Annals
of the

Missouri Botanical

Garden

Volume ХП ^
1925

With Twenty Plates and Nine Figures

Published quarterly at 8 West King Street, Lancaster, Pa., by the Board of Trustees
of the Missouri Botanical Garden, St. Louis, Mo.

Entered as second-class matter at the Post-Office at Lancaster, Pennsylvania,
under the act of March 3, 1879,

MISFOUR! BOTANICAL
GARDEN LIGRARY

Annals
of the

Missouri Botanical Garden

A Quarterly Journal containing Scientific Contributions
from the Missouri Botanieal Garden and the Graduate Labora-
tory of the Henry Shaw School of Botany of Washington Uni-
versity in affiliation with the Missouri Botanical Garden.

Editorial Committee
George T. Moore Benjamin M. Duggar

Information
The Annals of the Missouri Botanieal Garden appears four times during
the calendar year: February, April, September, and November. Four numbers
constitute a volume.

Subscription Price - - - zs 00 per volume
Single Numbers - - - - 00 each
The following agent is authorized to accept foreign subscriptions:
Wheldon & Wesley, 2, 3 and 4 Arthur St., New Oxford St.,
London, W. C. 2, England.

STAFF
OF THE MISSOURI BOTANICAL GARDEN

Director
GEORGE T. MOORE.
ВЕМЈАМІ M. Duaaar, EDGAR ANDERSON,
Physiologist, in charge x Geneticist.
Graduate Laboratory
VON SCHRENK, ELVA PUMPHREY MILLER,
Pathologist. Research nt.
JEssE M. GREENMAN, KATHERINE H. LEIGE,
Curator of the Herbarium. Secretary to the Director.
NELL C. Horne

Librarian and Editor of асе.

BOARD OF TRUSTEES
OF THE MISSOURI BOTANICAL GARDEN

President,
EDWARDS WHITAKER.
Vice-President,
EDWARD C. ELIOT

Second Vice-President,
GEORGE C. HITCHCOCK.

SaMvEL С. Davis. А. C. Е. MEYER.
Tuomas S. MAFFITT. PuiniP C. SCANLAN.
Epwarp MALLINCKRODT. Jonn Е. SHEPLEY.

EX-OFFICIO MEMBERS:
Henry Е. FAHRENKROG. FREDERICK Е. JOHNSO

— of the ina of Education Bishop of the DER. of Missouri
o

HERBERT S. HADLEY VicroR J. MILLER,
Chancellor of Washington University. Mayor of City the of St. Louis.
Grorce T. Moore

President of The Academy of воне of St. Louis.

TABLE OF CONTENTS

Monograph of the Genus Monardella. .Carl C. Epling
Studies on South American Labiatae. I
ОИ ИОД Carl С. Epling

Colloidal Sulphur as a Spray Material. . Н. С. Young
Pigment Studies with Special Reference to

Carotinoids in Fruits........... Grace E. Howard
The Thelephoraceae of North America.
СООО Cli E T E HE EL E. A. Burt

The Effect of Treating the Virus of Tobacco
Mosaic with the Juices of Various Plants
jx rd B. M. Duggar and Joanne K. Armstrong

Determination of Total Nitrogen in Plants
and Plant Solutions: A Comparison of
Methods with Modifications...... Emery R. Ranker

Colloidal Sulphur: Preparation and Tox-
NES Л Л УУЛ Lo UAR ei Se Lionel E. Tisdale

PAGE
1-106

107-132
133-143

145-212

213—357

359-366

367—380

381-418
419-424

Annals
of the
Missouri Botanical Garden

Vol. 12 FEBRUARY, 1925 No. 1

MONOGRAPH OF THE GENUS MONARDELLA'?
CARL CLAWSON EPLING
Instructor in Botany, University of California, Southern Branch

Formerly Rufus J. Lackland Research Fellow in the Henry Shaw School of Botany
of Washington University

INTRODUCTION

The following paper records the results of a study of the species
of Monardella, a Labiate genus of western North America. The
object of this investigation was to ascertain the relationships and
geographical distribution of the various elements which con-
stitute this natural group of plants. The work was done mainly
at the Missouri Botanical Garden during the years 1922-24,
but during its course the collections of the genus at the Field
Museum, the National Herbarium, the Gray Herbarium, the
Rocky Mountain Herbarium, the Colorado State Museum, the
herbaria of the Universities of Colorado, California, and Wash-
ington and of Leland Stanford University, Pomona College, the
Oregon Agricultural College, and the private herbarium of W. L.
Jepson were studied, in part by loan, in part by visit to the places
concerned. Reference is not made to all herbarium material
examined. Full citation of specimens is given, however, when
the citation concerned is of particular historical interest, when
geographical ranges are extended, or for other cogent reasons.

The author is indebted to the curators of the herbaria in which
material has been studied or from which loans have been made;

1 An investigation carried out at the Missouri Botanical Garden in the Graduate
Laboratory of the Henry Shaw School of Botany of Washington University and sub-
mitted as a thesis in partial fulfilment of the requirements for the degree of doctor
of philosophy in the Henry Shaw School of Botany of Washington University.

? Issued September 28, 1925.

Axx. Mo. Bor. Ganp., Vor. 12, 1925 (1)

[Vor. 12
2 ANNALS OF THE MISSOURI BOTANICAL GARDEN

it was only through their considerate coóperation that so repre-
sentative a collection of the genus was brought together. He is
partieularly indebted to Dr. George T. Moore, Director of the
Missouri Botanieal Garden, for the privileges of the splendid
library and herbarium of that institution, and especially to Dr.
J. M. Greenman whose helpful advice and unfailing courtesy
have both aided the progress and added very greatly to the
pleasure of the work.

HISTORY or THE GENUS

Michaux,! in 1803, described and illustrated a certain plant
which he named Pycnanthemum Monardella, the habitat of which
was said to be the high mountains of Carolina. The corolla was
neither described nor illustrated and in an observation at the
end of the description it was stated that the plant was of the
habit of Monarda fistulosa and that neither it nor the species im-
mediately preceding it were strictly congeneric with the other
members of the genus to which they were there assigned. The
species referred to was Pycnanthemum montanum Michx.

Bentham? later made use of the suggestion contained in Mi-
chaux’s comment and established the genus Monardella, using
for the generic name the specific designation employed by
Michaux. He renamed Michaux’s plant Monardella Caroliniana
and included within this new genus the second doubtful species,
Pycnanthemum montanum Michaux, which was renamed Monar-
della montana, and three previously undescribed plants collected
by Douglas in northwestern America, which were named M.
odoratissima, M. undulata and M. Douglasii. In describing the
renamed species, M. Caroliniana, Bentham referred to it the
plant described by Michaux and a plant described by Elliott,
repeating their descriptions with quotation marks but giving no
indication by the customary abbreviation, v.s." that he had
seen their specimens. Ор the contrary, he says: * Pycnanthemum
Monardella Pursh seen in Lambert's herbarium is very similar to

! Michaux, A. Flora санав 2:8, pl. 34, 1803.
? Bentham, С. Labiatarum genera et species, p. 331.

s 1834.
3 Elliott, S. Sketch of the Бома. of South Carolina and Georgia 2: 81. 1824.

1925]
EPLING—MONOGRAPH OF MONARDELLA 3

Monarda fistulosa but the corolla is wanting; thus also in Mi-
chaux's drawing; I have not seen Elliott’s specimen." !

Fourteen years later, Bentham,? who in the meanwhile had
seen additional material not available at the time of the first
revision, definitely referred Pycnanthemum Monardella Michx.,
which he had renamed Monardella Caroliniana, to synonymy
with Monarda fistulosa L. and remanded Pycnanthemum mon-
tanum to the genus to which it had been assigned originally.
Considering the extreme paucity and fragmentary nature of the
specimens with which he worked and the similarity of the calyces
of Pycnanthemum montanum, Monarda fistulosa, and Monardella
odoratissima, the previous error is not to be wondered at. As
reconstituted in 1848 the genus included four species of closely
related plants, namely, Monardella odoratissima, M. undulata,
M. Douglasii, and M. villosa, nor was a species of Monardella as
at present known or as known to Bentham, included in any
other genus by him. The genus thus described and constituted
was accepted by Gray? who enlarged it by the publication of six
new species and divided it into two sections. Later authors `
having to deal with the group accepted it in the same sense.

In 1906 Greene‘ reviewed the history of the genus, as outlined
above, but went no further than the original monograph of
Bentham, and in conclusion made the following statement: “ And
while later authors have remanded the type of Monardella to
an older genus and an older species even, the name has been
retained for what is now a large genus of western plants. The
viciousness of this method in nomenclature I long ago attempted
to point out; and I here, after long delay, propose a new name
for the western genus: a name made out of the old Monardella,
that is Madronella. I shall not attempt to transfer more than a
portion of the species, but here is a considerable number of them,
placing first in order what should be the type of the genus.”
There followed a list of the species described by Bentham, Gray,

1“Species dubia. P. Monardella Pursh! in Herb. Lamb. Monardae fistulosae
simillima est, sed corolla desunt; sic etiam in icone Michauxiana; plantam Elliot-
tianum non vidi." : y

2 Bentham, G. in De Candolle’s Prodromus systematis vegetabilis 12: 191. 1848.

3 Gray, A. Proc. Am. Acad. 11: 100. 1876
4 Greene, Е. L. Leaflets of Bot. Obs. and Crit. 1:168. 1906.

[Vor. 12
4 ANNALS OF THE MISSOURI т BOTANICAL GARDEN

and Greene, at the beginning of which was the species M. odora-
tissima.

By the phrase “type of Monardella” Greene referred to the
species Monardella Caroliniana which was the first cited by
Bentham. Even had Bentham been familiar with the more
recent concept of a ‘‘type” it is very doubtful that he would
have chosen as the type of a new genus a plant which he had not
seen and which he termed “species dubia." Furthermore, by
reason of the fact that Bentham himself, as early as 1848, and
not later authors," as erroneously stated by Greene, limited
the genus as at present understood, and since the genus was ac-
cepted in this sense for fifty-eight years without question, the
action of the latter is indefensible. It may be further observed
that the original complete and expressive generic diagnosis,
based upon a few battered plants, is still exactly applicable to
the genus which has been enlarged at least fivefold in number of
valid species. The adoption of M. odoratissima as the generic
type-species was a desirable step because it was the first described
of the true Monardellas and because it is the most widely dis-
tributed of any of the species.

Since 1906 the names Monardella and Madronella have both
been in use to describe the same group of plants, and the species
may be found listed under both names in the ‘Index Kewensis.'
Sinee the publieations of both Bentham and Greene are not
readily aecessible, no small amount of confusion has been oc-
casioned.

The first subdivision of the genus, as already mentioned, was
by Dr. Gray, in 1876, in a synopsis in which he described as one
section “‘ Macranthae laxiflorae nempe floribus in capitulo laziusculo
sat magis minus numerosis: corolla e calyce longe exserta: antherae
loculis ovali-oblongis divaricatis: perennes," including therein M.
macrantha and M. nana, and a second group “ Densiflorae et
multiflorae: calyce X4 to Y4 pollicari: antherae loculis brevioribus
minis divaricatis." These subdivisions were adopted by Briquet!
in his presentation of the family in *Die Natürlichen Pflanzen-
familien’ under the name Macranthae and Pycnanthae, a ter-
— ыш J. in Engler u. Prantl, Die Natiirlichen Pflanzenfamilien. IV. 3a. 309.

1925]
EPLING—MONOGRAPH OF MONARDELLA 5

minology which was followed by Abrams! in his revision of the
southern California species of Monardella.

MORPHOLOGY

The Monardellas are either annual or perennial. The annual
species are erect, herbaceous plants from six inches to two feet
in height, arising from a short tap-root from which spring small
lateral rootlets. They are, for the most part, plants of semi-
xerophytic habit, the leaves being few, small, rather thick, and
variously pubescent. The stems are columnar, obscurely four-
angled, slender and branched. The character of the branching
is of two kinds, terminal or basal. In the first type as illustrated
by M. candicans or M. lanceolata the branches occur chiefly in the
upper axils and are widely divaricate, but ascending, and when in
full flower form thus a corymbose group of inflorescences. The
second type, as illustrated by M. Breweri and M. exilis, are
branched throughout with the principal branches arising near
the base of the stem and ascending more or less parallel to it,
being themselves either simple or branched. Both types may
be seen in the same species, yet under the ordinary conditions of
its habitat a given species is characterized by one type.

The leaves of the annuals show little diversity, being of about
the same range of size, lanceolate or oblong, entire, shortly
petiolate and spreading, and subcinereous with a short close
pubescence or glabrate. The chief exception to this condition
is shown by the leaves of M. undulata. Here the leaves are ob-
lanceolate and markedly undulate or crisped. The venation is
pinnate but rather obscure in the annuals.

The stem of the perennials is generally decumbent, occasionally
trailing or even somewhat subterranean, as in the case of M.
macrantha. It is apparently, in many cases at least, a modifica-
tion of the first stem to be formed, which after flowering dies
back only part way to the base. In some species, as in M.
hypoleuca, the stem may become elongated to several feet, trailing
over and supported by brush. The branches arise from the
stem, being either distributed along it in a candelabra-like way,

1 Abrams, L. В. Mublenbergia 8; 26-44. 1912.

[Vor. 12
6 ANNALS OF THE MISSOURI BOTANICAL GARDEN

as often seen in M. linoides, or else from a well-defined crown as
in M. odoratissima. It is evident from the nature of the dis-
tribution of the perennial species that the habit is frequently a
function of quite varied environmental conditions. Like the
annuals, the stem of the perennials is provided with a stout tap-
root. In old plants the bark, which is smooth and brown, be-
comes checked and flakes away. The branches are variously
pubescent above and glabrate below. They may be either erect,
ascending, or decumbent, and either simple or branched, but when
branching the secondary branches are short, slender, and seldom
fertile. Their habit is more or less characteristic of the species.

As contrasted with the leaves of the annuals, the leaves of the
perennials are diverse in form, covering, and size. They may be
entire or serrate upon a single plant. In general, however, there
is a certain aspect about the leaf which permits its identification
with a certain species. In shape they may vary from ovate to
oblong, the extremes being, on the one hand, rotund, and, on the
other, linear-oblong; they are more frequently petiolate than
sessile, the petioles being always short in proportion.

The pubescence of the stem and leaves is equally diverse, being
silky-villous, woolly-villous, tomentose or hirsute, canescent or
cinereous, or may be very minute but dense, so as to cause a
silvery or glaucous appearance. The pubescence is of value in
distinguishing subspecies but by reason of its response to the
environment must be used with care as a basis for specific dif-
ferentiation. In general a close puberulence or pubescence is
characteristic of the forms of the drier interior, while forms ex-
hibiting a looser, more villous or tomentose covering are to be
found in closer proximity to the coast-line. The pubescence of
the stem is usually retrorse, but may occasionally point upwards.
In a similar way the trichomes of the upper surface of the leaves
may point either to the distal or proximal end. This fact has
been utilized by Abrams in separating two closely related forms,
but from a microscopic examination of copious material it is
clear that this character cannot be relied upon to effect other than
an arbitrary separation, for it has been observed that in numerous
cases the pubescence of the lower part of the leaf may point
downward, while that of the upper part may point upwards. At

1925]
EPLING—MONOGRAPH OF MONARDELLA T

the same time material from a given locality which is strikingly
similar in other respects and hardly to be separated may exhibit
both types of pubescence characters as shown by the collection
of M. linoides made by Purpus on Pah Ute and Argus Peaks. A
similar condition has been observed in other mints, as, for ex-
ample, the coastal forms of Monarda punctata.

Yellowish glandular punctations are common upon the leaves
of the genus, and in some cases stalked glands may be present.
The former are more apparent in some plants than in others,
often being obscured by the pubescence. The punctations vary
somewhat in dried material, chiefly in the degree to which the
leaves become pitted. Their size remains fairly constant with a
certain degree of variation in the frequency of their distribution.
No use of them has been made herein.

The inflorescence of the genus consists of a compact globose
head or glomerule of flowers borne terminally upon the branch
and subtended by a series of bracts arranged in more or less
opposite pairs. Rarely, two glomerules may be present, a smaller
one above the first. This has been observed only in M. undulata
and in M. odoratissima. The flowers are attached by a short
pedicel to a small disc-like structure which terminates the branch
and which may be considered a foreshortened and modified cyme.
Only the outermost flowers are bracteate. The bracts are of an
oval or lanceolate shape and appear membranous or foliaceous,
with a pinnate venation. They are of especial value in specific
diagnosis, since their form and consistency, their size and arrange-
ment relative to the calyces, their venation, and the nature of the
margin and pubescence are accompanied by well-marked dif-
ferences in other parts and within a given group of closely related
forms remain fairly constant throughout a wide range of dis-
tribution.

In consistency the bract varies from a white papery mem-
branous structure to a fleshy green form hardly separable from
the leaves of the stem. In any case the outermost are more
likely to be foliaceous than the inner. In some species the bracts
are erect and sheathe the cluster of flowers; in some, however,
they are partly or wholly reflexed. Few exceed the calyces by
more than half their length, and few are shorter than the calyces.

[Vor. 12
8 ANNALS OF THE MISSOURI BOTANICAL GARDEN

While the venation is ultimately pinnate in all, in some the mid-
vein is foreshortened to such an extent that it appears to be
wanting, and the lateral veins appear parallel. Frequently the
margin, which in most species is firm and green, becomes thin,
scarious, and whitened.

The calyx is tubular, in most species one-fourth to one-fifth as
wide as long, and bilabiate in most but appearing equidentate on
casual examination. The bilabiate condition of the calyx-limb
is especially noticeable in the subgenus Macranthae. The teeth
are somewhat shorter than the width of the tube and narrowly
triangular in form. The veins of the calyx are prominent but
hardly costate, giving it a striate appearance. While the aspect
of the calyx is much the same throughout, a careful study reveals
the fact that numerous small differences may be observed and
that within the limits to be expected are quite constantly associ-
ated with other characters. This is especially true of the ar-
rangement and number of the veins which vary from 10 to 15 in
the genus, and the conformation of the calyx-teeth and their
margins.

In the simplest case, the veins of the calyx terminate at the
apex of each tooth and at the base of each sinus, being ten in
number. In calyces with thirteen veins each of the two shallower
sinuses is provided with only one vein, while the three additional
veins are paired with each of those in the remaining sinuses. In
calyces with fifteen veins, in addition to the five veins terminating
in each tooth, each sinus is the terminus for a pair of veins. A
species characterized by calyces with ten veins will usually show
within the same glomerule calyces with eleven veins, rarely with
twelve; calyces with typically thirteen veins will be found to
vary between twelve and fourteen; those with fifteen will vary to
fourteen, rarely less. If a number of flowers from a given species
be examined the number of veins in the calyces will be found to
center about one of the three modes indicated.

In some species the margin of the tooth, which is ordinarily
bordered by a vein, becomes scarious and white and in a single
species prong-like.

The pubescence of the calyx is fairly constant but not suf-
ficiently so to offer a means of infallible specific diagnosis in the

1925]
EPLING—MONOGRAPH OF MONARDELLA 9

case of the perennials. This was shown to be the case by a
microscopic study of various closely related forms, judged by
other considerations, which nevertheless showed under the miero-
scope a considerable difference in the degree to which certain
trichomes might be developed, thus causing an apparent differ-
ence in kind when examined with a lens or the naked eye. The
- character is of value in separating subspecies, however.

In the nature of the corolla the genus presents several differ-
ences not readily observed unless subjected to careful exami-
nation. Thesubgenus Macranthae is characterized by the unusual
proportions existing between the corolla-tube and the limb, on
the one hand, and the corolla-tube and the calyx, on the other, as
well as the fact that the lobes of the corolla taper evenly to a
point. In the case of the subgenus Pycnanthae, the corollas are
much the same size, the size being nearly constant for a given
species, yet present valuable diagnostic indices in the degree to
which the lips are lobed and the shape of the lobe. The corolla
is definitely bilabiate, the posterior lip being two-lobed, the an-
terior three-lobed. The lobes are linear-oblong or linear-lanceo-
late. The shape of the lobes, whether nearly equal throughout
and ribbon-like or whether tapering noticeably, whether blunt or
whether rounded to a point, and the degree to which they are
coalesced, are characters which have been found fairly constant
for most of the species and are of help in specific differentiation.
The degree of exsertion of the corolla-tube from the calyx is also
of value but must be employed with caution. The presence or
absence of a retrorse pubescence within the tube at the base of
the stamens may be used as a specific character but when present
such a pubescence is variable.

The stamens are four in number, the two anterior exceeding the
posterior. They are attached to the corolla just within the throat,
their position varying but little. The filaments are fairly stout
and present little or no difference. They are often retrorsely
hispidulous but when this condition is present it is quite variable
on the same plant, and not infrequently a single filament may be
pubescent nearly to the apex while the remaining three are
glabrate.

The anthers and connective and the relation between the two

[Vor. 12
10 ANNALS OF THE MISSOURI BOTANICAL GARDEN

serve admirably in some cases as characters for specific differen-
tiation, and in the case of Macranthae are subservient to sub-
generic differentiation. The anther-sacs are two, subparallel or
divergent, being subconfluent above when markedly divergent.
In the annuals the degree of divergence, which is a function of the
development of the connective, is in general less and the connect-
ive is less developed than in the perennials. In the former the
shape of the anther-sac after dehiscence, together with the ap-
pearance of the connective, especially the conformation of its
lower margin, was found to be somewhat diverse among the
species but constant for a given species. In the perennials,
however, the connective presents much the same appearance in
all the species, being approximately equilateral when viewed
from the front, except in the subgenus Macranthae where the
angle of divergence approaches that of 180 degrees.

The ovary is four-parted, forming at maturity four smooth
brown nutlets, basally attached, oblong or oval in outline and
somewhat flattened. These are about two millimeters in length
and present few differences. The style is about the length of the
corolla, unequally bifid at the top and glabrous.

Considering the genus as a whole, the following morphological
criteria have been found most trustworthy in diagnosis of the
species: the habit and foliage within certain limits, the texture
of the bracts, the number of the calyx-nerves and the form of the
teeth, the conformation of the lips of the corolla and the structure
of the anthers.

RELATIONSHIPS AND GENERAL DISTRIBUTION

The species of Monardella, while forming a precisely circum-
scribed and very natural group as a genus, are not themselves so
easily capable of definition. This is particularly true of the
perennials, where very considerable degrees of variation may
occur on a single individual. In the absence of copious material
for study, or through failure to consider variations in connection
with their geographical relationships, or by reason of tendencies
to magnify differences between individuals while neglecting or
overlooking equally important likenesses, many of the variants

1925]
EPLING— MONOGRAPH OF MONARDELLA 11

which may be found in the perennials have been described as
species. If these variants, however, are studied in connection
with one another and in relation to their environment and geo-
graphical and ecological distribution, the distinctions upon which
such species rest will be found to effect no more than an arbi-
irary delimitation. Considered alone, the type specimens of
these species offer quite ample grounds, in many cases, for desig-
nation as species. Considered in connection with numerous inter-
grading and connecting forms, considered as living plants and
not as artifacts, followed from one area of occurrence to the next
throughout the range as organisms in contact with an ever-
changing environment, such variants will be found to merge
imperceptibly.

The criteria which had been described above have been found
to offer a means of separating the genus into certain groups which
are rather clearly defined, and which have a characteristic and
natural distribution, and which suggest at the same time a prob-
able phylogenetic sequence. These groups have been termed
species and in general correspond to the concept known as the
Linnaean species, a category of great value in demonstrating
ecological, geographical and morphological relationships.

To insist that such a category is homogeneous would doubtless
lead to error in numerous cases and is contrary to well-known
evidence; to ignore the variations which occur within such a
group would serve only to prevent a further understanding of it;
to recognize that such a group may be heterogeneous but never-
theless that its members are more closely related to each other
than to the other members of the genus and to so name these
elements that the relationship may be apparent in the name, just
as the relationship between species is indicated in the common
generic name, has seemed to the writer the most helpful course in
the case of the genus Monardella.

To determine the exact relationship existing between the ele-
ments of a species requires detailed analytical and synthetical
experiment. Such a course has not been possible i in the present
study. Proceeding on hol groun attempt has been
made to characterize the components of the polymorphic species
in so far as these components appear to represent certain evo-

[Vor. 12
12 ANNALS OF THE MISSOURI BOTANICAL GARDEN

lutionary tendencies within the species. Such categories have
been termed subspecies, in the sense that the groups thus des-
ignated merge imperceptibly, but when taken in the average
or typical aspect occupy a definite and characteristic habitat. It
is not without caution that the term subspecies has been thus
employed in the absence of genetical data and garden observation
and experiment. By way of illustration it may be assumed that
the several subspecies comprising the species M. villosa are de-
rived from a common stock which was at one time in occupancy
of the approximate area now occupied by the species. By reason
of isolation, or through climatic changes in geological time, or
for other reasons, the species once homogeneous may be thought
of as gradually differentiating into several closely related groups,
each with a characteristic habitat. These groups would form the
subspecies herein described, each exhibiting a tendency of evo-
lution from a common stock, which might very well be still
extant. Such subspecies do not vary merely by the presence or
absence of a single given character, but rather in the accumulation
of numerous small differences. The category known as the
variety has been employed to designate those forms which differ
from the typical, principally in the presence or absence of one or
two morphological characters of the magnitude commonly em-
ployed in taxonomy. Such forms, as herein understood, do not
possess a characteristic geographical or ecological distribution
distinct from the typical. It has been possible thus far for the
writer to bring only M. odoratissima under cultivation. From
the behavior of other Labiatae grown from seed or from trans-
plants from the field, particularly of the genus Monarda, it is
believed that the subspecies as herein described will be found to
be racially distinct and not impossibly a complex of variants
which differ genetically, the subspecies representing the modal
points of such a complex.

Related groups.—Each of the four most widely distributed
species, M. macrantha, M. lanceolata, M. villosa, and M. odora-
tissima, may be considered a center about which the other species
can be arranged according to resemblances. Four species,
of which are endemic, are more or less intermediate or of uncertain
relationship. The groups resulting from such an arrangement
are herein designated as ‘‘Sections” and are as follows:

1925]

EPLING—MONOGRAPH OF MONARDELLA 13
Section I Section III
M. macrantha M. villosa
?M. Palmeri M. lanata
Section II М. hypoleuca
M. viridis
M. lanceolata M saxicola
M. Breweri Vil dada
M. Pringlei м eon
И loucooephala ?М. ii
M. candicans Section IV
M. exilis M. odoratissima
M. Douglasii M. linoides
?M. undulata

The sections are all closely connected, yet the greater and more
significant differences of the first, as contrasted with the remain-
ing three, have led to its segregation as the subgenus Macranthae.
M. Palmeri is apparently an intermediate form. Of the remaining
three sections, the second, while closely knit and showing definite
lines of divergence, is still too closely connected with the other
two to be separated as a subgenus. A brief comparison of the
morphology of each of these groups will be made in order to in-
dicate the possible affinities. The conclusions drawn from such
a comparison are shown in fig. 1.

The habit of the subgenus Macranthae is not essentially dif-
ferent from that of the subgenus Pycnanthae but the stem is
slender and more or less rhizomatous and semi-subterranean,
giving rise to either decumbent or ascending branches, usually
few in number and located at the distal end. In those forms
which occur in acerose forests the stem trails along beneath the
detritis of the surface layer. In the forms of more xerophytie
habit this distinetion is partly lost, and in M. macrantha var.
arida, the variety most adapted to a xerophytic habitat, a definite
crown is formed and the branches often rebranch at the base,
thus forming a small tuft: the habit, in other words, of the sub-
genus Pycnanthae. It may be said, then, that there is a tendency
on the part of the most restricted of the species of this subgenus
to assume the growth-form which characterizes the perennials of
the subgenus Pycnanthae. Macranthae are further characterized

[Vor. 12

ANNALS OF THE MISSOURI BOTANICAL GARDEN

14

#9ртоцутпә *doqne

BL Bydeoooney, “N
LL L———————

9TlTXO*W

EM
v3wroseouvl'A | ———

|

suvoTpuvo*y
genuuy
A, | go 24f303 044
14949498 * N А

тэїҖитїд`'и

Vreer3noq *

: /

Jeu tse Tavropons *deqng

ETT 028]p'dsqns

von?[$ *deqne

m
unxojeupd*deqng

*TI93 TAxvd*deqne

Sr[9149nv*doqng

вәртоптт ‘и AN

jesse и

9911 $$13910pQ
40 99А3о3024

vuvogrou?4] * 19A
weo[[1TAna*aeqng

93919904719 'dsqne

4031943 * deqns |

33 21Заиц * 43418

Е: EA

wre:
8101} muag’ ed
wuwu*deqns

TEIUSH'XvA

e

que 1oeuno*deqne

| зтоотхез'н |
| S8TPTX TA*W

| v3vuvi ‘и En

еы

SS H |

»dK301044

| anuon |

А

| AVHLNVNOAd

r A

| BTL OF WAY *w Es TA t e энек кыре ig LU UT:
Ы m Um pee 3° 906304014

үдәшгед * н |

$0 244303704 d

fees
ED
Hi
te d
РЕ АЕ
TE
SER
SPFS
2533

:

3

g

In form and size of leaf there is great di

even on individual plants,

a hardly definable aspect which suggests

by a more coriaceous leaf than is genera

1925]
EPLING—MONOGRAPH OF MONARDELLA 15

In addition to the vegetative habit, the habit of the inflorescence
is distinctive. While fewer-flowered, given equal opportunity
for development, the inflorescences in Macranthae are larger in
proportion to the plant, due in part to the larger calyx and
corolla, and give a top-heavy appearance to the branch in typical
cases. The arrangement of the flowers in the glomerule is also
looser. However, the greatest difference between the two sub-
genera lies in the flower. The calyx is actually larger in Macran-
thae and different in proportion, and the corolla is of different
proportions, although in the more reduced forms a casual glance
would not distinguish it from the corolla of M. villosa for example.

A study of the corolla indicates either a progressive reduction
from the large corolla of M. macrantha subsp. eumacrantha to
the small corolla of subsp. nana var. arida, or a progressive de-
velopment in the other direction. It is believed by the author
that the series is à reduction series; first, because the vegetative
habit of the small-flowered species is correspondingly reduced and
adapted to the arid habitat; second, the corollas themselves
strongly suggest a reduction form, whence the specific adjective;
again, in the progressive color changes which accompany the re-
duction in size certain characters are lost rather than gained.

In resumé, then, we may state that M. macrantha is the least
modified of the species of the first subgenus and hence most like
the hypothetical common precursor of the two subgenera. M.
Palmeri, a restricted endemic, is intermediate between the sub-
genera. There may be observed in the subgenus two evolutionary
tendencies, namely, the adoption of a more compact growth-form
and a reduction and modification of the flower and foliage, both
attendant upon occurrence in a more arid habitat.

The annuals form a well-defined and interesting group. They
are all of essentially the same stature, growth-form, and foliage,
except M. undulata, and differ principally in the characters of the
inflorescence, namely, in the texture of the bracts and in the degree
to which the corolla is bilabiate. By reason of the fact that
M. lanceolata is least differentiated, it is looked upon as being
most like the common progenitor of the annuals. From this
basis it is possible to trace three suggestive lines of divergence.
The first to be considered will be that of the bract.

[Vor. 12
16 ANNALS OF THE MISSOURI BOTANICAL GARDEN

As previously stated, the bract of M. lanceolata is less differen-
tiated and not infrequently becomes foliar in nature. It is
lanceolate, acute, pinnately veined, green and opaque, but not
fleshy. Beginning here one may trace through M. Breweri, M.
Pringlei, M. candicans, M. exilis, and M. leucocephala a progressive
reduction of the midvein until the veins appear parallel, ap-
parently arising from the base of the bract, but in reality from a
much foreshortened mid-vein. More or less concurrent with the
reduction of the midvein is a reduction of the secondary veins, as
well as an increasing scarious nature of the bract. In the opposite
direction lies a very curious effect. In M. Douglasii the midvein
and secondary veins have become costate and thickened, the
latter becoming confluent on the margin. The intravenous tissue
has become homogeneous, translucent or even transparent, and
tough, suggesting isinglass. While this would appear an extreme
modifieation it should be noted that in many instances half of the
bract may be truly foliar.

Correlated with the differentiation of the bract is a differenti-
ation of the corolla. In M. lanceolata the corolla is bilabiate, the
lobes of the upper lip being coalesced for two-thirds of its length,
the lobes of the lower lip being free nearly to the base. A pro-
gression may be observed commencing with M. lanceolata through
M. candicans, M. Breweri, M. Pringlei to M. exilis and M. leuco-
cephala in which the corolla-lobes become more coalesced until
those of the upper lip are free for less than a quarter its length
while those of the lower lip are coalesced for about one-third of its
length. At the same time the corolla becomes less and less exserted
until in M. leucocephala it is hardly seen at a casual glance.

Again, the calyx-teeth of M. lanceolata are green, lanceolate and
acute. In M. Breweri and M. Pringlei the calyx-teeth are her-
baceous still, but slender and not infrequently become slightly
mucronate. In М. candicans they are herbaceous but frequently
scarious-margined. In M. exilis the scarious margin is well
developed and eonspieuous. In M. leucocephala the calyx-teeth
are terminated by a whitened recurved prong. It should be ob-
served that each of these lines of development is correlated with
occurrence in a more arid habitat, and since the plants are annuals,
presumably with a shortened vegetative cycle.

1925]
EPLING—MONOGRAPH OF MONARDELLA 17

M. undulata is a variable species of uncertain relationship,
annual in part, suggesting the annuals in some respects, in others
more closely allied to M. odoratissima. The oblanceolate crisped
leaf is unique in the genus. |

We may conclude, then, that the annuals of the subgenus
Pycnanthaeforma natural closely knitted group, showing increased
adaptation to a more arid habitat. Three correlated lines of
divergence may be observed, namely, a progressive modification
of the bract, of the corolla and of the calyx. M. undulata is of
uncertain relationship but lies closest to this section.

The perennials of Pycnanthae may be divided rather arbitrarily
into two sections by the nature of the bract, whether firm and
tending to foliar, or whether membranous. In addition the
corolla of the latter group is generally more bilabiate, the upper
lip being coalesced to a greater extent than in the former. The
first section is typified by M. villosa, the second by M. odoratis-
sima. M. villosa is regarded as being the least modified of any
of the species of this subgenus by reason of the subfoliar nature
of the bract, the slight degree to which the lobes of the corolla
have become coalesced, the bilabiate condition thus being less
pronounced, and because of its generally more mesophytic char-
acter. The occurrence of broadly ovate, obtuse, crenate-dentate
hairy leaves suggests strongly a relationship to M. macrantha var.
Hallii. Such a conclusion is strengthened by a comparison of the
leaves of the endemic M. thymifolia which occurs on Cedros
Island off the coast of Mexico. Facts of distribution support
the assumption that these three species (M. villosa, thymifolia,
macrantha) taken together indicate the probable nature of the
generic prototype.

Section III, as outlined above, is united by the perennial habit,
by the rhomboidal crenate-dentate leaves, in general glabrate
above and variously tomentose beneath, by the subfoliar or
thickened bract, and the little modified corolla. M. hypoleuca,
M. lanata, M. viridis and M. saxicola more nearly resemble each
other than they resemble M. villosa. M. cinerea and M. thymi-
folia are endemics of uncertain position but most nearly related
to M. villosa. The latter suggests the subgenus Macranthae in
the aspect of its foliage. The former suggests M. odoratissima

[Vor. 12
18 ANNALS OF THE MISSOURI BOTANICAL GARDEN

in bract character and habit. М. odoratissima and M. linoides,
while more closely related to each other, are nevertheless very
close to this section. They have been grouped separately, partly
for convenience in illustrating their distribution. Sections III
and IV are more heterogeneous than the two preceding and it
is more difficult to discern any continuous line of development in
them. In general it is true that, with an increasing occurrence
in an arid habitat, the leaves progress from ovate and crenate to
oblong or oblong-linear and entire and become more thickened
and leathery; the bracts from foliaceous become membranous or
chaffy; the lobes of the corolla become more coalesced.

A study of the genus as a whole "pe strongly that it is a
genus of mesophytic origin which exhibit
to an arid habitat.

Distribution.—Generally speaking it may be said that Section I
occupies the mountains of southern California and northern
Lower California; that Section II occupies the less arid parts of
the interior valley of California; that Section III occupies the
coast ranges of California and southern Oregon, while Section
IV occupies the high mountains surrounding the Great Basin.
The more detailed distribution may be better obtained by refer-
= to the accompanying charts than from a verbal description

ere.

In view of previous discussion relating to isolation as a factor
in the origin of species! it is desired to call attention to an apparent
correlation between the degree of relationship existing between
certain components of the genus and the degree to which these
components are associated geographically.

The species in which the widest range of variability is found and
in which the units are least readily defined is M. macrantha. It
is also a species of considerable geographical range, extending as
it does from the Santa Lucia Mountains, near the Monterey pen-
insula to San Pedro Martir in Lower California. The variations
in foliage and pubescence are considerable but are surpassed by

1 Jordan, D. S. The origin of species through isolation. Science N.S. 22: 545-
562. 1905; Lloyd, F. E. Isolation and origin of species. Ibid. 710-712. 1905;
Abrams, L. R. Theory of isolation as applied to plants. Ibid. 836-838. ;
Abrams, L. R. and Smiley, F. J. Taxonomy and distribution of Eriodictyon.
Bot. Gaz. 60: 115-133. 1915.

Lo] A.

1925]
EPLING—MONOGRAPH OF MONARDELLA 19

Fig. 2. Map of San Diego County, California, with an insert of southern and Lower Cali-

ia, showing the distribution of Macranthae; E, subsp. ; H, subsp. eumacrantha
var. Hallii; N, subsp. nana; T, subsp. nana var. tenuiflora; A, subsp. nana var. arida; P, M.
Palmeri. :

[V or. 12
20 ANNALS OF THE MISSOURI BOTANICAL GARDEN

- я 20 119 118 n?
T T T T T- T T
ТЕ E \
Dp ve v
,

116 115
T T

M. ундо

M 123 122 Yai 120 ns a

^ Р g
117 EET 115 c

Fig. 3. Map of California and adjacent Oregon (insert) showing dates ; : E
: " | ‚ showing distribution of M. villosa and allied species:
M. ею subsp. euvillosa; М, M. villosa subsp. neglecta; S, M. villosa subsp. Sheltoni; T, M. villosa subsp. sub
rata; Н, M. hypoleuca; L, M. lanata; A, M. viridis; U, M. saxicola; C, M. cinerea.

1925]
EPLING—MONOGRAPH OF MONARDELLA 21

the variations in corolla size. It was pointed out by Gray! and
more recently by Hall? that, although the extremes in corolla size
within the group are most diverse, they are connected by a con-
tinuous and graded series. As far as the size is concerned this
is true, but as shown by Abrams? there is apparently a certain
qualitative difference present, namely, the shape of the corolla-
tube, as well as a slight quantitative difference in the size of the
anther. These differences are correlated with differences in
foliage and pubescence so that two fairly well-defined groups may
be discerned. These groups were both given specific rank by
Gray (as M. macrantha and M. nana) who later, with the ac-
cession of more material reversed his opinion. They were con-
sidered distinct species by Abrams. Whether called species or
subspecies or varieties, the two groups are very closely connected
morphologically and are very closely associated geographically.
The group designated herein as subsp. ewmacrantha is found in
the Santa Lucia Mountains, the San Gabriel Mountains, the
San Bernardino Mountains, and the mountains of Lower Cali-
fornia at San Pedro Martir. In addition it is found in the San
Jacinto Mts. at low elevations and in San Diego County on
Palomar Mt., in the mountains near Julian, in the Cuyamaca
Mountains and in the Laguna Mountains, but in these places is
in association with the group designated herein as subsp. nana,
both forms apparently occurring in the same locality.

The subspecies nana, on the contrary, is confined largely to the
mountains of San Diego County, occurring in the typical aspect
only in the localities in which is also found subsp. ewmacrantha.
The possible exception may be found in the Orcutt collections at
Japa in Lower California. The author has been unable to as-
certain the location of this place. While being always associated
with subsp. eumacrantha, when in typical aspect, subsp. nana ex-
hibits two fairly well-defined varieties which together have a
separate geographical distribution, being found in the San Jacinto
and Santa Rosa Mountains at higher elevations.

1 Gray, А. Syn. Fl. М. Am., ed. 2, 21: 459 (suppl). 1886.

2 Hall, H. M. A botanical survey of San Jacinto Mountain. Univ. Calif. Publ.
Bot. 1: 109. 1902

3 Abrams, L. R. The Monardellas of southern California. I. Muhlenbergia 8:
26. 1912.

[Vor. 12

ANNALS OF THE MISSOURI BOTANICAL GARDEN

118
T

121
T

123

Ман Pun ot ас

"e

T

(insert), showing distribution of М. linoides: ·

Fig. 4. Map of California and

adjacent Lower California
» M. linoides subsp. stricta,

linoides subsp. eulinoides; S

1925]
EPLING—MONOGRAPH OF MONARDELLA 28

Since subsp. nana and its varieties tenuiflora and arida are
almost certainly derivatives of subsp. eumacrantha or a similar
form now extinct and since the relationship is still very close it is
of interest to find that the range of the former subspecies coincides
with & limited portion only of the range of the latter subspecies
and of particular interest to note that the chief deviation from
the range of subsp. eumacrantha is found in the two most highly
adapted varieties, namely, var. tenuiflora and var. arida.

The section which is next in order in the closeness of the re-
lationships of its components is section IV, composed of M.
odoratissima and M. linoides. This group is also of the widest
geographical distribution. As treated herein the first-named
species is divided into seven subspecies, the second into two sub-
species. In actuality the group represents an almost unbroken
series of intergrading forms, with but & poorly defined hiatus
between M. linoides and M. odoratissima. While the extremes
within M. odoratissima are much less than in M. macrantha,
nevertheless the modal points within the range of variation
stand out more clearly than is true of the varieties and sub-
species of that species. These modal points furthermore repre-
sent plants with characteristic geographical habitats which are
distinct but contiguous. It is chiefly in the intermediate geo-
graphical regions that intermediate morphological forms are
found. The same is true in the case of M. linoides.

M. villosa and its allies present a condition where differentiation
has proceeded further, where the connecting forms have disap-
peared and where the related species are separated by definite
and sometimes considerable geographical barriers. At the same
time there is occurring in M. villosa the same differentiation with
respect to geographical habitat which has apparently taken place
in the formation of this species and its allies. M. villosa extends
from northern San Luis Obispo County to southern Oregon, never
being found east of the Sierra Nevada as far as known. Its sub-
species intergrade but occupy characteristic and contiguous geo-
` graphical habitats. Just as in M. odoratissima, the intermediate
morphological forms are intermediate geographically. The only
other ally which is found in this range is M. viridis. M. viridis,
however, is most nearly connected to M. saxicola which is found

ANNALS OF THE MISSOURI BOTANICAL GARDEN

124

Bria z

МАР 4
Js
Me

ар | 4 NS pA
7 Le ment FOO SAS ,
UE E "AC ENS e ist es Cer
х Ме выс
ү =й ъ®°_ ad
AG rk \ c à
\чк крс е
» э рае d C A ӯ
? Q ae 70, Y A A
s y 1 el Lo 7
5r \ х PD 221 oe

“af
м. 5; P D
/ р ] 1
h ys "d (d oS Я a -*
Lc ERATUA 5: ›
2 ON

A » UA A

MILES

; E, M. iam

4cans.

B, M. Breweri;

.
„

n of Lower California, showing the distribution
C, M. cand:

M. lanceolata var. microcephala

io

L, M. lanceolata; Lm
P ringlei; W, M. leucocephala; D, M. Douglasii.

.
.

g. 5. Map of California, with an insert of a port

Fi
annu als

M.

1925]
EPLING—MONOGRAPH OF MONARDELLA 25

in the San Gabriel Mountains. M. lanata and M. hypoleuca are
more closely connected to each other than to any other species.
M. cinerea and M. thymifolia are species of uncertain relationship,
but most nearly allied to M. villosa. They are very restricted in
distribution, the former being found only on Mt. San Antonio
(Mt. Baldy) in southern California, the latter on Cedros Island
off the coast of Lower California.

The same situation exists in the case of the annuals and is even
more clearly seen there. As already stated it appears that M.
lanceolata is most nearly like that prototype from which these
species have evolved. It has also the widest range, from Lower
California to the Tehachapi, thence along the Sierra Nevada to
Shasta County. Within this range are also to be found the dis-
tributional areas of three other species which are themselves
clearly separated. "With the exception of this overlapping on
. the part of M. lanceolata, the ranges of the other allied species,
seven in number, are distinct. Of these, two, namely, M. leuco-
cephala and M. Pringlei, are very restricted in distribution, the
former being found on the plain of the San Joaquin River near
Merced, the latter in the Jarupa Hills near Colton. The areas
of distribution of all the species save M. Douglasii appear to be
continuous. In the case of this species, however, one area in the
Coast Range and foothills of the Bay Region is separated from the
second in the foothills of the Sierra in Yuba, Butte and Plumas
Counties by the valley floor of the Sacramento River. There
was found no apparent morphological difference between the
inhabitants of the two regions. One specimen of M. lanceolata,
very typical in aspect, was found to have been collected in ‘‘ Mont.
Cr.” Arizona. The location of this place could not be ascertained.
If the label was correct this would represent an unusual extension
of the species. M. undulata, which is most nearly allied to the
annuals, is confined to the coastal hills ranging from Santa Bar-
bara County to Point Reyes. There is nothing to suggest that
its area of distribution is not continuous.

It is apparent from the facts presented above that there does
exist in the genus Monardella a correlation between the degree to
which its components are related and the degree to which they are
separated geographically. It is strongly indicated that such

[Vor. 12
26 ANNALS OF THE MISSOURI BOTANICAL GARDEN

groups of species as the annuals or the allies of M. villosa have
arisen from a common stock which was at one time in possession
of the territory now occupied by the group as a whole and that
such species were formed after isolation of the chief variants within
this common ancestor. It is not inconceivable that even moder-
ate climatic changes on the Pacific slope would effect a similar
segregation of the subspecies of M. odoratissima into forms which,
after a period of isolation, would appear as distinct species of
distinct range.

Center of dispersal.—Adams,'! adopting suggestions from pre-
vious authors, has proposed the following criteria for the deter-
mination of centers of dispersal, in the absence of paleontological
evidence:

*1. Location of greatest differentiation of type

2. Loeation of dominance or greatest abundance of individuals.

3. Location of synthetic or closely related forms (Allen).

4. Location of maximum size of individuals (Ridgway-Allen).

5. Location of eene produetiveness and its relative stability,

in cro

6. Continuity я peces ss of lines of dispersal.

7. Location of least dependence upon a restricted habitat.

8. Continuity and directness of individual variations or modifica-
tions radiating from the center of origin along the highways

dispersal.

9. Direction indieated by bio-geographical affinities.
10. Sic MORE indie ated by annal migration routes, in birds
(Palmén
More recently Liehigiton and Shreve? have modified these
somewhat and have suggested the following as further criteria
applicable to plants:

1. Location of most rapid rate of growth.

2. Location in which a form is accompanied by the largest
number of individuals which are specifically distinct but
of the same growth form.

In the present case it has seemed desirable to restate these

criteria and to limit the group under immediate consideration to

1 Adams, C. С. Southeastern United States as a center of geographical distribu -

“on IE flora and fauna. Biol. Bull. 3: 115-131. 1902.

ivingston, B. E. and Shreve, F. The distribution of vegetation in the United
States as related to climatic conditions. Carnegie Inst. Wash. Publ. 284: 1992.

1925]
EPLING— MONOGRAPH OF MONARDELLA 27

one showing close affinities and a common growth-form. In the
ease of Monardella it seems not improbable that the annual
species have had a more recent origin than the perennial stock;
if this is true the center of dispersal need not necessarily coincide
with that of the perennial species. Since the annuals form a
natural group of close affinity the criteria were applied to them
separately. The species were considered units. Each of the
groups of perennials outlined above was similarly treated. By
means of such integration a closer approximation may be had to
the center of dispersal of clearly related categories. If the
centers of dispersal of such categories coincide, greater confidence
may be felt in stating the center of dispersal for the larger group.
At the same time secondary centers of dispersal may be disclosed
which would otherwise lie hidden.

CRITERIA FOR DETERMINATION OF A CENTER OF DISPERSAL.

Determination of:
. Region inhabited by the most units of the category studied.
. Region inhabited by the units which are most diverse mor-
phologically.
Region indicated as the focus of geographical paths of dis-
persal.
Region indicated as the center of dispersal of the units least
modified.
Region where the most units are most successful as judged
by:
a. greatest abundance of individuals,
b. greatest reproductive activity,
c. greatest vegetative vigor,
d. least dependence upon a restricted habitat.
. Region indicated by known center of closely allied categories.
. Region indicated as the center of evolutionary tendencies in
development as shown by progressive modification or in-
ereasing adaptation to a certain habitat.
These criteria were applied independently to the four sections
outlined in preceding paragraphs. Since the application of cer-
tain of these criteria presupposes a field study it was impossible to

Lung e Ne

са

— ©

[Vor. 12
28 ANNALS OF THE MISSOURI BOTANICAL GARDEN

draw any conclusions regarding these. A more extensive study
of the species in the field, following the lines of variation of each
species, is necessary for a satisfactory solution. From the con-
clusions which it was possible to draw it would appear that the
present center of dispersal of Monardella lies in California south
of the 35? parallel. There appear to be two subordinate centers,
each of which corresponds to the center of distribution of a sub-
genus: Macranthae centering in the mountains of San Diego
County, Pycnanthae in the mountains and foothills of Los Angeles
and Ventura Counties. The results obtained are shown in tablet.

It is generally believed that the western margin of the North
American continent had assumed substantially its present outline
by the beginning of the Eocene,! which was marked by a tropical
or subtropical climate as far north as Puget Sound. With the
Pliocene began a period of elevation of the continental margin
which reached its climax during the glacial period of the Pleisto-
cene, followed by a subsidence to somewhat less than its present
elevation. These movements and attendant climatic changes
affected profoundly the existent flora and fauna which by this
time had assumed a distinctly modern type.* There seems to be
evidence that in western North America the increase in elevation
varied at different times from 1500 feet along the coast: to 3000-
6000 feet in the Sierra and Cascade Ranges and in the interior
basin.*

An abundant precipitation and a corresponding increase of
snowfall resulted in an extensive glaciation which reached south-
ward along the Cascade Range into southern Oregon and the
Sierra Nevada of California and was also present in the Rocky
Mountains and the Wasatch Range of Utah.® 7 s This, the so-

1Schuchert, C. Paleogeography of North America. Bull. Geol. Soc. Am. 20:
427-606. 1910.

2 — ERP co events in the geologic history of California. Science N. S.

3 Salisbury, а D. Physical geography of the Pleistocene with reference to the
correlation of Pleistocene formations. In Willis & Salisbury, Outlines of Geologic
History. 306 pp. .

4 Osborn, Н. Е. TM age of mammals. 635 pp. 1910.

ë Salisbury, R. О. 1. c.

* Smith, J.P. l.c.
dg ет І. С. Geological history of Lake Lahontan. U. S. Geol. Surv. Monogr.

з Gilbert, ok K. Lake Bonneville. U. S. Geol. Surv. Monogr.1:1-438. 1890.

1925]

EPLING—MONOGRAPH OF MONARDELLA

рәшшпәуәр JON рәшшдәәр YON зовшвАпгу вәтәпәр
зо *ju*g-reuro[eq|-uo үвпотуполә јо ләўпәг) */,
suuin[oo 9AOq* eu ut pojvorput sy] АПз }вәлвәп

jo [9s1edsrp jo ләиәг) '9

peurunojop oN ^o
рәшішләзәр 40N "4

рәшгшләјәр 40N ә
peuruyp ON 9

peurunojop JON *o
рәшшшәзәр 40N 'q
ourprvu

poururojop JON "o

333148 Ч рәўәш}вәл 4889] ‘ә
Joža oArjonpoudoed ‘q

335849 ЗИ 4} ^9 это во) "M 'S "w|-Ioqp uwg-[pLiqer) ueg 'ejesoy vjuvg-reulo[|eq ‘в ОЯТ 9AT)€39S9A "в
10у
|-889900n8 JSOU SPUN JSO ‘Q
(my
(nsojptano sogna `W) (0ројоәоит) ИГ) |-ирләрштэ оурирлорш w ‘pəyıpow 4583]
рәшшләјәр JON uordel Avg [911qvr) uvg { uorzo1 vovureÁn?) yun jo [esrodsip јо 193u97) *р
[esiedstp jo sonuoA€
[911qer) ueg рәшшләјәр YON puqen ueg рәшгшләәр о [821418098 јо 51204 'e
(орд рурирлтш * ИГ pue
(nonojod fiy (»jv1d2202n9] UDH уют *p)
ошр ип | pue 102422 W) [д pue 21807600 'W) возит Ап)
799 989-9493) ueg BIUIOJTIVD ‘М "S uordo1 Avg ево €jueg-1ieuro[eq SHUN әѕләлтр JSON, 'Z
ourpreu ourp vovureÁn^)
-og uvg-[oniqer) изя €IUIOJITO/) "A, ‘S| -wung ueg-[oriqur) ueg вво eju'eg-reuro[eqsjrun јо Ioquinu 2897994») т
soroodsqns : и вәтәәйѕ yun вәәйв :4tu() seroedsqns pun
ЛІ uonoog III uonoog II uonoog І uonoog 901493115)

NOLLAHDLLSIG AO H3LLNÀO V GNIWHOLUXG OL VIHSLLIMO ЯО NOLLYOIIddY АЯ аямтухно SLIASTU DNIMOHS INVHO Y

I S'IGVIL

[Уот. 12
30 ANNALS OF THE MISSOURI BOTANICAL GARDEN

called Ice Age, was of long duration and is thought to have been
divided into several warmer inter-glacial epochs. At the begin-
ning of the Pleistocene the flora and fauna of this region were
essentially of a temperate type and included many surviving
forms. At the period of maximum glaciation, however, the biota
was reduced both in number and in kinds.! If the remains pro-
duced by the Conard Fissure are correctly placed in point of time,
they would indieate that the climate of Arkansas at that time was
much the same as British Columbia at present, with an analagous
fauna. The larch extended as far south as Georgia. The
climate of California at that time has been compared to the
present climate of the Olympie Peninsula, cool and rainy, and
productive of heavy forests. To the east of the ice-covered
Sierra and Cascade ranges lay a vast upland correspondingly
elevated, with a higher rainfall than at present, and characterized
by the great inland lakes Lahontan and Bonneville which were
fed from the rivers and glaciers of the Sierra Nevada and Cascade
Ranges and Wasatch Range.’ The musk-ox ranged as far south
as Salt Lake City, indicating in the north a tundra-like region.*
There is, then, fairly certain evidence of the repeated displace-
ments of the existing biota during the Pleistocene and a consequent
actual southward migration of both plants and animals together
with the reappearance of the survivors as the climate ameliorated.
As at present constituted Monardella is an inhabitant either of
semi-arid or arid situations in the transition zone, or of the chap-
arral. From a study of the apparent evolutionary tendencies it
has been suggested that the genus represents a group of plants of
mesophytic origin which has become more and more adapted to
such a habitat. It does not seem probable that the genus main-
tained its present range during the Pleistocene. It is obvious
that most of its present range east of the great valley of Cali-
fornia would have been uninhabitable by reason of the boreal

‘Osborn, Н. Е. The age of mammals, р. 500. 1910.
* Osborn, Н. Е. І. c. pp. 487-488.

.

1925]
EPLING—MONOGRAPH OF MONARDELLA 31

or sub-boreal nature of the stations. While the genus may have
existed at lower levels on the great plateau, the facts of present
distribution suggest rather that it has entered this area since the
glacial period. Nor does it seem probable that the present range
in California was maintained, in view of the fact that the genus
is at present rarely found even in the outskirts of the northwestern
hygrophytic forests which presumably extended much further
south during the Pleistocene. One is led consequently to the
conclusion that Monardella or its precursors migrated southward
at this time. Such a conclusion is supported by other facts of
present distribution.

There is evidence to suggest that, at the time of maximum ele-
vation during the Pleistocene, the Channel Islands off the coast
of southern California (? Lower California also) were continuous
with the present mainland and evidently supported a similar
biota. With the subsequent depression the connecting valleys
were flooded and eventually only the highest points which con-
stitute the present islands were left uncovered. The present
flora of these islands,! while essentially that of the present main-
land, nevertheless contains at least one genus no longer found
upon the mainland and numerous endemic species as well, sug-
gesting very strongly the preservation of certain types through
isolation.

Monardella has been reported from these islands twice, namely,
from Santa Catalina Island and from Cedros Island. M. lanceo-
lata was reported from Santa Catalina Island by Lyon. The
author has seen no material supporting this report nor was
Millspaugh? able to verify it after examination of Lyon's collec-
tion. If seen by Lyon it is not improbable that the plant was a
recent adventive. The only species definitely known to grow on
any of these islands is M. thymifolia Greene, endemic on Cedros

1 Watson, S. On the flora of Guadalupe Island, Lower California, Proc. Am.
Acad. 11: 105-121. 1876; Lyon, У. Г. The flora of our southwestern archipelago.
Bot. Gaz. 11: 197-205, 330-336. 1886; Greene, E. L. Botanical excursion to the
island of San Miguel. Pittonia 1: 74-93. 1887; Botany of Cedros Island 7
194-208. 1888; Supplementary list of Cedros Island vci Ibid. 266-269. 1889;
Vegetation of San Benito Islands. Ibid. 261-266. 188

2 Millspaugh, C. F., and Nuttall, L. W. Flora of ons Catalina Island. Field
Mus. Nat. Hist. Publ. 212. 1923.

[Vor. 12
32 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Island. Since M. thymifolia is most nearly allied to M. villosa,
yet separated from it by perhaps 750 miles, occurring as it does
at the southernmost extreme of the generic range, it may be in-
ferred that the common stock from which both have originated
had a much more southerly range than M. villosa now occupies.
Furthermore, the fact that M. macrantha subsp. eumacrantha
occurs in an isolated montane community as far south as San
Pedro Martir in Lower California suggests a former wider and
more southerly range for that species. М. linoides presents a
similar case.

From the inferences drawn in the preceding paragraphs we may
arrive at the following explanation of the present distribution of
the genus. The precursors of Monardella were pushed southward
during the glacial period of the Pleistocene. The surviving rep-
resentatives again migrated northward when conditions of
climate were ameliorated. The prototypes of each of the sections
previously discussed became established approximately in the
present centers of dispersal of each group and were variably
successful in maintaining themselves and increasing their range.
With increasing aridity of the climate among other factors, and
due in part to isolation, the present species have arisen, and of
these the most widespread and thus, perhaps, the most successful,
is M. odoratissima.

MONARDELLA Benth.

Monardella Benth. Lab. Gen. & Sp. 331. 1834; in DC. Prodr.
12: 190. 1848; Benth. & Hook. f. Gen. Pl. 2: 1185. 1876;
Gray, Proc. Am. Acad. 11: 100. 1876; Bot. Calif. 1: 593. 1876;
Syn. Fl. №. Am., ed. 2, 21; 356. 1886; Brig. in Engl. & Prantl,
Nat. Pflanzenf. IV. Abt. 3a, 309. 1896; Howell, Fl. Northwest
Am. 549. 1901; Nelson in Coulter & Nelson, Man. Cent.
Rocky Mts. 430. 1909; Jepson, Fl. West. Middle Calif., ed. 2,
363. 1911; Abrams, Muhlenbergia 8:26. 1912; Fl. Los Angeles,
ed. 2, 317. 1917; Davidson & Moxley, Fl. South. Calif. 312. 1923.

Madronella Greene, Leaflets Bot. Obs. 1: 168, 1906; Piper,
Contr. U. S. Nat. Herb. 9: 493. 1906; Frye & Rigg, Elem. Fl.
Northwest, 195. 1914; Piper & Beattie, Fl. Southeast. Wash-
ington, 216. 1914; FI. "анте Coast, 309. 1915; Rydberg,
Fl. Rocky Mts., ed. 2, 750. 1923.

1925]
EPLING—MONOGRAPH OF MONARDELLA 33

Annual or perennial herbs, of fragrant odor, with small entire
or serrate leaves, flowers borne in terminal, globose, bracteate
glomerules. Calyx tubular, narrow, 10-15-nerved, 5-dentate;
teeth triangular, subequal, erect; throat naked. Corolla small,
usually rose-purple, sub-bilabiate, the upper lip two-lobed, the
lower lip three-lobed, the lips subequal, plane, the lobes linear.
oblong. Stamens four, all fertile, the anterior pair exceeding the
posterior, or subequal, erect, distinct, not greatly exserted.
Anthers bilocular, the locules oval, subparallel to divaricate,
subconfluent above but distinct. Style shortly and unequally
bifid at the summit. Ovary four-parted, the nutlets oblong-oval,
smooth, brown at maturity.

The type species is Monardella odoratissima Benth.

KEY то THE SUBGENERA!

A. Limb of corolla 15-1/5 the length of the tube; calyces 10-25 mm. long

B. Limb of corolla 12-24 the length of the tube; бубе 5-10 mm. long.. PYCNANTHAE

Subgenus 1. МАСКАМТНАЕ Briq. in Engl & Prantl. Nat.
Pflanzenfam. IV. Abt. 3a, 309. 1896 (adapted from Gray, Proc.
Am. Acad. 11: 100. 1876); Abrams, Muhlenbergia 8: 26. 1912.

Stem slender, rhizomatous, the branches decumbent or as-
cending; glomerules loosely flowered, usually with less than 20
flowers, bracts oblong; calyx 10-30 mm. long, slender, 13-nerved;

1 The abbreviations used herein are as follows:
H—Baker Herbarium of Pomona College.
CAS—California Academy of Science.
CSM—Colorado State Museum, Denver.
DH—Dudley Herbarium of Leland Stanford ay:
FM—Field Museum of Natural History, Chicag
GH—Gray Herbarium of Harvard University.
J—Herbarium of W. L. Jepson.
KH—Kew Herbarium
MBG—Herbarium of the Missouri Botanical Garden, St. Louis.
NYS—Herbarium of the New York State um.
OAC—Oregon Agricultural College Herbarium, Corvallis.
RMH—Rocky Mountain Herbarium, University of Wyoming.
S—Herbarium of H. St. John.
UC—Herbarium of the University of California.
US—U. S. National Herbarium at Washington.

(Voc. 12
34 ANNALS OF THE MISSOURI BOTANICAL GARDEN

corolla red, yellowish or pallid, the limb !/2 to 1/5 the length of
the tube.'
$ Section I—MACRANTHAE
KEY TO THE SPECIES
A. Corolla 20-45 mm. long; leaves generally pubescent........ 1. M. macrantha
B. Corolla 15-18 mm. long; leaves glabrate on both surfaces...... 2. M. Palmeri

1. M. macrantha Gray, Syn. Fl. М. Am. ed. 2, 2!: 459 (suppl.).
1886. Hall, Univ. Calif. Publ. Bot. 1: 110. 1902.

Perennial from slender rhizomatous stems, the branches de-
cumbent or ascending, 10-30 cm. long, seldom branching, pu-
bescent with short recurved trichomes, or villous, purplish;
leaves subcoriaceous, variable on the same plant, the blades
.5-3 em. long, ovate to lanceolate, subeuneate at the base and
generally broadest about one-quarter their length from the base,
very obtuse, entire or obscurely crenate-serrate, glabrous to
villous or cinereous on petioles .5-1.5 em. long; glomerules 2-4
cm. broad, bracts oblong-elliptical, approximately equal to the
calyces, acute, membranous, purplish or whitish, sparsely villous,
ciliate; calyx variable in size on the same plant, 1.2-2.5 ст. long,

purplish or green, sparsely villous, teeth acute, slender, villous
within; corolla scarlet to yellowish or pallid, puberulent, the tube
greatly exserted, the limb 5-11 mm. long, the upper lip the
longer, the lobes coalesced more than half the length of the lip,
those of the lower lip nearly free; the anther 1-1.5 mm. wide, the
sacs widely divergent, the connective wider than the length of
the sac, the margin retuse.

KEY TO тнв SUBSPECIES
Corolla for the most part 35-45 mm. long; calyx 20-25 mm.
long

а, БИБ: AAA

! All measurements of flower parts herein given are based upon flowers which
were fresh and in full bloom when pressed and which were boiled in water prior to
study. It should be observed that flower parts which have withered naturally
Pres pressing never regain their full size but remain much shrunken. If the

ected uy p a drop of thin
br e, i i 3.4 J BM! +1 n н 2-4 11 2
ü dry. On ad addition of a drop of warm "water it may again be examined.

——

1925]
EPLING—MONOGRAPH OF MONARDELLA 35

a. Subsp. eumacrantha, nom. nov.
... Monardella macrantha Gray, Proc. Am. Acad. 11: 100. 1876;

Bot. Calif. 1: 593. 1876, 2: 476. 1880; Syn. Fl. М. Am., ed. 2,
2': 356. 1886; Abrams, Muhlenbergia 8: 28. 1912; Davidson
and Moxley, Fl. South. Calif. 313. 1923.

Madronella macrantha Greene, Leaflets Bot. Obs. 1: 169. 1906.

Blades of the leaves 1-3 cm. long, glabrate above and pubescent
beneath, rarely pubescent on both surfaces (villous in the variety) ;
glomerules 3-4 cm. broad, bracts seldom equaling the calyces,
purple; calyx in general 20-25 mm. long; corolla scarlet or yellow-
ish, in general 30-45 mm. long, the anthers 1.25-1.5 mm. wide.

Specimens examined:

CALIFORNIA: Big Sur, May-June, 1901, Davy 7436 (UC);
Santa Lucia Mts., June 10, 1909, Brandegee (UC); Cuyamaca
Mts., July 12, 1875, E. Palmer 295 (GH, Tyre; MBG); Cuya-
maca Mts., Sept. 1882, Orcutt 485 (GH); San Diego Co., Orcutt
(MBG); Santa Lucia Mts., 1885, Brandegee (GH); Tassajara Hot
Springs, Monterey Co., June 1901, Elmer 3228 (MBG; US);
Laguna Mts., June 28, 1919, Hastwood 9227 (GH); Smith Mt.,
San Diego Co., July 25, 1882, Orcutt (MBG); between Cuyamaca
and Julian, June 21, 1903, Abrams 3812 (US; MBG; GH) Julian
City July 15, 1875, Cleveland (GH) ; Cuyamaca Peak, 5000 ft., June
30, 1897, Reed (BH); Cuyamaca Lake, dry stony slopes, 4700 ft.,
June 27, 1923, Munz & Harwood 7241 (BH); Laguna Mts.,
July, 1889, Orcutt (US); Santa Lucia Mts., 1880,G. R. Vasey 487
(US); Cuyamaca Mts., 1875, E. Palmer 294 (US); Julian, Cleve-
land Nat. Forest, July 29-30, 1915, Hitchcock (US); Pine Hills,
July 29, 1915, Collins & Kempton 268 (US); West Fork Trail
near Sturdevant’s, San Gabriel Mts., 4250 ft., July 13, 1918,
Peirson 183 (J).

Lower CALIFORNIA: San Pedro Martir, Aug. 1903, Robertson
33 (UC, leaves small for the plant and pubescent on both sides) ;
Oallecitos, San Pedro Martir, 8000 ft., July 15, 1905, Goldman
1228 (US).

Var. Hallii Abrams, Mublenbergia 8: 29. 1912; Davidson
and Moxley, Fl. South. Calif. 313. 1923.

M. macrantha var. tenuiflora Hall, Univ. Calif. Publ. Bot. 1:
110. pl. 11. 1902 (neither of Watson nor Gray).

[Vor. 12
36 ANNALS OF THE MISSOURI BOTANICAL GARDEN

M. macrantha var. longiloba Abrams, Muhlenbergia 8: 29.
1912; Davidson and Moxley, Fl. South. Calif. 313. 1923.

Branches and leaves villous, the latter in general 2 cm. long or
more and tending to be ovate rather than lanceolate, frequently
subtruncate at the base and very obtuse; corolla frequently
yellowish, the limb being sometimes as long as 10-11 mm., the
lobes being correspondingly slender and very acute.

Specimens examined:

CALIFORNIA: Smith Mt., San Diego Co., July 19, 1890, Г. J.
Gray (MBG); chaparral belt of south side, canyon of the San
Jacinto R., 4300 ft., July 4, 1898, Hall 976 (UC, турв of var.
longiloba Abrams; US; fragment GH); San Jacinto Mts., 4000
ft., July 16, 1897, Hall 687 (UC); chaparral belt, San Jacinto
River, 4400 ft., June 19, 1897, Hall 669 (UC); Mill Creek, San
Bernardino Co., July 8, 1898, Parish 4578 (US); San Bernardino
Co., 1876, Parry & Lemmon 328 (GH; MBG); Palomar, May,
1901, Hall 1936 (MBG; US; type collection, TYPE in Dudley Herb.) ;
City Creek Road, San Bernardino Mts., 5000 ft., July 17, 1921,
Johnston 2858 (BH).

b. Subsp. nana (Gray), comb. nov.

Monardella nana Gray, Proc. Am. Acad. 11: 101. 1876; Bot.
Calif. 1:593. 1876; Syn. Fl. №. Am., ed. 2, 21:356. 1886; Abrams,
Muhlenbergia 8: 30. 1912; Davidson & Moxley, Fl. South.
Calif. 313. 1923.

M. macrantha var. nana Gray, Syn. Fl. N. Am. ed. 2, 21; 459
(suppl.). 1886; Hall, Univ. Calif. Publ. Bot. 1: 111. 1902.

M. villosa var. leptosiphon Torrey, Bot. Mex. Bound. 129.
1859 (not Gray, Bot. Calif. 1: 593. 1876; Syn. Fl. N. Am., ed.
2, 2:5 357. 1886).

М. nana var. leptosiphon Abrams, Mublenbergia 8: 31. 1912;
Davidson and Moxley, Fl. South. Calif. 313. 1923.

Madronella nana Greene, Leaflets Bot. Obs. 1: 169. 1906.

Blades of the leaves in general .5-1.5 ст. long, less commonly
2 cm. long, ovate rather than lanceolate with a tendency to be-
come truncate at the base, glabrous on the upper surface and
pubescent beneath or sparsely villous or cinereous throughout;
glomerules 2-3.5 em. broad, bracts usually somewhat longer than

1925]
EPLING—MONOGRAPH OF MONARDELLA 37

the calyces and often whitish; calyx in general 12-16 mm. long,
corolla pinkish or pallid, in general 20-35 mm. long, the anthers
] mm. wide.

Specimens examined:

CALIFORNIA: Cuyamaca Mts., 4500 ft., May 29, 1899, Hall 1202
(UC); mountains in back of San Diego, 1875, Cleveland (GH;
TYPE); San Felipe, 1881, Cleveland 770 (GH); between Cuyamaca
and Oriflamme Canyon, San Diego Co., June 28, 1903, Abrams
3941 (GH; MBG; BH; US); Laguna Mts., San Diego Co., June
28, 1919, Eastwood 9215 (US; GH); Laguna Mts., San Diego Co.,
July, 1889, Orcutt (MBG; US); Laguna Mts., 5700 ft., June 2,
1920, Spencer 1560 (BH); Laguna, June 14, 1894, Schoenefeld
8544 (US); Julian, 1880, Vasey (US); Smith Mt., San Diego Co.,
July, 1882, Orcutt (GH); Julian, San Diego Co., June 13, 1894,
Brandegee (UC) ; Smith Mt., July 15, 1890, Orcutt 2110 (US).

Lower CALIFORNIA: mountains of Lower California (near
Japa), July 5, 1884, Orcutt (FM).

Var. tenuiflora Gray, Syn. Fl. №. Am., ed. 2, 2:: 459. 1886
(suppl.); not of Hall, Univ. Calif. Publ. Bot. 1: 110. pl. 12. 1902.

Monardella tenuiflora Watson in Gray, Proc. Am. Acad. 17:
230. 1882.

M. nana var. tenuiflora Abrams, Muhlenbergia 8: 32. 1912;
Davidson & Moxley, Fl. South. Calif. 313. 1923.

M. macrantha var. pinetorum Hall, Univ. Calif. Publ. Bot. 1:
110. pl. 12. 1902.

Branches 10-15 mm. tall, short-pubescent; leaves cinereous-
pubescent, the trichomes usually appressed but frequently some-
what villous, blades averaging 1 cm., the petioles 4-5 mm.;
corolla pale yellow to almost white; the tube averaging 20 mm.,
barely tapering, the limb 6-8 mm. long, the lobes acute, narrower
at the base than towards the middle, somewhat exceeding the
stamens.

Specimens examined:

CALIFORNIA: Santa Rosa Mts., frequent in partly shady rocky
slopes in the chaparral belt, 5700 ft., June 26, 1922, Munz 5833
(BH); between Tahquitz and Round Valley, San Jacinto Mts.,
7500 ft., July 6, 1922, Munz 6033 (BH); San Jacinto Mts., 6000—

[Vor. 12
38 ANNALS OF THE MISSOURI BOTANICAL GARDEN

8000 ft., July 17, 1897, Hall 691 (UC; fragment in GH); San
Jacinto Mts., Aug. 1881, S. B. & W. F. Parish 327 (UC); San
Jacinto Mts., July, 1880, S. B. & W. Е. Parish 327 (GH, түре;
MBG; US); San Jacinto Mts., 8500 ft., July 2, 1895, A. W.
Anthony (UC); Fuller's Mills Mts., San Jacinto Range, 6500 ft.,
July, 1901, Hall 2559 (UC, this and the following approach subsp.
eumacrantha var. Hallii in the structure of the corolla but retain
the small anther; in size of corolla they are very similar to Elmer's
collection of M. macrantha at Tassajara); pine-clad slopes of
west side (San Jacinto Canyon), 6000 ft., June 24, 1901, Hall
2258 (UC); Tahquitz Valley, San Jacinto Mts., July 22, 1897,
Hall 725 (UC, 25479, TYPE of M. macrantha var. pinetorum Hall;
US); San Jacinto Mt., halfway between forks of Summit Trail
and Log Cabin on upper trail, 8000 ft., no date, Jepson 2322 (J);
Palm Canyon and return to Van Deventer's, May 17-June 1, 1901,
Jepson & Hall 1336 (J).

Lower CALIFORNIA: Japa, 9000 ft., July 5, 1884, Orcutt (GH).

Var. arida Hall, Univ. Calif. Publ. Bot. 1: 111, pl. 10. 1902.

M. nana var. arida Abrams, Muhlenbergia 8: 33. 1912; David-
son & Moxley, Fl. South. Calif. 313. 1923.

Low, branches less than 10 em. long, cinereous; leaves cinereous,
the blades 0.5-1.0 cm. long, the petioles often longer, averaging
about three-fourths the length of the blade; corolla pale yellow
to almost white; the tube less than 18 mm. long, scarcely tapering,
less than a millimeter wide, the limb 4-6 mm. long, the lobes oblong
rather than tapering, often obtuse, exceeding the stamens slightly.

Specimens examined: |

CALIFORNIA : desert region to the southeast of San Jacinto Mt.,
along Coyote Creek at 5000 ft., June, 1901, Hall 2127 (UC 25475;
MBG; US); Coyote Canyon, Santa Rosa Mt., 5000 ft., May,
1899, Hall 1180 (UC; RMH; US); between Vandeventer's and
Palm Canyon, San Jacinto Mts., 4000 ft., May, 1901, Hall 1852
(UC); Old Nicholas Canyon, Santa Rosa Mts., common in
clearings in chaparral, 5000 ft., July 1, 1922, Munz 5936 (BH).

M. macrantha was based originally upon the collections of both
Cleveland and Palmer, the first collected at “Julian City,” the
second in the “‘Cuyamaca Mts." In a note at the end of the

1925]
EPLING—MONOGRAPH OF MONARDELLA 39

original description Gray interchanged the order of the collectors’
names and the localities from whence the collections were made
and wrote “Southern part of California in the Cuyamaca Moun-
tains and near Julian City, D. Cleveland, E. Palmer." Both
eollections at the Gray Herbarium are mounted upon the same
sheet and are similar although the former is scant. It is evident
that the original description is drawn mainly from the Palmer
collection and since, of the two, it only is labeled in Gray's hand
'* Monardella macrantha n. sp." and is more copiously represented,
it is here considered the historical type.

In the same paper in which M. macrantha was published,
M. nana was described and united with it to form a generic
section. M. nana was based upon a collection by Cleveland in
the same year and same locality. The two species thus consti-
tuted were published again in the ‘Botany of California’ and in
the ‘Synoptical Flora.’ In the supplement to the second edition
of the latter, however, M. nana was made a variety of M. ma-
crantha, together with M. tenuiflora Wats. published in 1882.
This interpretation was accepted by Hall, who studied the plants
in the field and described two varieties as new, referring certain
plants to M. macrantha var. tenuiflora (Wats.) Gray. One of
the newly described varieties was M. macrantha var. pinetorum.
A comparison of the plants referred to var. tenuiflora with the
type specimen indicated to Abrams that this variety had been
misinterpreted by Hall and that M. macrantha var. pinetorum
Hall was in fact M. macrantha var. tenuiflora Gray. Abrams
therefore applied the name M. macrantha var. longiloba to the
plants referred by Hall to M. macrantha var. tenuiflora, and named
the plants referred by Hall to M. macrantha (not the variety)
var. Hallii, since they did differ from the type of M. macrantha
in the villosity of the foliage.

The chief character upon which var. longiloba was based lay in
the unusually long (10 mm. or more) lobes of the corolla which
were correspondingly slender and acute. The chief character
upon which var. Hallii was based lay in the abundant villosity
of the foliage. The present author agrees with Abrams that
M. macrantha var. tenuiflora Hall is not synonymous with M.
macrantha var. tenuiflora Gray. A careful study of the group,

[У ог. 12
40 ANNALS OF THE MISSOURI BOTANICAL GARDEN

however, has not demonstrated that var. longiloba and var.
Hallii may be segregated by any constant group of characters.

M. macrantha varies tremendously, and yet, as stated by Hall,
all gradations between the extremes may be found nor do any
very constant lines of cleavage appear to exist. The com-
binations of characters which appear are very puzzling. The
clearest hiatus appears between those groups herein described
as subspecies. Subsp. eumacrantha, on the one hand, is made up
of plants which are more or less lax in their habit, trailing, with
leaves which are comparatively large, being 2-3 cm. long, and
with flowers unusually large for the genus, being in general 30-45
mm. long and scarlet or yellowish. Subsp. nana, on the other
hand, has a more compact growth form, particularly in the plants
of more arid habitat, with smaller leaves and flowers which are
pinkish or yellowish-white and in size from 20 to 35 mm. long.
The pubescence of this group is more often dense and close,
giving a cinereous aspect to the plant.

In the first subspecies, while the range of variation is consider-
able, there does not seem to be any very definite occurrence of
concomitant characters save that, in general, plants with the
larger ovate villous leaves also have the elongated lobes of the
corolla. Such plants have accordingly been referred to var.
Hallii, and since the occurrence of the elongate lobes is more or
less connected with the villous leaves, var. longiloba is considered
synonymous with it. All gradations exist.

In the second subspecies two variants from the more typical
plant may be more clearly discerned and appear to have a dis-
tribution which is characteristic. The subspecies proper has
foliage which in general resembles that of subsp. eumacrantha and
like it varies from pubescent to villous, but is in general smaller,
ranging from 1 to 2 em. in length. The pubescence, however, is
closer and more cinereous. The corolla is smaller, and presents
one or two qualitative differences as well, namely, in the shape
of the tube, which is evenly tubular and less funnel-form, and in
the shape of the lobes, which are more slender and usually slightly
narrower at the base and longer in proportion to the length of
the tube. The anthers are distinctly smaller, averaging 1 mm.
in width. "These differences were noted by Abrams and used as

1925] í
EPLING— MONOGRAPH OF MONARDELLA 41

a means to segregate M. nana. Nevertheless, they are not con-
stant, the form described by Torrey as M. villosa var. lepto-
siphon clearly connecting the two subspecies.

The variety tenuiflora (var. pinetorum Hall) has smaller leaves
(1 em.), which are distinctly cinereous on both surfaces with a
corolla much exserted, the limb being 6-8 mm. long, the lobes very
acute. "The petioles of the leaves are somewhat less than half
the length of the blade. The variety arida has similar foliage
but smaller, the blades being .5-1 cm. long and the petioles often
three-fourths the length of the blade. The growth form is very
compact. The corolla is much less exserted, the limb being 4-6
mm. long, the lobes oblong-linear and rather blunt. All grada-
tions occur.

It is impossible from present knowledge of the group to judge
to what extent these forms represent racial differences and to
what extent they represent forms induced by the environment.
To determine this will require careful field study and experiment.

The present author has not seen the type of M. villosa var.
leptosiphon which was based upon a collection by Parry at “San
Felipe," but from the description there can be little doubt as to
the plant referred to, particularly so from the description of the
corolla and stamens. It was confused by Gray with a subspecies
of M. villosa common to central California, an error which was
perpetuated until attention was directed to it by Abrams, and in
the Gray Herbarium there is apparently no authentic material
of Torrey’s plant.

2. M. Palmeri Gray, Proc. Amer. Acad. 12: 82. 1877; Bot.
Calif. 2: 476. 1880; Syn. Fl. N. Am., ed. 2, 21: 357. 1886.

Madronella Palmeri Greene, Leaflets Bot. Obs. 1: 169. 1906.

Perennial from a slender, rhizomatous stem, the branches low,
ascending, 10-15 cm. long, purplish, scarcely puberulent; leaves
subcoriaceous, 1-2 em. long, oblong or lanceolate-oblong, entire or
nearly so, obtuse, glabrous, the midvein hardly perceptible,
tapering into a very short petiole or sessile; glomerules large in
proportion, 2.5-3 cm. broad, purplish, bracts exceeding the cal-
yces, oblong, the innermost linear-oblong, obtuse, thin and mem-
branous, reddish purple, puberulent; calyx 9-11 mm. long, red-

[V or. 12
42 ANNALS OF THE MISSOURI BOTANICAL GARDEN

dish purple, 13-veined, rather definitely bilabiate, the teeth
slender, 1.5-2 mm. long, hirsute within; corolla 15-17 mm. long,
the tube slender, twice the length of the limb, retrorsely hirsute
in the throat, the limb 5-6 mm. long, the lips about equal, the
lobes of the upper lips coalesced about one-half its length, those
of the lower lip approximately free, tapering slightly; stamens
about equal, the anthers less than 1 mm. wide, the anther-saes
divaricate, the angle about 90°, the margin of the connective
apparently entire, or if retuse, very slightly; nutlets 2.5 mm. long.
pecimens examined:

CALIFORNIA: Santa Lucia Mts., under the redwoods, Aug. 2,
1876, E. Palmer 859 (GH, түре; MBG; US); no locality given, .
Brandegee (GH); Santa Lucia Mts., July, 1880, Vasey 498 (US;
FM)

Subgenus 2. РүсхАхтнАЕ Briq. in Engler & Prantl, Nat.
Pflanzenf. IV. Abt. 3a, 309. 1906 (adapted from Gray, Proc.
Am. Acad. 11: 101. 1876); Abrams, Muhlenbergia 8: 26. 1912.

Stems various, decumbent or erect but not rhizomatous;
glomerules compactly flowered, usually with thirty or more
flowers; bracts variable, tending to ovate or lanceolate; calyx
5-10 mm. long, 10-15-nerved; corolla rose-purple or pallid, the
limb one-half to two-thirds the length of the tube.

KEY то THE SPECIES

A. Annuals; stem herbaceous and not suffrutescent.
a. Bracts fenestrate, i.e., the intravenous tissue like
И 18. M. Douglasii
b. Bracts membranous or scarious, but not fenestrate.
a Margin of the calyx teeth white-scarious or
terminating in a white, recurved cusp.
1. Teeth ending in a white recurved cusp. ...19. М. leucocephala
2. Teeth blunt, margin white-scarious.
I. Lower margin of connective entire;
calyz Бен 1:3. ees. 18. М. exilis
II. Lower margin of connective dis-
tinctly notched; calyx 13-nerved..... 17. М. candicans
0. Margin of the calyx teeth bounded by a distinct
vein, not scarious.
1. Leaves plane, lanceolate or oblong.
І. Bracts puberulent.
*Bracts acuminate, secondary veins
absent or not prominent........ 15. M. Breweri

1925]
EPLING—MONOGRAPH OF MONARDELLA 43

**Bracts acute, secondary veins green,

net-like, prominent. ..... р 14. M.lanceolata
П. Braets villous throughout. .......... 16. M. Pringlei
2. Leaves undulate or crisped, oblanceolate-
о АЕГОН E ae AS EE 12. М. undulata

B. Perennials; stem woody at the base and suffrutescent.
a. Leaves oblanceolate-oblong, undulate or erisped. 12. M. undulata
b. Leaves ovate to linear, plane.
a. Bracts reflexed, foliar in texture and shape. 4. M. villosa
B. Bracts erect, sheathing, firm or membranous,
but only the € pair foliar (see also M.
villosa subsp. neglecta).
l. Leaves їй, a strongly developed felt-like
tomentum on the under surface, glabrous
or lanate above.
I. Leaves glabrous above.............. 5. M. hypoleuca
II. Leaves lanate above............... 6. M.lanata
2. Leaves pubescent or glabrous, often paler
beneath, but never felt-like.
I. Leaves .5-.8 mm. long, more or less
eretate do ntate.
esp ensis pubescent, cinereous,
а 3. М. thymifolia
2 alah гк hoary with а
bluish COs ТУСЕ ober 9. M.cinerea
II. Leaves 1-several em. long, mostly

entire.

"Foliage and stem silvery white,
with a dense microscopic puber-
won leaves typically linear-

Sy CO Taare eT T] 11. М. linoides

“Foliage and stem glauco
some but not silvery white; leaves
typically lanceolate.

TBracts firm, neither mem-
branous nor chaffy; lobes of
blunt.

corolla
1Leaves v RM above, 2-5
wm, EE. ас... 8. М. зале а
езе ака on both
surfaces, seldom 2 сш.
AME TCU шык: и. 7. M. viridis

[Vor. 12
44 ANNALS OF THE MISSOURI BOTANICAL GARDEN

$ Section П. VILLOSAE

3. M. thymifolia Greene, Bull. Calif. Acad. Sci. 1: 211. 1886;
Gray, Syn. Fl. N. Am., ed. 2, 2': 459. 1886.

Madronella thymifolia Greene, Leaflets Bot. Obs. 1: 169. 1906.

Perennial, shrubby, 12-20 cm. tall, much branched, the lower
branches woody, covered with a grayish brown checking bark,
spreading, the upper branches erect, slender, pubescent; leaves
5-8 mm. long, ovate triangular, very obtuse, margin recurved,
entire or obscurely serrate, soft-pubescent, petioles broad, 1-2
mm. long; glomerules a centimeter or less in diameter, bracts ovate,
acute, equaling the calyces, herbaceous (Greene), becoming chaffy .
and brittle, pinnately veined, thinly pubescent, the margin sub-
ciliate; calyx 6-7 mm. long, thinly pubescent; corolla purplish
(Greene), 12-13 mm. long, the tube about 8 mm. long, the lips
subequal; the lobes of the upper lip coalesced about two-thirds
its length, those of the lower lip about one-fourth its length,
tapering but little, obtuse, anther-sacs divergent, the connective
equilateral, well developed, nutlets oblong-oval, 1.5 mm. long.

Specimens examined:

Lower CALIFORNIA: Cedros Island, 1859, Veatch (UC; GH);
Cedros Island, July-Oct. 1896, А. W. Anthony 143 (US; GH);
Cedros Island, March-June, 1897, Anthony 316 (GH; MBG;
US); Cedros Island, March 22, 1911, Rose 16162 (US).

The few specimens which are available for study are uniform
in appearance, but are for the most part past flower, the glomer-
ules being dry and chaffy and straw-colored. The bracts are
stated by Greene to be herbaceous. The only specimen examined
in which this point might be determined (US 313872) showed the
outer pair erect and subfoliaceous only, the inner being mem-
branous but firm, the veins being well developed. All were
tinged with rose and were noticeably pubescent and glandular.

4. M. villosa Benth. Bot. Voy. Sulph. 42. pl. 21. 1844; in
DC. Prodr. 12: 190. 1848; Gray, Proc. Am. Acad. 11: 101.
1876; Bot. Calif. 1: 593. 1876, excl. variety; Syn. Fl. N. Am.,
ed. 2, 2!: 357. 1886, excl. variety; Jepson, Fl. West. Middle
Calif., ed. 2, 364. 1911.

Perennial from decumbent stems, woody at the base, the bark

1925]
EPLING— MONOGRAPH OF MONARDELLA 45

brown in old stems, often checking; the branches erect, simple,
or often with short, usually sterile secondary branchlets: of
variable stature, 10-60 cm. tall, variably pubescent near ihe in-
florescence, gisbrete below; leaves exceedingly variable, in
general ovate, varying, on one hand, to rotund and, on the other,
to lanceolate, obtuse, obscurely crenate-serrate, or entire, or coarsely
dentate-serraté, usually of a firm texture, villous or villous-
tomentose to glabrate, blades 1-3 cm. long, rounded at the base,
but abruptly tapering into a petiole 0.5—1.0 em. long; glomerules
compact, 2-4 em. broad, bracts leaf-like in shape and texture, only
thei , about equal to the calyces,
reflexed, villous or tomentose, conforming to the pubescence of
the plant; calyx 7-10 mm. long, 13-nerved, scarious below, green
and shaggy-villous above, teeth ovate-triangular, villous without
and on the margins; corolla 10-18 mm. long, rose-purple to pallid,
the lobes linear-oblong and ribbon-like, blunt, those of the upper
lip coalesced usually about half its length, those of the lower lip
nearly or wholly free; stamens about the length of the lobes, the
filaments retrorsely hispidulous near the base, the anther sacs
divergent, the connective equilateral, well developed.

The following classification represents an attempt to char-
acterize what appear to be the principal tendencies of evolution
within M. villosa. In each subdivision of its range may be found
a form which is characteristic of that locality, and while forms
geographically distant may be quite diverse all intermediate
forms may be found and, taken as a whole, show a close corre-
lation between form and geographical position. This is true to
the extent that, after familiarity with the species and its distri-
bution as a whole has been attained, it is possible to place any
given plant within a comparatively short distance of its habitat
merely by inspection. There are not hard-and-fast lines between
these areas of distribution just as there may not be found any
constant morphological differences between the forms that in-
habit them. Yet taken in their typical aspect, as represented
by the modal points in the curve of their variability, it will be
found that each subspecies occupies a fairly distinct territory,
certainly of a degree of distinctness to be very suggestive. The
subsp. euvillosa, characterized by leaves which are rather large,

[V or. 12
46 ANNALS OF THE MISSOURI BOTANICAL GARDEN .

thinnish, ovate, sharply serrate and villous, is found along the
coast from the Santa Lucia range northward to Mendocino
County, the variety fransiscana being evidently a form of the
littoral. The next and most nearly allied subspecies with which
it intergrades imperceptibly is subsp. subserrata. This sub-
species is characterized by a usually smaller lanceolate leaf which
is typically lanceolate and is shallowly serrate. The pubescence
is villous but much finer, being frequently subtomentose. Subsp.
subserrata is found in the Salinas Valley and ranges northward
through the coastal valleys to the Bay region and Napa County
and northward to Mendocino County. It occurs occasionally
in the Sierra Nevada, and in southern Oregon but at low eleva-
tions.

Subsp. Sheltoni, characterized by a lanceolate, even narrower
leaf, which is nearly or quite entire and is glabrate, or puberulent
at most, is found chiefly in the Sierra Nevada at elevations as
high as 7300 feet and in the Siskiyou Mountains of northern
California and southern Oregon. Transitional forms occur on the
valley side of the coast range in Central California.

Subsp. neglecta is a very doubtful form which is herein treated
as a subspecies for convenience and for the purpose of directing
attention toward it. It occurs only in the vicinity of Mt.
Tamalpais.

KEY TO THE SUBSPECIES

A. Herbage variously pubescent.
1. Leaves ovate to roundish, villous and green rather
than canescent ог cinereous..................-.- subsp. euvillosa
2. mind spores rarely ovate, canescent or

box NEA. +a ba pow ane de Ek N . subserrata
B. иес айк ы at most.
1. Bracts leaf-like, not markedly ciliate............. subsp. Sheltoni
2. Bracts membranous, purple, ciliate. .............. subsp. neglecta

a. Subsp. euvillosa, nom. nov.

Monardella globosa Greene, Pittonia 5: 32. 1902.

M. involucrata Heller, Muhlenbergia 1: 35. 1904.
Madronella villosa Greene, Leaflets Bot. Obs. 1: 168. 1906.
Madronella globosa Greene, Leaflets Bot. Obs. 1: 169.. 1906.
Madronella involucrata Heller, Muhlenbergia 1: 136. 1906.

1925]
EPLING—MONOGRAPH OF MONARDELLA 47

Branches villous in the upper nodes, leaves ovate, commonly ser-
rate, the teeth of variable prominence, 1-3 cm. long, for the most
part 2-2.5 em., tending to become truncate at the base, rather
than cuneate; corolla 12-15 mm. long, the lobes of the upper lip
somewhat less than half its length, those of the lower lip some-
what less than its full length.

Specimens examined:

CALIFORNIA: Berkeley, July 4, 1880, Engelmann (MBG);
Woodside, San Mateo Co., June 9, 1919, Wattlier (GH; US);
mountains near Santa Cruz in shaded woods, 186-, Bolander
(GH); “Potrero, near San Francisco, July 30, 1868,” Kellogg &
Harford 775 (GH; MBG); Goat Island (San Francisco Bay),
June 8, 1882, Greene (FM); foothills west of Los Gatos, June 11,
1904, Heller 7497 (RMH; MBG; OAC; GH; US$; foliage varies
exceedingly on different sheets of this collection) ; San Mateo Co.,
Aug. 11-12, 1897, Congdon (GH); Santa Cruz, June 16, 1903,
Thompson (MBG); no locality, Coulter 541 (GH; cited by Ben-
tham); Pine Forest at Pacific Grove, Aug. 16, 1905, Coleman
(DH); Madrone Springs canyon (? Sonoma Co.), 2000 ft., Aug. 14,
1917, Abrams (DH); California Redwood Park, Santa Cruz Co.,
June 18, 1919, Shockley (DH); Crystal Springs, June 30, 1903,
Baker 3353 (GH; MBG); Bay-View Hills, June 13, 1912, East-
wood 349 (US; GH; MBG); Saratoga, Big Basin Road, June 23,
1915, Abrams 5273 (DH); Angel Island near San Francisco,
Vasey (СН); Borax Lake, 1865, Torrey 403a (GH); New Almaden,
1865, Torrey (GH); Mare Island, June 9, 1874, Greene 226 (GH);
Contra Costa, July, 1903, Elmer 4658 (MBG; US); Niles, Mor-
rison Canyon, June 20, 1897, Jepson 59h (J); Weldon Canyon,
Vaca Mts., Solano Co., June 1, 1891, Jepson 501 (J); Glen Echo,
Santa Cruz Co., June 17, 1896, Jepson 59g (J); Ft. Bragg, 1914,
Mathews 169 (J; typical); no locality, Hartweg 1913 (GH);
Monterey, Haenke (GH); near San Francisco, 1866, Kellogg (US) ;
sandy loam, Pajaro Hills, Monterey Co., June-July, 1899, Chandler
366 (US); San Leandro, Contra Costa Co., June 21, 1915, East-
wood 4743 (US); Santa Cruz Mts., July 9, 1913, Hitchcock 221
(US); N. Berkeley Hills, June 30, 1917, Walker 605 (US); Santa
Clara Co., June 1, 1895, Dudley 4200 (US); Santa Cruz Mts.,
July 22, 1882, Pringle (US).

[Vor. 12
48 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Var. franciscana (Elmer), comb. nov.

Monardella franciscana Elmer, Bot. Gaz. 41: 320. 1906.

Madronella franciscana Heller, Muhlenbergia 2: 244. 1906.

Branches villous to tomentose in the upper parts; leaves ovate to
rotund, thickish, obscurely serrate to entire, truncate at the base,
even subeordate, the blades commonly about 2 cm. long, villous
above, tomentose and canescent beneath; corolla 12-18 mm. long,
the lobes tapering somewhat, appearing as though foreshortened,
those of the upper lip one-third to one-half its length, those of
the lower lip somewhat less than its length, the throat more
ample than in subsp. euvillosa.

Specimens examined:

CALIFORNIA: Santa Lucia Mts., 1885, T. S. Brandegee (GH);
San Juan, back of Monterey, June 14, 1861, Brewer 713 (GH ; US);
no locality given, 1853, Gibbons (GH); Pine Mt., near San Simeon
Bay, San Luis Obispo Co., July 22, 1876, E. Palmer 361 (US;
MBG); San Mateo, July, 1903, Elmer 4766 (MBG; UC; US;
type collection of M. franciscana Elmer, түре in DH); Santa
Lucia Mts., Monterey Co., June, 1898, Plaskett (GH; US); San
Bruno Hills, near Ocean View, San Francisco Co., June 8, 1906,
Heller 8371 (MBG; GH; US); no locality given, 1876, E. Palmer
360 (US; suggests M. hypoleuca).

b. Subsp. subserrata (Greene), comb. nov.

Monardella subserrata Greene, Pittonia 5: 81. 1902.

M. tomentosa Eastwood, Bull. Torr. Bot. Club 30: 496. 1903.

M. villosa Howell, Fl. Northwest Am. 549. 1901.

M. villosa var. leptosiphon Gray, Syn. Fl. М. Am., ed. 2, 2:
357. 1886 (not Torrey).

M. mollis Heller, Muhlenbergia 1: 35. 1904.

Madronella mollis Heller, Muhlenbergia 1: 138. 1906.

Madronella gigantea Heller, in herb. (Heller 12395). 1916.

Madronella subserrata Greene, Leaflets Bot. Obs. 1: 169. 1906.

Stems villous or at least pubescent in the upper nodes; leaves
lanceolate, rarely ovate, commonly 2-2.5 ст. long, entire or shal-
lowly serrate, those in the upper nodes canescent from a villous-
like tomentum, varying to a short pubescence, more dense on the
lower surface, the trichomes soft and fine; corolla 15-18 mm. long,
the lobes ribbon-like, scarcely tapering, blunt, those of the upper

1925]
EPLING—MONOGRAPH OF MONARDELLA 49

lip one-half its length or more, those of the lower lip usually
equal to it.

Specimens examined:

CALIFORNIA: Arroyo del Puerto, Stanislaus Co., June 11, 1862,
Brewer 1253 (GH; US); Laytonville, Aug. 2, 1902, Eastwood
(CAS; ТУРЕ of M. tomentosa Eastwood); dry hills, Carmel R.,
Monterey Co., June 22, 1921, S. B. Parish 20043 (GH); near
Ladoga, Lake Co., June 8, 1919, Heller 13241 (MBG; GH; US);
Jolon, Monterey Co., Sept. 22, 1894, Eastwood (GH ; an unusual
form); Tuolumne City, 2800 ft., July 19, 1911, Abrams 4719
(DH; approaches the preceding in the degree of pubescence);
Ione, 200-500 ft., June, 1904, Braunton 1046 (MBG; US);
gravelly stream bank west of Proberta, Tehama Co., June 19,
1916, Heller 12395 (OAC; MBG; GH; US; type collection of
M. gigantea Heller) ; Sonoma Co., Samuels 162 (US) ; Mt. Diablo,
near Lake, Contra Costa Co., June 30, 1916, Abrams 5707 (DH);
road between Petrified Forest and Mark West, Sonoma Co.,
July 4, 1916, Abrams 5790 (DH); between Knight's Valley and
Mark West Springs, June 28, 1902, Heller 5791 (MBG; GH; US);
3 mi. west of Leesville, Colusa Co., June 6, 1916, Heller 12355
(GH; US; OAC; MBG); near Arnold's on Outlet Creek, Mendo-
cino Co., July 9, 1916, Abrams 5925 (DH); Jolon, Monterey Co.,
1880, Vasey 492 (US); Valley of Arroyo Seco, Monterey Co.,
May 30, 1861, Brewer 678 (GH; US); Searsville Ridge, Santz Cruz
Mts., June 2, 1914, Abrams 1700 (US); New Idria, San Benito
Co., July 24, 1861, Brewer 798 (US); Ukiah, Mendocino Co.,
June 20-July 3, 1898, Chesnut 377 (US); Round Valley (east of
Mt. Diablo?), July 25, Aug. 3, 1897, Chesnut 542 (US); moun-
tains of the upper Sacramento, 1845-7, Fremont’s 3rd Expedition
(US, 48119; MBG, 114315; GH); Grass Valley, Amador Co.,
July 10, 1894, Hansen 439 (US); Big Horse Mt., S. Fork Eel RA
July-Aug., 1892, Jepson 30p (J); grade to Howell Mt., Napa
В. basin, June 26, 1893, Jepson 50 т (J); Calistoga, June 3, 1923,
Jepson 9969 (J; M. villosa var. tomentosa Jepson); Hemlock,
July 17, 1897, Jepson (J.).

OrEGon: Roseburg, Oct. 2, 1881, Pringle (MBG); Grant’s
Pass, July 24, 1915, Canby 103 (OAC); 4 mi. north of Agnes,
June 25, 1917, Nelson 1502 (GH); by the river, Grant’s Pass,
July 2, 1887, T. Howell, in part (US, 43117).

[Vor. 12
50 ANNALS OF THE MISSOURI BOTANICAL GARDEN

M. subserrata Greene is based upon a specimen collected by
G. W. Dunn in Sonoma County, June, 1890. Only a photograph
of this plant has been seen by the author.

c. Subsp. Sheltoni (Torrey), comb. nov.

Monardella Sheltoni Torrey in Durand, Pl. Pratten., Jour.
Acad. Nat. Sci. Phila. II. 3: 99. 1855.

M. villosa var. glabella Gray in Bot. Calif. 1: 593. 1876; ee
Fl. N. Am., ed. 2, 2:: 357. 1886 (in part).

M. reflexa Howell, Fl. Northwest Am. 549. 1901.

M. dentata Rydb. Bull. Torr. Bot. Club 31: 641. 1904.

M. coriacea Heller, Muhlenbergia 1: 35. 1904.

Madronella coriacea Heller, Muhlenbergia 1:138. 1906.

Madronella dentata Rydb. Bull. Torr. Bot. Club 33: 150. 1906.

Madronella amabilis Heller, in herb. (12567). 1916.

Stems puberulent or glabrous in the upper nodes; leaves ovate,
tending to lanceolate, puberulent or nearly glabrous, blades com-
monly 2-2.5 cm. long, subcuneate at the base, entire or obscurely
serrate, tapering into a petiole 2-5 mm. long; bracts lanceolate,
reflexed, short-pubescent; corolla 12-20 mm. long, purple to
whitish, the lobes of the upper lip about half its length, those of
the lower its full length.

