb.

4: 77-112. 1929.

Povah, A. H. W. Helicostylum and Cunninghamella: two genera of the Muco-
rales new to the state. Ann. Rept. Mich. Acad. Sci. 17: 152-155. 1915.

. Ricker, P. L. Second supplement to new genera of fungi published since the year

1900, with citation and original descriptions. Journ. Mycol. 12: 60-67. 1906.

. Ridgway, R. Color Standards and Nomenclature. Wash., D. C. 1912.
. Rivolta, S. Dei Parassiti Vegetali. Torino, 1873.

Saccardo, P. A. Fungi veneti novi vel critici v. mycologiae venetae addendi.
Series XII. Michelia 2: 241-301. 1881.

. Fungi Italici Autographice Delineati. fig. 805. Patavii, 1877-1886.

. Syll. Fung. 17: 508. 1905.

1. Saito, K. Actinocephalum japonicum nov. gen. et nov. spec. Bot. Mag. Tokyo 19:

36-38. 1905.

. Saito, K. & H. Naganishi. Eine Neue Art von Cunninghamella. Bot. Mag.

Tokyo 29: 285-286. 1915.

. Stadel, O. Uber einen neuen Pilz, Cunninghamella Bertholletiae. Dissert. Kiel.

1-35. 1911.

. Swift, M. E. Contributions to a mycological flora of local soils. Mycologia 21:

204-221. 1929.

. Thaxter, R. On certain new or peculiar North American Hyphomycetes. I.

Oedocephalum, Rhopalomyces and Sigmoideomyces n. g. Bot. Gaz. 16: 14-26. 1891.
. A New England Choanephora. Rhodora 5: 97-102. 1903.

. Torrey, G. S. Les Conidies de Cunninghamella echinulata Thaxter. Bull. Soc.

Mycol. France 37: 93-99. 1921.

. Zycha, H. Kryptogamenflora der Mark Brandenburg, Pilze II, Mucorineae:

1-264. Leipzig, 1935.

ll

340 Fartowia, VoL. 2, 1946

EXPLANATION OF PLATE I.

All figures drawn with the aid of a camera lucida at a magnification of x1300 and
reduced in printing to an approximate magnification of x325.

A-F. Cunninghamella echinulata. Types of conidiophores and conidia produced on
M.D.A. agar.

G. Cunninghamella echinulata. Conidiophore and conidia produced on liquid me-
dium, showing attenuated lateral branches and long echinulations on conidia.

H-I. Cunninghamella echinulata. Vesicles and chlamydospores formed on submerged
mycelium.

J—M. Cunninghamella Bainieri. Types of conidiophores and conidia.

N. Cunninghamella Bainieri. Conidiophore showing arrangement of giant conidia
on specialized lateral branch.

Cunninghamella Bainierit. Types of conidia.

CuTTER: THE GeNuS CUNNINGHAMELLA 341

342 FarLowlia, Vout. 2, 1946

EXPLANATION OF PLATE II.
All figures drawn with the aid of a camera lucida at a magnification of x1300 and
reduced in printing to an approximate magnification of x325.
A-D. Cunninghamella Blakesleeana. Types of conidiophores and conidia.
E-G. Cunninghamella Bertholletiae. Types of conidiophores and conidia.
H. Cunninghamella elegans. Conidiophore and conidia.

343

CuTTER: THE GENUS CUNNINGHAMELLA

9

A On O |
08

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2 Qo

PurateE II

2(3) :345-383 FARLOWIA JANUARY, 1946

A COMPARATIVE STUDY OF.TORULOPSIS PULCHERRIMA
AND TAPHRINA DEFORMANS IN CULTURE '*

CATHERINE ROBERTS
INTRODUCTION

The concept of a morphologic or phylogenetic relationship existing be-
tween the yeasts and the fungi comprising the order Taphrinales (or
Exoascales) is not of recent origin. Reess (1870), De Bary (1884), and
Hansen (1904) all expressed such an opinion, based upon macroscopic and
microscopic resemblances in culture between the two groups of fungi. At
present, two widely used schemes of mycological classification, that of
Giumann and Dodge (1928) and that of Bessey (1935, 1942), differ radi-
cally in regard to the relative positions of the Taphrinales and the yeasts.
From a phylogenetic point of view, Gaumann and Dodge place the two
groups in the primitively simple Hemiascomycetes, but separate the yeasts
from the Taphrinales by relegating the former to the order Endomycetales.
On the other hand, Bessey regards the yeasts and the Taphrinales as being
widely separated, the latter representing an offshoot of the Pezizales (which
he assumes to be primitive), while the yeasts are regarded as ‘“‘the ultimate
degree of simplification in the Class Ascomyceteae” and are therefore placed
at the end of this group. In 1937 Guilliermond concluded that “. . . les
Exoascées que la plupart des mycologues modernes s’accordaient a séparer
radialement des levures et 4 rattacher aux Ascomycétes supérieurs .. .
nous apparaissent, au contraire, beaucoup plus proches des levures qu’on
ne le pensait et l’on en revint ainsi aux anciennes conceptions de Reess,
de Bary et Hansen.”

These concepts were based on similarities evident in culture, but it should
be emphasized that only one phase in the life cycle of Taphrina — the
saprophytic, budding stage — has been found to be readily capable of
‘development on artificial media. The parasitic, mycelial stage of this genus,
involving sexuality, has resisted all attempts at culture, exclusive of scat-
tered reports in the literature to be referred to later in more detail.

In brief, the life cycle of Taphrina deformans (Berk.) Tul., the causal
agent of peach leaf curl and the species investigated in this study, is as
follows: Asci are formed in the spring and summer on the surfaces of in-
fected leaves, and each normally produces eight haploid ascospores which
are discharged in a group. The ascospores fall to various parts of the host
plant and initiate a yeast-like budding or sprout conidial stage, in which

1 Portion of a thesis submitted to the Graduate Division, University of California in
partial satisfaction for the degree of Doctor of Philosophy, October, 1943. The valuable
advice and criticisms offered during the course of the investigation and in the prepara-

tion of the manuscript by Prof. J. T. Barrett, Dr. H. N. Hansen, Dr. E. M. Mrak, and
Prof. M. W. Gardner are gratefully acknowledged.

345

346 FartowiA, VoL. 2, 1946

form the organism overwinters, In the following spring when the leaf buds
are expanding, infection occurs through penetration by an infection hypha
produced by germination of the sprout conidium. It is believed (Eftimiu
1927, Fitzpatrick 1934, Martin 1940) that the occurrence of nuclear divi-
sion without cross-wall formation in this germinating cell is responsible for
the origin of the binucleate or dicaryon condition which is evident in the
cells of the intercellular mycelium subsequently developing throughout the
leaf tissue. From this mycelium is formed a subcuticular layer of rather
thick-walled binucleate cells, the ascogenous cells. Karyogamy occurs in
these, followed by the extrusion of the cell wall to form the ascus, which in
its development ruptures the cuticle. The diploid fusion nucleus divides
mitotically, one of the daughter nuclei passing into the ascus, which is then
cut off from the ascogenous cell by a cross wall. Meiotic and mitotic
nuclear divisions then occur, resulting in the formation of eight haploid
ascospores, and the cycle is completed. In investigations of other species
of Taphrina by Wieben (1927) the origin of the dicaryon was reported to
occur through the copulation of physiologically differentiated ascospores or
their derivatives, but in T. deformans only rare instances of copulation have
been noted (Mix 1929), and it is generally believed that this species is
homothallic.

As a group, the yeasts are readily culturable and may be maintained for
long periods of time on various types of artificial media, both natural and
synthetic. The majority of the yeasts are saprophytic organisms, but para-
sitic forms exist, both on plants and animals. Yeasts fall into two main
groups — the sporogenous or sporulating and the asporogenous or non-
sporulating forms. The sporulating yeasts pass through complete life cycles
in culture, and although the types of sexuality involved are greatly diversi-
fied, the formation of ascospores within asci at some stage in development
is a constant occurrence in all those organisms investigated. Those yeasts
which have never been known to produce spores in culture are known as
the imperfect yeasts and have been regarded by some as perfect yeasts
which have lost their sporulating ability. They are placed among the
Fungi Imperfecti, and, as would be expected, no attempt to relate them
one to another in a natural system is made, but they are merely classified
for convenience in identification. Their development in culture lacks any
type of sexuality or alternation of haploid and diploid generations and con-
sists merely of a predominantly budding type of vegetative reproduction,
which often exhibits a pseudomycelial tendency. The genus Torulopsis is
classified with the imperfect yeasts in the family Torulopsidaceae and the
sub-family Torulopsidoideae, the latter being characterized by the absence
of pseudomycelium, or, if present, by a very primitive type (Lodder 1934).

An important contribution to our knowledge of the imperfect yeasts was
made by Lodder in 1934. As a result of a detailed study of the physiology
and morphology of these organisms (exclusive of the pseudomycelial-
forming Mycotoruloideae), she erected a scheme of generic and specific
determination which has since been recognized as the standard scheme of

ROBERTS: TORULOPSIS AND TAPHRINA 347

classification for these non-sporulating forms. From the standpoint of the
problem of the relationship between the yeasts and the Taphrinales, this
work is significant because of the fact that if this key is used, the genus
Taphrina, as represented by its saprophytic budding stage in culture, will
fall into the genus Torulopsis. The importance of this can hardly be over-
emphasized, for it is possible, therefore, that isolates of Taphrina may be
identified as belonging to the genus Torulopsis. The differences in the
natural habitats of these two groups of fungi are not so great as to preclude
their being isolated from similar or identical material, for, although the
budding stages of the parasitic species of Taphrina are quite closely asso-
ciated with their hosts, the species of Torulopsis are quite widespread and
have been isolated from a great variety of living and non-living material.
In this connection, the investigations of Koch (1934) on the overwintering
of parasitic and saprophytic fungi on fruit trees stress the actual difficulty
involved in the identification of isolates of Taphrina deformans. Koch
states, “Among numerous other fungi appearing in isolations from buds
and bark of Prunus persica, an organism resembling Taphrina deformans
appeared on five occasions . . . as this organism is somewhat difficult to
distinguish from certain ‘budding’ saprophytic fungi, many of which appear
consistently in isolations from the bark of peach trees, it was necessary to
prove the identity of the organism by inoculation.”

Studies of the general cultural behaviour of species of Taphrina have
already been referred to, but some recent investigations of Torulopsis spp.
in culture are worthy of mention at this time. Punkari and Henrici (1933,
1935), through the employment of single cell technique, studied the cultural
variation occurring in several generations of Torulopsis pulcherrima (Lind-
ner). Saccardo and noted striking dissociation into pigmented and non-
pigmented colonial areas, The white variants, although showing a tendency
to “breed true,’ were never completely stable, and the authors concluded
that these phenomena could be explained by the occurrence of ‘“‘mutation-
like changes” which, in a few cases, appeared to be reversible. Three years
later, Porchet (1938), although apparently unaware of Punkari and Hen-
rici’s contributions, also reported the occurrence of dissociation in T. pul-
cherrima. Her results, based upon mass transfer technique, indicated that
the formation of the pigmented and non-pigmented types may be a re-
versible phenomenon. In 1936 there appeared a paper by Todd and Herr-
mann dealing with the life cycle of a yeast which is known as the causal
agent of yeast meningitis. This organism had been known as Torula his-
tolytica Stoddard and Cutler until Lodder (1934), considering the generic
name Torula Turpin to be invalid, designated it as Torulopsis neoformans
(Sanfelice) Lodder. Todd and Herrmann, however, on the basis of priority,
adopted the name Cryptococcus hominis Vuillemin and described the oc-
currence of heterogamous copulation and the formation of single-spored,
thick-walled asci in this organism. This. constituted the first report of
sporulation in the genus Torulopsis. Because of the transformation of the
zygote into the ascus, and the formation of only one ascospore per ascus,

348 FartowlA, Vov. 2, 1946

Todd and Herrmann designated the perfect stage of C. hominis as Debaryo-
myces hominis (Vuillemin) Todd and Herrmann. A complete life cycle was
then formulated for this species, but it should be pointed out that it was
based on isolated observations of various types of morphological structures,
the actual sequence of events in the developmental life cycle of individual
cells not having been observed. Confirmation of Todd and Herrmann’s
observations on the formation of single spores in this yeast was reported
by Giordano (1938-39) and Henrici (1941), the latter also citing a
paper by Redaelli, Ciferri, and Giordano (1937) in which sporulation was
described.

In 1938 Windisch described for the first time heterogamous copulation
and subsequent ascospore formation in Torudopsis pulcherrima, apparently
occurring in response to the effect of a Penicillium contaminant in the cul-
tures. In this organism, four spores were produced per ascus.

Two years later Windisch (1940) in a lengthy paper supplemented by
drawings and photomicrographs described in detail the life cycle of T. pul-
cherrima (in pure culture) with particular emphasis laid upon the method
of sporulation. In addition, observations were made upon another imperfect
yeast, Candida tropicalis (Castellani) Berkhout 2 and a discussion of the
relationship of these fungi to each other and to the Taphrinales was in-
cluded. Through actual observation of the developmental sequence of
individual cells in hanging drop cultures, Windisch was able to formulate
a life cycle for T. pulcherrima which, in brief, is as follows: Large thick-
walled cells containing large oil droplets (typical “pulcherrima cells’’)
which were interpreted as ascus mother cells (ascogenous cells) occurred in
old cultures; these gave rise to bud-like protrusions into which passed a
portion of the oil droplet and in which were subsequently formed four
ascospores; rupture of the wall of this protrusion (or ascus) liberated the
spores, the empty ascus wall remaining attached to the ascus mother cell;
copulation occurred between cells derived from germination of the asco-
spores and resulted in the subsequent formation of large thick-walled
“pulcherrima cells.” Nuclear staining revealed a binucleate condition in
all vegetative cells as well as in the ascus mother cells. These phenomena,
which were also similar to those of Candida tropicalis, were interpreted by
Windisch as being indicative of a definite relationship with the Taphrinales.
He therefore proposed that T. pudcherrima be incorporated in the genus
Candida as C. pulcherrima and “alle candidoiden Pilze mit gleichartigen
Entwicklungsgangen zusammenzufassen in der neuen Familie der Candi-
daceae, die in der Ordnung der Exoascales einzureihen wire.”

‘It should be noted that these proposals of Windisch were based upon a
comparison of the cultural behaviour of T. pulcherrima with previous de-
scriptions in the literature of the culturable as well as the parasitic stages
of Taphrina, especially those of Wieben on the heterothallic species, 7.

* The genus Candida is placed in the family Torulopsidaceae but because of abundant

production of pseudomycelium is included in the sub-family Mycotoruloideae, rather
than with Torulopsis in the Torulopsidoideae (Lodder 1934).

ROBERTS: TORULOPSIS AND TAPHRINA - 349

epiphylla Sadeb. and T. klebahni Wieben. Windisch made no comparative
studies on the behaviour of Taphrina spp. in culture.

In the present investigation, a study of the comparative cultural be-
haviour of one isolate of T. pulcherrima and several isolates of T. deformans
was made in order to obtain additional information which could be used
either in substantiation or rejection of the proposals of Windisch in regard
to the relationship between 7. pulcherrima and the genus Taphrina. T.
deformans was chosen for this purpose because its economic importance
has attracted the attention of a large number of investigators, whose cumu-
lative efforts have resulted in this species having become the best known
representative of the Taphrinales. It is, of course, obvious that a compara-
tive study of any two species based upon observations of only one or a few
isolates from each will not give a complete picture of the range of existing
differences or resemblances. The present results, therefore, are considered
of necessity only preliminary, but it is hoped that they may be used as a
basis for future comparative studies of these or similar organisms.

MATERIALS AND METHODS

SOURCES OF CULTURES

The strain of Torulopsis pulcherrima used in this investigation was kindly
supplied by Dr. E. M. Mrak of the Division of Fruit Products, University
of California, and had been isolated by him from the surfaces of ripe Con-
cord grapes obtained in Walnut Creek, California. This particular isolate
had been maintained in culture for several years prior to the present
investigation.

Isolates of Taphrina deformans were secured from infected leaves of
peach trees growing on the Berkeley campus and in Piedmont in April and
May, 1941, and in the Kirkman Nurseries in Bethany, California, in April,
1942. The peach trees from which isolations were made consisted of
~ ornamental types of unknown variety as well as the commercial varieties,
Yunnan and Shalil.

The usual method of obtaining Taphrina spp. in culture was employed
(Martin 1925, Mix 1935). This consisted of attaching, by means of melted
wax, fragments of infected leaves containing actively discharging asci to
the underside of a Petri dish cover which was then placed over the lower
half containing solidified agar. Ascospores in groups of eight were subse-
quently discharged upon the agar surface, and when transferred to agar
slants, each individual group budded readily, producing a colony which
represented a single ascus culture. In obtaining single spore cultures of
T. deformans, the eight ascospores derived from a single ascus were sepa-
rated one from another by means of a micromanipulator which is described
below.

Stock MEDIA

The cultures of both T. deformans and T. pulcherrima were maintained,
for the most part, at room temperature (20°-25° C.) on potato dextrose

350 Fartow1a, VoL. 2, 1946

agar with a pH range from 5.6 to 6.0. In addition, the vegetable agar of
Mrak, Phaff, and Douglas (1942) which consists of equal parts of carrots,
beets, cucumbers, and potatoes with 2% agar at a pH of ca. 5.7 was em-
ployed. Limited use was made of a medium composed of a juice extraction
from leaves of susceptible peach varieties, with the addition of 2 % dextrose
and 2% agar. Liquid media were not used for maintaining stock cultures.

SINGLE CELL TECHNIQUE

All single cell isolations of both organisms were carried out by means of
a locally made micromanipulator attached to the stage of a Leitz compound
binocular microscope. A single glass needle, the tip of which varied from
about 10 to 17 » in diameter, was clamped in the holder of the micro-
manipulator in such a position that its tip was between the low power
objective and the object in focus, at a magnification of 100x. Three dif-
ferent motions of the needle in two planes were made possible by three
screw adjustments. The culture to be used was streaked across a plate of
potato dextrose agar and incubated at room temperature. When growth
was evident macroscopically, a block of agar, 10 to 15 mm, square and
containing on one edge a portion of the margin of the streak, was cut out
and transferred to a sterile slide. This was placed under the low power
objective of the miscroscope, and the margin of the colony and the tip of
the glass needle were brought into focus. By moving the needle, a single
cell at the margin was then separated from the colony by being drawn or
pushed along the surface of the agar. Plate I, Fig. 1 shows a single cell
separated in such a manner from the margin of a colony of T. deformans.
After the cell had been moved to a distance of 2 to 5 mm. from the colony,
its position on the agar block was designated by making an indentation in
the agar around the cell by means of the tip of the needle. Four or five
isolations were made on each agar block, which was then placed in a sterile
moist chamber, and after one to seven days, depending upon the growth
rate of the organism, a small colony derived from the single cell was usually
evident under the dissecting microscope (Plate I, Fig. 2). Transfers were
accomplished by touching the colony with the tip of a sterile needle and
transferring the adhering cells to an agar slant. It was found possible, after
some practice, to make twenty-five to thirty single cell isolations with the
micromanipulator in an hour. The dilution plate method of isolating single
spores as described by Hansen and Smith (1932), although possessing the
advantage of greater rapidity, was not satisfactory for the fungi studied
because of the comparatively small size of the cells which did not permit
them to be easily distinguished under the dissecting microscope from nu-
merous particles apparent in the agar. A definite advantage of the micro-
manipulator method over the dilution plate method is that the use of the
compound microscope with its higher magnification made possible the isola-
tion of any individual cell desired, as well as its positive identification as a
thin-walled cell, a thick-walled cell, a rod-shaped cell, etc. A similar tech-
nique was used to obtain monosporous cultures of T. deformans from dis-

ROBERTS: TORULOPSIS AND TAPHRINA 351

charged groups of eight ascospores, only that here the ascospores were
separated one from another by drawing them out with the micromanipulator
needle in radial directions from the center of the group.

STAINING METHODS

Most of the observations on cellular morphology were made from un-
stained living material in water mounts. In a few instances, a stain of
cotton blue in lactophenol was used (Rawlins 1933), a .025% solution
being found the most satisfactory.

For demonstrating the nuclei in the cells of T. pulcherrima, Laustsen’s
modification of the Feulgen technique (Winge 1935) gave fairly satisfactory
results, as had previously been noted by Windisch (1940) for the same
organism. The method consists of smearing the organism on a clean slide
by means of a cover slip, immersing in chloroform for thirty seconds, im-
mersing in cold water for two minutes, hydrolyzing in IN-HCI at 60° C.
for five to fifteen minutes, washing in water, and staining in alcoholic
fuchsin for three hours. This method was not found to be satisfactory as a
nuclear stain for the cells of T. deformans. The iron-alum-haematoxylin
schedule of Rawlins (1933) gave only fair results, while the use of Mayer’s
haem-alum (Chamberlain 1932) was somewhat more satisfactory. This
later method consisted of smearing the organism on a clean slide which
had been coated with a thin film of albumin, fixing in formalin-acetic-alcohol
No. 2 (Rawlins 1933), washing in water, staining in Mayer’s haem-alum
overnight, and counterstaining with either erythrosin or eosin for one-half
minute.

PHYSIOLOGICAL TESTS

The comparatively few physiological tests undertaken were performed
in accordance with the methods described by Lodder (1934) in her mono-
graph on the imperfect yeasts for the determination of fermentative ability,
the determination of sugar assimilation by non-fermenting strains, and the
determination of nitrogen utilization.

PHOTOGRAPHY

The majority of photomicrographs were taken with a Zeiss Contax camera
with a Miflex attachment. In printing, the microfilms were enlarged 2x.

EXPERIMENTAL RESULTS AND THEIR INTERPRETATION

MACROSCOPIC VARIATION IN CULTURE

The general appearance of the growth of T. deformans on potato dextrose
agar showed a striking resemblance to that of T. pulcherrima on the same
medium. Colonies of both organisms may be characterized by the phrase
“veast-like growth,” which, although not specific, is nevertheless frequently
employed as a descriptive term for the budding stages of various unrelated
fungi.

S32 Fartowla, VoL. 2, 1946

After several weeks at room temperature on potato dextrose agar both
T. deformans and T. pulcherrima produced colonies which were smooth,
dull (sometimes slightly glistening), dry, slightly convex, and with predom-
inantly entire borders. The most important distinguishing character be-
tween the appearance of the two fungi in culture was in the pigment forma-
tion. Although both organisms normally develop reddish or pinkish pigment
of a non-carotinoid nature (Lodder 1934), that of T. deformans is non-
diffusible, while that of T. pulcherrima readily diffuses into the surrounding
medium. In addition to the diffusibility of the pigment, the two organisms
were also distinguished by the actual color of the pigments produced. The
pinkish color of the normal colonies of T. deformans was identified as pale
grayish vinaceous by using Ridgway’s color standards (1912), but sectors
of greater or lesser intensity of color frequently occurred. The normal
colonies of 7. pulcherrima produced varying shades of red, characterized
according to Ridgway as vinaceous russet, avellaneous, sorghum brown,
and army brown. In the unpublished description of this strain made at the
time of isolation, E. M. Mrak had characterized the colony color as “pale
pinkish buff” on wort agar, while Mix (1924) had described the appearance
of colonies of T. deformans on potato dextrose agar as “moist” and “shin-
ing” and their color as “pale vinaceous pink.”

The occurrence of cultural variation in the form of sectors was very fre-
quent in mass and single cell colonies of these organisms, and in addition to
the colors described above, T. deformans frequently produced mustard or
Naples yellow sectors as well as white or non-pigmented sectors which
darkened with age. 7. pulcherrima likewise produced abundant non-
pigmented sectors which darkened with age to colors characterized as
Tilleul-buff and vinaceous buff. Thus, the usage of the term “non-pig-
mented” as descriptive of white colonies or sectors for comparison with red
pigmented colonies is not a completely accurate one, for the white areas
may darken with age to shades of pale brown or buff. Likewise, the term
“red” is used with reservations, for no distinction is made among the
previously described varying shades of red developing in old cultures with
diffusible pigment.

Although smoothness appeared to be a normal character of the surface of
colonies, both young and old cultures of 7. deformans at times produced
roughened areas, some of which were in the form of sectors. These areas,
usually of the papillate type, were found to develop sporadically and seem-
ingly bore no relation to the colony color. Rough surfaces, both papillate
and crateriform, were also found in cultures of T. pulcherrima but were
usually confined to non-pigmented cultures which had darkened with age.
The papillate type was much more common in this organism than was the
crateriform.

These preliminary observations demonstrated the close resemblance
between the cultural appearance of T. deformans and T. pulcherrima,
suggesting the possibility of confusion in identification, especially if Lod-
der’s recent generic key for the determination of the non-sporulating yeasts

ROBERTS: TORULOPSIS AND TAPHRINA 353

is used. Differentiation of the genera on the basis of a diffusible pigment is,
of course, possible, but because of the production in culture of non-pig-
mented variants of both organisms, such types may be isolated directly
from nature..

ANALYSIS OF SINGLE SPORE CULTURES OF T, DEFORMANS

With a view toward making a more detailed study of the cultural varia-
tion of T. deformans, single spore cultures were obtained in 1941 from
discharged groups of ascospores in the manner previously outlined. From
a total of 215 ascospores discharged, 179, or approximately 83%, were
recovered, each being used as the starting point of a monosporous culture.
Each culture was designated by three symbols: a Roman numeral indicat-
ing the location of the host plant, a letter indicating the individual ascus, |
and an Arabic numeral indicating an individual ascospore discharged from
that ascus. As an example, X—A-—8 represented a culture derived from one
of eight ascospores which had been discharged from a single ascus pro-
duced on an infected leaf of a peach tree growing on the Berkeley campus.
Thirty-three discharging asci were used in these isolations, and of that num-
ber twenty-five discharged a complete set of eight spores, the other eight
asci discharging six or seven spores. Eight complete sets of monosporous
cultures out of the twenty-five were actually established in culture, while
of the remaining seventeen, two to seven ascospores from each ascus were
recovered.

The 179 established cultures, kept at room temperature on potato dex-
trose agar slants, were observed closely after several months’ time in order
to determine the extent of variation among the monosporous isolates from
individual asci. With reference to the findings of Wieben (1927) on the
segregation of sexually differentiated ascospores in the asci of the hetero-
thallic T. epiphylla and T, klebahni, both Mix (1935) and Fitzpatrick
(1934) have shown that in T. deformans the eight ascospores from one
ascus are sexually undifferentiated, although instances of “occasional” and
- “incomplete” copulation between cells from different monosporous cultures
were observed by Mix. This suggests that the fungus may possibly be
heterothallic, and it is conceivable, therefore, that segregation of other
factors could occur in the division of the diploid nucleus which might be
evident macroscopically as a definite ratio of cultural types among the
eight monosporous cultures derived from one ascus. No such ratio of
variants appeared, however, in any of the series of monosporous cultures
investigated, nor was there any evidence for the occurrence of definite ratios
of morphological or physiological variants. The fact appears to be well
established, therefore, that neither segregation of sex nor of other factors
occurs at the initiation of the haploid phase in the life cycle of T. deformans.

It is not to be inferred that variation was absent in the 179 monosporous
cultures, for it did occur, and quite abundantly, but not in any definite
ratio among the eight cultures derived from one ascus. The cultural varia-
tion observed was in the form of variously pigmented sectors arising

354 FarLOwIA, VoL. 2, 1946

sporadically, and often more than one type of sector would appear in the
same colony (Plate I, Figs. 3, 4).

These cultures were maintained as stock cultures on potato dextrose agar
for a period of two years, during which time they were transferred every

TABLE I
: me NORMAL
SECTORS
COLOR
CULTURE DATE OBSERVED ee :
a b c d
— XIL-P4 1942 x x
1943 x x
XIII-P2 1942 x
1943 x
XIII-O2 1942 x PAS
1943 bs
XITI-H1 1942 x x
1943 x = x
XTII-S4 1942 x x
1943 x x
XIII-l1 1942 x x x
1943 x =
XITII-T1 1942 x x x
1943 x
XIITI-P8 1942 x x
1943 x
XII-01 1942 x x
1943 x x
XIII-K3 1942 be x x
1943 x
XITI-K1 1942 x 24
1943 x x x
XITII-G4 1942 x
1943 x
XIII-E3 1942 x =
1943 = oA
VI-B1 1942 x =
1943 x x

Variations in pigmentation apparent in single spore cultures of 7. deformans in 1942
and in their sub-cultures in 1943. (a, pale grayish vinaceous; b, modifications in inten-
sity of a; c, white; d, mustard or Naples yellow.)

three or four months by mass inoculations from the colony, irrespective of
the sectoring. In 1942 recorded observations on the sectoring apparent in
each culture were made. This was repeated in 1943 on sub-cultures of
approximately the same age, and Table | is a tabulation of fourteen repre-
sentative cultures with comparative observations for the two years. It will
be noted that in three cases, the 1943 culture had the same macroscopic
appearance as did the 1942 ancestral colony, while in eleven cases, the

ROBERTS: TORULOPSIS AND TAPHRINA 355

1943 culture showed more, or less, or a different type of, variation than did
the ancestral colony. Because of the technique of transfer, these results are
of no significance in regard to the constancy of the variant types, but they
are regarded as evidence of the abundant production of varied types of
sectors in T. deformans maintained in culture.

ANALYSIS OF SINGLE CELL CULTURES OF T., PULCHERRIMA

Studies on the transmission * of the cultural variations observed in this
organism were undertaken by means of the single cell technique already
described. The original stock culture, which had been maintained on
vegetable agar at ca. 13° C. for several years, exhibited striking dissociation
into red and white areas after several weeks at room temperature on potato
dextrose agar. From both the pigmented and the non-pigmented areas of
this colony, mass transfers were made. The resulting colony from the white
sector developed no pigment, while that from the pigmented area again
produced a red colony with white sectors (Plate I, Figs. 5, 6). In all sub-
sequent work on variation, which involved nearly 350 individual isolations
during a period of 18 months, single cell technique was employed, and the
cultures were maintained on potato dextrose agar at room temperature.
The extent of this phase of the investigation makes it impossible to indi-
cate clearly all results on one chart, but Figure 1 represents diagram-
matically certain portions of the work. In these charts, each numeral rep-
resents an individual colony, and each group of numerals enclosed in a
bracket indicates all or part of a series of single cell colonies derived from
that portion of the parent colony at which the arrow originates. The actual
appearance of each colony is indicated by the letters W or R, the former
signifying a pure white or non-pigmented colony, and the latter signifying
a red or pigmented colony. When W and R are placed together under one
numeral, the colony so indicated was pigmented with one or more white
sectors.

Figures 1A and 1B are examples of the most characteristic and
frequent type of transmission of cultural variants found in T. pulcherrima.
It will be noted that in Figure 1A, single cells derived from an entirely
white colony give rise to pure white colonies. It will be noted in Figure
1B that cells derived from white sectors also give rise to entirely white
colonies; in addition, Figure 1B shows that cells taken from the pig-
mented portions of colonies with white sectors will give rise to pigmented
sub-colonies which again produce white sectors. It should be emphasized
that these colonial phenomena — the stability of the white variants and
the instability of the pigmented areas —— were found constantly, without
deviation, in 302 distinct single cell isolations, or in nearly 90% of the total
number isolated. (Plate I, Figs. 7, 8.) Consequently, the formation of pure

°“Transmission” is used here and elsewhere in this paper in the sense of the reap-
pearance of a cultural type in single cell sub-cultures. It is used in order to obviate any
confusion which might arise over the use of the word “inheritance” with its connotation
of sexuality.

356 FartowlA, VoL. 2, 1946

white colonies or of red colonies with white sectors usually could be pre-
dicted merely upon the basis of the portion of the parent colony from which
the single cell originated. The abundant production of totally white colonies
developing on the same medium which supported the growth of pigmented

on Gee Le aa ae 2 ge ES
W W | W W-. &@ W W W
Cy a ao es ee eS Oe ae ee as a ee ee kas 8
W Ww W W W W W W W W W W W WwW W W
A.
‘S ~ Je 20!
WR WR WR
‘se 8 7 gl
W W W WR WR
B.
we
WR
Sone Geet eee er as Sa a eS Se ar aa ae
WR W WR WR WR WR WR WR WR WR WR WR WR WR
Cc.

Fig. 1. Occurrence and transmission of cultural variants in single cell colonies of
Torulopsis pulcherrima, (Explanation in text.)

colonies indicates that the chemical composition of this substratum ap-
peared to play no role in pigment production (Porchet 1938), although
Beijerinck (1918) and Castelli (1940) have advanced the idea that the
formation of pigment in 7. pulcherrima is dependent upon traces of iron
salts in the medium.

However, three distinct exceptions to this phenomenon of variation were
found to occur in about ten percent of the cases. In the first type, twenty-

ROBERTS: TORULOPSIS AND TAPHRINA 357

four single cells, each derived from the marginal area of pigmented portions
of parent colonies, gave rise to wholly pigmented sub-colonies which failed
to develop any visible white sectors. One of these pure red colonies was
subcultured, the five derived single cell colonies also being pigmented and
failing to show any variation. Further subculturing from one of these re-
sulted in five more completely red colonies. Thus, the red cultural type
was not found to be completely unstable, although the possibility exists
that these apparently constant red types may have produced white sectors
in portions of the colony other than the surface layers,and consequently
were not apparent to the eye.

The second exception to the normal type of variation was the production
of two small brownish sectors in the red portion of a sectored colony de-
rived from a single cell. Single cells isolated from these brown sectors gave
rise to colonies of the normal red color. It is concluded that the brownish
sectors were merely temporary variants, the formation of which, therefore,
presumably did not involve a mutational change (Snyder and Hansen 1939).

The final type of irregularity in cultural behaviour involved the appear-
ance of a red pigment in single cell colonies which were derived either from
the white sectors of pigmented colonies (Figure 1C) or from pure white
colonies. It should be noted, however, that a preliminary examination in-
dicated that this pigment was perhaps not identical with the normal type
in that its color was usually a deeper red and it appeared to lack diffusibility.
This type of variation, although of comparatively rare occurrence, is never-
theless of utmost significance in regard to the interpretation of the results
of this study of cultural variation in T. pulcherrima as will be noted in the
following discussion.

In attempting to explain the occurrence and transmission of cultural
variation observed in single cell colonies of T’. pulcherrima, emphasis should
be placed upon the distinctive nature of the dissociation, in that white to
white and red to red and white appeared to be the normal behaviour. This
is in confirmation of the work of Punkari and Henrici (1933, 1935) who
state, ‘‘. . . red is much less stable than white. It has been possible to
maintain pure reds through three generations in 2 lines by selective sub-
culturing. But white sectors appear so frequently in red colonies that we
believe it will be impossible, under the conditions of this experiment to
maintain a pure red line indefinitely.”’ The lack of sporulation in their cul-
tures was justifiably regarded by Punkari and Henrici as proof of absence
of sexual reproduction, and they therefore concluded that the observed
dissociation bore no relation to hybridization or to any other phase in a life
cycle involving sexuality. ,

A possible explanation for the observed variation may lie in the investiga-
tions of Hansen (1938) on the dual phenomenon, in which a dual fungus is
regarded as one which is composed of two distinct cultural homotypes,
M (mycelial) and C (conidial), which may not be recognized until their
separation from the heterotype MC is accomplished by single spore or single
cell technique. Hansen stated that ‘the dual phenomenon is due to hetero-

358 FartowlA, VoL. 2, 1946

caryosis, i.e., that individual cells and individual spores of dual fungi con-
tain two genetically distinct types of nuclei,” and he further stated in 1942
that “two factors or mechanisms operate against the possibility of the uni-
nucleate condition being continuous or permanent,” the two factors being
mitosis and anastomosis. Heterocaryosis is believed to be initiated by ‘“‘(1)
mutation within a plurinucleate entity, and (2) by fusion or anastomoses
between cells having genetically unlike nuclei.” Hansen and Snyder (1943)
in discussing the relationship between the dual phenomenon and sex in a
species of Hypomyces found that in many cultures an M homotype of
extreme stability arose de novo in C homotypes with great frequency and
regularity and that the M type never reverted to the C type. They con-
cluded, as a result of crossing behaviour, that “the change of C to M is a
true genetic mutation, that it is a unidirectional one, apparently common
to fungi in general. . . .,” and “that this mutation may be the cause of
the dual phenomenon in all fungi where this condition occurs.” Although
the regular appearance of white sectors in red colonies of 7. pulcherrima is
certainly indicative of a unidirectional mutation, the actual proof of this
assumption would necessarily lie in crossing experiments involving the in-
heritance of the factor for white sectors. Although this has not been demon-
strated in this organism, it is nevertheless obvious that the stable white
variant obtained in culture may be considered analogous to the mutating
C homotype of these authors. In view of the uncertainty in regard to the
nuclear condition in this species, the designation of the original mass isolate
in regard to M and C would be a matter for speculation.

However, the exception to the normal dissociation involving the produc-
tion of red from white does not support the interpretation of the variation
in this organism as being an expression of the dual phenomenon because of
its reversibility. Rather, it is indicative of Punkari and Henrici’s ‘‘reversion
to ancestral forms” (1933), in which they found that a pure white colony
derived from a white sector gave rise to a mixed red and white subcolony.
However, if this pigment arising in white colonies is found to be of a differ-
ent nature than the normal type, the concept of reversion would not apply,
and the normal dissociation could then be regarded with a degree of cer-
tainty as an expression of the dual phenomenon. Studies on this phase of
the problem are now in progress.

One further point in regard to the production of pigment, regardless of
its nature, in colonies derived from white sectors is of significance. Actual
dissection of a three-month-old sectored colony revealed that the sector did
not extend throughout the entire depth of the colony. The relatively small
size of the colony and its sector made this relation rather difficult to deter-
mine, but further evidence of its existence was obtained by making mass
transfers of cells from different portions of the dissected colony. It was
found that transfers of cells from the base of the colony directly underneath
the white sector gave rise to subsequent pigmented colonies, indicating that
the white sector did not actually extend to the bottom of the colony. It is
conceivable that if a transfer needle were plunged rather deeply into a sector

RoBERTS: TORULOPSIS AND TAPHRINA 359

of this type, the cells transferred could be a mixture of both the non-
pigmented and the pigmented types, and that single cell isolations could
therefore give rise to both types. Further dissection of other sectored
colonies of both T. pulcherrima and T. deformans revealed that the depth
of the sectors may range from superficial layers of cells to an entire wedge
of the colony, emphasizing the fact that accurate interpretation of results
may be dependent upon the actual depth to which the transfer needle is
inserted into the sectored portions of the colony. In this connection,
Christensen (1940) has stated, “Sectors are not necessarily pure for type
because mutants may grow either over or under the parent, and subcultures
from such sectors may consist of the mutant plus the parent line.”” Whether
or not these facts may be correlated with the apparent reversion noted from
white to red is unknown, since the dissections were made on sectored col-
onies from other series; these facts are offered, however, as a possible
explanation, although, of course, they would not apply to the production
of pigment in colonies derived from pure white cultures.

Thus, it appears that because of the possibility of the occurrence of
reversibility, the dual phenomenon may not be the correct explanation for
the cultural variation occurring in T. pulcherrima. Regardless of inter-
pretation, however, the observed facts still point to this variation as being
of a distinctive type in that it has been found to be predominantly uni-
directional, stable, and predictable, and, as will be discussed later, is asso-
ciated with significant differences in cellular morphology.

ANALYSIS OF SINGLE CELL CULTURES OF T. DEFORMANS

The previously noted variants occurring in the form of colonial sectors
in monosporous cultures of T. deformans were subjected to a single cell
analysis similar to that made for T. pulcherrima in order to obtain addi-
tional information in regard to their transmission and stability. Because of
the number of monosporous isolates on hand, a series of single cell isola-
tions, each involving one or more generations, was made from each of six
monosporous sectored cultures; whereas in the analysis of the cultural
variants of T. pulcherrima, a single sectored colony had been used as the
basis for all subsequent single cell work.

The single cell colonies obtained in this manner exhibited, as a group,
the same range of variation in pigmentation which had been observed in
their ancestral monosporous cultures, as well as a sporadic tendency to
produce roughened surface areas which appeared to bear no relation. to the
pigmentation, and which were characterized by numerous small papillae.

Transmission of two (white and yellow) out of the three variants ob-
served was noted, while the third type, which was characterized by various
modifications in intensity of the normal pink colony color, did not appear
to be transmissible. It should be noted, however, that this third type was
often evident only as a slight deviation from the normal color, pale grayish
vinaceous. Consequently, although quite evident in its original appearance
as a colonial sector, its later recognition in sub-cultures involved some

360 FarLowiA, VoL. 2, 1946

difficulty. In spite of this, it should be emphasized that no positive evidence
was obtained for the transmission of this sector type (designated as d),
because all single.cells derived from 6 sectors gave rise to colonies the color
of which appeared to be essentially similar to the normal type (a). It is

Uxrrt-x6 |

ad, a

5!
a
A.
Pal ee. a Be pS gs ies ee ae
a,b a a a a c 6 o.4: ce ¢ ¢,a
CT 2. ot E i a
a,d a,d a c c c
B.

Fig. 2. Occurrence and transmission of cultural variants in single cell colonies of
Taphrina deformans. (Explanation in text.)

therefore concluded that sectors of the b type were temporary variants
which consequently were not. produced by mutational changes.
Transmission of the other two variants is shown in Figure 2 which
diagrammatically represents the single cell analysis made of two mono-
sporous sectored colonies, XIII-E6 and XITI-L1. In these diagrams, as
in those for T. pulcherrima, each numeral represents an individual colony,

ROBERTS: TORULOPSIS AND TAPHRINA - 361

and each group of numerals enclosed in a bracket represents a series of
single cell colonies derived from that portion of the parent colony at which
the arrow originates. The actual appearance of the colonies is indicated by
the following symbols: a, the normal color, pale grayish vinaceous; 5, modi-
fications of a either by greater or lesser intensity of color; c, non-pigmented
or white; and d, mustard or Naples yellow. When these letters are com-
bined under one numeral, that colony consisted of the color indicated by
the first letter, with sectors of the variant type(s) so designated.

Figure 2A exhibits three points worthy of note: (1) the transmission
of the yellow variant d; (2) the incomplete stability of d, as seen by its
dissociation in one colony to an a variant; and (3) the sporadic production
of 6 variants in the normal a type.

Figure 2B exhibits similar phenomena: (1) the transmission of the
white variant c; (2) the incomplete stability of c, as seen by its dissociation
into a variants; and (3) the sporadic production of variants 6 and d in the
normal @ type.

These observations point to the main distinction apparent between the
cultural variation of T. deformans and T. pulcherrima — namely, the regu-
larity with which the latter produces a single variant as contrasted with the
sporadic production of two variants by the former organism. In addition,
the apparent reversion of the white variant te the parent type as observed
in T. pulcherrima was of comparatively rare occurrence when contrasted
with the abundant reversions to the normal type occurring in the variants
of T. deformans. The two organisms are similar with respect to the produc-
tion of temporary variations in the form of non-transmissible sector types
—namely, the D variant of T. deformans and the brown variant of T.
pulcherrima.

It is therefore concluded that, while the observed cultural variation in
T. pulcherrima may possibly be regarded as an expression of the dual
phenomenon, no such interpretation can be inferred from the evidence
obtained from T. deformans. As has been previously noted, the regular
occurrence of the white variant in the red cultural type of T. pulcherrima
is indicative of a true mutation, but actual proof from inheritance studies
is lacking. Likewise, crossing experiments are not possible with the homo-
thallic T. deformans, so that the inheritance behaviour of the variants are
unknown. In addition, the single cell analyses performed with T. deformans
were not so extensive or of such long duration as to preclude the possibility
that the yellow and the white transmissible variants may also be only
temporary. As Snyder and Hansen (1939) have pointed out, “They (tem-
porary variations) may result from some factor in the environment .. .
and eventually revert to the parent type with the return of original con- .
ditions. Reversion, however, may be slow and the variants give a semblance
of permanency.” However, if they are to be regarded as true mutants, and.
the fact that they arose from monosporous cultures derived from haploid
uninucleate cells is good evidence that they may well be (Christensen 1940),
it should be emphasized that their spontaneous and unpredictable occur-

362 Fartowla, VoL. 2, 1946

rence would seem to preclude the possibility of their being an expression of
the dual phenomenon.

Microscopic EXAMINATION OF CULTURES

The general morphology of the cells of both organisms in culture was
found to be essentially similar to the descriptions given by previous in-
vestigators, such as those of Mix (1935) on T. deformans and those of
Lodder (1934) on T. pulcherrima.

Young cultures of both fungi maintained on potato dextrose agar at room
temperature were composed of round to oval, thin-walled cells which re-
produced vegetatively by typical budding. Both organisms produced cells
of similar shape, although those of 7. pulcherrima showed a tendency to
become more globose than did those of T. deformans. The range in cell size
was found to be similar to that previously reported for T. deformans
(2.4-6.3 x 3.6-8.6 u, Mix 1935) and for T. pulcherrima (3:0-4.5 x 3.5-6.5
wu, Lodder 1934), although somewhat larger cells were occasionally found
in young cultures of the latter. The main distinction in cellular morphology
between young cultures of the two organisms appeared to be in the produc-
tion of oil (or fat) globules in the round cells of Torudopsis and their absence
in the cells of Taphrina. These oil globules, although showing quite a range
in size, were characteristically quite large, often occupying almost the entire
cell. Such round cells containing a single large oil droplet have been desig-
nated by Lodder as typical “‘pulcherrima cells,” and, according to her, were
noted in the original description of the species by Lindner (1887). Nu-
merous other investigators (Beijerinck, 1918; Grosbusch, 1915; Punkari
and Henrici, 1933) of this organism have also noted them. In the present
to be much more abundant in older cultures, as will be discussed below.

Old cultures (one to three months) of both fungi were, in general. quite
similar microscopically and showed few significantly distinguishing char-
acters. In addition to the normal type of vegetative cells (Plate I, Fig. 9),
the abundant production of large, thick-walled, globose cells was noted in
both, those of 7. pulcherrima appearing as typical “pulcherrima cells,”
while those of T. deformans, although of similar shape, were often larger
and typically contained more than one oil globule (Plate I, Figs. 10, 11).
The size of these cells varied considerably in both organisms, but all were
larger than the vegetative budding cells, and some reached a diameter of
12 to 15 ». As previously mentioned, the occurrence of these large thick-
walled cells in T. pulcherrima has been noted by many investigators, while
Pierce (1900), Fitzpatrick (1934), Mix (1924, 1935), and Martin (1925,
1940) have observed them in 7. deformans. Their significance will be
discussed later.

SPORULATION

Previous reference has been made to the work of Windisch (1938, 1940),
in which sporulation was reported for the first time in T. pulcherrima. The

ROBERTS: TORULOPSIS AND TAPHRINA 363

results of the following microscopic investigation of this organism are
offered in partial confirmation of Windisch’s observations.

As previously noted, large thick-walled cells containing a single oil globule
were observed to be extremely abundant in old cultures of T. pulcherrima
on both potato dextrose and vegetable agar. In addition, it was found that
many of these cells possessed what appeared to be the attached remnant of
a cell wall. In view of Windisch’s interpretation of similar structures
representing stages in the life cycle of this organism, it was decided to in-
vestigate the species more fully for the occurrence of sporulation in culture.

It will be recalled that Windisch had observed in ‘‘pulcherrima cells” the
formation of bud-like protrusions into which passed a portion of the divided
oil globule and in which four spores were subsequently formed. The rupture
of the protruded daughter cell liberated the spores, the remnant of this cell
remaining attached to the mother cell, often for long periods of time.
Copulation between cells derived from the budding of the ascospores oc-
curred and resulted in the eventual formation of the large, thick-walled
“pulcherrima cells.”

Although not all stages in this life cycle were found, the following struc-
tures, believed to represent the sporulation process, were noted in both
mass and single cell cultures (Plate I, Figs. 12-14; Plate IT, Figs. 1-5):

Thick-walled “pulcherrima cells” with

1) papillate protrusions

2) attached daughter cells, some of which contained oil globules

3) attached daughter cells which contained one to three spore-like
bodies

4) attached remnants of ruptured daughter cells

Although not clearly evident in the photomicrographs, the small bodies
occurring in the attached daughter cells were readily distinguished from oil
globules by the fact that they were less refractive. In addition, a small
dark area, interpreted as a small oil droplet, is apparent in these bodies.
It is concluded that these structures are actually spores, and that all of the
structures listed above represent phases of the sporulation process. In this
connection, it should be noted that the morphology of these structures was
essentially similar to that described and illustrated by Windisch.

_ Windisch’s observations on the budding of the ascogenous cells were also

confirmed. He states, ‘“Die Mutterzelle sprosst wieder aus; noch wahrend
der erste Ascus aufsitzt, kann sie durch Sprossung eine oder mehrere Toch-
terzellen hervorbringen, von denen sich nicht sicher entscheiden lasst, ob
sie nun auch Asci werden oder Sprosszellen bleiben sollen.’’ Such a phe-
nomenon is illustrated in Plate II, Fig. 6, which represents the development
in twenty-four hours of a “pulcherrima (ascogenous) cell” with attached,
empty daughter cell (ascus), both of which had been separated together
from the margin of a colony by the micromanipulator. It will be noted that
in this particular case two cells have been formed from the ascogenous cell
at the opposite side from the empty ascus. No further growth occurred.

364 Fartowia, VoL. 2, 1946

Figure 6 of Plate IT is also of significance in that it reveals a morpho-
logical distinction between normal buds and those buds which are believed
to represent asci. It will be seen from this figure that the two vegetative
cells are somewhat constricted at their bases, while the assumed ascus has
a comparatively large, unconstricted base. This distinction between typical
buds and asci was quite characteristic in the cultures examined and was
used as a basis for differentiation in the absence of spore-like bodies.

Unfortunately, the isolate of T. pulcherrima used in these studies had
been maintained as a stock culture for several years prior to the present
investigation and appeared to be losing its sporulating ability. As a result,
comparatively few spores were found, and it was not possible to study the
actual sequence of events in the sporulation process of individual cells, as
did Windisch. Also, Windisch had found quite consistently spores in groups
of four, while in the present study, three spores were the most found in
one cell. Spore discharge and germination were not observed.

A few indications of copulation of vegetative cells were noted. One non-
pigmented colony was found to consist of cells which appeared to be copu-
lating with the formation of conjugation tubes (Plate IJ, Fig. 7). In an-
other non-pigmented colony, two cells were seen which were in such a
position as to be indicative of the copulation process.

Windisch observed sporulation in T. pulcherrima and subsequently sug-
gested the relationship of this organism to members of the genus Taphrina
on the basis of the latter’s sporulation as it occurs on the host plant. In
view of this fact, it was decided to make an intensive study of the cellular
morphology of 7. deformans in culture in order to determine whether °
Windisch’s views on the similarity of the two organisms could be further
substantiated.

One- to three-month-old cultures of the fungus on potato dextrose and
vegetable agar were examined microscopically and revealed the presence of
large thick-walled cells containing one to many oil globules, with, in addi-
tion, attached structures similar, if not identical, to those noted above for
T. pulcherrima (Plate II, Figs. 8-10; Plate I, Fig. 11).

The similarity of these structures to those observed in old cultures of
T. pulcherrima is readily seen from the photomicrographs, and it is believed
that the bodies occurring in the attached daughter cells of Taphrina may
also be regarded as spores, and that likewise the attached structures may
represent phases of the sporulation process. As in 7. pulcherrima, it will be
seen that each of the spore-like bodies contains what appears to be one or
two small oil globules. The discharge of these bodies was not observed.

The occurrence of this apparent sporulation in T. deformans in culture
has two significant aspects — first, it lends support to Mix’s report of the
occurrence of the diplophase of 7. deformans in culture, and secondly, it
further substantiates Windisch’s conception of a relationship between
T. pulcherrima and the Taphrinales.

In 1924 Mix observed large, thick-walled cells in old cultures and desig-
nated them as “‘resting spores.” He noted that in germination they ‘‘may

RoBERTS: TORULOPSIS AND TAPHRINA 365

burst, allowing the escape of a thin-walled protoplast, a behaviour similar
to that of the ascogenous cells found in the leaf,” and, in agreement with
Pierce (1900), he suggested that “the resting spores developed in culture
may indicate an approach to the ascigerous stage.” In 1929, he reported
in an abstract that the resting cells are believed to be ascogenous cells and
that “structures believed to be imperfect asci have also been found.” In
1935 further emergences from the germinating resting spores were noted
and interpreted as asci, some of which were found with “2 and 3 fully
formed spores.” Other workers, such as Martin (1940) and Fitzpatrick
(1934) have noted similar large, thick-walled cells in culture, but have not
ascribed to them the character of ascogenous cells, for no structures re-
sembling asci were found. Although Mix’s assumed asci were comparable
in size and shape to the normal asci occurring on the host, it is believed that
the smaller attached daughter cells containing spore-like bodies noted here
may. also be regarded as imperfect asci with ascospores, and that conse-
quently the large, thick-walled mother cells may be interpreted as ascoge-
nous cells. It is interesting to note that the large thick-walled cells were
found in both mass and monosporous cultures of T. deformans, indicating,
as Martin (1940) has recently pointed out, that “their formation is evi-
dently not dependent upon conjugation.”

It will be recalled that Windisch interpreted the ‘“pulcherrima cells” of
T. pulcherrima as ascogenous cells and the bud-like protrusions developing
into daughter cells as asci in which ascospores were formed. The similarity
between these structures occurring in culture and the ascogenous cells, asci,
and ascospores of species of Taphrina was then pointed out, and resulted
in the conception of a close relationship between T. pulcherrima and the
Taphrinales. It is believed that the results of the present study provide
further supporting evidence for this view, because of the fact that quite
similar types of cultural sporulation in both T. pulcherrima and T. de-
_formans have been observed.

COMPARATIVE CELLULAR MoRPHOLOGY OF THE CULTURAL cyees or TL.
PULCHERRIMA

Both Porchet (1938) and Windisch (1940), although aware of the varia-
tion in pigmentation occurring in T. pulcherrima, worked with mass rather
than with single cell cultures and consequently failed to obtain a complete
separation of the two pure cultural types. As a result, their observations
on cellular morphology were undoubtedly made, in part, on mixtures of
cultural types and are therefore of little significance in a study of the com-
parative cellular. morphology of the pigmented colonies and their non-
pigmented variants. On the other hand, Punkari and Henrici (1933, 1935)
by means of single cell technique were able to separate the white variant
from the pigmented portion, and, as a result of microscopic examination,
concluded (1933) that “no morphologic differentiation could be established
between red and white variants.” The evidence obtained from the present
investigation of this phase of the problem indicates, to the contrary, that

366 ¥ FARLOWIA, VoL. 2, 1946

some differentiation on the basis of cellular morphology is apparent between
the two cultural types.

Microscopic investigations were made of all of the single cell colonies
used in the work on cultural variation, each colony or sector being exam-
ined after it had been growing on potato dextrose agar for one month or
longer. The results of this study showed that the red pigmented cultural
type produced abundant thick-walled cells, often containing oil globules
(Plate II, Fig. 11), while the white variants, both in the form of sectors
and entire colonies, often lacked this tendency and consisted, for the most
part, of smaller, thin-walled cells reproducing vegetatively by typical
budding # (Plate II, Fig. 12). This phenomenon was first noted in the
original mass sectored culture on potato dextrose agar. The results of
subsequent microscopic examinations of single cell colonies derived from
this ancestral culture are tabulated as follows:

Number of Cultures
Containing Large
Number of | Number of Cultures Thick-walled Cells

Cultures Containing Large with Attached

Examined Thick-walled Cells Daughter Cells
Red Cultural Type 44 40 31
White Cultural Type 141 10 ; 3
Total “185 50 34

It should be noted that some cellular structures occurring in both the
red and white cultural types were rather difficult to classify because of
transitional stages in size and shape, due, it is believed, to the age of the
culture and the portion of the colony from which the cells to be examined
were taken. Furthermore, oil globules were not always apparent in the
large cells, and for this reason they have not been designated as “pulcher-
rima cells” in the above tabulation. It is believed, however, that the large,
thick-walled cells observed, either with or without oil globules, may be
regarded as similar to those cells interpreted by Windisch as ascogenous
cells, for many of them were found to possess attached daughter cells.
Likewise, the attached daughter cells are considered to be asci, for several
of these were found to contain spore-like bodies.

Because of the occurrence of large, thick-walled cells and attached
daughter cells in some of the white variants, this tabulated evidence cannot

“In 1918 Beijerinck, working with mass cultures, noted that certain cells of Saccharo-
myces pulcherrimus (Toruopsis pulcherrima) failed to produce fat globules. This was
considered to involve a mutational change; and such variants were designated as
S. pulcherrimus secundarius. Although fairly constant, these so-called mutants were
found to show in some cases a tendency to revert to the normal condition and were
apparently not related to any constant directional change in pigmentation. Hence it is
believed that this behaviour is not comparable to that which has been found in the
present study, although the possibility exists that the results would have been similar if
Beijerinck had employed single cell technique.

ROBERTS: TORULOPSIS AND TAPHRINA 367

be regarded as implying a definite, constant distinction between the two
cultural types on the basis of cellular morphology. It can be regarded,
however, as indicative of different morphological tendencies in the two
types. In view of Windisch’s interpretation of these cellular structures,
this evidence raises interesting implications in regard to the relationship
between the life cycle of this organism and its cultural variation. Certainly,
a tendency for the sexual stage to occur more frequently in the red cultural
type is indicated here, but the complete answer to this problem must await
the results of further investigations.

Although some secondary papillae and roughened surfaces were evident
in several single cell cultures of T. pudcherrima, no detailed study of them was
undertaken. It should be mentioned, however, that one pure white single
cell colony produced two large craters, and microscopic examination of
cells from both of them revealed the presence of long, septate mycelial
strands. This is in confirmation of Punkari and Henrici’s investigations
(1935), in which they noted that craters were only observed in white
colonies and stated that ‘“. . . microscopic examination of colonies with
craters always showed the presence of mycelium, which was not found in
smooth colonies.”’

In addition, twelve white colonies or sectors, when examined micro-
scopically, were found to consist of polymorphic cells, or of cells exhibiting
a pseudomycelial tendency, and it was further noted that these were always
associated with a roughened surface characterized by the formation of
numerous small papillae.

COMPARATIVE CELLULAR MORPHOLOGY OF THE CULTURAL TYPES OF T.
DEFORMANS

Similar microscopic investigations were carried out on all of the single
cell colonies of JT. deformans in order to determine whether any variation
in cellular morphology existed, and if so, whether it bore any relation to
the formation of the four cultural types previously designated on the basis
of pigmentation as a, b, c, and d. All colonies were examined after they had
been growing on potato dextrose agar slants for one month or more, and
recorded descriptions were made of the general appearance of the cells,
including the occurrence of large, thick-walled cells either with or without
attached daughter cells. In view of the fact that spore-like bodies were
found in several of these attached daughter cells, these cellular structures
were regarded as being indicative of the sporulation process and were con-
sequently interpreted as ascogenous cells, attached asci, and spores. As in
the microscopic investigations of T. pulcherrima, some difficulty was en-
countered in accurately interpreting all structures observed because of the
occurrence in some colonies of cells of transitional shape and size.

In general, it can be stated that these cellular structures indicative of
sporulation occurred as abundantly as in T. pulcherrima, being found in
62 out of a total of 159 cultures examined. (Compare with tabulation totals
for IT. pulcherrima, page 366.) In contrast to the observations on Torulopsis,

368 FaRLowlA, VoL. 2, 1946

however, no evidence was obtained in this study for a differentiation of
the cultural types on the basis of cellular morphology, for these structures
were found in representatives of all four cultural types. They appeared to
be somewhat more abundant in the normal a type, but it is believed that
they were found in sufficient numbers in the variants 6, c, and d to preclude
any generalization as to a constant relationship between sporulation and
pigmentation.

These results seem to indicate, therefore, that in T. deformans cellular
morphology, with regard to the occurrence of structures indicative of
sporulation, appears to be more constant than in T. pulcherrima, in the
sense that it shows no clear-cut differentiation into two types, each associ-
ated with a distinct pigmented cultural type. On the contrary, in Taphrina,
the occurrence of cells indicative of sporulation may appear in any culture
regardless of its pigmentation. It is possible that their occurrence may be
correlated with the age of the culture, but this is offered only as a theo-
retical consideration, for no detailed study was made of this phase of the
problem.

A tendency for mycelial production in culture, which is believed to be
more pronounced than has previously been reported, was also noted in
T. deformans. Long, septate hyphal strands very similar in appearance to
the intercellular mycelium of this organism in host tissue were observed
(Plate IIT, Figs. 1, 2). This tendency was noted in the normal type, as well
as in the } variant and was not found to be associated in every instance
with a roughened surface.

Other cultures of T. deformans were composed of elongated, rod-shaped,
or otherwise irregularly shaped cells which were either single or in pairs.
In addition, the formation of primitive pseudomycelium in the form of
short chains was noted by both the normal and irregularly shaped cells.
These pseudomycelial and polymorphic tendencies occurred in the normal
pigmented type as well as in the 6 variant and were found either in asso-
ciation with roughened surfaces or independent of them. A tendency to
produce elongated or rod-shaped cells was also noted in some of the smooth
colonies of the yellow variant d.

NUCLEAR CONDITION

Considerable difficulty was encountered in obtaining a satisfactory
nuclear stain for T, pudcherrima. The most successful stain employed was
Laustsen’s modification of the Feulgen technique, as described by Winge
(1935). Windisch had apparently obtained very satisfactory results with
this method, although no photomicrographs of stained preparations were
included in his paper. He states (1940): “Kernfarbungen nach Winge-
Feulgen mit Fuchsin erwiesen die Zweikernigkeit aller vegetativen Formen
sowie der Ascusmutterzellen.”

In the present study, cells from young, active cultures were the only ones
stained satisfactorily by this method, and consequently no information as

RoBERTS: TORULOPSIS AND TAPHRINA 369

to the nuclear condition of the large, thick-walled cells (ascogenous cells or
ascus mother cells) occurring in old cultures was obtained. However, it
was found that all vegetative cells, whether from mass or single cell,
pigmented or non-pigmented cultures, were stained in such a way that
they exhibited what is believed to represent a constant uninucleate con-
dition. This is in direct contrast to the binucleate condition caganien by
Windisch.

Although some differences in interpretation of the stained bodies could
be made, it will be seen from Plate III, Figs. 3, 4 that these stained prep-
arations cannot possibly be regarded as being indicative of a binucleate
condition, for each cell contains a single, deeply stained body. The difficulty
in interpretation appears to involve the nucleoli and the nuclear membranes.
A comparison of Plate III, Figs. 3, 4 with Figs. 11, 12, 13, 14, and 15 of
Plate II in Winge’s paper (1935) shows a definite similarity in the results
of employing this staining procedure, notwithstanding the fact that two
different yeasts were involved. Winge interprets, without question, the
deeply stained bodies as nuclei, but it is possible that the stained bodies in
our preparations may be nucleoli, because of the fact that some of them
appear to be surrounded by a definite, clear area which is not apparent in
Winge’s photomicrographs. This clear area may possibly represent the
extent of a single nucleus and the enclosed stained body, the nucleolus, but
because no membrane is visible at the periphery of the clear area, this
interpretation is doubtful. Regardless of the interpretation of the nuclear
details, the results of this staining method clearly indicate that the young
vegetative cells of T. pulcherrima are uninucleate.

In contrast, all of Windisch’s stained preparations revealed a . binucleate
condition of the cells. Consequently, he believed that the uninucleate,
haploid phase of the life cycle must be of very short duration, the deriva-
tives of the ascospores copulating very soon after spore discharge, resulting
in the establishment of the binucleate stage. Windisch actually observed
this fusion of ascospore derivatives (which appeared to involve dissimilar
partners and therefore can be considered as heterogamous copulation), but
- it should be emphasized that although this plasmogamy was found to result
in the dicaryon phase, the actual point in the life cycle at which karyogamy
occurred was not ascertained. Windisch had consistently observed a bi-
nucleate condition in the ascogenous cells and believed that these two
nuclei corresponded to the nuclear pair observed in the vegetative cells.
The possibility that the two nuclei were diploid daughter nuclei of a zygote
nucleus was offered, but because, under certain conditions, the ascogenous
cells could reproduce vegetatively by budding instead of forming asci, led
Windisch to prefer the assumption that the two nuclei in the ascogenous
cell were haploid. Because uninucleate ascogenous cells were never ob-
served, Windisch proposed the idea that karyogamy in T. pulcherrima may
occur in the ascus, and as evidence of a similar occurrence in the Taphri-
nales, he referred to the statement of Kniep (1928) that in Taphrina

370 FaRLowIA, VOL. 2, 1946

nuclear fusion occurs in the ascus. However, it has recently been definitely
established by Martin (1940) that karyogamy in T. deformans occurs only
in the ascogenous cells, a fact which has been generally regarded as typify-
ing the genus as a whole (Eftimiu 1927, Gdumann and Dodge 1928).

It is therefore believed that Windisch’s assumption of a similarity be-
tween T. pulcherrima and Taphrina spp. based upon the nuclear condition
in the life cycles of these organisms is incorrect. The possibility exists, of
course, that karyogamy does occur in the ascogenous cells of 7. pulcherrima
and that’ Windisch had failed to observe it; but until this fact is actually
known, a concept of a relationship between Torulopsis and Taphrina on the
basis of the nuclear behaviour known at present does not appear to be
justified. .

The Laustsen modification of the Feulgen technique was not found to be
satisfactory for demonstrating the nuclei in cells of T. deformans. None of
the stains employed was found to be completely successful, although the
use of Mayer’s haem-alum demonstrated the presence of a single stained
body in some of the sprout conidia from young cultures. Although no
detail was apparent in any of these bodies, they were regarded as
being indicative of a uninucleate condition, in view of the fact that sev-
eral investigators (Fitzpatrick 1934, Martin 1940, Mix 1935) have
found this to be the case in the majority of cells from young cultures of
T. deformans.

Because, as in Torulopsis, only active cells from young cultures were
successfully stained, no information on the nuclear condition of the larger,
thick-walled cells (ascogenous cells) was obtained, although Mix reported
them as being either uni- or binucleate (1935). In addition, Fitzpatrick
(1935) observed some binucleate sprout conidia taken from fairly old cul-
tures, while Wieben (1927) found in TapArina spp. binucleate conidia fol-
lowing copulation. In the present study, no binucleate cells were observed.

PHYSIOLOGICAL TESTS

Although comparative observations of the microscopic and macroscopic
appearances of the two fungi in culture represented the main problem in
the present investigation, a limited number of tests were performed in order
to determine some of the physiological differences between the two organ-
isms as well as between the variant cultural types of each organism.

One of the most important distinctions between the physiology of T. de-
formans and T. pulcherrima is that the former has been reported to lack
the ability to produce alcoholic fermentation (Mix 1924), while 7. pul-
cherrima is a species capable of fermenting glucose, fructose, and mannose
(Lodder 1934). Mix in 1924 used 10% solutions of dextrose, sucrose, and
maltose in potato broth in his fermentation tests and had obtained com-
pletely negative results. Confirmation of this work was obtained by the
failure of active mass and single cell cultures of JT. deformans to produce
any fermentation of 2% solutions in yeast juice of galactose, glucose,

eae |

ROBERTS: TORULOPSIS AND TAPHRINA Spl

maltose, lactose, and sucrose. In addition, the three variant types, 8, c,
and d were found unable to cause fermentation of any of these sugars.®

Because of this lack of fermentative ability in Taphrina, tests were per-
formed by means of the auxanogram method (Lodder 1934) in order to
determine what sugars this organism was capable of assimilating under
aerobic conditions. Positive results were obtained with glucose and sucrose,
and negative results with galactose, lactose, and maltose.

T. pulcherrima, on the other hand, was found to be capable of fermenting
a 2% solution of glucose. In addition, the appearance in some of the
sucrose solutions of a few gas bubbles was noted, which may be regarded
as being indicative of a slight tendency of this organism to ferment this
sugar. However, the explanation for the occurrence of gas bubbles in the
sucrose solutions may lie in the method of sterilization of the liquid media.
As Mudge (1917) pointed out, disaccharides may. become hydrolyzed by
sterilization in both the autoclave and the Arnold sterilizer, resulting in the
formation of monosaccharides in the medium. Mudge found this to be the
case with maltose and lactose, and the possibility exists that the sucrose
solutions used in the present tests were sterilized for a length of time suffi-
cient to induce a partial breakdown into the monosaccharides glucose and
fructose, both of which are fermented by this organism. It should be noted,
however, that, although Lodder in her standard work on generic and specific
determination of the imperfect yeasts lists glucose (fructose, mannose) as
the only sugar fermented by 7. pulcherrima, Grosbiisch (1915) found that
a strain of Torula rubefaciens (Torulopsis pulcherrima) was able to cause
a very slight fermentation of sucrose.

Galactose, lactose, and maltose gave negative results in all fermentation
tests with T. pulcherrima. In addition, no differences in fermentative ability
were found between the red and the white cultural types.

Nitrogen utilization tests were undertaken by means of the auxanogram
technique. Both the red and the white cultural types of T. pulcherrima
were found capable of utilizing only asparagine and peptone, although
Lodder had obtained, in addition, positive results with ammonium sulfate
and urea. In this connection, it is worthy of note that in the original un-
published identification of this isolate of T. pulcherrima, ammonium sulfate
was found by E. M. Mrak to be slowly utilized, while urea was not utilized
at all.

No positive results for the utilization of any nitrogen-containing sub-
stances by T. deformans were obtained by means of the auxanogram
method. In view of the fact that peptone is generally utilized as a nitrogen
source by yeasts, as well as ammonium sulfate (Henrici 1941), these results
are surprising. Moreover, in the sugar auxanograms, ammonium sulfate
was the only nitrogen source available in the basal medium and therefore
must have been utilized by T. deformans as evidenced by its growth in the

®° According to the well-known “Kluyver-Dekker laws” (as cited by Henrici 1941),
any yeast which can ferment glucose can also ferment fructose and mannose. For this
reason, fructose and mannose were not included in the fermentation tests.

372 FARLOWIA, VoL. 2, 1946

presence of glucose and sucrose. It is concluded that the complete explana-
tion of these negative results must therefore await the results of further
investigation. ;

In general, the physiological tests, although limited in scope, appear to
point to two facts worthy of consideration: 1) 7. pudcherrima can be dis-
tinguished from T. deformans on the basis of its ability to ferment certain
sugars, and 2) on the basis of these tests, no differences in physiological
behaviour were found between the two cultural types of T. pulcherrima —
and among the four cultural types of T. deformans.

DISCUSSION AND CONCLUSIONS

As evidenced by cultural similarities, the results of this investigation
point to the apparent interrelationship of a saprophytic yeast, Torulopsis
pulcherrima, and a parasitic fungus possessing a yeast-like saprophytic
stage, Taphrina deformans.

It will be recalled that this comparative study was undertaken with the
hope of obtaining additional information which could be used either in
substantiation or in rejection of Windisch’s concept of a relationship be-
tween T. pulcherrima and the genus Taphrina and his proposal to incor-
porate 7. pulcherrima in the order Taphrinales as Candida pulcherrima.
This proposal was based upon the similarities between the observed life
cycle, including sporulation, of T. pulcherrima and of C. tropicalis in cul-
ture and the previously described life cycles of Taphrina spp. in their para-
sitic, non-culturable stages. Windisch made no recorded observations on
the cultural characteristics of any species of Taphrina. In the present study,
the genus Candida was not investigated, and while T. deformans was the
only species of Taphrina used, it is believed to be representative of the
general cultural behaviour of the genus as a whole.

For purposes of comparison, the information obtained from a study of |
the two fungi in culture may be tabulated as follows:

That the two fungi are, in general, quite similar in cultural behaviour
is clearly evident from the above tabulation. In both organisms cultural
variation may occur in the form of variously pigmented, transmissible or
non-transmissible growth types; cellular morphology is of a similar type,
pseudomycelium or true mycelium may be found, and cellular structures
indicative of sporulation frequently occur. This latter fact is regarded as
one of importance for the following reasons: 1) it confirms Windisch’s
report of sporulation in T. pulcherrima and consequently justifies the re-
moval of this organism from the imperfect yeasts; 2) it supports Mix’s
observations on the occurrence of the diplophase and sexuality in T. de-
formans in culture; and 3) it offers additional support for the view of a
general relationship between 7. pulcherrima and the Taphrinales. More-
over, the finding of pseudomycelium and true mycelium in T. pulcherrima
is indicative of the fact that it does not properly belong in the sub-family
Torulopsidoideae as defined by Lodder (1934).

