Memoirs of the Torrey Botanical Club.

CONTENTS | t

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A study of the Lactariae of the United States. GERTRUDE SIM- :
MONS BURLINGHAM. Pp. 1-109. Issued Мау: 26, 1908. | :

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A Study of the Lactariae of the United States
GERTRUDE SIMMONS BURLINGHAM
CONTENTS
List oF ILLUSTRATIONS
INTRODUCTION
Scope and aim of the work
Collections of Zacfariae іп the New SE Botanical Garden

Collections е for examination
erbaria v

aen treatment of the genus

Arrangement of the genus in groups and sections

Distribution of species

Effect of latitude and elevation on distribution

Relation of humus to species distribution

РН Q QA ER EMI

Structure and internal morphology

Laticiferous vesse

Probable causes of ‘the change in color of the latex

olor-scheme and photography
SYNOPSIS OF SECTIONS AND GROUPS
EY TO KNOWN SPECIES OF LACTARIA IN THE UNITED STATES
KEYs TO INDIVIDUAL GROUPS AND DESCRIPTION OF EEN INCLUDING SYN-

ONYMS, DISTINGUISHING FIELD-MARKS, AND CRITICAL К
INDEX...
LIST OF ILLUSTRATIONS |
(The photographs are all four fifths of the natural size of the plants)

Бір. 1. Lactaria glaucescens Crossl.......... .......ӛ...-..... SEN
Fig. 2. Lactaria speciosa sp. nov, e:

№. 3. ctaria crocea sp. nov. А тулыр кт
Fig. 4- Lactaria delicata sp. nov DONC CNN VIP obs ree
Fig. 5. Lactaria agglutinata sp. поу........... т. RA
Fig. 6. Lactaria lanuginosa sp. nov ася 52-021
Tu 7. ctaria mucida sp. поу.......... essere PRE NODE MEN UM oS
Fig. 8. Lactaria $ubpurpurea Peck.... а ооо Van uH ee
Fig. 9, то. Lactaria ME Burl ана Тт о
Fig. 11. Lactaria cinerea ен т
Fig. 12. Lactaria "eech Meis KREE
Fig. 13. Lactaria Peckii sp. nov SO Ee тк еба
Fig. 14. Lactaria grisea Peck EE E
Fig. 15. Lactaria isabellina Burl EE eege

Memoirs of the Torrey Botanical Club, Volume 14, Мо. 1.
[Issued 26 Му 1908.

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2 LACTARIAE ОЕ THE UNITED STATES
INTRODUCTION
ScOPE AND AIM OF THE WORK

This study of the Lactariae has been limited to the United
States, since, with the exception of a list of thirteen species from
Nova Scotia,* and a few species from other provinces of Canada
preserved in the herbarium of the N. Y. State Museum at Albany,
no species have been reported from other parts of North America.
In pursuance of the study the aim has been to observe the living
plants in the field. Accordingly, for several summers, collections
were made in the vicinity of Lake Ontario in Oswego County,
New York. One season was spent in southern Vermont near
Newfane, and in the summer of 1907 a study of the southern dis-
tribution of the genus was undertaken in the mountains of North
Carolina about forty miles south of Asheville in the Pisgah Forest
reserve, where the Biltmore Forest School is located during the
summer.

The field study has been supplemented by the examination
of dried specimens collected in various states, many of which col-
lections are the property of the М. Y. Botanical Garden. Among
these should be mentioned the herbarium of the late Professor L.
M. Underwood, which contains species from Connecticut, Central
New York, Indiana, and Alabama ; the Earle collection from Con-
necticut, Alabama, Mississippi, and southeastern New York; the
Lactaria herbarium of N. M. Glatfelter, including the specimens
from which the determinations were made for his St. Louis list t
as well as several species from Pennsylvania; specimens collected
on Mt. Desert Island, Maine, by Miss V. S. White ; species from
Massachusetts collected by G. Morris; collections made in
Maine, Pennsylvania, District of Columbia, Virginia, and Tennessee,
by W. A. Murrill; some Delaware species collected by H. S.
Jackson; collections made near Gainesville, Florida, by H. S.
Fawcett; and specimens collected in Indiana, by G. W. Wilson.
Several other collections were loaned to the Botanical Garden,
thus facilitating the work very much. B. M. Longyear sent for

* MacKay, A. Н. Fungi of Nova Scotia: a provisional list. Proc. and Trans.
N. S. Inst. Sci. 11: 122-143. 1904.

f Preliminary list of higher fungi collected in the vicinity of St. Louis, Mo., from
1898 to 1905. Trans. Acad. Sci. St. Louis 16: 33-94.

LACTARIAE ОЕ THE UNITED STATES 3

examination the type specimens of his two Michigan species,
while Kauffman sent his collection of Lactariae from Michigan.
Through the courtesy of Dr. Bessey, I was enabled to examine the
type of L. vi//osus Clements. Professor L. R. Jones kindly loaned
the specimens of Lactariae іп the herbarium of the University of
Vermont, and Hanmer allowed me the privilege of examining his
entire collection of Lactariae from Connecticut, including the speci-
mens from which the determinations were made in the White list.*

Through the courtesy of Professor Peck, I was greatly assisted
in examining the herbarium of the N. Y. State Museum at Albany.
In addition to the New York specimens it includes species from
several other states. Here are found the Peck type specimens.
Professor Peck also furnished me with additional notes upon some
of these, as will appear under the species. I found that in the
Schweinitz herbarium, which is at the Academy of Natural Sciences
of Philadelphia, not a specimen of ZLac/aria remains. The Herbst
collection of fungi from near Allentown, Pennsylvania, has recently
been turned over to the Academy of Natural Sciences of Phila-
delphia, and, although it has not yet been systematically arranged, I
was able with the assistance of Mr. Stewardson Brown to pick out
the Lactariae which it contains. Only three specimens given in his
list are represented, but there are four in the collection which are
notlisted. Тһе Frost collection of fungi is now accessible at the
University of Vermont, at Burlington. It contains in good con-
dition specimens of /. Aygrophoroides B. & C., Г. scrobiculata
(Scop.) Fr., Г. zusu/sa Fr., and Г. pergamena (Swartz) Fr. ; in poor
condition, Г. ¢rivialis Fr. and Г. theiogala (Bull.) Fr. ; also, four
species indicated as new, one with a brief description. Тһе last are
so covered with mold, however, as to conceal the specific char-
acteristics.

Unfortunately, the specimens from which the determinations
were made іп the lists of Lactariae from some of the western states
have not been preserved. This is true of Bundy's Wisconsin
list of fungi,t Johnson's Mycological Flora of Minnesota,] the

* White, E. А. А preliminary report on the Hymeniales of Connecticut. Conn.
Geol. and es Hist. Surv. Bull. 3: 1-81.

T Geol. of Wis. Surv. 1: 396-4 1883.

+ Ball Minn. Acad, Nat. Sci. a 254-257. 1877.

4 LACTARIAE ОЕ THE UNITED STATES

Pacific Coast Fungi by Harkness and Moore, * and Morgan’s treat-
ment of Lactarius in his Mycologic Flora of the Miami Valley,
Ohio.T The specimens reported from Maryland by Miss Banning
are so perfectly reproduced in her Folio of Maryland Fungi, which
is in the State Museum at Albany, N. Y., that there is no doubt
as to the species which she had in hand, and I have not hesitated
to accept her list. With this exception, the distribution of species
as given in this paper has been based upon the examination of the
specimens themselves.
HISTORICAL
The genus Lactaria was established by Persoon іп 1797.1 Не
defines the genus as follows: “ Pileus fleshy, depressed, lamellae
latex-bearing. If the pileus is broken and the lamellae a little irri-
tated, they pour out a milky liquor for the most part acrid, in which
respect these fungi agree with some higher plants ; and this gives
me a handle so that I am able to separate them under a peculiar
genus from other agarics," etc. Under the genus he describes six
species, four of which are readily assigned; namely, Г. piperata
(L.) Pers., 2. lateritia Pers. [= L. deliciosa (L.) Fr.], L. pallida
Pers., Г. torminosa (Schaeff.) Pers. ; of these, Z. piperata (LJ Tem
stands as thetype of the genus by reason of precedence. In his later
books, Persoon returns to his earlier nomenclature, Agaricus /acti-
ии being used as equivalent to Lactaria, the species being written
trinomially ; nevertheless, to him belongs the credit of first recog-
nizing the genus. Roussel § in 1806 used Lactifluus as a generic
name. Regardless of these two attempts at separating the group
as a genus, Fries classified the Lactariae under Agaricus both in
the Observationes Mycologicae and in the Systema Mycologicum,
in the latter of which he introduced the tribal name Galorrheus.||
In the Epicrisis, f! published in 1838, however, he took up the
generic name Lactarius, the name by which the genus was hence-
forth designated until Schróter recognized the Lactaria of Persoon
in his treatment of the genus in Cohn's Kryptogamen- Flora v von

* Cat. Pacif, Coast F "e * 1880,

T Jour. Cincinnati Soc. Nat. Hist. 6 : 182-185. 1883.

і Tentamen SE ойы fungorum 63-65. 1797.
4 Flore du Calvados 66. 1806.

| Syst. Мус. 1: 61. 1821.

{Т Epicr. 333. 1838.

LACTARIAE OF THE UNITED STATES 5

Schlesien. * And Hennings has followed Schroter in his treatment
of the genus in the Engler & Prantl, Natürlichen Pflanzenfamilien.
Schróter has separated Lactaria plinthogala (Otto) ї and Zac-
taria ligniota Fr. under the generic name Гасатейа | upon the
basis of the yellow spores. But if the spore color is taken asa
generic character, one must include other yellow-spored Lactariae,.
namely L. deliciosa, L. subpurpurea, and related species, as well
as /. delicata, L. crocea, and L. trivialis, which are widely divergent
from Z. plinthogala апа Г. Потоа. Furthermore, Г. Gerardi, а
white-spored species, is so closely related to these two species
as to suggest an origin from them. Hennings, also, has described
one latex-bearing species from Africa under the generic name Lac-
tariopsis,§ the distinctive characteristic being the presence of a mem-
branaceous veil connecting the stem with the pileus. After a care-
ful study of the morphological and physiological characters of the
species forming the genus Гас?а??а as it now stands, I have come
to the conclusion that with the exception of the tribe Реигориз Fr.,
which is not represented in America, there is not sufficient differ-
entiation to form the basis of separation into smaller genera. Тһе
morphological characters which might serve as generic traits are
neither constant enough in any group nor sufficiently limited to the
one group. Taking for example the tomentose margin, we find it
combined with a viscid pileus іп Z. ¢orminosa (Schaeff.) Pers., and
with a dry pileus іп Z. deceptiva Peck; while, if Г. regalis Peck
shall prove to be a valid species or only a form of L. restma Fr., it
will give us in either case two physiologically and naturally related
plants, one with tomentose, the other with glabrous margin. Again,
among the species classified by Fries under Ше Ррегай, we find
glabrous forms like ZL. piperata (L.) Fr., and tomentose ones like
L. vellerea Fr., yet there is no doubt but that these two are very
closely related. In every group except the Piperatae there are
species showing a deepening color in the mature gills, together
with more or less pruinosity. The homogeneity of the genus is,

%3: 534. 1880.
t Agaricus plinthogalus Otto, Versuch Agar. 75. 1816.
Agaricus fuliginosus Ет. Syst. Myc. 1: 73. 1821.
Lactarius fuliginosus Ет. Epicr. 348. 1838.

pt Schles. 3: 544. 1889.
2 Bot. Jahrb. 30: 51. 1901.

6 LACTARIAE ОЕ THE UNITED STATES

then, very strong, and the most pronounced differences are physio-
logical and are lost in the dried specimens.

In the present paper I propose to arrange the species of the
genus in seventeen natural groups, each group being centered about
a spécies which, by reason of possessing in a marked degree the
characteristics found in the other species, shall stand as the type of
the group. In many instances, the first described species has been
found to combine the most of these characteristics. In speak-
ing of this as a natural arrangement, I mean that each group is
made up of those species which seem to be most closely related
both morphologically and physiologically, and often so closely
resemble each other as to be confused by the casual observer.
In many groups the relationship is so marked that it is easy to
conceive of the species as mutants from the type. Itis a grouping
which deals primarily with the living plants.

These groups I have in turn segregated into five sections:
Piperites, Sublimacina, Limacina, Russularia,and Dapetes. Each of
the last two embraces the species included by Fries in his so-
called “‘tribe’’* of the same name. Оп the other hand, the species
which he included under the “ tribe" Piperites, I have classified in
three sections of equal rank with Dapetes and Russularia. The
section Suó/imacina is made up of species possessing characteristics
of both the Piperites and the Limacina. It is not intended that this
grouping shall multiply the names to be used in classification, but
that it shall be an aid in the determination of species and in the
comprehension of relationship.

THE DISTRIBUTION OF THE LACTARIAE IN THE UNITED STATES
AND THE RELATION OF DISTRIBUTION TO LATITUDE,
ELEVATION, AND FOREST CONDITIONS

Some Lactariae have been reported from all of the states east
of the Mississippi River excepting Illinois and Kentucky. In the
region west of the Mississippi, fourteen species have been reported
from Missouri, three from Капзаз.? two from Colorado, one from
Nebraska, and ten from California.t This apparently limited dis-

* According to our - present conception of the term tribe, Fries used it it incorrectly.

He applied the name to each of the four groups into which he divided the genus,

namely: Piferites, Dapetes, Russularia, and Pleuropus. Hym. Eur. 422-439. 1821.
T I have seen only one specimen from Kansas and only a few from California.

LACTARIAE OF THE UNITED STATES 7

tribution of the genus in the West is explained by the fact that in
some of the western states no collections of fleshy fungi have been
made. But enough has been done to indicate that some species will
be found in any state, wherever the forest conditions are favorable.

Within the limits of the United States, latitude seems to affect
the distribution of Lactariae chiefly іп so far as it controls the char-
acter of the forest. Lactaria Indigo Schw., which grows in pine
woods, has been found from Florida to the Adirondack region.
Lactaria deliciosa, L. subpurpurea, апа Г. atroviridis have a like
distribution. Ош of thirty-one species which I found in North
Carolina, twenty-three occur as far north as Vermont, one has a
northern distribution in Europe, Z. Ресей has been found as far
north as Long Island, and six others have not been described
before. 2. salmonea Peck, from Alabama, and L. suġvellerea Peck,
have not as yet been found north of North Carolina.

Elevation also influences the distribution of species in propor-
tion as it produces different ecological conditions. At 500 meters
in Vermont, where the oaks are practically lacking, Г. piperata
(L.) Fr. and ZL. lactifiua L. rarely occur, while at 1,000 to 1,200
meters in North Carolina, where oaks and chestnuts form about
70 per cent. of the forest, these species are very abundant, but
they disappear as one approaches the spruce line. At this altitude
one begins to notice the combined effects of latitude and elevation.
The condition of spruce forests in the southern states differs from
that in the North. Тһе greater elevation of the dividing line
between oaks and spruces causes a greater daily range of temper-
ature and the nights may be too cold for certain spruce-loving
species. Оп this account it is probable that a species like Z. resima
Fr. will be found to be limited to northern forests. In the south-
ern mountains below the spruce line, on account of the great
humidity there will be not only a profusion of fungi but one might
expect to find certain species which would thrive only under such
moisture conditions. It is also possible that the unglaciated con-
dition of the southern states may account for the occurrence of
some species which seem to be limited to that region. This can
not be stated with certainty until further field work proves that
such species as L. speciosa, L. agglutinata, L. salmonea, etc., are
not found north of the glacial line. Not enough is known about

8 LACTARIAE OF THE UNITED STATES

the ZLactariae in California and Oregon to form а basis for
generalization.

The species of trees growing in a forest, then, seem to be the
greatest factor in determining what species of Lacfarta shall grow
there. In oak or chestnut woods one may be sure of finding Z.
lactiffua L. and L. piperata (L.) Fr. ; in beech woods, L. cinerea Peck
and often Г. ve//erea Fr. ; in hemlock woods, Г. deliciosa (L.) Fr.,
L. subpurpurea Peck, апа Г. deceptiva Peck; other species seem
to grow in any fairly moist deciduous forest. This limitation of
species to the vicinity of a particular species of tree is undoubtedly
largely due to the adaptation of the humus there found to the
growth of the fungus. Bourquelot * claims to have found іп Z. con-
troversa Fr., which grows atthe foot of poplars, a ferment analogous
to emulsin, which might indicate a symbiotic relation between the
two. He was not able to find such a ferment in Z. vellerea Fr.
Noack T states that he found a mycorhiza relationship between L.
piperata (L.) Fr. and Fagus silvatica and Quercus pedunculata,
also between L. vellerea Fr. and the beech.

THE STRUCTURE, INTERNAL MORPHOLOGY, AND PHYSIOLOGY OF THE
SPOROPHORE, INCLUDING COLOR CHANGES AND
REACTIONS

The Lactariae and Russulae are characterized by a vesiculose
structure, that is, by the presence of parenchyma-like cells scat-
tered among the filamentous hyphae. Fayod | designates these
tissues as fundamental, consisting of isodiametric sphaerocysts, and
connective tissue or slender hyphae. The sphaerocysts are
grouped in the filamentous tissue, and DeBary § suggested that
they might be enlarged branches of these hyphae constricted at
rather regular intervals, so that the branch might be compared to
a rosary. The sphaerocysts are sometimes grouped so as to
appear in a cross-section of the stem like a rosette, the center of
the rosette being the cross-section of a thin-walled tube which pur-

* Bourquelot, E. Présence d'un ferment analogue à l'émulsine dans les cham-
pignons et en see dans ceux qui sont parasites des arbres ou vivent sur le bois.
Bull. Soc. Myc. 54.

T Ueber darin s t Pilze. "Bot. Zeit. 47: 389. 1889.

f Ann. Sci. Nat. Bot. УП. 9: 322-330. 1889.

4 Morph. Fung. Mycet. and Bact. 298-301. f. 756. 1887. [Engl. transl. ]

LACTARIAE ОЕ THE UNITED STATES 9

sues a winding course through the fundamental tissue. The cor-
tical tissue is made up of the filamentous hyphae and bands of this
tissue run through the interior of the sporophore among the groups
of sphaerocysts. There are also oil-vessels which Fayod * regards
as derived from the connective tissue in a manner similar to the
origin of the laticiferous vessels іп the Рарауегасее. In the Zac-
tartae there is a well-developed system of latex-vessels, and it is the
presence of these that distinguishes the genus Lactaria from the
closely related genus Russula.

The latex-tubes are large thin-walled hyphae about 10-12 pin
diameter running through the bundles of connective tissue and some-
times approaching the groups of sphaerocysts but not coursing
through this tissue. Іп the stem the tubes extend longitudinally
and in the pileus they are for the most part parallel to the surface.
They do not anastomose so as to form a network, but are connected
now and then by short cross branches and in the trama they branch
more freely. Fayod considers these latex-vessels to be modified
sphaerocysts, since they sometimes end among the filamentous
hyphae like sphaerocysts and since it is not uncommon to find
these latter elongated and filled with granular colored protoplasm,
as well as cylindrical with hyaline contents. He also regards the
formation of the latex-vessels to be parallel with their-formation in
the higher plants.

According to Boudier} the latex is an emulsion of a resinous sub-
stance in a liquid which contains albuminoids, and it is this resinous
product which gives to the latex its characteristic taste. Errera
states that the latex-tubes do not contain glycogen. The chem-
istry of this milky fluid has not, however, been sufficiently worked
out. Errera advances the hypothesis that the fundamental tissue
is the place for the deposit of hydrocarbon material while the fila-
mentous hyphae are the routes along which the proteid material
returns to the developing organs. Fayod 5 has also observed that
glycogen is formed in the sphaerocysts and that the material nec-

* Cf. Godfrin, J. Homologie des hyphes vasculaires des agaricinés. Bull, Soc.
Мус. Fr. 18: 147-150.
udier, E. Des champignons au point de vue de leurs caractéres usuels chim-
iques et toxicologiques, 78-82. 1866,
{ Sur le glycogéne chez les Basidiomycétes 30, 31. 1585.
$ Fayod, /ос. cit.

10 LACTARIAE ОЕ THE UNITED STATES

essary for the formation of the glycogen is carried through the
filamentous hyphae and that later the nourishment returns along
the same route to the hymenium as it is needed. Errera further
suggests that the filamentous hyphae are elements of support.
This can be true, of course, only in an elementary way.

The cause of the change in color of the latex of certain species
of Lactariae has been somewhat recently studied Бу Bourquelot
and Bertrand.* They came to the conclusion that the phenom-
ena of coloration are often due to the presence of an oxidizing
ferment in the cells which acts upon a particular chromogen con-
tainedin the mushroom. Only two out of twenty species examined
showed the absence of ап oxidizing ferment. These were L. sub-
dulcis (Bull.) and Г. mitissima Fr. When the latex changes color
this chromogen is in solution in it and the oxidizing ferment con-
cealed also in the solution acts upon the chromogen as soon as
they come together in the air, as in the case of a wound, thereby
changing the chromogen bodies into а colored pigment. It is to
be assumed that in such species as Г. deliciosa (L.) Fr. the mush-
room is so rich in oxidizing substances that this color change will
result in the outer layers of cells. This view is upheld by the fact
that when a leaf adheres firmly to the surface the epidermis is
frequently colorless. The latex in the Dapetes is either colored in
the living plant or the change takes place so instantly when the flesh
is broken that one cannot perceive it. If the color is present in
the latex while within the plant, it is possible that the oxidizing
ferments are able to use the oxygen which enters the plant by res-
piration. If this theory should be true we might expect variations
of color under different temperatures and light conditions,+ since
these factors affect the respiration of the mushroom. But while
it has been shown that light diminishes respiration, certain species
are brighter-colored when growing in open woods. Z. theiogala

* Bertrand, G. Bull. Soc, Chim. Paris 111. 15: 793. 1896.

Bourquelot, E. & Bertrand С. Les ferments oxydants dans les champignons.
Bull. Soc. Myc. Fr. 12: 18-26. 1896; Sur la coloration des tissus et du suc de cer-
tains champignons au contact de l'air. Bull. Soc - Myc. Fr. 12: 27-32. 1896.

Bourquelot, E. Sur la présence générale, dans les champignons, d'un ferment
oxydant agissant sur la tyrosine ; sur le mécanisme de 1а coloration du chapeau de ces
végétaux. Bull. Soc. Myc. Fr. 13: 65-72. 1897.

f Bonnier & Mangin. Ann. Sci, Nat. Bot. VI. 17:210-302. 1884.

LACTARIAE OF THE UNITED STATES 11

(Bull. Fr. is nearly white when growing in dense shade but in a
more exposed place it varies from yellowish-salmon to fulvous.
In Г. /actiffua, however, which varies greatly in color, I have not
been able to detect any fixed relation between coloration and light.
It is thus probable that color depends upon the interaction of a
number of factors, and any one factor may not have the same
effect upon color in two different species. Іп Tricholoma Rus-
sula, Bourquelot * was unable to find an oxidase ferment, yet color
changes occur in the broken flesh, hence it seems possible that in
some cases the action of the air alone is sufficient to cause a color
change. Thus far, two oxidases have been identified in the mush-
rooms, tyrosinase and laccase, both of which occur in the Lactar-
iae. No work has been done along this line upon the American
species of Lac/aria, and much investigation is needed before the
chemistry and physiology of the genus will be understood.

The accompanying analysis of three species of Lactariae by
Margewicz is taken from Schenck's Handbuch der Botanik (4:
391. 1890) The analysis was made from fresh young speci-
mens. Bourquelot has made a study of the carbohydrates in sev-
eral species of Lac/aria. With the exception of 2. /асийиа (L.),
which contained volemite, mannite was present in all the species
which he tested in the dried state. He found that the proportion
of carbohydrate varies in different plants and in different specimens
of the same species and that drying produces important differences
both in the proportion and the nature of the substances. In Г.
piperata the trehalose disappears and is replaced by mannite. In
other species the glucose which is present as a trace in the fresh
plant is found in large proportions in the dried plant. Bourque-
lot's + analysis differs from that of Margewicz. From Gerard’s {
studies of the oils in 2. ve//erea апа Г. piperata, we find that the
former contains stearic acid, and the volatile acids, formic, acetic,
and butyric, while oleic acid is present in a free state. Lecithine
is probably present.

* Bull. Soc. Myc. Fr. 13: 65-72. 1

+ Bourquelot, Е, Matiéres sucrées continues dans les champignons, Bull, Soc.
Myc. Fr. 8: 196-208. 1892; Les matiéres sucrées chez les lactaires. Bull. Soc.
Myc. Fr. 5: 143-163.

E

1880.
i Sur les matières grasses de deux champignons. Bull. Soc.
Myc. Fr. 6: 115-128. 1890.

12 LACTARIAE OF THE UNITED STATES

In fresh plant. In the dry substance,

Analysis,

Solid. Water. | Proteid. | Fat. Маппие. | Sugar, Ash. Cellulose. ` Waste.

17 89.83 39.14 5.34 13.14 | 1.98 | 7.37 | 28.93 | 4.10

82 91.18 26.37 401 15.71 | 4.31 | 5.27 | 38.86 25.47

; 83 90.17 | 32.21 |6.01| 13.47 | 4.17 | 7.13| 30.30 | 5.81

2. piperata 77| 88.23 | 28.35 |4.72 12.17 | 4.13 | 8.43| 38.04 | 4.16
OI

|
о. 89.99 | 38.12 |7.37 12.91 | 1.49 | 8.14 | 27.42 | 4.55
8.90| 91.10 | 27.00 |4.20| 11.57 | 5.27 |848 | 39.27 | 4.21

стела шей
L. torminosa |

L. deliciosa

P pileus. S— stem.

ECONOMIC VALUE

So far as is known, none of the Zactariae are seriously poison-
ous, although some are considered somewhat poisonous, while a
large number of the species are edible. Whenever the edible
qualities are known, I have indicated it at the end of the descrip-
tion of the species, following, in every instance, the conservative
opinion.

S NOMENCLATURE

In the matter of nomenclature, I have followed the rules of
the American Code.* According to this, under canon 16 (c),
Lactarius Fr. is to be regarded as a homonym in the sense that it
is a variation from Lactaria Pers. merely in the spelling of the
word ; so that while the latter form must be retained because of
priority, the name Lactarius is considered as practically identical
with it, and the transfer of a species from Lactarius Fr. to Lactaria
Pers. is not to be looked upon as making new combinations. But
in all cases I have attempted to show which form was used by the
one describing the species, as well as to indicate any subsequent
correction. Forexample, “Zactaria pergamena (Swartz) Fr. Epicr.
340. 1838 [as Lactarius]. — Schrót. in Cohn, Krypt.-Fl. Schles.
3: 537. 1889," would. be explained as follows : The species was
described by Swartz under the specific name pergamena, trans-
ferred from Agaricus to Lactaria by Fries under the form Lactarius
pergamenus, and later corrected by Schröter to read Lactaria
pergamena,

* Bull. Torrey Club 34: 167-178. 1907.

LACTARIAE OF THE UNITED STATES 13

COLOR-SCHEME USED, AND EXPLANATION OF PHOTOGRAPHIC
METHODS

In the description of the colors of the mushrooms, I have used
the Répertoire de Couleurs published by the Société Frangaise des
Chrysanthémistes, which contains 365 color-plates, most of the
colors being produced in four tones. The plates represent living
rather than metallic colors and are for this reason admirably
adapted to the use of the mycologist. Іп the description of species
the numbers in parenthesis after the color refer to the plate and
tone in the Répertoire.

For the accompanying plates perfectly fresh representative
plants were photographed four-fifths natural size on orthochro-
matic plates.

The specimens photographed were all collected in the “ Pink
Beds " region, in the Pisgah Forest reserve, about forty miles south

of Asheville, North Carolina.
: Much об this work was completed under the direction of Pro-
fessor L. M. Underwood, whose assistance and inspiring encourage-
ment I am glad to acknowledge. Many thanks are due also to
Professor F. S. Earle for reading the manuscript.

14 LACTARIAE OF THE UNITED STATES

Order AGARICALES
Family AGARICACEAE
Genus LACTARIA

Sections, PIPERITES, SUBLIMACINA, LIMACINA, RUSSULARIA,
AND DAPETES

Groups, 17, as in following synopsis of sections and groups,
etc.
Synopsis of sections and groups
* Latex white at first (except іп 2. sa/monea іп XV)
а. PIPERITES, Gills becoming neither darker with age nor pruinose; latex usually
crid,
Pileus without a pellicle, absolutely dry.
Color white ОТ. РІРЕКАТАЕ.
lor dark. II. RUSTICANAE,
Pileus viscid when wet.

argin involute, densely tomentose or villose. III. TORMINOSAE.
B. SUBLIMACINA. Gills becoming somewhat deeper-colored with age and pruinose ;
pileus very viscid, т involute and more ог less downy-tomentose at first ;
latex less acrid than in
Latex white, желш У. AGGLUTINATAE.
Latex becomin, ow, V
Latex or wounds becoming lilac. VI. ASPIDEA
у. LIMACINA. Gills unchanged with age or sometimes slightly darker іп 7) ыша
and slightly ргиїпозе; pileus very viscid, margin absolutely naked; latex very
crid,
Pileas some shade of gray or brown or red. VIII. TRIVIALES.
Pileus some shade of yellow VII. INSULSAE.
6, RUSSULARIA. Gills decidedly darker when mature, and pruinose; latex mild or

ily acrid except in 2. rufa and 2. chrysorhea.
Pileus viscid when wet, never slimy, soon dry.
vm d т yellow ог red.

us grayi XI. CINEREAE.
па pale EE to fawn or mahogany. X. QUIETAE.
Latex becoming yellow, XII, THEIOGALAE.
Pileus not viscid when wet.
Surface glabrous, polished, or sometimes areolate.
esh thin, stem slender, XVII. CAMPHORATAE.
Flesh thick, firm, stem stout. XVI. LACTIFLUAE.
Surface pruinose to velvety.
r some shade of fulvous. XVI. LACTIFLUAE.
Color white or some shade of brown. XV. PLINTHOGALAE.
Surface minutely tomentose or Squamulose.
Color some shade of gray, XIV. GRISEAE.

Color Metam to fulvous or isabelline. XIII. HELVAE.

LACTARIAE OF THE UNITED STATES 15

** Latex bright-colored from the first

| г. DAPETES. Pileus тараны viscid when wet, spores yellowish, latex not very

| acrid ; plants е
Wounds becoming оын at least after some time. ІХ. DELICIOSAE.
Wounds not becoming greenish. ХУ. L. salmonea.

Key to the known species of Lactaria in the United States *

mu wu

Latex bright-colored from the first. А
atex white, unchanging. B
Latex becoming yellow, at least where in contact with the flesh. C
Latex becoming lilac, at least where in contact with the flesh. D
Latex becoming к or reddish, at least where in contact with the flesh. E
Latex becoming glaucous-gree F
Latex bright-colored from the first
1. Pileus more or less zonate. 2
Pileus azonate, whitish, latex salmon-colored. 57. L. salmonea,
2. Latex indigo-blue or paler 242 igo.
Latex dark-red. за. L. Дери: cram
Latex orange- terri then red-orange. . L. deliciosa.
Latex saffron-yell 31. 1. EC E
Latex white, unchanging
Pileus never viscid,
Pileus viscid when young or wet. %П
| 1
| 1. Latex very а 2
1 Latex mild or ae acrid. II
| 2. Pileus white. z 3
1 Pileus colored. 8
3. Surface glabrous. 4
Surface not glabro 6
4. Margin naked, plants large 5
Margin silky, plants small. 7. L. involuta.
3 5. Flesh tbin, gills becoming straw-colored, 1 mm. broad. 2. L. pergamena.
| Flesh thick, gills dichotomously branching, 2 mm. broad. I. Г. piperata.
| 6. et velvety-tomentose.
glabrous or torn, margin cottony-tomentose. 5. L. deceptiva.
7. Gifs ы. ‘oad. 3. L. vellerea.
Gills close, narrow, cinnamon-colored in drying. 4. Г. subvellerea.
8. Pileus glabrous, at least when mature. 9
Pileus scabrous-hairy, olivaceous. g. L. atroviridis.
9. Red. IO
Gray, mixed with brown, zonate, moist. 8. L. rusticana.
то. Umbonate, 44. L. rufa.

* [n the keys preceding the groups will be found the doubtful American species and
the European species which have been reported from the United States, but the occur-
rence of which is uncertain

LACTARIAE OF THE UNITED STATES

Without umbo. 47. L. rufula.

. Pileus iiie 12
ileus squamulose to floccose-squamulose. 2I
Pileus velvety, or minutely tomentose. : 26

. Gills not changing color where wounded. 13
Gills Sege color where wounded. 18

. Pileus fulvous to isabelline 14
Pileus Ке stem and gills concolorous. 53. Z. Sumstinei.
Pileus zonate when moist. 69. L. mutabilis.
Pileus azonate. I5

IS. Latex white or sometimes bluish-white to watery and thin. 16
Latex whey-colored, pileus fading. 67. Г. seriffua.
16, Pileus polished, not fadin I
ileus becoming areolate, fading, latex watery. 65. Z. rimosella.
17. or none. 66. L. subdulcis.
Aromatic, stronger in drying. 64. L. camphorata.
18. Gills becoming greenish where wounded. 19
ills becoming brown where wounded. 20
I9. Pileus about 3 cm. broad, umber-colored. 70. L. parva.
Pileus up to 7.5 cm. bind, grayish, often roughened. 71. L. varia.
20. Pileus azonate. 58. Г. /actifiua.
Pileus zonate. бо. L. ichorata.
21. Pileus fulvous to isabelline.
Pileus gray. 2
22. Odor none, latex white. 46. L. alpina,
Aromatic, latex watery. 43. L. helva.
23. Odor none. 24
ie бо. Г. glyciosma. |
24. Pileus = floccose-squamulose. gris
Pileus zo 25
25. Plants very weg? blue-black to slate-colored, stem glabrous. 49. Z. Bensleyae.
Plants larger, paler, stem pubescent. 51. L. Hibbardae.
26. Pileus azonate. 27
Pileus zonate, brick-red. 45. L. Pec£ii.
: 27. Wounds becoming brown or fulvous. 28
Wounds not changing color. 29
28. Pileus yellowish-buff, gills close, 62. L. luteola.
Pileus Vandyke-brown, corrugated. бі. L. corrugis.
29. Pileus brown, Е distant. 55. L. Gerardit.
Pileus pe fulvo 31
30. Gills distan wë? i. асана
; . L. subvelutina
& II. Pileus viscid when young or wet
I. Latex acrid.
Latex mild or slowly acrid.
2. Margin of pileus involute at first, with long tomentum
Margin of pileus involute at first, with tomentum дш I mm. long, soon naked.
Margin of pileus naked, pileus g 1
3.

Pileus whitish, entirely covered vis matted tomentum, d nd cilicioides. 3
Poni er, zonate. L. torminosā. `

ке їп м Center

Е.

d

LACTARIAE OF THE UNITED STATES 11

4. Pileus white, serm sparsely covered with coarse tomentum. 18. Z. lanuginosa.
l Pileus buff, papillate when wet, squamulose when dry. 17. L. agglutinata.
| Pileus olivaceous- SS 19. Г. /urfi.
5. Pileus die owish to orange.
| Pileu р 7
| Pileu et аго 28. Г. hysgina.
| 6. Zonate, spores тірі 24. L. insulsa.
l Azonate, spores white. 25. Г. affinis.
| 7. Wounds of the gills becoming glaucous-green. 8
Wounds not changing, pileus zoned with umber. ing. 27. L. circellata.
8. äs, gray to OE sometimes lilac- og ний yellow. 26. Г. trivialis
sepia, spores w 20. Г. doen
9. ак gray, gills not sui color where wounded. 39. L. cinerea.
ileus red. то
Pileus fulvous to pale-isabelline, plants small. II
Pileus brownish, obscurely zonate, fading. 38. L. paludinella.
IO. Flesh thick, pileus 4. L. quieta.
Flesh thin, pileus ses dh йд when moist. 35. L. nitida.
11. Umbo persisting dark, latex mild. 6. L. oculata.
Umbo concolorous, margin aca milk acrid. 37. L. minuscula.
с
Latex becoming yellow, at least where in contact with the flesh
1. Margin of young pileus with long tomentum. 2
Lt e ша pileus glabrous ог minutely tomentose.
2. Pileu 13. L. resima.
Pilens yellowish, zonate, stem with bright pits. 12. L. scrobiculata.
3. Latex mild. 68. L. isabellina.
Latex slowly acrid or bitterish. 4
tex acrid. 6
4. Pileus та pinkish-buff to fulvous. до. L. theiogala.
Von mate. 122%
5. тын ish, not becoming perceptibly darker. до. L. theiogala.
Заря whitish, then reddish-brown. ді. L. colorascens.
6. Pileus umber-brown, subtomentose. . 56. Г. subtomentosa.
Pileus some shade of yellow, glabrous. 7
1 7. Pileus very viscid when moist. 8
3 Pileus scarcely viscid, pallid, zoned with yellow spots. 42: 4. angele
1 8. Pileus saffron-yellow, latex becoming yellow-cadmium. 15. Г

Pileus ое, в salmon-tinted, latex becoming арыы ылу
. L. delicata.

Latex becoming lilac or heliotrope, at least where in contact with the flesh

I. Pileus covered with long tomentum, zonate. 14. L. speciosa.

Pileus glabrous 2

[ 2. Gray 3
| Yellow. 4
3- Azonate or faintly zoned, stem not spotted. 22. L. lividorubescens.

Conspicuously zoned, stem spotted, plants large. 23. L. maculata.

18 LACTARIAE OF THE UNITED STATES

4. Straw-colored, azonate, latex acrid. 20. 2. aspidea.
Sulphur-yellow, zonate, latex bitter. 21. L. aspideoides.

ex becoming salmon or reddish, at least where in contact with the fles
ти glabrous, not сөне ог zoned, 52. Г. jw
Pileus velvety, seal-brow 54. Г. ligniota.
Latex becoming glaucous-green * `
I. Pileus white, dry, latex very acrid. 6. L. glaucescens.

DESCRIPTIONS OF THE SPECIES, INCLUDING KEYS,
ARRANGED BY GROUPS
I. PIPERATAE

Pileus without a pellicle and absolutely dry, glabrous of
tomentose, flesh firm; plants for the most part large; gills not
becoming darker with age nor pruinose ; latex very acrid, white,
unchanging except in Lactaria glaucescens, in which it dries glau- |
cous-green.

In one species, Lactaria deceptiva, the margin is very involute,
and is covered with a roll of tangled fine cottony tomentum as in |
the Zorminosae.

Synopsis of species
Latex sip unchanging. :

Bien di abrou
Pileus pud gills very close, about 1 mm. broad. 2. L. pergamena.
Pileus thick ; gills dichotomously forking, 2 mm. broad.

т. L. piperata.
Pileus velvety-tomentose.

з. L. vellerea.
Gills close. 4. Г. subvellerea.
Pileus with a cottony roll оп margin, surface glabrous or torn
. L. deceptiva.

Plants small, pileus up to 5 cm. broad ; margin involute, minutely silky.
involuta.

Latex drying glaucous on broken flesh and gills. 6. b glaucescens.

I. ГАСТАВТА РІРЕКАТА (L.) Pers. Tent. Disp. Meth.
Fung. 64. 1797
Agaricus piperatus L. Sp. Pl. 1173. 1753; Fl. Suec. 373. 1745"
Not Agaricus piperatus а and В, L. Fl. Suec. 441. 1755.
Agaricus Listeri Withering, Nat. Arr. Brit. Pl. 4: 156. 19801.
[Ed. 4.]
* See also B 2 I, 19, and В II, 9.

LACTARIAE ОЕ THE UNITED STATES 19

Pileus fleshy, compact, convex-umbilicate, at length infundib-
uliform, white, azonate, dry, glabrous, 4—12 cm. or mere in diam-
eter, margin involute at first and naked, at length uplifted; gills
white or creamy-white, forking dichotomously, close, more or less
decurrent, arcuate at first, then extending upwards, only about 2
mm. broad ; stem white, equal, dry, often pruinose, solid and firm,
2-8 cm. long, up to 2 cm. thick, flesh white, unchanging or
becoming sordid ; spores white, subglobose, nearly smooth, 8-0 и;
latex white, unchanging, very acrid, abundant. Ва.

Has.: In oak woods or groves. July to October.

DISTRIB.: New York, Peck, Earle, Burlingham 20, 1905;
Connecticut, Underwood; New Jersey, Sterling ; Pennsylvania,
Mrs. Dallas; Maryland, Banning ; District of Columbia, Murrill,
1403; Virginia, 600 meters elevation, Murrill 272 and 377;
North Carolina, Burlingham,; South Carolina, Ravenel; Tennes-
see, 500 meters elevation, Murrill 517 В; Alabama, Underwood ;
Indiana, С. W. Wilson; Missouri, Glatfelter 345. It has also
been reported from the following states: Maine (White) ; Vermont
(Frost); Rhode Island (Bennett); Ohio (Morgan); Minnesota
(Johnson); Wisconsin (Bundy) ; California (Harkness & Moore).

Плозт.: Atkinson, Stud. Am. Fungi, /. 119; Banning, Folio
Md. Fungi, ai 83 ; Вапа, Champ. Nice, £l. 22. f. 1-5 ; Bel. Champ.
Tarn. ai 22; Bern. Champ. Roch. AX. 57. f. 2; Bolt. Geschichte,
pl. 21; Boyer, Champ. Comest. е: Ven. Fr. p/. 30; Britz. Lact.
f. 24; Bull Herb. Fr. pl. 200, Agaricus acris; Cooke, Br.
Fungi, ai 979; Cordier, Champ. Fr. pl. 28. f. г; Eng. & Prantl,
Nat. Pflanzenfam. 1'**: 7 110В; Fl Dan. M. 1132; Fries,
Sverig. Svamp. р/. 27; Hahn, Der Pilz-Sammler, f. 19, ed. 2;
Harzer, Pilze, ai 39; Krombh. Abbild. ai 56. f. 1-4; Lanzi,
Fung. Mang. AM. 54. f. 2 a, 6, c; Lorinser, Essb. und Gift.
Schwàm. A. о. f. 4; Mcllvaine, One Thous. Am. Fungi, ai 47.
f. 1.; Pat. Tab. Analyt. Fung. A. 110; Paulet, Traité Champ.
pl. 68. f. 3-2; Roumeg. Crypt. Illustr. Æ 144; Rich. & Roze,
Atl. Champ. AX. 40. f. 5-8 ; Schaeff. Fung. Bav. Icon. AM. 83,
Agaricus amarus; Sicard, Hist. Nat. Champ. ai 44. J. 225)
White, Conn. Geol. and Nat. Hist. Surv. Bull. 3: ai 9.

Exsic.: Ravenel, Fungi Caroliniani, fasc. 2, 5 р. Р.; Roume-
guére, Fungi Gallici 3679.

DISTINGUISHING FIELD-MARKS: The glabrous, white pileus, the

20 LACTARIAE OF THE UNITED STATES

close, dichotomously forking, rather narrow gills, and the thick |
flesh. Usually the latex remains entirely white, and the flesh is |
odorless.
In North Carolina I found plants agreeing in all other essentials |
with Lactaria piperata except that the latex dried a pale-yellowish, |
and the fresh plant when wet or when rubbed had the odor of A
crushed blackberries, and the gills were slightly less crowded.
This can scarcely represent more than a form of the species and |
on account of the odor, which is the distinguishing characteristic, I
will refer to it as form fragrans. It is wo, 79, 1907, of my North
Carolina plants. Gillet recognizes a form атаға, іп which the |
milk becomes yellowish іп drying, but the plant is odorless.
In Species Plantarum 1173. 1 753, Linnaeus gives as synonyms 2
of his piperatus the following : Amanita piperata alba, Dill. Giss. |
179; Fungus piperatus albus acris, Mich. Gen. 141; Haller, 34. n. |
І; and Fungus albus acris, C. Bauhin 371. No mention was made
by any of these writers of incarnate gills, Micheli describing his |
piperatus under unicolores albi, and Haller under gills white. Since |
the Дгфегай referred to by Linnaeus are all described as white, and |
no mention is made of zones or tomentose covering, this character |
of the gills seems scarcely sufficient evidence to indicate that E
Linnaeus had in mind Agaricus piperatus var. а and В, Fl. Suec |
441. 1755, when he described Agaricus piteratus in Fl. Suec. 375: —
1745, and Sp. Pl. 1173. 1753. If it were shown that the Agar-
icus piperatus L. Sp. Pl. 1173. 1753, were the same as Agaricus
piperatus L. Fl. Suec. 441. 1755, var. a and В, then Lactaria tormi-
nosa (Schaeff.) Pers. would be the real Zactaria piperata and Гас
taria torminosa would be reduced to a synonym. For, according |
to the present codes, Agaricus piperatus must be credited to Lin-
naeus whatever the type may prove to be, so there is no ground
for crediting Scopoli with the name, and, furthermore, Scopoli gives
Agaricus piperatus Sp. Pl. 1 173 and Fl. Suec. 441. 1755 as syn
onyms of his own piperatus. :
In conclusion, it seems most probable that the forms included |
under var. а and 3. Fl. Suec. 441. 1755, were found later by Lin-
naeus and incorrectly referred by him to Agaricus piperatus aS `
described in Sp. РІ. 1173. 1753, which was undoubtedly the |
esculent fungus piperatus of the ancient writers and the German |

“ Pfefferling."

Ера зуы а

LACTARIAE ОЕ THE UNITED STATES Si

2. LACTARIA PERGAMENA (Swartz) Ет. Epicr. 340. 1838. [As
Lactarius.] — Schrót. in Cohn, Krypt.-Fl. Schles.
3:.537.. 1889
Agaricus pergamenus Swartz, Kongl. Sv. Vet.-Akad. Nya Handl.

30: 90. 1809.

Pileus fleshy, pliant, thin, convex to plane or depressed, often
irregular and eccentric, and usually flexuous, white to creamy-
white when old, azonate, dry, glabrous, minutely wrinkled, 4-11
cm. broad ; gills white, then straw-colored, very close, thin, fork-
ing, adnate, horizontal, very narrow (г mm. broad); stem white,
equal or tapering downwards, glabrous, smooth, stuffed} but firm ;
spores white, minutely echinulate, elliptical, 8 x би; flesh white ;
latex white, abundant, acrid. 204е.

Нав.: In dry oak woods. August, September.

DISTRIB. : New York, Peck, Burlingham (120 meters elevation);
Vermont, Frost; Connecticut, Earle; Virginia, Murrill; North
Carolina, 1,000 meters elevation, Burlingham 39, 1907 ; Alabama,
Earle ; Ohio, Morgan, Dawson ; Tennessee, 500 meters elevation,
Murrill 517; Mississippi, Eae.

IrLusr.: Batsch, Elench. Fung. M. 73. f. 59, а, 6, с. Agaricus
piperatus; Britz. Lact. f. ro; Cooke, Br. Fungi, ai 978; Gillet,
Champ. Fr. A. 152 [389]*; Krombh. Abbild. ai 57. f. 1-6;
Lucand, Champ. Fr. ai 42. l

Dist FIELD-MARKS: This species is very closely
related to Lactaria piperata, and is often confused with it. Accord-
ing to Fries it differs from Lactaria piperata in having a stuffed
stem which at length becomes spongy, and is longer, tapers down-
wards, and is perfectly glabrous; in the thinner, pliant, elastic,
often irregular and eccentric, and for the most part, flexuous pileus,
which is at first convex and not umbilicate, then rather plane, and
wrinkled in a peculiar manner ; in the gills being adnate, very close
and very narrow, always straight and plane, and soon becoming
straw-colored. The latex is more abundant in the flesh than in
the gills.

Lactaria pergamena is not so common as Lactaria piperata.
The latex sometimes dries sulphury-white (74. 2. 2).

* The plates in Gillet’s Champ. Fr. were renumbered in 1890 and again іп the
final edition. In citation I have given the original number and in brackets the numbers
which occur in the subsequent revisions.

99 LACTARIAE OF THE UNITED STATES

3. LACTARIA VELLEREA (Fr.) Fr. Epicr. 340. 1838.

[As Lactarms.]. — Schrót. in Cohn, Krypt.-Fl.

Schles. 3: 538. 1880

Agaricus vellereus Fr. Syst. Myc. т: 76. 1821. |
Pileus fleshy, thick, firm, convex-umbilicate, with inrolled, then |
extended margin, depressed in the center, whitish, tinged in places |
with grayish or yellowish, dry, minutely tomentose, 6-11 cm. A
broad ; gills whitish, becoming cream-colored ог yellowish, chang- =
ing to brownish with injury, many forking near the stem, inter- |
veined, shorter gills often joining with longer, distant, appearing |

more so with age or in larger specimens, adnate to slightly decur-

rent, 2-5 mm. broad; stem whitish, tinged in places with yellow,
equal or rounded at the base, pruinose-velvety, 3-4 cm. long, |
18-25 mm. thick ; flesh white at first, staining yellowish; spores |
white, globose to ovate or elliptical, with only a few scattering |
blunt spines, uniguttulate, 5-6 » x 8 и ; latex white, acrid, unchang- |
ing, or if creamy soon becoming white again. Regarded as -
poisonous by some. 2
Нав.: Open deciduous woods. July, August, September. -
Півтків.: New York, Peck, Earle, Burlingham, Peck & Earle |

(Long Island) ; Vermont, Burlingham 49, 1906; Maine, White; |
Connecticut, Underwood, Hanmer 153; New Jersey, Earle; Vir- E.
ginia, Murrill 135; North Carolina, 1000-1200 meters elevation, |
Burlingham ror, 1907 ; Tennessee, 300 meters elevation, Murrill. :
526; South Carolina, Ravenel ; Alabama, Earle. 3
Irrusr. : Вапа, Champ. Nice, //. 22. f. 6-9; Bres. Fung. Mang. |
A 67; Britz. Lact. f. 26; Bull. Herb. Fr. pl. 558. f. G, Н, Ni ©

Cooke, Br. Fungi, ai 980; Cordier, Champ. Fr. ai 28. f. 27 -
Gillet, Champ. Fr. ai 153 [400] ; Hahn, Der Pilz-Sammler, ей. |
2. pl. 4. f. 17; Hussey, Illust. Br. Myc. 1: pl. 63; Klotzsch, ~
in Dietrich, Fl. Вог. ai, 469; Krombh. Abbild. pl. 57. f. 10-135 №

Lanzi, Fung. Mang. ai 54.7. 1; Lorinser, Essb. und Gift. Schwam. _
pl. 9. f. 5; Phoebus, Deutschl. Krypt. Giftgewachse, РА 4. А 3» -
4; Rich. & Rose, Atl. Champ. ad ҙа. f.1—3 ; Sicard, Hist. Nat.
Champ. 7/45. f. 239; Sowerby, Eng. Fungi, pl. год, Agaricus
Listert ; White, Conn. Geol. and Nat. Hist. Surv. Bull. 3: ai 9:
Exsic. : * Ravenel, Fungi Caroliniani, fasc. 2, 5p. p. ; Roume-

guère, Fungi Gallici 5579.

teo Pt

1

* In the Fungi Caroliniani Exsic. of Ravenel, fasc. 2, no. 5 in the ‘herbarium
of the N. Y. Botanical Garden, there are two specimens of Z, ve//erea Fr. оп the sheet
with 2. pizerata (L.) Fr.

|
|

LACTARIAE OF THE UNITED STATES 28

DISTINGUISHING FIELD-MARKS: Lactaria vellerea may be dis-
tinguished from Lactaria piperata (L.) Fr. by the short tomentum
on the pileus. In typical forms this is dense and gives the sur-
face a very velvety feeling, but it is sometimes less prominent.
The gills also differ in being distant and they stain brownish
where wounded, while the milk dries in gummy drops on the gills.
The gills do not for the most part fork dichotomously as in Lac-
taria piperata, but the shorter ones anastomose with the longer
ones. While the latex is practically unchangeable, it is noted by
Earle as becoming cream-colored (herb. N. Y. Bot. Garden 789,
826, 1906). Massee says the latex sometimes becomes pale-sul-
phur when the flesh is broken but soon changes to white.

Up to the time of Fries, this mushroom was confused with
so-called piperatus forms. Agaricus piperatus described by Pol-
lich (Fl. Pal. 3: 289) in 1777, is very clearly Г. vellerea of Fries.
This is the only description prior to Fries which makes mention
of the tomentose covering of the pileus. Sowerby figures it in
1797, then considering it to be the same as the mushroom de-
scribed by Dr. Lister (Ray Cat. Plant. 123. 1677). Bulliard's
Agaricus acris pl. 538. f. С, H, №, is undoubtedly 2. vellerea Fr.

4. LACTARIA SUBVELLEREA Peck, Bull. Torrey Club 25: 360.
1808. [As Lactarius]

Pileus fleshy, thin, convex, soon umbilicate, at length nearly
infundibuliform, white, becoming tinged or spotted with yellowish,
and when dried cinnamon-colored (323. 2. 1-4), azonate, dry, cov-
ered with velvet-like tomentum, 7.5-15 cm. broad, margin at first
and for some time very involute, at length spreading ; gills white
to pale cream-colored, staining amber-white (72. 2. 4) where the
latex dries, often forking, adnate or slightly decurrent, narrow ;
stem white, tapering slightly toward the base, dry, velvety-pubes-
cent, firm, 1.8-2.8 cm. long, 1.2-2.5 cm. thick ; flesh amber-
white, odor faint; spores subglobose, smooth, 6-6. 5 их 6.5-7.5 и,
latex pale cream-colored or whitish, very acrid, usually abundant.

Has.: On dry ground, mixed woods. July.

Disrris.: Alabama, Earle (type); North Carolina, Burling-
ham 7, 1907.

DISTINGUISHING FIELD-MARKS: This species differs from /.
vellerea, to which it is closely related, in the narrow close gills,
and in the finer velvety tomentum or pubescence covering the

24 LACTARIAE OF THE UNITED STATES

pileus and the stem. The latex is cream-colored or dries yellow-
ish on the elle, The change in color of the pileus and stem dur-
ing drying is a marked characteristic.

The type specimens are in the herbarium of the N. Y. Botanical
Garden. The specimen which I found in North Carolina has the
gills a little more distant, but otherwise appears like the type. It
occurred in oak-chestnut woods at ап elevation of 1000 meters.

5. ГАСТАВТА DECEPTIVA Peck, Ann. Rep. М. Y. St. Bot. 38: 125.
1885. [As Lactarius.] — Hennings, in Eng. & Prantl,
Nat. Pflanzenfam. 1! * *; 216. 1898.

Pileus fleshy, firm, convex-umbilicate, then expanded and cen-
trally depressed or somewhat infundibuliform, white or whitish, dry,
glabrous at first, becoming torn into fibers and cracked as the
pileus matures, 5-15 cm. broad, margin involute at first and covered
with a dense soft cottony tomentum which fills in the space between
the margin and the stem, then spreading or elevated and more or
less fibrillose; gills whitish or cream-colored, some forking, some-
what distant, adnate or decurrent, rather broad ; stem white, nearly
equal, pruinose-pubescent, solid, 2-8 cm. long, 10-40 mm. thick ;
flesh white ; spores white, globose to elliptical, echinulate, 8-9 и X
9-12 и; latex white, unchanging, acrid. Edible.

Нав. : On the ground in woods, especially near hemlock trees,
rarely in oak-chestnut woods. July, August, and September.

DISTRIB.: New York, Peck, Shear, Jackson, Burlingham; Maine,
White ; Vermont, Burlingham 21, 1906; Massachusetts, Vai/ ; Con-
necticut, Underwood & Earle, Hanmer 682 ; Pennsylvania, Herést ;
Virginia, Murrill до; North Carolina, 1000 to 1675 meters eleva-
tion, Burlingham 56, 1907; Alabama, Ear/e ; Ontario, Guillet.

Irrus;.: Atkinson, Stud. Am. Fungi, f. 120, 121, Lactarius
resimus ; White, Conn. Geol. and Nat. Hist. Surv. Bull. з: 76 8.

Exsic. : Shear, New York State Fungi 70.

DISTINGUISHING FIELD-MARKS: This species has been much
confused with Z. vellerea Fr. and Z. piperata (L.) Ег., but differs
decidedly from both. The mature glabrous form of L. deceptiva
resembles the latter in general appearance, but the texture of the
pileus, the broader and more distant gills, and the larger more
echinulate spores distinguish it from that species. The pileus of
L. vellerea is covered over the entire surface with a short velvety

tomentum, while the center of 7. deceptiva is practically glabrous

LACTARIAE OF

in the young state, and the
margin is covered with cot-
tony fibers forming a soft roll-
like veil. The surface for some
distance toward the center has
the texture and the feeling of
chamois-skin. In very wet
weather the cottony roll on
the margin sometimes feels
slightly slippery.

6. LACTARIA GLAUCESCENS
Crossl. Naturalist 1000:
5. 3/а1000. [As
Lactarius |

Pileus fleshy, firm, thick,
convex-umbilicate, then in-
dide e Geen white, becom-
ing sta or spotted with
gelieft buff, dry, glabrous,
4-9 cm. broad, margin іпуо-
lute at first, even, naked ; gills
maize-yellow (36. £. 1-2) at
first, becoming white as the
pileus expands, staining glau-
cous-green (253. 4. 1. ) уу vhere
the latex dries, forked, very
close, adnate, 1-1.5 mm. wide;
stem white, tapering down-
wards or sometimes nearly
equal, solid, firm, 3-9 cm.
long, 1.5 cm. thick at the base

nutely and Am echinu-
late, 5.5-7 м, ccm terete-
clavate, 40-60 их 7-8 и;
latex white, drying glaucdus-
green on the flesh and gills,

THE UNITED STATES

|
|
|
|
|
|
|
|
!
|
|
|
|

|

|

|

b

very acrid, abundant. (FIGURE I.)

17, 1907.

No.

Lactaria glaucescens Crossl.

FIGURE Г.

26 LACTARIAE OF THE UNITED STATES

Has. : On the ground in sandy loam or in vegetable soil, oak- .
chestnut woods. July and August. Lj

DISTRIB. : North Carolina, 1000 meters elevation, Burlingham
77, 1907. |

Плозт.: Crossland, New and Crit. Brit. Fungi, Naturalist |
1900: IO. f. 1-3. 2

DISTINGUISHING FIELD-MARKS : The gills are very narrow and |
so crowded when young that а space between them is scarcely -
visible, and while tinted dull-yellowish when young, they become |
white with age. Тһе abundant white latex dries glaucous-green, |
which characteristic separates the species from all the other mem- |
bers of this group. It differs from L. pergamena also by the thick | |
flesh of the pileus, and from Г. piperata іп its narrower and more |
crowded gills. It was abundant in the * Pink Beds," North Caro- |
lina, іп rather dry oak-chestnut woods, during the latter part ог“
July and August. It is probable that it grows elsewhere, but has |
been referred to L. piperata.

7. Lacraria INVOLUTA Soppit; Cooke, Handb. Br. Fungi, ed. 2
380. 1883. [As Lactarius]

Нав.: On the ground, in woods or in grassy places on the
edge of woods. July.

Пізтків.: Connecticut, Hanmer 1479 ; Alabama, Underwood
& Earle.

Плозт.: Cooke, Br. Fungi, ai 1194.

DIsTINGUISHING FIELD-MARKS: This differs from the other.
Piperatae forms in its small size, and in the minutely silky covering |
on the margin of the pileus. The gills are as closely crowded

tinguish it.

LACTARIAE ОЕ THE UNITED STATES 27

In the Hanmer specimen the spores vary much in size, and in
this as well as in the Alabama specimens they run a little larger
than noted by Cooke, although spores as small as in that also occur.
The spores are really the final test in determining this species and
care must be taken not to mistake for it small specimens of some
of the other species.

II. RusTicANAE

Pileus without a pellicle, as in the Piperatae, and never viscid,
though in some species somewhat moist; color always dark,
varying from gray to brown or reddish- brown, tinted with lilac,
while in Lactaria atroviridis the color is dark-green ; flesh compact ;
latex usually very acrid, white and unchanging, although the
wounds sometimes change color.

Synopsis of species
Pileus glabrous, at least at first.
Gray to brown with lilac tints.

Gills thin. . Lactaria rusticana.
Gills thick, pileus becoming rivulose-squamulose. Lactaria ffexuosa.*
uliginous to brownish-black. Lactaria plumbea.*

Pileus scabrous-hairy.
Olivaceous-green. 9. Lactaria atroviridis.

8. Lactaria rusticana (Scop.)
Agaricus rusticanus Scop. Fl. Carn. 2: 452. 1772.
Agaricus pyrogalus Bull. Herb. Fr. pl. 529. f. г. 1701; Hist.

Champ. 1: 487. 1809
Lactarius pyrogalus Fr. Epicr. 339. 1838.

Pileus fleshy, firm, broadly convex, then plane to somewhat
depressed, gray to livid-gray or brownish-gray, darker in the
center, zoned toward the margin, moist in wet weather, but not
viscid, glabrous, 3-5-6-5 cm. broad, margin glabrous, inrolled, then
arched in mature specimens ; gills cream to ochroleucous, rarely
forking, distant, adnate, up to 5 mm. broad; stem paler than the
EE silver-gray, tinted with ochroleucous, nearly equal, some-

es tapering downwards, glabrous, smooth, compact, 3-5 cm.
long, 6-10 or 16 mm. thick ; flesh white, compact, thick, unchang-
ing; spores white, globose to subglobose, echinulate, greenish-
hyaline, 6-6.5 и x 6-8 и; cystidia transparent, 16-20 и long, 5—10 и
at the base, abruptly tapering ; latex white, unchanging, very acrid.
Poisonous.

Нав. : On open grassy places in woods. July to October.

* European species, the occurrence of which in the United States is doubtful.