Specimens examined:

CALIFORNIA: Klamath R., Humboldt Co., 1500 ft., June, 1901,
Chandler 1522 (GH; MBG; US); Castella, Shasta Co., July 24,
1912, Eastwood 1369 (GH; US); Nevada Co., June 20-22, 1912,
Eastwood 581 (GH; MBG; US; type locality); Sierras, 1872, A.
Gray (GH); no locality stated, Bridges 307 (US; GH); no data
other than this: ‘‘M. Sheltoni Torr. in herb.," in Dr. Gray’s
hand; the author’s conception of M. Sheltini is based upon this
plant together with the description and other plants from the
type locality; Indian Valley, Plumas Co., Aug., 1896, Austin
(MBG); Summit, between Mad and Trinity Rivers, on Eureka
Red Bluff Road, July 22, 1916, Abrams 6181 (DH); between the
McCloud and Sacramento: Rivers, Shasta Co., June 24, 1916,
Heller 12447 (MBG; GH; US; OAC; distributed as a form of M.
gigantea Heller; varies considerably within the collection);
Shasta River hills near Klamath River, June 27, 1909, Butler 938
(MBG); Hornbrook, Siskiyou Co., July 6, 1903, Copeland 3497

1925]
EPLING—MONOGRAPH OF MONARDELLA 51

(GH; US; MBG); Miller's Ranch, summit between Gilroy and
Watsonville, May, 1903, A. D. E. Elmer 4647 (US; MBG; OAC);
Tassajara Hot Springs, June, 1901, Elmer 3224 (MBG; US);
Loma Prieta, Santa Clara Co., July 22, 1893, Dudley (DH);
slope above Round Meadow, Fresno Co., 7300 ft., July 25, 1914,
Smiley 588 (GH); Sisson, Aug. 20, 1889, Sheldon (MBG; NYS);
Grass Valley, 3000 ft., July, 1892, Hansen 439 (MBG); Green-
ville, July 12, 1907, Heller & Kennedy 8824 (US; GH; MBG);
Willow Creek, Humboldt Co., June 16, 1918, Abrams 7183
(DH); near Nevada City, July 14, 1905, Heller 8113 (GH; US;
type locality of M. Sheltoni); west side of Trinity River near
Willow Creek, Humboldt Co., 600 ft., July 9, 1911, Tracy 3476
(GH; US); Plumas Co., May, 1894, Ames (GH); Little Chico
canyon, May, 1896, Austin 803 (MBG; US); Russian River at
Healdsburg, July 8, 1902, Heller 5812 (MBG; GH; type collection
of M. coriacea Heller); Napa, 1899, Smyth (GH; approaches the
large-leaved forms of subsp. euvillosa, the same with the two
following); Ukiah, April 22, 1891, Fritchey (MBG); Calaveras Big
Tree Road, Aug. 1890, Jepson 40 c (J); Huntington Lake,
Fresno Co., 7000 ft., Grant 1157 (J); Vichey Springs, June 22,
1891, Fritchey (MBG); Summit, Butte Co., June 28, 1897, Austin
1126 (US); Clio, Plumas Co., Aug. 27, 1910, Eggleston 6213 (US);
Little Chico Creek, 2000 ft., July 5, 1900, Leiberg 5024 (US).
Овевом: Grant’s Pass, June 24, 1884, T. Howell 244 (GH; US);
by the river, Grant's Pass, July 2, 1887, T. Howell, in part
(OAC, 8981; MBG, 114312); gravel bar along Chetco River,
7 mi. above Harbor, Curry Co., July 19, 1919, Peck 8910 (GH;
MBG); by the river, Grant's Pass, July 2, 1887, T. Howell (MBG;
NYS; OAC); Snow Camp, Curry Co., 4000—4250 ft., July, 1916,
Thompson 7 (DH); Cherry Creek Flat, Klamath Co., Aug. 15,
1908, Rose 265 (DH); Grants Pass, June 20, 1886, Henderson 805
(OAC); Big Butte Creek Crossing, 30 mi. east of Medford,
Jackson Co., Aug. 27, 1916. Heller 12567 (MBG; GH; US; OAC;
type collection of M. amabilis Heller); Sykes Creek, Jackson Co.,
July 14, 1892, Hammond 325 (MBG); Takilma, Josephine Co.,
June 25, 1918, Peck 7981 (GH); Wimer, Jackson Co., July 14,
1892, Hammond 325 (US); Waldo, dry open forest of Pinus
ponderosa, 1 mi. west of town, July 18, 1924, Hall 11990 (UC).

[Vor. 12
52 ANNALS OF THE MISSOURI BOTANICAL GARDEN

M. dentata Rydb., based upon an unnumbered sheet in the -
Herbarium of the New York Botanical Garden, is unquestion-
ably the plant which grows in the lower altitudes of the Sierra
Nevada in central California. That it occurs on Gray’s Peak
in Colorado as reputed is very doubtful, and until further material
is forthcoming such an extension of the range should not be made.
It should be noted in this connection that Dr. Torrey visited the
central Sierras several years earlier and that it is not impossible
that an exchange of labels has taken place. "The present printed
label of the type sheet reads: “Herbarium of Columbia College,
New York. Plants collected on Gray's Peak, Colorado Territory
in August and September, 1872, by J. Torrey" and below in an
unidentified hand ‘‘Monardella odoratissima Bth." The label
is not contemporary with the collection. The plant thus labeled
is strikingly like the collection of M. villosa subsp. Sheltoni made
by Bridges in California (US; GH).

d. Subsp. neglecta (Greene), comb. nov.

Monardella neglecta Greene, Pittonia 5: 82. 1902.

Madronella neglecta Greene, Leaflets Bot. Obs. 1: 169. 1906.

Stems puberulent to glabrous in the inflorescence, purple;
leaves ovate or oblong, obtuse, the blades 1-1.5 cm. long, sub-
cuneate at the base, serrate in some at least, tapering into a
petiole 2-5 mm. long; bracts ovate, acute, membranous, only the
outer foliaceous, and reflexed, the innermost purple, pubescent to
glabrous, ciliate, about equal to the calyces; calyx 6-8 mm. long,
pubescent with spreading trichomes; corolla 12-14 mm. long,
rose-purple, the lobes of the upper hio about one-half its length,
that of the lower free nearly to the base.

Specimens examined:

CALIFORNIA: Mt. Tamalpais, 1876, Vasey (GH; US, TYPE of
M. neglecta Greene); Tiburon, Marin Co., June 9, 1912, East-
wood 315 (GH; US); Mt. Tamalpais, July 29, 1912, Eastwood 1617
(GH; MBG; US); Crystal Springs Lake, June 23, 1913, Suksdorf
312 (GH); south side of Mt. Tamalpais, July 17, 1913, Suksdorf
581 (GH); Rock Spring Trail, Tamalpais, July 2, 1905, K. Bran-
degee (UC); Tiburon, Marin Co., on a rocky slope facing San
Francisco Bay, June 3, 1909, Walker 1727 (GH; MBG; US).

The specimen collected by Vasey on Mt. Tamalpais is cited

1925]
EPLING—MONOGRAPH OF MONARDELLA 53

with the description of M. neglecta Greene. In the Greene
Herbarium a fragment of this plant is mounted on the same sheet
with a specimen collected by G. W. Dunn in Marin Co., July 22,
1890. The latter collection is designated as the type in Greene's
handwriting. Only a photograph of this sheet has been seen by
the author.

Subsp. neglecta is exceedingly variable, in habit, in pubescence,
and in the texture of the bracts. When most vigorous it is
scarcely to be distinguished from subsp. Sheltoni. А villous
form, scarcely separable from subsp. euvillosa presumably col-
lected with or near average plants, has been observed.

5. M. hypoleuca Gray, Bot. Calif. 2: 476. 1880; Syn. Fl. N.
Am., ed. 2, 21: 356. 1886; Abrams, Muhlenbergia 8: 39. 1912;
Davidson & Moxley, Fl. South. Calif. 314. 1923.

Monardella robusta Elmer, Bot. Gaz. 39: 46. 1905.

Madronella hypoleuca Greene, Leaflets Bot. Obs. 1: 169. 1906.

Perennial, suffrutescent, 30-50 cm. tall, erect or trailing,
“prostrate or supported by other growth” (Elmer), the older
stems glabrous, light brown, the bark checking longitudinally,
the younger branches simple, purplish, villous near the glomerule;
leaves rhomboidal-lanceolate, the smaller oblong, 2-4 cm. long,
entire, obtuse, green and glabrous above, the veins impressed,
covered beneath with a white felt-like tomentum, the margin revolute,
petioles 3-10 mm. long; glomerules 3-4 cm. broad, compact, the
bracts ovate, shorter than the calyces, membranous rather than
foliar, but firm, tomentose; calyx 6-7 mm. long, scarious below,
green and villous above, veins 13-15, teeth ovate-triangular,
acute; corolla pale lavender or white, 15-16 mm. long, the tube
10 mm. long, the upper lip equal to or somewhat longer than the
lower, the lobes about half its length, those of the lower lip one-
fourth its length, tapering but little, blunt; stamens well exserted,
the anthers divergent, the connective equilateral, well developed.

Specimens examined:

CALIFORNIA: mountain drive, Santa Barbara, Aug. 21, 1904,
Abrams 4149 (GH); dry ridges, Santa Monica Mts., Topanga
Canyon, June, 1907, Hasse; Rattlesnake Canyon, Santa Bar-
bara, Aug. 1902, Elmer 3728 (US; MBG; type collection of M. ro-

[Vor. 12
54 ANNALS OF THE MISSOURI BOTANICAL GARDEN

busta Elmer, TYPE in DH); San Bernardino Co. (Gray, Syn. FL),
1876, Parry & Lemmon 330 (GH; түре); San Juan Capistrano,
July, 1882, Nevin 688 (GH); Malibu Canyon, Los Angeles Co.,
Aug. 5, 1898, Barber (UC); Rincon Cr., Ventura Co., Sept. 19,
1922, Baer (BH); trail, Trabuco canyon to Santiago Peak, Santa
Ana Mts., Orange Co., 3000-4000 ft., Sept. 7, 1923, Munz 7742
(BH).

6. M. lanata Abrams, Muhlenbergia 8: 39. 1912; Davidson
& Moxley, Fl. South. Calif. 313. 1923.

Perennial, suffrutescent, 30—50 em. tall, erect, branching at the
base, the branches purplish, villous to lanate; leaves oblong, the
smaller even oblanceolate or subspatulate, 2—4 cm. long, very obtuse,
short-pubescent to lanate above, the veins not prominent, covered
beneath with a dense white felt-like tomentum, the margin strongly
revolute, petioles 3-10 mm. long; glomerules 3-4 em. broad, com-
pact, the bracts ovate, shorter than the calyces, membranous
rather than foliar, but firm, tomentose; calyx 6-7 mm. long,
scarious below, green above, lanately villous, veins 13-15, teeth
ovate, triangular, acute; corolla pale lavender or white, 15-17
mm. long, the tube 10 mm. long, the upper lip equal to or some-
what longer than the lower, the lobes coalesced about half its
length, those of the lower lip one-fourth its length, tapering but
little, blunt; stamens well exserted, the anthers divergent, the
connective equilateral, well developed.

Specimens examined:

CALIFORNIA: Descanso Grade near the top, July 19, 1906,
К. Brandegee (UC, 104626, түрк); Potrero Mts., July 23, 1883,
Orcutt 996 (GH; UC); San Diego Co., near Арш; July 10,
1912, Abrams 4896 (UC; US; BH; DH); Palomar Mt., back
of Pala, July, 1901, Schellenger (UC; upper surface of leaves
glabrate).

M. lanata is apparently distinct from M. hypoleuca which it
most nearly resembles. The specimen collected by Schellenger
suggests strongly a continuous range of variation between the
two. If such is found to be the case when M. lanata is better
— it would best be considered as a subspecies of M. hypo-

ca.

1925]
EPLING—MONOGRAPH OF MONARDELLA 55

7. M. viridis Jepson, Fl. West. Middle Calif., ed. 1, 465. 1901,
and ed. 2, 364. 1911.

M. ledifolia Greene, Pittonia 5:81. 1902.

Madronella ledifolia Greene, Leaflets Bot. Obs. 1: 169. 1906.

Madronella viridis Arthur, Torreya 21: 12. 1921.

Perennial from a short woody stem, branching at the base,
erect, or decumbent, the sterile branches numerous, short, the
fertile branches slender, 6-12 inches tall, whitish puberulent;
leaves rhomboidal lanceolate, 1-2 cm. long, obtuse, canescent with a
minute but dense pubescence, dark green above, white beneath, sub-
entire, the margin distinctly and narrowly revolute, subcuneate
at the base and narrowed to a margined petiole 2-3 mm. long;
glomerules 2 cm. broad or less, globose, the bracts about equal to
the calyces, membranous or subfoliar, firm, ovate-lanceolate,
acute, softly pubescent; calyx 7 mm. long, rather shaggy-pu-
bescent or villous; corolla 14-16 mm. long, the upper lip some-
what shorter, the lobes about half its length, those of the lower
lip nearly free, all tapering evenly, but blunt; stamens subequal,
about equal to the lips, the anthers small, .75 mm., the sacs di-
vergent, the connective equilateral.

Specimens examined:

CALIFORNIA: Sonoma County, ? 1894, Krauss (DH); Upper
Conn Valley, Napa Range, Oct. 13, 1894, Jepson 70d (J, TYPE;
fragment UC; турв of M. ledifolia Greene in Greene НетЪ.);
Mt. St. Helena, 1893, Jepson 32f (J); Mt. Hanna, Lake Co.,
July 15, 1897, Jepson 100j (J).

M. ledifolia Greene is based upon the type collection of M.
viridis. The species is distinct but apparently restricted to the
St. Helena range.

8. M. saxicola Johnston, Bull. South. Calif. Acad. Sci. 28: 19.
1921.

Perennial from decumbent, suffrutescent stems, branches erect,
30-40 em. tall, branching below if at all, purple, puberulent;
leaves rhomboidal-lanceolate, the smaller oblong, 2-5 cm. long,
obtuse, green and glabrous above, pubescent beneath with a
silvery microscopic tomentum, the margin revolute, the veins im-
pressed, the blade narrowed at the base to a petiole 2-7 mm.

[Vor. 12
56 ANNALS OF THE MISSOURI BOTANICAL GARDEN

long; glomerules 2.5-3 em. broad, bracts ovate, acute or obtuse,
about equal to the calyx, membranous, but firm and rather thick-
ish, pubescent; calyx 8-10 mm. long, rather slender, short-pu-
bescent to villous, veins 10-11, teeth ovate-triangular, acute,
hirsute within; corolla lavender to rose-purple, 14-17 mm. long,
the tube 10 mm., the lobes ribbon-like but tapering slightly,
blunt, those of the upper lip less than half its length, those of the
lower lip three-fourths its length or more; stamens well exserted,
the anther-sacs divergent, the connective equilateral, well de-
veloped.

Specimens examined:

CALIFORNIA: at start of old trail to the flats, near Brown’s
Flats, San Antonio Mts., 5000 ft., Sept. 1, 1918, Johnston 2133
(BH, түре; DH); south spur of Cucamonga Peak, common among
rocks on ridge crest, 5200 ft., June 30, 1918, Johnston 2050 (DH;
BH); head of Evey Canyon on Sunset Trail, San Antonio Mts.,
4750 ft., July 1, 1917, Johnston 1440 (BH).

Apparently distinct but very similar to M. viridis Jepson, from
which it differs chiefly in leaf character, the leaves being larger,
coarser, glabrous above, with a very minute, dense silvery to-
mentum on the lower surface. They are less cuneate and more
rounded at the base and the veins are noticeably impressed.
Since the author has seen no intergrading forms and since M.
saxicola is geographically quite distant from M. viridis, it has
seemed preferable to retain this as a species until it may be more
extensively studied.

9. M. cinerea Abrams, Muhlenbergia 8: 33. 1912; Fl. Los
Angeles, ed. 2, 317. 1917; Davidson & Moxley, Fl. South. Calif.
313. 1923. :

Perennial, woody at the base, the stem covered with a brown
bark which flakes and falls away; branches several, spreading,
9-15 em. long, cinereous; leaves ovate-lanceolate, 5-11 mm. long,
obtuse or acute, denticulate rather than serrate, sometimes entire,
cinereous or hoary on both surfaces with a soft subvillous pubescence
giving the whole plant a bluish cast, sessile; glomerules 1.5-2 cm.
broad, compact, purplish, bracts ovate to elliptical, acute or
shortly acuminate, about the length of the calyces, membranous,

1925]
EPLING—MONOGRAPH OF MONARDELLA 57

pinnately veined, sparingly villous, purplish; calyx 7-9 mm. long,
villous, 12-15-nerved, the teeth 2 mm. long, slender, acute,
villous within; corolla rose-purple, 13-14 mm. long, lobes lance-
olate, those of the upper lip two-thirds its length, those of the
lower lip the length of the lip, the lower stamens slightly exceed-
ing the lobes, a third longer than the upper pair, the anthers di-
vergent, the connective equilateral, well developed.

Specimens examined:

CALIFORNIA: Little Baldy, 9500 ft., (common among the
rocks,” Aug. 22, 1917, Johnston 1693 (GH; UC; BH); south
slope of Baldy, 6000 ft., dry rocky ground, July 4, 1917, Johnston
1422 (DH; GH; UC; BH); Icehouse Canyon, rocky places, 8300
ft., July 31, 1917, Johnston 1458 (BH; UC); N. Fork San Antonio,
8000 ft., July 28, 1917, Johnston 1571 (UC; BH); Mt. Baldy
Lookout, 7000 ft., June 20, 1917, Johnston 1261 (UC); Mt.
Baldy, 8750 ft., July 4, 1917, Johnston 1449 (UC); Mt. Baldy,
10,000 ft., July 4, 1917, Johnston 1420 (UC); Mt. San Antonio
(Mt. Baldy), 9000 ft. or more, July 24, 1901, Abrams 1928 (BH;
type collection, түре in DH); saddle between Baldy and Little
Baldy, dry rocky slopes, 9400 ft., July 21, 1922, Munz 6114 (BH);
Mt. Baldy, 7500 ft., June 8, 1918, Peirson 182 (J).

A very puzzling species of restricted habitat. The degree of
variability within its altitudinal distribution is considerable, and
while some forms suggest M. linoides subsp. stricta and others
M. odoratissima subsp. australis, it still appears to be distinct.

$ Section ПІ. OponaATISSIMAE

10. M. odoratissima Benth. Lab. Gen. et Sp. 332. 1834, in
DC. Prodr. 12: 190. 1848; Gray, Proc. Am. Acad. 11: 101.
1876; Bot. Calif. 1: 594. 1876; Syn. FI. №. Am., ed. 2, 2!: 357.
1886.

Perennial from a woody, often contorted and decumbent stem,
the bark dark brown, splitting and falling away; branches erect
or ascending, unbranched, seldom with short, erect, sterile branch-
lets in the upper nodes, of variable stature, 10-60 cm. tall, com-
monly 25-30 cm., pubescence close and short, canescent, or ciner-
eous or glaucous-like, but never silvery ; leaves lanceolate, varying,

[Vor. 12
58 ANNALS OF THE MISSOURI BOTANICAL GARDEN

on the one hand, to ovate and, on the other hand, to oblong,
1-3 em. long, entire, rarely obscurely serrate, pubescent or
glabrate, the pubescence close and short, never villous but canes-
cent, cinereous or glaucous-like, never silvery, sessile or tapering
behind to a petiole 1-3 mm. long (7 mm. in extreme forms);
glomerules 1-5 cm. broad; bracts membranous, ovate or rotund,
obtuse or acute, varibasy shaded with purple, densely short-
villous or tomentose, or puberulent, strongly ciliate on the margin
in, most; calyx 6-10 mm. long, 13-nerved, scarious below, green
and villous or hirsute above, teeth ovate, triangular, acute, hir-
sute within; corolla 1-2 em. long, rose-purple to pallid, the tube
retrorsely puberulent, the lobes lanceolate rather than linear-
oblong, rounded to a rather obtuse point, those of the upper lip
one-third to one-half its length, those of the lower lip seldom
equaling it, for the most part about three-quarters its length;
filaments retrorsely hispidulous at the base, the anther-sacs
divergent, the connective equilateral, well developed; nutlets
oval-oblong, about 2 mm.

Ккү TO THE SUBSPECIES

A. Leaves appearing nearly or quite glabrous, tending to
appear glaucous, usually more than 2 cm. long.
a. Bracts tending to ovate or rotund, pubescent......... subsp. euodoratissima
b. Bracts tending to elliptical or oblong, puberulent........... subsp. glauca
B. Leaves distinctly pubescent.
а. Leaves hoary, whiter beneath, pubescence dense in

typical specimens я eae aE а ene es subsp. discolor
b. Leaves cinereous or green, but not markedly whiter
beneath.
«. Bracts usually exceeding the calyces and short-
ССИ ЧА t cusa ae ds subsp. australis
8. Bracts equal to or shorter than the calyces,
acute or obtuse
1. Glomerule 5-1. 5 em. broad, seldom 2 em... . ....subsp. parvifolia
2. Glomerules 2-3 em. broad, seldom less.
I Calyces woolly; pubescence about the
same on both leaf surfaces.................. subsp. pallida
II. Calyces hirsute; pubescence on lower
leaf surface usually longer and soft. ...... .subsp. pinetorum

It was an unfortunate circumstance that the historical type
of M. odoratissima should have been collected at the extreme
northern station, not only for the species but for the genus and

1925]
EPLING—MONOGRAPH OF MONARDELLA 59

in a locality which is not readily accessible. By reason of the
great variability of the species, and because the described “type”
was an extreme form and but little known, the synonymy which
has grown up within the species is unusually large. If one were
to consider the historical type-sheets alone, he might construct
an artificial key which would satisfactorily separate not only
those sheets but not a few others which by themselves appear
"distinct." Yet when he came to apply such a key to extensive
herbarium material or to the plants in the field, it is the author's
opinion that the user of such a key would find himself driven to
the creation of a still larger number of new *'species" in order to
classify his material consistently.

By reason of the fact that the species covers a wide range of
territory, and since it does present differences in different parts
of the range, an earnest but unsuccessful effort was made to
find satisfactory criteria which would serve to divide the group
into two or more clean-cut divisions. The subspecies here de-
scribed represent the nearest approach to such an ideal. For
convenience they might be called species and treated as such, yet
it is the opinion of the author that no advantage would accrue,
so numerous are the connecting forms, so profound is the effect
of the environment, especially in numerous montane-desert sta-
tions where great extremes may occur within a relatively small
radius of map distance and so close are the relationships of
the subspecies.

a. Subsp. euodoratissima, nom. nov.

Monardella odoratissima Howell, Fl. Northwest Am. 550. 1901.

M. glabra Nutt., collected near Walla Walla, Mss., ex Benth.
DC. Prodr. 12: 190. 1848.

Madronella odoratissima Piper, Contr. U. 8. Nat. Herb. 9: 493.
1906; Piper & Beattie, Fl. Southeast. Washington, 216. 1914.

Branches thinly pubescent above, hardly cinereous; leaves
lanceolate, tapering at the base but subsessile, averaging 2 cm.,
green, appearing glabrous, but shortly and sparingly pubescent;
bracts ovate to rotund, about equal to the calyces, obtuse for
the most part, pubescent on the veins, ciliate, calyx woolly pu-
bescent around the teeth; corolla about 15 mm. long, the lobes
slender.

[Vor. 12
60 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Specimens examined:

WasHINGTON: near the narrows above Kettle Falls on the
Columbia, Douglas (TYPE at Kew, a portion and photograph at
MBG); Meyers Falls, Stevens Co., Aug. 22, 1902, Kreager 499
(US; GH; from the type locality and a very close match for the
type); Blue Mts., June 26, 1897, Horner 408 (US); Blue Mts.,
Columbia Co., July 26, 1897, Horner B408 (GH; the same as the
preceding ?); Blue Mts., Walla Walla Co., 5000 ft., Aug. 2, 1896,
Piper (GH); Blue Mts., Walla Walla Co., July 15, 1896, Piper
(BH); Colville Nat. Forest, 3500 ft., Aug. 6, 1912, Wright (US);
near Mt. Hood on the Columbia R., Aug. 4, 1920, Suksdorf
10568 (GH).

Onzcox: Clearwater, no date, Spaulding (GH); no locality or
date, Geyer 468 (GH; cited by Bentham); Pendleton, dry thicket,
July 16, 1915, Peck (GH); Blue Mts., between Ukiah and Long
Creek, 4600 ft., July 26, 1917, Lawrence 842 (DH; US); Gibbon
(Bingham Springs Station), 530 m., June 28, 1916, Eggleston
12845 (US); Milton, Umatilla Co., 1000 ft., Aug. 26, 1896, Brown
43 (RMH; MBG; US); Elgin, Aug. 15, 1899, Shear 6587 (US).

NEvADA: Monitor Range, 8000 ft., Sept.-Oct., 1878, Phillips
& Sargent (ОН); 7 mi. east of Ely, 2400-3000 m., Hitchcock 1306
(08); the two preceding are intermediate with subsp. glauca, but
are perhaps closer to this.

New Mexico: Mogollon Mts., on or near the west fork of the
Gila R., Socorro Co., 8500 ft., Aug. 23, 1903, Metcalfe 565 (US;
GH; MBG; can in no way be distinguished from the Blue Mt.
plants).

b. Subsp. discolor (Greene), comb. nov.

Monardella discolor Greene, Pittonia 2: 24. 1889 ; Howell,
Fl. Northwest Am. 550. 1901.

M. nervosa Greene, Pittonia 4: 322. 1901.

Madronella discolor Greene, Leaflets Bot. Obs. 1: 169. 1906;
Piper, Contr. U. S. Nat. Herb. 9: 493. 1906; Piper & Beattie,
Fl. Northwest Coast, 309. 1915.

Madronella nervosa Greene, Leaflets Bot. Obs. 1: 169. 1906 i
Piper, Contr. U. S. Nat. Herb. 11: 493. 1906.

Madronella odoratissima Greene, Leaflets Bot. Obs. 1: 169.
1906; Rydb. Fl. Rocky Mts., ed. 2, 751. 1922.

1925]
EPLING—MONOGRAPH OF MONARDELLA 61

Branches pubescent above, scurfy and cinereous, leaves ovate-
lanceolate, tending to ovate, rather abruptly tapering behind but
subsessile, averaging about 2 cm., often less, obscurely crenate-
serrate, hoary to cinereous with a dense minute tomentum; bracts
ovate or oblong, about equal to the calyces, obtuse, woolly-
pubescent in typical specimens, at least pubescent, the margins
strongly ciliate; calyx woolly-pubescent; corolla averaging about
13 mm., the lobes slender.

Specimens examined:

WASHINGTON: north fork of Cowiche Creek (Yakima region),
July 21, 1901, Cotton 464 (US); Alkali Lake, Douglas Co., 335
m., July 7, 1893, Sandburg and Leiberg 413 (US); no locality
stated, 1889, Vasey 466 (GH; US); Wenatchee, July 28, 1896,
Whited 195 (US); White Bluff Ferry, Upper Columbia River,
Aug. 11, 1892, Lake & Hull 705 (GH; MBG); Rock Island, July,
1893, 2000-3000 ft., Sandburg & Leiberg (US; MBG; RMH);
Yakima region, Cascade Mts., 1882, T. S. Brandegee 235 (US;
MBG; GH); Mt. Rainier, moraine of Cowlitz Glacier, in loose
rock, 5000 ft., Aug. 1895, Piper 2078 (US); Yakima, 1877, T. J.
Howell (GH); bluffs east side of Columbia, below the Chelan,
Oct. 13, 1880, Watson 328 (GH; a specimen on the same sheet
collected “оп the bluffs of the Columbia В. above the Chelan,
west side,” Oct. 12, 1880, by the same collector and bearing the
same number is very close to subsp. euodoratissima) ; stony bot-
toms of canyons, Rattlesnake Mts., 2500 ft., Aug. 2, 1902,
Cotton 760 (US; GH; MBG); along creek south of Ellensburg,
June 27, 1897, Whited 547 (US; OAC); gravelly banks of the
Yakima near Clealum, Aug. 13, 1889, Greene (FM; UC; type
collection of M. discolor Greene) ; gravelly shores of Yakima River,
June, 1897, Elmer 373 (US; MBG; annoted by Greene as being
“М. discolor Greene exactly and from near original station");
southeast side of Mt. Rainier, 7000 ft., Allen (GH); Ellensburg,
July, 1898, Elmer (MBG); Mt. Adams, 6000-7000 ft., Aug. 31,
1882, Suksdorf (FM); Umtanum Creek, Yakima Co., July 26,
1923, St. John 3108 (8); Coulee City, Grant Co., June 25, 1923,
St. John 3107 (S); 18 mi. north of Yakima, dry stony soil in belt of
Purshia and Artemisia tridentata, July 23, 1924, Hall 12009 (UC).

Orncon: Mt. Hood, among rocks, 6500 ft., Aug. 25, 1899,

[Vor. 12
62 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Barber (GH; RMH); The Dalles, June, 1881, T. Howell (OAC);
no locality, collected by (?) Cooper, Stevens Exp. (GH). The
following specimens from eastern Oregon are most nearly related
to M. odoratissima subsp. discolor but have the pubescence much
reduced, in this respect resembling subsp. glauca; the habit, shape
of the leaves and general aspect is that of subsp. discolor, however;
some suggest subsp. euodoratissima: Gilliam Co., near forks of
Cottonwood Canyon, 3400 ft., Sept. 7, 1894, J. B. Leiberg 885.
(US; GH; UC); near dry run between Bear Buttes and Button
Springs, Crook Co., 4400 ft., Aug. 24, 1894, J. B. Leiberg 797
(US; GH; FM; UC); White Horse Mts., Aug., 1901, D. Griffiths
& E. L. Morris 416 (US); near Guano Ranch, Harney Co., 4350
ft., July 24, 1896, F. V. Coville & J. B. Leiberg 9 (US).

c. Subsp. glauca (Greene), comb. nov.

Monardella glauca Greene, Pittonia 4: 321. Nov. 7, 1901.

M. modocensis Greene, Pittonia 4: 321. Nov. 7, 1901

M. purpurea Howell, Fl. Northwest Am. 550. Nov. 20, 1901.

M. rubella Greene, Pittonia 5: 84. 1902.

M. ovata Greene, Pittonia 5: 82. 1902.

M. ingrata Greene, Pittonia 5: 83. 1902.

Madronella modocensis Greene, Leaflets Bot. Obs. 1:169. 1906.

Madronella rubella Greene, Leaflets Bot. Obs. 1: 169. 1906.

Madronella glauca Greene, Leaflets Bot. Obs. 1: 169. 1906.

Madronella ovata Greene, Leaflets Bot. Obs. 1: 169. 1906.

Madronella oblongifolia Rydb. Bull. Torr. Bot. Club 36: 686.
1909; Fl. Rocky Mts., ed. 2, 751. 23.

Madronella sessilifolia Rydb. Bull. Torr. Bot. Club 36: 685.
1909; Fl. Rocky Mts., ed. 2, 751. 1923.

Madronella purpurea A. Nelson, Bot. Gaz. 52: 71. 1911.

Madronella ingrata Greene, Leaflets Bot. Obs. 1: 169. 1906.

Branches puberulent, usually glaucous-appearing, purple or
whitish; leaves ovate-lanceolate, elliptical or oblong, 1.5-4 em.
long, seldom truly ovate, variable on the same plant in size and
shape, narrowed at the base to a margined petiole 1-5 mm. long,
glabrate, commonly glaucous-appearing due to a microscopic
puberulence, the uppermost leaves commonly subsessile; bracts
ovate, with a tendency for the outer to become orbicular and
the inner oblong, puberulent, purplish, ciliate; calyx pubescent,

1925]
EPLING—MONOGRAPH OF MONARDELLA 63

hirsute around the teeth, seldom woolly; corolla 1-2 cm. long, the
lobes noticeably tapering.

Specimens examined:

OREGON: summit of Cascade Mts., Ashland-Klamath Falls
Rd., July, 1920, Peck 9265 (GH; MBG); coast mountains near
Waldo, June 13, 1884, T. J. Howell (US; GH; type locality of
M. purpurea Howell and not improbably the collection referred
to by him); Ashland, July 8, 1886, Henderson 805 (OAC) ; Cherry
Creek, 4150 ft., dry woods, July 23, 1899, Leiberg 4306 (US);
rocky peak 12 mi. southeast of Port Orford, July 25, 1919, Peck
8934 (MBG; GH); Siskiyou Mts., 12 mi. south of Waldo, July 2,
1918, Peck 8396 (GH); 8 mi. south of Waldo, June 14, 1904,
Piper 6234 (US); Eight Dollar Mt., Josephine Co., June 12, 1904,
Piper 5341 (US); Eastern Cascade Mts., Klamath Co., June 30,
1902, Cusick 2846 (GH; MBG; US); 2 mi. southeast of Oregon
Caves, Josephine Co., July 16, 1918, Peck 8369 (GH); Keno,
Klamath Co., July 9, 1920, Peck 9413 (GH) ; Buck Lake, Klamath
Co., July 24, 1897, Coville & Applegate 14 (US); Siskiyou Mts.,
July 8, 1887, T. Howell 1250 (NYS; OAC; MBG); Siskiyou, 4100
ft., June 26, 1920, Fischer 38 (US); Baker Co., dry run of canyon
of East Pine Creek, 3 mi. northeast of Cornucopia, Aug. 27, 1915,
Peck 5538 (GH); eastern Oregon, June 21, 1898, Cusick 1956
(GH; MBG; US; type collection of M. glauca Greene); Wallowa
Co., east of Tollgate Ranger Station, Wenaha Nat. For., 4500
ft., Aug. 3, 1916, Lawrence (US).

CaLrFORNIA: Mt. Shasta, Sept. 7, 1897, Canby 221 (GH; US);
Gold Lake, Sierra Co., 1960 m., Aug. 28-29, 1910, Eggleston 6272
(US; this and the next have the inflorescence characters of subsp.
pallida and the herbage of subsp. glauca) ; Gold Lake, Sierra Co.,
1960 m., Aug. 28-29, 1910, Eggleston 6269 (US); Siskiyou Co.,
Mt. Eddy, 4500 ft., July 16, 1915, Heller 12105 (US; GH; OAC;
МВС; US); Baxter Gulch, Trinity Co., June 17, 1914, Yates 441
(UC); Siskiyou Co., Mt. Hilt, Reaford (US); Shasta Co., near
Bald Mt., south of Fall River Mills, 3800 ft., June, 1903, Hall &
Babcock 4262 (DH); south fork of the San Joaquin River, Fresno
Co., 6700 ft., July, 1900, Hall & Chandler 688 (US; MBG);
Yreka, Aug., 1876, Greene (FM; authentic material of M. modo-
censis Greene); Buck Mt., near Summit, Humboldt Co., 5500 ft.,

[Vor. 12
64 ANNALS OF THE MISSOURI BOTANICAL GARDEN

July 31, 1912, Tracy (UC; US; suggests subsp. pallida); Butte
Co., Chico Meadows, 4000 ft., Heller 11604 (OAC; GH; US;
part of the collection suggests M. villosa subsp. Shelton?) ; Butte
Co., near Stirling, 3400 ft., June 7, 1913, Heller 10792 (GH ; US;
MBG); Siskiyou Co., Humbug Mt., 4000 ft., May 23, 1910,
Butler 1393 (US; RMH; МВС; GH); Gasquet, French Hill, Del
Norte Co., Sept. 14, 1912, Hastwood 2200 (US; GH); ridge between
Eagle & Bear Mt., Modoc Forest, 8000 ft., Aug. 18, 1918, Smith
1046 (J); Clear Creek, Butte Co., near Cherokee, 175 feet.,
Apr. 15-30, 1897, Brown 43 (US); Yreka, June 30, 1876, Greene
910 (MBG; authentic material of M. modocensis Greene) ; Sisson,
Siskiyou Co., 3555 ft., June 1-10, 1897, Brown 381 (MBG; US;
type collection of M. ovata, TYPE in US, an extreme form); Shasta
Springs, Siskiyou Co., June 13, 1905, Heller 8018 (GH; US);
Dorleska, Trinity Co., 6500 ft., July, 1909, Hall 8602 (US); Hot
Spring Valley, Plumas Co., July 7, 1897, Austin 1123 (US); Mt.
Bidwell, Modoe Co., Aug. 27, 1903, Manning 350 (US; suggests
subsp. discolor as it occurs in eastern Oregon) ; Tuolumne Meadows
on south gravelly slopes at base of granite dome, at cascades of
Dana Fork, 8700 ft., Aug. 5, 1923, Hall (UC); Middle Fork San
Joaquin River, 7600 ft., open slopes of Abies magnifica forest
near Hot Spr., Aug. 30, 1923, Hall 11891 (UC; inflorescence of
subsp. glauca, foliage of subsp. pallida).

NEVADA: Washoe Co., log railroad north of Verdi, 5300 ft.,
June 30, 1913, Heller 10891 (GH; US; MBG; type locality of
M. rubella; the plants of this collection elosely resemble the type) ;
Washoe Co., Hunter Creek, 6000 ft., Aug. 2, 1912, Kennedy (US;
GH; MBG); Washoe Co., Mt. Rose, 9300 ft., Aug. 26, 1911,
Heller 10346 (US; GH; MBG); Reno, 4500 ft., June 11, 1897,
Jones (MBG; US); Virginia City, 1863, Bloomer (US); Reno,
5000 ft., June 19, 1900, Stokes (DH); Genoa, Douglas Co.,
June 15, 1889, Lt. Bryan’s Exp. (MBG); Summit Lake Region,
July, 1901, Griffiths & Morris 328 (US); hills, northeast of Reno,
6000 ft., June 20, 1900, Stokes (US). The following have the
foliage of subsp. glauca but approach subsp. pallida in the nature
of the inflorescence: Hunter's Canyon, vieinity of Reno, 1350-
1500 m., July 18, 1913, Hitchcock 529 (US); Marlette Peak,
Washoe Co., 8000 ft., July, 1903, Hall & Chandler 4567 (US);

1925]
EPLING—MONOGRAPH OF MONARDELLA 65

King's Canyon, Ormsby Co., 1700-2000 m., June 21, 1902,
Baker 1113 (RMH; US; MBG; GH; referred to M. glauca by
Greene); Franktown, Washoe Co., 5000 ft., June 28, 1909,
Heller 9795 (GH). "The following are intermediate between the
longer-petioled forms of California and Nevada and the more
nearly sessile forms with oblong leaves, of Utah: Battle Moun-
tain, 1350 m., July 22, 1913, Hitchcock 576% (US); Allegheny
Creek, 8000 ft., Aug. 6, 1912, Nelson & Macbride 2172 (US;
MBG; GH; RMH); Gold Creek, 6300 ft., July 29, 1912, Nelson &
Macbride 2130 (GH; MBG; US); Ruby Mts. near Blaine P.O.,
9300 ft., Aug. 22, 1913, Heller 11109 (US; GH; MBG); Kingston
Canyon, Landor Co., Toiyabe Range, 7500 ft., July 28, 1913,
Kennedy 4202 (GH; the Toiyabe range proper is in Nye Co.);
Bunker Hill, Toiyabe Forest, 2250-3400 m., July 29, 1913,
Hitchcock 877 (US).

Montana: Mt. Logan, Aug. 1895, Shear 3164 (US).

Ipamo: Oneido Co., top of ridge on the west of Franklin Basin,
July 25, 1910, Smith 2286 (US); Caribou Forest, Aug. 4, 1913,
Young (US); Owyhee Co., Silver City, 7000 ft., July 19, 1910,
Macbride 434 (US; GH; RMH; MBG).

CoronApo: Elk Mountains, Pittsburgh, Aug., 1889, Eastwood
(CSM); Ouray, Ouray Co., Horsethief Trail, July 25, 1915,
Osterhout 5352 (RMH ; inflorescence of subsp. parvifolia).

Отан: Cottonwood Canyon near Salt Lake, Sept. 7, 1896,
Stokes (DH); Clayton Peak, 10,000 ft., Wahsatch Mts., Aug. 12-
26, 1903, Stokes (US; MBG); Fish Lake, 10,000 ft., Aug. 7, 1894,
Jones 5768 (MBG; US); Aquarius Plateau, 11,000 ft., Aug. 11,
1875, Ward 549 (MBG; US); Alta, 8500 ft., July 17, 1880, Jones
(US); 12-mile canyon, Wahsatch Mts., 2700 m., Sept. 3, 1907,
Tidestrom 477 (US); Aquarius Plateau, Aug. 5, 1905, Rydberg &
Carlton 7464 (US); Marysvale, Tate Mine, 11,500 ft., Aug. 28,
1894, Jones 5940 (US); Wahsatch Mts., 8200 ft., July 18-24,
1902, Pammel & Blackwood 3782 (GH; MBG); grassy slopes, La
Sol Mts., 9000-10,000 ft., July, 1899, Purpus 6694 (MBG; US);
City Creek Canyon, July 17, 1880, Jones (RMH); Big Cotton-
wood Canyon, 10,000 ft., Aug. 7, 1902, Cooper 348 (RMH); Mt.
Terrell, Wahsatch Mts., 3285 m., Aug. 27, 1908, Tidestrom 1829
(US); Big Cottonwood Canyon, 9000 ft., July 12, 1905, Garrett

[Vor. 12
66 ANNALS OF THE MISSOURI BOTANICAL GARDEN

1404 (US); Abajo Mts. (eastern range), 3000—3300 m., Aug. 17,
1911, Rydberg & Garrett 9723 (US; RMH); Pine Valley Mts.,
7000-8000 ft., May—Oct., 1898, Purpus 6198 (US); Mt. Nebo,
Aug. 15, 1905, Rydberg & Carlton 7706 (RMH; US; type collection
of M. oblongifolia Rydb., түре іп №. Y. Bot. Gard.); mountains
north of Bullion Creek, above Marysvale, July 23, 1905, Rydberg
& Carlton 7178 (RMH; GH; US); Wahsatch Mts., American
Fork, 1877, Hooker & Gray (GH); Alta, Wahsatch Mts., 10,000
ft., July 30, 1879, Jones 1109 (GH).

ARIZONA: Coconino Nat. For., 7900 ft., July 9, 1909, Pearson
245 (US; inflorescence that of subsp. parvifolia).

d. Subsp. pallida (Heller), comb. nov.

Monardella pallida Heller, Muhlenbergia 1: 36. 1904.

Madronella pallida Heller, Muhlenbergia 1: 138. 1906.

Branches scurfy-pubescent, cinereous, but not glaucous-appear-
ing, whitish rather than purple, leaves lanceolate-oblong, 2-3 cm.
long, somewhat rounded at the base and narrowed to a usually
margined petiole 2-8 mm. long, the upper seldom appearing sessile,
cinereous with a minute pubescence, but not glaucous-like;
bracts inconspicuous, broadly ovate, short-pubescent, often
woolly, purplish, ciliate, seldom exceeding the calyces, often
decurved but not reflexed; calyx short, woolly, often densely so
throughout, the glomerules hence appearing very compact; corolla
1-1.5 em. long, pallid, the lobes noticeably tapering; tube little
or not at all exserted.

Specimens examined:

CALIFORNIA: Round Meadow Camp, Sierra Nevada Mts.,
7000 ft., July, 1902, Grant 2030 (US); ridge near the lower end of
Donner Lake (south side), July 17, 1903, Heller 6959 (GH; US;
МВС; type collection of M. pallida Heller); Kaiser Crest, base of
south slope, 8600 ft., July 27, 1914, Smiley 616 (GH); Clio,
Plumas Co., 2080 m., Aug. 27, 1910, Eggleston 6281 (US) ; Gabbot
Meadow, Stanislaus Forest, Alpine Co., 1970 m., June 19, July
19, 1913, Eggleston 9689 (US); Gold Lake Trail, Clio, Plumas
Co., 1800 m., Aug. 27, 1910, Eggleston 6246 (US); Farwell Gap,
10,400 ft., Apr.-Sept., 1897, Purpus 6224 (GH; MBG; US);
above Giant Forest, Sequoia Nat. Park, Aug. 24, 1899, Copeland
19 (US); Grass Lake (Lake Tahoe Region), Aug. 8, 1909, Mc-

1925]
EPLING—MONOGRAPH OF MONARDELLA 07

Gregor 3 (GH; US); Butte Co., Summit above Jonesville, 7000
ft., July 29, 1917, Heller 12860 (MBG; OAC; US); Camp Echo,
Eldorado Co., on the Lineoln Highway, 7000 ft., A. A. Heller,
12186 (MBG; OAC); Prattville, Plumas Co., Heller & Kennedy
8774 (GH; US; MBG); Summit, Alpine Co., 9000 ft., Aug., 1892,
Hansen 438 (MBG); Bear River, 5500 ft., Amador Co., July 30,
1896, Hansen 1934 (US); Scott's Mt., 4000-7000 ft., Aug. 29,
1880, Engelmann (MBG); Horse Camp, Mt. Shasta, Aug. 12,
1920, Heller 13515 (US; MBG); Donner Pass, Placer Co., 7200 ft.,
July 26, 1919, Heller 13320 (GH; MBG; US); Eldorado Co., east
side of Ralston Peak, above Echo Lake, 7900 ft., Aug. 10, 1911,
Heller 12538 (GH; MBG; OAC; US); vicinity of Tuolumne
Meadows, Tuolumne Co., 8500-9500 ft., July 1902, Hall &
Babcock 3626 (DH ; perhaps better as subsp. glauca) ; Aspen Grove,
Lake Tahoe region, July, 1911, Hawver (US); Yosemite Nat.
Park, July 21-22, 1915, Hitchcock (US); Eagle Lake, Lassen Co.,
July 28, 1894, Baker & Nutting (RMH); south side of Mt. Shasta,
5000-10,000 ft., July 15-31, 1897, Brown 570 (US; MBG);
Plumas Co., 1876, Ames (MBG); Mt. Shasta, 6000 ft., Aug. 22,
1880, Engelmann (MBG); Colby, Butte Co., July, 1896, Austin
289 (US); headwaters of Hat Creek, Shasta Co., 2120 m., July 31-
Aug. 1, 1911, Eggleston 7457 (US); near Donner Pass, 1865,
Torrey 403b (US); Mt. Shasta, July 13-27, 1892, E. Palmer 2508
(US); Nevada Co., 6300 ft., Aug.-Sept., 1893, Carpenter (US);
Cisco, Placer Co., 6000 ft., July 1, 1910, Hall 8731 (US); Mt.
Shasta, 8000 ft., Aug.-Sept., 1902, Grant 796 (US); Mt. Shasta,
9000 ft., Aug. 28, 1889, Munson & Hopkins (US); McKinney’s,
Lake Tahoe, Aug., 1900, De Con 428 (US); Mt. Shasta, 5000-
9000 ft., Sept. 13, 1862, Brewer 1386 (US); Mineral King, Tulare
Co., 2750 m., July 31, 1891, Coville & Funston (US); Tar Gap
region, Tulare Co., Aug. 2, 1904, Culbertson 4448 (MBG; GH);
Angora Peak, north slope, 7600 ft., July 14, 1913, Smiley 7 (GH);
Sierra Nevada, 1875, Muir (MBG); Luthers Pass, 7800 ft.,
July 27, 1911, Abrams 4760 (US; GH); Truckee, Nevada Co.,
June, 1886, Sonne 284 (FM); Goose Valley, Shasta Co., June 29,
July 11, 1912, Eastwood 770 (GH; herbage of subsp. glauca and
inflorescence of subsp. pallida); Greenhorn Pass, 5000-6000 ft.,
Apr., Sept. 1897, Purpus 5582 (US); Truckee, 1750 m., July 14,

[Vor. 12
68 ANNALS OF THE MISSOURI BOTANICAL GARDEN

1913, Hitchcock 316 (US); Pine Ridge, Fresno Co., 5200 ft.,
June 15-25, 1900, Hall & Chandler 286 (MBG ; US; suggests subsp.
pinetorum somewhat); Rancheria Mt., north of Tuolumne R.,
8500 ft., July 24, 1909, Jepson 3404a (J); South Yallo Bolley,
July 2, 1897, Jepson 100% (J; “flowers nearly white—merely lilac-
tinged”); Mt. Shasta, near snow line, Aug. 4, 1894, Jepson 59}
(J.); Kennedy's meadow, Tuolumne Co., 6700 ft., Aug. 4, 1916,
Grant 896 (J; herbage of subsp. glauca and inflorescence of subsp.
pallida); Huntington Lake, Fresno Co., 7000 ft., July 20, 1917,
Grant (J); Campito Mt., White Mts., 10,700 ft., July 25, 1917,
Jepson 7286 (J).

Nevapa: Galena Creek, Mt. Rose, Washoe Co., 8000 ft.,
Aug., 1906, Kennedy 1236 (US). The following have the in-
florescence of subsp. pallida, or approach it more closely, but in
herbage suggest subsp. glauca: Mt. Rose, Washoe Co., 9700 ft.,
Aug. 26, 1911, Heller 10345 (GH; MBG; US); Clear Creek
Canyon, Ormsby Co., 2000-2615 m., July 22, 1902, Baker 1347
(MBG; GH; US).

OnEGON: Keno, Klamath Co., July 9, 1920, Peck 9413 (MBG);
Mt. Pitt (Jackson-Klamath Co.), Aug. 16, 1896, Gorman 457
(US); Gayhart Buttes, 2100 m., Aug. 7, 1896, Coville & Leiberg
266 (US).

e. Subsp. pinetorum (Heller), comb. nov.

Monardella pinetorum Heller, Muhlenbergia 1: 36. 1904.

Madronella pinetorum Heller, Muhlenbergia 1: 138. 1906.

Branches softly pubescent, not glaucous-appearing, leaves ovate
to lanceolate, 1.5-2.5 ст. long, somewhat rounded at the base,
tapering to a usually margined petiole 2-8 mm. long, the upper
seldom appearing sessile, softly pubescent, even to subvillous
principally on the under surface, the margin usually revolute and
obscurely crenate-dentate or entire; bracts inconspicuous but
erect, equaling the calyces, ovate, short-pubescent, purplish,
ciliate; calyx pubescent, the hairs spreading and not curling;
corolla 1.0-1.5 em. long, rose-color, the lobes noticeably tapering,
the tube exserted 1-2 mm.

Specimens examined:

CALIFORNIA: southern slope of Mt. Sanhedrin above the saw-
mill, July 19, 1902, Heller 5909 (MBG; GH; US); near Slap Jack

1925]
EPLING—MONOGRAPH OF MONARDELLA 69

Camp, west of alder springs, 5000 ft., Glenn Co., July 5, 1917,
Heller 12810 (OAC; US; MBG; GH; type collection of M. pine-
torum Heller); south fork Kaweah River, Tulare Co., July 22,
1904, Culbertson 4477 (GH; MBG); McCombers, Aug. 1, New-
berry (US); Longworthys, near North Fork, 4500 ft., July 16,
1912, Abrams 4951 (GH; DH; suggests M. villosa).

This group strongly suggests hybridization with M. villosa.

f. Subsp. parvifolia (Greene), comb. nov.

Monardella parvifolia Greene, Pl. Baker. 3: 22. 1901.

Monardella muriculata Greene, Pittonia 5: 84. 1902.

Madronella parvifolia Rydb. Bull. Torr. Bot. Club 33: 150.
1906.

Madronella muriculata Greene, Leaflets Bot. Obs. 1:169. 1906.

Monardella parviflora Nelson in Coulter & Nelson, Manual
Central Rocky Mts. 430. 1909 (not Greene; in error for M.
parvifolia Greene).

Branches scurfy-pubescent, cinereous, slender, seldom puberulent
and purplish; leaves lanceolate or oblong, 1-2 cm. long, tapering
at the base to a margined petiole 1-3 mm. long, cinereous, but
not glaucous-appearing, with a sparse, short pubescence; glomer-
ules small for the species, 1-2 cm. broad, bracts inconspicuous,
seldom exceeding the calyces, ovate, acute, or shortly acuminate,
pubescent, even shortly villous, ciliate; calyx 5-6 mm. long,
pubescent, sparingly villous around the teeth; corolla seldom ex-
ceeding a centimeter in length.

Specimens examined:

CALIFORNIA : foot of Mt. Whitney, 12,000 ft., Sept. (in flower),
1875, Rothrock 42 (US; GH); near Whitney Meadows, Aug. 20,
1891, Coville & Funston 1646 (US); along Big Cottonwood Creek
(Inyo Co.), Aug. 4, 1891, Koch 2158 (US); Ebbets Pass, on dry
mountain top, 8500-9000 ft., Aug. 1, 1863, Brewer 2006 (US);
Tuolumne Meadows, dry ledges, 8600 ft., July 20, 1907, Ware
2658 (GH); Mammoth Lakes just below Mary's Lake, 8900 ft.,
“In small openings in the subalpine forests of Abies magnifica
and Pinus monticola, with Chrysopsis Breweri, Ceanothus ve-
lutinus, Gilia aggregata and Haplopappus Bloomeri,' Aug. 30,
1923, Hall 11893 (UC); Junction Camp to Whitney, head of Kern
Canyon, 9500 ft., Aug. 1-12, 1900, Jepson 1055 (J).

[V or. 12
70 ANNALS OF THE MISSOURI BOTANICAL GARDEN

МЕУАРА: Morey Peak, 7000-8000 ft., May-Oct. 1898, Purpus
6369 (US); Star Peak, 8500 ft., Sept. 1867, Watson 826 (US; a
second collection bearing the same collection number was made
from E. Humboldt Range, 8000 ft., Aug. 1868; both are similar
and are the plants referred to by Watson in the Botany of the
King’s Expedition). The plant described by Greene as M. muri-
culata was from the same locality and the plant of Watson from
Star Peak resembles Greene’s type closely.

ARIZONA: near Sunset Peak, 6000 ft., July 13, 1901, Leiberg
5698 (US); Flagstaff, May—Oct. 1900, Purpus 8089 (US; MBG);
San Francisco Mts., crater at 10,000 ft., July 16, 1913, Goldman
2132 (US); Schulze's Ranch, 8000 ft., San Francisco Mts., July
7, 1891, MacDougal 404 (US; suggests subsp. ewodoratissima) ;
San Francisco Mts., 9000 ft., Aug. 3, 1898, MacDougal 363 (GH;
RMH; US); hillsides along the northern foot of San Francisco
Mts., 5550 ft., July 1, 1901, Leiberg 5620 (US; suggests subsp.
glauca); northern slopes of San Francisco Mts., 5500 ft., Aug. 29,
1901, Leiberg 5906 (US); near Kendrick Mts., 6600 ft., July 7,
1901, Leiberg 5653 (US).

CoronRADo: Black Canyon near Sapinero, 7200 ft., Sept. 1893,
Purpus 726 (FM); Black Canyon, 7000 ft., Aug. 1, 1901, Baker
678 (RMH; US; GH; MBG; type collection of M. parvifolia
Greene); Placerville, June 23, 1917, Payson 1097 (MBG); Black
Canyon, July 30, 1917, Bethel (CSM); El Late (Ute) Mts., Aug.
1875, Brandegee 1224 (MBG).

New Mexico: near Mogollon, Mogollon Mts., Socorro Co.,
Aug. 8, 1900, Wooton (US; RMH).

g. Subsp. australis (Abrams), comb. nov.

Monardella australis Abrams, Muhlenbergia 8: 34. 1912;
Davidson & Moxley, Fl. South. Calif. 313. 1923.

Branches decumbent or ascending, seldom erect, sparsely
pubescent, subvillous; leaves lanceolate or oblong, often acute,
green or cinereous, but not glaucous-like or silvery, 1-2.5 cm.
long, narrowed at the base to a petiole 1-3 mm. long; bracts
lanceolate, exceeding the calyces, short-acuminate, whitish or purple,
puberulent, frequently with a sparse pubescence in addition, the
margin ciliate, but not strongly; corolla about 1.5 cm. long, the
lobes slender, tapering, but not greatly.

1925]
EPLING—MONOGRAPH OF MONARDELLA 71

Specimens examined:

CALIFORNIA: San Bernardino Mts., 1880, Nevin (GH); San
Jacinto Mt., 8000-10,000 ft. (6000—10,000 ft. on some sheets),
July 22, 1917, Hall 713 (UC; DH; US); on trail to summit of
Round Valley, 9300 ft., July 11, 1908, Reed 2420 (US); in open
forests of Tamarack Valley, 9200 ft., July, 1901, Hall 2486 (UC;
BH; MBG; type collection of M. australis Abrams, TYPE in DH);
San Bernardino Mts., Aug. 1884, S. B. & W. F. Parish 462
(MBG); High Creek, San Bernardino Mts., 9100 ft., Aug. 23,
1923, Munz 7629 (BH); north side of Mt. Waterman, Aug. 29,
1917, Grinnell (BH); Tahquitz Valley (San Jacinto Mts.), no
data (BH); Box Springs on City Creek Road, 4800 ft., June 8,
1919, Johnston 2894 (BH); slope above Tamarack Valley, 10,000
ft., Sept. 7, 1922, Munz 6409 (BH); Little Bear Valley, 5300 ft.,
July, 1899, Hall 1298 (UC); San Jacinto Mt., Aug. 1881, S. B. &
W. F. Parish 462 (GH; MBG; FM; the latter is dated July, 1881,
from San Bernardino Mts., but bears the same number); Mt.
Grayback, San Bernardino Mts., June, 1880, Wright (GH);
Tahquitz Creek above Walters, San Jacinto Mts., July 10,
1909, Wilder 2 & 3 (UC); Deep Creek, San Bernardino Mts.,
July 30, 1901, Abrams 2046 (DH); Little Bear Valley, 5000 ft.,
July 22, 1897, Chandler (UC); Mill Creek Divide, San Bernardino
Mts., 8000 ft., Robertson 107 (UC); Tahquitz Valley, San Jacinto
Mts., July 10, 1909, Wilder 1 (UC); slope of San Jacinto Creek,
10,300 ft., Jepson 2814 (J); Tahquitz Valley, San Jacinto Mt.,
8000 ft., Jepson 2295 (J).

Some forms suggest M. macrantha var. tenuiflora but differ in
the flower.

11. M. linoides Gray, Proc. Am. Acad. 11: 101. 1876; Bot.
Calif. 1: 594. 1876; Syn. Fl. М. Am., ed. 2, 21: 357. 1886.

Perennial from a woody, usually decumbent, branching stem,
the bark light brown, checking and falling away; branches erect,
their arrangement often candelabra-like, rebranching below, if
at all, 30-50 cm. tall, pubescence minule, close, silvery; leaves
narrowly oblong, oblong, or narrowly lanceolate, 1-4 cm. long, acute
or obtuse, entire, narrowed to a petiole 2-4 mm. long, the upper-
most appearing sessile, the lowermost obovate or spatulate, all

[Vor. 12
72 ANNALS OF THE MISSOURI BOTANICAL GARDEN

covered with a minute silvery pubescence; glomerules 2-3 cm.
broad, bracts ovate to lanceolate, acuminate, equalling or exceed-
ing the calyces, scarious-membranous, whitish puberulent or
tinged with rose to purple, the margin subciliate; calyx 6-10 mm.
long, 13-nerved, even throughout and rather slender, closely pu-
berulent to rather sparingly hirsute, especially above, teeth
slender, hirsute within; corolla 12-15 mm. long, rose-purple or
pallid, the lobes slender, tending to oblong rather than lanceolate,
blunt, tube retrorsely puberulent within and without, the lips
subequal, the lobes of the upper about one-half its length, those
of the lower lip nearly or quite free, filaments retrorsely hispidu-
lous towards the base, but variable, anther-sacs divergent, the
connective equilateral, well developed.

KEY TO THE SUBSPECIES

А, . Bracts ovate to rotund, whitish...................... subsp. eulinoides
B. Bracts lanceolate, purple-colored. ....................... subsp. stricta

a. Subsp. eulinoides, nom. nov.

Monardella linoides Abrams, Muhlenbergia 8: 37. 1912;
Davidson & Moxley, Fl. South. Calif. 313. 1923.

M. viminea Greene, Pittonia 5: 85. 1902; Abrams, Muhlen-
bergia 8: 37. 1912; Davidson & Moxley, Fl. South. Calif. 313.
1923.

M. oblonga Greene, Pittonia 5: 83. 1902; Abrams, Muhlen-
bergia 8: 38. 1912; Davidson & Moxley, Fl. South. Calif. 313.
1923.

M. anemonoides Greene, Pittonia 5: 86. 1902.

Madronella linoides Greene, Leaflets Bot. Obs. 1: 169. 1906.

Madronella viminea Greene, Leaflets Bot. Obs. 1: 169. 1906.

Madronella oblonga Greene, Leaflets Bot. Obs. 1: 169. 1900.

Madronella anemonoides Greene, Leaflets Bot. Obs. 1: 169.
1906.

Herbage silvery, with a close minute puberulence, the leaves
narrowly oblong or narrowly lanceolate; bracts broadly ovate,
shortly acuminate, whitish puberulent, infrequently tinged with
purple, occasionally pubescent, surpassing the calyces, not in-
frequently enveloping them; calyx puberulent or sparingly hispid;

1925]
EPLING—MONOGRAPH OF MONARDELLA 73

lobes of the corolla tending to narrowly oblong, tapering but
little, the upper lip usually incised to about one-half its length.

Specimens examined:

CALIFORNIA: Oriflamme Mine, near San Diego, July 28, 1875,
E. Palmer 261 (2296) (GH, түре; MBG; UC); Oriflamme Canyon
near Cuyamaca, June 28, 1903, Abrams 3932 (US; GH; MBG;
BH); Tahquitz Valley, San Jacinto Mts., 7000 ft., June-July,
1901, Hall 2430 (US; UC; MBG); 20 mi. south of Palm Springs,
July 30, 1897, Hall 758 (UC); Coyote Canyon, Santa Rosa Mts.,
5000 ft., June, 1901, Hall 2137 (UC); eastern base of San Jacinto
Mts., June, 1901, Hall 2110 (UC); Santa Rosa Mts., Santa Rosa,
6500 ft., June 30, 1922, Munz 5921 (UC; BH); Big Morongo
Canyon, San Bernardino Mts., 3000 ft., June 15, 1894, S. B.
Parish 3009 (US; MBG); San Diego Co., 1889, Orcutt (US);
Laguna Mts., July, 1889, Orcutt (US). The following illustrate
the form described as M. viminea: McCoon’s Ranch near Poway,
400—500 ft., June 8-9, 1897, S. B. Parish 4421 (MBG; GH; US);
near San Diego, 1880, Vasey 491 (US, түрЕ of M. viminea
Greene); San Diego, May, 1906, T. S. Brandegee (FM); San
Diego, May 21, 1894, Brandegee (UC); ''along river bed," San
Diego Co., ?1878, Cleveland (GH). The following illustrate the
form known as M. anemonoides: Greenhorn Mts., 6000—7000 ft.,
Kern Co., June 7-15, 1888, E. Palmer 69 (MBG; US, type col-
lection of M. anemonoides Greene, TYPE in US); Pah Ute Peak,
5000-6000 ft., June, 1897, Purpus 5096 (UC; US; GH; MBG);
Argus Peak, Kern Co., 5000-6000 ft., June, 1897, Purpus 5098
(US; UC; GH; MBG); Cottonwood Cr., 7000-7500 ft., Inyo Co.,
Aug. 1896, Purpus 1947 (UC); road between Bishop & Andrews
Camp, Inyo Co., July, 1913, K. Brandegee (UC) ; Panamint Mts.,
Wild Rose Canyon, June 24, 1891, Coville & Funston 2045 (US);
Westgard Pass, between Deep Springs Valley and Big Pine,
Inyo Co., July 19, 1918, Ferris 1381 (DH). The following il-
lustrate the form known as M. oblonga Greene: Mt. Pinos, 6500
ft., July 7, 1904, Grinnell (UC, a good match for the type of
M. oblonga Greene); Tehachapi Mts., vicinity of Bisses station
(?Bissel), June 28, 1895, Dudley 476 (DH; US; UC); Griffin's,
Ventura Co., July, 1902, Eimer 3952 (GH; US); Frazier Mt.,
6000 ft., June 15, 1896, Dudley & Lamb (DH; BH); Mt. Pinos,

[Vor. 12
74 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Ventura Co., North Fork, 6000 ft., June 28, 1905, Hall 6461
(UC); Kaiser Crest, Fresno Co., 9700 ft., July 28, 1914, Smiley
646 (GH).

Lower CaLirornia: Palm Valley, June 3, 1883, Orcutt 382
(MBG; GH); San Pedro Martir, 7000 ft., May 6, 1893, T. S.
Brandegee (UC); “Mountains of the Peninsula," July 25, 1885,
Orcutt (UC); Cantites (?) Mts., July 26, 1883, Orcutt 927 (GH);
mountains of Lower Calif., July 25, 1883, Orcutt (FM).

b. Subsp. stricta (Parish), comb. nov.

Monardella linoides stricta Parish, Erythea 7: 96, 1899.

M. epilobioides Greene, Pittonia 5: 85. 1902; Abrams, Muhlen-
bergia 8: 35. 1912; Davidson & Moxley, Fl. South. Calif. 313.
1923.

M. epilobioides var. erecta Abrams, Muhlenbergia 8:36. 1912;
Davidson & Moxley, Fl. South. Calif. 313. 1923. -

Madronella epilobioides Greene, Leaflets Bot. Obs. 1: 196. 1906.

Herbage similar to that of subsp. typica but tending in some to
have a short sparse pubescence in addition to the silvery covering;
bracts lanceolate, distinctly short-acuminate, whitish, tinged with
rose, to a deep purple, puberulent or sparingly pubescent, the
margin subciliate; calyx puberulent to sparsely hispid; the lobes
of the corolla tending to lanceolate rather than oblong, the
upper lip more often cut to less than half its length.

Specimens examined:

CALIFORNIA: The following illustrate the form known as M.
epilobioides Greene: San Antonio Mts., July, 1896, Hall (RMH;
BH; US); 12 mi. west of Cajon Pass, Aug. 6, 1896, Hall 297 (US;
UC); the two preceding suggest M. viminea; San Bernardino
Mts., Aug. 1884, S. B. Parish 2077 (UC; түре of M. linoides var.
stricta Parish); Le Montaine, north of Big Pines, 7300 ft., July 5,
1922, “very abundant on open slopes but hardly in bloom at this
time" Peirson 3160 (J); San Bernardino Mts., Aug. 1881, S. B. &
W. F. Parish 462a (GH, suggests M . viminea) ; Mill Creek Canyon,
1904, Smith 105 (UC); Bear Valley, San Bernardino Mts., 6500
ft., June 22, 1894, S. B. Parish 3008 (UC; MBG; US; type col-
lection of М. epilobioides Greene, TYPE in US); Little Green
Valley, San Bernardino Mts., 7200 ft., July, 1904, Hall 2 (UC).
The following illustrate the form known as M. epilobioides var.

1925]
EPLING—MONOGRAPH OF MONARDELLA 75

erecta Abrams: Mt. San Gorgonio, 7500 ft., July 23, 1904, Grant
795a (FM; MBG; UC); Santa Ana R., San Bernardino Mts.,
6100 ft., July 27, 1906, J. & H. W. Grinnell 306 (US); Upper
Santa Ana Canyon, 8500 ft., July 26, 1906, Hall 7578 (RMH;
US; BH; MBG; UC); Fish Creek, San Bernardino Mts., 8700 ft.,
Sept. 1, 1921, Jaeger (BH); above Green Valley, July, 1899,
Hall 1362 (UC); on dry ridges, Bear Valley, Aug. 3, 1902,
Abrams 2861 (GH; MBG; UC; BH; US, type collection of M.
epilobioides var. erecta Abrams, TYPE in DH); east of Fish Camp,
San Bernardino Mts., 6700 ft., July 17, 1921, Johnston 2898
(BH, suggests M. australis Abrams); San Bernardino Mts., 7000
#., July 19, 1898, Hall 1021 (UC); South Fork Meadows, Santa
Ana Canyon, 8200 ft., Aug. 6, 1906, Hall 7676 (UC).

МЕУАРА: Lee Canyon, Charleston Mts., Clark Co., 8000 ft.,
July 25, 1913, Heller 10984 (US; MBG;GH). _

Arizona: Little Meadows, June 28, 1902, Stephens (UC).

Lower CALIFORNIA: Hansens, Sept. 18, 1884, Orcutt 1224
(GH); San Pedro Martir, 7000 ft., May 6, 1893, T. S. Brandegee
(UC); San Pedro Martir, Aug. 1903, Robertson 54 (UC); Oal-
lecitos, San Pedro Martir Mts., 8000 ft., July 15, 1905, Goldman
1240 (US).

After a careful study of a considerable mass of herbarium
material, representing M. linoides from most of the localities
where it is known to grow, the opinion has been formed that
some, at least, of the numerous forms have been caused by
hybridization. Whether this is true remains to be determined.
In the San Bernardino Mountains, especially, the data are very
confusing. One may pass by imperceptible gradations from the
form originally described by Dr. Gray as M. linoides to the plant
of the interior mountains described herein as M. odoratissima
subsp. glauca. Inasimilar way he may pass into M. odoratissima
subsp. australis or into the group described as subsp. stricta.
Until some evidence is forthcoming as to the nature of these
variants, whether partly or wholly environmental, whether
hybrid, due to the existence of several closely related subspecies,
or both, it seems desirable to retain the present arrangement
which will permit of fairly close determination of material.
Furthermore, the two groups as thus outlined have a distribution
which seems fairly consistent.

[Vor. 12
76 ANNALS OF THE MISSOURI BOTANICAL GARDEN

M. viminea Greene is a plant in which the stems are unusually
long, with long internodes, the leaves similarly elongate and
glabrate, and the calyx and bracts pubescent. In shape and
size the bracts are midway between subsp. eulinoides and subsp.
stricta. As it is found on the type sheet, the several stems have
been coiled, in order to gain space, the whole suggesting the wil-
lowy aspect from which the name was derived. Few collections
may be referred to this, and the author has seen no plants as ex-
treme as the plant of the type sheet.

Only a photograph of the type sheet of M. oblonga Greene has
been available for study. However, the collection of Grinnell
on Mt. Pinos (UC, 149541) is a very good match for both the
description and photograph; the type was collected “іп the
mountains south of Tehachapi," June 24, 1889, by Greene and is
at Notre Dame. In general, the more hairy calyx and broader
leaf characterize the plants of the more northern stations.

M. anemonoides Greene is an extreme form in which the bracts
are unusually developed in such a way as to envelop and conceal
the flowers. The bracts of the plant collected at the same time
and deposited at the Missouri Botanical Garden (the type is in
the U. S. National Herbarium) are not greatly above normal, and
apart from this collection the author has seen nothing to equal
the type sheet.

M. linoides subsp. stricta may, in a general way, be divided into
two forms, namely, one (M. epilobioides Greene) with but two or
three slender, erect, fertile branches (suggestive of M. viminea
Greene) which arise from a low tuft of sterile branches, with
elliptic-oblong leaves, and a second form (M. epilobioides var.
erecta Abrams) in which the fertile branches are numerous, sub-
equal, and more or less fastigiate, with leaves shorter and linear-
oblong.

§ Section IV. ANNUAE

12. M. undulata Benth. Lab. Gen. et Sp. 332. 1834; in DC.
Prodr. 12: 190. 1848; Gray, Proc. Am. Acad. 11: 102. 1876;
Bot. Calif. 1: 594. 1876; Syn. Fl. №. Am., ed. 2, 2:: 358. 1886;
Jepson, Fl. West. Middle Calif., ed. 2, 363. 1911.

Madronella undulata Greene, Leaflets Bot. Obs. 1: 168. 1906.

1925]
EPLING—MONOGRAPH OF MONARDELLA 77

Annual or perennial, usually forming a bush-like plant 20-40
em. tall, or erect, rather slender, the ascending branches un-
branched, branches purple, puberulent; leaves somewhat succu-
lent, oblanceolate-oblong, 2-5 cm. long, obtuse, glabrate, thinly
villous, or shortly pubescent, the margins undulate or crisped,
narrowed to a short petiole; glomerules compact, 2.5-3.5 cm.
broad, bracts broadly ovate or orbicular to elliptical, obtuse or
acute, variable in size, equal to the calyces or much exceeding
the flowers, submembranous with green or purple prominent
parallel veins, or scarious, glabrate to villous; calyx 5-9 mm.
long, variable on the same plant, 13-15-veined, tapering down-
wards, subglabrous and subscarious below, green or purplish and
villous above, or villous throughout, the teeth ovate-triangular,
obtuse, hairy, white within; corolla rose-purple, 14-20 mm. long,
the tube twice the length of the corolla or less, the throat ample
and hairy within, the upper lip somewhat shorter, the lobes co-
alesced two-thirds its length or more, those of the lower coalesced
one-third to one-fourth its length, linear-oblong or tapering;
anthers divergent, the connective equilateral, well developed.

CALIFORNIA: Gigling Monterey Co., June 1903, Elmer 4379
(OAO); along the railroad 2 mi. northeast of Del Monte, Mon-
terey Co., July 31, 1906, Heller 8426 (GH; US; MBG); Point
Reyes, Marin Co., July, 1903, Elmer 4611 (MBG; US); Point
Reyes, May 5, 1901, Eastwood (GH); Point Reyes, June 23, 1915,
Eastwood 4773 (GH); San Francisco, 1865, Bolander (MBG;
GH); San Francisco, near Lake Merced, August 5, 1913, Suksdorf
786 (GH); “California,” Hartweg (GH); Bardins, June, 1903,
Elmer 4379 (US; MBG); “Northern California," D. Douglas
(GH; a portion of the type collection cited by Bentham); Lake
Merced near San Francisco, June, 1905, K. Brandegee (RMH);
Clarke Creek, 10 mi. from San Luis Obispo, June 26, 1876, E.
Palmer 362 (GH; MBG; US); Arroyo Grande, San Luis Obispo
Co., June, 1887, Lemmon 4622 (GH); sea-shore hills, Feb. 3, 1882,
Summers (MBG; US); Nipomo, San Luis Obispo Co., sandbanks,
Bolander (GH); Nipomo, San Luis Obispo Co., Apr. 11, 1861,
Brewer 421 (US); Castroville, Monterey Co., K. Brandegee (US).

Var. crispa (Elmer), comb. nov.
Monardella crispa Elmer, Bot. Gaz. 39: 46. 1905.

[Vor. 12
78 ANNALS OF THE MISSOURI BOTANICAL GARDEN

A low shrubby plant 20-30 cm. tall, bush-like in appearance,
the branches simple or branching, lanately villous, the older parts
covered with a light brown checking bark; leaves oblanceolate-
oblong, 2-5 cm. long, as much as 1 cm. broad, very blunt at the
apex, rather succulent, pubescent, undulate or crisped, narrowed
into a short petiole; bracts ovate or roundish, villous; calyx 6-8
mm. long; corolla 12-14 mm. long, the lobes tapering and tending
to lanceolate rather than oblong.

Specimens examined:

CALIFORNIA: Surf, Santa Barbara Co., May, 1902, Elmer 3965
(MBG; US; type collection of M. crispa Elmer, түрк in DH);
Santa Maria, Santa Barbara Co., 1882, Jared (GH); Surf, Santa
Barbara Co., “оп beaches but most abundant on the hills,"
May, 1909, K. Brandegee (UC; GH; US); no locality stated,
. Coulter 586 (GH; cited by Bentham); San Luis Obispo, Summers
(GH); Point Reyes, May, 1906, Eastwood (US) ; Casmalia, Santa
Barbara Co., Eastwood (US).

13. M. Douglasii Benth. Lab. Gen. et Sp. 332. 1834; in DC.
Prodr. 12: 190. 1848; Gray, Proc. Am. Acad. 11: 102. 1876;
Bot. Calif. 1: 595. 1876; Syn. Fl. №. Am., ed. 2, 21: 357. 1886;
Jepson, Fl. West. Middle Calif., ed. 2, 363. 1911.

Monardella candicans var. venosa 'Torrey, Pacif. R. R. Rept. 4: -
123. 1857 (Whipple's Exp.).

Madronella Douglasii Greene, Leaflets Bot. Obs. 1: 168. 1906.

Annual, erect, with divarieate branches, 20-30 cm. tall, or often
simple, with a single terminal inflorescence; stems purplish, pu-
berulent; leaves 1-3 cm. long, lanceolate to linear-oblong, pu-
berulent, narrowed at the base to a petiole 1-5 mm. long; glomer-
ules 1.5-3 em. broad; bracts ovate-lanceolate, surpassing the
calyces, with a strong midrib and a well-defined marginal vein
formed by the confluence of the ascending lateral veins, the intravenous
tissue like isinglass when dry, transparent, the veins purple, rough-
pubescent, margin ciliate; calyx 7-9 mm. long, 15-nerved, pubescent
or hirsute, the teeth rigid, acute, subcuspidate, pubescent within;
corolla deep reddish purple, 11-12 mm. long, the tube somewhat
exserted, retrorsely puberulent, lips subequal, the lobes of the
upper coalesced more than half its length, those of the lower nearly

1925]
EPLING—MONOGRAPH OF MONARDELLA 79

free, slender, tapering slightly; the anther-sacs subparallel, con-
fluent behind, the connective scarcely wider than the filament.

Specimens examined:

CALIFORNIA: Moragua Valley (Bay Region), Aug. 1863, Bolander
2499 (GH; UC; US); plains of the Feather River near Marysville,
May 25, 1854, Bigelow (Whipple's Exp.) (GH; US; type collection
of M. candicans var. venosa Torrey); Oakland, May 29, 1892,
Brandegee (GH); Gilroy, June 16, 1896, Jepson (US; MBG; J);
Mt. Diablo, July, 1908, Elmer 4544 (MBG); Plumas Co., Austin
(MBG); Oakland Hills, 1865, Bolander (GH; MBG); near. San
Francisco, Kellogg (GH); Mt. Hamilton, July, 1905, R. I. Smith
(RMH); Cherokee, Butte Co., May, 1879, Bidwell (GH); Ala-
meda, 1876, Vasey (GH); Butte Co., 1882, Parry (UC); Chico
Valley, May, 1882, Parry (UC); Black Canyon (? Marin Co.),
July, 1885, K. Brandegee (UC); San Jose, May 20, 1897, Chipman
(US); locality not given, 1875, Vasey (US).

14. M. lanceolata Gray, Proc. Am. Acad. 11: 102. 1876;
Bot. Calif. 1: 594. 1876; Syn. Fl. №. Am., ed. 2, 2!: 357. 1886;
Hall, Univ. Calif. Publ. Bot. 1: 108. 1902; Jepson, Fl. West.
Middle Calif., ed. 2, 363. 1911; Abrams, Muhlenbergia 8: 41.
1912; Fl. Los Angeles, ed. 2, 318. 1917; Davidson & Moxley,
Fl. South. Calif. 314. 1923.

Monardella sanguinea Greene, Pittonia 5: 86. 1902.

M. acuta Greene, in Herb. Baker. 1193. 1902.

Madronella sanguinea Greene, Leaflets Bot. Obs. 1: 169. 1906.

Madronella lanceolata Greene, Leaflets Bot. Obs. 1: 169. 1906.

Annual, erect, 30-50 cm. tall, branching throughout, but with a
tendency to branch chiefly in the upper axils, the branches di-
varicate but curving upwards, puberulent, purplish; leaves lan-
ceolate, 3-4 cm. long, obtuse, entire, sparsely puberulent, nar-
rowed to a slender petiole 0.5-1.5 cm. long; glomerules 1.5-3 cm.
broad, bracts ovate-lanceolate, acute, surpassing the calyces, sca-
brous, membranous but green, pinnately veined with numerous
readily observed net-like secondary veins, the principal veins promi-
nent, often costate; calyx 6-8 mm. long, glabrous or scabrous, some-
times bristly at the sinuses, veins slender, typically 13, teeth ovate-
triangular, acute, hirsute within; corolla rose-purple, 12-15 mm.

[Vor. 12
80 ANNALS OF THE MISSOURI BOTANICAL GARDEN

long, the tube somewhat exserted, puberulent, the limb 3-5 mm.
long, the upper lip shorter, lanceolate in outline, the lobes co-
alesced one-half its length or more, those of the lower lip free
nearly or quite to the base, tapering slightly; anther-sacs diver-
gent at an angle of about 60?, the connective about as wide as
the filament, little developed and scarcely evident from behind;
nutlets oblong-oval, about 2 mm. long.
pecimens examined:

CALIFORNIA: Tallae, Lake Tahoe, 6300 ft., July 18, 1913,
Smiley 138 (GH); Tioga road above Aspen Valley, 6500 ft.,
Aug. 24, 1916, Smiley (GH); Yosemite Valley, Sept.-Oct. 1878, .
Phillips & Sargent (GH); Glendora, July 7, 1902, Abrams 2662
(GH; MBG); Cuyamaca, 4000 ft., June 30, 1917, Spencer 635
(GH); Cajon Pass, June 8, 1861, Cooper (GH; US); Shasta Co.,
between the McCloud and Sacramento Rivers, July 23, 1916,
Heller 12499 (GH; MBG); Oakgrove Canyon, Liebre Mts.,
3500—4000 ft., July 19-21, 1908, Abrams & McGregor 350 (GH);
Bloomington, June 2, 1917, S. B. Parish 11268 (GH; BH; MBG);
Colton, April, 1885, S. B. Parish 1750 (GH); locality and date
not given, Bridges 308 (GH); Fort Tejon, 1857-58, de Vasey 77
(GH); south fork, Kaweah River, Tulare Co., July 20, 1904,
Culbertson (Baker 4489) (GH; MBG); Nevada City, July 14,
1905, Heller 8114 (GH); no locality given, 1872, Gray (GH);
San Diego, 1875, Cleveland (GH; some of these approach var.
microcephala); Ramona Valley, San Diego Co., June 19, 1903,
Abrams 3773 (GH; MBG); Yosemite Valley, July, 1866, Bolander
6320 (GH; US); Calaveras Co., 1877, Hooker & Gray (GH);
foothills of the Sierra Nevada, 1865, Torrey (GH); Ojai Valley,
July, 1875, Rothrock 175 (GH); Snowdon Ranch, Calaveras Co.,
July-Aug., 1890, Jepson 50f (J); Hetch-Hetchy, ‘‘opens” on
valley floor, Jepson 3437 (J); Augustine’s ranch, Palomar, May
30, 1901, Jepson 1549 (J); Mineral King road, 5900 ft., Aug. 1-12,
1900, Jepson 1160 (J); Colton, June, 1882, S. B. & W. F. Parish
1430 (GH) ; southern California, 1876, Parry & Lemmon 331 (GH;
MBG); Tulare Mts., May, 1878, Lemmon 336 (GH); Middle Tule
River, 3000-4000 ft., April-Sept. 1897, Purpus 5040 (GH; MBG);
Middle Tule River, 4000-5000 ft., April-Sept. 1897, Purpus 5050
(GH; MBG); Newcastle Road, Plumas Co., May, 1894, Ames

1925]
EPLING— MONOGRAPH OF MONARDELLA 81

(GH); Yosemite Valley, June 28, 1911, 4000—5000 ft., Abrams 4562
(GH); Pitt to Baird, Shasta Co., July 25,1912, Eastwood 1439 (GH ;
MBG); West Point Bridge, 2300 ft., July 7, 1896, Hansen 1824
(MBG); Mariposa, June 14, 1903, Congdon (MBG); Nevada Co.,
July 14, 1905, Heller 8114 (MBG); Ranger Station, Amador Co.,
2000 ft., June, 1891, Hansen 128 (MBG); mountains above Clare-
mont, no date, Davis (MBG); Mojave River, June 1, 1876, Е.
Palmer 363 (MBG); Sierra Santa Monica, June, 1889, Hasse
(MBG); Mt. Lowe, Los Angeles Co., 1903, Grant 796a (MBG);
Strawberry Valley, San Jacinto Mts., 6000 ft., Aug. 28, 1896, Hall
340 (MBG); southern California, near the boundary, June, 1880,
S. B. Parish (MBG); San Bernardino Mts., 3000 ft., June 29,
1888, S. B. Parish (MBG; it is probably this collection which
was referred by Greene to M. sanguinea); Tigh’s ranch, San
Diego Co., July 4, 1875, E. Palmer 294 (MBG); North Fork and
vicinity, May 30-June 8, 1903, Griffiths 4628 (MBG); Madera,
July 10, 1904, Griffiths 6589 (MBG); Tehachapi Peak, June 28,
1895, Dudley 348 (OAC); Nevada (?City), no date, Pratten
(MBG); La Crescenta, 1897, Wislizenus 1307 (MBG); Dry
Canyon, July 15, 1917, Johnston 1912 (BH); Palomar, July, 1901,
Schellenger (BH); Strawberry Valley, San Jacinto Mts., 5200-
6000 ft., July, 1901, Hall 2527 (BH); Santa Ana River, San
Bernardino Mts., 6300 ft., Aug. 21, 1922, Munz 6149 (BH);
Little Chico Cr., 2000 ft., July 5, 1900, Leiberg 5022 (US);
Deer Cr. Canyon, Tehama Co. July 17, 1911, Eggleston
7277 (US); Breckinridge Range, Kern Co., 5000 ft., 1905,
Hopkins (US); between Temecula and Pala, July 10, 1915,
Collins & Kempton 224 (US); San Jacinto Plains, 1880, Vasey
490 (US); Tehachapi Pass, Kern Co., June 25, 1891, Coville &
Funston 1113 (US); Lyttle Cr., San Gabriel Reserve, 1800 m.,
April 29, 1898, Leiberg 3364 (US); Tejunga Wash, Los Angeles
Co., July 6, 1905, Grinnell (US); Yosemite Valley, Aug. I5
1872, Redfield 6497 (MBG); Yosemite Valley, 1886, Bolander
6320 (MBG); Forest Ranch, Butte Co., July 24, 1914, Heller
11628 (GH; OAC; US); Azusa, June 22, 1915, Macbride & Payson
731 (GH); Plumas Co., Ames (GH); San Bernardino Mts., S. B.
& W. F. Parish 405 (MBG); Palomar Mt., San Diego Co.,
. Sept. 14, 1922, Spencer 635 (BH); Etiwanda, June 16, 1921,

[Vor. 12
82 ANNALS OF THE MISSOURI BOTANICAL GARDEN

1500 ft., Munz 4658 (BH); San Dimas Canyon, July 4, 1915,
Davis (BH); San Jacinto, 6000 ft., July 15, 1898, Anthony (UC);
Little Bear Valley, San Bernardino Mts., 5000 ft., July 31, 1897,
Chandler (UC); head of Hemet Valley, 5000 ft., July 3, 1922,
Munz (UC); Julian, Aug. 4, 1892, Dunn (UC); Downieville,
Sierra Co., 1909, Kennedy 29 (UC); El Dorado Co., Aug. 1914,
K. Brandegee (UC); Rennies Sta., 3000 ft., June 27, 1897, Reed
(BH); San Joaquin Hills, Orange Co., July 13, 1901, Abrams 1788
(BH); Fort Tejon, June, 1881, S. B. Parish (J); Fish Cañon,
San Gabriel Mts., 1000 ft., July 1, 1919, Peirson 511 (J).

NEvaDA: Lake Tahoe, 6300 ft., Washoe Co., Aug. 8, 1906,
Kennedy 1459 (MBG); Kings Canyon, Ormsby Co., 1700-2000
m., June 30, 1902, Baker 1193 (GH; MBG; US); western
Douglas Co., 6250 ft., Hall & Chandler 4593 (UC; US).

ARIZONA: Mont Cr. (northern Ariz.), Aug. 9, 1894, Wilson
(UC, as “M. lanceolata Arizonica”’).

As it occurs in the southern part of its range, M. lanceolata
tends to become more slender, lower in height, with fewer
branches, these being in the upper axils and divaricate. The
glomerules are often reduced in size and the corollas are a deeper
color. It was such a plant as this which was described by Greene
as M. sanguinea. The author has been unable to find any dif-
ferences which might not be ascribed to the differing conditions
of its habitat. M. acuta Greene is a depauperate, unbranched
form bearing a single terminal glomerule, which is in all respects
that of M. lanceolata.

Typical M. lanceolata var. microcephala occurs only in the
extreme southern portion of the specific range. The author has
seen only one or two collections, even from the type locality,
which equal the type collection in the extreme reduction in the
size of the glomerules. All gradations occur, and it is a matter of
opinion as to where the line may be drawn between the variety
and the more typical plant. The writer has not seen the Orcutt
collection which formed the basis for M. peninsularis, but from
a comparison of the Orcutt material from the same locality with
Greene’s description, he feels certain that such a plant was rep-
resented, and that M. peninsularis is synonymous with the
variety microcephala. In describing var. microcephala Abrams

1925]
EPLING—MONOGRAPH OF MONARDELLA 83

differentiated between it and the typical plant, among other
things, by the absence of hispid hairs at the sinuses of the calyx.
Johnston in describing var. glandulifera notes a point of difference
between his plant and var. microcephala in that the hispid hairs
are present. Careful examination shows that these hispid hairs
about the base of the calyx teeth may be variously developed
in different collections of the typical plant and the variety, and
that while some sinuses may be naked, the calyx is rarely wholly
so. This is true of the type collection of M. lanceolata var.
microcephala and of the Orcutt collections. Furthermore, the
stalked glands upon which the var. glandulifera was based may
be found occasionally in both the typical plant and the variety
microcephala, more especially in the southern forms, although no
plants were observed in which they were as abundant as in the
Johnston collection. It accordingly seemed preferable to con-
sider Mr. Johnston’s plant as a form of var. microcephala. A
great diversity may be observed in the size of the glomerules
upon a single plant, either of the variety or the more typical
specimens, especially in the southern part of the range, such that
it would appear that while environmental conditions had been
favorable for the normal development of some of the glomerules
others had been stunted or even aborted. At the same time
vigorous plants with well-developed foliage usually have the
glomerules of about the same size.

The foliage varies considerably, and robust, rankly growing
plants may have quite a different aspect from those subjected to
drier less favorable conditions. This variation in the leaf form
is paralleled by an unusual development of the outermost bracts,
or the involucral pair of leaves, which in some plants reach a
length of several centimeters and thus form a reflexed foliaceous
involucre. All gradations may be observed. Such a plant
(N. C. Wilson, Mont Cr., Ariz., UC, 25461) has been named in
the herbarium by some one “М. lanceolata Arizonica.” It is not
peculiar to Arizona, however, but extends throughout most of
the specific range, occurring in Butte Co., the Yosemite, the San
Bernardino Mountains, and San Diego Co. It is considered
to be only an environmental form.

One plant of Heller’s collection, No. 12499 (OAC, 8932), while

[Vor. 12
84 ANNALS OF THE MISSOURI BOTANICAL GARDEN

similar in all other respects to M. lanceolata, bore reduced corollas,
with very small stamens. Since apparently normal (but smaller)
seeds were produced, it will be of interest to note whether this
variant has established itself. Мо other similar plant was found
among the numerous collections of M. lanceolata.

Var. microcephala Gray, Syn. Е. N. Am., ed. 2, 2: 459.
1886; Abrams, Muhlenbergia 8: 42. 1912; Davidson & Moxley,
Fl. South. Calif. 314. 1923.

Monardella peninsularis Greene, Pittonia 5: 87. 1902.

M. lanceolata var. glandulifera Johnston, Bull. South. Calif.
Acad. Sci. 18: 20. 1919.

Madronella peninsularis Greene, Leaflets Bot. Obs. 1: 169.

1906.

A form with the glomerules reduced in size to 1 cm. or less
broad, the bracts and flower parts being reduced accordingly.
The stems and branches are more slender and divaricately
branched. All gradations seem to occur.

M. peninsularis Greene is based upon a collection made by
Orcutt, June 6, 1885, in northern Lower California.

Specimens examined:

CALIFORNIA: Potrero, July 24, 1883, Orcutt 928 (GH, TYPE;
MBG); El Campo, dry valley, Aug. 14, 1917, Munz 1681 (BH);
Cameron's Ranch, Laguna, June 22, 1894, Schoenefeld 3692 (US);
Pine Valley, Aug. 12, 1894, Mearns 3983 (US); near San Diego,
1875, E. Palmer 257 (US); San Diego Co., April, 1890, Orcutt
(MBG); Pine Valley, San Diego Co., Orcutt (MBG); mountains
of San Diego Co., Aug. 1879, Orcutt 57 (GH); Brown's Flats,
San Antonio Mts., 4300 ft., Sept. 1, 1918, Johnston 2139 (UC;
BH, 4040; type collection of var. glandulifera Johnston, TYPE
in BH); ?Fish Creek, San Bernardino Mts., 6500 ft., July 10,
1906, Grinnell 261 (US).

Lower CaurrFonNIA: Hansen’s, July 30, 1883, Orcutt 929 (GH);
La Gralla, San Pedro Martir, 7000 ft., July 20, 1905, Goldman
1256 (US).

15. M. Breweri Gray, Proc. Am. Acad. 7: 386. 1867, and 11:
102. 1876; Bot. Calif. 1: 594. 1876; Syn. Fl. N. Am., ed. 2,
2': 357. 1886; Jepson, Fl. West. Middle Calif., ed. 2, 363. 1911.

1925]
EPLING—MONOGRAPH OF MONARDELLA 85

Monardella Elmeri Abrams, Muhlenbergia 8: 43. 1912; Fl.
Los Angeles, ed. 2, 318. 1917; Davidson & Moxley, Fl. South.
Calif. 314. 1923.

Madronella Breweri Greene, Leaflets Bot. Obs. 1: 168. 1906.

Annual, erect, 15-65 cm. tall, branching throughout, the
branches ascending, the lowermost longest, sometimes nearly
equal to the main stem, rebranching, cinereous-puberulent above;
leaves ovate-lanceolate or oblong, 1.5-8.5 em. long, tapering at
both ends, obtuse to slightly acuminate, puberulent, petioles
2-10 mm. long; glomerules 2-3 cm. in diameter, bracts broadly
ovate, little exceeding the calyces, abruptly acuminate to a cusp-
like point, the veins 5-9, arising from the base, subparallel and con-
verging at the tip, the midvein stronger and branching below, the
outermost bracts pinnately veined throughout, all thinly pubescent
or scabrous on the veins; calyx 6-8 mm. long, 14-15-nerved,
scarious below, the teeth acute, slender, unequal, hirsute within;
corolla 12-14 mm. in length, rose-color, the tube retrorsely pu-
berulent, the limb about 5 mm. long, the lips subequal, the lobes
of the upper lip coalesced about two-thirds of the length of the
lip or more, those of the lower lip coalesced almost one-third its
length, tapering somewhat, obtuse, the anther-sacs subparallel,
confluent behind above the connective, but little broader than
the filament; nutlets oblong-oval, 1.5-1.8 mm. long, grayish brown
and mottled.

Specimens examined:

CALIFORNIA: Monterey Co., Nacimiento River, Sept. 19, 1894,
Eastwood (GH); Santa Lucia Mts. (received at GH, July 22,
1898) Eastwood (GH, fragment); Lemmon's ranch, Cholame,
June, 1887, Lemmon 4548 (GH); Acton, June, 1902, Elmer 3681
(GH; MBG; FM; type collection of M. Elmeri Abrams, TYPE in
DH); Corral Hollow, Contra Costa Co., east side of north Coast
Range, east of Mt. Diablo, June 3, 1862, Brewer 1213 (GH,
TYPE; UC; US); edge of Antelope Valley near Neenach, June 6,
1896, Dudley & Lamb 4341 (DH); Hernandez, San Benito Co.,
June 13, 1903, Lathrop (DH); 3 mi. above Acton on Palmdale-
Saugus Road, June 12, 1918, Ferris 949 (DH); Sprague's, Liebre
Mts., Los Angeles Co., June 8, 1896, Dudley & Lamb 4341 (BH);
Lockwood Creek canyon, Mt. Pinos region, Ventura Co., June 24,

[Vor. 12
86 ANNALS OF THE MISSOURI BOTANICAL GARDEN

1896, Dudley & Lamb 4668 (BH); San Antonio River, July,
1880, Vasey 493 (US); San Miguelito Ranch, Santa Lucia Mts.,
June 14-20, 1901, Jepson 1647 (J); near Templeton, July 20,
1913, Abrams 5048 (DH).

While a well-marked species of fairly wide distribution, it has
been little understood and often confused with M. lanceolata.
Beyond the purplish, somewhat more scarious bracts of the Elmer
and similar collections, I can see no essential difference between
M. Breweri and M. Elmeri, certainly none sufficient to warrant
specific distinction. The species may be readily distinguished
from M. lanceolata by the more scarious acuminate bracts.

There are numerous ‘‘Corrals” and numerous ''Hollows"
throughout the state. ‘‘Corral Hollow,” however, is near Tesla
in Alameda County at an elevation of 1000-2000 ft. on the in-
terior side of the coast range.

16. M. Pringlei Gray, Proc. Am. Acad. 19: 96. 1883; Syn.
Fl. М. Am., ed. 2, 2': 459. 1886 (suppl.); Abrams, Muhlenbergia
8: 42. 1912; Fl. Los Angeles, ed. 2, 318. 1917; Davidson &
Moxley, Fl. South. Calif. 314. 1923.

Madronella Pringlei Greene, Leaflets Bot. Obs. 1: 169. 1906.

Annual, erect, 15—40 em. tall, cinereous-puberulent throughout,
occasionally shortly villous near the inflorescence, branching
throughout, the branches ascending, the lowermost longest,
sometimes nearly equal to the main stem, rebranching; leaves
ovate-lanceolate or oblong, 1.5-3.5 ст. long, tapering at both
ends, obtuse or slightly acuminate, pubescent, petioles 2-10 mm.
long; glomerules 2-2.5 cm. broad, bracts ovate, little exceeding
the calyces or equal to them, abruptly acuminate, veins 5-7,
arising from the base, subparallel and converging at the point, the
midvein stronger and branching below, the outmost bracts pinnately
veined, villous with fine trichomes; calyx 6-7 mm. long, 14-15-
nerved, scarious below, pubescent, villous above, the teeth nearly
equal, slender, acute, hirsute within; corolla 11-13 mm. long,
rose-color, the tube puberulent, the limb 3.5-4 mm. long, the
lobes of the upper lip coalesced two-thirds its length or more,
those of the lower coalesced about a third its length, tapering,
stamens exceeding the lips slightly, the anther-sacs divergent at

1925]
EPLING—MONOGRAPH OF MONARDELLA 87

an angle of about 90°, distinct, the connective about three times
the width of the filament, the margin entire; nutlets oblong-oval,
1.2-1.5 mm. long, grayish brown and more or less mottled.

Specimens examined:

CALIFORNIA: Colton, May, 1887, S. B. Parish 1881 (MBG);
locality not stated, 1881, Parry (GH); Colton, May 23, 1882,
Pringle (MBG; US; GH; түре); Colton, June 20, 1905, S. B.
Parish 5398 (GH); vicinity of San Bernardino, May 14, 1895,
S. B. Parish 3653 (GH; MBG; US); sandhills near Colton, June
20, 1907, S. B. Parish 6396 (DH); Declez Pass, Jarupa Hills,
June 20, 1904, Wilder 199 (BH).

The relationship between M. Breweri and M. Pringlei is very
close, and while some forms of one approach forms of the other,
yet one may definitely and readily place them in one category
or the other. The corolla of M. Рите? is constantly smaller
and of a different texture. The anthers, while similar, neverthe-
less show small points of difference, namely, a greater develop-
ment of the connective, which is in general broader than M.
Breweri, appearing more translucent, with the margin distinctly
convex; in the rear, above the connective, the anther-sacs appear
nearly or quite distinet. In addition to these rather obscure
points, the nature of the bract, its smaller size, different texture,
and woolly covering afford a ready means of distinction. While
of similar aspect the entire plant is more slender and of less robust.
habit than M. Breweri. In some collections there is a suggestion
of a scarious margin at the tip of the calyx teeth, the midrib being
prolonged slightly.

17. M. candicans Benth. Pl. Hartweg. 330. 1839; Gray,
Trans. Am. Acad. 11: 102. 1876; Bot. Calif. 1: 594. 1876;
Syn. Fl. М. Am., ed. 2, 2': 358. 1886; Jepson, Fl. West. Middle
Calif., ed. 2, 363. 1911.

Madronella candicans Greene, Leaflets Bot. Obs. 1: 168. 1906.

Annual, erect, 30-40 em. tall, branching from the upper nodes,
forming a corymbose group of inflorescences; branches seldom re-
branching, purplish, puberulent; leaves lanceolate to oblong-
lanceolate, 2-4 cm. long, obtuse, entire, puberulent or nearly
glabrous, narrowed to a slender petiole 0.3-1 em. long; glomerules

[Vor. 12
88 ANNALS OF THE MISSOURI BOTANICAL GARDEN

1.5-2.5 em. broad, subglobose, bracts broadly ovate, obtuse,
scarious, the veins subparallel and green, the secondary veins net-
like, evident, pubescent or puberulent, the margin villous-ciliate;
calyx 5-5.6 mm. long, scarious in the lower half, 13-nerved, sub-
glabrous below, villous above, teeth subequal, obtuse, the margin
narrowly scarious and terminating in an acute white scarious tip,
but not cuspidate, villous inside and out; corolla white, purple-
dotted in some, 10-11 mm. long, the tube but little exserted, re-
trorsely puberulent, the limb 4-5 em. long, the upper lip shorter,
the lobes coalesced for more than half the length of the lip,
those of the lower lip coalesced only at the base, the lobes of both
rather broad, tapering, anthers oblong in outline, divergent at
an angle of about 90°, quite distinct, the connective about three
times the width of the filament, the margin notched; nutlets
about 1.5 mm. long, mottled.

Specimens examined:

CALIFORNIA: Auburn, Plumas Co., 1894, Ames (GH); no
locality given, 1845-47, Fremont’s Exp. (СН); Consumnes River,
1866, Rattan 222 (US; GH); Yosemite, June 15, 1891, Fritchey
80 (MBG); Fresno Creek, Madera Co., June, 1915, Hall 10043
(GH; US; MBG; a very good match for the type); North Fork,
May 30-June 8, 1903, Griffiths 4619 (US; MBG); Volcano,
Amador Co., June 25, 1896, Hansen 1759 (MBG; US); Tollhouse,
Fresno Co., June 13, 1900, Hall & Chandler 26 (MBG; US);
Colfax, July 4, 1882, Jones 3458 (MBG; US); “Mt. Sacramento"
(mountains of Sacramento River), no date, Hartweg (GH;
labeled in Dr. Gray's handwriting “РІ. Hartweg. no. 191 1," type
collection from Herb. Benth.); Knight’s Ferry, Stanislaus Co.,
May 7, 1854, Bigelow, Whipple's Exp. (GH); Calaveras Co., May
18-30, 1895, Davy 1334 (UC); Greenhorn Range, Kern Co.,
June 2-10, 1904, Hall & Babcock 5005 (US; UC); Eldorado Co.,
Sweetwater Cr., June 2, 1908, K. Brandegee (UC); no locality,
Bridges 308 (US); Coloma, Eldorado Co., June 23-24, 1892, E.
Palmer 2373 (US); Mariposa Co., June 15, 1892, Congdon (US).

18. M. exilis (Gray) Greene, Pittonia 5: 86. 1902; Abrams,
Muhlenbergia 8: 43. 1912; Davidson & Moxley, Fl. South.
Calif. 314. 1923.

` 1925]
EPLING—MONOGRAPH OF MONARDELLA 89

Monardella candicans var. exilis Gray, Syn. Fl. N. Am., ed. 2,
21: 358. 1886; Bot. Calif. 2: 476. 1880.

Madronella exilis Greene, Leaflets Bot. Obs. 1: 169. 1906.

Annual, erect, 10-30 cm. tall, cinereous-puberulent, brownish-
purple, branching throughout, the branches flexuous, the lower-
most longest, nearly equalling the main stem in some, rebranch-
ing; leaves oblanceolate to oblong, 1.5-2.5 em. long, obtuse, nar-
rowed to a slender short petiole or sessile, puberulent; glomerules
1.5-2.5 em. broad, bracts ovate, usually surpassing the calyces,
scarious, but with green subparallel veins, with few or no secondary
veins, the margin white-scarious, terminating in a short scarious

abrupt acumination, the margin and back short-pubescent, the entire
bract а bright purple in some; calyx 5-5.5 mm. long, 15-nerved,
scarious in the lower half, sparingly pubescent, the teeth subequal
with a white-scarious margin and tip bristly without, hirsute within;
corolla white, 10 mm. long at most, the tube barely exserted, re-
trorsely pubescent, the limb 2.5-3 mm. long, the upper lip shorter,
the lobes coalesced for more than half its length, those of the lower
lip one-third to one-half its length, lanceolate, obtuse; stamens
shortly exserted, the anther-sacs divergent at an angle of about
60°, the connective about twice the width of the filament, its
margin entire.