RoBERTS: TORULOPSIS AND TAPHRINA 373

T. pulcherrima

T. deformans

Character of cultural
growth

Characteristic color of
colony

Occurrence of variation in
pigmentation

Occurrence of transmis-
sible variants

Occurrence of temporary
variants

General appearance of cells
from young cultures

General appearance of cells
from old cultures

Occurrence of cellular
structures indicative of
sporulation

Occurrence of differences
in cellular morphology
among the various cul-
tural types

Occurrence of a mycelial
tendency

Nuclear condition of
young vegetative cells

Fermentative ability

Occurrence of physiologi-
cal differences among the
various cultural types

Yeast-like

Red — non-carotinoid,
diffusible pigment

White, brown

Predictable occurrence of
white sectors

Unpredictable, rare occur-
rence of brown sectors

Oval-globose, thin-walled,
budding cells and “pul-
cherrima cells”’

Mainly “pulcherrima
cells”

Abundant in old cultures
and interpreted as asco-

genous cells, attached asci,
and ascospores

Evident in red and white
cultural types

Present
Uninucleate
Ferments glucose

Not evident

Yeast-like

Pink — non-carotinoid,
non-diffusible pigment
White, yellow, and vari-
ous modifications of pink
Unpredictable occurrence
of yellow and white sectors
Unpredictable, frequent
occurrence of sectors of
various modifications of
pink

Oval-globose, thin-walled,
budding cells

Similar large, thick-walled
cells but with many oil
globules

Abundant in old cultures
and interpreted as asco-
genous cells, attached asci,
and ascospores

Not evident

Present
Uninucleate
Incapable of fermenting

glucose or any other sugar
Not evident

fee

It will be noted from the above tabulation that the chief distinguishing

points between the two organisms appear to involve the type of pigmenta-
tion and the fermentative ability, but it is known that both fungi may
produce a non-pigmented growth type and that the genus Torulopsis con-
tains several species which lack the ability to produce alcoholic fermentation
(Lodder 1934). T. deformans, although customarily isolated from its host
by means of discharging ascospores, also may be isolated in its overwinter-
ing, saprophytic stage from peach bark or buds; it is possible that confusion
in identification would result if such isolates were studied according to the
present methods for the determination of yeasts, without the use of inocula-
tion tests.

Thus, in partial support of Windisch’s views, it can be said that the

374 FarLowIA, VoL. 2, 1946

cultural similarities evident between T. pulcherrima and T. deformans are
indicative of their interrelationship, an assumption which is believed to be
further justified by the sporulation phenomenon, which was previously
reported for T. pulcherrima by Windisch (1938, 1940) and which, in the
present study, appears to have been confirmed for that species, and, in
addition, to have been found in 7. deformans.

However, certain other facts, as yet unexplained, have been revealed by
this comparative investigation which do not lend themselves to a complete
substantiation of the proposals of Windisch.

Specifically, all of the evidence obtained in this study is indicative of a
constant uninucleate condition in the young vegetative cells of both T.
deformans and T. pulcherrima, in contrast to Windisch’s report of a con-
stant binucleate condition in T. pulcherrima (exclusive of an assumed, very
short, uninucleate phase). This discrepancy, coupled with Windisch’s belief
that karyogamy in this fungus occurs in the ascus, does not support the
concept of a relationship between Torulopsis and Taphrina based upon
nuclear behaviour. It is believed that perhaps the explanation for this
discrepancy may lie in the use of single cell technique in the present study,
and the subsequent distinctive type of variation in pigmentation which it
revealed. It was found, in contrast to the investigations of Punkari and
Henrici, that dissociation in T. pulcherrima into the two cultural types,
regardless of whether it is an expression of the dual phenomenon, was
apparently correlated with variation in cellular morphology, and that these
distinct morphological tendencies seemed to bear some relation to the
sporulation process, and therefore to sexuality. Consequently, until further
cytological investigations of this organism reveal the complete facts sur-
rounding the nuclear cycle, it seems unwise to point to the observed uni-
nucleate condition of both Taphrina and Torulopsis in culture as being
indicative of their interrelationship.

Thus, it is believed that these preliminary cultural investigations lend
support to the ideas advanced by numerous investigators in regard to a
general relationship between the yeasts and the Taphrinales. It is further
believed that they also support the recent, more specific proposal of Win-
disch in regard to a relationship between the Taphrinales and T. pulcher-
rima, as well as the removal of the latter from the imperfect yeasts. How-
ever, because our knowledge of the cytology, sexuality, and variation of
T. pulcherrima is still in an imperfect state, it is believed that. further
investigations of a wide number of isolates of this species are needed before
it can be placed with certainty into any of the present schemes of classifica-
tion of the perfect fungi.

It is therefore concluded that, although our present knowledge points to
a relationship between T. pulcherrima and the Taphrinales, until the life
cycle of the former has been completely worked out, especially with regard
to the nuclear behaviour and the relationship between sexuality and cultural
variation, the similarities recognized at present are not sufficient grounds
for adopting Windisch’s proposals of transferring T. pulcherrima to the

RoBERTS: TORULOPSIS AND TAPHRINA 375

genus Candida and of incorporating it in the Taphrinales in a new family,
the Candidaceae.

It should be added that no evidence was obtained from either organism
which would fully support the findings of Todd and Herrmann in regard to
their assumed life cycle of Cryptococcus hominis (Torulopsis neoformans )
and that consequently the transfer of T. pulcherrima to the genus Debaryo-
myces has not been considered.

SUMMARY

1. Both Torulopsis pulcherrima and Taphrina deformans presented a
similar yeast-like appearance when grown at room temperature on potato
dextrose agar.

2. Both fungi produced a reddish or pinkish non-carotinoid pigment,
that of 7. pulcherrima being diffusible and that of T. deformans being
non-diffusible.

3. In T. pulcherrima a predictable mutation-like change involving the
production of white sectors occurred with great regularity in pigmented
colonies. In T. deformans unpredictable sectoring involving a color change
from pink to yellow or white occurred frequently but sporadically. The
white variant of 7. pulcherrima and the yellow and white variants of T.
deformans were transmissible but did not possess complete stability.

4. Cellular morphology was essentially similar in cultures of both or-
ganisms, which were composed of single, ovoid to globose, budding cells.
In addition, a mycelial tendency was noted in both organisms.

5. In old cultures of both fungi, cellular structures indicative of sporula-
tion were noted and were interpreted as ascogenous cells, asci, and asco-
spores.

6. In T. pulcherrima a tendency for a differentiation of the red and the
white cultural types on the basis of cellular morphology was noted and was
found to be correlated with the sporulation phenomenon. In T. deformans
no such morphological differentiation among the three cultural types was
evident. |

7. A constant uninucleate condition was found in the young vegetative
cells of both fungi.

8. T. pulcherrima was found capable of fermenting glucose, while T.
deformans was found incapable of causing fermentation. No evidence was
obtained from either organism for the occurrence of any physiological
differences among their various cultural types.

9. On the basis of these results, the interrelationship of T. pulcherrima
and T. deformans is discussed, with especial reference to the recent proposal
of Windisch in regard to the removal of T. pulcherrima from the imperfect
yeasts to the Taphrinales.

DIVISION OF PLANT PATHOLOGY .

UNIVERSITY OF CALIFORNIA
BERKELEY, CALIFORNIA

376 FarLowI1A, Vov. 2, 1946

LITERATURE CITED

De Bary, A. Vergleichende Morphologie der Pilze, Mycetozoen, und Bacterien. Engel-
mann, Leipzig, 1884.

Beijerinck, M. W. Levures chromogénes. — Nouvelle réaction biologique du fer. Arch.
Neerland. Physiol. Homme et Anim. 2: 609-615. 1918.

Bessey, Ernst A. A Textbook of Mycology. Blakiston, Philadelphia, 1935.

——. Some problems in fungus phylogeny. Mycologia 34: 355-379. 1942.

Castelli, Tommaso. Considerazioni sulla Torulopsis pulcherrima. Arch. Mikrobiol. 11:
126-136. 1940.

Chamberlain, Charles J. Methods in Plant Histology. 5th rev. ed., University of
Chicago, 1932.

Christensen, J. J. The origin of parasitic races of phytopathogenic fungi through
mutation. Jn The Genetics of Pathogenic Organisms, Science Press (A.A.A.S.),
pp. 77-82. 1940.

Eftimiu, P. Contribution 4 l'étude cytologique des Exoascées. Le Botaniste 18: 1-154.
1927.

Fitzpatrick, R. E. The life history and parasitism of Taphrina deformans. Sci. Agr.
14: 305-326. 1934.

Gaumann, Ernst A. & Carroll W. Dodge. Comparative Morphology of the Fungi.
McGraw-Hill, N. Y., 1928.

Giordano, A. Studio micologico del Debaryomyces neoformans (Sanfelice) Red., Cif.
et Giord. e significato delle specie nella patologia animale. Mycopathologia 1: 274—
304. 1938-39.

Grosbiisch, J. Uber eine farblose, stark roten Farbstoff erzeugende Torula. Centbl.
Bakt. II, 42: 625-638. 1915.

Guilliermond, A. La sexualité, le cycle de développement, la phylogénie, et la classi-
fication des levures d’apres les travaux récents. Extrait des Ann. des Fermentations
2: 129, 257, 474, 540 (Paris, 1937).

Hansen, E. C. Grundlinien zur Systematik der Saccharomyceten. Centbl. Bakt. IJ,
12: 529-538. 1904.

Hansen, H. N. The dual phenomenon, Mycologia 30: 442-455. 1938.

. Heterocaryosis and variability. Phytopath. 32: 639-640. 1942.

& R. E. Smith. The mechanism of variation in imperfect fungi: Botrytis

cinerea. Phytopath. 22: 953-964. 1932.
& W. C. Snyder. The dual phenomenon and sex in Hypomyces solani {.
cucurbitae, Amer. Jour. Bot. 30: 419-422. 1943.

Henrici, Arthur T. The yeasts, genetics, cytology, variation, classification and identi-
fication. Bact. Rev..5: 97-179. 1941.

Kniep, Hans. Die Sexualitat der niederen Pflanzen. Fischer, Jena, 1928.

Koch, L. W. Studies on the overwintering of certain fungi parasitic and saprophytic
on fruit trees. Canad. Jour. Res. 11: 190-206. 1934.

Lindner, P. Wochenschrift f. Brauerei, 4: 853. 1887. (Cited from Lodder.)

Lodder, J. Die Hefesammlung des “Centraalbureau voor Schimmelcultures.” Beitrage
zu einer Monographie der Hefearten. II. Die anaskosporogenen Hefen. Erste Halfte.
K. Akad. Wet., Verhand. (2nd Sect.) D1. 32: 1-256. 1934.

Martin, Ella M. Cultural and morphological studies of some species of Taphrina.
Phytopath. 15: 67-76. 1925.

. The morphology and cytology of Taphrina deformans. Amer. Jour. Bot.

27: 743-751. 1940.

Mix, A. J. Biological and cultural studies of Exoascus deformans. Phytopath. 14: 217-
233. 1924.

. Further studies on the Exoascaceae. Phytopath. 19: 90. 1929. (Abstract).
————.. The life history of Taphrina deformans. Phytopath. 25: 41-66. 1935.
Mrak, E. M., H. J. Phaff, & H. C. Douglas. A sporulation stock medium for yeasts

and other fungi. Science 96: 432. 1942.

RoBERTS: TORULOPSIS AND TAPHRINA oii

Mudge, Courtland S. The effect of sterilization upon sugars in culture media. Jour.
Bact. 2: 403-415. 1917.

Pierce, Newton B. Peach leaf curl: its nature and treatment. U.S.D.A., Div. Veg.
Physiol. & Path. Bull. 20: 204 pp. 1900.

Porchet, Berthé. Contribution a l’étude de la levure Torulopsis pulcherrima. Ann. des
Fermentations 4(7): 385-405. 1938.

Punkari, Laila & Arthur T. Henrici. A study of variation in a chromogenic asporo-
genous yeast. Jour. Bact. 26: 125-138. 1933.

. Further studies on spontaneous variations of Torula pulcherrima. Jour.
Bact. 29: 259-267. 1935.
Rawlins, Thomas E. Phytopathological and Botanical Research Methods. Wiley,
: N. Y., 1933.

Redaelli, P., R. Ciferri, & A. Giordano. Debaryomyces neoformans (Sanfelice)
nobis, nov. comb., pour les espéce du groupe Saccharomyces hominis-Cryptococcus
neoformans-Torula histolytica. Boll. Sez. Ital., Soc. Intern. Microbiol. I-II: 1-7.
1937. (Cited from Henrici). [Corrected citation, 9(1-2): 24-28. 1937]

Reess, M. Botanische Untersuchungen uber die Alkoholgarungspilze. Leipzig, 1870.

Ridgway, Robert. Color Standards and Color Nomenclature. Author, Washington,
D. C., 1912.

Snyder, William C. & H. N. Hansen. The importance of variation in the taxonomy
of the fungi. 6th Pacific Sci. Congr., Proc. 4: 749-752. 1939.

Todd, Ramona L. & Walter W. Herrmann. The life cycle of the organism causing
yeast meningitis. Jour. Bact. 32: 89-103. 1936.

Wieben, Magdalena. Die Infektion, die Myzeliiberwinterung, und die Kopthton bei
Exoasceen. Forsch. Geb. Pflanzenkr. u. Immunitat 3: 139-176. 1927.

Windisch, Siegfried. Zur Kenntnis der Ascosporenbildung bei Torulopsis pulcherrima
(Lindner) Saccardo. Arch. Mikrobiol. 9: 551-554. 1938.

Entwicklungsgeschichtliche Untersuchungen an Torulopsis pulcherrima
(Lindner) Saccardo und Candida tropicalis (Castellani) Berkhout. Ein Beitrag zur
Systematik der Garungsmonilien. Arch. Mikrobiol. 11: 368-390. 1940.

Winge, O. On haplophase and diplophase in some Saccharomycetes. Compt. Rend.

Trav. Lab. Carlsberg, Ser. Physiol. 21: 77-111. 1935.

ae i : fs
1 A Cs

378

Fig.

Fig.

Fig.
Fig.

Fig.

Fig.

Fig.

Fig.
Fig.

Fig.

Fig.
Fig.

Fig.

Fig.

FarLowlA, VoL. 2, 1946

EXPLANATION OF PLATE I

1. 7. deformans. Glass needle of micromanipulator drawing out a single cell from
the margin of a colony. (135x).

2. T. pulcherrima. A day-old colony originating from a single cell withdrawm
from the margin of the parent colony by means of the glass needle of the micro-
manipulator. (135x).

3. T. deformans. A 3-month-old colony on potato dextrose agar originating from
a single ascospore. Note abundant sectoring. (Natural size).

4. T. deformans. A 3-month-old colony on potato dextrose agar originating from
a single ascospore. Note absence of sectoring. (Natural size).

5. 7. pulcherrima. A 3-month-old pigmented colony on potato dextrose agar
originating from a mass transfer of cells from the pigmented portion of the original
stock culture. Note formation of white sectors. (Natural size).

6. T. pulcherrima. A 3-month-old white colony on potato dextrose agar originat-
ing from a mass transfer of cells from a white sector in the original stock culture.
Note absence of pigmentation. (Natural size).

7. T. pulcherrima. A month-old pigmented colony on potato dextrose agar origi-
nating from a single cell of the pigmented portion of a sectored colony. Note
formation of white sectors. (Natural size).

8. T. pulcherrima, A month-old white colony on potato dextrose agar originating
from a single cell of a white sector. Note absence of pigmentation. (Natural size).
9. T. deformans. Normal type of vegetative cells (sprout conidia) from a 2-
month-old single cell culture. Unstained. (588x).

10. T. pulcherrima. Typical “pulcherrima cel!s” from a month-old, pigmented,
single cell culture. Note single oil globules, thick walls, and attached cell wall
remnants. Unstained. (1200x).

11. 7. deformans. Large thick-walled cell from a 4-month-old mass culture. Note
numerous oil globules and the attached remnant of a cell wall. Unstained. (1101x).
12. T. pulcherrima, ‘“Pulcherrima cells” from a pigmented, month-old, single cell
culture, showing a papillate protrusion. Unstained. (1200x).

13. T. pulcherrima. A “pulcherrima cell” from a pigmented, 4-month-old, mass
culture, showing the attached daughter cell. Note that both mother and daughter
cells contain a single oil globule. Unstained. (1101x).

14. T. pulcherrima. A large “pulcherrima cell” (11.2 « in diameter) from a culture
grown on peach leaf agar, showing an attached, intact, empty daughter cell. Un-
stained. (1200x).

‘ROBERTS: TORULOPSIS AND TAPHRINA 379

Pirate I

380

Fig.

Fig.

Fig.

Fig.

Fig.

Fig.

Fig.

Fig.

Fig.

Fig.

FaRLowI1A, VoL. 2, 1946

EXPLANATION OF PLATE II

ig.1. 7. pulcherrima. A large ‘“pulcherrima cell” from a pigmented, 2-month-old,

single cell culture, showing an attached, intact, empty daughter cell. Unstained.
(1200x).

-2. T. pulcherrima. A “pulcherrima cell” from a pigmented, 4-month-old, mass

culture, showing an attached daughter cell containing one spore-like body. Note
the single oil globules in the mother cell and in the spore-like body. Unstained.
(1101x).

3. T. pulcherrima. “Pulcherrima cells” from a pigmented, 3-month-old, mass cul-
ture, showing one cell which possesses an attached daughter cell containing two
spore-like bodies. Unstained.

4. T. pulcherrima. A “pulcherrima cell” with a single oil globule from a pigmented,
4-month-old, mass culture, showing the attached remnant of a daughter cell. Un-
stained. (1101x).

5. T. pulcherrima. A “pulcherrima cell” with a single oil globule and the attached
remnant of a daughter cell. From same culture noted in Fig. 4. Unstained. (1200x).
6. YT. pulcherrima. Growth after 24 hours from a single thick-walled cell with
attached empty daughter cell, both of which had been withdrawn together from
the margin of a colony. Note that only 2 cells were produced during this time.
Unstained. (588x).

7. T. pulcherrima. Cells from a non-pigmented, single cell colony which appear
to show evidences of copulation. The long outgrowths may be regarded as con-
jugation tubes. Stained with cotton blue in lactophenol. (867x).

8. T. deformans. Two large cells, one with many oil globules and a papillate pro-
trusion, and the other with a single oil globule and an attached, intact, empty
daughter cell. From a 4-month-old mass culture. Unstained. (1101x).

9. ¥. deformans. Cell from a 6-month-old single spore culture, showing a thick-
walled cell with an attached daughter cell containing a single oil globule (or per-
haps a spore-like body). Unstained. (1101x).

10. T. deformans. Large cell with attached daughter cell containing 2 spore-like
bodies. Note oil globules within spore-like bodies. From same culture noted in
Fig. 8. Unstained. (1101x).

11. 7. pulcherrima. Typical “pulcherrima cells” from a pigmented, month-old,
single cell colony. Unstained. (1200x).

12. T. pulcherrima. Thin-walled vegetative cells from a non-pigmented, 25-day-old,
single cell colony. Unstained. (1200x).

381

ToRULOPSIS AND TAPHRINA

ROBERTS

Pirate II

382 FaRLowIA, VoL. 2, 1946

EXPLANATION OF PLATE III

Fig.1. 1. deformans. .Mycelial tendency occurring in a 2-month-old, single spore,
smooth colony. Unstained. (588x).

Fig. 2. 7. deformans. Mycelial tendency occurring in a rough sector of a 3-month- old,
single cell colony. Stained with cotton blue in lactophenol. (588x).

Fig.3. 7. pulcherrima. Nuclear staining of cells from a 5-day-old, pigmented, mass
culture. (1200x).

Fig.4. 7. pulcherrima. Nuclear staining of cells from a 5-day-old, non-pigmented,
single cell culture. (1200x).

383

ToRULOPSIS AND TAPHRINA
Prate III

ROBERTS

2(3) :385-437 FARLOWIA | January, 1946

THE CYPERICOLOUS AND JUNCICOLOUS SPECIES
OF SCLEROTINIA

H. H. WHerzet !

During the spring and summer of 1930 I took the opportunity afforded
by sabbatic leave to collect and study species of the Sclerotiniaceae in
Europe. My travels carried me through England, Holland, Switzerland,
France, Germany, Denmark, Sweden and Norway. Through the kind co-
operation of colleagues ? in these countries I was enabled to make invaluable
collections and studies of many species in their natural habitat. I was also
afforded every courtesy and assistance in the examination of specimens of
these cup fungi preserved in the herbaria of most of the important botanical
institutions in the countries visited.

Stimulated by my early studies on the species of Sclerotinia found in
northeastern United States on species of Carex (Whetzel 1929) I took
special pains to collect and study the species of this genus occurring on
sedges and rushes in western Europe. I was exceptionally fortunate in
finding in nature most such species recorded by European mycologists.

A manuscript presenting the results of my studies on these species was
partially prepared during my stay in Europe. On my return, teaching
duties prevented the completion of this paper, and only recently have I
been able to take up the work again and complete the studies which are
here presented. This delay has not been without its compensations in
certain respects. During the years that have passed, while much of the
freshness and some of the details of my European observations have faded

*This manuscript was received and edited before Professor Whetzel’s death on No-
vember 30, 1944. Realizing that he might not live to see the appearance of his paper,
the author made all arrangements for its publication, including a generous gift to cover
the extra cost of the halftone plates.

* Acknowledgments: The researches on which this paper is based, especially those
undertaken in Europe in 1930, were made possible largely by grants from the Heckscher
Foundation for the Advancement of Research at Cornell University. I am especially
indebted to Dr. Cynthia Westcott, who as my research assistant gave invaluable help
on my European trip. For access to herbarium specimens and for expert advice and
assistance I am greatly indebted to Drs. David H. Linder and W. L. White of the
Farlow Herbarium; to Dr. John A. Stevenson of the U.S.D.A.; to Dr. W. G. Solheim
of the University of Wyoming; to Dr. J. Ramsbottom of the British Museum; to Dr.
A.-D. Cotton and Miss E. M. Wakefield of the Royal Herbarium, Kew; to Dr. J. A.
Nannfeldt, University of Upsala; and to Mr. Ernst Gram, Dr. C. Ferdinandsen, and
Mr. Johs. Grgntved of the University of Copenhagen. My best thanks are also due
Miss Charlotte Dill and Mr. Carlos Garcés for preparation of the drawings for the text
figures and to Dr. Rolf Singer of the Farlow Herbarium who has kindly prepared the
Latin descriptions. Mr. W. R. Fisher prepared the photographs. The writer is indebted
to so many other friends.and colleagues both in this country and in Europe for assistance
and favors that it is useless to attempt to list them all here. To everyone who has thus
contributed to the consummation of these investigations I here express my thanks and
gratitude.

385

386 Fartowia, VoL. 2, 1946

from my memory, there have come to hand, now and then, specimens and
information on species of this group that have corrected some of my earlier
conclusions, confirmed others, and have added three species to those known
to me at the time of my 1930 visit in Europe.

THE GENUS SCLEROTINIA

The generic name, Sclerotinia, should be restricted to those species in
which the apothecium arises from a tuberoid sclerotium as in S. sclerotiorum
(Lib.) De Bary, which is to be taken as the type of the genus. In this
genus the sclerotia originate and develop upon the surface of the substrate
or in lysigenous cavities within the organs of the suscept. They rarely and
only incidentally enclose in their pseudoparenchyma bits of the tissue of
the suscept. The sclerotia when freely formed upon the surface of the
substratum are approximately loaf-shaped or globose. Even those species
which form characteristically shaped sclerotia regularly within the organs
of their suscepts, when grown on artificial culture media produce sclerotia
which tend to be globose in shape. None of them are true parasites. They
must kill the cells of the invaded tissues before they are able to extract the
nourishment they seek. They are necrogenic saprophytes. They have no
conidial stage. All have spermatia (microconidia) produced endogenously
from the tips of obclavate spermatophores, fasciculately massed into muci-
laginous spermodochia, borne free on the aerial mycelium or enclosed in
lysigenous cavities in the tissues of the suscept.

The species here discussed are the only ones known to me on sedges and
rushes that may be properly referred to the genus Sclerotinia as above
characterized. A number of species of Discomycetes, recorded on various
species of these plants and referred to the genus Sclerotinia, do not qualify.
These in so far as they are known to me are listed and briefly discussed at
the end of this paper.

SPECIES TREATED IN THIS PAPER

As a stimulus to the reader to plunge into the details of my observations
and conclusions on the species treated in this paper, it may be well to
present here a list of these together with the sedges and rushes which they
inhabit. There are ten species occurring on forty-one different suscepts.
These species are largely restricted each to a distinct suscept or set of
suscepts. Only among the four species on Carex have there been found a
few cases of suscepts common to two or more.

On the Cyperaceae:

1. Sclerotinia duriaeana® (Tul.) Rehm in EUROPE only, on Carex
brizoides * L.; C. paniculata L.; C. disticha Huds.; C. chordorrhiza Ehrh.;

* Exercising his option under the rules, the author decapitalizes the first letter of
species names.

*The species names here used for the suscepts have been carefully checked by my
colleague Dr. R. T. Clausen.

WHETZEL: SPECIES OF SCLEROTINIA 387

C. acutiformis Ehrh. 2. Sclerotinia sulcata (Desm.) Whetzel in EUROPE
on Carex hudsonii A. Bennett; C. vulpina L.; C. gracilis Curtis; C. inflata
Huds.; C. brizoides L.; C. nigra (L.) Reichard; C. disticha Huds.; C.
paradoxa Willd.; C. paniculata L.; C. riparia Curtis, in NORTH AMER-
ICA on Carex stricta Lam.; C. prairea Dewey; C. interior Bailey; C. hys-
tricina Muhl.; C. riparia var. lacustris (Willd.) Kiiken; C. flava L.; C.
inflata var. utriculata (Boott) Druce; C. crinita Lam.; C. nebraskensis
Dewey. 3. Sclerotinia longisclerotialis Whetzel in NORTH AMERICA
only, on Carex prairea Dewey; C. interior Bailey; C. vesicaria L.; C. crinita
Lam.; C. retrorsa Schw.; C. oligosperma Michx. 4. Sclerotinia caricis-
ampullaceae Nyberg in EUROPE and NORTH AMERICA on Carex
aquatilis Wahl.; C. aquatilis var. alttior (Rydb.) Fern.; C. inflata Huds.
5. Sclerotinia vahliana Rostr. in EUROPE and NORTH AMERICA
on Eriophorum scheuchzeri Hoppe; E. angustifolium Roth.; E. vaginatum
L. 6. Sclerotinia scirpicola Rehm in EUROPE only, on Scirpus lacustris
L. 7. Selerotinia schoenicola n.sp. in EUROPE only, on Schoenus
nigricans L.

On the Juncaceae:

8. Sclerotinia curreyana (Berk. in Currey) Karst. in EUROPE only,
on Juncus effusus L.; J. conglomeratus L.; J. glaucus Ehrh.; J. filiformis
L.; J. communis E. Mey. 9. Sclerotinia juncigena (E. & E.) n.comb. in
NORTH AMERICA only, on Juncus effusus L. var. pacificus (Fern.)
Wieg. 10. Sclerotinia luzulae n.sp. in EUROPE only, on Luzula pilosa
Willd.

LIFE HISTORY

The life histories of the Sclerotinia species attacking sedges and rushes
are in their main features very much alike. The discharged ascospores,
carried by air currents, fall upon the flowers of their suscepts. Germination
and invasion of the flowers is followed promptly by death and browning of
the entire inflorescence, the affected culms never setting seed. The invading
hyphae now work rather slowly down the culm forming spermodochidia ®
(Text Figs. A, C, D) in the cortical tissues just beneath the epidermis.
The increasing mass of spermatia in these fruit bodies soon burst through
the epidermis in pearly white or brownish mucilaginous drops. Rain or
dew and possibly insects disseminate the spermatia to the surfaces of the
dying culms below wherein the sclerotia are now beginning to develop. The
mechanism of fertilization is unknown but that the spermatia do function
is certain. The sclerotia, at first evident as white soft cottony masses within
the culm, gradually harden, enveloping the medulla in a thin black rind.
This medulla is often pink at first, eventually becoming white. In the final
stages of maturity in late autumn, the sclerotia usually burst the confining
outer tissues of the culm. In some species they fall out of the culms onto
the water or moss beneath. In other species they remain partially enclosed,

For a definition of this term see Whetzel, Mycologia 35: 335-338. 1943.

388 FarLowlA, VoL. 2, 1946

exposing only a part of their surface. In one species, S. longisclerotialis
(Fig. 20), the sclerotia remain enclosed until decay of the culm tissues more
or less frees them the following spring. The natural shape of the sclerotium
is usually characteristic for each species, but is evidently determined to
some extent by the shape of the enveloping culm. The number of sclerotia
per culm varies not only with the species of Sclerotinia but to some extent,
and within limits, in most of the species of their respective suscepts.

The sclerotia of those species in which they are freed at maturity, float
on the surface of the water, eventually finding lodgment among the mass of
debris about the hummocks or shores of the swamp. In some species the
sclerotia remain upright in the dead culms throughout the winter. In S.
longisclerotialis the dead slender culms enclosing the sclerotia fall over and
settle to the bottom of the pools about the hummocks.

Development and maturity of the apothecia of all the species are closely
correlated with the growth and flowering of their respective suscepts early
in the spring. The ascospores are matured and discharged over a period of
two weeks or so while the flowers of their suscepts are opening and shedding
their pollen. Thus for the species of Sclerotinia attacking sedges and rushes
there is a single primary cycle of activities. So far as known, secondary
cycles do not occur. Each species presents certain peculiarities and varia-
tions from the life-history picture outlined above. These will be pointed
out in connection with the discussion of the different species. This outline
of their life history is inferred largely from observations on their seasonal
developments and the syndrome of the diseases they cause. Most of them
have been grown on potato dextrose agar. Much remains to be discovered
by further field studies and especially by carefully executed inoculation
experiments. Several of them, no doubt, have a more extensive suscept
range than my observations indicate. They appear to have been very in-
frequently collected, not only in North America but also in Europe. Con-
sidering their abundance and annual reappearance in the localities where
I have had them under observation, it is surprising how scanty in numbers
and quantities are the specimens in the herbaria which I have visited.

SPECIES ON THE CYPERACEAE

Of the seven species of Sclerotinia known to me to attack members of the
Cyperaceae, four occur on species of Carex, one on Scirpus, one on Erio-
phorum, and one on Schoenus. These appear to constitute a group of very
closely related species. All occur in much the same type of environment,
namely in wet marshes where water stands throughout the year or along
the margins of lakes, ponds, and quiet streams. They appear to be restricted
to the cooler parts of the north temperate zone and on up into the colder
arctic regions. They usually occur in great abundance in the localities
where I have found them both in America and Europe.

WHETZEL: SPECIES OF SCLEROTINIA 389

a

SPECIES ON CAREX

Four species of Sclerotinia pathogenic to Carex species have been de-
scribed. Of these S. duriaeana (Tul.) Rehm is known only from north-
western Europe, occurring widely on several species of Carex from France
northward to Lapland in Scandinavia; S. suicata (Desm.) Whetzel occurs
in both Europe and America. It appears to be generally distributed over
western Europe from south to north wherever its-Cavex suscepts occur. It
is this species which has been most frequently collected and preserved in
European herbaria but almost always erroneously labeled Sclerotinia du-
riaeana, with which it has generally been confused by European mycologists.
While S. longisclerotialis Whetzel infects many species of Carex in north-
eastern United States it apparently does not occur in Europe. The fourth
species, S. caricis-ampullaceae Nyberg, discovered in Finland in 1930 and
first described two years later, has been collected since in Sweden as well
as in North America in the high mountain swamps of Wyoming (Whetzel

«

_ and Solheim 1943). This is the largest and most striking of the four species

(Fig. 25). The apothecia of these four species differ chiefly in size and
somewhat in shape. The differences in color of the apothecia and in the
shape and size of asci and ascospores are scarcely of taxonomic significance.
The species are most clearly distinguishable by the shape and size of their
sclerotia together with the external appearance and arrangement of their
spermodochidia.

KEY TO THE SPECIES

1. Sclerotia short, more or less fusiform, falcate or kidney-shaped.
2. Spermodochidia in globose, equally spaced groups..... S. duriaeana
2. Spermodochidia in linear, scattered groups........... S. sulcata

1. Sclerotia long and slender.
3. Sclerotia of uniform diameter, truncate, more or less 3-angled
S. longisclerotialis
3. Sclerotia obclavate, drawn out above into a slender whiplike tip
S. caricis-ampullaceae
SCLEROTINIA DURIAEANA
Prate I & II

A clear and accurate concept of this species has long been beclouded by
errors in identification of certain stages in its development and by its fre-
quent confusion with S. suicata (Desm.) Whet. by mycologists subsequent
to its description by Tulasne in 1861. I have already presented (Whetzel
1929) in some detail the basic evidence for this confusion of the two species.
Critical examination of specimens preserved in European herbaria together
with observations in the field, in culture, and on preserved specimens of
both these species personally collected on numerous species of Carex in
Europe during 1930, has fully confirmed my earlier provisional conclusion
(Whetzel 1929: 15) that the two species have commonly been confused
under the name Sclerotinia duriaeana.

While I made numerous collections of the spermatial and sclerotial stages
of this species in most of the countries visited, I was in most places too late

FARLowIA, VOL. 2, 1946

390

PLATE I

WHETZEL: SPECIES OF SCLEROTINIA 39]

to find apothecia. Unfortunately I was unable to visit the type locality
where Durieu made his collections and observations on this species. I did,
however, make two collections of apothecia which appear to be those of
S. duriaeana, one near Rambouillet, France, (CUPP 31561) and another
near Lyngby, Denmark, (CUPP 31564) (see data and comments below
p. 395). In neither case were many apothecia found and the circum-
stances under which they were taken raises some question as to their
identity. Perhaps if I had found abundant fresh cups of both species, each
of unquestionable identity, I might have detected some differences of
specific significance in their apothecia. I doubt it. Such evidence as is
available offers little expectation that they are thus distinguishable. The
sclerotia of the two species cannot with confidence be told apart and the
spermatial stages alone provide ready and certain identification. (Compare
Figs. 1 and 13.) A more extensive comparison than I have been able to
make of these two species in artificial pure culture would almost certainly
also provide dependable evidence for distinguishing between them.

In view of the scattered, confused and incomplete published data and
descriptions of S. duriaeana, it appears desirable to present here a more
complete and accurate description of the species, interpreted in the light
of my own studies and observations.

SCLEROTINIA DURIAEANA (Tul.) Rehm, Hedwigia 21: 66. 1882.

Epidocium ambiens Desm. Ann. Sci. Nat. Ser. 3, 20: 231. 1853.

Peziza duriaeana Tul. Sel. Fung. Carp. 1: 103. 1861, and 3: 203, pl. 22, fig. 20-24.
1865.

Sphacelia nigricans sensu Sacc. Michelia 2: 131. 1880.

Sclerotium nigricans sensu Sacc. Michelia 2: 134. 1880.

Sphacelia ambiens Sacc. Syll. Fung. 4: 666. 1886.

Sclerotinia duriaeana Sacc. Syll. Fung. 8: 199. 1889.

Sclerotinia duriaeana (Tul.) Quélet (In error. See Notes p. 395.)

Hymenoscypha duriaeana Phill. Manual Brit. Discom. p. 115. 1893.

Myrioconium ambiens V. Hohnel. Mitt. Bot. Inst. Tech. Hochsch. Wien. 3: 50.
1926.

Spermodochidia (Figs. 1, 3 & 7) dull black in paired groups of 3-6 at
regular intérvals of 5-15 mm. along upper part of culm, 5-15 paired groups
per culm, groups oblong to subglobose when moist, 1-3 mm. long, arranged
parallel side by side lengthwise of the culm, usually on but two sides of the
3-angled culm, sometimes encircling the culm as in C. chordorrhiza (Fig. 2);
at maturity becoming swollen and rupturing the culm, discharging the
viscid, olivaceous mass of spermatia; spermatia minute, globose, 2—3 y in
diam., faintly olivaceous.

The spermodochidia are formed in lysigenous cavities in the palisade
parenchyma of the culm, between the vascular bundles. Each spermo-

Plate 1. Sclerotinia duriaeana. Fig. 1. (Left) diseased heads of Carex paniculata L.;
(right) healthy head, with spermodochidia and sclerotia in the diseased culms. CUPP
31562. Fig. 2. Diseased culms of C. chordorrhiza Ehrh. showing the spermodochidia.
CUPP 31563. Fig. 3. Culm of C. paniculata with spermodochidia and sclerotium from
De Thiimen’s Myc. Univ. 1557. All figures natural size.

392 FARLOWIA, VoL. 2, 1946

, bh to-g™ (455
PE Vacs ecoubee Hei S anni 406 49k 4

Tq

PuaTeE II

WHETZEL: SPECIES OF SCLEROTINIA 393

dochidium contains many spermodochia surrounded by masses of spermatia.
(See Tulasne 1865: pi. 22, fig. 21.)

Sclerotia (Fig. 4) one to several in each culm, from which they are dis-
charged at maturity through a slit in the culm over the sclerotium by the
toppling of the upper part of the culm which bending at this point opens
the sclerotial cavity; black, slender, fusiform, more or less curved when
dry, when fresh plump, fusoid, slightly 3-angled, varying in length and
thickness according to the size of the culm in which formed, 5-15 mm.
long. The sclerotia first appear within the culm as elongate, pink, cottony
masses of hyphae, which gradually darken externally as the dull black rind
is formed and the densely’packed thick-walled hyphae of the white medulla
takes form. '

Apothecia variable in size, depending upon the size of the sclerotia from
which they arise, usually but one from each sclerotium; cup, 5-10 mm. in
diam. occasionally larger, shrinking greatly on drying, deep goblet-shaped,
thin, fragile, light fawn to vinaceous brown, lighter beneath due to a fine
hyphal fuzz which covers the under surface when fresh; stipe rather short,
its length usually about the diameter of the fully open cup, nearly or quite
smooth; asci, according to Tulasne (1861: 106) are ‘“‘8-spored (spores
obliquely and closely uniseriate, seldom biseriate), cylindrical, obtuse,
straight, 8—9.5 » wide and about ten times as long.” Boudier (19.10: 274)
describes them as cylindrical, rather large, slightly attenuate toward the
base, 8-spored, pore blue with iodine, 210-230 x 7-10 yp; ascospores are
described by Tulasne (/.c.) as “ovate, inequilateral, obtuse at both ends
and muticous, smooth, unilocular, and filled with oil the greater part
measuring 10-15 x 6.5—7.5 yp, but some not exceeding 8—9:5 x 5 ».”” Boudier
describes them as oblong-fusiform, colorless, usually without internal gran-
ules, but sometimes cloudy or rarely with non-oily droplets, 14-18 x 6—7 up.
Tulasne (1865: pl. 22, fig. 24) describes the discharge of the ascospores
and their germination which he illustrates showing the direct formation on .
them of spermatophores and spermatia; paraphyses slender, slightly thick-
ened at the tips, hyaline, sparingly septate, vacuolate (Boudier 1907: pi. .
473, fig. 9; 1910: 274).

Hasirat: In culms of Carex species, growing in open swamps, marshes
and wet meadows in western Europe.

_ TYPE SPECIMENS: Collected by Durieu de Maisonneuve in culms of
Carex brizoides L. during the years 1855 to 1861 in the sandy meadows
along the river Ciron about the ruins of the Chateau de Fargues not far
from the towns of Langon and Bazas, France. There appears to be no

Plate 2. Sclerotinia duriaeana. Fig. 4. Photograph of Durieu’s type specimens of
the apothecium and sclerotia deposited in the Botanical Museum, Paris, sent me by
Roger Heim; natural size. Tulasne (Sel. Fung. Carp. 3: pl. 22, fig. 23) presents a good
sketch of the apothecium of this species. Fig. 5. Mycelial growth from bits of diseased
_ culms of C. paniculata on potato dextrose agar; 14 natural size. CUPP 31562. Fig. 6.
Mycelium and spermodochidial masses from tissue plantings from culms of C. chordor-
rhiza on potato dextrose agar; 4% natural size. CUPP 31563. Fig. 7. Spermodochidia
on culm of C. chordorrhiza, x6. CUPP 31563.

394 Fartow!A, Vow. 2, 1946

evidence that Tulasne preserved specimens of the materials on which his
description of the species was based. The specimens he had in hand appear
to have all been sent him by Durieu. Durieu, however, preserved what are
obviously duplicate specimens of those sent Tulasne. These specimens were
later, 1878, deposited in the cryptogamic herbarium of the Museum of
Natural History in Paris (Fig. 4). These specimens therefore may be
properly considered the type specimens of this species. These, together
with rather extensive notes in Durieu’s handwriting, were in 1930 mounted
on herbarium sheets. Photographic copies of three of these sheets were
kindly sent me by the curator, Dr. Roger Heim, after my visit to the mu-
seum. These are on file with my notes on this species. Duplicate specimens
of sclerotia and infected inflorescences bearing spermodochidia from Du-
rieu’s collection of June 24 and 26, 1860, are filed in CUPP 17044. I found
among Durieu’s specimens but one apothecium (Fig. 4), apparently one of
those developed from his first collection of sclerotia, July 1, 1855.

Since no species of Carex other than C. brizoides was found in Durieu’s
collections, and since this is the species pictured by Tulasne, it would ap-
pear that neither of them had in hand specimens of S. sulcata which they
might have confused with S. duriaeana. Tulasne’s (1861: 103) application
of the name Sclerotium sulcatum Roberge in Desmaz. to the sclerotial stage
of S, duriaeana was at the time a pardonable error. The only other name
of the sclerotial stage of this species which has been applied is Sclerotium
nigricans Sacc. given it by Saccardo (1880: 134, and 1899: 1153) under
the erroneous impression, however, that the specimens in hand were sclerotia
of Claviceps nigricans Tul.

DistRiBUTION: This species is known to occur only in Europe. It has
not been found in North America. Apothecia apparently appearing in May,
spermodochidia June to July, sclerotia July and August. It is known to me
from France, Belgium, Germany, England, Denmark and Norway.

ILLustTRATIONS: Tulasne, Sel. Fung. Carp. 3, pl. 22, fig. 20-24. 1865.
Boudier, Icon. Myc. 3: pl. 473. 1907.

MarteriaL Examinep: On Carex brizoides L. (C. arenaria in error): (1) Durieu’s
type specimens deposited in the Crypt. Herb. Mus. Nat. Hist., Paris (fully discussed
above). I also found what appear to be duplicate specimens (spermodochidia and
sclerotia) from Durieu’s collections in the Kew Herbarium labeled “Peziza Duriaeana
on Carex arenaria” from Cooke’s herbarium; — (2) Krieger, Fung. Sax. 1682, “am
grossen Winterberg,” Germany, August 1, 1898 (spermodochidia and sclerotia). On
Carex paniculata L.: Desmaziéres, Pl. Crypt. France. Ser. I, 20, “Epidocium ambiens
Desm. Not. XXI. En été, sur les tiges séches de divers Carex, 4 Ouystreham par M.
Roberge.” France (spermodochidia only) ;—De Thiimen, Mycotheca Univ. 1575,
“Epidocium ambiens Desm. f. Caricis paniculatae. Gallia. Grande Chartreuse pr. Gre-
noble. Aug. 1879, J. Therry.” (spermodochidia and sclerotia); Roumeguére, Fungi
Sel. 4682 “Sphacelia ambiens (Desm.) Sacc.” (spermodochidia only) ; his Fungi Gallici
1200, “Sclerotium nigricans (Tul.) Sacc.” (spermodochidia) all appear to have been
made up from one and the same original collection by Therry; — Roumeguére, Fungi
Gallici 3505, “Epidocium affine Desm. 3N.21. Chaumes du Carex paniculata. Tremonde
(Belgique). Reliq. Westendorp.” This is incorrectly determined. It is typical Epidocium
ambiens Desm. The spermodochidia (no sclerotia) are typical of those of S. duriaeana.
No date of collection is given; — Roumeguére, Fungi Sel. 4600, “Sclerotium sulcatum

WHETZEL: SPECIES OF SCLEROTINIA | - 395

Rob. A l’interieur des chaumes secs du Carex paniculata. Louette Saint Pierre (Bel-
gique). legit Aubert.” (sclerotia only). No date given; —CUPP 31561, swamp near
Rambouillet, France, May 6, 1930, Whetzel, Westcott & Duffrenoy. Several apothecia
found lodged on hummocks of Carex paniculata standing in a pool.of water. It is barely
» possible that some of these apothecia are these of S. sulcata. Ascospores discharged
from two of them (Figs. 5, 6) gave, however, cultures quite different from cultures of
S. sulcata (Figs. 17, 18) and like those from infected culms of the following collection;
— CUPP 31562, swamp in outskirts of Faaborg, Isle of Fyn., Denmark, June 27, 1930,
Whetzel & E. Gram (spermodochidia and sclerotia). On Carex disticha Huds.:
Roumeguére, Fungi Selecti. Exs. 5419, “Sclerotinia Duriaeana Tul. (conidies). Sur les
chaumes maladifs d’un Carex sp. King’s Lynn (Angleterre), Automne 1889.” Coll. Plow-
right (sclerotia and spermodochidia). A. J. Wilmott of the British Museum says the
Carex is C. disticha Huds.; — Phillips Herbarium, Brit. Mus., three collections on one
sheet. One is labeled ‘“Peziza Duriaeana Tul. King’s Lynn. July 1888,” presumably
collected by Plowright, showing typical spermodochidia. A. J. Wilmott identified the
suscept as Carex disticha Huds. Two others are sclerotia which bear apothecia, labeled
respectively: “Peziza Duriaeana Tul., growing from sclerotia gathered June 3, 1883 by
C. B. Plowright, King’s Lynn”; and “Peziza Duriaeana Tul. cultivated from sclerotia
on Carex arenaria by C. B. Plowright, June 1, 1883.” Dupl. in CUPP 33271. On Carex
chordorrhiza Ehrh.: Flora Suecica, 1636, under the name Myrioconium ambiens
(Desm.), the spermatial stage only. Collected by J. A. Nannfeldt in Torne Lappmark,
Jukk&sjarvi, at Abisko above the railway station August 9, 1928. Dupl. in CUPP 31620.
Apparently the first collection on this suscept; — CUPP 31563, marshy bogs near hotel
at Fjellsiter above Trondhjem, Norway. July 8, 1930, Whetzel & Jgrstad (spermodo-
chidia and sclerotia). On Carex acutiformis Ehrh. (?) (new suscept): CUPP 31564,
along the shore of Lyngby S6, Lyngby, Denmark. June 5 and June 17, 1930, H. H.
Whetzel (apothecia and sclerotia). The determination of the suscept by Grgntved of
the University of Copenhagen is in some doubt as the single culm enclosing sclerotia is
old and rotten.

In making the transfer of this species to the genus Sclerotinia, Rehm had
before him (his Ascomyceten 603) specimens of S. sudcata which he mis-
takenly took to be those of Tulasne’s species Peziza duriaeana (Whetzel
1929: 7).

The combination, Sclerotinia duriaeana (Tul.) Quél. appears to be a
strange error, originally made by Rehm in his ‘‘Discomyceten” in Rabenh.
Krypt. Flora 1: 820. 1893. He here lists this species as “Scl. Duriaeana
(Tul)” giving as a synonym, “Sclerotinia Duriaeana Quel. Bull. soc. myc.
I. pag. 115”. An examination of this reference discloses that the paper is
by Boudier (not Quélet) in which he merely lists “P[eziza] Duriaeana
Tul.” as an example of species to be included in Sclerotinia Fckl. which
Boudier makes a sub-genus of Ciboria Fckl. Later Boudier in his Icones
Myc. 4: 274 writes “Sclerotinia Duriaeana (Tul.) Quél.” without citing the
place where Quélet is assumed to have made the transfer. Succeeding au-
thors appear to have perpetuated the error.

Massee (1895: 283) cites as a synonym of Sclerotinia duriaeana, “Epi-
docium arabicus Desmaz. xxii. Not. in Ann. Sci. Nat., p. 19.” This species
name appears to be a strange error in the spelling of the name ‘“‘ambiens.”
The reference he gives is obviously that of the original place of publication
of Epidocium ambiens, the page being apparently that of a reprint instead
of the original.

The names Sphacelia nigricans and Sclerotium nigricans were erected by

396 FarLowI1a, VoL. 2, 1946

Saccardo who erroneously believed the sclerotia and spermodochidia in
specimens (of S. duriaeana) on Carex paniculata were the sclerotia and
spermoidea of Claviceps nigricans Tul. (Whetzel 1929: 9-10 and 1944:
426-428).

Von Hohnel (1926a: 50-51) erroneously lists Epidocium affine Desm. as
synonymous with his Myrioconium ambiens, believing it to be the same as
or a variety of Epidocium ambiens of Desmazieéres.

There is considerable variation in the measurements of asci and asco-
spores of S. duriaeana as recorded by different authors. I have made meas-
urements of asci and ascospores from three collections of apothecia with
the following results: (1) Plowright’s collection on C. paniculata: asci (6
meas.) 121.2—151.5 x 6—9.5 ; spores (50 meas.) 9.9-18.4 x 4.6-6.6 4, mode
12.5x5.3 pw, av. 12.7 x 5.4 p. (2) Whetzel’s collection on C. paniculata —
(CUPP, 31561): asci (30 meas.) 133.3-181.8 x 8.1-10.8., mode 157.5 x
8.1 pw, av. 158.9x8.3 w; spores (50 meas.) 10.5-14.5 x 4.6-7.2 «, mode
11.8x5.3 p, av. 11.9x5.4 w. (3) Whetzel’s collection on C. acutiformis
(CUPP, 31564): asci (40 meas.) 127.2-169.6 x 8.1-9.5 uw, mode 151.5 x 8.1
pf, av. 147.6 x 8.7 w; spores (80 meas.) 9.9-15.8 x 5.3-7.9 », mode 13.2 x 6.6
#, av. 13.3x 6.5 w. These measurements are distinctly smaller than those
given by Boudier, but compare favorably with those given by Tulasne.

Although I cite Boudier’s description and illustrations which he presents
under the name S. duriaeana, I do so with some reservations. He does not
specify the species of Carex on which his specimens occurred. His illustra-
tions might just as well be those of S. su/cata. His measurements for asci
and ascospores are rather large in comparison with the measurements of
asci and spores of S. duriaeana. They more nearly approximate the maxi-
mum measurements of these structures in S. sudcata.

Whoever was responsible (presumably Durieu) for the identification of
the Carex harboring the Peziza duriaeana which Tulasne had in hand, ap-
pears to have been in error. Culms from these collections bearing the typi-
cal spermodochidia and sclerotia as pictured by Tulasne (1865: pl. 22, fig.
20) have been critically examined by competent European taxonomists and
declared to be Carex brizoides L., not C. arenaria L.

It is to be noted that of the Carex species on which S, duriaeana is known
to occur only two are reported from North America. This may account for
its apparent absence from the Western Hemisphere. It would appear to be
most common and widely occurring on Carex paniculata and may yet be
found on this suscept where that occurs in the Western Hemisphere.

SCLEROTINIA SULCATA
Prates III anp IV anp Text Fic. A

This species is apparently the most common and widely distributed of
the Sclerotinia species on Carex. It occurs throughout northwestern Europe,
northeastern United States and in the Pacific Northwest. Long confused
with S. duriaeana, its distinct identity was first suggested by the writer
(Whetzel 1929: 15). In this paper I reviewed in some detail the evidence

WHETZEL: SPECIES OF SCLEROTINIA 397

then available to me. My study of the species in Europe in 1930 has fully
confirmed that conclusion. In that paper (/.c.: 19-24) I described it under
the name Sclerotinia duriaeana (‘affine form’), but anticipating the cor-
rectness of my belief that it was a species distinct from S. duriaeana I made
the combination S. sudcata (l.c.: 15). It is here again briefly characterized
under its proper name. |

SCLEROTINIA SULCATA (Desm.) Whetzel, Mycologia 21: 15. 1929.

Sclerotium sulcatum Roberge in Herb. Desm. (in part) Ann. Sci. Nat. Ser. 3, 16:
329. 1851.

Epidocium affine Desm. (in part) Ann. Sci. Nat. Ser. 3, 20: 232. 1853.

Claviceps (?) caricina Griffiths, Bull. Torrey Club 29: 300. 1902.

Sclerotinia duriaeana “ambiens form,” Whetzel, Mycologia 21: 15. 1929.

Spermodochidia (Figs. 8-14) scattered irregularly along the upper part
of the affected culm, usually confined to that part above the first sclerotium,
sometimes a few below between sclerotia; almost always confined to two
faces of the 3-angled culm; distinctly linear, 2-5 mm. long by % to 2 mm.
broad; dull olivaceous brown, inconspicuous especially in dry culms, split-
ting longitudinally when mature to release the spermatial ooze. Their struc-
ture is illustrated in Text Fig. A; spermatia globose about 2 » in diam.,
hyaline, produced endogenously and in succession from the tips of densely
clustered obclavate spermatophores, embedded in a mucilaginous matrix.

Sclerotia (Fig. 16) one to three in each culm, fusiform, ends blunt,
rounded, sulcate, at first pink, then smoky gray and finally black, variable
in size, depending on the slenderness of the culm in which formed, 20-25
mm. by 3—4 mm. in the large culms of C. riparia var. lacustris, 4-5 mm. by
144-2 mm. in C. interior. (For a more detailed description see Whetzel
1929: 20.) |

Apothecia (Fig. 15) usually but one from each sclerotium, short-stipitate,
fawn-colored; cups deep goblet-shaped to shallow funnel-shaped when fully
expanded, varying in size from 2 to 10 mm. broad, about half as deep,
smooth without; membranous; stipe relatively short, 5-20 mm. long,
smooth and concolorous with cup above, darker below; ascz slender, clavate,
apex rounded, pore J+, 8-spored, spores uniseriate, 106 meas. of asci from
6 apothecia from C. stricta 128-183 x 7.3.x 11 pw, mode 155.8x 9.16 p, av.
161 x 8.6 »; ascospores hyaline, ovoid, more or less inequilateral, 205 meas.
from 5 apothecia from C. stricta 8.8—-17.5 x 5.3-8.8 p, mode 13.1 x 7.09 gp,
av. 12.6x 6.9 »; paraphyses slender clavate, hyaline.

Hasitat: In culms of Carex species in swamps, wet, open marshes and
along banks of slow streams and ponds. Europe and North America.

LECTOTYPE SPECIMEN: Cornell University Pl. Path. Herb. 11515 on
Carex stricta Lam., McLean, N. Y., swamp just south of Lloyd Preserve.
April 26, 1921, Whetzel. Duplicate deposited in Kew Herbarium, Kew,
England.

DISTRIBUTION: Known to me in North America only from the region
about Ithaca, N. Y., Delaware and the Pacific Northwest (Oregon). It will
almost certainly be found when sought throughout northern United States

OL. 2, 1946

, Vv

TA

ARLOW

=
4

398

PLate III

WHETZEL: SPECIES OF SCLEROTINIA 399

and Canada. It is apparently common throughout northwestern Europe as
the records (below) of European specimens examined would indicate. The
apothecia appear early in the spring (April-May) at the time the Carex
suscepts are beginning to flower. The spermodochidia and sclerotia begin
to appear about a month later (June-July) and may be readily spotted in
the dead bleached culms standing among the healthy.
ILLUSTRATIONS: Whetzel, Mycologia 21: fig. 1-7, 14-19, 1929.

Materia EXAMINED: In connection with my original description of this species
(1929) I recorded the North American species of Carex then known to me, on which
S. sulcata occurs. Since that time no additional collections of this species of special
interest from North America have come into my hands. To round out the picture
I present here the records of European collections which I have examined and studied:
On Carex Hudsonii Bennett. (=C. stricta Good.): Desmaziéres, Pl]. Crypt. France,
Ser. II, 212. ‘“Epidocium affine” (spermodochidia only) ; — Lindhart, Fungi hungarici,
381, “Peziza Duriaeana Tul.” on Carex, Hansag, Nov. 1883 (sclerotia and spermodo-
chidia) ; — Rehm, Ascomyceten, 603 a, ‘Sclerotinia Duriaeana (Tul.)” on “Carex stricta
Good.”, Diessenhofen, Thurgau, Switzerland, Apr. 1881, “Dr. Winter’ (sclerotia and
apothecia) ; — Rehm, Ascomyceten, 603 b, “Sclerotinia Duriaeana (Tul.) Walesellen bei
Ziirich, Auf faulenden Halmen, Carex stricta, 4/1893. Dr. V. Tavel” (apothecia and
sclerotia) ; — Rabenhorst-Winter, Fungi Europaei, 2749. “Sclerotinia Duriaeana (Tul.)”
on Carex stricta Good., Diessenhofen, Switzerland, April 1882, H. Wegelin (apothecia
and sclerotia);— CUPP 31566, swamp near Fuer S6, Jagerhuset, Denmark, June 8, -
1930, Whetzel (spermodochidia and young sclerotia) ; —CUPP 31567, along shore of
Sjael S6, Denmark, June 20, 1930, Whetzel & Ferdinandsen (spermodochidia) ; —
CUPP 31568, swamp on outskirts of Copenhagen, near Soburg, June 21, 1930. Whetzel
& Groéntved (spermodochidia) ;— CUPP 31569, Ryggmossen near Upsala, Sweden,
May 28, 1931, K. G. Ridehies (apothecia and sclerotia). Dupl. in Herb. Univ. Upsala,
Sweden. On Carex vulpina L.: Desmaziéres, Pl. Crypt. France, Ser. I, Ed. 1, 2029
and Ser. I, Ed. 2, 1629. “Sclerotium sulcatum Rob. in herb.” (spermodochidia and
sclerotia) ; — CUPP 31576, Upper Grimley, brick pits, Worcestershire, England, August
8, 1930, Whetzel & C. Rea (spermodochidia). On Carex gracilis Curtis: CUPP 31570,
Upton-on-Severn, Worcestershire, England, August 8, 1930, Whetzel & C. Rea (spermo-
dochidia and sclerotia) ;—- CUPP 31571, Upper Grimley, brick pits, Worcestershire,
England, August 8, 1930, Whetzel & C. Rea (spermodochidia and sclerotia). On Carex
inflata Huds. (=C. rostrata Stokes): CUPP 31572, swamp in woods at Nosten, parish
of Bondhyrho near Upsala, Sweden, July 3, 1930, Whetzel & Nannfeldit (spermodo-
chidia). On Carex brizoides L.: CUPP 31573, swamp in woods at Nosten, parish of
Bondhyrho near Upsala, Sweden, July 3, 1930, Whetzel & Nannfeldt (spermodochidia).
On Carex nigra (L.) Reichard (=C. Goodenowii Gay) : CUPP 31574, swamp in woods
at Nosten, parish of Bondhyrho near Upsala, Sweden, July 3, 1930, Whetzel & Nann-
feldt (spermodochidia). On Carex disticha Huds.: CUPP 312575, along shore of
Lyngby Sé, Lyngby, Denmark, June 21, 1930, Whetzel (spermodochidia). On Carex
paradoxa Willd.: CUPP 31578, swamp near Fuer S6, Jagerhuset, Denmark, June 24,
1930, Whetzel & Buchwald (spermodochidia). On Carex paniculata L.: CUPP 31579,
marsh near Faaborg, Isle of Fyn, Denmark, June 29, 1930, Whetzel & E. Gram (sper-
modochidia). On Carex riparia Curtis: CUPP 317580, Knapp’s brickyard, Claines,
Worcestershire, England, August 7, 1930, Whetzel & C. Rea (spermodochidia and

Plate 3. Sclerotinia sulcata. Figs. 8-10. Diseased culms of C. gracilis showing dis-
eased inflorescence, spermodochidia, and sclerotia; natural size. CUPP 31571. Fig. 11.
Spermodochidia on culms of C. vulpina, x6. CUPP 31576. Fig. 12. Spermodochidia on
culms of C. paniculata, x6. CUPP 31579. Fig. 13. Diseased inflorescence, spermo-
dochidia and sclerotia on culm of C. prairea; natural size. CUPP 14947. Fig. 14.
Spermodochidia on culms of C. gracilis x6. CUPP 31571.

400 FarRLow1A, Vou. 2, 1946

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Text Fig. A. Cross section through a spermodochidium of Sclerotinia sulcata in culm
of C. paniculata about x325. Note that the cavity is formed by lysis of the paren-
chyma between the vascular bundles. Drawn by Carlos Garcés.

WHETZEL: SPECIES OF SCLEROTINIA 401

sclerotia) ; — CUPP 31581, valley of the Avon, south of Salisbury near Woodgreen,
Hampshire, England, August 4, 1930, Whetzel & Wm. Brown (spermodochidia. and
sclerotia). On Carex sp.: Herbarium of-C. Crossland (Herb. Kew, England), “Sclero-
tinia Duriaeana (Tul.) Quel. Among decaying Carex etc. Masham, (Yorkshire), Eng-
land. May 28, 1902” (sclerotia and apothecia).

The earliest name applied to any stage of this species is Sclerotium sul-
catum Roberge in herb. Desm. (1851). Two years later Desmaziéres de- .
scribed the spermodochidial stage under the name Epidocium affine. Both
of these names were based on mixtures of culms of Carex species and Schoe-
nus nigricans, the sclerotia and spermodochidia on the latter, being those of
a heretofore undescribed species of Sclerotinia. (See p. 421). These stages
as well as the apothecia of Sclerotinia sulcata were long confused by Eu-
ropean mycologists with those of S. duriaeana. The writer (Whetzel 1929:
15) appears to have been the first to suspect its specific identity.

The only recorded collection of this species in North America, prior to
1929, as far as I have been able to discover, was made by Griffiths and
Morris in August 1901 near Andrews, Oregon on Carex nebraskensis. Their
collection consists of sclerotia and spermodochidia only. Griffiths (1902)
described the fungus under the name “Claviceps? caricina sp. nov.” Groh
(1911) having had occasion to examine this collection by Griffiths and
Morris which had been distributed as West American Fungi 400, referred
it to Giissow who identified the fungus as Sclerotium sulcatum but errone-
ously referred the spermodochidia to Epidocium ambiens, the spermatial
stage of Sclerotinia duriaeana.

It would appear from the recorded collections that S. sudcata is more
common in Europe than S. duriaeana and goes to a greater number of Carex
species. Both species are recorded on C. brizoides, C. paniculata and C.
disticha. While Carex paniculata and its closely related species, C. brizoides,
C. disticha and C. chordorrhiza appear to be the favorite suscepts of
S. duriaeana in Europe, C. Audsonii and its North American counterpart,
‘C. stricta, appear to be the most common suscepts for S. sudcata in North
America.

SCLEROTINIA LONGISCLEROTIALIS
PLATE V

As far as I am aware, Sclerotinia longisclerotialis Whet. has been recorded
only from North America. I kept a sharp lookout for it wherever I col-
lected in swamps and wet marshes in western Europe during 1930, but no
specimens were found which might on any grounds be referred to this
species. No Sclerotinia species was taken there on any of the Carex species
on which S, longisclerotialis is known to occur. It seems therefore probable
that it is a species restricted to North America.

Although this species is described, illustrated, and discussed in full in my
1929 paper (Whetzel 1929: 24-30, fig. 10-13, 20-23) a brief description
- is here presented for the convenience of the reader. A complete list of the
species on which it is known to occur, including a new suscept made sub-
sequent to those already listed (Whetzel 1929: 30), is recorded below.

402 FarLowlA, VoL. 2, 1946

PLaTE IV

WHETZEL: SPECIES OF SCLEROTINIA 403

SCLEROTINIA LONGISCLEROTIALIS Whetzel, Mycologia 21: 24-30. 1929.

Spermodochidial groups (Figs. 20, 21) ovate, oblong, shining black, in
pairs at rather regular intervals along two faces of the culm just below the
inflorescence, rarely toward the base below the sclerotium, formed within
the cortical tissues of the culm, exposed by slits in the epidermis through
which ooze the mucilaginous masses of spermatia; spermatia globose 1-2 p
in diam., hyaline, produced in succession from the tips of obclavate sperma-
tophores. External appearance and internal structure essentially as in
S,. duriaeana (see Whetzel 1929: fig. 23).

Sclerotia (Figs. 19, 22) usually one, sometimes two or more, in culm,
remaining enclosed or slightly exposed by a slit in the epidermis (Fig. 20),
never freed as are the sclerotia in S. sulcata or S. duriaeana, at maturity
truncate, uniform in thickness, more or less 3-angled, striate due to pressure
of the vascular bundles of the culm during development, at first cottony,
becoming pinkish, then smoke gray, finally dull black without, the medulla
white; variable in length and diameter according to the size of the infected
culm, 1 cm. long by 1 mm. thick in C. interior, 7 cm. long by 2 mm. thick
in C. retrorsa and 3 to 10 cm. long in C. prairea.

Apothecia (Fig. 19) thistle-, funnel-, or goblet-shaped with long Sendet
stipe, one, rarely two or more, from each sclerotium; cup dark fawn-colored,
smooth without, enone! mouth courier never opening midely,
2-5 mm. broad; stipe slender, 10-50 mm. long, pale and smooth above,
dark tomentose below; asci long, cylindrical, attenuate below the middle,
136 meas. from 5 apothecia from C. prairea gave 170-230 x 8.8-14.3 pn,
mode 194x 10.4 p, av. 182.1 x 10.7 yw; ascospores strongly inequilateral,
flat or incurved on one side, 289 meas. from 5 apothecia from C. prairea
gave 12.6-22.8 x 5.3-12.5 pw, av. 18.3x 8.1 », mode 21x9 pw; paraphyses
slender, slightly clavate, septate.

In culture on potato dextrose agar forming an appressed feathery growth
of aerial mycelium (Whetzel 1929: fig. 20) with distinct minute hard
spermodochial masses. No sclerotia formed.

HasitaT: Open swamps and marshes where water stands throughout
the year.

TYPE LOCALITY: Swamp south of the Lloyd Preserve, McLean, N. Y.

DisTRIBUTION: U.S. A.: New York and Maine; Canada: Lake Tema-
gami, Ontario.

TYPE SPECIMEN: CUPP 10544 on Carex prairea aay May 16, 1918.
Whetzel.

MateErIAL EXAMINED: In addition to the specimens listed in 1929 (p. 30) on Carex
prairea Dewey, C. interior Bailey, C. vesicaria L., C. retrorsa Schw., and C. crinita Lam.
may be reported: On C. prairea: CUPP 17499, McLean, N. Y. May 27, 1929 (apo-

Plate 4. Sclerotinia sulcata. Fig. 15. Apothecia from culms of C. stricta; natural
size. CUPP 11515. Fig. 16. Sclerotia from culms of C. stricta; natural size. CUPP
14747. Figs. 17-18. Cultures from C. stricta on potato dextrose agar showing cottony
aerial mycelium, and sclerotia with spermodochidial masses respectively; 14 natural size.
CUPP 11515 and 14747. a

404

FarLowlA, VoL. 2, 1946

PLATE V

WHETZEL: SPECIES OF SCLEROTINIA 405

thecia) ; — CUPP 23536, McLean, N. Y. May 14, 1934 (apothecia). On Carex oligo-
sperma Michx.: CUPP 17783, Sand Point, and CUPP 17784, near Bear Island, Lake
Temagami, Ontario, Canada, both on September 16, 1929, H. H. Whetzel & Geo.
Thompson (spermodochidia and sclerotia).

This species is doubtless common and widely distributed in northern
United States and Canada. Localities where it occurs are readily discovered
by looking during late summer for the dead and bleached culms with their
characteristic spermodochidia and sclerotia.

Carex oligosperma Michx. is here for the first time recorded as a suscept
of S. longisclerotialis.

SCLEROTINIA CARICIS-AMPULLACEAE
PiaTeE VI

Of all the species of Sclerotinia known to me on sedges and rushes, S.
caricis-ampullaceae Nyberg is the largest, the most striking, and to the eye
of a mycologist, the most beautiful. This species was first brought to my
attention by Dr. John Nannfeldt of the University of Upsala. He sent me
in February, 1933 a collection taken June 25, 1932 near Lake Gardsjon in
Sweden. I recognized it at once as an undescribed species and proposed to
Dr. Nannfeldt that I publish a description of it, naming it in his honor.
Fortunately I delayed the preparation of the manuscript. A year later, in
February 1934, I received another specimen of this species from Dr. Nann-
feldt which had been collected in Finland. Along with the specimen he sent
me a preliminary description of the species prepared by the collector, Mr.
Wolmar Nyberg, who was publishing it under the name the species now
bears. Nyberg, amateur mycologist and banker, had discovered it in early
June 1930, in a sphagnum bog on his villa grounds near Vess6 in the parish
of Borga, where it appeared each succeeding spring. Nyberg’s description
of which he sent me a reprint appeared in the Finnish scientific journal
cited below. :

During the winter of 1937-38, Dr. W. G. Solheim of the University of
Wyoming told me of a remarkably large and beautiful species of Sclerotinia
on Carex aquatilis which his colleague, Dr. C. L. Porter, had discovered in
a high mountain swamp in the Medicine Bow Mountains, in June 1937
(Fig. 25). Duplicate specimens of Porter’s collection together with abun-
dant collections made by Solheim in 1939 and in succeeding years proved
it to be conspecific with the Scandinavian collections. The extraordinary
size and unusual form of the sclerotia of S. caricis-ampullaceae distinguish
it at once from the other species on Carex. Although known from but three
localities, its occurrence in Scandinavia and the high mountains of Wy-

Plate 5. Sclerotinia longisclerotialis. Fig. 19. Apothecia attached to the sclerotia,
from culms of C. prairea; natural size. CUPP 23536. Fig. 20. Spermodochidia and
enclosed sclerotia, from culms of C. interior; natural size. CUPP 15868. Fig. 21. Sper-
modochidia, x6, from culms of C. vesicaria. CUPP 15860. Fig. 22. Young sclerotia
exposed by cutting away the culm epidermis, x2, in culms of C. prairea. CUPP 15872.

406 FaRLow1A, VoL. 2, 1946

Pirate VI

WHETZEL: SPECIES OF SCLEROTINIA 407

oming suggests that it is probably common wherever its suscepts occur
throughout the sub-arctic isothermal zone.

An extensive treatment of this species is to be found in the paper by
myself and Solheim (1943).

SCLEROTINIA CARICIS-AMPULLACEAE Nyberg, Mem. Soc. pro Fauna et
Flora Fenn. 10: 20-23. 1934.

Spermodochidial groups (Fig. 24) when fresh, dark colored, tuberculate,
more or less spaced at intervals along the culm just below the inflorescence,
exposed by slits on two faces of the culm; spermatia hyaline, globose to
slightly ovate, 2.5—3 « in diam., produced successively from the tips of
clustered obclavate spermatophores, oozing forth in mucilaginous drops.

Sclerotia (Fig. 24) extraordinarily large, variable in size up to 20 cm..
long by 1 cm. thick near the base, ventricose below with rounded or bluntly
attenuated base, gradually elongated above into a slender whip-like apex,
developed within the slender 3-angled culm, the base firmly nestled in the
very base of the culm and the enclosing leaf bases, breaking forth through
a slit in the epidermis of the culm to expose the lower half or so, the slender
pointed tip remaining firmly enclosed, black without, sulcate, within pale
pink to white, firm.

Apothecia (Figs. 23, 25) usually several to many, fasciculate, arising at
one point on the exposed surface of the sclerotium, long-stipitate; cup
thistle-funnel-shaped becoming more or less expanded, vinaceous brown,
4—22 mm. in diam. by 7-16 mm. deep, outside finely tomentose, relatively
thin (1 mm.), membranous; stipe long, slender, flexuous, varying in length
depending upon the depth to which the sclerotium is immersed, only the
cup and upper part of the stipe protruding from the water or moss; asci
cylindrical, attenuated below, apex rounded and thickened, pore faintly J+,
192-213 x 11.8-13.5 y, 8-spored, spores uniseriate; ascospores ellipsoid,
hyaline, smooth, without oil drops, 13.5-16.9 x 8.5-10 yp; paraphyses fili-
form, hyaline, branched below, scarcely enlarged toward the tips, 1-2 p»
in diam. —

Hapirat: In culms of Carex aquatilis Wahl., C. aquatilis Wahl, var. altior
(Rydb.) Fern., and C. inflata Huds. in wet swamps and mossy bogs in
sub-arctic or high mountain regions.

TYPE SPECIMEN: On Carex inflata Huds. (=C. ampullacea Good.) , sphag-
num bog near Vesso, parish of Borga, Finland, June 19, 1931. Coll. W. Ny-
berg. Part of type deposited in CUPP 28912. Dupl. in Bot. Inst. Univ.
Upsala, Sweden.

DisTRIBUTION: Finland, Sweden, and Wyoming, U.S.A. Apothecia June—
July, spermodochidia and sclerotia late summer and autumn.

Plate 6. Sclerotinia caricis-ampullaceae. Fig. 23. Apothecia from sclerotia in culms
of C. aquatilis var. altior (specimen from preserving solution). CUPP 28910. Fig. 24.
Spermodochidia and sclerotia in culms of C. aquatilis var. altior; natural size. CUPP
29268. Fig. 25. Apothecia in their natural habitat; natural size.

408 FarLowIA, VoL. 2, 1946

MarTerRIAL EXAMINED: In addition to the type material, I have had in hand: On
Carex aquatilis Wahl.: — Collection made at Selet, near Lake Gardjon, prov. Vaster-
botton, parish of Lovanger, Sweden, June 25, 1932, G. Lohammar, CUPP 21965, (dupl.
in Herb. Bot. Inst. Univ. Upsala). On Carex aquatilis Wahl. var. altior (Rydb.)
Fern.: numerous collections (Whetzel & Solheim 1943: 397) made below Nash Fork
bridge near Univ. Wyoming Science Camp in the Medicine Bow Mts. (alt. 9,600 ft.),
Albany Co., Wyo. during the years 1937, 1939, 1940, 1941, and 1942, W. G. Solheim &
C. L. Porter. On Carex inflata Huds., Solheim, same locality, June 27, 1942. Dupli-
cates of all these collections are deposited in the herbarium of the University of Wy-
oming and the Plant Pathology Herbarium, Cornell University. The collection of
apothecia made June 26, 1939 on C. aquatilis var. altior has been distributed by Sol-
heim as 204, Mycoflora Saximontanensis Exsiccata.