28 LACTARIAE OF THE UNITED STATES

DISTRIB. : New York, Peck, Burlingham ; Vermont, Burling- |
ham r4, 1906; Ohio, Dawson ; Maryland, Banning. :
Irrusr.: Banning, Folio Md. Fungi, ai $6; Bull. Herb. Fr. A
529. f. г; Gillet, Champ. Fr. pl. 52 |162; 390]; Hahn, Der Pilz-
Sammler, f. 27 (poor); Krombh. Abbild. ad 14. f. 1-9; Lanz,
Fung. Mang. M. 55. f. 2. а, Ё, c, d; Noulet & Dass. Champ. 24
19. f. А; Pat. Tab. Analyt. Fung. A. 727; Rich. & Roze, Atl.
Champ. ad 37. f. 13-15 (poor); Sicard, Hist. Nat. Champ. 2/. 45.

f. 240.
DISTINGUISHING FIELD-MARKS: The nearly plane pileus, with
the margin remaining deflexed, the gray or brownish-gray zonate 4
surface, the absence of viscidity, and the distant rather yellowish
gills. In color it resembles both Lactaria circellata Fr. and Lac-
taria flexuosa Fr., from the former of which it may be distinguished
by the lack of viscidity, and from the latter by the persistently gla-
brous surface, the more regular margin, and the less distant gills.

9. LACTARIA ATROVIRIDIS Peck, Ann. Rep. М. У. St.
Mus. 42: 110. 1889 Ци.

Pileus fleshy, compact, nearly plane, soon depressed іп the =
center, olivaceous, azonate, dry, scabrous-hairy, sometimes crack-
ing into small areas, 6-10 cm. broad, margin involute, at length
spreading to uplifted; gills whitish, becoming mottled with dark-
green, especially where injured, sometimes forking near the stem, |
close, adnate to slightly decurrent, rather narrow ; stem colored `
like the pileus or paler, spotted, equal, dry, glabrous, firm, soon
hollow, 2-5 cm. long, 1-2 cm. thick ; flesh white ; spores creamy-
white, subglobose, echinulate, 7-8 и; latex white, unchanging, but
staining the gills green after some time, acrid.

Нав.: Deciduous woods. July and August.

Distris.: New York, Peck; Connecticut, Underwood & Earle;
District of Columbia, Murrill.

DISTINGUISHING FIELD-MARKS : Lactaria atroviridis closely re-
sembles Lactaria turpis (Weinm.) Fr., especially when dried, but
in the fresh condition it may be distinguished from it by the
absence of viscidity in wet weather, by its dark-green color, and |
by the change in color of the wounded gills.

The type specimens of this species аге in the herbarium of the |
N. Y. State Museum at Albany. The species seems to be rare

LACTARIAE OF THE UNITED STATES 29

DOUBTFUL SPECIES
LACTARIA PLUMBEA (Bull.) Fr. Epicr. 339. 1838. [As Lactarius. |

— Schróter in Cohn, Krypt.-Fl. Schles. 3: 536. 1889.

This species has been reported from New York by Peck ; from
Wisconsin by Bundy. Тһе specimens referred to Lactaria plum-
bea by Peck differ from the typical form in color and in the pres-
ence of a tomentum in the fresh condition. It seems to me doubt-
ful whether the New York specimens represent this species. They
appear much like Z. glyciosma but have no odor. Bundy’s speci-
mens are not preserved. The species is described in the Ann.
Rep. N. Y. St. Mus. 38: 127. 1885. A good figure of it may
be found in Вапа, Champ. Nice, //. 27. f. 1-5. Itis regarded as
poisonous.

LACTARIA FLEXUOSA (Fr.) Fr. Epicr. 338. 1838. [As Lactarius. |
— Schrót. in Cohn, Krypt.-Fl. Schles. 3: 536. 1889

This European species has been reported from North Carolina
by Schweinitz, from Vermont by Frost and by Burlingham. The
specimens which I assigned to this species do not correspond
closely enough to specimens which I have since received from
Romell, to justify citing them as this species. The Frost and
Schweinitz specimens are not "preserved. Тһе species is well fig-
ured by Cooke, Br. Fungi, AJ. 992.

III. ToRMINOSAE

Plants rather large, but the flesh not so compact as in the
Piperatae, and the pileus viscid when wet ; margin of the pileus in-
rolled at first and covered with long coarse tomentum, which may
be persistent or fugacious. The latex varies from intensely acrid
in L. torminosa to acrid in 2. speciosa.

Synopsis of species

Latex white, becoming yellow.

Pileus white, azonate. 13. Lactaria resima.
Pileus yellowish, zonate to subzonate. 12. Lactaria scrobiculata.
Latex white, wounds becoming heliotrope. 14. Lactaria speciosa.

Latex white, unchanging.
Marginal tomentum persistent.
Pileus zonate, some shade of yellow. 10. Lactaria torminosa.
Pileus azonate.
White to flesh-colored tinged with fuscous. 11. Галата cilicioides.*

* The latex sometimes very slowly turns yellowish.

30 LACTARIAE ОҒ THE UNITED STATES

Marginal tomentum fugacious.
Pileus white, zoned or spotted with red. Lactaria sanguinalis.*

10. LACTARIA TORMINOSA (Schaeff.) Pers. Tent. Disp. Meth. Fung.
64. 1797

Agaricus torminosus Schaeff. Fung. Bav. Icon. 4: 7 (index).

1774.
Lactarius villosus Clements, Bot. Surv. Neb. 4: 20. 1896.

Pileus fleshy, convex, depressed in the center with the margin
involute, finally nearly infundibuliform with the margin merely
deflexed, pale pinkish-yellow or pale-ochroleucous or ochraceous
tinged with incarnate, often zoned with deeper color, sometimes
nearly white and azonate, viscid when wet, 4-10 cm. broad, center
glabrous, margin persistently white-tomentose ; gills whitish or
cream-yellow tinged with pale-incarnate, some forking near the
stem, close, decurrent, thin, 5 mm. broad; stem paler than the
pileus, sometimes faintly spotted with yellow, equal or tapering
downwards, glabrous or pruinose, smooth, stuffed, becoming hol-
low, 3-7 cm. long, 2 cm. or less thick; flesh white, not changing
color; spores white, broadly elliptical, echinulate, uniguttulate,
hyaline, 8-10 их 6-8 p; latex white, unchanging, very acrid.
Poisonous.

Нав.: In deciduous or coniferous woods, in open or shady
places. August and September. А

Півтків.: New York, Peck, Underwood; Maine, Murrill,
White; Vermont, Burlingham 56, 1906, Burt; Connecticut,
Underwood ; Alabama, Earle; North Carolina, 1000 meters ele-
vation, Burlingham ; Colorado, 2900 meters elevation, Clements ;
Nebraska, Clements.

ý ә.
* European species, the occurrence of which іп the United States is doubtful.

елек

абы Фр ЕЕ

LACTARIAE ОЕ THE UNITED STATES 31

gewachse, ^. 5; Rich. & Roze, Atl. Champ. МА 37. Je 1-0)
Schaeff. Fung. Bav. Icon. A. 12; сага, Hist. Nat. Champ. pl.
43. f. 232; Sv. Bot. pl. 184; Sowerby, Eng. Fungi, p/. 103;
Venturi, Stud. Micol. AX. 6. f. 53, 54.

Exsic.: Clements, Cryptogamae Formationum Coloradensium
364; Herpell, Sammlung práparirter Hutpilze 46; Karsten,
Fungi Fennici, 508 ; Sydow, Mycotheca Marchica 7504.

DISTINGUISHING FIELD-MARKS: The large size, the persistent
long whitish tomentum on the margin, and the unchanging white
latex. The color of the pileus is some shade of yellow, some-
times tinted with incarnate, and it is usually zoned. The tomen-
tum often extends from the margin midway to the center of the
pileus. Occasionally, nearly white azonate forms occur.

After comparing the type specimens of Lactarius villosus
Clements with both American and European specimens of Lac-
taria torminosa (Schaeff.) Pers. I have concluded that it is without
question identical with the latter species. The spore-measure-
ments run somewhat larger than indicated by Clements and the
stem becomes hollow as shown both in the specimens and in the
photograph accompanying the sheet in the Crypt. Format. Colora-
densium.

This species is the Agaricus piperatus var. a L. Fl. Suec. 441.
1755, but not the Agaricus piperatus L. Sp. Pl. 1173. 1753, which
from the synonyms given by Linnaeus is plainly the same as Zac-
taria piperata of Fries. In Monogr. Suec. 2: 153, Fries de-
scribes the stem as “subtiliter adpresse tomentosus 2. glabratus,”
and Massee and Stevenson have followed Fries' description in this
particular; but in all specimens which I have examined the stem
appears to the naked eye as merely pruinose, and even with the
aid of a lens it can scarcely be described as tomentose.

тт. LACTARIA CILICIOIDES (Fr.) Fr. Ерісг. 334. 1838. [As
Lactarius.] —Hennings, in Eng. & Prantl, Nat.
Pflanzenfam. 1! **: 218. 1898
Agaricus tomentosus Otto, Versuch Agar. 74. 1816. Not Agari-
cus tomentosus Bull. Herb. Fr. pl. 138. 1782.
Agaricus cilicioides Fr. Syst. Myc. 1: 63. 1821.
Pileus fleshy, not very compact, convex, becoming plane with

92 LACTARIAE ОЕ THE UNITED STATES

the center depressed, sometimes nearly infundibuliform, pale-incar-
nate tinged with fuscous, azonate, viscid when wet, entirely covered

ee с сс ен аа

with matted tomentum, 5-10 cm. broad, margin involute, fibrillose- |

woolly ; gills white to yellowish, forking, close, decurrent, rather

_ narrow; stem whitish, sordid, never spotted or scrobiculate, equal, |

pruinose-silky when viewed with a lens, firm, stuffed, at length

somewhat hollow, 5-7.5 ст. long, up to 2.5 cm. thick; flesh |

yellowish -white ; spores white, minutely echinulate, broadly еШр- ч

tical, 84 x 6-7 p; latex white or pale yellowish-white, acrid, not
abundant, sometimes becoming more yellowish when exposed to
the air.

5 у DE S
Нав: ‘Woods and open places, especially near pine trees, |

Peck. September and October.

DISTRIB.: New York, Peck. It has also been reported from :

Maine by Ricker, and from Ohio by Morgan.
Пловт.: Britz. Lact. f. 2; Cooke, Br. Fungi, 77. 973; Schaeff.
Fung. Bav. Icon. 2/ 228, Agaricus crinitus.

ыт Эх Ас

+
|

DISTINGUISHING FIELD-MARKS: This species may Бе distin-
guished from Lactaria torminosa, to which it is closely related, :
by the shorter and more matted tomentum covering the entire :
pileus, by the paler or duller-colored azonate pileus, and by the `
less abundant latex. At the margin of the pileus the tomentum |
becomes somewhat longer. It is finer than the tomentum on the |

margin of Lactaria torminosa.

Agaricus intermedius Krombh. ai 58. f. 11-13 has been con- :
sidered a variety of L. cilicivides, but it seems to be more nearly `

Lactaria scrobiculata Scop. Massee, however, describes it as 8
distinct species.

Krombholtz describes an Agaricus tomentosus

pl. 40. f.17, 18 * as synonymous with Agaricus tomentosus of Otto, |
and Massee and Cooke have followed Krombholtz, but шеге
does not seem to me to be sufficient evidence that Krombholtz |

species is the same as that of Otto, while Otto’s description agrees

well with Agaricus cilicioides Fr. Тһе specific name given either

by Schaeffer or Otto was a homonym, hence cilicioides stands as the
specific name.

12. LACTARIA SCROBICULATA (Scop.) Fr. Epicr. 334. 1838. (А5
Lactarius.] —Schrót. in Cohn, Krypt.-Fl. Schles. 3: 341. 1889
Agaricus serobiculatus Scop. Fl. Carn. 2: 450.

1792.
Pileus fleshy, convex, soon depressed in the center, at length |

* Fig. 17 is lacking, and / 78 shows no tomentum.

LACTARIAE OF THE UNITED STATES 38

infundibuliform, yellowish to ochraceous or even reddish-yellow,
zoneless, or subzonate or conspicuously zonate, viscid when moist,
6-15 cm. broad, margin at first involute and tomentose, then
spreading and plane or upturned and nearly or quite naked ; gills
whitish or pale-yellowish, sometimes forking close to the stem,
cro wded, adnate or slightly decurrent, thin ; stem of the same color
as the pileus or paler, with brighter-colored elliptical or orbicular
scrobiculate spots, equal, glabrous, hollow, 3-7 cm. long, 1-2 cm.
thick ; spores white, minutely echinulate, 6.5—7 и x 8-10 4; latex
white, quickly changing to sulphur-yellow, acrid. Suspicious.

Нав.: In moist woods. July to November.

DISTRIB.: New York, Peck; Vermont, Frost; Connecticut,
Earle ; Alabama, Ват.

Плозт.: Вапа, Champ. Nice, AJ. 18. f. 3-6, Britz. Lact. f.
1; Cooke, Br. Fungi, ai, 977 ; Gillet, Champ. Fr. 2/. 154 [392];
Hahn, Der Pilze-Sammler, ed. 2. f. 27; Krombh. Abbild. ai 58.
f. 1-6 ; Lorinser, Essb. und Gift. Schwàm. 2/. 9. f. 6; Lucand,
Champ. Fr. A. 971; Pat. Tab. Analyt. ai 409; Schaeff. Fung.
Bav. Icon. AX. 227.

DISTINGUISHING FIELD-MARKS: Тһе yellowish, more or less
zonate pileus, the coarse rather conspicuous tomentum on the
margin of the young pileus, the bright-colored scrobiculate spots
on the stem, and the rapid change in the color of the latex from
white to sulphur-yellow. Тһе plants are large and the margin
becomes glabrous in the mature plant.

Lactaria scrobiculata is commonly described by European
writers as azonate. But Fries in Monogr. 2: 153, says “vulgo
azonus, interdum vero conspiciuntur zonae" Маззее and Stevenson
also describe it as sometimes zonate. The Alabama specimens are
conspicuously zonate in the dried state, while the Connecticut
specimens are zonate or subzonate. When the plants are grow-
ing in the open they fade more or less and this may account in
part for the difference in zonation.

13. ГАСТАВТА RESIMA (Fr.) Fr. Epicr. 336. 1838. [As Lacta-
rius.] — Schrót. in Cohn, Krypt.-Fl. Schles. 3: 542. 1880
Agaricus resimus Fr. Hym. Eur. 472. 1821.
Pileus fleshy, firm, deeply umbilicate with the margin involute,
at length infundibuliform with the margin arched or spreading,
whitish, soon faintly tinged with yellow, darker in the center,

34 LACTARIAE OF THE UNITED STATES

azonate, viscid when wet, glabrous except the extreme margin,
which is white-tomentose, at length entirely glabrous, 8 cm. or |
more broad; gills whitish or pale-cream, a few forking near the
stem, close, decurrent, about 4 mm. broad; stem whitish, spotted
with cream or dull-yellow, equal, glabrous, or pruinose at the top,
stuffed, becoming hollow, 3.5 cm. long, 2.5 cm. thick ; flesh white,
odor rather strong when fresh and persisting for a time; spores
white, globular to elliptical, echinulate 5-7 M X 7-9 p; latex
d

Re M

instantly changing to sulphur-yellow, acrid.

Нав.: In woods, chiefly in mountainous regions.

DISTRIB.: Vermont, Burlingham 110, 1906.

Intust.: Fr. Icon. ai 169. f. г.

DISTINGUISHING FIELD-MARKS : The whitish lily-shaped azonate
pileus with a fringe-like white tomentum on the extreme margin,
and the very rapid change in the color of the latex from white to
sulphur-yellow. Іп the mature plant the tomentum is often lack-
ing. The specimen found in Vermont was growing under a small |
spruce on the edge of a wood in a somewhat mountainous district.
Near the edge of the pileus were four or five faint ridges or corru-
gations running parallel to the margin.

In Мопорг. 2: 152, Fries says of the stem “sud lente pube
tenuissima vix perceptibili tectus" In other places the stem is
described as villous, but the statement in the Monograph is the
more accurate. Our specimen, however was simply pruinose, and
it is probable that the scarcely perceptible down mentioned by
Fries was little more than a pruinosity, although there might be а
variation in the nature of the covering under different environmental
conditions.

14. Lactaria speciosa sp. nov.

tinted with honey-yellow, with elliptical spots of flavous, some-
times scrobiculate, staining heliotrope where handled, equal, viscid

LACTARIAE OF

THE UNITED STATES

No. 40, 1907.

Lactaria speciosa Burl.

FIGURE 2.

36 LACTARIAE OF THE UNITED STATES

when wet, sometimes slightly tomentose at the Базе, stuffed, 3
becoming hollow, 2.5-7 cm. long, 1-1.5 cm. thick; flesh white, 7
staining heliotrope where wounded, odor none; spores white, |
elliptical, strongly echinulate, 8-8.5 их 9.5—10 Ж latex white, :
not plainly changing, but staining the flesh and gills heliotrope |
(188. 1. 1), mild to slightly acrid, abundant in young plants, |
scanty in old ones. (FIGURE 2.) f

Нав.: In oak woods, sandy soil, oak-chestnut woods, frequently T
by wood-roads, in dry as well as wet weather. August and.
September. 2

DISTRIB. : “ Pink Beds," North Carolina, 1000 to 1200 meters : 1
elevation, Burlingham 40, 1907 (type); Tennessee, 400 meters |
elevation, Murrill ; Virginia, 670 meters elevation, Murrill. |

DISTINGUISHING FIELD-MARKS: This species із readily recog- | |
nized by the pallid pileus and the honey-colored tomentum and 4
zones, together with the heliotrope color which injured parts |
assume. It was found abundantly during August in dry weather %
and іп dry soil. Тһе pileus is always zonate, the stem not at all E
or only slightly scrobiculate and the latex only slightly acrid.
The tomentum is darker at the tips of the hairs, which become
matted together in small clusters, thus standing out more prom-
inently against the paler color of the pileus.

azonate, and deeper- or brighter-yellow in color, while the spores
are smaller and much less echinulate.
DOUBTFUL SPECIES

LACTARIA REGALIS Peck, Ann. Rep. М. У. St. Mus. 26: 64. 1874
[As Lactarius |

mens represent a distinct species or whether they should be classe?
with Z. resz;ma. Тһе margin of this latter species is at length g!

LACTARIAE ОЕ THE UNITED STATES 37

brous and the stem is practically glabrous from the first. Although
the type specimens of 2. regalis do not show any very young plants,
some of them seem to be young enough to have retained the to-
mentum on the margin. Since, however, only the type collection
is known, I prefer to leave the species in doubt until further col-
lections can be made.

LACTARIA SANGUINALIS(Batsch) Schrót. [Lactarius controversus Fr. |

This species has been reported from California by Harkness &
Moore, but the specimens from which the determination was made
were not preserved. The species is described in МсПуате’$ One
Thousand Am. Fungi, 164. 1902, and is well figured by Batsch,
Elench. Fung. ai, 36. f. 201; by Fries, Sverig. Svamp. 2/. 20; by
Cooke, Br. Fungi, //. тооз; and by Gillet, Champ. Fr. ai 160
[387]. Вай.

LACTARIA PUBESCENS (Fr.) Fr. — This species was reported from
New York by Peck (Ann. Rep. N. Y. St. Mus. 31: 31. 1879),
but was ommitted from his list of Zactariae in the 38th Report,
1885.

IV. CROCEAE

Pileus very viscid when young or in wet weather, the margin
involute at first and covered with a short tomentum, some shade
of yellow, zonate, plants large, flesh rather compact ; gills darker
with age, and more or less pruinose; latex acrid, white, changing
to yellow.

Synopsis of species
Pileus maize-yellow tinted with yellowish-salmon, latex becoming sulphur-yellow.
16. L. delicata
Pileus saffron-yellow to orange-cadmium, latex becoming yellow-cadmium,
15. L. crocea

15. Lactaria crocea sp. nov.

Pileus fleshy, brittle, broadly convex-umbilicate with the mar-
gin involute, then expanding, depressed in the center with the margin
deflexed, saffron-yellow to orange-cadmium (49), subzonate, expal-
lent, very viscid when moist, glabrous, 5-10 cm. broad, margin in-
volute and pruinose-downy at first, then glabrous; gills pallid, then
honey-yellow (35), and at length yellowish-buff ( 310. г. г), be-
coming cadmium (47. 2. 7) where cut or bruised, seldom forking,
not close, adnate, 5-6 mm. broad; stem of the same color as the
pileus but paler, spotted, equal, glabrous or sometimes tomentose

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at the extreme base, stuffed,
becoming hollow, stout, 5

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thick ; flesh whitish, staining
yellow-cadmium where cut,
odor none; spores yellow-
ish, broadly elliptical, echi-
nulate, 7 и x 8-8.5 м; la-
tex white, scanty, slowly
becoming yellow-cadmium,
and staining the gills and
flesh, acrid. (FIGURE 3.)

Has.: In oak and chest-
nut woods, growing from
vegetable mold and dead
leaves. August and Sep-
tember.

DISTRIB. : North Caro-
lina, 1000 meters eleva-
tion, Burlingham 48, 1907
(type); Virginia, Murrill.

DISTINGUISHING FIELD-
MARKS: The beautiful
orange-cadmium color of
the viscid pileus and stem,
and the paler, rather distant
gills, which become yellow-
cadmium where wounded.
The zones are sometimes
very distinct, but more often
obscure, especially in old
age. Тһе latex is scanty
and the change in color is
detected mostly by the
change in the color of the
wounded gills and flesh.
The wounds never become
greenish as in Lactaria de-
lictosa. The much deeper
color of the plant, the more

|

39

72, 1907.

No.

Lactaria delicata Burl.

FIGURE 4.

40 LACTARIAE OF THE UNITED STATES

viscid pileus, and the more distant gills separate this species from |
Lactaria chrysorhea and Lactaria theiogala.

16. Lactaria delicata sp. nov.

Pileus fleshy, firm, convex-umbilicate, at length nearly infundib-
uliform, maize-yellow (36. 2. 3), tinted in the center with yellow-
ish-salmon (65), faintly but decidedly zonate, viscid and covered |
with gluten when wet, glabrous, 8-12 cm. broad, margin involute |
at first and covered with coarse short tomentum, then merely de-
flexed and glabrous; gills whitish, becoming maize-yellow with
age, some forking near the stem, close, slightly decurrent, 5-7 mm. E.
broad; stem whitish to maize-yellow tinted with yellowish salmon, -
more or less scrobiculate-spotted, spots of the same color as Ше.
rest of the stem or duller, equal or tapering downwards, glabrous, |
stuffed, becoming hollow, 4-5 cm. long, 1.5-2.5 cm. thick; flesh
white, odor strong ; spores tinted yellowish-salmon in mass, sub-
globose, echinulate, 7-8 м; latex white, becoming sulphur-yellow, |
acrid, scanty. (FIGURE 4.) ў

Нав.: In sandy loam and dense shade, oak and chestnut |
woods, July and August. a

Півтків.: “ Pink Beds," North Carolina, 1000 meters elevation, |
Burlingham. | 1

mature pileus, the rather persistent viscidity, the lily shape of the
. mature pileus, and the change in the color of the latex.

V. AGGLUTINATAE |
Pileus very viscid when young or in wet weather, margin invo-
lute at first and covered with a short tomentum, color from w!
to buff and umber with greenish tinge, becoming deeper in агу!

STATES

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49 LACTARIAE OF THE UNITED STATES

flesh rather firm, plants medium-sized to large; gills becoming |
deeper in color with age and more or less pruinose; latex acrid,
white, unchanging.

Synopsis of species
Pileus whitish.

nate, sparsely covered with coarse, short tomentum. 18. Z. lanuginosa.
Azonate, glabrous. L. subinsulisa,* |
Pileus buff, papillate when wet, squamulose when dry. 17. Г. agglutinata,
Pileus olivaceous-umber, blackening in drying. 19. Г. turpis. |
Pileus grayish-green, zonate, not blackening. L. blennia.t [

17. Lactaria agglutinata sp. nov.

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so when older, slimy-viscid when wet, with subrugose elevations
or papillae showing through the gluten, appearing squamulose to

, 6-

minutely pubescent af а the pubescence becoming less notice- |
able as the margin unfolds ; gills yellowish-buff (310. 2. г), some |
forking near the stem, close, slightly decurrent, 2—4 mm. broad; |
stem of the same color as the pileus or paler, sometimes spotted,
equal or tapering downwards, viscid when wet, glabrous, firm as
though solid, becoming spongy to hollow, 2.5-4 cm, long, 1-1.5
ст. thick ; spores creamy-white in mass (zo. 4. 1, 2), subglobose,
echinulate, 7-8 и; flesh white, odor somewhat like raw pumpkin ;
latex white, unchanging, acrid. (FIGURE 5.)

Нав.: Among dead leaves, in rather sandy soil, oak-chest-
nut woods, frequently under the flowering dogwood. August
and September.

Пізтків.: “ Pink Beds," North Carolina, 1000 meters eleva-
tion, Burlingham 59, 1907.

DISTINGUISHING FIELD-MARKS : The medium size, buff color,
and the papilliform and rugose elevations showing through thick,
glistening gluten when wet, and the squamulose appearance of the
pileus when dry. One peculiarity of the pileus is that if it becomes |
wet again after having dried, the squamules swell up and the sur- |
face appears papilliform and rugulose as at first.

is species seems to be quite distinct. While resembling the
Triviales in the very viscid character of the pileus, it also approaches
eer Doubtful species, me

T European species, the occurrence of which in the United States is doubtful.

LACTARIAE ОЕ THE UNITED STATES 43

the Zorminosae in the minutely tomentose condition of the margin
of the young pileus. The gills become somewhat pruinose in the
mature plant. While not abundant, the species was found several
times during August and September, in as many different places.

3 No. 67, 1907.

url,

Lactaria lanuginosa 1

FIGURE 6.

18. Lactaria lanuginosa sp. nov.

Pileus fleshy, firm, convex-umbilicate, then expanding and
depressed in the center, whitish to yellowish-flesh (68), zoned
especially toward the margin, where the zones are close and nar-
row, viscid, sparsely covered with agglutinated short white coarse
tomentum, 7-8 cm. broad, margin involute at first and the extreme

44 ТАСТАКІАЕ OF THE UNITED STATES

edge covered with fine short tomentum, at length glabrous and |
upturned ; gills whitish, at length maize-yellow (36. 2. 3) to yel-
lowish-buff (370. £. 2), seldom forking, close, slightly decurrent,
3-5 mm. broad; stem white, becoming yellowish-buff spotted
more or less with darker, equal or tapering downwards, viscid
when young or wet, minutely pubescent to hairy tomentose, then
nearly glabrous, firm, stuffed, then hollow, 3 cm. long, 1-2 cm.
thick ; flesh white, unchanging, odor rather strong ; spores white,
nearly globose, echinulate, 6-7 м; latex white, unchanging, scanty,
astringent, not very acrid. (FIGURE 6.)

Нав.: Among dead leaves under oak, maples, alder, and
Rhododendron. August and September.

Distris.: North Carolina, Burlingham.

DISTINGUISHING FIELD-MARKS: The large, whitish, zonate |
pileus, covered with short sparse, agglutinated tomentum. The 7
zones seem to be concentric translucent rings, which appear ||
cream-colored in contrast with the opaque white of the remainder
of the pileus. The tomentum is more noticeable when the viscidity
has disappeared. In the dried plant the rings are conspicuous and
bright-colored. Ki

I found this species in the “ Pink Beds," North Carolina, five |
times during August and September. It seems to be quite distinct |
from any other species. The type specimens, ло. 67, 1907, аге іп
the herbarium of the N. Y. Botanical Garden.

10. LACTARIA TURPIS (Weinm.) Fr. Epicr. 335. 1838. [As
Lactarius |

glabrous or agglutinated-fibrous, 6-12 cm. broad, margin involute
at first and yellow-villose, then glabrous; gills cream-colored, then |
darker yellow, becoming nearly black where bruised, then ash

* Report of Botanist published in advance,

LACTARIAE OF THE UNITED STATES 45

glabrous, smooth or somewhat scrobiculate, or sometimes merely
spotted, spots becoming nearly black in dried specimens, firm,
stuffed, occasionally becoming hollow when old, usually 3-4 cm.
long, 1.5-2.5 cm. thick ; flesh whitish, odor slight ; spores white,
globose to subglobose, echinulate, 6.5-8 и; latex white, unchang-
ing, acrid. Possibly edible.

Has. : On the ground in mixed woods, often near fir or spruce
trees. August and September.

DISTRIB. : New York, Peck; Vermont, Jones, Burlingham ;
Maine, White ; Connecticut, Hanmer ; Ohio, Beardslee ; it was also
reported from North Carolina by Schweinitz.

Плозт.: Cooke, Br. Fungi, p/. 987; Fl. Dan. pl. 1913; Fr.
Sverig. Svamp. M. бо; Gillet, Champ. Fr. 2/. 758 [397] ; Hahn,
Der Pilz-Sammler, ed. 2. f. 22; Harzer, Pilze, р/ бо; Krombh.
Abbild. a. бо. f. 1-6; Lorinser, Essb. und Gift. Schwàm. Р/ д.
f. 2; Lucand, Champ. Fr. ai 47; Phoebus, Deutschl. Krypt.
Giftgewachse, AM. б. f. 1-3 ; Roumeg. Crypt. Illustr. f. 720.

Exsic.: Karsten, Fungi Fennici 306; Sydow, Mycotheca
Marchica 7009.

DISTINGUISHING FIELD-MARKS : The yellowish-brown or oliva-
ceous color of young plants and the blackish color of mature
plants, the slimy condition of the whole mushroom in wet weather,
the blackening of the gills with injury or in drying, and in many
cases the grayish color due to the presence of the spores. The
pileus may be covered with villose fibers which are closely stuck
to the surface, and are not easily distinguishable, or it may be
practically glabrous. The yellow down or villosity on the margin
disappears in the mature plant, and is at no time conspicuous.

I have compared the type specimens of Lactarius sordidus
Peck with European specimens of Lactaria turpis and have not
been able to detect any specific difference. Some of the Euro-
pean forms show the stem to be slightly spotted, and some of the
specimens of Г. sordidus have slightly scrobiculate spots on the
stem. These spots become more prominent in drying.

DOUBTFUL SPECIES

LACTARIA SUBINSULSA Peck, Ann. Кер. М. Y. St.
Mus. 43: 19. 1890. [As Lactarius]
Pileus fleshy, firm, convex to nearly plane, umbilicate, whitish,
azonate, viscid, glabrous, 5-10 cm. oad, margin involute and

46 LACTARIAE OF THE UNITED STATES

slightly tomentose at first, soon naked ; gills whitish, close, adnate
to decurrent, narrow; stem whitish, not spotted, obscurely rugu-
lose-reticulated, 2.5-4 cm. long, 12-16 mm. thick; spores sub-
globose; latex white, unchanging, tardily acrid.

Нав.: Pine groves. August.

DISTRIB.: New York, Peck.

I have not seen this plant living, and the above description is
arranged from the original description and from the dried specimens.
The type specimens are in the herbarium of the N. Y. State
Museum at Albany. Тһе plant has not been found since the type
collection and the dried specimens so closely resemble Г. ай
Peck as to be indistinguishable from that species and at present I
am inclined to regard it as the same.

* LACTARIA BLENNIA (Fr.) Epicr. 337. 1838. [As Lactarius.]
— Hennings, in Eng. & Prantl, Nat. Pflanzenfam. 1“:
217. 1898

Agaricus aerugineus Lamarck, Fl. Fr. 1: Gor) 1775.

This species has been reported from Connecticut by White on
the basis of specimens collected by Hanmer, and from Minnesota
by Johnson. I have seen the Hanmer specimens and they are all
Lactaria turpis (Weinm.) Fr. The Johnson specimens are not
available for examination. For these reasons I omit this species
from the list of our known Lactariae. А description of the species
is given in McIlvaine's One Thousand Am. Fungi. The following
may further help in the recognition of the species should it be
found and will also enable one to avoid confusing it with related
species.

DISTINGUISHING FIELD-MARKS : The grayish-green viscid pileus,
zoned with drop-like concentric spots. It may be distinguished
from Lactaria turpis (Weinm.) Fr. by the paler-green color, by
the more inconspicuous down on the margin of the pileus and the
otherwise glabrous surface and by the wounds of the gills turning
cinereous. The gills do not become blackish in drying as do those
of L. turpis. Regarded as poisonous.

УТ. AsPIDEAE
Pileus very viscid when young or in wet weather, and the

margin minutely tomentose or pruinose-downy to nearly glabrous |

* The name of this species should be Lactaria aeruginea ( Lamarck )

ae A %

NE Mead саран ЕТ

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LACTARIAE OF THE UNITED STATES 41

when young, wounds becoming Шас; gills somewhat darker with
age and slightly pruinose.

While Lactaria aspidea апа Lactaria aspideoides have a decidu-
ous minute tomentum on the margin of the young pileus, the other
species are merely pruinose-downy to almost glabrous, and all
the species agree in being entirely glabrous when mature. This,
together with the extreme viscidity of the pileus and the lilac color
assumed by the wounds, indicates their relationship. Lactaria
speciosa seems to be more closely related to the Zorminosae by its
very long tomentose covering and less viscid surface.

Synopsis of species

Pileus gray.
Conspicuously zoned and spotted, 7-12 cm. broad. 23. L. maculata.
Faintly zoned or azonate, up to 8 cm, broad. 22. L. lividorubescens.
Pileus yellow.
Azonate, latex acrid. 20. L. аз аға.
Zonate, latex bitter. 21. L. aspideoides.

20. LACTARIA ASPIDEA (Fr.) Fr. Еркг. 336. 1838. [As
Lactarius.]— Hennings, in Eng. & Prantl, Nat.
Pflanzenfam 1'**:218. 1898

Agaricus aspideus Yr. Obs. Мус. 2: 180. 1818.
Agaricus roseo-violascens Lasch, Linnaea 3: 161. 1828.

Pileus fleshy, not thick, plane to convex, slightly papillate, at
length depressed, straw-colored, somewhat sordid, azonate, viscid,
3-10 cm. broad, margin at first with a zone of white silky decidu-
ous tomentum, then entirely glabrous ; gills white or cream, rather
close, somewhat thick, of various lengths, here and there connected
by branches,about 2 mm. broad; stem of the same color as the pileus
or paler, nearly equal, even, glabrous, stuffed, then hollow, 5-8
cm. long, up to 15 mm. thick ; flesh white, becoming lilac where
wounded; spores globular, echinulate, 8-10 и; latex white,
changing to lilac, acrid. Porsonous.

Нав.: In moist places under willows, and іп the border of
woods, September.

Півтків.: New York, Peck.

Птозт.: Lanzi, Fung. Mang. pl. 56. f. 2. а, б, с, d; Cooke,
Br. Fungi, a 1083 ; Agaricus pudibundus* Pico, Mem. Soc. Med.
Par. M. 12. 1780.

* This name was preoccupied. See Scop. Fl. Carn. 2: 452. 1772.

48 LACTARIAE ОЕ THE UNITED STATES

DISTINGUISHING FIELD-MARKS: The pale-yellow or straw-colored |
pileus, the absence of zones, and the acrid latex.

This species has been found but once in the United States.
Peck notes that the stem was not spotted in his specimens, and 2
that the spores were broadly elliptical or subglobose.

Lactarius uvidus Krombh., which has commonly been referred
to Lactaria aspidea Fr., does not seem to me to agree closely |
enough to warrant considering it this species. Krombholz de-
scribes the color of the pileus as “ braun," and that of the stem and |
gills as white.

21. LACTARIA ASPIDEOIDEs Burl. Bull. Torrey Club 34: 87.
1907. [As Lactarius] :

Pileus fleshy, rather firm, convex-umbilicate, then plane, becom- |

ing infundibuliform with age, sulphur-yellow (18) with narrow zones |

of butter-cup yellow (22), glabrous, 3-4.5 cm. broad, very viscid |
when wet, gluten thick and persisting, margin involute and minutely A
tomentose at first, then spreading and glabrous; gills whitish, |

then cream-colored spottéd with yellow, staining lilac where |
wounded, sometimes forking next the stem, close, adnate, but -
acute at the inner end, 4 mm. broad; stem sulphur-yellow, often |
spotted with butter-cup yellow, equal or abruptly smaller at the `
base, viscid when young or wet, glabrous, stuffed, becoming hol:
ow, 2-3.5 cm. long, 6-10 mm. thick; flesh whitish, changing t° |
faintly lilac where exposed to the air; spores white, globular to =
broadly elliptical, echinulate, 5-7 » x 7-8 и; latex white, becoming |
lilac where in contact with the broken flesh, taste bitter. E

Нав. : In grassy hillside sheep-pasture near small fir trees, after `
heavy rain and warm weather. September. 5"

Distris.: Vermont, 523 meters, Burlingham 115, 1906.

DISTINGUISHING FIELD-MARKS: The bright-yellow color of the |
entire plant, the very viscid zonate and glabrous pileus, the broad, `
short, chaff-like tomentum on the extreme edge of the young pileus, |
and the lilac color assumed by the wounded flesh or gills. e
latex does not seem to change color except where in contact with |
the flesh. This species is most closely related to Г. aspideus Fr, |
but differs in its brighter color, the zonate pileus, the smaller size, a
and the bitter latex. It can readily be distinguished from 2. spe : |
05а, since the pileus of the latter is covered with long tomentum |
and is pallid, with honey-colored zones. Z. aspideoides is also à
much smaller plant.

LACTARIAE OF THE UNITED STATES 49

22. Lactaria lividorubescens (Batsch)
Agaricus lividorubescens Batsch, Elench. Fung. 2: 51. ai 36. f.

202. 1780.

Agaricus uvidus Fr. Obs. Мус. 2: 191. 1818.
Lactarius uvidus Fr. Epicr. 338. 1838.
Lactarius livescens Passerini, Nuovo Giorn. Bot. Ital. 4: 105.

1872.

Pileus fleshy, firm, then more lax, rather thin, convex, then
plane to depressed, often umbonate, cinereous or brownish- -gray,
tinged with lilac, often faintly spotted and zonate especially when
young, viscid, glabrous, 4-8 cm. broad, margin at first inrolled
then spreading except the extreme edge, which remains inrolled
for some time and is at first white-pruinose to minutely downy ;
gills white to cream, quickly changing to lilac then violet when
wounded, sometimes a few forking next the stem, close, thin, of
various lengths, adnate to slightly decurrent, up to 5 mm. broad ;
stem white to cream, not spotted or only obscurely so, equal or
tapering upwards, a little viscid when moist, glabrous, sometimes
tomentose at the base, stuffed, becoming hollow, 4-8 cm. long,
6—12 mm. thick ; flesh white, changing to lilac where wounded ;
spores white, elliptical, echinulate, 7 м x 8-12; latex white at
first, changing to dark-lilac or violet where in contact with the
the flesh, acrid, sometimes bitter.

Нав.: On the ground in woods, in moist places. August
and September.

Півтків.: New York, Peck; Maine, White; Vermont, Bur-
їп йат 63, 1906 ; Connecticut, Earle, Benedict ; Maryland, Ban-

hos. : Banning, Folio Md. Fungi, ai $0; Batsch, Elench.
Fung. A. 36. f. 202; Bern. Champ. Roch. AJ. 37. f. г; Britz.
Lact. 7. 76 ; Cooke, Br. Fungi, ai 991 ; Gillet, Champ. Fr. ai,
48 (157; 399); Pat. Tab. Analyt. Fung. ai 209.

DISTINGUISHING FIELD-MARKS: The gray or brownish-gray,
very viscid pileus, the paler stem, and the lilac color which all
parts of the plant assume where touched. The pileus may be
faintly zonate but is never conspicuously zoned or spotted as in
Lactaria maculata Peck, and the plant is rarely as large as the
smallest specimens of Lactaria maculata.

Most European mycologists describe the latex as changing
color upon exposure to the air. In Monogr. 2: 162, Fries says,

50 LACTARIAE ОҒ THE UNITED STATES

“ Lac raro album persistit.” It was evidently on the strength of |

the unchangeable milk that Passerini described his specimens as
a distinct species, Г. Z/vescess. In the Vermont specimens I
repeatedly watched the latex for fifteen minutes or even longer,
and I could never detect any change except where the drop was
in contact with the broken flesh.

Krombholz’ description, accompanying his al, 57. f. 7—9,accords
with the characters of Lactaria lividorubescens but the plate does
not represent this species, nor does it agree with his description.
And ai 57. f. 14, 16 can scarcely be positively referred to Lac-
taria lividorubescens, hence I have omitted both of these in the
plates cited.

23. Lacraria MACULATA Peck, Ann. Вер. М. Y. St.
Mus. 41: 74. 1888. [As Lactarius]

Pileus fleshy, compact, convex-umbilicate, then depressed in the
center or at length infundibuliform, gray to lilac-gray, distinctly
zoned with concentrically arranged darker spots, viscid, glabrous,
7-5-12.5 cm. broad, margin involute, naked, then spreading ; gills
whitish or cream-colored, sometimes forking, close, adnate 10
decurrent ; stem of the same color as the pileus or paler, spotted,
equal or tapering downwards, glabrous, hollow, 2.5-5 cm. long,
10-16 mm. thick; flesh grayish becoming lilac where wounded ;
spores subglobose, echinulate, 10—12.5 и т diam. ; latex creamy-
white, becoming lilac, acrid and unpleasant.

Нав. : “Іп thin woods, and pastures " (Peck).

Distris.: New York, Peck ; Vermont, Morgan.

DISTINGUISHING FIELD-MARKS: This species is to be distin-
guished from Lactaria lividorubescens Batsch by its larger size, the
firmer flesh, and the conspicuously spotted-zoned pileus and the
absence of an umbo. The stem is also more spotted. 5

Lactaria maculata was first described as a variety of Lactari@

uvida Fr., under the name magnus, but after further field-work, 3

was separated as a distinct species. I have not found this species,
but from an examination of the type specimens at Albany, I con-
sider it to be distinct from L. /ividorubescens.

VII. ĪNSULSAE

Pileus very viscid, entirely glabrous, some shade of yellow, |

flesh firm, plants rather large ; gills becoming neither perceptibly
darker with age, nor pruinose ; latex very acrid, white, unchanging

LACTARIAE OF THE UNITED STATES 51

Synopsis of species

Pileus azonate, spores white. 25. Г. affinis.
Pileus zonate. :
Spores yellowish. 24. L. insulsa.
Spores white, L. зопата.*

24. LACTARIA INSULSA (Fr.) Fr. Epicr. 336. 1838. [As Lac-
tarius.] — Hennings, in Eng. & Prantl, Nat. Pflanzenfam.
pt 21O: 1608

Agaricus insulsus Ет. Syst. Myc. т: 68. 1821.

Pileus fleshy, firm, at length somewhat lax, convex-umbilicate,
becoming depressed in the center, coppery-orange ( 56) with alter-
nating zones of deeper and lighter tones, sometimes yellowish-buff
(370) zoned with coppery-orange, center coppery-orange to red-
ochre ( 32. 7. 4), viscid, glabrous, surface not polished, and often
marked with striae like the stem of а Russula, 4.5—10 cm. broad,
margin involute at first and remaining arched until the last; gills
white, becoming dull-reddish along the margins where rubbed,
frequently forking near the stem, adnate, becoming decurrent as
the pileus deepens, thin, fragile, 2 mm. broad ; stem paler than the
pileus, sometimes spotted with deeper color, tapering downwards,
glabrous, stuffed, then hollow, 1.5-5 cm. long, up to 10-12 mm
or more thick ; flesh white, no odor; spores pale Naples-yellow
(29..& f, 2), globose, теке echinulate, 7-8 и, rarely би; latex
white, unchanging, very a ficious.

Has. : On the eeng in rather open woods ог by trails in
mixed deciduous woods. July and August.

DisrRiB.: New York, feck ; Maine, White; Vermont, Burt;
North Carolina, 1,000 meters elevation, Burlingham 37, 1907 ; Ala-
Бата, Underwood, Earle ; Missouri, G/affelter 303.

Пловт.: Berk. Outl. A. 73. /. 2; Bres. Fung. Mang. ai 62;
Cooke, Br. Fungi, A. 975 (good); Gillet, Champ. Fr. ai 396;
Agaricus flexuosus Hussey, Шив, Br. Myc. A. 59 (good);
Krombh. Abbild. ф/. 72. f. 1-6 (good); Lanzi, Fung. Mang. ad
56. f. т, a, 6, c (good); Rich. & Rose, Atl. Champ. Fr. A. 37.
f. 10-12.

DisTINGUISHING FIELD-MARKS : The reddish or coppery-orange
color of the glabrous viscid pileus, which is marked more or less
prominently with zones, especially toward the naked, involute
margin, the paler, usually spotted stem, the whitish gills, and the
unchanging acridlatex. The color of the pileus is sometimes more

* European species, the occurrence of which in the United States is doubtful, |

59 LACTARIAE ОЕ THE UNITED STATES

yellowish, and according to Peck, may be nearly straw-colored,
but the prevailing color of the North Carolina and Alabama speci-
mens was like that described in the European plant.

25. LACTARIA AFFINIS Peck, Rep. N. Y. St. Cab. 23:
116. 1873.* [As Lactarius]
Lactarius platyphyllus Peck, Ann. Rep. N. Y. St. Cab. 23: 118.
1873

Pileus fleshy, firm, convex-umbilicate, then plane to depressed
in the center, ochraceous-yellow, azonate, very viscid in wet
weather, glabrous, 6-12 cm. or more broad, margin involute, then. |
merely arched, even, naked; gills whitish or cream-colored, not — |
changing with age, but often becoming pruinose in drying, many |
forking near the stem, not very close, adnate to slightly decurrent,
4-10 mm. broad; stem yellowish, paler than the pileus, often
spotted, nearly equal, viscid when wet, stuffed, becoming hollow,
.3-6 cm. long, 10-22 mm. thick ; flesh white, unchanging ; spores
whitish, globose to broadly elliptical, echinulate, 8 x той; latex
white, unchanging, acrid.

Нав.: On the ground in mixed balsam and maple woods.
August and September. (Pastures and copses, October, Peck.)

DisrRiB.: New York, Peck; Vermont, Burlingham 60, 1906 ;
Massachusetts, Morris ; Maine, White.

DISTINGUISHING FIELD-MARKS: The dull ochraceous, zoneless

green on the gills. The species is very closely related to Lactaria
insulsa Fr., but its constantly duller color and the absence of zones,
together with the broader and more distant gills, seem to separate
it as a distinct species.
Lactarius platyphyllus Professor Peck has decided is a large
form of Lactaria affinis, and I have accordingly given it as a SI" |
onym.
DOUBTFUL SPECIES
LACTARIA ZONARIA (Lamarck) Fr. Еркг. 336. 1838. - [As L4
tarius.| — Schröt. in Cohn, Krypt.-Fl. Schles. 3: 538. 1889
Agaricus zonarius Lamarck, Fl. Fr. 1: (108). 1778. |
This species has been reported from Vermont by Frost, Rhode
* Separate published in advance in [Ap] 1872. — и

"e

лаа TS ын адас сылы сынса ааа дын

LACTARIAE ОЕ THE UNITED STATES 53

Island by Bennett, New Jersey by Ellis, Ohio by Morgan, Wis-
consin by Bundy, Minnesota by Johnson, and from California by
Harkness & Moore. I have seen only the Ellis specimens, in Ellis
& Everhart, N. Am. Fungi, second series 7916, and they do not
resemble Z. sonaria. The species is closely related to Г. 7изшза
Fr., from which it differs in the firmer flesh, the unspotted stem, and
the white spores. It is regarded by some as poisonous.

VIII. TRIVIALES

The triviales are characterized by the slimy-viscid glabrous
pileus having the margin naked, and by acrid white latex. The
latex does not change color, but in some species it stains the flesh
or gills a pale dull gray-green. Г. trivialis seems to be the most
widely distributed species.

Synopsis of species
Pileus red 28. Lactaria hysgina.
Pileus not red.
Azonate ; wounds of the gills becoming greenish.
Pileus gray to putty-colored, spores yellow. 26. Lactaria trivialis.
Pileus sepia, spores white. 29. Lactaria mucida.
Zonate ; wounds of gills not changing color.
Pileus gray, zoned with raw-umber, fading to gray. 27. Lactaria circellata.

26. LACTARIA TRIVIALIS (Fr.) Fr. Epicr. 337. 1838. [As Lac-
tarius.]— Schrót. in Cohn, Krypt.-Fl. Schles. 3: 538. 1889

Agaricus trivialis Fr. Obs. Myc. 1: бр авт
Lactarius deflexus Lindblad, Monogr. Lact. Suec. 8. 1855.
Pileus fleshy, firm, depressed in the center with the margin
arched, then plane to infundibuliform, plum-colored or smoke-gray,
tinted with livid when young, fading, becoming putty- colored (377.
г. Г) to snuff-brown (303. 2. 7), ог center darker with lilac tints
(106), azonate, viscid, glabrous, up to 15 cm. broad, margin even,
thin edge involute for some time, pruinose at first; gills cream-
colored to yellowish, some forking, close, slightly decurrent, 5—7
mm. broad; stem cream-yellow, nearly equal but often inflated,
glabrous, smooth, becoming hollow, 4-12 cm. long, 1-2 cm. thick ;

unchanging, acrid.

Нав: In mixed or deciduous woods. July to September.
DISTRIB.: New York, Peck, Burlingham 3, 1905; Vermont,
Burlingham, Frost; New Hampshire, Р. Wilson ; Connecticut,

54 LACTARIAE ОЕ THE UNITED STATES 1

Earle, Hanmer 1826; Pennsylvania, Herést ; * Virginia, 7 50 meters, al
Murrill 173; Tennessee, 400 meters, Murrill 592; North Caro-
lina 1066 meters, Burlingham 13, 1907 ; Missouri, Glatfelter 1249. -

Плозт.: Britz. Lact. f. 9 ; Cooke, Br. Fungi, 77. 976 ; Krombh.
Abbild. ai 74. f. 17, 18; Lucand, Champ. Fr. ai 166.

DISTINGUISHING FIELD-MARKS: Тһе broad thick grayish
pileus, which is depressed in the center, viscid, glabrous, and with-
out zones; the paler stout stem ; the creamy-white to yellowish
gills, which become pale-greenish where the latex dries, or some-
times sordid where bruised ; and the very acrid milk. L. ¢rivialis
Fr. differs from Г. mucida Burl. in being very much larger, gray-
ish instead of sepia, and in having yellowish spores. *

One form of this species may be recognized. It has been |
described by Peck аз L. trivialis gracilis and differs in its small | :
size and slender character (pileus 2.5-5 cm. broad, stem equal to
or larger than the diameter of the pileus). Peck has described оле l
other variety, maculata but I should hesitate to consider this 2
form of 2, trivialis, or even a variety in the old sense, because І |
can find no mention of a zonate characteristic in the European | |
plant, and I have never found Z. trivialis with any indication of i
zones. І am inclined to regard this as either a specimen of 2.
circellata or as an undescribed species.

27. LACTARIA CIRCELLATA (Fr. Fr. Epicr. 338. 1838. [As
Lactarius.|— Hennings, in Eng. & Prantl, Nat. Pflan-
земат. 1'**: 216. 1808

margin even, inrolled and pruinose at first ; gills white to cream-
colored, margins ochraceous when bruised, forking near the stem, |
close, adnate with a decurrent tooth, appearing more decurrent
with age, up to 5 mm. broad; stem paler than the pileus, equ

e the cuticle ; spores white, subglobose, slightly есїїпї »
-5 t X 7-8 p, somewhat ine ; i ing, асп
7-8 м ewhat hyaline ; latex white, unchanging, ae" 4

* In the Herbst collection this is labeled Z. turpis (Weinm. ) Fr.

LACTARIAE ОЕ THE UNITED STATES 55

Нав.: In moist mixed woods. August and September.

DISTRIB. : Vermont, 500 meters elevation, Burlingham 50,
1906.
Плот. : Omphalomyces circellatus acris Batt. Fung. Arim. pl.
13. f. Р; Cooke, Br. Fungi, ai, 990 ; Gillet, Champ. Fr. ai 380 ;
Sowerby, Eng. Fungi, ai 203.

DISTINGUISHING FIELD-MARKS: This species may be distin-
guished from Lactaria pyrogala and Lactaria flexuosa by its viscid
pileus, and from Lactaria trivialis by its zonate pileus and white
spores. Although zoned and streaked with umber at first, the
pileus fades to an even rather pale gray when old Іп this state
it can be distinguished from 2. ¢rivialis by the absence of yellowish
color in the center of the pileus, the unchanging gills, and the
white spores.

28. LACTARIA HYSGINA (Fr.) Fr. Epicr. 337. 1838. [Аз Zac-
tarius.] — Hennings, in Eng. & Prantl, Nat. Pflan-
zenfam. 1'**; 216. 1898

Agaricus hysginus Fr. Syst. Мус. 1: 67. 1821.

Pileus firm, fleshy, convex, then umbilicate, becoming de-
pressed in the center, reddish-incarnate or blood-red-brown (337),
darker in the center, shading into paler and even incarnate (739.
1. Г) at the margin, fading with age, azonate, smooth, very viscid,
viscidity persisting for some time, glabrous, smooth, 5-8 cm.
broad, margin involute then arched and the extreme edge inflexed ;
gills white or creamy-white, becoming yellowish, often forking
near the stem, close, adnate to slightly decurrent, 3-5 mm. broad ;
stem paler than the pileus, or sometimes of the same color, spotted
with red or reddish-brown, nearly equal, viscid when wet, glabrous,
stuffed, becoming hollow; flesh white in the pileus, reddish next
the cuticle, faintly yellowish in the stem ; spores whitish, subglob-
ular to elliptical, echinulate, 6-8 p x 8-10; latex white, un-
changing, acrid.

Нав.: In moist woods, especially near spruce trees, often in
grassy places. July to September.

Півтків.: New York, Peck; Maine, White; Vermont, Bur-
lingham 57, 1906.

Плозт.; Britz. Lact. f. 15 (poor); Cooke, Br. Fungi, 2/. 989;
Fries, Icon. pl. 160. f. 2; Agaricus vietus Krombh. Abbild. ai
I4. 2 1$, 10.

56 LACTARIAE OF THE UNITED STATES

DISTINGUISHING FIELD-MARKS: Тһе red-brown azonate, very
viscid pileus, acrid latex, and the medium-large size of the plant.

29. Lactaria mucida sp. nov.

Pileus fleshy, rather thin, convex-umbilicate, then plane, at
length infundibuliform, warm-sepia (305. 2. 2, 3) in the center,
putty-colored ( 377) to stone-colored ( 372) on the extreme margin,
azonate, very viscid and shining when wet, glabrous, 3-9 cm.
broad margin, even at first, slightly wavy and striate in the old

FIGURE 7. Lactaria mucida Burl. No. ог, 1907.

plant › gills white, scarcely changing color with age, staining blue-
greenish-gray where the milk dries (249. Ё. т), sometimes forking
near the stem, close, adnate to slightly decurrent, acute at the
inner end, up to 7 mm. broad; stem of the same color as the
pileus or paler, tapering upwards, slightly viscid when wet, glab-
rous, sometimes with faint striae, stuffed, then hollow, 4-7 ©:
long, 7-10 mm. thick at the top, 10-15 mm. а! the base; flesh
white, odor none; spores white, broadly elliptical, echinulate,

) th

”_ 9 вы ит. я . .
-Вих 8-9.5 и; latex white, drying blue-

ills. ' 'reenish-gray on
gills and the broken flesh, acrid.

SS

(FIGURE 7.) 4

Нав.: Under hemlocks, іп wet weather. Late August an
September.

“I D

LACTARIAE OF THE UNITED STATES 57

DISTRIB. : Vermont, Burlingham Qr, 1906; North Carolina,
1000 meters elevation, Burlingham 92, 1907 (type).

DISTINGUISHING FIELD-MARKS: This species may be easily
recognized by the contrasting dark-sepia color of the center, the
whitish color of the margin of the pileus, and its slimy shining
appearance, as well as by the blue-greenish-gray color of the
dried latex. It closely resembles Lactaria cinerea Peck in size and
texture, but differs in the habitat, in the color of the pileus, and in
the change in color of the more acrid latex. While Z. cinerea is
most abundant in August, Z. mucida does not appear until late in
August and is most plentiful in September. І have found it only
under hemlock trees, while 2. cinerea seems to grow only under
beech trees. From 2. тотай it may be distinguished by the white
spores, the smaller size, the morelax flesh, and the absence of lilac
tints in the color of the pileus, which does not become yellowish in
fading.

IX. DELICIOSAE

Latex deep-colored and rathér mild, pileus viscid; wounds, or
even the entire plant when old, often becoming greenish ; spores
yellowish.

Three species have been described from Europe, Zactaria del-
ictosa, Lactaria sanguiffua, and Lactaria haemorrhea Lowe, the
first alone being common to Europe and America. Тһе remain-
ing three species have been reported only from North America.
All of the species resemble Lactaria deliciosa L. in many respects
and form with it a natural group of which it may be taken as the
type. Lactaria Chelidonium is most closely related to Lactaria
deliciosa, while Lactaria subpurpurea appoaches Lactaria sanguiflua.
Lactaria Indigo diverges more from the type. JL. salmonea Peck
seems to belong rather with the Fudiginosae.

Synopsis of species

Latex¥orange-colored. зо. Lactaria deliciosa.
Latex'saffron-yellow. 31. Lactaria Chelidonium.
Latex dark-red 32. Lactaria subpurpurea,

Latex indigo-colored. 33. Lactaria indigo.

58 LACTARIAE ОЕ THE UNITED STATES E

30. LACTARIA DELICIOSA (L.) Fr. Epicr. 341. 1838. [As Lac

tarius.| — Schrót. in Cohn, Krypt.-Fl. Schles. 3:
543. 1889
Agaricus deliciosus L. Sp. Pl. 1172. 1753.

Pileus fleshy, firm, convex-umbilicate, becoming plane, а |
length infundibuliform, orange, yellow-orange or paler, zoned with —

deeper orange, becoming paler with age, sometimes mixed with
grayish and greenish tints, viscid when wet, glabrous, surface

somewhat roughened, 5-12 cm. broad, margin even, glabrous, |
involute, then arched, at length upturned ; gills deep-orange with |

yellowish reflections, paler when old, and becoming greenish with
age or where bruised, many forking near the stem and shorter

gills forking into the longer, often connected with cross veins at the |
base, close, somewhat decurrent, rather narrow ; stem of the same E
color as the pileus, spotted with brighter orange, nearly equal, `

glabrous, or sometimes a little hairy at the base, smooth, stuffed,

becoming hollow, 2.5-10 cm. long, 8-12 mm. thick ; flesh yd- +
lowish, staining greenish next the gills and the exterior of the `
stem ; spores yellow, subglobose to elliptical, slightly echinulate, |
more ог less hyaline, 8-8.5 иж 8-11 и; latex orange to red- |

orange, aromatic and somewhat acrid. Ее.

Has. : In moist woods, especially under firs and hemlocks. |
August and September, or up to December and January in Florida. -

Distris.: New York, Peck, Fisher, Burlingham то, 1905) |
Maine, White; Vermont, Burlingham 44, 1906, Jones; Con-
necticut, Earle, Underwood ; Alabama, Earle ; Florida, Fawcett’
Colorado, 2680 meters elevation, Clements. It has also been
reported from North Carolina by Schweinitz and by Curtis ; from |

Ohio by Morgan; and from California by Harkness & Moore.

Плозт.: Atkinson, Stud. Am. Fungi, ai 35. f. г; Badham, |
Escul. Fung. Eng. ai 6. f. 2; Вапа, Champ. Nice, pl. 19. f. 1-3” |
Bern. Champ. Roch. ai 39. f. 1; Boyer, Champ. pl. 32; Вгезй“.
dola, Fung. Mang. ai 64; Britz. Lact. f. 17; Cooke, Br. Fungh |
pi. 982 (very good); Cordier, Champ. Fr. ai 25. f. г (poor); Eng. |
& Prantl, Nat. Pflanzenfam. т!**. 7. тто D; Fl. Dan. pl. ГІЗ
Gauthier, Champ. ai rr. f. 1 (bad); Gibson, Edible Toadstools and |
Mushrooms, ai 18; Gillet, Champ. Fr. ai 49 (166; 382] (мегу -
good); Hahn, Der Pilz-Sammler, f. 20; Harzer, Pilze, pl- 10.
(good); Hussey, Illust. Br. Fung. 1: A. 67; Krombh. Abbild. |

pl. 11 (good); Lanzi, Fung. Mang. ai 53. f. 2 (good) ; Lorinsefy

LACTARIAE OF THE UNITED STATES 59

Essb. und Gift. Schwam. A. 3. f. 5; Lucand, Champ. Fr. ai 167
(good) ; McIlvaine, ai 27. f. 3 (poor); Моше! & Dass. Champ. ai
18. f. A (bad); Rich. & Roze, Atl. Champ. Fr. X. 38. f. 1-5;
Rolland, Bull. Soc. Myc. Fr. 7: 2/ 2. f. 2; Schaeff. Fung. Bav.
Icon. ai гг; Sicard, Hist. Nat. Champ. A. 44. f. 237 (bad);
Sowerby, Eng. Fungi, ai 202 (good); Fries, Sverig. Svamp. ai.
б (very good); Venturi, Studi Micol. f. 55, 56; Vittad. Descr.
Fung. Mang. AX. 42; Viviani, Fung. Ital. ai 73.

Ехвіс.: Herpell, Sammlung prápariter Hutpilze 707;
Karsten, Fungi Fennici, 509 ; Sydow, Mycotheca Marchica 670.