Specimens examined:

CALIFORNIA: Lancaster, May, 1909, К. Brandegee (UC);
Palmdale, June, 1902, Elmer 3648 (US; GH; MBG); Mojave
River, June, 1886, S. B. Parish 898c (MBG; US); Walker Pass,
Apr.-Sept. 1897, Purpus 5347 (MBG; US); north fork Kern
River, June 7-15, 1888, E. Palmer 126 (GH; US; it was upon
the sheets of this collection at the National Herbarium that
M. exilis Greene was based); Mojave Desert, about 4000 ft.,
June 14, 1895, S. B. Parish 3784 (GH; US); Mojave River, 1876,
E. Palmer 364 (US; GH; type collection of M. candicans var.
exilis Gray, түре at GH, a fragment at DH); Mojave Station,
June 10, 1906, Hall & Chandler 7382 (US; RMH; BH; UC);
Rabbit Springs, 3000 ft., Apr. 29, 1906, Hall & Chandler 6772
(UC); Mojave River, near Hesperia, May 31, 1892, S. B. Parish
2460 (UC); Mojave River, Burcham’s ranch, May 29, 1901, S. B.
Parish 4909 (DH); vacant lot, Lancaster, Los Angeles Co., Ferris

[Vor. 12
90 ANNALS OF THE MISSOURI BOTANICAL GARDEN

925 (DH); Lancaster, June 4, 1896, Dudley & Lamb 4302 (BH);
Victorville, spring of 1917, Edwards (BH); Little Rock Creek,
sand flat, 3400 ft., Peirson 2416 (BH; J); mouth of Deep Creek,
Mojave Desert, May 19, 1921, Jaeger 1155 (BH); Mojave Desert,
May 17, 1882, Pringle (US).

While M. exilis resembles M. candicans in many ways, the
author is of the opinion that the relationships between М. can-
dicans, M. leucocephala and M. exilis, together with their re-
lation to the other annual species is better shown by its retention
as a species, rather than a subspecies or variety of M. candicans.
It may be distinguished from the latter by the habit of branching,
the smaller corolla and less exserted tube, the number of calyx
veins, and more especial by the nature of the bract, which,
despite the similarity, holds certain differences. These differences,
when learned, permit it to be readily recognized, so that one may
quickly separate a number of specimens, as they appear dried,
by observation of the bract alone, due perhaps to the more opaque
nature of the tissue, the greater crowding of the veins, the fewer
and less conspicuous secondary veins, and the white-scarious
margin and acuminate tip.

19. M. leucocephala Gray, Proc. Am. Acad. 7: 385. 1867;
ibid. 11: 102. 1876; Bot. Calif. 1: 595. 1876; Syn. Fl. N. Am.,
ed. 2, 2': 358. 1886; Jepson, Fl. West. Middle Calif., ed. 2, 363.
1911.

Madronella leucocephala Greene, Leaflets Bot. Obs. 1: 169.
1906.

Annual, erect, 15-20 cm. tall, cinereous-puberulent, branching
throughout, the branches regularly dichotomous, ascending, the
lowermost longest; leaves lanceolate or oblong-lanceolate, obovate
in one specimen, 1-1.5 cm. long, pubescent, the veins scarcely evi-
dent, on petioles 2-3 mm. long; glomerules 1.5 cm. in diameter,
subcorymbose, bracts ovate, orbicular or obovate, with a short
acumination, scarious, pure white, the veins parallel from the base
but not prominent, the cross veins few; calyx 5-6 mm. long, tap-
ering downwards, 1ó5-nerved, hirsute above, the teeth white, at-
tenuate into a spreading and recurved white cusp; corolla white,
5-5.5 mm. long, nearly included within the calyx, the upper lip

1925]
EPLING—MONOGRAPH OF MONARDELLA 91

shorter, incised less than half its length, the lower lip incised
about two-thirds its length, the lobes lanceolate, acute, the
middle lobe of the lower lip somewhat larger than the lateral
lobes; stamens included, those of the upper lip appearing sessile,
the anthers subsagittate, the anther-sacs subparallel, the con-
nective wider than the filament, indented; nutlets oblong-oval,
tapering toward the base, 2 mm. long, apparently only one ma-
turing.

Specimens examined:

CALIFORNIA: plains near Merced ‘‘near the river," June,
1866, Bolander 4845 (GH, турв; US; UC, four sheets); Merced,
June, 1878, Bush (GH); Merced Plains, July 15, 1896, Jepson
100h (J).

M. leucocephala is a very distinct and interesting plant; it is
to be regretted that it is not better known. The matured calyces
fall away, leaving a roughened cruciate receptacle which persists.

INDEX TO SPECIES

' Synonyms are printed in 108; species or genera maintained in this paper
are printed in Roman type; new species, subspecies, or combinations are indica
in bold face type.

Page Page
Macranthae. оао ПОЙ: су PREIS 46
Aigdionsla. -nas En R: o3 ЭМЛАШ o И 79
amab ee ee 50 OS EE CETTE TE EE 55
ПО Еа 72 Я LEER RC ROS ROLLE 90
BEES III и 85 Bol i viu teris 72
ONERE II I EI ICE 87 Е... 35
а АЕ 50 EE AEE A E E ros ev onn 62
бена -S EE AE OS 50 EDL Dae РИ 48
ие: а 60 ES Sr ga re eee ee 69
Dougie ла 78 ПР И eae LE
И: 74 > 52
uL PER ER К 89 BM orci Eran 60
J'OMMBNE, Е es 48 И еги О 72
ИШЛШИ асан OD IS 48 ИМОМ л огоо а 62
iso... eee 62 одогайза та. ................ 59, 60
ОРА Ке РИФИ
Meu: а. 53 АН: 66
o A E o 62 Pani oo GE PS rv XS 41

[Vor. 12

92 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Page Page
с: а c SRL EP LIES 6 lanceolata var. microcephala . 84
DUO... os. oe ak os EU 69 ОМИ 55
рша: u ove 84 Hoo DAE es lui 90
DIMUS sz xv OE eX 68 Basis uss eae ks 71
Pili. ues Е S6 | рр. 72
DUUM Ree ee SG о-у шу» 62 linoides subsp. eulinoides....... 72
а 62 linoides subsp. stricta........... 74
8 о 79 inoides Ver. siridia c... SS 74
СОМЕ. Ке У 62 с oss ec хо. eae 34
ШИЕ о УУ 48 E SEP ESE SCR OEE ET
il [oo SREB Soa ere QUE 44 macrantha subsp. eumacrantha . 35
те ача Rt 76 macrantha subsp. папа... ......
CER ES TE COE осока ое 46 macrantha var arida. ........... 38
о = i ess nee xl ICA 72 macrantha var Hallii............
WHS as es 55 ntha var. longiloba.......... 36
Monarda macrantha var. папа............. 36
ВИНОЙ А ое. 2 macrantha var. pinetorum........ 37
now. iu c cr D Ток» 7 macrantha var. tenuiflora........ 37
WHMIATQODE,. Lv а 32 macrantha var. tenuiflora......... 35
QUI ee ee ee ee er 79 MOBIN LITT ss 62
Meise cio or d ESSE 72 ра: 48
а, аи 70 TRO ies oe LIRE EP 1
ММ e Cue 84 QUEM. Тае 69
о СОО ОО еее 87 WES OES ае ERE CY 36
О E TOS EON СЭУ E 78 WONG WAP. ИТИЙ or a ss eR es 38
candicans var. exilis............. 89 nana var. leptosiphon............ 36
ndicans var. 0епоѕа............ 78 nana var. tenuiflora.......... s. 37
о а rS 2 ИШЕН уе ууз hg Oe 52
OMNI у. у ого Ыы. 56 ПОГРОЗ о CPP Fan V aS 60
а LIA S cv as oe es .. 80 SOMME. eS ees I ede es 72
PEDES I S Du P соле с ыл 77 DIOE о IU SA CHE dos 57
СОМА СООН 50 DUONG CRINGE S E 59
CCE Ss ее 60 odoratissima subsp. australis.... 70
О В зы Mai feet 2, 78 odoratissima subsp. discolor..... 60
ZEE as ea S 85, 107 odoratissima subsp. euodoratis-
ИОНИ В eo 44 io р. 59
epilobioides var. erecta........... 74 odoratissima subsp. glauca...... 62
lo OCUPA UAR, Cr DX соо 88 odoratissima subsp. pallida ...... n
Franciana: А 48 odoratissima subsp. parvifolia. .
GOONER IS eor Re у. 59 wives subsp. pinetorum. . s
мацы еее 0274 | £900. c ee ie
eats OVA лу weed ны 46 uh o o rA
hypolnm. овса 58 PUMA Lois Fee А 41
DUM is с ллы» 62 BENE РОО к 69
ИРОНИ o o a 46 a ack a аа E ng S 69
pn Ice A ee 54 DOMINE a ША 84
lanosollli. ооо АЯ 79 НОГИ Os au Ces LERRA 68
lei var. glandulifera. ...... 84 FUNEM. LI о: 86

1925)

EPLING—MONOGRAPH OF MONARDELLA 93

Page Page

UNNI Ir E G rod OI EE 62 VIEN vu CG Cin НИЯ 44, 48

т SERIO EE BO UV LULA Td 50 villosa subsp. euvillosa......... 46
Vul. О ea 58 villosa var. franciscana.........

Vis ECKE OU NA OT A d Jn 62 villosa subsp. neglecta.......... 52
MEUM. LL S EVA CER E, 79 villosa subsp. Sheltoni.......... 50
XN I Cox A es ad ra s 55 villosa subsp. subserrata........ 48
ORE ПИ TIT Ld 50 villosa var. glabella..............

RON LI LL LE О 48 villosa var. эи ЫЛ pui А 36, 48
UNE и it WM PEGE EEE e onm 72
НИИ NE 48 MA Li a eee ОАА 55
font Nore ЕЕ eink C eT A PYRE о 33, 42
Inque... ee. C624 14 AS 76 И A C ee
undulata var. crispa............ 77 INE ОЗ A Ty 2

[Vor.. 12, 1925]
94 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE

PLATE I

Map of western United States, showing distribution of M. odoratissima.
P —М. odoratissima subsp. pallida
E —M. odoratissima subsp. euodoratissima
D —М. odoratissima subsp. discolor
Po —M. odoratissima subsp. pinetorum
Pv —М. odoratissima subsp. parvifolia
A —М. odoratissima subsp. australis y
G —М. odoratissima subsp. glauca d

Mo. Bor. Garp., Vor. 12, 1925

PLATE 1

PWR ankes

EPLING—MONOGRAPH OF MONARDELLA

96 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE

PLATE 2
Monardella macrantha

Fig. 1. Leaf of var. Hallii, Х 2.
Fig. 2. Leaf of subsp. eumacrantha, x2
Fig. 3. Bract, x 5.
Fig. 4. Calyx of subsp. eumacrantha, X 4.
Fig. 5. Corolla of subsp. eumacrantha, Х 2.
Fig. 6. Limb of var. Hallii, x 2.
Fig. 7. Anthers, x 20.
Fig. 8. QN of var. Seni 9.
Fig. 9. lla of subsp. nana,

Fig. 10. RR of subsp. nana, x 2 (from type).
Fig. 11. Corolla of var. arida, X 2.

All drawings of M. Palmeri are from the type.

[Vor. 12, 1925]

PLATE 2

Mo. Вот. Ganp., Vor. 12, 1925

ANN.

EPLING—MONOGRAPH OF MON ARDELLA

[Vor. 12, 1925
98 ANNALS OF THE MISSOURI BOTANICAL GARDEN

ExPLANATION OF PLATE

PLATE 3
Monardella villosa

Fig. 1. Corolla of subsp. euvillosa, Х 5.

Fig. 2. Bract, X 5 (villosity not shown).

Fig. 3. Corolla of var. franciscana, X 5.

Fig. 4. Anthers, X 25.

Fig. 5. Calyx, X 10.

g. 6. Outer bract, X 5 (villosity not shown),

Fig.
Fig. 7-16. Leaves of various types, pubescence not shown.

Fig. 24. Corolla, x 5

. 26 А
Drawings made from type.

PLATE 3

ANN. Mo. Bor. Garp., Vor. 12, 1925

EPLING—MONOGRAPH OF MONARDELLA

[Vor. 12, 1925]
100 ANNALS OF THE MISSOURI BOTANICAL GARDEN

ExPLANATION OF PLATE

PLATE 4

Monardella hypoleuca

Fig. 1. Calyx, X 10.

Fig. 2. Anther, Х 25

Fig. 3. Corolla, x 5.

Fig. 4. Bract, X 4.

Fig. 5. Leaf, upper surface, X 2.

Fig. 6. Leaf, lower surface, X 2.
Drawings made from type.
M. lanata

Fig. 7. Corolla, X 5.
Fig. 8. Leaf, Х 2.
Fig. 9. Anthers, x 25.
Fig. 10. Calyx, x 10.
Fig. 11. Leaf, lower surface, X 2.
Fig. 12. Leaf, lower surface, X 2 (villosity not shown).
Fig. 13. Leaf, upper surface, X 2 (villosity not shown).
Fig. 14. Bract, x 5.
Drawings made from type collection.
М. saxicola

Fig. 15. Bract, x 5.
. 16. Leaf, upper surface, X 2.
Fig. 17. Leaf, lower surface, X 2
Figs. 18-19. Leaves, upper ө. % 3.

Figs. 24-96. Leaf, Wpber surface, T not shown in last two, Х 2.
Fig. 27. Leaf, lower surf: x4

Drawings made from type collection.

PLATE 4

ANN. Mo. Bor. Ganp., Vor. 12, 1925

TN 7

ФАТ 0 eer
Е АЕ var

EPLING—MONOGRAPH OF MONARDELLA

[V or. 12. 1925]

102 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 5

M. odoratissima

Fig. 14,

The corollas чи above are those whi

indicated.
inoides

Calyx, Х 5 (subsp. discolor).
Anthers, Х 50.
Nutlets, X 50.
Trichomes common to the genus.
Bracts, X 5.
Corolla of subsp. discolor, X 5
Gynobase after nutlets have fallen, х 50.
Corolla of subsp. euodoratissima, Х 5.
Tip of style, x 50.
Corolla of subsp. glauca, Х 5
Types of foliage, X 2 (pubescence not shown).
Corolla of subsp. pallida, х 5
Corolla of subsp. some x 5.
Corolla of subsp. australis, Х 5.

All gradations may be found.
Bract,

eel of їй, X 2 (pubescence not shown).
Corolla, x 5.

ch are common in the subspecies

ANN. Mo. Вот. Garp:, Vor. 12, 1925

PLATE 5

EPLING—MONOGRAPH OF MONARDELLA

[У от. 12, 1925]

104 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE

PLATE 6

M. undulata

Fig. 1. Cal х 10.

Fig. 2. P. ха ия not shown).

Fig. 3. Anther, X 2

Fig. 4. Bracts, X 5 I second outermost).

Fig. 5. Corolla, x

Fig. 6 I x ^
M. D

Fig. 7. Dad X 5 (the second innermost).
Fig. 8. Calyx, Х 10.

Fig. 17. Nutlets, x 10.
M. Breweri
Fig. 18. Bract, Х 5.

Drawings made from бур:

ANN. Mo. Bor. Ganp., Vor. 12, 1925 Р 6
LATE

EPLING—MONOGRAPH OF MONARDELLA

[Vor. 12, 1925]

106 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 7

liag
Anthers, X 25.
Fig. 6. Bract, x 5.

Fig. 7. Corolla, X 5.

Fig. 8. Calyx, x 10.

Fig. 9. Anthers, Х 25.

Fig. 10. Leaf, X 2

Fig. 1. Nutlets, x 10 (? mature).
Fig. 12. Bract, X 5

Drawings ida Pos type.

M. Pringlei
Fig. 13. Bract, X 5

Fig. 17. Leaf, X
Drawings made from type collection,

EPLING—MONOGRAPH OF MONARDELLA

STUDIES ON SOUTH AMERICAN LABIATAE. I

SYNOPSES OF THE GENERA TEUcRIUM, Rosmarinus, MAR-
RUBIUM, PRUNELLA, LAMIUM, LEONURUS, AND
LEONOTIS

CARL CLAWSON EPLING
Instructor in Botany, University of California, Southern Branch
Formerly Rufus J. Lackland doc, rch Fellow in the Henry Shaw School of Botany of
Washington University
FOREWORD

The botanical exploration of the past decade which has had
South America as its field has resulted in the accumulation
among other things of a considerable quantity of specimens of
the Labiatae of that region. These collections, as far as this
family of plants is concerned, have been critically studied in
but very few instances, and for the most part the specimens
have been determined only to the genus. The same is true of
the collections of earlier date as they are represented in the
larger herbaria of the United States. It has been the author’s
privilege to have placed at his disposal for study the collections
of South American Labiatae in the herbaria of the Missouri
Botanical Garden, the United States National Herbarium, the
Field Museum of Chicago, the Academy of Natural Sciences of
Philadelphia, the Gray Herbarium of Harvard University,
the University of California, and the New York Botanical Gar-
Чеп. The names of the principal collectors whose collections
are represented in these herbaria, together with the country
in which the collections were made and the year, are listed below.

J. Dombey Peru about 1780

H. B. Fielding Chili

C. F. Ph. von Martius Brazil 1817-1821
A. Bonpland Colombia

1 The following abbreviations are used herein: ASP, Academy of Natural Sciences
of Philadelphia; FM, Field Museum of Natural History; GH, Gray Herbarium;
MBG, Missouri Botanical Garden; NY, New York Botanical Garden; UC, University

of California; US, United States National Herbarium.
Issued September 28, 1925.

Ann. Mo. Bor. Garp., Vor. 12, 1925 (107)

108

Fr. Sello (Sellow)
C. H. Mertens

C. J. Bertero

Cl. Gay

W. Jameson

J. Tweedie

R. H. Schomburgk
G. Gardner
Wilkes Exped.
Commerson

H. F. A. von Eggers

Styles

T. Hartweg
Glaziou

P. Salzmann
J. Gillies

N. J. Anderson

J. Ball

H. H. Rusby &
Roy W. Squires
O. Buchtien

British Guiana
Brazil
Argentina-Chili

[Vor. 12

ANNALS OF THE MISSOURI BOTANICAL GARDEN

1819

(?)1827

1827
1828-1834

1838-1840
1838-1842

Uruguay and Argentina 1840

Ecuador

Argentina-Chili
Galapagos
Argentina
Venezuela
Ecuador

Chili

Bolivia

Brazil

Argentina
Venezuela
Argentina
Chili

Colombia-Ecuador

Uruguay
Chili
Paraguay
Bolivia
Galapagos
Argentina

Venezuela
Chili

1840

1841-43

(?)1844

1851

1854-1855
1855-1859
1856
1860
1866

1872
1874
1876
1882
1881-1883
1886
1887
1888-1890
1891-1892
1891
1891-1892

1896
1895-1896,
1903

1925]

EPLING—SOUTH AMERICAN LABIATAE. I

109

A. H. Moore Venezuela 1899
G. T. Hastings Chili 1900
K. Fiebrig Paraguay 1902
H. Pittier Colombia 1905-1906
A. Stewart Galapagos 1906
O. Buchtien Bolivia 1906-1919
K. Fiebrig Paraguay 1909-1910
A. Jahn Venezuela 1910
H. Pittier Venezuela 1913-1921
E. Hassler Paraguay 1913
P. Dusén Brazil 1914
Bro. Ariste-Joseph Colombia | 1914
O. F. Cook &
G. B. Gilbert Peru 1915
W. Fischer Argentina 1915
A. Weberbauer Peru 1915
P. Jórgensen Argentina 1915-1917
H. H. Rusby &
F. W. Pennell Colombia 1917
F. W. Pennell Colombia 1917-1918
J. N. Rose Ecuador 1918
Bro. Claude-Joseph Chili 1918-1923
A. S. Hitchcock British Guiana 1919
E. W. D. & M. M. Holway Chili 1919
A. S. & M. Kalenborn Peru 1919
H. A. Gleason British Guiana 1921
J. F. Macbride &
Featherstone Peru 1922
F. W. Pennell Colombia 1922
F. W. Pennell & E
E. P. Killip Colombia 1922
E. P. Killip &
Bro. Ariste-Joseph Colombia 1922
Bro. Ariste-Joseph Colombia 1922
E. P. Killip Colombia 1922
F. W. Pennell, E. P. Killip
& T. Hazen Colombia 1922
Venezuela 1923

W. E. Broadway

[Vor. 12

110 ANNALS OF THE MISSOURI BOTANICAL GARDEN

J. S. De La Cruz British Guiana 1920-23
G. H. Pring Colombia 1923
F. M. Macbride Peru 1923
H. E. Anthony &

G. H. Tate Ecuador 1923
A. S. Hitchcock Ecuador _ 1923
E. Werderman -Chili 1924

The present paper offers a synopsis of the species belonging
to the genera named in the title, together with their distribution
as represented by the above-named collections. With the ex-
ception of Teucrium, the genera are monotypic for this region,
and of these monotypic genera all but Prunella are introduced
and naturalized, some growing as wayside weeds. Prunella
vulgaris is cosmopolitan. Teucrium, however, is represented in
South America by approximately five species.

While recent years have added very greatly to the exsiccatae
available for study, nevertheless the fact remains that numerous
species are represented in the above-named collections by a
single specimen or not at all. Furthermore, the majority of
types and historical collections are to be found in European
herbaria. These types have not yet been studied by the author.
By reason of these facts, then, the proposed papers, of which
‚ this is the first, are not monographie in nature but in many
cases include tentative dispositions which may or may not be
changed upon accession of additional specimens or after study
of certain types. Accordingly, for the present, where doubt
exists, changes in nomenclature, which may eventually be neces-
sary, will be only indicated.

In spite of the fact that these notes are incomplete their
publieation is thought desirable by reason of their possible
bearing first upon field study and secondly upon herbarium
study of the collections already made. In the first instance it
is hoped that the descriptions and keys will permit of identi-
fication and study in the field and will stimulate the preservation
of variant forms and the collection of seed for garden study.
In the second instance it is hoped that the determination of
unnamed herbarium specimens will be greatly facilitated.

The work outlined in the following notes was done largely at

1925]
EPLING—SOUTH AMERICAN LABIATAE. I 111

the Missouri Botanical Garden during the summer of 1924.
The author is accordingly greatly indebted to the Director of
that institution for the excellent facilities afforded. He is no
less obligated to the directors of the various herbaria listed
above who have so generously loaned very valuable material.

TEUCRIUM (Tourn.) L.

Herbae fruticesve floribus in his speciebus ad ramorum ex-
tremitates sitis vel racemoso-spicatis vel patento-racemosis; ver-
ticillastris densis, 2—6-Йог из, bracteis differentibus; floribus
in axillis bracteorum foliosorum solitariis; calycibus tubulosis
vel campanulatis frequenter leviter inflatis, 10-venis, quinque-
dentatis, dentibus subaequalibus vel posticis latioribus; corollae
tubo brevissimo in calyce incluso, intus inannulato, frequenter
fauce paulo constricto, limbo pro rata longissimo, labro inciso,
in ealyce incluso vel paulo exserto, eius lobis labioli laterales
subaequantibus, labiolo trifido, lobo medio ovato, prominenter
declinato-patente, lateralibus oblongis minoribus; staminibus
quatuor, didymis, anticis longioribus, omnibus e labri sinu
exsertis, declinato-arcuatis; antheris rotundatis, thecis conflu-
entibus; stylo subaequaliter bifido; nuculis hilo prominentiore et
laterali, rugoso-reticulatis, maturis cohaerentibus.

CoNsPECTUS SPECIERUM
A. Flores in axillis bracteorum foliosorum solitarii.

а. Folia caulinaria subintegra, floralia saepius trilobata ........... 3. T. bicolor
b. Folia caulinaria 3-5-fida vel -lobata, floralia trifida.

a. Calycis lobi tubo aequilongi......... 60008068 2. T. nudicaule
B. Calycis lobi tubo duplo longiore............-- ey SUE ya PA зе

B. Flores in racemosis spicis extremis dispositi.
a. Folia obtusa; corolla fauce lata 3-3.5 mm........ n 6. T. tenuipes

b. Folia acuta; corolla fauce lata 2-2.5 mm.
а. Planta breviter et dense pubescens; corolla longa 9-12 mm. . 4. T. inflatum
B. Planta pilosa; corolla longa 10-15 mm........«- 5. T. palustre

1. T. cubense Jacq. Enum. Syst. Pl. Carib. 25. 1760, et
Select. Stirp. Amer. Hist. 1: 172, pl. 63, fig. 74. 1763.
Herba gracilis altitudine 20-50 em., in basi ramosa, ramis
adscendentibus, virgatis vel iterum ramosis, glabris vel puberulis,
quadratis, saepe canaliculatis, angulis acutis; foliis obovatis vel

[Vor. 12
112 ANNALS OF THE MISSOURI BOTANICAL GARDEN

oblanceolatis, 1—4 cm. longis, glabratis, obtusis, basi cuneatis et
in petiolam brevem coarctatis, eaulinariis maxime diversitatis,
vel in lobos tres obscure incisis, segmento medio majore, omnibus
saepius crenato-dentatis, dentibus 8-10, obtusissimis, vel in
segmenta linearia tria quinqueve partitis, floralibus saepius
3-fidis, lobis linearibus, subaequalibus, frequenter tamen foliis
eaulinariis similibus; floribus oppositis, solitariis, decussatim
instructis; calycibus ‘campanulatis, florentibus 5-8 mm. longis,
dentibus anguste lanceolatis, acutis, tubo duplo longioribus,
pedicellis 4-5 mm. longis; corollis 8-10 mm. longis, tubo 1.5-2
mm. longo, lobis labri oblongis, 2-2.5 mm., obtusis, labiolo
6-7 mm. longo, lobis lateralibus lobos labri subaequantibus,
lobo medio obovato, 6 mm. longo, in basi angustato; staminibus
ad faucem sitis, 7-9 mm. longis, stylo paulo longiore; nuculis
2.5-3.5 mm. longis, valde rugoso-reticulatis, fuscis, maturis
calycis tubum superantibus, hilo 1.8-2.3 mm. longo.

CoNsPECTUS SUBSPECIERUM

Planta glabra; calycis tubus maturus non auctus.......... subsp. Chamaedrifolium
Planta ge destin tubus maturus patenter auctus, forma hemisphaerii,
diimetro 4.5 ТА Se oEVIQ ee Ea subsp. cordobense

a. Subsp. Chamaedrifolium (Mill. comb. nov.

T. Chamaedrifolium Mill. Gard. Dict., ed. 8, n. 16. 1768.

T. laevigatum Vahl, Symb. 1: 40. 1790 (non Solander in
Russ. Nat. Hist. Aleppo, ed. 2, 2: 255. 1794).

Planta glabra, foliis maximam partem in lobos tres incisis,
segmento medio majore, crenato-dentatis, dentibus 8-10, ob-
tusissimis; calycibus florentibus 5-6 mm. longis, fructiferibus
7 mm. к, lobis acutis, in basi 1 mm. latis, tubo 2 mm. longo,
vix aucto; corolla caerulea vel albida.

The plant which occurs at Buenos Aires is apparently the
same as that which is distributed throughout the West Indies
and along the shores of the Gulf of Mexico from Florida to
Yucatan, the type locality of which is Cuba. The leaves may
vary greatly upon an individual or may be nearly of a kind.
At one extreme they may be merely dentate-crenate or crenately
8-10-lobed, the lobes being subequal and very blunt; at the
other extreme they may be irregularly incised into three prin-

1925]
EPLING—SOUTH AMERICAN LABIATAE. I 113

cipal obovate or oblanceolate segments, of which the middle one
is clearly the largest and usually crenately lobed though fre-
quently entire. The leaf subtending the lowest part of the in-
florescence is but rarely parted into linear segments. The floral
leaves, however, are for the most part three-parted into linear
segments but may occasionally be merely toothed.

Specimens examined:

ARGENTINA: Buenos Aires, 1837, Tweedie (NY).

b. Subsp. cordobense subsp. nov.

T. Grisebachii Hieron., nom. nudum in Gris. Symb. ad Floram
Argent. 275. 1879.

T. cubense Grisebach, 1. с. 275. 1879.

Planta puberula, cinerea, foliis maximam partem in segmenta
linearia tria quinqueve partitis; calycibus florentibus 5-7 mm. -
longis, fructiferibus 7-9 mm. longis, lobis obtusiusculis, in basi
1-1.2 mm. latis, tubo 2.5-3 mm. longo, patenter nuculis distento,
forma hemisphaerii, diametro 4-5 mm., dentibus nonnihil
auctis, super nuculos subconniventibus sinibus rotundatis;
corolla ‘‘luride rubescentibus" (Hieron.).

Specimens examined:

ARGENTINA: Sierra de Cordoba, Feb. 14, 1876, Hieronymus
390 (FM; US); El Sancho, Catamarca, 1700 m., Nov. 12, 1916,
Jörgensen 1296 (US), Мау 22, 1915, Jörgensen 1296 (MBG),
May 12, 1915, Jérgensen 1296 (GH); Ischilin, 1892, Kuntze
(US, түрк; NY); Cordoba, en las Quintas, Oct. 1883, Galander
(NY); Potrero de Lujan, Cordoba, Dec. 25, 1883, Galander (NY);
Cordoba, Dec. 1891, Kuntze (NY); Cordoba, Altos del Observa-
toro, Nov.-Dec. 1891, Kurtz 7260 (NY).

A note on the label of the Hieronymus collection at the Field
Museum reads “Аш Fuss der Cuesta de las Chacras zw. Devi-
saderos u. Taninga; Westseite der Sierra de Cordoba."

2. Т. nudicaule Hooker, Bot. Miscellany 2: 235. 1831.
? T. tripartitem Meyen, Reise 1:406. 1843 -
Suffrutex, altitudine 20-(?)40 cm., ramis numerosis e corona

lignosa, adscendentibus, virgatis, puberulis, quadratis, angulis

obtusis, internodiis ramorum fertilium quam foliis duplo triplove
longioribus, eis in ramis sterilium 2 cm. longis, in segmentis

[Vor. 12
114 ANNALS OF THE MISSOBUI ROTANICAL GARDEN

linearibus acutis profunde tripartitis, segmento medio paulo
longiore, eis in ramis fertilium 7-15 mm. longis, vel ad basim
vel ad medium tripartitis, omnibus puberulis; floribus oppositis,
decussatim instructis, in racemis extremis dispositis, racema
virgata vel ramosis, ramis distantibus, gracilibus, divaricatis;
bracteis foliosis; calycibus 3-5 mm. longis, campanulatis,
puberulis, lobis tubo aequilongis, lanceolatis, subacuminatis,
fructiferibus vix auctis, pedicellis 4-5 mm. longis elatis; corollis
12-15 mm. longis, extus pubescentibus, tubo 2-3 mm. longo,
ad bases staminum pubescente, lobis labri oblongo-ovatis 7
mm., labiolo 10-13 mm. longo, lobis lateralibus oblongis 5 mm.,
lobo medio oblongo-rotundato, 8-10 mm. longo, staminibus ad
faucem sitis, 12 mm. longis, ad basim hirtellis; stylo paulo
longiore; nueulis oblongis, 3 mm. longis, 1.2 mm. latis, atris,
asperulis, hilo nigro, parte dimidia quam nucula breviore.

The middle lobe of the lower lip is apparently of a deep
purplish eolor, the remainder of the corolla being lighter in shade
as in T. bicolor. The species is reported by Gay from Copiapo,
Coquimbo, and Arqueros. The type locality is uncertain.

Specimens examined:

Снил: Desert of Atacama, 1885-7, Geisse 71 (NY); no data,
Gay (GH).

Var. leucanthum (Phillipi), comb. nov.

T. leucanthum Phillipi, An. Univ. Chili 90: 565. 1895.

Planta 20 cm. alta, foliis 8-10 mm. longis, confertissimis, cor-
olla alba (Phillipi).

Not improbably a drought form of T. nudicaule. Phillipi
gives the following note: ‘‘ Las hojas son casi sesiles, las mas veces
mucho mas pequenas que en el 7. nudicaule i apenas partidas
mas alla del medio i las lacinias esteriores a menudo provistas
de un lobulo de modo que las hojas parencen casi quinquefidas;
las flores son axilares i solitarias i cortamente pediceladas. En
otros ejemplares las hojas son mas profundamente partidas, i
sus divisiones provistas de dos a cuatro lobulitos de cada
lado." His type is a collection made by Bouchers near Taltal
in October, 1887.

The label of the Ball collection at the Gray Herbarium bears

1925]
EPLING—SOUTH AMERICAN LABIATAE. I 115

the following annotations: ‘“‘ramis dense foliaceis floribus ap-
proximatis," ‘‘only three phanerogams were seen on the coast
at that place."

Specimens examined:

Снил: ex rupestribus chilensibus juxta Taltal, May, 1882,
Ball (US; GH; NY).

3. T. bicolor Smith in Rees, Cyclopedia 35: no. 25. 1819.

T. heterophyllum Cav. leones 6: 56. pl. 577. 1801 (non
T. heterophyllum L'Herit. Stirp. Nov. 84. fig. 49. 1784-85).

T. orchideum Lindl. Bot. Reg. 15: pl. 1255. 1829.

T. Cavenellesii Bert. ex Steud. Nom., ed. 2, 2: 674. 1841
(nom. nudum).

Т. chilense Desf. ex Steud. Nom., ed. 2, 2: 674. 1841 (nom.
nudum).

Frutex 2 m. altitudine, multo ramosis, foliosis, ramis tere-
tibus, cortice discedente, ramulis gracilibus, puberulis, quad-
ratis, angulis acutis, marginatis, internodiis plerumque foliis
subaequalibus vel brevioribus; foliis forma diversis, oblan-
ceolatis 1.5-2.5 em. longis, trivenis, obtusis, in basi ad brevem
petiolam angustatis, puberulis, margine revoluta, integra vel
diverse dentata vel lobata, maximam partem dentibus duobus
ad medium, his 1-2 mm. longis, ovatis, obtusis, divergentibus;
floribus in racemis extremis in axillis foliorum solitariis, de-
cussatim instructis; calycibus 5-7 mm. longis, lobis tubo aequi-
longis, lanceolatis, acutis, fructiferibus vix auctis, pedicellis 4—5
mm. longis elatis; corollis 18-20 mm. longis, extus pubescentibus,
tubo 3-4 mm. longo, ad bases staminum hirsuto, labri lobis
oblongis, obtusis, 5-6 mm., labiolo 12 mm. longo, lobis lateralibus
ovatis, majoribus quam labri lobis, 5-6 mm. longis, lobo medio
10 mm. longo, oblongo-rotundato, in basi subtruncato, purpureo,
corolla alia alba; staminibus ad faucem sitis, 15-18 mm. longis,
ad basim valde hirsutis, valde arcuatis; stylo 20 mm. longo,
ramis gracilibus, 2 mm. longis, nuculis obovatis, 2 mm. longis,
asperulis, fuscis, hilo 1.5 mm. longo.

T. bicolor and T. nudicaule are closely allied and appear to
have been derived from a common stock, the latter representing
an adaptation to a more arid habitat. Their distribution, as
far as known, supports this assumption.

[Vor. 12
116 ANNALS OF THE MISSOURI BOTANICAL GARDEN

T. bicolor is reported by Gay as common in the hills of the
central provinces.

Specimens examined:

Снил: Prov. de Chillon, Dec. 1869, ? Couthoy (FM); Ramon,
Nov. 25, 1920, Bro. Claude-Joseph 1275 (US); Styles (ASP);
Santiago, San Cristobel, Nov. 3, 1900, Hastings 119 (US); Gay
293 (US); Lota, Nov. 7, 1868, Cunningham (GH); Valparaiso,
Mertens (GH); Coquimbo, July-Aug. 1856, Harvey (GH); ex
regione inferiore Andium Chilensium in Convalle Aconcagua,
May 1882, Ball (GH; NY); Gay (GH); Bertero 689 (GH); Chili
boreale, 1827, Bertero 1352 (MBG); Valparaiso, Wilkes Exp.
(US; NY); Costa, Nov. 4, 1920, Bro. Claude-Joseph 1254 (US);
Isle St. Marys, Eights (US); Temuco, Dec. 5, 1919, Holway 200
(US); in the bush near Salto, Valparaiso, Sept. 21, 1895, Buchtien
(US); Prov. Colchagua, collector unknown 158 (NY); no data, Gay
293 (NY); Valparaiso, Gillies (NY).

4. Т. inflatum Swartz, Prod. Veg. Ind. Осе. 88. 1797.

(?) T. vesicarium Miller, Gard. Dict., ed. 8, sub Teucrio,
по. 17. 1768.

Herba, caule erecto e rhizomate repente, frequenter in nodis
inferioribus radicante, altitudine 40-90 cm., ramosis in axillis
superioribus, quadratis, canaliculatis, angulis obtusis, dense et
breviter pubescentibus; foliis ovatis vel lanceolatis, 4-12 cm.
longis, vix acuminatis, in basi truncato-cuneatis, margine sub-
duplicato-serrata, erenis 2-3 mm. longis, 1.5-2.5 mm. latis,
ovatis, subapiculatis, pagina superiore hirtellis, inferiore pal-
сев dense tomentellis, petiolis gracilibus, pubescentibus,
1-3 em. longis floribus in spicis extremis racemosis 5-20 cm.
longis, 1.5 cm. latis congestis, subspirale sed tamen 2-4 floribus
in pseudoverticillastris instructis; bracteis lanceolato-linearibus,
acuminatis, pubescentibus, pedicellos excedentibus; calycibus
campanulato-tubulosis, cano-pubescentibus, bilabiatis, 5-7 mm.
longis, inflatis, ore obliquo, dentibus 2 mm. longis, posticis in
` basi 1.2 mm. latis, triangulo-ovatis, acutis, anticis angustioribus,
acuminatis, omnibus conniventibus; corollis 9-11.5 mm. longis,
tubo 4-5 mm. longo, lobis labri triangulis, 1 mm. longis, acutis
sinu labri 2-3 mm. alto, labiolo 5 mm. longo, extus pubescenti-

1925]
EPLING—SOUTH AMERICAN LABIATAE. I 117

bus, lobis lateralibus 1 mm. longis, oblongis, obtusis, lobo
medio 3 mm. longo, rotundato-obovato; staminibus 7 mm.
longis, ad medium tubi sitis, infra medium hirtellis; stylo
staminibus paulo breviore; nuculis 2-2.5 mm. longis, fuscis,
valde rugoso-reticulatis, glabris, hilo 1.7-2 mm. longo.

The type locality of T. inflatum is Jamaica. The plant is
described as “calycibus inflatis, villosis." The type locality of
T. vesicarium is near Vera Cruz, Mexico. Miller's description
was drawn from plants grown in England from seed introduced
by Houston. The plant is described as “Calice vesicario"
and with *smooth branches." The type locality of Т. palustre
is cited as being between the mouth of the Sinu River and Car-
tagena in Colombia. The plants collected in this locality by
Pennell (which are distinctly pilose) correspond very closely to
Kunth’s description.

The plants of Jamaica and the West Indies are of the pu-
bescent type and are similar in every way to the plants growing
in Brazil, Paraguay, Ecuador and Argentina. On the other
hand, all specimens from Mexico which have been examined
were definitely pilose and were similar in every way to the
plants of Colombia.

Careful measurements of numerous flowers (after boiling) have
shown that the Mexican plants in addition to the pilose covering
have corollas which are uniformly larger, ranging in length from
10-16 mm. The calyces, too, are somewhat larger (6-9 mm.)
and the teeth are usually more acute. The corollas of the
West Indian plants and the South American plants other than
Colombian are 9-11.5 mm. long with calyces 4.5-7 mm. long.
The two groups do intergrade, however, and since their geo-
graphical distribution is distinct they are better considered
subspecies. Until it is possible to ascertain what plants were
actually described by Swartz, Miller, Kunth, and Sprengel, no
changes in nomenclature have been made.

Specimens examined:

Braziu: Prov. Sta. Catharina, Tubarao, Jan. 1889, Ule 1060
(US).

Ecuapor: Prov. Guayas, Milagro, 50 m., June 30-July 2,
1923, Hitchcock 20210 (US; NY); near Guayaquil, Hartweg 684
(NY); no data, Eggers (NY).

[Vor. 12
118 ANNALS OF THE MISSOURI BOTANICAL GARDEN

PARAGUAY: in region of Alto Parana R., 1909-10, Fiebrig
5604 (US; GH); in region of Lake Ypacaray, March, 1913,
Hassler 11462 (US; MBG; GH); Asuncion, 1888-90, Morong 179
(US; MBG; ASP; GH; NY).

ABGENTUNA: El Monte las ? Palan, Nov. 1917, Jórgensen
2239 (US; MBG; GH).

5. T. palustre Kunth in Humboldt, Bonpland & Kunth,
Nov. Gen. et Sp. Pl. 2: 306. 1817 (non T. palustre Lam. Fl.
Fr. 2: 441. 1778).

(?) T. hirtum Willd. ex Spreng. Syst. 2: 710. 1825.

Herba, caule erecto e rhizomate repente, frequenter in nodis
inferioribus radicante, altitudine 40-90 cm., ramosis in axillis
superioribus, quadratis, canaliculatis, iratis obtusis, pilis
patentibus vestitis; foliis ovatis vel lanceolatis, 4-12 cm. longis,
vix acuminatis, in basi truncato-cuneatis, margine subdupli-
cato-serrata, crenis 2-3 mm. longis, 1.5-2.5 mm. latis, ovatis,
subapieulatis, pagina superiore tenuiter pilosa, pilis 1 mm.
longis, inferiore pallidiore praecipue ad venos dense pilosa,
petiolis gracilibus pilosis 1-3 cm. longis; floribus in spicis extremis
racemosis, 5-20 cm. longis, 1.5 ст. latis congestis, subspirale
sed tamen 2-4-floribus in pseudoverticillastris instructis, brac-
teis lanceolatis-linearibus, pilosis, acuminatis, calyces paulo ex-
cedentibus; calycibus campanulato-tubulosis, inflatis, breviter
pubescentibus, pilis quoque 1 mm. longis ornatis, bilabiatis,
6-8 mm. longis, ore obliquo, dentibus 2 mm. longis, posticis in
basi 1.2 mm. latis, triangulo-ovatis, acutis, anticis angustioribus,
acuminatis, omnibus conniventibus; corollis 11-16 mm. longis,
pallide rubro-violaceis vel albis (Pennell), tubo 5-7.5 mm.
longo, fauce 2-2.5 mm. lato, lobis labri 1-1.5 mm. longis,
acutis, sinu labri 2.5 mm. alto, labiolo 5 mm. longo, extus pu-
bescentibus, lobis lateralibus 1.5 mm. longis, oblongis, obtusis,
lobo medio 3.5-4 mm. longo, rotundato-obovato; staminibus
8 mm. longis, ad tubi medium sitis, infra medium hirtellis;
stylo staminibus paulo brevioribus; nueulis 2.5 mm. longis,
fuscis, valde rugoso-reticulatis, glabris, hilo 2 mm. longo.

For a discussion of nomenclature see the note to T. inflatum.

Specimens examined:

1925]
EPLING— SOUTH AMERICAN LABIATAE. I 119

CoLoMB1iA: Dept. of Bolivar, Cienago de Oro, 50-100 m.,
meadow, corolla light pink-violet, Jan. 28, 1918, Pennell 4124
(US; GH); Dept. of Bolivar, tierra alta on Rio Sinu, March
7-10, 1918, meadow along river, 50-80 m., corolla pink-violet,
Pennell 4619 (US; MBG; FM; NY; type locality of T. palustre
Kunth).

6. T.tenuipes sp. nov.

Herba altitudine 20-40 cm., e rhizomate repente, caule
gracile, diametro 2 mm., pubescente, quadrato, angulis obtusis,
nodiis inferioribus nudis, in nodis superioribus ramosis, ramis
subdichotomis; foliis in nodis superioribus confertis, obtusis,
2-6 ст. longis, in basi truncato-cordatis, margine convexo, ir-
regulariter serrato-crenatis, crenarum culminibus 1.5-4 mm.
altis, obtusis, apiculatis, inter se 3-6 mm. distantibus, pagina
superiore molliter pubescente, inferiore pallidiore, pubescente
vel subtomentoso, petiolis gracilibus, 1-3.5 cm. longis; floribus
in spicis racemosis extremis, subspirale sed tamen 2-4-floribus
in pseudoverticillastris instructis, frequenter oppositis; bracteis
lanceolato-linearibus, acuminatis, villosis, pedicellos paulo ex-
cedentibus; calycibus 5.5-7 mm. longis, campanulato-tubulosis,
subvillosis, bilabiatis, paulo inflatis, ore obliquo, dentibus 2-2.5
mm. longis, posticis in basi 1.5-2 mm., triangulo-ovatis, acutis,
antieis angustioribus, fructiferibus vix conniventibus; corollis
11-14 mm. longis, tubo 4 mm. longo, fauce 3-4 mm. lato, lobis
labri 1.5-2 mm. longis, sinu labri 3-4 mm., labiolo 5-7 mm.
longo, lobis lateralibus 2 mm., oblongis, obtusis, lobo medio
3.5-4 mm. longo, rotundato-obovato; staminibus 10-11 mm.
longis, ad tubi medium sitis; stylo staminibus subaequale;
nueulis 2.5 mm. longis, fuscis, reticulato-rugosis, hilo 2 mm.
longo.

'The plants above described have previously been referred to
T. inflatum Sw. The convex obtuse leaves, with relatively
coarser teeth, the differences in habit and in pubescence, and
_ particularly the differences in the proportions of the corolla,
which more nearly resembles that of T. occidentale, have
seemed sufficient reasons for considering them as of a distinct
species.

(Vor. 12
120 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Specimens examined:

GaLAPAGOS Isr.: Charles Island, 1852, Anderson (GH);
Charles Isl., ‘соттоп among rocks," 446 m., Feb. 27, 1906,
Stewart 3342 (US, TYPE; MBG; GH); Charles Isl, 446 m.,
Oct. 9, 1905, Stewart 3343 (GH); Chatham Isl., Wreck Bay,
200 m., Jan. 27, 1906, Stewart 3345 (US; MBG; ASP; FM; GH);
Chatham Isl. Wreck Bay, ‘соттоп in open places," 200 m.,
Jan. 27, 1906, Stewart 3345 (GH); same, Feb. 23, 1906, Stewart
3344 (GH); Chatham Isl., southwest end, middle region,
June, 1891, Baur 164 (GH).

T. SconopoNiA Г. has been reported by Phillipi' with the
following note: “Es planta comun en la mayor parte de Ale-
mania, que parce haber llagado a Chili con semillas de forage.
El tallo alcanza a 65 cm. de alto, las hojas suelon tener 27-50
mm. de largo i son mui arrugadas. Los racimos son alargados
i flojos, multifloros i las flores amarillentas. No tiene semejanza
con ninguna otra planta chilena."

Rosmarinus (Tourn.) L.

Frutex floribus in racemis lateralibus brevibus dispositis,
calycibus ovatis-campanulatis, bilabiatis, labio superiore integro,
inferiore bifido; corollae tubo exserto, intus inannulato, fauce
dilato, limbo bilabiato, labiis subaequalibus, labro erecto,
emarginato, labiolo patente trifido, lobo medio maximo, con-
cavo, dependente; staminibus duobus (inferioribus) infra me-
dium eum dente parvo ornatis, arcuatis, e labro exsertis, ad
faucem sitis; stylo staminibus aequilongo, ramo superiore
brevissimo; uvali siccis, laevibus.

1. R. officinalis L. Sp. Pl. 23. 1753.

R. chilensis Molina, Saggio sulla Storia Nat. de Chili 158.
1782.

Frutex dense ramosus foliosusque altitudine circa 1 m.,
ramis teretibus, lignosis, glabratis, cortice discedente; foliis
linearibus, 1-3 em. longis, acutiusculis, mucronatis, sessilibus,
glabratis, subtus canescentibus, margine integra, valde re-
voluta; floribus paucis in racemis in ramulis lateralibus, 2-3

! An. Univ. Chili 90: 565. 1895.

1925]
EPLING—SOUTH AMERICAN LABIATAE. I 121

ст. longis dispositis, oppositis, in axillis bracteorum solitariis;
bracteis membranaceis, ovatis, pedicellis subaequalibus; caly-
cibus 4—4.5 mm. longis, purpurascentibus, ovato-campanulatis, .
bilabiatis, labio superiore subintegro, inferiore bifido, fauce
intus subnudo, pedicellis 2-3 mm. longis elatis; corollis 10-12
mm. longis, albis vel pallide caeruleo-purpurascentibus, tubo
exserto, 4-5 mm. longo, inannulato, fauce dilato, labro erecto,
4-5 mm. longo, bifido, lobis ovatis, divergentibus, obtusis,
labiolo 8-10 mm. longo, lobis lateralibus oblongis, obtusis,
erectis, subtortis, lobo medio magno, 7 mm. longo, concavo-
dependente; staminibus 10 mm. longis, arcuatis, e labro
exsertis; stylo staminibus paulo longiore.

Cultivated throughout Latin America and apparently natur-
alized locally.

R. chilensis Molina is not improbably this plant and not
Sphacele campanulata to which it was questionably referred by
Bentham. Kuntze and Briquet, acting upon Bentham's sug-
gestion, have referred it there definitely and have made the
combinations Alguelagum chilense and Sphacele chilensis. The
present author has been unable to obtain the first edition of
Molina’s work except in the French and German translations.'
. These compare closely, however, and in both the supplements
to the narrative portion are in Latin and are identical. This ap-
pendix contains a systematic arrangement of the plants named
in the narrative, grouped according to the system of Linnaeus.
Speaking of R. chilensis, on page 129 of the French edition, the
author says ‘‘Le romarin sauvage (5) etant tres refineux, sert,
comme plusieurs autres arbustes, pour les fouderies de cuivre."
The numeral refers to a footnote giving the latin name, R.
chilensis, which is listed in the systematic appendix under the
class ‘‘Diandria, Monogynia.” In the second edition of his
work Molina makes no mention of Rosmarinus but describes a
new genus Phytoris, now known as Sphacele, and it is with this
genus that Rosmarinus chilensis has been confused. This plant
is listed under the class “ Didynamia.” ; |

Since Molina states, on the one hand, that his Rosmarinus

! Molina-Brandis, Versuch einer Naturgeschichte von Chili, 309. 1786; Molina-
Gruvel, Essai sur l'histoire naturelle du Chili, 329. 1789. —

[Vor. 12
122 ANNALS OF THE MISSOURI BOTANICAL GARDEN

is used for the table, and, on the other, that Phyioxis is very
acid and known to the natives as the ‘‘devil’s shrub" (Alhue
Lahuen) and since he places the former under the class ‘‘ Dian-
dria" and the latter under the class ‘‘ Didynamia," it seems very
improbable that the two are synonymous and very probable
that Molina was correct in referring his earlier described plant
to Rosmarinus, & genus with which he was undoubtedly ac-
quainted at first hand.

Specimens examined:

PERU: Ollantéstembo, 3 3000 m., May 18, 1915, Cook & Gilbert
811 (US).

Bourva: Island of Titicaca (Sonnen Inseln), 384 m., Nov.
1910, Buchtien (US).

MARRUBIUM L.

Herbae perennes, saepius tomentosae vel lanatae foliis ru-
gosis, in basi rarius cordatis, saepe incisis, floralibus con-
formibus, flores superantibus; floribus in verticillastris axillaribus
densis; bracteis maximam partem subulatis calycem aequilongis;
calycibus tubulosis, 5-10-venis, aequalibus, dentibus 5-10,
acutis, subspinosis, subaequalibus, erectis vel saepius ad maturi-
tatem patentibus; corollis tubo incluso, intus nudo vel suban-
nulato, limbo bilabiato, labro erecto, subplano vel concavo,
integro vel breviter bifido, labiolo trifido, patente, lobo medio
latiore saepius emarginato; staminibus intra tubum corollae
inclusis, parvis, antheris bilocularibus, thecis divaricatis, sub-
confluentibus, omnes subconformibus; styli a brevibus,
obtusis; nuculis apice obtusis nec —:

1. M. vulgare Г. Sp. Pl. 583. 1753.

M. hamatum Kunth in Humboldt, Bonpland & Kunth,
Nov. Gen. et Sp. Pl. 2: 310. 1817.

Herba altitudine 60—90 cm., in basi densiore ramosa, ramis
adscendentibus vel erectis, iterum in axillis inferioribus ramosis,
juventate arachnoideo-canescentibus, ad maturitatem sordido-
pannosis, quadratis, angulis obtusis; foliis late ovatis, obtusis,
1.5-3 em. longis, in basi rotundato-truncatis vel subcuneatis,
frequenter ad petiolam angustatis, margine crispo-crenata,

1925]
EPLING—SOUTH AMERICAN LABIATAE. I 123

crenarum culminibus 1-1.5 mm. altis, inter se 2-3 mm. dis-
tantibus, pagina superiore viride, tenuiter arachnoidea, bullato-
rugosissima, inferiore dense cano-arachnoidea, petiolis 1-2 cm.
longis, dense cano-arachnoideis elatis; floribus 7-20 in glomerulis
globosis densis dispositis, glomerulis inter se 3-5 cm. distantibus;
calycibus maturis 5-6 mm. longis, fauce paulo dilatis intus hir-
sutis, 10-venis, lanatis, sessilibus, dentibus 10-12, subaequalibus,
patentibus, subspinosis et hamatis, lanatis; corollis albis, 7-8
mm. longis, tubo crasso, aequale, 5 mm. longo, nectarostegio e
pilis brevibus ad basim staminum constante, labiis aequilongis,
labro erecto, ad medium bifido, lobis oblongis, labiolo trifido,
lobis lateralibus triangulo-oblongis, lobo medio majore, oblongo,
erispo; staminibus 2 mm. longis, didymis, ad tubi medium sitis,
in basi pubescentibus; stylo incluso, ramis acutis; nuculis 2.5
mm. longis, fuscis, oblongis.

A cosmopolitan weed of semi-arid habitat, appearing along
roadways and in fields. Frequently found in uncultivated
regions but in association with human activities. The plants
may be solitary or gregarious or may form dense patches several
feet across, or frequently cover an entire slope.

Specimens examined:

BnaAzir: Minas Geraes, Caldas, Sept. 3, 1861, Regnell 938 (US).

ARGENTINA: General Боса, Rio Negro Valley, 250-360 m.,
Nov. 17, 1914, Fischer 114 (US; MBG; GH).

Согомв:А: Moritz 994 (US); Bogota, Bro. Ariste-Joseph
(US); Loacha, 2500 m., Oct. 1911, Bro. Apollinare & Bro.
Arthur 83 (US).

BourviA: ? Cotani, 2500 m., Nov. 1911, Buchtien 5881 (US).

Ecuapor: Ambato, Aug. 24-26, 1918, Rose 23382 (US; GH).

Рево: Mollendo, Aug. 5, 1901, Williams 2528 (US); Matu-
cana, 2461 m., Apr. 12-May 3, 1922, Macbride & Featherstone
400 (FM; MBG).

Снил: Valparaiso, highways, 1895, Buchtien (US); Santiago,
Nov., 1922, Bro. Claude-Joseph 1486 (US).

PRUNELLA L.

Herbae perennes floribus in spicis densis extremis, verticil-
lastris 6-floribus; bracteis rotundatis, persistentibus, calyces

[Vor. 12
124 ANNALS OF THE MISSOURI BOTANICAL GARDEN

aequantibus et cum eis imbricatis; calycibus tubuloso-cam-
panulatis, irregulariter sub-10-venis et reticulato-venulosis,
supra planis, bilabiatis, labio superiore plano, lato, truncato,
breviter tridentato, inferiore semibifido, segmentis lanceolatis,
fauce intus nuda; corollae tubo amplo, subexserto, adscendente,
intus ad basim pilis squamisve brevibus annulato, sub fauce
subtus inflato, ad faucem paulo contracto, labro erecto, galeato,
supra subcarinato, integro, labiolo dependente, lobis lateralibus
oblongis, deflexis, medio rotundato, concavo, crenulato; sta-
minibus e tubo exsertis, didymis, filamentis in basi adentilis,
glabris, apice praesertim superioribus breviter bidentatis, dente
inferiore antherifero; antheris sub labro per paria approximatis,
liberis, bilocularibus, loculis distinctis, divaricatis; stylo apice
bifido, ramis subulatis; nuculis oblongis, levibus.

I. P. vulgaris L. Sp. Pl. 600. 1753.

Brunella aequinoctialis Kunth in Humboldt, Bonpland &

Kunth, Nov. Gen. et Sp. Pl. 2: 323, pl. 162. 1817.
à, Herba perennis altitudine 15-50 cm., in basi ramosis, ramis
paucis, decumbentibus, saepe debilibus et prostratis, purpuras-
centibus, glabratis vel in angulis nodisque hispidulis, quad-
ratis, angulis obtusis; foliis oblongo-lanceolatis, 3-5 em. longis,
acutis vel saepius obtusis, in basi angustatis, frequenter etiam
subtruncatis, crassiusculis, supra glabris, infra in venis his-
pidulis vel utrinque hispidulis, margine integra vel obscure
repando-denticulata, petiolis 1.5-2.5 em. longis elatis; floribus
in spicis extremis cylindratis, 2-5 cm. longis; bracteis calyces
subaequantibus? rotundatis, abrupte acuminatis, viridibus vel
purpurascentibus, frequenter translucentibus, venis reticulatis,
margine ciliata; calycibus 6-12 mm. longis, purpurascentibus,
hispidis, pedicellis brevibus elatis; eorollis violaceis, frequenter
albis, 9-15 mm. longis, labro labiolo duplo longiore; nuculis
2 mm. longis, ellipticis.

A cosmopolitan plant appearing in fields and particularly
along grassy margins of roadways, or frequently in openings in
the forest. While cosmopolitan it appears more commonly in
temperate regions and at higher elevations.

Specimens examined:

1925]
EPLING—SOUTH AMERICAN LABIATAE. I 125

СотомвтА: Salento, Caldas, 2100-2500 m., “edge of forest
above S.," July 25-31, 1922, Pennell 8892 (US); San Cristobal,
moist brushy mountain slope, 2800 m., Sept. 30, 1917, Pennell
2307 (US; MBG; GH).

CHILI: Valdivia: highways, 1898, Buchtien (US); Temuco,
Jan. 1923, Bro. Claude-Joseph 1950 (US).

LAMIUM L.

Herbae in basi decumbentes, foliis infimis longe petiolatis,
parvis, mediis majoribus, in basi saepius cordatis, rugosis,
plerumque duplicato- vel inciso-dentatis; floralibus subcon-
formibus, superioribus minoribus, Gésdilicribuk; omnibus calyces
superantibus; floribus in verticillastris densis, inferioribus vel
omnibus remotis, superioribus saepius approximatis; bracteis
paucis, calyces brevioribus, subulatis vel rarius lanceolatis;
ealycibus tubulosis vel turbinato-campanulatis, sub-5-venis,
ore aequali vel saepius obliquo, dentibus 5, subaequalibus vel
superioribus longioribus, apice subulatis; corollis tubo incluso
vel saepius exserto, intus nudo vel piloso-annulato, limbo
bilabiato, labro ovato vel oblongo, galeato, in basi plerumque
angustato, labiolo fauce dilatato, lobis lateralibus ad margines
faucis truncatis vel rarius oblongis, appendicula dentiforme
auctis vel muticis, lobo medio emarginato, in basi contracto;
Staminibus e tubo exsertis, antheris per paria approximatis,
bilocularibus, loculis demum divaricatis, oblongis, extus hir-
sutis vel nudis; stylo apice subaequaliter bifido, ramis subulatis;
nuculis triquetris, angulis acutis, apice truncatis, laevibus vel
minute tuberculoso-rugosis.

1. І. amplexicaule L. Sp. Pl. 579. 1753.

Herba annua 10-30 ст. altitudine, in basi ramosissima, ramis
decumbentibus, glabriusculis, purpurascentibus, internodo se-
eundo tertioque frequenter parte dimidia quam caule toto
breviore, internodiis superioribus multo brevioribus, quadratis,
angulis marginatis; foliis mediis ovato-rotundatis, 1-2 cm. longis,
in basi subcordatis, irregulariter crenato-lobatis, obtusis, sub-
hirsutis, petiolis plerumque lamina duplo triplove longioribus;
foliis floralibus 1-2 em. longis, amplexicaulibus, subtus venis

Vor. 12
126 ANNALS OF THE MISSOURI BOTANICAL GARDEN

elevatis, hispidulis, supra plerumque villosis, crenato 5-8-
lobatis, lobis obtusissimis; floribus in verticillastris compactis,
inferioribus distantibus, superioribus approximatis; calycibus
5-6 mm. longis, tubulosis, hispidulis, dentibus lanceolato-
linearibus, apice acuminatis, tubo aequilongis; corollis rubro-
purpureis, 15 mm. longis, tubo multo exserto, arcuato, intus
nudo, fauce dilatato, limbo parte tertia quam tubo breviore,
labro integro, labiolo albido, rubro-punctato, appendiculis
lateralibus nullis, antheris hirsutis; nuculis 1.5-2 mm. longis,
albido-punctatis et colliculosis, oblongis, in basi hyalinis;
gynobasi parva, lobo postice subnullo.

A cosmopolitan weed appearing particularly in waste ground
along the margins of fields and in flat open places newly sur-
faced.

Specimens examined: .

Рево: Oroya, 3076-4000 m., in moist soil, Kalenborn 32 (US;
MBG); Rio Blanco, half ascending in rock slides, May 8-19,
1922, Macbride and Featherstone 673 (FM; MBG).

Ecvuapor: Ambato, Tunguragua, Dec. 1918, Pachano 67
(US).

Снил: Santiago, Sept. 1, 1921, Bro. Claude-Joseph 1355 (US).

LEONURUS L.

Herbae erectae, foliis maximam partem inciso-lobatis, in-
ferioribus rotundatis, floralibus angustioribus, omnibus flores
longe superantibus; floribus in verticillastris densis distinctis
dispositis; bracteis subulatis; calycibus 5-venis, turbinatis, ore
truncato, dentibus 5, apice subspinosis, demum patentibus;
corollae tubo incluso vel rarius exserto, intus nudo vel oblique
annulato, limbo bilabiato, labro oblongo, integro, in his speciebus
subplano, in basi angustato, labiolo patente trifido, lobis
lateralibus oblongis, medio obcordato vel subfurcato; staminibus
e tubo exsertis, antheris per paria approximatis, bilocularibus,
thecis parallelis transversalibus, valvulis nudis; stylo apice
subaequaliter bifido, ramis subulatis; nuculis laevibus, triquetris,
apice truncatis, angulis acutis.

EPLING—SOUTH AMERICAN LABIATAE. I 127

1. L.sibiricus L. Sp. Pl. 584. 1753.

Herba erecta altitudine 40-60 cm., virgata vel ramosa, ramis
lateralibus brevibus, glabra vel tenuiter pubescens, canaliculata,
quadrata, angulis obtusis; foliis infimis rotundato-ovatis, sub-
cordatis, obscure trilobatis, lobis irregulariter incisis, obtusis,
mediis profunde trilobatis, lobis iterum incisis, frequenter sub-
pedatis, superioribus subintegerrimis, omnibus ad petiolam
longam attenuatis, subglabris, subtus pallidioribus, junioribus
molliter pubescentibus, petiolis parte dimidia quam laminis
brevioribus; floribus 8-25 in verticillastris densis, 1-1.5 cm.
latis, infimis inter se distantibus 2-4 cm., superioribus approxi-
matis; bracteis subulatis, 4-5 mm. longis, pubescentibus;
calycibus 6-8 mm. longis, turbinatis, 5-venis, molliter pu-
bescentibus, dentibus subspinosis, subaequalibus vel duobus
superioribus majoribus, ore obliquo; corollis rubris (pink-
Pennell), 12-15 mm. longis, pubescentibus, tubo e calyce paulo
exserto, nectarostegio e pilis brevibus supra basim 2 mm. ob-
lique annulato, limbo 6-7 mm. longo, labro oblongo, integro,
labiolo oblongo, trifido, lobis lateralibus ovatis, medio majore,
5—6 mm. lato, obcordato vel subfurcato; staminibus didymis ad
faucem sitis sub galea adscendentibus, filamentis ad basim
glandulosis; nuculis 2 mm. longis.

Specimens examined:

СотомвтА: Cartagena, 1919, Bro. Heriberto 240 (US); Tras-
quillo on Rio Sinu, grove along river in valley, 90-120 m.,
corolla pink, March 5-6, 1918, Pennell 4611 (US; GH).

VENEZUELA: Lower Orinoco, Sacupana, April, 1896, Rusby
& Squires 25 (US; MBG; GH); Tovar, 1854-5, Fendler 887
(MBG; GH); Caracas, March 14, 1899, Moore 2 (GH).

Brit. Guiana: coast lands, June, 1889, Jenman 5391 (US).

BnaziL: Glaziou (US); Rio de Janeiro, Martius 864 (MBG;
GH); Rio de Janeiro, Wilkes Exp. (US).

Paraguay: Central Paraguay, 1888-90, Morong 769 (US;
MBG; GH); in region of Alto Paraná R., 1909-10, Fiebrig 5369
(US; GH).

ARGENTINA: Sta. Iquazu, Nov. 17, 1910, Rodriguez 445 (US).

Vor. 12
128 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Lronotis В. Br.

Herbae fruticesve foliis variis, verticillastris densis, multi-
floribus; bracteis subulatis, numerosis; calycibus ovato-tubu-
losis, 10-venis, apice incurvis, ore obliquo sub-10-dentato, dente
supremo majore; corollis speciosis, coccineis vel flavescentibus,
patente villosis, tubo saepius exserto, intus nudo vel incomplete
annulato, limbo bilabiato, labro concavo, erecto, integro,
labiolo brevi patente trifido, lobo medio vix majore; staminibus
sub galea adscendentibus, filamentis in basi inappendiculatis;
antheris per paria sub labro approximatis, bilocularibus, loculis
divaricatis, acutis; stylo lobo superiore brevissimo; nuculis
apice obtusis.

1. Г. nepetaefolia (L.) В. Br. Prodromus, 504. 1810.

Herba erecta altitudine 50-200 cm., virgata vel ramosa,
ramis lateralibus brevibus, puberula, И quadrata,
angulis obtusis; foliis late ovatis, 3-10 em. longis, maximam
partem 5-7 cm., membranaceis, obtusis sed tamen apice leviter
angustatis, Боба ИВ vel truneatis et ad petiolam abrupte
angustatis, inciso-crenatis, crenarum eulminibus 2-5 mm. altis,
obtusissimis, inter se 2-5 mm. distantibus, utrinque viridibus,
tenuiter cano-tomentosis vel glabris, floralibus oblongo-lanceo-
latis, petiolis gracilibus, longitudine foliorum aequilongis; ver-
ticillastris paucis, distantibus, globosis, saepe ultra 100-floribus,
maturis frequenter 6-7 mm. latis; bracteis linearibus calycibus
aequilongis, apice spinescentibus; calycibus tubulosis 12 mm.
longis, maturis 20-25 mm., pubescentibus, in basi attenuatis,
apice abrupte incurvis, ore obliquo, maturo clauso, dentibus 8,
supremo maximo, acuto, spinoso, tribus inferioribus lanceolatis,
aequilongis, acutissimis, rigidis, subreflexo-patentibus, laterali-
bus utrinque 2 (3), brevibus, acutis, erectis; corollis coccineis,
20-25 mm. longis, tubo paulo exserto, ра dilatato, nectaro-
stegio e pilis densis incomplete 1-3 annulato ad tubi medium
sito, limbo tubo aequilongo, labro oblongo, 8-12 mm., integro,
extus dense villosis, concavo, labiolo 4-5 mm. longo, glabro,
mox marcescente, trifido, lobis oblongis, medio vix laterales
superantibus; staminibus in galea inclusis, antheris divaricatis;
stylo glabro, breve exserto, ramo superiore .5 mm. longo,

1925]
EPLING—SOUTH AMERICAN LABIATAE. I 129

inferiore 3 mm.; nuculis oblongis 2.5-3 mm., gynobasis lobo
postieo sub anthesi elongato.

Specimens examined:

CoLoMBiA : Cartagena, 1919, Bro. Heriberto 155 (US); thickets
on slopes east of Viterbo, 1100-1300 m., Sept. 4, 1922, Pennell
10260 (ASP); Сов, Cauca, 1000-1200 m., Dec. 1905, Pittier
631 (US); Dagua, El Valle, Dagua Valley, 700-900 m., May 13,
14, 1922, Pennell 5630 (US; ASP).

VENEZUELA: Cristobal Colon, pasture, Jan. 5-Feb. 22, 1928,
Broadway 146 (US; GH); Tovar, 1854-5, Fendler 894 (MBG;
GH); Island of Margarita, El Valle, July 19, 1901, Miller 74
(US; MBG; GH); Caracas, 800-1200 m., Apr. 8, 1913, Pittier
6028 (US; GH).

Brit. Guiana: Vreed-en-Hoop, west bank of Demerara R.,
opposite Georgetown, wasteland along railroads, Nov. 10-12,
1919, Hitchcock 16712 (US; GH).

BrazıL: Bahia, Salzmann (MBG); Rio de Janeiro, 1837,
Tweedie (GH); Porto Dom Pedro II, Dusèn 880 (MBG).

PARAGUAY: Cordillera de Altos, Dec. 1902, Fiebrig 584 (GH);
Central P. in regione lacus Upacaray, Oct. 1913, Hassler 12315
(US; GH; MBG).

ARGENTINA: Buenos Aires, Posadas, Feb. 20, 1914, Vattuone
& Bianchi 39 (US); Mendoza, Gillies (GH).

[Vor. 12, 1925]
130 ANNALS OF THE MISSOURI BOTANICAL GARDEN

ExPLANATION OF PLATE

Teucrium cubense Jacq. subsp. cordobense Epling
Argentina

From the type specimen, Kuntze, in the United States National Herbarium.

ANN. Mo. Bor. GARD., Vor. 12, 1925 PLATE 8

({товэте
Drstrnion бо балда ашар (әл doi ud A р Soe

“Ai he orr. ay cani EPLING BO
71 :

Lu M^ ceto tekeng l

ARGENTINIEN Ze Dr Otio Kontze
" Freegan а. hals
T E) —
Lal

EPLING—SOUTH AMERICAN LABIATAE

[Vor. 12, 1925]
132 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE.
PLATE 9
Teucrium tenuipes Epling
Galapagos Islands

From the type specimen, Stewart 3342, in the United States National Herbarium
(drawing inverted).

ANN. Mo. Bor. Garp., Vor. 12, 1925

PLATE 9

uh

SITED STATES NATIONAL MUSEUM

A

EPLING—SOUTH AMERICAN LABIATAE

Annals
of the
Missouri Botanical Garden

Vol. 12 APRIL, 1925 No. 2

COLLOIDAL SULPHUR AS A SPRAY MATERIAL
H. C. YOUNG
Chief, Department of Botany and Plant Pathology,
Ohio Agricultural Experiment Station, Wooster, Ohio
Formerly Research Fellow in the Henry Shaw School of Botany of Washington University

In a recent study on the toxic property of sulphur it was
found that sulphur became toxic to fungi primarily after it had
been oxidized into the somewhat unstable compound, penta-
thionie acid. This compound is most stable and very highly
toxic to fungi at a hydrogen-ion concentration of Py 4.2-5.4.
It was further found that the more finely divided sulphurs, and
especially the precipitated forms, were more easily oxidized
and were therefore more active as fungicides at temperatures
and conditions at which the flowers, or flour, of sulphur ex-
hibited toxicity to only a very slight degree.

Since these findings with reference to conditions favorable
for toxic action were not in accord with conditions existing in
most of the generally used sulphur sprays, it was thought ad-
visable to test the comparative value of representative sprays in
a commercial way. The need for a more active sulphur summer
spray is urgent, in that lime-sulphur, because of foliage injury,
is so frequently unsafe even on apple; likewise unsafe to a
much greater degree on peach and cherry; and cannot be used
at all on many of the smaller fruits. The other sulphurs, such
as the various dusts, the dry mix and wettable sulphur, etc.,
are not sufficiently active to control fungi under all conditions.
This lack of adequate control in all seasons is true of apple
scab, a disease that is, however, ordinarily controlled with most
of the sulphur sprays. In the case of blotch or bitter rot, the
sulphur sprays are generally reported as ineffective. Com-
AxN. Mo. Bor. Ganp., Vor. 12, 1925 (133)

[Vor. 12
134 ANNALS OF THE MISSOURI BOTANICAL GARDEN

mercially, the spraying program for any one disease is expensive
and may become considerably more so when several diseases
in which the requirements are different are to be controlled.

It was hoped that a sulphur spray could be developed that
would not only control apple diseases but would also be ef-
fective on other types of fruit and vegetables.

PREPARATION OF MATERIALS

Two types of colloidal sulphur were prepared. An attempt
was made at the outset to prepare the materials by the method
recorded in the previous report. Because of the lack of appa-
ratus many changes had to be made. The precipitated sulphur
was prepared as follows: Ten gallons of lime-sulphur, 32?
Baumé, were placed in a 50-gallon barrel. To this were added
10 gallons of water and 1 pound of glue previously dissolved in
hot water. Sulphurie acid diluted with 3 parts of water was
then added until the reaction of the mixture was Py 4.2. The
mixture was allowed to settle and the supernatant liquid de-
canted off. 'The residue, having the consistency of cream,
contained 1 pound of active sulphur to the gallon and was
used as a spray by adding 5 gallons of the precipitate to 95
gallons of water.

Considerable difficulty was encountered in the preparation of
the colloidal soluble sulphur. Fifteen gallons of a saturated
solution of sodium thiosulphate (hypo) were slowly stirred into
5 gallons of concentrated H,SO,, using a wooden barrel as a
container. At this point the mixture should have been cen-
trifuged, but no centrifuge of sufficient size was at hand.
An attempt was next made to neutralize the mixture with NaOH
and then to use it directly. In this way all of the soluble
compounds were retained in the mixture, resulting in consider-
able burning when applied to foliage. Moreover, the high con-
centration of electrolytes precipitated most of the soluble
sulphur.

Later in the season the mixture of hypo and acid was filtered
through a fine sereen and the filtrate neutralized with lime-
торош, Н. C. The toxic property of sulphur. Ann. Mo. Bot. Gard. 9: 403—435.

1925]
YOUNG—COLLOIDAL SULPHUR AS A SPRAY MATERIAL 135

sulphur diluted 1 to 3. During the addition of the lime-sulphur,
which was slowly effected, a further quantity, 3 gallons, of con-
centrated hypo was introduced. By the addition of lime-sulphur,
hydrogen sulphide fumes evolved, while SO, was produced by
introducing hypo at the same time. When these two gases
meet, soluble colloidal sulphur is formed, provided the sulphur
dioxide is in excess. This latter condition was maintained.
This method gave a very concentrated solution of soluble sulphur
and should have been an excellent spray material had it been
possible to have centrifuged it. For the same reason as stated
above, the soluble sulphur was soon precipitated. Analysis
showed that one gallon of the final mixture contained 10 ounces
of active sulphur.

PLAN OF EXPERIMENT

In order to test the new material under as many conditions
as possible and to be reasonably certain that diseases would ap-
pear in one or more locations, the experiments were tried in
5 states, namely, New York, Pennsylvania, Michigan, Illinois,
and Virginia. In all of these states apple scab is frequently
prevalent. Except in New York State the spray was applied
only to apple. The materials were prepared at the New York
Agricultural Experiment Station and then shipped to the cooper-
ating stations. The spraying tests at the New York station
were incorporated into the spraying program of the Department
of Entomology. The 2 colloidal sulphur preparations were used
in 3 Greening orchards and in one orchard of the Rome variety.
Five regular applications were made, namely, the delayed dor-
mant, the pink, calyx, 2 weeks, and early August. In the same
orchards, tests were made of lime-sulphur 1 to 40 plain, also
with glue, with casein, sulphur glue (8-4-50 formula), sulphur
casein spray (8-4-50 formula), sulphur lead arsenate dust (90-
10 formula), sulphur lead arsenate dust (90-10 formula) with
glue and casein sticker, sublimed sulphur (16 pounds) with
casein spreader, the same with 16 pounds of lime and casein
spreader, ground sulphur with casein spreader, the same with
16 pounds of lime added, 85-15 dust, and the copper sprays, as
listed in table I. The colloidal and precipitated mixtures con-

[Vor. 12
136 ANNALS OF THE MISSOURI BOTANICAL GARDEN

tained 5 pounds of sulphur per 100 gallons of spray and applied
at the rate of 8 to 12 gallons per tree on blocks of 10 trees each.
Sulphur dusts were applied at the rate of 5 pounds per tree and
the copper dusts 4 pounds per tree.

On July 18, leaf counts were made on the Rome orchard,
Geneva, New York. The results are tabulated in table 1.
Some of the check trees of this orchard were covered with a
canvas during the spraying operation and all the trees used for
checks had been checks for several years. This rather abnormal
treatment of the checks may have resulted in the heavy initial
infection. Counts were also made in two near-by test orchards
of the Greening variety, the results of which showed practically
no disease on the treated plots and less than two per cent on
the check. Later in the season scab became more general on
the Greening checks. From all indications the initial infection
was not heavy, and a rather mild secondary infection occurred.
In the main, scab was not difficult to control in the vicinity of
Geneva, during the season.

TABLE I

RESULTS OF u^ COUNTS ON THE ROME а. №. У. AGR. EXP.
ТА., GENEVA, MADE JULY 18,

No. of No. of Per ce
Plot 'Treatment trees leaves infection
$ counted

1 Lime-sulphur 5 2500 .68
2 Lime-sulphur & Ca casenate 5 2500 .96
3 Sulphur & glue (84-50) 5 2500 2.5
E Sulphur & casein ( 5 2500 6.5
5 Colloidal sulphur 5 2500 .16
6 Check, trees covered 2 000 98.

T Check, trees uncovered 2 90.

8 90-10 dus 5 2500 T.I
9 90-10 dust & glue sticker 5 2500 6.1
10 90-10 dust & casein sticker 5 2500 5.4
11 Precipitated sulphur 3 1500 4.2
18 Special process sulphur 2 1000 40.

At harvest time all the fruit in all of the plots was counted
and a summary of results is given in table п. A more detailed
account of all the tests is given in the report of the Department
of Entomology, New York Agricultural Experiment Station,
Geneva, New York.

1925]
YOUNG—COLLOIDAL SULPHUR AS A SPRAY MATERIAL 137

TABLE II
COLLOIDAL AND PRECIPITATED SULPHUR COMPARED WITH OTHER
SPRAYS IN NEW YORK STATE
No. оЙАуег. no.|AVer. по.|Рег cent|Per cent
No. Treatment trees | apples | scabby | scabby | clean
exam-| per tree apples apples
ined per tree
1 |Spray, lime-sulphur..... 17 1683 11.4 1.18 88
5 |Spray, sublimed ры
а. 10 1415 29.5 2.08
6 ry e ae ipn
., 8 lb. lime...... 9 2265 46. 2.02
7 |Spray, ena sul-
hy o о: 6 1932 14. ‚12 96
ы 8 |Spray, colloidal sulphur. 6 1924 28. 1.45 91
Е 9 |Эргау, flour sulphur, 16
© OES et peo И, 9 2388 34 1.43
£ | 10 |Spray, flour MM 16
о Ib., 8 Ib. lime........ 10 | 1651 32.2 | 1.95
CI H Spray, flour Е er 7 3486 50.3 1.44
E 12 |Spray, flour sulphur
© RMB РЕЯ 9 | 1895 | 74.7 | 3.94
© 18 Dust, 90-10, 51b. per tree| 9 1143 19.0 1.65
= | 15 |Dust, 85-15, 51. рег tree| 8 1287 7. 53
is | 17 Dust, 77-13-10 lime cop-
т, 4 Ib. per tree..... 10 1846 | 224.8 | 12.17
21 |Dust, copper arsenate
cis II IQ, 10 1127 | 213. 18.89 | 54
22 |Check, no treatment.. 5 849 | 736. 86.81 5
23 |Check, 1 application be-
sides lime-sulphur, paige
PLA. nicotine a {
Beg VEU Kc Rue С 5 1474 | 334 22.68 | 36 -
1 АИЯ 1 1284 2.3 86.6
E| 4 [Precipitated sulp 1 | 2080 1.8 | 96.3
gaj 5 mos mph & glue 1 234 d "T
95] 7 я чалы
t 1 [Dust 90-10... ........- 4 5.0
St 4 Wettable bM АЕ + 4.8
> E 7 |Dust, Жл соррег : 19.75
tf р Collo idal sul h PE Ed 4 4.0
Ci E ncn а И 6 83. 07
1 DIE. os eke 1.7
ov| 3 |Wettable sulphur...... 5 A
ЕЕ 5 [Colloidal sulphur. ....-- 5 5.9
@ 2 1 апзї............ 2 2 2
"|n lee 13 90.7

Spray injury was also noted on all the sprayed plots. The
most severe burning resulted from the lime-sulphur spray.

[Vor. 12
138 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Occasionally some burning was noted with colloidal sulphur.
The copper sprays caused some leaf burning and an immense
amount of russeting of fruit. In general, leaves sprayed with
precipitated sulphur appeared best throughout the season. A
more detailed discussion of spray injury is given by Young and
Walton.!

The material sent to Michigan was applied by Professor W. C.
Dutton of the Department of Horticulture. A late frost caused
all the fruit to drop so that the experiment was discontinued
after the second application.

In Pennsylvania the materials were applied by Mr. R. C.
Walton, of the Department of Plant Pathology, Pennsylvania
State College. The tests were conducted on Stayman Winesap
apples in orchards owned by Mr. Eshelman and Mr. Keller.
Precipitated and colloidal sulphur, commercial lime-sulphur (1
to 40) with and without arsenate of lead, dry mix, and sulphur
dust were used. The sprays were applied with a 300-gallon
power sprayer equipped with 2 guns and giving a pressure of
250 pounds. In 1922 the trees were seriously affected with scab.

The delayed dormant spray was entirely of lime-sulphur and
applied April 18, when the first leaves were showing approxi-
mately 14 inch but had not started to spread. The ‘‘pink”’
application was made April 30, at which time about 1/5 to !/, of
the central buds were opening and the stems had separated.
On May 11 the ‘‘petal fall" application was made, when prac-
tically all the petals had fallen. The 10-day application was
made May 22. On May 23 very little scab could be found, not
even on the check plots. Scab began to show on the check
plot to a considerable extent on June 4, but was not serious.
The weather, prior to the opening of the blossoms, was very dry.
The rains were few and light, and conditions in general were
unfavorable for scab development. А 4- to 5-week spray was
applied June 20, and the late summer spray, August 6.

ing the 4- to 5-week spray the temperature was 100? F.
in the shade. "The lime-sulphur burned severely, and traces of
burning were noticed in the precipitated and dry-mix plots. On
August 6 the check trees were heavily infected with scab on
both fruit and foliage.

'Young, Н. C. and В. C. Walton. Spray injury to apples. Phytopath. 15:
405-416. pl. 14. f. 1. 1925.

1925]

139

YOUNG—COLLOIDAL SULPHUR AS A SPRAY MATERIAL

— ouy
E | | [ил
т eunf
ounf
с Avy
т Avy
І Av
ABW
єє rr | 9°8 Ar | S’St [1978 29°F | 6F'S |se'ereA4'r [88° "Ig _Хв 6. зеп 01-06] 6
€ 6| LI | 0'9 |Z°8 | SZ TO 8`6 | ZL°ST |19 STIIT 'eT|IS "9193 TG '* '3ueunen
d EARS Мет Oud x d. ONT кт
ЖЕ. 1 9`б1(6°? |815 |900 | 2 9' 9€ |£019' |85192 | от 9 х-га) 4
ЕЕ 20118 `9 | 0°55 |6801] 9'9 2' UI |57|$’ |8’ | l'€| ota) 9 |^ po ‘вгу
—1dnudms-ourT! qo
£u Z'8 |С | #55 |0951| 8's 8` РЕТТЕ 049 | он 6 05 әп
[yun I ON
cme —anydins-oury| 89
9 "any
0c Pune
55 ABW
ZI-
di Avy
e
EI ы £°9 |6°т | FSI (Ой 9T é Ie TESS" |À^ eS 99 D pevejydroei1g| $
VINVATASNNGd "W'TIIASLONWHV ‘ачуночо dVSANIM NVWAV.LS NVW'IHHSS 'O '8
TIZ | 441] 8°¢ | s'orjo'ee| e'99 [ecOT 865] 9°29 | 8' v6 Ad die 9'79 309019891}
as UE омон 4
ов! CC тес |» wo o $ аа [€ № г о _$ | mud[ns-eurg 9
29i EI IS LIES TOI Е *8' I6 | 66 AL [00:9 |8?° [69° |cOP| Ond еер mud
M -ins — [epropoo| _$
oot I (ЖА Ж | 6 gol 9° LS SLUTS LTS 16 100: oma 9 xmrÁdq| c6
8 ‘any
$5 AB
И Avy
т Avy
| А 85 "dy mudrns
III M СӨТЕ 990] Г A 9°21 | 67|?’ |?’ | ТТ 105 чу $ раза | T
зәзәтдә тэДот _ qyour KO eun snq punos|3maj| рүеәѕ | Yo oTq gods 1—30, "43S POW 3485 әш, [ох A ‘ON
Әәл, эл "HDI чау ста -un % ‘ON 9pts % 3008 % пш} 4o quos ju9o ЈӘ, d suorjeondd y quon et, Vd

VINVATASNNGd ‘ATTIASLONGUV

III WISVIL

GUVHOUO 4ӯБЯМІМ NVWAVLS ЧАТТУЯ

LE III
KELLER STAYMAN WINESAP ORCHARD, ARENDTSVILLE, PENNSYLVANIA

Plot| Treatment |APPlications| Percent seab — |» fruit 7$, sooty] % side| No.l% un. Red phis Codling
No. ‘ No.| 'Time Slight Mod. Sev. | Total Spot lotch scald |fruit| sound bug moth [roller ler icket
1 |Precipitated 5 |Арг. 20| 4.1 .4| .4|4.9| 17.6 4.7 .1 [1086 11.2 | 1.3| 2.3 WETA
sulphur Apr. 28
May 1
May 11
May 23
Aug. 3 ; ^ tol
2 |Dry-mix 5 |Ditto GH} 9i .41T.8 117.2 5.1 .6 |1052) 9.4 | .7 2.7 do 415.6
3 {Colloidal sul-
Е _5 |Ditto | 4.02) .59| .48| 5.09] 17.92 | 21.84 1176) 19.0 | 8.2) 3.4 | 1.2 | 6.3
5 |Lime-sulphur .5 |Ditto 1 ЗЕ Toy Е .9 11035] 11.4 |] .2 2.1 | 2.2 | 8.6
7 |Check—No
treatment 54.6 |24.4 [15.8 |94.8 | 67.6 | 72.8 1023| 66.3 |33.6]10.4 | 5.8 |17.7 | 21.1
S. C. ESHELMAN STAYMAN WINESAP ORCHARD, ARENDTSVILLE, PENNSYLVANIA
5 |Precipitated 6 |Арг. ss т а 19 1.6 11202 12.4 | 1.9] 6.3 A $5.9
May 11
-12
May 22
June 20
Aug. 6 p [LE ———
6a |Lime-sulphur —
No lead until
e 20 6 |Ditto 1.9 6 та I. 8 5.8 1260] 22.3 4.5) 8.2 б H12
6b |Lime-sulphur : Sues. DOM
Ars. le 6 Ditto | 3.1 8 SA ERI 1.1 ‚8 6.6 [1059] 22.0 | 6.510.7 5 E59
7 |Dry-Mix 6 Dito | 7.5 | 2.2) .6 110.3 | 2.6 .6 2.7 | 906| 21.8 | 4.912.6 SEO 144
8 eck—No | | Sighs b Voc
treatment... ^ 51.54/16.81/13.16/18.51| 13.72 | 9.8 .641| 25.4 | 8.7 6.0 | 1.7 |9.2
9 190-10 Dust 9 zt 11.27| .58| 1.7313.58| 5.49 4.62 347| 15.5 | 1.7| 8.6 Pek 3.8
ay
May 1
May 1
May 2
June
June 1
July 1
Aug. —

[9561

ТҮІЧЯШҮИ AVUdS V SV unHd'InS 'IVdIOTIOO—95NO00X

681

[Vor. 12
140 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Tests including both the precipitated and colloidal sulphur
were made in the Keller Stayman Winesap orchard. The spray
schedule was practically the same for each test. On August 3
the fruit on the ‘‘precipitated”’ plot appeared very fine and the
owners were very much pleased with the spray. Observations
made on September 28 indicated that the fruit sprayed with
precipitated sulphur, colloidal sulphur, and dry-mix sulphur was
more highly colored than that sprayed with lime-sulphur. The
results from both orchards are given in table пт.

The Illinois tests were conducted by Dr. H. W. Anderson, of
the Department of Horticulture, University of Illinois. In his
orchards scab did not show to any extent on the check trees.
He states that neither the precipitated nor colloidal sulphur con-
trolled blotch. Where the checks showed 98 per cent blotched
apples, the precipitated sulphur gave 48.4 per cent and the
colloidal 70 per cent. Lime-sulphur gave 26 per cent. A more
complete report of these tests was given by Dr. Anderson at a
meeting of the Illinois State Horticultural Society in December.

The experiments in Virginia were conducted by Professor
Schneiderhahn at Winchester. He reported that owing to the
dry season no disease developed on the check plots and he
therefore obtained no check on the material. He stated further
that he has never observed before such a striking correlation
between climatic factors and disease prevalence as exists in
this section, and also: “аё in spite of the fact that the trees
in these plots received approximately 14 gallons of spray ma-
terial per tree, there was no perceptible spray burn on leaves or
fruit and the variety is the Ben Davis,—the most susceptible to
spray burning. I am convinced that there is very little, if any,
danger from spray burning with the materials you sent me.
We have had exceptionally hot weather and the leaves of these
trees were nearly white after the spray dried, but no damage
resulted." :

Discussion or RESULTS

In general the past season cannot be considered one favorable
for scab development, especially in Illinois and Virginia. Since
nearly every type of sulphur spray gave commercial control of

1925] :
YOUNG—COLLOIDAL SULPHUR AS А SPRAY MATERIAL 141

scab, it might be said that climatic conditions favored the toxic
action of sulphur. The dry and warm season in both New
York and Pennsylvania undoubtedly increased the action of
sulphur.

Because of the type of season it is difficult to draw very
definite conclusions as to the value of one sulphur spray over
another. In the main, however, the precipitated sulphur was
not quite as effective as lime-sulphur, but more effective than
the dusts, dry mix, or wettable sulphur. Its value was greater
than lime-sulphur because in most every case no injury to
foliage resulted and in every case a higher percentage of clean
fruit was obtained. In New York, Pennsylvania, and Virginia,
the growers in whose orchards precipitated sulphur was used
were very well pleased with it. Whether it would be as ef-
fective in a season more favorable for scab development, is a
question that can be answered only by further trials. The
materials, both colloidal and precipitated sulphur, could not
be prepared as desired, because of the lack of equipment, and
it is thought that both forms would be considerably more
active and injury to foliage less if better purification methods
could be employed.

Since precipitated sulphur caused practically no injury to
apple foliage, it was thought advisable to try it on other
fruits, usirig the same concentration as for apples, namely, 5
gallons of the concentrated in 100 gallons of spray. Trees of
two-year-old peaches were sprayed heavily. The peach trees
were in a rapidly growing condition and therefore in а suc-
culent state when they are more susceptible to injury. They
were sprayed July 24, during a spell of warm weather.
Careful observations were made daily for 14 days, during
which time no leaves that showed spray injury could be found.

A similar experiment using colloidal sulphur was conducted
at the same time. There resulted in this case considerable
burning but not enough to be of economie importance. These
tests were continued until August 30, during which time the
applications of colloidal sulphur resulted in injury.

Both materials were tried on sweet and sour cherries, grapes
and plums. Slight injury was noted on the foliage of cherry

[Vor. 12
142 ANNALS OF THE MISSOURI BOTANICAL GARDEN

and severe burning on grapes where the colloidal sulphur was
used, but no injury whatever with the precipitated sulphur.
The injury eaused by the colloidal sulphur was undoubtedly
due to the soluble sodium salts that it contained.

During the summer, colloidal sulphur preparations sent to us
from the Texas Gulf Sulphur Company and the Riches-Piver
Company were sprayed on peaches of the same type and con-
ditions as mentioned above. No foliage injury resulted from
either compound. Both types spread and stuck very well. In
one ease at the times of application, about 3 P. M. and about
5 P. M., it began to storm and rain and continued intermit-
tently until morning. The following day the trees showed
white and very little of the spray had been washed off. The
precipitated sulphur was used in this same experiment and very
little remained on the trees. The Texas Gulf Sulphur Company
product had a hydrogen-ion concentration of 7.4, that of the
Riches-Piver 4.2. No test could be made at that time as to
the fungicidal value of the two products. The Herbert and
Herbert product was received too late for outside tests.

One of the most serious defects in both the precipitated and
colloidal sulphur was their sticking quality. The quantities of
CaSO, and other coarse insoluble matter prevented their close
adherence to the leaf surface. When prepared in a pure state
both compounds stick very well, the colloidal especially. The
sticker casein could not be used because of its reaction. Late
in the season Professor Dutton, of Michigan Agricultural
College, called my attention to semi-solid buttermilk as pre-
pared by the Consolidated Products Co., of Chicago, Ill. This
company kindly sent me a gallon of the product. This amount
was added to 50 gallons of spray mixture of precipitated sulphur.
This mixture was applied to peach trees, and stuck and spread
very well. The reaction of the mixture was not altered.

The possibility of having a general sulphur spray that can be
used on all fruits seems certain, especially in districts where
blotch and blister rot are not present. My products, prepared
in the manner they were, were for my own satisfaction, and
there is little doubt in my mind but that the commercia,
products listed above will prove to be more active fungicidallyl
providing their reaction point be near Pg 4.2-5.0.

1925]
YOUNG—COLLOIDAL SULPHUR AS A SPRAY MATERIAL 143

The work here reported represents some practical applications
of the sulphur project which has been pursued at the Missouri
Botanical Garden. The writer is greatly indebted to the New
York Agricultural Experiment Station, Geneva, and thanks are
due the members of the Department of Entomology for their
hearty cooperation, and especially to Mr. P. J. Parrott, for
making available facilities for work, and to Mr. G. P. MacLeod
for assistance in the preparation of materials. "Thanks are also
extended to the cooperators mentioned in the text and to the
Crop Protection Institute, under whose auspices the work was

one.

PIGMENT STUDIES WITH SPECIAL REFERENCE TO
CAROTINOIDS IN FRUITS:
GRACE E. HOWARD

Instructor in Botany, W ellesle
Formerly Jessie R. Barr Fellow in the Henry Shaw | repr of Botany of Washington

This investigation, concerned primarily with the carotinoids,
has been divided into 4 sections, as follows: (1) plastid studies;
(2) the use of the colorimeter in the determinations of relative
amounts of fruit pigment; (3) determinations of the presence of
certain pigments in various fruits; (4) ripening experiments.
These sections will be discussed in the order indicated.

PLASTID STUDIES
INTRODUCTION

The problem of the origin and structure of leucoplasts, chloro-
plasts, and chromoplasts has occupied investigators for many
years. The state in which the carotinoid pigments occur in
fruits, flowers, and leaves has been much under discussion. Are
these carotinoids in crystalline or granular form and more or less
free in the cell cytoplasm, or are they necessarily imbedded in
lipoid material and usually contained within a stroma or plastid?
If, as some workers believe, chloroplastids are the progenitors of
chromoplastids in which the carotinoid pigments must be con-
tained, how are we to account for the presence of these pigments
in the animal kingdom or in various groups of fungi in which
chloroplasts are never present?

In an attempt to throw some light on these problems micro-
scopic studies were made on plastids of green, yellow-orange, and

red-ripe fruits. Fruits were chosen because the plastids are
large and changes can be very easily noted during the ripening
process. In several kinds of fruits the decolorization of the
chromoplasts was accomplished, leaving behind а colorless
stroma ри: cecus to that „+ Ше оны Ошон the in-

Ann. Mo. Bor. GARD., Vor. 12, 1925 A “©

[Vor. 12
146 ANNALS OF THE MISSOURI BOTANICAL GARDEN T

vestigation evidence was accumulated to show that in some
fruits the carotinoids were in granular or crystalline form, while
in others these pigments were contained in plastids commonly
known as chromoplasts. No attempts were made to show just
where the color was located in the plastid.

SURVEY OF LITERATURE

The earliest work on plastids was concerned especially with
the study of starch grains. The investigations of von Mohl (755),
Böhm (57), Nägeli (58), and Sachs (762) showed that starch
grains appeared in a great many of the chloroplasts. More than
10 years before Schimper's experiments on plastids we have the
first work along this line by Kraus (72) who sought to show that
the yellow color of the tomato and the rose was originally green,
and that the yellow color originated from the green, also that the
green chlorophyll-containing bodies, with or without a continu-
ation of form, acquired a yellow color. He assumed that in this
ease the blue-green pigment disappeared and the original yellow
remained or even increased in amount. Kraus was the first
worker to prove that the yellow fruit pigments are identical with
those of this class in the chloroplasts of the leaf.

Another of the early workers was Millardet (76) who believed
that the chlorophyll bodies became more or less deformed just
prior to the disappearance of the chlorophyll. At the same time
a great number of these bodies losing their coherence, became
elongated and irregular, appearing separate and viscid. А little
later they became swollen and coalesced to form somewhat ir-
regular masses located in vacuoles of the cytoplasm. They as-
sumed a more or less yellow-orange tint, due to the presence of
small rods of *solanorubin." This colored substance was finely
granulated but coarser than that of the chlorophyll. In a later
stage these rods of solanorubin multiplied and increased in size.

Schimper (’83), in his epoch-making studies on the origin of
plastids, concluded that all plastids arose from the division of
preéxistent plastids, that they never arose de novo from the
protoplasm. These plastids were then transmitted only in the
female line, never through the sperm. He found chloroplasts
in the embryos of a great many of the higher plants, such as Hy-

1925]
HOWARD—CAROTINOIDS IN FRUITS 147

pericum, Tropaeolum, Evonymus, Helianthus, Acer, etc. Schim-
per showed that leucoplasts may be transformed into chloroplasts
by the chlorophyll becoming photosynthetically active or they
may remain as leucoplasts and serve in the building of starch
from food already assimilated. Finally, in various parts of the
plant, as, for example, in roots, leaves, flowers, and fruits, the
chloroplasts may become transformed into chromoplasts of
various shapes and diverse colors.

That chloroplasts arise from leucoplasts and chromoplasts from
chloroplasts, was Schimper’s view. He noted an exception to
this in the fruit of Symphoricarpus racemosus, which, in the ripe
condition, contained leucoplasts which were transformed chloro-
plasts. The latter phenomenon was found to occur in many
embryos. This writer assumed that the chloroplast, or the
photosynthetic chromatophore, is the most primitive of the 3
types of plastids. From the phylogenetic point of view the
leucoplast and chromoplast represented later development and
arose concurrently with an increase in tissue differentiation.

Meyer (’83) substantiated much of this work of Schimper’s,
so that often the theory of the origin of plastids from preéxistent
plastids is known as the Schimper-Meyer theory. This theory
of Schimper and Meyer has never been seriously called into
question in the higher cryptogams nor in the algae, but in phanero-
gams it has caused some discussion. Mikosch (’85) and Belzung
(87) could find no chromatophores in the resting seeds which .
they investigated. After examining ripening seeds, mature
seeds, and seedlings of a large number of plants they concluded
that growth of starch grains can take place without leucoplasts
and that chloroplasts are formed directly by differentiation of the
cytoplasm.

Courchet (’88) decided that ‘‘chloroleucites” and most of the
* chromoleucites" have an analogous structure. They consist of
a protein substratum or stroma in which are enclosed granules
either of chlorophyll or other pigments. Rarely, these pigments
are truly crystalline in nature. He believed with Schimper that
the “leucites” always resulted from the division of preéxisting
leucites and that they multiplied by direct division. Frequently
the chromoleucites arose independently of the colorless leuco-

[Vor. 12
148 ANNALS OF THE MISSOURI BOTANICAL GARDEN

leucites. He studied many different flowers and fruits, among
them the flowers of Aloe and the fruits of Evonymus japonicus
(arils) and Cucumis Citrullus.

Another worker to substantiate the Schimper-Meyer theory of
the origin of plastids was Chmielevsky (’90), who showed that in
Spirogyra the chromatophores of the male cell disappeared at
the time of cell conjugation, while those of the female cell divided
and gave rise to the chromatophores of the new plant. Davis
(99) observed the division of a chloroplast in a spore mother
cell of Anthoceros. Zimmermann (94), who worked primarily
on algae but also on ferns, discussed the chemical and physical
character of the chromatophore pigments. His views regarding
the development of plastids were the same as Schimper’s.

Lotsy (07) claimed that the Schimper-Meyer theory was not
sustained by sufficient evidence so far as higher plants are con-
cerned. He suggested that if hybrids could be obtained between
species with markedly different chromatophores, it might be
shown experimentally that chromatophores were transmitted
only in the female line. Senn (’08), in his extensive work on the
form and change of position of chromatophores, studied a great
many different algae, also the leaves and stems of spermatophytes,
but no fruits were investigated by him. He found that chromato-
phores change form and position in response to light, temperature,
water content, also chemical and mechanical influences. He ob-
served protoplasmic connections between plastids.

Sapehin (’11) observed that during sporogenesis in mosses each
archegonial cell contained 1 or 2 deeply staining bodies lying close
to the nucleus; these he considered to be chloroplasts. A si
condition казыр in Anthoceros. All the other Bryophytes and
Pteridophytes as well as the Spermatophytes examined showed
a considerable number of plastids in the archegonial cells. He
traced the division of the single chloroplast in the spore mother
cell of various mosses into 4, one going into each tetrad. The
ripe spore was found to contain several chloroplasts, as did all the
cells of the gametophyte and of the embryo sporophyte. Sapehin
(13), working with Lycopodium, showed that each cell of the
young antheridium contained numerous plastids, but the number
is reduced to one in the antherozoid mother cells. This work is

1925]
HOWARD—CAROTINOIDS IN FRUITS 149

very important as a substantiation of the Schimper-Meyer
theory, for it has sustained the idea of complete genetie continuity
of the plastids throughout the life cycle in various groups.

Rothert (12), in an extensive study of various vegetable
organs from many different plants, examined and described care-
fully the form and shape of the plastids. He found the pigment,
whether yellow, orange, red, or brown, to be in distinct granular
form, distributed in the stroma, probably dissolved in drops of
oil. Inone plant, a saprophytic orchid, the pigment was observed
to exist as needle-shaped crystals. He believed that chloroplasts
and chromoplasts contained the same pigments but in quantita-
tively different proportions. The carotin was present in distinct
grains but the chlorophyll was distributed evenly in the stroma,
the latter often colorless but sometimes pale green. His views
on the origin of plastids were very similar to those of Schimper,
that is, that they form an unbroken series. Molisch (13) be-
lieved that leucoplasts and chloroplasts may be transformed one
into the other, and that chromoplasts may be finally produced
from chloroplasts.