SPECIES ON ERIOPHORUM

As far as I have been able to find, there is but one species of Sclerotinia
reported on Eriophorum. This is Sclerotinia vahliana described by Rostrup
(1891: 607) from specimens collected by J. Vahl, S. Hansen and N. Harz
on botanical expeditions at various times on the west coast of Greenland.
The earliest collected specimen which Rostrup had before him when he
described this species is presumably one collected by J. Vahl for whom he
named the species. A specimen preserved in the Rostrup collections in the
University Botanical Museum, Copenhagen, bears the label “E. Rostrup’s
Svampesamling. 1. Sclerotinia Duriaeana Tul. (Carex rigida). Nonnese
5/1829 by J. Vahl.” Through the kindness of the director of the museum,
Johs. Grgntved, this specimen was loaned me in September 1927 (Fig. 28).
A critical examination satisfied me that this is not S. duriaeana and that
the suscept is a species of Eriophorum. Whether or not this is one of the
specimens on which Rostrup based his description it is now impossible to
say. Since this is earliest known collection of the species and was collected
by J. Vahl, I am designating it the type specimen of S. vahliana Rostr.

SCLEROTINIA VAHLIANA
Prate VII anp Text Fic. B

Rostrup gives Eriophorum scheuchgzeri as the suscept for all the collec-
tions he had from Greenland and lists, in addition to Vahl, two other col-
lectors, S. Hansen and N. Harz, who appear to have made their collections
during the years 1888 and 1889 (Rostrup 1891: 593). These specimens
were presumably deposited in Rostrup’s collections in the Botanical Mu-
seum in Copenhagen, but I have no record of having seen them when I was
there in 1930.

I know of but two other collections identified as S. vahliana. One of these
is a collection made by J. A. Nannfeldt in northern Sweden in 1927 on
Eriophorum angustifolium and another collected by Vestergren, e¢ al. in the
same region in 1903 on the same species and distributed under 730, in
Vestergren’s Micromycetes Rariores Selecti. These agree well in all their
characters with the type collection.

During the summer of 1930 I made collections of a Sclerotinia attacking
E. angustifolium (Fig. 26) in Denmark and in southern Sweden. I was able

WHETZEL: SPECIES OF SCLEROTINIA 409

to study only the early stages of the infection which was developing while
I was there during the months of June and July. I obtained the fungus in
pure culture (Fig. 27) and also found the sclerotia in early stages of de-
velopment in culms in the field. Such evidence as I was able to get, indicates
that the fungus in these collections is probably S. vahliana.

It seems desirable to present here a translation of Rostrup’s original
description.

“376. Sclerotinia Vahliana n. sp.

“Apothecia arising from a sclerotium, stipitate, brown or castaneous. Cup hemi-
sphaerical, length and height, 4-8 mm. Stipes flexuous, tomentose, striate below, 1-3 cm.
long, about 1 mm. thick. Asci cylindrical-clavate, long stipitate, length (including stipe)
150-170 w x 8-10 uw. Spores ellipsoid, 11-14x 4-6 w. Paraphyses numerous. Sclerotia
black, oblong, compressed, sulcate, 1 cm. long by 0.5 cm. thick.

“Between the leaf sheaths of Eriophorum Scheuchzeri Hoppe, Greenland (Vahl),
Umanak (S. Hansen), Egedesminde, Holstenborg, Godthaab (N. Harz).

“Sclerotia assembled, 2-3, between the lower leaf sheaths, with 3-4, deep furrows
along the outward turned, free side.”

Based upon Rostrup’s original description of the apothecia and my studies
on the specimens above referred to, the following would appear to be a
fairly complete description of this species.

SCLEROTINIA VAHLIANA Rostr. Jn Tillaeg til “Grgnlands Svampe (1888).”
Meddel. om Grgnland. 3: 607-608. 1891.

Spermodochidia not found.

Sclerotia (Fig. 28 and Text Fig. B) variable in size and shape, chelle like
in the form of a short, lobed half-cylinder or hollow hemisphere, up to
2—3 cm. in civciimference, about 2—3 mm. thick, black without, white within,
surface rough, irregularly and deeply creased, with low ridges, usually 3—4
in each culm, obviously originating within the culm but finally breaking
forth to complete its growth. This peculiar morphology of the sclerotia has
been recorded by Rostrup (1903: 315) who quotes a note by Ostenfeld, the
collector of specimens of this species from Iceland: “Sklerotierne hdjst
uregemaessigt bugtet-foldede” (Sclerotia very irregularly curved and
creased). The internal structure is essentially like that of other species of
Sclerotinia,

Apothecia (Figs. 29, 30) 1-several arising fasciouleely from some point
on the sclerotium, broadly and deeply cup-shaped, castanean brown, short
stipitate; cup up to 10 mm. broad by nearly as deep; stipe 1-3 cm. long x
1 cm. thick, flexuous, tomentose, darker than cup, black toward base; asci
cylindrical clavate, long-attenuate below, tip rounded, not thickenéd, 50
meas. (type) 109-164 x 7.8-10.4 », mode 151.7x9 yp, av. 144x 8.6 p, 8-
spored, spores uniseriate; ascospores long ellipsoid to fusoid, flat to concave
on one side, biguttulate, 50 meas. (type) 10.8-18 x 4.8-6.6 p, mode 12 x
5.4 pw, av. 13x 5.6 »; paraphyses numerous, slender, simple or branched,
3 » diam., slightly or not at all enlarged at the tips.

In culture (Fig. 27) on potato dextrose agar forming a sparse white
webby aerial mycelium; submerged mycelial mat dark olivaceous brown.

410 FARLOWwIA, VoL. 2, 1946

Pate VII

WHETZEL: SPECIES OF SCLEROTINIA 411

Sclerotia formed abundantly, irregularly discoid or oblong, flat or low loaf-
shaped (from diseased stems collected near Lyngby, Denmark).

HasitatT: In the culms of Eriophorum scheuchzeri Hoppe, FE. vaginatum
L., and E£. angustifolium Roth. in sphagnum bogs, apothecia, May—July;
sclerotia, late summer and autumn. An arctic and sub-arctic species.

DISTRIBUTION: Reported from Greenland (Rostrup 1891 and 1906), and
Iceland (Rostrup 1903 and Larsen 1932). Known to me from Denmark
and Sweden. Should be found in the arctic and sub-arctic zones of con-
tinental North America.

TYPE SPECIMEN (Fig. 28): The collection by J. Vahl on Eriophorum
(scheuchzeri ?) Nonnese, Greenland, May 1829. Deposited in the Botani-
cal Museum, Copenhagen, under the name, “Sclerotinia Duriaeana Tul.”
Photograph and microscopic mount deposited in CUPP 16275.

MATERIAL EXAMINED (in addition to the type specimen): On Eriophorum vagi-
natum L.: CUPP 31589, sphagnum bog along north shore of. Lyngby $6, Lyngby,
Denmark, June 7, 1930, H. H. Whetzel (early stages of infection) ;— CUPP 31591,
swamp in forest at Nosten, parish of Bondhyrho, near Upsala, Sweden, July 3, 1930,
H. H. Whetzel & J. A. Nannfeldt (early stages of infection). On Eriophorum angus-
tifolium Roth. (=£. polystachon L.): Vestergren, Micromycetes Rariores Selecti, 730.
Lapponica Tornensis, Vassijaure (Sweden), July 1903, A. Roman, N. Sylvén & T. Ves-
tergren (apothecia and sclerotia) ; — “Fungi Suecici”. University Upsala. Dupl. N. Y.
Bot. Gard. and CUPP 31606. Collected in Torne, Lappm. Jukkasjarvi. s:n. Abisko,
Sweden, June 30-July 5, 1927, J. A. Nannfeldt (apothecia and sclerotia) ; - CUPP
- 31590, swamp in forest near Nosten, parish of Bondhyrho, near Upsala, Sweden, July 3,
1930, H. H. Whetzel & J.A. Nannfeldt (early stages of infection).

This species is very distinct from the others in cyperaceous suscepts, be-
ing especially marked by the unusual form and development of its sclerotia.
While I think that the Sclerotinia which I found in its early infection activi-
ties in the culms of E. vaginatum and E. angustifolium in Denmark and
Sweden is probably S. vahliana, it is unfortunate that I was not early
enough in the localities to get the apothecia and did not remain late enough
in the summer to follow the development to mature sclerotia. I feel quite
sure that spermodochidia of some type would have developed during late
July or August in the infected culms. Tiny masses of densely protoplasmic,
large, thin-walled hyphae, which are present at intervals in the diseased
culms which I preserved (CUPP 31589) are characteristic of the initial
stages of spermodochidial development in the other species of the cyperi-
colous species of Sclerotinia.

The sclerotia of S. vakliana appear to have an unusual history of develop-
ment (see Text Fig. B). They are initiated as in the related species, within
the culms of the suscepts. I was able to observe only the early stages in

Plate 7. Sclerotinia vahliana. Fig. 26. Diseased culms of Eriophorum vaginatum ;
Y% natural size. CUPP 31589. Fig. 27. Culture from culms of £. vaginatum on potato
dextrose agar showing sclerotial formation; natural size. CUPP 31589. Fig. 28. Type
specimen in Botanical Museum, Copenhagen (revived dry material); natural size.
CUPP 16275. Figs. 29-30. Apothecia on sclerotia from £. angustifolium from Sweden
(revived from liquid preservative) ; natural size. CUPP 31606.

412 FARLOWIA, VOL. 2, 1946

this development. The sclerotia begin as elongate, cottony, white masses
at three or four places in the culm. As the young sclerotia begin to solidify
they become pink in color as do those of related species. In a few cases I
found evidences of the developing rind. I found no mature sclerotia, all
being still firmly enclosed within the culms. I observed that the infected
culms often appeared to be stouter and apparently to have been more
vigorous in their growth than were the healthy ones. This may have been
due to stimulation by the fungus during the first days of infection. As far
as I was able to discover, these sclerotia were developing only in the upper
internode of the culm in E. vaginatum. It seems almost certain, however,
that sclerotia are formed at intervals throughout the culms even to the base.
They apparently do not complete their development within the culm but,

Text Fic. B

Text Fig. B. Sclerotium of Sclerotinia vahliana from base of culm in crown of Erio-
phorum, (a) front and (b) back view, (c) from one higher up in culm; all x2.
Drawn by Miss Charlotte Dill.

bursting out, continue to grow in size. The mature sclerotia I have seen
bearing apothecia are obviously too large to have fully matured within
the culms.

Rostrup’s description of the sclerotia as ‘‘assembled between the lower
leaf sheaths” suggests that in the dwarfed Eriophorum plants of Greenland
and Iceland, they may be formed chiefly in the basal region of the culm.
However, I have seen specimens of sclerotia bearing apothecia which had
clearly developed in the upper reaches of the culm (e.g. Nannfeldt’s collec-
tion of 1927, Text Fig. Bc). Careful observations in the field may disclose,
in the case of sclerotia clustered in the crowns of the suscepts, that the base
of the leaves as well as the culm have been involved in the nourishment of
these sclerotia. Larsen (1932), in recording the occurrence of this species
in Iceland, cites collections by himself and by Ostenfeld and remarks, “On
tufts of Carex and Eriophorum on swampy ground. The pea-sized sclerotia
are often seated in the axils of the leaves, covered by the leaf sheaths”. It
is very doubtful that S. vahliana ever attacks Carex species. Larsen is either

WHETZEL: SPECIES OF SCLEROTINIA 413

in error or has confused some species of Sclerotinia actually occurring on
Carex in Iceland, with S. vahliana on Eriophorum.

SPECIES ON SCIRPUS AND SCHOENUS

A species of Sclerotinia on Scirpus lacustris L., long known to European
mycologists, chiefly in its sclerotial stage, was finally described by Rehm
(1896) under the name of Sclerotinia scirpicola. Although represented in
herbaria by but few collections, it is certainly common and widely distrib-
uted throughout northwestern Europe. It is not known from North Amer-
ica. No species of Sclerotinia is known to occur on other species of Scirpus.
Ferdinandsen & Winge (1911: 293) report a collection by Ade in Bavaria,
Germany, said to be this species on Scirpus silvaticus (“the spores finally
becoming 2-celled’”’). Since the spores of S. scirpicola are uniformly 1-celled,
it is very doubtful that Ade’s specimen was correctly determined. There is
however a species, pathogenic to the closely related Schoenus nigricans L.,
which appears to have been generally overlooked by mycologists.

SCLEROTINIA SCIRPICOLA
Pirate VIII ano Text Fic. C

This species is of peculiar interest because of the nice adaptation it has
made to its aquatic habitat. Its suscept is an inhabitant of the shallow
waters of lakes and ponds. Not only do the freed sclerotia float like tiny
corks upon the water but in the spring the apothecia develop on these where
they accumulate in thousands on the low flat shores as the water subsides.
Aside from the original description of the apothecial stage by Rehm (based
upon dry material) the papers by Ferdinandsen & Winge (1911 and 1913)
and Sydow’s (1912) appear to be the only contributions of importance
dealing with this species. During my stay at Lyngby, Denmark, during
June 1930, an exceptional opportunity was afforded for study of S. scrpicola
as it occurs in Lyngby S6. As a result of these studies a more complete
picture of this species can now be drawn.

SCLEROTINIA SCIRPICOLA Rehm in Rabenh. Krypt. Fl. 1°: 822, 1893.

Sclerotium roseum Moug. in Fries, Syst. Myc. 2 (Suppl.): 43, 1830. (=Elench. Fung.
3: 43. 1828).

Sclerotium roseum Kneiff in Duby, Bot. Gall. 2: 874. 1830.

Sphacelia scirpicola Ferd. & Winge, Biolog. Arbej. Tilegnede, Warming, Nov. 3, p.
290, 1911.

Myrioconium scirpi Syd. Ann. Myc. 10: 449. 1912.

Myrioconium scirpicola V. Héhn. Mitt. Bot. Inst. u. Tech. Hochsch. Wien 3: 50.
1926.2

Spermodochidial groups (Figs. 31, 32) more or less oval to elongate,
shallow, slightly raised, very variable in size, often minute, irregularly

® Ferdinandsen & Winge (1913: 24) admitted the logic of Sydow’s argument (in a
letter) for referring the conidial stage of S. scirpicola to his genus Myrioconium but
they did not actually make the combination M. scirpicola, as Von Héhnel seems to imply
in this 1926 paper.

414 FarLowIA, VoL. 2, 1946

“Prate VIII

-WHETZEL: SPECIES OF SCLEROTINIA 415

disposed along one side of the culm for some distance just below the heads,
dull black, rupturing by slits in the epidermis, from which the viscid mass
of spermatia are discharged; spermatia minute, globose, about 3.3 » in diam.
The dark color of the spermodochidial groups is due to the brownish color
of the massed spermodochia, the brown gummose contents of the epidermal
cells, and to brown-walled intercellular hyphae in some of these cells.

Sclerotia (Fig. 34) usually 8-10 per culm, the first one developing with
the appearance of the spermodochidia and just below them; successive
sclerotia appear a few inches apart as the invading hyphae spread down-
wards in the culm. The sclerotia originate as soft pinkish mycelial masses
in the loose central parenchyma of the culm, at maturity appearing as large,
longitudinally lobate, oblong, blunt, black, tuberoid bodies completely en-
closed in the culm. The culms, weakened at these points, are easily broken
by the wind, discharging the sclerotia into the water, where they float, and
drifting with the wind, accumulate, together with broken pieces of the culms,
in great patches along the leeward shores of the lake or pond. The sclerotia
vary much in shape and size, being usually somewhat longer than broad,
ranging from 2-10 mm. by 2—5 mm. in diameter. The form of the sclerotium
is striking, being elongately 2—3-lobed, the ends of the lobes blunt and
rounded. This compound appearance is more apparent than real, being due
to the structure of the culm. A cross section of the sclerotium discloses a
narrow outer black rind, one to two cells thick, enclosing a medulla of rather
loosely interwoven, relatively thin-walled, large hyphae, distinctly septate
and often constricted at the septa. The medulla long retains its bright pink
color, even after the formation of the rind, but eventually becomes white.
The lobation of the sclerotium is marked out in the cross sections by faint
brown lines, the persistent brownish-walled cells which formed the firmer
tissues surrounding the cavities originally filled with thin-walled pith cells
which have been largely digested by the developing sclerotium.

Apothecia (Figs. 34, 35) appearing in the spring at flowering time of the
Scirpus; 1-to several from each sclerotium, usually not more than 2, arising
at any point on the sclerotium, variable in size, when fully expanded 3-15
mm. in diam., av. 8-10 mm. The size of the apothecium apparently does
not depend entirely upon the size of the sclerotium for several, all usually
small, or one much larger than the others may arise from a single sclerotium;
cups deep goblet-shaped becoming nearly flat expanded, light fawn to vina-
ceous brown, lighter beneath, due to a coating of fine hyphal fuzz; stipe
usually short, slender, concolorous with under side of the cup, also covered
with a hyphal fuzz above, clothed below, especially at the base, with a dense
mat of long, dark hyphae which serve to anchor the large apothecia to the

Plate 8. Sclerotinia scirpicola. Fig. 31. Diseased culms and inflorescence of Scirpus
lacustris showing spermodochidia; about natural size. Fig. 32. Part of culm showing
spermodochidia, x6. Fig. 33. Showing method of inoculating pistils of S. lacustris for
study of invasion phenomena; about natural size. Fig. 34. Sclerotia and young
apothecia; natural size. Fig. 35. Sclerotia bearing mature apothecia; natural size. All
from specimens in CUPP 31607.

~

416 FarLowlA, VoL. 2, 1946

ONguypea——\

D en

be u
Des 3 jo 8 FOC U

ol

:

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O
&y

s

Og

S900

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cS

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aa oe
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Text Fig. C. Cross section through spermodochidia of Sclerotinia scirpicola about x400.
Note that each cavity is directly beneath a stomate, and is formed chiefly by crowd-
ing aside the palisade cells rather than by digesting them. Drawn by Carlos Garcés.

surrounding floating debris of broken culms; asci long, cylindrical, apex

blunt, of about the same diameter throughout, except at the base where .

they are rather sharply narrowed to about half the diameter of the apical

portion, 8-spored, spores uniseriate in the upper third of the ascus, 50 meas.

(fresh, in water) 133.3-181.8 x 8.1-10.8 », mode 157.5 x 9.5 uw, av. 153.2x

9.4 w; ascospores long oval, 100 meas. (fresh, in water) 11.2-17.8 x 5.3-7.9

#, mode 13.8 x 6.6 w, av. 14.4 x 6.6 » surrounded by a very evident mucilagi-

nous coat; on germination in water often producing spermatophores and

spermatia directly; paraphyses slender, not appreciably swollen toward the
tips, about 3.4 » in diam.

WHETZEL: SPECIES OF SCLEROTINIA 417

In culture on potato dextrose agar forming a thin, white, aerial, webby
coating over the surface of the agar; submerged mycelium feathery as in
cultures of S. longisclerotialis; no sclerotia formed. ;

Hastitat: In the culms of Scirpus lacustris L. in the shallow waters of
lakes and ponds. Apothecia developing on sclerotia stranded along the
shore line among sedges, grasses and other plants which afford continuous
shade and moisture during their development.

DISTRIBUTION: Throughout western Europe; very abundant in Den-
' mark (Ferdinandsen & Winge 1911: 293). Also reported from Finland,
Germany, Sweden, and England. Probably to be found in equal abundance
when sought in neighboring regions of northern Europe; not known from
North America. ;

TYPE SPECIMEN: The specimen upon which Rehm (1893: 823) based
his original description of this species, was found by him in Winter’s her-
barium. He says it bore the label ‘“Peziza tuberosa” with the notation
(transl.) “in May, the old culms of Scirpus lacustris, in our lakes break up.
When these fall into the water, this Peziza appears from sclerotia in a few
days, from every piece which with us very frequently contains Sclerotium
roseum (Moug.) Fries.” According to Rehm the specimen was probably
collected by Winter in Saxony near Zweibriicken in the Rheinpfalz (Ger-
many). I have been unable to discover where this type specimen (if indeed
it is still extant) is now deposited.

ILLUSTRATIONS: .Ferdinandsen & Winge, Saertryk af Biol. Arbej. Til-
egnede. Eug. Warming. Nov. 3, fig. 1-7. 1911.

MATERIAL EXAMINED: Exsiccati Specimens: In the Kew Herbarium there is a speci-
men of three pieces of culm each with an enclosed sclerotium labeled ‘“‘Sclerotium
roseum” with a brief written description signed by Persoon. The specimen is unnum-
bered, undated and locality where collected not indicated; — Mougeot et Nestler, Stirp.
Crypt. Vogeso-Rhen. 884 (1826). “Sclerotium roseum Kneiff-Pers.-Fries. in Litt.”
Specimens of this in the British Museum, in Kew, and at Cornell show sclerotia in
culms said to have been collected near “Argenoratum”, now Strassburg, in Alsace,
France. Collected by Kneiff in autumn. No date of collection given; — Kneiff et Hart-
mann; Pl. Crypt. Badens, 20, 1828. “Sclerotium roseum Kneiff (nov. sp.)”. This speci-
men in the British Museum consists of pieces of culms containing mature sclerotia
collected in canals along the Rhine near Kehl in Germany. Sept—Oct. No year of
collection given and no collector cited; — Klotzsch, Herb. Mycol. 244 (1842). “Scle-
rotium roseum Moug. Intra culmos Scirpi lacustris ad Dresden”. This specimen in Kew
Herbarium bears only the data above quoted. It consists of a piece of culm containing
a sclerotium ; — Roumeguére, Fung. Gallici Exs. 499, 1879. “Sclerotium roseum Kneiff.
Pers. Fries. in Litt. Moug.” This number was made up from material remaining in
Mougeot’s herbarium and is of the same origin as his Stirp. Crypt. Vogeso-Rhen. 884.
Specimens examined in the Kew Herbarium and in CUPP show a single sclerotium in
a piece of culm; — Otto Jaap, Fung. Sel. Exs. 755. Collected by Jaap in Brandenburg,
Triglitz in the Prignitz, Germany. June 15, 1916 (spermodochidia (?), sclerotia and
apothecia) ; — Flora Suecici 4222. Collected by J. A. Nannfeldt on shore of shallow arm
of Malaren. Bondkyrok s:n. Skarholmen, Uppland, Sweden. June 25, 1931 (apothecia).
Dupl. CUPP 32615; In Kew Herbarium a spec’men labeled: “Sclerotinia scirpico'a,
Ashgrove Loch, Shevenston, Ayrshire 26.2.21. Coll. D. A. Boyd.” Consists of a single culm
of S. lacustris containing a single sclerotium ; — CUPP 31607, along the shore of Lyngby
$6, Lyngby, Denmark, June 3-July 13, 1930, H. H. Whetzel (spermodochidia, sclerotia,
apothecia and cultures). Duplicate specimens from this collection have been distributed

418 FartowlA, Vov. 2, 1946

to the following herbaria: Kew, England; British Mus., London; Univ. Toronto, Can-
ada; Farlow Herb., Cambridge, Mass.; N. Y. Bot. Gard.; Univ. Mus., Ann Arbor,
Mich.; Missouri Bot. Gard., St. Louis, Mo.; Mycol. & Pl. Dis. Survey, Bur. Pl. Ind.,
Washington, D. C.; Univ. California, Berkeley; Univ. Iowa, Iowa City.

The abundance of apothecia in Lyngby Sé during my stay in Lyngby,
and the nearness of laboratory facilities for critical studies and culture
work, presented an opportunity not to be neglected. I devoted most of my
time to such studies, the results of which are summarized below.

Lire History STUDIES

The life history of S. scirpicola, while in general quite like that of its near
relatives on Carex species, presents some unique features largely due to the
habits of its suscept Scirpus lacustris. This Scirpus grows in the shallow
waters along the shores of the lake. It is most vigorous and fruits most
luxuriantly near the shore, the plants becoming more scattered, shorter and
usually producing fewer flowering culms as the plantation spreads out into
the deeper water. When we arrived at Lyngby, the first week in June 1930,
I found the apothecia already developing in great abundance from the
thousands of sclerotia lodged in debris on low-lying projections of the shore,
a few minutes’ walk from the laboratory of the Phytopathological Experi-
ment Station in which we were kindly afforded working facilities by the
Director, Mr. Ernest Gram. The shores of this shallow lake are fringed
with great patches of Scirpus lacustris and the prevailing southwest winds
of the previous fall and winter had deposited the sclerotia in immense
numbers along with the broken culms of Scirpus just above the receded
water level on every flat projection of the northeast shore of the lake and
on the water where they were anchored and shaded by a rich growth of
grasses and other shore-loving plants and shrubs. The open apothecia
reached their maximum abundance about the end of the first week in June.
A week later most of the apothecia were shrivelling or decaying, though
new ones were still to be found just sprouting from sclerotia here and there.
By the 17th of June, only an occasional apothecium was to be found. Many
heads of the Scirpus were already in bloom on June 2 when we arrived, but
blossoming reached its maximum about the time apothecia were to be found
in greatest abundance. The weather throughout June was warm, the skies
clear, and practically no rain fell. From the abundance of mature apothecia
at the beginning of June, we judge that they must have begun to open about
the middle of May. This would indicate that the period of apothecial pro-
duction normally extends over a period of three to four weeks.

The mature apothecia discharge their ascospores in tiny puffs which,
caught by gentle air currents among the shading vegetation, are wafted
upward, some of them eventually reaching the white extruded pistils of the
Scirpus inflorescence, where they lodge on the feathery stigmas. By sus-
pending open apothecia over the inflorescence (Fig. 33) of culms (with cut
ends in water) placed under an inverted large test tube, I was able to obtain
naturally inoculated pistils. These when removed and mounted on glass
slides the following morning, afforded excellent material for study of spore

WHETZEL: SPECIES OF SCLEROTINIA . 419

germination and penetration. The spores germinate promptly, sending out
from one end a stout simple germ tube. This penetrates the projecting
hyaline cells of the stigma, usually forming a swollen globose appressorium
where the tip has applied itself to the surface of the stigmatic cells. From
this appressorium a penetration tube is sent into the cell from which an
invasion hypha passes down into the tissues of the pistil. Infection is
promptly indicated by the browning and shrivelling of the pistils. Un-
inoculated pistils remain white and plump for several days under the same
conditions. The fungus having entered the ovary, growth seems to be
greatly slowed down. In the open, about two weeks after inoculation, one
or more of the peduncles supporting the individual spikes of the inflorescence
begin to turn brown, the mycelium having killed the spike and spread
downward into the peduncle through which it reaches the main stalk of
the culm.

A dark browning just below the point of attachment of the inflorescence
announces the invasion and infection of the culm proper. Visible infection
of the culm becomes evident about three weeks after inoculation, as Ferdi-
nandsen and Winge had already discovered. Rapid progress of the infection
down the culm, however, does not usually occur for another week or so.
It is evidenced by a hydrotic discoloration of the culm, with disappearance
of the chlorophyll and discoloration in the form of alternating broad bands
of a light and dark brown color. When this zonation has extended a foot
or so down the culm, the spermodochidial groups begin to appear in an
irregular line on one side of the culm in the brown necrotic tissues several
inches below the inflorescence (Fig. 31). These spermodochidial areas are
very variable in size, ranging from mere specks to elongate or irregularly
ovate, black, slightly raised patches just beneath the epidermis. They are
usually very numerous and the tissues in which they develop are generally
of a distinctly lighter brown color than the adjacent regions. They finally
rupture by slits, discharging the viscid masses of spermatia as the first
sclerotial initials are to be detected just below. The downward progress of
the fungus, indicated by hydrosis and zonate discoloration, now proceeds
rapidly with the successive formation of sclerotial initials at intervals of a
few inches. As many as ten to fifteen sclerotia may appear in a long vigor-
ous culm, according to Ferdinandsen and Winge. When we left Lyngby on
July 15, the first sclerotia were just beginning to appear occasionally in an
infected culm.

As the sclerotia mature, they rupture the epidermis over them by several
slits. The dead culms, weakened at these points, are gradually broken over
by the winds of late summer and autumn. The breaking up of the ice in
the spring by storms completes the breaking up of the culms, freeing the
sclerotia which float on the surface of the water and are thus transported
by the waves to their final resting place among the debris along the shore,
ready to germinate and produce apothecia under the favorable conditions
of temperature, shade, and moisture prevailing at the blossoming time of
the Scirpus.

420 FarLowlA, VoL. 2, 1946

Ascospores discharged onto potato dextrose agar or disinfested segments
of infected culm of Scirpus planted in the same medium give identical
mycelial growths. Growth both from ascospore sowings and from tissue
plantings is prompt and vigorous, like that of S. swcata. Aerial mycelium
is abundant, usually loose and cottony, at first pure white (never pink)
becoming denser with age and faintly brownish, often with large dense
masses of a dirty grey color, especially in test tube slants. The submerged
mycelium becomes blue black at the center of the colony with radiating
dark brown feathery strands. Spermodochia were not observed on’ this
medium. Sclerotia were only tardily developed on test tube slants but were
numerous in Petri dish cultures, being globose, black outside, pink where
they grow against the glass and more or less completely buried in the dense
aerial mycelium.

Although the morphology and life history of S. scirpicola are now per-
haps more fully known than those of any of the related species on cypera-
ceous suscepts, a number of features require further consideration.

It seems probable that some of the sclerotia may remain dormant through
the summer following their formation, as numerous hard, plump, ungermi-
nated sclerotia were found among the debris along the shore of Lyngby S6
long after the last apothecia had disappeared. On the other hand, these
sclerotia may not be capable of germination for some unknown reason,

The germinability of the sclerotia may be conditioned in some way by the
functioning of the spermatia. The writer’s original suggestion (Whetzel
1929: 18) that the microconidia probably function as do the spermatia of
the rusts has since been supported by the work of Drayton (1934) in his
work on Sclerotinia gladioli. Ferdinandsen & Winge (1911: 297) think the
microconidia may function as true conidia, causing secondary infections
later in the season when primary infections by ascospores are no longer
possible. This seems very doubtful though they cite some observations in
support of this hypothesis. The writer has had no satisfactory opportunity
to attempt a careful study of the germination and role of the spermatia,
but believes that S. scirpicola is a favorable species for such an investigation.
It occurs in great abundance in Lyngby within easy walking distance of
excellent laboratory facilities. It is to be hoped that Professor Ferdinandsen
or some of his students will undertake intensive investigation on this very
interesting and important problem, as they are most favorably situated for
such an undertaking.

The suggestion of Ferdinandsen and Winge that invasion by the fungus
takes place through the culm at some point just below the inflorescence
does not appear tenable. Ascospores shot on to culms at this point, in
moist chambers, were observed to germinate poorly or to produce spermatia
only. No germ tubes could be found penetrating the epidermis or entering
through the stomata, which are numerous all over the culm, while actual
invasion of the stigmas was easily demonstrated. The death of the pedicles
of infected spikes prior to the first evidences of infection in the culm itself
also argues for blossom invasion. While the very limited and preliminary

WHETZEL: SPECIES OF SCLEROTINIA 421

investigations which we have been able to make all point definitely to the
pistils of the Scirpus as the infection court, more extensive studies and
inoculation experiments should be made.

ORIGIN AND RELATIONSHIPS

Undoubtedly Sclerotinia scirpicola is very closely related to the species
on Carex. It differs from these chiefly in the characters of its spermodo-
chidia and sclerotia. On the whole, it approaches more nearly S. sulcata
rather than S. duriaeana, as suggested by Ferdinandsen and Winge who,
however, at that time were not aware of the fact that there were two distinct
species confused under the name S. duriaeana. Its similarity to S. sulcata
is most marked in its cultural characters. Its dense cottony growth of aerial
mycelium and large tuberoid sclerotia produced on potato dextrose agar
are scarcely to be distinguished from those produced by S. sudcata in pure
cultures. Sclerotial production on potato dextrose agar is, however, much
more tardy and less certain, while the growth of aerial mycelium is gen-
erally more abundant; the submerged mycelium is also more promptly and
extensively darkened. Its spermodochidial patches also simulate those of
S. sulcata in their irregular distribution on the culm. The form of its
sclerotia in nature is quite distinct but may possibly be due entirely to the
structure of the Scirpus culm. It is probably restricted in its pathogenism
to Scirpus. Many Carex species were common about the shores of Lyngby
S6, but only two or three apothecia arising from sclerotia of the Carex type
were found among the apothecia of S. scirpicola along the lake shore. Cul-
tures from one of these proved to be typical of S. duriaeana. Infected culms
of C. disticha in the same locality yielded typical cultures of S. sulcata.
Moreover, the period of apothecial development is distinctly later for S.
scirpicola than for the species on Carex. Apothecia of the species on Carex
were rarely to be found about Lyngby when we arrived at the beginning of
June, although infection of the culms of C. Hudsonzii by S. sulcata were dis-
covered in abundance shortly after our arrival. The blossoming period for
Carex species was also practically past. Since two to three weeks arg re-
quired for the development of visible infection after inoculation by S. du-
riaeana or S. sulcata in their Carex suscepts, the scarcity of apothecia of
these species when those of S. scirpicola were at their optimum abundance
is significant In spite of all this evidence as to the specific identity of
S. scirpicola, careful cross inoculation experiments should be made with
this form on species of Carex known to be suscepts for S. sudcata and S.
duriaeana. Here is another interesting problem for some energetic young
pathologist or mycologist who can avail himself of the exceptional oppor-
tunity afforded there at Lyngby.

The marked similarity of S. scirpicola to S. sulcata appears to be of con-
siderable significance in connection with my theory of the origin and evolu-
tion of this group of species on cyperaceous suscepts. It would seem that
S. scirpicola has been a relatively recent differentiation derived directly
from the European form of S. sudcata, which is probably also the mother

422 FaRLowIA, VoL. 2, 1946

PLATE IX

WHETZEL: SPECIES OF SCLEROTINIA 423

species of S. duriaeana. The characters peculiar to S. scirpicola may, in
part at least, be due to its adaptations to a different suscept.

While Mougeot is credited by Fries with having first applied the name
Sclerotium roseum to the original collecton made by Kneiff, there is sub-
stantial evidence that Kneiff himself is the legitimate author of the name.
Not only did Mougeot and Nestler credit Kneiff with this name on the
label of their Stirp. Crypt. Vogeso-Rhen. 884 (1826), but also Kneiff and
Hartmann, two years later, in distributing this sclerotial stage, in their Pl.
Crypt. Badens 20, credit the name to Kneiff. Subsequent writers in refer-
ring to this name are divided as to the legitimate author of it. Tulasne and
others follow Fries, but most of them credit Kneiff with the name. Sac-
cardo (1899: 1153) notes that the name “Sclerotium Knezffi” is applied to
this fungus by others (“Quibusdam”’). I have been unable to locate any
other reference to the fungus under this name.

Currey’s assumption (1857) that “‘Sclerotium roseum Kneiff” is the
sclerotial stage of “Peziza curreyana” was accepted by all subsequent stu-
dents of S. curreyana until Rehm (1893) pointed out the error in the note
to his description of Sclerotinia scirpicola. This confusion of the sclerotia
in the culms of Scirpus with those in the culms of Juncus as representing
one and the same species had not, however, escaped suspicion in the critical
mind of Tulasne (1861:, 105) who while admitting that the sclerotium in
Scirpus “. . . seems to agree completely with it (in Juncus) in its whole
structure though no one has yet found it in fruit” remarks “But we are
sorry that it has not been possible for us to test it by suitable culture, for
we have never seen it alive”.

SCLEROTINIA SCHOENICOLA
PraTE IX, Fics. 36-38

Desmaziéres appears to have been the only mycologist to refer to speci-
mens of what proves to be a heretofore unrecognized species of Sclerotinia.
He took the sclerotial and spermatial stages of this fungus on Schoenus
nigricans (Figs. 36-38) to be the same as those on species of Carex which
he designated respectively Sclerotium sulcatum and Epidocium affine, and
which were later (1929) oe by Whetzel as stages in the life cycle
of Sclerotinia sulcata.

Desmaziéres 7 distributed specimens of the culms of Carex vulpina and
acuta and culms of Schoenus nigricans all of which contained sclerotia very

7 Plantes Cryptogames France, Ed. 1, Ser. I, No. 2029 (1850); Ed. 2, Ser. II, No.
1629 (1853).

Plate 9. Sclerotinia schoenicola and S. luzulae. Fig. 36. Culms of Schoenus nigricans
showing spermodochidial patches, from Desmaziéres’ Plantes Crypt. France, Ser. II, 21;
about natural size. Fig. 37. Spermodochidial areas, x6. Fig. 38. Sclerotia of S. nOhGe:
nicola from culms of Schoenus nigricans from Desmaziéres’ Plantes Crypt. France, Ed. 2,
Ser. I, 1629; natural size. Fig. 39. Culms of Luzula pilosa from Krieger’s Fung. Sax.
2073 showing sclerotia of Sclerotinia luzulae (revived dry specimens) ; natural size.
Fig. 40. Portions of a sclerotium still enclosed in the culm, x6.

424 FarLowlA, VoL. 2, 1946

similar in character (see Whetzel 1929: 7) and which he collectively la-
beled “‘Sclerotium sulcatum Rob. in herb.” No date or locality is indicated
on the packets. Presumably Roberge was the collector. In specimen 2029
the culms were all mixed together. In his formal description of Sclerotium
sulcatum, Desmazieres (1851) does not mention Schoenus nigricans spe-
cifically, saying only ‘hab. in culmis siccis Caricum’’; but in the note fol-
lowing he says (transl.) “This fungus was found inside triangular stubble
of Carex vulpina and C, acuta and in species with cylindrical culms”’, the
last referring undoubtedly to S. nigricans which, in the later distributed
specimens (number 1629), were segregated into a separate packet and
labeled Schoenus nigricans. His description clearly applies primarily to the
sclerotia in the Carex culms, thus in effect leaving those in S. nigricans
without a name.

Desmazieres * later (1853) distributed the upper portions of the same
culms containing the spermatial fruit bodies as those containing the scle-
rotia (Ed. 1, Ser. I, 2029; Ed. 2, Ser. II, 1629) distributed in 1850. Here,
however, he separated the culms of Schoenus from those of Carex in differ-
ent packets. In his description of the spermatial fruit bodies (1853) he
gives these the name ‘“‘Epidocium affine Desmaz.” and gives both Schoenus
nigricans and Carex species as suscepts citing his exsiccati specimens Series
II, 27. His description, however, clearly applies primarily to the fruit bodies
on the Carex culms, thus again leaving those on Schoenus nameless.

The evidence clearly indicates that the sclerotia and the spermodochidia
on Schoenus nigricans belong to the same fungus. The spermodochidial
patches on the culms of S. nigricans are quite distinct in character from
those of E. affine and from those of the other Sclerotinia species on the
Cyperaceae. (Compare Figs. 36 and 37 with Figs. 11-14.) The sclerotia,
while very similar to those of S. swdcata, are rather more slender but the
spermodochidia are very different. Since the spermodochidia are taxonomi-
cally the most distinctive structures of the different species of Sclerotinia in
this group, I have no hesitation in giving to this species a distinctive name.

It is perhaps remarkable that this species apparently has not been col-
lected by mycologists since the days of Desmaziéres. It is probably not
rare but will be found by one seeking it in the proper localities. I unfor-
tunately encountered Schoenus nigricans in but one locality in Europe.
That was in late summer on the fens at King’s Lynn, England. No trace
of the’ Sclerotinia was found.

This species is here presented, chiefly to bring it to the attention of
European mycologists in the hope that someone will be stimulated again to
discover this very interesting form and make a more complete study of its
different stages. Only in this way may it be determined whether I am cor-
rect in assuming it to be distinct from other species occurring on members
of the Cyperaceae. Our present scanty knowledge of this form is summar-
ized in the following technical description.

“Plantes Cryptogames France, Ed. 3, Ser. II, No. 21 (1853).

WHETZEL: SPECIES OF SCLEROTINIA 425

SCLEROTINIA SCHOENICOLA, sp. nov.

Maculis spermodochidialibus majusculis, ovatis vel circularibus, numerosis, magni-
tudine variabilibus, 2-6 mm. in diametro, sub inflorescentiis infectis irregulariter secun-
dum unum culmi latus dispositis, pallide olivaceo-brunneis coloris atroolivacei massae
spermodochidiorum globosorum lurideque brunnei cellularum contextus suscipientis
gratia; structura interna spermodochidiorum structurae S. scirpicolae subsimili; sclerotiis
gracilibus, fusiformibus, apicibus obtusis instructis, strictis, rarius manifeste curvatis vel
inaequilateralibus sclerotiorum S. sulcatae modo, initio pallide roseis, dein nigrescentibus
ad superficiem, intus albis, fortiter arato-sulcatis, 8-10x 1-2 mm. in statu sicco, per
rimam culmi liberatis; apotheciis ignotis.

Spermodochidial patches (Figs. 36, 37) rather large, ovate to circular,
numerous, variable in size, 2-6 mm. diam., distributed irregularly along
one side of the cylindrical culm, just below the blighted head, pale oliva-
ceous brown, due to the dark olivaceous color of the massed globose sper-
modochia and dead brown cells of the suscept tissues; internal structure
essentially like that of the spermodochidia of S. scérpicola.

Sclerotia (Fig. 38) slender, fusiform, tips blunt, straight, rarely distinctly
curved or inequilateral as are the sclerotia of S. sudcata, at first pale pink,
becoming black externally, white within, strongly furrowed, 8—10 mm. long
by 1-2 mm. thick (dry), discharged by a slit in the culm.

Apothecia unknown.

Hasitat: Culms of Schoenus nigricans L.

DistTRIBUTION: Apparently known only from France. There is nothing
on the labels of Desmaziéres’ specimens (the three cited above) to indicate
where, when, and by whom they were collected. All were presumably from
material sent him by Roberge.

TYPE SPECIMENS: Desm. PI. Crypt. France. Ed. 1, Ser. I, 2029 (in part)
(1850), and Ed. 2, Ser. II, 1629 (in part) (1853), (sclerotia only); Ed. 3,
Ser. II, 27 (in part) (1853) (spermodochidia only).

SPECIMENS EXAMINED: The specimens from Desmaziéres’ Plantes Cryptogames France
listed above were examined in the Royal Herbarium at Kew and in the British Museum
in London, in the New York Botanical Gardens, in the Farlow Herbarium, Cambridge,
Mass., and in the Mycological Collections, Bur. Pl. Ind., U.S.D.A.

The life history of S. schoenicola is presumably similar to that of the
other species of Sclerotinia attacking cyperaceous suscepts. Invasion is
evidently by way of the flowers. The appearance of the spermodochidia
might suggest a near relationship to S. scirpicola, while from the shape and
size of the sclerotia one would rather infer that S. schoenicola is more nearly
related to S. duriaeana or S. sulcata.

SPECIES ON THE JUNCACEAE

There are, as far as I have discovered, but three species of the true
Sclerotinias attacking members of the Juncaceae. These are S. curreyana,
very common in culms of Juncus effusus in Great Britain and western con-
tinental Europe; a species in our Pacific Northwest, described as Ciboria
juncigena; and an undescribed species on Luzula pilosa known only from
one collection taken in Saxony, Germany. -

426 Fartowla, Vor. 2, 1946

PLATE X

WHETZEL: SPECIES OF SCLEROTINIA 427

SCLEROTINIA CURREYANA
PLATE X AND Text Fic. D

Tulasne in 1861 in his Selecta Fungorum Carpologia (1: 104-105) de-
scribed accurately and in some detail two species of Sclerotinia in the group
under consideration in this paper, and in 1865 (3: pl. 22) presented beauti-
ful illustrations of both..

Thus S. curreyana and S. duriaeana may well be regarded as the classical
representatives of the species of Sclerotinia occurring on sedges and rushes.
Of these two, S. curreyana has long been the best known to the English and
European mycologists since the time of its study by Berkeley and Currey
in 1857, because of its very common occurrence and the fact that the
apothecia are readily found arising from the sclerotia of the still standing,
bleached culms. There is strong circumstantial evidence, however, that it
was really first described by Fries in 1818 under the name of Peziza cibo-
rioides. Rehm (1893: 822) appears to have been the first to question
whether Peziza ciborioides of Fries (1824), to which Persoon four years
later (1822: 277) gave the new name Peziza friesii, may not have been the
Peziza curreyana which Currey described in 1857. A careful consideration
of all the evidence leads me to believe that Fries actually had this species
on culms of Juncus in view when he named and described P. ciborioides.
His description fits the species well, inadequate though it is. The fact that
his fungus occurred in culms in early spring in marshy places is very sug-
gestive if not confirmatory evidence for my opinion. However, since Fries’
specimens are not preserved and since the fungus is so well known under
the name suggested by Berkeley, I am not using the Friesian name which
current rules of nomenclature might justify. I found this species very
abundant in the region about London in the early spring of 1930 and so
had an unusual opportunity to observe and study it. I have searched for it
diligently since that time on Juncus effusus in the Cayuga Lake region of
central New York, but have never found it, nor is it recorded from anywhere
in North America, unless S. juncigena should prove to be conspecific. It
should be found throughout the Euro-Asian continent wherever its suscepts
occur.

SCLEROTINIA CURREYANA (Berk. in Currey) Karsten, Revisio Monogr. p. 123.
1885.

Peziza curreyana Berk. in Currey, Journ. Linn. Soc. 1: 147. 1857.

Peziza curreyi Berk. Outlines Brit. Fung. p. 370. 1860.

Rutstroemia curreyana Karst. Myc. Fenn. 1: 107. 1871.

Phialea curreyana (Bart.) Gillet, Discom. p. 211. 1879.

Sphacelia tenella Sacc. Miscellanea Mycologica in Venezia Ist. Atti, Ser. 6, 2: 448.
(Reprint 1: 14) 1884. (See Saccardo 1886: 666.)

Plate 10. Sclerotinia curreyana. Fig. 41. Diseased culms of Juncus effusus showing
spermodochidia; about natural size. CUPP 31655. Fig. 42. Spermodochidia, x6. Fig.
43. Apothecia arising from sclerotia within standing culm; natural size. CUPP 15578.
Fig. 44. Apothecia in various stages of development from sclerotia in culms in moist
chamber; natural size. CUPP 20198. Fig. 45. Apothecia fully expanded; cums showing
rupture slits over enclosed sclerotia, x2. CUPP 20198.

‘428 FartowiaA, VoL. 2, 1946

Peziza juncifida Nylander, Pezizas Fenniae in Sallsk. Fauna et Fl. Fenn. Forhandl.
Notiser (n. s. 7) 10: 39. 1869.

Hymenocypha curreyana Phill. Manual Brit. Discom. p. 16. 1887.

Placosphaeria junci Bubak, Ann. Myc. 4: 113-114. 1906.

Sphacelia curreyana Grove, Journ. Bot. Brit. and Foreign 50: 46. 1912.

Myrioconium tenellum V. Hohnel, Mitteil. Bot. Inst. Tech. Hochsch. Wien. 3: 50.
1926.

Spermodochidia (Figs. 41, 42) numerous, scattered irregularly along and
about the upper part of the diseased culms, usually most numerous along
one-half of the circumference of the culm, minute, pustulate, ellipsoid,
brown, slightly erumpent when fresh and moist, opening by a slit in the
epidermis; spermatia minute, globose, 2—2.5 » in diameter, produced from
the tips of obclavate spermatophores fasciculately clustered about a cen-
trum to form globose spermodochia as in other species of this group, but
also forming an irregular hymenium lining the cavity.

Sclerotia one to several in each culm, short, stout, cylindric with blunt,
rounded ends, 4-15 mm. or more long, by 2—4 mm. in diameter, externally
black or dark brown, coarsely sulcate, internally white when mature, of a
rosv pink color during development, embedded in the pithy interior of the
culm, disclosing in section the inclusion here and there of remnants of
undigested pith cells. The structure of the rind and medulla is essentially
like that of the sclerotia of S. scirpicola. The sclerotia are not freed from
the culms at maturity but remain embedded, apothecia developing in the
spring from the surface exposed by a slit in the epidermis of the culm over
the sclerotium.

Apothecia (Figs. 43-45) two to several from each sclerotium, short-
stipitate, pale tan to dark avellaneous brown; cup thin, deep goblet-shaped
to infundibuliform, often umbilicate, 2-12 mm. or more in diameter, margin.
entire, recurved, smooth without, rugose-fluted within; s#zpe as long as the
diameter of the cup, equally cylindrical, flexuous, solid, glabrous or slightly
hairy to tomentose at the base, somewhat darker in color than the cup; asci
long, slender, clavate, apex rounded, 38 meas. gave 47.6—78 x 4—5.4 u, mode
65x 5.4 pw, av. 65.2 x 5 w; 8-spored, spores uniseriate; ascospores slender,
allantoid, ends rounded, 99 meas. gave 7.9-14.5 x 1.3-2.9 pw, mode 11 x 2.9
p, av. 11.5x 2.5 pw; paraphyses “slender, pale brown at the slightly thick-
ened apex”, according to Massee (1895: 282). I have found it impossible
satisfactorily to make out the characters of the paraphyses from dry
material.

In culture on potato dextrose agar this fungus makes a dense cottony,
felty growth of pinkish aerial mycelium with eventual development therein
of more or less globose sclerotia which are at first a bright pink, and finally
form a white to pink medulla that is enclosed in an olivaceous brown rind.
The olivaceous brown masses of sporodochia form about the margin of the
mycelial mat.

HasitaT: In culms of Juncus, most common in Juncus effusus L. but
recorded also on J. conglomeratus L., J. glaucus Ehrh., and J. filiformis L.

WHETZEL: SPECIES OF SCLEROTINIA 429

in wet locations, meadows, heaths, moors, and along ditches. The apothecia
appear in early spring, followed by infection of the culms in which the
spermatia and sclerotia appear during midsummer and early autumn..

DISTRIBUTION: This species appears to be most common in Great Britain
judging from my own observations and from the numerous collections in
herbaria. Known also from Belgium (Lambotte 1887: 303) and Bommer
and Rousseau (1884: 133), France (Boudier 1905: 273-4) and Saccardo
(1889: 199), Finland (Nylander 1869: 39), Denmark (Lind 1913: 109),
Bohemia (Bubak 1906: 113, spermodochidia). Velenovsky (1934: 223)
records this species in Bohemia on Juncus glaucus and J. piscinam, also in
heads of wheat. His description and his figures would indicate that he had
some species quite different from S. curreyana. The spores as he describes
and figures them are certainly not the spores of this species.

TYPE SPECIMEN: There are a number of collections of this species de-
posited (in 1881) in the Royal Herbarium at Kew of which at least three
packets are from Currey’s personal herbarium and collected on ‘“Paul’s
Cray Common, Near Chislehurst in Kent”, the locality given by Currey
(1857) as the place where, on the 23rd of April 1856, he first discovered
the apothecia of Peziza curreyana. The label on these specimens says “On
Juncus conglomeratus” and is dated ‘“‘“May, 1856”. These specimens would,
therefore, appear to constitute the type material. Massee (1895: 283) says
that he had examined the type specimen.

MatTeERIAL EXAMINED: In addition to Currey’s type specimens, I have carefully exam-
ined the other twelve specimens in the Kew Herbarium, all of which were apparently
collected in England from 1856 to 1884. The labels for most of them are very sketchy.
They appear to be mostly specimens collected by (or sent to) Currey and Berkeley. In
the herbarium of the British Museum there is a sheet and a half of packets of this
species, .all originally in the collection of William Phillips. One specimen labeled ‘“co-
nidia” is of special interest, as showing that Phillips recognized the spermatial stage of
S. curreyana. These all appear to have been collected in England.

In addition to the above, I have had for study the following: On Juncus effusus L:.:
CUPP 15578, Worcester, England, October 18, 1926, C. Rea (sclerotia from which I
developed apothecia the following April); CUPP 31648, Wisley, Surrey, England,
February 10, 1930, Whetzel, Green & Cotton (apothecia) ; —CUPP 31654, Goslar am
Harz, Germany, Harzerberger Thal, Whetzel & Westcott, April 22, 1930 (apothecia) ;
— CUPP 31655, near Downton in Hampshire, England, August 2, 1930, Whetzel &
Brown (spermodochidia) ; — CUPP 31657, Sandhill bog, Worcestershire, England, Au-
gust 9, 1930, Whetzel & Rea (spermodochidia and sclerotia). On Juncus conglome-
ratus L.: CUPP 31658, shore of Loch Katrine near Stronachlar Pier, Scotland, August.
3, 1930, Mrs. Whetzel & Miss Westcott (spermodochidia) ; — CUPP 31659, near Plow-
right’s old home, not far from King’s Lynn, England, Whetzel (spermodochidia) ; —
CUPP 20198, Phytopathological Field Station, Slough, Buckingham, England, Decem-
ber 5, 1931, Mr. and Mrs. E. W. Mason (sclerotia from which I developed apothecia in
April at Ithaca, New York) ;— CUPP 31662, near London, England, May 1934,
William Brown (sclerotia) ; — Otto Jaap, Fungi Selecti Exs. 754 a and bd. Prov. Bran-
denburg, Triglitz in Prignitz, O. Jaap. (a) apothecia, May 30, 1916. (b) spermodochidia,
“Placosphaeria junci”, Bubdk, November 3, 1915;—-CUPP 31656, near Downton,
Hampshire, England, August 2, 1930, Whetzel & William Brown (spermodochidia) ; —
Sydow, -Mycotheca germanica 2358, near Burgholdinghausen, Kreis Siegen, Westfalen, -
Germany, May 18, 1924, A. Ludwig (apothecia).

430 FarLow1A, VoL. 2, 1946

2%
Va toae
Sit

{t)
a

Text Fic. D

WHETZEL: SPECIES OF SCLEROTINIA 431

This species is also said to occur on Juncus glaucus Ehrh., on J. filiformis
L. and on J. communis E. May. Saccardo (1886: 666) described the sper-
matial stage under the name Sphacelia tenella from a collection on Juncus
glaucus collected near Rouen, France. Mr. Carleton Rea (in a letter April
24, 1927) says the sclerotia of S. curreyana are common in the culms of
this species in the vicinity of Worcester, England. Bubak (1906: 113)
described what is apparently the spermatial stage of our fungus under the
name Placosphaeria junci in culms of Juncus filiformis and Nylander (1869)
has described the apothecial stage under the name Peziza juncifida on culms
of Juncus (compressi?) but I have not seen material on any of these suscepts.

In numerous records of collections of this fungus, the species of Juncus
on which it occurred is not given. The evidence available seems to indicate
‘very clearly that S. curreyana is most frequently to be found on Juncus
effusus L. and Juncus conglomeratus L.

The erroneous identification of the sclerotial stage of S. curreyana with
that of S. scirpicola by Currey and Berkeley has resulted in the citation of
Sclerotium roseum in many synonymies of S. curreyana (See p. 413). In his
transfer of Peziza curreyana to the genus Phialea, Gillet writes “Bart.” for
the author of the species. This appears to be an error for ‘‘Berk.” The
inclusion of Peziza juncifida Nyl. and Placosphaeria junci Bubak in the
synonymy of this species might be questioned, since I have not been able
to see authentic specimens of either. Nylander’s description of P. juncifida,
while brief, appears to me to justify the conclusion that he had apothecia
of S. curreyana in hand.

There would appear to be little question of the identity of Bubak’s fungus
with the spermatial stage of S. curreyana, since his sketch of a section
through the fruit body is quite in character with the spermodochidium of
this Discomycete (See Text Fig. D.) Bubak records his fungus on Juncus
filiformis, a species on which this Discomycete is not otherwise known
to occur.

The slender curved ascospores of S. curreyana are strikingly different
from the ascospores of any of the other species in the genus. The structure
of the spermodochidium is also different from that of the others. The dark
color is due in part to the dilute brownish color of the central hyphae of
the spermodochidia and brown hyphae in the covering epidermal cells. It is,
however, largely the result of the dark brown discoloration of the crushed
thin-walled cells of the parenchyma surrounding the spermodochidial cavity
giving the appearance of a spermogonial wall. This “wall” is further sug-
gestive of a true hyphal wall, since resting upon it is a spermatophore
hymenium, lining the cavity (Text Fig. D). This hymenial lining presents
a picture very different from that in the spermodochidia of the species in
Carex suscepts, where it is entirely wanting. However, just as in the Carex-

Text Fig. D. Cross section through a spermodochidium of Sclerotinia curreyana about
x200. Note that the cavity is formed by extensive lysis of the parenchyma between
the vascular bundles with collapse and browning of the surrounding cells to form a
pseudo-spermogonial wall. Drawn by Carlos Garcés.

432 FartowlA, Vor. 2, 1946

invading species, S. curreyana also produces globose spermodochidia which
lie typically in the central cavity of the hymenium-lined spermodochidium.
These peculiarities might be taken to indicate that it is not closely related
to the other species in the group here under consideration, but in all its
other characters it conforms very closely. Its nearest relationship would
seem to be with S. scirpicola if one considers the similarity of the sclerotia
of these two species.

In culture on potato dextrose agar S. curreyana presents a picture strik-
ingly like that of S. sulcata, the chief difference being the pinker hue of the
mycelial mat of the former. Both form sclerotia on this medium and the
medulla of the sclerotium of S. sulcata is nearly as pink as that of S. cur-
revana at comparable stages in its development.

It is remarkable in the light of Tulasne’s recognition (1861: 104) of the
spermatial stage in S. duriaeana that the spermatial stage of S. curreyana
was not described until 1884 (Saccardo) and not recognized as such until
1912 (Grove). Grove says the spermatia measure 3—5 » in diameter. Pre-
sumably his measurements were made from fresh material. Mine (2-2.5 ,)
were made under oil immersion from dry material soaked over night in
weak KOH. This difference in size is, however, hard to explain on these
grounds.

SCLEROTINIA JUNCIGENA

In May 1883, Suksdorf collected the apothecial stage of a little Dis-
comycete apparently growing on the surface of ‘dead stems of Juncus above
the water in Falcon Valley”, according to a note on the packets distributed
by him in his exsiccati, Flora of Washington. Ellis, who determined the
material, gave it the name Ciboria juncigena E. & E. Dr. J. R: Keinholz
advises me in a letter of June 7, 1942 that the name “Falcon Valley” ap-
parently was abandoned many years ago and the name changed to ‘Hell
Roaring Meadows”, which lies at the head of a creek of the same name. No
other collections of this form have been preserved as far as I can discover,
although Dr. S. M. Zeller of the Oregon Experiment Station writes me that
he has once come upon this fungus in the field. The species of Juncus is not
given by Suksdorf but it is probably J. effusus var. pacificus Fern. & Wie-
gand. Dr. Keinholz who kindly undertook to locate fresh material of this
fungus has thus far been unsuccessful. Its close resemblance to S. curreyana _
of Europe makes a critical study of living specimens highly desirable. Pend-
ing such a study it seems wisest to retain its present specific standing.

SCLEROTINIA JUNCIGENA (E. & E.) n. comb.
Ciboria juncigena E. & E., Proc. Acad. Nat. Sci. Phila., Nov. 30, 1894, p. 348.

Spermodochidia not seen.

Sclerotia one, possibly more, within a diseased culm, slender, cylindric,
_ with truncate, slightly rounded ends, up to 15 mm. long by 2 mm. in diam-
eter, externally black, finely sulcate, internally white when mature; appar-
ently lying free in a lysigenous cavity in the pith region of the culm; struc-
ture essentially like that of S. curreyana. The sclerotia are not freed from

WHETZEL: SPECIES OF SCLEROTINIA 433

the culm at maturity, nor are they apparently exposed by a slit on emer-
gence of the apothecia as in S. curreyana.

Apothecia usually but one from each sclerotium, long-stipitate, avellane-
ous (?); cup thin, membranous, shallow cup-shaped, sub-umbilicate, 4—5
mm. in diameter; Aymenium “wine colored”; stipe arising from the scle-
rotium within and penetrating the wall of the culm, relatively long, about
1 cm. by 1 mm. thick, lower half or more black, longitudinally wrinkled,
fibrillose, anchored at point of emergence to the surface of the culm by a
spreading dark hyphal mat, concolorous above with the cup; a@sci clavate-
cylindrical, (Ellis meas.) about 60 x 4-5 p, 8-spored, spores sub-biseriate;
ascospores allantoid, slender, distinctly curved, hyaline, (Ellis meas.)
7-8 x 1.25 yw, (Whetzel, 50 meas. in KOH) 7.2—10.8 x 1.2-2.4 yw, mode
8.4 x 2.4 p, average 8.86 x 1.83 1; paraphyses not distinguishable in mounts
from dried material.

HasitaT: In culms of Juncus (probably J. effusus L. var. californicus
Fern. & Wiegand) in wet marshes. The apothecia appear in May on stand-
ing dead culms.

DISTRIBUTION: Known only from the type locality. Falcon Valley (now
Hell Roaring Meadows), Washington State.

TYPE SPECIMEN: Exsiccati specimens distributed as 371 Flora of Wash-
ington. Collected by W. N. Suksdorf, May 31, 1883. I have seen two
packets, one deposited from the Ellis Collection in the N. Y. Bot. Gard.
herbarium and another in the Mycol. Coll. Bur. Pl. Ind., Washington, D. C.

It is with some hesitation that I treat Ciboria juncigena E. & E. as a
species distinct from S. curreyana. It is undoubtedly a Sclerotinia rather
than a Ciboria. Ellis entirely overlooked the presence of the sclerotia,
which was perhaps due to the absence of a slit in the culm over the scle-
rotium and the unusual anchorage of the stipe to the surface of the culm.
As far as the characters of the cup are concerned, it could hardly be dis-
tinguished from S. curreyana, but the long, slender stipe, its black color,
and the basal, dark hyphal mat anchoring it to the culm at the point of
emergence, together with the slender character of the sclerotium, may well
prove to be dependable evidence of its specific character. It is unquestionably
very closely related to S. curreyana and may indeed be more properly re-
garded as an American form of variety of that species. However, my experi-
ence with the closely related S. duriaeana and S. sulcata, together with the
fact of its long geographical isolation from the ancestral S. curreyana, in-
clines me to believe that a fuller study of S. juncigena in fresh condition
and in its spermatial stage will prove it to be a distinct species.

SPECIES ON LUZULA

SCLEROTINIA LUZULAE
PuaTE IX, Fics. 39 AND 40

W. Krieger distributed in his Fungi Saxonici 2073 culms of Luzula pilosa
L. in which are embedded slender black sclerotia of a fungus which he

434 FarLowlA, VoL. 2, 1946

labeled provisionally “?Sclerotinia Curreyana (Berk.) Karst.” The label
bears a brief description of the sclerotia with the remark that unfortunately
he had been unable to develop the fruit body of the Ascomycete, hence the
question mark before the name he was giving the specimen. He apparently
collected the specimens he distributed under this number in two different
localities and in different years, in Polenztale, Utterwaldergrunde, late May
1896, and near Konigstein, June 1897. In Rehm’s collections at Stockholm,
Sweden, there is in addition to Krieger’s exsiccati specimen 2073, another
packet labeled in Krieger’s handwriting bearing the same data, evidently
duplicate material.

I have been able to locate but one other collection which is in the her-
barium of the Mycological Collections, Bur. Pl. Ind., U. S. D. A., Washing-
ton, D. C. This bears the label ‘‘Herbarium G. Bresadola, 221 Sclerotium
roseum Moug. en feo oritur Scler. Curreyana Berk. Sclerotium Luzulae
Krieg. n. sp.? In den diirren Halmen von Luzula pilosa Willd. bei Kénig-
stein, 19, Juli, 1897. Lg. W. Krieger”. The name “Sclerotium luzulae
Krieg.” appears never to have been published. The specimen is apparently
a part of the 1897 collection distributed by Krieger in his Fungi Saxonici
2073. Krieger’s collections of a sclerotial form on Luzula pilosa would seem,
for the present, best regarded as a distinct species.

SCLEROTINIA LUZULAE sp. nov.

Spermodochidiis apparenter nullis; sclerotiis 1-3 vel pluribus, intra culmos natis et
maturitate ad superficiem soli ruptionis culmorum causa liberatis, 5-30x 1 mm., fusi-
formibus, acute abrupteque acuminatis unum ad latus, aqua imbutis obscure castaneis,
siccis nigris, indistincte arato-sulcatis, cortice uno e strato cellularum atro-tunictarum
consistente, medulla alba, ex hyphis dense compactis, multo minoribus diametro ordi-
nario specierum stirpis huius composita; apotheciis ignotis.

Spermodochidia apparently wanting.

Sclerotia one to three or more formed within the culms which are ruptured
at maturity, discharging the sclerotia onto the soil, 5-30 mm. long by 1 mm.
in diameter, fusiform, sharply and abruptly pointed at either end, dark
mahogany brown when soaked in water, black when dry, indistinctly fur-
rowed, not sulcate as in other species in this group, rind one layer of dark-
walled cells, medulla white, composed of densely packed hyphae which are
much smaller in diameter than those of the sclerotia of other species in
this group.

A pothecia unknown.

Hasitat: In culms of Lugula pilosa Willd. in Saxony, Germany.

DIstTRIBUTION: Known only from the type locality and very rare, accord-
ing to the collector.

TYPE SPECIMEN: Krieger’s Fungi Saxonici 2073, in culms of Luzula pilosa
Willd. in Polenztale, Utterwaldergrunde, May 1896 and near Konigstein,
June 1897, W. Krieger.

MATERIAL EXAMINED: I have carefully examined the specimens of the type collections

in the British Museum and in the CUPP Herbarium; also the specimen from the
herbarium of the Bur. Pl. Ind., U.S.D.A.

WHETZEL: SPECIES OF SCLEROTINIA 435

Krieger, on the label of his Fungi Saxonici 2073, refers the specimen
provisionally to Sclerotinia curreyana and cites Sclerotium roseum Fr. as a
synonym, obviously two errors. This species is probably more common
than Krieger surmises and should be sought wherever the suscept occurs.

The apparently complete absence of spermatial fruit bodies is puzzling.
I have scrutinized carefully all the culms which I find in Krieger’s collec-
tions but without finding any trace of spermodochidia. Their absence is
most unusual, this being the only species in this group, except S. vahliana
and S. juncigena, in which I have not found them. That I should have
found them in these two species is almost certain had I had specimens of
diseased culms collected during the summer.

EXCLUDED SPECIES

Several species of stromatic Discomycetes other than those described
above occurring on species of the Cyperaceae and Juncaceae have been put
in the genus Sclerotinia by various authors. None of these species with
which I am acquainted are, however, congeneric with the true species of
Sclerotinia as I interpret that genus. I list them here with some observations
and comments which may prove useful to my mycological colleagues until
such time as I may treat them more fully.

Sclerotinia aschersoniana P. Henn. & Pléttn. Verh. Bot. Vereins, Prov. Branden-
burg 41: IX, 1899.

This species is pathogenic in the ovaries of certain species of Carex,
notably in C. kudsonii A. Bennet (=C. stricta Good.) in Europe and on
C. stricta Lam. in North America. I have collected and cultured it fre-
quently from the latter species in swamps in central New York and once
from C. prairea Dewey in the same region. It is certainly not a Sclerotinia.
It infects the suscept by ascospore invasion through the stigma, converting
the ovary into a black stromatized mummy. It might be referred to the
genus Ciboria but its proper taxonomic relationships require further con-
sideration.

Sclerotinia utriculorum Boud. Bull. Soc. Myc. France 19: 196-197, pl. 8, fig. 6. 1903.

This is described as growing on the hard seeds of Carex davalliana in
May, in the Jura in France. Boudier says it differs from other species of
Sclerotinia on Carex in that the apothecia do not arise from sclerotia but
from the sclerified achenes of the Carex. As he makes no reference to
S. aschersoniana it is probable he was unaware of that species. As far as I
can judge, S. utriculorum Boud. is specifically the same as S. aschersoniana
P. Henn. & Plottn. Velenovsky (1934: 224) apparently regards this species
as identical with S. aschersoniana, citing the latter as a synonym of S. wféri-
culorum, but his reason for this is not clear.

436 FarLowl1A, VoL. 2, 1946

Sclerotinia vesicariae Giesenhagen, Ber. Bayer. Gesell. Erforsch. der Heim. Flora 11:
167-169. 1907.

This species is said to occur among (‘“inter’’) fallen dead seed of Carex
vesicariae in autumn, collected by Stechsee near Seehaupt, Bavaria. The
sclerotia are said to be irregularly cylindrical, rough, black, white within,
5-8 mm. long by 2 mm. in diameter. This would seem to indicate that this
species does not occur in the ovaries of the Carex, but there is little else to
indicate that it is distinct from S. aschersoniana. The asci are said to be
180-190 yu, considerably longer than those of S. aschersoniana which are
given as 90-110 y» in the original description; the width is about the same.
Ascus measurements, however, are so very variable in the stromatic Dis-
comycetes, especially the length, as to be of little specific significance. The
size and shape of the spores approximates those of S. aschersoniana. I am
inclined to regard S. vesicariae as specifically distinct.

Sclerotinia caricina Velenovsky, Monog. Discom. Bohemiae 1: 224; 2: pl. 22, fig. 40.
1934,
This species is said to occur on the roots of Carex muricata among Juncus
communis. It is not possible to tell from Velenovsky’s description and
figure whether this is a true Sclerotinia or not.

Sclerotinia paludosa Davidson & Cash, Mycologia 25: 271. 1933.

A critical examination of all the specimens on which this species is based,
discloses that the sclerotia shown in their fig. 7 are those of a Typhula. The
apothecia in the other specimens do not arise from sclerotia but from
stromatized leaf tissues. This fungus is undoubtedly a Rutstroemia. It is
a common species of wide distribution occurring on leaves and culms of:
various species of sedges, grasses, etc. The proper species name remains to
be determined. It is probably conspecific with the following species.

Sclerotinia heterocarpa Bennett, Ann. Appl. Biol. 24: 254. 1937.

While this species is described as occurring on grasses as the cause of
the “dollar-spot” of turf, what is without doubt the same species has been
discovered by the writer on leaves of Carex and several other plants. It was
earlier described under the name Ciboria armeriae by Von Hohnel (Frag.
Myk. 22, Mitt. No. 1122, p. 43. 1918). This fungus is clearly not a Scle-
rotinia. It is without doubt a species of Rutstroemia.

CoRNELL eee are
Irnaca, N, Y.
LITERATURE CITED

References cited in the text but not in the following list will be found in the
synonymy of the species referred to.

Bommer, E. & M. Rousseau. P. curreiana Tul. In Florule mycologique des Environs
de Bruxelles. Mem. Soc. Roy. Bot. Belg. 23: 133. 1884.

Boudier, E. Icones Mycologicae 1-4. 1905-1910.

Drayton, F. L. The sexual mechanism of Sclerotinia gladioli. Mycologia 26: 48-72.
1934.

WHETZEL: SPECIES OF SCLEROTINIA 437

Ferdinandsen, C. & O. Winge. Studier over en hidtil upaaagtet, almindelig dansk
Baegersvamp, Sclerotinia scirpicola Rehm. Biol. Arbejder Tilegnede, Eug.-Warm-
ing den 3. November 1911: 281-298. 1911.
. Uber Myrioconium Scirpi Syd. Ann. Myc. 11: 21-24. 1913.
Fries, E. Peziza ciborioides. In Observationes Mycologicae 2: 307. 1824.
Groh, Herbert. A new host for Claviceps. Mycologia 3: 37-38. 1911.
Hohnel, F. von, Uber Epidocium affine Desmaziéres. Mitt. Bot. Inst. u. Tech. Hochsch.
Wien 3: 50-51. 1926a.
. Uber Placosphaeria Junci Bubak. Mitt. Bot. Inst. u. Tech. Hochsch. Wien
3: 49-50. 1926b. é
Lambotte, E. Scler[otinia] curreiana Tul. In Fl. Myc. Belg. Suppl. 1, p. (303). 1887.
Larsen, P. 324 S. Vahliana Rostrup, Gronl. Svampe 1891, p. 607. In Fungi of Iceland.
In Rosenvinge & Warming, The Botany of Iceland 2°: 504. 1932.
Lind, J. Sclerotinia Curreyana (Berk.) Karsten. Jn Danish Fungi as represented in the
Herbarium of E. Rostrup, p. 108-109. 1913.
Massee, G. Sclerotinia Curreyana. In British Fungus-Flora 4: 282-283. 1895.
Persoon, C. H. (Peziza) Friesii. In Mycologia Europaea 1: 277. 1822.
Rostrup, E. Sclerotinia Vahliana Rostr. In Islands Svampe. Bot. Tidskr. 25°: 315.
1903.
. Fungi collected by H. G. Simmons on the 2nd Norwegian Polar Expedition,
1898-1902. In Rept. Second Norwegian Arctic Exped. in the “Fram” 1898, 9: 5.
1906. %
Saccardo, P. Sclerotium nigricans (Tul.) Sacc. In Syll. Fung. 14: 1153. 1899.
——. Sclerotinia Curreyana (Berk.) Karst. Zn Syll. Fung. 8: 198-199. 1889.
Tulasne, L. R. & C. Tulasne. Selecta Fungorum Carpologia 1: 103-105. 1861; 3: 203,
pl. 22. 1865. (Transl. by W. B. Grove 1: 106-108. 1931; 3: 202, pl. 22. 1931).
Velenovsky, J. Monographia Discomycetum Bohemiae 1: 1-436; 2: pl. 1-31. 1934.
Whetzel, H. H. North American species of Sclerotinia II. Two species on Carex, S.
duriaeana (Tul.) Rehm, and S. longisc!erotialis n. sp. Mycologia 21: 5-32. 1929.
. The spermodochidium, an unusual type of spermatial fruit-body in the
Ascomycetes. Mycologia 35: 335-338. 1943.
. Saccardo’s confusion of the spermatial stage of S. duriaeana and S. cur- |
reyana with the sphacelia stage of Claviceps nigricans. Mycologia 36: 426-428. 1944.
Wheizel, H. H. & W. G. Solheim. Sclerotinia caricis-ampullaceae, a remarkable sub-
arctic species. Mycologia 35: 385-398. 1943.