DISTINGUISHING FIELD-MARKS: Lactaria deliciosa varies іп
color from orange to pale-yellow and is sometimes conspicuously
zonedand again nearly azonate, but the coloris always brighter and
the zones are usually more marked than in Lactaria Chelidonium,
and, while the latex may at first be saffron-yellow, it soon becomes
red-orange and is always abundant. The color and the zonation
of the pileus are most decided in the young plant. The change in
the color of wounds from orange to greenish is often rapid, the
whole plant becoming greenish where bruised or as it becomes
old. At other times I have not been able to detect any greenish
tint to the gills or other parts of the plant either with age orinjury.
The green color was very prominent in the specimens sent from
Florida by Fawcett and it persisted in the dried mushroom,

31. ГАСТАВА CHELIDONIUM Peck, Ann. Rep. N. Y. St. Mus.
24: 74. 1872. [As Lactarius.] — Hennings, in Eng.
& Prantl, Nat. Pflanzenfam. 1!**: 218. 1898

Pileus fleshy, firm, convex, then plane with the center more or
less depressed, “ grayish yellow or tawny,” at length stained with
bluish and greenish, usually with two or three narrow zones near
the margin, slightly. viscid when wet, glabrous, 5-8 cm. broad,
margin involute at first and naked; gills saffron-yellow mixed
with gray, sometimes forking, close, ‘‘anastomosing or wavy at
the Базе,” adnate, then slightly decurrent, narrow ; stem of the
same color as the pileus, nearly equal, glabrous, becoming hollow,
2.5-4 cm. long, 10-12 mm. thick; flesh whitish, staining saffron-
yellow from the latex, then becoming bluish and at length

reenish ; spores yellowish, globular to broadly elliptical, echinu-
late, 7x 8 р (ди Peck); latex saffron-yellow, mild, scanty. Eae.

60 LACTARIAE OF THE UNITED STATES
Нав.: “ Sandy soil under or near pine trees ” (Peck); also in
dry spruce woods. 7
Distris.: New York, Peck, Earle; Vermont, Burlingham; |
Connecticut, Underwood; Alabama, Earle; it has also been re- |
ported from Vermont by Frost and from North Carolina Бу. |
Atkinson. :
Плоэт.: Atkinson, Stud. Am. Fungi, AX. 35. f. 2. :
DISTINGUISHING FIELD-MARKS: lt is a paler yellow and duller |
in color than Lactaria deliciosa, the flesh is firm, the pileus scarcely |
viscid, the zones marginal, the stem short, the gills narrow, and
the latex saffron-yellow rather than orange. It is usually found
in dry woods in the vicinity of pine trees, while Lactaria deliciosa
is most abundant in mossy wet woods, especially near hemlocks.
The type specimens are in the herbarium of the N. Y. State.
Museum at Albany.

32. ГАСТАВТА SUBPURPUREA Peck, Ann. Вер. М. Y. St. Mus.
_ 29: 43. 1878. [As Lactarius.] — Hennings, in Eng. — |
. & Prantl, Nat. Pflanzenfam. 1!**: 218. 1898
Pileus fleshy, convex umbilicate, then plane to infundibulifot
dark-red (Indian-lake, 705. 2. г) zoned with hydrangea-pink (77

pruinose at first, then plane to uplifted, gills colored like the pileus |
and fading and greenish with age, seldom forking, rather distant
slightly decurrent, 6-7 mm. broad ; stem of the same color as the
pileus, often spotted with dark-red, equal ог tapering upwards,
glabrous or pruinose, sometimes tomentose at the base, stuffed,
becoming hollow, 3-7 cm. long, 8-15 mm. thick; flesh whitish
to hydrangea-pink, instantly staining red next the gills and the
surface of the stem, after some time becoming greenish, 0401
ip етар, broadly elliptical, echinulate, often Kei |
guttulate, 7-5 u x 8-10 p; latex Mo - Li PED
mild. Edible. (тои 8) pene uec |
Нав.: Usually moist woods in the vicinity of hemloc
August, September, and October.
Пізтків, : New York, Peck, Burlingham 9, 1905; Vermo
Burlingham 109, 1906 ; Massachusetts, Morris; Connectic
Underwood ; North Carolina, Burlingham 51, 1907.

LACTARIAE ОЕ THE UNITED STATES 61

IrLusr.: Ann. Rep. М. У. St. Mus. 54: X. 70. f. 1-6.

DISTINGUISHING FIELD-MARKS: The peculiar mixture of dull
dark-red, hydrangea-pink, and grayish in the coloring of the plant,
and the dull dark-red latex. It has been found at elevations
varying from 130 meters in New York to 1000 meters in North
Carolina.

э en

FIGURE 8. Lactaria subpurpurea Peck. No, 51, 1907, Burlingham.

Bresadola’s figure of Lactaria sanguifiua Fr. in Fungi Triden-
tini 2: pl. 126. 1892, closely resembles Lactaria subpurpurea Peck,
but his description of the color and the taste of the milk as well as of
the color of the pileus would not indicate that he had that species
in hand. The figure, however, seems to accord with L. subpur-
purea much more than with Lactaria sanguiffua Pr.

33. Lacraria Innico (Schw.) Fr. Epicr. 341. 1838. [As Lac-
tarius.]— Hennings, in Eng. & Prantl, Nat. Pflanzenfam.
144%; 218. 1898
Agaricus lactifluus Indigo Schweinitz, Syn. Fung. Carol. Super.

бі. 1818.

62 LACTARIAE OF THE UNITED STATES

Pileus fleshy, firm, convex-umbilicate, becoming plane to infun- |

dibuliform, indigo-blue when wet, fading when drier, often with a |
grayish luster, zonate, sometimes spotted, 5—12 mm. broad, margin
naked; gills indigo-blue or paler, at length yellowish or sometimes |
tinged with green, some forking near the stem, close, somewhat |
decurrent, rather broad ; stem of the same color аз Ше pileus, often `
spotted, nearly equal, glabrous, smooth, firm, becoming hollow,
up to 5 cm. long, 1-2 cm. thick; flesh sky-blue to darker, often |
becoming greenish; spores yellowish, globular to broadly ellip-
tical, echinulate, 7 м x 8-0.5 м; latex dark-blue. Edible.

Has.: In dry pine or oak woods. July to October (and -
November in Florida).

Півтків.: New York, Peck; Connecticut, Underwood, Han- |

те’; Vermont, Jones ; Maryland, Banning ; Virginia, Murrill; | |
North Carolina, Schweinitz ; South Carolina, Ravenel ; Alabama, ` |
Underwood; Georgia, Harper; District of Columbia, Murrill

1522; Florida, Fawcett. It has also been reported from Pennsyl- ;
vania by Herbst. ”
Плозт.: Atkinson, Stud. Am. Fungi, ai 35. f. 3; Banning, |
Folio Md. Fungi, ai 82 (excellent); McIlvaine p/. ут. f. 2.
DISTINGUISHING FIELD-MARKS: The more or less indigo color
or tint which characterizes the entire plant.

X. QuiETAE 1

Pileus covered with a thin viscidity when moist, soon dry, su.
entire, glabrous, and margin naked, flesh more lax and the e :
usually smaller than in Limacina ; gills becoming decidedly GC 2
with age, and dusted with the spores; latex mild or only slighty |
acrid,

In these plants the viscidity disappears so quickly that it may ;
not be noticed in dry weather unless the mushrooms are collected |

in the morning while covered with dew.

Synopsis of species

* Pileus 2-4 cm. broad.

Latex mild.
Pileus mahogany-red, shining-viscid when moist, 35. Lactaria nitida. |
Pileus fulvous, fading, papilla persisting dark, 36. Lactaria we ШШ
Pileus brownish-drab to yellowish-brown, fading. 38. Lactaria paludinetti i
Latex acrid.

ET RES

Pileus fulvous, margin crenate to sulcate. 37. Lactaria minuscule.

DOM сайын сиал ааа ны ER EE ET E SET

LACTARIAE ОЕ THE UNITED STATES 63

** Pileus 5-15 cm. broad.

Latex mild.
Pileus dark-red to reddish-cinnamon. 34. Lactaria quieta.
Pileus pale leather-colored. Lactaria pallida. *

34. ГАСТАВТА QUIETA Fr. Epicr. 343. 1838. [As Lactarius.]—
Schrót. in Cohn, Krypt.-Fl. Schles. 3: 537. 1889
Agaricus quietus Fr. Syst. Myc. 1: 69. 1821.

Pileus fleshy, rather thick, depressed in the center, dark-red to
reddish-cinnamon, fading, obscurely zonate, viscid at first, soon dry,
glabrous, 5-8 cm. broad, margin arched, even ; gills white, becom-
ing pale brick-red, sometimes forking near the stem, close, adnate
to decurrent, 3-4 mm. broad; stem colored like the pileus, at
length rubiginous, equal, glabrous, smooth, stuffed, at length hol-
low, 5-8 cm. long, up to 15 mm. thick; flesh white, becoming
tinged with fulvous where broken, odor slight but disagreeable ;
spores white, echinulate, 8-тои x 6-7 p; latex white, unchanging,
mild. Ва.

Нав.: In deciduous woods.

Distris.: New York, Peck.

Плозт.: Cooke, Br. Fungi, ai 983.

DISTINGUISHING FIELD-MARKS: This species may be distin-
guished from the other dark-red or fulvous-colored species in this
group by the thicker pileus and stouter stem, and the faint zona-
tion. The viscidity soon disappears, and Romell says: “I have
never found this plant viscid.”

35. LacraRIA NITIDA Burl. Bull. Torrey Club 34: 89. 1907.
[As Lactarius]

Pileus fleshy, rather thin, convex, then depressed in the center,
umbonate, mahogany-red (335) more golden-red toward the mar-
gin, umbo persistently dark, otherwise fading when dry, shining-
viscid when wet, glabrous, smooth, 3 cm. broad, margin remaining
arched for some time, even; gills yellowish, becoming pruinose, some-
times forking, close, slightly decurrent, broad; stem mahogany-red,
equal or sometimes ventricose when growing in wet places, stuffed,
becoming hollow, 4.5-5 cm. long, 5-8 mm. thick ; flesh faintly
buff, turning a little reddish where cut; spores white, echinulate,
5-6 p. x 6-8 p ; latex white, unchanging, mild.

Нав.: In grassy sheep pasture near hemlocks and in moist

woods, September.

* European, occurrence in United States doubtful.

64 LACTARIAE ОЕ THE UNITED STATES

Півтків.: Vermont, 500 meters elevation, Burlingham 114, |
1906. a
DISTINGUISHING FIELD-MARKS: The mild milk and the ma- |
hogany-red color of the pileus and stem, together with the shining - |
varnished appearance of the pileus when moist and viscid, make | |
this little plant easily recognizable. m

The type specimens are in the herbarium of the N. Y. Botanical |
Garden. 1

36. LACTARIA OCULATA (Peck) Burl. Bull. Torrey Club 34: 8%
1907. [As Lactarius] 5
Lactarius subdulcis oculatus Peck, Bull. N. У. St. Mus. 67: 37.

Pileus fleshy, thin, convex, then depressed іп the сеп
umbonate, rich-fulvous in the center, shading to buff-fulvo
toward the margin, all except the papilla fading to pinkish w
age, viscid in dew or wet weather, glabrous, smooth, 1.5-2.5
broad, margin involute and pruinose at first, then arched
finally nearly plane, slightly crenate ; gills whitish, then yello |
at length pruinose, a few forking near the stem, close, slightly
decurrent, rather broad, stem buff at the top, sublatericeous bel "
the middle, equal, slightly viscid when wet, tomentose at the base,
stuffed, up to 6 cm. long, 5 mm. thick, flesh buff-whitish ; spo
white, broadly elliptical to subglobose, echinulate, 6-7.5 А X
8-0.5 м; latex white, unchanging, mild.

Нав, : Under pine, hemlock, spruce, and balsam-fir trees, ой
in moss. July to September. a

DisrRIB.: New York, Peck ; Vermont, 500 meters elevation
Burlingham 107, 1906.

Плозт.: Peck, Bull. М. У. St. Mus. 67: pl. 83. f. 20-26 И

DISTINGUISHING FIELD-MARKS: The thin viscidity and fulvous
color of the moist pileus, and the dark-fulvous or chestnut-colo
spot which remains in the center of the pileus when the рам.
dry. The viscidity and the expallent color separate this species |
from Г. subdulcis (Pers.) Fr., while its larger size, paler color, mild
milk, and persistent "eye-spot" serve to distinguish it from 1.
minuscula Burl, _

LACTARIAE ОЕ THE UNITED STATES 65

37. LACTARIA MINUSCULA Burl. Bull. Torrey Club 34: 88. 1907.
[As Lactarius |

Pileus fleshy, thin, broadly convex, with a small umbo, becom-
ing plane then somewhat depressed in the center, fulvous in the
center, cinnamon (323. 4. 1) toward the still paler margin, azonate,
viscid in wet weather, sometimes shining with viscidity, glabrous,
1-3 cm. broad, margin minutely crenate sometimes sulcate, often
slightly wavy, pruinose at first; gills whitish, seldom forking,
dow adnate or decurrent by a tootit, broad бог the thickness ep
the pileus ; - stem fulvous near the base but paler toward the pileus,

FIGURE 9. Lactaria minuscula Burl. FIGURE 10. Lactaria minuscula Burl.

No. 56, 1907. Aa, 56, 1907.

equal, glabrous, sometimes ок, at the eg when growing

in moss, stuffed, becoming hollow, 2.5-4.5 long, 3-4 mm
thick ; flesh isabelline-white ; spores pictas ылоо, slightly
echinulate, 6-8 м; latex white, unchanging, acrid. (FIGURES
9, 10).

Has.: In moist woods, in moss ог on decayed wood, under
yellow birches, black gum, and black oak. July and August.

Distris: New York, Peck; Vermont, 500 meters, Burling-
лат; North Carolina, 1,000 meters, Burlingham 56, 1907.

DISTINGUISHING FIELD-MARKS: This species differs from all
others in this group in its small size, crenate margin, and more
acrid latex. It may be distinguished from Г. subdulcis by its
viscid pileus, and by being expallent. It is frequently solitary.

66 LACTARIAE ОЕ THE UNITED STATES

38. LACTARIA PALUDINELLA Peck, Ann. Rep. N. Y. St. Mus. 38: І
133. 1885. [As Lactarius.] — Hennings, in Eng. &
Prantl, Nat. Pflanzenfam. 1'**: 214. 1808

Pileus fleshy, thin, convex, then plane-umbilicate to depressed |
їп the center, sometimes with a small umbo, brownish-drab ( 302.
t. 2) to dark-fawn ( 307),* expallent, slightly viscid when wet, |
glabrous, 12 mm. to 4 cm. broad, margin at length slightly stri-
ate ; gills white to cream-colored, becoming darker with age, pruin- |
ose, many forking near the stem, close, adnate or slightly decur-
rent, thin, up to 4 mm. broad ; stem of the same color as the pileus
or paler, nearly equal, glabrous, except at the base, which Б
slightly villose when growing in moss, stuffed, sometimes hollow, |
2-3 cm. long, 3-4 mm. thick ; flesh white, or tinted with the color aT
of the surface; spores white, subglobose, echinulate, 6.5-8.5 №
latex white, unchanging, mild.
Нав.: In marshy places іп woods, іп Sphagnum, or in decay-
ing leaves. August. 3
Пізтків.: New York, Peck ; Vermont, Burlingham ; North
Carolina, Burlingham 82, 1907.
-DISTINGUISHING FIELD-MARKS: The sordid-brown color or
mixture of brownish-drab and yellow-brown, which gives the
pileus a mottled, streaked, and subzonate appearance, and |
striatulate margin. The species is small and is rendered inc
spicuous by its dusky coloring. It occurs only in densely shad
places.

' DOUBTFUL SPECIES

LACTARIA PALLIDA Pers, Tent. Disp. Meth. Fung. 64. 1797:
This species has been reported from Rhode Island by Bennett
Connecticut by White, and Minnesota by Johnson. I have seen
Hanmer's zo. 1399, which was cited as pallida by White, and it
is not that species, but a pale form of Z. lactifiua (L.). The plants
from which the other determinations were made are not available
for examination, and there is a reasonable doubt whether Z. pallida
occurs within the United States. It is edible.

ХІ. CiNEREAE |
Pileus more viscid than іп the Quietae, of some shade of ore
perfectly glabrous, flesh lax and thin, plants rather fragile ; gilis
becoming darker with age and pruinose ; latex soon acrid.
GE
* A. yellowish-brown color, not fulvous.

LACTARIAE ОЕ THE UNITED STATES 67
Lactaria cinerea has been classed with the Lzmacinit by Sac-
cardo, but seems to belong rather to Russu/aria on account of the
less acrid latex, the less viscid pileus, the more lax flesh, as well
as the changing gills.
d Synopsis of species
Latex white, not staining gills. 39. Lactaria cinerea.
Latex white, becoming gray. Lactaria vieta. *
39. LacTARIA CINEREA Peck, Кер. N. Y. St. Bot. 24: 73. 1872.
[As Lactarius.] — Hennings, in Eng. & Prantl, Nat. Pflanzen-
fam. п'е: 217, 1498
Pileus fleshy, thin, lax, convex at first, soon umbilicate to de-
yressed in the center, at length infundibuliform, cinereous ( 358),

I

No. 105, 1907, Burlingham.

FIGURE 11. ZLactarta cinerea Peck,
darkest in the center, becoming pale toward the margin, fading,
occasionally zonate, viscid when wet, glabrous, 2-5 cm. broad,
margin involute at first, at length arched or uplifted, even ; gills
or where bruised, often appearing pruinose,
or 2 mm. of the

--

white, not changing со!
especially in dried plants, a few forking within 1
stem, close, adnate, rather narrow ; stem of the same color as the
* European species, the occurrence of which in the United States is uncertain.

68 LACTARIAE ОЕ THE UNITED STATES

pileus or paler, subequal but more often tapering upwards, fre-
quently tomentose at the base, otherwise glabrous, spongy, soon.
hollow, 3-8 cm. long, 8-16 mm. thick ; flesh white, not changing
color; spores white, subglobose, minutely echinulate, 5.5-7 И;
latex white, unchanging, very acrid after a few seconds on the
tongue. (FIGURE 11.)

Has. : Under beeches or among beech leaves. July and August. -

Півтків.: New York, Peck, Earle ; Connecticut, Underwood |
& Earle; Maine, White ; Vermont, Morgan, Burlingham 13, |
1006; North Carolina, 1000 meters, Burlingham 105, 1907.

DISTINGUISHING FIELD-MARKS: The uniform pale-gray color, |
the thin pileus, which is glabrous, viscid, and usually azonate, but -
sometimes faintly zonate from the margin half-way to the center, |
the spongy stem, which usually tapers upwards, and the white:
close gills, which often become pruinose and do not become |
stained from the latex. I have found Lactaria cinerea only under
or in the vicinity of beech trees. In North Carolina I rare |
failed to find several specimens of it wherever even a solitary |
beech tree was growing in the midst of the oak-chestnut woods.

and a few of North Carolina plants were faintly but decidedly
zonate, the zones being narrow and more prominent toward the.
margin. Specimens collected by Morgan had this same charac
teristic. Peck considered the species to be related to Z. vieta |

and it seems to belong in the group with this rather than with the
Limacini, where it is placed by Saccardo. Lactaria mucida seems
to stand between L. cinerea and L. trivialis, but is more closely re-
lated to the latter by the more acrid milk and the very slimy pileus.

=

DOUBTFUL SPECIES

LACTARIA VIETA (Fr.) Fr. Epicr. 344. 1838. [As Zactarius.] — с
Schrot. in Cohn, Krypt.-Fl. Schles. 3: 541. 1889

This species was reported from Ohio by Morgan in his My
cologic Flora of the Miami Valley, but as the specimens are
preserved it is impossible to verify the determination. His descrip
tion would indicate that he had the species which he thought. Е
. this reason I append the following description :

Е ОРНА ча ыры даа EE ын EES

LACTARIAE ОЕ THE UNITED STATES 69

Pileus fleshy, lax, thin, convex, subumbonate, then expanding,
umbilicate to infundibuliform, incarnate, or livid-gray, fading to
grayish, azonate, viscid when wet, finely silky when dry, 4-6 cm.
broad, margin even, involute at first, then arched to uplifted ; gills
white then yellowish, thin, close, somewhat decurrent, 2 mm. broad;
stem of the same color as the pileus, equal or tapering upwards,
glabrous, dry, stuffed, becoming hollow, 5-8 cm. long, 4-6 mm
thick; flesh whitish, odor faint, but somewhat pungent; spores 8
и, nearly globose, minutely echinulate ; latex white, becoming gray,
mild, then acrid. :

Нав.: Damp woods.

Пловт.: Britz. Lact. f. 21; Cooke, Br. Fungi, a 1000;
Fries, Icon. А 170. f. т; Gillet, Champ. Fr. ai 167. [gor] ;
Lucand, Champ. Fr. 2/. 96.

Exsic. : Sydow, Mycotheca Marchica 2727.

ХП. THEIOGALAE

Pileus slightly viscid when moist, soon dry, glabrous, margin
naked or minutely pruinose-downy in the young plant when viewed
with a lens, whitish to isabelline or fulvous or pinkish-buff; flesh
rather thick and firm at first, then thinner and lax ; gills becoming
darker with age, and more or less ргиіпове; latex bitter or acrid,
white, becoming yellow.

Synopsis of species
Latex very acrid, becoming golden-yellow.
Latex mild, bitterish,'then acrid, becoming sulphur-yellow.

Pileus whitish, entire plant becoming brownish-red

>
N

. L. chrysorhea.

wit e, azonate. 41. L. colorascens.
Pileus pinkish-buff to reddish terra-cotta or
vous, zonate. 40.

f L. theiogal
Pileus pallid, azonate. 40.

gala.
L. theiogala brevis.
40. LacTARIA THEIOGALA (Bull) Fr. Epicr. 342. 1838. [As
Lactarius.] — Schrót. in Cohn, Krypt.-Fl. Schles.
3: $44. 1889
Agaricus theiogalus Bull. Herb. Fr. ai 667. f. 2. 1793; Hist. І:

1809.
Hypophyllum lateritium Paulet; Paulet & Léveillé, Icon. Champ.
59.: 1855. |
Lactarius brevipes Longyear, Rep. Mich. Acad. Sci. 3: 59. 1901.
Lactarius brevis Peck, Bull. М. Ү. St. Миз. 04: 33. 1905.
Lactarius xanthogalactus Peck, Bull. Torrey Club 34: 346. 1907.

70 LACTARIAE ОЕ THE UNITED STATES

Pileus fleshy, firm and rather thick at first, then thinner and
more lax, convex, sometimes with a small umbo, sometimes slightly
umbilicate, then plane to depressed, incarnate-isabelline to yellow-
ish-salmon (65. 2. 1—4) or reddish-terra-cotta (гоо) to fulvous, more
or less zonate, occasionally much paler and nearly or quite azonate,
viscid when wet, glossy when dry, glabrous, 5—8 cm. broad, mar-
gin inrolled at first, and pruinose to very minutely pruinose-downy
on the extreme edge ; lamellae whitish to yellowish, reddish-brown

FIGURE 12. Lactaria theiogala ( Bull.) Fr.

i . hy
No. 44, 1907, Burlingnam.

when old or when injured, some forking near the stem, close, ad-
nate or a little decurrent, up to 4 mm. broad; stem colored like
the pileus or paler, sometimes faintly spotted, glabrous except 4
the base, which is often slightly tomentose, smooth, full and rather
firm, at length hollow, 3-7 cm. long, 7-13 mm. thick ; flesh white,
becoming yellow from the latex, odor pungent, disappearing ™
drying ; spores whitish, minutely echinulate, subglobose to broadly
elliptical, 6-7 их 8—9 м, latex white, changing to sulphur-yellow,
bitterish then acrid. Suspicious, : 12.

Нав.: In botl
or in oak woods.

(FIGURE 12.

1 dry and moist woods, under spruce and fir tree
August and September.

SES ^ To r И i - . А > /, te;

DISTRIB. : New Y ork, Peck, Earle, Burlingham ; Maine, Whites

LACTARIAE ОЕ THE UNITED STATES 71

Vermont, Jones, Frost, Burlingham 30, 1906 ` Massachusetts, Davis ;
Connecticut, Earle, Hanmer 2335; Pennsylvania, Herbst; Mary-
land, Shear ; Alabama, Earle; North Carolina, Burlingham 44,
1907 ; California, Patterson.

Плозт.: Вапа, Champ. Nice, //. 27. f. 14-16; Bull. Herb. Fr.
pl. 567. f. 2; Cordier, Champ. Fr. pl. 27. f. 2; Gillet, Champ.
Fr. M. 164. [396] ; Krombh. Abbild. A. г. f. 23, 24; Lanzi,
Fung. Mang. AX. 32. f. 3, а, b,c; Paulet & Lév. Icon. Champ. f.
71. f. 1-4; Rich. & Roze, Atl. Champ. Fr. M. 37. f. 7-9;
Sicard, Hist. Nat. Champ. A. 43. f. 233.

Exsic.: Ellis & Everhart, М. Am. Fungi 7915; Arcangeli,
Erb. Син. Ital. ser. 2, $07; Herpell, Sammlung prapariter Hut-
pilze 132.

DISTINGUISHING FIELD-MARKS: This species is somewhat diffi-
cult to determine from a description because of the rather wide
variations it shows in different habitats. The latex more or less
quickly turns to sulphur-yellow and the broken flesh has at first
a pungent and characteristic odor which is an assistance in dis-
tinguishing the plant. The milk and the odor, together with
the yellowish-salmon, sometimes more or less fulvous, color of the
faintly zonate, slightly viscid pileus, will serve as determining char-
acteristics for typical forms. The viscidity disappears very readily
and the pileus is often absolutely dry and shining.

In shady dry woods, I have found the pileus to be rather pale,
varying from whitish to pale-flesh color (7 36.1. 4) and from azonate
to obscurely zonate as in my #05. 30B, 1906, and 2, 1907, while in
wet localities or in the open, I have found the color more yellowish-
salmon or approaching fulvous, and the zones well marked. Vo.
68, 1907, which was growing in a little hollow made very wet by
rain, was a striking example of this tendency. Nos. 1012 and
1357, herb. N. Y. Bot. Garden, collected by Earle, are inter-
mediate forms between such pale forms as are represented by Т.
brevis Peck or L. brevipes Longyear, and the more distinctly
zonate forms. Z. brevis Peck seems to me to be an extreme form
of Г. theiogala (Bull.) Fr., and, although the original description
gives the pileus as azonate, the type specimens show in some
cases faint indications of zones. Further, the presence of tomentum
on the stem base of 2. theiogala is not constant, but may be found

12 LACTARIAE OF THE UNITED STATES

in either the pale azonate or the darker zonate forms. While the |
size of the pale forms is usually much less than that of the darker, 1
I have found typical forms of the species as small as the size |
given for Г. órevis Peck. Тһе stem of plants of Z. theiogala grow-
ing in dry woods is usually firm and stuffed. Since the distin- 1
guishing characteristics given for Z. órevis— white or whitish |
color, smaller size, solid or stuffed stem, and absence of tomentum
from the base of the stem — аге all present in Г. ¢hetogala under |
varying conditions of habitat, and intermediate forms exist between |
the typical Г. brevis Peck апа Г. theiogala (Bull) Fr., I am in- |
clined to regard the former as а form of the latter due to eco- |
logical factors. Z. brevipes Longyear I believe to be founded on |
immature specimens of a small pale form of 2. theiogala and prob- |
ably the same as Г. brevis Peck. 1
The type specimens of Г. xanthogalacta correspond to small 4
plants of Z. theiogala. This species was described from dried |
plants collected in California and the accompanying field-notes indi-
cated that the milk was yellow, upon which character the species |
was based. But since the latex of Z. theiogala often becomes |
yellow so quickly as to seem to be yellow from the first, and `
since the type specimens of Z. xanthogalacta resemble exactly 21
similar specimens of L. thetogala, I have no doubt as to their
identity with the latter species.

Fo

41. LACTARIA COLORASCENS Peck, Bull. N. Y. St. Mus. 94: 35:
1905. [As Lactarius]
Pileus fleshy, thin, nearly plane, becoming depressed in the
center, whitish at first, becoming brownish-red with age, geen
moist, glabrous, 2-5 cm. broad; gills whitish, becoming со e
like the mature pileus, close, thin, adnate, or slightly decurrent;
stem whitish, then brownish-red, equal, even, solid, 2.5-4 сш, on
4-6 mm. thick ; spores globose, echinulate, 8 и; milk white, be-
coming sulphur-yellow, bitter. E:
Нав.: Woods. August.
DisrRiB: New York, Peck, Atkinson (Long Island). S
DISTINGUISHING FIELD-MARKS: The change in color from pallid |
in young specimens to brownish-red in the mature plants, and Ge 1
bitter white milk which becomes sulphur-yellow upon exposure 19
the air. Peck says that the color of the mature plants is 91
to that of 2. camphorata.

Wu

LACTARIAE OF THE UNITED STATES 73

The type specimens which are in the herbarium of the N. Y.
State Museum at Albany are those taken by Atkinson at Port
Jefferson, L. I. The description does not indicate whether the
pileus is viscid or dry in wet weather, but the relationship seems
to be with Г. thetoga/a (Bull.) Fr.

42. LACTARIA CHRYSORHEA Fr. Epicr. 342. 1838. [As Lacta-
rius. |— Schrót. in Cohn, Krypt.-Fl. Schles. 3: 541. 1880
Pileus fleshy, firm, rather thick, convex-umbilicate, then infun-

dibuliform, whitish to yellowish tinged with flesh color, usually

zoned with golden-yellow or pale-orange, slightly viscid when
moist, glabrous, 2.5-10 cm. broad, margin involute at first and
covered with a minute down, then spreading and naked; gills
white then yellowish, some forking, close, adnate to slightly de-
current, thin; stem white, then colored like the pileus, sometimes
with bright-colored spots, pruinose, glabrous except at the base,
which is more or less villose, stuffed, then hollow, 2.5-8 cm. long,

6-15 mm. thick; flesh white, becoming yellow where injured ;

spores white, subglobose, minutely echinulate, 7-8 и; milk white,

Te golden-yellow, very à

: Mixed woods or groves. Ligne and September.
pits : New York, Peck; North Carolina, Atkinson, Bur-
lingham 100, 1907, 1000 meters.

Плозт.: Atkinson, Stud. Am. Fungi, f. 722; Cooke, Br. Fungi,
2. 984; Gillet, Champ. Fr. ai 50 (751; 379]; Hahn, Der Pilz-
Sammler, 5. f. 20; Krombh. Abbild. A. 12. f. 7-14; Lucand,
Champ. Fr. pl. 5.

DISTINGUISHING FIELD-MARKS: The white to yellowish color
of the plant, the indistinct zones which are made up of brighter-
colored spots concentrically arranged, and the acrid milk. The
zones and spots become more prominent with age and in drying,
and while the zones may be nearly lacking in the fresh plant, they
may be conspicuous in the dried specimen. The species is ap-
parently closely related to Г. theioga/a Fr., from which it is readily
distinguished by the brighter, yellow tones, the spotted character
of the zones, the absence of odor in the fresh plant, and the acrid
milk,

This species is classed by Saccardo with the Реғай, but it

seems to me to belong rather with Russu/aria on account of its

slightly viscid pileus and the changing color of the gills. If the

74 LACTARIAE OF THE UNITED STATES

pileus were more decidedly viscid, its acrid milk would indicate its
relationship with the .SuéZmac:ina.

XIII. HELVAE

Pileus dry, varying from velvety to flocculose, floccose-squam-
ulose, or squamulose, at least at first, color bay-red to fulvous,
fulvous-isabelline, and testaceous, flesh rather thin, and lax,
except in one species; gills becoming darker with age, and pru-
inose; latex white and unchanging, acrid, or watery and subacrid
to mild

Synopsis of species
Pileus azonate.
Latex white, very acrid.
ileus red, umbonate, minutely flocculose at first, soon gla-

brous. 44. L. rufa.
Pileus red, not umbonate. 47. Г. rufula.
Latex white, acrid ; pileus fulvous to isabelline, squamulose. 46. L. alpina.
Latex watery, mild or subacrid, rarely white; pileus testaceous,
fading to isabelline, floccose-squamulose. 43. L. helva.

Pileus zonate.
Latex white, acrid ; pileus brick-red to reddish-terra-cotta, velvety.
45. L. Peckü.
43. LACTARIA HELVA (Fr.) Fr. Ерісг. 347. 1838. [As Lactarius.]
— Schrot. in Cohn, Krypt.-Fl. Schles. 3: 539. 1889
Agaricus helvus Fr. Syst. Myc. 1: 72. 1821.
Lactarius aguiffuus Peck, Кер. №. Y. St. Миз. 28: 50. 1877.
Lactarius aquifluus brevissimus Peck, Rep. М. Y. St. Mus. 5I:

298. 1897.

Pileus fleshy, fragile, convex, then plane to depressed, subum-
bonate, testaceous to isabelline, expallent, azonate, dry, the whole.
surface broken up into floccose-granulose squamules, sometimes
rivulose, 5-15 cm. broad, margin involute at first, then spreading ;
gills white, then tinted with incarnate, finally yellow, often forking,
close, decurrent, 2-3 mm. broad ; stem pale-testaceous, equal, pru-
inose, pubescent at the base, stuffed, then hollow, 5-8 cm. long, !
cm. or more thick; spores globose, echinulate, hyaline, 6-7 #3
flesh of the same color as the pileus but paler, odor faint, sweet
persistent in drying; latex white, scanty, subacrid, more often
watery and mild or subacrid. Ж г.

Нав.: In mossy rather wet woods or marshes. “Іп pines,
frequently degenerate in swampy places ” (Fries).

Пізтків.: New York, Peck (Г. aquifluus); Massachusetts, |
Morris; Connecticut, Hanmer 1469; Pennsylvania, Herbst; 1e-

Када ааа жаздадым ын рада релі да.

ИО a MET HP PESE 1. E EE ИРИНА

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4

LACTARIAE ОЕ THE UNITED STATES 75

ported from North Carolina by Schweinitz, from Maine by Ricker,
from Ontario, Canada, by Guillet.

Плозт.: Bies. Fung. Trid. ai 39, 127; Britz. Lact. f. 30
(very poor); Cooke, Br. Fungi, 2/. 994.

Exsic. : Sydow, Mycotheca Marchica 2779.

DISTINGUISHING FIELD-MARKS: The rather large size, the tawny
buff-colored, dry, floccose-squamulose pileus, the usually watery
milk, and the aromatic odor, which persists in drying.

Fries describes the latex as white except when the plant is
growing in wet places. Romell says “ The milk is watery ; I have
never found it white” ; and Stevenson writes in British Fungi,
“Те occurs most frequently in marshes with watery juice." Thus
far the American form has been found only with watery juice, and
on this characteristic Peck has described it as a distinct species,
L. aquifluus. Не says “ I have never found this plant with a white
or milky juice, and therefore I am disposed to regard it not as a
variety of Г. he/vus, but as a distinct species." Тһе type spec-
imens of Lactarius aquifluus agree т form and color with the 2779
Sydow, with Hennings’ Berlin specimens, and with specimens sent
from Stockholm by Romell. Since also the European plant occurs
most frequently with watery juice, it would seem hardly possible
to separate our plant as a distinct species, but Z. helva may be con-
sidered as showing a variation in the character of the latex from
white to watery, the latter being the usual form. It is probable
that this difference in the latex is due to ecological conditions, but
the mushroom most commonly grows in wet mossy places, and
the prevailing form of the species scarcely seems to be degenerate,
as Fries suspected ; we probably have rather a hydrophilous plant,
which in the dry condition might have more scanty white latex.
The short-stemmed form I consider to be due to habitat.

44. LACTARIA RUFA (Scop.) Fr. Epicr. 347. 1838. [As Lacta-
rius.] —Schrot. in Cohn, Krypt.-Fl.Schles. 3: 539. 1889

Agaricus rufus Scop. Fl. Carn. 2: 451. 1772.

Pileus fleshy, not very compact, rather thin, convex, umbonate,
at length infundibuliform, bay-red to rufous, not fading, azonate,
dry, minutely flocculose-silky, then glabrous and shining, 5-10
cm. broad, margin involute at first, whitish-downy, then glabrous ;
gills ochraceous, then rufous, sometimes forking, close, somewhat

76 LACTARIAE OF THE UNITED STATES

decurrent, 3 mm. broad ; stem rufous, but often paler than the
pileus, nearly equal, dry, glabrous, or sometimes pruinose and
downy at the base, stuffed, firm, at length sometimes hollow, 5-10
cm. long, 6-10 mm. thick ; flesh pallid or tinged with pink, no
odor; spores white, subglobose to broadly elliptical, slightly
echinulate, 7-8; latex white, unchanging, very acrid. Very
potsonous.

Has.: “Low woods and swamps” (Peck). August.

Disrris.: New York, Peck ; Michigan, Longyear.

Плозт.: Cooke, Br. Fungi, //. 085; Eng. & Prantl, Nat.
Pflanzenfam. 1%%:/ rro А; Fr. Sverig. Svamp. ai тт; Gillet,
pl. 163 [391]; Gauthier, Champ. A. r2. f. г, 2 (poor); Hahn,
Der Pilz-Sammler, ed. 2. f. 15 (poor); Hussey, Illust. Br. Myc.
1: pl. 75; Krombh. Abbild. ai 39. f. 12-15 ; Lanzi, Fung. Mang.
pt. saf з @ be (coloring poor); Lucand, Champ. Fr. pl. 223;
НурорйуЙит torminosum Paulet, Icon. Champ. ai 22; Rolland,
Bull. Soc. Myc.Fr. 7: pl. 2. f.3; Rich. & Roze, Atl. Champ. Fr.
Bl. 37. f. 16-19.

Exsic.: Herpell, Sammlung prapariter Hutpilze 108%; Karsten,
Fungi Fennici 226; Krieger, Fungi Saxonici 480; Sydow,
Mycotheca Marchica 609 and 2720. :

DISTINGUISHING FIELD-MARKs: The dark-red, dry, umbonate
pileus, the whitish down on the incurved margin of the young

pileus, the very acrid milk, and the large size of the plant. The |

pileus is only minutely flocculose at first and soon becomes gla-

brous and shining. The species seems to be rare in the United
States. It has been reported from Vermont by Frost and from
Rhode Island by Bennett.

45. Lactaria Peckii sp. nov.

close, decurrent by a tooth, 3 mm, broad; stem dull pale reddish-

Ге
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‘Lo61 ‘7 сол” "ung 117227 аамррот Cfr 250514

THE UNITED STATES

LACTARIAE OF

78 LACTARIAE OF THE UNITED STATES

terra-cotta, usually paler than the pileus, sometimes spotted with
reddish-brown, becoming darker where bruised, nearly equal or
abruptly smaller at the base, glabrous or merely with a whitish
bloom, firm, stuffed, becoming hollow in the mature plant, up to
4.5 cm. long, 1-2 cm. thick; flesh tinted with terra-cotta ; spores

whitish, subglobose, strongly echinulate, 7-8 и; latex white, |

unchanging, astringent, then very acrid, abundant. (FIGURE 13.)
Нав.: In moist grassy wood-trails and open places near
brooks, or even in clay-banks by wood-road, in deciduous woods,
oak and chestnut predominating. July, August, and September.
DISTRIB.: New York, Peck (Long Island), Benedict (Staten
Island) North Carolina, 1000-1400 meters elevation, Burlingham
1, 1907 (type); Alabama, Baker.

DISTINGUISHING FIELD-MARKS: The reddish-brick-colored zo-
nate pileus with its short coarse pile-like tomentum, the madder-
brown color of the mature gills, and the abundant white milk,
which is very acrid after being on the tongue a moment. The
margin of the pileus is usually marked with darker furrows, possi-
bly from the pressure of the inrolled margin upon the gills. The
Zones are sometimes obscure, but are more often conspicuous. In
the dried plant the tomentum is inconspicuous.

A few specimens of this plant were found August 30, 1904, near
Smithtown, on Long Island, by Professor Peck, who from the first
was inclined to regard it as an undescribed species. In the follow-
ing summer one specimen was collected by Benedict on Staten
Island. But in 1907 in the “ Pink Beds,” North Carolina, this
was the most abundant mushroom throughout July and August.
It is frequently gregarious and sometimes cespitose in habit. 1
have seen as many as fourteen specimens growing within a few
square feet. What has proved by comparison to be the same
species was collected in Alabama in November, 1896, by C. F.
Baker, but as it was accompanied by no field notes it was then im-
possible to classify it.

I take pleasure in naming this species Peckii in recognition of
Professor Peck's extensive work upon the genus Lacfaria.

The type specimens are in the herbarium of the N. Y. Bota-
nical Garden.

LACTARIAE OF THE UNITED STATES 79

46. Lacraria ALPINA Peck, Ann. Кер. N. Y. St. Mus. 27: 96.
1875. [As Lactarius.] Hennings, in Eng. & Prantl,
Nat. Pflanzenfam. 1'** : 216. 1808

Pileus fleshy, thin, convex, then plane or depressed in the cen-
ter, sometimes papillate, fulvous to fulvous-isabelline, azonate, dry,
minutely squamulose, especially toward the center, 1.5-4 cm. broad,
margin involute, then merely arched, even; gills pallid, then yel-
lowish and pruinose, seldom forking, close, thin, decurrent, narrow ;
stem of the same color as the pileus, or slightly paler, equal, dry,
glabrous, stuffed, becoming hollow, 2-5 cm. long, 3-5 mm. thick ;
spores white, globose, echinulate, 6.5-8 и; latex white, unchang-
ing, acrid.

Has.: Borders of woods. July, August, and September.

DISTRIB. : New York, Peck, Earle 864; Vermont, Burling-
ham тоб, 1906; Maryland, Banning; District of Columbia, Mur-
rill 1491 ; Virginia, 914 meters, Murrill 17; Alabama, Earle.

Плозт.: Banning, Folio Md. Fungi (at N. У. St. Museum,
Albany), A. 79.

DISTINGUISHING FIELD-MARKS : The small size, the tawny-ochra-
ceous color, the squamulose pileus, and acrid latex. It differs
from Lactaria helva Fr. т its small size, in the pileus being merely
squamulose instead of floccose-squamulose, and in the milk never
being watery. It is of about the same size and color as Lactaria
subdulcis (Pers.) Fr.

47. LACTARIA RUFULA Peck, Bull. Torrey Club 34: 346. 1907.
[As Lactarius]

Pileus fleshy, firm, broadly convex, becoming somewhat in-
fundibuliform, without an umbo, brownish-red, azonate, dry, appar-
ently glabrous, 5-10 cm. broad; gills pinkish-yellow, becoming
darker with age and pruinose, close, adnate; stem colored like the
pileus but paler, equally or slightly tapering upwards, sometimes
with root-like extension, often with yellowish-brown strigose hairs
at the base, stuffed, 4-8 cm. long, 5-6 mm. thick; spores creamy-
white, globose, verruculose, 8-10 м; latex white, acrid.

Has.: Rich soil and leaf-mold under trees. March.

Distris.: California, Patterson & Nohara.

This species seems to be separated from L. rufa (Scop.) Fr.
chiefly in the absence of an umbo, and in a cespitose habit of
growth. I have not seen L. rufa living, but it seems to me that

80 LACTARIAE ОЕ THE UNITED STATES

L. rufula is so closely related to it as possibly to prove to be
only a form. The type specimens аге in the herbarium of the №.
Y. State Museum at Albany, and the above description has been
arranged from them and the original description.

XIV. GRISEAE

Plants rather small, varying in color from ash-gray to nearly
black ; pileus dry and covered with a short tomentum or merely
squamulose ; latex white, unchanging, and slowly acrid.

Synopsis of species
Pileus 2.5-5 cm. broad in mature plant, ash-gray to brownish-gray.
Azonate, stem glabrous,
Minutely squamulose or unpolished, aromatic. бо. L. glyctosma.
Minutely tomentose, no odor. 48. Г. grisea.
Zonate, stem white-tomentulose. . L. Hibbardae.
Pileus not over 15 mm. broad in mature plant, blue-black, then
zoned with slate-gray or finally with snuff-brown, scabrous-

pubescent.

CA
м со

49. L. Bensleyae.

FIGURE 14. Lactaria grisea Peck. No. 85, 1907, Burlingham.

48. Lacraria GRISEA Peck, Ann. Rep. N. Y. St. Mus. 23: 119
1873 (separate, 1872). [As Lactarius.] — Hennings,
in Eng. & Prantl, Nat. Pflanzenfam.
E406 T1898
Pileus fleshy, rather thin, firm at first, then lax, broadly convex
papillate, then depressed in the center, or at length infundibuliform,
with or without papilla, varying from slate-gray (362) to smoke-

LACTARIAE ОЕ THE UNITED STATES 81

gray (363), becoming yellowish with аре (putty-colored, 311),
azqnate, dry, minutely tomentose, becoming floccose-tomentose,
sometimes appearing squamulose to the naked eye, 1-5 cm. broad,
margin involute, then spreading, entire; gills white, becoming
cream-colored to honey-yellow, and pruinose, seldom forking,
close, adnate to slightly decurrent, broader than the thickness of
the pileus; stem of the same color as the pileus or paler, nearly
equal, dry, glabrous except at the base, which is sometimes pu-
bescent, stuffed, then hollow, 1.5-6 cm. long, 3-6 mm. thick ;
flesh white, unchanging, not aromatic; spores white, broadly
elliptical, echinulate, 6-7 их 8-9.5 и; latex white, unchanging,
slowly acrid. (FIGURE 14.)

Has. : In moist, mossy places in either coniferous or deciduous
woods, on the ground or on decaying logs. July, August, and
September.

DISTRIB.: New York, Peck, Earle, Benedict, Burlingham ;
Vermont, Jones, Burt, Burlingham 2, 1906 ; Maine, White ; Con-
necticut, Underwood & Earle, Benedict; North Carolina, 1000
meters, Burlingham 85, 1907.

DISTINGUISHING FIELD-MARKS: The gray, tomentose, azonate,
expallent pileus, the glabrous stem, and the lack of odor. While
the plants may be dark-gray at first, they usually become dull-
yellowish or putty-colored when mature. This species is closely
related to L. mammosa Fr., a European plant which has not been
found in the United States. Аз figured by Fries, Г. mammosa is
a larger, stouter plant than Г. grisea, it does not become yel-
lowish with age, and it has a white pubescence on the margin of
the young pileus, and the stem is pubescent. L. grisea is at first
uniformly gray and covered with gray tomentum, which later
becomes floccose and less evident.

49. Lacraria BENsLEYAE Burl. Bull. Torrey Club 34: 87. 1907.

[As Lactarius]
Pileus fleshy, firm, nearly flat with margin inrolled, papillate,
when older depressed in the center but the margin recurved,

(362), and finally with snuff-brown (303) toward the margin, the
center remaining nearly black, dry, surface covered with a dense,
minute, short, rather stiff pubescence, 2-15 mm. broad; gills

whitish, some forking near the stem, close, slightly decurrent, I
mm. wide or equal to the thickness of the pileus, stem somewhat

89 LACTARIAE ОЕ THE UNITED STATES

buff or putty-colored (377), covered more or less witha gray pruin-
osity, nearly equal, dry, glabrous, becoming hollow, 1 cm. or less
in length, 2-3 mm. thick; flesh of the pileus gray, of the stem
buff; spores slightly cream-colored, mostly globose, some slightly
elliptical, echinulate, 5-6.5 и, or rarely 6.5-8 и; latex white, un-
changing, acrid.
Has.: Woods, in black moist soil, under yellow birch and
spruce trees. July to September.
ANGE: Vermont, 500 meters, Bensley, Burlingham.
DISTINGUISHING FIELD-MARKS : The blue-black color of young
or wet plants, the minute size, the hirsute-downy covering of the
pileus, and the distinct zonation in the mature pileus. The species
is gregarious in habit and is sometimes cespitose. As тапу as |
35 were found in the type collection growing in an area of less |
than one square foot. Asa rule, the stem is so short that the |
pileus rests upon the ground, and the plants can be detected only |
by careful search. Type specimens of this species are in the her- |
barium of the N. Y. Botanical Garden and іп the herbarium of the |
2
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N. Y. State Museum at Albany. They аге xo. 38, 1906.

50. ГАСТАВТА Gtyciosma (Er) Fr. Epicr. 348. 1838. [As
Lactarius.|— Schrót. in Cohn, Krypt.-Fl. Schles. 3:
538. 1889
Agaricus glyciosmus Fr. Obs. Мус. 2: 1. 1818.

Pileus fleshy, thin, convex, then expanding, umbonate, at length
depressed, often without umbo, varying from dark-gray (367) to
gray (359) and snuff-brown (303), azonate, dry, “slightly innately- | |
squamulose, or unpolis ed," 2-5 cm. broad, margin involute, then |
spreading and striate ; gills straw-colored, becoming ochraceous, .
close, slightly decurrent, 2 mm. broad; stem of the same color |
as Ше pileus or paler and more yellowish, nearly equal, dry, |
glabrous or minutely downy, stuffed, becoming hollow, 2-5 cm. 2%
long, 4-8 mm. thick; flesh white, odor strong, aromatic; spores -
white, globose to broadly elliptical, echinulate, 6-7 их 6-8 и;
latex white, unchanging, slowly acrid. E 1

Has.: In woods, on the ground or on decaying wood. August | |
September, and October. - |

DISTRIB.: New York, Peck ; Vermont, Jones; Missouri, Glat- |
felter 1203.

Плозт.: Britz. Lact, f. 29 (poor); Cooke, Br. Fungi, p4 2011;

LACTARIAE OF THE UNITED STATES 83

Fries, Icon. A. 170. f. 3 (good); Krombh. Abbild. ai 39. f. 16-18
(poor, doubtful).

Exsic.: Karsten, Fungi Fennici 307; НегреП, Sammlung
praparirter Hutpilze 47.

DISTINGUISHING FIELD-MARKS: Although this is much the
same color as L. grisea, it can readily be distinguished from it by
the absence of tomentum, by the striate margin, and the pungent,
aromatic odor, which persists for some time. Тһе pileus is only
faintly squamulose, and sometimes seems to be merely roughened
and unpolished. It is never floccose-squamulose like the pileus
in mature forms of L. grisea.

According to Fries, the color is sometimes brick-red and again
has a violet tinge, but this has not been observed in the American
forms.

51. LacraRiA HiBBARDAE Peck, Jour. Myc. 14: 2. 1908. [As
tarius]

colored, some forking, close, adnate, thin, narrow; stem pinkish-
white, equal or slightly tapering upwards, glabrous below, whitish-
tomentose at the top, stuffed, 2.5-4 cm. long, 3-6 mm. thick ; flesh
whitish, odor weak or none; spores globose, 6-8 и; latex white,
unchanging, acrid. (Arranged from Peck’s description.)

Has.: On the ground under pine trees. October.

DisrRiB.: Massachusetts, Hibbard.

DISTINGUISHING FIELD-MARKS: This species may be distin-
guished from ZL. g/yciosma, which it most closely resembles, by the
tomentose rather than squamulose pileus, the faint zonation, the
absence of the odor peculiar to that plant, and by the tomentum
on the stem.

The type specimens are in the herbarium of the N. Y. State
Museum at Albany.

ХУ. Е АЕ

Pileus dry, pruinose to velvety, dark-brown, smoky-brown, ог
putty-colored to paler, flesh rather firm, plants of medium size ; gills
becoming darker with age, and pruinose ; latex mild or acrid,

84 LACTARIAE OF THE UNITED STATES

salmon-colored or white, either unchanging or changing to salmon- 4
pink, at least where іп contact with the broken flesh, or in one |
species becoming yellow. 5

The species іп this group show ав close a relationship as do
those in the Ласінае. Three of the species seem to be limited
to America.

Synopsis of species

Latex or wounds becoming salmon-pink or reddish.

Pileus azonate, unspotted, no odor.

i d

Pileus dark-brown, velvety. 54. L. ligniota.
Pileus snuff-brown or paler, pruinose. 52. L. plinthogala,
Pileus spotted, odor bad, L. аты
Latex and wounds not changing color,
Spores yellow. 53. ZL. Sumstinei.
white. su. Г. Gerardi.
Latex becoming yellowish. St
Pileus subtomentose. 56. L. subtomentosa.
Latex salmon-colored from the first, pileus whitish. 57. L. salmonea.

52. Lactaria plinthogala (Otto) :
Agaricus azonites Bull. Hist. Champ. 2: 497. 1809; Herb. Fr. |
pl. 567. f. 3. 1791. Probably not Lactarius azonites Gillet |
Agaricus plinthogalus Otto, Versuch Agar. 75. 1816. 1
Agaricus fuliginosus Ет. Syst. Myc. 1: 73. 1821.
Lactarius fuliginosus Fr. Epicr. 348. 1838.
Lactarius fumosus Peck, Ann. Кер. N. У. St. Mus. 24: 74

1872.

Lactariella azonites (Bull.) Schrót. in Cohn, Krypt.-Fl. Schles. 3:
544. 1889.
Pileus fleshy, convex then plane, sometimes with a small umbo,

depressed іп the center, then subinfundibuliform, raw-umber to

dingy yellow-brown ( 325, café au lait), snuff-brown (303. Ё 4) or

sometimes connected by vein-like reticulations, subdistant,
or slightly decurrent, about 5 mm. broad; stem of the same 00,
as the pileus, often whitish toward the base, nearly equal or taper
ing downwards, glabrous, pruinose, stuffed but firm, then hollow,

р ge pean species, the occurrence of which іп the United States is doubtfi

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LACTARIAE ОЕ THE UNITED STATES 85

3-7 cm. long, 6-12 mm. thick; flesh white, changing to reddish
or salmon where exposed to the air; spores yellow, mostly globu-
lar, echinulate, uniguttulate, 6.5-10/4; latex white, rarely chang-
ing color except where in contact with the flesh, where it becomes
salmon-pink, mild, then acrid. ;

Has.: In deciduous or mixed woods. July to September.

Distris.: New York, Peck, Earle, Peck & Earle (Long Island) ;
‘Maine, White; Vermont, Burlingham 41, 1906; Pennsylvania,
Murrill, Herbst ; North Carolina, Burlingham 80, 1907. It has
also been reported from Alabama by Atkinson.

Плозт.: Atkinson, Stud. Am. Fungi, f. 117; Bern. Champ.
Roch. pi. 38. f. 3; Britz. Lact. f. 33", 40; Bull. Herb. Fr. ai,
567. f. 3; Cooke, Br. Fungi, ai 996; Gillet, Champ. Fr. ai.
165 [384]; Harzer, Pilze, p/. 19; Krombh. Abbild. a 14. f.
10-12 (f. 12 very poor); Noulet & Dass. Champ. a 18. f. В.;
Pat. Tab. Analyt. Fung. pl. 322.

Exsic.: Herpell, Sammlung práparirter Hutpilze 73.

DISTINGUISHING FIELD-MARKS: For the most part, the pileus
is snuff-brown or coffee-and-milk-colored, and is dry, glabrous or
pruinose, and very smooth ; the gills become yellow and the spores
are yellow, while wounds of the plant become salmon-pink or red-
dish. Тһе species varies greatly in size, color, closeness of the
gills, and in the readiness with which the latex or wounds change
color.

In the * Pink Beds," North Carolina, I found small forms
growing in moss and shade, having the pileus nearly white, or
stone-colored in the center, and from 3-4 cm. broad, and the gills
white, close, and about 2 mm. broad, while the latex as well as
the wounds became salmon-pink, but since intergrading forms
occur it does not seem possible to separate it from the species.
At the other extreme, in sandy soil, large normally colored forms
occurred, having distant gills up to 10 mm. broad, the wounds
changing slowly but the latex unchanging except where in contact
with the flesh. Аза rule, the paler the plant, the more quickly the
wounds become salmon and the more likely the milk is to change
color. The small white form described above, ло. 60, 1907, and
the distant-gilled form, по. 69, 1907, аге in the herbarium of the
N. Y. Botanical Garden.

86 LACTARIAE ОЕ THE UNITED STATES

Gillet considered Agaricus azonites Bull. pl. 559 to be the type 7
of a distinct species from Lactaria fuliginosa Fr. and accordingly
described a Lactarius azonites based on that plate as the type.

53. Lacraria Sumstiner Peck, Bull. Torrey Club 32: 78. |
1905. [As Lactarius] 1

Pileus fleshy, rather thin, firm, convex, soon depressed іп the cen- |

ter, grayish to putty-colored ( 37 7), azonate, dry, glabrous, smooth,
or with wrinkles radiating from the center, 2.5-7.5 cm. broad, margin |
involute, then arched or spreading ; gills colored like the pileus,
distant, thin, decurrent, 1 cm. or more broad; stem colored like
the pileus, nearly equal, rather firm, dry, glabrous, 2.5-5 cm. long, |
6-12 mm. thick ; flesh whitish, not discolored with the milk; spores |
yellow, globular, echinulate, 7.5-10 м; latex white, unchanging, С
and not staining the flesh or gills, acrid. 4
Нав.: “ Grassy places in open woods.” Іп deciduous woods. |
July and August. 2
Пізтків.: Pennsylvania, Swmstine (type); District of Columbia, |
Murrill 1476 and 1523; Virginia, Murrill 1682; Missouri, Glat- |
ЕЙ ғ 1077.
DISTINGUISHING FIELD-MARKS: Тһе grayish or yellowish-gray

pinkish or salmon as іп Z. plinthori/a. It differs from Z. Gerardi
in its paler color, its yellow spores, in the gills being of the same
color as the pileus, and in the acrid taste of the latex.

The type specimens are in the herbarium of the Carnegie. 2
Museum at Pittsburg, Pa., and in the herbarium of the N. Y. St.
Museum at Albany.

54. LACTARIA LIGNIOTA | Monogr. 2: 177. 1863. [As
La t rius]

Lactarius fuliginosus majo | picr. 348. 1838.

Lactariella ligniota Schrii à Cohn, Krypt.-Fl. Schles. 3: 544
1880. )
Pileus fleshy, convex, | plane to slightly depressed, umbo-

nate, often wrinkled in the © =v, dark-brown, azonate, dry, РИ

nose-velvety, 3-8 cm. bro. от sometimes plicate ; gills WA?

to ochraceous, becominy or salmon where wounded, p

crowded, of various lengths, с urrent, about 5 mm. broad ; 5

Сори ли dE ТЕЧЫ

И ng ааа ыы ыма асы гр азасы

LACTARIAE ОҒ THE UNITED STATES 87

of the same color as the pileus, equal or abruptly smaller and pli-
cate at the apex, smooth, pruinose-velvety, stuffed, up to 8 cm.
long and 12 mm. thick ; flesh white, becoming pinkish or salmon
where wounded ; spores yellowish, globular, echinulate, 9-10»;
latex white, changing to salmon-pink where in contact with the
broken flesh, mild or finally slightly acrid.

Has.: On the ground in mossy wet woods, especially fir.
July to September.

Distris.: New York, Peck; Vermont, Burlingham 43, 1906,
Jones ; Connecticut, Hanmer 1945 ; New Jersey, Sterling; North
Carolina, Burlingham 20, 1907.

Плозт.: Atkinson, Stud. Am. Fungi, /. 776 ; Britz. Lact. f.
4; Fr. Icon. M. 171. f. І

DISTINGUISHING FIELD-MARKS: This species differs from /ас-
taria fuliginosa in its darker seal-brown color, the velvety cover-
ing of the pileus and stem, the more rugose surface of the pileus,
and in the larger average size of the plant.

Peck has described one “ variety” under this species, Z. /igniota
tenuipes, which differs in the small size and the slender stem. This
form has been found also in Connecticut by Hanmer 1479, and in Ver-
mont I found one specimen, 4 3, 3995. gro us = - nep
in spruce woods. Theform

Pileus up to 3 cm. broad, stem 2. 5-8 cm. long, 4 mm. ; thick,

55. Г.АСТАНА GERARDII Peck, Ann. Rep. N. У. St. Миз.
26: 65. 1874.* [As Lactarius.] — Hennings, in Eng.
& Prantl, Nat. Pflanzenfam. 1'**: 216. 1898
Pileus fleshy, firm, convex at first, often with a small umbo,
then plane or depressed, dark seal-brown, becoming golden-brown
or umber, or sometimes paler, azonate, dry, surface velvety,

edis or ventricose, stuffed, then hollow, 2. 5-5 cm cm. long, 4-20
thick ; flesh white, unchanging ; spores white, globular, pda
6. 65-91; latex white, unchanging, mild, then slightly acrid. Edible.

viria published in advance in April 1874. |

88 LACTARIAE ОЕ THE UNITED STATES

Has.: On the ground in woods or in open groves. July to
September.

Півтків.: New York, Gerard, Peck, Earle, Burlingham, Peck
& Earle (Long Island); Vermont, боо meters, Burlingham 28, |
1906; Pennsylvania, Murrill; District of Columbia, Murrill;
North Carolina, 1000 meters, Burlingham 23, 1907.

ILLUSTR.: Ann. Rep. М. У. St. Mus. 26: pl. 50. f. 12-16;
Memoir М. У. St. Mus. 3: ai 53. f. 12-16.

DISTINGUISHING FIELD-MARKS: This species is closely related
to Lactaria ligniota Fr., but can be distinguished from it by the
white spores, the unchanging color of the broken flesh or gills,
and the more distant gills.

The type specimens were collected near Poughkeepsie, N. Y.,
by W. R. Gerard, in whose honor the mushroom is named. They
are preserved in the herbarium of the N. Y. State Museum at
Albany.

56. Т.АСТАВТА SUBTOMENTOSA Berk. & Rav. Апп. and Mag.
Nat. Hist. ПІ. 4: 11. 1859. [As Lactarius.]— Hennings,
in Eng. & Prantl, Nat. Pflanzenfam. 1: 216. 1898.
Pileus convex, firm, umber-brown, dry, subtomentose, 5-7.5 2
cm. broad; gills white, distant, decurrent, broad; stem of the |
same color as the pileus except at the white base, hollow, 2.5 cm. |
long, 12 mm. thick ; latex white, becoming yellowish, acrid. a
Нав.: On the ground in swamps. September. 7)
DISTRIB. : South Carolina, Ravenel. 2
DISTINGUISHING FIELD-MARKS: Тһе dry umber-brown, sub- ; 3
tomentose pileus, and the acrid white latex, which becomes yellow |
on exposure to the air. %
I have not seen the type specimens of this species, which аге
іп the herbarium of the Royal Botanic Gardens at Kew. But the
species seems to be well defined and distinct from any other of our
species, and it is probable that further collections in South Carolina -
may discover more of these plants.

