-Vol1 ‘JANUARY, 1943. = No.1
- FARLOWIA.
A JOURNAL OF Cryprogamic BoTany
Eprrortat Boarp a5
DAVID H. LINDER, Editor
E. V. SEELER, Ja, Mg. Ed.
. R. Bartrotomew - -G. H. Ripa
__ F. Verpoorn
- 'W. H. Weston, Jr.
_R. M. Wuetpen
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CONTENTS OF VOL. I, NO. 1
PORMWORO. OVS) GUIE dt LUO ues os orca vn ck ook bs ae ee 1
Tue Arms or Furure ReskarcH in THE Atcan. By V. J, Chapman........ 5
Notss on New ENGLAND Augan, III. By Roy M. Whelden................ 9
Apive Mossus rrom New Guinma. By H. N. Dicon........ Pee A RO
Mossss or Papua, New Guinzga. By Edwin B. Bartram. Sara SS Seay meget 41
Tue Genera KickxEa, MaRrenseia, AND ComMANsIA (Phycomycetes).
EY PAU Pes SONOS obs hae y GV cate he bane Nae aE) CP aE OES 49
Basipium ForRMATION anp Sporn DiscHareH IN GrunosroraNerum Nipus-
Avis. By Alton Bi Prince. 7 2 5a er aus. See onee 79
Tan Genus PeciicunaRta (Thelephoraceae). By Donald P. Rogers....., oa 06
Soman Funeicotovs Fouwnat. By Rdgar ¥: Beeler, IR; «sep See sek ove wok 119
STupius IN THE Gmnvs cua TIT ( Discomycetes). By W. Datprence White. 135
FARLOWIA _
A JOURNAL OF CRYPTOGAMIC BOTANY
VOLUME 1
— 1943-1944 |
ee ar
PUBLISHED BY
FARLOW LIBRARY AND HERBARIUM OF HARVARD UNIVERSITY
: 20 Divinity AvVE., CAMBRIDGE, Mass.
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FARLOWIA
A JOURNAL OF CRYPTOGAMIC BOTANY
Vou. 1 JANUARY, 1943 oT NOt)
FOREWORD
The question may be raised: Why have a journal for taxonomic
cryptogamic botany? To those who query the desirability of inaugurat-
ing a periodical of this description, the following brief answer is offered.
Those who have been interested in cryptogamic botany in its taxonomic
aspects have long recognized the need of monographic studies, either
taxonomic, morphological, cytological, or ecological, which might con-
tribute to the systematic arrangement of the many and diverse groups
of organisms which constitute the non-vascular cryptogams. Likewise
it is evident that taxonomy is the keystone of all biological studies since
it provides the investigator in other fields of endeavor a label for the
specific unit of plant life that is being studied, thus making it possible
for future students to confirm or to amplify the previous investigations.
The stability of the name employed to label the organism depends to
a great extent on the thoroughness with which the special group has been
studied, how carefully the older and often imperfect records have been
evaluated, how widely acquainted with related groups is the writer, and
how well he has incorporated the information that may have been pro-
vided by morphology, cytology, genetics, ecology, chemistry and other
related fields. Unfortunately, our knowledge is still meager, but on the
basis of that which we do possess, it may be stated that accurate tax-
onomic investigations not only borrow from but contribute to related
branches of knowledge. At the same time, it may be said that this
discipline is a prime necessity for the validation of results, either from
a scientific point of view or, since all knowledge may be of eventual prac-
tical value, from the viewpoint of its contribution to human welfare.
Frequently the systematist, through his special knowledge of the organ-
isms that come to his attention, is in a position to save a considerable
amount of time for the chemist, the physician, the physiologist, or the
bio-physicist since he is able as a result of his taxonomic studies to
suggest a single species or a group of species that will satisfy the needs
of the investigator in the other fields of research. This contribution
is almost wholly forgotten because it is less dramatic than are the
results not infrequently secured by those, who without any attempt to
belittle them may be called test-tube biologists, who started their studies
where generations of taxonomists left off. Many examples could be
1
2 FARLOWIA, VOL. 1, 1943
chosen from the broad field of cryptogamic botany which would demon-
strate clearly the dependence of other branches of biological science on
the results of the many years of labor of the collectors and the taxonomists
and which would illustrate the value of the so-called useless information.
Probably no better example could be chosen than the situation that has
arisen as a result of the cutting off of America’s supply of agar-agar by
the Japanese who with their cheap labor and favorable localities have
a monopoly on the harvest and production of the algal material. With
the cessation of imports, commercial concerns have turned to the algae
of the American coasts for substitutes for the agar that they used in the
preparation of hand-lotions, ice cream, and other products. Commercial
needs are more easily supplied than are those of the bacteriologists, the
public health officers, and the plant and animal doctors who require a
substance of rather specific physical characteristics for the hardening of
the culture media that they employ for growing the pathogenic organ-
isms which they are studying in an effort to control diseases. It is im-
mediately apparent that agar plays a very important role in the welfare
and nutrition of this and allied countries. The acute shortage of this
material has made it necessary that substitutes be found at the earliest
possible opportunity. Fortunately, as a result of the activities of the
collectors, many of whom were amateurs who made the gathering of
representative algae a hobby, and of the taxonomists whose studies have
made known the morphology and structure of the algae, there is present
and available for use, either in the literature or in the various herbaria,
a large amount of information that is of immediate value. Those who
are today seeking a substitute for the Japanese agar have been spared
the large amount of pioneering work that has been done for them, and
can turn their attention without loss of time to the analysis of the more
likely species.
The above may be considered by some to be an excuse for the con-
tinuation of taxonomy, but taxonomists directly or indirectly have made
too many contributions to knowledge to apologize either for their exis-
tence or their errors. Rather it is intended to prophesy how little credit
will be given to the taxonomists and collectors and to emphasize the
reason for the relatively small amount of support received by them
today. It is this lack of support that has made it extremely difficult
for American systematists to publish the longer monographic studies of
which there is great need. Part of the difficulties arise from the fact
that the papers submitted for publication are in competition with those
dealing with current scientific fads or else those having more immediate
practical value and they are accordingly judged to be of less importance.
The other side of the story is that most journals are published by
academies, societies or institutions which for the most part must limit
the privilege of publication to their members or staffs, and since the
membership is comprised of many individuals with almost as numerous
FOREWORD 8
interests, articles must be limited in number and length in order that
all may have a fair opportunity to make public the results of their
investigations. To remedy this situation and to make possible the pub-
lication of longer taxonomic papers, it has been decided to publish the
present journal which for the time being will include articles up to
one hundred pages in length and that deal with taxonomy, cytology,
morphology and the related fields that contribute to the classification
of the non-vascular cryptogams. The first volume will be published
semi-annually over a period of two years, after which it is hoped that
the uncertainties brought about by the war will be removed and that
the support of the journal by cryptogamic botanists will make possible
the establishment of FaRLOwIA as a quarterly journal. Thus those
of us who have enjoyed the fruits of his contributions to the develop-
ment of American cryptogamic botany by his teachings or through the
use of the facilities of the herbarium and library that he had brought
together and dedicated to the advancement of this special field of botany,
may fittingly acknowledge in 1944 the centenary of the birth of William
Gilson Farlow.
Davip H. LInDER, Curator
FarRLow HERBARIUM
HarvarRD UNIVERSITY
1(1): 5-8 FARLOWIA January, 1943
THE AIMS OF FUTURE RESEARCH IN THE ALGAE
V. J. CHAPMAN
Since the beginning of this century very considerable progress has
been made in our knowledge of the algae. Up to then interest centered,
as was perhaps natural, around morphology and systematics, and at
the same time compilation of flords proceeded apace. Now, more atten-
tion is being given to the ecology, physiology and reproduction of the
algae, but these branches are still in their infancy. This change in
outlook does not imply that no future progress can occur in morphology
and systematics, but it does mean that the greatest advances of the
future can be expected to be made in these younger branches. For this
reason research ‘in them should be encouraged, though the older aspects
should not be neglected since they still offer many problems that remain
to be solved.
Morphology and systematics are an essential background to any
botanical subject, and the more complete our knowledge of them, the
more likelihood is there of sound and substantial advances in other
branches. Today one of the most important needs in algal systematics
would seem to be a continuation, with greater intensity, of the attack
being made upon the smaller microscopic algae, especially the marine
forms. In recent years much has been achieved in this direction. for
soil and fresh-water species and Pringsheim with his cultures (Biol.
Reviews 16: 198. 1941) has indicated the best method to be adopted.
Here one may perhaps express a hope that someone will soon publish
a book describing all these forms with the provision of a key for their
identification. There are still, especially in tropical and sub-tropical
marine habitats, a number of species which require investigation and
description. One may mention here the encrusting species of all groups:
these have been studied fairly thoroughly in Europe and to a somewhat
lesser extent in North America. Because they are so inconspicuous and
often are not found in fruit, relatively few phycologists have studied
them and many are regarded as rare. Persistent search in England has
shown that this is not correct and that the majority have a far wider
distribution than has been ascribed to them. This will no doubt be found
to be true in other countries. Our knowledge of such forms in tropical
and sub-tropical regions is exceedingly scanty and the position is not
much better in the southern hemisphere.
There are certain genera, such as Enteromorpha, which I have been
studying for some years, and Cladophora, which require to be investi-
gated with a view to monographic treatment. At present it is quite
impossible to identify with any degree of certainty some of the Clado-
phorae, more especially the tropical and sub-tropical forms. Valuable
studies of this character have been undertaken by Kylin (Lunds Univers.
Arssk. 20: 1924) for the Delessariaceae in the Rhodophyceae and by
5
6 Fartowia, Vou. 1, 1943
Parke (Hartley Bot. Lab. Pub. 9: 1933) for the Mesogloiaceae. Even
the common species of Fucus are in need of further investigation in
spite of excellent work upon them by the earlier phycologists, e. g.
Sauvageau. The salt marsh forms of this genus are particularly difficult
to determine and yet they play an important part in the algal ecology
of salt marshes. Numerous varieties of Fucus species have been described
and it is only by collecting and examining forms from many different
localities that their status will finally be determined.
If we turn from systematics to morphology a principal need would
seem to be a study of the developmental stages among members of the
Fucales. In recent years Delf (Journ. Bot. Lond. 75: 273. 1937, and
77: 129. 1939) and her school have done much towards elucidating
the development of the reproductive organs in the lesser-known species
of this group, but the young stages have been largely neglected, a fact
which can be ascribed to the difficulties of identifying them in the field
or with growing them in culture. It is important that this problem
should be overcome because it is of very considerable significance in
ecology. A very valuable start in this direction has been made by
Nienburg (Wiss. Meeres. Abt. Kiel 21: 51. 1931). Culture studies of cer-
tain algae have recently forced us to realize that we have not yet deter-
mined all those species which have very different forms for their sexual
and asexual generations. It has recently been shown (J. Behlan, Beih.
Biol. Pflanz. 27: 221. 1939) that Chlamydomonas variabilis and Carteria
ovata are alternate generations of the same species and also (P. Korn-
mann, Planta 28: 464. 1938) that the same applies to Halicystis ovalis
and Derbesia marina. These are comparable to the older classic example
of Cutleria and Aglaozonia. With the improved culturing technique we
must re-investigate the life-histories of many species with a view to ascer-
taining whether they behave similarly. Bliding (Svensk. Bot. Tidskr.
27: 233. 1933) has shown by such means that two well-marked forms
of Enteromor pha prolifera are simply two generations of a single species.
The investigations of Knight (Trans. Roy Soc. Edin. 56: 307. 1929),
Schussnig and Kothbauer (Oest. Bot. Zeit. 83: 81. 1934) and Papenfuss
(Bot. Gaz. 96: 421. 1935), which show that Ectocarpus siliculosus may
have a different life cycle in different parts of the world, lead us to the
conclusion that we must not be satisfied with a single study of a life
history, however complete it may appear to be. The Chaetophorales is
an order for which it is very desirable to have some advance in our
knowledge of life cycles and chromosomal conditions. Such knowledge
will prove to be of the greatest value phylogenetically.
In physiological studies the greatest advances will probably come
from biochemistry, especially studies concerned with the pigments. The
classification of the algae is primarily based upon pigmentation and
hence the discovery of any unusual features, e.g the phaeophycean pig-
ment fucoxanthin in Polysiphonia violacea, in Zygnema pectinata and
CHAPMAN: RESEARCH IN ALGAE 7
the Chrysophyceae (P. W. Carter, I. M. Heilbron, and B. Lythgoe) (Proc.
Roy. Soc. 128: 82. 1939), may lead to far-reaching systematic con-
clusions. Apart from this the réle of growth substances in the algae has
been but little studied and further investigation may show that they
play as important a part in algal development as they do in the phanero-
gams. Most phycologists would now subscribe to the existence of ‘sun’
and ‘shade’ algae, but there is not the same agreement upon the criteria
that determine to which group any given species should be assigned.
Further work upon this problem is necessary in order to arrive at a
definite working basis.
Since ecology is such a young branch of phycology there are innumer-
able problems, any one of which would repay study. Some, however,
are more outstanding or of greater urgency than others. So far there
has been too little attention paid to the operation of critical levels. The
existence of these was first discussed by Colman (Journ. Mar. Biol. Ass.
U. K. 18: 435. 1933) and since then by David (Ph.D. Thesis, Aberystwyth.
1941). There is no doubt that such levels are of prime importance
but further studies are essential before we can really understand their
significance. It is also probable that in different localities the critical
levels will vary and this should provide a clue to the dominant factors.
More or less associated with this problem is the non-tidal exposure factor.
The importance of this on salt marshes has already been emphasized
(V. J. Chapman, Journ. Ecol. 27: 160. 1939) but it has not yet been
studied for the rocky litoral. It is believed that the incidence of these
periods, when no inundation occurs, has a profound effect upon the
occurrence of many species, especially when the periods operate during
the germling phase. Whether the effect is due to actual lack of water
or to loss of water or to some other cause has also to be determined.
These remarks particularly apply to the fucoids and here we can appre-
ciate how important it will be when. we can distinguish with certainty
in the field from their very earliest stages, the germlings of the different
species. Up to the present, largely because of this difficulty, insufficient
attention has been paid to the germling phase. In the case of Ascophyl-
lum, for example, this would seem to be all-important because it is absent
from areas exposed to strong wave action: young plants are not to be
found in such places so that the inhibitory effect takes place at a very
early stage.
Studies of recolonization after denudation or on new surfaces, such as
those by Rees (Journ. Ecol. 28: 403. 1940), David (1. c.) and Boken-
ham (Ann. Nat. Mus. 9: 1938), will lead to important conclusions and
help us to understand the interrelations between different species on the
shore. Such work has already shown that colonization by Fucus is
commonly preceded by the development of an Enteromorpha mat. It has
been suggested that this mat provides the necessary humidity for the
Fucus germling, but it may equally well provide a suitable substrate
8 FaRLowIA, VOL. 1, 1943
for attachment. In any case the function of this mat in promoting the
attachment of germlings requires investigation. The part played in the
different communities by microscopic and encrusting species is not prop-
erly appreciated, though in this case it is partly due to a lack of adequate
systematic knowledge. Yet again studies of areas associated with peculiar
environmental conditions, such as that of the Baltic by T. Levring (Ph.D.
Thesis, Lund, 1940), cannot help but lead to important results. The
abnormal conditions induce morphological changes or ecological varia-
tions which can often be correlated with the operation of a single factor.
Such areas are not common, but it is important that they should be
investigated wherever they occur.
Studies of geographical distribution may be regarded as a part of
ecology and here there is almost a virgin field. Some work can be
carried out with the use of existing floras, but in many areas the floras
cannot be employed because of the unsatisfactory condition of their
systematics and uncertainty of synonymy. Plant geographical studies
of the phanerogams have led to extremely important conclusions, and
there is no reason to suppose that similar studies of the algae will not
do likewise.
In fossil botany the principal interest, so far as the algae are concerned,
would seem to be with that new group, the Nematophytales (W. H. Lang.
Phil. Trans. B. 227: 245. 1937). It is to be hoped that further material
will be discovered that will provide decisive proof as to whether they
were algae or not. If an algal nature is confirmed then they will take
their place as perhaps the most interesting algae of all time.
GONVILLE AND Catus COLLEGE
CAMBRIDGE, ENGLAND
1(1): 9-23 FARLOWIA January, 1943
NOTES ON NEW ENGLAND ALGAE
Ill. SOME INTERESTING ALGAE FROM MAINE?
Roy M. WHELDEN
Casual observation of collections of freshwater algae from the numerous
waters of Maine reveals: an algal flora of surprising richness; more
careful attention shows it to be well worth intensive study to which the
following notes may be considered only as an introduction.
The collections on which these notes are based have been selected from
several hundreds of vials of algal material gathered in every month of
the year through seventeen years. Beyond the statement that no collec-
tion noted here was made from any large body of water, either lake or
river, no description can be made to include the many habitats from
which collections were made. One might stress the fact that several
separate collections were made from what appeared to be the least promis-
ing places, such as a shallow rain pool formed in a foot-path. Despite
what seemed to be a sterile habitat, a sample nevertheless was taken and
later found to contain an abundance of Closterium macilentum, a large
percentage of which was in some stage of conjugation. This and other
habitats to be mentioned subsequently seem to justify the statement that
no place should be passed by too carelessly, since surprising discoveries
may be completely missed by such lack of attention.
As originally planned, collections were made as frequently as feasible
from as wide a variety of habitats as possible: from densely shaded
brooks or pools; from cold water and some from shallow pools in full
sunlight when the water at the time of collecting was extremely warm;
from crystal clear waters to some of deepest brown color. Final selec-
tion of material to be considered is due in part to its rarity, to the oc-
currence of anomalous forms, to noteworthy variability, and to stages
of reproduction.
MYXOPHYCEAE
The presence of blue-green algae was frequently noted in a large ma-
jority of the collections; in no case did these algae occur in any quantity.
One of the most interesting of these was Eucapsis alpina Clements and
Shantz (Fig. 1) which was found in abundance in a single station, a
vegetation-filled pool in which the water was only a few inches deep,
with several feet of slimy brown ooze beneath. The Eucapsis colonies
varied in size from small ones having only sixteen cells to large com-
pound groups of more than five hundred cells. The first specimens ob-
served were collected in early June; later they were found in abundance
up to mid-November. Many colonies were examined and the component
cells carefully measured. When mature the cells were of strikingly
uniform diameter. Different colonies showed a rather wide range of
1Contribution from the Laboratories of Cryptogamic Botany and the Farlow
Herbarium, Harvard University, no. 221. :
9
10 Fartowia, Vou. 1, 1943
cell size: many were formed of cells 2-3 » in diameter — var. minor
Skuja; others, found with the small ones, were 3-4 y» in diameter. Very
many more had cells 6—6.5 » in diameter; often the large-celled colonies
were composed of the largest number of cells. The presence of this
alga was observed at this station from 1927 to 1940. Not once, despite
repeated search, was it found in any other of the extremely numerous
habitats, some very like this one and only a few rods away. Frogs and
a variety of wading and swimming birds are frequent here; yet it would
seem that they have never successfully transported this alga elsewhere.
Occurring in the same marsh with Eucapsis, and in many other places
throughout the southern half of the state is Glaucocystis nostochinearum
Itzigs. This alga seems always to occur sparingly.
In one shallow marsh, specimens of Anabaena were found in small
numbers, always occurring singly and devoid of any obvious sheath.
The spherical cells comprising each filament were 1.5 » in diameter or
slightly less; the heterocysts were also spherical and 2—2.5 » in diameter;
akinetes were always remote from the heterocysts and were cylindrical,
4.5 » in diameter and 18.5-22 » long. While somewhat smaller in all
dimensions, this seems to be Anabaena minutissima Lemm. The only
specimens found were from this single station. Anabaena flos-aquae
(Lyngb.) Bréb. is found rather frequently throughout this region. Its
var. minor W. West was found in one place in large amounts. The
numerous filaments were of cells 2.7x 2.0 » in diameter. The heterocysts
were 2.8-4 », diameter, and the akinetes, present in large numbers, varied
from 5.9x 12 uw to 84x 22 p.
The many other species of blue-green algae found in this region may
never have been reported as occurring there. However they are of such
common occurrence there and elsewhere that their presence may be
expected. For the present at least they remain unnoted.
CHLOROPHYCEAE
Among the green algae one finds many of particular interest. One
of these was a species of Closteridium (Fig. 7) which occurred rather
abundantly in a small marsh pool. The majority of the solitary cells
were semi-circular in shape with one side straight or only slightly curved.
Each pole bears a single pointed spine which may be straight but fre-
quently is distinctly curved. The cell wall is thin. The single chloro-
plastid fills nearly the entire cell and contains a single pyrenoid and
also a single nucleus, the latter approximately centrally located in the
cell. These cells vary considerably in size, being usually between 75
and 95 » long (including spines) and 30-40 » in maximum diameter.
The spines are 11-13 ». long. The smallest cell measured was only 63 p
long and 23.5 » in diameter. Among the many cells seen were a large
number apparently in various stages of division. The most noticeable fea-
ture of these was the presence of two distinct pyrenoids, which probably re-
WHELDEN: New Encianp AcaE. III 11
sulted from the division of the single pyrenoid normally present. Once di-
vided the two pyrenoids seem to move apart subsequent to which there ap-
pears a distinct division of the chloroplastid, leaving a single cell with two
distinct plastids. Several of these were seen. Further stages showing
that actual cell division did occur were not found. Cells in which these
stages occurred were 100-105 y» long but with diameters of the same
magnitude as had the normal cell. Despite the rather larger dimensions
- and coarser habit this alga seems to agree quite well with descriptions
of C. siamensis (W. and G. S. West) Wille, reported in this country
by Prescott.
Occurring with the Eucapsis considered above and apparently also
restricted to the one pool, is a species of Tetraédron which has appeared
regularly in collections made throughout many years. The thick-walled
cells are triangular with sides generally strongly convex, or occasionally
straight or slightly concave. Each angle bears a single stout, curved,
straight or rarely slightly hooked spine. The single chloroplastid con-
tains no pyrenoid. The length of the cells, spines included, varied from
102-118 », without spines from 77-84 »; the breadth, without spines,
from 50-62 ». The length of the spines is 11.3-17 uy.
W. and G. S. West in 1895 described, without any figure, as Tetraédron
tortum, a species which has much in common with that considered here.
Their species was relatively narrower, 44 » being given as maximum
breadth, twisted, and with sharp spines about 30 » long. The specimens
herein described are much plumper, in end view generally presenting
sub-circular outline; they rarely, if ever, show a shape that could be de-
scribed as twisted, and they have much shorter spines. This combination
of characters seems sufficient to justify describing this as a new species,
Tetraédron mainensis n. sp. (Fig. 4).
Tetraédron cellulis magnis, tumidis, trigonis, lateribus convexis, vel rectis vel
rarissime plus minus concavis; anguli spinas crassas rectas, vel arcuatas vel plus
minusque uncinatas ferunt; membrana crassa glabra; cellulae cum spinis 102-118 p.
longitudine et sine spinis 77-84 yp. longitudine, 50-62 y. latitudine, 40-51 yp. crassi-
tudine; spinae 11.3-17 p longitudine.
The desmids have always received much attention, partly because of
the striking variety of their many species and their beautifully symmetti-
cal shapes. No less: interesting are the many problems they offer —
problems of distribution, of habitat, of variability, of reproduction have
frequently drawn attention. In the region considered in this paper,
desmids are one of the most abundantly encountered groups of algae,
and certainly are not devoid of interest.
One of the commonest species, occurring throughout the region, is the
ubiquitous Netrium digitus (Ehrenb.) Itzigs. and Rothe. Attention was
drawn to the numerous observed specimens by the wide range in size:
the smallest specimens were 80-85 » long and about 30 » broad, and a
well graduated series existed which varied from these smallest specimens
12 FARLOWIA, VOL. 1, 1943
up to those that are 300 » and more long and 90-100 » broad. The
cell outlines varied from sub-cylindrical to fusiform, with broadly
rounded to truncate ends.
Closterium occurs in abundance throughout the region. Collections
may frequently be made in which a single species occurs, but more
frequently specimens of this genus are found scattered among other
algae. Zygospores are not infrequently found, developing at all seasons
of the year. One pool gave Cl. rostratum Ehrenb. zygospores in April;
in July, zygospores of Cl. Ralfsii var. hybridum Rabenh. were found;
and in September Cl. setaceum Ehrenb. was conjugating freely. Usually
only a small part of the specimens are conjugating, but occasionally one
finds instances in which large numbers are seemingly forced to conjugate
at the same time. The Closterium macilentum Bréb. mentioned earlier
offers such a case. In this collection many of the zygospores departed
considerably from the spherical shape described by Brébisson for this
species, and were much larger. The example in Fig. 3 is 53x57 w and
rather vaguely angular in shape. The cells contributing to this were
420x 17 » and 460x15 ». The zygospores of Cl. Ralfsii var. hybridum
Rabenh. were spherical and 70-80 » in diameter.
Many interesting problems other than zygospore formation present
themselves here. Sigmoid forms occur very frequently; examination
shows that in some cases the cells which appear as sigmoid are really
of spiral form. In like manner the striae or ribs so characteristic of
many species may depart from the normal arrangement in which they
are parallel to the optical margin and run in spirals around the cell.
More frequently the spiralling is restricted to the apical portion of each
semi-cell. Another feature in the cells of this genus is the gypsum
particles found in the terminal vacuoles. Variation of these granules
is often great, as in a collection of Closterium angustatum Kitz., in
which some cells had vacuoles containing as many as sixteen long slender
granules, others 6 to 10 ovoid particles, others two to six broadly cyl-
indrical granules, and many but a single large nearly spherical one. The
granules in the two ends of a single cell might be quite different. Cells
from this same collection showed many with spiral ribs. Some are
shown in Fig. 5. Many other species have a similar range of variation
in granules. Abnormally shaped cells are not infrequently found in
collections of this genus. Already mentioned are the sigmoid and spiral
forms. Occasionally more abnormal forms appear. The two examples
of Closterium striolatum Ehrenb. shown in Fig. 8 are from a collection
in which many such cells occurred. In some the apical portion was almost
at a right angle to the cell axis; all degrees of bent cells were found
between these and quite normal cells. No cell was found in which
both ends were abnormal. In a few cases cells were found in the later
stages of division, with the younger semi-cells of the two daughter cells
showing the same type of bending.
WHELDEN: New ENGLAND ALGAE. III 13
Specimens of Spinoclosterium curvatum Bernard (Fig. 6) have ap-
peared in several widely separated parts of Maine. The cells vary some-
what, especially in the degree of curvature and the shape and orientation
of the apical spines. The latter are usually straight and from 17-21 p
long, but occasionally curved spines are found. Many specimens showed
terminal vacuoles containing 8-10 small granules. Cells were frequently
found containing specimens of a parasitic fungus (Olpidium sp. ?) so
frequently found in Closterium and other desmids.
Occasionally one finds a collection of desmids which is practically
pure, containing but a single species, in the most unexpected places.
One such was gathered from a shallow sand pit into which a small turf
of Polytrichum moss had slid. No algae were apparent in the clear
water some ten inches deep, yet examination showed the mosses covered
with masses of Euastrum oblongum (Grev.) Ralfs. Most of the cells
were very uniform in appearance, 155-175 » long and 82-90 yp broad.
One specimen merits notice, and is shown in Fig. 2 (right). First ob-
servation indicated that a zygospore was present; examination showed
it to be a cell in which division had gone to the formation of a large
somewhat spherical mass 108 pw in diameter with a coarsely irregular
surface and dense contents of which the nature could not be determined.
The specimen shown in Fig. 2 (left) presents certain troublesome points:
it is 161 » long and 80 » broad; in one semi-cell the upper lateral lobes
are rounded, suggesting EL. humerosum Ralfs; in the other the correspond-
ing lobes are subquadrate with retuse margins and are characteristic of
E. oblongum.
Anomalous cells result from arrested divisions in other desmids as
well as Euastrum. Fig. 10 shows two such cells of Cosmarium bioculatum
Bréb., together with a normal cell. In both abnormal cells shown and
in several other similar ones seen, specimens of a parasitic fungus
occurred.
Cosmarium subturgidum (Turn.) Schmidle is a rare species described
from India. A smaller forma minor has been found in Asia, Africa and
Australia. It is interesting to note that a form should appear in Maine,
where it occurred in considerable numbers in a small pool in a Sphagnum
bog near the coast. The cells were large, from 2.6—3.7 times as long as
broad, distinctly constricted, sinus obtuse, angled; semi-cells ovate with
truncately rounded ends; cell-wall smooth except in apical portion where
many coarse irregularly distributed pores appear. Vertical view circular.
The chloroplastids are parietal longitudinal bands, 7—8 per semi-cell, and
of these 3—4 are visible in face view. The cells are 130-145 » long, and
46-48 », in maximum diameter. The isthmus is 38-44 p» broad. These
cells are distinct from the species and form by their narrower shape
and by the presence of pores at the apex, and so are described as a
new form of this apparently rare species. (Fig. 9.)
Species of Micrasterias are among the commonest of the desmids of
14 FarLowIA, VOL. 1, 1943
this region, twenty-four species having been found, mostly in abundance
and widely distributed. All species show considerable variation in the
nature of the lateral lobing. Anomalous forms seem however to be
relatively rare. Two such cells of Micrasterias rotata (Grev.) Ralfs are
shown in Fig. 15. These occurred in a shallow slow-flowing brook of
clear cold water in which there were thousands of cells of this species.
The few anomalous forms were all of the same nature, represented by
individuals in which one semi-cell had the lobes much distorted and
sometimes nearly at right angles to the main axis of the cell. No evi-
dence of any disturbing factor could be discovered. Another collection
of Micrasterias proved to be of more than passing interest. It was made
from a small pool in a brook flowing very slowly through a swamp.
Few indeed were the cells found on examination of the material—a few
specimens of Closterium, one or two specimens of Staurastrum muticum
Bréb., and a somewhat larger number of Micrasterias specimens. All
the latter were of a single species, but no two were found to be exactly
alike, nor did any cell have its two semi-cells alike. In several cases
one semi-cell was quite definitely Micrasterias pinnatifida (Kiutz.) Ralfs;
in others a semi-cell suggested M. oscitans Ralfs. By far the greater
number of the semi-cells were not easily assigned to any species. Varia-
tions were particularly noticeable in the lateral lobes, which appeared
now broad, now narrow, nearly entire or provided with prominent apical
spines. The variations of the polar lobe were less frequent and often
less pronounced, and included an apical margin which varied from
straight to strongly convex, and with ends merely angular, or with one
tooth or two. Separating the lateral and apical lobes the incision varied
from extremely narrow to very broad. In many cases the two sides
of a semi-cell varied somewhat, though usually they were of the same
general pattern. No noticeable variations could be observed in the
other species of desmids present. Four specimens of this Micrasterias
pinnatifida (Kiitz.) Ralfs are shown in Fig. 16. The cells are 58-69 pu
long, 63-71 y» broad, and the isthmus is 8-12 » broad.
Four species of Xanthidium are here found widely scattered and in
abundance. These are X. armatum (Breb.) Rabenh., X. antilopaeum
(Bréb.) Kiitz., X. cristatum (Bréb.) Kiitz., and X. tetracentrotum Wolle.
Other species such as X. subhastiferum W. West and X. pseudobengalicum
Gronb. also occur, but infrequently. The commoner species hold par-
ticular interest because of the extreme variability which seems to be
exhibited. The specimens of X. tetracentrotum shown in Fig. 14 serve to
illustrate this very well. All these are from a gathering in early August
from a small pond. At the time of collecting, the plants were actively
dividing, filaments of 4-8 cells being found in abundance. All cells
were of normal size for the species, being 38-45 » long, 35-45 p» broad
without spines, and 52-65 p» with them, and 24-26 p» thick. The isthmus
was 10—-12.5 » broad. The cells bore a yellowish brown thickened area
WHELDEN: New Encuianp Atcae. III tis
in the middle of the lateral face; the scrobiculations thereon sometimes
formed a semicircle or crescent, but more frequently a complete circle.
Within the circle a few accessory scrobiculations often appeared. Great-
est irregularity attached to the spines, both in their number and their
disposition. The smallest number found was in those cells in which
each lateral angle bore a single spine. Other cells bore a pair of spines
in each angle, or eight spines for the entire cell. All variations between
these were found, the paired spines being in a horizontal plane. One
of these is shown in vertical view in Fig. 14, the semi-cell having two
spines on one angle and one on the other. The other semi-cell has un-
paired spines. In many specimens there were two spines to each angle,
but they were in a vertical plane. Often unpaired spines were found
above these, the cell shape then becoming subhexagonal instead of ellipti-
cal. Many cases were observed in which a prominent papilla appeared
instead of a spine. Some specimens presented a decidedly asymmetrical
appearance as a result of the very irregular disposition of the spines.
All the variations or irregularities noted were cells which could be con-
sidered mature, the central ornament being deeply colored or the cell
actively dividing.
Xanthidium antilopaeum (Bréb.) Kiitz. is much more widely distrib-
uted in this region than is X. tetracentrotum Wolle, several distinct varie-
ties being found. One of these is Irénée’s var. quebecense. Many speci-
mens of the latter are much larger than his, the largest measured being:
100 » long (with spines), 92 » (without spines), 106 » wide (with
spines), 90 » (without spines), and the isthmus 16—23 » broad. The
central scrobiculations varied greatly in different specimens, but most
frequently these were arranged in a broad ellipse within which many
others formed a second ellipse or were most irregularly disposed. Sev-
eral zygospores were found in the same collection, but none with empty
semi-cells attached. They were spherical and about 60 » in diameter,
without spines. The surface bore many simple straight spines 11-15 p»
long. It seems quite possible that these are zygospores of the numerous
var. quebecense cells present.
A pool of stagnant water in a large granitic ledge contained large
numbers of another form of Xanthidium antilopaeum. The many cells
examined are of very uniform appearance and represent a distinct
variety. They are of medium size, about one and one-quarter times
longer than broad, sinus at first linear, then divergent to a rounded
basal angle. Apex usually straight, less frequently slightly convex, in-
frequently faintly retuse, the lateral margins straight or slightly convex.
Angles rounded, each furnished with a pair of stout upwardly curved
(rarely straight) spines frequently yellowish-colored. In a few instances
a third spine may exist between or lateral to one or both pairs of basal
spines. Central thickened area brownish-yellow and circular or ellipti-
cal, and furnished with numerous scrobiculations which are variously
16 FarLowliA, VoL. 1, 1943
arranged. Sometimes these are very faintly visible or even completely
lacking. Vertical view elliptical with truncated ends, each bearing a
pair of parallel or slightly converging spines. Side view of semi-cell
subcircular. Length 74-80 »; breadth (with spines) 87-95 p, (without
spines) 60-68 y; thickness 45-48 y», breadth of isthmus 24-36 »; length
of spines 13-16 ». Distinct from other species and other varieties in
relative dimensions, in its straight apical margin, and the disposition
of the spines. This plant seems to have been first described in 1888 by
W. H. Hastings, presumably in his local newspaper, as Xanthidium
antilopaeum var. truncatum. Soon after W. West described the same
plant as X. Tylerianum, collected from Maine. About two years later
Hastings raised his variety to species rank, citing at the same time West’s
X. Tylerianum (misquoted as X. Taylorianum) as synonymous. It would
seem better to keep this desmid as a variety of X. antilopaeum; therefore it
becomes Xanthidium antilopaeum var. Tylerianum (W. West) Whel-
den comb. nov. (Fig. 17).
Arthrodesmus quadridens Wood is one which has frequently appeared
in collections from Maine and has always attracted considerable attention,
primarily because it seemed so different from all other species of the genus.
Recently Prescott and Scott, with good reasons, have removed this alga
from that genus, and established for it a new genus, Spinocosmarium.
They note that the species S. quadridens shows “a great variation in the
degree to which granules and processes are developed”. This is indeed
so. However, collections vary greatly in this respect. Frequently one
may examine scores of specimens in a single collection and find no
obvious variation. In another collection from a habitat closely re-
sembling the first, every specimen seems distinct in some way. The
most conspicuous variation occurs in the spines which are so striking
a feature of the lateral margin. These may be simple or once-furcate;
between cells in which all spines are simple and those in which all are
furcate, every conceivable combination of simple and furcate spines
occurs. In the furcate spines the two forks are usually nearly equal in
size, but instances do occur in which one fork may be reduced sometimes
to a degree where it becomes merely a minute tooth on one side of the
spine. Infrequently spines are borne at the apical angles and are usually
much smaller than the lateral ones. Generally the spines occur singly,
although occasionally two may be produced on an angle. All specimens
I have examined are alike in having a single large rectangular tubercle
just above the isthmus. The many other tubercles vary greatly both in
prominence and in number, especially in the lateral parts of the cells.
In rare cases coarse pores occur in abundance and irregularly distributed
in the wall between the tubercles. Generally this wall appears smooth.
The cells were 33-42 » long, 51-57 » broad (with spines) and 33-40 p»
without; the isthmus was 9-13 » broad; the cells were 24-26 w thick.
Three species of Staurastrum merit notice because of their zygospores.
WHELDEN: NEw EncuLanp ALGaeE. III 17
On three different occasions zygospores of St. punctulatum Bréb. have
been found in abundance, once in mid-May and twice in late summer.
The most noticeable feature of these zygospores is the broad range in
size, from 36 p» to 62 p» in diameter, the processes included. (Fig. 11.)
In several shallow pools Staurastrum Rotula Nordst. (Fig. 12) is a
species encountered frequently and generally in large numbers through-
out the warmer months, becoming particularly common in August and
September. During the entire period the rather large cells show a
certain amount of variation; the number of processes may be either eight
or nine. These rays usually are quite straight, those of the two semi-
cells being parallel. Not infrequently one finds specimens in which the
processes are curved outward. The margins of the processes are in
most instances coarsely crenulate, but occasional specimens are found
with perfectly smooth processes. The verrucae which are so character-
istic a feature of this species also vary, being usually rather large and
bidentate, in other cases small and conical, and occasionally completely
lacking. Some of the specimens have larger dimensions than those usually
given, being 42-48 » long, and up to 124 w broad (processes included).
Zygospores of this alga were found in very large numbers during
August and early September of 1929. They were spherical bodies 36-43 p.
in diameter, with many straight or slightly curved processes 28-32 yp
long. In some zygospores the apices of all processes were armed with
two or rarely three curved teeth; in other zygospores the ends of all
processes were simple and rounded; in some, processes of each type
occurred, The body of the zygospore was deep brown to black in color.
Fig. 12 shows three zygospores with different types of processes, and
two cells, one with crenulate and the other with smooth processes. In-
frequently zygospores of this species were found infected with a small
undetermined fungus. In these the processes were generally much
shortened (10-15 » long), coarser than normal, or even unciform. Such
infected zygospores were easily recognized as abnormal, even when the
parasitic body was not distinctly visible.
Staurastrum Johnsonit var. depauperatum G. M. Smith (Fig. 13) oc-
curs quite generally in this region, sometimes in abundance, together
with the species. Occasional specimens of zygospores of the present
species were found. They were spherical bodies of brownish color,
28-32 » in diameter and provided with 12-15 simple straight processes
16-20 yw» long.
Disregarding purely taxonomic problems and those dealing with the
life-histories of algae, interest in this group of plants has centered around
three main questions, their distribution in both a local and a broader
sense, the variations that they exhibit and possible explanations of them,
and the formation of sexual stages, with some attention being given to
the periodicity of their existence.
The appearance of sexual stages and especially of zygospores in the
18 Fartowia, VoL. 1, 1943
desmids, is often a rare occasion. To record their occurrence, therefore,
seems ever desirable, the more so when notes on any unusual circum-
stances attend this; with the accumulation of such cases an understanding
of some of the causes leading thereto may be had. Certainly my own
observations on zygospores, far more numerous than those recorded
above, lend no support to any statement that any one season favors
their appearance. Frequently one finds a collection in which many
different species of the algae present are fruiting freely, thus suggesting
that this event is a consequence of external factors. The collection in
which the zygospores of Staurastrum Rotula were so abundant is of this
sort, for in this ten different algae, including eight desmids, were forming
zygospores. In other cases, large numbers of algae may be present,
and only one fruiting. Collections in which a dozen species of Closterium
are abundant may show only one forming zygospores. It might then
be suggested that an internal factor might be operating.
The question of occurrence, or favorable habitats, may be briefly
mentioned. Everyone who has given serious attention to collecting algae
knows that there are problems involved, in deciding just where collecting
may be best. Nor can any rule be given for this. A collection may be
very poor in numbers and quantity, yet show rich variations in the
material that is present. Apparently identical habitats may show entirely
different results, one spot yielding abundant material while one nearby
is almost barren. Variations of this kind may be found in two collec-
tions made in a monotonously uniform habitat and only a foot apart.
It would seem that oftentimes the algae maintain a remarkably fine
adjustment to some factor or factors in their environment or within
themselves. All too frequently algae, especially desmids, may disappear
from some place with startling abruptness, not a trace remaining in a
habitat which twenty-four hours before gave very rich collections.
The question of variations in algae and the causes of them has at-
tracted considerable attention. Jacobsen (1875) was one of the first to
consider the nature and causes of variations so frequently observed in
desmids. He advanced several possible explanations and noted the wide
geographical range of the variations. De Wildeman (1887) paid at-
tention to this question, brought together may cases of variations noted
by previous workers and concluded that . . . “ce ne sera que par une
étude approfondie faite sur de nombreux matériaux de diverses prov-
enances que l’on pourra arriver 4 donner, a chaque espéce, la place qui
lui convient dans la série et 4 supprimer toutes les espéces dont la creation
est basée sur der caractéres secondaires.” Later de Wildeman (1895)
noted the frequency of occurrence of variations in the genera Euastrum
and Micrasterias. Cushman (1907) found the species of Micrasterias
to be very variable, and remarked that “as many of these so called
varieties are often seen forming one semicell of a specimen, the other
semicell of which is typical, it seems hardly wise to recognize them as
WHELDEN: New Encianp Atcag. III 19
good varieties.” Johnson (1894), Stange (1908), Playfair (1910),
Ducellier (1914, 1915, 1917, 1918), Woronichin (1926) and Huber-
Pestalozzi (1931) are among the many who haye given considerable
attention to the occurrence and extent of variations under natural condi-
tions, and offered various explanations as to their causes. More recently
Lefevre (1939) has studied the problem both in nature and more especially
in cultures. His extensive experiments on several genera and species of
desmids lead him to conclude that anomalies and monstrosities appear
much more frequently in cultures than in natural habitats. He is led to
state that “Observations based on material grown in culture can never
replace material observed in nature, but will serve only to supplement
these and may give proof to matters otherwise left very doubtful.”
The notes on variations recorded above are based on materials from
natural environment. They lead to certain observations. Monstrosities
such as those in Euastrum and Cosmarium do occur infrequently and in
many cases at least seem to be accompanied by, if not indeed a con-
sequence of, attack by a parasitic organism. It is quite possible that
in such abnormal forms as those of Micrasterias (Fig. 15) a parasitic
organism is present, but in a stage the presence of which is not yet ap-
parent. For certain it is that this genus is frequently infested with
parasitic organisms. Possibly such abnormal forms occur only if the
parasite enters just before division of the host begins, or in its early stages.
It has frequently been stated that the many variations in size, form,
ornamentation, and cell contents occur much more frequently in certain
genera. Undoubtedly this is true. It is also a fact that in many cases
the very nature of the cell makes it difficult to observe variations — small
smooth species of Cosmarium cannot offer as much in the way of variety
as ornate ones. Where obvious variations do occur it is seldom possible
to ascribe to them definite causes. Many factors outside the cell come
to mind. Abrupt changes in temperature, and large fluctuations in the
amount of substances dissolved in the water have been frequently
mentioned. Yet when one observes what happens in two small nearby
pools, or examines two collections from points near one another in a
single pool, where great variation appears in some of the species in one
and as great uniformity in the species in the other collection, one con-
cludes that such a factor cannot always operate. Nor do they seem
adequate to explain all observations. Many things indicate that there
is within the organisms themselves something which may lead to the
appearance of variations, and that. this together with other external
factors operate together perhaps in very subtle ways to bring about
the morphological changes which one observes. Only through observa-
tions accumulated slowly, and carefully supplemented whenever possible
with observations on controlled cultures, can a solution of these prob-
lems of algal behavior be hoped for.
Harvarp UNIVERSITY
CamBripcE, Mass.
20 Fartowia, VoL. Ll, 1943
LITERATURE CITED
Cushman, J. A. Ya, vin Sip
<— . cs
A
mettre
Ms
NW
ANN
£5
on
=
y:
PLatE 2.
74, . FarLow1A, VoL. 1, 1943
EXPLANATION OF PLATE 3
All drawings are made with the aid of a camera lucida. Each unit of the scale
equals 10 yp.
A. Martensella Corticii Thaxter. This species is set apart from others by the short
conidiophores that are produced on the surface of Corticium spp., and by the
fact that the phialides are produced on the upper surface of the sporocladia.
B. Coemansia erecta Bainier. The conidiophore illustrated here is rather atypical
since it is branched below the fertile zone and the sporocladia are somewhat
more closely aggregated than is usual. A typical conidiophore is shown in
Plate 2 A.
C. Coemansia ceylonensis Linder. This species, the smallest of the genus, is also
characterized by the small number of few-septate sporocladia.
D. Coemansia scorpioidea Linder. This species is readily distinguished by the
scorpioid arrangement of the sporocladia at the apex of the conidiophore as
well as by the elongate, 1-2 celled sporocladial stipes.
E. Coemansia spiralis Eidam, showing the spirally twisted upper end of the con-
idiophore.
F. Coemansia guatemalensis Thaxter. The slender conidiophores bearing the sporo-
cladial stipes at nearly right angles to the main axis, the sharply upcurved
basal cell of the sporocladium, and the dimensions of the spores all help to
separate this from other species of Coemansia.
75
LINDER: KICKXELLACEAE
PLATE 3.
76 FarLowlia, VOL. 1, 1943
EXPLANATION OF PLATE 4
All drawings were made with the aid of a camera lucida. Each unit of the scale
equals 10 y.
A. Coemansia ceylonensis Linder.
B. Coemansia scorpioidea Linder. A sporocladium from which most all spores have
fallen,
Coemansia erecta Bainier.
Coemansia guatemalensis Thaxter.
Coemansia Thaxteri Linder.
Coemansia kamerunensis Thaxter.
Martensella Corticii Thaxter. Immature sporocladia.
TOmAmMoO
Coemansia spiralis Eidam. An immature sporocladium showing the phialides
which are at first cylindrical.
I-J. Coemansia aciculifera Linder. Immature zygospore from sphagnum is shown
in figure I, while a discharged sporocladium and a single conidium, pointed at
both ends is illustrated in figure J.
K. Coemansia breviramosa Linder. An immature sporocladium, and three conidia.
L. Coemansia interrupta Linder. A mature, discharged sporocladium and 2 conidia.
M-N. Coemansia braziliensis Thaxter. A mature sporocladium with spores (M) and
immature sporocladia (N).
O. Coemansia reversa Van Tieghem & Le Monnier, showing sporocladia in differ-
ent early stages of development.
P. Kickxella alabastrina Coemans. A single sporocladium at an early stage of
development in which the phialides have produced sterigmata on which conidia
will be produced. Above the sporocladium are shown two conidia, of which
the left hand one has been drawn in optical section to demonstrate the annular
ring that has been laid down inside the conidial wall.
77
LINDER: KICKXELLACEAE
PLATE 4.
1(1) : 79-93 FARLOWIA January, 1943
BASIDIUM FORMATION AND SPORE DISCHARGE
IN GYMNOSPORANGIUM NIDUS-AVIS
ALTON E. PRINCE 2
During the course of life history studies of Gymnosporangium nidus-
avis, an extensive investigation was conducted on the process of teliospore
germination from basidium formation to discharge of the basidiospores.
For the necessary observations a special technique, described below, had
to be perfected. The general method employed for observing the various
stages of germination under the oil immersion lens is not new, but ap-
parently the adaptation devised here has not been employed previously
for this purpose. Shallow Van Tieghem cells 4 to 5 mm. high were
used to reduce the difficulties ordinarily encountered when attempting
. to obtain proper light. A small, moist mass of teliospores was placed
on a thin cover glass and spread out as much as possible; then another
cover slip, small enough to go inside of the cell was pressed onto the
telial mass, and the whole then inverted over the cell which contained
a small drop of distilled water. The glass slip served five purposes:
(1) It permitted the use of any number of mounts which could be kept
under ordinary laboratory conditions. (2) No regulation of the tem-
perature was necessary to prevent the water from condensing on the
under side of the cover glass. (3) The water of condensation formed
_ on the under side of the glass slip where it did not interfere with the
germination of the spores nor with the observations. (4) The teliospores
were kept in a thin layer under conditions of high humidity where they
could germinate and produce basidiospores normally and where they
could be observed readily under the oil immersion lens. (5) The difh-
culty described by Buller (1924, p.524) of getting the teliospores to germi-
nate and produce basidiospores normally was thus overcome. |
The mode of teliospore germination in G. nidus-avis does not differ
from other pucciniaceous rusts that have been investigated by various
workers. However, since it is believed that a more detailed study has
been made than previously, the facts found will be presented. In this
discussion the terminology of Linder (1940) has been adopted.
As Farlow (1880) and Thaxter (1891) reported, the thin-walled telio-
spores may germinate at both ‘ends after becoming separated from their
pedicels. When this takes place, the basidium of the lower cell passes
through the locus where the passage from the pedicel into the spore had
been during the growing period of the spore. Usually only one basidium
1 Contribution from the Laboratories of Cryptogamic Botany and the Farlow Her-
barium, Harvard University, No. 217.
“Tt is with pleasure that the author acknowledges the guidance and constructive
criticism of Dr. D. H. Linder and Dr. J. H. Faull during the entire progress of this
investigation and to Dr. W. H. Weston, Jr., for suggestions and criticism concerning
this manuscript.
79
80 Fartowia, VoL. 1, 1943
is produced by each cell of a spore, but occasionally more may start
to form. However, only one develops completely.
There are two types of basidia formed when teliospores germinate,
depending upon the environmental conditions. The first type of basid-
ium, which is considered normal, is characteristically formed by telio-
spores which are near the surface of the telium. In these, the basidia
are variable in length, uniform in diameter, sessile (Pl. 1, B), and
subsequently become divided into four cells, each cell of which normally
gives rise to a sterigma bearing a basidiospore. Occasionally normal
basidia are produced by spores deeply embedded in the telium, but when
this occurs the four cells of the basidium become separated at their
septa, a phenomenon often described by other investigators. The indi-
vidual cells, when allowed to germinate in water, instead of producing
basidiospores, proceed to germinate by germ tubes which are capable
of penetrating the host and causing infection. This, however, would ,
rarely if ever occur except in artificial inoculations since the individual
cells could not be dislodged from the gelatinous matrix of the telium.
Therefore, it can be concluded that basidiospores in nature constitute
the only structure for the dissemination of the rust from its telial hosts.
The second type of basidium is produced from teliospores deeply em-
bedded in the telial matrix when, as noted by Linder (1940), there is
formed a basidial stipe, that is a fifth cell of the basidium, and which
_may be 100 » or longer, the length depending upon the distance of the
teliospore from the air. The diameter of the stipe is usually about
one-third of that of the spore-bearing portion of the basidium, or if the
stipe is short it may be of the same diameter as the basidium. When
teliospores are deeply embedded, the basidial stipe enlarges only as it
approaches the surface or after it is out of the telial matrix (Pl. 1, A),
but even after this the spore-producing portion of the basidium may
sometimes become covered with water, after which the sterigmata usually
grow long enough to form their spores in the air.
During the active period of germination, the young basidium has a
colorless thin wall and is filled with a yellowish-orange protoplasm.
When a basidial stipe is produced, its appearance at first is very much
like that of a non-septate vegetative hypha and as the stipe elongates
only the outer portion remains filled with the colored protoplasm. The
portion next to the teliospore becomes hyaline, vacuolate, with occasional
globules of the yellowish-orange cell contents. Finally, when the basidial
stipe reaches the surface of the matrix, the enlarged portion of the
basidium is formed (Pl. 1, A), after which most of the colored proto-
plasm passes into the outer enlarged portion and is then cut off by a
septum at its base. The basidial stipe remains turgid to the extent that
it may support the whole enlarged end until after the basidiospores have
been discharged. From this stage the formation of sterigmata and
basidiospores is the same as when there is no basidial stipe. A wall
PRINCE: BASIDIOSPORE DISCHARGE Sl
first divides the basidium into two nearly equal parts and then the two
halves are likewise divided by a septum in each, resulting in the forma-
tion of the four basidial cells. A sterigma soon begins to form on one
of the cells or more than one sterigma may be produced simultaneously
on two or more cells, but there is no definite order for their formation.
Once a basidium becomes divided into its constituent parts, each segment
appears to be a separate unit more or less independent of the other
cells of the basidium.
The presence of a continuous wall around the basidium permits a
comparison of a basidium with a four-spored ascus, the outer continuous
wall being comparable to the wall of an ascus and the four cells, each
with its wall, to the ascospores. The correlation of this fact with the
cytological findings of Olive (1942) who established the presence of
astral rays during nuclear divisions in Coleosporium is additional evi-
dence for the theory proposed by Linder ce) that rusts originated
from an ascomycetous ancestor.
The sterigma starts as a protuberance from the surface of a basidial
cell (Pl. 1, B). Usually all four sterigmata arise from the same side
of the basidium, but the forms of the sterigmata are variable and may
be long or short as shown in Pl. 1, G and K, respectively. Long
sterigmata are often twisted or variously contorted. By the time a
sterigma has attained its maximum length which may be 50 » or more,
depending upon the distance of the basidium from the air, the sterigma
has a more or less tapered distal end from which a basidiospore begins
to form almost immediately. The spore at first appears as a globose en-
largement of the apex (PI. 1, C), then it grows more on one side and
becomes sub-ovoid (Pl. 1, D-E) and subsequently the growth rate of the
spore is increased greatly. During this period the yellow-orange cyto-
plasm can be seen rushing from the basidial cell through the sterigma
and into the spore where there is a microscopic maelstrom of whirling
cytoplasm. The flow of the colored protoplasm into the rapidly forming
spore is restricted to a central channel in the sterigma (PI. 1, E), there
being little or no movement of the hyaline, parietal protoplasm. The
opening from the sterigma into the basidiospore being approximately
1.5 » in diameter is large enough to permit the rapid passage of the
cell content. However, during the course of the migration of the cell
content, it was observed occasionally that the flowing might cease for
an instant, then a reverse flow take place, and after a moment, the out-
ward flowing into the spore is resumed.
The spore continues its growth until all, or nearly all, of the colored
protoplasm has passed from the basidial cell into the spore (Pl. 1, F
' and L), and by that time the spore has reached its maximum size. Very
soon afterwards a septum (PI. 1, G-H) is formed a short distance below
the spore in the sterigma, thus forming the apiculus, and following this
the basidial cell and the sterigma remain turgid and filled with the
*
82 Fartowia, VoL. 1, 1943
hyaline, vacuolate periplasm, some of which soon appears to become
much concentrated in the apex of the sterigma (Pl. 1, K). At the
same time similar appearing material which can be differentiated from
the yellowish and more dense content of the spore, can be seen in the
newly formed apiculus (Pl. 1, H).
The nature of the hyaline material is unknown, but that it plays an
important part in the discharge of the spore is almost certain. It is more
than possible that this substance is like the periplasm found in the ascus
where it is known to have a marked affinity for water, and in this way
a pressure is built up, thus bringing about the discharge of the ascospores.
This hyaline material may have a similar function in the sterigmata
and apiculi of the basidiospores of Gymnosporangium nidus-avis, that is,
it may take on water with the production of a pressure in both of these
extremities and result in the violent discharge of the basidiospores.
That the material may act as a plug to prevent the collapse of the sterig-
mata and spores after the discharge is not probable because there is a
definite wall over the apices of both the sterigmata and the apiculi and
therefore no plug is necessary. The septum between the spore and
sterigma has a middle lamella and as soon as the septum is formed the
spore seems to pass through a very brief maturation period which may
have a duration of a few seconds to a few minutes. About ten to twenty
seconds before the spore is discharged there is produced, just above the
apiculus, on the flattened edge of the spore, a droplet (PI. 1, I-J) that
appears to come from a hilum of some kind, but a diligent search has
failed to reveal any opening. However, at the place where usually one
large drop is given off, but where sometimes several small ones are
emitted and nearly always, whether it is only one or whether there are
several, they are usually given off from a fraction to about ten seconds
before the spore is discharged. The droplets were seen a few times to
flow down the sterigma prior to discharge of the spore, a fact reported
previously by Buller (1922) as occurring in abnormal cases, and in
addition the single drops occasionally are carried away on their spores.
After the droplets have been produced, and at the moment before dis-
charge, the spores vibrate as if under tension, but after the discharge
a definite wall is evident over the end of the sterigma and over the end
of the apiculus (Pl. 1, M, a), both the walls being a little less than
a micron thick, are distinctly convex showing that they are under pressure,
but the apiculus soon nearly disappears whereas the apex of the sterigma
remains rounded until the basidial cell bearing it subsequently collapses.
The actual discharge occurs so quickly it is not possible to see exactly
what happens, but from observations and photographs (Pl. 1) made
just before and after the discharge, and from the study of germinated
teliospores which have been fixed in 4% formalin and stained in
0.1% aqueous cotton blue it is possible to offer a theory that seems to
be a plausible explanation of this phenomenon. In a consideration of
PRINCE: BASIDIOSPORE DISCHARGE 83
this theory, it will be recalled from the preceding description that at
the earliest stage in the formation of the basidiospore, the apex of the
sterigma becomes enlarged to form a small sphere (Fig. 1, A and B),
and then grows more on one side than on the other so that as the spore
rapidly enlarges it becomes more or less reniform (Fig. 1, C).
SIAN Teg
Ficure 1.—Semidiagrammatic representation of basidiospore development and dis-
charge. The heavy outer line represents the primary wall common to the basidium,
sterigma and spore. The secondary wall is shown by the dotted areas within the
outer wall. The broken line at (a) represents the wall plate that apparently hydro-
lyzes and may exert a pressure which assists in spore discharge. For further explana-
tion see text.
When the spore reaches its mature size (Fig. 1, D), there is formed
a short distance from the spore in the sterigma a wall plate (Fig. 1, E)
and this closes the opening between the sterigma and the spore, pro-
ducing the apiculus. The formation of the disc-shaped wall plate is
followed by a wall that is laid down inside the primary wall and above
the disc by the spore, and below the disc by the sterigma (Fig. 1, F),
the result being that the wall plate becomes a middle lamella bordered
peripherally by the very tenuous original wall (Fig. 1, G) that is com-
mon both to the developing spore and to the sterigma. At this stage
in the development, the spore, sterigma, and basidium have a very
tenuous outer, primary wall with a thicker secondary wall on the inside.
When pressures are built up in the apiculus of the basidiospore and
in the apex of the sterigma by the periplasm, the original wall acts
84 FarLowia, VoL. 1, 1943
as a retaining membrane until the pressures become sufliciently great
to rupture it suddenly, thus permitting the instantaneous bulging of
the end wall of the sterigma and of the apiculus, and resulting in the
forceful discharge of the spore (Fig. 1, I). It is possible also that
just prior to spore discharge there is either a partial or complete hy-
drolysis of the middle lamella, (Fig. 1, H, a), and this would serve not
only as a structure for disjunction, but might even exert some pressure
against both the spore and sterigma.
A comparison of the method outlined above with the method of
spore discharge in Entomophthora (Thaxter, 1888) and Sclerospora
(Weston, 1923) leads one to believe that the discharge phenomenon is
essentially the same whenever it occurs in fungi. The reasons for mak-
ing such a comparison will be given in paragraphs to follow.
This discussion would not be complete without a consideration of the
droplet that is given off from the basidiospore. It is difficult to under-
stand or explain its existence, but it may be supposed that the emission
of the droplet is a means by which the pressures that must exist in the
spore and sterigma are equalized so that at the moment of discharge
the expanding surfaces of the sterigma and apiculus may bulge instan-
taneously with equal force.
That the droplets play a mechanical part (Ingold, 1939) in the dis-
charge of the spores is doubtful because nearly mature teliospores can
be placed in water and the spores will still be discharged in the normal
way. Even the droplet or droplets can be seen as they are given off
and this is interesting for another reason since it indicates, as has been
supposed, that the droplets contain certain substances in addition to water.
The facts ascertained by the writer tend to substantiate, in part, and
extend those recorded by previous investigators-among whom Brefeld
(1877) was apparently the first to see basidiospores of a Hymenomycete
discharged violently from their sterigmata, but he thought all the spores
were discharged at once as a result of a bursting of the ends of the
sterigmata, and that water drops were left on the apices of the sterigmata
after the spores had gone. However, Schmitz (1843) was the first to see
basidiospores of a Hymenomycete leave their sterigmata, but he did
not realize they were discharged since he definitely stated (p. 434)
that the spores fall from their sterigmata one at a time.
Subsequently Fayod (1889) in discussing this same problem mentions
that Zalewski (1883) confirmed Brefeld’s observations but that he,
Fayod, could not agree that the ends of the sterigmata were open after
the discharge of the basidiospores. Instead he observed that there was
formed at a hilum located at the septum, which separated the spore from
the sterigma, a droplet which grew to a certain size. This droplet, he
claimed, hydrolyzed the middle lamella and was followed by a sudden
distension of the end of the sterigma, a result of the sudden expansion
of the basidial content as indicated by the rounded end of the sterigma.
PRINCE: BASIDIOSPORE DISCHARGE 85
He also saw that the spores of a single basidium were not discharged
simultaneously as suggested by Brefeld (1877) but that they left their
sterigmata one at a time, and that the basidia remained turgid after
their spores were discharged.
The first to publish that basidiospores of rusts were forcibly discharged
was Klebahn (1904) who observed that, when fresh telial sori of Gym-
nos porangium were placed in conditions favorable for teliospore germina-
tion, basidiospores could be seen a few millimeters away from the sori;
thus he concluded that the sterigmata had the power to discharge their
spores and this experiment was confirmed by Falck (1904), and was later
repeatedly verified by Weimer (1917), Giddings (1918) and Buller
(1924).
Massee (1906) observed that in Hymenomycetes the mature basidio-
spore was cut off from the apex of the sterigma by a transverse septum
and that the parietal protoplasm remained in the sterigma. In addition
he stated that, when a spore was discharged, the force came from an
accumulation of water in the basidium which had an elastic wall and,
when the tension became great enough, the wall over the end of the
sterigma ruptured “in a circular manner just below the septum at its
apex, the elastic wall of the sterigma instantly contracts and forces its
contained water to strike the apical transverse wall, which is thrown
off along with the spore seated on it.” The present investigations show,
not that the end of the sterigma ruptures in a circular manner to leave
a pore when the spore is shot off, but that a septum is formed in the
sterigma, bordered on both surfaces by the wall of the apiculus and
sterigma, thus preventing the extrusion of water and making it impossible
for the septum to be shot off with the spore.
Dietel (1912) stated that the studies of Klebahn (1904) were the most
complete and accurate up to that time. He made further observations
on the discharge of basidiospores in different species of rusts. In the
case of a species of Coleosporium he reported that at the base of mature
basidiospores there appeared a droplet which enlarged rapidly until at
the end of 40 seconds it had reached a diameter of 9-10 ». This caused
the spores to be pushed a little to one side and apparently brought about
the rupture of the spores from their sterigmata because these basidio-
spores were observed to have suddenly discharged and carried their
droplets with them. In addition, Dietel found that there was a difference
in the time necessary for the production of basidiospores between species
in Puccinia and Coleosporium, the time from teliospore germination to
basidiospore discharge in the former was 244 to 3 hours while that for
Coleosporium was 34% to 4 hours, but in both genera a half hour was
all that was necessary for the formation and discharge of the basidio-
spores. The writer’s investigations in Gyminosporangium nidus-avis
confirm the amount of time it takes for the teliospores to germinate and
produce basidiospores although it was found that germination could be
86 Fartowia, Vou. 1, 1943
controlled to a great degree by temperature. Thus 8 to 10 hours were
necessary for germination at 5° C., whereas a minimum of only 2 hours
were necessary at a room temperature of about 24° C., but above this
temperature the heat definitely retarded germination. As for the ob-
servations of Dietel on the time it takes for the water drop to form, the
size and function of the droplet that is exuded from the basidiospores,
the author can only state that this is not in agreement with what is true
in G. nidus-avis, but there is no reason why the size of the drop and the
time necessary for its formation should not vary widely, depending upon
the fungus that is being studied. In G. nidus-avis, the single droplets
form in about 5 to 15 seconds, depending upon their size and never did
a droplet reach a diameter that equaled the width of the spore or more
than 8 » in diameter. The droplet shown in Plate 1, I is 5 » while the
one in Plate 1, J is approximately 3 p.
Coons (1912) in addition to presenting a nearly complete historical
resumé of the facts concerning spore discharge, demonstrated that ether
and chloroform inhibited basidiospore discharge in Gymnosporangium
juniperi-virginianae, and therefore concluded that the actual discharge
must be dependent upon protoplasmic activity unless the vapors of chloro-
form and ether affected the turgor relations within the spores and basidial
cells. Furthermore, Coons verified the fact that basidiospores are forcibly
discharged from their sterigmata following the formation of a droplet
which is excreted and that the actual discharge is the result of a rapid
bulging of the two flat-end walls of the apiculus and sterigma. The
presence of a cross-wall in the sterigma was again confirmed by Hiley
(1919) during his investigations of Fomes annosus.
The “Researches” of Buller contain most of our present knowledge
concerning basidiospore formation and discharge. Buller (1909) postu-
lated that basidiospore discharge in the Hymenomycetes was comparable
to but distinct from conidial discharge in the phycomycetous Empusa;
however, he was unable to establish this because he could not prove con-
clusively that there was a double wall between the spore and the sterigma,
and several years later he (1922, pp. 22-24) abandoned this theory
on the grounds that the septum between the spore and the sterigma could
not be seen and furthermore following certain experiments stated that a
septum was not necessary for the process of discharge. Earlier, how-
ever, Buller (1915) was successful in establishing the observation of
Fayod that a droplet was formed at the hilum of a basidiospore just
prior to the discharge. Buller (1924) in a comparison of the basidia in
Hymenomycetes and Uredinales stated that in rusts (1) at the beginning
of spore development, the basidiospores first form a projection, the
hilum, at one side of the point where the spore joins the sterigma;
(2) the axis of the spore is always inclined to the axis of the sterigma;
(3) the four spores are discharged from their sterigmata in succession,
not all at once; (4) the spores are shot a distance of 0.4—0.85 mm.;
PRINCE: BASIDIOSPORE DISCHARGE 87
(5) a droplet is formed at the hilum 10-40, often 15, seconds before
the spore is discharged; (6) after the droplet has reached a certain size
it is discharged with the spore; (7) the basidia or sterigmata do not
collapse at once after the discharge, neither do the sterigmata bear water
drops. My own observations substantiated by photographs shown in
Plate 1, do not confirm the first two statements of Buller that the hilum
is the first structure of the spore to be formed, or that the axis of the
basidiospores is always inclined to the axis of the sterigma, but on the
other hand there is no reason to believe that what is true in Gymnospor-
angium nidus-avis is true for all the rusts. There is neither a distinct
structure on the basidiospores of G. nidus-avis that can be called a hilum
in the sense that Buller has used the term, nor is the axis of the basidio-
spores always inclined with the axis of their sterigmata, but the two axes
often seem to be parallel except in an intermediate stage of growth. The
third point that the spores are not discharged all at once but in succession
confirms the original observations of Fayod (1889) in the Hymenomy-
cetes and of Dietel (1912) in the Uredinales, and was again shown to
be true during the progress of the present investigation. Concerning
the fourth point, the distance the spores are discharged, Dietel (1912)
reported that spores may be discharged 0.3 mm. vertically and up to
0.85 mm. horizontally, with the approximate velocity of 8 cm./sec. The
writer found that, if a greased cover glass was placed about 0.4 mm.
above a telial sorus that was discharging spores, a few would stick to
the glass, but at 0.5 mm. or higher no adhering spores could be found, —
although, if the glass cover was fixed so that it was barely above the
telium, many spores could be found on the under side. The last three
of Buller’s points confirm the important facts described by Fayod (1889)
in the Hymenomycetes and Dietel (1912) in the Uredinales, and it
should be stated that with the exceptions already given the writer has
again confirmed these facts. In G. nidus-avis the water droplet is ex-
creted from 5 to 15 seconds before the basidiospores are discharged,
the droplet may appear as a single large one that may reach a size of
7 p but is usually less than 5 p» in diameter, or several small droplets
may be given off in rapid succession, but only occasionally do the droplets
remain on the spore until the latter is discharged. There appears to
be no definite size for the droplet to reach before discharge takes place
in the species under consideration and, as has been previously men-
tioned, the basidia and sterigmata remain turgid for some time after
they have lost their spores.
In reference to the actual mechanism of spore discharge, previous
theories with the exception of the one presented by Fayod (1889) are
skillfully summarized by Ingold (1939, pp. 81-86) who supports the
suggestion of Buller (1922) that the discharge may be caused by the
force of surface tension found on the surface of the exuded droplet, but
after proving by mathematical formulae that there is more than enough
88 ; Fartowia, Vou. 1, 1943
energy stored on the surface of the droplet to account for the. initial
velocity of the spore, he has concluded with the statement that “it is not
easy to see how this energy can be mobilized to bring about discharge.”
The most recent consideration of the problems in basidiospore dis-
charge is given by Lohwag (1941) who states that there is no evidence
of a septum between the basidiospore and sterigma, a statement that
seems to confirm the observations of Buller (1922). In addition, Lohwag
presents a theory different from any Buller or Ingold have postulated.
Lohwag believes that, as in the discharge of spores from the ascus, the
force that discharges the basidiospores originates from a turgid basidium,
the pressure being developed by osmotically active substances. For the
discharge to take place, there must be a weak point somewhere between
the spore and sterigma and this point is found at the top of the sterigma
where it is attached to the spore. If this weak point is constructed sim-
ilarly to the apex of an ascus with an operculum, then if the pressure
from within becomes great enough or if the balance of the spore on the
sterigma is disturbed, a tearing of the spore from the sterigma will
take place. Only the slightest change in the balance of the spore on
the sterigma is necessary to bring about the tearing away of the spore
from the sterigma. An example of this is found in Calocera cornea in
which the spore is attached off-center to the sterigma resembling a lever
with unequal arms attached weakly at the fulcrum. The droplet is
exuded from the spore at the end of the short arm of the lever thus
causing the spore to alter its position and resulting in the breaking away
of the spore from the sterigma at the weakly constructed apex. At the
same time, the internal pressure of the basidium causes the spore and
its drop to be forcefully discharged. There is little evidence to warrant
the acceptance of this theory.
A better explanation for the method of spore discharge in Gymnospor-
angium nidus-avis can be arrived at by comparing the spore discharge
in this species with what is known about this phenomenon in Phycomy-
cetes. This is a comparison previously made by Buller (1909) but re-
jected later (1922) after he apparently failed to find in the sterigma
a double wall which is prerequisite to this theory of spore discharge.
Buller (1909) uses the drawings of Nowakowski as a basis of his
theory, while Thaxter (1888) stated that the findings of the latter de-
lineator had to be considered as abnormal for the Entomophthoraceae.
Thaxter (1888) has shown that the conidia of Entomophthora are covered
with a delicate membrane which is continuous with that of the conidio-
phore. He stated further that the conidia take on water until the base
of the spore presses downward against the columella, until sufficient
pressure is exerted to rupture the membrane holding the conidium to the
conidiophore and thus bringing about discharge. This method of spore
discharge has been further emphasized and accurately depicted by Weston
(1923) for Sclerospora philippinensis in which species the discharge of
PRINCE: BASIDIOSPORE DISCHARGE 89
the spores resulted from the sudden bulging of the end wall of both
sterigma and apiculus to break the surrounding membrane, the remains
of which are evident on the discharged conidia as a narrow ragged collar
at the point where the spore was attached to the sterigma. The com-
parison of conidial discharge in Entomophthora and Sclerospora with
basidiospore discharge in Gymnosporangium may not at first appear
valid, but if nuclear behavior prior to spore formation be excluded in
these groups and if only the morphology and the development of the
two types of spores be considered, it becomes obvious that the methods
of spore discharge are similar, if not identical. In both groups the
spores are formed by a bulging out of the apex of the spore-bearing
structures followed by the laying down of septa at the base of the spores
within the original walls that surrounded the apiculi and spore-bearing
structures. The spores in both groups are discharged forcefully and
the mechanics involved appear to be the same, that is, there are surfaces
of two structures exerting force against each other and these surfaces
are held together by a common membrane and not until it is ruptured
can the forces thus involved be transformed into motion. The only
structural difference between the two Phycomycetes mentioned and the
species of Gymnosporangium under consideration is that in the latter
there is a relatively thick middle lamella interposed between the spore
and sterigma. If it can be accepted as true that the method of discharge
in these forms is the same, and all the evidence seems to confirm this,
then the method of spore discharge presents a striking instance of in-
dependent and parallel development in two widely separated groups
of fungi.
SUMMARY
The developmental stages of germinating teliospores of Gymnospor-
angium nidus-avis Thaxter have been observed and photographed. The
basidium was found to have an inner and outer wall, the latter being
common also to the sterigma and spore, the inner wall common only to
each cell. It is suggested that in the light of the recent work of Olive
(1942) the basidium may be compared to an ascus with four spores.
Basidiospore discharge was found to be dependent, not on the presence of
a droplet as suggested by Lohwag and others, or on the sudden expansion
of the basidium as suggested by Fayod, but rather the process is com-
parable to the one found in the Entomophthoraceae and Sclerospora. An
explanation of the phenomenon of basidiospore discharge, based on these
investigations, is described and illustrated. The present theory of spore
discharge is compared with those of previous investigators.
CLEMSON COLLEGE
Ciemson, S. C.
90 FarLowl1a, Vou. 1, 1943
LITERATURE CITED
Brefeld, O. (1877). Botanische Untersuchungen tiber Schimmelpilze. Heft 3.
Arthur Felix, Leipzig.
Buller, A. H. R. (1909). Researches on Fungi I. Longmans Green, London.
. (1922). Researches on Fungi II. Longmans Green, London.
. (1924). Researches on Fungi III. Longmans Green, London.
Coons, G. H. (1912). Some Investigations on the Cedar Rust Fungus, Gymno-
Sporangium juniperi-virginianae. Nebraska Agr. Exp. Sta. Rept. 25: 215-245.
Dietel, P. (1912). Uber die Abschleuderung der Sporidien bei den Uredineen.
Mycol. Centralblatt 1: 355-359.
Falck, R. (1904). Die Sporenverbreitung bei den Basidiomyceten und der bio-
logische Wert der Basidie. Cohn’s Beitr. Biol. Pflanzen. 9: 1-82.
Farlow, W. G. (1880). The Gymnosporangia or Cedar-Apples of the United States.
Anniv. Mem. Boston Soc. of Nat. Hist.
Fayod, V. (1889). Prodrome d’une Histoire des Agaricines. Ann. Sci. Series 7.
9: 271-272.
Giddings, N. J. (1918). Infection and Immunity in Apple Rust. W. Va. Agr. Exp.
Sta. Bull. 170: 9.
Hiley, W. E. (1919). The Fungal Diseases of the Common Larch. Clarendon
Press, Oxford.
Ingold, C. T. (1939). Spore Discharge in Land Plants. Clarendon Press, Oxford.
Klebahn, H. (1904). Die Wirtswechselnden Rostpilze. Borntraeger. Berlin.
Linder, D. H. (1940). Evolution of the Basidiomycetes and its Relation to the
Terminology of the Basidium. Mycologia 32: 419-447.
Lohwag, H. (1941). Anatomie der Asco- und Basidiomyceten. Handbuch der
Pflanzen Anatomie 6 (35): 456-457.
Massee, G. (1906). Text-book of Fungi. Duckworth, London.
Olive, L. S. (1942). Nuclear Phenomena Involved at Meiosis in Coleosporium
Helianthi. Jour. Elisha Mitch. Sci. Soc. 58: 43-51.
Schmitz, J. (1843). Beitrige zur Anatomie und Physiologie der Schwamme.
Linnaea 17: 434.
Thaxter, R. (1888). The Entomophthoreae of the United States. Mem. Boston
Soc. Nat. Hist. 4: 143-144.
- (1891). The Connecticut Species of Gymnosporangium (Cedar-Apples).
Conn. Agr. Exp. Sta. Bull. 107: 2-6.
Weston, W. H. Jr. (1923). Production and Dispersal of Conidia in the Philippine
Sclerosporas of Maize. Jour, Agr. Res. 23: 239-278.
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.
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92 Fartowia, Vou. 1, 1943
EXPLANATION OF PLATE 1
A: Basidium with narrow stipe cell formed when the teliospore germinated deep
in the telial matrix. B: Teliospore germinating on the surface of the telial matrix
producing a basidium without an evident stipe cell. A sterigma has started to form
on the third basidial cell. C: An early stage in the formation of a basidiospore
showing the globose enlargement at the end of a sterigma. D-E: The young spore
grows more on one side, elongates upward becoming more or less reniform. F: Sterig-
ma bearing a fully enlarged spore. G-H: A septum, indicated by arrow, is formed
in the sterigma a short distance below the spore forming an apiculus, hyaline ma-
terial becomes concentrated in the apiculus. I-J: Hyaline material is evident (J)
at the apex of the sterigma and a droplet appears on the flattened side of the spore
just before discharge. K: The end of the sterigma after discharge has a definite
convex wall. L-M: Note hyaline material in the tips of the two sterigmata in focus,
and that after spore discharge (a) the apex is convex indicating pressure from within.
The two left hand spores are from one basidium, the right hand one from another.
Pictures taken 8 minutes apart.
PRINCE: BASIDIOSPORE DISCHARGE 93
PLATE 1
a <7
es 7
stew eee =
1(1): 95-118 FARLOWIA January, 1943
THE GENUS PELLICULARIA (THELEPHORACEAE) !
DonaLp P. RoceErs
Since the completion of my account of the genus Botryobasidium (Univ.
Iowa St. N. H. 17: 10-19. 1935) many additional specimens have been
collected or otherwise become available for examination, including
material of eight additional species; and the necessity of adoption of
the earlier generic name Pellicularia, and of a revised nomenclature for
some of the species, has become inescapable. The present treatment
is as inclusive as accessible material would permit, and the nomenclature
is that determined by strict application of the Rules to the established
or probable synonymy. Without any doubt there are species yet un-
described that belong in Pellicularia, and others, already described among
the multitude of fungi assigned to Hypochnus, Corticium, Peniophora,
or more unlikely genera, unrecognized or unrecognizable as members
of the present group. It is not to be expected that this discussion can
serve as a monograph of Pellicularia: some of the specific units as here
broadly defined may be found susceptible of dissection into smaller but
still natural units, names will be displaced on the legitimate ground
of priority, and other species will be added. The most that is to be
hoped for is that a nucleus may here be found around which to assemble
information concerning a natural and, from the standpoint of phytopathol-
ogy as well as mycology, important group of organisms.
It will be observed that components of Pellicularia have been drawn
impartially from Hypochnus, Corticium, and Peniophora. It has many
times been argued, and may now be regarded as settled, that Hy pochnus
as defined by looseness of hymenial texture (“lack of a true hymenium’’)
is a concept as unworkable as it is unnatural. That some components of
Peniophora are much more closely related to components of Corticium
than to others included in Peniophora must be fully as apparent to even
a casual student of those genera; and both genera, in their present in-
clusive sense, are to be tolerated only until a more natural arrangement
of species can be worked out. Pellicularia, like Aleurodiscus and
Vararia, is a genus, whereas Corticium and Peniophora are at present
only heterogeneous accumulations.
It has been considered unnecessary to cite data, or locations in herbaria,
for most specimens examined. Critical material has been identified
sufficiently in the text to indicate the bases for some of the taxonomic
conclusions arrived at. Herbaria are referred to, after citation of the
specimens, according to the scheme presented by Lanjouw (Chron. Bot.
5: 142-150. 1939.) : BPI — Pathological and Mycological Collections of
the Bureau of Plant Industry, U. S. Department of Agriculture; FH =
Farlow Herbarium, general collection; FH-B= its Burt collection;
FH-C = its Curtis collection; FH-H = its von Hohnel collection; FH-P =
* Contribution from the Department of Botany, Brown University.
95
96 FarRLowliA, Vou. 1, 1943
its Patouillard collection; FP = collections of the Division of Forest
Pathology, New Haven; [A =Mycological Herbarium of the University
of Iowa; MICH = Herbarium of the University of Michigan; OTB =
Herbarium of the Division of Botany, Central Experimental Farms,
Ottawa; SP = Herbario da Seccao de Fitopatologia, Instituto Biologico,
Sao Paulo, Brazil (specimens lent by H. S. Fawcett) ; TENN = Herbar-
ium of the University of Tennessee; TRT = Herbarium of the University
of Toronto; D.P.R. is here used for my own herbarium. Color names
followed by (R) are used in the sense of Ridgway.
Material examined was uniformly treated according to the highly
satisfactory KOH-phloxine technique introduced by Martin (Mycologia
26: 264. 1934.). Species not here illustrated have already been figured
in the cited discussion of Botryobasidium. All figures were drawn with
the camera lucida at 1960 and reduced in reproduction to 1000 X.
I am indebted to the Oregon General Research Council for grants in
aid of mycological collection and research carried on at Oregon State
College; to Dr. J. N. Couch of North Carolina, Dr. H. S. Fawcett of
the Citrus Experiment Station at Riverside, Professor F. O. Grover of
Oberlin, Dr. L. R. Hesler of Tennessee, Dr. J. H. Kotila of the United
States Department of Agriculture, Dr. G. W. Martin of Iowa, Dr. Mildred
K. Nobles of Ottawa, Mr. J. A. Stevenson of the United States Depart-
ment of Agriculture, Dr. George F. Weber of Florida, Dr. L. E. Weh-
meyer of Michigan, and Dr. W. L. White of Harvard for material here
treated; to Dr. W. H. Snell of Brown for counsel and encouragement
during the preparation of the present paper, and for provision of
ample opportunity and facilities for mycological research; to Dr. D. H.
Linder of Harvard and Dr. H. 5S. Jackson of Toronto for advice and
assistance in many matters and for rendering available the abundant
important material in their charge; and to Alpha Mae Rogers for the
preparation of illustrations and manuscript.
PELLICULARIA Cooke, Grev. 4: 116. 1876; 4: 134. 1876.
Corticium [Sect.] Botryedea Bourd. & Galz., Soc. Myc. Fr. Bul. 27: 247. 1911;
Hym Fr. 238. [1928]; Overh., Mycologia 26: 509. 1934,
Tomentella Sect. Tomentellastrum [Subsect.] Botrytes Bourd. & Galz., Soc. Myc.
Fr. Bul. 40: 137. 1924; Hym. Fr. 481. [1928].
Botryobasidium Donk, Nederl. Myc. Ver. Med. 18-20: 116. 1931; Rogers, Univ.
Iowa St. N. H. 17: 10. 1935.
Botryohypochnus Donk, Nederl. Myc. Ver. Med. 18-20: 118. 1931.
Type: P. Koleroga Cooke.
Fructification resupinate, mucedinoid or hypochnoid, reticulate-pellicu-
lar, finely granulose, under the lens more or less tufted, or even and
loose-membranous; hyphae strongly stainable in aniline blue, thick,
short-celled except for basal strands, branching at right angles and
often with the formation of cruciform cells, the ascending hyphae usually
several times cymosely divided, bearing the terminal basidia in more
or less candelabrum-like clusters, or in parasitic species sometimes rela-
Rocers: THE GENUS PELLICULARIA 97
tively short and little divided; basidia subcylindric, not greatly exceeding
in diameter the supporting cells, relatively short, bearing 4 or in several
species 6-8 sterigmata; spores smooth-walled or rarely asperulate or
spinulose, colorless or pale ochraceous; cystidia wanting, or present and
of various forms. Saprobes or facultative parasites,
Pellicularia is distinguishable by very short-celled, stout hyphae (of
which some portions in every species except P. chordulata attain a di-
ameter of 10-18 »), by right-angled branching of the mycelium, by very
stout basidia, and by mucedinoid texture.
The genus Botryobasidium Donk is a distinct and well characterized
systematic and phyletic unit, for which the need had long existed. Pelli-
cularia Cooke, of which P. Koleroga is thus far the only species, was
described by its distinguished author as subgelatinous, lacking basidia,
having echinulate and sessile spores, and falling somewhere near “Oidium”
(i. e., the imperfect stage of certain powdery mildews). Nevertheless,
P. Koleroga is a good Botryobasidium. There seems to be no earlier
genus named Pellicularia; and Miss Wakefield’s sketches of the type speci-
men of P. Koleroga, published by Burt, show both basidia and spores.
Consequently, Pellicularia cannot be disposed of as would be most appro-
priate — at least, not through invocation of Art. 57 or Art. 61 of the Rules.
Furthermore, Botryobasidium cannot be regarded as sufficiently well estab-
lished to deserve the application of the greatly overworked principle of
nomina conservanda. There is no choice but to reduce Botryobasidium to
synonymy, and to take up as soon as possible the valid name Pellicularia.
KEY TO THE SPECIES
EeSpores Tough: walled vec fe ee cee Wah et a ae Nn ee ale Z
1 Spores smooth walled sees. elie rns. cis aie. cit Pures tne cman umn Fae NE ct 4
2. Clamps present in subhymenial hyphae; spores asperulate, subglobose
1. chordulata
2s, lampan lacing were, Waiter ae ne atte me ren ss eet ee ee. 3
3. Spores subglobose, even or angular in outline, spinose ....... 2. isabellina
3. Spores ellipsoid, minutely asperulate ....................... 3. asperula
4c Cystidia present emergent er sane ee ee ee ee 5
4. Cystidia lackingae) ae iy ncn tin et none Are Sian ese ee 8
SeeCvstidigesep tale my ae ew ema eee es ce ee eee nd ie dere tie. sy 6
Sar Cystidia caseptate ma. tome reer een Ne ane ae ena MONT od. pee eh: Sa rete i
6. Clamps absent from cystidia and mycelium; cystidia stout, brown-
ENCLUSLCCS Stra. ee reign eter ae en ae 4. Langloisii
6. Clamps present on cystidia and mycelium; cystidia unencrusted or
with a few loose colorless mineral platelets, slender .. 5. cystidiata
7. Cystidia thick-walled at the base, becoming very thin-walled toward the
apex; spores thick-walled ...................20eeeeeee 6. ochroleuca
7. Cystidia thin-walled (or with walls uniformly slightly thickened); spores
thin-walled +c: eee: Me hag hae creer ela et, v2 ele 7. ansosa
Si, Glamps: present) grew, Meets: coer ieee tens a tere ei ye 8. subcoronata
S27@lamps. lacking uc. feuee eect anne Wt ete on coe ce Odea cade 8 9
9. Fructification formed on dead plant parts or on soil; saprobes ............ 10
9. Fructifications formed on living plant parts. or on soil in contact with living
plants: .parasitesm. ya.) ere; A Spee pen ee 13
98 FarLowlA, VOL. 1, 1943
10. Spores germinating by repetition, with large, truncate apiculus; sterig-
mata (2—) 4, very large .......0... 000. ccc cee 9. flavescens
10. Spores not germinating by repetition, with small apiculus; sterigmata
POLO > RN hogs os sir 84s buy dpa ot oe aie aan ee aa ll
1l. Spores 4-7 (-7.5) yu. long, subglobose to broadly ellipsoid-fusiform
10. pruinata
11. Spores 7-12 ». or more long, fusoid-cylindric to navicular ................ 12
12. Hyphae mostly 8 y, or less in diameter; spores slender-navicular,
ee) Or he os sab ee wid ede heer a6 sis eee 11. lembospora
12. Many hyphae 10 p. or more in diameter; spores variable, in most collec- |
Pe ome larger an (0 8 20 fe or alrcy ine vane eens 12. vaga
13. Spores subcylindric to elongate-ellipsoid, flattened or depressed on the inside,
slightly thick-walled ......0...0000.. 0000.00.00... 00000. 13. Koleroga
13. Spores relatively broader, ellipsoid, oblong-ellipsoid, or ovate. not at all de-
RCH CL AIM AWB oa ck is ks, deen nca parietal OR) WED REIN ou ed 14
14, Spores ellipsoid or oblong-ellipsoid ................... 14. filamentosa
14. Spores ovate, 5.2-7.8 X4.9-5.9 wo. ca ls seivisebiee des 15. C. praticola
1. Pellicularia chordulata sp. nov.
FIGURE 1,
Fructificatio tenuis, hypochnoideo-membranacea, alutacea; hyphae (3.5-) 4.5-6
(-8) p diam., basales vel in chordulis fasciculatae vel liberae, passim nodoso-septatae
(ansis anastomosis 3-4.5 » crassis), breviter articulatae. hyalinae vel paululo luteo-
tinctae, subhymeniales ubique nodoso-septatae, tenuiter tunicatae; basidia in cymis
regularibus producta, subcylindracea, 12.5-16-21 x 6-7-8 y, sterigmata 4 divergentia
2-4.5 (-6) pw longit. gerentia; sporae subglobosae, 4-5.5 x 44.5 uw, asperulae, tunica
leviter incrassata, apiculo conico, | y, longit.
Fructification thin, hypochnoid-membranous, under the binocular
minutely reticulate-poroid, even or irregularly granulose, readily sep-
arable, when dry Cartridge Buff (R) to a color between Light Buff (R)
and Warm Buff (R); all parts strongly stainable by aniline blue; hyphae
(3.5-) 4.5-6 (—8) yw in diameter with the thickest and thinnest portions
both among the free subicular strands, the basal in part compacted into firm
cords, hyaline or only slightly tinted, anastomosing but with only very
rare clamps, and in part distinct, short-celled, branching at right angles,
with clamps at many septa 3—4.5 p» thick, giving rise to thinner-walled
subhymenial branches, short-celled, with clamps throughout; basidia in
candelabrum-like clusters, subtended by strong proliferating clamps,
clavate-subcylindric or slightly barrel-shaped, 12.5—-16—21 x 6-7-8 4h,
bearing 4 divergent sterigmata 24.5 (—6) » long; spores nearly globose,
slightly flattened on the inside, 4—-5.5 x 44.5 py, the wall slightly thick-
ened and ochraceous-tinted, asperulate, the apiculus distinct, conic, hya-
line, 1 » long. |
On dead bark and wood of Populus tremuloides, Salix sp., and Tilia
americana.
Specimens examined: Ontario, Gull L. Pt., L. Temagami, VIII.20.36, R. Biggs
(TRT 10408) ; Sandy Inlet, L. Temagami, VIII.12.31, G. EF. Thompson (TRT 2464) ;
Rondeau Gov't Park, VIII.15.34, R. F. Cain (TRT 7286); Onto, Ten Mile Cr., west
of Toledo, VIHI.12.35, D. P. Regers 946, type, and 947; Iowa, Wellman, VIII.30.32,
L. W. Miller (1A, D.P.R. 948) .
Rocers: THE GENUS PELLICULARIA 99
A species readily distinguished by the asperulate (rather than spinose )
spores, clamp-bearing mycelium, and prominent subicular cords. In
spore-ornamention, in possession of cords, and in mycelium more delicate
than that of most species of Pellicularia, the present fungus approaches
Corticium sect. Humicola of Bourdot & Galzin. In the keys of the
Hyménomycétes de France it would be associated with C. sulphureum, or
might be sought near C. araneosum or (in Tomentella) in the same
bracket with T. testaceogilva. In Burt’s keys to Hypochnus the specimens
cited would be assigned to H. sparsus, which, however, has typical long
basidia, strong sterigmata, and the angular-spinose spores of a Tomentella,
and mycelium not approaching that of a Pellicularia.
2. Pellicularia isabellina (Fries) comb. nov.
Thelephora isabellina Fr., Epicr. 544. 1838.
Hypochnus isabellinus (Fr.) Fr., Summ. Veg. Scand, 337, 1849; Bres., Ann.
Myc. 1: 106. 1903; Burt, Mo. Bot. Gard. Ann. 3: 222. fig. 12. 1916; Wakef.,
Br. Myc. Soc. Tr. 6: 133. 1919; nec H. isabellinus sensu Schroet. in Cohn,
Krypt.-Fl. Schles. 3 (1): 417. 1888.
Corticium isabellinum (Fr.) Fr., Hym. Eur. 660. 1874; Mass., Linn. Soc. Bot.
Jour. 27: 149. 1890; nec C. isabellinum sensu Pat., Tab. Anal. 1: 16. fig. 23.
1883, nec sensu Eichl. ex Bres., Ann. Myc. 1: 98. 1903.
Hypochnus argillaceus Karst., Soc. Faun. Fl. Fenn. Med. 6: 13. 1881; Bidr.
Kanned. Finl. Nat. Folk 37: 164, 1882.
Lyomyces isabellinus (Fr.) Karst., Bidr. Kanned. Finl. Nat. Folk. 37: 153. 1882.
Zygodesmus argillaceus (Karst.) Karst., in Rabenh.-Wint., Fungi Eur. 3188.
1884; Bidr. Kanned. Finl. Nat. Folk 51: 420.1892.
Odontia tenerrima Wettst., Zool.-Bot. Ges. Wien Verh. 38: 178. 1888.
Tomentella flava Bref., Unters. 8: 11. pl. 1, fig. 11-14. 1889,
Hypochnus flavus (Bref.) Sacc., Syll. Fung. 9: 242, 1891.
Tomentella ochraceo-viridis Pat., Soc. Myc. Fr. Bul. 9: 134. 1893; Cat. Pl. Cel.
Tunisie 63. 1897.
Tomentella isabellina (Fr.) Hohn. & Litsch., Ak. Wiss. Wien, Math.-Nat. Kl.,
Sitzungsb. 115, I: 1570. 1906; Wiesner-Festschr. 78. 1908; Bourd. & Gallz.,
Soc. Myc. Fr. Bul. 40: 137. 1924; Hym. Fr. 482. fig. 121. [1928].
Botryobasidium isabellinum (Fr.) Rogers. Univ. Iowa St. N. H. 17: 11. pl. 2,
fig. 5. 1935.
Hypochnus isabellinus Fr., Obs. Myc. 2: 281. pl. 6, fig. 3. 1818.
Fructification arachnoid-pruinose, soon becoming granular, loose-
hypochnoid and relatively thick, at first whitish, soon buff or isabelline,
when dry Ivory Yellow (R) to Clay Color (R) and Sayal Brown (R),
under the binocular uniformly hypochnoid; hyphae short-celled, 6-14 p
in diameter, branching at right angles, without clamps, hyaline to ochra-
ceous, in older portions the basal strands usually brownish-ochraceous
and with bilamellate walls; basidia clavate-cylindric or clavate-obovate,
not constricted at the septum, 15-25 x8-11.5 yp, bearing 4 strongly di-
vergent sterigmata 3.5-6 » long and 2-3 » thick at the base; spores
mostly somewhat ochraceous, subglobose or subglobose-ellipsoid, in
occasional specimens subangular, 6—-10.5x5.5-9 yp, tapered or more
commonly abruptly giving rise to the lateral, truncate-conic apiculus,
the surface marked by scattered obtuse-cylindric or- obtuse-conic spines
100 FarLowia, Vou. 1, 1943
of variable length, in the specimens where least developed 0.25-0.75 p
long, in those where best developed 2—3.5 » and expanded at the base
so as to give a subangular outline to the spore.
American specimens on soil, on dead Pteridium latiusculum, on bark
and wood of Abies balsamea, Pinus virginiana, unidentified conifers,
Betula lutea, Liriodendron Tulipifera, Populus grandidentata, P. tremu-
loides, Salix sp., Tilia americana, and on other hardwoods.
Specimens examined: FinLanp, Mustiala, P. A. Karsten, Rabenh.-Wint. Fung. Eur.
1883, as Z. argillaceus (FH); SwepEN, Upsala’ Halmbyboda, 1853, E. P. Fries, as
‘Hypochnus s. Thelephora isabellind (FH-C); Ausrria, Steiermark, 1886, Eber-
staller, type of O. tenerrima (FH-H); Ecuapor, Quito, 1892, de Lagerheim, type and
paratype of JT. ochraceo-viridis (FH-P); also specimens from France, Colombia,
British Guiana, Nova Scotia, Quebec, Ontario. Manitoba, Massachusetts, Connecticut,
New York, Pennsylvania, Florida, Tennessee, Louisiana, Michigan, Iowa, and Oregon.
Except in spore-ornamentation, the abundant material at hand is not-
ably uniform. The spores with perfectly cylindric spines less than 34 p
in length would seem amply distinct from those whose long, conical
spines render the spore-body almost polyhedral, but the more abundant
intermediates, and the lack of other variation, render the setting up of
a new species for the long-spined forms a singularly pointless exercise.
The only possible basis for division would be shape of spines—conical
versus cylindrical—and even thus a natural cleavage could probably not
be found.
Donk published a genus Botryohypochnus for this species (Nederl.
Myc. Ver. Med. 18-20: 118. 1931.) but did not list the new binomial.
3. Pellicularia asperula sp. nov.
FIGURE 2.
Fructificatio tenuissima, haud continua, mucedinoidea, albida; hyphae enodosae,
angulis rectis ramosae, 4-11.5 (-14) y diam., basales e cellulis longis compositae,
interdum crasse tunicatae, luteae, fertiles creberrime ramosae, cymosae, angustiores;
basidia subcylindracea, ad apicem leviter expansa, 13.5-17.5 x 7.5-8 p, sterigmata 6-8
peripherica, 2.5-3 y. longit., gerentia; sporae ellipsoideae, uno latere subapplanatae,
crasse tunicatae, asperulae, 4.5-5.5 x 3.5-4.5 u.
Fructification delicate, discontinuous, mucedinoid, when fresh whitish,
Pale Neutral Gray (R), when dry about the same (i. e., white, and
allowing the substratum to show through); hyphae regular, without
clamps, 4-11.5 (—14) » in diameter, branching at right angles, usually
with the formation of cruciform cells, the basal long-celled, wide, often
with thickened and yellowish walls, the fertile portions with abundant
cymose branching, shorter-celled, more slender; basidia subcylindric,
somewhat expanded toward the summit, 13.5-17.5x7.5-8 y, truncate,
bearing at the periphery 6-8 divergent sterigmata 2.5-3 » long; spores
ellipsoid, somewhat flattened on the inside, fairly thick-walled, with
minutely asperulate non-amyloid surface, 4.5-5.5 x 3.5-4.5 p.
On fallen decayed hardwood limb.
Specimen examined: Cusa, Blanco’s Woods, Soledad, Cienfuegos, Santa Clara Prov.,
VII.1.41, W. L. White 603, type (FH).
Rocers: THE GENUS PELLICULARIA 101
A typical Pellicularia in all respects but the spores. These are very
abundant in the material cited, sometimes free, but often, as in other
species of the genus, held in groups by the collapsed, nearly invisible
basidia. Distinct mature basidia are very rare, and may vary more
widely than the description allows. The form of the spores makes them
unique in the genus, as does the type of wall-sculpturing; they resemble
the rough-walled conidia of some species of Aspergillus, but have apiculli,
and were seen attached to the sterigmata.
4. Pellicularia Langloisii (Pat.) comb. nov.
Hypochnus Langloisii Pat., Soc. Myc. Fr. Bul. 24: 3. 1908.
Peniophora magnahypha Burt, Mo. Bot. Gard. Ann. 12: 238. [1926].
FIGURE 3.
Fructification cream-colored, thin-hypochnoid, fragile, under the lens
- a continuous, minutely poroid, whitish layer, closely brown-setulose;
hyphae without clamps, very short-celled, branching at right angles to form
candelabrum-like fascicles, with many cruciform cells, the basal, and
the main axes, slightly thick-walled, straight and rigid, up to 10-19 p
in diameter, few, the lateral branches progressively decreasing in diam-
eter down to (3.5—) 6 »; cystidia formed by prolongation of the main
axes of the hyphal fascicles, tapering outward, obtuse, with a few septa,
encrusted with orange-brown granular material, 100-160 x 9-11 p, long-
emergent; basidia borne in symmetrical cymes, subcylindric,
11-15 x 6-7 p, bearing 4 sterigmata 3.5-5 (-6) » long; spores smooth,
hyaline, oblong, flattened or slightly depressed on the inner side (sub-
reniform), (5.5-) 6-8 x 3-4.5 (-5) up.
On decorticated hardwoods.
No material in addition to the type specimens has been encountered;
these are from FLoripa, Coconut Grove, R. Thaxter 57, P. magnahypha
(FH), and Louistana, St. Martinville, Langlois 2968, H. Langloisii,
(FH-P).
A typical and highly characteristic member of this genus, distinguish-
able from the other cystidiate species by incrustation, septation, and lack
of clamps on the cystidia. The brown incrustation persists in lactophenol
but disappears in KOH. The Louisiana specimen, according to Lang-
lois’s notes, had “dents jaunes sur un blanc subiculum”; at present it
is mostly disintegrated, and the Florida specimen alone shows all struc-
tural details.
5. Pellicularia cystidiata sp. nov.
FIGURE 4.
Fructificatio tenuis, pruinoso-hypochnoides vel mucedinoides, vix separabilis, sordide
albida, sub lente minute pilosa; hyphae 4.5-6-10.5 p, diam., majori ex parte repentes,
zygodesmatibus minutis ubique ornata, ramos breves atque plura cystidia emittentes;
cystidia apicem versus attenuata, pauciseptata, obtusa, tenuiter tunicata, (50—-) 70—
110x 6-10 »; basidia plerumque obovata, 13-15x9-11 yp, sterigmata 4 gracilia,
3.5-4.5 p. longit., gerentia; sporae fusiformes, apiculum versus curvulae (8—) 11-13
(-16) x 4-5 p.
102 FarLowia, VoL. 1, 1943
Fructification thin, pruinose-hypochnoid or loose and mucedinoid,
inseparable from the substratum or separable only in small bits, whitish
with a faint yellow tinge, lighter than Pale Olive Gray (R) when dry,
under the binocular loosely pilose-hypochnoid; hyphae short-celled, with
relatively minute clamps throughout, 4.5-6-10.5 » in diameter, mostly
repent, giving off, in a direction about perpendicular to the substratum,
fertile branches 1-6 cells long, and abundant cystidia; cystidia little
differentiated, thin-walled, colorless, tapering toward the apex, obtuse,
(50-) 70-110 x 6-10 , mostly 2-6-septate and with minute clamps, with
a few loosely attached plate-like mineral bodies; basidia barrel-shaped
to obovate, 13-15 x 9-11 y, bearing 4 delicate sterigmata 3.54.5 long;
spores subfusiform, curved toward the apiculus, thin-walled, colorless,
(8) 11-13 (-16) x4-5 un.
On Picea sp. and log of undetermined conifer.
Specimens examined: Ontario, Constance Bay, VI.1933, J. W. Groves (OTB
F6335), type; Connecticut, N. Bloomfield, VIII.28.38, H. G. Eno (FP 84049) ;
Iowa, Iowa City, V.2.1932, D. P. Rogers, 786.
Distinguishable from P. subcoronata, which likewise has clamps at
the septa, by the presence of cystidia, greater size of spores, lack of
regular and extensive candelabrum-like branching of fertile hyphae, and
relative inconspicuousness of the clamp-cells; distinguishable from P.
ansosa by the occurrence in P. cystidiata of septate cystidia, different
basidia, more delicate clamp-cells, and longer spores. The spores re-
semble somewhat those of P. ochroleuca, but are smaller and considerably
more tapered —especially toward the base —than in that species, and
are thin-walled whereas those of P. ochroleuca have strongly thickened
walls; the cystidia differ in every respect.
6. Pellicularia ochroleuca (Bres.) comb. nov.
Coniophora ochroleuca Bres. apud Brinkm., Westf. Prov.-Ver. Jahresber. 26:
130. 1898.
Peniophora ochroleuca (Bres.) Héhn. & Litsch., Ak. Wiss. Wien, Math.-Nat. K1.,
Sitzungsb. 117, I: 1107. fig. 6. 1908; Bourd. & Galz., Hym. Fr. 301. [1928].
Coniophorella ochroleuca (Bres.) Brinkm., Westf. Prov.-Ver. Jahresber. 44: 41.
1916. ;
Peniophora fusispora sensu Hohn. & Litsch., Ann. Myc. 4: 289. 1906; Bourd.
& Galz., Soc. Myc. Fr. Bul. 28: 391. [1913]; Hym. Fr. 300. [1928], quantum
ad descr. et specim., typo excl.; nec Hypochnus fusisporus Schroet. in Cohn,
Krypt.-Fl. Schles. 3 (1): 416. 1888 (= Pellicularia flavescens (Bon.) ).
FIGURE 5.
Fructification hypochnoid, when fresh sordid whitish or pale isabelline,
occupying small separate patches on the substratum, under the binocular
minutely interrupted, tufted, when dry between Warm Buff (R) and
Cream Color (R), pilose, readily separable; hyphae with prominent
clamps at most septa, (4.5-) 6-12 in diameter, short-celled, branching
at right angles but lacking cruciform cells, in part soon collapsed;
cystidia tapered evenly toward the apex, or up to the middle and with
Rocers: THE GENUS PELLICULARIA 103
the upper half subfusiform, obtuse, moderately thick-walled below, very
thin-walled above, continuous, borne on stout mycelium, subtended by
a prominent clamp, 95-360 x 6-12 ; basidia cylindric or thick-claviform,
somewhat constricted near the middle but not at the basal septum,
18-48 x 7-10 p, bearing 4 slender erect sterigmata 3-6 » long; spores
hyaline when fresh, becoming yellowish on drying, thick-walled, evenly
navicular-fusiform, obliquely apiculate, 9.5-18 x4-8 up.
American specimens on Pinus sp., Pseudotsuga mucronata, undeter-
mined conifer, and Acer sp.
Specimens examined: GERMANY, Rheinauerwald bei Rastadt, Baden, XI.1877,
Schroeter, as Hypochnus fusiger (FH-H); and Lengerich, Westfalen, X1.1897, Brink-
mann, type of C. ochroleuca (FH-B), and II.1899, Westf. Pilze 28 (FH); also
specimens from France, Ontario, Massachusetts, and Oregon.
A Pellicularia with aseptate cystidia thick-walled at the base and thin-
walled at the apex, much like those of Peniophora subalutacea. Under
low magnification the living fructification strongly resembles columnar
hoar-frost — and Peniophora chaetophora (Hohn.) Héhn. & Litsch. With
its relatively long basidia and its hyphae lacking cruciform cells P. och-
roleuca is far from typical of Pellicularia, but appears to belong in this
genus.
The type of H. fusisporus Schroet. is not among Hohnel’s specimens,
nor is there even a slide of it. As the discussion in the “Revision der
Corticieen in Dr. J. Schroter’s ‘Pilze Schlesiens’” (Ann. Myc., 1. c.) only
ambiguously indicates, the account of P. fusispora was drawn up from
a specinfen carrying Schroeter’s herbarium name Hypochnus fusiger.
That specimen and the type of Coniophora ochroleuca agree completely.
But the collection data for H. fusiger are not those published by Schroeter
for H. fusisporus, and it must be apparent from his description that the
two are not the same. The fructification of H. fusisporus is said by
Schroeter (1. c.) to be “ohne Endborsten” (i. e., without cystidia) , while
the specimen of H. fusiger has cystidia over 200 » long. The spores
of H. fusisporus are said to be “an beiden Enden stark verschmalert, fast
citronenférmig,” while the spores in fusiger are evenly tapered, and
more nearly banana-shaped; in H. fusisporus the spores “keimen leicht
und bilden oft an kurzen pfriemlichen Keimschlaiichen secundare Sporen,”
while no germination is observable in H. fusiger or in any of the ten
other specimens examined of the present species. It seems clear enough
that Schroeter’s H. fusisporus is Pellicularia flavescens (q. v.), and
that Corticium fusisporum in Brinkm. Westf. Pilze 53 truly represents
Schroeter’s species. For the cystidiate Pellicularia here described, then,
Bresadola’s name must be used.
7. Pellicularia ansosa Jackson & Rogers sp. nov.
FIGURE 6.
Fructificatio alutacea, hypochnoidea, sub lente obscure pilosa, continua; hyphae
conspicue nodoso-septatae, 7-10-15 y, diam., ansarum nonnullarum tunica exteriori
104 Fartowia, VoL. 1, 1943
incrassata ad 2.5 y; cystidia plerumque tenuiter tunicata, subcylindracea, hic illic
constricta, obtusa, continua, haud incrustata, 55-120 x 8-12.5 u; basidia sub-
cylindracea, subventricosa, 17-29 x 8-10 yw, sterigmata (4—) 6 recurva, 4-5 pu. longit.,
gerentia; sporae naviculiformes, attenuatae, tenuiter tunicatae, 8-9 x 4-5 y,.
Fructification buffy, about Cream Color (R), continuous, under the
binocular hypochnoid, obscurely pilose; hyphae distinct, firm, with large
clamps throughout, branching at right angles, 7-10-15 p» in diameter,
the basal long-celled, often sending out several branches from a single
cell, often with the outer wall of the clamp-cell strongly thickened (up
to 2.5 ), the subhymenial hyphae thin-walled and short-celled; cystidia
arising among the basidia, thin-walled or with walls only very slightly
and uniformly thickened, emergent, irregularly subcylindric to subfusi-
form, constricted here and there, obtuse, unincrusted, aseptate,
55-120 x 8-12.5 »; basidia arising in simple cymes, subcylindric, some-
what constricted near the middle, usually subventricose, 17-29 x 8-10 up,
bearing (4-) 6 sterigmata 4-5 » long and recurved; spores broadly
fusiform or navicular, curved and tapered laterally to the apiculus,
thin-walled. 8-9x4-5 pu.
On conifer wood and on bark of Picea sitchensis.
Specimens examined: BrivisH CoLtumBia, Cumshewa Inlet, Queen Charlotte I,
V.18.24, J. E. Bier (OTB F10894, TRT) ; Wasuincton, Deer L. Trail, Olympic Mts.,
VI.13.39, A. H. Smith 14344, type (MICH, TRT).
In its aseptate cystidia and clamp-bearing mycelium distinct from all
species except P. ochroleuca, from which it differs in the more distinct
mycelium, the thin-walled spores, the shorter and ventricose basidia,
and the cystidial walls not strongly thickened toward the base. P. ansosa
should be distinguishable from P. ochroleuca by examination with a
hand-lens; the present species is continuous and almost imperceptibly
pilose, where ochroleuca is areolate and bristly.
8. Pellicularia subcoronata (Héhn. & Litsch.) comb. nov.
Corticium subcoronatum Hohn. & Litsch., Ak. Wiss. Wien, Math.-Nat. K1.,
Sitzungsb. 116, I: 822. 1907; Wakef., Br. Myc. Soc. Tr. 4: 118. 1913; Bourd.
& Galz., Soc. Myc. Fr. Bul. 27: 249. 1911; Hym. Fr. 238. [1928]; Coker, El.
Mitchell Sc. Soc, Jour. 36: 174, 1921; Overh., Mycologia 26: 510. pl. 55,
fig. 11. 1934; Rick, Brotéria ser. trim. Ciénc. Nat. 3: 156. 1934.
Botryobasidium subcoronatum (Hohn. & Litsch.) Donk, Nederl. Myc. Ver. Med.
18-20: 117. 1931; Rogers, Univ. Iowa St. N. H. 17: 12. pl. 2, fig. 6, 1935.
FIGURE 7.
Fructification white or whitish when fresh, arachnoid or loose and
continuous, under the binocular delicately pilose to compact, byssoid,
when dry white or cream to buff, soft-membranous, separable but not
coherent in a pellicle; hyphae with large clamps (2-4 p» thick) through-
out, or more rarely wanting from occasional septa, branching at right
angles, the basal hyphae comparatively long-celled and straight, in part
with yellowish walls, the ascending branches short-celled, colorless,
4—7—-1] «; basidia borne in candelabrum-like clusters, subcylindric, usually
Rocers: THE GENUS PELLICULARIA 105
ventricose-inflated, (11.5—) 15-18 (—25) x 6-7.5-9 py, bearing 6-8 in-
curved sterigmata 3-4.5-6 » long; spores slender-fusoid, thickest near the
apiculus, long-attenuate to the obtuse apex, short-attenuate to the lateral
apiculus (i. e., slender-navicular), or rarely broad-navicular, or fusoid-
cylindric, 6.5-8.5 (—15) x (2-) 3-4 (-5) p.
Commonly on wood so completely decayed as to be difficult of identi-
fication, or on bark or sawdust; American collections on Abies balsamea,
A. Fraseri, Picea sp., Pinus resinosa, P. Strobus, Pseudotsuga mucronata,
Tsuga canadensis, Acer sp., Castanea dentata, Fagus grandifolia, Populus
sp., Quercus sp., numerous undetermined broad-leaved and coniferous trees,
and soil.
Specimens examined: GerMANy, Wannsee, Berlin, VI. 1894, P. Sydow, Myc. March.
4105, as Peniophora Greschikii, type of C. subcoronatum (FH-H); also from Nova
Scotia, Quebec, Ontario, Manitoba, British Columbia, Vermont, New Hampshire,
Massachusetts, Rhode Island, Connecticut, New York, New Jersey, North Carolina,
Missouri, Idaho, and Oregon.
Readily recognizable by the abundant large clamps and lack of
cystidia; distinct from other species also in the ventricose-subcylindric
basidia and the (usually) very slender spores. Two collections, aberrant
in the possession of only scattered clamps on the basal mycelium and of
spores 7—15x (1.5-) 2-3 p, nevertheless agree with typical material in
the form of the clamps, in constant occurrence of clamps at the base of
the basidia, in form of basidia, and in texture, and are placed here.
9. Pellicularia flavescens (Bon.) comb. nov.
Hypochnus flavescens Bon., Handb. 160. 1851; Fckl., Nass. Ver. f. Naturk. Jahrb.
25-26: 291. (Symb. Myc. 1 Nachtr. 3.) 1871.
Corticium flavescens (Bon.) Wint., in Rabenh., Krypt.-Fl. 1 (1): 329. 1882;
Mass., Linn. Soc. Bot. Jour. 27: 149. 1890; Hohn., Osterr. Bot. Zeits. 54:
428, 1904; Hohn. & Litsch., Ak. Wiss. Wien, Math.-Nat. K1., Sitzungsb. 115, I:
1607. 1906; 116, I: 835. fig. 17. 1907; Bourd. & Galz., Soc. Myc. Fr. Bul. 27:
247. 1911; Hym. Fr. 239. [1928]; Wakef. & Pears., Br. Myc. Soc. Tr. 6:
317. fig. 1920; nec C. flavescens Bres., Ann. Myc. 3: 163. 1905 (= C. teutobur-
gense Brinkm., Westf. Prov.-Ver. Jahresber. 44: 38. 1916.).
Hy pochnus fusisporus Schroet., in Cohn, Krypt.-Fl. Schles. 3 (1): 416. 1888.
Corticium fusisporum (Schroet.) Brinkm., Westf. Pilze 53. 1904; nec C. fusi-
sporum Cke. & EIll., Grev. 8: 11. 1879 (= Coniophora fusispora (Cke. & Ell.)
Sacc., Syll. Fung. 6: 650. 1888.).
Peniophora fusispora (Schroet.) Hohn. & Litsch., Ann. Myc. 4: 289, 1906, et
auctt. omnium, quantum ad typum, specim. descr. excl.
Coniophora ochracea Mass., Linn. Soc. Bot. Jour. 25: 137. pl. 47, fig. 13. 1889.
Corticium frustulosum Bres., Ann. Myc. 1: 98. 1903; Ann. Myc. 9: 425. 1911;
Bourd. & Galz., Soc. Myc. Fr. Bul. 27: 247. 1911; Hym. Fr. 239, [1928]; Rick,
Brotéria ser. trim. Ciénc. Nat. 3: 158. 1934.
Coniophora vaga Burt, Mo. Bot. Gard. Ann, 4: 251. fig. 8. 1917.
Corticium frustulosum var. intermedia Bourd. & Galz., Hym. Fr. 240. [1928].
Corticium fenestratum Overh., Mycologia 26: 510. pl. 55, fig. 5. 1934.
Botryobasidium flavescens (Bon.) Rogers, Univ. Iowa St. N. H. 17: 13. pl. 2,
fig. 8. 1935,
Botryobasidium ochraceum (Mass.) Donk ex Rogers, Univ. Iowa St. N. H. 17:
16. pl. 2, fig. 7. 1935.
106 Fartowia, VoL. 1, 1943
Fructification at first delicately pruinose-arachnoid, soon becoming
loosely and coarsely hypochnoid, on drying becoming, except in very
young specimens, distinctly tufted or reticulate-floccose, or rarely con-
tinuous, whitish, sordid whitish, through Naples Yellow (R) and Warm
Buff (R) to Tawny Olive (R); hyphae without clamps, branching at
right angles with many cruciform cells, the basal repent and in part
colored, with cells several times as long as wide, often thick-walled, up
to 10-12 w» in diameter, the remainder very short-celled, thin-walled,
colorless, 5—8—10 » in diameter; basidia clavate-cylindric, little attenu-
ated toward the base, 13-20-24x 8-12-15 yp, bearing 4 or sometimes
only 2 stout sterigmata 6-9-15 x 2-4 1; spores typically with an ellipsoid-
subglobose body, truncate apiculus, and obtuse distal false-apiculus,
being thus asymmetrically lemon-shaped, but sometimes without the
false-apiculus, and thus subglobose, 7.5-13-15 x (4.5—) 6-10 p, hyaline
or with ochraceous wall, germinating by repetition.
American specimens on Ustulina vulgaris, bark of living trunk of
Carpinus caroliniana, and on dead wood of Abies balsamea, Pinus Strobus,
Acer Negundo, A. spicatum, Betula sp., Carya sp., Juglans cinerea, Phyto-
lacca decandra, Populus tremuloides, Salix sp., Ulmus americana, and
unidentified hardwoods.
Specimens examined: Potanp, 1900, Eichler, type of C. frustulosum (FH-H);
Germany, Lengerich, Westfalen, X.1900, Brinkm. Westf. Pilze 53, as C. fusisporum,
and 155, as C, flavescens (FH); Austria, Fuckel Fung. Rhen. 2396 (herb. Barbey-
Boissier, Geneva) ; New York, Hudson Falls, IX.1.1915, Burnham, type of C. vaga
and C. fenestratum (FH); also specimens from Quebec, Ontario, Manitoba, Maine,
New Hampshire, Massachusetts, Pennsylvania, Florida, Iowa, Missouri.
Pellicularia flavescens is characterized among saprobic species by
clavate-cylindric basidia with usually four large, often finger-like or
ventricose sterigmata, by the germination of its spores by repetition, and
by the broad even- and slightly thick-walled spores with subglobose body
and truncate apiculus. In the earlier treatment, under Botryobasidium,
I distinguished flavescens and ochraceum by form of spores; and it
would be possible to segregate at least one more group on the same basis.
Examination of further material, however, has suggested the propriety
of combining rather than segregating the members of what now appears
to be a single variable species. The subglobose spores characteristic of
ochraceum differ in no respect except the absence of a distal protuberance
from the lemon- or spindle-shaped spores of flavescens; and since the
latter vary widely in the degree of development of this protuberance,
and intermediates between the two types occur, the separation seems
untenable. Coniophora vaga and Corticium fenestratum were in the
earlier discussion assigned to ochraceum because of the characterization
of the spores as merely “apiculate” or “broadly ovoid and rather strongly
apiculate” by Burt and Overholts respectively; but the spores of the
type are what may be called biapiculate—i. e., lemon-shaped — and
Rocers: THE GENUS PELLICULARIA 107
the same as those of C. flavescens sensu Héhnel & Litschauer and Bourdot
& Galzin. On the other hand, the spores of the type of frustulosum are
“fere globosa,” as insisted by Bresadola (1. c., 1911), and that species
is the same as ochraceum. The two, or three, smaller units are easier
to define or to insert in a key; but the one species is the natural group.
10. Pellicularia pruinata (Bres.) Rogers ex Linder, Lloydia 5: 170. 1942.
?Hypochnus coronatus Schroet. in Cohn, Krypt.-Fl. Schles. 3 (1): 418, 1888;
nec H. coronatus Bon., Hedw. 15: 76. 1876.
?Hypochnus Schroeteri Sacc., Syll. Fung. 6: 658. 1888; i. e., H. isabellinus
sensu Schroet. in Cohn, Krypt.-Fl. Schles. 3 (1): 417. 1888; nec H. isabel-
linus Fr., Summ. Veg. Scand. 337. 1849.
?Tomentella granulata Bref., Unters. 8: 11. pl. 1, fig. 15, 16. 1889; nec
Hypochnus granulatus Bon., Handb. 160. 1851.
?Hypochnus Brefeldii Sacc., Syll. Fung. 9: 243. 1891.
Corticium pruinatum Bres., Ann. Myc. 1: 99. 1903.
Corticium coronatum sensu Hohn. & Litsch., Ann. Myc. 4: 291. 1906; Ak. Wiss.
Wien, Math.-Nat. K1., Sitzungsb. 116, I: 832. fig. 15. 1907; Bourd. & Galz.,
Soc. Myc. Fr. Bul. 27: 248, 1911; Hym. Fr. 241. [1928]; an H. coronatus
Schroet.?
Botryobasidium coronatum sensu Donk, Neder. Myc. Ver. Med. 18-20: 117.
1931; Rogers, Univ. Iowa St. N. H. 17: 15. pl. 2, fig. 9. 1935.
?Botryobasidium granulatum (Bref.) Donk, Nederl. Myc. Ver. Med. 18-20:
118. 1931.
Corticium botryoideum Overh., Mycologia 26: 510. pl. 55, fig. 10. 1934.
Corticium isabellinum sensu Eichl. ex Bres., Ann. Myc. 1: 98. 1903; nec C.
isabellinum (Fr.) Fr., Hym. Eur. 660. 1874.
Fructification at first mucedinoid, under the lens arachnoid and minute-
ly tufted, becoming tufted-floccose or finally nearly continuous, lax and
easily separable but scarcely coherent, when fresh pallid or sordid-
whitish, when dry Pale Smoke Gray (R) through Olive Buff (R) and
Cream Buff (R) to Chamois (R); hyphae short-celled, with cruciform
branching, lacking clamps, the repent basal strands 10-18 (—29) yp, often
relatively thick-walled and ochraceous, the subhymenial 5—10 p, colorless,
with symmetrical cymose branching; basidia subcylindric, sometimes
slightly constricted above the middle, short, 14-23 x 7-9 (-10) p, bear-
ing (5~) 6 (-8) divergent, recurved sterigmata 3-5 (—6) p» long; spores
colorless, subglobose or short oblong-ellipsoid, to broad ellipsoid-fusi-
form, more or less attenuated toward the lateral apiculus, distally obtuse,
(3.5-) 4-7 (-7.5) x 3-5 (-6) p.
Mostly on wood or bark of angiosperm hosts, and best developed on
the underside of fallen logs. American specimens on Acer macrophyllum,
A. Negundo, A. rubrum, A. saccharum, Acer sp. indet., Alnus rugosa,
Betula lutea, Carpinus caroliniana, Fagus grandifolia, Kalmia sp., Liquid-
ambar styraciflua, Populus grandidentata, P. tremuloides, Prunus sero-
tina, Prunus sp. indet., Quercus alba, Q. borealis, Q. macrocarpa, Salix
spp., Ulmus sp., Abies balsamea, Pinus Strobus, Pseudotsuga mucronata,
Tsuga canadensis, Fomes pomaceus, indeterminate conifer and angiosperm
wood, and bare soil.
108 Fartowia, VoL. 1, 1943
Specimens examined: PoLanp, type of C. pruinatum (slide, FH-H); Germany,
Baden, im Niederwald bei Rastatt, Schroeter, as H. coronatus (FH-H); Austria,
det. Litschauer as C. coronatum; ENcLAND, det. Wakefield as C. coronatum; New
York, East Galway, VIII.4.1901, Burt, det. Bresadola as ‘C. isabellinum Eichler’,
det. Burt as C. vagum (FH-B); Pennsyzvania, Biglerville, . L. White 1070, herb.
L. O. Overholts 14503, type of C. botryoideum (FH); also specimens from Sweden,
Netherlands, Nova Scotia, Quebec, Ontario, Massachusetts, Rhode Island, Maryland,
Virginia, North Carolina, Ohio, lowa, Missouri, Washington, Oregon.
On the basis of a small number of selected specimens it would be
possible to dismember the group here described into several species. But
when a large number of specimens is compared, whatever lines may have
been tentatively drawn will disappear. The chief difficulty seems to
be that of separating the present species from P. vaga. For the present
study all specimens with distinctly navicular spores were placed with
the latter species, and those with spores both relatively short and quite
obtuse distally were grouped together in P. pruinata. It would require
considerable temerity to assert that this cleavage is a natural one; it is
at least fairly workable. Most collections of pruinata have thicker and
more strongly colored basal hyphae than most collections of vaga; most
spores in pruinata are under 6.5 yw long, while only very rarely are ma-
ture spores of vaga less than 7.5 3 pruinata is most abundant on hard-
woods and vaga on conifers. In the broad-leaved forests of the central
states pruinata is much the more abundant species; in the Northwest,
where vaga is extremely abundant, pruinata is almost entirely lacking;
in the mixed woods of the northeastern region both occur in some abund-
ance. An indication of distinctness, of considerable importance but little
use in determining specimens, lies in the imperfect stage. Three fructi-
fications already assigned to P. pruinata were discovered to be associated
with Oidium candicans (Sacc.) Linder, Lloydia 5: 183. 1942; in two of
these I found by microscopic examination that the hyphae giving rise
to the conidiophores bore also basidial tufts, and the connection is a
fore certain. As noted under P. vaga, that species has another Oidium
for its imperfect stage.
The nomenclature of the present species is more than commonly in-
volved. The accepted names have been those based on Hypochnus cor-
onatus Schroet., untenable from the first because of the earlier, and
quite different, H. coronatus Bon. Furthermore, of the two specimens
listed with Schroeter’s description (1. c.) the second has spores measur-
ing 9-12x 4-5 yw and is no different from C. botryosum Bres., here listed
under P. vaga; whether the first specimen (from Breslau) possessed the
“short elliptic or oval” spores originally described cannot now be de-
termined, since material of it is lacking in the Héhnel collections. The
specimen described by Schroeter as “H. isabellinus Fries 1818 (?)” is
apparently not in existence (cf. Hoéhnel & Litschauer, Ann Myc. 4: 289.).
As Schroeter suspected, and as Saccardo implied in renaming it H. Schroe-
teri, it.is quite probably not the Friesian species. Bresadola’s descrip-
Rocers: THE GENUS PELLICULARIA 109
tion of C. isabellinum (1. c.), and a specimen so determined by him
for Burt, show that for him C. isabellinum sensu Schroet. is the fungus
here under discussion. If this identity could be accepted, the correct
specific epithet for the fungus would be Schroeteri; but in view of the
relative accuracy of Schroeter’s descriptions of associated species there
appears to exist no sufficient reason to disregard nearly all the charac-
teristics which he published (“Haupthyphen bis 9 » breit . . . Basidien-
tragende Hyphen 3-4 p» breit . . . Basidien . . . etwa 5 yp breit, mit
vier Sterigmen”) in deciding the probable identity of his fungus. From
Brefeld’s prolixities (1. c.) it is just possible to surmise that T. granulata
is a Pellicularia; and since it would appear to resemble other known
species rather less than the present one, it is here doubtfully listed as a
synonym. Bresadola’s description of C. pruinatum applies well to the
species under discussion; a slide of the type, in the Héhnel collection,
shows hyphae such as belong to either C. coronatum sensu Hohn. &
Litsch. or P. vaga, but unfortunately no spores. Since the spores of
C. pruinatum as described are rather short for P. vaga, it appears safe
to include Bresadola’s fungus here, and to employ his specific name.
The round-spored extreme described by Overholts has already been
discussed in Univ. Iowa St. N. H. 17: 15.
ll. Pellicularia lembospora sp. nov.
FIGURE 8,
Fructificatio tenuissima, arachnoidea, grisea; hyphae enodulosae, 4.5-8 (-10.5) wu
crassit., angulis rectis ramosae; basidia obovata, 9-10 (-12) x 7-9 yu, sterigmata 6-8
curvula, 2.5-3 yp, longit., gerentia; sporae anguste naviculiformes, ad apiculum oblique
productae, 7-9 x 2.5-3.5 pu.
Fructification delicate and inconspicuous, grayish (Mineral Gray R)
when collected or barely pruinose, under the binocular loose-arachnoid;
hyphae without clamps, branching at right angles, 4.5-8 (—10.5) p, the
basal with walls slightly thickened; basidia borne in small, loose cymes,
subglobose to obovate, 9-10 (—12)x7-9 yp, at the base 44.5 » in
diameter, bearing inside the periphery of the rounded summit 6-8 curved
sterigmata 2.5-3 » long; spores slender-navicular, strongly laterally pro-
duced to the apiculus, narrowed some distance below the summit, straight
or slightly curved on the inner side, convex on the outer, 7-9 x 2.5-3.5 p.
On wood and bark of unidentified dicotyledonous species, and on
Bambusa vulgaris.
Specimens examined (all FH): British Gurana, Bartica, 1.18.24. D. H. Linder
731, type; XII.17.23, D. H. Linder 567. Cusa, Santa Clara Prov.: Blanco’s Woods,
Cienfuegos, VII.1.41, W. L. White 578; Guabairo, Cienfuegos, VII.10.41, VW. L. White
950; Central Soledad, Cienfuegos, VII.8.41, VW. L. White 824; Mina Carlota, Sierra
de San Juan, Trinidad Mts., VII.5.41, VW. L. White 749.
A tropical fungus, approaching P. vaga, but differing in the obpyriform
basidia, short sterigmata, slender spores strongly drawn out at the base,
and slender hyphae. It is more constant in its characters than most
110 Fartowia, VoL. 1, 1943
members of the genus. White’s 824 is associated with, and, as Dr. D. H.
Linder has found, connected with, the imperfect stage, Oidium tomentosum
(Berk. & Curt.) Linder, Lloydia 5: 204, 1942.
12. Pellicularia vaga (Berk. & Curt.) Rogers ex Linder, Lloydia 5: 170. 1942.
FIGURE 9.
Corticium vagum Berk. & Curt.. Grey. 1: 179. 1873; Massee, Linn. Soc. Bot.
Jour. 27: 148. 1890; Coker, El. Mitchell Sc. Soc. Jour, 36: 173. pl. 33,
fig. 9, 10. 1921; Burt, Mo. Bot. Gard. Ann. 5: 128. fig. 3 c, d. 1918; 13:
295. fig. 3 c, d. 1926, quantum ad typum et specim. nonnul.; Bourd. & Galz.,
Hym. Fr. 242. [1928], quantum ad typum, descr. specim. synon. excl.;
Overh., Mycologia 26: 511. pl. 55, fig. 4. 1934,
Corticium botryosum Bres., Ann. Myc. 1: 99. 1903; Hohn. & Litsch., Ak. Wiss.
Wien, Math.-Nat. Kl., Sitzungsb. 116, I: 833. fig. 16. 1907; Wakef., Br.
Myc. Soc. Tr. 4: 117. pl. 3, fig. 15-17. 1913; Bourd. & Galz., Soc. Myc. Fr.
Bul. 27: 248. 1911; Hym. Fr. 241. [1928]; Litsch., Soc. Sc. Skoplje Bul.
18: 176. 1938.
Botryobasidium vagum (Berk. & Curt.) Rogers, Uniy. Iowa St. N. H. 17:
17, 1935.
Fructification at first loosely and delicately arachnoid, becoming
continuous and hypochnoid, sordid whitish or pale buffy, under the
binocular distinctly ascending-tufted, when dry whitish in very young
specimens, or more often Ivory Yellow (R), Cream Buff (R), or rarely
Isabella Color (R); hyphae branching at right angles, (4.5—-) 6-10
(-15) y», the basal relatively long-celled, often yellowish and with bila-
mellate membranes, the basidiophorous colorless, with short and often
barrel-shaped cells, branching to form symmetrical candelabrum-like
cymes; basidia blunt-cylindric, sometimes slightly inflated either toward
the apex or toward the base, 13-22 (—27) x6.5-10 (-15) yp, bearing
rarely 4 or 5, mostly 6-8 stout, divergent, recurved sterigmata (3—)
4.5-6 x 1.5-2 w; spores smooth-walled, colorless, 7.5-12 (—17) x (2.5—)
3.5-5 (—5.5) mw fusoid, obliquely tapered to the apiculus, only slightly
curved on the inside, more strongly curved to strongly ventricose on
the outside, slightly or strongly tapered to the apex.
In America occasionally on humus or on rotting herbaceous stems
(Pteridium latiusculum), commonly on wood or bark of coniferous or
broad-leaved trees: Abies balsamea, A. magnifica, Larix occidentalis,
Libocedrus decurrens, Picea canadensis, P. sitchensis, Pinus contorta,
P. rigida, P. Strobus, Pseudotsuga mucronata, Taxodium distichum, Tsuga
canadensis, Alnus rubra, Betula alba, Castanea dentata, Fagus grandi-
folia, Kalmia latifolia, Populus balsamifera, Quercus spp., Salix sp.,
Tilia americana, and many other undetermined species.
Specimens examined: Poranp, III.1902, Eichler, type of C. botryosum (FH-B) ;
Germany, Glatz, Grunwalder Thal bei Reinerz, Schroeter, paratype of Hypochnus
coronatus Schroet. (FH-H); Sourn Carouina, Curtis, type of C. vagum (IFH-C) ;
also specimens from Netherlands, Austria, Nicaragua, Alaska, Nova Scotia, Quebec,
Ontario, British Columbia, Massachusetts, Rhode Island, New York, Pennsylvania,
Tennessee, Ohio, Wisconsin, Iowa, Missouri, Idaho, Washington, Oregon, and Cali-
fornia.
Rocers: THE GENUS PELLICULARIA lll
Pellicularia vaga is a taxonomic grouping rendered extremely difficult
both by its own variability and by the confusion introduced with the
inclusion, by Burt and others, of the common parasite P. filamentosa.
There is no important variation in gross appearance, in hyphae, or, ex-
cept for dimensions, in basidia; but the forms of the spores are such
that one who had before him only isolated extremes might well wish
to describe for them two, or even three or four, species. The spores of
all specimens here included in P. vaga are in some degree asymmetrically
fusiform — that is, what is called navicular. But besides variations in
absolute dimensions, this material shows great differences in the relation
of breadth to length; and some spores are decidedly obtuse, tapering
only a little from the broadest portion to the apex, while others are
apically cuneate. The extremes in form are to be characterized as
ventricose-navicular — that is, the ‘little boat’ with which the spores
are compared would have to be one with a fairly deep keel — and fusoid-
subcylindric — that is, narrow throughout, and only a little more so
at the ends than at the middle. A fresh preparation from the type of
C. vagum shows abundant spores 9—10.5 x 3-4—4.5 yp, subcylindric-navic-
ular, and near the slender extreme; it is matched by scattered collec-
tions from Ontario and the eastern states and by the Alaskan and many
Oregon specimens (fig. 9 i-j). The type of C. botryosum (fig. 9 a—d)
is near the other extreme, with spores (8—) 9-11.5x (4-) 4.5-5 w and
strongly tapered; it is matched by European and a good deal of eastern
American material. The intergradation in the whole series at hand is,
however, such that the separation of the group into taxonomic subdi-
visions seems unwarranted. Probably segregation on the basis of degree
of taper towards the ends of the spore would yield more usable lines of
separation than segregation on relative breadth; but it does not seem
probable that either would define natural — that is, phyletically divergent
— segregates.
Litschauer has written (1. c.) that C. botryosum is “not identical with
C. vagum Berk. & Curt.” There is no indication in his note whether he
desired to maintain separate species for the two extremes here de-
scribed, or whether he was still misled, as formerly (cf. Rogers,
l. c.), by Burt’s interpretation of C. vagum. As Bresadola observed in a
note to Burt (of January 18, 1920) attached to a sheet in the latter’s
herbarium, C. vagum sensu Burt included C. botryosum Bres., the form
subsequently described as C. botryoideum Overh., and C. Solani
(Prill. Del.) Bourd. & Galz. Bourdot’s description and a specimen from
Litschauer show that C. vagum as they had learned it through Burt bears
no resemblance either to C. botryosum or to the fungus of Berkeley
and Curtis.
In material at hand I have found basidia connected by unbroken series
of mycelial cells with the conidiophores of the imperfect stage, Oidium
Curtisit (Berk.) Linder, Lloydia 5: 201. 1942,
HZ FartowiA, VoL. 1, 1943
13. Pellicularia Koleroga Cooke, Grey. 4: 116, 134. 1876; Pop. Sc. Rev. 15:
164-167. pl. 135, fig. a-c. 1876; Linn. Soc. Bot. Jour. 18: 461, 463. 1881;
Fawcett, Jour. Ag. Res. 2: 231. fig. 1-3. 1914.
Erysiphe scandens Ernst, Estudios sobre las deformaciones . . . del arbol de
cafe en Venezuela. 16. fig. 5. 1878.
Hypochnus ochroleucus Noack apud Sacc., Syll, Fung. 16: 197. 1902; Stevens
& Hall, Ann. Myc. 7: 53. fig. 5-8. 1909.
Corticium Koleroga (Cooke) Hohn., Ak. Wiss. Wien. Math.-Nat. KI., Sitz-
ungsber. 119, I: 395. 1910; Burt, Mo. Bot. Gard. Ann. 5: 123. fig. 1. 1918;
13: 292. fig. 1. 1926; Wolf & Bach, Phytopath. 17: 710. fig. 10. 1927.
Corticium ochroleucum (Noack) Burt ex Peltier, Univ. Ill. Ag. Exp. Sta. Bul.
189: 290. 1915 (falso ut ‘ochraleucum’); nec. C. ochroleucum (Fr.) Fr.,
Epicr, 557. 1838.
Corticium Stevensii Burt, Mo. Bot. Gard. Ann. 5: 125. fig. 2. 1918; 13: 293.
fig. 2. 1926; Coker, El. Mitchell Sc. Soc. Jour. 36: 174. 1921.
Hypochnopsis ochroleuca Noack, Inst. Agron. Sao Paulo, Campinas, Bol. 9:
80. 1898 (mycelium sterile).
FIGURE 10.
Fructification a delicate, separable, subhypochnoid pellicle, sordid or
buffy; hyphae branching at right angles, without clamps, 4-12 » in
diameter, the basal mostly 5—8, or occasionally up to 10 p», long-celled,
little branched, sometimes colored, the subhymenial shorter-celled and
abundantly branched, rigid or early collapsed, bearing basidia in very
small and irregular cymose-branching clusters, the subbasidial cell often
thicker than adjacent hyphal cells and widened outward; basidia obpyri-
form, short-barrel-shaped, or rarely short-clavate and constricted (ap-
parently from failure of a septum to form at the constriction), 11.5-15
(-19.5) x8.5-10 », bearing 4 horn-like sterigmata 6-8 x 2-3 y; spores
very slightly thick-walled, subcylindric-fusiform to elongate subellipsoid,
in either case tapered to both ends, flattened or depressed on the inside
just above the apiculus, distinctly and laterally apiculate, 10.5-16 x 3-5.5
(—7) pe
On underside of living leaves of Citrus sinensis, Coffea sp., Diospyros
virginiana, Pittosporum sp., and Pyrus communis.
Specimens examined: Argentina, Misiones, Fawcett & Bitancourt (SP 2626) ;
Venezuela, Caracas, X. 1923, Pittier (BPI): Puerto Rico, Mayaguez, V.22.17, Thomas
(FH); North Carolina, Mt. Airy, IX.2.07, Reimer, det. Stevens as H. ochroleucus,
det. Burt as C. Stevensii (FH-B), a sterile specimen (sclerotial) with same data
(FH-B), and Indian Cr., Hesler (TENN 14344); South Carolina, Summerville,
VII.12.41, Jahnz (BPI).
Distinguished from P. filamentosa by the elongate spores with a
slightly but distinctly thickened and refractive membrane and with nar-
rower and thick-walled apiculus. The basidia also are usually distinctive,
tending toward the pyriform rather than short-clavate. In the small
amount of material available there is considerable variation in the form
of the spores. In the Puerto Rican collection, on coffee, they are slender
and almost fusiform, and so strongly depressed as to be slightly curved
(fig. 10 a—d); in the Venezuelan collection, likewise on coffee, they are
Rocers: THE GENUS PELLICULARIA 113
shorter and broader; in the South Carolina collection and the Mt. Airy
collection (fig. 10 e-f) they are still broader and more nearly ellipsoid.
There does not, however, appear to be sufficient basis for division into
two species, and the treatment of Wolf & Bach, who combined C. Stevensii
with C. Koleroga, is here adopted. Noack (in Saccardo, I. c.) described
the spores of his fungus as being “ovoideis . . . 812-10 x5%-6”; it may
be that his Hypochnus ochroleucus was not the same fungus as the one
described under that name by Stevens & Hall. In that event, H. ochro-
leucus, as well as C. ochroleucum Burt and C. Stevensti, which strictly
are based on his fungus, must be deleted from the synonymy here given;
but the nomenclature for the specimens here cited is not to be altered
on that account.
Amateurs of Lloyd’s “Myths of Mycology” will do well to compare
the present fungus, or description, with Cooke’s several accounts of its
structure and affinities (ll. cc.).
14. Pelliculatia filamentosa (Pat.) comb. nov.
Hypochnus filamentosus Pat., Soc. Myc. Fr. Bul. 7: 163. pl. 11, fig. 2. 1891
(30 Sept.) ; nec H. filamentosus Burt, Mo. Bot. Gard. Ann. 13: 320. 1926
(= Corticium sulphureum (Pers. ex Fr.) Fr.).
Hypochnus Solani Prill. & Del., Soc. Myc. Fr. Bul. 7: 220. fig. 1891 (31 Dec.) ;
Miller, Biol. Reichsanstalt Land- u. Forstwiss. Arb, 13: 205-8. pl. 1, 2. 1924;
et alibi.
Corticium vagum var. Solani Burt ex Rolfs, Science n. s. 18: 729. 1903.
Corticium Solani (Prill. & Del.) Bourd. & Galz., Soc. Myc. Fr. Bul, 27: 248.
1911.
Corticium vagum sensu Burt, Mo. Bot. Gard. Ann. 5: 128. fig. 3a. 1918, pro
parte; 13: 295. fig. 3a. 1926, p. p.; Petch, R. Bot. Gard. Peradeniya Jour.
7: 288. 1922 (falso ut ‘C. vagans Berk. & Curt.’); et Auctt. plur.; nec C.
vagum Berk. & Curt.
Corticium vagum subsp. Solani (Prill. & Del.) Bourd. & Galz., Hym. Fr. 242.
[1928].
Botryobasidium Solani (Prill. & Del.) Donk, Nederl. Myc. Ver. Med. 18-20:
117. 1931; Rogers, Univ. Iowa St. N. H. 17: 18. 1935.
Corticium areolatum Stahel, Phytopath. 30: 129. fig. 3, 4. 1940; nec C. areo-
latum Bres.. Mycologia 17: 68. 1925.
Oidium Citri Bondar, Lab. Path. Veg. Bahia Bol. 7: 78, fig. 36b. 1929, nomen
nudum.
Corticium microsclerotia Weber, Phytopath. 29: 565. fig. 4, 7. 1939, nomen
nudum.
FIGURE LI.
Fructification forming a delicate separable, flaky or thin-hypochnoid
pellicle, when dry white to buffy, or semitranslucent; hyphae branching
at right angles and with some cruciform cells, without clamps, 4.5-14
(-17) » in diameter, the basal long-celled and often somewhat colored,
the subbasidial colorless, short-celled, with often barrel-shaped segments,
branching abundantly and bearing the basidia in small imperfectly
symmetrical cymes; basidia subcylindric and barrel-shaped (widest
toward the middle) or obpyriform or clavate (widest at the summit),
(10-) 12-18 (-23) x 8-11 (-12.5) yw, bearing 4 sterigmata which
114 Fartowia, VoL. 1, 1943
arise as blunt knobs and become later horn-shaped, (3—) 5.5-12 (—20) x
1,5-3.5 (4.5) mw; spores ellipsoid or oblong-ellipsoid, thin-walled, flat-
tened on the inside, a little the broadest below the middle, truncate-
apiculate, 7-12.5x 4~—7 p, occasionally germinating by a stout promycel-
ium on which is borne a similar secondary spore.
On surface of living leaves or stems of Boerhavia erecta, Chimaphila
maculata, “sour orange seedlings” (? Citrus Aurantium), C. grandis c.
Marsh, C. sinensis c. Bahia, Dianthus Caryophyllus, Ficus Carica, Hydro-
phyllum virginicum, Nicotiana tabacum, Solanum Dulcamara, S. tuber-
osum, and an indeterminate member of the Amaryllidaceae.
Specimens examined: Denmark, Lyngby, Sjaelland, VI.21.10, J. Lind, as H. Solani
(FH); Brazil, Bondar (SP 725) and Fawcett & Bitancourt (SP 2499, 2516), all as
C. areolatum; Argentina, Fawcett & Bitancourt (SP 2632), as C. areolatum; Ecuador,
de Lagerheim, type and paratype of H. filamentosus Pat.; Panama, Fawcett (SA 149) ;
Nova Scotia (OTB 10326); Florida, Gainesville, [X.10.41, Weber, as C. micro-
sclerotia; also specimens from Ontario, Maine, Massachusetts, New York, and Iowa.
Distinguishable from the saprobic species by the flattened-ellipsoid
spores with a peculiar truncate, almost jagged, apiculus, by the four
usually long and thick sterigmata (or epibasidia; the evidence is not
clear), and by the small and usually imperfect cymes of basidia. The
spores of P. flavescens, which in some respects approaches the present
species, are subglobose or have a subglobose body with apical prolonga-
tion, and the basidiocarp is usually much thicker and more hypochnoid.
Obviously the substratum — parts of living plants, or occasionally soil
in contact with them — is the easiest identifying character, and it seems
to be a reliable one; during the present study of a large amount of
saprobic material no specimen has been encountered on another sub-
stratum which could be taken for P. filamentosa, nor have the typically
saprobic species ever been seen on living plant-parts. Leaving aside
the matter of substratum, however, the present species is readily dis-
tinguishable, as noted, by morphology alone.
The diseases associated with the various fungus names here listed in
synonymy seem distinct enough, but the fungi, at least in the material
at hand, show relatively slight variation. C. areolatum is the smallest
of the lot, with spores 7.59 x 4.5—5 yp, and correspondingly small hyphae
and basidia (fig. 11 e-g); the sterigmata, as throughout the present
species, are enormously variable within a single collection, and mature
ones run from 5.5 to 10 » in length and about 2.5 (or in other specimens
only 1-1.5) » in diameter, in contrast to the very large ones shown by
Stahel’s material (1. c.). A specimen forming a mildew-like growth on
sour-orange seedlings (from Panama), with no suggestion of the areolate
leaf-spot, is indistinguishable, as is a Massachusetts specimen on Chima-
phila. Corticium microclerotia (fig. 11 h-i) shows spores 8.5-9.5 x5 p,
and small basidia and long sterigmata (7-9 x 1.5-2 ,») similiar to those
of C. areolatum. The type specimen of H. filamentosus lies near the
upper end of the size-range; it is quite indistinguishable from material
Rocers: THE GENUS PELLICULARIA Ls
. examined growing on potato (fig. 11 a-d). Corticium praticola Kotila,
the species next discussed (q. v.) may belong with it. The similarity
of H. filamentosus and H. Solani would seem sufficiently apparent from
the illustrations, published (ll. cc.) in successive numbers of the same
journal; it is a wonder that Prillieux & Delacroix’s account was ever
allowed to appear, and a greater wonder that their name was not at
once reduced to synonymy. Differences in presence or absence of
sclerotia, and, a fortiori, in time or place of their formation, or in their
size, must be regarded as untrustworthy — at least, until comparative
studies have been further carried out. Among mycelia isolated from
the pathologically highly uniform and characteristic areolate leaf-spot of
Citrus, Stahel (1. c.) found some which formed sclerotia readily and
others which would under no conditions do so.
The sclerotial stage of P. filamentosa has been described under the
names Rhizoctonia Solani Kuhn, Krankh. Kulturgew. 224. 1858, and
R. microsclerotia Matz, Phytopath. 7: 116. fig. 3; pl. 2. 1917. The de-
scription of Corticium microsclerotia refers to that of Rhizoctonia micro-
sclerotia; but the earlier description, dealing with only the imperfect
stage, cannot form the basis for a basidiomycetous species. Since the
account of C. microsclerotia lacks the validating Latin diagnosis, Weber’s
name must technically be accounted a nomen nudum. Corticium areo-
latum Stahel is also invalidated, by the existence of the earlier homonym-
ous species of Bresadola. Bondar’s figure of Oidium Citri probably
represents the present fungus (as suggested by Stahel), and Bondar’s
material probably included basidia; but since the fungus is not described
by Bondar (1. c.), his name has no standing.
There is room for further study of the fungi here treated; obviously
a great deal more is known of the diseases produced by P. filamentosa
than about the fungus itself. It is at least arguable that on different
hosts and under different circumstances a single pathogen may give rise
to different pathological phenomena. It would appear preferable, then,
to maintain whatever differentiating terminology is needed for the various
diseases, but to admit taxonomic segregation within the pathogen only
when significant differences have been shown to exist between groups
of the fungi themselves. Further study may demonstrate such differ-
ences, and the necessity of subdividing P. filamentosa; they have not thus
far been shown, nor has the present study revealed them.
Since most of the literature concerning “C. vagum” and H. Solani as
a parasite is concerned with the host rather than with the fungus, it has
not been cited here.
SPECIES INQUIRENDAE
15. Corticium praticola Kotila, Phytopath. 19: 1065. fig. 5, 6. 1929.
“Hymenium loose, in minute placques formed by a more or less
dichotomous branching of the vegetative hyphae. Terminal cells develop
116 FarLowia, VoL. 1, 1943
into basidia bearing 1 to 4 sterigmata, the usual number being 3. Basidia
average 15.6 » long and 6.5 » wide. Sterigmata vary from 13 to 26.5
or more » long, average 18.8 », and taper gradually upward from point
of attachment to basidium, greatest diameter 2.6 p. Basidiospores
hyaline, ovate, apiculate, smooth-wall, from 5.2 to 7.8 » long and from
4.9 to 5.5 w wide, with average dimensions of 7.7 by 5.2 ».”” — Kotila, 1. c.
A parasitic form characterized by, among other things, extremely long
sterigmata (? epibasidia) and spores germinating by repetition. Like
similar material of P. filamentosa (from which it is not certainly dis-
tinguishable), and like P. flavescens, it might be included in either Pelli-
cularia or Ceratobasidium — in the former because of clear affinity, in
the latter by definition. The fungus was made the subject of intensive
study by Kotila; after examination of an excellently preserved portion
of his type material I find nothing to add to either his description or
his illustrations (1. c.). Because that material grew in a Petri dish, it is
not wholly comparable with specimens —e. g., of P. filamentosa — col-
lected in the field. Because the abundant basidia and spores now present
in it are now tightly stuck to the Petri dish cover, it seems possible that
their development was modified by water condensed there. I therefore
prefer to suspend judgment on the problems of its generic position and
its relation to P. filamentosa.
16. Corticium album Dastur, Indian Jour. Agr. Sc. 10: 92. pl. 1. 1940, nomen nudum.
From the plate cited and from two slides of the type it appears highly
probable that this fungus is a member of the genus Pellicularia. The
basidia are longer than in most species of the genus and apparently dis-
tinctly clavate, in both characters resembling those of P. flavescens.
Neither the figures, however, nor the sections provide a sufficient basis
for redescription or safe disposition of the fungus. As pointed out by
Mundkur (Current Sc. 9: 284. 1940), Dastur’s name is a later homonym
of C. album Britz., Bot. Centr. 71: 91. 1897— or would be except that
it lacks a Latin diagnosis, and hence was never validly published.
SPECIES EXCLUDENDAE
Four species included by Bourdot & Galzin in Corticium sect. Botryodea
are here excluded from Pellicularia:
C. sterigmaticum Bourd. = Ceratobasidium sterigmaticum (Bourd.) Rogers, Univ.
Iowa St. N. H. 17: 7. 1935.
C. cornigerum Bourd. = Ceratobasidium cornigerum (Bourd.) Rogers, Univ. Iowa
St. N. H. 17: 5. 1935.
C. terrigenum Bres. is probably likewise a Ceratobasidium; since no one appears
to have seen it since Bresadola, judgment must be suspended.
Hypochnella violacea (Auersw. in sched.) Schroet. has much in common with
Pellicularia and also, as pointed out by Martin (Iowa Ac. Sc. Trans. 44: 47.
[1938].) with Coniophora. At least until a comparative study can be made
of species of the latter genus, it seems best not to include it in Pellicularia.
DEPARTMENT OF BIOLOGY
AMERICAN INTERNATIONAL COLLEGE
SPRINGFIELD, MASSACHUSETTS
Rocers: THE GENUS PELLICULARIA lig iy;
Fig. 1-6. 1, Pellicularia chordulata: a-c, type; d-f, DPR 948. 2, P. asperula:
type. 3, P. Langloisii: Thaxter 57. 4, P. cystidiata: type. 5, P. ochroleuca: DPR
702 (Massachusetts). 6, P. ansosa: type.
118 Fartowia, Vou. 1, 1943
Fig. 7-11. 7, Pellicularia subcoronata: a-b. DPR 990 (Rhode Island) ; c—d,
DPR 989 (Massachusetts). 8, P. lembospora: a-c, type; d-f, DHL 567. a ge
vaga: a-d, type of C. botryosum; e-h, DPR 988 (Rhode Island); i-j, DPR 987
(Oregon). 10, P. Koleroga: a-d, Thomas; e-f, Reimer. 11. P. filamentosa: a—d,
OTB 10326; e-g, SP 725; h-i, Weber.
1(1): 119-133 FARLOWIA January, 1943
SEVERAL FUNGICOLOUS FUNGI?
Epcar V. SEELER, JR.
A small discoid parasite on Trichoglossum collected by the author
proved to be Hypomyces Geoglossi Ell, & Ev. which had been transferred
later to Eleutheromyces. Examination of authentic specimens uncovered
the confusion in herbaria and in the literature concerning it and an asso-
ciated group of species which grow on living or on decaying fungi. From
a study of a few of these species have come these notes. Included are
the following: a discussion and emendation of the genus Eleutheromyces
and a redescription of its type species, a redescription from the type of
Eleutheromycella mycophila von Héhnel, the first full description of
Sphaeronema blepharistoma Berk. and the establishing of it by priority
as the type of Hyalopycnis von Héhnel, the erection of the new genus
Micropyxis based on Hypomyces Geoglossi Ell. & Ev., and finally an
emendation of Sphaeronaemella Karsten and of its type species S. Hel-
vellae. All species discussed have been illustrated as completely as pos-
sible by camera lucida drawings. Abbreviations of herbarium names are
those recommended by Lanjouw in Chronica Botanica 5: 142-150. 1939.
Citations pertinent to synonymy and discussion are listed below each
name considered.
ELEUTHEROMYCES Fuckel. Symb. Myc. p. 183, pl. 4, fig. 52a—c. 1869.
Fuckel based this genus on Sphaeronema subulatum Fries which he
stated was an Ascomycete with /saria brachiata Schiim. as its conidial
phase. The type, or cotype, was therefore Fries’ collection distributed
as Scler. Suec. 325, and authentic material for the genus concept (deter-
mined by Fuckel) comprised Fungi Rhenani 773. |
There should have been no confusion as long as these specimens existed
for study and the writer can state with certainty that the fungus which
carried the name of Sphaeronema subulatum is not an Ascomycete but
a member of the Sphaeropsidales. It may prove to have an ascophorous
form, but none has appeared in the five collections from Europe and
thirteen from North America examined here. It should be pointed out
that all who claim to have seen asci have recorded the ciliate spores
(typical pycnospores) but have completely failed to record or recognize
any method other than asci for bearing these spores. While there have
been no asci ever observed in the present study there have been the
same pycnophores (PI. 1, fig. 2) clearly visible with spores attached in
every fructification examined. Perhaps the fact that Fuckel, and those
who later “saw” asci, entirely missed these ever-present pycnophores is
the most important evidence that, after Fuckel, they were playing follow
the leader. Each case will be considered in detail. Citations will be
found with the list of synonyms under the species E. swbulatus.
1 Contribution from the Laboratories of Cryptogamic Botany and the Farlow Herbar-
ium, Harvard University, no. 220.
119
120 Fartowia, Vou. 1, 1943
Fuckel was the first to report asci “52x 2.5 mic.” and he illustrated
one with a tiny sketch showing eight simple oval spores arranged in a
single row, and in his fig. b a single “endospore” is shown as biterminally
ciliate. He gave Isaria brachiata Schum. as the conidial phase and de-
scribed its conidia as “ciliate at both ends or nude.” Conidia from
Fuckel’s collection (sub E. subulatus) ex herb. Barbey-Boissier (FH)
are typical and are never ciliate. The other Fuckel specimen examined,
Fungi rhenani 773 (FH), exhibits the following fungi: Sphaeronema
subulatum Fr., Cryptophaeella Heteropatellae von Hohnel, and Isaria
brachiata Schiim. In 1892 Ellis devoted six figures to this species which
are the clearest illustrations to date; they all represent Sphaeronema
subulatum Fr. except for the fact that he has drawn a heavy black line
around eight appendiculate spores, the “mature ascus,” and he has shown
“peculiarly jointed paraphyses.” These “paraphyses” are typical
pycnophores of S. swbulatum Fr., a fact which Ellis obviously did not
recognize; and his “ascus” being surrounded with them as depicted would
indicate that asci grow among pycnophores. Specimens collected and
determined by Ellis as Elewth. subulatus have revealed no asci since Ellis
saw them, and all are typical S. subulatum Fr. Saccardo (1879) reported:
asci 8-spored, 50x 4 yp, spores 2-rowed, 5-6 x 1.75-2.5 p, often associated
with Isaria brachiata; the specimen was from Italy, collected by Spe-
gazzini. In 1883 Saccardo added the measurements: perithecia 1.5 mm.
high x 0.3 mm. broad, asci 48-52 x 2.5-3 p, spores “submonostichus,”
4—6x1.5 ». In a section of Rabh. Krypt. Fl. published the next year,
1884, Winter copied Saccardo’s data and Fuckel’s drawing. Later (1905)
as a rebuttal to von Hohnel, Saccardo quoted himself as having seen
asci in 1879; and in 1913, still answering von Hohnel, he refers to
“Winter, Schroter, Ell. & Ev., etc.” as evidence for the ascus; here he
suggests the possibility of separate perithecial and pycnidial forms —
whereas Ellis’ picture would indicate both in one. Jaczewski in his
monograph of Sphaeronema passed this species easily saying, “the asci
are so ephemeral that one never finds them in old specimens, but the
form of the spores is so characteristic as not to permit error.” Seaver
(1909) saw and drew the ciliate spores and assumed that they came
from asci as he noticed no other source. In combatting this ascus theory,
it should again be emphasized that no one of the men who described
asci has mentioned the important feature so conspicuous in his own
specimens, pycnophores bearing spores.
On the other side of the ledger, there are those who have searched
carefully for asci and failed to find them in authentic specimens left us
as evidence for the writings of earlier workers. These include: von
Hohnel (1902), Petrak and Sydow (1923), Petch (1935), and Grove
(1937). Von Héhnel and Petrak and Sydow saw the pycnophores and
said definitely that they thought that the ascus phase did not occur as
it had been described. And Petch (Shear to the contrary notwithstand-
SEELER: FUNGICOLOUS FUNGI 121
ing) hesitated to believe that Fuckel’s ascigerous form existed. The
present author’s findings are discussed under the type species Eleuth.
subulatus.
Finally there are Shear’s most recent (1940) and convincing argu-
ments for the retention of Eleutheromyces as an Ascomycete genus. These
are based entirely on the literature, since Shear himself states that his
packet of the cotype, Scler. Suec. 325, contained “no fructifications of a
Sphaeronaema,” and he does not mention having examined any other
specimens, not even Fungi rhenani 773 which Fuckel distributed.
It is hoped that from the above it has been shown that modern experi-
mental evidence taken from authentic specimens from two continents
has failed to support the presence of asci in this species. Petrak and
Sydow have given what the writer believes to be the correct interpreta-
tion of the situation: (translating from their German) “The circumstance
that several authors claim to have found asci of this fungus and also have
pictured such may easily be explained by the fact that the conidia, strongly
glued together by slime, hang together in shorter or longer rows, often
more or less parallel to each other. Such rows of spores were probably
regarded as asci.”
Petrak and Sydow, and Shear have shown that Sphaeronaema should
be reserved for the black sphaeropsidaceous forms based on S. cylindri-
cum (Tode) Fr., and should not include nectrioidaceous species. Grove’s
1937 combination is no longer tenable because Karsten’s Sphaeronaemella
had for its type S. Helvellae which is here shown to be an Ascomycete.
The von Hoéhnel genus Eleutheromycella seems related but as shown
elsewhere in this paper must remain distinct generically according to
present keys and systems of classification which are based on pycnidial
color. Therefore since Eleutheromyces as an Ascomycete was based on
confusion and misinterpretation of two unrelated Fungi Imperfecti, and
yet since the name is firmly associated in the literature and in herbaria
with its type, Sphaeronema subulatum Fr., and as the name does suggest
the apparent relationship to Eleutheromycella von Héhnel, I propose to
emend it as follows:
ELEUTHEROMYCES Fuckel, emend. Seeler.
(Sphaeropsidales, Nectrioidaceae)
Pycnidia single, superficial, light-colored, soft-leathery or gelatinous
and translucent when wet, walls and neck pseudoparenchymatous, conoid
with upper portion more or less extended into a tapering neck, internally
lined up to the mouth with simple or more often branched hyaline
conidiophores. Spores hyaline, typically one-celled and ellipsoid, apical-
ly ciliate, basally attenuate.
Type species: Eleutheromyces subulatus (Fr.) Fuckel, emend. Seeler.
Of the two other species which have been placed in this genus, the
first, E. longisporus Phill. & Plowr. (Grevillea 13: 78. 1885) has been
122 FARLOWIA, VoL. 1, 1943
transferred to Rhynchonectria by von Héhnel (Sitzb. Akad. Wiss. Wien
111: 1023. 1902) and has been carefully described and illustrated by
Grove (Journ. Bot. (London) 45: 171, pl. 485, fig. 3a—e. 1907). The
second, E. Geoglossi (E. & E.) Seaver is discussed elsewhere in this paper
under Micropyxis.
Eleutheromyces subulatus (Fr.) Fuckel, emend. Seeler.
? Sphaeria subulata Tode, Fungi Mecklenb. 2: 40, pl. 15, fig. 117a—e. 1791.
Sphaeronema subulatum Fries, Syst. Myc. 2: 536. 1822.— Petrak & Sydow, Ann.
Mycol. 21: 361. 1923.— Shear, Mycologia 32: 541-544, 1940.
Sphaeromyxa subulatum (Fr.) Sprengel, in Linn. Syst. Veg. (16 ed.) 4: 406. 1827.
Zythia subulata (Fr.) Schweinitz, Syn. Fung. Amer. Bor. p. 247, no. 2128. 1832.
Zythia compressa Schweinitz, Syn. Fung. Amer. Bor. p. 247, no. 2130. 1832.
Sphaeronema oxyspora Berkeley, “Decades of Fungi no. 136,” Journ. Bot. (Lon-
don) 6: 325. 1847.
Eleutheromyces subulatus (Fr.) Fuckel, Symb. Mycol. p. 183. 1869.— Sacc.,
Michelia 1: 50. 1879; Syll. Fung. 2: 455. 1883; 17: 779, 1905; 22: 1142, 1913.
— Winter, in Rabh. Krypt. Fl. 2: 93, 4 fig. p. 84. 1884. — Ell, & Ev., N. Amer.
Pyren. p. 93, pl. 14, fig. 6-12. 1892.— Jaczewski, “Monographie du genre
Sphaeronema Fr.”, Nouv. Mem. Soc. Imp. Nat. Moscou 15: 336. 1898.— von
Hohnel, “Fragm. Myk. no. 32,” Sitzb. Akad. Wiss. Wien 111: 1022. 1902. —
Seaver. Mycologia 1: 47, pl. 4, fig. 12-13. 1909. — Petch, Journ. Bot (London)
73: 186. 1935.— Shear, Mycologia 32: 546. 1940.
Sphaeronaemella oxyspora (Berk.) Saccardo, Syll. Fung. 3: 618. 1884.
Sphaeronaemella subulata (Fr.) Grove, Brit. Stem & Leaf Fungi 2: 115-116, fig.
54a—c. 1937.
PLATE I, Fic. 1-6.
Pycnidia discrete, clustered or scattered, when dry brittle and approxi-
mately “Raw Sienna” (Ridgway PI. III), when wet translucent and gela-
tinous, often with flecks or threads of dark brown on the basal portion,
extremely variable in size and shape, diameter of base 130-350 yp,
over-all height 220-3300 y, essentially conical, usually tapering grad-
ually upward, walls pseudoparenchymic of thick-walled polygonal cells
up to the ostiole which is fimbriate but usually hidden in a globular
mass of spores or entirely broken away. Spores borne on fine long
sterigmata which are drawn out from the apex or just below the nodes
of hyaline, simple or branched, septate conidiophores (1.5—3.5 » diam.)
which completely line the pycnidial cavity and neck canal. Spores not
catenate, hyaline, one-celled, ellipsoid body (4-5.5x1.5-2 p») with
attenuated filiform pedicel (2-3x 0.5 ») and apical seta (3-7x 0.5 yp).
Growing on various decaying agarics, boletes, polypores, and other
fleshy Basidiomycetes. Recorded range: North America; England to
eastern Russia; Italy to Scandinavia.
Type: Some specimens of Scleromyceti Sueciae 325 distributed by
Fries as seen by Petrak and Sydow (1. c.). This was a mixed collection
apparently, as the specimen in the Farlow Herbarium shows only Didymos-
tilbe sp., and yet the specimens examined prove that there is a well-
founded international concurrence in the concept of this common species.
SEELER: FUNGICOLOUS FUNGI 123
Specimens examined: (all in FH unless otherwise indicated) Ontario, Cain, TRT
4859; New Hampsuire, Shelburne, Farlow.— Chocorua, on Polyporus betulinus,
Sept. 1906, Farlow. — Chocorua, on Lenzites sp., Aug. 1913, Farlow. — VERMONT,
L. Dunmore, Sept. 1896, Farlow; MassacHusetts. Newton, Oct. 1878, Farlow. —
Waltham, Nov. 1885, Thaxter; Connecticut, New Haven, Sept. 1888, Thaxter ;
PENNSYLVANIA, ex herb. Schweinitz in herb. Curtis 2128.— Bethlehem, on Polyporus
frondosus, type (?) of Zythia compressa Schw., ex herb. Schw. in herb. Curtis;
Ouro, type (?) of Sphn. oxyspora Berk., in herb. Curtis. — Chillicothe, Aug. 1906,
Kellerman, det. Peck (CU 21356) ; SoutH Caro.ina, Soc. Hill, Dec., in herb, Curtis;
FINLAND, Karsten, Fungi Fenn. 370; Germany, Klotsch, Herb. Viv. Myc. 1463. —
Fuckel, Fungi Rhen. 773.— Fuckel, ex herb. Barbey-Boissier.— Rabenhorst, Fungi
Eur, 1334.
In order to bring together all pertinent information, this species has
been illustrated and redescribed here. Among the listed synonyms Tode’s
name is marked “?” even though Fries credited the species to him. Since
Tode’s specimen is no longer available for comparison, it is impossible
to be certain today what fungus he meant.
Eleutheromycella mycophila von Hoéhnel, Frag. Myk. nr. 178, Sitzb. Akad. Wiss.
Wien, etc. 117: 1023. 1908.— Grove, Brit. Stem and Leaf Fungi 2: 116. 1937.
— Shear, Mycologia 32: 546. 1940.
Eleutheris, Clements & Shear, Genera of Fungi, p. 371. 1931 (arbitrary change of
name).
PLATE I, Fic. 11-15,
Pyenidia single, parasitic (?), fungicole, submerged or erumpent,
black, smooth, soft-leathery when wet, spherical or deformed by position
of growth in host, those growing in pores being elongated, those under
the tough epidermis being flattened, body in vertical section 250-900 p
wide x 300-510 » high, wall completely pseudoparenchymatous of thin-
walled cells, neck from short-papillate to 150 » long and tubular (only
5 pycnidia measured), ostiole not fimbriate. Spores borne on fine, long
sterigmata which are drawn out from the apices or from just below the
nodes of hyaline, simple or branched, septate conidiophores (1.2—-2.5 p
diam. at base) which completely line the pycnidial cavity. Spores not
catenate, hyaline, typically one-celled, cylindrical-ellipsoid body 5-6.5
(-10) x 1.2-1.5 p», with attenuated filiform pedicel 2-3 » long and an
apical seta 12-30 x 0.5 yp.