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CONTENTS OF VOL. 2, NO. 4
THE Genus GrirFiTHsiA (Rhodophyceae) im Hawa. By Isabella A. Abbott . . 439
Func DomincEnses Novi vet Minus Cocniti. I. By Carlos E. Chardon . . . 455
THE BIOLOGY OF GYMNOSPORANGIUM Nipus-AvIs THAXTER. By Alton E. Prince . 475
THE BOLETINEAE OF FLorIDA witH Notes oN EXTRALIMITAL Species IV. THE

LAMELLATE Famities (Gomphidiaceae, Paxillaceae, and Jugasporaceae). By
BOE DIME ER cn he PRR ow, (oes og GS ere eG mai mer cg RY

Pees he Vola So en) i ees Cee 2 io: | Sa ak ae oe ae

os aS ee On | ee

Vol. 2, No. 3 was issued on January 16, 1946.

FARLOWIA

A JOURNAL OF CRYPTOGAMIC BOTANY

Vel 2a Jun, +1946°° << No. 4°

THE GENUS GRIFFITHSIA (RHODOPHYCEAE) IN HAWAII

IsABELLA A. ABBOTT

The small, inconspicuous plants of the genus Griffithsia (Ceramiaceae,
Rhodophyceae) are usually missed in general collections of algae. For the
most part they are microscopic as well as epiphytic and occur well-mixed
with other algae. In the subtropical waters of the Hawaiian Islands,
members of this genus are usually found in reef and rocky shore habitats.
They do not occur in abundance.

Griffithsia is fairly well known from the Atlantic and Indian Oceans,
Caribbean and China Seas, and little known from the Pacific. The first
record of the genus in the Hawaiian group was made by Reed in 1907,
with reference to the economic use of the plants by the native Hawaiians.
The algae are one of nearly seventy which were used to supplement the
diet of the Hawaiians. Some of the more common native names of mem-
bers of this genus are limu (literally translated as edible algae) moopuna-
a-ka-lipoa, and limu aupuu. The native names of this and other algae
vary from island to island, and sometimes on a single island.

Reinbold (1907) has listed Griffithsia thyrsigera ( = G. tenuis) from
Hawaii. MacCaughey (1917) and Neal (1930) have mentioned Griffithsca
ovalis Harvey as possibly being present in the Hawaiian Islands.

Five species of Griffithsia are recorded here from the Hawaiian group.
Four are listed for the first time from the area, and the presence of G.
ovalis is confirmed.

New extensions of ranges are listed for Griffithsia Binderiana Sonder,
an Australian species, G. tenuis C. Agardh, a cosmopolitan species, G. Met-
calfii Tseng, reported from China, G. ovalis Harvey from Australia, and
G. rhizophora Grunow from Ceylon, China, and the Malay Archipelago.

Cystocarps of Griffithsia tenuis and G. Binderiana are studied for the
first time.

The following key is based on characters which are present for the
greater part of the life history of the plants in the Hawaiian Islands.
Cystocarpic and spermatangial plants which furnish the best differences
among the species are infrequently found.

KEY TO THE SPECIES

A. Tetraspores between the articulations of the vegetative cells
B. Involucre surrounding tetraspores arising from vegetative cells; cells micro-
scopic, subcylindrical below, globose at tips ........ 1. G. ovalis.

439

440 FarRLowIA, VoL. 2, 1946

B. Involucre attached to tetraspore pedicel; vegetative cells macroscopic and
moniliform throughout ...... Ale REE rani: eee cl 2. G. Metcalfii.
A. Tetraspores borne on specialized branches, or at the tips of the filaments
C. Cells cylindrical, slender; all reproductive structures formed on lateral pedi-
Clee wehbe eee aes iG Soon a ey ye 3. G. tenuis.
C. Cells subeylindrical or globose
D. Reproductive structures in lateral, specialized branches, all involucrate

Pcrano ety pene oseer eee te cea ia carci eG hy acetone eon th Syn 4. G. Binderiana.
D. Tetraspores (other plants not seen) at the tips of globose cells; no in-
WiC NOP GN. oi gis bees: ose ok bo Sak 5. G. rhizophora.

1. Griffithsia ovalis Harvey, yew Australica 4:203. 1862.
Prater I, rics. 1-4; Prare II, rics. 1-2

Plants in well-formed tufts to 2 cm. in height, or creeping on other algae
and having erect filaments. Cells at the tips are oval to globular, 50-100 u
in diameter. Cells directly below are moniliform to subspherical, and at
the base are larger and subcylindrical to ellipsoidal, 500-700 p» in diam-
eter. Branches are occasionally fastigiate but predominantly dichotomous,
with the tips slightly convergent. Rhizoids are commonly found through-
out the plants which creep.

Tetrasporangia are borne between the articulations of the cells toward
the tip, and are pedicellate with four or more sporangia on a pedicel. The
mature tetrasporangia are about 40 » in diameter. A slightly incurved
involucre of 12-16 cells surrounds the tetraspores. No male or female
plants have been collected.

Type locality: King George’s Sound, Western Australia.

Distribution: Recorded previously only from Australia, and here re-
corded from the Hawaiian Islands.

Hawaiian specimens examined: Oahu Island: Kawela Bay, Abbott 770b, sterile on
Mesotrema; Waimanalo, north of Makapuu Point, Abbott 911a, sterile on Laurencia;
Diamond Head, below lighthouse, Abbott 899a, tetrasporic on Halimeda, Abbott 900,
sterile on Dictyosphaeria; Waikiki, near marine laboratory, Abbott 396, 474, 492, 501
on Sargassum, Abbott 502, 504, all tetrasporic.

The plants seem to be members of the Griffithsia rhizophora-Schousboei-
coacta complex, and although they have many vegetative characteristics
in common, studies of the male and female plants would be greatly desired.

Except for the small size of G. ovalis, the Hawaiian specimens also seem
related to Griffithsia corallinoides (L.) Batters (G. corallina (Lightf.)
C. Ag.). The involucrate tetrasporangial groups at the articulations are
characteristic of both species.

The specimens have been collected on volcanic rock as well as on coral
where they are much entangled with other small algae, chiefly Centroceras
clavulatum (C. Ag.) Montagne and Falkenbergia rufalanosa Harvey. Al-
though these collections have been made at different times of the year,
only tetrasporic plants have been found.

2. Griffithsia Metcalfii Tseng, Papers Mich. Acad. Sci. Arts and Letters, 27:111-116.

1942.
Prate II, rics. 3-6

Apsott: GRIFFITHSIA (RHODOPHYCEAE) IN HAwalrl. 441

Plant caespitose, to 5 cm. in height, the fronds free at the outer ends,
- subdichotomous in the upper parts of the filaments. Cells are conspicu-
ously moniliform, spherical, 500-700 » in the terminal cells, largest in
diameter through the center of the filaments, and subspherical to sub-
cylindrical basally. Attachment is by large, stout, basal rhizoids. Tenacu-
lar cells often connect branches near the base of the plant with each other.
Hair clusters are present in young specimens, and are quickly deciduous.

Spermatangial clusters are crowded between fertile vegetative cells below
the terminal ends of the branches, and are not involucrate. Tetrasporangia
bear a few involucre cells on the common pedicel. No cystocarpic speci-
mens were collected. ;

Type specimen: C. K. Tseng 938 in the herbarium of Tseng, from
Yenkehai, Hainan Island.

Hawaiian specimens examined: Oahu Island: Laie Bay at Makahoa Point, Abbott
25, tetrasporic, Abbott 662, spermatangial; Richard Cowan, 3 specimens from Nana-
kuli, tetrasporic; Edna Fassoth, from Waimanalo, sterile; Reed, without number, date,
or locality, sterile.

This species compares very favorably with the type specimen. It is
usually found with corallines in wave-swept areas about the Island of
Oahu. The familiar Griffithsia globulifera Harvey of the Atlantic is re-
lated vegetatively to this species, but differs in branching and location of
spermatangia. Griffithsia rhizophora Grunow of the Malayasian-Chinese
coasts resembles G. Metcalfii, but is a smaller plant and has branches
consistently convergent at the tips.

8 Griffithsia tenuis C. Agardh, Species Algarum 2(1):131. 1828.
Griffithsia thyrsigera (Thwaites) Askenasy, Algae, in Forschungsreise S.M.S.Ga-

zelle 4:36, pl. 9, figs. 1-4. 1889.
Griffithsia sp. Neal, Hawaiian Marine Algae, Bishop Museum Bull. 67:73-74.

1930.
Prate III, Fics. 1-7

Plant in freely branching loose tufts to 5 cm. in height, or prostrate and
creeping on other algae. The multinucleate filaments are irregularly
dichotomous, narrow in aspect, and non-forcipate. In the Hawaiian speci-
mens, subsecund branches rarely occur and thus this character, so promi-
nent in specimens from the Caribbean and China areas, is not a good
one here. Cells of the filaments are cylindrical to subcylindrical, 60-80 »
in diameter, becoming smaller at the tips where they are surrounded by
two or more clusters of hairs, the lower one long and six or more times
dichotomous, the upper one short and quickly deciduous.

The asexual plants are usually found to be separate from the sexual
plants, but in one collection tetraspores are found on the same plant as
the cystocarps (Abbott 472). Tetraspores occur in whorls of 4-12 at the
nodes of the upper filaments, each on a clavate or pyriform pedicel.
Deciduous involucral cells infrequently surround the tetraspores of the
Hawaiian specimens.

442 FARLOWIA, VoL. 2, 1946

Cystocarps are lateral on a one-celled pedicel, surrounded by an in-
volucre of 12-16 cells. The auxiliary cell of the carpogonium is cut off
by the supporting cell as reported for G. corallina by Kylin (1916), and
G. Metcalfii by Tseng (1942). Carpospores are formed by the repeated
division of the auxiliary cell on its outer face. They are pyriform and are
massed. Spermatangia are stalked, subcapitate, subglobose, without in-
volucre, on lateral specialized branches.

Type locality: Venice.

Distribution: Widely distributed in warm to subtropical waters; in
the Pacific area at points on the China coast, Fiji, and at Tutuila Island,
Samoa. .

Hawaiian specimens examined: Oahu Island: Laie Bay, Abbott 485, sterile;
Abbott 486, tetrasporic; Waimanalo, 3 miles north of Makapuu Point, Abbott 907a on
Dictyota, tetrasporic; Halona, Abbott 300, on Microdictyon Setchellianum, tetrasporic ;
Hanauma Bay, Abbott 50, tetrasporic, spermatangial; Wailupe, east of fishpond,
Abbott 808a on Dictyota, sterile; Kaalawai, Abbott 642, on Sargassum echinocarpum,
tetrasporic; Waikiki near marine laboratory, Abbott 472, tetrasporic and cystocarpic;
Abbott 491, spermatangial, cystocarpic, tetrasporic; Neal 23, carpogonial, tetrasporic;
Diamond Head, below lighthouse, Abbott 839, sterile, Abbott 896a on Dictyota, sterile,
Abbott 897a on Liagora, tetrasporic; G. P. Andrews without collecting data, tetra-
sporic; Kawela Bay, Abbott 732b on Padina, sterile. Maui Island: EF. Bailey 7, 5
specimens of which two are tetrasporic, others sterile.

The persistent dichotomous branching of the Hawaiian specimens is a
very misleading character since Griffithsia tenuis of other writers is taxo-
nomically characterized by having lateral, secund branches. One collection
of fifteen sets shows the secund habit of the species. These plants were
epiphytic on a small Sargassum echinocarpum, and on Champia sp. The
Hawaiian specimens appear to be more narrow in the size of the cells
and have smaller tetraspores than the Atlantic and China specimens. The
specimens collected at Hanauma Bay, in addition, show only four tetra-
spores in a whorl at the nodes and the pedicels are not typically clavate.
or pyriform as in the Waikiki specimens. These differences, however, may
perhaps be attributed to the tide pool habitat of the former.

The appearance of tetraspores and cystocarps on the same plant is not
rare for the Ceramiales (see Kylin, 1928, Drew, 1944), but is newly
reported for G. tenuis. Unfortunately, I have not studied the species
cytologically.

4. Griffithsia Binderiana Sonder, Bot. Zeit. 3:52, 1845.
Bornetia Binderiana (Sond.) Zanard., Iconogr. Phycol. Med.-Adr. 2:45. 1865
(cited by Bgrgeson, F. Danske Vidensk. Selsk., Biol. Medd. 19(10):16. 1945).
(See other references for this species by Sonder in literature cited.)

Pirate IV, Fics. 1-7

Plants tufted, to 4 cm. in height, much branched, subdistichous, dicho-
tomous, furcate at the tips. Cells are subcylindrical, 300-500 « in width,
2.5 to 3 times as long in the central parts of the filaments, cells be-
coming gradually smaller toward the terminal ends. Tetraspores are

ApBotT: GRIFFITHSIA (RHODOPHYCEAE) IN Hawall 443

borne on characteristic cuneate or ovoid, solitary, secund pedicels; the
tetraspores are surrounded by a prominent incurved involucre. Tetra-
spores are 50-60 » in diameter. Three or more tetrasporangia are clustered
on a common stalk. Rhizoids occur chiefly in the basal parts, but in some
ecological forms they may occasionally appear laterally throughout the
plant. .

Type locality: “ad litus occidentale Novae Holland.”

Hawaiian specimens examined: Oahu Island: near Elk’s club, Waikiki, Abbott 467,
sterile; Laie Bay at Makahoa Point, Abbott 468, tetrasporic; Abbott 659, spermatan-
gial; Abbott 660, cystocarpic; Abbott 661, tetrasporic; J. T. Conover at Barber’s
Point, tetrasporic; R. S. Cowan at Nimitz Recreation Beach, south Ewa, near Bar-
ber’s Point, tetrasporic; J. F. Rock from Honolulu Harbor, sterile. Maui Island:
Paia, E. Bailey without number, some tetrasporic, others sterile.

The prominent lateral branches bearing involucrate tetraspores are
most characteristic of this species, and fortunately the cystocarpic and
spermatangial plants also show this characteristic well. The development
of the cystocarp follows that of G. corallina, G. Metcalf and G. tenuis.
Spermatangial and cystocarpic plants have been found in the winter
months, the tetrasporic plants throughout the year. Spermatangial and
cystocarpic plants are reported for the first time.

Griffithsia Binderiana seems to be related to the oriental G. subcylindrica
Okamura, especially in the vegetative structure.. The cells of the latter
appear to be larger both in width and length. The Hawaiian plants com-
pare favorably with Harvey’s description of specimens of G. Binderiana
from Rottnest.

5. Griffithsia rhizophora Grunow ex Weber-van Bosse, Siboga Expeditie. Mono-
graph 59¢:313. 1923.

Puate I, Fics. 5-9

Frond to 2 cm. in height, caespitose. Branches are convergent at the
tips, otherwise subdichotomous and erect, or creeping. The cells are
moniliform except at the base where they are subcylindrical. The central
cells of the filaments are 400-500 u in diameter with the same length.
Terminal cells are somewhat less than 100 yp, basal cells 500 p» in diameter
and twice as long. Tetraspores without involucre.

Type specimen: Ferguson 316, in Algae of Ceylon. -
Distribution: Ceylon, Hainan, Malaya.

Hawaiian specimens examined: Oahu Island: Laie Bay, Abbott 466, tetrasporic;
Waimanalo, 3 miles north of Makapuu Point, Abbott 909a on Dictyota, sterile ;
Abbott 910a on Padina, sterile; Wailupe, east of fishpond, Abbott 802a on Dictyota,
sterile; Diamond Head, below lighthouse, Abbott 816a on Sargassum, sterile; Kaal-
awai, Abbott 663a on Padina, sterile; Kawela Bay, Abbott 715a on Halimeda, sterile.

The regular subdichotomous branching and the convergent tips of
this species separate it vegetatively from Griffithsia Metcalfii. The latter
plant is also larger. Of the Hawaiian species of Griffithsia studied, this

444 FarLowIA, VOL. 2, 1946

one is least known since only one fertile specimen was collected. The
identification, therefore, should be considered to be tentative.

Griffithsia rhizophora was named by Grunow on a herbarium sheet
(nomen nudum). Later, Madame Weber-van Bosse published it in the
list of Siboga Algae, crediting it to Grunow. Article 48, section 7 of the
International Rules states the “. . . name of the latter author must be
appended to the citation with the connecting word ex.” The listing of the
species should be as above.

The writer wishes to thank Dr. George F. Papenfuss of the University
of California who has helped greatly in checking literature she was not
able to see.

UNIVERSITY OF HAWarI
Hono tutu, T. H.

LITERATURE CITED

Agardh, C. A. Species Algarum. 2(1): 131. 1828.

Askenasy, Eugen. Algae in Forschungsreise S.M.S.Gazelle . . . 1874—bis 1876. 4: 36.
pl. 9, figs. 1-4. 1889.

Bérgesen, F. Marine algae of the Danish West Indies II. Dansk Bot. Arkiv. 3:
462-464. 1915-20.

Drew, K. M. Nuclear and somatic phases in the Florideae. Biol. Rev. Camb. Phil.
Soc. 19: 113-117. 1944.

Harvey, W. H. Phycologia Australica 4: 203. 1862.

Kylin, H. Die Entwicklungsgeschichte von Griffithsia corallina (Lightf.) Ag. Zeit. f.
Bot. 8: 99. 1916.

- Entwicklungsgeschichtliche Florideenstudien. Lunds Univ. Arsskrift,
N. F., 2: 24. 1928.

MacCaughey, Vaughan. The algae of the Hawaiian Archipelago. (Thrum’s) Ha-
waiian Almanac and Annual for 1918: 154. 1917.

Neal, Marie C. Hawaiian marine algae. Bishop Museum Bull. 67: 73-74. 1930.

Reed, Minnie. The economic seaweeds of Hawaii and their food value. Hawaii
Agr. Exp. Sta. Ann. Rept. for 1906: 87. 1907.

Reinbold, Th. Meeresalgen im Rechinger, K. Bot. und Zool. Ergebnisse einer wissen-
schaftlichen Forschungsreise nach den Samoainseln, dem Neuguinea-Archipel und
den Salomonsinseln. 1: 10. 1907.

Setchell, W. A. American Samoa: Pt. 1. Vegetation of Tutuila Island. Carnegie
Inst. Wash. 20: 159-160. 1924.

Sonder, O. G. Nova Algarum genera et species quas in itinere ad oras occidentalis
Novae Hollandiae collegit L. Preiss, Ph. Dr. Bot. Zeit. 3: 52. 1845.

in Lehmann, Johann Georg C. Plantae Preissianae, sive enumeratio
Plantarum quas in Australasia occidentali et meridionali-occidentali annis 1838—41
collegit L. Preiss. 2: 168. 1846-47.
. m Plantae Preissianae (reprint with different pagination, referred to by
J. Agardh) p. 21.

Tseng, C. K. Studies on Chinese species of Griffithsia. Papers Mich. Acad. Sci., Arts,
and Letters, 27: 111-116. 1942.

Weber-van Bosse, Anna. Liste des algues du Siboga. Siboga Expeditie. Mono-
graphie 59c (3): 313-314. 1923.

446 FarLowIA, VoL. 2, 1946

PLATE I

All figures are drawn with the aid of a camera lucida. Each unit of scale A equals
200 w; scale B, 100 w; scale C, 50 w; scale D, 10 pz.

Grifithsia ovalis Harvey. Figs. 1-4.
1. Vegetative cells with involucrate tetrasporangia between the articulations.
(Scale B)
2. Habit of repent specimen showing rhizoid. (Scale A)
3. Involucrate tetrasporangia near tip of plant. (Scale B)
4. Mature tetrasporangium on a pedicel. (Scale D)

Grifithsia rhizophora Grunow ex Weber-van Bosse. Figs. 5-9.

5. (Upper figure) Mature tetrasporangia. (Scale D); (lower figure) tetrasporangia
on pedicel. (Scale C)

Habit of typical specimen with forcipate tips. (Scale B)

Habit of repent specimen with rhizoids. (Scale C)

Tetrasporic plant. (Scale C)

Habit of specimen with nearly straight tips. (Scale B)

2D

ABBOTT: GRIFFITHSIA (RHODOPHYCEAE) IN HAwalt 447

Prate I

448 FartowlA, VoL. 2, 1946

PLATE II

All figures are drawn with the aid of a camera lucida. Each unit of scale A equals
200 w; scale B, 100 x; scale D, 10 pw.
Griffithsia ovalis Harvey. Figs. 1-2.

1. Habit of erect specimen showing fastigiate branching. (Scale A)

2. Habit of creeping specimen showing secund branching. (Scale A)
Griffithsia Metcalfii Tseng. Figs. 3-6.

3. Tetraspores between articulations of vegetative cells. (Scale B)

4. Tip of a frond. (Scale B)

5. Spermatangial cluster. (Scale D)

6. Tetrasporic cluster. (Scale D)

449

GRIFFITHSIA (RHODOPHYCEAE) IN Hawall

ABBOTT:

Pirate II

450

FarLow14, VoL. 2, 1946

PLATE III

All figures are drawn with the aid of a camera lucida. Each unit of scale B equals
100 uw; scale C, 50 uw; scale D, 10 wu.

Griffithsia tenuis C. Agardh. Figs. 1-7.

1.

2
3
4.
5

io a)

Whorl of four tetrasporangia. (Scale C)

. Habit of specimen with male cluster. (Scale B)
. Spermatangial cluster. (Scale D)

Habit of robust specimen, showing young tetraspore. (Scale C)

. Part of cystocarp, showing supporting cell derivatives and pyriform carpospores.

(Scale D)
Involucrate cystocarp on lateral pedicel. (Scale C)
Young gonimoblast with involucre. (Scale D)

451

AppoTt: GRIFFITHSIA (RHODOPHYCEAE) IN Hawatl

Pirate III

452

FartowlA, VoL. 2, 1946

PLATE IV

All figures are drawn with the aid of a camera lucida. Each unit of scale B equals
100 u; scale C, 50 yw; scale D, 10 x.

Grifithsia Binderiana Sonder. Figs. 1-7.

SD

. Habit of portion of tetrasporic plant. (Scale B)
. Spermatangial cluster with young involucre. (Scale D)

Spermatangial cluster on lateral pedicel, with surrounding involucral cells.
(Scale B)
Detail of mature cystocarp showing a few carpospores. (Scale C)

Tetrasporic plant with involucrate tetrasporangia. (Scale B)

(a) Young tetraspore; (b) mature tetrasporangium showing pedicel. (Scale D)
Position of cystocarp with regard to involucre on a lateral pedicel. (Scale B)

ABBOTT: GRIFFITHSIA (RHODOPHYCEAE) IN HAWAII 453

Pirate IV

2(4) :455-473 FARLOWIA Juty, 1946

FUNGI DOMINGENSES NOVI VEL MINUS COGNITI. I.

CarLos E. CHARDON

From May to November 1937, the writer was engaged by His Excellency
President Rafael L. Trujillo as head of a party in charge of a partial survey
of the natural resources of the Dominican Republic. Ample travel facilities,
which permitted him to cover most of the territory of that country, were
generously placed at his disposal.

A particularly interesting trip which lasted seven days was made on
horseback from Jarabacoa to San José de Ocoa, in the company of Prof.
H. A. Meyerhoff. The trail, which reaches an altitude of 2,400 meters,
leads across the Cordillera Central through large and little-explored pine
forests. A delightful temperate climate prevailed in these mountains and
several days were spent botanizing in this terra incognita to mycology.
Frequent collections were also made as time permitted, in other sections of
the country. In short, close to a thousand specimens of fungi were collected
which were sent to Prof. H. H. Whetzel to be added to the collections from
the Dominican Republic previously made by M. F. Barrus, F. D. Kern,
R. A. Toro, and the writer, and deposited in the herbarium of the Depart-
ment of Plant Pathology, Cornell University.

The present contribution to the fungous flora of the Dominican Republic
represents an addition of forty-four species, of which fourteen are appar-
ently new to science. The writer wishes to acknowledge his thanks to the
late Prof. H. H. Whetzel, of Cornell University, for encouragement and
helpful suggestions; to Dr. Fred J. Seaver, of the New York Botanical
Garden, for the determination of the Discomycetes and Hypocreales; and
to Miss Edith K. Cash, of the U. S. Department of Agriculture, for con-
tributing the description of a new species of the Phacidiales.

PEZIZALES

-_ -Jn Ciferri’s ““Micoflora Domingensis” (2) there are reported only three
species of Discomycetes proper, all of which are common and well-known
tropical forms. Ciferri’s record is based on the paper published by Berke-
ley in 1852, “Enumeration of some fungi from Santo Domingo” (1), which
was the first contribution published on the fungous flora of the Dominican
Republic. The species are: Cookeina sulcipes (Berk.) Kuntze, Cookeina
tricholoma (Mont.) Kuntze, and Phillipsia domingensis (Berk.) Berk.
Since the appearance of Berkeley’s paper, ninety-two years ago, and in
spite of the intensive collections made in the last twenty years by half
a dozen explorers, not a single cup-fungus has been added to Berkeley’s
original list. This group of fungi has been overlooked because of its sapro-

455

456 FartowlA, Vo. 2, 1946

phytic nature, the former mycologists having been interested in parasitic
forms only.

We have made an effort to collect cup-fungi and have contributed thir-
teen specimens, representing nine different species, of which seven are new
to the flora, and one is new to science. All the Discomycetes included below
were determined by Dr. Fred J. Seaver, of the New York Botanical Garden.

This brings the total of known Pezizales in the Dominican Republic to
ten species. In Puerto Rico, the number of species of this group reported
by Seaver and Chardon (4) is forty-seven. Our list, consequently, is a
mere beginning and could be multiplied many-fold by subsequent explora-
tions.

The most common species found was the cosmopolitan Cookeina sulcipes
forming beautiful conspicuous cups on dead sticks and debris in the shade.
Cacao plantations furnish ideal conditions for its growth and the cups are
found by the hundreds. Cookeina tricholoma and Phillipsia domingensis
require the same habitat but are far less common.

The attractive Ascobolus magnificus, common in Puerto Rico, was en-
countered twice, always on dung, in the shade. Erinella calospora and
E. longispora were both found on the bark of trees in the tropical region.

All the other species were found at much higher altitudes. On our trip
to Constanza Valley, at the high point in Loma Barrero, along the trail
just above Jarabacoa, we found two specimens on dung in a forest: Asco-
phanus granulatus and Patella theleboides. The altitude was 1150 meters.
Further along, a third species was found, Saccobolus Kerverni. All these
species are almost exclusively temperate species.

At Valle Nuevo, at an altitude of 2250 meters, where intense cold was
felt all the time, we found Patella coprinaria on dung in the meadows near
an old abandoned hut. This is the highest station at which a Discomycete
has been found in the West Indies. This is also a temperate-zone species.
It is not known in the tropical and subtropical zones, except for a single
collection in the Luquillo Mountains of Puerto Rico.

Ascobolus magnificus Dodge, Mycologia 4: 218. 1912.

First described from Puerto Rico, where it is common on cow dung.
Also known from the Cauca Valley (Colombia) and New York City, where
it has been cultivated in damp-chambers. Careful search for this species
was rewarded by the collection of two specimens. The cup-shaped apothecia
with the dark violet hymenium at maturity are typical.

On cow dung, Prov. Samana, in forests at Sabana la Mar, Chardon 703, July 5,
1937; Prov. La Vega, in shaded places near Jarabacoa, Chardon 1136, Sept. 13, 1937.

Ascophanus granulatus (Bull.) Speg., Michelia 1: 235. 1879.

Small apothecia, 1-2 mm. in diameter, pale orange, with the concave
hymenium orange. A common temperate-zone species.

On cow dung, Prov. La Vega, trail from Jarabacoa to Constanza, 1100-1200 m.,
Chardon 1139, Sept. 13, 1937.

CHARDON: FuNnciI DoMINGENSES I. 457

Cookeina sulcipes (Berk.) Kuntze, Rev. Gen. Pl. 2: 849. 1891.

One of the commonest and most conspicuous cup-fungi in the American
tropics; also known from Australia. The cups are deep orange to rose color,
attaining a diameter of 2 cms. or more. Specimens nos. 722 and 735 were
collected among many hundreds of cups found in a well-sheltered cacao
plantation, where moisture and shade afforded ideal conditions for develop-
ment. Again at El Hatillo (specimen no. 1117) the cups were very abund-
ant, literally covering portions of the floor of a dense, humid forest.

On dead wood and forest debris, Prov. Samanas, Cacao plantations near Sabana la
Mar, Chardon 722 & 735, July 5, 1937; Cacao plantations above Samana, Peninsula
de Samana, Chardon 739, July 6, 1937; Prov. La Vega, forests at El Hatillo, Chardon
1117, Aug. 28, 1937.

Cookeina tricholoma (Mont.) Kuntze, Rev. Gen. Pl. 2: 849. 1891.

Also known throughout the American tropics and Old World tropics.
Of less frequent occurrence than the preceding species, the specimen cited
below consisting of three apothecia. Cups red or deep red, externally
clothed with long hairs.

On dead wood, Prov. La Vega, forests at El Hatillo, Chardon 1231, Aug. 28, 1937.

Erinella calospora Pat. & Gaill., Bull. Soc. Myc. Fr. 4: 101, 1889.

On living bark of Jambos, Prov. La Vega, forests at El Hatillo, Chardon 1231,
Aug. 28, 1937.

Erinella longispora (Karst.) Sacc., Syll. Fung. 8: 507. 1889.
On dead wood, Prov. La Vega, forests, road to Cotui, Chardon 975, Aug. 6, 1937.

Saccobolus Kerverni (Crouan) Boudier, Ann. Sci. Nat. Ser. V. 10: 229. 1869.

This species, with the exception of a single collection from Puerto Rico,
occurs in the temperate regions of the world. It is common throughout
North*America and Europe, and is also found in Bermuda.

On cow dung, Prov. La Vega, trail from Jarabacoa to Constanza, 1100-1200 m.,
Chardon 1140, Sept. 13, 1937.

Patella coprinaria (Cooke) Seaver, N. Amer. Cup-Fungi 171. 1928.

Apothecia small, 2-3 mm. in diameter, bright yellow in color, bordered
with hairs. This is a temperate-zone species, occurring in the United States
and Europe. Its appearance in the temperate zone of Hispaniola may be
significant in elucidating problems of geographical distribution.

On cow dung in meadows, Prov. La Vega, Valle Nuevo, above Constanza, Cor-
dillera Central, 2250 m., Chardon 1227, Sept. 16, 1927

Patella theleboides (Alb. & Schw.) Seaver, N. Amer. Cup-Fungi, 170. 1928.

Apothecia densely crowded, small, 2-3 mm. in diameter, bright yellow.
Another temperate-zone species, reported in the United States, from Con-
necticut and Delaware to California; also in Europe.

On cow dung in the shade, Prov. La Vega, trail from Jarabacoa to Constanza,
1100-1200 m., Chardon 1140, Sept. 13, 1937.

458 FarLowlA, VoL. 2, 1946

PHACIDIALES

Nymanomyces Xolismae Cash sp. nov.
Ficures 1-4

Stromata sparsa, atra, pulvinata, subcarbonacea, subcircularia, 2-3 mm. in diam.,
epidermicalia, amphigena; ascomata hypophylla, singularia, subcircularia, 1-2 mm.
in diam., rima longitudinali vel lobis pluribus aperientia, hymenio brunneo; asci
elongato-clavati, octospori, 100-150 x 18-25 mu; ascosporae elongato-ellipsoideae vel
subclavatae, 1-3-seriatae, viridi-brunneae, 25-32 x 5-6 «; paraphyses filiformes, sub-
hyalinae; hypothecium atrum, plectenchymaticum, margine incrassato; pycnidia am-
phigena, loculata; pycnosporae parvae, hyalinae, ellipsoideae, 1.5—2 x 0.7 yp.

Stromata sparse, black, subcarbonaceous, round to angular, 2—3 mm. in
diameter, or occasionally elongate and up to 5 mm. long, pulvinate, am-
phigenous, developing in the epidermis, surrounded on the lower surface by
a broad, pale reddish, circular area, epiphyllous stromata sterile or pycnidial
only; ascomata hypophyllous, single, subcircular, elliptical, or elongate,
1-2 mm. in diameter, opening by a longitudinal slit or more often by sev-
eral lobes, the covering layer later breaking up and falling away, hymenium
dark brown; asci long-clavate, gradually narrowed to a long pedicel,
rounded at the apex, 8-spored, 100-120 (—150 x 18-20 (-25) p; asco-
spores unicellular, narrow-ellipsoid or subclavate, sometimes slightly con-
stricted near the center, obliquely uniseriate below, irregularly 2—3-seriate
above, greenish-brown, the granulose contents occasionally containing one
or two large hyaline guttules, 25-32 x 5-6 »; paraphyses filiform, numer-
ous, subhyaline to pale brownish, longer than the asci; hypothecial layer
narrow, black, plectenchymatic, dense, and very much thickened toward
the margin. Pycnidia amphigenous, when hypophyllous developing from
the same stromata as the apothecia, irregularly chambered, the locules
frequently becoming confluent and extending over most of the surface of
the stroma; spores minute, hyaline, ellipsoid, filling the locules, 1.5-2 x
0.7-1 », emerging through irregular fissures and forming a white mass on
the surface of the stroma; conidiophores not seen.

On Xolisma sp. (det. N. Y. Bot. Garden), Valle Nuevo, above Constanza, Prov.
La Vega, Sept. 16, 1937, no. 1189 (type) and Sept. 17, 1937, Carlos E. Chardon 1195.
Type specimen in the Mycological Collections of the Bureau of Plant Industry, Belts-
ville, Maryland; at Cornell University and in Chardon herbarium.

The only similar species described on an ericaceous host is Criella leuco-
thoes P. Henn., which occurs in Brazil on an undetermined species of
Leucothoe. No specimens of this species were available for comparison,
but it appears to differ from the fungus on Xolisma in the epiphyllous
stromata and much broader spores.

HYPOCREALES

Cordyceps militaris (L.) Link, Handb. 3: 347. 1833.

On pupa of an insect, Prov. Monte Cristi, forests back of Copey, Chardon 520,
June 8, 1937.

CHARDON: FuncrI DoMINGENSES I. 459

Hypocrea citrina (Pers.) Fries, Summa Veg. Scand. p. 185. 1849.
On dead wood, Prov. La Vega, trail from Maimon to Yuna River, Chardon 1030,
Aug. 13, 1937.

Hypocrea patella Cooke & Peck; Peck, Ann. Rep. N. Y. State Mus. 29: 57. 1878.
On dead wood, Prov. La Vega, road to Cotui, Chardon 966, Aug. 6, 1939.

Hypocrea rufa (Pers.) Fries, Summa Veg. Scand. p. 383. 1849.
On dead wood, Prov. La Vega, road to Bonao, Chardon 797, July 18, 1937.

Stilbocrea hypocreoides (Kalchbr. & Cooke) Seaver, Mycologia 2: 62. 1910.

Sphaerostilbe hypocreoides Kalchbr. & Cooke, Grevillea 9: 26. 1880.
Sphaerostilbe intermedia Ferd. & Winge, Bot. Tidssk. 29: 12. 1908.

On dead wood, Prov. Monte Cristi, forests back of Copey, Chardon 520, June 8,
1937.

- DOTHIDEALES

Dothidella domingensis Chardon sp. nov.
FicurREs 9-13

Stromata semper amphigena, atra, crustosa, conspicua, supra superficiem folii elevata,
fere rotundata, 1 mm. diam., partem magnam folii tegentia, dothideacea (e cellulis
distinctis fuscis composita) ; loculi 2-7 in quoque stromate, globosi, 96-120 wu diam.;
asci clavato-cylindracei, octospori, 107-124 x 15-18 mw; sporae apice et basi monostichae,
medio distichae, uniseptatae, hyalinae, longe ellipsoideae, una cellula majore obtusa,
altera minore subacuta, 24-27 x 7-8 yp, leves, sine tunica crassa; paraphyses carentes.

Stromata amphigenous only, black, crust-like, conspicuous, raised above
the surface of the leaves of the host, approximately circular, 1 mm. in
diameter, covering a large portion of the leaf, dothideaceous (made up of
distinct, dark-brown cells); locules several, 2-7 in each stroma, globose,
96-120 » in diameter; asci cylindrical-clavate, 8-spored, 107-124 x 15-18
u, With the spores uniseriate at the top and bottom, biseriate in the main
body of the ascus; spores 1-septate, hyaline, long-elliptical, with one cell
larger and with an obtuse end, the other cell smaller with a subacute end,
24-27 x 7-8 p, surface smooth, without a thick cell wall; paraphyses
absent. :

The fungus looks superficially like Dothidella tinctoria (Tul.) Sacc.,
common on Compositae in the upper subtropical and temperate zone of
the Andes of Venezuela and Colombia. In fact it looked identical to us,
until a microscopic study disclosed that it was a new species. The spores
were in asci (which is rarely observed in D. tinctoria where the spores are
loose from a very early stage). The ascospores are also different; they are
hyaline, 1-septate, with two unequal cells, and no thick cell wall was
found. In D. tinctoria the cells are equal, the spores larger, 26-30 x 5-6 p,
the contents granular, and a thick cell wall envelops the ascospore.

For the marked differences cited above, reference is made to Figures 12
and 13.

On Baccharis myrsinites (Lam.) Pers., along trail from Jarabacoa to Constanza,
Cordillera Central, 1300 m., Chardon 1142, Sept. 13, 1937 (type).

460 FarLowIA, VoL. 2, 1946

Trabutia conica Chardon, Mycologia 13: 292. 1921.

The type is Whetzel & Olive 658, from Mayaguez, Puerto Rico, on
Drepanocarpus lunatus, with which our material compares well. The
species is known to occur also in Venezuela (Chardon & Toro 474), on
Lonchocar pus punctatus. Not known elsewhere. The stromata are typically
conical, epiphyllous, circular, about 1 mm. across, black, shiny, uniloculate,
located between the cuticle and the epidermis. Spores globose, 9-11 yp,
inordinately placed in the ascus.

On Drepanocarpus lunatus (L.£.) G. Meyer, Prov. Puerto Plata, road to Blanco,
Ciferri 4871, June 15, 1931 (in Chardon’s herb.).

Trabutia Danthoniae Chardon sp. nov.

Stromata atra, graminicola, linearia, 1 mm. longa, 0.5 mm. lata, interdum con-
fluentia longioria et leviter elevata, subcuticularia; loculi 1-2 in quoque stromate,
applanato-lenticulares, 150-240 x 74-90 yu, primo stylosporas longas, filiformes, 25-28
x 1 w, posterius ascos gignens; asci clavati, octospori, 70-85 x 17-21 mw; sporae dis-
tichae, hyalinae usque subhyalinae, continuae, longe ellipsoideae, apice subacuto, 19-25
x 7-9 w; paraphyses praesentes.

Stromata small, black, graminicolous, linear, 1 mm. long, 0.5 mm. wide,
sometimes confluent and forming longer stromata slightly raised, sub-
cuticular; locules 1-2 in a stroma, flat-lenticular, 150-240 x 74-90 uh,
giving rise first to long, cylindrical stylospores, 25-28 x 1 yp, later to asci;
asci clavate, 8-spored, 70-85 x 17-21 yw, with the spores biseriate; spores
hyaline to subhyaline, 1-celled, long ellipsoidal with one end subacute,
19-25 x 7-9 »; paraphyses present.

This interesting species forms inconspicuous stromata on a grass which
is peculiar to and widespread in the higher elevations of the Cordillera
Central. The subcuticular character of the stroma makes the fungus fall
under Trabutia.

On Danthonia domingensis Hack. and Pilger, Prov. Azua, trail from Valle Nuevo
to Rio de las Cuevas, Cordillera Central, 1770 m., Chardon 1200, Sept. 17, 1937 (type).

Phaeotrabutia Smilacis Chardon sp. nov.

Stromata epiphylla, rotundata, nigra, lucida, conspicua, 1-3 mm. diam., inter
cuticulam et epidermidem folii definite insidentia; loculi generaliter singuli, applanati,
650-720 x 85-95 gw, supra clypeo denso atro, infra membrana tenui stromatica, brunnea
praediti, interdum loculi 2-3, 320-410 x 80-90 yw; asci clavati usque cylindro-clavati,
octospori, 123-135 x 29-38.5 m; sporae in parte praecipua asci distichae, continuae,
clare brunneae, ellipticae, 19-25 x 8.5-11 uw; paraphyses filiformes.

Stromata epiphyllous, circular, black, shiny, conspicuous, 1-3 mm. in
diam., distinctly located between the cuticle and the epidermis of the leaf;
locule single, flattened, 650-720 x 85-95 y, with a heavy black clypeus
above and a thin, brown, stromatic tissue below, sometimes divided into
2 or 3 locules, 320-410 x 80-90 y; asci clavate to cylindric-clavate, 8-
spored, 123-135 x 29-38.5 qu, with the spores biseriate in the main body
of the ascus; spores 1-celled, distinctly brown, elliptical, 19-25 x 8.5-11 BS
paraphyses present, filiform.

CHARDON: FUNGI DoMINGENSES I. 461

The genus Phaeotrabutia was recently erected by Dr. Carlos Garcés
Orejuela (Caldasia No. 2: 77. 1941) to take care of species like Trabutia,
possessing brown spores. In our Dominican material, the stromata lie
between the cuticle and the epidermis, and the spores are unicellular, dis-
tinctly brown in color.

On Smilax coriacea Spreng., Prov. Azua, trail from Valle Nuevo to Rio de las
Cuevas, Cordillera Central, 1600 m., Chardon 1202, Sept. 17, 1937 (type).

Catacauma Sabal Chardon sp. nov.
Ficures 14-17

Stromata amphigena, in utraque superficie folii pariter visibilia, angulata, leniter
elevata, sordide brunnea, conspicua, 2-3 mm. longa (axe principali axem folii sequenti),
1.5 mm. lata, subter epidermidem in maturitate distincte visibilem evoluta, mesophyl-
lum penentrantia; loculi plures, magni, 250-420 x 93-120 uw, applanati usque labyrinthi-
formes, supra clypeo atro denso praediti, infra et lateraliter sine membrana stromatica;
asci clavati, octospori, exigue pedicellati, 68-85 x 35-43 «; sporae in ascis irregulariter
conglobatae, continuae, hyalinae, naviculares, 28-30 x 10-12 m@, subgranulosae; para-
physes: filiformes, rarae.

Stromata amphigenous, equally visible on both surfaces of the leaves
of the host, angular, slightly raised, dirty brown in color, conspicuous,
2-3 mm. long (with the main axis following that of the leaf), 1.5 mm. in
width, originating beneath the epidermis, and in mature stromata the
epidermal layer clearly seen above, not penetrating the mesophyll; locules
several, large, 250-240 x 93-120 uy, flattened to labyrinthiform, with heavy
black clypeus above and no stromatic tissue below or on the sides; asci
clavate, 8-spored, short-pedicellate, 68-85 x 35-43 », with the spores irregu-
larly pressed in the ascus; spores 1-celled, hyaline, navicular, 28-30 x
10-12 p, with the contents slightly granular; paraphyses filiform, scarce.

The stromata are very characteristic; they are angular, dirty brown
in color. The stromata are subepidermal and the row of epidermal cells is
seen clearly above the clypeus. The spores are large, navicular. The
species, which is apparently new, is referred to Catacauma on account of
the position of the stromata in the leaf tissue.

On dead leaves of Sabal palm, Prov. Barahona, near sulphur springs beyond
Duvergé, Chardon 760, July 11, 1937 (type).

Phyllachora Acaciae P. Henn., Hedwigia 33: 233. 1894.

Widely distributed in tropical America on various foatinous trees and ©
shrubs; stromata very minute, punctiform, less than 0.5 mm. in diameter,
visible from both surfaces of the leaf, uniloculate; asci 60-70 x 20 w (in
our specimen); spores biseriate or inordinate, long-elliptical, 14-16 x
6-7 p.

On Acacia sp., Prov. Duarte, road to San Francisco de Macoris, Chardon & Toro
851, July 11, 1937.

Phyllachora Bourreriae Stevens & Dalbey, Bot. Gaz. 68: 54. 1919.
Our specimen has been compared with type material, which is Stevens
4149, from Vega Baja, Puerto Rico. Stromata amphigenous, black, circu-

462 Fartowia, VoL. 2, 1946

lar, 1.0-1.5 mm. in diameter, enclosing many locules; spores obliquely
uniseriate, elliptical, 8.5-12.5 x 7 uy.

On Bourreria sp., Prov. Puerto Plata, La Cumbre, Cordillera Septentrional, Char-
don 1129, Aug. 30, 1937.

Phyllachora Exostemae Chardon sp. nov.

Stromata minuta, inconspicua, rotundata, 0.4-0.8 mm. diam., atra, in epiphyllo
leniter arcuata, in hypophyllo vix visibilia, dispersa, non connata; loculi singuli, in
mesophyllo centraliter insidentes, globosi vel fere globosi, 480-670 m in diam., supra
clypeo atro, crasso, loculum non superanti praediti, stromate incrassato, atro circum-
dati; asci longe cylindracei, octospori, 95-110 x 7-8 «; sporae monostichae, continuae,
hyalinae longe fusiformes, 12-14 x 4-5 ms; paraphyses praesentes.

Stromata minute, inconspicuous, circular, 0.4-0.8 mm. in diameter,
black, slightly convex in the epiphyll, scarcely perceptible in the hypo-
phyll, scattered, not coalescing; locule single, centrally located in the
mesophyll, globose or approximately so, 480-670 p» in diameter, with a
thick black clypeus above, not exceeding the locule, and heavy black
stroma all around; asci long-cylindrical, 8-spored, 95-110 x 7-8 pu, with the
spores uniseriate; spores 1-celled, hyaline, long-fusoid, 12-15 mw; para-
physes present.

On Exostema sp., Las Caobas, San José de Ocoa, Chardon 781, July 10, 1937 (type).

Phyllachora gratissima Rehm, Hedwigia 31: 306. 1892.

The type is from Bafos, Ecuador, collected by Lagerheim, in 1891. This
is a well-known, conspicuous species, forming large, tar-like spots, 2-3 mm.
in diameter, on the upper surface of the leaves of the avocado. It is com-
mon in the higher elevations in Puerto Rico and in the subtropical zone
of the Andes of Venezuela and Colombia. The fungus seems to be restricted
to the humid subtropical zones. Our collection from the Dominican Re-
public is not an exception to this rule.

On Persea gratissima L., Prov. La Vega, trail from Jarabacoa to Constanza, 600 m.,

Chardon 1143, Sept. 13, 1937; Constanza Valley, 1150 m., Chardon 1170, September 15,
1937. —

Phyllachora insueta Sydow, Ann. Mycol. 23: 372. 1925.

The species was originally described from material collected in Grecia,
Costa Rica, on Serjania caracasana and distributed under Sydow’s Fungi
exot. exs. 662. Also known from various collections made by the writer in
Venezuela and Colombia.

The stromata in the Dominican material are typical: pustule-like, raised
above the leaf epidermis, yellowish-green, dotted with minute, black specks,
which are the clypei of the locules. In cross section, the locules are numer-
ous, labyrinthiform, with the leaf tissue hypertrophied around the locules.
The spores are ellipsoidal, uniseriate, 15 x 6 ». This is the first report of
the fungus from the West Indies.

On Serjania sp., Prov. Samana, Savanas, near Sabana la Mar, Chardon 709, July 5,
1937.

CHARDON: Funct DomINGENSESs I. | ; 463

Phyllachora Masini Toro, Sci. Survey Porto Rico & Virgin Isl. 8 (1): 53. 1926.

The type specimen is from the mountains above Yauco, Puerto Rico,
Whetzel, Kern & Toro 2482. The spots are black, circular, 1-2 mm. in
diameter, more conspicuous in the epiphyll, bordered by a reddish zone
of host tissue. The shape and measurement of the asci and spores agree
with the type material.

On Brunellia comocladifolia H. & B., Prov. La Vega, trail from Constanza to
Valle Nuevo, 1575 m., Chardon 1181, Sept. 16, 1937.

Phyllachora ocoensis Chardon sp. nov.
FIGuRES 18-21

Stromata amphigena, parva, punctiformia, atra, rotundata, 0.5-1.0 mm. in diam.,
in hypophyllo convexa, in epiphyllo tantum leniter elevata; loculi singuli, globosi,
300-360 x 250-278 , ostiolo parvo epidermidem versus muniti, undique (praesertim
infra supraque) membrana stromatica atra cincti; asci cylindrico-clavati, octospori,
95-110 x 17-19 yw; sporae distichae vel in asco aggregatae, continuae, hyalinae, longe
naviculares, 17-23 x 4.5-6.0 w, plasmate granuloso, homogeneo; paraphyses praesentes.

Stromata amphigenous, small, punctiform, black, circular, 0.5 to 1.0
mm. in diameter, convex in the hypophyll, only slightly raised in the epi-
phyll; locule single, globose, 300-360 x 250-278 p, with a small beak facing
the epidermis, surrounded on all sides (especially above and below) with
black, stromatic tissue; asci cylindrical-clavate, 8-spored, 95-110 x 17-19 p,
with the spores biseriate or crowded in the ascus; spores 1-celled, hyaline,
long-navicular, 17-23 x 4.5-6.0 », with the contents homogeneous; para-
physes present.

This small Phyllachora macroscopically resembles Ph. Acaciae P. Henn.,
a species of wide distribution in tropical America, but the stromata are
more conspicuous. In Ph. Acaciae they do not reach 0.5 mm. in diameter.
The spore characters are also different (in Ph. Acaciae they are 14-16 x
5 p»). A cross section of the leaf of our material shows that the base or
bottom of the locule is in the hypophyll, where it bulges out conspicuously,
while the upper portion of the locule has a small neck which faces the
epiphyll.

On leguminous shrub (with pinnate leaves), Las Caobas, road to San José de
Ocoa, Chardon 773, July 10, 1937 (type).

Phyllachora Ocoteicola Stevens & Dalbey, Bot. Gaz. 68: 57. 1919.

Our material has been compared with the type specimen, Stevens 4768,
from Monte Alegrillo, Puerto Rico. In the original diagnosis the spores
are described as “oblong to cylindrical, 17-54 », somewhat pointed at one
end.” This is a gross typographical error. We have reexamined the type,
as well as Stevens 1520, also from Puerto Rico, and they are 14-20 x 5-7 p,
with one end tapering. ;

In the specimen from the Dominican Republic, the stromata are black,
shiny, angular, 2-3 mm. across, surrounded by a zone of dead host tissue,
characters which compare well with the Puerto Rico material. The spores
are 17-21 x 7-8 p, a trifle larger, pointed at one end.

On Ocotea sp., Prov. La Vega, Constanza Valley, 1150 m., Chardon 1175, Sept. 14,
1937.

464 FartowiA, Vor. 2, 1946

Phyllachora panamensis Stevens, Ill. Biol. Monogs. 11 (2): 37. 1927.
FIGURES 22-27

The type material is Stevens 1351, Fungi of Panama. Our Dominican
material compares well with it in macroscopic characters, in the size and
shape of the asci, and in the ellipsoidal spores 15-17 x 6-8 pu, Which are ar-
ranged uniseriately in the ascus. It has also conidia which are small, globose
to elliptical, about 2 » long. Our specimen is not Phyllachora Roureae
Sydow from the Philippines, the description of which was based on conidial
material (Philippine Jour. Sci. 8 (4) sec. C: 227. 1923), distributed as
Sydow’s Fungi exot. exs. 187. We have examined this material and it
has filiform conidia (stylospores) which measure 24-46 x 1-1.5 p. (See
fig. 26.)

On Rourea glabra H.B.K., Prov. La Vega, near Maimén, Chardon 1064, Aug. 13,
1937.

Phyllachora Phaseoli (P. Henn.) Theiss. & Sydow, Ann. Mycol. 13: 507. 1915.

The type, Physalospora Phaseoli P. Henn. from Brazil, has not been
examined, but our material has been compared with numerous specimens
from Puerto Rico and with Sydow’s Fungi exot. exs. 667, from San José,
Costa Rica. The stromata are typical: punctiform, minute, in dense cir-
cular groups; the spores are 9-12 x 6-8 p, broad-elliptical.

On Phaseolus lunatus macrocarpus, Hato del Yaque, Ciferri 4842, Feb. 7, 1932;
on Phaseolus aff. adenanthus, road from Santiago to Puerto Plata, Ciferri 4575, Feb. Ss,
1931; on Phaseolus sp., Dist. Nacional, ravine at Km. 30, Duarte Road, Chardon 677,
June 30, 1939,

Phyllachora Scleriae Rehm, Hedwigia 39: 232. 1900.

Originally described from South America. A common species in Puerto
Rico, on Scleria; also known from Panama City (Chardon & Nolla 188),
Spores long-fusiform, 19-21.5 x 5 ». Probably present in other West Indian
islands.

On Scleria pterota Presl., Prov. Trujillo, Sabana de Guabatico, north of Guerra,

Chardon 553, June 20, 1937; Dist. Nacional, ravine at kilometer 30, Duarte Road,
Chardon 678, June 30, 1937.

Phyllachora viequesensis Orton & Toro, Sci. Surv. Porto Rico & Virgin Isl. 8 (1):

54. 1926.

Stromata typical of the Phyllachoraceae on grasses and sedges, several
loculate (mostly 2-3), the stromata formed in large air cavities beneath
the epidermis of the host. Spores obliquely uniseriate, long-fusiform, 18-20
x 6 mw; paraphyses present. Our material has been compared with type
material, Whetzel, Kern & Toro 2648, on the same host, from Vieques
Island, Puerto Rico.

On Cyperus ligularis L., Prov. Samana, beach at Sabana la Mar, Chardon 688,
July 4, 1937; Prov. Barahona, shores of Lake Rincon, near Cabral, Chardon 796,
July 11, 1937.

Phyllachora Paspalicola P. Henn., Hedwigia 48: 106. 1908.

Common throughout tropical America, where the host plant is widely

distributed. Spores lemon-shaped, 8-10 x 5—6 BM.

CHARDON: FuNcI DoMINGENSES I. 465

On Paspalum conjugatum Berg., Prov. Trujillo, San Cristobal, Kern & Toro 156,
Mar. 12, 1926; Prov. Azua, Las Caobas, San José de Ocoa, Chardon 779, July 10,
1937; Puerto Plata, ravine at Km. 37, road to Puerto Plata, Chardon 914, Aug. 1,
1937; Prov. La Vega, ravine, base of Loma Peguera, Chardon 1059, Aug. 13, 1937.

Rhagadolobium cucurbitacearum (Rehm) Theiss. & Syd., Ann. Mycol. 12: 275.
1914.

. Spots conspicuous, in the undersurface of the leaf, appearing like dusted

gunpowder. Reported from various Tropical American countries.

On Gurania sp., Prov. La Vega, trail from Jarabacoa to Constanza, 1100-1300 m.,
Chardon 1145, Sept. 14, 1937.

FIMETARIALES

Sordaria curvula De Bary, Morph. Phys. Pilze p. 209. 1866.
On cow dung, Prov. Azua, river forest, near San Juan de la Maguana, Chardon
623, June 27, 1937.

SPHAERIALES

Melanopsamma Milleri Chardon sp. nov.
Ficures 5-8

Perithecia superficialia, atra, carbonacea, fere 0.8 mm. in diam., 0.1 mm. alta,
hemisphaericalia usque conoidea, levia, non nitentia, Rosellinia similis, in strato ligneo
subimmersa, ostiolo distincto papilliformi praedita; muro perithecii duplici, e pariete
exteriore duro, carbonaceo, et interiore molli, brunneo composito; asci cylindracei,
breviter pedicellati, maximi, 170-210 x 12-17 uw, apice et basi membrana tenui pseudo-
parenchymatica e paraphysoideis composita tunicati, octospori; sporae ex parte dis-
tichae in asco dense compressae, uniseptatae, hyalinae, longe fusiformes, interdum
subarcuatae, ad septum constrictae, magnae, 59-64 x 9-12 yu, leves, utrinque acutae;
paraphysoidea filiformia.

Perithecia superficial, black, carbonaceous, about 0.8 mm, in diameter,
1.0 mm. in height, hemispherical to conical, with the surface smooth, not
shiny (Rosellinia-like), seated on the woody substratum where they are
slightly immersed, and provided with a pronounced papilliform ostiolum;
perithecial wall made up of a double wall, the outer hard, carbonaceous,
and the inner brown and soft; asci cylindrical, short-pedicellate, very large,
170-210 x 12-17 y, embedded in paraphysoids that are connected with a
fine pseudoparenchymatous tissue on the top of the perithecium and a
similar one at the base, 8-spored, with the spores partially biseriate, closely
pressed in the ascus; spores 1-septate, hyaline, long fusiform, sometimes
slightly arcuate, constricted at the septum, large, 59-64 x 9-12 p, smooth,
with the ends acute; paraphysoids filiform.

A beautiful species, picked up for a Rosellinia. The conspicuous black
perithecia are seated on the woody matrix with no subiculum or stroma.
The species is dedicated to Dr. Julian H. Miller, of the University of
Georgia, a distinguished student of the Sphaeriales, who confirmed the
determination of the fungus as a new species and its inclusion under
Melanopsamma. ;

On dead wood, La Vega, forests at El Hatillo, near Yuna river, Chardon 1094,
Aug. 28, 1937 (type).

466 FartowiA, VoL. 2, 1946

Lasiosphaeria pezizula (Berk. & Curt.) Sacc. Syll. Fung. 2: 195. 1883.
Sphaeria pezizula Berk. & Curt., Grevillea 4: 106. 1875.

Previously known from Puerto Rico, Bermuda, and continental North
America. (Determination made by Professor H. M. Fitzpatrick.)
On dead wood, La Vega, near Jarabacoa, Chardon 1229, Sept. 13, 1937.

Calyculosphaeria calyculus (Mont.) Fitzpatrick, Mycologia 15: 51. 1923. :
Sphaeria (Caespitosa) calyculus Mont., Ann. Sci. Nat. ser. II, 14: 322. 1840.
Sphaeria calyculus Mont., Syll. Crypt. p. 226. 1856.

For more details of this interesting species see Fitzpatrick (J.c.). The
type is from French Guiana, collected by Leprieur. (Determined by Dr.
J. H. Miller.)

On decaying fence posts, La Vega, Maimén River, near Maimén, Chardon 1009, Aug.
13, 1937.

Leptosphaeria Cecropiae Chardon sp. nov.

Maculae amphigenae, epiphyllae parvae, rotundae, 1.0-1.5 cm. in diam., centro
peritheciis aggregatis atratae, hypophyllae pustulatae, parvae, argenteae ut colore
folii, ostiolis peritheciorum minute punctulatae; perithecia pluria, 5-12, in quaque
macula, lageniformia, 240-300 x 215-270 uw, pariete brunnea praedita; asci cylindracei,
175-190 x 12-14 yw, octospori; sporae oblique monostichae, longe fusoideae, clare
brunneae, 3-septatae, ad septa constrictae, una cellula terminali attenuata; paraphyses
filiformes, numerosissimae.

Spots amphigenous, in the epiphyll appearing as small, circular, brown
spots, 1-1.5 cms. in diameter, with the center black due to the coalescing
the leaf, but showing minute black spots which are the necks of the peri-
thecia; perithecia several (5-12) on each spot, flask-like in section, 240-
300 x 215-270 », provided with a brown wall; asci cylindrical, 175-190
x 10-14 y», 8-spored, with the spores obliquely uniseriate in the ascus;
spores long-fusoid, distinctly brown, 3-septate, constricted at the septa
and one end cell tapering; paraphyses filiform, very abundant.

Macroscopically, the pustule-like spots which appear on the under-
surface of the leaf are typical. Both the asci and spores show beautifully
under the microscope.

On Cecropia peltata L., Kilometer 67, near La Cumbre, road to Bonao, Chardon
& Toro 832, July 11, 1937 (type).

Leptosphaeria Smilacis Chardon sp. nov.

Maculae magnae, in utraque pagina folii visibiles, conspicuae, orbiculares vel ellip-
ticae, plus minus 1.0 cm. in diam., vel confluentes et majores, subalbae, peritheciis
numerosis minutis punctatae, e zona atra definita marginatae; perithecia minuta, in
textura matricis immersa, globosa, 145-170 uw in diam., pariete definito e cellulis brun-
neis composito, non-ostiolata; asci cylindrico-clavati, octospori, parvi, 47-55 x 8-9 y, in
stipitem brevissimum contracti; sporae irregulariter conglobatae, brunneae, 3-septatae,
longe fusiformes, subcurvatae, 14-19 x 4-5 «; paraphyses praesentes.

Spots large, visible on both surfaces of the leaves of the host, conspicu-

ous, circular, or elliptical, about 1 cm. in diameter or larger through
coalescence, whitish, with numerous minute specks (perithecia) in the

CHARDON: FuNcI DoMINGENSES I. 467

spot, provided with a definite blackish margin; perithecia small, immersed
in the host tissues, globose, 145-170 » in diameter, with a definite cell
wall made up’ of brown cells, without a beak; asci cylindrical-clavate,
8-spored, small 47-55 x 8-9 p, short-stipitate, with the spores inordinately
pressed together; spores brown, 3-septate, long-fusiform, slightly curved,
14-19 x 4-5 w; paraphyses present.

The spots produced by this fungus are very conspicuous on both sur-
faces of the leaves of the host, with many minute specks and a definite
border. The asci are clearly seen under the microscope with the brown,
3-septate spores tightly pressed together.

On Smilax sp., Distrito Nacional, road from Ciudad Trujillo to Bocachica, Chardon
539, June 13, 1937 (type).

Didymosphaeria Trichostigmae Chardon sp. nov.

Maculae amphigenae, brunneae, fere orbiculares, 2-4 mm. in diam., margine in-
definita cinctate; perithecia in maculis densiuscule (6-13) disposita, semper epiphylla,
inconspicua, punctiformia, globosa, parva, 210-280 x 160-175 p, pariete cellulari
definito, circa ostiolum incrassato et obscuriore praedita; asci numerosi, fasciculati,
cylindracei, octospori, 95-133 x 8-10 mw; sporae inordinatae, bicellulares, elongatae,
brunneae, ad septa constrictae, 28-34 x 3-4 uw; paraphyses praesentes.

Spots amphigenous, light brown, approximately circular, 2-4 mm. in
diameter, with the margins indefinite; perithecia crowded in the spots
(6-15 in number), visible only in the epiphyll, inconspicuous, punctiform,
globose, small, 210-280 x 160-175 p, provided with a definite, brown,
cellular perithecial wall, which thickens around the region of the ostiolum
and becomes dark-brown, the numerous asci fasciculate inside the peri-
thecium; asci cylindrical, 8-spored, 95-135 x 8-10 y, with the spores in-
ordinate; spores 2-celled, long rod-like, brown, constricted at the septum,
28-34 x 3-4 mw; paraphyses present.

This is different from Linospora Trichostigmae Stevens, described on
the same host from Puerto Rico. In Clements and Shear (3, p. 68), there
are two genera in the Sphaeriaceae-Phaeodidymae which have close peri-
thecial characters: Stegastroma, with a clypeus, and Didymosphaeria with-
out a clypeus. Our specimen has no clypeus; the brown tissue occurring
around the region of the ostiolum is produced by a thickening of the
perithecial wall. Although the walls of this perithecial neck attains a thick-
ness of 50-60 p, it neither loses its cellular structure, nor becomes black,
and hence cannot be interpreted as a clypeus. The asci inside the peri-
thecia are basal-peripheral which fits with Didymosphaeria.

On Trichostigma octandrum (L.) H. Walt., Duarte, road to San Francisco de
Macoris, Chardon & Toro 853, July 11, 1937 (type).

Metasphaeria phyllachoracearum Petrak, Ann. Mycol. 29: 350. 1931.

Telimena domingensis Chardon, Jour. Dept. Agric. Porto Rico 13: 14. 1929.

The specimen cited below, Kern & Toro 156, was studied by the writer
in 1929 (J.c.) and described as a new species of Telimena, on the basis of
the spores being ‘‘fusoid, 3-septate, hyaline to greenish, with both ends
acute, 12-14 x 2-3 yp.”

468 FaRLowIA, VoL. 2, 1946

The researches of Dr. F. Petrak (Mykologische Notizen, XI, /.c.) have
thrown new light on this fungus. He did not see the type material, but
from examination of other material made available to him by Dr. R. Ciferri,
he concluded there was no Telimena, and that the asci and 3-septate, hya-
line, fusoid spores were the ascosporic stage of a Metasphaeria parasitic
on the stromata of certain Phyllachoraceae. A reexamination of the type
of Telimena domingensis Chardon has convinced the writer that he was in
error, and that Petrak’s interpretation is the correct one.

On old phyllachoroid stromata on Paspalum conjugatum Berg., San Cristobal,
Kern & Toro 156, March 12, 1926.

Pleospora Doidgeae Petrak, Ann. Mycol. 25: 295. 1927.
Dictyochorella Andropogonis Doidge, Bothalia 1: 66. 1921.

The synonymy of these two species was clearly established by Petrak
(l.c.). The type of Miss Doidge’s species is on Andropogon nardus from
Natal, South Africa. The species was found in Venezuela by the writer,
parasitic on various phyllachoroid stromata, and careful drawings were
made. (See Monogr. Univ. Puerto Rico, Ser. B, No. 2, plate XX XI, figs.
2-3, 1934.)

In the material from the Dominican Republic the perithecia are found
inside or adjacent to old phyllachoroid stromata; they are clearly provided
with a brown, cellular, perithecial wall; asci 65-85 x 8-10 p, 8-Spored,
with the spores inordinate; spores muriform, brown, with one of the central
cells longitudinally septate, 17-19 x 5-6 uw. This leaves no doubt as to
the identity of the species, which is the first record of its occurrence in
the West Indies.

On old phyllachoroid stromata, on Andropogon bicornis L., Distrito Nacional,
ravine al Kilometer 30, along Duarte Road, Chardon 672, June 30, 1937.

FUNGI IMPERFECTI

Phyllosticta Citharexyli Chardon sp. nov.

' Maculae amphigenae, conspicuae, fere orbiculares vel irregulares, 4-7 mm. in diam.,
zona decolorata membranae matrici emortuae, e pycnidiis numerosis atris punctata
compositae, margine definita, conspicua, leniter elevata rubro-violacea circumdatae;
pycnidia globosa vel subglobosa, in membrana folii epiphyllum versus immersa, parva,
93-120 x 72-81 yw, cellulis definitis composita; conidia unicellularia, hyalina, ovata,
8-9 x 5-7 uw.

Spots amphigenous, conspicuous, approximately circular or irregular
in outline, 4~7 mm. in diameter, made up of a discolored zone of dead host
tissue in which may be seen numerous small, black specks (the pycnidia),
the dead tissue being surrounded by a definite, conspicuous, slightly raised
margin of reddish violet tissue; pycnidia globose or nearly so, immersed
in the tissues of the leaf, facing the epiphyll, small, 93-120 x 72-81 uw, made
up of distinct wall cells; conidia 1-celled, hyaline, ovate in shape, 8-9 x
5-7 p.

On Citharexylon sp., Prov. Samana, Chardon 744, July 6, 1937 (type); Prov.
Santiago, San José de las Matas, Chardon 483, June 6, 1937.

CHARDON: FUNGI DOMINGENSES I. 469

Phyllosticta Stigmatophylli Chardon sp. nov.

Maculae fere orbiculares vel irregulares, amphigenae, 2-4 mm. in diam., epiphyllae
subalbae, centro pycnidiis numerosis laxe dispersis punctatae, linea definita, parum
elevata cincta, hypophylla sordide brunneae, sine linea definita; pycnidia dispersa,
immersa, globosa, 110-125 mw in diam., non-ostiolata, semper epiphylla; conidia
hyalina, unicellularia, ovoidea usque ellipsoidea, 7-9 x 5—6 4, numerosissima.

Spots nearly circular or irregular in outline, amphigenous, 2-4 mm.
across, whitish in the epiphyll, with numerous specks (pycnidia) scat-
tered in the center of the spot, with a slightly elevated, definite black mar-
gin which makes the spot conspicuous; in the hypophyll the spots are
dirty black in color, without a pronounced border; pycnidia scattered
over the infected spot, immersed globose, 110-125 yw in diameter, not
beaked, restricted to the upper side of the leaf; conidia hyaline, 1-celled,
ovoid to ellipsoidal, 7-9 x 5-6 », very numerous.

On Stigmatophylion sp., Prov. Santiago, San José de las Matas, Chardon 494,
June 6, 1937 (type).

Cylindrosporium Securidacae Chardon sp. nov.

Maculae amphigenae, angulares, parvae, 1.0-1.5 mm. in diam., centro brunneae,
margine obscuriore, parum elevato cinctae; acervuli subepidermicales, 100-120 uw in
diam.; conidia unicellularia, hyalina, longe bacillaria, arcuata, 17-25 x 3 mw, plasmate
homogeneo, in conidiophoris hyalinis acutis oriunda.

Spots amphigenous, angular, small, 1.0-1.5 mm. across, brown in the
center, with a slightly raised dark margin; acervuli subepidermal, 100-
120 » across; conidia 1-celled, hyaline, long rod-like, curved, 17-25 x 3 pn,
with the contents homogeneous, borne on hyaline tip-like conidiophores.

On Securidaca volubilis, Prov. Samana, beach at Sabana la Mar, Chardon 695,
July 4, 1937 (type).

INSTITUTO DE AGRICULTURA TROPICAL InstiTuTO NACIONAL AGRICOLA
UNIVERSIDAD DE PuERTO Rico San CRISTOBAL
MayactEz, Puerto RIco ReEpUsLica DoMINICANA

LITERATURE CITED

1. Berkeley, M. J. Enumeration of some fungi from Santo Domingo. Ann. Mag.
Nat. Hist., ser. II, 9: 192-203. 1852.

2. Ciferri, R. Micoflora Domingensis. Est. Agron. Moca (Républica Dominicana)
Ser. B, No. 14: 1-260. 1929.

3. Clements, F. E. & C. L. Shear. The Genera of Fungi. 496 p. Wilson Co. New
York. 1931.

4. Seaver, F. J. & C. E. Chardon. Mycology in Scientific Survey of Porto Rico and
Virgin Islands, 8 (1): 1-208. 1926.

470

FarLowIA, VoL. 2, 1946

EXPLANATION OF FIGURES

Figs. 1-4. Nymanomyces Xolismae sp. nov.

Figs.

1:

2.

5.
4.

Leaf of Xolisma sp. showing hypophyllous ascoma opening by a longitudinal
slit.

Cross section of leaf of ruptured ascomata in the hypophyll disclosing the
hymenium and epiphyllous sterile stromata.

Portion of the hymenium with asci and numerous paraphyses.

Individual ascospores.

5-8. Melanopsamma Milleri sp. nov.

ee

Perithecia, natural size.

Perithecia under binocular, showing papilliform ostiolum; seated and par-
tially immersed in the woody matrix.

An ascus and two paraphyses.

Ascospore.

13. Dothidella domingensis sp. nov.
Habitat sketch of fungus on leaves of Baccharis myrsinites.

Cross section of leaf showing position of compound stroma with several
locules.

. Ascus and spores.

Individual ascospores, showing septation, with one cell larger and obtuse,
and the other cell smaller, with a subacute end (note that there is no thicken-
ing of the cell wall).

Individual ascospores of Dothidella tinctoria showing differences from the
above (note the larger size, the ascospore divided into two cells of approxi-
mately the same size, and the thick cell wall).

CHARDON: FuNcGI DoMINGENSEs I. 471

472 Fartow14, VoL. 2, 1946

Figs. 14-17. Catacauma Sabal sp. nov.
14. Habit sketch of fungus on a portion of a leaf of Sabal sp.
15. Cross section of leaf showing amphigenous, loculate, angular stromata located
between the epidermis and the mesophyll.
16. Clavate ascus and paraphyses with ascospores inordinately pressed in the
ascus.
17. Two ascospores.

Figs. 18-21. Phyllachora ocoensis sp. nov.
18. Habit sketch of pinnate leaves, leguminous shrub with small punctiform
stromata.
19. Cross section of a pinna of the same with uniloculate stroma in the mesophyll.
20. Ascus and thread-like paraphyses.
21. Two individual ascospores.

Figs. 22-27. Phyllachora panamensis Stevens.

22. Habit sketch of fungus on leaves of Rourea glabra, showing angular stromata.

23. Cross section of leaf showing position of stromata and locules in the meso-
phyll.

24. Ascus and paraphyses.

25. Two individual ascospores.

26. Stylospores (Phyllachora Roureae).

27. Conidia (Ph. panamensis).

CHARDON: FuNcI DoMINGENSES I. 473

2(4):475—525 FARLOWIA Jury, 1946

THE BIOLOGY OF
GYMNOSPORANGIUM NIDUS-AVIS THAXTER

ALTON E. PRINCE !

HISTORICAL RESUME

The history of our knowledge of Gymnosporangium nidus-avis Thaxter
is varied and interesting. This has resulted from several circumstances.
(1) The rust has proved to be heteroecious, and we now know that it
affects many host species in both its aecial and telial phases. (2) The
elucidation of its hosts has proceeded slowly, at times uncertainly. (3)
The manifestations of the hosts carrying the telial phase are confusing,
for while usually they are witches’ brooms with juvenile, subulate leaves,
in other instances they are brooms with adult, scale-shaped leaves, and in
still others they are slight fusiform stem swellings or only simple twig,
branch or stem infections. A still further complication is now added as a
result of studies recorded in this paper which presents evidence that the
heretofore recognized G. juvenescens Kern is not distinguishable from
G. nidus-avis.