57. LACTARIA SALMONEA Peck, Bull. Torrey Club 25: 369.
1898. [As Lactarius] Y

Pileus fleshy, rather firm, thin, convex, soon depressed in the
center, otherwise nearly plane, white, becoming reddish where

LACTARIAE OF THE UNITED STATES 89

bruised, dry, somewhat velvety, 2.5-3.8 cm. broad, margin invo-
lute, then spreading, even ; gills pallid, becoming darker, brownish
in drying, close, adnate to decurrent; stem white, salmon-colored
within, nearly equal, occasionally eccentric, velvety, solid, about 2.5
cm. long, 3-6 mm. thick ; flesh becoming salmon where wounded ;
spores subglobose, echinulate, 7.5-0/4; latex salmon-colored.

Has.: In wet swampy places, usually on naked ground that
has been overflowed. August to October.

DISTRIB. : Alabama, Ват (type) ; Mississippi, Zarte.

DISTINGUISHING FIELD-MARKS: The white color of the plant,
the velvety covering, the short stem, and the salmon-colored latex.

This isthe only known species having from the first salmon-col-
ored milk. I have not seen this plant living and I am in doubt as
to its relationship. The dry velvety pileus and the salmon color
assumed by wounds would indicate a relationship with the P/intho-
galae, while the colored milk would tend to place it with the
Dapetes.

DOUBTFUL SPECIES

LACTARIA ACRIS (Bolt.) Fr. Ерісг. 342. 1838. [As Lactarius.]
— Hennings, in EN & Prantl, Nat. Pflanzenfam.
**: 216. 1898

This European species has been reported from North Carolina
by Schweinitz in Syn. Fung. Carol. Super. 60. 1818, but the
specimens have been destroyed by insects, and, since this is the
only station for which it has been reported, I hesitate to include it
in the list of species positively known in the United States. Из
relationship seems to be with Lactaria plinthogala. Thedistinguish-
ing marks are the ill odor, the spotted pileus, and the more red-
dish color assumed by the acrid latex. Itis well figured in Cooke,
Br. Fungi, p/. 1005.

XVI. LacrIFLUAE

Pileus dry, glabrous or velvety, some shade of fulvous, flesh
rather thick Set firm ; gills becoming darker with age and pruinose ;
latex abundant, sweetish or at least mild, unchanging, but the gills
and flesh sometimes becoming brown where injured.

The members of this group seem to be very closely related,
the species, in one instance, differing chiefly in the reaction of the
wounds upon exposure to the air. With the exception of the first
two, the species are all confined to America. T

90 LACTARIAE OF THE UNITED STATES

Synopsis of species

Pileus entirely glabrous.
Azonate, 58. Г. асийиа.
Zonate. бо. LZ. ichorata.

Pileus pruinose-velvety.

ills not staining brown where injured,
Distant. 59. Г. Aygrophoroides,
Close. 63. L. subvelutina.

Gills staining brownish where injured.
Pileus yellowish-buff, smooth or very slightly rugose. 62. Z. /uteola,
Pileus Vandyke-brown to dead-leaf, usually corru-

gated. бі. Г. corrugts.

58. Lactaria lactiflua (L.)

Agaricus lactifiuus L. Sp. Pl. 1172. 1753.
Agaricus oedematopus boob. Fl Carn, 2: 443. 1772. e
Agaricus testaceus Alb. & Schw. Consp. Fung. 200. 1805.
Agaricus volemus Fr. Syst. Myc. 1: 69. 1821.
Lactarius volemus Ет. Epicr. 344. 1838.
Pileus fleshy, firm, convex, then nearly plane or slightly `
depressed, fulvous (308), buff (309), brownish-terra-cotta (334) 9
brownish-orange, sometimes much paler, azonate, dry, glabrous,
smooth or at length rimose-rivulose, 5-13 cm. broad when
mature ; margin even, involute at first, then extended ; gills creamy- `
white, or tinged with the same color as the pileus, becoming |
darker with age, changing brownish where injured, often forking 2
or 3 mm. from the stem or midway to the margin, close, adnate,
2-5 mm. broad; stem of nearly the same color as the pileus кє S
paler, nearly equal, glabrous, pruinose, solid or sometimes becom-
ing hollow, 2-10 cm. long, 1-2 cm. thick; flesh whitish, chang-
ing brown where exposed to the air, odor strong, persisting;
spores white, globular, echinulate, 7-го и, cystidia 20—35 # long, |
colorless or yellowish; latex white, unchanging, mild, sticky, |
abundant. Edible, :

Нав. : In woods ог groves, especially іп the vicinity of oaks
July to October. 3

Пізтків.: New York, Peck, Earle, Burlingham 23, 19046
Maine, White, Murrill ; Vermont, Jones, Burlingham ; Mass
chusetts, Morris ; Connecticut, Earle, Underwood, Hanmer; Ме
Jersey, Ев; Pennsylvania, Herbst ; Maryland, Banning; D
trict of Columbia, Murril/ ; North Carolina, Atkinson, Burlingha®
14, 1907, 1,000-1,400 meters elevation ; Virginia, Murrill ; A

* See also ** Doubtful Species.”

LACTARIAE OF THE UNITED STATES 91

Бата, Baker, Earle, Underwood; Mississippi, Earle; Ohio,
Beardslee ; Missouri, Glatfelter 1078 ; Indiana, С. W. Wilson.
Плозт.: Вапа, Champ. Nice, ai 20. f. 1-3; Bel. Champ.
Tarn. oi 23; Boyer, Champ. A. 37; Bres. Fung. Mang. ai 66;
Britz. Lact. f. 6; Cooke, Br. Fungi, ai 000; Cordier, Champ. Fr.
pl. 26. f. 2; Ellrodt, M 6. f. г, 2, 3; Fr. Sverig. Svamp. MX.
го; Gillet, Champ. Fr. ai 170 [402] (form); Hahn, Der Pilz-
Sammler, ed. 2. pl. 4. f. 14; Hussey, Illust. Br. Myc. т: M. 87;
Krombh. Abbild. ai 39. /. 1-4 (coloring poor); Lucand, Champ.
Fr. pl. 145 ; Peck, Ann. Rep. М. Y. St. Mus. 48 : р/.ҙо; Pat. Tab.
Analyt. Fung. oi 323; Rolland, Bull. Soc. Myc. Fr.'7: pl. 2.7. 7;
Roumeg. Crypt. Illust. oi 742 ; Rich. & Roze, Atl. Champ. ai 38.
f. 6-12 (coloring poor); Schaeff. Fung. Bav. Icon. AM. 5; Sicard,
Hist. Nat. Champ. p/. 44. f. 236 (poor); Venturi, Studi Micol. ai.
6. f. 42-48 ; White, Conn. Geol. and Nat. Hist. Surv. Bull. 3:
pl. то

Exsic.: Herpell, Sammlung praparirter Hutpilze 77; Rou-
meguére, Fungi Selecti Exsic. 5223 ; Sydow, Mycotheca March-
ica 608 ; Thümen, Fungi Austriaci 872.

DISTINGUISHING FIELD-MARKS : The rather large size and thick
flesh of the plant, the glabrous, smooth, or sometimes rugose or
cracked surface of the pileus, the fulvous to golden-fulvous color
of both pileus and stem, the whitish gills which become brown
where injured and in drying, and the abundant, mild, sticky latex.
The latex drops from even slight wounds in the gills or flesh.

There is a great variation in color in different specimens of L.
lactiflua, but the other characteristics remain so constant that it
seems scarcely possible to separate valid varieties upon the color
basis. Two collections from Virginia (407 and 266 Murrill) are
described as pale cream-color, while in North Carolina I found the
color varying from pale-buff to a dark chestnut-red (Indian chest-
nut-red, 333). This difference in color does not seem to be due
to a shady or a sunny habitat. The rugose and the cracked
character of the pileus, however, is probably due to weather con-
ditions or to moisture content. In wet weather the pileus is more
often rugose, while in dry weather the surface is liable to become
cracked in areas. In North Carolina, while rain fell nearly every
day the rugose condition prevailed, the plants with the cracked

92 LACTARIAE OF THE UNITED STATES

pileus being found only later in the season when the weather was
drier. Consequently, the forms having a rugose pileus I would |
include as merely a normal variation of the species.
Linnaeus describes Agaricus lactifluus “ pileo plano carneo lac-
tescente, lamellis ruffis, stipite longo сагпео” This is rather brief
to indicate plainly the type that he was describing, but he cites
Haller’s “ Fungus flavo-ruffus in medio depressus, lacte non аст
manans” as synonymous with Agaricus lactifiuus. Haller
description applies without doubt to Z. volema Fr. He says “ not
rare in forests, easily recognized from its smoothness, firm flesh,
sweet milk such as is rare in fungi. In shape the pileus is ele-
gantly depressed in the center, with the margin roundly elevated.
The color is everywhere rufus-flavous, butyraceous, lamellae paler.
Not decaying, but becoming coriaceous. It is edible, with good -
taste. Ours is occasionally 4 in. in diameter.” (Haller, Helv.
50. 1742.) Lamarck in Fl. Ег.т: 106. 1748, describes Agaricus |
lactiffuus of Linnaeus іп terms which show his plant to have been |
L. volema Fr., and he cites Schaeff. ai 5. Тһе gills of Г. volema
Fr. become rufous with age or in drying so that there is nothing in `
Linnaeus’ description which disagrees with this species ; and from |
his citation there seems to be little doubt that his plant was iden- `
tical with 2. volema Fries.
Agaricus oedematopus Scop. does not seem to differ from А. -
volema Fr. and the description by Scopoli agrees more exactly |
with Fries’ description of Z. volema than it does with the
“variety ” oedematopus. And from Scopoli's description I have |
referred Agaricus oedematopus to L. laċtiflua (L.), or L. volema Е.

59. LACTARIA HYGROPHOROIDES Berk. & Curt. Annals and Mag: |
Nat. Hist. III. 4: 10. 18 59. [As Lactarius.| — Hennings,
in Eng. & Prantl, Nat. Pflanzenfam. 1'**: 214. 1898

Lactarius distans Peck, Ann. Rep. М. Y. St. Mus. 23: ИГ.
1873. [Separate in April, 1872. |
Pileus fleshy, firm, convex, then plane, depressed in the center |

Sometimes becoming infundibuliform, yellowish-buff (37 ec

(328), or fulvous (308), azonate, dry, very minutely DOT",

velvety, appearing as though pulverulent, sometimes rugose, 5011”

times rimose-areolate, 4-10 cm. broad, margin involute, then SI" `

LACTARIAE OF THE: UNITED STATES 93

ing or uplifted ; gills whitish to cream-colored or yellowish-buff, not
discoloring where injured, not forking, distant, sometimes con-
nected by rugose elevations, adnate to slightly decurrent, about 3
mm. broad; stem of the same color as the pileus, nearly equal,
glabrous or pruinose, stuffed or solid, 2-5 cm. long, 0.5-1.5 cm.
thick; flesh whitish, no odor ; spores white, globose to broadly
elliptical, minutely echinulate, 8-10; latex white, unchanging,
not staining the gills or flesh brownish, mild, not so abundant as
in Г. lactifiua (L.). Edible.

Нав.: Mixed woods. July, August, and September.

Distris.: Maine, Sprague (type collection); New York,
Peck, Underwood, Earle; Massachusetts, Francis ; Connecticut,
Earle 940; Delaware, Jackson; District of Columbia, Murrill
1487 ; Indiana, С. И. Wilson ; Missouri, Glatfe/ter ; Mississippi,
Earle

Itrusr.: Peck, Mem. М. Y. St. Mus. 3: e 53. f. 7-11.
DISTINGUISHING FIELD-MARKS: The yellowish-buff or fulvous
color of the plant, the pruinose-velvety covering of the pileus,
the usually short stem, the distant gills, which do not stain brown
where injured or in drying, and the white, mild latex.
type specimens of Lactaria hygrophoroides are іп the
herbarium of Harvard University and also of the Royal Botanic
Gardens, Kew, England. They are по. 6194 of Curtis's fungi.
_ Specimens of Lactarius distans Peck from the herbarium of the
N. Y. Botanical Garden, ло. 940, were sent to Kew Gardens for
comparison with the type specimens of L. hygrophoroides and were
pronounced by Mr. Massee to agree with the type.

бо. Т.АСТАВТА ІСНОКАТА (Batsch) Fr. Epicr. 345. 1838. [Аз
Lactarius.] — Hennings, in Eng. & Prantl, Nat. Pflan
zenfam. 1'** : 274. 1898
Agaricus ichoratus Batsch, Elench. Fung. 38. 1783.

Pileus fleshy, rather thin but opaque, rigid, then lax, plane to
depressed in ie center, fulvous, sometimes brown in the center,
zoned with testaceous, dry, glabrous, smooth, 5-8 cm. broad,
margin incurved, then. spreading ; gills white, ‘then ochraceous,

94 LACTARIAE ОЕ THE UNITED STATES

Has.: On ground in woods.

Distris.: Ohio, Loyd. Rare.

Ittust.: Batsch, Elench. Fung. ai тз. f. 60. a, 6; Britz. Lact.
f. 36 (coloring poor); Cooke, Br. Fungi, p/. гооо (good).

DISTINGUISHING FIELD-MARKS: This specimen resembles 2.
volema but differ in being zonate and more slender. Lloyd says
of the specimens which he found: ‘Stem solid but slightly
spongy, eccentric in the few specimens I saw. Pileus brown,
marked with zones. Milk white, acrid, not changing in color.”

These specimens are in the herbarium of the N. Y. State Mu-
seum at Albany. In the dried condition they resemble Z. /acti-
Биа except for the zonation and their smaller size.

бі. LACTARIA CORRUGIS Peck, Ann. Кер. М. У. St. Mus. 32: 31.
1880. [As Lactarius.] Hennings, in Eng. & Prantl,
Nat. Pflanzenfam. 1'**: 214. 1808

Pileus fleshy, firm, thick, convex, then depressed in the center,
color varying from Vandyke-brown (340. t. 4—7) in the center, to
mineral-brown (339. £ г) at the margin, sometimes paler, ap-
proaching more nearly dead-leaf (327. 2. г), azonate, dry, minutely
velvety, and appearing as though covered with a bloom, surface
more or less corrugated, 7-12 cm. broad, margin involute at first,
then arched or spreading ; gills cinnamon (323. 7. 3) when young,
paler when mature or tinted with honey-yellow, becoming fulvous-
brown where injured or when dried, sometimes forking, close,
adnate to slightly decurrent ; stem tinted with dead-leaf ( 327. 1. 7),
paler than the pileus, nearly equal, dry, pruinose in the upper
portion, minutely pubescent at the base, firm, solid, 6-7 cm. long,
2-2.5 cm. thick; flesh white, having only a slight odor; spores
white, globose, echinulate, 9-12 p; latex white, unchanging, mild
or slightly astringent, abundant. Edible,

Has.: Moist woods, especially in mixed oak-chestnut-maple
woods. August and September.

Distris.: New York, Peck; Connecticut, Webster, Under-
wood, Hanmer, 1936; New Jersey, Sterling ; Pennsylvania,
Herbst ; Delaware, Jackson ; North Carolina, Atkinson, Burling-
ham 18, 1907 (1000 meters); Tennessee, Murrill, 506; Alabama,
Earle ; Mississippi, Earle ; Missouri, Glatfelter.

Плозт.: Atkinson, Stud. Am. Fungi, f. 115.

DISTINGUISHING FIELD-MARKS: The rather large size of the ma- |

LACTARIAE ОЕ THE UNITED STATES 95

ture plants, the dark color of the pileus, which inclines more to
Vandyke-brown than to fulvous, the paler stem, the velvety-
pubescent covering of the pileus and the lower portion of the
stem, and the more or less corrugated condition of the pileus.
There is but little or no odor, and the wounds of the gills do not
turn brown so quickly as іп L. Јасна.

The species is closely related to Г. /acziffua. The specific name
refers to the corrugated surface of the pileus, but since L. /асийиа
occurs with a rugose pileus, the specific differences seems to me
to be in the velvety-pubescent covering of L. corrugis, the more
livid color of the plant, the absence of the strong odor which is
characteristic of Г. /actiflua, and the larger spores and cystidia. As
а rule the cystidia of Г. /actifiua are paler.

62. LACTARIA LUTEOLA Peck, Bull. Torrey Club 23: 412. 1896.
[As Lactarius]

Lactarius foetidus Peck, Bull. N. Y. St. Mus. 54: 949. 1902.

Pileus fleshy, firm, broadly convex or nearly plane, somewhat
depressed in the center, whitish to yellowish-buff (370), azonate
or sometimes with a depressed zone near the margin, dry, minutely
pruinose-velvety, sometimes slightly rugose, 5-8 cm. broad, margin
involute at first, then arched or spreading ; gills white, then yel-
lowish, becoming brown where injured, some forking near the stem,
close, adnate, or slightly decurrent, narrow ; stem of the same color
as the pileus, nearly equal, dry, somewhat pruinose-velvety, stuffed,
2-6.5 cm. long, 6-12 mm. thick; flesh whitish, staining brown
where injured; spores white, subglobose, echinulate, 7.5-8 м;
latex white, staining the flesh and gills brown, mild, abundant.

Нав.: Mixed woods, among fallen leaves, in dry or fairly
moist soil. July, August, and September.

DISTRIB. : Massachusetts, Webster (type specimens); New
York, Peck, Earle 840; Connecticut, Hanmer 534; Tennessee,
518 meters, Murrill 960 ; Ohio, Beardslee ; Missouri, Glatfelter ;
Mississippi, Earle.

Плозт.: Bull. М. Y. St. Mus. 67: X. 83. f. 7-11.

DISTINGUISHING FIELD-MARKS: The whitish to yellowish-buff
color of the pileus and stem, the minutely velvety covering, and
the white, close narrow gills which stain brown where wounded.
The latex is very abundant, and according to Peck becomes brown

96 LACTARIAE ОЕ THE UNITED STATES

upon exposure to the air. The color of the plant becomes a deeper
yellow in drying. It may be distinguished from Г. /лс иа by its
smaller size, the velvety, more yellow pileus, and by the different
odor. It resembles Z. hygrophoroides more closely, but can be
separated from that by the close gills, and by the brown color
which the milk and wounds assume.

After further collections of Г. /ueo/a, Professor Peck now con-
siders Z. foetidus Peck to be the same species, having found that
Г. luteola sometimes has a fetid odor; accordingly, L. foetidus is
reduced to а synonym. The type specimens are in the herbarium
of the N. Y. State Museum at Albany.

63. LacTARIA SUBVELUTINA Peck, Bull. М. Y. St. Mus. 75: 18.
1904. (Ав Lactarius]

Pileus fleshy, firm, convex then nearly flat, depressed in the
center, golden-fulvous, dry, minutely velvety, appearing pruinose
to the naked eye, sometimes wrinkled, 2.5—5 cm. broad, margin
inrolled, then spreading, even, sometimes wavy; gills Cream:
colored, becoming darker with age, not staining brown where
injured, a few forking, close, adnate to slightly decurrent, narrow ;
stem of the same color as the pileus or paler, equal, somewhat
velvety, smooth, solid, 1-2 cm. long, 4-8 mm. thick ; flesh white,
not staining brown; spores white, subglobose, nearly smooth
(minutely and sparsely papillate), 6.5-8 и; latex white, unchang-
ing, mild.

Нав.: “ Woods and open places.”

Пізтків.: New York, Peck (type); Ohio, Dawson 41; Mis-
souri, G/affelter 399.

; DISTINGUISHING FIELD-MARKs: The close gills and the unchang-
ing white latex which does not stain the gills brown.

The dried specimens of this plant closely resemble those of
L. luteola Peck, but the spores are smaller and smoother.

DOUBTFUL SPECIES
“ LACTARIUS SACCHARIUM ” Johnson, Bull. Minn. Acad. Sci. 1878:
337. 1878.
The type spcimens of this species I һауе not seen, and, 50 far

as is known, Johnson’s material from which he made his determi- |

nations has not been preserved. Since the type is not available and

LACTARIAE ОЕ THE UNITED STATES 97

since during the thirty years following the description no further
collections have been reported, I hesitate to regard it as a valid
species. But in order that the species may be established if it be
valid, I append the following, arranged from Johnson's description ;

ileus convex, papillate, at length expanded, depressed, then
infundibuliform, brownish, or slate, pruinose, then reddish-orange,
1.5-4 cm. broad; gills pale-orange, becoming brownish when old,
dimidiate or entire, sinuate, crowded, decurrent, narrow ; stem
orange, irregular, compressed, curved or straight, 1.5 cm. high,
2-6 mm. thick ; latex white, slightly sweet. Тһе plants are gre-
garious, or cespitose. Тһе pileus and the upper part of the stem
are milky, but the gills are not. It was found growing among
moss and grass, beneath poplar trees. July and August.

LacrARIA CarckEoLus Berk. Lond. Jour. Bot. 6: 315,316. 1847.
[As Lactarius | 297%)
Pileus fleshy, thin, convex, with depressed center, brown-buff,
dry, smooth, 7.5 cm. broad, margin repand, epidermis rimose ; gills
white, more or less connected by transverse veins, forked near the
edge, very distant, decurrent, up to 12 mm. broad; stem of the same
color as the pileus, 12 mm. thick and long; flesh white; latex
white, mild.
Нав. and Loc. : On the ground in woods, Waynesville, Ohio,
August 31 and September го, 1844, T. С. Lea. -
_ Berkeley says : “An extremely curious species, remarkable for
its few distant gills and the contrast between them and the brown-
buff stem. The pilei in all the specimens found at present аге
laterally confluent. It cannot.be confounded with any known
species.”’ с |
This species has not been reported since they were collected by
Mr. Lea, and the confluent рйеі would indicate them to be an ab-
normal growth, very possibly of Z. hygrophoroides, described by
Berkeley & Curtis in 1859 from Sprague material.

XVII. CAMPHORATAE

; Pileus dry, glabrous, usually smooth and polished, but some-
times becoming areolate or minutely roughened with pits, reddish-
brown, fulvous, or tawny, rarely grayish, plants not large, pileus
thin, flesh lax ; gills becoming darker with age and pruinose ; latex

mild or tardily acrid, white, or thin and watery, usually unchanging.

98 LACTARIAE OF THE UNITED STATES

Synopsis of species
Latex white, unchangi ng.
Odor aromatic in drying, pileus red-brown, 64. Г. camphorata. |
Odor none.
Pileus azonate.
ills not changing color when wounded.
Fulvous to isabelline, sometimes bay-
red. 66. Г. subdulcis.
Golden-fulvous, stem orange L. mitissima.T
Gills becoming greenish where wounded.
ileus about 3 cm. broad, umber-colored. 70. Z. parva.
Pileus up to 7.5 cm. broad, grayish,
а

often roughened. . varia.
Pileus zonate when moist, rather large. 69. Z. mutabilis.
Latex white, becoming yellowish or staining gills yellow,
uff.

pileus yellowish- 68. Z. isabellina.

Latex watery or whey-colored.
Pileus brownish-terra-cotta, fading, areolate. 65. Z. rimosella.
Pileus not becoming areolate
Margin even. 67. L. serifiua.
Margin crenate to sulcate, plant small, 64. L. camphorata fragilis

64. LACTARIA CAMPHORATA (Bull) Fr. Epicr. 346. 1838. [Аз
Lactarius.] — Schrot. in Cohn, Krypt.-Fl. Schles.
3: 535. 1880.
Agaricus camphoratus Bull. Herb. Fr. ФА 507. J. г. арал Hist.

Champ. 493. 1800.

Pileus fleshy, firm, rather thin, convex, often umbonate, at
length expanded, depressed in the center, but the margin still
arching, fulvous (308) to madder-brown (334), azonate, ФУ,
glabrous, 1-4 cm. broad, margin inrolled and pruinose at first, not `
striate ; gills whitish or flesh-colored (67), becoming reddish-brown, |
sometimes a few forking next the stem, close, adnate to slightly |
decurrent, rather narrow; stem of the same color as Ше pileus
or paler, nearly equal, sometimes flexuous, glabrous, pruinose
smooth, firm to spongy, 1-3 cm. long, 3-8 mm. thick; flesh of
about the color of the gills, unchanging, odor aromatic, becoming
more pronounced in drying; spores white, globose, echinulate,
6-7; latex white, unchanging, mild, abundant. 4.

Нав.: In woods, more abundant in moist mixed woods. Му
to October.

Пізтків.: New York, Pick, Earle, Burlingham ; Massachu-
setts, Davis, Morris ; Connecticut, Earle ; Vermont, Витйи ат.
10, 1906, Burt; Pennsylvania, G/atfe/ter ; Virginia, Af

* See also ** Doubtful Species.”

f European species, the occurrence of which in the United States is doubtful.

LACTARIAE ОЕ THE UNITED STATES 99

North Carolina, Burlingham 3, 1907; Tennessee, Murrill; Ala-
Бата, Earle.

Плозт.: Вама, Champ. Nice, 2/20. f. 11-13 (coloring poor);
Bern. Champ. Roch. ad. 38. f. 2; Bull. Herb. Fr. ai, CSI NS RE ge
Cooke, Br. Fungi, p/. 1013. f. а; Hahn, Der Pilz-Sammler, /.
12 (coloring poor); Krombh. Abbild. ai 39. f. 21-24; Sicard,
Hist. Nat. Champ. A. 43. f. 243 (poor); Venturi, Studi Micol.
pl. 13. f. 126 (poor).

DISTINGUISHING FIELD-MARKS: This species is of about the
size of Lactaria subdulcis and sometimes of nearly the same color,
but usually it is a darker-fulvous or more red-brown, and the
flesh is firmer. Тһе odor is usually faint at first but becomes
strong as it dries. То me the odor is like that of slippery-elm
bark. The pileus is polished in appearance and does not fade
with age nor become rimulose.

The European writers describe the pileus as zonate, but no
zonate specimens have been reported in the United States. In the

Б'е
Onoinal deccrrintian В
©

which would seem to indicate that the zonation was not marked by
a difference in color so much as by concentric wavesor furrows.

In North Carolina I found several times in the same place and
once in a different locality plants which were different in several
points from the typical Z. camphorata, but had the characteristic
odor both when fresh and in drying, though slightly fainter.
When dried they closely resemble small forms of Г. camphorata.
Until I have made further collections I hesitate to describe this
plant as a distinct species, but it seems to me to be more than a
variable form of Г. camphorata, and hence I shall assign it the rank
of subspecies under the following name and characterization :

64a. Lactaria camphorata fragilis subsp. nov.

Pileus soon depressed in the center, with or without an umbo,
very thin, snuff-brown (303) or burnt-umber (304) toward the
center, not fading, azonate, dry, glabrous, pruinose, sometimes
rugose in the center 2.5-3.5 cm. broad, margin crenate and some-
what sulcate; gills maize-yellow (36. 2. 4), not crowded, about
five times broader than the thickness of the pileus; stem burnt-
umber, stuffed, becoming hollow, 2-5 cm. long, 4-0 mm. thick ;
latex watery, mild, abundant. Other characters like those of Z.
camphorata.

100 LACTARIAE OF THE UNITED STATES

The type specimens are in the herbarium of the N. Y. Bot.
Garden and are ло. 33, 1907.

65. LACTARIA RIMOSELLA Peck, Bull. №. У. St. Mus. 105: 37
1906. [As Lactarius]

Pileus fleshy, thin, rather firm, convex-umbonate, then de-
pressed іп the center, brownish-terra-cotta (334) fading to red-
brown-terra-cotta ( 332. 7. 1) azonate, dry, glabrous, rugose from
the center, then cracking into minute areas, 3-6.5 cm. broad,
margin involute then arched, even, sometimes wavy ; gills whitish,
then somewhat ochraceous, and finally colored like the pileus, a
few forking near the stem, close, decurrent, 4—6 mm. broad; stem
colored like the pileus, but not so expallent, equal or tapering
upwards, pruinose at the apex, tomentose at the base, stuffed, then
hollow, 2-6.5 cm. long, 5-10 mm. thick; flesh isabelline when
young, colored like the pileus when old, odor faint, somewhat like
that of Z. camphorata ; spores white, broadly elliptical, echinulate,
7 x85; latex watery or colored like skimmed milk, taste mild,
but a little woody.

Has.: In mixed woods under beech trees among ferns; also
“оп bare soil in woods or on banks of earth by roadsides ” (Peck)
July and August.

Пізтків.: New York, Peck (type); Vermont, Burlingham 39,
1006. ;
Плозт.: Bull. N. Y. St. Mus. 105: pl. 95. f. 7—11.

DISTINGUISHING FIELD-MARKS : This species resembles Z. can-
phorata in size and color, but differs in the pileus becoming
rimulose-areolate, in its fading to a more pinkish-terra-cotta when
mature, or in any case as soon as collected, and in the thin bluish-
white or watery milk. Тһе odor, which is very faint, does not
become stronger in drying.

The type specimens are in the herbarium of the N. Y. State
Museum at Albany.

66. LACTARIA SUBDULCIS (Pers.) Fr. Epicr. 345. 1838.
[As Lactarius]
Agaricus lactifluus dulcis Bull. Herb. Fr. pl. 224 А, В. 1784.
Agaricus subdulcis Pers, Syn. Meth. Fung. 433, 434. 1801.
Lactarius subdulcis Fr. Epicr. 345. 1838.
Lactarius subserifluus Longyear, Rep. Mich. Acad. Sci. 1901: 57
59, 1902.

LACTARIAE OF THE UNITED STATES 101

Pileus fleshy, thin, firm, convex, papillate, becorhing depressed
to infundibuliform, fulvous (308), isabelline (309), or reddish-fulv-
ous, not fading, azonate, dry, glabrous, smooth, 1.5-5 cm. broad,
margin involute, then spreading, sometimes flexuous ; gills whit-
ish or tinted with isabelline, becoming pruinose, sometimes fork-
ing, close, adnate or decurrent by a tooth, up to 3 mm. broad ;
stem of the same color as the pileus or paler, nearly equal or
tapering upwards, glabrous, or sometimes slightly pubescent at
the base, dry, stuffed, becoming hollow, 2-7 ст. long, 2-6 mm.
thick ; flesh whitish or tinted with isabelline or fulvous, odor none ;
spores white, globular to broadly elliptical, echinulate, 7 x 8 м;
latex white, unchanging, mild or slowly acrid to bitterish. Ædible.

Нав.: In the woods or on the border of woods, common.
July to September.

DISTRIB. : New York, Peck, Earle; Vermont, Jones, Burling-
ham 9, 1906; Maine, White, Murrill ; Connecticut, Underwood ;
New Jersey, E//is, Murrill & Earle; Pennsylvania, Murrill ; Nir-
ginia, Murrill ; Alabama, Underwood & Earle ; Illinois, Glatfelter ;
Ohio, Beardslee; Missouri, Glatfelter ; North Carolina, Bur-
lingham.

Плозт.: Вапа, Champ. Nice, M. 20. f. 4-10 (poor); Bern.
Champ. Roch. ai 38. f. 1. (bad) ; Bolt. Geschichte, p/. 3; Britz.
Lact. f. зг; Bull. Herb. Fr. A4. 224. f. А, В; Cooke, Br. Fungi,
pl. 1002 (poor); Cordier, Champ. Fr. pl. 26. f. 1 (very bad);
Gillet, ai тут, [393] ; Hahn, Der Pilz-Sammler, f. 13 (poor) ;
Harzer, Pilze, al 53 (very bad); Lanzi, Fung. Mang. pl. 51. f.
3; Quél. Champ. Jura, ai. 17. f. 3 (bad); Roumeg. Crypt. Ilust.
f. 141 (uncolored); Sicard, Hist. Nat. Champ. А 44. /. 238;
Sowerby, Eng. Fungi, 2/. 204, Agaricus lactifiuus (?).

Exsic.: Ellis & Everhart, Fungi Columbiani 7736 ; Herpell,
Sammlung praparirter Hutpilze 48 ; Karsten, Fungi Fennici 510;
Thiimen, Fungi Austriaci 973 ; Sydow, Mycotheca Marchica 3320.

DISTINGUISHING FIELD-MARKS: The fulvous or isabelline color
of the pileus and stem, the rather thin but firm pileus, which is
dry, glabrous, and usually smooth, the lack of odor when fresh
or in drying and the mild then slightly bitterish-acrid taste of the
white latex. The plants are comparatively small and slender.

There seems to be much variation in the color of the plants as
well as in the form. I have found specimens which differed from
the typical Z. suódu/cis only in having the margin distinctly striate,

102 LACTARIAE OF THE UNITED STATES

or the surface rugose, and both of these I suspect to be common
variations in the plant. Gillet describes three “ varieties” of 2.
subdulcis, two of which Peck reports to have found in the United
States, but I have collected only one of these, rufa. These two
forms, which Gillet terms “ varieties," he describes as follows:
Stem spongy, cap becomes more concave, of a deep live russet-chestnut,

but unpolished, taste sweet.
Stem full, then hollow, cap russet-cinnamon, sublustrous, taste sweet,

then acrid, bitter. cinnamomea.,

Lactarius subserifiuus Longyear differs from the typical Z. sub-

dulcis only in the more distant gills and the crenate margin, and
the watery latex. It seems to me to be а depauperate form of 2.
subdulcis, due probably to ecological conditions, all of the speci-
mens having been collected in high, open woods. Mr. Longyeat
states that they were growing with small forms of Г. suddulcis.

67. LACTARIA SERIFLUA (DC.) Fr. Epicr. 345. 18438. [As Lac.
tarius.| — Schrót. in Cohn, Krypt.-Fl. Schles. 3:
534. 1889

Agaricus seriffuus DC. Fl. Fr. 6: 44. 1515.
Agaricus gynaecogalus Otto, Versuch Ара. 76- 1810.

_ Pileus fleshy, firm, not very thick, at first plane with the mar-
gin inrolled, then depressed with the margin elevated, brown-fawn

307), azonate, dry, glabrous, 5-8 cm. broad, margin sometimes
flexuous ; gills yellowish, close, decurrent 2~3 mm. broad ; stem
of the same color as the pileus, nearly equal, dry, glabrous, solid,
shorter than the breadth of the pileus, 4-7 mm. thick ; flesh whit-
ish; spores whitish, subglobular, echinulate, 7-8 м; latex semi-
transparent, resembling whey, mild (acrid, fide DC.), scanty.

Нав. : On ground in woods, in moist and shaded places. July
and August.

Distris.: New York (Long Island), Peck. It has also been
reported from Pennsylvania by Herbst.

Плозт.: Berkl. Outl. pl. 13. f. 4; Britz. Lact, f. 37 (poor);
Cooke, Br. Fungi, ai 1012; Krombh. Abbild. pl. до. f. 15, ТА
18; Lucand, Champ. Fr. pl. б (good); Peck, Bull. N. y. St
Mus. 105: pl 95. f. 7-11. |

Exsic. : Sydow, Mycotheca Marchica блг. ;

DISTINGUISHING FIELD-MARKS : Тһе whey-colored, semi-trans

LACTARIAE ОЕ THE UNITED STATES 103

parent latex is the prominent feature of this plant. In color it
varies ‘from brownish-fawn to cinnamon, or it may approach the
color of L. camphorata. Peck says that our plant has a slight
aromatic odor, but no mention of an odor is made by the European
writers, and it may prove not to be constant in the American plant

68. LACTARIA ISABELLINA Burl. Bull. Torrey Club 34: 88.
1907. [As Lactarius]

Pileus fleshy, not very thick, convex, then broadly convex, at
length infundibuliform, umbonate, red-fulvous in the center, buff
(309) toward the margin, all fading to buff when mature, azonate,
dry, glabrous, somewhat roughened and wrinkled in the center

FIGURE 15. Lactaria isabellina Burl. No. 38, 1907.

especially when old, 3—4.5 cm. broad, margin glabrous, even or
at length faintly striate, sometimes areolate-wrinkled ; gills pale-
yellowish, or tinted with the same color as the pileus, becoming
reddish where bruised, forking near the stem or midway to the
margin, close, thin, slightly decurrent, 3 mm. broad, or twice the
thickness of the pileus ; stem of the same color as the pileus, equal
or slightly tapering upwards, tomentose at the base, stuffed, be-
coming hollow, 4 cm. long, 6 mm. thick ; flesh white, staining yel-
lowish from the milk ; spores white, slightly echinulate, 6-7.5 # X
7-8.5 и; latex white, at length (after five minutes) becoming sul-
phur-yellow, or at least staining the gills and flesh yellow, astrin-
gent, then acrid, abundant. (FIGURE I5.

Has.: In leaf-mold or sphagnum, moist, mixed or spruce
woods. August and September.

Distris.: Vermont, 460 meters, Burlingham 85, 1906, type ;
North Carolina, 1500 meters, Burlingham 38, 1907.

104 LACTARIAE OF THE UNITED STATES

DISTINGUISHING FIELD-MARKS: This species differs from 2. suó-
dulcis, which it most closely resembles, by the richer buff color and
by the change in the color of the latex or bruised portions of the
gills and flesh. The type specimens are in the herbarium of the
N. Y. Botanical Garden.

69. .LACTARIA MUTABILIS Peck, Ann. Rep. N. Y. St. Mus. 43:
1800. [As Lactarius |

Pileus fleshy, thin, convex, then nearly plane, reddish-brown,
the disk and zones darker, zonate when moist, azonate when dry,
moist but not viscid, glabrous, 5-10 cm. broad; gills whitish,
becoming tinted with yellow in the mature plant, close, adnate,
broader than the thickness of the pileus; stem of the same color
as the pileus, equal or slightly tapering upwards, glabrous, stuffed,
2.5-5 cm. long, 6-10 mm. thick; flesh colored like the surface,
odorless; spores subglobose, rough, 7.5 м; latex white, unchang-
ing, mild.

Нав.: “ Low damp places.” June and September.

Disrris.: New York, Peck.

Плозт.: Peck, Ann. Rep. N. Y. St. Mus. 43: M. т. f. 1-4

DISTINGUISHING FIELD-MARKS: The large size, the reddish-
brown color, and the zonate condition of the moist pileus. Ace p
cording to Peck, the zones are made up of ** more or less confluent
spots." Тһе dried specimens resemble Z. suddulcis in color, but
they are much larger. E

The type specimens are in the herbarium of the N. Y. State
Museum at Albany. Ihave not seen this plant living and the
above description is arranged from the original description after
an examination of the type specimens.

70. ТАСТАВТА PARVA Peck, Ann. Кер. N. Y. St. Mus. 29: 44
1878. [Аз Lactarius.|— Hennings, in Eng. & Prantl, S
Nat. Pflanzenfam. 1!**: 216. 1898
_ Pileus fleshy, thin, broadly convex or nearly plane, then
slightly depressed in the center, sometimes with a small umbo,
first umber tinged with lilac, fading, azonate, dry, glabrous, 1.5-3
cm. broad, margin inrolled and pruinose at first, then spreading, 3
even ; gills whitish or yellowish, becoming pale dull-green wher |
injured, and at length brownish, some forking near the stem, close,
adnate to slightly decurrent, 2-4 mm. broad; stem paler than tie
pileus, equal or slightly tapering upwards, glabrous or merely

LACTARIAE OF THE UNITED STATES 105

pruinose, stuffed, 1.5-5 cm. long, 4-10 mm. thick ; flesh tinted
with umber, staining pale dull-green, then brownish where injured,
odorless; spores white, subglobose, slightly echinulate, 6.5-8 и;
latex white, unchanging, tardily but decidedly acrid.

Han: In woods on decaying wood. August and September.

Distris.:, New York, Peck (type); Vermont, Burlingham
101, 1906.

The plant has been collected also by Guillet in Toronto,
Canada.

DISTIN FIELD-MARKS: The pale dull-green color which
the wounded gills slowly assume serves to separate this species
from the other members of this group with the exception of L.
varia. It may be distinguished from the latter by its smaller size,
the prevailing umber color of the pileus and stem, and the per-
fectly glabrous, smooth surface of the pileus. For the most part
Г. parva grows on decaying wood іп moist woods, while Г. varia
grows on the ground.

The type specimens are in the herbarium of the N. Y. State
Museum at Albany.

71. LacraRIA VARIA Peck, Ann. Кер. N. Y. St. Mus. 38: 126.
1885. [As Lactarius. ] — Hennings, in Eng. & Prantl,
Nat. Pflanzenfam. 1!**: 216. 1898

Pileus fleshy, thin, convex, then plane, depressed in the center,
sometimes with a small umbo, gray to brown, often with lilac tints,
azonate, or rarely narrowly zonate on the margin, not viscid when

ing pale dull-greenish where wounded, close, adnate to decurrent ;
stem of the same color as the pileus or paler, equal, glabrous,
stuffed, firm or spongy, 2.5-6 cm. long, 4-8 mm. thick ; flesh
white, odorless; spores white, 7-8 м; latex white, unchanging,
slowly acrid.

Нав. : On the ground in moist woods. September.

Distris.: New York, Peck (type); Vermont, Jones; Massa-
chusetts, Davis; New Jersey, Sterling.

DISTINGUISHING FIELD-MARKS: This species
color that that characteristic alone cannot serve to distinguish it,
but the prevailing gray color together with the glabrous, dry,

varies so much in

D

106 LACTARIAE OF THE UNITED STATES

` sometimes glistening pileus, the change in the color of Ше
wounded .gills and the lack of odor will help to determine it. In
color it often approaches Г. parva Peck, but, judging from the
specimens of the latter which I found in Vermont and the type
specimens of Z. varia, the two species are plainly distinct, although
closely related. Тһе type specimens of 2. varia are a pale gray,
while Z. parva remains some shade of umber when dried. As I
have not seen L. varia living, I have arranged the above descrip-
tion from the original description and some additional notes from
Professor Peck and from observation of the type specimens which
are in the herbarium of the New York State Museum at Albany.

DOUBTFUL SPECIES

LACTARIA ILLACHRYMANS Berk. & Rav. Berk. & Curt. Ann. and
Mag. Nat. Hist. П. 12: 425. 1853. [As Lactarius |
“Small; pileus thin, firm, convex to plane, then umbilicate,

pale-fulvous, obscurely two- to three-zonate ; stipe solid, white ;

lamellae white, crowded, narrow, subdecurrent, not milk-bearing
when broken ; taste slightly aromatic, subacrid.”

Has. and Loc.: In swamps in South Carolina, Ravenel.

This species has been reported also from North Carolina by
Curtis. I have not seen the type specimens, but the coloration
and zonation given in the above description would indicate that
the species belongs to the Lactariae, and that the absence of milk
may have been due to the age of the plant or to the fact that when
the milk is scanty it will exude only immediately as a vigorous
mushroom is collected. I would suspect that this number repre-
sented a form of some plant which is under favorable and normal
conditions latex-bearing.

LACTARIA ALBIDA Peck, Rep. N. Y. St. Mus. 38: 126. 1885.
[As Lactarius]

" Pileus thin, plane or slightly depressed, glabrous, dry, white ;
lamellae subdistant, adnate or slightly decurrent, white, the inter-
Spaces venose; stem equal, solid, glabrous, white ; spores white,
0003 to .00035 in.; milk white, taste acrid.”

I have not seen this growing and the dried plant specimens do
not show distinguishing characteristics, Professor Peck says ІП 4
recent letter, replying to an inquiry regarding this species, “На

LACTARIAE OF THE UNITED STATES 107

found in the Catskills a small form of what I take to be Г. albidus
Pk. The species belongs to the section Russularia, and I am not
quite sure it is more than a white variety of 2. varius Pk., which
belongs to the Russuaria.” Further collections will undoubtedly
serve to place this either as a distinct species or as a form of Lac-
taria varia. :
DOUBTFUL SPECIES

Lactaria mitissima has been reported from Pennsylvania by
Herbst and from California by Harkness & Moore, but the spec-
imens from which the determinations were made have not been
preserved. The description given by Herbst seems to conform
very closely to Г. subdulcis. The species is described in Mc-
Ilvaine's One Thousand American Fungi, 181, and is figured in
Cooke's British Fungi, 2/. гоог. It may be distinguished from Z.
subdulcis by the more golden-fulvous color of the pileus and the
orange stem, and by the more abundant latex. In Fries, Monogr.
Hym. Suec. 2 : 180, the pileus is described as subviscid when moist.
If this condition exists, the species should be classed with the
Quietae. -

INDEX TO SPECIES AND GROUPS

(Synonyms in Italics)
Agaricus, volemus, 90
aerugineus, 46

zonarius, 52
AGGLUTINATAE, I4, 40

RISEAE, I4,
glyciosmus, 8 HELVAE, 14, 7
aecogalus, 102 Hypophyllum,
helvus, 74. lateritium, 6
Aysginus, 55 INSULSAE, 14, 50
'choratus, 93 CTARIA, I
Indigo, бі acris, 8
insulsus, 51 aeruginea, 46
intermedius, 32 affinis, 52
lactifiuus agglutinata, 41, 42
lactifluus dulcis, 100 albida, 106
Listeri, 18 alpina, 79
lividorubescens, 49 atroviridis, 28
Necator, 44 aspidea
oedematopus, до aspideoides, 48
атепиз, 21 4 Bensleyae, 8
piperatus, 18, 20 Я blennia, 46
plinthogalus, 84 Calceolus, 97
pyrogalus, 27 camphorata, 98
quietus, 63 mphorata fragilis, 99
resimus, 33 Chelidonium, 59
roseo-violascens, 47

chrysorhea, 73

LACTARIAE OF THE UNITED STATES 109

LACTARIA (continued) subpurpurea, Co, 61
grisea, 80 subtomentosa, 88
haemorrhea, 57 subvellerea, 23
helva, 74 subvelutina, 96
Hibbardae, 83 Sumstinei, 86
hygrophoroides, 92 theiogala, 69, 70
hysgina, 55 torminosa, I2, 30
ichorata, 93 trivialis
illachrymans, 106 turpis, 44
Indigo, бі varia, IO
insulsa Ge vellerea, 22, 24
involuta, 26 vieta, 68
oben 103 ‘ zonaria, 52
lactiflua, 90 Lactariella, 5
нн; 43 azonites, 84
ligniota, 86 | ligniota, 86
lividorubescens, 49 Lactariopsis, 5
_ luteola, 95 Lactarius, I
maculata, 50 : aquifluus, 7
minuscula, 6 aq. brevissimus, 74
icr 107 brevipes, 6
| mucida, 5 brevis, 69
: eoi 104 |. €ontroversus, 37
: nitida, 63 5
3 oculata, 64 . distans, 92
pallida, 66 foetidus, 95
| paludinella, 66 fuliginosus, 84
| ; fuliginosus major, 86
Peckii, 76, 77 fumosus, 84
amena, 2I livescens, 49
piperata, 12, 18 platyphyllus, 52
plinthogala, 5, 84 pyrogalus, 27
plumbea, 29 : saccharium,
pubescens, 37 sordidus, 44
eta, 63 Icis oculatus, 64
regalis, 36 subserifiuus, YOO
i uvidus, 49
rimosella, 100 villosus, 30, 31
ruf. volemus,
rufula, 79 xanthogalactus, 69
2 rusticana, 27 /ACTIFLUAE,
| salmonea, 88 PIPERATAE, 14, 18
E sanguinalis, 37 PLINTHOGALAE, 14, 83
| scrobiculata, 32 QUIET.
seriflua, 102 RUSTICANAE, 14, 27
speci 35 THEIOGALAE, 14, 09
subdulcis, 100 TORMINOSAE, 14, 29

2 | subinsulsa, 45 De 14, 53

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OBSERVATIONS ON `

— — CALIFORNIA VINE

ron
ee

Observations on the California vine disease
Овмоко BUTLER
(WITH PLATES 1-5)

I

Introduction

In the height of the summer of 1886, the grape-vines in Los
Angeles and bordering counties, in California, began to show very
marked and alarming symptoms of disease. The vineyards in
the environs of Anaheim, then a flourishing viticultural center,
were the most seriously affected. In fact, Anaheim is generally
considered the birthplace of the scourge that had thus suddenly
appeared and was, within the next few years, to devastate Los
Angeles and Orange counties. The Anaheim disease, as the new
malady was called before it received the name of California vine
disease, gradually decreased in violence in each succeeding year
after 1886, and, today, one may say that it is little feared by the
growers. To be sure, Anaheim is no longer a viticultural center,
but the vine is nevertheless extensively grown in places where the
malady existed in 1886, and there are vineyards in Los Angeles
County that have passed through the years of the epidemic with-
out serious loss.

But if the California vine disease is a malady of little economic
importance іп 1010, it was a very grave problem to face in 1886.
In August of that year F. W. Morse began an investigation of
the malady, under the direction of the director of the Agri-
cultural Experiment Station, University of California, and pub-
lished, a few months later, the first description that we have of
the California vine disease.

From this author we gather: that the disease first became ap-
parent by a failure of the vines to bud, or, as occurred more gen-
erally, in a noticeable backwardness in starting, which often ex-
tended to six weeks, the foliage of the vines thus late in leafing

111

112 OBSERVATIONS ON THE CALIFORNIA VINE DISEASE

out soon becoming blighted ; that the disease might affect the vine
at any time from early spring until the autumn; that it was
reputed to be, in some cases, very rapid in its action, affecting an
apparently healthy vine in the space of a day. Morse himself
had observed no such rapid changes, the changes noticed being
only “such as one may frequently note in any vineyard where
sudden variations in meteorological conditions occur, and sun-
burn or scald follows” ;* predominantly, however, the progression
of the disease was gradual.

The malady was characterized, to continue our quotations, by а
“drying up, and apparent burning of the foliage at all times,
up to the time of ripening of the fruit. ...2 Тһе leaves in
general have the very decided appearance of sunburn. When the
vineyard is seen as a whole the foliage looks withered, leaves
partially dried and wrinkled, and large parts of them have be-
come red, the affected part of each leaf assuming no regular
shape or particular position. The canes having the most upright
growth and those most exposed have the leaves the worst
affected; low growth is less troubled.’

“ Among the several varieties the Mission is, undoubtedly, the
worst affected. . .. Other varieties such as Golden Chasselas,
Sultana, Semillon and Sauvignon are affected to a greater от less
extent, and in about the order named."*

Morse believed that the mortality of the vines was "due 10
more or less accidental and local peculiarities of climate, soil,
moisture conditions, ес.’

In a report? submitted to the State Board of Viticultural Com-
missioners in 1890, Dowlen described the California vine disease
with care, and added some remarks on the anatomy of the cane
of affected vines which I shall have occasion to quote later. This

' Morse, F. W. Report of an examination into the phenomena and СМ:
of a supposed vine disease in Los Angeles County. Report of the viticultural
work during the seasons 1885 and 1886, College of Agriculture, University of
California, 176, 177. 1886

Loe. cit., 176.

Loc. cit; 183, 184.
в : # +; i ў
‘Report California Viticultural Commissioners for 1889-90, 57 €t SÉ: |

OBSERVATIONS ON THE CALIFORNIA VINE DISEASE 113

author did not advance any opinion as to the nature of the
disease.

Pierce’s important memoir! on the California vine disease ap-
peared in 1892 and contains many facts and figures of interest.
In describing the disease he says: “In the majority of diseased
vines, although not in all varieties, the leaf presents distinct char-
acters. They may be mentioned as constitutional and localized
characters.’

The general, or constitutional effects are “due to a failure in
the formation of chlorophyll, or degeneration of that once prop-
erly formed, in those portions of the leaf supplied by the finer
spiral vessels. These general effects are found to some extent
in nearly all varieties. . . .

“Тһе localized effects upon the leaf are most clearly seen in
the white varieties, and are especially distinct in the Muscat."
The leaves become more or less covered with yellow spots "in
that part of the parenchyma supplied with the finer spiral vessels.
These spots are often well defined, the outline being very sharp
and distinct. . . ."? j

“Тһе cane usually becomes bare of leaves before the wood
is properly ripened. Тһе end of the cane, being last to ripen, is
most immature, and soon after the leaves fall the unripened parts
turn black and become dry. This progresses more rapidly and
the dying is more complete when the leaves drop early... . . The
peculiar and unequal ripening of the cane is very marked."* f

In the roots “among the first signs of disease is а dis-
coloration and shrinkage in diameter of the finer root fibers, the
root hairs and cap. This progresses until the tissue begins to
decay. . . . The root, at last becoming wholly rotted, passes into
a brown, loose, amorphous mass,"

The fruit of diseased vines is markedly affected. “If the first
attack of the disease be violent the grape will sometimes fall from

CN
1 Pierce, N, В. The California vine disease, U. 5. Dept. Agr, Div. Veg.
Path. Bull. 2. 1802.

* Loc. сі, 41.

114 OBSERVATIONS ON THE CALIFORNIA VINE DISEASE

the bunch. This dropping of the fruit is not so strongly marked
and is less important than the drying of the berry upon the
bunch. . . . In some cases the growth of the berry is retarded.
... The drying of the fruit upon the vine is a leading effect of
the disease and is very general in all varieties and under all
Conditions "7

Pierce states that the California vine disease has а period of
incubation. He says “ In the affected district it is common to find
a vineyard of one variety looking perfectly healthy and the
adjoining vineyard of another variety badly affected or killed by
the disease. It may be that the vines are of the same age and
upon like soil. When we see a sharp line of this kind,” he con-
tinues, "drawn between varieties it is folly to say that the
disease has affected one and not the other, for it may occur that
the dying variety is found on all sides of the living one. It must
be admitted, then, that the disease has produced its effect upon
vines not yet showing those effects." Again, “it is also common
to find a few Mission vines scattered here and there in vineyards
of other varieties, they having been planted through oversight and
the mixing of cuttings. Where this has been the case, these
Mission vines have been singled out and killed by the disease 25
surely as if they were by themselves in adjoining vineyards. . -
The truth is, that all the vines have felt the same influence of the
disease, but on account of difference in hardiness some show this
influence earlier than others." Another, and тоге striking
feature of the malady is the overbearing of the vines “ while the
disease is incubating.” But overbearing is not a constant symp-
tom. Pierce notes the fact, and at the same time remarks:
“That overproduction has not always been noticed is but nega-
tive evidence, and its well-attested occurrence in a reasonable
number of cases is of more value than much negative evidence."

The California vine disease appears also to be transmitted 1n
cuttings,

Pierce does not come to any conclusion regarding the nature

* Lok єй а

OBSERVATIONS ON THE CALIFORNIA VINE DISEASE 115

In 1892 Viala and Sauvageau also published a note on the
California vine disease in the Comptes Rendus, and a memoir on
the Brunissure and the California vine disease in the Journal de
Botanique, which was republished later in the Annales de l'École
Nationale d'Agriculture de Montpellier. These authors described
the anatomical features of the California vine disease very
accurately, as I shall have occasion to point out later, and con-
cluded that this malady was produced by a myxomycete very
similar to the organism causing Brunissure ( Plasmodiophora
Vitis), but as it was infinitely more destructive they gave it
specific rank under the name Plasmodiophora californica.

In recent years Ravaz' has advanced the opinion that the
California vine disease and the Brunissure are one and the same
malady, both being due to overbearing, but the description I give
of the latter disease in chapter III will show that this view is
incorrect. If we modify, however, Ravaz's opinion to read that
the Brunissure killed a number of the vines during the epidemic
of 1886 in southern California, I think that it can then be sup-
ported on grounds. But I shall not attempt to advance the argu-
ments in favor of this modification of Ravaz's view; it would
lead me too far to do so adequately, and a brief presentation is,
in the nature of the subject, impossible.

II
Description of the California vine disease
А. MORPHOLOGY

The California vine disease affects primarily the leaves, fruit,
shoots, and canes. The arms and trunk reveal nothing of diag-
nostic value, and an examination of the roots is fruitful in con-
tradictory results.

The leaves, fruit, shoots, and canes show symptoms that are
constant from one variety, or species, of vine to another, the
observable discrepancies being due to the fact that the virulence

végétation de la vigne.

'Ravaz, L. Influence de la surproduction sur la
Ann. École Agric. Montpellier, II. 6: 5-41- 1906.
emarques sur le dépérissement de quelques vignes
. Progrés Agricole 44: 41-50, 71-73. 1905.

en Tunisie et en France.

116 OBSERVATIONS ON THE CALIFORNIA VINE DISEASE

|

|

A
|

|

PERCENTAGE OF VINES DISEASED.
|

т А
JUNE JUY А SEPT М

GURE 1.—Curve showing the general course of development of the Cali-
fornia vine disease,

OBSERVATIONS ON THE CALIFORNIA VINE DISEASE Ht

of the disease is subject to marked variations. These variations
are both individual and inter-individual. Тһе symptomatic differ-
ences between two plants are often found in replica on the shoots
ofa single vine. Тһе disease, in fact, appears to work, to use a
colloquialism, without rhyme or reason. It may affect а single
shoot, or several shoots; one side only, or the whole of a vine.
One vine may show all its characteristics, manifold though they
be, and another, a part, perhaps only one, of them. The disease
may never proceed beyond the first stages, a not uncommon
feature, or it may develop slowly until all its characters are
patent to the most casual observer; more often, however, its
development is rapid.

Тһе California vine disease, though it may appear at any time
of the growing season, usually develops to a noticeable extent
only when the vines are ripening their fruit, that is, at the most
critical period of their yearly development. This fact is brought
out very clearly in the accompanying graphic, which may be con-
sidered representative when 25 per cent. or more of the vines in
a vineyard are diseased. When less than twenty-five vines in
one hundred are affected, the curve may become either flatter or
sharper: flatter, if the malady develops gradually throughout the
growing season; sharper, if the disease develops very rapidly.
As soon, however, as one quarter of the vines become diseased it
tends to assume the form shown in the graphic.

The development of the disease is sporadic; and the malady
always develops in the following general manner whether few or
many vines are affected:

A certain number of scattering vines show the ominous symp-
toms; then more vines, unrelated positionally to the others, be-
come affected, and thus, by the continued addition of diseased
individuals, a vineyard becomes, to a greater ог less extent, in-
fested. There is no such thing, however, as spreading by ef
tiguity, no *oil spots." Such a thing as a diseased center, using
exist, nor is there in the
lves to

the term in its narrower sense, does not
different varieties of the grape-vine, if we confine ourse
the Viniferae, any marked difference in predisposition. One can-
not predict, as in the case of the powdery mildew,

for instance,

118 OBSERVATIONS ON THE CALIFORNIA VINE DISEASE

that the disease will first appear in this or that variety. We only
know that at A the disease may affect almost exclusively +; at
В that у is worse than ғ; and at C that у, и and v are equally,
or almost equally affected. But such observations as these are
only good for the season in which they are made; they may not
be true for the year before, nor yet in the year to come. Ina
word, the past is no criterion of the future: this is true whether
the vines become diseased suddenly or progressively, and I mean
by this latter term an intermittent addition rather than a natural
sequence of symptoms.

I have said that the past is no criterion of the future. This
statement, while true for the individual vine, is not necessarily
true for the vineyard as a whole. When a large percentage of
the vines in a vineyard are affected, not suddenly! but pro-
gressively, it is not infrequent that the disease reappears the
following years and becomes chronic, as it were, in a large
number of vines, though it is impossible to select with certainty
the vines in which it will be so, and gradually, in bad cases, 50
weakens them that they die.

From the behavior of diseased vines, both as individuals and
collectively, it appears that the California vine disease is suffi-
ciently variable in its mode of action to be considered as pos-
sessed of two forms, differing from one another in immediate
virulence. The one we may call apoplectic, from the suddenness
of its action ; the other chronic, from its lesser destructiveness and
predisposition to recurrence. These two forms, identical symp-
tomatically, conduce, the one rapidly, the other slowly, to the
same end, the death of the vine. It is, therefore, unnecessary, in
describing the disease, to state specifically which of these two
forms one has in mind, the line of demarcation between them
being, to all intents and purposes, but a line in point of time.

This being so, the following description determines the Cali-
fornia vine disease, provided the characters of the affected
organs are considered in conjunction with one another and not 25
separate and sufficient entities; for, and it cannot be too strongly
urged, the disease we are considering cannot be surely and safely

1 When a vine is affected салалары in all its shoots it en dies of `
chlorosis the following зеазо

OBSERVATIONS ON THE CALIFORNIA VINE DISEASE 119

identified except in the “all together.” Concretely : a vine show-
ing but a single symptom of the California vine disease is, in
itself, a doubtful subject.

Symptomatology

(а) Leaves.—The young leaves—on upper third of shoots—
generally show particular characters only in the case of the chronic
form of the California vine disease. When a vine is affected
apoplectically they either remain normal or assume the characters
common to the older and adult leaves or, when the shoots are
defoliated without death ensuing, form a healthy plume, as it
were, at their tips. In the apoplectic form the young leaves are
not indicative, but in the chronic form they are very often pre-
monitory symptoms; the vine first showing them may become no
further diseased, but it is practically certain that other vines in
the vineyard, if they are all of one variety and age, will, and
not only lightly, but severely. The characters shown by the
young leaves are, in the case of the chronic form, then of con-
siderable interest.

They are:

Case r——The leaves become pale in the intervenium, growth
ceases at the periphery— sometimes also between the veins—
and death ensues. Тһе tissues not immediately affected, not
having reached complete development, continue growing, the
leaves becoming paler in color, more or less convex, and, accord-
ing to the amount of dead tissue other than peripheral, variously
distorted. (PLATE I, FIGURES 1, 2, 3, 4.)

Case 2—The leaves, leaves more developed than those just
described but of the same coloring, do not become convex as а
Whole, but only in one or both wings of the petiolary sinus. This
distortion is accompanied by a sinking of the tissues between the
venation, and subsequent death.

n older leaves the symptoms of the disease are variable and
Cannot be accounted for by their position in regard to other
diseased leaves. Those leaves that are still in fairly active growth
may show the characters common to the young leaves described in
Case 2. In other cases, and coincident with the furrowing and
death of the wings of the petiolary sinus, there appear, to a

120 OBSERVATIONS ON THE CALIFORNIA VINE DISEASE

greater or less extent, between the veins of the remainder of the
leaf-blade, suffused yellow or red spots, which, when their
centers die, have the appearance of reddish brown maculations
surrounded with aureolae of red or yellow, аз the case may bei
(PLATE 2, FIGURE I.) If instead of, or coexisting with, the
spotting of the leaf we have a reddening or yellowing of the
intervenium, the dead tissue forms strips. Marginal discolora-
tion and death may occur, as in the adult leaves, but is not
so frequent.