No review of literature on the subject of plastids would be
complete without some mention of mitochondria or chondrio-
somes, employing these terms as synonymous. An enormous
literature has accumulated on the subject, both from the animal
and plant side. Only one phase of it will be given here, namely,
that which has to do with the theory that mitochondria are the
progenitors of plastids. A very good general review and biblio-
graphy of the subject was given by Cavers (14), and likewise by
Sharp (21). Among those who claim mitochondria to be the
progenitors of plastids Lewitsky (10) stands out as one of the
most important of the early workers. He found these bodies
present in all essential parts of the cytoplasm, not only in the
pollen mother cells but also in the pollen grains of Asparagus
officinalis. In the stem end of the embryo, the chondriosomes
changed to chloroplasts; on the other hand, in the root end they
were transformed to leucoplasts.

This work on mitochondria, so contrary to the ideas of Schimper
and Meyer, was criticized by the latter (Meyer, '11), on the
ground that if such bodies really existed, it was extraordinary

[Vor. 12
150 ANNALS OF THE MISSOURI BOTANICAL GARDEN

that he and Schimper should have overlooked them. Rudolph
(12) obtained no evidence of any relation between chondriosomes
and chloroplasts. He worked with a large number of plants
belonging to various groups, using the Benda killing method and
iron-haematoxylin stain with best success. He concluded from
his results that the value of all previous work was merely to show
the existence of such bodies in many different plants. Mottier
(18) was also unable to confirm Schimper's view, in a study of
the root-tips of Pisum sativum and Zea Mays, of the thallus cells
of Marchantia and of Anthoceros, and cells of the stems and leaves
of Elodea canadensis. In this work he found cells containing both
chloroplasts and chondriosomes, and his conclusions were that
the primordia of leucoplasts were not the so-called mitochondria.
He observed other granular bodies that did not become leuco-
plasts; these he considered mitochondria whose function could
not then be definitely formulated. He suggested that perhaps
these last are concerned in some processes of metabolism. Twiss
(19), working with Zea Mays and Preissia, was able to show an
unbroken series from mitochondria to plastids in the root-tips
of corn, while in Preissia this seriation was not so obvious
However, he believed that such definitely staining bodies as
mitochondria exist as normal constituents of the protoplasm.

Guilliermond has produced more evidence than any other
worker to show that mitochondria are progenitors of plastids.
He has published a number of papers, only a few of which can
be reviewed here. In an important work (Guilliermond, 713)
he described the occurrence of chondriosomes in the young asci
of Pustularia vesiculosa, in Penicillium, Botrytis, Endomyces,
yeasts, and in various Autobasidiomycetes. In a later paper of the
same year he found chondriosomes in a number of Pezizeae.
This was especially interesting since these forms contain an
abundance of carotinoids. In Pustularia he distinguished 3
kinds of reserve substances, glycogen, fat, and the so-called

“metachromatic corpuscles” of earlier writers. He claims to
show that the latter arise from chondriosomes in much the same
way as do the plastids of higher plants. His paper (Guilliermond,
'19) records the results of his extensive studies on the epidermal
cells of the petals of several different flowers, such as tulip and

1925]
HOWARD—CAROTINOIDS IN FRUITS 151

iris, and also the mesophyll cells of several fruits including the
berry of Asparagus. He observed the shape, form, movement,
and division of chondriosomes of living material in isotonic
solutions, and also studied material fixed by Benda’s method and
stained with iron-haematoxylin. A detailed explanation was
given of just how the different plastids were formed from the
chondriosomes. He argued that Schimper’s so-called protein
erystalloids were only the long-drawn-out ends of the dividing
chondriocontes. Upon examination they showed no optical
properties. In a later paper (Guilliermond, ’21), he described
the origin of leucoplasts from their primordia in root-tips of castor
bean, bean, pea, corn, and a gourd. He acknowledged the fact
already established by Rudolph, Sapehin, and Mottier that,
besides the primordia of plastids, there may be other similar
granules and rods present in a cell which do not become plastids.
Kozlowski (22), in a criticism, said that the whole subject of
mitochondria was too indefinite and needed clearing up. He be-
lieved that a worker started with a preconceived idea of what
chondriosomes should produce and then fitted them into his
particular problem.

There has been a little work devoted especially to the formation
of carotinoids. The Toblers (10) made an extensive study of the
nature and appearance of carotin in the fruit of Momordica
balsamina. They found that after reaching maturity all parts
of the pericarp quickly became green and showed rich starch
content. In the layers of the mesocarp near to the fibro-vascular
bundles and also in the endocarp, starch was especially abundant.
As the fruit ripened there was a decrease in the amount of starch,
and while the mesocarp was becoming yellow the endocarp was
turning a light rose-red color, which constantly increased. In
this region the deep red color of the aril finally became apparent.
The orange-yellow color characteristic of the mesocarp was due
to the presence of lens and needle-shaped crystals. In another
piece of work finished in the same year the Toblers (10а) con-
cluded that, in the absence of more conclusive evidence, the idea
of a direct genetie relation between chlorophyll and carotin must
be excluded. Using the same fruit, Momordica balsamina, they
decided that throughout the ripening process the formation of

[Vor. 12
152 ANNALS OF THE MISSOURI BOTANICAL GARDEN

carotin was connected with the decomposition of chlorophyll.
They attempted to show a possible relation between carotin
formation and the path of nutrition, since the basal part of the
fruit always remained green longest. They showed that if
respiration is prevented by covering the fruits with vaseline, the
ripening process is retarded.

The same workers (Toblers, '12) made comparative researches
on chlorophyll, carotin, starch, and sugar in 3 regions of the
carrot, namely, top, middle, and base. "They found the amount
of carotin increased in indirect proportion to the chlorophyll but
directly with the starch and sugar content. Age seems to deter-
mine the proportion of starch and sugar present and also the
amount of carotin. In old roots both starch and carotin de-
creased from the top downwards, although in younger ones starch
was in excess at the top. The sugar content always increased in
all regions with age. 'The progressive greening marked the
decrease of starch, the subsequent products of photosynthesis
being stored as sugar. These workers observed that carotin is
laid down in medullary rays. They remarked on the presence of
carotin, often considered a decomposition product of chlorophyll,
in close connection with such storage products as starch, sugar,
and oil. They concluded that the relations which govern the
appearance of carotin are very complicated.

Duggar (13) during some microscopic observations on the
formation of lycopersicin in tomato found that the development
of this carotinoid is preceded by the paling and ultimate disap-
pearance of the chlorophyll, and that following this change there
occurred a yellowish or orange cast in the chloroplastids. With
the disorganization of the chloroplast there appeared granules of
an orange carotin-like pigment. Accompanying these changes
typical lycopersicin crystals were formed, and as these increased
there seemed to be a decrease in the number of orange granules or
crystals.

Lubimenko (’14) believed that lycopersicin appears after the
decomposition of the chlorophyll, and especially so in those organs
which are characterized physiologically by very energetic oxi-
dations in the tissues. Lycopersicin, however, does not exist
within the chloroleucites of the fruit before the decomposition

1925]
HOWARD—CAROTINOIDS IN FRUITS 153

of the chlorophyll. Later (’16) he found during the conversion of
chloroplasts into chromoplasts that (1) chlorophyll and the ac-
companying pigments were decomposed; (2) yellow pigments ac-
cumulated and then (3) undergo alterations.

The decomposition of chlorophyll and the accompanying pig-
ments was brought about by oxidations, with the accumulation
of yellow pigments and of peroxidase, at the same time with an
increase in acidity.

MATERIAL AND METHODS

Green, partly ripe, and fully ripe fruits of the following plants
were used: Solanum Lycopersicum L., Rosa rugosa Thunb.,
Asparagus officinalis L., Capsicum annuum L., Lycium Maine
folium Mill., Я Treubii Engl. Исая were also
made on the following: yellow and red stages of Solanum Dul-
camara L. and Solanum Pseudo-capsicum L.; green and ripe fruit
of Sorbus sitchensis (Roem.) Piper, and Solanum carolinense L.;
red-ripe fruits of Rhus canadensis Marsh, Arisaema triphyllum
(L.) Schott, Citrullus vulgaris Schrad., Celastrus scandens L.
(arils), Evonymus americana L., Evonymus europaea L., Lonicera
sp., Viburnum Opulus L., Crataegus Phaenopyrum (L. f.) Medic.,
and Dracaena Godseffiana Hort.

In studying the green fruits water mounts were made of free-
hand sections, or in some cases strips of epidermis were used.
The same method was followed with the partly ripe and fully ripe
material when possible, but often the ripe fruits were too soft
to permit of such treatment. Under these circumstances smears
were made either of the mesocarp or of the juice alone. Many
of these sections were very satisfactory for study without staining.
However, to determine if chromoplasts of most fruits are really
plastids, killing and staining were resorted to.

A variety of killing and staining methods was tried, but only
some of the more satisfactory will be mentioned here. Free-hand
sections were killed by a treatment of 30 minutes in stock
chromo-acetic solution made up by Chamberlain’s (’15) formula,
washed in water for 1 hour, dehydrated in the usual grades of
alcohol, stained for 3-10 minutes in 1 per cent erythrosin (made
up in 70 per cent alcohol), run through the remaining alcohols to

[Vor. 12
154 ANNALS OF THE MISSOURI BOTANICAL GARDEN

xylol, and mounted in Canada balsam. A very satisfactory
method for staining chromoplasts was to kill in medium chromo-
acetic solution for 30 minutes, and then wash in water for 1 hour.
The sections were then placed directly into an aqueous solution
of 0.5 per cent acid fuchsin for 2-10 minutes, after which they
were transferred direct to 95 per cent alcohol, dehydrated in the
usual manner, and mounted in Canada balsam. This latter
procedure was modified to great advantage by mounting in
glycerine directly out of the stain. One of the most satisfactory
methods for staining chromoplasts of juicy fruits was to press out
on a slide first treated with albumin fixative a drop of the juice,
which was almost certain to contain an abundance of the plastids,
and these were allowed to dry 15-30 minutes. This material was
then killed on the slide for 30 minutes in the stock chromo-acetic
solution, washed for the same length of time, stained in acid
fuchsin, and mounted in glycerine as indicated above.

Paraffin sections 10 џ thick were cut with a rotary microtome
from material killed in stock chromo-acetic solution and stained
in Haidenhain’s iron-alum haematoxylin.

EXPERIMENTAL DATA AND DISCUSSION

The pepper fruit was studied in some detail. Spaces about
2 cm. in diameter were marked off on each of 3 peppers. These
fruits were then placed in an electric oven at about 25° C. to ripen.
The marked areas were examined during a period of 15 days while
the fruits were ripening, at 5-day intervals. In all 3 peppers the
results were approximately the same, so only one set of drawings
was made. These were camera-lucida drawings of free-hand
sections, mounted in water, showing the changes in shape of the
plastids, during the various ripening stages (pl. 10, figs. 1-3).
With the gradual change in color from green to yellow to orange,
and then to the red-ripe pepper color there was a gradual elonga-
tion of the plastid and in some of the red-ripe chromoplasts the
ends were actually pointed.

To determine whether the chromoplasts of red-ripe pepper were
really plastids they were stained with iron-haematoxylin. ge
of the epicarp containing many chromoplasts were killed and
stained in acid fuchsin by the method indicated above. The

1925]
HOWARD—CAROTINOIDS IN FRUITS 155

results were very satisfactory, another proof of the fact that red
pepper chromoplasts are really stainable bodies (pl. 10, figs. 4—5).
The chromoplasts stained in this material were mostly of the
elongated type. In fig. 4 the chromoplasts appeared to be in
various stages of division, and a tendency for them to collect
around the nucleus was observed.

Free-hand sections of green and yellow-orange tomatoes,
mounted in water, showed up the plastids very well. In the
green material the chloroplasts were of the ordinary shape; in the
yellow-orange a number of pigment granules beside the plastids
were visible, also a few small crystalline bodies. These pigment
granules were carotin-like in color, while the crystals had rather
a pinkish cast. The red-ripe fruit was very hard to section, and
so smears were made. In these, beautiful carmine-red, needle-
like crystals were observed. In addition, some granular pigment
material was present, but no real plastids of any kind (pl. 11,
figs. 1-3). Microtome sections made of the 3 stages of fruit
and stained with iron-haematoxylin gave well-stained plastids of
the green and yellow-orange fruit, but in the red-ripe fruit no
plastids appeared, and the crystals seen in the fresh material had
entirely disappeared, or were so distorted that they could not be
recognized. Attempts were made to stain these needle crystals
with acid fuchsin, but with no success, and indeed in the resulting
slides no crystalline forms could be found.

Free-hand sections of all 3 stages of the rose fruit showed up
the plastids very distinctly. In this fruit a very marked change
in the shape of plastids occurred. In the yellow fruit they were
elongated, in the red material much more so; in some cells they
looked almost like long irregular streaks of colored oil. To get
good results smears had to be made of the red-ripe fruit. Free-
hand sections and smears of the green, yellow-orange, and red-
ripe material were killed and stained in erythrosin as indicated
above. The green and yellow-orange plastids stained very
readily, but the red-ripe smears were not so satisfactory. It
was very difficult to get slides in which the chromoplasts retained
their exact shape during the staining process (pl. 10, figs. 6-8).

Green, yellow-orange, and red-ripe fruits of Lycium were
studied by the free-hand sections mounted in water; here a

[Vor. 12
156 ANNALS OF THE MISSOURI BOTANICAL GARDEN

gradual elongation of the plastids was observed, but the red-ripe
chromoplasts were not so markedly different as in some other
fruits. Smears of the red-ripe fruit juice were very interesting;
in these slides plastids of peculiar shape were observed, quite
different from those of the mesocarp itself. There is a striking
resemblance between these plastids and the chondriosomes of
Asparagus as pictured by Guilliermond (19), although the
chondriosomes of Guilliermond are much smaller. Smears, also
free-hand sections, of the red-ripe Lycium fruits killed and stained
in acid fuchsin showed plastids of very much the same shape as
those in the fresh material (pl. 12, figs. 1-5).

Great difficulties were experienced in attempts to make sections
of green Asparagus fruits, this being due to a hard, bony epicarp.
Smears of the mesocarp of fruits of 3 degrees of ripeness mounted
in water showed up plastids very well. Here again the red-ripe
chromoplasts were very unlike those of the green and yellow-
orange preparations. These plastids also are very much like
Guilliermond’s chondriosomes of Asparagus referred to above,
but several times larger. Smears made from the red-ripe meso-
carp, also from the juice, were stained in acid fuchsin, and in these
preparations much-better stained and more normally shaped |
chromoplasts were obtained from the juice smears than the
mesocarp. ‘There were some peculiarly shaped bodies in the
red-ripe juice due, no doubt, to disintegration processes (pl. 12,
figs. 6-9).

In green fruits of Aglaonema studied by means of free-hand
sections mounted in water, very distinct and typical chloroplasts
were observed. In the yellow fruit the plastids seemed to have
disintegrated, with the formation of carotin-colored crystals and
granules. In the red-ripe stage carmine-red crystals were
present, these being smaller but very much the same in shape and
appearance as those present in red-ripe tomato. No attempts
were made to stain any of this material (pl. 11, figs. 4—5).

Yellow-orange and red stages of Solanum Dulcamara (pl. 11,
figs. 6-7) and Solanum Pseudo-capsicum (pl. 13 fig. 9) and green
and ripe stages of Sorbus sitchensis (pl. 13, figs. 1-2) were
studied by free-hand sections and smears mounted in water.
Nothing unusual was noted about the plastids of Solanum Dul-

1925]
HOWARD—CAROTINOIDS IN FRUITS 157

camara or of Sorbus, but those of red-ripe Jerusalem cherry were
very much elongated with long-drawn-out points. In ripe S.
carolinense fruit no plastids were ever observed; the pigment,
although a carotinoid, was present in oily globules. Fair results
were obtained by staining smears of Solanum Dulcamara and red-
ripe fruits of Sorbus in acid fuchsin. Smear mounts made from
the juice of red-ripe Jerusalem cherry, stained in acid fuchsin,
showed very well-stained chromoplasts, exactly like those of the
fresh material. In the red-ripe fruit of Rhus canadensis nothing
like a plastid was ever found, just a mass of oily globules as in
Solanum carolinense.

In red-ripe fruits of Arisaema, Celastrus, Viburnum, Crataegus
(pl. 13, figs. 3-8), Evonymus americana (pl. 11, fig. 9), E. euro-
paea (pl. 14, fig. 2), Lonicera (pl. 14, figs. 4-5), and Dracaena (pl.
11, fig. 8) definite plastids were found in smears mounted in
water. Especially good results were obtained by staining juice
smears of Arisaema and Celastrus in acid fuchsin. The stained
chromoplasts of Celastrus were of exactly the same shape and
size as the unstained (pl. 18, fig. 3). Hvonymus europaea was
very interesting because the plastids were so very small in the red
arils and very difficult to find, no doubt due to the fact that they
disintegrate very early and the pigment is present in oily glob-
ules, much the same as in Rhus and Solanum carolinense (pl. 14,
fig. 2).

The work on Е YE was rather curtailed owing to lack
of material, only red-ripe fruit being available. Smears were
made of the mesocarp near the epicarp, where it remains green
even in the ripe fruit. In slides made from this region of the
mesocarp oval plastids were observed, some of which were green
and some pink. In smears made from the red-ripe pulp many
carmine-red needle-like crystals with uneven ends were observed,
and these were very similar to those in tomato and Aglaonema
- 14, figs. 6-7).

deter e whether t the сі lasts of a number of fruits are
i ‚равна these decolorization experiments were undertaken.
At first thought it seemed an easy matter to remove the color
from a chromoplast. Carbon bisulphide had long been known as a
ready solvent for carotinoids, especially lycopin (lycopersicin)

à [Vor. 12
158 ANNALS OF THE MISSOURI BOTANICAL GARDEN |

and carotin. As these two carotinoids were present in larger
proportion than xanthophyll in the fruits studied, carbon bi-
sulphide was the first solvent tried. The first thing to be done
was to get rid of all possible water from the tissue, as water and
carbon bisulphide are miscible only in small proportions. Small
pieces of sections of red-ripe tomato, Lycium mesocarp, and
Evonymus europaea arils were placed in 25, 50, 75, and 100
per cent acetone, 30 minutes for each grade. The color came out
and appeared in drops over the surface of cells. The acetone
was allowed to evaporate, after which, where these sections were
placed i in CS,, much color was drawn out over the tissue but noth-
ing could be determined in regard to the presence of plastids.
Several smears of red-ripe Lycium juice containing plastids were
placed in each of the following reagents, acetone, carbon bisul-
phide, chloroform, benzole, and petrol ether, and the effect of
each reagent observed. The slides standing in petrol ether and
those in carbon bisulphide were the only ones that gave up any
color to the liquid. On none of the slides could there be any
changes noticed in the chromoplasts. The pigment went out of
the tissue of red-ripe Lycium when placed in warm chloroform
for 12 hours, but the tissue was too badly broken up to be of any
use іп a study of plastids. Sections of red-ripe Lycium were
dehydrated by means of the usual grades of alcohol. The
color almost disappeared in absolute alcohol, but the plastid
structure was not clear.

After all these unsatisfactory results a method was tried which
proved very successful. For the extraction of all the chlorophyll
and carotinoid pigments in green leaves Tsweet ('06) recom-
mended petroleum ether and absolute aleohol (1 per cent absolute
alcohol in petroleum ether for dry leaves and 10 per cent for fresh
leaves). This method was applied to fruit tissue in these experi-
ments. Small pieces of the tissue were placed in vials contain-
ing petroleum ether and 1 per cent absolute alcohol and allowed
to stand for 48 hours, after which they were mounted in glycerine
and examined with the microscope. Red-pepper tissue gave very
good results with this treatment. On these slides colorless
plastids very much like those of the chloroplasts showed up in the
tissue. Fruits with very small chromoplasts gave fair results
but not so satisfactory as the pepper.

1925]
HOWARD—CAROTINOIDS IN FRUITS 159

From these studies the writer feels sure that at least in most
ripe fruits studied the amorphous pigment in a lipoid substratum
is contained in a stroma or kind of plastid. At least there is
some sort of an accumulation of the cytoplasm at these points.
In some fruits great diffieulty was encountered in staining the
chromoplasts. The best success was obtained by staining
smears of juice. Mention should be made of the fact that after
killing in chromo-acetic acid these chromoplasts in most of the
fruits appeared as colorless bodies of the original form and shape.
Very good slides were made showing stained chromoplasts of
red pepper, Lycium, Solamum Pseudo-capsicum, Arisaema, Ce-
lastrus, while rose and Asparagus gave only fair results. The de-
eolorization experiments seemed to show without a doubt that
the so-called chromoplast is a definite body or stroma and very
much like the chloroplast. These so-called chromoplasts increase
by direct division, which is another proof of their plastid-like
nature. In all of these fruits there was a variation in plastid
shape from globose to oval in the chloroplast, while the chromo-
plasts were elongated, irregular forms. Asparagus no doubt
showed the most remarkable plastids, for, as has been previously
mentioned, their similarity to mitochondria was great.

Lycopersicin crystals are present in the natural red-ripe fruit
of tomato, watermelon, and Aglaonema. Proof of the fact that
the pigment here is really in a crystalline state is as follows:
(1) These crystals disappeared when any attempt was made to
stain them, (2) no re-crystallization could be obtained by means
of the alcoholic-potash method of Molisch. These 3 fruits contain
a large percentage of lycopersicin, so that a generalization might
be assumed to the effect that lycopersicin, at least, is usually
found in a natural, crystalline form. Solanum Dulcamara might
seem to destroy the value of this generalization because the red-
ripe fruit in its natural condition shows no crystals, but exhibits
plastids of very much the same shape as the chloroplasts. How-
ever, it is quite possible that the pigment in S. Dulcamara is not
lycopersicin. In Rhus there is another interesting condition,
which is that the pigment of the red-ripe fruit appears in oil
globules, and at no time was I able to see plastids or crystals.
In the 6 kinds of natural red-ripe fruit in which lycopersicin was

[Vor. 12
160 ANNALS OF THE MISSOURI BOTANICAL GARDEN

found it appeared in 3 different conditions, namely: amorphous in
a lipoid substratum contained in a stroma; crystalline, free in the
cytoplasm; and in oily globules.

It is interesting to note that within the genus Solanum the
fruit carotinoids appear as crystals in tomato, in plastids in
Solanum Dulcamara, and as oil globules in Solanum carolinense.
Again, in Evonymus americana there are very definite chromo-
plasts, in Evonymus europaea the pigment is in oil globules.
Kars the genus Asparagus we have carotinoid pigments in

A. officinalis, while in A. Sprengeri the fruit pigment is entirely
anthocyanin. The same pigment conditions occur in the genus
Rhus, for the pigment of R. canadensis is a carotinoid, while in
R. glabra the pigment is anthocyanin.

А COLORIMETRIC METHOD FOR THE DETERMINATION OF THE
RELATIVE AMOUNTS OF PIGMENT IN FOUR VARIETIES
or ToMATOES

MATERIALS AND METHODS

The purpose of this phase of the investigation was to work out
a method by which a colorimeter of suitable type might be used
as a means for making quantitative determinations of the pig-
ments contained in tomatoes. Since the principal pigment in
tomato is lycopersicin, and CS, is a ready solvent for this car-
otinoid, this solvent was used in making the extractions for ex-
amination with the colorimeter. It was necessary to devise
methods for drying the tomato pulp, since no previous work
had been done along this line.

Four varieties of tomatoes were used, Ponderosa, Excelsior,
Globe, and Dwarf Champion. These were grown to maturity
in the experimental plots at the Missouri Botanical Garden.
Three methods were tested for drying the pulp, and the one
yielding the most deeply colored extract, as determined by the
colorimeter, was considered the best. For these preliminary de-
terminations no special varieties were employed; however, 2
classes of fruits were used: (1) red-ripe fruit; (2) red-orange
fruit. Tomatoes from each of these classes were selected, then
skinned by plunging into boiling water for a few minutes. From

1925]
HOWARD—CAROTINOIDS IN FRUITS 161

the central areas of these tomatoes slices were cut transversely
and weighed out in 20-gm. portions, each being carefully crushed
up in a mortar and spread out on filter-paper. From this point
3 distinct procedures were followed: (1) Using a dry-heat method,
the pulped fruit on filter-papers, 2 containing red-ripe tomato pulp
and 2 red-orange, were dried for 6 hours by means of an electric
fan, then brought to total dryness in a gas oven at a tempera-
ture varying from 40 to 50° C. (2) With the alcohol method
2 filter-papers of each of the 2 lots, as before, were successively
covered with 30, 50, and 70 per cent alcohol, remaining covered
with each grade for an interval of 30 minutes, after which the
alcohol was filtered off by means of a filter pump. "Then 95 per
cent aleohol was poured on and after evaporation the pulp was
entirely dry. (3) Using the acetone method, the same number
of filter-papers were successively covered with 50, 70, and 85 per
cent acetone, remaining covered with each grade for 30 minutes.
Then 100 per cent acetone was added and, as in the alcohol
method, upon evaporation the pulp was entirely dry. From
each filter-paper the pulp was carefully removed and extracted
in glass-stoppered bottles with 25 cc. of CS. After 4 days
these extracts were examined with a Duboscq micro-colorimeter.
The method followed will have to be described in some detail,
as a colorimeter of this type is not ordinarily used in this
manner.

One of the lower eups of the colorimeter was filled about one-
third full with the extract from the red-ripe acetone-dried ma-
terial, and the other was filled with an equal amount of the extract
from the red-ripe alcohol-dried material, then the colors were
matched and a reading taken on each scale. Two ''day-light"
globes were used as a source of illumination. A match was then
made between the alcohol-dried and heat-dried material and a
reading on the scale recorded. By this same method the extracts
of the 2 kinds of tomato pulp dried in the 3 different ways were
matched, and readings recorded in table r. Dilutions of 50 per
cent and 5 per cent were made of all the extracts matched with
the colorimeter, and readings taken. The smaller the scale
reading the greater the concentration of the extract, therefore
the more of extractable lycopersicin per unit of tomato pulp and

[Vor. 12
162 ANNALS OF THE MISSOURI BOTANICAL GARDEN

hence the better drying method. The results are recorded in
table 1.

TABLE I
COLORIMETRIC READINGS IN MM. MADE WITH A DUBOSCQ INSTRUMENT
TET 50 cc. extract, 5 cc. extract,
No dilution 50 cc. CS2 95 cc. CSa

Асе- Ace- Ace-

| Alcohol |Heat Бл Alcohol| Heat iond Alcohol| Heat tone
Red-ripe tomato| 2.5 a 7.8 | 29 ft В: LES : 2.0 | 1.7 | 1.5
-orange “ 8.4 1.24 0.8 2.6 0.9 1 0.8 1.8 0.9 10.5

From the results of this experiment acetone is the best drying
agent for tomato pulp, since after drying by this method the
most deeply colored extracts were obtained, as indicated by the
small readings on the scale. "With this problem of the drying
determined, so far as the 3 methods described were concerned,
the main part of the investigation was undertaken. It was ac-
cordingly required to determine the availability of this colori-
metric method in estimating the relative amounts of carotinoids
in these 4 varieties of tomatoes. In all of these experiments only
red-ripe fruits were used.

EXPERIMENTAL DaTA AND DISCUSSION

Experiment 1.—Red-ripe fruits from the 4 varieties of tomato,
Globe, Ponderosa, Excelsior, and Dwarf Champion, were skinned,
crushed in a mortar, allowed to evaporate 12 hours in evaporating
dishes, then placed on filter-papers in Buchner funnels and 5-cc.
amounts of acetone poured on at 30-minute intervals until 25 cc.
had been used. These filter-papers were then placed in evapor-
ating dishes where in a short time the pulp became entirely dry
and ready for extraction after being ground in a coffee mill. One
gram of each kind of dried, ground material was placed in a
ground-glass-stoppered bottle and extracted with 10 ce. CS:
for 4 days. Colorimetric determinations were then made by the
method indicated above. The results are recorded in table п.

1925]

аа Та Л READINGS IN

HOWARD—CAROTINOIDS IN FRUITS

TABLE II

163

MM.; ACETONE-DRIED RED-RIPE FRUIT,
. OF MATERIAL EXTRACTED 4 DAYS WITH 10 CC. C8:

S 50 cc. extract,
No dilution 50 сс. C8,
rf
Excelsior | Ponderosa PE tates HE Globe | Excelsior | Ponderosa UP dent eh Globe
0.50 0.70 0.95 1.07} 1.40 1.90 3.00 3.10
0.42 0.60 0.80 0.90 0.90 1.20 1.83 1.90
0.42 0.60 0.80 0.90 | 0.60 0.80 1,23 1.27

Experiment 2.—The same material and methods were used as
in experiment 1 (except no Dwarf Champion material was avail-
able). The results are recorded in table пт.

TABLE III

COLORIMETRIC READINGS IN MM.; ACETONE-DRIED qme. FRUIT,
5 GM. OF MATERIAL EXTRACTED 4 DAYS WITH 10

50 cc. extract,
50 ec. СӘ»

Excel-
sior

30 cc. extract,

No dilution 70 ec. СӨ»

coe Globe | Ponderosa
1.02
0.86

Globe
1.25 | 1.56 |
0.80 | 1.00

Experiment 3.—Juice from each of the 4 varieties of tomatoes
being tested was pressed out, filtered, and the precipitate dried
in a gas oven at 60° C. One-tenth of a gram of each kind of dried
material was extracted in ground-glass-stoppered bottles with
10 ec. CS, for 4 days. Colorimetric determinations were made
and recorded in table ту.

Globe Ponderosa

per E Ponderosa

4.32
2.13

1.03 1.42
0.865| 1.20

0.63
0.30

1.32
0.63

0.53
0.25

TABLE IV
COLORIMETRIC READINGS IN MM.; JUICE PRESSED OUT, FILTERED, DRIED

IN OVEN AT 60° C., .1 GM. OF MATERIAL EXTRACTED WITH 10 CC.
CS: (EXTRACTED 4 DAYS)
No dilution = д =
Dwa
Excelsior | Ponderosa Champion Globe | Excelsior | Ponderosa Champion Globe
1.70 3.50 8.30 |13.10| 3.50 8.40 29.80 137.80
0.88 1.80 4.30 .80| 0.41 1.00 3.50 4.50
0.84 1.60 4.00 6.20| 0.57 1.40 4.90 6.30

[Vor. 12
164 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Experiment 4.—Juice was pressed out from Excelsior, Globe,
and Ponderosa, then centrifuged, and the pigment was dried in à
dehydrater. Seven-tenths of & gram of each kind of material
was placed in a glass-stoppered bottle and each extracted with
20 сс. of СЪ, for 5 days. Colorimetric determinations were made
and recorded in table v.

TABLE V
COLORIMETRIC READINGS IN MM.; JUICE CENTRIFUGED AND DRIED IN

DEHYDRATER,.7 GM. OF MATERIAL EXTRACTED WITH 20 CC. CS
(EXTRACTED 5 DAYS)

ou 50 ec. extract, 30 cc. extract,
меи 50 ce. СЗ: 70 ее. CS2
Excel- Excel- Excel-
2 Globe| Ponderosa sus Globe| Ponderosa icis Globe| Ponderosa

0.38 | 0.73 | 1.16 0.13 | 0.83 | 1.33 | 1.02 2| 1.16| 2.4
0.33 | 0.63 1:00 0.80|0.66| 1.03 |0.05|0.06| 1.35

From these experiments Excelsior tomatoes seem without a
doubt to contain the largest amount of lycopersicin of any of the
varieties experimented with. There is too much irregularity of
results, as may be seen from the tables, to permit of any very
definite statement respecting the relative pigment content of the
other varieties. This is perhaps an indication of the fact that
these 3 varieties, namely, Ponderosa, Dwarf Champion, and
Globe, all contain very nearly the same amounts of pigment;
and this colorimetric method is only a rough means of deter-
mining relative amounts.

Experiment 5.—In an attempt to make a color standard for use
in quantitative determinations the question arose as to the sta-
bility, with elapsed time, of the color of dried tomato pulp; ac-
cordingly a time-factor test was made. The method followed
was to find some dye to match a СЗ, extract of the dry, ground-up
tomato pulp. One gram of this material was allowed to extract
for 4 days in 10 сс. of CS., then 15 сс. of this extract were diluted
with 3.5 се. of CS.. A very good match was obtained for this ex-
tract by using Medalia’s ('20) methods for colorimetric determin-
ations of H-ion concentration. Three cc. each of N/10 H,SO, and
N/10 KOH were placed in tubes to each of which 9 drops of
methyl red were added. These tubes, one in front of the other

1925]
HOWARD—CAROTINOIDS IN FRUITS 165

with a third one in the line containing CS., were matched with
another set of 3 tubes. In this second set, the first tube con-
tained the pigment extract, the second pure CS., and the third
Н.О. These tubes were placed in an ordinary wooden com-
parator. 'This comparison was made December 20.

On March 30 an extract of the same material of the same
strength was matched by the same method. To get a good
match this time 7 drops of methyl red had to be added to the
H.SO,, and 20 drops to the KOH. The extract now was more of
an orange-red tint than a true red.

Again, on May 22, an extract of the same strength was made
from the same material, but this time only 6 drops of methyl red
were added to the H.SO, and 20 drops to the KOH to give a good
match. The ground material from which the samples were
taken for testing had been kept in a paper envelope in a drawer
away from thelight. The conclusion is that dried, ground tomato
pulp changes in respect to pigment color very gradually, if kept
in the dark, during a period of 5 months. However, the lycoper-
sicin carmine-red shade disappeared much more rapidly than the
red-orange. |

DETERMINATIONS OF THE PRESENCE OF CERTAIN PIGMENTS IN
ARIOUS FRUITS

INTRODUCTION

Definite scientific knowledge of the kind and quantity of
earotinoid pigments found in fruits is rather limited, as may be
seen from a survey of the literature. The main reason for this
is the great difficulty with which the pigments are obtained in a
pure state. There are 2 general methods for quantitative esti-
mations, one the gravimetric, the other the colorimetric. The
gravimetric method is unsatisfactory for most workers because, in
any case, the amount of pigment is relatively small, hence the
necessity for such large amounts of the fruit employed. Again,
the large proportions of colorless impurities, which are very dif-
ficult to separate from the pigments, and the rapidity with which
these pigments oxidize cause trouble. The colorimetric method
is better adapted for general use, because smaller amounts of

[Vor. 12
166 ANNALS OF THE MISSOURI BOTANICAL GARDEN

material can be used, but it doesn't seem entirely satisfactory
on account of the difficulty with which the standards of really
pure carotin and xanthophyll can be prepared and kept in such a
state. Substitutes, such as 25 per cent alizarin in chloroform,
or a 0.2 per cent aqueous solution of К,Ст,О,, are used to good
advantage, because of their stability. Willstátter and Stoll (13),
using K.Cr.O; solutions, established a certain set of relations
between the standard 0.2 per cent K.Cr.0, and amounts of
carotinoids by means of which quantitative determinations of
carotin and xanthophyll might be made on any fruit.

In consideration of these rather unsatisfactory results, no actual
quantitative determinations were carried out. However, at-
tempts were made to use the colorimeter determinations in con-
nection with a color eode, to estimate the relative quantities of
pigments in certain fruits. Spectroscopic determinations were
made of a number of fruit extracts, and artificial crystallization
of these pigments within the tissues was accomplished.

Throughout this investigation the term lycopersicin suggested
by Duggar (13) has been used in place of the older term lycopin.

SURVEY OF LITERATURE

Citrullus vulgaris.—The first work done on the identification of
the watermelon pigment was that of the De Negris (79) who iso-
lated the pigment from the flesh of the fruit and called it rubidin.
Courchet ('88) determined that the natural crystals were the
same in form and color as those of tomato. Upon agitation with
ether the pulp gave almost instantly a beautiful yellow-orange
solution, which deepened rapidly in color in proportion to its
concentration. The crystals which formed voluminously upon
evaporation were needle-like in form or, more frequently, long,
slender, carmine-red sheets, which were often contained in sheaths
Montéverdé and Lubimenko (13) further confirmed the presence
of lycopersiein in watermelon. Lubimenko (14) gave spectro-
scopic determinations for petroleum ether and C$, extractions
of the pulp. Nevertheless, the work on this pigment has been
very limited, and about the only — = is the fact
that lycopersicin is present in large am

Capsicum annuum.—Pabst (92) ночи to eitis the pigment

1925]
HOWARD—CAROTINOIDS IN FRUITS 167

in pepper with carotin, but was unsuccessful. Kohl (’02), by
means of spectroscopic examinations, was able to find no differ-
ence between pepper and carrot pigments. Tschirch (04) found
the following absorption bands for pepper: (1) 517.0-501.0 џи;
(2) 486.0-467.0 uu; (3) 458.0-439.0 uu, no end absorption.
Duggar (113) obtained the characteristic, bright red lycopersicin
color in a СЗ, extract, which gave the distinctive lycopersicin
spectrum, 2 bands in the green and 1 in the blue, though no figures
were given for the limits of the bands. The name lycopersicin
was suggested by him to take the place of lycopin, because lycopin
has long been employed to designate a resinoid compound derived
from the bugle-weed. Lubimenko ('14) called the pigments of
pepper purified by various washings with 95 per cent alcohol a
lycopinoid, because its absorption bands were intermediate
between pure lycopersicin, which relation the following data show.
Daucus Carota, which is considered to contain practically no
pigment but carotin, gave a spectrum with 3 bands: (1) 533.0-
508.0 uu, (2) 489.0-472.0 uu, (3) 455.0-445.0 uu. Pepper showed
3 bands: (1) 560.0-540.0 uu, (2) 530.0-500.0 uu, and (3) 495.0-
480.0 uu. Areca Alicae, which he considered to give a very char-
acteristic lycopersicin spectrum, gave the following bands: (1)
565.0-540.0 uu, (2) 520.0-500.0 ши, (3) 480.0-470.0 uy. Van
Wisselingh (15), by means of the alcoholic-potash method of
Molisch (713), obtained the pigment in a crystalline form in the
tissue of the fruit. These crystals were mostly in the form of
aggregates, but others varying in form were present, and they
ranged in color from red (Klincksieck et Valette ('08), Code des
Couleurs, pp. 16, 21, also 76, 81, and 106, 127) to orange-red and
orange. No definite qualitative or quantitative determinations
for the pigments contained in pepper have been made.

Solanum Lycopersicum.—The red tomato pigment has been
studied by a great many workers, and from definite chemical
determinations we know now that it is identical neither with
carotin nor xanthophyll. Millardet (76), the first worker оп
tomato pigments, soon recognized that it was not identical with
either the red or yellow pigments of other fruits. A. and G. de
Negri (79) considered the pigment identical with ‘‘rubidin”
which they had isolated from watermelon. Ehring (96), Ar-

[Vor. 12
168 ANNALS OF THE MISSOURI BOTANICAL GARDEN

naud (’85), and Passerini (790) all believed the tomato pigment to
be carotin, as did also Tammes (00) and Kohl (02). Schunck
(03) suggested the name lycopin for the pigment which he was
able to extract from the tomato. By difference in spectroscopic
analysis and solubility, he decided this pigment was different
from his chrysophyll. Montanari ('04) submitted the pure crys-
talline lycopersicin to chemical analysis and obtained the first
evidence of the true relation between lycopersicin and carotin.
He obtained an average composition of C — 84.14 per cent and
H = 10.88 per cent. He considered this to correspond suf-
ficiently closely to Arnaud’s formula СН» for carotin, which had
not been disproved at that time. Molecular weight determina-
tions in benzene, using the cryoscopie method, gave a value of
698, from which he decided it must be a dicarotin with the
formula С.Н 76+

Willstatter and Escher (’10) showed that lycopersicin was iden-
tical in general composition and molecular weight with carotin,
—differing only in degree of solubility in certain solvents, in the
position of the absorption bands, in the form and color of the
crystals, and in the melting point. They determined lycopersicin
to be a true isomer of carotin. Out of 74 kgms. of tomato con-
serve (canned tomatoes) they obtained 11 gms. of crystalline
lycopersicin. Crystals of carotin were also obtained in small
amounts as a by-product. The analyses and molecular weight
determinations agreed very closely with the theoretical deter-
minations made by Willstatter and Mieg (07) for carrot. Mon-
téverdé and Lubimenko (’13) have confirmed these figures. From
the above work the principal pigment in tomato has been shown
to be lycopersicin, although carotin is present in small amounts.
Quantitative determinations for lycopersicin have been made.

Rosa rugosa.—Some variation occurs in the pigmentation of the
different species of rose as indicated by the literature. Tammes —
(00), the first worker, found the pigment constantly gave a blue
color with H.SO,, HCI, HNO,, phenol, and bromine water, thus
showing the presence of carotinoids. Kohl (02) noted the oc-
currence of carotin in Rosa sp. Tschirch (04), employing the
method of capillary analysis with Rosa canina, obtained a dark
orange zone, and his spectrum analysis gave the following bands:

1925]
HOWARD—CAROTINOIDS IN FRUITS 169

(1) 492.0-475.0 uy, (2) 462.0-445.0 uu, (3) 439.0-418.0 pu. Mon-
téverdé and Lubimenko (713) isolated lycopersicin crystals from
the dried fruit pulp, but they regarded this as a minor constituent
of the pigments. Lubimenko (’14) reported the presence of lyco-
persicin in the chromoleucites in the form of small needle crystals,
and included the rose in his list of fruits containing lycopersicin.
Van Wisselingh (15) by means of the Molisch (713) alcoholic-
potash method was able to obtain crystals of carotinoids from
Rosa rugosa.

Evonymus.—Courchet (’88) found the arils of Evonymus to
contain fusiform, orange-red chromoleucites in which the pig-
ment was present in the form of needle-like crystals which he
called crystallites. The usual blue color reactions for carotinoids
with Н.50, and HNO, were obtained. Tschirch (04), in his
capillary colorimetric analysis of the alcoholic extract of Е.
europaea, obtained several yellow to red-orange zones. Spectro-
scopic analysis gave the following bands: (1) 495.0-472.0 uy,
(2) 467.0-439.0 џи, (3) 430.0—415.0 uu, with no end absorption.
Tammes (700) examined the arils of E. latifolius and found that
the plastids gave the characteristic blue color with H.3O,,
HNO., НСІ, and bromine water. Lubimenko ('14) classed Е.
japonica as a fruit containing lycopersicin.

Solanum Dulcamara.—Thudichum (’69) classed the pigment of
this fruit as a lutein. Hartsen (’73) found a red substance in
granular form, crystallizable in tablets, which was the same sub-
stance that he found in T'amus communis and Asparagus officinalis.
Tammes (’00) obtained crystals by the Molisch (^13) alcoholic-
potash method. Lubimenko (’14) found the chief pigment to
be lycopersicin. Van Wisselingh (’15) also obtained crystals by
means of the Molisch alcoholic-potash method.

Sorbus.—The first worker on these fruits was Tammes (’00)
who reported the presence of carotinoids, as determined by the
characteristic reaction with H.SO, HCL, HNO,, phenol, and
bromine water. By means of the Molisch alcoholic-potash
method he was able to obtain yellow-red, pointed crystals,
single, or in small bundles. Kohl (02) reported the presence of
carotin in Sorbus Aucuparia and S. Aria. Van Wisselingh (715),
by various methods, found 3 types of crystals in the pericarp, but
he did not undertake to classify them.

[V or. 12
170 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Asparagus.—Thudichum (769) classified the pigments found
in this fruit with the luteins. Hartsen (73) found some red
substances in the form of granules that were crystallizable in the
form of tablets. These pigments were insoluble in Н.О, slightly
soluble in alcohol and ether, but especially so in benzene.

Aglaonema.—Tammes (700) determined the presence of caro-
tinoids in A. commutatum, by means of the characteristic color
reactions with H.SO., HCl, HNO,, phenol, and bromine water.
Van Wisselingh (15) obtained by using the Molisch alcoholic-
potash method, characteristic carotinoid crystals. Crataegus
pigments have been very little experimented with, though
Thudichum (769) classed them as luteins.

Other fruits.—On Solanum Pseudo-capsicum very little work
has been reported. Kraus (72) observed red and orange-yellow
pigment forms in fruit flesh of Jerusalem cherry, and also made
spectroscopic determinations of the alcoholic extract. Lycium
was found to contain carotinoids by Courchet (’88). In Lonicera,
Schimper (’83) stated that he found red and orange-yellow crystals
in the plastids. Courchet ('88) obtained what he called crystal-
lites in the fruit of Lonicera Caprifolium. Molisch (96) and
Kohl (02) obtained crystals by the alcoholic-potash method of
the former worker. Viburnum Opulus was worked on by Van
Wisselingh (15), who obtained carotinoid crystals by using the -
Molisch alkaline method but no classification was made of them.
Fritsch (’84), who alone has worked with Celastrus scandens, de-
scribed the shape of the chromatophores and observed that they
were colored blue and then dissolved with concentrated H.SO,,
also that iodine colored them blue-green. In Cucumis Melo,
Courchet (’88) examined the character of the pigments in the
plastids, then extracted the pigment and obtained, upon re-
crystallization, rhombohedral crystals and a few trapezoidal
forms orange-red in color. No work has been reported upon the
pigments of Arisaema, Dracaena or Rhus. In Momordica char-
antia fruit Duggar (’13) obtained spectroscopic and physiological
proof that the principal pigment of the carpels is carotin, but that
the aril is characterized by lycopersicin. Courchet (’88), from a
recrystallized ether extract of the aril of Momordica balsamina,
obtained carmine-colored needles, which he found to be identical

1925]
HOWARD—CAROTINOIDS IN FRUITS 171

in form and color with those obtained from tomato and water-
melon. The Toblers (10а) and Duggar (13) have confirmed
Courchet's observation that the red pigment in the aril of Momor-
dica balsamina is lycopersicin.

From this review of the literature it is evident that with the
exception of a few fruits such as tomato and pepper very little is
definitely known concerning just what carotinoids and in what
proportions they are present in fruits.

Daucus Carota.—The first yellow plant pigment to be isolated
in crystalline form was carotin from carrots. This pigment was
first described by Wachenroder (1826) and called carotin by this
worker. This investigation serves as the starting point for our
knowledge of the properties as well as the nomenclature of the
carotinoids. Vauquelin and Bourchadat (1830) were the next
workers, but not until the investigations of Zeise (47) was carotin
isolated from carrots in amounts sufficient for analysis. He dis-
covered its ready solubility in CS. and made the first analysis of
carotin, giving it the formula C.H, with a melting point of 68? C.
This formula was not accepted, however, due to the authority of
the next experimenter, Husemann (61), who worked extensively
with carrots. His methods were to press the juice from finely
grated carrots, and then add weak Н.50,, thus throwing down a
coagulum, which was partly dried and extracted with 80 per cent
СН.ОН, then this residue dried and extracted with CS. The
_ earotin crystals were then precipitated by the addition of ab-
solute alcohol, these being purified by washing with hot 80 per
cent ethyl alcohol and finally with absolute alcohol. Husemann
was the first to show the unsaturated nature of the carotin
molecule, although he regarded the chlorine and iodine derivatives
which he was able to make as substitution products. From these
analyses he proposed the formula C,,H:.O, and his figures were
accepted over those of Zeise (’47).

Arnaud (’85), the next investigator, pressed the juice from
grated carrots and precipitated the pigment by the addition of
lead acetate, then dried this precipitate in vacuo and extracted
with CS, Тһе residue was washed with cold petroleum ether,
and the pigment purified by crystallization from CS,, with ab-
solute alcohol, and it was then allowed to recrystallize spontane-

[Vor. 12
172 ANNALS OF THE MISSOURI BOTANICAL GARDEN

ously from cold petroleum ether. About 3 gms. of crystals per
100 kgms. of carrots were obtained in this way. His analysis of
freshly prepared crystals showed an average composition of
88.67 per cent carbon and 10.69 per cent hydrogen, thus defi-
nitely proving the correctness of Zeise's statement that carotin
is a hydrocarbon. This worker for the first time prepared the
crystalline iodine derivative of carotin. The elementary com-
position of this product and the composition of pure carotin led
Arnaud to give the formula СН; to carotin and C..H;sI: to the
iodine derivative. Kohl (02) did a great deal of detailed work
on the chemical and physical properties of carotin. His own
analyses of the crystalline pigment in carrot gave unsatisfactory
results, so he accepted Arnaud's formula as correct. His pub-
lication contains an extensive and valuable list of plants and
animals containing carotin

Schunck (703) РИ а very good yield of crystalline pig-
ment by drying the juice of grated carrot and dissolving in ether.
It crystallized in a form very much like the chrysophyll of Hartsen
(73) and Schunck (701), which was no doubt identical with
carotin. He made spectroscopic examinations in which the
bands corresponded very closely with those of chrysophyll.
Tschirch (04) cut into pieces carrot which had been soaked in
water for a day to remove all sugar, then cooked in water, covered
with alcohol for a day, and finally dried by gentle heating. These
dried pieces were then ground in a powder mill and extracted in
a Soxhlet with petroleum ether of boiling point 60° C. After
evaporation of the petroleum ether the residue was crystallized
out of alcohol. Later he found it unnecessary to cook the carrot
slices, and the soaked pieces were treated immediately with
petroleum ether in a Soxhlet. This ethereal extract was allowed
to evaporate spontaneously. After a few days the walls of the
glass container were covered with the characteristic, dark red to
orange, metallic carotin crystals. These were washed with ethyl
alcohol and dried under reduced pressure. The alcoholic solu-
tions gave a spectrum with three absorption bands: (1) 487.0-
470.0 uu, (2) 457.0-439.0 uu, (3) 429.0-417.0 uu. Solutions in
chloroform gave two bands: (1) 507.0-486.0 ши, (2) 473.0—454.0 ug.
Willstátter and Mieg (’07) definitely settled the composition of

1925]
HOWARD—CAROTINOIDS IN FRUITS 173

carotin and proved its identity with the pigments in leaf chloro-
plasts. They prepared the crystalline material in large amounts,
making several extractions with CS., which were in alcohol and
crystallized from petroleum ether. Their data show a mean
ratio of C:H of 1:1.406 for which the simplest formula is
C.H;. Molecular weight determinations in chloroform and СЗ»,
using the ebulloscopie method, showed an average of 536, which
corresponds exactly with the formula (C,H;). ог C,,H.,—which
is now accepted as the empirical formula for carotin. Euler and
Nordenson ('08) also isolated carotin from carrots in quantities
sufficiently large for analysis. "Their results confirmed the Will-
státter and Mieg (07) formula. The purified crystals were also
found to eontain xanthophyll, which was identified by the color
of the crystals and their differences in solubility. Escher (09),

in a very comprehensive paper, gave minute directions for making
quantitative determinations of carotin in carrot, methods in
which he used both large and small quantities. He confirmed
the work of Willstátter and Mieg, but was unable to determine
the structural formula, because of impurities in the crystalline
material, which he was unable to eliminate. Palmer and Eckles
(14) have shown the presence of xanthophyll in the carrot
root by the Tsweet (06) chromatograph method. Van Wisselingh
(15), using the microchemical methods, was unable to show the
presence of xanthophyll crystals in the carrot root. From a
survey of this work it is evident that carotin and xanthophyll are
the carotinoids in carrot, also that the empirical formula for
carotin has been definitely established, and that some quan-
titative determinations have been made.

MATERIALS AND METHODS

The fruits used were Lonicera, Lycium, Citrullus vulgaris,
Cucumis Melo, Aglaonema, Arisaema, Solanum Dulcamara,
Viburnum, Celastrus (carpels and arils), Solanum Lycopersicum,
Rosa, Asparagus, Evonymus americana, and E. europaea, Sorbus,
Rhus, Capsicum, Solanum Pseudo-capsicum, and Crataegus. These
were all fully ripe fruits.

For drying the fruit there was used a vegetable dehydrater
constructed by Dr. G. T. Moore after one designed by the U. S.

[Vor. 12
174 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Department of Agriculture (Pugsley, 17). For the colorimetric
determinations, the same colorimeter was used and the same
methods followed as in a previous part of the investigation. All
extractions were made with CS., the material being first ground
in a coffee-mill or mortar. Extractions were then made in
ground-glass-stoppered bottles. Klincksieck and Valette’s (’08)
"Code des Couleurs’ was used for color comparisons.

The instrument used for spectroscopic analysis was an A.
Kriss, Hamburg, spectroscope. Two 115-volt, 75-watt Mazda
lamps were used, one to illuminate the scale and the other as a
source of light for the collimator. All readings were taken with a
very narrow slit in the collimator tube. This type of instrument
has a movable mm. scale reflected into one of the tubes. The
70th mm. division was adjusted to the sodium flame spectrum,
and with this adjustment the flame spectrum of lithium fell at
51 mm., that of calcium at 61 mm., and of rubidium at 160.5 mm.
The conversion from mm. on the scale to wave lengths, the
standard unit for expression of the width of absorption bands
was accomplished by means of an interpolation curve. The
method for plotting this curve was taken from Landauer (’07).
The divisions of the scale were plotted along the abscissae on
coordinate paper; wave lengths from 400 to 750 uu formed the
ordinates. The divisions on the scale where the flame spectra
appeared for sodium, calcium, rubidium, and lithium, 60, 61,
160.5, and 51 mm., respectively, were converted into wave lengths
by using Rowland’s Table, Landauer (07). A smooth curve
was then constructed through these points. Measurements were
then made and these scale measurements were then converted
into wave lengths by means of the interpolation curve mentioned.

The methods used for crystallization of pigments in tissues were
2: (1) The Molisch (13) alcoholic-KOH. method, and (2) a modi-
fication of this. In the first method small strips of the fruit
tissue were placed in vials containing aleoholie-potash (40 per
cent C.H,OH by volume and 20 per cent KOH by weight).
These were then put in the dark at room temperature, 20? C.,
and for a short period out of every 24 hours the temperature was
raised to 75° C. They were kept for varying lengths of time until
crystallization appeared in the tissues, upon microscopic exam-

1925] ;
HOWARD—CAROTINOIDS IN FRUITS 175

ination. The second method, a modification, was proposed by
Van Wisselingh (15) to hasten crystallization. The same process
was followed as before, but 10 per cent by weight of KOH and
100 per cent glycerine was used and a constant temperature of
140? C. Inthis work a constant temperature of 95? C. was found
satisfactory. In both methods, after crystallization had taken
place, the crystals showed up more clearly if the pieces of tissue
were thoroughly washed in distilled H:O and mounted in glycerine.

EXPERIMENTAL DATA AND DISCUSSION

One gram of each kind of dried, ground fruit tissue was placed
in separate ground-glass-stoppered bottles, and 5 ec. of CS,
added to each and allowed to extract in the dark for 4 days. Then
spectroscopic and colorimetric determinations were made, also
comparisons with the color code. The results of the spectroscopic
analysis are given in table vr, those for colorimetrie determin-
ations in table vir, and for the comparison with the color code
in table уш.

Table vr shows that the pigment of tomato, watermelon,
Rhus, S. Dulcamara, Arisaema, and Aglaonema is largely lycoper-
sicin. These readings approximate those made by Willstátter
and Escher (10) for tomato. They show the limits of the 3
characteristic absorption bands of lycopersicin. The typical lyco-
persicin absorption bands are 2 in the green and 1 in the blue
region of the spectrum. The first band is never so distinct as the
second, which is a fact pointed out by Lubimenko (14) and found
to be true in this investigation. Pigments of Rhus and Arisaema
have never been shown to contain lycopersicin before, so far as
I was able to ascertain. Other carotinoids are also present in
these fruits, as demonstrated in the crystallization experiments.
When the CS, extracts from these fruits were mixed with equal
amounts of petroleum ether, in every case only a very little yellow
to orange color appeared in the petroleum ether layer, thus show-
ing the portion of carotin and xanthophyll to be rather small.

The pepper pigment requires some discussion. It has attracted
the attention of several workers because of contradictory results.
Palmer (’22) calls attention to the fact that the absorption bands
given by Tschirch (04) for the pepper pigment, namely, (1)

[Vor. 12
176 ANNALS OF THE MISSOURI BOTANICAL GARDEN

TABLE VI

SPECTROSCOPIC ANALYSIS OF VARIOUS itas EQUAL WEIGHTS OF
MATERIAL DRIED IN DEHYDRATER, POWDERED IN 5 AND
ACTED 4 DAYS WITH EQUAL AMOUNTS S OF CS
ABSORPTION BAND LIMITS ARE GIVEN IN ци

Material Band I Band II Band III
Aglaonema Treubii 562.0-548.5 524.0-513.2 495.2-483.2
Solanum Lycopersicum 557.0-536.5 520.0-503.0 485.0-477.5

us canadensis 557.0-536.5 20.0-503.0 485.0-477.5
Solanum Dulcamara 557.0-541.0 Кз ше 485.0-476.5
Arisaema triphyllum 557.0-541.0 520. 0-503.0 485.0-476.5
Citrullus vulgaris 557.0-541.0 522.5-505.0 485.0-476.5
Celastrus scandens (arils) 531.3-509.2 408.0-485.0
Solanum Pseudo-capsicum 528.5—509.2 494.5-480.0
Celastrus scandens MH 513.2-503.0 480.0-470.0
Evonymus america 528.2-513.2 494.5
равна га Sp. 522.5-509.2 494.5-480.0
Rosa rugosa 522.0 over, shaded no good

bands.
Evonymus europaea 531.3-513.2 483.5-490.0
Asparagus officinalis 531.3-511.3 501.3-488.8
Capsicum annuum 522.5-503.0 494.5—483.2
Lycium halimifolium 522.5-503.0 496.5-480.0
Cucumis Melo 531.3-513.2 501. :
Viburnum Opulus 513.2-503.0 480.0-476.5
Sorbus Айчына 513.3-517.5 500.0-485.0
TABLE VII

NER DETERMINATIONS MADE WITH A DUBOSCQ INSTRUMENT.
EADINGS GIVEN IN MM. NO DILUTIONS

.05 .01/Capsicum annuum 2.9 — .7|Citrullus vul-
5| garis

S. Duleamara 1.90 .47|E. americana 5.4 1. 10.5 2.8
C. scandens (arils) 2.15 .B4|Asparagus 6.8 1.9|Lo 11.88 3.03
S.L 2.30 .55|Lycium 8.2 2.2|E.europaea 13.4 3.7
ema .30 .57|Rosa 9.1 2.35.
C. scandens (car- Cucumis Melo 9.9 2.5| capsicum 13.9 3.9
pels) 2.35 .60 Vib 16.04 4.5
Sorb 42.4 11.9

517.0 џџ-501.0 uu, (2) 486.0 uu—467.0 uu, and (3) 458.0 uu-
439.0 џи, and those by Willstátter and Escher do not correspond
exactly, but that Tschirch’s (04) bands coincide more nearly
with the measurements made by Willstátter and Escher (10) for
carotin, which are 525.0 џџ-511.5 рь and 488.5 џџ-474.0 ци. In
а true lycopersicin spectrum the band furthest towards the red
end of the spectrum should lie at least as far over in the green as
550.0 ш. Lubimenko (14) does not class the pepper pigment
with the fruits he considers to contain lycopersicin, because its
absorption bands lie intermediate between pure lycopersicin and
pure carotin, and also because he was never able to crystallize
this pigment. In my work a good pure lycopersicin spectrum

1925]
HOWARD—CAROTINOIDS IN FRUITS 177

was never obtained. At times a very faint band appeared in the
characteristic position, but never with any degree of certainty.
The spectrum obtained always indicated the presence of carotin
and xanthophyll rather than lycopersicin. This may be due to
the presence of a mixture of lycopersicin and carotin, in which
the proportion of carotin is considerably greater than that of
lycopersicin, under which condition there is much difficulty in
getting distinet absorption bands. Even when greatly diluted
with СБ,, the СЪ, extracts of these 6 previously listed lycopersicin-
containing fruits exhibit a decided pink color. At equal dilutions
the other fruit extracts were orange in color. This interesting
phenomenon, due perhaps to dichromatism, is striking in a com-
parison of the pepper and tomato. From these 2 plants CS,
solutions of equal strengths are about the same color, but upon
dilution with CS, the tomato remains red, pink in great dilution,
while with corresponding strengths the pepper turns orange in
slight dilutions and yellow in greater dilution.

The spectrum analysis of the other fruit extracts require no
particular comment. They showed 2 bands, one in the green
and one in the blue region. This is a characteristic carotin
spectrum.

TABLE VIII
COMPARISON WITH a enn pea ET VALETTE “CODE DES COULEURS."
SAME вает AS U М TABLE VI. FIGURES TAKEN FRO

LE IN утаа & VALETTE'S "CODE"

Code Readings
Material ~
T 2 3
Solanum L ponti 31 56 56
Rhus canadens 66 66 66
Solanum Duleun ra : 36 51 56
Arisaema triphyllum 51 61 56
C us aris 46
Solanum Pseudo-ca 126 131 126
lastrus scandens (arils) 56
C'elastm dens ( 101 101 101
vonymus americana 101 pe ЧЕ
: 9
jum dou ш
Evonymus euro; 136
1 бе officinalis 111 111 111
Capsicum annuum 76 76 76
Lycium halimifolium 106 126 126
Cucumis Melo 91 096 91
Vi iburnum | Opulus А E AEN

[Vor. 12
178 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Table уш is of little value without the Klincksieck and Valette
‘Code des Couleurs. The smaller numbers represent the true
pinks and reds, the lycopersicin shades. Beginning with 76 the
red-orange tints come in, which are characteristic of carotin and
xanthophyll eolors. Again, evidence is obtained from this table
that the above-mentioned 6 fruits contain large proportions of
lycopersicin. An interesting point is the reading 76 for pepper,
which places it nearer the true lycopersicin color than any of the
other fruits which are known definitely to contain only carotin
and xanthophyll. There is an exception to this in the extract
from the arils of Celastrus with a reading 56, which I cannot
explain. |

In table уп, the determinations made with the colorimetre
show the relative order of intensity of the pigments contained in
the various fruits. According to their strength they have been
arranged in descending order in table уп, Rhus containing the
most concentrated pigment, and Sorbus the most dilute. :

Crystallization within the fruit tissues was attempted in order
to gain more knowledge of the kinds of pigments found within
certain fruits. Identification of pigments by means of crystals is
not very satisfactory, because of differences of opinion among the
various investigators as to the color, shape, and form in which
these pigments crystallize. Xanthophyll often crystallizes in
quadratic, often trapezoidal tables, frequently showing inden-
tation, or it may crystallize in lance or wedge-shaped prisms.
Sometimes the crystals are rhombic, almost hexahedral, with a
color variation from greenish yellow to rose. They are not in-
frequently similar to carotin, but usually contain less color, and
are often more pleochromatie.

Carotin erystals are rhomboidal in outline, tabloid or leaf-like
in form, and the color is frequently orange to orange-red, but
sometimes vermillion, with a steel-blue luster.

Lycopersicin yields brownish rose to carmine-red crystals,
usually in the form of needles or minute elongated prisms, and
often fissured at the ends. Sometimes crystal aggregates &re
formed, consisting of elongated prisms, at the center of which the
color is a bluish red. Often lycopersicin appears in long, fine,
hair crystals.

1925]
HOWARD—CAROTINOIDS IN FRUITS 179

Certain microchemical tests have been definitely established for
these З kinds of crystals. Those given here have been taken from
Molisch (713), Willstátter and Escher (10), Wilstátter and Mieg
(07), Van Wisselingh (15), and Palmer (22). All 3 kinds of
crystals turn blue when treated with H.SO, HNO., or HCl.
Lycopersicin crystals give more of an indigo-blue than carotin or
xanthophyll. IKI usually turns carotinoid crystals green, al-
though Van Wisselingh found an exception to this in some of the
crystals of the S. Dulcamara pigment. АП 3 give a blue color
with chlorozineiodide solution, and the same reaction with
bromine water. Xanthophyll erystals turn blue immediately,
and are not dissolved by 65-75 per cent H;SO,, while carotin
erystals turn blue only after the treatment with the acid has con-
tinued for some time, or when the acid is stronger. Lycopersicin
is dissolved in concentrated Н.ЗО.. Carotin and lycopersicin are
insoluble in phenol-glycerine (3 parts by weight of phenol and 1
part by weight of glycerine), but xanthophyll is readily soluble.

Xanthophyll crystals are entirely insoluble in low boiling pe-
troleum ether. The solubility in cold methyl alcohol is low,
greater in ethyl alcohol, a little greater in СӨ», and still greater in
ether. The solubility in acetone and chloroform is quite rapid.
Carotin crystals are almost insoluble in cold ethyl alcohol and
even less soluble in methyl alcohol, and they dissolve with dif-
ficulty in hot alcohols. Their solubility in low boiling petroleum
ether is poor, but somewhat greater in the higher boiling ones.
Acetone dissolves them with difficulty, even when hot. Benzole
dissolves them more easily, chloroform and CS, with great ease.
Lycopersicin crystals are less soluble than carotin in all the
carotinoid solvents. Ethyl and especially methyl alcohol are ex-
ceptionally poor solvents. Petroleum ether of low boiling point
dissolves only a small amount, while ether is a better solvent,
but CS, is much better. In CS: a 2 per cent solution may be
obtained. These crystals are insoluble in acetone and glacial
acetic acid.

According to the methods described above attempts were made
to crystallize the pigments in the tissue of each kind of fruit used
in this section of the work. In some fruits no crystals could be
obtained.

[Vor. 12
180 ANNALS OF THE MISSOURI BOTANICAL GARDEN

In tissue of Jerusalem cherry placed in alcoholic potash solution,
long, bright red needle-like crystals and a few aggregates were
observed after 16 days (pl. 18, fig. 2). Oftentimes small needle
crystals were seen within the cells. No success was obtained
with the Jerusalem cherry material placed in glycerine potash.
In tissue of Arisaema there was no crystallization after 21 days
in either alcoholic or glycerine potash. Tissue of Celastrus after
16 days showed rather short orange and yellow needle crystals
(pl. 15, fig. 5). Tissue of Sorbus placed in alcoholic potash solu-
tion yielded many very fine pale yellow-orange crystals (pl. 15,
fig. 7). Tissue of Lonicera under the same conditions showed
orange to yellow crystals (pl. 15, fig. 1), but broader than those
of Sorbus, and more like the xanthophyll crystals pictured by
Willstátter and Stoll (13).

The crystals formed in the pepper tissue were of special interest,
due to the uncertainty of the presence of lycopersicin. Strips of
tissue were placed in both potash solutions, but the best results
were obtained after 16 days in the alcoholic potash. The crystals
were of 2 types, the typical rhomboidal tabloid or leaf-like forms,
which were orange to orange-red, with a metallic luster so char-
acteristic of carotin, as described by Willstátter and Escher (’10);
and rose-red rosette-like crystals, which is one of the forms
ascribed to lycopersicin and one of the types found in tomato
(pl. 15, fig. 6).

A great variety of crystals was produced in the Asparagu& tissue
by the glycerine-potash treatment after 10 days. Some indica-
tions of carotin, xanthophyll, and lycopersicin were obtained (pl.
15, fig. 11). In Evonymus europaea crystals formed more rapidly
than in any other tissue, which is no doubt due to the fact that the
pigment is already present in an oily substrate distributed through
the entire cell. Van Wisselingh (’15) gives a very interesting
explanation of how these crystals are formed by the application
of the Molisch method. He says the pigments are dissolved
within the plastids in an oily substance, then through treatment
with alcoholic potash the plastid or stroma is destroyed and the
pigment rounds up into colored globules; the KOH then saponifies
the oily substance and the pigment is set free, crystallizing in the
alcohol, in which carotinoids are not very soluble. Heat no doubt

1925]
HOWARD—CAROTINOIDS IN FRUITS 181

increases the saponification process. When fruit tissues are
placed in either of these KOH solutions the first change to be
noted is the formation of colored globules. Since the plastids
in E. europaea, at the point of highest coloration of the aril,
are already broken down and the pigment is distributed in oily
globules, the addition of the alkali effects at once the saponifica-
tion of the lipoid material and the precipitation of the pigment in
a crystalline form, thus accounting for the rapid crystallization in
this tissue. The crystals appeared to originate from the colored
globules mentioned (pl. 14, fig. 1, pl. 17, fig. 4). After 5 days
in glycerine potash Crataegus showed very distinct carotin crys-
tals of a deep vermilion red with the characteristic metallic
luster (pl. 15, fig. 3). After 10 days in alcoholic potash there
were formed in the fruit of S. Dulcamara beautiful carmine crys-
tals of lycopersicin similar to those shown by Willstátter and
Mieg (707) (pl. 15, fig. 8, pl. 17, fig. 2). Good lycopersicin crys-
tals were obtained in the tissue of Rhus after treatment of 30
days with alcoholic potash (pl. 15, fig. 4). With Evonymus
americana an interval of only 7 days in the solution was re-
quired in order to produce many fine needle crystals, orange-
yellow in color (pl. 17, fig. 3). In Lycium the crystals observed
in the cells after a treatment of 7 days with alcoholic potash
were rather broad needles (pl. 15, fig. 2).

Some of the tests described earlier were applied to certain of
these crystals. Lycium, Celastrus, Sorbus, Aglaonema, Asparagus,
and Crataegus all gave a green color with IKI solution. With
the exception of Sorbus, crystals from the same fruits gave a blue
reaction after a treatment of 1 hour with 85 per cent H.SO,. In
Sorbus the crystals changed color almost immediately, thus in-
dicating the presence of xanthophyll rather than lycopersicin or
carotin. No results were obtained with bromine water nor acetic
acid. Phenol-glycerine dissolved some of the crystals in the
Sorbus tissue, but this reagent had no effect upon most of the
crystals.

From the results of this work it is difficult to draw conclusions
as to just which carotinoids are present in all of these fruits.
All the workers on fruit pigments have agreed that lycopersicin
is the principal carotinoid in tomato, watermelon, Aglaonema

[Vor. 12
182 ANNALS OF THE MISSOURI BOTANICAL GARDEN

(several species) and Solanum Dulcamara; while by some workers
pepper and rose are added to this list. The results of this work
confirm the presence of lycopersicin in tomato, watermelon,
Aglaonema Treubii and Solanum Dulcamara, adding to the list
Arisaema triphyllum and Rhus canadensis. The presence of lyco-
persicin crystals in pepper is certainly an evidence of the presence
of the pigment, the failure to get good characteristic lycopersicin
absorption bands no doubt being due to the small proportion of
the pigment in pepper tissue. I have found no indication of lyco-
persicin in rose, although Lubimenko (’14) lists the rose among
fruits that do contain it. In some fruits the spectroscopic work
and the crystallization are rather contradictory. In Jerusalem
cherry the long, pinkish red, needle crystals characteristic of
lycopersicin were found, while in Asparagus tissue rosette crystals,
also characteristic of lycopersicin, were observed. From the
spectroscopic determinations there was no indication of lyco-
persicin in either of these tissues. 3
me fruits were found to contain both anthocyanin and caro-
tinoid pigments. These were Crataegus, Rosa, Sorbus, Lonicera,
and Viburnum. In each fruit the anthocyanin pigments were
located in the cells of the epicarp. As these pigments are soluble
in the sap and not contained in plastids, these cells appeared as
if filled with a colored liquid. The mesocarp tissue when ex-
amined under the microscope contained many plastids. All of
these fruits when boiled in water lost their bright external colors,
but the carotin-colored plastids remained without any change.
Since the anthocyanin pigments when in combination with car-
otinoids in the same fruit were always in superficial cells, the
writer accepts the view that perhaps light is a strong factor in
effecting this type of distribution. It would seem that light is
more necessary for the production of anthocyanin than for caro-
tinoid pigments. No satisfactory experimental work has been
carried out during this investigation to substantiate this theory.
Observational evidence is both for and against the idea. The
most highly colored apples are raised in the regions of most
intense light, also the brightest colors of most fruits such as
peaches, plums, grapes, pears, strawberries, etc., are in the epicarp

1925]
HOWARD—CAROTINOIDS IN FRUITS 183

or close to the surface of the fruit, where the light is greatest.
Among roots we find certain ones that are highly colored with
anthocyanin, such as red beets, radishes, and turnips; in other
cases the pigments are carotinoids, as in carrot. According to
Wheldale (16) few definite experimental data have been accu-
mulated and the evidence is very conflicting. Some flowers and
fruits grown in the dark develop color while some do not. A
general survey of the distribution of anthocyanin shows without
a doubt that it occurs most often in the external tissue.

The beet root is à good illustration of the development of an
anthocyanin pigment practically in darkness. A few other in-
stances are recorded of pigment formation in the dark. Wiesner
(77) observed that potato sprouts, which formed in the light,
showed little, if any, yellow pigment, while those formed in the
dark developed from 30 to 150 per cent more pigment. Elfving
(82) and Immendorff (’89) found that carotinoids increased
greatly in leaves under conditions which depressed chlorophyll
formation, that is, low temperature and very diffuse light. Lubi-
menko ('14) maintained that light was not necessary for the for-
mation of lycopin. Duggar (13) proved that light is absolutely
unnecessary for the normal pigmentation of red tomatoes.

Evonymus europaea offers an interesting condition, for here
the arils colored with a carotinoid become bright red before the
colorless carpels have burst open. The carpels are later colored
by an anthocyanin. After the carpels break open they assume a
rosy red color. An experiment was tried in which some green
fruits of Evonymus europaea were placed in the dark and some in
the light at the same temperature and at the same humidity. No
satisfactory results were obtained as the small fruits dried up
without any decided appearance of color in the carpels, under
either set of conditions.

Gaultheria ovatifolia Gray, Symphoricarpus orbiculatus Moench,
Fragaria sp. (cultivated), Rubus spectabilis Pursh, Sambucus cal-
licarpa Greene, Vaccinium parvifolium Smith were found to
contain only anthocyanin pigments.

Two tests were made to determine what kind of pigments are
present in these fruits. (1) Each kind of fruit was boiled and in
every case a colored liquid was obtained and a decrease or entire

[Vor. 12
184 ANNALS OF THE MISSOURI BOTANICAL GARDEN

loss of color in the fruit itself. (2) Air-dried and powdered ma-
terial of each fruit was treated with CS, and allowed to stand for
2 weeks. No colored extract was ever obtained with any of the
fruits tested.

CARROT PIGMENTS

Experiment 1.—To determine the state in which carotin is
found naturally occurring in the carrot root, thin, free-hand
sections of mature carrot containing an abundance of crystals
were cut and placed in vials containing petroleum ether and
absolute alcohol (10 ec. of petroleum ether and 1 cc. of absolute
alcohol) maintained in darkness (pl. 15, figs. 9-10). Upon ex-
amination with the microscope after 48 hours no evidence of any
crystals or granules could be found in the tissue, and no trace of
color remained.

For the same purpose microtome sections, 10 џ in thickness, of
mature carrot material, killed in chromo-acetie and imbedded in
paraffin were stained in Delafield’s haemotoxylin as described by
Chamberlain (15). Upon examination under the microscope no
crystals or granules were observed in these slides.

Free-hand sections from the same region of the carrot were
placed in iodine and 70 per cent alcohol and left in darkness for
24 hours. Upon examination with the microscope after this
treatment no color bodies were observable in the tissue. From
these experiments the conclusion was drawn that in the mature
carrot root the pigment is in a crystalline or granular form free in
the cytoplasm and not contained in any plastid or stroma, other-
wise some evidence of a stroma would have appeared in the sec-
tions treated by the above 3 methods. The size, shape, and
general form of most of these color bodies under the microscope
indicated the presence of crystals rather than granules.

Experiment 2.—To study the question of where and how the
carotinoids in carrot are formed, very young carrot roots were ex-
amined both by means of free-hand and paraffin sections. The
microtome sections proved to be of little interest as the pigment
itself was dissolved by all of the methods used. Free-hand
sections were cut from very young carrots from 3 regions: (1) very -
close to the root tip, (2) half way between the root tip and the

1925]
HOWARD—CAROTINOIDS IN FRUITS 185

top, and (3) at the top of the root just where the stem joins the
root. Sections from these 3 regions were all tested for starch
with IKI. Sections from region (1) gave no test for starch, from
region (2) very.good starch tests. An abundance of leucoplasts
could be seen before the IKI was added, and in some sections a
peculiar arrangement was noted between the carotin-colored
bodies, which were sparingly present and the starch (pl. 16, fig.
9). There was a close connection between these starch grains
and masses of pigment, as indicated by the blue coloration and
the orange pigment. Leucoplasts containing starch inclusions
were also observed (pl. 16, fig. 12). Sections cut from region (3)
gave practically no starch test, but an abundance of chloro-
plasts were present and in addition many tiny carotin crystals
in the fibro-vascular bundles. Upon the addition of H.SO, these
erystals gradually turned blue, and green with IKI, two decisive
tests for carotin (pl. 16, fig. 1D.

zperiment 3.—Mature carrots often show a green coloration
near the top of the root. Sometimes this extends down into the
center of the vascular cylinder. Tests were made for starch and
sugar by means of IKI and Fehling's solution respectively. An
abundance of starch was present but no sugar, and more starch
was found to be present in the top region than in the bottom.
Sections cut about .5 mm. in thickness from mature carrots and
examined under the low power of the microscope showed very
distinctly that the arrangement of the great masses of pigment
followed the medullary rays (pl. 16, fig. 10).

Experiment 4.—To show the distribution relation between the
chloroplasts and carotin crystals, a mature carrot was divided into
4 regions, each 1 cm. wide. From each region free-hand sections
were cut, mounted in water, and examined under the microscope.
Beginning at the top these regions were numbered 1, 2, 3, and 4.
In each region sections were cut from the vascular eylinder and
the cortex. In region (1) the sections from the cortex showed
only chloroplasts, while those from the vascular cylinder showed
chloroplasts and some carotin crystals. In region (2) the cortex
sections showed only chloroplasts, while the vascular cylinder
only greenish yellow and orange crystals. In region (3) the cortex
sections contained both chloroplasts and carotin crystals, some

[Vor. 12
186 ANNALS OF THE MISSOURI BOTANICAL GARDEN

of the latter being of good size. In the vascular cylinder sections
many carotin crystals and a few chloroplasts were observed. In
region (4) the vascular cylinder sections contained many large
carotin crystals, while the cortex contained many very large
earotin crystals and also an abundance of smaller ones and
granules but no chloroplasts. These crystals were of various
shapes and forms with some variation as to color. They varied
in size from 2.1 to 3 u by 10.0 to 42.7 в. Color variations were
from pale orange to red (pl. 16, figs. 1-8).

Experiment 5.—This part of the work was undertaken to deter-
mine whether mitochondria are present or not in carrot. Both
mature and various stages of young carrot roots were killed by
Renand's IV B method, imbedded in paraffin, and stained in
Haidenhain's iron-haematoxylin (Cowdry, '18). No definitely
staining bodies resembling mitochondria could be found. Many
chloroplasts stained well in the region where root and stem join
(pl. 16, fig. 13).

From these experiments the following conclusions seem war-
ranted: (1) The carrot carotin is typically in a crystalline or
granular state, and not contained inastroma. This is opposed to
the conception of Schimper (’83) who considered the carrot
carotin to be contained in chromoplasts. (2) Carotin is laid
down as a storage product in the medullary rays. (3) The amount
of carotin crystals increases from the top of the root towards the
bottom, while the chloroplasts decrease. (4) No evidence could
be found of mitochondria in young carrot roots.

RIPENING EXPERIMENTS
EXPERIMENTAL DATA AND DISCUSSION

These experiments were undertaken in an attempt to determine
the most favorable temperature for rapid fruit ripening, that is,
to bring the fruit to the point of red ripeness. Very little work
has been done on temperature relations. Duggar (13) in some
experiments with tomatoes showed that light was unnecessary for
normal ripening, and that the most favorable temperature range
was 18-25° С. In another experiment he found that red peppers
ripen normally at higher temperatures.

1925]
HOWARD—CAROTINOIDS IN FRUITS . 187

Experiment 1.—Green fruits of Lycium halimifolium Mill. were
placed under 4 different conditions, 9 fruits in each lot. These
conditions were as follows: (1) wrapped in black paper in a glo
at 17-20? C.; (2) in front of west window at 18-22? C.; (3) in
incubator at 26—28° C.; (4) in an incubator at 32-35* C. The
results are given in table rx.

From these data the rapid factor in ripening fruits is shown to
be heat and not light, a temperature of from 26-28? C. being the
most favorable. "These results agree with those found by Duggar
(13), although he was working with pepper.

Colorimetric determinations were made of the red-ripe fruit
from each lot. These fruits were dried in a gas-drying oven at
60° C., ground up in a mortar, and extracted with CS, (0.3 gm.
of dried material in 4 cc. of CS, extracted for 4 days). The
methods were the same as those already indicated.

TABLE IX

COLOR CHANGES IN GREEN LYCIUM FRUITS UNDER 4 TEE D ele
ING CONDITIONS з G А PERIOD OF 5 DAYS. NOTES
LOT.

UITS IN EACH
cep Lot Lot Lot 4
Days 18-22? C. 26-28? 7. 32-35? C.

Wrapped in de On table in front In electric In electric

| paper put in desk | of west window oven oven
1 5 green dn 1 green 3 green

2 yellow bellows green : yellow 3 yellow ·

1 almost orange 1 ricas 2 nearly orange |3 orange

1 orange 5 orang

1 green 2 green 1 green 1 green
2 |3 yellow { yellow : yellow а yellow

2 orange orang: orange ora:

3 red-ripe 6 red-ripe 4 dps

1 green 1 green 1
3 3 ай 1 pale yellow 1 orange 2 Fellow

2 orange 4 yellow 7 red-ripe 6 red-ripe

3 red-ripe 2 orange

1 red-ripe

1 greenish yellow |4 yellow 1 yellow 2 green
4 3 yellow 4 orange 1 orange 1 yellow

1 orange 1 red-ripe 7 red-ripe 6 red-ripe

4 red.

1 greenish yellow |4 yellow 1 yellow
5 3 yellow 3 1 orange 1 yellow

1 orange 2 red-ripe 7 red-ripe 6 red-ripe

4

188 .

COLOR

[Vor. 12

ANNALS OF THE MISSOURI BOTANICAL GARDEN

CHANGES

TABLE X

IN GREEN PEPPERS UNDER 5 DIFFERENT RIPENING

CONDITIONS DURING A PERIOD OF 42 DAYS, 20 FRUITS IN EACH
T*

Lot 1 Lot 2 Lot 3 Lot 4 Lot 5
26-28? C. 22-24? C. 20-22? 16-18? C. 14-16? C.
Wrapped Unwrapped Unwrapped Unwrapped Wrapped
2 red-ripe 4 one-half or-|3 red-ripe 1 red-ripe 4 half orange
8 almost red-| ange to red-|5 almost red-|3 almost red-| to red-spot-
ripe spotted ripe d
7 red-orange |10 orange to|5 вида 1 one-half or-|10 orange to
6 |3 green, f red-spotted | red-ripe nge-red red-spotted
days | red-orange |6 green green, redjll one-fourth|6 green
spots spots orange-re
5 green 1 three-fourths
green
3 green
4 almost red- Е red-ri 4 red-ripe 4 red-ripe 2 almost red-
ri denies be-|5 almost гед-11 three- i
11 three-| ginning urths red-|9 three-fourths
fourths red-| spoil 1 three-fourths} ri red-ripe
12 ripe 9 almost red-| red-ripe 1 yellow 5 green, or-
days |2 few orange-| ripe 5 one-fourth |1 green-yellow| ange-red
2 three-fourths -ripe stri spots
1 spoiled red-ri 1 few yellow|2 green 3 green
1 one-fourth pots 1 spoiling
red-ripe green
2 red-ripe 3 red-ripe 1 red-ripe 2 red-ripe 4 almost red-
4 almost гед-6 almost red-|3 m red-|7 ик t red-| ri
ripe ripe 8 three-fourths
9 three-fourths] three- |5 three iourtbe 3 three fourth red-ripe
18 ipe fourths red-| red-ri red-ri; 2 one-third
days |2 few orange-| ripe 4 spoile 3 node. -ripe
spots |5 spoiled 8 few red spots
1 yellow
2 spoiled
red-ripe 5 red-ripe 6 almost red-|8 almost гед-|8 almost red-
almost red-|2 almost red-| ripe ripe i
ripe ripe 2 three-fourths|2 three-fourths|3 three-fourths
2 three- red-ripe red-ri red-ripe
days | fo 3 spoiled 3 one-half red-
TI тї
i x"
spoili
2 red-ripe -ripe -ripe -ripe -ripe
MI red Ji mes sod. aluo PED MOM [Ub рем red-
n pe ripe
days |1 three- 2 three-fourths|2 three-fourths|3 three-fourths
fo -ripe -ripe red-ripe
1 yellow
6 spoiled

* As soon as a pepper became red-ripe or spoiled a record was no longer kept of it.

1925]

HOWARD—CAROTINOIDS IN FRUITS

TABLE X (Cont.)

189

Lot 1 Lot 2 Lot 3 Lot 4 Lot 5
26-28? C. 22-24? C. 20-22? C. 16-18? C. 14-16? C.
Wrapped Unwrapped | Unwrapped | Unwrapped Wrapped
7 red-ripe 1 red-ripe 3 almost red-|3 red-ripe 4 almost red-
3 2 almost red- ripe 4 almost red-| ripe
days | pe 2 three-fourths} ripe 3 three-fourths
y red-ripe 1three-fourths| red-ripe
red-ri ellow
2 red-ripe 3 red-ripe 4 almost red-|5 almost red-
2 almost red-| ripe ripe
42 ripe 1 three-fourths|2 three-fourths
days red-ripe red-ripe
1 yellow

The results showed that fruits ripened under the condition
of lot 3 give a more deeply colored extract than any of the others.
From this it was concluded that 26-28? C. is the most favorable
temperature for ripening fruits of Lycium. The temperature
range doesn’t agree with that of Duggar (13) for tomato, but the
presence of a large percentage of lycopersicin in tomato and none
in Lycium may account for this difference.

Experiment 2.—Bell peppers of uniform size and greenness,
20 in each lot, were placed under 5 different conditions as to tem-
perature and light. Lot 1 was placed in an electric incubator
(26-28? C.), each fruit being first wrapped in brown paper to pre-
vent too much drying out; lot 2 on a table in front of a south win-
dow (22-24? C.) unwrapped; lot 3 on the floor (20-22° C.) un-
wrapped; lot 4 on the floor (16-18? C.) unwrapped; lot 5 on the
floor (14-16? C.) wrapped. Detailed color notes taken at 6-day
intervals are given in table x. From these results 22-24" C. is
the best temperature for ripening peppers, the higher tempera-
tures being more favorable, however, than the lower. Light is
not a controlling factor.

Spectroscopic determinations were made on samples from
peppers ripened under conditions of experiment 2. The methods
followed in this work have been previously described. The pulp
of the red-ripe fruits from each lot, after removal of the seeds, was
placed in the dehydrater until perfectly dry, then ground as fine
as possible with a coffee mill. Weighed amounts of this material
were then extracted with relatively equal portions of CS, for 2
days. The extract in all cases was too dense to show absorption

[Vor. 12
190 ANNALS OF THE MISSOURI BOTANICAL GARDEN
bands. The whole spectrum was blotted out from 652.2 uy over
to 412.5 ри, so that dilutions had to be made before the absorption
bands could be observed. The amount of dilution varied with
the different lots of peppers. The absorption bands in all lots
lay approximately in the region of the carotin and xanthophyll
spectrum. These spectroscopic analyses are given in table x1.
The most concentrated extract was believed to indicate the best
ripening condition.

TABLE XI

SPECTROSCOPIC ANALYSES OF PEPPER EXTRACTS ien EXPERIMENT 2.
LL MEASUREMENTS ARE IN y

Lot 1 2 3 4 5
Dilutions 2 cc. extract | 2 сс. extract | 2 сс. extract | 2 ec. extract | 2 сс. extract
10 ee. CS; | 11 ec. CS2 8 се. CS2 5 се. CS2 З се. CS2
Absorption | (1) 522.5- | (1) 523. 5- (1) 521.5- | (1) 523.2- | (1) 522.6-
bands 505. 507. 506. Х á
(2) кез 2- | (2) eid 5- | (2) 496.5- | (2) 497.5- | (2) 497.5-
83.0 2.0 482.0 483.0 483.0

The same conclusion was drawn from these results as from ex-
periment 2. Lot 2, with a temperature range of 22-24? C
offered the most favorable temperature for fast ripening. Lot 1
ripened at 26-28? C., the next most favorable. Lot 5, ripened at

14-16? C., gave a very weak extract needing only about one-
fourth the dilution of lot 2.

Experiment 3.—These extracts from experiment 2 were further
tested by comparison with Klincksieck et Valette’s ‘Code des
Couleurs.’ The method followed was to dip 3 equal-sized pieces
of filter-paper in each of the 5 extracts obtained from experiment
2. These were allowed to dry for 5 seconds and then a comparison
made with the charts. Three readings were made for each ex-
tract. In most cases these triplicate readings were exactly the
same. They are recorded in table хи.

TABLE XII

За EXTRACT FROM FRUITS RIPENED UNDER E OF EX-
PERIMENT 2. COLOR ний ТАКЕ N FROM KLINCKSIECK &
ALE "CODE"

Lot 1 2 3 4 5
26-28° C 22-24? C. 20-22? C. 16-18? С 14-16? C.
Readings
1 61 66 66 Тї 71
2 61 66 66 71 71
3 61 61 66 ra! 71

1925]
HOWARD—CAROTINOIDS IN FRUITS 191

Since the smaller the figures the deeper the color this experiment
is further proof that higher temperatures (22—24° C.) are necessary
for deep pigmentation of fruits during the ripening process.

Experiment 4.—Color changes on the individual pepper were
observed with green fruits placed in the dark at a temperature of
22-25? C. "The first changes were a few yellow spots, then dark-
ening of these spots to an orange, then to an orange-red. Finally
this orange-red color spread gradually over the entire fruit until
red-ripeness was reached. These same color changes were ob-
served in 12 individual fruits.

SUMMARY

1. During the ripening process of fruits a change occurred in
the general shape of the plastids from oval or subglobose to a
very much elongated spindle form. In some of the red-ripe
fruits the pigment was crystalline in form.

2. In most of the ripe fruits studied the pigment was found to
occur in a stroma or in a definite body. This pigment was in an
amorphous state contained in a lipoid substratum.

3. Lycopersicin was present in a crystalline form in the red-
ripe fruits of Solanum Lycopersicum, Cucumis Citrullus, and
Aglaonema Treubit.

4. A Duboscq micro-colorimeter was used with good results as a
means of determining relative amounts of pigments in tomatoes.
Of the 4 varieties studied, Ponderosa, Excelsior, Dwarf Champion,
and Globe, Excelsior was found by this method to contain the
most pigment.

5. Dried tomato pulp kept in the dark for a period of 5 months
lost very little color. This was proved by comparisons of the
CS, extract of the dried material, at certain intervals, with a
methyl red solution.

6. A characteristic lycopersicin spectrum was obtained with a
CS, extract of the pigments in the ripe fruits of the following:
Aglaonema Treubii, Solanum Lycopersicum, Rhus canadensis,
Solanum Dulcamara, Arisaema triphyllum, and Citrullus vulgaris.

7. A characteristic carotin spectrum was obtained with a CS,
extract of the following fruits: Celastrus scandens, Solanum
P psicum, Evonymus americana, Lonicera sp. Rosa rugosa,

[Vor. 12
192 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Evonymus europaea, Asparagus officinalis, Capsicum annuum,
Lycium halimifolium, Cucumis Melo, Viburnum Opulus, and Sor-
bus sitchensis.

8. Only anthocyanin pigments were found in the fruits of
Gaultheria ovatifolia, Symphoricarpus orbiculatus, Fragaria sp.,
Rubus spectabilis, Sambucus callicarpa, and Vaccinium parvi-
folium. Both anthocyanin and carotinoid pigments were found
in the fruits of Crataegus phaenopyrum, Rosa rugosa, Sorbus
sitchensis, Lonicera sp., and Viburnum Opulus.

9. The carrot carotin occurred in both crystalline and granular
forms, but free in the cytoplasm. It appeared to be laid down as
a storage product in the medullary rays.

10. The optimum temperature for the rapid ripening of Lycium
halimifolium fruits was 26-28? C. The optimum temperature for
the rapid ripening of pepper fruits was 22-24? C.

The writer wishes to express her appreciation of the valuable
suggestions and kindly criticisms of Doctor B. M. Duggar, under
whose direction the work was carried out. Thanks are also due
Doctor J. M. Greenman for the verification of all scientific
names, and to Doctor G. T. Moore for the privileges and facilities
of the Missouri Botanical Garden.

Graduate Laboratory, Missouri Botanical Garden.

BIBLIOGRAPHY

Arnaud, A. (’85). Recherches sur les matiéres colorantes des feuilles: identité de
la matière rouge orangé avec la carotine, СзН»:О. Compt. Rend. Acad. Paris
100: 751 -753. 1885.

а ). Recherches sur la composition ER la carotine, sa fonction

vus et за formule. 1634. 102: 1119-1122.

„ (87). Recherches "pep Ka ooo et x ph ае sur l'amidon et

les grains de chlorophylle. Ann. Sci. Nat 1 5:179-310. pl. 5-8. 1887.

Bohm, J. A. (57). Beitráge zur naheren Ke sA des Chlorophylls.

Wiss., Wien, math.- naturw. Kl., Sitzungsber. 1857. (Cited

Cavers, F. YA Chondriosomes (Mitochondria) and th

Ph; 3: 96-106, 170-180. 88 1914.

Ани C. J. (19). вано а о aid T e oe к

Chmiel itraege zur Mo e аы
каа Fons b i den niederen Pflanzen. Charkow) 1890. [Russian, cited

Courehet, Тә L. rr Ai 15-18. 1886. les chromoleucites. Ann. Sci. Nat. Bot. VII.
Cowdry, N. H N. H. NONE ыы logy of the Myxomycetes
iol. Lab.,

reference to
Biol. Bul. 35: 14-94. 1. 1918.
Davis, B. M B. e E ‘The, Боо r-cell of Anthoceros Bk Gaz. 28: 69-109.

1925]
HOWARD—CAROTINOIDS IN FRUITS 193

Duggar, B. M. (18). Lycopersicin, the red pigment. of the tomato, and the “еи
of conditions upon its development. Wash. Univ. Studies 1: 22-45. 1913.
Ehring, C. (96). Ueber M Farbstoff der дада (Lycopersicum esculentum).

Ein Beitrag zur Kenntniss des Carotins. Inaug. Diss. München. 35 pp
1896. [Bot. Ce Ка). 69: 154. 1897.)
Wi Fr. (82). Ueber eine Beziehung zwischen Licht und Etiolin. Bot. Inst.
ürzburg, Arb. 2 : va быз 1882. d by Kohl, '02.)
ur

r, H.
ied by Palmer, ’22.
uler, H., und Nordenson, E. (08). Zur Kenntniss des Moehrencarotins und seine
Be gle itsubstanzen. Hoppe-Seyler's Zeitschr. f. НО, Chem. 56: 223-225.

Fritsch, P. (84). Ueber TTA ча Stoffe des Zellinhalts. Jahrb. f. wiss.
Bot. 14: 185-231. pl. 11-13. 1884.
ON Compt: ke A. e 3). Nouv o observations sur le ЖЕТТИН des champignons.

ches sur Pe ine des gs н et le mode de foramen des pigments
xantho payee et carotiniens. Rev. Gén. Bot. 31: 635-770. pl. 10-45. 1919.
йыма: F. А. (73). Note a b “substance colorante des baies rouges. Compt.
Rend. Acad. Paris 76: 385.
Жасы А. (61). Ueber Carotin und Hydrocarotin. Liebig's Ann. d. Chem.

Immendorff, H. (s 9). Das Carotin in Pflanzenkórper und einiges v es grünen
_ Farbstoff des Е llkorns. Landwirt. Jahrb. 18: 507-520.
Klincksieck, P., et Valette, T. (08). Code des танін 81 pp. 2 f. 1908.
e sies G. E aa Un tersuchungen @ M г das Carotin und seine physiologische
ung in der nze.
Kozlowski, BS Critique vs l'hypothàse des chondriosomes. Rev. Gén. Bot.
у

9. pl. 1
Kraus, "с. v) Zur е der Chlorophyllfarbstoffe und ihrer Verwandten.
Spectralanalytische e Untersuchungen. 131 pp. 3 pl. Stuttgart, 1872.
ie (207). Spectrum analysis. 236 рр. 49f. 1907. [Translated by J. B.

lewis, G. (10). и die ирип in pflanzlichen Zellen. Ber. d.
deut. bot. Ges. 28: 538-546. pl. 17. 1910.
Lotsy, J. P. (07). и über 0 а Stammesgeschichte 1: 27-30. Jena,

Lubimenko, (14). Quelques recherches sur la lycopine et sur ses rapports
vec la chlorophylle Rev. Gén. Bot. 25: "474-493. 1914.
енсе n ormations of plastid pigments in living plant tissue. Acad.
Sci. Petrograd, Mem. [8] 33: 275 pp. 1916. [Chem. Abstr. 14: 1697-1698. 1920.]
Medalia, L. S. (20). Color standards for the colorime тіс measurements of H-ion
to pH о 5: 44 1920.

M cr Ae (n b pH as Chor меа ^ ian 2 Ti
eyer, A. ('83 oro orn. p. 7
о 95, tersuc eni über die Stürkekórner. 318 pp. 9 pl. 99 f.

—— (1 Bemerkungen zu G. Lewitsky: über die Chondriosomen in
pflanzlichen Zellen. Ber. d. deut. bot. Ges. 29: 158-160. 1911. ;
Mikosch, C. (85). Ueber die Entstehung der Chiorophyllkórner, Akad. d. Wiss.,
Wien, math.-naturw. Kl., xs ungsber. 92!: 72-109. 1885.

Millardet, A. (76). Note sur ne substance colorante nouvelle (solanorubine)
d T неди" ancy,

i roue Я e te ани Я phylls. Bot. Zeit. 13: 89-99, 105-

Molise ce 96 Die Krystallization und ze o t ariig rs Xanthophylls
+ = Y ы latte. Ber. d. deut. bot. -— A B pl.2. 1896.
temerity . Mikrochemie Pflanze. pp.
Montanari, ed. Matéria е, коой te róssa del pomidoro. Staz. Sper. Agrar.
Ital. 37: 909-919. 1904.
Montéverdé, N. A., et Lubimenko, V. N. (13). Recherches sur la formation de la

[Vor. 12
194 ANNALS OF THE MISSOURI BOTANICAL GARDEN

en eds chez les plantes. IV. Sur p тонос. её m lycopine. Acad.
etrograd, Bul. VI. 72: iM рі. 1-2. f. 1. 1913.

Mottier. D. M. (18). Chondriosomes and v еы ordia of chloroplasts and leuco-
plasts. Ann. Bot. 32: 91-114. pl. 1.

Nägeli, C. von (58). Die Stürkekórner.

— e and G. de £9. Délla matéria oar délla angüria e del pomidoro.

chim. ital. 9: 506-507. 1879.
рав. ^ (92). Zur Kenntniss der Früchte von Capsicum annuum. Arch.
. 230: 108-134. 92.

Palmer, L. 8. dy Carotinoids and related pigments. 316 pp. pl. 1-2. f. 1. 1922.

= а ckles, C. H. (14). Carotin.—The principal natural yellow pig-
ment of ill fat. Part II. Chemical and physiological relations of pigment of
milk fat to the carotin "e — of green plants. Mo. Agr. Exp. Sta.,
Res. Bul. 10: 339-387.

Passerini, М. ('90). Sulla Pc NIE chemica = е del pomidoro (Solanum
Lycopersicum L.). Staz. Sper. Agrar. Ital. 18: 545-572. 1890.

Pugsley, C. W. (17). A successful raisen е plant. U. 8. Dept. Agr.,

armers' Bul. 916: 1 -12. n 1-9. 1917.
DS d I. (13). Über Chromoplasten in vegetativen Organen. Acad. Sci.
e, Bul. Internat. B. 1912: P189 -336. 1913.
Rudolph, K. (12). Chondriosomen und Chromatophoren. Beitrag zur Kritik
hondriosomentheorien. Ber. d. deut. bot. Ges. 30: 605-629. pl. 18. f. 1.

UR
Sachs, J. (62). Ueber den Einfluss des Lichtes auf die as des Amylums in
den hin, AA. (D. U órnern. Bot. Zeit. 20: 365-373. 1862.

Schimper, A. F. W. (83), Ueber die Entwickelung der Chlorophyllkérner und
Farbkórper. Bot. Zeit. 41: 105- 112, 121-131, 187-146, 153-162. pl. 1. 1883.
———— — (95). Untersuchungen. über die Chlorophylikérper und die ihnen
homologen Gebilde. Jahrb. f. wiss. Bot. 16: 1-247. pl. 1-5. 1895.
таратат Бе A. (01). The yellow su Pores сүт eer ee эге иң a ee uen fee and

Lb (03). The xanthophyll group of yellow colouring matters. Ibid. 72:

nn, G. (208). Die rye und Lageveründerung der Pflanzen-Chromatophoren.
1

. (21). Ар introduction to cytology. 402 pp. 159 f. 1921.
Tammes, T. CHA. Ueber = Verbreitung des Carotins im Pflanzenreiche. Flora
* 1

; MB:
Tobler, С G. und F. (1 0). ‘Untersuchungen über Natur und Auftreten von Carotinen.
. Fruc “ee а Momordica Balsamina І. Ber. d. deut. bot. Ges. 28: 365-376.

————, (10а). den e — Tu nm der ананар bei der

bid. - 22:
Tsweet, M. SA 00). ыы аи d das Chlorophyll. Die
Awe an Ges. 24: 316- Sa

1925]
HOWARD—CAROTINOIDS IN FRUITS 195

bien seg, tà H. (1826). Ueber das Oleum radicis Dauci qoae i das Carotin,
n Carotinzucker und den officinellen suceus Dauci; e . f. Anthel-
aie Göttingen; Geiger's ;Mag. Pharm. 33: 144-172. ` 1831. (Cited by

Palmer, 722.)
Wheldale, M. (16). The anthoc йет ә pete of sgg cnr 318 pp. 1916.
Wiesner, J. (77). Ueber das tehung von Etiolin und
Chlorophyll in der Kartoffel. gene Bot er me 27: 7-11. 1877.
Magne R., und t H. H. (10). Uber den Farbstoff der Tomate. Hoppe-

р А 10.

———,‚ und WW. ('07). Ueber die gelben Begleiter des Chlorophylls.
Fat шй d. Ж 355: 1-28. 1

—— ——————., und Stoll, A T ^Y) Psp wer, über Chlorophyll. Methoden und

Ergebnisse. ;

Wisselingh, C. van Vi : die Nachweisung und das Vorkommen von Caroti-

. noiden in der Pflanze. Flora 107: 371-432. 1915.
Zeise, (47). Ueber das Carotin. Liebig's Ann. d. Chem. 42: 380-3 1847.
immermann, A. (94). Sammel-Referate aus dem Gesamm tgebiete der Zellenlehre.

9. Die ne to gr are ue og bot. Centralbl. 4: 90-101. 10. Die Augenfleck
Stigma. Ibid. 161-165. Elaioplasten, Elaiosphüren n und verwandte Körper
bet 165-169. 15. Die Балко und verwandten Körper. Ibid. 329-335.

[Vor. 12, 1925]
196 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 10
(Camera-lucida drawings, X 880)

Figs. 1-5. Capsicum annuum: figs. 1-3, successive changes in plastid shapes
during ripening; fig. 1, from green pulp; f 2, from yellow-orange pulp; fig. 3,
ise epicarp of ripe Íruit; fig. 4, from microtome sections of ripe pepper stained with

iron-haematoxylin; fig. 5, bom free-hand sections, stained in acid fuchsin.

Figs. 6-8. Rosa rugosa. Successive changes in plastid shapes during ripening:
fig. б from green pulp; fig. 7, from yellow-orange pulp; fig. 8, from ripe pulp.

ANN. Mo. Bor. Garp., Vor. 12, 1925 PLATE 10

HOWARD—CAROTINOIDS IN FRUITS

[Vor. 12, 1925]
198 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 11
(Camera-lucida drawings, Х 880)

Figs. 1-3. Solanum Lycopersicum: figs. 1 and 2, successive changes in plastid
shapes during ripening; fig. 1, from green pulp; fig. 2, from yellow-orange pulp;
fig. 3, from ripe pulp, bowing Snare crystals.