The history begins with two collections made by Ravenel along the
Santee Canal, South Carolina, in the spring seasons of 1848 and 1850
respectively. These specimens are now a part of the Curtis collection in
the Farlow Herbarium, and bear the original labels under the designation
Gymnosporangium Juniperi Lk. About thirty years later Farlow (1880)
called attention to them and decided that they were properly G. coni-
cum DC.? Since Ravenel’s time many collections of G. nidus-avis have
been made and there are scores of references to this species in the literature,
but the latter are of interest mainly from the standpoint of geographical
distribution. This resumé, however, will be restricted to a brief survey of
the taxonomic and biological contributions pertinent to the author’s studies.
Thus it includes only references to the works of Farlow, Thaxter, Arthur,
Kern, and the few others who have made additions of some significance.
In this connection, Farlow and Thaxter stand out as the pioneer con-
tributors. 7 |

Farlow (1880) was actually the first to give a description of G. nidus-
avis. This he did in his notable monograph on the Gymnosporangia of
the United States. He described it as an American representative of the
European G. conicum DC. on Juniperus virginiana, but he included speci-
mens of this rust characterized only by fusiform stem swellings and pul-

1 Thesis submitted in partial fulfillment of the requirements for the Degree of Doctor
of Philosophy at Harvard University. |

*The G. conicum DC referred to by Farlow (1880, pp. 23-24) is in my judgment '

what is now known as G. juniperinum (L.) Mart.

475

476 FarLowl4, VoL. 2, 1946

#

vinate telia. In the same paper he made mention of rust brooms on
Juniperus virginiana, but ascribed them to G. clavipes C. & P. The illus-
trations in his monograph (1880, Pl. 2, figs. 22-27), however, represent
perfectly what Thaxter subsequently described under the name G. nidus-
avis.

In a second paper, Farlow (1886a) stated that the broom rust on red
cedar previously referred to by him as G. clavipes likewise produces its
Roestelia on Amelanchier. The same year Farlow (1886b) confirmed the
previous report that his G. conicum was the cause of brooms on Juniperus
virginiana, although his cultures failed to produce aecia on Sorbus (one
of the known hosts of G. conicum in Europe). In spite of this failure
he was unshaken in his conviction that the rust was G. conicum, and he
expressed the belief that additional cultures would demonstrate Sorbus
to be a host. If this had been proven then to be the case, Thaxter prob-
ably would not have continued investigations which later led to the
establishment of a new species.

The work of Thaxter is fundamentally significant because of its biologi-
cal findings. He reported (1887) inoculations of Farlow’s G. conicum
on Amelanchier canadensis,* Aronia arbutifolia, Sorbus americana (Pyrus
americana) and Malus pumila (Pyrus Malus). The Amelanchier only
became infected, and on it were produced both spermogonia and aecia. As
an indication of the extreme variability exhibited by this species, he noted
that he was unable to distinguish the aecia from those of his Roestelia
lacerata y (G. globosum Farl.) even though the former matured two months
earlier than the latter. Thaxter (1889) reported two further series of
cultures. In the first, inoculations were made on Amelanchier canadensis,
Crataegus coccinea, Malus pumila (Pyrus Malus), and Sorbus. Again,
only the Amelanchier became infected and, as before, both spermogonia
and aecia were produced. The second series was restricted to cultures
made on Amelanchier. Following these experiments he stated that the
result was “similar in all respects to that obtained in previous years.”
From these investigations he reached the following significant conclusions:
“In relation to our ‘birds’ nest’ Gymnosporangium, the question of identi-
ties is still further confused by my cultures of this species, made during
three successive years with identical results. The Roestelia obtained

. was referred to R. cornuta, there being no other alternative among
described American Roestelias. . . . It should be noted that upon Sorbus
the ‘birds’ nest’ Gymnosporangium has given no result whatever. Further
careful cultures and observation together with a more extended examina-
tion of European specimens than I have been able to make, will of course
be necessary to confirm this supposition; but I am decidedly of the opinion
that our ‘birds’ nest’ species is a distinctly American form as yet un-
named.”

® The name Amelanchier canadensis should now be referred to A. arborea (Michx. f.)
Fernald, and the name A. oblongifolia, in part, to A. canadensis (L.) Medic. Rhodora
43: 559-567. 1941.

PRINCE: GYMNOSPORANGIUM Nipus-AVIS THAXTER 477

Two years later Thaxter (1891) published a description of the telial
and aecial stages of the new species under the name G. nidus-avis. He
also republished these descriptions in 1892 and 1893. Specimens of Thax-
ter’s cultures of G. nidus-avis are in the Farlow Herbarium and have been
carefully examined by me. The groundwork being thus firmly laid, it re-
mained for others to confirm and extend it.

During the period between 1908 and 1915, Arthur (1908) reported
inoculations on Amelanchier intermedia, and Malus pumila var. Whitney
Crab (Malus Malus var. Whitney Crab). The apple plant only became in-
fected. This produced both spermogonia and aecia on its leaves. In 1910
he reported on additional cultures made in 1909, when Amelanchier cana-
densis, Crataegus Pringlei, and Malus ioensis were inoculated. The Malus
and Crataegus became infected, the former producing spermogonia and
aecia, the latter spermogonia only. The Amelanchier remained uninfected.
In his report of cultures made in 1910, he stated that the fruit of
Amelanchier ovalis (A. vulgaris) and the leaves of Cydonia oblonga
(C. vulgaris) and Malus coronaria were inoculated but only the Amelan-
chier became infected, and it produced spermogonia and aecia. A second
series of inoculations was made later in the same year on identical specific
hosts. That time the Malus alone became infected, but with the pro-
duction of spermogonia only; the leaves died before aecia could be
developed. Finally Arthur (1915) reported his last cultures of this
rust. Amelanchier ovalis (A. vulgaris) was inoculated and both spermo-
gonia and aecia were produced. Arthur (1934) in summary has recently
listed the following names of plants that serve as hosts of G. nidus-avis:
for the aecial stage — Amelanchier canadensis, A. erecta, A. intermedia,
A. laevis, A. oblongifolia, A. ovalis (A. vulgaris), Crataegus Pringlet, Cy-
donia oblonga (C. vulgaris), Malus pumila (Malus Malus); for the
telial stage — Juniperus virginiana.

Dodge was the next important contributor. His most notable work
concerned teliospore formation and production, and this will be reviewed
in the section below dealing with life history of the fungus. He also
(1918b, pp. 294-295) published the results of cultures of G. nidus-avis
made between 1914 and 1917. A species each of Amelanchier and Malus
was inoculated in the spring of 1914 with inoculum from telia borne in
the axils of leaves on brooms, from the stems of brooms, and from sori
on the tree trunk. Some aeciospores from these cultures were used to in-
oculate a potted red cedar plant in June 1914. No external symptoms of
infection appeared in 1915. Two small sori became evident in May 1916.
By 1917 these two sori had coalesced and produced a telial mass two
inches long. This infection spread vertically five inches in three years.
On May 10, 1916 telia were collected in the field for inoculation purposes.
The sowings were made on Amelanchier ‘canadensis,’ May 17, and aecia
were produced June 17. A cedar was inoculated with spores from this
culture on June 21. On March 22, 1917, twenty-one days after being
taken from a cold frame, a telium appeared in the axil of one leaf. The

478 ; FarLowlA, Vo. 2, 1946

next year, on February 19, 1918, two more sori appeared. Thus under
favorable conditions telia may be produced the spring following inocula-
tion. Dodge (1918c) thus found that telial- sori of G. nidus-avis some-
times develop “on the blades of leaves.” Dodge (1931) found that G.
nidus-avis was very destructive to eastern red cedar trees. As emphasized
by him, this is very much in contrast to the general belief that it is only
the aecial host which is injured. According to him (1934) the common
red cedar is subject to attacks by G. midus-avis of such severity as to
cause the premature death of the suscept.

As a matter of minor interest, reference may properly be made to Clinton
and McCormick’s (1924) Petri-dish cultures of G. nidus-avis. Leaves of
Cydonia oblonga (C. vulgaris), Malus ioensis (Pyrus ioensis), Malus
pumila (Malus communis), and Malus pumila var. Wealthy (Pyrus Malus
var. Wealthy) were placed on a liquid medium in Petri-dishes and inocu-
lated with basidiospores of the fungus. Spermogonia developed on the
apple and quince leaves; aecia did not develop because of the unfavorable
method of culture. The other inoculations gave negative results.

Prince and Steinmetz (1940) were the next to investigate G. nidus-avis
when cultures of 1936 and 1937 were reported. In 1936, they inoculated
Amelanchier canadensis, A, laevis, A. stolonifera, A. canadensis «& A.
stolonifera, Malus pumila var. Wealthy (M. sp. var. Wealthy); all except
the apple became infected and produced spermogonia and aecia. In 1937
the plants inoculated were Amelanchier canadensis, A. laevis, A. stoloni-
fera, A. Wiegandii, A. canadensis < A, stolonifera, Cydonia ph ane (C.
vulgaris), Chaenomeles japonica (Cydonia japonica), Malus sp. and
Sorbus americana; all of these except the Chaenomeles, Malus and Sorbus
produced spermogonia and aecia. In addition to their eultuee experiments
they made careful taxonomic comparison of the broom-forming rust of
G. juvenescens on Juniperus horizontalis, with G. nidus-avis on Juniperus
virginiana, making use of their cultures, field collections, and specimens
obtained from various herbaria. No differences could be found in the
spermogonial, aecial or telial stages of these rusts. Hence the morphologi-
cal evidence clearly indicated that G. juvenescens differed in no respect
from G. nidus-avis. But culture work was necessary before a final con-
clusion could be drawn. Should culture experiments confirm the tentative
taxonomic conclusions of Prince and Steinmetz, the addition of Juniperus
_ horizontalis as a telial host of G. nidus-avis is of greater significance than
just the addition of another name to a host list. This is important because
this telial host extends from the Atlantic nearly to the Pacific coast (see map
p. 483) and this is a connecting link between the eastern and western
red cedars, Juniperus virginiana and J. scopulorum respectively, (the latter
being another recognized telial host of the so-called G. juvenescens).

The history of what, up until the present work, has been called Gymno-
sporangium juvenescens Kern will now be added to that of G. nidus-avis.
Arthur (1907) reported the first cultures of the western rust, which at that
time was determined as G. Nelsoni Arth. Telia from Juniperus scopulorum

Prince: GYMNOSPORANGIUM Nipus-AVIS THAXTER 479

were used as a source of inoculum for cultures which yielded spermogonia
on Amelanchier canadensis and Sorbus americana but no aecia were formed.
Crataegus Pringlei, Chaenomeles japonica (Pyrus japonica), and Aronia
melanocarpa (Aronia nigra) were inoculated but no infection occurred.
Arthur (1908) reported two series of cultures of “G. Nelsoni” later recog-
nized by him to be G. juvenescens, using inoculum from J. scopulorum.
In the first series, Amelanchier intermedia was inoculated and spermogonia
appeared, but because of injury to the plant no further development took
place. The second series was started about two weeks later using inoculum
from a second collection of telia on J. scopulorum. Sowings were made on
two plants of Amelanchier canadensis upon which were produced both
spermogonia and aecia.

In regard to these cultures Arthur (1908) made the following state-
ment: “The above results confirm and much extend the somewhat un-
certain work of last season. It leaves no further doubt that Roestelia
Nelsoni Arth. should be counted as a synonym of G. Nelsoni Arth. as
suggested in the original publication of the name.” This statement shows,
more than anything else, the difficulty early investigators had in definitely
placing the extremely variable species under consideration, since G. N elsoni
today is a valid species.

Arthur (1909) reported additional cultures of his “G. Nelsoni” using
inoculum from the usual source. Amelanchier erecta and Sorbus ameri-
cana were inoculated and spermogonia were produced on both hosts, but
aecia only on the Sorbus. Arthur (1912a) reported the last cultures. of
this “G. Nelsoni’” which were conducted during the spring of 1910. The
inoculum used was obtained from Juniperus virginiana, and Amelanchier
erecta, Cydonia oblonga (C. vulgaris) and Malus coronaria were inocu-
lated. Amelanchier was the only host successfully infected and on it
were produced spermogonia and aecia. Arthur (1912b) called attention
to the fact that previous cultures reported under G. Nelsoni should be re-
ferred to G. juvenescens Kern. ©

Kern (1911) described the aecial and telial stages of a broom-forming
rust on Juniperus scopulorum and J. virginiana under the name of G.
juvenescens. As aecial hosts he cites Amelanchier alnifolia, A. florida, A.
oreophila, A. polycarpa, and A. pumila. The distribution of the new species
is given as “British Columbia and Alberta southeast to the Black Hills
of South Dakota and to Colorado, and locally in Nebraska and Wisconsin.”

Arthur (1934) subsequently gave a similar description and range of
G. juvenescens. A number of hosts were added. For the spermogonial and
aecial phases are given Amelanchier alnifolia, A. Bakeri, A. canadensis,
_ A. florida, A. humilis, A. Jonesiana, A. mormonica, A. oreophila, A. poly-
carpa, A. pumila, A. spicata and Sorbus americana; as telial hosts Juniperus
horizontalis, J. scopulorum and J. virginiana.

According to hitherto apparently unpublished data found on specimens
and in letters now in the Farlow Herbarium, the late Dr. J. J. Davis
likewise carried out cultures with G. juvenescens in 1918. It was he who

480 FarLowI!A, VoL. 2, 1946

first recognized that brooms on Juniperus horizontalis were the result of
a rust infection. Using inoculum from this source, sowings were made on
Amelanchier oblongifolia on which both spermogonia and aecia were pro-
duced.

Fraser (1925) inoculated Amelanchier alnifolia with spores of G. juven-
escens collected on Juniperus horizontalis. Spermogonia and aecia de-
veloped.

During the spring of 1935, Steinmetz and Hilborn (Prince and Stein-
metz, 1940) using inoculum from J. horizontalis, which had previously
been determined as G. juvenescens by Dr. J. C. Arthur, inoculated an unde-
termined species of Amelanchier which resulted in the production of
spermogonia and aecia. They were unable to differentiate what was sup-
posed to be G. juvenescens by cultures from G. nidus-avis made on the
same Amelanchier plant. Prince and Steinmetz (1940) reported cultures
made in 1936 and 1937 with inoculum from Juniperus horizontalis on Ame-
lanchier canadensis, A. canadensis X A. stolonifera, A. laevis, A. stolonifera,
A. Wieganditi, Chaenomeles japonica (Cydonia japonica), and Cydonia
oblonga (C. vulgaris), Spermogonia and aecia were produced on all of
these hosts except Chaenomeles japonica. The aecia produced were in all
respects the same as those produced in parallel cultures of G. nidus-avis.
It should be noted that the Cydonia was successfully inoculated because
this host genus was not known previously as a host for G. juvenescens
but was for G. nidus-avis and had served until then as one of the reasons
which justified separating G. juvenescens from G. nidus-avis.

In consideration of the conflicting evidence presented by previous in-
vestigators, it has seemed desirable not only to study further the life
histories of G. nidus-avis and the fungus called G. juvenescens, but also
to make additional extensive inoculations in order to verify and add to,
as far as possible, the work of these investigators. Thus the present
studies were carried out with the hope of adding to the knowledge of
the life history of the organism.

TAXONOMY

The nomenclature of Gymnosporangium nidus-avis Thaxter is probably
less complicated than any other of the widely distributed North American
species belonging to this genus. This is certainly not because of the uni-
formity of symptoms produced by the fungus since no species of Gymno-
Sporangium other than G. nidus-avis causes more than one distinct type
of fasciation of its telial host and in addition produces a variety of symp-
toms on the branches and stems. In the aecial phase it is very similar to
several species occurring on the same aecial hosts, an example of which
was given by Thaxter (1889), when he pointed out the uncertainty of
distinguishing G. nidus-avis and G. clavariaeforme by gross appearance
alone. This and other similarities are in part the cause of the not in-
frequent mislabeling of collections found in herbaria. Therefore it can

PRINCE: GYMNOSPORANGIUM Nipus-AVIS THAXTER 481

be said that the present simplicity in the nomenclature of this species is
undoubtedly because of the thoroughness with which it was studied prior
to its publication by Thaxter, and to the completeness and accuracy of
his first description (1891). The following binomials have been applied
to the rust now known as Gymnosporangium nidus-avis Thaxter. The
phase in the life cycle to which the name was affixed is indicated by the
Roman numeral at the left. |

III Gymnosporangium nidus-avis Thaxter, Bull. Conn. Agr. Exp. Sta. 107: 6. 1891.

Ill G. clavipes C. & P. sensu Farlow, Anniv. Mem. Bost. Soc. Nat. Hist. pp. 21-22.
pl. 2, figs. 22-27. 1880.

Ill G. conicum Hedw. f. in DC. Flora Franc. sensu Farlow, Anniv. Mem. Bost.
Soc. Nat. Hist. pp. 23-24. 1880.

O,1 Roestelia nidus-avis Thaxter, Bull. Conn. Agr. Exp. Sta. 107: 6. 1891.

Ill Puccinia nidus-avis (Thaxt.) O. Kuntze, Rev. Gen. Plant. 3: 507. 1899.

Ill Tremella nidus-avis (Thaxt.) Arthur, Proc. Ind. Acad. Sci. 1900: 136. 1901.

O, I Aecidium nidus-avis (Thaxt.) Farlow, Bibl. Index 1: 68. 1905.

III Gymnosporangium juvenescens Kern, Bull. N.Y. Bot. Gard. 7: 448. 1911.

The reasons for including G. juvenescens Kern as a synonym of G. nidus-
avis will be presented in the sections to follow.

Crowell (1940) considered G. effusum Kern to be synonymous with
G. nidus-avis, and thus has introduced a problem that must be dealt with.
The writer has spent much time searching for G. effusum in South Caro-
lina but has not been successful. However, the type material of G. effusum
from the Arthur Herbarium has been very carefully examined and no close
resemblance could be found between its telia and those of G. nidus-avis.
In G. effusum the telia are elongate, broadly wedge-shaped, furcate, and
according to Kern (1911) may become lacunose. In addition, the telio-
spores are very long, narrow, without an obvious constriction at the sep-
tum, and with no evidence of a slightly enlarged apex of the pedicel,
characters which exclude this species from G. nidus-avis. To the writer,
G. effusum, in its telial phase, is more closely related to the European
G. Sabinae (Dicks.) Wint. than to any American species. Furthermore,
preliminary cultures of G. effusum reported by Arthur (1912b) show that
one alternate host is Aronia, and to check this many inoculations were
made without success with G. nidus-avis on the known species and some
varieties of Aronia. Therefore, because of the distinctive morphology of
the two fungi and because of the differences in host ranges, it is not
possible to include G. effusum as a synonym of G. nidus-avis. My own
description of Gymnosporangium nidus-avis follows:

Spermogonia epiphyllous, hypophyllous or amphigenous, sometimes |
fructicolous, rarely caulicolous in yellowish, yellowish-orange or orange-
yellow lesions, appearing on an average of twelve days after inoculation, —
the average size of the mature spermogonium being 111.7 < 86.7 p.

Aecia following spermogonia on an average of twenty-one days after
inoculation on the same host organs, the lesions 0.5 to 5.0 mm. in diameter

482 FaRLowIA, VOL. 2, 1946

on leaves, usually with 1 to 6 cornute peridia which become up to 2.5 mm.
in length. Peridia rupturing laterally or apically, finally becoming lacerate
to or nearly to the base. Peridial cells typically lanceolate-oblong in face
view remaining straight or nearly so when wet, 29.4-88.2 » in length,
11.7-36.7 » in width, thickness of side walls 2.9-5.8 », inner walls 5.8—
11.7 yw, outer walls 1.4—4.4 w; the outer walls of peridial cells are smooth,
the inner walls with rugose-verrucose sculpturing, and the side walls
rugose with interspersed verrucose markings. Aeciospores globoid to sub-
oval, 14.7-36.7 11.0-27.9 p» in size; walls 0.7-5.8 » thick, colorless to
chestnut brown, smooth to minutely verrucose and usually with 8-12,
rarely with 6 or 14 pores.

Uredia lacking. Telia foliicolous and caulicolous on dense or open
fasciations of branches, or caulicolous on more or less enlarged branch
or stem lesions; the ungelatinized telia pulvinate, hemispheric to oval,
occasionally Higulate. of a color slightly darker than chestnut brown.
Teliospores one to four-celled, usually with two cells, somewhat con-
stricted at the septa, ellipsoid to oval in shape, 19.1-60.2 & 8.8-27.9 pn;
walls smooth, hyaline to dark cinnamon brown, most often pale cinna-
mon brown, 0.2—2.2 » thick with one to three pores usually with an obvious
papilla; pedicels hygroscopic, often enlarging just below the apices, the
enlarged portion occasionally rupturing, leaving the spore with a ragged
fringe or collar.

GEOGRAPHICAL DISTRIBUTION

The geographical distribution of Gymnosporangium nidus-avis is limited
only by the ranges of its telial hosts. This is true because the pomaceous
hosts, according to Rehder (1940), are present throughout northern
America and extend deeply into Mexico, while the Juniperus hosts are
more restricted; therefore the aecial hosts need not be considered as fac-
tors affecting geographical distribution. The hosts of the telial phase of
G. nidus-avis, as is clearly illustrated in the map below, show a nearly
continuous and sometimes overlapping distribution in southern Canada
and the United States, except in the southwestern limits of Juniperus
virginiana and in the southeastern limits of J. scopulorum. Not shown on
the map and only slightly extending the geographic range is the report of
Hedgecock (1940) of G. nidus-avis on a new telial host, J. barbadensis
from near Gainesville, Florida. This report has been confirmed by the
writer who has examined a part of the specimen and has verified the
determination of the fungus name.

On the map (Fig. 1) are shown the geographical ranges of G. nidus-avis
and its telial hosts. The small circles represent the imporiant stations
where collections of the rust were made, and where specimens from these
stations have been examined and verified by the writer, while the crosses
represent stations reported in the literature that have not been verified
but at the same time are believed to be accurate. Whenever possible,

PRINCE: GYMNOSPORANGIUM Nipus-AVIS THAXTER 483

material was obtained to verify reports given in the literature, especially
when unusual characteristics for the specimens were cited.* It is believed,
however, that intensive collecting will show that this rust is in every state
and province within the range of the telial hosts. This assumption is based
on the writer’s observations which have extended over an area from Maine
to South Carolina and westward to southeastern Tennessee and north
central Ohio where it has been found to occur locally in most districts.

4

GEOGRAPHICAL RANGES
OF G.NIDUS- AVIS AND
ITS TELIAL HOSTS

O: Specimens seen
X: From literature

: J. virginiana

——-—: J. horizontalis

FicurE 1. The Geographical Ranges of G. nidus-avis and its Telial Hosts. Juniperus
virginiana L. including its var. crebra Fern. & Grisc. after Munns (1938); J. hori-
zontalis Moench after Rehder (1940) ; J. scopulorum Sarg. after Sudworth (1915).

The local occurrence of this rust, in relation to its wide dispersal, is
of interest and tends to confirm the reports of Kern e¢ al. (1929) who
stated that this fungus is rare in Pennsylvania, while Long (im litt.) and
Coker (1920) respectively reported it as not uncommon around Wash-
ington, D. C., and Chapel Hill, North Carolina. Other reports in cor-
respondence which have come to the writer bear out the above. Mrs. B.
N. Brown was unable to find the rust in the vicinity of Marysville, Ten-
nessee, although L. R. Hesler and the writer have observed this species
in the vicinity of Knoxville, Tennessee. I. L. Conners was unable to find
it around St. Catherines, Ontario; but in Saskatoon, Saskatchewan, the
late Dr. W. P. Fraser indicated that it is rather common. R. W. Bailey was
unable to locate the rust in the region around Ogden, Utah, while A. O.
Garrett reported that it could be found several miles south of Salt Lake

*The report of Ray (1940) of G. nidus-avis from Oklahoma has been found to be
based on a misdetermination and the specimen should be referred to G. globosum Farl.

484 FartowiA, VoL. 2, 1946

City; J. L. Forsburg found it to be not uncommon near Fort Collins, Colo-
rado, as did W. H. Long in the foothills of New Mexico; J. W. Hotson was
unable to find the rust in the region around Seattle, Washington, while
F. D. Heald indicated that this species was known only in its aecial phase
around Pullman, Washington.

The reasons for the localized occurrence of this rust are not at all clear,
especially when both aecial and telial hosts are in the same locality. How-
ever, from observations made during several seasons on Mt. Desert Island,
Maine, it would seem that distance from the telial host limits the in-
fection of Amelanchier. Evidence for this conclusion is furnished by the
fact that although the different species of Amelanchier are common
throughout the island, the only plants infected are within approximately
200 yards of Juniperus horizontalis, which is extremely local, being con-
fined to the rocky ocean bluffs, and in addition is the only telial host.
This is an excellent example of how the fungus may be limited to a small
area, although in this instance the range is determined by the distance of
the aecial host from the telial, and thus depends in part on the duration of
viability in the basidiospores and in part on physiographic factors. This
is in contrast to the findings of Thaxter (1891) and MacLachlan (1935b)
who found that the basidiospores of other species could infect aecial hosts
eight and six miles respectively from the source of inoculum. The writer’s
findings on Mt. Desert Island show that environmental factors may be
more important in limiting the dispersal of this species.

An additional factor tending to limit local distribution is furnished by
the relative period of maturation of the telial sori and the stage of de-
velopment of the leaves of the aecial hosts.

Thus, in the vicinity of Clemson, South Carolina, species of Amelanchier
are often found growing adjacent to cedars that are heavily infected with
the rust. An abundance of telia was produced early in April, 1941, and
gelatinization took place during four days of nearly continuous rain.
At the beginning of this rainy period, several artificial inoculations were
made on Cydonia oblonga growing in a sheltered habitat where the leaves
had already begun to unfold, but at the same time the species of Amelan-
chier growing in the open had not yet started to produce leaves. On the
plants artificially inoculated (Cydonia) the rust developed exactly the
same as it did on this host at the University of Maine and in the Arnold
Arboretum, yet on the species of Amelanchier not a single lesion was found
that had resulted from natural infection. In 1942 and 1944 similar condi-
tions reoccurred, and although artificial inoculations were successful on
both Cydonia oblonga and Amelanchier canadensis, again no natural in-
fections could be found. Here, then, is evidence that the aecial hosts must
be in a condition favorable for infection, and this must coincide with the
germination of the teliospores, all being more or less dependent on tem-
perature and humidity, which confirms the conclusions of Crowell (1940).

While the examples cited above tend to explain that the viability of
the spores, the physiography of an area, and the relative times of de-

PRINCE: GyMNOSPORANGIUM Nipus-AVIS THAXTER 485

velopment of foliage on the aecial hosts and telial sori may indeed play
a part in the local limitation of this species, there are still other factors
that have to be considered. Important among these are the effect of in-
dividual host susceptibility, and the occurrence of strains with varying
degrees of virulence within this species of rust. Although there is no evi-
dence from culture experiments that there are resistant strains of the
Juniperus hosts, yet at any time field observations can be made which
show that resistant hosts exist in quantities. As will be reported later,
there is evidence resulting from culture studies that there exist resistant
strains of the pomaceous or Juniperus hosts or both. In brief it can be
stated that the species has a very wide distribution characterized by areas
in which the rust is relatively scarce and in these areas the paucity of the
species may be explained by one or more of the above sets of factors.

LIFE HISTORY STUDIES

SPERMOGONIAL PHASE

MAcroscoPpicaL DEVELOPMENT

Heretofore very little has been published concerning the macroscopic
symptoms and signs of infections caused by the spermogonial and aecial
phases of Gymnosporangium nidus-avis on its pomaceous hosts. For re-
corded observations we are chiefly indebted to Thaxter (1887, 1889, 1891),
Arthur (1901-1915), and Kern (1911), but for the main part these are
brief and incidental to taxonomic delineations or limited life history
studies of the fungus. They are also incomplete because of an insufficient
number of specimens from the various hosts and geographic range of
the fungus. Kern, in his monograph on Gymnosporangium (1911) was
responsible for the enumeration of specific characters of G. midus-avis
and G. juvenescens, which at that time seemed sufficiently valid from
the materials at his disposal, and which could not be checked from the
life history studies then available. During the course of intensive studies
of G. nidus-avis, the writer assembled a wide range of material from sev-
eral geographical regions and carried out extensive series of culture ex-.
periments. Therefore it is possible at this time to give a more complete
and connected account of the symptoms and signs of infections caused
by this fungus on its pomaceous hosts than could have been done previously.

The first indication of infection, regardless of the organ affected, is a
light spot or lesion in the host. These lesions soon change to a distinct
yellow, then to a yellowish-orange which is more or less constant for most
hosts. The lesions soon attain their definitive size and remain thereafter
distinct from the adjacent uninfected host tissues, whether on leaf, stem
or fruit. Subsequently, with the formation of spermogonia, the lesions
lose their yellow color and become either orange-yellow or necrotic and
black. Spermogonia appeared in from six to fourteen days in the writer’s
cultures. These organs occur singly or in a group of two to ten or more on

486 . Fartowla, Vor. 2, 1946

a single lesion. They are usually epiphyllous, occasionally hypophyllous or
amphigenous on different lesions of the same leaf (Plate I, Fig. 3). In
the early stages of development, the spermogonia are a waxy, orange-yellow
on the lighter colored lesions. The waxy, colored appearance is caused
by the colored spermatia in a viscous spermogonial fluid. The exudate
is usually copious for the first five to ten days, during which time the
general macroscopic symptoms normally remain unchanged, when flies
and ants can be observed feeding on the exudate that has been shown by
Rathay (1883) to contain sugars. As demonstrated by Craigie (1927),
flies, attracted by the sweet spermogonial exudate, bring about cross-
spermatization in Puccinia Helianthi and it seems probable that the in-
sects mentioned above must perform the same function for this rust. It
is probable also that the spermatia are transferred by water.

Following cross-spermatization, the exudate rapidly diminishes in amount
external indication that cross-spermatization has been completed, after
which the exudate dries up entirely. During the period when spermatia ©
are being produced, if the spermogonia are carefully examined, a definite
protuberance can be seen emerging from the ostiole of each of these organs.
These protuberances are made up of ostiolar filaments,

Along with the usual sequence of events in the ontogeny of the spermo-
gonia, two distinct variations occur during the development of the in-
fections. The degree of variation is dependent upon the host involved.
Thus on Amelanchier alnifolia and A. intermedia, instead of the lesions
being the usual yellowish-orange, they are reddish-orange with a yellow-
ish-orange margin. The other important variation is the necrosis of lesions.

Necrosis as it is found in the lesions is variable and dependent upon
the susceptibility of the host under attack. On the very resistant hosts,
the lesions become necrotic soon after they develop. Indeed, this often
takes place before the production of the spermogonia. However, occa-
sionally some spermogonia are formed but these are small, and probably
non-functional as is indicated by their failure to produce an exudate.
Necrosis of the lesions occurred on the resistant and susceptible host
species, like Chaenomeles superba and Amelanchier stolonifera, but on
these it often may not take place until after the spermogonia and aecia
are produced. When aecia are formed on these hosts some at least are
non-functional, as is shown by the fact that the aeciospores produced did
not germinate in tests. A very striking type of necrosis appeared on
Sorbus scopulina, where the main part of the lesions remained alive; but
a narrow zone at their periphery died.

MIcROSCOPICAL DEVELOPMENT

Since the lesions and subsequent symptoms produced on the pomaceous
hosts are the result of infection by the wind-carried basidiospores from
the telia on the cedars, it seems desirable to follow more exactly the process

PRINCE: GYMNOSPORANGIUM Nipus-AVIS THAXTER 487

of infection and to study the subsequent development of the fungus within
the host tissue.

According to MacLachlan (1935b) and other investigators, the basidio-
spores of Gymnosporangium species are carried by wind during humid
spring weather. During this period the leaves of the alternate hosts are
just beginning to unfold, and are often in their most susceptible condition.
Therefore, when the basidiospores reach the host, germination proceeds,
after a short delay, by means of germ tubes.

The germ tubes of the basidiospores may penetrate directly after ger-
mination of the basidiospores, or the germ tubes may grow to some length
before they pass into the host, without the formation of appressoria, by
penetrating the cuticle and becoming established intercellularly. From this
it seems that the germ tubes must secrete a substance which dissolves the
cuticle, thus permitting them to pass through this layer. Certainly the
absence of appressoria seems to preclude the possibility of the fungus enter-
ing the host as a result of pressure exerted by the germ tubes. Infection
takes place between twelve and forty-eight hours after inoculation, or
perhaps even earlier under the most favorable conditions. The hyphae
grow rapidly between the cells and the rust soon becomes thoroughly estab-
lished in susceptible hosts, The intercellular hyphae vary considerably,
but are usually 1.5-4.5 » in diameter; they may often entirely fill the
intercellular spaces and frequently appear to isolate an individual host
cell from those adjacent. This fungus becomes intracellular only when
haustoria are formed. In any stage of development after the fungus has
become established, haustoria can be found in nearly all of the host cells
in the lesion, large cells often having two or more haustoria which vary
in shape from clavate, slightly curved to spiral.

Following the establishment of the fungus within a leaf, the spermogonial
initials appear as knots of hyphae between the upper epidermis and the
palisade layer or between the lower epidermis and spongy parenchyma
cells. In this stage of development, two to six days after inoculation, the
rust hyphae were seen to extend between the cells to a distance of a half-
millimeter from the hyphal plexus. During subsequent development some
of the hyphae in the center of the spermogonial initials elongate and appear
to act as buffer cells which force the inner cell walls of the epidermis against
the outer, and as a result of this pressure, the epidermis is forced outward.
The buffer cells are soon replaced, however, by other fungal elements that
arise from the young spermogonium. The writer’s investigations indicate
that the buffer cells play no part in the actual rupturing of the epidermis.
This is accomplished by the continued growth of the spermogonium, while
buffer cells seem to break down to make room for other developing spermo-
gonial structures. The organ soon becomes differentiated into the spermo-
gonial wall, from the bottom of which arise the spermatiophores and re-
ceptive hyphae, while at the same time the periphyses develop from the side
wall. The spermatiophores are specialized, short hyphae, their function
being to produce at their distal ends spermatia which are yellowish-orange,

488 FarLowlA, VoL. 2, 1946

ellipsoidal, and vary from 5.8—-8.8 x 0.7-2.9 ». The periphyses are hyaline,
septate, subulate structures which vary in length from 58.8-133.8 uw. In
addition to the periphyses are the receptive hyphae, but the latter structures
arise from deep in the bases of the spermogonia instead of from the sides.
The receptive hyphae are yellowish, uniform in diameter, non-septate, and
have rounded apices (Plate I, Fig. 5) to which, as the writer has observed,
spermatia fuse, but there was no evidence of the sub-cuticular receptive
hyphae described by Allen (1930, 1932).

By the time the spermogonium ruptures the epidermis of the host, sper-
matia have already been produced in the spermogonial cavity. Spermatia
may be liberated as early as six days after inoculation although the usual
period is about twelve days, by which time the spermogonia are sub-

has been completed, the spermogonial cavity becomes filled with elongated
spermatiophores and dried up spermatial exudate, histological evidence
that these organs have fulfilled their function.

The spermogonia have been used with success by Hunter (1936) as an
aid in determining the various species or genera of the Melampsoraceae.
With this in mind, it seemed desirable to make a comprehensive study of
the spermogonia of G. nidus-avis and of G. juvenescens since the latter spe-
cies has been accepted up to the present time. As is shown by measure-
ments of one hundred medial sections, these organs vary in size from 80.8—
171.9 » high by 64.7—-132.3 w in diameter, the average size being 111.7 x
86.7 ». The above one hundred measurements included fifty each of G.
nidus-avis and G. juvenescens. The average dimensions of the G. nidus-
avis spermogonia were 111.5 x 84.8 u while those of the G. juvenescens were
111.5 x 88.9 ». The evidence thus furnished by the nearly identical size and
morphology of the periphyses, spermatia, and the size of the spermogonia,
since these structures are relatively large, may accordingly be considered
as showing only minute variations. Furthermore, the time of appearance of
the spermogonia is the same in both species, and for this and the preceding
reasons, G. juvenescens is considered to be the same as G. nidus-avis.

Farlier in this chapter I have described the external symptom of necrosis
of some lesions. The histological symptomatology of the necrotic lesions
was studied in detail. This was undertaken because it became necessary to
determine whether or not necrotic lesions appearing on very resistant hosts
were caused by G. nidus-avis or some other fungus. Even preliminary ex-
aminations showed that necrotic lesions caused by the rust fungus had what
I have chosen to call an invasion zone (Plate I, Fig. 4). The invasion zone,
showing the reaction of the host against the rust, is characterized by having
a layer of host cells which are filled with a resin-like material that is vellow-
ish in unstained sections, but in stained preparations takes dyes readily and
stains deeply. The resin-like material appears the same in all respects as
that to be found in the leaf spots where it is scattered throughout the in-
fected area, whereas in the invasion zone of the necrotic lesions it is con-
centrated. Within the infected area of the leaf tissue the host cells may be

PRINCE: GYMNOSPORANGIUM Nipus-Avis THAXTER 489

entirely dead while those of the healthy leaf tissues outside of the invasion
zone are alive and have the usual arrangement of chloroplasts. Without
exception, when the invasion zone was evident around necrotic lesions, rust
hyphae could be found, but when the invasion zone was lacking, typical
rust hyphae could not be found. Therefore, necrotic lesions on hosts that
lacked invasion zones were considered to be caused by agents other than
G, nidus-avis. Correlated with this is the fact that when there were necrotic
lesions without there being an invasion zone on inoculated leaves, similar
lesions also occurred on uninoculated leaves. Thus by using the presence
of the invasion zone as a criterion for the presence of G. nidus-avis, it was
possible to exclude necrotic spots caused by other fungi.

As others have previously suggested, plants are able in some degree to
inhibit or prevent the growth of fungi. The presence of this invasion zone
in the resistant plants would, then, appear to be a reaction by the host to
prevent or limit the spread of the fungus since no hyphae were found be-
yond this barrier, whereas in susceptible host species in which no invasion
zone is formed, the fungus hyphae can be seen well beyond the limits of
the lesion.

AECIAL PHASE

MAcrROSsCcOPICAL DEVELOPMENT

The aecial phase starts at about the time the spermogonia are complet-
ing their function and becomes evident as a result of more or less hyper-
trophy accompanied by a change in the color of the affected host parts.
The color of that part of the lesion from which aecia finally emerge is
nearly always yellow, but frequently when the spermogonia are epiphyllous,
the under side of the leaf does not turn yellow until the formation of the
aecial fundament which is usually accompanied by hypertrophy. The
amount of hypertrophy varies in degree and occurrence according to the
relative susceptibility of the host, and according to the organ involved. In
the more or less circular leaf lesions, no enlargement of the host tissue takes
place until the aecia begin to form, at which time there is a slight swelling
of the lesion from which a single aecium or several aecia finally emerge.
The swelling is absent on resistant hosts where there is no indication of
aecia even though spermogonia are present. Some hypertrophy is evident,
however, if aecia do develop on resistant hosts; it is smaller when the aecia
remain embedded, and larger when they emerge.

There are two general types of leaf lesions, their shape depending on their
relative position with respect to the major veins. Most of the lesions occur
in the thin parts of the lamina where there are no major veins and the size
is influenced by the susceptibility of the host, it being smaller on the more
resistant species and larger on the more susceptible. On Amelanchier stolon-
ifera, a susceptible species, the spots vary from 0.5—2 mm. but on Amelosor-
bus Jacki (Plate I, Fig. 1), a very susceptible species, their diameters vary
from 2-5 mm. When lesions occur on leaf veins, there is always some

490 FaRLOWIA, VOL. 2, 1946

hypertrophy, with the vein as its long axis. These swellings are usually
hypophyllous, but may be evident on the upper leaf surface. A severe
vein infection often causes a distortion of the leaf (Plate I, Fig. 2), and in
addition the aecia may be amphigenous. Likewise, on affected fruits, the
amount of hypertrophy is not readily assessed. Infected fruits are not much
larger than the uninfected, but they are more or less distorted. When
young stems become infected, there is also more or less distortion and a
conspicuous enlargement (Plate I, Fig. 2). No comparison has been made
in this discussion of the macroscopic symptoms of G. nidus-avis and G.
juvenescens, because there were no variations in the developmental stages
of the two species that could be distinguished.

MrcroscopicAL DEVELOPMENT

The nuclear phenomena which occur during the transition from the
spermogonial phase to that of the aecial phase have not been entirely
worked out for any rust. Unfortunately the writer has not been able to
contribute to the elucidation of this phenomenon, but it is postulated that
cross-spermatization is followed by dicaryotization. This latter process
may begin when the haploid nucleus of the spermatium has passed into
the receptive hypha, and end when the haploid aecial fundament becomes
dicaryotic. It is then that the aecial phase is initiated. This stage marks
the beginning of a period of rapid development of the aecia which is cor-
related with a reaction of the host that is almost immediate. The apparent
host reaction is to increase the size and number of the cells of the lesion.
In this stage of development, hyphae are discernable considerably beyond
the limits of the infection zone where there is no visible host reaction.

Aecial primordia, each consisting of a plexus of haploid hyphae, appear
about the time the spermogonia are mature, are formed just below the
palisade cells in the leaves. Aecial primordia appear about six days after
cross-spermatization has been completed or about twenty-four days after
inoculation. ;

The formation of the hyphal plexus is accompanied by a breaking down
of host cells in the area, followed by the development of large thin-walled
cells approximately at the center of the primordium. The aecial hymenium
soon becomes visible in the lysigenous cavity formed by the dissolution of
the thin-walled interwoven hyphae. The peripheral and apical cells appear
as the initial peridial cells (Plate I, Fig. 6). The elongating peridium ap-
pears not to force its way out of the hypertrophied host tissue but to follow
a path of rust hyphae which in turn seem to break down the host cells in
advance of the apex. No evidence was found that would indicate that the
host cells were forced to one side by the apex of the peridium or that the
epidermis was actually split to allow the peridium an exit.

By the time the aecium reaches the surface of the host, the apical por-
tion is filled with a mass of loose spores. Only those in the process of forma-
tion remain attached to the clavate sporophores, The peridium and the
enclosed structures are all dicaryotic, whereas the hyphal plexus, which

Prince: GYMNOSPORANGIUM Nipus-AVIS THAXTER 491

surrounds the peridial envelope, is for the most part haploid. Since it is
formed before dicaryotization, this hyphal plexus is considered to be of a
more or less stromatic nature not properly belonging to the aecium. How-
ever, among the haploid cells there are some scattered hyphae which sec-
ondarily have become dicaryotic.

A careful study was made of the development and morphology of the
peridial cells and aeciospores because these structures were supposed to
show differences which enabled Kern (1911) and Arthur (1934) to distin-
guish between G. nidus-avis and G. juvenescens. The differences were to
be found in the method of peridial dehiscence, in peridial cell morphology
and aeciospore size.

My own observations show that the peridia, regardless of the origin of
‘noculum used for the cultures, may rupture apically soon after emergence,
or retain their cornute form until after elongation has been completed. This
phenomenon appears to depend upon whether or not they are alternately
wetted and dried or whether dehiscence is retarded during a dry season.
Cultures made with western material of G. juvenescens in Maine during
1938 developed exactly in the same manner as did the indigenous rust,
G. nidus-avis. The peridia dehisced apically soon after their emergence and
became lacerate to, or nearly to, the base. The same method of dehiscence
occurred with the cultures that matured the latter part of June and early
July 1939, in the Arnold Arboretum. However, cultures, regardless of the
origin of the inoculum, that matured later during almost drought conditions,
did not dehisce at all or only after collection. When the cornute form is
retained by the peridia, rupturing first occurs laterally, followed by the
rupture of the apex, after which the peridium becomes lacerate, its parts
usually remaining straight or only slightly divergent. In some cases only
the apex ruptures and the peridium retains its cylindric form. It was
found that such peridia were made up of cells with finer markings than
those peridia which become more lacerate. These findings do not agree
- with those of Kern (1911) and Arthur (1934) who reported that in G.
nidus-avis the peridium ruptures apically and becomes lacerate to the base,
while in G. juvenescens the peridium is tardily dehiscent at the apex, the
rupturing first occurring laterally, and finally the peridium becomes lacerate
nearly to the base. Because of the reported variation in the method of
peridial dehiscence, an attempt was made to explain this by a careful study
of the morphology of the cells of the peridium.

Kern (1911) described the peridial cells of G. juvenescens as “lanceolate-
oblong in face view, 16-23 x 65-110 p, linear-rhomboid in side view, 17—
27 » thick, outer wall rather thin, 1-1.5 p, smooth, inner and side walls
medium thick, 5—7 p, finely and closely verrucose-rugose with irregular and
oftentimes ridge-like papillae.” Arthur (1934) added that the cells remain
straight when wet. Kern (1911) described the peridial cells of G. nidus-
avis as “lanceolate in face view, 15-23 x 55-88 yu, linear in side view,
14-18 » thick, outer wall 1—-1.5 » thick, smooth, inner and side walls 5—7 u
thick, coarsely rugose with narrow ridges, with shorter often roundish papil-

492 FarRLowIA, VoL. 2, 1946

lae interspersed.” Arthur (1934) again added the important character that
these cells remain straight when wet.

Fischer (1891 and 1895) and Kern (1910) pointed out that peridial
cell markings were very important in differentiating between species of
Gymnosporangium. I have made extensive observations on many different
species of this genus and have likewise found this character to be the most
dependable for taxonomic delineations.

The apparent differences between the peridial cells of G. nidus-avis and
G. juvenescens as stated above are in the length, thickness, and wall mark-
ings. Accordingly, an attempt was made to confirm the constancy of these
characters. Samples of peridial cells from cultured material were measured.
The inoculum used to produce the aecial specimens originated from Sas-
katchewan, Colorado, Tennessee, Maine, Massachusetts, North and South
Carolina. The dimensions measured were length, width, thickness and
thickness of the side, the inner and the outer walls. Extreme variation was
encountered in the lengths and widths of the peridial cells from all of the
regions listed. Only cells from the side walls of the peridia with typical
oblong-lanceolate shape were measured. The results of these measurements
are recorded (Table 1) according to the Juniperus host from which the
inoculum originated to produce the fungus.

TABLE 1. Data ON PERIDIAL CELL DIMENSIONS OF G. nidus-avis TABULATED ACCORDING
TO THE ORIGIN OF THE INOCULUM USED TO OBTAIN THE AECIA

Origin of Length Width Thickness Wall Thickness
Inoculum bh a mn Side u Inner“ Outer u

J. horizontalis Sask. 29.4-88.2 11.7-25.0 14.7-36.7 2.9-5.8 5.8-8.8 1.4-4.4
J. horizontalis Maine 29.4-88.2 11.7-25.0 11.7-36.7. 2.9-5.8 5811.7 1.44.4
J. scopulorum Col. 35.2-88.2 -11.7-22.0 11.7-30.2 2.9-5.8 7.3-10.2 1.44.4
J. virginiana Mass. 36.7-88.2 14.7-23.5 11.7-33.8 = 2.9-5.8 7.3-11.7 1.4-4.4

No significant differences are evident that would separate the eastern
from the western rusts. Fischer (1891) and Arthur (1934) used only the
outer and inner wall measurements; therefore additional measurements
were made using only those dimensions. This time cultures were obtained
from inoculum from Maine and Saskatchewan on Juniperus horizontalis,
Colorado on J. scopulorum, and Maine and Massachusetts on J. virginiana
(Table 2).

TABLE 2. DATA ON OUTER AND INNER PERIDIAL CELL WALL MEASUREMENTS TABULATED
ACCORDING TO THE ORIGIN OF THE INocULUM USED To OBTAIN THE AECIA

Wall Thickness

Origin of Jnoculum Outer 4 Avg. w Inner wu AVE. &
J. horizontalis Maine ........ 4.4-14.7 8.5 0.7-4.4 25
J. horizontalis Sask. .....00¢, 58-147 8.9 0.7-4.4 2.6
J. scopulorum Colo. ......... 5.8-14.7 9.2 0.7-4.4 2
J. virginiana Maine ........ .. 4.4-14.7 8.5 0.7-4.4 2.5
J

. uirginzanga Mass. .....,,--:: 4.4-14.7 8.4 0.7-4.4 oF

PRINCE: GYMNOSPORANGIUM Nipus-Avis THAXTER 493

Measurements of the thickness of the walls also confirm the data in
Table 1, and hence these results offer further substantiation of the thesis
that G. nidus-avis and G. juvenescens should not be distinguished.

Since the above biometrical studies show no reason for the separation
of the two species, the investigations were carried further by careful com-
parisons of the sculpturing on the peridial cells. The usual markings ob-
served on peridial cells regardless of the origin of the inoculum has been
found to be rugose-verrucose. The side walls are rugose with some verru-
cose markings interspersed. The inner walls are rugose along the sides,
these rugae being extensions from those on the side walls. The centers of
the inner walls are usually covered with short ridge-like papillae and verru-
cose markings. The outer walls are smooth. The more resistant hosts
tended to produce aecia with more verrucose-rugose peridial cells than were
found on susceptible species. However, this was not constant enough to
indicate an expression of the effect of the host reaction on the morphology
of the fungus.

Just as the preceding peridial characters were studied, so have those fur-
nished by the aeciospores. Since the aecia of both G. nidus-avis and G.
juvenescens could be produced by inoculation on similar hosts, further
attempts were made to distinguish the species by their spore characters.
Therefore measurements were made of the spore dimensions and the thick-
ness of the walls, while at the same time counts were made of the number
of pores, and notations were made of spore wall color and markings.

No differences could be found in the color of the walls. The variation,
regardless of the origin of inoculum ranged from hyaline through cinna-
mon to chestnut brown or even darker, although some peridia were found
to contain nearly all hyaline spores. The number of pores were counted
only when they could be seen distinctly and it was found that the usual
number of pores per spore varied from eight to twelve, although very rarely
spores with six or fourteen pores were found.

Similarly, other characters to be found in the aeciospores regardless of
the origin of the inoculum show no significant differences which would war-
rant separating the two species. Thus the shapes of the spores range
usually from globoid to suboval although in any sorus it is possible to find
abnormal spores which are easily distinguished from those characterized as
normal. Just as the normal shape of the spores varies within relatively
narrow limits, so do the surface characters of the aeciospore walls. A
large series of observations on this point leads the author to conclude that
the spores may vary from smooth to minutely verrucose. Not only is there
this variation in the spores from the same aecium, but even using the same
spores, the appearance of the markings may vary with the mounting me-
dium used, — in lactophenol the markings stand out when present whereas
when the spores are mounted in water, the minutely verrucose spores may
appear smooth. Although the presence or absence of markings on the walls
of the aeciospores has served various authors for separating G. nidus-avis
from G. juvenescens, the present observations demonstrate that such dif-

494 FartowliA, VoL. 2, 1946

ferences are not significant. The thickness of the aeciospore walls is rela-
tively constant regardless of the origin of inoculum used to produce the
spores. On the basis of 408 measurements of both eastern material of G.
nidus-avis and of spores cultured from Colorado and Saskatchewan inocula,
it was found that the thickness of the walls of the former varied between
0.7 and 6.5 » but with a mean of 2.99 », whereas walls of the aeciospores
which were produced from western inoculum varied from 1.0-5.8 » with
a mean thickness of 2.7 p, or a difference of only 0.29 « between the aver-
ages. This difference is so small that it is not considered to be of any
taxonomic value for the separation of the eastern rust from the western.

Since other characters of the aeciospores have proved of little value in
separating the two species, a careful determination was made of the aecio-
spore dimensions. It was believed that these, if anything, might enable
one to distinguish the two species in question by following the examples set
by Levine (1923) who was able to distinguish physiological races of a rust,
and Colley (1925) who separated two closely related species by the use
of biometry. For these biometrical studies, notes and data were kept sep-
arate for all the kinds of inocula used throughout the cultural investigations.
The data in Table 3 are based on aeciospores produced from eastern inocu-
lum collected on J. virginiana var. crebra in Massachusetts, J. virginiana
from North Carolina, South Carolina, and Tennessee, while the western
inoculum was from J. scopulorum growing in Colorado and J. horizontalis
growing in Maine and Saskatchewan. Since the inoculum from all sources
was used on the same pomaceous hosts to obtain the aecia, it is believed
that any differences between the eastern and western inocula would become
evident. In Table 3 not only is there a comparison of the dimensions of
aeciospores produced by the eastern and western inocula, but also differ-
ences within each of these series.

TABLE 3, AECIOSPORE SIZE IN RELATION TO THE GEOGRAPHICAL ORIGIN OF INOCULUM
Usrep FoR AECIAL CULTURES

Origin of No. of Range of Mean Range of Mean

Inoculum Spores Lengths u Lengths u Widths « Widths u
1 DTC ee oe earn nes $00 17.0-32.0 23.4 14.0-26.0 20.3
WIRED ce ea acess ae 500 17.0-33.0 22.9 11.0-26.0 19.6
OUASS) oe eect a se ee 400 16.1-30.8 22.6 13.2-23.5 17.7
Maine ..:..6:.. 400 ‘17 .6-33.8 22.1 14.7-25.0 19.6
Dace en ot ee 400 14.7-35.2 23.5 11.7-27.9 19.3
Eloy etet at ae 400 14.7-36.7 Zeal 13.2-25.0 18.4

Average ........ 14.7-36.7 227 11.0-27.9 19.1

In this comparison it can be seen that the ranges in the lengths and
widths of the two sets of spore measurements are very similar, but what is
more important is that the differences in mean spore lengths and widths are
insignificantly small, the differences in the mean lengths being only 0.5 p,
while in the mean widths the variation between spores produced from the
eastern and western inocula is only 0.7 ». An analysis of the data in Table

Prince: GyMNOSPORANGIUM Nipus-AVIS THAXTER 495

3 shows that the greatest difference in the average spore lengths is between
the spores produced from the Colorado and Saskatchewan inocula, the dif-
ference being only 1.5 ». In the mean spore widths the greatest difference
of 1.9 » is between the aeciospores cultured from the Maine and Massa-
chusetts inocula. In consideration of the large size of the spores, the varia-
tions seem of little consequence, especially since in a comparison of the
aeciospores from the eastern inocula with those from the Maine and
Massachusetts inocula, the difference between the mean lengths was
equally as small, 0.5 », while in the spore widths the difference is 1.9 u
compared to 0.7 ». Similar differences exist between the spores produced
from the Saskatchewan and Colorado inocula. It is now possible to con-
clude that the aeciospore dimensions have no character which might permit
the separation of G. juvenescens from G. nidus-avis. However, the minute
tendencies toward variation do indicate the beginnings of biological special-
ization which becomes very evident when host susceptibility is taken into
consideration, a subject which will be discussed in a following section.
Here again is emphasized the very close similarity of the eastern and
western rusts. Thus it is shown here as in the preceding paragraphs of
this section that there are no differences either in the spermogonial or in
the aecial phases which might be considered large enough to delimit G.
juvenescens from G. nidus-avis. The careful comparison of the western
and eastern rusts was carried to the telial phase which is to be considered
in the section to follow.

THE TELIAL PHASE

MAcROSCOPICAL DEVELOPMENT

Gymnosporangium nidus-avis has been confusing as a species not only
in its aecial phase but also in its perfect or telial stage because of the
variability in its manifestations on various species of Juniperus. The ex-
treme in variation is shown in a statement by Arthur (1910) when he de-
scribed a collection of stem inhabiting telia from South Carolina as having
an appearance like a large Tremella and which was not accepted as a rust
until it was examined microscopically. As the result of extensive investi-
gations and observations, the writer has been able to coordinate the some-
times very diverse points of view and to extend the knowledge of the telial
phase of G. nidus-avis. Observations have been made from Maine to
South Carolina and west to Tennessee and Ohio and in addition numerous
herbarium specimens from the entire range (Fig. 1) of the rust have been
carefully examined.

The most characteristic symptom caused by the fungus G. nidus-avis is
the production of a fasciation of branches which when seen from a dis-
tance resemble a bird’s nest. These fasciations, being more commonly
known as brooms, appear to have their origin when a terminal or lateral
growing shoot becomes infected.

In this respect there are three expressions of fungus infection: the

496 Fartow14, VoL. 2, 1946

first a very dense broom with the subulate or juvenile form of leaves
(Plate II, Fig. 1), that may reach twenty-four inches or more in diameter,
and become fifteen or more years old. Many of the branchlets, arising
from about the same place on the main branch or stem, remain slender
throughout the life of the broom and retain most of their acicular leaves.
The second form of broom is similar to the first except that the foliage
remains normal. They do not appear as compact because the leaves
remain Closely appressed to the branches (Plate II, Fig. 2). Just why
some brooms have subulate and others scale-shaped leaves is not clear, but
both forms of brooms have been seen on the same tree and rarely a broom
with acicular leaves may be found with one branch retaining its normal
foliage. The third symptom of infection is different in general aspect from
the first two, it being characterized by the fewer, more widely separated
branchlets which retain their scale-shaped or sometimes subulate leaves
only at their extremities. The branchlets become somewhat enlarged in
diameter and very rough. The first symptom of infection occurs com-
monly on Juniperus horizontalis, J. scopulorum (Plate II, F ig. 4), J. vir-
giniana var. crebra® (Plate II, Fig. 1) and occasionally on J. virginiana ;
the second on all of these hosts, while the third type is the common one on
J. virginiana, although sometimes it occurs on J. virginiana var. crebra.

The fungus does not confine itself to the brooms but often invades the
supporting branch or stem and the mycelium may extend several feet
away from a broom. In addition, branches (Plate II, Fig. 3) and old
stems not associated with brooms may be infected. When branches and
stems are attacked by G. nidus-avis, the effect on the morphology of the
host may or may not be externally apparent. Small young branches and
stems may be slightly enlarged in diameter, the bark is usually very rough
with deep longitudinal and transverse fissures, or occasionally, as Dodge
(1918b) has reported, the bark may actually be split open. Sometimes,
however, young infected branches do not show enlargement, and the only
external evidence of the fungus is the presence of telial masses or the
oval scars left in the outer bark just after the telia fall. When infected
stems and branches become older, there is considerable enlargement, and
the deeply furrowed bark of the infected area may be seen. Hartley (1913)
reported an exceptionally large trunk lesion six feet eight inches long, and
another lesion where the fungus had continued to spread for thirty-two
years after the bark in the center had died. Similar symptoms are not
uncommon on old trees that have been infected for many years.

As there is a difference in the characters of brooms produced on the
telial host, there is also a difference in the occurrence of telia on branches
and stems. From observations and specimens examined, it has been found
that on Juniperus virginiana the telia usually occur on leafless parts of
branches and stems and this is apparently also true on J. barbadensis,

°The variety of Juniperus virginiana can be distinguished from the true species with
characters described by Fernald and Griscom (1935).

.

PRINCE: GYMNOSPORANGIUM Nipus-AVIS THAXTER 497

since the only collection of G. nidus-avis on this host that has been made
was on a branch. On J. virginiana var. crebra and J. horizontalis the telia
occur on both leafy branchlets and the older leafless branches and stems,
while on J. scopulorum no telia have been seen that were not associated
with a broom. This latter was at first thought to be a coincidence, but a
careful examination of all available specimens, both fresh and dried, failed
to give any evidence that telia were formed away from the brooms, and,
therefore, it was concluded that such telia do not occur, or if they do,
only rarely. Long (im litt.) has confirmed this conclusion from his observa-
tions of this rust on J. scopulorum.

MicroscoPICcAL DEVELOPMENT |

The external alterations in the host morphology are not usually a true
indication of the extent to which the rust has spread through a given host.
For this reason it is necessary to examine sections of the host microscopi-
cally to determine whether or not rust hyphae are present. Such investiga-
tions show that the mycelium is present throughout the host tissues of the
dense brooms and the large, yellowish-orange hyphae can be found through
the tissues of the bark where, in stems one to five years old, the intercellular
spaces of the cortex are much larger than in uninfected stems, thus making
this region somewhat thicker than normal. These intercellular spaces are
often crowded with rust hyphae which are also in the phloem, and cambial
region, and not uncommonly extend into the xylem along the medullary
rays, by which they may reach the pith, and in addition hyphae can be
seen occasionally in the tracheids. Hyphae often can be found also in the
subulate leaves of brooms, but even when brooms have a normal foliage,
the leaves are frequently invaded by the fungus. The mycelium may also
extend several inches away from the broom into the supporting branch
or in advance of a branch or stem lesion without causing external symptoms.
During the invasion of the host, the fungus mycelium spreads more rapidly
longitudinally along the main axis of the branch or stem than transversely.
The rate of spread longitudinally, as shown by Dodge (1931) is at least
six inches in three years.

The effect of the fungus on its telial hosts having been considered, a
treatment of vegetative and reproductive stages of the rust, G. nidus-avis,
in its telial hosts will be discussed in the paragraphs to follow.

The mycelium is yellowish-orange and very irregular in size but usually
3—5 w in diameter, penetrating and branching between the host cells, send-
ing the small clavate haustoria into the parenchyma cells of the host. The
hyphae can be seen also in the intercellular spaces of the cortex on young
stems and the spongy parenchyma of the leaves. At any time of year, but
most commonly. late in the fall, large knots of hyphae are produced which
represent the earliest beginnings of the telial primordia. Their presence
may be indicated externally by a slight discoloration, but usually they are
invisible to the unaided eye. There seems to be a slight amount of growth
in the hyphal knots which may well occur in brief warm periods during

498 Fartowla, VoL. 2, 1946

the winter. In early spring, definite telial primordia can be seen under
the phellogen of the stems and the epidermis of the leaves, and these
rapidly mature during the moist warm weather so that the telia are fully
developed at about the time the leaves of the alternate hosts are being
formed, the exact time of year varying considerably within the geographic
range of the fungus — by early March in Florida, but not until May or
early June in Maine. It is especially interesting to note that in both Fort
Collins, Colorado, and Saskatoon, Saskatchewan, these sori are completely
developed in early May. The telia appear from the nodes, internodes and
leaves on the dense brooms, regardless of host or whether the foliage is of
the scale-like (Plate II, Fig. 2) or acicular form, On the open stag-horn
brooms, however, they are mostly on the older leafless parts of the branch-
lets where the telia occur irregularly (Plate II, Fig. 3). This is also true
as to occurrence and position of the sori on the leafless parts of branchlets
in the dense brooms, and on branches and stems not connected with a
fasciated portion of the host. On young branches with smooth bark, the
telia emerge at the margins of telial scars of previous seasons or cause new
lesions, while on older branches and stems, the telia emerge from between
the fissures of the rough bark.

In reference to the characters of the telial sori, shape, size, and color
are of significance taxonomically. The first two are the most constant,
but while shape should be considered both before and after gelatinization,
size and color are of importance only before this takes place. The telia,
when in the fresh ungelatinized condition and when they occur on the
leafy, fasciated or normal branchlets, are pulvinate, hemispheric or oval,
or occasionally tend to be ligulate, but these sori always have smooth sur-
faces. Only rarely are the telia confluent on leafy shoots, but are commonly
seen in this condition on leafless parts of branches and stems. The telia
inhabiting these leafless bark surfaces are about the same in shape as
those on the brooms but tend to be larger and often coalesce, giving rise
to a narrow elliptical sorus. Ligulate telia are not uncommon on stems,
but in this instance they are never furcate, rough surfaced, or lacunose,
this combination of characters being descriptive of the telia belonging to
G. effusum and not G. nidus-avis. The dimensions of the ungelatinized
telia are 1-6.5 mm. in diameter and 0.5-5.5 mm. in height on brooms,
while on leafless, scaly-barked branches and stems they are 3—5 mm. long,
3—6 mm. wide and 2-9 mm. high. Upon gelatinization, the telia enlarge
considerably in height, but little in diameter and width or in length along
the stem. The highest gelatinized telium measured was 16 mm. In this
condition they are globoid, ellipsoid or sometimes ligulate. Dodge (1931)
reported a telium at least two inches in length which is longer than any the
writer has seen.

In addition to a study of the size and shape of the telial sori, their usual
color was determined by comparison with standard color charts in Maerz
and Paul (1930). Color is probably the most variable of the telial charac-
ters, but in the fresh condition just after emergence, it has been found to

Prince: GYMNOSPORANGIUM Nipus-Avis THAXTER 499

be reasonably constant, regardless of the host or position on the host. It
should be mentioned here that no telia appeared to be exactly chestnut
or cinnamon brown in the ungelatinized condition as reported by Kern
(1911) and Arthur (1934). By careful and repeated comparisons, the
color of ungelatinized telia on J. virginiana were found to be “Coromandel,”
on J. scopulorum, “Cordova,” and on J. horizontalis, “rustic brown.” All
three of these colors are slightly darker than chestnut. Fully gelatinized
telia were determined to vary from a light yellowish-orange to a darker
yellowish-orange or “Spa Tan.” Telia repeatedly gelatinized become cin-
namon-brownish. After one long period of gelatinization, such as during a
continued rain or after three or four brief periods of extension, the telia
drop from their host leaving oval scars which are at first yellowish-orange
from the remaining mycelium. Sooner or later, however, the scars become
covered by callous tissue formed by the host. Regardless of the host, while
the telia are still attached or immediately after they fall off, the colored
subtending tissue mentioned above is nearly always present. Telia that
emerge from the fissures of old bark often drop off or dry up after gela-
tinization without leaving-evidence of their presence.

Although the telia may be somewhat affected by changes in the sub-
stratum or by ecological conditions, the teliospores are little affected by
these factors, and therefore offer many important taxonomic characters.
Because of this, a study has been made of the spore development and
morphology, and a separate investigation. on the process of germination
which includes basidium and basidiospore formation and the discharge
of the latter (Prince, 1943). Fully formed teliospores can be found before
the host tissue, which covers the telium, has been ruptured, but if speci-
mens are collected early enough in the spring it can be seen that prior to
the formation of the teliospores there are telial primordia, the ontogeny
of which has been carefully described and illustrated by Dodge (1918a),
who pointed out that the terminal cells of the primordia elongate three to
five times their diameter and form buffer cells which are followed by telio-
spores that arise from penultimate cells. Upon the enlargement of the
teliospores and elongation of the pedicels, the host tissue covering the
telium is finally ruptured.

The mature teliospores are very irregular in shape, size, and number
of cells. Their shape varies from ellipsoid, tapering at both ends, through
oblong to oval with both ends more or less obtuse. The most common form
of teliospore is ellipsoid with both the upper and lower cells tapering
slightly, and almost without exception they are somewhat constricted at
the septum or septa which when present divides the spore into two or
more cells. The number of cells in each spore varies from one to four.
Mesospores, characterized by being one-celled, are commonly seen among
the spores of the telium while the three and four-celled teliospores are
rarely seen. The smooth teliospore walls are usually pale cinnamon-brown
but the walls of some spores may be hyaline while others are dark cinna-
mon-brown. Similarly, the thickness of the wall is variable also, ranging

500 FarLowI14, VoL. 2, 1946

from 0.4—-2.2 ». The walls of the two-celled teliospores have pores that
are nearly always papilliate. The pores are located at the septa, except
for the frequent presence of an additional apical pore in the upper cells.
There are no basal pores in the lower cells. In spores with three or four
cells, the pores are located as in the two-celled spores, while the aa are
apical i in mesospores.

At an early stage the pedicels become differentiated from the teliospores
which eventually become septate and enlarge in diameter while the stipes
for a long time retain their original diameter, but elongate. During the
final stages of elongation, the wall of the pedicel becomes thickened and by
the time the spore has matured it has become somewhat hygroscopic. The
top of the pedicels just below the spores often enlarge in diameter but never
become bulbous like those of G. clavipes. When the top of the pedicels
do enlarge upon gelatinization, this may continue to the extent that the
outer wall bursts leaving a ragged fringe attached to the base of the spore,
but with the inner wall surrounding the lumen remaining intact and con-
tinuous with the lower portion of the pedicel which does not break away
or deliquesce.

As is true with the characters of the aeciospores, those of the teliospores
do not furnish evidence that would lead one to consider G. juvenescens to
be distinct from G. nidus-avis, nor are there any valid differences that would
separate the eastern from the western strains. The only character that
might be considered valid is that on J. scopulorum the telia do not occur
on the old branches and stems away from the brooms, whereas on J. vir-
giniana the telia are produced on these parts of the hosts. This difference,
however, cannot be accepted because on J. Aorizontalis the telia occur on
both the leafy shoots of brooms and on branches not connected with brooms.
In addition to these facts, the morphology and size of the telia are the
same regardless of host, and therefore from investigations by the writer,
G. juvenescens is not a valid species.

In a letter dated July 18, 1941, Dr. Long agrees with the author that G.
juvenescens on J. scopulorum is the same as G. nidus-avis, but he writes
that he has seen G. juvenescens causing brooms with normal needles as
well as the acicular form “but in all cases the brooms were dense and not
of the open type that is found in certain areas of the east.” Dr. Long be-
lieves that the open type broom which occurs more commonly in the South
on Juniperus virginiana may be caused by a different rust than that which
causes the dense brooms that are more common on the northern cedars
(Juniperus virginiana var. crebra). His basis for this belief is that he ex-
amined 3040 “eastern red cedar” trees in and near the District of Colum-
bia. Of the trees examined, 382 were infected with G. nidus-avis. Data
recorded according to the type of symptom produced showed that on the
infected trees there were 58 trunk lesions, 1108 branch lesions, 484 brooms
of the open type, and 1 “poor little broom” with acicular leaves on a
stunted tree. During additional observations in Arkansas he found that
the cedars there also had only the open type of brooms. Additional char-

PRINCE: GYMNOSPORANGIUM Nipus-AvIS THAXTER 501

acters, he believes, that distinguish the southern from the northern rust,
are that his observations show that the bark under the telia is a golden
yellow during the development and gelatinization periods of the telia and
that there are many mesospores in southern telia. He seems to have com-
pared his extensive observations of southern specimens with a single col-
lection from Sharon, Massachusetts, and because the latter specimen
lacked certain characters present in the others he believes they are different.

These differences are invalidated, first because he did not recognize
the two kinds of eastern red cedars, J. virginiana and J. virginiana var.
crebra, the former of which, as has already been pointed out in this work,
usually forms open brooms when attacked by G. nidus-avis, while the latter
usually has dense brooms under the same conditions. In addition, the true
J. virginiana according to Fernald and Griscom (1935) may occur as far
north as eastern Massachusetts, but is most abundant from the coastal
plain of New Jersey south and westward; Thus the cedars in Washington,
D. C., and Arkansas are well within the range of the true species and not
the northern variety. Secondly, the orange coloration of the bark under
the telia is characteristic for at least two species of Gymnosporangium in
addition to G. midus-avis. Thirdly, during the present investigations, the
orange coloration under the telia has been observed on all the Juniperus
hosts of G. nidus-avis studied by the writer in the field; and finally, meso-
spores are not uncommon in the telial sori regardless of host or locality.
Therefore it seems evident that it is important not to separate G. nidus-
avis into geographical species or varieties by the broom types, coloration
of the host tissue under the telia, or by the presence of mesospores.

As a further test of the possibility that G. juvenescens could be distin-
guished from G. nidus-avis in the telial phase, it would be possible to do
it on the basis of teliospore dimensions. Accordingly, the lengths and widths
of one hundred spores were measured from each of seven geographical
localities; the collections coming from the following species of Juniperus
from the different localities:

J. horizontalis Moench. Bar Harbor, Maine

J. horizontalis Moench. Saskatoon, Saskatchewan

J. scopulorum Sarg. Fort Collins, Colorado

J. virginiana L. var. crebra Fern. Canton, Jamaica Plain, Norwood, and
& Grisc. Waltham, Massachusetts

J. virginiana L. Chapel Hill, North Carolina

Clemson, South Carolina
Montvale Springs, Tennessee

_ In addition, the measurements were based not on spores derived from -
a single telium, but from several telia on specimens collected or received
while these investigations were in progress, and only spores from dried
telia mounted in lactophenol were used. Thus it was possible to obtain

502 Fartowia, VOL. 2, 1946

from each host and locality a good representation of the degree of varia-
tion in the size of the spore, the basis of the data in Table 4.

Tas_e 4. Tue Size or TELIOSPORES OF G. nidus-avis FROM DIFFERENT TeLiaL Hosts
AND LOCALITIES

Spore Mean Spore Mean

Lengths Lengths Widths Widths

Hosts in us be inp Le
Juniperus
horizontalis (Me.) ...... 22.0-47.0 32.8 11.7-27.9 17.3
horizontalis (Sask.) ..... 25.0-63.2 37.1 11.7-27.9 16.5
scopulorum (Colo.) ....- 19.1-50.0 34.4 11.7=22.5 16.2
virginiana var. crebra
PARR ea haarata ur 26.4-51.4 35.8 13.2-23.5 17.0
virginiana (N. C.) ...... 23.5-55.8 38.3 10.2-25.0 17.4
virginiana (S.C,) ....... 22.0-60.2 41.7 10.2-22.0 16.2
virginiana (Tenn.) ...... 20.5-51.4 39.4 8.8-23.5 16.3
Total Range Total Range
in Spore Mean in Spore Mean
Lengths Length Widths Width

19.1-60.2 37,07 8.8-27.9 16.70

An analysis of the data in this table shows that the greatest difference
of 8.5 » in the mean spore lengths occurs between spores from Maine on
J. horizgontalis and spores from South Carolina on J. virginiana. In the
width measurements, the spores from South Carolina and those from Colo-
rado on J. scopulorum have the same mean, and the greatest variation
from these (1.2 ,) is in the spores from North Carolina on J. virginiana.
There is certainly no evidence in Table 4 to show that G. juvenescens is
different from G. nidus-avis when the mean lengths of the spores on J. vir-
giniana var. crebra {rom Massachusetts differs only by 1.4 » from those on
J. scopulorum from Colorado, and the mean widths differ only 0.8 ; the
first comparison in the spores from the north and south seems to bear out
Dr. Long’s argument, yet even this is invalidated when an analysis of the
data in Table 5 is made.