Growing, apparently parasitically, in sporophores of Polystictus versi-
color (L.) Fr.; pycnidia growing in pores of the host open downwards,
those embedded in trama open upwards with necks protruding through
the surface.
Type: Herb. v. Héhnel sheet 10203 (FH-H), Georgenberg, Purkersdorf,
Wiener-Wald, Austria, Aug. 2, 1907, collected by von Héhnel.
This is known only from the type specimen which is also the type of
the genus. It is strikingly like Eleutheromyces suwbulatus emend. in
the character of spores and conidiophores but differs from it in the black,
short pycnidia and the longer more slender spores. It is hoped that
the figures and this description from the type will eliminate some con-
fusion evident in the literature.
124 Fartowlia, VoL. 1, 1943
Hyalopyenis blepharistoma (Berk.) Seeler, comb. nov.
Sphaeronema blepharistoma Berkeley, “Notices of British Fungi no. 57” in Mag.
Hist. 6: 363. 1841.
Hyalopycnis vitrea von Hohnel (non Corda?), Hedwigia 60: 151-154. 1918.
Hyalopycnis hyalina v. H., Hedwigia 60: 151-154. 1918.
PLATE I, Fic. 7-10.
Pycnidia usually growing fairly close together, when dry pale yellow-
ish, transparent, “glassy,” amber, often collapsed on one another in a
confused tangle cemented by the spores, rarely standing singly with
erect necks; when wet nearly hyaline with milky opaque spore-mass
within, soft-gelatinous, size and shape variable (300-600 y over-all
height), a short, broad basal trunk below the swollen body (100-200 p»
diam.), tapering gradually into the neck (30-70 p» diam.) and flaring
again at the plume-like apex by the out-curving of the two or three ranks
of delicate, parallel, scarcely-septate filaments which make up the peri-
thecial wall and neck. Pycnospores one-celled, hyaline (yellow-amber
in mass), thin-walled, generally ovoid but very irregular in shape and
size (5—) 9-13 (-18) x (3—) 3.5-5.5 (—7) yp, occasionally budding re-
peatedly at the same point and thus forming cruciform clusters of cells,
or later their contents “encysting” as thick-walled chlamydospores about
two-thirds the length and the same diameter as the parent spore. Pycno-
phores arising from the central loosely plectenchymic base of the pycnidial
cavity, dendroidally branched, producing spores acrogenously.
Substrata: Russula adustus, Lactarius pargamenus, Limacium penarium,
Pleurotus sp., and indet. agarics.— Leotia lubrica— “nettle roots” .
(Urtica), carrot (Daucus); all of these somewhat decayed when found.
Whetzel (in litt.) says that it grew easily in culture on several media
including bean stems, various fleshy fungi, and agar media.
Type: on blackened Ag. (Russula) adustus, King’s Cliffe, England,
Sept. 1836, collected by M. A. Berkeley.
Lectotype: on decaying Russula or Lactarius, Aug. 1917, Chocorua,
New Hampshire, U. S. A., coll. & det. VW. G. Farlow. Parts of this speci-
men (FH) will be distributed to the herbaria of the following institu-
tions: Mycology and Disease Survey of the U. S. Dept. Agr., New York
Botanical Garden, University of Michigan, Kew Botanical Garden.
Specimens examined: NEw Hampsuire, Chocorua, on agaric, Sept. 14, 1907. coll.
& det. as Sphaeronema vitreum by Farlow. — Chocorua, on Russula or Lactarius, 1917,
Farlow; Massacuusetts, Cambridge, on carrot, Feb. 19, 1890, Farlow. (FH slide
1642); New York, Enfield Gorge near Ithaca, on Leotia lubrica, Aug. 14, 1942,
Whetzel, (CU 31538) — Richford, culture on sporophore of Favolus europaeus original
isolate from decaying Pleurotus sp. with Hypomyces sp., Aug. 14, 1904, Whetzel
(CU 2022); Encianp, King’s Cliffe, on blackened Ag. (Russula) adustus, Sept.,
Sphaeronema blepharistoma Berk., Berkeley’s British Fungi, Fasc. III, nr. 189
(cotype) — also a duplicate of the last, and a slide made from it by von Héhnel;
Austria, Wiener Wald, on decaying Limacium penarium, Sept. 16, 1903, von Héhnel,
(FH-H 10202) — Wiener Wald, on Lactarius pargamenus, Aug. 21, 1906, von Héhnel,
(FH-H 10200) ; Bonemta, near Flana, on agaric, Aug. 12, 1905, Bubak, (FH-H 10202).
SEELER: FUNGICOLOUS FUNGI 125
No one has recorded having seen Corda’s specimen of Sphn. vitrea.
His description has three differences from that of the present species.
First, he says that the “mature spore, seen from the side, is egg-shaped
and at the same time four-edged” the latter probably meaning in cross-
section; second, he pictures the spores as very uniform in size and
having thick walls; third, his enlarged view of the pycnidial mouth does
not show the wide-flaring pointed hyphae so characteristic for H. blephari-
stoma. All three differences can be explained by saying: one, Corda
observed plasmolized, shrunken pycnospores; @wo, he illustrated chlamyd-
ospores after the pycnospore wall had dissolved; three, his close-up of
the beak is of a very young pycnidium. All of these are begging the
point, however, and are speculative theories as Corda’s specimen may
well have been something else entirely, perhaps an Ascomycete.
To point out the fallacy in such speculation one needs only refer to
von Héhnel (1. c.) where he calls a fungus of his own collection Sphn.
vitreum Cda. without comparing Corda’s type and then establishes
Hyalopycnis vitrea on this (v. H.) specimen as genotype. He says it
differs from Sphn. blepharistoma Berk., though Berkeley (1841) became
equally convinced that Sphn. blepharistoma was synonymous with Corda’s
fungus. Unfortunately von Hohnel labelled the four packets in his
herbarium “Sph’nmella’ with the species name followed by “?” and
never indicated the genus “Hyalopycnis” on them or noted which he
considered as H. vitrea or H. hyalina. Slight variations in size of
pycnidia or spores seem to have been his species criteria, and these have
no significant value in this case. At any rate, all of von Hohnel’s speci-
mens represent the same species, so both his species names are here
subjected to Berkeley’s prior blepharistoma, specimens of which have
proved to be the same thing. The five American collections, which have
not been recorded heretofore, are indistinguishable from the European.
Until Corda’s type specimen is found and examined, for the present
it seems best to use Berkeley’s name which was published the same year
as Corda’s, 1837, and which has an exsiccati set to substantiate it. Since
von Héhnel’s specimens on which Hyalopycnis was based are really later
collections of Sphn. blepharistoma Berk. it is proposed that Berkeley’s
species be the type of the genus Hyalopycnis v. H.
Listed here as doubtful is Sphaeronema vitreum Corda, Icon. Fung.
1: 25, pl. 7, fig. 297. 1837, with the combinations Ceratostoma vitreum
(Cda.) Fries, Summa. Veg. Scand. p. 396. 1849 and Melanospora vitrea
(Cda.) Sacc., Syll. Fung. 2: 463. 1883.
MICROPYXIS Seeler, gen. nov.
(Discomycetes, Pezizales)
Ascomata sessilia in subiculo tenuo; parvissima, gelatinosa; excipulo hypharum
parallelarum; paraphysibus distinctis, infra ramosis non epithecio formantibus; ascis
breve-clavatis, sursum obtuse rotundatis, 8-sporis; sporis elongato-ellipsoideis vel basem
leniter attenuatis, hyalinis, 1- vel 2-cellulatis.
126 Fartowia, VoL. 1, 1943
Ascocarp sessile on a thin subiculum, very reduced and small, gela-
tinous; excipulum composed of parallel hyphae; hymenial elements
surrounded and covered by gelatinous material; paraphyses distinct,
branched below, not forming an epithecium; asci short-clavate, bluntly
rounded above, 8-spored; spores elongate-elliptical or tapered slightly
toward the base, hyaline, 1- or 2-celled.
Differs from other genera in the button-like hypothecium combined
with an entirely filamentous-gelatinous excipulum and the absence of
any true epithecium. It®probably belongs below the most reduced
members of the Helotiaceae sensu Keissler in Rabenhorst’s Krypt. Flora
8: 52. 1930. Beside the absence of any perithecial wall, the shape of
the asci and spores and the presence of true paraphyses exclude it from
the Hypocreales.
Type species: Micropyxis Geoglossi (Ell. & Ev.) Seeler.
Micropyxis Geoglossi (Ell. & Ev.) Seeler, comb. nov.
Hypomyces Geoglossi Ellis & Everhart, Journ. Myc. 2: 73. 1886.— Journ. Myc.
3: 114, 1887 [revised description] —N. Amer. Pyreno. p. 72. 1892.
Peckiella Geoglossi (Ell, & Ev.) Saccardo, Syll. Fung. 9: 944. 1891.
Eleutheromyces Geoglossi (Ell. & Ev.) Seaver, Mycologia 1: 48, pl. IV, fig. 10-11.
1909 —N. Amer. Flora 3: 5. 1910.
PLATE II, Fic. 1-7.
Ascocarps superficial, parasitic on hymenial surface of members of
the Geoglossaceae, growing close together, mostly distinct, rarely anasto-
mosing, pale yellow-brown, translucent and gelatinous when wet, shrunk-
en and brittle when dry, resembling microscopic droplets of dark resin,
at first spherical, finally up to three times as broad as high, 60-150 »
diam. x 60-70 » high, ascus layer 25-45 » deep; base (hypothecium)
short, of thick-walled gelatinous hyphae, pseudoparenchymatous, lower
part composed of hyphae which spread out, surrounding the tops of host
paraphyses and asci and wedging between them and connecting to the
bases of neighboring apothecia; excipulum formed of a single layer of
branching filaments in an amber-colored gelatinous substance; these
arch over the tops of the asci in young apothecia almost completely cov-
ering them; no true epithecium of fused tips of paraphyses; pale amber
gelatinous material covers the tops of the asci. Asci eight-spored, clavate,
short-stiped, stipes mostly broad, thickened, obtuse apex, 28-45 x 6.4-8.3
rarely a little longer. Paraphyses hyaline, filamentous, simple or once-
branched near base, 1.2-1.6 » diam., tips never branched, swollen,
1.9-3.2 » diam., slightly overtopping the asci and about as numerous as
they. Spores one-celled, hyaline, long elliptical or tapering toward base,
ends rounded, irregularly distichous, 7.7—10.5 x 2.5-3.2 p. No parts
blue with iodine.
Type: on Geoglossum (glabrum?), [= Trichoglossum Farlowi (Cke.)
Dur.], Newfield, New Jersey, collected by J. B. Ellis, Sept. 1876.
SEELER: FUNGICOLOUS FUNGI 127
Specimens examined: (all on Trichoglossum Farlowi except the first, all FH.)
Massacuusetts, Nantucket, on Trichoglossum Walteri (Berk) Dur., Oct. 10, 1941,
and on 7. Farlowi, Aug. 1938 and Sept. 1941, Seeler; New Jersey, Newfield, “on
Geoglossum (glabrum)?”. Sept. 1879, ex herb. Ellis, authentic (FH), and Aug.
7, 1887, Ellis, authentic (FH-E); Tennessee, Burbank, “on Geoglossum,” Aug.
1896, coll. & det. Thaxter (FH 677); NortH Carouina, Cranberry, “on Geoglossum,”
Aug. 1896, coll. & det. Thaxter (FH 7931).
Since this interesting little fungus grows and matures on living sporo-
phores of Trichoglossum it can without doubt be called parasitic. In
the fresh and mature condition it can be detected by the smooth brownish
tinge the many apothecia give to the usually velvet-black hosts on which
it has been found. But after the specimens have been dried, it is im-
possible to be certain of its presence or absence even with a twenty-
power hand-lens. It does not seem to affect its host beyond causing a
shrinkage in the normally inflated, curved tips of paraphyses. Several
attempts to grow this fungus in culture on potato-dextrose agar and on
Trichoglossum-decoction agar have been unsuccessful; ascospores dis-
charged on the agar from fruit bodies fastened above, swelled a little
and produced short buds but grew no farther. No evidence has been
found for relating this species with any imperfect form; see Howell,
Mycologia 31: 196, 207, 209, Fig. C.8, D.4. 1939 (re Stephanoma tetra-
coccum).
SPHAERONAEMELLA Karsten, emend. Seeler.
(Hypocreales, Nectriaceae)
Perithecia light colored, soft, with neck composed of parallel filaments,
mouth fimbriate. Asci eight-spored, subspherical, evanescent; para-
physes none; ascospores one-celled, hyaline to pale yellow; differing
from Melanospora in the hyaline spores. Originally described as belong-
ing in the Sphaeropsidales by Karsten in Hedwigia 23: 17. 1884, and
emended by Grove in Brit. Stem and Leaf Fungi 2: 115. 1937 to include
pycnidial forms with appendaged spores.
Type species: S. Helvellae Karsten, emend. Seeler.
Sphaeronaemella Helvellae Karsten, emend. Seeler.
Sphaeria Helvellae Karsten, Fungi Fenniae exs. 674. 1867.
Sphaeronaemella Helvellae Karsten, Hedwigia 23: 17. 1884.— Diedicke, Krypt.
Fl. Brand. 9: 694, 1914.
Sphaeronema Helvellae (Karst.) Jaczewski, “Monographie du genre Sphaero-
nema Fries,” in Nouv. Mem. Soc. Imp. Nat. Moscow 15: 302, fig. 1. 1898.
PLATE II, Fic. 8-12.
Perithecia superficial or with lower half sunk in host tissue, mostly
growing close together but discrete, sometimes with a very thin whitish
hyphal subiculum from which grow branched conidiophores bearing
apically, hyaline, one-celled, cylindrical conidia, 14-18 (—23) x 3.5-5 yp;
when dry yellowish-amber, collapsed with necks erect like fine white
bristles topped by small amber spore-balls; when wet 500-850 » over-all
128 Fartowia, Vou. 1, 1943
height, body nearly spherical 150-200 p» diam., quite hyaline except for
the pale yellowish spores inside. Body wall of tile-like angular cells
grading in a short collar into the purely parallel septate-hyphal tissue
of the long slender neck which terminates in a mouth fimbriate with
obtusely pointed hyphae. Asci sub-cylindrical, eight-spored, soon evanes-
cent; paraphyses none; ascospores, hyaline-yellow, one-celled, walls clear-
ly visible, reniform to oblong, 8.5-10.5 x4-4.5 (-5) pe
Type: on “half-dead” Helvella infula, Sept. 1867, Mustiala, Finland,
collected by P. A. Karsten.
Specimens examined: (all on Helvella infula) FINLAND, 1867 et 1883, Karsten,
Fungi Fenn. 674, cotype, (FH) ; New Hampsuire, Shelburne, ¥. G. Farlow (FH);
New York, Adirondack Mts., Sept. 12, 1914, C. H. Kauffman & E. B. Mains, det.
C. H. K. (MICH); Micuican, Wallace Lake, Aug. 19. 1930, coll. & det. 4. H. Povah,
FP 533 (MICH).
The ephemeral nature of the asci misled Karsten and others into think-
ing that this, the type of the genus, belonged to the Sphaeropsidales,
and several light-colored perithecial species were transferred into
Sphaeronaemella after the limitation of Sphaeronaema to black forms.
According to the system of Saccardo there is a place for this genus among
the Ascomycetes in the Nectriaceae, differing from Melanospora in having
“hyaline” spores. The spores in mass are clearly yellow, so that this
species could be considered merely as a light spored Melanspora, but
the author prefers to retain Sphaeronaemella, emended, at least until
obvious relationships can be recognized with species in the black spored
genus.
Recently Professor H. H. Whetzel has been kind enough to send me
notes and drawings which he made in 1904 of apparently this species,
as details agree in every respect except for slightly smaller spores, “about
7x3% yw.” At that time he noted sub-globose, evanescent asci and ex-
ternal conidia. He found his fungus “parasitic” on Hypomyces sp.
which grew on a decaying Russula ?, Enfield Center Glen, Aug. 27, 1904.
Unfortunately the specimen has not been found for comparison.
It should be noted that von Hohnel in his Frag. Myk. 58 (Hedwigia
60: 151. 1919), though unfamiliar with any specimen of it, surmised
from the descriptions that S$. Helvellae was an Ascomycete when he
founded the genus Hyalopycnis for nectrioidaceous forms.
130 FarLowlA, Vou. 1, 1943
EXPLANATION OF PLATE I
All drawings made with the aid of camera lucida.
Fig. 1-6. Eleutheromyces subulatus (Fr.) Fuckel, emend. Seeler.
1. Hyphae from margin of ostiole. > 1175.
2. Pycnophores and ciliate pycnospores. 1175.
3. Surface pattern of pycnidial wall cells. > 550.
4-5. Mature pycnidia drawn in moist condition. Broken line indicates size
of spore-ball at apex. >< 65.
6. Vertical section through pycnidium. Pycnophores line the entire inside of
wall > 65,
Fig. 7-10. Hyalopycnis blepharistoma (Berk.) Seeler.
7. Hyphae from margin of ostiole. > 1175.
8. Pycnophores and pycnospores. 1175.
9, a. Chlamydospores which form inside of pycnospores whose wall remnants.
adhere. x 1175.
b. Cruciform bud-spores from neck canal. > 1175,
a. Young pycnidium, ostiole still closed. > 65.
b. Vertical section of mature pycnidium. Pycnophores grow at base only of
cavity. Wall and neck are composed entirely of parallel scarcely septate
filaments whose tips are shown in fig. 7. 65.
c. Mature pycnidium, drawn in moist condition. 65.
10.
Fig. 11-15. Eleutheromycella mycophila v. Hohnel.
11. Pycnophores and ciliate pycnospores. > 1175.
12. Surface pattern of pycnidial wall cells. > 550.
13. Diagram to show position of pycnidia in sporophore of Polystictus versicolor.
~ IE.
14. Vertical section through pycnidium near upper surface of host. 65.
15. Vertical section through pycnidium in lower (pore) surface of host. Pycno-
phores line the entire inside of wall. x 65.
1, Drawn from TRT 4859 (FH). 2a. Sphaeronema oxyspora Berk. cotype in
Herb. Curtis. 2b. Klotsch, Herb. Viv. Myc. 1463 (FH). 2c. Soc. Hill, S. C. in Herb.
Curtis. 3. TRT 4859. 4. Shelburne, N. H., Farlow. 5. Fungi Rhenani 773 (FH).
6. TRT 4859. 7-10. Chocorua, N. H., 1917, Farlow; except pycnidia which are from
cotype of Sphaeronema blepharistoma Berk., Brit. Fungi 189 (FH). 11-15. Type
in Herb v. Héhnel.
SEELER: FUNGICOLOUS FUNGI Pow
PLATE 1
132 FarLowia, VoL. 1, 1943
EXPLANATION OF PLATE II
All drawings made with the aid of camera lucida.
Fig. 1-7. Micropyxis Geoglossi (Ell. & Ev.) Seeler.
1. Stages in the development of asci. > 1175.
2. Two mature asci. 1175.
3. Eight mature ascospores. X 1175.
4, Paraphyses. > 1175.
5. Habit diagram to show apothecia growing on surface of Trichoglossum hy-
menium. X 85.
6. Vertical section through a very young apothecium showing ascogonial coil
and beginnings of ascogenous hyphae among marginal paraphyses. Dotted
line to left indicates extent of gelatinous material which covers the apothe-
cium. 1175.
7. Vertical section through a nearly mature apothecium to show arrangement of
asci, paraphyses, and marginal filaments. Darker cells at base envelop
the curved Trichoglossum paraphyses. x 800.
Fig. 8-12. Sphaeronaemella Helvellae Karsten, emend. Seeler.
8. a. Diagrammatic sketch of exterior of mature perithecium to show tile-like
cells forming body, filaments forming neck. Dotted line at apex indicates
ball of spores. X 65.
b-c. Outlines of young and of mature perithecia. x 65.
9-10. Asci (spores immature) and mature ascospores. 1175.
1]. Apex of neck of perithecium showing ascospores in single row. 1175.
12. Conidiophore and conidia which grew on the outside of the perithecium.
x 1175.
1-7. Drawn from Nantucket, Mass. material. Seeler. 8, and 10-12. from Shelburne,
N. H., Farlow. 9. from cotype, Fungi Fenn. 674.
SEELER: FUNGICOLOUS FUNGI 138
PLATE II
1(1): 135-170 FARLOWIA January, 1943
STUDIES IN THE GENUS HELOTIUM, Il
HISTORY AND DIAGNOSIS OF CERTAIN EUROPEAN AND
NORTH AMERICAN FOLIICOLOUS SPECIES '
W. Lawrence WHITE
The object of the present paper is to present as thorough a taxonomic
study as available material will permit of the species of Helotium (sensu
latu)? occurring on the blades of fallen, decaying leaves of woody
dicotyledonous plants in Europe and North America. If such a paper
would be more than another incident in an historical background of in-
accuracy and confusion, and in itself a perpetuation of and an addition
to such a preéxisting condition, it must present more than the usual
series of species descriptions: it must undertake rather completely to re-
view and disentangle the whole literature record of the several species
which fall within the scope of its restricted subject.
To this end the study is first of all based on an examination of types,
for in a genus where taxonomic characters are as wanting as they are
in this one® many if not the great majority of the species simply cannot
be interpreted from the brief original diagnoses that have been presented
in the past; neither can their identity be established by later diagnoses,
which are often based on incorrect determinations, nor, except very rarely
[e. g., H. epiphyllum (Pers. ex Fr.) Karst], by general agreement or
usage. To illustrate by a single example: H. phyllophilum (Desm.)
Fr. was described by Desmaziéres from France in 1842, and the original
material, said to be on Acer and Fagus, was widely distributed to herbaria
as an exsiccati number. Accompanying it was a diagnosis sufliciently
vague (as is to be expected of that period) to apply almost equally well
to any of the species from numbers 7 to 13 inclusive in the present paper.
Having been listed on two substrate species it is possible that a mixed
fungus species is involved. The packets which I have examined from
several herbaria, however, all contain leaves of Fagus bearing good and
abundant apothecia of a single well-defined species. The point to be
made is that of eighteen later collections under the specific epithet of
Desmaziéres which have been available for examination— and despite
the general availability of the original material — five distinct species
are represented, none of which agrees with the original material on
Fagus leaves.
1Contribution from the Laboratories of Cryptogamic Botany and ‘the Farlow
Herbarium, Harvard University, no. 216.
2But not in the all-inclusive sense of early mycologists who placed here species
now correctly referred to Ciboria, Chlorociboria, Chlorosplenium, Rutstroemia, and
several other genera. In the first paper of this series (Mycologia 34: 154-179,
fig. 1-16. 1942). ten of the eighteen species described in Helotium by Peck were
excluded and placed elsewhere. Despite this gradual narrowing, however, the so-
called genus as it still is recognized (White, Amer. Midl. Nat. 28: 512-523. 1942) is
large and in the light of modern generic concepts probably diverse.
* Cf, Amer. Midl. Nat., 1. c.
135
136 FarLow1a, Vo. 1, 1943
Secondly, an attempt is made either to confirm or reidentify all of
the European (omitting H. epiphyllum) and American exsiccati speci-
mens, as well as others taken within this range, because, among several
reasons, these must be used as a basis for comparison for future deter-
minations by those who do not have access to the types, and further be-
cause they are the basis of our knowledge of the geographic and sub-
strate ranges for the species. The fallacy of basing any mycogeographical
studies on the now-existing records of the aforementioned H. phyllo-
philum is at once apparent. The effect of sorting out and redetermina-
tion of historical material on known geographical distribution may again
be illustrated by H. immutabile, which was described by Fuckel from
Germany in 1871. Removing H. immutabile from synonymy with H.
conformatum (Karst.) Karst., to which it is scarcely remotely related,
takes it out of Scandinavia, and a correction of the only previous report
of the species from North America takes it out of Canada; finding it in
herbaria under incorrect names takes it to Italy, Massachusetts, New
Jersey, and Minnesota.
Thirdly, the paper aims to provide a complete bibliography of each
species, except for the first, H. epiphyllum, for which European references
have been omitted for want of space, and to indicate whether or not they
are correct. This goes hand in hand with the examination of specimens
as indicated in the preceding paragraph and is necessary for the same
reasons. And there is still another factor involved: the amount of ex-
ploring for these forms has been so scanty that it becomes necessary to
search out the basis of each local report in order to get material for study.
This, of course, has been supplemented as much as possible by a study
of fresh material. At first thought it appears impossible to check any
appreciable number of literature references— but the literature is not
very extensive, and fortunately, type and other historic material of the
helotiaceous fungi has been well preserved and a fair concentration of it
is to be found in the various institutions of northeastern United States.
Having been privileged over a considerable period to work in the larger
herbaria of this region, especially those of Harvard, Cornell, and the
New York Botanical Garden, each generally rich in exsiccati material
and library facilities, and each excelling in its own particular private
collections — i. e., Harvard with its von Hodhnel and Patoulliard collec-
tions and others, Cornell with its Durand collection of 12,000 discomy-
cetous specimens including a fragment from nearly every specimen that
was the basis of a North American literature record up to about 1905,
and the New York Botanical Garden with its Massee herbarium of British
material —I feel that the opportunity for such a study has been ample.
To be sure, some type and otherwise important material has been un-
available, and a number of species therefore must be left for future
reference.
It is hoped that this treatment, together with those references indicated
Wuite: Stupies In Genus Hetotium. IIL 137
in the following key, may enable the investigator to determine most of
the foliicolous species of Helotium and Helotium-like fungi which occur
in central and western Europe and northeastern United States. . The re-
maining vast areas of both continents remain “untouched” as far as such
fungi as these are concerned.
A certain degree of substrate specificity permits the grouping of the
helotiaceous fungi for purposes of convenience as occurring on leaves,
petioles, wood, fruits, grasses, herbaceous stems, etc. Strict attention to
phylogeny might well be left until a complete monograph of a genus or
family can be prepared; to attempt it now, especially in view of the
relative lack of outstanding characters in these forms, either specific or
of higher category, could result only in a rapid series of name changes,
or at best a series of papers which could bear little or no meaning to
the non-specialist. Of primary concern is the placing of the individual
species on a sound taxonomic basis. With phylogeny ultimate, however,
the following remarks on the species of this paper may be pertinent.
The first three species, H. epiphyllum (Pers. ex Fr.) Karst. (plus
Ombrophila subaurea Cooke, which is appended as doubtfully distinct) ,
H. carpinicolum Rehm, and H. immutabile Fuckel, are surely very closely
allied. They undoubtedly form a natural taxonomic unit with certain
similar forms such as are usually referred to as H. lutescens (Hedw.)
Fr. or H. pallescens (Pers.) Fr., which occur on wood. It is certain
that H. epiphyllum is not confined to leaves, and some specimens now
in herbaria under the two names just mentioned may be referable to this
species. Number four, described as new, shows affinities with those pre-
ceding it but is somewhat uncertain. Number five, H. Conocarpi Seaver,
is included because it might be looked for here; as indicated later in
the discussion of that species it probably belongs in the Ciborioideae.
The sixth, presented as new, appears to be intermediate between the
H. epiphyllum group and the slender, stipitate, and more or less delicate
forms represented by numbers 7 to 13, inclusive. The latter group
most certainly forms a highly natural series and is allied with a group
consisting of H. scutula (Pers. ex Fr.) Karst., H. fraternum Peck, and
H. Dearnessii (Ellis) White, which occur on substrata other than leaf
blades. The punctiform species (Pezizella sensu Rehm) represented by
numbers 14 to 17 are of uncertain affinities — perhaps miscellaneous —
yet scarcely set off from the H. epiphyllum group by any obvious char-
acter except by the smaller dimensions, and from the H. caudatum group
(no. 7-13) by anything other than the lack of.a long, slender stipe.
As to materials and methods, all observations and drawings of micro-
scopic structures are based on material mounted in KOH-phloxine (Martin,
Mycologia 26: 261-265. 1934) and studied under the oil immersion
objective. The figures are drawn with the aid of a camera lucida to a
magnification of 1540 (with the exception of no. 4, which is x 1835) and
reduced slightly on reproduction. Descriptions of gross morphology and
4
138 Fartowia, Vou. 1, 1943
surface markings are based on material observed under a good binocular.
Technical color terms are those of Ridgway.
In the citation of literature, references known to be based on incor-
rectly determined material are enclosed in brackets. The basis for in-
dicating a record as incorrect or for allowing it to stand may be deter-
mined by a perusal of the material examined or of the discussion in the
treatment of the particular species.
The specimens examined are listed geographically and chronologically.
Whenever a specimen cited under a particular species has been filed or
distributed under an incorrect name the latter is given and is enclosed in
brackets. The herbarium in which each specimen examined is located
is indicated by an abbreviation in parentheses. The abbreviations are
those of Lanjou (Chronica Botanica 5: 142-150. 1939) with an addi-
tional letter added in many cases to designate the private collections of
the various institutions. The meanings of the latter will be apparent to
most workers who have reason to pay any attention to them; most of
them are listed in the first paper of this series (Mycologia 34: 154-179.
1942).
KEY TO THE SPECIES
1. Apothecia sessile, subsessile, or with stipe present but obscure or at least much
shorter than the diameter of the disc.
2. Apothecia large, fleshy, 1-4 mm. diameter.‘
3. Apothecia broadly sessile, watery and sordid when fresh, drying soiled-
ochraceous; paraphyses branched at the apex; North American.
4. H. contortum
3. Apothecia attached by a narrow central papilla or having a short stipe ;
paraphyses not branched at the apex.
4. Spores obovoid, regular and equilateral in outline, not larger than 10-
13 x 4-5 py.
5. Spores of the dimensions indicated above, slightly trapezoid-shaped ;
apothecia white when fresh, drying yellow or orange; on various
species in Europe and N. Am. ............ 3. H. immutabile
5. Spores slightly smaller, not at all trapezoid in outline; apothecia dark
brown when dried; on Conocarpus in Bermuda 5. H. Conocarpi
4. Spores more elongate, more or less fusoid. inequilateral or flattened on
one side, more than 13 y, long; apothecia drying pale to deep orange.
6. Spores narrow oblong-fusoid asymmetrical or slightly curved, 16-24 x
OO Mh dots tesa tech edaw ah en deci, 1. H. epiphyllum
6. Spores elongate-ellipsoid, scarcely fusoid, flattened on one side or
slightly curved, 13-16x44.5 4 ............ 2. H. carpinicolum
2. Apothecia minute, 0.1-0.8 mm. in diameter,
7. Apothecia lemon-yellow, thin, plane, puberulent; spores distinctly long and
narrow, 12-18x 18-26 » ....0.0.........0.0.. 17. H. puberulum
7. Apothecia and spores not entirely as above.
8. Apothecia up to 0.8 mm. in diameter, subfleshy, deeply concave, then
saucer-shaped, pale to ochraceous yellow, easily visible to the unaided
DVS hos epg hak beds Whe add Geo ae's 14. H. populinum
*Cf. also Pycnopeziza (White & Whetzel, Mycologia 30: 187-203. 1938; Whetzel
& White, Mycologia 32: 609-620. 1940), or if brown-spored, Phaeociboria: one species
occurring on Hedera Helix L. in Europe (1. c., p. 614).
WHITE: STUDIES IN Genus HELoTium. III 139
4
8. Apothecia not over 0.3 mm. diameter when fresh, translucent whitish or
brownish, drying dirty ochraceous or brownish and scarcely or not at
all visible to the unaided eye.
9. Paraphyses forked at the apex; spores 2-celled; North American.
16. Pezizella glagosa
9. Paraphyses not branched at the apex; spores always 1-celled.
10. Apothecia with a short but distinct stipe; asci 75-95 x 10-11.5 p;
spores obovoid, 12-15 x 44.5 u; North American.
6. H. translucens
10. Apothecia erumpent, then broadly sessile; asci 40-46 x 8.5-9 y;
spores oblong, 12-13 x 2.8-3.3 u.; on leaves of Betula in Europe.
15. H. tumidulum
1. Apothecia distinctly stipitate, the stipe slender, as long as the diameter of the
disc or longer. (Slender delicate species, white to yellow or pale ochraceous,
not over 2.5 mm. high).°
11. Apothecia with lower half of stipe black; spores small, narrow-oblong,
6-8 x 1.4-1.8 u; on leaves of Chamaedaphne in North America.
13. Phialea Cassandrae
11. Apothecia not with stipe as above; spores conspicuously larger.
12. Spores distinctly long and narrow, 28-36 x 3.5-4.5 %; North American.
7. H. fastidiosum
12. Spores not more than 25 y, long.
13. Hymenium drying cinereous; spores 16—18 x 3.5—4 4; North American.
9. H. Linderi
13. Hymenium drying pale ochraceous.
14. Spores 16-24 x 4.5-5.5 4; Europe and North America.
8. H. caudatum
14, Spores smaller, not more than 16 y, long.
15. Asci originating from croziers; spores narrow oblong-fusoid, dis-
tinctly 2-celled, 11.5-15 x 3.2-4 y.; European.
12. H. phyllophilum
15. Asci not originating from croziers; spores 1-celled.
16. Spores narrow, 14-16 x 3.5-4 yw; on leaves of Fagus in North
ATNOTION Rog. pitt Sak Pee ee eet ees ae 10. H. albopunctum
16. Spores broader, 11-14x3.8-5 wu; on leaves of Populus in
HrOpes ey. Ae tes a eed ee 11. H. phyllogenon
THE SPECIES
1. Helotium epiphyllum (Pers. ex Fr.) Fr., Summa Veg. Scand., p. 356. 1849, —
Cooke, Handb., p. 715. 1871.— Peck, New York St. Mus. Rep. 25 (1871):
98. 1873. — Cooke, Bull. Buffalo Soc. 2: 299. 1875. — Frost, Cat. Pl. Amherst,
p. 91. 1875. — Harkn. & Moore, Cat. Pacific Fungi, p. 37. 1880. — Bundy, Geol.
Surv. Wisconsin 1873-79 1: 401. 1883. — Peck in Day’s Cat., Bull. Buffalo Soc.
4: 220 (156). 1883. — Cobb, List Pl. Amherst, p. 47. 1887. — Sacc., Syll. Fung.
8: 227. 1889. — Ellis in Britt., Cat. Pl. New Jersey, Rep. St. Geol. 2: 550. 1890.
— Atk. & Schrenk, Journ. El. Mitch. Soc. 9: 106 (12). 1893.— Lindau in
Engl. & Pr., Nat. Pfl. 1*: 207. 1894.— Durand, Bull. Torrey Club 27: 483.
pl. 31, fig. 13. 1900. — Kauffm., Rep. Michigan Acad. 8: 28. 1906.— Millsp.,
West Virginia Geol. Surv. 5(A): 111. 1913. — Dodge, Journ. Wisconsin Acad.
17: 1034. 1914. — Kauffm., Rep. New York St. Mus, 68: 83. 1915. — Burnh.
°Tf brown, or if they become brown on drying, compare in Rutstroemia (White,
A Monograph of the Genus Rutstroemia (Discomycetes). Lloydia 4: 153-240, fig.
1-75. 1941); if white or bright colored, but larger and more robust, compare in
Ombrophila, especially O. clavus (Alb. & Schw.) Fr.
140 FARLOWIA, VoL. 1, 1943
& Lath., Torreya 23: 5, 1923.— Bisby et al, Fungi Manitoba, p. 62. 1929. —
Cummins, Papers Michigan Acad. 11: 111. 1930.— Stev.. Claytonia 3: 25.
1936. — Linder, Mycologia 29: 372. 1937.— Bisby et al, Fungi Manitoba &
Sask., p. 40. 1938.— Mains et al, Mycologia 31: 730. 1939.— Wehm.,
Canadian Journ. Res. 18(C): 547. 1940.
Peziza epiphylla Pers., Tent. Disp. Fung., p. 72. 1797. — Fr., Syst. Myc. 2: 137.
1822. — Schw., Naturforsch. Gesell. Leipzig Schr. 1: 122, 1822. — Schw., Trans.
Amer. Phil. Soc. 4: 175. 1834. — Curtis, Bot. North Carolina, p. 134. 1867. —
Cooke, Bull. Buffalo Soc. 2: 299. 1875.
Helotium aurantiacum Cooke ex Phill., Grevillea 92: 106. 1891.— Nom. nud.
Cooke in Britt., Cat. New Jersey Pl., Rep. St. Geol. 2: 550. 1889. — Sace.,
Syll. Fung. 10: 8. 1892.
Calycina aurantiaca (Cooke ex Phill.) Kuntze, Rev. Gen. Pl. 37: 448. 1898.
Calycina epiphylla (Pers. ex Fr.) Kuntze, Rev. Gen. Pl. 3?: 448. 1898.
Hymenoscypha epiphylla (Pers. ex Fr.) Rehm ex Kauffm., Papers Michigan
Acad. 9 (1928): 177. 1929. :
FIGURE Il.
Apothecia scattered, sparse to rather numerous, solitary or more
rarely in groups of 2-4, bright yellow to bright or dull orange when
fresh, all parts more or less concolorous, the color not changing markedly
on drying, fleshy, sessile to short-stipitate, reaching a diameter of 4 mm.
and a height of 1.8 mm., drying waxy to waxy-cartilaginous; stipe absent
or short, commonly merely a papilla or a narrowed basal portion of the
disc, more rarely cylindric and distinct, entirely smooth, pale yellow or
yellow when fresh, yellow to pale orange on drying; receptacle smooth,
concolorous with stipe, retaining its fullness on drying; dise opening
by a pore very early, fleshy, saucer-shaped, at maturity nearly or quite
plane, sometimes repand with maximum moisture; hymenium yellow
when fresh, on drying becoming deep yellow to orange or even a deep dull
red; margin thick, even; in section composed largely of a thick, compact,
homogenous tissue of thin-walled, short-celled hyphae 3-5 » diam. and
a more or less distinct ectal layer; ectal layer at least in the region
of the stipe consisting of thin-walled isodiametric cells 8-14 ,» diam.,
this structure continued to or nearly to the margin or gradually giving
way to a less well differentiated tissue made up of the slightly enlarged
and compacted and upwardly directed hyphal tips from the medullary
region; paraphyses long, straight, scarcely or not at all thickened at
the apex, 2-3 » diam.; asei originating from croziers, clavate, 90-130 x
9-12 yw, the pore faintly colored with iodine; ascospores fusoid or
oblong-fusoid, more or less asymmetrical or slightly curved, 1-celled,
16-23 x 4-5 yp, uniseriate, sometimes biseriate above, the content granular,
sometimes with two or three definite oil globules.
HasitaT: North American material on leaves of Abies (Cummins),
Acer, Alnus (Kaufim.), Amelanchier, Betula, Castanea (also on peri-
carps), Fagus, Populus, Quercus, and species indet.; also on forest debris
of various kinds: twigs of oak and other species, fern fronds, herbaceous
stems, and rabbit dung.
DISTRIBUTION: Quebec, Nova Scotia (Wehm.), New Hampshire, Massa-
Wuite: Stupies In Genus Heotium., III 141
chusetts, New York, New Jersey, Pennsylvania,* Maryland,* District of
Columbia,* West Virginia, Virginia* (Stev.), North Carolina, South
Carolina, Tennessee, Michigan, Wisconsin (Bundy), Manitoba (Bisby),
Montana, California (Harkn.). The best known and perhaps the most
widely distributed of the leaf-inhabiting species of Helotium. Common
both in North America and in Europe.
MATERIAL EXAMINED: Quebee: Duchesnay, County Portneuf, Aug. 25, 1938, H. H.
Whetzel & T. Sproston [as H. phyllophilum] (CU-P 27850).— Duchesnay. Aug. 26,
1938, R. F. Cain 11154 [as H. immutabile] (TRT).—N. H.: White Mts., Sept. 1884,
Miss Minns (CUP-D 8443). — Shelburne, Sept., 91 (FH).— Gilmanton, Oct. 12, 1942,
D. H. Linder (FH).— Mass.: Canton, 1925, D. H. Linder (FH).— Canton, July
1933, D. H. Linder (FH).—Swamp near Squam, Nantucket, Sept. 8, 1942, E. V.
Seeler 730a (FH).— Drake’s Marsh, Maddeket, Nantucket, Sept. 11, 1942, E. V.
Seeler 730b (FH).— Cushman, Sept. 12, 1942, T. Sproston (FH).—N. Y.: Enfield,
Aug. 20, 1935, W. L. White 2022 & H. J. Miller (FH).— White Lake, Adirondack
Mts., Sept. 2, 1936, Whetzel & White (CU-P 25519, FH).— Newcomb, Huntington
Wildlife Forest, Adirondack Mts., Sept. 6-17, 1941, Rolf Singer (FH).—N. J.:
Newfield, Sept. 18, 1875, as H. aurantiacum (CUP-D 8399. 8400, NY-E 940).— New-
field, Oct. 1875, J. B. Ellis, Type? of H. aurantiacum (CUP-D 3748).— Newfield,
1874, Ellis, Thiimen, Myc. Univ. 315, as H. epiphyllum (CUP-D 10376) .— Newfield,
Oct. 1, 1879, as H. aurantiacum (CUP-D 8401, NY-E 3332).— Newfield, July 28,
1883, Ellis, as H. epiphyllum (CUP-D 8442).—Newfield, July 1889, Ell. & Ev.,
N. Am. Famed 2328 (FH).—W. Va.: Nuttallburg, Aug. 1896, L. W. Nuttall, Ell. &
Ev., Fungi Columb. 1023 (CUP-D 10304, FH).—N. Car.: Cranberry, alt. 3250 ft.,
July—Aug. ’87, R. Thaxter (FH).— Blowing Rock, Blue Ridge Mts.. alt. 3500-5000
ft., July 20-Sept. 14, 1901, FE. J. Durand (CUP-D 12398).— S. Car.: Aiken, on damp,
fallen leaves of Cerasus, Dec. 1880, Ravenel 3128, as H. epiphyllum var. Cerasi.
[Have seen only a fragment and it is not typical of the species.] (CUP-D 3742).—
Tenn.: Grassy Patch, Great Smoky Mts. Nat. Park, elev. 4200 ft., Aug. 18, 1939,
D. H. Linder (FH).—Indian Gap, Great Smoky Mts. Nat. Park, Aug. 20, 1939,
D. H. Linder (FH).—Mich.: Nubish, Sept. 1917, FE. T. & S. A. Harper S199
(CUP-D 10828).— Marquette, 9/11/34, E. B. Mains 34-149 (FH).— Mont.: Echo
Lake, Flathead Nat. Forest, July 20, 1928, G. B. Cummins; comm. Dr, Kanouse
(MICH).
The species appears to exhibit, within well defined limits, a certain
amount of variation in form and structure of the apothecium, and an
occasional specimen, such as the one cited above from South Carolina
and another (not cited) taken at Cranesville, W. Va., Aug. 6, 1908, by
J. L. Sheldon (CUP-D 6471) has spores shorter and more obtuse than
are normal for the species, in this latter respect resembling H. carpinicolum
(described on the following pages). As I have presented H. epiphyllum
here, it is characterized by its rather large and fleshy, substipitate orange-
colored apothecia, long asci originating from croziers (cf. Ombrophila
subaurea, mentioned below), and distinctly elongate and fusoid spores.
Durand (1. c.) describes and illustrates a very distinct pseudoparenchy-
matous ectal excipulum extending around the apothecium from the sub-
stratum nearly to the margin. However, Lagarde (Ann. Myc. 4: 234-235,
fig. 53. 1906), presenting a detailed anatomical and morphological study
* fide Miss Cash from material in Pathol. & Mycol. Collections, U. S. D. A.
142 FarLowia, Vou. 1, 1943
of the species, presumably based wholly or largely on European material,
indicates that he never encountered this layer in the material he ex-
amined, although he did find the hyphal tips compacted in such a way
as to form an outer zone rather distinctly different from the tissue of the
broad middle region. In my own experience I have found the outer
layer to be always pseudoparenchymatous in the basal region at or near
the stipe, and sometimes extending well up toward the margin, but
scarcely in any case as distinctly defined as illustrated by Durand. The
latter is more typical of the two species following, H. carpinicolum and
H. immutabile, than it is of H. epiphyllum. Note the variation in form
shown by Nannfeldt (Nova Acta Reg. Soc. Sci. Upsala 4: 8?: pl. 14,
fig. 2-4. 1932) in photomicrographs of three different European collec-
tions. I have examined numerous collections from Europe and fail to
find any differences in the species as it occurs typically in Europe and
as it occurs in North America.
The species which I have here referred to synonymy with H. epiphyllum,
that is, H. aurantiacum, does not appear to have received any attention
since the original description by Phillips. Phillips lists the material
on which his description is based as “J. B. Ellis, No. 75” without date
or locality, and it is impossible at present to be sure which of the three
specimens cited above, if any, actually represents the type. That of Oct.,
1875, is marked “Type” on the portion which is in the Durand Herbarium.
It contains a single apothecium which is sessile and 2 mm. in diameter.
The Durand portion of the 1879 material has apothecia up to 4.5 mm.
in diameter, and of the Sept. 18, 1875, collection up to 1.3 mm. diam.;
in both they are sessile to substipitate. They all obviously represent
the same species and are typical H. epiphyllum.
Several varieties of H. epiphyllum remain to be investigated: var.
acarium Karst., var. candidum Karst., var. Ellisii Rehm, var. praeponens
Karst., and perhaps others.
Ombrophila subaurea Cooke, Bull. Buffalo Soc. Nat. Sci. 2: 300. 1875.
The material of Ellis, N. Am. Fungi 394, which presumably is type,
differs from typical Helotium epiphyllum only in that croziers are en-
tirely lacking. This is the only material I have seen with the spores
and other characters of H. epiphyllum, yet lacking croziers. Cooke’s name
may remain unaltered pending future investigation.
2. Helotium carpinicolum Rehm, Hedwigia 35: 146. 1896.— Sacc., Syll. Fung.
14: 764. 1899. — Boud., Hist. Classif. Discom. Eur., p. 113. 1907. — Gola, Erb.
Mic. Sace. Cat., p. 119. 1930.— Nannf., Nova Acta Reg. Soc, Sci. Upsal. 4:
8’: 298, pl. 14, fig. 1. 1932.—[Vel., Monogr. Discom. Bohem, 1: 204. 1934.]
FIGURE 2,
Fig. 1-3: 1, Helotium epiphyllum, from the type ? of H. aurantiacum (CUP-D
3748) ; 2, H. carpinicolum, from Rehm’s Ascom. 1157, type (FH); 3, H. immutabile,
from Fuckel’s Fungi Rhenani 2388 (FH). All drawings 1290.
143
StupIEs IN GENUS HEtotTium. III
WHITE
oe APT OLIE
A
Ficures 1-3.
144 FarLowl!A, VoL. 1, 1943
Apothecia sparse, solitary or more rarely in groups of two or three,
arising from the mid, lateral, and net veins of the leaf, sessile, reaching
a diameter of slightly more than 1 mm. in the dried condition and ex-
hibiting a bright orange color in all stages of development; at first
papillate, erumpent, then more or less erect, columnar, sometimes distort-
ed, enlarged at the apex, opening by a pore, very early becoming saucer-
shaped to plane, thick, fleshy, finally reaching a diameter of 0.7—1.2 mm.,
the largest up to 1.6 mm. on being moistened; at maturity attached by
a central papilla; receptacle concolorous with or frequently slightly
more yellowish than the hymenium when dry and slightly raised above
the substratum, when moistened lying closely adjacent to it; hymenium
in the dried condition plane to slightly convex, orange colored: “Ochra-
ceous Orange,” “Zinc Orange,” or “Apricot Orange” (R), on moistening
becoming pale yellow and strongly convex; margin smooth, in dried
specimens slightly upturned in young material, scarcely so with age,
lying closely adjacent to the substratum when moist; in section composed
of a thick medullary excipulum of closely interwoven, narrow, thin-
walled hyphae, and a distinct ectal layer rather thin and made up of
closely arranged, globular, thin-walled cells 7-12 » diam., this layer
grading above into a marginal tissue of parallel, rectangular cells
7-12x 3.5-5 p directed outward and upward in filaments ending in
cells with obtuse rounded ends, of the same diameter, and closely ap-
pressed to form a smooth external surface; paraphyses simple, not
enlarged at the apex, 3-3.5 » diam.; asci 90-110 x 11-13 B, arising
from crosiers; ascospores biseriate or sub-biseriate, ellipsoid-cylindric,
flattened on one side, 13-16x4-4.5 p, always one-celled, those from
dried specimens containing a small amount of scattered refractive granu-
lar material in each end.
Hagitat: On leaves of Carpinus Betulus L.; also reported on Fagus
sylvatica L.
DIsTRIBUTION: Germany; also reported from Czechoslovakia. Sept.—Oct.
MATERIAL EXAMINED: Germany: “Jungfernhaide bei Berlin. An faulenden Blittern
von Carpinus Betulus.” 9/1894, Sydow, Rehm, Ascom. 1157, type (FH, FH-P, NY-E).
— “Ad folia putrida Carpini Betuli. Marchia: Bredower. Forst pr. Nauen.” Oct. 1896,
P. Sydow, Rabenh.-Pazsch., Fungi Eur. et Extraeur. 427] (FH).
This species is extremely close to Helotium epiphyllum, but I am con-
vinced that it is distinct — and possibly local in occurrence. The two
German specimens cited above are strictly uniform in their characters,
and in comparison with most of the material that has been referred to
H. epiphyllum they are smaller, tend to be more definitely sessile, in the
dried condition remain more fully expanded, in section exhibit a more
distinct ectal excipulum of small isodiametric cells, have shorter asci,
and have smaller spores with ends rounded rather than pointed. Nann-
feldt’s (1. c.) photomicrographs of the apothecia of the two species in
section scarcely show any differences.
WHITE: STuDIES IN Genus HELotTium. III 145
_ It appears highly probable that some of the central European specimens
now in herbaria as H. epiphyllum will prove on closer examination to
be H. carpinicolum, and there is no reason to suppose that all of them
will be on Carpinus.
3. Helotium immutabile Fuckel, Symb. Myc., Nachtr. 1, p. 50. 1871.— Sacc.,
Padova. Soc. Venet. Trent. Sci. Atti 4: 137. 1875.— Phill., Man. Brit.
Discom., p. 162. 1887.—Sacc., Syll. Fung. 8: 241. 1889.— Massee, Brit.
Fung.-Fl. 4: 259. 1895. — Gola, Cat. Erb. Mic. Sacc., p. 120. 1930. — [Mycologia
31: 730. 1939.]
Helotium conformatum auct. non Karst.— Rehm in Rabenh. Krypt.Fl. 1°:
780. 1893.— Jaap, Abh. Bot. Ver. Prov. Brandenburg 52: 5. 1910. — Migula
in Thomé’s Krypt.-Fl. 10°: 1197. 1913.— Nannf., Nova Acta Soc. Sci. Upsal.
4: 8: 298, pl. 15, fig. 1. 1932.
Pachydisca immutabilis (Fuckel) Boud., Hist. Classif. Discom. Eur., p. 94. 1907.
— Ramsb., Brit. Myc. Soc. Trans. 4: 371. 1913.
FIGURE 3.
Apothecia sparse, solitary, short-stipitate, white when fresh, drying
yellow to orange, 0.8-1.5 mm. diameter; stipe short, distinct, 0.2-0.5 mm.
long, firm, straight, cylindric below and slightly enlarged above, some-
times obconic, abruptly or gradually expanded into the disc, smooth,
always concolorous with receptacle; dise of medium thickness, spreading,
changing but little on drying, plano-convex or repand; receptacle
smooth and white when fresh, pale yellow to pale orange on drying, not
wrinkled, sometimes marked with very fine superficial striae radiating
from upper part of stipe to margin, these disappearing on moistening;
hymenium when fresh white, convex, drying firmly waxy, plane to
slightly convex, pale yellow or in old herbarium specimens becoming
pale orange-red or deep reddish brown; margin even, subacute, at
maturity little or not at all elevated above the hymenium either when
fresh or dried; in section composed of a broad medullary region of
rather loosely interwoven, thin-walled, irregular, branched, septate hyphae
2.56 » diameter, and a thin well defined ectal layer composed of thin
walled compactly arranged globular cells 9-16 » diameter; paraphyses
once forked below the middle, scarcely enlarged at the apices, 3-3.5 yp
diameter; asei clavate, arising from croziers, 80-105 x 8-10 yp, the pore
faintly blue with iodine; ascospores uniseriate to sub-biseriate, 1-celled,
angular-obovoid, 10-13 x 3.8-4.6 y, the content rather finely and evenly
granular when fresh, homogeneous in herbarium specimens or with a
few minute scattered oil globules in the ends.
Hasitat: Apothecia arising from the veins, petioles, and parenchyma
of decaying leaves: in N. Am. on Robinia Pseudoacacia L. and sp.
indet.; European specimens listed on Populus nigra L., P. tremula L.,
Quercus pedunculata Ehrh., Quercus sp., and Ulmus campestris L.; re-
ported by Velenovsky on Betula sp., Car pinus sp., and Quercus sp., but his
fungus almost certainly was incorrectly determined.
DISTRIBUTION: Massachusetts, New Jersey, and Minnesota — June—Oct.
146 Fartowia, Vou. 1, 1943
23; in Europe from Germany, ? Czechoslovakia, Italy, and England —
October.
MATERIAL EXAMINED: North America: Mass.: “Ad fol: putresc: in humidis” June,
C. J. Sprague 1231, as Peziza indet. (FH-C).—On Robinia Pseudoacacia LL...
leaflets, racheae, and petioles, Wakefield, Oct. 23, 1942, D. H. Linder (FH).—N. J.:
On decaying leaves, etc., in muddy places, June & Sept. 1879, Ell. & Ev., N. Am.
Fungi 393 [as Ombrophila purpurascens Fr.] (FH).—Minn.: On decaying leaves
on wet ground, Purgatory Swamp, Hennepin Co., Sept. 20, 1918, E. J. D. [as Helotium
sp. but filed under H. epiphyllum] (CUP-D 10656).— Europe: Germany: “Ad
Populi tremulae folia putrida, raro. Aut. Boss pr. Eberbach.” Fuckel, Fungi Rhen.
2388, type (FH).— Portion of same collection ? Herb. Barb.-Boiss. 1215 (FH).—
“Auf Populus tremula. Zehlandorf bei Berlin.” Sydow. Myc. March. 678 [as H.
epiphyllum Pers. var. candidum Karst.| (FH).—“An Quercus Blattern. Berlin,
Wannsee.” 10,1894, Sydow, Myc. March. 4258 [as H. conformatum] (FH).— “Auf
Quercus pedunculata. Berlin, Wannsee.” 10,1895, Sydow, Myc. March. 4436 [as H.
conformatum] (FH).— “Auf faulenden Blattern von Quercus pedunculata. Branden-
burg: Bredower Forst bei Nauen.” 7-10-’06, Sydow, Myc. Germ. 600 [as H. con-
formatum] (FH).—TItaly: “Padova, in foliis putridis Ulmi campestris, Oct. 1878.”
Sacc., Myc, Ven. 1387 [as H. epiphyllum] (FH).
The distinctive character of the species is its trapezoid spores which
are symmetrical in outline and very constant in size and shape. The
apothecia of the one and only collection I have seen in the fresh condi-
tion were pure white. On drying they changed shape but little, and after
about three weeks in the herbarium have turned no darker than a very
pale yellow: the stipe and receptacle are “Cream Color” (R) and the
hymenium “Maize Yellow.” In the Minnesota collection cited, taken
about 25 years ago, the apothecia are bright yellow. In most of the
older herbarium material, however, they are pale to deep orange.
The species is of the same general aspect as H. carpinicolum to which
it corresponds closely in color, size, structure, and habitat. It differs,
however, in being somewhat larger, distinctly stipitate, and finally and
more definitely in its characteristic, though not pronouncedly, trapezoid
spores.
Some confusion has been caused by Rehm’s reference of this species
to synonymy under H. conformatum Karst. to which it is not at all re-
lated. Karsten’s species as indicated by his description and his Fung.
Fenn. Exsicc. 738 is a species of Rutstroemia and I hope to deal with it
in another paper in the near future. Rehm notes that he did not see
Karsten’s specimen and that his interpretation is based only on the
description. Study of Rehm’s description together with an examination
of the exsiccati material which he cites (except Ascom. 152, which is not
available) indicates that he actually was working with Helotium immuta-
bile, not H. conformatum. Nannfeldt’s illustration [sub nom. H. con-
formatum], which was made from Sydow’s Myc. Germ. 600, also repre-
sents H. immutabile.
Velenovsky in his Discomycetes of Bohemia illustrates side by side
two species, one of which he refers to H. conformatum with H. immutabile
WHITE: STUDIES IN GENUS HELOTIUM. III 147
indicated as synonymous, and the other he describes as new, H. trapezoid-
eum Vel. The material which he identifies with H. conformatum appears
almost certainly to be neither that species nor H. immutabile. On the
other hand his H. trapezoideum is on leaves of Populus as was Fuckel’s
type of H. immutabile; this, to be sure, is of doubtful significance, but
his drawings show the characteristically shaped spores of the latter and
leave little doubt that H. trapezoideum Vel. is referable to synonymy
under H. immutabile Fuckel.
Sydow’s Mycotheca Marchica 678 labeled H. epiphyllum (Pers.) Fr.
var. candidum Karst. is cited above as H. immutabile since it is in com-
plete agreement with the type of the latter. It is possible that it also
correctly represents Karsten’s variety, but no authentic material is avail-
able for comparison. The specimen reported from Quebec (Mycologia
31: 730) as H. immutabile appears inseparable from H. epiphyllum
though it has smaller spores. A Michigan collection sent me by Dr.
Kanouse (A. H. Smith 32-103) as H. conformatum, and presumably
representative of, though not one of, the three collections reported under
that name by her (Papers Michigan Acad. 19: 97, 1934) has spores that
are less regular in shape than those of H. immutabile and it also lacks
croziers, otherwise it appears similar. Reports of “H. conformatum
Karst.” from China have not been investigated.
4. Helotium contortum White, sp. nov.
FIGURE 4.
Apotheciis sparsis, sessilibus, 2-5 (-7) mm. diam. in statu vegeto mollibus, carnosis,
albis vel sordidis, hymenio convexo; apotheciis exsiccantibus tenuibus, 1-3 mm. diam.,
sordide-ochraceis, concavis; receptaculis levibus, complanatis basis latis adhesis; para-
physibus ad basem rectis vel l-ramosis, ad apicem inaequabiliter breve-ramosis; ascis
clavato-cylindricis, 70-90x 8-9 ; ascosporis uniseriatis vel biseriatis, obovatis,
8.5-12 x 44.5 uw.
Apothecia widely scattered or more rarely subgregarious, entirely
sessile; when fresh 2-5 (—7) mm. diam., soft, fleshy, watery, dirty white
to more typically dull dilute yellowish or brownish, all parts externally
and internally exactly concolorous, the hymenium convex; on drying
contracting to 1-3 mm., becoming thin, soiled-ochraceous, dish-shaped;
receptacle entirely smooth, attached by a broad central portion from
which a few whitish hyphae sometimes radiate superficially on the sub-
stratum, free toward the margin, turning away from the substrate on
drying; margin in the fresh condition cuneate, even, directed outward
and downward, on drying turning up, becoming inrolled; in section
compact, homogeneous, composed throughout of large thin-walled
isodiametric cells 15-30 » diam. or up to 45x25 yp except in an inter-
mediate zone just underneath the margin where they are narrower,
elongate in the direction from base to margin forming a narrow layer
of prosenchymatous tissue with globular cells on either side; paraphyses
once- or twice-forked below the middle and with two or three very short
148 Fartowlia, VoL. 1, 1943
branches at the immediate apex, the tips scarcely or not at all enlarged;
asci_ clavate-cylindric, 70-90x8-9 4, originating from croziers;
ascospores irregularly uniseriate or biseriate, obovoid, 1-celled, 8.5-12 x
44.5 p, the content not granular.
Hasitat: Apothecia arising from the veins and parenchyma of water-
saturated leaves of Acer sp. and Alnus sp. in low wooded swamps.
DistriBuTION: New York and Michigan. June 14—July 10.
MATERIAL EXAMINED: N. Y.: Labrador Lake, on Acer, July 10, 1937, Whetzel,
Viegas, & White (FH).— Mich.: Au Train Lake, Au Train, on Alnus, June 14, 1933,
A. H. Smith 33-476, type (OTB).
The species is characterized by the rather large, broadly sessile, fleshy
apothecia which contract and become more or less distorted on drying,
the branched paraphyses, and the pseudoparenchymatous structure of
the disc tissue.
5. Helotium Conocarpi Seaver & Waterston, Mycologia 34: 517, fig. 1, upper. 1942.
FIGURE 5.
Apothecia scattered to subgregarious, erumpent, when young and in
the dried condition, smooth, shiny, nearly black, subglobose and sub-
sessile and opening by a minute apical pore, later expanding, becoming
discoid, substipitate, fleshy, and at maturity dark brown to the naked eye,
0.5-0.8 mm. diameter (up to 3 mm. fide Seaver, perhaps when fresh) ;
stipe stout, scarcely more than a papilla, rarely cylindric and distinct,
barely raising the receptacle off the substratum, close to “Cinnamon Brown”
(R); receptacle marked and colored like the stipe, becoming smooth
and much darker toward the margin at least when dry; margin in dried
material smooth, entire, tumid and upturned until complete maturity then
approximately even with the hymenium; hymenium with a delicate lilac
tint (Seaver) when fresh ?, dark brown when dried; in section the
hymenium about 75 » thick, the exposed surface brown due to color in the
tips of the paraphyses; hypothecium scarcely distinct; medullary re-
gion composed of thin-walled, short-celled hyphae compact and parallel in
the region of the stipe, directed outward and upward toward the margin
and loosely interwoven in the subhymenium; eectal excipulum distinct
about 50 y» thick at the base, tapering to 25 » at the margin, composed
of isodiametric cells 5-14 ,» diameter, compact, with walls scarcely
thickened and very slightly gelatinously modified, the outer exposed cells
brownish and rather uneven thus imparting the furfuraceous external
surface; paraphyses simple or once or twice forked at or below the
middle, clavate-enlarged at the apex, longer than the asci, 3-4.5 » diam.,
distinctly brown; asei clavate-cylindric, 55-65 x 7-8 ,, not originating
from croziers; ascospores biseriate or sub-biseriate, more rarely in one
row, obovoid, 7.5-10 x 3.3-4 yw, the content not granular.
Hapirat: On dead leaves of Conocarpus erectus L.
DISTRIBUTION: Bermuda.
WHITE: STUDIES IN GENUS HELOTIUM. III 149
MATERIAL EXAMINED: St. David’s Island, Dec. 7, 1940, F. J. Seaver & J. M. Waterston
(FH).
The description presented above is based on that of Seaver, together
with an examination of a portion of one of his two original collections
(cited by him as 375 and 425), which he kindly sent me, and which has
been deposited in the Farlow Herbarium. In his letter to me, Dr. Seaver
questioned his published generic disposition of this species. I also would
question it: the dark brown color, coriaceous texture, and colored para-
physes preclude its acceptance in Helotium, even in the all-inclusive
sense in which I am treating that genus at the present time; and these
same characters place it in the Ciborioideae in the vicinity of Rutstroemia
and Pycnopeziza. It should eventually find a place in Pyenopeziza (see
White & Whetzel, Mycologia 30: 187-203, fig. 1-21. 1938; also Whetzel
& White, Mycologia 32: 609-620. 1940), where it differs only in minor
characters from P. pachyderma, occurring on the fallen leaves of various
trees in North America and Europe, and from P. Sejournei, occurring on
the leaves of Hedera Helix L. in central Europe. The brown paraphyses
and perhaps the darker color separate it from both P. pachyderma and
P. Sejournet, and from the latter it is further distinct in its smaller asci
and spores.
Sections demonstrate definitely that there is no trace of a stroma in
the leaf tissue, and careful search fails to reveal any conidial stage or
spermatia. Study of the species in pure culture is desirable.
6. Helotium translucens sp. nov. e
FIGURE 6.
Apotheciis sparsis vel subgregariis, stipitatis, subcarnosis, subhyalinis in statu
vegeto, dum autem exsiccantibus albido-discoloribus vel pallide ochraceis, usque ad
0.3 mm. altis ac disco 0.3-0.4 mm. diam., in statu sicco, colore eodem fere gaudentibus,
in statu vegeto et exsiccato, haud plicatis neque distortis; stipite distincto sed brevi,
mediocriter crasso, cylindraceo, levi, stricto, abrupte in discum dilatato; disco crasso;
receptaculo levi; hymenio pori ope aperto, mox applanato et exsiccante persistenter
plano; paraphysibus leviter clavato-dilatatis, ad apicem 3-3.5 yp, diam.; ascis cylindraceo-
clavatis, 75-95 x 10-11.5 u, poro iodi ope caerulescente; ascosporis uni-seriatis vel
sub-biseriatis dispositis, obovatis, unicellularibus, 12-15 x 44.5 w, intus vix vel non
granulatis.
Apothecia scattered to subgregarious, stipitate, subfleshy, practically
hyaline when fresh, drying discolored-whitish to pale ochraceous, up to
0.3 mm. high and 0.3 to rarely 0.4 mm. across the disc when dry, of
about same color and consistency on all parts in both fresh and dried
condition, not wrinkled or distorted on drying; stipe distinct though short,
of medium thickness, cylindric, smooth, straight, expanding abruptly into
the disc; dise thick; receptacle smooth; hymenium opening by a pore,
soon plane, remaining plane on drying; paraphyses slightly clavate-
enlarged, 3-3.5 » diam.; asei cylindric-clavate, the thickness carried well
down toward the base and rather abruptly narrowed at the septum,
79-95 x 10-11.5 p, the pore blue with iodine; ascospores uniseriate to
150 FarLow1A, VoL. 1, 1943
sub-biseriate, obovoid, 1-celled, 12-15 x 44.5 y, the content scarcely or
not at all granular.
Hasirat: Apothecia arising from the parenchyma, lateral- and net-
veins of leaves of Acer sp. and Quercus sp.
DISTRIBUTION: Massachusetts. November.
MATERIAL EXAMINED: On decaying leaves of Acer sp., swamp at east side of Great
Blue Hills, near Milton, Nov. 11, 1941, Linder, Seeler, & White, type (FH).—A
second collection, same substrate, etc., White (FH).— On fallen leaves Quercus, Stony
Brook Reservation, West Roxbury, Nov. 11, 1941, White (FH).
In habitat and in color and consistency of the apothecia Helotium
translucens is somewhat similar to H. caudatum (Karst.) Vel., but it
differs in occurring later in the season, being smaller in size though more
fleshy, and in having smaller asci and smaller and differently shaped
spores. The apothecia in the dried condition are somewhat lighter in
color than the surface of the leaves on which they occur, yet not strik-
ingly so, which together with their small size renders them practically
invisible to the unaided eye. The spores are similar to those of H.
immutabile.
7. Helotium fastidiosum Pack, Ann, Rep. New York St. Mus. 27 (1873): 107.
1875. — Sace., Syll. Fung. 8: 221. 1889. — Mains, Mycologia 31: 730. 1939. —
White, Mycologia 34: 165-168, fig. 6-9. 1942.
Calycina fastidiosa (Peck) Kuntze, Rev. Gen. Pl. 3°: 448. 1898.
FIGURE 7,
Apothecia small, slender, delicate, stipitate, 1-1.5 or rarely 3 mm.
high, 0.8-1.5 mm. across the disc; stipe very slender, subcylindric,
smooth, dilute white to practically hyaline, sometimes more or less dis-
colored toward the base, thickened slightly just below the disc, expanding
abruptly into disc, not changing on drying except for a deepening of
the color to a hyaline-yellow or pale creamy yellow, sometimes becoming
pale ochraceous especially on the lower portion; dise flat-expanded at
maturity, medium thin; receptacle smooth, dilute white to whitish-
yellow, rarely varying to pale ochraceous with age, not changing on
drying except for a slight deepening of color; hymenium slightly convex
with maximum moisture content, opaque, white or more often pale yellow,
on drying becoming plane to saucer-shaped, waxy, the color deepening
somewhat to pale or bright yellow, approximating “Apricot Yellow”
(R), or more rarely as dark as “Deep Chrome”; paraphyses simple,
slightly or not at all enlarged at apex, 2.5-3.5 » diam.; asei not orig-
inating from croziers, clavate, 80-100x 9-12 un; ascospores elongate,
clavate, slightly curved and rounded at the upper end, straight and
pointed below, 26-36 x 3.5-4.5 y, biseriate, each containing a single row
of from 6-10 oil globules.
Hasitat: On petioles and midveins of decaying leaves of Alnus incana
(L.) Moench., Alnus sp., and decaying catkins of indet. species, probably
Alnus.
DIsTRIBUTION: Quebec, New York, and Oregon. Aug.—Sept.
WHITE: StupieEs IN GENUS HELotTium. III iru
MATERIAL EXAMINED: See Mycologia 34: 167. 1942.
In general aspect it is similar to the several species following, but
the long spores distinguish it easily from all other known foliicolous
forms.
8. Helotium caudatum (Karst.) Vel., Monogr. Discom. Bohem. 1: 206, pl. 20,
fig. 35. 1934.
Peziza caudata Karst., Fungi Fenn. Exsicc. 547. 1866. — Karst., Not. Soc. Fauna
Fl, Fenn. 10: 144. 1869.
Helotium scutula (Pers. ex Fr.) Karst. var caudatum (Karst.) Karst., Not. Soc.
Fauna Fl. Fenn. 11: 234. 1871. — Karst., Bidr. t. Kannedom af Finl, Nat. o.
Folk 19: 112. 1871.— Migula in Thomé’s Krypt.-Fl. 3: 1200. 1913.
Helotium naviculasporum Ellis, Bull. Torrey Club 5: 46. 1874.— Cooke, Bull.
Buffalo Soc. Nat. Hist. 2: 299. 1875.—Sacc., Syll. Fung. 8: 211. 1889.
Ellis, Cat. Pl. New Jersey, p. 551. 1890.— Rehm in Rabenh. Krypt.-Fl. 1°:
780. 1893. — White, Mycologia 34: 169, fig. 3, 11. 1942.
Helotium saprophyllum Cooke & Peck in Peck, Ann. Rep. New York St. Mus.
29 (1875): 55. 1878.— Nom. nud. Cooke, Bull. Buffalo Soc. 3*: 23. 1875.
Sacc., Syll. Fung. 8: 227. 1889.— [Mycologia 33: 573. 1941.]
Calycina naviculaspora (Ellis) Kuntze, Rev. Gen. Pl. 3°: 448. 1898.
Calycina saprophylla (Cooke & Peck) Kuntze, Rev. Gen. Pl. 3°: 449. 1898.
Helotium sparsum Boud., Hist. Classif. Discom. Eur., p. 111. 1907.— Boud.,
Icones Myc. pl. 495. 1905-10.— Rea, Brit. Myc. Soc. Trans. 4: 317, pl. 9.
1913. — Ramsb., Brit. Myc. Soc. Trans. 4: 373. 1913.—Sacc., Syll. Fung.
22: 649. 1913. Kanouse, Papers Michigan Acad. 22 (1936): 119. 1937.
Phialea phyllophila (Desm.) Gill. var. Jaapii Rehm ex Jaap, Abh. Bot. Ver.
Prov. Brandenburg 49: 10. 1907.— Sacc., Syll. Fung. 22: 655. 1913.
FIGURE 8.
Apothecia scattered, sparse, delicate, slender-stipitate, hyaline-white
when fresh, becoming opaque and more or less yellowish or pale ochra-
ceous on drying, reaching a height of 0.75—1.5 mm. and a diameter across
the disc of 0.4-1.3 mm.; stipe slender, delicate, smooth, dilute white
when fresh, sometimes yellowish at the substratum, drying semi-opaque
and white to creamy yellow; dise at first elongate, thick, the outside
wine-glass shaped, plane, then expanding, at maturity of medium thick-
ness, emarginate; receptacle smooth, hyaline to dilute white when
fresh, drying opaque and yellowish; hymenium plane from the very
young stages to complete maturity, finally somewhat convex with maxi-
mum moisture content, tender, white, more opaque than stipe and re-
ceptacle, drying waxy and pale yellow to ochraceous, remaining plane
or nearly so; paraphyses simple, slightly or not at all enlarged at apex,
2-3 » diam.; asei clavate, not or very rarely originating from croziers,
85-95 (-110) x8-1l ,»; ascospores elongate, slightly curved, obtuse
above, more or less pointed at the lower end, 18-25 x 4.5-5.5 yp, typically
more or less filled with irregular refractive granules, sub-biseriate.
Hasitat: On fallen decaying leaves, the apothecia usually originating
from the midrib or veins, more rarely from petioles or parenchyma of
the leaf blade. American specimens on Acer sp., Alnus incana (L.)
Moench, Carya sp., Fagus grandifolia Ehrh., ? Prunus sp., and Tilia
152 Fartow1a, VoL. 1, 1943
americana L.; listed from Europe on Acer campestris, Alnus glutinosa,
Alnus sp., Betula sp., Corylus sp., Carpinus sp., Fagus sylvatica, Fagus
sp., Populus alba, P. canadensis, P. canescens, P. tremuloides, Populus
sp., Quercus sp., and Salix sp. I have observed it frequently in the
vicinity of Ithaca, New York, where it appears to be confined to low
moist woods and swampy areas in which the leaves are wet or at least
moist most of the time; alder swamps are a favorite habitat.
DisTRIBUTION: Common in New York from July through September,
also found in New Jersey, Michigan, and Ohio; common in Europe where
it is known from Finland, Czechoslovakia, Germany, France, and England.
MATERIAL EXAMINED: North America: N. Y. (All as Helotium saprophyllum) :
Lake Pleasant, on maple leaves, Peck, type of H. saprophyllum (CUP-D 5987, NYS).
— Indian Lake, August, Peck (CUP-D 5989, NYS).— Adirondack Mts., Peck (CUP-D
5990, NYS).— Lowville, on fallen leaves Tilia americana. Sept., Peck (NYS).—
Sand Lake, July, Peck (CUP-D 5832).— Ridgway, Orleans Co., on dead leaves in
woods, 1904, C. E. Fairman (CUP-D 462, NYS).— Lloyd Preserve, McLean, on de-
caying leaves Alnus incana, Oct. 20, 1935, White & Whetzel (FH).— Old Forge,
Adirondack Mts., on fallen leaves ? Prunus, Sept. 3, 1936 (CUP 25515, FH) —
Labrador Lake, on decaying leaves, mostly Acer, in wooded swamp, July 10, 1937,
White, 2981 (FH).— On Carya, midribs, lateral and net veins of leaves in good state
of preservation, Sept. 14, 1938, Whetzel, White, & Sproston [Atypical spores] (CUP
27890, FH, US).— Little Ravine, below Newfield, on petioles, midribs, lateral & net
veins of well decayed leaves of ? Ulmus, Sept. 14, 1938, Whetzel & White (CUP
27891, FH).—V. J. (All as Helotium naviculasporum): “Ad folia putrescentia
Aceris.” Ell., Fungi Nov.-Caes. 53: (NY).— Newfield, on maple leaves on wet ground,
Ell, N. A. F. 62 (CUP-D 2972, NY).— Newfield, on old leaves decaying in stagnant
water, 23 June, 1876, Ellis (CUP-D 8465).— Newfield, on decaying leaves lying on
wet ground in swamp, July, 1876, Ellis (FH).— Mich.: Marquette, on decaying
leaves. 9-10-34, E. B. Mains 34-144, as H. sparsum (FH).— Ohio: Preston, 1902,
A. P. Morgan [as H. phyllophilum] (CUP-D 2126).— Europe: Finland: “Mustiala,
pa formultnande aspblad, Okt. 1865.” Karst., Fungi Fenn. Exsicc. 547, type of
Peziza caudata (FH).— Germany: “An Spiraea Blattern,” Berlin, Steglitz. 10.1892,
Sydow, Myc. March. 3772 [as H. sordidatum] (FH).—“An Alnus glutinosa.”
Berlin, Wannsee, 10.1892, Sydow, Myc. March. 3768 [as H. phyllophilum] (FH).—
“An Populus alba,” Berlin, Lichterfelde, 10.1892. Sydow, Myc. March. 3770 [as H.
phyllophilum] (FH).—“Auf Fagus sylvatica.” Steglitz bei Berlin, 6.1893, Sydow,
Myc. March. 4163 [as H. phyllophilum] (FH).—“Auf Alnus glutinosa.” Berlin,
Wilmersdorf, 11.1895. Sydow, Myc. March. 4437 [as H. phyllophilum f. albescens]
(FH).— “Auf Populus canescens.” Berlin, Thiergarten, 11.1895, Sydow, Myc. March.
4438 [as H. phyllophilum] (FH).—“Pappelblattern.” Prater, Wien, 16/X/1901,
Von Hohnel [as H. phyllophilum] (FH-H).— “Auf den Nerven faulender Blatter von
Populus canadensis Michx. Proy. Brandenburg: Triglitz in der Prignitz.” 10.X.1905,
Jaap, Fungi Sel. Exsicc. 152 type of Phialea phyllophila var. Jaapii (FH).—
France: “Feuille et petioles Alnus.” Envy. le Rouen, 8-1889 [as Helotium scutula var.
albida] (FH-P).
Taxonomically this is one of the more difficult members of the genus,
differing from its neighbors in characters which are for the most part a
Fig. 4-9: 4, Helotium contortum, from type (OTB) ; 5, H. Conocarpi, from spec.
cited (FH); 6, H..translucens, from type (FH); 7, H. fastidiosum, from type
(CUP-D 5948) ; 9, H. Linderi, from type (FH). Fig. 4 1500 approx.; all others
x 1290: Soe ey | | au
StupiEs IN Genus HeEtotium. III
.
.
WHITE
Ficures 4-9,
154 FarLowlA, VoL. 1, 1943
matter of degree and not to be considered very reliable. First of all
it is doubtfully distinct from H. scutula (Pers. ex. Fr.) Karst., which is
common and widely distributed on dead herbaceous stems; while generally
more delicate and less deeply colored than the latter, it appears difficult
nevertheless to find any morphological characters (cf. White, Mycologia
34: 158, fig. 1-3, 166, fig. 6, 11. 1942) on which to make a really
meaningful distinction, and the separation here maintained, for the
present at least, is largely arbitrary, those forms on leaves being referred
to H. caudatum and those on herbaceous stems to H. scutula. The larger
spores appear to separate H. caudatum sufficiently from H. albidum (Rob.
ex Desm.) Pat. & H. albopunctum Peck. The beautiful and very ample
material of Petrak, Fl. Bohem. Morav. Exsicc. 236 issued as H. caudatum
has spores that are rather small for that species and is reserved for further
study.
9. Helotium Linderi sp. nov.
FIGURE 9,
Apotheciis dispersis, sparsis, stipitatis, ad 1.5 mm. altis, disco circa 1 mm. diam.,
in statu vegeto albis, discis exsiccantibus cinereis vel nigrescentibus; stipitibus
gracilibus, exsiccantibus hyalinis ochraceis, levibus, sublucidis; discis subcarnosis,
poris apertis, obpiriformibus, expansis; receptaculis levibus, hyalinis, in statu vegeto
ochraceis, cinereis vel subnigrescentibus; hymenio plano, exsiccante nigrescente et
subconcavo; paraphysibus simplicibus vel ad basem ramosis, subclavatis, circe 3 u.
diam.; ascis clavatis, 65-75 x 9-10 wu; crozier nul; ascosporis biseriatim dispositis,
elongatis, subclavatis, leviter curvatis, guttulis 3-6.
MATERIAL EXAMINED: Tenn.: On fallen leaves, Chimney Trail, Great Smoky Mt.
Nat. Park, elev. 3200 ft., Aug. 18, 1939, type, D. H. Linder (FH).— On fallen birch
leaves, moist brook bed, Indian Gap, Great Smoky Mt. Nat. Forest, Aug. 20, 1939,
D. H. Linder (FH).— Wise.: Blue Mounds, 8-1-08 (Univ. Wisc.-Herb., R. A. &
A. M. Harper 410).
Differs from all the other slender-stipitate species in that the hymenium
and upper part of the receptacle becomes dark on drying —varying from
WuiteE: StupiEs In Genus HeEtotium. III 155
cinereous to brownish or nearly black —commonly about the color of
the pore surface of Polyporus adustus (Willd.) Fr.
The two Tennessee specimens recorded above appeared in the report
of the 1939 foray as Helotium saprophyllum (Mycologia 33: 573. 1941).
The Wisconsin specimen appears to be the basis of the record by Dodge
(Trans. Wisconsin Acad. 27: 1035. 1914) under the name of H. sordid-
atum Karst. No material of the latter is available for comparison, but
that it does represent that species is extremely doubtful.
10. Helotium albopunctum Peck, Ann. Rep. New York St. Mus. 31(1877): 47.
1879; non (Desm.) Bucknall, Bristol Nat. Soc, Proc. 3: 137. 1882. — White,
Mycologia 34: 172, fig. 13. 1942.
Pezizella albopuncta (Peck) Sacc., Syll. Fung. 8: 276. 1889.— [Cummins,
Papers Michigan Acad. 11(1929): 111. 1930].
Hymenoscypha albopuncta (Peck) Kuntze, Rev. Gen. Pl. 3°: 485. 1898.
FIGURE 10.
Apothecia scattered, not numerous, sometimes subgregarious in groups
of two to five, usually not more than five on a single leaf, rarely rather
abundant, very small, slender, delicate, stipitate, up to 1 mm. high and
0.8 mm. diam. across the disc, usually smaller, when fresh creamy-white
to yellowish-stained, drying ochraceous-yellow, all parts approximately
concolorous, waxy-cartilaginous when dried; stipe very slender, smooth,
sub-shiny, sometimes very short, commonly about twice as long as the
diameter of the disc; dise rather thick; receptacle smooth, when dried
sub-shiny and not wrinkled; hymenium plane from the earliest stages
to complete maturity, not changing shape on drying; margin obtuse,
circular, even, not elevated above the hymenium in the fresh condition,
rarely very slightly so on drying; paraphyses simple or more often
once or twice branched at or below the middle, clavately enlarged at the
apex, 2.5-3.8 » diameter; asci 60—70x 8-10 yp, not originating from
croziers; ascospores biseriate, mostly lying parallel with ascus,
14-16 x 3.2-4 p, oblong, flattened on one side, straight or slightly curved,
rounded above and sometimes bent abruptly to one side, obtusely-pointed
at the lower end, always 1-celled, containing when fresh 2-4 conspicuous
oil drops, these usually irregularly contracted in old herbarium speci-
mens or sometimes breaking up to form several smaller, scattered, re-
fractive granules.
Hasirat: Apothecia arising from the parenchyma tissue, or less fre-
quently also from the lateral and mid-veins of fallen leaves of Fagus
grandifolia Ehrh.
DistRIBUTION: New York, West Virginia, Virginia, and Georgia. A
mid- to late summer species.
MATERIAL EXAMINED: N, Y.: Adirondack Mts., Peck, type (CUP-D 5915, NY, NYS).
— Fall Creek, Ithaca, Aug. 8, 1895, Durand [as Helotium phyllogenon on oak leaves]
(CUP-D 921).— Same locality. etc. [as H. phyllogenon on Carpinus(?) ] (CUP-D
922).— Labrador Lake, near Tully, Aug. 26, 1935, Whetzel & White (CU-P 24821,
FH).— Labrador Lake. Aug. 12, 1936, Whetzel & White 2555 (CU-P 25464, FH). —
156 FarLow1a, Vou. 1, 1943
Labrador Lake, Aug. 12, 1936, Whetzel & White 2613 (FH).— W. Va.: Albright,
Aug. 4, 1908, J. L. Sheldon [as H. phyllophilum] (CUP-D 6502).— Va.: Laing
Trail near Mt. Lake, Sept. 3, 1936, J. A. Stevenson & W. W. Diehl [as Phialea
phyllophila (Desm.) Gill. f. fagicola Sacc.] (BPI, FH).— Cascades, near Mt. Lake,
Sept. 4, 1936, E. K. Cash [as Phialea phyllophila (Desm.) Gill. f. fagicola Sacc.]
(BPI).— Ga.: Bobbin Mill, Athens, July 13, 1940. J. H. Miller. [as Phialea phyllo-
phila (Desm.) Gill. f. fagicola Sacc.] (BPI).
Additional work on the foliicolous forms since the publication of my
paper covering the Peck species, demonstrates that Helotium albo punctum
is fundamentally a stipitate species. Peck notes of the type: “The stem
is so short that the plant appears sessile.” On the basis of this state-
ment, together with an examination of Peck’s specimen which had given
satisfactory evidence of its accuracy, I had previously formed a concep-
tion of H. albopunctum as a sessile form. It was largely this that pre-
vented the identification of later collections cited above with Peck’s
species, even though some of them had been studied prior to the prepara-
tion of the previous paper. In the type there are scarcely any apothecia
with stipes more than 0.1 mm. long. The single Georgia specimen cited
agrees in this respect. However, in all the others they range generally
from 0.2-0.8 mm., with only a very exceptional one as short as the longest
in the type.
The West Virginia specimen cited was incorrectly reported in a list
by Millspaugh (W. Va. Geol. Surv. 5 (A): 111.1913) as H. phyllophilum.
The Virginia material (comm. Miss Cash) was listed among the 1937
foray collections (Mycologia 29: 372. 1937) as Phialea phyllophila
f. fagicola, and the Georgia collection (also comm. Miss Cash) was re-
ported by Miller (Pl. Dis. Rep., Suppl. 131: 47, 1941) under the same
name. Saccardo’s forma fagicola is unknown to me; it may be a synonym
of Peck’s albopunctum.
There appears to be only one previous literature record under Peck’s
name and based on material other than the original: It was reported by
Cummins (1. c.) on leaves of ? Cornus from Montana. This specimen
has been loaned me by Miss Kanouse, and it proves to be entirely foreign
to Helotium, though I am unable to place it.
11. Helotium phyllogenon Rehm, Hedwigia 24: 14, 1885. — Massee, Brit. Fung.-Fl.
4: 258. 1895. — Boud., Hist. Classif. Discom. Eur., p. 112. 1907. — Ramsb.,
Brit. Myc, Soc. Trans. 4: 373. 1913. — Gola, Cat. Erb. Mic. Sacc., p. 120. 1930.
Peziza epiphylla Pers. ex Fr. var. populina Lasch ex Rabenh., Flora 38: 267.
1855.
Phialea phyllogena (Rehm) Sacc., Syll. Fung. 8: 274. 1889.
Hymenoscypha phyllogena (Rehm) Kuntze, Rev. Gen. Pl. 3°: 485. 1898.
Helotium phyllophilum auct. non (Desm.) Fr.— Rehm in Rabenh., Krypt.-F1.
1*: 796. 1893. — Migula in Thomé’s Krypt.-Fl. 10°: 1203. 1913.
FIGURE 1], .
Apothecia known only in the dried condition, scattered, solitary, not
numerous, delicate, slender-stipitate, ochraceous yellow, all parts more
or less concolorous, reaching a height of 1.5 mm. and a diameter across
WuiIteE: StTupIEs IN GENUS HELoTIUM. III 157
the disc of 0.5 mm., on moistening reviving fully and becoming entirely
colorless; stipe very slender, delicate, cylindric, smooth, flexuous when
long, slightly wrinkled; dise at first piriform, opening by a pore, later
flat-expanded, of medium thickness; receptacle smooth, concolorous
with stipe; hymenium saucer-shaped, waxy, concolorous with receptacle
or varying toward reddish or brownish, becoming plane on moistening ;
margin subacute, the sterile tissue not extending above the asci, or at
other times very slightly so and abruptly acute; paraphyses once-forked
near the base, very slightly clavate above, 3-3.5 » diam.; asci clavate-
cylindric, rather broad at both base and apex, 65-75 x 8.5-11 yw, not
arising from croziers, the pore blue in iodine; ascospores uniseriate to
sub-biseriate, narrow oblong-ellipsoid, 11-14x 3.8-5 p, never septate,
the content homogeneous, not granular or containing oil globules.
Hasirat: Apothecia arising from net veins or occasionally lateral
veins of leaves of Populus, reported on P. nigra L. and Populus sp. A
fall species — October (Hungary) and November 1 (Austria). Rehm
(1. c., p. 797) mentions a German collection on beech leaves; this probably
is a misdetermination.
DistRIBUTION: Material studied from Germany, Austria, and Hungary;
reported from England by Massee.
MATERIAL EXAMINED: Germany: “Ad folia putrida Populi nigrae pr. Dreisen.”
1854, Rabenh., Klotz. Herb. Myc. 1920, Type of Peziza epiphylla var. populina (FH).
— Austria: Prater, Heustadlwasser, 1 Nov. 1902, Von Hoéhnel [as Helotium phyllo- -
philum] (FH-H).— Hungary: “An faulenden Pappel-Blattern bei Ungarisch-Alten-
burg (Ungarn).” 10/1883, Linhart, Rehm, Ascom. 768, type of H. phyllogenon (FH).
It does not appear possible to separate this species on external morpho-
logical characters from H. phyllophilum (Desm.) Rehm, H. albopunctum
Peck, H. cawdatum (Karst.) Vel., or H. fastidiosum Peck. They all have
distinctive spore characters and these, along with substratum and geo-
graphic range, should enable one to place a given specimen without
too much difficulty; the contrasting characters are indicated in the key.
Rehm’s conception of H. phyllophilum (Desm.) Karst. appears to be
based largely, if not entirely, on his own H. phyllogenon; the latter he
refers incorrectly to synonymy under the former. It appears further
that Rehm was more or less generally confused with regard to these two
species and certain similar or related forms. For example, he cites Ascom.
768, type of H. phyllogenon, and also Linh., Fungi Hung. 386 as both
representing “Sehr schéne Exemplare . . ”” of H. phyllophilum; his
Ascom. 768 (FH) typifies H. phyllogenon as a good species, and Linhart’s
386 (FH) represents H. caudatum (Karst.) Vel.
Two specimens in the Durand Herbarium as H. phyllogenon (921, 922),
both taken on beech leaves at Ithaca, represent instead H. albopunctum
Peck.
12. Helotium phyllophilum (Desm.) Fr., Summa Veg. Scand., p. 356. 1849. —
Karst.. Symb. Myc. Fenn., Not. Fauna Fl. Fenn. 11: 239. 1870.— [Sacc.,
158 Far.towlia, Vou. 1, 1943
Michelia 1: 442. 1878].—[Phill., Man. Brit. Discom., p. 162. 1887].—
[Massee, Brit. Fung.-Fl. 4: 257. 1895].— [Rehm in Rabenh., Krypt.-Fl. 1°:
796. 1893. — [Massee & Crossl., Fung. Fl. Yorkshire, p. 285. 1905]. — Boud.,
Hist. Classif. Discom. Eur., p. 111. 1907.—[Millspaugh, West Virginia
Geol. Surv. 5(A): 111. 1913].— [Ramsb., Brit. Mycol. Soc. Trans. 4: 373.
1913].— [Dodge, Journ. Wisc. Acad. 17: 1035. 1914].— [Vel., Monogr.
Discom. Bohem. 1: 205. 1934].— [Mycologia 31: 730. 1939]. — [Ibid.
33: 573. 1941].
Peziza phyllophila Desm., Pl. Crypt. France, ed. 1, 1159. 1842; Ann. Sci. Nat.
2: 17: 96. 1842. — Str., Nom. Fung., p. 431. 1862.
Phialea phyllophila (Desm.) Gill., Champ. France, Discom., p. 105. 1869. —
Sace., Syll. Fung. 8: 254, 1889.— Oud., Ned. Kruidk. Archief. 3: 2": 208.
1902. — Oud., Champ. Pays-Bas, p. 345. 1904. — [Mycologia 29: 372. 1937].
Pezicula phyllophila (Desm.) Karst., Myc. Fenn. pars prima, Bidr, t. Kann.
af. Finl. Nat. o. Folk 19: 167. 1871.
Hymenoscypha phyllophila (Desm.) Kuntze, Rev. Gen. Pl. 3°: 485. 1898.
Allophylaria phyllophila (Desm.) Karst., Rev. Monogr., Acta Soc. Fauna FI.
Fenn. 2°: 131. 1885.— Nannf., Nova Acta Soc. Sci. Upsal. 4: 8°: 290. 1932.
FIGURE 12.
Apothecia scattered, very small, slender, stipitate, in the dried condi-
tion nearly the same color as the beech leaves on which they occur and
scarcely visible to the unaided eye; when young consisting of a very
slender subhyaline stalk, soon broadening at the top, clavate, then open-
ing by a pore; stipe extremely slender, cylindric, delicate, smooth, yellow-
ish-hyaline in the dried condition, not over 0.2 mm. long but, in relation
to the size of the apothecium, of medium length, slender, and distinct;
dise of same color and consistency throughout, at least so in dried speci-
mens, hyaline-yellow, shiny, thin, cupulate to flat-expanded, reaching
a diam. of 0.2 mm.; paraphyses simple, slightly or not at all clavate
at the apex, about 3 » diam.; asei arising from croziers, clavate, 65-75 x
8-9 pw; ascospores biseriate, narrow oblong-fusoid, always distinctly
l-septate, 11.5-15 x 3.2-4 y, the ends rounded, the content homogeneous.
Hasirat: Apothecia arising from parenchyma, lateral- and mid-veins
of both dorsal and ventral surfaces of leaves of Fagus sp. (also Acer.
sp. ?).
DisTRIBUTION: France. Known with certainty only from the type,
most, if not all, other reports being in error.
MATERIAL EXAMINED: “Hab. in foliis semiputridis Acerum et Fagorum.” Desm.,
Pl. Crypt. Fr., type (FH, FH-C).
The portion of the original material which is in the Desmaziéres set
at the Farlow Herbarium consists of two fairly large-sized leaves of
Fagus, both in a good state of preservation. On both leaves the apothecia
are fairly numerous; on one they are on the upper surface and on the
other the lower surface. The apothecia are only slightly more yellowish
than the surface of the leaf on which they occur, this together with their
minuteness, rendering them nearly invisible to the unaided eye. Under
a good binocular they are of a dilute yellow, tending toward hyaline,
and all parts are of the same color and consistency. On being moistened
WHITE: Stupies IN Genus He totium. III 159
they become colorless or practically so. My conception of the species,
and also the description presented above, are based entirely on this speci-
men, together with what appears to be a part of the same original collec-
tion in the Curtis herbarium. Its characteristic feature would appear
to be its extreme minuteness, dilute color, and consistently 2-celled spores
whose crosswalls are easily visible in ordinary KOH-phloxine prepara-
tions. Also, while too small for freehand sectioning, the disc in crushed
mounts exhibits a large amount of tissue composed of hyphae with walls
thickened and adherent, and the paraphyses appear to break easily, so
that it is impossible to trace one from its base to its apex. Desmaziéres
also indicates Acer as a substrate for the original material. No Acer
leaves have been seen. It is possible that if some of the packets of the
Desmaziéres number contain Acer leaves they will exhibit another species
of Discomycete.
See also discussion under Helotium albopunctum and H. phyllogenon.
Eighteen herbarium specimens — all that have been available — under
the name phyllophilum have been examined. Of these, eight represent
H. caudatum (Karst.) Vel., four H. albopunctum Peck, three H. popu-
linum Fuckel, two H. epiphyllum (Pers. ex Fr.) Karst., and one H.
phyllogenon Rehm. These are as follows: Herb. Barb.-Boiss. 1381
(FH) = H. populinum — Syd., Myc. March. 3768 (FH) = H. caudatum.
— Syd., Myc. March. 3769 = H. populinum.— Syd., Myc. March. 3770,
4163, 4437, 4438 (FH) = H. caudatum.—Tichener’s Glen, Canandaigua,
N. Y., Sept. 7, 1896 (CUP-D 916) = H. populinum. — Enfield, N. Y.,.
Oct. 19, 1901, as H. phyllophilum fide Rehm (CUP-D 1192) =H.
epiphyllum ? — Prater, Wien, 16/X.1901 (FH-H) =H. caudatum.—
Coll. of 1/11/1902 (FH-H) = H. phyllogenon. — Preston, Ohio, 1902,
Morgan (CUP-D 2126) = H. caudatum.— Coll. of 1904. “Sudl. Wiener
Bechen” (FH-H) = H. caudatum.— Albright, W. Va., Aug. 4, 1908,
Sheldon (CUP-D 6502) = H. albopunctum. — Duchesnay, Quebec (FH)
=H. epiphyllum.— Three specimens comm. Miss Cash = H. albo-
punctum.
13. Phialea Cassandrae Kanouse, Papers Michigan Acad. Sci. 20(1934): 71. 1935.
FIGURE 13.
Apothecia solitary, sparse, stipitate, slender, delicate, up to 2 mm.
high when fresh, and up to 0.5 mm. across the disc; stipe extremely
long and thin, in the dried condition flexuous, smooth, shiny, the lower
half or more black, the upper part translucent-white, sub-bulbous at the
base; dise in dried material thin, spreading at maturity, all parts con-
colorous with upper part of stipe, externally smooth, subshiny; para-
physes simple or more rarely once or twice forked at or near the base,
clavate-cylindric or cylindric, thin-walled, rounded at the apex, about
same length as the asci, with one or two crosswalls near the base, the
long upper cell scarcely or not staining, 3-4.5 » diameter; asei clavate,
160 FarLowia, Vou. 1, 1943
30-42 x 5-7 pw, the pore faintly blue with iodine; ascospores biseriate,
oblong, more or less pointed below, 6-8 x 1.4-1.8 pu, the content not
granular.
Hasirat: On fallen leaves of Chamaedaphne calyculata (L.) Moench.
DistRIBUTION: Michigan. Several collections with dates ranging from
May 22 to June 17.
MATERIAL EXAMINED: Mud Lake Bog, Whitmore Lake, May 22, 1933, 4. H. Smith
33-82, det. Kanouse, authentic (MICH.).
The description presented above is adapted from that of Dr. Kanouse
(I. c.) together with an examination of the specimen cited, which she
kindly loaned me. This and the preceding five species (nos. 5-12)
appear to form a closely knit natural group, the small spores separating
it at once from all the others. Its superficial appearance is similar to
that of H. Linderi, but it is more slender and delicate, and the lower
part of the stipe is black whereas in H. Linderi the disc is dark on dry-
ing and the base of the stipe white. It may be characterized as one of
the most beautiful and distinct of species. Its gross characters are ex-
treme slenderness and black stipe, and its microscopic characters the
broad, cylindric paraphyses, the small asci proliferating abundantly from
conspicuous croziers, and the small spores.
14. Helotium populinum Fuckel, Symb. Myc., p. 316. 1869. —Sacc., Syll. Fung.
8: 247. 1889.— [Seaver, Mycologia 3: 62. 1911].
? Peziza punctiformis Grev., Scot. Crypt. Fl. 2: 63, fig. 1-3. 1824; non Pers.,
Syn. Meth. Fung., p. 674. 1801; non Fr., Syst. Myc. 2: 105. 1822. — Str.,
Nom. Fung., p. 433. 1862.— (Under Calycellina) Nannf., Nova Acta Soc.
Sci. Upsal. 4: 8’: 194, pl. 10, fig. 4. 1932.
Helotium punctatum Fr.. Summa Veg. Scand., p. 356. 1849. — Berk., Outl. Brit.
Fung., p. 372. 1860. — [Sacc., Fungi Ital., fig. 1357, 1883.1 — [Sacc., Michelia
168. 1877.)
Helotium Ilicis Phill., Man. Brit. Discom., p. 164. 1887.— Sacc., Syll. Fung.
§: 241. 1889. — Massee, Brit. Fung.-Fl. 4: 260. 1895.
Helotium punctiforme (Grev.) Phill, Man. Brit. Discom., p. 168. 1887. —
Massee, Brit. Fung.-Fl. 4: 259, 1895.— Massee, Fung. FI. Yorkshire, p- 285.
1905.
Pseudohelotium punctiforme (Grev.) Sacc., Syll. Fung. 8: 295. 1889.— Oud.,
Champ. Pays-Bas, p. 347. 1904.
Pezizella punctiformis (Grev.) Rehm in Rabenh., Krypt.-Fl. 1°: 664. 1893. —
Starb., Bihang till k. Vet.-Akad, Hand]. 21: 29, 1895.— [Kanouse, Papers
Michigan Acad. 19 (1933): 96. 1934.1] — Vel., Monogr. Discom. Bohem. 1:
159; 2: pl. 10, fig. 16. 1934.
Pezizella tumidula auct. —Rehn in Rabenh., Krypt.-Fl. 1°: 666, 650, fig. 1-5.
1893. — Von Héhn., Mitt. Bot. Techn. Hochsch. Wien 3: 72, 105. 1926.
Pezizella populina (Fuckel) Rehm in Rabenh., Krypt.-Fl. 1°: 668. 1893. —
[Von Hohn., Mitt. Bot. Techn. Hochsch. Wien 3: 69. 1926.]
Calycina Ilicis (Phill.) Kuntze, Rey. Gen. Pl. 3°: 448, 1898.
Calycina populina (Fuckel) Kuntze, Rey. Gen. Pl. 3°: 448. 1898.
Calycella Hicis (Phill.) Boud., Hist. Classif. Discom. Eur., p. 95. 1907.—
Ramsb., Brit. Myc. Soc. Trans, 4: 371. 1913.
Hyaloscypha punctiformis (Grev.) Boud., Hist. Classif. Discom. Eur.., p. 126.
1907. — Anon., Brit. Myc. Soc. Trans. 7: 9, 1921,
Waite: Stupies In Genus Hetotium. III 161
Micropodia populina (Fuckel) Boud., Hist. Classif. Discom. Eur., p. 128. 1907.
Hymenoscypha punctiformis (Grev.) Schrét., Krypt.-Fl. Schlesien 3°: 71. 1908.
— Migula in Thomé’s Krypt.-Fl. 10°: 1151. 1913.
Hymenoscypha populina (Fuckel) Migula in Thomé’s Krypt.-Fl. 10°: 1152. 1913.
Calycellina punctiformis (Grev.) Hohn., [Sitzb. Akad. Wissensch. Wien, Math.-
nat. KI. 1: 127: 601. 1918].—[Von Hoéhn., Mitt. Bot. Techn. Hochsch.
Wien 3: 105. 1926.]
Calycellina populina (Fuckel) Hohn. [Sitzb. Akad. Wissensch. Wien, Math.-
nat. Kl. 1: 127: 601. 1918].—[Von Héhn., Mitt. Bot. Techn. Hochsch.
Wien 3: 105. 1926.]
Phialina puberula auct.— Von Hoéhn., Mitt. Techn. Hochsch. Wien 3: 106.
1926.
FIGURE 14,
Apothecia usually numerous, solitary or more rarely in groups of two
or three, unevenly distributed over large portions of the substrate sur-
face, tending in places to be gregarious; at first minute, globose, dirty
yellowish-white when fresh, narrowed toward the base, enlarging, a pore
appearing at the apex, the cavity deep, becoming more shallow as the
disc expands; maturing subfleshy, up to 1.1 mm. diam. in the fresh
condition, contracting slightly on drying, commonly 0.3—-0.6 (—0.8) mm.
diam. in dried specimens, sessile; receptacle entirely smooth, white when
fresh, seated on the substrate by a narrow central portion, pale yellow
to ochraceous on drying; hymenium creamy-white when fresh, plane
to slightly concave, or with maximum moisture varying to strongly con-
vex, drying waxy, pale to dull yellow, variable but commonly near
“Ivory Yellow” (R) or “Ochraceous Buff,” saucer-shaped; margin
circular to rarely more or less lobed, smooth when fresh and in mature
specimens not elevated, turning upward on drying, smooth or becoming
pruinulose; in section rather thick over point of attachment, gradually
thinner toward the margin, the hymenium 60-70 » thick, hypothecium
scarcely differentiated, medullary region rather open in structure, composed
of thin-walled, septate, interwoven hyphae, grading rather indefinitely into
an ectal layer which is rather thick at the base, becoming thinner toward
the margin, about 30-50 » half-way between base and margin, compact,
composed of more or less isodiametric cells, 8-15 » diam., thin-walled
or the outer ones near the substratum very slightly thickened; paraphyses
simple, 3-5 septate, very slightly enlarged upward, 3-3.8 p at the apex;
asci clavate, originating from croziers, 50-70 x 6-9 yp, the apex notice-
ably pointed, the pore bright blue with iodine; ascospores biseriate
or sub-biseriate, irregularly oblong-fusoid, slightly curved or flattened
on one side, 1-celled, 10-13 x 2.7-3.4 p, the content homogeneous or more
rarely with a few small oil droplets.
Hapitat: American specimens on Amelanchier intermedia Spach.,
Fagus grandifolia Ehrh., Platanus occidentalis L., Robinia Pseudoacacta
L., Tilia americana L., and sp. indet.; European specimens recorded on
Acer platanoides, Castanea vesca, Corylus sp., Fraxinus pubescens, Pop-
ulus canadensis, Populus pyramidalis, Quercus sp., Robinia macrophylla,
162 Fartow1a, Voi. 1, 1943
R. Pseudoacacia, Salix alba, S. daphnoides, Tilia sp., Ulmus sp., and sp.
indet.
DISTRIBUTION: Specimens studied from Massachusetts, New York and
Pennsylvania; not otherwise known from North America. European
material examined from Germany, France, and England; not known from
elsewhere. Sept. 6-Oct. 9 in N. Y. & Pa., in quantity when found.
MATERIAL EXAMINED: Mass.: On racheae & petioles of Robinia Pseudoacacia,
Wakefield, Oct. 23, 1942, D. H. Linder (FH).—N. Y.: On beech leaves, Tichener’s
Glen, Canandaigua, Sept. 7, 1896, Durand [as Helotium phyllophilum] (CUP-D 916).
—On hasswood leaves, Jasin’s Gull, Sept. 5, 1902, Durand, as H. punctiforme f.
Tiliae (CUP-D 1727).— On Amelanchier intermedia, Lloyd Preserve, McLean, Sept.
6, 1935, H. H. Whetzel (CU-P 29137, FH, BPI).—On decaying leaves sycamore,
Cayuta Lake, Sept. 9, 1938, H. H. Whetzel & W. L. White 3432 (FH).—Penna.:
On decaying leaves. Sycamore Mills Road, near Media. Oct. 9, 1936. Whetzel (CU-P
27978, FH).— Europe: Germany: “An der obern Flache diirren Blitter von
Pepulus pyramidalis auf der Miinchau bei Hattenheim.” Herb. Barb.-Boiss. 1217,
type of H. populinum (FH).—“‘Ad folia putrida, non raro. Automno. Oestrich
(Nassau). leg. Fuckel.” [as Pseudohelotium puberulum] Herb. Barb.-Boiss. 1220 (FH).
— “In foliis Quercus putridis ad Sugenheim Bavariae. leg. Dr. Rehm.” Rabenh., Fungi
Eur. 1121 [as Peziza tumidula] (FH).—‘Auf Castanea vesca. Lichterfelde bei
Berlin. 9.1888. leg. P. Sydow.” Myc. March. 2445 [as Pezizella tumidula f. Castanae]
(FH).—“Auf Corylus-Blattern. Lichterfelde bei Berlin. 9-1890. leg. P. Sydow.”
Myc. March. 3476, as Helotium punctiforme (FH).— “An Corylus, Ulmus-Blattern.
Berlin, Lichterfelde. 10.1892 leg. P. Sydow.” Myc. March. 3769 [as H. phyllophilum]
(FH).—“An Robinia macrophylla. Berlin, Lichterfelde. 10.1892. leg. P. Sydow.”
Myc. March. 3780, as Pezizella punctiformis f. foliicola (FH).—“‘An Salix alba.
Berlin, Lichterfelde. 10.1892. leg. P. Sydow.” Myc. March. 378] as P. punctifermis f.
foliicola (FH).—“An Robinia pseudoacacia. Berlin, Lichterfelde. 10.1892. leg. P.
Sydow.” Myc. March 3782, as P. punctiformis f, petiolicola (FH).—“An Salix
daphnoides. Am Wannsee bei Berlin. 10.1893. leg. P. Sydow.” Syd., Myc. March. 3960,
as P. punctiformis (FH).— “Auf Robinia macrophylla, Lichterfelde bei Berlin. 10.1893.
leg. P. Sydow.” Myc. March. 4046, as P. punctiformis f. petiolicola (FH). — “Fau-
lende Blattern von Acer platanoides. Lichterfelde bei Berlin.” 10.1893, Sydow, Rehm,
Ascom. 1162 [as P. puberula] (FH).—‘“Auf faulen Blattern. Wiener-Wald. 1/X.
1909 v. Hohnel.” [as P. tumidula Rehm (non Desm.)?] (FH-H).— “Auf faulenden,
vorjahrigen Blattern von Populus canadensis Michaux. Prov. Brandenburg: Triglitz
in der Prignitz. 8.X. 1909. leg. Otto Jaap.” Fungi Sel. Exsicc. 364 [as Helotium
conformatum] (FH).— “Auf Blattern von Fraxinus pubescens. Brandenburg: Baum-
schulen zu Tamsel. 25.9.1909, leg. P. Vogel.” Syd., Myc. Germ. 902, as Pezizella punc-
tiformis (FH).—On decayed leaves of Tilia, Naedaschutz, Saxony, Oct. 6. 1912,
G. Freulich. as P. punctiformis Rehm non Grev. (FH-H).— France: “Normandie:
Sur de vieilles; Lebiscy pres Caen. Roberge.” [as Phialea phyllophila] (FH).—
“Fol. sicc. Aceris platanoidis, etc. Caen. Coll. Roberge.” [as Peziza lachnobrachya]
(FH-C).— England: Carlisle, Carlyle, as Helotium punctiforme (CUP-D 4198,
NY-M).— Terrington, St. Clement, C. B. Plowright, Phill., Elv. Brit. 88 [as H.
epiphyllum] (FH).—On Holly, Shrewsbury, Phill., Elv. Brit. 134 [as H. epiphyllum
var. Illicina (nom, nud.) ]. Type of H. Ilicis (FH).
Greville’s specific epithet of 1824, due perhaps to the fact that it is
descriptive of numerous minute species, has become a “dumping ground”
for almost any species that might be described as “punctiform.” Since
it has been the subject of so much controversy and misinterpretation,
and since the other specific names included in the synonym are so little
known, the following historical sketch seems necessary.
WHITE: STupDIES IN Genus Hetotium., III 163
Greville’s Peziza punctiformis was described from Scotland and said
to be on the leaves of Fagus and Quercus. The essential features of the
description are as follows:
“P. yellow, very minute, gregarious, punctiform, globose, at length plane, the
margin crenulate.
“Bright yellow but extremely minute, gregarious, globose at first, with a small
central opening, at length becoming quite plane, or even slightly convex, the extreme
margin very thin and crenulate. The substance of the plant is carnose, and the
center rather thick for its size. The whole is glabrous. Sporuliferous cells linear-
oblong. Sporules yellow, difficult to be perceived.
“Were it not for a considerable number growing together, this species might
escape the eye of a practical mycologist. I have met with it several times and have
ascertained its characters to be constant. It not infrequently accompanies Phacidium
coronatum and Sclerotium quercinum.”
In Greville’s accompanying illustrations the apothecia are gregarious
but not crowded and occur on the dorsal surface of an oak leaf, especially
scattered in the vicinity of the midrib and the larger lateral veins. They
are at first globose with a small distinct central pore and are attached
by a fairly narrow central portion. As they expand they are rather fleshy,
flattened-globose with a pore and an apparently deeply cupulate central
hymenial cavity. The receptacle does not narrow to a stipe or a stipe-
like attachment, but rather it narrows gradually from the margin to the
central part where it is seated on the substrate. The apothecia are in
all stages bright yellow throughout, thick and fleshy for their size, and at
certain stages they exhibit a slightly roughened margin.
The interpretation I have placed on Greville’s species is that of Rehm,
and it is the one that has been general in continental Europe. In favor
of this general European interpretation is the morphological similarity
of the form to that in Greville’s illustration; against it is the fact that of
the large number of specimens available none are known with certainty
to be on oak leaves, which is the correct substratum for Greville’s species.
Finally Von Hohnel considered Greville’s species in some detail and
rejected the common European interpretation of it. He noted that while
Greville’s species was described as occurring on both Fagus and Quercus,
and might therefore be a mixed species, yet the apothecia illustrated
were on an oak leaf and for this reason we should refer to the species
of Greville a form known to occur on oak. Unfortunately, when Von
Hohnel applied Greville’s name to a form occurring on oak, he chose
a species that varied rather conspicuously from that of Greville in its
gross morphology. ‘This species is treated in the present paper as num-
ber 17, Helotium puberulum.
In order to get an idea of the British interpretation of Greville’s puncti-
formis I have referred to Massee’s herbarium at the New York Botanical
Garden. I find there four British entries under that name, and as nearly
as can be determined they represent three different species. Only one
specimen is there; the other three entries consisting only of notes and
104 FaRLowIA, VoL. 1, 1943
drawings. The specimen is the same as the material referred to puncti-
formis on the continent and is marked by Massee “not the typical form.”
In view of the foregoing, Greville’s name punctiformis should be dis-
carded. It may have been applied originally to any one or more of
several minute foliicolous species, and since the original specimen is lost
it is unlikely that a non-controversial interpretation could ever be placed
upon it.
The material of Herb. Barb.-Boiss. 1217 appears to be part of that
on which Fuckel based his description of H. populinum. Its apothecia
are rather sparse, are only partially opened, and apparently are largely
immature. They tend to be more deeply colored —more of a grayish-
ochraceous —than most of the later material which I have referred
here, but otherwise there is no difference.
Phillips’ type of H. Ilicis, distributed in his Elv. Brit. 134, as repre-
sented by the specimen in the Farlow Herbarium, is beautiful material
showing well developed apothecia on the leaves of Ilex. They exhibit
the distributional pattern typical for H. populinum, that is, they are
numerous but more or less unevenly distributed over the leaf surface,
tending in places to become gregarious. They occur on both surfaces
of the leaf but are predominantly on the under (dorsal) surface. They
are fleshy, deep yellow, concolorous on all parts, waxy, and are in all
stages of development, the largest sometimes becoming somewhat lobate
and a few ranging up to 0.8 mm. in diameter, the hymenium plane,
but the margin rather thick and obtuse and elevated above the hymenial
surface.
Both H. populinum and H. Ilicis have been very much neglected, most
collections referable here having been identified with Peziza punctiformis
Grev. or one of its later synonyms. In fact the only record found of
the identification of later material with either populinum or Ilicis is that
of a collection made by Seaver and Bethel in Colorado on Populus
tremuloides in 1910 and reported doubtfully by Seaver as H. populinum
in Mycologia 3: 62. 1911. This specimen, kindly loaned the writer by
Dr. Seaver, proves to be Mollisina acerina (Mout.) Hohn. (Mitt. Bot.
Techn. Hochsch. Wien 3: 67, 104. 1926), and represents the only col-
lection of the species which I have known from North America, although
several have been examined from Europe.
The species as here defined appears to be constant and well delimited,
the ascospores, especially, presenting a remarkable uniformity in both
size and shape.
The following exsiccati and other specimens under names which I
have listed in the synonymy of Helotium populinum are incorrectly de-
termined and are here reidentified as follows:
Krieg., Fungi Sax. 88] [as Pezizella punctiformis| (FH) = Mollisina
acerina (Mout.) Hoéhn.— Syd., Myc. March. 4041, 4042, 4043, 4044,
4045 [as P. punctiformis {. foliicola] (FH).—= Mollisina acerina (Mout.)
WHITE: StTupIEs IN Genus HELotTium. III 165
Hohn. — Spec. in Durand Herb. [as P. punctiformis f. fructicola] (CUP-D
1315, 1775) = An unknown species. — Syd., Myc. Germ. 125 [as P.
punctiformis] — An unknown species. — Spec. reported by Kanouse (1. c.)
on elm leaves from Michigan [as P. punctiformis] (Mich.) = An un-
known species.
15. Helotium tumidulum (Rob. ex Desm.) Massee [Fungus Fl. Yorkshire, p. 285.
1905].
Peziza tumidula Rob. ex Desm., Ann. Sci. Nat. 3: 16: 325. 1851.— Str., Nom.
Fung. p. 437. 1862.
Pezizella tumidula (Rob. ex Desm.) Sacc., Syll. Fung. 8: 276. 1889. — [Rehm
in Rabenh. Krypt.-Fl. 1°: 650, fig. 1-5, 666. 1893.] Von Hohn., Sitz.-
ber. Akad. Wien. Math.-nat. 1: 127: 599. 1918.— Mitt. Bot. Inst. Techn.
Hochsch. Wien 3: 72. 1926.—[Vel., Monogr. Discom. Bohem. 1: 160; 2:
pl. Il, fig. 54. 1934.] — [Kanouse, Papers Michigan Acad. Sci. 19 (1933):
96. 1934.]
Mollisia tumidula (Rob. ex Desm.) Hoéhn., Sitz-ber. Akad. Wien, Math.-nat.
Kl. 1: 115: 1284. 1906.
Hyalinia tumidula (Rob. ex Desm.) Boud., Hist. Classif. Discom. Eur., p. 104.
1907.
Hymenoscy pha tumidula (Rob. ex Desm.) Kuntze, Rev. Gen. PI. 3°: 487. 1898. —
[Migula in Thomé’s Krypt.-Fl. 10°: 1153, pl. 173, fig. 5-8. 1913.]
FIGURE 15.
Apothecia scattered or subgregarious, not crowded, erumpent, sessile,
minute, white when fresh (Desm.), in the dried condition gelatinous-
cartilaginous, dilute ochraceous, about the same color as the leaves on
which they occur, rising only slightly above the leaf surface, cushion-
shaped with a slightly concave hymenium, not over 0.2 mm. diameter;
on moistening becoming thick, showing a tumid excipulum and a
slight, thick, lacerate margin raised above the hymenium; paraphyses
more or less adherent to each other, not staining, strongly enlarged at
the apex to 4-5 » diam.; asei subcylindric, the pedicel short, thick, the
apex noticeably narrowed then truncate, the pore bright blue in iodine,
40—46 x 8.5-9.2 p», originating from croziers which are obscure;
ascospores oblong, slightly inequilateral, biseriate in the ascus, 12-13 x
2.8-3.3 , the ends rounded, the content homogeneous or with a small
amount of granular material in each end, always 1-celled.
Hapitat: Apothecia on the parenchyma of dead leaves of Betula sp.
Reports of its occurrence on other species are in error. |
DisTRIBUTION: France. Known only from the material of Desmaziéres.
MATERIAL EXAMINED: “Au printemps, sur les deux faces des feuilles 4 demi-pourries
du Bouleau.” Desm., Pl. Crypt. Fr. ed. 1, 2011; ed. 2, 1611 (FH, FH-C).
The original material of this species is very ample and in good con-
dition. The apothecia in the dried condition, especially at maturity,
are not strongly raised above the leaf surface and are rather thin. The
excipulum protrudes beyond the margin of the hymenium and turns
inward, lying close to and entirely obscuring the ascus layer. When
viewed in the dried condition only the enclosing excipular tissue is seen,
166 Fartowia, Vou. 1, 1943
plus some slight evidence of a central pore or depression. On moistening
the margin becomes erect and appears rather thick and lacerate and is
elevated above the central hymenial cavity.
Though several workers since the time of Desmaziéres have referred
material to this species, there is no evidence that anyone except Von
Hohnel has made a critical examination of the type. Most, if not all,
of this later material so referred has been wrongly identified. The
history of the species follows briefly:
Saccardo (1889) transferred it to Pezizella but suggested the genus
Cyathicula for it because of its lacerate margin. This was entirely on
the basis of the original description. Rehm treated it in 1893, reporting
it from two localities in Germany, and listing decaying leaves of Quercus
as the substrate. It is certain, as shown later by Von Héhnel, that Rehm
did not have the Desmaziéres species. Massee (1905) reported it on
oak leaves from two localities in England and transferred it to Helotium.
These collections may be in the Massee herbarium at the New York
Botanical Garden but I have not seen them. There is scarcely a possi-
bility that they are correctly identified. In 1906 Von Hohnel published
a fairly good redescription based on the Desmaziéres type and transferred
the species to Mollisia. He also noted that Rabenh. 1121, which was
cited by Rehm, and which was partly, if not wholly, the basis of Rehm’s
conception of tumidula, did not agree with the type and should be the
basis of another species which he would call Pezizella tumidula Rehm
non Desm. He further noted that Syd. Myc. March. 1853 distributed as
Helotium tumidulum was not in agreement with the type. In 1918 he
again noted that tumidula had not been used in its correct sense by Rehm,
and following up this in 1926 he referred tumidula sensu Rehm to what
he called the true Peziza punctiformis of Greville and made punctiformis
the basis of a new genus Calycellina. In 1934 Velenovsky reported
tumidula, obviously in error as indicated by his drawings, from Czecho-
slovakia on Quercus, Fagus, and Carpinus.
Rehm’s interpretation of tumidula is confused. Since he mentions at
least two different collections it is possible that he was working with two
—or possibly more, species. The one specimen which he cites spe-
cifically (Rabenh., Fungi Eur. 1121) is, as indicated by the specimen in
the Farlow Herbarium, closely allied if not identical with Helotium
populinum. It agrees perfectly with populinum in the characters of its
paraphyses, asci, and spores; it differs however in that the apothecia
are smaller, are much thinner and more closely adherent to the sub-
stratum, in the dried condition are translucent-ochraceous and cartilagin-
ous, and in its occurrence on oak leaves. It is strongly suggestive of
H. lucellum (Karst.) Karst., of which I have not been able to get au-
thentic material for comparison. On the other hand, the spore charac-
ters indicated by Rehm for tumidula— which may or may not have been
taken from the Rabenhorst specimen — are not those of populinum, but
WHITE: Stupies IN GeNUs HELOTIUM. III 167
could represent either lucellum or. puberulum. A slide in the Von
Hohnel herbarium made from Rabenh. 112] shows spores of lucellum or
puberulum —not populinum. It appears then that Rabenh. 1121 might
be a mixture of two species, one of which is puberulum (or lucellum).
This would account for Von Héhnel’s reference of tumidula sensu Rehm
to synonymy under punctiformis, the latter as Von Hohnel understood
it, but not according to the treatment of others which is accepted in this
paper, actually representing puberulum.
The single North American specimen (on oak leaves in Michigan) that
has been reported as tumidula is incorrectly determined and represents a
species which I am unable to name. Those available from Europe are
all wrongly determined and are here reidentified as follows:
Rabenh., Fungi Eur. 1121 (FH) = An atypical specimen of H. popu-
linum or a closely related form.— Syd., Myc. March. 1853 (FH) =A
mixture of Mollisina acerina (Mout.) Hohn. and Hyaloscypha lachno-
brachya (Desm.) Nannf,—Syd., Myc. March. 2444, 2445 (FH) =
Helotium populinum Fuckel.—Syd., Myc. March. 2446 (FH) = Hya-
loscypha lachnobrachya (Desm.) Nannf. — Spec. in Von Hohnel Herbar-
ium [as Pezizella tumidula Rehm (non Desm.) ?, Wiener-Wald, 1/X.1909
= Helotium populinum Fuckel.
16. Pezizella glagosa (Ell. & Ev.) Sacc., Syll. Fung. 8: 276. 1887.
Peziza glagosa Ell. & Ev., Journ. Myc. 4: 56. 1888.
Hymenoscypha glagosa (Ell. & Ev.) Kuntze, Rev. Gen. Pl. 3°: 485. 1898.
FIGURE 16.
Apothecia minute, numerous, evenly distributed and not crowded,
sessile, thick, white or whitish when fresh, reaching a maximum diameter
of 325 p», drying concolorous with the substrate surface and not visible
to the naked eye; receptacle smooth, when fresh translucent-brownish,
the sides nearly vertical from substrate to margin, slightly constricted at
point of attachment, contracting on drying and becoming more or less
turbinate, darker in color, shiny; hymenium in fresh condition white,
convex, drying plane or slightly concave, brown, shiny; paraphyses
noticeably thin, once forked below the middle and again at the apex,
the apices 2 »; asci clavate, originating from croziers, 70-75 x 7-8 yp;
ascospores biseriate, inequilaterally oblong or oblong-fusoid, 2-celled,
11-13 x 3.2-3.9 pn, the ends obtuse.
HasitaT: On decaying leaves of Nyssa sylvatica Marsh.
DisTRIBUTION: Massachusetts (Nantucket) and New Jersey. August.
MATERIAL EXAMINED: Mass.: Madaket, Nantucket, Aug. 17, 1942, E. V. Seeler
(FH).—N. J.: Newfield, 7 Aug. 1887, Ellis, type (CUP-D 8706, FH-E).
The species is known only from these two specimens. Though the
host of the type material had never been determined, it appears by com-
parison to be the same as that of the Nantucket specimen.
The affinities appear to be with Helotium populinum and H. tumidulum.
Fartow1a, Vou. 1, 1943
Ficures 10-17.
WHITE: Stupies IN GENUS HELOTIUo. III 169
It has not, however, had a combination in Helotium and there can scarcely
be any gain in making one for it, since in any disposition of these species
which even remotely approaches finality none of them could be left in
Helotium.
17. Helotium puberulum (Lasch ex Rabenh.) Fuckel, Fungi Rhenani 1150. 1864.
Peziza puberula Lasch ex Rabenh., Flora 34: 583. 1851; non Berk. & Curt.
Grevillea 3: 155. 1875. — Str., Nom. Fung., p. 432. 1862.
Pseudohelotium puberulum (Lasch ex Rabenh.) Fuckel, Symb. Myc., p. 298.
1869. — Sacc., Syll. Fung. 8: 300. 1889.
Lachnella puberula (Lasch ex Rabenh.) Phill. Grecillen 18: 85. 1890.
Pezizella puberula (Lasch ex Rabenh.) Rehm in Rabenh., Krypt.-Fl. 1°: 665.
1893. —[Rehm, Hedwigia 35: 146. 1eGeu 2 [Ranouse. Papers Michigan
Acad. 19 (1933) : 96. 1934.]
Dasyscypha puberula (Lasch ex Rabenh.) Massee, Brit. Fung.-Fl. 4: 355. 1895.
Belonium sulphureo-tinctum Rehm, Hedw. 35: 146. 1896. — Sacc., Syll. Fung.
14: 785. 1899.— Jaap, Verh. Bot. Ver. Prov. Brandenburg 52: 5. 1910;
57: 9. 1915.
Urceolella puberula (Lasch. ex Rahenh.) Boud., Hist. Classif. Discom. Eur.,
p. 129, 1907.
Niptera sulphureo-tincta (Rehm) Boud., Hist. Classif. Discom. Eur., p. 141.
1907.
Hymenoscypha puberula (Lasch ex Rabenh.) Migula in Thomé’s Krypt.-Fl.
10°: 1151. 1913.
Peziza punctiformis auct.— Von Hoéhn., Sitzb. Akad. Wissensch. Wien, Math.-
nat. KI], 1: 127: 601. 1918.
Calycellina populina auct.— Von Héhn., Sitzb. Akad. Wissensch. Wien. Math.-
nat. Kl. 1: 127. 601. 1918.— Von Héhn., Mitt. Bot. Inst. Techn. Hochsch.
Wien 3: 105. 1926.
Phialina puberula (Lasch. ex Rabenh.) Héhn. [Mitt. Bot. Inst. Techn. Hochsch.
3: 106. 1926]. — (Under Hyaloscy pha) Nannf., Nova Acta. Soc. Sci. Upsal.
4: 8: 273. 1932.
FIGURE 17.
Apothecia rather numerous on localized areas of the leaf surface,
sometimes crowded in groups of 5—15 with occasional solitary ones scat-
tered beyond these groups, very small, appearing sessile but actually
with a short obscure stipe, the disc 0.1-0.4 (—0.7) mm. diam., when
fresh bright lemon-yellow, not contracting or changing color aanaeael Nk
on drying, all parts concolorous; stipe very short, thin, delicate, not
perceptible unless fruit body is removed from substrate and inverted;
disc turbinate at first, finally spreading and thin, more so on drying;
receptacle very finely puberulent, often finely radiately striolate, lying
against the substratum in the older ones, sometimes becoming pale yellow
or buff on drying or after several years in the herbarium; margin
Fig. 10-17: 10, Helotium albopunctum, from type (CUP-D 5915); 11, H.
phyllogenon, from Rehm’s Ascom. 768, type (FH); 12, H. phyllophilum, from spec.
in Desm., Pl. Crypt. Fr. 1159, type (FH); 13, Phialea Cassandrae, from spec. cited
(MICH); 14, Helotium populinum. from Herb. Barb.-Boiss. 1217, type (FH); 15,
H. tumidulum. from spec. in Desm., Pl. Crypt. Fr. 2011, type (FH); 16, Pezizella
glagosa, from type (CUP-D 8706); 17, Helotium puberulum, from spec. Rabenh.,
Kl. Herb. Myc. 1529, type (FH). All drawings x 1290. |
170 FarLowl14, VoL. 1, 1943
not elevated, rarely becoming very slightly so on drying, circular to
irregular or angular in outline; hymenium convex in the fresh condition
and bright yellow, on drying becoming plane, waxy, remaining yellow,
or in old herbarium specimens changing to pale citron to pale orange;
paraphyses rather stout, scarcely or not at all enlarged at the apex,
3-3.5 », diam.; simple or more rarely once branched near the base; asei
originating from croziers, short, clavate, 38-54x 6.5-8 »; ascospores
biseriate or irregularly crowded, elongate, straight or slightly irregularly
curved, sometimes somewhat irregular in outline, 1l-celled, 12-18x
1.8-2.6 p», the content homogeneous or more frequently with several
small oil globules in each end.
HasiratT: On decaying leaves of Quercus sp., rarely on other genera;
in the Nantucket collection cited below a few apothecia are on a leaf
which appears to be Vaccinium sp., and Thiim., Myc. Austr. 930 is said,
evidently correctly, to be on Carpinus sp. Reports in European literature
of its occurrence on leaves other than Quercus probably are for the most
part incorrect.
DistriBuTION: New Hampshire, Massachusetts, New York, Germany,
Czechoslovakia, and France; reported from England and perhaps other
European countries but material not examined. Oct. 3—Nov. 11 in N. Am.
MATERIAL EXAMINED: North America: N. H.: Gilmanton, Oct. 12, 1942, D. H.
Linder (FH).— Mass.: Nantucket, Oct. 12, 1941, E. V. Seeler (FH).— East side
of Great Blue Hills, near Milton, Nov. 11, 1941, D. H. Linder & E. V. Seeler (FH).
— Another collection of same locality, etc. VW. L. White (FH).—N. Y.: Coy Glen,
near Ithaca, Oct. 3, 1938, Whetzel, White, et al (FH).— Europe: Germany: “Ad
folia decid. acervata Quercus pr. Driesen.” Rabenh., KI. Herb. Viv. Myc. 1529, type
of Peziza puberula (FH).—“An faulenden Eichenblittern, Zehlendorf bei Berlin.
10.1892.” Sydow. Rehm, Ascom. 1158, type of Belonium sulfureo-tinctum (FH).—
“An Blattern von Quercus Rober, Triglitz (Prignitz). Jaap.” Rehm. Ascom. 11588,
as B. sulfureo-tinctum (FH-H).— “An Quercus-Blattern, Berlin, Lechtindorf, 10.1892,
leg. P. Sydow.” Syd., Myc. March. 3778, as B. sulfureo-tinctum (FH).—“An foliis
Quercus coccinea, Tamsel bei Custria, Mark Brandenburg. 10.10.1906. leg. P. Vogel.”
As B. sulfureo-tinctum (FH-H).—“. . . auf Quercus Rober L. bei Konigstein.
Oct. 1910. leg. W. Krieger.” As Helotium punctiforme (Grev.) vy. H. non Rehm sub
Pezizella (FH-H).— Czechoslovakia: “Bohemia sept.: Teplitz ad folia putrida
Carpini, vere 1872 . . .” Thiim., Fungi Austr. 930, as Pseudohelotium ‘puberulum
(FH).— France: “Ad folia Quercus. 13 Oct. 1849. Sent by Roussel.” [as Peziza
punctiformis] (FH-C).
The following specimens under names listed in the synonymy above
represent incorrect determinations: Herb. Barb.-Bois 1220 [as Pseudo-
helotium puberulum] (FH) = Helotium populinum Fuckel. —Rehm.,
Ascom. 1162 (FH) [as Pezizella puberula] (FH) = Helotium populinum.
— Spec. taken on oak leaves at Ann Arbor, Michigan, Oct. 21, 1928, and
reported by Kanouse (1. c.) as Pezizella puberula (FH) = Mollisina
acerina (Mout.) Hohn.
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FARLOWIA
A JOURNAL OF CRYPTOGAMIC BOTANY
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DAVID H. LINDER, Editor
E. Y. SEELER, Jr., Mg. Ed.
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CONTENTS OF VOL. 1, Se 2
Burma Mossss. By Edwin B. Bartram woe i dtp We eas ee ee Satekee kee
AppDITIONS To THE Moss Fiona oF NorTHERN Carte. By Edwin B. Bartram. 191
Tur SmxvaL-DImMorPHISM OF THE TROPICAL Mossis oF THE Genus Macro-
mitrium. By Marthe Ernsi-Schwarzenbach.... 0... .....00000: be a ae ot 195
Tap Morrnoioey, Taxonomy, AND Disrrisution or Coccripromms Immitis
Rixrorp & Giicurist. By EH. HE. Baker, HE. M. Mrak, and C. E. Smith.... 199
An InpDEx TO RAFINESQUE’S PUBLISHED TECHNICAL Wane FOR THE CELLULAR
Cuveroganas~ By be Med 6655266 02 ees a es 245
Norrs on tae Synonymy or Some Norra Ammrican ‘THELEPHORACEAE AND
Oruer Resvurinates. By D. P. Rogers and A. S. Jackson..... 01.00 ecco 263
Vol. 1, No. 1 was issued on February 1, 1943.
FARLOWIA
A JOURNAL OF CRYPTOGAMIC BOTANY
A*Z0) Fe | JuLY, 1943 No. 2
BURMA MOSSES!
EpwIn B. BARTRAM
During recent years Mr. F. G. Dickason has been collecting mosses in
Burma in cooperation between the Arnold Arboretum of Harvard University
and Judson College, Rangoon, Burma. No outline of the Burma moss
flora has ever been published so these numerous collections supplemented
by a small series from the Burma-Tibet Frontier made by Mr. F. Kingdon
Ward with the C. Suydam Cutting Expedition of the Field Museum of
Natural History and a few earlier collections of Mr. Dickason received
from Mr. A. J. Sharp of the University of Tennessee form the basis for an
important and significant study.
The area is an important one in relation to the geographical distribu-
tion of Indo-Malayan mosses and in some respects seems to represent
a center in which a number of unusual types have originated. Cleistostoma
ambigua (Hook.) Brid. and Penzigiella cordata (Harv., Hook.) Fleisch.
are instances.
One of the especially interesting facts resulting from these studies is
the occurrence in Burma of Pseudosymblepharis pervaginata Broth.,
Pseudopohlia bulbifera Williams, Anomobryum gemmigerum Broth. and
Caly ptothecium himantocladioides Bartr. It is altogether probable that
these four supposedly endemic Philippine species had their origin in
Burma. Such instructive range extensions illustrate again the strong
west to east trend of distribution extending from the Himalayas through
Indo-Malaya, the Philippines and the Pacific Islands to Hawaii. The
collections recorded here total 133 species and are probably broadly
representative of the Burma moss flora as a whole. Of this total 74
species or well over half the number are known from the Philippines.
Types of the new species are deposited in the Farlow Herbarium of
Harvard University and in the herbarium of the author.
Unless otherwise indicated in the following list the collector is Mr.
F. G. Dickason.
* Contribution from the Laboratories of Cryptogamic coeey and the Farlow Her-
barium, Harvard University, no. 160.
171
172 FartowiA, VoL, 1, 1943
SPHAGNACEAE
Sphagnum subsecundum Nees.
Mt. Victoria, Chin Hills, wayside, 9000 ft., no. 8512.
Distribution: Europe, Asia, North America.
FISSIDENTACEAE
Fissidens sylvaticus Griff.
Taunggyi, F. S. S., damp open slope, no. 8787-C-1; Kamayut, Rangoon,
damp shady ground, 60 ft., no. 9600.
Distribution: Siam, Sumatra, Celebes, Java, Borneo, Hongkong, Philip-
pines.
Fissidens nobilis Griff.
Sin Lum Kaba, no. 117.
Distribution: Himalayas, Sumatra, Java, Hongkong, Philippines.
DITRICHACEAE
Garckea phascoides (Hook.) C. Miill.
Rangoon, along ditch during the rains, 20 ft., no. 6536; Tavoy, moun-
tain side, 20 ft., no. 7983; Rangoon, shady area, 60 ft., no. 9602, 9603,
9604; Rangoon, Kamayut, damp ground, 30-40 ft., no. 8663, 8669.
Distribution: wide in southern Asia, Malaysia, Philippine Islands.
DICRANACEAE
Trematodon longicollis Michx.
Pyin Mana, Kyauk-Than-Ba, Pyin Ma Na, wet shady place, 1000 ft.,
no. 7122, ;
Distribution: eastern United States, Mexico, Cuba, Italy, Ceylon,
Philippines,
Campylopus gracilis (Mitt.) Jaeg.
Loimwe, Kentung State, epiphytic, 6500 ft., no. 9741-3; Nataung,
Kareni State, on tree, 5000 ft., no. 9421.
Distribution: Sikkim.
Campylopus laetus (Mitt.) Jaeg.
Mount Popa, on rock, 3500 ft., no. 6615.
Distribution: Khasia.
Campylopus umbellatus (W. Arn.) Bartr.
Sin Lum Kaba, 6000 ft., no. 109.
Distribution: wide and frequent through the Pacific Islands and
Malaysia.
Dicranodontium uncinatum (Harv.) Jaeg.
Pang Yang, Manglon State, 5500 ft., no. 9689.2, 9694-4; Pang Yang,
BarRTRAM: BuRMA MossEs d Lies
Manglon State, epiphyte, 5000 ft., no. 9647; Saw Chaung, Chin Hills,
on tree, 6000 ft., no. 8556.
Distribution: India, Siam, Java, Moluccas, Philippines.
Holomitrium Griffithianum Mitt.
Pang Yang, Manglon State, 5500 ft., no. 9689.3.
Distribution: Bhotan, Assam, Khasia, Formosa.
Dicranum gymnostomum Mitt.
Burma-Tibet Frontier, Adung Valley, 12000 ft., Abies-Rhododendron
forest, F. Kingdon Ward, 1930-31.
Distribution: Himalayas, Yunnan.
Dicranoloma fragile (Hook.) Broth.
Loi Se, Manglon State, epiphyte, 4000 ft. no. 9650; Pang Yang,
Manglon State, moist banks, 5000 ft., no. 9657, 9675; Mong Pawk,
Kengtung State, epiphytic, 4000 ft., no. 9825.1.
Distribution: Himalayas, Annam, Philippines.
LEUCOBRYACEAE
Leucobryum Hollianum Doz. & Molk.
Zokhua (Haka), dry banks, 4800 ft., no. 7489.
Distribution: Java, Borneo.
Leucobryum scalare C. Miill.
Tavoy, 18 miles down, on trees, 1000 ft., no. 80/2 in part; Mong Pawk,
Kengtung State, on ground in woods, 4000 ft., no. 9824; Pang Yang,
Manglon State, epiphytic, 5000 ft., no. 9673, 9659-C; Nataung, Kareni
State, 9422-B.
Distribution: Nepal, Malacca, Siam, Java, Sumatra, Borneo, Philip-
pines, Fiji.
Leucobryum neilgherrense C. Miill.
Loimwe, Kentung State, epiphytic, 6500 ft., no. 9741.2; Sin Lum Kaba,
no. 108.
Distribution: India, Ceylon, Sumatra, Java, Celebes, Borneo, Philip-
pines, China, Japan, Korea.
Octoblepharum albidum Hedw.
Rangoon, on trees, 30 ft., no. 6817.
Distribution: cosmopolitan at low altitudes in tropical regions.
CALYMPERACEAE
Calymperes peguense Besch.
Rangoon, Royal Lake, on palm tree, 30 ft., no. 8665; Rangoon, Royal
Lake, on mango tree, 40 ft., no. 8668 in part.
Endemic.
&
174 Fartowia, VoL. 1, 1943
Calymperes Hampei Bryol. Jav.
Toungoo, no. 101.
Distribution: Malay Archipelago.
POTTIACEAE
Molendoa burmensis Bartr. sp. nov.
Gracilis, dense caespitosus, caespitibus fusco-viridis, haud nitidis. Caules ca. 2 cm.
longi, flexuosi, inferne parce radiculosi. Folia sicca arcte crispata, humida erecto-
patentia, 2-2.4 mm. longa, lineari-lanceolata, breviter acuminata, integerrima;
marginibus uno latere anguste recurvis; costa pervalida, inferne ad 80 y, lata, percur-
rente; cellulae superiores leniter papillosae, irregulariter subquadratae vel ovales,
parietibus pallidis, incrassatis, basilares rectangulares, pellucidae. Caetera ignota.
Goteik, Hsipaw, on rocks, 2800 ft., no. 9521 type, 9585,
This species differs from M. Roylei (Mitt.) Broth. in several noticeable
particulars. The leaves are strongly crisped when dry, the margin often
recurved on one side below the middle and the costa twice as wide toward
the base. Without fruit its position in Molendoa is problematical.
Anoectangium Stracheyanum Mitt.
Taunggyi, F.S.S., on damp area, 5200 ft., no. 8788, 8779, 8787-C-2.
Distribution: Himalayas.
Trichostomum angustatum (Mitt.) Fleisch.
Taunggyi, F.S.S., damp ground, 4500 ft., no. 9071-A.
Distribution: India, Ceylon, Java, Celebes, Philippines, South China.
Pseudosymblepharis pervaginata Broth.
Pang Wai, Kentung State, 6000 ft., no. 9699.1.
Distribution: Philippines.
Timmiella anomala (Bry. Eur.) Limp.
Taunggyi, F.S.S., on rock, 5000 ft., no. 9359 in part.
Distribution: Europe, North America, Himalayas.
Tortella cyrtobasis Dix.
Taunggyi, F.S.S., on rock, 5000 ft., no. 9359a.
Distribution: Siam.
Hyophila involuta (Hook.) Jaeg.
Taunggyi, F.S.S., on rocks, 5200 ft., no. 8790, 8794; Taunggyi, F.S.S.,
on trees, 4500 ft., no. 9075; Taunggyi, F.S.S., open damp ground, 4500 ft.,
no. 9094.1; Rangoon, Kamayut, 40 ft., no. 8670; Rangoon, Mingaladon,
on bricks, 60 ft., no. 9610.1; Rangoon, Inya Road, on rocks, 60 ft.,
no. 9598.
Distribution: India, East China, Philippines.
Barbula indica (Schwaegr.) Brid.
Rangoon, open damp earth, 40 ft., no. 8661.
Distribution: India, Ceylon, Java, Borneo, Philippines, East China.
BARTRAM: BuRMA MOosseEs 175
Barbula constricta Mitt.
Taunggyi, F.S.S., damp ground and open slopes, 4500-5200 ft., no.
8789, 9063, 9097.
Distribution: Himalayas, Philippines.
Barbula consanguinea (Thw. & Mitt.) Jaeg.
Rangoon, Mingaladon, on bricks, 60 ft., no. 9601; Taunggyi, E.S:S5
open damp ground, 4500-5200 ft., no. 8787-A, 9094, 9062, 9099.1.
Distribution: Ceylon, Singapore, Java, Philippines.
Desmatodon involutus Bartr. sp. nov.
Pusillus, dense caespitosus, flavescenti-viridis, haud nitidulus. Caulis brevis, dense
foliosus. Folia sicca leniter contorta, humida erecto-patentia, oblongo-ovata, concava,
apice truncata vel rotundata, haud mucronata, inferiora minuta, superiora, ca. 1 mm.
longa; marginibus integerrimis, superne late inflexis; costa 35-40 p lata, infra
apicem soluta; cellulae superiores rotundato-quadratae, mamillosae, dorso laeves,
diam. 7-8 pu, basilares rectangulares, laevissimae, pellucidae. Folia perichaetialia
caulinis similia, seta 6-7 mm. longa, aureo-rubella, laevissima ; theca erecta, oblongo-
ovalis, deoperculata 1-1.2 mm. longa; peristomii dentes 16, erecti, pallidi, papillosi,
ad 150 p longi; operculum rubrum, conico-rostratum, ca. 0.5 mm. longum; annulus
latus, calyptra cucullata, spori laeves, diam. 7-8 yp.
Taunggyi, F.S.S., damp open slope, 5200 ft., no. 8787-B, 8795 type.
In Desmatodon this species is unique in the broadly rounded or truncate
leaves with the margins widely inflexed above. The costa in cross section
shows the typical structure with a stereid band on the dorsal side only;
the leaf cells are smooth on the back and mamillose on the ventral side
and there is nothing unusual in the peristome characters. On the whole
it seems more appropriate to include the species in Desmatodon than to
create a new ‘monotypic genus.
FUNARIACEAE
Entosthodon wichurae (Broth.) Fleisch.
Taunggyi, F.S.S., damp ground, 4500 ft., no. 9072, 9078, 9500.
Nataung, Kareni State, open dry ground, 5000 ft., no. 9427.
Distribution: Java.
Funaria hygrometrica Hedw. var. ealvescens (Schwaegr.) Bry. Eur.
Mt. Nwalabo, Tavoy, on tree, 5000 ft., no. 8205; Nataung, Kareni
State, open damp earth, 5000 ft., no. 9487; Wunba Taung, Chin Hills,
on rocks, 7900 ft., no. 8481, 8491.
Distribution: cosmopolitan in warm regions.
SPLACHNACEAE
Tayloria indica Mitt. :
Haka, on trees and rocks, 5100-7000 ft., no. 7401, 7583.
Distribution: Himalayas, Philippines.
176 FarLowiA, Vou. 1, 1943
BRYACEAE
Pseudopohlia bulbifera R. S. Williams
Taungteik (Falam), damp banks, 4500 ft., no. 7548; Yetagone, along
the damp, shady path, 4000 ft., no. 7801a. .
Distribution: Philippines.
Brachymenium acuminatum Harv.
Taunggyi, F.S.S., damp ground, 4500 ft., no. 9071.B, 9083.1,
Distribution: East India, Philippines.
Brachymenium nepalense Hook.
Taunggyi, F.S.S., no. 7891; Nataung, Kareni State, on trees, 5000 ft.,
no. 9426-A ; Pang Yang, Manglon State, on bank and epiphytic, 5000-5500
ft., no. 9686, 9689, 9693.1, 9815.2; 9814-A; Hopong, Hopong State,
no. 82766,
Distribution: East India, Siam, Sumatra, Java, Borneo, Philippines,
Celebes.
Anomobryum gemmigerum Broth.
Taunggyi, F.S.S., open damp area, 4500 ft., no. 9083.
Distribution: Philippines.
Anomobryum cymbifolium (C. Miill.) Broth.
Taunggyi, F.S.S., damp ground, 4500 ft., no. 9066.
Distribution: Himalayas, South India, Khasia, Java, Philippines,
Amboina.
Bryum argenteum Hedw. var. lanatum Bry. Eur.
Taunggyi, F.S.S., on damp ground, 4500 ft., no. 9079, 9089,
Distribution: cosmopolitan.
Bryum nitens Hook.
Taunggyi, F.S.S., on trees, 4500 ft., no. 9095.1.
Distribution: Nepal, Ceylon, Java.
Bryum coronatum Schwaegr.
Taunggyi, F.S.S., open damp slope, 4500 ft., no. 9088, 9099; Rangoon,
Inya Road, on bricks, 60 ft., no. 9599.
Distribution: cosmopolitan in tropical and subtropical regions.
Bryum porphyroneuron C. Miill. :
Taunggyi, F.S.S., damp ground, 4500 ft., no. 9503-A.
Distribution: India, Ceylon, Sumatra, Java.
Bryum ramosum (Hook.) Mitt.
Thandaung, usually found on rocks of cold places, 3500 ft., no. 6738.
Loimwe, Kentung State, epiphytic, 6500 ft., no. 9741.1.
BaRTRAM: BurMa MossEs 177
Rhodobryum giganteum (Hook.) Schp.
Mong Yang, Kengtung State, damp soil in forest, 3500 ft., no. 9862;
Mong Yang, Kengtung State, epiphyte, 3500 ft., no. 9863.4.1; Webula,
along damp trails, no. 7786; Taungteik (Falam), damp shady banks,
4000 ft., no. 7756; Taunggyi, South Shan States, along the path, 5300 ft.,
no. 8398; Nataung, Kareni State, on damp areas, 5000 ft., no. 9435;
Burma-Tibet Frontier, F. Kingdon Ward, Coll.
Distribution: Himalayas, Sumatra, Java, Borneo, Philippines, China,
Japan, Hawaii.
MNIACEAE
Orthomnium trichomitrium Wils.
Nataung, Kareni State, shady forest, 6300 ft., on trees, no. 9448.1.
Distribution: Sikkim, Khasia.
Mnium Thomsoni Schp.
Burma-Tibet Frontier, F. Kingdon Ward, Coll. 1930-31.
Distribution: Sikkim, Yunnan, Japan.
Mnium rostratum Schrad.
Webula, rocky wet places, 4000 ft., no. 7788; Pang Wai, Kengtung
State, 6000 ft., no. 9690, 9699.5; Pang Yang, Manglon State, epiphytic,
5000 ft., no. 9656; Loimwe, Kantung State, epiphytic, 6500 ft., no. 9741.
Distribution: cosmopolitan.
Mnium coriaceum Griff.
Haka, damp banks, 6200 ft., no. 7294.
Distribution: Himalayas, Khasia.
BARTRAMIACEAE
Bartramia Halleriana Hedw.
Wunba Taung, Chin Hill, no. 84826.
Distribution: Europe, North America, South America, Himalayas,
Yunnan, New Guinea, New Zealand.
Philonotis Griffithiana (Wils.) Mitt.
Nataung, Kareni State, on open damp earth, 5000 ft., no. 9485.
Distribution: Himalayas, Khasia.
Philonotis (Philonotula) aristifolia Bartr. sp. nov.
Dioica videtur. Densissime caespitosa, caespites superne laete viridi, intus fusci.
Caules ad 4.cm. alti. Folia late patentia, saepe falcata, angustissime lineari-lanceolata,
ca. 2 mm. longa, 0.3 mm. lata, sensim acuminata, longe aristata; margines leniter
recurvi, denticulati; cellulae superiores lineares, papillosae, inferne sensim latiores;
costa valida, in aristam denticulatam Jonge excurrens. Seta 3 cm. alta; theca
magna, inclinata vel horizontalis, fortiter plicata; peristomium duplex.
178 FarLowia, Vou. 1, 1943
Langoon, Insein, on damp earth, no. 8158; Mt. Nwalabo, Tavoy, on
damp earth, 5000 ft., no. 8203 type.
Differs radically from P. angusta Mitt. of Ceylon in the longer, narrower
more widely spreading aristate leaves with longer cells. The type collec-
tion is richly fruited and is an unusually handsome little plant for the
genus with a delicate silky appearance due to the slender spreading foliage.
Philonotis imbricatula Mitt.
Taunggyi, F.S.S., open damp area, 4500 ft., no. 9494.
Distribution: Ceylon, Hongkong, Philippines, Fiji.
Philonotis perlaxifolia Dix. ?
Rangoon, Royal Lake, on mango tree, 30 ft., no. 8666.
Distribution: Siam.
As this collection is sterile the determination is open to doubt. |
Philonotis faleata (Hook.) Mitt.
Taungteik (Falam), wet bank, 4500 ft. no. 7237, Falam, damp bank,
6000 ft., no. 7324.
Distribution: North India, China, Japan, Philippines, Hawaii.
Philonotis Turneriana (Schwaegr.) Mitt.
Taungteik, damp banks, 4500 ft., no. 7237.A; Naungkhio, Thibaw State,
on rocks in moist localities, 2800 ft., no. 9536.
Distribution: Himalayas, Khasia, J ava, Japan, China, Hawaii.
Philonotis speciosa (Griff.) Mitt.
Yetagone, along the damp shady path, 4000 ft., no. 7801.
Distribution: Nepal, Khasia, Philippines.
Fleischerobryum longicollis (Hampe) Loeske
Burma-Tibet frontier, F. Kingdon Ward, Coll.
Distribution: Himalayas, Java.
ORTHOTRICHACEAE
Macromitrium nepalense (Hook. & Grev.) Schwaegr.
Pang Yang, Manglon State, epiphyte, 5000 ft., no. 9816; Mong Pawk,
Kentung State, epiphytic in woods, 4000 ft., no. 9823.
Distribution: Nepal, Sikkim, Siam, Malay Peninsula, China, Philippines.
Macromitrium Dickasonii Bartr. sp. nov.
Caulis repens, ramis erectis, ca. 5 mm. altis, fusco-viridibus. Folia sicca arcte
crispata, humida erecto-patentia, apice incurva, 2.5 mm. longa, oblongo-lanceolata,
acuta, integerrima, profunde canaliculata; costa infra apicem evanida; cellulae
superiores minutae, obscurae, papillosae, basilares magis incrassatae, laevissimae,
marginales angustissime lineares, in costam versus breviores. Seta crassa, rubra,
laevissima, 6-7 mm. longa; theca oblongo-ovalis, laevis, macrostoma, castanea,
‘
BarRTRAM: BurMA Mosses 179
2-2.5 mm. longa; peristomii simplicis, dentes ad 165 u, longi, obtusi, sordide albidi,
papillosi; calyptra dense pilosa; spori minute papillosi, diam. ad 30 u.
Mogok, Apr. 1934, no. 102, type.
Near M. falcatulum C. Mill. of the Philippines but with elongated basal
cells, longer setae and longer capsules. ;
Macromitrium suleatum (Hook.) Brid.
Nataung, Kareni State, on trees, 5000 ft., no. 9431; Nataung, Kareni
State, on trees, shady forest, 6300 ft., no. 9447-B, 9448; Pang Yang,
Manglon State, moist bank, 5000 ft., no. 9654; Pang Yang, Manglon
State, 6000 ft., no. 9694-C; Haka, damp banks, 6200 ft., no. 7584.
Distribution: East India, Ceylon, Malay Peninsula, Borneo, Philippines.
Macromitrium goniorhynchum (Doz. & Molk.) Jaeg.
Kalaw, 5000 ft., no. 115.
Distribution: Khasia through Malaysia to the Philippines and New
Guinea.
RHACOPILACEAE
Rhacopilum Schmidii (C. Miill.) Jaeg.
Hopong, Hopong State, on trees, no. 8276; Taunggyi, F.S.S., on trees,
5000 ft., no. 8775; Mong Yang, Kengtung State, epiphyte, 3500 ft.,
no. 9876. ;
Distribution: Nilghiri, Tonkin, Philippines.
HEDWIGIACEAE
Cleistostoma ambigua (Hook.) Brid.
Sin Lum Kaha, 6000 ft., no. 124; Taunggyi, F.S.S., on rocks, 6000 ft.,
no. 7848; Nataung, Kareni State, on trees, shady slope, 5000 ft., no.
9425; Pang Yang, Manglon State, on trees, 5500 ft., no. 9642-A.
Distribution: Himalayas, West China, Siam, Assam.
TRACHYPODACEAE
Diaphanodon thuidioides Ren. & Card.
Mt. Victoria, Chin Hills, on trees, 10000 ft., no. 8523a.
Distribution: Sikkim.
Trachypus bicolor Reinw. & Hornsch.
Pang Yang, Manglon State, damp soil, 5000 ft., no. 9659-A; Pang Yang,
Manglon State, epiphytic, 5000 ft., no. 9815; Pang Yang, Manglon State,
6000 ft. no. 9713; Pang Wai, Kentung State, 6000 ft., no. 9699.4;
Nataung, Kareni State, on trees, 5000 ft., no. 9431.1; Nataung, Kareni
State, shady mountain slope on trees, 5000 ft., no. 9424; Mt. Victoria,
Chin Hills, on trees, 10000 ft., no. 8529.
Distribution: North India, Ceylon,:Siam, Java, Celebes, Philippines,
China, Japan, Formosa, Hawaii.
180 FarLtowia, VoL. 1, 1943
Trachypodopsis crispatula (Hook.) Fleisch.
Haka, on trees and banks, 6200-6500 ft., no. 7293, 7427; Taungteik
(Falam), mountain side, 5000 ft., no. 7262; Mt. Nwalabo, Tavoy, on
rocks, 5000 ft., no. 8196; Mt. Victoria, Chin Hills, on trees, 10000 ft.,
no. 8523; Loi, Maing, Sampu, Namkhok State, epiphytic in mossy forest,
7500 ft., no. 8356, a robust form with coarsely toothed leaves; Pang
Wai, Kentung State, 6000 ft., no. 9699.2; Pang Yang, Manglon State,
5500 ft., no. 9690.B.1; Namkham, no. 103.
Distribution: Himalayas, Yunnan, Ceylon, Andaman Islands, Philip-
pines.
MYURIACEAE
Myurium rufescens (Reinw. & Hornsch.) Fleisch,
Pang Yang, Manglon State, 6000 ft., no. 9714; Pang Yang, Manglon
State, epiphytic, 5000 ft., no. 9815.1; Nataung, Kareni State, on trees,
shady forest, 6300 ft., no. 9447-C.
Distribution: Khasia, Ceylon, Siam, wide in Malaysia, Philippines,
Australia, New Caledonia.
PTEROBRYACEAE
Penzigiella cordata (Harv.) Fleisch.
Haka, mossy forest, 7625 ft., no. 7500a; Pang Yang, Manglon State,
epiphyte, 5000 ft., no. 9818.
Distribution: Sikkim, Nepal, Khasia.
Endotrichella rigida Dix.
Nataung, Kareni State, on trees, shady forest, 6300 ft., no. 9446.
Distribution: Siam.
Pterobryopsis (Eupterobryopsis) caudata Bartr. sp. nov.
Dioica?, robusta, pallide virens, nitida. Caules secundarii ad 7 cm. alti, breviter
stipitati, simplices, superne longe attenuati, arcuati. Folia humida late patentia,
cochleariformia, ad 3 mm. longa, 2 mm. lata, e basi cordata late ovata, acuta; mar-
ginibus erectis, integris vel apice minutissime denticulatis; costa supra medium
folii evanida; cellulae superiores lineari-rhomboideae, infimae laxiores, aurantiacae,
alares nullae. Fructus ignotus.
Mt. Nwalabo, Tavoy, on trees, 5000 ft., no. 8202 type; Pang Yang,
Manglon State, 5500 ft., no. 9693; Nataung, Kareni State, on trees, shady
forest, 6300 ft., no. 9448.2.
A very unusual looking plant with the stems attenuate in a long, curved,
microphyllous, proboscoid-like tip. Through the well defined costa it is
probably nearest P. crassicaulis (C. Miill.) Fleisch. but the broad, short
pointed leaves not inflexed on the margins above and without any well
defined alar cells are thoroughly distinctive. I am not sure that nos.
9448.2 and 9693 belong here. In 9693 the stems are not caudate at the
tips and in 9448.2 the habit is more slender. Both show the leaves more
BaRTRAM: Burma MosseEs 181
slender than in the type collection and the upper margins are distinctly
serrulate. Otherwise the form and structure of the leaves are very
similar.
Pterobryopsis flexipes (Mitt.) Fleisch.
Nataung, Kareni State, on trees, shady forest, 6300 ft., no. 9446.1.
Distribution: Nilghiri, Ceylon.
Pterobryopsis acuminata (Harv.) Fleisch.
Sin Lum Kaba, no. 119,
Distribution: Sikkim, Nepal, Khasia, Nilghiri.
Pterobryopsis orientalis (C. Miill.) Fleisch.
Taunggyi, F.S.S., on rocks, 4700-5000 ft., no. 7872, 9021; Haka, on
trees, 6400 ft., no. 7443; Pang Yang, Manglon State, 5500 ft., no. 9685,
9695-A; Mt. Popa, 4000 ft., no. 125.
Distribution: Himalayas, Nilghiri, Assam.
METEORIACEAE
Papillaria fuscescens (Hook.) Jaeg.
Haka, on trees, 6200-7625 ft., no. 7296, 7426, 7499a, 7503; Thandaung,
Taungoo, on rocks, 4000 ft., no. 8180; Nawrig Heng, Yunnan, China,
no. 123; Nataung, Kareni State, on trees, shady forest, 5800-6300 ft.,
no. 9436, 9447-A, -
Distribution: Himalayas, Ceylon, Yunnan, Siam, Malaysia, Philippines.
Papillaria (Eupapillaria) lanosa Bartr. sp. nov.
Tenella, pallide lutescens vel subfusca, haud nitida. Caulis secundarius remote
et irregulariter ramosus, ad 5 cm. longus, flexuosus. Folia sicca laxe adpressa,
humida erecto-patentia, leniter plicatula, oblongo-ovata, plerumque raptim longissime
piliformiter attenuata, 2.5 mm. longa, 0.8 mm. lata, canaliculato-concava; margines
inferne late inflexi, superne erecti vel anguste incurvi, ubique remote et minute
denticulati; cellulae lineares, obscurae, dense et humiliter papillosae, infimae solum
laxiores, laevissimae, alares subquadratae, minutae, numerosae. Fructus unicus
visus; seta brevis, 1 mm. longa, crassa; theca exserta, erecta, ovato-globosa, 1 mm.
longa; operculum conico-rostratum, 1 mm. longum.
Pang Yang, Manglon State, epiphyte, 5500 ft., no. 9820, type.
In a general way this species is suggestive of P. Deppei (Hornsch.)
Jaeg. of tropical America but the distinctions are obvious. Here the
leaves are more abruptly narrowed to the capillary hair point and the
leaf cells much more obscure with numerous small papillae. In P. Deppei
the papillae are sharp and salient and average 3 or 4 to a cell while in
P. lanosa the papillae are low and so numerous as to almost obscure the
cell structure.
Aerobryopsis longissima (Doz. & Molk.) Fleisch.
Mong Yang, Kengtung State, epiphyte, 3500 ft., no. 9875, 9877.
Distribution: India, Ceylon, Malaysia, China, Philippines to Hawaii.
182 FarLowia, VoL. 1, 1943
Aerobryidium aureo-nitens (Hook.) Broth.
Pang Wai, Kentung State, 6000 ft., no. 9699.3; Taunggyi, F.5.S., on
rocks and trees, 4500-5000 ft., no. 9355, 9360, 9507-A; Pang Yang,
Manglon State, 5500 ft., no. 9688; Mt. Victoria, Chin Hills, on trees,
10000 ft., no. 8586; Taungteik (Falam), mountain path side, 5000 ft.,
no. 7257; Namkham, 4000 ft., no. 173; Sin Lum Kaba, no. 104; Mt. Popa,
4000 ft., no. 178.
Distribution: Himalayas, Khasia, Nilghiri.
Floribundaria floribunda (Doz. & Molk.) Fleisch.
Haka, mossy forest, epiphytic, 7625 ft., no. 7495; Pang Yang, Manglon
State, 5500 ft., no. 9694-B; N’Bapa, no. 111; Sin Lum Kaba, 6000 ft.,
no. 122,
Distribution: Africa, India, Malaysia, Japan, Tonkin, Philippines to
Hawaii.
Floribundaria aurea (Griff.) Broth.
Burma-Tibet Frontier, Valley of Tamai River, 3000 ft., F. Kingdon
Ward, Jan. 1931; Burma-Tibet frontier, Adung Valley, 12000 ft., upper-
most Abies-Rhodendron forest, F. Kingdon Ward, 1930-31.
Distribution: Himalayas, Bhotan, Java, Celebes, Ceram, Japan.
Barbella enervis (Mitt.) Fleisch.
Mong Yang, Kengtung State, epiphyte, 3500 ft., no. 9874, 9869,
9863.B.1; Mong Yang, Kengtung State, damp soil, 3500 ft., no. 9867.
Distribution: Himalayas, Ceylon, Philippines, Australia, Lord Howe
Island, New Caledonia.
Barbella Stevensii (Ren. & Card.) Fleisch.
Sin Lum Kaba, no. 112.
Distribution: Sikkim.
Barbella pendula (Sull.) Fleisch.
Mong Yang, Kengtung State, epiphytic, 3500 ft., no. 9860.
Distribution: Ceylon, Sumatra, Java, China, Japan, Formosa, Philip-
pines, Southern United States.
Barbella (Aerobryella) burmensis Bartr. sp. nov.
Tenella, lutescens, nitida. Caules secundarii elongati, tenues, flexuosi, remote
ramosi, ramis brevibus, vix 1 cm. longis, complanatis, obtusis. Folia caulina ovato-
lanceolata, acuminata, 1.3 mm. longa; margines plani, ubique denticulati; costa
valida, supra medium folii evanida; cellulae lineares, unipapillati; folia ramea sub-
similia, breviter acuminata, ad 1.8 mm. longa, 0.6 mm. lata, margines fortiter denticu-
lati. Caetera ignota.
Haka, mossy forest, 7625 ft., no. 7500d, type.
The branch leaves in this species are broader and shorter than in B.
javanica (Bryol. Jav.) Broth. and smaller and more crowded than in
B. spiculata (Mitt.) Broth.
BartrRaAM: Burma Mosses 183
Meteoriopsis ancistrodes (Ren. & Card.) Broth.
Mt. Popa, on trees, 4000 ft., no. 6616.
Distribution: Himalayas, Formosa.
Meteoriopsis squarrosa (Hook.) Fleisch.
Pang Yang, Manglon State, epiphyte, 5000 ft., no. 9818.1; Namkham,
MOL ds
Distribution: Himalayas, Ceylon, Malay Peninsula, Siam, Sumatra,
Java, Philippines.
Meteoriopsis reclinata (Mitt.) Fleisch.
Taunggyi, F.S.S., on trees, 4500 ft., no. 9499; Pang Yang, Manglon
State, epiphyte, 5000 ft., no. 9661, 9664; Mong Yang, Kengtung State,
epiphyte, 3500 ft., no. 9861-B, 9873.
Distribution: Nilghiri, Ceylon, Java, Celebes, Philippines, Formosa.
NECKERACEAE
Calyptothectum burmense Bartr. sp. nov.
Caulis secundarius ad 7 cm. longus, cum foliis 5-6 mm. latus, parcissime ramosus,
valde complanatus, luteo-viridis, nitidus. Folia fortiter complanata, horride patula,
haud undulata, vix auriculata, concava, oblongo-ovata, breviter acuminata, ca. 3.5 mm.
longa, 1.5 mm. lata, inferiora minora; margines erecti, superne denticulati; costa
supra medium folii evanida; cellulae anguste rhomboideae, 7-8 uy. latae, ca. 1 : 7,
parietibus firmis, pellucidis, infimae laxiores, fuscae. Caetera ignota.
Pang Yang, Manglon State, epiphyte, 5000 ft., no. 9803-B type, 9819.
In this species the stems are more strongly flattened and less branched
than in C. Ramos Broth. of the Philippines, the leaves are broader and
the lamina cells wider. No. 9819 shows the leaves more or less auriculate
and has scattered septate propagula in the upper leaf axils but otherwise
agrees with the type collection.
Calyptothecium himantocladioides Bartr.
Mt. Nwalabo, Tavoy, on rocks, 5000 ft., no. 8795.
Distribution: Philippines.
Calyptothecium Urvilleanum (C. M.) Broth.
Haka, on trees, 7400 ft., no. 7448; Burma-Tibet Frontier, Valley of
Tamai River, 3000 ft., F. Kingdon Ward, Coll., Jan. 1931.
Distribution: India, Malaysia, to Fiji, Caroline Islands, Samoa and
Tahiti.
Neckera crenulata Harv.
Taungteik (Falam), open mountain side, 5000 ft., no. 7265; Pang
Yang, Manglon State, 5500 ft., no. 9690-B.
Distribution: Himalayas, Khasia.
Neckera himalayana Mitt.
Haka, mossy forest, epiphyte, 7625 ft., no. 7502.
Distribution: Sikkim.
184 FarLowla, VoL. 1, 1943
Homaliodendron exiguum (Bryol. Jav.) Fleisch.
Pang Yang, Manglon State, 5500 ft., no. 9687.
Distribution: Himalayas, Ceylon, Malay Peninsula, Sumatra, Java,
Celebes, Philippines, Australia, New Guinea.
Homaliodendron microdendron (Mont.) Fleisch.
Haka, mossy forest, 7625 ft., no. 7500c.
Distribution: Sikkim, Khasia, Bhotan, Yunnan, Borneo, Philippines,
Formosa.
Homaliodendron flabellatum (Sm.) Fleisch.
Loi Naing, Sampu, Namkhok State, epiphytic in mossy forest, 7500 ft.,
no. 8355; Sin Lum Kaba, no. 110. Haka, mossy forest, 7625 ft., no.