In fully developed leaves the leaf-blade is not deformed. The
tissue between the veins, and the margin, also, very largely, be-
comes yellow or red, the discolored areas dying in time, the dead
tissue assuming a color which ranges from fewille-morte to gray,
according as the death has been rapid or slow. Instead, however,
of beginning as a general discoloration of the intervenium, the
disease may first appear as suffused greenish yellow spots, which,
enlarging and becoming more definite in outline, often merge
together, forming large maculations and stripes. These macu-
lations and stripes may die to the edge of the healthy tissue |
itself, but are more often surrounded by aureolae, which may be
red, red and yellow, or yellow alone. (PLATE 1; PLATE 2.)

The leaves near the base of the shoots sometimes show a slight
variation from the characters just described. Ав soon as the
spots appear between the veins, enlarge, and form stripes, the
remainder of the parenchyma becomes chlorotic. Death in the
diseased areas proceeds slowly and, when accomplished, the dead
parts being soft and crumbly, the leaf is beaten by the winds
into deeply incised fragments which hang together around the
petiole.

Diseased leaves—this remark is generally applicable—fall
sooner or later with, or without, their petioles. The fall of the
leaf prior to that of the petiole occurs, so far as I have been
able to ascertain, only when the intervenium becomes diseased
immediately around it. The death of the parenchyma then in-
volves the death of the apex of the leaf-stalk, and the blade
becomes severed from its support.

“In the varieties of the grape-vine producing white fruit, the апгеоіе аге
always yellow ; but in those bearing colored fruit they may be both yellow and
red on the same leaf, the predominant color varying with the variety.

OBSERVATIONS ON THE CALIFORNIA VINE DISEASE 121

(b) FRurr.—The fruit may be affected at any time from
setting to maturation. If the fruit is affected just after setting
the whole bunch withers and falls away; if attacked somewhat
later, on passing the hand over a diseased bunch, the berries will
come away often with, not infrequently without, their pedicels—
they may even fall of themselves. As the fruit grows older it
does not fall, but may shrivel or, if nearing maturation, ripen
imperfectly. This shriveling and imperfect maturation of the
fruit is a feature of the California vine disease. “ The drying of
the fruit upon the vine,” says Pierce, “is a leading effect of the
disease and is very general in all varieties and under all
conditions.’’!

(с) SHoots.—The effect of the disease оп the shoots depends
on the amount and suddenness of the defoliation, which, itself, is
a measure of the quality of the attack. If the vine is affected
apoplectically, the life of the shoots, or of the canes, will be more
endangered than if it be affected with the chronic form of the
California vine disease. The season of the year at which the
vines are affected is also a factor of some importance. Vines
that have suffered even complete apical defoliation on one, or
several, of their shoots in early summer, that is while they are
still growing vigorously, may not have them visibly damaged.
This is evidenced by the fact that such defoliated shoots continue
to elongate, after a period of rest, and throw out axillary foliage.

In the height of summer, and at maturation, defoliation
brought about by either form of the disease is more serious.
The growth of the vines has then normally ceased and their re-
cuperative powers аге low. In July, and to some extent, also,
in August, defoliation is followed by a progressive dying of the
shoots, the amount of death in each shoot being, as it were, a
register of its defoliation. When the disease affects the vines
after lignification has set in, and during maturation, the fall of
the foliage leaves the canes very imperfectly formed. The shoots,
instead of maturing properly, remain, to a greater or less extent,
teen. One side of a shoot will be mature, the other not.
Maturation may have proceeded normally at the base, and be

* Loc. сін, 53.

122 OBSERVATIONS ON THE CALIFORNIA VINE DISEASE

very disordered further up. Or again, lignification may be im-
perfect only around a few nodes, the internodes showing appar-
ently normal maturation, and conversely.

If one cuts a smooth cross-section of a shoot or cane that is
bearing diseased foliage, or has been defoliated, he will notice that
the wood is slightly discolored. The pith is best seen in longi-
tudinal section; it will be found discolored more or less discon-
tinuously, or turned completely brown, if the foliage has fallen.

(d) SPURS, ARMS, AND TRUNK.—Externally, the spurs, arms,
and trunk have the same appearance in diseased as in healthy
vines. This is no longer true when cross-sections are examined.

In the spurs the woody tissue of the previous year is discolored
and, often, more especially near the edge, zoned with darker lines.
In the arms the zones become less marked, and disappear before
reaching the trunk. The discoloration of the wood is much more
persistent, but hardly descends to the roots.

Besides the characters just mentioned, the arms and trunk may
show alterations due to die-back from old wounds, decay, and
various other injuries interfering with the circulation. The
tissues surrounding these impediments are generally affected in
such a manner that their differentiation from those that have
become diseased through the action of the California vine disease
is impossible. The characters found in the arms and trunk, in
the case of the California vine disease, are, therefore, of but
very little diagnostic value, and, I am inclined to think, hardly
worth the pains one must be at to find them.

(е) Roors.—The roots do not present any characteristic altera-
tions. They are either diseased, or not diseased, according to the
situation in which the affected vines are growing and the presence,
or absence, of root parasites, either animal or vegetable. If the
affected vines are suffering also from the Phylloxera Vastatrix,
Adoxus Vitis larvae, or nematode worms, the roots will appear
damaged; they may be decomposed by root-rot (Dematophora
Necatrix), or dry-rot, simply die, or, а frequent occurrence, be
entirely sound.

OBSERVATIONS ON THE CALIFORNIA VINE DISEASE 123

В. HISTOLOGY

TECHNIQUE.—In studying the anatomy of the California vine
disease I have employed the usual cytological methods. The
diseased tissues were prepared and placed, with rare exceptions,
in the fixing fluid directly in the field.

The fixative that gave me the best results is one per cent.
chromic acid. Flemming’s solution does not appear superior to
chromic acid as a fixative, and labors under the disadvantage of
blackening the tissues to such an extent that details are masked,
and decoloration of the sections in hydrogen peroxide has to be
resorted to to bring them out. Even then, however, the chromic
acid material gives preparations superior in sharpness and con-
trast. Sections made from alcoholic material give stained prepa-
rations that compare favorably with those obtained from chromic
acid material. Though alcohol is not so good a fixative as
chromic acid, I believe that it may nevertheless be advantage-
ously substituted for it in many cases. The matter occluding the
cell lumen in diseased tissues contains tannin, and is more or less
darkened by the latter fixative, which is quite a disadvantage in
all but very thin sections.

After fixing, the tissues were washed, passed into alcohol and
through bergamot oil into paraffin; or, if working partly in the
cold, from alcohol to chloroform, in which the paraffin was dis-
solved to saturation. Finally the chloroform was evaporated
off at бо degrees Centigrade.

The stain that gave me the best results for general purposes is
acid fuchsin, a concentrated aqueous solution of bichromate of
potash being used as the differentiator. This stain, prepared by
dissolving 20 grams of acid fuchsin in 100 cubic centimeters of
aniline water, gave me the best results when used in the fol-
lowing manner:

The sections to be stained, previously fixed to the slide, were
covered with warm acid fuchsin, which was kept steaming hot,
but not boiling, for a few minutes. The sections were then
washed and plunged into warm bichromate, washed again m
Water, and mounted.

* Cf. Zimmermann, Botanical Microtechnique, 197. 1893. [Humphrey trans.]

124 OBSERVATIONS ON THE CALIFORNIA VINE DISEASE

The various other stains or reagents that I have used were
prepared for the most part after the formulae given in Zimmer-
mann’s Botanical Microtechnique.

Lraves.—From the description I have given of the appearance
of diseased foliage it will be seen, upon close examination, that
leaves have always one or another characteristic in common, and
may be classified in two groups: first, those leaves in which the
maculations and striations occur in a field of green ; and secondly,
those leaves in which the spots and stripes occur in a field of
yellow. The young leaves that betray a diseased state only by
the convexity of their blade and a pale color naturally belong to
the second group, chlorosis being the characteristic of it.

This separation of the diseased leaves into two groups, while
of little value for descriptive purposes, and I made no use of it,
is of considerable assistance in their microscopic examination. I
found, as would be supposed, that corresponding to the observed
macroscopic differences, there existed microscopic differences.
These latter differences, though less pronounced perhaps than the
former, and more of degree than of kind, appear to best advan-
tage and in their truer value when considered separately from
one another. I have, therefore, considered in (a) the anatomy
of those leaves possessing diseased areas and healthy areas;
and in (b) the anatomy of those leaves that are entirely
pathognomonic.

(а) If one cuts a cross-section through a leaf, being careful to
include tissue in various stages of disease, it will present, under
the microscope, very nearly the following appearance:

The epidermal cells (of upper surface and lower surface also,
but to a markedly less degree in the latter) are full of matter
in the diseased areas which becomes less dense and gradually
disappears as one enters the healthier tissues. This matter may
be globoidal, homogeneous, or coarsely or finely granular. CR LATE
3, FIGURE 4; PLATE 4, FIGURES 3, 4, 5.) Тһе globoidal form 15
infrequent, homogeneity the rule; and both forms do not 89
beyond the areas of marked disease. When the globoidal form
is present it may pass over into the homogeneous, or disappear
as the healthy tissue is approached. The homogeneous deposit,

OBSERVATIONS ON THE CALIFORNIA VINE DISEASE 125

however, invariably changes in density, and not infrequently in
character, on nearing the healthy tissues.

In the palisade cells the deposit, as a rule, corresponds in
density with that in the abutting epidermal cells (when there
is a difference it will be in favor of the latter), but extends
greatly beyond the outer limit of the matter in these cells.
Homogeneity of the deposit is the rule, though it occurs now
and then in granular form, the size of the granules varying from
cell to cell, but not to any extent in the individual cell. The
incrusting of the primordial utricle of the palisade cells is a
striking and constant feature. The density of the incrustation
decreases, as one passes from diseased to apparently healthy
tissue. (PLATE 3, FIGURES 4, 6; PLATE 4, FIGURE 4.)

In the lacunose tissue the incrusting of the lumen is not so
regular or so marked as in the case of the palisade layer. The
row of lacunose cells adjoining the palisade tissue is more free
from it than the others, though there is no absolute constancy in
this matter. These cells contain not infrequently, though scat-
teringly, a few globules much smaller and constitutionally dif-
ferent from those in the epidermal cells, as will be shown sub-
sequently. In the remaining cells of the lacunose tissue the
deposit is either homogeneous or granular, dense or thin, corre-
sponding, in this respect, with the variations in the palisade layer.
(PLATE 3, FIGURE 4; PLATE 4, FIGURES 3, 4, 5-)

The chloroplasts are sometimes absent from the older portion
of the diseased areas ` when present their, degeneration is marked,
but, I should add, not unexceptionally, as sometimes palisade
cells may be found filled with dense homogeneous matter in which
chloroplasts, still containing starch, lie embedded. Degeneration,
however, is the rule, but is more noticeable and has progressed
further wherever the deposit in the cells is less dense; the chloro-
plasts may then be observed as protean plasmodium-like masses,
sometimes of considerable size, from aggregation of individuals.
As one progresses towards the healthier tissues, and with the
decrease in density of the deposit, the chloroplasts are generally
more difficult to observe, their resorption having usually pro-
gressed further. (PLATE 3, FIGURES 3, 4, б; PLATE 4, печ

235)

126 OBSERVATIONS ON THE CALIFORNIA VINE DISEASE

The degeneration of the chloroplasts, like the incrusting of the
primordial utricle, proceeds further in the palisade than in the
lacunose tissue. In the cells of the lacunose tissue the chloro-
plasts do not, as a rule, form large plasmodium-like aggregates.
They become vacuolate, but usually without distending to any
extent, and, in the row of cells next the palisade layer, when the
lumen is free, are inclined to fragment or degenerate into oil-
like bodies. (PLATE 3, FIGURES 3, 4; PLATE 4, FIGURES 3, 5.)

The vessels of the minor bundles of the leaves are occluded
more or less by granular or homogeneous matter, and the same
may be said of the bast, cortical parenchyma, collenchyma, and
epidermis of the main veins. (PLATE 3, FIGURE 5.)

Thylloses are often present in the vessels of the main vein.
(PLATE 4, FIGURE I.)

Viala and Sauvageau, in their study of the California vine
disease, remark that a “section cut through an apparently uni-
formly diseased area often shows breaches of continuity due to
healthy starch-replete cells which may be coextensive with the
diseased tissue.” "This observation, while perfectly correct,
applies only to diseased leaves taken from varieties of grape-
vines bearing black grapes. In these varieties diseased centers
may be surrounded directly by red aureolae,? and the cells іп
this reddened tissue are replete with starch. When therefore,
Ше aureolae of diseased centers are contiguous, or we have an
intervenar stripe dying irregularly, the prenecrotic color being
red, such an irregular alternation of starch-replete and starch-
free cells may occur. The presence of starch, however, is patho-
gnomonic, and not indicative of health, as the authors just quoted
believed. When, as occasionally happens, the dead tissue abuts
directly on that which is still green, without apparently any inter-
vening morbid cells, we do not find such a thing as starch-
replete and starch-free cells. In fact, in section, what appeared
macroscopically as a decided line of demarcation is certainly

шекер тенден.

1 Viala, Sa & Sauvageau, С. La Brunissure et la Maladie de Californie.
Ann, Ecole TR Montpellier, 7: тот. 1892. [Translation.]

“Тһе reddening of vine leaves, so far as I have meses is always accom
panied by a starch congestion. See Ravaz, L., & Roos, L. Le Rougeot de i
vigne. Progrès Agricole 44: 363-370, 392-398. 1905. А

OBSERVATIONS ON THE CALIFORNIA VINE DISEASE 137

not distinctly delineated; disorganization of the cell constituents
proceeds into the green tisue and starch, if present, is nowise
abundant.

(b) Examined in cross-section, those leaves in which the
appearance of the disease is followed by a progressive chlorosis
of the entire parenchyma differ from those in which it remains
normal, i. е., Case a., rather in degree of occlusion of the cells
and degeneration of their chloroplasts than otherwise.

The deposit in the epidermal cells (upper epidermis almost
exclusively) is either homogeneous ОГ granular, apparently
rarely globoidal. It is never very dense, though generally homo-
geneous in character where the tissue is longest diseased, and
becomes granular and finally disappears as one proceeds into the
surrounding chlorotic tissues.

The deposit in the palisade cells is usually homogeneous and
thin, or more or less granular, and proceeds well into the healthier
tissues. Starch is rarely to be found. The chloroplasts are
largely resorbed—those still remaining being smaller than in
normal tissue and reticulate—in the surrounding chlorotic tissues,
and, in the diseased spot itself, form only comparatively small
plasmodium-like aggregates. (PLATE 4, FIGURE 5.)

In the lacunose tissue the cells abutting the palisade layer are
the freest from deposit; they are frequently almost empty. The
remaining cells do not differ much in appearance from the
palisade cells; their lumen, however, is freer from deposit.

The deposit in the cells of the lacunose tissue is either homo-
geneous or granular.

Occlusion of the vessels of the smaller veins is not general,
nor do thylloses appear to occur in the vessels of the main veins.

I have just described the general microscopic appearance of
sections cut through diseased leaf tissue. I will now describe in
detail the anatomy of the pathognomonic tissues with the view
of determining as far as possible the nature of the catabolic
processes brought about in them.

In describing the California vine disease I said that this malady
may appear spontaneously as it were, от develop more or less
slowly. This statement applies not only to the individual vine
but to individual leaves.

128 OBSERVATIONS ON THE CALIFORNIA VINE DISEASE

We found that the leaf dies in spots or stripes, infrequently
without prenecrotic coloration, the rule being a reddening or
yellowing before death, even though all the stages are concurrent
as it were; we found that exceptionally the disease appears as
yellow maculations, isolated or running together, when death is
slow and confined to the diseased areas, even though the leaf
turns chlorotic—but I did not lay particular stress upon the point
that when death is very rapid, the dead tissue has a somewhat
glossy brick tint, that when less rapid it is more reddish brown
and matte and when slow, fawn-colored. These differences in
coloration of the dead tissue have, however, considerable anatom-
ical importance.

If one examines sections through material showing the color
characters mentioned above, he will obtain a conspectus of the
behavior of the chloroplasts. Thin sections must be cut, owing
to the opacity of the deposit in the lumen of the cells, when the
chloroplasts may be well brought out by acid fuchsin,—carbol
fuchsin and iron haematoxylin (the first gives the clearer
preparations) do not give as sharp a differentiation. Acid fuchsin
might almost be called a specific chloroplastid stain. Sections
placed in it for a few minutes, and then washed in bichromate | |
will show the chloroplasts deep red, the cytoplasm very faint ` —
rose, the other cell inclusions being practically colorless. ВУ |
means of this stain the chloroplasts may be studied without fear
of misinterpretation. Carbol fuchsin and iron haematoxylin,
the latter especially, did not appear to me quite so trustworthy `
and were soon discarded.

If we take, then, a series of sections through diseased tissues ,
that have died with various rapidities and stain them, preferably | S
in acid fuchsin, we shall find that the resorption, vacuolation
and plasmodium-like aggregation of the chloroplastids is, 10 4 E
certain extent, inversely proportional to the amount of lumes
occlusion. In the tissues that have died very rapidly the deposit
is homogeneous, dense, and the chloroplasts hardly show more
than a slight vacuolation and some appear, in optical section,
as hollow elliptical spheres; their center is not a vacuole, how
ever, but a starch grain, as the blue color they assume оп

А А еа де А IRR E В,

OBSERVATIONS ON THE CALIFORNIA VINE DISEASE 129

addition of an iodine solution readily shows. (PLATE 3, FIGURE
4.) This observation regarding the presence of starch in a few
chloroplasts applies only to those cases where the prenecrotic
coloring of the diseased tissues is red, but does not apply when
the change of hue is not apparent or yellowish. In these cases
the chloroplasts do not appear to contain starch, and their vacuo-
lation is more pronounced, which fact would lead one to suspect
that death does not really occur without some previous discolora-
tion of the tissues, however transient it may be.

When the death of the tissues is less rapid, vacuolation and
distension of the chloroplasts is marked, and one will observe
plasmodium-like aggregates here and there in the cells, but in
more important masses at the lower extremities. With the final
occlusion of the lumen all further changes are arrested. (PLATE
3, FIGURES 3, 6; PLATE 4, FIGURES 3, 4, 5 )

When the cells die with moderate rapidity, 1. e., when the sec-
tions are taken from typical diseased leaves, the vacuolation
and plasmodium-like aggregation of the chloroplasts seems to
reach a maximum. The homogeneous matter occluding the
lumen, though still dense, is light-colored.

As the rapidity of death still decreases, the tendency of the
chloroplasts to run together is less marked; though still vacuo-
late, they distend less and their resorption progresses further
and further. The density of the deposit in the cells also
decreases. (PLATE 4, FIGURE 5.)

Finally we come to the stage (diseased leaves that become
entirely chlorotic, the original diseased areas dying first and thus
remaining distinct) when sections through an autumn leaf near
its fall and a diseased leak ет not at at alb Ot Ver? little in
appearance; traces of chloroplasts may remain in both cases, and
the lumen may also be slightly occluded by homogeneous or gran-
ular matter. (PLATE 3, FIGURE 2; PLATE 4, FIGURE 5.)

The chloroplasts ар to the stage of plasmodium-like aggrega-
tion stain readily, but as their resorption progresses further,
they stain less readily and, in fact, at the final stage (autumn-
leaf stage may I not call it?), they hold Ше acid fuchsin less
readily—the other cell inclusions not at all—and decolorization
in potassium bichromate is unnecessary.

130 OBSERVATIONS ON THE CALIFORNIA VINE DISEASE

The facts that I have just related in regard to the behavior
of the chloroplasts refer almost exclusively to the cells of the
palisade tissue. In the cells of the lacunose tissue the chloro-
plasts become vacuolate, but remain small, as a rule, and their
resorption progresses gradually. Тһе occluding of the lumen of
these cells does not occur so rapidly and is rarely so dense or dark
in color, even in the most rapid cases of death, as that of the
palisade cells.

I remarked, in a previous passage, that the row of cells of the
lacunose tissue abutting on the palisade layer was very free, when
compared to the other cells of the same tissue, from deposit.
When the lumen of these cells is free from deposit the chloro-
plasts not only become vacuolate but fragment and, it would
appear, decompose with the formation of oil-like bodies, which,
when small, stain like the chloroplasts, but do not color, when
larger, as vividly, if at all, in acid fuchsin, which fact leads me to
believe that, if originally largely decomposing chloroplastid
remnants, they grow by accretion of other proteid substances;
this is brought out clearly when sections are stained with rosani-
line: the smaller bodies will appear red, the others violet. They
all stain, however, more vividly in safranin and eosin than the
chloroplasts themselves, which would tend to show that their
composition is fairly complex. |

During the course of my remarks on the behavior of the

chloroplasts- I have frequently made mention of the homogeneous — |

substance filling the cell lumen. The various stages of chloro-
plast degeneration we found to depend on the relative amount and
rapidity of production of this substance. It is therefore, im-
portant for us to determine the nature and origin of the homo- |
geneous deposit and its homologues, the globules and granulat ,
matter. This I will now attempt to do. e
From my observations on the degeneration of the chloroplasts
it plainly appears that the substance occluding the cell lumen is
not a product of their decomposition. That from the decomposi-
tion of the chloroplasts there appears to result, in some cases, the
formation of oil-like bodies is no contradiction to this statement
The latter form of decomposition is rare. Furthermore, the n

OBSERVATIONS ON THE CALIFORNIA VINE DISEASE 131

that the decomposition of the chloroplasts is in inverse ratio to the
density of the occluding matter, precludes the assumption that,
in the major cases, these bodies act as nuclei of condensation, like
crystals in supersaturated solutions. But if the chloroplasts play
no part in the formation of the homogeneous matter, or its homo-
logues, whence does it come? The answer to this question in-
volves considerable difficulties, and must, at best, be largely
hypothetical.

The deposits in the lumen of the cells are probably genetically
related, though it would appear, from their variation in form,
that their ultimate composition is somewhat different. They all
behave very similarly to reagents and stains. They dissolve in
Javelle water, but are not at all, or but little, affected by either
hydrochloric or sulphuric acid. А concentrated solution of potash
has practically no effect on the homogeneous and granular
matters, but will sometimes remove the globules in the epidermis.
Fixing diseased tissues in Flemming’s solution is apt to make the
homogeneous, granular, and globoidal matters so dark that, for
staining purposes, sections taken from such material are very
imperfect. Chromic acid, one per cent. solution, does not change
the color of the cell occlusions to any extent, and sections taken
from material fixed in it are hardly more opaque than those taken
from alcoholic material.

The deposits in the cells turn black when the sections are
placed in a saturated solution of iron acetate; the black color
Flemming’s solution imparts to them may be very largely removed
by peroxide of hydrogen.

Amongst the stains, iron haematoxylin’ is retained vigorously
by the deposits, but Bóhmer's haematoxylin is without effect.

“Тһе homogeneous matter colors deep red in safranin, takes eosin
readily, colors in erythrosin, tropaeoline oo, carmalum, rosaniline?
(dull red), orcein and hydrochloric acid.

From these reactions of the deposits it is clear that they con-

tain tannin (action of iron acetate, regeneration of osmic acid by

*I mordanted the sections in "liquor ferri sulfurici oxidati,” diluted with
two volumes of water, for twenty-four hours; stained in 1 per cen
of hematoxylin (Benda's) and differentiated in 20 per cent. ace

* An alcoholic solution of equal parts fuchsin and methyl violet.

t. solution
tic acid.

139 OBSERVATIONS ON THE CALIFORNIA VINE DISEASE

hydrogen peroxide), proteid matter (eosin, safranin) ; that they
contain .also a gum, somewhat similar to wound-gum, would appear
from their reaction to orcein and rosaniline.

The deposits in the cells are, then, complex in composition, but
it is not improbable that a decomposition product of starch forms
their predominant ingredient. The following facts strengthen
this view: |

The cell walls and cytoplasm of healthy tissues stain blue in
Bóhmer's haematoxylin, but pathognomonic tissues, on the other
hand, do not. If Bóhmer's haematoxylin and safranin are used
together the healthy tissues will appear as we have just described
them, but the cell walls and cytoplasm, when diseased, are red or
reddish. This would prove that a decided change takes place in
the cell walls of diseased tissues simultaneously with the change
in the cytoplasm, but as this change is visible only upon colora-
tion, it is not likely that the cell walls contribute materially to the
formation of the deposits in the lumen. The deposits must,
therefore, arise as decomposition products from one or more of
the cell contents. I believe that starch is the most important
contributing substance of them all, and circumstantial evidence
favorable to this view is not lacking.

We have seen that the more rapid the death of the leaf the
denser the occluding deposit. We have seen also that when the
tissues die suddenly without apparent prenecrotic coloration,
they are, nevertheless, as free from starch as tissues that become
previously colored. We have seen also that reddened tissues аге
replete with starch and practically free from deposit, but 10
sooner die than occlusion becomes noticeable and starch, to all
intents and purposes, absent ; we have seen further that the longe!
the period of chlorosis before death, the freer the cell lumen
from deposit, and starch, it is well known, is quite scarce in
chlorotic tissues. It appears to me, therefore, that the substance |
(of the nature of wound-gum) found in the cell lumen, as атап |
lar or homogeneous matter, is largely derived from starch.

The deposits we find in diseased cells do not appear to contain
pectic substances in any considerable amount, for Victoria blue `
and chloriodide of zinc do not give the typical reactions, and the

/
OBSERVATIONS ON THE CALIFORNIA VINE DISEASE 133

retention of methylene blue seems to be due to the presence of
tannin.

In my observations, just described, on the anatomy of diseased
leaves, I never mentioned that hyphae, plasmodia, and bacteria
were always absent from pathognomonic tissues: such, however,
is the case. In diseased tissues that have not been weathering
under conditions favorable to the growth of fungi or bacteria no
foreign organism of any kind is to be found. I must, therefore,
deny the existence of Plasmodiophora californica, but in so doing
I wish to impugn only the interpretation of Viala and Sauvageau,
not the correctness of their observations. The facts exist as
they saw them, but not as they interpreted them.

CaxEs.— The anatomy of the shoots of diseased vines does not
reveal any facts of importance until they begin to mature. Their
maturation is somewhat fantastic and the resulting canes appear
interspersed with green immature tissue. This immature tissue
constitutes the immature spots so characteristic of the California
vine disease and will, therefore, occupy our attention almost ex-
clusively in the following remarks.

I think it well, for clearness sake, to preface my observations on
the immature spot with a brief description of a healthy cane, as
seen in cross-section. We find, around the pith, a ring of w
composed of wood-fibers and large vessels interspersed radially,
at equal intervals, by the ligneous medulla; beyond the wood,
the cortex, corresponding to the wood fascicles, Ше basts
containing two to three or four rows of fibrous bundles and
separated by a parenchymatous and widening prolongation of
the medulla; beyond the bast, and separating it from the remain-
ing cortical tissues, the suber; beyond the suber, and capping the
basts, as it were, the pericycles; the other tissues, parenchyma,
collenchyma, and epidermis, have turned brown, and have more
or less collapsed.

In immature spots the departure from t Е
sketched it, is as striking as unexpected. Around the pith we
find the ring of wood to be of unequal, instead of equal, diameter,
and the wood fascicles to be of unequal development: at the
center, or to one side of the center, in the i

he normal, as I have

mmature spot, they

134 OBSERVATIONS ON THE CALIFORNIA VINE DISEASE

are very undersized and the vessels themselves fewer in number
and subnormal; the phloems corresponding to these undersized
wood fascicles are greatly reduced in size, free from bast-fiber
bundles, which normally should be 2 or 3 in number, and covered
with dwarfed pericycles. As one works towards the sides of the
immature spots the wood fascicles increase in size and assume
normal proportions; the phloem correspondingly increases in size,
but the production of the bast fibers is slow, and at first frag-
mentary ` the pericycles are soon of a size with those in matured
tissue. There is no production of suber in the typical immature
spot; it ends abruptly at the immediate edge of the matured
tissues. With the production of the suber, if the cane is perfect,
except for the immature spot, the bast fibers are produced
normally. Starch is present in the matured tissues, but not in
normal amount, if at all, in the immature spot.

If we now examine in greater detail the modifications occurring
in diseased canes, we will find that the above description applies
only to those canes in which the immature spot occurs solely on
shoots which are not only apparently but also morphologically
matured. These canes are, however, rather the exception than
the rule, and we find more frequently immaturity and morpho-
logically imperfect maturity forming, to macroscopic vision, im-
mature spots and maturity respectively. Furthermore, if one
examines a sufficient number of canes, he will find that the
immature spot is not morphologically constant: it may resemble
more closely a morphologically matured cane than the cane,
macroscopically speaking, on which it is found. I think it best,
therefore, seeing the confusion that is liable to arise in the mind
of the reader, to precede all further remarks by three comprehen-
sive definitions: he will then know exactly what I mean when I
speak of an immature spot; and my dual use of the term cane
will not be ambiguous.

C one (morphologically perfect).—Tissues exterior to the endo-
dermis brown, dead; suber strongly developed; phloem normally
developed; bast fibers never absent.

Cane (morphologically imperfect).—Tissues exterior to the
endodermis brown, dead ; suber well developed ; phloem perfectly.

бысы уы;

OBSERVATIONS ON THE CALIFORNIA VINE DISEASE 135

and imperfectly developed; bast fibers absent more or less over
extended spots.

Immature spot.—Any part of the cane (perfect or imperfect)
in which the cortical tissues, from epidermis to endodermis, still
preserve all the characters of live tissue.

With these definitions in mind, the reader will be better able
to appreciate the extent of the morphological changes that occur,
and I do not pretend to cite them all, in diseased canes.

Immature spot—(a) No suber is laid down in the immature
spot; it stops on one side beneath the pericycle, then jumps above .
it and runs out to the epidermis. Тһе phloem, at this point, con-
tains but one bast fiber bundle; the second phloem (immature
spot), contains but an imperfectly developed one, and the third
(one half the normal size), none. Where the immature spot
begins on the other side we have the following state of things:
the suber ends beneath the pericycle, but has attempted to push
through one end; there is then a break and an attempt to form
suber above the next pericycle. The phloem beneath the peri-
cycle under which the suber stops contains two rows of bast
fiber bundles, but five phloems further on they have completely
disappeared, and the bast itself is about one half its normal size.
(PLATE 5, FIGURE 1.)

(b) The suber stops, as a definite layer, at the edge of the
pericycle, but may continue for a while as a feeble thread. We
find the following striking anomaly in some cases: the suber, upon
arriving at the immature spot, jumps above the pericycle, and the
cortical parenchyma, for a certain distance, intermittently even,
divides in an attempt at suberization. The cell walls of the
phloem are generally discolored, in some cases markedly so, espe-
cially where abnormal suberization of cortical parenchyma has
occurred. Bast fibers are present; three or more bundles may be
observed in each phloem, generally two, sometimes only one.
The bast fiber bundles are always imperfectly developed, m
When three in number. The following condition is sometimes
met with: one phloem will contain three bundles of bast fibers,
the next none and the next two, or the conditions shown in
PLATE 5, FIGURE 3 may be observed.

(c) The suber is always produced. Then

ormal number of bast

186 OBSERVATIONS ON THE CALIFORNIA VINE DISEASE

fiber bundles seems to be produced and the phloem is of normal
size.

(d) The production of suber is scant, and occurs at about the
position normally occupied by the second bast fiber bundle. The
phloem is much reduced in size, and the bast fiber bundles are
obsolete. |

(е) The suber is continuous: it has not always formed immedi-
ately below the pericycle, but frequently several rows of phloem
cells beneath it.

(f) The suber is continuous in the immature spot and lies well
within the phloem. The bast fiber bundles, which are normally
developed at the edge of spot, gradually dwindle down to a
fragment of one, and disappear. Following the decrease in the
number of the bast fiber bundles there occurs a parallel decrease
in the size of the phloem; when the bundles disappear the phloem
is about one half its normal size. (PLATE 5, FIGURE 2.)

Cane.—The canes on which the immature spots I have just
described occurred were all morphologically imperfect. Могрһо-
logically mature tissue would interchange, by gradations, with
morphologically immature tissue, and the immature spots would
form a break in one or the other, or, as it were, the connecting
link between the two. Where the formation of the tissues was
the least perfect the wood fascicles and their corresponding
phloems were much undersized, and no bast fiber bundles were
formed.

І have just shown that in diseased canes the morphological
variations are considerable, and the reader will naturally expect,
as а consequence, considerable variation in the cell pathogno-
monics. These variations, though in themselves interesting, are
not sufficiently important to warrant particular mention, and
I shall, therefore, confine my attention to a general description,
taking for type a section through a cane in which the cells show
considerable disease. Тһе part of the cane most diseased will be,
as a rule, the immature spot.

Dowlen says, in describing the histology of diseased canes,
that in “those canes which have one side ripe and the other side
unripe, the tissues of the ripened portion are almost always wel!
supplied with starch—some starch will always be found—whilst

OBSERVATIONS ON THE CALIFORNIA VINE DISEASE 137

in the unripened portions the tissues will be altogether devoid of
starch. In the discolored areas of the woody bundles, the com-
ponents of the tissues are seen either to have their walls simply
stained brown or else the cell cavity is partially or wholly filled
up with a black brown deposit. . . . The larger ducts and vessels
are often seen to be more or less filled up with thylles, which are
developed sometimes to a great extent."!

These observations are correct. Canes taken from diseased
vines show a paucity of starch, and, when treated with 1 per cent.
iodine solution, give (macroscopically) no starch reaction at all.
In section taken through canes with immature spots, starch will
usually be found under the microscope, generally in the matured
tissues, though, contrary to Dowlen's observations, I have found
it in the immature spots, and in larger quantity when suber is
produced than when it is not. Its entire absence, however, І have
also observed. Тһе presence or absence of starch in the cortex
_ depends, I believe, on the production or nonproduction of the
suber. The presence of starch in the xylem, ligneous medulla
and pith near the protoxylem depends also, to a certain extent,
on the production or nonproduction of the cork—the relation,
however, is not so apparent.

The presence, or absence, of starch also bears а very close
relation to the quantity of brown granuloid, globoidal, or homo-
geneous matter found in the diseased cells. The production of
these homogeneous substances is proportional to the amount of
starch present. Тһе freer the cells from occluding matters, the
freer the sections from starch.

In a cross-section of an immature spot one will observe, suber
being present, the following condition of affairs:

The pith cells encircling the protoxylem are full of starch or
of starch and brown, more or less finely divided matter, which
run together into a
the cell lumen, the
d there as clear
re globoidal, or
t, or encom-

may become coarser, predominant, or even
pseudo-homogeneous mass and entirely fill
starch grains being perceptible only here an
spots. In other cases the brown masses аге mo
fill the space between the starch grains like a cemen

—

Viticultural Commissioners for 1889-

1 Dowlen, E. Report of Board of State
90, бо. 1800.

138 . OBSERVATIONS ON THE CALIFORNIA VINE DISEASE

pass the starch in a brown translucent film. Where the medul-
lary rays begin, and in the secondary wood, this occluding matter
is darker; the starch grains are blackened, or appear to lie in a
blackish matrix, or the lumen of the cells may be filled with a
homogeneous black mass. Sometimes before the medullary rays
reach the cortex the following change takes place: the starch
has largely disappeared from the cells, and the brown irregular
lining of their walls appears to be due to an incrusting of the
primordial utricle. In the cortex the medullary rays present the
following appearance: the lumen of the cells is more or less
filled with yellowish or brown homogeneous matter. The deposits
are darker where the bast fibers have not been produced. Brown
homogeneous matter occludes the greater number of the phloem
cells. The cells in the cortical parenchyma, collenchyma, and
epidermis may also be more or less occluded. Thylloses are gen-
erally present in the primary wood, and are not infrequently very
numerous in the secondary wood; they contain now and then
granuloids. The discoloration of the cell walls in the different
tissues is, except, perhaps, in the neighborhood of the cambial
layer, rather inconstant.

Outside the immature spot the occlusion of the cells is less
pronounced, though of the same general character.

The matter occluding the cells in the diseased canes does not
appear to differ, for the most part, from that filling the cells of
the leaves. Microchemically, the gummous substance in the cells
of the mesophyl does not differ from the occluding matter found
in the cells of diseased canes. This statement, though generally
true, needs to be qualified: the reactions characteristic of the
deposits found in the leaf are more constant in the cortex, and
especially in the phloem, than in the wood, in this latter tissue
a great deal of browning of the primordial utricle seems to be
due to death rather than to particular catabolic changes—hence
the normal appearance of the starch.

Roots.—The roots may be either healthy or show, without
structural modification, to a lesser or greater degree, the same
character of cell occlusion I described as occurring in the cells
of diseased canes, less the accompanying starch when decay js.
evident. |

OBSERVATIONS ON THE CALIFORNIA VINE DISEASE 139

The homogeneous form of the deposits may still persist in
roots in the last stages of decay. Pierce observed in very
decomposed roots corroded homogeneous matter. He says: “a
microscopical examination of the decaying outer parts of the
vascular bundles shows that the cell lumen is filled with a trans-
lucent amber-like deposit. This persists after the decay of the
cell wall, and presents rod-shaped, more or less irregular and
eaten, amber-like casts.’

To sum up our microscopic observations in a few words:
An examination of the various organs of a diseased vine—
leaves, canes, fruit, roots—fails to reveal the presence of any
parasitic organism. In all organs exhibiting signs of disease we
find, however, a polymorphous gummous substance, containing
proteid and tannic matters, that appears to be derived from
starch. In the leaves chloroplastid degeneration precedes the
occlusion of the cell lumen, or is arrested by it.

ПІ

The relationships of the California vine disease

I said before entering on the description of the California
vine disease that the symptoms shown by the various aérial
organs had to be considered in conjunction with one another,
and not singly, if this malady was to be identified with any
certainty. Such a statement as this implies that the symptoms
of the disease are also more or less common to other maladies ;
and this upon examination we find to be the case.

A comprehensive study reveals the interesting fact that the
California vine disease has features in common with the diseases
known as Folletage, Rougeot, Sun-scald, Brunissure, Shelling,
and Tetranychosis. What their common characteristics are І
shall now attempt to show, and this will be best accomplished by
short descriptions of the related maladies. |

Folletage —This disease may affect a vine either in part or in
its entirety, is very rapid in its action, and, as 4 rule, fatal.
Foéx tells us that all the vines in a vineyard are sometimes

ыы ы Е

"1 Pierce, М. В. Loc. cite, 53.

140 OBSERVATIONS ON THE CALIFORNIA VINE DISEASE

affected. Chauzit remarks that he has seen “ vineyards in which
one quarter of the vines were destroyed,"? and Professor Е. T.
Bioletti tells me that it has been known to destroy three quarters
of a vineyard. The symptoms of Folletage vary somewhat
with the rapidity of the attack; if the vines are affected and
killed within the space of a day, the leaves fade, curl, and dry;
but when the attack is less severe the seared leaves will be con-
fined more to the apex of the shoots, the lower leaves being
“much discolored, either with red or yellow spots or stripes.”
The shoots always die from the apex downward and the fruit
withers and dries up more or less according to the degree of its
maturity and the seriousness of the affection on the shoots upon
which it is borne.

The anatomy of Folletage does not appear to differ greatly
from that of Brunissure.

Folletage generally occurs only at midsummer, but may affect
vines as early as May.

The accredited cause of the malady is a rupture of equilibrium
between absorption and transpiration.

Rougeot* is a mild form of Folletage and Pierce says the
following description would apply to leaves of vines affected by
this malady:

“Тһе leaves of the dark varieties of grapes show a red dis-
coloration between the veins and at the margin. In the earlier
stages this color is faint, but later on the tissue lying between
the main veins becomes bright red, and still later dies and changes
to dull brown. Тһе death of the leaf usually begins at the mar-
gin, or in the center of the red stripes lying between the veins,
or it may involve both regions at once. The venation of the leaf
remains green in most instances, forming a symmetrical green
vem system after nearly all the intervening tissue is dead, of
has turned red or brown. Thus there are in these later stages
three distinct gradations of color in the affected leaves: (!
A brown and more or less dried margin, or bands of brown lying
between the main veins, or both; (2) a band of bright red bor-
dering the dead brown portion of the leaf; ( 3) normal gu

: Foéx, с. Cours complet de viticulture, 573. [ed. 4].

кыруы В. Revue de Viticulture 26: 50. 1006.
Pierce, N. B. Loc. cit., 106;

is Rougeot of authors pro parte.

HR
Е

OBSERVATIONS ON THE CALIFORNIA VINE DISEASE 141

tissues outlining the main venation of the leaf. All colors vary
according to the time since the first alteration took place. The
petiole is not involved at once in any evident change, but later
the leaf is cut off. A second variety of grape had leaves altered
in a somewhat similar manner to those of the dark varieties de-
scribed, but the bright colors did not prevail. There was little
to be seen of a third color on these leaves. The alteration
is almost directly from the normal green to a dull muddy brown,
as if the base colors were yellow and black. The dead tissue
occurs first at the margin, and in spots and stripes between
the main veins, rarely if ever touching a large vein. Between
this dead tissue and the green next the veins is sometimes a
slight transitional shade of yellow, which is nearly wanting in
many cases, the brown being directly joined to the green on
either side of the main veins. Where the intermediate yellow
line is wanting, the appearance of the leaf is very striking,
and differs in color from any diseased varieties noticed in
California. The pattern of the markings is, however, the same.
The difference observable is a varietal one. The leaves of a
variety of white grape were altered in the manner described for
the Muscat of Alexandria in California. In the early stages the
changes of the leaf are foreshadowed in faint yellowish spots im
the parenchyma, which become more pronounced as the trouble
advances. At this time the leaf may have a yellow speckled
appearance. The spots are yet somewhat cloud-like and illy de-
fined, and are rarely located upon a vein. As the discoloration
becomes more marked these cloud-like spots are better defined at
their margin and more and more of the parenchyma of the leaf
between the veins becomes involved. As the light yellow spots
enlarge the parenchyma at their center turns reddish brown and
dies. Later there is a brown central stripe between the veins
and at the margin of the leaf, and bordering this dead tissue 15 4
line of half-dead yellow tissue lying next the green bordering
the veins. All these markings are very distinct and well defined
in the later stages of the trouble. As the death of the tissue
between the veins progresses it gives to the green bands at the
veins the symmetrical appearance seen on the Muscat leaf in
California,"

Rougeot has been ascribed to the same cause as Folletage.

Sun-scald.— This malady is described as follows by Viala:

somewhat depressed
mes affected periph-

“ ^ А
Sun-scald appears as irregular intervenar,
feuille-morte maculations. Leaves are зотей

* Pierce, М. В. Loc. cit., 186.

142 OBSERVATIONS ON THE CALIFORNIA VINE DISEASE

erally: in these cases the discoloration gradually works inwardly
towards the petiole, the dead parenchyma assuming a dirty yel-
low or light brown color; in other leaves again the diseased
tissues form sinuate intervenar bands extending from the petiole
to the edge of the blade. The hairs of tomentose varieties are

tufts of powdery mildew to have sometimes been taken for them.
In some instances the leaves show small, more or less brownish
Брега. я

Іп severe cases Sun-scald gradates into Folletage.

Brunissure.—" Тһе disease first appears on the upper sur-
face of the leaves in the form of very small, very numerous
yellowish brown spots, in the case of the varieties of the vine
bearing white fruit, and as brown almost black punctuations, in
the case of those varieties bearing colored fruit. As these spots
are all very near one another, for they are separated only by the
ultimate ramifications of the fibro-vascular bundles, they run
together almost from the day of their inception. After coa-
lescence has taken place they form yellowish brown or dark
brown areas that cover the leaf-blades to a greater or less extent.
Some cover only the space of half an inch, while others cover 4
quarter, one-third, the half, and sometimes even the whole of
the leaf

“These maculations appear indifferently here and there upon
the blade of the leaf, now between the veins, now upon the tissues
adjacent to the veins, and across the latter; now along the edges
of the leaf, now at the center of the blade. In general they
form between the veinlets, encroaching upon the main veins and
the circumjacent tissues later."?

All the leaves do not become diseased at once. The basal
leaves are the first to become affected, and the apical leaves, even
when the shoots have ceased growing, are the last to become
diseased ; they may even, in mild cases, remain entirely healthy:

Brunissure has been studied by Viala and Sauvageau, Debray,
Prunet, Ducomet, and Ravaz.

Viala describes the appearance of diseased cells as follows:

ene D We 3
Bee: P. Les maladies de la vigne, 470 et seq. 1893 Ге. 31. [Translation
Ravaz, L. La Brunissure de la vigne. Ann. École Nat. Agric. Mon m.

II. 3: 175 et seg. 1904. [Translation.]

ИИ

d

OBSERVATIONS ON THE CALIFORNIA VINE DISEASE 143

“Tn the first stages of the disease the parasite develops more
especially in the palisade tissue; it invades the cells of the lacu-
nose tissue later, but is found only exceptionally in the epidermal
cells. One observes in sections taken through recently affected
tissue that healthy cells may contain starch in goodly amount,
but that in the cells that are being invaded, it is much less abun-
dant. The starch completely disappears with the spread of the
parasite throughout the cell. . . .

* Wherever the leaf blade is brown the tissues are infested,

worst affected tissues, however, the infection is general; all the
cells of the mesophyl are filled with the plasmodium.

plasmodium presents the appearance of a sponge. At other times
it lines the cell walls to a greater or less extent; this parietal
plasmodium is more or less finely vacuolate and may be compared
to fine lace-work. Protoplasmic strands sometimes join the
various parts of the plasmodium and may even anastomose in a

geneous, refringent, and oil-like; others that contain a large
central or more or less excentric vacuole; lastly, others that are
finely vacuolate and apparently composed of spongoid proto-
plasmic matter,"

What is the cause of Brunissure? Viala and Sauvageau be-
lieved that the vacuolate matter was a plasmodium, Plasmodio-
bhora Vitis, and the globoids, homogeneous and granular matters,
Products of decomposition. Debray considered that the globoids
and the homogeneous (cereous) matter as well and the vacuolate,
plasmodium-like masses were one and all phases in the develop-
ment of an organism, Pseudocommis Vitis. Prunet as à result

dui E

ооо ы
'Viala, P. Loc. cit., доз et seq. [ Translation.]

144 OBSERVATIONS ON THE CALIFORNIA VINE DISEASE

of his investigations determined that Brunissure was caused
by a fungus which he referred to the genus Cladochytrium
Nowakowski. Ducomet, on the other hand, concluded that the
vacuolate, plasmodium-like bodies were degenerate chloroplasts,
and the globoids, granular and homogeneous matters, products
of decomposition, and his results were confirmed later by Ravaz.
Both Ducomet and Ravaz claimed that Brunissure was a physi-
ological disease; they did not agree, however, as to its cause.
Ducomet believed that the disease was induced by rapid changes
of temperature, a sudden rise or fall of the thermometer being
causal. Ravaz argued that the malady was due to over-produc-
tion, and, in support of his thesis gave a great deal of data;
he further strengthened his opinion by asserting that he could
produce Brunissure at will.

Shelling.—In this disease “ the leaves at the outer extremities
of the shoots first show a yellow discoloration which follows
more or less continuously the outer margin. . . ." This “ yel-
low portion dies and turns brown," and the leaf, as a conse-
quence, curls at the edges. In older leaves and young leaves in
which foliar development is exceedingly slow the following char-
acters will be observed: “ Small irregular blotches of a dark color
appear between the veins, these enlarge rapidly, . . . and coalesce
to fill up the space between the veins which remain green or yel-
low. These changes occur so rapidly that the foliage seems 10
change color suddenly. The contrast between the green or light
yellow veins and dark purplish brown of the intervening tissues
gives a peculiar streaked appearance to the leaves. In the most
serious cases they curl up, become dry and brittle, and finally
drop from the vine, leaving it nearly bare.”? The most striking
feature of the disease is, however, the fall of the berries from the
pedicels.?

The ва а —
Lodeman, E. G. Some ра troubles of. Western Kë у York. Corne
Univ. Agric. Exp. Sta. Bull. 76
oe D. G. Diseases of zi grape in Western New York. Jour. МУС
891.

OBSERVATIONS ON THE CALIFORNIA VINE DISEASE 145

“As the season of ripening approaches, certain berries of the
affected clusters fall to the ground on account of the inability
of the main fibers and other connecting tissue of the fruit-stems
to sustain their weight. . . . The portions of the clusters first
affected are, so far as my observations go, invariably either the
lower extremity of the cluster as it hangs from the cane, or, in
the case of heavily shouldered clusters, the outer extremity of
the stem forming the shoulder. . . . Sometimes only one or two
berries may fall, but in other cases the drying and shriveling of
the stem gradually extends upwards, the affected portion being
plainly marked by the absence of the berries. . . . It often occurs
that not a berry remains hanging upon the bunch. .. . Some
clusters upon a vine seem to be more free from shelling than
others upon the same plant.’

The development of shelling is sporadic. Тһе disease affects
vines at the height of summer and is as selective as Folletage or
Rougeot. One, several, or all the shoots may be affected.

“It very commonly occurs that plants in certain portions of a
vineyard shell, while the large majority of them do not. | Тһе
line is sometimes so sharply drawn,” our authority continues,
“that the affected plant may be entirely surrounded by healthy
vines; and it is not uncommon to find a healthy vine in the midst
of those which shell. . . . Some clusters upon a vine seem to be
more free from shelling than others upon the same plant. This
seems to be due, in many cases, to its location upon the cane, but
there are so many exceptions that no definite rule can be laid

Wn. ... Another peculiarity which may sometimes be seen,
although cases of it are very rare, is the shelling of the berries
upon only one portion of the vine, as for instance those borne
upon the canes which spring from an arm, the difficulty thus
affecting only one half of the plant. One case was noticed in
which the clusters found upon one cane were the only ones which
suffered, amongst all those borne by the vine.”

The cause of Shelling is not definitely known, though the
Weight of the evidence points to defective nutrition.

Tetranychosis?—Young vigorously growing leaves become
convex and paler than normal; they may even become somewhat

maturity and the Viniferae are not entirely free from the trouble. Certain
Franco Х Americans behave similarly.

*Lodeman, E. С. Loc. cit., 413-415.

*Lodeman, Е. С. Loc. сй., 415.

* Maladie rouge.

146 OBSERVATIONS ON THE CALIFORNIA VINE DISEASE

spotted with dead tissue and dry up peripherally. Leaves that
have ceased rapid growth or are already fully developed become
yellowish green and more or less covered with maculations
between the veins. The spots spread over the intervenium, not
so much from individual development as from increase of
centers. The centers of the maculations die, but a running to-
gether of the dead areas to form stripes does not seem to pre-
cede the death of the whole surrounding parenchyma. As a
rule the leaves die from the edges inwards, and in severe cases
maturation of the shoots is interfered with; the canes appear
spotted or striped with immature tissue. The anatomy of patho-
gnomonic leaves may be briefly described as follows: |

A deposit, in the epidermal cells of the upper surface and, to a
less extent, of the lower surface, occurs in the most diseased areas
only. It is homogeneous to all intents and purposes, and rapidly
thins out in the healthier cells. In the palisade tissue, where.
the cells have not collapsed, they are either full of a homogeneous
deposit or with granular matter. The chloroplasts are small,
vacuolate, and scarce. The cells of the lacunose tissue in the row
next the palisade layer, a certain number of them at least, con-
tain disintegrated chloroplasts, but rarely globuloids. The re-
maining cells contain disintegrated chloroplasts as well as granu-
lar or homogeneous matter. The vessels of the small fibro-vas- |
cular bundles are occluded, more or less, with granular Of
homogeneous matter. The microchemical reactions of the de
posits are the same as in the case of the California vine disease.
| The disease is caused by Tetranychus Vitist and may be held
. In check by sulphur or the polysulphides.

We may conclude from the study of the relationships of the
California vine disease that this malady, while having шапу
ous of resemblance with other diseases, is, in diagnosing
Specimens, likely to be confounded only with Folletage, Tetra- |
nychosis or Sun-scald. I have just shown that all these disease — |

have at least one salient character that differentiates them from —

the California vine disease, and these characters are clearly.

eg + Ў

Мг. N. Banks, to whom I sent specimens for identification, could ва: e
ға from the material at hand, whether this Tetranychus was T. Vitis я
а n of T. telarius, but was of the opinion that it was T. Vitis Boisd :

OBSERVATIONS ON THE CALIFORNIA VINE DISEASE 147

observed on sufficiently fresh material that includes both the
apical and basal portions of the shoots.

IV

Nature of the California vine disease

When empiricists agree, their opinion may be taken as having
some foundation in fact. The grape-growers of California have
always considered the California vine disease as a “ top disease,”
and the facts undoubtedly support this opinion.

Nearly every observer who has written upon the California
vine disease has dwelt at length upon the foliar characteristics of
this malady, and either stated implicitly or expressly that it travels
downwards, not upwards. Morse observes that “ scattering vines
which appear not to have made good growth last year, are dried
up and dead to the roots, which in nearly all cases still contain
sap." Dowlen is of the opinion that the disease travels down-
wards. “The disease always travels downwards,” he says,
“both in vines and cuttings.” “Some cuttings were purposely
planted in an inverted position, still the result was the same; the
disease always started at the end which was naturally farthest
from the main stem, whether that end was placed in the air or
in the soil"? That the disease does not affect the roots is ап
opinion that has been largely held by viticulturists, if we may
judge from a letter published in the Pacific Rural Press of
October 20, 1888.3 “Тһе published accounts of the disease which
I have seen," writes Scribner, “ assert that the roots are perfectly
sound." But this is not exactly his opinion, for in the very next
sentence we find him saying—" In every case examined by Pro-
fessor Viala and myself we found the ultimate rootlets dead
often for a foot or more from their tips." Scribner does not |
state, however, whether the vines examined were in the first or
last stages of the disease, but I am inclined to believe, from the

ing the seasons 1885 and

' Morse, F. W. Report of the viticultural work dur
1886, College of Agriculture, University of California, 177. | 1886. Ca
Dowlen. Report of Board of State Viticultural Commissioners for 1889

90,

Cal.

1890.
3 Letter of Е. L. Scribner to Benj. Pratt, of Orange,

148 OBSERVATIONS ON THE CALIFORNIA VINE DISEASE

observations of Pierce, that they were in the latter, or, at least,
іп an advanced stage; for this writer says: “ The time when the
roots become diseased is difficult to ascertain, and will probably
not be known before the nature of the malady is determined.
. . . At present I incline to the opinion that the extremes of the
vine show the early signs of disease at nearly the same time.”
Viala and Sauvageau observe that “the disease gradually de-
scends towards the base of the shoots, becomes manifest in the
arms, the trunk, and later reaches the roots."?

My personal observations are in perfect accord with those of
the authorities just cited. I have observed that the roots of
diseased vines are not affected at the inception of the disease and
will take the longer to show signs of weakness the healthier the
vine, and the greater the amount of foliage still remaining in
normal function. ТЕ is evident that when a vine is affected in all
its shoots with a severe apoplectic attack, disorders in the finer
roots will immediately take place; the sudden die-back of the
shoots almost to the spurs will be followed by corresponding
death in the rootlets—but if we take a vine that is diseased only
in a few of its shoots, then its roots will not be different in ap-
pearance from those of the neighboring healthy vines. Visible
symptoms of disease in the roots do not precede the appearance
of disease in the shoots. When the roots decay it is because
they are in a weakened state and external conditions are ѕирег-
inducive; this is evidenced by the fact that roots will dry-rot in
one soil and soft-rot in another.

The visible seat of the disease being in the foliage, then, ue
popular opinion vouchsafes, and our own and other investigators
observations substantiate, it will be in the study of the behavior
of the diseased vines and of the anatomy of pathognomonic
tissues that we will find the answer to the question: What is the
nature of the California vine disease?

I shall attempt to answer this question.

Шш the preceding chapter I established that the California vine
disease does not differ in method of attack or propagation from
Folletage, and showed also that the foliar characteristics of the

* Loc, cit., 51.
* Loc. cit., 99.

OBSERVATIONS ON THE CALIFORNIA VINE DISEASE 149

two diseases are in a large measure similar. The importance of
this similarity is enhanced when the anatomical study of patho-
gnomonic tissues in the former malady fails to reveal the presence
of any foreign organisms, but simply shows a chloroplastid de-
generation inversely proportional to the rapidity of death, and an
occluding of the cell lumen by a gummous product probably due
to starch decomposition. When we also consider that Roze
is credited by Debray! with having found Pseudocommis Vitis
(Syn. Plasmodiophora Vitis) in leaves taken from vines attacked
by Folletage it can hardly be denied that the two diseases are very
closely related, for this supposed organism is the cause of
Brunissure, which malady I have shown to be undifferentiable
anatomically from the California vine disease. Furthermore, we
know that sun-scald may gradate into Folletage and, a fact also
of capital importance, that this malady is prevalent in southern
California—the home of the California vine disease.

The California vine disease, Folletage, Sun-scald, and Brunis-
sure are then very closely related. In fact, a close and compre-
hensive study of the anatomical features ef these diseases cannot
help but lead one to the conclusion that all four are due to the
some functional state in the vines themselves, outwardly indi-
cated, owing to the play of external agencies, by somewhat differ-
ent symptoms. This common functional state I shall call a lower-
ing of functional activity; and for the following reason: In the
California vine disease—one might say without grave inaccuracy
in all four diseases—the anatomical changes observed follow very
closely those occurring in autumn leaves, preeminently is this the
case when the sequence of changes is slow. Now it is well
known that the changes brought about in the leaves of deciduous
plants in autumn are due to a lowering of functional activity.
The rest these plants require after a certain period of growth
takes place normally at the end of such a period, provided condi-
tions are not conducive to its prolongation, without the determi-
nant intervention of external agencies. The róle of external
agencies is largely of secondary importance; they hasten or retard
autumnal changes in the leaves, but they will not bring them

'Debray, А. Bull. Soc. Bot. France 45: 256. 1898.

150 OBSERVATIONS ON THE CALIFORNIA VINE DISEASE '

about unless the plant has reached the proper stage of develop-
ment. But once the plants have reached the required stage in
their development then external agencies may hasten or retard
autumnal changes in the foliage. In countries with a cold winter
climate it is a matter of common knowledge that the weather
influences very largely the brilliancy of autumn tints, their period
of duration and time of development, but is not, I repeat once
more, the determinant factor: the plant must first have reached
а certain stage of functional activity.

The changes occurring in the autumn leaves of vines, vacuola-
tion and absorption of chloroplasts with, in cases, a slight pro-
duction of globoidal and homogeneous matters—are similar to
those observed in the California vine disease; one might say that
they represent a diseased area of a leaf affected by the latter
malady in a very mild form. If, therefore, in certain of its forms
the California vine disease becomes microscopically identical with
an autumn leaf it necessarily follows that Brunissure, Folletage,
and Sun-scald are also related to it. In autumn leaves, then, and
in the above maladies the same cause must be paramountly active.
And as the changes observed in autumn leaves are due to à
decrease in functional activity, the disease we are considering,
i. е, the California vine disease, may be said to be due to the
same cause. But the same functional inactivity need not neces
sarily be operative in all cases:

In autumn leaves the changes are due to a decrease of vegeta-
tive activity; in the case of Brunissure, to overbearing, 25
appears from the researches of Ravaz; in the case of Folletage
Sun-scald, and the California vine disease, to a rupture of equi-
librium between absorption and transpiration operating upon
vines weak in their power of absorbing and translocating water,
and brought about by external agencies favoring transpiration.

That it is really to a weakened state of the vine that the
characteristics of the disease above mentioned are due, may be
deduced from the conditions favoring their development. To
consider, however, only the California vine disease, these condi-
tions taken individually could not be held responsible for its `
development, but when considered as factors favoring the visual

OBSERVATIONS ON THE CALIFORNIA VINE DISEASE 151

manifestation of an organic weakness, they are worthy of
consideration.

The California vine disease develops with greater intensity in
sunshine than in shade, in wind-blown vineyards to windward
rather than to leeward, and soil fertility and texture are not
without influence. I shall examine more at length these factors,
and independently of one another, though in reality, they
cooperate more or less.

Shade.—The effect of shade on the development of Ше
disease is marked. Pierce notes that vines well shaded are but
slowly killed; Ша“ shade has a marked retarding influence upon
the work of the disease." My own observations confirm this
view. Vines shaded at the time the rupture of equilibium be-
tween transpiration and absorption is brought about in the open
vineyard, whether by sudden insolation or insolation plus
humidity or wind, are not affected except in extreme cases.

Insolation.—The effect of excessive insolation has been ob-
served by Morse, who remarks that “ many successive vines could
be found with dead spurs of last year upon the sunny side, and
not infrequently a line of dead wood extended with the fiber to
or near the surface of the ground. No shoots started from this
side. Suckering, if it occurred at all, came invariably from the
north side, where the greenest wood was always found; in ас,
I saw no line of dead wood upon this side.”

The deleterious effect of intense sunshine following a deposi-
tion of moisture is well known to all horticulturists. Morse
thinks that water of condensation is not without effect in the
burning of the foliage in some cases of the California vine
disease. He writes: Those vines “which are protected more ог
less by trees, present a scalded appearance; some leaves show
three different stages: about the margin, and extending an inch
or so outward, they will be perfectly red and dead; next comes
a zone of light green color, followed by another only slightly
lighter colored than the healthy part of the leaf. These are
usually most exposed to the sun. In the early morning large
drops of moisture, almost equal to that from a heavy rain, are

1 е
Loc. cit., 111.

* Loc. cit., 177.

152 OBSERVATIONS ON THE CALIFORNIA VINE DISEASE

found upon these vines, and it is probable that the hot sun scalds
the leaves before the water is evaporated."

“Tt is quite noticeable that leaves exactly similar to those found
injured in the open vineyard may also be found among the
scalded ones.”

Wind.—The effect of wind on the development of the disease I
have been able to follow closely. I have observed that the disease
appears under the following climatic conditions: When hot, still
mornings are followed by stiff breezes, the disease may be ех-
pected to appear, and if one walks through a vineyard that has
become affected from this cause, with his eye to the wind, he will
observe less disease than if he walk before the wind. The vines
are affected to windward, which would be expected were trans-
piration difficulties the cause of the disease.