Figs. 4-5. Aglaonema Treubii: fig. 4, from green pulp, unstained free-hand
sections, mounted in water; fig. 5, ripe ie showing natural lycopersicin crystals. |

Figs. 6 and 7. Solanum Dulcamaora. Unstained, free-hand sections, ниш
in water: fig. 6, from yellow-orange pulp; fig. 7, from epicarp of ri

Fig. 8. — Godseffiana. Unstained, free-hand sections, mounted in i.
from ripe frui

Fig. 9. Nr americana. From unstained smears of pulp of ripe fruit.

PraTE 11

1925

,

ANN. Mo. Bor. Garp., Vor. 12

HOWARD—CAROTINOIDS IN FRUITS

[Vor. 12, 1925]
200 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 12
(Camera-lucida drawings, X 880)

Figs. 1-5. Lycium halimifolium: figs. 1-3, successive changes in plastid shapes
during ripening; fig. 1, from green pulp; fig. 2, from yellow-orange pulp; fig. 3, from
red-ripe pulp; fig. 4, unstained chromoplasts from ripe juice; fig. 5, chromoplasts
stained with acid fuchsin from ripe juice.

Figs. 6-9. Asparagus officinalis: figs. 6-8, successive changes in plastid shapes
during ripening; fig. 6, from green pulp; fig. 7, from yellow-orange pulp; fig. 8,
chromoplasts stained with acid fuchsin from ripe juice; fig. 9, from ripe juice.

ANN. Mo. Bor. Garb., Vor. 12, 1925 PLATE 12

HOWARD—CAROTINOIDS IN FRUITS

[V or. 12, 1925]
202 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 13
(Camera-lucida drawings, Х 880)

Figs. 1 and 2. Sorbus sitchensis. From green and ripe pulp respectively.
Figs. 3 and 4. Arisaema triphyllum. From ripe juice and pulp respectively.
Fig. 5. Viburnum Opulus. From ripe pulp.
Fig. 6. Crataegus phaenopyrum. From ripe pulp.
Figs. 7 and 8. Celastrus scandens: fig. 7, from ripe juice stained with acid fuchsin;
fig. 8, from ripe pulp.
Fig. 9. Solanum Pseudo-capsicum. From juice of ripe fruit.

[Vor. 12, 1925]
204 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 14
(Camera-lucida drawings, X 880)

Fig. 1-3. Evonymus europaea: fig. 1, a, b, c, and d, successive о! in the
formation of crystals by the addition ol alcoholic potash in ripe tissue; fig. 2, from
ripe fruit showing chromoplasts; fig. 3, crystals obtained by the addition of alcoholic
potash.

Figs. 4 and 5. Lonicera sp. From pulp and juice respectively of ripe fruit.

Figs. 6 and 7. Citrullus vulgaris: fig. 6, from ripe pulp showing lycopersicin
crystals; fig. 7, from green portion of pulp, showing chloroplasts.

D., Vou. 12, 1925

ANN. Mo. Bor. Gar

HOWARD—CAROTINOIDS IN FRUITS

[Vor. 12, 1925]
206 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 15
(Camera-lucida drawings, X 880)

Fig.l. Lonicera sp. Yellow crystals obtained by the addition of alcoholie potash.
Fig. 2. Lycium halimifolium. Orange crystals obtained by the addition of
glycerine potash.
Fig. 3. Crataegus phaenopyrum. Deep red crystals obtained by the addition
of ки potash.
g.4. Rhus canadensis. Carmine-red crystals obtained by the addition of
а potash.
Fig. 5. Celastrus scandens. Orange and yellow crystals obtained by the addition
of alcoholie potash.
Fig. 6. Capsicum annuum: (a) carmine-red crystals obtained by the addition of
alcoholic potash; (b) orange crystals obtained by the addition of alcoholic potash.
g. 7. Sorbus sitchensis. Pale orange crystals obtained by the addition of
seat potash.
Fig. Solanum Dulcamara. Carmine-red crystals obtained by the addition
of sha potash..
. Figs. 9 and 10. Daucus Carota. Natural carotin crystals.
Fig. ll. Asparagus — Red-orange crystals obtained by the addition
of alcoholic potash.

ANN. Mo. Bor. Garp., Vor. 12, 1925 Prate 15
V wo o oo
| CAET уунар",
. |
2

[Vor. 12, 1925]
208 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 16
(No camera-lucida drawings)
Daucus Carota

Figs. 1-8. Free-hand sections, unstained, mounted in water from mature root:
figs. 1 and 2, from cortex and vascular cylinder respectively, near top of carrot;
figs. 3 and 4, from cortex and vascular cylinder respectively, from region just below
preceding figures; figs. 5 and 6, from cortex and vascular c ylinder respectively,
from region just below preceding figures; figs. 8 and 7, from cortex and vascular
cylinder respectively, from region nearest tip of root.

Fig. 9. From young roots, irregularly shaped orange bodies with starch grain
inclusions, treated with IKI.

Fig. 10. Cross-section .5 mm. thick, under low power, showing distribution of
stals.

Fig. 11. Cross-section of fibro-vascular bundle in young root, showing plastids
and carotin crystals.

Fig. 12. From young root, showing leucoplasts and starch inclusions.

Fig. 13. From young root, showing chloroplasts stained with iron-haematoxylin.

PLATE 16

ANN. Mo. Bor. Garb., Vor. 12, 1925

тыз

13

HOWARD—CAROTINOIDS IN FRUITS

[Vor. 12, 1925]
210 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE
PLATE 17
(All photomicrographs, Х 800)
Fig. 1. Solanum Lycopersicum. Natural lycopersicin кулаш in ripe fruit.

Fig. 2. Solanum Dulcamara. Lycopersicin crystals obtained by the addition of
alcoholic potash.

Fig. 3. Evonymus americana. Crystals obtained by the addition of alcoholic
tash

Fig. 4. Evonymus europaea. Crystals obtained by the addition of alcoholic

ANN. Мо. Bor. Gard., Vor. 12, 1925 PLATE 17

HOWARD-——CAROTINOIDS IN FRUITS

[Vor. 12, 1925]
212 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE

PLATE 18
Fig. 1. Dehydrater.
Fig. 2. Solanum Pseudo-capsicum. Crystals obtained by the addition of alcoholic

potash,
Fig. 3. Celastrus scandens, showing chromoplasts, unstained.

Q
м

IN FRUIT

S

HOWARD-—CAROTINOID:

SI GV Ig GZ6I ‘ZI "10A "auvr) тоң ‘о ‘мму

Annals
of the
Missouri Botanical Garden

Vol. 12 SEPTEMBER, 1925 No. 3

THE THELEPHORACEAE OF NORTH AMERICA. XIV!
EDWARD ANGUS BURT

Mycologist and Librarian to the Missouri Botanical Garden
Professor in the Henry Shaw School of Botany of
ashington University

PENIOPHORA

Peniophora Cooke, Grevillea 8: 20. pl. 122-125. 1879; басс.
Syll. Fung. 6: 640. 1888; Massee, Linn. Soc. Bot. Jour. 25
140. pl. 47, f. 14-19. 1889; Karsten, Finska Vet.-Soc. Bidrag
Natur och Folk 48: 421. 1889; Engl. & Prantl, Nat. Pflanzen-
fam. (1:1**): 119. 1898; Bourdot & Galzin, Soc. Мус. Fr. Bul.
28: 372. 1913; Burt, Mo. Bot. Gard. Ann. 1: 191, 193, 198.
1914; Rea, Brit. Ваза. 687. 1922.—Кпе а (in part) Bresadola,
Ann. Myc. 1: 99. 1903.—Includes Gloeopeniophora v. Hóhnel
& Litschauer, K. Akad. Wiss. Wien Sitzungsber. 116:815. 1907.
—Includes in part Gloeocystidiwm Karsten, Finska Vet.-Soc.
Bidrag Natur och Folk 48: 429. 1889.

Fructifications waxy, coriaceous, cartilaginous, membrana-
ceous, submembranaceous, floccose, or filamentous, always re-
supinate, effused, even; simple basidia with 2-4 white spores;
cystidia incrusted or not incrusted, present in the hymenium and
often more or less immersed in the substance; substance variously
differentiated in some species but not containing colored, stellate
organs.—Distinguished from Corticium by the presence of cys-
tidia.

The North American species of Peniophora are here arranged in
five groups according to the color of the substance of the fructi-
fication, nature of the cystidia, and presence or absence of gloeo-
cystidia. These groups are further subdivided to such degree as

1 Issued February 18, 1926.

Акм. Mo. Bor. Garb., Vor. 12, 1925 (213)

[Vol. 12
214 ANNALS OF THE MISSOURI BOTANICAL GARDEN

seems desirable by color of hymenium, adnation to substratum,
and incrustation of cystidia or of hyphae, into minor groups of so
few species that the characters of the component species of any
group within which a species seems to belong, may all be con-
sidered in determining the probable species of the specimen in
course of identification. An appalling amount of time and
labor has been required for the accumulation from sectional prep-
arations of the structural characters of the individual specimens
of species of Peniophora listed in this work. The older descrip-
tions of resupinate Hymenomycetes were based on so few definite
characters that a specimen in hand might seem to be referable
equally well to more than one published species, or that several
specimens in hand and certainly different specifically might all
seem referable to one species, judging from its published de-
scription. However, by the addition of the knowledge of the
definite features of structure characteristic of species, deter-
mining additional characters have been found for so many species
that the specific taxonomy of the large genus Peniophora in
North America becomes practicable. In the use of this work
sectional preparations of fertile specimens are necessary.

Of the 120 species of Peniophora described herein, 36 occur in
Europe as well as in North America and 11 others have been
already recognized as North American species. 'The remaining
73 species are unlike those which the writer has been able to
recognize among the known species from other regions of the
world and have therefore to be described as new. It is quite
probable that nearly all of these 73 species will bear the test of
study by foreign mycologists and be demonstrated eventually to
be really new, for most of them are of local occurrence, known
from a single collection and distributed with surprising uniformity
over the great area of North America which has in its different
regions such great differences in temperature, moisture, altitude,
and composition of its forests that the conditions are ideal for
the origin and survival of species of merely local distribution.
This is in accord with the fact that 9 of our new species occur in
more than one state and that others occur as follows:—7 from
Louisiana; 4 each from New York, British Columbia, Washing-
ton, Mexico, and Jamaica; 3 each from Vermont, Idaho, Oregon,

1925]
URT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 215

and Texas; 2 each from Canada, New Hampshire, Alabama,
Florida, Colorado, and Porto Rico; and 1 each from Virginia,
Georgia, Kentucky, Montana, Alaska, California, New Mexico,
Nicaragua, Cuba, and Bermuda.

It might be well for American students who use this work for
the determination of their gatherings of Peniophora to concen-
trate their attention in first attempts on the characters of the
species of wide distribution and on such of the new species as are
local in their respective regions.

Mycologists, with special knowledge of the Thelephoraceae in
every nation, work not only to make better known the fungous
floras of their own countries, but also to determine which of their
species occur in other countries. As an aid to such study in the
future and for checking my work, there is here given the following
list of foreign species of Peniophora, not known to me except from
the more or less satisfactory published descriptions, viz., Penio-
phora abietis, P. amaniensis, P. atro-cinerea, P. ранах Р.
bambusicola, P. carneola, P. Сћееѕзтатії, Р. cineracea, P. citrina,
P. coccinea, P. Coffeae, P. convolvens, P. Corsica, P. crustosa, P.
difissa, P. discoidea, P. Dussii, P. fimbriata, P. gigaspora, Р.
habgallae, P. leprosa, P. lilacea, P. Martelliana, P. mimica, P.
ochroleuca, P. orphanella, P. pirina, P. rimicola, P. rufomarginata,
P. sordidella, P. sororia, P. sparsa, P. subavellanea, P. subglebulosa,
P. sublaevis, P. subtilis, P. tomentella, P. tremelloidea, and P. ver-
micularis. It is probable that most of these species are of local
interest but I should like to have been sure that I have not re-
described any of them as new American species. Much time and
effort have been required to make this list as small as it is.

KEY TO ARRANGEMENT OF THE SPECIES
I. Substance not colored, with the usual cystidia, no gloeocystidia.

1. Hymenium white or w .
*At least small pieces separable when moistened.

a. Cystidia шегамей....;.. ccc e eect иене 1-7
b. Cystidia not inerusted. . ............ en е 8,9
**Closely adnate, not separable.
a. Antler-shaped paraphyses not present.
Суваа ТО 2. or eae Reine. 10-18
t{Cystidia rough-walled or denticulate.................. 14, 15
titCystidia not incrusted, even.......... een ee

b. Antler-shaped paraphyses ргеѕепі.......................:

[Vol. 12
216 ANNALS OF THE MISSOURI BOTANICAL GARDEN

2. Hymenium colored and the subhymenium also in a few species.
*Closely adnate, not separable.
a. Cystidia not incrusted.

etification not stratose...... . 28-30
ttFructification becoming stratose. . 81, 88
b. Cystidia incrusted...... . 83-38

**At least small pieces separable when moistened.
dia incrusted.

а. Cysti
THopbae ПИРИНЕН; О pee es ce 39-44
TtHy ph ti ted t obvi ly incrusted CAREN 45-52
b. Cystidia not incrusted.
Tiris о с Lait 53-56
{tHyphae not incrusted or not obviously incrusted.... 57-61
II. Substance not colored almost without exception, cystidia very
long, cylindric, thick-walled, not normally incrusted, often visible
through ce oe of the fructification, no gloeocystidia—the
и 62-68
ш. Substance not кайшы. gloeocystidia present as well as cystidia—the Е
gloeocystidial group.
1. At least small pieces separable when moistened.
*Cystidia inerusted. -
{Some cystidia 40-100 x 20-50 ,....................... 69
idin of thé ШИШ М рса Е jap
WC ations ВОС DU. ole si cea hs E Cm vLex
2. Closely adnate, not верагаре.............................. 84-90, Та
IV. Substance yellow or yellowish rather than dark-colored and often
bleached by potassium hydrate solution.
1. At least small pieces separable when moistened................ 91-97

4. Себу adnate, not eepirable. 1... узуу. orco her 98-100
V. Substance more or less dark-colored, the dark color retained in pre-
parations stained with eosin. Gloeocystidia sometimes present.
1. Fructifieation stratose.....
2. пса not stratose.
east small pieces separable when moistened. Compare also

**Closely adnate, not ла. аваас CUPIT. ae ; 111-120

(Including as 117-120 the Р. ‹ cinerea group with opaque zone
next to substratum and the largest cystidia on this zone.)

i: веза gigantea (Fr.) Massee, Linn. Soc. Bot. Jour. 25:

3:113. 1897; amen & Galzin, Soc. Myc. Fr. Bul. 28: 401.
1913; Rea, Brit. Basid. 693. 1922.

Thelephora gigantea Fries, Obs. Myc. 1:152. 1815;Syst. Myc.
1: 448. 1821.—Corticium giganteum Fries, Epicr. 559. 1838;
Hym. Eur. 648. 1874; Peck, N. Y. State Mus. Rept. 28: 52.
1876; Sacc. Syll. Fung. 6: 610. 1888

1925]
URT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 217

Illustrations: Fries, Icones Hym. 2: pl. 197, f. 3.

Fructifications broadly effused, hyaline, white, waxy, swelling
when moist and separable from substratum, when dry horn-like
and parchment-like, the hymenium even, pale pinkish buff, pale
olive-buff, or pallid mouse-gray in the herbarium, the margin
white, fibrillose, radiating, sometimes becoming free and curling
away from the substratum in drying; in section 100-500 џ thick,
not colored, with the broad layer towards the substratum com-
posed of crowded and more or less longitudinally arranged
hyphae so highly gelatinously modified that only the lumen and
cell contents usually show distinctly in preparations, about
3-5 y in diameter; cystidia incrusted, about 40-50 x 8-12 џ,
confined to the hymenium or a zone up to 100 џ broad; spores
hyaline, even, about 414-5 x 214-3 u as found in preparations.

Fructifications 3-30 cm. in diameter.

On bark and wood of dead conifers such as Pinus, Abies, and
Tsuga. In Europe, Canada to Texas, westward to the Pacific
states, in Mexico, and in Japan. June to January. Widely dis-
tributed and abundant locally.

P. gigantea may usually be recognized at sight by its occurrence
on coniferous bark in large, whitish or pinkish buff fructifications
of cartilaginous structure, separable from the substratum and
more or less curling away from it in drying.

Specimens examined:

Exsiccati: Bartholomew, Fungi Col., 2422, 4242, 4622; Ellis, N.
Am. Fungi, 410; Krieger, Fungi Sax., 117; Ravenel, Fungi Am.,
452; Fungi Car. 2: 38; Romell, Fungi Scand., 34; Sydow, Myc.
Germ., 553; de Thümen, Мус. Univ., 909.

Finland: Mustiala, P. A. Karsten, in Myc. Univ., 909.

Sweden: L. Romell, 97, 98, 349, and in Romell, Fung. Scand., 34.

Germany: Brandenburg, H. Sydow, in Sydow, Муе. Germ., 553;
Königstein, W. Krieger, in Krieger, Fungi Sax., 117.

Austria: Karwendel, Tirol, V. Litschauer; Sabai Tirol, V.
Litschauer.

Italy: Trient, G. Bresadola.

France: Fautrey, comm. by Lloyd Herb., 4354; Jura, N. Patouil-
lard (in N. Y. State Mus. Herb., and Mo. Bot. Gard. Herb.,
55907).

[Vol. 12
218 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Canada: Montreal, on timbers in a mill, R. J. Blair, 887, comm.
by L. O. Overholts, 4113 (in Mo. Bot. Gard. Herb., 55632);
Belleville, Ontario, J. Macoun, 253; Ottawa, J. M acoun, 48.

New Hampshire: Chocorua, W. G. Farlow, 32.

Massachusetts: Fall River, on floor beams in a mill, W. H. Snell
(in Mo. Bot. Gard. Herb., 57379).

New York: ex herb. Torrey, 112 (in N. Y. Bot. Gard. Herb., and
Mo. Bot. Gard. Herb., 61437); Beaver River, Adirondacks,
G. F. Atkinson, 4606; Mechanicville, C. H. Peck (in N. Y.
State Mus. Herb., and Mo. Bot. Gard. Herb., 55575).

New Jersey: Newfield, J. B. Ellis, in Ellis, N. Am. Fungi, 410.

Pennsylvania: Adelaide, on cross ties, H. v. Schrenk (in Mo. Bot.
Gard. Herb., 6685); State College, L. O. Overholts, 4810 (in
Mo. Bot. Gard. Herb., 56124).

Maryland: Takoma Park, C. L. Shear, 1266, and in Bartholomew,
Fungi Col., 2422.

Virginia: Rio, F. Gravatt (in Mo. Bot. Gard. Herb., 44042).

North Carolina: Biltmore, Е. Bartholomew, 5658 (in Mo. Bot.
Gard. Herb., 44216), and in Bartholomew, Fungi Col., 4622;
Chapel Hill, J. N. Couch, comm. by Univ. N. C. Herb., 4306
(in Mo. Bot. Gard. Herb., 57422). —

South Carolina: H. W. Ravenel, in Ravenel, Fungi Car. 2: 38;
Aiken, H. W. Ravenel, in Ravenel, Fungi Am., 452.

Georgia: Brunswick, H. v. Schrenk (in Mo. Bot. Gard. Herb.,
43887).

Florida: W. W. Calkins, 63 (in N. Y. Bot. Gard. Herb., and Mo.
Bot. Gard. Herb., 61446).

Louisiana: Shreveport, E. Bartholomew, in Bartholomew, Fungi
Col., 4242, and (in Mo. Bot. Gard. Herb., 4943).

Texas: Houston, H. W. Ravenel, 221; Quitman, W. H. Long,
12074, and comm. by C. J. Humphrey, 2552 (in Mo. Bot.
Gard. Herb., 55046 and 9804 respectively); Somerville, H. v.
Schrenk (in Mo. Bot. Gard. Herb., 42884).

Ev Bay City, J. R. Weir, 301 (in Mo. Bot. Gard. Herb.,

Wisconsin: Madison, on log of Pinus Taeda in timber yard, M. C.
Jensen, eomm. by C. J. Humphrey, 719 (in Mo. Bot. Gard.
Herb., 42729).

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 219

Minnesota: Cass Lake, J. R. Weir, 394 (in Mo. Bot. Gard. Herb.,
13981); St. Louis River, J. C. Arthur, L. Н. Bailey & Е. W. D.
Holway, 175 St. (in Mo. Bot. Gard. Herb., 4821).

Arkansas: Texarkana, on cross ties, H. v. Schrenk (in Mo. Bot.
Gard. Herb., 56378). :

Colorado: Tolland, Г. О. Overholis, 1834 (in Mo. Bot. Gard.
Herb., 54879).

Montana: Libby, E. E. Hubert, comm. by J. R. Weir, 11449 (in
Mo. Bot. Gard. Herb., 63276); Rockhill, E. E. Hubert, comm.
by J. R. Weir, 11966, 11979, 11984 (in Mo. Bot. Gard. Herb.,
63336-8).

Idaho: Coolin, J. R. Weir, 11097, comm. by U. S. Dept. Agr.,
Path. Мус. Coll., 1335 (in Mo. Bot. Gard. Herb., 62988);
Priest River, J. В. Weir, 80 (in Burt Herb.) and 5825, 6331,
11985, 12018, 14938 (in Mo. Bot. Gard. Herb., 58289, 55950,
63350, 63374, and 56800 respectively); Santa, E. E. Hubert,
comm. by J. R. Weir, 11603 (in Mo. Bot. Gard. Herb., 63304).

British Columbia: Hastings, J. Macoun, 24.

New Mexico: Gila National Forest, G. G. Hedgcock & W. H.
Long, comm. by C. J. Humphrey, 2524 (in Mo. Bot. Gard.
Herb., 21669).

Mexico: Orizaba, Nuevo, W. A. & E. L. Murrill, 767, comm. by
N. Y. Bot. Gard. Herb. (in Mo. Bot. Gard. Herb., 54649).

Japan: Sendai, A. Yasuda, 76 (in Mo. Bot. Gard. Herb., 56315).

2. P. globifera Ellis & Everhart, Am. Nat. 1897: 340. 1897;
Sacc. Syll. Fung. 14: 224. 1899.

Type: in N. Y. Bot. Gard. Herb. and a fragment in Burt
Herb.

Fructifications broadly effused, separable when moistened,
when dry horn-like, whitish to pale smoke-gray, the hymenium
with somewhat convex granules, bristling with the crowded
cystidia; in section 400-500 » thick, not colored, with a layer
about 200-300 » thick towards the substratum composed wholly
of densely interwoven, hyaline hyphae 3-5 а in diameter, with
the walls so gelatinously modified as to be very indistinct; no
gloeocystidia; cystidia heavily and coarsely incrusted, 30-70
X 12-15 в, very abundant both in the hymenium and im-

[Vol. 12
220 ANNALS OF THE MISSOURI BOTANICAL GARDEN

mersed throughout an outer zone up to 150 u thick; basidiospores

hyaline, even, 4-6 x 2-214 y.

Fructifications up to 10 cm. in diameter.

On bark of conifers. New York and Ontario, and in Montana,
Idaho, British Columbia, Oregon, and New Mexico. August to
October. Rare. .

P. globifera is either known only from the type collection or else
it is an extreme form of P. gigantea, for each of the more recent
gatherings cited below has some character approaching P. gi-
gantea. The distinctive features of the type of P. globifera are
whiter color, much more numerous and larger cystidia which are
also more coarsely incrusted, and a hymenial surface with some
convex granules like those of Grandinia granulosa. The original
description is erroneous in stating that the fructifications are
closely adnate and that the spores are globose and 3 џ in diameter.

Specimens examined:

Ontario: Ottawa, McKay’s Lake, J. Macoun, 175, type (in N. Y.
Bot. Gard. Herb.) and a specimen from the type collection,
comm. by J. Macoun; Harraby, E. T. & S. A. Harper, 680;
St. Lawrence Valley, J. M. acoun, 85.

New York: Mt. McIntyre, C. H. Peck (in N. Y. State Mus. Herb.,
and Mo. Bot. Gard. Herb., 55777).

Montana: Rockhill, E. Е. Hubert, comm. by J. R. Weir, 11960
(in Mo. Bot. Gard. Herb., 63317).

Idaho: Coolin, J. R. Weir, 11158M, 11491 (in Mo. Bot. Gard.
Herb., 63254, 63279); Priest River, J. R. Weir, 20.

British Columbia: Kootenai Mts. near Salmo, J. R. Weir, 517,
526 (in Mo. Bot. Gard. Herb., 5068, 19430).

Oregon: Portland, C. J. Humphrey, 6126.

New Mexico: Clouderoft, W. H. Long, 19524 (in Mo. Bot. Gard.
Herb., 44765).

3. P. arachnoidea Burt, n. sp.

Type: in N. Y. Bot. Gard. Herb. and Burt Herb.

Fructifications effused, very thin, fragile, small pieces separ-
able, white, becoming cartridge-buff in the herbarium, the hy-
menium continuous, not shining, fragile, loosely supported on an
arachnoid subiculum, the margin delicately fibrillose or arach-

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 221

noid; in section 150-300 y thick, not colored, 2-layered, with the

layer next to the substratum composed of very loosely arranged,

nodose-septate, hyaline hyphae 3-4 y in diameter and with the
hymenial layer about 50,» thick, compact; no gloeocystidia;
cystidia numerous, tapering, rough or granule-incrusted near the
tips, 4—6 y in diameter, protruding up to 40 y beyond the basidia;
spores hyaline, even, 3—4 X 2-2l5 y, copious.

Fructifications 1-4 cm. in diameter.

On bark of fallen limbs of Populus and Alnus. New Hamnahite,

New York, Alabama, and Oregon. October and November. Rare.
P. arachnoidea has the aspect of Corticium arachnoideum but

is a Peniophora. The hyphae and their arrangement are like

those of Р. cremea. The microscopic characters are so similar to
those of Coniophora byssoidea that the gatherings from northern
localities may possibly be white forms of the latter which I have
erroneously included under P. arachnoidea.

Specimens examined:

New Hampshire: Hanover, on Populus, G. R. Lyman, 27 (in
N. Y. Bot. Gard. Нењ, Mo. Bot. Gard. Herb., 61589, and
Burt Herb.).

New York: Ithaca, G. F. Atkinson, 2589, formerly referred by
me to C. byssoidea; Karner, on Populus, H. D. House (in N. Y.
State Mus. Herb., and Mo. Bot. Gard. Herb., 54376).

Alabama: Auburn, on Alnus, F. S. Earle, 97, type, comm. by
N. Y. Bot. Gard. Herb.; Montgomery County, R. P. Burke,
367 (in Mo. Bot. Gard. Herb., 57234).

British Columbia: Salmo, J. R. Weir, 472, 480 (in Mo. Bot. Gard.
Herb., 63357, 63387).

Oregon: White Pine, on hymenium of Thelephora terrestris, J. R.
Weir, 620 (in Mo. Bot. Gard. Herb., 13999).

4. P.inconspicua (B. & C.) Massee, Linn. Soc. Bot. Jour. 25:
149. pl. 47, f. 14. 1889; Sace. Syll. Fung. 21: 410. 1912.

Corticium inconspicuum Berkeley & Curtis, Linn. Soc. Bot.
Jour. 10: 336. 1868; Sacc. Syll. Fung. 6: 615. 1888.

Type: in Kew Herb. and Curtis Herb.

Fruetifications effused, thin, small, orbicular, gregarious, be-
coming confluent, membranaceous, small pieces separable, white

[Vol. 12
222 ANNALS OF THE MISSOURI BOTANICAL GARDEN

when fresh, becoming light buff in the herbarium, even, setulose
with the cystidia, the margin composed of radiating hyphae; in
section 250-300 џ thick, not colored, composed of densely inter-
woven, rather thick-walled and rigid, ascending hyphae 3-4 y in
diameter, not incrusted; no gloeocystidia ; cystidia incrusted,
fusoid, 50-60 x 12-15 и, scattered in the surface of the hymen-

ium; spores in a crushed preparation are hyaline, even, 4 x 3 y,

so few that they may not belong.

Fructifications about 2 mm. in diameter, becoming more or less
confluent over areas up to 3 cm. long and 1 cm. wide.

On bark of dead frondoselimbs. West Indies. December and
March.

P. inconspicua has small clustered fructifications becoming
confluent and very large cystidia scattered along the surface of
the hymenium and none wholly immersed. In one portion of
my sections the hyphae next to the substratum are slightly
brownish and suggestive of those of a resupinate Stereum but I
do not recall an effuso-reflexed Stereum of which P. inconspicua
may be the resupinate fructification.

Specimens examined:

Cuba: Mountain of Rangel, C. Wright, Fungi Cubenses Wrighti-
ani, 841, type (in Kew Herb., Curtis Herb., Mo. Bot. Gard.
Herb., Burt Herb., and an unnumbered portion in N. Y. Bot.
Gard. Herb.).

Porto Rico: Rio Piedras, J. R. Johnston, 1664, comm. by J. A.
Stevenson (in Mo. Bot. Gard. Herb., 13223).

5. P. galochroa Bresadola, Hedwigia 35: 200. 1896; Sacc.
Syll. Fung. 14:224. 1899.

Type: a part in Burt Herb.

Broadly effused, membranaceous, small pieces separable, from
white becoming pinkish buff, finally cracked and silky along the
crevices, the margin somewhat fimbriate at first, soon similar; in
section 250—400 ų thick, not colored, with hyphae rather stiff,

substratum, densely interwoven in the broad middle region, the
hymenial layer about 50 u thick; по gloeocystidia ; cystidia coarsely

1925]
URT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 223

incrusted, fusiform, 25-50 х 9-10 џ, barely protruding, usually
immersed in all parts of the hymenial layer; spores published by
Bresadola as 544-614 x 4-414 u but I find the type sterile.

Fructifications 3-6 cm. long, 15-21% ст. wide.

On bark of decaying branches and on wood. Brazil ur West
Indies. August to December. Rare.

In aspect P. galochroa is somewhat suggestive of бо
portentosum but much thinner and very different in structure by
the presence of cystidia which are confined to a hymenial layer
not more than 50 џ thick in the specimens studied. The speci-
mens from the West Indies, which I have referred to P. galochroa,
have subglobose spores about 3 и in diameter and are perhaps
a distinct species if P. galochroa has as large spores as published
by Bresadola, but ‘T find the portion of the type communicated
to me wholly sterile.

Specimens examined:

Brazil: Blumenau, A. Méller, type, from Bresadola.

Jamaica: Chester Vale, W. A. & E. L. Murrill, 331, 751, comm.
by N. Y. Bot. Gard. Herb.

Porto Rico: Rio Piedras, J. A. Stevenson, 2985 (in Mo. Bot. Gard.

Herb., 7799).

6. P. odontioides Burt, n. sp.

Type: in Mo. Bot. Gard. Herb.

Fructifications effused, very thin, arachnoid-membranaceous,
tender, small pieces separable when moistened, white, even, not
shining, the margin thinning out, fibrillose; in section 50-130 y.
thick, not colored, composed of thin-walled, loosely interwoven,
suberect hyphae about 4-414 y. in diameter, incrusted, becoming
collapsed; no gloeocystidia; cystidia of Odontia type, trans-
versely septate, cylindric-obtuse, 8 џ in diameter, protruding up
to 45 u, not incrusted or with a few incrusting granules; spores
hyaline, even, 9-12 x 4-415 y, copious.

Fructifications in fragments which are 2-3 cm. long, 5-6 mm.
wide.

On decaying frondose wood. Canada. July to September.

P. odontioides is distinguished among our thin, white species
by having large, cross-septate cystidia such as are common in

[Vol. 12
224 ANNALS OF THE MISSOURI BOTANICAL GARDEN

many species of Odontia where they are clustered together in the
granules, but in the present species such cystidia are distributed
along an even hymenium devoid of granules.
Specimens examined:
Canada: J. Macoun, 20; St. Lawrence Valley, J. Macoun, 14.
Manitoba: 52° 15’ north latitude, Swan River, G. В. Bisby, 1047,
type (in Mo. Bot. Gard. Herb., 59034).

7. P.exigua Burt, n. sp

Type: in Mo. Bot. Gard. Herb. and N. Y. Bot. Gard. Herb.

Fructifications effused, small, circular, gregarious, thin, some-
what membranaceous, tender, small pieces separable when
moistened, snow-white, even, contracting in drying and cracking
into polygonal masses 1-2 mm. in diameter, with the white
arachnoid subiculum visible on the sides of the fissures, the
margin narrow, white, arachnoid; in section 150-180 и thick,
not colored, with some hyphae densely arranged parallel with
the substratum and then ascending and loosely interwoven to the
hymenial layer, about 3 y in diameter, thin-walled, not nodose-
septate, perhaps slightly incrusted in the hymenial layer; no
gloeocystidia; cystidia inerusted, cylindric, 30-60 х 6-7 и, con-
fined to the hymenial layer and usually wholly immersed, a few
protruding up to 12 » beyond the basidia; spores hyaline, even,
4-5 х 2% u.

Fructifications 1-12 mm. in diameter—8 in an area 4 X 1
em. and probably becoming confluent.

On bark of dead, fallen limbs, about 12 mm. in diameter, of &
frondose species. Mexico. December.

P. exigua is distinguished among our species by its clustered,
small, aes cai fructifications which crack into small polygonal

бо ЧН examined:

Mexico: near Guernavaca, altitude 4500 m., W. A. & E. L. Mur-
rill, 377, type, and 378 (in Mo. Bot. Gard. Herb., 54474,
54473, respectively).

8. Р, laxa Burt, п.
Type: in Mo. Bot. Gard. Herb.

1925]
URT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 225

Fructifications broadly effused, thin, waxy-membranaceous,
loosely attached to the substratum by a cottony subiculum,
tender, small pieces separable, becoming pale ivory-yellow in the
herbarium, even, not much cracked, the margin thinning out,
fibrillose, with some mycelial strands; in section 200—300 y thick,
not colored, with the thin, compact hymenium supported by a
very broad layer of loosely interwoven, thin-walled, granule-
incrusted hyphae 114-2 y, rarely 3 y, in diameter, with the
hyphae more densely arranged in a middle zone of this layer; no
gloeocystidia; cystidia not inerusted or with only a few incrusting
granules, 414-6 y in diameter, protruding up to about 30-50 u
beyond the basidia, often capitate and 6-9 џ in diameter at the
apex; basidia up to 6 y in diameter, with 4 sterigmata; spores
hyaline, even, spherical, 4144-6 u in diameter, copious.

Fructifications 2-6 em. long, 1-214 cm. wide.

In woods on bark with the wood underneath wholly decayed.
British Columbia. December.

P. laxa is probably white when growing and assumed the pale
ivory-yellow tint in the herbarium; the aspect is like that of P.
arachnoidea but with globose spores. Р. sphaerospora of Europe
has similar spores but much coarser, erect hyphae and different
cystidia.

Specimens examined
British Columbia: Sidney, J. Macoun, 8, type (in Mo. Bot. Gard.

Herb., 5767).

9. P.humifaciens Burt, n. sp.

Type: in Burt Herb.

Fructifications effused, circular, thin, membranaceous, loosely
attached by white rhizomorphie strands from the substratum,
easily separable, white, becoming somewhat pale pinkish buff in
the herbarium, the margin thinning out, floccose; in section
150 y thick, not colored, with hyphae hyaline, up to 5 u in diameter
and coarsest next to the substratum, very loosely arranged,
branching and becoming 3 y in diameter towards the hymenial
layer, nodose-septate, with few incrusting granules; no gloeo-
= hymenial layer 30—40 р, thick, continuous; cystidia not

, 3314p in diameter at base, protruding 25-40 y,

[Vol. 12
226 ANNALS OF THE MISSOURI BOTANICAL GARDEN

tapering, attenuated to a long and very sharp point; basidia
with 4 sterigmata; spores hyaline, even, subglobose, about 214
X 2 u, copious.

Fructifications 2-214 em. in diameter.

On very rotten coniferous log—perhaps Thuja. Washington.
October. Rare.

P. humifaciens was so sparingly and loosely connected with the
substratum by the white, mycelial strands that the impact of a
hatchet against the log caused fructifications to fall away. P.
arachnoidea, a related species, has quite different hyphae and
cystidia and mode of attachment.

Specimens examined:

Washington: Chehalis, C. J. Humphrey, 6266, type.

10. P. candida (Pers.) Lyman, Boston Soc. Nat. Hist. Proc.
33: 167. pl. 20, f. 44-55, pl. 26, f. 188. Е. 1907.

Aegerita candida Persoon, Roemer Neues Mag. Bot. 1: 120.
1794 (imperfect stage); Syn. Fung. 684. 1801; Fries, Syst.
Myc. 3:220. 1829; Басс. Syll. Fung. 4: 661. 1886.—Sclerotiwm
Aegerita Hoffmann, Fl. Germ. 2: pl. 9, f. 1. 1795.—Peniophora
Aegerita (Hoffm.) v. Hóhnel & Litschauer, K. Akad. Wiss. Wien
Sitzungsber. 116: 814. 1907; 123:83. 1914; Sacc. Syll. Fung.
21: 410. 1912; Bourdot & Galzin, Soc. Мус. Fr. Bul. 28: 382.
1913; Rea, Brit. Ваза. 687. 1922.—Kneiffia farinosa Bresadola,
Ann. Myc. 1: 105. 1903; Sacc. Syll. Fung. 17: 178. 1905.

Illustrations: See Sacc. Syll. Fung. 19: 25, for numerous
figures of imperfect stage.

Fructifications effused, thin, adnate, very tender, at first fari-
nose, then forming a continuous hymenium, white to pale cream-
color, very minutely velvety under a lens, the margin thinning
out, indeterminate, usually with clusters of the minute, globose,
white, imperfect stage adjoining; in section 40-100 џ thick, not
colored, with hyphae suberect, thin-walled, collapsing, of ir-
regular outline, about 4 џ in diameter; no gloeocystidia; cystidia
incrusted, scattered, starting from the substratum, 40-100 x
6-12 u; spores hyaline, even, subglobose, 6-7 x 414-6 y.

Fructifications 2-4 cm. long, 1-2 cm. wide.

On decaying wood and fallen branches of Alnus, Populus, Acer,

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 227

Ulmus, etc., and on the ground. In Europe, and from Massa-
chusetts to Missouri. October and November. Rare.

The association of the effused, white fructifications of P. can-
dida, with the clustered, small, globose, white or cream-colored
fructifications—about 5 or 6 to a mm.—of the imperfect stage,
Aegerita candida, affords an easy means of recognizing P. candida.

Specimens examined:

Poland: Eichler, part of the type of Kneiffia farinosa, comm. by

Bresadola.

France: Allier, H. Bourdot, 19908.
New Hampshire: Hanover, G. E. Lyman.
Massachusetts: Arlington, A. P. D. Piguet, comm. by W. G.

Farlow, 33; Waverly, G. R. Lyman, two gatherings.
New York: Ithaca Flats, G. F. Atkinson.

Missouri: Creve Coeur, E. A. Burt (in Mo. Bot. Gard. Herb.,

56059), and F. P. McWhorter (in Mo. Bot. Gard. Herb., 57309).

11. P.cana Burt, n. sp.

Type: in Mo. Bot. Gard. Herb. and N. Y. Bot. Gard. Herb.

Fructifications effused, closely adnate, very thin, hypochnoid,
not forming an interwoven membrane, pilose under a lens, the
margin pruinose, indeterminate; in section 10-30 thick, not
colored, consisting of short, erect, simple or once- or twice-
branched hyphae 3-314 џ in diameter, not incrusted, not nodose-
septate, and of large cystidia; no gloeocystidia; cystidia heavily

inerusted, conical, 50-60 x 10-18 в, protruding 30-45 y, start-
ing from the substratum, very numerous; spores hyaline, even,
3-3V4 x 116 u as seen on basidia.

Fructifications fragmentary, with the fragments 116-2 cm.
long, 10-15 mm. wide.

On dark, brittle wood humus—probably of a frondose species.
Florida. March.

P. cana is so thin and hoary that it is likely to be regarded as a
Hyphomycete unless examined with the microscope. The large,
conical, incrusted cystidia and small spores distinguish it from
Р; albugo.

Specimens ined:

Florida: Cutler Hummock, W. A. Murrill, 82, type, and 83,

: [Vol. 12
228 ANNALS OF THE MISSOURI BOTANICAL GARDEN

comm. by N. Y. Bot. Gard. Herb. (in Mo. Bot. Gard. Herb.,
62102, 62103).

12. P. irregularis Burt, n. sp.

Type: in Mo. Bot. Gard. Herb.

Fructifications effused, very thin, adnate, flocculent, tender,
white, interrupted, somewhat lacunose, not shining, the margin
thinning out, with hyphae interwoven; in section 45-75 y. thick,
not colored, composed of interwoven, hyaline, inerusted hyphae
275 in diameter; no gloeocystidia ; Cystidia inerusted with
coarse granules, 15-22 x 8-12 y, barely protruding, confined to
the hymenium; spores hyaline, even, 415 x 214 u, borne 4 toa
basidium.

Fructification 314 em. long and broken off at one end, 1 cm.
wide.

On bark of a rotten frondose limb about 7 mm. in diameter.
Cuba. December.

P. irregularis is a thin, white species of flocculent texture
rather than waxy, with the dark substratum visible in small
Spaces not covered by the fructification.

Specimens examined:

Cuba: near Havana, C. J. Humphrey, 2953, type (in Mo. Bot.

Gard. Herb., 9010).

13. P. albofarcta Burt, n. Sp.

Type: in N. Y. Bot. Gard. Herb., Mo. Bot. Gard. Herb., and
Burt Herb.

Fructifications effused, adnate, dry, spongy-membranaceous,
light buff to pinkish buff in the herbarium, minutely velutinous
under a lens, even, but little cracked, the margin thinning out,
minutely tomentose; in structure 200-350 и thick, not colored,
composed of a broad layer of loosely interwoven, rather rigid
hyphae 3-314 и in diameter, not incrusted, not nodose-septate,
and of a dense hymenial layer about 100 y thick; no gloeocystidia;
cystidia incrusted, slender-fusiform, 50-90 x 6-9 и, protruding
up to 30 y, numerous in all parts of the hymenial layer; spores
hyaline, even, spherical, 3-4 u in diameter, only few found but
seem to belong.

1925]
URT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 229

Fructifications in fragments 5 mm.-2 em. long, 5-10 mm. wide.

On very rotten wood of stump of orange tree (Citrus). Louisi-
ana. December.

The fructifications of P. albofarcta are scarcely distinguishable
in color from the rotten wood upon which grown. "The occurrence
on Citrus wood, velvety hymenium, globose spores, and thick and
loosely interwoven pales seem good, distinctive characters.

Specimens examin
Louisiana: Point 4 la Hache, A. B. Langlois, 894, type (in N. Y.

Bot. Gard. Herb., Mo. Bot. Gard. Herb., 63729, and Burt

Herb.).

14. Р. longispora (Pat.) v. Hóhnel, Ann. Myce. 3: 325. 1905;
Bourdot & Galzin, Soc. Мус. Fr. Bul. 28: 392. 1913; Rea, Brit.
Basid. 690. 1922.

Hypochnus longisporus Patouillard, Jour. de Bot. 1894: 221.
1894; Басе. Syll Fung. 11: 130. 1895.—Kneiffia longispora
(Pat.) Bresadola, Ann. Мус. 1: 105. 1903.

Fructifications widely effused, thin, pubescent, hypochnoid,
not separable, white, becoming pale smoke-gray to pale olive-
buff in the herbarium, the margin thinning out; in structure
30-120 » thick, not colored, composed of кареги, loosely ar-
ranged, thin-walled, rough-walled, SRE NONE 1 hyphae 21%-
3 win diameter, and of cystidia; no la acicular,
rough-walled, 40-80 x 3-4 y, protruding up “to 60 ш; spores
white in spore collection, even, 6-15 X 214-3 y.

Fructifications 3-10 cm. long, 1-5 ст. wide.

On bark and decaying wood of frondose species usually—es-
pecially Populus—rarely on conifers. In Europe and Africa and
from Maine to Louisiana, Montana to Washington, and in the
West Indies. July to March. Frequent.

P. longispora is well marked by its thin, white fructifications,
hyphae, and needle-shaped cystidia rough or somewhat barbed
with minute erystals, and the slender spores. There are few
resupinate species which may be more confidently recognized.

Specimens examined:

Sweden: Lappland, L. Romell, 316; Stockholm, L. Romell, 411.
Poland: Russian Poland, Eichler, comm. by С. Bresadola.

[Уо]. 12
230 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Austria: Innsbruck, Tirol, V. Litschauer.

France: Allier, H. Bourdot, 4073, 20807; Aveyron, A. Galzin,
11842, 17636, comm. by H. Bourdot, 20861, 20862.

England: Doncaster, E. M. Wakefield (in Мб. Bot. Gard. Herb.,
57118).

Maine: Kittery Point, В. Thaxter & E. A. Burt.

New York: East Berne, C. H. Peck (in N. Y. State Mus. Herb.,
and Mo. Bot. Gard. Herb., 14859).
Florida: Royal Palm Hummock, W. A. Murrill, 105, comm. by
N. Y. Bot. Gard. Herb. (in Mo. Bot. Gard. Herb., 62105).
Louisiana: Baton Rouge, C. J. Humphrey, 2529 (in Mo. Bot.
Gard. Herb., 20690); St. Martinville, A. B. Langlois, br, dk.
Montana: Columbia Falls, C. J. Humphrey, 7239 (in Mo. Bot.
Gard. Herb., 12526).

Idaho: Coolin, J. R. Weir, 11255, 11541 (in Mo. Bot. Gard.
Herb., 63259, 63296).

Washington: Olympia, C. J. Humphrey, 6339.

Cuba: C. G. Lloyd, 436 (in Mo. Bot. Gard. Herb., 55169).

Jamaica: Blue Hole, W. A. Murrill, 231, comm. by N. Y. Bot.
Gard. Herb.

Grenada: Grand Etang, R. Thazter, comm. by W. G. Farlow, 5.

15. Р. asperipilata Burt, n. sp. ,

Type: in Burt Herb. and U. 8. Dept. Agr. Herb.

Fructifications effused, very thin, closely adnate, snow-white,
velvety, the margin thinning out; in section 45-60 y. thick, not
colored, composed of somewhat erect, loosely interwoven, thin-
walled hyphae 3 y in diameter, not incrusted, occasionally nodose-
septate, which terminate in cystidia and clusters of basidia form-
ing a hymenium barely continuous; no gloeocystidia; cystidia
hair-like, slender, tapering to a sharp point, conspicuously den-
ticulate or rough, 30-60 x 3-5 y, protruding up to 50 и beyond
the basidia, very numerous; basidia 4-spored; spores hyaline,
even, subglobose, 314-4 y in diameter.

Fructifications 1-214 cm. in diameter in the fragmentary
specimens known to me.

On rough bark of a frondose species. Louisiana and Texas.
April and May. Rare.

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 231

P. asperipilata is a delicate, white, hypochnoid species covering
very rough decaying bark. It is noteworthy by the abundant,
needle-shaped, thin-walled cystidia with denticulate sides and by
the globose spores.

Specimens examined:

Louisiana: St. Martinville, A. B. Langlois, 44, comm. by Lloyd
Herb., 2395, and 1225, comm. by W. G. Farlow (in Mo. Bot.
Gard. Herb., 44067).

Texas: Houston, H. W. Ravenel, 265, type (in U. S. Dept. Agr.
Herb. and Burt Herb.).

16. P.albugo Burt, n. sp.

Type: in Burt Herb.

Fructifications longitudinally effused, filmy-pruinose, adnate,
whitish, pale smoke-gray in the herbarium, even, the margin in-
determinate, pruinose; in section 25-50 р. thick, not colored, with
the basidia and cystidia starting directly from the substratum or
with only very short, erect, intervening hyphae 214-3 џ in diam-
eter, thin-walled, collapsing; no gloeocystidia; cystidia not in-
crusted, 40-50 x 4144-6 u, protruding up to 40 u; spores white
in spore collections, even, 5-8 x 3-414 u, borne 4 to a basidium.

Fructifications 5-8 em. long, 114-3 em. wide.

Under side of deeaying frondose wood. Louisiana. December
and April.

P. albugo is a whitish, pruinose, filmy growth resembling in
aspect the young sterile mycelia which are sent in for determina-
tion in nearly all extensive series of specimens, but in this instance
Mr. Langlois took spore falls on glass from the specimens—a
highly commendable method of saving time, which is wasted
when sectional preparations are made of mere mycelia, and also
of keeping rubbish from preservation in the herbarium. Р. al-
bugo is related to P. detritica of France but has less membra-
naceous fructifications and more elongated spores.

Specimens examined:

Louisiana: St. Martinville, A. B. Langleis, ba, type, and dl.

17. P. albula Atkinson & Burt, n. sp.
Type: in Burt Herb.

[Vol. 12
232 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fructifications long-effused, adnate, thin, tender, small pieces
separable, white, becoming light buff when old and in the her-
barium, somewhat granular, becoming cracked into polygonal
masses 1-2 mm. in diameter, the margin thinning out; in section
70-200 » thick, not colored, composed of suberect, thin-walled,
branching hyphae about 3 и in diameter, occasionally nodose-
septate, not incrusted, sometimes slightly brownish near the
substratum; no gloeocystidia; cystidia not incrusted, 3 и in di-
ameter, tapering towards the apex, protruding 10-20 y, sometimes
very few and inconspicuous; spores hyaline, even, 4-6 х 214-
З р.
Fructifications 2-20 em. long, 1-2 cm. wide.

On bark of fallen decaying branches on the ground, of Alnus,
Acer, Tilia, Populus, and other frondose species. Canada to
Alabama and westward to Washington. July to February.
Frequent.

P. albula belongs near P. Sambuci on account of its white color,
somewhat granular hymenium, and minute cystidia which are not
incrusted and sometimes so few and inconspicuous that they may
possibly be overlooked, and the specimen referred to Corticium.
P. albula differs from P. Sambuci in not having the hyphae in-
crusted in a subhymenial zone and in having them sometimes
slightly brownish towards the substratum.

Specimens examined:

Exsiccati: Ellis, №. Am. Fungi, 409, under the name Corticium
calceu

m.

Canada: J. Macoun, 5, type; Beechwood Cemetery, other locality
not given, J. Macoun, 58; Ottawa, J. Macoun, 9.

Maine: Kittery Point, В. Thazxter & Е. A. Burt.

New Hampshire: Chocorua, W. G. Farlow, D (in Mo. Bot. Gard.
Herb., 56132).

Massachusetts: Sharon, A. P. D. Piguet, 137, comm. by Farlow
Herb. (in Mo. Bot. Gard. Herb., 59628); Wayland, A. B. Sey-
mour, T 8 (in Mo. Bot. Gard. Herb., 19550).

New York: Albany, H. D. House, 3 gatherings (in N. Y. State
Mus. Herb., and Mo. Bot. Gard. Herb., 57447, 59683, 59686) ;
Ithaca, several collections by G. F. Atkinson, H. L. Jackson,
C. O. Smith, and Van Hook, comm. by G. F. Atkinson, 8028,

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 233

8069, 8072, 8235, 14392, 14393; Karner, H. D. House (in N. Y.
State Mus. Herb., and Mo. Bot. Gard. Herb., 54373); New
York, Bronx Park, L. M. Underwood (in N. Y. Bot. Gard.
Herb., Mo. Bot. Gard. Herb., 61592, and Burt Herb.).

New Jersey: Newfield, J. B. Ellis, J 87, D 81, comm. by Farlow
Herb. (in Mo. Bot. Gard. Herb., 8266, 14689, 7458), and in
Ellis, N. Am. Fungi, 409.

Pennsylvania: Philadelphia, A. S. Rhoads, comm. by L. O.
Overholts, 2679 (in Mo. Bot. Gard. Herb., 5919).

Maryland: Takoma Park, C. L. Shear, 1271, 1272.

District of Columbia: Chive Chose, J. R. Weir, 372 (in Mo. Bot.
Gard. Herb., 17649).

Virginia: Chain Bridge, A. S. Rhoades, comm. by L. O. Overholts,
3969 (in Mo. Bot. Gard. Herb., 54986).

Florida: W. W. Calkins, comm. by U. S. Dept. Agr. Herb.

Alabama: Montgomery, R. P. Burke, 127, 157 (in Mo. Bot.
Gard. Herb., 5499, 44964).

Iowa: Woodbine, C. J. Humphrey & C. W. Edgerton, comm. by
C. J. Humphrey, 6511, 6546 (in Mo. Bot. Gard. Herb., 11063,
11276).

Washington: Bingen, W. N. Suksdorf, 894, 900.

18. P. Sambuci (Pers.) Burt, n. comb.

Corticium Sambuci Persoon, Roemer Neues Mag. Bot. 1: 111.
1794; Fries, Epier. 565. 1838; Hym. Eur. 660. 1874; Berkeley,
Outlines Brit. Fung. 276. 1860; Massee, Linn. Soc. Bot. Jour.
27:137. 1890; Wakefield, Brit. Myc. Soc. Trans. 4: 115. pl. 3,
f.1,2. 1913; Rea, Brit. Ваза. 677. 1922.—Thelephora Sambuci
Persoon, Syn. Fung. 581. 1801; Муе. Eur. 1: 152. 1822 (in
subgenus Corticium).—H ypochnus Sambuci (Pers.) Sace. Syll.
Fung. 6:656. 1888.—Thelephora sera Persoon, Syn. Fung. 580.
1801; Муе. Eur. 1: 151. 1822 (in subgenus Corticium).—Cor-
ticium serum (Регз.) Bresadola, І. В. Ассай. Agiati Atti III. 3:
112. 1897; Bourdot & Galzin, Soc. Мус. Fr. Bul. 27:246. 1911.

Fructifications effused, closely adnate, incrusting, not separ-
able, snow-white or chalk-white, sometimes becoming pale
cream-color in the herbarium, granular and pruinose, the margin
thinning out; in section 100-250 » thick, not colored, composed

[Vol. 12
234 ANNALS OF THE MISSOURI BOTANICAL GARDEN

of suberect, somewhat interwoven, thin-walled, incrusted hyphae

25-3 и thick, occasionally nodose-septate; по gloeocystidia;

cystidia not incrusted, tapering towards the apex, Зи in di-

ameter, protruding 10-30 u beyond the basidia; spores hyaline,

even, 4-514 x 3-4 y.

Fructifications 3-10 cm. long, 1-3 em. wide, often surrounding
small twigs.

On bark and wood of fallen Sambucus and other frondose
species. In Europe and throughout North America. Through-
out the year. Very common.

This species is very common, and has become so well known to
mycologists early in their work, under its original name Cor-
ticium Sambuci that there has been a reluctance, which I feel
also, to call it Peniophora Sambuci, which its structure really
requires. Its cystidia have been termed sterile basidia and cys-
tidioles, but they differ in no morphological respect from the
cystidia of other species of Peniophora. The species occurs in
especially fine condition on Sambucus, and it is well to use such
specimens as standards for comparison.

Specimens examined:

Exsiccati: Bartholomew, Fungi Col., 3135, 4715; Brinkmann,
Westfálische Pilze, 10; Cavara, Fungi Longobardiae, 63, 213;
Cooke, Fungi Brit., 408; Ell. & Ev., Fungi Col., 607, under the
name Corticium scutellare; Krieger, Fungi Sax., 523; Romell,
Fungi Scand., 35; Roumeguere, Fungi Gallici, 2911; Westen-
dorp, Herb. Crypt. Belge, 588.

Sweden: Stockholm, L. Romell, 129, 200, and in Romell, Fungi
Scand., 35. : 25

Germany: Saxony, Krieger, Fungi Sax., 523; Westphalia, in .
Brinkmann, Westfálische Pilze, 10.

Austria: Tirol, Innsbruck, V. Litschauer.

Italy: Rome, G. Bresadola; Brixia, A. Marozzi, and Papia, F.
Mae in Cavara, Fungi Longobardiae, 63 and 213 respec-
tively.

England: Forden, E. Vize, in Cooke, Fungi Brit., 408.

Belgium: Courtrai, in Westendorp, Herb. Crypt. Belge, 588.

France: Fautrey, comm. by Lloyd Herb., 4326, 4357; Bois de
Vincennes, N. Patouillard (in N. Y. Bot. Gard. Herb., and

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 235

Mo. Bot. Gard. Herb., 55906); Rouen, in Roumeguere, Fungi
Gallici, 2911.

Canada: J. Macoun, 21, 26; Lake Rosseau, E. T. & S. А. Н, arper,
754; Ottawa, J. Macoun, 370.

Maine: F. L. Harvey (in N. Y. State Mus. Herb., and Mo. Bot.
Gard. Herb., 55804).

New Hampshire: Chocorua, W. G. Farlow, 2, and C3 (in Mo. Bot.
Gard. Herb., 43893).

Vermont: Grand View Mountain, E. A. Burt; Middlebury, E. A.
Burt, 4 gatherings; Weybridge, E. A. Burt.

Massachusetts: East Wareham, С. L. Shear, 2903 (in Mo. Bot.
Gard. Herb., 15448).

New York: Clarksville, C. H. Peck, comm. by N. Y. State Mus.
Herb. (in Mo. Bot. Gard. Herb., 18281); Constableville, C. H.
Peck, comm. by N. Y. State Mus. Herb., T4 (in Mo. Bot.
Gard. Herb., 54571); Ithaca, G. F. Atkinson, 942, 4567, 8047,
22960; Lyndonville, C. E. Fairman, comm. by N. Y. State
Mus. Herb. (in Mo. Bot. Gard. Herb., 57512); Marcellus,
L. M. Underwood, 62 (in N. Y. Bot. Gard. Herb., and Mo. Bot.
Gard. Herb., 61595); New York, W. A. Murrill (in N. Y. Bot.
Gard. Herb., and Mo. Bot. Gard. Herb., 61595).

New Jersey: Newfield, J. B. Ellis, in Ell. & Ev., Fungi Col., 607.

Pennsylvania: Trexlertown, W. Herbst, 11.

Louisiana: Baton Rouge, Edgerton & Humphrey, comm. by C. J.
Humphrey, 5662; St. Martinville, A. B. Langlois, dd, 37,
comm. by Lloyd Herb., 2384, and 1944, comm. by W. G. Far-
low (in Mo. Bot. Gard. Herb., 44065).

Michigan: Ann Arbor, C. H. Kauffman (in Mo. Bot. Gard. Herb.,
5584); Gogebie County, E. A. Bessey, 372 (in Mo. Bot. Gard.
Herb., 56636).

Missouri: Creve Coeur, L. O. Overholts (in Mo. Bot. Gard. Herb.,
63704) ; St. Louis, E. A. Burt (in Mo. Bot. Gard. Herb., 63466,
58335).

Kansas: Stockton, E. Bartholomew, 5815, 8206 (in Mo. Bot.
Gard. Herb., 16709, 62175), and in Bartholomew, Fungi Col.,
4715; Rooks County, E. Bartholomew, 2045 (in Mo. Bot. Gard.
Herb., 4827).

Washington: Bainbridge Island, E. Bartholomew, in Bartholo-

[Vol. 12
236 ANNALS OF THE MISSOURI BOTANICAL GARDEN

mew, Fungi Col., 3135; Olympia, C. J. Humphrey, 6311;
Sedro-Woolley, C. J. Humphrey, 7464.

California: Claremont, J. M. Johnston, comm. by L. О. Over-
holts, 3645 (in Mo. Bot. Gard. Herb., 54699); Santa Catalina
Island, L. W. Nuttall, 522b (in Mo. Bot. Gard. Herb., 57622).

Mexico: Guernavaca, W. A. & E. L. Murrill, 891, comm. by N. Y.
Bot. Gard. Herb. (in Mo. Bot. Gard. Herb., 54548); Orizaba,
W. A. & E. L. Murrill, 756, comm. by М. Y. Bot. Gard. Herb.
(in Mo. Bot. Gard. Herb., 54644).

19. P. Thujae Burt, n. sp.

Type: in Burt Herb.

Fruetifications effused, thin, closely adnate, white, sometimes
becoming cartridge-buff in the herbarium, the margin thinning
out; in section 40-150» thick, not colored, with the hyphae
loosely arranged, suberect, interwoven, 2-3» in diameter,
nodose-septate, thin-walled, becoming incrusted in a subhymenial
zone; no gloeocystidia; cystidia hair-like, not incrusted, Зи in
diameter at the base, tapering upward and sometimes some-
what capitate at apex, protruding 20-30 » beyond the basidia;
spores white in a spore collection, 4-5 x 3 y.

Fructifications 2-6 cm. long, 1-2 em. wide, on trunks of
Thuja, more rarely on Juniperus and Pinus Strobus. Canada to
Massachusetts, and westward to Missouri. July to October.
Occasional. |

P. Thujae may be recognized by its thin, white fructifications
on white cedar, with microscopic structure as stated. It differs
from Peniophora Sambuci in having thinner fructifications, not
becoming granular and pruinose, cystidia numerous, and in oc-
currence on a coniferous substratum.

Specimens examined:

Canada: J. Macoun 62; St. Lawrence Valley, J. Macoun, 68, 80;
Ottawa, J. Macoun (in N. Y. State Mus. Herb., and Mo. Bot.
Gard. Herb., 55813), and 45, and J. M. Macoun (in N. Y.
State Mus. Herb., and Mo. Bot. Gard. Herb., 56078) ; Quebec,
Hull, J. Macoun, 173.

Vermont: Middlebury, E. A. Burt, 3 gatherings, one of which is

e type.

1925] :
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 237

Massachusetts: Magnolia, W. б. Farlow, f; Newton, W. G.

Farlow.

New York: Ithaca, G. F. Atkinson, 14416; North River, C. H.
Peck, comm. by N. Y. State Mus. Herb., T5 (in Mo. Bot.
Gard. Herb., 54569).

Missouri: St. Louis, E. A. Burt.

20. P. montana Burt, n. sp.

Type: in Burt Herb.

Fructifications effused, thin, adnate, tender, whitish to ivory-
yellow, widely cracked in drying and showing the loose subiculum
on the sides of the crevices, the margin thinning out, somewhat
floccose; in section 200-225 u thick, not colored, composed of
loosely interwoven, thin-walled, hyaline hyphae 4-5 їп di-
ameter, not incrusted, not nodose-septate, of irregular outline;
no gloeocystidia; cystidia hair-like, not incrusted, conical, taper-
ing to a sharp apex, 6-9 » in diameter at the base, protruding
up to 40 u; spores hyaline, even, cylindric, slightly curved, 12-
14 X 4-5 y.

Fragmentary fructification 4 cm. long, 1 cm. wide.

On badly decayed coniferous wood at an altitude of 10,000 ft.
Colorado. July. Rare.

P. montana is noteworthy by having spores as large as those of
P. mutata, but the fructifications are thinner and more tender
than those of P. mutata and occur on coniferous wood and have
no gloeocystidia.

Specimens examined:

Colorado: Ouray, C. L. Shear, 1188, type.

21. P.terricola Burt, n. sp.

Type: in Burt Herb.

Fructifications effused, thin, closely adnate, somewhat mem-
branaceous, white, not waxy, the margin indeterminate, thinning
out; in section 100-200 » thick, not colored, composed of sub-
erect, branching hyphae 3-6 џ in diameter, incrusted, densely
interwoven and with more or less sand intermixed; no gloeocys-
tidia; cystidia not incrusted, cylindric, 4—6 џ in diameter, pro-
{тийш 20-50 и beyond the База; spores hyaline, even, 4-6
х 38-4 y.

[Vol. 12
238 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fructifications received in fragments but probably about 1-3
em. in diameter.

On ground in mixed woods. New York and Louisiana. April
and June.

The fructifications of P. terricola contain so much of the sand
from the earth substratum that it is difficult to secure sections
or to distinguish the fructification proper from its vegetative
mycelium. The occurrence in small white patches on the ground,
and the characters of spores and cystidia may enable recognition
of this species which is probably common.

Specimens examined:

New York: Ithaca, G. F. Atkinson, 22658, 22659, type.
Louisiana: St. Martinville, A. B. Langlois, bq.

22. P. magnahypha Burt, n. sp.

Type: in Mo. Bot. Gard. Herb. and Farlow Herb.

Fructifications interruptedly effused, thin, adnate, between pale
drab-gray and pale vinaceous-fawn, contracting in drying into
small, more or less completely separated masses, not waxy, the
margin thinning out; in section 150-180 џ thick, not colored,
composed of erect hyphae 9-10 » in diameter which start from
the substratum at points 30-100 y apart, branch repeatedly into
branches of smaller diameter, are sparingly granule-incrusted,
and terminate in large clusters of basidia and one or a few cys-
tidia forming a hymenium; no gloeocystidia; cystidia not in-
crusted, septate, 9 » in diameter, protruding up to 60 p beyond
the basidia; basidia with 4 sterigmata; spores hyaline, even,
9-10 x 6 y.

Fructifications up to 4 em. long, 2 cm. wide.

On decaying wood of a frondose species. Florida. Autumn.

While preliminary inspection of P. magnahypha with a lens
does not promise more than any one of the great number of little-
differentiated, perplexing, whitish resupinate species difficult
to identify yet doing an important work in splitting up complex
organic compounds, nevertheless its structural characters are
unique. The combination of coarse, scattered, trunk-like, erect
hyphae with the main trunk hypha or some of its principal
branches protruding through and beyond the flat-topped cluster

1925]
URT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 239

of basidia as a transversely septate cystidium should lead to the
ready recognition of this species when sectional preparations are
studied.
Specimens examined:
Florida: Cocoanut Grove, В. Тражег, 57, type (in Mo. Bot.
Gard. Herb., 43947).

23. P. exilis Burt, n. sp.

Type: in Mo. Bot. Gard. Herb.

Fructifications longitudinally effused, very thin, closely adnate,
pale olive-buff, even, somewhat velutinous, the margin thinning
out, indeterminate; in section 30-60 и thick, not colored, com-
posed of erect, bushy-branched hyphae 3 y in diameter, ascending
from the substratum, soon terminating in basidia and cystidia,
not nodose-septate, with very little, if any, incrustation; no
gloeocystidia; cystidia hair-like, irregular, flexuous, 30 x 414-
5 и, protruding up to 20 u, few and scattered; basidia simple,
with 4 short sterigmata; spores hyaline, even, 415-5 X 214-3 y.

Fructifications 1-6 em. long, about 1 cm. wide.

On bark of decaying branches of frondose species in moist
virgin forest. Mexico. January.

The fructifications of P. exilis occur as a thin, downy, gray
coating on very rotten branches 1-2 cm. in diameter. The pale
olive-buff color should be helpful in separating this species from
the great number more white in color.

Specimens examined:

Mexico: Guernavaca, W. A. & E. L. Murrill, 385, comm. by
N. Y. Bot. Gard. Herb. (in Mo. Bot. Gard. Herb., 54467);
Orizaba, W. A. & E. L. Murrill, 757, type, and 780, comm. by
N. Y. Bot. Gard. Herb. (in Mo. Bot. Gard. Herb., 54619,
54620).

24. P. livida Fries in herb. under Corticium, n. sp.

Peniophora serialis in part of v. Hóhnel & Litschauer, Bourdot
& Galzin, and Rea.—Not Corticium seriale Fries of Fries Herb.—
Not Xerocarpus Cacao Karsten, Hedwigia 29: 271. 1890.

Type: in Herb. Fries, determined by E. Fries as Corticium
lividum.

[Vol. 12
240 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fructifications longitudinally effused, closely adnate, waxy-
soft, variable in color, pale olive-gray and pale olive-buff to fawn-
color in the herbarium, glabrous, even, not cracked usually,
rarely with a few fissures from contraction in drying, the margin

inning out; in section 75-500 џ thick, not colored, composed of
densely interwoven, rather erect hyphae about 3 и in diameter,
indistinct, with the wall gelatinously modified; no gloeocystidia;

cystidia not incrusted, tapering to a sharp apex, 314-6 y in di-

ameter, protruding up to 40 џ beyond the basidia; spores hyaline,

even, 4-5 x 115-2 y.

Fructifieations 3-12 cm. long, 1-3 em. wide.

Generally on old, decaying, coniferous wood, rarely on frondose
wood. Europe, Louisiana, and British Columbia. Throughout
the year.

P. livida may be best recognized by its close resemblance in
aspect to even specimens of common Corticium lividum Pers.,
from which the presence of cystidia separate it. Р. livida is one
of the 3 species which European mycologists, following Bresadola,
have been inclined to regard as sufficiently meeting the original
description of Corticium seriale that one could ignore the fact
that the species concerned do not agree in structure with one
another, nor with any of the specimens in Kew Herbarium or
Fries Herbarium, determined by Elias Fries as Corticium seriale.
With regard to the appliability of the original description of
Corticium seriale, it emphasizes rimose and testaceous fructi-
fications which are not characters of P. livida. It might solve
the problem of Corticium seriale Fr. to search in Sweden for a
true Corticium which is testaceous and rimose and could be com-
pared with the specimen in Kew Herbarium determined by Fries
—something more like Corticium Cacao Karst, which has the
hymenium somewhat deteriorated in my specimen so that I can-
not be quite positive as to its genus from this specimen alone
but seems to me to be a true Corticium.

Specimens examined:

Sweden: E. Fries, type, the thinner and paler of the specimens in
Herb. E. Fries, determined by E. Fries as Corticium lividum;
L. Romell, 108, 109; Femsjé, L. Romell, 410; Stockholm,
L. Romell, 198, 326, 345, 362.

1925]
T—THE THELEPHORACEAE OF NORTH AMERICA. XIV 241

Austria: Tirol, Innsbruck, V. Litschauer, 3 specimens under the
name Р. serialis.

Louisiana: Bogalusa, C. J. Humphrey, 5547.

British Columbia: Revelstoke, C. W. Dodge, 1639 (in Mo. Bot.
Gard. Herb., 58784); Sidney, J. Macoun, 9 (in Mo. Bot.
Gard. Herb., 5768); Victoria, J. Macoun, 541 (in Mo. Bot.
Gard. Herb., 63728).

25. P. phyllophila Massee, Linn. Soc. Bot. Jour. 25: 150.
1889; басе. Syll. Fung. 9: 238. 1891; Rea, Brit. Basid., 697.
1922.

Asterostromella epiphylla v. Hóhnel & Litschauer, К. Akad.
Wiss. Wien Sitzungsber. 116: 773. text f. 3. 1907.

Type: type distribution in Ravenel, Fungi Am., 457, under the
name Corticium epiphyllum.

Fructification broadly effused, thin, closely adnate, not at all
separable, whitish, becoming olive-buff in the herbarium, the
margin thinning out; in section 40-80 u thick, not colored, com-
posed of suberect, interwoven, branching, thin-walled hyphae
2 in diameter, not incrusted, not nodose-septate, bearing clus-
ters of basidia and branching paraphyses; also present occasional,
tapering cystidia not incrusted, 30-45 x 4-10 џ, usually im-
mersed, occasionally protruding up to 32 џ beyond the basidia;
paraphyses colorless, branching at the hymenial surface into an
antler-shaped form with very slender prongs; spores published
by v. Höhn. & Lits. as 10-22 x 112-3 y.

Fructifications up to 5 cm. in diameter.

On fallen frondose leaves in South Carolina and on fallen de-
caying frondose limbs in Florida and Central America.

P. phyllophila is apparently a tropical species occurring more
frequently on epidermis of small fallen twigs and ranging north-
ward to South Carolina. I have studied specimens of the type
distribution in the copies of Ravenel, Fungi Americana, of
Farlow Herbarium, Missouri Botanical Garden Herbarium,
United States Department of Agriculture Herbarium, and Burt
Herbarium, and find these specimens to be the same in structure
and all showing the distinctive antler-shaped paraphyses em-
phasized by v. Hóhnel & Litschauer, and also tapering, non-

[Vol. 12
242 ANNALS OF THE MISSOURI BOTANICAL GARDEN

inerusted cystidia which are presumably what Massee really
saw. І see no reason for displacing the specific name given by
Massee for the later combination proposed by v. Hóhnel &
Litschauer. The basidia are so young that I found none bearing
sterigmata nor spores and only twice slender spores 12-15 X
3 p.
Specimens examined:
Exsiccati: Ravenel, Fungi Am., 457, type distribution, under the
name Corticium epiphyllum.
South Carolina: Aiken, H. W. Ravenel, in Ravenel, Fungi Am.,
457.
Florida: W. W. Calkins.
Central America: Panama Chagres, F. L. Stevens, 1300 (in Mo.
Bot. Gard. Herb., 63521).

26. P. piliseta Burt, n. sp.

Type: in N. Y. Bot. Gard. Herb., Mo. Bot. Gard. Herb., and
Burt Herb.

Fructifications longitudinale effused, thin, somewhat mem-
branaceous, tender, small pieces separable when moistened,
whitish cream-color in the herbarium, not cracked, not shining,
the margin thinning out, with the hyphae interwoven; in section
100-120 u thick, not colored, composed of ordinary, interwoven,
thin-walled hyphae about Зи in diameter, not incrusted nor
nodose-septate, and of a system of hyaline tissue about 1 in
diameter, not taking stain, branching like the coarser tissue of
Corticium investiens and with its delicate antler-shaped branches
barely visible in the hymenial surface; no gloeocystidia; cystidia
not incrusted, cylindric, obtuse, 414-6 y. in diameter, protruding
30-45 u, confined to the surface of the hymenium; spores hyaline,
even, cylindric, biguttulate, 9-11 x 4-414 в, copious.

оек 715 em. long, broken off at one end, 10-15 mm.
wide.

On a very rotten, small, frondose limb about 1 em. in diameter.
Porto Rieo. June.

P. piliseta is noteworthy by having in addition to an ordinary
hyphal system in its fructification an additional system, inter-
mixed with че first, of delicate, branching organs not taking

1925]
URT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 243

stain, such as is more distinctly visible, because coarser, in P.
phyllophila, Hypochnus pallescens, Corticium investiens and
Grandinia granulosa, and whose peripheral branches are more or
less visible in the surface of the hymenium as antler-shaped
paraphyses. Р. mexicana has coarser hyphae and more hypoch-
noid surface.
Specimens examined:
Porto Rico: Martin Pifia, Rio Piedras, J. R. Johnston, 971 a, type
(in N. Y. Bot. Gard. Herb., Mo. Bot. Gard. Herb., 63243, and
Burt Herb.).

27. P. mexicana Burt, n. sp.

Type: in N. Y. Bot. Gard. Herb. and Mo. Bot. Gard. Herb.

Fructifications longitudinally effused, adnate, dry, hypochnoid,
thin, cream color in the herbarium, even, velutinous under a
lens, not cracked, the margin thinning out; in section 140 yu
thick, not colored, composed of even-walled, rather rigid, loosely
arranged, branching hyphae 5-7 и in diameter, not incrusted,
not nodose-septate, which ascend obliquely from the substratum
and bear a dense hymenium containing numerous cystidia and
branching, filiform paraphyses (or perhaps conidiophores); no
gloeocystidia; cystidia minutely incrusted or rough, tapering,
60-100 x 5-9 в, protruding 40-60 u; spores (perhaps conidia)
hyaline, even, 6-714 X 4-5 y, copious.

Fructifications 4 em. long and broken off at both ends, 6 mm.
wide.

In depressed places on very rotten frondose wood. Mexico.
January.

The dry, cream color or buff fructifications of hypochnoid
texture, very coarse hyphae, large cystidia, and branching par-
aphyses or conidiophores in the surface of the hymenium are
characters which should make this species recognizable, although
my inability to demonstrate basidia convinces me that the type
is in a conidial stage somewhat comparable with that of Corticium
roseum.

Specimens examined:

Mexico: Orizaba, Nuevo, altitude 3600 m., W. A. & E. L. Mur-

rill, 773, type (in Mo. Bot. Gard. Herb., 54633).

[Vol. 12
244 ANNALS OF THE MISSOURI BOTANICAL GARDEN

28. Р. ludoviciana Burt, n. sp.

Type: in Burt Herb., and Farlow Herb. probably.

Fructifications effused, adnate, very thin, buff-yellow, dark-
ening to cinnamon-buff in the herbarium, hymenium subvelu-
tinous, not waxy, not cracking, the margin thinning out, paler;
in section 40-75 thick, not colored, composed of suberect,
branching, granule-incrusted, hyaline hyphae 3-4 y in diameter :
no gloeocystidia; cystidia hyaline, not incrusted, protruding
18-25 » beyond the basidia; spores hyaline, even, 4-5 x 214-
316 в, somewhat flattened on one side.

Fructifications 1-214 em. long, 14-114 ст. broad, becoming
confluent.

On rotting decorticated wood of frondose species. Louisiana
and Michigan. August and April. Rare.

P. ludoviciana closely resembles in aspect P. flammea and,
like the latter, is not separable from the substratum and gives no
noteworthy color changes when the sections are treated with
potassium hydrate solution. Prolonged search has failed to find
any immersed cystidia. P. sulphurina has larger, cracked fruc-
tifications with shining hymenium and yellow subiculum.

Specimens examined:

Louisiana: St. Martinville, A. B. Langlois, 1919, type, comm. by

W. G. Farlow.

Michigan: Vermilion, A. H. W. Povah, 369 (in Mo. Bot. Gard.

Herb., 13921).

29. P. fusca Burt, n. sp.

Type: in Mo. Bot. Gard. Herb.

Fructifications effused, closely adnate, very thin, drying ecru-
drab to drab, velvety, even, the margin not determinate, thinning
out; in structure 35—45 y thick, not colored, composed of loosely
arranged, suberect, hyaline hyphae more or less incrusted, 314-
4 р in diameter under the incrustation, not nodose-septate; no
gloeocystidia nor conducting organs; cystidia hair-like, not in-
crusted, 7-124 in diameter at the base, protruding 40-125 u
beyond the basidia; basidia with 4 sterigmata; spores copious,
hyaline, even, 6-7 x 3% y.

Fructifications 2-6 em. long, 1-2 em. wide, becoming larger by
confluence,

1925]
URT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 245

On very rotten, decortieated and probably frondose wood.
Alabama. June to October. Only 2 gatherings known.

P. fusca is a thin species of mucedinous aspect, like P. longi-
spora but well characterized by its drab color, large cystidia, and
moderately large spores. Р. cinerea is sometimes of the same
color but is less mould-like when viewed with a lens and with
quite different structure and microscopic characters.

Specimens examined:

Alabama: Montgomery County, R. P. Burke, 508, type, and 836

(in Mo. Bot. Gard. Herb., 57301 and 63125 respectively).

30. P. gilvidula Bresadola, Mycologia 17: 70. 1925.

Type: in Weir Herb.

Fructifications broadly effused, closely adnate, waxy, pinkish
buff in the herbarium, here and there somewhat cracked, pruinose,
the margin thinning out; in section 150-250 u thick, not colored,
2-layered, the layer next the substratum 75-150 и thick, com-
posed of densely arranged hyphae about 4-5 y in diameter, not
incrusted, which are longitudinally interwoven in the type,
hymenial layer 75-100 и thick, composed of densely arranged,
erect, coarse tissue; no gloeocystidia; cystidia not incrusted,
6-8 џ in diameter, protrading 30-60 u beyond the basidia, not
numerous, confined to the hymenium; basidia with 4 sterigmata;
spores white in the mass, even, 5-6 х 214-314 u, copious.

Fructifications 8-15 em. long, 3-5 em. wide.

On wood of log of Pinus ponderosa. Montana. September.

P. gilvidula has no especially distinctive character. The oc-
currence on Pinus ponderosa wood, buff color, thick hymenial
layer, coarse hyphae, and small spores constitute the group of
distinguishing characters. I have included under P. gilvidula a
specimen from the same place which has the layer next to the
substratum composed of erect hyphae.

Specimens examined:

Montana: Evaro, J. R. Weir, 23305, type (in Weir Herb.) and

426 (in Mo. Bot. Gard. Herb.).

31. Р. zonata Burt, n. sp.
Type: in Mo. Bot. Gard. Herb.

[Vol. 12
246 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fructifieations widely effused, closely adnate, thick, layered
or zonate within, avellaneous, pruinose, contracting in drying and
cracking into more or less connected masses about 1 mm. in
diameter, the margin thinning out; in section 700 y. thick, prob-
ably stratose but perhaps with merely a hymenium of several
(4 in the type) layers or zones, not colored, composed of densely
arranged hyphae about 214-3 in diameter, with somewhat
gelatinously modified and indistinct, not sharply defined as
tramal, and hymenial layers; no gloeocystidia; cystidia not
incrusted, З и in diameter at the base, tapering to a sharp apex,
protruding up to 30 џ, very numerous in the surface of the
hymenium; spores hyaline, even, curved, 414 x 2% y, copious.

Portion of fructification 7 cm. long, 4 em. wide, broken off at
one end and on sides.

On decayed coniferous wood. Oregon. March. :

The cystidia are so small and so very numerous in the hymenial
surface and the season when collected—March—so early in the
year that it is possible that this species is a stratose Corticium
just starting a new outer stratum on its fructification, but I do
not recognize it as a Corticium at present known to me. No
matter what the genus may prove to be, the thick, somewhat
liver-colored fructifications of layered or stratose structure, and
notably cracked, should always make this species easy to recog-
nize.

Specimens examined:

Oregon: Corvallis, S. M. Zeller, 2252, type (in Mo. Bot. Gard.

Herb., 63030).

32. P.laminata Burt, n. sp.

Type: in Burt. Herb.

Fructifications broadly effused, thin, adnate, not separable,
cream-buff to warm buff, pubescent, somewhat tubercular, at
length cracking into small masses 2-3 to a mm., the margin
thinning out, fibrillose; in section 75-140 и, rarely 200 џ, thick,
not colored, becoming stratose, 1-6 strata, each composed of a
supporting layer of loosely arranged, erect, hyaline hyphae 3-
315 р in diameter, thin-walled, collapsing, not incrusted, and of a
compact hymenial layer; no gloeocystidia; cystidia not incrusted,

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 247

hair-like, cylindric, obtuse, 3-314 » in diameter, protruding up
to 30 р, beyond the basidia; basidia 4-spored ; spores hyaline, even,
415 x 3-314 в, copious.
Fructifications 2-8 cm. in diameter.
On bark and wood of fallen decaying trunk of Pinus Strobus.
Vermont. December. Rare.

P. laminata is so suggestive in color and general aspect of the
very common Corticium investiens that it is possible P. laminata
has been passed by as a thin, young specimen of C. investiens,
but the structure of these two species is quite different. The
color of P. laminata does not fade in the herbarium; my gathering
of nearly thirty years ago still has the color originally noted.

Specimens examine
Vermont: Middlebury, Е. А. Burt, уре.

33. Р. guttulifera (Karst.) Saec. Syll. Fung. 9: 240. 1891;
Bourdot & Galzin, Soc. Муе. Fr. Bul. 28: 400. 1913.

Gloeocystidium guttuliferum Karsten, Finska Vet. Soc. Bidrag
Natur och Folk 48: 430. 1889.

Type: a portion in Burt Herb.

Fructifications broadly effused, closely adnate, thin, becoming
light buff to pinkish buff and chamois-colored in the herbarium,
more or less studded with minute, hard, globular masses of res-
inous color which are visible under a lens but dissolve and disap-
pear in aqueous mounts, the margin indeterminate, thinning out;
in section 50-160 » thick, not colored, with the hyphae erect,
branching, 3-5 и in diameter, not incrusted; no gloeocystidia;
cystidia heavily incrusted, often obtuse, 40-90 х 10-15 u, pro-
truding up to 60 u; spores white in a spore collection, even, de-
pressed on one side, 7-10 X 3-4% y.

Fructifications 2-5 cm. long, 1-214 cm. wide.

On decaying wood of Populus, Betula, Acer, and undetermined
frondose species. In Europe, and from Canada to Louisiana
and westward to Oregon. May to January. Rare.

The type specimen of P. guttulifera differs from P. pubera in
having no gloeocystidia whatever and in bearing on its surface
minute, globular, shining masses of such aspect as occur on tips
of the granules in Odontia sudans. Such masses are also borne

[Vol. 12
248 ANNALS OF THE MISSOURI BOTANICAL GARDEN

on specimens from France communicated by Bourdot, and they

are stated to be borne on the cystidia—this in addition to the

usual incrustation of these cystidia. Since the resinous-colored

masses disappear in the liquids to which they are subjected in

sectioning and making aqueous preparations for microscopic

study, I am inclined to regard the presence of these masses as

perhaps due to weather conditions prevalent when the specimens

bearing them were collected—a helpful, confirmatory specific

feature when present, but not a necessary morphological character

of P. guitulifera. Hence I have included under P. guttulifera,

specimens which have spores 7-10 x 3-414 y, lack gloeocystidia,

and have the aspect of P. pubera.
Specimens examined:

Finland: Mustiala, P. A. Karsten, type of Gloeocystidium gut-
tuliferum, under the label Gloeocystis gultulifera.

Sweden: Femsjó, E. A. Burt; Góteberg, L. Romell, 295.

France: Allier, St. Priest, H. Bourdot, 6656, 8458.

Canada: Ottawa, J. Macoun, 180 a.

Maine: Kittery Point, В. Thazter & E. A. Burt, 2 gatherings.

New Hampshire: Shelburne, W. G. Farlow, 3.

Vermont: Middlebury, E. A. Burt.

New Jersey: Newfield, J. B. Ellis (in N. Y. Bot. Gard. Herb.,
and Mo. Bot. Gard. Herb., 61398).

Alabama: Montgomery County, R. P. Burke, 212, 478 (in Mo.
Bot. Gard. Herb., 57083, 57295).

Louisiana: A. B. Langlois, 256, comm. by U.S. Dept. Agr. Herb. -

Ohio: Lancaster, W. A. Kellerman, 168, comm. by U. S. Dept.
Agr. Herb.

British Columbia: Agassiz, J. Macoun, 129.

Oregon: Corvallis, W. A. M urrill, 940, comm. by N. Y. Bot.
Gard. Herb., 55715.

34. P. flavido-alba Cooke, Grevillea 8: 21. pl. 125, f. 14.
1879; Басс. Syll. Fung. 6: 644. 1888; Massee, Linn. Soc. Bot.
Jour. 25:151. 1889.

: in Kew Herb.
: Fructifications broadly effused, thin, closely adnate, cracking
in drying, becoming cartridge-buff to pinkish buff in the her-

1925]
URT— —THE THELEPHORACEAE OF NORTH AMERICA. XIV 249

barium, setulose with the large cystidia, the margin indeterminate,

thinning out; in section 75-250 y. thick, not colored, composed of

densely interwoven, hyaline hyphae about Зи in diameter, not
inerusted, and of very numerous large cystidia, many of which
are often tilted in all directions; no gloeocystidia; cystidia heavily

inerusted, cylindric-fusiform to conical, sharp-pointed, 60-120

X 12-18 y, numerous in all regions to the substratum, protrud-

ing up to 50 џ beyond the basidia; spores hyaline, even, white in a

spore collection, 444-6 x 2-86 u.

Fructifications 5-15 em. or more long, 2-5 em. wide.

On bark of decaying limbs of Carya, Liguidambar, Myrica,
Quercus, Salix, Vitis, and other frondose species. South Carolina
to Louisiana, West Virginia and Ohio to Arkansas, and in the
West Indies. July to April. Common.

P. flavido-alba resembles in aspect P. pubera with which it was
eonfused by v. Hóhnel & Litschauer in their study of specimens
distributed by Ravenel and by Ellis in their exsiccati, but differs
sharply from P. pubera in absence of gloeocystidia and in having
smaller spores. Its spores are smaller than those of P. guttulifera;
it lacks layered structure, and the cystidia are much larger than
in either P. Вазепей ог P. Rowmeguerit. There may be ob-
served in sectional preparations a curious tilting of many cys-
tidia, some towards the right and some towards the left while
most are erect and the tilting is at varying angles, being occasion-
ally quite parallel with the substratum. Such tilting is unique
among the species of Peniophora known to me and is best shown
by the immersed cystidia in sections from the thicker fructi-
fications. The type specimen in Kew Herbarium is on the same
substratum, Myrica, as the specimens distributed in Ravenel,
Fungi Am., 226, and impressed me as probably being from the
same gathering.

Specimens examined:

Exsiccati: Bartholomew, Fungi Col., 4741; Ellis, N. Am. Fungi,
1209; Ell. & Ev., N. Am. Fungi, 3412; Ravenel, Fungi Am.,
226.

South Carolina: P. H. Rolfs, 1622, 1625.

Georgia: Atlanta, E. Bartholomew, 5677, 5689 (in Mo. Bot.
Gard. Herb., 44253); Darien, H. W. Ravenel, 2529, type (in

[Vol. 12
250 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Kew Herb.) and specimens in Ravenel, Fungi Am., 226, and
, N. Am. Fungi, 1209.

Florida: W. W. Calkins, comm. by W. G. Farlow (in Mo. Bot.
Gard. Herb., 44634), and 628, comm. by W. G. Farlow (in Mo.
Bot. Gard. Нев. 44641, 44254); New Smyrna, С. б. Lloyd,
2128; Tallahassee, Е. Bartholomew, 5722 (in Mo. Bot. Gard.
Herb., 44256).

Alabama: Auburn, F. S. Earle & C. F. Baker, 2217 (in N. Y.
Bot. Gard. Herb., and Mo. Bot. Gard. Herb., 61344); Mont-
gomery County, В. P. Burke, 68, 147, 164, 169, 237, 444, 463,
465, 475, 667 (in Mo. Bot. Gard. Herb., 18395, 7552, 44963,
44959, 57105, 57271, 57284, 57286, 57293, 63089).

Louisiana: Abita Springs, A. B. Langlois, 2684; Baton Rouge,
Edgerton & Humphrey, comm. by C. J. Humphrey, 5665; New
Orleans, E. Bartholomew, in Bartholomew, Fungi Col., 4741,
and 5765 (in Mo. Bot. Gard. Herb., 44265); A. B. Langlois,
460 (in N. Y. Bot. Gard. Herb., and Mo. Bot. Gard. Herb.,
61476); St. Martinville, A. B. Langlois, 2680, comm. by
Lloyd Herb., 3529, and 1954, 2679, aq. bt, cm, and cn.

West Virginia: Ellis Coll., 48 (in N. Y. Bot. Gard. Herb., and
Mo. Bot. Gard. Herb., 61395).

Ohio: Cincinnati, C. G. Lloyd, 4515, 4526, 4806.

Kentucky: Crittenden, C. G. Lloyd, 3115; Mammoth Cave, C. G.
Lloyd, 1602, and in Ell. & Ev., N. Am. Fungi, 3412.

Arkansas: Bigflat, W. H. Long, 19894 (in Mo. Bot. Gard. Herb.,
6387).

Cuba: San Antonio de los Baños, Havana Province, Earle &
Murrill, 88, comm. by N. Y. Bot. Gard. Herb.; Pinar del Rio
Province, Earle & Murrill, 241, comm. by N. Y. Bot. Gard.
Herb.; Santiago de las Vegas, H. Hasselbring (in N. Y. Bot.
Gard. "Herb. and Mo. Bot. Gard. Herb., 61468).

Porto Rico: Rio Piedras, J. A. Stevenson, 8366, 5582, 6068 (in
Mo. Bot. Gard. Herb., 7574, 6944, 54685).

Jamaica: Hall’s Delight, P. S. Earle, 184, comm. by N. Y. Bot.
Gard. Herb.

35. P. vernicosa Ellis & Everhart in herb., n.
Type: in N. Y. Bot. Gard. Herb., Mo. Bot. dad Herb., and
Burt Herb.

1925]
URT—THE THELEPHORACEAE OF NORTH AMERICA, XIV 251

Fructifieations long and broadly effused, very thin, closely
adnate, pinkish buff in the herbarium, even, somewhat puberu-
lent and setulose under a lens, not cracked, the margin thinning
out, indeterminate; in section 30-45 y. thick, not colored, com-
posed of densely ео, hyaline hyphae about 3 u in di-
ameter, indistinct; no gloeocystidia; cystidia incrusted, fusiform,
40-50 X 10-15 u, protruding up to 50 и beyond the База;
spores hyaline, even, 4-5 x 3-314 y.

Fructifications 10-12 cm. long, 2-3 em. wide.

On dead pieces of Celtis. Florida and Louisiana. August and
March.

The 3 gatherings under the name P. vernicosa in Ellis Col-
lection of New York Botanical Garden and duplicates of these
communicated to me directly by Langlois seem to be thin forms
of 3 species, 2 of which are well known. The type of P. vernicosa
shows the location of the fructification by the pinkish buff color .
of the area covered, somewhat varnish-like effect produced, and
cystidia visible under a lens. There is the bare possibility that
P. vernicosa may be demonstrated to be the very early stage of
P. flavido-alba but my knowledge of the latter does not at present
warrant such a conclusion.

Specimens examined:

Florida: Cutler Hammock, W. A. Murrill, 86, comm. by N. Y.

Bot. Gard. Herb. (in Mo. Bot. Gard. Herb., 62082).
Louisiana: St. Martinville, A. B. Langlois, 1965, type (in N. Y.

Bot. Gard. Herb., Mo. Bot. Gard. Herb., 63726, and Burt

Herb.).

36. P.texana Burt, n. sp.

Type: in Mo. Bot. Gard. Herb.

Fructifications long and broadly effused, adnate, thin, even,
not shining, drying between cream-buff and pinkish buff and
cracking transversely, the margin indeterminate, thinning out;
in section about 100» thick, not colored, with the hyphae in-
distinct, interwoven, 3-85 и in diameter, not incrusted; cystidia
inerusted, conical, often tilted, not colored, 45-55 x 10-12 y,
protruding beyond the basidia up to 45 џ; no gloeocystidia nor
conducting organs; spores copious, hyaline, even, 414-6 x 3-
416 y.

[Vol. 12
252 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fructifications up to 25 em. long, 5 cm. broad.

On bark of Juniperus sabinoides. Texas. October. Only the
type collection known.

Although occurring on bark of Juniperus, P. texana is not at
all related to P. laevigata and seems rather to belong in the group
of species of which P. flavido-alba is best known. The occur-
rence on Juniperus, the large expanse of the fructifications, and
large cystidia and spores should afford recognition of P. ехала.

Specimens examined:

Texas: Austin, W. H. Long, 21070, type (in Mo. Bot. Gard.

Herb., 55134).

37. P.flammea Burt, n. sp.

Type: in Burt Herb. and N. Y. Bot. Gard. Herb.

Fructifications effused, adnate, very thin, olive-ocher to deep
chrome, fading to clay color in the herbarium, hymenium often
with some granules, the margin thinning out, paler; in section
50-90 y. thick, not colored and with no color changes by potas-
sium hydrate solution, with hyphae 3 y in diameter, interwoven
next to the substratum but suberect, branching and granule-
incrusted towards the hymenium; no gloeocystidia; wholly im-
mersed cystidia incrusted, 15-60 x 5—10 и, few and scattered;
hair-like cystidia not incrusted, 3-5 и in diameter at base, pro-
truding 20-30 u beyond the База, are scattered in the surface
of the hymenium; spores hyaline, even, 314-5 x 114-214 y.

Fructifications 1-10 em. long, 5 mm.-214 em. broad.

On rotting wood and bark of frondose species and on under
side of rotting leaves of Sabal. Florida, Alabama, Texas, Cuba,
and Bermuda. March to June. Probably rare.

P. flammea has the intense yellow color of Corticium chrysocreas
and Odontia Wrightii but, unlike these species, its sections do not
become vinaceous and then bleach when treated with potassium
hydrate solution and the structural details of the sections are
quite different also. Peniophora sulphurina is yellow and has
small spores, but the fructification of P. flammea is as closely
adnate to, and inseparable from, the substratum as that of P.
cinerea.

Specimens examined:

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 253

Florida: Tarpon Springs, W. A. Murrill, 216, comm. by N. Y.
Bot. Gard. Herb. (in Mo. Bot. Gard. Herb., 62121).

Alabama: Montgomery, R. P. Burke, 3, 158 (in Mo. Bot. Gard.
Herb., 17431, 44962).

Texas: Austin, W. H. Long, 524.

Cuba: C. G. Lloyd, 421 (in Mo. Bot. Gard. Herb., 55172); El
Yunque Mt., Baracoa, L. M. Underwood & F. S. Earle, 1215,
type, comm. by N. Y. Bot. Gard. Herb.

Bermuda: Paget Swamp, H. H. Whetzel, Abe (in Mo. Bot. Gard.
Herb., 58905).

38. P. isabellina Burt, n. sp.

Type: in Burt Herb.

Fructifications longitudinally effused, very thin, closely adnate,
not at all separable, between light pinkish cinnamon and avel-
laneous, not shining, becoming somewhat minutely cracked, the
margin thinning out; in section 50-75 u thick, not colored, com-
posed of innumerable cystidia and densely arranged hyphae
212-3 и in diameter, indistinct; no gloeocystidia; cystidia in-
crusted, 30 X 6 и, protruding up to 12 и, fusoid, usually starting
from the substratum; spores 6 X З y. present but so few found
that they may not belong.

Fructification 8 em. long and broken off at both ends, 1 cm.
broad.

On dead canes of blackberry (Rubus), and perhaps on other
frondose wood. Virginia and Alabama. June to September.

P. isabellina is as closely adnate as P. cinerea and P. versicolor,
from both of which it differs in not being colored in section. The
occurrence of the type on blackberry stems may be helpful in
recognizing this species, but several other species also occur on
blackberry stems. 'The specimen from Alabama, referred to
P. isabellina, is proebuy specifically distinct.

Specimens examin
Virginia: Woodstock, C. L. Shear, 1191, type.

Alabama: Montgomery County, R. P. Burke, 62 (in Mo. Bot.

Gard. Herb., 18207).

39. P. coccineo-fulva (Schw.) Burt, n. comb.
Phlebia coccineo-fulva Schweinitz, Am. Phil. Soc. Trans. М. S.

[Vol. 12
254 ANNALS OF THE MISSOURI BOTANICAL GARDEN

4: 165. 1832; Saec. Syll. Fung. 11: 112. 1895.—Corticium
rhodellum Peck, М. Y. State Mus. Rept. 42: 122. 1889.—Peni-
ophora rhodella (Peck) Sace. Syll. Fung. 9: 239. 1891.— Penio-
phora Karstenit Massee, Linn. Soc. Bot. Jour. 25: 153. 1889.
—Corticium calotrichum Karsten, Rev. Мус. 10: 73. 1888; Soc.
pro Fauna et Fl. Fenn. Meddel. 16: 21. 1888; Icones Hym.
Fenn. 3: 7. pl. 4, f. 71. 1891; Sace. Syll. Fung. 6: 617. 1888;
9: 232. 1891.—Peniophora rhodochroa Bresadola, Mycologia 17:
70. 1925.—Peniophora leprosa Bourdot & Galzin, Soc. Myc.
Fr. Bul. 28: 394. 1913.

Type: in Schweinitz Herb. and Farlow Herb.

Fructifications effused, adnate, becoming russet to Natal
brown in the herbarium, sometimes cracked, the margin paler;
in section typically vinaceous russet but sometimes not colored,
150—400 и thick, 2-layered, the layer next to substratum 100-
300 u thick, composed of loosely interwoven, thin-walled hyphae
4-8 u in diameter, with many rough-walled or incrusted, the
hymenial layer very dense, typically colored, bearing the cys-
tidia; cystidia hyaline or slightly colored, incrusted, 40-80 X
10-14 y, protruding up to 50 u; spores hyaline, even, 4-5 x 2-
215 у.

Fructifications 4-10 cm. long, 2 cm. broad. |

On rotting wood and bark of Juglans, Quercus, and other fron-
dose species, rarely on conifers. Canada to Alabama and west-
ward to British Columbia and California, and in Mexico; occurs
in Europe also. July to December. Frequent.

P. coccineo-fulva has been confused with P. velutina, from which
it differs when best developed, in more intense color, the vinaceous
subhymenial layer often showing this color on edges of cracks in
the fructification, and in the incrusted hyphae. Paler specimens
which are not otherwise distinguishable from P. velutina I have
now referred to P. coccineo-fulva when they have the large, in-
crusted hyphae of the latter, for the European concept of P.
velutina, as shown by specimens under this name in Kew Herba-
rium and communicated to me by Bourdot, Bresadola, Romell,
and Litschauer, has the hyphae not incrusted, with the exception
of additional specimens from Bresadola and Romell which they
distinguished as different from Р. velutina by labelling as “ Peni-

1925]
URT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 255

ophora velutina Fr. f. pallidior," and which I cite below as P. coc-

cineo-fulva. These European specimens have exactly the same

structure as the authentic specimen of Corticium calotrichum
sent to me by Karsten, who noted the large rough hyphae in the
description in Icones Hym. Fenn. 3:7, but the hyphae are really
granule-incrusted.

Specimens examined:

Exsiccati: Ell. & Ev., N. Am. Fungi, 2019, under the name
Peniophora velutina; Ell. & Ev., Fungi Col, 707, under the
name Peniophora velutina; Rabenhorst, Fungi Eur., 3231,
under the name Corticium alneum, the type distribution of
Peniophora Karstenii.

Finland: Mustiala, P. A. Karsten, authentic specimen of Cor-
ticium calotrichum, and in Rabenhorst, Fungi Eur., 3231.

Sweden: Femsjó, L. Romell, 421; on Fagus, Hangnen, Femsjó,

| E. А. Burt.

Germany: Brinkmann, comm. by Bresadola as Peniophora velutina
Fr. f. pallidior.

France: Aveyron, A. Galzin, 26563, comm. by H. Bourdot,
32878, authentic specimen of Peniophora leprosa.

Canada: Hull, Quebec, J. Macoun, 197; Lambeth, Ontario,
J. Dearness, D 172b (in Mo. Bot. Gard. Herb., 5482); Granton,
J. Dearness, 966 (in Mo. Bot. Gard. Herb., 22582); Ottawa,
J. Macoun 197, 291, and J. M. Macoun, 230 (in N. Y. State
Mus. Herb., and Mo. Bot. Gard. Herb., 55756, 55920, 56081
respectively).

New Brunswick: Campobello, W. G. Farlow, 2 (in part).

Maine: Boarstone Mountain, Piscataquis County, W. A. Mur-
rill, 2404 (in N. Y. Bot. Gard. Herb., and Mo. Bot. Gard.
Herb., 61354).

New Hampshire: Chocorua, W. G. Farlow, 43 (in Mo. Bot. Gard.
Herb., 43972); North Conway, A. S. Rhoads, 10 (in Burt
Herb., and Mo. Bot. Gard. Herb., 56979).

Vermont: Middlebury, E. A. Burt, 2 gatherings.

New York: Albany, H. D. House (in N. Y. State Mus. Herb.,
and Mo. Bot. Gard. Herb., 59703); Alcove, C. L. Shear, 1309;
Floodwood, C. H. Peck би N. Y. State Mus. Herb., Burt
Herb., and Mo. Bot. Gard. Herb., 55986); Hudson Falls,

[Vol. 12
256 ANNALS OF THE MISSOURI BOTANICAL GARDEN

S. H. Burnham, 36 (in Mo. Bot. Gard. Herb., 54457); Ithaca
Flats, G. F. Atkinson, 3090; Karner, H. D. House (in N. Y.
State Mus. Herb., and Mo. Bot. Gard. Herb., 54368) ; Lyndon-
ville, C. E. Fairman, type of Corticium rhodellum Gn М. X.
State Mus. НегЬ.); Westport, C. Н. Peck (in N. Y. State Mus.
Herb., and Mo. Bot. Gard. Herb., 55968).