For the data in Table 5, the lengths and widths of fifty spores from
telia on J. virginiana var. crebra were measured from each of twelve sta-
tions in Massachusetts. The telia are different from any used for the data
in Table 4. All twelve stations are within a twenty-mile radius of Boston.
First, it will be noted that there is nearly as great a spread in the mean
lengths and widths of the spores produced on the one host within a limited
area as was found throughout the whole geographic or host range of G.
nidus-avis. Next, it can be seen that there is as large a difference (8.5 »)
in the lengths of the spores from Newton and Canton, which are only
about ten miles apart, as there is between the spores from South Carolina
and Massachusetts to which attention was called in the preceding table.
Therefore it may be concluded that G. juvenescens cannot be distinguished

PRINCE: GYMNOSPORANGIUM NipuUs-AVIS THAXTER 503

TaBLE 5, THE S1zE OF TELIOSPORES FROM Juniperus virginiana vaR. crebra IN EASTERN

_ MAssACHUSETTS
Spore Mean Spore Mean
Station Lengths Lengths Widths Widths
in uw be inp be
Beverly os oo 28.5—53.8 39.21 14.2-23.7 17.51
Medford ........ .. 23.7-55.4 41.45 14.2-23.7 18.03
Brighton ......... 23.7-45.9 36.86 11.1-31.6 20.41
Newton ......... 22.1-42.7 34.96 15.8-25.3 20.25
Hammond’s Pond
Brookline ...... 31.6-44.3 36.70. 12.7-28.5 18.98
Chestnut Hill
Brookline ...... 25.3-49.0 r . 39:39 11.1-23.7 18.82
Waltham ........ 31.6-50.6 39.36 15.8-25.3 18.67
Jamaica Plain .... 26.9-44.3 36.38 15.8-23.7 18.04
QUINCY ieee dee et 31.6-55.4 43.19 14,2-30.1 20.41
Walpole ......... 25.3-50.6 37.02 14.2-26.9 20.41
Gantonitect= er 31.6-53.8 43.51 14.2-28.5 19.78
Sharon: veamece 26.9-57.0 39.23 14.2-25.3 19,93
Total Range Total Range
in Spore Mean in Spore Mean
Lengths Length Widths ° Width
22.1-57.0 38.76 11.1-31.6 19.30

from G. nidus-avis on teliospore size and neither can the southern rust be
differentiated from the northern rust by this method. The total range in
the teliospore lengths and widths of G. nidus-avis, then, may be said to
vary from 19.1-60.2 » x 8.8-31.6 », with a mean length of 37.9 » and a
mean width of 18.0», In addition, after a comparison of average spore
measurements in Tables 4 and 5, of the Massachusetts material, it can be
seen that spore measurements from a single specimen or even one locality
are not enough to give an accurate representation of spore size. For ex-
ample, 35.8 x 17.0 » (Table 4) is the average spore size for Massachusetts
material because specimens from throughout the state have been included,
whereas 38.8 x 19.3 u, (Table 5) is representative of spores from only the
eastern portion of the state. Moreover, neither of these are as accurate for
the species as a whole as 37.1 x 16.7 » in Table 4, this being the average size
of spores from throughout the geographic range of the fungus.

THE HOSTS AND THEIR RELATIVE SUSCEPTIBILITY TO
G. NIDUS-AVIS

Among the many problems facing the phytopathologists are the subjects
of immunity and susceptibility of potential or actual phanerogamic hosts.
of the various rusts. The importance of these studies may be realized when.
it is noted that the alternation of hosts in the life cycles of some rusts:
gives the phytopathologist one of the few means of control of these fungi,
since by removing one of the hosts to a safe distance, the alternate host

504 FaRLowIA, VOL. 2, 1946

may be protected to a great extent. For this reason, then, the present
studies were undertaken in order to determine what hosts are most sus-
ceptible to attack by the fungus and which are resistant or immune. From
the information obtained during this study, it is now possible to state
definitely which pomaceous trees may be grown near the Juniperus species
and which are susceptible to G. nidus-avis and, therefore, unsuited for orna-
mental use. Thus the matter of planting horticultural projects may be
undertaken with a degree of foresight not previously possible and with a
consequent reduction of waste effort.

As far as can be determined from the literature, only a limited number
of investigations have been carried out in order to determine the host
plants of G. nidus-avis. The earliest records that we have are those of
Farlow and Thaxter. Following these earliest records are those of Arthur,
Kern and others who have increased the list of susceptible hosts. A sum-
mary of these investigations is given in Table 6.

In the above table the results of previous experiments on both G. nidus-
avis and G. juvenescens have been tabulated in parallel columns to facili-
tate comparison. It can be seen that according to previous investigations it
was thought that G. nidus-avis could be distinguished from G. juvenescens
on only four hosts: Crataegus Pringlei and Malus coronaria as hosts of GC.
nidus-avis but not of G. juvenescens, and Amelanchier intermedia and
Sorbus americana as hosts of G. juvenescens but not of G. nidus-avis.
These and other purported differences have been considered sufficient to
keep these two species distinct. However, it should be pointed out that
when the results of different investigators are compared, there is an incon-
sistency which makes one question the advisability of such a distinction.
It only needs to be pointed out that Prince and Steinmetz (1940) were
able to inoculate successfully Cydonia oblonga with spores of G. juvenes-
cens whereas Arthur (1912) was not. Similarly with Malus pumila var.
Wealthy, Clinton and McCormick (1924) were able to obtain infection
while Prince and Steinmetz were not, yet the writer in subsequent experi-
ments was successful with his inoculations on this host.

In addition to the above brief discussion of previous investigations, it
seems desirable to give a complete list of the pomaceous hosts of G. nidus-
avis which includes those reported also for G. juvenescens.

Farlow and Thaxter, the two original investigators, were able to culture
the rust successfully on only Amelanchier canadensis (L.) Medic., and an
undetermined species of the same genus. Arthur, who reported cultures
extending over the period from 1907 to 1915, successfully inoculated
Amelanchier erecta Blanchard, A. intermedia Spach. A. ovalis Med., Cy-
donia oblonga Mill., Crataegus Pringlei Sarg., Malus coronaria (L.) Mill.,
M. toensis (Wood) Brit., M. pumila Mill. var. Whitney Crab, and Sorbus
americana Marsh. Kern (1911) reported additional hosts without cul-
tures. They are Amelanchier alnifolia Nutt., A. florida Tindl., A. oreophila
Nels., A. polycarpa Greene, and A. pumila Nutt. G. nidus-avis was reported
by the same author as cultured on Amelanchier vulgaris Moench. Davis

PRINCE: GYMNOSPORANGIUM Nipus-AVIS THAXTER 505

TABLE 6
_ HOSTS INVESTIGATORS DATE G.NIDUS-AVIS G.JUVENESCENS DATE INVESTIGATORS
Anelanchier
einifoliad.. cise cnuseuecsene Mistetale ercielaterelenett eters coveccccecs © cesee L924 «.e+ Fraser
canadensis ...... ». Thaxter ......,1887 ..... eo ecics cee ae + ..... 1907 .... Arthur
" soeveces TMARCOD 0.0504 1889 12.6 Ho cbeeroeees * oneee 1908 ...., Arthur
es apres Pit ATCDUEL ers feel O10 4 ve oe eile te AS Det ct ee oer er eee i
‘2 eeeeevean Dodge eeeese eee 1918 @eeoe + eeeeneeeerrsewernaer ee ee eee eaee eeeoseveon
a eoeeee Prince &

Steinmetz .. 1940 .10e FH cececccrvccevseseeesnrevvesssssessnee
On Ch yan poorodnt 2 a raaanooo Dee aoc ae Godcpsbedn + ..+. 1909 .... Arthur
bees poe meee se huy tee estes 1 OLD: cisieie! | = ike cletaievaielsials + ..... 1912 .... Arthur
intermedia ........ Arthur ....... 190Gb eccc r notes see . * 4... 1908 .... Arthur

laevis ........ -seee Prince & :

Steinmets .. 1940 1.60 PF cccccrccsnscrsccccsssvsnnensesscscsoce
oblongifolia, ici. ccecatesescccs Aialavarsteseiaveroisiete ane iorielereee crs - > «oeeee.1918..... Davis
OWSLTS Ge cee AMthUr sc cagieen Lolessies Slee selec eel eaealec E's etale eae sa an tele tetereyeierctoss

a me are pee ae Arthur ....... 1) Ave Gos GAC peislavetele einarere mictaretereerelelersielete lenis :
stolonifera ....... Prince & :

Steinmetz ., 1940 .... % wscccccsceres Ce One he oe dneereceseree
canadensis
x A. stolonifera .. Prince &

Steinmetz .. 1940 1... % ssacscveres aiatiolelaliniats a's laie 6 pio eset ele
Wiegandli .....c006 Prince &

' Steinmets ... 1940 .:25 0 4. (oe. ccctteccdecucauente Rea tinccligie heise
_ Aronia
arbutifolia ....... Thaxter ...... 1887 1... 7 ssccsccnvcvesccavees Ba eee lecaveieiels etre ois :
MELANOCAFPA weeccaeerrescerer label cicieioletejsle ese (s/s) eivieletsteyereiele sooes 7 eoeee 1907 .... Arthur
Chaenomeles
japonica .......... Prince & ‘
Steinmetz .. 1940 1... - ssseveces « | esesee 1907 .... Arthur
Cydonia
oblonga ..... Beescc Arthur ....... LOLS eo PU Weeics cclte ) tesa el Obes s6 arthur
4 Bites tae -». Clinton &
MeCornlclkth, U92dte stat sc shies « ccisisie's ee a ele sistas sae atlep dae oes 67s 4
u Sie eivslgte sistas Prince &
Steitimet so 5 LOGO creche sale etnies aisles e'nle o/c s.¢ 6 lecwlsisislse alse cite c wie
Crataegus
coccinea ......ceee Thaxter ...... TODO a ere ee epics «cise tee es.cets Be oodE AAC ORB OUDOOUC
Pringlel ...c.csceve Arthur ..... 66 RIO ae Pe wee ccecvee - sees 1907 .... Arthur
Malus
coronaria ..,.++¢+. Arthur ....... VOU 2 srl ce Min ee cies sels & — seoee 1912 .... Arthur
foensi8 ......ece0s Arthur ...... we L GLO crete atte, ainle atolais cleo e wis is oe slejefuje't.cie' <laiw cess tiv 4-6 .
. eoeceeeeces Clinton & :

Matebelok) :. Sgr tre ein Sc cc ne aeeedccm eras tasdees cesmss :
punila ee rene eeves » Thaxter ....0 LBB7 ice * sevecccercensesssvesceessseccsncs eens
pumila Clinton &
var. Wealthy ...... McCormick .. 1924 .... 4 ssssevas Blevareteta elele elereisiearewieleiniers(a sla’ele te
pumila
var. Wealthy ...... Prince &

Steinmetz .. 1940 1... —- saceccene GOGO RODOCI OOOO ORO ADECMOOUOE
pumila
var. Whitney Grad . Arthur ....... 1908 1.26. ® ceescassves Parsee cascssereneeee esses

Sorbus
americana ......... Farlow ....... 1886 1... = cesceccece # covee 1907 .o.. Arthur
w wpeevecree Thaxter eueeee 1887 eeee oni eeecven eave + esene 1909 eave Arthur
“ Sis clcece ee URAXCOE:) oc ciatew LOGS” veislel “ue aicicincsicccccs cence eda cscs Bre eaerterycia
Reta ce ne ict ‘Prince &

Steinmetz we NORD 6 ose oi Soca ccce ew cercccanstenvsassesesecons

(1918 unpublished) cultured the rust on A. oblongifolia Roem. Dodge
(1918b) successfully inoculated undetermined species of Amelanchier and
Malus. Clinton and McCormick (1924) cultured the rust on leaves of
Malus pumila Mill. var. Wealthy in Petri dishes. Fraser (1925) success-
fully inoculated Amelanchier alnifolia Nutt. Arthur (1934) added to the
_host list without reference to cultures Amelanchier Bakeri Greene and
A. humilis Wieg. Crowell (1940) reported Malus fusca (Raf.) Schneid.

506 FarLtowlA, VoL. 2, 1946

while Prince and Steinmetz (1940) supplemented the host list of G. nidus-
avis by means of cultures on A. laevis Wieg., A. stolonifera Wieg., and
A. Wiegandii Nielsen.

In a consideration of the above data it is possible to state that G. juvenes-
cens as a distinct species does not have a firm foundation, thus substan-
tiating the conclusion arrived at as a result of the morphologic and biometric
studies. However, the conflicting data already mentioned indicated the
desirability of carrying on the investigations under conditions that were
as near alike as possible. The facilities and extensive collection of living
pomaceous hosts in the Arnold Arboretum of Harvard University offered
an excellent opportunity to round out adequately our knowledge of the
variation in susceptibility of the pomaceous plants.

As a source of inoculum, material was obtained from the following hosts
and localities: — Juniperus virginiana var. crebra from Canton, Waltham,
and Barnstable, Massachusetts; J. virginiana from Chapel Hill, North
Carolina, Clemson, South Carolina, and Knoxville, Tennessee; J. hori-
zontalis infected with the rust was obtained from Bar Harbor, Maine, and
from Saskatoon, Saskatchewan. Brooms caused by the fungus on Juniperus
scopulorum were obtained from Fort Collins, Colorado.

Immediately after collection or on arrival by post, the telial material
was placed in a refrigerator which was maintained at a temperature of
about 6° C. Under these conditions teliospores of New England material,
freshly collected, would retain their viability up to six weeks. The telio-
spores of the southern material received by post, however, retained their
maximum viability for only four weeks. After five weeks less than 50%
of them would germinate. This fact made it necessary in the latter instance
to use cuttings and potted plants forced in the greenhouse for inoculation
experiments, because the spring season at the stations in the South from
which the telia were sent was more than four weeks in advance of the
spring season in Massachusetts. Cuttings were tried but they were not
found to be satisfactory even though infection was obtainable on susceptible
plants. For this reason potted plants of Amelanchier stolonifera and potted
grafts of Malus pumila var. Wealthy were prepared in numbers for use
during these investigations. These two hosts were chosen because in previ-
ous experiments the former was found to have been susceptible in all
trials, the latter, because there was a question as to its actually being a
host of G. nidus-avis.

The inoculum was prepared some time before it was to be used. Tests
were made by observing the germination of spores in Van Tieghem cells
or by observing the amount of germination of the teliospores just before
the inoculum was used. Usually the latter method was followed because
the percentage of viability of the teliospores when left en masse was more
accurate than when the spores were separated and placed in Van Tieghem
cells.

The inoculum was made by thoroughly wetting with distilled or rain
water, the telial material to be used, and then placing the telia in a moist

PRINCE: GYMNOSPORANGIUM Nipus-AviIs THAXTER 507

container for at least two hours. Often material to be used for a fore-
noon’s inoculations was prepared late the night before and left under
cool conditions, that is, at 18 to 20° C. and by morning basidiospores
would begin to appear. When cultures were to be made in the afternoon,
telial material was placed in moist chambers early the same day, thus
keeping a careful check on all inoculum used. When ready for use, a
thick suspension of germinating teliospores was made in small, wide-
mouthed, stoppered bottles from which inoculum was removed and painted
on both sides of the leaves or on the surface of the fruits when the latter
were present. All of the inoculated branches were protected during the
infection period with celluloid cylinders similar to those described by
Hubert (1916) but Sphagnum was used to plug the ends instead of cotton,
and the chambers were then covered with paper bags after which the in-
oculated branches were marked with numbered tags. The paper bags
served a twofold purpose. They protected the inoculated plant parts from
the direct rays of the sun and prevented rapid evaporation from the
Sphagnum plugs. Normally the chambers were allowed to remain in place
from thirty-six to seventy-two hours, but during the unseasonably cold
weather in the spring of 1940, they were left on for five days without
causing injury to covered plant parts. After the chambers were removed,
the suscepts were examined at frequent intervals until ready for collection,
or until the subject was considered to be immune. In most cases, the
plants listed as immune were observed for four weeks before a decision

was made.
THE DETERMINATION OF VARIATION IN Host SUSCEPTIBILITY

Considerable work has been done from the time of De Bary up to the
present to determine the variation in susceptibility of economic plants to
rust diseases, but nothing has been recorded as to the variation in sus-
ceptibility of pomaceous hosts of Gymnosporangium nidus-avis.

To express my observations with respect to this phenomenon, standards
were established according to which data were recorded for these suscep-
tibility studies. Five arbitrary categories were adopted; they are similar
to those employed by other uredinologists but differ, through necessity, in
some details. Their designations and descriptions follow: (1) ZJmmune.
No evidence of infection observed. (2) Very resistant. Necrotic spots pro-
duced, usually with some abortive spermogonia, always without aecia;
the presence of an invasion zone clearly evident in a transverse section
the leaf (Plate I, Fig. 4). (3) Resistant. Lesions sometimes becoming
necrotic; spermogonia and aecia taking longer to develop than on the more
susceptible hosts. (4) Susceptible. Spermogonia forming within twelve
days, aecia within four to five weeks; a few lesions occasionally becoming
abortive or even necrotic; lesions with an average of one to five aecia.
(5) Very susceptible. Time for the development of spermogonia and aecia
about the same as in four; no lesions becoming necrotic; lesions with an
average of six or more aecia.

508 FARLOWIA, VOL. 2, 1946

As a result of the writer’s investigation with artificial inoculations made
in the Arnold Arboretum, it is now possible to add to the already rather
extensive host list. These new hosts are Amelanchier amabilis Wieg., A.
asiatica (Sieb. and Zucc.) Endl. var. sinica Schneid., A. Bartramiana
(Tausch) Roem. x A. laevis Wieg., A. florida Lindl., A. grandiflora Rehder
forma rubescens Rehder, A. oblongifolia Roem. var. micropetala Robins.,
A, sanguinea (Pursh) DC., A. sera Ashe, A. spicata (Lam.) K. Koch,
Amelosorbus Jackii Rehder, Chaenomeles lagenaria (Thunb.) Lindl., C.
superba (Frahm) Rehder, Sorbaronia Dippelii (Zab.) Schneid., Sorbus
americana Marsh., S. americana var. fructo-albo auct., S. americana Marsh.
var. micrantha Dum.-Cours., S. Aria (L.) Crantz var. salicifolia auct., S.
aucuparia L., S. aucuparia L. var. edulis Dieck, S. aucuparia L. var. xantho-
carpa Hartw. and Ruempl., S. discolor (Maxim.) Hedl., S. japonica
(Decne.) Hedl. var. callocarpa Rehder, S. latifolia (Lam.) Pers., S. scopu-
lina Green, and S. thuringiaca (Ilse) Fritsch.

It became obvious during the progress of this investigation that there
were considerable variations in the degree of susceptibility of some of
the hosts, not only in the different host species but also in different plants
belonging to the same species. For this reason, all of the plants inoculated
and found to be susceptible in some degree are listed in Appendix I and
the variations in the susceptibilities of the hosts are indicated by symbols.
The origin of the telia used for inoculum is shown with reference to host
and geographical station.

Some plants, as Amelanchier canadensis, A. humilis, A, sanguinea, A.
sera, A. stolonifera, Chaenomeles lagenaria, and Sorbus scopulina were
found to exhibit the same degree of susceptibility to G. nidus-avis regard-
less of range of telial host. Other plants, however, show a variation in
their reaction to the fungus. The cultures on Amelanchier amabilis tend
to separate the eastern rust from the western. This host was found to be
susceptible to the rust from Colorado and Saskatchewan, but was resistant
to the rust from Maine, and immune to the rust from Massachusetts. The
fact that there was a difference in the degree of susceptibility of the aecial
host when it was inoculated with spores from widely separated localities,
strongly suggests that different strains of the fungus have arisen through
geographic segregation. Additional evidence of this is shown by inocula-
tions on Amelanchier grandiflora forma rubescens, A. oblongfolia, A. spi-
cata, Amelosorbus Jackii, Sorbus americana and other species listed in
Appendix I.

The segregation of geographic strains is only one factor. A complication
results from the fact that there is also evident a tendency for basidiospores
from a given telial host to infect one pomaceous host to a different degree
than do the basidiospores from another telial host. Here then is evidence
of fungus strains. Thus, according to the results in Appendix I the rust
strain on Juniperus virginiana can be distinguished on Amelanchier asiatica
var. sinica, A. Bartramiana x A. laevis, and Sorbus japonica var. callo-
carpa. Similarly, the strain from Juniperus horizontalis can be separated on

PRINCE: GYMNOSPORANGIUM Nipus-AVIsS THAXTER 509

Sorbus americana var. micrantha and Amelosorbus Jacki. Finally, the
strain on Juniperus scopulorum can be separated on Amelanchier florida,
Amelosorbus Jacki, and Cydonia oblonga.

While there is evidence that there is some host specialization, it does
not seem that this should be taken to indicate the desirability of erecting
large numbers of strains as has been done for the graminicolous species of
rusts. In addition another examination of Table 6 shows clearly that ex-
periments with a limited number of hosts may lead to false conclusions.
Examples of this are shown by the inoculations on three-plants of Ame-
lanchier spicata numbers 5215—A, 12635—A, and 12635-C. In this instance .
A. spicata 5215—A is immune to the spores from Juniperus horizontalis
from Maine, but is susceptible to infection by spores from the same host
from Saskatchewan. On the other hand, Amelanchier spicata 12635—A and
12635-C are offspring from the same parent. This shows a variation in
the susceptibility within an individual host species. The same phenomenon
is shown in the hybrid Chaenomeles superba 13017 and 18184, and in the
species Sorbus aucuparia 17264—A, 17682—D, 17682-C, and 17266-E. The
concept is thus made more confusing by the fact that individual host
species show variations in the degree of susceptibility.

In spite of the confusion that seems to be added by the factor of indi-
vidual host variation in susceptibility, it is possible to coordinate these
variables if we consider the average of all of the inoculated susceptible
hosts to inocula from the four geographic regions. In addition, there is
confirmation of the supposition that there are different strains of G. nidus-
avis. Data showing the average degree of susceptibility of all the sus-
ceptible hosts that were inoculated are given in Table 7.

TABLE 7
TABLE 7. DATA SHOWING THE AVERAGE DEGREE OF SUSCEPTIBILITY OF ALL OF THE

INOCULATED SUSCEPTIBLE:-Hosts To Four DIFFERENT INOCULA

J. vir- J. scopu- J. hori- J. hori-
Telial Hosts giniana lorum zontalis zontalis
Origin of
Inoculum Mass. Colo. Maine Sask.
Degree of .
Susceptibility
of Aecial Hosts Percent Percent Percent Percent
Very Susceptible ......... 0 Z 0 3
Susceptible: .0 242. ..0.0... 40 44 32 46
Resistant os cee eer 19 1G 52, 26 21
Very Resistant .......... 15 15 7 9
Jamies tere ee ee eee 26 29 35 21

The above values mean that 2% of the inoculated plants were very
susceptible to the rust from Juniperus scopulorum, 44% were susceptible,
10% were resistant, 15% were very resistant, and 29% were immune.

510 FartowlA, VoL. 2, 1946

These data show that degrees of susceptibility or resistance are of the
same order for all of the different inocula, but a slightly greater percentage
of the hosts were immune to the inoculum from Maine than to the inocula
from the other three stations. This confirms my interpretation of the data
presented in the preceding table, that there is evidence of host and geogra-
phical strains of G. nidus-avis. There is also a slight variation in the
degrees of susceptibility of the hosts to inocula from Massachusetts and
Colorado, and Colorado and Saskatchewan. In addition, although the
values given in Table 7 are expressed in the degrees of susceptibilities of
the plants to the fungus, these values may also be interpreted as the ability
of the fungus to attack these various hosts, since the same plants were
used throughout.

The differences shown, even though slight, indicate that there is some

ca
A ORIGIN OF INOCULUM FROM
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i CULTURED

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AECIOSPORE LENGTHS

Grapu A. A Comparison of Aeciospore Lengths. Aeciospores Cultured on Similar Hosts
with Inoculum from Four Localities

variation in the virulence of these inocula when they are placed on the
various hosts. There is still additional evidence of the presence of biologi-
cal strains within G. nidus-avis to be found in the time necessary for
development of the spermogonia and aecia. The data concerning this are
presented in Table 8. Again the amount of variation in the host reaction
to the fungus from different areas and different hosts is slight, but the
variation, where it exists, indicates the presence of biological strains.
The sizes of the aeciospores furnish evidence of the presence of biologi-
cal strains. The aeciospores measured came from cultures made with
inocula from Maine, Massachusetts, Saskatchewan, and Colorado. The
lengths and widths of these aeciospores are compared in Graphs A and B.
The aeciospore measurements represented here are taken from the data
given in Table 7. In Graph A where the spore lengths are compared, the

Prince: GYMNOSPORANGIUM Nipus-Avis THAXTER 511

Massachusetts strain varies slightly from the Saskatchewan. Those from
Maine and Colorado are very similar, but vary slightly from the two pre-
ceding strains. There is a greater degree of variability shown in the spore
widths than in the lengths. Graph B illustrates this, and shows that the
variability is not large enough to change the conclusions already drawn.

Even though the degree of variability shown by these comparisons is
small, when one considers this variability and that which exists in the
differences in aecial host susceptibility, and in the time of development,
and finally, the variability in the manifestation of the symptoms of disease
on the telial hosts, the only interpretation that can be advanced is that

150

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! WHICH AECIOSPORES WERE
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AECIOSPORE WIDTHS

Grapx B. A Comparison of Aeciospore Widths. Aeciospores Cultured on Similar Hosts
with Inoculum from Four Localities

there are definitely biological strains of G. nidus-avis. However, there is
no evidence to show that more than one species is represented here, or
that the one species can be separated into varieties or even forms. Cer-
tainly G. juvenescens has been shown to be morphologically identical
with G. nidus-avis. More conclusive evidence which bears out this state-
ment is that aeciospores from Amelanchier grandiflora forma rubescens,
which were produced following an inoculation with basidiospores from
telia on Juniperus scopulorum, were used to inoculate successfully Juni-
perus virginiana var. crebra, thus removing any doubt as to the ability
of the fungus to infect both coniferous hosts, and thereby establishing
the identity of G. juvenescens with G. nidus-avis.

In addition to the plants found to be susceptible to attack by the fungus,
a number of the plants inoculated were found to be entirely immune. From

512 Fartowia, VoL. 2, 1946

TasLe 8. A COMPARISON OF THE DEVELOPMENTAL PERIODS OF G. nidus-avis
ACCORDING TO THE ORIGIN oF INOCULUM AND THE TELIAL Hosts

Stage - First Average
Origin of Inoculum of Appearance in
and Telial Host Rust in Days Days
Maine 0 +16. 12.7
J. horizontalis ie 28-45 33.0
Saskatchewan — ~ tg aig) © 11.8
J. horizontalis 1 25-36 30.5
Colorado Tg 10-16 13.3
J.scopulorum .. 25-41 a
Massachusetts 0) 9-16 12.7
J. virginiana 7 : _
var. crebra ee 28-45 33.0
N. Carolina 0 6-14 | 8.9
J. virginiana i fa 32 ee:
S. Carolina 0 14 14 ;
J. virginiana 1 31-43 33.3
Tennessee 0 10-14 13
J. virginiana zi : 31 31

the economic standpoint it is as necessary to know the immune plants as
the susceptible. It is certain that any of the plants listed as immune could
be planted adjacent to the susceptible species of Juniperus without danger
of being infected by G. nidus-avis. However, persons interested in the
planting of these combinations of hosts should keep in mind the works of
Crowell (1934a and 1934b) and MacLachlan (1935a and b) who have
dealt with other species of Gymnosporangium which infect both pomaceous
and Juniperus hosts, since species found in the present studies to be im-
mune to attack by Gymnosporangium nidus-avis may possibly be subject
to infection by G. Juniperi-virginianae, G. globosum, or G. clavipes.

ECONOMIC IMPORTANCE AND POSSIBILITIES OF CONTROL

When compared with the three species of Gymnosporangium that com-
monly are called the cedar-apple rusts, G. nidus-avis is actually of little
economic importance since only under most favorable conditions does it
infect the apple trees. Yet even though it is of little importance in orchard
practice, it is a fungus that must be taken into consideration by the land-
scape architect who employs both its aecial and telial hosts for ornamen-

Prince: GYMNOSPORANGIUM Nipus-AvIS THAXTER 513

tal plantings, often so close together that heavy cross-inoculations result
that may cause severe damage to both hosts.

In the species of Gymnosporangium that cause the cedar-apples that
are so conspicuous in the spring, it is the aecial hosts that are heavily
damaged, whereas in the case of G. nidus-avis it is the telial host that
suffers most. Dodge (1931, 1934) pointed out that cedars (Juniperus
spp.) are actually killed following attacks by this rust, but Hartley (1913)
indicated that when cedars succumbed following attacks by this rust, it
was not the primary cause. Observations by the author support the
views of Hartley rather than those of Dodge. It is clear, however, that
the rust does cause considerable malformation of seriously infected trees,
weakening them, and then by making openings in the protective corky
bark, permits the entrance of wood destroying organisms. The most
severely infected cedar tree that the author has had occasion to study
occurs in the vicinity of Clemson, South Carolina. This tree is about eighty
years old, as determined by increment borings, and has more than one
hundred brooms. The trunk and branches are infected from the ground
line upward so that in the spring when gelatinization of the sori takes
place, the tree is nearly covered with the telia. In spite of this heavy in-
fection, however, the tree shows no external symptoms of an early demise
but when the increment borings were made, the first ten to fifteen years
of annual growth at the center showed evidence of decay. Similarly, in
several different localities where branches with brooms have been found
dead or dying, the rust did not appear to be the primary cause, although
there is the possibility that the fungus mycelium had penetrated beyond the
obvious lesions and this in some instances may have resulted in partial
girdling that so interfered with the vegetative activities of the broom as
to weaken it during conditions that might have been adverse for the tree
as a whole. Whatever may have been the cause of the death of the
brooms, the result of infection by this fungus is to spoil the appearance
by making distorted or malformed trees out of what would otherwise be
symmetrical ornamentals.

The only course open to the tree surgeon or landscape gardener after
a cedar has become infected, is to remove the broom as early as possible
after it has become evident and possibly to remove infected parts near
the point of attachment of the broom. Frequently this treatment may be
inadvisable when a whole limb must be removed since the loss of a limb
may damage the appearance of the’ tree more than the presence of an
infected branch. Thus while tree surgery may in special cases prove to
be desirable, the best method for preserving the health and beauty of
ornamental cedar trees is to prevent their infection by not planting sus-
ceptible alternate hosts within three hundred feet, preferably more, of
the host, and to practice control measures such as those suggested for
Gymnosporangium clavipes by MacLachlan and Crowell (1937). The
best method of all, where Juniperus and the pomaceous hosts must be
planted near each other for desired effects, is to plant only pomaceous

514 FartowiA, Vov. 2, 1946

hosts that have been shown to be immune, a list of which will be found in
- Appendix II. Among the names will be found a variety of shrubs and trees
which should permit plantings with Juniperus species in almost any habitat
and allow considerable latitude in the choice of types of trees.

SUMMARY

The first collection of Gymnosporangium nidus-avis was by Ravenel
on Juniperus virginiana in South Carolina during 1848, while Farlow
(1880) was the pioneer contributor to our knowledge concerning it, but
not until 1891 did Thaxter establish this rust as a new American species.
Arthur and Kern (1908 to 1915), Dodge (1918a—c) and Prince and Stein-
metz (1940) were the main contributors to life history studies of this
fungus; the latter two authors were unable to distinguish between the
western G. juvenescens and the eastern G. nidus-avis,

A summary of investigations and the results obtained follows:

1. The taxonomy of the fungus has been clarified. As a result of mor-
phological studies and inoculation experiments, it has been conclusively
demonstrated that G. juvenescens is a synonym of G. nidus-avis,

2. The geographic distribution of its telial (Juniperus) hosts and aecial
(pomaceous) hosts has been determined.

3. The symptomatology of the spermogonial, aecial and telial phases
has been defined.

4. Results of studies on the microscopical development and morphology
of the vegetative and reproductive phases in the spermogonial, aecial and
telial phases are presented in detail.

5. The pomaceous host range and varietal host susceptibility has been
clarified as a result of cultures with inoculum from four telial hosts repre-
senting seven localities, five eastern and two western,

6. The economic importance is considered, and possibilities of control of
the fungus are presented.

ACKNOWLEDGMENTS

It is with sincere pleasure that the author acknowledges his indebtedness
to Dr. David H. Linder for his thoughtful guidance and suggestions during
the progress of these investigations. The author wishes to express similar
appreciation to Professor Emeritus J. H. Faull under whom these studies
were initiated and whose special interest made them possible. Acknowledg-
ments are due staff members of the Farlow Herbarium and Reference Li-
brary and the Arnold Arboretum for their willing assistance in several
phases of the work. Thanks are due the following individuals who kindly
supplied fresh telial material for cultures from the stations indicated: A. E.
Brower, Bar Harbor, Maine; the late Dr. W. P. Fraser, Saskatoon, Sas-
katchewan; J. L. Forsberg, Ft. Collins, Colorado; L. R. Hesler, Knoxville,
Tennessee; L. S. Olive, Chapel Hill, North Carolina; R. C. Russell, Sas-

PRINCE: GYMNOSPORANGIUM Nipus-AVIS THAXTER 515

katoon, Saskatchewan; L. Shanor, Clemson, South Carolina; and F. H.
Steinmetz, Cape Elizabeth and Bar Harbor, Maine. Thanks are due G. G. «
Hedgcock, Gainesville, Florida, and W. W. Ray, Stillwater, Oklahoma for
sending specimens for comparison and especially to G. B. Cummins for the
loan of specimens from the Arthur Herbarium at Purdue University, La-
fayette, Indiana.

CLEMSON AGRICULTURAL COLLEGE

CLEMSON, SOUTH CAROLINA

AppenpIx I. Tue Hosts or G. nidus-avis AND THEIR VARIATION IN SUSCEPTIBILITY AS
DETERMINED BY ARTIFICIAL INOCULATIONS

Mass. Maine Colo. Sask. N.C. SEG:

wh wt ue Whe J. Sis
Hosts virginiana horizon- scopu- horizon- virgin- virgin-
v.crebra _ talis lorum talis tana tana
Amelanchier
alnifolia 15700 Si 5
amabilis 7058 I R S 5
astatica
var. sinica 15694-C R S 5 S
Bartramiana
x A. laevis 4405 Soa 5 S
canadensis 19100-B S S) S
florida 11315-A I I VR I
grandiflora
forma rubescens 20678-A 5 5 S VR
humilis 6817 5 S S i)
intermedia 21920-B 5 S
laevis 21447-B VS
laevis P—193-38 Spe I I I
oblongifolia 18265-A
oblongifolia 18265-B R I I R
oblongifolia
var. micropetala 13192 R
sanguinea 6620-B 5 S S) S
sera 11579 Ss) 5 S 5
spicata 5215-A 5 J 5 S
spicata 12635-A I I I 5
spicata 12635-C S iS)
stolonifera 644-30 5 > S) R
stolonifera (potted) S S S S S S
Amelosorbus :
Jackit 17943-B R 5 VS I
Chaenomeles
lagenaria 50098 S S S s
x superba 13017 I I Hel: I
x superba 18184 I I VR R

*S = Susceptible; VS = Very Susceptible; R = Resistant; VR = Very Resistant;
I = Immune
Numerals refer to Arnold Arboretum specimen numbers.

516 ’ FarLowiA, VoL. 2, 1946

APPENDIX I (continued)

Mass. Maine Colo. ~ Sask, “Sy. c §.¢,

oe a J. J. a: J,
Hosts virginiana horizon- scopu- horizon- virgin- virgin-
v.crebra —_ talis lorum talis tana iana
Cydonia
oblonga 15648 I VR s I S
oblonga at U. of Me. R S
Malus
pumila
var, Wealthy (potted) VR VR
Sorbaronta
Dippelit 170-25 I I I R
Sorbus
americana at U. of Me. I S
americana 22831 VR VR
americana
var. fructo-albo 20931 VR
americana
var. micrantha 22381 I R I R
Aria
var. salicifolia 222-27 VR
aucuparia 17264-A R I I R
aucuparia 17682—-D R I I
aucuparia 17266-E R S S 5
aucuparta 17682-C VR VR VR VR
aucuparia
var, edulis 21427-A VR R R R
aucu paria
var. xanthocar pa 224-27 R I VR
discolor 10o8-29-B S R 5 5
discolor 20010 R R I 5 R
R

discolor 21426-A
japonica
var. callocar pa 7703 5 I
latifolia 19861-A Vv V
scopulina 18452 R R R
R I
I R

_—

x thuringiaca 19847
x thuringiaca 19847-B

Ann

Number species and
varieties inoculated 42 34 41 37 5 2

*S = Susceptible; VS = Very Susceptible; R = Resistant; VR = Very Resistant;
I = Immune
Numerals refer to Arnold Arboretum specimen numbers.

PRINCE: GYMNOSPORANGIUM Nipus-AvIs THAXTER

a7

AppEnpIx II. A List of PLANTS INOCULATED AND FouND TO BE IMMUNE TO

Gymnosporangium nidus-avis

Aronia
arbutifolia (L.) Elliott
arbutifolia (L.) Elliott forma macro-
phylla (Hook.) Rehder
melanocarpa (Michx.) Elliott
prunifolia (Marsh.) Rehder

Chaenomeles

japonica (Thunb.) Lindl.
Cotoneaster

tomentosa (Ait.) Lindl.

Crataegomes pilus
grandiflora (Sm.) Bean

Crataegus
austromontana Beadle
Brittonii Eggl.
chrysocarpa Ashe
coccinioides Ashe
crus-galli L.
Douglasii Lindl.
fecunda Sarg.
Fischeri Schneid.
Holmesiana Ashe
intricata Lange
macrosperma Ashe
micracantha Sarg.
mollis Scheele
Oxyacantha L.
phaenopyrum (L.f.) Medic.
Pringlei Sarg.
pruinosa (Wendl.) K. Koch
punctata Jacq.
rivularis Nutt.
scabrida Sarg.
spathulata Michx.
viridis L.

Malus
angustifolia (Ait.) Michx.
arnoldiana (Rehder) Sarg.
astracanica Dum.-Cours.
baccata (L.) Borkh.
baccata (L.) Borkh. var. Dartmouth

Crab
bracteata Rehder
coronaria (L.) Mill.
coronaria (L.) Mill. var. elongata Reh-
der

fusca (Raf.) Schneid.
glabrata Rehder

zoensis (Wood) Brit.

ioensis (Wood) Brit.
(Schneid.) Rehder

lancifolia Rehder
x “Mathews Crab”

platycarpa Rehder

prunifolia (Wild.) Berkh. var. fructu-
coccinea auct.

pumila Mill.

Sargenti Rehder

Sieboldii (Reg.) Rehder

Soulardi (Bailey) Brit.

spectabilis (Ait.) Borkh.

sylvestris Mill.

Zumi (Mats.) Rehder var. calocarpa
(Rehder) Rehder

Mespilus
germanica L.
Photinia

villosa (Thunb.) DC. var. sinica Reh-

der & Wilson
Pyrus

amygdaliformis
(Guss.) Bean

communis L.

nivalis Jacq.

Pashia D. Don

phaeocarpa Rehder

serrulata Rehder

syriaca Boiss.

ussuriensis Maxim. var. hondoensis
(Kikuchi & Nakai) Rehder

Sorbopyrus

auricularis (Knoop) Schneid. var. bul-
biformis (Tatar) Schneid.

Sorbus

alnifolia (Sieb. & Zucc.) K. Koch

americana Marsh. var. nana auct.

Aria (L.) Crantz

commixta Hedl.

decora (Sarg.) Schneid.

hybrida L.

intermedia (Ehrh.) Pers.

latifolia (Lam.) Pers. ~

Matsumurana (Mak.) Koehne

pohuashanensis (Hance) Hedl.

rufo-ferruginea (Schneid.) Schneid.

forma flena

Vill. var. cuneifolia

518 FarLowIA, Vou. 2, 1946

LITERATURE CITED

Allen, Ruth F. 1930. A cytological study of heterothallism in Puccinia graminis.
Jour. Agr. Res. 40: 585-614.

. 1932. A cytological study of heterothallism in Puccinia coronata. Jour.
Agr. Res. 45: 513-541.

Arthur, J. C. 1901 a. Generic nomenclature of cedar apples. Proc. Ind. Acad. Sci.
1900: 136.

- 1901 b. New species of Uredineae —I. Bull. Torrey Bot. Club 28: 665.
. 1907. Cultures of Uredineae in 1906. Jour. Myc. 13: 203.

. 1908. Cultures of Uredineae in 1907. Jour. Myc. 14: 18.

. 1909. Cultures of Uredineae in 1908. Mycologia 1: 239-240.

. 1910. Cultures of Uredineae in 1909. Mycologia 2: 213-240.

. 1912 a. Cultures of Uredineae in 1910. Mycologia 4: 26.

. 1912 b. Cultures of Uredineae in 1911, Mycologia 4: 61-63.

———. 1915. Cultures of Uredineae in 1912, 1913, and 1914. Mycologia 7: 61-89.

————.. 1934. Manual of the Rusts in United States and Canada. Lafayette, In-
diana.

Bliss, D. E. 1931. Physiologic specialization in Gymnosporangium. Phytopath. 21:
111.

. 1933. The pathogenicity and seasonal development of Gymnosporangium
in Iowa. Iowa Agr. Exp. Sta. Bull. 106: 340-392.

Clinton, G. P. & Florence A. McCormick. 1924. Rust infection of leaves in
Petri dishes. Conn. Agr. Exp. Sta. Bull. 260: 492.

Coker, W. C. 1920. Notes on the lower Basidiomycetes of North Carolina. Jour.
E. Mitchell Sci. Soc. 35: 118.

Colley, R. H. 1925. A biometric comparison of the Urediniospores of Cronartium ribi-
cola and Cronartium occidentale. Jour. Agr. Res. 30: 283-291.

Coons, G. H. 1912. Some investigations of the cedar rust fungus, Gym. juniperi-
virginianae. Neb. Agr. Exp. Sta. Rept. 25: 215-245.

Craigie, J. H. 1927. Discovery of the function of pycnia of the rust fungi. Nature
120: 765-767.

Crowell, I. H. 1934 a. The hosts, life history and control of the cedar-apple rust
fungus, Gymnosporangium juniperi-virginianae Schw. Jour. Arnold Arb. 15:
165-232.

. 1934 b. The hosts, life history and control of Gymnosporangium clavipes
C. & P. Jour. Arnold Arb. 16: 368-409.

. 1940. The geographical distribution of the genus Gymnosporangium.
Can. Jour. Res. 18: 469-488.

Curtis, M. A. 1867. Geological and Natural History Survey of North Carolina.
Pt. III. Botany. p. 121. Raleigh, N. C.

Dodge, B. O. 1918 a. Studies in the genus Gymnosporangium. I. Mem. Brooklyn
Bot. Gard. 1: 128-140.

. 1918 b. Studies in the genus Gymnosporangium. II. Bull. Torrey Bot.
Club 45: 294-295.
1918 c. Studies in the genus Gymnosporangium. III. Mycologia 10:

182-193.
. 1931. A destructive red-cedar rust disease. Jour. N. Y. Bot. Gard. 32:
101-108. '
- 1934. Witches’ brooms on southern white cedar. Jour. N. Y. Bot. Gard.
35: 41.

Farlow, W. G. 1880. The Gymnosporangia or cedar apples of the United States.
Anniv. Mem. Boston Soc. Nat. Hist. pp. 1-38. pl. 1-2.
- 1886 a. Development of Roestelia from Gymnosporangia. Bot. Gaz. 11:
189-190.

PRINCE: GYMNOSPORANGIUM Nipus-Avis THAXTER 519

. 1886 b. The development of the Gymnosporangia of the United States.
Bot. Gaz. 11: 234-241.
. 1905. Bibliographical Index of North American Fungi. 1: 68. Carnegie
Inst. Washington, D. C.
Fernald, M. L. & L. Griscom. 1935. Three days of botanizing in southeastern
Virginia. Rhodora 37: 131-133. pl. 332-333.
Fischer, Ed. 1891. Uber Gymnosporangium Sabinae (Dicks.) and Gymnosporangium
confusum (Plowright). Zeitschr. Pflanzenkr. 1: 271.
. 1895. Die Zugehérigkeit von Aecidium penicillatum. Hedwigia 34: 1-6.
Fraser, W. P. 1925. Culture experiments with heteroecious rusts in 1922, 1923, and
1924. Mycologia 17: 84.
Giddings, N. J. 1918. Infection and immunity in apple rust. W. Va. ae Exp. Sta.
Bull. 170: 9.
Hartley, C. 1913. Bark rusts of Juniperus virginiana. Phytopath. 3: 249.
Hedgcock, G. G. 1940. Notes on distribution of fungi collected in southeastern
United States in 1938 and 1939. U.S. Dept. Agr. Plant Dis. Rept. 24: 322.
Hubert, E. E. 1916. Celluloid cylinders for inoculation purposes. Phytopath. 6:
447-450.
Hunter, Lillian M. 1936. Morphology and ontogeny of the spermogonia of the
Melampsoraceae. Jour. Arnold Arb. 17: 115-132.
Kern, F. D. 1910. The mon Daley, of peridial cells of the Roesteliae. Bot. Gaz.
49: 445-452.
. 1911. A biologic and taxonomic study of the genus Gymnosporangium.
Bull. N. Y. Bot. Gard. 7: 391-483.
, H. W. Thurston, C. R. Orton, & J. F. Adams. 1929. The rusts of
Pennsylvania. Penn. Agr. Exp. Sta. Tech. Bull. 239: 20.
Levine, M. N. 1923. A statistical study of the comparative morphology of biologic
forms of Puccinia graminis. Jour. Agr. Res. 24: 539-567.
MacLachlan, J. D. 1935 a. The hosts of Gymnosporangium globosum Farl. and
their relative ‘susceptibility. Jour. Arnold Arb. 16: 98-142.
. 1935 b. The dispersal of viable basidiospores of the Gymnosporangium
rusts. Jour. Arnold Arb. 16: 411-422.
. 1936. Studies on the biology of Gymnosporangium globosum Farl. Jour.
Arnold Arb. 17: 1-24.
, & I. H. Crowell. 1937. Control of the Gymnosporangium rusts by
means of sulphur sprays. Jour. Arnold Arb. 18: 149-163.
Maerz, A. & M. R. Paul. 1930. A Dictionary of Color. New York.
Millspaugh, C. F. & L. W. Nuttall. 1896. Flora of West Virginia. Publ. Field
Mus. Bot. Ser. 1: 129.
Munns, E. N. 1938. The Ee of important forest trees of the United States.
U. S. D. A. Misc. Publ. 287:
Palmer, E. J. 1925. Synopsis of ud American Crataegus. Jour. Arnold Arb. 6:
5-28.
Prince, A. E. 1943. Basidium formation and spore discharge in Gymnosporangium
nidus-avis. Farlowia 1: 79-93.
, & F. H. Steinmetz. 1940. Gymnosporangium rusts in Maine, and their
host relationships. Univ. Me. Stud. 2d Ser. No. 50: 3-10.
Rathay, E. 1883. Untersuchungen iiber die Spermogonien der Rostpilze. Denkschr.
K. Akad. Wiss. Wien. Mat.-Nat. Kl. Abt. 2. 46: (1-51) 22-23.
Ray, W. W. 1940. Notes on Gymnosporangium in Oklahoma. Mycologia 32: 572.
Rehder, A. 1940. Manual of Cultivated Trees and Shrubs. New York.
Sudworth, G. 1915. The cyfress and juniper trees of the Rocky Mountain Region.
U.S. D. A. Dept. Bull. 207.
Thaxter, R. 1887. VII. On certain cultures of Gymnosporangium with notes on
their Roesteliae. Proc. Amer. Acad. Arts and Sci. 22: 259-268.

520 FarLowiA, VoL. 2, 1946

1889. Notes on cultures of Gymnosporangium made in 1887 and 1888.
Bot. Gaz. 14: 163-172.
1891. The Connecticut species of Gymnosporangium (cedar apples).
Conn. Agr. Exp. Sta. Bull. 107: 2-6.
. 1892. The Connecticut species of the Gymnosporangium (cedar apples).
Conn. Agr. Exp. Sta. Rept. 1891: 164.
1893. Fungi described in recent reports of the Connecticut Experiment
Station. Jour. Myc. 7: 278-280.
Weimer, J. L. 1917. Three cedar rust fungi. Their life histories and the diseases
they produce. Cornell Univ. Agr. Exp. Sta. Bull. 390: 523-524.

Sa

Fig.
Fig.
Fig.
Fig.
Fig.

Fig.

FarLowIA, VoL. 2, 1946

EXPLANATION OF PLATE I

1. Leaves and fruit of Amelosorbus Jackii infected by G. nidus-avis, the abundant,
large lesions indicating the extreme susceptibility to inoculum from Colorado.

2, Leaves and stems of Amelanchier canadensis bearing numerous aecia, evidence
that this host is very susceptible to G. nidus-avis from Saskatchewan, Canada.

3. Portion of a cross section of a leaf of Amelanchier stolonifera, illustrating am-
phigenous spermogonia of G. midus-avis.

4. Invasion zone bordering healthy tissue in portion of Wealthy apple leaf sec-
tion. Note abortive spermogonium.

5. A longitudinal section of a single spermogonium showing a darkly stained
receptive hypha among the periphyses.
6. A longitudinal section of a young aecium.

523

GYMNOSPORANGIUM Nipus-AvIs THAXTER

PRINCE

PraTeE I

524 FarLowlA, VOL. 2, 1946

EXPLANATION OF PLATE II

Fig. 1. A dense broom caused by the telial stage of G. nidus-avis on Juniperus vir-
giniana var. crebra.

Fig. 2. Foliicolous telia on leaves of Juniperus horizontalis.

Fig. 3. Caulicolous telia of G. midus-avis on infected branch of Juniperus virginiana.

Fig.4. A dense broom caused by the telial stage of G. nidus-avis on Juniperus
scopulorum.

PRINCE: GYMNOSPORANGIUM Nipus-AVIS THAXTER

Prate II

525

2(4) :527-567 FARLOWIA Jury, 1946

THE BOLETINEAE OF FLORIDA WITH NOTES ON
EXTRALIMITAL SPECIES

IV. THE LAMELLATE FAMILIES (GOMPHIDIACEAE
PAXILLACEAE, and JUGASPORACEAE)

R. SINGER

As Lenzites is close to Coriolus in the Aphyllophorales, so, in the Bole-
tineae, some lamellate families come closer to certain genera of the Bole-
taceae than other tubulose groups, e.g. Paxillus seems to be closer to
Gyrodon than Strobilomyces or Porphyrellus. The relationship between
Paxillus and the Boletaceae was actually recognized long ago, but the re-
lationship between the Gomphidiaceae and the Boletineae had been doubted
by some mycologists (including Gilbert), and it was necessary to draw at-
tention to the similarities between these groups. (See Ann. Mycol. 40:.555.
1942.) As for the Jugasporaceae, they seem to.be intermediate, according
to their characters, between the Boletineae and the Agaricineae (Rho-
dophyllus, Rhodogoniosporaceae; Rhodocybe, Tricholomataceae). For
reasons we shall point out later (p. 548), we have always preferred to
consider them as a family of the Boletineae.

The following monograph of the three families named above is written
in the same pattern as the three preceding monographs on Florida Bole-
tineae.! For a general introduction and a key to the families, the reader
is again referred to Part I (Farlowia 2: 97. 1945). In addition to the
acknowledgments in my first monograph (/.c.), I wish to express my thanks
to Dr. L. R. Hesler, University of Tennessee, and especially to Dr. A. H.
Smith, University of Michigan, for the loan of important material of
Gomphidius without which I would never have been able to prepare that
genus for North American Flora, a manuscript on which the following
treatment of the Gomphidiaceae of Florida is based.

GOMPHIDIACEAE R. Maire (1933)

Characters of the family: Pileus glabrous or tomentose or farinaceous,
viscid to glutinose or more rarely dry, small to rather large; hymenophore
lamellate, consisting of rather thick, decidedly decurrent gills with fre-
quently obtuse edges (less so in subgenus Chroogomphus), waxy-subgela-
tinous to tender-fleshy consistency, and rather thick trama; lamellae sub-
distant to distant, more or less arcuate, gray to fuliginous when mature;
spore print black; spores melleous to gray under the microscope, fusoid to
subcylindric, smooth; cystidia large, projecting, usually distinctly in-
crusted, sometimes colored and with very thick walls (Gomphidius to-

1 Singer, R. The Boletineae of Florida with notes on extralimital species. I. Farlowia
2(1): 97-141. 1945; II. Farlowia 2(2): 223-303. 1945; III. Amer. mid]. nat. (in press).

527

528 FarLowiA, VoL. 2, 1946

mentosus), numerous to very rarely scattered (when old); trama of the
gills more or less bilateral, not always clearly divergent, and very fre-
quently partly interwoven, at least the hymenopodium interwoven but
the trama never homogenously interwoven, nor intermixed, nor regular 7) ;
hyphae without clamp connections; stipe versiform, equal or swollen, etc.,
in most species and specimens with a discolored base which is pink or rich
lemon yellow to orange yellow both inside and on the surface; veil present
or absent, more often present, and then either entirely glutinous, or tender
and fibrillose, or mealy and consisting of a loose pseudoparenchyma (Cysto-
gomphus), sometimes forming an annulus; glandulae on the surface of the
stipe present in one subgenus of Gomphidius (Laricogomphus); growing
on earth and forming mycorrhiza with conifers, practically all species
extratropical.

KEY TO THE GENERA AND THEIR SUBDIVISIONS

A. Veil consisting of spherocysts. Gen. Cystogomphus
A. Veil, if present, not consisting of spherocysts. Gen. Gomphidius
B. Context of the pileus ochraceous to orange (though at times in young speci-
mens rather pallid), more rarely salmon to pink; veil constantly present,
consisting of strictly parallel, pigment-incrusted hyphae, macroscopically
never entirely glutinous, never hyaline; subhymenium filamentous-intermixed

and dense to very dense; mediostratum rudimentary in young specimens,

indiscernable in adult ones. Subgen. Chroogomphus
C. Pileus dry to subviscid in wet weather, not shining in dry weather, more
or less tomentose to fibrillose. Sect. Floccigomphus

C. Pileus viscid in wet weather, shining in dry weather, not tomentose and
not fibrillose except for traces of the veil on the margin.
Sect. Viscogomphus

B. Context of the pileus white, more rarely pink to salmon, or becoming pink
on exposure; veil absent, or consisting of subparallel-subinterwoven, thin
hyphae which are not incrusted by pigment, macroscopically hyaline to white,
and often entirely glutinous, sometimes blackening with age; subhymenium
filamentous-cellular to filamentous, moderately dense; mediostratum of young
specimens distinct, less distinct in older ones.

D. Veil visible only in the primordium, fugacious and not leaving any traces
in adult specimens; dermatocystidia of the stipe fasciculate and the
fascicles forming glandulae which make the stipe appear fibrillose or
furfuraceous (though these macroscopical characters are not always easy to
ascertain) ; mycelium (of G. maculatus) connected with Larix-mycorrhiza.

Subgen. Laricogomphus

D. Veil covering the lamellae of the young specimens and leaving more or
less distinct traces even in old specimens; dermatocystidia of the stipe
not fasciculate, no fibrils or glandulae made up of dermatocystidia present
on the surface of the stipe; mycelium in nature not connected with Larix

but forming mycorrhiza with other conifers. Subgen. Myxogomphus
E. Spores (with the exception of immature and aborted ones) longer
than 14 uw. Sect. Macrosporus

E. Spores, or majority of spores, shorter than 14 yu, or just reaching 14 wu.
Sect. Microsporus

* See also observations for subgenus Chroogomphus, sect. Viscogomphus, p. 529.

SINGER: THE BOLETINEAE OF FLORIDA 529

Cystogomphus Sing., Ann. Mycol. 40: 51. 1942.

Characters of the genus: these are evident in the key above. Type spe-
cies Cystogomphus Humblotii Sing. No species is recorded in Florida.

Gomphidius Fr., Epicrisis, p. 319. 1938.

Characters of the genus: these are evident in the key above. Type spe-
cies Gomphidius glutinosus (Schaeff. ex) Fr.

Subgen. Chroogomphus Sing., (in print).

Characters of the subgenus: these are evident in the key above. Type
species Gomphidius rutilus (Schaeff. ex Fr.) Lundell & Nannf.

Sect. Floccigomphus Imai, Jour. Fac. Agr. Hokkaido Imp. Univ. 43: 285. 1938.

Characters of the section: these are evident in the key above. Type
species Gomphidius tomentosus Murr. Other species G. stbiricus Sing.;
G. leptocystis Sing. In Florida none of these occurs.

Sect. Viscogomphus Imai, Jour. Fac. Agr. Hokkaido Imp. Univ. 48: 284. 1938.

The characters of this section are obvious in the key above. The type
species is the same as in subgenus Chroogomphus.

There have been various and seemingly controversial statements con-
cerning the subhymenial and tramal structure of this section as compared
with other sections. As for the trama, Fayod (1889) says it is bilateral
quite generally in Gomphidius, and expresses surprise that De Seynes’
figures do not show any bilateral trend. Lohwag (1937) finds the trama
typically bilateral (truly bilateral as we call it in this paper) in G. gluti-
nosus, and not distinctly bilateral, becoming irregular in G. rutilus. Elrod
and Blanchard (1939) studied G. maculatus and found that the trama
is “slightly and loosely interlaced, with a tendency toward a subparallel
condition in many places. There is no sign of a differentiated mediostratum
and laterostratum.”’ In regard to the subhymenium the situation is still
more obscure. Fayod (J.c.) says there is none. Lohwag (/.c.) and Singer
(1942) state that there is one, and that it is pseudoparenchymatic in G.
rutilus, and filamentous in G. glutinosus. Elrod and Blanchard (1939)
and also Singer (1938) state that it is filamentous in G. maculatus. There
is in all species, no doubt, a more or less definite layer immediately be-
neath the hymenium, but more recent re-examination by the writer tends
to modify the above statements, and partly reverse them. It is true that
in a not too thin section of the lamella of G. vinicolor the subhymenium
appears to be coarsely cellular, but more detailed examination of the base
of the hymenium shows a dense tissue of strongly interlaced, curved, irregu-
lar, strongly coherent hyphae which look cellular where they run for a
short distance in a vertical direction. However, very few of the individual
hyphae are short enough to be called subisodiametric and then they are
very small, or very irregular, e.g. the widened hyphae from which two

‘

530 Fartowl4, VoL. 2, 1946

branching hyphae arise, or the basal cells of the basidia. The subhymenium
is not well differentiated from the underlying formation, and, in a sense,
it may be stated that there is none. If we reserve the name subhymenium
for a thin set of hyphae, continuing the basidia from their base downward,
we cannot call it pseudoparenchymatic. Jt is filamentous-intermixed. As
for the subhymenium of G. glutinosus, we may say that on superficial ex-
amination it always appears to be filamentous, but when the subhymenial
cells are examined carefully and individually, they frequently turn out
to consist partly of chains of very small globose bodies, and the same con-
dition has been observed recently by the writer in G. maculatus.

The trama proper is not a priori different in G. vinicolor, G. glutinosus,
G. roseus, and G. maculatus (the hymenophoral anatomy of G. rutilus
does not differ from that of G. vinicolor). The difference pointed out by
Lohwag and Peringer (1937) is not so striking, and more a matter of
degree and of individual development. The mediostratum of G. vinicolor
is very strongly reduced, so much so that no darker zone is seen in the
pallid middle layer, called, as a whole, mediostratum by the above-cited
authors. The irregular, interlaced accompanying zone between this and
the subhymenium is called lateral stratum by all authors though I should
think that it would rather deserve the name hymenopodium, since most of
the looser central portion of the trama (Lohwag and Peringer’s medio-
stratum) actually corresponds to the loose lateral stratum of the boletes.
This will also make understandable the complicated tramal structure of
G. glutinosus. In this latter species, we found a well-developed, melleous
mediostratum, and there is no choice, in this case, but to call the hyaline
looser layers at its sides the lateral stratum. The next-following layer
between the lateral stratum and the subhymenium proper differs from the
lateral stratum in being again more interlaced and dense. This layer is
neither a part of the lateral stratum nor a part of the subhymenium. It is
the hymenopodium, Lohwag and Peringer (/.c.) in the case of G. glutino-
sus, do not distinguish between this and the subhymenium proper, and,
consequently, what they call subhymenium is the subhymenium plus the
hymenopodium.

The gradual difference, therefore, between the trama of G. glutinosus
and G. vinicolor (and G. rutilus) consists in the stronger or weaker develop-
ment of the mediostratum proper on one hand, and the more or less decided
tendency toward an irregular, interlaced condition in all or some layers on
the other. It seems that other species may be intermediate between these
extreme cases, for in my notes I described the trama of G. roseus exactly
as it is in G. glutinosus though without mentioning the existence of a
mediostratum, and as we saw above, Elrod and Blanchard did the same
in the case of G. maculatus. However, it is obvious that the structure of
the trama in the Gomphidiaceae, as in general in most Boletineae, is sub-
ject to drastic changes during the individual development of the carpo-
phore, and our specimens may have been somewhat too old. Though the
photographs published by Elrod and Blanchard do not show much as far

SINGER: THE BOLETINEAE OF FLORIDA 531

as the exact structure of the various layers goes, Lohwag and Peringer’s
figures 11-13 show this difference in different stages.

Another important fact not indicated in the literature on Gomphidiaceae,
is the change in color of the spores when these are preserved in the her-
barium for a considerable time. There is a specimen called Gomphidius
viscidus [described here as G. rutilus (Schff.) Fr. ssp. alabamensis] col-
lected ninety-five years ago and preserved in the Curtis Herbarium (FH)
with a spore print on blue paper attached to it. The spores of the print
are now ferruginous macroscopically, and greenish melleous under the
' microscope. This discoloration should be kept in mind when older her-
barium material is described. I have never seen any of Fayod’s specimens
of Gymnogomphus Fay., but the description suggests that some old speci-
mens of G. rutilus or of its group were misinterpreted and a new genus
was created because of this fact.

KEY TO THE SPECIES

A. Mycelium pink-salmoneous; base of the stipe usually not markedly discolorous

(yellow).
B. Middle portion of the cystidia strongly thick-walled, apex and base thin-
walled. 1. Gomphidius vinicolor (in Florida: ssp. jamaicensis)

B. Middle portion of the cystidia not markedly thick-walled, wall of the cystidia
nearly equal, thin to very slightly thickened (not more than 1.7 » at most).
Gomphidius ochraceus

A. Mycelium never pink-salmoneous, but pale yellowish white to “Isabella color”;
base of the stipe frequently markedly discolorous, yellow.
2. Gomphidius rutilus (in Florida: ssp. alabamensis)

1. Gomphidius vinicolor Peck ssp. jamaicensis (Murr.) Sing. comb. nov.
Gomphidius jamaicensis Murr. Mycologia 10: 69. 1918.
Gomphidius alachuanus Murr., Jour. Elisha Mitch. Sci. Soc. 55: 367. 1939.

Prate I, FIG. 4-7.

Pileus between “Natal brown” and “Army brown” with some tints of
“light vinaceous cinnamon” to “Mikado brown” intermixed, or almost
“chestnut,” viscid, during dry periods “liver brown,” “carob brown,’ or
sometimes somewhat darker than these and shining, sometimes pallescent
on the disk and becoming “Kaiser brown” or even “ferrugineous,” glab-
rous, naked, smooth, conic-convex, then convex, eventually usually more.
or less flattened, or with depressed center, obtuse, (24)—50-(98) mm. diam.
— Hymenophore between ‘“‘Prout’s brown” and “mummy brown,” initially
dirty ochraceous-alutaceous, eventually black from the spores, the lamellae
broadest in the middle (4-10 mm.), subclose to subdistant, arcuate, de-
current, intermixed (one lamellula after either one or two through-lamel-
lae), slightly transversely venose, only occasionally and very sparsely
forked; spore print black. — Stipe “light vinaceous cinnamon” to “buff
pink,” or “vinaceous rufous,” the base usually “cinnamon”? to “clay color,” -
‘ight ochraceous buff,” or “ochraceous buff,” sometimes at places becoming
“Jasper red” or “orange cinnamon” to “cacao brown,” not viscid, glabrous

532 Fartowla, VoL. 2, 1946

except for the veil, smooth but more or less grooved when drying, naked,
gradually tapering downward, or subequal, solid, 36-100 x 4-15 mm.;
veil either forming a definite, fibrillose-woolly, narrow annulus which is
concolorous with the remaining surface of the stipe, or reduced to a mere
fibrillose zone at the apex of the stipe; mycelium pink. — Context of the
pileus and upper portion of the stipe “pale ochraceous salmon”’ to “‘ochrace-
ous salmon,” or “pinkish buff” in pileus and “cinnamon buff” in upper
part of stipe, in the base “primuline yellow” with a tinge of “yellow ocher”
or “clay color,” often somewhat umber immediately below the cuticle,
fleshy, rather firm, not as soft and watery as in G. glutinosus; odor none,
or agreeable when quite young, recalling apples or apple sauce; taste mild.

Spores melleous-grayish to gray, varying from a pure cinereous to fuli-
ginous gray, ellipsoid to fusoid, not cylindric, smooth, (in carpophores
with the spores of small average size, the short spores are usually the
broadest, and the long ones the slenderest), 17-21.8 x 5.8-8; basidia
(2)—4-spored, 34-46 x (8.2)-9.2-l3u: cystidia numerous, on edge as well
as on sides of lamellae, slightly incrusted on the inside of the walls in
many of them, the incrustation melleous, and in almost all of them there
is a strong resinous or occasionally granular, fulvous-castaneous incrusta-
tion that covers the apices, otherwise smooth, thick-walled, the wall hyaline,
thinner or quite thin at the apex which is frequently capitate, rarely clavate,
and always broadly rounded, cylindric or slightly fusoid below, and there
the walls reaching 2-5.5 » in diameter but becoming thin near the very
base, the entire length varying between 115 and 150 p, the diameter from
13.5 to 22.5u, the base reaching into the lateral stratum, the upper portion
projecting well beyond the level of the sterigmata; subhymenium filamen-
tous-intermixed; ¢rama consisting of an indistinct mediostratum reduced
to a central string of a few parallel or subinterwoven hyphae which are all
hyaline and cylindric, but most of the trama proper (i.e. excluding the
hymenopodium) formed by a hyaline, loosely arranged, partly subirregular
layer that with age becomes homogeneous and increasingly irregular while
any trace of a mediostratum disappears; Aymenopodium obscured and
thickly covered by an adhesive (sordid melleous brown) pigment that
precipitates in form of an amorphous intercellular pigment when the prepa-
ration is boiled in 10% KOH, differing in this regard from the trama
proper, and differing, also, in its much denser arrangement, the hyphae
being more variable in diameter and running in all directions to form a
subintermixed broad zone along the subhymenial layers; all hyphae with-
out clamp connections; pellicle of repent, subparallel-interwoven, with
fulvous-castaneous incrustation, elongate, clampless hyphae; mycelium
consisting of filamentous hyphae which are beset with button-like orna-
mentations.

Chemical reactions: KOH on pileus, little reaction; on context, between
“pale rosolane purple’ and “thuilite pink,” becoming “spinel red’; in
base, “rich brown.” — NH3;-~vapors on context, “liseran purple” and ‘‘pale
rosolane purple’; in base olive. —NH.,OH on pileus, little reaction; on

SINGER: THE BOLETINEAE OF FLORIDA 533

context between “pale rosolane purple” and “thuilite pink,’’ becoming
“spinel red”; in base brownish. — H2SQu,, little reaction but the treated
portions of the base becoming deep fulvous yellow when dried subsequently.
—FeS0O, in all parts, ‘deep olive” to “dark olive.”

Habitat: In woods, in Florida under Pinus taeda, possibly also P. aus-
tralis, and P. palustris, but not observed in pure stands of any of the latter
species; on sandy soil, gregarious, fruiting from November to January.

Distribution: North Florida and Alabama, also in the West Indies (Ja-
maica).

Material studied: Fla.: Alachua Co., Gainesville, Murrill et al. F 10213, F 18431,
F 20040 (all FLAS, det. Murrill) ; Planera Hammock near Gainesville, Murrill et al.
det. Murrill F 19517, F 20086 (FLAS) type of G. alachuanus, Newnan’s Lake,
East of Gainesville, West & Murrill F 18402 (FLAS); my own collections compared
with the type and partly from the type locality: F 1657, F 1657a, F 1657b, F 1657c,
F 1657d, F 1657e (all FH).— Ala.: Lee Co. Auburn, Fall 1897, F. S. Earle & C. F.
Baker (G. viscidus) (NY).— Jamaica, W.I.: Cinchona, in 5000 ft. elevation, Earle
(352), type (NY).

This species is divided into several geographic races and mycoecotypes
(in the sense of Singer, Sovietsk. Botan. 1940, (5-6):262-269). The
subtropical (G. alachuanus) and tropical-montane (G. jamaicensis) form,
having subclose to subdistant lamellae and forming mycorrhiza with pines
of the section Australes, should properly be called ssp. jamaicensis while
the northern and western forms occurring with such pines as Pinus resinosa,
and P. radiata, and differing morphologically in having more distant lamel-
lae and frequently umbonate pileus, should be distinguished under separate
subspecific names. The northern subspecies of G. vinicolor Murr. has the
same characteristic cystidia with thick walls in the middle portion as has
been observed in the southern form. It should not be confused with G.
ochraceus Kauffm. and allied forms such as G. superiorensis Kauffm. &
Sm., which also have pink mycelium and similar colors under certain con-
ditions and in certain stages, but lack the thick cystidial walls. G. tomento-
sus Murr., which has thick-walled cystidia, has the walls nearly equally
thick over the entire length of the cystidia, and not merely in the middle.
G. rutilus and allied forms (ssp. alabamensis, the form called G. furcatus
by Kauffman non Peck) is easily distinguished by the color of the mycelial
tomentum and the thin-walled cystidia.

2. Gomphidius rutilus (Schaeff. ex Fr.) Lund. & Nannf., Fung. Exs. Suec. 409. 1937.
Agaricus rutilus Schaeff. ex Fr., Syst. Mycol. 1: 315. 1821.
Cortinaria rutila S. F. Gray, Nat. Arr. Brit. Pl. 2: 629. 1821.
Gomphidius viscidus (L. ex) Fr., Epicrisis, p. 319. 1838.
Paxillus pubescens Ellis, Bull. Torr. Bot. Cl. 6: 76. 1876. (= ssp. alabamensis)
Gomphidius testaceus (Fr.) Britz., Hymen. Siidbay. 4: 133. 1885. (=var.

testaceus)

Gomphidius litigiosus Britz., Hymen. Siidbay. 9: 14. 1893. (= var, litigiosus)
Gomphidius alabamensis Earle, Torreya 2: 54. 1902. nomen nudum (= ssp.

alabamensis)
Gomphidius viscidus forma columbiana Kauffm., Mycologia 17: 122. 1925.

Ssp. typicus.

534 _ Fartowia, Vou. 2, 1946

The type subspecies, the most common Gomphidius in Europe and in
parts of the Rocky Mountain region, does not occur in Florida. It differs
only slightly from the following subspecies (alabamensis) in having the
pileus almost constantly umbonate, and in its geographic area.

Var. testaceus Fr., Epicr., p. 319. 1838 (ut varietas G. viscidi)

This is a small umbonate form with very light, almost white mycelium,
and with the stipe lacking the yellow color of the surface and of the con-
text. I have collected it under pines in mixed beech woods in Europe,
but not in this country. Good typical specimens are preserved in the
Hoehnel Herbarium (FH), from Tullnerbach-Pressbaum, Wiener Wald,
Austria (F. v. Hohnel B 1467). The specimens described and illustrated
under the name of Gomphidius viscidus var. testaceus Fr. in Icones Far-
lowianae, pl. 70, are a form of G. ochraceus Kaufim. G. viscidus fa. testacea
Kavina seems to belong either to var. pulcher or var. fulmineus. .

Var. pulcher Killerm. Denkschr. Bay. Bot. Ges. 21: 58. 1940 (ut varietas G. viscidi)
Var. litigiosus (Britz.) Sing. comb. nov.

These two varieties, with differently colored pilei, are both described
from Bavaria, not reported from America.

Var. fulmineus Heim, Treb. Mus. Ciénc. Nat. 15: 68. pl, 1, fig. 3. 1934 (ut varietas
G. viscidi) ,

This was described from Spain; it is possibly specifically different.