Soil texture and fertility.—Pierce observes that the rapidity
with which vines succumb to the California vine disease depends
upon the physical condition of the soil. Dividing the soils of the
state into (1) “Heavy soils, including the red and black adobe
and clay soils; (2) the gravelly soils; (3) the fine loose soils,
including the sandy loams and the sands and fine sedimentary

deposits of the river bottoms,"? he finds that “If conditions.

of age and variety are the same, the power of any vine to
resist disease is about as follows upon the three classes of soils :
(1) Least resistance upon coarse gravelly soils; (2) medium
resistance upon soils of a heavy and compact nature; (3) greatest
resistance upon level soils which are loose and sandy but not
infertile."*

The róle of soil texture on the development of the California
vine disease I have been able to follow particularly well in one
instance. Іп a vineyard already old and subject to the daily blast
of the trade wind, I found that the disease first appeared where
the soil was heaviest, developing later where an admixture of
sand and fine gravel made it more open and penetrable, and this

despite the fact that the free moisture was approximately the.

same in both cases.

nts ete irit

EE а 0

Morse, Е. W. Loc. cit., 1595.
2 Loc. cil., 96.
* Loc. cit., 98.

1
і
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E
;
д
3
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3
$

OBSERVATIONS ON THE CALIFORNIA VINE DISEASE 153

ConcLuston.—The facts gathered during the course of this
paper point to the conclusion that the California vine disease is
due to some weakness in the functions of absorption and translo-
cation of water becoming manifest when conditions favoring
transpiration are marked. To say that the disease is due to a
rupture of equilibrium between absorption and transpiration does
not conflict with any recorded observations. This is true whether
one considers the disease from the point of view of the effect of
external agencies upon its general development, or from the point
of view of its development upon individual vines, or from the
point of view of its symptomatology, anatomy, and relationships.

CORNELL UNIVERSITY,

ITHACA, NEw YORK.

Explanation of plates 1-5

PLATE 1
1,2, 3, 4. Young diseased leaves of Vitis vinifera, var. Missi
eaf of V. vinifera, var. Muscat of Alexandria, showing ыы stria-
tions and death of tissues at edge of blade.

PLATE 2
Leaves showing various stages of disease.
1,2, 3." V. vinifera, var. Mission.
4 5. Г. vinifera, var. Berger.
PLATE 3
1, 2, 3, 4, 6. Sections through palisade tissue of leaves in various stages of
disease,

5. Cross-section of a small fibro-vascular bundle.

PLATE 4
. Cross-section of part of а fibro-vascular bundle of a main vein showing
development of thylloses in the vessels. 4
Section of healthy leaf showing normal appearance of chloroplasts e
3,4,5. Sections through palisade tissue showing various stages of diseas

PLATE 5

1, 2, 3. Cross-sections of diseased cane nicis
B, phloem; C, cortex; F, bast fiber WE М, medulla; P, регісу

fibers; К, Фейн rays; 5, suber; X, xylem

Мем. TORREY CLUB VOLUME 14, PI :
UME 14, PLATE'I

DISEASE

BUTLER: CALIFORNIA VINE

Mem. TORREY CLUB VOLUME 14, PLATE 2

ASE

BUTLER: CALIFORNIA VINE DISE

MEM. TORREY CLUB VOLUME 14, PLATE 3

VOLUME 14, PLATE 4

Mem. TORREY CLUB

el

SN =
152

du

BUTLER: CALIFORNIA VINE DISEASE

Мем. ToRREY CLUB

DISEASE

BUTLER: CALIFORNIA VINE

VOLUME 14, PLATE §

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a

A monograph of the genera Chaetomium and Ascotricha* -
А. H. CHIVERS
(WITH PLATES 6-17)
INTRODUCTION
In 1902 the writer attempted to arrange and classify a con-
siderable number of specimens of Chaetomium which were then
in his herbarium at Hanover, New Hampshire, and which were
collected by him in various localities in New England. At that
time the only available monograph was the well-known work of
Zopf (113) which contained descriptions of ten species only.
While it was possible to identify certain forms in the collection,
others were laid aside until more information could be obtained.
The study of these first collections, however, aroused so much
interest that the writer was led to continue his work on these
fungi, and since that’ time he has been engaged in the preparation
of an illustrated monograph of the genus.

It became evident from further examination of the literature
that no adequate work on the genus was available, and that from
а systematic standpoint it had become greatly confused owing to
the multiplication of species which are either synonyms, in many
Cases even of species not belonging to this genus, ог are described
without recognizable figures or full and comparative descriptions.

Some time after, when this work was well under way, a mono-
graph of the genus by Bainier (3) appeared, in which twenty-
two species and three varieties were described and illustrated,
some of which proved to be American, while twelve species
and two varieties were described as new. This monograph,
although in some respects more comprehensive, was nevertheless
like that of Zopf by no means complete. No mention was made
of work by American authors or with two exceptions of English
literature on the subject, while the repeated use for new species
of names already preoccupied introduced a further element of
confusion.

* Contributions from the Cryptogamic Laboratory of Harvard E rd
ss

No. LXXVI, Memoirs of the Torrey Botanical Club, Volume 14, No. 3-
June то, 1915.
155

156 MONOGRAPH ОЕ CHAETOMIUM AND ASCOTRICHA

Up to the present time there has been no further attempt too
make a comprehensive review of the genus or to collate the
American forms, with the exception of the revision of the Chaeto-
miaceae by Miss Palliser in the North American Flora, where
seventeen species are enumerated including three unpublished
names. |

The writer feels therefore that а thorough and complete
study of the subject is much to be desired, and while he cannot
hope that the following revision can be final, he has spared no
pains to ascertain the facts in every case so far as has been possible,
and to make clear by figures and descriptions the specific char-
acteristics of the individual species which in his opinion should be
recognized.

The postponement of the final publication for a considerable
time on account of unavoidable interruptions and delay caused
by the preparation of plates seems in the end an advantage,
for in the course of his work upon these widely distributed fungi, E
the writer has been able to examine a very large series of specimens.
from various herbaria and exsiccati, and to cultivate many species - SS
from diverse sources on various media, over long periods of time |
and through many successive generations. Аз а result of this '
examination, numerous forms һауе been added to those previously Se
recorded from America, and a number of new species have been-
recognized. In this connection it may be mentioned that all of X
these forms, with six exceptions, have been extensively cultivated 52
in а pure condition and that it has been possible to determine with E
accuracy their range of variation as well as their salient зресшс 22 |
characteristics. 71 m

Those who have given attention to species of Chaefomum — |
must be aware of the more or less unsatisfactory results 19 be SE
obtained by working with dry herbarium material. Unless the
specimens are carefully protected by pill boxes or other convenient
receptacles, the characteristic appendages of the perithecia Ei
become broken, and the thin, brittle, perithecial walls disintegrate.
It is the exception to find herbarium and exsiccati material
protected. Оп the contrary it is most often mounted on "`
Page with no protection whatsoever, or enclosed in paper envelope
Which soon collapse and press against the plants, and it is

ма ES

MoNoGRAPH OF CHAETOMIUM AND ASCOTRICHA 157

these reasons that specimens are so often broken and damaged.
In many cases, therefore, it is unfortunate that new species names
have been freely applied to such forms as have seemed to differ
from those already named, before it was determined with cer-
tainty that such differences as existed were characteristic of the
living plant and not peculiar to the dry specimen. Moreover,
since all species of Chaetomium which have come to the attention
of the writer lend themselves to easy culture on various media,
it would seem highly desirable that they should be carefully
studied in all stages of development in order that their true
characteristics may be determined.

As a result of a thorough review of the literature one hundred
and fourteen species names and fourteen variety names have
been found which have been applied to forms supposedly Chae-
tomia. In the present revision a considerable number of changes
have been made. Names which were first used in connection
with good species have been retained, while many others have
been listed as synonyms thereto. For reasons given elsewhere, a
considerable number have been excluded, and the writer feels*that
the use of such names should be discontinued. Certain new
names representing new forms heretofore unrecognized have been
added to the list of known species. Nevertheless according to
the best judgment of the writer, the genus should include not
more than twenty-eight species.

А careful study of the genus Bommerella has led to its in-
clusion under Chaetomium, while on the other hand Ch. chartarum
(Berk.) Winter and Ch. pusillum E. & E., which possess similar
characteristics, yet are markedly different from those of the typical
species of Chaetomium, have been included in a separate genus
to which the name A scotricha, first given by Berkeley to the plants
which Winter later renamed Ch. chartarum, has been apphed.

In connection with his work the writer has been especially
fortunate in being able to examine exsiccati in several herbaria
and to study a large number of type specimens which have been
generously contributed by many individuals both in this country
and abroad. Sets of specimens as complete as possible and
containing as many of the writer's forms as are available have
been distributed to the Royal Botanic Gardens at Kew, the

158 MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

Cryptogamic Herbarium at Berlin, the University of Padua,
the New York Botanical Garden, and to the Cryptogamic
Herbarium at Harvard University. Cultures of several species
have been sent to the Centralstelle fiir Pilzkulturen at Amsterdam.
At this point the writer wishes to acknowledge his indebted-
ness to those who have aided substantially in the completion of
this work; to Dr. Roland Thaxter, especially, who has at all
times been ready to encourage and give freely of his time and
material, and whose helpful suggestions and guidance have made
this work possible; to Professor W. G. Farlow for the use of his
herbarium and literature from his private library; to Professor
С. Е. Atkinson, Dr. J. H. Faull, Dr. L. W. Riddle, Dr. I. F.
Lewis, Dr. A. F. Blakeslee, Dr. George R. Lyman, Mr. R. H.
Colley, and Mr. А. Т. Speare, for material for cultures; to Dr. Р.
А. Saccardo, Mr. George Massee, Dr. С. Lindau, Dr. Charles Н.
Peck, Dr. J. Dearness, Dr. P. Hariot, Dr. F. Cavara, Dr. C. H.
Kauffman, Dr. L. H. Pammel, and Dr. Elam Bartholomew, for
valuable type material and authentic specimens; to Mr. A. B.
Seymour and to Mr. Piguet for assistance in reference work.
This investigation was begun in the Cryptogamic laboratories
of Harvard University under the guidance of Dr. Roland Thaxter,
and completed in the botanical laboratories of Dartmouth College.

THE GENUS CHAETOMIUM, HISTORICAL REVIEW

The contributions of a systematic nature which have been
made to the genus Chaetomium are numerous and cover a con-
siderable period dating back to 1817. In the brief summary
which follows it will be possible to discuss only those which are
most important and to call attention іп a very general way to а
host of minor contributions which have been made by writers
little acquainted with the characteristics of the forms in this
genus.

In 1817 Gottfried Kunze (50) published a description of a
hitherto unrecognized genus and gave to it the name Chaetomium
(хойтира, a hair). It is of interest to note his characterization
of the genus, since he clearly discerned some features which
many later writers have disregarded, while on the other hand, he
made errors in describing structures which later writers have

MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA 159

corrected. According to his description the perithecia were
globose, membranaceous, clothed on all sides with opaque hairs,
and at length became pierced by an opening at the summit.
The spores were pellucid and mingled in a gelatinous mass. The
fungus appeared like an inverted Муготесит of Tode. The
spherical or steeple-shaped fruiting body consisted of a cuticular
integument, trimmed with long, mostly stiff opaque hairs envelop-
ing a more or less globular, gelatinous mass in which spores were
mixed. Under the influence of moisture the semi-transparent
enclosing walls opened by a central pore and drew back more or
less. The wall closed as it became dry and reopened with re-
turning moisture. This happened as long as the perithecium
contained spores.

It may be seen from the above description that Kunze saw
clearly an important fact, namely that the perithecium possessed
a pore through which the spores were discharged, but that on
the other hand he failed to interpret aright its internal structure.
It is not difficult for one acquainted with the appearance of the
structure within the perithecial wall to understand how, with
comparatively slight magnification, he mistook these structures
for a mass of jelly and failed to discover the asci of which the
mass is made up. Hence the fact that asci were present in
these plants was entirely overlooked.

Ch. globosum Kze. (50) was named and described as charac-
teristic and typical of the genus. According to his own state-
ment Kunze was familiar with several other species of Chaetomium,
and it was his intention to publish a monograph at a later date,
but with the exception of a description of an interesting species
under the name Ch. elatum Kze. (51) this author published
nothing further on Chaetomium. :

In spite of the fact that not less than thirteen new species
Were described by Ehrenberg (28), Fries (38), Wallroth (109),
and Schweinitz (91) during the years immediately following
Kunze's work, no contribution of value was made until 1837.
Writers up to this time had not discovered the presence of asci
nor had they considered the size of spores an aid to the identi-
fication of species. In many diagnoses of new forms the fact
that the genus had been characterized by Kunze as possessing
_ ап ostiole seems to have been entirely overlooked.

160 MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

With the publication of his Icones by Corda (21, 22, 23) in 1837,
1838 and 1840 came quite a new era in the history of the genus.
This author amended the original description given by Kunze as
follows: ‘‘Peridium membranaceous, at length opening at the
apex by a pore, clothed on the outside with opaque hairs, sup-
ported by a more or less well-developed, fibrous hypothallus:
spores grouped together, ascomorphic, pedicellate, at length
discharged as simple powdery spores. Pedicels without mucous.”
It may be readily seen that from the amended description one is
able to form a much more accurate judgment of the characteristics
of the genus.

Corda was the first to study the internal structure which other
authors had been content to call a gelatinous mass. He observed
this to be made up of bodies which he called asci, though according
to his description he was misled in believing that these bodies
functioned as pedicels on which the spores were borne. Corda’s
descriptions are for the most part illustrated by figures which
are elaborate for his time, and while it is not possible to deter-
mine with certainty all of the forms with which he dealt, several
may be easily recognized. Of the seven new species described
Ch. indicum and Ch. murorum stand out as those to be retained,
while the remaining five have been referred to other genera,
treated as synonyms, or excluded.

Between the time of Corda and the appearance of Zopf's
monograph in 1881 descriptions of at least thirty-two new species
appeared. In fact so many contributions were made that it will
be possible to mention here only those which furthered the ac-
curate knowledge of the genus. In 1849 Fries (39) called attention
to the fact that in Chaetomium the spores are formed in typica
asci, though the asci are rarely conspicuous. This is the first
recognition of the true character of the perithecial contents which
Corda had mistaken for spore-bearing pedicels and which all-
other writers had described as a gelatinous mass including spores
Fuckel (42) and Cook (16) also made valuable contributions in
1869 and 1873, respectively, in their descriptions of two new species
under the names Ch. crispatum and Ch. funicolum.

The well-known monograph of Zopf (113), to which reference ` —
has been made on a preceding page, marked the appearance of а

MONOGRAPH ОЕ CHAETOMIUM AND ASCOTRICHA 161

work of an entirely different nature from that of many preceding
authors, and it is only to be regretted that the monograph could
not have been made more complete by its author. Zopf recognized
the necessity of making descriptions clear and complete, of
including details, and of presenting figures with the descriptions.
It will be remembered that in his monograph Zopf divided his
species into two subgenera, Chaetomium and Chaetomidium.
Under the first subgenus he described as new and figured Ch.
spirale; renamed, described, and figured Kunze's Ch. globosum
under the name Ch. Kunzeanum; described and figured Ch.
murorum Cda., Ch. pannosum Wallr., Ch. crispatum Fckl., and
Ch. indicum Cda.; described Ch. cuniculorum Fckl., Ch. elatum
Kze.; redescribed and figured his own species, Ch. bostrychodes,
which had already appeared at an earlier date. Under the
subgenus Chaetomidium he described and figured Ch. fimeti
Fckl.

Since 1881, writers have for the most part been content with
publishing scattered new species and varieties, and no one has
attempted a complete survey of all which have appeared, or a
revision of the badly confused literature. Among these forms,
Which number at least forty and which represent the work of
nearly as many authors, two only can be regarded with certainty
as good species; Ch. contortum, described by Dr. Peck (67) in
1896 and Ch. simile, by Massee and Salmon (59) in 1902. During
the year 1910, however, Miss Helen L. Palliser (65) wrote her
revision of the Chaetomiaceae of North America, and in the
same year Bainier (3) published his monograph of the genus.
The work of both these authors includes the study of a con-
siderable number of forms.

In Palliser's revision seventeen species are enumerated,
fourteen of which had previously appeared in publications of
other authors. Ch. aterrimum is described for the first time
under a name given it by Ellis and Everhart, while Ch. cochliodes,
Ch. spirochaete, and Ch. flexuosum are described as new. At the
end of her paper four species are enumerated as doubtful forms.
It may be well to state here the more important differences in
arrangement between Palliser’s paper and that of the present
writer. In Palliser’s revision Ch. caninum Е. & Е. stands as а

162 MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

species, rather than as a synonym of Ch. bostrychodes Zopf; Ch.
lanosum Peck and Ch. olivaceum С. & E. appear as species, rather
than as synonyms to Ch. globosum Kze. Тһе name Ch. Ellisianum
Sacc. is retained for the plant which the present writer has re-
named Ascotricha pusillua (E. & E.); Ch. contortum Peck is treated
as a synonym to Ch. crispatum Fuckel, rather than as a distinct
species; Ch. melioloides C. & P. is recorded as a species, rather than
as a synonym of Ch. indicum Cda.; Ch. sphaerospermum С. & E.
is treated as a species of the genus Chaetomium rather than as

Ascotricha chartarum Berk.; Winter's name, Ch. setosum, is re-

tained and Ch. indicum of Zopf, but not of Corda, appears as a
synonym thereto.

The monograph of the genus Chaetomium by Bainier contains
a brief historical sketch and review of the work on development;
a characterization of the genus; a description (with figures)
of twenty-two species and three varieties, twelve species and two
varieties of which are described under new names as follows:
Ch. megalocarpum, Ch. contortum (not of Peck), Ch. spirilliferum,
Ch. undulatum, Ch. setosum (not of Winter or of Ellis & Ever-
hart), Ch. comosum, Ch. glabrum (not of Berkeley & Broome),
Ch. tortile, Ch. formosum, Ch. formosum, var. ovatum, Ch. formosum,
var. neglectum, Ch. caprinum, Ch. torulosum and Ch. rigidulum.
The most striking characteristic of Bainier's monograph is the
multiplication of species and the fact that several species and
varieties have been made from Ch. globosum Kze. and Ch. bostry-
chodes Zopf, both of which are variable forms. It should here
be noted that three of the names used by Bainier, viz., Ch. con-
tortum, Ch. setosum, Ch. glabrum, must, if the priority rule is to
be followed, be changed, since Dr. Peck had previously used the
name contortum; Winter, and later Ellis and Everhart, had made
use of the name setosum, and Berkeley and Broome had used the
name glabrum.

CHARACTERS OF THE GENUS CHAETOMIUM

wall membranaceous, brittle, distinctly cellular, provided wit!

appendages in the form of variously modified hairs. Mycelium |

Es

MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

163

in the form of a densely aggregated mass of fungus threads

radiating in ropy strands from the point of origin.
haped or rarely linear

walled, delicate, stalked, evanescent, club-s

and cylindrical, eight-spored

frequently olive-brown, typically lemon-shap

Asci thin-

Spores single-celled, colored, most

KEY TO THE SPECIES OF CHAETOMIUM

I. Terminal hairs unbranched.

А;

В.

n

D

Terminal hairs straight or и во. Ѕрогеѕ
uërg triangular.

Terminal hairs flexuous, circinate at their
tips.

Terminal hairs slender, about 4и in thick-
ness, with open, circinate tips. Plant blue-
gray to blue-black.

Terminal hairs stout, about ти in thickness,
coarsely encrusted, with closed, circinate
tips 2-3 times re-curved. Plant gray-

ack,

Terminal hairs tortuous, consisting of succes-
sively reversed loops with connecting
arch ips circinate

es -
Terminal hairs stout, 15 4 in thickness at
1

crown of terminal arch. тсһез short
thecium large with diameter as great as

1050 д.

Terminal hairs slender. about бы іп thick-
ness at crown of terminal arch. rches
long and graceful, loops far apart. Peri-
thecium not exceeding 340 и in diameter.

Terminal hairs rather slender, about 7.5 и а

mated. т not exceeding 350 и
in diam

- Terminal sued tortuous or spirally re

Terminal hairs contorted as well intri-
cately апа irregularly Vise coiled,
usually with arched circinate tips. Регі-
thecia subglobose. Plant gray-black.
enbinal hairs coi s

bose. Plant blue-

M

“

yrs

8. Ch.

pay oy

g

Q
=

с
с
=

g

trigonosporum (p. 166).

. murorum (p. 166).

‚ circinatum (р. 168).

. contortum (р. 169).

. simile (p. 169).

‚ crispatum (р. 171).

‚ tortile (р. 172).

convolutum (p. 173).

164

Terminal hairs more slender, finely coiled
above into a spiral of small diameter which

tinuously arcuate from base to incurved

Terminal hairs incurved at tips, spores ir-
regularly ovate. Plant gray, pale olive
or golden yellow.

Terminal hairs circinate or slightly convo-
lute at tips. Spores Plant
gray to pale olive.

Terminal hairs 1-3 spirally convolute at
tips. Spores with shape of a section of
an orange. Plant olive yellow.

II. Terminal hairs constantly and conspicuously
at least not

fusiform.

small, not ex ceeding 160 и in
der Plant black at maturity, gray-
green in old age
Terminal hairs of two types; (a) stout,
ranching by obtuse angles with branch-
lets reflexed; (b) more slender, irregularly
ed, branching by
Perithecium not
Plants black

when young, black at
never dichotomously

form of stiff, spine-like

Perithecium extremely

lant ochraceous.

branched, with branches
ork,

spirall

MoNoGRAPH op CHAETOMIUM AND ASCOTRICHA

9. Ch. subspirale (p. 173).

10. Ch. aureum (p. 174).

II. Ch. fusiforme (р. 175)-

12. Ch. trilaterale (р. 175).

13. СВ. funicolum (р. 176).

14. Ch. indicum (р. 178).

15. Ch. elatum (p. 180).

16. Ch. spinosum (р. 187).

17. Ch. ampullare (p. 187).

18. Ch. cuniculorum (p. 188).

MonoGRAPH OF CHAETOMIUM AND ASCOTRICHA 165

A. Terminal hairs undulate or loosely, spirally

coiled.
perenne! hairs undulate, extremely fine and

low, narrow above. Plant gray or olive
yellow, at maturity golden yellow.
Terminal hairs undulate or loosely spirally
coiled. Branches few. Plant gray, olive-
chocolate-brown. 20. Ch. globosum (р. 100).
В. Terminal Su convolute in definite spirals.
а. Perithecia gray when young, becoming
in old age
x. Perithecia subglobose or ovate.
Terminal hairs slender, about 5 и in
thickness near their tips, with 6-14
coils. Spores large, 9 X 7 m, rich
olive-yellow to olive-brown 21. Ch. spirale (p. 199).
Terminal hairs stout, biet: ou in
thickness near tips with 10-18
coils. Sporesolive-brown, 7 X би. 22. Ch. aterrimum (p. 200).
Terminal hairs regularly coiled with
5-7 convolutions, or rem
coiled with 2-3
frequently produ cing branches.
Spores elliptical in face view, hya-
line or only slightly colored,

. Ch. sphaerale (p. 189).

-
©

7.4-6 и. 23. Ch. bostrychodes (р. 201).
y. Perithecia elongated.
Spores in face view four-sided and
four-angled. 24. Ch. quadrangulatum (p. 202).
Spores lemon-shaped or globose. 25. СВ. caprinum (р. 203).
b. Perithecia green or golden yellow.

convo!

А: eg delicate, either coiling
in spirals or twisting irregularly.
Plant olive-green.

Terminal hairs of ee types; (a)
long, straight, tapering; (b) slen-
der, straight below, coiling above
spirally with about 5-7 convolu-
tions; (c) stout, coarse, straight

26. Ch. cochliodes (p. 204).

: 27. Ch. angustum (p. 206).
у. Perithecia elongated, bottle-shaped.

166 MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

ove. Plant golden-yellow. 28. Ch. torulosum (p. 207).
1. Chaetomium trigonosporum (Marchal) comb. nov.

Bommerella trigonospora Marchal, Bull. Soc. Roy. Bot. Belgique

24:1. 1885; 28: 261—271. pl. 10. 1889.

PLATE 6, FIGS. 6-11

Black. Perithecia of medium size, globose or subglobose to
somewhat elongated, 315 X 221 и (250-340 X 160-262), pro-
vided with long, slender, straight or curved, black cirrhi, seated
on a mat of hyphae which are olive-yellow to olive-brown. Lateral
hairs rather numerous, comparatively short, spine-like, tapering,
below dark olive-brown, minutely roughened, conspicuously and
evenly septate, at base about 4.7 и п thickness; above faded, pale
yellow, obscurely septate, smooth, near tip colorless. Terminal
hairs straight, unbranched, longer than the lateral ones, dark
rich olive, regularly septate to near the tips, smooth or only
slightly and obscurely roughened, at base about 4.7 и in thick-
ness, at tip pale yellow or colorless, without septa. Asci narrowly
and irregularly club-shaped, 8-spored, 50 X 8 и, pars sporif. 32 и.
Spores when young greenish and filled with highly refractive
globules, when mature rich olive-yellow to dark olive-brown, in
face view irregularly triangular, 8.9 X 5.7 и (8.1-9.7 X 4.8-6.4);
when seen edgewise nearly oval, 3.6-4 и broad.

Оп rabbit dung, North Carolina, Herb. В. Thaxter (Chivers
No. 6). Type locality: in heath near Aerschot, Belgium;
on dung of hare.

So far as the writer is aware this is the only form with straight,
unbranched, terminal hairs to be included in this genus. The
species has been grown in varied cultures for many years and
seems to be a true Chaetomium both in its life history and specific
characteristics. While the spores which are here roughly tri-
angular differ in shape from those of the greater number of species,
the variation is no greater than in those of Ch. quadrangulatum
where they are four-sided and four-angled.

2. CHAETOMIUM MURORUM Corda, Icones т: 24. pl. 7, $. 203В-
1837; Icones 2: 29. pl. 13, {. 103. 1838

Chaetomium comatum, var. helicotrichum Saccardo, Michelia I:

222. 1878. |

MoNoGRAPH OF CHAETOMIUM AND ASCOTRICHA 167
Chaetomium griseum Cooke, Grevillea 1: 175. 189%,

PLATE 8, FIGS. 6-10

Blue-black. Perithecia of medium size, globose or ovate
with a bluntly pointed base, 278 Х 267 и (243-337 X 206-337).
Lateral hairs long, graceful, flexed, insensibly tapering to a point,
delicate when young, when mature about 7.5 in thickness, .
dark olive-brown near base, gradually fading at tip, conspicuously
granular-roughened, or nearly smooth, clearly or obscurely
septate. Terminal hairs variable with age; when young stout,
about 5.5 u in thickness at the middle of their length, not circi-
nately curved at tips, but broadly arched throughout their length;
at maturity slender, about 4 іп thickness, gracefully flexed ог
nearly straight, ending in a graceful arch with circinate tip, dark
rich olive-brown, sparsely and irregularly septate, smooth or
roughened by irregular projections, in old age becoming still
darker, frequently losing their circinate tips and tending to
become wavy throughout. Asci broadly and irregularly club-
shaped, 8-spored, 53 X 19 и, pars sporif. 34 и. Spores filled with
greenish refractive globules when young, when mature dark
olive-brown, globose-ovate to narrow elliptical, apiculate at
both ends or apiculate at one end and umbonate at the other,
frequently collapsing by а longitudinal furrow, 12.6 X 8.14
(1 1.3-12.9 X 8.1), when seen edgewise, compressed, 6.4 и broad.

Exsiccati.—Fung. Europ., Series II, Edit. nova, ІП, 234.
Sub Ch. Kunzeanum Zopf: Fung. Gall. XLV, 4436.

This species is extremely common, growing on dung of various
animals, especially on dog dung, and on very varied substrata
from all parts of New England (Chivers No. 13). Reported also
from Montana, by Ellis and Everhart (Anderson No. 651);
from England, by Massee and Salmon; and from Germany, by
Zopf. Type locality: Prague; on damp walls. :

It may be seen from the above that Ch. griseum described
by Cooke in 1873 and Ch. comatum, var. helicotrichum by Sac-
cardo іп 1878 are considered as synonyms of Corda's species.
А careful study of the former has been made from type material
received from the Royal Botanic Gardens at Kew and the char-
acteristics noted seem to agree in every way with those of eh
murorum, though Cooke stated that the threads of his species
Were stouter and the spores larger and colorless. Through the
kindness of Dr. Saccardo it has also been possible to examine

168 MonoGRAPH OF CHAETOMIUM AND ASCOTRICHA

type specimens of his variety and while he found a similarity to
Ch. murorum only in the circinate tips of the hairs, the present
writer has found all the characteristics of the plants examined to
be typical of Corda’s species. On the contrary there seem to
be no reasons for considering the form related in any way to Ch.
comatum, since the hairs are not branched.

Only Ch. elatum Kze. could be obtained from exsiccati speci-
mens in Rabenhorst’s Fungi Europaei No. 234.

Ch. murorum Cda. may at first sight be confused with Ch.
circinatum, but may easily be distinguished from it by the smaller
diameter of the terminal hairs which are only 5.5 и in thickness,
and which are flexed not sinuous, slightly if at all recurved at
their tips, and more minutely and inconspicuously roughened.

3. Chaetomium circinatum sp. nov.
PLATE 8, FIGS. 1-5 ;

Gray-black. Perithecia of medium size, ovate or globose,
325 X 312 u (270-344 X 258-335). Lateral hairs long, flexed,
graceful, sparsely and irregularly septate, at base olive-brown,
roughened, 5.6 in thickness, at the apex smooth, pale olive,
slender. Terminal hairs dark, nearly opaque, olive-brown oF
brownish-black, sinuous, 7 шіп thickness, at the base irregularly
encrusted, at the apex smooth or only slightly encrusted, circinate,
two to three times re-curved. Asci irregularly club-shaped,
8-spored, 70 X 25 и, pars sporif. 50 и. Spores ovate or lemon-
shaped, apiculate or umbonate, olive-brown, 14.3 X 8.9 и (12.9-
I5.3 X 8.1-9.7)

А rare species having appeared only once on a piece of old bur-
lap in a refuse heap, Worcester, Massachusetts (Chivers No. 12).

This species, while similar in certain respects to Ch. murorum,
may be easily distinguished from it by the characteristic terminal
hairs which are sinuous instead of flexed, roughened by crystals
of calcium oxalate, which are irregularly clustered along their
entire length, circinately recurved at the tips, the last coil of
Which often springs out to one side or the other from the plane
of coiling.

MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA 169

4. CHAETOMIUM сохтовтом Peck, Rep. New York State Mus.
Nat. Hist. 49: 24. 1896
PLATE 9, FIGS. 10-12

Black. Perithecia large, globose ог subglobose, 875 X 1050 и.
Lateral hairs numerous, straight or flexed, clearly septate, smooth,
some with equal diameter and olive-yellow throughout, others
thicker (about 5.6 и) and dark olive-brown at base, gradually
tapering and fading toward tip. Terminal hairs dense olive-
brown to black, without visible cross walls, roughened throughout
with blunt, flat-topped projections, nearly straight below, con-
torted above into loops which are separated by short, abrupt
arches, terminating іп an arch with circinate, recurved tip, 15 и
in thickness at the crown of the terminal arch. Asci * fugacious."
Spores when young hyaline, refractive, filled with refractive
globules, when mature dark rich olive-brown, irregularly lemon-
shaped, not always symmetrical, sometimes apiculate at ends,
sometimes barely angular, 11.6 X 9.4 и (10.5-12.5 X 7.5-10.5).

So far as the writer is aware this species has never been found
except in the type locality: Woodside, New York; on bulbs of
lilies, Lilium longiflorum (F. C. Stewart).

On account of the fact that only two mounts of the type
material received from Dr. Peck have been available, it has been
impossible to determine the characteristics of the asci, for although
the perithecium in one mount was broken and the young asci
Were exposed, they could not be accurately studied. It has seemed
desirable, therefore, to quote from the original description re-
garding the asci.

This species has certain features in common with CA. cris-
batum, Ch. simile, and Ch. tortile. From the first it differs in the
Sreater dimensions of the perithecium, and the greater width
and more regular looping of the terminal hairs. From Ch. simile
it differs in the greater dimensions of the perithecium, the greater
width and more compact looping of the terminal hairs and the
greater size of its spores. From Ch. tortile it differs in the greater
dimensions of the perithecium, in the greater width and more
regular looping of its terminal hairs and the greater size of its spores.

5. CHAETOMIUM SIMILE Massee & Salmon, Ann. Bot. 16: 71.
pl. 4, f. 8,0. 1902
Chaetomium glabrum Bainier, Bull. Soc. Myc. France 25: 214.
bl. 21, f. 1-4. 1910.

170 MONOGRAPH ОЕ CHAETOMIUM AND ASCOTRICHA

PLATE 9, FIGS. 1-4

Dark gray to black. Perithecia of medium size, globose or
subglobose, frequently of greater diameter horizontally, 231-
295 X 231-327 и. Lateral hairs not differentiated, but appearing
about the perithecium as a tomentum of yellow, semi-trans-
parent, fine and delicate mycelial threads. Terminal hairs
dense olive-brown to black, with low power smooth, but with
immersion lens irregularly thickened with extremely minute
spines, irregularly and sparsely septate, nearly straight below,
contorted above into loops which are separated by long, graceful
arches, terminating in an arch with circinate tip, 5.6-7 шіп thick-
ness at the crown of the terminal arch. “Asci cylindrical, about
80 X 9-10 yu, 8-spored." Spores monostichous, when young hya-
line, filled with refractive greenish globules, when mature dark rich
olive-brown, subglobose or very broadly elliptical, some clearly
apiculate at one end, barely so at the other, others apiculate
only at one end and rounded at the other, то X 8 u (9.5-10.5
X 7-5-8.9), when seen edgewise, compressed, 6.4 и broad.

Type locality: Kew, England; on dog dung.

Although it has been impossible to examine type material of
the plant which Bainier found on dog dung and described under
the name Ch. glabrum in 1910, the writer is convinced that it is
identical with Ch. simile. The measurements of the perithecia
and spores are the same for both species and the development
of the dark area near the top of the perithecium which Bainier
noted as unusual appears in all species of Chaetomium where
the wall remains so transparent that the darkened spore mass
may be seen.

Through the kindness of the Royal Gardens at Kew the writer
has been enabled to study type specimens of Ch. simile, but
as no asci could be found in mounts made from this material, the
original description has been quoted and the figure of the ascus
has been copied.

At first sight one may confuse this species with Ch. contortum,
Ch. crispatum and Ch. tortile. It differs from these, however, in
the terminal hairs which are of much smaller diameter and which
юа long, graceful arches between their loops. It differs also
from Ch. contortum in the smaller size of the perithecium and
ev both Ch. contortum and Ch. crispatum in its slightly smaller

es.

MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA 171

6. CHAETOMIUM CRISPATUM Fuckel, Symb. Myc. 90. 1869
Sphaeria crispata Fuckel, Fung. Rhen. 2022. 1867.
Chaetomium streptothrix Quélet, Mém. Soc. d’Emul. Montbeliard
1875: 103. pl. 4, f. 40. 1876.

PLATE 9, FIGS. 5-9

Gray to gray black. Perithecia of medium size, globose or
subglobose with a bluntly pointed base. 263 X 253 4 (198-320
X 183-350). Lateral hairs numerous, straight or slightly flexed,
long, slender, gradually tapering to a point, smooth, regularly
septate, about 4 u in thickness and dark olive-brown at base,
fading to:yellow and becoming colorless near the tip, frequently
breaking at maturity and giving to the surface of the perithecium ,
а coarse, rough appearance. Terminal hairs dense olive-brown
to black, rather evenly roughened with minute spines throughout,
below about 4 и in thickness, straight or slightly curved, septate,
twisting or coiling above into an irregular spiral, near the tip
forming alternate loops and arches, ending in an arch with circi-
nate tip, irregularly and obscurely septate above, enlarging to
7.5 in thickness at the crown of the terminal arch. Asci 8-
spored, long, narrow, cylindrical, 80-100 X 8-10 и, pars sporif.
65-80 и. Spores monostichous, hyaline and filled with greenish
refractive globules when young, when mature, dark rich olive-

rown, lemon-shaped, apiculate at both ends or broad and apicu-
late at one end and slightly more pointed and less conspicuously
apiculate at the other, 11.7 X 8.8 u (11.3-12.1 X 8.1-9.7), when
seen edgewise, compressed, lenticular, 6.4 и broad.

This species has been found frequently on various substrata
from different localities in New England (Chivers No. 9). Re-
ported also from England, and various localities in Europe. Type
locality: in the cellar of L. Fuckel; on rotting potatoes. |

Exsiccati.—Fung. Sax. XXIV, 1167. Sub Sphaeria crispata
Fuckel: F ung. Rhen. 2022. |

In 1876 Quélet described under the name Ch. streptothrix fungi
Which he found on rotting potatoes and which he thought might
be the ascosporic stage of Peronospora infestans. While there are
nO figures accompanying his brief and somewhat indefinite -
scription, it seems probable that it is identical with Fuckel's
species for the following reasons: the hairs are described as tortuous,
the specimens were found on rotting potatoes, a place where СВЕ
is peculiarly likely to find Ch. crispatum, and ће name which һе
gave indicates the same type of twisted hairs.

.

172 MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

Ch. crispatum differs in its peculiarly contorted terminal
hairs from all other species which the writer has studied, with the
exception of Ch. contortum, Ch. simile and Ch. tortile. It has a
smaller perithecium and terminal hairs of much smaller diameter
and with more irregular coilings than in Ch. contortum; terminal
hairs of slightly greater diameter and with much more irregular
coilings, and spores of greater diameter than in Ch. simile; a
smaller perithecium, more regularly contorted hairs and larger
spores, than in Ch. tortile.

The writer has seen specimens of the exsiccati mentioned
above and has made and examined mounts from them.

7. CHAETOMIUM TORTILE Bainier, Bull. Soc. Myc. France 25: 214.
Ы. 22. 1910
PLATE 9, FIGS. 13—17

On rabbit dung, Hanover, New Hampshire (Chivers No. 1)
Type locality: Bainier makes no mention of the place where he
found this species or the substratum on which it grew.

Ch. tortile is in certain respects similar to Ch. contortum, Ch-
crispatum, and Ch. simile. The terminal hairs are much more
complicated in their twistings than in those of the other species
mentioned and more slender than those of Ch. contortum. The
Spores are smaller than those of Ch. contortum or Ch. crispatum.

MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA 173

8. CHAETOMIUM CONVOLUTUM Chivers, Proc. Am. Acad. 48: 85.
1912
PLATE 14, FIGS. 9-12
Blue-gray. Perithecia of medium size, globose or subglobose,
244 X 232и (236-254 Х 224-240), frequently provided with
cirrhi, destitute of differentiated rhizoids. Lateral hairs com-

10, are open and of large diameter below, but toward the
extremity constantly and evenly decrease in size and become
more and more closely appressed. Авсі club-shaped, 8-spored,
56.4 X тои, pars sporif. 27.4 и. Spores when young colorless
with granular contents, when mature, pale dull olive, ovate or
lemon-shaped, bluntly pointed at either end, slightly apiculate,
8-8.4 X 6.4 и, when seen edgewise, 4.8-5.6 и broad.

Cultivated on mouse dung from Germany (Chivers No. 18).

This is a rare form, having appeared but once. The species
may be identified by the distinct blue color of the plant when
seen with the naked eye or hand lens, and by the long, spreading
and drooping, terminal hairs, the long series of coils tapering
abruptly to a blunt point.

9. CHAETOMIUM SUBSPIRALE Chivers, Proc. Am. Acad. 48: 84.
1912
PLATE 12, FIGS. 12-17
Perithecia rather large, elon-

twisting near the end into a very small and tightly coiled ae
Terminal hairs slender, graceful, obscurely septate, smooth,

174 MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

below dark olive, straight and about 3.7 и in thickness, above
fading to light olive-yellow or becoming colorless, апа spirally
coiling at first tightly, finally becoming extended and drawn out
into a twisted thread. Asci club-shaped, 8-spored, 45 X 9.7»,
pars sporif. 24 и. Spores pale olive, lemon-shaped, apiculate at
both ends, 6.4 X 5.2-5.6 u.

Frequent in cultures of various substrata from New England.
Appearing in cultures of dung from Holland and South America
(Chivers No. 27).

This species may be easily distinguished by its characteristic
hairs; the lateral ones being short, straight, dark below, tightly
coiled into a spiral of small diameter, hyaline and refractive at
the tips; the terminal ones slender, at first tightly coiled into a
fine delicate spiral, later elongated, twisted rather than coiled,
and giving the appearance of woolly threads.

10. CHAETOMIUM AUREUM Chivers, Proc. Am. Acad. 48: 86.
1012
PLATE 11, FIGS. 12-17

Gray, pale olive, becoming yellow, at length golden-yellow.
Perithecia minute, globose or subglobose, often bluntly point
at the base, 127 X 115 (110-140 X 105-123), without differ-
entiated rhizoids, provided with a long, slender, arched or re-
curved cirrhus. Lateral hairs numerous, slender, straight or
flexed, regularly and distinctly septate, olive-yellow, minutely
roughened, with nearly equal diameter throughout, 3.5 шіп thick-
ness, broadly arched at tips. Terminal hairs olive-yellow, regu-
larly septate, minutely roughened, straight or slightly re-curved,
at base about 3.8 u in thickness, at tip nearly straight or incurved.
Asci club-shaped, 8-spored, 42 X 10 и, pars sporif. 26 и. Spores
when young filled with refractive globules, when mature olive-
brown, irregularly ovate, apiculate at both ends, 9.8 X 5.44
(9.4-11 X 4.7-5.6).

On paper, dung and other materials of various kinds from
New England (Chivers No. I). In cultures of old paper from
Java (R. Thaxter).

г Тһе small size and characteristic golden yellow color clearly
distinguish this species from all others except Ch. trilaterale and
Ch. fusiforme. From the former of these it differs in that the
spores are discharged in long black cirrhi, in the comparative |
obscurity of the perithecial hairs at maturity, in the incurved |

MONOGRAPH ОҒ CHAETOMIUM AND ASCOTRICHA 175

tips of the terminal hairs, and in the irregular, oval shape of its
spores. From the latter species it differs also in producing long,
black cirrhi, in the incurved extremities of its terminal hairs and
in the size of its spores and their irregular oval shape.

The name of this species suggests the characteristic golden-
yellow of the mature plant, but the color usually disappears as
the plants grow old or as they are dried for herbarium specimens,
and the highly colored perithecium becomes black. Moreover
the cirrhi of spores are frequently so long that they overturn the
perithecium.

II. CHAETOMIUM FUSIFORME Chivers, Proc. Ат. Acad. 48: 87.
1012
PLATE II, FIGS. 7-11

Gray or pale olive. Perithecia minute, globose or ovate
with a bluntly pointed base, 120 X 102 и (116-123 X 101-125),
without cirrhi, producing at base a few yellow rhizoids. Lateral
hairs numerous, slender, flexed, not spirally convolute, regularly
and distinctly septate, olive-yellow, minutely roughened, about
2.5 ш in thickness at base. Terminal hairs stouter and more
darkly colored than the lateral hairs, minutely roughened, olive-
brown, си апа distinctly septate, nearly straight or in-
curved, about 3.7-4 и in thickness at base, at tip circinate or
slightly convolute. Asci club-shaped, 8-spored, 48 X II и, pars
sporif. 32 u. Spores when young filled with refractive globules,
when ‘mature olive-yellow or olive-brown, long, narrow, some-
what fusiform, round or apiculate at the ends, 15.8 X 5.4 и (15-16

4.8-5).

A rare species having appeared only oncé on paper from
Alabama, Herb. R. Thaxter (Chivers No. 3).

The long narrow spores distinguish this form from all other
species of Chaetomium. In general characteristics it most nearly
resembles Ch. aureum and Ch. trilaterale, but differs from both
in the long, slender, fusiform spores.

12. CHAETOMIUM TRILATERALE Chivers, Proc. Am. Acad. 48: 87.
12
19 2
PLATE 11, FIGS. 1-6

Olive-yellow. Perithecia minute, globose or ovate with <
bluntly pointed base, 106 X 94 и (100 X 90-97), without сит

176 MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

or differentiated rhizoids. Lateral hairs numerous, graceful,
rather long, regularly and distinctly septate, golden-yellow, at
base straight, minutely roughened and about 2.8 и in thickness
near tips, smooth and 1-3 spirally convolute. Terminal hairs
stouter and more darkly colored than the lateral ones, dark olive-

pars sporif. 26 џи. Spores when young hyaline with obscure
globules, when mature rich olive-yellow to olive-brown, having
the shape of a section of an orange, slightly apiculate at both
ends, 9.5 X 5.5 u (8.9-9.7 X 5.2-6).

This species has appeared only once, on paper, from New
England, Herb. R. Thaxter (Chivers No. 2).

This species has certain characteristics іп common with
Ch. aureum and Ch. fusiforme. From the former it differs in the
more numerous, stout, 1-3 spirally convolute, terminal hairs;
the spirally coiled lateral hairs; the smaller size and unusual
shape of the spores. From the latter species it differs in the
convolute lateral hairs, the shape of its spores and their smaller
size.

13. CHAETOMIUM FUNICOLUM Cooke, Grevillea т: 176. 1873

Chaetomella Сатайй Mattirolo; Savoia, П Ruwenzori т: [3]. pl. 3,
11535: 1909.

Chaetomium Bartholomaei Saccardo & Sydow; Saccardo, Syll.
Fung. 14:490. 1899.

Chaetomium setosum Ellis & Everhart, Am. Nat. 31: 340. 1897.

PLATE 7, FIGS. 0-19

Black. Perithecia small, ovate to globose, 149 X 147#
(130-157 X 130-158), firmly attached to the substratum by
dark olive to black rhizoids, frequently provided with long,
straight or curved cirrhi. Lateral hairs comparatively numerous,
smooth or irregularly roughened by short, blunt projections, stiff,
spine-like, dark olive-brown to black nearly to tip, hyaline and
crumpled at tip, rarely and obscurely septate, about Ap in thick-
ness at base. Terminal hairs forming an especially dense, com-
pact head, dichotomously branched with narrow, acute angles,
frequently alternately constricted and inflated throughout,
roughened over entire length by spines and irregular projections,
near base dark olive-brown to black and about 6.3 и іп thickness,

MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA 177

fading to light brown, pale olive, or colorless tips, at maturity
bearing on the most remote branches clusters of refractive needles.
Asci club-shaped, 8-spored, 33.7 X 8.1 и, pars sporif. 16 и. Spores
when young greenish hyaline, refractive, filled with globules,
when mature dark, rich, olive-brown, egg-shaped to lemon-
shaped, slightly more pointed at one end, apiculate at both ends,
6.3 Х 4.7 и (5.6-6.4 Х 4-4.8).

ExsiccATI.—Sub Ch. setosum ЕП. & Ev.: Fung. Columb. XII,
1126; М. A. Е. 24 Ser. XX XV, 3423.

A rather common species, having been found in cultures of
brazil nuts at Cambridge, on straw from Nairobi, Congo, on corn
stalks from Cambridge, and on old fruit from Germany (Chivers
No. 16). Reported by Peck on an old broom at Albany, New
York. Reported also as Ch. setosum E. & E. by Ellis, and
as Chaetomella Cavallii by Mattirolo on paper. Type locality:
British Museum; on twine (W. Carruthers).

The writer is indebted to the Royal Botanic Gardens at Kew
for type specimens of this species which was first described by
Cooke in 1873 as Ch. funicolum or “Twine Bristle Mould,” and
which has since been found and reported by several writers in
various localities. Although the species seems to have been
clearly characterized by Cooke, Ellis (35) redescribed it in 1897
under the name Ch. setosum, in apparent ignorance of the fact that
Winter (112) had previously used this combination for an entirely
different plant. Authentic specimens examined in the exsiccati
mentioned below are found to be identical with Ch. funicolum

ооКе. In 1901 Saccardo changed the name to Ch. Bartholomaei
in honor of the collector who sent the specimens to Ellis.

Although no specimens have been available the writer is
convinced that the plants which M. Mattirolo (61) described and
figured in 1909 under the name Chaetomella Cavalli аге identical
With Cooke's species. It was discovered in considerable numbers
оп а piece of paper which wrapped some moss. Mattirolo stated
that, considering the present confusion of literature in the genera
Chaetomella and Chaetomium, he thought it best to assign his
Species to the genus Chaetomella. ln spite of every attempt,
he could not determine whether the spores were produced T nee
perithecia or pycnidia or whether in an ascus or on а basidium.
His three figures show a young perithecium, a branched terminal

178 MONOGRAPH ОҒ CHAETOMIUM AND ASCOTRICHA

hair, and several spores, all of which are identical with corre-
sponding structures in Ch. funicolum Cke.

This species may be confused on the one hand with Ch. elatum
from which it differs by the small size of its perithecium and
spores. On the other hand it will be confused with CA. indicum.
In Ch. funicolum the perithecium during its early development is
clothed throughout with long, straight, stiff, black, spine-like
hairs which are more conspicuous than in Ch. indicum and whose
tips frequently appear above the mass even at maturity. Later
the terminal hairs (of one type only) grow up, branching by
narrow, acute angles, until a compact head is formed. Тһе
greenish-gray and powdery appearance of the plants is due in
part to the large number of colorless hair tips and also to the
bundles of crystals on the ultimate branches. These charac-
teristics, present also in Ch. indicum, make it more difficult to
separate the two species.

14. CHAETOMIUM INDICUM Corda, Icones 4: 38. pl. 7, f. 104.
1840
Chaetomium melioloides Cooke & Peck; Peck, Rep. New York
State Mus. Nat. Hist. 27: 106. 1875.
Chaetomium setosum Winter, Hedwigia 26: 16. 1887.

FLATS 7, Fics. 1-8

perithecium and which do not form a dense mass, stout, dichoto-
mously branched with branches reflexed and roughened by
spine-like projections, at base dark olive-brown to black and
about 7.5 и in thickness, fading only slightly or becoming hyaline
at the terminal branches; (b) hairs which appear later, forming at
first a tuft about the ostiole, profusely branched by narrow acute
angles, branches never refl
inflated, light olive-brown or yellow, finely roughened, terminal — .
acicular or prismatic crystals. |

MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA 179

Asci club-shaped, 8-spored, 30 Х 9.4 и, pars sporif. 16.5 д. Spores
hyaline when young and filled with refractive greenish globules,
when mature dark, rich olive-brown, ovate to lemon-shaped,
slightly apiculate at one or both ends, 5.5 X 4.5 и (5.3-7 X 4.5-
5-6), when seen edgewise, compressed, 4.2 ш broad.

On culture in laboratory, Cambridge, Massachusetts, R. Thax-
ter (Chivers No. 14). Reported by Spegazzini from Parque de la
Plata, Argentine, and by other authors from France and Germany.
Reported also as Ch. melioloides by Cooke and Peck, on old stems
of Indian corn, North Greenbush, New York, and as Ch. setosum
by Winter on branches of Berberis buxifolia in Patagonia. Type
locality: India, Tenasserim, Maulmain; on rotten paper (Dr.
Helfer).

While this species seems to have been clearly described and
figured by Corda, it has since been re-described by Cooke and
Peck in 1875 as Ch. melioloides, and by Winter in 1887 as Ch.
setosum.

The writer must here acknowledge his indebtedness to Dr.
Charles Peck for his generosity in furnishing specimens of Ch.
melioloides as well as several other of his species. In answer to
inquiry regarding the possible identity of Ch. melioloides with
Corda's СВ. indicum, Dr. Peck kindly sketched with a camera
typical hairs and spores, and wrote as follows: “The two species
certainly run close together and if I were inclined to overlook small
differences I could easily make myself believe that they are forms
of опе species. 1 notice, however, that the spores of СЁ. indicum
are more elliptic as required by the description of the species, and
that they run a little longer than in our plant. But I never would
think of describing them as acute at each end. Can this be а
mistake in the description or is it due to culture modifications?
After a very careful examination of the two plants, the present
writer is convinced that the two are identical and that Dr. Feck:
name should appear as а synonym to Ch. indicum. The hairs as
sketched by Dr. Peck and the spores as studied in mounts from
the type specimen are typical of Corda’s species.

Through the kindness of M. Hariot the writer has been able
to study Ch. setosum Winter, and it has been found that the plant
thus named is identical in every way with Ch. indicum Cda. It
Produces the same dichotomously branched hairs with widely

180 MONOGRAPH ОЕ CHAETOMIUM AND ASCOTRICHA

spreading branchlets, and spores of the same shape and size.
The statement made by Palliser in North American Flora (p. 63),
where Ch. indicum Zopf is listed as a synonym under Сл. setosum
Wint., to the effect that Ch. setosum Wint. is the same as Ch.
indicum Zopf, but not the same as Ch. indicum Cda., is ‘not
easily understood, since Zopf in Nova Acta (p. 279) simply copied
Corda's description of Ch. indicum, and made no statements
which seem in any way at variance with Corda's original descrip-
tion.

On the one hand this species may be confused with Ch. elatum,
from which it differs in the small size of its perithecium, the
spreading or reflexed branches of the terminal hairs, and by its
spores which have much smaller dimensions. Оп the other hand
it may be confused with Ch. funicolum, especially when only the
mature stage is studied. In its younger condition Ch. indicum
is covered throughout with straight, spine-like hairs, the lateral
ones only of which are visible at maturity. Later terminal hairs
appear with branches spreading or reflexed, and still later, hairs
which branch by narrow angles and which grow up among the
branches of those already formed. In this way a dense head is
formed consisting of innumerable branches, and the identity of
the hairs is more or less concealed. Тһе plant finally has a
greenish-gray and powdery appearance similar to that of Ch.
funicolum Cke., due in part to the large number of refractive hair
tips and also to the clusters of crystals on the ultimate branches.

I5. CHAETOMIUM ELATUM Kunze & Schmidt, Deutsch. Schwám.
3. 1818; Fries, Syst. Myc. 3:253. 1829

Chaetomium atrum Link; Linnaeus, Spec. Plant. Ed. 5, 1: 40. 1824-

Chaetomium atrum, var. distinctum Roumeguére, Rev. Myc. 8:
198. 1886. Not Chaetomium atrum, var. Therryana Roume-
guére & Patouillard, Rev. Мус. 5: 29. 1883.

Chaetomium comatum (Tode) Fries, Syst. Myc. 3: 253. 1829.
Not Chaetomium comatum, var. helicotrichum Saccardo,
Michelia 1: 222. 1878.

Chaetomium comatum, var. ligni Roumeguére, Fung. Gall. LXIV,
6309.

Chaetomium Fieberi Corda, f. chartarum Roumeguére, Fung. Gall.
LIX, 5827. 1801.

MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA 181

Chaetomium Fieberi Fuckel, Symb. Myc. 90. 1869. Not Ch.
Fieber? Corda.

Chaetomium glabrescens Ellis & Everhart, Proc. Acad. Nat. Sci.
Phil. 1893: 130. 1893.

Chaetomium graminicolum Fuckel, F ung. Rhen. VII, 647. 1863.

Chaetomium graminis Rabenhorst, Bot. Zeit. 34: 569. 1851.

Chaetomium lageniforme Corda, Icones т: 24. pl. 7, f. 2034.

1637.

Chaetomium Libertii Roumeguére & Patouillard, Rev. Myc. 5:
15. pl. 35, f. 2. 1883; Fung. Gall. XXIV, 2376.

Chaetomium pannosum Wallroth, Flora Crypt. German. 2: 267.
1833.

Chaetomium velutinum Ellis & Everhart, Jour. Myc. 1:90. 1885.

Conoplea atra Sprengel, Syst. 4: 554.

Conoplea atra Persoon, Syn. Fung. 1: 235. 1801.

Sphaeria comata Tode, Fung. Mecklenb. 2:15. 1791.

PLATE 6, FIGS. I-5

Black. Perithecia large, ovate, 435 X 391 и (418-500 Х 334-
451), seated on a subiculum of dark olive-brown to black rhizoids
which most frequently hold the perithecia in place through old
age. Lateral hairs numerous, long, slender, graceful, unbranched,
near the base dark olive-brown to black, coarsely roughened and
about 5 и in thickness, gradually tapering and fading to slender,
Pale to hyaline smooth tips which are obscurely septate. Ter-
minal hairs extremely coarse, conspicuously roughened through-
out with irregular pyramidal projections and blunt spines, once
ог twice dichotomously branched with the branches widely
Spreading and often reflexed and once or twice irregularly forked
by an acute angle, at base black and about 9.5 р in thickness,
tapering and fading to slender, hyaline tips. Asci broadly and
irregularly club-shaped, 8-spored, 64-75 Х 17 и, pars sporif. 34 и.

pores hyaline to light olive when young, when mature dark,
rich, olive-brown, lemon-shaped, apiculate at both ends, 12.8
X 8.7 u (12-13 X 8.4-9.5), when seen edgewise, compressed,
7.4 и broad.

ExsiccATI.—Brit. Fung. I, 49; Deutsch. Schwäm. 184; Erb.
Crit. Ital. ХУП and XVIII, 877; Fung. Aust. Х, 989; Fung.
Bavar. Х, 927; Fung. Brit. I, тоо, and ІП, 290; Fung. Carol. Ш,
66; Fung. Columb. VII, 621, and XI, 1034; Fung. Europ. VI,
529, XII, 1147, and XXVI, 2527; Fung. Fenn. ІХ, 820, and

182 MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

X, 980; Fung. Gall. I, 66, and XV, 1428; Fung. Rhen. VII,
646; Fung. Rossiae II, 83; Fung. Sax. XVII, 834; Herb.
Crypt. Belg. I, 83; Herb. Myc. I, 58; Klotzsch. Herb. Viv.
Myc. XI, 1032; Myc. March 343, 4534, 4751; Myc. Univ.
ҮШІ, 758; М. A. Е. VI, 560; РІ. Crypt. de Fr. Set. I, М, 237;
Rehm. Asc. 247b; Schweiz. Krypt. XI, 526; Sub Ch. atrum
Link: Fung. Gall. IV, 325; М. & М. Stirp. XV, 1483; РІ.
Crypt. de Fr. Set. III, II, 86. Sub Ch. atrum Link, var. dis-
tinctum Roum.: Fung. Gall. XXXIX, 3883. Sub Ch. atrum
Link, var. Therryana Roum. et Pat.: Fung. Gall. ХХУ, 2496.
Sub Ch. comatum (Tode) Fr.: Erb. Crit. Ital. Series II, XVI, and
XVII, 834; Fung. Gall X, 975; Jaap, Fung. 372; Krypt.
Vind. 1814; Misc. rar. (Vestergren) LVII-LVIII, 1420; Myc.
Ital. IX, 841; Myc. March. 4016; Myc. Venet. VII, 636. Sub
Ch. comatum (Tode) Fr., var. ligni Roum.: Fung. Gall. 64, 6309.
Sub Ch. Fieberi Cda., var. chartarum Roum.: Fung. Gall. LIX,
5827. Sub Ch. graminicolum Fckl.: Fung. Rhen., VII, 647.
Sub Ch. graminis Rabh.: Klotzsch. Herb. Viv. Myc. XVI, 1555-
Sub Ch. lageniformeé Cda.: Herb. Myc., Edit. Nova, VI, 521.
Sub Ch. Libertii Roum. et Pat.: Fung. Gall. XXIV, 2376. Sub
Ch. murorum Cda.: Fung. Europ. Series II, Edit. nova, ІП, 234.
Sub Ch. pannosum Wallr.: Fung. Europ. Series II, XXI, 2025;
Fung. Gall. I, 61; Herb. Myc., Edit. Nova., VIII, 748; Klotzsch.
Herb. Viv. Myc. XVI, 1556; Myc. March. 1548, 4752; Myc.
Ital. XV, 1475.

This is a very common and widely distributed species, having
appeared on substrata of very varied character such as old rope,
straw, paper, barrel hoops, old brooms, and on the dung of
different animals. The writer has found this species on such
materials from many localities in New England, and from the
Carolinas, Maryland, California, Illinois, New Jersey, and Penn- `
sylvania; also from Switzerland, Germany, and the Galapagos
(Chivers No. 15). Тһе species has also been reported by many
other authors from different localities in North America, from
England, Scotland, Russia, and from many places in Europe.

There is no species of Chaetomium, with the possible excep-
tion of Ch. globosum Кге. which has received so much attention
as Ch. elatum, and while the characteristics described by Kunze

MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA 183

and figured by Greville seem to be very clear, the form has been
re-described under many different names and confused many times
with other species.

In 1818 Kunze gave the name СЙ. elatum to specimens which
were collected in the spring at Halle, Germany, on dead leaves of
Typha and Sparganium, and which also occurred commonly on
dried stems and leaves of the Aroidaceae and Gramineae. He
called attention to the simple, unbranched hairs of the perithecium
near the base, and the very long, rigid, branched hairs above, and
while he gave no measurements of spores, he described their shape
as oviform or globose.

In 1826 Greville (43) published a diagnosis of Kunze's Ch.
elatum accompanied by six figures, and while the description
contains no more information than that of Kunze, the figures are
instructive. ;

In 1824 H. F. Link (54) published a note included in a brief
comment on the Conopleae, to the effect that Conoplea atra
Persoon is Chaetomium atrum, and in 1853 Desmaziéres (26)
made the following statement: “According to specimens which
we have received from M. Persoon, his Conoplea atra is surely a
Chaetomium and not the Myxotrichum chartarum Fries (38, р. 349]
thought it to be." It should here be stated that in connection
with his specimens Desmaziéres listed Conoplea atra Persoon (е а
Synonym under Ch. atrum Link, and indicated that һе had verified
the Synonymy. Тһе very generous specimens of Ch. atrum, which
were distributed by Desmaziéres, have been examined and found
to be typical in every way of Ch. elatum Kze. It may be seen,
therefore, that Conoplea atra Persoon is identical with Ch. atrum
(Persoon) Link, and with Ch. elatum Kze.