7500.
Distribution: India and Ceylon to Malaysia and Japan and through the
Pacific Islands to Hawaii.
Thamnium subseriatum (Hook.) Mitt.
Haka, mossy forest, 7625 ft., no. 7499, 7500.
Distribution: Himalayas, Ceylon, Philippines.
THUIDIACEAE
Claopodium nervosum (Harv.) Fleisch.
Mt. Popa, damp places, 3000 ft., no. 6632; Taunggyi, F.S.S., on damp
ground, 5000 ft., no. 8781.1.
Distribution: Himalayas, Ceylon, Sumatra, Java, Celebes, Philippines,
China, Korea, Japan.
Thuidium investe (Mitt.) Jaeg.
Ah-Shai-Myin, Ah-Naught-Myin, Lawk-Sawk State, no. 8701-A.
Distribution: Philippines.
Thuidium tamariscellum (C. Miill.) Bryol. Jav.
Taunggyi, F.S.S., on damp ground, 4500 ft., no. 9070, 9085.
Distribution: Nilghiri, Tonkin, Siam, Java, Philippines.
Thuidium Meyenianum (Hampe) Bryol. Jav.
Gotiek, on rocks, 2800 ft,, no. 9524; Naungkhio, Hsipau, on rocks,
2800 ft., no. 9535; Pang Yang, Manglon State, epiphyte, 5000 ft., no.
9674.
Distribution: Himalayas, Ceylon, Annam, Philippines, Sumatra, Java,
New Guinea, Solomon Islands.
Thuidium glaucinoides Broth.
Mong Yang, Kengtung State, damp bank, 3500 ft., no. 9865.
Distribution: Tonkin, Malaysia, China, Formosa, Philippines, New
Hebrides, Fiji, Samoa.
BarTRAM: BurRMA MosseEs 185
Thuidium cymbifolium (Doz. & Molk.) Bryol. Jav.
Pang Yang, Manglon State, epiphyte, 5000 ft., no. 9803-A; Taungteik
(Falam), open mountain side, 5000 ft., no. 7263; Haka, mossy forest,
7625 ft., no. 7498; Mt. Popa, epiphyte and on banks, 3500 ft., no. 6630;
Pang Wai, Kengtung State, 6000 ft., no. 9699; Sin Lum Kaba, no. 116.
Burma-Tibet Frontier, F. Kingdon Ward, 1930-31.
Distribution: wide in Malaysia, Formosa, Philippines, Lord Howe
Island, New Guinea, China, Japan.
Actinothuidium Hookeri (Mitt.) Broth.
Mt. Victoria, 10000 ft., on tree, no. 8531.
BRACHYTHECIACEAE
Brachythecium Buchanani (Hook.) Par.
Taunggyi, F.S.S., on trees, rocks, and ground, 4500-5000 ft., no. 8773.1,
8781, 9081, 9090; Pang Wai, Kengtung State, 6000 ft., no. 9695-B;
Nataung, Kareni State, no. 9426-B; Mt. Popa, Myin Gyan, on rocks,
3000 ft., no. 6632; Sin Khoa (Haka), on tree trunks, 4000 ft., no. 7617;
Sin Lum Kaba, no. 120.
Distribution: Himalayas, Assam, Philippines, China, Japan.
The above collections are all sterile but probably belong here.
Brachythecium longicuspidatum (Mitt.) Jaeg.
Zokhua (Haka), 4800 ft., no. 7490a.
Distribution: Sikkim.
ENTODONTACEAE
Erythrodontium julaceum (Hook.) Par.
Taunggyi, F.S.S., on rocks, trees and banks, 4500-5200 ft., no. 8792,
9061, 9064, 9068, 9084, 9091, 9092, 9090.1, 9507-B; Namkok, on tree,
3000 ft., no. 9400.
Distribution: India, Assam, Tonkin, Yunnan, Ceylon, Sumatra, Java,
Celebes, Philippines.
Trachyphyllum inflexum (Harv.) Gepp
Naunghkio, Hsipaw State, on tree trunk, 2800 ft., no. 9574 in part.
Distribution: India, Philippines, New Caledonia.
Campylodontium flavescens (Hook.) Bryol. Jav.
Pang Yang, Manglon State, epiphyte, 5000 ft., no. 9817.
Distribution: India, Malay Peninsula, Ceylon, Annam, Sumatra, Java,
Philippines, Celebes, Australia.
Rozea pterogonioides (Hook.) Jaeg.
Mt. Victoria, Chin Hills, on trees, 10000 ft., no. 8540.
Himalayas, Yunnan.
186 Fartowia, VoL. 1, 1943
Entodon flavescens (Schwaegr.) Jaeg.
Taunggyi, F.S.S., on trees, 4500 ft., no. 9098.
Distribution: Nepal.
Much more slender than the plant illustrated in Schwaegrichen’s plate
but apparently not specifically distinct.
Entodon plicatus C. Miill.
Pang Yang, Manglon State, epiphyte, 4500-6000 ft., no. 9659-B, 9665,
9672, 9712; Mong Yang, Kengtung State, epiphyte and on soil, 3500 ft.,
9861-a, 9871-A, 9871-B, 9872.1; Taunggyi, F.S.S., on trees, 4500 ft.,
no. 8780-A, 9095; Hopong, Hopong State, no. 8276a; Naungkhio, Thibaw,
on rocks, 2800 ft., no. 9540.1; Zokhua (Haka), 4800 ft., dry banks, no.
7490.
Distribution: Himalayas, Ceylon, Celebes, Philippines.
PLAGIOTHECIACEAE
Stereophyllum tavoyense (Hook.) Jaeg.
Mt. Popa, on rocks, 3000 ft., no. 6654 in part; Mt. Popa, 4000 ft., no. 106.
Distribution: India.
SEMATOPHYLLACEAE
Clastobryum Dickasonii Bartr. sp. nov.
Dioicum? gracile, lutescenti-viride, nitidum. Caulis repens, irregulariter sub-
pinnatus, ramis brevibus, ad 6-7 mm. longis, caudato-attenuatis. Folia erecto-
patentia, oblongo-lanceolata, 1.2 mm. longa, 0.4 mm. lata, acuminata, integerrima,
fortiter concava, ecostata; cellulae lineares, haud incrassatae, ca. 1 : 8, laevissimae,
infimae fuscae, alares 3-4, magnae, oblongae. Fructus ignotus.
Saw Chaung, Chin Hills, on tree, 6000 ft., no. 8556a, type.
In this species the leaves are relatively broader and shorter acuminate
than in C. caudatum (Bryol. Jav.) Fleisch. of Java, the cells are shorter
and not incrassate and the reduced leaves of the caudate tips are short
acuminate. C. caudiforme Dix. of India is described as having strongly
denticulate leaves and very narrow cells.
Acanthocladium baculiferum Dix.
Nataung, Kareni State, open damp ground, 5000 ft., no. 9422-4;
Nataung, Koreni State, on trees, shady slope, 5000 ft., no. 9428; Nataung,
Kareni State, on trees, shady forest, 5300 ft., no. 9445; Pang Yang,
Manglon State, on a rock, 5500 ft., no. 9813-B, 9814-B; Mong Pawk,
Kengtung State, epiphytic and on ground in woods, 4000 ft., no. 9824.1,
9825.
Distribution: Assam.
As no. 9428 and 9445 in the above series are in fruit it is possible to
add to the original description of this interesting species, previously
known only from Assam, the following sporophyte characters. Peri-
-chaetial leaves erect, not strongly convolute, the inner lightly plicate,
BARTRAM: BuRMA MosseEs 187
oblong-ovate, rather abruptly acuminate, denticulate toward the extreme
apex. Seta slender, smooth, reddish, strongly flexuose, to 3.5 cm. long.
Capsule nodding, strongly asymmetrical, constricted under the oblique
, mouth when dry, urn about 2 mm. long, brown, gibbous on the upper
side, abruptly constricted to a short neck. Peristome normal. Lid and
calyptra not seen.
Acanthocladium benguetense Broth. var. latifolium Bartr. var. nov.
Ramis vix attenuatis. Folia caulina latiora, ad 0.75 mm. lata, abrupte acuminata,
subintegra.
Nataung, Kareni State, on tree, shady forest, 5300 ft., no. 9444, type.
Although the differences between the Burma plants and those of the
Philippines are quite marked the distinctions seem hardly wide enough
to merit a new species. The Burma plant is more robust than the type
from the Philippines and the branchlets are scarcely attenuate; the stem
leaves are more broadly ovate, more abruptly acuminate and subentire.
Brotherella erythrocaulis (Mitt.) Fleisch.
Burma-Tibet Frontier, Adung Valley, uppermost Abies-Rhododendron
forest, F. Kingdon Ward, 1930-31.
Distribution: Sikkim, Bhotan, Yunnan.
Sematophyllum tristichulum (Mitt.) Fleisch.
Pang Yang, Manglon State, rock, 5000 ft., no. 9822.
Distribution: Khasia, Assam, Ceylon, Siam, Java, Annam, Philippines.
Sematophyllum subhumile (C. Miill.) Fleisch,
Pang Yang, Manglon State, damp soil, 5000 ft., no. 9659-A.1.
Distribution: India.
Taxithelium nepalense (Schwaegr.) Broth.
Rangoon, Royal Lake, on damp earth, 40 ft., no. 8667.
Distribution: India, Ceylon, Sumatra, Java, Assam, Borneo, Philip-
pines, Amboina.
HYPNACEAE
Hypnum plumaeforme Wils.
Mong Yang, Kengtung State, epiphyte and on damp soil, 3500 ft.,
no. 9863-A, 9863-B, 9866, 9870; Pang Yang, Manglon State, epiphyte,
5000 ft., no. 9645.
Distribution: Korea, Japan, China, Formosa, Tonkin.
Ectropothecium dealbatum (Hornsch. & Reinw.) Jaeg.
Taunggyi, F.S.S., on tree, 5000 ft., no. 8780-A-1.
Distribution: Sumatra, Java, Borneo, Philippines.
188 FarRLowliA, VoL. 1, 1943
Isopterygium Textori (Lac.) Mitt.
Wunba Taung, Chin Hill, no. 8482-A.
Distribution: South India, Malay Peninsula, Annam, Borneo, Philip-
pines, Formosa, China, Japan.
Isopterygium albescens (Schwaegr.) Jaeg.
Tavoy (18 miles down), on trees, 1000 ft., no. 8012 in part.
Distribution: Himalayas, Sumatra, Java, Borneo, Philippines, Celebes,
Pacific islands to Hawaii.
Ptilium crista-castrensis (Hedw.) DeNot.
Burma-Tibet frontier, Adung Valley, 12000 ft, F. Kingdon Ward,
1930-31.
Distribution: Europe, North America, Japan, Yunnan, Sikkim, Caucasus.
HYLOCOMIACEAE
Leptohymenium tenue (Hook.) Schwaegr. -
Nataung, Kareni State, on trees, shady slope, 5000-6300 ft., no. 9429,
9446.2; Pang Yang, Manglon State, on trees, 5000-5500 ft., no. 9642-A.1,
9644; Falam-Haka, banks along paths, 6000 ft., no. 7292; Haka, mossy
forest, epiphytic in shade, 7625 ft., no. 7501; Burma, without locality,
no. 7427-A,
Distribution: North India, Khasia, Philippines.
Foreauella orthothecia (Schwaegr.) Dix.
Taunggyi, F.S.S., epiphytic, damp ground, and on dry slope, 4500 ft.,
no, 9060, 9067, 9498; Mong Yang, Kengtung State, epiphyte, 3500 ft.,
no. 9872; Naungkhio, Thibaw, on rocks, 2800 ft., no. 9540; Mt. Popa,
4000 ft., no. 105; N’Bapa, no. 107.
Distribution: Himalayas, South India, Siam, Philippines.
' Macrothamnium macrocarpum (Hornsch. & Reinw.) Fleisch.
Ah-Shai-Hymin, Ah-Naught, Hymin Lawk Sawk State, on tree, 7600 ft.,
no. 8647, a slender form; Mt. Victoria, no. 8531-4; Taungteik (Falam),
mountain path side, 5000 ft., no. 7258; Haka, on rocks and trees, 6200
ft., no. 7701; Taunggyi, F.S.S., open damp ground and on trees, 4500—
5900 ft., no. 8102, 8773, 9076, 9082, 9087; Pang Yang, Manglon State,
epiphyte, 5000 ft., no. 9646, slender form.
Distribution: East India, Ceylon, Sumatra, Java, Borneo, Philippines,
Hawaii.
Macrothamnium submacrocarpum (Hampe) Fleisch.
Haka, mossy forest, epiphytic, shady, 7625 ft., no. 7497.
Distribution: Himalayas, Khasia.
BAaRTRAM: BuRMA MosseEs 189
POLYTRICHACEAE
Atrichum flavisetum Mitt.
Taunggyi, F.S.S., damp ground and rocks, 4500-5000 ft., no. 9065,
9080.1, 9093.
Distribution: Himalayas, Philippines, Japan.
Atrichum undulatum (Bry. Eur.) Limp.
Taunggyi, F.S.S., on damp ground and rocks, 5000 ft., no. 8782, 9359.
Distribution: wide in north temperate zone, Japan, China, Cochinchina.
Pogonatum Junghuhnianum (Doz. & Molk.) Bryol. Jav.
Taunggyi, F.S.S., on damp ground, 4500 ft., no. 9074, 9077, 9080,
9086; Nataung, Kareni State, damp open slope, 5000 ft., no. 9420.
Distribution: Sikkim, Siam, Java, Celebes, Borneo, Philippines.
Pogonatum inflexum Lindb.
Mt. Victoria, Chin Hills, on rock, 5000 ft., no. 8511; Thandaung,
Taungoo, on rocks and damp places, 3400-4500 ft., no. 6730, 8167;
Taungteik, (Falam), open mountain side, 5000 ft., no. 7264; Mt. Nwalabo,
Tavoy, on rocks and shady dry slope, 5000 ft., no. 8203-A, 8204; Zokhua
(Haka), banks, 4800 ft., no. 7687.4; Pang Yang, Manglon State, epiphyte,
5000-5500 ft., no. 9655, 9689.1.
Distribution: Japan, Formosa, China, Siam.
Pogonatum gymnophyllum Mitt.
Haka, mossy forest, 7600 ft., no. 7577.
. Distribution: Himalayas, China, Tonkin, Formosa, Philippines, Celebes.
Pogonatum microstomum (R. Br.) Brid.
Wunba Taung, Chin Hill, on rocks, 7900 ft., no. 8482; Webula, along
the path where there is damp and shade, no. 7785; Taungteik (Falam),
damp banks, 4500 ft., no. 7549.
Distribution: Himalayas, Khasia, Nilghiri, Ceylon, West China, Philip-
pines.
Polytrichum xanthopilum Wils.
Burma-Tibet Frontier, Adung Valley, 12000 ft., F. Kingdon Ward, Coll.
Distribution: Sikkim. ;
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1(2): 191-194 FARLOWIA July, 1943
ADDITIONS TO THE MOSS FLORA OF NORTHERN CHILE!
EpwIN B. BARTRAM
Only ten species are represented in the collections made by Dr. Ivan M.
Johnston in the montane and coastal regions of northern Chile during
late 1925 and early 1926 yet the series is an unusually instructive one.
The following list includes two new species and four records new to Chile,
as far as my knowledge goes. The types of the new species are in
my herbarium and duplicates are in the Farlow Herbarium, Harvard
University.
POTTIACEAE
Barbula replicata Tay].
Antofagasta Province, Dept. Taltal, Aguada Panulcito, along trail to
the old Andacolla Mine, on slopes above water hole, circa lat. 24° 49’ S.
On earth on dryish cliff wet only by occasional fog, near waterhole,
Dec. 5, 1925, J. M. Johnston, 16 in part.
Pottia flavipes Mont.
Atacama Province, Dept. Copiapo, head of Quebrada de San Miguel,
lat, 27° 25’ S., long. 69° 36’, in very alkaline vega in dry quebrada, grows
where soil is white with alkali, Sierra San Miguel, alt. circa 2700 m.,
Nov. 8, 1925, J. M. Johnston, 2.— Atacama Province, Dept. Copiapo,
Quebrada de San Miguel above Los Marayes, lat. 27° 28’ S., long. 69° 43’,
along slow streamlet in vega, Sierra San Miguel, alt. circa 1500 m.,
Nov. 7, 1925, J. M. Johnston, 8.— Atacama Province, Dept. Copiapo,
head of Quebrada de San Miguel, lat. 27° 23’ S., long. 69° 33’, upper-
most vega, bleak, Sierra San Miguel, alt. circa 3800 m., Nov. 8, 1925,
I. M. Johnston, 9. — Atacama Province, Dept. Copiapo, between Portezuelo
Tolar and Carrizalillito, in small slightly alkaline vega, lat. 27° 37’ S., long.
69° 42’, alt. circa 3000 m., Nov. 9, 1925, J. M. Johnston, 22.— Atacama
Province, Dept. Copiapo, Quebrada Tolar, lat. 27° 34’ S., long. 69° 42’,
uppermost vega, Sierra San Miguel, alt. circa 3600 m., J. M. Johnston,
7.—— Atacama Province, Dept. Vallenar, vicinity of Laguna Valeriano,
circa lat. 29° 3’ S., long. 69° 52’ W., common in vega at head of lake,
alt. circa 4000 m., Jan. 8-10, 1926, J. M. Johnston, 23.— Atacama
Province, Dept. Copiapo, Sierra San Miguel, Vega de San Pedrito, lat.
27° 22’ S., long. 69° 45’ W., in slightly alkaline vega, alt. circa 3000 m.,
Nov. 5, 1925, J. M. Johnston, 3.— Atacama Province, Dept. Copiapo,
Sierra San Miguel, Aguada del Tobaco, lat. 27° 23’ S., long. 69° 46’ W.,
in vega, alt. circa 3000 m., Nov. 6, 1925, J. M. Johnston, 13.— Atacama
Province, Dept. Copiapo, Quebrada de Paipote, Pasto Grande, in slightly
alkaline vega, alt. circa 2500 m., J. M. Johnston, 14.— Northern Argen-
tina: Andes of northwestern San Juan, headwaters of Rio de la Tagua,
* Contribution from the Laboratories of Cryptogamic Botany and the Farlow Her-
barium, Harvard University, no, 173.
191
192 FarLowliaA, Voi. 1, 1943
circa lat. 29° 25/35’ S., long. 69° 50/55’ W., bleak meadow, in wet
places only, circa 4000 m. alt., J. M. Johnston, 15.
I am tempted to believe that Pottia flavipes is a valid species well dis-
tinguished from P. Heimii (Hedw.) by the golden yellow setae and
nearly or quite smooth leaf cells. Typical P. Heimii is well represented
in Fuegia but the above series is so obviously different, even to the naked
eye, that it seems unwise to combine them in one specific concept. The
dentation of the leaves, the length of the setae and the shape of the
capsules are all unstable and variable characters of relatively little
diagnostic importance. ©
Crossidium Roseae (Williams) comb. nov.
Pterogoneurum Roseae Williams, Bull. Torrey Bot. Club 42: 394, 1915.
Antofagasta Province, Dept. Taltal, Aguada Panulcito, along trail to
old Andacolla Mine, on slopes above waterhole, circa lat. 24° 49’ S., on
earth on dryish cliff wet only by an occasional fog, near waterhole,
Dec. 5, 1925. — I. M. Johnston, 16 in part.
New to Chile. These plants are identical with the original collection
from Peru. The costa is not lamellose on the inner face but covered with
filaments 4—5 cells high with the end cell larger, entire and conical. This
structure suggests that the species should be placed in Crossidium.
BRYACEAE
Bryum megalothecium Ther.
Antofagasta Province, Dept. Taltal, Aguada del Panul, circa lat.
24° 47’ S., on cliff wet by streamlet at springs, Dec. 4, 1925, 7. M.
Johnston, 5. — Antofagasta Province, Dept. Taltal, vicinity of Aguada del
Migual Diaz, circa lat. 24° 35’ S., moist soil under rock near waterhole,
Dec. 1-4, 1925, J. M. Johnston, 10.
Bryum flexisetum Mitt.
Atacama Province, Dept. Vallenar, vicinity of Laguna Valeriano, Jan.
8-10, 1926, J. M. Johnston, 28.
New to Chile.
ORTHOTRICHACEAE
Orthotrichum Johnstoni Bartr. sp. nov. FIG. 1-5.
Autoicum, laxe caespitosum, caespitibus fuscescenti-viridibus. Caulis flexuoso-
erectus, ad 3 cm. altus, ramosus. Folia sicca laxe adpressa, humida patentia, 5-6 mm.
longa, anguste lanceolata, acuminata; marginibus inferne anguste recurvis, superne
planis, integris; costa infra summum apicem evanida; cellulae superiores majus
incrassatae, valde pluri-papillosae, basilares lineares, laevissimae. Theca immersa,
sicca plicata, stomata superficialia; exostomii dentes sicca recurvi, processus endostomii
16, anguste lineares; calyptra sparse pilosa; spori 18-24 wu.
This species has a superficial resemblance to some of the North Ameri-
can forms of O. Lyellii Hook. & Tayl. but is thoroughly distinctive in the
BARTRAM: Mosses OF NORTHERN CHILE 193
autoicous inflorescence and the longer peristome. Vicinity of Agua Grande
(“Cachinal de la Costa” of Philippi) near Antofagasta — Atacama
provincial boundary, circa lat. 26° 2’ S., on bushes on flat just back of
high fog-bathed sea cliffs, Dec. 16-18, 1925, J. M. Johnston, 17, type.
FABRONIACEAE
Juratzkea seminervis (Kunz.) Lor.
Antofagasta Province, Dept. Taltal, El Rincon, just north of Paposo,
along trail to old Paranas Mine, circa lat. 24° 59’ S., under low bushes
on rich fog-drenched ridge in fertile belt, circa 400 m., Dec. 7, 1925, I. M.
Johnston, 6.
AMBLYSTEGIACEAE
Hygroamblystegium austro-fluviatile (C. Miill.) Broth.
Atacama Province, Dept. Chanaral, vicinity of Potrerillos, circa lat.
26° 27’ S., long. 69° 30’ W., in cold, bleak meadow, slightly alkaline,
Aguada Pelada, alt. circa 3600 m., Oct. 24, 1925, J. M. Johnston, 19.
This determination is based on a specimen from Chile named by Thériot.
I have not seen the original of Hypnum austro-fluviatile C. Mill. from
South Georgia. The entire leaf margins and the less sharply defined
area of inflated cells in the decurrent auricles will distinguish these
plants from Cratoneuron filicinum (Hedw.) to which they bear a close
superficial resemblance.
Hygroamblystegium crassicostatum Bartr. sp. nov. FIc. 6-8.
Dioicum? Caules elongati, ramosi, rigidi, fluitantes. Folia erecta, adpressa, ovata,
acuminata, ad 1.5 mm. longa; marginibus planis, ubique minutissime denticulatis;
costa pervalida, 100-120 y, lata, percurrens; cellulae hexagonae, plus minus elongatae,
basilares subrectangulares, infimae laxiores. Caetera ignota.
I feel but little hesitation in referring this collection to Hygroambly-
stegium. The species is unique in the Chilean flora but seems to be
closely allied to H. irriguum (Wils.) Loeske var. spinifolium (Schp.)
Grout of North America from which it may be sharply distinguished by
the rigidly erect leaves and the very strong percurrent costa uniform in
width to near the apex. 7
Atacama Province, Dept. Copiapo, Quebrada del Mono at forks below
Mina Maricunga, lat. 27° 5’ 5., long. 69° 27’ W., in water in vega, alt.
circa 3000 m., Quebrada de Paipote, J. M. Johnston, 20, type. — Atacama
Province, Dept. Copiapo, Quebrada de Cerro Bravo at Incaguasi, lat. 25°
42’ S., long. 69° 20’ W., in pool in bleak vega, 2 cm. of ice broken in
collecting this at 8 A. M., Nov. 1926, I. M. Johnston, 1.
Hygrohypnum peruviense Williams
Atacama Province, Dept. Vallenar, vicinity of Laguna Valeriano, circa
lat. 29° 3’ S., long. 69° 52’ W., in water in cold vega at head of lake,
alt. circa 4000 m., Jan. 8-10, 1926, J. M. Johnston, 4.
194, FarLowi4, VoL. 1, 1943
New to Chile. Although not identical with the type material I think
this collection may safely be referred here. The leaves are smaller,
strongly falcate-secund, hooked at the tips of the branches and more
slenderly acuminate but otherwise closely similar in areolation and costal
structure.
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Fig. 6-8, Hygroamblystegium crassicostatum Bartr. 6, Plant x 1.4. 7, Leaf X 25.2.
8, Upper leaf cells and margin x 315.
1(2) : 195-198 FARLOWIA July, 1943
THE SEXUAL DIMORPHISM OF THE TROPICAL MOSSES
OF THE GENUS MACROMITRIUM
MarTHE ERNST-SCH WARZENBACH
Sexual dimorphism occurs in several mosses but its expression is par-
ticularly clear in the genus Macromitrium which grows in the tropics
of the Old and the New World. In 1920 Max Fleischer (Ber. Deut. Bot.
Gesell. 38) noted that the spores of the dioecious species are of unequal
size while those of the monoecious ones are of equal size. He found that
the dioecious plants had dwarf male plants living on the leaves of the
normal large female ones. The dwarf males arose from germinating
spores that had fallen on the leaves. He looked for a relation between
sex and spore dimorphism and believed erroneously, as will shortly be
demonstrated, that the dwarf males resulted from the germination of the
big spores.
The following questions now need to be discussed in the light of new
experiments:' 1) How do distribution of the sexes, and dimorphism
behave in a larger number of species of different origin? 2) Is the
determination of sex genotypic or phaenotypic? 3) Is the difference in
size of the spores an expression of sexual dimorphism?
Statistical investigations on the variation of the spores have been
carried out with herbarium material of thirty-nine species from Java,
India, Ceylon, Japan, Australia, the islands of the Pacific, Africa, Central
and South America. The specimens have also been examined for dwarf
males, and male and female gametangia. The ten monoecious species that
were investigated proved to be homosporous, the measurements of their
spores gave the results graphically represented in figure 1 which is
characterized by having a single peak. Twenty-seven dioecious species
produced normal sized female plants and epiphytic dwarf males, seven
of them were homosporous and twenty were more or less distinctly
heterosporous as shown in figure 2 which clearly shows the two peaks.
Thus it can be said that in the dioecious species dimorphism of the
stems is always clearly evident while spores are mostly but not always
dimorphous. Macromitrium Blumei has a special position in that its
spores are not distinctly heterosporous since, as is shown in figure 3,
there are not two definite peaks. According to Denning, it is polyoical,
i. e. it has partly dimorphic dioecious and partly normal sized monoecious
plants.
The study of the development of the spores of a heterosporous species
from Java has shown that the spore-mother-cells first produce all spores
of an equal size. They soon escape from the tetrad formation and later
*Vide M. Ernst-Schwarzenbach: Zur Kenntnis des’ sexuellen Dimorphismus der
Laubmoose, — Arch, der Julius Klaus-Stiftung Ziirich 14, 1939; — Weitere Mitteil-
ungen tiber den sexuellen Dimorphismus der tropischen Laubmoosgattung Macro-
mitrium. 2. Jahresber. Schweiz. Gesell. f. Vererbungforsch. Arch. der Julius Klaus-
Stiftung Ziirich 17, 1942.
195
196 Fartowla, VoL. 1, 1943
the differentiation begins and results in big and small spores whose rela-
tion to a tetrad cannot be seen. The ratio of big to small spores is
about 1:1.
The relation of spore dimorphism and sexual determination is of
special interest and can only be ascertained by culture experiments. With
material from Java, especially M. salakanum, one-spore cultures could
be made in hanging drops of nutritive solution. The big as well as the
small spores germinated well. On the protonemata of the small spores,
dwarf males grew in two to three months and produced ripe antheridia
about two months later. The cultures remained alive for about one and
one-half years, exceptionally three and a half years, repeatedly forming
new secondary protonemata with dwarf males on each. Thus the pos-
sibility that an early male phase is followed later by a female phase is
excluded and the genotypical method of male determination of the small
spores is evident.
The products of the germination of the large spores gave considerable
trouble. The protonemata from the big spores also produce small stems
but these soon differ from the dwarfs by their attenuate appearance
(Fig. 5). They are much less numerous and grow much more slowly
than the dwarfs, but eventually they are quite similar to the females
grown under natural conditions.
The greatest difficulties were encountered in raising the plants that
originated from the big spores to the fertile state. Only in one culture
did several plants bear numerous archegonia. Similarly, the Macro-
mitrium tufts from the natural habitats, cultured at Ziirich for four years,
very seldom formed archegonia and sporogonia. Therefore the evidence
that the normal-sized plants are really female-determinate had to be ob-
tained by another method: by the regeneration of leaves, since if the leaves
of the big female plants constantly give big plants, these are genotypically
female-determinate, whereas if they also give stems with antheridia, they
are genotypically monoecious. The result of the cultures was very clear:
the leaves of big female plants produced protonemata which always gave
rise again only to big plants of normal size. Thus it is evident that
genotypical sex determination of M. salakanum is bound to sexual
dimorphism.
The expression of sexual dimorphism of M. salakanum is different
during its development since in the spores there is a difference in size
and in color: the female-determinate spores, of a mean diameter of
36.57 mp, are green, while the male-determinate spores, of a mean diameter
of 19.60 », are yellow. The primary protonema growing from the spore
depends so much on the environment that no dimorphism can be observed.
It is only later on when the young stems form new secondary protonemata
that dimorphism is noticeable again: the female has more the appear-
ance of rhizoids while the male resembles chloronema.
The most distinct morphological and physiological dimorphism is
ERNST-SCHWARZENBACH: SEXUALITY IN MACROMITRIUM 197
found in the moss stems: the dwarfs are 14-1 mm. high, the orthotrope
parts of the female stems are 2-3 mm. The length of the leaves of the
male is 0.30—0.45 mm. and that of the female leaves is 1.5-2.5 mm.
The size of an organ is generally determined by the size and number
of the cells. Measurements have shown that the differences in the organs
of M. salakanum are not dependent on the cell size because it is never
smaller, perhaps even larger, in the males than in the female. The size
of the organs is determined by the cell number: in the leaves of the
dwarfs, their mean number in length is 33.5, and in width 23.2 whereas
the female leaves are 204 cells in length and 74.7 cells in width. There-
fore the number of the cells in the male plant is only a small part of the
number in the female.
SUMMARY
The dioecious species always have dwarf male individuals which are
the only ones to produce antheridia. If the normal sized stems, or the
regeneration of their leaves, produce sexual organs, they are archegonia.
Not all but many of the dioecious species show dimorphism in the
spores, heterospory, whilst the monoecious are always homosporous.
Species of Macromitrium from Java have shown their dimorphism to be
determined genotypically. The small spores give dwarf males and the
large spores produce large female plants. Sexual dimorphism is ex-
pressed in the spores, secondary protonema, and stems. The difference
in the size of the organs depends on the number of cells per organ.
INSTITUT FUR ALLGEMEINE BOTANIK
ZURICH, SWITZERLAND.
198 Fartowia, VoL. 1, 1943
OOOO ovo D.PX900°0 OO
scene co oe ao
Fig. 1. Diagram showing size vs. distribution of the homosporous species,
Macromitrium Reinwardti. Fig. 2. Diagram showing size vs. distribution of the
heterosporous species, Macromitrium. salakanum. Fig. 3. Diagram showing size vs.
distribution of the indistinctly heterosporous species, Macromitrium Blumei. Fig. 4.
Male dwarf on the leaf of a female plant from Java. Fig. 5. a. Dwarf male plants,
and b. Female plants from cultures of the same age, and drawn to the same magnifica-
tion ( 22.4).
1(2): 199-244 FARLOWIA July, 1943
THE MORPHOLOGY, TAXONOMY, AND DISTRIBUTION
OF COCCIDIOIDES IMMITIS RIXFORD AND GILCHRIST 1896 !
kK. E. Baker, E. M. Mrax, anp C. E. Situ 2
The fungus, Coccidioides immitis Rixford and Gilchrist 1896, is the
causative agent of a disease, coccidioidomycosis (Dickson, 1937a), hav-
ing two principal clinical manifestations, namely: an acute pulmonary
infection which is sometimes accompanied by erythema nodosum, and
usually heals spontaneously, and a generalized infection, coccidioidal
granuloma, which frequently terminates fatally. When the generalized in-
fection occurs, it is nearly always as a dissemination from a preceding
primary pulmonary infection.
The first case of coccidioidal granuloma was described by Posadas
(1892, 1900) and Wernicke (1892). They described the morphology
of the parasite in human tissues in considerable detail. Rixford and
Gilchrist (1896) gave a more complete morphological description of the
parasitic phase of C. immitis in their study of the first two North American
cases. Ophiils and Moffitt (1900) isolated a white fungus from their
patient, the third North American case of coccidioidal granuloma to be
reported. They proved the etiological relationship of the fungus and
gave a brief description of both the parasitic and cultural phases of
the organism,
Ophiils (1905) gave the first complete description of C. immitis. He
described in detail the formation of endospores by cleavage planes in
animal tissues and the formation of chlamydospores in culture. Since
Ophuls’ description, numerous investigators have made many morpho-
logical studies, notably: Wolbach (1904), MacNeal and Taylor (1914),
Fonseca and Area Leao (1927), Fonseca (1928), Ahlfeldt (1929), Al-
meida (1929; 1930a,b, 1932a,b,c, 1934a,b) Agostini (1932), Ciferri
(1932), Moore (1932, 1938), Castellani and Jacono (1933), Takahashi
(1933), Ota and Kawatsure (1933), Benham (1934), Ciferri and Redaelli
(1936), Jordan and Weidman (1936) and Dickson (1937b). Most of
these investigators have added little to Ophiils’ description.
Ciferri and Redaelli (1934a,b, 1936) reported that if spherules of a
certain strain were cultured semianaerobically by the technic of MacNeal
and Taylor (1914), an isogamous or heterogamous copulation sometimes
occurred prior to sporulation. This finding, however, lacks confirmation.
Lack (1938) devised a method for producing spherules from chla-
mydospores “in vitro.” He incubated chlamydospores semianaerobically
in a medium composed of glucose broth and partially coagulated egg
albumin. Chlamydospores grown in this manner enlarged to form
spherules rather than germ tubes as in ordinary cultures. Both semi-
‘This investigation was supported by a grant from the Rosenberg Foundation.
*From the Department of Public Health and Preventive Medicine, Stanford: Univer-
sity School of Medicine and the University of California.
199
200 Fartowia, VoL. 1, 1943
anaerobiosis and partially coagulated egg albumin were essential to
success.
Baker and Mrak (1941) reported that spherules (termed culture spher-
ules) are formed by certain strains in old cultures, slide cultures and
in a semi-solid medium containing 0.05% copper sulfate. They con-
cluded that for these particular strains the culture spherules form a
normal phase in the life cycle of the fungus.
Although a considerable number of papers have been published on the
morphology and life history of C. immitis, the information is still incom-
plete. There has been no extensive comparison of strains isolated in
widely separated areas. The taxonomy is confused because of the lack
of morphologic information and the anomalous relationships of this or-
ganism. Furthermore, except for the paper of Ciferri and Redaelli
(1936), published in Italian, there is no complete review of the literature
concerning the morphology, taxonomy, and distribution of this organism.
The purpose of this investigation has been to compare morphologically
fifteen strains of C. immitis secured from widely separated areas in order
to obtain information of taxonomic significance and to obtain evidence
regarding the systematic position of the fungus.
EXPERIMENTAL PROCEDURE
Fifteen strains of C. immitis representing isolates from various clinical
types of coccidioidomycosis and various geographical locations were stud-
ied. The data concerning these cultures are tabulated in Table I.
Animal and culture inoculations were made with suspensions of chla-
mydospores and hyphal fragments in physiological saline solution. Each
suspension was prepared by shaking a loopful of a month-old Sabouraud’s
glucose agar culture in 5 cc. of physiological saline solution. Experi-
mental animals were usually inoculated with 0.5 cc. of suspension, a dose
more than sufficient to insure infection.
The studies of the gross morphology were made on single-spore cultures
obtained by the method of Hanson and Smith (1932), and the three media
used were potato dextrose agar (see Rawlins, 1933), Sabouraud’s glucose
agar (see Lewis and Hopper, 1939), and a synthetic agar similar to a
medium proposed by Stewart and Meyer (1932) except that the 1%
ammonium chloride was reduced to 0.1%, and 2% agar was added.
All studies of the morphology in animal tissues (parasitic stages) were
made on material obtained from mice or guinea pigs experimentally in-
fected with cultures number 4 and 46. These cultures represented the
typical and extreme forms under cultural conditions. However, all strains
were inoculated into experimental animals and were found to be patho-
genic and to present a uniform morphology even though some strains had
been held in culture for several years. The mice, inoculated intraperi-
toneally, usually died the second week following inoculation. Guinea pigs,
inoculated intratesticularly, were killed after three to four weeks. Fresh
BAKER, Mrak & SMITH: CoccIDIOIDES IMMITIS 201
material was examined by mounting pus or infected tissue fragments in
cotton blue lacto-phenol (see Rawlins, 1933).
Tissues for cytological studies were fixed for 48 hours in the following
fixatives: Bouin’s solution, Allen’s modification of Bouin’s solution
(PFA3), Zenker’s solution, Carnoy’s solution, Karpachenko’s solution,
Flemming’s strong solution, Flemming’s strong solution diluted 1 : 2,
Flemming’s weak solution, 10% formalin in 0.85% sodium chloride solu-
tion, Miiller’s solution, and formalin-acetic acid-alcohol solution (formula
number 2 of Rawlins, 1933). Sections 10 » thick were used in most of
the cytological studies. They were stained with Heidenhain’s iron hema-
toxylin (schedule given by Rawlins, 1933, except that 0.5 instead of 2%
hematoxylin solution was used), Flemming’s triple stain, crystal violet,
methylene blue, thionin, and Feulgen’s stain. Usually no counterstain
was used, although occasionally erythrosin or eosin was tried. Differen-
tial destaining was very difficult. In most cases the cytoplasm retained
the basic nuclear stains as tenaciously as the nuclei. The best nuclear
staining was obtained with tissue fixed in Flemming’s strong solution
diluted 1 : 2 followed by Flemming’s triple stain.
Usually culture material was mounted in cotton blue lacto-phenol. Slide
cultures were prepared by dipping slides, sterilized by immersion in al-
cohol followed by burning off the excess alcohol, into melted agar (usually
potato dextrose agar). These agar-coated slides were then placed on
bent glass rod supports in petri dishes containing a piece of sterile, moist
blotting paper. After the agar solidified, a drop of spore suspension was
placed on the slide. Finally the petri dishes were sealed with “parafilm”
and stored at room temperature. After development, the excess agar on
the bottom of the slide and around the culture was removed. The cultures
were sometimes studied by adding a drop of cotton blue lacto-phenol and
a cover slip. Other cultures were fixed in various fixatives, washed and
after the excess agar was removed, dried, and stained. For staining the
thallus, the slides were stained in 1% aqueous cotton blue with 1%
acetic acid and destained slightly in acidified 95% alcohol to remove
excess stain from the agar before dehydrating and mounting. Spore
germination was studied in moist chamber slides.
Culture material for cytological observations was fixed for 48 hours
in the following fixatives: Bouin’s solution, Carnoy’s solution, Zenker’s
solution, Flemming’s strong solution diluted 1 : 2 or formalin-acetic acid-
solution. The last was the most satisfactory. When other fixatives
were used, differential destaining was not obtained. Numerous stains were
tried but only iron hematoxylin gave satisfactory preparations. The
nuclei of germinating spores were stained most satisfactorily in concen-
trated hematoxylin solution (2-4%) for about 5 minutes, followed by a
short differentiation in 4% ferric alum. In contrast, the nuclei of ma-
ture mycelia and chlamydospores were stained best in dilute hematoxylin
for about 24 hours followed by relatively long differentiation in 4% ferric
202 Fartowia, VoL. 1, 1943
alum. Other staining technics tried were the same as those described
for animal tissues.
Microchemical tests as described by Rawlins (1933) were made on
both culture and tissue material for cellulose, chitin, callose, glycogen,
starch, and fat.
MORPHOLOGICAL OBSERVATIONS
Gross Morphology
Hansen and Smith (1932) and Hansen (1938) have shown that variation
occurs in pure cultures of certain imperfect fungi. They have established
that pure cultures of these fungi are mixtures of two stable types and
that these types can be segregated by making series of single spore cul-
tures. Whether this phenomenon occurs in C, immitis is a subject for
speculation.
A series of 25 single spore cultures of each of the fifteen cultures listed
in Table I were made on synthetic agar. Some of the cultures exhibited
slight differences. However, in subsequent single-spore cultures prepared
from representative single-cell cultures of these strains, these differences
were not constant. In one case, a third set of single-cell cultures gave
the same results as did the second. To obviate the possibility that re-
peated culturing had eliminated variants, a series of 50 single spore cul-
tures was made directly from the pus of a coccidioidal granuloma patient.
All of the cultures appeared to be identical.
The gross morphological differences among the various strains on all
three media were very striking. Photographs showing the strain differ-
ences on the three media incubated in the light are shown in Plate I, figs.
1 and 2; Plate II, fig. 1. The differences were most striking on potato
dextrose and Sabouraud’s glucose agar. For example, on Sabouraud’s
glucose agar the colony surfaces ranged from an almost flat, smooth col-
ony with a tough tenaceous surface to a convex cottony colony, the aerial
growth of which fragmented very readily. There were all gradations
between these two types. Strain number 27 tended to form finger-like
processes extending from a central knob which superficially resembled
coremia. Similar structures were also observed in strain number 40, but
instead of arising from a central prominence they were scattered over
the surface of the flat central portion of the colony, assuming a counter-
clockwise spiral arrangement.
In some strains (numbers 4 and 10) variations which appeared to be
sectoring, were observed. However, subcultures from these sectors de-
veloped normally. If the original sectored cultures were incubated for
two or three months, the entire surface of these colonies became covered
with the downy type of growth characteristic of the sectors.
It has been shown by Coons and Levin (1926), Harter (1939), and
Snyder and Hansen (1941) that light favored sporulation and darkness,
mycelial development in the Fusaria and certain other fungi. On the
TABLE 1—HISTORY OF CULTURES
Cul- | Date of
ture Isola- : Geographical
No. tion Isolated by Location Clinical History Previous Investigations Remarks
+ 1938 Dr. C. E. Smith, Stanford | Kern Lake, Calif. |Coccidioidal granuloma, Baker and Mrak (1941) Fatal
Univ. School of Medicine, culture from abscess on arm
San Francisco, California
7 1938 Dr. C. E. Smith, Stanford | Yolo Co., Calif. {Primary coccidioidomycosis Recovered
with erythema nodosum,
culture from sputum
8 1938 Dr. C. E. Smith, Stanford | Bakersfield, Calif.|Primary coccidioidomycosis Baker and Mrak (1941) Recovered
with erythema nodosum,
culture from sputum
10 1938 Kern County Health Arvin, Calif. Coccidioidal granuloma
Dept. without cutaneous lesions,
culture from pleural fluid :
11 1938 Kern County Health Oildale, Calif. Primary coccidioidomycosis Recovered
Dept. without erythema nodosum,
culture from sputum
15 1938 Dr. C. E. Smith, Stanford | Tulare, Calif. Primary coccidioidomycosis Illness originally di-
with erythema multiforme, agnosed as small-pox.
culture from sputum Recovered
17 1932(?) |Dr. C.-F. Lehman, San Texas Coccidioidal granuloma, Ota and Kawatsure (1933) {Culture from Dr. F.
Antonio, Texas culture from lesion on left Kawatsure (1934), Ciferri} D. Weidman
fore) and Redaelli (1936)
23 1924(?) |Dr. D. J. Davis, Univer- ? Coccidioidal granuloma Ota and Kawatsure (1933) |Culture from Dr. F.
sity of Chicago, Chicago, D. Weidman
27 4 Dr. A. Castellani, London 2 Coccidioidal granuloma (?) Castellaniand Jacono (1933)|Culture from Dr. P.
School of Tropical Med. Ciferri and Redaelli (1936)| Redaelli
London, England
28 ? Dr. A. Castellani, London | The Balkans Coccidioidal granuloma Castellani and Jacono (1933)|Culture from Dr. P.
Ciferri and Redaelli (1936)| Redaelli
29 1S Dr. A. Castellani, London | Naples, Italy Coccidioidal granuloma Castellani and Jacono (1933)|Culture from Dr. P.
Ciferri and Redaelli (1936)| Redaelli
40 1939 Dr. C. E. Smith, Stanford | Bakersfield, Calif. |Primary coccidioidomycosis Recovered
with erythema nodosum,
culture from sputum
41 1939 E. E. Baker, Stanford Stockton, Calif. |Bovine coccidioidal granu- Baker and Mrak (1941)
Univ. School of Medicine loma culture mediastinal
San Francisco, California lymph node
46 1933 Dr. du Bray, St Lukes Probably San Coccidioidal granuloma Lack (1938) Lack and Fatal
Hospital, San Francisco, |’ Jeequie Valley, Cronkite (1940)
California alif.
63 1939 Dr. C,. E. Smith, Stanford Roly neotte NA Coccidioidal granuloma Recovered
alif.
SILINW] SAGI01dI990)
HLINS Y MVUy “Wav
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204 Fartowia, Vou. 1, 1943
other hand, Yarwood (1937) showed that sporulation of certain of the
downy mildews was more abundant in the dark, and that there was a
diurnal effect. Most investigators who have studied C. immitis, grew
the fungus in the dark of an incubator. Sets of cultures of C. immitis
were maintained in both light and darkness but no significant differences
were noted. Some of the cultures showed somewhat more abundant de-
velopment in the light while others developed slightly better in the dark.
There was no difference in the microscopic morphology.
The strains appeared to be quite stable. There was no correlation
between gross morphology and clinical history, geographic source of
isolation, or length of time in culture. Although the gross morphological
differences observed in the fifteen strains were marked, they were not
sharply defined but seemed to form a continuous series ranging from
smooth surfaced to cottony and even mealy surfaced types. The differ-
ences were observed in “‘mass transfer” cultures as well as in the single-
spore cultures. The taxonomic significance of the gross morphological
differences will be discussed later.
Microscopic Morphology
Discussion of the microscopic morphology is divided into that of the
organism as a parasite in animals (termed “parasitic phase”), and in
culture (termed “culture phase”). These terms are used solely for con-
venience. Most strains showed a remarkable uniformity in microscopic
morphology, the differences being quantitative rather than qualitative.
Some strains produced a greater amount of surface mycelium and conse-
quently relatively greater numbers of chlamydospores. Chlamydospores
were of two types, cylindrical and spherical, and were either terminal or
intercalary, frequently catenulate. Some strains produced more of the
spherical type than others, but this character was not constant. The fol-
lowing description is based almost entirely upon the morphology of strains.
number 4 and 46. These two strains represented the extremes in mor-
phologic variation, as will be pointed out later. Most strains, however,
were not distinguishable microscopically from number 46. The uniform-
ity in the morphology of the parasitic phase was even greater than of
the cultural phase.
Parasitic Phase: Either in the tissues of man, naturally infected with
C. immitis, or in the tissues of experimentally infected animals, the ma-
ture fungus is a spherical structure, usually termed a spherule, containing
numerous endospores (Plate II, figs. 2, 6, 7, 9, 10; Plate III, fig. 1).
The nature of the spherule will be discussed later. It is surrounded by
a refractile double wall which may be 2 » thick. The thin inner wall
is composed of chitin and the thick outer wall of a substance which does
not react with any stains used and is not chitin, cellulose, or callose as
determined by the procedures described by Rawlins (1933). It is sug-
gested that this outer wall is composed of some polysaccharide although
BAKER, Mrak & SMITH: CoccipIoIDEs IMMITIS 205
attempts to adapt Molisch’s test for carbohydrates to microscopic use
failed. Usually the spherule wall is smooth but sometimes may be cov-
ered with numerous excrescences. These structures may be fine (Plate
Ill, fig. 9) or club-like (Plate III, fig. 2). The fine excrescences have
been observed by many investigators and have been termed “prickles”
by Dickson (1937b). The club-like excrescences formed around the
spherules of C. immitis have been reported by Almeida (1934b) and
Henrici (1940). These structures, termed “actinomycetoid forms” by
Henrici, are not peculiar to any type of fungus. They have been ob-
served by Almeida surrounding Paracoccidioides braziliensis, Asper-
gillus fumigatus and Trichosporium pedrosoi in human and experimental
infections with these fungi. Henrici (1940) stated that they are also
found in sporotrichosis and have been obtained experimentally with cer-
tain acid-fast bacteria and actinomycetes. He considered them to be the
result of an allergic state developed following infection.
The spherules are quite variable in size, ranging from about 15 to 80 »
in diameter. Schenken and Palik (1942) report a very large spherule
measuring 262 p» in diameter with a capsule 21 » thick. Mature spherules
are usually found in giant cells or histiocytes of the host (Plate II, figs.
2 and 3). The endospores contained within the spherule may be spher-
ical or sub-spherical (Plate II, fig. 2; Plate III, fig. 1) or irregular in
shape (Plate II, figs. 6, 7,9, and 10). The endospores may be as small
as 2 » or as large as 5-10 p» in diameter. Occasionally a spherule is
seen containing only a few large endospores. Plate III, fig. 5 shows such
a spherule containing three endospores each about 40 » in diameter. The
exact significance of such a structure is not known but it is supposed that
in this case cleavage ceased after the formation of the second cleavage
plane. The small irregular endospores have a thin wall and probably
represent immature spores. The larger endospores usually have a thick
wall resembling that of spherules (Plate III, fig. 7).
The endospores are released by rupture of the wall of the spherules.
In view of the variation in size and shape of the endospores, it is reason-
able to assume that when the spores are first formed they are irregular
and then tend to attain a more nearly spherical shape. Presumably the
spores enlarge and finally rupture the spherule wall because of increased
pressure on the wall. The released spores develop into immature spher-
ules, and sporulation again occurs by the formation of successive cleavage
planes. There appears to be no relationship between size of the imma-
ture spherules and the initiation of spore formation. Plate II, fig. 5
shows a small spherule about 15 » in diameter in which the first cleavage
plane has formed, and Plate II, fig. 8 shows a very large spherule about
80 » in diameter in which cleavage planes have not as yet appeared. In
Plate II, figs. 6 and 7 are shown spherules in which only a few cleavage
planes have been formed. In these spherules the spore initials are still
quite irregular. The protoplasm in immature spherules may fill the
206 Fartowia, Vou. 1, 1943
entire structure (Plate II, fig. 8) or be distributed around the periphery
(Plate II, figs. 4 and 11). Cleavage planes may divide the protoplast
first into two, then four, etc., irregular segments (Plate II, fig. 9). These
spore initials may then enlarge to form a few large endospores (Plate III,
figs. 1 and 5) or additional cleavage planes may divide them into many
small spores (Plate II, fig. 10). In the tissues of guinea pigs the spher-
ules are usually non-vacuolated and the cleavage planes may divide them
into many small spores. In contrast, in mice the spherules are usually
vacuolated and the radial cleavage planes cut the peripheral protoplasm
into segments as shown in Plate II, fig. 9 (although this photograph was
taken of material from a guinea pig, it is more characteristic of that from
infected mice).
Ciferri and Redaelli (1934a,b, 1936) reported that one strain of the
fungus studied by them underwent conjugation, either isogamous or
heterogamous, prior to sporulation. This phenomenon was observed in
semianaerobic cultures prepared from pus of an experimentally infected
guinea pig according to the technic of MacNeal and Taylor (1914).
They stated that the protoplast of one cell passed through a conjugation
bridge and fused with the protoplast of the other gamete. However, they
were unable to secure sufficient material for cytological study. In pus
from a guinea pig infected with strain number 40, structures resembling
the figures of Ciferri and Redaelli were seen (Plate III, figs. 3, 4, 6, 7, and
8). In all of the structures, both halves contain protoplasm. Plasmog-
amy does not appear to have taken place either in the structures in which
spores have not yet formed (Plate III, figs. 3 and 4) or in those in which
spores have formed (Plate III, figs. 6, 7, 8). In fact, in Plate III, figs.
6 and 7, spores are present in one cell and the accompanying cell still
contains protoplasm indicating definitely that plasmogamy has not oc-.
curred. Unfortunately the occurrence of these structures in tissue is so
infrequent that staining procedures to determine the possible occurrence
of karyogamy could not be carried out. However, in view of the absence
of proof of karyogamy it is not wise at present to consider these structures
as sexual. “Dumb-bell” shaped structures and swollen cells joined by
a tube are seen in old cultures of all strains (Plate III, fig. 10). It
seems likely that the structures seen in guinea pig pus may have orig-
inated directly from injected culture material and not by fusion of
spherules. It is not possible to state whether the structures observed by
Ciferri and Redaelli originated in this manner or whether they were
actually the result of a sexual process.
The morphological changes accompanying the transformation of cul-
ture material (chlamydospores, oidia, and hyphal fragments) have not
been investigated. There is some evidence that when culture material is
injected into an experimental animal, the chlamydospores merely “round
up” and enlarge into spherules. Ophiils (1905) stated that unless chla-
mydospores are present, culture material is not capable of producing
Baker, Mrak & SmitTH: CoccipioiEs IMMITIS 207
spherules when injected into experimental animals. However, Chope
(unpublished data quoted by Dickson, 1937b) showed that very young
cultures containing no chlamydospores could also produce spherules.
The observations of Chope are as follows:
“|. after the injection of the suspension of chlamydospores the outline of the
hyphae may be intact on the third day but . . . the chlamydospores are somewhat
rounded and their outer capsule is well defined. At forty-eight hours some of these
chlamydospores are well rounded out although still connected together as in the
hyphae. At sixty hours the spherules are well developed and are as large (about
30 1) as the immature spherules are likely to be when observed in pus from a
patient. By seventy-two hours some of the spherules are filled with endospores,
and in ninety-six hours some of them have burst, showing the manner in which
the endospores are liberated and how the ‘envelope’ is formed.”
Considerable time has been devoted to a study of the nuclear changes
accompanying the development of spherules, but they have not been
studied to the extent desired because of the small size of the nuclei and
the difficulty in obtaining satisfactory stained preparations. In most of
the preparations, the cytoplasm retained the nuclear stains as tenaciously
as the nuclei. However, sections of mouse lung fixed in Flemming’s
strong solution diluted 1:2 and stained with Flemming’s triple stain
permitted some observations. The nucleus is characteristically spherical
and about 1.5 » in diameter. It has a rather delicate membrane which
stains with gentian violet and a very small eccentrically placed nucleolus
about 0.2 » in diameter which stains with safranin (Plate V, figs. 2 and
3). It was not possible to make chromosome counts or to study the de-
tails of nuclear division because of the small size and because only very
few dividing nuclei were seen in the material examined. It is assumed
that the nuclei divide by the usual mitosis but the division figures were
too small and indistinct for accurate interpretation. The smallest spores
(or immature spherules, about 7 » in diameter), seen free in the tissues
contain two nuclei (Plate VI, fig. 1), which divide (Plate VI, fig. 2) sim-
ultaneously. This process is repeated, giving successively four, eight,
etc., nuclei (Plate VI, figs. 3, 4,5, and 6). The number of nuclei formed
in this structure could not be determined beyond about eight nuclei. In
a large spherule about 50 p» in diameter (Plate VI, fig. ©) numerous nuclei
are seen scattered throughout the peripheral protoplasm. It was not
possible to follow the nuclear changes during the formation of cleavage
planes. In Plate VI, fig. 7 is shown a large spherule divided into numer-
ous irregular segments containing a variable number of nuclei. These
segments appear similar to the protospores of some of the Mucorales.
Probably some of these segments normally would be subdivided further
before formation of mature endospores. Plate VI, fig. 8 shows a mature
spherule containing mature endospores with thick walls. These endo-
spores contain from one to four nuclei. In one spore a dividing nucleus
can be seen. It seems probable that all of the nuclei in a developing
spherule normally divide simultaneously, for if they divide independently
it is likely that more dividing nuclei would have been observed.
208 Fartowia, VoL. 1, 1943
Although Moore (1932), Dodge (1935), and Stewart and Kimura
(1940) stated that the spherules are asci, no evidence of “free cell”
formation was observed. The cytological observations are essentially in
agreement with the observations of Ciferri and Redaelli (1936). They
stated, however, that the endospores are uninucleate, while according to
the present observations they may be uninucleate, or irregularly multi-
nucleate.
Satisfactory preparations showing nuclear changes during germination
of endospores were not obtained. However, it seems possible that these
changes are the same as in germinating chlamydospores (Plate IV, figs.
13 and 14; Plate V, fig. 1; Plate VIB, fig. 1) which will be described later.
Cultural Phase: When pus is planted on artificial culture media, the
immature spherules and the endospores, either free or contained within
a spherule, germinate by the formation of one or more germ tubes (Plate
IIT, figs. 11, 12, 13, 14). Very soon after the formation of a germ tube,
which occurs as early as four hours after inoculation, branches may form
(Plate III, figs. 11 and 12). Septation may occur either before (Plate
IIT, fig. 12) or after branching (Plate III, fig. 13). After twenty-four
to forty-eight hours a complex network of hyphae is formed. In young
cultures (four to seven days) the hyphae are 2-4 » in diameter and ir-
regularly septate (Plate III, figs. 15 and 16). After about seven to ten
days, many of the cells of the aerial hyphae develop into oidia and
chlamydospores (Plate III, figs. 18 and 19; Plate IV, figs. 1, 2, 3, and 4).
Oidia probably represent stages in the formation of chlamydospores since
they are similar in structure to chlamydospores, differing only in having
slightly greater size and lacking a thick wall. These oidia are not strictly
comparable to the conidia of the genera Oidium and Oospora, in that they
may be either terminal or intercalary. There may be several oidia to-
gether in a chain or there may be short chains separated by ordinary
hyphal cells. The oidia mentioned by Ophiils (1905), MacNeal and
Taylor (1914), Wolbach (1904) and others are in reality not oidia but
chlamydospores. The oidia and chlamydospores usually disarticulate
readily. At about the same time that the oidia and chlamydospores are
formed some of the hyphal cells become swollen at one end resembling
the clavate cells of the “racquet” mycelium of the dermatophytes (Plate
III, fig. 17). These cells are thin-walled and quite variable in size. The
oidia are about 2-4 » in diameter and may attain a length of 6-8 p». The
chlamydospores are generally cylindrical or with sides slightly convex and
measure 2—4 x 2-4 » or 2-4 x 3-6 ys (Plate III, figs. 18 and 19). However,
they are at times ellipsoidal to nearly spherical. Strain number 4, in
particular, produces this type of chlamydospore commonly measuring
up to 10 » and occasionally 20 y». in diameter (Plate IV, figs. 2 and 3).
Sometimes chlamydospores are seen which appear to have been formed
by septation of spindle-shaped cells into two, three, and even four cells
(Plate II, fig. 19; Plate VII, fig. 5). The composition of the medium
has no effect upon the characteristics of the chlamydospores.
Baker, Mrak & SmitH: CoccipioIpes IMMITIS 209
When chlamydospores are placed on fresh media they germinate by
the formation of germ tubes in much the same manner as do endospores
(Plate IV, figs. 5 and 6). Subsequent development is identical with
that of endospores.
The cells of the hyphae usually contain one or two nuclei (Plate VII,
figs. 2 and 3). The chlamydospores are usually either uni- or binucleate
(Plate VII, figs. 4 and 5) but some of the larger chlamydospores may
contain four nuclei (Plate VII, fig. 4). When the chlamydospores
germinate, germ tubes usually form before the first nuclear division
(Plate VII, fig. 1). The nucleolus in the nuclei of germinating chlamy-
dospores is considerably larger than that in the nuclei of spherules,
endospores, mature hyphal cells, or chlamydospores, and ranges from
about 0.75 to 1.0 » in diameter (Plate IV, figs. 13 and 14; Plate V, fig. 1).
There are either one or two nuclei in the cells formed by septation of
the young hyphae.
It is generally considered that the only vegetative reproductive struc-
tures produced by C. immitis growing in culture are oidia and chlamydo-
spores. MacNeal and Taylor (1914), and Ciferri and Redaelli (1936)
observed continued multiplication of spherules when pus from infected
animals was inoculated into a medium composed of Tyrode’s solution to
which was added slices of sterile tissues. Lack (1938) showed that under
conditions of semianaerobiosis chlamydospores developed into spherules
when inoculated into beef infusion broth containing partially coagulated
ege albumin. These two technics attempted to simulate conditions in the
animal body.
However, Baker and Mrak (1941) have shown that certain strains of
the fungus produce structures in culture closely resembling the spherules
found in animal tissues. These are not carried over from animal tissues
because strain number 4 has been maintained in culture since 1937. These
culture spherules range in size from about 10 to 20 » in diameter, and
are characteristically spherical but sometimes oval. They are terminal
or intercalary. Endospores develop by the formation of successive cleav-
age planes in the culture spherule. Mature spores are usually irregular
in shape but occasionally may be spherical. When the spherules are
placed in liquid media or on agar, they germinate by the formation of
germ tubes.
Subsequent to the above investigation, one immature culture spherule
in which the first cleavage plane had formed was observed during exam-
ination of a four-month-old culture of strain number 46 (Plate IV, fig. 7).
Most of the stained preparations of culture spherules proved inadequate
for cytological study. In a slide culture fixed in formalin-acetic acid-
alcohol solution and stained with iron hematoxylin, a few spherical chla-
mydospores without cleavage planes were seen (Plate VII, figs. 6, 7, and
8). Structures with two, four, and eight nuclei were observed. Unfor-
tunately, the entire development could not be studied.
210 FarLtowra, Vou. 1, 1943
In one-month-old slide cultures of strain number 4, fixed in formalin-
acetic acid-alcohol solution and stained with 1% aqueous cotton blue
solution and mounted in balsam, structures resembling zygospores were
observed (Plate IV, figs. 8, 9, 10, 11, and 12). The earliest stage re-
sembled nothing more than hyphal fusion (Plate IV, fig. 8). Later the
bridge joining the two hyphae became broader and, finally, the structure
became spherical and assumed a thick wall (Plate IV, fig. 12). The
supporting hyphae were then empty. Sometimes the smooth wall, re-
sembling the wall of the spherules (tissue and culture), was quite thick.
The explanation of these structures is somewhat difficult but it is possible
that they represent primitive zygospores lacking the usual suspensors;
on the other hand, they may be only an unusual form of hyphal fusion
that has formed chlamydospores.
Thus far, attempts to reproduce these structures have been unsuccessful
and consequently cytological studies have not been made. Since these
structures have been observed only in a few fixed and stained slide cul-
tures, germination experiments have not been possible.
LIFE CYCLE
The life cycle of C. immitis has always been a problem because of the
great difference between the parasitic and the cultural cycles. The ten-
dency has been to consider that this fungus possesses two distinct life
cycles depending upon the environment (Moore, 1932; Ciferri, 1932:
Dodge, 1935; Ciferri and Redaelli, 1936; Stewart, 1942). According to
this concept, the fungus reproduces in the parasitic state by means of
endogenous spore formation within the spherules and in the saprophytic
state by formation of oidia and chlamydospores.
At the present time culture spherules have been observed on only four
strains, numbers 4, 8, 41, and 46, and constantly on only one strain,
number 4. The factors affecting the production of culture spherules are
not known. According to Baker and Mrak (1941), the culture spherules
did not appear until after active growth of the fungus had ceased because
of drying, “staling products” in the medium or both. In a semi-solid
medium containing copper sulfate, the culture spherules were formed
before drying alone could have become an important factor. However,
it may be that drying increased the concentration of copper to a toxic
level, thereby restricting hyphal development. No definite explanation
was given for the failure of other strains to produce culture spherules
under the conditions described. All strains were pathogenic and morpho-
logically similar. The suggestion was offered that strains not producing
culture spherules had lost this ability or that they were less susceptible
to conditions inducing spherule formation. The behavior of strain num-
ber 41 suggested that the explanation lies in a loss of ability to produce
spherules in culture.
The finding of spherules produced in culture eliminates the need for
- Baker, Mrak & SmitH: Coccipiowes IMMITIS 2h
postulating two distinct life cycles. It is hoped that in the future the
conditions necessary for the production of culture spherules by all strains
will be found. A diagrammatic representation of the life cycle of C.
immitis is given in Plate VIII. When an endospore is placed on arti-
ficial media, it germinates, forming a mycelium composed of septate
hyphae. Later some of the cells of the hyphae are differentiated into
oidia and chlamydospores which are capable of germination on fresh
media to repeat the cycle. In some strains (number 4 in particular) cer-
tain of the hyphal cells enlarge, develop a thick wall and produce endo-
spores by cleavage. These endospores are also capable of germinating
to repeat the cycle. If the chlamydospores and oidia (and probably cul-
ture spherules and endospores) gain access to animal tissues, either by
injection or through the respiratory tract, they undergo a series of im-
perfectly known changes to form mature spherules containing endospores.
According to Chope (see Dickson, 1937b) this process consists of en-
largement of the chlamydospores with the formation of endospores. This
sequence of events is made more probable by the process of culture
spherule formation which occurs in a similar manner. The endospores
formed in mature spherules are capable of either repeating the cycle in
tissues or beginning a new cycle in culture.
In this concept of the life cycle, there is still a remarkable difference
between the cultural cycle and the parasitic cycle. However, certain
analogies and assumptions will aid in making them less divergent. The
essential difference between the complete cultural cycle (with culture
spherule production) and the parasitic cycle is the complete absence of
hyphal development in the latter. If the parasitic condition is considered
as abnormal for the development of the fungus, possibly the development
of the fungus has been reduced to a form which produces the greatest
number of reproductive structures with the least synthesis of new proto-
plasm. In other words, the environmental conditions imposed by animal
tissues completely restrict hyphal development, which is less profitable
from a reproductive standpoint. This is not an uncommon phenomenon
since it occurs in Sporotrichum, Histoplasma, Blastomyces, etc., in animal
tissues and in certain cultural conditions. A similar process also occurs
in Empusa in which the conidia germinate to form a second conidium
when the first conidium is placed in unfavorable environmental condi-
tions (see Fitzpatrick, 1930).
TAXONOMY
From a taxonomic standpoint, C, immitis presents a difficult problem.
Posadas (1892) believed that he was studying a protozoan and discussed
a possible relationship with the Gregarines of the Psorospermia (Sporo-
zoa) but did not arrive at any definite conclusion and did not propose
a name. Wernicke (1892) studied material from the same case but he
212 Fartowia, VoL. 1, 1943
also did not propose a name.* No cultures were made by either Posadas
or Wernicke.
In 1896, Rixford and Gilchrist published the results of a study of the
first two North American cases of coccidioidal granuloma and for the
first time gave a definite name to the organism. They also believed that
they were dealing with a protozoan parasite and after consultation with
the protozoologist, Stiles, described the genus Coccidioides and the two
species, C, immitist and C. pyogenes.
In 1898, at the First Latin-American Medical Congress, Cantén pro-
posed the name Posadasia sphaeriformis. This name is not valid be-
cause it is antedated by C. immitis and because the proceedings of this
Congress were mimeographed but not formally published.
Ophiils (1905) showed that the two species of Rixford and Gilchrist
were identical since the differences could be observed in experimental
animals inoculated with a single strain of the fungus. He rejected the
name C. immitis because Rixford and Gilchrist described a protozoan,
tentatively proposing to name the fungus either Oidium coccidioides or
O. protozoides in spite of the fact that he considered the structures found
in animal tissues to be asci. It must be pointed out that Ophiils’ usage
of Oidium is not comparable to the original description of Oidium.
Verdun (1907) created the combination Oidium immite, for the first
time using the name immitis in a mycologic sense. Brumpt (1910), in
the first edition of his Précis de Parasitologie, used the name C. immitis.
He placed the fungus in the Hyphomycetes and stated that it showed
relationships with certain of the Ascomycetes (Endomyces and Saccharo-
myces).
Hartmann and Schoo (1912), working in Holland, studied an autopsy
specimen from a patient with a disease diagnosed as cancer. Later, Hart-
mann (1912) briefly described Blastosporidium Schooti as the cause of the
disease and placed it in the Haplosporidia. He commented upon the
possibility of its identity with C. immitis. Brumpt (1927 and later edi-
tions) considered Blastosporidium Schooii to be identical with C. immitis.
Unfortunately, however, Hartmann and Schoo failed to obtain cultures.
Hartmann’s figures and description are not entirely typical of C. wnmitis.
It is possible that the two organisms are not identical, a view already
pointed out by Ciferri and Redaelli (1936). In addition to suggesting
‘Tt is frequently stated (Almeida, 1933; Dodge, 1935; Ciferri and Redaelli, 1936;
and others) that Posadas and Wernicke used the generic terms Megalosporidium and
Megalocytosporidium for Coccidioides. However, upon examination of their papers
(Posadas, 1892, 1900; Wernicke, 1892) we were unable to find these names. At the
present time the exact origin of these names has not been determined.
“The description of C. immitis is usually credited to Stiles. However, Rixford
and Gilchrist (1896) published Stiles’ letter containing suggestions regarding the
nomenclature and systematic relationships of this organism. This letter contained no
description of the organism. Rixford and Gilchrist did give a description and as-
sumed full responsibility for it. Thus the correct nomenclature should be Coccidioides
immitis Rixford and Gilchrist 1896 and not C. immitis Stiles as is frequently given.
Baker, Mrak & SmitH: Coccipioies IMMITIS 213
a relationship between the two organisms, Hartmann, for the first time,
suggested that C. immitis is a Phycomycete.
In the second edition of his Précis de Parasitologie, Brumpt (1913)
introduced a new name in the literature, namely: Mycoderma immite.
He made this change in spite of the fact that C. immitis never forms a
yeast-like pellicle, which is the most important characteristic of the “med-
ical” genus Mycoderma as defined by Vuillemin (1931). He also re-
tained this name in the 1927 edition although he returned to the name
C. immitis in subsequent editions.
Although, as stated above, Hartmann first suggested the relationship
of C. immitis with the Phycomycetes, Langeron (1922) is usually credited
with this suggestion since he noted in animal tissues the formation of
endospores by the process of cleavage, and suggested placing this organ-
ism in the Oomycetes near the Chytridiales.
In 1927 Castellani created a new genus Blastomycoides to include
Blastomyces Gilchrist and Stokes, Coccidioides Rixford and Gilchrist,
Oidium Link (in part), Cryptococcus Kiitzing (in part), and Zymonema
Beurmann and Gougerot (in part). In this genus he included three
species: B. immitis, B. dermatitidis, and B. Tulanensis. The first two
species are probably identical. B. Tulanensis has been shown to be
identical with Blastomyces dermatitidis (Martin, 1939) and is entirely
unrelated to C. immitis. Castellani grouped C. immitis with these other
organisms because he considered the endospores to be granules or fat
droplets rather than spores. Castellani and Jacono (1933) discontinued
the use of the name Blastomycoides and placed C. immitis in the genus
Geotrichum, following Agostini (1932).
Mazza and Parodi (1927) published a report of a case of coccidioidal
granuloma of the larynx. They considered the organism to be identical
with that of Posadas and Wernicke but did not make cultures. In the fol-
lowing year Fonseca (1928), after studying the sections from the case
of Mazza and Parodi, established a new genus and species Pseudococcidi-
oides Mazzai for the parasite. He suggested that both Pseudococcidioides
and Coccidioides are related to the genus Protomyces. In a previous
paper (Fonseca and Ledo, 1927) a relationship between Coccidioides and
Protomyces is mentioned, with the suggestion that a search be made for
C. immitis in this group of plant parasites.
Almeida (1932b,c, 1934a) after comparing material from the case of
Mazza and Parodi with material from cases of coccidioidal granuloma
(human and experimental) concluded that the genus Pseudococcidioides
should be reduced to synonomy with Coccidioides. He pointed out that
Posadas (1892) and Rixford and Gilchrist (1896) had figured forms of
C. immitis identical with Pseudococcidioides Mazzai. In fact, Rixford
and Gilchrist’s description of C. pyogenes bears a great resemblance to
Fonseca’s description of Pseudococcidioides Mazzai.
Until 1929 it was generally considered that C. immitis occurred in
214 FarRLowl1A, VoL. 1, 1943
Brazil. Almeida (1929, 1930a,b) made a comparative study of C. immitis
from the United States and of the supposed C. immitis from Brazil and
showed that they are not identical. He established the genus Paracoc-
cidioides for the Brazilian fungus with the species, braziliensis. For a
complete discussion of this fungus see Almeida (1933), Redaelli and
Ciferri (1937), and Moore (1938).
Moore (1932) concluded that the spherules found in tissue are asci
and created the family Coccidioideaceae in the Endomycetales. He
recognized the species C. immitis for the North American fungus and
returned the Brazilian fungus to this genus with the species name esferi-
formis originally proposed by Cantén (1898) for C. immitis (see Ciferri
and Redaelli, 1936). Later Moore (1935) included in this family Pos-
adasia (Canton) Moore (an invalid name for Histoplasma Darling (see
Ciferri and Redaelli, 1936), Paracoccidioides Almeida, and Rhinospor-
idium Minchin and Fantham. Ciferri (1932) independently of Moore,
established the family Coccidioideaceae, placing it in the Chytridiales
and including in it Coccidioides Rixford and Gilchrist and Paracoccidi-
oides Almeida.
Ota and Kawatsure (1933) published a study of fungi causing blasto-
mycosis. Included in this group of fungi were C. immitis and an organ-
ism previously described by Ota (1926; in Japanese) as Scopulariopsis
americana Ota. Ciferri and Redaelli (1936) studied one of the four
cultures of this fungus (stated by Ota and Kawatsure to be identical) and
found it to be a slightly atypical culture of C. immitis.
Dodge (1935) followed Moore in placing Coccidioides in the family
Coccidioideaceae Moore, but reduced Coccidioides esferiformis to syn-
onomy with C. immitis.
In 1936 Ciferri and Redaelli published an extensive study of fifteen
strains of C. immitis. They reaffirmed Ciferri’s (1932) belief of a rela-
tionship with the Chytridiales. They removed Paracoccidioides from
the family Coccidioideaceae and added Rhinosporidium, at the same time
discussing the relationships between Coccidioides and various Chytridiales
(many of which Fitzpatrick, 1930, considered as doubtful chytrids) and
proposing a possible phylogenetic scheme. They recognized the single
species C. immitis with the three varieties: typicus, metaewropeus and
Pipkini.
It is apparent that there has been considerable controversy regarding
the nature of the spherule. Most investigators have described and figured
spore formation by the process of cleavage. The present study indicates
that the spores both of tissue and culture spherules are formed by cleavage
and not by “free cell” formation. In the opinion of the authors, there is
little doubt but that the spherules are sporangia and not asci.
In view of this formation of endospores by cleavage, C. immitis cannot
be placed in the Ascomycetes despite its septate mycelium. Neither.
basidiospores nor clamp-connections have been observed by previous in-
Baker, Mrak & SmiTH: CoccipIoIDEs IMMITIS 215
vestigators or by the authors. Since these structures are the chief basis
for the differentiation of the Basidiomycetes, it appears impossible to in-
clude C. immitis in this group of fungi. Inclusion of this fungus in the
Fungi imperfecti has been the proposal of various investigators, notably:
Castellani (1927), Agostini (1932), Castellani and Jacono (1933), and
Ota and Kawatsure (1933). An attempt was made to classify C. immitis
in this group using the keys of Clements and Shear (1931). Since no
conidia (except oidia-like structures which probably represent a stage
in the formation of chlamydospores) are formed, the only possible rela-
tionship would be with the Mycelia sterilia. However, fungi in this group
possess no reproductive structures of any kind, whereas C. immitis forms
chlamydospores. Also, the production of sporangia in the host tissues
and occasionally in culture would eliminate any relationship with this
group, the members of which are generally considered to be imperfect:
representatives of the Ascomycetes. Thus, it is impossible to classify
C. immitis in the Fungi imperfecti.
The most probable relationship is with the Phycomycetes because of
the formation of the sporangium. However, C. immitis presents certain
characteristics inconsistent with such a relationship. For example, the
mycelium is septate from the initiation of hyphal development, a char-
acteristic of the Ascomycetes and Basidiomycetes. Sporangia are rare
in culture and are not borne on erect sporangiophores as is characteristic
of the terrestrial Phycomycetes.
The exact systematic position of C. ummitis among the Phycomycetes
presents further difficulties. Since C. immitis has a well developed
mycelium and grows readily on very simple media, relationship with the
higher Oomycetes or Zygomycetes might be expected. However, the
sporangium is not typical of the Oomycetes or Zygomycetes. There are
no definite sporangiophores, a condition not found above the level of the
Pythiaceae of the Peronosporales. In culture, the sporangia are terminal
or intercalary, solitary, or in chains. Again this condition is not found
above the level of the Pythiaceae and not at all in the Zygomycetes. The
sporangium is thick-walled, a condition not found in either the Oomycetes
or Zygomycetes, although the upper half of the sporangium in both
Pilobolus and Pilaira is thickened (Fitzpatrick, 1930).
If only the parasitic cycle of C. immitis is considered, this fungus is
reminiscent of the Chytridiales in that the entire thallus is transformed
into a single reproductive structure. Ciferri and Redaelli (1936) have
attempted to trace a relationship with this group. However, there are
certain objections to this view. C. immitis has a highly developed, sep-
tate mycelium and grows well on very simple media. In contrast, the
Chytridiales produce motile zoospores and are, for the most part, obli-
gate parasites, with the mycelium, when present, evanescent. Ciferri and
Redaelli point out that certain members of the Chytridiales (Catenaria,
Hypochytrium) have septate mycelia. While Catenaria does have a
216 FartowiA, VoL. 1, 1943
primitive mycelium, it is by no means comparable to that of C. immitis.
Hy pochytrium has a well developed mycelium, but Fitzpatrick (1930) ex-
presses some doubt as to its relationship with the Chytridiales. Ciferri
and Redaelli did not observe motile spores and are of the opinion that
because of the environment (animal tissues) C. immitis has lost the ability
to produce motile zoospores. However, the endospores produced by
C. immitis in culture are also aplanetic. They state that Hypochytrium
infestans produces zoospores, some of which lose their flagella before
emergence. However, none of the Chytridiales form only aplanetic
zoospores. Most of them are obligate parasites or at least have not
been cultured. Recently Karling (1934) has cultured Catenaria An-
guillulae on boiled fluke eggs and vegetables. Macrochytrium botrydioides
is saphrophytic on rotten fruit in water according to Minden’s (1902)
description. According to Fitzpatrick (1930) the latter fungus is a
doubtful member of the Chytridiales. In contrast to the predominately
obligate parasitism and simple structure of the great majority of the
Chytridiales, C. immitis is not an obligate parasite and produces a com-
plex thallus on very simple media. Thus, it is difficult to accept a rela-
tionship with the Chytridiales.
Fonseca (1928) described Pseudococcidoides Mazzai, now known to be
identical with C. immitis, and made the following statement (translated)
concerning its systematic position:
“Pseudococcidioides mazzai is probably best placed in the family Protomycetaceae
in the Hemiascomycetes which are transitional between the Phycomycetes and the
Ascomycetes. It is near Coccidioides from which it is distinguished primarily by
the mode of formation of spores which separate in groups within the membrane
of the mother cell.”
Fonseca based the separation of Pseudococcidioides from Coccidioides
on the formation of endospores in the former by cleavage of the cyto-
plasm arranged around the periphery of the spherule. He did not study
cultures of this organism because none were obtained. Almeida (1932 b,c,
1934a) pointed out that this type of sporulation also occurs in C. immitis.
Rixford and Gilchrist (1896), Ophiils (1905), and others as well as
the authors, have also observed this type of sporulation in C. immitis.
It was this cleavage of peripherally arranged cytoplasm that led Fonseca
to suggest a relationship with Protomyces.
Biiren (1915) has shown that in Protomyces the delicate intercellular,
septate mycelium bears large, globose to ellipsoidal, thick-walled, uni-
cellular chlamydospores which may be either terminal or intercalary.
After a period of rest, chlamydospores germinate by rupture of the thick
exospore and the extrusion of the thin endospore as a globose or cylin-
drical sack. When the sack is fully formed, it contains a peripheral
multinucleate layer of protoplasm which is soon divided by cleavage
planes into many small uninucleate segments. The nuclei in these seg-
ments divide twice, forming four nuclei, and eventually four uninucleate
Baker, Mrak & SMITH: CoccipioiDes IMMITIS 217
spores. Finally these spore tetrads break up and the spores aggregate
in a clump near the tip of the extruded sack. After release, the spores
copulate in pairs. The nuclei migrate into the copulation bridge but it
is uncertain whether or not they fuse. When these spores reach the
epidermis of the host, they put out germ tubes which penetrate between
the epidermal cells, and repeat the cycle. The exact position of meiosis
in the life cycle is not clear.
The question of a possible relationship between the genera Protomyces
and Coccidioides appears to lie in the mode of formation of endospores
and their subsequent development. In Coccidioides the spores are un-
doubtedly sporangiospores. At the present time the life cycle of Pro-
tomyces is not clear. Biiren considers the cells cut out of the peripheral
cytoplasm to be asci and the four spores formed in them to be ascospores.
He terms the sack extruded from the chlamydospores a “synascus.” He
believes that meiosis probably occurs in the formation of the spore tetrads.
Gaiimann and Dodge (1928) offer another possible interpretation as
follows:
“If one wishes to avoid this explanation (meiosis during formation of spore
tetrads), one must assume that the quartering of the spore mother cells does not
have the significance of a meiosis and that the spore mother cells, like the proto-
spores of the Mucoraceae and Synchytriaceae, represent only an intermediate stage
not connected with a change of nuclear phase. Meiosis then, as in the Dipodascaceae
and Endomycetaceae, would be sought in connection with the nuclear fusion in the
beginning of nuclear divisions at the emergence of the endospore. Like the ascus
of Dipodascus, the endospore would correspond to an ordinary ascus with still
definite sporangial character, but encysted like the zeugites of many parasitic
Basidiomycetes.”
In both of these interpretations, the spores are assumed to be ascospores
and therefore could not correspond to the endospores of C. immitis. On
the other hand, Gwynne-Vaughn and Barnes (1937) interpret the chla-
mydospore of Protomyces as a resting sporangium and place this genus
in the order Protomycetales near the Chytridiales and Ancylistales.
Fitzpatrick (1930) summarizes the situation as follows:
“The position of the family Protomycetaceae in the natural system is in doubt,
any disposition of it being to an extent dependent upon the interpretation given
to the structures and phenomena just described. Unless the extruded endospore
be regarded as a many-spored ascus there would seem to be no reason for incorpo-
rating these forms in the Ascomycetes.”
In addition to the genus Protomyces, the family Protomycetaceae con-
tains two other genera: (see Gatimann and Dodge, 1928) Protomycopsis
and Taphridium. In both Protomyces and Protomycopsis the spores
are delimited in the extruded sack but the spores do not copulate in Pro-
tomycopsis as they do in Protomyces. In Taphridium the spores are
delimited in the thick-walled chlamydospore before extrusion of the endo-
spore. In at least one species the spores have been described as fusing
in pairs after their escape.
218 FarLowia, VoL. 1, 1943
If C. immitis is compared with these three genera, certain similarities
as well as differences can be seen. In Protomyces and Protomycopsis
the spores are delimited in the extruded sack while the spores in C. immitis
and Taphridium are delimited within the original chlamydospore.°
Taphridium differs from C. immitis in that the spores of Taphridium pass
into an extruded sack before release while the spores of C. immitis are
released directly by rupture of the sporangium. The released spores of
Protomyces and some species of Taphridium copulate in pairs while those
of C. immitis and Protomycopsis do not copulate (unless the observations
of Ciferri and Redaelli (1936), of a fusion of cells before sporulation in
a strain of C. immitis, is confirmed). C. immitis differs from the other
genera in several important respects. C. immitis grows readily on simple
media while the other genera are obligate parasites on higher plants or at
least have not been cultured. C. immitis produces small cylindrical
chlamydospores which are incapable of functioning as sporangia in cul-
ture but which are probably able to do so in animal tissues.
On the basis of present information, it appears possible to suggest a
relationship with the three genera, Protomyces, Protomycopsis, and
Taphridium, even though members of these genera have not yet been
cultured. C. immitis may even be included in the family Protomycetaceae
although this would be unwise without more information regarding the
details of the life histories of these four genera. Moore (1932) estab-
lished the family Coccidioidiaceae placing it in the Endomycetales.
Ciferri and Redaelli (1936) amended the description of this family and
placed it, as well as the family Protomycetaceae, in the Chytridiales. No
attempt will be made at the present time to further amend the description
of the family Coccidioidiaceae or to merge it with the Protomycetaceae
although this latter change may be desirable in the light of future data.
The following statement from Fitzpatrick (1930) probably best sum-
marizes any attempt at an exact taxonomic treatment of either the family
Protomycetaceae or C. immitis:
“as treated here the Protomycetaceae are regarded as of unknown relationship.
They may represent an isolated line of development with a phycomycetous an-
cestry,...”
Emmons and Ashburn (1942) have recently described a new species of
fungus, Haplosporangium parvum, recovered from several genera of wild
rodents in Arizona. C. immitis was also isolated from the same genera
of rodents and in some cases both fungi were obtained from one animal.
From their description no morphological similarities are apparent be-
tween H. parvum and C. immitis. An antigen prepared from H. parvum
produced more positive skin reactions in individuals giving positive
coccidioidin skin tests than in individuals giving negative coccidioidin
® The term chlamydospore has been used to designate the structure in which the
spores of the various Protomycetaceae are formed. In the present discussion this
term is also applied to the spherule of C. immitis for comparison.
Baker, Mrak & SmitH: CoccipioIDEs ImMMITIS 219
tests. On the basis of this, speculation is made of a possible genetic rela-
tionship between H. parvum and C. immitis. While such a relationship
may be possible, more evidence is needed, particularly in view of known
cross-serological reactions between unrelated - organisms.
On the basis of available information the genus Coccidioides may be
defined as follows:
Genus Coccidioides Rixford and Gilchrist 1896; emend 1943.
Synonomy: Posadasia Canton 1898
Oidium of Ophiils, 1905, not Link.
Blastosporidium Hartmann 1912 (?)
Mycoderma (in part) of Brumpt, 1913, not Persoon
Blastomycoides Castellani, 1927 or 1926
Pseudococcidioides Fonseca, 1928
Geotrichum (in part) of Basgal, 1931, not Link
Scopulariopsis (in part) of Ota, 1928, not Bainier
Glenospora of Castellani and Jacono, 1933, not Berkeley and Curtis
Colonies on solid media circular, raised, or flat, with or without a
prominent central boss; surface cottony, powdery, or smooth and mem-
branous; color white becoming brown with age. Thallus in animal
tissues reduced to a single cell; in culture branched and well developed;
hyphae absent in animal tissues; in culture septate at all stages of de-
velopment, 2-4 » in diameter. Vegetative reproduction in animal tissues
by formation of holocarpic thick-walled sporangia (termed spherules),
up to 80 » in diameter, number of sporangiospores variable, few to sev-
eral hundred, thin or thick-walled, spherical to globose or irregular in
shape, 2-3x 2-3 p, occasionally up to 5-10x 5-10 p», formed by suc-
cessive cleavage planes; in culture sporangia rarely formed, when present
terminal or intercalary, thick-walled, spherical to globose, ranging up to
20 » in diameter, number of sporangiospores variable, fewer than in tissue
sporangia, sporangiospores usually thin-walled, occasionally thick-walled,
irregular, spherical or globose 2—3 x 2-3 p; oidia globose, spherical or
cylindrical 2-4.x 2-6 p, occasionally up to 20 » in diameter. Sexual
reproduction not definitely established.
Type species: C. immitis Rixford and Gilchrist 1896.
Most investigators have made no attempt to differentiate species within
the genus Coccidioides. However, Ciferri and Redaelli (1936) estab-
lished three varieties: C. immitis var. typicus; C. immitis var. Pipkini;
and C. immitis var. meteuropeus. The first variety represents the usual
form of the fungus. The second variety is said to form (translated)
“| .. a morphological complex with forms similar to those of Scopulari-
opsis; an atypical experimental histopathological picture in the guinea
pig.” An examination of a subculture (number 17) from this strain
showed this culture to be similar to all of the other strains studied. The
third variety was based upon the geographical location of its isolation,
Naples, Italy.
The advisability of describing culture number 4 as a new species was
220 Fartowia, Vou. 1, 1943
considered. However, in spite of the fact that it forms sporangia regu-
larly in culture while the other strains form sporangia rarely or not at
all, it seems inadvisable to describe an additional species. While the
various strains show considerable and constant differences, there are no
well differentiated colony types. Instead, the fifteen strains appear to
form an integrated series, each strain differing a little from the other.
Various physiological properties to be reported later were similar for
all strains. The ability to utilize various carbon and nitrogen compounds
is similar for all strains (Baker and Smith, 1942). The effect of pH on
growth is also similar. Coccidioidin tests made by Smith and Baker (un-
published data) on two sensitive individuals using antigens prepared
from all of the fifteen strains gave identical reactions to the various anti-
gens. Thus, in the opinion of the authors, there is no justification for
the description of additional species or varieties of Coccidioides on the
basis of the present information, although it is possible that in the future
they may be discovered.
On the basis of available information, the synonymy of Coccidioides
immitis Rixford and Gilchrist 1896 follows:
Synonomy: Coccidioides pyogenes Rixford and Gilchrist 1896
Posadasia esferiformis Canton 1898
Oidium protozoides Ophils 1905
Oidium coccidioides Ophiils 1905
Oidium pyogenes Verdun 1907
Oidium immite Verdun 1907
Blastosporidium Schooti Hartmann 1912 (?)
Zymonema immitis Mello and Fernandes 1918
Mycoderma immite Brumpt 1913 Second Edition
Blastomycoides immitis Castellani 1927 or 1926
Pseudococcidioides Mazzai Fonseca 1928
Scopulariopsis americana Ota 1926
Geotrichum dermatitidis Basgal 1931
Geotrichum immite Agostini 1932
Geotrichum louisianoideum Castellani, in Castellani and Jacono 1933
Glenospora metaeuropea Castellani, in Castellani and Jacono 1933
Coccidioides immitis var. typicus Ciferri and Redaelli 1936
Coccidioides immitis var. Pipkini Ciferri and Redaelli 1936
Coccidioides immitis var. meteuropeus Ciferri and Redaelli 1936
Characteristics are those of the genus.
DISTRIBUTION
Most investigators have assumed that C. immitis develops in soil and
is distributed in dust. Up to the present time, the fungus has been iso-
lated from soil in three different areas as will be mentioned below. How-
ever, the distribution of this fungus must be studied chiefly from the oc-
currence of reported cases of coccidioidomycosis and from the results of
skin testing with coccidioidin testing (see Smith, 1940) for justification
of this type of data.
Baker, Mrak & SMITH: CoccIDIOIDES IMMITIS yvalh
Although the first case of coccidioidomycosis (coccidioidal granuloma)
was reported by Posadas (1892, 1900) and Wernicke (1892) in Argen-
tina, practically all of the subsequent cases have been recorded in the
United States, particularly in the southern San Joaquin Valley region of
California. Beck (1931) summarized the distribution of 286 cases of
coccidioidal granuloma occurring prior to June 1, 1931.
California
The distribution of the fungus in California has now been fairly well
established. Stewart and Meyer (1932) reported isolation of the fungus
from soil collected near Delano in the San Joaquin Valley. Smith and
Baker (1941) isolated C. tmmitis from two soil samples collected in San
Benito County located in the eastern part of the coast range of mountains
bordering the San Joaquin Valley. The lower San Joaquin Valley has
long been considered an endemic area. Rixford’s (1894) first case of
coccidioidal granuloma originated in that area and subsequent investiga-
tions of Gifford (1936, 1942), Gifford, Buss and Douds (1937), Dickson
(1937a,b, 1938a,b), Dickson and Gifford (1938) and Smith (1940) have
provided confirmatory evidence. They studied the acute primary type
of infection which has an incubation period of about 8-21 days. Conse-
quently, the area in which the infections were acquired can be fixed with
considerable certainty.
A rather large number of infections have been acquired south of the
Tehachapi Mountains. Kessel (1941) stated that 31 of 100 patients hav-
ing coccidioidal granuloma, seen at the Los Angeles County Hospital, ac-
quired their infections in Southern California. Three infections are
known to have occurred in San Diego County. Two infections definitely
occurred in Riverside County (Thurber, 1929; Kessel, 1941).
In general, fewer cases are reported from the south coastal counties.
Schwalenberg (1929) reported an infection acquired near Santa Barbara.
Cases have also been recognized in Ventura County. Shelton (1942)
reported several cases from Camp Roberts located in Monterey and San
Luis Obispo Counties. In addition he performed coccidioidin tests on
888 newly arrived troops and retested the nonreactors three months later.
During the three-month period, fourteen of this group had become. in-
fected as indicated by their having acquired sensitivity to coccidioidin.
Davis, Smith and Smith (1942) reported an epidemic of acute primary
coccidioidomycosis resulting from an exposure to dust in San Benito
County.
There have occurred in Contra Costa and Yolo Counties a few isolated
cases of primary coccidioidomycosis which apparently originated in these
areas. However, C. immitis appears to be practically absent from the
northern part of the San Joaquin Valley and the Sacramento Valley as
well as the northern part of the state and the north coast range and the
Sierra Nevada regions.
223 FarLtowia, VoL. 1, 1943
Arizona
Emmons (1942) isolated C. immitis from five of one hundred and fifty
soil samples collected in the desert near the village of San Carlos. He
was also able to isolate cultures of this fungus from rodents trapped in
this same area. Phillips (1938), Woolley (1938), Brown (1939), Far-
- ness and Mills (1939), and Farness (1941) have reported cases of both
primary coccidioidomycosis and coccidioidal granuloma originating near
Tucson and Phoenix. Aronson, Saylor and Parr (1942) have shown a
high incidence of positive reactions to coccidioidin among Indian school
children on certain Indian Agencies near Phoenix and Globe.
Texas
Caldwell (1932, 1942), Smith and Waite (1934), Smith (1942), and
Lehmann and Pipkin (1935) reported cases of coccidioidal granuloma
from Texas in which the infections had been acquired in that state.
Schulze (1942) has reported the presence of an endemic area in western
Texas as indicated by the occurrence of cases of primary coccidioido-
mycosis.
United States other than California, Arizona, and Texas
Burkhead (1922), Van Cleve (1936), and Duckett and Fredeen (1936)
reported cases of coccidioidal granuloma in Kansas; however, careful
residence histories were not obtained. Duckett and Fredeen’s patient, a
boy of five, had lived in Texas and may well have acquired the infec-
tion there.
Stiles, Shahan and Davis (1933) recorded a case of coccidioidal granu-
loma in a heifer born and raised in Colorado but a complete residence
history and the origin of the feed was not available.
Lynch (1920) reported a case of coccidioidal granuloma from Charles-
ton, South Carolina. The patient, a Negress, had lived her entire life on
an island off the coast of South Carolina and in Charleston. However,
the fact that no additional cases have been recorded in this area suggests
that she may have acquired the infection from contaminated material
originating in California, Arizona, or Texas.
McDonald (1934) reported a case of coccidioidal granuloma from
Louisiana. She stated that the patient had spent his entire life in that’
state except for nine months of World War service, four months of which
was overseas. However, she did not state where the other five months
of service were spent.
Our discussion has included only reports of cases which might provide
information regarding the distribution of C. immitis in nature. While
these cases would indicate that C. tmmitis is fairly widely distributed in
the United States, the residence histories are not adequate and additional
cases must be proven before other endemic areas can be established.
Schenken and Palik (1942) have summarized much of the literature on
cases reported in areas of the United States other than California.
BAKER, Mrak & SmiITH: CoccipiomeEs [MMITIS pps)
South America
Four cases of coccidioidal granuloma have been recorded in South
America. Posadas (1892) and Mazza and Parodi (1927) reported two
cases and Nifo (1938) summarized the information on two additional
cases, all of which occurred in the Chaco region of Argentina. Cases of
coccidioidal granuloma reported from Brazil are paracoccidioidal granu-
loma (see p. 214).
Europe
Hartmann and Schoo (1912), working in Holland, published an autopsy
report on a patient said to have died of cancer, who may have had coccidi-
oidal granuloma., If this case could be verified, it would be the first
case to be reported from Europe. Ciferri and Redaelli (1936) and the
authors consider the diagnosis of coccidioidal granuloma open to ques-
tion. Redaelli and Ciferri (1934) have reviewed the data on the Eu-
ropean cases. They mentioned three cases occurring in Naples, Italy.
The fungus isolated from one of these cases was described as Glenospora
meteuropea by Castellani and Jacono (1933), but was shown by Redaelli
and Ciferri (1934) and Ciferri and Redaelli (1936) to be C. immitis.
They also mentioned another patient from the Balkans from which Cas-
tellani and Jacono isolated Glenospora meteuropea but indicated that
this culture was not C. immitis. In 1939 we received a culture from
Redaelli labeled: “Coccidioides immitis var. meteuropeus (= Glenospora
metaeuropeus Castellani) (culture originally from Castellani, of Balkan
origin).” This culture (our number 28) is definitely C. immitis, but
whether it is a subculture of the Balkan culture of Glenospora metaeur-
opeus, which Ciferri and Redaelli stated is not C. immitis, is of course,
uncertain. Redaelli and Ciferri also mentioned a case originating in
Hamburg, Germany, but stated that the diagnosis is doubtful.
Other Areas
Redaelli and Ciferri discussed a patient from Australia who appeared
to have acquired the infection from California vegetables. They pointed
out that this case needs verification. They gave no references to the
original reports of the Hamburg and Australian cases and we have been
unable to find any reference to them in the literature.
Fennel (1935) reported a case in Hawaii. The patient, a full-blooded
Hawaiian, had never been out of the islands. The author discussed the
possibility of the introduction of the fungus into this area by means of
fresh vegetables from California. Additional cases will have to be re-
corded in Hawaii before it can be considered an endemic area.
There are a few reports of cases of coccidioidal granuloma from other
areas, but a critical examination of the data makes it apparent that the
investigators were mistaken in their identification of the organism. For
example, Chou and Reiss (1931) reported a case of coccidioidal endo-
ophthalmitis from China. Examination of the figures makes it quite
224 Fartowia, Vou. 1, 1943
evident that the investigators were dealing with some other fungus.
Burgess (1929) reported coccidioidal granuloma in a patient from Mon-
treal, Canada, whose infection he attributed to handling fresh vegetables
from California. However, he was unable to obtain cultures, and his
description of the clinical aspects and of the parasite are not convincing.
The distribution of C. immitis (and coccidioidomycosis) may be sum-
marized as follows: Endemic areas occur in the southern half of Cali-
fornia, particularly in the San Joaquin Valley, and in certain parts of
Arizona and Texas. Possibly other endemic areas exist in the United
States, but the evidence is insufficient for any definite conclusions to be
made. Outside of the United States it seems likely that an endemic area
exists in South America (Argentina) and possibly in Italy (Naples).
SUMMARY
The gross morphology of fifteen strains of Coccidioides immitis Rixford
and Gilchrist 1896 was studied in single spore culture on potato dextrose,
Sabouraud’s and synthetic agars. Cultures of single strains on a given
medium were quite constant but there was considerable difference in
the gross morphology of the same strains on the three media. There was
considerable variation in the gross morphology of the different strains
when cultured on the same medium. Incubation of cultures in either light
or dark had no effect on the gross morphology.
The microscopic morphology of the same strains was studied both in
animal tissues and in culture. There was remarkable similarity in the
morphology in animal tissues of all of the strains. In culture the sim-
ilarity was somewhat less but the differences are quantitative rather than
qualitative and are not considered of taxonomic significance. A discus-
sion of the life cycle of this fungus was given. .
The taxonomic position of the genus Coccidioides Rixford and Gilchrist
1896 was discussed and a description given on the basis of present in-
formation. It was suggested that this fungus is a Phycomycete and may
be related to members of the Protomycetaceae. It was concluded that
the genus is represented by the single species, Coccidioides immitis Rix-
ford and Gilchrist 1896.
The distribution of Coccidioides immitis Rixford and Gilchrist 1896
was discussed. It was concluded that endemic areas exist in California,
Arizona, and Texas in the United States, the Chaco region of Argentina
and possibly in Italy.
Baker, Mrak & SMITH: CocciIDIOIDEs IMMITIS 220
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trophic). Arch. Dermatol. and Syphilol. 31: 586-588.
Lewis, G. M. and Hopper, M. E. 1939. An introduction to medical mycology.
315 pp. Year Book Publ., Chicago.
Lynch, K. M. 1920. Coccidioidal granuloma, including the first reported case east
of the Mississippi. South. Med. Jour. 13: 246-249.
McDonald, C. 1934. A study of Coccidioides immitis. Jour. Lab. and Clin. Med.
20: 2-8.
MacNeal, W. J., and Taylor, R. M. 1914. Coccidioides immitis and coccidioidal
granuloma. Jour. Med. Res. 30: 261-274.
Martin, D. S. 1939. Serologic diagnosis of fungus infections. Proc. Third Internat.
. Cong. for Microbiol., New York, p. 516-517.
Mazza, S. and Parodi, S. E. 1927. Micosis laringea con pardsitos andlogos
a los “megasporideos” de Posadas. Bol. Inst. Clin. Quir. 3, No. 26. Cited by
Nino (1938).
Minden, M. von. 1902. Ueber Saprolegniineen. Centr. Bakt. 2 Abt. 8: 821-825.
Moore, M. 1932. Coccidioidal granuloma: a classification of the causative agent,
Coccidioides immitis. Ann. Missouri Bot. Gard. 19: 397-428.
. 1935. A morphological and physiological study of two species of Posadasia,
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1938. Blastomycosis, coccidioidal granuloma and paracoccidioidal granu-
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organisms and clinical types. Arch. Dermatol. and Syphilol. 38: 163-190.
228 FarLowl1A, VoL. 1, 1943
Nino, F. L. 1938. Contribucion al estudio de las blastomicosis en la republica
Argentina. Boll. Inst. Clin. Quirurg. 14: 591-1042.
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149-172.
Phillips, E. W. 1938. Presence of coccidioidal infection in Phoenix. Southwest.
Med. 23: 48-51.
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156 pp. Wiley, New York. 4
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Schultze, V. E. 1942. Acute coccidioidomycosis in West Texas. Texas State Jour.
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Schwalenberg, 1929. Cited by Beck (1931).
Shelton, R. M. 1942. A survey of coccidioidomycosis at Camp Roberts, California.
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Snyder, W. C. and Hansen, H. N. 1941. The effect of light on taxonomic char-
acters in Fusarium. Mycologia 33: 580-591.
Stewart, R. A. 1942. The biologic challenge of Coccidioides immitis. Proc. Sixth
Pacific Sci. Congr. (Univ. of Calif. Press) 5: 811-813.
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Dis. 66: 212-217.
and Meyer, K. F. 1932. Isolation of Coccidioides immitis (Stiles) from
the soil. Proc. Soc. Exp. Biol. and Med. 29: 937-938.
BakER, Mrak & SMITH: CoccipIoweEs IMMITIS 229
Stiles, G. W., Shahan, M. S. and Davis, C. L. 1933. Coccidioidal granuloma in
cattle in Colorado. Jour. Am. Vet. Med. Assoc. 82: 928-930.
Takahashi, S. 1933. Experimentalle Untersuchungen tiber Coccidioides immitis.
Arch. Dermat. u. Syphil. 168: 597-610.
Thurber. 1929. Cited by Beck (1931).
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-37: 54-55.
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Wernicke, R. 1892. Ueber einen Protozoenbefund bei Mycosis fungoides (?)
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Wolbach, S. B. 1904. The life cycle of the organism of “dermatitis coccidioides.”
Jour. Med. Res. 13: 53-60.
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Yarwood, C. E. 1937. The relation of light to the diurnal cycle of sporulation of
certain downy mildews. Jour. Agric. Res. 54: 365-373.
230 FarLowia, VoL. 1, 1943
EXPLANATION OF PLATE I.
Coccidioides immitis. Photographs of fifteen strains.
Fig. 1, Cultures on potato dextrose agar.
Fig. 2. Cultures on Sabouraud’s agar.
Baker, Mrak & SmiITH: CoccipioIDEs IMMITIS 231
Priate I,
232
FarLowiaA, VoL. 1, 1943
EXPLANATION OF PLATE II.
Coccidioides immitis. With the exception of figure 1, all figures are of strain 46
and are shown at a magnification of x 495.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
1.
2.
3.
4.
5.
8.
9.
Photograph of fifteen strains cultured on synthetic agar.
Mature spherule containing endospores. Section of guinea pig testis stained
with Harris’ hematoxylin.
Small spherule within a histiocyte. Section of guinea pig testis stained with
Harris’ hematoxylin.
Spherule with large vacuole. Macerated mouse lung mounted in cotton
blue lacto-phenol.
Spherule showing first cleavage plane. Macerated mouse lung mounted in
cotton blue lacto-phenol.
Spherule showing early stages of cleavage. Macerated mouse lung mounted
in cotton blue lacto-phenol.
Spherule showing early stages of cleavage. Macerated mouse Jung mounted
in cotton blue lacto-phenol.
Large spherule (about 80 uy. in diameter) in which no spores have formed.
Macerated mouse Jung mounted in cotton blue lacto-phenol.
Spherule with peripherally arranged protoplasm cut by radial cleavage planes.
Macerated mouse lung mounted in cotton blue lacto-phenol.
10. Large spherule with many small, irregular spores. Macerated mouse lung
mounted in cotton blue lJacto-phenol.
11. Spherule with large eccentric vacuole. Macerated mouse lung mounted in
cotton blue lacto-phenol. '
Baker, Mrak & SmitH: CoccipiomwEs IMMITIS 233
Prate II.
234
Fartowia, VoL. 1, 1943
EXPLANATION OF PLATE IIL.
Coccidioides immitis. Figures 1 and 9-18 are of strain 46, figures 3-8 are of strain
40, and figure 19 is of strain 4; all are at a magnification of 495.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
2
Zz;
3.
4,
Spherule with large endospores. Macerated mouse lung mounted in cotton
blue lacto-phenol.
Club-like processes radiating from the cell wall. Section of a mediastinal
lymph node of a cow stained with Harris’ hematoxylin.
Dumbbell-shaped structure resembling conjugation. Pus from guinea pig
testis mounted in cotton blue lacto-phenol.
Two spherules apparently joined, resembling conjugation. Note the ap-
parent evagination into one spherule. Pus from guinea pig testis mounted
in cotton blue lacto-phenol.
Spherule with three large endospores. Pus from. guinea testis mounted in
cotton blue lacto-phenol.
Three spherules, two containing endospores, joined by short tubes suggestive
of conjugation. Two spherules, one containing endospores, joined by a
tube suggestive of conjugation. Pus from guinea pig testis mounted in
cotton blue lacto-phenol.
Large spherule containing three large endospores joined to a smaller spherule
by a narrow tube. Suggestive of conjugation. Pus from guinea pig testis
mounted in cotton blue lacto-phenol.
Two spherules containing endospores joined and with an opening between
them. Pus from guinea pig testis mounted in cotton blue lacto-phenol.
Spherule with fine spine-like excrescences radiating from the cell wall.
Section of guinea pig testis stained with Harris’ hematoxylin.
- Dumbbell-shaped structures and cells joined by a tube. From an old cul-
ture. Mounted in cotton blue lacto-phenol.
. Germinating endospore showing first branch. Unstained.
. Germinating endospore. Unstained.
. Germinating endospore showing branching and septation of germ tube. Un-
stained.
. Endospores in a large spherule germinating through the spherule wall. Un-
stained.
. Mycelium in a four day old slide culture. Mounted in cotton blue lacto-phenol.
. Mycelium in a four day old slide culture. Mounted in cotton blue lacto-phenol.
. Clavate cells from a one month old culture. Mounted in cotton blue
lacto-phenol.
. Chain of chlamydospores interspaced by empty hyphal spaces. From one
month old culture. Mounted in cotton blue lacto-phenol.
. Spindle-shaped chlamydospores from a one month old slide culture. Stained
in 1% aqueous cotton blue.
Baker, Mrak & SmiTH: CoccipDIoIDEs IMMITIS 235
Puate III,
236
FarRLowIA, VoL. 1, 1943
EXPLANATION OF PLATE IV.
Coccidioides immitis. Figures 1-3 and 8-12 are of strain 4; figures 4-7 and 13-14
are of strain 46. Figures 1-12 are shown at a magnification of 495, 13-14 at 1800.
Fig. 1. Chain of chlamydospores interspaced by empty hyphal spaces, From a one
Fig.
Fig.
Fig.
2.
16.
, 44.
month old slide culture. Stained in 1% aqueous cotton blue. %
Chain of chlamydospores containing both spherical and cylindrical spores
from a one month old slide culture. Stained in 1% aqueous cotton blue.
Spherical chlamydospores from a one month old slide culture. Stained in
1% aqueous cotton blue.
Chain of oidia from a month old slide culture. Mounted in cotton blue
lacto-phenol.
Germinating chlamydospore. Unstained.
Germinating chlamydospore. Unstained.
Terminal culture spherule with one cleavage plane. Mounted in cotton blue
lacto-phenol.
Early stage of zygospore-like structure. Slide culture stained with 1%
aqueous cotton blue.
Early stage of zygospore-like structure. Slide culture stained with 1%
aqueous cotton blue.
. Later stage of zygospore-like structure. Slide culture stained with 1%
aqueous cotton blue.
. Later stage of zygospore-like structure. Slide culture stained with 1%
aqueous cotton blue.
. Zygospore-like structure with thick wall. Slide culture stained in 1%
aqueous cotton blue.
Nuclei in germinating chlamydospores. Stained with iron hematoxylin.
Nuclei in germinating chlamydospores. Stained with iron hematoxylin.
BakKeR, Mrak & SMITH: CoccIpIOIDES IMMITIS Bot
238 FarLowia, VoL. 1, 1943
EXPLANATION OF PLATE V. |
Coccidioides immitis. All figures are of strain 46 and at a magnification of x 1800.
Fig. 1. Nuclei in germinating chlamydospores. Stained with iron hematoxylin.
Fig. 2. Large spherule showing nuclei. Stained with Flemming’s triple stain.
Fig. 3. Large spherule showing nuclei. Stained with Flemming’s triple stain.
Baker, Mrak & SMITH: CoccipIoIDpEs IMMITIS 239 ©
PLATE V. ;
240
FaRLow1A, VoL. 1, 1943
EXPLANATION OF PLATE VI.
Coccidioides immitis. All figures are of strain 46 and shown at a magnification
of x 1700.
Fig. 1. Recently released endospore (or small spherule) containing two nuclei.
Stained with Flemming’s triple stain.
Fig. 2. Small spherule showing two dividing nuclei. Stained with Flemming’s triple
stain. :
Fig. 3. Small spherule with four nuclei. Stained with Flemming’s triple stain.
Fig. 4. Small spherule with eight nuclei. Stained with Flemming’s triple stain.
Fig. 5. Small spherule with eleven nuclei. Stained with Flemming’s triple stain,
Fig. 6. Large spherule with a central vacuole and with numerous nuclei in the peri-
pheral cytoplasm. Stained with Flemming’s triple stain.
Fig. 7. Large spherule with protoplasm cut up into irregular multinucleate segments.
Stained with Flemming’s triple stain.
Fig. 8. Mature spherule containing uni- to multinucleate endospores. One endo-
spore contains one dividing nucleus. Stained with Flemming’s triple stain.
241
Baker, Mrak & SmitH: Coccipioipes IMMITIS
PuatTe VI.
242
FarRLowia, VoL. 1, 1943
EXPLANATION OF PLATE VII.
Coccidioides immitis. All figures are of strain 46 and are shown at a magnification
of x 1700.
Fig. 1. Germinating chlamydospore with a single dividing nucleus. Stained with
iron hematoxylin.
Fig. 2. Uni- and binucleate hyphal cells of young culture. Stained with iron hema-
toxylin.
Fig. 3. Uni- and binucleate hyphal cells of young culture. Stained with iron hema-
toxylin.
Fig. 4, Uni- and multinucleate chlamydospores. Stained with iron hematoxylin.
Fig. 5. Uni- and binucleate chlamydospores. Stained with iron hematoxylin.
Fig. 6. Binucleate spherical chlamydospore. ‘Stained with iron hematoxylin.
Fig. 7. Spherical chlamydospores with four nuclei. Stained with iron hematoxylin.
Fig. 8. Spherical chlamydospore with eight nuclei. Stained with iron hematoxylin.
Baker, Mrak & SmitH: Coccipioipes ImMMITIS 243
Puate VII.
244, Fartowia, VoL. 1, 1943
GERMINATING ENDOSPORES
MATURE TISSUE
SPHERULE Bi
A
sant \
CHLAMYDOSPORES
_cOMPLETE<~ ‘4S a
Qn PLANES IN pu
j=
4 DEVELOPING TISSUE
SPHERULES we
CLEAVAGE PLANES IN
DEVELOPING CULTURE
SPHERULES
K
(a)
MATURE CULTURE iy 5a
o, +
"
LIFE CYCLE
OF
COCCIDIOIDES IMMITIS RIXFORD AND GILCHRIST 1896
Priate VIII.
1(2).: 245-262 FARLOWIA July, 1943
AN INDEX TO RAFINESQUE’S PUBLISHED TECHNICAL
NAMES FOR THE CELLULAR CRYPTOGAMS !
E. D. MERRILL
Somewhat over a year ago | located an entirely overlooked botanical
paper” published by Rafinesque in 1834. This contained a considerable
number of validly published generic and specific names, none of which had
even been listed in our standard indices. An examination of other works
published by Rafinesque indicated also that some of these had never been
indexed, while here and there in works that had been indexed other over-
looked technical names were found. These findings impressed me with the
desirability of preparing and publishing a comprehensive Index Rafines-
quianus, chiefly because of the homonym rule in the International Code
of Botanical Nomenclature. After the lapse of more than a century
it did seem to be desirable at least to list the Rafinesque names; this
task should have been done a half century earlier. .
When the slips were prepared, approximating 12,000, those appertain-
ing to the vascular cryptogams and the cellular cryptogams were segre-
gated. The first study was devoted to the vascular crytogams wherein |
attempted to determine the actual status of each generic name and each
binomial.? The present paper is devoted to a consideration of the names
proposed by Rafinesque for the cellular cryptogams, but unlike his names
for the ferns, I have, in general, in the algae, fungi, lichens, and mosses,
not attempted to determine the status of the various entities. The final
study will be a consideration of the very: numerous entries appertaining to
the phanerogams, but this will be a time-consuming task and may not be
completed for a year or two. In the meantime it was thought to be worth
while to prepare a list of those names, most of them entirely overlooked
for more than a century, appertaining to the cellular cryptogams, and
to publish it in a place where it would be available to specialists con-
cerned with a study of these groups. In the cellular cryptogams it is chiefly
the algae and the fungi that are involved, for Rafinesque’s contributions
to the elucidation of the other major groups were fortunately slight.
With few exceptions Rafinesque’s descriptions of his numerous new
genera and new species of cellular cryptogams are short, vague, indefinite,
and hence highly unsatisfactory. Yet his names, as far as he prepared
and published descriptions for his proposed entities, are validly pub-
lished under all accepted rules of nomenclature, and should at least be
listed. Most of the names that he proposed for the cellular cryptogams
‘The publication of this article out of order of receipt is made possible through a
grant from the Arnold Arboretum.
* Merrill, E. D. A generally overlooked Rafinesque paper. Proc. Am. Philos.
Soc. 86: 72-90. 1942; see also Merrill, E. D. Rafinesque’s work from the stand-
point of world botany, op. cit. 1943. (Jn press.)
* Merrill, E. D. New names for ferns and fern allies proposed by C. S. Rafinesque,
1808-1838. Am. Fern Jour. 33: 41-56. 1943.
245
246 FartowiA, VoL. 1, 1943
have been overlooked by succeeding botanists concerned with the study
of the algae, fungi, lichens, and mosses. In the following list the hitherto
overlooked Rafinesque names are marked with an asterisk. In such biblio-
graphic work as I have been able to accomplish in a field in which I am
not qualified to work as a specialist, I have noted less than forty names
in all categories, that have ever been cited by specialists. In checking
Rafinesque’s names I must admit, however, that my bibliographic work
has scarcely exceeded the consultation of certain standard indices, such
as the Gesamtregister zum I Teil of the first edition of Engler and Prantl’s
Die Natiirlichen Pflanzenfamilien (1909), the indices to the few volumes
of the second edition of that work that have been issued appertaining to
the cellular cryptogams (1924-1938), the indices to Saccardo’s Sylloge
Fungorum, and Mussat’s “Synonymia generum specierum subspecierum-
que in Vol. I—XIV descriptorum” Syll. Fung. 15: i-viii. 1-455. 1901.
Of a total of about three hundred eighty new names in all categories
proposed by Rafinesque for all groups of cellular cryptogams, about three
hundred forty fall in the group of hitherto overlooked ones. In the list
of generic names proposed by Rafinesque as new, totalling about one
hundred thirty-four, only about thirty-one seem ever to have been men-
tioned or reduced to synonymy by Rafinesque’s successors. Almost no
attention has been given to his proposed new species. The percentage
of overlooked Rafinesque names for the cellular cryptogams is very much
higher than is the case for the phanerogams, clearly indicating that the
specialists in the classification of algae and fungi have rather generally
ignored Rafinesque’s crude attempts at taxonomic work in these groups.
Any specialist who will familiarize himself with Rafinesque’s very short,
generalized, and totally inadequate descriptions will probably agree to
the general proposition that it is impossible safely to place most of his
entities in synonymy or otherwise.
These lists have been prepared not with the idea that any specialist may
be intrigued with attempting to determine the status of the many inade-
quately characterized genera and species, but as indicated above, because
of the accepted homonym rule. It is realized that the conservative botanist
will disapprove this bibliographic attempt in his belief that nothing that
Rafinesque published in botany is worthy of consideration. Yet this atti-
tude does not obviate the fact that the homonym rule is official, and that
no matter how bad Rafinesque’s descriptions usually are, his validly pub-
lished names must be considered. It is difficult to see how a valid argu-
ment against listing Rafinesque’s names can be supported, no matter how
gladly this or that botanist would eliminate them. Under the homonym
rule should a modern botanist select a generic name or a binomial that
had been proposed and validly published by Rafinesque over a century ago
for one entity, and apply the same name to the entity that he considers to
be new, then the later name would be invalid and would have to be
changed. The modern taxonomist should be able to determine whether
MERRILL: RAFINESQUE’S NAMES 247
or not a certain name has been previously used, but until such names are
at least listed, he has no way of doing this. He should not be penalized
merely because those bibliographers concerned with the compilation of
comprehensive indices have, in the past, overlooked a considerable num-
ber of Rafinesque’s published works. As to generic names, there is, of
course, the principle of nomina generica conservanda that may be invoked
to preserve later homonyms, but additions to this already long list re-
quires appropriate action from some properly constituted International
Botanical Congress. There is no method of eliminating properly pub-
lished binomials. Because of the approved beginning dates for the names
of the fungi (1821-32, Fries’ Systema mycologicum) for other than the
Uredinales, Ustilaginales, and the Gasteromycetes, probably many of
Rafinesque’s proposed new genera may safely be ignored.
No matter what one may think of the character of Rafinesque’s work
and no matter if one would prefer that all of Rafinesque’s numerous bo-
tanical publications should be officially outlawed, only an International
Botanical Congress can take such action; and in the past only a few pub-
lished works have been officially banned. One illustration of a botanical
taboo is Gandoger’s Florae Europae, which was officially eliminated*; and
I suspect that the chief reason for this action was that Gandoger’s work
appeared in the form of lithographed handwriting rather than actually
printed; but all of Rafinesque’s published works were printed. As long
as such action has not been taken in reference to Rafinesque’s botanical
publications his work will have to be accepted and specialists will have
to do the best that they can with his descriptions. My personal opinion
is that no matter how unorthodox and how poor an individual author’s
work may be, the official banning of the work of this or that author is
extremely dangerous.
In this paper no serious attempt has been made to determine the status
of Rafinesque’s proposed genera and species for the cellular cryptogams,
as I did for the vascular cryptogams, for in the groups covered I do not
consider myself competent to express opinions except in manifestly clear
cases. I do feel, however, that with judicious field work in the neighbor-
hood of Palermo, Sicily, a competent algologist could place a consider-
able number of Rafinesque’s new genera and species of algae; and this
in spite of the short and usually inadequate descriptions, and as far as
he illustrated his species, the very poor figures. In the algae I note that
several of the Rafinesque entities that he definitely placed in that group,
actually represent organisms in the animal kingdom. Thus Pexisperma
lutescens Raf., as figured by him, is an alconyrian coral, probably a
Paragorgia; Phytelis radiata Raf. may represent a large species of Num-
mulites; Helmyton spiralis Raf. might be a species of Spirobus or a
“Gandoger, M. Flora Europae terrarumque adjacentium; sive. Enumeratio
plantarum per Europam atque totam regionem mediterraneam cum insulis atlanticis
sponte crescentium, novo fundamento instauranda. Paris. 27 volumes in 13, 1883-91.
248 FarLowia, Vou. 1, 1943
Serpicula; Siphytus obconicus Raf. probably represents a hydroid, and
S. alternus Raf. is definitely a plumularian hydroid; Chledipole tubulosa
Raf. apparently represents a species of sponge, and Myrsidrum clavatum
Raf. might belong in the same general group of animals. It is only fair
to Rafinesque to note that he considered the Corallina and the Spongidia
to represent algae, placing them in his group II Fucidia in the Algosia
(Algae) in his scheme of classification (see Rafinesque, Anal. Nat. Tabl.
Univ., pp. 206-209. 1815). Rafinesque’s descriptions of the algae are
so short and so generalized that sometimes one can scarcely say what
major group is represented: the Chlorophyceae, Phaeophyceae, or Rhodo-
phyceae.
The situation in reference to Rafinesque’s numerous new genera and
species of fungi is scarcely more promising than it is with the algae, and
for the same reasons, although here illustrations of some of his species
exist. In his Life of Travels, p. 45. 1836, Rafinesque states that he under-
took a voyage from Sicily to the United States in 1815, taking passage on
the “Union of Malta” to New York, and taking with him most of his
fortune and collections:
“50 boxes containing my herbal, cabinet, collections and part of my library. I took
all of my manuscripts with me, including 2000 maps and drawings, 300 copper
plates, &c. My collection of shells was so large as to include 600,000: specimens
large and small. My herbal was so large that I left a part of it.”
This material was all lost in Rafinesque’s shipwreck off Fishers Island
near New London, Connecticut, November 2, 1815. Rafinesque did, how-
ever, later receive from Palermo a set of proofs of a part of the illustra-
tions included on the 300 copper plates mentioned above. These he pre-
sented to the New York Lyceum of Natural History, and they are now
preserved in the library of the New York Botanical Garden. This partial
set is the only known extant copy.
Mr. W. R. Gerard’ examined these proofs in 1885 and states that this
New York set consists of twenty-nine plates on which figures of forty-six
new genera and new species of North American plants appear; these illus-
trations were based on specimens collected or observed by Rafinesque
while he was a resident of Philadelphia, 1802-04. Mr. Gerard lists the
names on these figures verbatim. Thirty-one of the figures represent
fungi, and the remainder flowering plants. Regarding the fungus figures,
Gerard states:
“Most of the figures of fungi seem to have been drawn from memory—possibly
from imagination, and all are quite crudely executed. Those relating to known
genera bear only a remote resemblance to the objects they were designed to illus-
trate, and not one could possibly be of aid in the determination of a species. .. .”
“The small collection of plates is perhaps of interest in connection with the
bibliography of American botany, and as unique relics of one of its earliest, as well
as one of its most eccentric students. The figures of flowering-plants may be of use
°Glerard], W. R. Reliquiae Rafinesquianae. Bull. Torr. Bot. Club 12: 37-38.
1885.
MERRILL: RAFINESQUE’S NAMES 249
in helping to elucidate the author’s vague descriptions, but the plates of the fungi will
only serve to show what a useless mass of rubbish would certainly have been inflicted
upon mycological science had not the fortunate shipwreck of 1815 intervened.”
However, in passing, it is well to note that all of the new genera and
most of the species of fungi that Rafinesque attempted to illustrate on
these lost plates, had already “been inflicted upon mycological science”
for their technical descriptions had been published previous to Rafinesque’s
loss of the plates in 1815. In listing Rafinesque’s genera and species of
fungi I have included references to those names appearing in Gerard’s list.
Rafinesque originated seventy-four new generic names for various
groups of fungi that he considered to be new, or, in a very few cases,
where he proposed new names for genera previously designated by other
authors under names that he did not approve. Until very recently, I judge
that the mycologists have largely ignored these proposed new genera. In
1933, however, Fischer, in Engler & Prantl, Nat. Pflanzenfam. ed. 2, 7a:
84, 1933, actually recognized one of Rafinesque’s generic names as valid,
when he accepted Colonnaria Raf. (1808) as the designation of a certain
group of fungi which he segregated from Clathrus (Micheli) Pers. He
gives the synonymy as Laternia auctt. in part, and Linderia G. H. Cun-
ningham (1931). On what inadequate data most of Rafinesque’s generic
entities in the fungi must be interpreted is illustrated by the following
very inadequate original descriptions, the only ones of Colonnaria pub-
lished by him: “Colonnaria (urceolata, truncata, &c) divided into four
pillars, united at the top, which bear the seeds in the margins. Found in
Penn.” (Med. Repos. II 5: 355. 1808). In the following year this was
translated into French and published thus: “5. Colonnaria urceolata,
truncata; dividée en quatre piliers réunis par le sommet, portant les
semences sur leurs bords. Se trouve en Pensilvanie.” (Jour. Bot. (Paris)
2: 176. 1809).
Incidentally, crude figures of both of these species of Colonnaria exist
in the unique set of proofs of Rafinesque’s lost copper plates, now pre-
served in the library of the New York Botanical Garden. Gerard® com-
ments on these crude illustrations as follows:
“As for the figures of Colonnaria, they bear a general resemblance to Laternea
columnata, in that they have a receptacle composed of four branches united at the
top and base; but here the analogy ends, for C. truncata is surmounted by a perfectly
square, sharp-cornered, open box, full as wide as the lower extremity of the fungus,
and the edge of which is decorated with what looks like a cylindrical moulding. The
other species, C. urceolata, has somewhat the same habit, but differs in the substitu-
tion of a wide circular ring or collar for the “box”—its edge likewise being finished
off with the ornamental moulding. The species of neither of the genera [Aedycia
Raf. and Columnaria Raf.] are provided with a volva, and both of those in the last-
mentioned genus [Colonnaria] remind me very strongly of certain quaint objects
which I have seen in collections of Japanese ceramics. I venture to say that no
so absurd fungi as these ever were, or ever will be, found on the face of the earth.”
®°Glerard], W. R. op. cit. 38.
250 Fartowia, Vor. 1, 1943
With this indictment of Rafinesque’s own figures of his two species of
Colonnaria, one wonders if Fischer used good judgment in accepting
Colonnaria Raf. (1808) as a valid generic name, with the sinking of
Linderia G. H. Cunningham (1931) in its synonymy. Incidentally,
Gerard states on the preceding page that most of Rafinesque’s figures of
fungi “seem to have been drawn from memory — possibly from imagina-
tion—.” As a phanerogamic botanist, were I dealing with a description |
of a flowering plant relatively as short and indefinite as that of Rafin-
esque’s appertaining to Colonnaria, and not supported by extant speci-
mens, my inclination would be to ignore the name, and in this case retain
Linderia G. H. Cunningham; and yet, in spite of the extraordinary Rafin-
esque figures that Gerard discusses, the probabilities are very great that
Rafinesque actually did have specimens of Linderia in mind when he wrote
his very brief description of Colonnaria, and when he made his very crude
and very inaccurate illustrations of the two species. In view of the
universal acceptance of the name Lindera Thunb. (1783) for a genus of
the Lauraceae, I believe that Linderia G. H. Cunningham (1931) is in-
valid, so perhaps after all Fischer should be followed in his interpreta-
tion of Colonnaria Raf.
The lichenologists and the bryologists fare much better than do the
algologists and the mycologists. In the lichens Rafinesque proposed one
new generic name, Scyphorus Raf. to replace Scyphophorus Ach., casually
mentioned Epidrolithus which might be a lichen or possibly an alga or a
fungus (Rafinesque merely states “a curious water lichen, rather than a
Tremella” from which one suspects a species of Leptogium), and pro-
posed but did not describe Leptuberia amorpha, “a small crustaceous
lichen.”” He described only one moss, Heterodon bryoides Raf. The full
description is: “Heterodon (bryoides) small moss with peristome 8-den-
tated, dentatures unequal. It grows in waters of New-Jersey.” Short
as this description is, it seems probable that Heterodon Raf. = Fissidens
Hedwig, for Fissidens is about the only genus of mosses occurring in New
Jersey that has aquatic representatives; yet as Mr. E. B. Bartram notes,
the peristome of Fissidens has more than eight teeth. His only other ven-
ture in this group was to propose the new generic name Diphas (Raf., Am.
Month. Mag. Crit. Rev. 1: 429. 1817) to replace Diphascum Eaton
(Diphascium Ehrh.).
In the following list, where Rafinesque’s new names of cellular crypto-
gams are arranged alphabetically under the Algae, Fungi, Lichenes, and
Musci, his apparently hitherto overlooked technical names are indicated
by an asterisk. Where reductions are made, the authority is cited. Where
only a name appears, without a description, the abbreviation nom.
(nomen nudum) is added, and where descriptions were supplied, no
matter how short, the corresponding abbreviation is descr. (descriptio).
Where no technical descriptions were published, but brief comments are
_made which might supply a clue to the identity of a proposed genus or
MERRILL: RAFINESQUE’S NAMES 251
species, the abbreviation nom. swhnud. (nomen subnudum) is used. Where
a Rafinesque description was repeated verbatim, when certain of his papers
were republished in other periodicals, this repetition is indicated by the
abbreviation iter. (iteratio) ; this does not apply to the series appearing
in the Medical Repository, in English, which were republished in French
in the Journal de Botanique. In those cases where Rafinesque published
two different names for the same entity at the same time, as he frequently
did, the abbreviation nom alt. (nomen alternum) indicates this for the
alternate name. To avoid ambiguity between the two runs of the Jour-
nal de Botanique, in both of which Rafinesque’s papers appear, the
Journal de Botanique 1-2 (1808-09) and 1-4 (1813-16), the former is
indicated as Jour. Bot. (Paris), and the latter as Jour. Bot. (Desv.), for
the reason that inexperienced bibliographers not infrequently confuse the
two. The first was entitled “Journal de botanique rédigé par une
société de botanistes,” and the second “Journal de botanique, appliquée
a l’agriculture, a la pharmacie, a la médicine et aux artes.” Both were
published in Paris. Desvaux is indicated as the principal editor of the
latter and probably also edited the former.
ALGAE’
*Acinaria Raf., Jour. Phys. Chim. Hist. Nat. 89: 107. 1819, descr.; iter., Isis von
Oken 1820, 1: Lit. Anz. p. 243. 1820; Neogen. p. 4. 1825, nom. = Chara Linn.
*coccifera Raf., 1. cc., descr.
*flexuosa Raf., 1. cc., descr.
*Jatifolia Raf., 1. cc., descr.
*salicifolia Raf., 1. cc., descr.
Rafinesque’s material was from the Ohio, Missouri, Mississippi, and Red Rivers.
In his Neogenyton p. 4. 1825, he states that his Acinaria, with Leiacina Raf.,
belongs in the Characeae.
*Amasperma Raf., Précis Découv. Somiol. p. 47. 1814. deser.; Jour. Bot. (Desv.)
4: 273, 1814, descr.; Anal. Nat. Tabl. Univ. p. 209. 1815, nom.
*floculosa Raf., Précis Découv. Somiol. p. 47. 1814, descr.; Jour, Bot. (Desv.)
4; 274, 1814, descr.
*monilia Raf., 1. cc., descr., Sicily.
*torulosa Raf., l. cc., descr., Sicily.
Anevriton Raf., Specch. Sci. 2: 167. 1814, descr.; Anal. Nat. Tabl. Univ. p. 208.
1815, nom.
*marginatum Raf., op. cit. p. 168, descr., Sicily.
*Aporenia Raf., Anal. Nat. Tabl. Univ. p. 209. 1815, nom.
Arthrodia Raf., Specch. Sci. 1: 89. 1814, descr.; Anal. Nat. Tabl. Univ. p. 209.
1815, nom. = Roya W. & G. S. West (1896) fide Printz, Closterium Nitzsch.,
sensu lat.
*linearis Raf., l. c., descr., Sicily.
*Bryonopsis Raf., Specch. Sci. 2: 89. 1814, nom. nov. = Bryopsis Lamouroux.
*Catenaria Raf., Med. Repos. II. 5: 351. 1808, nom.; Jour. Bot. (Paris) 2: 168.
1809, nom., non Sorokin.
*arenaria Raf., 1. cc., nom.
7As noted in the introduction, those generic and specific names in these lists that
are marked with an asterisk appear very generally to have been overlooked by students
of the cellular cryptogams ever since they were published over a hundred years ago. °
252 FartowiA, VoL. 1, 1943
*concatenata Raf., |. cc., nom.
*vagabunda Raf., 1. cc., nom.
The whole original statement is: “Catenaria, (arenaria, vagabunda, concatenata,
&c.) intermediary between some confervae and fuci. On the sea shores.”
[Eastern United States. ]
Ceramium Lyngbe.
*callithamnium Raf., Précis Découv. Somiol. p. 48. 1814, descr., Sicily.
*opacum Raf., l. c., descr., Sicily.
*squamosum Raf., |. c., descr., Sicily.
Chantransia DC.
*dichlora Raf., Précis Découv. Somiol. p. 47. 1814, descr.; Jour. Bot. (Desv.)
4: 273. 1814, descr., Sicily.
Chara Linn.
*fetidissima Raf., Med. Repos. II. 5: 354. 1808, nom.; Jour. Bot. (Paris) 2: 174.
1809, nom. Eastern United States.
*patens Raf., 1. cc. nom. Eastern United States.
Characias Raf., Anal. Nat. Tabl. Univ. p. 209. 1815, nom. nov. = Chara Linn.
*Chledipole Raf., Car. Nuov. Gen. Sp. Sicil. p. 95. 1810, descr. = seq.
*lobata Raf., 1. c., descr., Sicily.
*tubulosa Raf., |. c. t. 30. f. 10, descr., Sicily.
*Chledripole Raf., Specch. Sci. 2: 167. 1814, nom. = Chledipole Raf.
*Colophermum Raf., Précis Découv. Somiol, p. 49. 1814, descr.; Jour. Bot. (Desv.)
4: 275. 1814, descr.; Anal. Nat. Tabl. Univ. p. 209. 1815, nom.
*floccosum Raf., 1. cc., descr., Sicily.
Conferva (Linn.) Lagerh,
*isacella Raf., Précis Découv. Somiol. p. 46. 1814, descr.; Jour. Bot. (Desv.)
4; 273. 1814, descr., Sicily.
*Deloxus Raf., Anal, Nat. Tabl. Univ. p. 208. 1815, nom.
*Deraphytus Raf., op. cit. p. 209. nom.
Dictilema Raf., Précis Découv. Somiol. p. 48. 1814, descr.; Jour. Bot. (Desv.) 4:
274, 1814, descr.; Anal, Nat. Tabl. Univ. p. 209. 1815, nom. = Microdictyon
Decaisne (1839), fide Printz.
*glomerata Raf., Précis Découv. Somiol. p. 48. 1814, descr.; Jour. Bot. (Desv.)
4: 274. 1814, descr., Sicily.
*xanthosperma Raf., 1. cc., descr. Sicily.
*Ecpexis Raf., Atl. Jour. 1: 200. 1833, descr.
“fluviatilis Raf., 1. c., descr. Schuylkill River, Philadelphia.
*Ectosperma Raf., Précis Découv. Somiol. p. 47. 1814, nom.; Jour. Bot. (Desv.) 4:
274. 1814, nom., non Vaucher.
*longiuscula Raf., 1. cc., descr. Sicily.
*Ectospermia Raf. Anal. Nat. Tabl. Univ. p. 209. 1815, nom. = Ectosperma Raf.
*Endonema Raf., Anal. Nat. Tabl. Univ. p. 209. 1815, nom.
*Endosperma Raf., Car. Nuov. Gen. Sp. Sicil. p. 91. 1810, descr.; Anal. Nat. Tabl.
Univ. p. 208. 1815, nom.
*agoregata Raf., |. c., descr. Sicily.
*globosa Raf., 1. c., descr. Sicily.
*Episperma Raf., Précis Découv. Somiol. p. 48. 1814, descr.; Jour. Bot. (Desv.) 4:
274. 1814, descr.
*micramnia Raf., l. cc., descr. Sicily.
*Helmyton Raf., Car. Nuov. Gen. Sp. Sicil. p. 90. 1810, descr.
*glomeratum Raf., |. c., descr. Sicily.
*spiralis Raf., l.c., t. 20 f. 7, descr. Sicily. (Probably belongs in Vermes,
Serpicula. )
*Isophlis Raf., Car. Nuov, Gen. Sp. Sicil. p. 93. 1810, deser.
*concentricus Raf., l. c., ¢. 20. f. 3, descr. Sicily.
MERRILL: RAFINESQUE’S NAMES 253
*Laminaria Raf., Med. Fl. 2: 221. 1830, nom. sub Fucus, non Lamx.
“Some sp. - - - are edible, such as Flucus] edulis, dulcis, saccharinus,
esculentus, palmatus, belonging in the N. G. Laminaria.”
*Leiacina Raf., Neogen. p. 4. 1825, descr. = Chara Linn.
*capitata Raf., l. c., descr. (Chara capitata Ell.) = Chara capitata Elliott.
*lucida Raf.. 1. c., descr. [Eastern United States].
*Leptorima Raf. Car. Nuov. Gen. Sp. Sicil. p. 94. 1810, descr.; Anal. Nat. Tabl.
Univ. p. 208. 1815, nom.
*nivea Raf., 1. c., descr. Sicily.
*oculata Raf., 1. c., descr. Sicily.
*undulata Raf., 1. c. ¢. 20. f. 4, descr. Sicily.
*Lichiton Raf., Specch. Sci. 2: 168. 1814, descr.
*mamillaris Raf, 1. c., descr. Sicily.
*phyteloides Raf., |. c., descr. (Phytelis tuberculata Raf.). Sicily.
*tuberculatum Raf., |. c., descr. Sicily.
*Merasperma Raf., Med. Repos. II. 5: 350. 1808, descr.; Jour. Bot. (Paris) 2: 167.
1809, descr. = Oxytrema Raf. fide Raf., Jour. Phys. Chim. Hist. Nat. 89: 107.
1819; iter., Isis von Oken 1820, 1: Lit. Anz. p. 243. 1820.
*bifurcata Raf., 1. cc., nom. Pennsylvania.
*cylindrica Raf., 1. cc., nom. Pennsylvania.
*dichotoma Raf., 1. cc., nom. Pennsylvania.
*Mesasperma Raf., Anal. Nat. Tabl. Univ. p. 209. 1815, nom.—= Merasperma Raf.
*Myriosydrum Raf. Anal. Nat. Tabl. Univ. p. 208. 1815, nom.= Myrsidrum Raf.
*Myrsidrum Raf., Car. Nuov. Gen. Sp. Sicil. p. 97. 1810, descr., non Bory.
*Bursa Raf., op. cit. p. 98, descr. (Alcyonium Bursa Linn.)
*clavatum Raf., 1. c., ¢. 20 f. 12, descr. Sicily.
*dilatatum Raf., op. cit. p. 99, descr. Sicily.
*effusum Raf., op. cit. p. 98, descr. Sicily.
*ramosum Raf., |. c., descr. Sicily.
*vermilara Raf., 1. c., descr. (Lamarckia vermilara Oliv., Fucus fungosus Desf.,
Ulva tomentosa Linn., Fucus tomentosus Spach). Sicily.
*Nemalectra Raf., Neogen. p. 4. 1825, descr.
*calida Raf., 1. c., nom. “in hot springs” [Eastern United States].
*fuscata Raf., 1. c., nom.
*plumosa Raf., |. ¢., nom.
*Nostocus Raf., Anal. Nat. Tabl. Univ. p. 208. 1815, nom. nov. [Nostoc Linn.].
*Opospermum Raf., Précis Découv. Somiol. p. 48. 1814, descr.: Jour. Bot. (Desv.)
4: 274, 1814, descr.
*nigrum Raf., 1. cc., descr. Sicily.
*Orimanthis Raf., Car. Nuov, Gen. Sp. Sicil. p. 95. 1810, descr.; Anal. Nat. Tabl.
Univ. 208. 1815, nom.
*foliacea Raf., 1. c., descr. Sicily.
*vesciculata Raf., l. c., deser. Sicily.
*Oxytrema Raf., Jour. Phys. Chim. Hist. Nat. 89: 107. 1819, descr.; iter., Isis von
Oken 1820, 1: Lit. Anz. p. 243. 1820. [United States].
Pexisperma Raf., Car. Nuov. Gen. Sp. Sicil. p. 89. 1810, descr.; Anal. Nat. Tabl.
Univ. p. 208. 1815, nom. = Tetraspora Link (1809) fide Printz.
*amplectens Raf., 1. c., descr. Sicily.
*depressa Raf., Précis Découv. Somiol. p. 46. 1814, descr.; Jour. Bot. (Desv.)
4: 273. 1814, descr. Sicily.
*dichrosperma Raf., Car. Nuov. Gen. Sp. Sicil. p. 89. 1810, descr. Sicily.
*lutescens Raf., 1. c. t. 20, f. 1, descr. Sicily.
*sputo Raf., 1. c. ¢. 20. f. 2, descr. Sicily.
*truncata Raf., |. c., descr. Sicily.
254 FarLowiA, VOL. 1, 1943
*Phaxantha Raf., Car. Nuov. Gen. Sp. Sicil. p. 99. 1810, descr.; Anal. Nat. Tabl.
Univ. p, 208. 1815, nom.
*lichenoides Raf., |. c., descr. Sicily.
*Phoracis Raf., Car. Nuov. Gen. Sp. Sicil. p. 99. 1810, descr.; Anal. Nat. Tabl.
Univ. p. 208. 1815, nom.
*filicina Raf., 1. c., descr. Sicily. Regarding this, Rafinesque queries “an Fucus
filicinus Jaq. Wulf. & Gmel.?”
*Phycerus Raf., Specch. Sci. 2: 168. 1814, descr.; Anal. Nat. Tabl. Univ. p. 209.
1815, nom.
*bifidus Raf., l. ¢., descr., Sicily.
*Phylictis Raf., Car. Nuov. Gen. Sp. Sicil. p. 91. 1810, descr.; Anal. Nat. Tabl.
Univ. p. 208. 1815, nom.
*bifurcatus Raf., op. cit. p. 92, descr. Sicily.
*cervicornis Raf., l. c., descr. Sicily.
*cuneiformis Raf., l. c., descr. Sicily.
* dichotomus Raf., op. cit. p. 91, descr. Sicily.
*Jatifolius Raf., op. cit. p. 92, descr. Sicily.
*polypodioides Raf., 1. c., descr. Sicily.
*subfistulosus Raf., l. c., descr. Sicily.
*undulatus Raf., 1. c., descr. Sicily.
*Physidrum Raf., Car. Nuov. Gen. Sp. Sicil. p. 96. 1810, descr.
*agpregatum Raf., op. cit. p. 97, descr. Sicily.
*hyalinum Raf., op. cit. p. 97, descr. Sicily.
*pisiforme Raf., 1. c. p. 96, descr. Sicily.
*rubescens Raf., op. cit. p. 97, t. 20. f. 11, descr. Sicily.
*Physotris Raf., Car. Nuov. Gen. Sp. Sicil. p. 97. 1810, descr. ; Anal. Nat. Tabl. ©
Univ. p. 208. 1815, nom.
*capitata Raf., Précis Découv. Somiol. p. 46. 1814, descr.; Jour. Bot. (Desv.)
4: 273. 1814. descr. Sicily.
*olomerata Raf., Car. Nuov. Gen. Sp. Sicil. p. 97. 1810, descr. Sicily.
*Physydrum Raf., Anal. Nat. Tabl. Univ. p. 208. 1815, nom. = Physidrum Raf.
*Phytelis Raf., Car. Nuov. Gen. Sp. Sicil. p. 93, 1810, descr.; Anal. Nat. Tabl.
Univ. p. 208. 1815, nom.
*atra Raf., op. cit. p. 94, descr. Sicily. ,
*sranulata Raf., 1. c., descr. Sicily.
*macrocarpa Raf., |. c., descr. Sicily.
*radiata Raf., op. cit. p. 93. t. 20. f. 3, descr. Sicily.
*sulcata Raf., op. cit. p. 94. ¢. 20. f. 6, descr. Sicily.
*tuberculata Raf., l. c. (as tubercolata), descr. Sicily. = Lichiton phyteloides.
Raf., fide Rafinesque.
*Phyxalium Raf., Anal. Nat. Tabl. Univ. p. 208. 1815, nom.
*Plaxarthrus Raf., Anal. Nat. Tabl. Univ. p. 209. 1815, nom.
*Porula Raf., Anal. Nat. Tabl. Univ. p. 208. 1815, nom.
*Potarcus Raf., Am. Monthly Mag. Crit. Rev. 3: 356. 1818, descr.; 4: 208. 1819,
nom.; Jour. Phys. Chim. Hist. Nat. 89: 107. 1819, descr.; iter., Isis von Oken
1820, 1: Lit. Anz. p. 243. 1820.
*bicolor Raf., l. cc., descr. Ohio River.
*Sclernax Raf., Car. Nuov. Gen. Sp. Sicil. p. 90. 1810, descr.; Anal. Nat. Tabl.
Univ. p. 208, 1815, nom.
. *lutescens Raf., 1. c., descr. Sicily.
*truncata Raf., |. c., descr. Sicily.
*Siphorus Raf., Car. Nuov. Gen. Sp. Sicil. p. 96. 1810, descr.; Anal. Nat. Tabl.
Univ. p. 208. 1815, nom.
*alternus Raf., |. c., ¢. 20. f. 9., deser. Sicily.
*fasciculatus Raf., |. ¢., descr. Sicily.
MERRILL: RAFINESQUE’S NAMES 255
*Siphytus Raf., Car. Nuov. Gen. Sp. Sicil. p. 95. 1810, descr.
*filiformis Raf., op. cit. p. 96, descr. Sicily.
*hexodon Raf., |. c., descr. Sicily.
*obconicus Raf., 1. c. t. 20. f. 8., descr. Sicily.
*Spermipole Raf., Car. Nuov. Gen. Sp. Sicil. p. 88. 1810, descr.; Anal..Nat. Tabl.
Univ. p. 208. 1815, nom.
*effusa Raf., op. cit. p. 89, descr. Sicily.
*Stereonema Raf., Anal. Nat. Tabl. Univ. p. 209. 1815, nom.
*Stypnion Raf., Ann. Nat. Ann. Synop. p. 16. 1820, descr.
*fluitans Raf., 1. c., descr. Ohio River.
Vaucheria DC.
*dichotoma Raf., Précis Découv. Somiol. p. 47, 1814, descr.; Jour. Bot. (Desv.)
4: 274. 1814, descr. Sicily.
*flexuosa Raf., 1. cc., descr. Sicily.
*stricta Raf., 1. cc., descr. Sicily.
*Vermilaria Raf., Anal. Nat. Tabl. Univ. p. 208. 1815, nom.
Volvox Linn.
*fuscus Raf., Specch. Sci. 2: 68. 1814, descr. Sicily.
*ovalis Raf. l. c., descr. Sicily.
*Zonaria Raf. Anal. Nat. Tabl. Univ. p. 208. 1815, nom.
FUNGI
Actigea Raf., Précis Découv. Somiol. p. 52. 1814, descr.; Anal. Nat. Tabl. Univ.
p. 211. 1815, nom.; Med. Fl. 2: 222. 1830, nom., nota — Scleroderma Pers. fide
Saccardo. = ? Scleroderma Pers. fide Fischer.
multifida Raf., Précis Découv. Somiol. p. 52. 1814, descr. New Jersey = Scler-
oderma flavidum Ell. & Ev. fide Saccardo.
sicula Raf., l. c., descr. Sicily = Scleroderma geaster Fries, fide Saccardo.
* Actigena Raf., Jour. Bot. (Paris) 4: 276 (err. 176). 1814, descr. = Actigea Raf.
*multifida Raf., l. c., descr. New Jersey = Actigea multifida Raf.
*sicula Raf., l. c., descr. Sicily = Actigea sicula Raf.
Actinophora Raf., Med. Repos. II. 3: 423. 1806, nom.; op. cit. 5: 355, 1808, nom.;
Jour. Bot. (Paris) 2: 176. 1809, nom.; Précis Découv. Somiol. p. 51. 1814,
descr.; Jour. Bot. (Desv.) 4: 275. 1814, descr.; Anal. Nat. Tab. Univ. p. 211.
1815, nom.; Fischer in Engl. & Prantl, Nat. Pflanzenfam. ed. 2. Ta: 119. 1933,
gen. dub.
*aurantiaca Raf. 1. cc., nom., Précis Découv. Somiol. p. 51. 1814, descr.; Jour.
Bot, (Desv.) 4: 275. 1814, descr.; Gerard, Bull. Torr. Bot. Club 12: 37. 1885,
nom. Delaware, Pennsylvania.
* Actycus Raf., Chloris Aetn. p. 13, 180, nom.
*siculus Raf., 1. c., nom. Sicily.
Aecidium Pers.
*citrinum Raf., Stat. Gen. Sicil. p. 28. 1810, nom. Sicily.
Aedycia Raf., Med. Repos. II. 5: 358, 1808, descr.; Jour. Bot. (Paris) 1: 222.
1809, descr.; Anal. Nat. Tabl. Univ. p. 211. 1815, nom. = Mutinus Fries, fide
Saccardo.
*alba Raf., 1. cc., descr.; Gerard, Bull. Torr. Bot. Club 12: 37. 1885, nom.
Pennsylvania.
*rubra Raf., 1. cc., descr. Pennsylvania.
Agaricus Linn.
*aterrimus Raf., Jour. Bot. (Desv.) 1: 236. 1813, descr. Delaware = Amanita
aterrima Raf.
*azureus Raf., 1. c., descr. Delaware = Amanita azurea Raf.
*ellipticus Raf., Am. Monthly Mag. Crit. Rev. 3: 356. 1818, nom. Alleghany
Mountains or Ohio,
256 FarLowiaA, VoL. 1, 1943
*miptica Raf. ex M’Murtrie, Sketch. Louisv. p. 230. 1819, nom. Kentucky.
*surrectus Raf., Casket 1831: 423, J. f. 1831, descr. Pennsylvania.
*Alveolinus Raf., Anal. Nat. Tabl. Univ. p. 211. 1815, nom.
Amanita Pers.
*atterima Raf., Précis Découv. Somiol. p. 49. 1814, descr.; Jour. Bot. 4: 275. 1814,
nom. Delaware = Agaricus aterrimus Raf,
*azurea Raf., 1. cc., descr. nom. Delaware — Agaricus azureus Raf.
*Anastomaria Raf., Ann. Nat. Ann. Synop. p. 16. 1820, descr.
*campanulata Raf., |. c., descr. New York.
*dimidiata Raf., 1. c., descr. New York.
*Annularia Raf., Anal. Nat. Tabl. Uniy. p. 211. 1815, nom.
*Astrocitum Raf., Med. Repos. II. 3: 423, 1806, nom. = Astrycum Raf.
*dimidiatum Raf., |. c., nom. New York = Astrycum dimidiatum Raf.
*multifidum Raf., 1. ¢., nom. Pennsylvania, New Jersey = Astrycum multifidum
Raf. :
*quinquefidum Raf., 1. c., nom. Pennsylvania, New Jersey = Astrycum quinque-
fidum Raf,
Astrycum Raf., Med. Repos. II. 5: 355. 1808, nom. “tribe of the licoperdoideous”
fide Rafinesque = Actigea Raf. fide Saccardo.
*dimidiatum Raf., 1. c., nom.; Jour. Bot. (Paris) 2: 175. 1809, nom.; Gerard,
Bull. Torr. Bot. Club 12: 37. 1885, nom. Pennsylvania, New Jersey.
*multifidum Raf., 1. ec., nom.; Gerard, 1. ¢., nom. Pennsylvania, New Jersey.
*quinquefidum Raf., |. cc., nom. Pennsylvania, New Jersey.
Boletus Dill.
*“hematoporus Raf. ex M’Murtrie, Sketch, Louisv. p. 230. 1819, nom. Kentucky.
*pusillus Raf., Chloris Aetn. p. 13. 1815, nom. Sicily.
Cerophora Raf., Med. Repos. II. 5: 355. 1808, nom.; Jour. Bot. (Paris) 2: 176.
1809, nom.; Précis Découv. Somiol. p. 49. 1814, descr.; Jour. Bot. (Desv.) 4:
275. 1814, descr.; Anal. Nat. Tabl. Univ. p. 211. 1815, nom. = Hericium Pers.
fide Saccardo.
*capitata Raf., Précis Découv. Somiol. p. 50. 1814, descr.; Jour. Bot. (Desv.) 4:
275. 1814, descr.; Gerard, Bull. Torr. Bot. Club 12: 37. 1885, nom. New Jersey.
*clavata Raf., Med. Repos. II. 5: 355. 1806, nom.; Jour. Bot. (Paris) 2: 176.
1809, nom.; Précis Découy. Somiol. p. 49, 1814, descr.; Jour. Bot. (Paris) 4:
275. 1814, descr.; Gerard, Bull. Torr. Bot. Club 12: 37. 1885, nom. New Jersey.
*dichotoma Raf., Med. Repos. II. 5: 355. 1808, nom.; Jour. Bot. (Paris) 2: 176.
1809, nom.; Gerard, Bull. Torr. Bot. Club 12: 37. 1885, nom. Central Atlantic
United States.
*fastigiata Raf., 1. cc., nom. Central Atlantic United States,
*globosa Raf., 1. cc., nom. Central Atlantic United States.
*globularis Raf. ex Gerard, Bull. Torr. Bot. Club 12: 37. 1885, nom. Central
Atlantic United States, prob. = C. globosa Raf.
*minuta Raf., 1. cc., nom. Central Atlantic United States.
*pyriformis Raf., 1. cc., nom.; Gerard, 1. c., nom. Central Atlantic United States.
*ramosa Raf. ex Gerard. Bull. Torr. Bot. Club 12: 37. 1885, nom. Central Atlantic
United States,
*thamnoides Raf., 1. cc., nom. Central Atlantic United States.
Clavaria Vaill.
*bicolor Raf., Med. Repos. II. 5: 363. 1808, descr.; Jour. Bot. (Paris) 1: 233. 1809,
descr.; Gerard, Bull. Torr. Bot. Club 12: 37, 1885, nom. Virginia.
*citrina Raf., op. cit. 362, 233, descr. Pennsylvania.
*citrinofusca Raf., 1. cc., descr.; Gerard, |. c., nom. Pennsylvania.
*driophylla Raf., Jour. Bot. (Paris) 1: 233. 1809, descr. = C. dryophylla Raf.
*dryophylla Raf., Med. Repos. II. 5: 363, 1808, descr.; Gerard, 1. c., nom. Penn-
sylvania.
‘
MERRILL: RAFINESQUE’S NAMES 25%
*lepidorhiza Raf., 1. cc., descr. Maryland.
*oxantha Raf., Précis Découv. Somiol. p. 51. 1814, descr.; Jour. Bot. (Desv.)
1: 237. 1813, descr.; op. cit. 4: 276 (err. 176). 1814, nom. Maryland.
*rubescens Raf., Précis Découv. Somiol. p. 51. 1814, descr.; Jour. Bot. (Desv.)
1: 237. 1813, descr.; Jour. Bot. (Desv.) 4: 276 (err. 176). 1814, nom. Penn-
sylvania.
*tricolor Raf., Med. Repos. II. 5: 363. 1808, descr.; Jour. Bot. (Paris) 1: 233.
1809, descr.; Gerard, Bull. Torr. Bot. Club 12: 37. 1885, nom. Maryland.
Colonnaria Raf., Med. Repos. II. 3: 423. 1806, nom.; op. cit. 5: 355. 1808, descr.;
Jour, Bot. (Paris) 2: 176. 1809, descr. = Clathrus (Mich.) Pers., fide Saccardo;
Fischer in Engler & Prantl, Nat. Pflanzenfam. ed. 2, 7a: 84. 1933 (= Linderia
G. H. Cunningham, 1931), genus validus fide Fischer 1. c. See p. 249-250.
truncata Raf., I. cc., nom.; Gerard, Bull. Torr. Bot. Club 12: 37, 1885. nom. Penn-
sylvania = Clathrus truncatus Bosc, fide Saccardo.
urceolata Raf., l. cc., nom.; Gerard, 1. c., nom. Pennsylvania = Clathrus columnatus
Bosc, fide Saccardo.
*Cyathela Raf., Am. Monthly Mag. Crit. Rev. 4: 208. 1819, nom.
*Cyathella Raf., Anal. Nat. Tabl. Univ. p. 211. 1815, nom.
*Cynicus Raf., Anal. Nat. Tabl. Univ. p. 211. 1815, nom.
*Dedalea Raf., Anal. Nat. Tabl. Univ. p. 211. 1815, nom. (haud Daedalea Pers.! )
*Dicarphus Raf., Med. Repos. II. 3: 423, 1806, nom.; op. cit. II. 5: 355. 1808,
nom. subnud.; Jour. Bot. (Paris) 2: 176. 1809, nom. subnud.; Anal. Nat. Tabl.
Univ. p. 211. 1815, nom. = ? Hydnum Linn., fide Saccardo.
*rubens Raf., 1. cc. nom.; Gerard, Bull. Torr. Bot. Club 12: 37. 1885, nom.
Pennsylvania.
*Dicteridium Raf., Anal. Nat. Tabl. Univ. p. 211. 1815, nom.
*Druparia Raf., Med. Repos. II. 5: 358. 1808, descr.; Anal. Nat. Tabl. Univ. p. 211.
1815, nom. = Drupasia Raf.
*clobosa Raf., Med. Repos. II. 5: 358. 1808, descr. Pennsylvania.
*rosea Raf., l. c., descr. Delaware.
*violacea Raf., l. c., descr. Pennsylvania.
*volvacea Raf. ex Gerard, Bull. Torr. Bot. Club 12: 37. 1885, nom.
Drupasia Raf., Jour. Bot. (Paris) 1: 223. 1809, descr.; Fischer in Engler & Prantl,
Nat. Planzenfam. ed. 2, 7a: 119. 1933, gen. dub. = Druparia Raf.
*olobosa Raf. 1..c., descr. Pennsylvania.
*rosea Raf. 1. c., descr. Delaware.
*violacea Raf. 1. c., descr. Pennsylvania.
Dycticia Raf. Med. Repos. II. 5: 355. 1808, nom.; Jour. Bot. (Paris) 2: 176. 1809,
nom. = Clathrus (Mich.) Pers. vel. aff. fide Saccardo; Fischer in Engl. &
Prantl, Nat. Pflanzenfam. ed. 2, 7a: 84. 1933 = ? Clathrus (Mich.) Pers.
*clathroides Raf., 1. cc., nom. subnud. Delaware.
Endacinus Raf. Précis Découy. Somiol. p. 51. 1814, descr.; Jour. Bot. (Desv.) 4:
276 (err. 176). 1814, descr. = Polysaccum DC. in Desport & DeCandolle fide
Saccardo = Pisolithus A. & S.
tinctorius Raf., 1. cc., descr. Sicily = Polysaccum pisocarpum Fries, fide Sac-
cardo = Pisolithus tinctorius (Pers.) Coker.
*Endaematus Raf., Specch. Sci. 2: 105. 1814, descr.
*albus Raf., 1. c., descr. Sicily.
*Endematus Raf., Anal. Nat. Tabl.. Univ. p. 207. 1815, nom. = Endaematus Raf.
Endoconia Raf., Jour. Phys. Chim. Hist. Nat. 89: 106. 1819, descr.; iter., Isis von
Oken 1820, 1: Lit. Anz. p. 243. 1820—Hyphymocyteae gen. incert. fide
Saccardo.
*leucomela Raf., 1. cc., descr. Kentucky.
*stuposa Raf., |. cc., descr. New York.
*Endonius Raf., Am. Monthly Mag. Crit. Rev. 4: 208. 1819, nom.
258 Fartowia, Vou. 1, 1943
*Epixyla Raf., Med. Repos. II. 3: 423. 1806, nom.
*clavata Raf., 1. c., nom. Central Atlantic United States.
*cylindrica Raf., 1. c., nom. Central Atlantic United States.
*lineata Raf., 1. c., nom. Central Atlantic United States.
*Eriosperma Raf., Med. Repos. II. 5: 356. 1808, nom. subnud.; Jour. Bot. (Paris)
2: 177. 1809, nom. subnud.
“alba Raf., 1. cc.. nom. Pennsylvania.
*fugax Raf., 1. cc., nom. Pennsylvania.
*Eudacnus Raf., Anal. Nat. Tabl. Univ. p. 211. 1815, nom.
*Favaria Raf., Anal. Nat. Tabl. Univ. p. 211. 1815, nom.
Gelatina Raf., Med. Repos. II. 3: 423. 1806; op. cit. 5: 356. 1808, descr.; Jour.
Bot. (Paris) 2: 177. 1809, descr. = Tremella Dill., fide Saccardo.
*alba Raf., 1. cc., nom. Eastern United States,
*aurantiaca Raf., |. c. Il. 3: 423. 1806, nom. Eastern United States.
*candida Raf., 1. c., nom. Eastern United States.
“flava Raf., 1. c., nom. Eastern United States.
*foetidissima Raf., op. cit. II. 5: 356. 1808, nom.; Jour. Bot. (Paris) 2: 177.
1809, nom. Eastern United States.
*lutea Raf., 1. cc., nom. Eastern United States.
*rubra Raf., op. cit. II. 3: 423. 1806, nom.; op. cit. 5: 356. 1808, nom.; Jour. Bot.
(Paris) 2: 177. 1809, nom. Eastern United States.
*Gelatinaria Raf., Anal. Nat. Tabl. Univ. p. 211. 1815, nom. (non Roussel !)
*Gemmularia Raf., Jour. Phys. Chim. Hist. Nat. 89: 106, 1819, descr.; iter., Isis von
Oken 1820, 1: Lit. Anz. p. 243. 1820; Med. Fl. 2: 270. 1830, nom.; Atl. Jour.
1: 181. 1833, descr. = Tucahus Raf. = Rugosaria Raf.
*albida Raf., Atl. Jour. 1: 182, 1833, descr., nom. alt. Tucahus albidus Raf.
Pennsylvania, Ohio, Kentucky.
*leviuscula Raf., Jour. Phys. Chim. Hist. Nat. 89: 106. 1819, descr.; iter., Isis von
Oken 1820, 1: Lit. Anz, p. 243. 1820; Atl. Jour. 1: 181. 1833, descr., nom. alt.
Tucahus leviusculus Raf. Virginia, Kentucky.
*rimosa Raf., Atl. Jour. 1: 181. 1833, descr., nom. alt. Tucahus rimosus Raf.
Virginia, North Carolina.
*rugosa Raf., Jour. Phys. Chim. Hist. Nat. 89: 106. 1819, descr.; iter., Isis von
Oken 1820, 1: Lit. Anz. p. 243. 1820; Atl. Jour. 1: 181. 1833, deser., nom. alt.
Tucahus rugosus Raf. Maryland, South Carolina.
*Geoglossum Raf., Anal, Nat. Tabl. Univ. p. 211. 1815, nom. (an Persoon ?)
*Gymnopus Raf., Anal. Nat. Tabl. Univ. p. 211. 1815, nom. (an Persoon ?)
Hectocerus Raf., Med. Repos. II. 3: 423. 1806, nom. = Hericium Pers., fide Saccardo.
*clavatus Raf., 1. c., nom. Central Atlantic United States.
*dichotomus Raf., 1. c., nom. Central Atlantic United States.
*globosus Raf., 1. c., nom. Central Atlantic United States.
*pyriformis Raf., ]. c., nom. Central Atlantic United States.
*thamnoides Raf., 1. c., nom. Central Atlantic United States.
Hepataria Raf., Med. Repos. II. 5: 350. 1808, nom.; Jour. Bot. (Paris) 2: 168. 1809,
nom. = Tremella Dill. vel aff. fide Saccardo.
*cuneata Raf., 1. cc., nom. Central Atlantic United States.
*erecta Raf., 1. cc., nom. Central Atlantic United States.
Hericium Pers.
“grande Raf., Jour. Bot. (Desv.) 1: 236. 1813, descr.; Précis Découv. Somiol.
p. 50. 1814, descr.; Jour. Bot. (Desv.) 4: 276 (err, 176) 1814, nom. New Jersey.
Hydnum Linn.
*aurantinum Raf.. Précis Découy. Somiol. p. 50, 1814, descr. Delaware — H. auran-
tium Raf.
*aurantium Raf., Jour. Bot. (Desv.) 1: 237. 1813, descr.; op. cit. 4: 276 (err. 176).
1814, nom. Delaware.
MERRILL: RAFINESQUE’S NAMES 259
barbatum Raf., Med. Repos. II. 5: 363. 1808, descr.; Jour. Bot. (Paris) 1: 234.
1809, descr.; Gerard, Bull. Torr. Bot. Club 12: 37. 1885, nom. Delaware —
Hydnum sp. dub. fide Saccardo.
caerulescens Raf., Med. Repos. II. 5: 363. 1808, descr.; Jour. Bot. (Paris) 1:
234. 1809, descr.; Gerard, Bull. Torr. Bot. Club 12: 37. 1885, nom. (cerule-
scens). New Jersey = Hydnum sp. dub. fide Saccardo (as coerulescens).
citrinum Raf., |. cc., descr.; Gerard, |. c., nom. Pennsylvania = Hydnum sp. dub.
fide Saccardo.
dilatatum Raf., 1. cc., descr.; Gerard, 1. c., nom.:Pennsylvania-= Hydnum sp.
dub. fide Saccardo.
*puniceum Raf., Jour. Bot. (Desv.) 1: 237. 1813, descr.; Précis Découv. Somiol.
p. 50. 1814, descr.; Jour. Bot. (Desv.) 4: 276 (err. 176). 1814, nom. Delaware.
Hydromycus Raf., Med. Repos. II. 5: 356. 1808, nom. subnud.; Jour. Bot. (Paris)
2: 178. 1809, nom. subnud. = Dacryomyces Nees fide Saccardo.
*aquosus Raf., 1. cc., nom. New Jersey, Pennsylvania.
*tremelloides Raf., 1. cc., nom. New Jersey, Pennsylvania.
Hypolepia Raf., Med. Repos. II. 5: 356. 1808, nom. subnud.; Jour. Bot. (Paris)
2: 178. 1809, nom. subnud. = Dacryomyces Nees fide Saccardo.
*difformis Raf.; 1. cc., nom. Central Atlantic United States.
*igniarias Raf., Med. Repos. II. 5: 356. 1808, nom. Central Atlantic United States.
*ignarius Raf., Jour. Bot. (Paris) 2: 178. 1809, nom. Central Atlantic United States.
Isaria Pers.
*odora Raf., Jour. Bot. (Desv.) 1: 237. 1813, descr.; Précis Découv. Somiol. p. 51.
1814, descr.; Jour. Bot. (Desv.) 4: 276 (err. 176). 1814, nom. Maryland.
*Lamyxis Raf. Ann. Nat. Ann. Synop. p. 16. 1820, nom. provis. sub Sisostrema
globularis Raf.
Leptopora Raf., Med. Repos. II. 5: 355. 1808, descr.; Jour. Bot. (Paris) 2: 177.
1809, descr. = Poria Pers. fide Saccardo.
*difformis Raf., 1. cc., nom. Central Atlantic United States.
*nivea Raf., 1. cc., nom. Central Atlantic United States.
*stercoraria Raf., 1. cc., nom. Central Atlantic United States.
Lycoperdon (Tourn.) Pers.
*coccineum Raf., Jour. Bot. (Desv.) 1: 237. 1813, descr.; Précis Découv. Somiol.
p. 51. 1814, descr.; Jour. Bot. (Desv.) 4: 276 (err. 176). 1814. nom. Delaware.
*violacinum Raf., (Lycoperdum) Jour. Bot. (Paris) 4: 276 (err. 176). 1814, in
syn. [Pennsylvania] = Omalycus violacinus Raf.
Lycopodium Linn. :
*violacinum Raf., Jour. Bot. (Desv.) 1: 236. 1813, descr. sub Mycastrum siculum
Raf., — Lycoperdon intended. Pennsylvania = Omalycus violacinus Raf.
Merulius Pers.
*undulatus Raf., Jour. Bot. (Desv.) 1: 237. 1813, descr., Précis Découv. Somiol.