Kauffman described a f. columbiana which, as he thought, represents
the American form of G. viscidus (G. rutilus ssp. typicus in our nomencla-
ture). However, the examination of his herbarium shows that he mixed
two different species under this name, one being an umbonate form of G.
ochraceus, the other (including the type specimen) being G. rutilus ssp.
typicus. It appears that Kauffman was under the impression that the
European G. rutilus always is a large plant, which is definitely not true.
Thus there remain only two differences between the European and the
American form which escaped Kauffman. One is the frequently brighter
color of the base of the American specimens, a character that aside from
its vagueness does not seem to be quite constant. The other is a phyto-
sociological character which could possibly be physiological: the American
form is associated with pines of the series Imsignes while the European
form is associated with pines of the series Lariciones. If it could be
demonstrated by experiments that the European form does not form
mycorrhiza with Pinus contorta as readily as it does with P. silvestris, it
would make the American form a mycoecotype lacking morphological
differences (comparable with the races of rusts). However, as long as
such physiological differences have not been shown, I prefer to list f.
columbiana as a plain synonym of G. rutilus ssp. typicus,

SINGER: THE BOLETINEAE OF FLORIDA 535

Subsp. alabamensis (Earle ex) Sing. subsp. nov.
Gomphidius alabamensis Earle, Torreya 2: 54. 1902 (nomen subnudum)
Paxillus pubescens Ellis, Bull. Torr. Bot. Cl. 6: 76. 1876.
Prats I, Fic. 1-3.

Tomento myceliali cremeo-albo vel melleo-isabellino; basi plerumque laete flavo
in siccis; umbone plerumque nulla; cystidiis tenuitunicatis, incrustatis; ceterum Gom-
phidio rutilo et G. vinicolori analogus.

Pileus-“light russet vinaceous,” “cinnamon buff,” “vinaceous cinnamon,”

“testaceous,” sometimes reaching “chocolate,” “Hay’s brown” or “dark
vinaceous brown,” assuming at times colors like “pecan brown” or “our-
plish vinaceous” after drying, usually a lighter reddish-brown in dried
specimens than G. jamaicensis, glabrous, more rarely with cracking pellicle
showing orange flesh between the fragments, viscid, naked, shining when
dry, convex to plane, eventually often depressed and turbinate, obtuse,
rarely indistinctly umbonate, 30-100 mm. broad, mostly between 30 and
70 mm.— Lamellae “avellaneous,”’ “clay color,’ “ochracéous tawny,”
later “testaceous,” “snuff brown,” or “sayal brown,” and finally darker
sooty from the spores, arcuate, then descendant, broad, subclose to dis-
tant, entire, thick, inserted with very short ones, occasionally some forked
near the stipe, decurrent. Spore print “hair brown” to “deep grayish olive”
in thin to moderate layer, very nearly black in good print. — Stipe vary-
ing from sordid whitish to “Congo pink” or subconcolorous with the veil
-which is “ochraceous buff” or “capucine buff,” or streaked with vinaceous
brown to almost tawny, with a bright yellow color very often at the very
base of the dried stipes, mycelium cream-white to melleous-yellow or
“Isabella color,” strigose, the upper surface fibrillose and often showing
a cortina-like annulus at the apex, glabrescent below, the veil often spurious,
the outer fibrils somewhat slippery-sticky-subviscid in very wet weather
but never gelatinous, inside solid, the shape equal with attenuate base, or
fully equal and cylindric, or occasionally compressed, or tapering from the
apex downward, straight or curved, 25-90 x 8-18-(40) mm. — Context
of the pileus thick in the center, thin on margin, varying from almost
white (though usually showing a slight brownish-orange tinge) to “buff
pink,” soft; of stipe “Congo pink” to subconcolorous with the pileus, the
base mostly more or less yellow inside. Odor none to slight. Taste mild.

Spores (16)—18—21 x 5.8-6.5 mw, subcylindric-fusiform, smooth, melleous,
grayish-melleous; basidia 4-spored, 40-55 x 11-12.8 3 cystidia abundant,
cylindric to cylindric-ventricose, or subcapitate, with narrowed base and
rounded apex, incrusted or more rarély not incrusted, with or without
granular contents, 95-150 x 12-18 p», on sides and edges; subhymenium, —
hymenopodium, lateral stratum, and mediostratum as in G. jamaicensis
(the subhymenium perhaps very slightly looser); cuticle of incrusted re-
pent filamentous hyphae; Ayphae without clamp connections.

Habitat: In mixed and pine woods under Pinus (species of the series
Insignes and Australes), terrestrial, gregarious, fruiting from October to
January.

536 FarLowIA, VoL. 2, 1946

Distribution: Maryland to South Carolina and west to Alabama and
Tennessee, south to north Florida. In Florida rare, and observed only
once in the western part of Alachua Co.

Material studied: Fla.: Near Seven-Mile-Church, 7 mi. west of Gainesville, Jan-
uary 8, 1940, W. A. Murrill & J. R. Watson, det. Murrill (G. alachuanus) (ex Herb.
Erdman West, NY).—N.J.: Newfield, Oct. 10, 1887, on ground in pine woods,
Authentic material of P. pubescens Ell. (NY).—Md.: South River near Annapolis,
Oct. 6, 1919, J. R. Morris, comm. H. A. Kelly (276), det. L. C. C. Krieger (Gom-
phidius spec.) (MICH); Takoma Park, Oct. 12, 1918, Kauffman (G. furcatus).—
N. C.: Chapel Hill, Oct. 1912, W. B. Cobb 582 (NY).—S. C.: Society Hill, Curtis
2855 (FH).—Tenn.: Ball Camp Pike, Nov. 1937, Hesler & Sharp 10986, det. A. H.
Smith (G. viscidus) (MICH); Dec. 25, 1941, L. R. Hesler (G. viscidus f. columbiana)
14120 (MICH); near Clinton, Nov. 11, 1934, L. R. Hesler 7792, det. A. H. Smith
(G, vinicolor) (FH, MICH).—Ala.: type of G. alabamensis, Auburn, Dec. 1900,
ELS. & F. S. Earle (2845) (NY).

Earle published G. alabamensis in a key to the species of Gomphidius,
but without a formal description. The characters given in the key do not
determine the species as such, but notes with some of his specimens which
he called G. viscidus, and the specimen which we consider the type of
G. alabamensis, do not leave any doubt as to what Earle had in mind
when he used the binomial Gomphidius alabamensis. The specimens of
G. furcatus Peck, however, belong rather to G. maculatus than to this
form, and we think, therefore, that Kauffman was wrong in identifying
G. alabamensis with G. furcatus.

EXTRALIMITAL SPECIES

Gomphidius ochraceus Kauffm., Mycologia 17(3): 119. 1925.
Gomphidius superiorensis Kauffm. & Smith, Papers Mich. Acad. Sci. Arts & Lett.

17: 170. 1933. (a variety)
A polymorphous species, close to G. rutilus, not occurring in Florida.
Subgen. Laricogomphus Sing., (in press)

Characters of the subgenus: these are evident from the data in the key.
Type species G. maculatus (Scop. ex Fr.) Fr. Not occurring in Florida.

Subgen. Myxogomphus Sing., (in press)

Characters of the subgenus: these are evident from the key. Type
species G. glutinosus (Schaeff. ex Fr.) Fr.

Sect. Macrosporus Sing., (in press)

Characters of the section: these are evident from the key. Type species
G. glutinosus (Schaeff. ex Fr.) Fr. This and the other species of this
section, viz. G. septentrionalis Sing., G. nigricans Peck, G. Smithii Sing.,

SINGER: THE BOLETINEAE OF FLORIDA 537

G. subroseus Kaufim., G. roseus (Fr.) Karst. do not occur as far south
as Florida.

Sect. Microsporus Sing., (in press)

Characters of the section: see key. Type species G. oregonensis Peck,
a western species. Another species may occur in Europe, none in Florida.

SPECIES EXCLUDENDAE
Gomphidius foliiporus Murr.
This is Phylloporus rhodoxanthus ssp. foliiporus (Murr.) Sing.
Gomphidius rhodoxanthus (Schwein.) Sacc.

This is the American type of Phylloporus rhodoxanthus (Schw.) Bres.

PAXILLACEAE R. Maire (1902)

Characters of the family: pileus subtomentose to tomentose, sometimes
viscid, small to large, the margin initially inrolled; hymenophore lamellose,
but the lamellae very frequently connected by anastomosing veins and
ridges, or the sides of the lamellae venose-rugose, more rarely the anasto-
moses broad and numerous and then the hymenophore resembles that of
Merulius, arcuate, usually rather narrow, yellow, orange, or light tan to
brown; spore print from nearly white to chamois or brownish; spores
rather small to medium (3-11.5 » long), smooth, ovoid, subglobose, ellip-
soid or oblong-ellipsoid; cystidia present or absent; trama more or less
bilateral; hyphae constantly with clamp connections; stipe very frequently
eccentric to lateral or wanting; veil, said to be present in Paxillus argen-
tinus, mostly none.. Habitat: on earth or on wood, or on sawdust, rarer in
the tropics but not entirely confined to extratropical regions, not forming
mycorrhiza with specific flowering plants.

The Paxillaceae are closely related to the Boletaceae, especially the genus
Gyrodon. Aside from the more individualized lamellae (even if they are ~
connected by anastomoses), the more involute margin of young speci-
mens, and the biological differences (Gyrodon and most of the Boletaceae
being mycorrhizal fungi while the Paxillaceae do not seem to be dependent
on symbiosis with specific forest trees, which does not mean that they
will never and under no circumstances whatever form anything comparable
with mycorrhiza-relationship), there are some minor differences that sep-
arate this family from Gyrodon. There is, for example, a slight but distinct
difference in the chemical reactions (see the descriptions of the species).
The greenish-yellow color of the hymenophores in Gyrodon is indicative
of its close relationship with the Boletaceae. There is also the occur-
rence, in some species of Gyrodon, of autoxidation causing a blue dis-
coloration of the hymenophore and the flesh. This latter reaction is,
although not a constant feature of all Gyrodon, a character common to

538 FarLtow1A, VoL. 2, 1946

many groups of the Boletaceae, while in the Paxillaceae this blue discolora-
tion is constantly absent. There also appears to be some difference between
the structure of the tube walls of Gyrodon and the trama of the lamellae
in Paxillus. In the former, the mediostratum is somewhat denser and
somewhat colored while the lateral stratum is distinctly divergent and’
hyaline in young material. In Paxillus, we found the lateral stratum of our
sections definitely strongly, interwoven, not clearly diverging, and of a
’ rather different aspect when compared with the lateral stratum of the
Gyrodon spp. —so different, indeed, that it is possible to consider it as
non-homologous with the lateral stratum of the Boletaceae. Instead, the
homology is with the hymenopodium of the Gomphidiaceae (see p. 529),
. which, however, may be difficult to prove, since the middle layer is always
homogenous. In this family, as well as in the Gomphidiaceae, the struc-
ture of the hymenophore trama changes strongly while the carpophores
mature.

I cannot see any good reason for splitting the genus Paxillus into several
genera, merely on the, for the most part imaginary, differences in tramal
structure. These species are otherwise so similar, exactly as the Gomphidii,
that there is no doubt about their close natural affinities. This is confirmed
by the chemical reactions registered by the author. One has only to com-
pare the reactions of Paxillus involutus (see Ann. Myc. 40: 58. 1942) with
these of P. panuoides (see this paper, p. 542). Crepidotus, which has been
compared with Paxillus panuoides, belongs in an entirely different sphere
of affinities, being very close to Ripartites (which also has been considered
as belonging to Paxillus by Quélet, Ricken, and others), and some other
genera. Phylloporus is too close to certain Boletaceae, especially Xero-
comus, to be retained in the Paxillaceae.

The Paxillaceae and the Gomphidiaceae are no doubt related, as may be
expected of two lamellae-bearing families within the suborder Boletineae.
Their tramal structure has some analogies. But biologically, in spore color,
spore size, in chemical characters, and, most important, in clamp connec-
tions, they are consistently different. In many ways the Gomphidiaceae
are comparable with the Strobilomycetaceae, and the Paxillaceae with the
Boletaceae.

KEY TO THE GENERA, SECTIONS, AND SPECIES KNOWN TO THE AUTHOR

A. Spore print at least “chamois” (Ridgway), or somewhat deeper brownish; lamel-
lae more or less anastomosing, at least in some portions of the hymenophore.
Gen. Paxillus

B. Spores up to 7.5 mw large; cystidia none.
C. Stipe present, large, tomentose. Sect. Atrotomentosi
D. Trama of the lamellae lilac, in NH,OH blue.
Paxillus polychrous
D. Trama of the lamellae not as above. Paxillus atrotomentosus

C. Stipe present, and then lateral and short, or absent.
Sect. Panuoides

SINGER: THE BOLETINEAE OF FLORIDA 539

E. Spores 3.8-6 x 3-4.5 4. Odor none or not remarkable.
3. Paxillus panuoides

E. Spores 3-4 x 1.7-2 wu. Odor sometimes persistent, disagreeable.
Paxillus Curtisii

B. Most spores more than 7.5 w long; cystidia present.
Sect. Involuti
Paxillus involutus

A. Spore print almost white (somewhat cream colored in thick layer) ; lamellae not
or little anastomosing (though strongly forked), not venose on their sides.
Gen. Hygrophoropsis

F. Spores 4.8-8 x 2.7-4.8 uw; basidia 19-40 x 5-9.5 uw; pileus 35-85 mm. in diam-
eter; stipe central or somewhat eccentric, very rarely strongly eccentric to
lateral; habitat in coniferous woods or in mixed woods under conifers, on
moss beds, or on coniferous wood, or on naked earth.

5. Hygrophoropsis aurantiaca

F. Spores 3.3-4.8 x 2.5-3.3 uw; basidia 16-20 x 3.7-6.8 w; pileus up to 31 mm. in
diameter; stipe strongly eccentric, lateral, or entirely wanting; habitat on
sawdust, in hollow trunks (e.g. of Naseer silvatica) but not on coniferous
trunks, also occurring on old frondose logs in dense hammocks.

4. Hygrophoropsis tapinia

Paxillus Fr., Gen. Hymen., p. 8. 1836.
Ruthea Opat., Comm. Bol., Wiegm. Arch. 2: 4. 1836.
Tapinia Karst., Hattsv., p. XXIII. 1879.
Tapinella Gilbert, Bolets, p. 67. 1931.
Rhymovis Rabenh., Kryptogamenfl. 1: 453. 1844.
Paxillopsis Gilbert, Bolets, p. 86. 1931 (nom. nud.), non Lange, Fl.
Agar. Dan. 4: 47. 1939,
Plicaturella Murr., N. Amer. Flor. 9: 172. 1910 (sec. Murrill).

Characters of the genus: these are evident in the key above. Type
species, P. involutus (Batsch ex Fr.) Fr. The only species thus far ob-
served in Florida is P. panuoides.

The nomenclatorial problem in the case of the genus Paxillus i is a rather
simple one. The genus Paxillus and the genus Ruthea, based on the same
plants, were both proposed in 1836, and it certainly was a good idea to
put Paxillus on the list of momina generica conservanda as proposed by
R. Maire to be accepted by the next Botanical Congress, since it may
turn out that Opatowski’s little-known name has a few days’ priority
over the generally accepted name Paxillus. Unfortunately, Gilbert’s chap-
ter about this family has caused many wrong citations and erroneous con-
ceptions due to some inaccuracies in his book “Les Bolets.” In his chapter
on the “Histoire Taxonomique du Genre Paxillus” (p. 64-69), he says:

“Persoon . . . a explicitement créé le genre Rkymovis sur le P. involutus,
pour les seules espéces €troitement affines aux Bolets. Quoique puisse en
dire Fries . . . , ce genre Rkymovis a la priorité. . . . Il est bien curieux

de constater que Earle (1909) et Murrill ne Vaient pas reprise.” Per-
soon, however, did not validly describe a genus Rkymovis though he may
ches had the intention of doing so in the future. “Si plures species,” he

540 FartowlA, VoL. 2, 1946

says (Mycologia Europaea 3: 63), “inveniuntur, huic [z.e. Agaricus involu-
tus| conformes, tunc ex iis genus distinctum efformari posset, RAymovis *
forte denominandum.” It is inconceivable that anyone who has read this
could possibly consider Rkymoxis as a published genus. It must be as-
sumed that Earle and Murrill did read what Persoon said. They, therefore,
knew that RAymovis was not published before 1844 by Rabenhorst, i.e.
eight years after Fries published Pavxdllus. Gilbert continues: ‘Berkeley
(1836) crée lui aussi, sur P. involutus, un genre Tapinia qui sera repris

comme sous-genre par Fries (1838) ... Karsten (1879) .. . démem-
brait ce Paxillus (= Tapinia): Paxillus pour le P. involutus et ses affines,
et Tapinia pour le T. panuoides. . . . Il est donc nécessaire de rejeter le

terme générique Tapinia, d’ailleurs illégalement employe.” If we check
on Gilbert’s “Histoire” again, it turns out that Berkeley in 1836 did not
use Tapinia as a genus but as a subgenus. Karsten’s Tapinia, therefore,
is legal, though in my opinion not worthy of generic distinction. This
whole situation was clear enough in the synonymy given in North American
Flora 10(3) : 146-147. 1917, and did not need to be “embrouillé” as late
as 1931.

Sect. Atrotomentosi Sing. sect. nov.
Stipite praesente, vellereo; sporis parvulis; cystidiis nullis.

Characters of the section evident from the key above and the Latin diag-
nosis. Type species P. atrotomentosus (Batsch ex Fr.) Fr. Other species
P. polychrous Sing. None of them found in Florida.

P. atrotomentosus is a well-known species, frequent in North America
but not reaching Florida. Specimens from New Hampshire show the fol-
lowing chemical reactions, recorded here for comparison with P. panuoides.
NH,OH on the surface of the pileus coloring the tomentum and the pallid
surface of the context, “dull lavender” to “slate purple” but leaving the
cuticle proper unchanged brown in most cases (at least in old specimens) ;
“dull lavender” to “slate purple” on the context of the pileus and the stipe
but changing more slowly than on the surface of the pileus. —KOH on
the surface of the pileus, a dark spot with a violet to lilac ring; on the con-
text of the stipe, greenish; on the tomentum of the stipe, blackish. —
H.SO, on the surface of the pileus where previously treated with KOH
or NH,OH becoming “yellow ocher,” then “Buckthorn brown” in a few
seconds, changing more slowly and more indistinctly at places where not
previously treated with KOH or NH,OH.— FeSO, darker brown, and
eventually greenish black on the pileus, and dull greenish gray on the
context of the stipe, eventually blackish.

Sect. Panuoides Sing. sect. nov.
Pileo subsessili.

°“Rhymovis” was a printing error, and was corrected to Rkymoxis by Persoon in
the Errata corrigenda of his Mycologia Europaea, v. 3, a fact which escaped Raben-
horst who is the only one who used Persoon’s name.

SINGER: THE BOLETINEAE OF FLORIDA 541

Characters of the section evident from the key above and the Latin
diagnosis. Type species P. panuoides (Fr. ex Fr.) Fr., the only species
known in Florida. Another species is P. Curtisii Berk. apud. B. & C.

3. Paxillus panuoides (Fr. ex Fr.) Fr., Epicrisis, p. 318. 1938.

Agaricus panuoides Fr. ex Fr., Syst. Mycol. 1:273. 1821.

Agaricus lamellirugis D. C. ex Secr., Mycogr. Suisse 2: 457. 1833.

Agaricus croceolamellatus Let., Ann. Sci. Nat. sér. II. 3: 94. 1835.

Cantharellus Dutrochetii Turpin apud Mont., Ann. Sci. Nat., sér. II, 5: 342. 1836.

Rhymovis panuoides Rabenh., Kryptogamenfl. 1: 453. 1844.

Paxillus rudis Berk. & Curt., Ann. Mag. Nat. Hist. ser. III. 4: 296. 1859.

Paxillus ligneus Berk. & Curt., Jour. Linn. Soc. 9: 423. 1867.

Crepidotus croceolamellatus Gill, Champ. Fr. Hym., p. 557. 1878.

Tapinia panuoides Karst., Hattsv., p. 452. 1879.

Paxillus acheruntius (Humb. ex) Schroter, Jahresb. Schles. Ges. fiir 1884, p. 300.
1885.

Paxillus lamellirugus Quél., Enchir., p. 93. 1886.

Paxillus ionipus Quél., Assoc. Fr. Avanc. Sci., Congrés d. Toulouse, p. 588. 1887.

Tapinia lamellosa (Sow. ex) Murr., N. Amer. Fl. 10(3): 146. 1917.

Tapinella panuoides Gilbert, Bolets, p. 67. 1931.

?Gomphus pezizoides Pers., Mycol. Europ. 2: 10. 1825.

?Cantharellus olivaceus Schw., Trans. Amer. Phil. Soc. ser. II. 4: 296. 1832 (sec.
Murrill).

Pileus dusky ochraceous to olive ochraceous, “cartridge buff,” ‘“cream
buff,” “chamois,” “honey yellow,” “Isabella color,” “colonial buff,” “deep
colonial buff,” sometimes the basal tomentum purplish-violet, the entire
surface pubescent to more or less tomentose but usually glabrescent,
sometimes rivulose, conchoid, or spathulate-substipitate, dimidiate or with
a short, indistinct, concolorous, lateral stipe, 20-85—(100) mm. — Hymen-
ophore concolorous or, especially when young and fresh, more yellow,
reaching a color between ‘‘Naples yellow” and “mustard yellow,” eventu-
ally more dirty tan, or ‘‘chamois” from the spore dust, easily separable
from the context, lamellae decurrent or radiating from an eccentric point,
rather narrow or narrow, crisp, anastomosing by veins, at least near the
base of the pileus, and sometimes strongly anastomosing-lamellose and
corrugate because of veins which transversely run down the sides of the
lamellae, occasionally some forked without forming anastomoses, rather
close to close; spore print ‘“chamois.’’ — Context cream color to whitish,
soft, rather thin to thin, but sometimes gradually thickening to rather
thick near the point of attachment; taste mild or very slightly bitterish;
odor none, or not remarkable.

Spores 3.8—6 x 3—4.5 p, mostly around 4.8 x 3.5 » in America, 5.8 x 3.5 p
in Europe, melleous-pallid or pale melleous, smooth, thin-walled, ovoid-
ellipsoid; basidia 19-26 x 4.5-6.2 p; cystidia and cheilocystidia none;
trama of the lamellae bilateral, the mediostratum and the lateral stratum
equally dense or the former denser, especially farther away from the edge,
splitting along the mediostratum on increase of pressure; mediostratum
composed of subparallel-subinterwoven hyphae in young stages and near

542 FarLowIA, VoL. 2, 1946

the edges, of more truly interwoven hyphae in age and near the ground of
the lamellae, but always with a distinct axillar orientation; lateral stratum
mainly directed obliquely toward the subhymenium but increasingly inter-
woven, irregular, and even intermixed with age because of the presence of
broad, sometimes clavate hyphae near the subhymenium; cilia of the mar-
ginal tomentum of the pileus formed by 4—5 » thick, long hyphae; hyphae

Chemical reactions: KOH on the surface of the pileus, olive; on lamellae
and context, olive to dull deep yellow; NH;—vapors on the surface of the
pileus, strongly lilaceous-drab, livid; H2SO, on the surface of the pileus,
dull, deep yellow, little reaction elsewhere; FeSO, on context slightly olive.

Habitat: On coniferous wood, on trunks and stumps in the forests as well
as on structural wood in houses, on bridges, in mines, etc. where it is apt
to develop fertile though mostly resupinate fruiting bodies in perfect dark-
ness; preferring pine. Fruiting in summer to late fall if growing in the
open in the northern states, elsewhere also found fruiting in winter and
spring. Usually gregarious, frequently cespitose.

Distribution: Nearly a cosmopolite, but rarer or partly absent in the
tropics. Known from North, Central and South America, Europe, Asia,
Africa, and Australia. In Florida not common.

Material studied: Fla.: Alachua Co., Cary Forest, Nov. 1938, Erdman West &
W. A. Murrill, det. Murrill, F 17708 (FLAS).—N. H.: August 1860, Blake, det. Curtis
740 (FH).—Mass.: Harvard, on coniferous wood, July 1944, Harrie Dadmun &
R. Singer (det. R. Singer) (FH); Murray, Sprague (581) (FH); type of P. ligneus
B. & C. Murray, Sprague (1149) (FH); Floating Bridge, Lynn, August 1913, £. B.
Blackford (FH); Arlington, Oct. 1907, Piguet, det. Farlow (FH).—Conn.: type of
P. hirtus B. & C., Wright 234 (FH).—N. Y.: Enfield Gorge, Sept. 1903, C. H. Kauff-
man (FH).—S. C.: Santee Canal, Ravenel (1038) (FH).—Ga.: NortheastGa.
near Tallulah Falls, Tannery Brook, August 1901, A. B. Seymour, det. Farlow (FH).
—Tenn.: Great Smoky Mts., Greenbrier, July 1939, L. R. Hesler (FH); Grassy
Patch, July 1940, A. J. Sharp, det. L. R. Hesler (FH).— Ala.: “Alabama superioris”
Dec. 1855, Peters (925) (FH).— Ore.: Wygand State Park, Hood River Co. Oct.
1937, W. B. Cooke (9675).—Cal.: Berkeley, February 1933 (FH).— Mexico: Botteri,
det. Curtis (as P. ligneus) (2) (FH).— Brazil: Sao Leopoldo, 1932, Rick (FH).—
Sweden: Upsala, 1853, E. P. Fries (FH).—U.S.S.R. (former Finland): Kellomaki,
Sept. 1940, R. Singer 5-789 (LE).

The European plant usually has slightly larger spores, but this character
is by no means constant, Excellent colored figures of this have been pub-
lished, best of all is that in Lange, Flor. Agar. Dan. 4: pl. 134, fig. B.
Elrod and Blanchard’s photomicrograph (Mycologia 31: 697. fig. 1, D.
1939) of the gill trama gives an idea of the structure of a not quite young
though not very old specimen, but few details are visible. Josserand (Bull.
Soc. Myc. Fr. 48(2):112. pl. 13, fig. 3. 1932) published a better figure,
showing exactly what we have seen and described above.

SINGER: THE BOLETINEAE OF FLORIDA 543

EXTRALIMITAL SPECIES

Paxillus Curtisii Berk. apud Berk. & Curt., Ann. Mag. Nat. Hist. ser. II. 12: 423.
1853.

Paxillus corrugatus Atk., Stud. Amer. Fung., p. 170. 1900.

This species has smaller spores than P. panuoides, as is shown by exam-
ination of the type (on pine wood in Georgia), collected by Ravenel in
September 1848, # 853, preserved at the Farlow Herbarium. In fact
these spores are the smallest known in the Boletineae. Since this species
occurs from New England to Georgia, it is possible that it will be dis-
covered in north Florida.

Sect. Involuti Sing. sect. nov.
Pileo lateraliter vel excentrice vel subcentraliter stipitato; stipite glabro; sporis
volumine mediocri gaudentibus; cystidiis praesentibus.

Characters of the section: see the key p. 539, and the above Latin
diagnosis.

Paxillus involutus (Batsch ex Fr.) Fr. Epicrisis, p. 317. 1838.

This common and widely distributed species is remarkable for its ab-
sence in Florida. It occurs in Uruguay where it has been collected by
Herter, determined as Paxillus argentinus, which strongly differs in having a
veil. The trama of these specimens is divided into an intermixed-inter-
woven hymenopodium, and a subregular mediostratum or rather trama
proper, consisting of brownish, 3.5-9 » broad hyphae, while in the hymeno-
podium the hyphae are hyaline to brownish and still more variable in
diameter; all hyphae with clamp connections; spores 8—11.5—(13) x 5-7.5
p, the wall occasionally up to 1 » thick; basidia 4-spored, about 31 x 7.8-
9.8 »; cystidia hyaline to brownish, versiform, 34-65 x 13-20 »; hyphae
of the cuticular layer of the pileus repent, long-cylindric. For the chemical
reactions of European specimens, see Ann. Myc. 40: 58. 1942.

SPECIES EXCLUDENDAE

Paxillus miniatus Rick, Broteria 5: 19. 1906,

The type of this species does not exist in Brazil; authentic material
kindly sent by J. Rick does not fit in the original diagnosis, and is a
species of the Lentineae with smooth, hyaline spores. We therefore con-
sider P. miniatus as a dubious species, and propose P. russuloides Petch
as the type species of the genus Phyllobolites Sing. P. lateritius is also
thought to belong in Phyllobolites.

Paxillus pubescens Ellis, see under Gomphidius rutilus ssp. alabamensis.

Paxillus pinguis Hook. fil. aud Berk., Hook. Journ. Bot. 3: 41. 1851.

544 Fartowia, Vor. 2, 1946

This is, as is shown by the unpublished colored figure of Hooker (FH),
one of the subspecies of Phylloporus rhodoxanthus, and so are certainly
Paxillus flavidus Berk., Lond. Journ. Bot. 6: 315. 1847, Paxillus rhodoxan-
thus (Schw.) Ricken, Blatterpilze. 1: 95. 1911, Paxillus Tammii Pat.,
Tab. Anal. 4: 161. 1885, Paxillus Pelletieri (Lév.) Velen., Cesk. Houb.,
p. 356. 1920, and Paxillus paradoxus (Kalchbr.) Cooke, Grevillea 5: 6.
1876. Paxillus sulphureus Berk. is most probably another species of Phyl-
loporus.

Paxillus prunulus (Scop. ex Fr.) Quél., Enchir., p. 92. 1886.
See Clitopilus prunulus Jugasporaceae, p. 551 of this paper.

Paxillus reniformis Berk. & Rav. apud Berk. & Curt., Ann. Mag. Nat. Hist. ser. II.
12: 424. 1853.

This species described from North Carolina has been compared with
Paxillus panuoides by Murrill, N. Amer. Fl. 10(3): 146. 1917. However,
the type (FH) has verrucosely punctate spores of 8.3-10.2 x 5.5-6.5 y,
and all the main characters of a Crepidotus. The combination Crepidotus
reniformis (B. & R.) Sing. comb. nov. is proposed. C. versutus Peck is
probably the same species.

Hygrophoropsis R. Maire apud Martin-Sans, L’Empoisonnement, p. 99.
1929,
Cantharellus subgen. Hygrophoropsis Schréter in Cohn, Krypt. FI.
Schles. 3(1): 511. 1889.
Merulius S.F. Gray, Nat. Arr. Brit. Pl. 1: 636. 1821 (non Fries).

Characters of the genus: see key, p. 539. Type species, Hygrophoropsis
aurantiaca (Wulf. ex Fr.) Maire apud Martin-Sans [Cantharellus auran-
tiacus (Wulf.) ex Fr.]

The species entering this genus were formerly considered to belong in
Cantharellus, but since they are fundamentally different in many regards
from the type species of that genus, they were transferred by Studer,
Maire, and others to Clitocybe. The writer formerly agreed with this point
of view though Hygrophoropsis did not show close relationship with any
group of Clitocybe. It appears that the characters in common with Paxillus
are much more striking than the ones that suggested the comparison of
Hygrophoropsis with Clitocybe. The soft context is typical for the Bole-
tineae, and the tramal structure is strongly reminiscent of Paxillus and
Clitopilus. The small, slightly yellowish spores are similar to these of
Paxillus panuoides. The discovery of a second species, H. tapinia, with
smaller spores and an external appearance frankly suggesting Paxillus
Curtisti but never met with in Clitocybe, makes the affinity between Hygro-
phoropsis and Paxillus an established fact.

4. Hygrophoropsis tapinia Sing. spec. nov.

Pileo pallide aurantiaco, admodum excentrico vel astipitato, tomentoso, infundibuli-
formi-irregulari, carnoso atque tenui, usque ad 35 mm. lato; lamellis aurantiacis,

SINGER: THE BOLETINEAE OF FLORIDA 545

immutabilibus, furcatis, haud anastomosantibus nec corrugatis, acie obtusa instructis,
angustis, descendentibus vel arcuate distincte profundeque decurrentibus; sporis in
cumulo albidis, sub microscopio hyalinis vel pallide luteolis, 3.3-4.8 x 2.5-3.3 yw;
basidiis 16-20 x 3.7-6.8 mw; cystidiis nullis; hyphis fibuligeris; stipite pumilo vel nullo,
concolori; carne pallidiore, immutabili, molli, inodora. Habitatio: In truncis cavis,
ad sarmenta in dumetis, etc., Septembri et Octobri mensibus, Florida, U.S.A.

Pileus “pale orange yellow,” very eccentric to astipitate, tomentose, in-
fundibuliform or irregular, fleshy but rather thin, up to 35 mm. broad. —
Hymenophore ‘“capucine orange,” becoming brownish orange, unchanging
on drying and on injury, strongly forking but not anastomosing nor corru-
gate, smooth with blunt edges, narrow, decurrent; spore print white or
whitish on white paper in thin layer. — Stipe small, concolorous, or want-
ing. — Context paler than the surface, unchanging, soft, inodorous.

Spores hyaline to pale yellow under the microscope, 3.34.8 x 2.5-3.3 p;
basidia mostly 4—-spored, 16-20 x 3.7-6.8 yw; sterile bodies in the hymenium
not infrequent (aborted basidia?), extremely variable in shape, usually
numerous on edge; typical cystidia absent; trama consisting of hyaline,
thick (0.74) or thin-walled, often slightly gelatinized hyphae, bilateral
with more parallel-subinterwoven hyphae in the axillarly arranged medio-
stratum (respectively trama proper), and with more irregularly arranged
but mostly straight hyphae in the accompanying layers (lateral stratum
or hymenopodium) ; clamp connections numerous.

Habitat: On dead trunks of frondose trees and on sawdust in dense
hammocks, never under or on conifers. September and October.

Distribution: South Florida.

Material studied: Fla.: Dade Co., Simpson Park, Miami, on sawdust in dense tropi-
cal hammock vegetation, October 1942, R. Singer F 1037 (FH); Brickell’s Ham-
mock, Miami, on a dead trunk in tropical hammock, October 1942, R. Singer, F 1273
(FH). c

This species is very close to H yegrophoropsis aurantiaca from which it
is distinguished mainly by the smaller spores and the habit of fruiting
which suggest Paxillus Curtisii. The description of a British species, Paxz-
lus Fagi, may suggest to some our Florida species, but it is too poorly
characterized and moreover the surface is said to be pallid. Paxillus auran-
tiacus Ellis probably belongs in this group but it reminds one rather of
Hygrophoropsis aurantiaca than of H. tapinia.

5. Hygrophoropsis aurantiaca (Wulfen ex Fr.) R. Maire apud Martin-Sans,
L’Empoisonnement p. 99. 1929.

Cantharellus aurantiacus Fr., Syst. Myc. 1: 318. 1821.

Merulius aurantiacus Persoon ex S. F. Gray, Nat. Arr. Brit. Pl. 1: 636. 1821.
Cantharellus Ravenelii Berk. & Curt., Ann. Mag. Nat. Hist. ser. II. 12: 425. 1853.
Clitocybe aurantiaca Studer, Beiblatt z. Hedwigia 39:(7). 1900.

Chanterel alectorolophoides (Schaeff. ex) Murrill, N. Amer. Flora 9(3): 169. 1910.

Var. typica

546 Fartowia, VoL. 2, 1946

The type variety is a common species in the northern states of this
country, in Canada, in Europe, especially in the northern and central part
of that continent, and in Asia. I have collected typical specimens in all
these regions but not farther south than the Caucasus Mts. in Asia. D. H.
Linder has collected good specimens in Virginia, U.S.A., as did L. R. Hes-
ler in Tennessee, and Ravenel collected the type of Cantharellus Ravenelii
B. & C. in South Carolina. This is definitely the most southerly place
where this species has been found in this hemisphere. The type is in rather
good condition, and appears to belong here, as suggested by Murrill. The
spores of the type variety are 4.8-8 x 2.74.8 #, ellipsoid to cylindric or
rarely reniform, thin-walled, or somewhat thick-walled, hyaline to slightly
yellowish-hyaline; basidia 19-40 x 5-9.5 p; cystidia none, or rare, but
sometimes some aborted basidia present; the trama consists of a medio-
* stratum (trama proper) with distinctly subparallel-interwoven thicker-
walled hyphae which are more loosely arranged than in the hymenopodium ;
the latter strongly irregular, the hyphae running in all directions, many of
them obliquely toward the subhymenium; the hyphae making up the
hymenopodium are denser and slightly thinner and also have somewhat
thinner walls than the hyphae of the mediostratum; the latter is distinctly
continued above the dorsal end of the lamellae just below the context of
the pileus as can be observed in most Boletineae. Paler to partially white
carpophores are not infrequently observed in northern Europe, and have
been described under various varietal names. For a complete description
of this species, see Ann. Myc. 41: 22. 1943, under Clitocybe aurantiaca.
The type variety has not been observed as far south as Florida, We have
observed only a single specimen of what we think is a dwarf form or a
smaller race of this species since it is regularly stipitate and has the spores
of the general measurements of this species. However, it was found on
frondose trunk in low hammock, far from any conifers and this makes it
more remarkable.

Var. nana Sing. var. nov.

Carpophoris minusculis, gracillimis et habitatione in truncis cavis arborum frondo-
sarum in silva palustri subtropicali a typo differt.

Pileus brown, velutinous-tomentose, dry, convex, 6 mm. broad; margin
smooth, initially involute.— ZLamellae reddish Orange, orange, narrow,
forked, subobtuse at the edge, decidedly decurrent, descendant or arcuate,
rather close; spore print not obtained. — Stipe dirty orange brownish, sub-
velutinous, solid, subequal, soft, 9 x 1 mm. — Context subconcolorous with
the surfaces, soft, mild to the taste, inodorous. .

Spores hyaline, 6-8 x 3.5-3.7,, ellipsoid to subcylindric, smooth, non-
amyloid; basidia 20-30 x 5-7.7 uw; cystidia none seen; ¢rama as described
in the type; tomentum of the pileus consisting of loosely interwoven hyphae,
some of them slightly rough (from gelatinous matter), filamentous-cylin-
dric, hyaline, 3-7 » in diameter; all hyphae with clamp connections,

Chemical reactions: KOH, NH,OH, and FeSO,: little reaction.

SINGER: THE BOLETINEAE OF FLORIDA 547

Habitat: In the hollow trunk of living Nyssa sylvatica, in low hammock
vegetation, far from conifers. Solitary in summer.
Distribution: Known only from the type locality.

Material studied: Fla.: Highlands Co., Highlands Hammock State Park, Sept. 1942.
R. Singer, F 640, type (FH).

SPECIES INCERTAE SEDIS

Hygrophoropsis albida (Fr.) R. Maire, Publ. Inst. Botan. 3(4): 58. 1937.
Cantharellus albidus Fr., Syst. Mycol. 1: 319. 1821.

This species has been described from Sweden. I have a few times col-
lected specimens which I used to attribute to it, but since I have not
recently restudied any of these, I hesitate to refer them here, for the
available dried material is not well enough preserved to allow more de-
tailed anatomical studies. Cantharellus albidus (Hygrophoropsis albida)
does not occur in Florida. Maire thinks it belongs in this genus. He lists
it in the family Leptotaceae, but the type genus of that family, Leptotus,
belongs in the Cyphellineae, and has nothing in common with Hygrophorop-
Sis, aS a special study of the Leptotaceae showed (see R. Singer, The
Laschia-complex. Lloydia 8: 188. 1945.)

JUGASPORACEAE (Kiihner ut trib.) Sing. (1936)

Characters of the family: pileus subglabrous to sericeous, smooth or
more rarely venose, not viscid or scarcely viscid, small or rather large,
the margin initially frequently involute, the color as in the whole carpo-
phore usually pallid, white, more rarely light gray; hymenophore lamellose,
lamellae decurrent if there is a stipe; spore print pink; context with more
or less distinct farinaceous odor, more rarely subinodorous; stipe central
eccentric, lateral or absent; veil none; spores hyaline to pale stramineous
under the microscope, with 5-10 longitudinal ridges or veins or blunt
angles formed by flattened stripes running from the hilar end to the apex,
without germinative pore, thin-walled, the wall easily collapsing and the
folds thus formed sometimes resembling additional ridges, young spores
more smooth and rounded, ovoid, ellipsoid, fusoid, asymmetrical; cystidia
none; hyphae of the cuticle and the surface of the stipe (if present) re-
pent; trama consisting of a mediostratum of interwoven to subirregular
hyphae with a generally axillar arrangement, and becoming more parallel-
regular when seen at a point closer to the edge, conspicuously intermixed
near the ground (dorsal part) of the lamellae, looser in the mediostratum,
and the hyphae thicker than in the lateral stratum (hymenopodium) where
some hyphae run obliquely toward the hymenium. The hyphae generally
are irregularly interlaced, and in neither the mediostratum nor the lateral
stratum are they as loosely arranged as in Boletus. The species are homo-
thallic (sec. Kiihner & Vandendries). All hyphae are without clamp con-
nections.

548 FartowlA, VoL. 2, 1946

Habitat: On the soil and on various organic material, wood, dung, dead
Cormophyta, etc.

The Jugasporaceae form a very natural family. They have important
characters in common with the Paxillaceae: the general appearance and
biology, the shape and size of the spores, the structure of the trama (though
the bilateral character of the latter is still less evident in the Jugasporaceae
than it is in the Paxillaceae), the soft context, the occurrence of poroid
hymenophores in aborted carpophores, the absence of cystidia (if Paxillus
involutus is exempted), and the initially involute margin. On the other
hand, it cannot be denied that there is a slight analogy with the Rhodogonio-
sporaceae, in the color of the spores in prints, the odor, and the frequent
occurrence of pigmentless species. It is a fact that the sessile as well as
the smaller stipitate forms of RkodophAvyilus and Clitopilus are hard to
distinguish if one has to rely on macroscopical characters only. It is just
as true that some Rhodophylli with very rounded angles on the spores are
equally hard to distinguish from the Tricholomataceae, such as Rhodocybe
(cf. Lloydia 5(2): 110. 1942). Lange relates Clitopilus prunulus to the
Paxilli by putting it in a neighboring genus Paxillopsis Lange, but this
latter genus is complex, containing all kinds of atypical Paxilli, including
even Rhodopaxillus. Ricken, following Quélet, made Clitopilus a straight
synonym of Paxillus, but his generic conception of Paxillus was still very
broad, evidently under the influence of Quélet and Fries. Lundell, in
Fungi Exsiccati Suecici #137 says: ““Modern authors have placed it in the
genus Paxillus. Lange justly eliminates Clitopilus altogether, and placed
the smooth-spored species with Paxillus, and the angular-spored ones
with Eccilia.” Since Lundell understands Paxillus in a somewhat restricted
sense, probably in approximately the same way as we do, his classification
of Clitopilus prunulus within a restricted genus Paxillus goes much farther
than I would go considering the differences between Clitopilus and Pavxil-
lus, especially in the spore color and clamp connections. Fayod (1889)
comes closest to the point of view expressed in my papers since 1936. He
has the following arrangement: Series D: Tribe 25 Goniosporés (the genera
now considered as Rkodophyllus) ; Tribe 26 Paxillés (the genera here com-
bined in the genus Paxillus and the genus Gomphidius); Tribe 27 Fusi-
sporés (the genera here combined in Clitopilus). Fayod also admits the
affinity of the Paxillés with certain boletes, but so did Opatowski in 1836.

The position of the Jugasporaceae in the classification of the Agaricales
is one of the most difficult problems in spite of the large number of facts
now available. If the similarity with the Rhodogoniosporaceae and Rho-
docybe (in the first case the external similarity, in the latter case the
similarity of the spores *) is considered to show affinity, we arrive at the

* The spores of Rhodocybe alutacea Sing. ined., a species closely related to R. trun-
cata (Fr.) Sing. comb. nov. ssp. subvermicularis (Maire) Sing. (Rhodopaxillus trun-
catus (Fr.) Maire var. subvermicularis Maire from North Africa, are at the same
time warty because of thickenings of the wall, and when seen from above, obtusely
applanate forming a hexagon. This latter character reminds one strongly of Clitopilus,

SINGER: THE BOLETINEAE OF FLORIDA o-549

alternative solutions of this problem: either the Jugasporaceae are at the
same time related with the Rhodogoniosporaceae (Rhodophyllus, Rhodo-
cybe) on one hand, and the Paxillaceae on the other hand, or the Rhodo-
goniosporaceae and Rhodocybe would better be added to the Boletineae in
the larger sense, or the Jugasporaceae could be combined with the family
Rhodogoniosporaceae (Rkodophyllus, Rhodocybe). In view of the strong
arguments in favor of an affinity with the Paxillaceae, and the plurality of
possibilities suggested by the similarities with the Rhodogoniosporaceae
and Rhodocybe, we think it best to leave the Jugasporaceae within the
Boletineae at present, and wait for new facts to be found that might throw
more light on the presumed affinity with other pink-spored agarics. It
may well be that this affinity is not more than a certain coincidental simi-
larity in spore shape.

Since the type concept, now obligatory in nomenclature, excludes the
elimination of Clitopilus, and also its reestablishment in a modified sense
as formerly advocated by R. Heim and myself, I cannot but follow the ex-
ample of Konrad and Maublanc, R. Maire, Josserand, and others who use
the genus Clitopilus in the way it is used inthe present paper. T he ques-
tion of whether the smaller species should be kept in a separate genus
(Octojuga Fay.), or combined with Clitopilus, is, as I have emphasized
on several occasions, a question of the definition of the word genus. As it
seemed before the entrance of certain intermediate species into the taxo-
nomic picture of this family, the larger and the smaller species were well
separated by several correlated characters, i.e. the striation and the size
of the spores, the general appearance, and the habitat. Since then, however,
it has turned out that minute species like Omphalia Giovanellae Bres.
(see Mycologia 34: 66. 1942) and Omphalia scyphoides Fr. belong here,
species with central or eccentric stipe like Clitopilus prunulus, and also
growing on the soil. The difference in the spores does not seem to be so
important now since I have studied many species and specimens, and find
that only the length of the spores is consistently though slightly different
in the two groups. Two Indo-Chinese species, one of them (C. crispus)
very similar to C. prunulus, have strongly 8-10—ribbed spores instead of
6—ribbed ones, and besides the spores are shorter than in C. prunulus. I
therefore do not think that the differences as they appear now are sufficient
for the establishment of two genera within the family Jugasporaceae, and,
like Josserand (Bull. Soc. Myc. Fr. 53: 209-213. 1937), 1 admit. (¢.c.)
only one genus Clitopilus.

The longitudinal striation of the spores of the Jugasporaceae cannot be
identified with the ridges of the spores in Boletellus. In the latter case, the
optical section through the short axis of the spore shows terete spores with

and as a matter of fact, Kiihner has recently stated that in his opinion Clitopilus
prunulus is related to certain species of Rhodopaxillus as he calls the clampless species
which we refer to Rhodocybe in spite of the lack of pseudocystidia, e.g. Rhodocybe
fallax (Quél.) Sing. Syn: Rhodopacxillus fallax (Quél.) R. Maire. See Bull. Soc. Linn.
Lyon, Sept. et Oct. 1945, no. 7-8.

550 FartowlA, VoL. 2, 1946

the veins or lamellose ridges projecting like thorns or wings, while in the
Jugasporaceae we have a regular hexagon (with the angles slightly pro-
tracted or mucronate in Clitopilus prunulus) or heptagon, octagon, etc.
In the latter case, the angles often are so obtuse that the striation of the
spores is extremely difficult to observe, and only the outline of an optical
section with the longer axis vertically directed towards the objective re-
veals the character of the spore. This character may also be obscured by
the fact that young spores are more round in this position, and some do
not seem ever to reach the striate stage. This is the case with Clitopilus
scyphoides, and, to a lesser degree, C. Passeckerianus. But even then the
color of the spore print should easily exclude the genus Omphalia, and the
lack of clamp connections should exclude Clitocybe, Dochmiopus, etc.

Clitopilus (Fr.) Quél., Champ. Jura p. 120. 1872, sensu Patouillard,
Hym., Eur., p. 113. 1887.
Agaricus trib. Mouceron Fr., Syst. Myc. 1: 193. 1821.
Agaricus trib. Clitopilus Fr., Epicrisis, p. 148. 1836, non Syst. Mycol.
Hexajuga Fayod, Ann. Sci. Nat., sér. VII. 9: 389. 1889.
Octojuga Fayod, Ann. Sci. Nat., sér. VIT. 9: 390. 1889.

Characters of the genus: they are the same as for the family. The type
species is Clitopilus prunulus (Scop. ex Fr.) Quél. (Agaricus prunulus Fr.)

KEY TO THE SECTIONS AND SPECIES KNOWN TO THE
AUTHOR

A. Carpophore medium to rather large, pileus 30-130 mm. broad, fleshy, with the
external appearance of a larger Clitocybe, or Paxillus involutus; spores with 6
longitudinal furrows between 6 longitudinal folds or ridges, and therefore three-

striate when seen in profile, (7)-10-14 uw long: growing on the soil.
Sect. Prunuli

6. Clitopilus prunulus
A. Carpophore smaller than indicated above, or spores smaller; spores either ridged
as above, or seemingly smooth but appearing polygonal when seen from one end
(with the longer diameter vertically directed toward the objective), 5.5-11.5 4
long, ridges or angles variable in number, usually more than 6, often up to 10;
growing on the soil (mostly sand), or on various debris, rotting or rotten wood,
manure, etc.
B. Stipe present, constantly persistent. Sect. Scyphoides
C. On horse manure in mushroom cellars; stipe very small, less than 3 mm.
long. Clitopilus Passeckerianus
C. Not on manure, only in the open; stipe usually larger.
D. Pileus gray. 7. Clitopilus Giovanellae
D. Pileus white.
E. Spores with scarcely projecting ridges or folds, merely angular
when seen from one end. 8. Clitopilus scyphoides
E. Spores with strongly (0.3-0.4 m) projecting longitudinal ridges or
folds.
F. Pileus fleshy with crenate margin. Clitopilus crispus

F. Pileus thin with smooth, entire margin.
Clitopilus orcelloides

SINGER: THE BOLETINEAE OF FLORIDA iat

B. Stipe absent, or spurious, or disappearing with age, or inconstant.
Sect. Pleurotelloides

G. On horse manure, in white-mushroom cellars. (Rare astipitate form of
Clitopilus Passeckerianus)
G. On substrata other than manure.

H. Pileus rugose-venose; growing on the wood of evergreen trees in
tropical hammock. 9, Clitopilus venososulcatus

H. Pileus smooth, or transparently striate. 10. Clitopilus pleurotelloides

Sect. Prunuli (Quél. ut sectio subgeneris Orcellae generis Paxil, Ench., p. 92, 1886)
em. :
Carpophoris mediocribus vel majusculis, carnosis, distincte stipitatis; sporis 6 rugis
longitudinalibus instructis, 10-14 u longis. Ad terram.

Characters of the section: they are evident from the key and the Latin
diagnosis above. Type species as in the genus Clitopilus.

6. Clitopilus prunulus (Scop. ex Fr.) Quél., Champ. Jura, p. 120. 1872.

Agaricus prunulus Scop. ex Fr., Syst. Mycol. 1: 193. 1821.

Agaricus orcellus Bull. ex Fr., Syst. Mycol. 1: 180. 1821.

Clitopilus orcellus Quél., Champ. Jura, p. 120. 1872.

Paxillus prunulus Quél., Enchir., p. 92. 1886.

Paxillus prunulus var. orcella Quél., Enchir., p. 92. 1886.

Hexajuga prunulus Fayod, Ann. Sci. Nat., sér. VII. 9: 289. 1889.

Rhodosporus prunulus Schroter, in Cohn, Krypt.-Fl. Schles. 3(1): 618. 1889.
Pleuropus prunulus Murr. N. Amer. FI. 10(2): 104. 1917.

Pleuropus obesus Murr., N. Amer. Fl. 10(2):; 105. 1917.

Paxillopsis prunulus Lange, Flora Agar. Dan. 4: 48. 1939.

Pileus whitish, whitish-gray, whitish-cream, sometimes with spots, dots,
or zones, sometimes scarcely subviscid during rains, but usually quite dry,
mostly finely velutinous or innately mealy, convex but very soon flatter and
depressed though usually very irregular, with involute, eventually irregu-
larly-lobed margin, and obtuse, rarely indistinctly umbonate disc, smooth,
30-130 mm. broad. — Hymenophore white or whitish, then pale incarnate,
easily separable from the context of the pileus, lamellae frequently not
straight, moderately close to close, rather narrow or narrow (up to 4 mm.
broad), deeply decurrent; spore print “fawn color.” — Stipe white or con-
colorous with the pileus, slightly fibrillosely striate, glabrous or subglabrous,
often with tomentose base, naked, versiform, solid, but softer inside, 26—50
x 6-21 mm. — Context white, soft, with the consistency of a bolete; taste
mild, farinaceous; odor strong, farinaceous (of fresh flour, or flowers of
Berberis vulgaris).

Spores elongate-ovoid to mostly subfusoid, with distinct suprahilar de-
pression, with (5)—6-(7) longitudinal brims or edges which usually are
more or less projecting, forming low ridges or folds with furrows running
between them, the angular outline well-marked in an optical section with
the long diameter of the spore vertically directed towards the objective
(use chloral hydrate as medium), hyaline to hyaline-stramineous, asym-

552 Fartowia, VoL. 2, 1946

metric, (7)—10-14 x 5-6 », most frequently 10.5-12 x 5—5.8 pu; basidia 4-
spored, 30-36 x 8.3-9.8 »; cystidia and cheilocystidia none; trama of the
lamellae intermixed-subbilateral, the mediostratum differing from the lateral
stratum respectively the hymenopodium ® in consisting of large, irregu-
larly arranged elements in the basal (dorsal) portion of the lamellae, and
generally in being less densely interlaced, and much thicker; toward the
edge of the lamellae these differences become less distinct, the medio-
stratum becoming more regular and its elements more elongate and narrow,
and even the hymenopodium becoming slightly less interlaced, its hyphae
showing a slight but indistinct tendency to diverge toward the hymenium,
5.5—6 in diameter; hyphae of the cuticular layer of the pileus repent,
cylindric-filamentous; all Ayphae consistently without clamp connections.

Chemical reactions: NH,OH, yellowish-brown-vitreous. — FeSO, no
reaction. — Guaiacol reacting only in the base of the stipe, and even there
weakly and slowly. — Methylparamidophenol, negative to weak, or
moderate reaction in stipe, no reaction or only a weak one on the pileus,
the edges of the lamellae usually becoming distinctly deep umbrinous, in
young specimens the surface of the pileus and the edges of the lamellae
becoming distinctly deep violet but the rest of the carpophore remaining
negative.

Material studied: Fla.: Alachua Co. Gainesville, some collections by W. A. Murrill
(Pleuropus obesus) (FLAS) ; by R. Singer F 1867, F 1867a (FH). —N. H.: Chocorua,
August 1904, Farlow (FH).— Mass.: Canton, August 1925, D. H. Linder (C. orcellus)
(FH) ; Wakefield, August 1943, R. Singer (FH).—N. Y.: Newcomb Co., Mt. Goode-
now, September 1941, R. Singer (FH).—Ia.: Iowa City, September 1936, G. W. Mar-
tin (FH).—Tenn.: Highlands, August 1936, L. R. Hesler & A. J. Sharp (FH). —
Canada: N. B., October 1903, det. Farlow (FH). — U.S.S.R.: Moscow region, Mi-
khailowskoe, June 1896, Buchholz (FH); Leningrad region, near Ryabovo, August
1938, R. Singer (Hexajuga prunulus) (LE).— Spain: Catalonia, Salardi, June to
October 1934, R. Singer (BA).— Numerous collections of the fresh fungus in Czecho-
slovakia, Austria, Bavaria, and France, and abundant dried material from all Euro-
pean countries in the European herbaria. — Asiatic part of the U.S.S.R., Caucasus,
Abkhazia, Saken, August 1928, R. Singer; Caucasus, Guzeripl, August 1935, L. N. Vas-
silieva (KAZ).

This is an easily identified species, and one of the less-known but ex-
cellent edible mushrooms. Clitopilus orcellus is only an insignificant form
of this species. Its occurrence in Gainesville, south of the 30th parallel,
probably marks the southernmost limit of the species.

° What I call the hymenopodium, has been described by others as the subhymenium,
e.g. by R. Maire for Clitopilus cretaceus Mre. (See also p. 538 and 547 of this paper.)

SINGER: THE BOLETINEAE OF FLORIDA 553

Sect. Scyphoides Sing. sect. nov.

Carpophoris vix mediocribus, aut sporis minoribus quam 10 yw et tunc carpophoris
nonnunquam mediocribus vel submajusculis, constanter distincte stipitatis.

Characters of the section: compare the key p. 550, and the above Latin
diagnosis. The type species is Clitopilus scyphoides (Fr.) Sing., more
exactly var. typicus fa. typicus, as defined below, p. 554.

7. Clitopilus Giovanellae (Bres.) Sing., Mycologia 34: 66. 1942.
Omphalia Giovanellae Bres., Fung. Trid. 1: 9, pl. 5, fig. 2. 1881.

Pileus gray, sometimes somewhat shining, sericeous, smooth, but often
subrugulose on the margin, and also slightly transparently striate at the
extreme margin when wet and quite mature, not viscid, convex with a
deep umbilicus in the center from the beginning, eventually becoming
flatter to infundibuliform, 6-8 mm. broad (5-14 mm. according to Bresa-
dola). — Hymenophore pale gray, easy to separate from the context of
the pileus, lamellae very thin and slender, deep decurrent, arcuate, sub-
close to close, narrow; spore print pinkish. — Stipe grayish-white to gray,
somewhat silky, central, or sometimes somewhat eccentric, subequal, very
thinly and finely velutinous toward the base in many specimens, but in
our Florida material never seen with white tomentum at the base, as de-
scribed by Bresadola, solid or stuffed, 5-12 x 1-1.5 mm. — Context pallid,
very brittle when dried, tender when fresh; odor very slightly farinaceous,
almost inodorous in most cases; taste distinctly farinaceous but not bitter.

Spores 6-7 x 3.5—4 yw, smooth, with eight or less longitudinal flattenings
bordered by obtuse or almost rounded edges, not or hardly visible even in
chloral hydrate but the outline of an optical section clearly octagonal (or
hepta- or hexagonal) when the longer diameter of the spore is pointing
up toward the objective (like an egg seen “from above’), fusoid to ovoid,
hyaline to stramineous-hyaline, at least the larger spores with a supra-
hilar depression, non-amyloid; basidia 16-25 x 6.8-8.3 yw; cystidia and
cheilocystidia none; gill trama partly intermixed; all Ayphae without clamp
connections.

Habitat: Widely scattered on sandy places, in May and June (June and
July in Italy).

Distribution: With certainty known only from Italy and north Florida,
probably widely distributed but easily overlooked since it is rare and small,

-and often occurs in places where commonly no fungi are expected.

Material studied: Fla.: Alachua Co., Gainesville, June 1943, R. Singer, F 2294
(FH); May 1943, W. A. Murvill, det. R. Singer, F 2072 (FH).—lItaly: authentic,
from Gocciadoro, province of Trento, July 1899, G. Bresadola (NY).

Among the smaller species of Clitopilus, sect. Scyphoides, this species
is well-determined by its color and the more or less central stipe, its occur-
rence on sunny, sandy places in spring. There is a minor difference between
our Florida material and the Italian plant, i.e. the lack of the white tomen-

554 FaRLowlA, Vou. 2, 1946

tum on the base of the stipe of the former. This is an interesting parallelism
to what we have observed in the following species, C. scyphoides. In neither
case should too much importance be attributed to this frequently variable
character. Otherwise, our specimens agree perfectly with the authentic
material cited above. C. Giovanellae is new for this continent.

8. Clitopilus scyphoides (Fr.) Sing. comb. nov. (sensu lato).

Agaricus scyphoides Fr., Syst. Mycol. 1: 163. 1821 (equals var. typicus f. typicus).

Agaricus mutilus Fr., Syst. Mycol. 1: 191. 1821 (equals var. typicus {. mutilus).

Omphalia scyphoides Quél., Champ. Jura, p. 99. 1872.

Pleurotus mutilus Gill., Champ. Fr. Hymen., p. 344. 1878.

Omphalina scyphoides Quél., Enchir., p. 42. 1886.

Clitopilus submicropus Rick, Broteria, 6: 77. 1907 (equals var. submicropus).

Omphalina floridana Murr., Mycologia 33: 440. 1941 (equals var. floridanus).

Pleuropus minimus Murr., Lloydia 5: 144. 1942 (equals var. floridanus).

Clitopilus omphaliformis Joss., Bull. Mens. Soc. Linn. Lyon 10. 1941 (fide
Locquin, zbid. 13: 107. 1944).

Var. typicus forma typicus

The European type of this species does not occur in Florida, but it is
here redescribed for comparison with the American varieties. Pileus white,
convex and umbilicate or infundibuliform from the beginning, not hygro-
phanous or subhygrophanous but with a dry, sericeous, very thin and
sometimes indistinct sericeous covering, eventually undulate but with more
or less regularly circular outline, 5-25 mm. broad. Hymenophore white
to creamy white, eventually pallid to pinkish in fertile specimens, separable
from the context of the pileus, lamellae narrow (up to 1.5 mm. broad),
arcuate or horizontal-linear, or descendant, always decidedly and mostly
deeply decurrent, subclose to crowded. Stipe white, slightly sericellous,
typically short, 4-10 x 0.5-1.8 mm. (up to 2 mm. at the typically white
tomentose, frequently thickened base), equal or subequal, flexuous or
straight (typically more frequently flexuous), solid or stuffed, becoming
hollow in age. Context white, thin; odor farinaceous or none; taste fari-
naceous and mild. Spores ellipsoid-oblong, or in smaller spores with a
rhomboid outline, typically narrower than in var. floridanus, 7.5-11.5 x
4—5.5 m, otherwise as in the variety; basidia mostly 4-spored, 17-28 x
6.5—-8.5 mw; cystidia and cheilocystidia none; trama as in var. floridanus;
cuticle of appressed, long, cylindric, smooth, clampless, 2.5-3.5 p» thick
hyphae, making up the sericellous covering; neither the spores nor the
hyphae are amyloid.

Habitat: Solitary or in small to large groups in moist or mossy meadows
or fields, but also on the soil or on very decayed stumps in the woods,
May to October, through most of Europe but I have never verified the
indications from Siberia and South Africa. I have, however, made fresh
collections in Bavaria, near Munich, where it is frequent in May and
June on meadows, and I have examined the material in Lundell & Nann-
feldt, Fungi Exsiccati Suecici (LE) as well as two specimens from the

SINGER: THE BOLETINEAE OF FLORIDA 555

Hohnel Herbarium (FH), one from Jenbach, V. Héhnel B.2145, one from
near Trento, F. v. Héhnel.

Var. typicus f. mutilus (Fr. /.c.) Sing. comb. nov.
Clitopilus omphaliformis {. calathinoides Locquin, Bull. Mens. Soc. Linn. Lyon
18: 107. 1944.

This form does not occur in Florida, but it is here redescribed for com-
parison with other forms and varieties, and because it has so often been
misinterpreted. Pileus white, subhygrophanous, very slightly transparently
striatulate when wet, not viscid, very minutely subfarinaceous-subsericel-
lous, not circular but elongate in outline when old, often becoming lobate,
tongue-shaped, or labiate-depressed, very eccentric to lateral, and in the
latter case assuming the general appearance of Arrhenia auriscalpium (but
larger), with initially incurved margin, 10-14 mm. broad. Hymenophore
white, eventually somewhat pink, lamellae crisp, with rather obtuse edge
when young, forked or simple, rather narrow, subrounded-attenuate or
arcuate to the stipe, descendant-decurrent, close. Stipe white, solid, short
white tomentose at the base, erect and straight, occasionally flexuous, very
minutely mealy-sericellous, subequal, 5-17 x 1-1.5 mm. Context white,
fragile, inodorous. Spores 6.8-8.5 x 3.5-5 mw as in the var. floridanus ;
basidia up to 26.5 x 5-8y, 4-spored; cystidia and cheilocystidia none;
trama intermixed, its bilaterality has not been checked by the author;
hyphae without clamp-connections.

Habitat: On mossy north border of a mixed (Pinus, Picea, Betula) wood,
growing among mosses (or actually on decayed needles scattered among
these? ).

Distribution: Certainly in Scandinavia, France, Finland, and also in the
Leningrad region of the U.S.S.R., near Mga, August 21, 1938, R. Singer
& L. N. Vassilieva L37, 5-596 (LE). The above description was. gathered
from this latter collection. This form probably occurs in most of Europe.

Var. floridanus (Murr. /.c.) Sing.
PLATE I, Fic. 8-11.

Pileus white, pure white when dry, becoming sordid-pallid when old
and wet, innately fibrillose-sericellous, smooth, or scarcely noticeably stri-
atulate when wet, sometimes somewhat uneven, with the margin initially
incurved, convex and umbilicate from the beginning, the umbilicus very
deep, eventually the pileus deeply depressed, 8-20 mm. broad. — Hymeno-
phore white or creamy white, eventually becoming pinkish in the fertile
carpophores, easily separable from the context of the pileus, lamellae arcu-
ate, decurrent to deeply decurrent, narrow (about 1 mm. broad), subclose
to close, simple; spore print pale and dirty pink. — Stipe white, whitish,
or sordid-pallid when old, glabrous or somewhat innately white-fibrillose,
smooth, even, the base hardly noticeably mycelioid, versiform but usually
as long as the pileus is broad, or longer, rarely shorter, and subequal, solid

556 FarLow1A, VoL. 2, 1946

or stuffed, sometimes becoming hollow, sometimes curved, 5-20 x 1-2.5
mm.— Context white, thin, very brittle when dried; odor farinaceous, or
sometimes none; taste farinaceous, and slightly bitterish to mild.

Spores often agglutinate in fours, or in pairs, 6-10 x 3.7—5 pz, ellipsoid
to ventricose-subfusoid, smooth, hexa- to octagonal because of (6)—8 flat-
tened longitudinal ribs limited by obtuse or somewhat rounded edges which
are not or hardly visible even in chloral hydrate, but become evident in an
optical section with the long diameter of the spore pointing upward to the
objective (like an egg seen “from above’), the wall thin and very easily
collapsing, especially along the longitudinal edges, and then often appear-
ing longitudinally ridged, some with a slight suprahilar depression, some
only with an applanation, non-amyloid, hyaline to stramineous-hyaline;
basidia 17-26.5 x (5)—6.7-9.5 1, 4—spored, sometimes some 2- to 3—spored ;
cystidia and cheilocystidia none, but in some specimens many basidia abor-
tive; gill-trama intermixed- subbilateral, non-amyloid, the hyphae without
clamp connections; cuticle of the pileus (the thin silky covering) consisting
of long, cylindric, appressed, smooth, more or less interwoven, clampless
hyphae of 2—5.3 » diameter, forming a loose to subdense tissue.

Chemical reactions: FeSO,, somewhat grayish, almost negative. —
Methylparamidophenol, negative.

Habitat: On open lawns, on bare ground and among grasses, also on
roadsides and near the hammocks, never in the hammocks, fruiting from
May to October. Gregarious.

Distribution: Common all over Florida.

Material studied; Fla.: Dade Co., Miami SW, 15th Road, October 1942, R. Singer
F1216/II (FH); Highlands Co., Highlands Hammock State Park, August 1942,
R. Singer F 465 (FH); Alachua Co., Gainesville, several collections, among them
type of Omphalina floridana, W. A. Murrill F 16223 (FLAS) ; and type of Pleuropus
minimus Murr., W. A. Murrill, F 19980 (FLAS, FH).

Var. submicropus (Rick l.c.) Sing. comb. nov.

Pileus as in var. floridanus, but more distinctly sericeous or sericeous-
fibrillose, smooth, convex with depressed center, larger, 20-30 mm. in
diameter. — Hymenophore as in var. floridanus, close to crowded, some-
times somewhat forked; spore print pale and dirty pink. — Stipe white or
pallid, innately fibrillose to somewhat mealy, especially on the apex, smooth,
the bases connected by a very thin mycelioid membrane, solid, equal, but
frequently with slightly thickened base and apex, sometimes flexuous,
10-25 x 2-4 mm. — Context white, very brittle when dried and also more
or less fragile when fresh, rather thin; odor farinaceous; taste slightly
bitter to mild.

Spores often agglutinate in pairs or in four, 5.5-6.8 x 3.5—4 p, shaped
as in var. floridanus; basidia 17 x 6.8 »; cystidia and cheilocystidia none
seen; hyphae without clamp connections.

Habitat; In the hammocks (tropical hammock formations in Florida),
on small debris and sticks, gregarious to cespitose, fruiting in Florida in
October.

SINGER: THE BOLETINEAE OF FLORIDA 557

Distribution: Tropical Florida and Brazil, possibly all over tropical
America.

Material studied: Fla.: Dade Co., Matheson Hammock, October 1942, R. Singer
F 1252 (FH).— Brazil: type, Rio Grande do Sul, 1906, J. Rick (FH).

Rick indicates the spores as being 6 x 2.5 », but in the type specimens I
have found many foreign spores as well as many collapsed spores of the
Clitopilus, and some of these fit in Rick’s measurements. However, the
non-collapsed spores of Rick’s specimen perfectly agree with the above
measurements taken from my Florida collection.

Var. aff. floridanus, ined.

Pileus as in the type variety but less distinctly sericellous. Hymeno-
phore as in the type variety but often sterile. Stipe as in the type variety
but less silky, more elongate in general and more frequently straight,
sooner becoming hollow in large specimens, 3—-17—(24) x 0.7-2 mm., the
base glabrous in some specimens, white tomentose in others (inconstant).
Context white, not very fragile, but very brittle when dried; odor fari-
naceous only when the carpophores are squeezed. Spores 7.7-9 x 4.5-5.2 p
(specimen from North Dakota); dasidia 17.5-27.5 x 6.5-8.5 p, 2-, 3-,
or 4—spored (very variable); cystidia and cheilocystidia none; trama and
cuticle as in var. floridanus.

Habitat: On well-watered lawns on the soil, also in swamps on decayed
leaves and wood, and on burned-over prairie sod, fruiting from June to
August, gregarious.

This is another form, close to var. floridanus (Murr.) Sing., but obvi-
ously not entirely identical with it, found in the northern and western
states.

Material studied: Vt.: Middlebury, July 1897, E. 4. Burt (FH).— Mass.: Woburn,
August 1943, W. L. White, det. Singer (sterile form) (FH).—N.D.: Kulm, June
1923, J. F. Brenkle (under a new unpublished name), N. Dakota Fungi 1613.

Sect. Pleurotelloides Sing. sect. nov.

A sectione praecedente stipite inconstanter formato vel haud persistente aut rudi-
mentario vel complete absente differt.

Characters of the section: these are evident from the key p. 551, and
the above Latin diagnosis. Type species: Clitopilus pleurotelloides (Kiihn.)
Sing.

9. Clitopilus venososulcatus Sing. spec. nov.

Pileo pallide albo, radiato-sulcato et venoso, farinaceo, sessili vel subsessili, 12 23
mm. lato; lamellis concoloribus, satis angustis, postice attenuatis, confertis; sporis in
cumulo Poreole 8-8.5 x 4.5-5 w, substramineis, striatulis; stipite nullo vel abana seater
usque ad 1.3x0.6 mm.; carne inodora. Habitatio: Ad lignum emortuum prope Miami
Floridae, U. S. A.

Pileus pallid white, radially venose and sulcate, mealy, not viscid, sessile
or subsessile, 12-23 mm. broad. — Hymenophore white or pallid, becoming

558 FarLowlA, VoL. 2, 1946

dotted with pink from the spores, lamellae rather narrow (1.5-2 mm.
broad), close, attenuate behind; spore print pinkish on white paper. —
Stipe absent or rudimentary, and then concolorous and minutely mealy, not
larger than 1.3 x 0.6 mm.— Context white, inodorous, very thin, very
fragile.

Spores hyaline to pallid-stramineous, ellipsoid or short ventricose-sub-
fusoid, or somewhat ovoid, longitudinally substriate because of flattened
ribs limited by obtuse edges, in optical cross section obscurely angular,
hexagonal or octagonal, 8—8.5 x 4.5—-5 yw; basidia 20-21 x 7-7.5 wa; cystidia
and cheilocystidia none seen; all hyphae without clamp connections.

Habitat: On dead portions of standing trunks of living Ficus aurea, and
also on logs from the same host. October.

Distribution: Coastal Hammock in south Florida.

Material studied: Fla.: Dade Co., Matheson Hammock, October 29, 1942, R. Singer
F 1344, type (FH).

This species is close to C. pinsitus (Fr.) Josserand and C. pleurotelloides
(Kihn.) Sing. It differs from both in the venose surface, and from the
latter species also in being larger.

10. Clitopilus pleurotelloides (Kiihn.) Sing. comb. nov.

Octojuga pleurotelloides Kiihner, Le Botaniste 17: 158. 1926.

Octojuga variabilis (Fr. sensu Karst. p. p., non al.) Fayod, Ann. Sci. Nat., sér.
VIL. 9: 390. 1889.

Clitopilus variabilis Josserand, Bull. Soc. Myc. Fr. 53: 212. 1937.

Octojuga Fayodi Konr. & Maubl., Icon. Sel. Fung. 6: 234. 1924-36.

Geopetalum viticola Murr., Bull. Torr. Bot. Club 67: 2. 1940.

Pleurotus viticolus Coker, Journ. E. Mitch. Sci. Soc. 60: 92. 1944.

Pileus pallid white, frequently becoming somewhat dirty grayish cream
in age, opaque, white tomentose in the zone nearest to the substratum,
smooth or transparently striatulate when moist, at least in the marginal
zone when quite mature, glabrous but sericeous or finely villous when
seen under a lens, circular, semicircular, or reniform, convex, sometimes
attached by a resupinate portion and the free margin horizontally deflexed,
the margin initially incurved, then repand and straight and occasionally
radiately splitting, largest diameter of the non-resupinate portion of the
pileus 1.5-10 mm. — Hymenophore pallid, lamellae moderately broad to
very broad, subclose to rather distant, inserted, with entire or slightly un-
even edges; spore print pale cinnamon pink. — Stipe absent, or present
in very young specimens and then up to 1 mm. long, but soon becoming
rudimentary and button-like, or entirely disappearing, concolorous, sub-
pruinose to sericeous. — Context concolorous, very fragile, brittle when
dried; odor none or very slight, farinaceous; taste mild.