In 1829 Fries (38) gave to the Sphaeria comata of Tode (103)
the new name Ch. comatum, and the use of this and of Кипге 5
older name, Ch. elatum, has varied with later authors. In many
Cases the former has been arranged as a synonym under the latter,
but Saccardo (79, p. 221) and several other authors prefer the
use of Ch. comatum (Tode) Fr., with Ch. elatum listed as a synonym
thereto. While Fries published no figures there seems to be no
doubt that his form is identical with Kunze's species, and since
adequate description and figures of this plant under the name

184 MONOGRAPH ОЕ CHAETOMIUM AND ASCOTRICHA

Ch. elatum were published by Kunze and Greville previous to the
time Fries introduced the name comatum into the genus, and so
far as the present writer is aware, no figures under the name Ch.
comatum have ever appeared, it seems best to arrange Ch. coma-
tum as a synonym under Ch. elatum Kze.

In 1833 this plant was re-described by Wallroth under the
name Ch. pannosum. Zopf (113) in 1881 listed and figured Wall-
roth’s species, and stated in connection with his description of
Ch. elatum Kze. that the two forms differed on account of the fact
that in the former the mycelium was red brown, while in Ch. elatum
it was a golden color. His figures of Ch. pannosum, however, show
characteristics identical with those of Ch. elatum, and even though
the present writer, in all cultures of this plant, has found the
mycelium varying only from white to yellow and greenish-yellow,
he is inclined to disagree with Zopf regarding the separation of
these forms into two species, and rather to list Ch. pannosum аз а
synonym under Ch. elatum. The writer is led to question if
color-producing bacteria which so easily contaminate cultures,
or some change such as peculiar fading induced by the environ-
ment or substratum, would not account for the appearance which
Zopf found in his cultures.

The species was re-described and figured by Corda in 1837
as Ch. lageniforme, and again by Rabenhorst in 1851 as Ch.
graminis. While Rabenhorst speaks of the hairs of his species
as somewhat simple and not intertwined, it has been found from
а study of authentic exsiccati specimens, in Klotzsch. Herb. Мус.
No. 1555, that they are no less branched nor intertwined than in
Ch. elatum.

Fuckel (40) listed this plant in 1861 as Ch. graminicola and
distributed specimens under that name in Fungi Rhenani Хо. 647.
It should be stated here that Fuckel used Rabenhorst’s name in
connection with his specimens. It has been impossible to find
any reason for the use of the name graminicola. The writer is
led to the conclusion that graminis is the basis for this usage and
that Fuckel has mistaken this for graminicola. The form to
which Fuckel in 1869 called attention in the Symbolae Myco-

logicae (p. 90) under the name Ch. Fieberi is without question

' identical with Ch. elatum.

MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA 185

In 1883 Roumeguére and Patouillard redescribed the species
as Ch. Liberti. The writer has examined authentic exsiccati
specimens of this form in Fung. Gall. No. 2376, and it is not
clear to him why in their diagnosis, the authors compared and
contrasted it with Ch. crispatum, since it does not bear the slightest
resemblance to that species.

In 1885 Ellis gave the name Ch. velutinum to specimens which
were collected by Carpenter on a damp maple log, but in North
American Pyrenomycetes (32, p. 124) the same author lists Ch.
velutinum as a synonym under Ch. pannosum. According to the
arrangement used by the present writer Ellis's name must be listed
as a synonym to Ch. elatum.

In 1886 Roumeguére applied the name Ch. atrum Link, var.
distinctum, to this species which he found growing isolated and
arranged in parallel lines on poplar wood. While the preparation
Which could be obtained from Fung. Gall. No. 3883 was broken
and somewhat scanty, it was possible to identify and study the
spores and fragments of the branched hairs. The characteristics
of these structures were found to vary in no way from those of
Ch. atrum Link, and therefore from Ch. elatum Kze. That peri-
thecia should be scattered or that they should appear arranged in
lines along the substratum would seem to be conditions which
could hardly warrant the separating of a variety from the type
form, since such characteristics are likely to appear in any species,
especially when the substratum is more or less furrowed as it is
in the case of stems.

In 1891 Roumeguére distributed specimens of the same plant
in Fung. Gall. No. 5827, under the name Ch. Fieberi Cda., f.
chartarum. No description was published at that time, however,
and so far as can be learned, no mention or diagnosis has since
been made: Тһе present writer has examined authentic speci-
mens and has found them to be characteristic of Ch. elatum Kze.
in all details. Two years later, in 1893, the same author used
the name Ch. comatum (Tode) Fr., var. ligni, in Fung. Gall. No.
6309, for plants which he found on wood. Here again no descrip-
tion is to be found with the specimens and no subsequent нағы
of them has appeared. It does not seem advisable to retain this
name and set apart a variety from the type form simply on

186 MONOGRAPH ОЕ CHAETOMIUM AND ASCOTRICHA

account of the fact that it was found on wood, especially since a
study of the authentic exsiccati specimens shows all structures to
be typical of Ch. comatum (Tode) Fr. which, as we have seen, is
synonymous with Ch. elatum Kze.

Finally Kunze's species was redescribed by Ellis in 1893
under the name Ch. glabrescens. It has been possible to examine
type specimens of Ellis's species which are now in the possession
of Dr. Farlow at Cambridge, and while the mounts contain
for the most part rather fragmentary remains of perithecia, and
while the hairs in most cases are broken below the point of origin
of the first branches, still in other cases clusters of terminal hairs
- show characteristics identical with those of Ch. elatum Кге.

The spores also are typical of Kunze's species both in shape and
size.

All exsiccati mentioned above have been carefully examined and
with two exceptions have been found to be typical of Ch. elatum.
However, only plants of Chaetomidium fimeti (Fkl.) Zopf could

be obtained from exsiccati specimens of Ch. atrum Link in Fung.
Gall. IV, 325, and of Ch. atrum Link, var. Therryana Roum. et
Pat. in Fung. Gall. XXV, 2496. Іп the specimens of Ch. panno-
sum Wallr. in Myc. Ital. XV, 1475, perithecia of Ch. murorum
Cda. appeared in considerable numbers.

The following exsiccati specimens also consist wholly or in
part of Ch. elatum Kze.: Ch. chartarum Ehr. in М. А. Е. No. 1541;
Ch. Fieberi Cda. in Fung. Gall. No. 6409, Herb. Myc. No. 165,
and Myc. Ital. No. 1288; Ch. Fieberi Cda., var. chartarum Roum.
in Fung. Gall. No. 5827; Ch. Kunzeanum Zopf in Myc. March.
No. 3246; Ch. lanosum P. in Fung. Gall. No. 4437; Ch. murorum
Cda. in Fung. Europ., Series II, Edit. nova, ІП, 234; C^. oli-
vaceum Cke. and Ellis in Myc. Univ. No. 1942 and in Fung.
Columb. No. 512.

No other species with which this should be confused have
come to the attention of the writer with the exception of Ch.
indicum Cda. and Ch. funicolum Cke., from both of which it
differs in its large size, coarse appearance, and in the greater
size of its spores,

MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA 187

16. CHAETOMIUM SPINOSUM Chivers, Proc. Am. Acad. 48: 86.
1912
PLATE 10, FIGS. I-7

angles with it, or backward from it. Later the primary branches

есоте branched until an intricate system is formed. Branches
light yellow to colorless, covered with delicate spines. Asci
very narrowly club-shaped, 41 X 7.5 и, pars sporif. 22 и. Spores
when young filled with granules and globules, greenish, refractive,
when mature pale olive, egg-shaped, 5.9 X 3-9 и (5.6-6.4 X 3.2-4).

Growing in cultures of dung from Buenos Ayres, В. Thaxter
(Chivers No. 7),

This is apparently a rare species having appeared but once.
The egg-shaped spores and the branched, terminal hairs are
peculiar to the species. From the dark, stiff, spine-like shafts
of the terminal hairs arise slender, delicate, irregularly swollen
and constricted outgrowths, from which secondary branches arise
Which elongate and precede the cirrhus of spores as it forms.
In this way a support is formed for the spore mass.

17. CHAETOMIUM AMPULLARE Chivers, Proc. Am. Acad. 48: 86.
I9I2
PLATE 10, FIGS. 8-12

Ochre. Perithecia rather large, extremely elongated, flask-
shaped, 489 X 147 и (456-532 X 137-167), producing at —
a copious, black mass of spores and olive-yellow rhizoids. saa
airs comparatively few, long, slender, graceful, straight or nearly
50, insensibly tapering, at base dark olive-brown, smooth or та
minutely roughened and about 7.5 и in thickness, above brig :
Yellow, terminating in rather long, colorless, refractive, thin an
More or less collapsed tips. Terminal hairs smooth, very long,

188 MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

graceful, insensibly tapering, straight for a long distance above
the base, distinctly and regularly septate, at base about 7.5 шіп
thickness, from base to apex successively golden-brown, golden-
yellow, pale yellow, hyaline, terminating in colorless, elongated,
more or less collapsed tips, producing in the upper portions
branches which are frequently septate and only slightly colored
at base, hyaline and delicate above, and which in turn produce
branches of like nature. Asci club-shaped, 8-spored, 45 X 9.7 №,
pars sporif. 23 и. Spores bright olive-yellow, umbonate at either
end, lemon-shaped, 8.1-8.0 Х 6.4 и.

On culture of sail cloth from Lowell, Massachusetts (Chivers
No. 4). On dung from North Carolina (R. Thaxter).

This species is clearly characterized by the very much elon-
gated bottle-shaped perithecium, and by the terminal hairs which
are drawn out into long, hyaline, tangling, easily collapsible
threads, the branches of which may be very easily overlooked
except with higher magnifications.

18. CHAETOMIUM CUNICULORUM Fuckel, Symb. Myc. 89. 1869

Brownish-black. Perithecia spherical, ovate or egg-shaped,
370 и. Terminal hairs few, dark brown at base, lightly colored
at tips, twice as long as the perithecium, straight below, stiff,
septate, 5-7 шіп thickness, seldom branching at the base, more ог
less encrusted with calcium oxalate, thickly entangled, forming a
compact mass about the ostiole, some remaining simple, others
branching dichotomously at their summits and anastomosing
with each other by their free ends. Lateral hairs straight, rigid,
tapering, unbranched. Asci club-shaped, 8-spored. Spores dark
olive-brown, elliptical or spindle-shaped, barely apiculate at
the ends, 10-12 X 7-9 u

ExsiccaTI.—Fung. Rhenan. 1961.

Type locality: Freinweinheim, Germany; on rabbit dung.
Reported also by Bainier as occurring very commonly on dung of
rabbit.

In spite of repeated attempts it has been impossible to obtaina |

satisfactory mount from the type material of Ch. cuniculorum

Fckl. from the set of Fungi Rhenani at Harvard University: | 2 |
Тһе specimens consist of two rabbit pellets, exposed to the leav ^^

of the book. These pellets are thickly covered with perithecia of 7.

Sordaria, but at best only fragments of perithecia of Chaetomium |

could be obtained. In several cases many bases of hairs could

MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA 189

be distinguished which were very dark to black below, yellow and
finally hyaline and refractive above, clearly septate, and irre-
gularly branched. These hairs were studied with extreme care
with the hope that anastomosing branchlets might be found, but
none could be seen.

The species was first described by Fuckel, in the Symbolae
Mycologicae, in 1869, and again by Zopf in his monograph in
Nova Acta in 1881, and was finally redescribed and figured by
Bainier in his monograph in 1910. Since the writer has never
found this form and the available type specimens have not been
entirely satisfactory, he has been obliged to rely on those men-
tioned above for his information. Zopf in comparing and con-
trasting this species with “Ch. Kunzeanum” has stated that the
two resemble each other in respect to the dense mass of terminal
hairs, but differ in respect to the shape and size of the spores.
Bainier has stated that the perithecia are pure white during their
earlier stages, and that the terminal hairs may be divided into
two groups, those which are 5.6 и in diameter, long and rigid,
and those only ішіп diameter and only half as long as the first
type. He described the spores as dark bluish-gray or greenish.

I9. CHAETOMIUM SPHAERALE Chivers, Proc. Am. Acad. 48: 84.
1912
PLATE II, FIGS. 18-23
Grayish-yellow, olive-yellow, with age golden-yellow. Peri-
thecia rather large, globose or subglobose, evenly rounded at
base, distinctly narrowed above, 312 X 276 и (300-329 X 262-
300), frequently provided with short, black cirrhi, without rhi-

base about 3.7 іп thickness, others wavy, rather short, un-
branched, at base about 2.8 и in thickness. Terminal hairs long,
th, irregularly

when young filled with refractive greenish hyaline glo |
mature dark olive-brown, lemon-shaped to globose, apiculate or
umbonate at both ends, 7.3-8.1 X 6.4 и.

190 MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

А rare species having appeared only once in a culture of cater-
pillars from Reading, Massachusetts (Chivers No. 28).

The perithecium, globose below and conspicuously narrowing
above, is peculiar to this species. Тһе slender delicate hairs
and the entire absence of differentiated rhizoids are also sig-
nificant characteristics.

20. CHAETOMIUM GLOBOSUM Kunze, Myc. Hefte т: 15, 16. f. 0-4.
1817

Chaetomium affine Corda, Icones 4: 37. pl. 8, f. ror. 1840.

Chaetomium amphitrichum Corda, Icones 4: 37. pl. 8, f. 103:
1840.

Chaetomium Araliae Corda, Icones 4: 37. pl. 8, f. 102. 1840.

Chaetomium chartarum Ehrenberg, Sylv. Myc. Berol. 15, 27.
1818.

Chaetomium cymatotrichum Cooke, Grevillea 12: 21. 1883.

Chaetomium Elasticae Koorders, Verhandd. d. K. Akad. v. Weten-
schappen te Amsterdam (Tweede Sectie) 13“: 185. f. 10.

1907.

Chaetomium Fieber? Corda, Icones 1: 24. pl. 7, f. 293c. 1837.

Chaetomium Fieber? Corda, var. chlorina Saccardo, Myc. Venet.
X, 906, 1876; Michelia 1: 27. 1877.

Chaetomium Fieberi Corda, *** Saccardianum Bommer & Rous-
seau, Misc. Мус. т: 17. 1884. Not Chaetomium Feiert
Fuckel.

Chaetomium Kunzeanum Zopf, Nova Acta Acad. Leop.-Carol. 42:
278. M. 1$. 1881.

Chaetomium Kunzeanum, var. chlorina “Mich. " Bull. Soc. Мус.
France 25: 202. pl. 13, f. 1-4. 1010.

Chaetomium Kunzeanum, var. steet Bommer & Rousseau,
Bull. Soc. Roy. Bot. Belgique 23: 207. 1884.

_ Chaetomium lanosum Peck, Вер. New York State Mus. Nat.
Hist. 28:64. 1876.

Chaetomium macrosporum Saccardo & Penzig, Michelia 2: 591»
1882.

A

Chaetomium megalocarpum Bainier, Bull. Soc. Myc. France 25:
202. pl. 16, f. 1-4. тото.

Chaetomium olivaceum Cooke & Ellis, Grevillea 6: 96. pl. 100, f.38:
1878. |

МОХОСЕАРН ОҒ CHAETOMIUM AND ASCOTRICHA 191

Chaetomium olivaceum, var. chartarum Ellis & Everhart, N A

Pyren. 125. 1892. Sa,
Chaetomium olivaceum, f. ch

‚ f. chartarum Roumeguére, Rev. f

p. g ev. Myc. II:
Chaetomium oospora Beauverie, Ann. Univ. Lyon, Nouv. Sér. I

3: 201-218. 1900. x
répond orientale Cooke, Grevillea 5: 103. pl. 86, f. 11. 1877.

aetomium setosum, Bainier, Bull. Soc. Myc. France 25: 209.
^ pl. 18, f. 3-7. 1910.

aetomium spirilliferum Bainier, l. c. 2

C4 «€. : 207. pl. E

an 5 "o pL 10, J: 1—4.

Chaetomium undulatum Bainier, 1. с. 25: 208. pl. 16, f. 4-7- 1910.

PLATE IO, FIGS. 13-16; PLATE 15, FIGS. 9-14
T cde green, chocolate brown, or olive-brown. Perithecia
uh A ys variable in shape, somewhat elongated or subglobose
untly pointed base, when young yellow, translucent,

Кашу һы obscurely апа remotely septate, minutely roughened
Sep bi : Ке at base rather dark olive-brown with maximum thick-
slightly fle area и, light olive or hyaline at tip, straight or only
Ce wr , or more slender and undulate or even kinked.
eile airs extremely numerous, and intricately interwoven
Ces, a neat, rather compact head, ог in age spreading and
ида в еуеп 10 the substratum, ог to the hairs of nei i
ecia, slender, graceful, without septa, minutely roughened

lue davon hickness and dark at
, dark olive through greater part of length, with tapering and
неа late or kinked. Asci
gularly club-shaped, 8-spored, 64 X 138, pars sporif. 37 и.

h several large, refractive

| Ехятссат!.—Еипе. Gall. XLV, 4438; Klotzsch Herb. Viv.
ус. Х, 959. Sub. Ch. chartarum Ehrb.: Fung. Austro-Ameri-
cani, 193; Fung. Brit. IV, 328; Fung. Gall. XI, 1090; Micro.-

H

199 MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

Fung. Brit. 475; N. A. F. 2nd Series, 1541. Sub Ch. Fieberi
Cda.: Fung. Gall. LXV, 6409; Herb. Myc., Edit. Nova, II, 165;
Myc. Ital. XIII, 1288. Sub Ch. Fieberi Cda., var. chartarum
Roum.: Fung. Gall. LIX, 5827. Sub Ch. Fieberi Cda., f. lignicola
chlorina Sacc.: Myc. Venet. X, 906. Sub Ch. Kunzeanum Zopf:
Fung. Gall. XLV, 4436; Fung. Longob. I, 31; Myc. March.
3246. Sub Ch. lanosum Peck: Fung. Gall. XLV, 4437. Sub
Ch. olivaceum C. & E.: Fung. Columb. VI, 512; Myc. Univ. XX,
1942; N. A. Е. I, 56. Sub Ch. olivaceum С. « E., var. chartarum
Roum.: Fung. Gall. L. 4930.

A very common species of Chaetomium, having been found on
substrata of very varied nature in nearly all countries (Chivers
No. 26). Type locality: Leipzig; on dead stalks and leaves of
various plants.

Probably no species of Chaetomium is of so common occurrence
as Ch. globosum, and certainly no other has been given so much
attention both from a systematic and a morphological point of
view. It was first described and figured by Kunze as a type
form of his new genus, having appeared often in company with
Sphaeria doliolum on dead stalks and leaves of various plants, in
damp wet places in Leipzig, during the spring and autumn. It
was described by that author as a globular fungus, a fourth of a
line in size, black when in fresh condition, and entirely covered
with hairs. His four very simple figures illustrate: a compara-
tively young plant; a later stage when under the influence of
moisture a punctiform opening appears at the crown of the
fungus; a group of somewhat spherical transparent spores which
were mixed with a gelatinous mass; and a cross-section through
the body of the closed fungus.

While, as may be seen, Kunze's description is of a general
and indefinite nature, and his figures are quite inadequate for а
sure identification of the form with which he worked, it seems
reasonable to conclude that it is the same which Zopf described
and figured in his monograph as Ch. Kunzeamum. For reasons
not stated, but presumably on account of the fact that the term
globosum might be applied to any one of several species, Zopf
believed Kunze's form to be poorly named, and considered the
alternative of retaining the old name though unsatisfactory ог of m

MONOGRAPH ОЕ CHAETOMIUM AND ASCOTRICHA 193

giving an entirely new name, in which latter case he would be
disregarding a proper respect of the original author. He finally
named Kunze’s species Ch. Kunzeanum in honor of the founder
of the genus. It would hardly seem that this change was justi-
fied, and in the present monograph, therefore, Zopf's Ch. Kunze-
anum appears as a synonym under Ch. globosum Кге.

In 1818 Ehrenberg added the name Ch. chartarum, the de-
scription of which is indefinite and without figures. No mention
is made of spores and the statements regarding the hairs and
perithecia are of a most general nature. A description and
figures of this species were published by Corda (23) in 1840, and
there seems to be little doubt that the plant in question is iden-
tical with Ch. globosum of Kze. Since Ehrenberg’s time certain
writers have retained the name Ch. chartarum, but as it is reason-
ably certain that Ehrenberg’s species is identical with that of
Kunze, it seems most reasonable to list Ch. chartarum as а synonym
under Ch. globosum Kze.

In 1837 and 1840, in the Icones Fungorum, Corda described
and figured four new species of Chaetomium: Ch. Fieberi, Ch.
afine, Ch. amphitrichum, and Ch. Araliae. The simple, wavy,
undivided hairs and the characteristic spores as described and
figured would indicate that Ch. Fieberi is beyond a doubt identical
with Ch. globosum Kze. The recognition of the three remaining
species is not so easy, however, but it seems entirely possible that
all these are forms of the same plant and should be listed under
Ch. globosum Kze. Corda himself stated that Ch. affine is similar
to Ch. globosum but was to be distinguished from it by the inverted
egg-shaped spores. Zopf listed this in his monograph as а syn-
onym to his Ch. Kunzeanum. Corda stated that Ch. Araliae and
Ch. amphitrichum are distinguished from all other species of the
genus described up to that time by the light color of their eom
but in his diagnoses these spores are given characteristics not
unlike those of other species. In two of the three diagnoses under
consideration he failed to give the sizes of spores, a fact which
makes а sure determination more difficult. If his figures and
descriptions are studied with care it will be found that, with the
exception of the egg-shaped spores of Ch. affine, and slight differ-
ences in the shape of the hairs in the general figures of the three

‚ 194 MonoGRAPH OF CHAETOMIUM AND ASCOTRICHA

species, the forms are very similar, and while it is singular that
he should call these species new unless he felt satisfied that they
possessed sufficient distinguishing characteristics, especially since
he had already named Ch. Fieberi, which has all the characteristics
of Ch. globosum, still it does not seem unreasonable to the present
writer to place these forms as synonyms under Ch. globosum
Kze., on the ground that Corda was dealing with different stages
of the same plant.

In 1876 Dr. Peck redescribed Kunze’s species under the
name Ch. lanosum. Mounts of original specimens which were
received from Dr. Peck show the perithecia and spores typical
in all their characteristics of the younger stages of Ch. globosum
Kze. It may be well at this point to call attention to the fact
that the Ch. lanosum Peck, in Roumeguére’s Fung. Gall. No.
4437, is identical with the plant which Zopf described under the
name Chaetomidium fimeti (Fckl.).

The writer can hardly agree that it is desirable to apply the
name Ch. Fieberi Cda., Ғ. lignicola chlorina to the plant which
Dr. Saccardo found on moist wood, and which he described in
1876, for the form seems in every way to be typical of Ch. glo-
bosum Kze. If one examines young stages of Kunze’s species one
frequently finds hairs which are straight for a considerable por-
tion of their length, then taking a somewhat hooked turn, beyond
which they become wavy. The same hooked characteristic is
mentioned by Saccardo in his original description, but it may
be seen from an examination of original material received from
Dr. Saccardo, and from authentic specimens in Myc. Venet. X,
No. 906, that such is true not of the mature perithecia, where the
hairs are like those of Ch. globosum, but in the young immature
plants. As for the color, it may be said that it is typical of
Kunze’s species, and that the plant was growing on moist wood
seems to be insufficient cause for setting it apart as a variety,
especially since Ch. globosum will thrive on almost any substratum
so long as it is moist.

In 1877 Dr. Cooke described Ch. orientale, mounted material
of which has been placed at the disposal of the writer through the
kindness of the Royal Botanic Gardens at Kew. An examination
of that material shows, in spite of the fact that the perithecia аге

MONOGRAPH ОЕ CHAETOMIUM AND ASCOTRICHA 195

rather fragmentary, characteristics which are those of Ch. glo-
bosum, namely, olive-colored perithecia, simple flexuous hairs,
and globose spores with slightly apiculate ends. The figure
accompanying Cooke’s original description shows the same
characteristics.

In 1878 Cooke and Ellis gave the name Ch. olivaceum to a
form which they found on rotting stems of Erigeron. It was at
first thought by the present writer that this name could be associ-
ated with a species which he has found frequently and which has
apparently been recognized for the first time by Palliser (67) as
distinct from Ch. globosum Kze. This does not seem to be true
however, and from detailed study of the original description and
an examination of mounts made from the type material, one is
forced to conclude that Ch. olivaceum C. & E. is identical with
Ch. globosum Kze. It should be stated that the plants in each
of the specimens of type material in three different herbaria are
past maturity. Dark, rich olive, umbonate spores, and larger,
much more lightly colored, apiculate spores, are to be found in
both Ch. olivaceum and Ch. globosum, the former type appearing
more commonly in Ch. olivaceum, the latter in Ch. globosum. The
hairs in Cooke and Ellis’s material, while old and for the most
part frayed out, are typical of Kunze’s species.

As a result of careful study of type specimens of Ch. macro-
sporum, which was described by Saccardo and Penzig in 1882,
and which has been received from Dr. Saccardo, it has been
found that this species possesses characteristics sufficiently
similar to those of Ch. globosum to warrant placing it as a synonym
under that name. According to Saccardo's description the hairs
of the perithecium are olivaceous and flexuous, but the өрежев
measure 14—16 и. The original material was set apart ue
two parcels; the one containing plants which prove to be typical
of the young Ch. globosum, the other typical mature plants of the
same species. The spores which belong to the mature perithecia
do not have the size indicated by Saccardo, but measure, as for
Ch. globosum Kze., 10.5-12.5й X 9-5 №. There аге, a
intermingled with these plants the perithecia and spores of Ch.
murorum Cda. Here may lie an explanation for the error made

by the original writers, for the spores of Ch. murorum correspond

196 MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

exactly to the measurements recorded in the diagnosis, 14-16 ш
X 7-8 ш. The presence of Ch. murorum Cda. may also explain
the statement that the species is somewhat allied to Ch. crispatum
Fckl, for the perithecia are old and the hairs are very dark,
almost opaque, and except for the circinately coiled tips, have
somewhat lost their true characteristics.

It becomes evident from a study of the original description of
Ch. cymatotrichum made by Dr. Cooke in 1883, and from an
examination of mounts of type material supplied by the Royal
Botanic Gardens at Kew, that this is identical with Ch. globosum
Kze.

Тһе writer has recently received from Dr. Saccardo specimens
of Ch. Fieberi Cda., subspecies Saccardianum, described by
Bommer and Rousseau in 1884 (78), and which according to -
Saccardo (84, p. 86) is equivalent to Ch. Saccardianum. While
the plants are not in the best of condition it has been possible to
determine with a reasonable degree of certainty that this variety
is identical with Ch. globosum Kze. It is stated in the original
description that the perithecia finally collapse and become cup-
shaped. This is true in nearly all of the perithecia examined.
The terminal hairs have been worn away and the shiny, black
perithecial walls have collapsed. It is also stated that the hairs
bear rough spherical conidia. While it is not possible to determine
the origin of these conidia which are scattered among the rhizoids
and perithecial hairs, the writer has no hesitation in concluding
that these are entirely foreign to the Chaetomia, and are the
fruiting bodies most probably of a Cunninghamella. The small
number of perithecia which could be found still clothed with
hairs were typical in every way of Ch. globosum Kze.

In 1884 Bommer and Rousseau (8, p. 207) published a new
variety to Ch. Kunzeanum Zopf under the name fimicola.
These authors compared their variety with Ch. chartarum, Ch.
globosum, and Ch. Fieberi, and while type material is not at
hand, it would seem from their description that the variety could
well be included under Ch. globosum Kze. Marchal (55), refer-
ring to this variety as forma fimicola, called attention to the fact
that the spores, measuring 14 X 8.6 и, were a little larger than in
the species above mentioned and that this plant could not be |

MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA 197

identified with them as Zopf had listed them in his monograph.
The larger size of the spores, however, seems an insufficient reason
for setting apart this variety, for in Ch. Kunzeanum Zopf, the
spores are found to vary in size from 9и to 13и in length and
from 8и to 9.5 и in width.

An examination has been made of mounts from original
material which was described in 1892 by Ellis as Ch. olivaceum
C. & E., var. chartarum, and which had already been listed and
distributed without description by Roumeguére, in Rev. Myc.
II: 130 and Fung. Gall. No. 4930, in 1889, as f. chartarum. The
spores have been measured and found rather to correspond with
the somewhat variable spores of Ch. globosum Kze. As to the
fact that the color varied from nearly black to mouse color and
even to greenish-yellow, it may be said that such variations may
easily be found in one and the same culture, and may be attributed
to a difference in age. It hardly seems advisable to set plants
with such a variable and uncertain characteristic apart from the
type species.

During the same year Beauverie published an account of his
work with cultures of the conidial form which Saccardo (77)
called Oospora crustacea. In the month of June Beauverie found
perithecia in tube cultures of potato which dated from January,
and which were very tightly closed with cotton and covered with
caoutchouc, so that no communication between the media and
the outside air was possible. The white carpet of conidial vege-
tation was found to be spotted with small dark green bodies
which were recognized as Chaetomium, and to which the name
Ch. oospora was given. At two different times the writer has
found conditions similar to those to which Beauverie has called
attention in connection with his work. Cultures which were
supposed to have been pure and had apparently remained uncon-
taminated for a considerable period showed in both cases ES
organization of perithecia which have finally developed into
plants typical of Ch. globosum. While the writer has never
examined type material of Beauverie's form, he is led to believe
from his own experience and from Beauverie's description, that
Ch. oospora is identical with the above named species of Kunze.

While only the original description and figures of Ch. Еазисае

198 МОХОСЕАРН ОЕ CHAETOMIUM AND ASCOTRICHA

published by Koorders (49) in 1907, have been available for
study, the writer has concluded that this plant should be included
under Ch. globosum Kze. In fact at the end of his diagnosis
Koorders has stated that undoubtedly his species is related to
Ch. Kunzeanum, which, as Zopf stated, has a very wide distribu-
tion on various substrata.

Bainier in his * Monograph of Chaetomium” (3) has treated the
varied forms of Ch. globosum Kze. as distinct species, and has listed
and described them under the following names: Ch. megalocarpum,
Ch. setosum, Ch. spirilliferum, and Ch. undulatum. It is the opin-
ion of the present writer that these should not be separated from
the type form, since they prove to be extremely variable when
grown in cultures.

In addition to the above mentioned forms it is possible that
Ch. Douglasii, described by Schweinitz in 1834, and Ch. Fiebert
Cda., var. macropoda, described by Spegazzini in 1898, should
be listed here, but type specimens of these are not available and
the description in both cases is of such a general nature that it is
hardly possible to determine the identity of these forms. They
will be found listed, therefore, among the doubtful species.

The above is as complete an account of the synonymy con-
cerned with this species as the writer is able to give, and it may
be well to state here that he has arrived at the above conclusions
only after long experience in cultivating the variable forms of the
species which have been gathered from nearly all parts of the
world, and only after a thorough study of exsiccati and type
specimens so far as they have been available.

From an examination of the above exsiccati it has been found
that in several the specimens contain plants which belong to
other species and in some cases at least to other genera. In
the following only Ch. elatum Kze. could be found: Ch. chartarum
Ehrb. in Micro.-Fung. Brit. No. 475; Ch. Fieberi Cda. in Fung.
Gall. No. 6409; Ch. Fieberi Cda., var. chartarum Roum. in Fung.
Gall. No. 5827; Ch. Kunzeanum Zopf іп Мус. March. No. 3246,
and Ch. olivaceum C. &. E. in Myc. Univ. No. 1942. Тһе speci-
men under the name Ch. chartarum Ehrb. in F ung. Austro-
Americani, Хо. 193 contains plants of Ch. chartarum (Berk)
Winter, or Ascotricha chartarum according to the nomenclature |

DT

MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA 199

used in this paper. That under the same name in N. A. F.
No. 1541 consists in part of Ch. globosum Kze., of Ch. elatum Kze.,
and of CA. cochliodes Pall. No plants could be obtained from
Ch. chartarum Ehrb. in Fung. Brit. No. 328, while from specimens
under the same name in Fung. Gall. No. 1090 only Stachybotrys
lobulata could be obtained. Тһе specimen under the name
Ch. Fieberi Cda. in Herb. Myc. No. 165 and Myc. Ital. No. 1288
is in part Ch. globosum Kze., and in part Ch. elatum Kze. Only
Ch. murorum Cda. could be obtained from mounts of Ch. Kunse-
anum Zopf, in Fung. Gall. No. 4436, while the specimens under
the name CZ. lanosum Peck, in Fung. Gall. No. 4437 consist for
the most part of Chaetomidium fimeti (Fckl.) Zopf, with a con-
siderable number of perithecia of Ch. elatum Kze. intermingled.
Ch. olivaceum C. & E. in Fung. Columb. No. 512 is found to
consist partly of Ch. globosum Kze., of Ch. elatum Kze., and in
part of Chaetomidium fimeti (Fckl.) Zopf.

21. CHAETOMIUM SPIRALE Zopf, Nova Acta Leop. Carol. Akad.
42: 275. pl. 19, f. 21-26. 1881
Chaetomium Spirochaete Palliser, N. A. Flora 3:61. 1910.

PLATE 12, FIGS. 5-8

Dark brown to black. Perithecia of medium size 150-300 и,
globose or ovate with a bluntly pointed base, seated on dark
olive-yellow to brown rhizoids. Lateral hairs long, graceful,
nearly straight or slightly flexed, very gradually tapering toward
the tip, septate throughout, at base 3-5.5 u in thickness, dark
olive-brown, sometimes smooth but more frequently roughened
by irregular hyaline bodies of varying size and shape, becoming
smooth above and fading to a colorless or pale yellow tip. Ter-
minal hairs sparsely septate, dark, rich olive-brown, roughened
by minute spines and warts, slightly paler and somewhat less
roughened near the tips, straight or only slightly bent below for
300-370 u of their length, 4-6 in thickness at base, spirally
coiled above with 6-14 turns. “Asci club-shaped, with a short
stalk, pars sporif. 34-43 и long." Spores lemon-shaped, slightly
apiculate at either end or irregularly oval or spherical, dark
rich olive-yellow to olive-brown, 9 X 7 и (6-12 X 5.6-9), when
seen edgewise, 5.5-7 и broad.

The species above described is apparently not common, and
has never appeared іп any collections made by the writer. Туре
locality: near Berlin; on horse dung.

200 MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

Through the kindness of the New York Botanical Garden the
writer has been enabled to examine type specimens of Ch. spiro-
chaete which were collected on cotton roots at Ames, Iowa, by
L. H. Pammel, and which were named Ch. spirochaete by Palliser
in her monograph of the genus. These seem to be identical with
Zopf's species. The envelope containing the type specimens
bears the name Ch. bostrychodes Zopf, for which this material
had previously been mistaken. On a slip of paper within the
envelope, however, was written in the form of a note: ‘‘Spores
10 X 7u, dark brown, too large for bostrychodes although same
shape.” The large size and dark color of the spores are charac-
teristics which easily separate this form from Ch. bostrychodes
Zopf, and identify it with Ch. spirale Zopf.

At first sight Ch. spirale Zopf might easily be confused with
Ch. bostrychodes Zopf, which it so closely resembles so far as the
hairs are concerned, but from which it differs markedly in spore
characteristics. The characteristics of the asci are quoted from
Zopf's original description.

22. CHAETOMIUM ATERRIMUM Ellis & Everhart; Palliser, N. А.
Flora 3: 62. 1910
PLATE I2, FIGS. 1—4

Gray black to black. Perithecium of medium size, ovate or
subglobose, 275 X 200 u (190-300 X 160-230), seated on olive-
yellow or olive-brown rhizoids. Lateral hairs numerous, long,
slender, graceful, straight or slightly flexed, septate, gradually
tapering, minutely roughened throughout or only near the base,
below olive-yellow or dark olive-brown, 4-5 и in thickness at
base, above pale yellow or hyaline. Terminal hairs very rarely
producing small abortive branches, opaque, olive-brown or black
and roughened by minute spines throughout, irregularly septate,
at base 5-6 иіп thickness, at tip 8-11 шіп thickness, below straight
or slightly flexed, above coiling 10-18 times (usually about 15)
in a long, close, regular, cylindrical spiral, 45-60 и in diameter.
Asci not visible. Spores olive-brown, lemon-shaped, slightly
apiculate at either end, 7.5 X Dan (6.5-8 X 6.4), when seen
edgewise, compressed, 4-8 и broad.

This is ап exceedingly rare form apparently known only from
the type locality: Rockport, Kansas; on damaged wheat іп а
stack, 1891 (Bartholomew 448).

MONOGRAPH ОҒ CHAETOMIUM AND ASCOTRICHA 201

Through the kindness of the New York Botanical Garden it
has been possible to examine type specimens of this interesting
species. The large diameter of the terminal hairs and their
exceedingly long and regular cylindrical spiral distinguish this
from all other species.

23. CHAETOMIUM BOSTRYCHODES Zopf, Abh. Bot. Ver. Prov.
Brandenburg 19: 173; 1877. Nova Acta Acad. Leop.-Carol.
42: 277. pl. 20, f. 14-26. 1881
Chaetomium caninum Ellis & Everhart, Jour. Myc. 4: 79. 1888.
Chaetomium formosum Bainier, Bull. Soc. Myc. France 25: 215.

pl. 20, f. 6-9. 1910.
Chaetomium formosum, хат. neglectum Вашег, 1. с. 25: 217.

pl. та, Т. 8. 1910.
Chaetomium formosum, хат. ovatum, Bainier 1. с. 25: 216. pl. 20,

f 1-5. 1910.

PLATE 13, FIGS. I-II

Steel gray. Perithecia of medium size, extremely variable
in shape, broadly ovate, globose or nearly cylindrical, generally
with a bluntly pointed base, 340 X 220 и (168-350 X 131-230),
frequently provided with black, straight or re-curved cirrhi.

teral hairs not numerous, encrusted, clearly and evenly septate,
tapering, at base dark olive-brown and about 3.8 и in thickness,
at tips pale yellow or hyaline, frequently collapsed. Terminal
hairs encrusted and roughened with spine-like projections through-
out, at base straight or very slightly flexed, dark olive-brown to
black and about 44 in thickness, slightly less colored at tips,
always more or less spirally coiled but in this respect extremely
varied. In the type either regularly coiled with seldom more than

in turn are septate and spirally coiled. Asci short, stout, club-
shaped, 8-spored, 50 X 12 и, pars sporif. 24и. Spores when
young greenish, hyaline, with granular contents, when mature
pale with olive-brown tint, oval to nearly spherical, clearly or

лгі only in this species, 74 X hi бу АРТА» um
seen edgewise, compressed, 4.8 и broad.

Exsiccatt.—Fung. Europ. Edit. Nova, Series П, XXXIV,
3340; Myc. March. No. 43. |

209 MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

An extremely common species having appeared in cultures of
dung of many animals from nearly all parts of New England,
Florida, Chili, Venezuela, and China (Chivers No. 21). Ке-
ported also from many localities in middle western and western
United States, and from England and Germany. Туре locality,—
Berlin and Góttingen; on decaying animal and vegetable sub-
stances (carcasses, potatoes, and rabbit dung).

In 1888 Ellis and Everhart (31) gave the name Ch. caninum
to material found on dog dung in Louisiana. Later, however,
the same authors (32) listed this name аз a synonym under Ch.
bostrychodes Zopf.

Ch. bostrychodes has proved to be a most variable species and
before extensive cultures had been made the separation of the
variable forms into distinct species seemed possible. А careful
examination of plants grown under cultivation has been sufficient,
however, to convince the writer that this should not be done,
for there is no line which can be drawn between one variable form
and another. While no material is available of Ch. formosum,
Ch. formosum, var. neglectum and Ch. formosum, var. ovatum, all
of which were described by Bainier in his monograph of Chae-
tomium, the writer is strongly of the opinion that they should be
considered rather as variations of the type form.

he specimen in Fung. Europ. No. 3340 has been examined
and found to be typical of Ch. bostrychodes Zopf.

24. CHAETOMIUM QUADRANGULATUM Chivers, Proc. Am. Acad.
48: 85. 1012
PLATE 14, FIGS. 1-8

Gray. Perithecia rather large and elongated, barrel-shaped,
403 X 294 и (333-456 X 243-350), frequently provided with one
or more very long cirrhi, producing a mass of dark olive rhizoids
near base. Lateral hairs numerous, slender, straight, regularly
and distinctly septate, at the base dark olive to black, minutely
roughened and about 7.5 шіп thickness, in upper part pale yellow
or hyaline, smooth. "Terminal hairs of two types; (a) unbranched,
minutely roughened, below dark olive-brown to black, straight,
septate, above coiling in the form of a spiral with 2-6 convolu-
tions, near tips light olive-yellow or colorless, sparsely septate;
(0) showing tendency to twist in spiral fashion near the middle of «
their length, or curved or sometimes nearly straight throughout,

MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA 203

many with a single coil near the middle of their length, finely
roughened throughout, below dark olive-brown to black, and
about 7.5 и in thickness, clearly septate to near tips which are
hyaline (along the hairs of this nature several branches may be
produced, the branches partaking of the nature of the hairs).
Asci club-shaped, 8-spored, 39 Х 9.7 и, pars sporif. 21 и. Spores
when young greenish, hyaline with small refractive globules,
when mature pale olive, when seen in face view four-sided and
four-angled, either nearly square with equally rounded corners or
with depressions in the four sides, in which latter case there are
generally two acute angles at opposite ends which mark the length
of the spore, while the angles at the sides are broad and obtuse,
7.3 X 6.3 и (6.4-8 X 5.6-6.4), when seen edgewise ovate, 3-4 и
broad. А

Cultivated on dung from Cambridge, Massachusetts (Chivers
No.29). Appearing also on dung from Chile and from Little Swan
Island, Gulf of Mexico.

This species may be easily identified by its spores, which when
seen in face view are four-sided and four-angled, but when seen
in profile are oval. Ch. quadrangulatum and Ch. trigonosporum
are the only species known to the writer which possess spores
with angles, the former having spores clearly quadrangular, the
latter clearly triangular.

25. CHAETOMIUM CAPRINUM Bainier, Bull. Soc. Myc. France 25:
223. pl. 24. 1910
PLATE 13, FIGS. 18-22

eel gray. Perithecia large, tall, greatly elongated with
greatest width just above the base, then narrowing wit the
upper portion cylindrical, 580 X 2154 (400-580 X 200-228),
seated on a subiculum of dark olive-brown to black rhizoids, with
perithecium wall composed of cells greatly elongated in а plane
parallel to the long axis of the perithecium, often provided with
short, stout, black cirrhi. Lateral hairs rather numerous, tapering,
septate, smooth or minutely roughened with spines, some uncuti-
cularized, colorless and easily collapsible, others stoutet, at base
dark olive-brown and about 6.6 шіп thickness, fading and tapering
gradually to a colorless, collapsed tip. Terminal hairs almost
entirely obscured at maturity by the mass of spores, smooth or
remo tely septate,
at base straight or flexed, dark olive-brown, about 7.5 # 1n thick-
ness, fading toward tip, coiling above irregularly or in the form

204 MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

of a spiral, producing here and there along their convolutions short-
branches which in turn are septate, tapering and irregularly coiled.
Asch club-shaped, 8-врогей, 50 X 10 и, pars sporif. 24 и. Spores
when young greenish, hyaline, filled with granular contents, when
mature pale with slight olive tint, ovate to globose, sometimes
minutely apiculate at both ends, 6.6 X 5.1 и (5.6-8.1 X 4.8-5.6),
when seen edgewise, compressed, 4.5 ш broad.

On muskrat dung, Massachusetts; rat dung, Liberia, Herb. R.
Thaxter; on dog dung, Connecticut (Chivers No. 20). Type
locality: Paris(?); on goat and sheep dung.

It is possible that this species may be identical with Ch.
rostratum, described by Spegazzini in 1899, but type specimens
of his form are not available and it is impossible to decide with
certainty from a study of the original description. Spegazzini’s
species has been listed, therefore, among the doubtful forms.

The species may be easily distinguished from Ch. bostrychodes
Zopf, which it most nearly resembles, by the extreme length of
the perithecium, a characteristic which has proved constant
through cultures including many generations.

26. CHAETOMIUM COCHLIODES Palliser, N. A. Flora 3: 61. 1910”
Chaetomium flexuosum Palliser, N. A. Flora 3: 61. 1910.
PLATE 15, FIGS. 1-8

Grayish-green, brilliant green, often losing color and becoming
dark brown in dry herbarium material. Perithecia globose ог
subglobose with bluntly pointed base, 340 X 295 и (318-360 Х
273-310), forming black cirrhi in comparatively few cases only,
producing a heavy mass of stout, dark olive-brown to blac
rhizoids which anchor the plants firmly to the substratum. Lat:
eral hairs numerous, irregularly and sparsely septate, evenly rough-
ened with extremely fine projections, some hairs rather stout,
tapering, straight or evenly bent, at base about 5.3 шіп thickness
and dark olive-brown to black, above faded yellow, at tips fre-
quently hyaline, other hairs loosely and irregularly twisted
through their entire length, still others straight for a long distance,
then spirally twisted. Terminal hairs very numerous and finally
interwoven forming a thick, massive, shaggy head, always of
two types; (a) thick, stout, projecting beyond the dense portion
of the head, then becoming spirally coiled with about three ОГ `
four convolutions, dark olive-brown, almost black below, fading |
above to light yellow or becoming colorless, evenly roughened — |
throughout and without visible septa, at base about 7.54 E

MoNoGRAPH OF CHAETOMIUM AND ASCOTRICHA 205

thickness; (b) slender, sometimes coiling in spirals, at other times
irregularly twisted or undulate, medium olive to light yellow,
lighter than those of type (a), about 2.5 и in thickness at base.
Asci irregularly club-shaped, 8-spored, 88 X Іт џ, pars sporif.
32 и. Spores when young colorless, filled with refractive globules,
when mature dark, rich olive-brown, lemon-shaped, apiculate
at both ends, 9.5 X 7.2 и (8.9-9.7 X 6.4-8.4), when seen edge-
wise, compressed, 6.4 и broad.

ExsiccATI.—Sub Ch. chartarum Ehrb.: М. A. Е. 1541 (in part).

On paper, straw, dung and leaves from many localities in
New England. Also from middle western and southern United
States, Java and Ceylon (Chivers No. 24). Type locality: New-
field, New Jersey; on paper (Ellis & Everhart in N. A. F. No.
I541 in part.)

Type specimens of the plant which Palliser in her monograph
named CA. cochliodes and which are to be found in North Ameri-
can Fungi, No. 1541, under the name Ch. chartarum Ehrb. have
been examined by the writer. It has been discovered for the
first time by Palliser that this is a separate species to be set
apart from Ch. globosum Kze. It has been found frequently
and, while its color as well as its general appearance at maturity
may resemble Ch. globosum Kze., careful study іп cultures and
during its earlier development furnishes sufficient evidence that
it is to be reckoned with as a distinct species.

It seems hardly advisable to consider Ch. flexuosum, which
was described by Palliser (65) іп 1910, a distinct species, but
rather as a dilapidated condition of Ch. cochliodes. Тһе type
specimens of this plant have been found to be in extremely poor
condition. Through the kindness of Professor Kauffman and

r. F. J. Seaver, the writer has been provided with sufficient
material of the original gathering to enable him to arrive at
the conclusion that the plant is none other than Ch. cochliodes.
Mounts of such material show, in the first place, that the pant
are old and much weather-beaten. In most cases perithecia
bear only rhizoidal hairs and stumps of the lateral and terminal
hairs. One perithecium which was most perfect of all available
shows its top to be covered with a large number of hairs which
are stout, straight for a considerable distance, then loosely spirally
coiled once or twice, at which point by far the largest number are

206 MONOGRAPH ОЕ CHAETOMIUM AND ASCOTRICHA

broken. There seems to be every reason to believe that the hairs,
had not so many been broken away by disintegration, are sufficient
in number and suitable in characteristics to make up a head typi-
cal of Ch. cochliodes. The spores are also typical of that species,

Ch. cochliodes Palliser may be confused, especially when fully
developed, with Ch. globosum Kze. It may be most easily dis-
tinguished, in its earlier life history, when the terminal hairs show
clearly two types; the one straight, stout below, spirally coiled
above; the other much more slender, loosely spirally coiled,
twisted or only wavy, and entangling the stouter hairs.

27. Chaetomium angustum sp. nov.
PLATE 16, FIGS. 6-І2

Rich olive-green. Perithecia rather large, subglobose, with a
bluntly pointed base, 304 X 266 и (280-300 X 240-290), forming
cirrhi in comparatively few cases, producing a dense mass 0
dark olive-brown to black, undulate rhizoids. Lateral hairs
very numerous; obscurely and sparsely septate, either stout,
tapering, dark, olive-brown to black at base and about 6.6 in
thickness, fading above and becoming pale yellow, hyaline at
tips, or slender, olive-yellow and flexed or irregularly, spirally
twisted. Terminal hairs of three types: (a) straight, long, taper-
ing, dark olive-brown to black at base, light yellow to hyaline
at tips, minutely roughened throughout with spines, obscurely
septate or continuous, at base about 6.7 шіп thickness; (0) slender,
graceful, below straight or only slightly flexed and about 3.8 и in
thickness, in upper portions coiling spirally with about 5-7 con-
volutions which are comparatively small in diameter, minutely
roughened throughout, sparingly septate or continuous, at base
dark olive-brown, fading toward the tips; (c) stout, coarse,
roughened throughout, sparingly and obscurely septate, below
straight, very dark olive-brown, at base about 7.5 и in thickness,
above spirally coiled at first regularly coiled with 2-3 convolutions,
often becoming irregular and retaining only a single coil near the
middle of their length from which, frequently, one or two branches
arise. Asci irregularly club-shaped, 8-spored, 50 X 14 и, pars `
sporif. 23 u. Spores when young greenish, hyaline, filled with
irregular, refractive globules, when mature dark olive-brown,
чеге бирд кы apiculate or umbonate, 10 X 8и (9.7-10.5 X 7-37

On culture of bat dung from Porto Rico (Chivers No. 25).
This species may be confused with Ch. cochliodes Palliser, ОП

MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA 207

the one hand, and on the other hand with CA. globosum Kze.
From the former it differs in its straight terminal hairs, and in
the fact that the terminal hairs never form a dense head. From
the latter it differs also in its loose head of terminal hairs and in
the fact that it possesses straight and stout, coarse, spirally coiled
terminal hairs.

28. CHAETOMIUM TORULOsUM Bainier, Bull. Soc. Мус. France
25: 224. Ph 23. 1910
PLATE 16, FIGS. 1-5

Golden-yellow. Perithecia of medium size, rather tall and
elongated, vase-shaped, 355 X 170 (290-140 X 150-187), fre-
quently provided with short, stout, blue-black cirrhi. Lateral
hairs numerous, varying in length, delicate, smooth, generally
slightly curved or bent, clearly and regularly septate, near base
dark olive, about 3.7 и in thickness, gradually fading above to
yellow and ending in extremely slender, colorless, crumpled tips.
Terminal hairs almost entirely concealed at maturity by the spore
mass, long, slender, graceful, irregularly flexed, clearly and
regularly septate to near the tips, conspicuously constricted at,
and inflated between, the septa, dark olive-brown at base, about
3.8 и in thickness, very gradually fading and tapering for a long
distance, terminating in a long colorless, refractive thread. Asci
irregularly club-shaped, 8-spored, 40 X IO y, pars sporif. 18 и.
Spores when young hyaline, filled with refractive globules, when
mature very dark, rich olive-brown, lemon-shaped, apiculate,
extremely variable in size, 9 X 7.6 ы (73-113 Х 6.4-9), when
seen edgewise, compressed, 4.5 и broad.

In cultures of old paper, muskrat, rat and rabbit dung from
various localities in New England (Chivers No.5). Type locality:
Paris(?). i

This species may be easily distinguished from Ch. ampullare,
which it most nearly resembles, by the fact that the perithecia
are less elongated, the lateral hairs much more numerous and
shorter, and the terminal hairs less rigid and conspicuously con-
stricted at the septa.

Species DUBIAE
Unfortunately it has been impossible to identify certain forms
which have been described from time to time, on account of the
fact that their characteristics are not given in sufficient detail

208 MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

and no type material is at hand for examination. Since the
present writer cannot speak with certainty regarding these. it
has seemed best to list them as doubtful species.

In 1834 Schweinitz (91) described Ch. Douglas as a very
remarkable fungus allied to Sphaeria comata. In the herbarium
of Curtis, now at Harvard University, there is a reference to this
species and a very simple figure of it, but no specimen. Sac-
cardo, in the Sylloge (p. 229), has listed this as a doubtful form.
From a study of the very general description it seems possible
that this may be identical with Ch. globosum of Kunze.

Ch. lanatum, which was described by Quelét (70) in 1876 and
to which reference has already been made, may possibly be
identical with Ch. globosum Kze. Quelét’s description is of а
very general nature, but his figures though simple show sinuous
hairs like those of Ch. globosum, and spores similar in shape to
those of the same species.

Little is known of the form which Spegazzini (94) found on
dog dung in Italy, and which he described in 1878 as Ch. ster-
coreum. Іп the following year Saccardo (74) figured this and in
1894 Cavara reported it, but without further comment.

In 1881 Spegazzini (95) contributed a new form under the
name Ch. microsporum. While no type specimens of this species
are at hand it would seem that it might be identical with. Ch.
globosum Kze., with the exception that the spores, which measure
5-6.5 X 4-5 и, are smaller than in Kunze's species.

It is impossible to determine from Karsten's (48) description,
even though reasonably complete, the exact nature of the species
which he described in 1888 under the name Ch. humanum, and
so far as can be learned no author has since found or identified it.

The writer has little knowledge of the form which Starbáck
(97) described in 1889 as Ch. discolor, and no specimens of the
original material are at hand. It would seem, however, from
a study of the description and diagrammatic figures, that with the
exception of the smaller size of the spores the plant in question
might be Ch. globosum Kze.

The original description and two general figures of Ch. varium
described by Delacroix (25) in 1897 recall the characteristics of 1
Ch. globosum Kze., but on the other hand figure 4 represents Ше | |

MONOGRAPH ОЕ CHAETOMIUM AND ASCOTRICHA 209

type of branching which is found in Ch. elatum Kze. The spores
as figured are equally characteristic of either species. In a foot-
note Delacroix stated that the plant was allied to Ch. lanosum Peck,
and to Ch. comatum (Tode) Fr., but such a condition could hardly
be possible since those species differ so widely.

Reference has already been made to the possible identity
of Ch. Fieberi var. macropoda, described by Spegazzini (96,
р. 250) іп 1898, and Ch. globosum Kze. Saccardo (85) ге-
described this variety giving a much clearer diagnosis, and here
again there is marked correspondence with Kunze’s species.
The only variation seems to lie in the spores which, as described
by Saccardo, are slightly narrower than those of Ch. globosum.
Since original specimens are not at hand it is impossible to come
to a satisfactory conclusion regarding this variety.

It has already been pointed out that Ch. rostratum, published
by Spegazzini (96) in 1899, may possibly be identical with Ch.
caprinum Bainier, but since Spegazzini's description is not entirely
clear, and no type material is at hand, it is impossible to decide
the matter with certainty. This species has been reported by
A. P. Morgan (62), who found it on rabbit dung at Preston, Ohio,
and who kindly forwarded a specimen of his collection to the
present writer. Тһе perithecia obtainable from the specimens
were very much broken, however, and only their most general
characteristics could be made out.

SPECIES EXCLUDENDAE

As a result of a thorough examination of the literature con-
cerning described species of Chaetomium, and notes published in
connection with exsiccati, a considerable number of names have
been found which should be excluded from further use. In some
cases the plants to which the names were originally applied have
apparently never been described, and there is every reason to
believe that their characteristics will never be clearly understood.
In other cases the published account is so brief and inadequate
that it is impossible to arrive at a satisfactory conclusion regarding
the form in question, while the descriptions of several other species
indicate clearly that they are not Chaetomia. The following list
includes all names whose further use should be discontinued for
the reasons above mentioned.

210 MONOGRAPH ОҒ CHAETOMIUM AND ASCOTRICHA

In 1818 Ehrenberg (28, pp. 15, 27) contributed a new species,
Ch. gelatinosum, which, according to Zopf (113, p. 204), is either
an undeveloped condition of some other Chaetomium, or more
probably a Myxotrichum, and which is placed by Saccardo (79)
among his doubtful species. The original description, containing
only a few of the most general statements, and with no accompany-
ing figures, is entirely inadequate.

Fries (38) in describing a new species in 1829 under the name
Ch. pusillum, overlooked one of the most salient characteristics
of the genus pointed out by Kunze, namely,—the presence of an
ostiole. This plant, possessing a minute, spherical, closed peri-
thecium, with very short, stiff, opaque, bristle-like hairs, and
producing spores which are at least one-septate, could hardly be
classified as a Chaetomium. The same form has been observed
by many authors since Fries’s time, and has been given names as
follows: Acanthostigma Chaetomium Auersw. (1); Caelosphaeria
exilis (Alb. et Schw.) басс. (79); Niesslia Chaetomium (Cda.)
Auersw. (1); Niesslia exilis (Alb. & Schw.) Wint. (110); Niessha
pusilla (Fries) Schroeter (89); Nitschkia exilis (Alb. & Schw.)
Fckl. (42, p. 165); Peziza aterima Lasch; Sphaeria Chaetomium Cda.
(21); Sphaeria exilis Alb. & Schw. (2, p. 44); Sphaeria exospori-
oides Desm. (26, No. 126); Venturia Chaetomium (Cda.) Ces. &
DeNot. (13). The present writer has seen authentic specimens
of Ch. pusillum, which were distributed by Fries in the Sclero-
myceti Sueciae XXVIII No. 272, and has found them to consist
of small, black, naked pustules, not in the least resembling Chae-
tomia, scattered over the surface of the pine needles. Specimens
under the same name and with the same characteristics have been
distributed by Rehm in Ascomyceten No. 1762.

In 1833 Wallroth published, in the Flora Cryptogamica (109),
eight new species which must be placed in the group of excluded
names. They are as follows: Ch. Alchemillae, Ch. circinans, Ch.
coccodes, Ch. depressum, Ch. epiphyllum, Ch. oxysporium, Ch.
Potentillae, and Ch. strigosum.

. Майго’ original description of Ch. Alchemillae (109, р. 873)
is incomplete and leaves one uncertain regarding the exact
nature of the form. That part of the description which is clear,

however, would seem to indicate that the plant was a Ven-

MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA 211

turia. Saccardo (79, 84) has recorded it under the name Venturia
Alchemillae (Grev.) B. & Br., while other authors have given
names as follows: Asteroma Alchemillae Grev. (37, 44); Coleroa
Alchemillae (Grev.) Wint. (110, р. 199); Dothidea Alchemillae
Rabh.; Dothidea ceramioides Duby (27); Stigmatea Alchemillae Fr.
(39, р- 423).

The plant to which Wallroth (109, р. 266) gave the name Ch.
circinans is clearly not a Chaetomium. Saccardo (75, 79) has
given to this form the name Venturia Kunzei, while other authors
have applied the following names: Coleroa Chaetomium Rabh.
(71, No. 1456); Dothidea Chaetomium Kze. (51); Stigmatea Chae-
tomium (Kze.) Fr. (39; p. 422).

The original description of Ch. coccodes, as given by Wallroth
(109, p. 265), is very inadequate and it seems impossible to arrive
at any satisfactory conclusion regarding it. Zopf (113, p. 205) has
stated without further explanation that it probably represents a
Chaetomium, but is rather to be considered as a slightly developed
stage probably of Ch. crispatum Fckl.

Regarding Ch. depressum of Wallroth (109, р. 266) it may be
said that the original description is hardly that of a Chaetomium,
but rather of an Excipula, since the perithecia are described as
depressed-globose, sometimes oval, rough at the base, whence
arise short, rigid, black hairs, exposing at the apex, where they are
wide open, a disc of sporophores. |

The original diagnosis of СЁ. epiphyllum Kze. appeared among
the species of Wallroth (109, p. 265) mentioned above, as a note
rather than as a description. The name was used by Kunze
in connection with specimens which he enclosed in a letter to
Wallroth, but so far as can be learned the species has never been
described. Specimens of this species distributed in Klotzsch.
Herb. Myc. No. 1347 are found to consist of minute, smooth,
black pustules.

Ch. oxysporium Wallr. (109, р. 242) is merely mentioned by Wall-
roth in a description of a plant which he called Fusarium Chae-
tomium. It is a name which he himself had privately used
for this form at an earlier date, but which he had never published.

The original description of Ch. Potentillae Wallr. (тод, р. 266),
while of a brief and general nature, would hardly indicate a true

212 MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

Chaetomium. The perithecia were described as minute, black,
with short, rigid hairs. Saccardo (79, р. 594) has listed this
form under the name Venturia Potentillae (Fr.) Cke. (18). Other
authors have placed it under the following names: Coleroa Poten-
tillae (Fr.) Wint. (110, p. 199); Dothidea Potentillae Fr. (37, p.
563); Stigmatea Potentillae Fr. (39, р. 422).

The very brief diagnosis of Ch. strigosum Wallr. (109, p. 265)
is vague and insufficient, and unaccompanied by figures. Raben-
horst (71, Хо. 1309) has called this Ceuthospora phaeocomes.

Schweinitz (91, p. 265) in 1834 added the new name Ch.
Typhae to the genus, but the original description is scarcely more
than a note. Type specimens from the Curtis herbarium, now
at Harvard University, have been examined by the writer, but
no material could be found on the stalks of the Typha. Con-
sidering the meagre description, there seems to be little reason
for the continued use of this name or of the name Ch. Typhinum,
which Schweinitz (91, p. 310) later used for this species.

In 1845 Léveillé (52) published diagnoses of two supposedly
new forms; Ch. Cumingii, which he found on fallen leaves near
Manila, and Ch. viride, on fallen grasses in Paraguay. The
diagnoses, unaccompanied by figures of any kind, are brief and
of such a general nature that one is unable to gain any satis-
factory knowledge regarding the real nature of the plants in
question. It would seem that the use of both names may well
be discontinued.

The description of Ch. hispidum, published by Fries (39.
р. 405) іп 1849, is brief, calling attention only to the most general
characteristics and to the hemispherical shape of the perithecium.
No measurements of structures are given and it is impossible to
arrive at a satisfactory conclusion regarding his material. The
fact that the asci are four-spored would, in any case, exclude
this species from Chaetomium.