_ p. 50. 1814, descr. Maryland.
Morchella Dill.
*cava Raf., Car. Nuov. Gen. Sp. Sicil. p. 88. 1810, descr.; Stat. Gen. Sicil. p. 28.
1810, nom.; Chloris Aetn. p. 14, 1815, nom. Sicily.
*odorata Raf., Fl. Ludovic. p. 12. 1817, descr. Louisiana.
Mycastrum Raf., Jour. Bot. (Desv.) 1: 236. 1813, descr. —= Omalycus Raf, fide
Desy., Jour. Bot. (Desv.) 4: 276 (err. 176). 1814 = Actigea Raf. fide Saccardo.
*siculum Raf., 1. c., descr. Sicily. -
*Nidularia Raf., Anal. Nat. Tabl. Univ. p. 211. 1815, nom. (an Fries ?)
Omalycus Raf., Précis Découv. Somiol. p. 52. 1814, descr.; Anal. Nat. Tabl. Univ.
p. 211. 1815, nom. = Scleroderma Pers. fide Saccardo.
*erosus Raf., Précis Découv. Somiol. p. 52. 1814, nom. nov., Lycoperdon com-
planatum Desv., Jour. Bot. (Desv.) 4: 276 (err. 176). 1814.
260 FartowiA, Vo. 1, 1943
violacinus Raf., l. cc., descr., nom. (Lycoperdon violacinum Raf.). Pennsylvania =
Scleroderma violacinum De Toni, fide Saccardo.
*Onygena Raf., Anal. Nat. Tabl. Univ. p. 211. 1815, nom. (an Persoon ?)
Peziza Dill.
*alborufa Raf., Med., Repos. IJ. 5: 362. 1808, descr.; Jour. Bot. (Paris) 1: 232.
1809, descr.; Gerard, Bull. Torr. Bot. Club 12: 37. 1885, nom. Pennsylvania.
*atrata Raf., l. cc., descr. Pennsylvania, New Jersey.
*cupularia Raf., Med. Repos. II. 5: 362. 1808, descr. Delaware = seq.
*cupularis Raf., Jour. Bot. (Paris) 1: 232. 1809. descr. Delaware.
*depressa Raf., 1. cc., descr.; Gerard, Bull. Torr. Bot. Club 12: 37. 1885, nom.
Pennsylvania.
*globulosa Raf., 1. cc., descr.; Gerard, |. c., nom. Pennsylvania,
*lupularia Raf. ex Gerard, Bull. Torr. Bot. Club 12: 37. 1885, nom.
*ochrochlora Raf., |. cc., descr.; Gerard, |. c., nom. New Jersey.
*pulcherrima Raf., |. cc., descr.; Gerard, |. c., nom. Pennsylvania.
*smaragdina Raf., Jour. Bot. (Paris) 1: 232. 1809, descr. Pennsylvania.
*smeraldina Raf., Med. Repos. II. 5: 362. 1808, descr.; Gerard, Bull. Torr. Bot.
Club 12: 37. 1885 (smiraldina) Pennsylvania = P. smaragdina Raf.
*Pherima Raf., Am. Monthly Mag. Crit. Rev. 4: 208. 1819, nom. = Phorima Raf.
*Phiala Raf., Anal. Nat. Tabl. Univ. p. 211. 1815, nom. (an Phialea Fries ?)
Phorima Raf., Med. Repos. II. 3: 423. 1806, nom.; op. cit. 5: 355. 1808, descr.;
Précis Découv. Somiol. p. 49. 1814, descr.; Jour. Bot. (Desv.) 4: 275. 1814,
descr.; Anal. Nat. Tabl. Univ. p. 211. 1815, nom. = Favolus Beauv. fide Desvaux,
Jour. Bot. (Desv.) 4: 275. 1814.
*betulina Raf., Med. Repos. II. 3: 423. 1806, nom.; op. cit. 5: 355. 1808, nom.;
Jour. Bot. (Paris) 2: 177. 1809, nom.; Gerard, Bull. Torr. Bot. Club 12: 37.
1885, nom. Central Atlantic United States.
*boletoides Raf., Med. Repos. IJ. 3: 423. 1806, nom. Central Atlantic United States,
*coccinea Raf., op. cit. I]. 5: 355. 1808, nom.; Jour. Bot. (Paris) 2: 177. 1809,
nom, Central Atlantic United States.
*difformis Raf., op. cit. II. 3: 423. 1806, nom. Central Atlantic United States.
*minuta Raf., op. cit. I]. 5: 355. 1808, nom.; Précis Découv. Somiol. p. 49. 1814,
descr.; Jour. Bot. (Paris) 2: 177. 1809, nom.; Jour. Bot. (Desv.) 4: 275. 1814,
descr. Central Atlantic United States.
*Phyllops Raf., Fl. Ludovic. p. 172. 1817, nom. Louisiana.
*Piemyeus Raf., Jour. Bot. (Desv.) 1: 236. 1813, nom. subgen. sub Mycastrum
siculum Raf. = Piesmycus Raf.
Piesmycus Raf., Med. Repos. II. 5: 355. 1808, descr.; Jour. Bot. (Paris) 2: 176.
1809, descr. = Lycoperdon (Tourn.) Pers. vel aff. fide Saccardo; gen. dub.
fide Fischer in Engl. & Prantl, Nat. Pflanzenfam. ed. 2, 7a: 72. 1933.
*nigrescens Raf., 1. cc., nom. Central Atlantic United States.
*violaceus Raf., 1. cc., nom. Central Atlantic United States.
*Polystema Raf., Ann. Nat. Tabl. Univ. p. 211. 1815, nom.
*Poronea Raf., Ann. Nat. Tabl. Univ. p. 207. 1815, nom.
Priapus Raf., Med. Repos. II. 3: 423. 1806, nom.; op. cit. 5: 355. 1808, nom. subnud. ;
Jour. Bot. (Paris) 2: 176. 1809, nom. subnud.—= Phallus Linn. vel aff. fide
Saccardo.
*niveus Raf., l. cc., nom. subnud. Virginia.
*Pyrenium Raf., Anal. Nat. Tabl. Univ. p. 211. 1815, nom.
Pyrisperma Raf., Med. Repos. II. 3: 423. 1806, nom.; op. cit. 5: 355. 1808; nom.
subnud.; Jour. Bot. (Paris) 2: 177. 1809, nom. subnud. = Tuber (Micheli)
Fries vel aff. fide Saccardo.
*hypogea Raf., 1. cc., nom. New Jersey.
*Ramaria Raf., Anal. Nat. Tabl. Univ. p. 211. 1815, nom. (an Persoon ?)
>
MERRILL: RaFINESQUE’s NAMES ; 261
Riella Raf. ex Sacc., Syll. Fung. 15: 323. 1901, in syn. = Rimella Raf. = Tylostoma
Pers. vel aff. fide Saccardo.®
Rimella Raf., Am. Monthly Mag. Crit. Rev. 4: 208. 1819, nom.; Jour. Phys. Chim.
Hist. Nat. 89: 106. 1819, descr.; iter., Isis von Oken 1820, 1: Lit. Anz. p. 243.
1820 = Tylostoma Pers. vel aff. fide Saccardo.
*obovalis Raf., Jour. Phys. Chim. Hist. Nat. 89: 106. 1819, descr.; iter., Isis
von Oken 1820, 1: Lit. Anz. p. 243. 1820. Virginia and along the Ohio River.
*Rotula Raf., Anal. Nat. Tabl. Univ. p. 211. 1815, nom.
*Rugosaria Raf., Atl. Jour. 1: 181. 1833, nom. alt. Gemmularia Raf. = Tucahus Raf.
*Sarcanthia Raf., Fl. Ludovic. p. 172. 1817, nom. Louisiana.
*Sarcomelas Raf., Fl. Ludovic. p. 172. 1817, nom. Louisiana.
Sarconemus Raf., Anal. Nat. Tabl. Univ. p. 210. 1815, nom. “genus omnino incertum’
Saccardo.
*Sisostrema Raf., Ann. Nat. Ann. Synop. p. 16. 1820, nom., ut videtur sphalm. =
Sistotrema Pers.
*olobularis Raf., 1. c., descr. New York.
*Sphaeropsis Raf., Anal. Nat. Tabl. Univ. p. 207. 1815, nom. nov. = Sphaeria Pers.
Stemastrum Raf., Med. Repos. II. 5: 355. 1808, nom. subnud.; Jour. Bot. (Paris)
2: 177. 1809, nom. subnud. Virginia = Geaster Mich. aff. fide Saccardo.
*Boscii Raf., 1. cc., nom. subnud. Virginia.
*Stericium Raf., Am. Monthly Mag. Crit. Rev. 4: 208. 1819, nom.
*Stipiza Raf., Ann. Nat. Tabl. Univ. p. 211. 1815, nom.
Thelephora Ehrh.
*aurantiaca Raf., Med. Repos. II. 5: 363. 1808, descr.; Jour. Bot. (Paris) 1: 233.
1809, descr. Central Atlantic United States.
*nigrescens Raf., Jour. Bot, (Desv.) 1: 237. 1813, descr.; Précis Découv. Somiol.
p. 50. 1814, descr.; Jour. Bot. (Desv.) 4: 276 (err. 176). 1814, nom. Penn-
sylvania.
*revoluta Raf., 1. cc., descr.; Jour. Bot. (Desv.) 4: 276 (err. 176). 1814, nom.
Pennsylvania.
*semicampanulata Raf., Med. Repos. II. 5: 363. 1808; Jour. Bot. (Paris) 1: 233.
1809, descr. Pennsylvania.
*Trichidium Raf., Anal. Nat. Tabl. Univ. p. 211. 1815, nom. nov. Trichia Hall.
Tuber (Micheli) Fries
*rufescens Raf., Jour. Bot. (Paris) 1: 235, 1813, nom.; Précis Découv. Somiol.
(p. 3 of cover), nom. Sicily.
*Tuberium Raf., Anal. Nat. Tabl. Univ. p. 211. 1815, nom. nov. Tuber (Micheli)
Pers.
*Tucahus Raf., Med. Fl. 2: 270. 1830, descr.; Atl. Jour. 1: 1833, descr. = Gemmu-
laria Raf. = Rugosaria Raf. fide Raf.
*albidus Raf., 1. cc., nom., descr. Pennsylvania, Ohio, Kentucky.
*leviusculus Raf., 1. cc.. nom., descr. Carolina.
*rimosus Raf., Atl. Jour. 1: 181. 1833, descr. Virginia and North Carolina.
*rugosus Raf., 1. cc., nom., descr. Maryland to South Carolina.
Uredo Pers.
*citrina Raf., Car. Nuov. Gen. Sp. Sicil. p. 88. 1810, descr. Sicily, on Dianthus
dubius.
Volvaria Fries
*coccinea Raf., Med. Repos. II. 3: 423. 1806, nom.; Gerard, Bull. Torr. Bot. Club
12: 37. 1885, nom. Maryland = Volvycium coccineum Raf.
Volvycium Raf., Med. Repos. II. 5: 358. 1808, descr.; Jour. Bot. (Paris) 1: 222.
1809, descr.; Anal. Nat. Tabl. Univ. 211. 1815, nom. = Battarrea Pers. aff. fide
>
*In all of Rafinesque’s published papers that I have examined I did not detect the
name Riella Raf. It is perhaps published in the work of some other mycologist;
unfortunately Saccardo cites no reference.
262 Fartowia, Vou. 1, 1943
Saccardo = gen. dub. fide Fischer in Engl. & Prantl, Nat. Pflanzenfam. ed.
2, Za: 119. 1933.
*coccineum Raf., 1. cc., descr.; Gerard, Bull. Torr. Bot. Club 12: 37, 1885, nom.
Maryland.
Xylissus Raf., Med. Repos. II. 5: 356. 1808, nom. subnud.; Jour. Bot. (Paris) 2:
177. 1809, nom. subnud. = Dacryomyces Nees vel aff. fide Saccardo.
*cylindricus Raf., 1. cc., nom. Pennsylvania.
*lineatus Raf., 1. cc., nom. Pennsylvania.
*oblongus Raf., 1. cc., nom. Pennsylvania.
*Zonilia Raf., Anal. Nat. Tabl. Univ. p. 211. 1815, nom.
LICHENES
*Epidrolithus Raf., Life of Travels p. 136. 1836, nom. Pennsylvania. An Leptogium
(Ach.) S. Gray ?
*Leptuberia Raf., Med. Repos. II. 5: 350. 1808, nom.; Jour. Bot. (Paris) 2: 167.
1809, nom.
*amorpha Raf., 1. cc.. nom. Pennsylvania.
*Scyphorus Raf., Anal. Nat. Tabl. Univ. p. 208. 1815, nom. nov. Scy phophorus
Acharius,
MUSCI
*Diphas Raf., Am. Monthly Mag. Crit. Rev. 1: 429, 1817, nom. nov. Diphascum
Eaton = Diphyscium Ehr.
*Heterodon Raf., Med. Repos. II. 5: 350. 1808, nom.; Jour. Bot. (Paris) 2: 167.
1809, nom. = ? Fissidens Hedwig.
*bryoides Raf., 1. cc., descr. New Jersey.
ARNOLD ARBORETUM
Harvarp UNIVERSITY
1(2) : 263-328 FARLOWIA July, 1943
NOTES ON THE SYNONYMY OF SOME NORTH AMERICAN
THELEPHORACEAE AND OTHER RESUPINATES 1
D. P. Rocers anp H. S. Jackson
In the pages which follow it is proposed to record changes which ap-
pear to be necessary in the nomenclature of species in certain genera of
North American resupinate basidiomycetes, mainly in the Thelephoraceae,
which have come to our attention during the past eight or ten years.
Through correspondence and personal conference the authors recently
came to realize that they were independently covering much the same
ground and obtaining comparable results. It was, therefore, decided to
pool such of their findings to date as have to do with nomenclatorial
changes. This joint paper is the result.
The cosmopolitan distribution of many species, particularly in the
north temperate regions, has perhaps not been fully appreciated. Many
American species assumed to be endemic are found to be the same as well
known European species. The thelephoraceous flora of northeastern and
northwestern North America, for example, is surprisingly like that of
Sweden, Finland, and the mountainous region of central Europe; and the
mycobiota of central North America shows corresponding similarities to
that of the more western and southern parts of Europe. This situation
has made necessary a considerable number of changes in names now cur-
rent in America. Anyone who describes North American species as new
and has not an extensive knowledge of the European forms is taking a
grave risk of merely adding to the synonymy. Then, too, methods of
study which place disproportionate emphasis on the characters brought
out by the examination of cross sections, and which obscure or disregard
the more fundamental characters of basidia and spores and the more
detailed morphology of the hyphae, have often resulted in unnecessary
duplication of names. An object of this study has been to discover iden-
tities between European and American species, and between species de-
scribed from incomplete observations; by such reduction in the number
of current names the taxonomic confusion within the groups in question
is reduced, and the amount of information available concerning any one
form is increased.
Furthermore, any student who has devoted considered attention to taxo-
nomic study of the resupinate genera of Thelephoraceae is soon forced to
the realization that all too little regard has been given by former students
of the group to the exact application of nomenclatorial rules. Many of
the names in common use in Europe and in America are clearly synonyms
or are obviously untenable for one reason or another. A good deal of
current usage is contrary not only to the Rules but to non-legal reason.
* Joint contribution from the Laboratories of Cryptogamic Botany and the Farlow
Herbarium, Harvard University, No. 201, and the Department of Botany, University
of Toronto,
263
264, FarLowia, VoL. 1, 1943
In several cases the same specific name, with the same type, is found serv-
ing as the basis for binomials in two different genera. Even more com-
monly a binomial is ascribed to a mycologist who had no intention of
publishing a new taxonomic entity; but who was merely transferring, or
redescribing, an older species; frequently the later worker is said to have
established a type, even though he clearly cited the name and description
of the earlier. A second object of this study has been to determine, so
far as is possible, the correct name for the species treated; it is hoped
that these two objects, the taxonomic and the nomenclatorial, may con-
tribute to a more stable ultimate nomenclature.
To the close student of the group it is also evident that there is great
need for generic revision. At present it is obvious that in many instances
closely related species are classified in different genera. Real relation-
ships are often found to cut across family lines. Revision in this group
should, however, proceed slowly and progressively. No person as yet
possesses the wide and accurate knowledge of species which would be
necessary before any sweeping revision could be proposed. It is clear,
however, that certain groups of species, part of which may now be in-
cluded in one genus and part in another, show obvious relationship based
on characters believed to be of fundamental importance. As such groups
come to be well understood they should be segregated.
It is not the purpose of the present paper to attempt any revision. Cer-
tain segregates, largely from Corticium, which have already been proposed,
are here accepted and certain transfers not previously made are provided.
The segregates which are accepted include Ceratobasidium Rogers, Pelli-
cularia Cooke (= Botryobasidium Donk), Vararia Karsten (= Astero-
stromella Hohn. & Litsch.), and Trechispora Karsten (= Sistotrema sensu
Donk and Rogers p.p.). These genera, we feel, represent natural groups
of species.
The genus Gloeocystidium, commonly in use by many European stu-
dents, does not, as at present delimited, necessarily bring together related
species. We prefer to leave such species in Corticium for the present, as
Burt has done. However, since there is no sharp distinction between
gloeocystidia and cystidia, a number of species included by some authors
in Gloeocystidium but by others in Peniophora, e.g., P. tenuis (Pat.)
Massee, will here be found in the latter genus rather than in Corticium,
when their relationship seems to be with better characterized Peniophora
species. Similarly, several American species which have been described
in Peniophora, having projecting gloeocystidia but showing no marked
affinity to that genus (e.g., P. albo-straminea Bres.), have been transferred
to Corticium. While such procedure is recognized as merely a temporary
expedient, it serves to bring all definitely gloeocystidiate but non-cystidiate
forms together in one genus, with the recognition that that genus, like
some others at present, is merely a convenient repository.
Tomentella, as has already been pointed out by Rogers, must replace
Rocers & JAcKSON: SYNONYMY OF THELEPHORACEAE 265
Hypochnus sensu Burt. No attempt is made at this time to include trans-
fers for the American species which properly belong there. To do this
arbitrarily without a thorough revision of the genus as a whole would
only add unnecessary names to the ultimate synonymy. Since, however,
certain rough-spored species included in Hypochnus by Burt and others
are not Tomentellas, these are provided for in other genera.
The present work includes Sebacina of the Tremellaceae, Tulasnella and
Gloeotulasnella of the Tulasnellaceae, and in the Thelephoraceae the older
genera Aleurodiscus, Asterostroma, Coniophora, Hypochnus, Corticium
(in the broadest sense), and Peniophora. No attempt has yet been made
to examine the other genera critically and no claim is made to complete-
ness or finality in the genera mentioned. In Corticium and Peniophora,
for example, many of the southern and subtropical species have not been
reviewed and in the latter genus the section Coloratae has not been con-
sidered. We are including only such changes as have for one reason or
another come to our attention and concerning which we are in general
agreement. Further study of some of the species included may reveal
that we have erred on the side of ultraconservatism and it may later be
found desirable, when more abundant material is available for study, to
restore to specific rank certain species which are here treated as synonyms.
The present conservative treatment in such cases, however, has the ad-
vantage of emphasizing obviously close relationship.
The series of papers on North American Thelephoraceae prepared by
the late Dr. E. A. Burt constitutes the only monographic treatment of the
North American species of the Thelephoraceae available. Since Burt’s
treatise enumerates or (with a few exceptions) accounts for all the North
American names recorded previous to the dates of publication of the sev-
eral parts, this pioneer work is taken as the basis of the present contri-
bution. In the series of items making up the body of this paper the first
name in each paragraph heading is the name to be found in Burt if he
dealt with the species at all, and in most cases is followed by a second,
accepted name, separated by an = sign. In its proper place the accepted
name is printed in bold-face type.
To the corrections to names used by Burt have been added: (a) correc-
tions to names, new or old, which have been recorded for North America
since the publication of Burt’s account; (b) a few corrections involving
older American names which were not accounted for by Burt. Additions
to the North American flora are not included except as they may be inci-
dentally mentioned in connection with corrections to Burt’s identification
of paratypes or other cited collections. Similarly, additions to the synony-
my of American species are listed only if the species is included for
other reasons.
Burt included, in his monographic treatment of the Thelephoraceae,
the genus Sebacina of the Tremellaceae (along with other resupinate gen-
era) and Tulasnella of the Tulasnellaceae. These genera have since been
266 FarLowl1A, VoL. 1, 1943
re-monographed for North America — Sebacina by McGuire and Tulas-
nella by Rogers. In the present work, names in Sebacina have been
treated only where it has been possible to add to the results reported by
McGuire; for most of the species included in Burt the reader is referred
to McGuire’s recent monograph in Lloydia 4: 1-43. 1941; Tulasnella is
treated in the same way as outlined for the Thelephoraceae. Changes in
names used by Burt or changes found necessary for names used in other
later discussions are here brought together even though this involves some
repetition of information already available.
The arrangement is alphabetical by genera and species without refer-
ence to the families listed.
Authorities for species have been cited more fully than is customary or ,
ordinarily necessary. This is done in order that the citation may fulfill
its chief function, which is to set forth the history and especially the
typification of the species. To take an example from the synonymy listed
under Coniophora atrocinerea (p. 273): to write “Thelephora olivacea
Pers.,” as is allowable under the Rules (Art. 46, 48, Rec. XXXII), would
conceal the fact that the species was originally described by Fries and
should be interpreted after consultation of his treatment. To write: “T.
olivacea Fries ex Pers.,” as is probably also allowable, would imply that
Fries had described the species under Thelephora olivacea, whereas in
fact it was Persoon who first ascribed it to that genus. The form “T.
olivacea [Fries] ex Pers.” indicates: 1) that the species is a “pre-Friesian”’
one, first validly published by Persoon, and 2) that Fries described it in
another genus. The forms “Coniophora olivacea Karst.,” “C. olivacea
(Pers.) Karst.,” and “C. olivacea ([Fries] ex Pers.) Karst.” are the three
allowable forms of citation for the species when it is transferred to Conto-
phora; the most complete is the one used throughout the present paper,
although a shorter one would serve where it was desired to refer to the
fungus without entering upon a consideration of its history. “Coniophora
olivacea Fries” is a false citation: Fries never assigned the species to the
genus Coniophora; and “C. olivacea (Fries) Karst.” is equally incorrect,
since not Fries, but Persoon, first published the species, in the sense of
the Rules.
In their indispensible book “Hyménomycétes de France” Bourdot &
Galzin often used binomials for groups of fungi of subspecific rank. This
practise, not allowable under Art. 12 and Art. 28 of the Rules, has fre-
quently led to misunderstanding of the authors’ intent: later workers have
used as names of species binomials which Bourdot & Galzin published
for subspecies only. In the “Hyménomycétes de France” the authors
(p. iv) gave the explanation of their practise: the rank of a group is
indicated by typography, and instead of writing “subsp.” they printed
the name in slanted type. Thus, for example, Peniophora media, P. lurida,
and P. attenuata, published on p. 294, are not species, but subspecies of
P. subalutacea. In the series of papers in the Bulletin of the Société
Rocers & JACKSON: SYNONYMY OF THELEPHORACEAE 267
Mycologique de France which constitute their first general treatment of
the “Hyménomycétes de France” the same typographical system is used;
there, however, the rank of subspecies, varieties, and forms is usually
indicated also by the customary abbreviations. The precise concept or
definition of a subspecies held by Bourdot & Galzin need not be consid-
ered in evaluating the nomenclatorial status of their names, since it is
clear that the illegitimate binomials in question are intended for groups
of rank inferior to the species. In the present paper, therefore, such
names are written out in full so as to retain the rank assigned to them
by these authors — as, for example (on p. 274), “Coniophora arida subsp.
fumosa (Karst.) Bourd. & Galz.”
In the lists of synonyms which follow, the sign (!!) following the ref-
erence indicates that the type has been examined, while the sign (!) means
that an authentic specimen has been seen. Brackets ([ ]) around dates
are used where a presumably correct date has been substituted for that
carried by the title page of the work in question, and elsewhere, when
emphasis is considered desirable.
ACKNOWLEDGMENTS
The writers are greatly indebted to Dr. David Linder for having made
available, without restriction, the unique facilities of the Farlow Herba-
rium and Library. We are also under obligation to Mr. J. A. Stevenson,
Bureau of Plant Industry; Dr. F. J. Seaver, New York Botanical Garden;
Dr. L. O. Overholts, Pennsylvania State College; Dr. H. D. House, New
York State Museum; and to Drs. J. M. Greenman and C. W. Dodge of
the Missouri Botanical Garden, for the loan of types and other critical
collections from the herbaria under their care. From the late Professor
Victor Litschauer of Innsbruck, Austria, we have received much critical
information with reference to European species and he has freely shared
authentic materials from his herbarium.
It seems fitting also to acknowledge our indebtedness to the late Abbé
Bourdot for having provided, in Bourdot and Galzin’s “Hyménomycetes
de France,” the most useful summary of the species of lower basidiomy-
cetes that has yet appeared.
ALEURODISCUS
ALEURODISCUS APICULATUS Burt = A. mirabilis (Berk. & Curt.) Hohn.
Psilopezia mirabilis Berk. & Curt., Linn. Soc. Bot. Jour. 10: 364. [1868] (!!);
Massee, Jour. Mycol. 6: 179. 1891.
Cortictum peradeniae Berk. & Br., Linn. Soc. Bot. Jour. 14: 69. 1873.
Aleurodiscus javanicus P. Henn., Monsunia 1: 139, 1900 (!!).
Aleurodiscus peradeniae (Berk. & Br.) P. Henn., Monsunia 1: 139. 1900; Burt,
Missouri Bot. Gard. Ann. 10: 187. 1923.
Corticium javanicum (P. Henn.) Sacc. & Syd., Syll. Fung. 16: 189, 1902; nec
C. javanicum Zimm., Centralbl. Bakt. Parasit. 2 Abt. 7: 102. 1901.
Aleurodiscus usambarensis P. Henn., Bot. Jahrb. 34: 43. 1904 (!!).
Aleurodiscus spinulosus P. Henn., Bot. Jahrb. 38: 107. 1905 (!!).
268 Fartowia, VoL. 1, 1943
Corticium usambarense (P. Henn.) Sacc. & Sacc., Syll. Fung. 17: 168, 1905.
Aleurodiscus mirabilis (Berk. & Curt.) Hohn., K. Akad. Wiss. Wien Math.-Nat.
Kl. Sitzungsb. 118, I: 818. 1909.
Corticium spinulosum (P. Henn.) Sacc. & Trott., Syll. Fung. 21: 403, 1912.
Aleurodiscus apiculatus Burt, Missouri Bot. Gard. Ann. 5: 186, fig. 4. 1918 (!!).
Aleurodiscus albo-roseus Bres., Ann. Mycol. 18: 46. 1920 (!!).
Aleurodiscus Peteloti Pat., Soc. Mycol. Fr. Bul. 40: 31. 1924 (!!).
Aleurodiscus salmoneus Pat., Acad. Malgache Mem. 6: 11. pl. 1. 1927.
Aleurodiscus sinensis Teng & Ling, Contr. Biol. Lab. Sci. Soc. China 8: 273.
fig. 19. 1933.
Aleurodiscus pallide-roseus Litsch., Symb. Sinica II Theil, p. 41. fig. 1. 1937.
It is highly probable that the synonymy here given is not yet complete.
Burt’s figures are excellent, and a sufficient basis for identification. The
sublunate, biapiculate spores do not occur in any other species of Aleuro-
discus examined. The abundant, thick-walled, long-appendiculate acan-
thophyses are equally characteristic. Massee assigns P. mirabilis to
Aleurodiscus Oakesii (Berk. & Curt.) Cooke, from which, however, the
type differs in being attached for nearly its entire area and in the pos-
session of densely spinulose acanthophyses and in the absence of pseudo-
physes.
ALEurRopIscus crassus Lloyd = A. candidus (Schw.) Burt
Thelephora candida Schw., Nat. Ges. Leipzig Schrift. 1: 110. 1822 (!!); Fries,
Elenchus Fung. 1: 189. 1828; nec T. candida (Schw.) Fries, Syst. Orb. Veg.
p. 82. 1825; Schw., Am. Phil. Soc. Trans. n. s. 4: 166. [1832]. (= Merisma
candidum Schw., Nat. Ges. Leipzig Schrift. 1: 110. 1822; Tremellodendron
candidum (Schw.) Atk., Jour. Mycol. 8: 106. 1902.)
Thelephora candidissima Schw., Am. Phil. Soc. Trans. n. s. 4: 167. [1832].
Stereum candidum (Schw.) Fries, Epicr. Syst. Mycol. p. 552. 1838.
Aleurodiscus candidus (Schw.) Burt, Missouri Bot. Gard. Ann. 5: 188. 1918.
Aleurodiscus crassus Lloyd, Mycol. Writ. 6: 928. 1920 (!!).
Based on a single collection made in Oregon by H. C. Gilbert, A. crassus
is best interpreted as an unusually thick, broadly spreading condition of
the common A. candidus. The substratum is not stated but appears to
be the thick bark of some deciduous tree — probably Quercus Garryana,
the common substratum for the species in western Oregon. The type col-
lection agrees well with other western collections assigned to this species.
ALEURODISCUS CREMEUS Burt = A. Bertii Lloyd
Aleurodiscus cremeus Burt, Missouri Bot. Gard. Ann. 5: 199. 1918 (!!); nec
A. cremeus Pat., Soc. Mycol. Fr. Bul. 31: 73.1915 (!!) (= Stereum lepra Berk.
& Br., Linn. Soc. Bot. Jour. 14: 67. 1873 (!!); A. lepra (Berk. & Br..) Hohn.
& Litsch., K. Akad. Wiss. Wien Math.-Nat. KI]. Sitzungsb. 117, I: 1098. 1908).
Aleurodiscus Bertii Lloyd, Mycol. Writ. 7: 1288. 1924.
Aleurodiscus Burti Trott. in Sacc., Syll. Fung. 23: 525. 1925.
Because of the existence of the earlier homonym (cf. also Lloyd, l.c.),
Lloyd’s name must replace Burt’s. Unless an earlier description in an-
other genus exists, the species is probably autonomous. It is character-
ized by hard, compact structure, by gloeocystidia showing yellow, granular
Rocers & JAcKSON: SYNONYMY OF THELEPHORACEAE 269
content (in 1941), and acanthophyses whose outer portions show no
lumen whatever.
ALEuRODIScUS GranTu Lloyd = A. amorphus (Pers.) Rabenh. ex Cooke
Peziza amorpha Pers., Mycol. Eur. 1: 269. 1822.
Thelephora amorpha (Pers.) Fries, Elenchus Fung. 1: 183. 1828.
Corticium amorphum (Pers.) Fries, Epicr. Syst. Mycol. p. 559. 1838; Sacc., Syll.
Fung. 6: 606. 1888.
Nodularia balsamicola Peck, New York State Mus. Rept. 24: 96. pl. 4, fig. 23-26.
1872. :
Cyphella amorpha (Pers.) Quél., Ench. Fung. p. 215. 1886.
Aleurodiscus amorphus (Pers.) Rabenh. ex Cooke, Grevillea 3: 137. 1875;
Schroet. in Cohn, Krypt.-Fl. Schles. 3(1) : 429. 1888; Hohn. & Litsch., K. Akad.
Wiss. Wien Math.-Nat. KI]. Sitzungsb. 116, I: 799. 1907; Burt, Missouri Bot.
Gard. Ann. 5: 180. 1918; Pildt, Ann. Mycol. 24: 211. 1926; Bourd. & Galz.,
Hymén. de France p. 331. [1928].
Aleurodiscus Grantii Lloyd, Mycol. Writ. 6: 927. 1920 (!!).
(Aleurodiscus amorphus (Pers.) Rabenh., Fungi Eur, Exs. 1824. 1874 (nomen
nudum); Hedwigia 13: 184. 1874 (nomen nudum).
A. Grantii was based on two collections from Washington and one from
British Columbia. These have all been examined. The species was ad-
mittedly distinguished from A. amorphus on gross characters. Lloyd
writes: “Microscopic characters as in A. amorphus.” The differences
noted do not seem suflicient to justify specific segregation from a species
which, when a large series of collections is compared, shows considerable
variation in the form and margin of the fruiting bodies.
The genus Nodularia was unmistakably described and figured by Peck
in 1872. The name Aleurodiscus was first used, by Rabenhorst, in 1874;
but neither on the label of his exsiccatum nor in the summary of the same
in Hedwigia is there a word of description. The genus was therefore not
published by him. Cooke’s rambling discussion includes nothing that
can be considered a generic diagnosis. In it he pointed out the identity
of the validly published Nodularia and the undescribed Aleurodiscus —
and saw fit to retain the unpublished name. This choice, unwarranted as
it was by any information in his possession, and unjust as it was, consti-
tutes publication of the name Aleurodiscus under Art. 42 (2) of the Rules,
as has been pointed out by Donk (Bot. Gard. Buitenzorg Bul. III 17:
160. 1941.). Since there is a Nodularia Link, Schrad. Neu. Jour. Bot.
4: 9, 1809; Lyngbye, Tent. Hydrophytol. Dan. p. 99. 1819 (and a Nodu-
laria Mertens apud Juerg. Alg. Aquat. dec. XV, no. 4, 1822, which, being
a member of the Nostocaceae, was not nomenclatorially existent in 1872),
Cooke’s wrong choice proves to be the correct one to follow. It is curious
that the various monographers who, aware of the identity of Nodularia
Peck and Aleurodiscus, employed the later name, never felt bound to
give any reasons for doing so.
ALEURODISCUS HELVEOLUS Bres. Excluded.
This species, originally described by Bresadola (Mycologia 17: 71.
1925 (!!); Burt, Missouri Bot. Gard. Ann. 13: 306. 1926) from a single
270 | Fartowia, VoL. 1, 1943
collection made by J. R. Weir on Salix lasiandra at Spokane, Washington,
November 20, 1920 (Weir 16312), may be stricken from the list of
American species of Alewrodiscus on the grounds that its real relationship
is with the fungus which, in North America, is commonly referred to
Clavaria contorta Holmsk. ex Fries. This fungus is common on Salix in
the Temagami Lake region, Ontario, and specimens are in the herbarium
of the University of Toronto which bear some fruiting bodies which agree
very well with the type of Bresadola’s species, the fructifications of which
are more tuberculate and less well developed than in most collections.
The basidia and spores as well as other microscopic characters seem iden-
tical. Coker (The Clavarias of the United States and Canada p. 87.
1923.) places C. contorta as a doubtful synonym of C. fistulosa Holmsk.
ex Fries.
ALEuROpIscus succINEUS Bres. = A. diffissus (Sacc.) Burt
Peniophora diffissa Sacc., R. Soc. Bot. Belg. Bul. 28: 79. pl. 4, fig. 2. 1889;
Syll. Fung. 9: 239. 1891.
Aleurodiscus succineus Bres., Mycologia 17: 71. 1925 (!!); Burt, Missouri Bot.
Gard, Ann. 13: 309. 1926.
Aleurodiseus diffissus (Sacc.) Burt, Missouri Bot. Gard. Ann, 18: 485. 1931;
Pilat, Soc. Mycol. Fr. Bul. 49: 328. 1933.
Aleurodiscus sajanensis (Murashk. in litt.) Pilat, Hedwigia 71: 328. fig. 1-3.
1931 (11).
A study of the literature listed above shows that only three collections
are involved. Two of them were made in Siberia on Rhododendron
dahuricum and the third, on which A. succineus was based, is reported as
on Arbutus Menziesii from Oregon. In the latter the fructifications are
evidently younger than those in Murashkinsky’s collection on which Burt’s
transfer and Pilat’s species were based. There is no question, however,
that the two are identical. The species is readily distinguished by the
abundant narrow acanthophyses which become yellowish with age, the
presence of gloeocystidia and the sub-allantoid spores. Burt (1926) de-
scribes the spores of A. swccineus as ellipsoid 10 x 5 » and (1931) those
of the Siberian collection as globoid 5.5-6 » (!). They are more nearly
as described by Pildt (1931) 10-15 x 4-5.5 y, though spores as large
as 18 x 7 p» are present in the Murashkinsky collection examined.
ALEuROpISCcUS ZELLERI Burt = Peniophora aurantiaca (Bres.) Hoéhn. & Litsch.
Corticium aurantiacum Bres., Fungi Trident. 2: 37. 1892 (!!).
Kneiffia aurantiaca (Bres.) Bres., Ann. Mycol. 1: 104. 1903.
Peniophora aurantiaca (Bres.) Héhn. & Litsch., K, Akad. Wiss. Wien Math.-Nat.
Kl. Sitzungsb. 115, I: 1586. 1906; Burt, Missouri Bot. Gard. Ann, 12: 310.
[1926]; Bourd, & Galz., Hymén. de France p. 320. [1928].
Peniophora lepida Bres., Mycologia 17: 70. 1925 (!!); Burt, Missouri Bot. Gard.
Ann. 12: 295. [1926].
Peniophora Sheari Burt, Missouri Bot. Gard. Ann. 12: 268. [1926] (!!).
Aleurodiscus Zelleri Burt, Missouri Bot. Gard. Ann. 13: 309. 1926 (!!).
A. Zelleri was based on a tuberculate or at most only narrowly effused
form of P. aurantiaca not infrequent in the northwest. The type of P.
Rocers & JAcKSON: SYNONYMY OF THELEPHORACEAE 271
lepida is reported as growing on Salix; actually, the wood is Alnus. It
is best interpreted as an unusually vigorous development of P. awrantiaca
with all the essential structures, but lighter colored than usual, and sterile.
P. Sheari is, like P. lepida, a thick fructification with very numerous
heavily incrusted cystidia; its spores are more nearly oblong, or cylindri-
cal, than in typical specimens of P. awrantiaca, but are related by inter-
grades with the more usual ellipsoid form.
ASTEROSTROMA
ASTEROSTROMA BICOLOR Ellis & Everh. = A. andinum Pat.
Asterostroma andinum Pat., Soc. Mycol. Fr. Bul. 9: 133. 1893 (!!).
Asterostroma bicolor Ellis & Everh., Acad. Nat. Sc. Phila. Proc. 1893: 441. 1894
(!!); Burt, Missouri Bot. Gard. Ann. 11: 32. 1924.
Asterostroma spiniferum Burt, Missouri Bot. Gard. Ann. 11: 33. 1924 (!!).
Asterostroma gracile Burt, Missouri Bot. Gard. Ann. 11: 34. 1924 (!!).
The type of A. gracile shows a very delicate loose-arachnoid hymenium;
in the types of A. andinum and A. spiniferum the hymenium is continuous,
but branches of the stellate bodies emerge, appearing like the setae of a
Hymenochaete. In none of these are the asterophyses different from those
of A. bicolor; in all they are long-stalked, long-rayed (up to at least 125 p
in A. andinum and A. spiniferum, at least 150 » in A. bicolor) and rela-
tively slender-rayed, often somewhat asymmetrical, commonly with fork-
ing or uneven branching. In the preparations examined, subulate, cysti-
dioid bodies were observed in the types of A. bicolor, A. andinum, and
A. gracile, but not in A. spiniferum, where Burt reports them; probably
further search would have demonstrated them there also. There are no
differences other than those here noted; presumably A. bicolor is the name
given to well developed specimens, the other names referring to younger
fructifications.
ASTEROSTROMA GRACILE Burt = A. andinum Pat.
See under A. bicolor.
ASTEROSTROMA OCHROSTROMA Burt = Asterodon ferruginosus Pat.
Asterodon ferruginosus Pat., Soc. Mycol. Fr. Bul. 10: 130. 1894 (!!) (ut ‘fer-
ruginosum’) ; Bourd. & Galz., Hymén. de France p. 398. [1928].
Hydnochaete setigera Peck, New York State Mus. Rept. 50: 113. 1897 (!).
Asterostroma ochrostroma Burt, Missouri Bot. Gard. Ann. 11: 34. 1924 (!!).
Asterostroma ochrostroma was described from a sterile, immature fruc-
tification of Asterodon ferruginosus. There is considerable doubt whether
Asterodon ought to be maintained as a separate genus; but the matter lies
outside the scope of this paper. For the gender of names in -odon see
Art. 72(2) or any Greek lexicon.
Laurila (Soc. Zool.-Bot. Fenn. Vanamo Ann. Bot. 10 (4): 15. 1939.)
gives “Aciella tomentosa (Schrad.) Karst.” as a synonym of A. ferrugino-
sus. His citation is obscure, but probably refers to Acia (Aciella) tomen-
272 FarLowia, VoL. 1, 1943
tosa (Schrad.) Karst. (Bidr. Kanned. Finl. Nat. Folk 48: 362. 1889.), a
name by which Karsten replaced the earlier Acia ferruginea (Pers.) Karst.
Deferring consideration of a remarkable collection of nomenclatorial
complexities, one may well suspend judgment concerning the propriety of
superseding Patouillard’s specific epithet until it appears at least probable
that Schrader’s Hydnum tomentosum referred to the present fungus rather
than to a Caldesiella. As for Hydnum ferrugineum Pers., Fries very gen-
erously disposed of that name by reducing it to synonymy under a newly
erected H. ferruginosum Fries, and by describing a stipitate H. ferrugin-
eum Fries by which Persoon’s name is now preoccupied.
ASTEROSTROMA SPINIFERUM Burt — A. andinum Pat.
See under Asterostroma bicolor Ellis & Everh.
BOTRYOBASIDIUM
Botryospasipium Donk — Pellicularia M. C. Cooke
The American species of Botryobasidium were first brought together by
Rogers (Univ. Iowa Studies Nat. Hist. 17: 10-19. 1935.). In a more
recent paper (Iarlowia 1: 95-118. 1943.) the species of this group have
again been reviewed; since the generic name Pellicularia Cooke ante-
dates Botryobasidium Donk they have of necessity been treated under
Pellicularia, For the sake of completeness and to conform to the stated
scope of the present contribution (p. 264), the new names are listed
below. References to other names which have been used in American
literature will be found in their proper alphabetical position in the fol-
lowing pages.
Botryobasidium coronatum (Schroet.) Donk = Pellicularia pruinata (Bres.) Rogers.
Botryobasidium flavescens (Bon.) Rogers = Pellicularia flavescens (Bon.) Rogers.
Botryobasidium isabellinum (Fries) Rogers = Pellicularia isabellina (Fries) Rogers.
Botryobasidium ochraceum (Massee) Donk = Pellicularia flavescens (Bon.) Rogers.
Botryobasidium Solani (Prill. & Del.) Donk = Pellicularia filamentosa (Pat.)
Rogers.
Botryobasidium subcoronatum (Hohn. & Litsch.) Donk — Pellicularia subcoronata
(Hohn. & Litsch.) Rogers.
Botryobasidium vagum (Berk. & Curt.) Rogers = Pellieularia vaga (Berk. & Curt.)
Rogers.
CERATOBASIDIUM
CERATOBASIDIUM PLUMBEUM Martin = C, atratum (Bres.) Rogers
Corticium atratum Bres., Hedwigia 35: 290. 1896 (!); Rick, Brotéria Sér. Trim.
Ciénc. Nat. 3: 154. 1934.
- Tulasnella metallica Rick, Brotéria Sér. Trim. Ciéne. Nat. 3: 169. 1934 (!), saltem
pro parte.
Ceratobasidium plumbeum Martin, Mycologia 31: 513. fig. 21-27. 1939. (!!).
Ceratobasidium atratum (Bres.) Rogers ex Martin, Lloydia 4: 262. 1941.
It would be difficult to imagine a resupinate species more distinctive
than the present one, or more complete neglect of its characteristics than
that exhibited in the description of C. atratum. It is not a “species valde
Rocers & JACKSON: SYNONYMY OF THELEPHORACEAE 208
Corticio caeruleo affinis” (Bres.), nor does it show any similarity what-
ever to that fungus. It is well and fully described and figured by Martin
as Ceratobasidium plumbeum.
The species has a wide distribution in the western hemisphere, as noted
by Martin (1941). It appears to be quite common in northeastern North
America, seven collections having been made in Ontario. Collections
from Quebec and California have recently been recognized as this species.
Authentic material of 7. metallica Rick includes both the present species
and Peniophora rimicola (Karst.) Hohn. & Litsch.; from Rick’s descrip-
tion it seems probable that originally he had in hand the Ceratobasidium
rather than the Peniophora.
CONIOPHORA
CONIOPHORA ATROCINEREA Karst. — C. olivacea ([Fries] ex Pers.) Karst.
Thelephora olivacea [Fries] ex Pers., Mycol. Eur. 1: 143, 1822 (ante diem 1 Jul.) ;
Fries, Elenchus Fung. 1: 197. 1828.
Thelephora Sistotremoides Schw., Nat. Ges. Leipzig Schrift. 1: 109. 1822 (fortasse
post diem 18 Jul.) (!!); Fries, Elenchus Fung. 1: 198. 1828.
Thelephora umbrina Fries, Elenchus Fung. 1: 199. 1828; nec 7. umbrina Pers.,
Syn. Fung. p. 578. 1801.
Odontia Sistotremoides (Schw.) Fries, Epicr. Syst. Mycol. p. 529. 1838.
Hypochnus olivaceus ({Fries] ex Pers.) Fries, Summa Veg. Scand. p. 337. 1849;
Karst., Bidr. Kanned. Fin]. Nat. Folk 25: 320. 1876; et conf. Bres., I. R. Accad.
Agiati Atti IIT 3: 116. 1897; Rogers, Mycologia 31: 299. 1939.
Hypochnus umbrinus (Fries) Fries, Summa Veg. Scand. p. 337. 1849; Quél., Fl.
Mycol. p. 2. 1888, quantum ad typum; Burt, Missouri Bot. Gard. Ann. 3: 213.
1916, 13: 323. 1926, quantum ad typum, specim. descr. synon. excl.; Wakef.,
Br. Mycol. Soc. Trans. 6: 132. 1919, tantundem; nec H. umbrinus (Pers.)
Wallr., Fl. Crypt. Germ. 2: 312. 1833 (= Himantia umbrina Pers., Syn. Fung.
p. 704. 1801; Alb. & Schw., Consp. Fung. p. 373. 1805; Pers., Mycol. Eur. 1:
89. 1822.).
Corticium brunneolum Berk. & Curt., Grevillea 2: 4. 1873 (!!).
Corticium leucothrix Berk. & Curt., Grevillea 2: 4. 1873 (!!).
Corticium olivaceum ([Fries] ex Pers.) Fries, Hymen. Eur. p. 660. 1874.
Hymenochaete Ellisii Berk. & Cooke, Grevillea 4: 162. 1876; Hohn. & Litsch., K.
Akad. Wiss. Wien Math.-Nat. Kl]. Sitzungsb. 116, I: 782. 1907.
Corticium Ellisii (Berk. & Cooke) Cooke, Grevillea 8: 89. 1880.
Coniophora atrocinerea Karst., Soc. Faun. Fl. Fenn. Med. 6: 12. 1881, et apud
Thiim. Myc. Univ. 1806. 1881 (!!); Burt, Missouri Bot. Gard. Ann. 4: 260. 1917.
Coniophora fumosa Karst., Soc. Faun. Fl. Fenn. Med. 6: 13. 1881 (!!) (“Corti-
cium fumosum Fr. pr.p.”; num typus ?); nec Coniophora fumosa sensu Massee,
Linn. Soc. Bot. Jour. 25: 139. 1889.
Coniophora umbrina (Fries) Karst., Bidr. Kanned. Finl. Nat. Folk 37: 160. 1882;
Burt, Missouri Bot. Gard. Ann. 4: 256. 1917.
Coniophora olivacea ([Fries] ex Pers.) Karst., Bidr. Kanned. Finl. Nat. Folk
37: 162. 1882; Burt, Missouri Bot. Gard. Ann. 4: 257, 1917.
Coniophora furva Karst., Bidr. Kanned. Finl. Nat. Folk 37? 244. 1882 (!!).
Coniophora brunneola (Berk. & Curt.) Sacc., Syll. Fung. 6: 648. 1888.
Coniophora Ellisii (Berk. & Cooke) Sacc., Syll. Fung. 6: 648. 1888.
Coniophora leucothrix (Berk. & Curt.) Sacc., Syll. Fung. 6: 648. 1888.
Coniophorella atrocinerea (Karst.) Karst., Bidr. Kanned. Finl. Nat. Folk 48: 438.
1889; Bourd. & Galz., Hymén. de France p. 364. [1928].
274 Fartowia, VoL. 1, 1943
Coniophorella olivacea ([Fries] ex Pers.) Karst., Bidr. Kanned. Finl. Nat. Folk
48: 438. 1889; Bourd. & Galz., Hymén. de France p. 362. [1928].
Coniophora Sistotremoides (Schw.) Massee, Linn. Soc. Bot. Jour, 25: 133. 1889;
Burt, Missouri Bot. Gard. Ann. 4: 249. 1917; 13: 312. 1926, quantum ad typum.
Coniophora fulvo-olivacea Massee, Linn. Soc. Bot. Jour. 25: 134. 1889 (!!) ; Karst.,
Hedwigia 30: 300. 1891.
Coniophora Karstenii Massee, Linn. Soc. Bot. Jour. 25: 134. 1889 (!!).
Coniophorella umbrina (Fries) Bres., Ann Mycol. 1: 111. 1903; Bourd. & Galz.,
Hymén. de France p. 363. [1928]; Laurila, Soc. Zool.-Bot. Fenn. Vanamo Ann.
Bot. 10(4): 13. 1939,
Coniophora arida subsp. fumosa (Karst.) Bourd. & Galz., Soc. Mycol. Fr. Bul. 39:
113. 1923; Hymeén. de France p. 359, [1928], quantum ad typum.
Coniophorella olivacea subsp. fulvo-olivacea (Massee) Bourd. & Galz., Soc. Mycol.
Fr. Bul. 39: 116. 1923; Hymén. de France p. 362. [1928].
Coniophora sibirica Burt, Missouri Bot. Gard. Ann. 18: 485. 1931 (!!); Pilat,
Soc. Mycol. Fr. Bul. 49; 296. 1933.
Tomentella umbrina (Fries) Litsch., Soc. Mycol. Fr. Bul. 49: 52. 1933, quantum ad
typum; Donk, Neder]. Mycol. Ver. Med. 22: 29. (= Bot. Mus. Herb. Univ.
Utrecht Med. 9: 29.) 1933, tantundem.
Coniophorella umbrina var. olivacea ({Fries] ex Pers.) Rick, Brotéria Sér. Trim.
Ciénc. Nat. 3: 167. 1934.
(Thelephora umbrina Pers, var. lignatilis Alb. & Schw., Consp. Fung. p. 281. 1805.
(Hypochnus olivaceus Fries, Obs. Mycol. 2: 282. 1818.
(Coniophora fusca Karst. ex Thiim., Myc. Univ. 2112. 1883 (nomen nudum) ; nec
C. fusca sensu Massee, Linn. Soc. Bot. Jour. 25: 139. 1889.
As Laurila has pointed out, C. atrocinerea, C. olivacea and C. umbrina
are only stages in the development of a single fungus. In C. atrocinerea
the color is relatively grayish (lighter than Buffy Brown (R) in the type)
and the cystidia are still short and have mostly hyaline or pallid, narrow,
unincrusted distal portions (cf. Burt’s figure) ; a few, however, are nearly
mature and dark and broad through much of their length. In C. olivacea
the color is darker and less ashy (Isabella Color (R) in a specimen from
Bresadola, between that and Tawny Olive (R) in one from Karsten),
partly from darkening of the hymenial elements, and partly from the ac-
cumulation of the spores; the cystidia are mature through most of their
length, although the tips of the majority are still growing, and still narrow
(cf. Burt’s figure). In C. umbrina the hymenium is still darker (Sac-
cardo’s Umber (R) in a specimen from Bresadola), and the cystidia
mostly dark throughout and broadened and spatulate at the ends (cf. Burt’s
figure). Finally, in C. Sistotremoides (type) and C. Karstenii (type),
the cystidia are not only completely darkened but immersed. There are
no differences with respect to spores, basidia, or mycelium, nor in the
mature portions of the cystidia, except in C. sibirica, where the spores are
a little smaller and the basal mycelium darker than in most material.
Perhaps the majority of collected specimens fall between C. olivacea and
C. umbrina. The affinities of the present fungus with species of Conio-
phora lacking cystidia are very great, and the segregation of Coniophor-
ella, of which it is the type, is purely an artificial and mechanical treatment.
Bresadola has attempted to base two species, a Coniophora and a “Hy-
Rocers & JACKSON: SYNONYMY OF THELEPHORACEAE BES
pochnus,” on T. olivacea; as has been pointed out by Rogers (l.c.), the
description in Fries’s “Elenchus,” and to a lesser extent also the first valid
publication, in Persoon, apply only to the Coniophora. A number of
authors have likewise attempted to base two species on 7’. umbrina. Thus
Burt has a “Hypochnus umbrinus (Fries) Burt” and a “Coniophora um-
brina Alb. & Schw. ex Fries,” both based on a description by Fries (Elen-
chus Fung 1: 199, 1828.) and both having listed as a synonym Corticium
umbrinum Fries (Hymen. Eur. p. 658. 1874.). Aside from the absurdity
of this double treatment, and from the fact that H. umbrinus had already
been published by three different authors, the first time at least for quite
another fungus, objection to such taxonomy can be made on the grounds
of the original description, which, reading “Sporidiis sordide ferrugineis
. . . Hine optima Coniophora. Observantur insuper sub lente setulae
fuscae . . .” can apply only to the present species. Furthermore, it is not
correct to attribute this 7. umbrina to Albertini & Schweinitz; their T. um-
. brina, like all the species for which they gave no authority, is the species
of Persoon (cf. Consp. Fung. p. vi, vii). To T. umbrina Pers., described
as terricolous, they added a lignicolous variety which Fries cited under
T. umbrina, “nec T. umbrina Pers.” (Elenchus Fung. 1: 199. 1828.) —
that is, the variety is “Alb. & Schw.,” but the species is “Fries” as Fries
himself wrote it in the index to the “Systema.”
CONIOPHORA AVELLANEA Burt = Corticium confluens Fries
Thelephora confluens Fries, Syst. Mycol. 1: 447. 1821.
Corticium confluens (Fries) Fries, Epicr. Syst. Mycol. p. 564. 1838; Burt, Mis-
souri Bot. Gard. Ann. 13: 220. 1926; Bourd. & Galz., Hymén. de France p. 212.
[1928], cum synon. plur.
Hypochnus confluens (Fries) Bon., Handb. p. 159. 1851, quantum ad typum.
Coniophora avellanea Burt, Missouri Bot. Gard. Ann. 4: 251. 1917 (!!).
Corticium rubellum Burt, Missouri Bot. Gard. Ann. 13: 232. 1926 (!!).
Burt was apparently led to describe this species as a Coniophora by the
circumstance that its spore-print is somewhat colored. Corticium rubellum
is based on a fructification of a strong rosy color, such as is not uncommon
in this variable fungus. See Bourdot’s discussion of coloration (l.c.).
CONIOPHORA BYSSOIDEA (Pers. ex Fries) Karst. = Peniophora byssoides (Pers. ex
Fries) Bres. ;
Thelephora byssoides Pers. ex Fries, Syst. Mycol. 1: 452. 1821.
Athelia strigosa Pers. 8 muscigena Pers., Mycol. Eur. 1: 83. 1822.
Hypochnus strigosus 8 filamentosus Wallr., Fl. Crypt. Germ. 2: 312. 1833.
Corticium lacunosum Berk. & Br., Ann. Mag. Nat. Hist. IV 11: 343. 1873 (!!).
Corticium byssoideum (Pers. ex Fries) Fries, Hymen. Eur. p. 659. 1874.
Hypochnus byssoideus (Pers. ex Fries) Quél., Soc. Bot. Fr. Bul. 26: 231. 1879.
Coniophora byssoidea (Pers. ex Fries) Karst., Bidr. Kanned. Finl. Nat. Folk 37:
160. 1882; Burt, Missouri Bot. Gard. Ann. 4: 263. 1917.
Hypochnus muscorum Schroet. in Cohn, Krypt.-Fl. Schles. 3(1) : 418. 1888 (!).
Hypochnus setosus Schroet. in Cohn, Krypt.-Fl. Schles. 3(1): 418. 1888 (!).
Tomentella obducens Karst., Bidr. Kanned. Finl. Nat. Folk 48: 421. 1889 (!).
Diplonema sordescens Karst., Bidr. Kanned. Finl. Nat. Folk 48: 430. 1889 (!).
276 Fartowia, VoL. 1, 1943
Peniophora sordescéns (Karst.) Sacc., Syll. Fung. 9: 240. 1891.
Hypochnus obducens (Karst.) Sacc., Syll. Fung. 9: 243. 1891.
Amphinema sordescens (Karst.) Karst., Hedwigia 32: 61. 1893.
Peniophora byssoides (Pers. ex Fries) Bres. apud Brinkm., Westf. Prov.-Ver.
Jahresber. 26: 130. 1898 (falso ut “byssoidea”) ; Hohn. & Litsch., K. Akad.
Wiss. Wien Math.-Nat. Kl]. Sitzbungsb. 117, I: 1084. 1908; Bourd. & Galz.,
Hymén. de France p. 297. [1928].
Zygodesmus pubidus Ellis & Everh., Torr. Bot. Cl. Bul. 27: 50. 1900 (!!).
Kneiffia Tomentella Bres., Ann. Mycol. 1: 103. 1903.
Coniophorella byssoidea (Pers. ex Fries) Bres., Ann. Mycol. 1: 111. 1903.
Peniophora muscorum (Schroet.) Hohn., Ann. Mycol. 3: 325. 1905.
Peniophora byssoidea subsp. Tomentella (Bres.) Bourd. & Galz., Soc. Mycol. Fr.
Bul. 28: 391 [1913]; Hymén. de France p. 297. [1928].
Kneiffia muscorum (Schroet.) Rick, Brotéria Sér. Trim. Ciéne. Nat. 3: 72. 1934.
This species is a member of Coniophora only if that genus be arbi-
trarily defined so as to include all resupinate Thelephoraceae with smooth
tinted spores — such as Corticium confluens Fries, Peniophora incarnata
(Pers. ex Fries) Karst. and its relatives, and species of Alewrodiscus —a
sense in which no one has ever used it, and as artificial a generic concept
as could be imagined.
CONIOPHORA CEREBELLA Pers. = C. puteana (Schum. ex Fries) Karst.
Thelephora puteana Schum. ex Fries, Syst. Mycol. 1: 448. 1821.
Coniophora cerebella Pers., Mycol. Eur. 1: 155. 1822; Burt, Missouri Bot. Gard.
Ann. 4: 240. 1917; Bourd. & Galz., Hymén. de France p. 358. [1928].
Corticium puteanum (Schum. ex Fries) Fries, Hymen. Eur. p. 657. 1874.
Coniophora puteana (Schum. ex Fries) Karst., Sallsk. Faun. Fl. Fenn. Not.
n. s. 6: 370. 1868.
To judge from the synonymy given by most authors (there being no
material at hand to determine what Persoon, Schumacher, or Fries really
had), Coniophora puteana is the correct name here; both Burt and Bourdot
acknowledge the equivalence of C. puteana and C. cerebella. Many more
names have been given to the largely sterile fungus as it occurs on
worked timbers.
ConIOPHORA CORRUGIS Burt = Corticium corruge (Burt) Burt in sched.
Coniophora corrugis Burt, Missouri Bot. Gard. Ann. 13: 310. 1926 (!!).
(Corticium corruge Burt in sched., in Baker, Pacific Slope Fungi 3570, nomen
nudum; Overh., Mycologia 11: 249. 1919, nomen nudum.
Corticium corruge (Burt) Burt in sched., nom. ined.
This characteristic species is best considered a Corticium with vernicose-
pellicular hymenium over a byssoid subiculum. It is probably a member
of Bourdot & Galzin’s section Membranacea. Typical, true species of
Coniophora have a thick, firm, often lamellate spore-membrane, and an
apiculus more or less peg-like in appearance and distinct from the adjacent
wall, as though driven in from outside. A hyaline-spored Coniophora is
quite conceivable. Although its spores are nearly always hyaline, this
is not such a species.
-
Rocers & JacKSON: SYNONYMY OF THELEPHORACEAE Pa
CONIOPHORA CORTICOLA Overh. = C. alboflavescens (Ellis & Everh.) Héhn. & Litsch.
Corticium alboflavescens Ellis & Everh., Acad. Nat. Sci. Phila. Proc. 1893: 324.
1894 (!1).
Coniophora alboflavescens (Ellis & Everh.) Hohn. & Litsch., K. Akad. Wiss.
Wien Math.-Nat. K]. Sitzungsb. 116, I: 791. 1907.
Coniophora corticola Overh., Mycologia 30: 274. 1938 (!!).
C. alboflavescens was included by Burt as a synonym under Coniophora
Polyporoidea (Berk. & Curt.) Burt (q.v.). It is amply distinct and a
good Coniophora, while C. Polyporoidea is not. It is well described by
Overholts (l.c.) as C. corticola and is readily recognizable by the globose
spores. The spores of Coniophora Harperi Burt, which are also globose,
are much smaller; those of Coniophora mustialaensis (Karst.) Massee are
blue (see next item).
CoNIOPHORA CYANOSPORA Rogers = C. mustialaensis (Karst.) Massee
@
Hypochnus mustialaensis Karst., Sallsk. Faun. Fl. Fenn. Not. n. s. 8: 222. 1871 (!!).
Corticium mustialaense (Karst.) Fries, Hymen. Eur. p. 705. 1874.
Lyomyces mustialensis [!] (Karst.) Karst., Bidr. Kanned. Finl. Nat. Folk 37:
154. 1882.
Hypochnopsis mustialensis (Karst.) Karst., Bidr. Kanned. Finl. Nat. Folk 48:
442, 1889.
Coniophora mustialaensis (Karst.) Massee, Linn. Soc. Bot. Jour. 25: 139. 1889.
Coniophora cyanospora Rogers, Univ. Iowa Studies Nat. Hist. 17: 25. 1935 (!!).
It is difficult to imagine how Karsten’s species could have been compared
by Bresadola (Ann. Mycol. 9: 426. 1911.) with any of the forms which
pass as Odontia, and even more difficult to see it, as there listed, in the
company of Corticium sulphurellum Hohn. & Litsch., which is waxy,
closely adnate, clear yellow, and hyaline-spored. It differs from Calde-
siella viridis (Alb. & Schw. ex Fries) Pat., to which Bourdot assigned it
(Hymén. de France p. 409. [1928].), in the smooth spore walls and in
other respects. Hydnum Sobolewskii Weinm. (Flora 15: 452. 1832.)
may well be this species; but the description provides no assurance that
the superficially similar Caldesiella viridis is not the fungus referred to.
Amaurodon viridis sensu Schroet. (in Cohn, Krypt.-Fl. Schles, 3 (1):
461. 1888) is probably this species rather than the Caldesiella.
With age the darker parts of the fructifications in this species become
Grape Green (R) to Vetiver Green (R), the margin remaining about
Chartreuse Yellow (R).
CoONIOPHORA DRYINA (Berk. & Curt.) Massee = Peniophora dryina (Berk. & Curt.)
comb. noy.
Corticium dryinum Berk. & Curt., Grevillea 1: 179. 1873 (!!).
Xerocarpus laeticolor Karst., Bidr. Kanned. Finl. Nat. Folk 37: 137. 1882 (!).
Coniophora crocea Karst., Rev. Mycol. 9: 10. 1887 (!).
Corticium laeticolor (Karst.) Sacc., Syll. Fung. 6: 636. 1888.
Coniophora laeticolor (Karst.) Karst., Bidr. Kanned. Finl. Nat. Folk 48: 436.
1889; Burt, Missouri Bot. Gard. Ann. 4: 261. 1917.
Coniophora dryina (Berk. & Curt.) Massee, Linn. Soc. Bot. Jour. 25: 135. 1889;
Burt, Missouri Bot. Gard. Ann. 4: 253. 1917.
278 Fartowia, Vou. 1, 1943
Peniophora crocea (Karst.) Hohn. & Litsch., K. Akad. Wiss. Wien Math.-Nat. K1].
Sitzungsb. 115, I: 1574. 1906; Bourd. & Galz., Hymén. de France p. 278 [1928].
Peniophora tabacina Burt, Missouri Bot. Gard. Ann. 12: 334. [1926] (!!).
Coniophorella laeticolor (Karst.) Pilat, Soc. Mycol. Fr. Bul. 49: 296. 1933.
Peniophora dryina (Berk. & Curt.) comb. nov.
The external appearance of the fungus is that of a Coniophora, but the
structure of both spores and basidiocarp is quite different, and not essen-
tially different from that of various fungi now assigned to Pentophora.
The spores in particular are thin-walled and with an apiculus quite differ-
ent from the peg-like structure of a typical Coniophora. Except in older
specimens, where most of the elements are somewhat stained, the spore-wall
is colorless, such tint as is present arising from the brassy yellow color of
the contents. There is no important difference in this respect between
specimens assigned by Burt to Peniophora tabacina and those assigned
to Coniophora laeticolor. Fresh material of the species showed only
spores with colorless walls; presumably old embedded spores, even in
living specimens, would become somewhat stained.
The species is apparently very rare in Europe; neither Massee nor
Hohnel & Litschauer nor Bourdot & Galzin report other material than
Karsten’s two types. In addition to those listed by Burt, American speci-
mens have been seen from Virginia, Missouri, Idaho, and Ontario, and
in some numbers from Oregon.
ConiopHora FLAVA Burt = Peniophora flava (Burt) comb. nov.
Coniophora flava Burt, Missouri Bot. Gard. Ann, 4: 261. 1917 (!!).
Peniophora flava (Burt) comb. nov.
Readily recognizable, as Burt has pointed out, by the brilliant color,
separable membranous texture, and incrusted cystidia. These organs are
somewhat unusual in being quite thin-walled throughout, although in-
crusted at maturity, — not the common situation. They arise as more
or less ventricose gloeocystidium-like bodies, and become fusiform with
age. The granular coloring material has a brassy-yellow cast like that
of Peniophora dryina in some phases. The spores are thin- and appar-
ently hyaline-walled, with yellow resinous inclusions, 4.5-6 X 3.5 p.
CONIOPHORA FLAVOMARGINATA Burt = Mycoacia Himantia (Schw.) Miller & Boyle
Clavaria byssacea Roth ex Pers., Mycol. Eur. 1: 172. 1822; Bourd. & Galz., Soc.
Mycol. Fr. Bul. 26: 216. 1910. :
Hydnum Himantia Schw., Nat. Ges. Leipzig Schrift. 1: 104. 1822.
Hydnum subfuscum Peck, New York State Mus. Rept. 40: 55. 1887.
Odontia Himantia (Schw.) Bres., Ann, Mycol. 1: 85. 1903; Brown, Bot. Gaz. 96:
665. 1935.
Coniophora flavomarginata Burt, Missouri Bot. Gard. Ann. 13: 311. 1926 (!!).
Clavaria Himantia (Schw.) Bourd. & Galz., Hymén. de France p. 122. [1928].
Oxydontia Himantia (Schw.) Miller, Mycologia 25: 363. 1933.
Mycoacia Himantia (Schw.) Miller & Boyle, Univ. Iowa St. Nat. Hist. 18:
44, 1943.
(Clavaria byssacea Roth, Usteri Ann. Bot. 1: 11. pl. 1, fig. 5. 1791.
Rocers & JACKSON: SYNONYMY OF THELEPHORACEAE 279
Burt’s fungus has spore-walls thin and colorless or only slightly yellow-
ish, and is certainly not a Coniophora. It is very common throughout the
autumn and winter in western Washington and Oregon, growing in chinks
in the bark of standing trees of Quercus Garryana. The scarcity of col-
lections, which drew Burt’s comment, is almost certainly attributable to
the fact that on this substratum the fungus nearly always appears imma-
ture, or abortive. On dead sticks, mosses, and forest litter in the same
region the teeth are well developed, and the assignment to Schweinitz’s
H. Himantia is not difficult; such material agrees closely with Roth’s de-
scription and figure of C. byssacea.
From Persoon’s description also, C. byssacea should be the same as H.
Himantia, to which it is antecedent by perhaps half a year. _ Since we
have no warrant for taking the asterisk which precedes the name in Per-
soon as the sure sign of a subspecies (cf. Persoon, Syn. Plant. 1: x. 1805;
and 1: 174, where names so marked are referred to as species), “byssacea”
is probably the correct specific name, rather than “Himantia.” The genus
is not so readily determined. The fungus is certainly not a Coniophora
(as in Burt), nor a Hydnum sensu str., nor a Clavaria (as in Bourdot &
Galzin and in Persoon), nor, in any acceptable sense, an Odontia. Whether
Donk’s Mycoacia (or Miller’s Oxydontia) is a natural genus is a question
scarcely within the province of the present paper. It appears doubtful
that the fungus under discussion here is congeneric with either Hydnum
setosum Pers., the type of Oxydontia, or Hydnum fusco-atrum Fries, the
type of Mycoacia. However, until the genera of resupinate Hydnaceae can
be carefully correlated with appropriate sections or genera of the Thele-
phoraceae, it will not be possible to make more than a tentative place
for this somewhat anomalous species; and for the interim it is as well
left under Mycoacia Himantia.
Suksdorf 889, listed as a paratype of Burt’s species, is Aleurodiscus
_disciformis (DC. ex Fries) Pat.
CoNIOPHORA LAETICOLOR (Karst.) Karst. Peniophora dryina (Berk. & Curt.)
See under Coniophora dryina (Berk. & Curt.) Massee
ConropHora Potyporowwea (Berk. & Curt.) Burt = Corticium Polyporoideum
Berk. & Curt.
Corticium Polyporoideum Berk. & Curt., Grevillea L: 177. 1873 (? !!).
Coniophora Polyporoidea (Berk. & Curt.) Burt, Missouri Bot. Gard. Ann. 4:
247. 1917.
Hypochnus Polyporoideus (Berk. & Curt.) Overh., Mycologia 30: 275. 1938.
The spores of this species are asperulate and slightly colored, but are
not those of a Coniophora nor of a Tomentella. The species of Corticium
brought together by Bourdot & Galzin in the section Humicola will some
day be removed from the genus to their own place and this species will
go with them. Corticium Polyporoideum differs from the nearest species,
C. albo-ochraceum Bres., most obviously in the broader, blunt spores.
280 FaRLowiA, VOL. 1, 1943
Coniophora alboflavescens Ellis & Everh., which Burt reduced to syn-
onymy with the present species, is distinct, and a good Coniophora (see
under Contophora corticola Overh.).
CONIOPHORA SISTOTREMOIDES sensu Burt = C. suffocata (Peck) Massee
Corticium suffocatum Peck, New York State Mus. Rept. 30: 48. 1878 (!!)3 nec
C. suffocatum Bourd. & Galz., Soc. Mycol. Fr. Bul. 27: 263. 1911.
Coniophora suffocata (Peck) Massee, Linn. Soc. Bot. Jour. 25: 138. 1889; Burt,
Missouri Bot. Gard. Ann, 4: 254, 1917.
Coniophora Sistotremoides sensu Burt, Missouri Bot. Gard. Ann. 13: 312. 1926;
nec Thelephora Sistotremoides Schw. (= Coniophora olivacea ([{Fries] ex
Pers.) Karst.) 4
Hypochnus flavo-brunneus Dearn. & Bisby in Bisby et al., Fungi Manitoba p. 90.
1929 (!!).
Schweinitz’s material of Thelephora Sistotremoides shows the enormous
cystidia and the other characters of Coniophora olivacea, and is therefore
listed in synonymy with that species. C. suffocata Peck may then be re-
vived. Only a slide of Peck’s type was available for study, but in it the
fungus is well preserved, and there can be little doubt of its microscopic
characters. In Bisby et al., The Fungi of Manitoba and Saskatchewan
p. 75. 1938, H. flavo-brunneus was listed as a synonym of C. suffocata. It
agrees reasonably well with the type slide. Whether C. suffocata, C.
Betulae (Schum.) Karst., and some of the related species are distinguish-
able from each other, and from C. arida (Fries), is a question whose answer
is not attempted at this time. Perhaps the spores in C. suffocata are of
more even diameter than those of C. arida. For C. Sistotremoides (Schw.)
Massee see under Coniophora atrocinerea Karst.
CONIOPHORA UMBRINA (Fries) Karst. = C. olivacea ([Fries] ex Pers.) Karst.
See under Coniophora atrocinerea Karst.
ConropHora vaca Burt = Pellicularia flavescens (Bon.) Rogers
Coniophora vaga Burt, Missouri Bot. Gard. Ann. 4: 251. 1917.
Pellicularia flavescens (Bon.) Rogers, Farlowia 1: 105. 1943, ubi synon.
CORTICIUM
CorTICIUM ABEUNS Burt
Comment on this gloeocystidiate form is included here, not because the
species necessarily goes into synonymy, but because we have studied it
rather intensively and have found the description misleading and the
paratypes confused. The species was described by Burt (Missouri Bot.
Gard. Ann. 13: 250. 1926.) as having copious subglobose spores
6-7x4-6 yw. It is evident that this description cannot have been made
from the designated type, which is a collection made on coniferous wood
by Dr. R. P. Burke at Montgomery, Alabama (Burke 229). In that
collection the spores are 8.5-10.5x4.5-5.5 p, flattened and appearing
straight on one side with lateral apiculus; definitely rounded on the op-
posite side. Another feature not made clear in the description is the
Rocers & JacKSON: SYNONYMY OF THELEPHORACEAE 281
presence of substratal hyphae 2—2.5 » in diameter which become thick-
walled.
These features fit reasonably well the characters of Gloeocystidium
ochroleucum Bres. ex Bourd. & Galz. (= Gl. ochroleucum Bres. & Tor-
rend, nec Corticitum ochroleucum Fr., nec C. ochroleucum Bres.) which
was described as on coniferous wood from Portugal. We have seen only
collections from France on deciduous-leaved hosts assigned to this species
by Bourdot. While these are comparable, we hesitate to combine the
species without having seen the type from Portugal. In any case, Burt’s
specific name is the valid one in Corticium.
Of the seventeen paratypes assigned by Burt to this species, only three
resemble the type at all closely. These are Atkinson 2595, Macoun 812,
and Yasuda 12. We have examined all of the other collections and found
that none are comparable with the type. While several of them cannot
be identified, the following have been named: the collection from Maine
(Murrill 1938) and two of the British Columbia collections (Macoun 496
and 490) are all Corticium porosum Berk. & Curt.; the New Hampshire
(Snell 626) and the Wisconsin record (Humphrey 617) are Corticium
livido-caeruleum Karst.; the two Peck collections from New York are
both Cortictum Litschaueri Burt; and the New Mexico record (Long
21046) is Corticium punctulatum Cooke. None of these species have
anything in common with C. abeuns except the possession of gloeocystidia.
It seems probable that Burt’s description was taken largely from the New
Mexico collection rather than from the designated type.
CORTICIUM ALBO-STRAMINEUM (Bres.) Overh. = C. geogenium Bres.
See under Peniophora albo-straminea Bres.
CorRTICIUM APICULATUM Bres. = C. tessulatum Cooke
Corticium tessulatum Cooke, Grevillea 6: 132. 1878 (!!); Burt, Missouri Bot.
Gard. Ann. 13: 210. 1926.
Corticium illaqueatum Bourd. & Galz., Soc. Mycol. Fr. Bul. 27: 238. 1911 (!);
Hymen. de France p. 192, [1928]; Burt, Missouri Bot. Gard. Ann. 13: 236. 1926.
Corticium rhizophorum Bourd. & Galz., Soc. Mycol. Fr. Bul. 27: 238. 1911 (!).
Corticium apiculatum Bres., Mycologia 17: 68. 1925 (!!); Burt, Missouri Bot.
Gard. Ann. 13: 238. 1926; Overh., Mycologia 21: 280. 1929; nec C. areolatum
Bres., Mycologia 17: 68. 1925 (1!!).
Corticium illaqueatum f. rhizophorum (Bourd. & Galz.) Bourd. & Galz., Hymén.
de France p. 192. [1928].
While differing considerably in the degree of development of the subicu-
lum, the types of the three species listed above agree so closely in essential
microscopic characters that they are best considered all one species, for
which the name C. tessulatum Cooke has priority.
All of the seven collections cited by Burt as C. apiculatum have been
examined. None are like the type with the possible exception of the one
from British Columbia, which is close but doubtful. The assignment by
Burt of C. areolatum Bres. (q. v.) as a synonym of C. apiculatum Bres.
is found to be incorrect. The types of the two species differ widely,
282 FarLowiA, VoL. 1, 1943
especially in spore characters. Overholts (1.c.) has commented on this
species but the collection to which he refers (Overh. 3167) is Peniophora
albula Atk. & Burt. (q. v.).
CORTICIUM ARACHNOIDEUM Berk,
See under Corticium centrifugum (Léy.) Bres.
CORTICIUM AREOLATUM Bres.
This species, described by Bresadola (Mycologia 17: 68. 1925.) from
a collection made by J. R. Weir on Alnus tenuifolia at Priest River, Idaho
(Weir 23387), was listed by Burt (Missouri Bot. Gard. Ann. 13: 238.
1926.) as a synonym of C. apiculatum Bres. (q.v.). Study of the type
collection, however, shows that it is amply distinct and appears to be a
good species. None of the other collections cited by Burt under C. apicu-
latum are like it. Collections are now at hand, however, from British
Columbia on Salix, and from Idaho on Populus. Several collections made
in Ontario on Thuja occidentalis seem referable here.
Corticium areolatum Stahel, Phytopath. 30: 129. 1940, is Pellicularia
filamentosa (Pat.) Rogers, Farlowia 1: 113, 1943.
Corticitum ATKINSON Burt = Trechispora coronifera (Hohn. & Litsch.) comb. nov.
Gloeocystidium coroniferum Hohn, & Litsch., K. Akad. Wiss. Wien Math.-Nat.
KI. Sitzungsb. 116, I: 825. 1907.
Corticium Atkinsonii Burt, Missouri Bot. Gard. Ann. 13: 208. 1926.
Trechispora coronifera (Hohn. & Litsch.) comb. nov,
See under Sistotrema.
CorticlumM BEeRKELEYI Cooke = Peniophora aspera (Pers.) Sacc.
? Thelephora granulosa Pers. ex Fries, Syst. Mycol. 1: 446, 1821.
Thelephora aspera Pers., Mycol. Eur. 1: 153. pl. [5], fig. [4]. 1822.
Hydnum granulosum (Pers. ex Fr.) Pers., Mycol. Eur. 2: 184, 1825, quantum ad
typum Persoonianum.
Thelephora setigera Fries, Elench. Fung. 1: 208. 1828; Gen. Hymen. p. 17. 1836.
Hyphoderma spiculosum Wallr., Fl. Crypt. Germ. 2: 576. 1833.
Hyphoderma asperum (Pers.) Wallr., Fl. Crypt. Germ. 2: 577. 1833.
? Grandinia granulosa (Pers. ex Fries) Fries, Epicr. Syst. Mycol. p. 527. 1838;
Bourd. & Maire, Soc. Mycol. Fr. Bul. 36: 74. 1920; Bourd. & Galz., Hymén.
de France p. 408. [1928]; Bourd., Soc. Mycol. Fr. Bul. 48: 217. 1932.
Kneiffia setigera (Fries) Fries, Epicr. Syst. Mycol. p. 529. 1838.
Corticium setigerum (Fries) Karst., Bidr. Kanned. Finl. Nat. Folk 37: 143. 1882.
Cortictum myxosporum Karst., Bidr. Kanned. Finl. Nat. Folk 37: 240. 1882 (!!);
Hohn. & Litsch., K. Akad. Wiss. Wien Math.-Nat. Kl. Sitzungsb. 115, I:
1555. 1906.
Corticium latitans Karst., Rev. Mycol. 10: 74. 1888.
Kneiffia latitans (Karst.) Karst., Icon. Sel. 3: 7. fig. 73. 1889; Laurila, Soc. Zool.-
Bot. Fenn. Vanamo Ann. Bot. 10(4): 10. 1939.
Corticium Berkeleyi Cooke ex Massee, Linn. Soc. Bot. Jour. 27: 133. 1890 (!!);
Burt, Missouri Bot. Gard. Ann. 13: 183. 1926.
Corticium Chusqueae Pat., Soc. Mycol. Fr. Bul. 9: 134. 1893 (!!).
Peniophora trachytricha Ellis & Ev., Acad. Nat. Sc. Phila. Proc. 47: 413. 1895 (!!).
Kneiffiella aspera (Pers.) Underw., Torr. Bot. Cl. Bul. 24: 206. 1897.
Kneiffiella setigera (Fries) P. Henn. in E. & P., Nat. Pflanzenfam. 1'**: 140.
[1898].
Rocers & JACKSON: SYNONYMY OF THELEPHORACEAE 283
Kneiffiella latitans (Karst.) P. Henn. in E. & P., Nat. Pflanzenfam. 1***: 141.
[1898].
Pycnodon asper (Pers.) Underw., Torr. Bot. Cl. Bul. 25: 631. 1898 (falso ut
‘asperum’).
Odontia Acerina Peck, New York St. Mus. Rept. 53: 847. 1900.
Neokneiffia aspera (Pers.) Earle apud Mohr, U. S. Nat. Herb. Contr. 6: 203. 1901.
Peniophora setigera (Fries) Hohn. & Litsch., Ann. Mycol. 4: 289. 1906; Bourd. &
Galz., Hymén. de France p. 309. [1928].
Peniophora latitans (Karst.) Hohn. & Litsch., K. Akad. Wiss. Wien Math.-Nat.
K]. Sitzungsb. 115, I: 1554. 1906.
Peniophora aspera (Pers.) Sacc., Fl. Ital. Crypt. Hymen. p. 1182. 1916.
? Asterostromella granulosa (Pers. ex Fries) Bourd. & Galz., Hymén. de France
p. 396. [1928], saltem quantum ad typum, descr. icon. excl.
Odontia vesiculosa Burt ex Povah, Mich. Acad. Papers 9: 262, 1929 (!!).
Odontia setigera (Fries) Miller, Mycologia 26: 19. 1934.
Kneiffia setigera var. trachytricha (Ellis & Ev.) Rick, Brotéria Sér. Trim. Ciénc.
Nat. 3: 70. 1934.
? Vararia granulosa (Pers. ex Fries) Laurila, Soc. Zool.-Bot. Fenn. Vanamo Ann.
Bot. 10(4) : 15. 1939,
(Thelephora granulosa Pers., Syn. Meth. Fung. p. 576. 1801.
The type collection of Corticitum Berkeleyi was distributed in Ravenel’s
Fungi Americani 225; material in three different sets of this work, includ-
ing a fragment in Burt’s own herbarium, is the same fungus as that de-
scribed by Bourdot & Galzin and by Miller as, respectively, Peniophora
setigera and Odontia setigera. In his account of C. Berkeleyi Burt makes
no mention of the minutely papillose hymenium nor of the very large,
septate cystidia which are, nevertheless, clearly present. It therefore
appears that Burt drew up his description not from the type but from
quite different specimens present in his herbarium, specimens of a fungus
closely allied to Corticium bombycinum (Somm.) Karst. Whatever de-
cision is finally reached concerning C. Berkeleyi sensu Burt, there is no
doubt concerning the status of C. Berkeleyi Cooke.
Determination of the correct name for the latter fungus is not a simple
matter. Bourdot has reported (l.c., 1932) that Persoon’s type of T.
granulosa is the same as P. setigera. The simplest manner of dealing
with the problem would then be to adopt a binomial formed from T.
granulosa Pers. ex Fries, the earliest valid name in the long synonymy of
the species. But so many fungi have been identified (incorrectly) with
T. granulosa and G. granulosa that the use of the name (even though cor-
rectly) for another species would be bound to result in considerable con-
fusion. This is good but insufficient reason for rejecting the epithet
“sranulosa.” What is more important is that it is not allowable to define
T. granulosa merely from Persoon’s concept and his specimen; that species
was “‘validated” (as some would have it) by publication in Fries’s “Sys-
tema,” and must forever be seen through Fries’s eyes. Now in the “Epi-
crisis” and again in the “Hymenomycetes Europaei,” Fries wrote (under
G. granulosa) that his concept “includes several species . . . ; I have now
before my eyes the noblest form, the var. ochracea of Albertini & Schwei-
284, Fartowia, VoL. 1, 1943
nitz.” Whatever may have been Persoon’s species, then, Fries’s was an
omnium gatherum, incapable of definition and typified by nothing or, if
“noblest form’ can be translated “type,” by an alien fungus. It is there-
fore hereby proposed that Thelephora granulosa Pers. ex Fries be con-
sidered a nomen dubium and that it and all binomials based on it be
dropped from mycological nomenclature.
Thelephora aspera Pers. was repeatedly acknowledged by Fries, as well
as by later authors, as identical with 7. setigera Fries; the account of the
latter in the “Elenchus,” and of Kneiffia setigera in the “Hymenomycetes
Europaei,” may in this connection well be compared with Persoon’s de-
scription of 7. aspera. On Fries’s authority, then, P. aspera is the correct
name for the present species so long as it is retained in Peniophora.
CorTICIUM BOTRYOIDEUM Overh. = Pellicularia pruinata (Bres.) Rogers
Corticium botryoideum Overh., Mycologia 26: 510. 1934.
Pellicularia pruinata (Bres.) Rogers ex Linder; Rogers, Farlowia 1: 107. 1943,
ubi synon,
CorTICIUM CALCEUM sensu Romell & Burt Nomen confusum.
(Thelephora calcea Pers. * glebulosa Fries, Elenchus Fung. 1: 215, 1828 (!!).
(Corticium glebulosum (Fries) Bres., Fungi Trident, 2: 61. 1898.
(Peniophora glebulosa (Fries) Sacc. & Syd., Syll. Fung. 16: 195. 1902; Burt,
Missouri Bot. Gard. Ann. 12: 282. [1926]; Bourd. & Galz., Hymén. de France
p. 288, [1928].
(Kneiffia glebulosa (Fries) Bres., Ann. Mycol. 1: 104. 1903.
(Corticium calceum sensu Romell & Burt, Missouri Bot. Gard. Ann. 13: 203. 1926;
nec C. calceum (Pers.) Fries, Epicr. Syst. Mycol. p. 562. 1838. (= Thelephora
calcea Pers., Mycol. Eur. 1: 153. 1822).
(Peniophora cretacea Bourd. & Galz., Hymén. de France p. 288. [1928].
All of the names here listed are based either directly on the type speci-
men of T. calcea * glebulosa or else on that name as published by Fries
and hence indirectly on its type. It is now evident, as Bourdot’s de-
scription and figure of P. cretacea strongly suggest, as Litschauer has
clearly stated (Svensk. Bot. Tidskr. 32: 285. 1938; cf. also Lund. &
Nannf., Fungi Exs. Suecici 465. 1937, and Litsch., Ann. Mycol. 39: 134.
1941), and as examination of the material itself fully confirms, that Fries’s
type consists of the cystidiolate Peniophora Romellii growing over the
cystidiate fungus which has since 1902 been treated under the name Penio-
Phora glebulosa. Burt has denied that any cystidia are present in this
specimen; but his own slides show them. Thelephora calcea * glebulosa
and all its synonyms here listed are nomina confusa, not applicable to
any one organism. (cf. Peniophora glebulosa sensu Bres.).
It can be deduced from the above, since Burt did not observe the cystidia
in the designated type, that his intended concept of “C. calceum Fries
emend. Romell and Burt” was to apply to the cystidiolate component of
the complex, which is a clearly identifiable species now properly called
Peniophora Romellii Litsch. (ex Bourd., Soc. Mycol. Fr. Bul. 48: 212.
1932 = Cortictum cretaceum Romell ex Lund. & Nannf., Fungi Exs. Suec.
Rocers & JAcKSON: SYNONYMY OF THELEPHORACEAE 285
465. 1937; nec C. cretaceum [Fries] Cooke ex Sacc., Syll. Fung. 11: 128.
1895, quod Thelephora cretacea Fries, Obs. Mycol. 1: 153. 1815.). This is
borne out by an examination of the Swedish collections cited by Burt and
made by Romell or Romell and Burt. The two joint collections and
Romell’s 321, 322, and 325, together with one uncited Romell collection
323, are all P. Romellii, which is evidently common in Sweden. On the
other hand, three of Romell’s collections: 185, 211, and 324, together with
8 (? plus 3 poor and somewhat uncertain) of the 22 American collections
cited are Corticium furfuraceum Bres. (Mycologia 17: 69. 1925 (!!);
Burt, Missouri Bot. Gard. Ann. 13: 242. 1926; = C. subpallidulum
Litsch., Svensk. Bot. Tidskr. 32: 284. 1938 (!!)5 cf. also Lund. & Nannf.,
Fungi Exs. Suecici No. 571. 1938.).
P. Romellii is apparently rare in North America and does not appear
among the American collections cited by Burt; this, together with the
fact that half the American collections are as stated above, tends to con-
firm Litschauer’s statement (l.c.) that Burt’s description, and hence his
actual concept, more nearly fits C. subpallidulum (i.e. C. furfuraceum
Bres.) than C. cretaceum Romell in sched. (i.e. P. Romellii Litsch.). It
is worthy of note that neither Burt nor Bresadola mentioned in their de-
scriptions the highly characteristic capitate cystidioles so well described
and figured (as Képfchenparaphysen) by Litschauer for C. subpallidu-
lum (l.c.).
The remaining American collections cited by Burt show little con-
sistency. Four of them (the three Canadian collections made by Macoun
and one of thé Vermont collections) prove to be a possibly undescribed
Peniophora related to P. livida Burt and extremely common in Ontario.
Three of the six New York collections (Peck 9, 20, and Atkinson 941) are
Corticium suecicum Litsch. (apud Lund. & Nannf., Fungi Exs. Suecici
464, 1937; = C. calceum sensu Bourd. & Galz., Hymén. de France p. 237.
[1928]). The Wisconsin record (Harper 853) is C. incrustans Hohn.
& Litsch.; the Pennsylvania collection (Overh. 4809) has coronate Urni-
gera basidia, and two others (Michigan, Humphrey 1662, and Idaho,
Weir 6350) cannot be identified.
* Both C. furfuraceum and P. Romellii are cystidiolate but since it is a
matter of indifference, at present, whether such forms be included in
Corticium or Peniophora, neither is here transferred.
The true Corticium calceum (Pers.) Fries is, according to Bresadola,
who indicates by an exclamation point that he has seen Persoon’s material,
Sebacina calcea (Pers.) Bres. (cf. McGuire, Lloydia 4: 23. 1941.).
CortTiclumM cANUM Burt=C. Galzini Bourd.
Corticium Galzini Bourd., Rev. Sc. Bourb. 23: 11. 1910; Bourd. & Galz., Hymén.
de France p. 194. [1928] (!).
Corticium canum Burt, Missouri Bot. Gard. Ann. 13: 206. 1926 (!!).
C. Galzini is included in the group Pellicularia of Bourdot & Galzin.
Bourdot’s description gives the spore measurement as 3—4.5 x 1.25—2.5 yp.
286 FarLtowiA, VoL. 1, 1943
An examination of several collections assigned to this species from the
Bourdot herbarium shows considerable variation in spore size and shape.
The type collection of C. canum Burt has spores 3.5—4.5 x 2-3 yp, slightly
broader than any European collection we have seen. The variation is so
slight, however, that it seems unnecessary and undesirable to attempt to
recognize another species in this characteristic but very difficult group
of species.
Eight paratypes are listed by Burt for C. canum. Of these only one,
Humphrey 6375, is like the type. The others are as follows: Atkinson
2563 is Corticium coronilla Hohn. (q.v.); Langlois 168 is Cortictum
sphaerosporum (R. Maire) Hohn. & Litsch.; Weir 21 is Cortictum lacteo-
lum Bourd.; Humphrey 6413 is Corticium microsporum (Karst.) Bourd.
& Galz.; Weir 11101 is Corticium furfuraceum Bres.; Shear 1063 and Weir
447 have not been determined but are none of these.
CorTICIUM CENTRIFUGUM (Léy.) Bres. Untenable.
The taxonomy of those species of Corticium sect. Pellicularia Bourd. &
Galz. which cluster about C. arachnoideum sensu Bres. and C. centrifugum
sensu Bres. can be finally reduced to a satisfactory system only by com-
parative study of large numbers of specimens; and such relative finality
has as yet been attained by no one. Since several of the names involved
in this complex are in common use it has seemed desirable to include in
these notes as clear a statement of the general situation as is possible,
without attempting any solution at this time. As a basis for the dis-
cussion which follows a list of the names which must be considered is
given below together with the essential references, arranged chrono-
logically.
? Corticium arachnoideum Berk., Ann. Mag. Nat. Hist. 13: 345, 1844 (!) ; Massee,
Linn. Soc. Bot. Journ. 27: 135. 1890; nec C. arachnoideum sensu Bres.; Burt,
Missouri Bot. Gard. Ann. 13: 184. 1926; et auctt.
? Hypochnus centrifugus Tul., Sel. Fung. Carpol. 1: 114. 1861. Vix Rhizoctonia
centrifuga Lév., Ann. Sc. Nat. Bot. II. 20: 225. 1843.
Hypochnus bisporus Schroet. in Cohn, Krypt.-Fl. Schles. 3(1) : 415. 1888 (2 Jun.).
Corticium Pezizoideum Ellis & Everh., Jour. Mycol. 4: 74. 1888 (Aug.) (!!); nec
C. Pezizoideum (Schw.) Schrenk, Torr. Bot. Cl. Bul. 21: 388, 1894. :
Corticium centrifugum sensu Bres., Ann. Mycol. 1: 96. 1903; Héhn., Osterr. Bot.
Zeitschr. 54: 427. 1904; Ann. Mycol. 3: 188. 1905; Burt, Missouri Bot. Gard.
Ann. 13: 206. 1926; Bourd. & Galz., Hymén. de France p. 197. [1928]; nec
C. centrifugum (Weinm.) Fries, Hymen. Eur. p. 658. 1874 (= Thelephora
centrifuga Weinm., Hymen. Gasterom. Ross. p. 392. 1836; Coniophora centrifuga
(Weinm.) Karst., Bidr. Kanned. Finl. Nat. Folk 37: 160. 1882).
Corticium bisporum (Schroet.) Hohn. & Litsch., Ann. Mycol. 4: 289, 1906; Bourd.
& Galz., Soc. Mycol. Fr. Bul. 27: 240, 1911; Wakef. & Pears., Br. Mycol. Soc.
Trans. 8: 216. 1923.
Corticium decipiens Hohn. & Litsch., K. Akad. Wiss. Wien Math.-Nat. KI]. Sitzungsb.
117, I: 1116. 1908 (!!).
Corticium consimile Bres., Mycologia 17: 68. 1925 (!!).
Corticium centrifugum var. soredioides Bourd, & Galz., Soc. Mycol. Fr. Bul. 27:
240. 1911; Hymén. de France p. 198. [1928].
Rocers & JACKSON: SYNONYMY OF THELEPHORACEAE 287
Corticium centrifugum var. macrospora Bourd. & Galz., Soc. Mycol. Fr. Bul. 27:
240. 1911; Hymén. de France p. 198. [1928].
Corticium centrifugum subsp. fugax Bourd. & Galz., Hymén. de France p. 198.
[1928]. (Tomentella fugax Karst. in sched., ex Hohn. & Litsch., K. Akad.
Wiss. Wien Math.-Nat. Kl]. Sitzungsb. 116, I: 1572. 1906, pro synon.).
Corticium centrifugum subsp. bisporum (Schroet.) Bourd. & Galz.. Hymén. de
France p. 199. [1928].
Corticium centrifugum var. tenuis Donk, Nederl. Kruidk. Arch. 1930: 84. 1930.
Non Fusisporium Kuhnii Fckl., Fung. Rhen. Suppl. Fasc. 5, 1920. 1867 (!!);
Symb. Mycol. p. 371. 1869.
An analysis of this series indicates that this much is clear: 1) The name
C. centrifugum is untenable (since C. centrifugum Fries must be a Conio-
phora or a Serpula [ = Gyrophana] ). 2) There is probably no relation
between R. centrifuga Lév. and any of the fungi to which the name C.
centrifugum. has been applied by mycologists. 3) H. centrifugus Tul.
(Léveillé’s publication being invalid for any perfect basidiomycete) was
probably quite a different thing from the Corticium with which Bresadola
identified it (“flocci septis illis frequentissime utuntur quae ampulla
laterali aucta videntur . . . nempe geniculat[i] seu imperfecte articulat[i]”
—Tulasne, l.c.). 4) The one specimen at hand determined by Bresadola
as C. centrifugum agrees extremely well with Schroeter’s description of
H. bisporus, in having bisporous basidia, as in other points. 5) Of two
authentic specimens of C. arachnoideum (either, or perhaps each, being
possibly a part of the type), neither is the fungus identified with C. arach-
noideum by Bresadola and later authors. One (in the general collections
of the Farlow Herbarium) bears perfect basidia and is C. octosporum
Schroet. (i. e., Trechispora Brinkmanni (Bres.) ). The other (in the Curtis
collection), with a few mature basidia but no recognizable spores, has
no clamps whatever, and in all visible respects but this lack of clamps
and the presence of four sterigmata to the basidium is similar to Bresa-
dola’s specimen of C. centrifugum (cf. Rea, Br. Basid. p. 676. 1922, where
C. centrifugum is treated as a synonym of C. arachnoideum). 6) C. Pezi-
zoideum and C. decipiens differ only in size, the former having all elements
approximately twice as large as in the latter. 7) Fusisporium Kuhnivi is
as described by Fuckel, and not, as Hohnel has stated, C. centrifugum.
8) A considerable series of specimens examined includes most of the
variations which characterize Bourdot’s subspecies and varieties; but these
characters are recombined in so many ways that scarcely any two speci-
mens agree, and only rarely can a collection be surely identified with one
of the described subdivisions. 9) Corticium consimile, based on a single
collection made on Larix occidentalis by J. Weir (10808) in Idaho, was
omitted from Burt’s treatment of the genus. _ It is close to Corticium cen-
trifugum sensu Bres. as that species is usually understood, differing chiefly
in the complete absence of clamps. Bresadola’s type is identical with a
specimen of Corticium decipiens determined by Litschauer, but differs
from the type of that species in thicker and less clavate basidia, shorter-
celled and wider-branching mycelium, and more ellipsoid and less flat-
tened spores.
288 Fartowia, Vou. 1, 1943
Attempts at segregation from the complex should be made only after
considerable further study, and on a number of correlated characters.
For example, unless an indubitable Berkeley type can be shown to have
clamps throughout (and basidia not Urnigera-like), C. arachnoideum can-
not be separated on the possession of abundant clamps. Presence or ab-
sence of rare clamps on the basal mycelium is a difficult and probably not
valid basis of segregation of a species from C. decipiens (and C. Pezi-
zoideum) ; in some specimens the first clamp has been demonstrated only
after a search of twenty minutes. Another anomaly is that 2-spored ma-
terial (i.e., C. bisporum sensu stricto) shows clamps, where 4-spored
(e.g., C. decipiens) may not. It is quite possible that there may be no
natural planes of cleavage within the group. The available valid names
include C. arachnoideum, C. bisporum, C. Pezizoideum, C. decipiens, and
C. consimile. As a temporary expedient the oldest available combination,
C. bisporum, might be used as a name under which to file collections be-
longing in this complex. If one objects to including 2-spored forms with
those having 4-spored basidia in the same species, C. bisporum could be
used for the former and C, Pezizoideum for the latter. Conceivably C.
arachnoideum (non sensu Bres.) might replace C. Pezizoideum, or both
names, when Berkeley’s type specimen is settled upon and examined.
Corticium conFINE Bourd. & Galz. sensu Burt
Only one American collection is recorded by Burt (Missouri Bot. Gard.
Ann. 13: 246. 1926.) under this name. An examination of this collec-
tion (from Middlebury, Vermont) in the Burt herbarium shows that it is
Trechispora coronifera (Hohn. & Litsch.). The description was obviously
taken from the Vermont collection and not from the Bourdot collections
cited. So far as the Burt record is concerned this species may be stricken
from the North American flora.
CorTICIUM CONSIMILE Bres.
This species was omitted from Burt’s treatment of the genus. See un-
der Corticium centrifugum (Lév.) Bres.
CortIcItuM corRoNILLA Hohn. = Trechispora Brinkmanni (Bres.) comb. nov.
Corticium coronilla Hohn., Biggs, Mycologia 29: 686. 1937; Dodge, Mycologia
30: 133. 1938.
Odontia Brinkmanni Bres., Ann. Mycol. 1: 88. 1903.
Trechispora Brinkmanni (Bres.) comb nov.
See under Sistotrema.
CortIcIluM cRUSTACEUM (Karst.) Karst. =C. contiguum Karst.
Corticium contiguum Karst., Soc. Faun. Fl. Fenn. Acta 2(1): 39. 1881 (!!);
Bidr. Kanned. Finl. Nat. Folk 37: 150. 1882.
Xerocarpus crustaceus Karst., Hedwigia 35: 45. 1896 (!!).
Stereum crustaceum (Karst.) Sacc. & Syd., Syll. Fung. 14: 215. 1899.
Corticium crustaceum (Karst.) Karst., Bidr. Kanned. Finl. Nat. Folk 62: 93. 1903;
Burt, Missouri Bot. Gard. Ann. 13: 196. 1926; Bourd. & Galz., Hymén. de
France p. 211. [1928]; Laurila, Soc. Zool.-Bot. Fenn, Vanamo Ann. Bot. 10(4):
5. 1939,
Rocers & JAcKSON: SYNONYMY OF THELEPHORACEAE 289
Gloeocystidium contiguum (Karst.) Bourd. & Galz., Soc. Mycol. Fr. Bul. 28: 362.
[1913]; Hymén. de France p. 255. [1928], saltem quantum ad typum.
Corticium subcinereum Burt, Missourt Bot. Gard. Ann. 13: 277. 1926 (!!).
The original description of C. contiguum reads as follows: “Hymenium
contiguum, calceum; sporae longit. 4-6 mmm., crassit. 3-4 mmm. Ad
lignum Juniperi.” In the second reference a redescription in Swedish
is given which, translated, reads as follows: “Fruiting layer coherent,
chalk color, spores abundant. On Juniper wood. Finland (Mustiala)
_. . Effused, adnate, waxy, the periphery similar: spores oval, 4-6 mmm.
long, 3-4 mmm. broad.” It should be noted that both descriptions give
the substratum as Juniper wood. It is evident that these descriptions can-
not refer to the species described as Gl. contiguum by Bourdot & Galzin.
In the herbarium of the New York Botanical Garden (Ellis collection)
there is a specimen labeled, in Karsten’s hand, “Cortictum contiguum
Karst. Mustiala, ad Juniperinum. Aug. 1866. P. A. Karsten.” An ex-
amination of this specimen, which may be considered part of the type,
reveals that the substratum is not Juniperus as Karsten himself labeled it,
but is the wood of some deciduous-leaved tree, and the fungus is identical
with the better known Corticium crustaceum Karst. which occurs chiefly,
if not exclusively, on broad-leaved trees. Laurila (l.c.) lists C. contt-
guum under C. crustaceum, with the qualification: “sensu Karstenii pro
max. parte.” Two of four collections of C. contiguum from Karsten’s
herbarium which we have examined are the same as C. crustaceum. A
third is C. porosum Berk. & Curt. (see under C. stramineum Bres.), which
may have been the same as the “Specim. orig.!” to which Bourdot & Galzin
refer in the second line of their account of Gl. contiguum. The fourth is
Gloeocystidium Karstenii Bourd. & Galz. (Hymén. de France p. 254,
[1928] = Corticium ochraceum sensu Karst.; nec C. ochraceum Fries).
With this evidence before us we can see no alternative to replacing C.
crustaceum Karst. with C. contiguum Karst.
The published type of C. swbcinereum Burt (Macoun 37) is certainly
typical C. crustaceum. The description, however, does not well describe
the type specimen. In the Burt herbarium two specimens are marked
“type” in Burt’s hand. One of these is the Kansas collection (Bartholo-
mew 8702) cited by Burt. It is evident that the description was drawn
from the Kansas collections, which are all alike, and which would appear
to represent a form of Peniophora cinerea (Pers. ex Fries) Cooke in
which the cystidia are not developed. The Massachusetts collection cited
by Burt is a grandinioid Corticium with rough spores.
Corticium cRUSTULINUM Burt —=C. amianthinum Bourd. & Galz.
Corticium amianthinum Bourd. & Galz., Soc. Mycol. Fr. Bul. 27: 260. 1911 (!).
Corticium crustulinum Burt, Missouri Bot. Gard. Ann. 13: 209. 1926 (!!); nec
C. crustulinum Bres., Ann. Mycol. 18: 47, 1920.
Corticium albo-ochraceum subsp. amianthinum (Bourd. & Galz.) Bourd. & Galz.,
Hymén. de France p. 231. [1928] (ut ‘amiantinum’.).
290 Fartowia, Vou. 1, 1943
One of the small number of species characterized by ampulliform swell-
ings on the mycelium. The brittle yellow crust formed by the hymenium
and subhymenial hyphae is somewhat thicker than in the authentic speci-
men of C. amianthinum, and the subicular mycelium is on the average
somewhat more slender; but the differences seem too slight to justify
nomenclatorial recognition. Bourdot’s species is certainly related to C.
albo-ochraceum Bres. but differs, according to the material at hand, in
having abundant rather than scanty subiculum, in crustose rather than
hypochnoid hymenium, in size of spores and basidia, and in spores even-
walled (at least, typically) and colorless rather than asperulate and
brownish.
CortTIcIUM EFFUSCATUM Cooke & Ellis = Vararia effuscata (Cooke & Ellis) comh.
nov.
Corticium effuscatum Cooke & Ellis, Grevillea 9: 103. 1881 (!); Burt, Missouri
Bot. Gard. Ann. 13: 248. 1926.
Asterostromella effuscata (Cooke & Ellis) Bourd. & Galz., Hymén. de France
p. 396. [1928].
Vararia effuscata (Cooke & Ellis) comb. nov.
Since poorly differentiated dichophyses, such as occur in this species,
are not always clearly distinguishable from dendrophyses, the presence
of these bodies alone might not constitute sufficient grounds for including
C. effuscatum in Vararia. However, the basidia here are of a sort ap-
parently found in no other group —ventricose, elongate, emergent far
beyond the surface of the hymenium, and bearing slender erect sterig-
mata; in addition to the gloeocystidia there are present thin-walled, acute
cystidia with homogeneous content; in these points, and in the spores, it
agrees with other indubitable species of Vararia. Such coordination of
characters is conclusive where a single one might not be.
CorticluM EFFUSUM Overh, = Stereum Murraii (Berk. & Curt.) Burt
Thelephora Murraii Berk. & Curt., Linn. Soc. Bot. Jour. 10: 329. 1868 (!!).
Corticium Murrayi (Berk. & Curt.) Pat. apud Duss, FI. Crypt. Antilles Fr. p. 228.
1904.
Stereum Murraii (Berk. & Curt.) Burt, Missouri Bot. Gard. Ann. 7: 131. 1920,
ubi synon. (falso ut “Murrayi”).
Corticium effusum Overh., Mycologia 22: 238. 1930 (!!).
Stereum Murraii fa. tuberculosum (Fries) Pildt, Czech. Acad. Agr. Ann. 5: 399.
1930.
The difference in appearance of the type collection of Overholts’s species
from that of typical specimens of Stereum Murraii is best attributed to
occurrence on much decayed decorticated wood, and to age. The col-
lection is evidently young, just beginning to sporulate. The gloeocys-
tidia in the hymenium are elongated vesicular bodies and their occurrence
may be duplicated in the type of Stereum Murraii and in other collections.
The essential microscopic details are identical in the two forms.
Rocers & JACKSON: SYNONYMY OF THELEPHORACEAE 291
CorTICIUM ERMINEUM Burt = Aleurodiscus amylaceus (Bourd. & Galz.) comb. nov.
Corticium amylaceum Bourd. & Galz., Soc. Mycol. Fr. Bul. 27: 259. 1911 (!);
Hymén. de France p. 228. [1928].
Corticium ermineum Burt, Missouri Bot. Gard. Ann. 13: 182. 1926 (!!); Overh.,
Mycologia 21: 280. 1929,
Corticium sociatum Burt, Missouri Bot. Gard. Ann. 13: 192. 1926 (!!).
Aleurodiseus amylaceus (Bourd. & Galz.) comb. nov.
A comparison of an authentic collection of C. amylaceum Bourd. & Galz.
(probably a portion of the type) made by Galzin (6803) on Juniperus
(ex Herb. H. Bourdot, Champ. de l’Aveyron 7490), with the types of the
two species described by Burt leaves no doubt that all should be assigned
to the same species. The differences noted by Burt in his discussion under
C. ermineum disappear when a series of specimens are compared. In
addition to the types of C. ermineum and C. sociatum eight collections on
Thuja plicata from the western United States (Oregon, Montana, Idaho)
and British Columbia, together with three collections on Thuja occidentalis
from widely separated localities in Ontario, have been available for study
in the University of Toronto herbarium. This series of specimens shows
considerable variation in gross appearance attributable primarily to differ-
ences in thickness and age. Burt stresses the absence of clamp connec-
tions in both species described by him. Clamps are, however, present in
each of the type collections.
As in the case of Corticium subgiganteum Berk. (q. v.), it is the type of
basidium that is of paramount importance in deciding the generic posi-
tion of C. amylaceum, not the absence of some especially distinctive type
of paraphysis. The species is readily recognized by the thin white or
cream fructifications, pelliculose or thin membranous hymenium, basidia
of the type characteristic of Alewrodiscus accompanied by hyphal para-
physes, and the copious amyloid spores which are broadly ellipsoid,
8-12 x 5-8 p.
Burt assigned a collection made in Vermont on coniferous wood to
C. ermineum. This collection proves to be Corticium geogenium Bres.
(see under Peniophora albostraminea Bres.). A collection from Manitoba
assigned to C. sociatum by Burt, while certainly not this species, has not
yet been identified.
CorticIUM FENESTRATUM Overh. = Pellicularia flavescens (Bon.) Rogers
Corticium fenestratum Overh., Mycologia 26: 510. 1934.
Pellicularia flavescens (Bon.) Rogers, Farlowia 1: 105. 1943, ubi synon.
Corticium fuscostratum Burt and C. ocHROLEUCUM Bres.
Corticium ochroleucum Bres., Fungi Trident. 2: 58. 1898 (!); Bourd. & Galz..
Hymén. de France p. 193. [1928]; nec C. ochroleucum (Fries) Fries, Epicr.
Syst. Mycol. p. 557. 1838; nec C. ochroleucum (Noack) Burt ex Peltier, Univ.
Ill. Agric. Exp. Sta. Bul. 189: 290. 1915.
Corticium fuscostratum Burt, Missouri Bot. Gard. Ann. 13: 299, 1926 (!!).
Because of the earlier use of the name by Fries, Burt’s binomial takes
292 FarLtowiaA, VoL. 1, 1943
precedence over C. ochroleucum Bres., the name currently used in Europe
for this common species.
CorRTICIUM ILLAQUAETUM Bourd. & Galz. = C. tessulatum Cooke
See under Corticium apiculatum Bres.
Burt (Missouri Bot. Gard. Ann. 13: 236. 1926.) recorded but one
American collection (Langlois 203). While agreeing with thicker forms
of this species in general appearance, this shows spores cylindrical rather
than subpyriform and the identification is doubtful.
Cortictum INcANUM Burt — C. Tulasnelloideum Hohn. & Litsch.
Corticium Tulasnelloideum Hohn. & Litsch., K. Akad. Wiss. Wien Math.-Nat.
KI. Sitzungsb. 117, I: 1118. 1908 (!!); Bourd. & Galz., Hymén. de France
p. 235. [1928]; Rogers, Univ. Iowa Studies Nat. Hist. 15(3): 20. 1933.
Corticium incanum Burt, Missouri Bot. Gard. Ann. 13: 205. 1926 (!!).
Hypochnus Tulasnelloideum (Hohn. & Litsch.) Rea, Br. Mycol. Soc. Trans. 12:
222, 1927.
Of the six paratypes of C. incanum cited by Burt, none are like the type.
Two of the Canadian records, Macoun 34 and 36, are Corticium coronilla
Héhn., as is the North Carolina collection (Couch 4225). The New
York record collected by House is close to C. sphaerosporum (R. Maire)
Hohn. & Litsch. The third Canadian collection, Macoun 32, is a possibly
undescribed Corticium belonging in the section Athele of Bourdot &
Galzin. The collection from Vermont has not been identified.
The species is, however, a common one having a wide range in North
America. Specimens are now at hand from Quebec, Ontario, British
Columbia, New Hampshire, Massachusetts, Connecticut, Rhode Island,
New York, New Jersey, Georgia, Ohio, Iowa, Missouri, and Oregon.
CorTICIUM INVESTIENS (Schw.) Bres. = Vararia investiens (Schw.) Karst.
Radulum investiens Schw., Am. Phil. Soc. Trans. n. s. 4: 165. [1832] (!!).
Corticium alutarium Berk. & Curt., Grevillea 2: 4. 1873 (!!).
Xerocarpus alutarius (Berk. & Curt.) Karst., Bidr. Kanned. Finl. Nat. Folk 48:
418. 1889.
Thelephora subochracea Peck, New York State Mus. Rept. 46: 109. 1893 (!!).
Corticium investiens (Schw.) Bres., I. R. Accad. Agiati Atti III 3: 110. 1897;
Burt, Missouri Bot. Gard. Ann. 13: 283. 1926.
Vararia investiens (Schw.) Karst., Bidr. Kanned. Finl. Nat. Folk 62: 96. 1903;
Martin, Iowa Acad. Sci. Proc. 44: 48. [1938].
Asterostromella investiens (Schw.) Héhn. & Litsch., K. Akad. Wiss. Wien Math.-
Nat. Kl. Sitzungsb. 117, I: 1083. 1908; Bourd. & Galz., Hymén. de France
p. 395. [1928].
(Vararia alutaria (Berk. & Curt.) Karst., Soc. Faun. Fl. Fenn. Med. 16: 2. 1888,
nomen nudum (genus nondum descriptum).
(Langloisula spinosa Ellis & Everh., Jour. Mycol. 5: 68. pl. 10, fig. 1, 2. 1889 (!!)
(? nomen confusum) ; Héhn., K. Acad. Wiss. Wien Math.-Nat. Kl. Sitzungsb.
128, I: 537. 1919.
The genus Langloisula was published for what Ellis believed to be a
member of the Mucedineae; nevertheless, as von Hohnel was able to judge
from the published figure, the specimen collected by Langlois shows the
Rocers & JacKson: SYNONYMY OF THELEPHORACEAE 293
dichophyses characteristic of an Asterostromella. Furthermore, the type
both superficially and in its dichophyses, hyphae, and occasional thin-
walled cystidia does not differ greatly from Radulum investiens Schw.
What appear to be immature basidia are present but basidiospores are
apparently wanting; even so, there is reasonable evidence that Schweinitz’s
species is the one present in the Langlois collection. The thick-walled
colored conidia described by Ellis occur abundantly, borne singly at the
ends of usually long pedicels; but even after long-continued examination
it has not been possible to demonstrate a connection between them and
the mycelium of the basidiomycete, nor on any of their pedicels the clamps
that characterize the hyphae of R. investiens. These conidia resemble the
spores of Chromosporium viride Corda sufficiently to suggest that the para-
sitic component may be a species of Chromosporium. Furthermore, the fruc-
tification is (as noted by Ellis) brittle, as though disintegrating, in con-
trast to the moderately tenacious hymenium ordinarily found in this
species, and there are other indubitable hyphomycetes present. It is
therefore probable, although far from proved, that Langlois’s specimen
consists of parasitized material, and that Ellis’s generic and specific diag-
noses took about equal cognizance of the fungus host and its most con-
spicuous parasite. Under the circumstances it seems best for the present
to consider both Langloisula and L. spinosa to be nomina confusa, and
to retain the later name Vararia.
Vararia was originally published by Karsten as a subgenus of Xero-
carpus, for the single species X. alutarius (1.c.). When he elevated the
group to the rank of genus it still contained only the single species, this
time treated under the older specific name “investiens.” Since Astero-
stromella has the same type (1.c.) it can be only a synonym of Karsten’s
genus. Most authors have used the later name: Donk (Nederl. Mycol.
Ver. Med. 18-20: 191. 1931.) and Laurila (Soc. Zool.-Bot. Fenn. Vanamo
Ann. Bot. 10(4): 15. 1939.) have properly used Vararia; Burt has scat-
tered the species among Corticium, Peniophora and Hypochnus; Overholts
treats one species in Stereum.
In the original publication of C. alutarium three specimens were listed.
Of these the third, Michener’s Pennsylvania collection (Curtis 6349) is
the present species; the second, from Sprague (Curtis 5335) is Penio-
phora filamentosa (Berk. & Curt.) Burt; the first (Curtis 5169) is not in
the Curtis herbarium — at least, not in its proper place. Furthermore, it
seems likely that the number is a mistake, or that a third specimen did
not exist: in Curtis’s lists of fungi sent to Berkeley, 5169 was assigned to
an indeterminate Agaricus. Whatever may have been the case, Burt (I. c.)
designated the Michener collection as the type, and it should be so treated.
CorTICcIUM INVOLUCRUM Burt — Sebacina deminuta Bourd.
Sebacina (Bourdotia) deminuta Bourd., Assoc. Franc. Avanc. Sci. 45: 575.
1922; Rogers, Univ. Iowa Studies Nat. Hist. 15(3): 13. 1933; 17: 41. 1935;
McGuire, Lloydia 4: 39. 1941.
294, | Fartowia, Vor. 1, 1943
Corticium involucrum Burt, Missouri Bot. Gard. Ann. 13: 271. 1926 (!!).
Bourdotia deminuta (Bourd.) Bourd. & Galz., Hymén. de France p. 50. [1928].
As has already been pointed out by Rogers, and by McGuire, Corticium
involucrum Burt may be eliminated from the list of North American
species of Corticium.
CorTICIUM JAMAICENSE Burt = Vararia Peniophoroides (Burt) comb. nov.
Hypochnus Peniophoroides Burt, Missouri Bot. Gard. Ann. 3: 234, 1916 (!!).
Corticium jamaicense Burt, Missouri Bot. Gard. Ann. 13: 273. 1926 (!!).
Vararia Peniophoroides (Burt) comb. nov.
Burt’s species, both of which are based on collections made in Jamaica,
are characterized by compact buffy fructifications, abundant loosely
branched yellow dichophyses, large fusiform gloeocystidia, and subglobose
spores which are yellowish, thick-walled, marked by a prominent apiculus
on the flattened side, and ornamented by blunt hemispherical or short
ridge-like warts. In H. Peniophoroides (represented by quite old ma-
terial) the contents of the gloeocystidia have formed a hyaline, refrac-
tive, vitreous body, cracked, and conforming roughly to the gloeocystid-
ium. In C. jamaicense (represented by excellent specimens) the con-
cretion is brown and less dense. There are no other differences; and
there is no great doubt that the two names apply to the same fungus.
There is no indication on Burt’s specimens of the color of the fresh spores;
but those of the closely related V. effuscata are rosy in mass when fresh,
and fade as those of the present species may well have done.
Asterostromella rhodospora Wakef. (Kew Bul. Misc. Inf. 1915: 372.
1915.), to judge from the description and figures, may well be the same
species. The spore measurements published are the same as those given
for Burt’s species. Actually the spores in the American collections are
larger, running about 7.5-10x 7-9 yp. This difference is not in itself
significant but since it is not feasible to ask for a loan of the type under
present conditions, it seems best not to base a new combination for this
fungus on Miss Wakefield’s species.
The Langlois collection cited by Burt as H. Peniophoroides is not that
species. Donk (Nederl. Myc. Ver. Med. 22: 35. 1933.) cites H. Penio-
phoroides as a synonym of Hypochnus pallescens (q.v.) but it is distinct.
Cortictum Koteroca (Cooke) Hohn. = Pellicularia Koleroga Cooke
Corticium Koleroga (Cooke) Hoéhn.; Burt, Missouri Bot. Gard. Ann. 5: 123. 1918;
13: 292. 1926; Wolf & Bach, Phytopath. 17: 705. 1927.
Pellicularia Koleroga Cooke; Rogers, Farlowia 1: 112. 1943, ubi synon.
CortTIcIuUM LACTEUM Fries Nomen dubium.
In the Burt herbarium, now at the Farlow Herbarium, there are nine-
teen American collections referred to C. lacteum Fries. These have all
been carefully examined. Twelve of them are Corticium radiosum Fries.
Gloeocystidia and the characteristic spores of that species have been dem-
onstrated in each. It is noticeable that most of the collections referred
Rocers & JacKSON: SYNONYMY OF THELEPHORACEAE 295
to above, which prove to be C. radiosum, are rather old and in poor con-
dition. The failure of Burt to observe the gloeocystidia, which are uni-
formly present, may perhaps be explained by his insistence on the exam-
ination of sectioned preparations. The gloeocystidia are readily demon-
strated in properly prepared crushed mounts. Two collections prove to
be Peniophora albula Atk. & Burt (q.v.). The remaining five have not
been identified but appear to be all different from each other and none of
them agrees at all closely with either C. radiosum or the type of C. lacteum
as designated by Burt (Missouri Bot. Gard. Ann. 13: 212. 1926.). In
the herbarium of the Missouri Botanical Garden there are eighteen of
the collections cited by Burt as C. lacteum. Eight of these are C. radiosum
and seven have not been identified, including two which are in very poor
condition but may be C. radiosum. Three of the Canadian collections
made by J. Macoun are as follows: 165 is, in part, C. punctulatum Cooke;
286 is C. albocremeum Hohn. & Litsch.; and 349 is C. vellereum Ellis & Cr.
Litschauer (Osterr. Bot. Zeitschr. 88: 109. 1939.), after having exam-
ined collections in the Romell herbarium which had been determined by
Burt, has concluded that C. lacteum Fries sensu Burt is “Gloeocystidium
alutaceum (Schrad.) Bourd. & Galz. (=C. radiosum Fries).” In the
Burt herbarium there are five collections from the Romell herbarium
(only three of which are cited); all of these are C. radiosum. With this
evidence we can not do other than concur in Litschauer’s conclusion.
Having disposed of C. lacteum sensu Burt, we may now consider other
concepts. C. lacteum sensu Bourdot & Galzin and of Litschauer is quite
a different fungus, better known under the name Corticium tuwberculatum
Karst. Cortictum lacteum sensu Fuckel (Symb. Mycol. 2 Nachtr.: 8.
1873.) should be the same as Peniophora candida Lyman (Science n. s.
25: 290. 1907.). The proper interpretation of C. lacteum (Fries) Fries,
(Epicr. Syst. Mycol. p. 560. 1838.) is quite another matter. It is based
on Thelephora lactea Fries (Syst. Myc. 1: 452. 1821.), which Fries de-
scribed only in the vaguest terms. A fragment of a specimen in the Fries
herbarium, determined by Fries as T. lactea, was sent to Burt by Romell.
It was this specimen which Burt designated as the type and which was to
represent his intended concept. As noted above, his actual concept proves
to be C. radiosum. This specimen has been studied and compared with
the type of Thelephora bombycina Sommerf. (Fl. Lapp. Suppl. p. 284.
1826; =Cortictum bombycinum (Sommerf.) Karst., Hedwigia 32: 120.
1893.). The two are the same fungus. Other and later specimens, as-
signed by Fries to C. lacteum, represent other species; in particular the
authentic specimen at Kew which Burt discusses is quite different, having
more slender hyphae without clamps. It is probable that Thelephora
lactea and the names based on it were no more than designations for
various soft creamy corticioid fungi; it is, therefore, here proposed that
they be in consequence rejected, as nomina dubia, under Art. 63 of
the Rules.
296 Fartowia, VoL. 1, 1943
The alternative is quite clear. Unless there is evidence that some other
specimen of Fries’s under Thelephora lactea is the actual specimen from
which he drew up his description, the specimen designated by Burt as the
type becomes the type, and the name Corticium lacteum, concerning which
there has been so much confusion, replaces the better known C. bomby-
cinum which has never been in doubt. It appears more likely that mere
chance determined that Thelephora lactea should come into conflict with
T. bombycina than that Fries ever had a sharply defined concept of the
former. Frequently it is advantageous to attribute to Fries greater taxo-
nomic acumen than he may have possessed; here it seems more desirable
to recognize confusion, and to reject its results.
CorTIcIUM LAETUM (Karst.) Bres., sensu Burt
An examination of all the American collections cited by Burt (Missouri
Bot. Gard. Ann. 13: 223. 1926.) as this species reveals that the Michigan
collection is Peniophora aurantiaca (Bres.) Hohn. & Litsch., the one from
New York is Cortictum confluens Fries and the North Dakota collection
cannot be named but is neither of the above nor C. laetum. No American
collections have come to light which can be properly referred to this char-
acteristic species. The name should, therefore, be dropped for the present
as a component of the North American flora.
CorTICIUM OCHROFARCTUM Burt — C. pallidum Bres.
Corticium pallidum Bres., Fungi Trident. 2: 59. 1898 (!).
Gloeocystidium pallidum (Bres.) Hohn. & Litsch., K. Akad. Wiss. Wien Math.-Nat.
Kl. Sitzungsber. 116, I: 838. 1907; Bourd. & Galz., Hymén. de France p. 258.
[1928].
Gloeocystidium argillaceum sensu Hohn. & Litsch., Wiesner-Festschr. p. 67. 1908;
nec Corticium argillaceum Bres., Fungi Trident. 2: 63. 1898.
Gloeocystidium pallidum subsp. argillaceum sensu Bourd. & Galz., Soc. Mycol. Fr.
Bul. 28: 363. [1913]; Hymén. de France p. 258. [1928], quantum ad specim,
Corticium ochrofarctum Burt, Missouri Bot. Gard. Ann. 13: 275. 1926 (!!).
The type of Corticium ochrofarctum is in poor condition, but super-
ficially and in all observable microscopic characters exactly equals the
only paratype (Weir 11122) collected the same day and place, which is
good Corticium pallidum. Both collections are from Idaho. This species
has been found to be very common in Ontario as shown by about fifteen
collections so assigned in the herbarium of the University of Toronto. It
is also known from Massachusetts and Oregon and doubtless has a wide
distribution in North America. Corticium Tsugae Burt, which is also
common in Ontario, is closely related. The spores are, however, consist-
ently shorter than in C. pallidum.
Corticitum OvERHOLTsII Burt = Peniophora duplex Burt
Peniophora duplex Burt, Missouri Bot. Gard. Ann. 12: 298. [1926] (Feb.) (!!).
Corticium Overholtsii Burt, Missouri Bot. Gard. Ann. 13: 245. 1926 (Sept.) (!!);
Overh., Mycologia 21: 281. 1929.
Rocers & JACKSON: SYNONYMY OF THELEPHORACEAE 297
The type of C. Overholtsii consists of a young hymenium, just beginning
to sporulate, growing over the surface of an old fructification, probably of
the same fungus. The species was described as possessing neither gloeo-
cystidia nor cystidia. Gloeocystidia are, however, abundant in the type.
While no cystidia were demonstrated in the type collection, all other struc-
tures correspond with those of Peniophora duplex. C. Overholtsii may
best be interpreted as a young growth in which the cystidia have not yet
fully developed.
P. duplex, in its essential structures, is so much like Stereum Pini
(Schleich. ex Fries) Fries as to suggest a close relationship. The differ-
ences are mainly in color of the younger fructifications and in color and
consistency of the older ones. P. duplex at full maturity does not re-
semble at all closely the brown, cracked fructifications characteristic of
S. Pini as it occurs on Pinus Strobus in America or on Pinus sylvestris in
‘urope. It is possible that further study, involving cultures, may show
that P. duplex is best interpreted as a growth phase or variety of S. Pini.
For the present, however, it seems advisable to retain it as a separate
species and leave it in Peniophora where it is most likely to be sought.
CorTICIUM PACTOLINUM Cooke & Hark. Excluded.
Corticium pactolinum Cooke & Hark., Grevillea 9: 81. 1881 (!).
Chromosporium pactolinum (Cooke & Hark.) Cooke, Grevillea 16: 72. 1888.
C. pactolinum is not accounted for in Burt’s Thelephoraceae, presum-
ably because he accepted Cooke’s conclusion that the species was a
Chromosporium.
The Harkness material at hand consists of a thick, compact, cracked
layer of ochre-colored spores, with heavy walls and with the oblique
apiculus of a basidiospore. According to information supplied by Lee
Bonar, the “fructification” on which this species was based consists of a
heavy deposit of the basidiospores of Inonotus Leei Murr.; and certainly
the spores compare favorably with those of the related /. hispidus Karst.
Ellis and Everhart (Jour. Mycol. 6: 79. 1890) ascribe this and two other
species of Chromosporium to Mucronoporus Andersoni Ellis & Everh.
There is here no necessity to pass upon the specific distinctness of I. Leei
or M. Andersoni. It seems clear that C. pactolinum was described from
the spore fall of a polypore (Jnonotus Karst., Xanthochrous Pat.) and
has no place in the Thelephoraceae.
CoRTICIUM PARAPHYSATUM Burt = Aleurodiscus paraphysatus (Burt) comb. nov.
Corticium paraphysatum Burt, Missouri Bot. Gard. Ann. 13: 177. 1926 (!!).
Aleurodiscus paraphysatus (Burt) comb. nov.
This species is based on a collection from Cuba. Three paratypes are
listed, all from the same region, and, like the type, all collected by C. J.
Humphrey. The type specimen shows, scattered through the hymenium,
subglobose bodies, filled with brown resinous material, some appearing
isolated, some stalked; the trama consists in large part of delicate tubules
298 Fartowia, Vou. 1, 1943
of mineral, contorted, about 1 » in diameter, from which all sign of an
enclosed hypha has disappeared. A recent collection made by W. Law-
rence White in Cuba is certainly the same fungus, is in much better con-
dition, and shows the same tubules as well as numerous stout capitate-
cylindric cystidia which at first are colorless but ultimately develop a
brown content which presumably is what is preserved in the weathered
or disintegrating spheres of the type. Spores are numerous, hyaline,
even-walled, clavate, obliquely long-attenuate to the base, 14-18 x 7—7.5 p.
Corresponding bodies occur in the type, scarcely recognizable as spores
since the few that are present have absorbed color from the resin-bearing
structures. A few mature basidia are present in the recent collection and
are clearly the stocky, truncate-cylindric sort found in many species of
Aleurodiscus. The species should be readily recognizable from the spores,
cystidia, and tubules.
CorTIcCIUM PECTINATUM Burt = Vararia pectinata (Burt) comb. nov.
Corticium pectinatum Burt, Missouri Bot. Gard. Ann. 13: 286. 1926 (!!).
Vararia pectinata (Burt) comb. nov. ;
Superficially and structurally this species closely resembles Burt’s
C. racemosum (q.v.). The basidia, ventricose-clavate, long-emergent,
with fragile sterigmata, are much larger in C. pectinatum. The dicho-
physes are the most delicate and obscure encountered, with branches under
0.5 »; and without the confirmation of the basidial type the position of
the species would be doubtful. Gloeocystidia are abundant and conspicu-
ous in the type. The spores, which are difficult to find, are, however, not
infrequent in some preparations. They are fusoid-clavate, strongly tapered
and sharply geniculate below the middle, 11—16.5 x 44.5 yp.
CorRTICIUM PEZIZOIDEUM Ellis & Everh.
Ellis & Everhart’s species was omitted from Burt’s account of the genus.
See under Corticium centrifugum (Lév.) Bres.
CortTIcIuM PILOsuM Burt = Peniophora subalba (Burt) comb. nov.
Corticium pilosum Burt, Missouri Bot. Gard. Ann. 13: 262. 1926 (!!); nec Penio-
Phora pilosa Burt, Missouri Bot. Gard. Ann. 12: 291. [1926].
Corticium subalbum Burt, Missouri Bot. Gard. Ann. 13: 267. 1926 (!!).
Peniophora subalba (Burt) comb. nov.
The types of both species are on Alnus from adjoining Southern states.
They are identical in gross appearance and in microscopic detail. The
appearance of slender paraphyses noted in the original descriptions seems
to be due to collapsed basidia and not to any special structure. The
difference in spore size recorded in the descriptions is not borne out by
an examination of the types. There are cylindrical spores, slightly flat-
tened on one side, with lateral apiculus, measuring 10-13 x4.5-5 yp, in
both type collections.
The species is a gloeocystidiate Peniophora related to Peniophora incar-
nata (Pers.) Karst. and Peniophora aurantiaca (Bres.) Hohn & Litsch.,
Rocers & JACKSON: SYNONYMY OF THELEPHORACEAE 299
and intermediate between the two in the character of the spore. The
cystidia are of the type characteristic of the two species mentioned above
but are very scarce and easily overlooked. They have, however, been
demonstrated in the types of both the species described by Burt.
All of the seven paratypes cited by Burt as C. pilosum have been ex-
amined. Not one is like the type. The single paratype from Georgia
listed under C. subalbum is an immature Aleurodiscus, probably A. botry-
osus Burt.
Corticium Pruni Overh. = C. porosum Berk. & Curt.
See under C. stramineum Bres.
CorTICIUM RACEMOSUM Burt — Vararia racemosa (Burt) comb. nov.
Corticium racemosum Burt, Missouri Bot. Gard. Ann. 13: 287. 1926 (!!).
Vararia racemosa (Burt) comb. nov.
The dichophyses (‘“‘antler-shaped” paraphyses) by which Burt deter-
mined the kinship of this fungus to Corticium investiens (Schw.) Bres.
are obscure and very short-branched, but present. What confirms the
assignment of the species to Vararia is the form of the basidia — some-
thing like those of Alewrodiscus, something like those of the Urnigera
group, but exactly like nothing but those of other Vararia species. They
arise at some depth within the trama, in clusters, as irregular to long-
ellipsoid bodies, and continue by a very slender apical prolongation far
beyond the general level of the hymenium; at the truncate summit are
borne four long, straight, nearly erect sterigmata. The species is dis-
tinguishable from others of the genus by the abundant thin-walled fusi-
form gloeocystidia and the ellipsoid spores.
CORTICIUM ROSEOPALLENS Burt = C. incrustans Hohn. & Litsch.
Corticium incrustans Hohn. & Litsch., K. Akad. Wiss. Wien Math.-Nat. KI.
Sitzungsb. 115, I: 1602. 1906 (!!); Rogers, Univ. Iowa Studies Nat. Hist. 17:
27. 1935.
Corticium roseo-pallens Burt apud Lyman, Boston Soc. Nat. Hist, Proc. 33: 173.
1907 (?!!); Burt, Missouri Bot. Gard. Ann. 13: 240. 1926; Bourd & Galz.,
Hymén. de France p. 215. [1928].
Rogers (l.c.) has already pointed out the identity of the species listed
above. It is not certain that the Vermont specimen designated as the type
by Burt in 1926 is actually the type (i.e., the basis of the 1907 description
in Lyman) ; but there seems no reason to doubt that it correctly represents
the species.
CoRTICIUM ROSEUM (Pers. ex Fries) Fries = Aleurodiscus roseus (Pers. ex Fries)
Hohn. & Litsch.
Thelephora rosea Pers. ex Fries, Syst. Mycol. 1: 451. 1821.
Corticium roseum (Pers. ex Fries) Fries, Epicr. Syst. Mycol. p. 560. 1838; Burt,
Missouri Bot. Gard. Ann. 13: 224. 1926; Bourd. & Galz., Hymén. de France
p. 227. [1928].
300 FarLowia, VoL. 1, 1943
Lyomyces roseus (Pers. ex Fries) Karst., Bidr. Kanned. Finl. Nat. Folk 37: 153.
1882.
Hypochnus roseus (Pers. ex Fries) Schroet. in Cohn, Krypt.-Fl. Schles. 3(1):
417. 1888.
Peniophora rosea (Pers. ex Fries) Massee, Linn. Soc. Bot. Jour. 25: 146. 1889.
Aleurodiscus roseus (Pers. ex Fries) Hohn. & Litsch., K. Akad. Wiss. Wien
Math.-Nat. Kl. Sitzungsb. 115: I: 1568. 1906; Pildt, Ann. Mycol. 24: 219.
1926, ubi synon.
Corticium roseum and its relatives form a small group of species having
large Aleurodiscus-like basidia and characteristic branched paraphyses in
the hymenium. Because the paraphyses are not of the usual type found
in Aleurodiscus some recent authors, including Burt, have preferred to
leave these species in Cortictum. The species are, however, as shown by
the character of the basidia, more properly classified in Aleurodiscus
where they form a sharply circumscribed group. Pilat (1.c.) has placed
them in the subgenus Lyomyces. Two species are recognized in Europe,
the present one and A. polygonioides (Karst.) Pilat. Burt (1. c.) includes
the latter as a synonym of C. roseum, a disposition which seems ex-
tremely doubtful.
CoRTICIUM RUBELLUM Burt = C. confluens Fries
See under Coniophora avellanea Burt.
CORTICIUM SEPTENTRIONALE Burt = C. Litschaueri Burt
Corticium Litschaueri Burt, Missouri Bot. Gard. Ann. 13: 259. 1926 (!!).
Corticium septentrionale Burt, Missouri Bot. Gard. Ann. 13: 257. 1926 (!!).
A comparison of the types of the two species described by Burt leaves
no doubt as to their identity. Since there is no provision in the Inter-
national Rules which makes page priority compulsory we have chosen to
retain the name C. Litschaweri rather than C. septentrionale, for two
reasons: the type collection of the former species is in much better con-
dition than that of the latter; then, too, it seems fitting to conserve the
specific name which honors the late Professor Victor Litschauer, eminent
among the recent European students of the group, and one to whom both
authors of the present contribution are deeply indebted for assistance and
authentic materials.
CorticiuM sociaTUM Burt = Aleurodiscus amylaceus (Bourd. & Galz.)
See under Corticium ermineum Burt.
Cortictum Stevens! Burt = Pellicularia Koleroga Cooke
Corticium Stevensii Burt, Missouri Bot. Gard. Ann. 5: 125. 1918; 13: 293. 1926.
Pellicularia Koleroga Cooke; Rogers, Farlowia 1: 112. 1943.
CoRTICIUM STRAMINEUM Bres. = C. porosum Berk. & Curt.
Corticium porosum Berk. & Curt. apud Berk. & Br., Ann. Mag. Nat. Hist. Ser. 5,
3: 211. 1879; Massee, Linn. Soc, Bot. Jour. 27: 121. 1890; Wakef., Br. Mycol.
Soc. Trans. 4: 341-342. [1914].
Corticium stramineum Bres. apud Brinkm., Hedwigia 39: (221). 1900 (!!);
Brinkm., Westf. Pilze 18. 1900; Burt, Missouri Bot. Gard. Ann. 13: 258. 1926.
Rocers & JACKSON: SYNONYMY OF THELEPHORACEAE 301
®
Gloeocystidium stramineum (Bres.) Hohn. & Litsch., K. Akad. Wiss. Wien Math.-
Nat. KI. Sitzungsb. 116, I: 765. 1907.
Gloeocystidium contiguum sensu Bourd. & Galz., Hymén. de France p. 255. [1928] ;
nec Corticium contiguum Karst., Soc. Faun. Fl. Fenn. Acta 2(1): 39. 1881 (!!).
Gloeocystidium porosum (Berk. & Curt.) Wakef. ex Bourd. & Galz., Hymén. de
France p. 253. [1928].
Corticitum vesiculosum Burt, Missouri Bot. Gard. Ann. 13: 266. 1926 (!!).
Corticitum Pruni Overh., Mycologia 21: 282. 1929 (!!).
Gloeocystidiellum porosum (Berk. & Curt.) Donk, Nederl. Mycol. Ver. Med.
18-20: 156. 1931.
The validity of the use of the name Corticium porosum for this species
has been questioned by Burt (in litt. to Farlow) because of uncertainty
with reference to the type collection, and he has used Bresadola’s name,
C. stramineum. C. porosum was described in a paper by Berkeley and
Broome dealing with “British Fungi.” It was, however, credited to “Berk-
eley and Curtis,” presumably because Curtis had sent a collection from
Venezuela which Berkeley considered to be the same as a British collec-
tion from Aboyne. It is clear from the original account, however, that
the British collection was the intended type since, following the Latin
description, the first sentence of the comment reads as follows: “Aboyne.
Apparently the same with specimens from Venezuela.” When Miss Wake-
field looked into the situation she found two collections in the Berkeley
herbarium, one from Venezuela bearing the number 203, the other from
Glamis (Forfarshire?). No collection from Aboyne could be found.
The Glamis collection, however, answers reasonably well to the original
description while the Venezuela collection does not and she concludes
that “.. . it seems justifiable to regard the Glamis specimen as authentic,
even if it is not the actual type...” In the meantime Massee had com-
mented on this species and cites collections from Venezuela and Aboyne |
with no mention of a collection from Glamis. He writes: “Type in Herb.
Berk. n.4034,” without making it clear whether or not this number refers
to the Aboyne collection. It would seem evident that a collection from
Aboyne was present in the Berkeley herbarium in 1890 and, unless there
is more evidence than is given by Miss Wakefield that the Aboyne and
Glamis collections are one ang the same, this might seem to cast doubt
on the authenticity of the Glamis collection. Two courses seem open:
either the name Corticium porosum should be considered for the present
a nomen dubium or Miss Wakefield’s interpretation accepted and the
Glamis collection in the Berkeley herbarium be accepted as authentic.
Since, however, 1) it is obvious that Berkeley intended the name Cor-
ticlum porosum to apply to a British collection, and 2) there is now in
the Berkeley herbarium a specimen which answers well to his description
even though it may not be the original type, it seems desirable for the
present to concur in Miss Wakefield’s interpretation and accept the Glamis
collection as authentic. It is admittedly quite possible that a different
viewpoint might be found necessary in case the Aboyne collection should
later be disinterred.
302 FartowiA, Vou. 1, 1943
*
The spores of this species are not only amyloid but also minutely rough-
ened. The latter character seems not to have been previously noted. All
collections which we have seen, however, show minute roughening of the
spore wall when mounted in water and examined under an oil immersion
lens. When KOH is used in the mounting fluid, however, the roughening
may not be evident or may gradually disappear. Different collections
vary in the degree of roughening of the spores. Those with evident rough-
ening have been commonly referred to Gl. contiguum sensu Bourd. & Galz.
Four collections from the Bourdot herbarium have been examined. The
spore walls in each show definite roughening but there seems to be
no essential difference between these collections and those of C. porosum.
Four collections also of C. contiguum from the Karsten herbarium have
been examined. Only one of these, and that not the type, is at all com-
parable to the concept of Bourdot & Galzin. Two of them, including the
type, are Corticitum crustaceum Karst. (q. v.). The fourth is Gloeocystid-
ium Karstenti Bourd. & Galz. It seems evident that Bourdot & Galzin’s
concept was not based on the type collection.
Corticium porosum is designated by Donk as the type of Gloeocys-
tidiellum Donk, published as a name for Gloeocystidium sensu Hohn, &
Litsch. (Weisner-Festschr. p. 58. 1908.) , nec Gloeocystidium Karst. (Bidr.
Kanned. Finl. Nat. Folk 48: 429, 1889.).
Bourdot & Galzin included Gloeocystidium clavuligerum Hohn. & Litsch.
as a synonym of Gl. contiguum. An examination of the three collections
in the von Hohnel herbarium reveals that one is typical C. porosum; one
is Gl. furfuraceum Hohn. & Litsch. and the type is not sufficiently like
either to justify including it as a synonym.
Corticium vesiculosum Burt is a form with somewhat shorter and broad-
er gloeocystidia than usual and with spores having evidently roughened
walls, though this latter feature was not noted by Burt. It matches per-
fectly collections referred to Gl. contiguum by Bourdot. Corticium Pruni
Overh. is a form with somewhat more abundant subiculum than usual and
with spores that appear smooth unless closely examined. There is con-
siderable variation in the character of the gloeocystidia in C. porosum and
the differences noted by Overholts do not*seem significant when a large
series of specimens are compared.
CorTICIUM SUBALBUM Burt = Peniophora subalba (Burt)
See under Corticium pilosum Burt.
CorTICIUM SUBCINEREUM Burt = C, contiguum Karst.
See under Corticium crustaceum Karst.
Cortictum suscoronatuM Hohn. & Litsch. = Pellicularia subcoronata (H6hn. &
Litsch.) Rogers
Corticium subcoronatum Hohn. & Litsch.; Overh., Mycologia 26: 510. fig. 11. 1934.
Pellicularia subcoronata (Hohn. & Litsch.) Rogers, Farlowia 1: 104, 1943,
ubi synon.
Rocers & JACKSON: SYNONYMY OF THELEPHORACEAE 303
CORTICIUM SUBGIGANTEUM Berk. = Aleurodiscus subgiganteus (Berk.) Hohn.
Corticium subgiganteum Berk., Grevillea 2: 3. 1873; Burt, Missouri Bot. Gard.
Ann. 13: 215. 1926, ubi synon.
Aleurodiscus subgiganteus (Berk.) Héhn., K. Akad. Wiss. Wien Math.-Nat. KI.
Sitzungsb. 121, I: 342. 1912,
This species was left in Corticium by Burt, presumably because of the
lack of any distinctive type of paraphysis. The character of the basidium,
however, should be the primary determining factor in deciding the generic
position of this species. C. subgiganteum appears to be a true Aleuro-
discus.
The name Michenera artocreas Berk. & Curt., which Burt includes in
the synonymy of this species, is properly based on its imperfect stage;
so is Artocreas Micheneri Berk. & Curt. For a note on the vicissitudes of
these names, see Martin, Wash. Acad. Sci. Jour. 30: 378. 1940. Aleuro-
discus Micheneri (Berk. & Curt.) Massee, Linn. Soc. Bot. Jour. 25: 120.
1889, is both by typification and by description applicable only to the
imperfect stage of A. swbgiganteus.
CoRTICIUM SUBINCARNATUM Peck = Peniophora subincarnata (Peck) Litsch.
Corticium subincarnatum Peck, New York State Mus. Rept. 42: 124. 1889 (!!);
nec Corticium subsulphureum Karst., Soc. Faun. Fl. Fenn. Med. 6: 12. 1881.
Peniophora subincarnata (Peck) Litsch., Osterr. Bot. Zeitschr. 88: 117. 1939.
Burt (Missouri Bot. Gard. Ann. 12: 329, [1926].), included Corticium
subincarnatum as a synonym of Peniophora subsulphurea (Karst.) Hohn.
& Litsch. Litschauer has pointed out the differences between the two
species. All the American collections cited by Burt prove to be P. sub-
incarnata with the exception of that from Idaho (Weir 14), which is typi-
cal P. subsulphurea. A second American collection of the latter species
made at Priest River, Idaho, by Weir on Larix occidentalis, and determined
by Bresadola, is in the herbarium of the Bureau of Plant Industry. Both
species, then, occur in North America. Burt’s description, however, as
pointed out by Litschauer, is obviously drawn from collections properly
referred to P. subincarnata.
The two species are quite similar in general appearance and both occur
also in Europe. Litschauer calls attention to certain color differences
which, however, are not evident in old dried specimens. The chief differ-
ence discernible in the dried material is in the shape and size of the spores.
In P. subincarnata the spores are ellipsoid, 3.5-5x 2-2.5y.. In P. sub-
sulphurea the spores are cylindrical 6-8 x 1.5-2.5 w. Four collections
made by Karsten have been seen. The one cited by Burt is on Pinus
sylvestris and the fungus is P. subsulphurea. A collection distributed as
Xerocarpus subsulphureus Karst., in Roumeg. F. Sel. Exsicc. 4307 on
Pinus sylvestris and another distributed under the same name in Rabenh.-
Wint. Fungi Europ. 2723 on Abies are both P. subsulphurea. A Karsten
collection on Abies in the von Hohnel herbarium is, however, P. sub-
incarnata. It is evident then, that both species occur in Finland, Kar-
304 Fartowira, VoL. 1, 1943
sten’s original description cites the substratum as “ligno vetusto pineo.”
While the type of Karsten’s species has not been seen, it seems reasonable,
from the above evidence, to accept the current concept of Litschauer and
of Bourdot and refer to P. subsulphurea the collections with the longer
cylindrical spore. Specimens from the herbaria of both these authorities
have been examined and are consistently the same.
CortTIcIuM SULPHUREUM sensu Burt = C. bicolor Peck
Ozonium croceum [Kunze] Pers., Mycol. Eur. 1: 86. 1822.
Athelia citrina Pers., Mycol. Eur. 1: 86. 1822; nec Corticium citrinum (Pers.)
Fries, Hymen. Eur. p. 655, 1874 (= Thelephora citrina Pers., Mycol. Eur. 1:
136, 1822.).
Alytosporium croceum ({Kunze] ex Pers.) Link in Linn., Sp. Pl. ed. 4 (Willd.)
6(1): 24. 1824,
? Sporotrichum flavissimum Link. in Linn., Sp. Pl. ed. 4 (Willd.) 6(1): 9. 1824.
Sporotrichum croceum ({Kunze] ex Pers.) Wallr., Fl. Crypt. Germ. 2: 279. 1833.
Corticium bicolor Peck, Buffalo Soc. Nat. Hist. Bul. 1: 62. 1873 (!!); Burt,
Missouri Bot. Gard. Ann. 13: 291. 1926.
Lyomyces byssinus Karst., Soc. Faun. Fl. Fenn. Med. 11: 137. 1884 (!!).
Tomentella byssina (Karst.) Karst., Bidr. Kinned. Finl. Nat. Folk 48: 420. 1889.
Corticium byssinum (Karst.) Massee, Linn. Soc. Bot. Jour. 27: 133. 1890; Bourd.
& Galz., Hymén. de France p. 200. [1928].
Corticium croceum ([Kunze] ex Pers.) Bres., I. R. Accad. Agiati Atti III 3: 112.
1897; Bourd. & Galz., Hymén. de France p. 201. [1928]; nec C. croceum (Pat.)
Sace., Syll. Fung. 11: 124. 1895 (= Aleurodiscus croceus Pat., Soc. Mycol.
Fr. Bul. 9: 133, 1893.).
? Corticium byssinum var. microsporum Bres., Ann. Mycol. 1: 96, 1903; Litsch.,
Bot. Inst. Techn. Hochsch. Wien Mitt. 4: 90. 1927.
Corticium flavissimum sensu Bres. apud Bourd. & Galz., Soc. Mycol. Fr. Bul, 27:
242, 1911; Hymén. de France p. 200. [1928]; an Sporotrichum flavissimum
Link ?
Spicaria croceum ([Kunzel] ex Pers.) Oudem., Enum. Syst. Fung. 1: 281. 1919.
Corticium sulphureum sensu Burt, Missouri Bot. Gard. Ann. 13: 177. 1926; an
Fries, Hymen. Eur. p. 650. 1874, pro parte ?; nec Thelephora sulphurea Pers.
ex Fries.
Corticium byssinum var. plur. Bourd. & Galz., Hymén. de France p. 200. [1928].
(Himantia sulphurea Pers., Roem. Neu. Mag. Bot. p. 122, 1794; nec C. sulphureum
Pers.; nec Thelephora sulphurea (Pers.) Pers. .
(? Sporotrichum flavissimum Link, Ges. Naturf. Fr. Berl. Mag. 7: 34. 1816.
(Sporotrichum croceum Kunze, Mykol. Hefte 1: 81. 1817.
(Athelia citrina Pers., Traité sur les Champ. p. 67. 1819.
There must be considerable uncertainty concerning the early synonymy
of such ubiquitous fungi as those treated in Burt under the names Cor-
ticium sulphureum and Hypochnus fumosus; and the present rather full
treatment is given only because most of the names listed enter into the
problem of the nomenclature of these two fungi. There seems to be no
disagreement among the various mycologists who have treated H. fumosus
sensu Burt (q. v.) that it is actually C. sulphureum Pers. The chief ques-
tion is whether Thelephora sulphurea Pers. ex Fries is the same fungus
as Persoon’s Corticium. Leaving aside the problem whether Persoon’s or
Fries’s fungus determines the application of the name, one may well at-
Rocers & JACKSON: SYNONYMY OF THELEPHORACEAE 305
tempt to discover whether there was ever any divergence between the con-
cepts of those two authors. To judge from Fries’s treatment in the “Hy-
menomycetes Europaei,” there was in 1874, Burt supported his assertion
that Fries’s concept was other than Persoon’s by his account of a single
specimen of C. sulphureum from Fries at Kew which, he wrote, is the same
as his (Burt’s) concept of the species. Burt’s slide is preserved; it shows
no spores; but the subhymenial hyphae are distinct enough, and are often
quite wide, often abruptly swollen at the septa, and clearly nodose-septate.
The Friesian specimen cannot, then, be C. sudphureum sensu Burt, which
has slender, uniform hyphae without clamps; it may well be C. sulphureum
Pers. which has such mycelium as here described.
Corticium byssinum and its varieties, C. bicolor, and probably C. flavis-
simum, have no significant differences beyond color, extremely variable
here, as attested by Bourdot & Galzin’s varieties of C. byssinum. Unless
S. flavissimum can be better established than it is at present, C. bicolor
is the oldest name.
Burt cites Warrensburg, N. Y., as the type locality for C. bicolor. In
the original description, however, Center (now Karner) New York, is
given as the place of collection. In the herbarium of the New York State
Museum there is a sheet of mounted specimens which is marked “type.”
On this sheet beside each piece is marked-either the number “1” or “2”.
On the label the information is furnished that “1” refers to collections
made at Center and “2” to collections made at Warrensburg. It is evident
that Burt overlooked this point. Fortunately the two collections are of
the same species.
Corticium vacum Berk. & Curt. = Pellicularia vaga (Berk. & Curt.) Rogers
Corticium vagum Berk. & Curt., Burt, Missouri Bot. Gard. Ann. 5: 128. 1918;
13: 295. 1926.
Pellicularia vaga (Berk. & Curt.) Rogers ex Linder; Rogers, Farlowia 1: 110.
1943, ubi synon.
Corticium vacum var. SoLANI Burt = Pellicularia filamentosa (Pat.) Rogers
Corticium vagum var. Solani Burt ex Rolfs, Science n. s. 18: 729. 1903; Missouri
Bot. Gard. Ann. 13: 295. 1926.
Pellicularia filamentosa (Pat.) Rogers, Farlowia 1: 113. 1943, ubi synon.
Corticium vesicuLosum Burt = C. porosum Berk. & Curt.
See under Corticium stramineum Bres.
CortTICIUM VINOSOSCABENS Burt = Corticium Macounii Burt
Corticium Macounii Burt, Missouri Bot. Gard. Ann. 13: 256. 1926 (!!).
Corticium vinososcabens Burt, Missouri Bot. Gard. Ann. 13: 267. 1926 (!!).
Corticium maculatum Litsch., Osterr. Bot. Zeitschr. 77: 124. 1928 (!!); J. Weese,
Eumycetes Sel. Exs. 336 (!).
An examination of the three fungi listed above leads to the conclusion
that all are the same. The differences in gross appearance, particularly
those brought out in the two descriptions furnished by Burt, disappear
when a larger series of collections is available. The type of C. Macouni
306 Fartowia, VoL. 1, 1943
is on decorticated firm wood with little development of subiculum. That
of C. vinososcabens is on bark, with relatively abundant development of
loose subiculum. Collections are now available in which both conditions
are present in the same material. A series of collections of C. maculatum
received from Litschauer shows the same variation.
There are no essential microscopic differences noticeable in the three
types. Careful study of the type and comparable collections of C. vinoso-
scabens fails to reveal any structures answering to the “vesicular bodies”
mentioned by Burt. There are present hyphal paraphyses approaching
gloeocystidia mentioned in the description of C. Macounii and figured by
Litschauer for C. maculatum.
The other collections cited by Burt have been examined... Of the collec-
tions assigned to C. vinososcabens, the Wisconsin record is correct but the
New York record proves to be Gloeocystidium Karstenii Bourd. & Galdz.,
-a common species in northeastern North America on Populus, but not
recognized by Burt as such. Those referred to C. Macounii include two
specimens of C. confluens Fries (New Hampshire and New York) and
one (Macoun 86) the same as the type.
GLOEOCYSTIDIUM
GLOEOCYSTIDIUM ALBO-STRAMINEUM (Bres.) Overh. = Corticium geogenium Bres.
See under Peniophora albo-straminea Bres.
GLOEOTULASNELLA
GLOEOTULASNELLA OPALEA Rogers = G, traumatica Bourd. & Galz.
Gloeotulasnella traumatica Bourd. & Galz., Soc. Mycol. Fr. Bul. 25: 32. 1909
(!!); Hymén, de France p. 64. fig. 43. [1928]; Rogers, Ann. Mycol, 31: 197.
1933.
Gloeotulasnella opalea Rogers, Ann. Mycol. 31: 198. 1933 (!!).
Upon comparison of the types, the distinctions upon which G. opalea
was founded vanish. The hypobasidia of G. traumatica are not ovoid, but
long-stalked and subglobose; the long spores described for that form are
free epibasidia. Additional specimens are at hand from Ontario and Iowa.
HYPOCHNUS
HypocuHnus atsus Burt = Aleurodiscus albus (Burt) comb. nov.
Hypochnus albus Burt, Missouri Bot. Gard, Ann. 13: 319. 1926.
Aleurodiseus albus (Burt) comb. nov.
The type of Burt’s species, while certainly not a Tomentella, possesses
a combination of characters which make its true relationship difficult to
determine. The spores are colorless, non-amyloid, when mature with
minutely verrucose wall-markings running about 12 granules to 10 p
(not echinulate as described by Burt). They are obovate, slightly flat-
tened on one side toward the base, 10-13 x 8-9.5 » and not globose, 7.5
Rocers & JACKSON: SYNONYMY OF THELEPHORACEAE 307
in diameter, as described. The cystidia are prominent, projecting nearly
their whole length, 125-200 » long, 8-10 » wide at the middle and
10-17 p» at the base, gradually tapering to an acute apex. The walls of
the cystidia are thick, longitudinally laminate, with narrow lumen and
rugose on the surface. The basidia are cylindric or slightly enlarged
below, 24—26 x 8-10 p, with 4 sterigmata which are at first stout cylindric
and slightly divergent, becoming arcuate, 7-10 » long and 3 » in diameter
at the base. Not much could be made of the structure. A few fine
subicular hyphae with clamps are present, 1—2.5 » in diameter. An abun-
dance of fine particles is present but these seem not to be part of any
structure and no evidence could be obtained of the occurrence of “para-
physes delicate, branching in antler-shaped form.” There is certainly no
structure present which would suggest relationship with Peniophora phyl-
lophila Massee (q. v.), with which Burt compares this species.
The cystidia are of a type not to be expected in Alewrodiscus; they are
such as might be found in Peniophora sec. Tubuliferae subsect. * * of
Bourd. & Galz. But 1) the composition of the fruiting-body, consisting
as it does of a few delicate contorted basal hyphae, imbedded in a mass
of mineral material, and giving rise near the substratum to scattered
basidia, 2) the structure of the basidia, imbedded subglobose bodies
giving rise to thick clavate prolongations which work to the surface and
bear relatively large sterigmata, and 3) the rough-walled spores are all
foreign to any section of Peniophora, and typical of Aleurodiscus. There
scarcely seems to be any limit to the sterile organs that can be associated
with aleurodiscoid structure: witness A. Bertii Lloyd and A. aurantius
(Pers.) Schroet. with gloeocystidia, A. macrosporus Bres. and A. sub-
cruentatus (Berk. & Curt.) Burt with relatively thin-walled cystidia, and
Corticium paraphysatum Burt (q.v.) with capitate cystidia, not to men-
tion the commoner dendrophyses, acanthophyses, and pseudophyses. In
Aleurodiscus the cystidia of A. albus constitute, from the standpoint of
present knowledge of the genus, an innovation: in Peniophora the structure
would make the species wholly a misfit.
HyYpPocHNus FIBRILLOSUs Burt = Corticium fibrillosum (Burt) comb. nov.
Tomentella trigonosperma sensu Hohn. & Litsch., Osterr. Bot. Zeitschr. 58: 334.
1908; Bourd. & Galz., Hymén. de France p. 512. [1928], quantum ad descr. et
specim.; nec Corticium trigonospermum Bres., Ann. Mycol. 3: 163. 1905.
Hypochnus fibrillosus Burt, Missouri Bot. Gard. Ann. 3: 238. 1916 (!!).
Tomentella fibrillosa (Burt) Bourd. & Galz., Hymén. de France p. 513. [1928].
Corticium fibrillosum (Burt) comb. nov.
Since the binomial Tomentella trigonosperma is based on C. trigono-
spermum Bres., even though it has come to be applied to quite a different
species it must be relegated to synonymy and Burt’s specific name taken
up. The species is out of place in Tomentella and closely related to cer-
tain rough-spored species of Corticium, hence a new combination is
proposed.
308 FarLowia, VoL. 1, 1943
Von Héhnel and Litschauer transferred the specific name to Tomentella
in the belief that Bresadola’s Corticium and their fungus were the same.
They described (1.c., and K. Akad. Wiss. Wien Math.-Nat. K]. Sitzungsb.
117, I: 1090-1091. 1908.) a round-angular stage and a globose-spinulose
stage in the development of the spores, placing together what a long series
of European and American specimens show to be constant and separate
organisms. The error is partially explained by the material in von
Hohnel’s herbarium. The sheet for T. trigonospermum carries five
packets: two of Corticium trigonospermum, two containing C. fibrillosum,
and one with a fungus whose spores are smooth and round. Furthermore,
intermixed with C. fibrillosum in one packet is a large proportion of
Peniophora byssoides (Pers. ex Fries) Bres., in which the mycelium is
strikingly similar but the spores oblong and smooth. Presumably the
smooth-spored specimens are the source of the confusion; the rough-spored
specimens show no transitional forms.
HypocHNus FILAMENTOSUS Burt = Corticium sulphureum (Pers. ex Fries) Fries
See under Hypochnus fumosus sensu Burt.
HypocHNus FLAVO-BRUNNEUS Dearn. & Bisby = Coniophora suffocata (Peck) Massee
See under Coniophora Sistotremoides sensu Burt.
Hypocunus FuMosus sensu Burt = Corticium sulphureum (Pers. ex Fries) Fries
Phlebia vaga sensu Fries, Hymen. Eur. p. 625. 1874; Bres., I. R. Accad. Agiati
Atti IIT 3: 105. 1897; an Fries, Syst. Mycol. 1: 428, 1821 ?; nec Corticium
vagum Berk, & Curt., Grevillea 1: 179. 1873.
Thelephora sulphurea Pers. ex Fries, Syst. Mycol. 1: 452. 1821, sp. Himantiae e
synon. excl.; Elenchus Fung, 1: 204. 1828, excl. synon.
Athelia sericea Pers., Mycol. Eur. 1: 85. 1822.
? Thelephora fumosa [Fries] Pers., Mycol. Eur. 1: 147. 1822.
Hyphoderma sulphureum (Pers. ex Fries) Wallr., Fl. Crypt. Germ. 2: 577. 1833.
Hypochnus sericeus (Pers.) Wallr., Fl. Crypt. Germ. 2: 310. 1833.
Corticium sulphureum (Pers. ex Fries) Fries, Epicr. Syst. Mycol. p. 561. 1838
(falso ut ‘sulfureum’); Bres., I. R. Accad. Agiati Atti III 3: 111. 1897; Burt,
Missouri Bot. Gard. Ann. 13: 177. 1926, quantum ad typum Persoonianum,
descr. & spec. Burt. excl.; Bourd. & Galz., Hymén, de France p. 234. [1928].
? Corticium fumosum ([Fries] ex Pers.) Fries, Epicr. Syst. Mycol. p. 562. 1838;
Ic. Fung, 2: 98. pl. 198, fig. 3. 1884.
Odontia fusca Cooke & Ellis, Grevillea 9: 103. 1881.
Lyomyces sulphureus (Pers. ex Fries) Karst., Soc. Faun. Fl. Fenn. Med. 9: 54. 1882.
Coniophora sulphurea (Pers. ex Fries) Quél., Ench. Fung. p. 212. 1886; Massee,
Linn. Soc. Bot. Jour, 25: 132. 1889.
? Tomentella Menieri Pat., Tab. Anal. 2 ser. p. 32. fig. 580. 1886.
Odontia vaga sensu Karst., Soc. Faun. Fl. Fenn. Med. 14: 86. 1887; an Phlebia
vaga Fries ?
? Thelephora Menieri (Pat.) Sacc., Syll. Fung. 6: 547. 1888.
Hypochnus sulphureus (Pers. ex Fries) Schroet. in Cohn, Krypt.-Fl. Schles. 3(1) :
417. 1888.
Tomentella sulphurea (Pers. ex Fries) Karst., Bidr. Kanned. Fin]. Nat. Folk 48:
419. 1889.
? Coniophora fumosa sensu Massee, Linn. Soc. Bot. Jour. 25: 139. 1889; nec C.
fumosa Karst., Soc. Faun. Fl. Fenn. Med. 6: 13. 1881 (=C. olivacea ([Fries]
ex Pers.).
Rocers & JacKSON: SYNONYMY OF THELEPHORACEAE 309
Phlebriella vaga sensu Karst., Hedwigia 29: 271. 1890; an Phlebia vaga Fries ?
Odontia tenuis Peck, New York State Mus. Rept. 44: 134. [1891].
Caldesiella vaga sensu Pat., Essai Taxon. p. 120. 1900; an Phlebia vaga Fries ?
Hypochnus vagus sensu Kauffm., New York State Mus. Bul. 179: 88. 1915; an
Phlebia vaga Fries ?
Hypochnus fumosus sensu Burt, Missouri Bot. Gard. Ann. 3: 239. 1916; an T.
fumosa [Fries] Pers. ?
Hypochnus filamentosus Burt, Missouri Bot. Gard. Ann. 13: 320. 1926 (!!)$ nec
H. filamentosus Pat., Soc. Mycol. Fr. Bul. 7: 163. 1891.
(Corticium sulphureum Pers., Obs. Mycol. 1: 38. 1796.
(Thelephora sulphurea (Pers.) Pers., Syn. Fung. p. 579. 1801; nec Himantia sul-
phurea Pers., Roem. Neu. Mag. Bot. p. 122. 1794 (= Ozonium croceum [Kunze]
Pers., Mycol. Eur, 1: 86. 1822.).
(Hypochnus fumosus Fries, Obs. Mycol. 2: 279. 1818.
As noted under Corticium sulphureum sensu Burt (q. v.), there has
never been any doubt that the present fungus (i.e., Burt’s H. fumosus) is
Persoon’s 7. sulphurea; and whether or not Fries treated only this fungus
as T. sulphurea (he probably had a variety of things), the one Friesian
specimen available for study makes it highly probable that Persoon’s
fungus was also a part of Fries’s assemblage (cf. Massee, l.c.). Of the
other names, only PAlebia vaga Fries is of equal date; and it is not avail-
‘able for use in the genus Corticium. While there is little doubt that P.
vaga was applied in the “Hymenomycetes Europaei” to the present fungus,
the description in the Systema gives no indication of its limits in 1821.
The specific name “fumosa” dates from 1822; but its application is even
more doubtful. F'ries’s description and especially his figure of C. fumo-
sum seem rather to apply to a Coniophora; and Karsten (1881) published
a species in that genus which he equated to “Corticium fumosum Fries
pr. p.” C, sulphureum seems clearly the proper name for the fungus; if
it be for any reason rejected, a name based on Athelia sericea must be used.
HYPOCHNUS ISABELLINUS (Fries ex Pers.) Fries — Pellicularia isabellina (Fries ex
Pers.) Rogers
Hypochnus isabellinus (Fries ex Pers.) Fries; Burt, Missouri Bot. Gard. Ann. 3:
222. 1916.
Pellicularia isabellina (Fries ex Pers.) Rogers, Farlowia 1: 99. 1943, ubi synon.
Hypocunus Lanctoisn Pat. = Pellicularia Langloisii (Pat.) Rogers
Hypochnus Langloisii Pat., Soc. Mycol. Fr. Bul. 24: 3. 1908.
Pellicularia Langloisii (Pat.) Rogers, Farlowia 1: 101. 1943, ubi synon.
The Patouillard species, based on a collection from Louisiana made by
Langlois, was omitted from Burt’s treatment.
HYPOCHNUS PALLESCENS (Schw.) Burt = Vararia pallescens (Schw.) comb. nov. -
Thelephora pallescens Schw., Am. Phil. Soc. Trans. n. s. 4: 167. [1832] (!!).
Thelephora insinuans Schw., Am. Phil. Soc. Trans. n.s. 4: 167. [1832] (!!).
Corticium Thelephoroides Ellis & Everh., Jour. Mycol. 1: 88. 1885 (!!).
Corticium sordulentum Cooke & Massee, Grevillea 16: 69. 1888.
Stereum insinuans (Schw.) Sacc., Syll. Fung. 6: 586. 1888.
Coniophora sordulenta (Cooke & Massee) Sacc., Syll. Fung. 6: 650. 1888.