Spores 5.5-9 x 3.5-5.8 p (according to Kiihner up to 12 x 6.2y),
stramineous-hyaline, non-amyloid, ellipsoid to ovoid in outline but slightly
hexagonal, octagonal, or decagonal with flattened to very slightly concave
sides when seen in optical cross section, from one end; basidia 25-29 x

SINGER: THE BOLETINEAE OF FLORIDA 559

7.5-8.5 uw, 4-spored or a few 2-spored; cystidia and cheilocystidia none;
trama non-amyloid; hyphae of the sericeous covering of the pileus cylin-
dric, subinterwoven with some semi-erect ends, 2-4 yp thick, all hyphae
without clamp connections.

Habitat: On dead cortex of tree trunks, on logs and dead branches,
vines, (Quercus, Eucalyptus, Salix, according to R. Maire, Cornus florida
and Vitis spec. according to Coker), and also on fallen needles of conifers,
dead thallus of mosses (Hypnum according to Kiihner), on bark of Zizy-
phus jujuba, also on herbaceous stems; June to December.

Distribution: Europe, Asia, and America, possibly cosmopolitan but
often overlooked or misdetermined.

Material studied: Fla.: type of Geopetalum. viticola, Alachua Co., Arredonda, 1938,
Erdman West & W. A. Murrill (FLAS).— USSR: (formerly Finland), Kellomaki,
Sept. 1940, R. Singer 5-836 (LE).—India: Ladhar, Sheikhupura, Sultan Ahmed,
August 1944 (det. Singer), 1097 (FH).

Fayod based his name on an observation by Karsten, Mycol. Fenn. 3:
112. 1876: “Occurrit forma sporis minoribus (longit. 6 mm., crassit.
3-4 mm.), subhyalinis, qvalem in ramulis Salicis Capraeae prope Abo
mense Aug. legimus.” Abo is now Turku, Finland, and it may be of in-
terest that I collected this species in Finland (or what formerly was
Finland) on debris of rotten moss among sand near the railway station
of Kellomaki, September 1940. There is, therefore, little doubt that
Fayod was right in attributing this variety to what we now call Clitopilus
pleurotelloides. But the type variety of Agaricus variabilis sensu Karsten
is, as I have been able to confirm on the basis of Karsten’s exsiccatum
of this species distributed by him under #511 of the Fungi Fennici, de-
termined correctly in the sense of the majority of the authors, z.e. it is
a typical Dochmiopus (Agaricineae) with rough spores. Kihner was,
therefore, justified in erecting a new name for the Clitopilus reserving the
epithet “variabilis” for the Dochmiopus. The American collections do not
differ sufficiently to separate them from C. pleurotelloides. They are, per-
haps, slightly thinner with the lamellae accordingly broader and the sur-
face of the pileus accordingly more striatulate, but striations are frequent
in European specimens also, and the size of the lamellae and the diameter
of the flesh vary considerably. The spores are slightly smaller in the types
than they are in the European specimens I have studied, but Coker, who
has re-collected Murrill’s form in North Carolina, gives spore measure-
‘ments which are exactly intermediate to almost identical with my Euro-
pean measurements (6.7—9 x 4.8-5.8 for the Kellom&ki collection, 5.5—-8 x
3.5—4.8 » for Murrill’s type). This again shows that the size of the spores
is variable in this country as well as in Europe, though not more so than
in some other species of this same genus. Murrill observes that “not
one individual was found entirely resupinate, nor was there a stipe in the
entire lot.” If this latter observation should hold true for all American
collections, including the youngest stages, it would be possible to distin-

560 FarLowI!A, Vor. 2, 1946

guish Murrill’s species as a form or variety of C. pleurotelloides. I doubt
very much that the astipitate character of the American form is constant
since it is not constant in the European plant.

EXTRALIMITAL SPECIES

Clitopilus crispus Pat., Bull. Soc. Myc. Fr. 29: 214. 1913.

The type has been described from Indo-China. These specimens and
an excellent colored picture are preserved at the Farlow Herbarium. There
is also a spore print on blue paper which when transferred on white paper
now is between “fawn color” and “wood brown.” The mature spores are
stramineous under the microscope, or, in dense piles, succineous, ellipsoid
with some tendency to become hexagonal, with seven to ten (mostly
eight) angles reinforced by longitudinal ribs and with furrows between
them when seen in an optical cross section from one end, the ribs pro-
jecting about 0.4 p, 6.8-8.8 x 4.3-5.2 . This species is by far the stout-
est of the small-spored Clitopili, but, nevertheless, belongs in the section
Scyphoides. .

Clitopilus orcelloides Pat. & Demange, Bull. Soc. Myc. Fr. 26: 40. 1910.

The type, preserved in the Patouillard Herbarium (FH), comes from
Indo-China. It has spores exactly as described by Patouillard, i.e. ellip-
soid-subhexagonal or subrhomboid, or equally ovoid-ellipsoid, octagonal
when seen from above, and then the angles reinforced by longitudinal
veins which in the optical cross section look like knots, 5.5—7.5 x 3.7-5.3 p,
sometimes as short as 6 x 5.2 p, the ribs projecting up to 0.3 4; hymenium
and trama not very distinct in these specimens. There is an excellent
colored picture in the unpublished “album” of V. Demange, executed
by a native artist, and representing the type specimens in fresh condi-
tion. These vividly recall Clitopilus scyphoides var. floridanus from which
the dried specimens are difficult to distinguish. However, the more angular
spores, many of which are ornamented with ribs, cause me to consider
C. orcelloides a different though related species of the section Scyphoides.

Clitopilus Passeckerianus (Pilat) Sing., Mycologia 34: 66. 1942.
Pleurotus Passeckerianus Pilat, Atl. Champ. Eur., Pleurotus Fr., Ser. A, 9-10:
49. 1935.
Pleurotellus Passeckerianus Konr. & Maubl., Icon. Sel. Fung. 6: 361. 1938.
Octojuga Passeckeriana Sing., Ann. Mycol. 40: 61. 1942.

This is a frequent weed-fungus in white-mushroom beds, and in my
opinion is slightly different from C. pinsitus as well as from C. scyphoides
var. typicus {. mutilus and C. pleurotelloides. For more descriptive data
see the original account of Pilat, (1935) with photographs, and also
Singer, (1942).

SINGER: THE BOLETINEAE OF FLORIDA 561

SPECIES IMPERFECTLY KNOWN

Clitopilus pinsitus (Fr.) Josserand, Bull. Soc. Myc. Fr. 53: 210. 1937.
Agaricus pinsitus Fr., Syst. Mycol. 1: 184. 1821.

This species, originally described from Sweden, has been redescribed by
Josserand from France. Though it is impossible to prove that Josserand’s
interpretation is the only correct one, it seems to me that it is the best
emendation of an otherwise superfluous Friesian species. I have not col-
lected it myself, nor has it been found in America. It differs from C.
pleurotelloides in being much larger and in having close, narrow lamellae.

- SPECIES DUBIAE

Clitopilus cretatus (Berk. & Br.) Sacc., Syll. 5: 702. 1887.

Described from England, this species is thought by Lange to be the
same as Pleurotus mutilus, i.e. Clitopilus scyphoides var. typicus fa. muti-
lus. However, R. Maire (Publ. Inst. Botan. 3(4): 83. 1937) refers it to
his genus Clitopilopsis because of Massee’s indication on the spores of the
type specimen of Agaricus cretatus Berk. & Br., Ann. Mag. Nat. Hist.
ser. III. 7: 373. 1861. In Massee’s paper (Grevillea 21: 81. 1893) are
some not quite trustworthy measurements, and it may be unwise to take
his word for it that these spores are not of the Clitopilus-type. A re-
examination of the types at Kew will decide whether C. cretatus is another
synonym of C. scyphoides, or a good species in some other genus.

| SPECIES EXCLUDENDAE

Clitopilus abortivus (Berk. & Curt.) Sacc., Syll. 5: 701. 1887.

This species, common in north Florida in winter, does not belong in
this genus, but must be called Rhodophyllus abortivus (B. & C.) Sing.,
Rev. d. Mycologie 5: 9. 1940. Other species formerly called Clitopilus,
but having angular spores of the Rkodophyllus-type, must also be placed
in the genus Rhodophyllus rather than Clitopilus. These are C. undatus
Fr. (ut Agaricus), C. cancrinus Fr. (item), C. albogriseus Peck (item)
(Rhodophyillus albogriseus (Peck) Sing., recte), and many others.

Clitopilus pallidus Heim, Treb. Mus. Ciénc. Nat, Barcelona 15(3): 109. 1934.

Since Clitopilus is now used in the sense of Patouillard and Maire, the
species of Clitopilus em. Heim, l.c. must go into Clitopilopsis Maire
(Agaricineae). The type species of Clitopilopsis is C. pallida (Heim)
Maire from Spain. Another species to be excluded is Clitopilus togoensis
Henn., also considered to belong in Clitopilus em. Heim by Heim (1934)
and Singer (1936), and certainly belonging in the Agaricineae rather than
in the Jugasporaceae.

562 Fartowia, VoL. 2, 1946

GENERA EXCLUDENDA

The following genera formerly considered to be allied to the boletes
or other groups of the Boletineae, are now excluded from this suborder.

Filoboletus Henn. (belongs to the Tricholomataceae, Agaricineae).
Polyporoletus Snell (belongs to the Scutigeraceae, Clavariineae).
Fistulina Bull. ex. Fr. (belongs to the Fistulinaceae, Cyphellineae).
Volvoboletus Henn. (is a parasitized or abnormal Amanita).

GENERA DUBIA

Fistulinella Henn.
Melanomphalia Lange.
Gymnogomphus Fayod.

Harvarp UNIVERSITY
CamprIDGE, Mass.

THE MOST IMPORTANT LITERATURE ON THE BOLETINEAE,
ESPECIALLY THOSE OF FLORIDA

_ Atkinson, G. F. Studies on American Fungi, Mushrooms. p. I-VI, 1-275. Ithaca.
1900.

Barbier, M. Les Bolets. p. 1-30. Besangon. 1908.

Beardslee, H. C. New and interesting fungi. Mycologia 26: 253-260. 1934.

Beck, G. v. Mannagetta, Versuch einer systematischen Gliederung der Gattung
Boletus L. em. Zeitschr. Pilzk. 2: 141-149. 1923.

Beeli, M. Contribution nouvelle 4 l’étude de la flore mycologique du Congo. Bull.
Soc. R. Bot. Belg. 58: 203-216, pl. 15-16. 1926.

Bresadola, G. Iconographia Mycologica. 19: pl. 901-944. Mediolani. 1931.

Coker, W. C. & A. H. Beers. The Boletaceae of North Carolina. p. I-VIII, 1-96,
pl. 1-65, and front. Chapel Hill. 1943.

Elrod, R. P. & D. L. Blanchard. Histological studies of the Boletaceae and related
genera. Mycologia 31: 693-708. 1939.

Farlow, W. G. (edited by E. A. Burt) Icones Farlowianae. p. I-X, 1-120, pl. 1-103.
Cambridge, Mass. 1929.

Fayod, V. Prodrome d’une histoire naturelle des Agaricinées. Ann. Sci. Nat. Bot.
VII. 9: 181-411. 1889.

Fries, E. M. Systema Mycologicum. 1: 385-395. 1821.

Fries, E. M. & C. T. Hok, Boleti Fungorum Generis Ilustratio. 14 pp. 1835.

Frost, C. C. Catalogue of Boleti of New England, with description of new species.
Bull. Buff. Soc. Nat. Sci. 2: 100-105. 1874.

Gilbert, E. Les Bolets, p. 1-255. Paris. 1931.

Gray, S. F. A Natural Arrangement of British Plants, 1: 1-824. 1821.

Heim, R. Notes sur la flore mycologique Malgache III. Rev. d. Mycol. 1: 3-18.
1936; VII. Rev. d. Mycol. 4: 5-20. 1939.

Josserand, M. Sur la nature de la trame dans les genres Paxillus et Phylloporus.
Bull. Soc. Myc. Fr. 48: 112-117, pl. 13. 1932.

. Notes critiques sur quelques champignons de la region Lyonnaise.

Bull. Soc. Myc. Fr. 53: 175-230, pl. 6. 1937.

rn

SINGER: THE BOLETINEAE OF FLORIDA 563

Kallenbach, F. Die Rohrlinge in Pilze Mitteleuropas 1 (1-19): 1-138, pl. 1-52.
1926-38.
Kauffman, C. H. The genus Gomphidius in the United States. Mycologia 17: 114-
126, pl. 12-14. 1925.
Kavina, K. Ceske Slizaky. Trav. Mycol. Tchécoslov. 2: 1-8, 1 pl. 1924.
Karsten, P. A. Enumeratio Boletinearum et Polyporearum Fennicarum systemate
novo dispositarum. Rev. Mycol. 3: 16, 1881.
Killermann, S. Pilze aus Bayern. Denkschr. Bayer. Bot. Ges. Reg. 16: 1-23. 1925.
Konrad, P. Un dernier mot sur la classification des Bolets. Bull. Mens. Soc. Linn.
Lyon 3: 1-4. 1934.
Konrad, P. & A. Maublanc. Icones Selectae Fungorum. 4-6. 1924-1937.
Lohwag, H. & M. Peringer. Zur Anatomie der Boletaceae. Ann. Mycol. 35: 295-331.
1937.
Murrill, W. A. The Boleti of the Frost Herbarium, Bull. Torrey Bot. Cl. 35: 517-
526. 1908.
. The Boletaceae of North America. Mycologia 1: 4-18; 140-160. 1909.
. Agaricales. (Boletaceae) N. Amer. Fl. 9(3): 133-161. 1910.
. New Boletes. Mycologia 30: 520-525. 1938.
. Three new Boletes. Mycologia 31: 110-112. 1939.
. Addition to Florida fungi. Bull. Torrey Bot. Cl. 67: 57-66. 1940.
. New fungi from Florida, Lloydia 5: 136-157. 1942.
. Florida Boletes (Boletaceae). Mim. Contrib. Herb. Univ. Fla. Agric.
Expt. Sta. p. 1-6. 1942.
. More new fungi from Florida. Lloydia 6: 207-228. 1943.
—————. More fungi from Florida. Lloydia 7: 303-327. 1944.
Opatowski, G. Commentatio Historico-naturalis de Familia Fungorum Boletoideorum.
Diss. inaug. Berlin 1836 (also in Wiegmann’s Archiv. 2: 1-34. 1836).
Patouillard, N. & C. F. Baker. Some Singapore Boletinae. Jour. Straits Branch
R. A. Soc. 78: 67-72. 1918.
Peck, C. H. Genus Boletus. Rep. N. Y. State Cab. 23: 127-133. 1872.
. The Boleti of the United States. Bull. N. Y. State Mus. 2(8): 73-166.
1889, =
Persoon, C. H. Mycologia Europaea 2: 123-148. 1825.
Pilat, A. in Kavina, K. & A. Pilat. Atlas des Champignons de l'Europe: Pleurotus.
Praha, 1935.
Quélet, L. Enchiridion Fungorum in Europa et praesertim in Gallia vigentium, p.
I-VI, 1-352. 1886.
————.. Flore Mycologique de France. p. I-XVII, 1-492. Paris. 1888.
Rostkovius, F. W. T. in Sturm, J. Deutschlands Flora III 5: 1-132, pl. 1-48. 1844.
Saccardo, P. A. Sylloge Fungorum. 5, 6, 9, 11, 14, 16, 17, 19, 20, 21, 23. Patavii.
1887-1925.
Secretan, L. Mycologie Suisse. 3: I-VIII, 1-760. Genéve. 1833.
Singer, R. Sur les genres Ixocomus, Boletinus, Phylloporus, Gyrodon et Gomphidius.
Rev. de Mycol. 3: 35-53, 157-177. 1938.
. Notes sur quelques Basidiomycétes. VI. Rev. de Mycol. 5: 3-13. 1940.
. Das System der Agaricales I. Ann. Mycol. 34: 286-378. 1936; II. 40:
1-132. 1942.
. New genera of fungi. Mycologia 36: 358-368. 1944.
. The Boletineae of Florida. I. Farlowia 2: 97-141. 1945. II. Farlowia
2: 223-303. 1945. III. Amer. Midl. Nat. (in press).
Slipp, A. W. & W. H. Snell. Taxonomic-ecologic studies of the Boletaceae in northern
Idaho and adjacent Washington. Lloydia 7: 1-66, pl. 1-8. 1944.
Smotlacha, F. Monografie ceskych hub hribovitych (Boletinei). Vestn. €esk. spol.
nauk 2(8): 1-73, 1911.
Snell, W. H. Notes on Boletes I. Mycologia 24: 334-341. 1932; II. Mycologia 25:

564 Fartowia, VoL. 2, 1946

221-232. 1933; III. Mycologia 26: 348-359, pl. 41. 1934; IV. Mycologia 28:
-13-23. 1936. V. Mycologia 28: 463-475. 1936.
Tentative keys to the Boletaceae of the United States and Canada.
Publ. No. I. Rhode Island Bot. Cl. 1-25. 1936.
. The genera of Boletaceae. Mycologia 33: 415-423. 1941.
. New proposals relating to the genera of the Boletaceae. Mycologia 34:
403-411. 1942.
——— & E. A. Dick, Notes on boletes VI. Mycologia 33: 23-37. 1941.
Yates, H. S. A comparative histology of certain Californian Boletaceae. Univ. Calif.
Publ. Bot. 6(10):; 221-247, pl. 21-25. 1916.

* ' p
hi t
7 ‘ Lae 3 I
+ i ‘ ’ caer certs
' Ps i .
ae . ‘ ees, tne AR rer r
: i c eet : * = a
ei 4 , i he ; ais : re ’ es nl hes
; ale es rs mi ; ‘
; G F . = :
x > ‘ .

i

le

566 FartowI1A, VoL. 2, 1946

EXPLANATION OF PLATE I

All figures are drawn at a magnification of 2000.

Fig. 1-3. Gomphidius rutilus (Fr.) Lund. & Nannf. ssp. alabamensis (Earle) Sing.
1. Cystidium.
2. Basidium.
3. Two spores, (left) frontal view; (right) in profile.
Fig. 4-7. Gomphidius vinicolor Peck ssp. jamaicensis (Murr.) Sing.
4. Two spores, (left) nearly-frontal view; (right) in profile.
5. Hyphae of the remainders of the veil on the apex of the stipe, running in
parallel strands, with strong pigment-incrustation.
6. Basidium.
7. Cystidium with the partially thickened wall.

Fig. 8-11. Clitopilus scyphoides (Fr.) Sing. var. floridanus (Murr.) Sing.

8. Two spores, (above) in frontal view; (below) in profile with the hilar end
above; the line A... B indicates the longitudinal axis, C ...D the short
axis.

9. Three spores, seen from above (the longitudinal axis A... B_ vertically
directed toward the lens), the short axis indicated corresponding to fig. 8,

‘lower spore, as C...D. The upper spore shows 6 flattened sides and 6
angles, central one 7, the lowest 8 (which is the normal number in this form).

10. Basidium.

11. Hyphae of the sericeous covering of the pileus.

SINGER: THE BOLETINEAE OF FLORIDA

SINGER PrLaTeE I

567

2(4): 569-580 FARLOWIA July, 1946

INDEX TO VOLUME 2

INDEX TO AUTHORS AND TITLES

Abbott, Isabella A. The genus Griffithsia (Rhodophyceae) in Hawaii, 489-453.

Bartram, Edwin B. New mosses from Tierra del Fuego, 309-319.

Chardon, Carlos E. Fungi Domingenses novi vel minus cogniti. I, 455-473.

Cutter, Victor M., Jr. The genus Cunninghamella (Mucorales) , 321-343.

Harris, Hilda F. The correspondence of William G. Farlow during his student days
at Strasbourg, 9-37.

Linder, David H. In honor of William Gilson Farlow, 1-7.

Patrick, Ruth. A taxonomic and ecological study of some diatoms from the Pocono
plateau and adjacent regions, 143-221.

Prince, Alton E. The biology of Gymnosporangium nidus-avis Thaxter, 475-525.

Roberts, Catherine. A comparative study of Torulopsis pulcherrima and Taphrina
deformans in culture, 345-383.

Singer, Rolf. The Boletineae of Florida with notes on extralimital species. I. The
-Strobilomycetaceae, 97-141; The Boletineae of Florida with notes on extralimital
species. II. The Boletaceae (Gyroporoideae), 223-303; The Boletineae of Florida
with notes on extralimital species. IV. The lamellate families (Gomphidiaceae,
Paxillaceae, and Jugasporaceae, 527-567; Notes on Farlow’s Agaricales from
Chocorua, 39-51.

Taylor, Wm. Randolph. William Gilson Farlow, promoter of phycological research
in America 1844-1919, 53-70.

Verdoorn, Frans. Farlow’s interest in an international abstracting journal, 71-82.

Weston, William H., Jr. Dr. Farlow’s influence on mycology, 84-95.

Whelden, Roy M. Note on Argentine Charas, 305-308.

Whetzel, H. H. The cypericolous and juncicolous species of Sclerotinia, 385-437.

569 »

570 FartowlA, Vo. 2, 1946

INDEX TO GENERA AND SPECIES

New names and the final members of new combinations, and the pages on which
they occur, are in bold-faced type; in taxonomic papers, where the main treatment
of a genus or species is given, the page number only is in boldface. Synonyms and
the pages on which they occur are printed in Ztalics.

Achnanthes 166-170, 213; subgen. Ach-
nanthes 166, 169; subgen. Achnanthi-
dium 166; subgen. Microneis 166;
coarctata 166; exigua 167; hungarica
167; lanceolata 166, 167, 169, 211, 212;
lanceolata var. apiculata 167; lanceolata
var. Oestrupii 169; lanceolata var. ro-
busta 168; lanceolata var. rostrata 168;
Lewisiana 166, 168; linearis 167, 168,
207; microcephala 167, 169; minutis-
sima 167, 169; minutissima var. crypto-
cephala 169; Oestrupii 166; Oestrupii
var. parvula 169; Peragallii 166, 169;
Stewartii 169-170; Suchlandtii 168

Achnanthidium subgen. 166; coarctatum
166; hungaricum 167 ; lanceolatum 167;
lineare 168; microcephalum 169

Actinocephalum 323, 325; japonicum 323,
334

Aecidium nidus-avis 481 -

Agaricineae 527, 562

Agaricus trib. Clitopilus 550; trib. Mou-
ceron 550; cretatus 561; croceolamel-
latus 541; involutus 540; lamellirugis
541; mutilus 554; orcellus 551; panu-
oides 541; paradoxus 280; Pelletieri
280; pinsitus 561; prunulus 550, 551;
rhodoxanthus 280, 283; rutilus 533;
scyphoides 554; Tammii 280, 282; vari-
abilis 559

Allospori sect. 107, 138

Amanita 562; rubescens 272; sp. 250

Amblystegiaceae 316-318

Amblystegium excurrens 317

Amphipleura 170; pellucida 170

Amphora 195-196, 213; atomus 184;
ovalis 195; ovalis var. pediculus 195;
perpusilla 196

Ananae sect. 107, 122

Angustiporini subsect. 259, 260, 267

Anomoeoneis 213, 214

Anthoceros 62

Aporpiellus subgen. 251, 252

Arrhenia auriscalpium 555

Ascobolus magnificus 456

Ascophanus granulatus 456

Asterionella 153; formosa 153; Ralfsii
153, 207, 209, 212

Atrotomentosi sect. 538, 540

Bacillares sect. 176, 181

Bacillaria phoenocenteron 176; ulna 154;
viridis 195

Bartramiaceae 315

Batrachospermum 68

Biblarium leptostauron 151

Blakesleea 324

Boletaceae 101, 223-303, 527, 538

Boletellus 98, 103, 104, 105, 107, 122-139,
258, 549; sect. Allospori 107, 138; sect.
Ananae 107, 122, 139; sect. Chrysen-
teroidei 107, 130, 139; sect. Dictyo-
podes 107, 136; sect. Ixocephali 107,
135; sect. Mirabiles 107, 128; sect.
Retispori 107, 130; ananas 97, 104,
122, 123-126, 127, 128, 131, 133, 137,
139; annamiticus 126; betula 104, 137,
138; chrysenteroides 118, 128, 130, 131,
133, 134; costatus 138; cubensis 123,
127, 128; emodensis 122, 126, 127, 128,
283; guadelupensis 123; jalapensis 131,
135; lignatilis 123; lgulatus 139; Lin-
deri 104, 130, 131, 134; mirabilis 129,
300; obscure-coccineus 123, 127; pal-
lescens 123; paradoxus 138; picti-
formis 104, 130, 131, 134; pictiformis
var. fallax 131, 132, 133; porphyrius
122, 126, 127, 128, 131; projectellus
129, 130; pustulatus 139; retisporus
104, 130; Russellii 98, 136, 137; singa-
porensis 104, 135; subflavidus 120;
turbinatus 130, 134

Boletineae 97-141, 223-303, 527-567

Boletinellus 243; intermedius 246; meruli-
oides 243, 247; purpureus 249

Boletinus 98, 100, 225, 226, 229, 250;
251-257, 258; subgen. Aporpiellus 251,
252; subgen. Eu-Boletinus 251, 252;
sect. Cavipedes 251, 252, 253; sect.
Palustres 251, 252; sect. Solidipedes 251,
253, 257; sect. Spectabiles 251, 252;
amabilis 251, 253, 256, 257; appendicu-
latus 257; asiaticus 252; Benoisii 253,
256; Berkeleyi 254; castanellus 257,
296; cavipes 251, 252, 255; decipiens
253, 254, 255, 296; flavoluteus 257,
260, 278; floridanus 254; glandulosus
257, 278; grisellus 253, 256; Lakei
253, 256, 257; merulioides 257; ochra-

INDEX TO VOLUME 2

ceoroseus 253, 256, 257; oxydabilis
253, 256; paluster 252; pictus 251, 252,
253, 255, 256; porosus 245, 247, 249,
257; porosus var. opacus 247; puncta-
tipes 257; solidipes 257; spectabilis 251,
252; squarrosoides 257, 295; subgrisel-
lus 257; subtomentosus 296

Boletochaete 225, 300

Boletogaster 103, 122, 138; jalapensis 135;
Russellii 136

Boletoideae subfam. 227

Boletopsis corrugata 258; singaporensis
135

Boletus 224, 225, 228, 230, 240, 243, 257,
258, 285, 286, 287, 298, 299, 300, 547;
subgen. Porphyrosporus 115; sect. Fa-
vosi 106; acidus 271; aeruginascens
259; alabamensis 293; albidipes 44, 45,
272; alboater 105; albus 42, 271; ama-
bilis 256; amarellus 46; ananaeceps 123;
ananus 103, 122, 123, 126; appendicula-
tus 250; badius 299; Bellinii 42; betula
103, 138; bicolor 300; brachyporus 243,
249; brasiliensis 298; Braunii 241;
brevipes 41, 44, 265, 267; caespitosus
249; castaneus 237; castaneus fa. pur-
purinus 236; cavipes 251; chrysenter-
oides 103, 133; chrysenteron 103, 128,
134; cinereus 110; circinans ssp. lepto-
pus 42; coccineus 123, 126; communis
134; costatisporus 115; costatus 138;
crassus 242; cubensis 127; cyanescens
235; cyanescens 6 fulvidus 237; deci-
piens 100, 254; Earlei 239; edulis 250,
279, 290, 294; emodensis 126; ferru-
gineus 297; flavidus 260; flavus 260;
floccopus 110, 113; fraternus 300; Fros-
tii 258; fulvidus 237, 299; fumosipes
116, 118, 119, 133, 240; gossypinus 110;
gracilis 103, 121; granulatus 40, 41, 44,
267, 271, 276; granulatus var. albidipes
272; Grevillei 259; griseus 99, 300; gua-
delupensis 128, 131, 133; gyrosus 249;
hieroglyphicus 296; hirtellus 273, 274,
276, 277; hirtellus var. mutans 277;
hirtellus var. siccipes 274, 277; hydri-
ensis 110; illudens 287, 292, 293, 295;
impolitus 300; innixus 299; intermedius
246; isabellinus 123, 126; jalapensis
103; Junghuhnii 297; lacteus 235; la-
cunosus 122, 139; Lakei 257; lanatus
296; lateralis 247; Leguei 296; lepiota
110; lignatilis 127; lignicola 299; lividus
243, 249; lividus ssp. alneti 249; lividus
ssp. labyrinthicus 249; lividus var. ru-
bescens 249; luridellus 100; luteus 258,

571

264, 270; Morganii 138; Murraii 256;
nebulosus 115, 116; nigrellus 105; ob-
Scure-coccineus 127; pallescens 126;
pictus 256; Pierrhuguesii 46; piperatus
46; placidus 42, 271; porphyrosporus
106, 115; porphyrosporus var. minor
116; projectellus 103, 129; pseudo-
granulatus 44, 265, 268; pseudoscaber
115; pulverulentus 250, 300; punctipes
277; pustulatus 139; retisporus 130;
Rostrupii 139; Roxanae 299; rubellus
99, 127, 250; rubellus ssp. caribaeus
100; rubescens 249; rubinellus 45-47;
rubricitrinus 99; rubrotubifer 46; ru-
focastaneus 237, 239; Russellii 103, 136;
scaber 99; sistotrema 243, 249; sordidus
103, 118, 119; spadiceus 297; sphaero-
Sporus 243; Spraguei 256; squamatus
127, 128; squarrosus 110; strobilaceus
106, 110; strobiloides 110; stygius 110;
subluteus 269; subtomentosus 128, 296;
subtomentosus ssp. spadiceus 297; sub-
tomentosus ssp. spadiceus var. lanatus
297; subtomentosus var. Leguei 296;
sudanicus 241; sulphureus 300; testa-
ceus 237; tomentosus 277, 296; triden-
tinus 260; tristis 115, 118; tropicus 241,
242; tumidus 300; umbrosus 115; Van-
derbiltianus 46; variegatus 236; versi-
color 127; viscosus 265, 267

Bornetia Binderiana 442

Bovini sect. 259, 278

Brachytheciaceae 318

Brachythecium arenarium 318; collinum
318; longidecurrens 318; micro-col-
linum 318; rivulare 318

Brasiliensis sect. 287, 298

Breutelia angustiretis 315; chrysura 315;
integrifolia 315

Brevistriatae subgen. 187, 188

Bryaceae 312-315

Bryum andicola 315; austro-chilense
313; brachycarpum 314; caespiticum
314; dicarpum 313; fuegianum 313;
Hatcheri 314; Lechleri 315; macro-
sporum 313; obscurum 313; pauper-
culum 314; pseudothyridium 314;
Roivainenii 312; vernicosum 314;
zeballosicum 314

Calliergidium austro-stramineum 317

Calliergon sarmentosum 318

Calliergonella complanata 319

Caloneis 172, 194; Lewisii 172; Schu-
manniana 172; silicula var. alpina 172;
trinodis 172

572

Calyculosphaeria calyculus 466

Camptochaete sect. Rigodiella 316; ar-
busculans 316

Candida 375; pulcherrima 348, 372; tropi-
calis 348, 372

Candidaceae 375

Cantharellus 544; subgen. Hygrophorop-
sis 544; albidus 547; aurantiacus 544,
545; Dutrochetii 541; olivaceus 5341;
Ravenelii 545, 546

Capitatae subgen. 187, 189

Catacauma Sabal 461

Catagonium 318 :

Cavipedes sect. 251, 252, 253

Centroceras clavulatum 440

Ceramiaceae 439-453

Ceramiales 442

Ceriomyces alabamensis 293, 295; betula
138; flavimarginatus 293; fumosipes
116; Housei 296; illudens 293; jala-
pensis 135; mirabilis 128, 129; pro-
jectellus 129; pseudoboletinus 290, 292;
Russellii 136; sordidus 119

Chalciporus 47; piperatus 46

Champia sp. 442

Chanterel alectorolophoides 545

Chara 305; Curtisii 306; delicatula 306;
foetida 305, 306; hydropitys 306;
leiopitys 306; leptopitys 306; Martiana
305, 306; vulgaris 208

Choanephora 324, 336; cucurbitarum 336

Chorisodontium lanigerum 309; sericeum
309

Chroogomphus subgen. $27, 528, 529

Chrysenteroidei sect. 107, 130, 139

Ciboria Armeriae 436; Juncigena 425,
432, 433

Cladopodiella fluitans 209

Claudopus 50, 51; byssisedus 51

Clavariineae 562

Claviceps caricina 397, 401; nigricans 394,
396

Clitocybe 544, 550; aurantiaca 545, 546;
Pelletieri 280

Clitopilopsis 561; pallida 561

Clitopilus 50, 98, 544, 548, 549, 550-561;
sect. Pleurotelloides 551, 557; sect.
Prunuli 550, 551; sect. Scyphoides
550, 553, 560; abortivus 561; albo-
griseus 561; cancrinus 561; cretaceus
552; cretatus 561; crispus 549, 550,
560; Giovanellae 99, 550, 553, 554;
omphaliformis 554; omphaliformis fa.
calathinoides 555; orcelloides 550, 560;
orcellus 551, 552; pallidus 561; Pas-
seckerianus 550, 551, 560; pinsitus 558,

FarLowlA, VOL. 2,

1946

560, 561; pleurotelloides 551, 557,

558, 559, 560, 561; prunulus 544, 548,

550, 551; seyphoides 550, 554,
561; scyphoides var. floridanus 99,
554, 555, 556, 560; scyphoides var.
submicropus 556; scyphoides var. ty-
picus fa. mutilus 555, 560, 561; scy-
phoides var. typicus fa. typicus 553,
554; submicropus 554; togoensis 561;
undatus 561; variabilis 558; venoso-
sulcatus 551, 557

Clitopilus trib. 550

Cocconeis 165-166; aspera 197; lineata
165; placentula 165, 207, 208, 209;
placentula var. lineata 165, 211, 212

Cocconema cistula 197; tumidum 198

Coelopus 231

Colletonema vulgaris 171

Complexae subgen. 187, 194

Compsopogon 61

Conferva flocculosa 149; pectinalis 160

Cookeina sulcipes 455, 456, 457; tricho-
loma 455, 456, 457

Cordyceps militaris 458

Coriolus 527

Cortinaria rutila 533

Crepidotus 538, 544; croceolamellatus
541: reniformis 544; versutus 544

Cricunopus 257; luteus 257, arg

Criella leucothoés 458

Cryptococcus hominis 347, 348, 4375

Cunninghamella 321-343; africana 323,
324, 326, 327; albida 323, 334; Bainieri
323, 324, 326, 328, 334, 335, 336; Ber-
tholletiae 323, 324, 326, 329, 330, 331
332, 333, 334; Blakesleeana 323, 324,
326, 329, 330, 334; dalmatica 323, 335,
336; echinata 323, 329, 330; echinulata
323, 324, 325, $26, 327, 328, 329, 330,
332, 334; elegans 323, 324, 326, 330,
331, 332, 333, 334, 335; mandshurica
336; microspora 323, 336; polymorpha
323, 335, 336; ramosa 323, 324, 335;
verticillata 323, 327, 329, 330; sp. “A”
329

Cyclotella 147-148; comta var. radiosa
147, 207; Meneghiniana var. rectangu-
lata 147, 148; Meneghiniana var. stelli-
gera 148; stelligera 147, 148

Cylindrosporium Securidacae 469

Cymbella 196-199, 213; amphicephala
196, 197; aspera 196, 197, 208; cistula
196, 197; cuspidata 196, 197; Ehren-
bergii 196, 197; gracilis 209; Hauckii
196, 197; microcephala 196, 197; navi-
culiformis 196, 198; pediculus 195;

INDEX TO VOLUME 2

scotica 196, 198, 208, 209; sinuata 196,
198; tumida 196, 198; turgida 196, 198;
ventricosa 196, 198, 208, 211
Cyphellineae 547, 562
Cystogomphus 528, 529; Humblotii 529
Cystopleura 202, 214; turgida 202; Zebra
var. proboscidea 202; Zebra var. sax-
onica 202
Cystopus 91

Daedalea merulioides 247

Debaryomyces 375; hominis 348

Decipientes sect. 176, 187

Decussatae sect. 176, 177

Delesseria 24; sinuosa 64

Dendroligotrichum squamosum 319

Denticula Tabellaria 203

Desmogonium 164

Diadesmis biceps 182

Diatoma fenestratum 148

Dicranaceae 309

Dictyochorella Andropogonis 468

Dictyopodes sect. 107, 136

Dictyosphaeria 440

Dictyota sp. 442, 443

Didymodon ampliretis 310; diaphano-
basis 310

Didymosphaeria 467; Trichostigmae 467

Diploneis 174, 213; ovalis 174; Smithii
174

Distantes subgen. 187, 188

Divergentes subgen. 187, 192

Dochmiopus 550, 559

Dodgea 255

Dothideales 459-465

Dothidella domingensis 459; tinctoria
459

Drepanocladus 318; exannulatus 208

Eccilia 50, 51, 548

Echinella circulare 149

Entoleiae 176, 182

Entoloma 50, 51; rhodopolium 51

*Epidocium affine 394, 396, 397, 399, 401,
423; ambiens 391, 394, 395, 396, 401;
ambiens fa. Caricis-paniculatae 394;
arabicus 395

Epithemia 202; proboscidea 202; saxonica
202

Erinella calospora 456, 457; longispora
456, 457

Eriocorys 108; strobilacea 106, 110; stro-
bilacea var. floccopus 110

Eu-Boletinus subgen. 251, 252

Eucatagonium politum 319

Eunotia 155-165, 210, 213; alpina 160;

573

amphioxys 203; arcus var. curta 156,
157; arcus var. tenella 164; bidens 162;
diodon 157; elegans 155, 157; exigua
156, 157; faba. 157, 158; fallax 156,
158; flexuosa 155, 158, 163, 210; for-
mica 155, 158; gracilis 156, 158; im-
pressa 161; incisa 164; indica 156, 158;
lunaris 156, 159, 210, 211; lunaris var.
capitata 156, 159; Meisteri 156, 159,
207, 212; microcephala 157, 159, 208,
212; monodon 156, 160; monodon var.
major 155, 160, 208, 210; Naegelii 156,
160, 208, 209, 212; pectinalis 156, 160,
207, 210, 211; pectinalis var. minor 160,
207; pectinalis var. minor fa. im-
pressa 161; pectinalis var. recta 161;
pectinalis var. undulata 161, 210, 211;
praerupta 156, 162; praerupta var.
bidens 157, 162; praerupta var. curta
162; praerupta var. inflata 162; ro-
_ busta 157, 162, 207, 208, 212; ros-
tellata 156, 163; septena 157, 163; sep-
tentrionalis 156, 163, 210; sudetica 156,
163; suecica 157, 163; tautonensis 155,
163, 207, 213; tenella 156, 164, 210;
tridentula 159; tridentula var. permi-
nuta 159; trinacria 156, 164; valida
156, 164; veneris 157, 164, 165, 208,
209. 21d 79

Euryporini subsect. 259, 278

Euryporus 251; cavipes 251

Exilaria Vaucheriae 152

Falkenbergia rufalanosa 440

Favolus purpureus 249

Favosi sect. 106

Filoboletus 562

Fimetariales 465

Fistulina 562

Fistulinaceae 562 —

Fistulinella 562

Flammula rhodoxanthus 280;
280.

Flammulina velutipes 264

Floccigomphus sect. 528, 529

Fragilaria 150-153; bicapitata 150, 210,
212; brevistriata 150, 151; capucina
var. mesolepta 150, 151; constricta
152; construens 150, 151; crotonensis
150, 151, 211; elliptica 150, 151, 210;
grandis 161; leptostauron 150, 151;
mesodon 148; pectinalis var. undulata
161; pinnata 150, 151; undata 152;
undata var. lobata 150, 151; undata
var. y 152; Vaucheriae 150, 152; vires-

Tammii

574

cens 150, 152, 207, 210, 211,
virescens var. capitata 152

Frostiella 103, 122, 137; betula 138; Rus-
sellit 136

Frustulia 170; bipunctata 192; cuspidata
177; lanceolata 180; maior 194; ovalis
195; pellucida 170; rhomboides var.
saxonica 171; rhomboides var. saxonica
fa. capitata 171

Fucus 62, 91; evanescens 68

Funaria chiloensis 312; pungens 311

Funariaceae 311-312

Fungi Imperfecti 468-469

212}

Gaillonella italica 147

Ganoderma 102, 121

Gastroboletus 225, 226

Genuini sect. 106, 108

Geopetalum viticola 558, 559

Gigartina compressa 24

Gomphidiaceae 102, 527-537

Gomphidius 98, 100, 528, 529-537, 548;
subgen. Chroogomphus. 527, 528, 529;
subgen. Laricogomphus 528, 536; nib
gen. Myxogomphus 528, 536; sect.
Floccigomphus 528, 529; sect. Macro-
‘sporus 528, 536; Microsporus 528, 537;
sect. Viscogomphus 528, 529; alaba-
mensis 533, 535, 536; alachuanus 531,
533, 536; foltporus 280, 282, 537;
furcatus 533, 536; glutinosus 529, 530,
532, 536; jamaicensis 531, 533, 535;
leptocystis 529; litigiosus 533; macu-
latus 529, 530, 536; nigricans 536;
ochraceus 531, 533, 534, 536; orego-
nensis 537; rhodoxanthus 280, 537;
roseus 530, 537; rutilus 529, 530, 531,
533, 536; rutilus ssp. alabamensis
531, 535, 543; rutilus ssp. typicus 533,
534; rutilus var. fulmineus 534; ruti-
lus var. litigiosus 534; rutilus var.
pulcher 534; rutilus var. testaceus 534;
septentrionalis 536; sibiricus 529;
Smithii 536; subroseus 537; superior-
ensis 533, 536; testaceus 533; tomen-
tosus 527, 529, 533; vinicolor 529, 530,
533, 536; vinicolor ssp. jamaicensis
531; viscidus 531, 533, 536; viscidus
var. fulmineus 534; viscidus var. testa-
ceus 534; viscidus fa. columbiana 533,
534, 536; viscidus fa. testacea 534; sp.
536

Gomphonema 199-202, 213; acuminatum
199; acuminatum var. coronatum 199;
acuminatum var. laticeps 200; acum-
inatum var. pusillum 200; angustatum

FartowliA, VoL. 2, 1946

var. obtusatum 199, 200; augur 199,
200, 208; auritum 201; capitatum
200; constrictum 199, 200; constrictum
var. capitatum 200; coronatum 199;
gracile 199, 200; gracile var. auritum
201; gracile var. lanceolata 201; gra-
cile var. naviculoides 201, 211; laticeps
200; micropus 202; naviculoides 201;
parvulum 199, 201, 207, 211, 212; par-
vulum var. micropus 202, 211; sphaero-
phorum 199, 202; turris 199, 202

Gomphus pezizoides 541

Gonatobotrys microspora 336

Gracilaria 24

Graciles sect. 107, 119

Granulati sect. 258, 259, 260

Griffithsia 439-453; Binderiana 439, 440,
442, 443; corallina 440, 442, 443; coralli-
noides 440; globulifera 441; Metcalfii
439, 440, 441, 442, 443; ovalis 439,
440; rhizophora 439, 440, 441, 443,
444; subcylindrica 443; tenuis 439,
440, 441, 442, 443; thyrsigera 439, 441;
Sp. 441

Grimmia apocarpa 311;

Grimmiaceae 310-311

Grunowiae sect. 203

Guepinia lutea 79; merulina 79

Gymnocybe Tammii 280

Gymnogomphus 531, 562

Gymnosporangium clavariaeforme 480;
clavipes 476, 481, 500, 512, 513; conz-
cum 475, 476, 481; effusum 481, 498;
globosum 476, 483, 512; Juniperi 475;
juniperinum 475; Juniperi-virginianae
512; juvenescens 475, 478, 479, 480,
481, 485, 488, 490, 491, 492, 493, 495,

scabripes 310

500, 501, 502, 504, 511, 514; Nelsoni
478, 479; nidus-avis 475-481-525;
Sabinae 481

Gyrodon 225, 226, 229, 230, 243-250,
527, 537, 538; subgen. Paragyrodon
242; caespitosus 249; capensis 250;

castanellus 250; Emilei 250; filiae 250;
fusipes 250; immutabilis 250; inter-
medius 244, 246, 284; Ledermannii 250;
lividus 243, 244, 249; lividus ssp. alneti
249; lividus ssp. labyrinthicus 249;
merulioides 244, 245, 246, 247, 248, 249,
255, 257, 284; Mikhnoi 249; miramar

250; Mougeotii 250; Oudemansii 250;

placidus 250; proximus 244, 246, 247;
purpureus 249; Rompelii 244, 245, 246,
247, 249, 284; rubellus 250; rubrescens
249; sistotrema 243; sistotremoides
243, 249; volvatus 250; sp. 244, 249

INDEX TO VOLUME 2

Gyrodonteae trib. 230, 231

Gyrodontoideae subfam. 226, 230

Gyroporoideae 223-303

Gyroporus 225, 226, 229, 230, 231-240;
albisulphureus 240; asprellus 239; at-
roviolaceus 232, 239; bisporus 240;
caespitosus 239; castameus 231, 232,
233, 237, 299; castaneus var. fulvidus
237; cyanescens 231, 232, 234, 235,
236; cyanescens var. lacteus 235; de-
flexus 239; Earlei 239; fumosiceps 240;
jamaicensis 240; Jacteus 235; lividus
243; pisciodorus 240; porphyrosporus
115, 239; praeanisatus 240; purpuri-
nus 232, 236, 237, 238; Rhoadsiae 240;
roseialbus 232, 233, 234, 235; rufus
239; scaber 239; stramineus 240; sub-
albellus 232, 233, 234; umbrinisqua-
mosus 231, 234, 235

Halimeda sp. 440, 443

Hantzschia 203; amphioxys 203; elongata
203

Haplotrichum albidum 334

Heterostichae sect. 176, 184

Hexajuga 550; prunulus 551, 552

Himantidium arcus var. curtum 157;
bidens 162; exiguum 157; gracile 158;
majus 160; minus 160; veneris 164

Hirtellini subsect. 259, 273

Hookeriaceae 316

Hygroamblystegium 316-317; filum 317;
filum var. compactum 317; fuegianum
317; fuegianum var. excurrens 317;
fuegianum var. gracilis 317; fuegia-
num var. secundum 317; fuegianum
var. Skottsbergii 317

Hygrocybe 49

Hygrophoropsis 100, 539, 544-547; al-
bida 547; aurantiaca 539, 544, 545;
aurantiaca var. nana 546; aurantiaca
var. typica 545; tapinia 539, 544, 545

Hygrophoropsis subgen. 544

Hymenoscypha Curreyana 428; Duriae-
ana 391

Hypnum 559;
leucocytus 319

Hypomyces 358

Hypochnus 39

Hypocrea citrina 459; patella 459; rufa
459 -

Hypocreales 458-459

austro-stramineum 317;

Involuti sect. 539, 543
Ixechinus 223, 225
Ixocephali sect. 107, 135

575

Ixocomus 40, 225, 253, 258; sect. Piperatt
47; sect. Pseudotsugini 257; Bellini
42; cembrae 277; flavus 260; granula-
tus 42; hirtellus 274; Lakei 257; luteus
258, 270; piperatus 46; punctipes 277;
sibiricus 260

Jugasporaceae 102, 527, 544, 547-562
Krombholzia 262; porphyrospora 113

Lactarius sect. Plinthogali 48; acer 48;
azonites 48; fuligineus 48; lignyotus
48; nigroviolascens 47-48

Lamprothamnus 305

Lanceolatae sect. 203

Laricogomphus subgen. 528, 536

Larigni sect. 259

Lasiosphaeria pezizula 466

Latiporini subsect. 259, 260

Laurencia 440

Leccinum 99, 104, 225, 228, 229, 239,
262; chalybaeum 99, 258; constrictum
235; rubropunctum 258

Lemanea 68

Lembophyllaceae 316

Lenzites 527

Leptonia 50

Leptoporini subsect. 259

Leptosphaeria Cecropiae 466; Smilacis
466

Leptotaceae 547

Leptotus 547

Leskea nitida 318

Leucobolites 231

Leucoconius 231

Leucogyroporus deflexus 239

Leucoporelleae trib. 230, 231

Liagora sp. 442

Lineolatae sect. 176, 177

Linospora Trichostigmae 467

Macromitrium coriaceum 315; macro-
sporum 316; Rusbyanum 316

Macrosporus sect. 528, 536

Maiores subgen. 187, 193

Megalodonta Beckii 207

Megaporini subsect. 259

Melampsoraceae 488

Melanomphalia 562

Melanopsamma Milleri 465

Melosira 146-147; ambigua 146; italica
146, 147, 210; roeseana 146, 147; roese-
ana var. epidendron 147, 211, 212;
varians 146, 147

Meridion 149-150; circulare 149, 210,

576

211; circulare var. constricta 150, 211;
constrictum 150

Merulius 537, 544; aurantiacus 545

Mesoleiae sect. 176, 184

Mesotrema 440 :

Metasphaeria phyllachoracearum 467; sp.
468

Microdictyon Setchellianum 442

Microneis subgen. 166

Microsporus sect. 528, 537

Mielichhoferia leptoclada 312; multiflora
312

Minusculae sect. 176, 183

Mirabilis sect. 107, 128

Molendoa andina 310; boliviana 310;
fuegiana 309, 310; Herzogii 310

Mouceron trib. 550

Muratella 323, 325; elegans 323, 324, 327,
328; sp. 328

Myrioconium ambiens 391, 395, 396;
Scirpi 413; Scirpicola 413; tenellum
428

Myxocollybia velutipes 264

Myxogomphus subgen. 528, 536

Navicula 170, 176-187, 209, 213; sect.
Bacillares 176, 181; sect. Decipientes
176, 187; sect. Decussatae 176, 177;
sect. Entoleiae 176, 182; sect. Hetero-
stichae 176, 184; sect. Lineolatae 176,
177; sect. Mesoleiae 176, 184; sect.
Minusculae 176, 183; sect. Orthostichae
176, 177; affinis 173; americana 181;
amphigomphus 174; amphirhynchus
173; ampliata 174; anglica 178; an-
gusta 178; atomoides 185; atomus 183,
184; bacilliformis 185; bacillum 182;
bisulcata 173; Brebissonii 192; capitata
180; cincta 177, 178; cincta var. an-
gusta 178; cocconeiformis 184; con-
cava 182, 183; contenta var. biceps
182; crassinervia 171; cryptocephala
177, 178; cryptocephala var. pumila
177, 179; cryptocephala var. veneta
177, 179; cuspidata 177; decussis 178,
179; dicephala 178, 179; essox 193;
exigua var. capitata 178, 179; fasciata
189; festiva 183, 184, 210, 212; Flo-
towii 183; gentilis 194; gibba 202;
globulifera 177, 180; halophila var.
minor 187; hassiaca 182; hungarica
var. capitata 177, 180; iridis 173;
Keeleyi 182, 183, 211, 212; lanceolata
178, 180; mesolepta var. stauroneifor-
mis 190; minima var. atomoides 185,
207, 210, 211; muralis 183, 184; mutica

FarLowlA, VoL. 2, 1946

185; nodosa 188; perpusilla 182, 183,
211, 212; placenta 177; placentula 211;
poconoensis 182, 183; pseudoscuti-
formis 184; pupula 185; pupula var. ba-
cillaroides 186; pupula var. capitata
186; pupula var. rectangularis 186;
quadripartita 178, 180, 212; radiosa
178, 180, 208; radiosa var. tenella 177,
181, 208, 209, 211; rhomboides 170;
rhomboides var. amphipleuroides 170;
rhynchocephala 178, 181; Roteana 185,
186, 207, 212; scutiformis 184; semen
187; Smithii 174; stomatophora 192;
subatomoides 185, 186, 207, 212; sub-
linearis 189; tenella 181; Thingvallae
179; transversa 194; trinodis 172;
Trochus var. biconstricta .172; turgida
202; variostriata 185, 186; veneta 179;
viridula var. linearis 178, 181; vitrea
184

Neidium 173-174, 213; affine 173; affine
var. amphirhynchus 173; bisulcatum
173; bisulcatum var. undulatum 173;
iridis 173; iridis var. amphigomphus
173, 174; iridis var. ampliatum 173,
174; iridis var. vernalis 173, 174

Nemalion 63

Nitella 305; flexilis 208

Nitzschia 203-204, 213, 214; sect. Gruno-
wiae 203; sect. Lanceolatae 203; sect.
Nitzschiellae 203, 204; acicularis 204;
amphibia 203, 204; elongata 203;
Hantzschiana 203, 204, 211; Hantz-
schiana fa. subserians 204, 211, 212;
palea 203, 204, 210, 212; palea var.
debilis 204, 210, 212; palea var. tenui-
rostris 204; sinuata var. Tabellaria 203,
209; vivax var. elongata 203

Nitzschiellae sect. 203, 204

Nolanea 50

Nostoc 62

Nymanomyces Xolismae 458

Octojuga 549, 550; Fayodi 558; Passe-
ckeriana 560; pleurotelloides 558; vari-
abilis 558

Odontidium 148; anceps 148, 211; hie-
male var. mesodon 148, 210, 211

Oedocephalum albidum 334; echinula-
tum 323, 326, 327

Omphalia 550; Giovanellae 549; 553;
scyphoides 549, 554

Omphalina floridana 554, 556; scyphoides
554

Orthostichae 176, 177

Orthotrichaeceae 315-316

INDEX TO VOLUME 2

Padina sp. 442, 443

Palustres sect. 251, 252

Panuoides sect. 538, 540

Paragyrodon 226, 230, 242-243; sphae-
rosporus 242 i

Paragyrodon subgen. 242

Parallelestriatae subgen. 187, 188

Parasitici sect. 287, 297

Patella coprinaria 456, 457; theleboides
456, 457

Paxillaceae 102, 527, 537-547, 549

Paxillopsis 539, 548; prunulus 551

Paxillus 98, 100, 527, 538, 539, 540, 544,
548; sect. Atrotomentosi 538, 540;
sect. Involuti 539, 543; sect. Panuo-
ides 538, 540; acheruntius 541; argen-
tinus 537, 543; atrotomentosus 538, 540;
aurantiacus 545; corrugatus 543; Cur-
tisii 539, 541, 543, 544, 545; Fagi 545;
flavidus 280, 544; infundibuliformis
284; involutus 538, 539, 540, 543, 548,
550; ionipus 541; lamellirugus 541;
lateritius 543; lignews 541; miniatus
543; panuoides 538, 539, 540, 541, 543,
544; paradoxus 280, 544; Pelletiert 280,
544; pinguis 280, 543; polychrous 538,
540; porosus 247; prunulus 544, 551;
prunulus var. orcella 551; pubescens
533, 535, 336, 543; reniformis 544; rho-
doxanthus 280, 544; rudis 541; russu-
loides 543; sulcatus 280, 283; sulphureus
544; Tammii 280, 544

Peziza ciborioides 427; Curreyana 423,
427, 429, 431; Curreyi 427; Duriaeana
391, 394 395, 399; Friesii 427; juncifida
428, 431; tuberosa 417

Pezizales 455-457

Phacidiales 458

Phaeogyroporus 225, 226, 231, 240-242;
Braunii 240, 241, 242; tropicus 241,
242

Phaeoporus 105, 115; porphyrosporus
106, 115; pseudoscaber 106

Phaeotrabutia Smilacis 460; sp. 461

Phialea Curreyana 427

Phillipsia domingensis 455, 456

Philonotis acicularis 315; luteola 315;
parallela 315; vagans 315; varians 315

Phlebopus 225, 228, 229, 258; capensis
250; sulphureus 250, 300

Phyllachora Acaciae 461, 463; Bour-
reriae 461; Exostemae 462; gratissima
462; insueta 462; Masini 463; ocoen-
sis 463; Ocoteicola 463; panamensis
464; Paspalicola 464; Phaseoli 464;

ae

Roureae 464; Scleriae 464; viequesen-
sis 464

Phyllobolites 107, 139, 543.

Phylloporus 97, 223, 224, 225, 227, 229,
257, 279-284, 285, 286, 287, 291, 293,
295, 296, 298, 538, 544; bogoriensis 280;
infundibuliformis 284; intermedius
246, 284; lariceti 250, 251, 284; malac-
censis 121, 284; paradoxus 280, 284;
Pelletieri 279; rhodoxanthus 244, 251,
280, 282, 283, 284, 296, 537, 544; rho-
doxanthus ssp. americanus 280, 282,
283, 284; rhodoxanthus ssp. bogorien-
sis 280, 282, 284; rhodoxanthus ssp.
europaeus 279, 280, 282, 284; rhodo-
xanthus ssp. foliiporus 280, 281, 282,
537; Rompelii 247, 284; squarrosoides
284, 295; sulcatus 280

Phyllosticta Citharexyli 468; Stigma-
tophylli 469

Physalospora Phaseoli 464

Pinnularia 187-195, 210, 213; subgen.
Brevistriatae 187, 188; subgen. Capi-
tatae 187, 189; subgen. Complexae 187,
194; subgen. Distantes 187, 188; sub-
gen. Divergentes 187, 192; subgen.
Maiores 187, 193; subgen. Paralleles-
triatae 187, 188; subgen. Tabellariae 187,
191; acrosphaeria 188; acrosphaeria
var. turgidula 188; amphicephala 189;
borealis 188; Braunii var. amphi-
cephala 189, 190; Brebissonii 192;
brevicostata 188; Burkei 189; capi-
tata 180; cincta 178; Clevii 193; com-
mutata 195; divergens 192, 193; di-
vergens var. parallela 193; essox 193;
fasciata 188; gentilis 194; gibba 191;
gibba var. rostrata 192; Hilseana
189, 190, 209, 212; interrupta 189,
190; legumen 192, 193; maior 194;
maior var. transversa 194; mesolepta
var. stauroneiformis 189, 190; nobilis
194; nodosa 188; obscura 192, 193,
210, 212; ovalis 174; parallela var.
divergens 193; stomatophora 191, 192;
streptoraphe 194; subcapitata 189,
190; subcapitata var. paucistriata 189,
191; subcapitata var. stauroneiformis
189, 190, 191; sublinearis 189; sub-
stomatophora 192; sudetica 195; viri-
dis 194, 195; viridis var. elliptica 195;
viridis var. fallax 195; viridis var.
sudetica 195, 207

Pinuzza 257; flava 257

Piperati sect. 47, 259, 278

Piptocephalis 323

578

Placosphaeria Junci 428, 429, 431

Plagiotheciaceae 318-319

Plagiothecium nitidum 318

Pleospora Doidgeae 468

Pleuropus minimus 554, 556; obesus Bai
$52; prunulus 551

Pleurotelloides sect. 551, 557

Pleurotellus Passeckerianus 560

Pleurotus mutilus 554, 561; Passeckeri-
anus 560; viticolus 558

Plicaturella 539

Plinthogali sect. 48

Polyporoletus 562

Polyporus Maxonii 279

Polytrichaeceae 319

Polytrichum commune 319

Porphyra 24

Porphyrellus 98, 103, 105, 106, 113, 115-
122, 527; sect. Graciles 107, 119;
sect. Tristes 107, 115; alboater 122°
Cookei 122; gracilis 104, 113, 120,
121; indecisus 122; iseunosus (21,131;
malaceenats 104, 120, 121, 131, 284:
nigrellus 122; porphyrosporus 115;
pseudoscaber 103, 104, 105, 106, 115,
119, 121, 133, 239: pseudoscaber ssp.
cyaneocinctus 116, 119, 123; subfla-
vidus 99, 104, 120, 122, 131; tristis
104, 118

Porphyrospori 106

Porphyrosporus subgen. 115; subflavidus
104

Prachtflorella microspora 336

Prunuli sect. 550, 551

Psathyra 118

Psathyrella 118

Pseudoboleti sect. 287, 299

Pseudoleskia 316; fuegiana var. gracilis
317; fuegiana var. Skottsbergii 317

Pseudophyllopori sect. 287, 296

Pseudotsugini sect. 257

Psiloboletinus 226, 250-251, 284; lari-
ceti 250

Pterospori sect. 106, 114

Ptilota serrata 64

Puccinia Helianthi 486; nidus-avis 481

Pulveroboletus 97, 225, 228, a205 240.
240, 258; auriporus 300; corrugatus
258; gentilis 300; retipes 230

Retispori sect. 107, 130

Rhacomitrium crispulum 311; limbatum
311; ptychophyllum 311; scabrifolium
311

Rhagadolobium cucurbitacearum 465

Rhodocybe 527, 548, 549; alutacea 548;

Fartowia, VoL. 2, 1946

fallax 549;
cularis 548.
Rhodogoniosporaceae 527, 549
Rhodopaxillus 548; fallax 549;
tus var. subvermicularis 548
Rhodophyceae 439-453
Rhodophyllus 50, 51, 527, 548, 549, 561;
abortivus 561; albogrisens 561; Far-
lowii 49-51; salmoneus 50; Pease
551
Rhopalodia 202-203, 214; gibba 202
Rhymovis 539, 540; panuoides 541
Rhymoxis 540
Rigodiella sect. 316
Ripartites 538
Roestelia 476; cornuta 476; lacerata y
476; Nelsoni 479; nidus-avis 481
Rosellinia sp. 465
Rostkovites 257, 258;
278; granulatus 257;
Ruthea 539
Rutstroemia 436; Curreyana 427

truncata ssp. subvermi-

trunca-

californica 277,
hirtellus 274

Saccharomyces pulcherrimus 366; pul-
cherrimus secundarius 366

Saccobolus Kervernii 456, 457

Saitomyces 325; japonicus 334

Sargassum echinocarpum 442; sp. 440,

443

Sauloma tenella 316

Schistidium sect. 311

Scleroderma 224, 298

Sclerotinia 385-437; Aschersoniana 435,
436; Caricina 436; Caricis-ampullaceae
387, 389, 405, 407; Curreyana 387, 423,
425, 427, 429, 431, 432, 433, 434, 435;
Binineane 386, 389, 391, 394, 395, 396,
399, 401, 403, 408, 411, 421, 423, 425,
427, 432, 433; Dubaans Wee haan
form" 397 ; Gladioli 420; heterocarpa
436; Juncigena 387, 432, 433, 435;
fonerclerotale 387, 388, 389, 401, 403,
405, 417; Luzulae 387, 433, 434; palu-
dose 436; Schoenicola 387, 423, 425;
Sarateals 387, 413, 417, 418, 420, 421,
423, 425, 428, 431, 432; ealavatiorin:
386 ; sulcata 387, 389, 394, 395, 396,
397, 399, 401, 403, 420, 421, 423, 425,
432, 433; utriculorum 435; Vabhane,
387, 408, ‘409, 411, 412, 413, 435; Vesi-
cariae 436

Sclerotiniaceae 385

Sclerotium Kneiffii 423; Luzulae 434;
nigricans 391, 394, 398; roseum 413,
417, 423, 431, 435; sulcatum 394, 397,
399, 401, 423, 424

INDEX TO VOLUME 2

Scutigeraceae 562

Scyphoides sect. 550, 553, 560 .

Secotium excavatum 123, 126

Sematophyllaceae 319

Sematophyllum aureo-nitidum 319; cal-
lidum 319

Sigmoideomyces 324

Solidipedes sect. 251, 253, 257

Sordaria curvula 465

Spectabiles. sect. 251, 252

Sphacelia ambiens 391, 394; Curreyana
428; nigricans 391, 395; Scirpicola 413;
tenella 427, 431

Sphaeria calyculus 466; pezizula 466 -

Sphaeriales 465-468

Sphaerostilbe hypocreoides 459;
media 459

Sphenella obtusata 200; parvula 201

Sphenosira epidendron 147

Splachnaceae 312

Stauroneis 174-176; amphicephala 175;
anceps 174, 175; anceps var. amphice-
phala 175; anceps var. gracilis 175;
anceps var. linearis 175; gracilis 175;
Kriegerii 174, 175; linearis 175 ; phoeni-
centeron 174, 176, 210; pygmaea
175; Roteana 186; Smithii 174, 176

_Stauroptera gibba 191

Staurosira construens 151

Stegastroma 467

Stenoporini subsect. 259, 278

Stenopterobia 204-205; intermedia 204

Stilbocrea hypocreoides 459

Strobilomyces 98, 103, 104, 105, 106,
108-115, 258, 527; sect. Genuini 106,
108; sect. Pterospori 106,114; ananae-
ceps 123; annamiticus 126; coccineus
123, 125; confusus 108, 113, 114;
costatisporus 104, 115; echinatus 114;
excavatus 123; fasciculatus 139; floc-
copus 103, 104, 106, 108, 109, 110, 112,
113, 114, 130; indicus 113; lepidellus
115; ligulatus 139; montosus 114; nigri-
cans 108, 113, 114; pallescens 97, 123,
125, 139; paradoxus 138; polypyramis
114; porphyrius 126; pterosporus
104, 114, 115; rufescens 139; strobila-
ceus 106, 110, 112, 113; velutipes 105,
106, 108, 113, 119

Strobilomycetaceae 97-141, 538

Subtomentosi sect. 287, 297

Suillellus pictiformis 131

Suilloideae 226, 250, 251

Suillus 98, 225, 227, 229, 230, 231, 250,
253, 257-279, 299; sect. Bovini 259,
278; sect. Granulati 258, 259, 260; sect.

inter-

579

Larigni 259; sect. Piperati 47, 259,
278; subsect. Angustiporini 259, 260,
267; subsect. Euryporini 259, 278;
subsect. Hirtellini 259, 273; subsect.
Latiporini 259, 260; subsect. Lepto-
porini 259; subsect. Megaporini 259;
subsect. Stenoporini 259, 278; acidus
271; aeruginascens 259; albidipes 45,
261, 267, 272; americanus 260; atrovio-
laceus 239, 279; australis 98; Bellinii
267; bovinus 243, 278; Brauniit 241;
brevipes 41, 44, 45, 259, 261, 265, 267,
271, 272, 273; brevipes var. aestivalis
267, 271; brevipes var. pseudogran-
ulatus 268, 269; brevipes var. typicus
265, 268; cantharelloides 279; casta-
neus 237, 278; cembrae 259, 274, 277,
278; cothurnatus 260, 261, 264, 265,
268, 269, 270, 271; cothurnatus ssp.
aestivalis 261, 263, 265; cothurnatus
ssp. hiemalis 261, 263, 264, 265; cya-
nescens 235, 278; decipiens 254; flavi-
dus 260, 269; flavoluteus 260; flavus
260; granulatus 40, 41, 43, 44, 250, 260,
261, 267, 272, 273; granulatus ssp.
leptopus 42, 43, 44, 45, 261, 267, 272;
granulatus ssp. Snellii 40, 41, 42, 45,
261, 267, 270, 272; granulatus ssp.
typicus 41, 42, 44, 261, 272; Grevillei
259; hirtellus 98, 259, 268, 273, 274,
277; hirtellus ssp. cheimonophilus
274, 276; hirtellus ssp. thermophilus
274, 275; hirtellus ssp. typicus 277;
hirtellus var. mutans 274, 277; hygro-
phanus 279; intermedius 246; jamai-
censis 240, 279; lanatus 296; luteus 41,
258, 260, 265, 266, 268, 269, 270, 271,
273, 275; Maxoni 279; piperatus 46,
278; piperatus var. amarellus 46, 278,
300; placidus 42, 43, 45, 250, 260,
271; placidus fa. americana 43; placi-
dus fa. Pint cembrae 43; placidus fa.
Pint halpensis 43; placidus fa. Pinz
strobi 43; punctatipes 257, 277; punc-
tipes 259, 262, 263, 270, 273, 277; ru-
binellus 47, 278; sibiricus 260; sawbal-
bellus 232, 279; subaureus 260; sublu-
teus 260, 264, 265, 269, 270, 271;
tridentinus 260; variegatus 278

Surirella 205-206, 213, 214; Amphioxys
206; angusta 205; biseriata 205; gra-
cilis 205; guatemalensis 205; interme-
dia 204; Moelleriana 205, 206; Palmeri
206; splendida 205, 206; tenera 205,
206; tenera var. nervosa 206; tenera
var. Palmeri 206

580

Synedra 153-155, 160; acicularis 204;
acus 153; acus var. delicatissima 153;
danica 155; debilis 204; delicatissima
153; familiaris 154; lunaris 159; luna-
ris var. capitata 159; minuscula 153,
154; oxyrhynchus 155; palea 204;
Ramesti 155; rumpens 153, 154; rum-
pens var. familiaris 154; rumpens var.
scotica 154; ulna 153, 154, 210; ulna
var. danica 155, 208, 211, 212; ulna
var. oxyrhynchus 155; ulna var. Ram-
esli 155

Tabellaria 148-149, 213; fenestrata 148,
208, 209, 210, 211; flocculosa 149, 207,
208, 209, 210, 211, 212

Tabellariae subgen. 187, 191

Taphrina deformans 345-383; epiphylla
349, 353; Klebahnii 349, 353

Tapinella 539; panuoides 540, 541

Tapinia 539, 540; lamellosa 541; panu-
oides 540, 541

Tayloria magellanica var. gigantea 312

Telimena domingensis 467, 468; sp. 468

Thamnocephalis 324

Tibiella 214; punctata 213

Tomentella 39

Tortula pseudorobusta 310; rivularis
310; robusta 310; robusta var. laxa
310

Torula histolytica 347; rubefaciens 371

Torulopsis neoformans 347, 375; pul-
cherrima 345-383

Trabutia conica 460; Danthoniae 460

Tremella nidus-avis 481; sp. 495

Tricholomataceae 527, 548, 562

Tristes sect. 107, 115

Truncocolumella 255

Tryblionella gracilis 205

Tubiporus porphyrosporus 115

Tuomeya 68

Tylopilus 105, 106, 121, 122, 225, 228,
230, 240; alboater 122; conicus 99;
felleus 121; gracilis 121; plumbeovio-
laceus 258; Rhoadsiae 240; subflavidus
120; tabacinus 240

Typhula 436

FARLOWIA, VoL. 2, 1946

Uloporus 243; lividus 243, 249; lividus
var. rubescens 249

Vanheurckia 170-172, 212, 213; rhom-

amphipleuroides 170, 211, 212; rhom-
boides var. crassinervia 171, 207, 208,
209, 212; rhomboides var. crassinervia
fa. capitata 171, 208, 212; vulgaris 171
Versipellis 258
Viscipellis 258; lutea 258; variegata 258
Viscogomphus ‘sect. 528, 529
Volvoboletus 562

Wormskioldia 24

Xanthoconium 223, 225, 228, 230; stram-
ineum 234, 240

Xerocomoideae 227, 251, 279

Xerocomopsis 284

Xerocomus 128, 224, 225, 227, 230, 258,
279, 284-300, 538; sect. Brasiliensis
287, 298; sect. Parasitici 287, 297; sect.
Pseudoboleti 287, 299; sect. Pseudo-
phyllopori 287, 296; sect. Subtomen-
tosi 287, 297; amarellus 300; armenia-
cus 300; auriporus 300; badius 258,
287, 299; Boudieri 299; brasiliensis
258, 297, 298; chrysenteron 297; chry-
senteron var. versicolor 300; conifer-
arum 286, 287, 288, 297; flaviporus
300; gentilis 300; griseus 300; hemi-
xanthus 288, 292, 296; Housei 288,
296; hypoxanthus 288, 289, 293; illu-
dens 281, 285, 286, 287, 289, 290, 293,
295, 296; illudens ssp. typicus 289, 294;
illudens ssp. xanthomycelinus 289,
294, 295; impolitus 285, 300; Junghuh-
nii 297; lanatus 288, 296; lignicola
299; Linderi 288, 296; mirabilis 129,
300; parasiticus 224, 297, 298; pruin-
atus 300; pseudoboletinus 288, 290;
pulverulentus 300; radicans 300; rubel-
lus 300; spadiceus 286, 297; squarro-
soides 250, 255, 257, 288, 293, 295,
296; subilludens 100; subtomentosus
281, 285, 286, 297; subtomentosus var.
Leguet 297; subtomentosus ssp. spadi-
ceus 297; sulfureus 300; tumidus 300;
versicolor 300; Zelleri 297; sp. 300

MANUSCRIPT

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PUBLICATIONS FOR SALE

CLINTON, G. P. North American Liman 1904. (Proc.
Bost. Soc. Nat. Hist.) Unbound... .. . os 4. $ 2,00

FARLOW, W. G. Icones Farlowianae. Illustrations of the Larger
Fungi of Eastern North America. 103 colored plates by J.
Bridgham and L. C. C. Krieger. Folio. 1929. Cloth. . . . 50.00

JOURNAL OF MYCOLOGY. vol. 5, nos. 2, 3, 1889; vol. 6,
1890-91; vol. 7, 1891-94. Unbound. Thelot .... . . . 750

PERSOON, C. H. Index Botanicus to his Mycologia Europaea,
prepared by Dr. and Mrs. D. P. Rogers and Dr. E. V. Seeler, Jr.
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SEYMOUR, A. B. Host Index of the Fungi of North America.
Paes 1929. CBE a 7.50

THAXTER, R. Contribution towards a Monograph of the Laboul-
beniaceae. (Mem. Amer. Acad. Arts & Sci.) Unbound.

I. 1896. (Out of a

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THAXTER, R. Entomophthoreae of the United States. 1888.
(Mem. Bost. Soc. Nat. Hist.) Unbound . . .... + 3.00

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