New Jersey: Belleplain, C. L. Shear, 1242; Newfield, J. B. Ellis,
in Ell. & Ev., N. Am. Fungi, 2019, and Fungi Col., 707, and
(in Burt Herb., N. Y. Bot. Gard. Herb., and Mo. Bot. Gard.
Herb., 57337, 63455).

Maryland: Takoma Park, C. L. Shear, 1335.

Pennsylvania: Nazareth, Schweinitz, type of Phlebia coccineo-fulva
(in Herb. Schweinitz and Farlow Herb.).

Alabama: Auburn, comm. by Alabama Biological Survey; Mont-
gomery, Е. P. Burke, 72, 188, 635 (in Mo. Bot. Gard. Herb.,
17582, 57068, 63072).

Ohio: Cincinnati, C. G. Lloyd, 2808.

Michigan: Ann Arbor, C. H. Kauffman, 35 (in Mo. Bot. Gard.
Herb., 20025); New Richmond, C. H. Kauffman, 26 (in Mo.
Bot. Gard. Herb., 16386); Vermilion, A. Н. W. Povah, 5 (in
Mo. Bot. Gard. Herb., 9225).

Wisconsin: Lake Geneva, E. T. & S. A. Harper, 834, 961;
Madison, C. J. Humphrey & M. C. Jensen, 681 (in Mo. Bot.
Gard. Herb., 10275).

Colorado: Pike’s Peak, G. G. Н edgcock, comm. by C. J. Humphrey,
2554 (in Mo. Bot. Gard. Herb., 9782).

Idaho: Priest River, J. R. Weir, 131 (in Mo. Bot. Gard. Herb.,
15762), and 16809, type of Peniophora rhodochroa (in Weir
Herb. and Mo. Bot. Gard. Herb., 63690).

British Columbia: Kootenai Mts. near Salmo, J. В. Weir, 635
(in Mo. Bot. Gard. Herb., 21995).

California: Big Wash Cafion, Santa Catalina Island, L. W. Nut-
tall, 889, comm. by Field Mus. Nat. Hist. Herb. (in Mo. Bot.
Gard. Herb., 57650). |

Arizona: Coronado Nat. Forest, G. G. Hedgcock, comm. by C. J.
Humphrey, 2547 (in Mo. Bot. Gard. Herb., 9906).

Mexico: Jalapa, W. A. & E. L. Murrill, 144, comm. by N. Y.
Bot. Gard. Herb. (in Mo. Bot. Gard. Herb., 6962).

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 257

40. P.laevis (Fr.) Burt in R. Fries, R. Sci. Soc. Gothoburgens
Actis IV. 3: [36]. 1900; in Peck, N. Y. State Mus. Bul. 54: 954.
1902; v. Hóhnel & Litschauer, K. Akad. Wiss. Wien Sitzungsber.
115: 1550. 1906; Bourdot & Galzin, Soc. Мус. Fr. Bul. 28:
398. 1913; Rea, Brit. Basid. 692. 1922.

Thelephora laevis Fries, Elenchus Fung. 1: 206. 1828. Not
T. laevis Persoon.—Corticium laeve Fries, Epicr. 560. 1838;
Hym. Eur. 649. 1874.—Kneiffia laevis (Fries) Bresadola, Ann.
Myc. 1: 99. 1903.

Type: authentic specimen in Kew Herb.

Fructifications effused, membranaceous, adnate, separable
from the substratum when moistened, drying light pinkish cin-
namon to buff-pink and ochraceous buff, the margin radiately
fibrillose; in section not colored, 300—400 y. thick, with the hyphae
3-414. in diameter, not colored, granule-incrusted, densely
erowded together and running parallel with the substratum and
then ascending obliquely into the hymenium; cystidia incrusted
or not incrusted, 40-60 х 415-9 џ, protruding up to 30 y, con-
fined to the hymenial layer; spores white in à spore collection,
even, 414-6 x 214-3 u.

Fructifications 2-10 em. long, 2-4 cm. broad.

On bark of frondose species. Europe, New Brunswick to
Texas and westward to Washington and Oregon, in Cuba and
in Island of Guam and in Japan. July to October. Not com-
mon.

Peniophora laevis is one of the species which Karsten under-
stood as Corticium radiosum and sent under this name to Fries,
as shown by the specimens in Herb. Fries determined by Karsten,
and preserved by Fries without comment. Bresadola collected
the species occasionally and communicated to me duplicates under
the herbarium name Peniophora albo-gilvida. The above speci-
mens agree in aspect with the authentie specimen of Corticium
laeve from Fries in Kew Herb. and also agree with it in the details
of microscopic structure including incrusted hyphae, smaller than
those of Peniophora coccineo-fulva and more compactly and more
longitudinally arranged than those of P. sanguinea. P. affinis
does not have its hyphae at all incrusted.

Specimens examined:

[Vol. 12
258 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Sweden: authentic specimen (in Kew Herb.); on Betula, L.
Romell, 122; Gottenburg, L. Romell, 120; Stockholm, L. Ro-
mell, 358.

Finland: P. Karsten, 32 (in Fries Herb., under the name Corticium
radiosum) ; Mustiala, P. Karsten, under the name C. radiosum,
comm. by Bresadola, and also on Alnus under the name
Peniophora velutina.

Russian Poland: Eichler, 107, comm. by Bresadola.

France: Allier, St. Priest, H. Bourdot, 8981, under the name P.
Evchleriana.

Italy: Trient, Alps Mts., Bresadola, two specimens.

New Brunswick: Campobello, W. G. Farlow, 2.

New Hampshire: Chocorua, W. G. Farlow, 12 (in Burt Herb.)
and C 35, C 38, 40 (in Mo. Bot. Gard. Herb., 48963, 43967,
43971).

Vermont: Middlebury, E. A. Burt, three gatherings.

Massachusetts: Magnolia, W. G. Farlow; Williamstown, W. G.
Farlow, 9.

New York: East Galway, E. A. Burt; East Schaghticoke, C. H.
Peck (in N. Y. State Mus. Herb., and Mo. Bot. Gard. Herb.,
95758); East Schodack, C. H. Peck, 12; Hague, C. H. Peck, 2;
Ithaca, G. F. Atkinson, 4598; North Greenbush, H. D. House,
14.234 (in N. Y. State Mus. Herb., and Mo. Bot. Gard. Herb.,
44733); Snyder, C. H. Peck, 16.

New Jersey: Newfield, J. B. Ellis, 2020, comm. by W. G. Farlow,
22 (in Mo. Bot. Gard. Herb., 7943).

Virginia: Crabbottom, W. A. Murrill, 169, 259 (in N. Y. Bot.
Gard. Herb., 61557, 61568).

Alabama: Montgomery County, R. P. Burke, 129, 218, 813 (in
Mo. Bot. Gard. Herb., 11034, 57085, 63115).

Texas: Gonzales, C. L. Shear, 1231.

Kentucky: Crittenden, C. G. Lloyd, 10113 (in Mo. Bot. Gard.
Herb., 58689).

Ohio?: locality not stated, C. G. Lloyd, 4195.

Michigan: New Richmond, C. H. Kauffman, 47 (in Mo. Bot.
Gard. Herb., 3259).

Wisconsin: Blue Mounds, E. T. & S. A. Harper, 943; Lake
Geneva, E. T. & S. A. Harper, 836; Palmyra, A. O. Stucki, 53.

1925]
URT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 259

Missouri: Perryville, R. A. Studhalter & L. O. Overholts, 2706
(in Mo. Bot. Gard. Herb., 44290).

Nebraska: Lincoln, C. L. Shear, 540.

Idaho: Priest River, J. R. Weir, 608 (in Mo. Bot. Gard. Herb.,
63196).

British Columbia: Sidney, J. Macoun, 10 (in Mo. Bot. Gard.
Herb., 5728).

Washington: Bingen, W. N. Suksdorf, 764; Arlington, C. J.
Humphrey, 7610.

Oregon: Eugene, C. J. Humphrey, e um S. M. Zeller,
1985 (in Mo. Bot. Gard. Herb.,

Cuba: Ceballos, C. J. Неру. ri

Island of Guam: Edwards, comm. by J. R. Weir, 10765 (in Mo.
Bot. Gard. Herb., 56238).

Japan: Mt. Mili; Prov. Awaji, A. Yasuda, 62 (in Mo. Bot.
Gard. Herb., 56138).

41. P. subiculosa Burt, n. sp.

Type: in Mo. Bot. Gard. Herb. and N. Y. Bot. Gard. Herb.

Fructifications effused, somewhat membranaceous, tender,
small pieces separable when moist, with the hymenium drying
cartridge-buff, pulverulent, here and there cracked and showing
the whitish subiculum which is pale chamois-colored next to the
substratum and connected with chamois-colored marginal my-
celial strands ог cords; in section 400—500 џ thick, not distinctly
colored, with the hyphae loosely interwoven, hyaline, 4 и in
diameter, not nodose-septate, granule-incrusted in all regions
with large crystalline granules; cystidia heavily incrusted, 20-60
X 9 u, protruding up to 15 в, confined to the hymenium; spores
hyaline, even, 3-314 x 21% u, borne 4 to a basidium.

Fructifications 2-3 cm. long, 1 em. broad.

On humus of frondose wood. Mexico. December. Only one
collection known.

P. subiculosa is related to P. Burtii but differs from it in having
larger and incrusted cystidia and all hyphae heavily incrusted.

Specimens examined:
Mexico: Guernavaca, W. A. & E. L. Murrill, 396, type (in N. Y.

Bot. Gard. Herb., and Mo. Bot. Gard. Herb., 54550).

[Vol. 12
260 ANNALS OF THE MISSOURI BOTANICAL GARDEN

42. P. septocystidia Burt, n. sp.

Type: in N. Y. Bot. Gard. Herb., Mo. Bot. Gard. Herb., and
Burt Herb.

Fructifieations effused, the small patches becoming more or
less confluent, membranaceous, separable, between warm buff
and cinnamon-buff in the herbarium, somewhat tubercular
through conforming to the irregularities of the substratum, the
margin byssoid, with some mycelial strands; in section 250—400
thick, not colored, 2-layered, the layer next to the substratum
much the thicker, composed of very loosely interwoven, in-
crusted hyphae 4-5 in diameter under the incrustation, not
nodose-septate, the hymenial layer dense, 35-45 м thick; no
gloeocystidia; cystidia incrusted with a few, large, somewhat
colored granules, transversely septate, 5 и in diameter under the
incrustation, protruding 30-35 и, scattered along surface of
hymenium; spores hyaline, even, cylindric, curved, 5-7 x 214-
3

[he
Fructifications 5 mm.—214 cm. in diameter after confluence.
On decaying bark and humus. West Indies. January.

P. septocystidia is somewhat related to P. sanguinea, P. Burti,
and P. subiculosa, but is of different color, with very coarse
hyphae and noteworthy cystidia. |

Specimens examined:

Jamaica: Troy and Tyre, Cockpit Country, W. A. Murrill &

W. Harris, 860, type (in Burt Herb., N. Y. Bot. Gard. Herb.,

and Mo. Bot. Gard. Herb., 61490).

43. Р. canadensis Burt, n. sp.

Type: in Burt Herb. i

Fructification effused, adnate, dry, hypochnoid, small pieces
separable when moistened, cream-color in the herbarium, even,
tomentose under a lens, not shining, the margin thinning out, of
finely interwoven hyphae; in section 300-350 y thick, not colored,
stratose, each of the two strata composed of loosely arranged,
erect, branching, nodose-septate, somewhat incrusted hyphae
4-6 u in diameter, which are slightly colored near the substratum
and hyaline elsewhere, and of a more compact hymenial layer
containing cystidia; no gloeocystidia; cystidia incrusted, cylin-

1925]
URT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 261

dric, 50-90 х 6-9 u, protruding up to 45 в, very numerous in
the hymenium; basidiospores hyaline, even, 7-8 X 4-5 y, copious,
four to а basidium; other spherical spores 814—4 џ in diameter
are present in addition to immersed basidiospores in the buried
hymenium.

Fructification 21% cm. long, 2 ст. wide, broken off at both ends.

On wood of coniferous log and bark of Frazinus. Canada and
New York. September and October.

The type of P. canadensis somewhat resembles P. pubera in
aspect but has texture more suggestive of Coniophora byssoidea.
Such aspect, together with the coarse hyphae, large spores, and
numerous large cystidia should fix the species. Unfortunately,
the type consists of but a single piece of the dimensions stated,
which was present in a packet of Corticium bombycinum. The
New York gathering consists of a group of very small fructi-
fications only one stratum thick.

Specimens examined:

Canada: locality not given, J. Macoun, 60 (in part), type.
New York: Ithaea, G. F. Atkinson, Cornell Univ. Herb., 8282.

P. cremea Bresadola, Fungi Trid. 2: 63. pl. 178, f. 2.
1898; Sacc. Syll. Fung. 16: 195. 1902; Bourdot & Galzin, Soc.
Mye. Fr. Bul. 28: 396. 1913; Wakefield, Brit. Myc. Soc.
Trans. 5: 131. 1914; Rea, Brit. Ваза. 691. 1922.

Kneiffia cremea Bresadola, Ann. Мус. 1: 100. 1903.—An
Corticium Eichlerianum Bresadola, Ann. Мус. 1: 95. 1903?

Type: in Bresadola Herb. p authentic specimens in
Burt Herb.

Fructifications broadly effused, membranaceous, separable,
white or cream-color to ochraceous buff and darkening somewhat
in the herbarium, sometimes cracking when dry and showing the
white subiculum, the margin white and cobwebby; in section
100-300 y, rarely 500 и, thick, not colored, composed of a broad
layer next to the substratum of thick-walled, hyaline, erect hyphae
414-6 ų in diameter, branching at а wide angle, sometimes dichot-
omously, more or less granule-incrusted towards the hymenial
layer; hymenial layer dense, bearing protruding cystidia even-
walled or incrusted about the apex and containing also immersed,

[Vol. 12
262 ANNALS OF THE MISSOURI BOTANICAL GARDEN

incrusted cystidia when thickened; protruding cystidia cylindrie
or tapering towards the apex, 5-10 u in diameter at the base,
protruding up to 60 & beyond the basidia; immersed cystidia
40-60 Х 9-10 u; no gloeocystidia; spores white in a spore col-
lection, even, 5-8 x 215-3L5 y.

Fructifications 4-15 em. long, 2-4 em. wide.

On bark-covered and decorticated branches of frondose species
on the ground. In Europe, Canada to Louisiana, and westward
to the Pacific States and in Japan and in Natal, Africa. May to
January. Infrequent but widely distributed.

P. cremea is readily recognizable among the species of the
northern United States and Canada by its thick, white or creamy
fructifications which have small spores, lack gloeocystidia, and
are 2-layered with the thick under layer composed of coarse,
loosely arranged, erect hyphae branching at an angle of towards
60° and often dichotomously. These hyphae and their arrange-
ment are distinctive. P. mutata has the same aspect and color
but differs by much longer spores and the presence of gloeo-
cystidia. P. velutina has its hyphae ascending obliquely to the
hymenial layer.

Specimens examined:

Sweden: L. Romell, 196; Femsjé, L. Romell, 218.

Germany: Westphalia, Lengerich, Brinkmann, 341, determined
and communicated by Bresadola.

Russian Poland: Eichler, determined and communicated by

Bresadola.

Austria: Tirol, Gries, V. Litschauer; Innsbruck, V. Litschauer;

Stiermark, V. Litschauer.

France: Aveyron, M. Galzin, 13292, comm. by H. Bourdot, 20856.

England: Doncaster, E. M. Wakefield (in Mo. Bot. Gard. Herb.,
57126).

Canada: J. Macoun, 24.

New Hampshire: Chocorua, E. A. Burt.

Vermont: Bristol, E. A. Burt, two gatherings; Middlebury, E. A.

Burt, two gatherings.

Massachusetts: Magnolia, W. G. Farlow, a; Sharon, A. P. D.

Piguet, comm. by W. G. Farlow.

New York: East Galway, E. A. Burt, three gatherings; Bergen

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 263

Swamp, Genesee County, H. D. House (in N. Y. State Mus.
Herb., and Mo. Bot. Gard. Herb., 57473).

New Jersey: Newfield, J. B. Ellis, 68 (in N. Y. Bot. Gard. Herb.,
and Mo. Bot. Gard. Herb., 63425).

District of Columbia: W. A. Murrill, 1496 (in N. Y. Bot. Gard.
Herb., and Mo. Bot. Gard. Herb., 63453, 63465).

Alabama: Montgomery County, В. P. Burke, 800 (in Mo. Bot.
Gard. Herb., 63108).

Louisiana: St. Martinville, A. B. Langlois, k, 1886, 1963, 2631
(in Burt Herb., N. Y. Bot. Gard. Herb., and Mo. Bot. Gard.
Herb., 63456, 63503).

Michigan: Gogebie County, E. A. Bessey, 321 (in Mo. Bot.
Gard. Herb., 56543).

Montana: Rexford, E. E. Hubert, comm. by J. R. Weir (in Weir
Herb., and Mo. Bot. Gard. Herb., 63246).

Idaho: Coolin, J. R. Weir, 11499, 11575 (in Mo. Bot. Gard.
Herb., 63261, 63303), and an unnumbered specimen (in Weir
Herb., and Mo. Bot. Gard. Herb., 63247); Priest River, J. В.
Weir, 609 (in Mo. Bot. Gard. Herb., 63197).

Manitoba: Swan River, G. R. Bisby, 1049 (in Mo. Bot. Gard.
Herb., 59036); Winnipeg, G. R. Bisby, 1117 (in Mo. Bot. Gard.
Herb., 59040).

British Columbia: Sidney, J. Macoun, 23, 28, 73, 82, 104, 834 (in
Mo. Bot. Gard. Herb., 5757, 55335, 5758, 5759, 55337, 55334).

Washington: Bingen, W. N. Suksdorf, 867; Chehalis, C. J.
Humphrey, 6260; Everson, C. J. Humphrey, 7453; Kalama,
C. J. Humphrey, 6205

Oregon: Corvallis, S. M. Zeller, 1867 (in Mo. Bot. Gard. Herb.,
56871); Eugene, C. J. Humphrey, 6088.

California: Palo Alto, W. A. Murrill, 1173, comm. by М. Y. Bot.
Gard. Herb. (in Mo. Bot. Gard. Herb., 55706); Santa Catalina
Island, Grand Canyon, L. W. Nuttall, 1060, comm. by Field
Mus. Herb. (in Mo. Bot. Gard. Herb., 58883).

Japan: Sendai, A. Yasuda, 46 (in Mo. Bot. Gard. Herb., 56160);
Mt. Mikuma, Prov. Awaji, A. Yasuda, 53 (in Mo. Bot. Gard.
Herb., 56161).

Africa: Natal, Durban, P. A. van der Bijl, 612 (in Mo. Bot. Gard.
Herb., 59377).

[Vol. 12
264 ANNALS OF THE MISSOURI BOTANICAL GARDEN

45. P. velutina (DC) Cooke, Grevillea 8: 21. pl. 125, f. 15.
1879; Басс. Syll. Fung. 6: 644. 1888; Massee, Linn. Soc. Bot.
Jour. 25: 152. 1889; Bourdot & Galzin, Soc. Myc. Fr. Bul. 28:
398. 1913; Rea, Brit. Basid. 692. 1922.

Thelephora velutina De Candolle, Fl. Fr. 6: 33. 1815; Fries,
Elenchus Fung. 1: 203. 1828.—Corticium velutinum (DC)
Fries, Epicr. 561. 1838; Hym. Eur. 650. 1874.—Kneiffia velutina
(DC) Bresadola, Ann. Мус. 1: 100. 1903.

Fructifications broadly effused, membranaceous, separable,
becoming vinaceous buff to fawn color in the herbarium, minutely
_ velvety, the margin whitish, often extended in branching mycelial
strands; in section not colored, 250—500 u thick, composed of
loosely interwoven hyphae up to 5-8 in diameter, not in-
crusted, only very rarely nodose-septate; cystidia incrusted,
40-100 X 8-15 u, wholly immersed in the hymenial tissue or
protruding up to 50 p; spores white in spore falls, even, 414-514 X
215-3 y.

Fructifications 3-20 cm. long, 2-15 em. broad.

On decaying limbs and logs of such frondose species as Fagus,
Quercus, Castanea, Populus, etc., more rarely on coniferous wood.
Throughout Canada and the United States and in Europe. May
to December. Frequent. :

P. velutina may be recognized by its large and rather thick
fructifications of pinkish or vinaceous color when dry, separable
from the substratum when moistened, by frequent presence of
marginal mycelial strands, and by the coarse, non-incrusted
hyphae—often up to 8» in diameter—present in sectional prep-
arations near the substratum.

Specimens examined:

Sweden: L. Romell, 121, 133; Stockholm, L. Romell, 137.
Poland: Eichler, from Bresadola.

Austria: Tirol, V. Litschauer.

France: Cormatin, F. Guillemin, 10, in part; St. Priest, Allier,

Н. Bourdot, 20859.

Canada: J. Macoun, 231 , comm. by W. G. Farlow (in Mo. Bot.
Gard. Herb., 14763); Ontario, Casselman, J. Macoun, 366.
New Hampshire: Chocorua, W. G. Farlow, 71 (in Mo. Bot.

Gard. Herb., 43973).

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 265

Vermont: Ripton, E. A. Burt.

Massachusetts: R. J. Blair, comm. by Г. О. Overholts, 3812 b
(in Mo. Bot. Gard. Herb., 54994).

New York: Alcove, C. L. Shear, 1198; East Galway, E. A. Burt,
two gatherings: Floodwood, C. H. Peck (in N. Y. State Mus.
Herb., and Mo. Bot. Gard. Herb., 55967), E. A. Burt; Karner,
H. D. House (in N. Y. State Mus. Herb., and Mo. Bot. Gard.
Herb., 54393).

New Jersey: Alpine, P. Wilson, 29 (in Mo. Bot. Gard. Herb.,
54748).

Pennsylvania: State College, L. O. Overholts, 3326 (in Mo. Bot.
Gard. Herb., 9533).

Alabama: Montgomery County, R. P. Burke, 419 (in Mo. Bot.
Gard. Herb., 57259).

Tennessee: Elkmont, C. H. Kauffman, 73 (in Mo. Bot. Gard.
Herb., 54330).

Michigan: New Richmond, C. H. Kauffman, 54 (in Mo. Bot.
Gard. Herb., 11996); Seney, C. J. Humphrey, 1596 (in Mo.
Bot. Gard. Hub. 17541).

Wisconsin: Madina, C. J. Humphrey, 2156 бп Mo. Bot. Gard.
Herb., 6729).

Illinois: Аай C. J. Humphrey, 2044 (in Mo. Bot. Gard. Herb.,
21525).

Montana: Bernice, E. E. Hubert, comm. by J. R. Weir (in Mo.
Bot. Gard. Herb., 63250); Yellowstone, F. S. Wolpert, comm.
by J. R. Weir, 3934 (in Mo. Bot. Gard. Herb., 55179).

Colorado: Pike's Peak, G. G. Hedgcock, comm. by C. J. Humphrey,
2543 (in Mo. Bot. Gard. Herb., 20783).

Idaho: Priest River, J. R. Weir, 618 (in Mo. Bot. Gard. Herb.,
63200).

British Columbia: Kootenai Mts., near Salmo, J. R. Weir, 512
(in Mo. Bot. Gard. Herb., 3772); Sidney, J. Macoun, 34, 42
(in Mo. Bot. Gard. Herb., 55341, 55345).

Washington: Bingen, W. N. Suksdorf, 703.

Oregon: Grant's Pass, J. R. Weir, 8687 (in Mo. Bot. Gard.
Herb., 63199).

New Maxion: Tyom Experiment Station, W. H. Long, 21898 (in
Mo. Bot. Gard. Herb., 55121).

[Vol. 12
266 ANNALS OF THE MISSOURI BOTANICAL GARDEN

46. P. affinis Burt, n. sp.

Name without description in Peck, N. Y. State Mus. Bul. 54:
954. 1902.

Type: in Burt Herb.

Fructifications broadly effused, membranaceous, adnate, sepa-
rable when moistened, drying light buff to pinkish buff and light
pinkish einnamon, often cracked and showing the paler subiculum
in the erevices, the margin paler, radiately fibrillose; in section
not colored, 300-500 u thick, with the hyphae hyaline, 3-5 u in
diameter, not at all incrusted, arranged densely and longitudi-
nally in a broad layer along the substratum and then ascending
obliquely into the hymenial layer; cystidia incrusted or not in-
crusted, 30-60 х 5-8 u, protruding up to 30 u, occurring in the
hymenial layer only; spores white in a spore collection, even,
416-6 х 21$-3 y.

Fructifications 3-20 сто. long, 2-4 em. broad.

On bark and decorticated logs and limbs of frondose species.
Canada to New York and westward to Oregon, and also in
Europe. July to October. Common.

P. affinis is related in aspect to P. laevis and has hyphae of the
same diameter and arrangement as those of the latter species but
not at all inerusted. "The fructifications of P. affinis are usually
thicker than those of P. laevis, less adnate to the substratum,
paler and more cracked. Pale specimens of P. sanguinea crack
into somewhat similar areas but show a somewhat colored, floc-
cose subiculum in the fissures. The hyphae of P. affinis are of
smaller diameter than those of P. velutina.

Specimens examined:

Exsiccati: Reliq. Farlowianae, 343, under the name Peniophora
laevis

Sweden: L. Romell, 123, 124, both under the name P. velutina.

Austria: Tirol, V. Litschauer, under the name Р. laevis.

France: Allier, H. Bourdot, 8579, under the name P. laevis.

Canada: J. Macoun, 76, comm. by N. Y. State Mus. Herb. (in
Mo. Bot. Gard. Herb., 57510).

Quebec: Hull, J. M. acoun, 220.

Ontario: Jefferson, G. H. Graham, comm. by Univ. Toronto
Herb., 674 (in Mo. Bot. Gard. Herb., 44924).

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 207

Maine: Kittery Point, В. Thaxter & E. A. Burt.

New Hampshire: Chocorua, W. G. Farlow, 35 and two unnum-
bered specimens in Burt Herb., Reliq. Farlowianae, 343, and
C 36, 41, 69 (in Mo. Bot. Gard. Herb., 43964, 43969, 43970 re-
spectively), E. A. Burt, three gatherings; North Conway, L. O.
Overholis, 5106 (in Mo. Bot. Gard. Herb., 56356).

Vermont: Middlebury, Е. A. Burt, type and another gathering.

Massachusetts: North Scituate, W. G. Farlow.

New York: Albany, H. D. House (in N. Y. State Mus. Herb. and
Mo. Bot. Gard. Herb., 14835); East Galway, E. A. Burt, six
gatherings; Jamesville, L. M. Underwood (in N. Y. Bot. Gard.
Herb., and Mo. Bot. Gard. Herb., 63419); Karner, H. D. House
(in N. Y. State Mus. Herb., and Mo. Bot. Gard. Herb., 54348,
54352, 54371); Oneida, H. D. House (in N. Y. State Mus.
Herb., and Mo. Bot. Gard. Herb., 59681); Snyder, C. H. Peck
(in N. Y. State Mus. Herb., and Mo. Bot. Gard. Herb., 55757);
Syracuse, L. M. Underwood, 116 (in N. Y. Bot. Gard. Herb.,
and Mo. Bot. Gard. Herb., 61485); West Fort Ann, S. H.
Burnham, 12 (in Mo. Bot. Gard. Herb., 44002).

Tennessee: Elkmont, C. H. Kauffman, 68 (in Mo. Bot. Gard.
Herb., 1680). `

Illinois: Glencoe, E. T. & S. A. Harper, 650, 820.

Wisconsin: Madison, C. J. Humphrey, 2159 (in Mo. Bot. Gard.
Herb., 4597).

Oregon: Corvallis, W. A. Murrill, 1011, comm. by N. Y. Bot.
Gard. Herb. (in Mo. Bot. Gard. Herb., 55714).

Jamaica: Farr, 1617 (in N. Y. Bot. Gard. Herb., and Mo. Bot.
Gard. Herb., 61489). This reference is doubtful for the
hymenium is in poor condition.

47. P.inflata Burt, n. sp.

Type: in Burt Herb. and probably in N. Y. Bot. Gard. Herb.

Fructifications effused, thin, tender, soft, membranaceous,
separable, brittle when dry and cream color to cream-buff, the
subieulum and margin white and cottony; in section 150 џ thick,
not colored, 2-layered, consisting of (1) a layer next to substratum
75 a thick of loosely arranged, thin-walled, lax, hyaline hyphae
214-3 » in diameter bearing short lateral branches, each with 2

[Vol. 12
268 ANNALS OF THE MISSOURI BOTANICAL GARDEN

moniliform inflations, and of (2) a hymenial layer of erect hyphae
densely arranged, and of numerous cystidia in all regions of this
layer; no gloeocystidia; cystidia incrusted or not incrusted, 15-24
X 3-3 в, protruding up to 18 beyond the База; spores
colorless, even, 3 X 2-214 y, flat on one side, copious.

Fraotificatións 8—4 ст. long, 1-114 em. wide.

On very rotten wood. Jamaica. December. Probably rare.

P. inflata is so loosely attached to the substratum that careful
handling is necessary to prevent fructifications from becoming
detached from the wood during examination. The pair of monil-
iform inflations on short lateral branches of hyphae of the
hyphal layer shows distinctly in sectional preparation and
promises to be as helpful a character in the recognition of this
species as the details of hyphal structure in Stereum purpureum,
Corticium investiens, Grandinia granulosa, and others.

Specimens examined:
Jamaica: Hope Gardens, W. A. Murrill, 4, type, comm. by N. Y.

Bot. Gard. Herb.

48. Р. Sheari Burt, n. sp.

Type: in Burt Herb.

Fructification effused, rather thick, membranaceous, drying
pinkish buff, somewhat tubercular, somewhat velvety, not waxy,
the margin becoming somewhat free and curling up in drying,
separable from the substratum when moistened; in section 800-
1000 » thick, not colored, 2-layered, the layer next to the sub-
stratum up to 800-900 u broad and composed of densely inter-
woven, hyaline hyphae not incrusted, not nodose-septate, thick-
walled, Зи in diameter, the hymenial layer 100-150 u broad,
containing throughout great numbers of slender, rough-walled or
minutely inerusted cystidia 30-45 x 4-6 џ; no gloeocystidia;
basidia with 4 sterigmata; spores petia even, 10-12 x 6-7 y.

Fructifications 3 mm.-2 em. in diamete

On dead Alnus. Blue Mt., Oregon. Pese азн гаге
and local.

The fructifications apparently originate as сай ана {тот
lenticels in the bark and spread laterally over more or less circular
areas and become confluent. The occurrence on Alnus, tuber-

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. xiv 269

cular surface, numerous and small cystidia confined to the
hymenial layer, and spores 12 X 6 y. form a distinctive group of
characters.

Specimens examined:
Oregon: Blue Mt., C. L. Shear, 797, type.

49. P.Ravenelii Cooke, Grevillea 8: 21. pl. 124, f. 12. 1879;
Sacc. Syll. Fung. 6: 643. 1888; Massee, Linn. Soc. Bot. Jour.
25: 150. 1889.

Type: in Kew Herb.

Fructifications broadly effused, adnate, thin, small pieces sepa-
rable when moistened, becoming pale pinkish buff to pinkish buff
in the- herbarium, and somewhat cracked, the margin thinning
out; in section 100-300 u thick, not colored, composed of erect
and densely interwoven hyaline hyphae and very numerous
cystidia in all regions of the fructifications and having a somewhat
layered arrangement in thick specimens; no gloeocystidia; cystidia
heavily and coarsely incrusted, conical, with apex obtuse or
barely acute, 30-40 x 12-18 » when deeply immersed, or 30 x
8-10 y. in the hymenium; spores white in a spore collection, even,
4-5 x 2-3 y.

Fructifieations 2-8 em. long, 1-3 em. wide.

On bark and wood of decaying logs of Quercus and other
frondose species. District of Columbia to Mexico, in the Island
of Guam, and in Japan. July to January. Frequent.

' P. Ravenelii is distinguished by its small spores, coarsely in-

crusted, short cystidia with broad base, and absence of gloeo-

cystidia. Р. Roumeguerii is similar in aspect but becomes much
thicker and has longer, slenderer, and more taper-pointed
cystidia and is more distinctly layered.

Specimens examined:

Exsiccati: Ravenel, Fungi Am., 720, under the name Corticium
laeve; Ravenel, Fungi Car. 2: 39, under the name Corticium
laeve.

District of Columbia: Takoma Park, C. L. Shear, 1345.

South Carolina: H. W. Ravenel, type (in Kew Herb.), and in
Ravenel, Fungi Car. 2: 39.

Georgia: Darien, H. W. Ravenel, in Ravenel, Fungi Am., 720;

[Vol. 12
270 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Tallulah Falls, A. B. Seymour, comm. by W. G. Farlow, 13
(in Mo. Bot. Gard. Herb., 44597).

Florida: Brooksville Hammock, W. A. Murrill, 166, comm. by
N. Y. Bot. Gard. Herb. (in Mo. Bot. Gard. Herb., 62115);
Cocoanut Grove, В. Thazter, 96 (in Farlow Herb., and Mo.
Bot. Gard. Herb., 43924); Daytona, R. A. Harper, ? (in Mo.
Bot. Gard. Herb., 54539); New Smyrna, W. A. Murrill, 6,
comm. by N. Y. Bot. Gard. Herb., 62087.

Alabama: Auburn, F. S. Earle & C. F. Baker (in N. Y. Bot.
Gard. Herb., and Mo. Bot. Gard. Herb., 61345).

Louisiana: Bogalusa, C. J. Humphrey, 5495 (in Mo. Bot. Gard.
Herb., 13882); St. Martinville, A. B. Langlois, 2689, 2693
(in N. Y. Bot. Gard. Herb., and Mo. Bot. Gard. Herb., 61457,
61436), and 2692, ar, as, bp, bs, ci, and co.

Mexico: Orizaba, W. A. & E. L. Murrill, 765, comm. by N. Y.
Bot. Gard. Herb., 54647.

Island of Guam: Edwards, comm. by J. R. Weir, 10775 (in Mo.
Bot. Gard. Herb., 56239).

Japan: Prov. Awaji, Mt. Mikuma, A. Yasuda, 39, 56, 79 (in
Mo. Bot. Gard. Herb., 56156, 56159, 56313).

50. P. Roumeguerii Bresadola in litt., n. comb.

Corticium Roumeguerii Bresadola, Fungi Trid. 2: 36. pl. 144,
f. 1. 1892; Roumeguére, Rév. Myc. 15: 31 pag. sep. pl. 136,
f. 18 b. 1893; Sacc. Syll. Fung. 11: 125. 1895.—Kneiffia
Roumeguerii Bresadola, Ann. Мус. 1: 102. 1903.—Corticium
Mollerianum Bresadola in Saccardo, Soc. Brot. Bol. 11: 13.
1892.—Peniophora Molleriana (Bres.) Saccardo, Soc. Brot. Bol.
11: 13. 1892; Басс. Syll. Fung. 11: 128. 1895; v. Hóhnel &
Litschauer, K. Akad. Wiss. Wien Sitzungsber. 117: 1092. 1908;
Bourdot & Galzin, Soc. Myc. Fr. Bul. 28: 401. 1913; Wake-
field, Brit. Мус. Soc. Trans. 5: 132. 1915; Rea, Brit. Basid.
693. 1922.

Type: type distribution in Roumeguere, Fungi Gallici, 506.

Fructifications broadly effused, adnate, becoming rather thick,
small pieces separable when moistened, whitish at first, becoming
pale pinkish buff to pinkish buff in the herbarium, the margin —
thinning out; in section 100-700 y thick, not colored, becoming

1925] ; P
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 271

layered in thick specimens, composed of erect and interwoven,

closely agglutinate hyphae 2-3 » in diameter and of very nu-

merous cystidia; no gloeocystidia; cystidia incrusted, 35-80 x

8-12 u, acute, numerous in all layers except next to the sub-

stratum; spores hyaline, even, 4-6 x 2-3 y.

Fructifications 3-8 em. long, 1-4 em. wide.

On bark of logs of Quercus, Eucalyptus, Citrus, Ficus, and other
frondose species, rarely on conifers. In Europe, and in Alabama,
Louisiana, Missouri, Idaho, British Columbia to California, and
in the West Indies. May to February. Not common.

P. Roumeguerii is possibly a synonym of P. Ravenelii, as I
formerly regarded it, but the numerous specimens which have
been studied lead me to believe that while of the same aspect,
spore characters, and substratum, P. Roumeguerii eventually
becomes twice as thick as P. Ravenelii, more closely agglutinate,
and its cystidia longer and slenderer in proportion to isque thick-
ness. The error of v. Hóhnel & Litschauer, loc. cit.,
stating the year of publication of P. Molleriana as 1891 т prob-
ably led more recent European authors into reducing P. Roume-
диети to synonymy while it really has priority.

Specimens examined:

Locality not stated: G. Bresadola, authentie specimen under the
name Peniophora Roumeguerii Bres.

Italy: Trient, G. Bresadola, authentic specimen of Peniophora
Molleriana.

France: Aveyron, A. Galzin, 17908, comm. by H. Bourdot, 16898.

England: Symond's Yat, E. M. Wakefield (in Mo. Bot. Gard.
Herb., 44759).

Alabama: Montgomery County, R. P. Burke, 364 (in Mo. Bot.
Gard. Herb., 57232).

Louisiana: Baton Rouge, Edgerton & Humphrey, comm. by C. J.
Humphrey, 5646, 5648; St. Martinville, A. B. Langlois, 1346,
comm. by W. G. Farlow, 2675, 2683, 2970, cj, ck, and another
specimen, comm. by Lloyd Herb., 3042.

Missouri: Creve Coeur Lake, L. O. Overholts, 3165 (in Mo. Bot.
Gard. Herb., 5709).

Idaho: Santa, E. E. Hubert, comm. by J. R. Weir, 12001 (in Mo.
Bot. Gard. Herb., 63363).

[Vol. 12
272 ANNALS OF THE MISSOURI BOTANICAL GARDEN

British Columbia: Sidney, J. Macoun, 379 (in Mo. Bot. Gard.
Herb., 55330).

Oregon: Tidewater, S. M. Zeller, 1983 (in Mo. Bot. Gard. Herb.,
58760).

California: Berkeley, C. J. Humphrey, 5987, 5990; Redding, C. J.
Humphrey, 6038; Santa Barbara, O. M. Oleson, 10.

Cuba: Horne (in N. Y. Bot. Gard. Herb., and Mo. Bot. Gard.
Herb., 61464).

Porto Rico: Rio Piedras, J. A. Stevenson, 5792 (in Mo. Bot. Gard.
Herb., 54693); Sabana Llana, J. A. Stevenson, 6058 (in Mo.
Bot. Gard. Herb., 54686); Vega Baja, J. A. Stevenson, 5693
(in Mo. Bot. Gard. Herb., 54692).

51. Р. hiulca Burt, n. sp.

Type: in Burt Herb. and probably in N. Y. Bot. Gard. Herb.

Fructifications long and widely effused, thick, membranaceous,
separable when moistened, becoming light buff to warm buff in
the herbarium, widely cracked, the margin determinate, some-
what tomentose; in section 250-1400 џ thick, not colored, 2-
layered, with a very thick layer next to the substratum of
densely interwoven, longitudinally arranged and somewhat as-
cending thin-walled, hyaline hyphae 3-4 u in diameter, not in-
crusted, not nodose-septate and with the hymenial layer thinner
—only 100-200 „ thick—and containing in all portions very nu-
merous cystidia; no gloeocystidia; cystidia incrusted, somewhat
conical, 30-50 x 6-12 u, very numerous, wholly immersed or
protruding up to 30 u; basidia with 4 sterigmata; spores hyaline,
even, 415-5 x 3 y.

Fructifications 4-12 em. long, 2-4 em. wide— perhaps larger for
all specimens received are fragmentary.

On bark and decaying wood of frondose species. Mexico and
the West Indies. November to May. `

P. Миса has large, conspicuous fructifications with somewhat
the color and aspect of P. mutata and P. Roumeguerit. The
absence of gloeocystidia and the smaller spores distinguish it from
the former, and the comparatively thin hymenial layer to which
cystidia are restricted and the very thick layer of interwoven
hyphae running in all directions, rather than predominantly
erect, from P. Roumeguerii.

1925]
URT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 273

Specimens examined: ;

Mexico: Jalapa, W. A. & E. L. Murrill, 192, comm. by N. Y.
Bot. Gard. Herb. (in Mo. Bot. Gard. Herb., 54549).

Bermuda: S. Brown, N. L. Britton & F. J. Seaver, 1507, comm.
by N. Y. Bot. Gard. Herb.

Jamaica: Castleton Gardens, W. A. & E. L. Murrill, 71, type,
comm. by N. Y. Bot. Gard. Herb.; Mandeville, A. E. Wight,
comm. by W. G. Farlow.

52. P. phosphorescens Burt, n. sp.

Type: in Burt Herb. and probably in Farlow Herb.

Fructifications effused, membranaceous, separable, becoming
clay-color to avellaneous in the herbarium, and widely cracked
into rectangular portions about 5 mm. in diameter, which curl
up somewhat from the substratum along the fissures and show
the whitish, cottony subiculum, the hymenium waxy, somewhat
tubercular and minutely spotted in the type, the margin thinning
out; in section 300—500 џ thick, not colored, 2-layered, with the
layer next to the substratum composed of loosely interwoven
hyphae 3-344 in diameter, the hymenial layer up to 200 y
thick, composed of densely arranged hyphae and cystidia; no
gloeocystidia; cystidia incrusted, 70-100 x 12-18 y, fusiform,
acute, sometimes tilted, immersed throughout the hymenial layer,
few protruding; spores hyaline, even, subglobose, 4-5 X 3-314 u;
said to be phosphorescent when collected.

Fructifications probably large, for pri consist of frag-
ments 7 x 116 em., and 114-3 em. in diamete

On rotten wood of fence post and ас bark of frondose
species. Jamaica. October to December.

P. phosphorescens may be recognized by the thick, clay-colored
fructifications which contract in drying so as to crack into rec-
tangular masses about 5 mm. in diameter, separated from one
another by rather wide fissures. The thick, hymenial portion of
each mass is so weakly attached to the substratum by the loose
subieulum that these masses curl upward along their edges and
may occasionally become wholly detached. The cystidia are
suggestive of those of P. flavido-alba but all other characters of
these two species are different. Phosphorescence has been re-
corded for but few fungi.

[Vol. 12
274 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Specimens examined:
Jamaica: A. E. Wight, type, comm. by W. G. Farlow; Castleton
Gardens, F. S. Earle, 240, comm. by N. Y. Bot. Gard. Herb.

53. P. sanguinea (Fr.) Bresadola in v. Hóhnel & Litschauer,
K. Akad. Wiss. Wien Sitzungsber. 115: 1588, 1589. 1906; Bour-
dot & Galzin, Soc. Мус. Fr. Bul. 28:395. 1913; Rea, Brit. Basid.
690. 1922.

Thelephora sanguinea Fries, Elenchus Fung. 1: 203. 1828.—
Corticium sanguineum Fries, Epicr. 561. 1838; Hym. Eur. 650.
1874; Icones Hym. 2:97. pl. 198,f. 2. 1877; Sace. Syll. Fung. 6:
612. 1888; Wakefield, Brit. Myc. Soc. Trans. 4: 119. we 8; f.
18-20. 1913.—Kneiffia sanguinea (Fries) Bresadola, Ann. Myc.
1: 101. 1903.—Corticium glabrum Berkeley & Curtis, Grevillea
1: 178. 1873; Басс. Syll. Fung. 6: 620. 1888; Massee, Linn.
Soc. Bot. Jour. 27: 142. 1890.—(In part) Corticium Petersii
Berkeley & Curtis, Grevillea 1: 177. 1873.

Fruetification effused, somewhat membranaceous, tender,
dragon's-blood red, substance arachnoid, the margin byssoid or
fibrillose and often connected with mycelial strands of blood-red
color which stain the wood red, hymenium drying light buff and
pinkish buff to buff-pink; in section 200—500 u thick, not colored,
with the hyphae loosely arranged, 3-6 и in diameter, and with
some granule-incrusted, rarely nodose-septate; cystidia hair-like,
not incrusted usually, about 414 y in diameter, protruding 20-
30 u; spores white in spore collection, even, 4-5 x 2-214 y.

Fructifications 2-10 cm. long, 1-4 em. wide.

On dead wood and fallen branches especially of conifers.
Europe, New Hampshire to Louisiana, and in Oregon. July to
January. Infrequent.

P. sanguinea and P. miniata may be recognized by the blood-
red color of the young fructifications, the more or less numerous
red mycelial strands, and the wood stained red. Later in fertile
stage the hymenium tends toward a buff color with a tinge of red.
In section P. sanguinea shows granule-incrusted hyphae more or
less numerous among other even-walled hyphae, while P. miniata
contains no incrusted hyphae.

Specimens examined:

1925]
URT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 275

Exsiccati: Ell. & Ev., Fungi Col., 1020, under the name Corticium
radiosum.

Sweden: L. Romell, 130; Stockholm, L. Romell, 136.

Austria: Tirol, V. Litschauer.

France: F. Fautrey, from Lloyd Herb., 3308.

New Hampshire: Chocorua, W. G. Farlow, 10, E. A. Burt, 8, 4.

New York: Hudson Falls, S. H. Burnham, 21 (in Mo. Bot. Gard.
Herb., 54490); Karner, H. D. House, 14.157 (in Mo. Bot.
Gard. Herb., 44704); Oneida, H. D. House (in N. Y. State
Mus. Herb., and Mo. Bot. Gard. Herb., 57434); Wymantskill,
C. H. Peck (in N. Y. State Mus. Herb., and Mo. Bot. Gard.
Herb., 56051).

New Jersey: Newfield, J. B. Ellis, in Ell. & Ev., Fungi Col., 1020.

Pennsylvania: State College, L. O. Overholts, 3422 (in Mo. Bot.
Gard. Herb., 54476).

South Carolina: Society Hill, types of Corticium glabrum, Curtis

. Herb., 2404 (in Curtis Herb.) and 3719 (in Kew Herb.).

Florida: W. W. Calkins, 845 (in Burt Herb., Farlow Herb., and
Mo. Bot. Gard. Herb., 63421).

Alabama: Peters, 847, under the name Corticium miniatum (in
Curtis Herb., 5225), and Peters, 473, one of the types of Cor-
ticium Рите (in Curtis Herb., 4509).

Louisiana: St. Martinville, A. B. (put lois, 2 704.

Oregon: Corvallis, S. M. Zeller, 1860 (in Mo. Bot. Gard. Herb.,
56868).

54. P. limonia Burt, n. sp.

Type: in Mo. Bot. Gard. Herb.

Fructifications broadly effused, compact, fleshy-membranace-
ous, small pieces separable when moistened, cream-bufi, not
cracked, the margin byssoid and with some radiating, cream-buff
mycelial strands; in section 200 y. thick, not perceptibly colored,
2-layered next to the substratum, with very coarse, heavily in-
erusted, loosely arranged, longitudinally interwoven hyphae
6-9 и in diameter, and with the hymenial layer 75 џ thick and
composed of erect tissues; no gloeocystidia; cystidia not in-
crusted, 45 x 414 u, tapering to a sharp apex, protruding 20-
27 » beyond the basidia; spores hyaline, even, 3-4 X 275 в.

[Vol. 12
276 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fructifications 215-4 em. long, 1-14 cm. wide.

On bark of decaying Robinia neo-mexicana. New Mexico.
August

P. limonia has the color of P. sulphurina and P. carnosa but
differs from both by its occurrence on frondose bark and very
coarse, heavily incrusted hyphae. The hymenial layer does not
crack and flake away from the substratum like that of P. sul-
phurina. Treatment of sections with potassium hydrate solution
causes no color changes.

Specimens examined:
New Mexico: Sulphur Canyon, W. H. Long, 21405, type (in

Mo. Bot. Gard. Herb., 55146).

55. Р. amoena Burt, n. sp.

Type: in Mo. Bot. Gard. Herb.

Fructifications long and broadly effused, thin, adnate, small .
pieces separable when moistened, cream-color in the herbarium,
even, glabrous, the margin thinning out, of finely interwoven
hyphae; in section 120 u thick, not colored, with the hyphae near
the substratum compactly interwoven, about 3 y in diameter;
an incrusted subhymenial zone present, formed of numerous
inerusted bodies side by side; no gloeocystidia; cystidia of the
hymenial surface not incrusted, 7-9 и in diameter, protruding
up to 45 в; База rather large, 25-30 x 5-6 и, with 4 sterig-
mata; spores hyaline, even, 12-15 X 4-6 y, copious.

Fructifieations probably large, for pieces broken off at one end
and one side are 5-6 em. long, 115-2 em. wide.

On a soft wood of a frondose species. British Columbia.

P. amoena forms cream-colored, somewhat waxy fructifications
on decorticated logs of a pale soft wood—perhaps Populus. The
spores are so large as to afford a valuable specific character.

Specimens examined:

British Columbia: Sidney, J. Macoun, 7, type (in Mo. Bot. Gard.

Herb., 5766).

56. P. firma Burt, n. sp.
Type: in Burt Herb.
Fructifications effused, rather thick, dry, firm, membranaceous,

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 277

small pieces separable when moistened, cream-buff in the her-
barium, even, not cracked, the margin thinning out, fibrillose;
in section 300-500 џ thick, not colored, with the hyphae 4-6 y. in
diameter near the substratum, densely interwoven, ascending
and becoming finer, sometimes incrusted towards the hymenial
layer; no gloeocystidia; cystidia not incrusted, tapering upward
to a sharp point, 4—5 u at base, protruding 20-35 y, confined to
surface of hymenium, numerous; spores hyaline, even, 4-5 X
2V$-3 y.
Fructifications 2-5 em. long in pieces broken off at both ends,
3—5 em. wide.
On rotten wood of Alnus (?) and on bark of Robinia neo-
mexicana. Washington and Arizona. September and October.
P. firma resembles P. Rowmeguerii in general aspect but its
cystidia are slenderer than those of P. Roumeguerii, not incrusted,
and present in the hymenial surface only.
Specimens examined:
Washington: Arlington, C. J. Humphrey, 7609, type.
Arizona: Santa Catalina Mountains, Coronado National Forest,
G. G. Hedgcock & W. H. Long, comm. by C. J. Humphrey,
2555 (in Mo. Bot. Gard. Herb., 12262).

57. P. miniata (Berk.) Burt, n. comb.

Thelephora miniata Berkeley in Hooker, Eng. Flora 2°: 168.
1836; Brit. Fungi, No. 251. 1843. See v. Hóhnel & Litschauer,
K. Akad. Wiss. Wien Sitzungsber. 115: 1588. 1906.

Type: authentic specimen in Berkeley, Brit. Fungi, 251.

Fructification effused, somewhat membranaceous, tender,
English red, substance arachnoid, the margin byssoid or fibrillose
and often connected with mycelial strands of blood-red color;
hymenium drying pinkish buff to buff-pink and diónamon-rufous:
in section 150-300 » thick, not colored, the hyphae loosely ar-
ranged, 3-6 y. in diameter, not incrusted, rarely nodose-septate;
cystidia few, hair-like, not incrusted, 326-426 in diameter,
protruding 20-30 ш; spores hyaline, even, 4-414 x 2-214 y.

Fructifications 2-10 cm. long, 1-2% em. broad.

On fallen limbs, usually of conifers. In England, New Hamp-
shire to Louisiana, and in Washington and Oregon. July to De-
cember. Infrequent.

[Vol. 12
278 ANNALS OF THE MISSOURI BOTANICAL GARDEN

The twenty gatherings cited below have been separated from
P. sanguinea by the absence of incrusted hyphae in their sectional
preparations. In the original description of Т. miniata, Berkeley
stated, ‘‘This most elegant species differs so much from T. san-
guinea Fr., in being most highly colored where exposed to light,
while in the portions to which light has not free access it is nearly
white, and in not tinging the wood whereon it grows with its
own color, that an inspection of specimens renders it almost im-
possible to consider it the same." Fifteen of the twenty speci-
mens referred below to P. miniata on account of absence of
hyphal inerustation have the hymenium red and only five
pinkish buff, while none of the twenty specimens show the wood
stained red.

Specimens examined:

Exsiecati: Berkeley, Brit. Fungi, 251, authentic Specimen of
Thelephora miniata Berk.

England: Berkeley, Brit. Fungi, 251.

New Hampshire: Chocorua, E. A. Burt, 1, 2.

New York: Albany, H. D. House (in N. Y. State Mus. Herb.,
and Mo. Bot. Gard. Herb., 14830, 55201), and H. D. House &
J. Rubinger (in N. Y. State Mus. Herb., and Mo. Bot. Gard.
Herb., 17797); Karner, H. D. House, comm. by N. Y. State
Mus. Herb. (in Mo. Bot. Gard. Herb., 54347, 54357, 54377);
Newtonville, C. H. Peck (in N. Y. State Mus. Herb., and Mo.
Bot. Gard. Herb., 55969, 55989) ; North Elba, C. H. Kauffman, 5
(in Mo. Bot. Gard. Herb., 6719); Schuylerville, C. H. Peck,
comm. by N. Y. State Mus. Herb., T 17, T 25 (in Mo. Bot.
Gard. Herb., 54570, 54657).

Georgia: Tallulah Falls, A. B. Seymour, from Farlow Herb., D
(in Mo. Bot. Gard. Herb., 44609).

Louisiana: St. Martinville, A. B. Langlois, bu.

Washington: Chehalis, C. J. Humphrey, 6273; Hoquiam, C. J.
Humphrey, 6409.

Oregon: Granite Pass, J. R. Weir, 11183 (in Mo. Bot. Gard.
Herb., 63252).

58. P. Burtii Romell, n. sp.
Type: in Burt Herb. and Romell Herb.

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. ХІУ 279

Fructifications effused, somewhat membranaceous, tender,
hymenium drying warm buff usually but sometimes whitish to
cartridge-buff, sometimes cracked and showing the cottony sub-
stance, the margin byssoid or fibrillose and sometimes connected
with antimony-yellow mycelial strands; in section 200-300 y
thick, not colored, with the hyphae loosely arranged, hyaline,
rarely nodose-septate, with some incrusting granules in the sub-
hymenium; cystidia hair-like, not incrusted, tapering, 3-4 » in
diameter, protruding up to 25 в, not numerous; spores hyaline,
even, 4414 x 2-21 u.

Fructifications 2-7 cm. long, 1-2 em. broad.

On wood and fallen limbs of frondose species in woods. Ver-
mont to Louisiana and in Ohio, Michigan, and Montana. July
to October. Rare.

This species is noteworthy by the antimony-yellow or ochra-
ceous mycelial strands or eords which grow from under the bark
and connect with the fructifications. The presence of cystidia
separates this species from Corticium sulphureum which has
yellower fructifications and not as large mycelial cords when
present. ;

Specimens examined:

Exsiccati: Ellis, N. Am. Fungi, 933, under the name Corticium
radiosum.

Vermont: Middlebury, E. A. Burt.

Massachusetts: Sharon, A. P. D. Piguet, 136, comm. by Farlow

Herb. (in Mo. Bot. Gard. Herb., 59627).

New York: Ithaca, H. H. Whetzel, comm. by Cornell Univ.

Herb., 13760.

New Jersey: Newfield, J. B. Ellis, in Ellis, N. Am. Fungi, 933.
Virginia: Crabbottom, W. A. Murrill, 239 (in N. Y. Bot. Gard.

Herb., Burt Herb., and Mo. Bot. Gard. aga 61560).
Alabama: Auburn, Alabama Biological Surv
Louisiana: Bogalusa, C. J. Humphrey, 542; St. Martinville,

A. B. Langlois, cl.

West Virginia: Paw Paw, C. L. Shear, 1179.
Ohio: C. G. Lloyd, 3823, type.
Michigan: New Richmond, C. H. Kauffman, 34 (in Mo. Bot.

Gard. Herb., 23060).

[Vol. 12
280 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Montana: Evaro, J. R. Weir, 415 (in Mo. Bot. Gard. Herb.,
14772).

59. P. subapiculata (Bres.) Burt, n. comb.

Corticium subapiculatum Bresadola, Mycologia 17: 69. 1925.

Type: in Weir Herb.

Fructifications broadly effused, adnate, small pieces separable
when moistened, waxy, becoming ivory-yellow to pinkish buff in
the herbarium, even, only rarely eracked, the margin thinning
out, pruinate; in section about 150 x thick, not colored, composed
of interwoven hyaline hyphae 314-414 u in diameter, not in-
crusted, only rarely nodose-septate; no gloeocystidia; cystidia
hair-like, not incrusted, cylindric, obtuse, 3-414 u in diameter,
protruding 10-40» beyond the basidia; spores hyaline, even,
4-6 x 2-3 y.

Fructifications 8-12 em. long, 1-4 em. wide.

On decaying logs of Pinus, Abies, and Lariz—usually on the
wood. Idaho and British Columbia. June to September.

P. subapiculata resembles P. Weiri in color and general aspect
but has no gloeocystidia and smaller cystidia and spores.

Specimens examined:

Montana: Evaro, J. R. Weir, 414 (in Mo. Bot. Gard. Herb.,
63720); Kalispell, E. E. Hubert, comm. by J. R. Weir, 11957
(in Mo. Bot. Gard. Herb., 63312).

Idaho: Clarkia, A. S. Rhoades (in Weir Herb., 16928, type);
Coolin, J. R. Weir, 11086 (in Mo. Bot. Gard. Herb., 63245);
Priest River, J. R. Weir, 52, 130 (in Mo. Bot. Gard. Herb.,
63718), and E. E. Hubert, comm. by J. R. Weir, 12020 (in Mo.
Bot. Gard. Herb., 63375).

British Columbia: Kootenai Mountains, near Salmo, J. R. Weir,
476 (in Mo. Bot. Gard. Herb., 63719).

60. P. sordida (Karst.) Burt—not in the sense of Brinkmann
or Bresadola.

Corticium sordidum Karsten, Soc. pro Fauna et Flora Fennica
Meddel. 9: 65. 1883; Finska Vet.-Soc. Bidrag Natur och
Folk 48: 413. 1889; Sacc. Syll. Fung. 6: 631. 1888; Massee,
Linn. Soc. Bot. Jour. 27: 140. 1890. Compare v. Hóhnel

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 281

& Litschauer, K. Akad. Wiss. Wien Sitzungsber. 117: 1088.
1908.

Type: authentic specimen in Burt Herb.

Fructifications longitudinally effused, small portions separable
when moistened, in the herbarium young specimen pale olive-
buff and older specimen wood-brown, contracting in drying and
eracking into small rectangular masses about 1 mm. in diameter,
separated by rather wide crevices and showing the paler floccose
subieulum, the margin thinning out; in section 150-300 џ thick,
not colored, becoming stratose, each stratum 2-layered, with the
layer towards the substratum composed of loosely arranged,
suberect, branching hyphae 414-5 y, rarely 6 р, in diameter, not
incrusted, not nodose-septate, and the hymenial layer compact,
75 р thick; no gloeocystidia; cystidia not incrusted, cylindric,
obtuse, 4—6 u in diameter, protruding up to 30 y, none wholly
immersed; spores copious, hyaline, even, 415-6 x 2-3 y.

Fructifications 3 cem. X 7 mm. and 24% x 1 em. in the two
fragmentary pieces from Karsten, 3-10 em. long, 7-15 mm. wide
in an American specimen.

On decorticated wood of P. sylvestris and P. Strobus on the
ground. Finland and New York. October. Rare.

Brinkmann distributed in his “Westfälische Pilze," 8, under the
name of Peniophora sordida (Karst.) Brinkmann, a specimen which
was later referred by Bresadola to Peniophora serialis. I have
not seen this specimen. Von Hóhnel & Litschauer accepted this
reference, loc. cit., and placed Corticium sordidum Karst. as a
synonym of P. serialis. 'The study of other specimens of the
P. serialis complex shows that none of these others have the
strueture of authentic Corticium sordidum although somewhat
resembling the old stage in general aspect. The problem with
me for a time was whether P. cremea is distinct from P. sordida,
but P. cremea occurs on frondose wood, is not cracked into rec-
tangular, completely separated masses, and has larger cystidia,
some of which are incrusted and wholly immersed. The Karsten
specimens of C. sordidum are in some places composed of a single
stratum 150 y thick of 2 layers and in others of 2 strata with
thickness together of 240—300 џ.

Specimens examined:

7 [Vol. 12
282 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Finland: Mustiala, P. A. Karsten, authentic specimen of Cor-
ticium sordidum.

New York: Karner, H. D. House, 14.188 (in Mo. Bot. Gard.
Herb., 44722).

61. P. Burkei Burt, n. sp.

Type: in Mo. Bot. Gard. Herb.

Fructifications broadly effused, thin, adnate, membranaceous,
tender, small pieces separable when moistened, cream-buff in the
herbarium, somewhat tubercular, conforming to the inequalities
of the rough bark upon which growing, somewhat cracked in
drying, the margin thinning out, of finely interwoven hyphae;
in section 120-180 џ thick, not colored, with the hyphae suberect,
loosely interwoven, thin-walled, 3 џ in diameter, nodose-septate,
not incrusted; по gloeocystidia; cystidia not incrusted, subulate,
90 X 414-5 u, protruding up to 20 u; spores hyaline, even, 6-7
X 4-5 y, copious.

Fructifications probably large—4 em. long, 2-214 cm. wide in
pieces broken off at both ends and on one side.

On rough, frondose bark. Alabama. October.

P. Burkei has some resemblance in aspect to P. cremea but has
a more tubercular hymenium, slenderer and more erect hyphae,
and larger spores.

Specimens examined:
Alabama: Montgomery County, R. P. Burke, 474, S (in Mo.

Bot. Gard. Herb., 57292).

62. P. glebulosa Bresadola, Fungi Trid. 2: 61. pl. 170, f. 2.
1898; Басс. Syll. Fung. 16: 195. 1902; Bourdot & Galzin, Soc.
Myc. Fr. Bul. 28:384. 1913; Rea, Brit. Basid. 688. 1922.

Not Thelephora calcea Fries var. glebulosa Fries, Elenchus Fung.
1:215. 1828.—Not Corticium calceum Fries, Epicr. 562. 1838;
nor Hym. Eur. 652. 1874.

Type: in Bresadola Herb. and Burt Herb.

Fructifications widely effused, thin, closely adnate, whitish,
pinkish buff, pale olive-buff, or cream color, pubescent with the
Neue becoming eracked into small areas when dry, the margin

out; in section 50-200 » thick, not colored, composed
ке. of eystidia and rather erect, interwoven, hyaline,

1925]
URT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 283

thin-walled hyphae 1-3 ь in diameter, not incrusted; cystidia
thick-walled, with very narrow lumen which is often much larger
at apex of cystidium, even where immersed, or sometimes with
some granular incrustation near protruding apex, 60-150 x 7-
10 p, protruding up to 50-100 u, very numerous throughout the
fructification, not dissolved by treatment of sections with potas-
sium hydrate; spores white, even, cylindrie, slightly curved,
6-9 x 116-2 y.

Fructifications 3-15 cm. long, 1-6 em. wide.

On wood of decaying conifers, rarely on bark, and on frondose
species. In Europe, from Canada to New Jersey, in Nebraska,
Colorado, Montana and Manitoba to British Columbia and
Oregon. May to November. Common locally.

P. glebulosa has distinctive cystidia to which Bourdot & Galzin
have given the term cystidioles. These cystidia are elongated,
cylindric, even throughout their whole length usually, but some-
times with a little incrusting matter near the apex of the pro-
truding part, and with a very thick wall—so thick that the axial
lumen containing protoplasmie contents is merely a line which,
however, is often greatly expanded at its peripheral end in the
apex of the cystidium where the latter becomes thin-walled and
fragile. The cystidia of P. glebulosa are not at all dissolved or
only partially by the potassium hydrate treatment to which sec-
tions are subjected. In the original description Bresadola
states that P. glebulosa is the same as authentic Corticium calceum
var. glebulosum Fries. I believe this to be an error, for a frag-
ment of authentic C. calceum var. glebulosum communicated to
me by Bresadola and the original specimens so labelled in Fries
Herbarium, all of which I studied, have no cystidia whatever and
agree in all respects with a true Corticium collected at Femsjó,
the original station, by Romell and myself.

Specimens examin
Sweden: Femsjé, Romell & Burt, three gatherings; Lappland,

L. Romell, 405; Stockholm, L. Romell, 199.

Austria: Stubai, Tirol, V. рай
Italy: Trient, G. Bresadola, t;
England: Symond's Yat, Е. TM. Wakefield (in Mo. Bot. Gard.

Herb., 57120).

[Vol. 12
284 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Canada: Billings Bridge, J. Macoun, 113.

Quebec: Hull, J. Macoun, 246.

Maine: W. A. Murrill, 2139Y5 (in №. Y. Bot. Gard. Herb., and
Mo. Bot. Gard. Herb., 61424); Kittery Point, R. Thaxter &
E. A. Burt; Piscataquis County, W. A. Murrill, 2142, 2653 (in
N. Y. Bot. Gard. Herb., and Mo. Bot. Gard. Herb., 61349,
61441).

New Hampshire: Chocorua, W. G. Farlow (in Mo. Bot. Gard.
Herb., 19554) and 11.

Vermont: Middlebury, Е. A. Burt, four gatherings.

New York: Altamont, E. A. Burt; East Galway, E. A. Burt,
three gatherings; Farmington, E. Brown, 116 (in N. Y. Bot.
Gard. Herb. and Mo. Bot. Gard. Herb., 61451); Ithaca,
G. F. Atkinson, 8236, 8284; Sandlake, C. H. Peck, comm. by
N. Y. State Mus. Herb., T 15 (in Mo. Bot. Gard. Herb.,
54568).

New Jersey: Newfield, J. B. Ellis, under the herbarium name
P. gracillima (in N. Y. Bot. Gard. Herb., and Burt Herb.).

Nebraska: Long Pine, C. L. Shear, 1056.

Colorado: Geneva Creek Canyon, alt. 8000-14000 ft., F. J.
Seaver & E. Bethel (in N. Y. Bot. Gard. Herb., and Mo. Bot.
Gard. Herb., 61467) ; Golden, L. O. Overholts, 1752 (in Mo. Bot.
Gard. Herb., 54881). |

Montana: Bernice, J. R. Weir, 12000, 12006 (in Mo. Bot. Gard.
Herb., 63362, 63366); Darby, E. E. Hubert, comm. by J. R.
Weir (in Mo. Bot. Gard. Herb., 63248); Hecla, E. E. Hubert,
comm. by J. R. Weir, 11416 (in Mo. Bot. Gard. Herb., 63263);
Libby, E. E. Hubert, comm. by J. R. Weir, 11443 (in Mo. Bot.
Gard. Herb., 63273).

Idaho: Coeur d'Alene, J. R. Weir, 11974, and E. E. Hubert,
comm. by J. R. Weir, 11991 (both in Mo. Bot. Gard. Herb.,
63328 and 63354 respectively); Coolin, J. R. Weir, 11562 (in
Mo. Bot. Gard. Herb., 63301); Priest River, E. E. Hubert,
comm. by J. R. Weir, 12029 (in Mo. Bot. Gard. Herb., 63381),
and J. R. Weir, 350, 6362 (in Mo. Bot. Gard. Herb., 7853,
55952) and 54.

Manitoba: Norway House, G. R. Bisby, 1458, 1464, 1476 (in
Mo. Bot. Gard. Herb., 61640, 61646, 61658).

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 285

British Columbia: Agassiz, J. R. Weir, 364 (in Mo. Bot. Gard.
Herb., 16407); Comax, J. Macoun, 622 (in Mo. Bot. Gard.
Herb., 55333); Kootenai Mts., Salmo, J. R. Weir, 485, 538
(in Mo. Bot. Gard. Herb., 17619, 1738) ; Sidney, J. Macoun, 22,
41, 64, 97 (in Mo. Bot. Gard. Herb., 5682, 55342, 5742, 55343);
Squamish, J. Macoun (in Mo. Bot. Gard. Herb., 55180);
Vancouver Island, J. Macoun, 356, 357 (in Mo. Bot. Gard.
Herb., 55331, 55332); Victoria, J. Macoun, 576 (in Mo. Bot.
Gard. Herb., 63502).

Washington: Bingen, W. N. Suksdorf, 699; Hoquiam, C. J.
Humphrey, 6374; Kalama, C. J. Humphrey, 6139; Renton,
C. J. Humphrey, 6633; Tacoma, W. A. Murrill, 145, comm.
by N. Y. Bot. Gard. Herb. (in Mo. Bot. Gard. Herb., 55726).

Oregon: Corvallis, S. M. Zeller, 1813 (in Mo. Bot. Gard. Herb.,
56333) ; Eugene, C. J. Humphrey, 6086.

63. P. verticillata Burt, n. sp.

Type: in Mo. Bot. Gard. Herb.

Fructifications effused, thick, membranaceous, separable, whit-
ish to ecru-drab in the herbarium, even the margin whitish,
rather thick, cottony; in section 1300 џ thick, not colored, con-
sisting of (a) a layer 500 џ thick next to the substratum of densely,
longitudinally arranged hyaline hyphae about 3-314,» in di-
ameter, and of (b) a zonate hymenial layer 800 џ thick con-
taining many elongated cystidia; no gloeocystidia; eystidia cylin-
dric, 150-200 x 6-7 u, with 4-9 bands of incrusting matter,
protruding up to 45 u; no spores found.

Fructifications 114-214 em. long, 1-2 em. wide.

On rotten coniferous wood. Oregon. March.

The cystidia of P. verticillata are of the thick-walled cylindric
type occurring in P. glebulosa but without as narrow a lumen, nor
with the latter abruptly, greatly enlarged near the apex. The
bands of incrusting matter on the cystidia are a unique character
of the type but are not retained in glycerine mounts of sections.
The very broad layer of longitudinally arranged hyphae along the
substratum and the very thick, separable fructifications tending
to ecru-drab are probably the more distinctive characters of this
species, which is distinct from P. (Gloeocystidium) pallidula.

[Vol. 12
286 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Specimens examined: ·

Oregon: Waltersville, C. C. Epling & J. B. Shorett, 600, type,
comm. by S. M. Zeller, 2317 (in Mo. Bot. Gard. Herb.,
63041). ;

64. P. crassa Burt, N. Y. State Mus. Rept. 54: 155. 1901.

Stereum Karstenii Bresadola, I. R. Accad. Agiati Atti III. 3:
108. 1897; Bourdot & Galzin, Soc. Myc. Fr. Bul. 37: 126.
1921.—Not Peniophora Karstenii Massee, Linn. Soc. Bot. Jour.
25: 153. 1889.—Not Phanerochaete odorata Karsten, Finska
Vet.-Soc. Bidrag Natur och Folk 48: 427. 1889.—Corticium
ochroleucum, in part, of Berkeley & Curtis, Grevillea 1: 165.
1875, but not of Fries.

Type: in Burt Herb.

Fructifications broadly effused, becoming thick, somewhat
fleshy, light buff to pinkish buff, separable from the substratum
when moistened if thick, the margin somewhat tomentose, deter-
minate; in section 500-1500 џ thick, not colored, 2-layered, with
the layer next to the substratum 200-300 и, thick, composed of
densely interwoven, rather thick-walled and stiff, non-incrusted
hyphae 3-414 u in diameter, and with the hymenial layer 300-
1200 y. thick, more or less zonate, and composed of erect hyphae
and cystidia; no gloeocystidia; cystidia even or sometimes some-
what incrusted, cylindric, flexuous, 100-500 x 414-6 р, pro-
truding up to 30 u beyond the База, present in all parts of the
hymenial layer, destroyed and dissolved by potassium hydrate
treatment of sections; basidia 4-spored; spores white in spore
collection, even, curved, 414-6 x 114-2 y.

Fructifications 3-20 cm. long, 1-4 em. wide.

On decorticated, decaying logs of Pinus, Abies, Picea, Tsuga,
and Pseudotsuga. In Europe and from Canada to Alabama and
westward to the Pacific states. Common.

P. crassa is certainly cogeneric with P. glebulosa, belongs in the
same group of species, occurs on the same substrata and is prob-
ably equally destructive to wood. Its fructifications are thicker
than those of P. glebulosa and crack into larger masses. The
cystidia have thinner walls and larger lumen than those of P.
glebulosa and are noteworthy for the destructive action of potas-

1925]
URT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 287

sium hydrate on them, so that it can not be safely used in clearing

and swelling the sections. Lactic acid should be used instead.

I have included under P. crassa the two European specimens of
Stereum Karstenii cited below, because of agreement in all char-
acters except the much greater thickness of the latter and their
eurling away from substratum at the margin and separation of
the whole fructification in a sheet-like mass. American speci-
mens of P. crassa range from 500 to 1000 в. thick and have the
margin closely adnate to the substratum. Perhaps there is
specific difference between P. crassa and Stereum Karstenii.

Specimens examined:

Exsiccati: Ellis, N. Am. Fungi, 331, under the name Corticium
ochroleucum var. spumeum; Ravenel, Fungi Car. 3: 33, under
the name Corticium ochroleucum.

Hungary: A. Kmet, type of Stereum Karstenii from Bresadola.

France: Aveyron, A. Galzin, 20064, comm. by H. Bourdot, 20799.

Canada: J. M pes. 42; Quebec, J. Macoun, 260; Ottawa, J.
Macoun, 248, in

New Hampshire: gi W. G. Farlow, 23, and an unnum-
bered specimen.

Vermont: Middlebury, E. A. Burt, two gatherings; Ripton, Е. A.
Burt, type.

Massachusetts: Magnolia, W. G. Farlow, e; Sharon, A. P. D.
Piguet, 189, comm. by Farlow Herb. (in Mo. Bot. Gard. Herb.,
59360).

New York: Floodwood, E. A. Burt, C. H. Peck, 2; Ithaca, G. F.
Atkinson, 8008; Keene, C. H. Peck, comm. by N. Y. State
Mus. Herb., T 1 (in Mo. Bot. Gard. Herb., 54554); North
Elba, C. H. Peck, comm. by N. Y. State Mus. Herb., T 9 (in
Mo. Bot. Gard. Herb., 54555); Sylvan Beach, Oneida County,
Н. D. House (in N. Y. State Mus. Herb., and Mo. Bot. Gard.
Herb., 7460, 8293).

New Jorio Newfield, J. B. Ellis, in Ellis, №. Am. Fungi,’ F331.

Pennsylvania: State College, L. O. Overholts, 3631 (in*Mo. Bot.
Gard. Herb., 54703).

North Carolibé: H. W. Ravenel, 1521 (in Curtis Herb., 1763,
under the name Corticium ochroleucum var. erimosum).

South Carolina: H. W. Ravenel, in Ravenel, Fungi Car. 3: 33,

[Vol. 12
288 ANNALS OF THE MISSOURI BOTANICAL GARDEN

and (in Curtis Herb., 2169, under the name Corticium ochro-
leucum).

Alabama: Auburn, comm. by Alabama Biological Survey.

Idaho: Addie, E. E. Hubert, comm. by J. R. Weir, 11976 (in Mo.
Bot. Gard. Herb., 63329); Coolin, J. R. Weir, 11558 (in Mo.
Bot. Gard. Herb., 63299); Priest River, J. R. Weir, 108, 378,
6351 (in Mo. Bot. Gard. Herb., 16060, 21353, 55951) and
3, 24, 46, 50, 56.

British Columbia: Kootenai Mts., near Salmo, J. R. Weir, 455,
498 (in Mo. Bot. Gard. Herb., 8760, 21632) ; Revelstoke, C. W.
Dodge, 1654 (in Mo. Bot. Gard. Herb., 58788); Sidney, J.
Macoun, 63, 393 (in Mo. Bot. Gard. Herb., 5741, 55325).

Washington: Kalama, C. J. Humphrey, 6214 (in Mo. Bot. Gard.
Herb., 20431).

Arizona: Flagstaff, W. H. Long, 21386 (in Mo. Bot. Gard. Herb.,
95140); Fort Valley Experiment Station, W. H. Long, 19624
(in Mo. Bot. Gard. Herb., 20133).

65. P.subalutacea (Karst.) v. Hóhnel & Litschauer, K. Akad.
Wiss. Wien Sitzungsber. 115: 1601. 1906; Bourdot & Galzin,
Soc. Мус. Fr. Bul. 28: 387. 1913; Wakefield, Brit. Myc. Soc.
Trans. 5:133. 1914; Rea, Brit. Basid. 688. 1922.

Corticium subalutaceum Karsten, Soc. pro Fauna et Flora
Fennica Meddel. 9:65. 1883; Finska Vet.-Soc. Bidrag Natur och
Folk 48: 414. 1889; Sace. Syll. Fung. 6: 636. 1888; v. Hóhnel
& Litschauer, K. Akad. Wiss. Wien Sitzungsber. 115: 1560.
1906.—Kneiffia subalutacea (Karsten) Bresadola, Ann. Mye. 1:
104. 1903.

ү authentic specimen or perhaps part of type in Burt

erb.

Fructifications long and widely effused, very thin, closely
adnate, pale olive-buff to pinkish buff in the herbarium, hymenium
loose and rather rough under a lens, the margin thinning out; in
section 30-100 џ thick, not colored, with the hyphae interwoven,
rather rigid and thick-walled, about 214 u in diameter, not in-
crusted, cylindric, thin-walled, 414-6 u, protruding up to 60и
beyond the basidia, often starting from the substratum, some-
times somewhat clustered at slight elevations of the hymenium;

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 289

spores hyaline, even, narrowly cylindrie, slightly curved, about
415—7 X 1V$ y.

Fructifications 3-10 em. long, 1-3 cm. wide.

On decaying pine wood. Europe, New Jersey to Louisiana,
and in Washington. July to March. Rare.

The cystidia of P. subalutacea place it in the group with P.
glebulosa and P. crassa. It is thinner than either of these. It
may be distinguished from thin forms of the former by the thin-
walled cystidia which have a lumen of nearly uniform diameter
which is not abruptly and greatly enlarged near the apex of the
cystidium.

Specimens examined:

Sweden: Femsjé, E. A. Burt.

Finland: Mustiala, P. A. Karsten, authentic specimen.

Poland: Eichler, comm. by G. Bresadola.

" Austria: Tirol, V. Litschauer.

England: Baslow Foray, A. D. Cotton, comm. by E. M. Wakefield

(in Mo. Bot. Gard. Herb., 44583).

France: Aveyron, A. Galzin, 2444, comm. by H. Bourdot, 8007.

New Jersey: Newfield, J. B. Ellis, 7510, comm. by W. G. Farlow
(in Mo. Bot. Gard. Herb., 1794).

Maryland: Takoma Park, C. L. Shear, 1030.

Alabama: Montgomery County, R. P. Burke, 635 (in Mo. Bot.

Gard. Herb., 63071).

Louisiana: St. Martinville, A. B. Langlois, 0.
Washington: Mt. Paddo, W. N. Suksdorf, 726.

66. P. odorata (Karsten) Burt, n. comb.

Phanerochaete odorata Karsten, Finska Vet.-Soc. Bidrag Natur
och Folk 48: 427. 1889. Not Stereum odoratum Fries, Epicr.
553. 1838.—Not Stereum Karstenii Bresadola, I. В. Accad.
Agiati Atti III. 3: 108. 1897.

Type: in Burt Herb. from Karsten and probably in Karsten
Herb.

Fructifications narrowly effused, small, pulvinate, somewhat
convex, becoming longitudinally confluent, adnate, dry, felty,
cartridge-buff to pale pinkish buff, velvety, the margin thick,
entire; in section 500-1000 в, thick, not colored, at length zonate _

[Vol. 12
290 ANNALS OF THE MISSOURI BOTANICAL GARDEN

or stratose, composed of a layer next to the substratum of inter-

woven, tough, hyaline hyphae 3-4 џ in diameter, and of 1-4 hy-

menial layers; no gloeocystidia; cystidia not incrusted, cylindric,

80-150 х 6-9 u, protruding up to 80 и beyond the basidia, not

destroyed by potassium hydrate treatment; basidia with 4

sterigmata; spores hyaline, even, 12-15 x 4-6 y, copious.

Fruetifications 5 mm.—214 em. long, 3-10 mm. wide, rarely
5-10 ст. long by confluence.

On decorticated decaying wood and fence rails of Pinus albi-
caulis, P. contorta, P. flexilis, P. Murrayana, P. silvestris, Abies
grandis, Lariz, Pseudotsuga, and Thuja. In northern Europe,
and in Wyoming, Montana, Idaho, British Columbia, Wash-
ington, and Arizona. Frequent.

P. odorata may be recognized by its small, thick, pulvinate, dry,
velvety, pallid fructifications on old, weathered, blackened, coni-
ferous wood, by large spores, and stratose fructifications which
have even cystidia not affected by the potassium hydrate treat-
ment of sections. Karsten referred his specimens to Stereum
odoratum Fries, and Bresadola included the Karsten specimens
under his Stereum Karstenii Bres., of which I regard the type to
be a gathering made by Kmet in Hungary.

Specimens examined:

Finland: Mustiala, P. A. Karsten, type of Phanerochaete odorata.

Sweden: Bedaro, L. Romell, 412; Lappland, L. Romell, 413; Stock-
holm, L. Romell, 369.

Montana: Anaconda, J. R. Weir, 583 (in Mo. Bot. Gard. Herb.,
63173); Bernice, E. E. Hubert, comm. by J. R. Weir, 12011 (in
Mo. Bot. Gard. Herb., 63322); Hecla, E. E. Hubert, comm. by
J. R. Weir, 11405 (in Mo. Bot. Gard. Herb., 63260) ; Choteau,
J. A. Hughes, comm. by J. R. Weir, 5824 (in Mo. Bot. Gard.
Herb., 55649); Libby, E. E. Hubert, comm. by J. R. Weir,
11351 (in Mo. Bot. Gard. Herb., 63259); Melrose, E. E. Hu-
bert, comm. by J. R. Weir, 11427, 11433, 11439 (in Mo. Bot.
Gard. Herb., 63261, 63274, 63279); West Butte, J. A. Hughes,
comm. by J. R. Weir, 5496 (in Mo. Bot. Gard. Herb., 55647).

ies Fox Park, J. R. Weir, 10018 (in Mo. Bot. Gard. Herb.,

Idaho: Bonanza, G. G. Hedgcock, comm. by C. J. Humphrey,

1925]
URT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 291

2527, in part; Coolin, J. R. Weir, 11526 (in Mo. Bot. Gard.
Herb., 63291); Priest River, E. E. Hubert, comm. by J. R.
Weir (in Mo. Bot. Gard. Herb., 63258).

British Columbia: Kootenai Mts., Salmo, J. R. Weir, 536 (in
Mo. Bot. Gard. Herb., 22598).

Washington: Mt. Paddo, W. N. Suksdorf, ?29.

Arizona: Coronado National Forest, Santa Catalina Mountains,
G. G. Hedgcock & W. H. Long, comm. by C. J. Humphrey, 2544
(in Mo. Bot. Gard. Herb., 63534).

67. P. pilosa Burt, n. sp.

Type: in Burt Herb.

Fructifications effused, thin, closely adnate, hypochnoid, be-
coming pale olive-buff in the herbarium, the margin thinning out;
in section 40—60 џ thick, not colored, composed of loosely arranged,
thin-walled hyphae 214-3 и in diameter, not incrusted, and of
cystidia starting from the substratum; no gloeocystidia; cystidia
not incrusted, thin-walled, cylindric, 60-100 x 416-7 в, pro-
truding up to 70 џ beyond the basidia, often constricted near the
tip and terminating in an ovoid-shaped body; basidia 4-spored;
spores hyaline, even, 6-8 X 4-414 u, copious.

. Fructifications fragmentary, with the largest fragment 215 cm.
long, 1 em. wide.

On decaying coniferous wood. New York and Alabama. Prob-
ably rare.

P. pilosa forms a gray, downy covering on old weathered pine
wood, with the basidia not forming a compact hymenium. In
aspect this species somewhat resembles P. tenuis but there are
no gloeocystidia, and the numerous long, cylindric cystidia, some-
times terminating in a single spore-shaped end and sometimes in
a short row of 2 or 3, are distinctive.

Specimens examined:

New York: East Galway, E. A. Burt; Ithaca, G. F. Atkinson,

14416, type.

Alabama: Montgomery, R. P. Burke, 154 (in Mo. Bot. Gard.

Herb., 3650).

68. Р. Peckii Burt, n.
Type: in Burt Herb. ied probably i in N. Y. State Mus. Herb.

[Vol. 12
292 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fructifications broadly effused, somewhat membranaceous,
separable from the substratum in small portions when moistened,
thin, becoming cartridge-buff to cream-buff in the herbarium, not
shining, eracking in drying, the margin thinning out ; in section
60-360 » thick, not colored, composed throughout of suberect
hyphae about 4. in diameter, not incrusted and occasionally
nodose-septate, of elongated flexuous cystidia, and of great
numbers of subglobose, even chlamydospores 415-5 X 4u;
cystidia not incrusted, flexuous, elongated, with somewhat the
aspect of gloeocystidia, 30-100 х 6-9 џ, in all regions of fruc-
tification, many starting from the substratum, tapering upward,
protruding up to 40 » beyond the База; basidia with 4 sterig-
mata; basidiospores hyaline, even, subglobose, 5-6 x 41% u.

Fructifications 2-6 em. long, 1-3 em. wide.

On bare ground in woods and on bark and wood of decaying
Alnus, Betula, Populus, Quercus, and Ceanothus, rarely on a
coniferous substratum. Canada to Massachusetts and westward
to Washington. July to March. Occasional.

P. Peckii is placed in the species group with P. glebulosa on
account of the large, even-walled cystidia which are more flexuous
than those of the latter species and with more the aspect of gloeo-
cystidia, but I have not yet demonstrated by granular contents
that they are gloeocystidia. P. Peckii is distinguished by the
great number of subglobose spores distributed throughout the
whole fructification in sections studied.

Specimens examined:

Canada: J. Macoun, 1 8, 51; Lower St. Lawrence Valley, J.
Macoun, 5.

Massachusetts: Cherry Brook, E. A. Burt & A. B. Seymour;
Magnolia, W. G. Farlow (in Burt Herb. and Mo. Bot. Gard.
Herb., 44066); Sharon, A. P. D. Piguet, comm. by W. G. Far-
low, 11 (in Mo. Bot. Gard. Herb., 55590) and by Farlow Herb.,
132 (in Mo. Bot. Gard. Herb., 59622); Wellesley, L. W.
Riddle, 11.

New York: Ithaca, G. F. Atkinson, 5089; Karner, H. D. House,
comm. by N. Y. State Mus. Herb. (in Mo. Bot. Gard. Herb.,
94361, 55204); Westport, C. H. Peck, 6, type; White Plains,
W. H. Ballou, 1 (in Mo. Bot. Gard. Herb., 55030).

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 293

Michigan: Marquette, C. J. Humphrey, 1870 (in Mo. Bot. Gard.
Herb., 11089).

Wisconsin: Blue Mounds, Е. T. & S. A. Harper, 895.

Washington: Bingen, W. N. Suksdorf, 741, 907.

69. Р. heterocystidia Burt, n. sp.

Type: in Burt Herb.

Fructifications broadly effused, separable from the substratum
when moistened, becoming cracked in drying and often loosening
from substratum along the fissures, whitish when young, becoming
light drab, cinnamon drab or vinaceous drab, the margin often
paler; in section not colored or with only the hymenial layer
clay-colored or brownish, 200—400 » thick, 2-layered, the layer
next to substratum usually broad, composed of loosely interwoven,
somewhat ascending or longitudinally arranged, hyaline, nodose-
septate hyphae 3-414 џ in diameter, the hymenial layer 40-80 y.
thick, composed of cystidia, gloeocystidia, and erect hyphae
usually slightly colored near plane of origin from the under layer;
cystidia consisting of both usual incrusted cystidia 25-35 X
6-8 в, distributed in all parts of the outer layer, and of very large
cystidia up to 40-100 х 20-50» which start from the base—
often somewhat colored—of the hymenial layer; gloeocystidia
slender, flexuous, 40-60 х 5-6 u, between the basidia; basidia
with 4 sterigmata; spores from spore collection white, even,
cylindric, 12-15 x 3144-4% y.

Fructifications 2-7 cm. in diameter.

On fallen limbs of gray birch, beech, maple, Carpinus, Mag-
nolia, and other frondose species. Canada to Mississippi and
westward to Missouri and in Mexico. June to March. Com-
mon.

P. heterocystidia resembles Corticium laeve Pers.(=C. evolvens
Fr.) in color but is a true Peniophora, readily distinguished from
our other separable species by having incrusted cystidia of the
usual size, other very large cystidia up to 20-50 y. in diameter,
and gloeocystidia. Bresadola and von Hóhnel & Litschauer con-
fused this species with P. carnea, from which it differs in being
much thicker, separable from the substratum when moistened,
and being colored within on the hymenial side instead of next

[Vol. 12
294 ANNALS OF THE MISSOURI BOTANICAL GARDEN

to the substratum. Rarely a fructification may have the

hymenium vary somewhat hydnaceous.
Specimens examined:

Exsiccati: Ellis, N. Am. Fungi, 716, under the name Corticium
glabrum, 717a of some copies, under the name Corticium sub-
giganteum.

Canada: J. Macoun, 8, 10, 14, 46.

Ontario: Lake Rosseau, E. T. & S. A. Harper, 755; Ottawa, J.
Macoun, 110.

Vermont: Middlebury, E. A. Burt, type, and 2 other gatherings.

Connecticut: Central Village, J. L. Sheldon, 23, comm. by N. Y.
Bot. Gard. Herb.

New York: Bemis Heights, C. H. Peck (n N. Y. State Mus.
Herb., under the name Stereum albobadium); Bronx Park,
Class in Mycology (in N. Y. Bot. Gard. Herb., Mo. Bot. Gard.
Herb., 61392, 61430, and Burt Herb.); Kirkland, H. D. House
(in N. Y. State Mus. Herb., Mo. Bot. Gard. Herb., 59685,
and Burt Herb.); Snyders, C. H. Peck (in N. Y. State Mus.
Herb., and Mo. Bot. Gard. Herb., 56018); Syracuse, L. M.
Underwood, in some copies of Ell. & Ev., Fungi Col., 221, under
the name Corticium glabrum; White Plains, W. H. Ballou (in
Mo. Bot. Gard. Herb., 55034).

New Jersey: Newfield, J. B. Ellis.

Pennsylvania: Meadville, E. C. Smith, comm. by L. O. Over-
holts, 8337 (in Mo. Bot. Gard. Herb., 59475); West Chester,
Everhart, Haines, Jefferis & Gray, in Ellis, N. Am. Fungi, 716.

Distriet of Columbia: Washington, C. L. Shear, 1257a, 1260.

Mississippi: Ocean Springs, L. M. Underwood (n N. Y. Bot.
Gard. Herb., and Mo. Bot. Gard. Herb., 61484).

Ohio: Cincinnati, C. G. Lloyd, 191 ; 2790, 4518; Norwood, C. G.
Lloyd, 2274. ,

Indiana: Millers, E. T. & S. A. Harper, 962; Union County,
M. F. & L. O. Overholts & B. Fink, comm. by L. O. Overholts,
4204 (in Mo. Bot. Gard. Herb., 55636).

Illinois: Cypress, C. J. Н. umphrey, 1347 (in Mo. Bot. Gard. Herb.,
42923); Glencoe, E. T. & S. A. Harper, 662, 646.

Kentucky: Crittenden, C. G. Lloyd (in Lloyd Herb., 1411, and
Mo. Bot. Gard. Herb., 55626).

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 295

Missouri: Columbia, B. M. Duggar, 265, 288, 400, 472; Pacific,
B. M. Duggar (in Mo. Bot. Gard. Herb., 63417); near St.
Louis, E. A. Burt (in Mo. Bot. Gard. Herb., 63418).

Mexico: Tepeite Valley near Guernavaca, W. A. & E. L. Mur-
rill, 400, comm. by N. Y. Bot. Gard. Herb. (in Mo. Bot. Gard.
Herb., 54551).

70. P. borealis (Peck) Burt, n. sp.

Peniophora disciformis (DC) Cooke var. borealis Peck in
Harriman Alaska Exped. 5. The Fungi of Alaska, 43. 1904;
Басс. Syll. Fung. 17: 175. 1905.

Type: in Mo. Bot. Gard. Herb.

Fructifications effused, thick, membranaceous, separable, be-
coming light buff in the herbarium, velvety with the numerous
cystidia, the margin thinner, entire, clay-color, free in some places;
in structure 60 џ thick, not colored, composed of hyaline hyphae
2u in diameter, not incrusted, densely and longitudinally ar-
ranged along the substratum and then curving obliquely out-
ward to form the hymenial layer, and of occasional slender gloeo-
cystidial organs with enlarged clavate or pyriform tips up to
414-7 u in diameter; cystidia incrusted, cylindric, 60-75 x 6-
9 u, confined to the hymenial surface but in great numbers there,
protruding nearly their whole length beyond the basidia; a single
detached spore is hyaline, even, 10 X 8 џ, but may be foreign.

Fructifications 5 тт. 216 cm. long, 5 mm.-10 mm. wide.

On bark of small decaying twigs of a frondose species—perhaps
Alnus. Alaska. June.

P. borealis has aspect somewhat suggestive of P. aurantiaca but
is more buff-colored, with darker margin becoming free, and with
cystidia so long and numerous as to be very conspicuous when
viewed with a lens. The abundance of these cystidia is so great
as to be a very important character in the recognition of this
species by preliminary BED po

Specimens examin
Alaska: Aqua Dulce River, Yakutat Bay, W. Trelease, 583, type

(in Mo. Bot. Gard. Herb., 5006).

71. P.lepida Bresadola, Mycologia 17: 70. 1925.
Type: in Weir Herb

[Vol. 12
296 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fructifications broadly effused, thick, waxy-membranaceous,
separable from the substratum, somewhat horn-like and requiring
moistening for a short time before sectioning, pinkish buff to
light ochraceous-buff in the herbarium, somewhat pulverulent,
the margin finally free and rolling up from the substratum; in
section 500-600 в. thick, not colored, composed of densely ar-
ranged hyphae 3-31% џ in diameter, which run longitudinally
along the substratum and then curve obliquely into the hymeni-
um; between the hyphae occur numerous slightly more deeply
staining elongated organs of the nature of conducting organs or
slender gloeocystidia; cystidia incrusted, cylindric, 6-8 y. in diam-
eter, protruding ир to 30 џ beyond the basidia, very numerous
in the hymenial surface, the incrusted part about 20-45 y long;
no spores found.

Fructification 9 em. long, 314 cm. wide.

On a dead stub about 214 em. in diameter, of Saliz sp. Idaho.
June.

P. lepida has some resemblance to P. gigantea but is not quite
as gelatinous in consistency as P. gigantea and occurs on Saliz.
The slender conducting hyphae or gloeocystidia should aid in
recognizing the species. The broad layer of hyphae arranged
longitudinally along the substratum and then curving outward
into the hymenium is very like that of a resupinate Stereum but
I recall no pileate Stereum of similar structure.

Specimens examined:

Idaho: National Forest, 50 miles east of Orofino, A. S. Rhoads (in

Weir Herb., 16744, type).

72. P. Kauffmanii Burt, n

Type: in Mo. Bot. Gard. Нењ. and probably in Kauffman
Herb.

Fructifications long-effused, rather thin, adnate, small portions
separable when moistened, between pale pinkish buff and tilleul-
buff, not cracked, not waxy nor shining, the margin determinate,
thinning out; in section 300-350 в. thick, 2-layered,. with both
layers of abont equal thickness and the hymenial layer somewhat
honey-yellow, the layer next to the substratum not colored, com-
posed of loosely and longitudinally interwoven, thin-walled,

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 297

hyaline hyphae about 3 р in diameter, of irregular outline; hy-
menial layer composed of densely arranged, erect hyphae, gloeo-
cystidia, and cystidia; gloeocystidia flexuous, 45-100 X 4-7 p;
eystidia incrusted when wholly immersed, cylindric, obtuse,
30-45 x 6-8 в, protruding 20-30 и beyond the База in in-
erusted, or more usually, non-inerusted form, not abundant;
_ spores hyaline, even, curved, 8-12 x 214-3 y, with pointed and
tapering base, copious.

Fructifications 2-10 em. long but broken off at one end, 1-214
em. wide.

On decaying limbs of Fagus. Kentucky. September. Prob-
ably local.

Among our few species of Peniophora which have gloeocystidia,
P. Kauffmanii should be readily recognized by its occurrence on
beech, buff color, structure of 2 equal layers, and small, incrusted
cystidia.

Specimens examined:

Kentucky: Harlan, C. H. Kauffman, 69, type (in Mo. Bot. Gard.

Herb., 22827).

73. Р. alba Burt, n. sp.

Type: in Burt Herb.

Fructifications effused, very thin, somewhat membranaceous,
small pieces separable when moistened, white, even, not shining,
somewhat cracked by contraction in drying, the margin t inni
out; in section 80-100 и thick, not colored, with the hyphae
loosely arranged near the substratum, suberect, branching, about
Зи in diameter, not incrusted, only rarely nodose-septate; gloeo-
cystidia curved, 30-45 x 314-414 u, usually starting from the
substratum; cystidia not incrusted or with some incrusting
granules, thin-walled, 4-5 р in diameter, protruding up to 30 y
beyond the basidia; spores hyaline, even, 4-5 X 21% у.

Fructifications fragmentary and not showing ends nor more
than one side; such fragments 5 cm. long, 10-15 mm. wide...

On bark of dead cedar or spruce. Canada. September.

P. alba seems possible of recognition among our many whitish
species of Peniophora by its pure white color, presence of gloeo-
cystidia in addition to cystidia, and occurrence on coniferous bark.

[Vol. 12
208 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Specimens examined:
Canada: locality not given, J. Macoun, 57, type.

74. P.tenella Burt, n

Type: in Mo. Bot. Gard. "Herb. and Farlow Herb.

Fructifications effused, white, tender, thin, loosely attached,
separable when moistened, velvety, setulose with the large
cystidia, the margin indeterminate, thinning out; in section
150-200 u thick, not colored, composed of a denne hymenial
layer 75-90 u thick, borne on a loosely interwoven layer composed
of thin-walled, hyaline hyphae 3-4 џ in diameter, nodose-septate,

granule ted; hymenial layer ма of basidia,
gloeocystidia, and incrusted cystidia; gloeocystidia numerous,
flexuous, tapering from the base, 45-75 x 5-8 u; cystidia very
large, heavily incrusted, conical, 60-100 х 15-20 в, wholly im-
mersed, or protruding beyond the База up to 75 в; spores
copious, hyaline, even, 744-9 х 3-4 y.

Fructifications 1-2 cm. in diameter.

On coniferous bark. New Hampshire and Massachusetts.
September and October. Rare.

P. tenella is distinguished from P. pubera by occurrence on
coniferous, rather than frondose, substratum, by being so loosely
attached to the substratum that small portions needed for sec-
tioning may be separated from the substratum when moistened,
and by the loosely interwoven hyphal layer equalling or exceeding
in thickness the hymenial layer and containing no gloeocystidia
nor cystidia.

Specimens examined:

New Hampshire: Chocorua, W. G. Farlow, type (in Mo. Bot.

Gard. Herb., 7617).

Massachusetts: Cambridge, A. P. D. Piguet, comm. by W. G.

Farlow, 30.

75. P. duplex Burt, n. sp.

Type: in Burt Herb.

Fructifications small, effused, thin, adnate, somewhat mem-
branaceous, small pieces separable when moistened, becoming pale
pinkish buff in the herbarium, even, not cracked, not shining, the

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 299

margin narrow, radiate-fibrillose; in section 100-200 џ thick,
not colored, the hyphae with walls gelatinously modified, in-
distinct, about З и in diameter, ascending, densely crowded to-
gether and interwoven and with numerous gloeocystidia present;
gloeocystidia sometimes pyriform but usually more elongated,
20-45 x 6-9 и, in all regions of the fructification; cystidia in-
crusted, cylindric, 25-30 x 416-5 и, protruding up to 20,
beyond the basidia, confined to the hymenium; spores hyaline,
even, flattened on one side, 5 X 215 y.

Fructifications received in fragments 1-2 cm. long, 5-10 mm.
wide.

On bark of Pinus austriaca (cult.. New York. October.

In general aspect P. duplex suggests small fructifications of
P. gigantea but not curling away from the substratum at all.
The cystidia are smaller than those of P. gigantea and the latter
does not have gloeocystidia.

Specimens examined:

New York: Shelter Island, W. G. Farlow, type.

76. P. mutata (Peck) Bresadola in Bourdot & Galzin, Soc.
Myc. Fr. Bul. 28: 399. 1913.

Corticium mutatum Peck, N. Y. State Mus. Rept. 43:67. 1890.

Type: in N. Y. State Mus. Herb.

Fructifications broadly effused, membranaceous, fleshy, thick,
separable when moistened, drying white to pinkish buff, some-
times centrally tubereulose or with raduloid teeth and occasion-
ally with radial folds, sometimes cracking in drying and showing
the white, fibrillose subiculum in the fissures, the margin white,
radially byssoid; in structure 300-1000 u thick, composed of
loosely arranged, ascending, thin-walled, hyaline hyphae 3-4 в
in diameter, occasionally nodose-septate; gloeocystidia pyriform,
15 X 7 y, more or less numerous, sometimes grown out into elon-
gated, flexuous form up to 100 X 4-5 p, occurring as hyphal
ends or branches in the sybhymenium; cystidia incrusted or not
incrusted, 50-100 x 6-15 u, sometimes not protruding beyond
the basidia and sometimes so few present as to be found only
after examination of several sections; basidia 4-spored, with
short, thick, knob-like sterigmata; spores hyaline, even, cylindrie,
8-16 x 3-4 y.

[Vol. 12
300 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Fructifications 3-7 em. long, 1-3 em. broad, sometimes larger
by confluence.

Common on bark of decaying logs and fallen branches of
Populus and also on Tilia, Quercus, Acer, and other frondose
species. Canada to Alabama, westward to Idaho, in Europe and
in Japan. July to November and in April.

P. mutata is a thick, somewhat fleshy, white, or whitish species
occurring usually on bark of fallen poplar and basswood and
showing in sectional preparations pyriform gloeocystidia, cystidia,
and spores usually about 12 x 316 в. This species approaches
the genus Radulum in thickness of the fructification, its obliquely
ascending hyphae, and in occasional specimens having some
raduloid teeth; such specimens have the aspect of Radulum
orbiculare but differ from it by presence of the pyriform bodies
and cystidia.

Specimens examined:

Exsiccati: Ellis, N. Am. Fungi, 717 a, under the name Corticium
subgiganteum, 719, under the name Corticium laeve; Ell. & Ev.,
Fungi Col., 308, under the name Corticium laeve.

Austria: Langenschónbich, F. v. H” hnel, comm. by V. Litschauer.

Canada: J. Macoun, 7, 52; Lower St. Lawrence Valley, J.
Macoun, 11, 81; Ottawa, J. Macoun, 40, 41, and 136 (in N. Y.
State Mus. Herb., and Mo. Bot. Gard. Herb., 55924) ; Ontario,
Harraby, E. T. & S. A. Harper, 685.

Maine: Piscataquis County, W. A. Murrill, 2451 (in N. Y. Bot.
Gard. Herb., and Mo. Bot. Gard. Herb., 61402); Portage,
L. W. Riddle, 16.

Vermont: Middlebury, E. A. Burt, two gatherings.

New York: Ithaca, G. F. Atkinson, 22965; Karner, H. D. House,
14.158, 14.160 (in part), and two unnumbered specimens (in
Mo. Bot. Gard. Herb., 44712, 44706, 54367, 55215); Sevey,
C. H. Peck, type (in N. Y. State Mus. Herb.); Shokan, C. H.
Peck, T24 (in Mo. Bot. Gard Herb., 54660); Slingerlands,
C. H. Peck, T 23 (in Mo. Bot. Gard. Herb., 54659) ; Syracuse,
L. M. Underwood, 48 (in N. Y. Bot. Gard. Herb., and Mo.
Bot. Gard. Herb., 61409, 61433); White Plains, W. H. Ballou
(in Mo. Bot. Gard. Herb., 10458).