310 FarLowliA, VoL. 1, 1943
Coniophora insinuans (Schw.) Massee, Linn. Soc. Bot. Jour. 25: 138, 1889.
Corticium pallescens (Schw.) Massee, Linn. Soc. Bot. Jour. 27: 129. 1890.
Stereum duriusculum sensu Bres., Ann. Mycol. 6: 43. 1908; Overh., Mycologia 30:
279. 1938 (falso ut “S. duriusculum Bres.”); nec S. duriusculum Berk. & Br.,
Linn. Soc. Bot. Jour. 14: 66. 1875.
Hypochnus Thelephoroides (Ellis & Everh.) Burt, Missouri Bot. Gard. Ann. 3:
235. 1916.
Hypochnus pallescens (Schw.) Burt, Missouri Bot. Gard. Ann. 4: 267, 1917, ubi
synon.
Asterostromella dura Bourd. & Galz. apud Bourd. & Maire, Soc. Mycol. Fr. Bul.
36: 74. 1920; Hymén. de France p. 396. [1928] (!).
Dichostereum durum (Bourd, & Galz.) Piladt, Ann. Mycol. 24: 223, 1926.
Vararia pallescens (Schw.) comb. nov.
The fructification as ordinarily seen is composed largely of fine dicho-
physes. The basidia are ventricose-tubular and long-exserted, as in the
other members of this genus — never claviform as figured by Bourdot and
by Pilat — but their structure is made out only with some difficulty. The
spores may be sinuolate or appear almost smooth in KOH; iodine brings
out the strongly amyloid roughenings of the spore-wall. Asterostromella
dura is based on a thick, stratose, probably perennial phase of this species
which is occasionally found on partly buried wood or on old roots. It
is such a form that Overholts described as “Stereum duriusculum Bres.”
Hypocunus PENIOPHOROIDES Burt = Vararia Peniophoroides (Burt)
See under Corticium jamaicense Burt.
Hypocunus Potyporoimeus (Berk. & Curt.) Overh. = Corticium Polyporoideum
Berk & Curt.
See under Coniophora Polyporoidea (Berk. & Curt.) Burt.
Hypocunus THELEPHOROIDES (Ellis & Everh.) Burt = Vararia pallescens (Schw.)
See under Hypochnus pallescens (Schw.) Burt.
PENIOPHORA
PENIOPHORA ADMIRABILIS Burt Nomen confusum.
P. admirabilis Burt (Missouri Bot. Gard. Ann. 12: 304. [1926] (! !).)
consists of Tulasnella bifrons Bourd. & Galz. (Soc. Mycol. Fr. Bul. 39:
264. [1924].), growing over the surface of overwintered fructifications
of one or more thelephoraceous fungi. Burt’s description is obviously
drawn from sections, and components of the complex substratum of old
fungi are included. In so far as they have any objective counterpart,
then, the characters of P. admirabilis “were derived from two or more
entirely discordant elements . . . erroneously supposed to form part of
the same individual” (Int. Rules, Art. 64), and the name is not a synonym
of either 7. bifrons or any thelephoraceous form.
PENIOPHORA ALBA Burt = Peniophora Greschikii (Bres.) Bourd. & Galz.
Corticium rude Karst., Bidr. Kanned. Finl. Nat. Folk. 37: 143, 1882 (!!); nec
C. rude Pat., Soc. Mycol. Fr. Bul. 31: 74, 1915.
Corticium Greschikii Bres., Rev. Mycol. 12: 109. 1890 (!!).
Rocers & JACKSON: SYNONYMY OF THELEPHORACEAE 311
Gloeocystidium rude (Karst.) Hohn. & Litsch., K. Akad. Wiss. Wien Math.-Nat.
Kl. Sitzungsb. 115, I: 1558. 1906.
Peniophora subcremea Hohn. & Litsch., K. Akad. Wiss. Wien Math.-Nat. Kl.
Sitzungsb. 115, I: 1600. 1906 (!!); Burt, Missouri Bot. Gard. Ann. 12: 303.
[1926]; Bourd. & Galz., Hymén. de France p. 302 [1928].
Peniophora alba Burt, Missouri Bot. Gard. Ann. 12: 297. [1926] (!!).
Peniophora rudis (Karst.) Bourd. & Galz., Hymén. de France p. 302. [1928];
Litsch. apud Lund. & Nannf., Fung. Exsicc. Suec. 365. 1936; nec P. rudis (Pat.)
Sacc. & Trott., Syll. Fung. 23: 539. 1925.
Peniophora Greschikii (Bres.) Bourd. & Galz., Hymén. de France p. 300. [1928].
Peniophora subcremea var. subuncinata Bourd. & Galz., Hymén. de France p. 302.
[1928].
Corticium Greschikti Bres., based on a single collection made by Gres-
chik in Hungary, has evidently been frequently misinterpreted, most re-
cently by Litschauer (Osterr. Bot. Zeitschr. 88: 116. 1939.), as shown
by his having placed Peniophora hastata Litsch. in synonymy with it.
The types of the two species are quite different, the former agreeing well
in essential characters with specimens commonly referred to P. rudis or
P. subcremea. One of the most characteristic features of the majority
of such collections is the presence of a subicular layer of variable thick-
ness, of thick-walled subcapillary hyphae 2.5-3.5 » in diameter, clamp-
bearing and often parallel-conglutinate. In occasional collections these
basal hyphae are very scarce and easily overlooked. The type of C.
Greschikii is such a collection.
P. alba was based on a single Canadian collection made by J. Macoun
which shows the essential features of P. swhcremea and P. Greschikit.
Burt describes both cystidia and gloeocystidia, but these seem to be only
aspects of a single structure.. The cystidia are obtuse, ventricose-subulate,
thin-walled, and emergent for nearly their entire length. Typical spores
are subcylindric, 5-6 x 2-2.5 »; Bourdot’s variety seems to be based on
spores obliquely drawn out at the apiculus. The types of the four species
here placed in synonymy agree with each other in spore measurements,
almost to the micron.
The species is not uncommon in Canada and the northern United States.
Seven collections from Ontario are so assigned in the herbarium of the
University of Toronto. Burt recorded two collections under P. subcremea,
both of which are the same thing. Three of the seven collections recorded
by Burt under P. subapiculata Bres. (q. v.) prove to be this species.
PENIOPHORA ALBO-STRAMINEA Bres. = Corticium geogenium Bres.
Corticium geogenium Bres., Ann. Mycol. 1: 99. 1903 (!!).
Gloeocystidium inaequale Hohn. & Litsch., K. Akad. Wiss. Wien Math.-Nat. KI.
Sitzungsb. 116, I: 826. 1907 (!!).
Corticium inaequale (Hohn. & Litsch.) Sacc. & Trott., Syll. Fung. 21: 401. 1912.
Gloeocystidium tenue subsp. inaequale (Hohn. & Litsch.) Bourd. & Galz., Soc.
Mycol. Fr. Bul. 28: 365. [1913]; Hymén. de France p. 257. [1928].
Peniophora albo-straminea Bres., Mycologia 17: 69. 1925 (!!); Burt, Missouri
Bot. Gard. Ann. 12: 305 [1926].
Corticium albo-stramineum (Bres.) Overh., Mycologia 30: 275. 1938; nec Corti-
312 Fartow1a, Vou. 1, 1943
cium albo-stramineum (Bres.) Wakef., Br. Mycol. Soc. Trans. 4: 118. 1913
(= Hypochnus albo-stramineus Bres., Ann. Mycol. 1: 110. 1903.).
Gloeocystidium albo-stramineum (Bres.) Overh., Mycologia 30: 276. 1938; nec
G. albo-stramineum (Bres.) Hohn. & Litsch., K. Akad. Wiss. Wien Math.-Nat.
Kl. Sitzungsb. 117, I: 1097. 1908 (=H. albo-stramineus Bres.).
This species is a gloeocystidiate form, usually with prominently project-
ing cylindrical gloeocystidia, as was pointed out by Overholts. It may
be recognized among related species by the non-amyloid, ellipsoid spores
6-8.5 x 44.5 p, not appreciably flattened on one side, with slightly thick-
ened walls. It may occur on wood or bark of either coniferous or decidu-
ous trees. The species has apparently not been collected frequently in
North America. Two or three collections were made by Weir at the
type locality of P. albo-straminea, all on Alnus tenuifolia. One collection
made in Ontario has been assigned to this species. The Vermont collec-
tion cited by Burt under Corticitum ermineum (q. v.) belongs here. It
has been reported correctly (as C. albo-stramineum (Bres.) Overh.) from
Manitoba by Bisby e¢ al. (Fungi of Manitoba and Saskatchewan p. 75.
1938.). The collection from California listed by Burt as P. albo-stra-
minea is sterile and cannot be identified, but is certainly not the present
species,
Von Hohnel & Litschauer (K. Akad. Wiss. Wien Math.-Nat. KI.
Sitzungsb. 117, I: 1087. 1908.), on the basis of the description and a
specimen (Jaap. Sachsenwald. 1904) received from Bresadola labeled
Gloeocystidium Bourdotii Bres. n. sp., which they assumed to be the same
as G. praetermissum var. Bourdotii Bres. (Ann. Mycol. 6: 44. 1908.),
place the latter in synonymy under G. inaequale. The collection cited
has been examined and is certainly that species. However, Bourdot &
Galzin (Hymén. de France p. 252. [1928].) list this variety, which is
based on collections from Allier, France, under Gloeocystidium luridum
(Bres.) Hohn. & Litsch. No authentic collections from France are
available for study.
PENIOPHORA ALBULA Atk. & Burt = Corticium albulum (Atk. & Burt) comb. nov.
Peniophora Torrendii Bres. apud Torrend. Brotéria Ser. Bot. 11: 77. 1913 (!!);
Bourd. & Galz., Hymén. de France p. 315. [1928]; nec Corticium Torrendii
Bres., I. R. Accad. Agiati Atti III, 8: 131. 1902 (= Gloeocystidium Torrendii
(Bres.) Bres., Brotéria Ser. Bot. 11: 81. 1913.).
Peniophora albula Atk. & Burt apud Burt, Missouri Bot. Gard. Ann. 12: 231.
[1926] (!!).
Peniophora Burkii Burt. Missouri Bot. Gard. Ann. 12: 282. [1926] (!!).
Corticium albulum (Atk. & Burt) comb. nov.
The so-called cystidia described for this species are best interpreted
as gloeocystidia which may project slightly. Among forms so character-
ized this species may be readily recognized by the spores which are non-
amyloid, obovate, slightly flattened at one side below the middle, with
lateral apiculus, 6-8 x 4-5 y; these tend to cling together in groups of
two to four. The gloeocystidia are often rather scarce, flexuous or more
commonly subfusiform, 30-50 x 6-7 p, tapering to an obtuse apex.
Rocers & JAcKSON: SYNONYMY OF THELEPHORACEAE 313
PENIOPHORA ALLESCHERI (Bres.) Sacc. & Syd. =P. mutata (Peck) Hohn. & Litsch.
Corticium mutatum Peck, New York State Mus. Rept. 43: 69. 1890 (!).
Corticitum Allescheri Bres., Fungi Trident. 2: 62. 1898 (!!).
Peniophora Allescheri (Bres.) Sacc. & Syd., Syll. Fung. 16: 194. 1902; Burt,
Missouri Bot. Gard. Ann. 12: 301. [1926].
Kneiffia Allescheri (Bres.) Bres., Ann. Mycol. 1: 100. 1903.
Peniophora mutata (Peck) Hohn. & Litsch., K. Akad. Wiss. Wien Math.-Nat. KI.
Sitzungsb. 115, I: 1580. 1906; Burt, Missouri Bot. Gard. Ann. 12: 299, [1926].
Gloeopeniophora Allescheri (Bres.) Hohn. & Litsch., K. Akad. Wiss. Wien Math.-
Nat. Kl. Sitzungsb. 117: I: 1082. 1908.
Peniophora cremea subsp. Allescheri (Bres.) Bourd. & Galz., Hymén. de France
p. 304. [1928], quantum ad typum, descr. excl.
Peniophora cremea var. Allescheri (Bres.) Litsch., Osterr. Bot. Zeitschr. 88: 117.
1939, quantum ad typum.
Bresadola cites specimens from Bavaria, collected by Allescher, and
from Italy. Since the fungus is named after Allescher, his collection is
properly to be taken as the type. A part of it is in the Burt herbarium;
it agrees extremely well, as Burt has pointed out, with the upper figure
in Bresadola’s plate; and equally well with Peck’s C. mutatum. Von
Hohnel & Litschauer (1908), and Bourdot have both indicated that two
species were included in the original material, the present one and P.
cremea (Bres.) Sacc. & Syd. Bourdot’s subspecies and Litschauer’s
variety are described from the “cremea’ portion; nevertheless, being
based on “Allescheri,” and hence on its type, they must be included here.
PENIOPHORA ALUTARIA Burt = P. pallidula (Bres.) Bres.
Gonatobotrys pallidula Bres., Ann. Mycol. 1: 127. 1903 (!!).
Gloeocystidium oleosum Hohn. & Litsch., K. Akad. Wiss. Wien Math.-Nat. KI.
Sitzungsb. 116, I: 827. 1907 (!!).
Gloeocystidium pallidulum (Bres.) Hohn. & Litsch., Osterr. Bot. Zeitschr. 58:
471. 1908.
Corticium oleosum (Hohn. & Litsch.) Sacc. & Trav., Syll. Fung. 19: 437. 1910.
Peniophora pallidula (Bres.) Bres. apud Bourd. & Galz., Soc. Mycol. Fr. Bul.
28: 390. [1913]; Hymén. de France p. 296. [1928].
Peniophora pallidula var. regenerans Bourd. & Galz., Soc. Mycol. Fr. Bul. 28:
390. [1913]; Hymén. de France p. 296. [1928].
Peniophora alutaria Burt, Missouri Bot. Gard. Ann. 12: 332. [1926] (!!).
Peniophora laminata Burt, Missouri Bot. Gard. Ann. 12: 246. [1926] (!!).
This is apparently one of the most common species in northern North
America, although it was not recognized by Burt, except under his new
names to which were assigned only three specimens all told. The two
Canadian collections made by J. Macoun and cited by Burt as Peniophora
Odontioides Burt (1. c. p. 224) are this species and this fact perhaps
accounts for his having described the cystidia of the former as septate,
since they are not so in the type collection. P. pallidula has also been
encountered among collections assigned by him to Peniophora sub-
apiculata (Bres.) Burt (q. v.).
The species is quite variable in character of both basidia and spores,
as is suggested by the varieties described by Bourdot & Galzin (I. c.).
314 FarLowliaA, VoL. 1, 1943
P. laminata Burt corresponds to the var. regenerans Bourd. & Galz. P.
alutaria is based on a form at least as common in America as the typical
one, a form whose cystidia show fewer septa and less irregularity in
diameter, whose spores are consistently larger, and which bears in the
hymenium numerous slender acicular cystidia with a sharp, barb-like
terminal incrustation —like those illustrated by Miller (Mycologia 26:
pl. 3, fig. 4. 1934.) and by Bourdot (Hymén. de France, fig. 114) for
Odontia arguta. There is some doubt whether P. pallidula is separable
from O. arguta; it is retained for the present.
PENIOPHORA ARACHNOIDEA Burt = P. é¢remea (Bres.) Sacc. & Syd.
Corticium sordidum Karst., Soc. Faun. Fl. Fenn. Med. 9: 65. 1882 (!).
Corticium cremeum Bres., Fungi Trident. 2: 63. 1898 (!).
Peniophora cremea (Bres.) Sacc. & Syd., Syll. Fung. 16: 195. 1902; Burt,
Missouri Bot. Gard. Ann. 12: 261. [1926]; Bourd. & Galz., Hymén. de France
p. 303. [1928].
Corticium Eichlerianum (Bres.) Bres., Ann. Mycol. 1: 95. 1903 (!!).
Kneiffia cremea Bres., Ann. Mycol. 1: 100. 1903.
Peniophora sordida (Karst.) Hohn. & Litsch., K. Akad. Wiss. Wien Math.-Nat. KI.
Sitzungsb. 115, I: 1559. 1906 (Oct. ?); Burt, Missouri Bot. Gard. Ann. 12:
280. [1926]; Bourd. & Galz., Hymén. de France p. 305. [1928]; nec P. sordida
(Schroet.) Hohn. & Litsch., Ann. Mycol. 4: 290. 1906 (June).
Peniophora cremea var, glaucescens Bourd. & Galz., Soc. Mycol. Fr. Bul. 28:
396. [1913].
Peniophora Eichleriana (Bres.) Bourd. & Galz., Soc. Mycol. Fr. Bul. 28: 397.
[1913]; nec P. Eichleriana sensu Bourd. & Galz., Hymén. de France p. 306.
[1928].
Peniophora arachnoidea Burt, Missouri Bot. Gard. Ann. 12: 220. [1926] (!!).
Peniophora cremea var. Eichleriana (Bres.) Bourd. & Galz., Hymén. de France
p. 304. [1928].
Cerocorticium albissimum Rick, Brotéria Sér. Trim. Ciénc. Nat. 3: 169, 1934 (!).
The type of P. arachnoidea Burt is as here indicated. Some of the
other collections assigned to P. arachnoidea by Burt are P. byssoides
(Pers. ex Fries) Bres. (see under Coniophora byssoidea) ; others are a
white form of the latter which may well be P. byssoides subsp. Tomentella
(Bres.) Bourd. & Galz. if that subspecies can be recognized as distinct
from P. byssoides. The description provided by Burt would appear to
be a composite one, certainly not drawn exclusively or even mainly from
the designated type.
Corticium sordidum, as represented by an authentic specimen in Burt’s
herbarium and also by Bourdot’s description, is a name applied to older
specimens of C. cremeum; both Bourdot and Burt remark the stratifica-
tion of their specimens. The largest subicular hyphae in the Karsten
specimen studied occasionally attain a diameter of 12 », and have thicker
walls than ordinarily occur in P. cremea, but most hyphae are exactly
the same; for the most part, this material and the Bresadola material of
P. cremea agree well. The cystidia in C. sordidum are mostly engulfed
by the thickening mycelium, and the tips which protrude are mostly
broken; but the organs are the same in the two specimens.
Rocers & JACKSON: SYNONYMY OF THELEPHORACEAE 315
Having already transferred Schroeter’s Hypochnus sordidus to Penio-
phora (see under P. Peckit), von Héhnel & Litschauer discovered Kar-
sten’s earlier Corticium sordidum and likewise published it as a Penio-
phora, renaming the earlier fungus P. sordidella. The binomial had then
been published twice before Burt sought to create it again (1. c.). Be-
cause of the existence of P. sordida (Schroet.) Hohn. & Litsch., Bresa-
dola’s later C. cremewm furnishes the name for the species.
Von Hohnel & Litschauer (K. Akad. Wiss. Wien Math.-Nat, KI.
Sitzungsb. 117, I: 1082. 1908.) reduced C. Eichlerianum to synonymy
under Peniophora velutina ({DC.] ex Pers.) Cooke. To this Bresadola
objected (Ann. Mycol. 9: 425, 1911.), asserting that there were no
cystidia present. In the type specimen, however, there are cystidia,
differing in no way from those of P. cremea; they are somewhat scattered,
and for the most part broken. Bourdot & Galzin, having published
P. Eichleriana on the basis of the type specimen of Bresadola’s Corticium,
in 1928 reduced that form to varietal status under P. cremea and sought
to use the specific name P. Fichleriana for another collection, identified
by Bresadola as C. Eichlerianum, but quite a different species. Clearly
that other species must have another name.
PENIPHORA ARGENTEA Ellis & Everh. ex Burt Excluded.
P. argentea Ellis & Everh. in herb. ex Burt (Missouri Bot. Gard. Ann.
12: 346. [1926].) is based on a thin sterile fungus growth whose dis-
torted, conical seta-like structures and color-reaction in KOH seem to be
those of an abortive Hymenochaete. The silvery surface layer which gave
the specific name appears to be amorphous; at least in stained prepara-
tions there is no indication of protoplasm or hyphal structure in it.
The name should be dropped.
PENIOPHORA ASPERIPILATA Burt = P. longispora (Pat.) Hohn. :
Hypochnus longisporus Pat., Jour. de Bot. 8: 221. 1894 (!!).
Kneiffia longispora (Pat.) Bres., Ann. Mycol. 1: 105. 1903.
Peniophora longispora (Pat.) Hohn., Ann. Mycol. 3: 325. 1905; Burt, Missouri
Bot. Gard. Ann. 12: 229. [1926]; Bourd. & Galz., Hymén. de France p. 298.
[1928].
Peniophora longispora var. mycelialis, var. gloeocystidiata, var. clavispora, var.
cylindrospora Bourd. & Galz., Soc. Mycol. Fr. Bul. 28: 392. [1913]; Hymén.
de France p. 298. [1928].
Peniophora asperipilata Burt, Missouri Bot. Gard. Ann. 12: 230. [1926] (!!).
An examination of the portions of the type collection of P. asperipilata
in the Burt herbarium and in the herbarium of the United States Depart-
ment of Agriculture leaves no doubt that this is P. Jongispora. The char-
acteristic elongated subfusiform spores of that species are present in both
portions. Burt described the spores as “subglobose, 344-4 » in diam-
eter”; no such spores could be found. The paratype, Lloyd Herb. 2395,
is the same species,
316 FarLtowiA, VoL. 1, 1943
PenropHora Burk Burt = Corticium albulum (Atk. & Burt)
See under Peniophora albula Atk. & Burt
PENIOPHORA CANADENSIS Burt =P. polonensis (Bres.) Hohn. & Litsch.
Kneiffia polonensis Bres., Ann. Mycol. 1: 103. 1903 (!!).
Peniophora polonensis (Bres.) Hohn. & Litsch., Ann. Mycol. 4: 292. 1906 (falso
ut “poloniensis”) ; Bourd. & Galz., Hymén. de France p. 315. [1928].
Peniophora canadensis Burt, Missouri Bot. Gard. Ann, 12: 260. [1926] (!!).
Two collections of P. canadensis are recorded by Burt. Both are identi-
cal with the Bresadola species. The fungus is apparently common in
northeastern North America, as shown by over thirty Ontario collections
in the University of Toronto herbarium.
PENIOPHORA DELECTANS Overh. = P. calothrix (Pat.) comb. nov.
Corticium calothrix Pat., Cat. Rais. Pl. Cel. Tunisie p. 59. 1897 (!!).
Peniophora glebulosa subsp. Pirina Bourd. & Galz., Soc. Mycol. Fr. Bul. 28:
387. [1913].
Peniophora Pirina (Bourd. & Galz.) Bourd. & Galz., Hymén. de France p. 290.
[1928].
Peniophora delectans Overh., Mycologia 26: 513. 1934 (1!!).
Peniophora calothrix (Pat.) comb. nov.
Readily distinguishable among the species of Peniophora sect. Tubuli-
ferae by the cystidia, which have the lumen capillary below and expanded
towards the tip, leaving the outermost portion thin-walled except for a
narrow region extending along one side and over the summit, forming a
sort of hood of thicker material. Not common, but collected on both
coasts, in Iowa, and as far south as Tennessee.
PENIOPHORA DISSOLUTA Overh. = P. chaetophora (Hoéhn.) Hohn. & Litsch.
Hypochnus chaetophorus Hohn., K. Akad. Wiss. Wien Math.-Nat. KI. Sitzungsb.
111, I: 1007. 1902 (!!).
Peniophora chaetophora (Hohn.) Hohn. & Litsch., K. Akad. Wiss. Wien Math.-
Nat. KI, Sitzungsb. 116, I: 748. 1907; Bourd. & Galz., Hymén. de France p.
287. [1928].
Peniophora dissoluta Overh., Mycologia 26: 514.1934 (!!).
The fungus described by Overholts from Pennsylvania is a thin young
growth of P. chaetophora, like the margins of older and more mature
collections. The species is common in eastern Canada and western
Oregon, and probably elsewhere also. It is well represented by von
Hohnel & Litschauer’s figure (I. c.).
PENIOPHORA FLAMMEA Burt = P, ludoviciana Burt
Peniophora ludoviciana Burt, Missouri Bot. Gard. Ann. 12: 244. [1926] (!!).
Peniophora flammea Burt, Missouri Bot. Gard. Ann. 12: 252, [1926] (!!).
The fructification of this species is brilliant greenish yellow — Primu-
line Yellow, Old Gold, Olive Lake (R) — and soft-waxy when fresh; on
drying the color becomes a brownish buff; and since it is the latter
shades that are described (1. c.), the species has been hard to identify
because of the striking appearance of fresh material. The cystidia are
Rocers & JACKSON: SYNONYMY OF THELEPHORACEAE 317
for some time even and thin-walled, but finally become incrusted; in the
latter condition they become the sole diagnostic character of P. flammea.
The types of both species show both stages in cystidial development and
both represent a thin form of the species which commonly occurs in
thicker fructifications than allowed by Burt’s descriptions.
PENIOPHORA GLEBULOSA (Fries) Sacc. & Syd. Nomen confusum,
See under Corticium calceum sensu Romell & Burt.
PENIOPHORA GLEBULOSA sensu Bres. & auctt. = P. gracillima Ellis & Everh. sp. nov.
in sched.
Peniophora glebulosa sensu Bres., Burt, Missouri Bot. Gard. Ann, 12: 282. [1926];
Bourd. & Galz., Hymén. de France p. 288. [1928]; nec Thelephora calcea *
glebulosa Fries (nomen confusum).
(Peniophora gracillima Ellis & Everh. ex Hohn. & Litsch., K. Akad. Wiss. Wien
Math.-Nat. KI], Sitzungsh. 116, I: 743. 1907, pro synon.
Peniophora gracillima Ellis & Everh. sp. nov. in sched.’
Under Corticium calceum sensu Romell & Burt will be found a list of
names all of which are based directly or indirectly on Thelephora calcea
Pers. *glebulosa Fries, the type of which is found to consist of a mixture
of two species, and which is hence a nomen confusum.
The cystidiate species of this complex, which has been commonly known
in Europe and America as Peniophora glebulosa, is therefore without a
valid name. The herbarium name of Ellis & Everhart, which has ap-
peared in the literature (Hohn. & Litsch. 1. c.), is available and seems
appropriate. It is here published, not as a new name but as a new
species, in order that the type may be definitely fixed.
PENIOPHORA GLOBIFERA Ellis & Everh. =P. gigantea (Fries) Massee
Peniophora gigantea (Fries) Massee, Linn. Soc. Bot. Jour. 25: 142. 1889; Burt,
Missouri Bot. Gard. Ann. 12: 216. [1926]; Bourd. & Galz., Hymén. de France
p. 318. [1928], ubi synon.
Peniophora globifera Ellis & Everh., Amer. Nat. 31: 340. 1897 (!!); Burt,
Missouri Bot. Gard. Ann. 12: 219. [1926].
Kneiffia globifera (Ellis & Everh.) Rick, Brotéria Ser. Trim. Ciéne. Nat. 3: 77. 1934.
A comparison of the type of P. globifera with a series of collections of
P. gigantea from Europe and North America leaves little doubt that the
former is merely a variation of the latter. It seems an unnecessary and
very doubtful species.
PEeNiopHORA KAUFFMANII Burt = P. heterocystidia Burt
Peniophora heterocystidia Burt, Missouri Bot. Gard. Ann. 12: 293, [1926] (!!).
Peniophora Kauffmanii Burt, Missouri Bot. Gard. Ann. 12: 296. [1926] (!!).
2 Fructificatio sub lente subtiliter pilosa, cystidiis setulosa, continua; hyphae
2.5-3.5 y. diam., rigidae, nodoso-septatae; cystidia subcylindracea, obtusa, 55-135 x
8-10 pw, in partibus’ basalibus et mediis crassiter tunicata, apicem versus canalicula
abrupte expansa, ut tunica tenuis fit; basidia cylindraceo-clavata, 18-20 x 44.5 u,
sterigmata 4 gerentia; sporae cylindraceae, subrectae vel leniter curvae, 7-9 (—11) x
cere
In ligno Aceris putrido, Newfield, N. J., Aug. 1885, J. B. Ellis, typus, in Farlow herb,
318 Fartowia, Vou. 1, 1943
P. Kauffmanii was described from a single collection made by Kauff-
man in Kentucky. No mention was made of large cystidia (or compact
aggregations of crystals) such as are usually characteristic of P. hetero-
cystidia, and they do not occur in the portion of the type at the Missouri
Botanical Garden. In the portion at the Farlow herbarium, however, and
another received through Dr. Mains from the University of Michigan
herbarium, they are clearly present in certain areas. There are no other
differences, and the species are synonymous. P. heterocystidia is closely
related to P. mutata (Peck) Héhn. & Litsch. (see under P. Allescheri
Bres.).
PENIOPHORA LAEVIS sensu Burt = P. affinis Burt
Peniophora laevis sensu Burt apud Peck, New York State Mus. Bul. 54: 954. 1902;
Missouri Bot. Gard. Ann. 12: 257. [1926]; Bourd. & Galz., Hymén. de France p.
307. [1928]; nec Thelephora laevis Pers. (Syn. Fung. p. 575. 1801.) ex Fries,
Syst. Mycol. 1: 451. 1821 (= Corticium laeve Pers., Roem. Neu. Mag. Bot. p.
110. 1794; Fries, Epicr. Syst. Mycol. p. 560. 1838; Burt, Missouri Bot. Gard. Ann.
13: 280. 1926; Bourd. & Galz., Hymén. de France p. 183. [1928]; Thelephora
evolvens Fries, Syst. Mycol. 1: 441. 1821; Corticium evolvens (Fries) Fries
Epicr. Syst. Mycol. p. 557. 1838.).
Kneiffia laevis sensu Bres., Ann, Mycol. 1: 100. 1903.
Peniophora affinis Burt, Missouri Bot. Gard. Ann. 12: 266. [1926] (!!).
Peniophora laevis var. affinis (Burt) Litsch., Osterr. Bot. Zeitschr. 88: 117. 1939.
,
As in dealing with Thelephora umbrina, Burt here attempted to make
a single published name do duty for two distinct species in two genera.
Thelephora laevis Pers. ex Fries seems to be sufficiently well known; both
Burt and Bourdot have described it under Corticium laeve and the former
had a specimen from Persoon in his herbarium. Fries did not, of course,
alter the diagnostic characters or circumscription of the Thelephora
when he transferred it back to Persoon’s original name, Corticium laeve.
But sometime down the years he sent to Berkeley under that name a mis:
determined specimen with cystidia; that specimen is the basis of Burt’s
description of P. laevis; but the type of the Corticium is still the type of
T. laevis Pers, ex Fries (i. e., as described in Burt under C. laeve).
Fortunately Burt also published P. afinis, whose affinity to “P. laevis”
is so great that they cannot be distinguished; the comb. nov. then drops
out and the sp. nov. takes its place. It may be mentioned here, as indi-
cated in the synonymy listed above, that there is only page priority in
Fries to justify the use of Corticium evolvens to replace the more com-
monly used binomial Corticium laeve. Since page priority is not rec-
ognized by the Rules, and since C. laeve is well established by usage, we
have not included a discussion of the latter in this series of notes.
Corticium calotrichum Karst., cited by von Héhnel & Litschauer (K.
Akad. Wiss. Wien Math.-Nat. Kl. Sitzungsb. 115, I: 1550. 1906.) as
synonymous with P. laevis, according to authentic material is wholly
different.
Rocers & JACKSON: SYNONYMY OF THELEPHORACEAE 319
PENIOPHORA LAMINATA Burt = P. pallidula Bres.
See under P. alutaria Burt.
PENIOPHORA LEPIDA Bres. = P. aurantiaca (Bres.) Hohn. & Litsch.
See under Aleurodiscus Zelleri Burt.
PEeNropHorRA LIMONIA Burt = P. sanguinea (Fries) Hohn. & Litsch.
Peniophora sanguinea (Fries) Hohn, & Litsch., K. Akad. Wiss. Wien Math.-Nat.
KI. Sitzungsb. 115, I: 1588. 1906; Burt, Missouri Bot. Gard. Ann. 12: 274.
[1926], ubi synon; Bourd. & Galz., Hymén. de France ‘p. 312. [1928].
Peniophora sanguinea subsp. anaemacta Bourd. & Galz., Soc. Mycol. Fr. Bul. 28:
395. [1913]; Hymén. de France p. 313. [1928] (!).
Peniophora limonia Burt, Missouri Bot. Gard, Ann. 12: 275. [1926] (!!).
Peniophora miniata sensu Burt, Missouri Bot. Gard. Ann. 12: 277. [1926]; an
Thelephora miniata Berk. in Smith, Engl. Fl. 5(2): 168. 1836 (?!!); Corti-
cium miniatum (Berk.) Berk., Grevillea 1: 178. 1873. ? ; vel C. miniatum
Cooke, Grevillea 9: 2, 1880. ?
According to Bourdot, P. sanguinea subsp. anaemacta is a cream-colored
form with the “micrographic characters of P. sanguinea”; a specimen
from him is marked: “Certissime cum P. sanguinea conjuncta!”’ Speci-
mens answering this description are at hand from Iowa, where the red
form seems not to develop, and elsewhere; the type of P. limonia is just
such a fructification. The type distribution of T. miniata (Berk., Br.
Fung. 251) shows only sterile mycelium, as already noted by von Hohnel
& Litschauer (1. c.); its true position is uncertain, but it probably be-
longs here.’ Fries assigned the same fungus to C. sanguineum (Summ.
Veg. Scand. p. 334, 1849.). Specimens referred by Burt to P. miniata
show the whole range of variation exhibited by P. sanguinea, including
hyphal incrustation and staining of the woody substratum.
It seems easily possible that the present species should derive its name
from Thelephora bolaris Pers. (Mycol. Eur. 1: 138. 1822.), based on
Sowerby’s plate 291 of Auricularia aurantiaca. When Fries published
his Thelephora sanguinea he had not yet seen that plate (cf. Elenchus
Fung. 1: 219. 1828.); but at the time of compiling the index to the
“Systema” he apparently had done so, for he assigned both T. bolaris and
A. aurantiaca to his later T. sanguinea (cf. Syst. Mycol. 3, index 54,
185), as he did later in the “Epicrisis’ (p. 561). Sowerby’s plate
assuredly resembles the present species, about as much as does Fries’s
own plate (Ic. Fung. 2: 97, pl. 198, fig. 2. 1884.). But by the time
he was writing the “Hymenomycetes Europaei,” Fries had reverted to
the opinion that Sowerby’s—and hence Persoon’s— fungus was a
Phlebia; and the description bears out this surmise just sufficiently that
for the present the later Friesian name is here retained.
PENIOPHORA MAGNAHYPHA Burt = Pellicularia Langloisii (Pat.) Rogers
Peniophora magnahypha Burt, Missouri Bot. Gard. Ann. 12: 238. {1926].
Pellicularia Langloisii (Pat.) Rogers, Farlowia 1: 101. 1943, ubi synon.
PENIOPHORA MINIATA sensu Burt = P. sanguinea (Fries) Hohn. & Litsch.
See under Peniophora limonia Burt.
320 Fartowia, VoL. 1, 1943
PENIOPHORA ODORATA sensu Burt = Peniophora luna Romell in sched. sp. nov.
Stereum odoratum sensu Karst., Bidr. Kanned. Finl. Nat. Folk 25: 309. 1876;
nec S. odoratum Fries, Epicr. Syst. Mycol. p. 553. 1838 (= Thelephora odorata
Fries, Syst. Mycol. 1: 445. 1821.).
Xerocarpus odoratus sensu Karst., Rev. Mycol. 3(9): 22. 1881; Bidr. Kanned.
Finl. Nat. Folk 37: 137, 1882, typo excluso.
Phanerochaete odorata sensu Karst., Bidr. Kainned. Finl. Nat. Folk 54: 186. 1894
(!), typo excluso.
Peniophora odorata sensu Burt, Missouri Bot. Gard. Ann. 12: 289. [1926] (!);
Bourd. & Galz., Hymén. de France p. 295. [1928], typo excluso.
Nec Stereum Karstenii Bres., I. R. Accad. Agiati Atti III 3: 108. 1897.
Peniophora luna Romell in sched. sp. nov.’
No Friesian specimens are at hand to verify independently the state-
ments of various authors (including Karsten himself, |. c., 1894), that the
“odorata” of Karsten is not the Friesian species; but Fries’s descriptions,
and especially the long one (Fries, Elenchus Fung. 1: 207. 1828.),
tend to confirm that opinion. “Semper effusa, demum ad plures ulnas
truncos ambiens”; “numquam rimoso-incisa, demum tota ut corium a
truncis solubilis” —these words do not well describe the present notably
pulvinate form, which only rarely, and only tardily, becomes confluent
over even small areas, and which is often rimose. It is, of course, not
permissible to choose a type for P. odorata from among Karsten collec-
tions so labeled, as Burt has done; the Karsten references clearly indi-
cate that it is Fries’s species with which he intended to deal. Since an
appropriate name was at hand, on Romell material of the present
species, it is here published — and as a new species, rather than a new
name, in order that the type may be fixed beyond question. Stereum
Karstenii, published by Bresadola for “Xerocarpus odoratus Karst.
(nec. Stereum odoratum Fr.),” is apparently not available for the fungus
here described; it is probable that Karsten, like Burt, confused P. crassa
with the present species; Bresadola’s description applies rather to the
former (see under Peniophora verticillata Burt).
PENtopHORA Pecku Burt = Corticium punctulatum Cooke
Corticium punctulatum Cooke, Grevillea 6: 132. 1878 (!!); Burt, Missouri
Bot. Gard. Ann. 13: 179. 1926.
Hypochnus sordidus Schroet. in Cohn, Krypt.-Fl. Schles. 3(1): 418. 1888.
Peniophora Eichleri Bres. apud Sacc. & Syd., Syll. Fung. 16: 194. 1902 (!!).
Kneiffia Eichleri (Bres.) Bres., Ann. Mycol. 1: 101. 1903.
Hypochnus cremicolor Bres., Ann. Mycol. 1: 109. 1903.
*Fructificatio crassa, subpulvinata, margine distincto, hirsuto, hymenio alutaceo,
sub lente subtiliter piloso, continuo vel rimoso; hyphae tenuiter tunicatae, nodoso-
septatae, 2-4 y, diam., demum crasse tunicatae, compactae, paralleles; cystidia clavato-
cylindracea, (60—) 90-230 x 7.5-12 uw, vel 30-75 yu, emergentia, vel immersa, canalicula
ad basim angusta, apicem versus gradatim expansa, tenuiter uni- vel pauciseptata;
basidia clavato-cylindracea, 40-85 x 7.5-9.5 u, inferne crasse tunicata, sterigmata 4
gerentia 7-9.5 x 2-3 u.; sporae sublunatae, utrinque obtusae, apiculo laterali obtusoque
praeditae, 15-18 x 4.5-7 ys. — Suecia: Lappland, VIII.17.10, L. Romell 4/3, typus;
Bedaré, XII.25.10, L. Romell 412; Stockholm, VI.20.09, L. Romell 369 (ut Chaeto-
cortic, fusisporum) ; specim, omnia in herb. Burt in Farlow Herb.
Rocers & JACKSON: SYNONYMY OF THELEPHORACEAE 321
Hypochnus albo-stramineus Bres., Ann. Mycol. 1: 110. 1903 (!!).
Peniophora sordida (Schroet.) Hohn. & Litsch., Ann. Mycol. 4: 290. 1906 (Jun.) ;
K. Akad. Wiss. Wien Math.-Nat. Kl]. Sitzungsb. 117, I: 1097. 1908; nec Penio-
phora sordida (Karst.) Héhn. & Litsch., K. Akad. Wiss. Wien Math.-Nat. KI.
Sitzungsb. 115, I: 1559. 1906 (Oct.) ; Burt, Missouri Bot. Gard. Ann. 12: 280.
[1926]; Bourd. & Galz., Hymén. de France p. 305. [1928]; (= Corticium
sordidum Karst., Soc. Faun. Fl. Fenn. Med. 9: 65. 1882.).
Peniophora sordidella Hohn. & Litsch., K. Akad. Wiss. Wien Math.-Nat. KI.
Sitzungsb, 115, I: 1559. 1906; Bourd. & Galz., Hymén. de France p. 262. [1928].
Peniophora sphaerospora Hohn. & Litsch., K. Akad. Wiss. Wien Math.-Nat. KI.
Sitzungsb. 115, I: 1600. 1906 (!!).
Gloeocystidium Eichleri (Bres.) Hohn. & Litsch., Osterr. Bot. Zeitschr. 58: 331.
1908.
Tomentella albo-straminea (Bres.) Hohn. & Litsch., Osterr. Bot. Zeitschr. 58:
476. 1908.
Gloeocystidium albo-stramineum (Bres.) Hohn. & Litsch., K. Akad. Wiss. Wien
Math.-Nat. KI. Sitzungsb. 117, I: 1097. 1908; Bourd. & Galz., Hymén. de
France p. 262. [1928]; nec G. albo-stramineum (Bres.) Overh., Mycologia 30:
276. 1938 (= Peniophora albo-straminea Bres., Mycologia 17: 69. 1925.).
Corticium albo-stramineum (Bres.) Wakef., Br. Mycol. Soc. Trans. 4: 118. 1913;
nec C. albo-stramineum (Bres.) Overh., Mycologia 30: 275. 1938 (= P. albo-
straminea Bres.).
Gloeocystidium cremicolor (Bres.) Bres. apud Bourd. & Galz., Soc. Mycol. Fr. Bul.
28: 368. [1913].
Gloeocystidium albo-stramineum var. causseanum Bourd. & Galz., Soc. Mycol. Fr.
Bul. 28: 368. [1913]; Hymén. de France p. 262. [1928] (!!).
Peniophora Peckii Burt, Missouri Bot. Gard. Ann. 12: 291. [1926] (!!).
Gloeocystidium albo-stramineum subsp. Eichleri (Bres.) Bourd. & Galz., Hymén.
de France p. 263. [1928].
Gloeocystidium albo-stramineum subsp. sphaerosporum (Hohn. & Litsch.) Bourd.
& Galz., Hymén. de France p. 262. [1928].
Gloeocystidium albo-stramineum subsp. cremicolor (Bres.) Bourd. & Galz., Hymén.
de France p. 263. [1928].
Corticium punctulatum is readily recognizable by the rough- and thick-
walled ellipsoid to subglobose spores, the colorless, obtuse, subcylindric
gloeocystidia, and the loose mycelium, thick-walled near the substratum,
thinner above, with prominent proliferating clamps throughout. There
are certainly other species, closely related to this, with either the same
structure or spores of similar description; but strangely enough they are
not included among the type and authentic specimens indicated (by
exclamation points) above. As in many gloeocystidiate fungi, the degree
of exsertion of the gloeocystidia is determined largely by the maturity of
the fungus, and varies within a single specimen; relative abundance and
preservation of mycelium depends on the same factor. Most of the type
of C. punctulatum has bo:h mycelium and gloeocystidia partly disinte-
grated, but they are present, and characteristically shown about the
margins.
It will be observed that authentic material of H. sordidus and H. cremi-
color was not at hand for examination. Material of P. sordidella in the
von Héhnel herbarium and of G. cremicolor from Bourdot’s herbarium,
a22 Fartowia, Vou. 1, 1943
and the original decriptions, were used to determine the nature of those
species.
PENIOPHORA PERTENUIS (Karst.) Burt =P. tenuis (Pat.) Massee
Corticium tenue Pat., Rev. Mycol. 7: 152. 1885; Tab. Anal. Fung. 1: 203. fig.
462. 1886.
Peniophora praetermissa Karst., Bidr. Kanned. Finl. Nat. Folk 48: 423. 1889 (!!).
Peniophora tenuis (Pat.) Massee, Linn. Soc. Bot. Jour. 25: 149. 1889; Burt,
Missouri Bot. Gard. Ann. 12: 317. [1926].
Corticium pertenue Karst., Hedwigia 29: 270. 1890 (!).
Corticium praetermissum (Karst.) Bres., Ann. Mycol. 1: 100. 1903.
Kneiffia tenuis (Pat.) Bres., Ann. Mycol. 1: 105. 1903; nec K. tenuis Pat., Soc.
Mycol. Fr. Bul. 9: 132. 1893.
Gloeocystidium praetermissum (Karst.) Hohn, & Litsch., K. Akad. Wiss. Wien
Math.-Nat. KI. Sitzungsb. 115, I: 1556, 1565. 1906.
Gloeocystidium tenue (Pat.) Hohn. & Litsch., Wiesner-Festschr. p. 70. 1908;
Bourd. & Galz., Hymén. de France p. 256, [1928].
Gloeocystidium tenue subsp. praetermissum (Karst.) Bourd, & Galz., Soc. Mycol.
Fr. Bul. 28: 364. [1913]; Hymén. de France p. 256. [1928].
Peniophora pertenuis (Karst.) Burt, Missouri Bot. Gard. Ann. 12: 315. [1926].
Gloeocystidium caliciferum Litsch., Osterr. Bot. Zeitschr. 77: 126. fig. 4. 1928
(!!); 88: 115. 1939; Bourdot, Soc. Mycol. Fr. Bul, 48: 213. 1932.
The P. tenuis complex is an abundant, variable, and difficult group.
There is no material of Patouillard’s species in his herbarium, and no
record of anyone’s having seen authentic material except himself. How-
ever, the descriptions and figure cited seem to define it quite precisely.
He says “cystidia cylindric,” and shows them approximately so; there-
fore in preliminary sorting, specimens belonging to the tenuis-complex
were assigned to P. tenuis sensu stricto if the cystidia were long-emergent,
of nearly uniform diameter, and scarcely capitate. A specimen of P.
pertenuis whose data agree with those originally published, and which in
consequence is taken to be the type, has its cystidia emergent, ventricose,
quite slender above, and with a few mineral granules on the button-like
apex. In preliminary study of this complex considerable numbers of
specimens with similar cystidia were tentatively assigned to Karsten’s
species. There is much intergradation in various details between these
two extreme phases, and only extremes are at all readily separable.
Three authentic specimens of P. praetermissa were at hand; the best of
these showed both types of cystidia in the one specimen. There are no
other useful grounds for segregation; the species are therefore combined.
Litschauer published under the name G. caliciferum a clear and careful
account of a fungus characterized by unique caliciform gloeocystidioid
bodies (for which see his illustration) in addition to the structures
earlier ascribed to P. tenuis. Material exactly like his type has been
examined from several localities in North America. However, caliciform
bodies like those described by Litschauer are connected by numerous
intergrading structures with simple globoid gloeocystidia; and both simple
and highly differentiated structures occur in all the forms here included
under P. tenuis. Furthermore, on two occasions Professor Litschauer
Rocers & JAcKSON: SYNONYMY OF THELEPHORACEAE 393
wrote that he had come to regard the whole complex as a single species,
and that Bourdot was of the same mind. Although Bourdot previously
(Soc. Myc., 1. c.) and Litschauer subsequently (1939) recognized the
" autonomy of G. caliciferum, our ample material indicates the correctness
of the opinion here quoted from Litschauer,
PENIOPHORA PHYLLOPHILA Massee — Vararia phyllophila (Massee) comb. nov.
? Athelia epiphylla Pers., Mycol. Eur. 1: 84, 1822.
? Thelephora epiphylla (ers.) Fries, Elenchus Fung. 1: 226. 1828.
? Hypochnus epiphyllus (Pers.) Wallr., Fl. Krypt. Germ, 2: 309. 1833.
Corticium epiphyllum sensu Rav., Fung. Amer. Exs. 457. 1881; an A. epiphylla
Pers. ?
Peniophora phyllophila Massee, Linn. Soc. Bot. Jour, 25: 150. 1889 (!!); Burt,
Missouri Bot. Gard. Ann. 12: 241. [1926].
Asterostromella epiphylla (Pers.?) Héhn. & Litsch., K, Akad. Wiss. Wien Math.-
Nat. KI. Sitzungsb. 115, I: 773. fig. 3. 1907,
Vararia phyllophila (Massee) comb. nov.
Vararia phyllophila is readily separable by the extremely delicate,
flagellate outer branches of the dichophyses, somewhat as in the left-
hand member of von Hohnel & Litschauer’s fig. 3d, but more decidedly
curved toward the tips. No gloeocystidia, basidia, or spores were found
in the Farlow Herbarium copy of Ravenel, but a portion in von Héhnel’s
herbarium of the specimen in the Berlin copy shows them all —the
basidia not in the least as figured, the spores quite like the right-hand
member of fig. 3b, lunaie-acicular, 17-21 x 2.5-3.5 p, the gloeocystidia
with yellowish granular content. From the description, the Ravenel
fungus might well be Persoon’s, and since C. epiphyllum seems not to
have been used for any other species from the time of its publication,
Massee’s substitution of a new name seems to have been a bit high-handed.
There is no definite evidence, however, that P. phyllophila occurs in
Europe; on the probabi.ity that that will continue to be the case, the
later epithet is reluctantly accepted.
a
PeniopHora PIceEINA Overh. := Corticium Pini-canadensis (Schw.) comb. nov.
Radulum Pini-canadensis Schw., Am. Phil. Soc. Trans. n.s. 4: 164, [1832] (!!).
Peniophora Piceina Overt., Mycologia 22: 238. 1930 (!!).
Corticium Pini-canadensis (Schw.) comb. nov.
This very characteristic species, well described by Overholts (1. c.),
is so common that it is surprising that it has appeared in the literature
so infrequently. Specimens are at hand from Ontario (20 collections)
and Quebec, Maine, New Hampshire, Massachusetts, Connecticut, New
York, Tennessee, and Peansylvania. The New York record is based on
a collection (Cornell Univ. 8072) listed by Burt as a paratype of Penio-
phora albula Atk. & Burt.
The structures described by Overholts as cystidia seem best interpreted
as gloeocystidia, hence the transfer to Corticium rather than to Penio-
phora. The portion of the type of Radulum Pini-canadensis in the Curtis
herbarium is in an excellent state of preservation. The species occurs
324, FarLowiA, VoL. 1, 1943
not only on conifers, where it is most frequently found on small branches
on the ground, but also on the bark of a variety of deciduous hosts.
PENIOPHORA SHEARI Burt = P. aurantiaea (Bres.) Hoéhn. & Litsch.
See under Aleurodiscus Zelleri Burt.
PENIOPHORA sorDIDA (Karst.) Héhn. & Litsch. = P. cremea (Bres.) Sacc. & Syd.
See under Peniophora arachnoidea Burt.
PENIOPHORA STRATOSA Burt = P. Roumeguerii (Bres.) Hohn. & Litsch.
Corticium Roumeguerii Bres., Fungi Trident. 2: 36. 1892 (!!).
Peniophora Molleriana (Bres. in litt.) Sacc., Soc. Broteriana Bol. 11: 13. 1893 (!).
Kneiffia Roumeguerii (Bres.) Bres.. Ann. Mycol. 1: 103. 1903.
Peniophora Roumeguerii (Bres.) Héhn. & Litsch., K. Akad. Wiss. Wien Math.-
Nat. KI. Sitzungsb. 115, I: 1581. 1906; 116, I: 789. 1907; Burt, Missouri Bot.
Gard. Ann. 12: 270. [19261]; Bourd. & Galz., Hymén. de France p. 316. [1928].
Peniophora stratesa Burt, Missouri Bot. Gard. Ann. 12: 333. [1926] ('!)s; nee
P, stratosa Petch, Roy. Bot. Gard. Peradeniya Ann. 9: 293. 1925.
The type specimen of P. stratosa is only an especially thick fructifica-
tion of a species which typically is quite thick; a slide in von Héhnel’s
collection of authentic material of P. Molleriana is equally stratose.
The original description of Kneiffia subascondita Bres. (Ann. Mycol. 1:
102. 1903.) is a good picture of the present species, and strongly sug-
gests that, in reducing P. swbascondita to subspecific rank under P.
Roumeguerii, Bourdot & Galzin (Hymén. de France p. 317. [1928].)
erred only in retaining it in any rank as distinct. However, a paratype
of P. subascondita in Burt’s herbarium differs in having much larger
spores and clamp-bearing mycelium, and in other respects. For want
of a specimen agreeing with the description, P. subascondita is not here
listed as a synonym.
PENIOPHORA SUBAPICULATA (Bres.) Burt = Corticium subapiculatum Bres,
Corticium subapiculatum Bres., Mycologia 17: 69. 1925 (!!).
Peniophora subapiculata (Bres.) Burt, Missouri Bot. Gard. Ann. 12: 280, [1926].
Burt transferred C, subapiculatum Bres. to Peniophora. An examina-
tion of the type fails to reveal any cystidia. All of the six additional
collections assigned to this species by Burt have been examined. Three
of them prove to be Peniophora Greschikii (Bres.) Bourd. & Galz. (see
under P. alba Burt), and two are Peniophora pallidula Bres., neither of
which resembles C. subapiculatum at all closely. Only one (Weir 414)
from Evaro, Montana, is like the type. It is evident that Burt’s descrip-
tion was not drawn from the type specimen and it is desirable, therefore,
to restore Bresadola’s species in Corticium. :
PENIOPHORA SuBCREMA Hohn. & Litsch. = P. Greschikii (Bres.) Bourd. & Galz.
See discussion under Peniophora alba Burt.
_ PENIOPHORA SUBSULPHUREA (Karst.) Héhn. & Litsch.
See discussion under Corticium subincarnatum Peck.
Rocers & JACKSON: SYNONYMY OF THELEPHORACEAE 325
PENIOPHORA TABACINA Burt:= P. dryina (Berk. & Curt.)
See under Coniophora drvina (Berk. & Curt.) Massee.
PENIOPHORA TENUISSIMA Pec =P. pubera (Fries) Sacc.
Thelephora pubera Fries, Elenchus Fung. 1: 215. 1828.
Hyphoderma puberum (F-ies) Wallr., Fl. Krypt. Germ. 2: 576. 1833.
Corticium puberum (Fries) Fries, Epicr. Syst. Mycol. p. 562. 1838.
Hymenochaete pubera (Fries) Lév., Ann. Sci. Nat. Bot. III 5: 152. 1846.
Hypochnus puberus (Fries) Bon., Handb. p. 159. 1851.
Peniophora pubera (Fries) Sacc., Syll. Fung. 6: 646. 1888; Burt, Missouri Bot.
Gard. Ann. 12: 313. [1926]; Bourd. & Galz., Hymén. de France p. 316. [1928].
Hypochnus subtilis Schroct. in Cohn, Krypt.-Fl. Schles. 3(1): 418. 1888 (!!).
Peniophorella pubera (Fries ?) Karst., Bidr. Kanned. Finl. Nat. Folk 48: 427.
1889. ;
Kneiffia pubera (Fries) Bres., Ann. Mycol. 1: 102. 1903.
Peniophora subtilis (Schoet.) Hohn. & Litsch., Ann, Mycol. 4: 290. 1906; K.
Akad. Wiss. Wien Ma:h.-Nat. KI]. Sitzungsb. 116, I: 837. 1907.
Peniophora tenuissima Peck, New York State Mus. Bul. 157: 114. 1912 (!!).
Peck’s species was not included nor accounted for by Burt in his
treatment of the genus. The type is quite typical of the less ceraceous
collections of P. pubera.
Except for the type, all specimens cited as P. subtilis by Hohnel &
Litschauer, and all in Héhnel’s herbarium under that name, are P.
argillacea Bres. The type, however, is a thin but perfectly characteristic
specimen of P, pubera. Presumably one of the misdetermined collec-
tions is the basis for the inclusion, by Bourdot & Galzin (1. c., p. 279) of
P. subtilis as a synonym of P. argillacea.
PENIOPHORA THUJAE Burt =P. Sambuei (Pers.) Burt
Thelephora sera Pers., Mvcol. Eur. 1: 151. 1822.
Thelephora Sambuci Pers, Mycol. Eur, 1: 152. 1822.
Hypochnus serus (Pers.) Fries ex Wallr., Fl. Crypt. Germ. 2: 310. 1833 (falso ut
“sereus”) ; Karst., Bidr. Kanned. Finl. Nat. Folk 25: 320. 1876.
Corticium Sambuci (Pers.) Fries, Epicr. Syst. Mycol. p. 565. 1838; Donk, Nederl.
Mycol. Ver. Med. 18-20: 145. 1931.
Hypochnus Sambuci (Pers.) Bon., Handb. p. 159. 1851.
Corticium serum (Pers.) Fries, Hymen. Eur. p. 659, 1874; Bres., I. R. Accad.
Agiati Atti III 3: 112. 1897; R. E. Fries, Soc. Goth. Acta IV 3: 37. 1900;
Bourd, & Galz., Hymén. de France p. 203. [1928].
Lyomyces serus (Pers.) Karst., Rev. Mycol. 3(9) : 23. 1881 (ut “Lyomices”).
Lyomyces Sambuci (Pers.) Karst., Bidr. Kanned. Finl. Nat. Folk 37: 153. 1882.
Kneiffia sera (Pers.) Karst., Hedwigia 28: 195. 1889.
Corticium cretaceum [Fri2s] Cooke ex Sacc., Syll. Fung. 11: 128. 1895.
Peniophora Sambuci ()ers.) Burt, Missouri Bot. Gard. Ann, 12: 233. [1926].
Peniophora Thujae Burt, Missouri Bot. Gard. Ann, 12: 236. [1926] (!!).
There are other synonyms, pre-Friesian, with which it seemed un-
necessary to burden the list. While it is probable that what Fries and
Karsten described under C. serum and H. serus was Corticium bomby-
cinum (Sommerf.) Karsi:., such a misinterpretation of Persoon’s species
would not alter the synonymy (cf. R. E. Fries, Bresadola). Bresadola
in his discussion places an exclamation point after 7. Sambuci, thereby
326 Fartowia, Vou. 1, 1943
presumably indicating that he had seen authentic material. It is the
consensus of opinion that both specific names are equally available; but
since 7’, sera has been repeatedly confused with other fungi (cf. Fries’s
synonymy, and Rogers, Mycologia 31: 300. 1939.), “Sambuci” seems
the preferable specific name. As to the preferable genus, there seems
little choice between Bourdot’s and Burt’s treatment. The fungus is
extremely common in eastern North America on a variety of coniferous
and broad-leaved hosts. P. Thujae has no morphological characters by
which to distinguish it; and specimens exactly like the type occur on
wood of broad-leaved as well as coniferous species.
PENIOPHORA TRACHYTRICHA Ellis & Evy. =P. aspera (Pers.) Sacc.
See under Corticium Berkeleyi Cooke.
PENIOPHORA VERSATA Burt = Corticium versatum (Burt) comb. nov.
Peniophora versata Burt, Missouri Bot. Gard. Ann. 12: 305. [1926] (!!).
Corticium versatum (Burt) comb. nov.
This species is a form in which the gloeocystidia may in part project
prominently, There seems only one structure involved and the species
is best included among the gloeocystidiate forms in Corticium where it
would naturally be sought.
PENIOPHORA VERTICILLATA Burt =P. erassa Burt
Stereum Karstenii Bres., I, R. Accad, Agiati Atti III 3: 108. 1897; Bourd. & Galz.,
Soc, Mycol. Fr. Bul. 37: 126. 1921; Hymén. de France p. 385. [1928]; nec
Peniophora Karstenii Massee, Linn. Soc. Bot. Jour. 25: 153. 1889 (= Phlebia
coccineo-fulva Schw., Am. Phil. Soc. Trans. n.s. 4: 165. [1832]; Peniophora
coccineo-fulva (Schw.) Burt, Missouri Bot. Gard. Ann. 12: 253. [1926]);
neque Xerocarpus odoratus sensu Karst., Rev. Mycol. 3(9) : 22. 1881 (= Penio-
phora luna Romell in sched. sp. nov. in hac re).
Peniophora crassa Burt ex Peck, New York State Mus. Rept. 54: 155, 1901;
Missouri Bot. Gard. Ann. 12: 286. [1926].
Lloydella Karstenii (Bres.) Hohn. & Litsch.. K. Akad. Wiss. Wien, Math.-Nat. K1.,
Sitzungsb. 115, I: 1568. 1906.
Peniophora verticillata Burt, Missouri Bot. Gard. Ann. 12: 285. [1926] (!!).
Stereum Karstenii fa. incrustata Pilat, Soc. Mycol. Fr. Bul. 49: 44. 1933.
P. verticillata was described from a single collection made in Oregon
by S. M. Zeller. A comparison of this with a series of collections of
P. crassa leaves no doubt as to their identity. The thick layer of sub-
stratal hyphae stressed by Burt, in his description of the former, is an
unusual development due to the soft irregular surface of the substratum.
Such a layer is often present in P. crassa when it occurs on bark. Burt
states that no spores could be found. There is an abundance of the small
allantoid spores characteristic of P. crassa in the type specimen. The
character and arrangement of the incrusting granules on the projecting
cystidia, stressed by Burt in his description of P. verticillata, and from
which the name is derived, seem unessential. The cystidia in P. crassa
may or may not be incrusted.
Rocers & JACKSON: SYNONYMY OF THELEPHORACEAE B27
Bresadola’s name was published to replace Xerocarpus odoratus sensu
Karst. (nec Stereum odoratum Fries). Since, however, Karsten probably
had two fungi confused in his X. odoratus (cf. under Peniophora odorata
sensu Burt), the type of S. Karstenii must be the material Bresadola had
in hand when describing the species. The line between Stereum and
Peniophora is not always clear; this fungus appears to the present
authors —as to Burt—to be a Peniophora.
SEBACINA
SEBACINA DEGLUBENS (Berk. and Curt.) Burt =S. inerustans (Pers. ex Fries) Tul.
Thelephora incrustans Pers. ex Fries, Syst. Mycol. 1: 448, 1821,
Sebacina incrustans (Pers. ex Fries) Tul., Linn. Soc. Bot. Jour. 13: 36. [1871];
Burt, Missouri Bot. Gard. Ann. 2: 752. 1915; McGuire, Lloydia 4: 12. 1941,
ubi synon.
Corticium deglubens Beik. & Curt., Grevillea 1: 166, 1873 (1!).
Sebacina deglubens (Berk. & Curt.) Burt, Missouri Bot. Gard. Ann. 2: 755. 1915.
The type of C. deglubens is merely a rather thin fructification of S.
incrustans growing over dead wood. Burt seems not to have been too
happy in maintaining it as distinct.
SEBACINA FIBRILLOSA Burt =- Ceratobasidium fibrillosum (Burt) comb. nov.
Sebacina fibrillosa Burt, Missouri Bot. Gard. Ann, 13: 335. 1926 (!!).
Ceratobasidium fibrillosum (Burt) comb. nov.
Superficially appearing like a water-soaked sterile mycelium. Basidia
stalked-obovate, quite like those of a Sebacina except that they lack septa.
The (correct) impression that the fungus is related to the Tremellaceae
is increased by the regular germination of the.spores by repetition. In
many respects this species is quite distinct from the published species of
Ceratobasidium (cf. Rogers, Univ. Iowa Studies Nat. Hist. 17: 4. 1935.) ;
most obviously, in the possession of subcylindric cystidia.
SEBACINA POLYSCHISTA (Bers. & Curt. in herb.) Burt ? = Corticium cremoricolor
Berk, & Curt.
Corticium cremoricolor Berk. & Curt., Grevillea 1: 180. 1873 (!!); Burt,
Missouri Bot. Gard. Ann. 13: 218. 1926.
Sebacina polyschista (B2rk. & Curt. in herb. sub Corticio) Burt, Missouri Bot.
Gard. Ann. 13: 338. 1926 (!!).
The type and only specimen is in very poor condition, and its determina-
tion must remain doubtful. Assuredly, however, it is not a Sebacina;
and it appears to be a .arge-spored form of C. cremoricolor. It should
never have been disinterred!
SEBACINA SCARIOSA (Berk. & Curt.) Burt ? Nomen confusum.
Corticium scariosum Bek. & Curt., Grevillea 2: 3. 1873 (Jul.) Qi!); nec C,
scariosum Berk. & Br., Linn. Soc. Bot. Jour. 14: 71, 1873 (Dec.).
Corticium secedens Sacc., Syll. Fung. 6: 635. 1888.
Sebacina scariosa (Berk. & Curt.) Burt, Missouri Bot. Gard. Ann. 2: 762. 1915.
328 FarLow1a, Vo. 1, 1943
The type and only collection is superficially much like Peniophora
gigantea, and is so labeled in the Curtis herbarium. The surface, above
the horny layer, is covered with a loose mealy crust of irregularly ar-
ranged large globose cells, sometimes wrinkled, but not cruciate-septate,
apparently joined as are the conidia of Cystopus. Not certainly one
fungus; probably an old fructification of P. gigantea or a similar form
overgrown by a mold.
SISTOTREMA
SISTOTREMA sensu Denk and Rogers p. p. = Trechispora Karst.
The genus Sistotrema was earlier enlarged (cf. Rogers, Univ. Iowa
Studies Nat. Hist. 17: 19-23. 1935.) to include all species having the
Urnigera type of basidia, whatever the form of the basidiocarp. It has
seemed advantageous to separate the resupinate from the reflexed or stipi-
tate forms. The genus 7'rechispora Karst. is available for the former group
of species. Changes found necessary in the names used in that earlier paper
are here listed for the sake of completeness. Cross reference to other names
used in American literature which properly belong in Trechispora will
be found in the proper alphabetical position in the preceding pages.
More complete synonymy will be presented in a paper now in course
of preparation.
SISTOTREMA CORONILLA (Hoéhn.) Donk = Trechispora Brinkmanni (Bres.)
See under Corticium coronilla Hohn.
SISTOTREMA SUBTRIGONOSPERMUM Rogers = Trechispora subtrigonosperma (Rog-
ers) comb. nov,
TULASNELLA
TULASNELLA CALOSPORA (Boud.) Juel = Gloeotulasnella calospora (Boud.) Rogers
Tulasnella calospora (Boud.) Juel, Bih. Svensk, Vet.-Akad. Handl., Afd. 3, 23(12) :
23. 1897 (!!); Burt, Missouri Bot. Gard. Ann. 13: 328. 1926; Bourd. & Gallz.,
Hymén, de France p. 57. [1928].
Gloeotulasnella calospora (Bourd.) Rogers, Ann. Mycol. 31: 201. 1933, ubi
synon.
This species was placed in Gloeotulasnella because of consistency and
basidial structure.
TULASNELLA EICHLERIANA Bres. = T. violea (Quél.) Bourd. & Galz.
Hypochnus violeus Quél., Assoc. Frang. Avanc. Sci. 1882: 401. 1883 (!).
Tulasnella violea (Quél.) Bourd. & Galz., Soc. Mycol. Fr. Bul. 25: 31. 1909;
Hymén. de France 56. [1928]; Burt, Missouri Bot. Gard. Ann. 6: 256. 1919;
Overh., Torrey Bot. Club Bul. 49: 166. 1922 (falso ut “Violae”); Rogers,
Ann. Mycol. 31: 184. 1933, ubi synon.
Tulasnella Eichleriana Bres., Ann, Mycol. 1: 113. 1903 (!); Burt, Missouri Bot.
Gard. Ann. 6: 255. 1919,
The identity of the two species included by Burt has already been noted
by Rogers.
DEPARTMENT OF BIOLOGY DEPARTMENT OF BOTANY
AMERICAN INTERNATIONAL COLLEGE UNIVERSITY OF TORONTO
SPRINGFIELD, MASSACHUSETTS ToRONTO, CANADA
Rocers & JACKSON: SYNONYMY OF THELEPHORACEAE . 329
INDEX
Names accepted, or concerning whose status no judgment is expressed, are printed
in roman; synonyms, or names rejected for other reasons, are printed in italic; new
names are printed in bold-face. The number of the page of special treatment is
printed in bold-face.
Acia
ferruginea, 272
tomentosa, 271
Aciella
tomentosa, 271
Aleurodiscus, 265, 267, 269, 307
albo-roseus, 268
albus, 306
amorphus, 269
amylaceus, 291
apiculatus, 267
aurantius, 307
botryosus, 299
Bertii, 268, 307
Burti, 268
candidus, 268
crassus, 268
cremeus, 268
croceus, 304
diffissus, 270
disciformis, 279 ;
Grantii, 269
helveolus, 269
Javanicus, 267
lepra, 268
macrosporus, 307
Micheneri, 303
mirabilis, 267
Oakesii, 268
pallide-roseus, 268
paraphysatus, 297
peradeniyae, 267
Peteloti, 268
polygonioides, 300
roseus, 299
sajanensis, 270
salmoneus, 268
sinensis, 268
spinulosus, 267
subcruentatus, 307
subgiganteus, 303
succineus, 270
usambarensis, 267
Zelleri, 270
Alytosporium
croceum, 304
Amaurodon
viridis, 277
Amphinema
sordescens, 276
Artocreas
Micheneri, 303
Asterodon, 271
ferruginosus, 271
Asterostroma, 265, 271
andinum, 271
bicolor, 271
gracile, 271
ochrostroma, 271
spiniferum, 271
Asterostromella, 264, 293
dura, 310
effuscata, 390
epiphylla, 323
granulosa, 283
investiens, 292
rhodospora, 294
Athelia
citrina, 304
epiphylla, 323
sericea, 308
strigosa
8 muscigena, 275
Auricularia
aurantiaca, 319
Botryobasidium, 264, 272
coronatum, 272
flavescens, 272
isabellinum, 272
ochraceum, 272
Solani, 272
subcoronatum, 272
vagum, 272
Bourdotia
deminuta, 294
Caldesiella
vaga, 309
viridis, 277
Ceratobasidium, 264, 272
atratum, 272
fibrillosum, 327
plumbeum, 272
Cerocorticium
albissimum, 314
Chromosporium
pactolinum, 297
viride, 292
330 FarRtow!A, Vou. 1, 1943
Clavaria alboflavescens, 277
byssacea, 278 albo-ochraceum, 279, 290
contorta, 270 subsp. amianthinum, 289
fistulosa, 270 albo-stramineum, 311, 321
Himantia, 278 albulum, 312
Coniophora, 265, 273, 276, 278 Allescheri, 313
alboflavescens, 277, 280 alutarium, 292
arida, 280 amianthinum, 289
subsp. fumosa, 267, 274 amorphum, 269
atrocinerea, 266, 273 amylaceum, 291
avellanea, 275 apiculatum, 281
Betulae, 280 arachnoideum, 286
brunneola, 273 areolatum, 281, 282
byssoidea, 275 argillaceum, 296
centrifuga, 286 aurantiacum, 270
cerebella, 276 Atkinsonii, 282
corrugis, 276 atratum, 272
corticola, 277 Berkeleyi, 282
crocea, 277 bicolor, 304
cyanospora, 277 bisporum, 286
dryina, 277 bombycinum, 283, 295, 325
Ellisii, 273 botryoideum, 284
flava, 278 brunneolum, 273
flavomarginata, 278 byssinum, 304, 305
fulvo-olivacea, 274 var. microsporum, 304
fumosa, 273, 308 byssoideum, 275
furva, 273 caeruleum, 273
fusca, 274 calceum, 284, 285
Harperi, 277 canum, 285
insinuans, 310 calothrix, 316
Karstenii, 274 calotrichum, 318
laeticolor, 277 centrifugum, 286
leucothrix, 273 subsp. bisporum, 287
mustialaensis, 277 subsp. fugax, 287
olivacea, 266, 273, 280 var. macrospora, 287
Poly poroidea, 277, 279 var. soredioides, 286
puteana, 276 var. tenuis, 287
sibirica, 274 Chusqueae, 282
Sistotremoides, 274, 280 citrinum, 304
sordulenta, 309 confine, 288
suffocata, 280 confluens, 275, 276, 296, 306
sulphurea, 308 consimile, 286
umbrina, 273, 275 contiguum, 288, 302
vaga, 280 coronilla, 286, 288, 292
Coniophorella, 274 corruge, 276
atrocinerea, 273 cremeum, 314
byssoidea, 276 cremoricolor, 327
laeticolor, 278 cretaceum, 284, 325
olivacea, 274 croceum, 304
subsp. fulvo-olivacea, 274 crustaceum, 288
umbrina, 274 crustulinum, 289
var. olivacea, 274 decipiens, 286
Corticium, 264, 265, 279, 280 deglubens, 327
abeuns, 280 dryinum, 277
albo-cremeum, 295 effuscatum, 290
Rocers & JACKSON: SYNONYMY OF THELEPHORACEAE 331
effusum, 290
Eichlerianum, 314
Ellisti, 273.
epiphyllum, 323
ermineum, 291, 312
evolvens, 318
fenestratum, 291
fibrillosum, 307
flavissimum, 304
fumosum, 273, 308
furfuraceum, 285, 286
fuscostratum, 291
Galzini, 285
geogenium, 291, 311
glebulosum, 284
Greschikii, 310
illaqueatum, 281, 292
fa, rhizophorum, 281
incanum, 292
inaequale, 311
incrustans, 285, 299
investiens, 292, 299
involucrum, 293
jamaicense, 294
javanicum, 267
Koleroga, 294
lacteolum, 286
lacteum, 294
lacunosum, 275
laeve, 318
laeticolor, 277
laetum sensu Burt, 296
latitans, 282
leucothrix, 273
Litschaueri, 281, 300
livido-caeruleum, 281
Macounii, 305
maculatum, 305
microsporum, 286
miniatum, 319
Muraii, 290
mustialaense, 277
mutatum, 313
myxosporum, 282
ochraceum, 289
ochrofarctum, 296
ochroleucum, 281, 291
octosporum, 287
oleosum, 313
olivaceum, 273
Overholtsti, 296
pactolinum, 297
pallescens, 310
pallidum, 296
paraphysatum, 297, 307
pectinatum, 298
peradeniyae, 267
pertenue, 322
Pezizoideum, 286
pilosum, 298
Pini-canadensis, 323
Polyporoideum, 279
porosum, 281, 289, 300
praetermissum, 322
Pruni, 301
puberum, 325
punctulatum, 281, 295, 320
puteanum, 276
racemosum, 298, 299
radiosum, 294, 295
rhizophorum, 281
roseopallens, 299
roseum, 299
Roumeguerii, 324
rude, 310
rubellum, 275
Sambuci, 325
sanguineum, 319
scariosum, 327
secedens, 327
septentrionale, 300
serum, 325
setigerum, 282
sociatum, 291
sordidum, 314, 321
sordulentum, 309
sphaerosporum, 286, 292
spinulosum, 268
Stevensii, 300
stramineum, 300
subalbum, 298
subapiculatum, 324
subcinereum, 289
subcoronatum, 302
subgiganteum, 303
subincarnatum, 303
subpallidulum, 285
subsulphureum, 303
suecicum, 285
suffocatum, 280
sulphurellum, 277
sulphureum, 304, 305, 308
sulphureum sensu Burt, 304
tenue, 322
tessulatum, 281
Thelephoroides, 309
- Torrendii, 312
trigonospermum, 307
_Tsugae, 296
tuberculatum, 295
332 Fartowia, VoL. 1, 1943
Tulasnelloideum, 292
umobrinum, 275
usambarense, 268
vagum, 305, 308
var. Solani, 305
vellereum, 295
versatum, 326
vesiculosum, 301
vinososcabens, 305
Cyphella
amor pha, 269
Dichostereum
durum, 310
Diplonema
sordescens, 275
Fusisporium
Kuhnii, 287
Gloeocystidiellum, 302
porosum, 301
Gloeocystidium, 264, 302, 306
albo-stramineum, 312, 321
var. causseanum, 321
subsp. cremicolor, 321
subsp. Eichleri, 321
subsp. sphaerosporum, 321
alutaceum, 295
argillaceum, 296
Bourdotii, 312
caliciferum, 322
clavuligerum, 302
contiguum, 289, 301, 302
coroniferum, 282
cremicolor, 321
Eichleri, 321
furfuraceum, 302
inaequale, 311
Karstenii, 289, 302, 306
luridum, 312
ochroleucum, 281
oleosum, 313
pallidulum, 313
pallidum, 296
subsp. argillaceum, 296
porosum, 301
praetermissum, 322
var. Bourdotii, 312
rude, 311
stramineum, 301
tenue, 322
subsp. inaequale, 311
subsp. praetermissum, 322
Torrendii, 312
Gloeopeniophora
Allescheri, 313
Gloeotulasnella 265, 306
calospora, 328
opalea, 306
traumatica, 306
Gonatobotrys
pallidula, 313
Grandinia
granulosa, 282
Gyrophana, 287
Himantia
sulphurea, 309
umobrina, 273
Hydnochaete
setigera, 27]
Hydnum
ferrugineum, 272
ferruginosum, 272
fusco-atrum, 279
granulosum, 282
Himantia, 278
setosum, 279
Sobolewskii, 277
subfuscum, 278
tomentosum, 272
Hymenochaete, 315
Ellisii, 273
pubera, 325
Hyphoderma
asperum, 282
puberum, 325
spiculosum, 282
sulphureum, 308
Hypochnopsis
mustialensis, 277
Hypochnus, 265, 306
albo-stramineus, 312, 321
albus, 306
bisporus, 286
byssoideus, 275
centrifugus, 286
chaetophorus, 316
confluens, 275
cremicolor, 320
epiphyllus, 323
fibrillosus, 307
filamentosus, 309
flavo-brunneus, 280
fumosus, 304, 308
isabellinus, 309
Langloisii, 309
longisporus, 315
muscorum, 275
mustialaensis, 277
obducens, 275
olivaceus, 273
Rocers & JACKSON: SYNONYMY OF THELEPHORACEAE
pallescens, 294, 309
Peniophoroides, 294
Poly poroideus, 279
puberus, 325
roseus, 300
Sambuci, 325
sericeus, 308
serus, 325
setosus, 275
sordidus, 315, 320
strigosus
8 flamentosus, 275
subtilis, 325
sulphureus, 308
Thelephoroides, 310
Tulasnelloideum, 292
umbrinus, 273, 275
vagus, 309
violeus, 328
Inonotus
hispidus, 297
Leei, 297
Kneifha
Allescheri, 313
aurantiaca, 270
cremea, 314
Eichleri, 320 P
glebulosa, 284
globifera, 317
laevis, 318
latitans, 282
longispora, 315
muscorum, 276
polonensis, 316
pubera, 325
Roumeguerti, 324
sera, 325
setigera, 282
var. trachytricha, 283
subascondita, 324
tenuis, 322
Tomentella, 276
Kneiffiella
aspera, 282
latitans, 283
setigera, 282
Langloisula, 292
spinosa, 292
Lloydella
Karstenii, 326
Lyomyces, 300
byssinus, 304
mustialensis, 277
roseus, 300
Sambuci, 326
serus, 325
sulphureus, 308
Merisma
candidum, 268
Michenera
artocreas, 303
Mucronoporus
Andersoni, 297
Mycoacia, 279
Himantia, 278
Neokneifhia
aspera, 283
Nodularia, 269
balsamicola, 269
Odontia, 277
Acerina, 283
arguta, 314
Brinkmanni, 288
fusca, 308
Himantia, 278
setigera, 283
Sistotremoides, 273
tenuis, 309
vaga, 308
vesiculosa, 283
Oxydontia, 279
Himantia, 278
Ozonium
croceum, 304, 309
Pellicularia, 264, 272
filamentosa, 272, 282, 305
flavescens, 272, 280, 291
isabellina, 272, 309
Koleroga, 294, 300
Langloisii, 309, 313
pruinata, 272, 284
subcoronata, 272, 302
vaga, 272, 305
Peniophora, 264, 265, 285, 310
admirabilis, 310
alba, 310
albo-straminea, 264, 311, 321
albula, 282, 295, 323
affinis, 318
Allescheri, 313
alutaria, 313
arachnoidea, 314
argentea, 315
argillacea, 325
aspera, 282
asperipilata, 315
attenuata, 266
aurantiaca, 270, 296, 298
Burkii, 312
333
334
FarLowia, Voi. 1, 1943
byssoides, 275, 308, 314
subsp. Tomentella, 276, 314
ealothrix, 316
canadensis, 316
candida, 295
chaetophora, 316
cinerea, 289
coccineo-fulva, 326
crassa, 320, 326
cremea, 313, 314
subsp. Allescheri, 313
var. Allescheri, 313
var. Eichleriana, 314
var. glaucescens, 314
cretacea, 284
crocea, 278
delectans, 316
diffissa, 270
dissoluta, 316
dryina, 277, 278
duplex, 296
Eichleri, 320
Eichleriana, 314
filamentosa, 293
flammea, 316
flava, 278
gigantea, 317, 328
glebulosa, 284, 317
subsp. Pirina, 316
globifera, 317
gracillima, 317
Greschikii, 310, 324
hastata, 311
heterocystidia, 317
incarnata, 276, 298
Karstenii, 326
Kaufmanii, 317
laevis, 318
var. affinis, 318
laminata, 313
latitans, 283
lepida, 270
limonia, 319
livida, 285
longispora, 315
var. clavispora, 315
var. cylindrospora, 315
var. gloeocystidiata, 315
var. mycelialis, 315
ludoviciana, 316
luna, 320, 326
lurida, 266
magnahypha, 319
media, 266
miniata, 319
Molleriana, 324
muscorum, 276
mutata, 313, 318
Odontioides, 313
odorata, 320
pallidula, 313, 324
var. regenerans, 313
Peckii, 320
pertenuis, 322
phyllophila, 307, 323
Piceina, 323
pilosa, 298
Pirina, 316
polonensis, 316
praetermissa, 322
pubera, 325
rimicola, 273:
Romellii, 284
rosea, 300
Roumeguerii, 324
rudis, 311
Sambuci, 325
sanguinea, 319
subsp. anaemacta, 319
setigera, 283
Sheari, 270
sordescens, 276
sordida, 314, 321
sordidella, 315, 321
sphaerospora, 321
stratosa, 324
subalba, 298
subalutacea, 266
subapiculata, 311, 313, 324
subascondita, 324
subcremea, 311
var. subuncinata, 311
subincarnata, 303
subsulphurea, 303
subtilis, 325
tabacina, 278
tenuis, 264, 322
tenuissima, 325
Thujae, 325
Torrendii, 312
trachytricha, 282
velutina, 315
versata, 326
verticillata, 326
Peniophorella
pubera, 325
Peziza
amorpha, 269
Phanerochaete
odorata, 320
Rocers & JACKSON: SYNONYMY OF THELEPHORACEAE doo
Phlebia, 319
coccineo-fulva, 326
vaga, 308
Phlebriella
vaga, 309
Psilopezia
mirabilis, 267
Pycnodon
asper, 283
Radulum
investiens, 292
Pini-canadensis, 323
Rhizoctonia
centrifuga, 286
Sebacina, 265, 266, 327
calcea, 285
deglubens, 327
deminuta, 293
fibrillosa, 327
incrustans, 327
polyschista, 327
scariosa, 327
Serpula, 287
Sistotrema, 264, 328
coronilla, 328
subtrigonospermum, 328
Spicaria
croceum, 304
Sporotrichum
croceum, 304
flavissimum, 304
Stereum
candidum, 268
crustaceum, 288
duriusculum, 310:
insinuans, 309
Karstenii, 320, 326
fa. incrustata, 326
lepra, 268
Muraii, 290
_ fa. tuberculosum, 290
odoratum, 320
Pini, 297
Thelephora
amorpha, 269
aspera, 282
bolaris, 319
bombycina, 295
byssoides, 275
calcea, 284
* glebulosa, 284, 317
candida, 268
candidissima, 268
centrifuga, 286
citrina, 304
confluens, 275
cretacea, 285
epiphylla, 323
evolvens, 318
fumosa, 308
granulosa, 282, 284
incrustans, 327
insinuans, 309
lactea, 295
laevis, 318
Menieri, 308
miniata, 319
Muraii, 290
odorata, 320
olivacea, 266, 273
patllescens, 309
pubera, 325
puteana, 276
rosea, 299
Sambuci, 325
sanguinea, 319
sera, 325
setigera, 282
Sistotremoides, 273, 280
subochracea, 292
sulphurea, 304, 308
umbrina, 273, 275, 318
var. lignatilis, 274
Tomentella, 264
albo-straminea, 321
byssina, 304
fibrillosa, 307
fugax, 287
Menieri, 308
obducens, 275
sulphurea, 308
trigonosperma, 307
umbrina, 274
Trechispora, 264, 328
Brinkmanni, 287, 288, 328
coronifera, 282, 288
subtrigonosperma, 328
Tremellodendron
candidum, 268
Tulasnella, 265, 328
bifrons, 310
calospora, 328
Eichleriana, 328
metallica, 328
Violae, 328
violea, 328
Vararia, 264, 293, 299
alutaria, 292
effuscata, 290, 294.
granulosa, 283
336
investiens, 292
pallescens, 309
pectinata, 298
Peniophoroides, 294
phyllophila, 323
racemosa, 299
Xanthochrous, 297
FARLOWIA,
VoL. 1, 1943
Xerocarpus
alutarius, 292
crustaceus, 288
laeticolor, 277
edoratus, 320, 326
subsulphureus, 303
Zygodesmus
pubidus, 276
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CONTENTS OF VOL. 1, NO. 3
Norms on Aucat Nomenciarure. III. By George F. Papenfuss............ 337
New Sprcius AND VARIETIES oF Wisconsin ALGAB. By G. W. Prescott...... 347
MICROAPLANOAPORNS OF VaucHERIA. By Gilbert M. Smith........ a ee 387
te Nugusermp FIELD oF Sropy wirH THn Description or a New Licuen FROM
Catrrornia. By Albert W. C. Ds TOT Oe i grog gs ce 391
Marine Foner: Tuer Taxonomy AND Brotocy. By E£. S. Barghoorn and
DEON Ro Hes se GM ce se tl SE Sak ss Re .. 895
Srupms in THe Genus ZycosaccnaRomycus I. By Walter J. Nickerson...... - 469
A New GENUs OF THE ScLEROTINIACEAR, By H. H. Wheizel............005 483
Vol. 1, No. 2 was issued July 26, 1943.
FARLOWIA
A JOURNAL OF CRYPTOGAMIC BOTANY
VoL. 1 JANUARY, 1944 No. 3
NOTES ON ALGAL NOMENCLATURE.
Ill, MISCELLANEOUS SPECIES OF CHLOROPHYCEA®"
PHAEOPHYCEAE AND RHODOPHYCEAE
GEorRGE F. PAPENFUSS
The first two papers in this series! dealt with certain questions of
nomenclature pertaining primarily to South African marine algae. The
present article is concerned mainly with extra-South African species which
came under consideration in connection with the writer’s study of the
South African marine flora.
Udotea Flabellum (Ell. et Soland.) Lamouroux, Nouv. Bull. Sci. Soc. Philom.
3: 186. 1812.
Udotea Flabellum (Ell, et Soland.) Howe, Bull. Torrey Bot. Club 31: 94. 1904.
Corallina Flabellum Ellis et Solander, Nat. Hist. Zooph. p. 124. pl. 24, fig.
A-C. 1786.
U. Flabellum is the only species which Lamouroux included in Udotea
when he founded the genus in 1812. In 1904, Howe made the combina-
tion U. Flabellum (EIl. et Soland.) Howe on the grounds (Howe, 1907,
p. 495, note) that Lamoroux did not really include the species in Udotea
until 1816 and when he did so he called it U. flabellata instead of U.
Flabellum. A. and E. S. Gepp (1911) in their monograph of the Udoteae
and other writers have accepted Howe’s reasoning and have croted him
with the combination U. Flabellum.
Since 1812 is the accepted date of publication of Udotea ee
and since Corallina Flabellum Ellis et Solander was the only example
cited by Lamouroux as representative of his new genus, there seems to
be little question that he intended the citation to serve as a transfer. In
his paper of 1812, Lamouroux also erected several other genera of marine
algae which are still valid, such as Halimeda, Galaxaura, Liagora, Jania,
Amphiroa and Melobesia with examples of known species cited under
each. If the precedent set by Howe were to be followed, none of the
species referred to these genera by Lamouroux could be considered as
having been effectively transferred. Many writers, including Howe
1Proc. Nat. Acad. Sc. 28: 446-451. 1942; Amer. Jour. Bot. 30: 463-468. 1943.
337
338 FarLowla, Vo. 1, 1944
(1920), have, however, credited Lamouroux with certain of these com-
binations, as for example, Amphiroa Tribulus, Jania rubens and others.
It would seem, therefore, that Lamouroux should be regarded as the
author of the binomial Udotea Flabellum.
Codium dichotomum (Huds.) S. F. Gray, Nat. Arr. Br. Pl. 1. p. 293, 1821.
Codium dichotomum (Huds.) Setchell, Univ. Calif. Publ. Bot. 16: 357. 1931.
Spongia dichotoma Hudson, Fl. Angl. ed. 1. p. 489. 1762.
Codium tomentosum (Huds.) Stackhouse, Ner. Brit. ed. 1. p. XXIV. 1797.
Fucus tomentosus Hudson, Fl. Angl. ed. 2. p. 584. 1778.
This species was known for almost a century and a half as Codium
tomentosum until Setchell pointed out in 1931 that the plant should be
known as C. dichotomum (Huds.) Setch. Setchell gives a detailed ac-
count of the nomenclatural history of the species but in his review of the
literature overlooked S. F. Gray’s Natural Arrangement of British Plants
in which the combination C. dichotomum was made by Gray (1821).
Codium prostratum Levring, Lunds Univ. Arsskr. N. F. Avd. 2, 34(9): 16. fig.
8a—d, pl. 4, fig. 11. 1937.
Codium Vaughani Boérgesen, K. Danske Vidensk. Selsk. Biol. Medd. 15(4) :
70. fig. 24, 25. 1940.
From Bergesen’s (1940) description and figures of Codium Vaughani,
it seems clear that this species from Mauritius is identical with the South
African plant which Levring (1937) had described three years earlier
as C. prostratum. The species is very common in certain localities along
the east coast of South Africa and its occurrence in Mauritius is not
surprising.
‘Dictyota dichotoma (Huds.) Lamour. var. intricata (Ag.) Greville, Alg. Brit.
p. 58. 1830.
Zonaria dichotoma var. intricata Agardh, Sp. Alg. 1 (1). p. 134. 1820.
Dictyota dichotoma var. implexa (Desf.) S. F. Gray, Nat. Arr. Br. Pl. 1.
p. 341. 1821,
Dictyota dichotoma var. implexa (Lamour.) J. Agardh, Sp. Alg. 1. p. 92. 1848.
Fucus implexus Desfontaines, Fl. Atl. 2. p. 423. 1800.
Dictyota implexa (Desf.) Lamouroux, Nouv. Bull. Sci. Soc. Philom. 1: 331.
1809; Journ. de Bot, 2: 43. 1809.
This plant is usually referred to as Dictyota dichotoma var. implexa
(Lamour.) J. Ag. but the epithet implexa should be credited to Desfon-
taines who in 1800 described the alga as Fucus implexus. Lamouroux
in 1809 transferred Fucus implexus to Dictyota but made no claim to
the authorship of the species. Credit for the ternary combination D.
dichotoma var. implexa must go, as far as the writer has been able to
ascertain, to S. F. Gray who in 1821 reduced D. implexa (Desf.) Lamour.
to the status of variety under D. dichotoma. J. Agardh, who is usually
credited with the trinomial, made the transfer in 1848(a), some twenty-
seven years after it had been made by Gray. However, according to
Article 58 of the International Rules of Botanical Nomenclature (1935),
PapenFruss: ALGAL NOMENCLATURE 339
the ternary name D. dichotoma var. implexa (Desf.) S. F. Gray (1821)
is invalidated by D. dichotoma var. intricata (Ag.) Grev. (1830), which is
based on Zonaria dichotoma var. intricata Ag. (1820).
Zonaria subarticulata (Lamour.) Papenfuss comb. nov.
Dictyota subarticulata Lamouroux, Nouv. Bull. Sci. Soc. Philom. 1: 331. 1809;
Journ. de Bot. 2: 41. 1809.
Dictyota interrupta Lamouroux, Ann. Mus. Hist. Nat. 20: 272. pl. 12, fig. 1. 1813.
Fucus interruptus (Lamour.) Turner, Fuci 4. p. 114. pl. 245. 1819.
Zonaria interrupta (Lamour.) Agardh, Sp. Alg. 1 (1). p. 137. 1820; Syst. Alg.
p. 268. 1824; J. Agardh, Sp. Alg. 1. p. 111. 1848 (in part); Harvey, Phyc.
Austr. 4. pl. 190. 1862 (excl. certain syns.).
Phycopteris interrupta (Lamour.) Kiitzing, Sp. Alg. p. 564. 1849 (in part);
Tab. Phyc. 9. p. 27. pl. 67, fig. 1. 1859.
Zonaria Turneriana J. Agardh, Ofvers. Kgl. Svensk. Vetensk.-Akad. Férhandl.
27: 438. 1871; Till Alg. Syst. Afd. 1. p. 48. 1872.
Phycopteris angustata Kiitzing, Tab. Phyc. 9. p. 28. pl. 69, fig. 1. 1859.
In 1809 Lamouroux described Dictyota subarticulata, giving as the
source of his material “Ind. Or., Nov. Hol.” In his later treatment of
Dictyota, in his Essai (1813), Lamouroux substituted the specific epithet
interrupta for subarticulata and at the same time gave as the locality
“Ind. Orient.” only, omitting the “Nov. Hol.” In 1820 Agardh trans-
ferred D. interrupta to Zonaria and the species has since been known as
Z. interrupta. Since the specific name subarticulata antedates interrupta,
the binomial Z. interrupta must yield to the new combination Z. sub-
articulata (Lamour.) Papenf.
In 1834, Suhr recorded a South African alga under the name Z. in-
terrupta, and subsequent writers have generally regarded the South African
plant as representative of Z. interrupta. This plant was described as
Phycopteris cuneata by Kiitzing in 1859 but the name was not accepted,
owing, no doubt, to the fact that J. Agardh in 1871 contended that the
South African plant actually is the Dictyota interrupta (or D. subarticu-
lata) of Lamouroux and the Australian plant a different (new) species
which he described as Zonaria Turneriana. J. Agardh’s identification of
the South African alga with Dictyota interrupta seems to be based pri-
marily upon comparison with Lamouroux’s (1813) figure of D. interrupta
and is unconvincing. Although differing anatomically, the Australian
and South African plants are very similar in habit and it would be im-
possible to settle the identity of D. interrupta purely on the basis of
Lamouroux’s description and figure.
As has been mentioned, Lamouroux in 1809(a) gave as the source of
his material of D. subarticulata the East Indies and Australia, while in
1813, when he called the species D. interrupta, he cited the East Indies
only. As far as the writer is aware, no subsequent collectors have ob-
tained this species, or one resembling it, from the East Indies, which sug-
gests that this locality may have been given in error. On the other hand,
the evidence that the material in question had come from Australia and
340 FartowlA, VOL. 1, 1944
not from South Africa is twofold: firstly, in his article in the Journal
de: Botanique, Lamouroux (1809b) gave the names of Thouin and De la
Billardiére as the persons from whom he had received the material of
D, subarticulata. The writer can find no record of either of these men
having collected in South Africa but it is known that De la Billardiére
did collect in Australia. Secondly, in the paper containing the descrip-
tion of D. swbarticulata, Lamouroux (1809a) recorded a number of species
from Australia but not a single one from South Africa.
Zonaria cuneata (Kiitz.) Papenfuss comb. nov.
Phycopteris cuneata Kiitzing, Tab. Phyc. 9. p. 27. pl. 67, fig. 2. 1859.
This South African species has passed under the name Zonaria in-
terrupta for more than a hundred years, but as was pointed out in the
preceding paragraph this name applies to an Australian plant. Kiitzing in
1859 described the South African plant as Phycopteris cuneata but the
name has been considered as synonymous with Z. interrupta.
Zonaria Harveyana (Pappe ex Kiitz.) Areschoug, Phyc. Cap. p. 26. 1851.
Phycopteris Harveyana Pappe ex Kiitzing, Sp. Alg. p. 564. 1849; Tab. Phyc.
9. p. 26. pl. 66, fig. 2. 1859.
Homoeostrichus multifidus J. Agardh, Anal. Alg. Cont. 1. p. 15. 1894.
Zonaria multifida Harv. ex J. Agardh, Sp. Alg. 1. p. 109. 1848; Till Alg. Syst.
Afd. 1. p. 46. 1872 (not Z. multifida (Sm.) Agardh, Sp. Alg. 1(1). p. 135.
1820).
The genus Homoeostrichus was erected by J. Agardh in 1894 to receive
four species which had previously been included under Zonaria. The
first species listed under Homoeostrichus, and presumably the type of the
genus, is a South African plant which J. Agardh had described in 1848
as Zonaria multifida, a manuscript name of Harvey.
The primary feature distinguishing Homoeostrichus from Zonaria is a
difference in structure of the superficial tissue of the frond: in Homoeo-
strichus the cells composing this layer are of the same width as those of
the medulla, while in Zonaria the superficial cells become divided by a
vertical longitudinal wall and in transverse sections of the frond two
superficial cells consequently correspond in width to each of the under-
lying medullary cells.
It is obvious that this distinction between the two genera, even if con-
stant, is of minor importance and it seems best to unite Homoeostrichus
with Zonaria. Justification for the suppression of Homoeostrichus is
furnished, furthermore, by the fact that in Z. Tourneforti, the type of
Zonaria, the superficial cells are not always divided by a vertical longi-
tudinal wall while in H. multifidus, the type of Homoeostrichus, these
cells not infrequently are divided by such a wall.
As has been stated, H. multifidus is based on Zonaria multifida Harv.
ex J. Agardh (1848a). The latter binomial is invalidated, however, by
the earlier homonym of Z. multifida (Sm.) Agardh (1820). It is there-
PapenFuss: ALGAL NOMENCLATURE 341
fore necessary to employ the specific name Harveyana under which the
plant was described by Kiitzing in 1849 as Phycopteris Harveyana.
Zonaria spiralis (J. Ag.) Papenfuss comb. nov.
Homoeostrichus spiralis J. Agardh, Anal. Alg. Cont. 2. p. 89. 1894.
Zonaria flabellata (Okam.) Papenfuss comb. nov.
Homoeostrichus flabellatus Okamura, Icon. Jap. Alg. 6(6). p. 57. pl. 279. fig.
I1-13. 1931.
Homoeostrichus Sinclairii (H. et H.) J. Ag., Okamura, Icon. Jap. Alg. 4(10).
p. 192. pl. 199, fig. 1-4. 1923.
In consequence of the subordination of Homoeostrichus to the synonymy
of Zonaria, it is necessary to transfer the preceding two species to the
latter genus.
Ecklonia radiata (Ag.) J. Agardh, Sp. Alg. 1: 146. 1848.
Laminaria radiata Agardh, Syn. Alg. Scand. p. XIII. 1817; Sp. Alg. 1. p. 113.
1820.
Fucus radiatus Turner, Fuci 2. p. 161. pl. 134. 1809 (not F. radiatus Good. et
Woodw., Trans. Linn. Soc. 3: 202. 1797).
This species is usually referred to as Ecklonia radiata (Turn.) J. Ag.,
the specific epithet being based on Fucus radiatus Turner. Since this
binomial of Turner (1809) is invalidated by the earlier homonym of F.
radiatus Good. et Woodw. (1797), the authorship of the species must be
credited to Agardh who in 1817 called the plant Laminaria radiata.
Ecklonia biruncinata (Bory) Papenfuss comb. nov.
Laminaria biruncinata Bory, Dict. Class. Hist. Nat. 9: 190. 1826; in Duperrey,
Voy. Coquille p. 101. pl. 10. 1828 (as L. bironcinata).
Ecklonia exasperata (Turn.) J. Agardh, Sp. Alg. 1. p. 146. 1848.
Fucus radiatus 8 exasperatus Turner, Fuci 2. p. 161. 1809.
In a recent article the writer (Papenfuss, 1940, p. 210) gave his reasons
for considering this plantas an independent species rather than as a
variety or a form of Ecklonia radiata. He used the name E. exasperata,
which is based on Fucus radiatus 8 exasperatus Turner (1809), but if this
entity is treated as a species, it is necessary to employ the name biruncinata
under which the plant was described by Bory in 1826 as Laminaria
biruncinata.
Himanthalia elongata (L.) S. F. Gray, Nat. Arr. Br. Pl. 1. p. 389. 1821.
Himanthalia elongata (L.) Setchell, Univ. Calif. Publ. Bot. 16: 358. 1931.
Fucus elongatus Linnaeus, Sp. Pl. ed. 1, 2. p. 1159. 1753.
Himanthalia lorea (L.) Lyngbye, Tent. Hydr. Dan. p. 36. 1819.
Fucus loreus Linnaeus, Syst. Nat. ed. 12, 2. p. 716. 1767.
This species was known as Himanthalia lorea for more than a century
until Setchell pointed out in 1931 that the specific epithet elongata ante-
dates lorea and the plant should be known as H. elongata (L.) Setch.
Setchell, however, overlooked the fact that the combination H. elongata
had already been made by S. F. Gray in 1821.
342 FaRLOWIA, VoL. 1, 1944
Zanardinula Andersoniana (Eaton ex J. Ag.) Papenfuss comb. nov.
Prionitis Andersoniana Eaton ex J. Agardh, Sp. Alg. 3(1). p. 159. 1876; Kylin,
Lunds Univ. Arsskr. N. F. Avd. 2, 37(2): 11. 1941.
Zanardinula australis (J. Ag.) Papenfuss comb. nov.
Phyllotylus australis J. Agardh, Ofvers. Kgl. Svensk. Vetensk.-Akad. Férhandl.
4: 9, 1848,
Prionitis australis (J. Ag.) J. Agardh, Sp. Alg. 2(1). p. 188. 1851; Kylin, Lunds
Univ. Arsskr. N. F. Avd. 2, 37(2): 12. pl. 3, fig. 10. 1941.
Zanardinula linearis (Kylin) Papenfuss comb. nov.
Prionitis linearis Kylin, Lunds Univ. Arsskr. N. F. Avd. 2, 37(2): 12. pl. 4,
fig. 11. 1941.
Zanardinula filiformis (Kylin) Papenfuss comb. nov.
Prionitis filiformis Kylin, Lunds Univ. Arsskr. N. F. Avd. 2, 37(2): 13. 1941.
The change in name of the four preceding species is necessitated by
the fact that the generic name Prionitis J. Agardh (1851) is invalidated
by Prionitis Adanson, a genus of flowering plants described in 1763.
J. De Toni (1936) has proposed the name Zanardinula for the genus
of algae.
Gigartina Hystrix (Ag.) Setchell et Gardner, Univ. Calif. Publ. Bot. 17: 295. 1933
(excl. syn. Kiitz.).
Grateloupia Hystrix Agardh, Sp. Alg. 1(2). p. 223. 1822; in Presl, Reliquiae
Haenkeanae p. 9. 1825.
Chaetangium Hystrix (Ag.) Kitzing, Sp. Alg. p. 793. 1849.
Gigartina Radula var. Radula J. Agardh, Sp. Alg. 2(1). p. 278. 1851 (in part).
Gigartina Radula var. Hystrix (Ag.) J. Agardh, Sp. Alg. 3(1). p. 203. 1876 (in
part).
This species was described by Agardh in 1822 from material collected
by Haenke. As the source of the material, Agardh cited (with a query)
the Cape of Good Hope. The species was later included by Agardh in
the list of algae collected by Haenke (cf. Presl, op. cit. p. 8-12) on the
Spanish expedition under the command of Malaspina. In 1849 Kiitzing
transferred Grateloupia Hystrix to the genus Chaetangium. J. Agardh in
1851 included the plant in his var. Radula of Gigartina Radula but in
1876 made it an independent variety under G. Radula. Setchell and
Gardner in 1933 restored the entity to specific rank under Gigartina. It
should be noted, however, that none of the writers subsequent to Agardh
has seen the type or authentic material of Grateloupia Hystrix.
While examining the South African algae in Herb. Agardh, search was
made for the type of Grateloupia Hystrix but no specimen with this in-
scription in the hand of C. Agardh was to be had. The specimens which
J. Agardh referred to Gigartina Radula var. B Hystrix occur under the
following numbers: 23825 (“e C. B. Spei dedit Turner’), 23826-23829,
23831 (“Pappe”), 23833 (“ded. Turner”). Of these specimens, num-
bers 23825 and 23829 are representative of Gigartina polycarpa (Kiitz.)
Setch. et Gard. while the remaining numbers are of G. bracteata (Gmel.)
Setch. et Gard.
PapENFuss: ALGAL NOMENCLATURE 343
Agardh’s description of Grateloupia Hystrix is very inadequate and
the identity of the species can only be established from an examination
of the type, presumably in the Bohemian Museum at Prague. Since South
Africa was not visited by the Malaspina Expedition, it seems most un-
likely that the Cape was the source of Haenke’s material of G. Hystrix.
It is not unlikely that the material was collected at Monterey, California,
where Haenke apparently collected several marine algae (cf. Kylin, 1941,
pp. 10, 12, 16, 28).
In this connection it might be pointed out that in his list of the algae
of Haenke, Agardh described two species of Cystoseira, C. australis and
C. caudata, which seem to have escaped notice entirely. Another record
of interest is that of Fucus compressus. This little-known species was
first described by Agardh in 1824. As the source of his material, Agardh
gave “in mari australi” but he gave no information as to the collector.
Since several of Haenke’s algae were first described by Agardh in 1822
or in 1824, it seems likely that Haenke might have been the collector of
the type material of F. compressus. It is not improbable, moreover, that the
specimens were secured at Monterey, the source of several of Haenke’s
specimens which were incorrectly designated by Agardh as having come
from “mari australi” (cf. Kylin, 1941). If this is correct, F. com pressus
may prove to be representative of a species of Fucales of the Californian
coast.
Gymnogongrus vermicularis (Ag.) J. Agardh, Sp. Alg. 2(1). p. 323. 1851 (excl.
syn. Gracilaria concinna Mont.)
Sphaerococcus vermicularis Agardh, Syn. Alg. Scand. p. XVII. 1817; Sp. Alg.
1(2). p. 311. 1822. .
Fucus vermicularis Turner, Fuci 4. p. 61. pl. 221. 1819 (not Fucus vermicularis
Gmelin, Hist. Fuc. p. 162. pl. 18, fig. 4. 1768).
Chondrus capensis Kiitzing, Sp. Alg. p. 738. 1849 (excl. syns.); Tab. Phyc.
17. p. 17. pl. 59, fig. a, b. 1867 (not Gymnogongrus capensis (Ag.) J. Ag.).
This South African alga is currently known as Gymnogongrus vermicu-
laris (Turn.) J. Ag., the specific epithet being based on the Fucus vermicu-
laris of Turner. The name Fucus vermicularis Turner (1819) is invalid,
however, since it is a later homonym of Fucus vermicularis Gmelin (1768).
According to Article 69 of the International Rules of Botanical Nomen-
clature (1935) the binomial Gymnogongrus vermicularis would be valid,
however, if the specific epithet were treated as a new name of Agardh?
(1817, p. xvii) — Sphaerococcus vermicularis — and not as a transfer
from Fucus vermicularis Turner.
The source of Turner’s Fucus vermicularis is unkown and some un-
certainty has existed as to whether his specimen actually is representative
of the South African plant. An examination of the specimen (in Herb.
Kew) has convinced the writer, however, that Turner’s plant is repre-
sentative of Gymnogongrus vermicularis.
* Agardh (loc. cit.) cites Turner’s plate of Fucus vermicularis which indicates that
parts of volume 4 of Turner’s Fuci appeared at least as early as 1817. The title page
of this volume is dated 1819.
344, FarLowla, Vo. 1, 1944
Botryocladia Skottsbergii (Bgrg.) Papenfuss comb. nov.
Chrysymenia Skottsbergii Bgrgesen, in Skottsberg, Nat. Hist. Juan Fernandez
and Easter Island 2(3) Botany: 307. fig. 49, 50. 1920.
As can be seen from Bégrgesen’s (1920) description and figures of
Chrysymenia Skottsbergii, the species is representative of Botryocladia
Kylin (1931), a segregate from the genus Chrysymenia.
Gastroclonium ovatum (Huds.) Papenfuss comb. nov.
Fucus ovatus Hudson, Fl. Angl. ed. 1. p. 468. 1762.
Gastroclonium ovale (Huds.) Kiitzing, Phyc. Gen. p. 441. 1843.
Fucus ovalis Hudson, Fl. Angl. ed. 2. p. 573. 1778.
Fucus vermicularis Gmelin, Hist. Fuc. p. 162. pl. 18, fig. 4. 1768 (not Fucus
vermicularis Turner, Fuci 4. p. 61. pl. 221. 1819).
As was pointed out above, under Gymnogongrus vermicularis, the
binomial Fucus vermicularis Gmelin (1768) invalidates F. vermicularis
Turner (1819). In tracing the present status of F. vermicularis Gmelin,
it was of interest to find that this name is included in the synonymy of
Gastroclonium ovale (Huds.) Kiitz., which is based on Fucus ovalis
Hudson. Since F. vermicularis Gmelin (1768) antedates F. ovalis Hudson
(1778) by ten years, it is obvious that the specific epithet of Hudson
has no legal status.
When describing Fucus ovalis in 1778, in the second edition of his
Flora Anglica, Hudson made no reference to the first edition (1762) of
this work. Examination of the first edition shows, however, that the
species was originally described in this edition under the name Fucus
ovatus. Since this name antedates F. vermicularis Gmelin by several
years and since it seems to be the oldest name for the species, it becomes
necessary to make the combination Gastroclonium ovatum (Huds.). There
is no clue in the second edition of Flora Anglica as to Hudson’s reasons
for substituting the name ovalis for ovatus but it seems likely that he con-
sidered the former more appropriate. Subsequent writers have invariably
adopted the name ovalis and the binomial Fucus ovatus has, with few
exceptions, not been included in the synonymy of the species. Of the
numerous works consulted, the synonym F. ovatus Huds. was met with
only in Withering’s Arrangement of British Plants (ed. 3, 4. p. 113. 1796;
ed. 6, 4. p. 136. 1818 — the only editions available to the writer).
The writer wishes to express his appreciation to Professor Lee Bonar
for granting the facilities of the Botanical Department of the University
of California. The study was made during the tenure of a Fellowship
eranted by the Carnegie Corporation of New York.
DEPARTMENT OF BOTANY
UNIVERSITY OF CALIFORNIA
BERKELEY, CALIFORNIA
Papenruss: ALGAL NOMENCLATURE 345
LITERATURE CITED
Agardh, C. A. Synopsis Algarum Scandinaviae. Lund. 1817.
Species Algarum 1(1). Lund. 1820.
————. Ibid. 1(2). Lund. 1822.
————. Systema Algarum. Lund. 1824.
Algae, in Presl, Reliquiae Haenkeanae. Pragae. 1825-1830.
Agardh, J. G. Species Algarum 1. Lund. 18482. |
Nya Alger fran Mexico. Ofvers. Kgl. Svensk. Vetensk.-Akad. Forhandl.
4: 5-17. 1848b.
Species Algarum 2(1). Lund. 1851.
Om Chatham-darnes Alger. Ofvers. Kgl. Svensk. Vetensk.-Akad. Forhandl,
27: 435-456. 1871.
Till algernes systematik. Afd. 1. Lunds Univ. Arsskr, 9: 1-71. 1872.
Species Algarum 3(1). Lund. 1876.
Analecta algologica. Cont. 1. Lunds Univ. Arsskr, 29: 1-144. 1894a.
Ibid. Cont. 2. Ibid. 30: 1-98. 1894b.
Areschoug, J. E. Phyceae Capenses. Uppsala. 1851.
Begrgesen, F. Marine algae from Easter Island, in Skottsberg, The Natural History
of Juan Fernandez and Easter Island 2(3), Botany: 247-309. Uppsala. 1920.
. Some marine algae from Mauritius. Kgl. Danske Vidensk. Selsk., Biol.
Medd. 15(4). 1940.
Bory de Saint Vincent, J. B. Laminaria. Dict. Class. Hist. Nat. 9: 187-191.
Paris. 1826.
Cryptogamie, in Duperrey, Voyage Autour du Monde-Coquille. Paris. 1828.
Desfontaines, R. Flora Atlantica, 2. Paris. 1800.
De Toni, J. Noterelle di Nomenclature Algologica. VII. Primo Elenco di Floridee
Omonime. Published by the author. Brescia. 1936.
Ellis, J., & D. Solander. The Natural History of Many Curious and Uncommon
Zoophytes, Collected from Various Parts of the Globe. London. 1786.
Gepp, A., & E. S. Gepp. Codiaceae of the Siboga Expedition including a mono-
graph of Flabellarieae and Udoteae. Siboga Exp. Monogr. 62. Leiden. 1911.
Gmelin, S. G. Historia Fucorum. St. Petersburg. 1768.
Goodenough, S., and T. J. Woodward. Observations on the British Fuci, with
particular descriptions of each species. Trans. Linn. Soc. 3: 84-235. 1797.
Gray, S. F. Natural Arrangement of British Plants, 1. London. 1821.
Greville, R. K. Algae Britannicae. Edinburgh and London. 1830,
Harvey, W. H. Phycologia Australica 4. London. 1862.
Howe, M. A. Notes on Bahaman algae. Bull. Torrey Bot. Club 31: 93-100. 1904.
- Phycological notes. II]. Further notes on Halimeda and Avrainvillea.
Ibid. 34: 491-516. 1907.
—_—_——. Algae, in Britton and Millspaugh, The Bahama Flora. New York, 1920.
Hudson, G. Flora Anglica, ed. 1. London. 1762.
——. Ibid. ed. 2. London. 1778.
Kiitzing, F. T. Phycologia Generalis. Leipzig. 1843.
———. Species Algarum. Leipzig. 1849.
Tabulae Phycologicae 9. Nordhausen. 1859.
Ibid. 17. Nordhausen. 1867.
Kylin, H. Die Florideenordnung Rhodymeniales. Lunds Univ. Arsskr. N. F. Avd.
2, 27(11). 1931.
Californische Rhodophyceen. Ibid. 37(2). 1941.
Lamouroux, J. V. F. Observations sur la physiologie des algues marines, et de-
scription de cing nouveaux genres de cette famille. Nouv. Bull. Sci. Soc. Philom.
1: 330-333. 1809a.
Exposition des caractéres du genre Dictyota, et tableau des espéces qu’il
renferme. Jour. de Bot. 2: 38-44. 1809b.
346 FarLowia, Vor. 1, 1944,
Extrait d’un mémoire sur la classification des polypiers coralligénes non
entierement pierreaux. Nouv. Bull. Sci. Soc. Philom. 3: 181-188. 1812.
. Essai sur les genres de la famille des thalassiophytes non articulées. Ann.
Mus, Hist. Nat. 20: 21-47, 115-139, 267-293. 1813.
—_———. Histoire des Polypiers Coralligénes Flexibles. Caen. 1816.
Levring, T. Verzeichnis einiger Chlorophyceen und Phaeophyceen yon Siidafrika.
Lunds Univ. Arsskr. N. F. Avd. 2, 34(9). 1937.
Linnaeus, C. Species Plantarum, ed. 1, 2. Stockholm. 1753.
————. Systema Naturae, ed. 12, 2. Stockholm. 1767.
Lyngbye, H. C. Tentamen Hydrophytologiae Danicae. Kobenhavn. 1819,
Okamura, K. Icones of Japanese Algae 4. Tokyo. 1916-23.
——. Ibid. 6. Tokyo. 1929-32.
Papenfuss, G. F. Notes on South African marine algae. J. Bot. Notiser 1940:
200-226. 1940.
Setchell, W. A. Some early algal confusions. Univ. Calif. Publ. Bot. 16: 351-366.
1931.
Setchell, W. A. & N. L. Gardner. A preliminary survey of Gigartina, with special
reference to its Pacific North American species. Ibid. 17: 255-340. 1933.
Stackhouse, J. Nereis Britannica, ed. 1. Bath. 1795-1801.
Suhr, J. N. Uebersicht der Algen, welche von Hrn. Ecklon an der siidafrikan-
ischen Kiiste gefunden worden sind. Flora 17: 721-735, 737-743. 1834.
Turner, D. Fuci 2. London. 1809.
———.. Fuci 4. London. 1819.
Withering, W. An Arrangement of British Plants, ed. 3, 4. Birmingham. 1796.
———. Ibid. ed. 6, 4. London. 1818.
1(3) : 347-385 FARLOWIA January, 1944
NEW SPECIES AND VARIETIES
OF WISCONSIN ALGAE
G. W. Prescott
The history of glaciation in central North America, together with other
geological and geographical factors, has been such that Wisconsin is
richly endowed with a great variety of lakes, ponds, and drainage systems.
A considerable number of lakes, especially in the northern uplands of the
state, are land-locked, while many others are the flowage or drainage type.
The countless bodies of water, so characteristic of the north central states,
exhibit marked differences in their chemical and physical properties.
These dissimilarities together with certain hydrographical features make
possible a varied and abundant algal flora in Wisconsin which has be-
come well known through the published reports of G. M. Smith (1916,
1918, 1920, 1924).
For more than three decades E. A. Birge and C. Juday, together with
their co-workers have carried on limnological studies of the inland lakes
of Wisconsin. It is in relation to their work that Smith (1920-1924)
undertook his survey of the plankton algae, making rather exhaustive
studies of the flora, especially from the lakes in which Birge and Juday
had been conducting their investigations. Professor Juday, Director of
the Trout Lake Limnological Laboratory, invited the author to extend
the survey of Wisconsin algae from both taxonomic and ecological view-
points. Consequently, special attention has been directed toward the
attached or. littoral algae. Collections and field work were carried on
during three summers, mostly in the northeastern and northwestern lakes
regions. A full report on this survey, which will also include the algae
previously reported from the state, is forthcoming.
Many forms which appeared in the author’s collections and in others
loaned to him, are new records for North America. The following thirty
species and varieties apparently are novelties. The desmids were col-
lected and identified but because they are so numerous, descriptions of
them are deferred for a subsequent report. The diatoms are being studied
by Mr. Paul Conger of the Carnegie Institution, Washington, D. C. All
the other algal groups were considered, however, and they are represented
in the state by approximately 1,025 species and varieties, exclusive of
the desmids.
The author is grateful to Dr. Dorothy Schullian and to Dr. Hannah
Croasdale for their valuable assistance in the preparation of the Latin
diagnoses.
347
348 FarLowiA, VoL. 1, 1944
PHYLUM CHLOROPHYTA
A. CHLOROPHYCEAE
CHLAMYDOMONADACEAE
Chlamydomonas polypyrenoideum, sp. nov.
PL. 1, Fic. 1-2,
Cellulae ovoideae vel ellipsoideae, carentes papilla apicali et vagina gelatinosa;
chloroplaste calyce denso parietali cum incisione alta in medio; pyrenoidibus multis
(12-16), sparsis; macula oculari non observata; cellulis 8-10 u lat., 9-12 w long.
Cells ovoid to ellipsoid, without an apical papilla; gelatinous sheath
lacking; chloroplast a dense parietal cup with a deep median invagination ;
pyrenoids many (12-16), scattered; pigment spot not observed; cells
8-10 p» in diameter, 9-12 y long.
Rare in the plankton of Weber Lake, Vilas County.
The differentiating character of this species is the large number of
pyrenoids. Some other Chlamydomonas species have several such organs
but the form of the chloroplast, the shape of the cell and other morphologi-
cal differences prevent the Wisconsin form from being assigned to them.
TETRASPORACEAE
Tetraspora lamellosa, sp. nov.
pL. 1, Fic. 3.
Thallus irregulariter lobatus et bullatus; libere natans; cellulis sphaericis, in pari-
bus dispositis, cum membranis crassis et vaginis lamellatis gelatinosisque, distinctis
et non confluentibus in gluten colonicum; pseudociliis delicatissimis, 20-30-plo
diametro cellulae longioribus; chloroplaste lamina dense parietali paene totum mem-
branum tegente, cum una pyrenoide; cellulis 9-10.5 » lat.
Thallus irregularly lobed and saccate; free-floating; cells spherical, in
twos, with thick walls and gelatinous lamellate sheaths, distinct and not
confluent with the colonial mucilage; pseudocilia very fine, 20-30 times
the diameter of the cell in length; chloroplast a dense parietal plate cov-
ering almost the entire cell wall, with 1 pyrenoid; cells 9-10.5 » in
diameter.
The species differs from others by the possession of distinct lamellated
cell sheaths and the extraordinarily long pseudocilia. In the former char-
acter it resembles some species of Gloeocystis.
Schizochlamys compacta sp. nov.
PL. 1, FIG. 4-6,
Massa plantae microscopica, cum glutine solido, homogeneo et claro tegumento
limitato; cellulis globosis cum apice conspicuo glutinis in uno latere locato, dividente
fissura membrani quae unum fragmentum, rare 2, relinquit; apice glutinis persistente
in antiquo membrano matris, cellulis filialibus habentibus singulos apices glutinis
plerumque contingentes; una chloroplaste parietali cum pyrenoide; cellulis 7.4-1] p
lat.
Plant mass microscopic, mucilage firm, homogeneous and bounded by
a definite tegument; cells globose with a conspicuous cap of gelatine at
Prescott: New WIsconsin ALGAE 349
one side, dividing by a splitting of the cell wall leaving one fragment
(rarely two); the mucilage cap persisting on the old mother cell wall,
daughter cells with respective caps of mucilage usually in contact; one
parietal chloroplast with a pyrenoid; cells 7.4-11 y» in diameter.
Rare in a small pool within a Sphagnum bog near Manitowish River
fish hatchery, Vilas County.
This plant should be compared with S. delicatula West which forms
but a single fragment of the cell wall upon liberation of daughter cells.
It differs in the definiteness of shape of the mucilage, in the larger cells,
and in the mucilage cap on the cell wall which seems to be a constant
character. ;
ULOTRICHACEAE
Ulothrix cylindricum sp. nov.
PL. 1, Fic. 7.
Fila longa, curva, et leviter implicata; cellulis elongato-cylindricis, 11-12.5 y, lat.,
2%4-3-plo longioribus diametro, membranus parallelis et nullas constrictiones mem-
branis transversis habentibus; chloroplaste fascia lata, paene pari longitudine cellulae
et plicata ad mensuram % peripheriae parietalis; pyrenoidibus 2-5.
Filaments long, curved, and'slightly entangled; cells elongate-cylindric,
11-12.5 » in diameter, 214-3 times as long as wide, the walls parallel
and without constrictions at the cross walls; chloroplast a broad band,
nearly equal to the cell in length and folded about 34 of the wall circum-
ference; pyrenoids 2-5.
In the plankton of Cecelia Lake, Vilas County.
This species should be compared with U. aequalis Kuetz. which has
thick walls, shorter cells, fewer pyrenoids and a different form of chloro-
plast. Many species of this genus have been described and there is a
great confusion in the synonymy. It is highly possible that the Wisconsin
plants agree with some previously described varieties of known species.
The proportions of the cell, the form of the chloroplast and the number
of pyrenoids seem to separate our plants from described species as far as
determined by available herbarium material.
Geminella crenulatocollis sp. nov.
pL. 1, Fic. 8.
Planta filum uniseriatum cellularum irregulariter ovoidearum, subquadratarum, vel
oblongarum cum membranis lateralibus emarginatis, crenulatis, aut undatis, truncatis
aut late polis rotundatis, cum rugis et iugis interdum in membranis lateralibus visis;
cellulis inclusis in vagina lata gelatinosa, in paribus linearibus, sed saepe aequaliter
locatis; chloroplaste Jamina parietali irregulariter formata et plicata, quae paene
totum cylindricum in membrano fingit, saepe monstrante iugum aut labrum alatum
radialiter ad membranum extendens; pyrenoide una, in centro locata; cellulis 12-15 p
lat., 18-24 p long.
Plant a uniseriate filament of irregularly ovoid, subquadrate or oblong
cells with emarginate, crenulate, or wavy lateral walls, truncate or broadly
rounded at the poles, with folds and ridges sometimes present in the lateral
walls; cells inclosed in a broad gelatinous sheath, in linear pairs, but
350 FarLowia, VoL. 1, 1944
often evenly spaced; chloroplast an irregularly shaped, folded parietal
plate which almost makes a complete cylinder within the wall, often show-
ing a ridge or wing-like flange extending radially toward the wall; pyrenoid
single, centrally located; cells 12-15 » in diameter, 18-24 p long.
In an acid swamp near Woodruff, Oneida County.
This species should be compared with G. mutabilis (de Bréb.) Wille
from which it differs in the irregularly creased cell wall and the shape
of the chloroplast. The determination of this plant has been confirmed
by observation made on specimens discovered in collections from Che-
boygan County, Michigan.
Hormidiopsis ellipsoideum sp. nov.
PL. 1, Fic. 9-10.
Cellulae transversim ellipticae, collocatae in seriebus linearibus in globis 4 cellulas
continentes, quoque globo incluso vagina gelatinosa lata, hyalina homogeneaque;
chloroplaste fascia parietali latitudine pari longitudine cellulae sed totum mem-
branum non cingente, cum una pyrenoide; cellulis 8 vu. lat., 5.5 w long.; filo 14.8 w lat.
Cells transversely elliptic, arranged in linear series in groups of four,
each group inclosed by a wide, hyaline and homogeneous gelatinous
sheath; chloroplast a parietal band as wide as the length of the cell but
not entirely encircling the wall, with 1 pyrenoid; cells 8 » in diameter,
9.9 p long; filament 14.8 » wide.
In a Sphagnum bog lake near Long Lake, Vilas County.
This plant appeared in but one collection. It is so distinctive that it
seems worthy of description, however. Hormidiopsis is a little under-
stood genus containing, apparently, but one other species, H. crenulata
Heering. To this genus Heering has also questionably assigned a plant
previously described by Borge as Ulothrix moniliformis. Our plant has
the cell shape and approximate size of the latter, but differs in the ar-
rangement of the cells. In U. moniliformis the cells have a continuous
filamentous arrangement, inclosed by a sheath. All of the Wisconsin
specimens had the cells arranged in crooked filaments in linear groups
of four, each group with an individual sheath. Also in our specimens
the chloroplast contained a pyrenoid, a character which is typical for
the genus as described by Heering. This does not preclude our plants
from the genus Hormidiopsis since it is one in which members of the
Ulotrichaceae show considerable variation.
CHAETOPHORACEAE
Stigeoclonium pachydermum sp. nov.
PL. 2, FIG. 1-3,
Planta filum rectum ramosissimumque ex multis ramis rhizoidalibus deorsum emi-
nentibus; ramis irregulariter dispositis et plurimum forma distantibus, plerumque
alternis, sed cum uno ramo surgente protinus super alterum in latere adverso fili
aut in plano rectis angulis posito, saepe evolvente ex cellulis brevibus et ligneo vaso
similibus; multis ramis spinosis et deorsum irregulariter eminentibus evolventibus
ex parte superiore axis principalis, qui rami in acumina obtusa fastigantur et saepe
Prescott: New WIscoNsIN ALGAE 351
finiuntur in cellulis aliquantum dilatatis, quadrangulato-globosis sporangialibusque
quae et intercalares prope terminos ramorum brevium possunt esse; cellulis axis
principalis 19.5—-21 yu, lat., ramorum 15-16 yp. lat., cylindricis, etiam 20-plo longioribus
diametro, aut brevibus et ligneo vaso similibus in eodem filo; membrano cellularum
axis principalis 3-4 yw crass.
Plant an erect, much branched filament from numerous, downward pro-
jecting rhizoidal branches; branches irregularly disposed and extremely
varied in form, mostly alternate, but with one branch arising immediately
above another on the opposite side of the filament or in a plane at right
angles to it, often developing from short, barrel-shaped cells; many thorn-
like and irregular downward projecting branches developing from the
upper part of the main axis, branches tapering to blunt points; frequently
ending in somewhat enlarged; quadrangular-globose sporangial cells which
may be intercalary near the ends of short branches also; cells in the main
axis 19.5-21 » wide, in the branches 15-16 » wide, cylindrical and up to
20 times longer than wide, or short and barrel-shaped in the same fila-
ment; the wall of the cells in the main axis 3-4 p» thick.
In the shallow water of High Lake, Vilas County.
The chief characteristics of this plant are the irregularly arranged and
crooked branches and the thick cell walls. In its coarse habit it resem-
bles S. lubricum which is, however, a species with opposite branching.
Rhizoidal types of branches appear both near the base of the plant and
toward the apices of the main axes.
Draparnaldia Judayi sp. nov.
PL. 2, FIG. 4-5.
Planta mollissimo aquosoque glutine investita; axe principali composito ex cellulis
gracilibus, paulo inflatis aut cylindricis cum constrictionibus in membrana transverso
locatis; ramis secundariis lateralibus absentibus; fasciculis parvorum ramorum ad-
versis aut tortis, surgentibus angulis rectis ex regione media cellularum axis, fasci-
culis paulum ramosis et cum rachide viso tantum in maioribus et bene formatis;
fasciculis saepe simplicibus aut si ramosis habentibus ramulos unae aut duarum
cellularum, apicibus plurimorum ramulorum finientibus in setis validis rectisque quae
saepe bulbosae in basi sunt, nonnumquam surgunt ex latere aut dichotome divisae
sunt; choloroplaste fascia maiorem partem membrani in cellulis ramulorum angusta
in cellulis axis principalis quae sunt tegente; 12-15.2 y, lat., 30-40 wp long.
Entangled about overhanging grass in a Sphagnum bog pool near
Rhinelander, Oneida County.
One of the most distinctive characteristics of this species is the whorled
arrangement of the very simple and much reduced fascicles of branches,
arising, as they do, from the midregion rather than from the joint of the
main axial cell. The position of the branchlet origin is remindful of
Draparnaldiopsis alpinis Smith and Klyver. In that species, however,
the stalked fascicles arise from short, differentiated cells in the main fila-
ment. Another peculiarity is the form and location of the setae. These
may be terminal, one or two arising from a non-tapering apical cell, or
lateral, in which case they take the place of a dichotomous branch. The
setae are similar in morphology to those of Chaetonema.
352 FarLowiA, Vou. 1, 1944
OEDOGONIACEAE
Oedogonium crenulatocostatum var. cylindricum fa. major fa. nov.
Forma maior quam ordinaria; cellulis vegetativis 25.9 y, lat., 88.8 wu long.; oogoniis
1-3, obovoideo-ellipsoideis aut cylindrico-oblongis, aperientibus poro superiore, 42.5-
44 y, lat., 63-74 » long.; oospora ovoideo-ellipsoidea, membrano crasso, strato medio
cum circa 16 costis in longitudinem positis quae satis leves, non crenulatae sunt,
37-39 yw lat., 55.5-57 uw long.
A form larger than the typical; vegetative cells 25.9 » in diameter,
88.8 » long; oogonia 1-3, obovoid-ellipsoid or cylindric-oblong, opening
by a superior pore, 42.544 y in diameter, 63-74 » long; oospore ovoid-
ellipsoid, wall thick, middle layer with about 16 longitudinal ribs which
are quite smooth (not crenulate), 37-39 » in diameter, 55.5-57 yw long.
Floating in the back water of Lost Canoe Lake, Vilas County.
Our specimens are stouter and somewhat larger throughout than the
typical variety.
Oedogonium K jellmanii var. granulosa yar. nov.
PL. 5, FIG. 1-3,
Cellulae vegetativae cylindricae, 19.5-23.4 u lat., 109-117 yu. long., oogonio solitario,
ellipsoideo, superiore poro aperiente, 53-55 pv, lat., 97-102 p. long.; oospora ellipsoidea,
oogonium non replente, membrano tria strata habente, exterius leve, medium cum
circa 22 costis in longitudinem positis quae crenulatae granulataeque, saepe inter-
ruptae, sed sine anastomose sunt, 50.7-53 uw lat., 74-76 w long.
Vegetative cells cylindrical, 19.5—-23.4 « in diameter, 109-117 yp» long,
oogonium solitary, ellipsoid, opening by a superior pore, 53-55 » in
diameter, 97-102 » long; oospore ellipsoid, not filling the oogonium,
wall of three layers, the outer smooth, the middle layer with about 22
longitudinal ribs which are crenulate and granular, frequently interrupted,
but not anastomosing, 50.7-53 y» in diameter, 74-76 p long.
Attached to old wood in a roadside tarn near Boulder Junction, Vilas
County.
This plant should be compared with Oe. margaritiferum Hirn which
has similarly marked oospore walls in both the outer and middle layers.
The variety differs from the typical form of Oe. Kjellmanii in its larger
size and in the different marking of the spore wall.
Oedogonium oviforme var. gracile fa. nov.
PL. 5, FIG. 4-5...
Forma cum cellulis vegetativis gracilioribus et oogoniis minoribus quam in planta
ordinaria; cellulis vegetativis 7.6-9.2 lat., 103.6-125 ww long.; oogonio solitario,
ellipsoideo-ovoideo, 44.446 y, lat., 51.8-53 y. long.; oospora ellipsoidea, paene oogon-
ium replente, membrano levi, 40.7 u. lat., 49.9 u long.; cellulis antheridialibus 9.2 u
lat., 11.1 yw long.; spermatibus duobus, cum divisione horizontali.
A form with more slender vegetative cells and smaller oogonia than
the typical plant; vegetative cells 7.6-9.2 » in diameter, 103.6-125 yu
long; oogonium solitary, ellipsoid-ovoid, 44.4-46 » in diameter, 51.8-
53 p» long; oospore ellipsoid, nearly filling the oogonium, wall smooth,
Prescott: New WIScoNSsIN ALGAE 353
40.7 » in diameter, 49.9 » long; antheridial cells 9.2 » in diameter,
11.1 » long; sperms 2, division horizontal.
Attached to submerged aquatics in a slough cut off from Lost Canoe
Lake, Vilas County.
Oedogonium Smithii sp. nov.
PL. 5, FIG. 6-7.
Macrandrosum; dioeciosum (?); cellulis vegetativis cylindricis aut irregulariter
inflatis, 3.7-8 p lat., 13-25 p» long.; oogonio solitario, late pyriformi-fusiformi cum
inflatione laterali. secondaria in parte superiore, operculato, cum divisione in medio,
lato, 22-25 y, lat., 27-32 pw long.; oospora depresso-globosa, paene oogonium a latere
replente, membrano levi, 16-18 yu, lat., 12.9-14 yp, long.; antheridio (?).
Macrandrous; dioecious (?); vegetative cells cylindrical or irregu-
larly inflated, 3.7-8 » in diameter, 13-25 » long; oogonium solitary,
broadly pyriform-fusiform with a secondary lateral inflation in the upper
portion, operculate, division median, wide, 22-25 y» in diameter, 27-32 p
long; oospore depressed-globose, nearly filling the oogonium laterally,
wall smooth, 16-18 p» in diameter, 12.9-14 » long; antheridium (?).
Rare in a roadside pond near Genoa City, Washburn County. This
plant is named for Dr. Gilbert M. Smith.
This species should be compared with Oe. inconspicuum Hirn. It is
distinguished by the pyriform oogonium with its lateral inflations.
Oedogonium microgonium sp. nov.
PL. 5, Fic. 8.
Macrandrosum; dioeciosum (?); cellulis vegetativis clare capitellatis, 8-9.5 u lat.,
18-33 ». long.; in basi cellulis elongatis, 16-18 p. long.; oogonio solitario, rare cum
duobus, sed paululo latiore quam cellulae vegetativae, globoso aut depresso-globoso,
operculato, cum divisione superiore, 11-12.9 y, lat., 10-11.2 yp, long.; oospora globosa,
oogonium replente, membrano levi, 11-12 y, lat.; cellulis antheridialibus non observatis.
Macrandrous; dioecious (?); vegetative cells distinctly capitellate,
8-9.5 » in diameter, 18-33 » long; basal cell elongate, 16-18.5 » long;
oogonium solitary (rarely two together), but very little wider than the
vegetative cells, globose or depressed-globose, operculate, division su-
perior, 11-12.9 » in diameter, 10—11.2 » long; oospore globose, filling
the oogonium, the wall smooth, 11-12 w in diameter; antheridial cells
not observed.
In several soft water lakes and acid swamps, Vilas County.
This species should be compared with Oe. Howei Tiffany which is larger
and which has an inferior division of the oogonium. Although many
individuals were examined antheridia were not found.
Oedogonium spheroideum sp. nov.
PL. 5, Fic. 9.
Macrandrosum; monoeciosum; cellulis vegetativis longis ac cylindricis, 16-19 pu
lat., 115-155 p long.; oogonio solitario, late ellipsoideo-subgloboso, operculato, cum
divisione superiore, 55-64 yw lat., 80-87.5 » long.; oospora spheroidali, membrano
crasso, membrano exteriore cum 12-15 costis in longitudinem positis, 57-60 p lat.,
57-60 uw long.; cellulis antheridialibus 1-4, protinus infra oogonium locatis, aut
sparsis, spermatibus 2, cum divisione horizontali, 19-21 yp, lat., 16 y long.
304 Fartowia, VoL. 1, 1944
Macrandrous; monoecious; vegetative cells long and cylindrical 16-19 p»
in diameter, 115-155 » long; oogonium solitary, broadly ellipsoid to
subglobose, operculate, division superior, 55-64 » in diameter, 80-87.5 Mm
long; oospore spheroidal, the wall thick, outer membrane with 12-15
longitudinal ribs, 57-60 » in diameter, 57-60 » long; antheridial cells
1-4, immediately below the oogonium, or scattered, sperms two, the di-
vision horizontal, 19-21 » in diameter, 16 » long.
Attached to grass in a roadside swamp near Fishtrap Lake, Vilas County,
This species should be compared with Oe. sol Hirn which is smaller
and has an oospore with the middle rather than the outer layer of the
wall ribbed.
Oedogonium Sawyeri sp. nov. ;
PL. 5, FIG. 10,
Nanandrosum; gynandrosporosum; cellulis yegetativis cylindricis, validis, 30-33.3 U.
lat., 66.6-81 yp. long.; oogonio solitario, paene globoso, 63-66.6 p lat., 55-60 w, long,
aperiente poro superiore; cellula suffultoria inflata; oospora globosa, membrano ex-
teriore cum claro iugo spirali, continuo a polo ad polum, axe sporae converso in
angulo c, 30 graduum an axe oogonii in longitudine viso, 50-55 u. lat. (iugo incluso) ;
plantis masculinis in cellula suffultoria locatus; cellulis antheridialibus 8-10 vu. lat.;
androsporangiis 25.9 p. lat., 14-18 » long.
Nanandrous; gynandrosporous; vegetative cells cylindrical, stout, 30-
33.3 » in diameter, 66.6-81 » long; oogonium solitary, nearly globose,
63-66.6 » in diameter, 55-60 » long, opening by a superior pore; sufful-
tory cell swollen; oospore globose, outer membrane with a prominent
spiral ridge, continuous from pole to pole, the axis of the spore turned
at an angle of about 30 degrees from the longitudinal axis of the oogon-
ium, 50-95 » in diameter (including ridge) ; male plants on the suffultory
cell; antheridial cells 8-10 » in diameter; androsporangia 25.9 » in
diameter, 14.8 » long.
In a cut-off from Lost Canoe Lake, Vilas County. This plant is named
for Mr. Webster Sawyer.
Oe. Sawyeri should be compared with the smaller idioandrosporous
Oe, latviense (Tiff.) Tiffany and Oe. spiripennatum Jao. The latter has
a median rather than a superior pore.
Oedogonium sinuatum fa. seriatum fa. nov.
Cellulae vegetativae undulatae et capitellatae, cum 4 undulationibus in medio,
22-25 yp, lat., 48-59.2-(140) yw long.; oogoniis 5 in seriebus, globosis, operculatis,
cum divisione inferiore, 62-67 py. lat., 62-72 u long.; oospora globosa, oogonium non
replente, membrano levi, 55.5 yu lat.; plantis masculinis pusillis paene semper con-
densis in cellula suffultoria; cellulis antheridialibus exterieribus (?), 10.9 w lat.
Vegetative cells undulate and capitellate, with 4 median undulations,
22-25 in diameter, 48-59.2-(140) » long; oogonia five in series,
globose, operculate, division inferior, 62-67 » in diameter, 62-72 p» long;
oospore globose, not filling the oogonium, wall smooth, 55.5 in diameter;
dwarf male plants usually crowded on the suffultory cell; antheridial cells
exterior (?), 10.9 » in diameter.
Prescott: New WIsconsin ALGAE 355
Attached to overhanging grass in a Sphagnum bog ditch near highway
between Woodruff and Rhinelander, Oneida County.
This plant should be compared with Oe. undulatum (Bréb.) A. Braun
with which it has some features in common. The chief characteristic of
the Wisconsin plants is the decidedly seriate arrangement of the oogonia.
Oedogonium Kozminskii sp. nov.
PL. 5, Fic. 11.
Nanandrosum; idioandrosporosum; cellulis vegetativis cylindricis vel paulo capi-
tellatis, 14-15.6 pw lat., 50-60 » long.; oogonio solitario, globoso, operculato, cum
divisione supramediana, 46—50.7 y, lat., 46-53.7 y, long.; oospora globosa vel depresso-
globosa membrano levi, 43.9-45 uy. lat., 39-42 pw. long.; cellula suffultoria haud dilatata
vel paululo; plantis masculinis pusillis unicellularibus in oogonio locatis, 11.7 y, lat.,
13-14 y, long.
Nanandrous; idioandrosporous;-. vegetative cells cylindrical to slightly
capitellate, 14-15.6 » in diameter, 50-60 » long; oogonium solitary,
globose, operculate, division supramedian, 46-50.7 » in diameter, 46—
53.7 » long; oospore globose or depressed-globose, wall smooth, 43.9-45 pu
in diameter, 39-42 uw long; suffultory cell not or scarcely enlarged; dwarf
male plants unicellular, on the oogonium, 11.7 » in diameter, 13-14 y long.
Attached to grass in a bog near Arbor Vitae Lake, Vilas County. This
plant is named for Dr. Zygmunt Kozminski.
This species is quite unlike any other in the genus. The shape and
markings of the oospore together with the operculate oogonium and the
numerous two-celled male plants, form a combination of characteristics
which makes this species unique.
Oedogonium oelandicum var, contortum yar. nov.
PL. 5, Fic. 13-14.
Varietas ordinaria differens in filis contortis quae nonnumquam breves spiras form-
ant, multae quarum in uno filo possunt reperiri; cellulis vegetativis 11-12 yp, lat.,
distincte capitellatis; oogonio multo breviore quam lat., depresso-globoso, 29-30 yp.
lat., 20-21.5 p. long.; oospora depresso-globosa, membrano levi, 25-27 y, lat., 18-20 »
long.; plantis masculinis pusillis unicellularibus in oogonio locatis,
A variety differing from the typical form by its contorted filaments which
sometimes form short spirals, several of which may occur in one filament;
vegetative cells 11-12 » in diameter, distinctly capitulate; oogonium much
shorter than wide, depressed-globose, 29-30 m in diameter, 20—21.5 p
long; oospore depressed-globose, the wall smooth, 25-27 » in diameter,
18-20 » long; dwarf male plants unicellular, on the oogonium.
Attached to grass in a small lake near Barber Lake, Sawyer County.
Oedogonium polyandrium sp. nov.
PL. 5, Fic. 12.
Nanandrosum; idioandrosporum (?); cellulis vegetativis paulo capitellatis, 4-5.4 uv.
lat., 14-30 long.; oogonio solitario, ovoideo vel late ellipsoideo, operculato, cum
apertura superiore, 17-19 p» lat., 27-29 uw long.; oospora ovoidea, oogonium paene
replente, membrano exteriore sporae cum scrobiculationibus crassis sparsisque vel
cum puteis brevibus, stratis medio et interiore membrani sporae levibus, 15-17 wu
356 FarLowla, VoL. 1, 1944
lat., 22-25 » long.; plantis masculinis pusillus bicellularibus, numerosis, in oogonio
locatis, stipe 4.5 u lat., 14-16 y long.; cellula antheridiali exteriore (?).
Nanandrous; idioandrosporous (?) ; vegetative cells slightly capitellate,
4-5.4 » in diameter, 14-30 » long; oogonium solitary, ovoid or broadly
ellipsoid, operculate, opening superior, 17-19 » in diameter, 27-29 Mm
long; oospore ovoid, nearly filling the oogonium; outer spore wall with
coarse, sparsely arranged scrobiculations or shallow pits, middle and
inner layers of the spore wall smooth, 15-17 » in diameter, 22—25 y long;
dwarf male plants two-celled, numerous, on the oogonium, stipe 4.5 pu
in diameter, 14-16 » long; antheridial cell exterior.
Attached to grass in a bog near Arbor Vitae Lake, Vilas County.
This species is quite unlike any other in the genus. The shape and
markings of the oospore, together with the operculate oogonium and the
numerous two-celled male plants form a combination of characteristics
which makes this species unique. ’
HyYDRODICTYACEAE
Pediastrum biradiatum var. emarginatum fa. convexum fa. nov.
PL. 2, FIG. 6.
Colonia perforata (clathrata), cellulis bilobatis in peripheria, lobis bifurcatis,
margine interiore lobularum convexo; cellulis periphericis adiunctis tantum in parte
inferiore marginum lateralium, lobis cellularum interiorum solummodo emarginatis;
cellulis 11.7 y, lat., 9.7-10 uw long.; colonia 16 cellularum 44-50 v. lat.
Colony perforate (clathrate), cells bilobed at the periphery, the lobes
bifurcate, the inner margin of the lobules convex; peripheral cells ad-
joined along the lower part of their lateral margins only, lobes of inner
cells merely emarginate; cells 11.7 » in diameter, 9.7-10 » long; 16-celled
colony 44-50 y» wide.
Rare in a Sphagnum bog near Woodruff, Oneida County.
Pediastrum muticum var. crenulatum var. nov.
PL. 2, FIG. 7.
Varietas ordinaria differens membranis crenulatis vel inaequaliter undulatis et in
finitimis et in exterioribus superficiebus liberis locatis; colonia magna, cum etiam
134 cellulis (34~31-27-21-15-6) ; cellulis 18-24.5 p. lat., colonia 167 uw lat., 340 uw long.
A variety differing from the typical form by having crenulate or irregu-
larly wavy walls on both the adjoining and outer hes surfaces; colony
large, as many as 134 cells (34-31-27-21-15-6); cells 18-24.5 » in
diameter, colony 167 » wide, 340 » long.
Rare in the plankton of Lost Canoe Lake, Vilas County.
Pediastrum quadricornutum sp. nov.
pL. 1, Fic. 11.
Colonia paene integra, cum parvissimis intersticibus formatis ex marginibus retusis
nonnullarum cellularum; colonia oblonga, rare subcirculari (7-1) (16-11-5) (16-
9-7), composita ex 8-32 cellulis alte bilobatis quae habent sinum altum angustumque
formatem duas lobas maiores ita incisas ut fingant lobulas obtuse rotundatas, duae
centrales quarum omnino aut paene coniunctae sunt, sic sinum extrinsecus claudentes,
Prescott: New WISCONSIN ALGAE B57
duae lobulae laterales quarum tangunt lobulas laterales cellularum contiguarum;
cellulis interioribus paene eiusdem formae quam cellulae periphericae sed cum lobulis
minus prominentibus et nonnumquam absentibus, cum membrano tantummodo emar-
ginato aut paene recto; cellulis 10.5-18 yw lat.; colonia 8 cellularum 154 y, lat.
Colony nearly entire, with minute interstices formed by the retuse mar-
gins of some cells; colony oblong, rarely subcircular (7-1) (16—11-5)
(16-9-7), composed of 8-32 deeply bilobed cells which have a deep,
narrow sinus forming two major lobes, the lobes incised to form bluntly-
rounded lobules, the two central lobules in contact or nearly so, thus
closing the sinus outwardly, the two lateral lobules in contact with the
lateral lobules of the adjoining cells; interior cells about the same shape
as the peripheral cells but with the lobules not so prominent, sometimes
wanting, with the wall merely emarginate or nearly straight; cells 10.5—
18 » in diameter; eight-celled colony 154 pw wide.
Plankton in Beaver Lake and Trostel Lake, Vilas County.
According to Bigeard (1936) such a form as this would be included in
P. tetras (Ehr.) Ralfs, a species which has many variations. The dis-
tinctive shape of the peripheral cells, and the narrow, closed sinus are
constant characters and justify separating the Wisconsin plants from
that species.
OOCYSTACEAE
Qocystis pyriformis sp. nov.
PL. 1, Fic. 12-13.
Cellulae late pyriformi-ovoideae, cum apiculatione conspicua uno polo, altero late
rotundato; consociatae in familiis duarum vel quattuor cellularum; chloroplaste
solida et parietali cum una pyrenoide; cellulis 14-16 y, lat., 16-19 y, long.; colonia
4 cellularum 36 yp, lat., 48.8 » long.
Cells broadly pyriform-ovoid, with a prominent apiculation at one pole,
the other end broadly rounded; united in families of 2-4; chloroplast
massive and parietal with one pyrenoid; cells 14-16 » in diameter, 16—
19 w long; colony of four cells 36 » in diameter, 48.8 » long.
Plankton in cedar swamp near Antigo, Langlade County.
This species should be compared with O. apiculata W. West, a much
smaller plant with broadly elliptic cells. The pyriform shape of the
Wisconsin plants is the chief differentiating characteristic.
Tetraédron asymmetricum sp. nov.
PL. 1, Fic. 14,
Cellula forma quadrangulata, incisa sine symmetria ita ut formet duas lobas maiores
quae isthmum sinu late aperto praetectum habent; duae lobae maiores paulum bilo-
batae, lobulis 2-3 spinis brevibus praefixis; cellulae 10-18 » in mensura longissima.
Cell quadrangular in outline, unsymmetrically incised to form two major
lobes with an isthmus bordered by a widely open sinus, the two major
lobes slightly bilobed, the lobules tipped with 2—3 short spines; cells
10-18 » in their longest dimension.
Plankton in Big Clam Lake, Burnett County.
358 FarLowia, Vou. 1, 1944
This species should be compared with T. irregulare (Reinsch) de Toni
which it more nearly resembles than any other.
Tetraédron bifurcatum var. minor var. nov.
pL. 1, Fic. 15.
Forma ab ordinaria differens in mensura minore et quod lobas bifurcatas habet,
cum lobulis firmioribus et brevi spina praefixis; cellulae 22.5 u in diametro maximo.
A form differing from the typical by its smaller size and in having the
lobes bifurcated, the lobules rather stout and tipped with a short spine;
cells 22.5 » in their maximum diameter.
Plankton in Okauchee Lake, Waukesha County.
Tetraédron cruciatum var. reductum var. nov.
pL. 1, Fic. 16.
Cellula plana, irregulariter cruciformis vel nonnumquam trilobata, lobis bifurcatus,
lobulis brevi spina praefixis; margines cellulae in duobus lateribus oppositis concavi,
in alteris lateribus recti aut tantum paululum concavi; cellulae 28-30 vu. lat., etiam
54 w% in mensura longissima.
Cell flat, irregularly cruciform or sometimes three-lobed, the lobes
bifurcate, the lobules tipped with a short spine; margins of the cell con-
cave on two opposite sides, straight or only slightly concave on the other
sides; cells 28-30 » in diameter, up to 54 yu in the greatest dimension.
Plankton in Lost Canoe Lake and in Manitowish River, Vilas County.
This form differs from the typical in its broad, bilobed processes and
in having the lateral walls much less concave or emarginate. It differs
from var. polyfurcatum G, M. Smith in the form and number of processes,
Tetraédron enorme yar. pentaédricum var nov.
PL. 1, Fic. 17.
Cellulae forma quinquelaterali, lateribus rectis vel paulo convexis, paribus pro-
cessuum angustorum bifurcatorumque in omnibus planis extendentibus et brevibus
spinis praefixis; cellulae 50-55 uy. lat., processibus non inclusis.
Cells five-sided in outline, the sides straight or slightly convex, with
pairs of narrow, bifurcated processes extending in all planes, the processes
tipped with short spines; cells 50-55 » in diameter without processes.
Rare in the plankton of Sweeney Lake, Vilas County.
This form differs from the typical plant in its straight margins and
narrow, bifurcated processes extending from the angles. It is similar to
Borge’s figure of T. enorme (Ralfs) Hansgirg, which we judge to be
atypical of that species.
Tetraédron limneticum var. gracile var. nov.
pL. 1, Fic. 18.
Cellulae pyramidatae vel tetragonales, angulis in processus bifurcatos extentis qui
2-3 spinis firmis praefixi sunt; margines cellulae concavi inter processus; bases pro-
cessuum contiguae ita ut corpus cellulae vix visibile sit; cellulae 40-46.8 y. lat.,
processus 6-8 y, lat.
Prescott: New WIsconsin ALGAE 359
Cells pyramidal, or tetragonal, the angles extended into bifurcate pro-
cesses which are tipped with 2-3 stout spines, the margins of the cell
concave between the processes; bases of the processes adjoining so that
there is scarcely a cell body; cells 40—46.8 » in diameter; processes 6-8 p,
wide.
Plankton in Elcho Lake, Langlade County.
The processes of this form are so narrow and the cell body so greatly
reduced that it approaches the genus T’reubaria.
Tetraédron lobulatum var. crassum var. nov.
PL. 2, FIG. 8.
Cellulae pyramidatae, tetragonales, angulis leviter productis ita ut forment pro-
cessus comparate latos fortesque qui bilobati sunt, lobulis bifurcatis et in acuminibus
brevibus, curvis, spinosisque finitis; cellulae 25-30 py, lat.
Cells pyramidal, tetragonal, the angles slightly produced to form rela-
tively wide, stout processes which are bilobed, the lobules bifurcate and
ending in short, curved, spine-like tips; cells 25-30 » in diameter.
Plankton in Lost Canoe Lake and in Manitowish River, Vilas County.
Tetraédron regulare var. granulata var. nov.
PL. 3, Fic. 1.
Cellulae magnae, pyramidatae, lobis latis fortibusque cum marginibus convexis
instructis, angulis late rotundatis et spina solitaria brevique praefixis quae multo
reducta esse potest adeo ut tantummodo papilla sit; membranum punctatum et parvis
granulis aut asperitatibus copertum; cellulae 35-51.8 y, lat.
Cells large, pyramidal, the lobes broad and stout, with convex margins,
the angles broadly rounded and tipped with a single short spine which
may be reduced to a mere papilla; wall punctate and covered with small
granules or roughenings; cells 35—51.8 » in diameter.
Plankton in Flambeau Lake, Vilas County; Little Round Lake, Sawyer
County.
Tetraédron regulare var. incus fa. major fa. nov.
PIs osckIGs 2.
Cellulae pyramidatae, tetragonales marginibus rectis vel leviter convexis, angulis
productis ut spinas longas fortesque forment; cellulae 35-50 yu lat. spinis inclusis;
spinae 12-13.5 y. long. :
Cells pyramidal, tetragonal, the margins straight or slightly convex,
the angles produced to form long, stout spines; cells 35-50 y» in diameter
including spines; spines 12—13.5 p» long.
Plankton from Sweeney Lake; in tow from Manitowish River, Vilas
County.
ScCENEDESMACEAE
Scenedesmus opoliensis var. contacta var. nov.
PL. 3, Fic. 3.
Planta constans ex 4 cellulis naviformibus in serie singula instructis, per 34 longi-
tudinem parietum lateralium coniunctis; spinis cellularum terminalium, 1 vel 2
360 FarLowI!A, VoL. 1, 1944
utroque in polo, longius curvatisque; spinis cellularum interiorum brevibus rectisque,
cellulis 6-8 p. diam., 20-24 uw long.
Plant consisting of four naviculoid cells arranged in a single series,
adjoined along 34 of the length of their lateral walls; spines on terminal
cells either one or two at each pole, long and curved; spines on the inner
cells short and straight; cells 6-8 yu in diameter, 20-24 py long.
Plankton in Carpenter Lake, Vilas County.
Tetradesmus Smithii sp. nov.
PL. 3, FIG. 4-5.
Cellulae leviter arcuatae aut lunatae aut subcylindratae, parum coartatae in
regione polorum qui sunt late rotundati, locatae in globis quattuor cellulas conti-
nentibus, axibus longis parallelis et membranis convexis adiunctis solum in regione
media, membranis externis liberisque concavis aut paene rectis; cellulae a polo
visae sphaericae, in quadrangula dispositae; chloroplastis lamina parietalis; cellulae
3.7—-4 yw. lat., 27-29 w long.
Cells slightly arcuate or crescent-shaped or subcylindrical, slightly nar-
rowed toward the poles which are broadly rounded, in groups of four
with their long axes parallel and with the convex walls adjoined in the
midregion only, outer free walls concave or nearly straight; cells in end
view spherical, arranged in a quadrangle; chloroplast a parietal plate;
cells 3.7—4 » in diameter, 27-29 yu long.
Rare in the plankton of Devil’s Lake, Burnett County. This species
is named for Dr. Gilbert M. Smith.
Tetradesmus Smithii is very similar to some forms of Quadrigula, but
the arrangement of the cells is very definite and the colonies are without
a gelatinous sheath or envelope.
CONJUGALES
ZYGNEMATACEAE
Spirogyra pseudofloridana Prescott, n. sp.
TEXT Fic. 1.
Fila cellularum fortium cylindratarumque membrana plane finita habentium,
51-60 » lat., 3-5-plo longioribus diametro; chloroplastes 4-(5) angustae, laxe spirales
aut paene rectae, 4-114 revolutiones in cellula facientes; coniugatio scalariformis
facta per inclinationem geniculatum fili ita ut cellulae in iuxtapositione ponantur,
gibbis lateralibus (sed non tubulis) nascentibus in cellulis gametangialibus quae
breves et ut videtur cellulis vegetativis abscissae fiunt et graviter corporibus oleaceis
inter coniugationem onustae sunt, cellulis fertilibus leviter tumidis; zygospora ellip-
soidea cum polis anguste rotundatis, mesospora fulva atque granulationibus irregu-
laribus et rugis (non reticulatis) asperata, 63-70 uy, lat., 100-120 u long.
Filaments of stout cylindrical cells with plane end walls, 51-60 » in
diameter, 3-5 times their diameter in length; chloroplasts 4—(5), nar-
row, loosely spiralled or nearly straight, making 14-11% turns in the cell;
conjugation scalariform by geniculate bending of the filament to bring
the cells into juxtaposition, lateral bulges but not tubes forming on the
gametangial cells which become shortened and apparently cut off from
the vegetative cells, becoming heavily packed with oil bodies at the time
Prescott: NEw WIScoNSIN ALGAE . 361
Ficure 1. Spirogyra pseudofloridana Prescott n. sp. X 250.
of conjugation, the fertile cells becoming slightly swollen; zygospore
ellipsoid with narrowly rounding poles, the mesospore brown and rough-
ened with irregular granulations and wrinkles (not definitely reticulate),
63-70 p» in diameter, 100-120 yp long.
Floating clots in a lagoon-like cove, North Trout Lake, Vilas County.
This species should be compared with S. illinoisensis Transeau and S.
floridana Transeau. It is smaller than the former and has fewer chloro-
plasts. From the latter it differs chiefly in the decoration of the mesospore
layer of the zygospore wall.
PHYLUM CHRYSOPHYTA
A. XANTHOPHYCEAE
STIPITOCOCCACEAE
Stipitococcus apiculatus sp. nov.
PL. 3, FIG. 6-10.
Cellulae ovoideae vel fusiformes, apiculatae, stipite tenui coniunctoque instructae;
chloroplastes 1-3; corpus oleaceum et filus rhizoidalis in apice locati; cellulae gre-
gales, adiunctae turbinibus circum desmidibus filamentosis, nonnumquam 2 vel 3
cellulis in serie in una stirpe locatis, protoplastibus filamentis rhizoidalibus connexis;
cellulae 3.8-4 y, lat., 18-36 p long., stipe incluso.
Cells ovoid or fusiform, apiculate, with a slender attaching stipe;
chloroplasts 1-3; an oil body and a rhizoidal thread at the apex; cells
gregarious, attached in whorls about filamentous desmids, sometimes two
or three cells in a series on one stalk, the protoplasts connected by rhi-
zoidal threads; cells 3.8—4 « in diameter, 18-36 » long including the stipe.
On Hyalotheca filaments; pond near Silver Lake, Langlade County.
362 FarLtowia, Vo. 1, 1944
The arrangement of the cells in dense, transverse zones about the host
filament gives a very distinct appearance to this species. The occurrence
of two or three cells in a series on a single attaching stipe is unlike any
of the described species. It is thought that this condition must be the
result of in situ germination of swarmers.
Stipitococcus crassistipatus sp. nov.
pL. 3, Fic. 11-14.
Cellula late (nonnumquam anguste) in forma ampulla figurata, coartata in parte
anteriore in collum breve latumque et reductum in parte posteriore in stipitem
crassum et 1.5-2 u lat.; protoplastis ovalis vel subglobosa, 2 chloroplastibus laminatis
instructa; 7.6-8 wu lat., 18-20 » long.
Cell broadly (sometimes narrowly) flask-shaped, narrowed anteriorly
into a short, broad neck and reduced posteriorly to a thick stipe, 1.5-2
wide; protoplast oval to subglobose, with two laminate chloroplasts ;
7.6-8 w in diameter, 18-20 p» long.
On filamentous algae in a pool on beach of Crystal Lake, Vilas County.
This species differs in having a stipe which is much stouter and less
tapering than in the other described forms. It should be compared with
St. capense Prescott which has a very similarly shaped lorica but a very
slender, tapering stipe. Also compare with Derepyxis amphora Stokes
which it superficially resembles.
OPHIOCYTIACEAE
Ophiocytium desertum var. minor var. nov.
pL. 3, Fic. 15-16.
Cellulae coniunctae, cylindratae aut in forma farciminis figuratae, stipite basis
habente partem connexa, planatam similemque disco, polo anteriore late rotundato,
4-9 yw lat., 58-65 pw long.
Cells attached, cylindrical or sausage-shaped, basal stipe with a flat-
tened disc-like attaching organ, anterior end broadly rounded, 4—9 » in
diameter, 58-65 yp long.
In a roadside swamp near Boulder Junction, Vilas County.
This variety differs from the typical form only in its distinctly smaller
size.
Ophiocytium elongatum var. major var. nov.
PL. 3, FIG. 17.
Varietas ordinaria differens in mensura multo maiore; cellulae libere natantes,
cylindricae, rectae, curvae, aut uno polo truncato hamatae, altero spinam fortem,
non undulatam, habente; chloroplastes 16 disci parietales; cellula 10-12 yu lat.,
400-420 long., stipe (10-12 y, long.) non incluso.
A variety differing from the typical by its much greater size; cells
free-floating, cylindrical, straight, curved or hooked at one end which
is truncate, the other bearing a stout spine, not coiled; chloroplasts 16
parietal discs; cell 10-12 » in diameter, 400-500 » long without the
stipe which is 10-12 » long.
In a swamp near Rhinelander, Oneida County.
Prescott: New WIscoNnsIN ALGAE 363
This variety should be compared with O. cochleare (Eich.) A. Braun
which differs in its smaller size, arcuate or coiled cells. O. elongatum
W. & G. S. West, originally described from Burma, is sometimes coiled,
although not closely, and its dimensions are greater than for any of the
other species in this genus. Our specimens are mostly curved or bent
near one end and are not coiled.
B. CHRYSOPHYCEAE
MALLOMONADACEAE
Mallomonas pseudocoronata sp. nov.
pL. 3, Fic. 18.
Cellulae fusiformi-ellipticae, anguste rotundatae ambobus polis; squamae trans-
verse elliptico-rhomboidales aut scutulatae, non imbricatae, in seriebus spiralibus
transversisque positae; in polo anteriore formantes coronam proiecturarum acute
praefixarum circum aperturam flagelli; acus duorum geberum, alterae fortes longae-
que formantes floccum posteriorem, alterae breves spinosaeque, recurvatae super
superficiem; cellulae 20-25 p, lat., 48-50 p long., acibus inclusis.
Cells fusiform-elliptic, narrowly rounded at both poles; scales trans-
versely elliptic to rhomboidal or diamond-shaped, not imbricate, in spiral
transverse series; at the anterior end forming a corona of sharply pointed
projections about the flagellum opening, needles of two kinds, stout long
ones forming a posterior tuft, and short spine-like recurved ones over the
surface; cell 20-25 » in diameter, 48-50 » long including the needles.
Plankton in Lake McKenzie, Burnett County.
The collar or ‘crown’ at the anterior end which is formed of spine-like
scales, together with the shape and arrangement of the needles in the
posterior region differentiate this species.
Mallomonas urnaformis sp. nov.
PL. 3, FIG. 19.
Cellulae ellipticae et coronam collumve squamarum erectarum in polo anteriore
circum foramen flagelli habentes, squamis membrani rectangularibus in seriebus
transversis longitudinalibusque positis; squamae paucae immutataeque in polo pos-
teriore locatae; acus paucae, longae, tenuesque, passim divertentes, aequaliter sparsae;
cellulae 11-14 y, lat., 25-30 pw long., acibus non inclusis,
Cells elliptic with a corona or neck of erect scales at the anterior end
about the flagellum pore, scales of the membrane rectangular in transverse
and longitudinal series and with a few modified scales at the posterior end;
needles few, long and slender, diverging in all directions, evenly scattered;
cells 11-14 » in diameter, 25-30 p» long, without needles.
Tow from Boulder Lake, Vilas County.
The arrangement of the rectangular scales in regular transverse series
is unique in this species. ‘
RHIZOCHRYSIDACEAE
Lagynion reductum sp. nov.
PL. 3, Fic. 20-21.
364 FarLowia, Vor. 1, 1944
Lorica globoso-conica, fusco-fulva, habens, aperturam densatione anulari circum-
datam per quam extendit filus tenuis protoplasmatos; protoplastus globosus; chroma-
tophorus unus, in uno latere cellulae positus; 10 py alt., 11.5 w lat.
Lorica globose to conical, dark brown, with an opening surrounded by
an annular thickening through which a fine protoplasmic thread extends;
protoplast globose; chromatophore one, laying along one side of the
cell; 10 » high, 11.5 » in diameter.
Growing on filaments of Tribonema in a ditch near Rhinelander, Oneida
County.
This organism is remindful of Heterolagynion Oedogonii Pascher
(Ber. Deutsche Bot. Ges., 30: 157, figs. 1-4, 7-9. 1912) which is, how-
ever, colorless and much larger than our species.
Lagynion triangularis var. pyramidatum var. nov.
PL. 3, FIG. 22.
Lorica anguste pytamidata, marginibus inaequalibus aut subtriangularibus, fasti-
gata in collum breve longumve marginibus fere parallelis instructum; 10.8-12 u lat.,
15-20 yp, alt.
Lorica narrowly pyramidate with uneven margins, or subtriangular,
tapering to a short or long neck with nearly parallel margins; 10.8—12 p
wide, 15-20 w» high. ;
On filamentous algae in Cardinal Bog, Trout Lake, Vilas County.
This variety differs from the typical form in being narrower, propor-
tionately taller and in having a longer neck.
PHYLUM EUGLENOPHYTA
EUGLENACEAE
Euglena breviflagellum Prescott and Gojdics sp. nov.
PL. 3, FIG. 23-24,
Cellulae leviter metabolicae, elongato-fusiformes et spiraliter tortae curvataeve,
raro rectae ellipticae in transverso visae, nonnihil coartatae in parte anteriore et
truncatae, in parte posteriore coartatae lenius ita ut caudum longum forment; mem-
branum subtiliter spiraliterque striatum; flagellum breve, in mensura par sextae
parti longitudinis cellulae; paramyla corpora duarum varietatum, 6-8 laminae magnae
concavae alveataeve a latere dispositae et axi longo parallelae, pellicula leviter un-
dulata supra et aliis anulis discis similibus numerosis et parvis irregulariter per
cellulam sparsis; chloroplastes disci numerosi ovoideique regulariter per cellulam
dispositi; pyrenoides nullae; macula ocularis elliptica, formata ex granulis coccineis
irregulariter locatis; cellulae 120-145 uy long., 10-12 y lat.; corpora paramyla magna
18 w long.; grana paramyla parva 5 y, lat., 7 uw long.
Cells slightly metabolic, elongate-fusiform and spirally twisted or
curved, seldom straight, elliptic in cross section, rather abruptly narrowed
anteriorly and truncate, posteriorly narrowing more gradually to form
a long caudus; membrane finely and spirally striate; flagellum short,
about one-sixth of the length of the cell; paramylon bodies of two sorts,
6-8 large concave or trough-shaped plates laterally arranged and parallel
with the long axis with the pellicle slightly undulate over them, and other
Prescott: NEw WISCONSIN ALGAE 365
numerous and small disc-like rings irregularly scattered throughout the
cell; chloroplasts numerous ovoid discs evenly distributed throughout
the cell; pyrenoids lacking (?); eye-spot elliptic, composed of irregu-
larly arranged crimson granules; cells 120-145 w long, 10-12 » in
diameter; large paramylon bodies 18 » long; small paramylon grains
5 p» wide, 7 p» long.
Tychoplankton in Trilby Lake, Vilas County; also found in a pond
near Woods Hole, Massachusetts, August, 1935 (Gojdics).
The plate-like and laterally arranged paramylon bodies are distinctive.
The organism continually twists and turns in its movements and in so
doing causes the paramylon plates to telescope or overlap.
Euglena elastica sp. nov.
PL. 3, FIG. 25-27.