In 1851 Bonorden (9) described Ch. ciliatum and Raben-
horst published in the Bot. Zeitung (9: 453) a description of
Ch. Braunii. In the same year Preuss (69) described Ch. concin-
natum and Ch. tomentosum, and in 1852 added still another
name, Ch. signatum.

It may be seen from a study of Bonorden's description that the

MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA 213

plants which he called Ch. ciliatum are not Chaetomia, since the
spores are cylindrical and provided at both ends with delicate
cilia. Saccardo in the Sylloge Fungorum (3: 684) has given this
plant a place under the name Dinemasporium ciliatum (Bon.).

Sections of the authentic material of Ch. Braunii Rabh., in
Klotzsch. Herb. Myc. No. 1554, show the fruiting bodies to be
stromatic, cellular, passing far down below the surface of the host
tissue, black and coriaceous, covered with short, black, spine-like
hairs.

The three species, Ch. concinnatum, Ch. tomentosum, and Ch.
signatum, are described only in a most general way without meas-
urements or illustrations. Rabenhorst, in Linnaea (24: 144), has
questioned whether Ch. tomentosum is sufficiently different from
Ch. pannosum Wallr., and has stated that in his specimen the hairs
are soft, not rigid. A study of Rabenhorst’s Ch. tomentosum
in Klotzsch. Herb. Myc. No. 1856 shows his plants to be Ch.
murorum Cda. No author, with the exception of Rabenhorst,
has ever recorded the re-appearance of these forms and it would
seem that the use of these names could well be discontinued.

In 1853 Strauss (99) published a diagnosis of what he con-
sidered а new species of Chaetomium under the name Ch. nivale,
and Montagne (бі) in 1856 added the name Ch. raripilum. The
original description of Ch. nivale, together with the figures
which Strauss published in connection with it, furnish sufficient
evidence that he was dealing with a genus other than Chaetomium.
This plant has been listed by Saccardo (80, р. 855) under the
name Acanthostigma nivale (Str.). It is also evident from. a
study of Montagne's description that his plant is not a species
of our genus. It is given a place by Saccardo in the third volume
of the Sylloge (p. 322), under the name Chaetomella raripila
(Mont 3.

The description and simple figure of Ch. fimeti, published by
Fuckel (40, p. 491) in 1861, are those of the plant which Zopf
(113, p. 280) described and figured under the name Chaetomidium
fimeti, and which was still later described and figured by Вапиег
(3, p. 192) under the same name. Exsiccati specimens of this
form are to be found in Rehm Ascom. Хо. 991, and Fung. Sax.
No. 370.

214 MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

It has been possible to study type specimens in Fung. Rhenani
No. 1572, of Ch. paucisetum, which was named by Fuckel (41)
in 1866. The perithecia have been found to be globose, closed,
black, firmly adnate to the surface of the substratum, and naked
or bearing only a few short, scattered, slender, spine-like hairs.
In Die Pilze Deutschlands (110, p. 65) this form is given the name
Chaetomella atra (Fckl.).

In 1873 Berkeley and Broome (7) described Ch. glabrum
and Ch. rufulum. Тһе figures and descriptions of the first
named species assure one that the plant is not a Chaetomium.
It has been described by Cooke (19) as Orbicula perichaenoides,
and by Saccardo as Anixia perichaenoides (79, p. 35). So far as
Ch. rufulum is concerned there is no indication, either in the
description or the figures, of the presence of perithecial hairs.
The asci and the rough, spherical spores are not typical of Chae-
tomium.

Through the kindness of Dr. Saccardo it has become possi-
ble to examine type specimens of Ch. calvescens (76), described by
that author in 1878. The specimens, however, are in such a con-
dition that it is impossible to identify the plant in question
with a great degree of accuracy. Only in one perithecium of the
many which were mounted have any hairs been found, and in
that case they are few and rather stiff and spine-like. In one
perithecium the remains of a neck like that of Melanospora
appears. The honey-yellow color of the apparently mature
spores would indicate that the form is not a Chaetomium. In
1882, it was listed by its author (79, p. 227) under ''Species
Desciscentes."' +

Karsten (47) іп 1881 described a plant under the name Ch.
Лтзефит which produced many spores in the ascus. No species
typical of Chaetomium have been found to contain more or less
than eight spores in their asci, and it seems reasonable to conclude
that forms with a varying number of spores should be listed in
another genus.

In 1882 Rehm (73) added the new name Ch. Polypori to the
genus, and while authentic specimens are not at hand the original
description indicates that these plants are not Chaetomia since
the perithecia are minute, and clothed with very short, acute

MONOGRAPH ОҒ CHAETOMIUM AND ASCOTRICHA 215

setae. The asci are cylindrical and contain subrotund or spherical
spores arranged in a single row, and filiform paraphyses are
produced among the asci.

During the following year 1883, Roumeguére and Patouillard,
in Rev. Myc. (5: 29), applied the name Therryana to a
variety of Ch. atrum Link. A study of the original description
and the accompanying figures together with an examination of
authentic specimens in Fung. Gall. No. 2496, are sufficient to
convince the writer that this variety has nothing in common
with Ch. atrum but is identical with Chaetomidium fimeti (Fckl.)
Zopf. During the same year Therry (102) made mention of Ch.
exile, and whether he used this name by mistake or whether he
purposely used it for a species which he intended to describe later,
is uncertain. In either case it would seem advisable to exclude
this, since, so far as can be learned, no reference has since been
made to it, and no description has been published.

In 1888 Roumeguére gave to a plant in Fung. Gall. No. 4438,
the name Ch. globosum Kze., f. chartarum. The variety was
not described, however, and so far as can be learned no mention
or description has since appeared. From a study of mounted
material from the authentic specimens, this plant is found to be
identical with Chaetomidium fimeti (Fckl.) Zopf. :

In 1892 Dr. Cavara (12) described under the name Ch. Monte-
martinii a form which has characteristics similar to those of
СЪ. Сабли: Sace. Ва the cam, however mounts of the
type material received from Dr. Saccardo, and also of authentic
exsiccati specimens in Fung. Longob. No. 228, show more clearly
the exact nature of the plant. The perithecia, naked or scantily
clothed with a few very slender, short hairs, are globose, flattened
at the base, while from the top extends a cylindrical, curved
neck, 70-80 in length. At the top of the neck an ostiole 1s
located. In spite of several attempts to obtain asci none have
been found. The honey-yellow spores which are produced in
the perithecium and which in mass possess à decided orange
tinge are poured from the ostiole in enormous numbers whenever
pressure is brought to bear on the perithecium. From these
observations it may be seen that this plant is not a Chaetomium.
In 1894 Ellis and Everhart (34) described as new Ch. palli-

216 MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

dum, and through the kindness of Mr. J. Dearness the study of
this form from original material has been made possible. This
plant seems to possess characteristics entirely in common with
those of Melanospora, and ought therefore, as suggested by Palliser
in the North American Flora (65, p. 64), to be listed in that genus.
Ch. marchicum, described by Lindau (53) two years later, also
possesses characteristics of a Melanospora. According to the
original description and figures a mouth or ostiole is formed at
the top of the perithecium and this is surrounded by a few stiff
bristles. In this area the adjacent wall cells become somewhat
elongated. Saccardo (83, p. 627) has given to this form the
name Sphaeroderma marchicum.

In 1898 Ellis and Everhart (36) described as new Ch. Abieti-
num, and Hennings (46, pp. 153 and 154) described two new
species using the names Chaetomium ? importatum and Ch. мей-
icola respectively. i

Through the kindness of the New York Botanical Garden,
plants from the original specimens of Ch. Abietinum E. & E.
have been placed at the disposal of the writer, and a detailed study
of that species has been made. Тһе conclusion has been reached
that the plants in question cannot be included in our genus. Іп
mounts of the type material the perithecia are black, opaque and
somewhat coriaceous, firmly adnate to the substratum of wood
upon which they are growing, lacking an ostiole, and very densely
covered throughout with short, stout, spine-like hairs typical
more of Coleroa or Venturia than of Chaetomium. According to
the original description the species is allied to Ch. sphaerospermum
E.&E. Тһе bristles, however, are not branched and the sporidia
are rather larger and only slightly compressed. At first sight
the spores are similar to those of Ch. sphaerospermum E. & Е.
which in this paper has been listed аз a synonym under Ascotricha
chartarum Berk., but with more careful study one finds that the
spores are only slightly compressed and lacking in a girdle.

А study of the two authentic specimens kindly sent by Dr.
Lindau, to which Hennings gave the name Chaetomium ? im-
portatum and Ch. laeliicola respectively, is sufficient to convince
one that neither of these plants can be regarded as а Chae-

tomium. The first named species appears as dark, chocolate- | SE

MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA 517

brown or black, irregular pustules, with here and there stiff, dark
bristles near the margin of the pustules. It has little resemblance
to a Chaetomium. Mounts of authentic specimens of Ch.
laeliicola, to which Saccardo (85, р. 429) has later referred as
Ch. laeliicolum, show the plants to be in the form of irregular
pustules erupted through the epidermis of the host, from which
arise stout, dark, septate hairs. Sections of the same material
show the fruiting bodies to be sunk deeply below the surface of
the tissue of the substratum on which they grow.

Іп 1900 Cocconi (15) gave the name Ch. papillosum to а
plant which he found on rotting wood in a field at Bologna, and
which resembles a Magnusiella in certain of its characteristics,
and Ch. murorum Cda. in others, yet differs from both in some
respects. He described and figured this form as having a small,
globose perithecium, perforated at the apex by an ostiole, and
with hairs diverging in all directions. At the apex the peri-
thecium was rather scantily supplied with abruptly incurved
hairs. The asci were cylindrical, with their spores arranged in a
single row. According to Cocconi experiments with cultures of
this plant demonstrated on the one hand the formation of a
Pycnidial stage which produced two-celled spores, and on the
other, the formation of branched conidiophores bearing spores
in a clump at the tips of the branches.

In 1902 Massee and Salmon (59) published a diagnosis of Ch.
arachnoides. At first sight mounted specimens of this form,
which were received from the Royal Botanic Gardens at Kew,
may be easily mistaken for а Chaetomium, but with more careful
€xamination characteristics appear which are sufficient to exclude
it from this genus. At the top of the perithecium is a long neck
resembling that of a Melanospora, and it is from the neck that
the hairs arise which form at maturity a tangled mass, and which
Closely resemble the hairs of Ch. murorum Cda. The spores
at maturity are honey-yellow in color, a characteristic unusual
for Chaetomium spores.

There are three scattered nomina nuda which have been
found in literature unaccompanied by description or figures, and
regarding which little is known. Material under the name of
Ch. Bromelliae Schw. has been found in the Curtis herbarium,

218 MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

but no further mention or description has been published. The
plants here included are evidently not Chaetomia, for they consist
of spherical or elongated, black pustules, firmly attached to the
substratum by a broad base. There are no hairs with the excep-
tion of a few bases of stout hyphae at the sides of the fruiting body.
No spores could be obtained. The material in the Curtis her-
barium which, according to a note, came from the herbarium of
Schweinitz, and which undoubtedly represents authentic speci-
mens, appears to be an Epicoccum.

In the Nomenclator Fungorum of Streinz (100, p. 183) is found
the following: “Chaetomium Montagnei Brond. in litt. ad M."
This manuscript name, apparently used by Brondeau in a letter
to Montagne, seems never to have been published. In the refer-
ence quoted it is said to be identical with Ch. murorum Cda.

In the Curtis herbarium is to be found under No. 5978
the name Ch. subnudum B. & C. Examination of material here
contained, which must have been collected previous to 1872,
shows the fruiting bodies to be of the nature of pustules sunken
in the tissues of the host, black and round or elongated. Тһе
spores are dark bronze and one-septate. It is evident that the
material is not that of a Chaetomium and so far as can be learned
no description has ever appeared.

THE Genus AscorRiCHA, HISTORICAL REVIEW

A description of the genus Ascotricha was first published by
М. J. Berkeley in the Annals of Natural History (5, р. 257).
The characteristics were there cited as follows: “Peridium thin,
at length bursting, clothed with dark, sub-pellucid, even, obscurely
jointed hairs; sporidia simple, contained in linear asci. Super-
ficial, at length free or only supported by the investing thallus,

ack." In connection with his account of the genus, he gave à
very full and complete description of a single species, 45. chartarum,
and illustrated the same with six figures. Only one other form
similar to that published by Berkeley is known. This was first
described by Ellis in 1890, in the Proceedings of the Academy of
Natural Sciences of Philadelphia, page 220, under the name
Ch. pusillum.

In 1871 Cooke included this genus in the Handbook of British

хезм дай

MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA 219

Fungi (2: 653), and defined it as follows: perithecium thin, free,
astomate, seated on loose, branched, conidiophorous threads;
sporangia linear, containing dark, elliptical sporidia.

These fungi should not be considered as true Chaetomia, but
rather as species of an allied genus. The spherical perithecia,
almost naked below and constricted at the top into a neck from
which the terminal hairs arise, the curious stiff or whip-like am-
pullate hairs bearing large numbers of conidia, and the disc-
shaped spores of one of the species, afford a sufficient basis for
separating. these forms from the genus Chaetomium. The earlier
writers mentioned above were not aware that the perithecia of
Ascotricha were provided with an ostiole, for Berkeley stated that
the perithecia finally burst, and Cooke described them as astomous.
Otherwise their descriptions are reasonably clear, and it therefore
seems best to restore Berkeley’s original name Ascotricha to his
species, and to change the name of Ch. pusillum E. & E. to Asco-
tricha pusilla.

CHARACTERS OF THE GENUS ASCOTRICHA
Perithecium when young subconical, dark green and trans-
lucent, when mature spherical or nearly so below, constricted
above into a narrow and distinct neck, perforated by a central
ostiole. Differentiated rhizoids lacking. Perithecial wall brittle,
membranaceous. Lateral hairs colored and branched, varying
in number, sometimes numerous, but at times almost wanting.
Terminal hairs numerous, arising from the region of the neck,
branched sympodially, and distinguished by the prese of
clavate ampullae. Asci linear, cylindrical, 8-spored. Spores
arranged in a single row in the ascus, simple, dark colored, dis-

charged from the ostiole in a black cylindrical cirrhus.

КЕҮ TO THE SPECIES OF ASCOTRICHA

Perithecial hairs and conidiophores slender, frequently
gtacefully flexed, sparingly branched. Conidia 5.5 Х
3.5 и, smooth or nearly зо. Ascospores ovate to egg-

ped, 5.5 X 44 nA T г. As. pusilla (p. 220).

Perithecial hairs and conidiophores stout, profusely

ties ia 6 Хун, roughened minute

hed. Coni
wart-lik jecti isc-shaped, 8 X
e projections. Ascospores disc-s1Ap 2. As. chartarum (р. 222).

7-5 и.

220 MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

Ascotricha pusilla (Ellis & Everhart) comb. nov.
Chaetomium pusillum Ellis & Everhart, Proc. Acad. Nat. Sci..

Phil. 1890: 220. 1800.

Chaetomium Ellisianum Saccardo & Sydow, Syll. Fung. 14: 491.

1899.

PLATE 17, FIGS. 13-21

Black. Perithecia small, globose, the body somewhat broader
than long, constricted above to form a distinct neck, 151 X 1664
(135-190 X 139-202), extruding long, black cirrhi, rather firmly
adnate to the substratum and associated with numerous conidio-
phores. Lateral hairs when present sparsely scattered over the
upper portion of the perithecium wall, slender, whip-like, more or
less flexed or gracefully recurved to form a loop, tapering almost `
imperceptibly, evenly and remotely but obscurely septate, slightly
swollen at the septa, near the base dark olive-brown to blac
and about 5.6 in thickness, becoming pale olive at the tips,
unbranched except very rarely when the short main axis pro-
duces an extremely long, whip-like outgrowth just above the
origin of which the axile portion terminates in a club-shaped,
hyaline ampulla. Terminal hairs arising in the region of the
neck, extremely long, slender, whip-like, graceful, smooth and
even, with the exception of slight angular swellings at the septa,
flexed and often recurved forming a loop, septate but so dark
below that the septa are nearly obscured, very dark olive to
black and about 5.6 шіп thickness at the base, almost imperceptibly
tapering and fading toward the pale olive tip, unbranched except
in rare cases, where branching takes place in the same manner
as described above for the lateral hairs. Asci extremely deli-
cate, linear, cylindrical, 8-spored, 60 X би, pars sporif. 43:
Spores monostichous, hyaline when young, when mature dark
olive-brown, ovate, roughly egg-shaped, rounded at both ends,
56 X 4.2 џ (5.4-6 X 3.5-4.5), when seen edgewise, compressed,
3.2 и broad.

Оп barrels in cellar, Cambridge, Massachusetts, and Kittery
Point, Maine, herb. К. Thaxter; on barrel hoops and packing
boxes in cellar in Hanover, New Hampshire, and Amesbury,
Massachusetts (Chivers No. 41). Reported also by G. F. Atkin-
son from Alabama, growing on old paper.

Type locality: as Ch. pusillum Е. & E., Newfield, New Jersey;
on basswood barrel bottom in cellar. Also Manhattan, Kansas;
on an old churn in cellar (Kellerman 1437).

The original material of this species was collected by Ellis

MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA 221

at Newfield, New Jersey, and described by him as Chaetomium
pusillum, in apparent ignorance of the fact that this combination
had already been employed by Fries (38) for a different plant.
For this reason a new name Ch. Ellisianum was proposed by
Saccardo in the fourteenth volume of the Sylloge, p. 491, but
since for reasons already stated it has seemed best to recognize
the genus A scotricha, it becomes necessary to return to the original
specific name used by Ellis, and to employ a new combination in
designating this species.

Conidial growth precedes the development of the perithecia.
The conidiophores when young are grayish-green to the naked eye,
while at maturity they are black. Under magnification they
appear long and slender, rather sparingly branched, at the base
dark olive-brown to black and about 3.5 и in thickness, tapering
and fading toward the tip, one to several times sympodially
branched, the individual axes producing a lateral branch, above
the point of origin of which the axis terminates in a clavate,
hyaline ampulla. Тһе lower branches in turn become verticil-
lately branched, and bear large clusters of ovate to globose
conidia which are hyaline when young, olive-brown at maturity,
and measure 5.4 X 3.6 и, appearing smooth except with ап
immersion lens when a slight irregularity of the wall is apparent.
In the mounts most favorable for study the threads of the my-
celium are woven together into rope-like strands, along which
conodiophores branch forth in large numbers and perithecia take
their origin.

The species appears to be by no means rare in this country
and may frequently be found often in company with other inter-
esting forms growing on the bottom of barrels which have stood
for some time in a damp situation, as for example on a cellar
bottom. It may quickly be recognized by its jet black color and
the characteristic mingling of perithecia and conidiophores with
black masses of conidia and ascospores. This species may easily
be distinguished from Ascotricha chartarum, which it most nearly
resembles, by the following differences: the smaller perithecia,
the extremely long, whip-like hairs which are frequently grace-
fully looped, but seldom branched, and the much smaller spores,
Which are egg-shaped instead of disc-shaped, and without a girdle.

222 MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

ASCOTRICHA CHARTARUM Berkeley, Ann. Nat. Hist. І: 257. pl. 7,
1838

Ascotricha Zopp: (Вош.) Peyronel, Ann. Myc. 12: 459-470. f. 1-3.
1014.

Chaetomium Berkeleyi Schroeter; Cohn, Krypt.-Fl. Schlesien 3:
284. 1894.

Chaetomium chartarum Winter, Rabenhorst’s Kryptogamen-Flora
1: 157. f. 4-7. 1885. Not Chaetomium chartarum Ehrenberg,
Sylv. Berol. 27. 1818.

Chaetomium delicatulum Roumeguére, Rev. Myc. 7: 22. 1885.

Chaetomium sphaerospermum Cooke & Ellis, Grevillea 8: 16.

1879.
Chaetomium Zopfii Boulanger, Rev. Сеп. Bot. 9: 25. pl. 1-3. 1897.
Dicyma ampullifera Boulanger, 1. с. 9: 17. pl. 1-3. 1897.
Sporotrichum sp. Boulanger, 1. c. 9: 17. pl. 2. 1897.

PLATE 17, FIGS. 1-12.

Black. Perithecia rather small, globose or subglobose, con-
stricted into a neck at the upper extremity, frequently flatten
at Ше base, 1924 high X 198и broad (126-232 X 135-236),
discharging black spore cirrhi many times longer than the peri-
thecium, producing no differentiated rhizoids, but seated among
and upon the stalks and branches of the conidiophores. Lateral
hairs rather numerous, very variable in length but nearly always
short, tapering near the tips only, branched and ampullate as are
the terminal hairs, very dark olive-brown to black below, about
3.2 и in thickness near base, colorless and frequently septate near
their tips. Terminal hairs extremely variable in length, straight,
stiff, dark olive-brown to black except at the ampullae and extreme
tips, remotely septate, frequently with an angular swelling at the
septum, profusely and sympodially branched, each individual
axis producing one or two lateral branches beyond the point of
origin of which the axis terminates in a somewhat swollen, club-
shaped colorless ampulla. Asci linear, cylindrical, 8-spored,
65 ХІ Та, pars sporif. 45 и. Spores monostichous, when young
pale olive with a greenish tinge, when mature dark olive-brown
to black, regularly or irregularly circular in face view, lenticular, `
with girdle apparent when seen in profile, 8.1 X 7.7 в (7.279 x
72-81), when seen edgewise 4.5-5.4 и broad.

_Exsiccati.—Fung. Europ. Edit. nova, Series И, ХХУ, 2472;
Micro-Fung. Brit. IV, 348, 355 and V, 474; Myc. Ital. I, 685

MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA 223

Myc. March. 69. Sub Ch. delicatulum Roum.: Fung. Gall.
XXXII, 3143.

On cardboard, Cambridge, Massachusetts, Herb. R. Thaxter.
On barrel bottoms from various localities in New England (Chivers
Хо. до). Reported also from different parts of the United States,
South America, and Germany. Type locality: King's Cliffe,
England; on a candle box.

The species was first observed in America by Ellis and was
communicated to Cooke who described it as Ch. sphaerospermum,
as indicated in the above synonymy. Тһе writer has been able
to examine specimens of the original material collected by Ellis,
and has found it to be identical in every respect with Ascotricha
chartarum.

Although this species appears to be so clearly characterized it
has been repeatedly re-described, first in 1879 by Cooke and
Ellis as above mentioned.

In 1885 Winter in his Pilze Deutschlands (110, p. 157) includes
this species under the name Ch. chartarum, in apparent ignorance
of the fact that this combination had already been used by Ehren-
berg (28, p. 27) for a wholly different plant, and in the same year
Roumeguére added still another name to the list by re-describing
this plant as Ch. delicatulum, since an examination of the descrip
tion given by him, as well as a study of the authentic specimens
distributed in Fung. Gall. No. 3143, leave no doubt as to its
synonymy.

Schroeter (89) in 1894, being aware of the previous use of the
name Ch. chartarum by Ehrenberg, but following Winter M his
generic reference of the present species, completed. the list of
synonymy by adding yet another name Ch. Berkeley.

In 1897 the form was redescribed under the new name Ch.
Zopfii by Boulanger (10), to whom we owe a Very complete account
of its morphology and development. T his author made excellent
figures which place the identity of the plant which he had before
him beyond question.

During the publication of this paper the writer has noted an
article by Peyronel (68), in which he transfers this species from
the genus Chaetomium to that of Ascotricha, still retaining Bou-
langer's specific name. Ав there seems to be no doubt that Ch.

224 MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

Zopp? Boulanger is identical with Ascotricha chartarum, described
by Berkeley in 1838, it seems to the present writer more desirable
to return to the use of the original name given by Berkeley than
to continue the names given by more recent authors.

As in Ascotricha pusilla conidial growth precedes the formation
of perithecia. The conidiophores to the naked eye are greenish-
gray when young and black at maturity. Under magnification
they are dark olive-brown to black near their base and about Su
in thickness, clearly but irregularly septate and very profusely
and sympodially branched, each individual axis producing a single
branch, or forking and producing two symmetrical branches
beyond the point of origin of which the axis terminates in a color-
less, clavate ampulla. This may be several times repeated and the
peripheral branches thus produced may form an irregular whorl,
from which arise clusters of conidia which are roughly spherical,
ovate or egg-shaped, roughened by minute warts, hyaline when
young and olive-yellow or olive-brown when mature, 6.1 X 5.14
(5.4-7-2 X 3.6-5.9). |

This species is most often found on paper and cardboard, but
like As. pusilla, the only form with which it is in danger of being
confused, it occurs frequently on barrels or boxes in cellars. A
comparison shows at once that in the case of the former the регі-
thecium is of greater average size, the hairs are much stiffer,
straighter, and more profusely branched, the spores are larger
and easily distinguished by the fact that they are circular in face
view, but lenticular with a girdle at the margin, when seen in
profile.

MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA 225

LITERATURE CITED
In the following bibliography only the more important references
have been included and no attempt has been made to list the nu-
merous references which indicate simply the distribution of the various
species.
Auerswald, B.
I. Synopsis Pyrenomycetum europaearum; Gonnerman & Raben-
horst, Mycologia Europaea. 5,6: [1]-[30]. 1869.
Albertini, J. B., & Schweinitz, L. D. de.
2. Conspectus fungorum in Lusatiae superioris agro Niskiensi
crescentium. Leipzig. 1805.
Bainier, G.
3. Mycothéque de l'Ecole de Pharmacie XXX. Monographie des
Chaetomidium et des Chaetomium. Bull. Soc. Myc. France
25: 191—237. pl. 10-26. 1910.
Beauverie, J.
4. Études sur le Polymorphisme des Champignons. Influence du
milieu. Ann. Univ. Lyon, Nouv. Sér. I. 3: 1-266. f. 1-75.
1900.
Berkeley, M. J |
5. Notices of British Fungi. Ann. Nat. Hist. 1: 257-264. pl. 7, 8.

1838.
Berkeley, M. J., & Broome, C. E.
6. Notices of British Fungi. Ann. and Mag. Nat. Hist. П. 7: 97-
102; 176-180. pl. 5-7. 1851.
7. Notices of British Fungi. Ann. and Mag. Nat. Hist. IV. п:
339-349. pl. 7-10. 1873.
Bommer, Е., & Rousseau, М.

8. Florule mycologique des environs de Bruxelles. Bull. Soc.
Bot. Belgique 23: 15-365. 1884.
Bonorden, H. F.
1851.

9. Handbuch der allgemeinen Mykologie. Stuttgart.
Boulanger, E.
10. Sur une forme conidienne nouvelle dans le genr
Rev. Сеп. Bot. 9: 17-26. pl. 1-3. 1897-
Brondeau, L. de.
11. Note sur le Chaetomium chartaru
4: 999-1001. 1857.
Cavara, F.
12. Fungi Longobardiae exsiccati. Padua. 1892.

e Chaetomium.

m Ehrb. Bull. Soc. Bot. France

226 MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

Cesati, V., & Notaris, G. de.

13. Schema di classificazione degli Sferiacei italici aschigeri рій o
meno appertenenti al genere Sphaeria nell’ antico significato
attribuitogli da Persoon. Comment. Soc. Crittogam. Ital. 1:
177-240. 1863.

Chivers, А. H.

14. Preliminary diagnoses of new species of Chaetomium. Proc.

Am. Acad. 48: 83-88. 1912.
Cocconi, G.

15. Intorno ad una nuova specie di Chaetomium. Mem. R. Accad.

Sci. Ist. Bologna 5: 683-688. pl. 8, f. 1-0. 1900.
Cooke, M. C.

16. British Fungi. Grevillea т: 174-180. 1873.

17. Cocoa-Palm Fungi. СгеуШеа 5: 101-103. 1877

18. New British Fungi. Grevillea 6: 71-76. pl. 97, P 16. 1877.

19. New British Fungi. Grevillea 8: 1-11. 1879.

20. Australian Fungi. Grevillea 12: 21. 1883.

Corda, A.

21. Icones Fungorum 1. Prague. 1837.

22. Icones Fungorum 2. Prague. 1838.

23. Icones Fungorum 4. Prague. 1840.

Dangeard, P. A.
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: 330-333. pl. 73. 1907.
басы" М. с.
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Desmaziéres, J. B. H. J

26. Plantes cryptogames de France. Lille. 1853.
Duby, J. A.

27. Botanicon Gallicum 2. Paris. 1830.
Ehrenberg, C. G.

28. Sylvae mycologicae berolinenses. Berlin. 1818.
Eidam, E.

29. Zur Kenntaiss der Entwicklung bei den Ascomyceten. Cohn's
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Ellis, J. В., & Everhart, В. М.

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31. New Species of Fungi from various Localities. Jour. Мус. 4:
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32, Se American Pyrenomycetes. Newfield, New Jersey. `
1 А e и

MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA 221
33. New Species of North American Fungi from various localities.
Proc. Acad. Nat. Sci. Philadelphia 1893: 128-172. 1893.
34. New d of Fungi from various localities. Proc. Acad. Nat.
Sci. Philadelphia 1894; 322-386. 1894.
35. New species of Fungi from various localities. Am. Nat. 31:
340. 1897.
36. New species of Fungi from various localities. Bull. Torrey
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Fries, E.
37. Systema Mycologicum 2. Lund. 1823.
38. Systema Mycologicum 3. Greifswald. 1829.
39. Summa _ vegetabilium Scandinaviae. Stockholm & Leipzig.
1849.
Fuckel, I.
40. Enumeratio Fungorum Nassoviae. Series 1. Wiesbaden. 1861.
41. Fungi rhenani. Hedwigia 5: 23—30. 1866.
42. Symbolae mycologicae. Wiesbaden. 1869.
Greville, R. K
43. Scottish Cryptogamic Flora 4. Edinburgh.
44. Flora Edinensis. Edinburgh. 1824.
Hansen, E. C.
45. De danske Gjédningssvampe (Fungi
skabelige Meddelelser 1876-78: 207-354. pl. 4-9
Hennings, P
46. Die іп den Gewáüchsháusern
beobachteten Pilze. Abh. Bot
109-176. pl. 1, 2. 1898.
Karsten, P. A.
47. Enumeratio Fungorum
aestate 1861 lectae.

1826.

fimicoli danici). Viden-
1876.

des Berliner botanischen Gartens
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et Myxomycetum in Lapponia Orientali
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1882.
48. Symbolae ad Mycologiam Fennicam. XXVII, Meddel. Soc.

. Fl. Fenn. 16: 33-36. 1888.
Koorders, 5. Н.
49. Botanische Untersuchungen über einige іп Java vorkommende
Pilze, besonders über Blatter bewohnende, parasitisch аш-
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1907.
Kunze, G., & Schmidt, J. K.
50. Mykologische Hefte т. Leipzig 1817.

51. Deutschlands Schwimme. Leipzig. 1818.

228 MonoGRAPH ОҒ CHAETOMIUM AND ASCOTRICHA

Léveillé, M. J. H.

52. Champignons exotiques. Ann. Sci. Nat. Bot. III. 3: 38-70.
1845.

Lindau, G.

53. Zwei neue deutsche Pilze. Hedwigia 35: 56-57. /. А-Е. 1896.
Link, H. F.

54. Linnaeus, C., Species Plantarum, Ed. 5, т. Berlin. 1824.
Marchal, E.

55. Champignons coprophiles П. Décade d'espèces nouvelles pour
la Belgique. Bull. Soc. Roy. Bot. Belgique 237: 59-61. 1884.

56. Воттетейа, nouveau genre des Pyrénomycétes. Bull. Soc.
Roy. Bot. Belgique 247: 164-165. 1885.

57. Bommerella, nouveau BEE des Pyrénomycétes. Rev. Myc. 8:
IOI. 1886

58. Champignons coprophiles de Belgique. V. Note sur le Bom-
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271. pl. 10. 1880.

Massee, G., & Salmon, E. S.
59. Researches on Coprophilous Fungi. ІІ. Ann. Bot. 16: 57—93. bl.
4, 4. t002.
Мане о.
бо. Fungi; Savoia, Il Ruwenzori t. Milan. 1909.
Montagne, C.

61. Septiéme centurie de plantes cellulaires nouvelles tant indig-

enes qu’ exotiques. -Ann. Sci. Nat. Bot. IV. 5: 333-374. 1856.
Morgan, A. P.

62. Pyrenomycetes scarcely known in North America. Jour. Myc.

10: 226-228. 1904.
Oltmanns, F.

63. Über die Entwickelung der Perithecien in der Gattung Chaeto-
mium. Bot. Zeit. 45: 193-200, 200-218, 225-233, 249-254»
265-271. 1887.

Oudemans, C. A. J. A.

64. Aanwunsten voor de Flora Mycologica van Nederland IX en X.
Nederlandisch Krinkundig Archief. II. 4: 267. pl. 6, f. 14
1885.

Palliser, H. L.

65. Chaetomiaceae. North American Flora 3: 59-64. 1910.

66. Report of the botanist. Rep. New York State Mus. Nat.
Hist. 27: 15 uS: P а, 0875.

MonoGRAPH ОҒ CHAETOMIUM AND ASCOTRICHA 229

67. Report of the botanist. Rep. New York State Mus. Nat.

Hist. 49: 19-83. pl. 44-49. 1896.
Peyronel, B.

68. Osservazioni critiche e sperimentali su alcune specie del genere
Dicyma Boul. e sui loro stati ascofori. Ann. Myc. 12: 459-
470. f. 1-3. 1914.

Preuss, G. T.

69. Uebersicht untersuchter Pilze, besonders aus der Umgegend von

Hoyerswerda. Linnaea 24:99-153. 1851; 25: 79-80. 1852.
Quélet, L.
70. Les Champignons du Jura et des Vosges. III. Мет. Soc
d'Emulation de Montbeliard 1875: 1-128. pl. 1-4. 1876.
Rabenhorst, L.
71. Klotzschii herbarium vivum mycologicum. 1851.
Reinke, J., & Berthold, G.

72. Die Zersetzung der Kartoffel durch Pilze. Untersuch. Bot. Lab.

Göttingen т: 46-51. pl. 4. 1879.
Rehm, H.

73. Beiträge zur Ascomyceten-Flora der deutschen Alpen und

Voralpen. Hedwigia 21: 97-103, 113-123. 1882.
Saccardo, F. A.

74. Fungi Italici autographice deliniati. Padua. 1877-1886.

75. Fungi Veneti novi vel critici. Series V. Nuovo Gior. Bot. Ital.
8: 161-211. 1876. SE

76. Fungi novi ех herbario Prof. Doct. Р. Magnus Berolinensis.
Michelia 1: 117-132. 1878.

77. Fungi Veneti novi vel critici vel mycologi
(adjectis nonnullis extra-venetis). Series XIII.
528-563. 1882.

78. Fungi belgici lecti а cl. Dominis Elisa Bommer ес
Rousseau. Misc. mycol. 2: 435-463- 1884.

79. Sylloge Fungorum т. Padua. 1882.

80. Sylloge Fungorum 9. Padua. 1891.

81. Sylloge Fungorum 11. Padua. 1895.

82. Sylloge Fungorum 12. Padua. 1897.

83. Sylloge Fungorum 14. Padua. 1899.

84. Sylloge Fungorum 15. Padua. 1901.

85. Sylloge Fungorum 16. Padua. 1902.

86. Sylloge Fungorum 17. Padua. 1904

87. Sylloge Fungorum 10. Padua. 1919.

88. Sylloge Fungorum 22. Padua. 1913-

ae Venetae addendi
Michelia 2:

Maria

230 MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

Schroeter, J.

89. Pilze; F. Cohn, Kryptogamen-Flora von Schlesien 3. Breslau.

1894.
Schmitz, F.

90. Untersuchungen über die Zellkerne des Thallophyten. Sitzungs-
ber. niederrhein. Ges. für Naturhist. und Heilkunde in Bonn
345-376. 1879.

Schweinitz, L. D. von.

91. Synopsis fungorum in America boreali media degentium.

Trans. Amer. Phil. Soc. 4: 141-316. pl. 19. 1832.
Smith, A. L., & Carleton, R.

92. The fungi of germinating farm seeds. Trans. Brit. Myc. Soc.
1: 182. pl. 19. 1902.

93. Fungi new to Britain. Trans. Brit. Myc. Soc. 2: 31-40.
1903.

Spegazzini, C.

94. Fungi coprophili Veneti. Michelia 1: 222. 1878.

95. Fungi Argentini additis nonullis Brasiliensibus Montevidien-
sibusque Pug. 4: 98. Хо. 112. 1881.

96. Fungi Argenti novi v. critici. Anales del Mus. Nac. Buenos
Aires 6: 81-365. pl. 4, 5. 1899.

Starbück, K.

97. Anteckningar ófver nàgra skandinaviska Pyrenomyceter. Ві-
hang till Kongl. Svenska Vet.-Akad. Handl. 143, No. 5: 1-18.
pl. т. 1889.

Stewart, Е. С.

98. Notes on various plant diseases. VI. Chaetomium contortum on

barley seedlings. Bull. N. Y. Ag. Exp. Station Bull. 164:

99. Pilze; J. Sturm, Deutschlands Flora 3. Nuremberg. 1853
Streinz, W. M.

100. Nomenclator Fungorum. Vienna. 1862.
Taubenhaus, J. Т

о1. Chaetomium root rot. Delaware College Ag. Exp. Station,
Bull. 106: 24-27. 1914. [Illustrated.]

erry, J. J.

102. Contributions mycologiques. Ann. Soc. Bot. Lyon 10: 209.
1883.

3
Tode, H. J.
_ 103. Fungi Mecklenburgenses selecti 2. Lüneburg. 1791.
s PET nus ms я i

MonoGRAPH OF CHAETOMIUM AND ASCOTRICHA 231

Van Tieghem, Ph.
104. Sur la développement du fruit des Chaetomium et la prétendue
sexualité des Ascomycétes. Compt. Rend. 8r: 1110-1113,

105. Sur le développement du fruit des Ascodesmis, genre nouveau de
l'ordre des Ascomycétes. Bull. Soc. Bot. France 23: 271-279.
1876.
106. Nouvelles observations sur le développement du périthéce des
Chaetomium. Bull. Soc. Bot. France 23: 364-366. 1876.
107. Sur le développement de quelques Аѕсотусёќеѕ, Bull. Soc. Bot.
France 24: 96-105. 1877.
108. Remarque au sujet du développement des Chaetomium. Bull.
Soc. Bot. France 20: 317-318. 1882.
Wallroth, Е. б.
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Winter, G.
110. Die Pilze Deutschlands, Oesterreichs und der Schweiz. П.
. Abtheilung. Rabenhorst’s Kryptogamen-Flora 1. 1887.
111. Exotische Pilze. IV. Hedwigia 26: 6-18. 1887.
Zopf, W.
112. Untersuchungen über Chaetomium. Abh. Bot. Ver. Prov. Bran-
denburg 19; 170-173. 77
113. Zur Entwickelungsgeschichte der Ascomyceten.
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Zukal, Н.
114. Mykologische Untersuchungen. Anlage des Perithecium von
Chaetomium | crispatum Fuckel. Denkschr. Kaiser. Akad.
Mathem.-Naturwiss. Klasse 51: 21-36. pl. 1-3. 1886.

Chaetomium.
1881.

`

232 MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

EXSICCATI
The writer's conclusions have for the most part been based on the
exsiccati specimens in the Cryptogamic Herbarium at Harvard Univer-
sity.
Berkeley, M. J. British Fungi.
Cavara, F. Fungi Longobardiae Exsiccati.
Cooke, M. C. Fungi Britannici.
Curtis, M. A. Collection.
Desmaziéres, J. B. H. J. Plantes Cryptogames de France.
Ellis, J. B., & Everhart, B. M. North American Fungi. Fungi
Columbiani.
Fries, Е. Scleromyceti Sueciae.
Fuckel, L. Fungi Rhenani.
Jaap, O. Fungi Selecti Exsiccati.
de Jaczewski, A. Fungi Rossiae Exsiccati.
Karsten, P. A. Fungi Fenniae Exsiccati.
Krieger, K. W. Fungi Saxonici.
Kunze, G., & Schmidt, J. К. Deutschlands Schwamme.
Mougeot, J. B. & J. A., & Nestler, C. Stirpes Cryptogamae Vogeso-

Rabenhorst, L. Fungi Europaei. Klotzschii Herbarium vivum

e Н. Ascomyceten.
Rick, 7. Fungi Austro-Americani.
Roumeguére, С. Selecti Gallici Exsiccati.
Saccardo, D. Mycotheca Italica.
Saccardo, P. А. Mycotheca Veneta.
de Thumen, Е. Fungi Austriaci Exsiccati. Mycotheca Universalis.
Vestergren, Т. Micromycetes Rariores Selecti.
ill, A. Fungi Bavarici Exsiccati.
Vize, І.Е. Micro-Fungi Britannici.
Wartmann, B., & Schenk, B. Schweizerische Kryptogamen.
Westendorp, С. D., & Wallays, A. C. F. Herbier Cryptogamique Belge.
Zahlbruckner, A. Kryptogamiae Exsiccatae, editae a Museo Palatino
"Vindobonensi (Fungi by Keissler, C.).
Zopf, W., & Sydow, P. Mycotheca Marchica.

MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA 233

EXPLANATION OF PLATES 6-17
Unless otherwise indicated all figures are drawn with the following magnifica-
tions: perithecia X 47, lateral and terminal hairs X 180, asci X 390, spores X 390.

PLATE 6
CHAETOMIUM ELATUM Kunze & Schmidt
Fic. т. Mature саки
Fic. 2. Lateral hai
Fic. 3. Terminal ae
Fic. 4. Ascus.
Fic. 5. Spores. (a) Young spores with oil globules; (5) mature spores seen in
face view; (c) mature spore seen edgewise

CHAETOMIUM TRIGONOSPORUM (Marchal) Chivers
Fic. 6. Mature perithecium showing the relative size of the spore mass, Ж 25:
Fic. 7.. аво репера
Fic. 8. Latera
FIG. о. PEERY vay
Fic. 10. Asci а: variation in shape.
Fic. іі. Spores. (a) Young spores with oil RES (b) mature spore seen in
face view; (c) mature spores seen edgewise

PLATE 7
CHAETOMIUM INDICUM Corda
Fic. і. Mature perithecium
Fic. 2. Terminal hair мы the characteristic dichotomy and reflexed branches.
Fic. 3. Distal жа of a terminal hair showing the encrustation of calcium
oxalat

FiG. 4. Terminal ге of a different character showing dichotomy and narrow
acute angles.

Fic. 5. Distal portion of the same hair at a much older stage when its surface
has become encrusted with calcium oxalate.

Fic. 6. Lateral hair.

Fic. 7. Ascus. J

Fic. 8. Spores. (а) Young spores with ой € (b) mature spores seen in
face view; (c) mature spore seen edgewi

CHAETOMIUM FUNICOLUM Cooke
Fic. 9. Mature perithecium.
FIG. ro. Mature perithecium with slightly greater magnification,
ation in which the terminal hairs do not form a dense m
i ІІ. Lateral hair.
165. I2, 13. Terminal hairs showing dichoto :
Fics. 14, 15. Distal portion of un inal d irs showing the encrustation “
calcium oxalate

eas a vari-

Fics. 16, 17. Asci
Fics, 18, са _ Spores. (а) Young spores Ms oil globules;
een ед

(b) mature spores

that the Ken is not constant.

234 MoNOGRAPH OF CHAETOMIUM AND ASCOTRICHA

PLATE 8
CHAETOMIUM CIRCINATUM Chivers
Fic. 1. Mature perithecium.
Fic. 2. Terminal hair.
Fic. 3. Terminal hair showing an extreme case of circinate coiling.

. Ascus.

Fic. 5. Spores. (a) Young spore with oil globules; (b) mature spore; (c) mature
spore showing the characteristic furrow caused by the collapse of the
spore wall.

CHAETOMIUM MURORUM Corda

Fic. 6. Mature perithecium.

Fic. 7. Terminal hair.

Fics. 8,9. Asci showing their varying shape.

Fic. то. Spores. (а) ‘Young spores with oil globules; (b) mature spores; (c) mature
spore showing the characteristic furrow caused by the collapse of the
spore wall.

PLATE 9
CHAETOMIUM SIMILE Massee & Salmon

Fic. 1. Mature perithecium.

Fic. 2. Terminal hair. ;

Fic. 3. Азсиз, copied from the original drawing by Маззее and Salmon, Х 400
and reduced by one half.

Fic. 4. Spores. (a) Young spores with oil globules; (b) mature spores seen in
face view; (с) mature spore seen edgewise.

CHAETOMIUM CRISPATUM Fuckel

Fic. 5. Mature perithecium.
Fics. 6, 7. Terminal hairs showing the variation in the manner of coiling.
Ascus

8 қ
Fic. 9. Spores. (а) Young spores with ой globules; (b) mature spores seen in
face view; (c) mature spore seen edgewise.
CHAETOMIUM CONTORTUM Peck
- Fic. ro. Mature perithecium.
Fic. іі. Terminal hair. |
IG. 12. Spores. (a) Young spores with ой globules; (6) and (с) mature spores.
CHAETOMIUM TORTILE Bainier
Fic. 13. Mature perithecium.
Fic. 14. Terminal hair.
Fic. 15. Terminal hair, distal portion showing the variation in the manner of
coiling.

Fic. 16. Ascus.
Fic. 17. Spores. (a) Mature spores seen in face view; (b) seen edgewise.
PLATE IO

CHAETOMIUM SPINOSUM Chivers
Fic. 1. Mature perithecium.
М; M жараны шы аба D "

wine th tasinia Mir cirrhus of spores
(5555 Р “ы 4, ъв 5 ада E

MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA 235

Fic. 3. Terminal hair with branches.

Fic. 4. Distal portion of a branch of the -— hair.

Fics. 5, 6. Asci in different stages of developm

Fic. 7. Spores. (a) Young spores with oil mg (5) mature вроге.

CHAETOMIUM AMPULLARE Chivers

Fic. 8. Mature perithecium showing the relative size of the spore mass, X 25.
. Mature perithecium.
Fic. 10. Terminal hair.

м
e
ю

: cus.
FIG. 12. tee? (а) Young spores with oil globules; (5) mature spores seen In
face view; (c) seen edgewise.
CHAETOMIUM GLOBOSUM Kunze

Fic. іҙ. А germinating spore showing the cast-off exospore and the аламан?

endospore 390.

Fic. 14. A slightly date Ae showing the origin of the mycelial branches from
the endospo

Fig. 15. A germinating | spore in which the endospore does not appear аз а DÉI:
sistent vesi

cle.
Fic. 16. A diagrammatic drawing of the mycelium, showing its radial oven:

PLATE ІІ
CHAETOMIUM TRILATERALE Chivers

Fic. І. Mature perithecium.
Fic. 2. Mature perithecium under greater magnification, X 94.

FiG. 3. Lateral hair.
Fic. 4. Terminal hair. px
FIG. 5. Ascus. A ш
Fic. 6. Spores. (а) Young spores with oil eme (5) mature spores eg
face view; (с) mature spore seen edgewise
CHAETOMIUM FUSIFORME Chivers
Fic.

7. Mature perithecium
Fic. 8. Mature perithecium оди greater magnification, X 94.
9. Terminal hai

. Ascus. wise,
FIG. 11. Spores. (а) Mature spores seen in face view; (b) seen edgewise.

CHAETOMIUM AUREUM Chivers
Fic. 12. Mature perithecium
Fic. 13. Mature perithecium здік greater шарий " eg
Fic. 14. Mature perithecium showing the relati
of the spore cirrhus, Х 25.
FIG. 15. Terminal hair.

i rvature

(8) mature spores seen in

FIG. 17. Spores. (a) Young spores with oil globules;
face view; (c) mature spore seen edgewise..

236 MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

CHAETOMIUM SPHAERALE Chivers
Fic. 18. Mature peritheciu
Fic. 19. Mature Een showing the relative size of the spore mass, о.
Fic. 20. Lateralhairs. (а) Short and sinuous; (b) somewhat longer, nearly straight
and unbranched; (c) still longer and branched.
Fic. 21. Terminal hair.

Fic. 22. Ascus.
FIG. 23. ie (a) Young spores with oil globules; (b) mature spores seen in
face view; (c) seen edgewise
PLATE І2

CHAETOMIUM ATERRIMUM Ellis & Everhart
Fic. 1. Mature perithecium.
Fic. 2. Lateral hair.
Fic. 3. Terminal hair.
Fic. 4. Spores. (a) Mature spores seen in face view; (b) mature spore seen edge-
wise.
CHAETOMIUM SPIRALE Zopf

Fic. б. Mature perithecium.

6. Terminal hai
Fic. 7. Lateral hair.
Fic. 8. Spores. (a) Mature spores seen in face view; (b) mature spore seen edge-

wise
PLATE 13
CHAETOMIUM BOSTRYCHODES Zopf

Fics. І, 2, 3. Mature perithecia showing the characteristic variation.
Fic. 4. Terminal hair, branched and irregularly convolute.
Fic. 5. Terminal hair, regularly spirally coiled.
Fic. 6. Terminal hair, branched and regularly spirally coiled with ten coils.
Fic. 7. Portion of a terminal hair showing the encrustation of calcium oxalate.
Fics. 8,9, 10. Asci, showing different stages of development.
Fic. 11. Spores. (а) Mature spores seen in face view paris a characteristic

: r tive area at each end; (b) mature spores seen edgewise.

CHAETOMIUM SUBSPIRALE Chivers
Fic. 12. Mature perithecium
Fic. 13. Mature Байдалы showing the relative size of the spore mass, Х 15.
Lateral hair.

cus.
Fic. 17. Spores. (а) Young spores with oil globules; (5) mature spores seen in
face view; (c) seen edgewise.

CHAETOMIUM CAPRINUM Bainier
Fic. 18. Mature perithecium
Fics. 19, 20. Terminal hairs "i branches.

Fic. 22. iid (a) Young spore with oil globules; (5) mature spores seen іп
е іт egenen :

SI ANP YH

мн m
NH O

Se be ym

MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA 237

PLATE 14
CHAETOMIUM QUADRANGULATUM Chivers
Mature а with the greater portion of the spore mass removed.
Mature peritheciu
Mature Н ірді; showing the relative size of the spore mass, Х 15.
A spirally coiled terminal hair.
An irregularly convolute terminal hair with branches.
Ascus seen in face view.
The same ascus seen edgewise.
Spores. (a) Mature spores seen in face view; (b) seen edgewise; с.)
grouped in a characteristic manner.

CHAETOMIUM CONVOLUTUM Chivers
Mature perithecium.
Terminal hair.
Ascus с
Spores. (a) Young spore with ой globules; (b) mature spore seen in
face view; (c) mature spores seen edgewise.

PLATE 15
CHAETOMIUM COCHLIODES Palliser
Mature peritheciu
Mature Steeg showing the relative size of the spore mass, Х 15.
Lateral hair.
A slender, spirally coiled terminal hair.
A slender, sinudus terminal hair with branches.
A stout, spirally coiled terminal hair.
Ascus. |
Spores. (а) Young spores with ой globules; (b) mature spore seen in
face view; (c) seen edgewise.

CHAETOMIUM GLOBOSUM Kunze
Mature perithecium
Mature "do тэ beatos the relative size of the spore mass.
Terminal h

Ascus.
Young spores with oil globules.
Mature spores.
PLATE 16
CHAETOMIUM TORULOSUM Bainier
Mature perithecium.
Lateral hairs
Terminal "Ups with branches.
Ascus. ;
Е " mature
Spores. (a) Large spore when young with oil globules; (b) p bules; :
spores seen in face view; (с) ашай врогез when young w™ ed
in face view; ч) spores seen edgewise.

(d) small mature spores seen

238 MONOGRAPH OF CHAETOMIUM AND ASCOTRICHA

CHAETOMIUM ANGUSTUM Chivers

Fic. 6. Mature perithec
Ре. A A straight rg hair. For convenience the hair is figured in two

Fic. 8 A Mines terminal hair spirally coiled near its tip.

Fic. 9. А stout terminal hair spirally coiled near its tip.

Fic. 10. Ап irregularly convolute terminal hair with a branch. For convenience
the hair is figured in two parts

cus.
Fic. 12. Spores. (а) Young spores with oil globules; (b) and (c) mature spores
seen in face view; (d) seen edgewise.

PLATE 17

ASCOTRICHA CHARTARUM Berkeley
Fic. 1. Mature perithecium.
Fic. 2. A perithecium developing from the GASE itae strand, X I5..
Fic. 3. Terminal hair, showing its ampullate swellin
Fic. 4. Terminal hair of slightly different а
Fic. 5. Conidiophore with its clusters of conidia, X 180.
Fic. 6. A portion of a conidiophore under greater magnification, Х 390.
Fic. 7. А small portion of а conidiophore with four conidia, X 390.

IG: 8. Conidia, X 390.

Fic. 9. Се developing from the оа mycelial strand, Х 25.

І. DH
Fic. 12. Spores. (а) Seen in face view; (5) seen edgewise showing their girdle.

ASCOTRICHA PUSILLA (Ellis & Everhart) Chivers
Fic. 13. Mature perithecium
Fics. 14, 15. Terminal hairs жены their ampullate swellings.
Fic. 16. Conidiophore with its cluster of conidia, Х 180
Fic. 17. Conidiophore of а slightly different character, Х т
Fic. 18. А portion of a МОТА under greater оман Ч Х 780.
Fic. 19. Conidia, X 300
Fic. 20. Ascus.

Fic. 21. Spores. (а) and (6) seen in face view; (c) edgewise.

names, the final

INDEX

members of new combinations, and the numbers of the

ew
pages on which the descriptions are to be found are in bold fa

адет EE Chaetomium, 210
та

Anixi сы enoides,

Ascotricha eog

posi, БА 210, 220, 224
a Alche millae, 211
Bom

merella oe pc 166
Caclospheria exilis,
euthos phaeo

Chaetonidium fimeti, 161, 186,
213,

е а ш, gen 216
affine sete
Alchemillae, 2
amphitri dee: um 193
ampulla, ode 207

mda de 217
Arali

185,
m m, 180
„маг, Therryana, 180,
aureum, 174, 176
B

175,
artholomaei, е 177
Berkeleyi, 22
bostryc

h аг ag 162, 200, 201, 202,

Braunii, 212, 213
Bromelliae, 2

calvescens, 214, 15
caninum, бт, 201, gar
caprinum, 162, 203,
chartarum, 157, s d зо, 191, 193,
196, dais > 205, 222, 223
ciliatum, 212, 213
circinans, ec? 21I
ircina I
coccodes, 210
cochliodes, 161, Е 205, 206

т99 e
matum, 180, 182, 183, 184,

comatum, var. helicotrichuim,

167,
comatum, var. ligni, 180, 1
comosu
concinatum, ara, 2
contortum, т б D фы

214
arum, 162, 108, 216, 218,
Ss

104. 199,

186, 215
, 182, 185
182, 186,

186, 209
166,

182, 185

170, 172

Chaetomium ае
раё

173

ir aen 210, 211

exile,

Fiebti, Th 184, 190, 192, 193. 194,
196, 198, 1

pce var. chartarum, 180, 182,

190, te

formosum. 162, 201,2
formos m, var. leui у 162, 201,

formosum, var. ovatum, 162, 201,
funicolum, 160, 176, 177, 178, ы

18
fusiforme, 174, 175, 176

2
199, 205, 206, 207. 208,
obosum, var. 1
аааз ат 181, 182, 184
minis, 18 à 84
um, 167
hipte 212
oe |
are mc? p^ 162, 178. 179, 180,
unzeanum, 161, 167, 186, 189, 190,
192, 193. A 1 e =

240 INDEX

Chaetomium Kunzeanum, var. fimicolum, | Chaetomium spirochaete, 161, 199, 200
st

er ,
laeliicola, 216, Streptothrix, 171
lageniforme, a, 162. 184 strigosum, 210, 212
lanatu subnudum, 218
lanosum, 162, 186, 190, 192, 194, 199, subspirale, 1
tomentosum, 212, 2I
eroski с? oe 185 tortile, 162, 169, 170, 172
озроги 0, 195 torulosum, 162, 20
marchicam yas nio trigonosporum, 166, 203
meg 162, 190, 198 co a SE 175
е ош. Gg 178, 179 Тур
microsporum, T ug
Montagne, 218 undulatum, pris I9I, 198
amc ans 21 vari о
rum, 160, GE 166, 167, 168, velutinum, 181, 185
уду 186, 195, 196, 199, 213, 217, viride, 2
21 Zopfii, 2 223
‚ 213 Coleroa Alchemillae, 2II
m, 162, 186, 100, 195, 199 Chaetomium, 211
на, var. chartarum, Е. & E., Potentillae, 212

Con а 181, 183:
olivactum, f. chartarum Roum., 191 | Dicyma ampullifera, 22
I Dine

oospora, 101, I97 rium ciliatum, 213
orientale, 191, 194 Dothidea vae ag aic 211
oxysporium, 210, 211 8, ZII
pallidum, 215 Chaetomium, 211
nosum, 161, 181, 182, 184, 185, Potentillae, 212
186, 21 rium Chaetomium, 21
papillosum, 217 yxotrichum chartarum, 183
ucisetum, 214 Niesslia EN 210
Polypori, 214 exilis,
Potentillae, 210, ec
pusillum ES зА 218, 210, 220, | Nitschkia exilis,
Oospora crustac
pusillum Fries, 2 Orbicula perichaenoides, 214
quadran — atum, p 202 P тераа infestans, 171
raripilum, 213 ima Me
rigidulum, 1 Sheets ү. ena 210
rostratum, 204, 209 Sphaeri mata, 181, 183, 208
сат Sphaeria crispata,
setosum Бай: gei тот, 108 Sphaeria exilis, 21
setosum E. & E., ы Sphaeria rioi
setosum Wint., e 178, 179, 180 Sphaeroderma marchicum, 216
si natum, 212, 213 h trys lobu
imile, 161, 169, 170, 172 Stigmatea Alchemillae, 211
sphaer: haetomium,

Cha
sphaerospermum, 162, 210, 222, Potentillae, 212
22 Venturia Alchemillae, 211
spinosum, Chaetomium, 210

pirale, 161, 199, 200
spirilliferum, 162, 19r, 198

nzei,
Potentillae, 212

Mem. Товкку Сілув VOLUME 14, PLATE 6

IDT

1-5. CHAETOMIUM ELATUM Ком2Е & ScHM

` VERS
6-11. CHAETOMIUM TRIGONOSPORUM (MarcHat) Сн!

Мем. Torrey Сгов VOLUME 14,

1-8. CHAETOMIUM INDICUM CORDA

TNI Т ЧЕ
9-19. CHAETOMIUM FUNICOLUM Cook

PLATE

/

Мем. Torrey CLUB VOLUME 14, PLATE 8

E

1-5. CHAETOMIUM CIRCINATUM CHIVENS

6-10- CHAETOMIUM MURORUM Corps

Мем. Torrey Стов VOLUME 14, PLATE 9

3

Ce
%

а: =
x

NAM ақы

CR О “

OLS
SAS
SA

т ге & SALMON
1-4. CHAETOMIUM SIMILE MassEE X 9^

РАТИ TUCKEL
сә. CHAETOMIUM CRISPATUM p s:
Е "т TUN
10-12. CHAETOMIUM CONTORTU®. s
13-17. CHAETOMIUM TORTILE BAINIE

ECK

Мем. Torrey Стов VOLUME 14, PLATE 10

IVERS

1-7. CHAETOMIUM SPINOSUM Си

^ CHIVERS

8-12. CHAETOMIUM AMPULLARE CHIVE
UT КОМЕ

13-16. CHAETOMIUM GLOBOSL M Kt

Мем. Torrey CLUB
VOLUME 14, PLATE II

\TERALE CHIVERS
ISIFORME CHIVERS
UREUM CHIVERS
SPHAERALE CHIVERS

1-6. CHAETOMIUM TRIL:
1-11. CHAETOMIUM FL
12-17. CHAETOMIUM A
18-23. CHAETOMIUM

Mem. T
Mem. Torrey Сто
z -UB
VOLUME
UME 14 PLATE
1 ATE 12

Lis & ЕУЕВНАКТ

CHAETOMIUM ATER RIMUM Er

I-4.
ДОРЕ

5-8 ` цер
5-8. CHAETOMIUM SPIRALE

Мем. TorREY CLUB VOLUME 14, PLATE 13

c х?
АНАМ
ЖЖ ARS
{2

42

d 2 nm
=> журу?
тт РИ
ХЕ AOA

CHAETOMIUM BOSTRYCHODES ДОРЕ
SUBSPIRALE CHIVERS

CAPRINUM BAINIER

1-11.
12-17. CHAETOMIUM
18-22. CHAETOMIUM

Mem. Torrey Стов VOLUME 14, PLATE 14

EISE ws
dee
w е

KU
Kar

-1
=
ғ“

га

1-8. CHAETOMIUM QUADRANGLE LATUM CHIVER
UTUM CHIVERS

9-12. CHAETOMIUM CONVOL

Мем. TORREY CLUB \ ) ME LATE
OLUME I4, E 15

Ж; ЖАНА

22 ys N

ХА

2
A
+9 а = ` f
CX)
A ДҚ nih 2
қазы éi dh
VS, $,
4% жам
(b

З /
722
VC

Sy

NC

А у
wan
apt

са. Cé а
22 УЕ =

422 SR

TOMIUM COCI ILIODES PALLISER

1-8. CHAE
UM GLOBOSUM KUNZE

9-14. СНАЕТОМІ

Мем. Torrey Crus V
OLUME 14, PLATE 16

2

——À

s е
A
/ “ЙІ! VANNS

1-5. CHAETOMIUM TORULOSUM Вліхтек
6-12. CHAETOMIUM ANGUSTUM CHIVERS

Мем. TonREY CLUB

VOLUME 14, PLATE 17

D

Dee АРЕН pem

WENT.
nni Me Wi Mc Ж

un

ay x

Yi

М
va

А

1-12. ASCOTRICHA CHARTARUM BERKELEY |
13-21 ASCOTRICHA PUSILLA (ELLIS & EVERHART) CHIVERS

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