New Jersey: Newfield, J. B. Ellis, in Ellis, N. Am. Fungi, 7178,
719, and Ell. & Ev., Fungi Col., 308.

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 301

Florida: W. W. Calkins.

Alabama: Montgomery, R. P. Burke, 13 (in Mo. Bot. Gard.
Herb., 17193).

Ohio: Preston, C. G. Lloyd, 1655; West Elkton, L. O. Overholts,
3164 (in Mo. Bot. Gard. Herb., 5712).

Indiana: Crawfordsville, D. Reddick, 6, 8, 18, 14.

Illinois: River Forest, E. T. & S. A. Harper, 631.

Michigan: Ann Arbor, C. H. Kauffman.

Wisconsin: Blue Mounds, E. T. & S. A. Harper, 871.

Minnesota: Princeton, C. J. Humphrey, 899 (in Mo. Bot. Gard.
Herb., 21044).

Missouri: Columbia, B. M. Duggar, 561; Meramec Highlands,
P. Spaulding (in Mo. Bot. Gard. Herb., 63746); Pickering,
E. Bartholomew, 6425 (in Mo. Bot. Gard. Herb., 55195);
St. Louis, N. M. Glatfelter, 1377, comm. by N. Y. Bot. Gard.
Herb.; E. A. Burt (in Mo. Bot. Gard. Herb., 44073).

South Dakota: Custer, J. R. Weir, 10019 (in Mo. Bot. Gard.
Herb., 55799).

Idaho: Priest River, J. R. Weir, 37; St. Maries, J. R. Weir, 560
(in Mo. Bot. Gard. Herb., 63179).

Japan: Bungo, A. Yasuda, 107 (in Mo. Bot. Gard. Herb.,
57024).

77. P. Allescheri Bresadola, Fungi Trid. 2: 62. pl. 172.
1898; Sace. Syll. Fung. 16: 194. 1902.—But not as understood
by Bourdot & Galzin and by Wakefield.

Kneiffia Allescheri Bresadola, Ann. Мус. 1: 100. 1903.—
Gloeopeniophora Allescheri (Втез.) v. Hóhnel & Litschauer, K.
Akad. Wiss. Wien Sitzungsber. 117: 1082. 1908.

Type: in Bresadola Herb. and Burt Herb.

Fructifications broadly effused, membranaceous, fleshy, thick,
drying white to pinkish buff, sometimes contracting in drying,
eurling away from the substratum more or less at the fissures
and showing the white, fibrillose subiculum, the margin white,
byssoid; in structure 300-1000 u thick, composed of obliquely
ascending and interwoven, hyaline hyphae more or less incrusted,
3-6 и in diameter; gloeocystidia elongated, flexuous, 40-100 x
4-7 р, often continued beyond the deeply staining portion as an

[Vol. 12
302 ANNALS OF THE MISSOURI BOTANICAL GARDEN

undifferentiated hypha, numerous in the subhymenium; cystidia
incrusted or not incrusted, 30-60 х 6-10 u; spores hyaline, even,
10-13 х 3-4 y.

Fructifications 2-10 сш. long, 1-3 em. broad.

On bark of fallen limbs of Populus and other frondose species.
Canada to New York and westward to Washington, in West
Indies, and in Europe.

The type of P. Allescheri and specimens of similar structure
cited below differ so slightly from P. mutata that I have separated
them from the latter only by all their gloeocystidia being of
slender elongated form and perhaps specially differentiated
middle portions of hyphae, while the gloeocystidia of P. mutata
are terminal portions of hyphae and hyphal branches which are
in many cases pyriform and in others afford indication by a pyri-
form base of having finally grown out from a pyriform body into
an elongated gloeocystidium. It may be that when someone
can keep under observation and examination a specimen of
P. mutata during its season, he may find that pyriform gloeo-
cystidia are present abundantly up to the time of copious spore
production and then finally all become elongated so that the
fructification would be referable to P. Allescheri. In this event
P. Allescheri will become a synonym of P. mutata by priority
of the latter.

While P. mutata has become correctly understood in Europe
through my exchanges with Bresadola there is a misunderstanding
there concerning P. Allescheri. Von Нӧһпе! & Litschauer
studied the original specimen of P. Allescheri in Bresadola Herb.
and state, loc. cit., that this consists of a mixture of fructifications
of P. cremea and P. Allescheri, the latter as described by Bresadola
and figured in his plate. The specimen shared with me by Bresa-
dola is in such close agreement with the plate that the colored
drawing of the upper figure may have been made from it, and it
agrees also with the description. Its data as to collection is given
“аа corticem Fagi silv. Bavaria. Allescher.” The portions of the
original specimens communicated to Bourdot & Galzin and to
Miss Wakefield are apparently of the P. cremea component,
referred to by v. Hóhnel & Litschauer.

Specimens examined:

1925]
URT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 303

Sweden: L. Romell, 439 (in Mo. Bot. Gard. Herb., 44305) ; Stock-
holm, L. Romell, 102.

Germany: Bavaria, Allescher, comm. by Bresadola, part of type.

Canada: Rideau Park, J. Macoun, 825; Ottawa, J. Macoun, 138,
in part; Quebec, Ironsides, J. Macoun, 256.

New Hampshire: Jackson, W. H. Snell, 624 (in Mo. Bot. Gard.
Herb., 59246).

Vermont: Middlebury, Е. A. Burt.

New York: Glasco, Ulster County, P. Wilson, 40 (in Mo. Bot.
Gard. Herb., 54747); Ithaca, H. E. Stork, 6 (in Mo. Bot. Gard.
Herb., 56643), G. F. Atkinson, 8031, 22761, D. Reddick, by
Cornell Univ. Herb., 20567, Van Hook, Cornell Univ. Herb.,
8011, and Wright, Cornell Univ. Herb., 8353.

Ohio: C. G. Lloyd, 3920; College Hill, C. G. Lloyd, 3121 (in Lloyd
Herb., Burt Herb., N. Y. Bot. Gard. Herb., and Mo. Bot.
Gard. Herb., 61419).

Indiana: Crawfordsville, A. R. Bechtel, 10 (in Mo. Bov. Gard.
Herb., 59648).

Michigan: Ann Arbor, C. H. Kauffman, 12; New Richmond,
C. H. Kauffman, 31 (in Mo. Bot. Gard. Herb., 9864).

British Columbia: Vancouver Island, J. Macoun, 355 (in Mo.
Bot. Gard. Herb., 55323).

Washington: Bingen, W. N. Suksdorf, 702. `

West Indies: Grenada, Grand Etang, R. T'haxter, comm. by W. С.
Farlow, 4.

78. P. subcremea v. Hóhnel & Litschauer, К. Akad. Wiss.
Wien Sitzungsber. 115: 1600. text f. 6. 1906; Басе. Syll. Fung. ©

: type ibution in Rabenhorst, Fungi Eur., 3230, under
the name Corticium lacteum.

Fructifications effused, thin, closely adnate, becoming cartridge-
buff to ivory-yellow in the herbarium, not cracked, the margin
thinning out; in section 40-150 y thick, not colored, composed of
suberect, bushy-branched hyphae 2-3 in diameter, not in-
crusted, only occasionally nodose-septate, and of flexuous gloeo-
cystidia 40 x 4u, starting from the substratum in the type;
cystidia not incrusted, 414-6 y in diameter, protruding up to

[Vol. 12
304 ANNALS OF THE MISSOURI BOTANICAL GARDEN

40 u beyond the basidia; spores hyaline, cylindric, 315—416 x
2-214 u, copious.

Fructifications 114-8 em. long, 114-5 em. wide.

On bark and wood of Pinus. Finland, Montana, and Manitoba.
September to November. Rare.

The specimen from Manitoba is on bark of a frondose species,
but agrees well in other respects with the specimens on pine.
The small spores are a distinguishing character of P. sub-
cremea.

Specimens examined:

Exsiccati: Rabenhorst, Fungi Eur., 3230, type distribution, under
the name Corticium lacteum.
Finland: Mustiala, P. A. Karsten, in Rabenhorst, Fungi Eur.,

3230.

Montana: Anaconda, E. E. Hubert, comm. by J. R. Weir, 12010

(in Mo. Bot. Gard. Herb., 63370).

Manitoba: Winnipeg, G. R. Bisby & I. L. Conners, 1183 (in Mo.

Bot. Gard. Herb., 59047).

79. P. admirabilis Burt, n. sp.

Type: in Burt Herb.

Fructifications broadly effused, adnate, thin, membranaceous,
small pieces separable, becoming cartridge-buff in the herbarium,
fibrillose, not shining, even, with but few small cracks, the margin
thinning out, with its hyphae loosely interwoven; in section
180-240 y. thick, not colored, composed of suberect, thin-walled
hyphae 3142-4 y. in diameter, not incrusted, of gloeocystidia both
elongated and vesicular, and of large chlamydospores; cystidia
cylindric, incrusted, up to 105 X 9 y, confined to the hymenium,
somewhat disorganized by potassium hydrate solution; vesicular
gloeocystidia and vesicular spaces up to 45 х 30 u; chlamydo-
spores as seen singly on hyphal branches are up to 15 X 9 y;
basidiospores white in spore collection, even, 6-7 x 3 y, borne
4 to a basidium.

Fructifications 3-10 em. long, 1-2 em. wide.

On decaying wood of stump of Ulmus. New York. May.

P. admirabilis is well marked among our species which have
gloeocystidia by the presence of large imbedded spores.

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 305

Specimens examined:
New York: Oneonta, E. A. Burt.

80. P. versata Burt, n. sp.

Type: in Burt Herb.

Fruetifieations effused, thin, membranaceous, small pieces
separable when moistened, becoming pinkish buff in the her-
barium, not waxy, the margin thinning out, fibrillose; in section
150-300 y. thick, not colored, composed of suberect, interwoven
hyphae about 3 џ in diameter, with walls somewhat gelatinously
modified, and of gloeocystidia; gloeocystidia numerous, flexuous,
35-55 x 6-8 u; cystidia not incrusted, tapering towards the
apex, 6 џ in diameter, protruding up to 40 y, scarcely or perhaps
not at all distinguishable from gloeocystidia; basidia with 4
sterigmata; spores hyaline, even, 4-5 x 214-3 y.

Fructifications 2-5 mm. long, 5-15 mm. wide.

On slightly decayed red fir planks and cross-ties. Washington.
September and October.

The fructifications of P. versata stand out conspicuously against
the blackened timber upon which they occur. The resemblance
of the protruding cystidia to gloeocystidia and the possibility
that they have the function of gloeocystidia or may be gloeo-
cystidia functioning as cystidia should enable this species to be
readily distinguished from species containing cystidia and gloeo-
cystidia quite distinct from each other.

Specimens examined:

Washington: Chehalis, C. J. Humphrey, 6285; Edmonds, C. J.

Humphrey, 7623, type.

81. P. albo-straminea Bresadola, Mycologia 17: 69. 1925.

Type: in Weir Herb.

Fructifications orbicular, finally confluent and broadly effused,
thin, very tender, small pieces separable when moistened, be-
coming between cartridge-buff and ivory-yellow i in the herbarium,
somewhat cracked, even, the margin pruinate; in section 60-90 u
thick, not colored, composed of somewhat loosely arranged
hyphae 3-5» in diameter, occasionally nodose-septate, and of
gloeocystidia; gloeocystidia flexuous to zigzag, 40-60 x 444-6 u,

[Vol. 12
306 ANNALS OF THE MISSOURI BOTANICAL GARDEN

usually starting from the substratum and wholly immersed,
sometimes protruding beyond the База; ¢ystidia, if really
distinct from gloeocystidia, not incrusted, 414-8 џ in diameter,
protruding up to 30 џ beyond the basidia; spores hyaline, even,
5-71 х 3-4 y.

Fructifications 2-8 em. long, 112-3 ст. wide.

On wood and bark of decaying Alnus tenuifolia and Quercus
californica. Idaho and California. October.

The gloeocystidia, sometimes of zigzag form, and cystidia,
which are possibly only protruding portions of gloeocystidia, are
marked characters of the type specimen which should afford
recognition of P. albo-straminea if these are constant specific
characters. However, the buff color, Alnus substratum, and
presence of gloeocystidia should suffice. In No. 17069 a foreign
mycelium of coarse hyphae is underneath the fructification
proper.

Specimens examined:

Idaho: Priest River, J. R. Weir, 17069, type, and 16818 (both in

Weir Herb.).

California: Massack, Plumas National Forest, A. S. Rhoads, 17

(in Mo. Bot. Gard. Herb., 56986).

82. P. Taxodii Burt, n. sp.

Type: in Burt Herb.

Fructifications effused, very thin, closely adnate, whitish to
pale olive-buff in the herbarium, the hymenium loose and rather
hypochnoid under a lens, the margin indeterminate, thinning
out; in section 45-60 y. thick, not colored, composed of erect,
branching hyphae 2-21 u in diameter, not incrusted, not nodose-
septate, and of numerous cystidia, gloeocystidia, and crystalline
matter; cystidia not incrusted, thin-walled, tapering to a sharp
apex, 6-8 џ in diameter, protruding 20—40 y. beyond the База,
often starting from the substratum; gloeocystidia often not dis-
tinguishable from the cystidia except by granular, deeply staining
contents, protruding up to 20-40 u beyond the База; spores
hyaline, even, 7-714 x 3-314 y.

Fructification 7 em. long, 115-2 em. wide.

On decorticated top limb of prostrate top of Тахофит dis-

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 307

tichum left in the swampy woods. Texas. September. Prob-
ably local.

P. Taxodi has thin grayish fructifications on the blackened,
weathered wood of a prostrate tree top left in lumbering opera-
tions. It is difficult to distinguish cystidia from gloeocystidia in
the sections unless the organs lacking contents which take the
stain are cystidia and the deep-staining and more numerous
bodies gloeocystidia, for both start from the substratum, protrude
beyond the hymenium, and taper to a sharp point.

Specimens examined:

Texas: Beaumont, C. J. Humphrey, 5947, type.

83. Р. investiens Burt, п. sp.

Type: in N. Y. Bot. Gard. Herb., Mo. Bot. Gard. Herb., and
Burt Herb.

Fructifications longitudinally effused, thin, adnate, small
pieces separable when moistened, becoming cartridge-buff in the
herbarium, even, not cracked, not shining, the margin thinning
out, minutely tomentose; in section 180 y thick, not colored, with
a broad layer of densely interwoven hyphae 214-3 џ in diameter,
thin-walled, not incrusted, not nodose-septate; gloeocystidia
flexuous, 25-40 x 4—5 р, immersed in the hymenium; cystidia
not incrusted, 9 џ in diameter, protruding up to 60u beyond
the basidia; basidia with 4 prominent sterigmata; spores hyaline,
even, 12-13 x 3-314 u—one spore seen is 15 X 6 y but per-
haps does not belong.

Fructifications 8 cm. long, 1 cm. wide.

On decaying stem of palmetto. Bermuda. December.

The presence of the gloeocystidia in the hymenial layer and
not also in the interwoven hyphae near the trama, together with
the long spores and occurrence on palmetto, should enable the
recognition of P. investiens.

Specimens examined:

Bermuda: Stewardson Brown, N. L. Britton & F. J. Seaver, 1324,
type (in N. Y. Bot. Gard. Herb., Mo. Bot. Gard. Herb.,

63730, and Burt Herb.).

84, P. incarnata (Pers. Karsten, Hedwigia 1889: 27. F.
1889; Finska Vet.-Soc. Bidrag Natur och Folk 48: 424, 1889;

[Vol. 12
308 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Massee, Linn. Soc. Bot. Jour. 25: 147. Jl. 1889; Sacc. Syll.
Fung. 9: 241. 1891; Bourdot & Galzin, Soc. Муе. Fr. Bul.
28: 404. 1913; Rea, Brit. Ваза. 694. 1922.

Thelephora incarnata Persoon, Syn. Fung. 573. 1801 ; Myce.
Eur. 1:130. 1822 (Corticium); Fries, Syst. Myc. 1: 444, 1821;
Elenchus Fung. 1: 219. 1828.—Corticium incarnatum (Pers.)
Fries, Epicr. 564. 1838; Hym. Eur. 654. 1874 ; Berkeley,
Brit. Fung. 275. 1860; Berk. & Curtis, Grevillea 2: 4. 1873 ў
Peck, №. Y. State Mus. Rept. 24: 80. 1872 ; Morgan, Cin-
cinnati Soc. Nat. Hist. Jour. 10: 201. 1888; Sacc. Syll. Fung. 6:
625. 1888.—Kneiffia incarnata (Pers.) Bresadola, Ann. Myc. 1:
103. 1903.—Gloeopeniophora incarnata (Pers. v. Hóhnel &
Litschauer, K. Akad. Wiss. Wien Sitzungsber. 116:816. 1907.—
Peniophora aemulans Karsten, Finska Vet-Soc. Bidrag Natur och
Folk 48: 425. 1889; Sacc. Syll. Fung. 9: 239. 1891.

Fructifications effused, closely adnate, drying pinkish cinnamon
to warm buff, eracking, the margin sometimes paler, thinning
out; in section 100-250 џ thick, not colored, composed of hyaline,
thin-walled hyphae 2-3 y in diameter, densely interwoven along
the substratum and then becoming suberect and extending be-
tween more or less numerous gloeocystidia and some cystidia;
gloeocystidia sometimes broadly ovoid, 30-45 x 10-15 в, usu-
ally more cylindrie and narrower, 30-60 x 6-10 ш; cystidia be-
coming incrusted, 30-45 x 6-10 y, rarely protruding beyond
the basidia; basidia with 4 sterigmata; spores hyaline, even,
cylindrie, flattened on one side, 6-10 x 3-414 u.

Fructifications 2-10 cm. long, 1-2 cm. broad, sometimes in
scattered, small, tubercular growths 2-5 mm. in diameter on
lenticels of small limbs.

On wood and bark of fallen limbs of frondose species usually.
Europe, Canada to Alabama, and westward to the Pacific states,
and in Japan. Throughout the year. Common.

P. incarnata is recognizable by its closely adnate, reddish
fructifications, spores 6-9 x 3-4 и, and abundant gloeocystidia.
Sometimes one has to search several sections before finding an
incrusted cystidium. The spores run slightly smaller in most
American gatherings than in the fewer European specimens which
I have seen and are with us usually only about 6-8 x 3 y.

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. ХІУ 309

Specimens examined:

Exsiccati: Cooke, Fungi Brit, 7; Reliq. Farlowianae, 342;
Ravenel, Fungi Am., 140; Romell, Fungi Scand., 33

Sweden: Stockholm, L. Romell, 67, 100, 101, and in Romell,
Fungi Scand., 33.

Finland: Mustiala, P. A. Karsten, authentie specimen of P.
aemulans.

Austria: Tirol, V. Litschauer, four specimens.

Italy: locality not given, G. Bresadola, two specimens.

England: Knys Lynn, C. B. Plowright, in Cooke, Fungi Brit., 7;
Yorkshire, E. M. Wakefield (in Mo. Bot. Gard. Herb., 57124).

Newfoundland: Bay of Islands, A. C. Waghorne, 5, 165 (in Mo.
Bot. Gard. Herb., 43987, 5010).

Ontario: Harraby, Lake Rosseau, E. T. & S. A. Harper, ?91.

Maine: Piscataquis County, W. A. Murrill, 1861 (in N. Y. Bot.
Gard. Herb., Burt Herb., and Mo. Bot. Gard. Herb., 61591);
Orono, P. L. Ricker, 621.

New Hampshire: Chocorua, W. G. Farlow, c89 (in Mo. Bot.
Gard. Herb., 43974), and in Reliq. Farlowianae, 342.

Vermont: Middlebury, E. A. Burt, six gatherings; Ripton, Е. A.
Burt, two gatherings.

Massachusetts: W. G. Farlow, 4; Sharon, A. P. D. Piguet, 5,
two unnumbered specimens, comm. by W. G. Farlow (in Mo.
Bot. Gard. Herb., 55220, 55446, 55600).

New York: Albany, H. D. House, six gatherings (in Mo. Bot.
Gard. Herb., 57412, 57457, 57519, 59684, 59700, 63448);
Alcove, С. L. Shear, 246; Fulton, А. E. Fivaz, comm. by А. Н.
W. Povah, 136 (in Mo. Bot. Gard. Herb., 58158); Ithaca,
G. F. Atkinson, 3034, C. J. Humphrey, C. O. Smith, comm. by
G. F. Atkinson, 8225, Van Hook, comm. by G. F. Atkinson,
8066, H. H. Whetzel, Plant Path. Herb., 12228 (in Mo. Bot.
Gard. Herb., 60599); Orient, В. Latham, 144 (in Mo. Bot.
Gard. Herb., 44230) ; Orient Point, R. Latham, comm. by N. Y.
State Mus. Зе (in Mo. Bot. Gard. Herb., 55815, 55922);
Syracuse, L. M. Underwood, 2, 89 (in N. Y. Bot. Gard. Herb.,
and Mo. Bot. Gard. Herb., 61567, py Vaughns, 8. H.
Burnham, 14 (in Mo. Bot. Gard. Herb., 54500).

Pennsylvania: Center Hall, E. West, comm. iem L. O. Overholts,

[Vol. 12
310 ANNALS OF THE MISSOURI BOTANICAL GARDEN

3661 (in Mo. Bot. Gard. Herb., 54702); State College, L. O.
Overholis, 4807 (in Mo. Bot. Gard. Herb., 56340) ; Trexlertown,
W. Herbst, 28, and comm. by Lloyd Herb., 3611.

Maryland: Takoma Park, C. L. Shear, 1337.

District of Columbia: Washington, C. L. Shear, 1257, 1267.

West Virginia: Paw Paw, C. L. Shear, 1174.

South Carolina: Aiken, H. W. Ravenel, in Ravenel, Fungi Am.,
140.

Florida: Royal Palm Hammock, W. A. Murrill, 123, 136, comm.
by N. Y. Bot. Gard. Herb. (in Mo. Bot. Gard. Herb., 62098,
62099).

Alabama: Auburn, F. S. Earle, 2299 (in N. Y. Bot. Gard. Herb.,
and Mo. Bot. Gard. Herb., 61416); Montgomery County,
R. P. Burke, 19, 275, 630, 664, 666, 679, 699 (in Mo. Bot.
Gard. Herb., 16700, 57158, 63099, 63076, 63101, 63097, 63103).

Kentucky: C. G. Lloyd, 1878.

Ohio: College Hill, Aiken, comm. by C. G. Lloyd, 2327.

Wisconsin: Madison, W. Trelease (in Mo. Bot. Gard. Herb.,
44314); Palmyra, A. O. Stucki, 39.

Iowa: Woodbine, C. J. Humphrey & C. W. Edgerton, comm. by
C. J. Humphrey, 6566 (in Mo. Bot. Gard. Herb., 20691).

Missouri: Bismarck, L. O. Overholts (in Mo. Bot. Gard. Herb.,
63454); Perryville, L. O. Overholis, 2687 (in Mo. Bot. Gard.
Herb., 44287).

Kansas: Phillips County, E. Bartholomew; Rooks County, E.
Bartholomew, 2046 (in Mo. Bot. Gard. Herb., 4842, 44313).
Colorado: Mancos, G. G. Hedgcock, comm. by C. J. Humphrey,

2551 (in Mo. Bot. Gard. Herb., 9783).

New Mexico: Tyom Experiment Station, W. H. Long, 21564 (in
Mo. Bot. Gard. Herb., 55142).

ас. Farragut Вау, W. Trelease, 582 (in Mo. Bot. Gard. Herb.,

Washington: Bingen, W. N. Suksdorf, 715, 745, 760, 765, 881,
882, 904.

Japan: Prov. Bungo, A. Yasuda, 119, 123 (in Mo. Bot. Gard.
Herb., 59470, 59474).

85. Р. aurantiaca Bresadola in Bourdot & Galzin, Soc. Myc.
Fr. Bul. 28: 402. 1913 ; Rea, Brit. Basid. 694. 1922.

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 311

Corticium aurantiacum Bresadola, Fungi Trid. 2: 37. pl. 144,
f. 2. 1892; Saec. Syll. Fung. 11: 126. 1895.—K neiffia auran-
tiaca Bresadola, Ann. Мус. 1: 103. 1903.—Gloeopeniophora
aurantiaca (Bres.) v. Héhnel & Litschauer, K. Akad. Wiss. Wien
Sitzungsber. 117: 1094. 1908.

Type: probably in Bresadola Herb.; authentic specimen in
Burt Herb.

Fructifications effused, beginning as small, convex outgrowths
at lenticels of the bark, spreading so as to form circular patches
which become confluent, adnate, bright orange-pink to orange-
chrome, fading in the herbarium to light pinkish cinnamon and
light buff, the margin white at first, radiating; in section not
colored, 150-250 » thick, with the hyphae densely and longi-
tudinally arranged in a rather broad layer next to the substratum
except at the points of emergence from the lenticels, hyaline,
thin-walled, 3-4. in diameter; gloeocystidia 30-60 x 6-9 u,
abundant in the convex portions; cystidia rough-walled, pointed,
up to 45 X 8 п, sometimes protruding 30 џ beyond the База,
more often wholly immersed and 30 x 4-54; База large,
60 x 10-12 u, often protruding beyond the immature basidia
when fruiting and bearing 4 sterigmata; spores hyaline, even,
12-16 x 6-12 y.

Fructifications 1-5 mm. in diameter at first, then laterally
confluent over areas 1-10 cm. long, 44-2 em. broad.

On dead Alnus of various species. Labrador to North Caro-
lina, westward to northern United States and Canada to British
Columbia and Oregon, and in Europe. August to November.
Common.

P. aurantiaca is easily recognized by its occurrence on dead
twigs of alder, in bright incarnate or orange-red fructifications
with large spores up to 15 x 10 v. These spores are usually
borne copiously and show well in erushed preparations. To
demonstrate the gloeocystidia and cystidia it is necessary to
examine sections cut through the convex or papilliform points of
origin of the fructifications. Sometimes examination of many
sections is necessary for demonstration of the cystidia. Failure
to cut the sections from places above stated led me to refer
gatherings of this species to Corticium laetum for some of my cor-

[Vol. 12
312 ANNALS OF THE MISSOURI BOTANICAL GARDEN

respondents. C. laetum may occur on Alnus and has color and

spores like P. aurantiaca.
Specimens examined:

Exsiecati: de Thümen, Myc. Univ., 112, under the name Cor-
ticium incarnatum; Linhart, Fungi Hung., 438, under the name
Peniophora incarnata.

Sweden: L. Romell, 62. |

Austria: Tirol, V. Litschauer, three specimens, E. Rehm, in Мус.
Univ., 112.

Hungary: Petrozsény, G. Linhart, in Linhart, Fungi Hung., 438.

Italy: Trient Alps, G. Bresadola, authentic specimen.

England: Lyndhurst, Hamp., E. M. Wakefield (in Mo. Bot.
Gard. Herb., 57127).

Labrador: The Strait, A. C. Waghorne, 5 (in Mo. Bot. Gard.
Herb., 43986).

Newfoundland: Bay of Islands, A. C. Waghorne, 341 (in Mo. Bot.
Gard. Herb., 5012).

New Brunswick: Campobello, W. G. Farlow, 1.

Canada: J. Macoun, 17, 116; Carleton's Place, J. Macoun, 158;
Ottawa, J. Macoun, 25.

Maine: Costigan, W. A. Murrill, 1766 (in N. Y. Bot. Gard.
Herb., and Mo. Bot. Gard. Herb., 61407); Kittery Point, R.
Thaxter & Е. A. Burt; Portage, L. W. Riddle, 13; Westbrook,
P. І. Ricker, 977.

New Hampshire: Chocorua, W. G. Farlow, c6 (in Mo. Bot. Gard.
Herb., 44124); Hanover, G. В. Lyman, 24 (in N. Y. Bot. Gard.
Herb., and Mo. Bot. Gard. Herb., 61587).

Massachusetts: Weston, A. B. Seymour, T34 (in Mo. Bot. Gard.
Herb., 15759).

New York: Childwold, G. F. Atkinson & B. M. Duggar, Cornell
Univ. Bot. Dept. 5056; Hudson Falls, S. H. Burnham, 28 (in
Mo. Bot. Gard. Herb., 54491); Karner, H. D. House, 14.164
and another specimen, comm. by N. Y. State Mus. Herb. (in
Mo. Bot. Gard. Herb., 44713, 54372).

Pennsylvania: Bear Meadows, L. O. Overholts, 2677 (in Mo. Bot.
Gard. Herb., 20277).

North Carolina: Chapel Hill, W. C. Coker, 4705 (in Mo. Bot.
Gard. Herb., 57425).

125]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 313

Michigan: Gogebic County, E. A. Bessey, 184 (in Mo. Bot. Gard.
Herb., 56581); Vermilion, A. H. W. Povah (in Mo. Bot. Gard.
Herb., 18274).

Montana: Evaro, J. R. Weir, 410 (in Mo. Bot. Gard. Herb.,
21617); Missoula, J. R. Weir, 349, 425 (in Mo. Bot. Gard.
Herb., 6105, 14765).

Idaho: Addie, J. R. Weir, 12005 (in Mo. Bot. Gard. Herb.,
63321).

Manitoba: Norway House, G. R. Bisby, 1460 (in Mo. Bot. Gard.
Herb., 61642).

British Columbia: J. Macoun, 752, comm. by J. Dearness (in
Mo. Bot. Gard. Herb., 12027); Agassiz, J. R. Weir, 359 (in
Mo. Bot. Gard. Herb., 16760); Salmo, J. В. Weir, 515 (in
Mo. Bot. Gard. Herb., 14170); Sidney, J. Macoun, 1, 72, 752
(in Mo. Bot. Gard. Herb., 5755, 55339, 55318).

Washington: Kalama, C. J. Humphrey, 6134; Olympia, C. J.
Humphrey, 6304; Seattle, W. A. Murrill, 141, and an un-
numbered specimen, comm. by N. Y. Bot. Gard. Herb. (in
Mo. Bot. Gard. Herb., 55731, 55733).

Oregon: Corvallis, S. M. Zeller, 1906 (in Mo. Bot. Gard. Herb.,
56882).

86. Р. pubera (Fr.) басс. Syll. Fung. 6: 646. 1888; Massee,
Linn. Soc. Bot. Jour. 25: 149. 1889; Bourdot & Galzin, Soc.
Mye. Fr. Bul. 28: 400. 1913; Rea, Brit. Basid. 693. 1922.
Thelephora pubera Fries, Elenchus Fung. 1: 215. 1828.—Cor-

ticium puberum Fries, Epicr. 562. 1838; Hym. Eur. 652. 1874;
Patouillard, Tab. Anal. Fung. 1: 66. f. 152. 1883; Bresadola,
Fungi Trid. 2: 38. pl. 145. f. 1. 1892.—P. pubera forma villosa
Bresadola, I. R. Accad. Agiati Atti 3: 113. 1897.—Kneiffia
pubera (Fr.) var. villosa Bresadola, Ann. Мус. 1: 101. 1903.

Fructifications effused, closely adnate, white, becoming dirty
whitish to light buff and pinkish buff in the herbarium, and widely
cracked, the hymenium even, setulose with the large cystidia,
the margin indeterminate, thinning out; in section 45-400 y
thick, not colored, composed of rather crowded, erect hyphae
2-4 y in diameter, thin-walled, not incrusted, and of gloeocystidia
and incrusted cystidia; gloeocystidia flexuous, 30-60 x 415-9 u;

[Vol. 12
314 ANNALS OF THE MISSOURI BOTANICAL GARDEN

cystidia incrusted, conical, pointed, fusiform, 50-90 x 8-20 р,
wholly immersed or protruding up to 50 в; spores hyaline, even,
depressed on one side.

Fructifications 2-6 em. long, 1-3 ст. wide.

On decaying wood, logs and limbs of frondose species, rarely on
conifers. In Europe and from Canada to Louisiana and west-
ward to British Columbia and Oregon. May to January. Com-
mon.

P. pubera is characterized by having gloeocystidia, large, con-
ical, heavily incrusted cystidia, and spores usually 7-9 x 3144-4 y.
The gloeocystidia show well in my permanent mounts in glyc-
erine, after the sections have been stained with eosin and stood
. for а few hours in glycerine. I did not find an authentic specimen
of P. pubera in Kew or Fries Herbaria but specimens received
under this name from Bresadola, Litschauer, Romell, and Miss
Wakefield have gloeocystidia in every specimen and other char-
acters as stated and show agreement in the European concept
of this species. Specimens with the other characters of P. pubera
but lacking gloeocystidia should be compared with P. guttulifera.
In North America, P. pubera forms thinner fructifications than
in Europe and is sometimes paler, drying rarely whitish or with
a slight yellowish tint.

Specimens examined:

Sweden: Góteborg, L. Romell, 174.

Germany: Westphalia, Lengerich, W. Brinkmann, authentic
specimen of P. pubera Fr. f. villosa from Bresadola.

Austria: Tirol, Innsbruck, V. Litschauer, two specimens; Stubai,
V. Litschauer.

Italy: Trient, G. Bresadola.

Great Britain: S. Wales, Swansea, E. M. Wakefield (in Mo. Bot.
Gard. Herb., 57123).

Canada: Quebec, Hull, J. Macoun, 388.

New Hampshire: Chocorua, W. G. Farlow, 6, 6b, 27, and 29 (in
Burt Herb.), and an unnumbered specimen and 152 (in Mo.
Bot. Gard. Herb., 7843, 55244). |

Massachusetts: Magnolia, W. G. Farlow, two specimens.

Rhode Island: Woonsocket, W. H. Snell, 7M, 8M (in Mo. Bot.
Gard. Herb., 56805, 56806).

1925]
URT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 315

New York: Ithaca, G. F. Atkinson, 8227, 14363, and C. Thom,
comm. by G. F. Atkinson, 14372; Karner, H. D. House, 14.165
in part (in Mo. Bot. Gard. Herb., 44715).

New Jersey: Newfield, J. B. Ellis, three specimens (in N. Y. Bot.
Gard. Herb., and Mo. Bot. Gard. Herb., 61401, 61443, 63467).

Maryland: Takoma Park, C. L. Shear, 1129, 1158.

District of Columbia: Takoma Park, C. L. Shear, 964.

Virginia: Crabbottom, W. A. Murrill (in N. Y. Bot. Gard. Herb.).

Florida: W. W. Calkins, 860 (in N. Y. Bot. Gard. Herb., and Mo.
Bot. Gard. Herb., 61455).

Alabama: Montgomery County, R. P. Burke, 207, 208, 277, 847,
375, 376, 377, 424, 460, 468, 658, 662, 806, 807 (in Mo. Bot. Gard.
Herb., 57080, 57081, 57159, 57218, 57245, 57244, 57243, 57261,
57282, 57287, 63085, 63087, 63109, and 63110 respectively).

Louisiana: St. Martinville, A. B. Langlois, 2685, cl.

Kentucky: Harlan, C. H. Kauffman, 66 (in Mo. Bot. Gard. Herb.,
16346).

Wisconsin: Blue Mounds, Miss A. C. Stucki, 38; Madison, C. J.
Humphrey, 2488 (in Mo. Bot. Gard. Herb., 11277).

Missouri: Creve Coeur Lake, L. O. Overholts, 3166, 661 (in Mo.

. Bot. Gard. Herb., 5708, 5710).

Montana: Anaconda, Mt. Hagan, J. R. Weir, 11253 (in Mo. Bot.
Gard. Herb., 63256).

Idaho: Coolin, J. R. Weir, 11505, 11516 (in Mo. Bot. Gard.
Herb., 63286, 63289).

Manitoba: Winnipeg, G. R. Bisby, 1345 (in Mo. Bot. Gard.
Herb., 60555).

British Columbia: Cormac, J. Macoun, 658 (in Mo. Bot. Gard.
Herb., 55328); Sidney, J. Macoun, 788 (in Mo. Bot. Gard.
Herb., 55329); Vancouver Island, Oak Bay, J. Macoun, 600
(in Mo. Bot. Gard. Herb., 55327); Victoria, J. Macoun, 564
(in Mo. Bot. Gard. Herb., 55326).

Washington: Chehalis, C. J. Humphrey, 62

Oregon: Corvallis, S. M. Zeller, 1851, cs 1904 (in Mo. Bot.
Gard. Herb., 56863, 56864, 56880).

87. P. pertenuis (Karsten) Burt, n. comb.
Corticium pertenue Karsten, Hedwigia 29: 270. 1890; Finska

[Vol. 12
316 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Vet.-Soc. Bidrag Natur och Folk 51: 226. 1892; Saec. Syll.
Fung. 9: 234. 1891; v. Hóhnel & Litschauer, K. Akad. Wiss.
Wien Sitzungsber. 115: 1556. 1906—(In part) Gloeocystidium
praetermissum (Karst.) v. Hóhnel & Litschauer, K. Akad. Wiss.
Wien Sitzungsber. 115: 1565. 1906.—An Peniophora praeter-
missa Karsten, Finska Vet.-Soc. Bidrag Natur och Folk 48:
423. 1889?

Type: probably in Karsten Herb. and fragment in Burt Herb.

Fructifications long-effused, closely adnate, not separable
when moist, very thin, waxy, even, white or whitish, drying pale
pinkish buff to cream color, the margin thinning out; in section
60-150 џ thick, not colored, with the hyphae 3-4 y in diameter, ©
erect, branching, not incrusted, bearing the compact hymenium;
gloeocystidia numerous, variable in form, often tapering, 20-60
х 6-8 ш; cystidia hair-like, not incrusted, 4-5 u in diameter,
protruding up to 20 џ beyond the basidia, few and scattered;
spores copious, hyaline, even, curved, 7-10 x 4-5 y.

Fructifications 2-10 em. long, 1-2 cm. wide.

On decaying coniferous wood. In Europe, in Canada to District
of Columbia, and in Oregon, Jamaica and Bermuda. July to
November. Rare in North America.

The principal characteristics of P. pertenuis aiding in its recog-
nition are occurrence in thin, whitish, waxy fructifications on
old decaying coniferous wood, presence of gloeocystidia, and hair-
like non-incrusted cystidia which are not destroyed in any degree
by the treatment of the sections with potassium hydrate solution,
and the curved spores. P. tenuis differs in having its cystidia
incrusted at the tip. I have not seen an authentic specimen of
P. praetermissa but the specimen sent to me under this name by
Bresadola and one of the two specimens from Litschauer have
their cystidia almost completely disintegrated by the potassium
hydrate treatment in clearing and swelling the tissues of sections,
as occurs also in P. glebulosa. Hence, I think that P. praeter-
missa may eventually be regarded by European mycologists
as specifically distinct from P. pertenuis.

Specimens ined:

Finland: Mustiala, P. A. Karsten, portion of type, comm. by

Bresadola, also authentie specimen on Picea from Karsten.

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 317

Sweden: Stockholm, L. Romell, 116, 138, 162, 168, 164, 183, 191,
193, 208, 212, 215.

Austria: Natters, Tirol, V. Litschauer, under the name Gloeo-
cystidium praetermissum.

Canada: Ottawa, J. Macoun, 42, 313; St. Lawrence Valley, J.
Macoun, 41.

New Hampshire: Chocorua, W. G. Farlow, 5.

New Jersey: Newfield, J. B. Ellis (in N. Y. Bot. Gard. Herb. and
Mo. Bot. Gard. Herb.).

District of Columbia: Washington, C. J. Humphrey, 2525 (in
Mo. Bot. Gard. Herb., 20383).

Montana: Trego, J. R. Weir, 11969 (in Mo. Bot. Gard. Herb.,
63227).

Idaho: Coolin, J. R. Weir, 11512 (in Mo. Bot. Gard. Herb.,
63287); Priest River, J. R. Weir, 139 (in Mo. Bot. Gard.
Herb., 63468).

- Oregon: Portland, С. J. Humphrey, 6127.

Washington: Falcon Valley, W. N. Suksdorf, 724.

Bermuda: Paget Marsh, on Sabal, H. H. Whetzel, Ao, Abd (in
Mo. Bot. Gard. Herb., 58719, 58906).

Jamaica: Troy and Tyre, W. A. Murrill & W. Harris, 1053, in
part, comm. by N. Y. Bot. Gard. Herb.

88. P. tenuis (Pat.) Massee, Linn. Soc. Bot. Jour. 25: 149.
1889.

Corticium tenue Patouillard, Rev. Mye. 7: 152. 1885; Tab.
Anal. Fung. 1: 203. f. 462. 1886; Sacc. Syll. Fung. 6: 632.
1889.—Kneiffia tenuis (Pat.) Bresadola, Ann. Myc. 1: 105. 1903.
— Gloeocystidium tenue (Pat.) v. Hóhnel & Litschauer, Wiesner
Festschr. Wien, 70. 1908; Bourdot & Galzin, Soc. Myc. Fr.
Bul. 28: 364. 1913.

Fructifications effused, closely adnate, thin, white, drying
whitish to pale pinkish buff, somewhat pruinose, the margin
thinning out; in section 60-180» thick, not colored, with the
hyphae erect, branching, 3-44 in diameter, thin-walled, not
incrusted; gloeocystidia 20-60 x 6-8 р, flexuous; cystidia hair-
like, cylindric, 46, in diameter, protruding 20—45 р beyond
the basidia, incrusted about the tip; spores hyaline, even, curved,
8-10 x 4416 {he

Fructifications 2-6 cm. long, 1-2 ст. wide.

[Vol. 12
318 ANNALS OF THE MISSOURI BOTANICAL GARDEN

On decaying wood and bark of frondose species more usually.
Europe and Massachusetts. July to December. Rare.

P. tenuis is doubtfully distinct from P. pertenuis, having the
same aspect and microscopical characters except that some of
the cystidia have incrusting granules at the tips, as shown in
the figures by Patouillard cited above.

Specimens examined:

Germany: Westphalia, Lengerich, W. Brinkmann, comm. by

Bresadola.

Austria: Tirol, V. Litschauer.
France: Allier, St. Priest, H. Bourdot, 6530; Aveyron, A. Galzin,

11689, comm. by H. Bourdot, 18554.

Massachusetts: Brookline, Hammond's Pond, G. R. Lyman, 183.

89. P.serialis (Bres.) v. Hóhnel & Litschauer, К. Akad. Wiss.
Wien Sitzungsber. 116: 777. 1907.

Kneiffia serialis Bresadola, Ann. Myc. 1: 101. 1903 (in part);
Sydow, Myc. Germ., 1. 1903.—Not X erocarpus Cacao Karsten,
Hedwigia 29: 271. 1890.—An Corticium seriale Fries, Epicr.
563. 1838?—Corticium seriale (Bres.) Bourdot & Galzin, Soc.
Myce. Fr. Bul. 27: 253. 1911 (Forme 2).

Type: type distribution in Sydow, Myc. Germ., 1.

Fructifications long and widely effused, thin, closely adnate,
very variable in color, smoke-gray and pale olive-buff to wood-
brown in the herbarium, even, sometimes cracked; the margin
thinning out, indet inate; in section 60-180 ү. thick, not colored,
composed of densely arranged, erect hyphae about Зи in di-
ameter, with the outer portion of the wall gelatinously modified
and indistinct, and of some scattered, yellowish or brownish,
somewhat spherical masses 9-12» in diameter, immersed near
the substratum; gloeocystidia in the unusual form of irregular,
immersed spherical masses 9-12 » in diameter; cystidia not in-
crusted, tapering to a sharp apex, 3-5 џ in diameter, protruding
up to 30u; spores hyaline, even, curved, 4-6 x 1-2 y.

Fructifications 3-12 cm. long, 2-5 cm. wide.

On decaying wood of logs of Pinus, Abies, Tsuga, and Thuja.
Europe, New York, and Washington. August to May.

P. serialis resembles in aspect P. Cacao and Corticium lividum;

1925]
URT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 319

from the latter, the more common species, it is distinguished by
its cystidia and from both by the immersed, spherical, colored
masses near the substratum, such as were described and figured
for another species with larger spores as gloeocystidia by von
Hóhnel & Litschauer in K. Akad. Wiss. Wien Sitzungsber. 116:
838. 1907.
Specimens examined:
Exsiccati: Sydow, Myc. Germ., 1.
Sweden: Femsjó, E. A. Burt, two gatherings.
Germany: Brandenburg, P. Sydow, type distribution, in Sydow,
Myc. Germ., 1, and comm. by Bresadola.
New York: Floodwood, C. H. Peck, 8.
Washington: Sedro Woolley, C. J. Humphrey, 7538.

90. P. typhicola Burt, n. sp.

Type: in Burt Herb.

Fructifications effused, thin, adnate, somewhat membrana-
ceous, tender, between whitish and pale olive-buff in the her-
barium, even, not shining, not cracked, the margin thinning out,
indeterminate; in section 60-75 ų thick, not colored near the sub-
stratum, composed of suberect, densely interwoven, thin-walled
hyphae 2-3 p in diameter, indistinct, and of incrusted cystidia
and a few gloeocystidia; gloeocystidia flexuous, 25-30 x 4-6 в,
few present; cystidia incrusted, 40 x 15 и, immersed, starting
from the substratum; paraphyses with filiform tips about 14-1 р
in diameter, with 1 or 2 lateral branches but not antler-shaped,
in surface of hymenium; spores hyaline, even, 8-12 x 315-4 у,
two to a basidium.

Fructifications 2-10 mm. in diameter.

On dead Typha latifolia. New York.

This specimen was at first doubtfully referred to P. phyllophila
which it resembles in aspect and somewhat in structure, but it is
thicker, more dense, has gloeocystidia, and does not have con-
spicuous antler-shaped paraphyses. Reference to Epithele Ty-
phae, which I have been unable to find in our North American
species, is precluded by the absence of hyphal fascicles.

Specimens examined:
New York: Ithaca, G. F. Atkinson, 261, type.

У [Vol. 12
320 ANNALS OF THE MISSOURI BOTANICAL GARDEN

91. P. filamentosa (Berk. & Curtis) Burt in Coker, Elisha
Mitchell Scientif. Soc. Jour. 36: 162. pl. 32, f. 5,6. 1921.

Corticium filamentosum Berkeley & Curtis, Grevillea 1: 178.
1873; Басс. Syll. Fung. 6: 619. 1888; Massee, Linn. Soc. Bot.
Jour. 27: 154. 1890.—(In part) Corticium Petersii Berkeley &
Curtis, Grevillea 1: 177. 1873.—Peniophora unicolor Peck,
М. Y. State Mus. Rept. 43: 66. 1890; Sace. Syll. Fung. 9:
239. 1891.—Ап Corticium radicatum Р. Hennings, Pilze Ostaf-
rikas, 54. 1895; Басе. Syll Fung. 14: 222. 1899? See v.
Hóhnel & Litschauer, K. Akad. Wiss. Wien Sitzungsber. 117:
1093. 1908.

Type: in Kew Herb. and Curtis Herb.

Fructifications broadly effused, membranaceous, loosely ad-
nate, separable when moistened, soft, drying Isabella color to
buffy citrine, the margin and subiculum concolorous with, or a
little paler than, the hymenium, often extended into, or con-
nected with, branching mycelial strands or cords; in section
150-400 » thick, colored, with the hyphae loosely interwoven,
thin-walled, 3-4 y in diameter, densely incrusted with ochraceous
granules which are not soluble in lactic acid preparations, but
dissolve quickly when sections are treated with potassium hydrate
solution and leave the sections bleached, after first becoming
vinaceous; cystidia incrusted, 40-50 х 6-8u, protruding up to
40 u, confined to the hymenial layer ; Spores white in spore col-
lection, even, 3-5 x 2-3 y.

ructifications 2-10 cm. long, 1-3 em. broad; sometimes much
larger on logs by confluence longitudinally.

On decaying wood and logs and fallen limbs of frondose species.
Germany, Canada to Alabama, and westward to Arizona, in
Mexico, the West Indies, and Japan. July to January. Com-
mon.

Although colored like a Coniophora, P. filamentosa is easily
recognized by its marginal mycelial strands, small and white
spores, and hyphae incrusted with ochraceous granules which are
soluble in the 7 per cent solution of potassium hydrate with
which sections are usually treated. Since the original description
of Corticium Petersii combines the characters of both Peniophora
sanguinea and P. filamentosa, and one of the types is of one species

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 321

and the other of the other, C. Petersit has been reduced to syn-

onymy and C. filamentosum of the same authors adopted for the

present species.
Specimens examined:

Exsiccati: Ravenel, Fungi Car. 5: 28, the type distribution of
Corticium Раетзи.

Germany: Hannover, Engelke (in Mo. Bot. Gard. Herb., 43481,
under the name Peniophora radicata).

Canada: J. Macoun, 298; Lower St. Lawrence Valley, J. Macoun,
6, 28, 38.

New Hampshire: Franconia, W. G. Farlow, 26.

Vermont: Middlebury, E. A. Burt, two gatherings.

New York: Albany, L. O. Overholts, 3389 (in Mo. Bot. Gard.
Herb., 10179); Altamont, C. H. Peck (in N. Y. State Mus.
Herb., and Mo. Bot. Gard. Herb., 55211); Bolton, C. H. Peck,
15; Bolton Landing, C. H. Peck (in N. Y. State Mus. Herb.,
and Mo. Bot. Gard. Herb., 55975, 56021); Cazenovia, L. M.
Underwood, 46 (in N. Y. Bot. Gard. Herb., and Mo. Bot.
Gard. Herb., 61405); East Berne, C. H. Peck (in N. Y. State
Mus. Herb., and Mo. Bot. Gard. Herb., 56016); East Galway,
E. A. Burt, two gatherings; East Schagticoke, C. H. Peck (in
N. Y. State Mus. Herb., and Mo. Bot. Gard. Herb., 56022);
North Elba, C. H. Kauffman, 9 (in Mo. Bot. Gard. Herb.,
16769); North Greenbush, H. D. House, 14.235 (in N. Y. State
Mus. Herb., and Mo. Bot. Gard. Herb., 14840, 44734); Hudson
Falls, S. H. Burnham, 23 (in Mo. Bot. Gard. Herb., 54487);
Ithaca, G. F. Atkinson, 7892; Staten Island, W. H. Ballou (in
Mo. Bot. Gard. Herb., 10276); Syracuse, L. M. Underwood,
type of Peniophora unicolor (in N. Y. State Mus. Herb.).

New Jersey: J. B. Ellis, comm. by W. G. Farlow (in Mo. Bot.
Gard. Herb., 44645); Newfield, J. B. Ellis, 1518, comm. by
W. G. Farlow (in Mo. Bot. Gard. Herb., 14645); Orange,
L. M. Underwood (in N. Y. Bot. Gard. Herb., and Mo. Bot.
Gard. Herb., 61576).

Maryland: Takoma Park, C. L. Shear, 1097.

North Carolina: Chapel Hill, J. N. Couch, Univ. N. C. Herb.,
4607 (in Mo. Bot. Gard. Herb., 57423).

Alabama: Peters, type (in Kew Herb., and Curtis Herb., 6119),

[Vol. 12
322 ANNALS OF THE MISSOURI BOTANICAL GARDEN

and in Ravenel, Fungi Car. 5: 28; Montgomery County, R. P.
Burke, 425, 660 (in Mo. Bot. Gard. Herb., 57270, 63086).

Kentucky: Crittenden, C. G. Lloyd (in Lloyd Herb., 10113, and
Mo. Bot. Gard. Herb., 65627).

Ohio: C. G. Lloyd, 3883; Cincinnati, C. G. Lloyd, 4504.

Michigan: New Richmond, C. H. Kauffman, 49 (in Mo. Bot.
Gard. Herb., 3734).

Illinois: Riverside, E. Т. & S. A. Harper, 852.

Missouri: Columbia, B. M. Duggar, 447.

Arizona: Fort Valley Experiment Station, W. H. Long, 21121 (in
Mo. Bot. Gard. Herb., 55138).

Mexico: Jalapa, W. A. & E. L. Murrill, 178, 327, comm. by N. Y.
Bot. Gard. Herb. (in Mo. Bot. Gard. Herb., 44969, 54496).
Cuba: C. G. Lloyd, 420 (in Mo. Bot. Gard. Herb., 55173) ; Omaja,

C. J. Humphrey, 2782 (in Mo. Bot. Gard. Herb., 14850).
Jamaica: Troy and Tyre, W. А. Murrill & W. Н. arris, 911, comm.
by N. Y. Bot. Gard. Herb.
Japan: Prov. Bunga, A. Yasuda, 113 (in Mo. Bot. Gard. Herb.,
59463).

92. P. viticola (Schw.) v. Hóhnel & Litschauer, K. Akad.
Wiss. Wien Sitzungsber. 116: 779. text Job 41997.

Thelephora viticola Schweinitz, Naturforsch. Ges. Leipzig
Schrift. 1:107. 1822; Am. Phil. Soc. Trans. N. S. 4:168. 1832 ;
Fries, Elenchus Fung. 1: 205. 1828.—Corticium viticola Fries,
Epier. 561. 1838; Ѕасе. Syll. Fung. 6: 617. 1888; Massee,
Linn. Soc. Bot. Jour. 27: 146. 1890; Coker, Elisha Mitchell
Scientif. Soc. Jour. 36: 172. pl. 33, f. 6. 1921.—Corticium croci-
creas Berkeley & Curtis, Grevillea 1: 178. 1873; Sacc. Syll.
Fung. 6: 616. 1888. Not С. crocicreas Massee nor v. Hóhnel &
Litschauer.—Corticium subaurantiacum Peck, N. Y. State Mus.
Rept. 43: 67. 1890; Sacc. Syll. Fung. 9: 230. 1891.

Type: in Schweinitz Herb.

Fructifications effused, thin, adnate, soft, small portions separ-
able when moistened, the tomentose subiculum and margin
ochraceous orange, the hymenium even, grayish to buff-yellow
and pruinose; in section 150-400 u thick, with the denser and
broader subhymenial region ochraceous orange and the more

1925]
URT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 323

loosely interwoven region next to the substratum yellow, the
loosely interwoven hyphae thin-walled, 2-4 џ in diameter, not
nodose-septate, incrusted with colored granules which give the
color to the fructification, and are destroyed and dissolved by the
action of potassium hydrate solution leaving the sections bleached ;
no gloeocystidia; cystidia not incrusted, thin-walled, cylindric,

6-9 и in diameter, protruding 25-40 u beyond the basidia; basidia

with 4 sterigmata; spores white in collection on slide, even,

7-8 X 4-5 y.

Fructifications 1-2 cm. in diameter, becoming confluent over
areas 3-8 em. long, 2-5 em. wide.

On bark and wood of decaying Vitis, Abies, Acer, and Fagus.
Vermont to North Carolina, Kentucky, and Arkansas. July to
October. Abundant locally.

P. viticola is conspicuous by the large, brilliant orange fruc-
tifications with paler, pruinose fertile hymenium which occur on
rotting large stems of the wild grape and on logs in deep mountain
forests. The bleaching of the sections through destruction and
solution of the incrusting pigment granules is common also to
P. filamentosa, from which P. viticola differs in more orange color
and larger spores.

Specimens examined:

Exsiccati: Ravenel, Fungi Car. 3: 34.

Vermont: Bread Loaf, E. A. Burt; Little Notch, E. A. Burt;
Middlebury, E. A. Burt, determination as Corticium sub-
aurantiacum confirmed by Peck.

New York: Ampersand, C. H. Peck (in N. Y. State Mus. Herb.,
T 20, and Mo. Bot. Gard. Herb., 54638); Clear Water, G. F.
Atkinson, 5043 (in Cornell Univ. Herb.); Floodwood, E. A.
Burt; Lake Placid, W. A. & E. L. Murrill, 104 (in N. Y. Bot.
Gard. Herb., and Mo. Bot. Gard. Herb., 57344); Long Lake,
A. H. W. Povah, 13 (in Mo. Bot. Gard. Herb., 9084); Marcy
Trail, C. H. Peck (in N. Y. State Mus. Herb., T 19, and Mo.
Bot. Gard. Herb., 54637); North Elba, C. H. Kauffman, 8 (in
Mo. Bot. Gard. Herb., 6686); Ray Brook, C. H. Peck (in N. Y.
State Mus. Herb., T 21, and Mo. Bot. Gard. Herb., 54639) ;
Undercliff, comm. by Univ. Wis. Herb., 45.

North Carolina: Salem, Schweinitz, type (in Herb. Schweinitz).

[Vol. 12
324 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Alabama: Peters, in Ravenel, Fungi Car. 3: 34, and as Corticium
crocicreas Berk. & Curtis, type (in Kew Herb. and in Curtis
Herb., 4542).

Kentucky: Mammoth Cave, C. G. Lloyd, 1601, 2661, and another
specimen comm. by Ellis Herb.

Arkansas: Fordyce, C. J. Humphrey, 5799.

93. P.sulphurina (Karst.) v. Hóhnel & Litschauer, K. Akad.
Wiss. Wien Sitzungsber. 115: 1573. 1906.

Tomentella sulphurina Karsten, Finska Vet.-Soc. Bidrag
Natur och Folk 48: 420. 1889.—H ypochnus sulphurinus (Karst.)
Sacc. Syll. Fung. 9: 243. 1891.

Type: authentic specimen—perhaps part of type—in Burt Herb.

Fructifications effused, adnate, the hymenium drying clay
color, thin, brittle, even, here and there cracked and showing the
mustard-yellow subiculum, the margin fibrillose-byssoid, mus-
tard-yellow; in section 150—400 и thick, pale yellow, with the
hyphae loosely arranged, thin-walled, 46 in diameter, oc-
casionally nodose-septate, some hyphae granule-incrusted; cys-
tidia hair-like, not incrusted, 3-6 » in diameter, protruding up to
30 u, not numerous; spores hyaline, even, 3-4 X 2-21% ү.

Fructifications 2-6 em. long, 1-2 em. broad.

On coniferous bark usually. In Finland, from New Hampshire
to Alabama and westward to British Columbia and Oregon.
August to November. Rare.

The American gatherings referred to P. sulphurina are a little
paler than the European and the sections lose most of their color
when floated on alcohol in sectioning so as to become not dis-
tinetly colored in section. In other respects our specimens agree
so well with the authentic specimen that I believe they should be
included in this species. Potassium hydrate solution does not
change the color of the sections to vinaceous and bleach them as it
does sections of P. filamentosa.

Specimens examined:

Finland: Jalasjárvi, authentic specimen from P. A. Karsten.
New Hampshire: Chocorua, W. G. Farlow (in Mo. Bot. Gard.

Herb., 7872).

New York: Fall Creek, G. F. Atkinson, 7993; Ithaca, E. J. Petey,
comm. by C. J. Humphrey, 471; Rainbow, C. H. Peck (in

1925]
URT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 325

N. Y. State Mus. Herb., T 32, and Mo. Bot. Gard. Herb.,
54656) ; Yates, C. Н. Peck, 33.

Pennsylvania: Delaware Water Gap, W. A. Murrill, comm. by
N. Y. Bot. Gard. Herb. (in Mo. Bot. Gard. Herb., 61469).

` New Jersey: Newfield, J. B. Ellis, comm. by У. С. Farlow (in
Mo. Bot. Gard. Herb., 7764, 14290, 14770).

District of Columbia: Takoma Park, C. L. Shear, 1348.

Alabama: Montgomery, R. P. Burke, 145 (in Mo. Bot. Gard.
Herb., 10359).

Kentucky: Crittenden, C. G. Lloyd, 3120.

South Dakota: Sylvan Lake, Custer, J. R. Weir, 10011 (in Mo.
Bot. Gard. Herb., 55792).

Idaho: Priest River, J. E. Weir, 30.

British Columbia: Squamish, J. Macoun, 497, 534 (in Mo. Bot.
Gard. Herb., 55182, 55181).

Oregon: Eugene, C. J. Humphrey, 1051.

94. Р. carnosa Burt, n. sp.

Type: in Burt Herb., Mo. Bot. Gard. Herb., and N. Y. State
Mus. Herb.

Fructifications long and broadly effused, thick, fleshy-mem-
branaceous, adnate, barium-yellow to honey-yellow, of the same
color within where cracked, the margin determinate, thinning
out, somewhat radiate-fibrillose; in section 400-700 џ thick,
colored like the hymenium in thick sections but very thin sec-
tions hyaline, somewhat zoned, composed of a very broad hyphal
layer bearing а hymenial layer 50-60 y thick, the hyphae hyaline,
5-6 y. in diameter; no gloeocystidia; cystidia hair-like, not in-
crusted, tapering to a sharp tip, 4» in diameter at the base,
protruding up to 30 » beyond the basidia, very numerous in the
hymenial surface; basidia 4-spored; spores white in spore col-
lection, even, 4-5 x 2-215 y.

Fructifications 3-12 cm. long, 114-6 em. wide.

On bark and wood of coniferous logs such as Pinus, Abies,
Picea, Pseudotsuga, Juniperus, and Larix, rarely on frondose
species. In mountains of New England, New York, Minnesota,
and British Columbia and Montana to New Mexico. May to
October. Common in the Rocky Mountain forests.

[Vol. 12
326 ANNALS OF THE MISSOURI BOTANICAL GARDEN

P. carnosa may be recognized at sight by its large, thick,
yellow fructifications occurring on coniferous logs in forests of
the White Mountains, Adirondacks, and the Rocky Mountains.
The abundant cystidia are too small to be visible with a lens,
hence it is necessary to examine sections with a microscope to
recognize the species as a Peniophora rather than a Corticium.
It does not have the mustard-yellow subiculum of P. sulphurina
nor does its hymenial layer flake away from the substratum as in
the latter.

Specimens examined:

Maine: Piscataquis County, W. A. Murrill, 2311 (in N. Y. Bot.
Gard. Herb., and Mo. Bot. Gard. Herb., 61596).

New Hampshire: Chocorua, E. A. Burt; Intervale, L. O. Over-
holts, 5039 (in Mo. Bot. Gard. Herb., 56351); North Conway,
L. O. Overholts, 4732 (in Mo. Bot. Gard. Herb., 56117).

New York: Hague, C. H. Peck, type (in Burt Herb., N. Y. State
Mus. Herb., and Mo. Bot. Gard. Herb., 56019); North Elba,
C. H. Kauffman, 4 (in Mo. Bot. Gard. Herb., 21307).

Minnesota: Cass Lake, J. R. Weir, 392 (in Mo. Bot. Gard. Herb.,
12436).

Montana: Fortine, E. E. Hubert, comm. by J. R. Weir, 12013 (in
Mo. Bot. Gard. Herb., 63323); Missoula, J. R. Weir, 383 (in
Mo. Bot. Gard. Herb., 20892). :

Idaho: Meadow Creek, E. E. Hubert, comm. by J. R. Weir,
11669, 11671 (in Mo. Bot. Gard. Herb., 63308, 63310) ; Priest
River, E. E. Hubert, comm. by J. R. Weir, 11738 (in Mo. Bot.
Gard. Herb., 63311), J. R. Weir, 355, 9100 (in Mo. Bot. Gard.
Herb., 10883, 55955), and 26, 32, 51, 57, 62, 69.

British Columbia: Kootenai Mts. near Salmo, J. R. Weir, 450,
458, 464, 467, 468, 469, 479, 523, 530 (in Mo. Bot. Gard. Herb.,
8767, 9121, 12613, 12287, 8766, 12534, 12907, 20976, and 16078
respectively).

Washington: Olympia, C. J. Humphrey, 6291.

New Mexico: Cienega Canyon, W. H. Long, 21470, 21515, 21561
(in Mo. Bot. Gard. Herb., 55148, 55149, 55150); Sulphur
Canyon, W. H. Long, 21411 (in Mo. Bot. Gard. Herb., 55147);
Tyom Experiment Station, W. H. Long, 21935 (in Mo. Bot.
Gard. Herb., 55151).

1925]
URT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 327

95. P.citrinella (B. & C.) Burt, n. comb.

Corticium citrinellum Berkeley & Curtis, Linn. Soc. Bot. Jour.
10: 336. 1868; Sacc. Syll. Fung. 6: 616. 1888; Massee, Linn.
Soc. Bot. Jour. 27: 147. 1890.

Type: in Curtis Herb. and probably in Kew Herb.

Fructifications effused, thin, tender, small pieces separable
when moistened, barium-yellow, cracked and showing a byssoid,
barium-yellow subiculum, the margin thinning out, sometimes
with barium-yellow mycelial strands; in section 120—300 y thick,
barely barium-yellow when but slightly magnified but not per-
ceptibly colored under high magnification and wholly bleached
by treatment with potassium hydrate solution, 2-layered, with
the broader layer next to the substratum and composed of loosely
interwoven hyphae 212-3 џ in diameter under the incrustation
of scattered, coarse granules, not nodose-septate, and with the
hymenial layer 90 и thick, compact; no gloeocystidia; wholly
immersed cystidia few, incrusted, 15 X 9 u; protruding cystidia
hair-like, short, 3-414y in diameter, protruding up to 12y;
spores hyaline, even, 3-4 X 2-3 y.

Fructifications 1-3 cm. long, 4-1 cm. wide.

On bark of logwood limb on the ground. West Indies. October
to March.

P. citrinella belongs in the group of species with P. sulphurina,
P. limonia, P. Burtii, and P. subiculosa, and appears distinct from
each of these when specimens are compared with one another.
It has priority over all the others as a species. Its distinguishing
combination of characters is barium-yellow color, cracked hy-
menium showing subiculum of the same color, color bleached by
treatment with potassium hydrate solution, short cystidia, and
occurrence on аиа bark.

Specimens
Cuba: C. Wright, nd type (in Curtis Herb.); Pinar del Rio

Province, Earle & Murrill, 381, comm. by N. Y. Bot. Gard.

Herb.; Santa Clara Province, Earle & Murrill, 427, comm. by

N. Y. Bot. Gard. Herb.

Jamaica: Hope Gardens, F. S. Earle, 164, comm. by N. Y. Bot.

Gard. Herb.

[Vol. 12
328 ANNALS OF THE MISSOURI BOTANICAL GARDEN

96. Р. Sacchari Burt, n. sp.

Type: in Mo. Bot. Gard. Herb.

Fructifications longitudinally effused, adnate, thin, somewhat
membranaceous, noted as yellow when growing but now clay
color in the herbarium, even, cracking into polygonal masses
about 4% mm. in diameter which may curl away from the sub-
stratum more or less and show the exposed tissue colored like
the hymenium, the margin thinning out, of finely interwoven
hyphae; in section 75-110 џ thick, with the thin sections not
colored appreciably, composed of densely arranged, suberect
hyphae about 3 џ in diameter, not incrusted, not nodose-septate;
no gloeocystidia ; cystidia not incrusted, tapering towards the
apex, 5-6 y. in diameter, protruding up to 30 u; spores hyaline,
even, 314-4 x 215-3 y.

Fructifications 8 cm. long, 10-13 mm. wide.

On cane trash of Saccharum officinarum. Porto Rico. January.

P. Sacchari was listed by Johnston & Stevenson, Dept. of
Agr. Porto Rico Jour. 1: 227. 1917, as Peniophora sp. It has
not been received from other sources and is apparently a species
local to Porto Rico or with a preference for a sugar-cane sub-
stratum. It should be conspicuous by its yellow color. P.
citrinella is thicker and has incrusted cystidia.

Specimens examined:

Porto Rico: Rio Piedras, J. A. Stevenson, 1204, type (in Mo. Bot.

Gard. Herb., 11787).

97. P. medioburiensis Burt, n. sp.

Type: in Burt Herb.

Fructifications effused, thin, felty, small pieces separable when
moistened, becoming in the herbarium between light grayish
olive and deep olive-buff, the margin thinning out and some-
times paler; in section 200-300 y. thick, colored like the surface,
somewhat zonate, composed of suberect, thin-walled, even-
walled hyphae 3 џ in diameter and of incrusted hyphae 5 р in
diameter over the incrustation; no gloeocystidia; cystidia usually
not incrusted, sometimes granule-incrusted, cylindric, obtuse,
6-8 y. in diameter, protruding up to 30 beyond the basidia;
basidia with 4 large sterigmata up to 6 и long; spores white in
spore collection, cylindric, 8-14 X 414-6 y.

1925]
URT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 329

Fructifications 5 mm.-3 em. long, 5 mm.-1)4 em. wide.

On wood and bark of fallen, rotten limbs of Carya. Middle-
bury, Vermont. July. Seen but once.

This species is of felty or fibrillose structure like some species
of Hypochnus, not at all waxy, and of dull, olivaceous color so as
to be very inconspicuous on the fallen decaying limbs on which
found. So few species of Peniophora have spores up to 14 џ long
that the spore dimensions should prove in this instance an im-
portant character for recognition of P. medioburiensis.

Specimens examin
Vermont: Battell FENE Middlebury, E. A. Burt, type.

98. P. subsulphurea (Karst.) v. Höhnel & Litschauer, K.
Akad. Wiss. Wien Sitzungsber. 115: 1580, 1592. 1906; Bourdot
& Galzin, Soc. Myc. Fr. Bul. 28: 388. 1913.

Corticium subsulphureum Karsten, Soc. pro Fauna et Fl. Fen-
nica Meddel. 6: 12. 1881; Saec. Syll. Fung. 6: 632. 1888;
Massee, Linn. Soc. Bot. Jour. 27: 148. 1890.—Xerocarpus
subsulphureus Karsten, Finska Vet.-Soc. Bidrag Natur och Folk
37: 138. 1882; 48: 417. 1889. See v. Hóhnel & Litschauer,
K. Akad. Wiss. "ien Sitzungsber. 117: 1093. 1908.—Corticium
subincarnatum Peck, N. Y. State Mus. Rept. 42: 122. 1889;
Sacc. Syll. Fung. 9: 232. 1891.

Type: authentie specimen in Burt Herb., and specimen cited
by Karsten in Roumeguere, Fungi Gall., 4307.

Fruetifieations longitudinally effused, adnate, at first citron-
yellow, soon cinnamon-buff, even, pulverulent, the margin
thinning out, citron-yellow; in section 150-400 y thick, with the
hymenial layer and that next to the substratum usually slightly
colored, the hyphae suberect and branching or loosely interwoven,
thin-walled, hyaline, 3-4 y in diameter, nodose-septate, not in-
erusted except perhaps with a few minute grains in the sub-
hymenium; hymenium becoming 2-layered; cystidia hair-like,
thin-walled, even or with a few incrusting granules, 4-5 p in
diameter, protruding up to 40 џ; spores white in spore-collection,
4-5% x 2-215 u.

Fructifications 4-10 cm. long, 1-3 em. broad.

On decaying decorticated wood of Pinus and Abies in mountain

[Vol. 12
330 ANNALS OF THE MISSOURI BOTANICAL GARDEN

forests. Europe and northern United States and Canada west-
ward to Idaho and Manitoba. July to October. Uncommon.
This species may be recognized by its cinnamon-buff, yellow-
margined, closely adnate fructifications which occur, so far as
known at present, only on bare wood of spruce and pine. I
cannot understand how von Hóhnel & Litschauer, loc. cit., could
have regarded P. subsulphurea as perhaps not specifically distinct
from the Hannover specimens of P. radicata (P. filamentosa)
which have their hyphae heavily incrusted with a yellow matter
soluble in potassium hydrate solution, and much larger, more
incrusted cystidia, and fructifications only loosely adnate when
present on decorticated wood and margined with conspicuous
mycelial strands.
Specimens examined:
Finland: Mustiala, P. A. Karsten, authentic specimen of Xero-
carpus subsulphureus.
Sweden: L. Romell, 182.
France: Aveyron, A. Galzin, 21 033, comm. by H. Bourdot,
18426.

Canada: Lower St. Lawrence Valley, J. Macoun, 62.

New York: Cascade, C. H. Peck (in N. Y. State Mus. Herb.,
T. 31, and Mo. Bot. Gard. Herb., 56072); Clear Lake, G. F.
Atkinson, 5048; Floodwood, Е. A. Burt, C. H. Peck, 5, and an
unnumbered specimen (in N. Y. State Mus. Herb., and Mo.
Bot. Gard. Herb., 56017); North Elba, C. H. Peck, type of
Corticium subincarnatum (in N. Y. State Mus. Herb.).

Minnesota: Vermilion Lake, E. W. D. H. olway, 155, the Corticium
epichlorum of Geol. & Nat. Hist. Survey of Minn. but not C.
epichlorum Berk. & Curtis (in U. S. Dept. Agr. Herb. and Burt
Herb.).

Montana: Melrose, E. E. Hubert, comm. by J. R. Weir, 11431 (in
Mo. Bot. Gard. Herb., 63270).

Idaho: Priest River, J. R. Weir, 14.

Manitoba: Norway House, G. R. Bisby, 1478 (in Mo. Bot. Gard.
Herb., 61660).

99. P. martiana (Berk. & Curtis) Burt, n. comb.
Corticium martianum Berkeley & Curtis, Grevillea 1: 179.

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 331

1873; Peck, М. Y. State Mus. Rept. 30:48. 1879; 40:76. 1887;
Saec. Syll. Fung. 6: 633. 1888; Massee, Linn. Soc. Bot. Jour.
27:144. 1890.

Type: type distribution in Ravenel, Fungi Car. 5: 30.

Fructifications widely effused, rather thick, somewhat tuber-
cular or rugose, waxy, drying cinnamon to liver-brown and burnt
umber and so hard as to require prolonged moistening before
sectioning, the margin thinning out; in section 200—400 u thick,
colored like the hymenium, becoming dark vinaceous when
treated with potassium hydrate solution, 2-layered, with the layer
next to the substratum composed of longitudinally arranged,
honey-yellow hyphae 3 in diameter, and with the hymenial
layer thicker, denser, darker, and composed of densely inter-
woven hyphae and scattered cystidia; no gloeocystidia; cystidia
incrusted, conical, 30-45 x 8-12 u, wholly immersed or pro-
truding up to 30 u beyond the basidia; spores hyaline, even,
4416 x 2V$-3 y.

Fructifications 1-10 ст. long, 1-4 cm. wide.

On very rotten wood of frondose species—Betula and Populus
noted. Massachusetts to Alabama and in Ohio and Idaho.
September to November. Rare.

P. martiana is usually blood-red in color, with substance of
the same color, and with surface so tubercular or with so irregular
folds as to suggest a Phlebia. It is likely to be confused with
Phlebia hydnoidea Schw., from which it is sharply distinct by
the more toothed surface and slightly colored cystidia of the
latte

анн examined:

Exsiccati: Ravenel, Fungi Car. 5: 30, type distribution.
Massachusetts: Murray (in Curtis Herb., 6251).
New York: Ithaca, H. L. Jackson, comm. by Cornell Univ.

Herb., 18659; Keene Valley, Adirondack Mts., W. G. Farlow.
West Virginia: Eglon, C. G. Lloyd, 02678.

Florida: R. Thazter, 14 (in Mo. Bot. Gard. = ., 43934).
Alabama: Peters, in Ravenel, Fungi Car. 5

Ohio: College Hill, C. G. Lloyd, 1659.

Idaho: Coolin, J. R. Weir, 11116 (in Mo. Bot. Gard. Herb..

63251).

[Vol. 12
332 ANNALS OF THE MISSOURI BOTANICAL GARDEN

100. P. alutaria Burt, n. sp.

Type: in Burt Herb.

Fructifications effused, closely adnate, very thin, not at all
separable, clay color in the herbarium, somewhat lacunose in
some places, somewhat granular where thickest, the margin
thinning out; in section 120-160 р, thick, showing a little color
when only little magnified and giving the characteristic color to
the fructification but hyaline under high magnification, com-
posed of densely arranged, interwoven, suberect hyphae 3-314 y.
in diameter, not incrusted ; no gloeocystidia ; cystidia of two kinds:
cylindric, hair-like, flexuous cystidia 3-314 и in diameter, not
incrusted protrude up to 30 џ beyond the База and sometimes
have capitate tips; smaller incrusted cystidia 10 x 3 и are pres-
ent at surface of hymenium; basidia 4-spored; spores white in
spore collection, even, subglobose, 3-314 x 3 y.

Fragmentary pieces of fructifications are up to 5 cm. long,
2 cm. wide.

Type on wood of hardwood log of a frondose species in moun-
tain woods, also on Lariz. Vermont and Michigan. November.
Rare.

P. alutaria seems possible of recognition by its clay color,
closely adnate fructifications, and small spores and cystidia.

Specimens examined:

Vermont: Little Notch, Bristol, E. A. Burt, type.
Michigan: pole yard, Escanaba, C. J. Humphrey, 1788 (in Mo.

Bot. Gard. Herb., 42931).

101. P. separans Burt, n. sp.

Type: in Mo. Bot. Gard. Herb. and Dodge Herb.

Fructifications broadly effused, adnate, somewhat membra-
naceous, small pieces separable when moistened, between pale
ochraceous buff and avellaneous in the herbarium, even, some-
what cracked and showing the darker substance in the sides of
the fissures, the margin thinning out, slightly darker, some-
what radiately fibrillose, adnate; in section 300-350 и thick,
colored, stratose, each stratum 2-layered, the supporting layer
composed of densely and longitudinally interwoven, slightly
colored hyphae 3-314 и in diameter, the hymenial layer 75-

1925]
URT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 333

120 y. thick, composed of densely arranged, erect tissue; no gloeo-
cystidia nor conducting organs; cystidia incrusted, 40-50 x
8-15 в, numerous, immersed, starting from the base of the
hymenial layer; spores hyaline, even, 8-10 x 2-3 y.

Fructifieations probably large, for those studied are 4 cm.
long by 4 cm. wide and broken off on three sides.

On bark of coniferous log. British Columbia. September.

P. separans has some resemblance in color and aspect to
P. ciliata and resupinate Stereum sanguinolentum, but the stouter,
wholly immersed cystidia distinguish P. separans from the former
species, and the presence of cystidia and lack of conducting
organs from the latter. 'The type has two strata, the other
specimen only a single stratum of two layers.

Specimens examined:
British Columbia: Porcupine Creek, south of Beavermouth, C. W.

Dodge, 1702, type, and 1704 (in Mo. Bot. Gard. Herb., 58797,

58798, and in Dodge Herb.).

102. P. stratosa Burt, n. sp.

Type: in Mo. Bot. Gard. Herb.

Fructifications broadly effused, adnate, thick, stratose, some-
what cartilaginous-coriaceous, hard when dry, becoming pinkish
buff to light ochraceous buff in the herbarium, cracking in drying
and showing the stratose context, the margin thinning out; in
section 700 џ thick, pale yellowish, composed of 8 strata in the
type, with the hyphae hyaline, densely interwoven and con-
glutinate, about 2-214 y in diameter; cystidia incrusted, conical,
45-55 x 10-13 в, protruding up to 40 в, present in all strata but
more abundant and conspicuous in the outer half of the fructi-
fication and less distinct and perhaps becoming absorbed in the
more deeply buried strata; spores copious, hyaline, even, 4-5 X
2-2Y5 y.

Fructification 8 cm. long, 316 em. wide in the single piece con-
stituting the type, which has natural margin on one side only and
was broken from a larger mass.

On Quercus densiflora and Eucalyptus. California and Mexico.
September.

P. stratosa is related to P. similis but has larger cystidia and
spores.

[Vol. 12
334 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Specimens examined:

California: Pinehurst, E. E. Bethel, 26273 (in Mo. Bot. Gard.
Herb., 55437); Redwood Park, W. H. Long, 1851 4, type (in
Mo. Bot. Gard. Herb., 55065).

Mexico: A. Dampf, comm. by J. R. Weir, 63537 (in Mo. Bot.
Gard. Herb., 63710).

103. P. tabacina Burt, n. sp.

Type: in Burt Herb.

Fructifications effused, adnate, tawny olive to snuff-brown,
the hymenium becoming cracked and showing in the fissures the
concolorous subiculum, the margin thinning out, colored like the
hymenium; in structure 150—400 ų thick, tawny olive throughout,
2-layered, with the layer next to the substratum composed of
loosely interwoven, even-walled, colored hyphae 3-316 y in
diameter, nodose-septate, not incrusted, and the hymenial layer
about equal in thickness to the other, with its hyphae densely
crowded together in a palisade layer and bearing basidia and
sterigmata and containing some somewhat colored spores;
cystidia not incrusted, cylindric, 6-8 » in diameter, protruding
up to 80 џ; basidiospores hyaline, even, 6-9 x 214-3 и, copious;
slightly colored spores 9 x 3 џ are present in the deeper portion
of the hymenial layer of the type specimen.

Fructifications 2-9 cm. long, 1-214 em. broad.

On decaying coniferous wood and bark of logs. Wisconsin,
Colorado, Washington, and Oregon. July to November. Rare.

P. tabacina is distinguished by its tobacco color throughout and
hyphae and cystidia lacking incrustation. It lacks the radiate
filamentous margin of P. filamentosa of somewhat similar color
as well as the hyphal incrustation of the latter. The presence
of colored spores in the subhymenium is suggestive of Stereum
rugisporum, a species of the same color, occurring on coniferous
. Substrata in the same regions, and more abundant material may
show that P. tabacina is the thin, first-stratum stage of the
latter, but the fructifications at hand are closely adnate to the
substratum rather than loosely connected with it by the to-
mentose layer characteristic of many resupinate Stereums.

Specimens examined:

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 335

Wisconsin: Oconto Falls, C. J. Humphrey, 9445 (in Mo. Bot.
Gard. Herb., 57176).

Colorado: Ouray, C. L. Shear, 1185, type.

British Columbia: Agassiz, J. R. Weir, 330 (in Mo. Bot. Gard.
Herb., 63728); Sidney, J. Macoun, 19 (in Mo. Bot. Gard.
Herb., 5734).

Washington: Olympia, C. J. Humphrey, 6343; Seattle, C. J.
Humphrey, 6456; Sedro-Woolley, C. J. Humphrey, 7578 (in
Mo. Bot. Gard. Herb., 10753).

Oregon: Corvallis, on prune bark, Mrs. E. B. Zeller, comm. by
S. M. Zeller, 1871 (in Mo. Bot. Gard. Herb., 56872); Eugene,
C. J. Humphrey, 6096.

104. P. fusco-marginata Burt, n. sp.

Type: in Burt Herb. and probably in Lloyd Herb.

Fructifications long-effused, membranaceous, separable, be-
coming pinkish buff to warm buff in the herbarium, not waxy nor
cracked, the extreme margin byssoid, fuscous, colored like the
supporting hyphal layer next to the substratum; in section 300-
320 y. thick, colored next the substratum, 2-layered with (1) the
layer next to the substratum composed of longitudinally ar-
ranged hyphae 4-5 in diameter, not incrusted, not nodose-
septate, fuscous along the substratum, becoming colorless above,
and (2) the hymenial layer of equal thickness, composed of color-
less, erect hyphae somewhat granule-incrusted in an incrusted
zone; no gloeocystidia; cystidia not incrusted, 6 y in diameter at
base, tapering to the apex, protruding up to 30-40 р. beyond the
basidia; spores hyaline, even, 5-6 x 3-315 в.

Fructifications 1-10 em. long, the largest broken off at both
ends, 44-21% cm. wide.

On bark of fallen decaying frondose limbs. Florida and
Louisiana. June and July. ;

P. fusco-marginata has the unusual character of a colored layer
of coarse, fuscous hyphae running over the substratum and only
more or less completely covered by the buff, fertile portion of the
fructification, so that the protruding colored portion forms a dis-
tinctive fuscous margin. The Florida specimen is sterile and too
young for confident reference.

[Vol. 12
336 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Specimens examined:
Florida: Snapper Creek Hammock, W. A. M. urrill, 226, comm. by
N. Y. Bot. Gard. Herb. (in Mo. Bot. Gard. Herb., 62083).
Louisiana: St. Martinville, A. B. Langlois, 1947 and 1 00, type,
comm. by Lloyd Herb., 2771.

105. P. similis (B. & C.) Massee, Linn. Soc. Bot. Jour. 25:
147. 1889.

Corticium simile Berk. & Curtis, Linn. Soc. Bot. Jour. 10:337.
1868; Sacc. Syll. Fung. 6:631. 1888.

Type: in Kew Herb. and Farlow Herb., and a fragment in
Burt Herb.

Fructifications broadly effused, adnate, becoming light buff to
cream color in the herbarium, somewhat velutinous, cracked, the
margin thin; in section marguerite-yellow and darker next to
the substratum but with yellow color bleached by action of
potassium hydrate solution on the sections, 200—500 и thick in
the type but finally up to 2 mm. thick, composed of densely ar-
ranged, erect hyphae 3 џ in diameter, and of great numbers of
cystidia; cystidia incrusted, not colored, conical or fusiform,
15-25 х 6-8 в, very numerous in all regions; spores hyaline,
even, allantoid, 4 x 1 u, borne 4 to a basidium.

Fructifications “spreading for several inches." Fragmentary
specimens examined are 1—4 cm. in diameter.

On under side of frondose logs and fallen limbs. Florida,
Mexico, West Indies, and Japan. October to March. Prob-
ably common.

P. similis closely resembles Corticium portentosum in aspect,
and I am unable to distinguish it from the latter except by exam-
ination with the microscope which reveals the abundant, small,
colorless cystidia. Р. tephra is closely related but does not
form as thick fructifications, and its fructifications are less
cracked, darker-colored in section, with darker, thicker-walled,
more erect and more crowded hyphae, and slightly larger spores.

Specimens examined:

Florida: Cutler Hammock, W. A. M urrill, 63, comm. by N. Y.

Bot. Gard. Herb. (in Mo. Bot. Gard. Herb., 62093); Royal

Palm Hammock, W. A. Murrill, 112, 113, 119, 122, 125, all

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 337

comm. by N. Y. Bot. Gard. Herb. (in Mo. Bot. Gard. Herb.,
62094—62098, 62110).

Mexico: Guernavaca, W. A. & E. L. Murrill, 537, comm. by
N. Y. Bot. Gard. Herb. (in Mo. Bot. Gard. Herb., 54553);
Orizaba, W. A. & E. L. Murrill, 777, comm. by N. Y. Bot.
Gard. Herb. (in Mo. Bot. Gard. Herb., 54612); Xuchiles, near
Cordoba, W. A. & E. L. Murrill, 1211, comm. by N. Y. Bot.
Gard. Herb. (in Mo. Bot. Gard. Herb., 54600).

Cuba: C. Wright, 543, type (in Kew Herb., Farlow Herb., and
Burt Herb.), C. G. Lloyd, 432 (in Mo. Bot. Gard. Herb., 55171);
Alto Cedro, Earle & Murrill, 433, 553, both comm. by N. Y.
Bot. Gard. Herb.; Ceballos, C. J. Humphrey, 2678, 2813, 2834
(n Mo. Bot. Gard. Herb., 9087, 14855, 14837); Managua,
Earle & Murrill, 42, comm. by N. Y. Bot. Gard. Herb.; San
Diego de los Вайоз, Earle & Murrill, 203, 260, 303, all comm.
by N. Y. Bot. Gard. Herb.

Porto Rico: Rio Piedras, J. A. Stevenson & R. C. Rose, 6530 (in
Mo. Bot. Gard. Herb., 55072).

Bermuda: B. & J. Dodge, comm. by N. Y. Bot. Gard. Herb.

Jamaica: Cinchona, W. A. & E. L. Murrill, 598, 658, comm. by
N. Y. Bot. Gard. Herb.; Troy and Tyre, W. A. Murrill & W.
Harris, 886, 1053, in part, comm. by N. Y. Bot. Gard. Herb.

Grenada: Grand Etang, В. T'haxter, comm. by У. С. Farlow,
7, 8.

Japan: Mt. Tsukikuma, Prov. Bungo, A. Yasuda, 100 (in Mo.
Bot. Gard. Herb., 57018).

106. P. Seymouriana Burt, n. sp.

Type: type in Mo. Bot. Gard. Herb. and probably in Farlow
Herb.

Fructifications long and broadly effused, thin, closely adnate,
small portions separable when moistened, Verona brown to
mummy-brown or fuscous, somewhat velvety, cracking into
small areas, the margin determinate, entire; in structure 60-
180 y. thick, colored throughout like the hymenium, composed of
erect, colored, densely interwoven hyphae 3 y in diameter, not
incrusted, not nodose-septate, and of cystidia in all regions; no
gloeocystidia; hymenial surface velvety through very numerous

[Vol. 12
338 ANNALS OF THE MISSOURI BOTANICAL GARDEN

branched paraphyses having final branches 1 v in diameter;
cystidia incrusted, 20-35 x 12-15 y, usually wholly immersed;
spores not found.

Fructifications 12 em. long and broken off at ends, 3 em. wide.

On fallen decaying branches of undetermined frondose species.
Georgia and Cuba. August and April. Probably rare.

P. Seymouriana has general aspect suggestive of a resupinate
Hymenochaete or the effused stroma of an Н ypoxylon. The fruc-
tifications are thinner than those of P. tephra, with less numerous
cystidia and with the much darker hymenium becoming cracked
like that of Hymenochaete corrugata.

Specimens examined:

Georgia: Glen Ella, Tallulah Falls, A. B. Seymour, type, comm.

by Farlow Herb., G (in Mo. Bot. Gard. Herb., 44613).

Cuba: C. G. Lloyd, 145 (in Mo. Bot. Gard. Herb., 55495).

107. P. laevigata (Fr.) Massee, Linn. Soc. Bot. Jour. 25: 149.
Je. 1889; Karsten, Finska Vet.-Soc. Bidrag Natur och Folk 48:
426. O. 1889; Bourdot & Galzin, Soc. Myc. Fr. Bul. 28: 408.
1913; Rea, Brit. Basid. 696. 1922.

Thelephora laevigata Fries, Elenchus Fung. 1: 224. 1828.—
Corticium laevigatum Fries, Epicr. 565. 1838 ; Hym. Eur. 656.
1874; Басс. Syll. Fung. 6: 628. 1888.—Хегосатриз Juniperi
Karsten, Rev. Myc. 3°: 22. 1881.—Kneiffia laevigata (Fr.)
Bresadola, Ann. Myc. 1: 104. 1903.

Fructifications effused, thin, snuff-brown, drab, or pale drab-
gray, adnate, small pieces separable from the bark when moist-
ened, becoming cracked when dry, the margin at length free;
in section brown, 200 џ thick, composed of very numerous, colored
cystidia and thin-walled, hyaline hyphae 2-4u in diameter;
cystidia colored, cylindric-clavate or fusiform, 25-50 x 5-6 u,
thiek-walled and rough above or perhaps somewhat incrusted,
very numerous in all regions and giving their color to the trama
as & whole; spores hyaline, even, 7-8 x 3-4 u.

Fructifications 214-12 cm. long, 144-4 em. broad.

On bark of Juniperus. Canada, New York, and Europe.
April and September. Rare.

This species may be recognized by its occurrence on Juniperus,

1025]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 339

brown color within, and abundance of colored cystidia. European
authors record it on bark of living Juniperus communis, but the
data with the two American specimens which I have seen gave
merely the kind of substratum, one of these being J'uniperus
virginiana.
Specimens examined:
Exsiccati: de Thiimen, Myc. Univ., 2014, authentic specimen
from Karsten of Xerocarpus Junipert.
Sweden: L. Romell, 104, 105, 106; Femsjó, L. Romell, 407.
Finland: Mustiala, P. Karsten, in de Thümen, Myc. Univ., 2014.
Italy (?): locality not given, G. Bresadola.
England: Buckden, Yorkshire, E. M. Wakefield (in Mo. Bot.
Gard. Herb., 57119).
Canada: J. Macoun, 24.
New York: Orient, Long Island, R. Latham (in Mo. Bot. Gard.
Herb., 58907, and Burt Herb.).

108. Р. tephra (B. & C.) Cooke, Grevillea 8: 20. pl. 128, f. 6.
1879; Sace. Syll. Fung. 6: 643. 1888; Massee, Linn. Soc. Bot.
Jour. 25: 143. 1889.

Corticium tephrum Berk. & Curtis, Linn. Soc. Bot. Jour. 10:
336. 1868.

Type: in Kew Herb., and in Curtis Herb. mounted on left of
card, that on the right is Stereum albobadium.

Fructifications effused, adnate, between tilleul-buff and drab,
becoming drab in the herbarium, somewhat velutinous, the
margin thin, adnate, concolorous; in section brown throughout,
zonate, 400—550 р. thick, composed of erect, flexuous, thick-walled,
somewhat colored hyphae 3-4 y in diameter, densely crowded
together, and of very numerous cystidia; cystidia coarsely in-
crusted, conical, sometimes fusiform, 15-25 X 6-9 y, protruding
up to 9-12 y, not colored, very numerous, throughout the whole
fructification; spores hyaline, even, 5 Х 214-3 y.

Fructifications 2-6 ст. long, 14-2 cm. broad.

On dead wood of frondose species. Mexico, Cuba, Porto Rico,
and Bermuda. October to January.

Former accounts of P. tephra are erroneous because they were
partly based on a gathering of resupinate Stereum albobadium.

[Vol. 12
340 ANNALS OF THE MISSOURI BOTANICAL GARDEN

P. tephra belongs in the group with P. laevigata and P. pruinata

but does not have the colored cystidia of the former nor the

pruinose hymenium of the latter. The Australian specimen from

Berkeley under the name P. tephra, in N. Y. Botanical Garden

Herbarium, has colored cystidia and is more probably P. laevigata.
Specimens examined:

Mexico: Motzorongo, near Cordoba, W. A. & E. L. Миті,
997, comm. by N. Y. Bot. Gard. Herb. (in Mo. Bot. Gard.
Herb., 54623). ‘

Cuba: C. Wright, type (specimen in Curtis Herb. mounted on
left side of сага); Ceballos, C. J. Humphrey, 2692 (in Mo. Bot.
Gard. Herb., 21942); Ciego de Avila, Earle & Murrill, 592,
comm. by N. Y. Bot. Gard. Herb.; Herradura, Earle & Mur-
rill, 143, comm. by N. Y. Bot. Gard. Herb.

Porto Rico: Bayamon, J. A. Stevenson, 6760 (in Mo. Bot. Gard.
Herb., 55059).

Bermuda: Agricultural Station, H. H. Whetzel, Ak (in Mo. Bot.
Gard. Herb., 58909).

109. P. pruinata (B. & C.) Burt, n. comb.

Stereum pruinatum Berk. & Curtis, Linn. Soc. Bot. Jour. 10:
332. 1868; Sacc. Syll. Fung. 6: 583. 1888; Massee, Linn. Soc.
Bot. Jour. 27: 198. 1890.

Type: in Kew Herb. and Farlow Herb.

Effused, adnate, drying pale neutral gray to drab-gray,
pruinose, cracking when thick, the margin very thin; in section
fuscous throughout, becoming zonate and finally 1 mm. thick,
composed of densely arranged, erect, colored hyphae Зи in
diameter and of very numerous cystidia in all regions of the
section; cystidia incrusted, fusiform, 18-22 x 6-12 w; spores
— even, subglobose, about 3-414 x 216-3 in the few

ound.

Fructifications probably cover large areas, for those are 5-10
X 1-5 em. and fractured on 3 or all sides in the specimens seen.

On rotting hardwood logs. Florida, Alabama, Mexico, and
the West Indies. June to March. Occasional.

Dried specimens have the livid or cinereous color of some
forms of P. cinerea but with surface of rather more velvety tex-

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA 341

ture, often not cracked at all or, when cracked, into areas rangin

down to about 5 mm. in diameter. The fructifications of P.

pruinata are much thicker than those of P. cinerea and darker

throughout. When moistened, small pieces may be separated
from the bark for sectioning.
Specimens examined:

Florida: Cocoanut Grove, R. Thaxter, 77 (in Farlow Herb., and
Mo. Bot. Gard. Herb., 43897); Otter Creek, C. J. Humphrey,
6703 (in Humphrey Herb.); Palm Beach, H. von Schrenk (in
Mo. Bot. Gard. Herb., 43043).

Alabama: Montgomery County, R. P. Burke, 374 (in Mo. Bot.
Gard. Herb., 57242).

Mexico: Moian; near Cordoba, W. A. & E. L. Murrill, 988
(in Mo. Bot. Gard. Herb., 54621); Orizaba, W. A. & E. L.
Murrill, 764, in part, comm. by N. Y. Bot. Gard. Herb. (in
Mo. Bot. Gard. Herb., 54635).

Cuba: C. Wright, 193, type (in Farlow Herb., and Kew Herb.);
Alto Cedro, Santiago de Cuba Province, Earle & Murrill, 516,
518, 544, 555, comm. by N. Y. Bot. Gard. Herb.; Ceballos,
C. J. Humphrey, 2815.

Porto Rico: Mount Morales, near Utuado, Mrs. E. G. Britton &
D. W. Marble, 1204, comm. by N. Y. Bot. Gard. Herb. (in
Mo. Bot. Gard. Herb., 61486).

Jamaica: Hope Gardens, W. A. Murrill, 2, comm. by N. Y.
Bot. Gard. Herb.; Moneague to Union Hill, W. A. Murrill,
1176, comm. by N. Y. Bot. Gard. Herb.

110. P. rimosissima (B. & C.) Burt, n. comb.

Corticium rimosissimum Berkeley & Curtis, Am. Acad. Arts
& Sci. Proc. 4: 124. 1858; Sacc. Syll. Fung. 6: 639. 1888;
Massee, Linn. Soc. Bot. Jour. 27: 122. 1890.—An Stereum
umbrinum Berk. & Curtis?

Type: type distribution in C. Wright, Plants of U. S. North
Pacific Expl. Exp., 110.

Fruetifieations broadly effused, rather thick, dry, mem-
branaceous, separable in rather large pieces, pliant when dry,
now bister in the herbarium, not shining, even, cracking through
the colored hymenium into polygonal masses 1-4 to a mm. and

[Vol. 12
342 ANNALS OF THE MISSOURI BOTANICAL GARDEN

showing the underlying pale substance, the true margin un-
known; in section 360—450 џ thick, colored in the hymenial layer,
with the basal layer composed of obliquely ascending, loosely
interwoven, thin-walled, hyaline hyphae 3-4 v in diameter, not
inerusted, not nodose septate, and of thick-walled, non-staining,
hyaline organs 414 џ in diameter, not incrusted, whose pointed
tips protrude as cystidia up to 12 y beyond the basidia; spores
hyaline, even, 6 x 414 u—few found and may not belong.

Fragmentary fructifications not having margin are 4 cm. long,
2 cm. wide.

On dead eane. Nicaragua.

P. rimosissima is closely related in color and structure to
Stereum umbrinum but has colorless cystidia not incrusted and
only 414-6 u in diameter, and thinner fructifications which are
not known yet to occur reflexed.

Specimens examined:

Nicaragua: C. Wright, type (in U. S. Dept. Agr. Herb.).

111. P. Weiri Bresadola, Mycologia 17: 70. 1925.

Type: in Weir Herb.

Fructifications long and broadly effused, thin, closely adnate,
becoming cream-buff to chamois in the herbarium, even, some-
what cracked, the margin thinning out; in section 150 y. thick,
concolorous with, and giving the color to, the fructification,
composed of densely interwoven, rigid, slightly colored hyphae
2-314 u in diameter, not incrusted ; gloeocystidia flexuous or
sometimes filamentous, 30-75 x 3-5 ш; cystidia not incrusted,
thin-walled, cylindric, obtuse, 6-8 u in diameter, protruding up
to 40-50 u beyond the basidia, not numerous; basidia with 4
sterigmata ; spores hyaline, even, cylindric, 6-8 x 3-314 p,
copious.

Fructifications 5-12 cm. long, 2-4 em. wide.

On wood of decaying logs of Pinus monticola. Idaho. Sep-
tember.

The gloeocystidia of P. Weiri are unusual in their position,
since they are occasionally oblique or parallel with the substratum,
and more elongated then than when in the more usual, erect
position, nor did they become visible in my sections stained with

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 343

eosin until the sections have cleared somewhat in the permanent

glycerine mount. The color of the densely interwoven tissue of

the fructification should aid in recognition of the species.
Specimens examined:

Idaho: Priest River, J. R. Weir, 23345, type (in Weir Herb.).

112. P. Farlowii Burt, n. sp.

Type: in Burt Herb.

Fruetifieations effused, closely adnate, rather thick, pale
olive-buff in the Бойын even, somewhat cracked and showing
the tissue to be horn-like and somewhat resin-colored (pecan-
brown) where exposed on sides of the fissures, the margin thin-
ning out, composed of finely interwoven hyphae; in section 250—
350 u thick, somewhat colored, inclosing some portions of the
substratum, composed of densely interwoven and conglutinate
hyphae 2-3 in diameter, not incrusted, not nodose-septate,
indistinct; no gloeocystidia; cystidia incrusted, 30-70 x 12-15 и,
protruding up to 30 y, few and scattered; spores hyaline, even,
4 x2 y.

Fructifications in fragments 2-3 cm. long, 2 cm. wide.

On very rotten frondose wood. New Hampshire. September.

P. Farlowii shows in the dried specimen a pale olive-buff
hymenium covering a horn-like, somewhat resin-colored under-
lying layer; the cystidia are so large as to be a good distinctive
character.

Specimens examined:

New Hampshire: Chocorua, Bowditch Swamp, W. G. Farlow, 16,
type.

113. P. colorea Burt, n. sp.

Type: in Burt Herb.

Fructifications longitudinally effused, very thin, closely adnate,
light drab, not shining, even, the margin thinning out, indeter-
minate; in section 70-80 џ thick, light drab, 2-layered, with a
layer sone the substratum about 30 thick, of densely longi-
tudinally interwoven, somewhat colored hyphae about Зи in
diameter, indistinct, conglutinate, and with a colored hymenial
layer of erect basidia, paraphyses, and cystidia; no gloeocystidia;

[Vol. 12
344 ANNALS OF THE MISSOURI BOTANICAL GARDEN

cystidia incrusted, slightly colored, fusiform, 24-33 x 12-15 Uy
few, immersed in the hymenial layer; spores of a crushed prep-
aration cylindric, hyaline, even, curved, 8-10 x 2-3 и.

Fructifications 3-9 em. long, 1-114 em. wide.

On bark of dead branches about 1-14 em. in diameter, of
frondose species. Louisiana. December.

P. colorea belongs near the P. cinerea group of very variable
species. It may well prove that P. colorea is not a specifically
distinct member of this group when more abundant material
from southern Louisiana is available, but it seems to me distinct
now by the longitudinal layer next to the substratum, light drab
color throughout, few, large, slightly colored cystidia which are
confined to the hymenial layer, and by the slender, elongated
spores.

Specimens examined:

Louisiana: St. Martinville, A. B. Langlois, ch, type.

114. P. decorticans Burt, n. sp.

Type: in Burt Herb.

Fructifications long-effused, closely adnate, very thin, growing
on the wood, spreading longitudinally and laterally between the
wood and bark, loosening the latter, pale pinkish buff and pale
gull-gray to whitish, pruinose, with occasional tubercules in some
specimens; in section brownish throughout, 50-90 » thick, not
zonate, composed of densely arranged, interwoven, slightly
colored, erect hyphae 3 y in diameter, with no darker and opaque
zone next to the substratum; cystidia few, incrusted, ovoid to
subglobose, up to 20-25 x 15 и, seen only in the region next to
the substratum; paraphyses with slender, antler-shaped branches
protrude from hymenium; spores hyaline, even, slightly curved,
8-9 X 3 u, few seen.

Fructifications 1-2 em. wide, 2 em.—6 m. long, on under side of
dead branches along which the loosened bark eurls back laterally.