Cellulae maxime metabolicae et in statu natandi formam constanter mutantes, max-
ima ex parte fusiformes, saepe, autem, media in parte inflatissimae atque postice
anticeque abrupte coartatae, ad apices conice rotundatos paululum attenuatae; saepe,
autem, parte basali inflatissima nodiformique, in caudum numquam extensa; flagello
circa % longitudinem cellulae habente; chloroplastis. multis, irregulariter ovatis;
sine pyrenoidis (?); paramyli granis bacillis multis brevibusque, per cellulam quasi
aequaliter dispersis; stigmate irregulari atque parte in anteriore cellulae laterali;
cellulis 9.5-11 4 diam. (cellula videlicet extensa), 76-100 u long.
Cells highly metabolic and constantly changing shape when in motion,
mostly spindle-shaped but frequently much swollen in the midregion and
abruptly narrowed anteriorly and posteriorly, tapering slightly to coni-
cally rounded apices, but often with the basal end much swollen and
knob-like, never extended into a caudus; flagellum about 2% the length of
the cell; chloroplasts many, irregularly ovoid bodies; pyrenoids lack-
ing (?); paramylon bodies numerous short rods scattered rather evenly
throughout the cell; pigment-spot an irregularly shaped body, lateral in
the anterior end; cells 9.5-11 p» in diameter (when the cell is stretched
out), 76-100 » long.
In a small pool near Plum Creek at Sayner Fish Hatchery, Oneida
County.
Euglena minuta sp. nov.
PL. 3, FIG. 28-30. °
Cellulae maxime metabolicae, fusiformes ad subpyriformes; postice in cacumen
breve, obtusum, saepe curvatum productae; membrana levi (?); flagello %4 longi-
tudinem cellulae habente; chloroplasto uno, laminiformi, pyrenoidum habente; para-
myli granis bacillis multis parvisque aut granulis elongato-ovatis; cellula 5-6 wu
diam., 12-13.5 pw. long.
Cells highly metabolic, fusiform to somewhat pyriform; produced pos-
teriorly into a short, blunt, often curved tip; membrane smooth (?);
flagellum 34 the length of the cell; one plate-like chloroplast with a
pyrenoid; paramylon bodies many small rods or elongate-ovoid grains;
cells 5-6 » in diameter, 12-13.5 yp long.
Tychoplankton in Muskellunge Lake, Vilas County.
This is the smallest species of the genus recorded in North America.
The highly metabolic cells with a single chloroplast and small rod-like
366 Fartowla, VoL. 1, 1944
paramylon bodies make this organism different from other described
species.
Euglena oxyuris var. minor var. nov.
Bis S$, igs Sl,
Cellulae paulum metabolicae in statu natando formam constantem saepissime re-
tinentes, elongato-cylindricae tortaeque, postice ad caudem brevem efficiendam quasi
abrupte attenuatae; periplasto longitudinaliter striato; chloroplastis multis, disci-
formibus; paramyli granis 2 anulis magnis complanatisque, uno anteriore uno pos-
teriore quam nucleo centrali; cellulis 15-18 y. diam., 77-85 yp long.
Cells slightly metabolic, mostly keeping a constant shape in move-
ments, elongate-cylindric and twisted, tapering posteriorly rather abruptly
to form a short tail piece; periplast longitudinally striated; chloroplasts
numerous, disc-like; paramylon grains as two large, flattened rings, one
anterior and one posterior to the central nucleus; cells 15-18 p» in di-
ameter, 77-85 p long.
In shallow water of High Lake, Vilas County.
This variety differs from the typical form by its smaller size, the latter
being twice as large (30-45 y in diameter, 375-490 p long).
Phacus anacoelus yar. undulata fa. major fa. nov.
PL. 4, Fic. 1.
Cellulae late ovoideae, finientes in parte posteriore in caudo longo et oblique di-
recto; margines cellulae ruga alta in ambobus lateribus posita instructi quae duos
gibbos latos format; multo maior quam forma ordinaria; 64 p. lat., 111-115 p, long.
Cells broadly ovoid, ending posteriorly in a long, obliquely directed
caudus; margins of cell with a deep crease on either side forming two
broad bulges; much larger than in the typical form, 64 » in diameter,
111-115 pw long.
Plankton in Spider Lake, Vilas County.
This variety is very similar in size and shape to the typical form but is
differentiated by the prominent lateral creases or incisions which throw the
margin into broad undulations.
Phacus asymmetrica sp. nov.
PL. 3, FIG. 32.
Cellula irregulariter ovoideo-fusiformis et leviter spiralis in partibus anteriore et
posteriore; extenta in parte posteriore in caudum curvatum obtuseque praeacutum;
coartata in parte anteriore et habens duos gibbos inaequales in ambobus lateribus
apicis positos; periplastis subtiliter per longitudinem striata; corpora paramyla duo
anuli crassi torsique, plerumque per transversum cellulae locati; chloroplastes disci
ovoidei numerosi parvique; cellula 22-25 p. lat., 50-53 pw long.
Cell irregularly ovoid-fusiform and slightly spiral in the posterior and
anterior portions; extended posteriorly into a curved, bluntly pointed
caudus; narrowed anteriorly and with two irregular bulges on either side
of the apex; periplast finely striated longitudinally; paramylon bodies
as two thick twisted rings, usually lying transversely in the cell; chloro-
plasts numerous small ovoid discs; cell 22-25 p» in diameter, 50-53 p long.
In a roadside fosse near Shell Lake, Washburn County.
Prescott: NEw WIscoNsIN ALGAE 367
This species should be compared with P. Raciborski Drez. which is
‘much more slender and more nearly symmetrical. P. asymmetrica is ir-
regular in outline and in some instances seems more like an Euglena.
The cells are flattened, however, and show periplast features and para-
mylon grains which are like Phacus, especially in regard to the arrange-
ment of the latter. Besides the two large rings of paramylon there are
numerous smaller bodies which sometimes are densely packed in the cell.
Phacus Birgei sp. nov.
PL. 3, FIG. 33.
Cellula late ovalis, producta in parte posteriore ut formet caudum longum fastiga-
tumque qui obliquus axi longitidinali cellulae est, late rotundata in parte anteriore;
flagellum in longitudine cellulae par; periplast tenuissime striata; margines cellulae
acute incisae quattuor incisuris utrimque positis; corpora paramyla una magna et
numerosae parvae laminae circulares; chloroplastes multi disci ovoidei; macula ocu-
laris (?); cellulae 50-60 y», lat., 70-80 p. long.
Cell broadly oval, produced posteriorly to form a long tapering caudus
which is oblique to the longitudinal axis of the cell, broadly rounded
anteriorly; flagellum as long as the cell; periplast very finely striated;
margins of the cell sharply notched with four small indentations on either
side; paramylon bodies one large and numerous small circular plates;
chloroplasts many ovoid discs; eye-spot (?); cells 50-60 yw in diameter,
70-80 yp long. |
In a small pond near Genoa City, Walworth County. This species is
named for Dr. E. A. Birge.
The lateral creases, together with the shape and size of this species,
present a combination of characteristics which separate it from other
described Phacus.
Phacus chloroplastes sp. nov.
PL. 3, FIG. 34.
Cellulae late pyriformes; productae in parte posteriore ut caudum rectum vel
levissime deflexum forment; late rotundatae in parte anteriore cum papilla in medio
posita; periplastis per longitudinem striata; margo cellulae integer; chloroplastes
aliquot ligamenta parietalia quae parallela axi longo cellulae sunt; corpora paramyla
duo magni tenuesque anuli aut laminae leviter torsae quae per longitudinem cellulae
positae sunt; macula ocularis in medio regionis apicalis locata; cellula 20-22 u lat.,
29-31 y. long.
Cells broadly pyriform; produced posteriorly to form a straight or
very slightly deflected caudus; broadly rounded anteriorly with a median
papilla; periplast longitudinally striated; margin of the cell entire;
chloroplasts several parietal bands lying parallel with the long axis of
the cell; paramylon bodies two large, thin rings or slightly twisted plates
lying lengthwise in the cell; eye-spot median in the apical region; cell
20-22 p in diameter, 29-31 p long.
Plankton in an inlet to Trout Lake, Vilas County.
This species should be compared with P. hispidula (Eich.) Lemm. which
it resembles in general shape. The distinctive features of the Wisconsin
368 Fartowia, Vou. 1, 1944
species are the shape and the arrangement of the chloroplasts which are
quite unlike any of the other described members of the genus.
Phacus chloroplastes fa. incisa fa. nov.
PL. 4, FIG. 2-4,
Cellulae pyriformes vel napiformes in lineis extremis visae, fastigatae repente in
longum caudum rectum acutumque; periplastis per longitudinem striata; margo
cellulae instructus duabus incisuris utrimque locatis; cellulae 25-26 y, lat., 38-40 u.
long.
Cells pyriform or napiform in outline, tapering suddenly to a long,
straight, sharp caudus; periplast longitudinally striated; margin of the
cell with two sharp notches on either side; cells 25-26 » in diameter,
38-40 p. long.
In a cedar swamp, Lincoln County.
Phacus crenulata sp. nov.
' PL, 4, FIG. 5.
Cellula ovoideo-pyriformis, extenta in parte posteriore ut formet caudum leviter
fastigatum et acute praefixum; para anterior late rotundata sed bilobata cum papilla
convexa inter lobas posita; flagellum par cellulae in longitudine aut paulo longius;
margines cellulae distincte crenulati aut undulati; periplastis per longitudinem lineis
undulatis striata; corpora paramyla duo disci circulares; cellulae 14-15 y lat.,
34-36 yp long.
Cell ovoid-pyriform, posteriorly extended to form a gradually tapering,
sharp-pointed caudus, anterior end broadly rounded but bilobed with a
convex papilla between the lobes; flagellum as long as the cell or a little
longer; margins of the cell distinctly crenulate or undulate; periplast
longitudinally striated with undulating lines; paramylon bodies as two
circular discs; cells 14-15 » in diameter, 34-36 y long.
Plankton in a cedar swamp, Lincoln County.
This species should be compared with P. costata Conrad which has
spiral, entire striations rather than longitudinal wavy ones. The anterior
end is different in that species also. P. setosa var. crenata Skv. has spiral
striations.
Phacus pseudoswirenko sp. nov.
PL. 4, Fic. 8,
Cellulae orbiculares in lineis extremis visae, repente coartatae in parte posteriore
et productae ut forment caudum brevum acutumque qui ad sinistrum curvatur; pars
anterior late rotundata; flagellum in Jongitudinem corpori fere par; periplastis per
longitudinem striata, incisura laterali alta acutaque in medio lateris dextri posita
(rare una etiam in latere sinistro); corpus paramylon lamina magna circularisque;
cellulae 30-33 y lat., 37-40 » long.
Cells orbicular in outline, abruptly narrowed posteriorly and produced
to form a short, sharp caudus which curves to the left; anterior end broadly
rounded; flagellum about as long as the body; periplast longitudinally
striated, with a deep, sharp, lateral notch medianly located on the right
side (rarely one on the left side also); paramylon body a large circular
plate; cells 30-33 » in diameter, 37-40 p long.
Prescott: New WIScoNSsIN ALGAE 369
Plankton in a cedar swamp, Lincoln County, and in several ditches
and small ponds.
This species should be compared with P. Swirenko Skvortz., a form
which is about the same size but which has entire margins and a caudus
which turns to the right.
Phacus Segreti var. ovum var. nov.
PL. 4, FIG. 9.
Cellulae maiores quam in species ordinaria late ovoideae atque rotundatae in
partibus et anteriore et posteriore, caudo carentes; periplastis per longitudinem
striata; corpora paramyla duo disci anulares; cellulae 28-30 yu. lat., 39-41 p long.
Cells larger than in the typical form, broadly ovoid and rounded both
posteriorly and anteriorly, without a caudus; periplast longitudinally
striated; paramylon bodies two annular discs; cells 28-30 y in diameter,
39-41 p long.
In a swamp near Lauderdale Lakes, Walworth County.
Phacus spirogyra var. maxima var. nov.
PL. 4, FIG. 6-7.
Cellulae ovoideae vel quasi-oblongae, inaequaliter spirales aut tantummodo torsae,
repente coartatae in parte posteriore in longum caudum rectum curvumve; late ro-
tundatae in parte anteriore (anguste rotundatae a latere visae), cum gibbo claro in
medio posito; periplastis per longitudinem seriebus spiralibus granulorum gem-
meorum striata; chloroplastes disci numerosi; corpora paramyla duo anuli magni
ellipticique; cellulae 35-40 yp, lat., 70-80 p long.
Cells ovoid to somewhat oblong, unsymmetrically spiral or merely
twisted, abruptly narrowed posteriorly into a long straight or curved
caudus; broadly rounded anteriorly (narrowly rounded when seen from
the side), with a prominent median protrusion; periplast longitudinally
striated with spiral rows of pearly granules; chloroplasts numerous discs;
paramylon bodies two large doughnut-like rings; cells 35-40 » in di-
ameter, 70—80 p long.
In a roadside fosse near Shell Lake, Washburn County.
This variety is peculiar in the shape of the body which is not very
much flattened. In side view it is narrower, however, than when seen
in front view. The form of the paramylon bodies and the periplast deco-
rations are similar to those in Euglena spirogyra var. marchica. This
type of periplast decoration is uncommon in the genus Phacus. While
most specimens had but two large paramylon rings some individuals were
found with three. The cell is irregularly twisted in the anterior end.
This species should be compared with P. striata France.
Drezepolski’s description of P. spirogyra is not entirely complete and
since it has been impossible to see type specimens of that species, the
Wisconsin forms are assigned here with a question.
370 FarLowia, Vou. 1, 1944
Lepocinclis glabra fa. minor fa. nov.
PL, 4, Fic. 10.
Cellulae late ovoideae, caudo brevi papillaeque simili instructae, late rotundatae
in parte anteriore et finientes in gibbo bipapillato per quem emergit flagellum; flagel-
lum in longitudine fere corpori par; periplastis levis; paramylon ex duobus liga-
mentis semicircularibus compositum, curvatis in transverso in peripheria cellulae,
uno utrimque locato; chloroplastes numerosi disci ovales; cellulae 14-16 y, Iat.,
20-22 w» long., minores quam in forma ordinaria.
Cells broadly ovoid, with a short papilla-like caudus, broadly rounded
anteriorly and ending in a bi-papillate protrusion through which the
flagellum emerges; flagellum about as long as the body; periplast smooth;
paramylon in the form of two semicircular bands, curving transversely
at the periphery of the cell, one on either side; chloroplasts numerous,
ovoid discs; cells 14-16 » in diameter, 20-22 » long, smaller than the
typical form.
Tow sample from a cedar swamp in Lincoln County.
Trachelomonas hexangulata var. repanda var. nov.
PL. 4, FIc. 11.
Distans a specie ordinaria in marginibus lateralibus loricae convexioribus et mar-
ginibus lateralibus partis posterioris concavioribus adeo ut apiculatio obtusa in parte
posteriore proferatur; lorica 14.4-15 uw, lat., 36-38 » long.
Differing from the typical form in having the lateral margins of the test
more convex and the posterior lateral margins more concave so that a blunt
apiculation is produced posteriorly; test 14.4-15 » in diameter, 36-38
long.
In tow samples from a cedar swamp in Lincoln County.
Trachelomonas mammillosa sp. nov.
PL. 4, Fic. 12.
Testa spherica; ore flagelli in inflatione mammillata, per costam cincta crassa,
anularemque, quae extremitatem, anteriorem circumplectitur; membrana levi, sub-
flava; testa 24-26 py, diam.
Test spherical; flagellum aperture in a mammillate swelling which
is surrounded by a thick annular ridge encircling the anterior end; wall
smooth, yellowish; test 24-26 » in diameter.
Tychoplankton in Stevens Swamp near Trout Lake, Vilas County.
This species is distinguished by the prominent ridge which encircles
the test anteriorly, and the papillate swelling about the flagellum aperture.
It should be compared with T. peridiniformis Skv. reported from Man-
churia.
Trachelomonas superba yar. spinosa yar. nov.
PL. 4, Fic. 13.
Lorica late ovalis; membranum spinosum, parte posteriore spinis multo longioribus
et acutius praefixis cincta; apertura flagelli torque carens sed coronula spinarum
acutarum erectarumque circumdata; lorica 32.5-36 yp. lat., 39-48 wu long.
Test broadly oval; wall spiny, with the posterior part beset with much
longer and more sharply pointed spines; flagellum aperture without a
Prescott: New WIsconsInN ALGAE Ryall
sollar but encircled with a coronula of sharp, erect spines; test 32.5-36 p
in diameter, 39-48 yp long.
Plankton from Muskellunge Lake, Vilas County.
This variety differs from the typical form by the longer spines at the pos-
terior pole and the coronula about the flagellum aperture. Also the size is
greater in the variety.
PHYLUM PYRROPHYTA
GYMNODINIALES
GYMNODINIACEAE
Gymnodinium caudatum sp. nov.
PL. 4, Fic. 14-16.
Cellulae magnae, ovoideae aut inverse conicae aut similes turbini in forma, late
rotundatae in polo anteriore, coartatae et productae in caudum curvum in polo pos-
teriore; plurimum complanatae in parte dorso-ventrali; sulcus transversus clarus
in medio positus, spiraliter ad sinistrum versus; chromatophori numerosae et luteo-
fulvae laminae, ovoideae elongataeve, radialiter dispositae; macula ocularis prope
sulcum in hypocono posita; sulcus per dimidiam longitudinem hypoconi extendens
et paulum in epiconum; cellula 65-70 y, lat., 104-118 y, long.
Cells large, ovoid to inversely conical or top-shaped, broadly rounded
at the anterior end, narrowed and produced into a curved caudus at the
posterior pole; very much flattened dorso-ventrally; transverse furrow
prominent and median, spirally turned to the left; chromatophores numer-
ous golden-brown, ovoid or elongate plates, radially disposed; eye-spot
present near the sulcus in the hypocone; longitudinal furrow extending
about half the length of the hypocone and for a short distance into the
epicone; cell 65—70 yp in diameter, 104-118 p long.
In a Sphagnum bog near Rhinelander, Oneida County.
The large size of this species and the narrowed hypocone which forms
a distinct caudus posteriorly are its distinguishing features.
PHYLUM CYANOPHYTA
CHROOCOCCALES
CHROOCOCCACEAE
Glaucocystis duplex sp. nov.
pL. 4, FIG. 17-19.
Colonia 8-16 cellularum sphaericarum membrano sphaerico cellulae matricalis
multo dilatato circumdata; chromatophori duae massae stellatae habentes corpora
vermiformia et caeruleo-viridia ex duobus locis separatis extendentia; cellulae 40-44 y.
lat., colonia 150-170 y, lat.
Colony of 8-16 spherical cells inclosed by a much enlarged spherical
mother-cell wall; chromatophores two stellate masses with vermiform,
blue-green bodies radiating from two separate points; cells 40-44 » in
diameter, colony 150-170 p in diameter.
Among dense clots of algae in Manitowish River, Vilas County.
/
Sie Fartowl!A, Vou. 1, 1944 .
This species differs from G. nostochinearum Itz. in the spherical shape
of the cell and in the dual arrangement of the chromatophores. From
G. cingulata Bohlin it differs in the morphology of the cell wall. That
species is spherical but the wall has a median annular thickening. The
chromatophores are numerous and parietal. Some forms of G. nostochi-
nearum Itz, are described as spherical but other features of the Wisconsin
plants seem to warrant separating them from that species.
Glaucocystis oocystiformis sp. nov.
PL. 4, FIG. 20.
Cellulae solitariae (aut in coloniis positae ?) late ellipticae, densationibus glandu-
losis membrani in polis instructae; chromatophori numerosi, pulvinaribus in _peri-
pheria cellulae circum vacuolam centralem, sphaericum, non coloratam positis (?) ;
cellulae 20-27.3 y, lat., 40-45 p long.
Cells solitary (or in colonies ?) broadly elliptic, with nodular thicken-
ings of the cell wall at the poles; chromatophores numerous, irregular
pads at the periphery of the cell about a central, spherical, colorless
vacuole (?); cell 20-27.3 » in diameter, 40-45 yp long.
Plankton from an inlet to Trout Lake, Vilas County.
This species differs from the others in the shape of the cell, the form
of the chromatophores, and in its possession of polar nodules. Whether
the absence of colonial association, or the retention of autospores within
the mother-cell wall is a constant feature is undetermined, but in all cases
observed the cells were solitary. In the developmental stages of G. nos-
tochinearum Itz., as described by Hieronymus (1892), the chromatophores
show much the same character as exhibited in G. oocystiformis. Since
no other type of chromatophore expression or arrangement was found in
the Wisconsin plants there is no indication that the forms in our collec-
tions are but a developmental stage of some other species.
‘
HORMOGONALES
OSCILLATORIACEAE
Lyngbya latissima sp. nov.
PL. 4, FIG. 21.
Plantae solitariae, in aliis algis impeditae; trichomata recta, in apicibus non fasti-
gata; cellulae discis similes materiam tenuiter aequaliterque granosam habentes,
37-40 p lat, 3.7-7.4 uw long.; vagina dense (3.7-5 u.), lamellata, stratis externis
rugosis et senectute asperatis; filamentum 44-58 y, lat.
_ Plants solitary, entangled among other algae; trichomes straight, not
tapering at the apices; cells disc-like with contents finely and evenly
granular, 37-40.7 » in diameter, 3.7—7.4 long; sheath thick (3.7-5 »),
lamellate, with outer layers wrinkled and roughened in age; filament
44-58 p in diameter.
In Sweeney Lake, Vilas County; Little Clam Lake, Burnett County. .
This very large species should be compared with Oscillatoria princeps
Vauch. when the latter is in the hormogonial condition and inclosed by
Prescott: New WIsconsin ALGAE 373
a sheath. In such a case the sheath is not thick and lamellated as in
L. latissima, nor are the trichomes so long (usually). L. gigantea de-
scribed by Lewis, Zirkle, and Patrick (1933) and referred by Drouet
(1938) to Oscillatoria princeps Vauch., differs in having thin, smooth
sheaths characteristic of the hormogonial phase of the latter species. Un-
like Oscillatoria princeps, our plants are not at all tapering at the apices
and the contents of the cell are not coarsely granular.
L. latissima should be compared with L. Hummelii Borge, a large species
which is, however, smaller than the former, and one which has longer
cells. Also the sheath in Borge’s species is thinner and apparently not
lamellate as in the Wisconsin plants.
NOSTOCACEAE
Anabaena wisconsinensis sp. nov.
PL. 2, Fic. 9-10.
Trichomata planctonica, recta aut subflexuosa, solitaria aut (saepius) ad fasciculos
lamelliformes parvos laxosque efficiendos parallele aggregata, sine vagina, ad apices
non attenuata; cellulis quadratis vel cylindratis, ad dissaepimenta constrictis, 3.6—
4» diam., 3.6-10.8 » long., intus pseudovacuolas magnas praedentibus; heterocysti-
bus sphaericis aut compresso-globosis, 3.6—4.2 4 diam., una sola singulo in trichomate
partem mediam occupante; gonidiis elliptico-ovatis ad late ovatis, ab heterocystibus
remotis, primum constantibus ex 3 cellulis vegetativis dilatatis seriatim dispositis,
quarum una sola plerumque maturescit ut omnia trichomata unicum gonidium habeant,
si plura autem, gonidia, bina sunt, 7.2-8 » diam., 10-13 yw long.
Trichomes planktonic, straight or slightly flexuous, solitary or (more
often) aggregated in parallel fashion to form small, loose, flake-like bun-
dles, without a sheath, not tapering at the apices; cells quadrate to cylin-
drical, constricted at the cross walls, 3.6-4 » in diameter, 3.6-10.8 yp
long; cell contents with large pseudovacuoles; heterocysts spherical or
compressed-globose, 3.6-4.2 » in diameter, only one in each trichome,
centrally located; gonidia elliptic-ovoid to broadly ovoid, remote from
the heterocysts, beginning their development as a series of three enlarged
vegetative cells, usually only one of which matures so that each trichome
has but a single gonidium, when more than one, occurring in pairs, 7.2—-8 p,
in diameter, 10-13 » long.
Plankton in Post Lake, Langlade County.
This plant is remindful of Aphanizomenon flos-aquae (L.) Ralfs. It
should be compared with Anabaena aphanizomenoides Forti from which
it differs in the smaller size throughout, the ovoid shape of the gonidia
and their location remote from the heterocysts rather than adjacent to
them as in that species.
ALBION COLLEGE
ALBION, MicH.
374 FarLowIia, VoL. 1, 1944
LITERATURE
Allegre, Charles & T. L. Jahn. (1943). A survey of the genus Phacus Dujardin
(Protozoa) ; Euglenoidina. Trans. Amer. Micros. Soc. 62 (3): 233-244.
Bigeard, E. (1935-1936). Les Pediastrum d’Europe. Etude biologique et sys-
tematique. Rev. Algol. 7: 1-94; 327-418.
Conrad, W. (1933). Revision du genre Mallomonas Perty (1851) incl. Pseudo-
mallomonas Chodat (1920). Mem. Mus. Roy. d’Hist. Nat. Belgique, Mem. No.
56: 1-82.
. (1934). Materiaux pour une monographie du genre Lepocinclis Perty.
Arch, Protist. 82 (2): 203-249.
. (1935). Etude systematique du genre Lepocinclis Perty. Mem. Mus. Roy.
d’Hist. Nat. Belgique, 2nd Ser. 1: 1-84.
Deflandre, G. (1926). Monographie du genre Trachelomonas. Nemours Edition.
Drezepolski, R. (1925). Przycznek do zanjomosci polskich Euglenin. Kosmos,
Bull. Soc. Pol. Nat. Leopol. 50: 173-270.
Drouet, F. (1938). The Oscillatoriaceae of Southern Massachusetts. Rhodora
40: 221-241; 255-273.
Hazen, Tracy E. (1902). The Ulotrichaceae and Chaetophoraceae of the United
States. Mem. Torrey Bot. Club 11: 135-250.
Hieronymus, G. (1892). Beitrage zur Morphologie und Biologie der Algen. I.
Glaucocystis nostochinearum Itz. Cohn’s Beitr. Biol. Pflanzen 5: 461-495.
Lewis, I. F., C. Zirkle & Ruth Patrick. (1933). Algae of Charlottesville and
vicinity. Jour, Elisha Mitchell Sci. Soc. 48: 207-222.
Prescott, G. W. & Hannah T. Croasdale. (1937). New or noteworthy fresh
water algae of Massachusetts. Trans. Amer. Microsc. Soc, 56: 269-282.
Skvortzow, B. W. (1925). Zur Kenntnis der Mandschurischen Flagellaten.
Beih. Bot. Centralbl. 41: 311-315.
(1925). Die Euglenaceengattung Trachelomonas Ehrenberg. Sungari
Station zu Harbin der Gesell. zur Erforschung der Mandschurei 1 (2): 1-101.
(1928). Die Euglenaceengattung Phacus Dujardin. Ber. Deutsche Bot. .
Ges. 46: 105-125.
Smith, G. M. (1916). New or interesting algae from the lakes of Wisconsin.
Bull. Torrey Bot. Club 43: 471-483.
(1916). A preliminary list of algae found in Wisconsin lakes. Trans.
Wis. Acad. Sci. Arts and Letters 18: 531-565.
(1918). A second list of algae found in Wisconsin lakes. Trans. Wis.
Acad. Sci. Arts and Letters 19: 614-654.
(1920). Phytoplankton of the inland lakes of Wisconsin, Part I. Bull.
Wis. Geol. and Nat. Hist. Surv. 57.
(1924). Ibid. Part II. Ibid.
& F. D. Klyver. (1929). Draparnaldiopsis, a new member of the algal
family Chaetophoraceae. Trans. Amer. Microsc. Soc. 48: 196-203.
Swirenko, D. (1914). Zur Kenntnis der russischen Algenflora. I. Die Eugle-
naceengattung Trachelomonas. Arch. Hydrobiol. Plank. 9: 630-647.
Tiffany, L. H. (1937). Oedogoniales. North American Flora 11: Part 1.
Transeau, E. N. (1914). New species of green algae. Amer. Jour. Bot. 1: 289-301.
(1915). Notes on the Zygnemales. Ohio Jour. Sci. 16: 17-31.
West, W. & G. S. West. (1907). Freshwater algae from Burma, including a few
from Bengal and Madras. Annals Roy. Bot. Garden Calcutta 6 (2): 175-260.
snes
376
FarLow1a, VoL. 1, 1944
EXPLANATION OF PLATE 1
. Chlamydomonas polypyrenoideum sp. nov. 650.
. Tetraspora lamellosa sp. nov. > 500.
. Schizochlamys compacta sp. nov. > 600.
. Ulothrix cylindricum sp. nov. x 500.
. Geminella crenulatocollis sp. nov. X< 650.
. Hormidiopsis ellipsoideum sp. nov. >< 650.
. Pediastrum quadricornutum sp. nov. X 650.
. Oocystis pyriformis sp. nov. > 500.
. Tetraédron asymmetricum sp. nov. > 1500.
. Tetraédron bifurcatum var. minor var. nov. >< 650.
. Tetraédron cruciatum var. reductum var. nov. > 650.
. Tetraédron enorme var. pentaédricum var. nov. 650.
. Tetraédron limneticum var. gracile var, nov. >< 650.
377
Prescott: NEw WIsconsin ALGAE
Cet
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aa i : Zp pe ieee
Tees A Pe FAD
5 Af, ea “ Ge ::
ees eet / és H #
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Pate 1
378 Fartowia, VoL. 1, 1944
EXPLANATION OF PLATE 2
Fig. 1-3. Stigeoclonium pachydermum sp. nov.; 1, * 250; 2, * 500; 3, x 1000.
4-5. Draparnaldia Judayi sp. nov. 600.
6. Pediastrum biradiatum var. emarginatum fa. convexum fa. nov. < 600.
7. Pediastrum muticum var. crenulatum var. nov. > 700.
8. Tetraédron lobulatum var. crassum var. nov. X 1000.
9-10. Anabaena wisconsinensis sp. nov. >< 500.
Prescott: NEw WIscoNsIN ALGAE 379
380 Fartowia, VoL. 1, 1944,
EXPLANATION OF PLATE 3
Fig. 1. Tetraédron regulare var. granulata var. nov. 500.
2. Tetraédron regulare var. incus fa. major fa. nov. 500.
3. Scenedesmus opoliensis var. contacta var. nov. >< 850.
4-5. Tetradesmus Smithii sp. nov.; 4, < 400; 5, < 1000.
6-10. Stipitococcus apiculatus sp. nov.; 6, X 700; 7-10, x 2000.
11-14, Stipitecoccus crassistipatus sp. nov. >< 1000.
15-16. Ophiocytium desertum var. minor var. nov. >< 500.
17. Ophiocytium elongatum var. major var. nov. X 500.
18. Mallomonas pseudocoronata sp. nov. X 500.
19. Mallomonas urnaformis sp. nov. X 1000.
20-21. Lagynion reductum sp. nov. > 500.
22, Lagynion triangularis var. pyramidatum var. nov. X 650.
23-24. Euglena breviflagellum Prescott and Gojdics, sp. nov. x 650.
25-27. Euglena elastica sp. nov. X 500.
28-30. Euglena minuta sp. nov. x 500.
31, Euglena oxyuris var. minor var. nov. < 500.
32. Phacus asymmetrica sp. nov. X 600.
33. Phacus Birgei sp. nov. >< 500.
34. Phacus chloroplastes. x 600.
Prescott: New WISCONSIN ALGAE 381
Piate 3
382
FarLowla, VoL. 1, 1944
EXPLANATION OF PLATE 4
Phacus anacoelus var. undulata fa. major fa. nov. < 600.
. P. chloroplastes fa. incisa fa. nov. >< 600.
. Phacus crenulata sp. nov. > 600.
. Phacus spirogyra var. maxima var. nov. 500.
. Phacus pseudoswirenko sp. nov. X 600.
. Phacus Segreti var. ovum var. nov. >< 600.
. Lepocinclis glabra fa. minor fa. nov. 550.
. Trachelomonas hexangulata var. repanda var. nov. X 550.
- Trachelomonas mammillosa sp. nov. X 550.
. Trachelomonas superba var. spinosa var. nov. X< 750.
. Gymnodinium caudatum sp. nov. 550.
. Glaucocystis duplex sp. nov.; 17, X 250; 18, 19, & 350.
. Glaucocystis oocystiforme sp. nov. 650.
. Lyngbya latissima sp. nov. 375.
383
Prescott: NEw WIsconsin ALGAE
PLATE 4
384 Fartowia, Vou. 1, 1944
EXPLANATION OF PLATE 5
Fig. 1-3. Oedogonium Kjellmanii var. granulosa var. nov.; 1, 2, < 275; 3, « 550.
4-5. Oedogonium oviforme fa. gracile. < 600.
6-7. Oedogonium Smithii sp. nov.; 6, 500; 7, « 600.
8. Oedogonium microgonium sp. nov. X 600.
9. Oedogonium spheroideum sp. nov. X< 300.
10. Oedogonium Sawyeri sp. nov. >< 550.
1l. Oedogonium Kozminskii sp. nov. < 360.
12. Oedogonium polyandrium sp. nov. 550.
13-14. Oedogonium oelandicum var. contortum var. nov. X 600.
tm ip <
2. ay
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1(3): 387-389 FARLOWIA January, 1944
MICROAPLANOSPORES OF VAUCHERIA
GILBERT M. SMITH
The asexual reproduction of Vaucheria by means of multiflagellate
zoospores produced singly within sporangia is well known to all. So, °
also, is the production of an aplanospore instead of a zoospore within a
sporangium. To judge from the literature, the transverse segmentation
of the contents of a filament to form a uniseriate row of thick-walled
aplanospores (hypnospores) is of relatively infrequent occurrence, and
is found only in soil-inhabiting species of Vaucheria. Stahl (1879), the
first to describe these hypnospores in detail, holds that their formation
is induced by a drying out of the substratum. However, terrestrial species
growing in the vicinity of Stanford University show that production of
hypnospores is not always correlated with drying out of the soil. In
central California the terrestrial species form hypnospores during the
winter rainy season when the soil is saturated with water, but only when
the minimal temperatures are near the freezing point. Dangeard (1940)
has recently stated that the siphonaceous terrestrial algae producing
hypnospores are not members of the genus Vaucheria. This conclusion
is based upon his inability to obtain sex organs when the algae were
grown in culture. Dangeard’s studies raise the question as to whether
or not spores may be formed in vegetative portions of filaments of
V aucheria.
Specimens of Vaucheria Gardneri Collins collected in a seepage pool
from Lagunita, an artificial lake on the Stanford campus, show that
spores may be formed in vegetative portions of a filament, and reveal
a type of spore not previously described for the genus. These spores
were present in many of the filaments collected on April 15, 1943. The
station was revisited on April 18 to collect additional material, but in
the meantime most of the algae had been killed by oil sprayed on the
pool to kill mosquito larvae. Because of this it was impossible to obtain
material showing the manner in which the spores are formed or their mode
of germination after they are ripe.
Approximately 75% of the filaments in the collection bore sex organs,
the remainder did not. Of the latter more than half of them contained
small spores throughout their entire length. These filaments resemble
fruiting specimens of Phyllosiphon and at first it was thought that they
should be considered an aquatic species of that genus. Eventually the
spores were also found in filaments with sex organs and thus definitely
establishing that the vegetative filaments with spores were those of
V. Gardneri.
The type of spore, previously unrecorded for Vaucheria, is an aplano-
spore and the term microaplanospore is proposed to distinguish it from
the large aplanospore formed singly within a sporangium. Microaplano-
spores are formed throughout the length of a filament, not in sporangia
387
388 FartowiA, VoL. 1, 1944
separated from the remainder of a filament by a transverse wall. All of
the protoplasm of a filament divides to form microaplanospores and there
are no remnants of cytoplasm. between them. The distribution of the
spores within a filament is uneven; they are densely packed in certain
portions of a filament, in other portions they lie a short distance from
one another (Fig. 1). This uneven distribution is thought to be an
artifact caused by manipulation of the filaments in mounting them for
examination under the microscope.
Fig. 1. Portion of a filament of Vawcheria Gardneri containing microaplanospores.
xX 325. Fig. 2. Microaplanospores. x 1300.
The microaplanospores are spherical, 5.5-6.5 in diameter, and have
a wall somewhat thinner than the wall of the cell containing them. The
protoplast contains a single irregularly lobed chloroplast and one or two
droplets of oil adjacent to the chloroplast (Fig. 2). Nuclei were not
observed, but the small size of the spores suggests that they are uninuc-
leate. There are no pores or other openings in walls of filaments con-
taining microaplanospores. Thus it is obvious that liberation of spores
is by disintegration or decay of the parent cell wall. For reasons given
above material was not available for study of germination of micro-
aplanospores.
The Phyllosiphonaceae, a family universally recognized by phycologists,
differ from all other families of the Siphonales in that they produce
many small aplanospores within vegetative portions of filaments. The
discovery of microaplanospores in Vaucheria Gardneri shows that pro-
duction of many small aplanospores within vegetative portions of fila-
ments is a feature common to Phyllosiphonaceae and to at least one mem-
ber of the Vaucheriaceae. This might be taken as a justification for
uniting the two families, but such a procedure seems unwarranted when
SMITH: MICROAPLANOSPORES OF VAUCHERIA 389
sexual reproduction is taken into consideration. All known members
of the Vaucheriaceae regularly form antheridia and odgonia; whereas
no antheridia, odgonia, or any other kind of sex organ has ever been
observed in any of the Phyllosiphonaceae. Thus, pending discovery of
sexual reproduction in Phyllosiphon and related genera, it is better to
retain the family Phyllosiphonaceae.
STANFORD UNIVERSITY
CALIFORNIA
BIBLIOGRAPHY
Dangeard, P. (1940). Sur la prétendue reproduction des Vaucheria par des acinétes
et des spores amiboides et sur la nouveau genre Asterosiphon. Compt. Rend.
Acad. Sci. Paris 210: 719-721.
Stahl, E. (1879). Ueber die Ruhezustaénde der Vaucheria geminata. Bot. Zeitg.
Sa? 1oo-137e lepl
1(3) : 391-393 FARLOWIA January, 1944
A NEGLECTED FIELD OF STUDY WITH THE DESCRIPTION
OF A NEW LICHEN FROM CALIFORNIA
ALBERT W. C. T. HERRE
I suggest that students of fresh water and aerial algae take up seriously
the study of certain life forms that in this country have been greatly neg-
lected. I refer to those algae found in lichens, whether they are one of
the two essential lichen components or merely play a secondary part. Ap-
parently no one in this country is working upon them with a view to
finding out the changes they incur when enslaved by a fungus, as in
ordinary lichens, or the way they are affected when more or less dominant
as in the gelatinous lichens or other groups where the alga gives form
and character.
European botanists long ago isolated the algae of some lichens and
determined them with more or less exactitude. At various times pure
cultures of different lichen algae and fungi have been made; sometimes
these have later been synthesized and lichens grown until fruiting con-
dition was reached. There is ample room for more such studies, but
studies of a different character are likewise needed.
There are two other lines of research waiting for algologists to take
up in connection with lichens. First, little or nothing has been done in
this country upon the ecological and physiological changes or adapta-
tions (and their resultant effect upon morphology) which an alga must
sometimes undergo if it is to flourish or even survive as a prime factor
in making what we call a lichen. That such changes do occur and are
relatively common is well known to the few who have examined large
numbers of lichens under the microscope. That they are quite unknown
to the students of algae is not generally appreciated. For example, algol-
ogists are familiar with Trentepohlia and its orange red color, but they
are not generally aware that in lichens containing this alga the orange
red disappears and the alga is green. The student of lichens must there-
fore recognize the presence of Trentepohlia by other criteria than color.
Nostoc, when growing in lichens, apparently never has heterocysts; in
some lichens it forms characteristic chains, but in others it becomes com-
pacted into masses very different from ordinary Nostoc, while in still
others it breaks up into very small groups which can hardly be called
chains. Scytonema, too, breaks up into tiny groups or solitary cells,
and does not at all resemble the pictures in the books. Thirty-six years
ago my friend and teacher, Dr. Alexander Zahlbruckner, called my at-
tention to the fact that Rivulariaceae (Calothrix and Rivularia) lost their
basal heterocysts when growing in lichens, and were therefore hard
to recognize.
Gloeocapsa is a common and well known free living alga, found in
stagnant water and damp places. Algologists say the cells are always
globose, more or less, and are never elongate or develop rhizoids. In
391
392 Fartowia, Voi. 1, 1944
the new species of lichen described later, the alga is Gloeocapsa, section
Xanthocapsa. In the interior of the lichen the algal cells are from 6
to 10 » in diameter, arranged in groups 15 to 25 » in diameter, surrounded
by a thick gelatinous sheath. Near the under surface of the lichen the
Xanthocapsa groups are much larger, and just within the surface some
of the cells are seen to be somewhat elongate, instead of globose. From
this stage they become two or three times as long as broad; a septum
develops and the tip elongates still more until it projects beyond the
lower margin of the lichen. This tip, which may or may not develop
further septa, continues to lengthen until it becomes a very elongate,
slender, and tubular rhizoid. The basal portion is the dark blue-green
of the typical Gloeocapsa or Xanthocapsa color, with its characteristic
gelatinous coat, but the burrowing tip soon becomes colorless in its outer
part. The whole under surface of the lichen is covered with these rhizoids,
which may be 5 mm. or more in length and form a dense mat that binds
together the loose grains of the desert surface and holds the thallus firmly
in position. In all other crustaceous lichens known to me the rhizoids
are hyphal (fungal) threads only. I have no doubt that this is merely
one instance of many that would give algologists new light upon the
adaptions and physiology of algae.
Second, algologists should critically examine the algal constituents of
various lichens and establish their correct identity. This is particularly
needed in the case of those lichens containing blue-green algae. Many,
if not most, of the published records have merely been copied from one
author to another, without a thorough comparative examination of genera
and authentic species, a practice of which I have myself been guilty.
Recent studies of large numbers of lichens sent from all over the United
States, for identification, often reveal that lichens said to have Nostoc
as their alga do not contain Nostoc at all. This is particularly true of
members of the Pannariaceae, in which Scytonema or Xanthocapsa may
be mistaken for Nostoc. For example, in Pannaria lepidiota, distrib-
uted by Cummings and Seymour as No. 122, Decades of North American
Lichens, from Mt. Tamalpais, California, the alga is Xanthocapsa, not
Nostoc as given in all published works.
In a number of lichens containing Gloeocapsa (Xanthocapsa) or
Seytonema, I have also found Aphanocapsa, a blue-green alga not hitherto
reported as occurring in any lichens. Whether it is an intrusive in the
same way that Nostoc enters the hepatic Anthoceros and various foliose
lichens, whether it is the sole algal component of a lichen, or whether it
shares that onus with Yanthocapsa or Scytonema, is another problem for
the algologist to solve. To conclude, our knowledge of the algal con-
stituent of several families of lichens is but fragmentary, or even er-
roneous, and needs to be investigated by algologists who should settle
their systematics and explore their hidden ecology.
All this is preliminary to the description of a new desert lichen from
Herre: New Lichen From CALIFORNIA 393
California. Various workers, especially Dr. H. E. Hasse, have collected
and studied the earth lichens of California deserts, and it was assumed
that they were all known. The discovery of one belonging to a genus
hitherto unknown in this country is therefore of some interest.
Anema Dodgei Herre sp. nov. (Pyrenopsidaceae)
Thallus monophyllus, squamulosus, pseudoparenchymaticus, crassus, salebrosus,
inequalis, usque ad 5 mm. latitudine, substratum arcte obducens, late rhizino-affixus,
niger, aut virescenti-niger vel opace olivaceo-umbrinus rigidus, cartilagineus, made-
factus turgidus, subgelatinosus; gonidia Xanthocapsa sunt.
Apothecia parva vel mediocria, primum punctiformia, demum dilatata et urceolata
demum lecanorina, margine thallino prominulo, usque ad 1.25 mm., lata rufa vel
fusco-rufa; asci cylindrici; sporae octonae, monostichae, simplices, late ovoideae
10-22 y. longae, 5.5-11 yw crassae. Spermatia parva, bacilliformia aut subellipsoidea.
Ad terram desertorum, areas usque ad 8 cm. in diametro formans. Riverside and
San Bernardino Counties, California.
Thallus monophyllous, thick, the squamules unequal in size, reaching
a width of 5 mm. and forming a regular crust with rough irregular surface,
pseudoparenchymatous within and closely adherent to the substratum by
numerous rhizoids which reach a length of 5 mm. or more. Color, black
or greenish black to dull olive brown; when dry the thallus is rigid and
cartilaginous, becoming turgid and subgelatinous when wet. The alga
is Xanthocapsa, with cells 6-10 » in diameter, forming groups 15-25 p
in diameter, and surrounded by a thick gelatinous sheath.
The apothecia, reaching a width of 1.25 mm., are punctiform at first,
becoming open and urceolate to lecanorine, with prominent thalline
margin, the disk red to dark red; the epithecium is yellow, the thecium
colorless, about 125 p wide, the hypothecium gray or dusky gray, about
150 » wide; the asci are cylindrical; the paraphyses jointed, coherent.
Spores simple, colorless, 8, in a single row, broadly ovoid, 5.5-11 yw by
10-22 ». Spermatia small, bacilliform or slightly ellipsoid.
Apparently an abundant earth lichen in the deserts of Riverside and
San Bernardino Counties, California, forming crustose patches which
may reach a width of 8 centimeters, more or less. Collected by Francis
Drouet and J. Francis Macbride of the Field Museum, in 1941; No. 4621,
type material, on barren soil a mile southwest of Essex, San Bernardino
County, altitude about 1000 feet; Nos. 4624, 4626, and 4627, co-type
material, on open places in the desert about 2 miles east of Danby, San
Bernardino County; No. 4711, co-type material, on barren ground in
open scrub forest just west of Desert Center, Riverside County; altitude
about 900 feet.
Named for Dr. Carroll W. Dodge, of the Missouri Botanical Garden,
in recognition of his contributions to the study of lichens.
STANFORD UNIVERSITY
CALIFORNIA
ia. 22. 7 Te. 3;
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32 ~ 7
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S a “odes
1(3) : 395-467 FARLOWIA January, 1944
MARINE FUNGI: THEIR TAXONOMY AND BIOLOGY ‘
E. S. BaRGHooRN AND D. H. LINDER
INTRODUCTION
Although thousands of species of fungi are known from what may be
considered terrestrial habitats, and a much lesser number from freshwater
habitats, very few have been described from marine environments, al-
though a number of the latter do exist. Cotton (1), in 1907, in summar-
izing the situation, noted that fifteen species had been described as marine
forms, and to these he added a sixteenth. Of these species, six were mem-
bers of the phycomycetous order Chytridiales and ten of the ascomycetous
sub-class Pyrenomycetes. Since Cotton’s paper, some of the species have
been relegated to synonymy, but in 1915 Sutherland (14, 15, 16) de-
scribed fourteen new species of Pyrenomycetes, two of which were the
types of new genera. A year later, he (17) added a number of new species
of Fungi Imperfecti. Petersen (6) made a study of the chytridiaceous
forms parasitic on algae, and more recently Sparrow (11, 12, 13) has
added to our knowledge of that group. The only marine fungi that have
been published to date as occurring on plant remains, other than algae, are
three species of Ophiobolus described by Ellis (2), Saccardo (10), and
Mounce and Diehl (5). The last named authors described Ophiobolus
halimus which apparently caused considerable damage to the eelgrass,
Zostera marina. Subsequently Renn (7, 8, 9) demonstrated that a species
of Labyrinthula (Myxomycete), later determined by Young (19) as
L. macrocystis Cienk., actually was the agent that caused the destruction
of eelgrass over a considerable geographic range. In view of these later
findings, there arises the thought that Ophiobolus halimus may be a sec-
ondary parasite or indeed might even be saprophytic, thus placing the
species in the same category as those which are to be considered in the
present paper.
With the exception of the papers of Saccardo, Ellis, and Mounce and
Diehl, there is, as far as the authors are aware, only one published ref-
erence to the occurrence of marine fungi on phanerogamic plant remains.
This is a brief note by Johnson and McNeil (4) indicating the presence
? This study is a joint endeavor arising from a project undertaken by E. S. Barghoorn
on the microbiological factors responsible for the decomposition of submerged plant
remains in the sea. D. H. Linder has made all contributions to the taxonomy and re-
lationships of the fungi described. Problems encountered in the isolation and cultur-
ing of the various species have been worked out together. E. S. Barghoorn has un-
dertaken the field work, has gathered together most of the specimens and pertinent
collection data, has made the isolations of pure cultures and has carried out the ex-
perimental studies.
The present paper is one of a contemplated series of investigations on the various -
microbiological, chemical and physical factors involved in the decomposition or
preservation of submerged plant materials.
395
396 FarLowla, VoL. 1, 1944
of a seawater fungus in the decomposition of hardwood timbers in the
port of Sydney, Australia. No description of the fungus is made beyond
mention of the fact that it was referred to the class Ascomycetes and that
“some timbers are weakened by a softening of the surface fibres to a depth
of one-quarter of an inch of truewood in a period of seven years
immersion.”
The fact that a score or more of species have been described as occur-
ring in the sea is of importance since it shows that fungi not only tolerate
salt water, but indeed that marine conditions furnish a normal habitat
for the relatively small number of fungi that have become adapted to it.
The fungi considered in this study differ from the majority of those
already mentioned in that they do not occur on algae, but on wood and
plant remains in the sea. All of the species were obtained from specimens
of wood that had been either submerged continually from five months to
one year and were below low tide level for the entire period, or were ex-
posed in the inter-tidal zone where they were subjected twice a day to im-
mersion in seawater. Under these conditions, it is highly improbable
that the fungi are terrestrial forms which penetrated the wood after it
had been removed from the sea, or before it had been immersed. This
fact becomes evident when it is realized that the majority of the samples
were examined within one to three days of their removal from the marine
location, a time much too short to permit the formation and fruiting of
the ascigerous stage, and even too brief a time for the establishment of
any extensive colonies of the conidial phases. Another convincing line
of evidence that these fungi are normally marine in their occurrence is
the fact that the majority of them are very tolerant of sea water, even to
the extent of growing on artificial media made with sea water of three
times the normal saline concentration, as will be shown later in this study.
For the present, the collection data given below may serve to give some
insight into the conditions under which the various samples of wood and
cordage have become inoculated by the fungi.
SPECIMEN COLLECTION DATA
Speci- Type of
men Location Submerged Removed Conditions Specimen
1 Old Concord Wharf, Sept. 23/41 May 19/42 4 ft. below low Test block.
Portsmouth, New water.
Hampshire.
2 Searsport, Maine. Sept. 30/41 May 26/42 5 ft. below low Test block.
water.
3 Woods Hole, Mass. Oct. 13/41 June 13/42 7-8 ft. below low Test block.
water. 5 ft. above
mud.
4 Baylies Wharf, Fall Feb. 1942 July 18/42 8 ft. below low Test board.
River, Mass. water.
5 Tide flats, No. 1¢ June, 1942 Covered by 8ft.of Driftwood em-
Truro, 200 ft. from water at high tide. beddedin sand.
shore at low tide.
6 Chelsea, Mass. 1918 May, 1942 Near high tide line. Pile chip.
Speci-
men
7
29
30
BarcHoorn & LINDER: MARINE FUNGI
SPECIMEN COLLECTION DATA— Continued
Location
Same as 5.
Same as 5.
Same as 5.
Woods Hole, Mass.
B, & M. Bridge,
Portsmouth, N. H.
Bucksport, Maine.
Provincetown,
Mass.
Woods Hole, Mass.
Salem, Mass.
Saybrook, Conn.
Provincetown,
Mass
Newburyport,
Mass.
Woods Hole, Mass.
Rockland, Maine.
Thomaston, Maine.
Provincetown,
Mass.
Provincetown,
Mass.
Provincetown,
Mass.
Provincetown,
Mass.
Provincetown,
Mass.
Provincetown,
Mass.
Provincetown,
Mass.
Bridgeport, Conn
Submerged
?
?
?
March 6/42
Dec. 29/41
Dec. 19/41
1933?
April 6/42
May, 1942
May 1/42
?
May 25/42
?
March 10/42
Sept., 1938
a
1900 or 1901
Dec. 8/41
Removed
June, 1942
June, 1942
June, 1942
Aug. 5/42
Aug. 25/42
Aug. 25/42
July 24/42
Sept. 5/42
Sept. 18/42
Oct. 1/42
July 24/42
Oct. 16/42
Aug, 24/42
Nov, 18/42
Nov. 27/42
Sept. 24/42
Sept. 24/42
Sept. 24/42
Sept. 24/42
Sept. 24/42
Sept. 24/42
Sept. 24/42
Dec. 21/42
Conditions
Same as 5.
Same as 5.
Same as 5.
Below low water
at all times.
Below low water
at all times.
Below low water
at all times.
2 ft. above low
water.
Below low water
at all times,
Below low water
at all times.
Below low water
at al] times.
Tide flats, Covered
8 ft, at high water.
Below low water at
all times. Salinity
low.
Intertidal zone.
Below low water at
all times.
Below low water at
all times.
3 ft. above low
water.
3 ft. above low
water.
3 ft. above low
water.
3 ft. above low
water.
4-5 ft. above low
water.
2 ft. above low
water.
2 ft. above low
water.
Below low water.
397
Type of
Specimen
Driftwood.
Driftwood.
Driftwood.
Test board.
Test block.
Test block.
Pile chip. Cold
Storage Wharf.
Test board.
Test board.
Test board.
Timber em-
bedded in sand
Test board.
Manila rope.
Test block.
Back board for
test blocks.
Submerged
stump.
Submerged
stump (bark).
Submerged
stump.
Submerged
stump.
Jetty bulkhead
E. Breakwall,
Provincetown.
Driftwood.
Driftwood.
Test board.
Since these fungi appear to be terrestrial forms that have become
adapted to marine conditions, it may not be amiss at this time to point
out briefly some of the modifications that have occurred which appear to
make these forms better adjusted to their present environment.
One of
these changes which, with few exceptions, appears to hold for all species
belonging in the Pyrenomycetes, is that the ascus wall breaks down at an
early stage, very shortly after the ascospores have been delimited. In
398 FarLowla, VoL. 1, 1944
some cases, the ascus wall can be observed to be deeply eroded by the
time that the spores have been delimited. Exactly why dissolution of
the ascus wall is so common is difficult to conjecture, although it may be
presumed that with the breakdown of the wall, there is an increase in the
amount of hygroscopic material which, in addition to that already formed
within the ascus, aids in discharging the spores into the aquatic environ-
ment. When it is realized that the greater number of terrestrial species
are dependent for dispersal upon the forceful discharge of the spores into
the air where they are carried by air currents, it is apparent that a sub-
stitute method would be more effective for the dispersal of spores of the
marine species. The deliquescence of the ascus wall, then, would seem
to answer this need. In addition to this almost general characteristic,
there is another that immediately draws attention, viz., the presence of
processes on the spores that serve to keep the spores suspended for a
greater length of time in the water. Thus, Peritrichospora, which appears
to be closely allied to the terrestrial genus Ceriospora, is characterized
by numerous cilium-like processes that are produced around the middle
septa of the spores (Pl. V) and formed after the spores have been freed
from the ascus. In addition to the cilium-like processes, the spores have
an appendage at either end which soon becomes modified and eventually
deliquesces. These appendages appear to serve for the attachment of
the spores to the new substratum since when the spores are mounted in
water, the softened ends often cling to the cover glass or slide and can be
removed from their point of attachment only after considerable agitation
of the mount. Similarly, the spores of Remispora, while not having the
cilium-like processes, are equipped at each end with a pair of appendages
which appear to serve both for the attachment of the spores to the sub-
stratum, as well as to assist in suspending the spores in water and facili-
tating their dispersal. Unlike the appendages of the spores of Peritricho-
spora, those of Remispora appear to be in some way connected with the
astral rays of the nucleus since they are already formed within the ascus,
and, while the ascospores are immature, nearly surround them (PI. III,
figs. 7-10). As another example of adaptation in the pyrenomycetous
genera, the species of Halophiobolus furnish excellent illustration of their
suitability for aquatic dissemination. In these species, the spores (Pl. VI)
are filamentous and slender and sink less readily than would the rounded
or ellipsoid spores. While such filamentous spores are found in the dis-
comycetous genus Vibrissea which grows on decaying wood or leaves in
freshwater habitats, and are very common in the hypocreaceous genus
Cordyceps which, with few exceptions, parasitizes insects and is thus ter-
restrial, the two ends of the spores of the species of Halophiobolus are
equipped with modified tips which are somewhat gelatinous and thus are
capable of attaching the spores to any substratum with which they come
in contact. Indeed, it is very difficult because of the adhesiveness of the
tips, to discharge the spores from a glass pipette.
BarcHoorn & LINDER: MARINE FUNGI 399
A similar difficulty is encountered in handling the spores of several
other species for the following reason. Surrounding the spore there is a
jelly-like sheath or wing-like structure (PI. III, fig. 14) which becomes
more and more diffluent upon the release of the spores into water. Owing
to the tenacity of the jelly, the spores tend to cling to any object with
which they come in contact, even to such smooth surfaces as glass. A
spore modification closely resembling this is reported in the case of an
aquatic (freshwater) Ascomycete described by Weston (18) in which he
showed that the jelly firmly attaches the spore to the substratum or to any
surface upon which it settled. Thus it can be seen that there are special
modifications of the ascospores which not only aid in keeping the spores
suspended in water, but which also assist in holding them in contact with
any solid substratum which they encounter in the water. Among the
conidial forms, perhaps the most outstanding is Orbimyces (PI. I, fig. 5-6),
the conidia of which are dark and spherical or subspherical, and equipped
with finger-like processes which serve as a sort of parachute to keep the
conidia from sinking too rapidly. This modification of the conidium
parallels that recently described by Ingold (3) for a number of fresh-
water Fungi Imperfecti which also are characterized by their form and
the number of appendages they bear. The morphological response of
Orbimyces to its aquaeous habitat is so similar to that of the several species
described and delineated by Ingold that there seems no reason to deny
that Orbimyces is a form morphologically adapted to marine conditions,
just as are the other species to their freshwater habitat.
The morphological modifications of the ascospores and conidia lead
one to speculate as to the ancestry of these forms and the réle that they
play in the scheme of things. Of the former, there is so little evidence
on which to base theories that it does not seem worth while entering into
a discussion. The most that can be said is that these forms belong in
the same genera or in genera closely related to those that occur in ter-
restrial habitats. As regards the place that these forms take in the nat-
ural cycle of events, it would seem that they act more or less in the same
capacity as wood-rotting fungi in forests. The action on plant remains
does not appear to be as rapid as that brought about on land, but un-
questionably they play an important réle since many of the forms are
strong destroyers of cellulose, pectic substances, and other carbohydrates,
and to a lesser extent of lignin. Their economic significance to man is
at present being further studied, but from the evidence now available there
is not the least doubt that these forms are instrumental in speeding the
rate of decay of wooden piling and particularly of cordage by the suc-
cessive and continuous decaying of the outer parts that are submerged
intermittently or all the time in salt or brackish water. Since rope is
essentially cellulose, certain of these fungi play a very important part
in its deterioration. One of our collections, already listed, demonstrates
this fact most clearly for the samples of rope not only were penetrated
400 FarLowi, VoL. 1, 1944
in all directions by fungous hyphae, but the fungi were producing large
numbers of fruiting bodies all over the surface of the substratum and
producing spores in great numbers, as evidenced by the number of ma-
ture asci still present in a goodly proportion of the fruiting bodies. But
sufficient has been written at this stage to indicate the potential importance
of at least a fair proportion of these fungi. More extensive studies on
their physiology are being made, the results of which will appear later
in this and in subsequent papers.
ACKNOWLEDGEMENTS
The writers are deeply indebted to Dr. William F. Clapp and Miss
Ruth Lindquist, formerly of the Clapp Laboratories at Duxbury, Mass.,
for their keen interest in this problem as well as for their continued co-
operation in sending test boards for examination as soon as possible.
To Dr. Frederick S. Hammett, Director of the Marine Experimental
Station of the Lankenau Hospital Research Institute, appreciation and
thanks are extended for his hearty codperation and interest in these in-
vestigations and for making available the facilities of the Laboratory at
North Truro, Mass., where the use of laboratory space and equipment
greatly aided field collection and preparation of material for later study.
The writers wish to thank Dr. Alfred C. Redfield and Dr. Bostwick
Ketchum of the Woods Hole Oceanographic Institute for supplying speci-
mens of cordage of known conditions of exposure to sea water.
E. S. Barghoorn is indebted to the American Academy of Arts and
Sciences at Boston for grants-in-aid which have enabled him to carry on
extensive field work and other activities in the prosecution of this work.
BIBLIOGRAPHY
1. Cotton, A. D. Notes on marine Pyrenomycetes. Trans. Brit. Myc. Soc. 3:
92-99. 1907.
2. Ellis, J. B. & B. M. Everhart. New fungi. Journ. Myc. 1: 148-[150]-154.
1885,
3. Ingold, C. T. Aquatic Hyphomycetes of decaying alder trees. Trans, Brit.
Myc. Soc, 25: 339-417. pl. 12-17. 48 text figs. 1942.
4. Johnson, R. A. & F, A. McNeil. Supplementary Rept. No. 2. Maritime Serv-
ices Board of N. S. Wales. p. 36. 1941.
5. Mounce, I. & W. W. Diehl. A new Ophiobolus on eelgrass. Canad. Journ.
Res. 11: 242-246. ] pl. 1934. ,
6. Petersen, H. E. Contributions a la connaisance des Phycomycétes marins (Chy-
tridineae Fischer). Oversigt Kgl. Danske Vidensk. Selskabs. Forhandl. 1905:
439-488. 11 figs. 1905.
7. Renn, C. E. A mycetozoan parasite of Zostera marina. Nature 135: 544-545.
1935.
8. ————.. Persistence of the eel-grass disease and parasite on the American
Atlantic Coast. Nature 138: 507-508. 1936.
9, —____.. The wasting disease of Zostera marina. I. A phytological investiga-
tion of the diseased plant. Biol. Bull. 70: 148-159. 6 figs. 1936.
10. Saccardo, P. A. Sylloge Fungorum 2: 350. 1883.
BarRGHOORN & LinpER: MARINE FuNcI 401
11. Sparrow, F. K. Observations on marine Phycomycetes collected in Denmark.
Dansk. Bot. Arkiv 8: 1-24. 4 pls. 1934.
12, ————.. Biological observations on the marine fungi of Woods Hole waters.
Biol. Bull. 70: 236-263. 3 pl. 35 text figs. 1936.
13, ————. Aquatic Phycomycetes exclusive of the Saprolegniaceae and Pythium.
Univ. Mich. Stud. Sci. ser. 15: 1-785. 1 pl. 68 figs. 1943.
14. Sutherland, G. K. New marine Pyrenomycetes. Trans, Brit. Myc. Soc. 5:
147-154, pl. 3. 1915,
15. ————. New marine fungi on Pelvetia. New Phytol. 14: 33-43. 4 figs. 1915.
16. ————. Additional notes on marine Pyrenomycetes. New Phytol. 14: 183-
193. 5 text figs. 1915.
17. ————. Marine Fungi Imperfecti. New Phytol. 15: 35-38. 5 text figs. 1916.
18. Weston, W. H. Observations on Loramyces, an undescribed aquatic Ascomy-
cete. Mycologia 21: 55-76. pl. 8-9. 1929.
19. Young, E. L., WI. Labyrinthula on Pacific Coast eel-grass. Canad. Journ.
Res. 16C: 115-117. 1938.
I. CLASSIFICATION OF THE MARINE FUNGI
Davin H. LinpEeR
In undertaking the classification of the marine fungi that occur on
plant remains, the writer had little idea of the number of species that
would be encountered in so short a time, nor did he realize how great
would be the diversity of forms that would be discovered. The adapta-
tion of the species in some cases has been rather striking, as has already
been pointed out in the introductory pages, and it is this very factor that
makes difficult the task of relating the genera and species to those already
described from the terrestrial habitat. These points have impressed on
the writer the need for careful study of the species, and yet despite the
best of intentions, it has been necessary to describe numerous new species
or even genera, since, with but few exceptions, it has been impossible
to place the organisms in genera or species that have been described to
date. It is recognized, however, that with additional collections and a
greater assemblage of facts with which to work, forms will be found that
will throw considerably more light on the relationship between these
species that are about to be described and those that have already been
described from a terrestrial habitat. Such information, while highly
desirable, is not at this stage a prerequisite and for this reason the present
efforts should be considered merely as providing the names for those
organisms with which the student of the biology, physiology, and eco-
nomic importance of these fungi has to deal.
There have been difficulties in connection with the study of these marine
fungi, not the least of which has been the paucity of material for examina-
tion. Occasionally also, a colony that appeared on gross examination to
consist of one species, proved on sampling to be a mixture. Thus it early
became evident that in order to study these forms and to make a careful
record of their characters, it was first necessary to dissect out the pycnidia
402 Fartowla, VoL, 1, 1944
or perithecia and make temporary microscopical preparation from which
the structures could be studied and measured, and drawings made under
the low power of the microscope and with the aid of the camera lucida.
Once this preliminary work had been accomplished, the specimen, mount-
ed in acid-fuchsin-lactophenol, was gently crushed while being observed
under the low power of the microscope, after which further observations
were made of the more detailed structures. When these studies were com-
pleted, the specimens were remounted in glycerine-phloxine by the Diehl'
double cover glass method, and these slides, it is hoped, will serve as a
permanent reference for future investigators.
As a matter of convenience, the fungi are arranged in this paper more
or less in accord with Saccardo’s system:
Fungi Imperfecti
Sphaeropsidales
Moniliales
Pyrenomycetes
Amerosporae: |-celled spores
Didymosporae: 2-celled spores
Phragmosporae: 3- or more celled spores
Scolecosporae: filamentous, septate or non-septate spores.
It is obvious that this arrangement has very little if any phylogenetic
significance. The most that can be said in its favor is that it allows for
the more ready location of a desired species, a point of more importance
to the practical worker than to the taxonomic mycologist who is, or should
be, interested in the natural relationship of the different organisms. Be-
cause the writer feels that these marine fungi are of considerable economic
importance, convenience has governed the order of presentation of
the species.
FUNGI IMPERFECTI
SPHAEROPSIDALES
Phialophorophoma Linder, gen. nov.”
Pycnidiis atris, subcarbonaceis, subglobosis vel ellipsoideis, ostiolatis, immersis;
conidiophoris hyalinis, simplicibus vel infra ramosis; conidiis endogenis, hyalinis, ellip-
soideis, simplicibus.
Pycnidia black, subcarbonaceous, subglobose or ellipsoidal, ostiolate
and immersed in the substratum. Conidiophores hyaline, simple or
branched below, lining the cavity of the pycnidium. Conidia endogenous,
hyaline, ellipsoidal, 1-celled.
The genus receives its name from the fact that the conidiophores, like
those of Phialophora, produce the conidia endogenously and thus im-
mediately distinguish this sphaeropsidaceous fungus.
Diehl, W. W. Science, N. S. 69: 276. 1929.
? The writer wishes to acknowledge his thanks to Dr. Rolf Singer for his considerable
assistance in preparing the Latin descriptions.
BaRGHOORN & LINDER: MARINE FUNGI 403
Phialophorophoma litoralis Linder, sp. nov.
Plate I, figs. 7-9.
Pycnidiis in maculis atratis, immersis, atris et subcarbonaceis (vel atro-fuscis sub
microscopio) late-ovoideis vel ellipsoideis, 175 » longitudine, 118 yu. crassitudine,
116 y altitudine, ostiolo excluso; ostiolis late-conicis vel papilliformibus, leniter ex-
sertis. Conidiophoris (phialidiis) hyalinis, 7-10.5 x (2.5)-3-3.5 uw. sursum attenuatis
et ad apicem torque cupuliformi terminatis; conidiis hyalinis, ellipsoideis, piriformi-
bus, vel crepiduliformibus, plerumque biguttulatis, 2.5-3.5 x 1.5 pu, endogenis.
Pyenidia formed in the blackened area of the woody substratum, im-
mersed, broadly ovoid or ellipsoid, black, subcarbonaceous (deep fuscous
under the microscope) 175 yw long, 118 » wide, 116 » high excluding the
eccentric, broadly conoid or papilliform ostiole. Conidiophores (phia-
lides) hyaline, 7—10.5 x (2.5)-3-3.5 yw, tapering upward to a short cy]-
indrical neck that is surmounted by a flaring cup-shaped collar. Conidia
hyaline, ellipsoid to slipper- or pear-shaped, mostly biguttulate, 2.5-
3.9 x 1.5 uw, produced endogenously.
Specimens examined: Massachusetts, Provincetown, on oak bark about
two feet above the low tide level, Sept. 24, 1942, E. S. Barghoorn, 24;
Charleston, pilings (oak?) in Boston Harbor, May, 1942, comm. Ruth
Lindquist. Type.
This species and genus is characterized by the endogenous production
of the conidia. Judging by the fact that in other groups the endogenous
phialospores represent spermatia, it is not unlikely that this species rep-
resents the haploid phase of some Pyrenomycete.
Diplodia orae-maris Linder, sp. nov.
Plate I, figs. 1-3.
Pycnidiis pro parte immersis, ellipsoideis, membranaceis vel vetustis subcarbonaceis,
supra atris vel atro-fuscis, infra, in parte immersa leniter coloratis, 270 y, longitudine,
160 w altitudine, 150 y crassitudine; conidiophoris hyalinis 1-cellulatis, 6.5-8.5 x 2.5—
3.5 yu, cellulis hyalinis, bullatis, singillatim oriundis; conidiis ovoideis vel ellipsoideis,
brunneis, uniseptatis, 7.5-8.25 x 3.5—-4 yp.
Pycnidia partially immersed, ellipsoid, membranous or with age be
coming subcarbonaceous, black or deep fuscous above, lighter colored
in the lower submerged portions, 270 » long, 160 » high, 150 » wide, the
ostiole broadly papillate. Conidiophores hyaline, 1-celled, 6.5-8.5 x
2.5-3.5 p, arising singly from the hyaline, bullate cells that line the
greater part of the pycnidial cavity. Conidia ovoid to ellipsoid, brown-
ish, 2-celled, 7.5-8.25 x 3.5-4 p.
Specimen examined: California, Long Beach, on driftwood, Feb., 1943,
V.G. Dethier (c). Type.
MONILIALES
Botryophialophora Linder, gen. nov.
Mycelio aerio albo, bombycino; conidiophoris (phialidiis) plerumque tribus vel
compluribus ex cellula amplificata, globosa oriundis sive conidiophoris singulariter
oriundis, ergo cellula globosa frequenter destitutis; conidiis hyalinis conglobatis.
4.04, FarLowla, Vo. 1, 1944
Sterile mycelium white or light colored, cottony (in culture). Conidio-
phores (phialides) usually three or more from an enlarged globose or
subglobose basal cell which is produced pleurogenously on the main
hyphae or their branches; occasionally the conidiophores are produced
singly when they may be either pleurogenous or else acrogenous at the
ends of short side branches. Spores hyaline or light colored and tending
to aggregate in globose clusters.
Botryophialophora marina Linder, sp. nov.
Plate I, fig. 4.
Mycelio aerio albo vel vetusto dilute fusco, bombycino, ramoso, distanter septato,
5-10 y, diametro; conidiophoris (phialidiis) hyalinis, plerumque 3-20 e cellula am-
plificata, globosa vel subglobosa pleurogena oriundis, primum simplicibus deinde infra
ramosis, vasiformibus, 3.5-8.5 x 1.5-2.5 u.; conidiis hyalinis, globosis, 2-3 y, diametro,
vacuolatis, frequenter in capitulis globosis albidisque aggregatis.
Aerial mycelium white or becoming dilute fuscous with age, cottony
and tending to become repent, loosely branched, distantly septate, 5-10 p
in diameter. Conidiophores (phialides) hyaline, rarely 1-2 borne
pleurogenously or acrogenously on the ends of short branches, usually
from 3-20 or more arising from a globose or subglobose enlarged cell
which arises pleurogenously on the main hyphae, at first simple becoming
branched below, flask- or vase-shaped, 3.5-8.5x 1.5-2.5 yp. Conidia
globose or subglobose, hyaline, 2-3 » in diameter, usually with a large
central vacuole, tending to aggregate in white spherical heads.
Specimens examined: Massachusetts, North Truro, on driftwood cov-
ered by eight feet of water at high tide, FE. S. Barghoorn, 8. Type.
The above description was drawn from the fungus as it appeared in
culture. It seems quite likely that the species on its natural substratum
and especially in its native environment would be so inconspicuous as to
be overlooked. Furthermore, the fungus, as is evidenced by the fact that
the spores are small and are produced in phialides, may in the future
prove to be the conidial or spermatial stage of some ascomycetous species.
Orbimyces Linder, gen. nov.
Mycelio aerio fusco, atro-olivaceo vel subnigro, distanter septato; conidiis pleuro-
genis in dentibus sporogenis brevibus, globosis, nitide-nigris coronam ex 3-4 ap-
pendiculis digitatis, fuscis, septatis consistentem et circum basem turgidam appendi-
culae unicae erectae dispositis.
Aerial mycelium light fuscous to dark olivaceous or nearly black, dis-
tantly septate. Conidia pleurogenous on short sporogenous teeth, globose
shining black and bearing a crown of three to four fuscous, finger-like
appendages that are arranged radiately around the somewhat swollen base
of a single central appendage which, like the radiate appendages, is
fuscous and septate.
Orbimyces spectabilis Linder, sp. nov.
Plate I, figs. 5-6.
Coloniae fuscae vel atro-olivaceae, nonnihil jubatae; mycelio bombycino laxo un-
dulatoque, sparse ramoso et interdum anastomosante, distanter septato, 1.5-3.5
BarRGHOORN & LINDER: MARINE FUNGI 405
diametro; conidiis pleurogenis in dentibus sporogenis, subglobosis, 24~33 x 23-29 Lbs
nitide-nigris et coronam e 3-4 appendicularum digitiformium, fuscarum, septatarum,
consistentem et circum basem turgidam appendiculae unicae erectae dispositis.
Colonies fuscous to dark olivaceous, somewhat tufted. Mycelium
cottony loose and undulate, sparsely branched and occasionally anas-
tomosing, distantly septate, 1.5-3.5 » in diameter. Conidia borne pleu-
rogenously onthe lower parts of the hyphae on short inconspicuous
hyaline sporogenous teeth, readily deciduous, subglobose, 24-33 x 23-
29 », shining black and bearing at the distal end a crown consisting of
3-4 fuscous, septate, finger-like appendages that are arranged around the
somewhat swollen base of the central erect appendage that is of similar
length, color, and septation as the radiate members.
Specimen examined: Massachusetts, Charleston, on piling (oak?)
‘in Boston Harbor, May, 1942, communicated by Ruth Lindquist. Type.
This genus and species receives its name from the fancied resemblance
of the conidia to the orb which is a part of English royal regalia.
Alternaria maritima Sutherland, New Phytol. 15: 46-47. figs. 5-7. 1916.
Plate II, fig. 1.
“Mycelium diffused; conidiophores erect, simple, rarely slightly
branched, brown; conidia in simple or branched chains, pyriform with
neck-like apex, dark brown, verrucose, 30-50 » x 12-18 p.”
Specimen examined: Massachusetts: Woods Hole, isolated from test
block submerged seven to eight feet below low tide level on Oct. 13, 1941,
and removed June 13, 1942, W. F. Clapp, communicated by E. S. Barg-
hoorn, 3a.
The present specimen seems to agree closely enough with that described
by Sutherland, whose description is quoted above, to consider the two as
belonging to the same species. The spores of the specimen communicated
by Barghoorn, however, are smooth to slightly roughened and measure
(16.5)—21.5-38-(60, including the neck-like appendage) x (8.25)—10-
11.5-(13.5) ». Since species of Alternaria and Macrosporium show such
extreme variation in culture, with age, and in different environmental con-
ditions, the slight variation just mentioned does not seem to warrant the
erection of a new species in an already large genus.
Sutherland reports that in England this species is a saprophyte on
Laminaria fronds, and this would indicate that the species could grow on
wood where it might readily attack pectin and cellulose, among other
carbohydrates. :
Helicoma maritimum Linder, sp. nov.
Plate II, figs. 2-6.
Mycelio sterili in substrato plerumque immerso, ramoso, dilute vel atre fusco;
conidiophoris magnitudine mutabilibus, usque ad 100 y. longitudine, 3-5 yu. diametro,
vulgo ramulis rectis vel curvulis mycelio sterili oriundis, distanter septatis; conidiis
fuscis vel atro-fuscis, plerumque adumbratione aliquantulum quadrata gaudentibus
406 FartowI!A, Vou. 1, 1944
4. to 10-septatis, septis atro-fuscis, filis in spiras 1% vel bis convolutis, 8.5-10 p
diametro, spiris 23-26.5 y. diametro.
Sterile mycelium mostly in the substratum or spreading over its sur-
face, branching, dilute to deep fuscous. Conidiophores variable in size, up
to 100 » long, 3-5 p» in diameter, usually appearing as hyaline or dilute
fuscous, erect or irregularly bent branches from the sterile mycelium,
distantly septate. Conidia deep fuscous or almost black, often with a
squarish outline, 4- to 10-septate, the septa deep fuscous or black, the
spore filament 8.5—10 » in diameter, 134—2 times coiled, the coil 23-26.5 »
in diameter.
Specimens examined: Maine, Searsport, on test block immersed five
feet below low tide level on Sept. 30, 1941, and removed May 26, 1942,
W. F. Clapp, communicated by E. S. Barghoorn, 2; Massachusetts,
Woods Hole, on test block immersed seven to eight feet below low tide
level on Oct. 13, 1941, and removed June 13, 1942, W. F. Clapp, com-
municated by E. S. Barghoorn, 3a; Woods Hole, on test board submerged
below low tide level Mar. 6, 1942, and removed Aug. 5, 1942, W. F. Clapp,
communicated by E. S. Barghoorn, 10b.
Although the majority of the Helicosporeae are terrestrial, there is
little doubt of this species belonging in the marine habitat since in a
freshly removed test board that had been immersed for eleven months,
the spores of this fungus were discovered on the wood under and around
incrusting algae, sponge spicules and other debris.
Helicoma salinum Linder, sp. nov.
Plate II, figs. 8-11.
Coloniae effusae, sparsae; conidiophoris ramulis brevibus, dilute fuscis mycelio
sterili oriundis, cellula terminali obscuriore atque atro-fusco, cellula extrema globoso-
inflata est, atro-fusca; conidiis acrogenis, maturitate fuscis praeter cellulas duas basales
dilutius coloratas, 11—26-septatis, ad septa leniter constrictis, filis in spiras 1%4—4%4-
convolutis, 5.5-8 y, diametro, ad basem attenuatam 3.5-4.5 p. diametro, spiris 28-39 p,
diametro.
Colonies effuse, sparse. Conidiophores as rather short slender side
branches from the repent mycelium, light fuscous except for the terminal
cell which is darker or else when it is globose, inflated and deep fuscous.
Conidia acrogenous, at first darkest near the apex but later, except for the
one or two lighter colored basal cells, becoming uniformly fuscous,
11- to 26-septate, slightly constricted at the dark septa, 154 to 444 times
coiled, the filament 5.5-8 « in diameter, gradually tapering to 3.5-4.5 p
in diameter near the base which is rounded-truncate; the coiled spore
28-39 p» in diameter. a
Specimens examined: Maine, Bucksport, on test block below water at
all times, submerged Dec. 19, 1941, removed Aug. 25, 1942, W. F. Clapp,
communicated by E. S. Barghoorn, 13; Massachusetts, North Truro, on
driftwood covered by eight feet of water at high tide, June, 1942, E. S.
Barghoorn, 5. Type; Connecticut, Saybrook, on test board below low
tide level at all times, immersed May 1, 1942, removed Oct. 1, 1942, W.
F. Clapp, comm. E. S. Barghoorn, 17.
BarcHoorn & LiInDER: MARINE FUNcI 407
This species in the younger stages resembles H. maritimum, but may
in most cases be readily distinguished by the fact that the conidia tend to
coil in three instead of two planes, and by the more numerous coils and
_ septations of the spores.
Speira (Cattanea) pelagica Linder, sp. nov.
Plate II, fig. 7.
Mycelio sterili in substrato immerso, hyalino, ramoso; conidiophoris hyalinis, sim-
plicibus vel rarissime septatis, 16-28 x1.5-2 4; conidiis atro-brunneis, plerumque
24.5-30 x 16.5-24.5 , versiformibus, primum cellulis in ordinibus tribus, raro duobus
vel quattuor, ex cellula unica basali oriundis deinde saepe irregulariter multiseriatis,
septis atro-brunneis vel sub-atris, cellulae 5-6.5 x 5—5.75 wu. metientibus.
Sterile mycelium mostly within the substratum, inconspicuous, hya-
line or locally brownish and branched. Conidiophores hyaline, simple
or very rarely septate, slender, mostly 16-28 x 1.5—2.5 ». Conidia dark
brown under the microscope or black under the hand lens, mostly 24.5—
30 x16.5—24.5 y, at first nearly flat with a single cell giving rise to three
rows of 6 to 7 cells, rarely two or four rows, but later through division
of the cells the spores tend to become terete, irregularly multiseriate and
to assume very irregular shapes, occasionally branched and mitten-shaped,
the septa dark brown to nearly black, the cells variable in size but mostly
5—6.5 x 5-53.75 p.
Specimen examined: Massachusetts, on driftwood covered by eight
feet of water at high tide, July 24, 1942, E. S. Barghoorn, 18. Type.
The conidia of this species in the early stages of formation are typical
of the section Cattanea of Speira and the above description is based
mainly on this phase. Occasionally spores are found in which there has
been considerable proliferation of the cells and in such instances they are
strongly reminiscent of Sporodesmium cellulosum Sacc.
PYRENOMYCETES
AMEROSPORAE
Samarosporella Linder, gen. nov.
Peritheciis nigris, carbonaceis, immersis, globosis, ostiolatis; ostiolis papilliformibus
et paullulum exsertis; paraphysibus absentibus; ascis fusoideis, apicibus rotundatis
instructis; ascosporis 4-8, hyalinis, elongato-ellipsoideis vel fusoideis, alatis; asco-
sporis abortivis non alatis et multum minoribus.
Perithecia imbedded, black and carbonaceous, globose and ostiolate.
The ostiole papilliform and slightly exserted beyond the substratum.
Paraphyses absent. Asei fusoid with rounded apices, deliquescing, 4—8-
spored and usually with some of the spores aborted. Ascospores hyaline,
elongate-ellipsoid or somewhat fusoid, with irregular hyaline wings; the
aborted ascospores much smaller, ovoid or ellipsoid and devoid of hya-
line wings or else with very rudimentary ones.
This genus, because of the winged spores, appears to be related to
Samarospora of Rostrup (Bot. Centralbl. Beih. 3: 3. 1893) but from that
408 - Fartowra, Vou. 1, 1944
genus it differs by the perithecia having an ostiole and by the asci being
fusoid rather than globose or subglobose. It should be noted that
Samarospora is also an aquatic species which grows on Potamogeton.
Samarosporella pelagica Linder, sp. nov.
Plate III, figs. 1-4.
Peritheciis nigris, solitariis et sparsis, carbonaceis, in substrato immersis, sub-
globosis, plus minusve 170 p. crassitudine, 195 y, altitudine, ostiolatis; ostiolis papilli-
formibus et paullulum exsertis; ascis fusoideis, apicibus rotundatis instructis,
140-165 x 25-30 yp, deliquescentibus; ascosporis 4-8, hyalinis, elongato-ellipsoideis
vel nonnihil fusoideis, alatis, (18)-25-30—(39.5) x 7.5-10.5-(11.5) yp alis exclusis;
ascosporis abortivis non alatis, 15-20x 7-8 u.
Perithecia black, solitary and distantly scattered, carbonaceous, im-
mersed in the substratum, subglobose or globose + 170 p thick, 195 p
in height, ostiolate. The ostiole papillate or somewhat flattened above,
slightly exserted. The asci fusoid, somewhat rostrate with a rounded
apex, 140—165 x 25-30 y, deliquescing as the spores mature. Ascospores
4-8 in the ascus, usually some of them aborted, hyaline, elongate-ellipsoid
or somewhat fusoid, (18)—25—30—(39.5) x 7.5-10.5-(11.5) y, excluding
the hyaline wings which are somewhat undulate and of irregular width;
the aborted ascospores not winged or the wings vestigial, ellipsoid, 15—20 x
7-8 p.
Specimen examined: Massachusetts, Provincetown, on driftwood cov-
ered by eight feet of water at high tide, July 24, 1943, E. S. Barghoorn,
18. Type.
DIDYMOSPORAE
Ceriosporopsis Linder, gen. nov.
Peritheciis nigris (atro-brunneis sub microscopio) , membranaceis et collabescentibus
siccando, plerumque in substrato immersis, substrato attrito autem superficialibus,
ovoideis vel elongato-ellipsoideis, ostiolatis, ostiolis excentricis, rotundato-conicis vel
papilliformibus; paraphysibus absentibus; ascis non visis, sed octo-ascosporis fre-
quenter coacervatis; ascosporis hyalinis vel dilute coloratis, ellipsoideis, uniseptatis
et ad apices ambos appendiculatis; appendiculis hyalinis, robustis fastigatisque
demum dehiscentibus vel deliquescentibus qua ex re sporae demum utrinque trunca-
tae sunt.
Perithecia black, or dark brown under the microscope, membranous
and collapsing on drying, mostly buried in the substratum but becoming
exposed by erosion of the decayed wood, ovoid to elongate-ellipsoid,
ostiolate; the ostiole eccentric, rounded-conoid or papilliform. Para-
physes absent. Asci not seen but probably 8-spored since the spores occur
frequently in bundles of eight. Ascospores hyaline or dilutely colored,
two-celled and with a stout hyaline and tapering elongate appendage at
each end, the appendages dehiscent or deliquescent and then the asco-
spores are truncate at each end.
This genus strongly resembles Ceriospora but differs by the irregular
shape of the membranous, collapsing perithecia, the deliquescent asci,
and the deliquescent appendages of the ascospores. In view of the fact
that the asci of this species soon deliquesce, Saccardo and Marchal (Rev.
BarcHoorn & Linper: MARINE Func 409
Myc. 7: 148. 1885) may have misinterpreted their genus Tiarospora when
they placed it in the Sphaeropsidales and the spores that were described
as conidia of the imperfect genus were actually liberated by the early
dissolution of the ascus wall. That the spores of Ceriosporopsis are asco-
spores is clearly indicated by the frequent occurrence of the spores in
clusters of eight and by the fact that in neither of the appendages is there
any evidence of a canal that would indicate connection with a conidio-
phore. The genus appears, because of the structure of the wall and the
nature of the spores, to be related to Bombardia or to the Sordariaceae.
Ceriosporopsis halima Linder, sp. nov.
Plate III, figs. 10-12.
Peritheciis nigris vel sub microscopio atro-brunneis, membranaceis, plerumque in
substrato immersis vel substrato attrito superficialibus, ovoideis vel elongato-ellipsoi-
deis, axi longiora superficie substrati parallelo, 175-400 x 90-300 u, ostiolatis; ostiolis
excentricis, rotundato-conicis vel papilliformibus; ascis non visis; ascosporis hyalinis
vel dilute coloratis, uniseptatis et ad apices ambos appendiculatis, 22.5-26-(35) x
8-10 y appendiculis exclusis, appendiculis hyalinis, robustis fastigatisque 14-20 x
5-7 wu, demum dehiscentibus vel deliquescentibus, qua ex re sporae truncatae sunt.
Perithecia black to the naked eye or dark brown under the microscope,
membranous, buried in the substratum or becoming superficial through
the erosion of the matrix, ovoid to elongate-ellipsoid, variable in shape
and size, 175-400 x 90-300 yu, the long axis parallel to the surface of the
substratum; the ostiole conspicuous, rounded-conoid or papillate, eccen-
tric. Ascospores ellipsoid, hyaline or very dilutely colored, 22.5—26-
(35) x 8-10 p, 2-celled, each cell usually with a single large vacuole, with
stout hyaline, tapering, straight or curved appendages at each end, the
appendages somewhat gelatinized and either deliquescent or dehiscent.
Specimens examined: New Hampshire, Portsmouth, on test block
below water at all times, immersed Dec. 29, 1941, removed Aug. 25, 1942,
W. F. Clapp, comm. E. S. Barghoorn, 11; Massachusetts, Newburyport,
on test board in brackish water, submerged May 25, 1942, and removed
Oct. 16, 1942, W. F. Clapp, comm. E, S. Barghoorn, 19a; Connecticut,
Saybrook, on test block below water at all times, immersed Dec. 19, 1941,
removed Aug. 25, 1942, W. F. Clapp, comm. E. S. Barghoorn, 17. Type.
Dr. Barghoorn has found by single spore culture that this species is
the ascigerous stage of Helicoma salinum which is described in the present
paper among the Fungi Imperfecti.
Remispora Linder, gen. nov.
Peritheciis solitariis vel nonnihil gregariis, in substrato immersis, dilute coloratis
cremeis vel isabellinis, membranaceis, subglobosis, conspicue ostiolatis, ostiolis
truncato-conicis, eccentricis; paraphysibus absentibus; ascis late fusoideis nonnihil
apiculatis, 3-8 sporas efferentibus. Ascosporis ovoideis, ellipsoideis vel elongato-
ellipsoideis, hyalinis, uniseptatis, ad apicem utrumque bi-appendiculatis; appendiculis
subgelatinosis divergentibusque. -
Perithecia solitary or somewhat gregarious, imbedded in the sub-
stratum, light colored, about isabellinous or cream-colored, subglobose,
410 Fartowia, VoL. 1, 1944
membranous and with a pronounced truncate-conoid, eccentric ostiole.
Paraphyses absent. Asci broadly fusoid with a slight apiculus, tardily
deliquescent, 3—8-spored, usually with some of the spores aborted. Asco-
spores ovoid, ellipsoid or elongate-ellipsoid, hyaline, 1-septate, with two
hyaline, broad and tapering subgelatinous appendages at each end, the
appendages diverging and nearly at right-angles to the long axis of the
spore, and later deliquescing or dropping off.
This genus is characterized by the mustache-shaped appendages at each
end of the two-celled spores, and these with the light-colored, membranous
perithecial wall, seem ample to segregate the species from Massarinula.
It seems probable that the appendages serve, at least temporarily, as a
means of keeping the spores suspended in the water and hence assist in
the dispersal of the species. The name is derived from remus — oar,
and spora.
Remispora maritima Linder, sp. nov.
Plate III, figs. 5-9.
Peritheciis solitariis vel subgregariis, in substrato immersis, cremeis yel isabellinis,
membranaceis, subglobosis, 230-500 x 225-500 u.; ostiolis truncato-conicis, excen-
tricis; ascis late fusoideis et nonnihil apiculatis, 48-65 x 12-20 45 ascosporas 3-8
gerentibus; ascosporis ovoideis, ellipsoideis vel elongato-ellipsoideis, hyalinis, uni-
septatis, ad septa non constrictis, (17.5)-—21.5-27-(31.5) x (8)—10-11.5—(12) w=. ap-
pendiculis exclusis, ad apicem utrumque bi-appendiculatis, subgelatinosis, fastigatis,
divergentibus.
Perithecia solitary or somewhat gregarious, immersed in the sub-
stratum, cream colored or isabellinous, membranous and collapsing on
drying, subglobose, 230-500 x 225-500 », with a truncate, conoid, and
eccentric or nearly lateral ostiole. Asei broadly fusoid and somewhat
apiculate, tardily deliquescent, 48-65 x 12-20 ». Aseospores 3-8 per
ascus, not infrequently some of them aborted, ovoid, ellipsoid, or elongate-
ellipsoid, hyaline, 1-septate, not constricted at the septum, (17.5)—21.5—
27—(31.5) x (8)—-10—-11.5-(12) p excluding the appendages; appendages
in pairs at each end, stout and tapering, somewhat gelatinous, divergent
and somewhat recurved so that they are approximately at right angles
to the long axis of the spore.
Specimens examined: Maine, Thomaston, on back board for test blocks
below water at all times, immersed Sept. 1938, removed Nov. 27, 1942, W.
F. Clapp, comm. E. S. Barghoorn, 22B; Massachusetts, Provincetown,
jetty bulkhead of breakwall three feet above low tide level, built 1900-01,
removed Sept. 24, 1942; Provincetown, on driftwood two feet above low
tide level, Sept. 24, 1942, E. S. Barghoorn, 29. Type.
Unlike most appendiculate species described in this paper, the ap-
pendages of this one appear not to be secondary formations, but rather
seem to be remnants of the ascus protoplasm that in some way have been
differentiated during the formation of the spores. When the spores are
mounted in acid-fuchsin lactophenol and examined under the oil immer-
sion lens, definite striae may be seen, as may also remnants of the undiffer-
entiated ascus protoplasm (plate III, 9b.).
BarRcGHoorN & LINDER: MARINE FUNGI All
Amphisphaeria maritima Linder, sp. nov.
Plate III, figs. 13-16.
Peritheciis nigris, carbonaceis, subsphaericis vel late rotundatis et nonnihil conicis,
usque ad 200 » diametro, superficialibus vel partim immersis; ascis cylindricis,
54.5-60 x 8-13 yu, octosporis, distichis; ascosporis brunneis, uniseptatis, ad septa
leniter vel haud constrictis, 16.5x5 uw.
Perithecia black and carbonaceous, subspherical or broadly rounded
and conical, up to 200 » in diameter, ostiolate, superficial or else only
partly immersed in the substratum. Asei cylindrical with the apical wall
somewhat thickened, 54.5—60 x 8-13 p, 8-spored, distichous. Ascospores
brown, uniseptate, not or only slightly constricted at the septa, 16.5 x5 up.
Specimens examined: Maine, Bucksport, on test block below water at
all times, immersed Dec. 19, 1941, removed Aug. 25, 1942, W. F. Clapp,
comm. E, S. Barghoorn, 13; Massachusetts, Woods Hole, on test block
seven to eight feet below low tide level, immersed Oct. 13, 1941, removed
June 13, 1942, W. F. Clapp, comm. E. S. Barghoorn, 3a. Type.
Lentescospora Linder, gen. nov.
Peritheciis solitariis vel sparsis, membranaceis et collabescentibus siccando, nigris
vel sub microscopio atro-brunneis, ovoideis vel subcylindraceis, minute ostiolatis;
paraphysibus absentibus; ascis non visis sed probabiliter deliquescentibus maturitate ;
ascosporis hyalinis, ellipsoideis vel elongato-ellipsoideis, 0- vel l-septatis, raro 2-sep-
tatis et ad septa leniter constrictis, parietibus ad apices ambos incrassatis et sub-
gelatinosis. :
Perithecia solitary or scattered, membranous and collapsing on dry-
ing, black to the naked eye or dark brown under the microscope, ovoid
to subcylindrical, with a minute papillate ostiole. Paraphyses absent.
Asci not seen and are probably soon deliquescent. Ascospores hyaline,
ellipsoid or elongate-ellipsoid, 0- or 1-septate, rarely 2-septate, somewhat
constricted at the septa, the walls of the ends of the spores thickened and
somewhat gelatinized.
It is probable that this genus is closely related to both Ceriosporopsis
and to Remispora since the perithecia of all are membranous and col-
lapsing, and the spores are hyaline, two-celled, and with modified ap-
pendages, although in this genus the appendages may be considered as
being reduced to a mere thickening and gelatinization of the apical walls
of the spores (PI. IV, fig. 3). The ostiolar neck is much reduced and
consists of one or two rows of cells arranged vertically around the opening
of the perithecium (PI. IV, fig. 2). These differences, in the light of our
present incomplete knowledge of related forms, seem to warrant the erec-
tion of the above described genus, the name of which is derived from
lentesco = to stick (referring to the gelatinized apices of the spores)
and spora.
Lentescospora submarina Linder, sp. nov.
Plate IV, figs. 1-3.
Peritheciis solitariis vel sparsis, membranaceis et collabescentibus siccando, nigris,
sub microscopio autem atro-brunneis, ovoideis vel subcylindraceis, 225-320 x 80-130 py,
412 FarLowiA, VoL. 1, 1944
ostiolis exiguis, papillatis et supra substratum paullo exsertis; ascosporis hyalinis,
ellipsoideis vel elongato-ellipsoideis, 23-28 x 6-10 u, 0- vel 1-septatis, raro 2-septatis
et ad septa leniter constrictis, parietibus ad apices ambos incrassatis et subgelatinosis.
Perithecia solitary or scattered, membranous and collapsing on dry-
ing, black to the naked eye but dark brown under the microscope, ovoid
to subcylindrical, 225-320 x 80-130 y», with a minute, papillate, central
and scarcely exserted ostiole. Ascospores hyaline, ellipsoid or elongate
ellipsoid, 23-28 x 6-10 p, 0- or 1-septate, rarely 2-septate, somewhat con-
stricted at the septa, apical walls of one, or most frequently both ends of
the spores, thickened and somewhat gelatinized.
Specimen examined: Connecticut, Bridgeport, on test board sub-
merged at all times, immersed Dec. 3, 1941, removed Dec. 21, 1942, WV.
F. Clapp, comm. E. S. Barghoorn, 30. Type.
Halosphaeria Linder, gen. nov.
Peritheciis solitariis vel subgregariis, immersis, ovoideis, membranaceis, superne
atro-fuscis vel subnigris, infra subhyalinis, ostiolatis; ostiolis excentricis et obliquis,
atris, crassis et breviter cylindricis; paraphysibus absentibus; ascis late clavatis;
ascosporis quattuor, ellipsoideis, hyalinis, bi-cellulatis, cellula quaque vacuola unica
magna instructa, sporis ad septum leniter vel haud constrictis tribusque appendiculis
hyalinis, gracilibus ornatis quarum duae vulgo ad septum unaque frequenter ad
unum apicem nascuntur.
Perithecia solitary or subgregarious, immersed in the substratum, ovoid,
membranous, dark fuscous or nearly black above; subhyaline below,
ostiolate; the ostiole eccentric and oblique, black, stout and_short-
cylindrical. Paraphyses absent. Asci 4-spored, broadly clavate. Asco-
spores ellipsoid, hyaline, two-celled, each cell usually with a single large
conspicuous vacuole, the spores not or only slightly constricted at the
septum and provided with (2)—3—(4) slender hyaline appendages of which
two are usually at the septum and one at one end.
Halosphaeria appendiculata Linder, sp. nov.
Plate IV, figs. 4-7.
Peritheciis solitariis vel subgregariis, immersis, ovoideis, membranaceis, superne
atro-fuscis vel fuligineo-atris, infra subhyalinis, 425-500 x 317-330 w, ostiolatis;
ostiolis excentricis obliquisque, atris, crassis et breviter cylindricis; ascis 4-sporis,
late clavatis, plus minusve 55x 18.5 4; ascosporis ellipsoideis, 25-29 x 9-12 u, hya-
linis, 2-cellulatis, cellula utraque vacuola simplici magna conspicuaque instructa,
sporis non vel leniter constrictis ad septum tribusque appendiculis hyalinis gracilibus
ornatis quarum duae vulgo ad septum unaque frequenter ad unum apicem nascuntur,
appendice terminali interdum nulla.
Perithecia solitary or subgregarious, immersed in the substratum,
ovoid, membranous, dark fuscous to nearly black above, shading to sub-
hyaline below, 425-500 x 317-330 y, ostiolate; the ostiole eccentric and
oblique, black, stout and short-cylindrical. Asci 4-spored, broadly clavate,
+ 55x18.5 ». Ascospores ellipsoid, 25-29x9-12 yp, hyaline, two-
celled, each cell with a large conspicuous vacuole, the spores not or but
slightly constricted at the septum and provided with (2)—3-(4) slender,
hyaline, tapering appendages, 11-15 x 1-1.5 », of which two are usually
at the septum and one at one end of the spora or occasionally the terminal
appendage may be lacking.
BarcHoorn & LINDER: MARINE FUNGI 413
Specimen examined: Massachusetts, Newburyport, on test board in
brackish water, below low water at all times, submerged May 25, 1942,
removed Oct. 16, 1942, W. F. Clapp, comm. E. S. Barghoorn, 19c. Type.
PHRAGMOSPORAE
Leptosphaeria orae-maris Linder, sp. nov.
Plate IV, figs. 8-11.
Peritheciis solitariis vel subgregariis, immersis, late ovoideis vel subglobosis, mem-
branaceis, superne aterrime fuscis vel atris, ad latus inferum pallescentibus, 160-240 p.
longitudine, 140-200 yp, latitudine, 125-200 », altitudine, ostiolatis; ostiolis atris, brevi-
bus, papillatis, leniter exsertis ultra substratum; paraphysibus ramosis, hyalinis, in-
terdum anastomosantibus, longitudine variabili sed longioribus ascis, 1-1.5 B dia-
metro; ascis clavatis, primum crasso-tunicatis deinde tenui- -tunicatis, octosporis, dis-
ec parte sporogena 58 » longa et 8-10 y, lata} ascosporis late fuscoideis, fuscis,
l-septatis, tarde 3-septatis, constrictis ad septum centrale; parietibus levibus vel sub-
asperulatis, 18-22 x 5-6.5 u.
Perithecia solitary or subgregarious, immersed in the substratum,
broadly ovoid to subglobose, membranous, very dark brown or black
above and becoming lighter colored below, 160-240 p, long, 140-220 yp
wide, 125-200 » high, ostiolate; the ostiole black, short, papillate and
slightly exserted beyond the substratum. Paraphyses hyaline, branched
and occasionally anastomosing, of variable length but exceeding the length
of the asci, 1-1.5 » in diameter. Ascus clavate, at first thick-walled but
later becoming thin-walled, 8-spored, distichous, the sporogenous part
58 x 8-10 ». Asecospores broadly fusoid, brown, 1-septate, tardily be-
coming 3-septate, constricted at the middle septum, the wall smooth to
asperulate, 18-22 x 5-6.5 p.
Specimen examined: California, Long Beach, on driftwood, Feb. 1943,
V.G. Dethier (B). Type.
Cooke and Plowright (Grevillea 5: 120. 1877.) described Sphaeria
maritima, later transferred to Leptosphaeria by Saccardo (Syll. Fung. 2:
73. 1883) as occurring on Juncus maritima. Because of its habitat, there
was the possibility that this might be the same as the present species, but
instead, the spores proved to be 5-septate and yellowish rather than tardily
3-septate and brown. Leptosphaeria muralis Sacc., not from a marine
habitat, in general appearance seems to be closely related to this species
but the perithecia are entirely black and the spores are smaller.
Sphaerulina orae-maris Linder, sp. nov.
Plate IV, figs. 12-15.
Peritheciis solitariis vel sparsis, membranaceis, partim immersis, atro-fuscis vel
subatris superne, dilute fuscis infera in parte, globosis vel subglobosis, 200 uw. cc.
diametro; ascis clavatis, tristichis, 72 x 10-14 4; ascosporis 8, fusoideis utrinque ob-
tusatis, inaequilateralibus, hyalinis, 3-septatis, haud vel Jeniter constrictis ad septa,
parietibus levibus, 26-30 x 5-6.5 y.
Perithecia solitary or scattered, membranous, partially immersed in
the substratum, subglobose or globose, dark brown or nearly black above,
becoming light fuscous below, approximately 200 » in diameter. Para-
414, FarLowl14, Vou. 1, 1944
physes lacking. Asci clavate, tristichous, 72x 10-14 ». Ascospores
8, fusoid and bluntly rounded at both ends, inequilateral, hyaline, 3-septate,
not or only slightly constricted at the septa, with smooth walls, 26-30 x
5-6.5 pb.
Specimen examined: California, Long Beach, on driftwood, Feb. 1943,
V.G. Dethier. Type.
Peritrichospora Linder, gen. nov.
Peritheciis solitariis vel sparsis, majorem ad partem immersis, globosis, carbonaceis,
nigris, ostiolatis; ostiolis minutissimis, papillatis, apicalibus; paraphysibus absentibus;
ascis fusoideis vel late fusoideis; ascosporis 8, fusoideis, biappendiculatis, hyalinis,
fuscis circum septum centrale, aut quinque-septatis aut ]-septatis, atque 1-3 pseudo-
septis additis ad utrumque latus, multi-crinitis circum septum centrale, appendicibus
terminalibus hyalinis, subcylindricis, apicibus obtusis.
Perithecia solitary or scattered, for the most part immersed in the sub-
stratum, globose, carbonaceous, black, ostiolate; the ostiole extremely
small and papillate, central. Paraphyses absent. Asci fusoid or broadly
fusoid. Ascospores 8, fusoid, biappendiculate, hyaline but becoming
brownish around the central septum, either 5-septate or l-septate with
1—3 pseudosepta on each side of the septum, polytrichous around the
septum; the terminal appendages hyaline, subcylindrical and with ob-
tuse apices,
This genus may readily be separated from others that have hitherto
been described by the numerous cilium-like appendages around the cen-
tral septum and by the carbonaceous spherical perithecia which have pro-
portionately very small ostioles.
Peritrichospora integra Linder, sp. nov.
Plate V, figs. 6-9.
Peritheciis solitariis vel sparsis, majorem ad partem immersis, globosis, 225-260 u
diametro, nigris, ostiolatis; ostiolis minutissimis, papillatis, apicalibus; ascis fusoideis
vel late fusoideis, 82-140 x 25-30 wu; ascosporis 8, fusoideis, biappendiculatis, hyalinis,
fuscis circum septum unicum centrale, pseudo-septis 2-6, multi-crinitis circum septum
centrale, crinibus haud longioribus dimidio ascosporae, cc. % y diametro ad basin,
sporis (appendicibus exclusis) 21,5-33 x 6.5-8.3 , appendicibus terminalibus hyalinis,
subcylindricis, 10-13.5 yp, longis, apicibus obtusis.
Perithecia solitary or scattered, partially immersed in the substratum,
globose, 225-260 » in diameter, black, ostiolate; the ostioles very small
and papillate, apical. Asci fusoid or broadly fusoid, 82-140 x 25-30 u.
Ascospores hyaline, 8, fusoid, with an elongate, 10—-13.5 » long, hyaline,
subcylindrical appendage at each end, l-septate and with numerous cilium-
like appendages just above and below the septum, the cilium-like ap-
pendages about half the length of the spore.
Specimen examined: Massachusetts, Woods Hole, on test board below
water at all times, immersed Mar. 6, 1941, removed Aug. 25, 1942. W.
F. Clapp, comm. E. S. Barghoorn, 10. Type.
BarcHoorn & LinpER: MARINE FUNGI 415
Peritrichospora lacera Linder, sp. nov.
Plate V, figs. 1-5.
Peritheciis solitariis vel sparsis, majorem ad partem immersis, globosis, plus minusve
338 p diametro, nigris, ostiolatis; ostiolis minutissimis, papillatis, apicalibus; ascis
fusoideis vel late fusoideis, plus minusve 75 x 22 ; ascosporis 8, fusoideis, biappen-
diculatis, hyalinis, fuscis circum septum centrale, 5-septatis, tarde multi-crinitis circum
septum centrale, crinibus haud longioribus cellula unica, cc. %4 y, diametro ad basin,
sporis (appendicibus exclusis) 41—53-(62.8) x 11.5-15 yp, appendicibus terminalibus
hyalinis, subcylindricis, 9.5-16.5-(21.5) w longitudine, 3.3-4 y diametro ad basin,
1.75 y, diametro ad apicem obtusum.
Perithecia solitary or scattered, about 24 immersed in the substratum,
globose, + 338 » in diameter, black, ostiolate; ostioles very small and
scarcely papillate, apical. Aseci fusoid or broadly fusoid, + 75 x 22 p.
Ascospores 8 in the ascus, fusoid, biappendiculate, hyaline, brownish
around the middle septum, 5-septate and tardily becoming polytrichous
around the central septum, the cilium-like appendages about equalling
the length of the spore cell and approximately 34-1 » in diameter at the
base, the spores (excluding the appendages) 41—53—(62.8) x 11.5-15 yp,
the terminal appendages hyaline, subcylindrical, 9.5-16.5-(21.5) yw in
length, 3.3-4 » in diameter at the base, 1.75 w» in diameter at the ob-
tuse apex.
Specimen examined: Massachusetts, Provincetown, on driftwood, two
feet above the low tide level, Sept. 24, 1942, E. S. Barghoorn, 28b. Type.
This species differs from the preceding in that the ascospores even while
within the ascus are clearly 5-septate, whereas in P. integra, the spores
become 1]-septate just before liberation from the ascus, and the two re-
sulting cells then have pseudosepta or annular thickenings of the cytoplasm
which indicate where septa may form, as they do when the spore germi-
nates. This species, P. lacera, produces the lateral, cilium-like appendages
only after the perispore (PI. V, figs. 4-5) has been ruptured and shed,
the perispore tearing at the first septum on either side of the central one
and then peeling off the spore and appendage after the fashion that a glove
is removed from the fingers of a person’s hand.
SCOLECOSPOREAE
HALOPHIOBOLUS Linder, gen. nov.
Peritheciis solitariis vel gregariis, late ampulliformibus, elongato-ellipsoideis vel
subcylindraceis, membranaceis vel subcarbonaceis, atris, fuscis, vel laete coloratis vel
supra fuscis et infra laete coloratis; ostiolis conoideis vel elongato-cylindraceis et
rectis vel undulatis; aparaphysatis, ascis clavatis vel elongato-clavatis, octosporis;
ascosporis hyalinis, filiformibus, vulgo non septatis sed interdum pluriseptatis, ad
extrema utraque appendicula hyalina, conoidea ornatis,
Perithecia solitary or gregarious, flask-shaped to elongate ellipsoid or
subcylindrical, membranous to subcarbonaceous, black, fuscous or light
colored, the perithecia often dark above and light colored below; the
ostiole conical to elongate and cylindrical, straight or undulate, deep
fuscous or black. Paraphyses absent. Asci elongate-clavate, 8-spored.
Ascospores hyaline, elongate and slender, cylindrical or filamentous,
416 FARLOWIA, VoL. 1, 1944
mostly non-septate at first and only becoming septate on germination (one
species 15—18-septate), provided at each end with a conspicuous hyaline,
conoid, or somewhat inflated and conoid appendage.
As the name implies, this genus, which resembles Ophiobolus, has to
date been collected only in marine habitats. It is distinguished from
Ophiobolus by the lack of paraphyses, by the predominantly membranous
perithecia, and by the characteristic hyaline appendage at each end of
the ascospore. The appendages, as already mentioned in the introduc-
tion, have an adhesive quality which enables the spores to become at-
tached to the substratum.
KEY TO THE SPECIES OF HALOPHIOBOLUS
A. Neck of ostiole longer than the height of the perithecium.
B. Ascospores 74-82.5 y, long; perithecia cylindrical and paralleling the sur-
face of the substratum: ... 46. 6.60 cees java dwseunn 1. H. cylindricus
BB. Ascospores more than 100 uw long.
C. Perithecial walls black, opaque, 11.5-21.5 y, thick; perithecia imbedded
Gr MMOMMMCI MN eh. ol oe eet oe ed eas ee Rae 2. H. opacus
CC. Perithecial walls subhyaline or light fuscous, (15)-16.5-30 py thick;
perithecia imbedded .......................00005- 3. H. longirostris
AA. Neck of ostiole shorter than the height of the perithecium.
D. Ascospores 15-18-septate and breaking into uniseptate segments. On Zos-
ee AIMEE eet ae ik nt foe wedi a ce eee eae 4, H. maritimus
DD. Ascospores at most pseudoseptate or only becoming septate on germination.
E. Tips of ascospores suddenly inflated and rounded-truncate; appendage
inflated and tapering to a slender flanged beak; on buried leaf-sheaths
SEE te leer hs ua ats cee Ree dpe valcaty Oe 5. H. medusa
EE. Tips of ascospores rounded or truncate, not inflated; appendages conoid
or elongate conoid.
F. Perithecial walls thick, blackish brown. On Zostera marina.
6. H. halimus
FF, Perithecial walls dark near the surface of the substratum, nearly hya-
line below and only tardily becoming black ...... 7. H. salinus
The type species is Halaphiobolus opacus.
1. Halophiobolus cylindricus Linder, sp. nov.
Plate VI, figs. 12-14.
Peritheciis immersis, cylindraceis vel subcylindraceis et parallelis superficie sub-
strati, plus minusve 762x123 yu, sub microscopio fuscis, membranaceis, cellulis ex-
tremorum duorum subquadratis vel angulatis, cellulis ceteris elongato-rectangulatis;
ostiolis atris, cylindricis, usque ad 550 wu longitidine, 24-27 y, diametro, plectenchy-
maticis, ad basem bulboso-amplificatis, pseudoparenchymaticis et 50-57 p diametro;
ascis deliquescentibus; ascosporis breviter filiformibus, hyalinis, curvatis vel nonnihil
spiraliter tortis, ad extrema utraque appendice hyalina, elongato-conoidea, 8.5-15 y.
longitudine ornatis.
Perithecia immersed in the substratum, cylindrical or subcylindrical,
averaging around 762 x 123 yp, fuscous under the microscope, membranous,
the cells at the two ends and around the swollen base of the ostiolar neck
subquadrate or angular (pseudoparenchymatous), the remaining cells
elongate rectangular or subcylindrical; the ostiole black, cylindrical, up
BARGHOORN & LINDER: MarInE FUNGI 417
to 550 » long, 24-27 » in diameter, plectenchymatous, arising from a
bulbous pseudoparenchymatous base 50-57 » in diameter. Asci soon
deliquescing and leaving the spores in clusters of eight. Ascospores
short filiform, hyaline, curved or somewhat spirally twisted, 7482.5 x 5 p,
terminated at each end by a hyaline, elongate-conoid appendage, 8.5-11.5 p»
long.
Specimens examined: Massachusetts, Provincetown Harbor, on upper
portion of stump, Sept. 24, 1942, E. S. Barghoorn, 23. Type.
This species, like the following one, is reminiscent of Linospora and
Robergea in that the asci, as shown by the fascicle of ascospores, parallel
the long axis of the perithecium, but the perithecia being without a clypeus
separates this species from the former genus, and lacking paraphyses, it
is excluded from the latter.
2. Halophiobolus opacus Linder, sp. nov.
Plate VI, figs. 1-5.
Peritheciis immersis vel erumpentibus, atris, subsphaericis vel ovoideis, 198-264 x
148-214 y, (in substrato immersis ellipsoideis vel subcylindraceis et parallelis superficie
substrati) ; ostiolis excentricis, cylindraceis vel nonnihil attenuatis, atris, usque ad
300 x 28-33 w; parietibus peritheciorum 11.5-16.5-(21.5) u crassitudine, strato intimo
e cellulis hyalinis vel subfuscis consistente, parietibus tenuibus; strato medio crasso,
pseudoparenchymatico, atro opacoque; strato externo hyphis elongatis subbullatis
composito e quibus hyphae atrae tenues oriuntur et in substratum penetrant; ascis
elongato-clavatis, curvatis, mox deliquescentibus; ascosporis filiformibus, rectis, curva-
tis, vel nonnihil spiraliter tortis, hyalinis, 155-190 x 3.3-4 wp, continuis, multivacuo-
latis, vel germinantibus 4~7 septatis, ad extrema utraque appendice hyalina, conoidea
instructis, 7.5-8.5 p. longitudine.
Perithecia occurring singly or scattered, superficial or immersed,
black, subglobose to ovoid, 198-264 x 148-214 » or, when immersed in
the substratum not infrequently cylindric-ellipsoid with the cylindrical or
somewhat tapering, elongate ostiole arising from one end, the ostiole up
to 300 x 28-33 p; walls of the perithecium 11.5-16.5—(21.5) u thick, the
inner layer composed of thin-walled, subhyaline to light fuscous cells,
the middle layer thick, pseudoparenchymatous and deep fuscous to black
and opaque; the outer layer of rather closely interlaced and somewhat
bullate hyphae which give rise to more slender dark hyphae that either
ramify in the substratum or, in superficial specimens, form a loose, hairy
covering. Asei when young are cylindric-clavate and curved, and at ma-
turity soon deliquesce to liberate the eight ascospores. Paraphyses ab-
sent. Ascospores filiform, straight, curved, or somewhat spirally twist-
ed, hyaline, 155-190 x 3.3-4 y, non-septate and multivacuolate until
germination when they may become 4~7-septate, ending in hyaline conoid
appendages, 7.0-8.5 pw long.
Specimen examined: Massachusetts, Provincetown, on piling two feet
above low tide level, July 24, 1942, E. S. Barghoorn, 14. Type.
In addition to the above perfect stage that was found occurring in
nature, this fungus when grown on artificial media, produces chains of
spores which may attain a length of 350-400 yp, the individual cells meas-
uring (11.5)—16.5-21.5 x (8)-13-16.5 » with the central cells larger than
418 Fartowia, Vou. 1, 1944
the terminal. Occasionally the chains of cells are furcate at the base,
rarely branched above. As a rule the cells are at first hyaline but with
age they become brownish. These chains of spores have their counter-
part within the agar as intercalary chains of varying length and in the
natural substratum as single bladder-like cells or as short chains of two
to five cells which are usually globose but when their diameter reaches
the limit of size set by the diameter of the lumen of the tracheids, become
elongate and ellipsoidal.
3. Halophiobolus longirostris Linder, sp. nov.
Plate VI, figs. 6-7.
Peritheciis semper in substrato immersis, solitariis, subhyalinis yel dilute fuscis,
globosis vel late ellipsoideis, ostiolis mediis, atris, elongatis subrectis vel irregulariter
curvatis, 594-700 x 41-58 u.; parietibus peritheciorum (15)-16.5-29.5 y. crassitudine ;
strue intima hyalina cellularum isodiametricarum compacte aggregatarum; strato
medio crasso, hyalino, e cellulis aliquantulum rhomboideis efformato; strato ex-
terno tenui subhyalino vel dilute fusco, e cellulis cylindricis nonnihil bullatis con-
sistente; ascis elongato-clavatis, mox deliquescentibus; ascosporis filiformibus, rectis,
curvatis vel nonnihil spiraliter tortis, hyalinis, 155-200 x 3.3—4 uw, aseptatis, ad extrema
utraque appendice hyalina, conoidea, 7-8.5 y. longitudine ornatis.
Perithecia always (?) immersed and occurring singly or scattered,
subhyaline or dilute fuscous, globose or subellipsoid, the ostiole central,
black, elongate, nearly straight or else somewhat geniculately bent as it
penetrates to the outside of the woody substratum, 594-700 x 41-58 yp,
the external hyphae here and there growing out into the cells of the wood
and filling them with dark plectenchymatous hyphae; walls of the peri-
thecium (15)—16.5-29.5 » thick, the inner layer hyaline of closely aggre-
gated nearly isodiametric cells with rich protoplasmic content, — the cen-
tral layer thick, hyaline and composed of thin-walled, inflated, somewhat
diamond-shaped cells, — the outer layer 1-2 cells thick and of slightly
inflated, subhyaline to dilute fuscous subcylindrical cells among which
are often enmeshed the crushed, discolored cells of the substratum. Asci
when young cylindric-clavate, curved, at maturity soon deliquescing to
leave the spores in bundles of eight. Ascospores filiform, straight,
curved or somewhat spirally twisted, hyaline, 155-200 x 3.3-4 p, non-
septate, terminated by a hyaline conoid appendage, 7.0-8.6 » long at
each end.
Specimen examined: Maine, below low tide level at all times, on test
block immersed Sept., 1938, and removed Nov. 27, 1942, W. F. Clapp,
communicated by E. S. Barghoorn, 22.
Older specimens, because of the darker color, resemble H. opaca de-
scribed above. However, as shown in plate VI, figures 3 and 7, the struc-
ture of the perithecium wall of the two species is quite distinct, this species
having large hyaline, diamond-shaped cells in the central layer, while
H. opaca has a relatively thin, opaque black central layer.
Within the cells of the substratum, this species also produces bladder-
like cells either singly or in short chains, their size being limited by the
diameter of the lumen of the tracheids.
BarcHoorN & LINDER: MARINE FUNGI 419
4. Halophiobolus maritimus (Sacc.) Linder, comb. nov.
Ophiobolus maritimus Saccardo, Syll. Fung. 2: 350. 1883; Mounce, I. & W.
W. Diehl. A new Ophiobolus on eelgrass. Canad. Journ. Res. 11: 245. 1934.
This species was described as occurring on Zostera marina in the North
Sea. The septation and segmentation of the ascospores distinguish it
from other species of the genus.
5. Halophiobolus medusa (Ell. & Ev.) Linder, comb. nov.
Ophiobolus medusa Ellis & Everhart, Journ. Myc. 1: 150. 1885; Mounce, I. &
W. W..Diehl. Canad. Journ. Res. 11: 243-245. figs. 2, 6c-d. 1934.
The species was described as growing on the buried leaf-sheaths of
Spartina in New Jersey. The habitat would suggest that this fungus is
also a salt or brackish water inhabitant, and like many of the other species
mentioned in this paper, may well be a cellulose destroying form.
6. Halophiobolus halimus (Mounce & Diehl) Linder, comb. nov.
Ophiobolus halimus Mounce & Diehl. Canad. Journ. Res. 11: 242-246. figs.
1, 3-5, 6a-b, 7-9. 1934.
Like H. maritimus, this species was found on Zostera marina, but in
New Brunswick where it was thought to be a causative agent in the wasting
disease of eelgrass. As stated in the introduction, the discovery of Laby-
rinthula over a more widespread area suggests that H. maritimus may be
secondary or even saprophytic. At the same time, the need for further
research on the réle of this species in connection with wasting disease
is indicated.
7. Halophiobolus salinus Linder, sp. nov.
Plate VI, figs. 8-11.
Peritheciis in matrice immersis vel substrato eroso liberatis, subglobosis vel late
rotundate ampulliformibus, atris (fuscis sub microscopio) vel supra atris fuscisve,
infra pallide fuscis, 275-300 » diametro; ostiolis breviter et late conoideis vel papil-
latis; parietibus e cellulis amplis angulatis consistentibus; ascis primum fusoideis et
ad apicem rotundatum attenuatis, deinde elongato-clavatis et mox deliquescentibus,
octosporis; ascosporis hyalinis, filiformibus, curvatis vel nonnihil spiraliter tortis,
3.3 » diametro in parte centrali, ad extrema attenuata 1.5 yw diametro, et utrinque
appendice hyalina, conoidea, 7.5-8.5 py. longa ornatis.
Perithecia single or scattered, imbedded in the matrix or exposed by
erosion of the surrounding decayed wood, membranous and collapsing on
drying, subglobose to broadly rounded flask-shaped, black (fuscous under
the microscope) or black above and dilute fuscous below, 275-300 p» in
diameter; the ostiole short and broadly conoid or papillate; the walls of
the perithecium composed of large, relatively thin-walled angular cells.
Asci when young broadly fusoid and tapering to a rounded tip but when
mature elongate-clavate and soon deliquescing to liberate the 8 ascospores.
Ascospores hyaline, filiform, curved or slightly spirally twisted, 180-
200 » long, 3.3 « diameter at the middle and tapering to 1.5 » in diameter
at the base of the hyaline, conoid appendages, 7.5-8.5 » long which
terminate either end of the spores.
420 Fartowl1a, Voi. 1, 1944
Specimens examined: New Hampshire, Portsmouth, on test block four
feet below low tide level, submerged Sept. 23, 1941, removed May 19,
1942, W. F. Clapp, communicated by E. S. Barghoorn, 1d; Portsmouth,
on test block below water at all times, immersed Dec. 29, 1941, removed
Aug. 25, 1942, W. F. Clapp, communicated by E. S. Barghoorn, 11;
Massachusetts, Fall River, on test board eight feet below low water, im-
mersed Feb. 1942, removed July 18, 1942, W. F. Clapp, communicated
by £. S. Barghoorn, 4; Provincetown, on submerged stump three feet
above low tide level, Sept. 24, 1942, E. S. Barghoorn, 26; Woods Hole,
on manila rope in intertidal zone, E. S. Barghoorn, 20; Connecticut,
Saybrook, on test board below low water level at all times, immersed May
1, 1942, removed Oct. 1, 1942, W. F. Clapp, communicated by 2... &.
Barghoorn, 17.
This species, perhaps the commonest of our eastern coast, strongly re-
sembles H. halimus as regards the shape and size of the perithecia. How-
ever, it differs from that species by the color of the perithecial walls which
only tardily become all dark, and by the shorter length of the ascospores
which in H. halimus are 260-308 4 long.
Harvarp UNIVERSITY
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422
FarLowla, Vor. 1, 1944
EXPLANATION OF PLATE I
All figures are drawn with the aid of a camera lucida. Each unit of scales B, C
and E equals 10 wu; the unit of scale D equals 200 Ube
Figs. 1-3. Diplodia orae-maris Linder.
Fig.
1. Pycnidium shown with the substratum dissected from one side. The upper
surface is black and the lower surface is light colored. (>< 25, scale D)
2. Characteristic hyaline conidiophores, one bearing an immature conidium,
arising from the somewhat inflated cells that line the greater part of
the interior of the pycnidium. (> 800, scale E)
3. A representative group of colored, two-celled pycnidiospores. (>< 800,
scale E)
4. Botryophialophora marina Linder.
4. A portion of the repent or scandent mycelium bearing two clusters of
phialides, each of which arises from a single basal cell. The globose
conidia are produced semi-endogenously. (X< 1000, scale C)
Figs. 5-6. Orbimyces spectabilis Linder.
5. Characteristic conidiophores anastomosing at the point where they cross and
bearing above on a short hyaline sporogenous tooth the characteristic
appendaged conidium. (X 425, scale B)
6. Two shining black conidia with their fuscous appendages. The lighter col-
ored spot on the lower part of each conidium represents the place of at-
tachment of the sporogenous teeth. (> 425, scale B)
Figs. 7-8. Phialophorophoma litoralis Linder.
7. A representative group of pycnidiospores. (>< 1000, scale C)
8. Pycnidium shown with the substratum dissected from one side. The surface
cells of the substratum have been darkened through the action of the
fungus. (> 25, scale D)
9. Phialides, of which the middle one of the three right hand phialides is shown
with a pycnidiospore escaping from the cup-shaped terminal portion. On
left is shown a_ well eee phialophore bearing three phialides.
(< 1000, scale C)
BarcHoorn & LINDER: MARINE FUNGI 423
PiateE |
424, Fartowia, VoL. 1, 1944
EXPLANATION OF PLATE II
All figures are drawn with the aid of a camera lucida, and are reproduced at a
magnification of 425. Each unit of the scale equals 10 Us
Fig. 1. Alternaria maritima Sutherland.
1. a. The short, slightly colored, inconspicuous conidiophores bearing branching
chains of conidia.
b. Single conidium with an exceptionally long apical cell.
Figs. 2-6. Helicoma maritimum Linder.
Showing conidiophores and conidia and their variation in shape and dimen-
sions. Fig. 6 depicts the more irregular type of conidium that is found in
cultures while the remaining figures illustrate the more or less squarish
outlines that appear more characteristic of the fungus as it occurs in nature.
Fig. 7. Speira pelagica Linder.
A group of four conidia, of which the upper right hand one illustrates the
variation that may be observed in the older conidia.
Figs. 8-11. Helicoma salinum Linder.
Conidiophores and conidia drawn to show the variations of the fungus as it
grows in culture. Fig. 11 shows the conidiophore with the dark, inflated
terminal cell upon which the conidium is directly borne, a character that
appears when the fungus is cultured on agar. »
425
BarcHoorn & LINDER: MARINE FUNGI
Pirate II
426
Fartowia, Voi, 1, 1944
EXPLANATION OF PLATE III
All figures are drawn with the aid of a camera lucida. Each unit of scales A, B
and C equals 10 y; the unit of scale D equals 200 Ue
Figs.
Figs.
Figs.
Figs.
1-4. Samarosporella pelagica Linder.
1. The globose, black, carbonous perithecium shown with half the substratum
dissected away. (> 25, scale D)
2. Asci with ascospores nearly delimited but clearly showing one aborted spore
(left), and an ascus with nearly mature ascospores (right). (> 350,
scale A)
3. An abortive ascospore with an abortive wing. (> 1000, scale C)
4. Two ascospores showing variation in the size of the hyaline winged ap-
pendages. (> 1000, scale C)
5-9. Remispora maritima Linder.
5. A subglobose, cream-colored perithecium showing the lateral position of the
conoid ostiole which protrudes beyond the substratum. ( 25, scale D)
6. Three ascospores, of which the lowest one still retains the characteristic
mustache-like hyaline appendage. ( 425, scale B)
7. An ascus with three differentiating spores, of which one is small and some-
what aborted. (>< 425, scale B)
8. An ascus with mature ascospores. (>< 425, scale B)
9. a. An ascospore with appendages still connected at the ends and including
a remnant of the ascus protoplasm. (>< 1000, scale C)
b. An ascospore still showing remnants of the ascus protoplasm attached to
the appendages. (> 1000, scale C)
c. An ascospore with characteristic appendages. (>< 1000, scale C)
10-12. Ceriosporopsis halima Linder.
10. A black, elongate-ellipsoid perithecium dissected from the substratum. (X 25,
scale D)
11. A group of four ascospores of which the two central ones have lost the deli-
quescent hyaline appendages. (X 425, scale B)
12. An ascospore shown at a greater magnification to illustrate the large central
vacuoles. (>< 1000, scale C)
13-16. Amphisphaeria maritima Linder.
13. A nearly superficial perithecium as seen from above. (> 25, scale D)
14. Four dark-colored, uniseptate ascospores. (> 425, scale B)
15. An immature ascus showing the ascospores, already becoming septate, nearly
differentiated. (> 425, scale B)
16. Two asci with distichously arranged ascospores. (>< 425, scale B)
427
MaRINE FUNGI
.
°
BaRGHOORN & LINDER
Pirate III
428 FarLowia, VoL. 1, 1944
EXPLANATION OF PLATE IV
All figures are drawn with the aid of a camera lucida. Each unit of scales C and
E equal 10 »; the unit of scale D equals 200 y.
Figs. 1~3, Lentescospora submarina Linder.
1, At the left, a perithecium partially exposed and showing its position in the
substratum; at the right, a perithecium as seen from above. (x 25,
scale D)
2. The ostiole and adjacent cells as seen from above. (> 800, scale E)
3. Four ascospores to illustrate variations in the shape, size, and the thickening
of the apical walls. (> 800, scale E)
Figs. 4-7. Halosphaeria appendiculata Linder.
4. Two perithecia as seen from the side and the end, showing the blackened
upper part and the lighter colored under portions. Note the stout, short-
cylindrical and obliquely attached ostiole. (X25, scale D)
5. Three ascospores to show the variation in number and attachment of the
appendages. (>< 800, scale E)
6-7. Mature and immature asci. (>< 800, scale E)
Figs. 8-11. Leptosphaeria orae-maris Linder.
8. A broadly ovoid, immersed perithecium which is dark fuscous above and
lighter colored below. (> 25, scale D)
9. A thick-walled immature ascus at the left and a thinner-walled mature ascus
at the right containing 0- to l-septate ascospores. (>< 800, scale E)
10. Four ascospores with colored walls and with one to three septa. The septa.
in excess of the median one are usually formed after the spores are dis-
charged from the ascus. (> 800, scale E)
11. A typical branched paraphysis. (> 800, scale E)
Figs. 12-15, Sphaerulina orae-maris Linder.
12. A partially imbedded perithecium, dark-colored where exposed and light-col-
ored where buried in the substratum. (>< 25, scale D)
13. A group of four ascospores. (> 1000, scale C)
14, A mature ascus with eight spores. (>< 1000, scale C)
15. An immature ascus drawn to show the thickened wall. (> 1000, scale C)
429
MarRINE FUNGI
& LINDER
BaRGHOORN
Priate IV
430 FARLOWIA, VoL. 1, 1944
EXPLANATION OF PLATE V
All figures are drawn with the aid of a camera lucida. Each unit of scales B and
C equals 10 y; the unit of scale D equals 200 Ue
Figs. 1-5. Peritrichospora lacera Linder.
1, A large subglobose, shining-black, carbonous perithecium which is partially
buried in the substratum and which produces but a small inconspicuous
ostiolar neck. (> 25, scale D)
2. A cluster of eight ascospores liberated by the dissolution of the ascus wall.
(X 425, scale B)
3. Two ascospores, of which the right-hand one has just begun to form cilia
around the central septum and the left-hand one with elongate and now
more slender cilia. (X< 425, scale B)
4-5. Young ascospores with the ruptured exospore which breaks away from the
cells and peels off with the appendages making them appear longer than
they actually are. These spores are younger than those shown in figure 3.
(X 1000, scale C)
Figs. 6-9. Peritrichospora integra Linder.
6. A partially immersed, shining-black perithecium with the characteristic small
ostiole. (> 25, scale D)
7. Two asci containing immature ascospores. The ascus on the left is the more
mature. (> 1000, scale C)
8. Two uniseptate ascospores showing the coloring of the walls at the septa,
the vacuolate cell contents that give the appearance of additional septa,
and the long, hyaline appendages. (>< 1000, scale C)
9. Conidial chains formed in culture. (>< 425, scale B)
431
Marine FUNGI
BarcHoorn & LINDER
Z
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Pree es
PLATE V
432 FaRLowiA, VoL. 1, 1944
EXPLANATION OF PLATE VI
All figures are drawn with the aid of a camera lucida. Each unit of scales B and C
equals 10 1; the unit of scale D equals 200 Ue
Figs. 1-5. Halophiobolus opacus Linder.
1. Perithecium as seen in sectional view. The perithecium has become super-
ficial through the erosion of outer portions of the substratum. (x 25.
scale D)
2. A portion of the perithecial wall to show the black and nearly opaque middle
layer bordered internally by elongate colored cells, and externally by the
thin layer of cells which give rise to mycelium that normally penetrate into
the substratum. (>< 1000, scale C)
3. A perithecium with part of the substratum dissected away. The outer
mycelial threads have also been removed. (> 25, scale D)
4. A typical ascospore with hyaline, sticky terminal appendages. (X 425,
scale B)
5. A very immature ascus. (x 425, scale B)
Figs. 6-7. Halophiobolus longirostris Linder.
6. A perithecium as seen in sectional view to show the relatively thick sub-
hyaline wall and the long ostiolar neck from which the mycelium has here
and there grown out to fill adjacent cells of the substratum. (x 25,
scale D)
7. A section of the perithecial wall showing the thin inner layer of slender,
elongate protoplasm-filled cells, the thick middle layer composed of angu-
lar, enlarged, subhyaline cells, and the thin outer layer of darkened and
closely intertwined cells. (>< 1000, scale C)
Figs. 8-11. Halophiobolus salinus Linder.
8. Two perithecia to illustrate the variation in size and shape. The upper por-
tions are dark fuscous or nearly black and shading off to buff-colored
below. (> 25, scale D)
9. Three characteristic ascospores. (>< 425, scale B)
10. Detail of the hyaline terminal appendages of which that on the left has
begun to deliquesce. (> 1000, scale C)
11. Chain of conidia formed in culture on agar. (> 425, scale B)
Figs. 12-14. Halophiobolus cylindricus Linder.
12. A perithecium showing the subcylindrical shape, the swollen base of the
long protruding ostiolar neck. (> 25, scale D)
13. Two ascospores with characteristic elongate, tapering, hyaline appendages.
— (XK 425, scale B)
14. A terminal appendage more highly magnified. (1000, scale C)
433
Marine FUNGI
BaRGHOORN & LINDER:
PuateE VI
434, FarLowia, VoL. 1, 1944
II. BIOLOGICAL ASPECTS
E. S. BARGHOORN
Microbiological studies in recent years have clearly demonstrated the
significance of bacteria as active agents in many of the decomposition
processes which occur in submerged plant remains (Waksman, 1936;
Waksman and Stevens 1929, Waksman et al. 1933, Allgeier et al. 1932).
The possible réle of fungi in such processes in the sea has received little
attention, however, except in the case of certain fungal parasites (Spar-
row 1936, Renn 1936). In the degradation of organic matter in the sea
the actual existence of saprophytic truly marine fungi has been seriously
questioned by certain authors (Waksman 1934). Indeed, it is commonly
assumed that the lower availability of oxygen in conjunction with a less
vigorous fungal flora render unlikely any considerable fungal action on
organic matter at appreciable depths under water. The depletion of avail-
able oxygen by saprophytic bacteria and other microérganisms working on
the surfaces of decaying organic detritus would a priori support the view
that fungal action is greatly reduced if not inhibited under such conditions.
A survey of the literature, however, reveals a paucity of information
concerning the microbiological, microchemical and histological changes
which occur in submerged plant materials, particularly wood, which have
been immersed in salt, brackish or fresh water for considerable time.
Not only are the histological changes resulting from the decomposition
processes incompletely known, but the chemical composition of specific
plant constituents in bottom deposits is not well understood.
Recent microchemical and anatomical study of certain submerged woods
of recent geological age indicate that the relative significance of purely
chemical and of microbiological factors in decomposition is difficult to
determine (Bailey and Barghoorn 1942). In the latter investigation wood
buried deeply in estuarine silt for presumedly not less than 2000 years
was found to have undergone very extensive chemical degradation involv-
ing the loss of 95 per cent of its original cellulosic constituent and the
major portion of its pentosan complexes but with little or no change in
lignin content (Jahn and Harlow 1942). Histological and microchemi-
_ cal examination failed to reveal the presence of microérganisms inducing
decay, although the fungal hyphae of terrestrial wood-rotting forms in-
fecting the wood before it was submerged were well preserved and
easily demonstrated.
Non-mineralized fossil woods of far greater antiquity such as certain
deeply buried coniferous remains from the auriferous gravels of Cali-
fornia, on the other hand, show a remarkable preservation of both physi-
cal and chemical structure (Mitchell & Ritter 1934, Barghoorn and Bailey
1938). In material such as this, although of Miocene geological age,
it is evident that neither microbiological nor extensive chemical action
has occurred.
BarcHoorn & Linper: Marine Func 435
Many deposits of greatly-altered plant materials, such as lignite, brown
coal and certain peats are highly modified both chemically and physically
and yet exhibit little or no indication in most cases as to the influence of
microdrganisms inducing specific modifications, although the initial
changes are usually interpreted as biological in origin. (Waksman 1936,
Thaysen 1930).
A large proportion of plant materials which are preserved either as
isolated fossils, or in extensive deposits of varying thickness, has been
exposed at one stage or another to the conditions of submergence in salt,
brackish, or fresh water. It is desirable, therefore, to investigate the
initial changes which occur in plant materials upon continued exposure
to these conditions in order to establish the first steps in the long sequence
of events which lead to the ultimate preservation, degradation or com-
plete decomposition.
A convenient and significant starting place for such studies has been
found in the changes which occur in plant materials exposed to the sea
and variously modified conditions of the marine environment.
It is well known, although perhaps not as widely recognized as might
be expected, that plant materials of economic importance such as wood
and cordage undergo a steady deterioration in sea water which cannot
be assigned to the activities of marine borers, or other similar organisms.
In the case of wooden pilings the obvious depredations of Teredo and
Limnoria are well known. However, a conspicuous softening occurs in
the outer parts of piling and submerged wood which clearly does not
result from the activity of these wood destroying marine animals. Sim-
ilarly, cordage, particularly of untreated fiber, quite rapidly deteriorates
upon continuous exposure to sea water and less rapidly with intermittent
submergence. The often quoted “preservative” effect of sea water is
certainly not to be noted in materials constantly exposed to marine con-
ditions, although the salt treatment of timbers, effected by drying after
prolonged immersion in sea water, does serve to retard the activities of
terrestrial wood rotting organisms as well as the development of the air-
borne spores of fungi which normally attack damp, readily available
organic materials such as wood and its various constituents.
In an early phase of these investigations on the decomposition of wood
submerged in sea water it became evident that fungi are extremely com-
mon in such materials under aerobic conditions, though totally absent
under anaerobic conditions. Accordingly contemplated bacteriological
studies in the decay of submerged wood were set aside and efforts made
to isolate and culture the fungi, which comprise a far more conspicuous
and vigorous element of the microflora involved in the decomposition.
The occurrence of abundant perithecia and ascospores of many species
of the fungi facilitated the isolation of pure cultures. Other forms have
been brought into culture from their conidial stages. Certain species of
undoubted marine occurrence have been found only in their vegetative
436 Fartowla, VoL. 1, 1944
phase in decaying cordage and wood. Experimental studies on the bi-
ology of fourteen species of this interesting group of fungi have been made,
other forms having been isolated more recently. The present paper deals
in particular with the physiological behavior of seven representative
species, all of which have been isolated by spore culture from their
ascospore stages.
MATERIALS AND METHODS
The fungi with which this study is concerned were obtained from speci-
mens of wood or cordage exposed for varying lengths of time to the full
salinity of sea water or to the brackish waters of estuaries. In general,
two types of materials were used, viz., specimens of wood or cordage of
known conditions and duration of submergence, and specimens of wood
and cordage of undetermined length of exposure. In all cases the fungi
were isolated from a natural substratum which was either continuously
submerged in water or covered for the greater part of the interval elaps-
ing between successive high tides. A summary of the specimen collection
data is included in the introduction.
The “test” boards and blocks referred to in this paper were supplied
through the courtesy of the William Clapp Laboratories in Duxbury,
Mass. These blocks, chiefly of fresh-sawed hard pine, had been exposed
for six to twelve months at a depth of two to eight feet below low water
and had not been exposed to the air from the time of their submergence
until the date of removal. Abundant perithecia on the surface of or em-
bedded in many of the blocks provided an excellent source of material
for spore isolations. The majority of the blocks were examined and used
as sources of inoculum within three or four days after their removal
from the water. This brief period of time precludes the possibility of
perithecium production by any contaminating terrestrial fungi. Further
proof that perithecium production is commonly vigorous under sea water
was obtained by sectioning specimens of wood freshly collected from
Cape Cod Bay and Provincetown Harbor. Sections of such material,
bearing perithecia with viable spores, also show the presence of a very
extensive and vigorous mycelium in the wood. Comparable material has
recently been obtained from Chesapeake Bay, Maryland, and Biscayne
Bay, Florida.
In addition to wood, specimens of submerged rope and cordage were
examined for attack by fungi. All rope samples which have been studied
whether of treated or untreated material show varying degrees of infec-
tion and deterioration by fungi. In cases in which perithecia occurred
on or embedded in the rope the species have been found identical with
those obtained from wood blocks. Although studies are now in progress
on the mode and rate of attack of the marine fungi on both wood and
cordage it is desirable to point out here that rope, because of its fibrous
texture, appears to be more readily attacked than wood by the cellulose
destroying members of the group.
BARGHOORN & LINDER: Marine FUNGI 437
In addition to the collection data tabulated in the introduction the fol-
lowing locations along the Atlantic Coast have yielded specimens of wood
(or cordage) bearing perithecial stages of marine fungi:
Newfoundland Neponset
Corner Brook Quincy
: Vineyard Haven
Maine Weymouth
ee Connecticut .
Portland South Groton
Rockland New York
Wiscasset City Island
Massachusetts Rosebank, L. I.
Beverley Maryland
ree Gibson Island,
acne Chesapeake Bay
Edgartown :
Gloucester Florida
Marshfield Biscayne Bay
In isolating certain of the fungi from the natural substratum great
difficulties were encountered in freeing the germinating spores from bac-
teria. The poured plate method using nutrient agar made up in sea water
usually proved useless owing to the very rapid spread of motile marine
bacteria which frequently destroyed the spores even before their germina-
tion occurred. Modifications of the sprayed-spore technique (Wehmeyer
1923) were likewise of little value. The acidification of the spore sus-
pension to retard bacterial activity completely inhibited germination of
the fungal spores. The best results were obtained with stab cultures
made by detaching a small mass of spores from an exuding or crushed
perithecium and thrusting it with a streaking motion deep into agar. The
spores, germinating beneath the surface of the agar, rapidly developed a
bacteria-free mycelium. In certain cases, however, the rapid spread of
bacteria on the surface of the agar made it necessary to invert the stab-
culture plates, free the agar from the dish and cut blocks from the lower
surface in order ultimately to free the fungus from bacteria. The ex-
cessive motility of certain of the bacteria, in conjunction with a definite
agar-liquefying activity, permitted their development along agar-glass
interfaces, although they were unable to penetrate deeply into free agar.
In making the stab cultures, therefore, the needle was not allowed to pene-
trate completely through the agar layer. This precaution prevented the
spread of bacteria on the lower surface of the culture. The medium most
useful for spore germination with a minimum of bacterial growth was a
three per cent crude unwashed agar made up in sea water.
In determining the reaction of the fungi to the substratum, to tempera-
ture, and to hydrogen-ion concentration various agars were used. All of
the media, with the exception of the fresh water and concentrated sea
water, were solidified with three per cent Bacto agar made up in filtered
sea water with added nutrient.
438 Fartowia, VoL, 1, 1944
Stock cultures were maintained on a four per cent agar containing 0.5
per cent malt extract. Other agars were made up as follows:
Cellulose agar. One per cent regenerated cotton cellulose prepared by
the method of Northrup (1919), pH 7.47.
Maltose, Xylose and Galactose agars. Sugar concentration two per
cent, pH 7.4.
Starch agar. Two per cent purified corn starch, pH 7.5.
Asparagine agar. Four-tenths per cent asparagine Merck, pH 7.2.
Pectin agar. Five-tenths per cent lemon pectin purified by the method
of Baxter (1925) pH 6.5. The pH of the pectin agar was adjusted to
7.2 before use by the addition of dilute sodium hydroxide. This medium
doubtless contains many degradation products of pectin hydrolysis.
Wood flour agar. 0.75 per cent finely powdered white pine filings.
The wood dust was extracted with boiling chloroform, and repeated treat-
ments with benzol ‘and absolute alcohol to remove, insofar as possible,
the resins and terpenes. The treated wood was washed repeatedly with
boiling distilled water and dried on a Buchner funnel. pH 7.4.
Fresh water agar. Glass distilled water four parts, tap water one part.
pH 7.2. Tap water contained traces of organic matter in addition to
traces of inorganic salts.
Sea water agar. Filtered sea water with no added nutrient, pH 7.4.
Concentrated sea water agar. Prepared from sea water evaporated to
exactly one-third its original volume, with no added nutrient. A portion
of the calcium precipitated out during evaporation. pH 8.2.
The agar used for the study of the influence of temperature was made
with four-tenths per cent malt extract and no additional nutrient. The
experiments on the influence of hydrogen-ion concentration were run with
five per cent malt extract agar; the medium was acidified by the addition
of hydrochloric acid and made alkaline by addition of sodium hydroxide.
In studying the rate of radial growth of the fungi in culture a more or
less standardized procedure was followed. Pure cultures of the various
species were maintained by repeated transfers on a stock culture medium
in Petri dishes. Small blocks of this medium, one to two millimeters
square were used as inoculum in the various growth studies. The blocks
were cut from the outer third of the spreading mycelium. Rate of radial
growth was determined by measuring the spread of the mycelium on Petri
dish cultures containing the different media. After inoculating the plates
the colonies were allowed to grow for three to six days and then measured.
Measurements were made radially in three directions from the edges of
the inoculum to the outer extreme of the mycelial mat. The plates were
then incubated for a varying length of time, in most cases six to ten days,
and measured again. The rate of radial growth was taken as the increase
in radius of the colony in millimeters per day between the first and second
measurements. The value for any one measurement was obtained by
averaging the three readings.
*pH values are for media after sterilization.
BarcHoorn & LInDER: MARINE FUNGI 439
In eliminating from measurement the growth for the first three to six
days, the influence of nutrient carried over in the inoculum was greatly
reduced.
Although radial increase of a colony is a rather arbitrary index of the
amount of growth of a mycelium it is by and large a more practical means
of measurement than determinations of dry weight increase. As far as
was practicable in this study the radial increase of the colony was roughly
correlated with the apparent visible density of the fungal mat. In all
but a few cases extensive radial spread was quite closely associated with
abundant branching and production of hyphae. The cases in which this
correlation did not obtain, were those of “starvation media” such as the
fresh water and pure sea water cultures. Certain species on these two
media produced extremely sparse but far-ranging mycelia. These facts
must be taken into account in considering the rate of growth on certain
of the media deficient in available nutrient.
In preliminary experiments to determine the availability of different
carbohydrates it was found that the depth of medium in the culture dish
exerted considerable influence on the rate of spread of the mycelium.
Certain species spread more rapidly on deep cultures while others spread
more rapidly on shallow cultures. In order to counteract this effect which
is particularly significant in the study of the influence of temperature and
hydrogen-ion concentration, the plates were poured to as uniform depth
as possible and then sorted before inoculation so that a given species was
grown on media of the same depth.
The growth studies in this investigation were made with fourteen se-
lected species of fungi obtained either by spore isolation or direct inocula-
tion from infected material. Inasmuch as the latter source of cultures
always presents the uncertainty of possible contaminating organisms not
originally present in the material, only species obtained from spore isola-
tions are considered in detail. Of these, seven species were selected as
representative of the marine fungi as a group. The seven species cover
fairly well the morphological range of the forms known at present and
probably represent fairly well the physiological diversity of this interest-
ing group of fungi.
RELATION TO SALINITY
As indicated by the collection data summarized in the introduction, the
marine fungi occur naturally in both brackish water and sea water of
normal salinity. In view of their undoubted derivation from terrestrial
forms it seemed interesting to determine their reaction in culture to ex-
tremes of salinity as some measure of their adjustment to the marine
environment. For this purpose five species were chosen and grown on
three different media: fresh water, normal filtered sea water and sea water
evaporated to one-third its original volume. The three series of experi-
ments were run in duplicate, simultaneously and at the same temperature.
440 FarLowliA, VoL. 1, 1944
TABLE 1, EFFECT OF SALINITY ON RATE OF RADIAL GROWTH
Rapist Growrs in Miuumerers Per Day
Fresh water Sea water Concentrated
pH 7.2 pH 7.4 Sea water pH 8.2
Amphisphaeria maritima .55 .80 12
Helicoma salinum .68 1.14 31
Peritrichospora integra 79 .96 88
Ceriosporopsis halima 1.33 1.17 54
Halophiobolus opacus 52 3.10 .07
Halophiobolus salinus 2.74 3.16 07
In Table 1 it will be noted that two species, Amphisphaeria maritima
and Ceriosporopsis halima develop more rapidly in fresh water than in sea
water, while the remainder grow radially at a more rapid rate in normal
sea water. In concentrated sea water, the growth rate for all is consider-
ably decreased although Ceriosporopsis halima and Peritrichos pora integra
grow at nearly one-half the rate determined in fresh water. An important
factor in the experiment which is not expressed in the table is the relative
denseness of the mycelia. In all cases the hyphal production per colony
area was conspicuously greater in normal sea water than in either fresh
water or concentrated sea water. In fresh water agar the hyphal mats
were more hyaline and the hyphae less branched, particularly in the outer
third of the colony. In concentrated sea water the hyphae branched ab-
normally, the secondary hyphae being crooked and much shortened. In
this latter medium it might be presumed that the higher pH value would
adversely affect growth. However, as will be shown later, a pH of above
8.0 is more favorable for growth in culture than a pH below 8.0 in the
case of at least three species.
Although Amphisphaeria maritima and Ceriosporopsis halima in the
above experiments showed a more rapid radial growth in fresh water than
sea water, neither of these species shows preference for such conditions in
nature as far as collection data and field study indicate. Indeed, their
occurrence in fresh water is to be doubted, although further collections
may demonstrate their presence in the non-saline portions of estuarine
rivers. Of the five species studied in respect to salinity reactions all but
Halophiobolus opacus and Peritrichospora integra have been collected
from brackish waters as well as the open sea.
Two species of the marine fungi, Halophiobolus opacus and Remispora
maritima were collected in conditions in which a fairly high salt content
presumably occurred in the natural substratum. Both of these forms were
found, in the perfect stage, on wooden pilings in such a situation as to
be exposed to the drying action of wind and sun for many hours at low
tide, although completely submerged at high tide. Remispora maritima
in particular was collected on wood exposed to the air for six to seven
hours between tides. Such conditions would suggest a high tolerance to
salinity as well as to sudden changes in salinity such as would occur with
recurring incoming tides,
BarcHoorn & LInDER: MARINE FUNGI 441
In general, considering both ecological factors and the results from
experimental cultures, there is considerable evidence that the marine
fungi possess an adaptation to or tolerance of the saline conditions in
the environment of the sea.
RELATION TO SUBSTRATUM
In view of the natural occurrence of the fungi on wood, rope and other
plant materials of similar chemical composition, a series of experiments
were run to determine the relative availability of certain carbohydrates.
For this purpose the sugars maltose, xylose and galactose and the carbo-
hydrates cellulose, starch and pectin were employed. An artificial medium
containing wood in a finely divided state was also tried in an effort to
establish in a rough quantitative manner the relative wood destroying
powers of the various species. A medium containing nitrogen in the
form of asparagine was used to determine the availability of organic
nitrogen. The latter series of experiments is rather interesting inasmuch
as the fungi under natural conditions very commonly become established
on material covered by surface growths of various benthonic organisms,
presenting upon decomposition a readily available supply of organic
nitrogen.
In determining the reaction of the various forms to particular artificial
media, the experiments were run in such a way as to provide uniform
conditions for the medium being tested. This procedure was used in
preference to conducting a group of experiments synchronously with a
particular species on the whole series of media. The former method
probably provides more reliable data on the relative availability of a
given substratum to different species, although uniformity of conditions
for a given species are sacrificed to a certain extent.
Two isolates of Ceriosporopsis halima were used in all the experiments
on reaction to substratum, to pH, and to temperature. One of these, Heli-
coma salinum, was isolated as the imperfect stage while the perfect stage,
Ceriosporopsis halima, was later obtained from ascospore isolations. These
two strains of the same organism show a decided similarity in their reac-
tions to different substrata, hydrogen-ion concentration and temperature
differences. Helicoma salinum was isolated from a wood block bearing
an abundance of its gray-black helical spores. In culture, the mycelium
produced similar spores in large numbers. Subsequently, on another
wood specimen, perithecia of Ceriosporopsis halima were found in large
numbers. Spore cultures from the latter yielded mycelia and helical
spores identical with those of the Helicoma. In view of this, the two iso-
lates were carried along in all cultural experiments. The results furnish
interesting evidence from a physiological point of view for their taxonomic
treatment as the perfect and imperfect stages of the same species.
442 Far.towia, VoL. 1, 1944
RATE OF RADIAL GROWTH ON ARTIFICIAL MEDIA
Following the general procedures employed by Brown (1923), Linder
(1929), Howell (1940) and other workers who have contributed data
on the rate of growth of fungi in culture, the rate of spread of the mycelium
was used in this study as an index of the availability of the substratum
for growth. As has been pointed out previously, this procedure is open
to some error, but, by and large, it is the most practicable means of de-
termining the vigor of a fungal colony in response to available food ma-
terials and particularly so in the case of synthetic media containing in-
soluble materials such as cellulose, wood or lignin. A summary of the
statistical data derived from the growth of the fungi on various nutrient
media is shown in Table 2.
TABLE 2, RATE OF RADIAL GROWTH IN MM PER DAY ON
NUTRIENT MEDIA
|
o Y Bae
g 8 8 4 Bite ee
es =s 8 8 8 8 8&8 8
2 @ 32 ©» 8 8 2B 3s
o 2. © fH & €& 4 E
Amphisphaeria maritima 82.40 1% .385 .82 §©6.82 =§=.08 31
Helicoma salinum 1.56 1.55 91 1.17 1.86 1.50 .83 1.44
Peritrichospora integra 1.17 = .05 52 1.04 .97 1.18 1.01 1.18
Ceriosporopsis halima 1.09 1.10 52 1.09 1.20 1.88 .47 .96
Halophiobolus opacus 2.61 12 140 54 2.50 1.82 2.80 .84
Halophiobolus salinus 4.47 1.90 2.07 2.76 4.07 3.48 2.97 3.34
Halophiobolus cylindricus 1.22 .87 (76 .76 1.10 .90 87 1.08
Phialophorophoma litoralis 62 1.09 .77 .55 .91 1.08 .95 .39
1. Cellulose
A carbohydrate invariably present in the natural substrata from which
these fungi have been isolated is cellulose. As might be expected, there-
fore, both growth and spore production are favorable on cellulose media.
The availability of cellulose in supporting the radial growth of seven
species in culture is shown in Fig. 1. It will be noted that Halo phiobolus
salinus, of the species represented, not only shows the greatest absolute
rate of growth but also the greatest relative radial growth on cellulose.
Similarly Halophiobolus cylindricus and Peritrichospora integra make
their most rapid development on cellulose of the various carbohydrates
supplied as substrata. For the remaining species cellulose provides one
of the more favorable carbohydrate media, and only in the case of Amphi-
sphaeria maritima and possibly Ceriosporopsis halima is growth slower
than on a medium containing maltose as the sole carbon source.
The ability of the fungi to produce cleared spots on cellulose agar is
not closely related to their rate of radial growth on cellulose as com:
pared to other substrata. Thus, certain cultures of Peritrichospora in-
tegra produced definite, cleared “halos” on cotton cellulose plates within
a period of 72 to 96 hours whereas cultures of Halophiobolus salinus on
BarcHoorn & LinpDER: MarINE FUNGI 443
the same medium produced poorly differentiated clear zones, despite the
fact that cellulose supported a more rapid growth than any other carbo-
hydrate supplied in the medium. Similarly, the cellulose digesting power
of Halophiobolus cylindricus, indicated by rate of growth on this sub-
stratum, was not demonstrated by the formation of a conspicuous clear
zone in the plates. On the contrary, cultures of Amphisphaerta mart-
tima on cellulose plates, although of slow growth, produced cleared areas
adjacent to the mycelial mat despite the fact that the rate of growth on
this substratum was less than that on maltose, xylose, pectin and even
wood flour. Hence, it may be reasoned from the evidence derived from
such culture experiments that the intensity of cellulase activity of the
various forms in culture is not clearly indicated by their relative rate
of growth on cellulose agar media. More critical measure of the cellulose
digesting powers of selected members of the marine fungi is at present
being studied through their enzyme production and ability to attack wood
under experimental conditions.
In this connection it is of interest to note that the ability of an organism
to attack cellulose is not always clearly demonstrable by its development
on synthetic media containing regenerated cellulose. This behavior is
probably due in part to the fact that synthetic cellulose media contain
modified forms of cellulose, such as oxycellulose and other degradation
products as well as to the fact that cellulose destroying organisms in
nature develop on substrata containing a wide variety of carbohydrate
compounds, which, singly or in combination, are favorable to a more
“normal” and vigorous development. It has been observed that in the
case of many Basidiomycetes, growth is unsatisfactory on plate cultures,
whereas extensive cellulose decomposition is actually effected in the
medium, as may be shown by analysis (Norman and Fuller, 1942).
The influence of cellulose and other substrata on the color of the col-
onies and spore production is shown in Table 3. It will be noted that
cellulose produces, in general, an effect similar to that of xylose and of
wood flour, particularly the latter. There is, however, no specific in-
fluence or particular effect exerted by the presence of cellulose on the
color of the colonies; nor is there anything to suggest that the availability
of cellulose, or any other carbohydrate supplied is indicated by the in-
tensity of pigment formation in the mycelial mat or in the adjacent
medium. For example, Peritrichospora integra, a form possessing marked
cellulose digesting power, produces no colored products on either cellulose
or wood flour agars, although the mycelium production and rate of growth
are greatest on these substrata.. Amphisphaeria maritima is interesting
also in this connection, since it produces a characteristic brown tawny color
on cellulose, xylose and wood flour and remains white on maltose agar,
a medium which supports the most rapid radial growth as well as very
abundant mycelial development.
Data regarding the effect of cellulose and other substrata on spore
444, FarLowla, VoL. 1, 1944
production are very unsatisfactory owing to the failure of most forms to
produce spores in Petri dish cultures within a reasonable period of time.
However, Ceriosporopsis halima, which produces conidiospores freely on
a wide variety of substrata, attained its greatest spore production on
cellulose and wood flour agars. Similarly Halophiobolus opacus, al-
though normally producing few spores, sporulated quite vigorously on
cellulose and wood flour with virtually no spore production on other
substrata. In Ceriosporopsis halima and Halophiobolus opacus it is evi-
dent that spore production is not influenced by deficiency of available
carbohydrate since both of these forms grow with considerable speed and
vigor on cellulose and wood flour (Table 2). Hence, although the evi-
dence is somewhat meagre there is no indication that a deficiency of avail-
able carbohydrate stimulates spore production. In this regard it is in-
teresting to note that on galactose and asparagine, Ceriosporopsis halima
made a feeble growth and produced no spores. |
2. Maltose
Although the disaccharide maltose furnished a very favorable sub-
stratum for certain species of the fungi, cultures of Halophiobolus opacus,
Peritrichospora integra and Halophiobolus salinus made a conspicuously
poor growth on this carbohydrate. These effects were so pronounced
in the case of P. integra and H. opacus that several series of cultures were
run in order to reduce the possibility of experimental error. The addi-
tional cultures, however, yielded consistent results. If reference is made
to the cellulose digesting power of these forms illustrated in F ig. 1, it will
be noted that the three species show definite negative correlation in re-
spect to the availability of cellulose as contrasted with maltose. Whether
or not this has any general significance is difficult to determine from the
small number of cases studied; certainly it is not true of the forms Halo-
phiobolus cylindricus and Ceriosporopsis halima for which both maltose
and cellulose are readily available. However, it is interesting to note
that in the helicosporous fungi imperfecti studied by Linder (1929), two
species, Helicomyces scandens and Helicondendron tubulosum, made their
most rapid growth on cellulose and their poorest growth on maltose, a
very similar response to that exhibited by the marine fungi Peritricho-
spora integra and Halophiobolus opacus.
Maltose provided the most favorable substratum for rapid radial growth
in cultures of Amphisphaeria maritima and Phialophorophoma litoralis,
for both of which forms cellulose is also an available nutrient. A. mari-
tima, however, despite the rapid radial growth on maltose did not pro-
duce a dense mycelium such as formed on wood flour, xylose or cellulose
agars. Moreover, the pronounced brown color of the colonies on the
three latter media is not developed on maltose. Phialophoro phoma
litoralis, on the other hand, not only made its most rapid radial growth on
maltose, but also developed an extremely thick felty mycelial mat.
TABLE 3. INFLUENCE OF SUBSTRATUM ON COLONY COLOR AND SPORE PRODUCTION. COLONIES 18-21 DAYS OLD
Figure Denotes APPROXIMATE PERCENT OF SPORE PRopUCTION
Amphisphaeria maritima
Peritrichospora integra
Ceriosporopsis halima
Halophiobolus opacus
Halophiobolus salinus
Halophiobolus cylindricus
Phialophorophoma litoralis
Cellulose
Brown 0
White 0
Gray 100
Light olive
green 10
Yellow
brown 0
Cottony
white 0
Mouse
gray 0
Maltose Galactose
White 0 White 0
White 0 Brown
gray 0
Very light White or
gray 10 hyaline 5
White 0 White 0
Faint yellow Green
brown 0 gray 0
Cottony Cottony
white 0 white 0
Green Green
gray 0 gray 0
Xylose
Dark
brown 0
Light
gray 0
Gray 30
White 0
Yellow
brown 0
Cottony
white 0
Rust red
brown 0
Starch Pectin Asparagine
White 0 Buff 0 White 0
Light Gray Cottony
gray 0 green 0 white 0
Tron gray 50 Light White or
gray 75 hyaline 5
Yellow Brown White or
brown 0 gray 0 hyaline 0
Mouse Dark brown White 0
gray 0 gray 0
Cottony Cottony White 0
white 0 white 0
Brown Gray 0 Light
gray 0 gray 0
Wood
Flour
Dark
brown 0
White 0
Gray 100
White 10
Yellow
brown 0
Cottony
white 0
Green
gray 0
IONOY ANIMVI, -YaGNIT ¥ NYOOHIUVG
SPV
446 FarLowI1A, VoL. 1, 1944
The growth of the two isolates Ceriosporopsis halima and Helicoma
salinum on maltose, as compared to cellulose, is interesting in view of
the nearly identical relative growth despite the difference in the absolute
rates of growth of the two strains. In Fig. 1, it will be noted that the
Helicoma isolate shows an equal growth rate on the two media. This is
also the case with the ascospore isolation of C. halima. However, the
actual rate of growth of the strain isolated from the conidial stage is ap-
proximately 30 per cent greater than that from the ascospore stage. This
similarity in relative growth rates for the two strains of C. halima on
different substrata is not confined to maltose and cellulose as can be
noted in the graphs in Fig. 1. The difference in absolute growth rate of
the two strains is likewise nearly of the same order of magnitude on the
various other media.
Insofar as the color of colonies and the production of spores is con-
cerned maltose shows no particular influence. In general, pigment forma-
tion was conspicuously less pronounced than on cellulose agars and in
the case of Amphisphaeria maritima, Peritrichospora integra, Halophio-
bolus opacus and Halophiobolus cylindricus the mycelia were totally de-
void of color, consisting of hyaline or cottony white mats (Table 3).
3. Galactose
Galactose as a source of carbohydrate for fungi has been variously in-
terpreted in several studies on the development and rate of growth of
fungi in culture. Horr (1936), by dry weight determinations, found that
galactose as the sole carbohydrate source provided a poor substratum for
two species of Penicillium and Aspergillus. Recently Edgecombe (1938)
demonstrated that galactose is readily available to certain forms, though
not to others, as evidenced by contrasting rates of radial growth on
agar media.
The availability of galactose to the group of fungi under discussion
here is significant not only in the light of previous studies but also in
view of their ability to hydrolyze and penetrate the pectic materials of
the isotropic intercellular substance of woody tissues. Although the
ability to break down pectic compounds in the intercellular substance
is not a sure indication that the galactose constituent of the pectic ma-
terial can be utilized it seemed desirable to test this possibility by com-
paring growth on galactose with growth on pectin media.
Insofar as such a correlation between pectin utilization and growth on
galactose is concerned, the results are indifferent, if not negative, as can
be noted in Fig. 1. Indeed, Ceriosporopsis halima which made its most
rapid growth on pectin agar developed very poorly on galactose. How-
ever, it is interesting to note that galactose provided a better source of
nutrient than maltose for the more active cellulose digesting forms, Peri-
trichospora integra, Halophiobolus opacus, and Halophiobolus salinus:
while in only two species, Amphisphaeria maritima and Cerios poropsis
halima was galactose inferior to all other carbohydrates as a food source.
BarcHoorn & LINDER: MARINE FUNGI 447
The nature of the mycelial development on galactose was quite ab-
normal in certain forms, which developed curiously twisted and “gnarled”
hyphae on galactose agar, although exhibiting a moderate radial growth.
- On galactose the average diameter of the hyphae in Peritrichospora in-
tegra, Halophiobolus opacus, and Halophiobolus salinus was less than on ~
any other carbohydrate medium.
Pigment formation on galactose agar plates after three weeks incubation
was less than on any other carbohydrate substratum, except in cultures
of Peritrichospora integra. The latter form developed a peculiar brown
gray color on galactose which was distinctly in contrast to the pigmenta-
tion produced on any other medium.
Spore production on galactose occurred only in cultures of Ceriospor-
opsis halima. The helical spores (Helicoma salinum) were produced very
sparsely and examination showed an almost complete absence of the usual
dark pigmentation of the spore wall. The spores, moreover, were curi-
ously abnormal in shape, lacking the usually complete curvature. In this
respect the conidiospores closely resembled those produced on media de-
ficient in nutrient, such as plain sea water agar (Table 1).
In general the experimental evidence indicates that galactose is a rela-
tively poor nutrient for the growth of these fungi but as determined by
rate of radial growth, it is available in some measure to all of them.
4. Xylose
The presence of xylan in the secondary wall of woody cells in conjunc-
tion with the rather ready microbiological decomposition of the pentosan
and hexosan hemicelluloses suggest that xylose might provide an available
substratum for wood inhabiting fungi. In this study d-xylose proved a
rather indifferent source of carbohydrate for the various marine forms. In
no case did xylose support more vigorous growth than certain other car-
bohydrates, while in cultures of Halophiobolus cylindricus and Phialo-
phorophoma litoralis development was actually slower than on any other
simple substratum. Amphisphaeria maritima is the only species of the
entire group which made a conspicuously rapid and vigorous growth on
xylose, the mycelium of this form not only spreading extensively but also.
branching profusely in a dense hyphal mat or cushion.
The influence of xylose on pigment formation is interesting in the case
of Phialophorophoma litoralis which developed an intense rust red-brown
color on this medium although showing no similar pigmentation on other
substrata including even such complete nutrient as potato agar. Amphi-
sphaeria maritima produced a dark brown pigmentation both in the my-
celium and in the medium on xylose substratum (Table 3).
Conidiospore production by Ceriosporopsis halima was fairly heavy on
xylose agar although less than a third of that on cellulose or wood flour
(Table 3).
448 Fartowia, VoL. 1, 1944
5. Starch
Starch is among the carbohydrates not infrequently present in the carbo-
hydrates comprising the natural substratum of these marine fungi and it
is therefore not surprising to find that it supports a vigorous vegetative
development.”
As shown in Fig. 1 starch was readily utilized by all of the species
studied and particularly by Halophiobolus opacus, Halophiobolus salinus
and Ceriosporopsis halima. In the case of the Helicoma salina isolate, the
most rapid radial growth on the various media used in this study occurred
on starch agar. As indicated by radial growth and hypha production,
starch proved to be an excellent carbohydrate source for the development
of all three species of Halophiobolus.
In none of the cultures was starch a definitely inferior food source for
these fungi, although Peritrichospora produced a rather sparse mycelium
on this medium, despite relatively rapid radial growth.
For determining the diastatic activity of the marine fungi on plate cul-
tures of starch media a simple iodine color test was used. Week-old cul-
tures were covered with dilute IKI solution until the characteristic blue
color reaction of starch developed in the agar. Differences in intensity of
staining in and around the portions of the medium penetrated by the
mycelium, indicated clearly the extent and the degree of degradation of
the starch. In certain cases starch digestion was sufficiently advanced
near the point of inoculation to result in a clear or colorless zone. All
species of the marine fungi investigated by this technique showed a well ©
developed diastatic activity.
Spore production by Ceriosporopsis halima was considerably higher on
starch than on maltose, galactose or xylose, although only about one half
that on cellulose. The spores showed their normal black pigmentation.
6. Pectin
Thin sections of infected wood in either the radial or tangential longi-
tudinal plane show that all of the fungi described in this study are capable
of penetrating the cell walls of woody tissues in passing from lumen to
lumen of adjacent cells. In this process the hyphae penetrate the pectic
compounds of the isotropic intercellular substance as well as the highly
lignified primary walls and thick secondary walls. In passing trans-
versely through the wall, as will be described in more detail later, the
hyphae narrow down to a tenuous thread and grow directly through the
various wall layers, widening out again immediately upon entering the
lumen of the adjoining cell.
A priori it would appear that pectin hydrolyzing enzymes would neces-
sarily be produced in order that the hyphae might penetrate the inter-
cellular pectic substances. Such action, moreover, presumably indicates
*The starch used in the growth experiments was a purified corn starch carefully
freed of proteinaceous substances and hemicelluloses.
BarcHOooRN & LINDER: MARINE FUNGI 449
that the hydrolytic products may be utilized by the fungus. The experi-
ments using pectin as a substratum demonstrate that this is the case.
In Fig. 1 it will be noted that all seven species including both strains of
Ceriosporopsis halima make a favorable growth on pectin agar. In no
case is pectin an unavailable or even inferior source of nutrient, while in
cultures of Peritrichospora integra and one strain of Ceriosporopsis halima
it supports a very rapid radial growth.
It is interesting to note that the response to both cellulose and pectin
exhibited by this group of marine fungi is quite similar to that found
in eight species of the helicosporous Fungi Imperfecti (Linder 1929).
Certain members of the two groups are of course very closely related but
it is a striking fact that the nutritional behavior should be so similar in
view of other considerable physiological divergences between the marine
and terrestrial members of the two groups.
A factor which should be noted in experiments using pectin agar is
the ready hydrolytic decomposition of pectin during the process of steam
sterilization. Even in alkaline sea water pectin agars registered a pH
of 6.5 after autoclaving for fifteen minutes. This lowered pH was difh-
cult to readjust inasmuch as the addition of dilute sodium hydroxide would
not produce a permanent change in pH until after the solution had cooled
to below 50° C.. Above this temperature the change in pH produced by
sodium hydroxide was transitory, the reaction rapidly tending toward acid
again. It is clear therefore that certain decomposition products of pectic
origin are present in the medium even after the above precautions have
been taken. These facts, however, do not invalidate conclusions drawn
as to the availability of pectic substance to the fungi, particularly when
the experimental data is correlated with the action of the fungi on a
natural substratum containing pectic compounds.
The effect of pectin on colony color and spore production is most pro-
nounced in cultures of Peritrichospora integra and Ceriosporopsis halima.
Peritrichospora integra produced a dark gray-green pigmentation both in
the hyphae (submerged hyphae chiefly) and in the medium. The in-
tensity of pigmentation in the pectin cultures of this form was greater
than in any other medium with the exception of the complex potato-
dextrose agar. Ceriosporopsis halima exhibited no unusual color reactions
on pectin although the amount of pigmentation was great. Correspond-
ingly, spore production was vigorous, being about 75 per cent of that on
cellulose agar.
7. Asparagine -
The availability of simple organic compounds of nitrogen in a syn-
thetic medium was determined by using substratum containing small
quantities of asparagine. This highly mobile nitrogenous compound is
present in nearly all living plant tissues though probably virtually absent
in the normal environment of these saprophytic fungi.
450 Fartowla, VoL. 1, 1944
The availability of asparagine is perhaps not satisfactorily determined
by this series of experiments owing to the great deficiency or complete
absence of carbohydrate in the substratum. However, a selective effect
is manifested by the very diverse responses made by various species as
shown in Fig. 1.
The extraordinary radial growth of Halophiobolus opacus on asparagine
agar must be interpreted in correlation with the characteristics of the my-
celium on this medium. Although radial growth was extensive the col-
onies were very sparsely branched, produced no aerial hyphae and ap-
peared quite hyaline in transmitted light. On both cellulose and starch
agar, the radial growth of this species was somewhat less than on aspara-
gine, but the colonies developed a more luxuriant and vigorous my-
celial mat.
Asparagine afforded the poorest substratum for growth in cultures of
Ceriosporopsis halima and Amphisphaeria maritima. Other species ap-
pear to utilize asparagine to a somewhat greater degree although in none
of the colonies was the hyphal mat vigorously developed. In fact, the
data for growth on asparagine as indicated by radial extent of the colonies
is somewhat misleading owing to the consistently sparse development of
the hyphae in this medium. Another consistent reaction of the various
forms to this substratum was the deficient development of aerial hyphae.
Only in the case of Amphisphaeria maritima and Halophiobolus cylin-
dricus, both of which invariably form bushy cushion-like colonies, was
there any appreciable formation of aerial hyphae.
Conidiospore formation by Ceriosporopsis halima was almost completely
suppressed on asparagine agar, although a few nearly colorless and aber-
rant spores developed.
Pigment formation in the colonies was likewise reduced to a minimum
on asparagine. With the sole exception of Phialophorophoma litoralis
the colonies were white or hyaline in both reflected and transmitted light.
It can be stated with little qualification that asparagine is decidedly
deficient as a nutrient for the development of these fungi, and that rate
of radial growth is a poor index of total growth on this substratum unless
it is correlated with the abundance of mycelium formation.
8. Wood Flour
Despite minor changes in resistance to fungal decay and in the chemical
condition of wood induced by autoclaving (Schmitz 1919) it is reasonable
to suppose that the growth of a fungus in an agar medium containing
finely divided wood might indicate to some extent the wood decaying
powers of the organism. A priori, it would seem that even active wood
destroyers might grow more slowly on wood than on synthetic media con-
taining only that constituent of the wood most readily available to them.
Thus cellulose destroying microérganisms might be expected to make a
more vigorous development on “purified” cellulose than on plant fibers
BarcHoorn & LinpER: Marine FUNGI 451
containing, in addition to cellulose, relatively large amounts of resins,
waxes, lignin or other encrusting and protecting substances normally pres-
ent in untreated woody tissues. Similarly, organisms which readily break
down the hemicelluloses of the secondary wall of woody cells might pre-
sumably make a more rapid growth on synthetic substrata containing
xylose and arabinose. The growth of wood destroying fungi in laboratory
cultures on various wood components is probably, however, never as
rapid nor vigorous as development on the normal substrata in nature.
Not only are the complex factors in the ecology of the environment vastly
different but also the accumulation of inhibitory metabolic products is
almost inevitable in laboratory experiments. Despite these experimental
difficulties, however, it is clear that growth on wood in culture gives some
reasonable index of the potentialities for growth in wood under natural
conditions.
The wood flour used in these experiments consisted of finely divided
particles obtained by filing clear-grained sections of seasoned white pine.
The resulting flour or dust was extracted for resins, washed repeatedly in
solvents and finally prepared as a 0.75 per cent suspension in agar.
Growth on this medium was surprisingly rapid and vigorous in the
case of all but two species, viz., Halophiobolus opacus and Phialophoro-
phoma litoralis, colonies of which not only increased in diameter slowly
but also developed scanty mycelia. .
The most striking development on the wood flour substratum was made
by Peritrichospora integra, cultures of which grew with great vigor, as
indicated both by rate of radial increase (Fig. 1) and the production of
a luxuriant hyphal mat. Moreover, colonies of this form after a week
or ten days development produced a faint but definitely cleared zone in
the medium, indicating the partial or complete destruction and dissolu-
tion of the smaller wood particles dispersed in the agar. This latter
effect is indirect evidence of the utilization of lignin by the fungus, an
ability which many of these fungi unquestionably possess as will be
shown later.
Three of the seven species, viz., Halophiobolus salinus, Peritrichospora
integra and Halophiobolus cylindricus exhibited a greater rate of radial
growth on wood flour than on maltose agar, an interesting response in
view of the relative solubility of the carbohydrates in the two media.
The color of the colonies on wood flour was quite similar to that on
cellulose agar except in the case of Halophiobolus opacus which failed
to form any discernible pigment (Table 3). Amphisphaeria maritima
produced a very intense dark brown pigment both in the mycelium and
in the substratum, a reaction similar to that produced on xylose.
Conidiospore production in cultures of Ceriosporopsis halima was more
vigorous on wood flour than on any other artificial substratum with the
possible exception of potato-dextrose agar. In addition, the spores were
nearly jet black in color, an appearance intensified by their occurrence
I aund1y
CELLULOSE
MALTOSE
GALACTOSE
XYLOSE
STARCH
PECTIN
ASPARAGIN
WOOD FLOUR
CELLULOSE
MALTOSE
GALACTOSE
XYLOSE
STARCH
PECTIN
ASPARAGIN
WOOD FLOUR
CELLULOSE
MALTOSE
GALACTOSE
XYLOSE
STARCH
PECTIN
ASPARAGIN
WOOD FLOUR
2
€
RADIAL GROWTH IN MILLIMETERS PER DAY _ ~ - a
= 5 PS a °| °
Pk ‘i re a 1 "id = i ‘ N ese |
atic E |
ees ‘3 Saas ,
| aaa a a ae a ee
ee Saas eee Se See Rea hee eT
ef ities Mieke oe a I ee SS ke
ae Paes eee sat ie ER
Pi eee esos 8 50.
Fig. 2. Transverse section of the outer edge of a hard pine test block showing ad-
vanced stages in fungal hydrolysis of the cell wall caused by Ceriosporopsis
halima. The enzymatic activity of the hyphae is almost exclusively confined
to the central layers of the secondary wall. The more highly lignified inner
and outer layers of the secondary wall are relatively unaffected at this stage
of decomposition. Specimen number 1l. X 420.
Fig. 3. Transverse section of a spruce test board attacked by Halophiobolus longi-
rostris. Note the long neck of the deeply embedded perithecium. Specimen
number 22. > 50.
Fig. 4. Radial longitudinal section of the same wood specimen shown in Figure 3.
Note the complete destruction of cell wall substance in the region surrounding
the perithecium. Within the tracheids of the wood are spherical inflated cells
produced by hyphae in the lumina. Specimen number 22. 50.
Fig. 5. Transverse section of hard pine test block attacked by Ceriosporopsis halima
showing early stages in the breakdown of the cell wall by hyphal action. The
area shown is two growth rings in from the outer exposed surface of the wood.
Note the restriction of hyphal cavities to the central layers of the secondary
wall. Specimen number ll. X 230.
467
MARINE FUNGI
BarcHoorn & LINDER
Pirate VII
1(3): 469-481 FARLOWIA January, 1944
STUDIES IN THE GENUS ZYGOSACCHAROMYCES I.
TRANSFER OF PELLICLE-FORMING YEASTS
TO ZYGOPICHIA 1
WALTER J. NICKERSON 2
That the taxonomy of even the sporogenous yeasts is in an unsatis-
factory state is a statement admitting of little debate. The important
work of Stelling-Dekker (1931) served to bring considerable order out
of chaos, but the need is still felt for monographic treatments of the vari-
ous genera of yeasts. In considering only the organisms of which cul-
tures were available to her, Stelling-Dekker followed a practice which
has much in its favor. Practically, however, it results in almost com-
plete neglect of species not included in the monograph by all except
specialists in a particular group. One function of a monographer is
the critical consideration of as many of the forms assigned to a group
as is possible. Since it is manifestly impossible for one person to have
access to cultures of all the species in a large genus, the monographer is
forced, by necessity, to make decisions based solely upon descriptions in
the literature. However, the scanty descriptions in some of the older
literature are frequently valueless, and the names should be cleared from
the books as nomina nuda, nomina dubia, etc., as the case may be, unless
an authentic culture is at hand for a redescription.
The yeasts are a poorly defined assemblage of fungi, chiefly Ascomy-
cetes, with a monophyletic standing in the Endomycetaceae favored by
some workers, and a biphyletic treatment embracing the Exoascaceae
sponsored by others. It seems best to think of a “yeast-state” much as
one today thinks of a “colloidal-state” rather than of “a colloid.” This
“no-man’s land,” which does not even grant its workers a distinctive
name (!), is one cultivated chiefly by techniques borrowed from bac-
teriology and mycology as the need may arise. In the past, adherence
to the International Rules of Botanical Nomenclature has only too fre-
quently been neglected. And a somewhat exclusive adoption of bac-
teriological techniques has resulted at times in the neglect of valuable
taxonomic tools such as the range of variation, morphology of spores
and of asci, geographical distribution, etc.
In presenting this paper as the first in a series intended to constitute
a monograph of the genus Zygosaccharomyces, the author is aware of
some, at least, of the difficulties and limitations imposed by the very na-
ture of the organisms involved. Use must be made of any distinguishing
* Aided in part by a grant from the Faculty Research Fund of Wheaton College.
? The writer, now serving with the Army of the United States, wishes to thank
Drs. D. H. Linder and W. L. White for their help on problems of nomenclature, Drs.
Mrak and Wickerham for kindly supplying some of the cultures used, and Dr. H.
W. Anderson, University of Illinois, for sending type slides of Z. Chevalieri var.
Andersonii with which comparisons were made.
469
470 FarRLowl1A, VoL. 1, 1944
character available, be it morphological or physiological, for the organ-
isms in question do not fall into clearly defined, discrete species, but
present a picture of individuals more or less concentrated into groups
(species) with gradations in all directions away from any given center
of concentration. A character which shows a normal statistical distri-
bution is in general of little separatory value taxonomically. For instance,
if spore sizes within a genus fall along a normal distribution curve when
graphed, then this particular character will not serve alone to differentiate
groups. On the other hand, a character exhibiting discontinuous variation
within a genus serves readily to divide a genus into two groups: i. e., those
that have and those that have not. The “centers of concentration” men-
tioned above grow out of the overlapping of the curves of discontinuous var-
iation. Possibly no microbiologist could be found who would seek to con-
ceal the fact that the species concept in regard to yeasts is a fairly intangi-
ble thing; but for practical purposes, possibly to parallel or interweave
with the development of a comparative biochemistry, it is essential that we
be agreed upon principles of intra-generic classification. The inter-specific
gradation which exists must be recognized as such and not serve to cause
the creation of numerous “poor” species. The existence of strains of
microorganisms is clearly recognized by all as a difficult question taxo-
nomically; a strain is essentially any isolate of a micro-organism that is
in any way distinguished from its “parent” species; usually it is distin-
guished in some comparative way, as larger, smaller, more vigorously
fermenting, etc. However, in the case of some micro-organisms, where
parasitism is involved, one is able to distinguish clearly among forms
identical with respect to the usual morphological and physiological cri-
teria. See Leonian and Lilly (1943) for an interesting discussion of
the species-complex problem in microbiology. Perhaps the existence of
strains serves to indicate the direction in which the production of dis-
continuous variation within a genus is progressing with time, and per-
haps it is a result of an improvement in our ability to make distinctions.
The subgenus Zygosaccharomyces Barker, following Klocker (1924)
and Stelling-Dekker, is rather large, with the literature on it scattered.
Approximately 60 species have been described since Barker (1901)
founded the genus. To date, Stelling-Dekker’s treatment has been most
complete with a consideration of some 20 species. During the past 5
years the writer has gathered together some 90 cultures including about
40 of the described species. From these cultures, a few stand out by
reason of their not meeting the generic requirements of Zygosaccharo-
myces. While the writer, in this paper, does not intend completely to
delimit the genus, he is in thorough agreement with Klécker and with
Stelling-Dekker in excluding all organisms that form films promptly in
ordinary sugar-containing liquid media. Before taking up the problem
of the relationships existing among species properly belonging to Zygo-
saccharomyces, the writer hopes to simplify the problem by transferring
NICKERSON: ZYGOSACCHAROMYCES 471
forms which do not meet the requirements of the genus. The remainder
of this paper will be devoted to a discussion of 5 pellicle-forming yeasts
that have been assigned at one time or another to Zygosaccharomyces,
but which should be classified in Zygopichia.
METHODS
Sources of the organisms studied in this paper are listed under the
species concerned in the section on taxonomy. Stock cultures were car-
ried as stab-cultures in wort agar (Difco) with 1% added Bacto-agar.
Culture studies employed wort and wort agar slants; for liquid cultures
a synthetic medium described later in the paper was also used with the
addition of 1.5 gm. Bacto-asparagine per liter. Measurements on vegeta-
tive cells were made, using an ocular micrometer, after 48 hours growth;
averages of the measurements are reported. Sporulation occurred read-
ily by seven days on wort agar slants except in the case of Z, Chevaliert,
H-175, which responded to culture on Sabouraud’s dextrose agar (Difco)
with a 5% addition of Aspergillus filtrate prepared according to Nick-
erson and Thimann (1943). Fermentation tests were performed with
Durham tubes; if dextrose, fructose, and mannose were not fermented,
no other sugar was investigated. Nitrate utilization was studied in a
liquid medium using KNOs as the sole nitrogen source; a test for nitrite
was also performed; a discussion of the value of examining solely for
the presence of nitrite will be found later in the paper. Ethanol utiliza-
tion was studied using the synthetic medium referred to but omitting
glucose, peptone, and asparagine; recently boiled absolute alcohol was
added aseptically after autoclaving to give the medium a final concen-
tration of 2%. A discussion of adaptation and tolerance to increasing
solute concentrations, including ethanol, will be found later in this paper.
Gelatin liquifying ability tested with stabs and plates of wort gelatin
(pH 4.8) and malt extract (Difco) gelatin (pH 5.8). Maximum tempera-
ture for growth studied in wort and in the synthetic medium.
EXPERIMENTAL
In view of the findings of Baars (1930) on the sulfate reducing bac-
teria, wherein he showed that Vibrio estuarii is merely a salt-adapted
form of Spirillum desulfuricans, it was thought worthwhile to investigate
some of the yeasts discussed in this paper. As will be pointed out,
Zygosaccharomyces Guilliermondi is quite similar to Zygopichia Cheva-
lieri, being distinguished chiefly by its reported ability to grow in media
containing 25% NaCl. Since the possibility exists, at least, that Z.
Guilliermondi is merely a salt-adapted form of Z. Chevalieri, the following
experiments on adaptation of various members of Zygopichia were
undertaken.
To a basic medium consisting of: 3.0 g. (NH) 2SOx, 2.0 g. KH2POx,
0.25 g. CaCls, 0.25 g. MgSOu, 20.0 g. glucose, 2.0 g. Bacto-peptone, 0.10 g.
472 FarLowI1A, VoL. 1, 1944
yeast extract, and distilled water to make 1 liter, various amendments
were added as indicated in the respective experiments. The usual meth-
ods for quantitative measurement of yeast growth—counting of cells or
weighing of cells—are difficult to use with these film-forming species;
in most cases only a qualitative statement is given as to the extent of
growth; in a few cases, quantitative measures of metabolic products—
chiefly total acid production—have been made. All experiments were
kept at a room temperature of about 20° C.
In tables 1-3 are found data obtained from duplicate experiments on
subjecting yeasts to various environments. All cultures were inoculated
from vigorous growths on wort agar and observed at intervals for pellicle
formation; usually the lower concentrations supported a good growth.
After varying intervals of time, higher concentrations not showing growth
were inoculated from the next lower concentration showing a pellicle;
a sterile flask of the higher concentration in question was also inoculated
—data reported are based on the inoculation into this previously ster-
ile flask.
The concentrations employed for the various amendments are calcu-
lated in three ways in table 1: as per cent final concentration, as molarity,
TABLE 1—SALT TOLERANCE
Total NaCl
concentra- Z. Chevalieri M.1 2. Chevalieri v. Andersonii H. 177
tion
2%
0.34 M pellicle in 5 days, sediment only, 10 days.
0.61 M
5%
0.85 M pellicle in 10 days. sediment only, 25 days,
1.53 M
8% no growth in 15 days; pellicle in | no growth in 25 days; no growth in
1.36 M 1 day when inoculated from 5%. 5 days when inoculated from 5%.
2.45 M
10% * | no growth in 15 days; pellicle in
1.7M 3 days when inoculated from 8%.
3.06 M
15% no growth in 20 days; faint sediment
2.55 M in 5 days when inoculated from
4.59 M 10%; no pellicle.
20% no growth in 25 days; no growth in
3.4 M 10 days when inoculated from sedi-
6.12 M ment of 15%.
and as molarity < 1.8 to represent the effective molarity for the develop-
ment of osmotic pressure by a binary electrolyte. It is indicated in table
1 that the strain of Z. Chevalieri used, while able during growth to be-
NICKERSON: ZYGOSACCHAROMYCES 473
TABLE 2—SUGAR TOLERANCE AS SHOWN BY PELLICLE FORMATION
Total concentration of glucose
Organism
2% 6% 10% 14% 18% 22% 30% 42%
0.11M| 0.33M]| 0.55M| 0.77M| 0.99M/ 1.22M| 1.65M/ 2.33M
Z. Chevalieri v. 2 2 2 2 2 3 3 7
Andersonti days days days days days days days days
come adapted to increasing concentrations of solute, has an upper limit
to this adaptation in the region of 4M solute concentration. Thus, while
Zygosaccharomyces Guilliermondi is probably just a salt-adapted form
of Z. Chevalieri, there is a difference in their salt tolerances of about
3M effective solute concentration. Under the conditions of the experi-
ment and with the strain employed, Z. Chevalieri var. Andersonii ex-
hibited no ability to form a pellicle on even 2% NaCl.. By 10 days it
had formed good sediment, but no trace of pellicle.
In the alcohol series (table 3), each of the three organisms formed a
TABLE 3—ALCOHOL TOLERANCE
Total
alcohol Z. Chevalieri var. Z, Chevaliert M.1 4. farinosa ATCC 2252
concen- Andersonii H.177
tration
2%
0.43 M pellicle in 6 days. pellicle in 5 days. pellicle in 6 days.
5% no growth, 14 days; pel- | pellicle in 8 days. pellicle in 6 days.
1.09 M licle on inoc. from 2 % in
1 day.
10% no growth, 16 days; pel- | no growth, 10 days; pel-
2.17 M licle in 5 days on inoc. | licle in 2 days on inoc. | pellicle in 6 days.
from 5%. from 5%.
15% no growth, 18 days; pel- | no growth, 12 days; pel- | no growth, 10 days; pel-
3.26 M licle islands in 5 dayson | licle in 6 days on inoc. | licle in 2 days on inoc.
inoc. from 10%. from 10%. from 10%.
20% no growth, 18 days; no | no growth, 16 days; | no growth, 15 days;
4.34 M growth 5 days on inoc. | faint sediment only on | pellicle islands in 5 days
from 15%. inoc. from 15%. on inoc. from 15%.
well-developed pellicle that began to ascend the walls of the flask or
tube shortly after being formed. Castelli used this pellicle phenomenon
as a characteristic serving to differentiate Z. chiantigiana from Z. Cheva-
lieri but it is evident that such an ability is common to most, if not all,
members of the genus.
TABLE 4—A COMPARISON OF MEMBERS OF Zygopichia
Z. Chevalieri Z. Chevalieri Z. Chevalieri Zygosacch. Z. farinosa Z. chiantig-
v. Andersonii v. Barkeri Guilliermondii iana
Pellicle in liquid + + + + + +
media
Fermentation - - - - +d,1,m -
Conjugation heterogamic heterogamic physiol. heterogamic iso- and heterogamic
heterogamic heterogamic
Spore shape hat-shaped or spherical and spherical and hat-shaped or spherical or oval spherical or
spherical smooth smooth spherical flattened
Spore size + 2.5 up +2.0u 44.3 p +2.0 4 2x3uyn + 284
Cell size in wort 34x48.5 uw 4x6.5y 4-5 x 5-7.8 uw 3-5 x 6-9 uw 2-5 x 4-16 2.4-3.6 x 7.2-9.6 pu
2-3 days old
Mycelium on agar + _ — +. = +
Gelatin liquefac- +21 +21 —20 —60 —60
tion in days
Nitrite test _ - * + - -
Ethanol utilization; + + * > + +
pellicle formation
Max. temp. for bud- 42-43 39-40 32.5 34 * 37
ding in wort in °C.
Remarks: cells often cells elongate in grows in presence cells soon become
1,5-2.2 x 12-22 p
old culture
of 25% NaCl
elongate
central oil globule
in spores
* No data
PLY
PROL ‘T “IOA ‘VIMOTUVA
NICKERSON: ZYGOSACCHAROMYCES 475
ZYGOSACCHAROMYCES vs. ZYGOPICHIA
Following Stelling-Dekker and Klécker, the distinctions between Zygo-
saccharomyces and Zygopichia, genera having gametic conjugation pre-
ceding the formation of spores, are as follows: |
Zygopichia: in wort a dry, dull pellicle within a short time (24-72
hours) after inoculation; spores angular, hemispherical, hat-shaped,
or spherical.
Zygosaccharomyces: no pellicle in wort, or at most a soft, slimy pellicle
when fermentation is nearly complete, which is a considerable time
(2-3 weeks or more) after inoculation; renniform, hat-shaped, or
spherical spores.
ZYGOPICHIA Klécker, Die Garungsorganismen, 3 Auflage, Berlin-Wien. 1924.
The genus was established by transferring Zygosaccharomyces Cheva-
liert Guilliermond to it as the type. It bears the same relation to Pichia
that Zygosaccharomyces bears to Saccharomyces. In addition to the
generic characters brought out in the above comparison, there are the
following: fermentation, none or only very weak, nitrite tests negative,
nitrate assimilation poor, cells of varying shape with a tendency towards
pseudomycelial formation.
KEY TO THE SPECIES OF ZYGOPICHIA *
1. On wort, a thin, smooth pellicle; no. fermentation
2. Spores hat-shaped; pseudomycelium formed on wort agar
1. Z. Chevalieri
2. Spores smooth only; no pseudomycelium formed
3. Spores small, about 2.0 4 diameter; heterogamic conjugation.
2. Z. Chevalieri vy. Andersonii
3. Spores large, greater than 4.0 ». diameter; gametes morphologically similar
3. Z. Chevalieri v. Barkeri
1. On wort, a thick, dry, wrinkled pellicle; fermentation of glucose, fructose, and
mannose only 2. FObaee tis 4. Z. farinosa
1. Zygopichia Chevalieri (Guilliermond) Klécker, Die Giérungsorganismen, p. 289-
290, Berlin-Wien. 1924.
Zygosaccharomyces Chevalieri Guilliermond, Ann. Sci. Natur. II Botan. 19:
1, 1914; C. R. Soc. Biol. Paris 1: 442. 1911.
Pellicle formed promptly in wort, no fermentation of any sugar, wort
gelatin or malt extract gelatin liquefied within 21 days. Cells in wort
3-5x4-8.5 p, pseudomycelium frequently formed. Asci arise after
heterogamic conjugation or parthenogenetically; spores round, hemi-
spherical or angular, approximately 2.5 w in diameter. Nitrite negative;
good growth and pellicle formation with ethanol as the sole carbon
source. Maximum temperature for budding, in wort, 42-43° C.
Isolation: by Guilliermond from Biliwine (made from tubers of Osbeckia grandiflora)
in West Africa.
Cultures examined: Guilliermond Lab. de Parasit. no. 617, through Harvard Col-
lection no. 175; E. M. Mrak no. 1, orginally from Tanner’s Collection.
® See also table 4 for a comparison of the species.
476 Fartowia, Vou. 1, 1944
Klécker’s disposition of this species was approved by Stelling-Dekker,
and nothing found in this investigation warrants further discussion here.
A study of adaptation to various chemical environments in the growth of
this species will be reported later in this paper.
2. Zygopichia Chevalieri var. Andersonii Nickerson nom. nov.
Zygosaccharomyces bisporus Anderson, Journ. Infect. Dis. 21: 341, 1917, nec
Zygosaccharomyces bisporus Naganishi, Bot. Mag. Tokyo 31: 107-116, 1917.
Pellicle formed promptly in wort, no fermentation of any sugar, gelatin
liquefied within 21 days. Cells on wort agar average 4x6.5 p; no
pseudomycelium formed on agar. Asci arise after heterogamic conju-
gation or parthenogenetically; spores round and smooth, approximately
2.0 » diameter; very little variation from this average. Nitrite negative;
good growth and pellicle formation with ethanol as the sole carbon source.
Maximum temperature for budding, in synthetic medium and in wort,
39-40° C.
Isolation: by H. W. Anderson from human feces.
Culture examined: Guilliermond Lab. de Parasit. no. 569, through Harvard Co)-
lection no, 177.
An interesting coincidence occurred in 1917 when two men in widely
separated parts of the world independently gave the same name to yeasts
isolated by them. H. W. Anderson, in Urbana, Illinois, described one
Z. bisporus, and H. Naganishi, in Tokyo, described the other. The first
monographic treatment (Stelling-Dekker) after publication of these names
accepted Naganishi’s species and did not mention Anderson’s. It is quite
clear that Anderson and Naganishi did not describe the same yeast, for
the latter’s species complies with Stelling-Dekker’s emendation of the’
genus Zygosaccharomyces, while the former’s, as here shown, does not.
Since it is not possible to determine which specific name has priority, the
writer has been guided by Stelling-Dekker’s acceptance of Naganishi’s
name.*
The culture of Z. Chevalieri var. Andersonii examined is believed to
be a transfer of Anderson’s original isolate. It is identical in all respects
with Anderson’s description. While this organism is closely similar to
Z. Chevalieri, it is sufficiently different to warrant the varietal standing
proposed. Similarities between the two forms are in regard to pellicle,
absence of fermentation, gelatin liquefaction, mode of origin of asci, and
approximate cell and spore sizes. Points of difference are that the pro-
posed variety has smooth spores only, no pseudomycelial formation, and
a maximum temperature for budding of 39-40° C. (this latter point a
very small one, however).
3. Zygopichia Chevalieri var. Barkeri Nickerson var. nov. Yeast G, Pearce and
Barker, J. Agric. Sci. Cambridge 3: 55, 1908.
(From description given by Pearce and Barker.) In mediis maltatis membrana
cito formatur; nullum saccharum fermentatur. Cellulae in mediis maltatis 4-5 x
“International Rules, Art. 57.
NICKERSON: ZYGOSACCHAROMYCES 477
5-7.8 wu; in culturis altis multae cellulae non nihil longiores. Asci post conjuga-
tionem per cellulas formae similis pariunt; parthenogenesis quoque evenit. Sporum
globosum leveque longitudo media lineae mediae 4.3 2; spora post conjugationem
formata solum in una duae cellulae reperiuntur. Calor maximus gemmandi 32.5° C.
(Description as given by Pearce and Barker.) Pellicle formed prompt-
ly in wort; no fermentation of any sugar. Cells in wort 4-5 x 5-7.8 yp;
elongated cells in old cultures. Asci arise after conjugation of morpho-
logically similar cells, or parthenogenetically. Spores round and smooth,
average 4.3 » in diameter; when formed after conjugation, are found in
only one of the two gametes. Maximum temperature for budding 32.5° C.
Isolation: by Pearce and Barker from bottled English cider.
Culture examined: known only from the literature.
This organism was considered by Klécker as belonging in Zygopichia,
but he did nothing about its name. It is closely similar to Z. Chevalieri
in pellicle formation, absence of fermentation, and approximate cell‘ size
in young cultures, with elongate cells in old cultures. Sufficient differ-
ences for the proposed varietal standing are found in possession of
“physiological heterogamy” only, formation of only smooth spores, spore
size, and the low maximum temperature for budding (32.5° C.).
4. Zygopichia farinosa (Lindner) Klocker, loc. cit.
Saccharomyces farinosus Lindner, Woch. f. Brau. 11: 152, 1894.
Pichia farinosa (Lindner) Hansen, Centralbl. f. Bakt. 12: 529-538, 1904.
Zygosaccharomyces farinosus (Lindner) Papadakis, C. R. Soc. Biol. Paris.
86: 447, 1922.
Prompt formation in wort of a thick, dry, wrinkled pellicle; ferments
glucose, fructose, and mannose weakly. Gelatin not liquefied; nitrite
test negative. Growth and pellicle formation with ethanol as sole car-
bon source. Cells in wort 2-5 x 4-16 p, single or in budding chains; no
pseudomycelium. Spores round to oval, and smooth, 2x3 y; formed
after isogamic or heterogamic conjugation or parthenogenetically. Spores
have a refractile granule in center.
Isolated: by Lindner in Danzig from beer; by Saito from soy bean sauce.
Culture examined: ATCC no. 2252; originally from Tanner Collection.
This organism has had a rather checkered career. Discovered by
Lindner, who considered it a Saccharomyces, it was shown by Hansen in
1904 to belong to Pichia on the basis of pellicle formation. This yeast
was considered by Guilliermond as late as 1920 to have spores produced
without conjugation, yet his student, Papadakis, in 1922 showed that a
gametic conjugation was clearly involved, preceding spore formation.
Klécker, in 1924, on establishing the genus, removed this species from
Zygosaccharomyces along with Z: Chevalieri. Since Stelling-Dekker has
reaffirmed this species, and nothing has turned up in the course of this
investigation to raise any question, nothing more need be said.
478 Fartowl1A, VoL. 1, 1944
DOUBTFUL SPECIES
1. Zygosaccharomyces Guilliermondi Dufrénoy, Bull. Soc. Myc. Fr. 43: 272, 1927.
(Description as translated from Dufrénoy.) Prompt pellicle forma-
tion in wort; no fermentation; gelatin not liquefied within 20 days. Asci
originate after heterogamic conjugation or parthenogenetically; spores
hat-shaped or smooth, averaging 2.0 » diameter. Young cells in wort
3-5 x 6-9 u; pseudomycelium formed on agar. Will grow in media con-
taining 25% NaCl. Maximum temperature for budding is 34° C.
Isolation: by Dufrénoy in Corsica from the brine of the normal citron fermentation.
Culture examined: known only from the literature.
Dufrénoy used Lindner’s relatively inaccurate “little” method for study-
ing fermentation of this yeast, and observed a slight fermentation of
fructose, but none of glucose. Since it is highly unusual, if indeed pos-
sible, for an organism to ferment any sugar and yet not ferment glucose,
it is here considered that this yeast actually has no fermentative ability.
This form is closely similar to Z. Chevalieri in regard to pellicle forma-
tion, origin of asci, size and shape of spores, pseudomycelial formation,
and lack of fermentation. It is doubtful that sufficient differences exist
between the two to warrant varietal standing. It is a question whether
this species should not be regarded merely as a salt adapted form of
Z. Chevalieri, in view of the results of experiments on adaptation re-
ported early in this paper. The time elapsed (20 days) in the judging
of gelatin liquefaction as negative is somewhat shorter than one usually
considers sufficient when working with yeasts. Preliminary findings
which will be published at a later date indicate that high osmotic pressures
have an inhibitory effect on proteolytic exoenzymes involved in gelatin
liquefaction by two of the yeasts discussed here. This finding may pos-
sibly be of interest in view of the results obtained by Johnson et al (1942)
on the effect of barometric pressure on the luciferase mechanism of
phosphorescent bacteria. In any event gelatin liquefaction is an un-
satisfactory character on which to differentiate a species because of our
lack of knowledge concerning its distribution among yeasts and the in-
fluence environment may play upon it. In spite of this close similarity
between Zygosaccharomyces Guilliermondi and Zygopichia Chevalieri,
the author does not choose to reduce the former to synonymy with the
latter at the moment. Dr. Dufrénoy, in a personal communication, in-
forms me that his culture is possibly still in existence in the collection
of Prof. Guilliermond at the Sorbonne. Until such time as the culture
can be examined or is known definitely to be lost, decision may be reserved.
2. Zygopichia chiantigiana Castelli, Archiv f. Mikrobiol. 9: 449-468, 1938.
(Description as translated from Castelli.) In malt after 24 hours at
25° C., cells globose or elongate, often slightly curved in the form of
a sausage; cells usually contain a refractile globule; single or in groups
of 5-6 cells. Average dimensions 3.8-4.5 x 7-9.5 ». After 5 days cells
elongate, frequently forming pseudomycelia. On malt agar, young cells
NICKERSON: ZYGOSACCHAROMYCES 479
3.8-5x 7-9 3; many cells possess extruded tubes after 3 days. By 7
days heterogamic conjugation abundant with some asci formed par-
thenogenetically; usually 2 spores per ascus, spores globose or liek
flattened, usually containing a central refractile globule; average size
2.6-3 p. On liquid media, as wine must or malt, a pellicle is rapidly
formed, smooth and gray-white at first, it soon becomes strongly wrinkled
and hazelnut in color. There is no fermentation of any sugar. Growth
with ethanol is positive; scanty assimilation of nitrates; gelatin is not
liquefied in 60 days; maximum temperature for budding Si: GC.
Isolated: by Castelli from wine musts from the classic Chianti region and hills
near Florence.
Culture examined: known only from the literature.
Castelli in his description of this species pointed out the close sim-
ilarity to Z. Chevalieri, but was of the opinion that his organism differed
sufficiently to warrant a new specific name. However, the characters used
by Castelli to separate this yeast do not seem sufficiently strong to the
writer to warrant such a separation. Cell size is a quite variable quan-
tity with yeasts, and it is well known that the composition of the medium
plays an important réle in this variation. He is in error, in the writer’s
‘opinion, in saying that Z. Chevalieri forms only a weak pellicle (velo
sottile) on liquid media. The writer has repeatedly obtained films with
authentic cultures of Z. Chevalieri that could be described by the words
used by Castelli for the pellicle made by his yeast. Gelatin liquefaction
is a rather uncertain character for differentiating yeasts, as previously
pointed out. Rather than reduce this organism to synonymy with Zygo-
pichia Chevalieri, the author prefers to follow the course chosen pre-
viously, and await the time when it may be possible actually to examine
this yeast.
3. Zygopichia Chevalieri var. fermentati Saito. |
Such a species is listed in the 1940 catalogue of cultures issued by the
Centraalbureau voor Schimmelcultures at Baarn. The author has been
unable to find any reference in the literature to the name. It is listed
here solely for the sake of completeness.
EXCLUDED SPECIES
The following species were placed by Klécker in Zygopichia. Stelling-
Dekker reclaimed them for Zygosaccharomyces, for, though they may
produce a pellicle on liquid media, it is only after considerable time
and under special conditions (as with high salt concentrations).
Zygosaccharomyces japonicus (Saito) Guilliermond, Rev. Gen.. Bot. 20: 32, 1908.
Zygosaccharomyces japonicus var. soya (Saito) Stelling-Dekker, J. c. 1931.
Zygosaccharomyces major var. salsus (Takahashi and Yukawa) Stelling-Dekker,
I. c. 1931,
A study of cultures of these organisms, obtained from the American
Type Culture Collection, or from the Northern Regional Research Lab-
480 FarLowiA, VoL. 1, 1944
oratory of the Dept. of Agriculture, showed that a pellicle is not formed
promptly in ordinary sugar-containing liquid media. Stelling-Dekker’s
exclusion of these species from Zygopichia seems entirely justified.
DISCUSSION
At present the only character on which the genera Pichia and Hansenula
are separated is that of nitrate utilization. As Mrak et al (1942) have
pointed out, this is hardly a satisfactory condition, and they have re-
vived, in part, Baltatu’s (1939) suggestion of combining these genera as
subgenera under some single generic heading.
Baltatu lumped Pichia and Hansenula together as subgenera of Myco-
derma but his change has not been unequivocally received. To the
writer it seems a mistake to place sporogenous yeasts in subgenera under
an asporogenous genus. It would be a much better practice to combine
the sporogenous genera and to allow the imperfect or asporogenous
genera, in this case Mycoderma, to stand until their perfect stages have
been clearly demonstrated.> Using grape must, Baltatu claimed to have
induced sporulation in fourteen species of Mycoderma. While it is highly
possible that the future might see the discovery of spores in other indi-
viduals of this imperfect group, it is preferable to transfer at such time
the species involved to the perfect group rather than to align them in
their entirety at the present time.
Since the name Pichia has priority over Hansenula in point of time
of establishment and would take precedence in the event of a merging
of these genera, the writer feels justified in adding to the literature the
combinations proposed in this paper. While data on nitrate utilization
are not available on two of the forms discussed, the writer is of the
opinion that the close similarity between the proposed variety Barkeri
and the parent species is sufficient justification for its inclusion here.
To the author, it seems more satisfactory to test for the presence of
nitrite than to report on the extent of growth with nitrate as the sole
source of nitrogen. Excellent tests for nitrite, a reduction product of
nitrate, exist (c. f. Manual of Methods, 1942). By using one such
(sulfanilic acid and a-naphthylamine), it was found that all cultures
of Zygopichia examined were completely negative; 3 species of Pichia
examined were also negative, while 4 species of Hansenula and one of
Zygohansenula gave the brilliant reddish color of a positive test for nitrite,
signifying nitrate reduction. The usual difficulty in testing for nitrate
utilization is that in eliminating sources of nitrogen other than nitrate
from the culture medium, one invariably eliminates all sources of growth-
nutrilites. Since the test is not to see if the yeast can grow in the ab-
sence of “bios,” and since it is rather troublesome and expensive for
routine work to make up a solution containing the known growth nutrilites
as pure chemicals, the author feels it is preferable to permit yeast growth
* See International Rules, Art. 57.
NICKERSON: ZYGOSACCHAROMYCES 481
by adding a trace of yeast extract (0.10 g/liter) to a medium consisting
otherwise of salts, glucose, and KNO3. Yeasts negative for nitrite make
a meager growth, while yeasts that reduce nitrate flourish. Since the
results of the nitrite test are clear-cut, it obviates the necessity of de-
scribing the amount of growth in words. In cases where the nitrite test
was negative, it was found, using the zinc dust test, that nitrate still ex-
isted as such in the medium, thus precluding the rather obscure possi-
bility that nitrate had been reduced beyond the nitrite stage.
WHEATON COLLEGE
Norton, Mass,
BIBLIOGRAPHY
Baars, J. K. Over sulfaatreductie door bacterien. Thesis, Delft. 1930.
Baltatu, G. Mycoderma als echte Saccharomyceten. Zentralbl. f. Bakt. II, 101:
196-225. 1939.
Guilliermond, A. The Yeasts. New York, 1920. (Translated by F. W. Tanner.)
Johnson, F. H., D. Brown & D. Marsland. A basic mechanism in the biological
effects of temperature, pressure and narcotics. Science 95: 200-203. 1942.
Klécker, A. Die Garungsorganismen. Berlin-Wien. 1924.
Manual of Methods for pure culture study of bacteria. Geneva, N. Y., 1942.
Leonian, L. H. & V. G. Lilly. Induced autotrophism in yeast. Journ. Bact. 45:
329-339. 1943.
Mrak, E. M., H. J. Phaff, R. H. Vaughn & H. N. Hansen. Yeasts occurring
in souring figs. Journ. Bact. 44: 441-450. 1942.
Nickerson, W. J. & K. V. Thimann. The chemical control of conjugation in Zygo-
saccharomyces II. Am. Journ. Bot. 30: 94-101. 1943.
Stelling-Dekker, N. M. Die Sporogenen Hefen. Amsterdam. 1931.
1(3): 483-488 FARLOWIA January, 1944
A NEW GENUS OF THE SCLEROTINIACEAE
H. H: WHetzex *
My conservative critics will doubtless raise an eyebrow at this presenta-
tion of another monotypic genus in that group of stromatic inoperculate
Discomycetes for which I have erected the family Sclerotiniaceae.” When
I began my studies on these forms, now some 30 years ago, mycol-
ogists treating any of these species commonly tossed them more or less
indiscriminately into one of three or four genera, most frequently into
Sclerotinia or Ciboria, genera erected by Fuckel back in 1869. My in-
‘vestigations on these forms indicate that there are at least 15 or 20
well marked generic groups in this family. Those which appear to me
to have acceptable generic status are: Sclerotinia’ Fuckel 1869; Ciboria®
Fuckel 1869; Rutstroemia Karsten 1871 (in part) emend. Rehm 1893 ;4
Stromatinia Boudier 1907;5 Phaeociboria von Héhnel 1918;® Lambertella
von Héhnel 1918 (see also Whetzel 1943); Monilinia Honey 1928;
Septotinia Whetzel 1937; Ovulinia Weiss 1940; and Martinia Whetzel
1942. It will be evident to my incredulous reader that I still have a
substantial number of genera up my sleeve which I hope to spring upon
the mycological public shortly. Of the ten above listed genera, four are
as yet monotypic, but as I discovered in the case of Lambertella, they
are likely to lose that status as more intensive searches in the field uncover
undescribed species of the Sclerotiniaceae.
With this orientation of my critical mycological fellows, I shall now
proceed to add another monotypic genus to this growing family.
A COPROPHILOUS SPECIES
While on a collecting trip with some of my students at Malloryville,
New York, on June 22, 1942, I picked up a small dung-ball, about half
an inch long by a quarter of an inch in diameter among leaf mold on
the hillside about the pitcher plant bog. The dung-ball looked like that
of a partridge. It bore several tiny brown apothecial cups borne on the
tips of long hair-like stipes. Ascospore shootings on potato dextrose
agar gave a striking, black, indeterminate stroma (Fig. 2-3) from which
there presently developed numerous long-stipitate apothecia like those
on the original substrate.
1 Acknowledgments: I am grateful to Drs. David H. Linder and W. Lawrence White
for critical suggestions in the preparation of this paper and to Dr. Rolf Singer who
has kindly written the Latin descriptions.
* See Whetzel, 1943, p. 18, footnote 2.
® Both these genera need to be revised and emended with Sclerotinia sclerotiorum
(DBy) Fckl. and Ciboria caucus (Rebent.) Fckl. to be taken as type species
respectively.
“See White, 1941, p. 163, 169.
5 To be restricted to forms similar to the type species, S. rapulum Boud.
®*See Whetzel and White, 1940, p. 614. :
483
484 FarLowlia, Vor. 1, 1944
The dung-ball, meanwhile in a moist chamber, failed to produce more
apothecia and was discarded. The striking characters of the apothecia
which developed so abundantly from the stromata produced on the agar
substratum lead to a careful study of the species which proves to be quite
unlike any other of the fifteen or more genera of the Sclerotiniaceae
known to me. I am, therefore, describing it as a new species and plac-
ing it in a new genus.
Coprotinia gen. nov.
Stromate indistincto, nigro, 1-2 mm. crasso; strato corticali 1— vel pluri-stratoso,
ex hyphis dense intertextis, atris consistente, medullam includente. Structura stro-
matis structurae sclerotiorum Botrytis cinereae simillima. Carpophoris spermatigeris
haud visis. Statu conidiali nulla. Apotheciis gregariis, longis, stipitibus gracilibus
institutis, plus minusve fuscidulis; disco exiguo prae stipitis longitudine, margine fortiter
recurvato in maturis, stipite criniformi, apicibus hypharum glanduliformibus ornato;
ascis minutissimis, clavatis, basin versus gradatim attenuatis, apice incrassatis, poro
iodi ope leviter caerulescente; ascosporis bi-seriatim dispositis, in apicem asci con-
gregatis, minutis, elongate ellipsoideis, uni-cellulatis, hyalinis; paraphysibus cylin-
draceis et tenuiter tunicatis.
Stroma (on potato dextrose agar) indeterminate, black, 1-2 mm. thick;
rind (external and submerged) one to several layers of densely inter-
woven, slender, dark brown to black hyphae, enclosing a medulla; medulla
of rather closely interwoven, slender, thin-walled hyphae embedded in
a rubbery transparent matrix. Stromatic structure very like that of the
sclerotia of Botrytis of the cinerea type. Spermatial fruit-bodies not seen.
Conidial stage wanting. Apothecia gregarious, long, slender-stipitate,
some shade of brown; disc small in comparison with length of stipe, the
margin strongly recurved at maturity, stipe hair-like, the surface adorned
with scattered glandular hyphal tips; asci very small, clavate, tapering
gradually to the base, apex thickened, pore faintly J++; ascospores
biseriate, crowded into the swollen apical part of the ascus, minute,
long ellipsoid, one-celled, hyaline; paraphyses cylindric and thin-walled.
Coprotinia minutula sp. nov.
Stromate diffuso, indistincto, nigro, 1-2 mm. crasso; strato corticali ex hyphis
dense intertextis, atro-fuscis, gracilibus consistente et uno vel nonnullis stratis com-
posito, extus asperulato; medulla ex hyphis intertextis, ramosissimis, anastomosantibus,
gracilibus, septatis, hyalinis, 3-3.5 u crassis efformata. Carpophoris spermatigeris
haud observatis. Statu conidiali nullo. Apotheciis numerosis e superficie tota
stromatis nusquam non enatis, longissime stipitatis; receptaculo 1-2 mm. in diametro,
atrocastaneo ad primam maturitatem, vetustis expallentibus usque ad colorem
pallide fulvidum, tenui, margine prima juventute fortiter recurvato; stipite
gracillimo, criniformi, usque ad 20 mm. longo et % mm. crasso, sursum pallido,
atro deorsum, apicibus hypharum glanduliformibus ornato; ascis brevibus, crassius-
culis, clavatis, saepe conspicue inflatis sub ipso apice, 31-47 x 3.64.8 uw, apice ro-
tundato, incrassato, poro iodi ope leviter caerulescente; ascosporis 8, minutis, bi-
seriatim dispositis et in tertiam superiorem inflatam congregatis, elongate ellipsoideis,
4.8-6.0 x 1.5-2 y, hyalinis; paraphysibus cylindraceis, tenuiter-tunicatis nec septatis
nisi ad basin, 3-3.5 x 33-45 uw.
Stroma (on potato dextrose agar) diffuse, indeterminate, black, 1-2
mm. thick (Fig. 2-3) ; rind (external and submerged) composed of densely
Joe
i ea ee aaa
Fig. 1-3. Coprotinia minutula: 1, Apothecia produced in culture, X 2; 2,
Stroma formed on potato dextrose agar, photographed from upper surface, X 1; 3,
Same as 2 but photographed with Petri dish inverted, x 1.
486 Fartowla, Vor. 1, 1944
interwoven dark brown, slender hyphae, one to several layers thick, the
external portion here and there double with narrow strip of medulla be-
tween the two layers; the surface rough and where not imbedded in the
agar covered with a felty layer of loose brown hyphae; medulla of rather
densely interwoven, much branched and anastomosed, slender, thin-walled,
5
Fig. 4-6. Coprotinia minutula: 4, Diagram of section through disc showing
its general form at complete maturity, < 80 approx.; 5, Longitudinal section from
lower part of stipe showing structure of stipe and protruding hyphal tips, x 1290;
6, Asci, paraphyses, and ascospores, « 1290.
septate, hyaline hyphae, 3-5.5 » in diam., embedded in a transparent
rubbery matrix. Spermatial fruit-bodies not observed. Conidial stage
wanting. A pothecia arising in large numbers from all over the surface
of the stroma (Fig. 1), very long stipitate; receptacle 1-2 mm. in diam.,
dark chestnut brown when first mature, bleaching out to “pale ochraceous
tawny” (R) with age, thin, the margin very soon strongly recurved giving
the appearance of tiny toadstools; stipe very slender, hair-like, up to
20 mm. long by 1% mm. thick, pale above, dark below, adorned with
short glandular hyphal tips; asci short, stout, clavate, often markedly
WHETZEL: NEw GENUS OF SCLEROTINIACEAE 487
inflated or swollen just below the tip, 31-47 x 3.6-4.8 yu, apex rounded,
thickened, pore faintly J+-; ascospores 8 per ascus, minute, biseriate and
crowded into the swollen upper third of the ascus, narrow ellipsoid,
4.86.0 x 1.5-2.0 » hyaline; paraphyses cylindric, thin-walled and aseptate
except at the base, 3-3.5 x 33-40 yp.
Hasirat: Collected only once on a small dung-ball of some unkown
animal at Malloryville, N. Y., June 22, 1942. Known chiefly from
stromata and apothecia developed in cultures derived from ascospore
discharge on potato dextrose agar and on sterilized wheat in laboratory
at Ithaca, N. Y.
Type Specimen: C. U. Pl. Path. 31585. Stromata developed on potato
dextrose agar and on sterilized wheat kernels. —
This little discomycete presents several distinctive characters. Copro-
philous species in the Sclerotiniaceae are rare. The only other species
known to me to occur on dung is the form recorded by me” but collected
by R. F. Cain on rabbit dung near Lac St. Joseph, Quebec, on August
25, 1938 (C. U. Pl. Path. 29646). That form I have referred to Mar-
tinia panamaensis Whetzel. The species here described is, however, quite
different since the apothecia develop from a diffuse stroma rather than
from tiny distinct hemisphaerical sclerotia, and the spores are hyaline
instead of colored.
In certain of its aspects, Coprotinia minutula, reminds one at once of
certain species of Lambertella. It cannot, however, be referred to that
genus, since its stroma is of a more highly developed type, being in the
structure of its rind and that of its medulla much more like the sclerotium
of a Botrytis of the cinerea type. This, together with its colorless spores,
sets it off not only from Lambertella but also from Martinia, although
its apothecia, asci and ascospores are strikingly like those of the latter
genus. It is perhaps most closely related to Rutstroemia, but the slender
' hair-like stipes of the apothecia would seem to deny reference of it to
that genus. One of its most peculiar characters is the gland-like hyphal
tips which adorn the stipe throughout its length and over the underside
of the receptacle. These are of variable length, usually of 1 to 5 short
cells, the apical cell being swollen and filled with an olivaceous-green
substance of an oily character (Fig. 5).
DEPARTMENT OF PLANT PATHOLOGY ~
CornELL UNIVERSITY
IrHaca, NEw York
7™See Whetzel, 1942, p. 589.
488 FartowlA, Vou. 1, 1944
LITERATURE CITED
Boudier, E. Histoire et classification des Discomycetes d’Europe, p. 1-221. (Paris)
1907,
Fuckel, L. Symbolae mycologicae, p. 1-459. (Wiesbaden) 1869,
Hohnel, F. von. Humarina tetrica Quel. In Rehm: Ascomycetes Exs. Fasc. 56
and 57. Ann Myc. 16: 219-220. 1918.
Honey, E. E. The monilioid species of Sclerotinia. Mycologia 20: 127-157. 1928.
Weiss, Freeman. Ovulinia, a new generic segregate from Sclerotinia. Phytopath.
30: 236-244. 1940.
Whetzel, H. H. Septotinia, a new genus of the Ciborioideae. Mycologia 29:
128-146. 1937.
. A new genus and new species of brown-spored inoperculate Discomycetes
from Panama. Mycologia 34: 584-591. 1942.
. A monograph of Lambertella, a genus of brown-spored inoperculate Dis-
comycetes. Lloydia 6: 18-52. 1943.
and White, W. L. Mbollisia tetrica, Peziza Sejournei and the genera Phaeo-
ciboria and Pycnopeziza. Mycologia 32: 609-620. 1940.
White, W. L. A monograph of the genus Rutstroemia (Discomycetes). Lloydia
4: 153-240. 1941.
1(3) : 489-490 FARLOWIA January, 1944
BOOK REVIEWS
Tue BoretaceaE or NortH Carouina. By William Chambers Coker and Alma
Holland Beers. Chapel Hill, University of North Carolina Press, 1943. viii, 96 pp.
pl. .1-65, frontispiece. $7.00.
The authors include three genera in this treatment: Boletus, Boletinus, and Strobilo-
myces, all understood in a conservative sense, i.e., Boletus, by far the largest, with
seventy species indicated and described for North Carolina, Boletinus with five species,
and Strobilomyces with one species. The book is abundantly illustrated with many
good and some excellent photographs, six plates in colors, five plates of spore draw-
ings and one text illustration, showing the anatomy of several species. Some of the
colored pictures are quite natural, and thus very helpful to the beginner. The micro-
scopical pictures, however, are much too schematic and therefore of comparatively
little help. Cystidia (with one exception), basidia, tramal hyphae, the covering
layer of pileus and stipe, as well as chemical reactions are neither illustrated nor
mentioned in the descriptions. The exact color of the spore print, a frequently neg-
lected but very valuable character, is correctly indicated in nearly all descriptions.
Two keys, both mainly based on macroscopical characters, are useful for everyone
who tries to determine Boletes, although in a few instances they may lead the less
experienced into trouble because they are not dichotomous, and therefore difficult
to use.
Those who are not primarily interested in taxonomy may regret the lack of com-
mon names in the book, as well as the lack of indications concerning the practical
interest of the species described. The authors mention only casually that “this is
one of the safest groups ... for the beginner to experiment with, as only a few
species are now thought to be poisonous.” The warning to keep hands off the whole
Miniato-olivaceus group and all of the Luridi can be heartily subscribed to. Many
a lay reader, however, may desire more information on the value of the single species
for gastronomic purposes, while foresters and ecologists would appreciate more spe-
cific data on the important problem of mycorrhizal relationship between forest trees
and Boletaceae.
The taxonomist, on the other hand, will find it difficult to follow the authors in
many of their solutions of questions of nomenclature as well as in the general ar-
rangement of the species. He will also regret the little space that has been devoted
to synonymy, and the absence of Latin diagnoses for the new species and varieties
proposed (Boletus felleus var. minor var. nov., B. satanas var. americanus var. nov.,
B. viridiflavus spec. noy., B. sub-fraternus spec. nov., B. parvulus spec. nov., B. amer-
icanus var. reticulipes var. nov., B. subaureus var. siccipes var. nov.). The interpre-
tation of some species of Boletus does not seem to be in accord with the original
concept of their authors, as, for example, B. luridus Schaeffer which is evidently
different from the plant described under this name in Europe, as the authors them-
selves seem to feel.
Notwithstanding certain shortcomings mentioned above, the book contains much
valuable information for a wide variety of people interested in the study of Higher
Fungi, and should be warmly welcomed not only in North Carolina, but also in other
Eastern States, and especially in the South where the Boletaceae-flora, though rich
and interesting, has never been monographed before. —R. S. and D. lak, bes
PLANTS AND Vitamins. By W. H. Schopfer [translation by N. L. Noecker] Waltham,
Mass. Chronica Botanica. 1943. ix, 293 pp. illus. $4.75.
It may appear unusual to review in a taxonomic journal a book which deals primar-
ily with a physiological subject, but after reading the book on plant vitamins, one
can not help but be impressed by the part that taxonomy has had in the development
of this field of research, and conversely, one is equally struck by the service that a
knowledge of vitamins may render to taxonomy by making possible the more normal
489
490 FarLowla, Vou. 1, 1944
growth of organisms that have hitherto been grown on incomplete nutrient substrata.
Anyone who has tried to determine fungi from cultures can readily realize what it
would mean to have a medium on which these organisms would grow and behave as
they do in nature. Thus, for those who deal with organisms in culture, whether or
not they have sufficient chemical knowledge to understand the chemistry of vitamins,
there may be found ample reason for reading this book and garnering facts that
will be of service in their investigations since the author deals not only with the réle
of vitamins in the nutrition of the higher and lower plants, but also introduces facts
and suggestions on the part these substances play in symbiotism and parasitism. But
lest it be thought that only taxonomists may find food for thought in this excellently
presented and up-to-date summary of our knowledge of plant vitamins, it should be
emphasized that throughout the work the author, while dealing with the occurrence,
nature, and function of vitamins and kindred substances, has throughout presented
many suggestions which should inspire further research in the allied fields of physi-
ology, biochemistry, and genetics. Biologists are greatly indebted to the author for
this valuable compilation of facts that have been presented in a careful yet stimu-
lating manner, and to the publisher and translator who have made this work available
in English at a time when biological discoveries may mean so much. — D. H. L.
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MANUSCRIPT
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Contributors should strive for a clear concise style of writing. A limit of 100
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Vol. 1 JULY, 1944 No. 4
FARLOWIA
_ A JOURNAL oF CrypToGaMic BoTANy
EptroriaL Boarp
DAVID H. LINDER, Editor
E. V. SEELER, Jr., Mg. Ed.
J. R. BartHOoLOMEW R. Sincer
E. B. Bartram F. VerRDOoRN
N. F. Conant W. H. Weston, Jr.
H. F. Harris
R. M. WrEetpen
PUBLISHED BY
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Four numbers compose one volume of approximately five hundred pages. i
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Subscriptions and remittances should be addressed to FARLOWIA, 20 Divinity
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period.
CONTENTS OF VOL. 1, NO. 4
Marine Puyrocrocrapuy. By Valerie May.. Ss Gace eo ae
GALAXAURA, A Dir.opiontic FLoRIDEAN GENUS WITHIN THE Oeine NEMALIO-
nates. By Nils Svedelius........... Ce erat Oe ae a ieG ee es ees 495
MorruHotocic AND Puysiotocic InvesticaTions In Diatoms. By Sten
Wiedling. . Pincge eee ceed cl eres ey ee ee yikes 501
NEw AND NotEwortTHy Pururerinse Mosses. By Edwin B. Bartram ii vee OU:
On Two New Sprores or Mncaceros wits Norns on M. ARACHNOIDEUS, M.
Denticutatus, M. GIGANTEUS, AND M. Granvis. By L. P. Khanna...... 515
SruprEs oN Po.ysticrus Crrcinatus AND Its RELATION To THE Burt-Rot or
Spruce. By Roger Gosselin: .........ececececneees Bo Peat a eee .. 525
Norss on Wisconsin Parasitic Funai. IV. By H.C. Greene............ .. 569
A Discussion or Taxonomic Crrreria IN THE a ete gs By Alma J.
WGOn §. Ooo eee eRe, hn, Cocoa Pade a a yk eee ahs CV eee 583
SruDIESs IN THE GENUS HELOTIUM, IV. By W. Lawrence White. . Viiercs ORE
WPEURG: Ce re races ees pias aen Haha Peoioa ae Ms cee ee
Inprex To VotuME1........ ak, acs Bae ea co veers viet ete OM
Vol. 1, No. 3 was issued on January 15, 1944.
CORRECTION
Add 9 INDEX TO AUTHORS AND TITLES, Farlowia 1(4): 625, July, 1944.
Ernst-Schwarzenbach, Marthe. The sexual dimorphism of the tropical mosses
of the genus Macromitrium, 195-198.
Aor
Wha -
FARLOWIA
A JOURNAL OF CRYPTOGAMIC BOTANY
wk | JuLy, 1944 No. 4
: MARINE PHYTOGEOGRAPHY
VALERIE May
From a comparative study of marine floras we may at some date be
able to demonstrate the evolution of sea plants in both time and space.
At present it is impossible to complete this comparative study since
the algal exploration of many parts of the world is incomplete. Further,
algal taxonomy lags far behind the corresponding study of. land plants.
Thus any theory of phytogeography based on our present knowledge
of marine plants rests on an insecure foundation.
Variation in algal communities, within small areas, can be explained
satisfactorily in terms of changes in exposure to dessication or heavy
seas, changes in mineral and gaseous supplies, in intensity of light and
heat, depth of water, and the nature of the sea floor. On this there are
superimposed seasonal changes in the flora and interaction or competition
between organisms.
_ When a comparison is made of more widely separated districts, how-
ever, such explanations are inadequate. The original conception of
algal distribution was that each species had a centre of growth from
which it was disseminated following the lines of the coast, each species
being influenced by local factors such as depth of water. This idea ac-
counted for local richness in certain groups of the flora and also the
similarity between the algal floras bordering an ocean. Exceptions were
quickly noted, however, many plants (e.g. Codium tomentosum) being
- reported as cosmopolitan in distribution. Later work, as explained be-
low, has suggested that this supposed cosmopolitan distribution may, in
at least some cases, be due to taxonomic errors.
By 1849 we find Harvey coming to the conclusion that temperature
zones were highly important. Thus, in number of species, the “browns”
(Melanophyceae) predominated in the tropics, the “reds” (Rhodophy-
ceae) reached a maximum in the temperate zones, and the “greens”
(Chlorophyceae) were polar or universal.
Zoning by temperature may well be a misnomer for zoning by light
intensity, as has been emphasized by Tilden.
Besides this zoning by latitude, a certain amount of data indicates
zoning by longitude; for instance in the Codiaceae, A. & E. 5. Gepp (1911)
state that five large genera are widely distributed in both hemispheres,
491 .
492 FarLowla, VoL. 1, 1944
seven genera are in the Eastern Hemisphere only, one in the West Indies
region and one in the Mediterranean Sea. Only very few species occur
in both hemispheres. This grouping would seem due to some barrier
(past or present) preventing east-west migration.
Discontinuity of distribution has been attributed to the presence, or
past existence, of barriers such as land masses, ocean deeps, or sandbank
deserts. Likewise, wide dispersal has been explained as due to distri-
bution by birds, ships, etc., and by ocean currents (speed as well as di-
rection being important because the time of life of motile spores is
limited). Svedelius (1924) suggested that the present distribution of
algae may be due to geological changes in land and sea allowing migra-
tion by way of water connections from the Indian Ocean to the Mediter-
ranean Sea via Suez, and from the Pacific to the Atlantic Ocean via
Panama. Our present knowledge is too meagre to test this theory.
Fossil evidence is unsatisfactory because algal forms are usually soft,
and hence their remains are scarce except for the specialized calcareous
seaweeds of many so-called “coral” reefs. Lucas (1927) reported the
discovery of Bytholotrephis gracilis James & Hall, from Victoria, Aus-
tralia, “apparently identical” with forms found in the Lower Ordovician
beds of North America. Another fossil species was like one found in
England. Lucas assumed that Bytholotrephis was an “elemental or gen-
eralised type,” dominant in England, North America, Australia, etc.
This idea of an elemental or generalized type represents a theory that
at one time uniform conditions and marine floras existed throughout the
world. Later more temperate conditions prevailed north and south of
the tropics and the floras of these zones, subjected to similar environ-—
ments, changed along similar lines. Finally, it was suggested that, in
the frigid zones, we would have identical representatives which were
absent from temperate or torrid zones. This argument may hold for
plankton (which are not considered in this article) but Gain (1912)
has shown that great differences exist between the marine floras of the
Arctic and Antarctic. For instance, he noted that not one species of
the large groups of Fucaceae and Laminariaceae was common to both
zones, although all species of these groups belong to cool regions.
May (1940) applied Jaccard’s Coefficient of Correlation to the marine
floras (Chlorophyceae and Melanophyceae only) of Australia, New Zea-
land, Britain, and North America; also of Australia-Cape of Good Hope.
From figures published by Gain (reference given above) the coefficient
percentages for the Arctic and Antarctic floras are:
Chlorophyceae 10.5 Melanophyceae 6.9 Rhodophyceae 7.8
All these coefficients are surprisingly alike and show that the degree of
uniformity of the floras of the places examined is fairly constant. The
_ correlation tends to be higher among the Chlorophyceae than the Melano-
phyceae. The Chlorophyceae include more so-called “cosmopolitan”
species.
May: Marine PHYTOGEOGRAPHY .493
It seems the wide distributions reported for so-called “cosmopolitan”
plants may very well be due to faulty systematic work. Setchell (1914)
examined Scinaia furcellata, previously reported as “cosmopolitan.” He
found that this species was in reality limited in distribution. Plants re-
ported under this specific name 4s occurring in other districts were either
different species or even different genera. Thus the “cosmopolitan”
distribution resolved itself into there being a group of more or less closely
related species, each occupying its own limited region.
Ectocarpus confervoides, on the other hand, appears to be truly cos-
mopolitan, but different phases of its life history predominate in different
places. The reason for this is not understood, although the appearance
of many algae is known to change with habitat, i. e. the form assumed
depends on the environment.
Weber van Bosse (1904) reported that many characters used in classi-
fying plants “were not implicitly to be relied upon in all species.” These
characters included manner of ramification, diameter of conceptacles,
and the number or manner in which they appear on the joints, characters
which are still widely used. Perhaps the present unsatisfactory con-
dition of algal systematics could best be overcome by more care being
paid to the criteria used in classification. Certainly we need more
knowledge of the specific limits of the plants we deal with before a
phytogeographic comparison between floras can be satisfactory.
To conclude, phytogeography of marine plants is an almost untouched
field. Without more work of systematic nature and further collections
being made, this study is unable to advance beyond vague. hypotheses.
All we can say with certainty is that latitude and the presence or absence
of barriers (either geological or present-day) between suitable habitats
appear to control the distribution of marine vegetation.
- NATIONAL HERBARIUM
SYDNEY, AUSTRALIA
BIBLIOGRAPHY
Gain, L. La Flore Algologique des Régions Antarctiques et Subantarctiques.
Deuxiéme Expédition Antarctique Francaise (1908-1910) Commandée par le
Dr. Jean Charcot. 218 pp. 8 pl. Masson, Paris. 1912.
Gepp, A. & E. S. Gepp. The Codiaceae of the Siboga Expedition. Monographie
LXII of: Uitkomsten op Zoologisch, Botanisch, Oceanographisch en Geologisch
Gehied. 150 pp. 22 pl. Leiden. 1911.
Lueas, A. H. S. On Additional Occurrence of Bytholotrephis in Victoria. Mem.
Nat. Mus. Melbourne No. 7: 157-158. pl. 14. 1927.
May, V. A Comparison between Marine Floras. Contrib. N. 5S. Wales Nat. Herb.
1 (2): 94-98. 1940.
Setchell, W. A. The Scinaia Assemblage. Univ. Calif. Publ. Bot. @ (5): 79-152.
pl. 10-16. 1914.
Svedelius, N. On the Discontinuous Geographical Distribution of Some Tropical
and Subtropical Marine Algae. Ark. f. Bot. 19 (3): 70 pp. 1924.
Weber van Bosse, A. & M. Foslie. The Corallinaceae of the Siboga-Expedition.
Monographie LXI of: Uitkomsten op Zoologisch, Botanisch, Oceanographisch en
Geologisch Gehied. 110 pp. 16 pl. Leiden. 1904.
1(4): 495-499 FARLOWIA July, 1944
GALAXAURA, A DIPLOBIONTIC FLORIDEAN GENUS
WITHIN THE ORDER NEMALIONALES
Nits SVEDELIUS
One of the most remarkable discoveries in the sphere of floridean re-
searches was the observation of M. A. Howe (1917, 1918) that in the
genus Galaxaura there is a structural dimorphism of sexual and tetra-
sporic plants. This was previously quite unknown in this algal group
and the discovery was the more remarkable since this genus belongs to
the order Nemalionales among the Florideae, which order comprehends
especially haplobiontic types, i. e., such types in which the reduction
division occurs immediately after the fertilization of the female nucleus
and in which there is therefore no tetraspore formation whatsoever. This
structural dimorphism in respect to the anatomy was naturally quite un-
known to F. R. Kjellman when, in the year 1900, he published his com-
prehensive treatise of the genus Galaxaura and to a great extent based
the diagnoses of the sections in particular on anatomical characters. These
sections of Kjellman contain either purely sexual or purely tetrasporic
plants. After Howe’s discovery, this classification was quite untenable
and it is now an important task for algological research to unite Kjell-
man’s different species which are, as we now know, only different gen-
erations of one and the same species. The consequence of all this is
that a close investigation of the development of Galaxaura is highly neces-
sary, the more so as the divergent anatomy of the sexual and the tetra-
sporic plants also must be made clear from the developmental point of view.
I have delivered a contribution to the solution of these problems in a
paper “Zytologisch-entwicklungsgeschichtliche Studien tiber Galaxaura,
eine diplobiontische Nemalionales-Gattung” (1942) and I will here give
a short account of its main results. This seems to be all the more re-
quired as Galaxaura in many other respects too proved to be a new and
rather divergent type of organization amongst the Florideae.
By my investigations, I believe I have established that the sexual and
the-tetrasporic plants are also distinguished by a certain difference re-
garding their ramification and succession of shoots. The sexual plants
have a more lively ramification caused by proliferations from new grow-
ing points, whereas the tetrasporic plants have poorer, mainly dichotomous
ramification and a’ more continuous apical growth. This seems to depend :
upon the fact that the fertile shoots of the sexual plants are commonly
almost totally thrown off after fertilization, whereas the tetrasporic shoots
are not deformed to such a high degree by the formation of tetraspores,
but can survive and continue their growth.
The dimorphism regarding the anatomy that Howe first observed is
restricted to the assimilating layer of the cortex, and depends on a very
early differentiation, just below the growing point, which appears in such
a way that the cortex of the tetrasporic plants produce still another cell-
layer besides those belonging to the sexual plants. The assimilating cells
495
496 Fartowia, VoL. 1, 1944
of the tetrasporic plants are supported by special stalk-cells and the cells
lying between these stalk cells develop in another way. Either they
become stretched laterally with the result that the assimilating cells are
isolated from each other so that small cavities are formed between them,
or else they develop into big, rounded, colorless cells between the stalk-
cells.
Earlier statements about the cystocarp of Galaxaura are very scarce
and, moreover, quite false. Thus the statement that the cystocarp of
Galaxaura has an involucre or pericarp formed from the lowest cell of
the carpogonial branch is incorrect and derived from a false observation.
In this, Galaxaura differs from both Scinaia and Gloiophloea, Pseudo-
scinaia and Actinotrichia. On the other hand, Galaxaura in this respect
agrees with Chaetangium (Martin 1939).
The side branchlets of the three-celled carpogonial branch develop very
early and become large, and rich in plasmatic content, but with very
weakly staining nuclei. Further, all the cells of side branchlets get fer-
tilized diploid nuclei. After fertilization, the diploid nucleus either
enters directly into the hypogynous cell (G. corymbifera) or first divides
once in the carpogone itself, after which the daughter nuclei migrate into
the different inferior cells (G. Diesingiana). In the emigration of the fer-
tilized nucleus from the carpogone, Galaxaura corresponds with Scinaia
and Chaetangium, but differs from these genera in an important respect:
no immediate reduction division occurs in Galaxaura, which thus is a
diplobiontic genus within the Nemalionales having special diploid tetra-
sporic plants with the reduction division at tetraspore formation. This
I have plainly observed in G. Diesingiana and G. tenera.
When the diploid nuclei have entered the hypogynous cell, a lively
nuclear division arises, and these diploid nuclei enter all the cells of the
branchlets of the carpogonial branch except the empty carpogonium itself.
Open pore connections now arise between the hypogynous cell and the
cells of the side branchlets. The gonimoblast filaments soon begin to
develop and elongate in different directions, namely: 1) straight upwards,
i.e., through the basal part of the carpogonium; 2) laterally from the
side branchlets of the hypogynous cell; 3) from the first or lowest cell
of the carpogonial branch. It ought to be pointed out that there is a
differentiation between such gonimoblast filaments as grow straight up-
wards and begin the carpospore formation immediately, and those that
later on grow out laterally, giving rise to secondary gonimoblast centra.
Finally, there are also formed sterile nutritive filaments which grow
downward and seek out the bigger leading cells in the center of the alga,
which cells become twined round by these nutritive filaments. All gonimo-
blasts, the primary as well as the secondary, have, however, a common
ostiole through the calcified cortical layer. The unity of the cystocarp
is maintained in spite of several distinct gonimoblast centers.
Regarding the formation of the carpospores, it is to be observed that,
after they have been dispersed, regeneration generally takes place so that
SVEDELIUS: GALAXAURA ' 497
new sporangia are formed underneath the emptied ones. On the other
hand, no paraphyses are formed, just as in Scinaia, Statements about
paraphyses in Galaxaura in the literature are false, as are the statements
about a special pericarp formed by filaments from lower cells of the
carpogonial branch. ‘
Thus the formation of the cystocarp in Galaxaura has been found to
be a type which is quite new among the Rhodophyceae, and characterized
by the fact that the whole carpogonial branch receives diploid nuclei, and
that the whole carpogonial branch also forms starting-points for the
gonimoblast. The carpogonial branch can be comprehended as a com-
pound auxiliary system. The whole gonimoblast can be divided into
primary and secondary gonimoblasts whereby the primary is the one
which is formed in connection with the central fusion-cell, and the sec-
ondary the one originating from the centers which have proceeded from
the horizontally extending gonimoblast filaments. :
Galaxaura is also remarkable for the fact that its spermatangia are
formed in deep cavities, a very exceptional case among the Rhodophyceae.
An investigation of the development has proved, however, that the very
earliest stages correspond rather well with those of Scinaia, but in Scinaia
the spermatangial branches protrude and push between the epidermal
cells to become superficial. In Galaxaura the development proceeds in
another direction. The branching is especially lateral, and the sur-
rounding cortex. cells stretch around the spermatangial branchlets with
the result that they very early become depressed in a deep cavity, the
walls of which are covered by the spermatangial branchlets. However
different the spermatangial branches in Scinaia and Galaxaura may still
seem in fully developed stages, when the development has been clarified,
they can without difficulty be classed under a uniform scheme or type
of organization.
The genus Galaxaura belongs among the very few genera amongst the
Nemalionales which are characterized by the formation of tetraspores.
As it has now been proved that Galaxaura is a real diplobiont, there is of
course no reason to suppose anything else than that the tetraspore forma-
tion is also combined with a reduction division. I have also been able
to prove this reduction division in two species belonging to two different
sections, namely G. Diesingiana and G. tenera. It proceeds normally
and the nucleus of the tetraspore mother-cell passes through prophase
with spireme and diakinesis with about ten bivalents (gemini).
The position of the tetrasporangia varies in different species and on
this a division of the genus Galaxaura into sections may be founded better
than on the older one which was. based mainly upon anatomical char-
acters. Preliminarily, I divide Galaxaura into three sections, namely:
Sect. 1. Rhodura (Kjellm.) Sved. emend., characterized by quite sessile sporangia
formed terminally or laterally upon rather long assimilation filaments. The sporangia
never regenerate.
498 Fartowia, Vou. 1, 1944
Sect. 2. Acrosporangiatae Sved., characterized by terminal sporangia formed upon
assimilating cells from a special small stalk-cell. After the dispersion of the tetra-
spores, the stalk-cell forms a new tetrasporangium in which the nucleus passes through
a new reduction division and a new tetrad is formed. This process is repeated
many (5-6) times, and after each regeneration the basal part of the regenerated
sporangium always remains as a collar at the base of the new sporangium. Some-
times a new and purely vegetative cell is interpolated between the sporangia.
Sect. 3. Pleurosporangiatae Sved., characterized by the constantly lateral sporangia
originating from the assimilating cells by means of a special stalk cell. Regenera-
tion occurs here also but, in contrast to the section Acrosporangiatae, only once.
The repeated regenerations combined with repeated reduction divisions
is very remarkable and with no equivalent in any other floridean genus.
An investigation of the closely related chaetangiaceous genera Scinaia,
Chaetangium, and Galaxaura has shown that the first two genera are
haplobiontic and lack tetraspore generation, whereas Galaxaura is diplo-
biontic with special tetrasporic plants. According to statements in the
literature, Gloiophloea and Pseudoscinaia are very probably haplobionts,
whereas, as I have now shown, Galaxaura is diplobiontic with special
tetrasporic plants. Actinotrichia is also diplobiontic. Thus it is evident,
just as I have already pointed out in my Scinaia paper of 1915, that we
must be very careful to use haplo- and diplobiontic organization as a sys-
tematic character for characterizing the larger groups. Closely related
genera behave differently in this respect and in the genus Chantransia
even different species behave differently. Thus it is impossible to make.
haplobiontic organization into a main character for the order Nemalionales
even if the majority of its genera are haplobionts. It seems to me that
this character must not be accorded higher taxonomic value than, for
example, homospory and heterospory for the taxonomy of the Pterido-
phyta. We may say that in the order Nemalionales we find amongst the
Rhodophyceae the transition types between haplobiontic and diplobiontic
organization, that is, the organization which later becomes the predom-
inant one of the higher Red Algae.
Regarding the special taxonomy of the family Chaetangiaceae I here
give a revised classification in which the two subfamilies Scinaiae and
Chaetangieae are maintained with partly altered diagnoses:
Subfamily I. Scinaieae (Trevis.) Sved. emend.
The lowest cell of the carpogonial branch forms a sterile pericarp around
the gonimoblast, i.e. the cystocarp is provided with a wall. The spermatangia
are generally superficial.
A. Haplobionts: Scinaia, Gloiophloea, Pseudoscinaia.
B. Diplobiont: Actinotrichia.
Subfamily II. Chaetangieae (Kuetz.) Sved. emend.
The sterile pericarp about the gonimoblast lacking; gonimoblast developing
from nearly all cells of the carpogonial branch, the cystocarp is thus without
wall. Spermatangia are immersed in cavities.
A. Haplobiont: Chaetangium.
B. Diplobiont: Galaxaura.
It appears from this system that diplobiontic organization must have
SVEDELIUS: GALAXAURA 499
arisen at different occasions during the evolution of the Chaetangiaceae.
This seems to me to be of great interest and it is also of fundamental im-
portance for the real understanding of the evolution of the plants from
haploid to the prevailing diploid organization.
Unrversity oF UppsaLa
UppsaLa, SWEDEN
LITERATURE CITED ©
Howe, M. A. (1917). A note on the structural dimorphism of sexual and tetra-
sporic plants of Galaxaura obtusata. Bull. Torrey Bot. Club 43: 621-624.
(1918). Further notes on the structural dimorphism of sexual and tetra-
sporic plants in the genus Galaxaura. Mem. Brooklyn Bot. Gard. 1: 191-197.
pl. 3, 4.
Kjellman, F. R. (1900). Om floridé-slagtet Galaxaura, dess organografi och syste-
matik. K. Svenska Vet. Akad. Handl. 33 (1): 109 p. pl. 1-20.
Martin, M. T. (1939). The structure and reproduction of Chaetangium saccatum
(Lamour.) J. Ag. II. Female plants. Jour. Linn. Soc. London (Bot.) 52:
115-144, pl. 1.
Setchell, W. A. (1914). The Scinaia assemblage. Univ. Calif. Publ. Bot. 6 (5):
79-152. pl. 10-16.
Svedelius, N. (1915). Zytologisch-entwicklungsgeschichtliche Studien iiber Scinaia
furcellata. Nova Acta Reg. Soc.-Scient. Upsaliensis. Ser. IV. 4 (4): 1-55.
- (1939). Uber den Bau und die Entwicklung der Spermatangiengruben
bei der Florideengattung Galaxaura. Bot. Notiser 1939: 591-606.
- (1942). On the development of the cystocarp in the genus Galaxaura
and the auxiliary cells in the order of Nemalionales. Blumea. Suppl. II
(Dr. A. A. Weber van Bosse Jubilee Volume) p. 72-90. :
- (1942). Zytologisch-entwicklungsgeschichtliche Studien tiber Galaxaura,
eine diplobiontische Nemalionales-Gattung. Nova Acta Reg. Soc. Scient. Up-
saliensis. Ser. IV. 13 (4): 1-154. r
Weber van Bosse, A. (1913-28). Liste des Algues du Siboga. Siboga-Expeditie
Monog. 59 (a—d): 1-533. pl. 1-16.
*
Wa. aa ee ee Go) ls fee”. eee ee ee Wg? gee ge eee nl eT Sr gre ae Cae Pas eel Sap. ake RL a ee Prin oe ey |
Ba oN “ oy recy een Ee Pe ae ee AOS SSR eT. aetna cee aay ne ete Ae yee Air Shel Mie ae : A kt ie he
1(4) : 501-502 FARLOWIA July, 1944
MORPHOLOGIC AND PHYSIOLOGIC INVESTIGATIONS
IN DIATOMS
STEN WIEDLING
Having earlier studied culture methods for planktonic fresh-water algae,
especially diatoms, the author in 1934 started an experimental investiga-
tion of the Nitzschiae with the intention of finding out the conditions of
auxospore formation in these diatoms. Both fresh-water forms and those
from brackish and salt water were cultured (Botaniska Notiser 1941:
37. 1941). The list of species according to the determinations of Dr.
Astrid Cleve-Euler, the grand old lady of Scandinavian diatomology, com-
prises the following forms: Nitzschia ovalis Arnott fa., N. subtilis var.
paleacea Grun., N. palea var. debilis (Kg.) Grun., N. palea var. tropica?
Grun., N. Riuvemeiona Hilse, N. Kiitzingiana var. exilis Grun., and N.
communis Rabenhorst.
In all, about twenty-five strains were kept and some of them are still
being cultured. Three standard substrates were used, all of them based
on agar and inorganic salts; ferric citrate, however, was added as an iron
source since Carsten Olsen in 1929 demonstrated its superiority to inor-
ganic iron salts in plant cultures. For the fresh-water forms, distilled
or tap water was added and for the other forms surface water from the
Sound was found suitable (cf. Harvey, J., Marine Biol. Assoc. 23:
499, 1939).
The results of this investigation on the reproduction of the Nitzschiae
were partly not according to expectation (Naturwiss. 31: 115. 1943).
Several strains as those of N. subtilis var. paleacea Grun., N. Kiitzingiana
Hilse, and N. palea var. debilis (Kg.) Grun. did not show any decrease
in the apical length, which instead was constant. This experience is an-
other evidence against the universal significance of the MacDonald-Pfitzer
rule for diatom variation (J. D. MacDonald, Ann. Mag. Nat. Hist. 3: 1.
1869; E. Pfitzer, Sitzungsber. Niederrhein. Ges. Natur-Heilkde, Bonn
1869: 71; Bot. Zeitung 27: 779. 1869). Other diatoms with constant
apical length were earlier found by Allen and Nelson (N. closterium
W. Sm. fa. minutissima, Journ. Mar. Biol. Assoc. 8: 421. 1910) and
Geitler (Eunotia pectinalis var. minor (Kiitz.) Rabenh., Arch. Protistenk.
78: 1. 1932). The forms with a constant apical length may represent a
certain stage in the phylogenetic development of diatoms. Evidently our
views on diatom variability and reproduction must be thoroughly revised.
Another interesting item was the formation of cell aggregates (“col-
onies”) of a typical appearance (“forma multiplex’) in the otherwise
solitary N. Kiitzingiana var. exilis Grun. (Botaniska Notiser 1940: 403.
1940). This form also has a specific tendency of passing to a skeleton-
free phase (Botaniska Notiser 1941: 33. 1941). The skeleton disappeared
after the normal form had been cultivated for several years, during which
time the decreasing average apical length had been followed down to a
501
502 FarLtowia, Voi. 1, 1944
certain minimum size which had been reached without auxospore forma-
tion. The cause of this might be that the strains in question have, during
cultivation, been obtained in clones which consequently lacked sexual
partners, in this case necessary for the formation of auxospores. This is
in contrast to the skeleton-free forms described by Richter, Bachrach,
Geitler, and Stosch which were probably produced through environmental
influence on the forms growing vegetatively.
The skeleton-free types, although not a part of the normal develop-
ment, are of a great interest since they may show resemblance to the phylo-
genetic ancestors of diatoms, i.e. unicellular brown-colored algae without
a silica skeleton. The common origin of the Bacillariophyta and Phaeo-
phyta is thus likely.
The morphologic results have as yet been presented only as preliminary
communications but the author hopes to be able to issue a more complete
and definite report of these parts of his investigations.
An interesting feature in the physiologic investigation was the demon-
stration of antagonism between the sulphanilamides and p-aminobenzoic
acid (Naturwiss. 29: 455. 1941; Science 94: 389. 1941; Botaniska Notiser
1941: 375. 1941). This is the first time that such an antagonism was
found to exist in other organisms than bacteria. According to Fildes’
theory (Lancet 238: 955, 1940), p-aminobenzoic acid is thus also an
essential metabolite for these autotrophic algae. In the experiments on
the antagonism between sulphanilamides and p-aminobenzoic acid, it ap-
peared that sulphanilamide inhibited growth more than sulphapyridine,
which was in its turn more growth inhibiting than sulphathiazole. In the
presence of p-aminobenzoic acid, however, conditions were the reverse.
‘From this it was interpreted that sulphapyridine and sulphathiazole also
influenced other biochemical processes than those connected with the
p-aminobenzoic acid (cf. Nature 150: 290. 1942). This opinion has
found support in observations of other experiments with other organisms
(West, Proc. Soc. Exp. Biol. Med. 46: 369. 1941; Dorfman & Koser,
Journ. Infect. Dis. 71: 241. 1942; Teply et al, Journ. Pharmacol.
77: 207. 1943).
Astra CENTRAL LABORATORIES
SODERTALJE, SWEDEN
oT, 5: eine Se
1(4) : 503-513 FARLOWIA July, 1944.
NEW AND NOTEWORTHY PHILIPPINE MOSSES
_Epwin B. BARTRAM
Since the “Mosses of the Philippines” was published in 1939, several
important and quite extensive collections have added a number of sig-
nificant species to the local moss flora which are here placed on record.
In the following list of twenty-six species, seven are apparently new to
science, twelve are species previously unrecorded from the Islands, and
self-explanatory remarks accompany the remaining seven.
The supplementary collections from which these records are derived
total well over 400 numbers. Only the more noteworthy collections are
listed here but the remainder is important from the standpoint of local
distribution. The collectors to whom we are indebted and the principal
sources of the material are as follows: Luzon and Mindoro — Mr. José
V. Santos in 1938; Mindanao, Palawan and Mindoro — Mr. Liborio E.
Ebalo, 1939-1940; Mindanao — Mr. A. Lynn Zwickey, Expedition of the
Arnold Arboretum, Harvard University, and the Botanical Garden of the
University of Michigan, 1938. I am further obligated to Mr. Santos for
his labors in packeting and labelling the specimens accumulated by the
University of Michigan so that they would be available for study. The
collections from Mindanao, Mindoro and Palawan are especially valuable
as much field work remains to be done in these areas, especially Mindanao,
and it may be long years before these prolific regions are again open to-
further exploration.
With these additions the Moss Flora of the Philippines is expanded to
931 species representing 184 genera. It is highly probable that the total
will exceed 600 species when the possibilities of the Archipelago are more
fully developed.
The types of the new species are in the writer’s herbarium and in the
herbarium of the University of Michigan. A complete set of the series
collected by Mr. Zwickey in Mindanao including the duplicate types will
also be found in the Farlow Herbarium, Harvard University.
FISSIDENTACEAE
Fissidens (Crispidium) Zwickeyi Bartr. sp. nov. Fic. 1-4,
Dioicus. Caulis 5 mm. altus. Folia circa 10 juga, sicca falcato-flexuosa, oblongo-
- ovata, obtusa, breviter apiculata, ad 1.6 mm. longa, 0.5 mm. lata, elimbata, lamina
dorsalis ad basin costae rotundata, margines ubique crenulati, cellulae hexagonae,
chlorophyllosae, laevissimae, diam. 10 yp, costa percurrens vel infra apicem soluta.
Fructus terminalis, seta 2-3 mm. alta, superne arcuata, theca inclinata, deoperculata
0.8 mm. longa, sicca sub ore contracta.
Mindanao: Lanao Prov., vicinity of Olangu near Momungan, alt. 400—
500 m., in thin layer of clay on moist rock, 25 Nov. 1938, A. L. Zwickey,
B38.
Near F, sylvaticus Griff. but I think clearly distinguished from this
species by the more broadly pointed leaves, the essentially smooth lamina
cells and the inclined or horizontal capsules.
503
504. FarRLowliA, VoL. 1, 1944
Fissidens lagunensis Bartr. nom nov.
Fissidens diversiretis Bartr. Philipp. Journ. Sci. 68 (1-4): 21. 1939.
I find that the name Fissidens diversiretis was used twice prior to its
publication in the “Mosses of the Philippines.” Brotherus describes I.
diversiretis from Yunnan Prov., China! followed by F. diversiretis Dixon
from Fiji? in 1930. Since F. diversiretis Broth. has priority, I propose
the above new name for the Philippine plants.
Fissidens Wichurae Broth. & Fleisch. Hedw. 38: (127). 1899.
Mindanao: Lanao Prov., vicinity of Dansalan, alt. 700-800 m., on clay
hummock, forest floor, Nov. 23, 1938, A. L. Zwickey, 794.
New to the Philippine Islands.
Distribution: Java.
DICRANACEAE
Angstroemia orientalis Mitt.
Luzon: Mountain Prov., Mt. Data, 26 March 1938, Josée V. Santos, 937a,
in part.
Previously known in the Philippines from only a single locality on
Mt. Pulog.
LEUCOBRYACEAE
Leucophanes Massarti Ren. & Card. Rev. Bryol. 23: 99. 1896.
Mindanao: Lanao Prov. Ulango Mountain, on bark of tree, 26 Feb. to
30 March 1941, Liborio E. Ebalo, 1176.
_New to the Philippine Islands.
Distribution: Java.
This collection, which matches almost exactly Fleischer’s Musc. Frond.
Arch. Ind. no. 205 from Java, is distinguished by the longer leaves
(7-9 mm.) in contrast to the short leaves (3-6 mm.) of L. octoblephari-
oides Brid.
CALYMPERACEAE
Syrrhopodon hispido-ciliatus Bartr. nom. nov.
Syrrhopodon perarmatus Bartr. Farlowia 1 (1): 42. 1943 non Syrrhopodon perar-
matus Broth. Journ. Proc. Roy. Soc. N. S. Wales 49: 133. 1915.
Mindanao: Zamboanga Prov., Muralong Mountain near Kabasalan, on
dead log, 27 Nov. 1940, Liborio FE. Ebalo, 737.
New to the Philippine Islands.
Distribution: New Guinea.
Mr. Dixon has kindly called to my attention that the name S. perarmatus
is preoccupied by a New Hebrides moss as indicated above, hence the
change.
It is an interesting coincidence that this species should turn up in
Mindanao so soon after its discovery in Papua, New Guinea. It will be
*Symbolae Sinicae, Part IV, p. 11, 1929.
* Proc. Linn. Soc. N. S. Wales 55 (3): 270. 1930.
BARTRAM: PHILIPPINE MossEs 505
readily separated from S. philippinensis Bartr. by the narrower, rigidly
erect-spreading leaves acute at the apex and the more densely hispid costa.
Calymperes porrectum Mitt. Linn. Soc. Journ. (Bot.) 10: 172. 1869.
Calymperes salakense Besch. Ann. Sci. Nat. ser. 8. 1: 302. 1895.
Mindanao: Lanao Prov., vicinity of Dansalan, alt. 700-800 m., on trunk
of tree, Oct. 6, 1938, A. Lynn Zwickey, 268.
Distribution: Malakka, Java, New Guinea, Samoa.’
A recent study of some Samoan collections indicates that C. porrectum
Mitt. and C. salakense Besch. are identical. As Mitten’s name has unques-
tioned priority I am reducing C. salakense to synonymy.
When the “Mosses of the Philippines” was published I had seen no
material of this species from the local area. The Mindanao collection
is a robust form showing the upper leaf margins coarsely and distantly
toothed, the teniolae well in at the leaf shoulders and the cancellinae
slightly rounded above, but these variations are of slight importance. In
all the material of this species that I have seen, the abnormal leaves are
mixed with the normal ones and reduced above to a bare costa which is
gemmiferous at the tip, suggestive of pseudopodia.
Calymperes strictifolium (Mitt.) Roth, Hedwigia 51: 127. 1911.
Syrrhopodon strictifolius Mitt. in Seemann FI. Vit. p. 388. 1873.
Syrrhopodon tuberculosus Dix & Ther. Journ. Linn. Soc. (Bot.) 43: 303. 1916.
Mindanao: Lanao Prov., vicinity of Davis Falls, alt. 200-300 m., moist
crevices and beneath “overhangs” of rocks, 29 Oct. 1938, A. Lynn
Lwickey, 552. |
Distribution: Borneo, Fiji, Samoa.
Comparison with a part of the type collection of Syrrhopodon stricti-
jolius Mitt. fails to reveal any practical distinctions between this species
and Calymperes tuberculosum (Dix. & Ther.) Broth. so there is no al-
ternative but to reduce the latter name to synonymy.
Calymperes (Eucalymperes) Ebaloi Bartr. sp. nov. Fic. 5-10.
Caespites densi, sordide virides, haud nitidi. Caules brevissimi, 4-8 mm. alti,
dense foliosi. Folia sicca et humida erecta, leniter flexuosa, haud contorta, ad 16
mm. longa, e basi breve ovata longe et anguste ligulata, limbata, sensim late acuta,
‘vix 0.25 mm. lata, lamina inferne sensim contracta, longe supra basin angustissima
vel nulla; marginibus ubique et remote denticulatis, ad basin dense spinoso-serratis;
costa valida, percurrente, dorso ad apicem solum denticulata; cellulae superiores
laevissimae, subquadratae, diam. 10-12 y.; cancellina bene definita, supra acutata,
teniola ad basin indistincta, in lamina validiuscula, fere ad apicem attingens, intro-
marginata. Seta tenuis 12 mm. longa, laevis; theca cylindrica, deoperculata 1.5 mm.
longa. Caetera ignota.
Mindanao: Zamboanga Prov., Lilimbrog Mountain near Kabasalan, on
tree trunk, 24 Nov. 1940, Liborio Ela Ebalo, 664.
Widely distinct from C. serratum A. Br. in the petiole-like base, the
intramarginal leaf border and the straight or only slightly flexuose leaves.
I have an undescribed species from Fiji with similar leaves in which the
oy hoe ig spall aii CR al ac
306 FarLowia, VOL. i 1944,
lamina is gradually contracted below to a stretch of bare costa, then wid-
ened abruptly again at the shoulders of the leaf base, but in the Fiji plants
the leaves are shorter without an intramarginal border, the lamina cells
papillose and the leaf shoulders not spinose-serrate.
MNIACEAE
Orthomniopsis japonica Broth. Ofv. Finska Vet.-Soc. Forh. 49 (10): 1. 1905-06.
Luzon: Bontoc Subprov., June-July 1911, Father M. Vanoverbergh,
1282.
Distribution: Assam, New Guinea, Japan.
Through the kindness of Dr. Quisumbing I have seen part of the above
collection which is unquestionably this species. For some reason it was
not included in the material sent from the Bureau of Science as a basis
for the study of the local moss flora.
BARTRAMIACEAE
Philonotis evanidinervis Fleisch. Laubmoosfl. Java 2: 412. 1902-04.
Mindanao: Lanao Prov., vicinity of Dansalan, alt. 700-800 m., on rock,
edge of stream, secondary forest toward Remain, 28 October 1938, A. Lynn
Zwickey, 539. Ss
New to the Philippine Islands.
Distribution: Java.
An interesting extension of this localized species previously known only
from several localities in West Java.
RHACOPILACEAE
Rhacopilum magnirete Bartr. sp. nov. Fic. 11-14,
Sat robustum, caespites late extensi, lutescentes, haud nitidi. Caules intertexti,
fere ad apicem dense rufo-tomentosi, irregulariter ramosi. Folia late ovata, acuta,
breviter cuspidata, 2 mm. longa, 1 mm. lata. sicca fortiter inflexa, humida late patentia;
margines plani, supra medium irregulariter denticulati; costa tenuis, superne valde
angustiora, sensim in cuspidem attenuata, saepe infra apicem dissoluta; cellulae
superiores majusculae, pellucidae, laeves, hexagonae, 22-25 y. longae, parietibus
firmis, basin versus paullo majores, internae longiores. Amphigastria multo minora,
ovato-lanceolata, sensim acuminata, costa longe excurrente. Caetera ignota.
Mindanao: Lanao Prov., Saguiran Mountain, on tree trunk, 26 Feb. to
30 March 1941, Liborio E. Ebalo, 1124.
This species is possibly nearest R. Schmidi (C. M.) Jaeg. but the
densely tomentose stems, more coarsely toothed leaves, the slender costa
ending in the shorter point or even disappearing below the apex, together
with the larger pellucid and distinct upper leaf cells are distinctive fea-
tures. From R. spectabile Reinw. & Hsch. it will be separated by the less
robust habit, larger lamina cells, less coarsely toothed leaf margins and
the absence of the area of lax juxtacostal cells in the leaf base.
THUIDIACEAE
Herpetineurum toccoae (Sull. & Lesq.) Card. Beihefte Bot. Centralb. 19 (2):
127. 1905.
Anomodon toccoae Sull. & Lesq. Musc. Bor, Am. (Ed. 1) no. 240, 1856.
a
BARTRAM: PHILIPPINE MosseEs Serre | OE:
Luzon: Malinao, Atimanan, Tayabas Prov., on ground, 25 Oct. 1927,
José V. Santos, 862.
New to Philippine Islands.
Distribution: Africa, Asia, Ceylon, Sumatra, Java, Celebes, New Cale-
donia, Formosa, South America, Guatemala, Mexico, Southern United
States,
It is not surprising to find this almost cosmopolitan species in the
Philippines. Although not common anywhere it seems to occur spe-
radically in temperate and tropical regions throughout the world.
PTEROBRYACEAE
Endotrichella nematosa Bartr. sp. nov. FIG. 15-17.
Robusta,. flavescens, nitida. Caules secundarii ad 5 cm. alti, complanati, densi-
folii, circa 8 mm. lati, superne saepe attenuati et prope apicem propagula numerosa
aurantiaca filiformia articulata instructi. Folia 5 mm. longa, 1.5 mm. lata, oblongo-
lanceolata, ecostata, plicata, parum auriculata, in acumen cuspidatum raptim an-
gustata; margines infra recurvi, superne distanter fortiter serrati, cellulae superiores
anguste rhomboideae, perincrassatae, basilares lineares, infimae fuscae, fortiter
porosae, alares oblongae, laxae, numerosae. Perichaetium parvum; theca exserta,
leniter inclinata, deoperculata breviter oblonga, 1.8 mm. longa, 1 mm. lata; seta circa
2 mm. longa. Caetera ignota.
Panay: Mt. Bulilao, Capiz Prov., June 1919, A. Martelino & G. Edano,
35812a. Caroline Islands: Ponape Island, Kuporujo, grows on top of
the mountain on epiphytic orchid, 13 March 1936, Masahiko Takamatsu,
Herb. B. P. Bishop Museum, 66la (TYPE).
An outstanding species in the caudate tips of the secondary stems which
are abundantly felted with erect, filiform, orange-brown, articulated brood
filaments as in some of the species of Cyathophorella.
Garovaglia Zwickeyi Bart. sp. nov. FIG. 18-20.
Rigida, aureo-viridis, nitidula. Caules circa 4 cm. alti, sat gracili. Folia dense
conferta, erecta, arcte appressa, late orbiculato-ovata, haud undulata, fortiter plicata,
breviter cuspidata, 2.5 mm. longa, 2 mm. lata; margines plani, leniter denticulati;
costae nullae; cellulae breviter rhomboideae, parietibus firmis, alares rectangulares,
numerosae. Perichaetium Jonge exsertum, folia perichaetialia abrupte breviter
cuspidata, superne erosa, theca immersa. Caetera ignota.
Mindanao: Lanao Prov., vicinity of Olangu near Momungan, alt. 400—
500 m., on trunk of tree, 10 November 1938, A. Lynn Zwickey, 700.
A very distinct species in the short, broad, closely appressed leaves
giving the rather slender, subterete stems a catenulate appearance due to
the reflexed upper leaf margins. It may be compared in a broad way
with G. rigida Dix. of Siam but is widely different in the much shorter and
relatively broader erect appressed leaves with the margins weakly
denticulate. 3
NECKERACEAE :
Neckeropsis nitidula (Mitt.) Fleisch. Laubmoosfl. Java 3: 882. 1906-08.
Luzon: Tayabas Prov., Danlagan, Guinayagan, on tree branch, 19 Oct.
1936, José V. Santos, 482.
vi fe neh
508 Fartowia, VoL. 1, 1944
New to Philippine Islands.
Distribution: China, Japan, Formosa, Tonkin.
Notwithstanding my remarks on page 245 of the “Mosses of the Philip-
pines” I am happy to definitely record the presence of this species in the
local area.
Pinnatella ambigua (Bryol. Jav.) Fleisch.
Mindoro: Pandan Mountain, Bulalacao, on stone, 6-11 Nov. 1939,
Liborio E. Ebalo, 239.
Distribution: Bhota, Burma, Sumatra, Java.
Although the plates in the “Bryologia Javanica” representing P. ambigua
and P. Kuhliana respectively show clearly the distinctive difference in
the shape of the stem leaves, unfortunately I failed to grasp this distinc-
tion when studying the Philippine collections and as a result confused the
two species. P. ambigua is at once distinguished by the stem leaves grad-
ually narrowed from an ovate base to a relatively slender acuminate point
in bold contrast to the oblong-ovate abruptly short-pointed leaves of
P. Kuhliana.
P. ambigua is apparently rare in the Philippines. Fleischer cites a col-
lection from Luzon in the Laubmoosfl. Java, p. 919 but the only speci-
men I have seen is the one cited above.
Pinnatella Kuhliana (Lac.) Fleisch. Hedwigia 45: 80. 1906.
New to the Philippine Islands.
Distribution: Sumatra, Java, Ceram, Celebes, New Guinea, Fiji, Samoa,
Tahiti.
As explained above the description, figures and collections given under
the heading of P. ambigua in the “Mosses of the Philippines” should all
be referred to Pinnatella Kuhliana. This species does not seem to have
been recorded from the Philippines but a careful check of all the avail-
able specimens shows that it is widely distributed throughout the archi-
pelago and decidedly more frequent than P. ambigua.
Pinnatella mucronata (Lac.) Fleisch. Hedwigia 45: 50. 1906.
Palawan: Tagbarungis Mt., vicinity of Puerto Princesa, on tree trunk,
14 February 1940, Liborio E. Ebalo, 489.
New to the Philippine Islands.
Distribution: Sumatra, Java, Borneo, Ceram, Celebes, New Guinea,
Samoa.
HOOKERIACEAE
Daltonia armata Bartr. sp. nov. FIG. 21-24,
Pallide viridis, nitidula. Caulis ad 1 cm, altus. Folia 3.5 mm. longa, 1 mm.
lata, oblongo-ovata, longe piliformiter aristata, erecto-patentia, sicca leniter contorta,
ubique angustissime limbata, integerrima; costa tenuissima, vix dimidiam partem folii
attingens; cellulae superiores elongate hexagonae, 40-60 x 10-16 u, parietibus
tenuiusculis, basilares lineares. Seta 3 mm. longa, fere ad basin argute tuberculosa;
theca erecta, deoperculata 1.2 mm. longa; calyptra fimbriata.
BarTRAM: PHILIPPINE MossEs : 509
Mindanao: Lanao Prov., vicinity of Dansalan, alt. 700-800 m., on culm
of climbing bamboo, Sacred Mountain, 3 Nov. 1938, A. Lynn Zwickey, 638.
The leaves of D. armata are similar in shape to those of D. aristifolia
Ren. & Card. of Java but the resemblance ends here. In D. armata the
upper lamina cells are oval-hexagonal up to 60 » long and the setae are
densely armed throughout with high tubercular-like spines up to 25 » high.
PLAGIOTHECIACEAE
Plagiothecium neckeroideum Br. & Schp. var. javense Fleisch. Laubmoosfl.
Java 4: 1168. 1920.
Luzon: Mountain Prov., Mt. Pawai, Benguet, on ground, 27 March 1938, |
José V. Santos, 1035.
Distribution: Sikkim, Java.
In this collection the setae are 3-4 cm. long, thus agreeing enh the
description of Fleischer’s var. javense. In the absence of any other well
marked differences it seems probable that the variety is hardly more than
a robust. form of the species.
SEMATOPHYLLACEAE
Heterophyllum Santosii Bartr. sp. nov. FIG. 25-28.
Autoicum, pallide aureum, nitidum, Caulis prostratus, elongatus, regulariter pin-
natim ramosus, ad 9 cm. longus, ramis vix 1 cm. longis, leniter flexuosis, apice fal-
cato. Folia caulina falcata, 2.5-3 mm. longa, e basi late oblonga raptim lineari-
lanceolata, superne spinoso-serrata; marginibus inferne integris, anguste revolutis;
costa bina, brevissima; cellulae superiores lineares, parietibus, hyalinis, firmis,
basilares longiores, angustiores, infimae laxiores, fuscae, valde porosae, alares num-
erosae, oblongae, hyalinae vel fuscae, haud vesiculosae. Folia ramea minora, breviora,
angustiora, sensim acuminata, argute serrata, apicem versus saepe rugoso-undulata.
Seta 37 mm. longa, rubella, superne tenuissima; theca suberecta, deoperculata 2.5 mm.
longa. Caetera ignota.
Luzon: Mountain Prov., Mt. Data, Benguet, on tree trunk, 26 March |
1938. José V. Santos, 957.
A splendid species, possibly allied to H. brachycarpum (Mitt.) Fleisch.
of Japan but widely different in the elongated stems regularly and closely
pinnate forming narrow fronds when moist. The unusually long seta
shown by the only available sporophyte is also a distinctive feature. The
genus adds a new and interesting element to the local flora from a region
which has been carefully worked over by many previous collectors, so
Mr. Santos may well be proud of his discovery.
Glossadelphus similans (Bryol. Jav.) Fleisch. E. & P. Nat. Pflanzenfam, Musci
p. 1093. 1908.
Luzon: Mountain Prov., Mt. Pawai, Benguet, on dead log, 27 March
1938, José V. Santos, 1027.
New to Philippine Islands.
Distribution: Java.
510 FarLowlA, VoL. 1, 1944
Glossadelphus hermaphroditus Fleisch. Laubmoosfl. Java 4: 1359. 1920.
Mindanao: Lanao Prov., vicinity of Guassi-Dansalan Road, alt. 700-
800 m., on boulder in stream bed near Togaya, 2 Nov. 1938, A. Lynn
Zwickey, 618.
New to Philippine Islands.
Distribution: Java.
Pseudohypnella verrucosa (Doz. & Molk.) Fleisch. Laubmoosfl. Java 4: 1368.
1921,
Mindanao: Lanao Prov., Mt. Makaturing, alt. 800-1000 m., on rotting
wood, 22 Oct. 1938, A. Lynn Zwickey, 489.
New to Philippine Islands.
Distribution: Ceylon, Java, Banka.
POLYTRICHACEAE
Pogonatum macrophylloides Broth. Mitt. Inst. allgem. Bot. 7 (2): 140. 1928.
Luzon: Mountain Prov., Mt. Pawai, Benguet, on ground, 27 March 1938,
José V. Santos, 1026.
New to Philippine Islands.
Distribution: Borneo.
BusHKILL, Pike Co.
PENNSYLVANIA
ah
5.
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512, FarLtowlA, Vo. 1, 1944
EXPLANATION OF PLATE
Fig. 1-4. Fissidens Zwickeyi Bartr.
1. Plants « 1.
2. Leaf x 20,
3. Apex of leaf « 200.
4. Capsule x 7.
Fig. 5-10. Calymperes Ebaloi Bartr.
2: Plant x 1,
6. Leaf x 4.
7. Apex of leaf x 50.
8. Leaf base x 20.
9, Upper cells and margin x 200.
10. Part of cross section from upper half of leaf x 200.
Fig. 11-14. Rhacopilum magnirete Bartr.
ll. Part of plant x 1.
12. Leaf x 15.
13. Apex of leaf x 40.
14. Upper leaf cells and margin x 200.
Fig. 15-17. Endotrichella nematosa Bartr.
15. Plant x 1.
16. Part of propagulum x 200.
17, Leaf < 7. ‘
Fig. 18-20. Garovaglia Zwickeyi Bartr.
18. Plant x 1.
19. Leaf « 16.
20. Upper leaf cells and margin x 200.
Fig. 21-24. Daltonia armata Bartr.
2): Plant < 7,
22. Capsule x 9,
23, Leaf x 10.
24. Upper leaf cells and margin x 200.
Fig. 25-28. Heterophyllum Santosii Bartr.
25. Plant x \%.
26. Stem leaf « 13.
27. Branch leaf « 13.
: 28. Basal angle of leaf x 80.
Figures 1-28
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1(4) : 515-523 FARLOWIA July, 1944
ON TWO NEW SPECIES OF MEGACEROS WITH NOTES ON
M. ARACHNOIDEUS, M. DENTICULATUS, M. GIGANTEUS,
AND M. GRANDIS
L. P. Kanna
For the material which forms the basis of this note, the writer is in-
debted to Mrs. E. A. Hodgson, Wairoa, P. O. Hawkes Bay, New Zealand,
to whom he wishes to express his thanks. Thanks are also due to Dr.
Eduardo Quisumbing, Curator, Philippine National Herbarium, Manila,
-and to Dr. R. E. Holttum, Director, Botanical Gardens, Singapore, for
_ the loan of specimens from their herbaria.
Megaceros Hodgsoniae sp. nov. ; Fics. 1-6.
Planta monoica. Sectio transversa 5-6 cellis excelsitato, solida. Cellae super-
ficiales 30-70 X 25-45 y.. Involucre geminatum, 5-6 mm. longitudine, 0.5-1.5 mm.
latitudine, cylindricum, attenuatum. Capsula 8-15 mm. longitudine, 0.25-0.35 mm.
latitudine. Sporae pallidae, verrucolosae, 30-40 Une
Monoecious. In patches. Thallus up to 25 mm. long, nearly flat,
smooth, divided into broad obovate lobes, more or less rounded at the
apex with margin nearly entire; transverse section 5-6 cells high in the
middle, without lacunae; surface cells 30-70 x 25-45 p. Involucre fre-
quently geminate, 5-6 mm. long and 0.5-1.5 mm. broad, cylindrical,
slightly narrowing at truncate apex. Capsule 8-15 mm. long and
0.25—0.35 mm. broad, wall cells thickened, light brown. Spores 30-40 p,
pale, verrucose. On Popa Rock in creek in deep shade, Kiwi, April, 1935.
From the comparative chart of the species of Megaceros Campbell 1907
appended, the following species agree with the form described in having
the verrucose type of spore surface: M. arachnoideus Stephani 1916,
M. denticulatus (Lehman 1857) Stephani 1916, M. fuegiensis Stephani
1923, M. martinicensis Stephani 1916. The present form differs from
M. martinicensis by the shorter capsule and the larger thallus, from
M. arachnoideus, M. denticulatus and M. fuegiensis by the shorter capsule
and the structure of the thallus. It is therefore necessary to create a new
species for which the name M. Hodgsoniae is suggested.
Megaceros Zotovii sp. nov. Fics. 7-10.
Planta monoica. Sectio transversa 10-12 cellis excelsitato, solida; cellae super-
ficiales 30-45 x 15-25 y. Involucre solitarium, cylindricum, attenuatum, 12-16 mm.
longitudine, 1.5-2 mm. latitudine. Capsula 40-60 mm. longa, 0.5 mm. lata. Sporae
pallidae, verruculosae, 30-35 wu.
Monoecious. In patches,:light green. Thallus up to 50 mm. in di-
ameter, fan-shaped, with obovate lobes, rounded at the apex with margin
finely crenate, transverse section 10-12 cells high in the middle, without
lacunae; dorsal epidermal cells 30-45 x 15-25 p. Involucre 12-16 mm.
long and 1.5-2 mm. broad, cylindrical, narrowing towards the nearly
truncate apex. Capsule 40-60 mm. long and 0.5 mm. broad. Spores
30-35 y, pale, somewhat irregular in shape, verrucose. Elators un-
branched, 0.2-0.3 mm. long. Epidermal cells of the capsule somewhat
515
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516 FarLtowl1a, Vou. 1, 1944
thickened. Cehatorawa Saddle, Tarama Mts., New Zealand, 26 Nov.
1933, V. D. Zotov. :
The above described species agrees with the following in having ver-
rucose spores: M. arachnoideus Stephani 1916, M. denticulatus (Lehmann
1857) Stephani 1916, M. fuegiensis Stephani 1923, M. Hodgsoniae Khanna
and M. martinicensis Stephani 1916. The present form differs from M.
denticulatus and M. Hodgsoniae by its longer involucre and capsule, from
M. arachnoideus, M. fuegiensis and M. martinicensis by the bigger in-
volucre and other characters of the thallus.
Megaceros giganteus (Lehmann et Lindenberg) Stephani Fics. 11-15.
Monoecious. In light green patches. Thallus up to 50 mm. in di-
ameter, divided into obovate lobes, depressed in the center, margin slightly
ascending-irregular; transverse section 8-10 cells high in the middle,
without lacunae; surface cells 8-20 x 20-30 ». Involucre 8-15 mm. long
and 1.0-1.5 mm. broad, cylindrical, slightly narrowed towards the apex.
the mouth lobulate. Capsule 40-60 mm. long and 0.4—0.8 mm. broad,
epidermal cells of the capsule, long, narrow, highly thickened, and some
cells with brownish contents. Spores 30-35 p, pale, asperulate. Elators
long and slender, sometimes 0.5 mm. in length, unbranched. Antheridia
singly on prominent stalks. Water course, north of Field Flat, 8 Oct.
1933, V. D. Zotov.
The above described species agrees with the following in having the
same type of spore surface: M. caledonicus Stephani 1916, M. celebensis
Stephani 1916, M. crassus Stephani 1916, M. endiviaefolius (Montagne
1856) Stephani 1916, M. giganteus (Lehmann et Lindenberg 1834)
Stephani 1916, M. Jamesonii (Taylor 1848) Stephani 1916, M. lacerus
(Nees 1844.) Stephani 1916, M. longispirus (Carrington & Pearson 1887)
Stephani 1916, M. monandrus Stephani 1916, M. Novae-guineae Stephani
1916, M. Novae-zelandiae Stephani 1916, M. pallens (Stephani 1892)
Stephani 1916, M. parvisporus Stephani 1916, M. solidus Stephani 19106.
The present species differs from M. caledonicus, M. celebensis, M.
crassus, M. lacerus, M. monandrus, M. Novae-guineae, M. pallens, and
M. solidus by the larger thallus and involucre; from M. endiviaefolius
by its shorter capsule; from M. Jamesonii by its smaller spore and other
vegetative characters; from M. lacerus by the shape and the other char-
acters of the thallus. M. giganteus, M. Novae-zelandiae and M. parvi-
sporus and the above described species differ from each other in minor
characters which are not sufficiently distinct to be of specific value. The
writer therefore is of the opinion that these may be considered as a single
species — M. giganteus (Lehman et Lindenberg 1834) Stephani 1916.
_ The writer has been unable to consult Stephani’s material of these forms
and is therefore unable to state whether differences exist which would
supplement the already existing description. Judging from the char-
acters so far given, however, he is of the opinion that there is only one
species, and the other two listed are environment forms without spe-
cific validity.
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KHANNA: MEGACEROS Syke
Megaceros grandis (Angstrom) Stephani FIcs. 16-20.
Monoecious. On rotten log, mixed with mosses. In light green patches.
Thallus 12-20 mm. in diameter, fan-shaped or nearly so; slightly de-
pressed in the center, the margin ascending; divided into numerous nar-
row linear lobes, dorsal surface rough with several fine lamellae; surface
cells 35-55 x 20-40 yp, transverse section 5 cells high in the middle, with-
out lacunae. Involucre 4.5-6.5 mm. long and 0.8-1.0 mm. broad, cylin-
drical, narrowing to the almost truncate apex. Capsule 12-40 mm. long
- and 0.3-0.5 mm. broad, epidermal cells long, thick-walled. Spores
25-35 p, pale green, granular; elators unbranched, 0.18—-0.2 mm. long.
Antheridia borne singly in the antheridial cavity. Stream side—on a
log in a bush about 3000 ft. above sea level; on Tauhara, New Zealand.
From the comparative chart of Megaceros Campbell, 1907, the following
species agree with that described above in having the same type of spore
surface: M. grandis (Angstrom 1873) Stephani 1916, M. vescoanus
Stephani 1916, M. vincentianus (Lehmann et Lindenberg 1834) Ste-
phani 1916.
The present species can be separated from M. vincentianus by its
shorter capsule and other structures of the thallus; M. grandis, M.
vescoanus and the above differ from each other in minor characters which
are not sufficiently distinct to be of specific value. The writer, therefore,
is of the opinion that these may be considered as a single species —
M. grandis (Angstrom 1873) Stephani 1916.
Megaceros denticulatus (Lehmann) Stephani Fics. 21-25.
Monoecious. In overlapping patches. Thallus variable, up to 55 mm.
long and 40 mm. broad, with irregular branches, rounded at the apex,
with margin irregular; from the ventral surface marginal buds are given
off; the upper superficial cells 25-30 x 10-20 ». Involucre 5-7 mm.
long and 0.8 mm. broad, narrowing towards the apex, the mouth lobulate.
Capsule 16-20 mm. long and 0.5 mm. broad, brownish. Spores 35-40 p,
pale yellow, verrucose. Capsule wall cells highly thickened.
From the description this plant appears to be M. denticulatus (Leh-
mann 1857) Stephani 1916— though the present form differs by the
larger involucre and the smaller spores — but these differences within a
narrow limit are not sufficiently distinct to be of specific value.
Megaceros arachnoideus Stephani FI¢s. 26-29.
Monoecious. In patches. Thallus 15 mm. in diameter, divided into
broad obovate lobes, with margin lacerate; dorsal surface rough with
several leaf-like lamellae; transverse section 10-14 cells high in the
middle, without lacunae; upper superficial cells 35-45x 20-45 p. — In-
volucre 6-10 mm. long and 0.6-1.2 mm. broad, narrowing towards the
apex, mouth lobulate. Capsule 25-35 mm. long and 0.3—0.4 mm. broad,
brownish. Spores 25-40 », somewhat irregular in shape, greenish yellow,
verrucose. Elators long and slender, sometimes 0.2 mm. in length. On
damp earth by creek by [Mols St.?] New Zealand, 26 Dec. 1932,
kK. A. Hodgson.
518 FarLtowl14, VoL. 1, 1944
This form agrees with M. arachnoideus Stephani 1916 but shows minor
differences.
University OF RANGOON
InpIA
REFERENCES
Campbell, D. H. Studies on some Javanese Anthocerotaceae, I. Ann, Bot. 21:
467-486. pl. 44-46. 1907.
Stephani, F. Species Hepaticarum 5: 945-957. 1916.
—_————. Species Hepaticarum 6: 424. 1923.
alatifrons Steph.
amoenus Steph,
aneuraeformis Steph.
arachnoideus Steph.
caledonicus Steph.
callistictus Steph.
carnosus Steph,
celebensis Steph.
columbianus Steph.
crassus Steph.
cristisporus Steph.
denticulatus (Lehm.) Steph.
endiviaefolius (Mont.) Steph.
flagellaris (Mitt.) Steph.
flavens (Spruce)
Suegiensis Steph.
giganteus (Lehm. & Lindenb.)
Steph.
gracilis (Reichardt) Steph.
grandis (Angstr.) Steph.
guatemalensis Steph.
jamaicensis Steph.
Jamesonii (Tayl.) Steph.
lacerus (Nees) Steph.
laciniatus (Schw.) Steph.
leptohymenius (Tayl.) Steph.
longispirus (Carr. & Pears.)
Steph.
martinicensis Steph.
mexicanus Steph.
minarum (Nees) Steph.
monandrus Steph.
monospirus Steph.
muriculatus Steph.
Novae-guineae Steph.
Novae-zelandiae Steph.
Nymanii Steph.
pallens (Steph.) Steph.
parvisporus Steph. e
salakensis Campb.
schizophyllus (Gottsche)
Steph.
solidus Steph.
Stahlii Steph.
tjibodensis Campb.
tosanus Steph. . ’
Vescoanus Steph.
vincentianus (Lehm. & Lin-
denb.) Steph.
Wiemanii Steph.
KHANNA: MEGACEROS
APPENDIX
SPECIES OF MEGACEROS
Size of Invo- Cap-
plant lucre_ sule
mm. mm. mm.
20x3 7 30
20x3 7 30
40x 5 5 30
40x25 7 30
20x3 5 40
20 x3 3 15
30 5 50
20 x 12 8 30
15x5 12 40
20 6 40
40 15 70
50 2 20
60 x 7 7 90
40x 5 a 40
30x 5 8 60
30x 8 5 40
45 x7 10 40
20 5 40
30x10 10 30
40x 5 9 60
50x10 10 40
40x10 10 50
40 7 40
40 5 40
50x10 10 40
50 x 20 5 50
10x 4 4 40
15x4 4 40
20 5 40
30 5 20
35 7 25
30 5 45
20x3 5 40
40 5 30
30 4 35
25x5 5 40
30x 5 10 40
30 6 40
15 5 20
20x 4 6 20
30 15 80
50x15 10 90
30 i 20
40 x7 7 40
40 x 6 5 50
20x10 15 50
Spore
27 uw, small papillae in
center
27 pn, umbonate
36 », umbonate
36 uw, yellow, verrucose
27 uw, Minutely asper
22 uw, muricate -
27 uw, yellow, muricate
27 uw, minutely asper
36 uw, umbonate
36 uw, asper
27 uw, yellow, papillate
45 yu, yellow, verrucose
36 yu, asper
18 nz, muricate
27 w, muricate
36 yw, verrucose
27 uw, asper
36 u, papillate
27 uw, granulate
27 uw, papillate
27 u, papillate
54 yw, asper
36 yu, asper
— u, muricate
36 yu, papillate
36 », minutely asper
‘27 uw, dark, verrucose
27 yp, papillate—
27 w, yellow, papillate
27 », minutely asper
27 yw, papillate
27 uw, Muricate
27 yw, asper
27 w, asper
36 yu, papillate
27 uw, asper
18 uw, asper F
28 uw, minutely echinate
27 uw, tuberculate !
36 uw, asper
27 wu, yellow, densely
papillate
27 wu, papillate
27 uw, muriculate
18 yn, granular
27 w, granular
27 wu, umbonate
219
Distribution
Brazil
Guadeloupe
Brazil
New Zealand
New Caledonia
Brazil
Australia
Celebes
Colombia
Tasmania
Costa Rica
New Zealand
Chile
Fiji
Peru
Tierra del Fuego
New Zealand
Australia
Tahiti
Guatemala
Jamaica
Peru
Africa
Mexico
New Zealand
Tasmania
Martinique
Mexico
Brazil
Asia ,
New Caledonia
New Caledonia
New Guinea
New Zealand
New Guinea
New Zealand
Samoa
Java
Cuba
Guadeloupe
Java
Java, Sumatra
Japan
Tahiti
Guadeloupe, St.
Vincent
Brazil
520 FarLtow1a, Vou. 1, 1944.
EXPLANATION OF FIGURES 1-15
In figures 1 and 11, the scale equals 1 millimeter; in all other figures the scale
equals 20 yu.
Megaceros Hodgsoniae. Figs. 1-6.
1. Young plant showing sporophyte.
2. Dorsal epidermal cells.
3-4, Spores.
5. A portion of an elator.
6. Capsule wall cells.
Megaceros Zotovii. Figs. 7-10.
7. Dorsal epidermal cells.
8. Different views of spores.
9. Part of elator.
10. Capsule wall cells.
Megaceros giganteus. Figs. 11-15.
11. Thallus showing sporophyte.
12. Dorsal epidermal cells.
13. Spore.
14, Elator.
15. Capsule wall cells.
Se eg ee ae
Se ee Na
“Scale
521
KHANNA: MEGACEROS
)
Figures 1-1
eae se Le ne ea eer ee Cae ee ee ee ee ee x ee 7, = * ih sap ee we Of a. See OG wll) Se Te, a
522 Far.towlia, VoL. 1, 1944
EXPLANATION OF FIGURES 16-29
In figures 16, 21, and 26, the scale equals 1 millimeter; in all other figures the
scale equals 20 u.
Megaceros grandis. Figs. 16-20.
16. A young plant.
17. Dorsal epidermal cells of the thallus.
18. Spore.
19. A portion of an elator.
20. Epidermal cells of the capsule wall.
Megaceros denticulatus. Figs. 21-25.
21. A portion of the lobe of the thallus showing the irregular margin.
22, Epidermal cells of the thallus.
23. Capsule cell walls.
24. Spore.
25. Portion of an elator.
Megaceros arachnoideus. Figs. 26-29.
26. Young plant.
27. Dorsal epidermal cells of the thallus.
28. Spores.
29. A portion of an elator.
KHANNA: MEGACEROS
Ficures 16-29
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: ere ' Ss ene
1(4): 525-568 FARLOWIA July, 1944
STUDIES ON POLYSTICTUS CIRCINATUS AND ITS RELATION
TO BUTT-ROT OF SPRUCE
Rocer GossE.Lin!
During the writer’s study of the butt-rots of spruces in the Province of
(Quebec, it was found that the most important fungi causing butt-rots are
Porta subacida, Polyporus balsameus, Polyporus Schweinitzii, and Poly-
porus circinatus. This last was chosen for study because of the irregular
occurrence of the species and of its importance locally in causing ex-
cessive windfall.
It soon became evident that what had been determined as P. circinatus
actually included two distinct fungi. One was characterized by its bright-
er color and by the fact that it occurred chiefly on the ground; the other
was characterized by the darker color of the sporophores and by the fact
that it occurred mostly on the trunks of the trees. These differences be-
tween the two forms which were so obvious in the field, led to the inves-
tigations of the taxonomy. It was found that the two organisms could
be separated microscopically by the character of the setae present in the
hymenium and this led to their determination as Polyporus tomentosus
Fr. and Polyporus circinatus Fr.
From a review of the literature it appears that there has been consid-
erable confusion as to the identity of these two forms. This confusion
has arisen since Fries described the two species, P. tomentosus in 1821 and
P. circinatus in 1848, and it came from the fact that he described P. tomen-
tosus as having a homogeneous context, yet according to specimens iden-
tified by Fries, both P. tomentosus and P. circinatus have a duplex context.
The only difference, excepting the color and the shape, are the straight
setae for the former and the predominantly curved setae for the latter.
Peck (1878) described a new species from New York, which he called
Polyporus dualis. His species was sessile or short, lateral-stemmed and
of rather dark color. Later, Ellis and Everhart (1889) examined speci-
mens from Peck and found the curved setae characteristic of P. circinatus
Fr. Cooke (1886) and Saccardo (1888) have given P. dualis Pk. as a
synonym of Polystictus circinatus (Fr.) Cke. So it is fairly safe to say
that Peck’s type specimen is P. circinatus Fr.
In 1882, Karsten transferred Polyporus tomentosus to Polystictus. Later
(1889) he segregated the new genus Onnia from Polystictus, placing em-
phasis upon the setal character of the hymenium. He distinguished
O. tomentosus from QO. circinatus by the homogeneous context of the
former. Ellis and Everhart (1889) examined Karsten’s specimen later
and found straight setae for the plant he designated as Onnia tomentosus.
Although Ellis and Everhart’s illustration (1889) did not make this point
1 Thesis submitted in partial fulfillment of the requirements for the degree of Doctor
of Philosophy at Harvard University.
Contribution from the Laboratories of Cryptogamic Botany and the Farlow Her-
barium, Harvard University, no. 225.
529
526 FartowlA, Vou. 1, 1944
clear, Karsten’s specimens show that there is a duplex context, hence his
Onnia tomentosus is the same as P. tomentosus Fr. Little is known about
his O. circinatus.
While Karsten was creating the genus Onnia, Ellis and Everhart (1889)
created the genus Mucronoporus based on Polyporus circinatus. They
described straight setae for M. tomentosus and curved ones for M. cir-
cinatus. Subsequent mycologists have considered these species synony-
mous with the two Friesian ones. |
In 1900, Patouillard made a new genus Xanthochrous based on P. cir-
cinatus. He described the setae as being straight. Apparently his X.
circinatus is P. tomentosus Fr.
Murrill (1904) refers to Fries’ authentic specimens, stating that “the
type of Polyporus tomentosus at Upsala corresponds in all respects with
the plants found in America, having the same kind of spines and a dual
context.” Since Nannfeldt has reported in a personal letter to Haddow
(1941) that there are no type specimens of either of Fries’ species at
Upsala, it is difficult to understand to which specimen Murrill refers. It
is worth observing, however, that he attributed to his Friesian specimen
of P. tomentosus a double context. Murrill (1916) comprises under the
single species Coltricia tomentosa both P. tomentosus and P. circinatus
of Fries.
There apparently was much confusion in Lloyd’s mind in regard to
the status of these two species. In 1908, he described P. circinatus Fr.
as having straight setae, as in P. tomentosus Fr. Later in 1912, he stated
that P. tomentosus Fr. was unknown from the United States and ques-
tioned whether P. circinatus Fr. was really distinct from P. tomentosus
Fr., and finally suggested that “there are three (stipitate) forms of Poly-
porus tomentosus Fr.: first (typical), with thin context, straight setae;
second (circinatus, American), thick context, straight setae; third (cir-
cinatus, European and American), curved setae.”
Lloyd (1908), on account of the resemblance of P. dualis Pk. to Fries’
figure of P. triqueter, suspected it to be P. triqueter of Europe. Bresadola,
(1920) following Lloyd’s opinion, considered P. dualis Pk. to be a
synonym of P. triqueter Fr. Fries described P. triqueter in 1838, while
P. circinatus was described only in 1848. He did not mention any rela-
tionship between the two species, hence P. triqueter Fr., if different from
P. dualis Pk. as seems likely, cannot be used as a synonym for P. circinatus
and there is little need to consider it in this discussion. |
Neuman (1914) reported P. tomentosus from Wisconsin but gave no
description of the microscopic characters. He considered P. dualis as a
synonym of P. tomentosus and for that reason the exactitude of his iden-
tification may be questioned.
Lloyd (1920) described a new species, Polyporus peakensis Lloyd, from
a single specimen collected by Hedgcock at Pikes Peak in Colorado. Later,
Shope (1931) stated it was conspecific with P. circinatus Fr.
GossELIN: STUDIES ON POLYSTICTUS CIRCINATUS 527
Sattory and Maire (1922) believed that P. circinatus is only a variety
of P. tomentosus, a point of view shared by Haddow (1941).
Konrad and Maublanc (1926) under Fomes (Xanthochrous) circinatus
furnished a picture of curved setae and described it as straight or curved.
We previously mentioned the same thing for the wood-loving forms we
found in the field. It should be P. circinatus Fr. Bourdot and Galzin
(1927) made the same observations on what they called P. circinatus ae
Shope (1931), Overholts (1933) and Lowe (1934) followed Lloyd’s
opinion and do not recognize P. tomentosus in the American Flora.
Furthermore, Overholts states that all American forms should be referred
under the name P. circinatus. All three described the typical form as
having straight setae because they have been led astray by the statement
of Fries that the species was characterized by the homogeneous context
of the fruiting bodies. Concerning P. dualis Pk., Shope considered it
the same as P. circinatus, while Overholts gave a varietal rank because of
the curved setae which in the typical form are straight. Lowe considered
it a good species.
Jérstad and Juul (1938) worked out the taxonomy of the species with
Norwegian material. They recognized two varieties to P. tomentosus Pr.:
P. tomentosus var. americanus and P. tomentosus var. circinatus. They
made the variety americanus because of Lloyd’s statement (1912) that
P. tomentosus Fr. does not exist in the American flora, but only a related
form. The variety circinatus was suggested for P. circinatus Fr. because
they said there is no definite line of demarcation between Fries’ species.
They make no mention of Sartory and Maire who made the same va-
riety in 1922.
Haddow, in 1941, based all his discussion on specimens of the Curtis
Herbarium identified and labelled by Fries himself. He recognized the
duplex context for both P. tomentosus and P. circinatus; the former had
straight setae and the latter curved ones. Haddow did not agree that
the curved setae character was sufficient to make a new species as did Fries,
thus he called it P. tomentosus var. circinatus.
As a resumé, the following table shows the real position of each specimen.
Author Polyporus tomentosus Fr. Polyporus circinatus Fr.
- Fries (1851) Polystictus tomentosus
Peck (1878) P. dualis
Cooke (1886) Polystictus tomentosus Polystictus circinatus
Saccardo (1888) Polystictus tomentosus Polystictus circinatus
Karsten (1889) . Onnia tomentosus Onnia circinatus?
Ellis & Everhart (1889) Mucronoporus tomentosus Mucronoporus circinatus
Hennings (1898) Polystictus tomentosus Polystictus circinatus
Patouillard (1900) Xanthochrous circinatus
Murrill (1904) Coltricia tomentosa Coltricia tomentosa
~ Lloyd (1908) (1912) Polyporus tomentosus Polyporus dualis
Polyporus circinatus
Ames (1913) ; Polystictus tomentosus Polystictus circinatus
Neuman (1914) Polyporus tomentosus
Polyporus dualis
528 FarLowia, Voi. 1, 1944
Author Polyporus tomentosus Fr. Polyporus circinatus Fr.
Bresadola (1920) Polyporus dualis
Polyporus triqueter
Sartory & Maire (1922) Polyporus tomentosus P. tomentosus var. circinatus
Konrad & Maublanc (1926) Fomes (Xanthochrous)
circinatus
Bourdot & Galzin (1927) Polyporus circinatus
Killerman (1928) Polystictus tomentosus Polystictus circinatus
var. triqueter
Shope (1931) Polyporus circinatus
Polyporus dualis
Overholts (1933) Polyporus circinatus Polyporus circinatus
var. dualis
Lowe (1934)- Polyporus circinatus Polyporus dualis
Jérstad & Juul (1938) Polyporus tomentosus P. tomentosus var.
circinatus
P. tomentosus var.
americanus
Haddow (1941) Polyporus tomentosus P. tomentosus var.
circinatus
It is clear, from what has been written above, that the tangled taxonomy
of the two types would not have resulted had it not been for Fries’ early
statement that Polyporus tomentosus possessed a homogeneous context,
whereas a study of his specimens by the writer and others, has shown
that the context of the sporophore is duplex in both types, the first of
which should be recognized as P. tomentosus (with straight setae) and
the second P. circinatus (with curved or straight setae in the same fructi-
fication). Assuming for the time being that the two species belong in
the genus Polyporus, it appears to the writer that Fries was correct in
making these separations, since, as it has been stated above, the two species
are obviously different when seen together in the field (Plate I, fig. 1, 2).
It is for this reason that the writer considers it desirable to follow Fries
in recognizing these species as distinct rather than to follow Sartory and
Maire, and Haddow in reducing P. circinatus to varietal rank under
P. tomentosus.
The generic position of these two species would be greatly simplified
if they were kept in the genus Polyporus as has been done by many, al-
though Murrill (1916) and Donk (1933) and others have tended to divide
the genus into smaller units and probably rightly so. On the basis of
priority, Fries’ genus Polystictus (1851) must be accepted for this and
related species, since P. tomentosus was specifically mentioned as an ex-
ample of the genus and therefore can be taken to represent the type of the
genus. For this reason, it seems best to consider that Polyporus tomen-
tosus and Polyporus circinatus should be known as Polystictus tomentosus
(Fr.) Fr. and Polystictus circinatus (Fr.) Cke.
DISTRIBUTION
Since they have first been described by Fries, P. tomentosus and P.
circinatus have been found abundantly in Asia, Europe and America.
GOSSELIN: STUDIES ON POLYSTICTUS CIRCINATUS 529
Because frequently they have been misnamed, the writer has prepared
from the literature a list of localities and has tried to indicate the dis-
tribution under P. tomentosus Fr. and P. circinatus Fr. Where there is
any question of the accuracy of the determination, the author name is °
followed by a (?).
Nova Scotia
New Brunswick
Locality Polystictus tomentosus Polystictus circinatus
(Fr.) Fr. (Fr.) Cke.
Africa Lloyd(?) (1920)
Austria Lloyd (1908)
China Teng (1932)
Canada
Prince Ed. Is. Ell. & Evh. (1889)
Ontario Lloyd (1908), Faull (1922)
Manitoba & Bisby (1938)
Saskatchewan
McKay(?) (1904, 1908)
Hay (1903, 1905)
Wehmeyer (1940)
Hay (1903, 1905)
Germany All. & Schn. (1886), Hen- Hennings (1898) Murrill
nings (1898), Murrill (1904) (1904)
Japan Shirai(?) (1927
Sweden Fries (1863), Lloyd (1908, Fries (1863), Lloyd (1908),
United States
California
Colorado
Dist. of Columbia
Florida
Idaho (northern)
1912), Murrill (1904, 1908) Maurrill (1908)
Lloyd (1911, 1915)
Kauffman (1921), Lloyd (1908,
1916, 1920), Murrill (1904),
Seaver & Shope (1936),
Shope (1931)
Lloyd (1922)
Lloyd (1920)
Boyce (1938), Hubert (1929, Haskell & Wood(?) (1930),
1931), Martin (1929) Hubert (1931)
Murrill (1915)
Towa Wolf (1931)
Maine Lloyd (1923), Murrill (1904) White(?) (1902), Ricker (?)
(1902)
Maryland Lloyd(?) (1908)
Massachusetts Lloyd (1907, 1908, 1911, 1912,
1913, 1914 1915, 1921),
Murrill (1904)
Michigan Kauffman (1911), Lloyd
(1912), Longyear (1904),
Overholts (1916), Povah
(1935)
Minnesota Hubert (1924, 1931), Lloyd Haskell & Wood (1930)
(1920, 1922) Martin (?)
(1925)
Montana Hubert (1931)
New Hampshire Murrill (1904)
New Jersey Murrill (1904) Ellis(?) (1890), Ell. &
Everh.(?) (1889)
New York Burnham & Latham (1914, Peck (1878, 1894)
1924), Lloyd (1913, 1915,
1916, 1920, 1923), Murrill
(1904), Peck (1869, 1880,
1893, 1899) , Saccardo (1888)
‘eteahaeil oe he rele iL ae
530 Fartowia, Vou. 1, 1944
Locality Polystictus tomentosus Polystictus circinatus
(Fr.) Fr. (Fr.) Cke.
North Carolina Curtis (1867), Lloyd (1914) Atkinson & Schrenk(?)
(1893)
Pennsylvania Overholts (1933)
Rhode Island Bennett (1888)
Tennessee (east) . Hesler (1929)
Vermont Lloyd (1908, 1911, 1915),
Murrill (1904)
Washington Hubert (1931), Lloyd (1916)
West Virginia Lloyd (1908, 1920), Mur-
rill (1904) _
Wisconsin Dodge (1914), Neuman
(1914), Overholts (1916)
Wyoming Seaver & Shope (1935, 1936)
To complete this list, the writer has plotted on a map the North Amer-
ican localities of the specimens of P. circinatus (Fr.) Cke. kept in the
Farlow Herbarium. It seems that this species reaches its southern limit
at the 36th parallel of latitude (Plate I, fig. 3). It is probably limited
by the temperature and if so it could be expected farther south in the
mountains.
History AS A PARASITE
Many writers have discussed the taxonomy of the two Friesian species
since 1878, and still more mycologists have collected them in the forests,
but very few, either in the Eastern or Western Hemisphere, have worked
out the parasitism of these species.
In the Eastern Hemisphere, Jgrstad and Juul (1938), after a discussion
of the taxonomy of the species, questioned whether P. tomentosus Fr.
has any parasitic tendencies and mentioned the fact that in Sweden it has
been found on a root of spruce which did not present any rot. However,
the ability of P. circinatus to cause rot is not doubtful and they gave the
general characters of the rot as it occurs in Sweden and mentioned the
similarity that this. particular one has with that caused by Fomes pini.
The first in North America to report this species as a parasite was
Dr. Faull (1922) who found that, in Ontario, it was causing a root- and
butt-rot in spruce, hemlock and eastern white pine. Giving the descrip-
tion and habits of the sporophores, he identified them as being P. tomen-
tosus F'r., but his herbarium specimens for that region show that P. cir-
cinatus Fr. was also abundant. He mentioned the fact that the infection
was very much localized and further added that in the Otter District of
Ontario a high percentage of white pine is affected.
Hubert (1931) reported that P. circinatus attacked roots and butt sec-
tions of western white pines in northern Idaho, but considered it to be
a wound fungus and commonly associated with fire scars. He made a
comparative study of P. circinatus and Trametes Pini and clearly de-
scribed the similarities and differences between the two rots based on the
macroscopic and microscopic characters of the two different decays, and
the differences in the cultural characters of the two pathogens.
GOSSELIN: STUDIES ON PoLysTICTUS CIRCINATUS 531
Christensen (1940) published a brief note on P. circinatus, giving the .
cultural characteristics and a description of the decay and of the sporo-
phores and mentioning the importance of this disease in northern Min-
nesota as the cause of windthrow and premature death of the trees.
From this brief resumé of the literature it is apparent that although
we have a certain body of facts in regard to the type of decay, the micro-
scopical and macroscopical characters of the decayed wood, and of the
cultural characters of the fungus, we have very little knowledge concern-
ig the parasitic nature of the fungus, of geographical or other factors
influencing infection, and little information about the biology of Polystic-
tus circinatus. For these reasons and since the fungus seems to be of
local importance because of its effect on the spruce forests of the Province
of Quebec, the writer decided to make more intensive studies of the fungus
in an endeavor to discover further facts bearing on the biology of the
fungus and its relation to the host. At the same time, it was hoped to
explain the relation of other butt-rot fungi to their hosts.
STUDIES ON THE HOST-PARASITE RELATIONS
FIELD OBSERVATIONS
In undertaking this study of Polystictus circinatus, an intensive survey
was made to determine the occurrence of this speciés in the Province of
Quebec, especially along the north shore of the St. Lawrence River and in
the Gaspé Peninsula as well as in the Laurentide National Park area.
On the Gaspé Peninsula, and more especially along the northern side,
the fungus appears to be of rather frequent occurrence, whereas on the
south side it is somewhat scarcer. On the northern side of the Gaspé
Peninsula in the vicinity of Mt. Albert, infection? was locally very high,
(100%), in the York River area infection amounted to 95%, and at Lake
Ste. Anne approximately 50%. On the north shore of the St. Lawrence
River, near Lake Walker, the infection in the valley was about 20% and
on the mountain slope up to 80-90%. On the other hand, in the Lauren-
tide National Park, no evidence of infection could be discovered, even
though that region is intermediate between Ontario, where Faull (1922)
states that the fungus is very abundant, and the North Shore of the
St. Lawrence.
The high degree of rotting of spruces by Polystictus circinatus indicates
that this fungus may be of considerable importance in the Province of
Quebec, and the gap in the distribution of the species represented by the
complete or nearly complete absence of the fungus in the Laurentide Na-
tional Park indicates that there may possibly be some ecological factor
involved which not only would furnish information on the distribution of
P. circinatus but might also help to explain host-parasite relations in the
heavily rotted areas.
* As shown by the obvious symptoms of butt-rot at stump section.
552 FartowiA, VoL. 1, 1944
For the understanding of the behavior of the disease, it would be of
interest to know if this disease occurs on trees weakened by age, crowding,
competition with other species, by storm or insect damages.
Christensen (1940), according to observations made in Minnesota, states
that P. circinatus can infect trees as young as fifteen to twenty-five years
old. From the writer’s observations it can be stated that trees of forty
years age presented visible decay at the stump section. Since they must
have been infected earlier, it is probable that the age of infection given
by Christensen is near the truth. According to the same observations,
Christensen stated that “The fungus does not merely accompany stagnation
and senescence but, if present at all, is likely to be one of the primary
causes of such conditions.” The writer agrees with Christensen that it
sometimes happens that P. circinatus will kill the tree, but he does not
think that it will always bring a premature senescence of the trees. From
the data obtained from the Gaspé and North Shore regions, it is shown that
the average age for the infected trees in Gaspé is 90 years, while it is 130
years for the North Shore region. The average age for sound trees is
respectively 85 and 114 years. Thus, it seems significant. that, on the
whole, infection by P. circinatus appears to delay senescence.
The effect of crowding does not seem to have much influence on the
occurrence of the disease. The disease was found where tree cover was
very dense or loose, fruiting bodies having been found in places where the
canopy was so dense that herbaceous plants could not grow, as well as in
places where the canopy was sufficiently open to permit the growth of these
covering plants. Hence, crowding is not a determining factor for the
disease. Infection occurred either in pure stands of spruces or in mixed
stands of spruce and fir. But in these mixed stands, infection has rarely
been found when the percentage of firs was higher than 60%. This might
be explained by the fact that the fir grows where soil is richer and where
infection of spruce is relatively infrequent. Incidentally, P. circinatus
was not found in stands of mixed soft and hardwoods, although P. tomen-
tosus occurred in such stands. Again this may be explained by the fact
that infection is less frequent in richer soils. Competition between host
species does not seem to favor any tendency for infection. It can be
stated, rather, that infection is most commonly found in pure stands where
conditions for the growth of the trees are less favorable.
In the Gaspé region and on the north shore of the St. Lawrence River,
observations were made on the occurrence of the butt-rot; there was no
correlation between spruce sawfly epidemic, which was very severe, and
the rotting of spruce by P. circinatus. At Mount Albert and at Lake Ste.
Anne the black spruce was rotted, whether damaged or undamaged by the
spruce sawfly, and the disease was found independently on both of them.
Furthermore, in places like Petite Riviére-a-Marte and York River where
there was up to 95% and 100% infection by the disease, the damage by
the spruce sawfly was nonexistent. Except for the spruce sawfly epidemic
GOSSELIN: STUDIES ON POLysTICTUS CIRCINATUS 533
we have not seen, during our work, any sort of insect attack. It is there-
fore fairly evident that the rot caused by P. circinatus is not related to, or
a result of, insect attacks.
Storms which break branches or make scars on the trunks of trees do
not appear to have any influence on infection by P. circinatus, since infec-
tion takes place through underground parts. Nevertheless, a large num-
ber of forest soils in Quebec being thin, the rooting of trees is superficial
and hence, under the action of the wind the trees oscillate and some small
roots may break off and infection by P. circinatus may take place. Yet
careful studies did not furnish evidence of increased infection. In addi-
tion, it may be remarked that in no case did entrance of the fungus into
the wood take place through other parts than those underground, or which
were at least in contact with the duff. Hubert (1931) states that this
disease is often associated with fire scars but we believe that those scars
had to be in contact with the ground. In places where infection was up
to 85% and 100%, we have not seen a single case in which infection oc-
curred at other places than those mentioned above. However, from inocu-
lation experiments carried out in Duchesnay, it has been found that if the
mycelium comes in contact with a scar on the trunk, it will invade the
wood cells. Thus, it can be suspected that under natural conditions, the
spores of the fungus must germinate in the humus.
In our North Shore experiments we have organized some kind of rudi-
mentary meteorological stations in order to determine if temperatures,
maximum and minimum, the quantity of rainfall, the relative humidity of
the air, and the evaporation rate would be reflected in the number of in-
fected and non-infected stands. The different measurements obtained at
the two places had no significance, and none of the meteorological factors
could be correlated with infection or non-infection of the stand.
As a rule we can state that the fungus invades very young trees and is
seen in old ones, but the occurrence of this disease is not correlated with
any crowding, competition or damage of any sort, or with local meteoro-
logical factors.
EFFECTS OF TEMPERATURE AND HUMIDITY ON SPOROPHORE PRODUCTION
Bearing in mind that the practical aspect of the subject is the control
of the disease, field observations were made to determine the factors bring-
ing about the fructification of the fungus, since the control of a disease
is sometimes attempted by the removal of the fruiting bodies.
During the summer of 1941, which the writer spent in Shelter Bay on
the North Shore of the St. Lawrence River, it was only on the 15th of
August that he found sporophores of Polystictus circinatus. As they are
produced at about the same time of the year, he has tried to correlate the
maximum and minimum temperature and the relative humidity of the air
with the appearance of sporophores. For this purpose the following fig-
ures of temperature and humidity have been brought together in table I.
534 FartowisA, Vou. 1, 1944
Table I
Givine THE Maximum AND MINIMUM TEMPERATURE AND THE RELATIVE HUMIDITY
or THE Arr IN SHELTER Bay, FROM JULY THE 15TH TO AUGuUsT THE 21sT 1941
Temperature Temperature
Date Max. Min. Humidity Date Max. Min. Humidity
15 July 72.0 51.0 86 4 August 57.0 49.0 65
A sez 55.0 48.0 55 ele 59.0 47.0 50
Arg ae 64.0 53.0 88 te 58.0 45.0 58
1 Blea 64.0 54.0 58 pony 67.0 40.0 57
19 46.0 70 eat 54.0 40.0 79
De 72.0 48.0 74 ae 52:0 39.5 82
ps ay 82.0 50.0 82 ke 65.3 45.0 67
A ahh, 93.0 61.0 69 i Pe A 38.6 53
pe Ua 92.0 64.0 51 12 : 68.0 47.5 74
~ | ia 61.0 54.0 79 1s el 49.0 38.0 86
ae 69.0 49.0 60 ist 58.5 35.5 93
a ea 70.0 47.0 37 year 66.5 32.0 ref
7) aed 72.0 47.0 33 ll bee ed 64.5 51.5 94
29 «“ 71.0 53.0 47 i EF eh ia 62.3 43.3 82
Ft ae 61.0 51.0 43 bees f 60.5 36.5 74
Eth 58.0 48.0 94 iL earl 70.0 B1.5 65
1 August 61.2 45.0 68 5 A a 60.3 48.3 100
Batt 59.0 53.5 100 Bikwak ts 67.0 33.8 76
5 eS oe 48.0 84
It can be seen from the graphs of maximum and minimum temperatures
(Graph 1) that the average of both has a tendency to be lower after the
15th of July. Also, it can be seen that the maximum temperature after
the 30th of July averaged 60° F. and fell below 50° F. on the 15th of
August. It seems that the 60° F. temperature is the critical one for
initiating the fructification of P. circinatus. Since the figures given in
Table I and in Graph 1 are for air temperature it should be pointed out
that the temperature of the soil and of the tree would most likely be
neither as low nor as high as the air temperature. Although those data
have been taken only during one season and at one locality, they seem to
indicate fairly well that when the average temperature of the day ap-
proaches the minimum temperature for mycelial growth, the vegetative
phase is checked and the reproductive stage is initiated a short time before
fruiting bodies actually appear.
Reference to Graph 2, presenting the percentage of relative humidity of
the air against the time of the year make it evident that after the first day
of August and up to the time fruiting bodies appeared, the percentage of
relative humidity did not go below 50% and averaged 81.7%, while before
that date, from the 15th of July up to August, the relative humidity was
as low as 35% and averaged 64.3%. From the 15th of August up to
GrapH 1. (Above) The relation of maximum and minimum temperatures to the
appearance of sporophores.
GrapH 2. (Below). The relation of atmospheric humidity to the appearance of
sporophores.
GOSSELIN: STUDIES ON PoLysTicTUS CIRCINATUS Sao
TEMPERATURE
Appearance of the first sporophore
Degrees Fahrenheit
4
{
7? i
duly he Anon,
HUMIDITY
Appearance of the first sporophore
Percentage of relative humid lity of ihe air
1
t
July * August
GrapH 1, 2
536 Fartowia, Vou. 1, 1944
September, the best time for sporophore production, the relative humidity
was never lower than 65%, with an average of 85.7%. We have remarked ©
that the sporophores come on trunks or roots always very close to the soil,
a fact that substantiates the evidence tending to show that a relatively high
humidity favors not only growth, but at a later date, when vegetative
growth has been checked, also favors the production of fruiting bodies.
Just how much the percentage of relative humidity becomes a limiting
factor cannot be asserted from the data we have, but it is at least certain
that an average of around 80% is favorable.
As far as light is concerned, it can be said that the sporophores are
more generally found in diffused light. However, in one case, one was
found exposed to the afternoon sunlight. From laboratory studies it has
been found that the related P. tomentosus grows better in diffused than
in bright light or in darkness.
Although not conclusive, these data furnish good indications that the
appearance of the sporophores of P. circinatus is correlated principally
with temperature and humidity and that the time of fruiting corresponds
with the period when the low temperature checks vegetative growth and
stimulates reproductive growth. This period also correlates with the
time when the average relative humidity is on the increase and thus is
furnished the water necessary to enable the fungus to complete the fruiting
stage and to ripen the sporocarps. Exposure to light appears, if it is im-
portant at all, to play only a very secondary réle in initiating the forma-
tion of the fruiting body. However, it is possible that once the fructifica-
tions are above the surface of the soil, light may have some influence on
the formation of the hymenium and on the production of the spores. If
this is so, since the fruiting bodies are produced mostly in the deep shade,
it is evident that only very low intensities of light are needed to exert an
influence on fructification.
Errects on Host
In addition to some ecological data concerning the production of sporo-
phores, field observations have also given some information concerning
the effects of this disease on its host as, for instance, the external charac-
ters which diseased trees present and the kinds of breaking the rotted,
fallen trees will present.
The writer has observed that in early stages of infection the trees do
not present externally any signs of disease, but as the infection reaches an
advanced stage, the foliage of the trees turns pale yellow-green. Another
external character which sometimes accompanies the advanced stage
of decay is the exudation of resin at the base of the trees. Although
these characters are not specific to the infection by P. circinatus, they help
to spot the disease in regions where it is known to occur. It has already
been stated that field observations indicate that the infection of the host
takes place through the lateral roots, as is shown by the fact that when
subjected to strong winds the trees blow over very readily and in a char-
GOSSELIN: STUDIES ON PoLysTICTUS CIRCINATUS DOU
acteristic manner. By the comparison of trees infected by P. circinatus and
of those attacked by Poria subacida, this point can easily be brought out.
In this latter species, it was found that penetration was through the central
or tap root and from there the rot progressed into the stump leaving the
lateral roots unharmed; as a result the trees broke off about one to two
feet above ground level. In P. circinatus, on the other hand, when the
lateral roots have been weakened by decay the trees blew over and at the
same time the tap roots were pulled from the ground. Further studies
substantiate the above evidence. For example, at Shelter Bay in the
valley level where both fungi occurred, the trees were blown over in a
manner characteristic of trees infected by the two species of fungi. In this
connection it should be pointed out that in the valley the soil is sufficiently
deep so that the spruces develop both tap and lateral roots. On the moun-
tain slopes however, where the rock is covered only by a layer of humus,
and as a result tap roots are not developed, Poria subacida was absent
and only P. circinatus was evident. Under such conditions as are present
in the mountain habitat, only lateral roots could be infected. Thus there
is furnished the added evidence that P. circinatus is confined to the lateral
roots, at least during the early stages of growth.
GROWTH STUDIES
During the course of the studies in connection with penetration of the
host by the fungus it became evident that it was desirable to make a com-
parative study of the growth of trees infected by P. circinatus*® and non-
infected ones, and at the same time to endeavor to discover the relations
between the habitat and the infection of the trees as illustrated by their
rate of growth. A survey was also made to find out whether or not there
was any correlation between the age of the tree and the incidence of butt-
rot. For this purpose studies were made in five localities, namely: the
valley and mountain slope and summit in the vicinity of Lake Walker on
the North Shore of the St. Lawrence River, and in the vicinity of York
River and Lake Ste. Anne on the Gaspé Peninsula where ecological condi-
tions seemed to be similar.
In these studies sections were made at D.B.H. of trees infected by
P. circinatus and those not infected. In each type, the age of the tree was
determined by a count of the annual rings and the diameter of the tree
measured for each ten-year interval. The trees were segregated in age
classes: (40-60) (70-90) (100-120) ... etc. For each year class and
for each locality the diameter of the average tree was calculated and from
that the area of D.B.H. section was obtained for each ten-year interval.
* During the discussion on the growth rate of the trees, the terms “trees infected
by P. circinatus” and “trees non-infected by P. circinatus” have been used having in
mind the fact that the trees do or-do not present visible signs of decay. It cannot
be stated that trees which had no decay caused by P. circinatus were not-in mycor-
rhizal-association with this fungus, nor can it be said that no other mycorrhizal
fungus is present.
2
538 FarLowiA, VoL. 1, 1944
We assumed that the surface of the section of a conifer at breast height
increased in proportion to the increase in volume of the tree. The growth
rate was obtained by subtracting the logarithm of the surface of the sec-
tion of two subsequent decades. Appendix I gives the area at D.B.H.
section for each ten years and for each classification, Appendix II gives
the corresponding growth-rate. In Graph 5, the rate of growth of infected
and non-infected trees for the 130-150 year class have been plotted for
different sites. From this graph it can be seen that in the York River and
the Valley areas where the soils are favorable, the rate of growth of non-
infected trees is greater than that of infected trees. Contrarily, on the
East Slope and Mountain Top sites where the soil is shallow and leached
and hence less favorable, the rate of growth of the infected trees is greater
than those that are not infected. The significance of these facts will be
discussed shortly. At this point it may be objected that there are not
valid grounds for drawing any conclusions since the trees of each class
may have encountered different ecological conditions during the different
years of their growth. To answer this objection, the average rate of
- growth of trees for a given site and of infected and non-infected trees,
together with the rate of growth of individual trees in each classification,
was plotted by age (Graph 3) and by year (Graph 4). A comparison of
these two graphs demonstrate that although there are minor differences,
there is in general a very close agreement between the results derived by
the two methods. Thus, whichever method is employed, the results are
approximately the same and equally significant. For convenience and as
a resumé of Appendix II, Table II has been prepared to show for each age
class and for each region the percentage of number of decades in trees
rotted by P. circinatus in which the rate of growth was higher than those
not rotted by this particular fungus. Accompanying these percentages
are symbols which indicate whether the area at D.B.H. section of the
average tree rotted by P. circinatus is greater, equal, or smaller than that
of the non-rotted trees.
Table II
Gaspé Peninsula Lake Walker Area
Age class L. Ste. Anne York River Valley East Slope Mountain Top
40-60 50% [] 0% -- 67% ++
70-90 100% + 86% + 75% ++ 29% []
100-120 30% ++ 70% -- 4%+ 56% + 64% ++
130-150 42% —— 42% — 54% + 42% ++
160-180 388% —— 74% ++
190-210 38% ——
Average 62% 538% 38% 65%. - AT%
Area of rotted trees definitely higher ++, somewhat higher +, indifferent [ ],
somewhat smaller —, definitely smaller ——.
GrapH 3. The average rate of growth (heavy lines) and the growth of individual
trees (light lines) of the 130-150 year class plotted by age.
Grapu 4. The average rate of growth (heavy lines) and the growth of individual
trees (light lines) of the 130-150 year class plotted by year.
ee ee
Growth rate
0.9 fr 0.9 -
aa
0.8 og f : \
O7- i O7- \ ——. Trees infected by P,circl
\ . ——— Trees non-infected by P.circinatus Or ae ee Mme Pare 0g \ Nett eet eal a tN a Bee ono Trees infected by P, circinatus
‘ \ ==---+ Trees infected by P. circinatug ;
0.6 0.6 F
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o
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10 1872 1882 1392 1902 1912 1922 1932 1942
: Date
GraPH 3 GRAPH 4
SALYNIDUI) SALIILSATOG NO SAIGNLS :NITASSON
6€S
540
Fartowia, VoL. 1, 1944
AGE CLASS 130 = 150
GRAPH 5
ri
York River 1
w %
0.45 + ‘ _
g J
qq
0.25 + \ "———~ Trees non-infected by P.circinatus |
\ o----* Trees infected by P.circinatus
-
|
| t mae 1 1 ? . : aie SEES mote oe)
P ce = ?
R Mountain Top Aaa
E \ a J
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10 30 60 70 90 110 130 160
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GOSSELIN: STUDIES ON POLYSTICTUS CIRCINATUS 541
Reference to Appendix I and Table II shows that in the two localities:
York River and Valley, the area at D.B.H. section of the average rotted
tree by P. circinatus is lower than that of the non-rotted trees except in
age classes 70-90 and 100-120, but in all other localities the rotted trees
have larger area at D.B.H. section than the non-rotted ones. Thus, it
can be concluded that either the infection of the tree by this fungus is
beneficial to the host or else that this fungus invades rapidly growing trees.
This last hypothesis has to be discarded because the data from York River
and Valley regions (Graph 5) show that the reverse is true. Examination
of growth rates, Appendix II and Table II more especially, show that in all
cases there always has been a period during the life of the tree when the
growth-rate of trees infected by P. circinatus was higher than the non-
infected ones, except for the age class 40-60 of York River. The per-
centages given in Table II show how long was this period. To explain why
in York River and in the valley of the Lake Walker site, the area at D.B.H.
section of trees infected by P. circinatus is smaller, it may be said that
in these two regions the conditions for growth of the trees were not as poor
as in the other regions, and that the growth of young non-rotted trees was
at the beginning very good, while for some unknown reason it was some-
what delayed in the trees rotted by P. circinatus. Thus. starting with a
larger capital, the area at D.B.H. continued to be larger throughout the
life of the tree even though the rate of growth was smaller during a few
decades. On the other hand, in the other localities where the conditions
of tree growth were poorer, the initial capital of the non-rotted trees was
smaller or only slightly greater and was later exceeded by the faster
growing but rotted trees.
In general it can be said that when there is little sub-soil, as under the
conditions existing in the mountain slope and top, the trees show a larger
diameter when rotted than when not rotted; on the contrary, when there
is a considerable depth of sub-soil such as exists in the valleys, then the
reverse holds true, but in all cases, with one exception, rotted trees always
present a. higher rate of growth, at least for a time, than do the non-
rotted ones.
Soi, CONSTITUTION
Not having found any correlations between the occurrence of the disease
and the state of crowding or competition between species, or the appear-
ance of any insect epidemics, to explain the different relations between
fungus and host in the various habitats, we have tried by soil analysis to
correlate the occurrence of this disease with some salt deficiency. For
this purpose we have used the method given by Hester, Blume and Shelton
(1937) for rapid soil analysis. The results are shown in Appendix III
which have been made to illustrate the effect of the concentration in the
GrapH 5. The rate of growth and area at D.B.H. section of infected and non-
. infected tree at four different sites.
542 FarLtowia, VoL. 1, 1944
soil of phosphorus, CaO, NH4, and potassium on the percentage of rotted
trees in the stand. The conclusions are briefly summarized as follows:
1) — Phosphorus: From 0% to 30% infection, increasing concentra-
tion of phosphorus is correlated with increasing infection of the stand, but
for higher infection than 30% no correlation can be made with the con-
centration of this element.
2) — Calcium: From 30% to 80% infection, increasing concentration
of calcium seems to be correlated with increasing infection of the stand,
but outside of these percentages of infection, nothing can be correlated.
3) — Ammonium: From 10% to 80% infection, increasing infection
of the stand is fairly well correlated with increasing concentration of am-
monium in the soil.
4) — Potassium: Up to 50% infection, decreasing potassium concentra-
tion in the soil accompanies increasing infection of the stand.
5) — H-ion concentration: In all the cases seen, infection was never
found in stands where the humus had a greater H-ion concentration than
pH 3.5. It seems that the average H-ion concentration has a tendency
to be lower in stands with a higher percentage of infection, although this
does not seem very specific. From culture studies, we know that this
fungus can grow in artificial media having a pH between 3.5 and 8.6.
The medium with a pH of 6.4 supported the fastest growth, but since there
were not any cultures intermediate between pH 4.5 and pH 6.4, it cannot
be stated that pH 6.4 is the optimum for growth of the fungus. Never-
theless, the optimum pH appears to be around pH 6.4. The pH of the
humus of almost all infected stands are comprised between pH 3.58 and
pH 4.40. Thus, we can state that generally the mycelium infects trees
when conditions are not the optimum for the growth of the fungus, and
accordingly the infection of trees could be interpreted as a reaction of the
fungus against adverse conditions.
Thus, in so far as the percentage of rotted trees in the stand is not too
high, the increasing degree of infection of the stand can be correlated
with increasing concentration in the soil of phosphorus and ammonium,
and decreasing concentration of potassium and calcium.
Now, instead of correlating singly the elements with the degree of in-
fection of the stand, we will consider the elements by groups of two: phos-
phorus and calcium, ammonium and potassium.
1) — Phosphorus and calcium: It can be seen that, when infection is
at or below 30%, the phosphorus concentration increases and calcium
concentration decreases. From 30% to 50% infection, the phosphorous
concentration decreases while calcium concentration increases. This goes
on up to 70% infection. From that degree of infection up to 100%,
calcium concentration will act in the same manner as phosphorus con-
centration. :
2) — Ammonium and potassium; Ammonium and potassium concen-
_ trations always act in opposite ways, except between 0% and 10%, 50%
eae ee eee ar ee pe EE De ee ER eS eg ae ee ee i ees ae ie ae
GOSSELIN: STUDIES ON PoLysTICTUS CIRCINATUS 543
and 60% infection of the stand. So, it seems that as long as there are
opposite (or mirror) correlations between phosphorus and calcium con-
centrations in the soil, the stand will have only weak infection. But as
soon as these two elements correlate in the same sense, the stand is sus-
ceptible to a high degree of infection. In general, however, the correla-
tion between ammonium and potassium does not seem so important.
NEEDLE ANALYSIS
According to experiments done at the Black Rock Forest, New York,
the salt concentration in the leaves, or needles, is nearly proportional in
pure culture to the concentration of these elements in the soil. So, to
complete the information given by the soil analysis, we have analyzed
needles of spruces from localities in which there were different degrees
of rotting. The potassium, ammonium and phosphorus percentage of
the dry weight increase in the needles with the increase of infection up
to 30% (see Appendix IV). Even if in the soil the potassium concentra-
tion is low, the percentage of dry weight of this element in the needles is
relatively high. It should be remarked also that the data in the three
cases are about similar.
Although there are some indications from the soil analysis that increas-
ing phosphorus and ammonium and decreasing potassium and calcium
will favor rotting, those indications are valid only in the cases of low
percentages of rotting. There must be some other factors which, in higher
percentages, will govern the infection, but unfortunately we have not been
‘able to determine such factors. Even the needle analysis does not give
much information for greater rotting than 30%.
EVIDENCE OF SYMBIOTIC RELATIONSHIP
If soil and needle analysis do not give much information as to the
factors determining the occurrence of rot, they do furnish good evidence
that this parasitic fungus forms a symbiotic association with its host before
becoming parasitic. The evidence is furnished by the fact that, as has just
been pointed out above, under unfavorable conditions existing on the moun-
tain top, rotted trees show greater growth than the non-rotted trees, whereas
in the valleys where the essential nutrient elements have accumulated, the
reverse is true and indicates that the symbiotic relationship is only effective
when there is some mineral deficiency. The evidence is also furnished
by the fact that infection takes place by the lateral roots and that sporo-
phores of P. circinatus have been found associated with tree rootlets. The
results of needle analysis also tend to show that there is a symbiotic rela-
tionship between the host and the fungus.
As stated previously, the introduction of the fungus into the host is
made by way of the underground parts of the tree. Thus, the only possi-
ble way for the mycelium to penetrate the host will be either by under-
ground scars, by direct penetration through the bark or symbiotic asso-
ciation with the tree. We have dug out about a dozen rotted trees in Ste.
ee
544, FarLowl1A, Vout. 1, 1944
Catherine and in Shelter Bay, and there are only a few cases where infec-
tion took place through scars on the roots. In all the other cases we
could not find any particular place where the mycelium would have pene-
trated. Rather, we have seen the rot behaving exactly as a river. The
main body of the river is the main lateral root and all or many of the
secondary roots bring rot to the main lateral roots, and these meet in the
trunk. Dr. Faull, in a personal communication, stated that he followed
the rot up to the very tip of a root. The writer has made similar observa-
tions in Shelter Bay. Hence the mycelium must have penetrated directly
through the bark or by symbiotic association. We do not have any evi-
dence that it came through the bark, while we do have some for the sym-
biotic association hypothesis.
At least twice, sporophores of P. circinatus were found growing out of
nests of rootlets. Hatch (1937) reported that sporophores coming out
of rootlet-nests are quite frequent in the usual mycorrhizal fungi, and
therefore since P. circinatus has twice been found connected with rootlet-
nests, the evidence for its mycorrhizal relationships is further strengthened.
As has been previously stated, in the Gaspé and in the Lake Walker area
the growth rate of rotted trees was higher than that of healthy ones. It
is accepted that trees with mycorrhizae will have a larger rate of growth
than those without, at least when the mycorrhizal relationship is efficient.
On the other hand, we have seen that in the Valley (North Shore) of the
St. Lawrence River and on York River (Gaspé), the diameter was smaller
for rotted trees than for healthy ones. This has been correlated with the
fact that in these two places the soil is richer in nutrient salts and better,
apparently, for tree growth without the aid of P. circinatus as a partner,
at least when it does not cause any rot.1_ We have seen also that the
humus of these two places had less phosphorus than the others had.
Rosendahl (1941) found that when the phosphorus concentration of soil
was low, although mycorrhizal relationship was established, the intake of
nitrogen and potassium was less than in similar seedlings grown with an
adequate supply of phosphorus. Therefore if symbiotic relationship is
to be of greatest benefit to the tree, then a suitable concentration of phos-
phorus is required.
TaBLe III
THe CONCENTRATION OF THE Various MINERAL ELEMENTS IN THE Soi Is GIVEN
ACCORDING TO THE DIFFERENT LOCALITIES STUDIED
pH Kk NH, Fr CaO
Locality Inf Non InfNon Inf Non Inf Non Inf Non
Gaspé Peninsula
Lake Ste. Anne 3.81 3.79 31 35 23 19 5.00 3.638 165 1381
York River 3.80 38.74 39 35 26 24 4.75 6.50 97 260
L. Walker area
Valley 3.74 4.07 20 22 20s 22 2.60 12.50 80 40
Mountain Top 4.06 3.75 40 8 13 23 12.50 4.20 80 24
East Slope 3.95 3.95 5 23 15 32 12.50 10.25 40 48
‘Refer to footnote page 537.
Pst. ee ee
GOSSELIN: STUDIES ON POLYSsTICTUS CIRCINATUS 545
As seen from Table III the writer’s observations substantiate this con-
clusion since in the valley of the Lake Walker area and York River area
in Gaspé, the concentration of phosphorus was relatively low and this was
correlated with a low percentage of length of time beneficial to rotted
trees. It can be seen also that at Lake Ste. Anne where the concentration
of phosphorus was not optimum, the infection in age class 100-120 was
beneficial for only 30% of the time (Table II). The growth rates have
been discussed in greater detail in a previous paragraph.
Another good character which seems to indicate the symbiotic relations
of P, circinatus with the trees, is the salt content of the needles. Routien
and Dawson (1943) suggest “that mycorrhizae increase the salt absorbing
capacity of the roots . . . The increased salt uptake may then be reflected
in a greater rate of growth.” Finn (1942) stated: “It is seen that nitrogen
and potassium were taken up in larger quantities by the inoculated seed-
lings than by the uninoculated ones. Phosphorus was absorbed in a
slightly larger quantity by the inoculated seedlings than by the seedlings
which were not inoculated.” Thus, it is admitted now that plants with
mycorrhizae will generally absorb more nitrogen, potassium and phos-
phorus than those without.
In order to check the mycorrhizal nature of P. circinatus, we have
chemically analyzed spruce needles to find the percentage of these three
elements in the needles both in rotted and non-rotted trees.
The needle analysis reveals that the phosphorus percentage in relation
to the dry weight of the needles is lower for rotted trees than for healthy
ones in the general average and in the data from York River, while it is
higher at Lake Ste. Anne and at Chandler. Data from Finn (1942) reveal
that the phosphorus intake by the mycorrhizal seedlings is higher than for
the non-mycorrhizal ones, but the percentage to the dry weight is smaller
even though the intake is larger. The difference, as shown by Finn, is
0.020% while the writer found a difference of only 0.011%. On York
River this percentage is lower or only 0.007% while at Lake Ste. Anne it
is higher or 0.049% and at Chandler 0.054%. This, however, has not
definite significance because when there is a arenes amount of phosphorus
in the needles, there is a correspondingly greater concentration of this
element in the soil. Thus, it cannot be unqualifiedly concluded that the
infection by P. circinatus was beneficial to the trees although there seems
to be evidence tending to indicate that such is the case.
As in the phosphorus relations, the percentage of nitrogen in the needles
of rotted trees is higher, in the general average, than in the needles of
healthy ones. Although the general average is higher for rotted trees
than for healthy ones (Appendix VI), the percentage at Lake Ste. Anne
and at York River are lower, respectively of 0.002% and 0.003%, al-
though the soil analysis from these places (Appendix V) shows that soil
under rotted stands had a higher concentration of ammonium salts than
the healthy ones. Even if the nitrogen percentage of the needles is lower
546 FarLowl1A, VoL. 1, 1944
(0.002%) for rotted trees than for healthy ones, this difference is so
small that it probably is of little significance. However it is interesting
to note that nitrogen concentration in the needles is higher for rotted
trees than for healthy ones only when the concentration of nitrogen is
lower in the soil. For example, at Lake Ste. Anne and on York River,
the rotted stands had higher concentration of ammonium salt than the
non-rotted ones and the reverse was true for the percentage of nitrogen
in the needles. Exactly the contrary is found at Chandler. Neverthe-
less there seem to be indications that the infection by P. circinatus is some-
times beneficial to the host as a result of increasing the intake of nitrogen.
For the percentage of potassium in the needles, the analysis revealed
that, in the general average, this percentage is lower for rotted trees than
for non-rotted ones; however in the rotted trees of Lake Ste. Anne and
of Chandler this percentage is much higher. This low percentage in
rotted trees seems to be prevalent almost exclusively at York River. At
that place we have found percentages as low as 0.062, 0.071, 0.085, and
0.094. In none of the other places were percentages found as low as
these, and it was remarked that these low percentages came with heavily
rotted trees. In Gaspé only 22% of the infected trees were rotted as
high as D.B.H. section, but of the trees which have a low percentage of
potassium 60% were rotted up to the D.B.H. section. In general the
potassium percentage of the needles is low in all rotted trees of York
River, 0.236% compared with 0.372% at Chandler and 0.418% at Lake
Ste. Anne. This excessively low average might perhaps be explained by
the fact that 88% of the most heavily rotted trees were from York River
region. It can be seen from soil analysis that, at Lake Ste. Anne, al-
though the concentration of potassium in the soil was smaller for infected
stands than for healthy ones, the percentage of this element in the needles
was greater. It is logical to suppose that the infection, at least at Lake
Ste. Anne, has been beneficial to the trees for larger intake of potassium.
Thus if the data from York River region are excepted, we can state
that the infection of the trees with P. circinatus has increased the intake
of mineral salts. This beneficial effect should not be interpreted too
categorically, but rather as a probability. It should be remembered also
that the needles analyzed might, in many instances, have come from trees
in which the parasitic have exceeded the beneficial effects of the previous
symbiotic association. This would have lowered the average percentage
for the rotted trees and might explain why, in the York River region where
the trees are more heavily rotted than in the other regions, the percentages
of all elements are so low.
The field observations, in addition to some facts concerning the be-
havior of the disease caused by P. circinatus, has brought out facts which
seem to indicate fairly well that this fungus penetrates into its host by
symbiotic association. Each fact in itself is not very conclusive but they
all substantiate the same conclusion.
See ae eee ae ae a eee Or ye ee Ree Mee to ee
GOSSELIN: STUDIES ON PoLystTicTUS CIRCINATUS 547
EFFECTS OF SEEDLING INOCULATION WITH P. CIRCINATUS
Having in mind all the evidence from field observations that P. circinatus
was penetrating into the host through symbiotic association, the writer
has tried to establish the mycorrhizal relationship between spruce seedlings
and the fungus under laboratory control.
First, we have tried to culture seedlings under aseptic conditions with
the Hatch culture chambers, but the experiment did not succeed. Later,
loamy-clay soil was obtained from Saskatchewan where no trees are grow-
ing, and consequently in which there are no mycorrhizae forming fungi.
This was suggested by the experiments done by Hatch (1937). After
having sterilized and stratified spruce seeds, they were put in the ice box
for two weeks in order to obtain a better germination. The loamy-clay
soil was placed in wooden boxes previously sterilized and the seeds were
sown. After germination in July, 1942, the seedlings grew in a green-
house until January, 1943, when they were placed under the snow to allow
a dormant period. At the beginning of April, 1943, they were dug out
of the snow and put in the cold room (33° F.) for fifteen days to make
the transition between the temperatures of the snow cover and the green-
house less abrupt. After two weeks in the greenhouse, when the buds
were bursting, half of the seedlings were inoculated on May 1, 1943, with
cultures of P. circinatus, while the other half was reserved to serve
as a check, although maintained under the same growing conditions.
After three months of growth, namely on August 31, the seedlings from
both series were removed and studied to determine what difference ex-
isted. It was immediately obvious (Plate II) that the inoculated seedlings
had not only made a greater growth but that the needles were of a deeper -
green color and that the primary roots had produced more secondary roots.
To determine the relative growth of the inoculated seedlings and those
not inoculated, they were measured from the primary node to the apex.
The results of these measurements are presented in Graph 6 in which the
actual figures are represented by crosses for the non-inoculated seedlings
and circles for the inoculated ones. These figures are based on the meas-
urements of a hundred seedlings for each series. It is immediately obvious
that the non-infected seedlings show a fairly definite curve, whereas the
inoculated ones follow essentially the same curve, although there is some
deviation from the curve presented by the non-inoculated seedlings. This
deviation, it seems, may be explained by the fact that the period of growth
after inoculation was so short that complete mycorrhizal relationship had
not yet become established. Despite this erratic behavior, however, it
is quite obvious that even in so short a time as three months, the fungus
did stimulate the growth of the spruce seedlings.
A further examination of the seedlings shows that while the roots of
the uninoculated ones are not infrequently longer than the inoculated ones,
nevertheless the latter were characterized by the production of a greater
number of secondary rootlets which were frequently branched. In the
eee
FES Se Ag pelt
eo) dh tae le Ne en ois
hs
548 FarLtowia, VoL. 1, 1944
uninoculated plants these secondary rootlets, comparatively speaking, were
very rarely branched. Thus, it can be seen that the fungus in this in-
stance has added to the efficiency of the root system for absorption. This
becomes all the more evident when it is seen that, despite shorter roots
with increased secondary rootlets, the secondary internode of the stem
of the inoculated seedlings, on the basis of ten specimens taken at random,
totaled to 142.5 mm. whereas in the non-inoculated seedlings that were
measured in the same way, the total was 113 mm. Furthermore, the
28
265
24-
Frequency
-
r
q
\y/ peat as
lL 1 L l 1 i js a! _J
ll 12 13 #14 «#215 16 «#17 «#618 «#619 @
Length of internode in millimeters
GrapH 6. The relative rate of growth of inoculated (0) and non-inoculated (+)
seedlings.
inoculated seedlings bore 727 needles and the uninoculated 548 needles,
or respectively 51 and 48.5 needles per cm. of internode.
In addition to these obvious differences between the two series, there is
the further difference that is furnished by the darker color of the needles
of infected seedlings. The needles of non-inoculated seedlings in contrast
to those inoculated were mostly yellowish green and in addition show
the presence of anthocyanins, an indication, according to Mitchell (1934),
that there is a phosphorus deficiency in the soil. Since the inoculated
trees did not show this symptom of deficiency, it seems likely that the
fungal symbiont helped to make available a larger phosphorus supply. Un-
fortunately the information is not available to enable us to make any
statement as to how the process takes place.
GOSSELIN: STUDIES ON POLYSTICTUS CIRCINATUS 949
The obvious differences observed in the size of inoculated and un-
inoculated seedlings, and the greater number of needles per centimeter
of internode for the inoculated than for the uninoculated seedlings, led
us to examine the microscopical characters of rootlets of the inoculated
ones. Rootlets which we considered as having mycorrhizal relationship
on account of their larger size were killed and imbedded in paraffin. It
can be seen from the sections that the mycorrhizal relationship is of the
regular type. The Hartig net is two to three cells deep, without apparent
intracellular invasion. The mantle is homogenous (Plate III, fig. 1)
and somewhat parenchymatous in structure, about three to four cells deep,
with a smooth margin from which arise very few hyphae which penetrate
into the surrounding cells. Sections made of rootlets found in a rootlet
nest (Plate III, fig. 2) under a sporophore of P. circinatus show essen-
tially the same type of mycorrhizal relationship. The mantle is homo-
venous, although thicker, parenchymatous in structure, with a smooth
margin. The tannin layer is not very apparent, and locally seems to be
two cells thick. The Hartig net is not so thick as the type we obtained
by inoculation, but essentially the same type of mycorrhizal relationship
is evident in the artificially and the naturally invaded rootlets. Thus,
although the experiment was not done under aseptic conditions, it seems
very probable that the endophyte here concerned is P. circinatus.
When all characters are taken into consideration, it seems safe to con-
clude that the fungus has a beneficial effect on its phanerogamic partner,
at least in the early stages. Also these experimental results tend to sub-
stantiate the conclusions already drawn from field observations.
The observations that we have made so far, either on growth rates or
on soil or needles analysis, seem to indicate that the infection of the trees
by P. circinatus behaves as if the fungus was mycorrhizal. However, in
our northern forests where the pH of the soil is relatively high and where
nitrogen is chiefly supplied as ammonium salt, it would seem that the
trees must be in mycorrhizal association to live under such conditions.
We have seen that trees infected with P. circinatus behave, in comparison
with trees not infected by this species, much in the same manner as trees
that are in association with known mycorrhizal fungi. This relation be-
tween the fungus and the tree exists until some unfavorable factor, as yet
unknown, upsets the balance between the symbionts. When this happens,
then it would seem that the fungus readily becomes parasitic, and the
mycelium, instead of remaining intercellular, as in the Hartig net, pene-
trates into the cells (Plate III, fig. 3) and travels backward along the
rootlets, killing them as it proceeds. This would then explain the reason
why the butt-rot seems to have its inception in the lateral roots rather
than through wounds or through the tap root. In Polystictus circinatus,
then, we have a butt-rot fungus that in the earlier stages of the develop-
ment of the trees, instead of being a parasite, actually is beneficial until
the host reaches a certain age, which might indicate senescence, the age
being determined by local ecological conditions.
FSAI CBee
990 FarLtowia, Vou. 1, 1944
SUMMARY
The white pocket butt-rot of conifers caused by Polystictus circinatus
(Fr.) Cke. has been studied because of its local importance in the Province
of Quebec and because of its peculiar behavior.
Field observations have been made to correlate the occurrence of this
disease with some ecological factors. The disease was localized in such
definite loci of infection that it was thought that it was tied up with con-
ditions of temperature, inséct epidemics, mechanical damage, or with the
chemical nature of the soil. However, none of those factors appeared to
be responsible for the occurrence of the disease. i
At the same time as the ecological factors were discarded as having
no correlation with the disease, the field observations brought up evidence
that this disease penetrated into its host by mycorrhizal association. Al-
though none of the facts furnished any conclusive evidence, they all sub-
stantiate the same conclusion,
To bring more evidence, the writer has tried to establish mycorrhizal
relationship between spruce seedlings and the fungus. The results indi-
cate that the fungus has really formed a mycorrhizal association of the
usual type.
If P. circinatus penetrates into its host through symbiotic association,
and there is presumptive evidence that such is the case, this fact will open
an entire new field in forest pathology. It will associate the two concepts
of symbiotism and parasitism which were previously separated by air-tight
walls. Those who are specifically interested in symbiotic problems will
have to consider the pathological aspect of the question and the forest
pathologists will have to be better acquainted with symbiotism in order
to deal correctly with certain of their problems. Foresters will no longer
be able to consider the butt-rots only as a reducing factor for the volume
of the tree but will have to determine under which conditions the damage
from parasitism exceeds the beneficial effects of symbiotism on trees, if it
is demonstrated later, as it is not at all illogical to suppose, that butt-
rotting fungi penetrate into the host by mycorrhizal relationship. These
fungi in the Province of Quebec are all quite abundant; before trying to
eliminate them by short cycle rotations, it will be necessary to know what
part they are playing in the biological complex of our forest soil. Future
researches will have to determine under which conditions parasitism
follows the symbiotic association.
This field is new and wide open. It will certainly be of considerable
interest to those who will have the advantage of working on such prob-
lems, and it promises to be at the same time of practical interest to foresters.
GOSSELIN: STUDIES ON POLYSTICTUS CIRCINATUS 551
ACKNOWLEDGMENTS
The writer wishes to thank Dr. David H. Linder, under whose direc-
tion this work was done, for giving his time so spontaneously, for his
imaginative suggestions on possible hypotheses, his objective criticism of
experiments, and his correction of the manuscript. He also wishes to
thank Professor Emeritus J. H. Faull under whom this study was initiated
and who furnished the first knowledge on this disease; and to Dr. Gast
of the Harvard Forest, who has added to the writer’s knowledge of mycor-
rhizal problems, for constructive criticism of this study, and for informa-
tion concerning methods for the chemical analysis of spruce needles.
The writer wishes also to thank the officials of the Department of Lands
and Forests of Quebec for having made possible this study by the financial
help they provided for the academic studies and for the field investiga-
tions; among these officials the writer wishes to thank especially M. A.
Bédard, Deputy-Minister, M. F. Boutin, Chief of Forestry Service, and
Dr. R. Pomerleau, Director of the Bureau of Forest Pathology, under
whom the writer has been working during all this study.
He wishes also to thank Professor Z. Rousseat of Laval Forestry School
for his unfailing interest in this study and for the laboratory facilities he
extended to the writer at the Forestry School. ~
__ The writer is indebted to Dr. Amiot for the help he so kindly gave
during chemical analysis of needles, to M. Y. Garand, Soil Analyst in the
Department of Colonization, for advice on soil analysis, and to M. G.
Bolduc, Forest. Ranger, for his whole-hearted help during field and lab-
oratory work.
The writer is also indebted to Dr. W. H. Gileson, Superintendent of the -
Experimental Farm, Indian Head, Saskatchewan, who so kindly furnished
the prairie soil.
HarvaArD UNIVERSITY
Camepripce, Mass.
552 FartowlA, VoL. 1, 1944
Appendix I |
AREA OF D.B.H. SECTION FOR EACH AGE CLASS
Dara FROM YORK River, GASPE
|
Age Class: I
(40-60) (70-90) (100-120) | (130-150)
Number of trees: |
(1) (6) (13) (19) (25) (24) (2) (4)
A B A 5 A B A B
{i 34 1.27 1.34 0.61 1.09 0.58 1.03 0.21
5.35 4.13 4.48 2.41 3.06 1.61 4.97 0.58
20.37 8.44 7.96 5.48 5.88 2.77 ROL aT I15
49.15 18.24 11.08 9.45 7.98 4.01 13-87 1.61
40.65 16.73 14.29 14.08 9.80 5.62 20.39 2.24
58.67 23.00 17-00. 17°91 11.65 7.34 26.10 2.77
21.40 22.20 13.87 9.10 0,30 3.90
22.20 26.36 16.05 11.08 4.80 4.84
26.36" 18.38 13.04 46.20 5.89
20.89 14.72 53.00 F288
25.20 17.91 31.50 8.61
19.62 Poe 9.81
77.00 11.08
Data From Lake STE. ANNE, GASPE
Age Class:
(40-60) (70-90) cea
Number of trees: \
(7) (3) (23) iQ) (4) (3)
A B A B A B
0.54 0.67 0.39 0.26 0.12 0.39
1.76 1.91 1.47 1.21 0.50 1.34
3.26 Peay § 2.96 2.96 1.03 2.77
5.09 5.88 5.22 | 5.62 1.99 5.35
7.18 7.64 7.64 8.44 2.96 8.44
7.80 10.15 12.43 4.72 12.04
12.84 16.27 6.88 16.27
13.87 18.63 9.28 21.14
19.87 12.24 27.80
14.08 32.80
19.62 42.10
44.30
A—tTrees not infected (see footnote page 537).
B—Trees infected with P. circinatus.
GOSSELIN: STUDIES ON PoLysticTus CIRCINATUS
Appendix I (continued)
DATA FROM THE VALLEY, LAKE WALKER AREA
Age Class:
(100-120) (130-150) (160-180)
Number of trees:
(44) (5) (45) (23) (7) (9)
A B A B A B
0.20 0.32 0.18 0.16 0.23 0.04
0.67 La 0.72 0.58 0.98 0.11
1.40 2.68 1.54 1.34 2.24 0.26
2.50 5.09 2.68 2.32 4.36 0.42
3.90 7.80 4.12 3.68 Tale) Oat
5.48 10.71 5.15 5.36 1k.27- 7106
8.12 13.04 7.80 7.03 16.05 1.47
10.538 15.82 9.98 9.10 21.14 ° 1.99
12.84 20.12 12.64 11.46 26.65 2.59
16.73 20.88 14.93 13.66 32.15 3.386
22.78 17.44 15.82 Sholo yA ie
20.12 18.15 43.21 5.22
23.00 19.87 49.72 6.02
55.45 6.88
60.61 7.64
67.38
Data FROM THE East Store, Lake WALKER AREA
Age Class:
(40-60) (70-90) (100-120) (130-150)
Number of trees:
at) (10) (2) (11) (6) (5) (1)
A Bas A B A B A B
0.42 0.77 0 bbs" 0.18 0.20 0.18 0.20 0.18
2.41 1.99 0.42 0.46 Oot 40). 72 0.62 0.62
3.06 4.12 ieQesiy lL 1.40 1.48: a Eas gee 3) el
5.09 8.12 1.68 1.91 2.0 412.56 2.24 1°76
7.18 TA | aie 8.26 4.12 2.16 2,16
4.24 4.97 4.60 6.30 4.24 4.48
5.62 7.68 5.48 8.44 5.48 6.59
6.88 10.34 eUGs Wy. 7 1 6.88 8.61
7.96 15.82 9.98 138.04 8.61 11.46
12.04 15.16 10.58 13.24
12.64 14.50
15.16 15.82
16.97 17.44
18.63 19.62
A—tTrees not infected.
B—tTrees infected with P. circinatus.
B1s73)
(190-210)
(2) (3)
A B
0.10 0.01
0.29 0.04
0.54 0.06
1.15 0.13
1.99 0.26
3.36 0.39
4.36 0.62
6.02 0.82
7.18 1.09
9.10 1.40
11.65-. «1.76
18.87 ~ 2,93
16.97 2.59
20.87 3.06
23.81 3.46
26.92 3.79
30.25 4.24
33.78 4.36
37.83 4.60
(160-180)
(1) (1)
A B
O48) O48
0.39 0.50
On77) IS
124% 2.24
1764. O79
2.24 8.35
287. wag
3.47 9.63
4.48 12.64
5.35 14.50
6.16 16.73
7.03 19.62
8.61 22.19
9.63 24.65
10.16 26.36
11.46 28.79
504
FarLtowi14, VoL. 1, 1944
Appendix I (concluded)
Dara FROM THE Mountain Top, LAKE WALKER AREA
Age Class:
(70-90)
Number of the trees:
(21) (3)
>
ie
CNORNH OO
EOE Ean] y
DarAWnAI DMO
—
=)
or
on
—_
Age Class:
(40-60)
Number of trees:
(1) (1)
A B
0.6020 0.5104
0.5806 0.3110
0.1982 0.1956
0.1018 0.1016
0.1594 0.13880
GROWTH RATE BY AGE CLASSES
(100-120)
(31) (1)
A B
0.16 0.138
0.58 0.39
1.21 0.50
Dae 2.87
3.26 4.48
4.84 7.03
7.03 10.16
9.09 14.29
1427 17.44
12.74 20.37
15.16 23.81
15.37 25.79
Appendix II
Data FROM YoRK River, Gaspé
(70-90)
(13)
A
0.5246
0.2498
0.1438
0.1104
0.0980
0.0774
0.0158
0.0748
(19)
B
0.5864
0.3574
0.2366
0.1730
0.1044
0.0932
0.0748
(100-120)
(25)
A
0.4484
0.2842
0.0956
0.1260
0.0750
0.0756
0.0634
0.0590
0.0554
0.0818
A—Trees not infected (See footnote page 537).
B—tTrees infected with P. circinatus.
(24)
B
0.4436
0.2354
0.1608
0.1460
0.1160
0.0934
0.0858
0.0706
0.0526
0.0852
0.0396
(130-150)
(13) (1)
A B
0.16 0.20
0.58 0.98
1.09 1.99
1.83 4.12
2.87 6.59
4 24 10.16
5.89 13,24
7.96 15.82
9.98 17.44
12.43 19.62
14.72 21.14
17.20 22.19
19.62 23.81
24.65
(130-150)
(2) (4)
A B
0.6826 0.4520
0.3388 0.2960
0.2072 0.1468
0.1670 0.1422
0.1072 0.0932
0.0646 0.1484
0.0602 0.0940
0.1238 0.0848
0.0592 0.0956
0.0648 0.0694
0.0634 0.0566
0.0342 0.0532
GOSSELIN: STUDIES ON PoLysTicTUS CIRCINATUS
Appendix II (continued)
DaTA FROM LAKE STE. ANNE, GASPE
Age Class:
(40-60) (70-90)
Number of trees:
(7) (3) (23) (9)
A B ay B
0.5142 0.4556 0.5822 0.6716 0
0.2682 0.2718 0.3036 0.3884 0.
0.1938 0.2168 0.2464 0.2780 0.
0.1492 0.1134 0.1654 0.1770 0.
0.0358 0.1238 0.1682 0.
0.1014 0.1168 0.
0.03386 0.0588 0.
0.0180 0.
0
0
Data FROM THE VALLEY, LAKE WALKER AREA
Age Class:
(100-120) (180-150) (160-180)
Number of trees:
(44) (5) (45) (23) (7) (9)
A B A B A B
0.5166 0.5576 0.6020 0.5706 0.6274 0.4682
0.3216 0.3672 0.3328 0.3628 0.3604 0.3522
0.2500 0.2794 0.2398 0.2392 0.2898 0.2128
0.19388 0.1850 0.1878 0.2002 0.2170 0.2014
0.1480 0.1378 0.1442 0.1636 0.1956 0.1878
0.1704 0.0854 0.1324 0.1176 0.1536 0.1544
0.1128 0.0840 0.1072 0.1118 0.1198 0.1310
0.0862 0.1044 0.1024 0.1004 0.1006 0.1140
0.1152 0.0162 0.0726 0.0762 0.0814 0.0888
0.0368 0.0672 0.0638 0.0626 0.1024
0.0622 0.0596 0.0658 0.0620
0.0580 0.0394 0.0610 0.0580
0.0474 0.0454
0.0386
0.0460
A—tTrees not infected.
B—Trees infected with P. circinatus.
A eh eRe OE PEI EC Rt ee ee ER pe Ta
555
(100-120)
(4) (3)
A B
. 63874 0.5408
6168 0.38162
2854 0.2858
1726 0.1980
2024 0.1542
1640 0.13808
1298 0.1138
1200 0.1192
.0610 0.0714
. 1440 0.1084
0.0224
(190-210)
yee)
A B
0.4748 0.4860
0.2724 0.2182
0.3296 0.38204
0.2384 0.2818
0.2278 0.1742
0.1126 0.2096
0.1406 0.1158
0.0774 0.1262
0.1026 0.1100
0.1076 0.0978
0.0756 0.1044
0.0876 0.0632
0.0794 0.0730
0.0678 0.0542
0.0536 0.0386
0.0504 0.0490
0.0478 0.0118
0.0492 0.02382
996 FarLowiA, Vou. 1, 1944.
Appendix II (concluded)
Data FROM THE East SLoPE, LAKE WALKER AREA
Age Class:
(40-60) (70-90) (100-120) (130-150) (160-180)
Number of trees:
(9) (1) (10) (2) (11) (6) (5) (1) (1) (1)
A B A B A B A B A B
0.7574 0.4152 0.4312 0.4082 0.5166 0.6020 0.4862 0.6666 0.4570 0.5682
0.1040 0.3166 0.3892 0.3992 0.3216 0.3128 0.3098 0.2652 0.2980 0.338
0.2214 0.2940 0.2130 0.2208 0.1856 0.2284 0.2444 0.1846 0.2428 0.2860
0.1492 0.2162 0.2454 0.1804 0.2182 0.1500 0.2544 0.1186 0.2290
0.1848 0.1694 0.1592 0.1842 0.1280 0.1516 0.1044 0.1500
0.1220 0.1894 0.0766 0.1268 0.1116 0.1678 0.1178 0.1460
0.0884 0.1292 0.1618 0.1084 0.0968 0.1160 0.0826 0.1092
0.0630 0.1846 0.0984 0.0854 0.0970 0.1244 0.1112 0.1180
0.0814 0.0654. 0.0874 0.0628 0.0774 0.0598
0.0794 0.0394 0.0614 0.0622
0.0790 0.0878 0.0572 0.0690
0.0490 0.0422 0.0878 0.0536
0.0405 0.0512 0.0488 0.0456
0.0234 0.0292
0.0522 0.0282
Data FROM THE Mountain Top, Lake WALKER AREA
Age Class:
(70-90) (100-120) (130-150)
Number of the trees:
(20) (3) (31) (1) (13) (1)
A B A B A B
0.5726 0.4546 0.5706 0.4560 0.5706 0.6800
0.3422 0.4814 0.3194 0.1110 0.2736 0.3098
0.1978 0.1826 0.2826 0.7606 0.2260 0.3166
0.2128 0.1878 0.1476 0.19388 0.1940 0.2034
0.1608 0.1744 0.1718 0.1960 0.1702 0.1882
0.1442 0.0934 ~ 0.1620 0.1600 0.1424 0.1150
0.1390 0.1288 0.1118 0.1480 0.1310 0.0772
0.1892 0.0930 0.0864 0.0984 0.0422
0.0634 0.0676 0.0954 0.0512
0.0654 0.0678 0.07382 0.0326
0.0062 0.0346 0.0676 0.0210
0.0572 0.0306
0.0150
A—Trees not infected.
B—tTrees infected with P. circinatus.
GOSSELIN: STUDIES ON POLYSTICTUS CIRCINATUS
Appendix III
997
CONCENTRATION IN P.P.M. OF PHospHORUS, PorassiuM, CALCIUM AND AMMONIUM
AND H-10N CONCENTRATION OF THE HuMUS OF STANDS OF VARIOUS
DrGREES OF Rorrine
Percentage of
rotted trees in
the stand
Phosphorus Potassium Calcium Ammonium 5) a i
0% 5 aly .(3.82)* 35 (10) 154 (104) 33 (13) 3.83 (0.
5% 6.87 (4.06) 47 (5) 110 (78) 23 (2) 3.58 (0.
10% 6.25 (3.64) 39 (4) 300 (50) 19 (5) 4.22 (0
15% 5.0 40 165 (69) 20 3.70
20% 8.55 (3.33) 35 (5) 272 (78) 21 (4) 4.37 (0
30% 10.00 (3.54) 33 (6) 53 (38) 24 (5) 3.72 (0
50% 5.09 (8.45) 34 (14) 222 (99) 26 (2) 4.20 (0
60% 7.81 (3.98) 40 (8) 111 (67) 27 (10) 3.76 (0
70% 4.38 (0.63) 47 (5) 53 (19) 25 (5) 3.92 (0
80% 8.75 (8.57) 42 (24) 300 (41) 30 (7) 4,27 (0
90% 3.82 (0.82) 33 (12) 93 (67) 26 (8) 3.76 (0
95% 8.75 (4.61) 40 (10) 165 (85) 27 (3) 3.93 (0
100% 2.17 (2.05) 32 (13) 300 (50) 23 (7) 5.38 (1
* Standard deviation.
Appendix IV
PERCENTAGE OF THE Dry WerIGHT oF THE NEEDLES OF PHOSPHORUS,
PoTassIUM AND NITROGEN
Percentage of
rotted trees in
the stand
om oo ooo
Phosphorus
.188% (0.055)*
.146% (0.032)
.143% (0.027)
157% (0.042)
194% (0.017)
184% (0.028)
166% (0.030)
148%
0.148% (0.029)
0.
0.153% (0.028)
182% (0.007)
* Standard deviation.
ooo
ooo O'S ore
Potassium
.3839% (0.70)
.277% (0.053)
.278% (0.060)
.404% (0.082)
.545% (0.091)
. 306% (0.034)
.216% (0.070)
.198%
.232% (0.135)
.264% (0.115)
.294% (0.038)
Nitrogen
29)
06)
.5)
03)
.09)
37)
25)
06)
seis
16)
25)
02)
0.658% (0.062)
0.602% (0.051)
0.664% (0.042)
0.404% (0.082)
0.744%
0.631% (0.062)
0.620% (0.051)
0.626%
0.652% (0.093)
0.661% (0.061)
0.781%
558 Fartowia, Vou. 1, 1944
Appendix V
Tue pH Anp THE ConcENTRATION OF K, NHy, P, CaO In THE SorL OF THE VARIOUS ©
Loca.ities STUDIED
pH Kk NH, P CaO
Locality Inf Non InfNonInfNon Inf Non Inf Non
Gaspé Peninsula
Gaspé Park 4.95 3.81 41 38 20 21 2.75 4.13 290 72
Lake Ste. Anne 3.81 3.79 3135 2319 5.00 3.63 165 131
Chandler 4.24 4.01 87 32 22 28 5.75 4.13 260 216
York River 3.80 3.74 39 35 26 24 4.75 6.50 97 260
Petit Pabos River : 4.60 4.65 438 40 19 25 5.00 12.50 3800 250
West Branch Pabos River 3.81 31 26 3.63 182
Pecan River 4.06 3.90 34 25 25 30 4.88 38.13 250 250
Four Lakes (Marsoui) 3.85 25 25 12.50 40
Petite Riviére-a-Marthe 4.30 3.85 5035 1717 5.00 4.38 250 80
North Side Mt. Albert 6.45 .2.90 -20 50.30 30° 0.75: 3:75 350. -40
Lac des Américains Trail 5.15 4.20 48 40 20 23 1.25 38.75 350 250
East Side Mt. Albert 6.45 3.90 20 50 380 30 0.75 3.75 350 40
Lake Walker Area
Valley 3.74 4.07 20 22 20 22 3.23 12.50 80 40
Mountain Top 4.06 3.75 40 8 18 23 12.50 4.13 80 24
East Slope 8.95 3.938 5 23 15 32 12.50 10.25 40 48
Philibert Slope 3.81 30 18 8.75 56
Appendix VI
Per Cent or Dry WEIGHT OF THE MINERAL ELEMENTS IN THE NEEDLES FROM THE
DIFFERENT LOCALITIES STUDIED
Phosphorus Potassium Nitrogen
Locality Inf Non Inf Non Inf Non
Gaspé Park 0.148% 0.271% 0.765%
Lake Ste. Anne 0.224% 0.175% 0.418% 0.298% 0.671% 0.673%
Chandler 0.169% 0.115% 0.372% 0.344% 0.650% 0.622% ©
York River 0.164% 0.171% 0.236% 0.351% 0.631% 0.634%
General Average 0.164% 0.175% 0.264% 0.317% 0.654% 0.615%
560 FarLtowlaA, Vou. 1, 1944
EXPLANATION OF PLATE I
Fig. 1. Polystictus tomentosus (Fr.) Fr. Side-view to show the distinct stipe which
is nearly central.
Fig. 2. Polystictus circinatus (Fr.) Cke. Side-view to illustrate the sessile, applanate
fruiting bodies characteristic of the species.
_ Fig. 3. Map showing the distribution of P. circinatus. It will be observed that the
species has a predominantly northern distribution.
GOSSELIN: STUDIES ON POLysTICTUS CIRCINATUS
361
3 Map showing the North American distribution of Polystictus circinatus Fr,
@ References from literature; @ Specimens from Farlow Herbarium;
* Places where the author's studies were made.
Priate [
562 FarLowia, VoL. 1, 1944
EXPLANATION OF PLATE II
Shadowgraph taken from seedlings of Picea rubra and reproduced at one-half of
actual size. Specimens on the extreme left of upper and lower row show branching.
Note that the inoculated seedlings (below) have longer internodes above the first
year node, that the needles are more numerous and that the root system is more robust
and for the most part more abundantly provided with secondary rootlets.
GOSSELIN: STUDIES ON POLYSTICTUS CIRCINATUS
PiateE II
564 FarLowia, VoL. 1, 1944
EXPLANATION OF PLATE II
Fig. 1. Cross-section of rootlet of Picea rubra inoculated with P. circinatus, to show
the mantle of pseudoparenchymatous tissue and the Hartig net which sur-
rounds all the cortical cells.
Fig. 2. €ross-section of rootlet of Picea mariana from rootlet nest from which the
sporophore of P. circinatus had arisen. Notice the distinct pseudoparen-
chymatous mantle at the left, and the beginning of the formation of the
Hartig net.
Fig. 3. Cross-section of a small root of Picea mariana which has begun to show root
rot. Note the accumulation of dark tannin globules in most of the cells and
also the faintly showing mycelium that has penetrated into the interior of
the central cells after having passed through the walls.
GOSSELIN: STUDIES ON POLYSTICTUS CIRCINATUS 565
PiateE III
566 FarLowia, VoL. 1, 1944
BIBLIOGRAPHY
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Atkinson, G. F. and H. Schrenk. (1892). Some fungi of Blowing Rock, N. C.
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Bennett, J. L. (1888). Plants of Rhode Island. p. 80.
Bisby, G. R. et al. (1938). Fungi of Manitoba and Saskatchewan. p. 82, 84.
Bourdot, H. and A. Galzin. (1927). Hyménomycetes de France. p. 631.
Boyce, J. S. (1938). Forest Pathology. p. 372, 393, 427.
Bresadola, G. (1920). Selecta mycologica. Ann. Mycol. 18: 67.
Burnham, S. H. and R. A. Latham. (1914). The flora of the town of Southold,
Long Island and Gardiner’s Island. Torreya, 14: 215.
- (1924). The flora of the town of Southold, Long Island and Gardiner’s
Island. Torreya, 24: 29.
Christensen, C. M. (1940). Observation on Polyporus circinatus. Phytopath.
30: 957-963.
Cooke, M. C. (1886). Praecursores ad monographia Polypororum. Grevillea, 14:
77, 82.
Curtis, M. A. (1867). Geological and Natural History Survey of North Carolina.
Part III. Botany. Raleigh, North Carolina Inst. for the Deaf, Dumb and
Blind. p. 97.
Dodge, B. O. (1914). List of fungi of Kewaunee County, Wisconsin, Journ.
Wisconsin Acad. Sci. Arts and Let. 17 (2): 827, 829.
Donk, M. A. (1933). Revisio der Niederlandischen Homobasidiomycetae-aphyl-
lophoracea. II. Amsterdam. p. 1-278.
Ellis, J. B. and B. M. Everhart. (1889). Some new species of hymenomycetous
fungi. Journ. Myc. 5: 28, pl. 8.
Ellis, J. B. (1890). Catalogue of the Plants of New Jersey. p. 483.
Faull, J. H. (1922). Forest pathology, in Rept. Minister Lands and Forests, Ont.
1921: 259-266.
Finn, R. F. (1942). Mycorrhizal inoculation of soil of low fertility. Black Rock
Forest Papers (N. Y.) 1 (19): 116-117.
Fries, E. M. (1821). Systema Mycologicum. 1: 341, 351.
————. (1836-1838). Epicrisis systematis Mycologici, seu synopsis Hymeno-
mycetum. p. 474.
(1848). Fungi natalenses. p. 8.
. (1851). Novae Symbolae Mycologicae, in Acta Reg. Soc. Sci. Upsala,
3 (1): 70-71. 1851. Vol. date 1855.
. (1863). Monographia Hymenomycetum Suecia. 2: 268.
Haddow, W. R. (1941). On the history and diagnosis of Polyporus tomentosus
Fries, Polyporus circinatus Fries and Polyporus dualis Peck. Trans. Brit. Myc.
Soc. 25 (2): 179-190.
Haskell, R. J. and J. I. Wood. (1930). Diseases of plants in the United States
in 1929. Plant Disease Reporter. Suppl. 75: 66.
Hatch, A. B. (1937). The physical basis of mycotrophy in Pinus. Black Rock
Forest Bull. No. 6: 1-68.
Hay, G. U. (1903). New Brunswick Fungi. Bull. Nat. Hist. Soc. New Brunswick
5: 117.
-. (1905). Additions to the plants of New Brunswick. Bull. Nat. Hist. Soc.
New Brunswick, 5 (3): 363.
Hennings, P. (1898) in Engler & Prantl. Die Natirlichen Pflanzenfamilien,
BOR: 167. 1755 10s
Hesler, L. R. (1929). A preliminary report on Polypores of Eastern Tennessee.
Journ. Tenn. Acad. Sci, 4: 9.
eS Toe ee Bee ee ee ee et eee. ee) a A ye
Se ee ey at Se ee
GOSSELIN: STUDIES ON POLysTICTUS CIRCINATUS 567
Hester, J. B., J. M. Blume, and F. A. Shelton. (1937). Rapid chemical tests for
coastal plain soils. Virginia Truck Exp. Sta. Bull. 95: 1433-1486.
Hubert, E. E. (1924). The diagnosis of decay in wood. Journ. Agr. Res. 29:
sol, 567. :
. (1929). A root and butt rot of conifers caused by Polyporus circinatus
Fr. Phytopath. 19: 745-747.
(1931). Outlines of Forest Pathology. p. 351.
Jiead: I. and J. G. Juul. (1938). Ratesopper pa levende naletraer. (Rot fungi
on living conifers). Meddelelser fra det Norske Skogforséksveseen. 6 (3):
300-496.
Karsten, P. A. (1889). Finlands Basidsvampar. p. 326.
Kauffman, C. H. (1911). Unreported Michigan Fungi for 1910. Rept. Mich.
Acad. 13: 230.
(1921). The mycological flora of the higher Rockies of Colorado. Papers
Michigan Acad. 1: 119.
Konrad, P. and A. Maublane. (1926). Icones selectae fungorum. 5: pl. 457.
Lloyd, C. G. (1907). Letter No. 14: 5.
- . (1908). Myc. Notes Polyporoid Issue 1: 2, fig. 195-199,
oo CLO)». Letter Nov 303 I,-2! 6) 11.
——, (1912). Letter No. 38: 6; No. 39: 4; No. 42: 5; No. 43: 4.
———. (1913). Letter No. 44: 1; No. 45: 3; No. 47: 1.
————, (1914). Letter No. 49: 1, 5.
————. (1915). Letter No. 54: 1; No. 58: 1; No. 60: 1, 4, 7.
————. (1916). Letter No. 61: 1, 6; No. 62: 3; No. 63: 2, 8.
———. (1920). Myc. Notes 6: 933, 949, 953, 964, 978, 979, 1014, 1018.
———. (1921). Myc. Notes 6: 1083.
————. (1922). Myc. Notes 7: 1127, 1160.
(1923). Myc. Notes 7: 1187, 1188.
Rompvoat B. O. (1904). ae . |
enna a : ail m5 bd x si a a ie Pe al ly ; ? ;
a a ne % zt ‘ian — nee | Lee : 4 . seas & = |
See eA sith Se
= ieee i Cx +
r ENGES
say oat
oa
fo
1290.
Figures 1-9 .
604, FarLowia, Vou. 1, 1944
2. Pezizella culmigena Sacc., Ann. Myc. 12: 292. 1914. — Sacc., Syll. Fung. 24°:
1189. 1928.
FIGURES 6-9.
Apothecia subgregarious, rather numerous, sessile, attached by a broad
central portion or at least appearing so, free at the margin, yellow, about
0.6-1.1 mm. diam. when dry, not perceptibly larger when moistened,
when dry fairly thin, plane to saucer-shaped, when moistened somewhat
thicker, cushion-shaped; receptacle smooth, when dry turned slightly
off the substratum at the margin, slightly olive-brown toward the base,
pale yellow above, “Chamois” to “Honey Yellow,” on moistening re-
maining nearly the same color, lying closely adjacent the substrate;
margin circular, even, rarely slightly lobed or angular, obtuse, rather
thin, when dry even with the hymenium or slightly elevated, when
moistened directed outward and downward against the substratum, con-
colorous with upper part of receptacle; hymenium when dry plane,
waxy, concolorous with receptacle and margin, becoming slightly convex
on moistening; in section 170-200 y» thick from substrate to hymenial
surface; medullary region very compact, of indeterminate structure, con-
taining a large amount of crystalline material; ectal layer distinct, well
differentiated, about 30 ,« thick at the base, somewhat thinner toward
the margin, composed of compact, more or less angular, thin-walled
cells 9-12 » diam., colorless except for those in the outer layer which
in the basal region of the apothecium are brown and have walls slightly
thickened, those at the margin of the disc clavate; paraphyses simple,
colorless, thin-walled, non-staining, slightly clavate at the apex, 3-4 p
diam.; asei originating from croziers, narrow, clavate, 65-68 x 6-7 p;
ascospores biseriate in the upper part of the ascus, uniseriate below,
cuneate, rounded above, tapering to an obtuse point below, straight or
slightly curved, the sides sometimes slightly irregular in outline, 8.5-10 x
2.42.6 gw, 1-celled, lacking oil globules.
Known only from the type: On decaying culms of Phragmites communis Trin.,
Weisskirchen, Mahren, Bohemia, May 1914, F. Petrak (FH).
Saccardo in his original description indicates that this species is close
to Helotium citrinulum. Superficially the apothecia of the type (Fig. 6)
differ but slightly from those of H. citrinulum (Fig. 1) in being less bright-
ly colored than most specimens of the latter and in being slightly grayish
on the lower portion of the receptacle. Microscopically the species is
distinct from H. citrinulum in its wedge-shaped spores (Fig. 9), long and
narrow asci (Fig. 8) and different structure (Fig. 7). The hyphae of
the medullary excipulum do not stain well, the walls are thin and appear
to become confluent, and the tissue contains a large amount of obscur-
ing mineral matter (Fig. 7). It is not certain whether the structure is
fundamentally prosenchymatous or pseudoparenchymatous. In contrast,
the corresponding region in H. citrinulum (Fig. 2) is rather loose and
open and the hyphae are distinct. Also in Pezizella culmigena (Fig. 7)
the ectal layer is more clearly defined, more rind-like, and more com-
pact than in Helotium citrinulum, and the outer layer of cells in the basal
WHITE: StupIEs In Genus HELotTium. IV 605
portion of the apothecium is brown, whereas in H. citrinulum the entire
tissue is practically colorless. It is possible that the two species are less
closely allied than their superficial resemblances would at first indicate.
3. Helotium midlandensis sp. nov.
FIGURES 10-13.
Apotheciis sparsis vel subgregariis, plerumque solitariis, rarius trinis vel octonis,
stipitatis, flavis vel aurantiacis, 0.3-0.7 mm. in diam. et eaque altitudine; stipite sat
zracili, cylindrico, levi, pallide flavo; disco apiriente depressione minuta citoque ap-
planata, subcarnoso, remanente solido in siccis; receptaculo stipiti concolori, in siccis
minute-ericeo-striolato de margine usque ad apicem stipitis, levi in humidis; hymenio
plano vel patelliformi, rarius leviter convexo, flavo vel aurantiaco; margine levi, obtuso,
circulari vel angulato ex pressione marginum aggregatorum attingentium; paraphysi-
bus, simplicibus, interdum unum ramum prope basim gerentibus, cylindraceis, 3-5 sep-
tatis, 2.43.2 y. in diam.; ascis e “croziers” natis, clavatis, 45-65 x 6.5—-6 .; ascosporis bi-
seriatim vel subbiseriatim dispositis, unicellulatis, plus minusve ovoideis, 7-10x2.6—-3.2 ys.
Apothecia known only from dried specimens, scattered to subgregari-
ous, usually solitary, more rarely in clusters of three to eight, stipitate,
in general aspect yellow or orange-yellow, 0.3-0.7 mm. across the disc
and of about the same height; stipe rather slender, cylindric or essen-
tially so, smooth, pale yellow; dise opening by a minute depression,
very soon plane, subfleshy, retaining its original fullness on drying; re-
ceptacle concolorous with stipe, when dry showing fine, silky super-
ficial striae radiating from top of stipe toward margin, smooth when
moistened; hymenium plane to patelliform, more rarely slightly con-
vex, yellow or orange-yellow; margin smooth. obtuse, circular, or when
crowded becoming angular from mutual pressure; in section: stipe
about 180 » thick, very compact, composed of narrow, parallel hyphae
about 3 » diam. with narrow lumen and thickened, confluent walls, those
near the surface turning outward and ending in a thin indefinite zone
12-18 p» thick and composed of isodiametric cells 3-6 » diam.; medullary
region of disc fairly compact, of thin-walled hyphae 2.5-3.5 p», much
branched and interwoven; ectal layer compact, 40-50 » thick, composed
of narrow, parallel hyphae 3—4.5 » diam., except at or near the surface
where a few are of somewhat larger diameter and the cells shorter;
subhymenium not differentiated from the medullary layer; paraphyses
simple or occasionally once-branched near the base, cylindric, 3—5-septate,
2.4—3.2 » diam.; asci originating from croziers, clavate, 45-65 x 6.5—7 p;
ascospores biseriate or subbiseriate, |-celled, irregularly obovoid, 7-10 x
2.6-3.2 p.
Habitat: One specimen (Iowa, Grumbein) on petioles and larger leaf
veins of old leaves of Quercus sp.; all others on old pods of Gleditsia
triacanthos L,
Distribution and specimens examined: Ohio: Morgan 801, reported incorrectly
(Morgan, Journ. Mycol. 8: 184, 1902; Seym., Host Index, p. 416. 1929) as Helotium
discretum Karst. (NY-E).— Iowa: Homestead, Sept. 26, 1931, G. W. Martin 5186,
type (FH).— Hills, Dec. 3, 1933, G. W. Martin 5191 (FH).— (With Typhula
juncea), North Liberty, Oct. 4, 1941, M. L. Grumbein (FH).— Kansas: Lawrence,
1890, W. C. Stevens 59, incorrectly labeled Helotium herbarum (NY-E).
606 Fartowia, VoL. 1, 1944
This species has much the aspect of Helotium immutabile Fuckel and
other members of the H. epiphyllum group (cf. White, Farlowia 1: 137,
139-147. 1943) to which it belongs. It is distinguished among these
species by its small size, the comparatively slight parenchymatous differ-
entiation of the ectal layer of the disc, small asci, and small egg-shaped
spores. Compare the following species.
4. Helotium erraticum sp. nov.
FicuRES 14-18.
Apotheciis paucis, sparsis vel subgregariis, stipitatis, albis vel cremeo-flavis, usque
ad 2.5 mm. latis et 1 mm. altis, siccando flavescentibus, cartilagineis; stipite crasso,
plus minusve cylindraceo, levissimo, albo-hyalino, lutescente siccando; disco depres-
sione minuta aperto, mox applanato, carnoso, levi, albo, siccando lutescente vel rube-
scente; hymenio albo, plano, siccando lutescente vel rubescente, opaco vel plus minusve
pellucido, cupuliformi; margine levi, obtuso, — Stipite consistente ex hyphis parallelis,
densis, 2.5-5 y. crassis, tenuitunicatis, et ad basin et secundum superficiem expositam
brevioribus et 4-10 y, crassis, membranis crassiusculis praeditis; excipulo ectali disci
indefinito, admodum compacto, ad basin crasso, marginem versus tenuiore, ca. 40-50 p,
crasso in parte media inter basin et marginem; regione medullari minus compacta;
subhymenio vix distincto a medulla; paraphysibus longis, rectis, simplicibus vel
rarissime semel furcatis prope basin, ad apicem clavatis, ascos longitudine paululum
superantibus, 2.8-3.5 . crassis; ascis e “croziers” natis, clavatis, 90-120 x 9-11.5 wu;
ascosporis biseriatim dispositis, subtus autem unam tantum seriem formantibus, ob-
longis vel obovoideis, subcurvatis vel unum ad latus applanatis atque utrinque sub-
attenuatis, interdum plus minusve crepiduliformibus, praecipue cum membrana asci
iam compressae sunt, uni-cellularibus, 11-17 x 3.5-4 y, guttulis olei globulosis con-
spicuis binis apicalibus instructis. Habitatio: Ad folia putridissima arborum
frondosarum.
Apothecia few, scattered or rarely subgregarious, stipitate; when fresh,
white to creamy yellow, up to 2.5 mm. diameter and 1 mm. high, more
often 0.5-1.0 mm. across the disc, and 0.6—0.8 mm. high; in drying, con-
tracting slightly, becoming yellow or dull reddish yellow, “Chamois” to
“Cinnamon-Rufous” or “Orange-Cinnamon,” cartilaginous and of more
or less the same color and consistency throughout; stipe stout, cylin-
dric or essentially so, entirely smooth, hyaline-white when fresh, drying
translucent-yellow; dise opening by a minute depression, soon plane,
fleshy, contracting slightly on drying, more or less saucer-shaped, re-
maining thick; receptacle smooth, when fresh white, drying translucent-
yellow below, similar toward the margin or of a brighter and more
opaque yellow, or sometimes uniformly yellow or reddish; hymenium
when fresh white or nearly so, plane, on drying becoming bright yellow
Fig. 10-13, Helotium midlandensis: 10, Diagrammatic representation of sec-
tion through apothecium, x 110, from type (FH); 11, Section through ectal ex-
cipulum, Martin 5191, « 1290 (FH); 12, Asci and paraphyses, from type, x 1290
(FH); 13, Ascospores from type, x 1290 (FH).
Fig. 14-18, Helotium erraticum: 14, Diagrammatic representation of section
through apothecium, from type, x 78 (FH); 15, Section through ectal excipulum,
from CU-P 27850 (FH); 16, Asci and paraphyses, from CU-P 29658, x 1290 (FH);
17, Ascospores, from CU-P 29658, « 1290 (FH); 18, Ascospores, from TRT /7611,
_< 1290 (FH).
en ea,
>
a |
ca.
WL. White del.
Ficures 10-18
607
608 FartowiA, Vou. 1, 1944
to reddish, opaque or more or less translucent, saucer-shaped; margin
even, obtuse, drying slightly elevated and very slightly undulate or
wrinkled; in section: siipe composed of very compact, parallel hyphae,
2.5-5 pw diameter, thin-walled except at the base and along the exposed
surface where the cells are shorter and thicker, 4-10 » diameter, with
walls slightly thickened and confluent; ectal excitpulum of disc indefinite.
very compact, thick at the base, thinner toward the margin, about 40-50 p
thick half way between base and margin, the hyphae similar to those
of the stipe, the outermost ones similarly somewhat larger in diameter
and with walls slightly thickened, forming a thin rind layer extending
nearly to the margin; medullary region slightly less compact, of thin-
walled hyphae 2.5—-5 » diam., much branched and interwoven in all
directions; swbhymenium scarcely differentiated from the medullary re-
gion; paraphyses long, straight, unbranched, or very rarely once-branched
close to the base, clavate at the apex, extending somewhat beyond the
asci, 2.8-3.5 » diam.; asei originating from croziers, clavate, 90-120 x
9-11.5 4; ascospores biseriate above, uniseriate below, oblong or obo-
void, slightly curved or flattened on one side, slightly pointed toward the
ends, sometimes appearing more or less slipper-shaped especially when
crowded in the ascus, 1-celled, 11-17 x 3.5—4 p», with a conspicuous and
rather large oil globule in each end.
Habitat: On much decayed leaves of various frondose trees (as listed
below) ; one collection on old pods of Robinia Pseudoacacia L.
Distribution and specimens examined: Quebee: On Acer sp., Populus sp., & sp.
indet., Duchesnay, Aug. 25, 1938, H. H. Whetzel & T. Sproston, previously reported
(Mycologia 31: 730. 1939) as Helotium phyllophilum and (White, Farlowia 1: 141,
159. 1943) as H. epiphyllum (CU-P 27850, FH).— Duchesnay, Aug. 26, 1938, R. F.
Cain 11154, reported (Mycologia, l.c.) as H. immutabile and (White, ].c., p. 141,
147) as H. epiphyllum, spec. comm. H. S. Jackson (FH).— New York: On Acer sp.,
Coy Glen, near Ithaca, Oct. 3, 1938, H. H. Whetzel & W. L. White 3416 (FH).— On
Acer sp. ?, Coy Glen, Oct. 3, 1938, Whetzel & White 3421, type (FH).— On Amel-
anchier sp., Hamamelis sp., & Ulmus sp., Malloryville, near Ithaca, Oct. 18, 1941,
H. H. Whetzel & J. Niederhauser (CU-P 29658, FH).— On pods of Robinia Pseudo-
acacia L., Newfield Gorge, near Ithaca, Oct. 26, 1941, H. H. Whetzel & T. Sproston
(CU-P 29666, FH).— Ontario: On Acer sp., Brewer Lake, Algonquin Park, Sept.
5, 1939, R. F. Cain, comm. Jackson, TRT 17611 (FH).
This is another member of the Helotium epiphyllum group and must
be cautiously separated from such species as H. epiphyllum, H. immuta-
bile, H. carpinicola, and H. midlandensis. These species appear to be
distinct and taxonomically satisfactory, but may be somewhat difficult
for one who has had no experience with them. They cannot be sepa-
rated with any degree of certainty macroscopically, but under the micro-
scope are distinct on a number of characters, the final and most impor-
tant being spore size and shape.
5. Helotium albidum (Rob. ex Desm.) Pat., Tab. Anal. Fung. 4: 173, fig. 382a-d..
1885.
Peziza albida Rob. ex Desm., Ann. Sci. Nat. 3: 16: 323. 1851.
Helotium scutula (Pers. ex Fr.) Karst. var. albidum (Rob. ex Desm.) Karst.,
Myc. Fenn. I, Bidr. Kann. Finl. Nat. Folk 19: 112. 1871.
WHITE: StTupIEs IN GENUS HELoTIUM. IV 609
Phialea albida (Rob. ex Desm.) Gill., Champ. France, Discom., -p. 105. 1879-1883.
Hymenoscypha albida (Rob. ex Desm.) Phill., Man. Brit. Discom., p. 138. 1887.
FIGURES 19-24.
Apothecia arising from black areas l-several cm. long on the leaf
petiole, 6-20 on each such area, scattered to subgregarious, stipitate, in
the dried condition up to about 1.2 mm. high and 1.5 mm. across the
disc, the larger ones expanding to 2 mm. when moistened, in the fresh
condition creamy white or ivory white (Desm.) or with maximum mois-
ture content more dilute in color, drying with stipe and outside of disc
essentially concolorous, “Cinnamon” or “Pinkish Cinnamon,” and the
hymenium “Orange-Cinnamon”; stipe cylindric, slender but not deli-
cate, minutely scabrous when dry; disc subfleshy, spreading, saucer-
shaped when dry; receptacle when dry finely and closely lined with
superficial radiating striae; margin even, when dry conspicuously ele-
vated above the hymenium; hymenium drying firmly waxy; in section:
stipe composed of a broad central core, hyaline, of thin-walled hyphae
about 4-6 » diam., and a thin, yellowish, cortical layer 4-5 cells thick
and composed of short rectangular cells; medullary region of disc com-
pact, composed of hyphae similar to those making up the central core
of the stipe; hypothecium not noticeably differentiated; ectal layer not
sharply distinguished, consisting of a zone about five cells thick made
up of large hyphae 8-10 » or rarely up to 15 w diam., over which is a
thin external covering of narrow more or less crushed or collapsed hyphae
of smaller diameter; paraphyses simple, usually 3-septate, slightly en-
larged toward the apex, about 3 » diam.; asei clavate-cylindric, not
originating from croziers, 90-100 x 10-12 »; ascospores uniseriate
below, biseriate above, 1-celled, 13-17 x 4—5 p, rounded above, very
slightly tapering toward the lower end, slightly curved or flattened on
one side, obtuse-pointed below, the content granular.
On fallen, decaying petioles of Fraxinus excelsior in France. Known with certainty
only from the material of Desmaziéres: Pl. Crypt. Fr., ed. 1, 2004; ed. 2, 1604 (FH).
This is a member of the Helotium scutula complex in which many
forms, varieties and species have been described. It is being allowed to
stand as distinct from A. scutula or H. caudatum, the former on herba-
ceous stems and the latter on leaves. It is more robust than H. caudatum
and differs from both it and H. scutula in having spores which are
smaller in size and more obtuse at the lower end. If it proves to be
confined to Fraxinus petioles where it causes blackening, and if spore
and other characters are constant, then it may be maintained as a species.
The references listed above are only those pertinent to the synonymy.
There are many European records, but they are based largely upon
spcimens better referred to H. caudatum or H. scutula.
6. Helotium scutula (Pers. ex Fr.) Karst. var. fueatum (Phill.) in Rehm in Rabenh.
Krypt.-Fl. 1°: 793. 1893. — Massee, Brit. Fung.-Fl. 4: 254, 1895. — Boud., Hist.
Classif. Discom. Europe, p. 114. 1907.
Peziza fucata Cooke & Phill.; Cooke, nom. nud., Grevillea 4: 132, pl. 65, fig. 300.
1876.
610 FarLowl1A, Vou. 1, 1944
Hymenoscypha scutula (Pers. ex Fr.) Phill. var. fucata Phill., Man. Brit. Discom.,
p. 137. 1887.
Phialea scutula (Pers. ex Fr.) Gill. var. fucata (Phill.) Sacc., Syll. Fung. 8:
266. 1889.— Seaver, Bull. Lab. Nat. Hist. State Univ. Iowa 6°: 101. [Oct.
1911], spec. & descr. excluded. — Oud., Enum. Syst. Fung. 2: 1004, 1920. —
Seym., Host Index, p. 292. 1929, based on Seaver’s report (l.c.).
FIGURES 25-30.
Apothecia scattered to subgregarious, fairly numerous, stipitate,
hyaline-white when fresh, or sometimes yellowish-stained, 1-3 mm. high,
up to 2.8 mm. across the disc though usually smaller, drying yellowish
ochraceous, all parts approximately concolorous in both fresh and dried
conditions, the disc contracting somewhat in drying, usually not over
1 mm. diam. in herbarium specimens; stipe slender, usually about
equalling the diameter of the disc, originating underneath the epidermis
of the substratum, scarcely noticeably erumpent, smooth, usually more
or less broadened at juncture with disc; dise at first subglobose, open-
ing by a pore, finally expanding, more or less saucer-shaped, of medium
thickness; receptacle smooth when fresh, very minutely radiately fur-
rowed when dry; hymenium becoming plane at maturity and with
maximum moisture content, drying slightly to pronouncedly concave;
margin thick, obtuse; in section: stipe with a rind about 30 » thick
or about 5-6 cells thick, composed of longitudinally parallel hyphae
4-8 » diam. divided into rectangular cells 13-19 p» long with walls
slightly thickened and confluent, this rind enclosing a large central core
of parallel, compact hyphae 2.5-3.5 » diam. with walls slightly thick-
ened, divided into cells 25-50 » long; disc composed of an ectal layer
of hyphae similar to those of the stipe and a relatively reduced medul-
lary region of more loosely interwoven hyphae; ectal layer about 50 p
thick at the base, becoming thinner toward the margin, consisting of
an outer zone similar to that of the stipe and a broader inner zone of
radiately parallel hyphae similar to those of the central part of the stipe;
paraphyses simple or frequently once or twice branched at or more often
below the middle, usually near the base, 3—5-septate, slightly clavate
at the apex and there 3-3.5 » diam.; asei originating from croziers
which break rather easily, clavate, 118-135 x 12-15 »; ascospores
biseriate, 1l-celled, obtuse and rounded above, flattened on one side or
slightly curved, narrowed downward, acute at the lower end, 24-34 x
5—6.8 , each end beset with 1 to several small, inconspicuous cilium-
like processes, content with numerous conspicuous refractive granules.
Known from only three collections: New York: On old stems of Polygonum
robustius (Small) Fernald (det. Fernald) lying in film of water in swamp, Cayuta
Fig. 19-24, Helotium albidum, all from type material, Desm. Pl. Crypt. Fr.,
Ed. I, 2004 (FH): 19, Diagrammatic representation of section through an apothecium,
< 40; 20, Section showing blackening of the exposed cell walls over the area of the
petiole on which the apothecia are seated, the epidermis of the petiole having earlier
sloughed off, « 1290; 21, Section showing structure of outermost tissue of stipe,
exposed surface at left, taken from near but not at the base, x 1290; 22, Section
showing structure of disc tissue, < 1290; 23, Asci and paraphyses, x 1290; 24,
Ascospores, X 1290.
wre, Reape Ne ae ts
W.L. White del.
Ficures 19-24
FarLowiaA, VoL. 1, 1944
612
Ficures 25-30
WHITE: STUDIES IN GENUS HELoTIUM. IV 613
Lake, Sept. 1, 1936, H. H. Whetzel & W. L. White (CU-P 25494, FH, US).— Same
substrate and locality, Sept. 9, 1938, Whetzel, White, et al (CU-P 27863, FH, US).—
England: “On dead stems of Polygonum lying in water.”, Shrewsbury, Phill. Elv.
Brit. 120, type (CU-PD 11122, FH).
It would be scarcely possible for three collections to agree more com-
pletely than do those cited above. Although the substratum for the
British specimen is insufficient for complete identification, there is no
doubt about its being Polygonum, and the pieces of stem in the packets
agree so closely with those of the American collections that one is
tempted to guess that they represent a closely allied species. The evi-
dence is that this fungus does exist as a distinct entity within the taxo-
nomically difficult Helotium scutula complex of which it is a member.
Its distinguishing characteristics are the presence of croziers at the
bases of the asci, relatively large asci, large and robust spores, and
occurrence on decaying, water-saturated stems of Polygonum. Though
this variety may prove to be confined to Polygonum, this does not mean,
however, that all members of the Helotium scutula group found on
Polygonum should be referred here. On the contrary several collec-
tions are at hand where the substratum is Polygonum and the fungi
are typical Helotium scutula. A specimen on Polygonum from Iowa
(CU-PD 553), reported by Seaver under the varietal name, represents
typical material of the species.
Cultures were made from both of the American collections, using mass
ascospore discharge on potato dextrose agar. The spores germinated
readily, becoming 1-3 septate in the process, and developing a mat of
sterile mycelium. The cultures were maintained only in test tubes at room
temperature, but there developed no conidial stage, spermatia, stromata
or other. structures of taxonomic value.
7. Helotium Dearnessii (Ell. & Ev.) White, Mycologia 34: 167. 1942,
Peziza (Phialea) Dearnessii Ell. & Ev. nom. nud., N. Am. Fungi, second series
2624. 1891.
Phialea Dearnessii Ell. & Ev., Proc. Acad. Nat. Sci. Philadelphia 1893: 146.
[Feb. 28] 1893. — Sacc., Syll. Fung. 11: 403. 1895. — Seym., Host Index, p. 558,
(559 from error in host det.). 1929. — J. H. Miller, Pl. Dis. Rep. Suppl. 131:
47, Aug. 15, 1941, based on an incorrect determination.
Hymenoscypha Dearnessii (Ell. & Ev.) Kuntze, Rev. Gen. Pl. 37: 485. 1898.
FIGURES 3]—34.
Apothecia scattered to subgregarious, not crowded, appearing sessile
or nearly so, in the dried condition ochraceous, all parts concolorous
0.3-0.7 mm. diam. across the disc, when moistened becoming more yel-
Fig. 25-30, Helotium scutula var. fucata, all except fig. 30 from Phill. Elv.
Brit. 120, type (FH): 25, Diagrammatic representation of section through apothecium,
< 76; 26, Section showing structure of outermost tissue of stipe, x 1290; 27,
Section showing structure of outermost tissue of disc, X 1290; 28, Asci and para-
physes, x 1290; 29, Ascospores, 1290; 30, Ascospores germinated on potato
dextrose agar, slightly more than x 1290 (CU-PD 25494).
614 Fartowia, Vor. 1, 1944 —
lowish the largest reaching as much as 1.0 mm.; stipe obscure, scarcely
long enough to raise the central part of the disc off the substratum,
rarely longer, ochraceous, rarely darker at the base; dise rather fleshy,
saucer-shaped when dry, becoming cushion-shaped when moistened; re-
ceptacle smooth; hymenium opening by a minute depression, soon
plane, remaining plane or becoming somewhat concave on drying, con-
vex when moistened; margin even, slightly elevated when dry, when
moistened expanded and recurved, wedge-shaped; in section: stipe
originating underneath the cortex of the host, erumpent, cylindric,
usually protruding only slightly beyond the cortex, very compact,
homogeneous; ectal layer of disc thick, indefinite, in general composed
of an inner layer originating at the top of the stipe, about 40 » thick at
the base, tapering towards the margin, colorless, composed of compact
parallel hyphae about 3 » diam. with walls scarcely or not at all thick-
ened, these hyphae turning obliquely outward to form a thicker outer
brownish zone and there becoming somewhat larger, shorter-celled, with
walls thickened and confluent; medullary region thin, rather loose, the
hyphae rather narrow, thin-walled, interwoven, becoming erect towards
the hymenium; hypothecium not differentiated from medullary layer;
paraphyses mostly simple, rarely once-branched below the middle,
gradually enlarged from base to apex, about 3-3.5 » diam. at apex;
asci not originating from croziers, clavate, 100-108 x 9-13 ,; asco-
spores biseriate, elongate, pointed at the ends, attenuate below, 1-celled,
containing a row of 2-9 oil globules, with a delicate cilium 2.5-3.5 p
long at the lower end, sometimes also at the upper end, 30-35 x 44.5
not including the cilia.
Known only from the following collections: Quebec: On old herbaceous dicotyle-
donous stems, Aug. 25, 1938, J. VW. Groves (FH, OTB).— Ontario: On old stems of
Steironema ciliatum (L.) Raf., London, May [24], 1890, J. Dearness 1713, type
(FH-E).— Additional material from same place, June [13], 1890, J. Dearness 1713,
Ell. & Ev. N. Am. Fungi, 2nd ser. 2624 (FH).
According to a letter from Dearness to Farlow, he first collected the
species May 24, 1890 and recorded the substrate as Monarda didyma.
Ellis described it as on Monarda and it was so recorded in Seymour’s
Host Index. Dearness apparently returned to the same place June 13,
1890, collected additional material which he sent Ellis for his N. Am.
Fungi, and decided that the host was Steironema ciliatum. A very ample
portion of the June 13th material, in addition to the portion in N. Am.
Fungi, was sent to Farlow and is now in the Farlow Herbarium.
Fig. 31-34, Helotium Dearnessii, all from Dearness, 1713, coll. June 1890 (FH) ;
31, Diagrammatic representation of section through apothecium, x 64; 32, Section
showing details of structure of ectal excipulum, « 1290; 33, Asci and paraphyses,
> 1290; 34, Ascospores, < 1290.
Fig. 35-40, Helotium gemmarun, all from CU-P 251781 (FH): 35, Diagram-
matic representation of section through an apothecium, * 60; 36, Section showing
structure of outer layer of stipe, < 1290; 37, Section showing structure of medullary
and ectal layers of disc, x 1290; 38, Hyphal tips from margin of disc, « 1290; 39,
Asci and paraphyses, * 1290; 40, Ascospores, X 1290.
FIGuRES
Barr ey ieee
WL. White del.
a |
31-40
616 FarLtowiA, VoL. 1, 1944
The collection recorded by Miller (1. c.) from Georgia on Stetronema
ciliatum, of which he kindly loaned me a fragment, proves to be the
much more common and widespread species, Helotium scutula (Pers.
ex Fr.) Karst., to which H. Dearnessii is surely closely related but clearly
distinct in its long, attenuate spores.
The two specimens of Helotium Dearnessii recorded above agree closely
in their characters except that in the Ontario material the apothecia are
- all so short-stipitate that on superficial examination they appear sessile,
whereas in the Quebec specimen most of them are distinctly stipitate;
also the cilia, which are difficult to see in the former are rather distinct
in the latter.
6. Helotium gemmarum Boud., Bull. Soc. Myc. France 4: 81, pl. 17, fig. 2. 1888. —
Boud., Rev. Myc. 11: 167. 1889.— Boud., Icones Mycol. 3: pl. 493. 1905-
1910. — Boud., Hist. Classif. Discom. Europe, p. 112. 1907.
Phialea gemmarum (Boud.) Sacc., Syll. Fung. 8: 271. 1889. — Oud., Enum. Syst.
Fung. 2: 61. 1920. — J. H. Miller, Pl. Dis. Rep. Suppl. 131: 47, 1941.
FIGURES 35-40.
Apothecia scattered to subgregarious, small, delicate, slender, stipitate,
white, in the fresh condition up to 1.5 mm. high (up to 5.0 mm. fide
Boud.) and 0.4-1.4 mm. across the disc; stipe slender, slightly en-
larged upward, when fresh hyaline to dilute white, smooth at the base,
minutely puberulent above, not changing perceptibly on drying; dise
opening by a pore, at first infundibuliform, at length expanding, finally
plane, thin, on drying quickly contracting to become more or less funnel-
shaped with margin turned inward; receptacle minutely and sparsely
puberulent, dilute white when fresh, remaining so on drying or vary-
ing towards dingy cream-color; hymenium always about same color as
receptacle, flat at full maturity and with maximum moisture content,
otherwise more or less deeply concave; margin minutely puberulent,
even with the hymenium in fresh, mature specimens, erect or turned in-
ward in dried material; paraphyses not numerous, simple or more
rarely once-branched below the middle, not enlarged at the apex; asci
small, cylindric, clavate, 45-50 x 5-7 »; ascospores small, obpiriform,
6-9 x 2.5-3 py.
Habitat: Appearing in early spring on the scales of buds of various
species of Populus (listed below) which had fallen the previous season
and lodged under and among debris on wet ground.
Distribution and specimens examined: New York: On P. candicans Ait., Lloyd
Preserve, McLean, Apr. 26, 1936, H. H. Whetzel, D. H. Linder, & W. L. White (CU-P
25181, FH).— On P. candicans, Lloyd Preserve, McLean, May 2, 1937, W. L. White
et al (FH). — On P. candicans, Lloyd Preserve, McLean, May 18, 1940, H. H. Whetzel,
W. L. White, & J. Niederhauser (CU-P 29086, FH).— Pennsylvania: On Populus
sp. ?, May 5, 1935, H. H. Whetzel (FH).— Georgia: On P. deltoides Marsh., Agri-
cultural Campus, Athens, Apr. 29, 1940, J. H. Miller (GA 583).— France: Type
locality. Boudier indicated that it was frequent on the old bud scales of Populus
nigra L. in the vicinity of Paris in February. Notes and drawings by Massee, made
from Boudier material, are in the New York Botanical Garden but are not accompanied
by a specimen.
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{ Ree Ln als 3 » a — 5 lial
WHITE: Stupies 1n Genus HELotium. IV 617
There appears to be no European material of this species available
in the herbaria of this country. However, it is well marked as to
morphological characters, substratum, and season of occurrence. Com-
parison of North American specimens with the illustrations of Boudier
leaves no reason for doubt as to its identity. Two other species with
similar characteristics — white, slender, with small spores, occurring in
the spring in moist habitats — and possibly more or less closely related,
are Phialea Cassandrae Kanouse, occurring in Michigan on the leaves of
Chamaedaphne calyculata (L.) Moench (see White, Farlowia 1: 159.
1943), and Helotium amenti (Batsch ex Fr.) Fuckel (see Boudier, Icon.
Mycol., pl. 496. 1905-10), of widespread occurrence on female catkins
of Salix sp.
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1(4): 619-624 FARLOWIA July, 1944
REVIEWS
UEBER DIE BEDINGUNGEN DER MYKORRHIZABILDUNG BEI KIEFER UND
FICHTE. By Erik Bjorkman. Uppsala. Symbolae Botanicae Upsaliensis.
Vol. VI, no. 2. 1942,
When B. Frank (1885) likened the common fungus root of a tree to a factory op-
erating for its own profit, he was not unmindful that the tangled and swollen root
tips of a tree with their covering mantle of fungous skin, and their cakes of mycelia
between the cells, look like witches brooms or other gall formations caused by a
parasitic fungus. He considered that the fungus obtained its food from the host
tree and in return provided the tree with water and nutrients for growth, which the
fungus extracted from the soil through its widespread mycelial threads. This inter-
pretation did justice to what one can see, and itis supported by much else: many
nutrients are deficient in the raw forest humus (mar) (Stahl 1900); the fungus root
is built like an organ for the exchange of substances (Mangin 1910); Neger (1913)
found more sugar in roots from trees which habitually have fungus roots than in
the case of ash and many other plants; Rexhausen (1920) found plenty of fungous
starch (glycogen) in the mycelium of root fungi and more sugar or starch in in-
fected roots than in other roots of the same tree; a number of large fungi of the
forest known to form mycorrhiza (Melin 1922-25 et al) follow their host trees as a
“dolphin follows a ship.” Lange (1923), also Thesleff (1919), Romell (1921) and
Melin (1925) did not admit that the mycorrhizal agency can consume cellulose or
the more common energy-giving foods, except sugar; mycorrhizae can be abundant
in almost completely humus-free soil, such as volcanic ash (McDougall and Jacob
1927); soluble dyestuffs can penetrate the fungus membrane and mycorrhizal net
right into the cells of the host tree (Endrigkeit 1937); conifer seedlings without or
almost without mycorrhizae have shown a lower phosphorus content and most of
them also a lower nitrogen and calcium content than plants with abundant mycorrhizae
grown in the same soil (Hatch et al 1937).
Frank soon abandoned his first explanation of fungus-infected tree roots. Led
astray by the peculiar pine vegetation and by some experiments with tree seedlings
which he could not explain correctly, he concluded that the fungus gets no nourish-
ment from the root, but like other soil fungi lives on humus, and only lives in the
mycorrhizal covering in return for an ample rental, especially of nitrogenous sub-
stances. This interpretation was adopted more or less faithfully by most of those
who wished to believe that mycorrhizae are useful to the tree. Fungi came to appear
as inseparable thralls of the host plant, and one cared little about what pleasure
they could get from being tenants in the roots. Nor did Stahl care either, although
his explanation of fungus-infected roots of vegetables, after another manner, came
near to the old theory of Frank. Quite generally one has hesitated to believe that
fungi that do not penetrate into the root cells and there become digested, can do
the tree any other service than common soil fungi do by forming useful substances
or consuming harmful ones. Some have wished to attribute the benefits of root fungi
to this (Tubeuf 1896, Rayner 1934, Burges 1936, and Lindquist 1939) but it leaves
the formation of mycorrhizae unexplained, as Endrigkeit (1937) and Hatch (1937)
have pointed out. Others have not been able to escape the thought that the tree’s
mycorrhizal builders live on their hosts. It used to awaken doubt that the mycorrhiza
can be of any use to the tree, since biologists have become increasingly unwilling to
conceive of any peaceful exchange to mutual advantage outside the realm of human
beings. Frank’s Danish pupil Sarauw (1893) and Stahl’s pupil Weyland (1912)
could believe no more than R. Hartig, Ramann McDougall (up to 1927) and others,
that the mycorrhizal partner of trees is anything more than a harmless sponger.
Melin tried in 1925 to unite the current opinions by sifting out differences between |
other kinds of mycorrhizae and the common tree mycorrhizae. He had to class fungi
as parasites or saprophytes, but showed that they invade root cells more frequently
619
rrre rane
620 FarLtowliA, Vou. 1, 1944
than generally appears, and there they are consumed. Thereby the trees, in exchange
for growth substances and carbohydrates, receive nitrogenous foods rich in carbon
which the fungi have formed from substances in the raw humus that are resistant
to decomposition. Thus neither of the partners needs to get the short end of the
bargain, even if one calculates the calories exchanged. The theory failed of proof,
for, with culture in mir, conifer seedlings with their mycorrhizae showed that they
required ordinary nitrogenous nutrients; they could therefore not live on the nitro-
genous staples of the mir (Hesselman and Melin 1927). In addition, Hatch (1937)
found that seedlings get along just as well without fungi, with both the “more com-
plicated” nitrogenous compounds (peptone and nucleic acid) which Melin had
utilized in the experiment on which he had based his conclusions.
Hatch in 1937 forgot what Melin’s experiment with different carbon sources had
shown, and suggested that the mycorrhiza-forming fungi were consumed with small
amounts of growth substances, “and perhaps sometimes carbohydrates”. Endrigkeit
the same year expressed the opinion that the mycelial net in the fully-developed
mycorrhiza cannot obtain any nourishment from the host plant. The reviewer, on
the other hand (Romell 1939) held that there was an exchange of substances ac-
cording to Frank’s first belief, and supported this hypothesis with new reasons: a
fungus does active duty in place of roots and root hairs and does not need to draw
on the plant more than the ordinary root system would do.
In order to be able to extract nutrients for growth from the soil as efficiently as
possible in competition with all other fungi, the mycorrhizal fungi ought to have
some kind of monopoly on easily digested energy-yielding food. Otherwise there seems
to be no one since Rexhausen (1920) who has given Frank’s 1885 hypothesis any
clear support.
But here comes a young Swedish investigator and rejuvenates in new and im-
proved form the very oldest theory for tree mycorrhiza as a useful structure. He does
this by solving a riddle which has baffled investigators the world over during a gen-
eration. The mystery for which Bjérkman has sought the solution is the peculiar
variation in abundance of mycorrhizae. They are commonly found abundantly in
moderately good and better mar humus, but sparingly in the most infertile mar and
often also in mull, where they sometimes are absent.
Recently only Hatch (1937) seems to have interpreted the fact, as far as it now
goes, after the classical manner, namely that mycorrhizae are formed when the trees
need them because they do not have enough mineral nutrients. This is no factual
proof; Melin (1922-27) tried to obtain one and suggested that root fungi did not
thrive in pronounced, but only in slightly acid mull (an unsupported supposition)
and had insufficient “virulence” in bad mar. The word virulence is borrowed from
medical science, literally meaning poisonousness, and connotes capacity to attack.
But this power to attack is supposed to be explained by the word virulence. Of such
a contorted phraseology Heribert-Nilsson says: “Such an explanation is as scant
an aid as if a physician should stand up and declare that the cause of infantile
paralysis lies in the susceptibility of man for infantile paralysis.” The word virulence
is yet not altogether empty of meaning. Back of it lies the thought that the tree
root and fungus are in a feud with each other. But the thought carried at once
difficulties. Melin (1922) first supposed, as was natural, and as later McDougall did,
that the fungus must be more virulent in order to penetrate the cells than to grow
between them. The next year he had to contradict this and suppose something as
strange as that a less virulent fungus goes between the cells, but a more virulent
one can be forced by the host plant to hold itself in the cell walls. Melin’s specu-
lative fabric has nevertheless appeared to be the best to be had, and M. C. Rayner
has gratefully elaborated it further.
Bjérkman sought first a foundation by collecting statistics on the abundance of
mycorrhizae in different forest types. This yielded him as little as his less diligent
predecessors had found, as far as providing a key or even any clue, and the fifty
pages which deal with this serve best some academic requirement. Inoculated pure
REVIEWS 621
cultures repaid his efforts better, but still the work would have had almost the same
value without them. The scientific importance of the study lies partly in the
ecologically planned experiments where uninoculated pine or spruce seedlings were
grown in the same soil (rich mull, better and poorer mar and peat) with their natural
content of soil life, or at least of sources of life, and partly in the theory to which
the experiments led. It shows that Bjoérkman’s discovery is not a result of modern
technique and the work of mycorrhiza investigators of recent times, but could have
been made fifty years ago if someone then had been equally industrious and had had
the vital spark.
Six soils were tested in the experiment, some without additions, others under
different degrees of illumination up to four soils with different additions of nutrients,
and three soils under varying light and additions of nitrogen and phosphorus fer-
tilizers. -Mycorrhizae were formed abundantly even in rich beech forest mull, after
a moderate mixture of sawdust which decreased the nitrogenous food content. It is,
therefore, not as Melin believed, that no mycorrhizae thrive in well-developed mull.
Mycorrhizae are formed abundantly also in the soils showing the highest nitrogen
content (far higher than are found in nature in the richest mull) if only phosphorus
is low and light is strong enough. Lindquist (1932) also was mistaken when he
thought that nitrate formation is to be looked upon as a “change in dominance” in the
fungous flora of the soil of such a kind that mycorrhizal formation is thereby retarded.
During the entire course of the experiments, three things were decisive: light, and
the nitrogen and phosphorus contents of the soil. Thus mycorrhizae were formed in
weak light (1/16 or sometimes 1% of full sunlight). With increased light there were
steadily better seedlings with more mycorrhizae. On the contrary there were most
mycorrhizae (but not the best seedlings) with moderate deficiency of nitrogen and
phosphorus or both. Greater deficiency of nitrogen resulted in weak seedlings with
few or no mycorrhizae. The same results were obtained in inoculated pure cultures
when there was great deficiency of phosphorus. In soils richly fertilized with nitrogen
and phosphorus the plants grew strongest, but had no mycorrhizae. The interaction
between light and contents of nitrogen and phosphorus led to intricate correlations
which are shown clearly in the report by means of three-dimensional curves.
The action of light was known before, as a result of another study by Bjérkman
(1940) together with Hesselman. The per cent of mycorrhizae in connection with
light gives curves which appear logarithmic and call to mind Stalfelt’s photosynthesis
curves for pine and spruce. Bjorkman guessed even in 1940 that there might be a
connection between assimilation and mycorrhizal formation. This Melin (1927)
had rejected as a possible cause for the stronger plants in Hesselman’s experiment
having more abundant mycorrhizae; but he did not find the interpretation worth
discussing seriously.
Out of the guesses of 1940 eventually resulted the following: the mycorrhizae seek
soluble carbohydrates in the root, and they consume them for energy, but they find
them only if there is a surplus of carbohydrate in the root. Whether or not there
is any surplus depends on the head-start photosynthesis (carbohydrate formation)
has attained in the plant before the formation of albuminous substances. This ad-
vantage will be small or nil if there is an abundance of all plant foods including
nitrogen and phosphorus, so- that there will not need to be a shortage of materials
when albuminous substances are built up. Then there will be little for the fungi to
seek after in the root, and mycorrhizae will be formed sparingly or not at all. On
the other hand, should there be a deficiency of nitrogen or phosphorus foods, there
can easily arise a surplus of carbohydrates so that the formation of albuminous sub-
stances falls behind photosynthesis. With a moderate lack of nitrogen or phosphorus
or both, mycorrhizae can form abundantly, but if there is a serious lack of nitrogen
or phosphorus, sooner or later photosynthesis will become weak also (phosphorus
especially strongly influences carbohydrate formation in plants) so that the surplus
of assimilated carbohydrate becomes less for that reason, and mycorrhizae are formed
more rarely. Similar reactions occur if light becomes weak. Mycorrhizal formation
ee we ee pee
622 FarLtowia, Vou. 1, 1944
is, expressed briefly, a result of and a sign that there is a certain surplus of energy-
giving nourishment in the host plant.
Bjorkman’s theory is this simple, but it is confirmed not only by his own involved
curves, but also by what is otherwise known. A direct support for it is the higher
sugar content that Rexhausen found in mycorrhizal roots. Bjorkman himself has
found a still clearer direct support for his theory. In two experimental series he
determined the amount of “reducing substance” in the roots of seedlings (in plain
language — simple kinds of sugar; also many other substances according to Bjork-
man). He who takes time to plot the data against the percentages of mycorrhizae
(which Bjorkman should have done himself) finds a beautiful correlation. It is
especially noteworthy that the relationship is so similar in two different soils.
However, not even such a prop for a theory is any final proof of it, and questions
remain. How does the fungus know that there is surplus sugar in the root? Does
it go and feel here and there, or does the root give off sugar or something else tempt-
ing when assimilation begins to satiate it? Still there does not seem to be room
for any major error in Bjorkman’s theory.
It is clear, although Bjérkman has not wished to say so, that Melin’s virulence
theory has a reversed picture of reality. Whether mycorrhizae are formed or not
depends more on the host plant than on the fungus, rather than as Melin and others
have believed. The old scoffing belief that mycorrhizae are formed when the plant
needs them came much nearer the truth, for a surplus of energy-giving food, which
actually is the deciding factor, is conditioned in turn by a deficiency of mineral
nutrients (c/N ratio). Among recent investigators Hatch came nearest the truth
(as Bjorkman mentions rather cursorily) by his hypothesis that mycorrhizae are
formed when there is too low a content in the plant of one or another of a number
of nutrient substances.
Bjorkman’s work throws new light not only on the problem with which he has
been especially occupied. The solution he found to this question falls at once as a
judgment of Solomon on the old strife between the “nitrogen theory” and “salt
theory” of the ecology of mycorrhiza. It shows both theories to be more or less
correct, but erects in their places a new, simple and beautiful theory, with an inner
strength which no other mycorrhizal theory has had, for it gives an actual well-
grounded explanation which appears axiomatic when it has once been clearly
apprehended.
The new theory should be called the Frank-Bjérkman, for it is a deepened and
expanded form of Frank’s old one. According to it the mycorrhiza comes more than
ever to appear like a factory for mutual profit. False must be called the whole view-
point now for some time considered the only scientific one the world round: that
altogether too human fable of the brutal burglar or the company formed by two
gangsters who steal from each other. The exchange of substances in the mycorrhiza
appears instead as a peaceful exchange where one party gains from the other’s un-
necessary surplus, by offering goods he lacks in his activity, or if one will, by hiring
and paying an industrious servant. Probably never before has anyone shown so
clearly how the ways of nature can lead to a symbiosis between plants to their mutual
advantage, and how this living together in many cases must inevitably take on this
form just because an exchange serves its needs. Bjorkman’s work is destined to
make a deep imprint on biological thinking in due time. His theory may furnish
the key to the mystery of the symbiosis of alder and its nodule former, and perhaps
also that of legumes and their root bacteria.
A theory of such breadth as Bjérkman’s raises the question of exactly how it came
about. One can read that it grew out of the hypothesis that the organisms forming
mycorrhizae live on their host trees, and from Bjérkman’s interpretation of 1940 based
upon it, of the effect of light upon mycorrhizal formation. However it is far from
there to the complete theory. One thing and another suggest that Bjorkman almost
brought out his work without its real point. Probably he came upon the solution
when he read in the literature about the influence of phosphorus on the sugar con-
REVIEWS 623
tent of beets and other plants, and about nodule formation on roots of legumes where
the carbon-nitrogen ratio is at present considered to be the controlling factor. In
such cases it is proof that there can be a certain value to the learning they are wont
to have in Swedish doctorate dissertations. It in no wise follows from the above,
however, that Bjérkman got his theory without effort, or even that almost anyone
could have hit upon the solution by reading the literature. Others before this have
brushed lightly past the key Bjorkman found, but have not seen it.
If spruce seedlings die out on mull rich in nitrate, then it cannot reasonably be
attributed to their failure to get the mycorrhizae they need (Lindquist 1932). When
they do not have mycorrhizae it is on the contrary a proof that they do not lack any
nutrient that is consumed in further elaboration of the first formed carbohydrates,
and it is probably because they lack light.
Black mycorrhizae are no sign that the soil is sick or cold so that a cutting must
~be made (Lindquist 1937). It indicates only weak light, that is, the stand is un-
usually dense for northern Sweden. It must always remain a difficult task to judge
the soil by the mycorrhizae, since poor mycorrhizal formation can depend either upon
high nitrogen and phosphorus nutrition, or too little of either or both, or too scant
light. Now perhaps it can be done in certain cases, thanks to what we know through
Bjorkman. — Lars-Gunnar Romell in Svenska Skogsvardsféreningens Tidskrift. Haft
IV. 1942, [Translated by H. I. Baldwin]
RECHERCHES SUR LES CERAMIACEES DE LA MEDITERRANEE OCCIDEN- |
~ TALE. By G. Feldmann-Mazoyer. Algiers, Imprimerie Minerva, 1940 (1941).
510 pp. 191 text figs. 4 pls.
In this important memoir, Mme. Feldmann-Mazoyer has presented the results of
many years of research prosecuted in the Laboratory of Botany of the Faculty of
Sciences of Algiers and in the marine biological stations of the Faculty of Sciences
of Paris at Banyuls and Villefranche.
The researches of the author deal with: 1) the cytology of the Ceramiaceae, 2) the
morphology of their vegetative structures, 3) their reproduction, and 4) their tax-
onomy. In these four series of researches, the author has obtained new results and
has clarified questions which until now have remained obscure.
In the parts devoted to cytology, the membrane, the nucleus, the cytoplasm and
its elements, and the products elaborated by the cell (special floridean starch, os-
mophilic granulations, basophilic bodies, . ..) are studied successively. The author
makes many new contributions to the knowledge of the floridean cell, notably as it
concerns the structure of the membrane, the morphology and the variations of the
plastids, the structure and metabolism of floridean starch, etc. She has particularly
studied the secretory cells, “bromuques,” which according to Sauvageau contain free
bromine. By.new microchemical methods she has been able to establish the absence
of free bromine in these cells. Also, the study of intercellular relations has permitted
her to uncover in the axial cells, structures which are active in the translocation of
dissolved nutrient substances, the structures playing the réle that had been attributed
to the cells themselves. She has been able to confirm experimentally the role at-
‘tributed to. the synapses in these phenomena.
In the portion devoted to the morphology of the vegetative structures, the author
has been able to clarify structures heretofore poorly known. She has shown in par-
ticular that the verticillate epidermis of species of Griffithsia correspond to the trich-
oblasts of the Rhodomelaceae and has demonstrated their function as an absorp-
tive layer.
In the part treating with reproduction, the author has been able to describe the
development of the reproductive organs in certain genera and species in which this
was poorly known or even completely unknown.
The systematic part constitutes a veritable monograph of the Ceramiaceae of the
western Mediterranean. Using the results of researches enumerated above, the author
624 FarLowi4, Vou. 1, 1944
has ben able to establish a more rational classification of the family based principally
on the structure of the procarp, in part on the manner of development of the gonim-
oblast, and in part on the cytological structure. She has established three new tribes
and modified the limits of many others, has created three new genera and suppressed
four. She has described some new species and on the contrary has suppressed a
large number, the study of very abundant living material having permitted her to
determine that these so-called species represented only seasonal or ecologic variations.
In each genus, very carefully prepared dichotomous keys permit the determination
of the species which are later described in detail. Numerous illustrations facilitate
further and render more certain this determination.
The work of Mme. Feldmann will remain the basis of all studies of the Ceramiaceae,
not only in the western Mediterranean, but of the entire world. Its great merit is
that it was founded not only on herbarium material, but more especially on living
plants, and that it has correlated, thanks to this fact, the numerous physiological and
cytological characters with the purely morphological concepts of the earlier algologists.
—R. Maire [Translated by D.H.L.]
1(4) 3 625-641
FARLOWIA
July, 1944
INDEX TO VOLUME 1
INDEX TO AUTHORS AND TITLES
Baker, E. E., E. M. Mrak and C. E.
Smith. The morphology, taxonomy,
and distribution of Coccidioides im-
mitis Rixford and Gilchrist 1896, 199—
244.
Barghoorn, E. S. and David H. Lin-
der. Marine fungi: their taxonomy
and biology, 395-467.
Bartram, Edwin B.. Additions to the
moss flora of northern Chile, 191-194;
Burma mosses, 171-189; Mosses of
Papua, New Guinea, 41-47; New and
noteworthy Philippine mosses, 503-513.
Chapman, V. J. The aims of future
research in the algae, 5-8.
Dixon, H. N. Alpine mosses from New
Guinea, 25-40.
Gosselin, Roger. Studies on Polystictus
circinatus and its relation to butt-rot
of spruce, 525-568.
Greene, H. C. Notes on Wisconsin
parasitic fungi, IV, 569-581.
Herre, Albert W. C. T. A_ neglected
field of study with the description of a
new lichen from California, 391-393.
Jackson, H. S. see Rogers, D. P. and
H. S. Jackson.
Khanna, L. P. On two new species of
Megaceros with notes on M. arachnoi-
deus, M. denticulatus, M. giganteus,
and M. grandis, 515-523.
Linder, David H. Foreword, 1-3; The
genera Kickxella, Martensella, and
Coemansia, 49-77; see also Barg-
hoorn, E. S. and D. H. Linder.
May, Valerie. Marine phytogeography,
491-493.
Merrill, Elmer D. An index to Rafi-
nesque’s published technical names for
the cellular cryptogams, 245-262.
Mrak, E. M. see Baker, E. E., E. M.
Mrak and C. E. Smith.
Nickerson, Walter J. Studies in the
genus Zygosaccharomyces. I, Transfer
625
of pellicle-forming yeasts to Zygopich-
ia, 469-481.
Papenfuss, George F. Notes on algal
nomenclature. III. Miscellaneous spe-
cies of Chlorophyceae, Phaeophyceae
and Rhodophyceae, 337-346.
Prescott, G. W. New species and va-
rieties of Wisconsin algae, 347-385.
Prince, Alton E. Basidium formation
and spore discharge in Gymnosporan-
gium nidus-avis, 79-93.
Rogers, D. P. The genus Pellicularia
(Thelephoraceae), 95-118.
Rogers, D. P. and H. S. Jackson.
Notes on the synonymy of some North
American Thelephoraceae and _ other
resupinates, 263-336.
Seeler, Edgar V., Jr.
colous fungi, 119-133.
Smith, C. E. see Baker, E. E., E. M.
Mrak and C. E. Smith.
Smith, Gilbert M. Microaplanospores
of Vaucheria, 387-389.
Svedelius, Nils. Galaxaura, a diplobi-
ontic floridean genus within the order
Nemalionales, 495-499.
Whelden, Roy M. Notes on New Eng-
land algae III. Some interesting algae
from Maine, 9-23.
Whetzel, H. H. A new genus of the
Sclerotiniaceae, 483-488.
Whiffen, Alma J. A discussion of tax-
onomic criteria in the Chytridiales,
583-597.
White, W. Lawrence. Studies in the
genus Helotium, III. History and di-
agnosis of certain European and North
American foliicolous species, 135-170;
Studies in the genus Helotium, IV.
Some miscellaneous species, 599-617.
Wiedling, Sten. Morphologic and
physiologic investigations in diatoms,
501-502.
Several fungi-
626
FarLtowia, VoL. 1, 1944
InpEX 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 italics.
Acanthocladium baculiferum 186; ben-
guetense var. latifolium 187
Acanthorrhynchium papillatum 46
Achlyogetonaceae 585 ;
Acia ferruginea 272; tomentosa 271
Aciella tomentosa 271
Acinaria 251; coccifera 251; flexuosa 251;
latifolia 251; salicifolia 251
Acroporium diminutum 39; oxyporum 45
Acrosporangiatae 498
Actigea 255, 259; multifida 255; sicula
255
Actigena multifida 255; sicula 255
Actinomorpha aurantiaca 255
Actinothuidium Hookeri 185
Actinotrichia 496, 498
Actycus siculus 255
Aecidium citrinum 255
Aedycia 249, 255; alba 255; rubra 255
Aerobryidium aureo-nitens 182
Aerobryopsis 36; longissima 181
Agaricus 293; aterrimus 255, 256; az-
ureus 255, 256; ellipticus 255; miptica
256; surrectus 256
Aglaozonia 6
Alcyonium Bursa 253
Aleurodiscus 95, 265, 267-271, 276, 307;
albo-roseus 268; albus 306, 307; amor-
phus 269; amylaceus 291, 300; api-
culatus 267, 268; aurantius 307; Bertii
268, 307; botryosus 299; Burti 268;
candidus 268; crassus 268; cremeus
268; croceus 304; diffissus 270; dis-
ciformis 279; Grantii 269; helveolus
269; javanicus 267; lepra 268; macro-
sporus 307; Micheneri 303; mirabilis
267, 268; Oakesii 268; pallide-roseus
268; paraphysatus 297; peradeniae
267; Peteloti 268; polygonioides 300;
roseus 299; sajanensis 270; salmoneus
268; sinensis 268; spinulosus 267; sub-
cruentatus 307; subgiganteus 303;
succineus 270; usambarensis 267; Zell-
eri 270, 319, 324 |
Allophylaria phyllophila 158
Alternaria maritima 405
Alveolinus 256
Alytosporium croceum 304
Amanita aterrima 255, 256; azurea 253,
256
Amasperma 251; flocculosa 251; monilia
251; torulosa 251
Amaurodon viridis 277
Amblystegiaceae 193
Amerosporae 407
Amoebochytrium 592, 596
Amphinema sordescens 276
Amphiroa 337; tribulus 338
Amphisphaeria maritima 411, 440, 442-
447, 450, 451, 453, 455, 457, 458
Anabaena 10; aphanizomenoides 373;
flos-aquae 10; flos-aquae var. minor 10;
minutissima 10; wisconsinensis 373
Anastomaria campanulata 256; dimidiata
256
Anema Dodgei 393
Aneuriton 251; marginatum 251
Angstroemia orientalis 504
Annularia 256
Anoectangium Stracheyanum 174
Anomobryum cymbifolium 176; gemmi-
gerum 171, 176
Anomodon toccoae 506
Anthoceros 392
Aphanistis 591, 993
Aphanizomenon flos-aquae 373
Aphanocapsa 392
Aporemia 251
Arthrocormus Schimperi 29
Arthrodesmus quadridens 16
Arthrodia 251; linearis 251
Artocreas Micheneri 303
Ascochyta Thaspii 580
Ascophyllum 7
Aspergillus 101, 446, 471; fumigatus 205
Asterodon ferruginosus 271
Asteromella astericola 572; Asteris 572
Asterophlyctis 591, 592, 594
Asterostroma 265, 271, 272; andinum
271, 272; bicolor 271, 272; gracile
271; ochrostroma 271; spiniferum 271,
272
Asterostromella 264, 293; dura 310; ef-
fuscata 290; epiphylla 323; granulosa
283; investiens 292; rhodospora 294
Astrocitum dimidiatum 256; multifidum
256; quinquefidum 256
INDEX TO
Astrycum dimidiatum 256; multifidum
256; quinquefidum 256
Athelia citrina 304; epiphylla 323; sericea
308, 309; strigosa 8 muscigena 275
Atractylocarpus comosus 28; dicranoides
28
Atrichum flavisetum 189; undulatum 189
Auricularia aurantiaca 319
Barbella burmensis 182; enervis 182;
javanica 182; pendula 182; spiculata
182; Stevensii 182
Barbula consanguinea 175; constricta
175; indica 174; replicata 191; Wis-
selii 30
Bartramia cubica 33; Halleriana 177;
quadrata 34
Bartramiaceae 33, 177, 506
Battarrea 261
Belonium sulphureo-tinctum 169, 170
Bescherellea Eyrtapus 35; Cyrtopus var,
papuana 35
Blastocladiales 593
Blastomyces 211, 213
Blastomycoides 213, 219; dermatitidis
213; immitis 213, 220; Tulanensis 213
Blastosporidium 219; Schooii 212, 220
Blyttiomyces 590-593, 595
Boletus hematoporus 256; pusillus 256
Bombardia 409
_ Botryobasidium 95, 96, 97, 264, 272, 273;
coronatum 107, 272; flavescens 105,
106, 272; granulatum 107; isabellinum
99, -272; ochraceum 105, 106, 272;
Solani 113, 272; subcoronatum 104,
272; vagum 110, 272
Botryocladia Skottsbergii 344
Botryohypochnus 96, 100
Botryophialophora 403; marina 404
Botrytis 577; cinerea 484, 487, 577
Bourdotia deminuta 294
Brachyhymenium acuminatum 176; nepa-
lense 33, 176
Brachytheciaceae 37, 44, 185
Brachythecium Buchanani 185; longi-
cuspidatum 185
- Braunfelsia dicranoides 28
Breutelia Romeri 34
Brotherella erythrocaulis 187
Brotherobryum Dekockii 28
Bryaceae 32, 44, 176, 192
Bryonopsis 251
Bryopsis 251
Bryum argenteum var. lanatum 176;
coronatum 44, 176; crispo-capillare
VOLUME 1 627
33; flexisetum 192; megalothecium
192; nitens 176; pachycladum 33; por-
phyroneuron 176; ramosum 176; scle-
rodictyon 33
Bytholotrephis 492; gracilis 492
Caldesiella 272; vaga 309; viridis 277
Callicostella Karnbachii 44
Calocera cornea 88
Calothrix 391
Calycella callorioides 601; Jlicis 160
Calycellina 160, 166; Roipuliies 161, 169;
punctiformis 161, 166
Calycina aurantiaca 140; epiphylla 140;
fastidiosa 150; Ilicis 160; naviculaspora
151; populina 160; saprophylla 151
Calymperaceae 30, 42, 173, 504-506
Calymperes daruense 43; Ebaloi 505;
Hampei 174; moluccense 43; Motleyi
43; peguense 173; porrectum 505; sala-
kense 43, 505; serratum 505; stricti-
folium 505; tuberculosum 505
Calyptothecium burmense 183; himan-
tocladioides 171, 183; Ramosii 183;
Urvilleanum 183
Campylodontium flavescens 185
_Campylopus austro-subulatus 28; gracilis
172; laetus 172; novae-guineae 28;
umbellatus 172
Carteria ovata 6
Catenaria 215, 251, 592, 596; Anguillulae
216; arenaria 251; concatenata 252;
vagabunda 252
Catenarioideae 592
Catenochytridium 591-593, 595
Ceramium 252; callithamnion 252; opac-
um 252; squamosum 252
Ceratobasidium 116, 264, 272, 327; atrat-
um 272; cornigerum 116; fibrillosum
327; nlambede 272, 273; feriginat
cum 116
Ceratostoma vitreum 125
Cercoseptoria Blephiliae 578
Cercospora Alni 579, 580; Barbareae
579; Boutelouae 579; briareus 581;
gentianicola 581; incarnata 581; Nas-
turtii var, Barbareae 579; Violae 581.
Cercosporella Pycnanthemi 578
Ceriospora 398, 408
Ceriosporopsis 408, 411; halima 409,
440-442, 444-451, 453-455, 457
Cerocorticium albissimum 314
Cerophora 256; capitata 256; clavata 256;
dichotoma 256; fastigiata 256; globosa
256; globularis 256; minuta 256; pyri-
628 . Fartowia, Vou. 1, 1944
formis 256; ramosa 256; thamnoides
256
Chaetangiaceae 498, 499
Chaetangiae 498
Chaetangium 496, 498; Hystrix 342
Chaetocladium 54
Chaetocorticium fusisporum 320
Chaetomitrium 30; acanthocarpum 44;
orthorrhynchum 44; torquescens var.
barbatum 37
Chaetonema 351
Chaetophoraceae 350, 351
Chantransia 498; dichlora 252
Chara 251-253; capitata 253; fetidissima
252; patens 252
Characias 252
Chlamydomonadaceae 348, 349
Chlamydomonas polypyrenoideum 348;
variabilis 6
Chledipole 252; lobata 252; tubulosa 247,
paivs
Chledripole 252
Chlorociboria 135
Chlorophyceae 10-19, 348, 349
Chlorophyta 348-361
Chlorosplenium 135
Chondrus capensis 343 :
Chromosporium pactolinum 297; viride
292
Chroococcaceae 371, 372
Chroococcales 371, 372
Chrysophyceae 7, 363, 364
Chrysophyta 361
Chrysymenia Skottsbergii 344
Chytridiaceae 584, 590, 591
Chytridiales 583-597
Chytridioideae 590-593, 595, 596
Chytridium 587, 590-595; Lagenaria 587
Ciboria 135, 483
Cirriphyllum cameratum 38; cirrosum 37;
novae-guineae 37; subenerve 38
Cladochytriaceae 592, 595, 596
Cladochytrioideae 592
Cladochytrium 584, 592, 596; hyalinum
596
Cladophora 5
Cladopodanthus 43; heterophyllus 43;
muticus 43
Claopodium nervosum 184
Clastobryum caudatum 186; caudiforme
186; Dickasonii 186
Clathrus 249, 257; columnatus 257; trun-
catus 257
Clavaria bicolor 256; byssacea 278, 279;
citrina 256; citrinofusca 256; contorta
270; driophylla 256; dryophylla 256;
fistulosa 270; Himantia 278; lepido-
rhiza 257; oxantha 257; rubescens 257;
tricolor 257
Cleistostoma ambigua 171, 179
Closteridium 10; siamensis 11
Closterium 12-14, 18, 251; angustatum
12; macilentum 9, 12; Ralfsii var. hy-
bridum 12; rostratum 12; setaceum 12;
striolatum 12
Coccidioides 212-214, 216, 217, 219, 220;
braziliensis 214; esferiformis 214; im-
mitis 199-244; immitis var. metaeuro-
peus 214, 219, 220, 223; immitis var.
Pipkini 214, 219, 220; immitis var.
typicus 214, 219, 220; pyogenes 212,
213, 220
Coccidioidiaceae 218
Codiaceae 491
Codium dichotomum 338; prostratum
338; tomentosum 338, 491; Vaughani
338
Coemansia 49-57, 59, 61, 68; aciculifera
57, 65, 67; braziliensis 57, 64; brevi-
ramosa 57, 62; ceylonensis 56, 57,
60; erecta 57, 62, 63, 65; guate-
malensis 57, 64, 65; interrupta 57,
62; kamerunensis 57, 63, 65; pecti-
nata 51, 55, 57, 66-68; repens 68;
reversa 53, 55, 57, 60, 61; scorpioidea
55-57, 66; spiralis 57, 60, 61; Thax-
teri 57, 66, 67
Coemansiella 50, 57; alabastrina 57
Coenomyces 592, 596
Coleosporium 81, 85
Colonnaria 249, 250, 257; truncata 249,
257; urceolata 249, 257
Colophermum floccosum 252
Coltricia tomentosa 526, 527
Conferva isacella 252
Coniophora 116, 265, 273-280, 287; albo-
flavescens 277, 280; arida 280; arida
subsp. fumosa 267, 274; atrocinerea
266, 273, 274, 280; avellanea 275, 300;
Betulae 280; brunneola 273; byssoidea
275, 314; centrifuga 286; cerebella 276;
corrugis 276; corticola 277, 280; crocea
277; cyanospora 277; dryina 277, 279,
325; Ellisii 273; flava 278; flavomar-
ginata 278; fulvo-olivacea 274; fumosa
273, 308; furva 273; fusca 274; fusi-
spora 105; Harperi 277, insinuans 310;
Karstenii 274; laeticolor 277, 278;
leucothrix 273; mustialaensis 277;
ochracea 105; ochroleuca 102, 103;
Pe eee Sa ery TE ee er Ce en Mee Rags ys, ©
InDEX TO VOLUME 1 629
olivacea 266, 273, 274, 280, 308; Poly-
poroidea 277, 279, 310; puteana 276;
sibirica 274; Sistotremoides 274, 280,
308; sordulenta 309; suffocata 280,
308; sulphurea 308; umbrina 273, 275,
280; vaga 105, 106, 280
inum 277; effuscatum 290; effusum
290; Eichlerianum 314, 315; Ellisii
273; epiphyllum 323; ermineum 291,
300, 312; evolvens 318; fenestratum
105, 106, 291; fibrillosum 307, 308;
flavescens 105, 107; flavissimum 304,
vy eh ee Re ee ae eee
esha Bag
305; frustulosum 105, 107; frustulosum
var. intermedia 105; fumesum 273, 308,
309; furfuraceum 285,°286; fuscostrat-
um 291; fusisporum 103, 105; Galzini
285; geogenium 281, 291, 306, 311;
glebulosum 284; Greschikii 310, 311;
ulaqueatum 281, 292; illaqueatum fa.
rhizophorum 281; inaequale 311; in-
canum 292; incrustans 285, 299; in-
vestiens 292, 299; involucrum 293, 294;
isabellinum 99, 107-109; jamaicense
294, 310; javanicum 267; Koleroga
112, 113, 294; lacteolum 286; lacteum
Coniophorella 274; atrocinerea 273; bys-
soidea 276; laeticolor 278, 279: ochro-
leuca 102; olivacea 274; olivacea subsp.
fulvo-olivacea 274; umbrina 274; um-
brina var. olivacea 274
Conjugales 360, 361
Coprotinia 484; minutula 484—487
Corallina Flabellum 337
Coralliochytrium 591, 592, 595
Cordyceps 398
Coronella 49-51, 57; nivea 49, 50, 57
Corticium 59, 95, 264, 265, 279, 280-306;
sect. Athele 292; sect. Botryodea 96;
abeuns 280, 281; albo-cremeum 295;
alboflavescens 277 ; albo-ochraceum 279,
290; albo-ochraceum subsp. amianthin-
um 289; albo-stramineum 281, 311, 312,
321; albulum 312, 316; album 116,
Allescheri 313; alutaceum 59; alutarium
292, 293; amianthinum 289, 290; amor-
_phum 269; amylaceum 291; apiculatum
281, 282, 292; arachnoideum 282,
286-288; araneosum 99; areolatum
113-115, 281, 282; argillaceum 296;
Atkinsonit 282; atratum 272; auranti-
acum 270; Berkeleyi 282, 283, 326;
bicolor 304, 305; bisporum 286, 288;
bombycinum 283, 295, 296, 325; botry-
oideum 107, 108, 111, 284; botryosum
108, 110, 111; brunneolum 273; bys-
sinum 304, 305; byssinum var.. micro-
sporum 304; byssoideum 275; caerule-
um 273; calceum 284, 285, 317; calo-
thrix 316; calotrichum 318; canum 285,
286; centrifugum 282, 286, 287; cen-
trifugum subsp. bisporum 287; cen-
trifugum subsp. fugax 287; centrifugum
var. macrospora 287; centrifugum var.
soredioides 286; centrifugum var. tenuis
287; Chusqueae 282; citrinum 304;
confine 288; confluens 275, 276, 296,
300, 306; consimile 286-288; contiguum
288, 289, 301, 302; cornigerum 116;
coronatum 107, 109; coronilla 286, 288,
292, 328; corruge 2763; cremeum 314,
315; cremoricolor 327; cretaceum 284,
285, 325; creceum 304; crustaceum
288, 289, 302; crustulinum 289; de-
cipiens 286-288; deglubens 327; dry-
294-296; lacunosum 275; laeticolor
277; laetum 296; laeve 318; Jatitans
282; leucothrix 273; Litschaueri 281,
300; livido-caeruleum 281; Macounii
305, 306; maculatum 305, 306; micro-
sclerotia 113-115; microsporum 286:
miniatum 319; Murrayi 290; mustia-
laense 277; mutatum 313; myxosporum
282; ochraceum 107, 289; ochrofarctum
296; ochroleucum 112, 113, 281, 291,
292; octosporum 287; oleosum 313;
olivaceum 273; Overholtsii 296, 297;
pactolinum 297; pallescens 310; pal-
lidum 296; paraphysatum 297, 307;
pectinatum 298; peradeniae 267; per-
tenue 322; Pezizoideum 286-288, 298;
pilosum 298, 299, 302; Pini-canaden-
sis 323; Polyporoideum 279, 310;
porosum 281, 289, 299, 300-302, 305;
praetermissum 322; praticola 98, 115;
pruinatum 107-109; Pruni 299, 301,
302; puberum 325; punctulatum 281,
295, 320, 321; puteanum 276; racem-
osum 298, 299; radiosum 294, 295;
rhizophorum 281; roseopallens 299:
roseum 299, 300; Roumeguertt 324;
rubellum 275, 300; rude 310; Sambuci
325; sanguineum 319; scariosum 327;
secedens 327 ; septentrionale 300; serum
325; setigerum 282; soctatum 291, 300;
Solant 111, 113; sordidum 314, 315,
321; sordulentum 309; sphaerosporum
286, 292; spinulosum 268; sterigmat-
icum 116; Stevensti 112, 113, 300;
stramineum 299, 300, 301, 305; sub-
album 298, 299, 302; subapiculatum
630
324; subcinereum 289, 302; subcoron-
atum 104, 105, 302; subgiganteum 291,
303; subincarnatum 303, 324; sub-
pallidulum 285; subsulphureum 303;
suecicum 285; suffocatum 280; sul-
phurellum 277; sulphureum 99, 113,
303, 304, 305, 308, 309; tenue 322;
terrigenum 116; tessulatum 281, 292;
teutoburgense 105; Thelephoroides 309 ;
Torrendii 312; trigonospermum 307,
308; Tsugae 296; tuberculatum 295;
Tulasnelloideum 292; umbrinum 275;
usambarense 268; vagum 108, 110, 111,
113, 115, 305, 308; vagum subsp. Solani
113; vagum var. Solani 113, 305; vel-
lereum 295; versatum 326; vesiculos-
um 301, 302, 305; vinososcabens 305,
306
Cosmarium 19; bioculatum 13; sub-
turgidum 13; subturgidum fa. minor 13
Cratoneuron filicinum 193
Crossidium Roseae 192
Cryptococcus 213
Cryptophaeella Heteropatellae 120
Cutleria 6
Cyanophyta 371-373
Cyathela 257
Cyathella 257
Cyathicula 166
Cyathophorella 507
Cylindrochytridium 593, 595
Cylindrochytrium 593
Cylindrosporium Gei 577; triflori 577
Cynicus 257
Cyphella amorpha 269
Cyrtopodaceae 35
Cystopus 328
Cystoseira australis 343; caudata 343
Dacryomyces 259, 262
Daedalea 257
Daltonia aristifolia 509; armata 508,
509; Macgregorii 37
Dangeardia 591, 593, 594
Dasyscypha puberula 169
Dedalea 257
Deloxus 252
Deraphytus 252
Derbesia marina 6
Derepyxis amphora 362
Desmatodon involutus 175
Desmotheca apiculata 31; cymosa 31
Diaphanodon thuidioides 179
Dicarphus 257; rubens 257
Dichostereum durum 310
FarLowiA, Vou. 1, 1944
Dicnemonaceae 28
Dicnemos 29; calycinus 29; rugosus 29;
undulatifolius 28
Dicoccum nebulosum 578
Dicranaceae 26, 172, 504
Dicranodontium uncinatum 172
Dicranoloma fragile 173; laevifolium 28;
novo-guinense 28
Dicranoweisia alpina 27; fastigiata 27;
papuana 27
Dicranum gymnostomum 173
Dicteridium 257
Dictilema 252; glomerata 252; xantho-
sperma 252
Dictyota dichotoma var. implexa 338, 339;
dichotoma var. intricata 338, 339; im-
plexa 338 ; interrupta 339; subarticulata
339, 340
Didymosporae 408-413
Didymostilbe 122
Dilophospora Alopecuri 577
Diphas 250, 262
Diphascium 250
Diphascum 250, 262
Diphyscium 262
Diplodia orae-maris 403
Diplonema sordescens 275 —
Diplophlyctis 587, 591, 592, 595
Diplophlyctoideae 590, 591, 592, 595,
596
Dipodascus 217
Dispira cornuta 53
Ditrichaceae 26, 172
Ditrichum Colijnii 26; spinulosum 26;
spirale 26
Draparnaldia Judayi 351
Draparnaldiopsis alpinis 351
Druparia globosa 257; rosea 257; violacea
257; volvacea 257
Drupasia globosa 257; rosea 257; violacea
257
Dycticia clathroides 257
Ecklonia biruncinata 341; exasperata
341; radiata 341
Ecpexis fluviatilis 252
Ectocarpus confervoides 493; siliculosus
6
Ectosperma longiuscula 252
Ectospermia 252
Ectropothecium aureum 40; dealbatum
187; dentigerum 39; eleganti-pinnatum
40; laticuspes 39; longicapillare 46;
Micholitzii 46; Moritzii 40; plumosum
40
OF EE OE ATO MEL ee eae Sie
INDEX TO VOLUME I 631
Eleutheris 123
Eleutheromycella 121; mycophila 119,
123
Eleutheromyces 119, 121; Geoglossi 122,
126; longisporus 121; subulatus 119-
122, 123
Empusa 86, 211
Endacinus 257; tinctorius 257
Endaematus albus 257
Endematus 257
Endochytrioideae 591
Endochytrium 591, 592, 595, 596
Endocoenobium 592, 595
Endoconia leucomela 257; stuposa 257
Endomyces 212
Endomycetaceae 469
Endonema 252
Endonius 257
Endosperma aggregata 252; globosa 252
Endotrichella arfakiana 36; Campbelliana
36; nematosa 507; rigida 180
Enteromorpha 5, 7; prolifera 6
Entodon flavescens 186; plicatus 186
Entodontaceae 45, 185
Entomophthora 84, 88, 89
Entomophthoraceae 88
Entophlyctaceae 590, 591, 592, 595, 596
Entophlyctis 586, 587, 591, 592, 595
Entophlyctoideae 590-592, 595, 596
Entosthodon wichurae 175
Epidrolithus 250, 262
Episperma micramnia 252
Epixyla clavata 258; cylindrica 258;
lineata 258
Eriosperma alba 258; fugax 258
Erysiphe graminis 569; scandens 112
Erythrodontium julaceum 185
Euastrum 13, 18, 19; humerosum 13; ob-
longum 13
Eucamptodon 29; novae-guineae 29
Eucapsis 9-11; alpina 9; alpina var.
minor 10
Eudacnus 258
Euglena breviflagellum 364; elastica
365; minuta 365; oxyuris var. minor
366; spirogyra var. marchica 369
Euglenaceae 364-371
Euglenophyta 364-371
Eunotia pectinalis yar. minor 501
Exoascaceae 469
Fabroniaceae 193
Favaria 258
Favolus 260; europaeus 124
Fissidens 250, 262; Brassii 41; bryoides
41; diversiretis 504; geniculatus 26;
kurandae 26; lagunensis 504; micro-
cladus 41; nobilis 172; papuensis
41; rupicola 26; subspathulatus 25;
sylvaticus 172, 503; Wichurae 504;
Zwickeyi 503
Fissidentaceae 25, 41, 172, 503, 504
Fleischerobryum longicollis 178
Floribundaria aurea 182; floribunda 182
Fomes annosus 86; circinatus 527, 528;
Pini 530 ,
Foreauella orthothecia 188
Forsstroemia rigida 35
Fucus 6, 7; compressus 343; elongatus
341; filicinus 254; fungosus 253; im-
plexus 338; interruptus 339; loreus
341; ovalis 344; ovatus 344; radiatus
341; radiatus B exasperatus 341; to-
mentosus 253, 338; vermicularis 343,
344
Funaria hygrometrica var. calvescens 175
Funariaceae 175
Fungi Imperfecti 402
Fusarium 577; moniliforme 578; tricinct-
um 578
Fusisporium Kuhnii 287
Galaxaura 337, 495-499; corymbifera
496; Diesingiana 496, 497; tenera 496,
497
Garckea phascoides 172
Garovaglia papuana 36; Pauerlenii 36;
rigida 507; subintegra 36; Zwickeyi
507
Gastroconium ovale 344: ovatum 344
Geaster 261
Gelatina 258; alba 258; aurantiaca 258;
candida 258; flava 258; foetidissima
258; lutea 258; rubra 258
Gelatinaria 258
Geminella crenulatocollis 349; muta-
bilis 350
Gemmularia 258, 261; albida 258; levius-
cula 258; rimosa 258; rugosa 258
Geoglossum 126, 258
Geotrichum 213, 219; dermatitidis 220;
immite 220; louisianoideum 220
Gigartina bracteata 342; Hystrix 342;
polycarpa 342; Radula 342; Radula
var. Hystrix 342; Radula var. Radula
342
Glaucocystis cingulata 372; duplex 371;
nostochinearum 10, 372; oocystiformis
372
Glenospora 219; metaeuropea 220, 223
Gloeocapsa 391, 392
Gloeocystidiellum 302; porosum 301
‘ti Ne ee
632 Fartowia, Vou. 1, 1944
Gloeocystidium 264, 302, 306; albe-
stramineum 306, 312, 321; albo-stra-
mineum var. causseanum 321; albo-
stramineum subsp. cremicoler 321;
albo-stramineum subsp. Kichleri 321;
albo-stramineum subsp. sphaerosporum
321; alutaceum 295; argillaceum 296;
Bourdotii 312; caliciferum 322, 323;
clavuligerum 302; contiguum 289,
301, 302; coroniferum 282; cremicolor
321; Eichleri 321; furfuraceum 302;
inaequale 311, 312; Karstenii 289, 302,
306; luridum 312; ochroleucum 281;
oleosum 313; pallidulum 313; pallidum
296, 313; pallidum subsp. argillaceum
296; porosum 301; praetermissum 322;
praetermissum var. Bourdotii 312; rude
311; stramineum 301; tenue 322; tenue
subsp. inaequale 311; tenue subsp.
praetermissum 322; Torrendii 312
Gloeocystis 348
Gloeopeniophora Allescheri 313
Gloeosporium 571,572; cinctum 571, 572;
Oncidii 572
Gloeotulasnella 265, 306; calospora 328;
opalea 306; traumatica 306 = *
Gloiophloea 496, 498
Glossadelphus hermaphroditus 510; sim-
ilans 509
Gonatobotrys pallidula 313
Gracilaria concinna 343
Grandinia granulosa 282, 283; granulosa
var. ochracea 283
Grateloupia Hystrix 342, 343
Gymnodiniaceae 371
Gymnodinium caudatum 371
Gymnogongrus capensis 343; vermicularis
343, 344
Gymnopus 258
Gymnosporangium 85, 89; juniperi-vir-
ginianae 86; nidus-avis 79-90
Gyrophana 287
Halicystis ovalis 6
Halimeda 337
Halophiobolus 398, 415; cylindricus
416, 442-447, 450, 451, 454; halimus
416, 419, 420; longirostris 416, 418,
453, 463; maritimus 416, 419; medusa
416, 419; opacus 416, 417, 418, 440,
442, 444-448, 450, 451, 453-455, 458:
salinus 416, 419, 440, 442, 444-448,
451, 453-455, 458, 462
Halosphaeria 412; appendiculata 412
Hansenula 480
Haplosporangium parvum 218, 219
Haplosporidia 212
Hectocerus 258; clavatus 258; dichotomus
258; globosus 258; pyriformis 258;
thamnoides 258
Hedwigiaceae 179
Helicodendron tubulosum 444
Helicoma Curtisii 458; maritimum 405,
407; Miilleri 458; salinum 406, 409,
440-442, 446-448, 454, 458 ©
Helicomyces scandens 444
Helmyton glomeratum 252; spiralis 247,
252
Helotium 135-170, 599-617; albidum 154,
599, 608; albopunctum 139, 154, 155-
157, 159; album 601; amenti 617;
aurantiacum 140-142; callorioides 601,
602; carpinicola 137, 138, 141, 142,
145, 146, 608; caudatum 137, 139, 150,
151, 154, 157, 159, 609; citrinulum
600-602, 604, 605; citrinulum var.
Seaveri 600, 601; citrinum 599; con-
formatum 136, 145-147, 162; Conocarpi
137, 138, 148; contortum 138, 147;
cyathoideum 599; Dearnessii 137, 613,
616; discretum 605; epiphyllum 135-
139, 141, 142, 144-147, 159, 162, 599,
606, 608; epiphyllum var. acarium 142;
epiphyllum var. candidum 142, 146,
147; epiphyllum var. Cerasi 141; epi-
phyllum var. Ellisii 142; epiphyllum
var. Illicina 162; epiphyllum var. prae-
ponens 142; erraticum 606; fastidi-
osum 139, 150, 157; flexucsum 600,
601; fraternum 137; fructigenum 599;
gemmarum 616; herbarum 599, 605;
Ilicis 160, 162, 164; immutabile 136-
138, 141, 142, 145-147, 150, 606, 608;
Linderi 139, 154, 160; lucellum 166,
167; lutescens 137; midlandensis 605,
608; naviculasporum 151, 152; pal-
lescens 137; phyllogenon 139, 155, 156,
157, 159; phyllophilum 135, 136, 139,
141, 152, 156, 157, 159, 162, 608; phyl-
lophilum fa. albescens 152; populinum
138, 159, 160, 162, 164, 166, 167, 170;
puberulum 138, 163, 167, 1693; puncta-
tum 160; punctiforme 160, 162, 167,
170; punctiforme fa. Tiliae 162; sapro-
phyllum 151, 152, 155; scutula 137,
154, 599, 609, 613, 616; scutula var.
albidum 152, 608; scutula var. caudatum
151; scutula var. fucatum 6093 sor-
didatum 152, 155; sparsum 15], 152;
translucens 139, 149, 150; trapezoide-
bgt neiraced
INDEX TO VOLUME 1 633
um 147; tumidulum 139, 165-167; vir-
gultorum 599
Helvella infula 128
Hepataria 258; cuneata 258; erecta 258
Hericium 256, 258; grande 258
Herpetineurum toccoae 506
Heterodon 250, 262; bryoides 250, 262
Heterolagynion Oedogonii 364
Heterophyllum brachycarpum 509; San-
tosii 509
‘Himanthalia elongata 341; lorea 341 .
Himantia sulphurea 304, 309; umbrina
273
Himantocladium cyclophyllum 37
Histoplasma 211 :
Holomitrium Griffthianum 27, 173;
noyae-guineae 27
Homaliodendron exiguum 184; flabel-
latum 184; microdendron 184
Homoeostrichus 340; flabellatus 341;
multifidus 340; Sinclairii 341; spiralis
341
Hookeriaceae 37, 44, 508
Hookeriopsis utacamundiana 37
Hormidiopsis crenulata 350; ellipsoide-
um 350
Hormogonales 372
Hyalinia tumidula 165
Hyalocomiaceae 188
Hyalopyenis 119, 128; blepharistoma
124, 125; hyalina 124, 125; vitrea 124,
125
Hyaloscypha lachnobrachya 167; puber-
ula 169; punctiformis 160
Hyalotheca 361
Hydnochaete setigera 271
Hydnum 257; aurantinum 258; auranti-
um 258; barbatum 259; caerulescens
259; citrinum 259; dilatatum 259; fer-
rugineum 272; ferruginosum 272; fusco-
atrum 279; granulosum 282; Himantia
278, 279; puniceum 259; setosum 279;
Sobolewskii 277; subfuscum 278; to-
mentosum 272
Hydrodictyaceae 356, 357
Hydromycus 259; aquosus 259; tremel-
loides 259
Hygroamblystegium austro-fluviatile 193;
crassicostatum 193; irriguum var.
spinifolium 193
Hygrohypnum peruviense 193
Hylocomiaceae 188
Hymenochaete 271, 315; Eillisii 273; pu-
bera 325
Hymenoscypha albida 609; albopuncta
155; citrinula 600; Dearnessii 613; epi-
phylla 140; glagosa 167; phyllogena
156; phyllophila 158; populina 161;
puberula 169; punctiformis 161; scutula
var. fucata 610; tumidula 165
Hyophila involuta 174
Hyphochytrium 215, 216; infestans 216
Hyphoderma asperum 282; puberum 325;
spiculosum 282; sulphureum 308
Hypnaceae 39, 46, 187
Hypnodendraceae 34
Hypnum austro-fluviatile 193; plumae-
forme 187
Hypnochnella violacea 116
Hypochnopsis mustialensis 277; ochro-
leuca 112
Hypochnus 95, 265, 293, 306-310; albo-
stramineus 312, 321; albus 306; argil-
laceus 99; bisporus 286, 287; Brefeldii
107; byssoideus 275; centrifugus 286;
chaetophorus 316; confluens 275; coro-
natus 107, 108, 110; cremicolor 320,
321; epiphyllus 323; fibrillosus 307;
filamentosus 113-115, 308, 309; flave-
scens 105; flavo-brunneus 280, 308;
flavus 99; fumosus 304, 308, 309; fusig-
er 103, fusisporus 102, 103, 105; gran-
ulatus 107; isabellinus 99, 107, 108,
309; Langloisii 101, 309; longisporus
315; muscorum 275; mustialaensis 277 ;
obducens 275; ochroleucus 112, 113;
olivaceus 273, 274; pallescens 294, 309,
310; Peniophoroides 294, 310; Poly-
poroideus 279, 310; puberus 325; rose- -
us 300; Sambuci 325; Schroeteri 107-
109; sericeus 308; serus 325; setosus
275; Solani 113, 115; sordidus 315,
320, 321; sparsus 99; strigosus 6 fila-
mentosus 275; subtilis 325; sulphureus
308; Thelephoroides 310; Tulasnello-
ideum 292; umbrinus 273, 275; vagus
309; violeus 328
Hypolepia 259; difformis 259; igniarias
259; ignarius 259
Hypomyces 124, 128; Geoglossi 119, 126
Inonotus hispidus 297; Leei 297
Isaria 61; brachiata 119, 120; felina 61;
odora 259
Isophlis concentricus 252
Isopterygium albescens 188; minutirame-
um 46; Textori 188
Jania 337; rubens 338
Juratzkea seminervis 193
634,
Kickxella 49-52, 54-57; alabastrina 49,
50, 54-57, 58
Kickxellaceae 49-56-77
Kneiffia Allescheri 313; aurantiaca 270;
cremea 314; Eichleri 320; glebulosa
284; globifera 317; laevis 318; latitans
282; longispora 315; muscorum 276;
polonensis 316; pubera 325; Roume-
guerti 324; sera 325; setigera 282, 284;
setigera var. trachytricha 283; subas-
condita 324; tenuis 322; Tomentella
276
Kneifhiella aspera 282; latitans 283; seti-
gera 282
Labyrinthula macrocystis 395
Lachnella puberula 169
Lactarius pargamenus 124
Lagynion reductum 363, 364; triangu-
laris var. pyramidatum 364.
Lamarckia vermilara 253
Lambertella 483, 487
Laminaria 253, 405;
radiata 341
Lamyxis 259
Langloisula 292, 293; spinosa 292, 293
Laternea 249; columnata 249
Leiacina 251, 253; capitata 253; lucida
253
Lentescospora 411; submarina 411
Lenzites 123
Leotia lubrica 124
Lepocinclis glabra fa. minor 370
Leptodontium erectifolium 30; kinaba-
luense 30; Warnstorfii 30
Leptogium 250, 262
Leptohymenium tenue 188
Leptopora 259; difformis 259; nivea 259;
stercoraria 259
Leptorima nivea 253; oculata 253; un-
dulata 253
Leptosphaeria muralis 413; orae-maris
413 :
Leptuberia amorpha 250, 262
Leucobryaceae 29, 43, 173, 504
Leucobryum 29; arfakianum 29; Bow-
ringii var. sericeum 44; Hollianum 173;
neilgherrense 173; scalare 173; steno-
phyllum 29
Leucodontaceae 35
Leucophanes candidum 44; Massarti 504;
octoblepharioides 504
Liagora 337
Lichiton mamillaris 253; phyteloides 253,
254; tuberculatum 253
Limacium penarium 124
biruncinata 341;
Fartowla, Vout. 1, 1944
Linderia 249, 250, 257
Linospora 417
Lloydella Karstenii 326
Lycoperdon 260; coccineum 259; com-
planatum 259; violacinum 259, 260
Lycopodium violacinum 259
Lyngbya gigantea 373; Hummelii 373;
latissima 372, 373
Lyomyces 300; byssinus 304; isabellinus
99; mustialensis 277; roseus 300; Sam-
buci 325, 326; serus 325; sulphureus
308
Macrochytridioideae 591
Macrochytrium 591; botrydioides 216
Macromitrium 195-198; angustifolium
44; Blumei 195;_ brachystele 31;
Dickasonii 178; falcatulum 179;
goniorhynchum 179; leucoblastum 31;
lonchomitrium 31; longipes 31; nepa-
lense 178; ruberrimum 31; salakan-
um 196, 197; semipellucidum 44; sub-
megalocladum 31; sulcatum 179
Macrothamnium macrocarpum 188; sub-
macrocarpum 188
Mallomonadaceae 363
Mallomonas pseudocoronata 363; ur-
naformis 363
Martensella 49-52, 54, 57, 58, 59, 61, 68;
Corticii 55-57, 59, 61; pectinata 50,
51, 55, 57, 58, 59, 68; spiralis 60
Martinia 483, 487; panamaensis 487
Massarinula 410
Mastopoma Armitii 38
Megaceros 515-523; alatifrons 519; amoe-
nus 519; aneuraeformis 519; arachno-
ideus 515-517, 519; caledonicus 516,
519; callistictus 519; carnosus 519;
celebensis 516, 519; columbianus 519;
crassus 516, 519; cristisporus 519; den-
ticulatus 515-517, 519; endiviaefolius
516, 519; flagellaris 519; flavens 519;
fuegiensis 515, 516, 519; giganteus
516, 519; gracilis 519; grandis 517,
519; guatemalensis 519; Hodgsoniae
515, 516; jamaicensis 519; Jamesonii
516, 519; lacerus 516, 519; laciniatus
519; leptohymenius 519; longispirus
516, 519; martinicensis 515, 516, 519;
mexicanus 519; minarum 519; monan-
drus 516, 519; monospirus 519; muricu-
latus 519; Novae-guineae 516, 519;
Novae-zelandiae 516, 519; Nymanii 519;
pallens 516, 519; parvisporus 516, 519;
salakensis 519; schizophyllus 519;
solidus 516, 519; Stahlii 519; tjiboden-
InpEx TO VOLUME 1 635
sis 519; tosanus 519; Vescoanus 517,
519; vincentianus 517, 519; Wiemanii
519; Zotovii 515
Megachytriaceae 592
Megachytrium 592, 596
Megalocystosporidium 212
Megalosporidium 212
Melanospora 128; vitrea 125
Melobesia 337
Merasperma 253; bifurcata 253; cylin-
drica 253; dichotoma 253
Merisma candidum 268
Merulius undulatus 259
Mesasperma 253
Meteoriaceae 181
Meteoriopsis ancistrodes 183; reclinata
183; squarrosa 183
Michenera artocreas 303
Micrasterias 13, 14, 18, 19; oscitans 14;
pinnatifida 14; rotata 14
Microdictyon 252
Microdus brasiliensis 26; papuanus 26;
pomiformis 26
Micropodia populina 161
Micropyxis 119, 122, 125; Geoglossi
126
Mitochytridium 591, 592, 595
Mniaceae 177, 506
Mniodendron Hellwigii 34
Mnium coriaceum 177; rostratum 177;
Thomsoni 177
Molendoa burmensis 174; Roylei 174
Mollisia 166; tumidula 165, 166
Mollisiella citrinula 600
Mollisina acerina 164, 167, 170
Moniliales 403-407
Monilinia 483
Morchella cava 259; odorata 259
Mortierella 53; tuberosa 51, 66
Mucoraceae 217
Mucronoporus 526; Andersoni 297; cir-
cinatus 526, 527; tomentosus 526, 527
Mutinus 255
Mycastrum 259; siculum 259, 260
Myceliochytriaceae 596
Mycoacia 279; Himantia 278, 279
Mycoderma 213, 219, 480; immite 213,
220
Myriosydrum 253
Myrsidrum 253; Bursa 253; clavatum 248,
253; dilatatum 253; effusum 253; ra-
mosum 253; vermilara 253
Myuriaceae 180
Myurium 46; rufescens 180
Myxophyceae 9
Neckera crenulata 183; himalayana 183
Neckeraceae 37, 183, 507, 508
Neckeropsis nitidula 507
Nemalectra calida 253; fuscata 253;
‘plumosa 253
Nemalionales 495-499
Neokneiffia aspera 283
Neolindbergia falcifolia 35
Neottiospora arenaria 574
Nephrochytrium 591, 592, 595
Netrium digitus 1]
’ Nidularia 259
Niptera sulphureo-tincta 169
Nitzschia closterium fa. minutissima 501;
communis 501; Kiitzingiana 501; Kutz-
ingiana var. exilis 501; ovalis 501;
palea var. debilis 501; palea var. trop-
ica 501; subtilis var. paleacea 501
Nodularia 269; balsamicola 269
Nostoc 253, 391, 392
Nostocaceae 373
Nostocus 253
Nowakowskia 591, 592, 594,
Nowakowskiella 584, 592, 596
Obelidioideae 590, 592
Obelidium 591, 592, 594,
Octoblepharum albidum 173
Odontia 277; Acerina 283; arguta 314;
Brinkmanni 288; fusca 308; Himantia
278; setigera 283; Sistotremoides 273;
tenerrima 99, 100; tenuis 309; vaga
308; vesiculosa 283
Oedogoniaceae 352-356
Oedogonium crenulatocostatum var. cylin-
dricum fa. major 352; Howei 353;
- inconspicuum 353; Kjellmanii 352;
Kjellmanii var. granulosa 352; Koz-
minskii 355; latviense 354; margar-
itiferum 352; microgonium 353;
oelandicum var. contortum 355; ovi-
forme var. gracile 352; polyandri-
um 355; Sawyeri 354; sinuatum fa.
seriatum 354; Smithii 353; sol 354;
spheroideum 353; _ spiripennatum
354; undulatum 355
Oidium 97, 208, 212, 213, 219; candicans
108; Citri 113, 115; coccidioides 212,
220; Curtisii 111; immite 212, 220;
pyogenes 220; protozoides 212, 220;
tomentosum 110
Olpidiaceae 585
Olpidium 13, 587
Omalycus 259; erosus 259; violacinus
259, 260
636 FarLtowlA, Vou. 1, 1944
Ombrophila 139; clavus 139; purpuras-
cens 146; subaurea 137, 141, 142
Onnia 525; circinatus 525-527; tomentos-
us 525-527
Onygena 260
Oocystaceae 357-359
Oocystis apiculata 357; pyriformis 357
Oospora 208
Ophiobolus 395, 416; Aalimus 395, 419;
maritimus 419; medusa 419
Ophiocytiaceae 362, 363
Ophiocytium cochleare 363; desertum .
var. minor 362; elongatum 363; elon-
gatum var. major 362
Opospermum nigrum 253
Orbimyces 399, 404; spectabilis 404
Orimanthis foliacea 253; vesciculata 253
Orthomniopsis japonica 506
Orthomnium trichomitrium 177
Orthorrhynchium philippinense 37; Rid-
leyi 37
Orthotrichaceae 31, 44, 178, 192
Orthotrichum Johnstoni 192; Lyellii 192
Oscillatoria princeps 372, 373
Oscillatoriaceae 372
Ovulinia 483
Oxydontia 279; Himantia 278
Oxytrema 253
Ozonium croceum 304, 309
Pachydisca immutabilis 145
Pannaria lepidiota 392
Papillaria Deppei 181; fuscescens 181;
lanosa 181
Paracoccidioides 214; braziliensis 205,
214
Peckiella Geoglossi 126
Pediastrum biradiatum var. emarginatum
fa. convexum 356; muticum var.
crenulatum 356; quadricornutum
356; tetras 357
Pellicularia 95, 96-118, 264, 272; ansosa
97, 102, 103, 104; asperula 97, 100;
chordulata 97, 98; cystidiata 97,
101, 102; filamentosa 98, 111-113-
116, 272, 282, 305; flavescens 98, 102,
103, 105, 106, 114, 116, 272, 280, 291;
isabellina 97, 99, 272, 309; Koleroga
96-98, 112, 294, 300; Langloisii 97,
101, 309, 319; lembospora 98, 109;
ochroleuca 97, 102-104; pruinata
98, 107, 108, 272, 284; subcoronata
97, 102, 104, 272, 302; vaga 98, 108-
110-111, 272, 305
Penicillium 446
Peniophora 95, 264, 265, 285, 293, 310-
327; admirabilis 310; affinis 318;
alba 310, 311, 324; albo-straminea 264,
281; 291, 306, SLL, 321: albula 282,
295, 312, 316, 323; Allescheri 313, 318;
alutaria 313, 314, 319; arachnoidea
314, 324; argentea 315; argillacea
325; aspera 282-284, 326; asperipilata
315; attenuata 266; aurantiaca 270,
271, 296, 298, 319, 324; Burkii 312,
316; byssoides 275, 276, 308, 314;
byssoides subsp. Tomentella 276, 314;
calothrix 316; canadensis 316; can-
dida 295; chaetophora 103, 3163 cin-
erea 289; coccineo-fulva 326; crassa
320, 326; cremea 313, 314, 315, 324;
cremea subsp. Allescheri 313; cremea
var. Allescheri 313; cremea var. Eich-
leriana 314; cremea var. glauwcescens
314; cretacea 284; crocea 278; delec-
tans 316; diffissa 270; dissoluta 316;
dryina 277-279, 325; duplex 296,
297; KEichleri 320; Eichleriana 314,
315; filamentosa 293; flammea 316,
317; flava 278; fusispora 102, 103,
105; gigantea 317, 328; glebulosa 284,
317; glebulosa subsp. Pirina 316; globi-
fera 317; gracillima 317; Greschikii
105, 310, 311, 324; hastata 311; hetero-
cystidia 317, 318; incarnata 276; 298;
Karstenii 326; Kauffmanii 317, 318;
laevis 318; laevis var. affinis 318;
laminata 313, 314, 319; latitans 283;
lepida 270, 271, 319; limonia 319;
livida 285; longispora 315; longispora
var. clavispora 315; longispora var. cyl-
indrospora 315; longispora var. gloeo-
cystidiata 315; longispora var. myceli-
alis 315; ludoviciana 316; luna 320,
326; lurida 266; magnahypha 101, 319;
media 266; miniata 319; Molleriana
324: muscorum 276; mutata 313, 318;
ochroleuca 102; Odontioides 313; odo-
rata 320, 327; pallidula 313, 314, 319,
324; pallidula var. regenerans 313,
314; Peckii 320, 321; pertenuis 322;
phyllophora 307, 323; Piceina 323;
pilosa 298; Pirina 316; polonensis
316; praetermissa 322; pubera 325;
rimicola 273; Romellii 284, 285; rosea
300; Roumeguerii 324; rudis 311;
Sambuci 325; sanguinea 319; san-
guinea subsp. anaemacta 319; setigera
283; Sheari 270, 271, 324; sordescens
276; sordida 314, 315, 321, 324; sor-
didella 315, 321; sphaerospora 321;
ee ey en re ys 2c ee ee ee Ae ee a ee ae an CGM ee, Wee ee ae, ye a Eee ee ee ne eee
TEETED HGS hy Se PRE ete VOCRT Ge tr orem OmvRr Mametgy (Tee Wer a tart) dls tee eee
InpEx To VOLUME 1 | 637
stratosa 324; subalba 298, 302; sub-
alutacea 103, 266; subapiculata 311,
313, 324; subascondita 324; subcremea
311, 324; subcremea var. subuncinata
311; subincarnata 303; subsulphurea
303, 304, 324; subtilis 325; tabacina
278, 325; tenuis 264, 322; tenuissima
325; Thujae 325, 326; Torrendii 312;
trachytricha 282, 326; velutina 315;
versata 326; verticillata 320, 326
Peniophorella pubera 325
Penzigiella cordata 171, 180
Peritrichospora 398, 414, 448; integra
414, 415, 440, 442-447, 449, 451, 454,
455, 458; lacera 415
Peronospora alta 580
Pexisperma 253; amplectens 253; de-
pressa 253; dichrosperma 253; lutes-
cens 247, 253; sputo 253; truncata 253
Pezicula phyllophila 158
Peziza albida 608; alborufa 260; amorpha
269; atrata 260; caudata 151, 152;
cupularia 260; cupularis 260; Dear-
nessti 613; depressa 260; epiphylla
140; epiphylla var. populina 156, 157;
fucata 609; glagosa 167; globulosa
260; lachnobrachya 162; lupularia 260;
ochrochlora 260; phyllophila 158; pu-
berula 169, 170; pulcherrima 260; punc-
tiformis 160, 163, 164, 166, 169, 170;
smaragdina 260; smeraldina 260; smir-
aldina 260; tumidula 162, 165
Pezizella 137, 166; albopuncta 155; citrin-
ula 600; culmigena 604; Dactylidis
600; epiphylla var. populina 156, 157;
glagosa 139, 167; populina 160; pu-
berula 162, 169, 170; punctiformis 160,
162, 164, 165; punctiformis fa. folii-
cola 162, 164; punctiformis fa. fruti-
cola 165; punctiformis fa. petiolicola
162; tumidula 160, 162, 165-167; tumi-
dula fa. Castanae 162
Phacidium coronatum 163
Phacus anacoelus var. undulata fa. major
366; asymmetrica 366, 367; Birgei
367; chloroplastes 367; chloroplastes
fa. incisa 368; costata 368; crenulata
368; hispidula 367; pseudoswirenko
368; Raciborski 367; Segreti var.
ovum 369; setosa var. crenata 368;
spirogyra 369; spirogyra var. maxima
369; striata 369; Swirenko 369
Phaeociboria 138, 483
Phaeoseptoria 576; Calamagrostidis 576
Phallus 260
~
Phanerochaete odorata 320
Phaxantha lichenoides 254
Pherima 260
Phiala 260
Phialea albida 609; Cassandrae 139, 159, |
617; Dearnessii 613; gemmarum 616;
phyllogena 156; phyllophila 158, 162;
phyllophila fa. fagicola 156; phyllo-
phila var. Jaapii 151, 152; scutula var.
fucata 610
Phialina puberula 161, 169
Phialophora 402
Phialophorophoma 402; __litoralis
403, 442, 444, 445, 447, 450, 451, 454,
463
Philonotis angusta 178; aristifolia 177;
evanidinervis 506; falcata 178; Griffith-
jana 177; imbricatula 178; perlaxi-
folia 178; speciosa 178; Thwaitesii 34;
Turneriana 178
Phlebia 319; coccineo-fulva 326; vaga
308, 309
Phlebriella vaga 309
Phlyctidiaceae 586, 590, 591
Phlyctidioideae 590, 591 |
Phlyctidium 591, 594
Phlyctochytrium 590-595
Phoracis filicina 254
Phorima 260; betulina 260; boletoides
260; coccinea 260; difformis 260; min-
uta 260
Phragmosporae 413
Phycerus bifidus 254.
Phycopteris angustata 339; cuneata 339,
340; Harveyana 340, 341; interrupta
339
Phylictis bifurcatus 254: cervicornis 254;
cuneiformis 254; dichotomus 254; Jati-
folius 254; polypodioides 254; sub-
fistulosus 254; undulatus 254 :
Phyllachora 575; graminis 575; oxalina
573
Phyllops 260
Phyllosiphon 387, 389
Phyllosiphonaceae 388, 389
Phyllosticta 572; guttulatae 573; Heli-
anthi 573; solidaginicola 573, 574;
solidaginis 574
Phyllotylus australis 342
Physidrum aggregatum 254; hyalinum
254; pisiforme 254; rubescens 254
Physocladia 592, 596
Physoderma 569; Claytoniana 569
Physorhizophidium 591, 594
638
Physotris capitata 254; glomerata 254
Physydrum 254
Phytelis atra 254; granulata 254; macro-
carpa 254; radiata 247, 254; sulcata
254; tuberculata 253, 254
Phyxalium 254
Pichia 475, 477, 480; farinosa 477
Piemycus 260
Piesmycus 260; nigrescens 260; violaceus
260
Pilaira 215
Pilobolus 215
Piloecium 46; pseudorufescens 45
Pinnatella ambigua 508; Kuhliana 508;
mucronata 508
Piptocephalis 53, 54
Pisolithus 257; tinctorius 257
Plagiotheciaceae 186, 509
Plagiotheciopsis oblonga 45
Plagiothecium neckeroideum var. javense
509
Plaxarthrus 254
Pleurosporangiatae 498
Pleurotus 124
Podochytrium 591, 594
Pogonatum gymnophyllum 189; inflexum
189; Junghuhnianum 189; macrophyl-
loides 510; microstomum 189
Polychytrium 592, 596
Polyphagoideae 590, 592, 594, 596
Polyphagus 587, 592, 595
Polyporus 528; adustus 155; balsameus
525; betulinus 123; circinatus 525-528;
circinatus var. dualis 528; circinatus
var. triqueter 528; dualis 525-528;
frondosus 123; peakensis 526; Schwein-
itzii 525; tomentosus 525-528; tomen-
tosus var. americanus 527, 528; to-
mentosus var. circinatus 527, 528; tri-
queter 526, 528
Polysaccum 257; pisocarpum 257
Polysiphonia violacea 6
Polystema 260
Polystictus 525; circinatus 525-568; cir-
cinatus var. triqueter 528; tomentosus
527-530, 532, 536; versicolor 123
Polytrichaceae 189, 510
Polytrichum 13; xanthopilum 189
Poria 259; subacida 525, 537
Poronea 260
Porula 254
Posadasia 214, 219; esferiformis 220;
sphaeriformis 212
Potarcus bicolor 254
Pottia flavipes 191; Heimii 192
Fartow1a, Vou. 1, 1944
Pottiaceae 30, 174, 191
Priapus 260; niveus 260
Prionitis 342; Andersoniana 342; austral-
is 342; filiformis 342; linearis 342
Prionodontaceae 35
Protomyces 213, 216-218
Protomycopsis 217, 218
Pseudococcidioides 213, 219; Mazzai 213,
216, 220
Pseudohelotium puberulum 162, 169,170;
punctiforme 160
Pseudohypnella verrucosa 510
Pseudopiloecium 45, 46; scabrisetum
45 |
Pseudopohlia bulbifera 171, 176
Pseudoscinaia 496, 498
Pseudosymblepharis pervaginata 171, 174
Psilopezia mirabilis 267, 268
Psorospermia 211
Pterobryaceae 36, 180, 507
Pterobryopsis acuminata 181; crassicaul-
is 180; caudata 180; flexipes 181;
orientalis 181
Pterogoneurum Roseae 192
-Ptilium crista-castrensis 188
Puccinia 85; Andropogonis 580; Andro-
pogonis var. Polygalina 580; Eleochar-
idis 580; Helianthi 571; Liatridis 570,
571; rubigo-vera 577
Pycnodon asper 283
Pycnopeziza 138, 149; pachyderma 149;
Sejournei 149
Pyrenium 260
Pyrenomycetes 407—420
Pyrisperma 260; hypogea 260
Pyrrophyta 371
Quadrigula 360
Radulum investiens 292, 293; Pini-cana-
densis 323
Ramaria 260
Ramularia Equiseti 578
Remispora 398, 409, 411; maritima
410, 440
Rhacopilaceae 179, 506
Rhacopilum magnirete 506; Schmidii
179, 506; spectabile 506
Rhaphidostichum 46; loriforme 45; Pullei
38
Rhinosporidium 214
Rhizidiaceae 586, 590-594, 596
Rhizidioideae 590, 591, 594, 596
Rhizidiopsis 591, 594
Rhizidium 587, 591, 592, 594
Rhizochrysidaceae 363, 364
INDEX TO VOLUME 1
Rhizoclosmatium 591, 592, 594
Rhizoctonia centrifuga 286, 287; micro-
sclerotia 115; Solani 115
Rhizophlyctis 586, 591, 592, 594.
Rhizophydium 587, 591, 593, 594.
Rhizopus nigricans 51
Rhizosiphon 591, 592, 595
Rhodobryum giganteum 177
Rhodura 497
Rhynchonectria 122
Rhynchostegiella’ menadensis 45; papu-
ensis 44
-Rhynchostegium menadense fa. gracile
38
Riella 261 '
Rimella 261; obovalis 261
Rivularia 391
Robergea 417
Rosenscheldia 573; Heliopsidis 572, 573
Rotula 261
Roya 251
Rozea pterogonioides 185
Rugosaria 258, 261
Russula 128; adustus 124
Rutstroemia 135, 139, 146, 149, 483, 487
Saccharomyces 212, 475, 477; farinosus
477
Saccomyces 591, 592, 595
Samarospora 407, 408
Samarosporella 407; pelagica 408
Sarcanthia 261
Sarcomelas 261
Sarconemus 261
Scenedesmaceae 359, 360
Scenedesmus opoliensis var. contacta 359
Scherffeliomyces 591, 592, 595
Schistomitrium heterophyllum 43
Schizochlamys compacta 348; delicatula
349
Schlotheimia Campbelliana 32; emargin-
ato-pilosa 27, 32; longiseta 32; Mac-
gregorii 32; rubiginosa 32; subrubi-
ginosa 32
Sciadocladus celebensis 34; Kerrii 34;
novae-guineae 34; Wisselii 34
Scinaia 496-498; furcellata 493
Scinaiae 498
Sclernax lutescens 254; truncata 254
Scleroderma 255, 259; flavidum 255;
geaster 255; violacinum 260
Sclerospora 84, 89; philippinensis 88
Sclerotinia 483
Sclerotiniaceae 483-488
Sclerotium quercinum 163
639
Scolecosporeae 415—420
Scopulariopsis 219; americana 214, 220
Scyphophorus 250, 262
Scyphorus 250, 262
Scytonema 391, 392
Sebacina 265, 266, 327, 328; calcea 285;
deglubens 327; deminuta 293; ftbril-
losa 327; incrustans 327; polyschista
327; scariosa 327
Sematophyllaceae 38, 45, 186, 509
Sematophyllum revolutum 38; subhumile
187; tristichulum 187
Septochytrium 592, 596; variabile 596
Septogloeum oxysporum 576, 577
Septoria 575; Andropogonis 576; Andro-
pogonis var. Sorghastri 576; Ane-
mones 980; Cacaliae 581; cylindrica
580; emaculata 576; Nabali 581; Sene-
cionis-aurei 581; Senecionis-sylvatici
. 581; Thlaspii 575°
Septosperma 591, 594
Septotinia 483
Serpula 287
Siphonaria 591, 592, 594
Siphorus alternus 254; fasciculatus 254
Siphytus filiformis 255; hexodon 255;
obconicus 248, 255
Sisostrema 261; globularis 259, 261
Sistotrema 261, 264, 328; coronilla 328;
_ subtrigonospermum 328
Solutoparies 591, 594
Speira pelagica 407
Spermipole effusa 255
Sphaeria 261; Helvellae 127; maritima
413; subulata 122 =
Sphaerococcus vermicularis 343
Sphaeromyxa subulatum 122
Sphaeronema 120, 121, 128; dblephari-
stoma 119, 124, 125; cylindricum 121;
Helvellae 127 ; oxyspora 122; subulatum
119, 120, 122; vitreum 124, 125
Sphaeronaemella 119, 121, 127, 128;
Helvellae 119, 121, 127, 128; oxyspora
122, 123; subulata 122; vitrea 125
Sphaeropsidales 402, 403
Sphaeropsis 261; demersa var. foliicola
574; foliicola 574
Sphaerulina orae-maris 413
Sphagnaceae 25, 172
Sphagnum sericeum 25; subsecundum 172
Spicaria croceum 304
Spinoclosterium curvatum 13
Spinocosmarium 16; quadridens 16
Spirillum desulfuricans 471
640 FarLowia, VoL. 1, 1944
Spirogyra floridana 361; illinoisensis 361 ;
pseudofloridana 360
Splachnaceae 175
Spongia dichotoma 338
Sporodesmium cellulosum 407
Sporonema 575
Sporophlyctidium 591, 592, 594
Sporophlyctis 592, 595
Sporotrichum 211; croceum 304; flavis-
simum 304
Stagonospora 575; Paspali 575; simplicior
fa. Andropogonis 575
Staurastrum 16; Johnsonii var. depauper-
atum 17; muticum 14; punctulatum
17; Rotula 17, 18
Stemastrum 261; Boscii 261
Stephanoma tetracoccum 127
Stereonema 255
Stereophyllum tavoyense 186
Stereum candidum 268; crustaceum 288;
duriusculum 310; insinuans 309; Kar-
stenii 320, 326, 327; Karstenii fa. in-
crustata 326; lepra 268; Murraii 290;
Murraii fa. tuberculosum 290; oderatum
320, 327; Pini 297
Stericium 26]
Stigeoclonium Jubricum 351; pachy-
dermum 350
Stipitococcaceae 361, 362
Stipitococcus apiculatus 361; capense
362; crassistipatus 362 j
Stipiza 261
Stromatinia 483
Stypnion fluitans 255
Syncephalastrum 52, 55, 56; racemosum
54
Syncephalis 52, 54-56, 58; pycnosperma
54, 55; reflexa 55, 56; tenuis 54, 55;
Wynneae 54, 55
Synchytriaceae 217, 585
Synchytrium aureum 580
Syrrhopodon albovaginatus 30, 42; asper
30; Beccari 30; bornensis 30; hispido-
ciliatus 504; Mulleri 43; perarmatus
42, 504; philippinensis 43, 505; stricti-
folius 505 ; tuberculosus 505; Wallisii 30
Taphridium 217, 218
Taphrina lutescens 569
Taxithelium capillarisetum 39; dimorpho-
phyllum 39; glossoides 39; instratum
46; nepalense 39, 187; nitidulum 39;
novae-guineae 39; selenithecium 39
Tayloria indica 175
Tetradesmus Smithii 360
Tetraédron 11; asymmetricum 357;
bifurcatum var. minor 358; cruciatum
var. polyfurcatum 358; cruciatum var.
reductum 358; enorme 358; enorme
var, pentaédricum 358; irregulare
358; limneticum var. gracile 358;
lobulatum var. crassum 359; main-
ensis 11; regulare var. granulata
359; regulare var. incus fa. major
359; tortum 11
Tetraspora 253; lamellosa 348
Tetrasporaceae 348, 349
Thamnium subseriatum 184
Thelephora amorpha 269; aspera 282,
284; aurantiaca 261; bolaris 319; bom-
bycina 295, 296; byssoides 275; calcea
284; calcea * glebulosa 284, 317; can-
dida 268; candidissima 268; centrifuga
286; citrina 304; confluens 275; cre-
tacea 285; epiphylla 323; evolvens 318;
fumosa 308, 309; granulosa 282-284;
incrustans 327; insinuans 309; isabel-
lina 99, 100; lactea 295, 296; laevis
318; Menieri 308; miniata 319; Mur-
raii 290; nigrescens 261; odorata 320;
olivacea 266, 273, 275; pallescens 309;
_ pubera 325; puteana 276; revoluta
261; rosea 299; Sambuci 325; san-
guinea 319; semicampanulata 261;
sera 325, 326; setigera 282, 284; Sisto-
tremoides 273, 280; subochracea 292;
sulphurea 304, 308, 309; umbrina 273,
275, 318; umbrina var. lignatilis 274
Thuidiaceae 184, 506
Thuidium cymbifolium 185; glaucinoides
184; investe 184; Meyenianum 184;
tamariscellum 184
Thyridium 30; gracile 43
Tiarospora 409
Timmiella anomala 174
Tomentella 264, 279, 306; sect. Tomen-
tellastrum subsect. Botrytes 96; albo-
straminea 321; byssina 304; fibrillosa
307; flava 99; fugax 287; granulata
107, 109; isabellina 99; Menieri 308;
obducens 275; ochraceo-viridis 99, 100;
sulphurea 308; testaceogilva 99; trig-
onosperma 307, 308; umbrina 274
Tortella cyrtobasis 174
Trachelomonas hexangulata var. repanda
370; mammillosa 370; __ peridini-
formis 370; superba var. spinosa 370
Trachyphyllum inflexum 185
Trachypodaceae 35, 179
Trachypodopsis auriculata 35; crispatula
180
[INDEX TO VOLUME 1
Trachypus 36; bicolor 179; perplicatus
35
Trametes Pini 530
Tranzschelia Pruni-spinosae 576
Trechispora 264, 328; Brinkmanni 287,
288, 328; coronifera 282, 288; sub-
trigonosperma 328
Trematodon longicollis 172
Tremella 258
Tremellodendron candidum 268
Trentepohlia 391
Treuberia 359
Tribonema 364
Trichia 261
Trichidium 261
Trichoglossum 119; Farlowi 126, 127;
Walteri 127
Trichosporium pedrosoi 205
Trichosteleum Boschii 46; hamatum 46
Trichostomum angustatum 174
Tuber 260; rufescens 261
Tuberium 261
Tucahus 258, 261; albidus 258, 261;
leviusculus 258, 261; rimosus 258, 261;
rugosus 258, 261
Tulasnella 265, 266, 328; bifrons 310;
calospora 328; Eichleriana 328; metal-
lica 272, 273; Violae 328; violea 328
Tylostoma 261
Typhula juncea 605
Udotea flabellata 337; Flabellum 337,
338 :
Ulothrix aequalis 349; cylindricum 349;
moniliformis 350
Ulotrichaceae 349, 350
Ulva tomentosa 253
Urceolella puberula 169
Uredo citrina 261
Uromyces Fabae 576
Vararia 264, 293, 299; alutaria 292; ef-
fuscata 290, 294; granulosa 283; in-
vestiens 292; pallescens 309, 310;
pectinata 298; Peniophoroides 294,
310; phyllophila 323; racemosa 299
Vaucheria dichotoma 255; flexuosa 255;
Gardneri 387-389; stricta 255
Vermilaria 255 —
Vesicularia elegantula 40
Vibrio estuarii 471
Vibrissea 398
Volvaria coccinea 261
Volvox fuscus 255; ovalis 255
Volvycium 261; coccineum 261, 262
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Leen fe ee ee eh ee Seer i
641
Warburgiella 38; leptocarpa 38; papuana
38, 39; subpapuana 38
Webera rubripila 32
Xanthidium 14; antilopaeum 14, 15; anti-
lopaeum var. truncatum 16; antilopae-
um var. Tylerianum 16; armatum
14; cristatum 14; pseudobengalicum
14; subhastiferum 14; Taylorianum 16;
tetracentrotum 14, 15; tetracentrotum
var. quebecense 15; Tylerianum 16
Xanthocapsa 392, 393
Xanthochrous 297; circinatus 526, 527;
tomentosus 526
Xanthophyceae 361-363
Xerocarpus alutarius 292, 293; crustaceus
288; laeticolor 277; odoratus 320, 326,
327; subsulphureus 303
Xylissus 262; -cylindricus 262; lineatus
262; oblongus 262
Yeast G 476
Zarnardinula 342; Andersoniana 342;
australis 342; filiformis 342; line-
aris 342 :
Zonaria 255; cuneata 340; dichotoma
var. intricata 338, 339; flabellata 341;
Harveyana 340; interrupta 339, 340;
multifida 340; spiralis 341; sub-
articulata 339, 340; Tournefortii 340;
Turneriana 339
Zonilia 262
Zygnema pectinata 6
Zygnemataceae 360
Zygodesmus argillaceus 99, 100; pubidus
276
Zygohansenula 480
Zygopichia 469-475-481 ; Chevalieri 471-
475, 477-479; Chevalieri var. Ander-
sonii 472-476; Chevalieri var. Bark-
eri 474-476; Chevalieri var. fermentati
479; chiantigiana 473, 474, 478; fari-
nosa 473-475, 477
Zygorhizidioideae 591
Zygorhizidium 591, 594
Zygosaccharomyces 469-481; bisporus
476; Chevalieri 475-477; farinosus 477:
Guilliermondii 471, 473, 474, 478;
japonicus 479; japonicus var. Soya
4.79; major var. salsus 479
Zymonema 213; immitis 220
Zythia compressa 122, 123; subulata 122
MANUSCRIPT
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