On Quercus Garryana, Acer macrophyllum, and Rhus diversi-
loba. Washington and Oregon. February to December. Com-
mon locally. =

P. decorticans differs from P. cinerea, P. nuda, P. caesia, and
P. violaceo-livida in not being so dark as to be opaque next to the

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 345

substratum. Its most noteworthy character, by which it may
be recognized at a glance, is its curious habit of forming the
fructification on bark-covered limbs between the bark and the
wood, so that the loosened bark—very noticeable on Quercus
limbs—curls back, disclosing the fructification closely adnate on
the wood. The antler-shaped branching paraphyses occur in
P. phyllophida also.
Specimens examined:
Washington: Bingen, W. N. Suksdorf, 910, type, 756, 757, 758.
Oregon: Corvallis, C. Epling (in Mo. Bot. Gard. Herb., 60183),
S. M. Zeller, 1769, 2258 (in Mo. Bot. Gard. Herb., 56846,
63028).

115. P.nuda (Fr.) Bresadola, I. R. Accad. Agiati Atti III. 3:
114. 1897; Bourdot & Galzin, Soc. Myc. Fr. Bul. 28: 405.
1913; Rea, Brit. Basid. 695. 1922.

Thelephora nuda Fries. Syst. Мус. 1: 447. 1821.—Corticium
nudum Fries, Epier. 564. 1838; Patouillard, Tab. Anal. Fung.
2: 33. f. 582. 1887; Sacc. Syll. Fung. 6: 626. 1888.—Penio-
phora ochracea Masses; Linn. Soc. Bot. Jour. 25: 150. 1889,
but not Corticium ochraceum Fries.

Illustrations: Patouillard, loc. cit.

Fructification effused, closely adnate, very thin, pale drab-
gray, pale purplish gray or pale gull-gray, pruinose, waxy, crack-
ing in drying; in section brownish, darker and opaque next the
substratum, 75-160 и thick, the hyphae densely interwoven,
rather erect, Зи in diameter, somewhat colored; cystidia in-
crusted, in all regions of the fructification, usually about 20-25
X бы, larger near the substratum and sometimes up to 15 y in
diameter; spores hyaline, even, curved, 4144-9 x 214-3 y, re-
ported larger by European authors.

Fructifications 2-6 x 1-2 cm.

On fallen limbs of frondose species such as Acer, Quercus,
Populus, etc. Canada to Texas, in Europe and Japan. April

7 :

I have seen no authentic specimens of P. nuda, but the fnm
pean concept of this species differs from P. cinerea in having the
fructifications more whitish gray in color, more broadly effused,

[Vol. 12
346 ANNALS OF THE MISSOURI BOTANICAL GARDEN

and less evidently formed by confluence of several small fructi-

fieations and with some cystidia near the substratum of greater

diameter than those elsewhere. I have seen no spore collections,

and it is possible that the spore measurements given above are

too small, since they are based on spores found in preparations of

sections. |
Specimens examined:

Exsiccati: Ravenel, Fungi Am., 454, under the name Corticium
ochraceum.

Canada: Ottawa, J. Macoun, 26.

Vermont: Middlebury, E. A. Burt.

New York: Alcove, C. L. Shear, 1306; Altamont, E. A. Burt.

New Jersey: Newfield, J. B. Ellis (in N. Y. Bot. Gard. Herb.,
and Mo. Bot. Gard. Herb., 61338).

Maryland: Takoma Park, C. L. Shear, 1358.

Virginia: C. L. Shear, 1181.

South Carolina: Pinopolis, in Ravenel, Fungi Am., 454.

Georgia: Atlanta, E. Bartholomew, $981 (in Mo. Bot. Gard.
Herb., 63459).

Florida: Daytona, R. A. Harper, 5 (in Mo. Bot. Gard. Herb.,
54538).

Alabama: Auburn, F. S. Earle (in N. Y. Bot. Gard. Herb., and
Mo. Bot. Gard. Herb., 63415).

Louisiana: Baton Rouge, C. W. Edgerton, 830.

Texas: Beaumont, C. J. Н umphrey, 5936.

Japan: Province Bungo, N. N akayma, comm. by A. Yasuda, 125
(in Mo. Bot. Gard. Herb., 59471).

116. P. argentea Ellis & Everhart in herb., n. sp.

Type: in N. Y. Bot. Gard. Herb. and Mo. Bot. Gard. Herb.

Fructifications effused, closely adnate, thin, pallid mouse-gray
to drab-gray, pruinose, cracked in drying, the margin darker and
thinning out; in section brown and opaque with exception of the
hyaline hymenial layer, 150 џ thick, with the hyphae densely
interwoven, thick-walled, stiff, 3-314 p in diameter, colored as in
Hymenochaete, not incrusted; cystidia not incrusted, partially
destroyed and rendered nearly invisible by potassium hydrate
solution, tapering upward to a point, protruding up to 30 p,

1925]
URT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 347

6-7 y in diameter, often colored for 20 џ at the base and there
with the aspect of buried setae; basidia deteriorated; no spores
found.

Fructifications 4-8 cm. long, 1-11 em. broad.

On bark and decorticated wood of decaying Fraxinus. Louisi-
ana. January. Probably rare.

This species has the color and aspect of P. nuda and P. caesia
but differs from both of these and also from P. cínerea in having
its opaque basal layer 120 џ thick, comprising the whole thickness
of the fructification except the hymenium, and in having its
hyphae thick-walled and distinct and colored as in Hymeno-
chaete. The cystidia differ from those of the species just named
and also P. pruinata in not being incrusted and are noteworthy
by being attacked and partially dissolved by 7 per cent solution
of potassium hydrate to such a degree that they are best studied
when sections are mounted in lactic acid.

Specimens examined:

Louisiana: St. Martinville, A. B. Langlois, 1758, type (in N. Y.

Bot. Gard. Herb., and Mo. Bot. Gard. Herb., 63416).

117. P. violaceo-livida (Sommf.) Bresadola in Bourdot &
Galzin, Soc. Мус. Fr. Bul. 28:405. 1913; Rea, Brit. Ваза. 695.
1922.

Thelephora violaceo-livida Sommerfelt, Fl. Lapp. Suppl. 283.
1826.—Corticium violaceo-lividum (Sommf.) Fries, Epicr. 564.
1838; Hym. Eur. 655. 1874; Басс. Syll. Fung. 6: 627. 1888.

Fructifications somewhat effused, closely adnate, rather thick,
tubercular, pale mouse-gray to drab-gray, often round; in section
brownish, 100-300 р. thick, becoming zonate within, darker and
opaque next to the substratum, the hyphae somewhat colored,
densely arranged, erect; cystidia incrusted, 20-30 X 6-9 u, dis-
tributed in all regions, very numerous; spores hyaline, even,
curved, 6-9 x 2144-4 в, as found with sections.

. Fructifications 14 x 144-2 cm., often with the component
masses rounded, 5-7 mm. in бизан.

On fallen limbs of Salix, Prunus, Fraxinus Castanea, and
Quercus. Canada to Louisiana. March to October. Rare.

'The concept of this species presented by Bresadola, which has

348 ANNALS OF THE MISSOURI BOTANICAL GARDEN

become generally accepted in Europe, is followed here except

that I have referred to this species effused fructifications with

tuberculate surface, thick and zonate within, as well as fructi-

fications consisting of aggregations of small, round masses. The

specimen received from Bresadola has the latter form and is on

Prunus Cerasus; one from Romell on Saliz, the substratum first

cited for the species, has a similar zonate structure within and a

tubercular surface but is more effused than that from Bresadola.
Specimens examined:

Lappland: Sommerfelt, authentic specimen under. the name
Thelephora fallax var. violaceo-livida (in Herb. Fries). _

Sweden: L. Romell, 71. |

Austria: Hall in Tirol, V. Litschauer.

Italy probably: locality not stated, G. Bresadola.

Canada: Ontario, Ottawa, J. Macoun, 27, 131.

Vermont: Middlebury, E. A. Burt, two gatherings. |

Massachusetts: near Boston, E. A. Burt.

New Jersey: Newfield, J. B. Ellis (in Mo. Bot. Gard. Herb.,

9).

Maryland: Takoma Park, C. L. Shear, 1027.

District of Columbia: Soldiers Home, C. L. Shear, 1116.

Louisiana: Baton Rouge, Edgerton & Humphrey, comm. by C. J.
Humphrey, 2521.

118. P. cinerea (Pers.) Cooke, Grevillea 8: 20. pl. 123, f. 8.
1880; Saec. Syll. Fung. 6: 643. 1888; Bourdot & Galzin, Soc.
Myc. Fr. Bul. 28: 407. 1913; Rea, Brit. Basid., 696. 1922.

Corticium cinereum Persoon, Roemer Neues Mag. Bot. 1:
111. 1894; Fries, Epicr. 563. 1838; Hym. Eur. 654. 1874.—
Thelephora cinerea § Corticium Persoon, Syn. Fung. 579. 1801;
Myc. Eur. 1:148. 1822; Fries, Elenchus Fung. 1:221. 1828.—
Kneifía cinerea (Fr.) Bresadola, Ann. Мус. 1: 103. 1903.—
Corticium fumigatum de Thümen, Torr. Bot. Club Bul. 6: 95.
1876; Myc. Univ., 513. 1876.—Thelephora lilacina Schweinitz,
Am. Phil. Soc. Trans. N. S. 4: 168. 1832.—Peniophora lilacina
(Schw.) Massee, Linn. Soc. Bot. Jour. 25: 147. 1889.

Illustrations: Fries, Icones Hym., pl. 198, f. 4; Cooke, loc. cit.;
Patouillard, Tab. Anal. Fung. f. 251.

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 349

Fructifications effused, closely adnate, very thin, in small
patches becoming confluent, lurid, ashy in various shades as pale
drab-gray, pale mouse-gray, and cinnamon-drab, pruinose, waxy,
becoming cracked in drying; in section 50-100 џ thick usually,
brownish, darker and opaque near the substratum, the hyphae
densely interwoven, 3 y in diameter, somewhat colored; cystidia
inerusted, 25-40 x 414-9 u, distributed throughout the section;
spores hyaline, even, cylindric, 6-9 x 2-3 в, borne 4 to a ba-
sidium.

Fructifications 2-5 x 144-1 cm.; when scattered 2-5 mm. in
diameter.

On fallen limbs of Alnus, Acer, Prunus, Pyrus, Quercus, and
most other frondose and coniferous species. "Throughout North
America, West Indies, Europe, southern Africa, and Japan—
probably cosmopolitan. Our commonest species. Throughout
the year.

P. cinerea may be recognized by its resemblance to a thin coat
of ashy gray or slightly tinted paint on the bark of fallen limbs;
the substance of the sections is brownish when viewed with a
hand lens, and dark and opaque next the substratum under the
compound microscope. P. caesia, P. nuda, and P. violaceo-livida
must be cautiously separated from P. cinerea, for all are closely
related.

Specimens examined:

Exsiccati: Berkeley, Brit. Fungi, 63, 64; Ellis, N. Am. Fungi, 21,
under the name Corticium fumigatum, 610; Ell. & Ev., Fungi
Col., 610, 805, under the name C. fumigatum; de Thimen,
Myc. Univ., 513, type distribution of C. fumigatum, 1206;
Sydow, Myc. Germ., 205.

Sweden: L. Romell, 69, 70.

England: in Berkeley, Brit. Fungi, 63, 64; Kew Gardens, E. M.
Wakefield (in Mo. Bot. Gard. Herb., 57121).

Germany: Brandenburg, in Sydow, Myc. Germ., 205; Berlin,
P. Magnus (in N. Y. State Mus. Herb., and Mo. Bot. Gard.
Herb., 55803).

Austria: Lengerich, Brinkmann, comm. by G. Bresadola; Tirol,
three specimens, comm. by V. Litschauer.

Italy: Trento, G. Bresadola; Vallambrosa, Cavara, comm. by
G. Bresadola.

[Vol. 12
350 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Newfoundland: Bay of Islands, A. C. Waghorne, 989 (in Mo. Bot.
Gard. Herb., 5009).

Canada: J. Macoun, 8, 9, 50.

Quebec: Hull, J. Macoun.

Ontario: London, J. Dearness, 169a, 169c (in Mo. Bot. Gard.
Herb., 11350, 5629); Ottawa, J. Macoun, 334.

Maine: Portage, L. W. Riddle.

New Hampshire: Chocorua, W. G. Farlow, 147 (in Mo. Bot.
Gard. Herb., 55262) and three specimens in Burt Herb.;
North Conway, A. S. Rhoads, 8 (in Mo. Bot. Gard. Herb.,
56977), W. H. Snell, 627 (in Mo. Bot. Gard. Herb., 59294).

Vermont: Middlebury, E. A. Burt, nine gatherings.

Massachusetts: Arlington, E. A. Burt, A. P. D. Piguet, comm.
by W. G. Farlow (in Mo. Bot. Gard. Herb., 43959); Billerica,
E. A. Siegler (in Mo. Bot. Gard. Herb., 55035); Boston, E. A.
Burt; Stoneham, C. L. Shear, 1239.

Connecticut: Portland, G. P. Clinton (in Mo. Bot. Gard. Herb.,
43945).

New York: Albany, H. D. House (in N. Y. State Mus. Herb.,
and Mo. Bot. Gard. Herb., 15955, 57517, 59673, 59690,
59695), L. O. Overholts, 3388 (in Mo. Bot. Gard. Herb., 6989),
C. H. Peck (in N. Y. State Mus. Herb., and Mo. Bot. Gard.
Herb., 55987, 57516, 57518); Alcove, C. L. Shear, 248, 1006,
1100, 1138, 1216, 1300; Carrollton, C. H. Peck (in Mo. Bot.
Gard. Herb., 56012); East Galway, E. A. Burt; Greenbush,
C. H. Peck (in N. Y. State Mus. Herb., 74, and Mo. Bot. Gard.
Herb., 55776); Hudson Falls, S. H. Burnham, 19 (in Mo. Bot.
Gard. Herb., 54504); Ithaca, б. F. Atkinson, 674, 8218, Н. 8.
Jackson, Cornell Univ. Herb., 14394, C. O. Smith, comm. by
G. F. Atkinson, 8223; Karner, H. D. House (in N. Y. State
Mus. Herb., and Mo. Bot. Gard. Herb., 54369); Knox, C. H.
Peck (in N. Y. State Mus. Herb., and Mo. Bot. Gard. Herb.,
95751); Menands, C. H. Peck (in N. Y. State Mus. Herb., and
Mo. Bot. Gard. Herb., 55805); Middle Grove, E. A. Burt;
Van Cortland Park, New York City, C. H. Peck (in N. Y.
State Mus. Herb., and Mo. Bot. Gard. Herb., 55977) ; Orient,
R. Latham, 181 (in Mo. Bot. Gard. Herb., 44227); Selkirk,
C. H. Peck (in N. Y. State Mus. Herb., and Mo. Bot. Gard.

1925]
BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 351

Herb., 55773); Van Etten, Tioga County, W. C. Barbour,
1365 (in N. Y. Bot. Gard. Herb., and Mo. Bot. Gard. Herb.,
61400); West Albany, C. H. Peck (in N. Y. State Mus. Herb.,
and Mo. Bot. Gard. Herb., 55749); Westport, C. H. Peck (in
N. Y. State Mus. Herb., and Mo. Bot. Gard. Herb., 55750);
White Plains, L. M. Underwood (in N. Y. Bot. Gard. Herb.,
and Mo. Bot. Gard. Herb., 61588); Willsboro Point, C. O.
Smith; West Fort Ann, S. H. Burnham, 17 (in Mo. Bot. Gard.
Herb., 44046).

New Jersey: Newark, H. S. Jackson; Newfield, J. B. Ellis (in
Mo. Bot. Gard. Herb., 4818), 1076, 1078, comm. by W. G.
Farlow (in Mo. Bot. Gard. Herb., 14762, 7459), comm. by
N. Y. Bot. Gard. Herb. (in Mo. Bot. Gard. Herb., 61448), in
Ellis, N. Am. Fungi, 21, 610, in Ell. & Ev., Fungi Col., 610,
805, and de Thümen, Myc. Univ., 513, 1206; Belleplain, C. L.
Shear, 1165. -

Pennsylvania: Bethlehem, Schweinitz, type of Thelephora lilacina
(in Farlow Herb. and Kew Herb.).

Maryland: Takoma Park, C. L. Shear, 962, 1028, 1076, 1162, 1849.

Distriet of Columbia: Takoma Park, C. L. Shear, 515 (in N. Y.
State Mus. Herb., and Mo. Bot. Gard. Herb., 55808), 1353;
Washington, C. L. Shear, 1200, 1258.

Virginia: Park Lane, W. H. Long, 18509 (in Mo. Bot. Gard.
Herb., 55061).

North Carolina: Blowing Rock, G. F. Atkinson, 4328, 8030.

Georgia: Atlanta, E. Bartholomew, 5676 (in Mo. Bot. Gard.
Herb., 44252).

Florida: New Smyrna, W. A. Murrill, 5, comm. by N. Y. Bot.
Gard. Herb. (in Mo. Bot. Gard. Herb., 62085).

Alabama: Auburn, F. S. Earle, unnumbered specimens and 42
(in N. Y. Bot. Gard. Herb., and Mo. Bot. Gard. Herb., 61399,
61452), and F. S. Earle & C. F. Baker (in N. Y. Bot. Gard.
Herb., and Mo. Bot. Gard. Herb., 61428); Montgomery and
Montgomery County, R. P. Burke, 9, 11, 42, 120, 449, 456,
458, 461, 469, 513, 818 (in Mo. Bot. Gard. Herb., 16360, 22340,
21100, 19555, 57277, 57280, 57283, 57288, 57303, 63117).

Louisiana: Baton Rouge, Edgerton & Humphrey, 5727a, 5666.

Tennessee: J. R: Weir, 7558 (in Mo. Bot. Gard. Herb., 55464).

[Vol. 12
352 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Ohio: Norwood, C. G. Lloyd, 1576.

Indiana: Crawfordsville, A. R. Bechtel, 12 (in Mo. Bot. Gard.
Herb., 59660) ; Millers, E. T. & S. A. Harper, 939.

Illinois: Barry, Н. W. Anderson (in Mo. Bot. Gard. Herb., 55966);
Cypress, C. J. Humphrey, 1359 (in Mo. Bot. Gard. Herb.,
22522); River Forest, E. T. & S. A. Harper, 676, 757; River-
side, E. T. & S. A. Harper, 677.

Michigan: Ann Arbor, C. H. Kauffman (in N. Y. Bot. Gard.
Herb., and Mo. Bot. Gard. Herb., 61396); Gogebie County,
E. A. Bessey, 56, 78, 188, 216, 236, 371 (in Mo. Bot. Gard.
Herb., 56545, 56549, 56580, 56546, 56590, 56635); Michigan
Agricultural College, B. O. Longyear (in Mo. Bot. Gard. Herb.,
55704); New Richmond, E. W. Hartwell (in Mo. Bot. Gard.
Herb., 58163); Vermilion, A. H. W. Povah, 242 (in Mo. Bot.
Gard. Herb., 58163).

Wisconsin: Blue Mounds, comm. by Univ. Wis. Herb., 28;
Madison, E. Bartholomew 6652 (in Mo. Bot. Gard. Herb.,
57039), M. C. Jensen, comm. by C. J. Humphrey, 2432 (in
Mo. Bot. Gard. Herb., 4835), and W. T'release (in Mo. Bot.
Gard. Herb., 4816, 43988, 43989).

Minnesota: Lake Itaska, E. L. Jensen, 5 (in Mo. Bot. Gard.
Herb., 12530); Univ. Farm Campus, St. Paul, E. L. Jensen, 3
(in Mo. Bot. Gard. Herb., 4203).

Missouri: Columbia, B. M. Duggar, 572, 574; Creve Coeur Lake,
L. O. Overholts, 3159 (in Mo. Bot. Gard. Herb., 5714).

Nebraska: Lincoln, C. L. Shear, 1054, 1058, 1342.

Colorado: Golden, E. Bethel & L. O. Overholts, 1744 (in Mo. Bot.
Gard. Herb., 54870).

Manitoba: Winnipeg, A. H. R. Buller, comm. by G. R. Bisby,
878, and G. R. Bisby, 1348 (in Mo. Bot. Gard. Herb., 58995,
and 60554 respectively).

British Columbia: Salmo, J. R. Weir, 444 (in Mo. Bot. Gard.
Herb., 6243); Sidney, J. Macoun, 6, 775 (in Mo. Bot. Gard.
Herb., 5765, 55324).

Washington: Bingen, W. N. Suksdorf, 700, 701 ‚ 721, 744, 759,
861, 885, 918, 954, 960, 963; Corvallis, S. M. Zeller, 2262 (in
Mo. Bot. Gard. Herb., 63033) ; Chelan, J. R. Weir, 5490 (in
Mo. Bot. Gard. Herb., 58260) ; Kalama, C. J. Humphrey, 6219;

1925]
URT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 353

Washougal, E. H. T'urk, comm. by S. M. Zeller, 2630 (in Mo.
Bot. Gard. Herb., 63057).

California: Berkeley, comm. by W. A. Setchell, 1032 (in Mo. Bot.
Gard. Herb., 44241); Stanford University, C. F. Baker, 12;
Sierra Nevada Mountains, W. H. Harkness, 1025 (in Kew
Herb., under the name Peniophora carnea Berk. & Cke.).

Mexico: Guernavaca, W. A. & E. L. Murrill, 858, 407 (in N. Y.
Bot. Gard. Herb., and Mo. Bot. Gard. Herb., 54470, 54532).

Cuba: San Antonio de los Bafios, Havana Province, Earle &
Murrill, 73, comm. by N. Y. Bot. Gard. Herb.

Porto Rico: Rio Piedras, J. A. Stevenson, 2451, 2920, 3067, 5581,
5638 (in Mo. Bot. Gard. Herb., 9185, 3125, 9055, 6957, 54585).

Jamaica: Chester Vale, W. A. & E. L. Murrill, 334, comm. by
N. Y. Bot. Gard. Herb.; Cinchona, W. A. & E. L. Murrill,
696, comm. by N. Y. Bot. Gard. Herb.; Troy and Tyre,
W. A. & E. L. Murrill, 894, comm. by N. Y. Bot. Gard.
Herb.

Africa: Stellenbosch, Cape Colony, P. A. van der Bijl, 326 (in Mo.
Bot. Gard. Herb., 63397).

Japan: Mt. Mikuma, Province Awaji, A. Yasuda, 4 (in Mo. Bot.
Gard. Herb., 55666).

119. P.caesia Bresadola in Bourdot & Galzin, Soc. Myc. Fr.
Bul. 28: 406. 1913; Rea, Brit. Basid., 695. 1922.

Corticium caesium Bresadola, Fungi Trid. 2: 39. pl. 145, f. 2.
1892; Sacc. Syll. Fung. 11: 126. 1895.

Illustrations: Bresadola, loc. cit.

Type: authentic specimen in Burt Herbarium.

Fructifications broadly effused, closely adnate, very thin, pale
mouse-gray to pale purplish-gray, delicately pruinose, cracked in
drying, the margin similar; in section brownish, 40-90 џ thick,
dark and opaque next to substratum; hyphae densely interwoven,
somewhat colored; cystidia near the substratum 15-25 x 10-20 y,
inerusted, becoming slightly colored, not numerous; spores hy-
aline, even, curved, 6-8 х 214-3 u as found in preparations of
sections, probably larger in spore falls.

Fructifications 2-10 x 1-2 em.

On fallen limbs of Syringa, Betula, Quercus, and other frondose

[Vol. 12
354 ANNALS OF THE MISSOURI BOTANICAL GARDEN

species. Vermont to District of Columbia, in Missouri, and in

Europe. March to December. Rare.

P. caesia is more widely effused than P. cinerea, is not formed
by confluence of many small fructifications, and has much the
color and aspect of P. nuda but differs from the latter in absence
of the numerous, small буз

Specimens examin
Exsiccati: Наше, Fungi Gallici, 2910, under the name

Corticium incarnatum, 3213, under the name Corticium cinereum.
Austria: Vienna, comm. by V. Litschauer.

Italy: Trient, G. Bresadola, authentie specimen.

France: in Roumeguére, Fungi Gallici, 2910, 3213.

Vermont: Lake Dunmore, E. A. Burt.

Distriet of Columbia: Washington, Department Grounds, on
Syringa vulgaris, C. L. Shear, 1264, in part, and an unnumbered
specimen.

Missouri: Columbia, B. M. Duggar, 448.

120. P. carnea (Berk. & Cooke) Cooke, Grevillea 8: 21. pl.
124, f. 11. 1879; Басс. Syll. Fung. 6: 644. 1888; Massee, Linn.
Soc. Bot. Jour. 25: 151. 1889.

Corticium carneum Berkeley & Cooke, New York Acad. Sci.
Ann. 1: 179. 1878; Linn. Soc. Bot. Jour. 17: 141. 1878.

Type: in Kew Heb.

Fructification effused, closely adnate, thin, ochraceous flesh-
color, drying avellaneous and cracked, the margin whitish and
fibrillose; in section brownish, 100-120 и thick, with a dark,
semi-opaque zone next to the substratum; hyphae densely inter-
woven, 3-314 u in diameter, slightly colored, somewhat longi-
tudinally interwoven next to the substratum; cystidia incrusted,
of two kinds—very large cystidia resembling conical or subglobose
crystalline masses 45-75 x 30-75 и are seated on the opaque
zone, other cystidia 25-35 х 6-8 џ are scattered throughout the
region between the dark zone and the surface of the hymenium;
gloeocystidia flexuous, 40-50 x 4-414 u, not numerous; spores
hyaline, even, slightly curved, 8-12 x 3-4 в

Fructifications 1-6 em. long, 12-2 cm. Ко.

On logs and fallen, decaying, frondoselimbs. Texas and Cuba.
March. Rare.

1925]
URT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 355

The thin, closely adnate fructifications of P. carnea, brownish
within and with a broad, dark, opaque zone next to the sub-
stratum, place this species in the P. cinerea group. It is remark-
able by having, in addition to the ordinary kind of cystidia, very
much larger cystidia which finally become, by the accretions of
mineral matter, very large masses of mineral nature with very
coarse grains on the exterior of the mass. In the Cuban gathering
which I have referred to this species, when a small portion of the
hymenial surface was moistened with alcohol and then with
water preparatory to removal of a bit of the fructification for
sectioning, the moist hymenium became punctate with minute
depressions, probably by presence at those points of the large
buried cystidia. This may prove a useful test for preliminary
sorting out, without examination by the microscope, of the rare
P. carnea from the more common P. cinerea of nearly similar
aspect. P. heterocystidia has cystidia of two kinds, like those of
P. carnea but thicker, readily separable from the substratum
when moistened, and with a narrow brown zone in the middle of
its sectional preparations and with a loosely interwoven hyaline
zone next to the substratum. The specimen in Kew Herbarium,
collected on fir in the Sierra Nevada Mountains, California, by
Harkness, 1025, and referred by Cooke to P. cinerea does not
have the large cystidia of his type and is P. cinerea instead.

Specimens examined:

Texas: Galveston Bay, H. W. Ravenel, 78, type (in Kew Herb.).
Cuba: San Diego de los Baños, Pinar del Rio Province, Earle &
Murrill, 333, comm. by N. Y. Bot. Gard. Herb.

SPECIES TOO INCOMPLETELY DESCRIBED FOR LOCATION AMONG
PRECEDING SPECIES

Peniophora convolvens Bresadola, Ann. Myc. 18: 48. 1920.

**Elongato-effusa, ceraceo-membranacea, pallida vel avellanea,
ambitu similari, demum libero-convoluto; hymenio demum late
rimoso, interstitiis fibrillosis; sporis hyalinis, obovatis, 6-7 x 5-
б ш; basidiis clavatis, 40-45 x 6-7 ш; cystidiis saepe immersis
vel usque ad 45 u prominentibus, 9-12 y. crassis.

“Нар. ad ligna, St. Croix, Americae centralis. Raunkiaer."

[Vol. 12

356 ANNALS OF THE MISSOURI BOTANICAL GARDEN

P. gigaspora Massee, Linn. Soc. Bot. Jour. 25: 152.
Басс. Syll. Fung. 9: 238. 1891

1889;

“Latissime effusa, ambitu fimbriata albicans; hymenio pallido,
velutino, sicco indurato, contiguo; cystidia fusoidea, 80-120 x
30-40 u; sporae oblongo-ellipsoideae, 18-20 x 10 y.

“N. Providence, Bahamas.

*On decorticated wood, forming thin, continuous, broadly
effused patches, somewhat resembling P. velutina, but differing in

cystidia and spores."

(To be concluded)
INDEX TO SPECIES OF PENIOPHORA

Page Page
Wer NRA Seas eee 304 с RS ME
т, RENE ENE SUS OS apa Ee 226 fI. o eee ES AE 320
AMIE Ч 208 AIMÉ. o s a пана 276
АИША Qe ONES ERI MO Sum. o И 252
BI S ea hae eek ES 297 favido-alba.. aiio rere 248
и... 228 Е 244
alboustraminea os. ck у fusco-marginata.............. 335

SOME EE oc Sf а 5065656
ABRE. i gs лу ууа 301 и. ек аса 216

а C res 332 а LL eee eee Be
ПИОН; V or LS TELE 216 PUTS o oues Ce CIR 245

с. Ll IL LL es cass 220 о ое
о ее 346 globifera. ...... eres 219

BIDUIDUSER о... 230 ск.
SNO V LV eee ees 310 heterooystidía. ..............- 293
boreali... o o os S ST 295 RERO oie Ecco 272
о 282 т, .. v roc 225
nx CE SR Ga 218 ИО ОИ а
ME а КЖ 353 ШОН 6 055s А ЕР 221
SER S су а ул CE Dre e А P 267
о 260 о LII LII re PY as ed

ИЛИШ IUE а EAR. X d 226 с г сс ow cere TRES
TENES a Ou Un dE 354 Па: 253
И o oss 254, 286

SUME S oS IC in 327 о ООО ОТО МУТ

о С IDEE 253 :

DONDE o coo РАО А see
танне WE ANE uo v A 257
к. 355 laminata . 946
pose RUM er а. 286 [Uc omen d M eee
COMER. S Ba VLL ILLINS 261 lepida. РАО АКСАРА 296
са 344 ООШ ir E

disciformis милы - МЧА eL SONUS гу EPI es
uer ic cals В и 275

DMO. АИИ

9 еее ено CC eae NM RON буе d qo

1925]

BURT—THE THELEPHORACEAE OF NORTH AMERICA. XIV 357

Page
И 238
а (Ч РЕНАТА 330
medioburiensis............... 328
DERI ОНА 243
WING Oe ТН К С а 277
ОН а. 270
О. 237
In Iovi wore O d
DU И ВАРТИ 345
ODORE I EEE Ee CL 345
a A уе 223
ОЙОН СОЛУУ AE AR E UE E. 289
ТИИ ERE TEE E EPI a 291
ур, 315
pace ЕО ИА rode
о RL Par ee Sno ary Gs s
bete TR UNA LU st ALLE CAEL ESE E НИИ 242
ИМА C LI PILLS LAUR
ргаеіеттівга.................... 316
ИЕЛДЕ а iri ERRARE
т Е ЕО Corea cee 313
ооо. 313
ВАО VENTER IDEA 269
rimosissima , . .................. 841
BOSE С ard 270

Page
ieptooystdia. О ce tx và 260
SCIRE Ус бузуу CE di Eat do. 2
Seymouriana................. 3
SRM ei ea s ERA XN 268
SUPE РУО аала 336
NONI а ДЕ 280
о PETE Я В И 333
ТАДДАУ БУ И А СЕ 288
Му 280
ас АННА
РМО УССР 259
ть ДАТА A 329
ОРИ r.i douce inh 324
Tabu dA oov Lad ee 334
TuS C rU Uses i CR CE 306
Dd iios улла ei C
BU КИО О UU CE TY 317
We roi LA a de CUR M 339
КОЛИ ERAS FAAVAE MO.
ip Ps EO ARO EA Y DESI A KDE 251
ИР 236
о НО РНЕ 319
СИТИ ИИА 320
рост О 264
а 250
РРО ko es Vike X 305
IEE ded OLAS PE күл саа 322
м 342
ЗИ нь 245

Annals
of the
Missouri Botanical Garden

Vol. 12 NOVEMBER, 1925 No. 4

THE EFFECT OF TREATING THE VIRUS OF TOBACCO
MOSAIC WITH THE JUICES OF VARIOUS PLANTS

B. M. DUGGAR
Physiologist to the Missouri Botanical Garden, in Charge of Graduate Laboratory
Professor of Plant Physiology in the Henry Shaw School of Botany of
# Washington University

AND
JOANNE KARRER ARMSTRONG
Formerly Research Assistant to the Missouri Botanical Garden

During the period that the mosaic disease of tobacco has been
under investigation in this laboratory, many attempts have been
made to transfer the disease, through cross inoculation, to the
pokeweed, Phytolacca decandra. Five operators have at different
times endeavored to make the transfers referred to, each em-
ploying à somewhat different technique. In no instances have
results been obtained that might be interpreted as a positive
indication of the susceptibility of the pokeweed to the tobacco
form of mosaic.

Although at that time no experiment had been made here in
the direction of testing the possibility of transfer through insect
agency, it seemed quite probable that the pokeweed might be
immune. It occurred to the junior author that it might be of
interest to determine the effect of the juice of the pokeweed on
the tobaeco virus. Accordingly, a preliminary experiment was
arranged, and it was determined to include in this the effect upon
the tobacco mosaic agency of both the juice from healthy poke-
weed and that from pokeweed affected with the mosaic peculiar
to that host.

Juiee or erude sap was then prepared from (1) healthy poke-
weed, (2) pokeweed affected with mosaic, and (3) tobacco plants

ANN. Mo.B or. Garp., Vor. 12, 1925 (359)

[Vor. 12
360 ANNALS OF THE MISSOURI BOTANICAL GARDEN

affected with tobacco mosaic. In each case the procedure was as
follows: The younger shoots and leaves were ground up in a meat
or food grinder and the resulting semi-fluid mass was thrown into
a muslin bag and the juice pressed out. The juice was then fil-
tered through absorbent cotton. The undiluted filtered juice
was then.used as indicated later. Adequate precautions were
taken as to the cleanliness of all vessels involved. The meat
grinder and other articles required were, after preliminary
cleaning, soaked several hours in hot water containing 16 per
cent formaldehyde, and afterwards boiled in plain water. In
these and in later experiments where pipettes and other glass-
ware were required, such vessels were invariably dry-sterilized.

In the preliminary experiment, 1 part of the tobacco juice from
diseased plants was mixed with 99 parts of each of the 2 types of
pokeweed juice, and as a partial control on the dilution and the
infectivity of the virus, 1 part of the same diseased tobacco juice
was added to 99 parts of sterile distilled water. Further control
was not used at the time owing to a scarcity of suitable plants
for the inoculation work. The fluids as diluted for inoculation
were permitted to stand 3-4 hours prior to their use. The tech-
nique of inoculation is described later. Twenty healthy young
tobacco plants were inoculated with each of the 3 lots of material,
4 cc. of inoculum being used for each lot. The inoculations were
made June 12, and all of the 20 control plants (diseased juice 1
part, water 99 parts) were diseased by June 23, while none of the
40 plants inoculated with the diseased tobacco juice made up in
pokeweed juice showed the least indication of mosaic on June 30,
when the experiment was closed. These results were so definite
that it was determined to repeat the work and to use a variety of
plant juices when opportunity offered.

The method of inoculation employed in the preliminary experi-
ments and in that subsequently reported is one that has been
adopted as a standard in this laboratory and it requires a brief
description. It is preferred to employ young plants with stems
about 2 inches high. When possible, the plants to be inoculated
are arranged in place about 1 week in advance so that sporadic
Occurrences of the disease may be promptly detected. Three

inoculations are made in éach plant, 1 on the stem near the ground,

1925]
DUGGAR & ARMSTRONG—TOBACCO MOSAIC VIRUS 361

another farther up and in a leaf axil—so as to reach the young
bud—and the other in the immediate region of the terminal bud.
The quantity of inoculum required is about 2 cc. for each 10
plants. A large drop of the inoculum may be placed with a glass
rod on the spot desired, and with a small inoculating needle many
prieks are made through this, thus working the fluid into the
tissue. Before passing to the next plant the needle is flamed, then
dipped in alcohol, and burned off. The usual precaution is
taken with respect to touching plants, or if plants must be
touched, the hands are cleansed between operations. Because
no disease has appeared in any lot of plants, it is not safe to
assume freedom from disease, so that every plant in any lot is
treated as though it might convey disease. Inoculated plants
are kept under observation for 28-30 days in all cases where more
than preliminary data are required.

Preparing the crude sap as already described, experiments were
arranged with shoots and leaves of pokeweed, Jimson weed
(Datura Stramonium), geranium (Pelargonium sp.), cotton, and
squash, with Irish potato tubers, sweet-potatoes, and apples of
the variety Ben Davis. In each case the juice from diseased
tobacco plants was diluted, as shown in table I, with the juice
from the plant the influence of which was to be tested. In these
experiments, the mixed juices were allowed to remain at room
temperature about 2 hours, and then placed in a refrigerator,
at 3° C., until used the following morning (15-18 hrs.). The
undiluted j juices used as control were similarly exposed.

From the table it is again clear that pokeweed juice effectively
inactivates the agency of tobacco mosaic in a relatively short time.
Even when the virus is diluted only 5 times with the pokeweed
juice the inhibition is complete. Inactivation is shown by the
juice of Jimson weed when the latter is in relatively high concen-
tration, and the juice of the geranium is also to some extent ef-
fective. On the other hand, cotton, squash, potato, sweet-
potato, and apple exert no injurious influence at the concentra-
tions tested.

It became of much interest to determine if the influence of
pokeweed juice especially might result from some relatively
simple chemical factor, or if it might be far more complex, possibly

[Vor. 12
362 ANNALS OF THE MISSOURI BOTANICAL GARDEN

analogous to agglutination. The possibility that the reaction of
the juice might be an important factor suggested itself, but
colorimetric tests made it clear that the H-ion concentration was

TABLE I
EFFECT OF VARIOUS PLANT JUICES xs THE PATHOGENICITY OF Ыы
TOBACCO MOSAIC VIRUS. TEN PLANTS WERE INOCULATED I
EACH eed
Juices constituting inoculum Dilution Number diseased
Diseased tobacco, dist. water 1:10 All (in 15 days)
Diseased tobacco, dist. water 1:100 Ал a 11 days)
Diseased tobacco, pokew 1:5
Diseased tobacco, weed 1:10 Hone
Diseased tobacco, pokeweed 1:25 None
Diseased tobacco, pokew 1 None
Diseased tobacco, pokeweed 1:75 None
Diseased tobacco, pokeweed ер None
Pokeweed None 1 (after 4 wks.)
(apparently accidental)
Diseased кымы, Jimson 1:10 АП (in 9 days)
Diseased tobacco, Jimson | : 100 None
Jimson None None
Diseased tobacco, Irish potato 7109 АП (in 9 days)
Diseased tobacco, Irish potato — 1:100 АВ n 9 days)
Irish potato None
Diseased tobacco, sweet-potato 1:10 All ( (in 11 days)
tobacco, sweet-potato : 100 9
Dionne ри b. er J Gn 15 days)
o пао geranium : in
Diseased tobacco, geranium F 5
Erina y tob 1 err АП Gi 14 days)
Diseased tobacco, apple T2 in ys
Diseased tobacco, apple 1:100 irem ws days)
Apple None
Diseased tobacco, tobacco : 10 All "in 12 days)
tobacco, tobacco | : 100 All (in 12 days)
Healthy whee o gyr Nu: 13 die)
acco, cotton 0 in ys
Diseased tobacco, Dae w oe in 13 даун)
: ,8q 2 in ys
Diseased tobacco, squash 1:100 All (in 9 days)
Squash None None

higher in the apple juice than in any other tested, and this last
exerted no injurious influence. Moreover, experiments then in
progress, to be reported elsewhere, indicated a relatively high
tolerance of th virus t ls. Inview of the follow-
ing facts, namely, that pokeweed is a host for a form of mosaic,
that Jimson weed is credited with several forms of mosaic, and

‘Employing electrometric methods, a subsequent test made by Mr. Е. В. Ranker

and Miss Fanny Fern Smith gave a Py of approximately 6.0 for the juice of the
pokeweed used.

1925]
DUGGAR & ARMSTRONG—TOBACCO MOSAIC VIRUS 363

that even geranium exhibits a disease tentatively classed as a
mosaic, the influence of the juices of these plants must be re-
garded as specific with reference to the tobacco mosaic agency.
However, in order to determine if pokeweed juice possesses
general germicidal properties, cultures in small Erlenmeyer
flasks were arranged with sterile and unsterile (natural) juice,
and both lots were inoculated with Aspergillus niger. The fungus
grew promptly and profusely on both lots, indicating no general
inhibiting effect.

The toxicity of pokeweed juice was further tested, using as an
indicator the growth of Bacterium prodigiosum Lehm. & Neum.
In carrying out the experiments with this organism a strong bac-
terial emulsion was prepared from a fresh nutrient agar slant
culture. This emulsion was then added to pokeweed јшсе,! or
to an equal quantity of distilled water, as control. These mix-
tures were allowed to stand for 2 hours, after which suitable di-
lutions were prepared and plates were poured. Every possible
care was taken with the samples employed in the preparation of
the dilution cultures to take the sample from the vessel in such
manner that the pipette did not come in contact with the walls
of the vessel where organisms might occur which had not had
free access to the fluid used. In the first series of cultures the
original bacterial emulsion was diluted (1) 1:5 with pokeweed
juice; (2) 1:25 with pokeweed juice; and (3), for control, 1: 25
with distilled water. The results are given in detail in table п.

The result of the foregoing test is convincing proof that no
toxicity of pokeweed juice is exhibited toward this species of
bacteria. In explanation of the table it should be pointed out
that the data with the 2 dilutions of pokeweed juice are closely
comparable, while the result from dilution with distilled water
shows apparently a very much smaller number of bacteria
present. This, however, was anticipated and is in harmony with
results obtained from a preliminary series not here reported in
detail. This preliminary series brought out the fact that, on

1 The pokeweed juice used in these experiments was prepared as previously de-
scribed except that in this case, in order to minimize contaminations, the pokeweed
leaves and shoots were first treated for 4 hours with 20 per cent Javel water, after

were washed in sterile distilled water.

[Vor. 12
364 ANNALS OF THE MISSOURI BOTANICAL GARDEN

TABLE II

EFFECT OF POKEWEED JUICE ON BACTERIUM PRODIGIOSUM; EXPOSURE
TO JUICE 2 HOURS

Plate No. colonies Average.
Treatment dilution 4 days ' | colonies per ec.
1/10 Numerous
Bacterial emulsion diluted 1/100 768
1:5 with pokeweed juice 1/1000 39 | 51,980
1/10000 4
1/10 960
Bacterial emulsion diluted 1/100 108 | 10,800
1:25 with pokeweed juice 1/1000 12 ;
Bacterial emulsion diluted 1/10 71
1 : 25 with distilled water 1/100 14 | 1,055
1/1000 0

standing, an emulsion of this bacterium in distilled water shows
an apparent decrease in the number of organisms. It is not the
purpose of this paper to determine the cause of this decrease
though it may be a simple aggregation phenomenon. For veri-
fication of this observation another experiment was carried out,
the results of which are briefly given in table ш. In this experi-
ment the bacterial emulsion was diluted, as will be seen, with
distilled water and with pokeweed juice to the same extent.
Plates were poured immediately after the bacterial emulsions
were diluted, and then after intervals of 1 and 2 hours respectively.

TABLE III
BACTERIUM PRODIGIOSUM IN DISTILLED WATER AND IN POKEWEED
JUICE
: No. colonies, No. colonies,
Treatment No. colonies, after exposure of | after exposure of
ves 1 hour 2 hours
Bacterial emulsion diluted
1:25, distilled water 20,300 —— 7,050
emulsion diluted
1:25, pokeweed juice 29,850 44,050 42,500

No discussion of table пт is required further than to point out
again the diminution, on standing, of the number of colonies in
the case of the emulsion diluted with distilled water. The figures
for the emulsion diluted with pokeweed juice are sufficiently
comparable when it is recalled that this is not filtered juice, and

1925]
DUGGAR & ARMSTRONG—TOBACCO MOSAIC VIRUS 365

accordingly represents a minute suspension, in itself a sufficient
cause for some variation in numbers.

These experiments were further suggestive of a specific in-
activating effect of the pokeweed juice towards the agency of
mosaic disease. The cases of geranium and of Jimson weed
have not been more closely analyzed.

It seemed desirable to determine the possible relation of the
larger colloidal particles in the pokeweed juice to inactivation,
but up to the present only a few preliminary experiments have
been made. These, however, are not without suggestion.
Pokeweed juice was filtered through a cylindrical, porcelain
atmometer cup under a pressure of one-half atmosphere, and the
filtrate thus obtained was used in treating the mosaic virus sub-
sequently used in inoculations, as in table 1. When the relation
of virus to filtrate was 1 : 10, the incidence of infection was 9
out of 10 plants inoculated; when the relation was 1 : 100, the
virus was completely inactivated. Filtration through such a
filter seems therefore to reduce the effect of the juice. It may
be stated that the filter employed permits the mosaic virus to
pass through but does not permit Bacterium prodigiosum.

Some of the same lot of pokeweed juice was diluted with an
equal quantity of distilled water and then centrifuged for 30
minutes at 1500 revolutions per minute. The effect of the super-
natant liquid on the virus was determined, as before, by inocu-
lating 10 tobacco plants. With the relation of the diseased juice
to this diluted liquid 1:10, 2 plants developed the disease, again
indicating some loss of inactivation capacity.

It seemed conceivable that inactivation of the mosaic virus
might be due to adsorption. It was not possible to determine
this merely by increasing relatively the quantity of diseased to-
bacco juice, since this dilutes the pokeweed juice. Concen-
tration of the diseased tobacco juice by evaporation would also
be subject to criticism from another angle. Further work on
the chemical or physical nature of this inactivation is planned.

For the moment the data in the 3 experiments given in table ту
show further the influence of the ratio of diseased juice to poke-
weed juice as developed through infection experiments, and thus
supplement table 1. The experiments were made under con-

[Vor. 12
366 ANNALS OF THE MISSOURI BOTANICAL GARDEN

ditions similar to those reported in table r, but at a different time.
The progressive increase in the incidence of disease with the
increasing amount of virus locates rather definitely, in conjunc-
tion with the data of table 1, the inactivation capacity of the
pokeweed juice.

TABLE IV
Juices constituting inoculum Dilution Result
Diseased tobacco, pokeweed 1:7 2 diseased (in 8 days)
Diseased tobacco, pokeweed 5:1 8 diseased (in 14 days)
Diseased tobacco, pokeweed 10:1 9 diseased (in 9 days)

Upon the completion of the experiments with pokeweed juice
reported in table r, the results were regarded, and apparently
properly so, as a simple case of inactivation. Nevertheless, it
was considered desirable to ascertain if in the plants used in
experiments 3-8 inclusive there might exist such a mild form of
the disease as to be essentially without external symptoms. Ас-
cordingly, leaves of these plants were used directly in the inocu-
lation of fresh young tobacco plants, but without visible result.
Likewise, one-half (5) of each lot of plants used in experiments
3-8, table 1, were inoculated with diseased tobacco juice, un-
treated, and the entire lot of 30 plants exhibited the characteristic
mosaic symptoms in less than 15 days, indicating, as anticipated,
not the least suggestion of a modification of susceptibility to
mosaic.

The specific nature of this inactivation of the tobacco mosaic
virus, especially by pokeweed juice, has seemed of sufficient im-
portance to justify the projection of further experiments in the
direction of ascertaining the physieal and chemical properties of
this juice.

DETERMINATION OF TOTAL NITROGEN IN PLANTS
AND PLANT SOLUTIONS: A COMPARISON OF
METHODS WITH MODIFICATIONS

EMERY R. RANKER
Rufus J. Lackland Fellow, Missouri Botanical Garden

The analyst who has had considerable experience in the deter-
mination of nitrogen is impressed by the fact that some sub-
stances are analyzed with more ease and accuracy than other
substances. Many factors are involved, not the least of which
is the combination of the various forms of nitrogen in the sample
to be analyzed. For example, if we consider the determination
of total nitrogen in whole plants (peas, wheat, barley, ete.) in-
cluding the residual nutrient solutions in which they grew,
amino-, amide-, and some ammonia-nitrogen are present in the
plants in addition to nitrate- and possibly ammonia-nitrogen in
the residual solutions. In such a case the determination of total
nitrogen presents problems that challenge the accuracy of the
various methods used.

Two methods are used generally: (1) some modification of the
Devarda method, and (2) the official salicylic-thiosulphate
method (Assoc. Off. Agr. Chemists, '21, p. 8; I, 28). The latter
method has been criticized severely by several investigators whose
data have been interpreted as indicating the limitations and
defects of any method based on the reduction of nitrates in acid
medium. Further reference will be made to these criticisms after
the presentation of data. On the other hand, the Devarda
method for total nitrogen in the presence of organic matter is
time-consuming, since (1) there is a preliminary alkalin e distil-
lation with the alloy to collect the ammonia obtained from the
reduction of the nitrates and other substances acted upon in the
process (Allen, "15; Davisson, '18), and (2) the organic matter
and remaining nitrogen must be subjected to a Kjeldahl digestion
followed by a second distillation into the same or a second lot of
standard acid. Unfortunately, then, neither of these methods is
entirely satisfactory for the determination of total nitrogen in
plants and plant solutions when the two are present in the same
sample.

In a study of nitrogen fixation by higher plants the total ni-
trogen content of the plants and residual nutrient solutions was

Axx. Mo. Bor. Gard., VoL. 12, 1925 (367)

[Vor. 12
368 ANNALS OF.THE MISSOURI BOTANICAL GARDEN

determined. The official salicylic-thiosulphate method was used.
Extremely inaccurate results were obtained. From nutrient
solutions containing respectively 400, 300, 200, 100, 50, 25, and
10 mgs. of nitrogen per 950 cc., only 68, 67, 30, 62, 62, 54, and
27 per cent of the nitrogen was recovered. From solutions con-
taining smaller quantities of nitrogen than 10 mgs. the amounts
recovered were usually less than for the blank, that is, there was
a loss of nitrogen from the reagents used. A total of about 60
determinations was made, and there was no agreement among
the results obtained. The greatest losses of nitrogen occurred
from those samples containing whole plants in addition to the
residual nutrient solutions. In several cases there was a visible
evolution of nitrogen dioxide fumes, and duplicate determinations
varied widely. These data tend to corroborate the statements of
those investigators who have criticized the official salicylic-
thiosulphate method.

In spite of these criticisms, however, it was thought advisable
to attempt some modifications that might overcome the dif-
ficulties encountered. Several possibilities were tried, and from
the results of these tests it was tentatively determined that the
inaccuracy of the method was due, primarily, to the presence of
water at some stage during the process of acid digestion. Certain
details of manipulation seemed to influence the determination to a
limited degree.

A modification of the official method was devised, and its ac-
curacy for the determination of total nitrogen was tested out on
the various forms of nitrogen. For purposes of comparison, simul-
taneous determinations on samples from the same stock, measured
by the same pipettes and at the same temperature, were made
by & modification of the Devarda method. The procedure for
this comparison method was as follows: (The procedure for the
modified official method will be given later.)

I. If organic matter is not present.—Place the sample in an 800-сс.

jeldahl flask; add 1.0 gm. of Devarda alloy for every 60-70 mgs. of
nitrate-nitrogen present!; add a small piece of paraffin; make up to
' Subsequent to this investigation the work of Burrell and Phillips (’25) has been

1925]
RANKER—DETERMINATION OF TOTAL NITROGEN IN PLANTS 369

a total volume of 150 cc.; add 6 cc. of 10 per cent sodium hydroxide;
connect to the distillation apparatus and distill into standard acid at
slow boiling for 1 hour. Titrate the standard acid to neutrality.

J organic matter is present.—Proceed as in I; continue distil-
lation to as low a volume as is safe; disconnect. Add 45 cc. of con-
centrated suphuric acid and 10 gms. of anhydrous sodium sulphate;
digest for 1 hour after the copper color appears (the mixture remains
milky due to aluminum and zinc precipitates); cool; make up to an
estimated volume of 400 cc. and distill as for the modified official

All the determinations reported were made of the following
forms of nitrogen:

1. Nitrate-nitrogen as NaNO; which was twice recrystallized
from Merck's blue-label grade. "The solution used was such that
1 ce. was equivalent to 1 mg. of nitrogen.

2. Ammonia-nitrogen as (NH,4)4SO, which was recrystallized
from Merck's blue-label grade. The solution used was such that
1 сс. was equivalent to 1 mg. of nitrogen.

3. Amino-nitrogen as glycine, an Eastman Kodak Company
product. The solution used was such that 1 cc. was equivalent
to 0.91 mg. of nitrogen.

4. Amide-nitrogen and amino-nitrogen as asparagine, a Merck
product. The solution used was such that 1 cc. was equivalent
to 0.95 mg. of nitrogen.

5. Total organie plant nitrogen as found in 6-day-old wheat
seedlings; 10 such seedlings produced 7 mgs. of nitrogen with but
little variation from this average. It would have been some-
what more accurate had smaller seeds been used, in which case
the larger number used would have appreciably decreased the
variation from the average.

'The acid and alkali used in titration were standardized against
benzoie acid obtained from the U. S. Bureau of Standards (sample
№. 39B). А total of 190 determinations, including the prelimi-
naries, was made. The results obtained were subjected to a
statistical analysis, the data of which, expressed as average
percentages, are given in table r. The probable error of the
arithmetical mean was calculated by the formula

(2-255)

(Vor. 12

370 ANNALS OF THE MISSOURI BOTANICAL GARDEN
TABLE I
RECOVERY OF TOTAL NITROGEN
(average percentages and probable error)
Recovery b Recovery by
Devarda CA PS modified. oH
Sample determined Sample | Sample | Sample Sample o e
vae ри» M не
n t
to dryns. Е to TRA WM
50 mgs. nitrate-N. 97.7 98.4 100.4 62.2 8
+.19 +.03 +.06 +.38
100 mgs. nitrate-N. 68.5* 72.4* 99.2 17.8 13
99.91 +.10 +.81 3
+.24
50 mgs. nitrate-N. | 98.5 99.4 98.0 36.7 12
10 mgs. ammonia-N. + .23 +.22 +.06 1.80
50 mgs. nitrate-N. | 97.7 ‚ 99.4 99.3 51.7 12
10 mgs. amino-N. +.14 +.04 +.14 +4.03
50 mgs. nitrate-N. | ar 25-20 S is Б 15
- =. >: =: d.
10 mgs. Vende:
50 mgs. nitrate-N. | 99.6 98.8 100.2 64.6 12
7 mgs. plànt-N.f +.09 +.09 +.32 4.23
50 mgs. nitrate-N.
ese samples were not determined, as during evapo-
: EM i PSU. N.T | ration heavy NO; fumes were given ой.
50 mgs. nitrate-N. | 100.8 99.7 14
7 mgs. plant-N.f || +.56 +.69
50 eia ects i2 99.1 6
7m К | + .52 + .62
1 се. EXP N aOH
50 mgs. nitrate-N. | 98.1 99.5 92.7 33.2 10
0.5 gm. sucrose + .64 +.00 +.90
* 1.0 gm. of Devarda alloy used.
1 Supplied as the imis content of ten 6-day-old wheat seedlings.
12.0 gms. of Devarda allo

oy
|| This solution was adjusted to АН prior to determination.

From a study of the data of table 1 certain facts are evident:

1. The presence of free water in the sample is the determining
factor for the accuracy of a method dependent upon the reduction
of nitrates in aeid medium (the minimum amounts of water that
may be present and not decrease the accuracy of such a method

1925]
RANKER—DETERMINATION OF TOTAL NITROGEN IN PLANTS 371

have not been determined—it is recommended, however, that
no more than just a trace of free water be present).

2. When the sample is practically dry the official method as
here modified is somewhat more accurate for the determination
of total nitrogen than the Devarda method used.

3. The modified method is aceurate for the determination of
amino-, amide-, ammonia-, nitrate-, and total plant-nitrogen and
combinations of these forms of nitrogen in plants and plant
solutions.

4. If sugar is present in abundance a slight loss of nitrate-
nitrogen may occur, due to the reducing action of the sugar. This
loss would be very slight in actual practice since the nitrate-
nitrogen content of plants is small.

The procedure for the modified official method as used in this
investigation is as follows:

Place the sample in an 800-cc. Kjeldahl flask; adjust to neutrality
or make slightly alkaline; if water is present evaporate just to dryness
on a water bath under vacuum. Add 35-40 cc. of salicylic acid mix-
ture (1.0 gm. of salicylic acid to 30 cc. of concentrated nitrogen-free
sulphuric acid); mix thoroughly and allow to stand for at least an
hour with occasional shaking (if organic matter is present, stopper
tightly with a rubber cork and allow to stand over night). Add 5
gms. of sodium thiosulphate and heat for 5 minutes with a low flame;
cool; add 7-10 gms. of anhydrous sodium sulphate and a pinch of
copper sulphate. Digest for an hour at the boiling point after the
solution clears; just before the solution solidifies dilute to an estimate
volume of 400 сс.; cool completely. Add a small piece of paraffin,
100 сс. of a saturated solution of sodium hydroxide, and a piece of
mossy zinc; connect immediately to the distillation apparatus and
distill 150-200 cc. over into standard acid during a period of 1 hour.
Titrate the standard acid to neutrality with standard alkali and cal-

the amount of nitrogen present.

Subsequent to this investigation 380 determinations were
made by the above method with an accuracy equivalent to that
here reported. Some of the samples determined had a total
nitrogen content as high as 400 mgs., in which case the amount of
salicylie acid mixture was increased to 45 cc. Some of the
samples determined had a total nitrogen content of 2 mgs. per
950 ec. of solution; the solutions (950 cc. in the sample) were
evaporated just to dryness on a water bath under vacuum and
an accurate recovery was obtained. Most of the determinations

[ Уот, 12
372 ANNALS OF THE MISSOURI BOTANICAL GARDEN

were made on samples composed of whole plants of wheat, barley,
or peas, plus the residual nutrient solutions in which they grew.
If this modified method be in error all attempts to locate that
error have failed and any suggestions or criticisms are welcome.

Attention to detail is essential to accurate determinations; for
this reason it is well to mention a few details of manipulation that
have been found of value:

1. It is the habit of some analysts to wash down the neck of
the Kjeldahl flask during the process of digestion; in the interests
of personal safety and certainty of results this should be avoided.
Such a practice introduces water into the sample and operates
against the advantages of a previous evaporation under vacuum.
If sulphur collects in the neck of the flask it can be removed
easily by heating gently and uniformly the neck of the flask in
the open flame; by using a strong hot flame during the last hour
of digestion the accumulation of sulphur is almost entirely
avoid

2. If excess solid organie matter is present in the sample it
may be necessary to increase the amount of salicylie acid mixture
used in order to maintain a liquid condition of the contents during
the first part of digestion.

3. The open flame should be allowed to come in contact with
only that portion of the flask which is covered by the solution.
Use an asbestos ring to prevent this (Paul and Berry, ’21).

4. In the presence of organic matter, the tendency of the
digesting mixture to foam and spew out presents an irritating
problem. This loss by foaming has been entirely overcome in
this laboratory by allowing the acid to thoroughly disintegrate
the solid portions of organie matter, without heat, over an ap-
proximate 12-hour period It is advisable to redistribute the
acid by occasional shaking. It is necessary to stopper the flasks
tightly with rubber corks to prevent the absorption of ammonia
fumes. The practice used by the author has been to add the
acid and allow the mixture to stand over night. Of the 380 de-
terminations mentioned above, not one has been lost = to
foaming when this method was followed.

5. For evaporation of the samples a 5-hole, анына. gas-
heated water bath is used; the flasks are held on this bath in an

1925]
RANKER—DETERMINATION OF TOTAL NITROGEN IN PLANTS 373

inclined position by 2 notched wooden supports and connected to
an ordinary water filter-pump by a series of 5 corks (rubber)
and 4 Y-tubes. А 3-liter safety bottle is placed between the
filter-pump and the flasks on the water bath to prevent the en-
trance of water into the flasks when, for any reason, the water
pressure becomes reduced. In operation, a partial vacuum is
quickly developed and ebullition proceeds at a rapid rate, provid-
ing all rubber connections are sufficiently thick-walled to with-
stand the vacuum developed. Evaporate the sample just to
dryness, not to an ash-dry condition. Release the vacuum slowly
before removing the flasks; if released rapidly the flasks will
crack.

6. A few Pyrex glass beads or small angular pieces of broken
Pyrex glass placed in the flask will facilitate evaporation and
subsequent digestion; they may be used over and over again.

umping” during distillation is a question of concentration
sod relative abundance of insoluble substances present—at a
dilution of 400 cc. practically no *bumping" was experienced.

8. To determine the correct amount of alkali to use in distil-
lation the following test is of value: When ready to distill, add 2
drops of phenolphthalein indicator; add the paraffin, sodium
hydroxide, and zinc; after the flask is connected to the distillation
apparatus and the flame is adjusted, shake the flask vigorously
and if the correct amount of alkali has been added the pink color
of the indicator will flash through the solution for 1 to 2 seconds
and disappear. If the pink color lasts for more than 2 seconds
it is advisable to add more alkali; if the color disappears in less
than 1 second a useless excess of alkali is present. Once adjusted
the amount of alkali remains practically constant as long as the
amount of acid used in digestion is not varie

In the first part of this paper reference was made to certain
criticisms against various methods which are based on the re-
duction of nitrates in acid medium, and it was also shown that
those criticisms against the official method (Assoc. Off. Agr.
Chemists, '21) were satisfactorily confirmed in this laboratory.
These same criticisms are not applicable, however, to the modified
method here reported. A critical study of the data upon which
these criticisms are based reveals the fact that the moisture
content of the samples was not adequately controlled.

(Vor. 12
374 ANNALS OF THE MISSOURI BOTANICAL GARDEN

It is pertinent to this investigation, therefore, although no
attempt has been made to go into the literature, that all such
criticisms which did come to notice! be considered. In doing
so it is to be remembered: (1) that the accuracy of the modified
official method here proposed has been tested out for plants and
nutrient solutions only, (2) that, while no difficulty is anticipated
in using this method on other biologieal substances in watery
medium (for example, soil extracts), no broad generalizations are
advanced until these tests are actually made, (3) that a criticism
of the principle upon which this method is based, that is, the
acid reduction of nitrates, is also an indirect criticism of the
method here used, though the particular method under criticism
may be that of Jodlbauer (Metge, 718, p. 30), Förster (Krische, ’06,
p. 71), ete.

After the completion of this investigation a reference to the
work of Allen (15) became available. Allen's work was very
carefully done and is referred to often. In his summary, referring
to reduction methods, he concludes that ‘‘of such procedures
only the modified Devarda and aluminium reduction methods
gave promise of meeting our requirements." The data presented
indicate that these methods (that is, 2 Devarda methods and 1
aluminium reduction method) were the only ones actually tested
out by Allen (15). He rejected the acid methods of nitrate
reduction, apparently upon the evidence presented by Mitscher-
lich and Herz (’09), as follows:

* Mitscherlich and Herz conducted an extended investigation on
the perfection of an accurate Kjeldahl method which would include
all forms of nitrogen, in course of which they studied, among са
sources of error, the question of the reduction of nitric nitrogen.
Using phenolsulfonic acid and zine dust, sodium hydroxide, and zinc-

iron dust, Jodlbauer's method, and Férster’s method, they were
unable to obtain the theoretical amount of ammonia from nitrate.”
(Allen, 715, p. 522).

Reference was then made to the work of Mitscherlich and Herz
(09), whose data is an excellent tribute to accurate procedure.

1 After this paper had been sent to the publishers a reference to the work of W. E.
Loomis (Am. Soe. Hort. Sci. Proc. 1924: 365-370) was found. From the data of
7 reported determinations the official salicylic-thiosulphate method is discarded
by Loomis. The criticisms advanced against this method and against the principle
of the acid reduction of nitrate are essentially the same as the criticisms previously
advanced by Gallagher (’23), which will be considered later in this paper.

1925]
RANKER—DETERMINATION OF TOTAL NITROGEN IN PLANTS 375

For the purposes of this study, however, attention must be called
to certain facts pertaining to their report (709):

1. Of the 40 pages devoted to the report, only 12 (pages 307-
318) are concerned with a study of any method based on the
reduction of nitrates in acid medium.

2. Through their data they give no detailed statement of pro-
cedure for any of the many methods they used, except for that
modifieation of the Devarda method which they recommend
(Mitscherlich and Herz, '09, p. 280). They refer to the other
methods by name only (for example, Fórster, Jodlbauer, etc.),
and it is logical to assume, therefore, that the procedures for the
methods used were those in common use at the time. For
example, the Fórster method in use at that time was in part as
follows:

“0.5 g Salpeter (50 ccm der Lösung 10:1000) werden in einem
Kjeldahlkolben mit 15 ccm einer 6%igen Phenolschwefelsiure oder
mit 15 cem einer 6%igen Salizylsiure-Schwefelsiure vermischt,
. . ." etc. (Kirsche, '06, pp. 71-72).

a procedure for the Jodlbauer method is very similar:

. . : 250 cem Wasser werden mit 25 ccm Phenolschwefels&ure
im ‘ikjeldahl- Kolben versetzt e nach Zugabe einiger Sandkórnchen
móglichst weit eingedampft. . . ." etc. (Metge, '18, p. 20, no. 13).
Attention is called to the presence of water in the inae deter-
mined by these methods.

3. Finally, attention must be called to the fact that the deter-
minations upon which Mitscherlich and Herz ('09) based their
conclusions and to which Allen (15) referred were made upon
samples in water solutions or upon soil extracts, some of which
were very dilute. Most of the tables presented have headings
similar to the following examples:

“ Analyse einer есета mittels der Zink-Eisenstaub-
Reduktionsmethode”’ [table 37, p. 3

* Versuche zur quantitativen Reduktion des Salpeterstickstoffs
einer Kaliumnitrat-Lésung”’ e 38, p. 317].

* Stickstoffbestimmung in e m Bodenextrakt nach der Zinkeisen-
staub-Reduktionsmethode (table. 36, p. 315].

In explanation of the data in table 34, p. 313, the following occurs:

* Nach oer Reduktion der auf je 1 L verdünnten Mengen der

iumsalpeterlósung durch zwei Stunden langes Erhitzen auf fast
100? unter pee von 5 g Zinkstaub und 5 cem Phenolschwefelsáure,
nach darauffolgendem Eindampfen und Destillieren wurden folgende

[Vor. 12
376 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Ammoniakmengen entsprechend cem N/50 H,SO, im Destillat fest-
gestellt:" [table 34, page 313].

In a few cases the solutions apparently were evaporated and
theoretical recovery of nitrogen was not obtained (Mitscherlich
and Herz, '09, p. 316). In most of these cases some acid was
added (table 32, p. 312; table 35, p. 314) prior to evaporation and
in the other cases no statement can be made due to the absence
of specific details of procedures.

From a study on the determination of nitrites and nitrates in
plants Strowd (20) concluded that “the determination of ni-
irates in plants by finding the difference between the Kjeldahl-
Gunning-Arnold method and the Kjeldahl method modified to
include nitrates is unsatisfactory.” Strowd attempts to explain
this discrepancy on the basis that ''appreciable amounts of
nitrate were apparently reduced without zinc and salicylic acid."
These deductions were based on 2 determinations only (Strowd,
’20, table 1); one being a determination of the nitrate-nitrogen
content of a “Риге NaNO; solution," and the other a “Риге
NaNO; solution + nitrate-free plant extract." By calling at-
tention to these facts it is not implied that the difference between
the nitrogen obtained by the Kjeldahl-Gunning-Arnold method
and the modified official method, as here reported, would be
accurate for the determination of nitrate-nitrogen only—this
is another problem. The purposes of this investigation are served
by calling attention to the presence of water in the samples deter-
mined (Strowd, ’20).

For the determination of total nitrogen in plants Gallagher
(23, p. 67) concludes that ‘‘nitrate reductions are invariably es-
sential in dealing with Kjeldahl estimations of plant products,
since the plant contains nitrates at nearly all stages of growth.”
He recommends a Devarda method to accomplish this and dis-
cards the principle of acid reduction of nitrates on a purely
theoretical basis as follows:

“In the estimation of nitrates by means of acid reducing agent a
serious objection exists when amino groups, whether in the form of
amino acids, or of proteins, etc., be present. The first step in the
reduction of nitrates would appear to be the formation of nitrites.
In the presence of acid, nitrous acid is formed, which, of course, will

quickly react with the amino groups of amino acids, ete., with liber-
ation of nitrogen.”

1925]
RANKER—DETERMINATION OF TOTAL NITROGEN IN PLANTS 377

Under certain conditions, and for the method (Ulsch) involved
this may be a true statement of fact, but it is not true as a gener-
alized criticism of the reduction of nitrates in acid medium. In
acid medium, for example, concentrated sulphuric acid, in the
absence of water, and in the presence of either phenol or salicylic
acid, the above generalization probably does not hold true.
Skeleton equations to illustrate the transformation of the nitric-
acid radicle into ammonia may be somewhat as follows, in the
case of phenol:
(1) C,H,.OH + HNO; - C,H4.OH.NO; 4- Н.О
(2) С.Н ..ОН.М№О, + ЗН. = С Н..ОН.МН, + 2Н.0
(3) C4,H,OH.NH; + NaOH = C;,H,.OH.ONa + NH;
А very similar transformation takes place with salicylie acid;
the reduction process is hastened by the addition of zine or
sodium thiosulphate. By equation (1) it is evident that nitrie
acid is responsible for the nitrification of the phenol- or salicylic-
acid molecule. The nitric acid is obtained from the action of the
sulphuric acid on the nitrate present:
(4) NaNO; + Н,80, = NaHSO, + HNO;
If water is present in the sample to be determined it is probable
that the nitric acid will be diluted below the point at which it can
quantitatively nitrify the phenol or salicylic-acid molecule; if so,
the instability of the nitric-acid molecule will be manifest and
there will be a loss of nitrogen-dioxide gas:
(5) 4HNO; 2% 2H;,0 + 4NO; F О,
In fact, in many cases when even small amounts of water are
present, visible amounts of nitrogen-dioxide fumes may be given
off. The above generalization by Gallagher (723, p. 64) is based
on the reported data of 2 determinations only; the first one being
the analysis of ‘‘. . . . a solution! containing nitrate only, ...
and the second béitig the analysis o . & Solution containing
14.84 mg. of nitrate nitrogen and 9.13 е, of nitrogen in the form
of asparagine. . . ." Не obtained a nitrogen loss of approxi-
mately 20 per cent.

1 Italics ours,

[Vor. 12
378 ANNALS OF THE MISSOURI BOTANICAL GARDEN

SUMMARY

For the determination of total nitrogen in plants and plant
solutions the official salicylic-thiosulphate method has proved
inadequate. The inaccuracies of this method are demonstrated
to be due, primarily, to the presence of water in the sample under
analysis. A modification of the official salicylic-thiosulphate
method is proposed and certain details of manipulation are dis-
cussed. Under the conditions of this investigation this proposed
modified method has proved to be approximately twice as rapid
and just as accurate as a modified Devarda method used for
comparison purposes. Some of the criticisms advanced against
the principle of acid reduction of nitrates are reviewed.

It is with pleasure that the author acknowledges his indebted-
ness to Dr. B. M. Duggar, whose kindly interest, many sug-
gestions, and keen criticisms have been a constant source of
strength in this investigation. Acknowledgements also are
made to Dr. George T. Moore for the use of the facilities of the
Missouri Botanical Garden, and to Dr. E. S. West, Dr. Paul S.
Rider, and Dr. Roland La Garde for the timely suggestions and
assistance they have gladly given.

Graduate Laboratory, Missouri Botanical Garden.

LITERATURE CITED

Allen, E. В. (15). The determination of nitric nitrogen in soils. Jour. Ind. and
Eng. Chem. 7: 521-529. 1915.

Association of Official Agricultural iere (21). Official and tentative methods
of analysis. Washington, D. C.,

Burrell, R. C., abd Phillips, T. G. Pu Tue determination of nitrate nitrogen in
me em Biol. Chem. 65: 229-234. 1925.

ee, Р: Н, Сар Estimation of nitric-nitrogen and total-nitrogen in plant
tissue extracts. Jour. Agr. Sci. 13: 63-68. 1923.

Krische, P. ('06). Das agrikulturehemische Kontrollwesen. Leipzig, 1906.

E m (18). Laboratoriumsbuch für Agrikulturchemiker. Halle (Saale),

Made. E. A., und Herz, P. (09). Eine quantitative Stickstoffanalyse für
sehr geringe Mengen. Landw. Jahrb. 38: 279-318. 1909.
Paul, A. E., and Berry, E. Н. (21). The Kjeldahl nitrogen method and its modi-
cations. Assoc. Off. Agr. Chemists, Jour. 5: 108-132. 1921.
Strowd, W. н. (20). Тһе ссе of nitrites and nitrates in plant tissue.
Soil Sci. 10: 333-342.

[Vor. 12, 1925]
380 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE

PLATE 19
Apparatus used for the evaporation of the samples under vacuum.

6I ALVIg

СТ “TL “TOA “ачую "Log ‘о ‘NNY

EN

AL NITROC

T

`ВМТМАТТОМ OF TO

R—DETE

X
4

RANKE

COLLOIDAL SULPHUR: PREPARATION AND TOXICITY

LIONEL EARL TISDALE
Formerly Fellow in the Henry Shaw School of Botany of Washington University!

There is urgent need for a fungicide which may be used effec-
tively, without danger of injury, on a variety of plants under
diverse conditions.

Sulphur has long been known to have fungicidal properties.
Previous to 1880 it was almost the only fungicide in use, doubtless
due, in large part, to its abundance and low cost. In the years
1848, 1852, and 1853, during the serious outbreaks of the grape-
vine mildew in France (Bourcart, '13) it played an important
part as a fungicide. The diversity of the forms and compounds
of sulphur has had an important bearing upon the use of this
substance as a fungicide. The application of a mixture of lime
and sulphur for the control of grape mildew was recorded as
early as 1833 (Scott, '08), before the use of fungicides became
general. Lime sulphur is at present the form of sulphur most
widely employed and it ranks high among the standard spray
mixtures. However, the use of this spray has been limited to a
certain extent because of the caustic effect upon the foliage of
certain plants, due to the soluble sulphide content. Serious
burning is often reported from its use on apples, which are among
those plants least susceptible to this type of injury. The applica-
tion of this preparation to stone fruits is usually considered un-
safe, and it cannot be satisfactorily used on the small fruits and
many other plants.

A number of theories have been proposed respecting the toxic
constituent of the different forms of sulphur, and until recently
the problem was a subject of considerable speculation. A
complete review of the earlier investigations has been given in a
recent paper by Young (’22). This investigator demonstrated
that regardless of the form of sulphur employed as a fungicide,
whether as a compound or as uncombined sulphur, the lasting
toxicity is due to an oxidation product of the sulphur itself.
He further proved that the toxic substance is liberated most

1 A fellowship established by the Crop Protection Institute for the investigation
of sulphur as a fungicide
Анн. Mo. Bor. Garp., Vor. 12, 1925 (381)

[Vor. 12
382 ANNALS OF THE MISSOURI BOTANICAL GARDEN

rapidly from sulphur in a very finely divided state, that is, from
eolloidal sulphur.

EXPERIMENTAL

Since the experimental evidence gained from laboratory tests,
up to the present time, has shown that colloidal sulphur has
greater fungicidal value than other forms of sulphur, it was
thought important to study the methods of its preparation.
The primary object was to perfect a method or methods which
would serve as a basis for commercial production of colloidal
sulphur. After studying a variety of preparations, those materials
which indicated sufficient promise were thoroughly tested for
injury on the foliage of a variety of plants growing in the green-
house. The plants used in these tests were the following: peach,
potato, tomato, squash, cucumber, cantaloupe, geranium, to-
baeco, sweet-potato, and rose. Several of these plants were
found to be very susceptible to injury by certain sulphur com-
pounds. In those cases in which injury resulted a further
study was made in an effort to eliminate the injurious property.
The toxicity of the materials was then tested, using the per-
centage of germination of spores of a variety of economic fungi
as indicators. The Van Tieghem cell or the modified hanging-
drop culture method used by Young (’22) was employed in this
work. The following organisms were used: Botrytis Alii, B. cine-
rea, Colletotrichum Соззури, Glomerella cingulata, Gloeosporium
venetum, Macrosporium sarcinaeforme, Sclerotinia cinerea, and
Ustilago Hordei. The spores were taken from cultures 10-15 days
old, grown on dextrose-potato agar prepared by the method of
Duggar, Severy, and Schmitz (17), except in the case of Ustilago
Hordei. 'The spores of the latter organism were obtained from
smutted barley heads, freshly collected in the field.

The culture solution used in the hanging drops and in which
the sulphur materials were suspended, except in cases otherwise
indicated, was a slightly buffered mixture prepared according
to the method of Karrer and Webb (20) and of Young (22),
as follows: Stock solutions of M/5 mannite in M/10 phosphoric
acid and of M/5 mannite in M/5 sodium hydroxide were prepared.
Equal quantities of the M/5 mannite-M/10 phosphoric acid were

1925]
TISDALE—TOXICITY OF COLLOIDAL SULPHUR 383

placed in each of ten flasks and ely in

of M/5 mannite-M/5 sodium hydroxide were added The flasks
were plugged with cotton, sterilized at 15 pounds pressure for 15
minutes, and allowed to stand for a few hours. Titrations made
by the colorimetric method (Clark,’20) showed the mixture to have
the following range of hydrogen-ion concentrations: Pg 2.4, 3.4,
4.2, 5.0, 5.4, 5.8, 6.2, 6.8, 7.4, 8.4.

THE PREPARATION OF COLLOIDAL SULPHUR FROM SODIUM
THIOSULPHATE AND SULPHURIC ACID

It has been shown by Young (722)! and others that when a
saturated solution of sodium thiosulphate (hypo) is mixed with
concentrated H.SQ,, a colloidal sulphur is formed having ex-
tremely small particles. In order to free the sulphur from the
acid solution, it was coagulated with concentrated NaCl and
centrifuged out of suspension. The coagulum was then peptized
with water. By repeating the coagulation, centrifuging, and
peptizing, a pure sample of colloidal sulphur is obtained—except
for traces of the salt. This form of colloidal sulphur was found to
have a high fungicidal value, but the method of preparation
proved to be impractical for its production on a commercial
basis.

There remained one other possible method of remedying the
acid-hypo mixture, so that it might be used as a spray; namely,
to neutralize it with an alkali. Young (25) also encountered
difficulties in this process. He first used NaOH to neutralize his
hypo-acid mixture and found this alkali to be unsatisfactory
because the concentration of the electrolyte soon precipitated
the soluble sulphur. He later used lime sulphur to neutralize
the mixture and prepared the material as follows: Fifteen gallons
of saturated hypo were slowly stirred into 5 gallons of concentrated
H.SO, in a wooden barrel. The mixture was filtered through a
fine screen and the filtrate neutralized with lime sulphur diluted
one to three with water. During the addition of the lime sulphur,
which was slowly effected, a further quantity of 3 gallons of
concentrated hypo was introduced. Analysis of the final mixture

! Young gives a review of the earlier investigations upon this method of preparing
colloidal sulphur.

[Vor. 12
384 ANNALS OF THE MISSOURI BOTANICAL GARDEN

showed that 1 gallon contained 10 ounces of soluble sulphur.
He suggested that if the mixture could have been centrifuged at
this point it would have made an excellent spray. As a result
of the conditions, however, the soluble sulphur soon precipitated.

When the hypo and HSO; are mixed according to the method
just described, there is a large quantity of insoluble sulphur
formed which is thrown out in the form of a gummy mass.
This insoluble mass must be filtered off, as has already been
indicated, before further procedures. It appeared logical to
suspect that this waste could be prevented to a large extent by
the introduction of certain glutinous colloids which might be
assumed to act as protective colloids. Likewise, the introduction
of such colloids might be assumed to exhibit further protective
action by preventing the precipitation of the soluble sulphur
when electrolytes are added to neutralize the mixture.

E. von Meyer and Lattermoser (Taylor, ’21), in 1897, were
the first investigators to recognize that the addition of a very
stable sol to a less stable sol, that is, an emulsoid to a suspensoid,
prevented the precipitation of the latter by salts. The latter
author reached the conclusion that the addition of very stable
colloids, such as albumin, gelatin, agar, or gum arabic, to a silver
sol, prevented precipitation by electrolytes until the stable
colloid is gelatinized. In 1902, Zsigmondy (’09) investigated
this “protective” action of colloids quantitatively by means of
his gold sol, which in pure aqueous solution is itself very stable,
but is also very sensitive to salts. The degrees of protection of
various substances, such as gelatin, casein, egg-albumin, dextrin,
starches, cane sugar, etc., were expressed by the ‘‘gold number."
Further discussion in this paper will be limited to the use of
protective colloids in the preparation and utilization of spray
materials. | :

Various organic and inorganic substances have been recom-
mended by different investigators for increasing the spreading
and adhesive qualities of fungicides and insecticides. Though
many of the early investigations dealt primarily with the spread-
ing and adhesive qualities of sprays, it is reasonable to assume
that the physical qualities were also improved in many cases.

Glue was one of the first substances used as a spreader. Mil-

1925]
TISDALE—TOXICITY OF COLLOIDAL SULPHUR 385

lardet and Davis (Moore, ’21) in 1885 used glue with Bordeaux
mixture in the treatment of mildew of the vine, thinking that
glue increased the efficiency of the mixture. Lowe (’96) found
that glue added to lead arsenate, in the proportion of 2 quarts
to 45 gallons of spray, gave satisfactory results in spreading the
spray evenly over the foliage of cottonwood. Surface ('05) used
glue to increase the adherence of lime-sulphur spray.

More recently Jones (19) has obtained a patent covering the
use of glue as a stabilizer for oil emulsions so that they will mix
with lime-sulphur solutions. Yothers and Winston (’24) state
that such substances as casein, skimmed milk powder, gelatin,
corn meal, wheat flour, and starches are as effective as glue in
rendering oil emulsions miscible with lime-sulphur solutions.

Various kinds of soaps have been very commonly employed
as spreaders. Gillette (90) was one of the first investigators to
report the use of soap as a spreader for insecticides, namely,
Paris green and London purple. Washburn (91), Galloway
(92), and Swingle (’94) are other early investigators to recom-
mend the use of soaps.

A little later those interested in the field began to investigate
the use of spreaders quantitatively. Van Slyke and Urner (’04)
showed by chemical analysis that different kinds of soaps varied
greatly in water content. Therefore, it was concluded that the
addition of a definite quantity of soap to sprays may or may not
increase their spreading qualities.

Vermorel and Dantony (’10) studied the surface tension of
soap solutions in relation to their ability to spread over a surface.
They state that whether or not the addition of soap to fungicides
will increase their spreading qualities depends upon the method
used in the preparation of the mixture. Parker (711) recom-
mended the addition of soap to certain arsenical sprays to retard
settling and likewise increase spreading qualities.

Lovett (’18, '20) made a rather extensive study of the use of
spreaders with arsenical sprays. He concluded that the ability
of a solution to hold arsenate in suspension, while not necessarily
a criterion as to its value as a spreader, does indicate a physical
quality in the solution much to be desired in a spreader. Various
organic substances were tested, such as caseinate, glue, gelatin,

[Vor. 12
386 ANNALS OF THE MISSOURI BOTANICAL GARDEN

sage tea, soap bark, and starches. He arranged the materials
tested in the order of their merit, based on compatibility, effi-
ciency, availability, cost, and ease of preparation, as follows:
caseinate, glue, gelatin, and soap bark. A number of inorganic
compounds were also tested. Oil emulsion gave very promising
results, while other inorganic materials were unsatisfactory.

Moore (’21)!, in a quantitative study of the spreading and
adherence of arsenical sprays, found that the addition of materials
similar in chemical constitution to the leaf surface causes the
spray mixture to form a continuous film over the foliage. Various
protein substances and plant infusions gave effective spreading
on leaves with surfaces of cellulose, even when they are strongly
. eutinized. The suspensions containing small-sized particles were
found to adhere better than those of larger-sized particles.

Recently, Brinley (23) has recommended a method of prepar-
ing colloidal lead arsenate. The method consists in precipitating
the lead arsenate in the presence of a protective colloid, such as
gelatin, by the chemical reaction between lead nitrate and di-
sodium arsenate. Не found that the method of mixing and
diluting the chemicals was a very important factor in obtaining a
solution of greatest stability.

Since most of the evidence in the foregoing references points
to organic substances as being more effective than inorganic
substances in preserving the stability of solutions, it was thought
desirable to test the hypo-H.SO, mixture as prepared with
certain glutinous colloids. Thus dextrin, peptone, starch, gelatin,
and glue were tested in varying percentages. Dextrin and pep-
tone did not prove of any value in preserving the stability of
the solution. While starch possesses certain desirable character-
istics, gelatin and glue proved to be more satisfactory. The
results obtained with gelatin were slightly less desirable than
those obtained with glue. Therefore, considering the materials
upon their merits, as to efficiency, cost, and availability, glue
should be recommended in preference to gelatin.

The hypo and H;SO, were diluted in a number of ways before
mixing and the stability of the resulting solutions was tested
as prepared with glue. Likewise the stability of the solution was

1 А review of the history of spreading is given.

1925]
TISDALE—TOXICITY OF COLLOIDAL SULPHUR 387

tested as prepared with different proportions of the chemicals.
Smaller quantities of the hypo in proportion to the HS0; gave
best results. The most satisfactory results were obtained by
introducing the glue immediately after the chemicals were mixed.
If the mixture is allowed to stand any length of time before the
glue is added, the stability of the solution is lost; and, further,
the introduction of glue to either chemical before mixing results
in the loss of a considerable quantity of sulphur, the latter being
thrown out of solution in an insoluble mass.

Since temperature has been shown to be an important factor
in the preparation of other colloidal solutions, as well as sulphur,
the effect of employing warm solutions was tested, with thor-
oughly satisfactory results. Wackenroder (46) observed that
low temperatures caused a large part of the sulphur particles to
settle out of solution.

In order to neutralize the hypo-H.SO, mixture, while at the
same time preserving the stability of the solution and likewise
the toxie property of the colloidal sulphur, it was necessary to
employ various alkalies. After preliminary tests, strong alkalies
were eliminated as unsatisfactory. Sodium carbonate and some
other weak alkalies proved satisfactory. Raffo ('08) used
Na,CO, to neutralize a sulphur suspension, made from hypo
and H;SO,, after the sulphur had been centrifuged out of sus-

pension and again peptized with water. The greater part of
the sulphur was precipitated, leaving a stable solution of sulphur
containing a little МазЗО.. The use of glue prevented, to a large
extent, the precipitation when weak alkalies were added.

After testing various methods as indicated in the foregoing
discussion, the method which proved most satisfactory was as
follows: Fifty gms. of hypo were dissolved in 40 cc. of water and
warmed to 40-50? C.; 40 се. of H,SO,, specifie gravity 1.84,
were measured into a 500-cc. glass cylinder. The warm saturated
solution of hypo was added slowly to the H.SO, with occasional
stirring. Eighty cc. of warm water (30-40? C.) were immediately
added to the mixture, followed by the same quantity of 1 per
cent glue solution of the same temperature. 'The temperature,
as indicated in each case, is a very important factor and one on
which, to a large extent, the stability of the solution depends.

* \ [Vor. 12
388 ANNALS OF THE MISSOURI BOTANICAL GARDEN

'The mixture was allowed to stand 48 hours, after which time the
reaction was adjusted to Pa 4.2 with a saturated solution of
Ма.СО; and then aerated for 30 minutes. The aeration is
necessary to free the solution of traces of SO; which is very
injurious to the foliage of plants. The addition of water as
indieated is a very important step in the process. It changes
the physical state of the solution to such an extent that the
solution will remain stable for several days without the addition
of a protective colloid. In this connection, Sobrero and Selmi
(Taylor, '21), in 1850, investigating colloidal sulphur formed
by the reactions of SO, and H;S in water, make the following
remark: ‘‘If water is added to it [colloidal sulphur] it divides
up, forming an emulsion from which it does not separate out,
even on prolonged standing [several months]. ”

A saturated solution of dibasic sodium phosphate may be
used for neutralizing the mixture. It may be added to the hypo-
acid mixture immediately, thereby eliminating the necessity of
allowing the mixture to stand 48 hours before adjusting the
reaction.

After the two solutions were thoroughly tested on the foliage
of plants, toxicity tests were made. The method employed in
preparing the sulphur suspensions for testing was essentially
the same as that used by Young (22) in his toxicity studies, and
was as follows: Three test-tubes were provided with pipettes that
extended through the cork stoppers to the bottom of the tubes.
By this means drops could be transferred readily to the hanging-
drop cells. Ten cubic-centimeters of the slightly buffered
. Solution, Py 4.2, were added to each tube. This gave a duplicate
series, one tube each for the two sulphur suspensions to be tested
and one for the control culture. One cubic centimeter of the
sulphur suspensions was added to the tubes.

The technique of planting the hanging-drop cultures was the
same as that employed by Webb (21) in his germination studies
and later slightly modified by Young (’22), and was as follows:
Ground-glass rings were cemented to glass slides by means of
parawax and petrolatum. Two of these rings were placed on
each slide and 3 slides constituted a series for each organism.
A large drop of the sulphur suspension to be tested for toxicity

1925]
TISDALE—TOXICITY OF COLLOIDAL SULPHUR 389

was placed in the bottom of the two cells. Another drop was
placed on each slide and three slides constituted a series for each
organism. A large drop of the sulphur suspension to be tested
for toxicity was placed in the bottom of the two cells. Another
drop was placed on a clean sterile glass slide. A definite spore
suspension was made in the drop. By means of a small sterile
glass rod a small part of the drop was transferred to a clean sterile
cover-glass and spread out in the form of a smear. The cover-
glass was inverted and sealed air-tight by means of petrolatum
over the glass cell. The cultures were incubated at 23° C. and
germination counts made at the end of 18 hours. The results are
recorded in table 1.

TABLE I

TOXICITY OF CERTAIN FORMS OF COLLOIDAL SULPHUR. THE FIGURES
INDICATE PERCENTAGE OF GERMINATION!

Organism
Form of sr
sulphur Botrytis | Colletotrich- | Glomerella msi a Sclerotinia
cinerea um Gossypii| cingula руно
Control—
without
sulphur 80 65 76 94 60
Colloidal :
sulphur
neutralized
NaCO; 5 / 0 : 8 44 2
Colloidal
sulphur
—
wit
Na HPO, 15 4 12 57 5

1 Average of duplicate tests run at two different times.

The results in this table show that both colloidal sulphur
solutions tested were toxic to all the organisms used. The solu-
tion prepared with disodium phosphate was slightly less toxic
than that prepared with Na4;CO;. This can be accounted for
in that it required more of the phosphate to neutralize the
mixture. Therefore, there was less sulphur in the mixture.

[Vor. 12
390 ANNALS OF THE MISSOURI BOTANICAL GARDEN

THE PREPARATION OF COLLOIDAL SULPHUR FROM LIME SULPHUR

Since the introduction of lime sulphur as an orchard spray
serious injuries to foliage of plants have been frequently reported
as a result of its use. Injury is often more frequent when the
spray is used in combination with certain arsenicals. This is
shown by a survey of the literature from various agricultural
experiment stations. A number of methods for the elimination
of these difficulties have been suggested by different investiga-
tors, all of which relate primarily to the method of diluting,
mixing, and applying the spray, or to the time of application, or
the evidence of inferior compounds. Since lime sulphur is
composed largely of sulphides (pentasulphides) the caustic
action on foliage of plants has been attributed to these soluble
compounds. A few investigators have suggested that the injuri-
ous properties can be eliminated by precipitating the soluble
sulphides with salts or acids. These methods, however, have
not come into general use.

Wallace (710), ina study of lime-sulphur spray injury, precipi-
tated commercial lime sulphur, diluted 1-30 with water, by
applying it with a carbonic-acid gas sprayer. The solution was
allowed to stand in contact with the gas one-half hour before
the application of the spray. A heavy white precipitate was
formed, and this did not cause injury to apple and peach foliage
when used alone as a spray. However, when lead arsenate was
introduced into the mixture before the precipitation, the resulting
material used on young peach trees caused almost complete
defoliation.

Stewart (12), in a similar study, pointed out that lime-sulphur-
arsenical injury could be eliminated by precipitating the mixture
with iron sulphate, or “copperas,” added at the rate of 3%
pounds to 50 gallons of 1.01 spray solution. The mixture was
found to be very disagreeable to handle, due to its tenacious
and blackening qualities. Also serious discoloration of fruit
resulted from its use, and the effectiveness of arsenicals was
reduced. Waite (’10) had previously reported the use of iron
sulphate and copper sulphate in the preparation of self-boiled
lime sulphur.

Safro (713), after a study of lime sulphur spray injury, con-

1925]
TISDALE— TOXICITY OF COLLOIDAL SULPHUR 391

cluded that the effect was due to the soluble sulphide content of
thespray. He found that the injury could be reduced by render-
ing the sulphides insoluble, that is, by precipitating them. His
results indicated that the injury could be reduced by the addition
of either iron, copper, or zinc sulphate, the salt being first dis-
solved in water, in the proportion of 4 pounds to 100 gallons of
diluted (1 to 20) lime-sulphur solution. Carbon dioxide and
H.SO, were also used, the substances being introduced until
the remaining sulphides gave the spray an amber color. The
carbon dioxide entirely eliminated the injury.

More recently, definite methods have been given for precipitat-
ing lime sulphur for commercial use. Ramsay and Cooke (722)
describe a method of preparing colloidal sulphur from lime sulphur
which has been used effectively as a spray in Australia. The
` method recommended is as follows: Ten gallons of home-made
lime sulphur, 26? Baumé, were diluted with 25 gallons of water
in a barrel of 40 or 50 gallons capacity. Using a porcelain or
earthenware vessel, 6 pints of strong commercial H;SO, were
poured slowly into 9 pints of cold water and the solution allowed
to cool. The cold diluted H;SO, was then added to the diluted
lime sulphur, 2 or 3 ounces at a time, stirring well after each
addition of acid, until the typical yellow color of the original
lime-sulphur disappeared and until the addition of H;SO, pro-
duced no further precipitation of sulphur. The precipitated
sulphur was allowed to settle for a day or two and the clear
liquid above was siphoned or decanted off. Three pounds of
cheap glue were dissolved in just sufficient hot water to render
the glue soluble, and, while still hot, stirred thoroughly into the
precipitated sulphur. For use, the mixture thus obtained was
diluted with water to 250 gallons. This gave approximately 5
pounds precipitated sulphur per 100 gallons of spray.

After making a study of the toxicity of the various ingredients
of decomposed lime sulphur, Young (’22) showed conclusively
that the lasting toxicity was due to the precipitated sulphur.
This form of precipitated sulphur was found to be about as
toxic to parasitic fungi as “hydrophobic” colloidal sulphur.
Later (25) he prepared colloidal sulphur from lime sulphur as a
summer spray. The results from the use of this material in

[Vor. 12
392 ANNALS OF THE MISSOURI BOTANICAL GARDEN

combating apple scab were very encouraging. His method was.
as follows: Ten gallons of lime sulphur, 32? Baumé, were placed
in a 50-gallon barrel. To this were added 10 gallons of water
and 1 pound of glue, the latter previously dissolved in hot water.
Sulphurie acid diluted with 3 parts of water was then added until
the reaction of the mixture was P, 4.2. The mixture was allowed
to settle and the supernatant liquid decanted off. The mixture
eontained about 1 pound of active sulphur per gallon. Five
gallons of the precipitate were diluted with water to 95 gallons
before using as & spray.

Since the recent uses of precipitated lime sulphur as a spray
have given encouraging results, it was considered important to
make further studies upon its preparation with the object of
producing a mixture of greater stability. Likewise from previous
tests of toxicity (Young, '22), the mixture would be expected to
exhibit greater fungicidal properties. There were several factors
which were thought to be important in the preparation, namely:
(1) the choice of acids and the dilution of these; (2) the concentra-
tion and method of diluting the lime-sulphur solution; (3) the
temperature; and (4) the protective colloid, its concentration,
and the method of introduction.

The lime sulphur used for this work was a freshly made solu-
tion, prepared according to the Van Slyke method, that is, in
the proportion of 80 pounds sulphur, 36 pounds lime (CaO),
and 50 gallons of water, these ingredients being boiled together
one hour. Preliminary tests were made with a number of acids
and phosphates to determine which of those were most desirable
for precipitating the freshly made lime sulphur. The use of
phosphates, of certain acids, and also of strong solutions of acids
gave unsatisfactory results and these procedures were eliminated.
Weak solutions of HNO; and HSO; gave very promising results.
Glue, which proved to be the most satisfactory in the previous
preparations, was used as a protective colloid. The concentration
of the glue, the method of introducing it, the dilution of the lime
sulphur, and the temperature of the solutions employed in the
preparation were found to be the important factors upon which,
to a large extent, the stability of the solution depended.

Since the preliminary tests showed that the dilution of the

1925]
TISDALE—TOXICITY OF COLLOIDAL SULPHUR 393

acid used in precipitating lime sulphur had an important bearing
upon the stability of the solution, further tests were made in
order to determine what strength of HNO; and HSO, could be
most satisfactorily employed. Twenty-five cc. of lime-sulphur
solution were measured into each of 10 glass beakers of 300 cc.
capacity. The beakers were numbered 1-5 inclusive, which gave
a duplicate set, one set each for the two acids to be tested.
The mixtures were then adjusted to the reaction Pg 4.2, with
dilutions of HNO; and HSO; as follows:

No. 1. Acids diluted 1-3 with water.

No. 2. Acids diluted 1-5 with water.

No.3. Acids diluted 1-10 with water.

No. 4. Acids diluted 1-20 with water.

No. 5. Acids diluted 1—40 with water. |
The resulting solutions showed a striking range in stability.
The precipitated sulphur in solutions No. 1 settled out of suspen-
sion within a few hours after it was prepared. Solutions No. 2
remained stable several days. Solutions No. 3 remained stable
about 2 weeks. Solutions No. 4 remained stable several months.
Solutions No. 5 were less stable than No. 3. The solutions
prepared with HNO; possessed slightly greater stability in all
cases than those prepared with H;SO, These results show
conclusively that the strength of the acid used in precipitating
the sulphur is an important factor in producing a product of the

desired stability.

After determining the proper dilution of acid to use, further
studies were made of different procedures in making the prepara-
tion. The method which proved the most satisfactory is as
follows: Fifty cc. of freshly made lime sulphur were measured
into a glass beaker of 600 cc. capacity. One gram of dry flake
glue was dissolved in 200 cc. of water and mixed with the lime
sulphur while warm (35-40? C.). Ten cc. of HNO; were diluted
with water in a separate container. The diluted HNO; was
then added to the diluted lime-sulphur mixture very slowly,
with stirring, until the reaction Py 4.2 was obtained. The
mixture was allowed to set for several hours in the open container,
with occasional stirring, to remove the H,S. This gave a final
dilution, in respect to the original lime-sulphur solution, of 1 to 9.

[Vor. 12
394 ANNALS OF THE MISSOURI BOTANICAL GARDEN

The mixture contained approximately 1.8 per cent sulphur.
Colloidal sulphur prepared by this method will remain stable for
several months. Sulphuric acid diluted 1-20 may be used to
precipitate the lime sulphur mixture. However, as previously
indicated, the solution as prepared with H;SO, is less stable.
In determining the toxicity of this form of colloidal sulphur,
the mixtures prepared with HNO; and H.SO, were compared
with lime sulphur precipitated by the СО, in the air, the latter
being effected as follows: Ten cc. of lime sulphur solution were
diluted with 60 cc. of water and this solution was poured into a
large open dish and allowed to decompose. The precipitate
was then taken up in 70 cc. of water. One cc. of each of these
3 precipitated lime-sulphur solutions was then added to 5 cc.
of the slightly buffered mannite-H,;PO,-NaOH mixture of Pa 4.2
-. (except in tests with Macrosporium sarcinaeforme, where M /5 man-
nite was used), and the toxicity determined as before. The
organisms used and the results are recorded in table п.

TOXICITY es PRECIPITATED LIME-SULPHUR SOLUTIONS. THE fk kad

INDICAT (AVERAGE OF CULTURE
RUN AT 2 DIFFERENT TIMES)
Organism
Form of Botrytis
sulphur cinerea | Glomerella eaten Sclerotinia | Ustilago
cingulata | sarcinae- cinerea Hordei
forme
Control—without
sulphur 84 64 88 59 73
ime sulphur solu-
за Od
24 10 10 0 1
ты sulphur solu-
чон itated
th Hs. 55 13 30 0 3
Ман sulphur solu-
on itated
by CO, in 71 — — — 21

From the results exhibited in table тт it is clear that the toxicity
of the sulphur prepared by the different methods is related to

1925]
TISDALE—TOXICITY OF COLLOIDAL SULPHUR 395

the stability of the sulphur suspensions. The sulphur precipitated
from lime sulphur by the CO, in the air settles out of suspension
in à very short time, and is therefore the least toxic of the three
forms of sulphur tested. The solution prepared with HNO,
possesses the greatest stability and likewise exhibits the greatest
toxicity.

The mixtures prepared with HNO; and H,SO, were thoroughly
tested on the foliage of plants already named growing in the green-
house. They were found to spread over the surface of the
foliage evenly and adhere well; in fact it was difficult to remove
the films by washing with a strong stream of water from the hose.

PREPARATION OF COLLOIDAL SULPHUR BY MEANS OF 80, AND HS

The formation of colloidal sulphur by the reaction between
SO. and HS in water was first recorded by Berthollet (1798).
The reaction was later studied by Wackenroder (’46) who dis-
covered pentathionic acid and prepared it by passing HS into
a saturated solution of SO;, always keeping an excess of the latter.
Lewes (81), Spring (782), Shaw (88), and Débus (’88) are other
early contributors to the study of the properties of pentathionic
acid formed by this process. The reaction of SO, and H;S has
been studied more recently by Riesenfeld and Feld (21) in a
study of polythionie acid and polythionate.

Selmi and Sobrero (50) (also earlier investigators) made
rather extensive studies of colloidal sulphur formed by the
reaction of SO, and H:S. A review of the work is given by
Odén (13). That there is more than one state of colloidal
sulphur formed by this process is shown by Selmi (52) who
prepared colloidal sulphur as follows: Sulphur dioxide was passed
into distilled water until a saturated solution was formed. Hydro-
gen sulphide was then passed into the saturated solution of SO;,
care being taken not to have an excess of HS, as it precipitates
the soluble sulphur. The solution was then centrifuged to remove
the larger particles and the supernatant liquid coagulated with
sodium chloride. Тһе coagulum was then peptized in water.
Further evidence of the two forms of colloidal sulphur is given
in the papers cited above. Young (22) has designated the two
forms, according to their degree of hydration, as hydrophilic

[Vor. 12
396 ANNALS OF THE MISSOURI BOTANICAL GARDEN

and hydrophobic colloidal sulphur. "There are in all probability
several grades of fineness in the hydrophobie forms. This is
shown in the work of Odén (13).

The colloidal sulphur prepared from SO, and HS in this in-

vestigation consists largely of the hydrophobic form. The method
used was as follows: A rubber stopper having two holes was
provided with two thistle tubes, over the mouths of which
finely perforated parchment paper was securely fastened. The
tubes were inverted into a 2-liter flask containing 1500 cc. of
water, the stopper closing the mouth of the flask. Sulphur
dioxide (compressed liquid) and H.S (made from FeS and 1 to
1 muriatic acid) were passed simultaneously into the water
through the thistle tubes, keeping the sulphur dioxide slightly
in excess. After three hours a slight excess of H.S was intro-
duced to free the solution of SO:. The suspension thus formed
contained 12 per cent sulphur, which had great stability. In
fact, there was but very little settling out after standing three
months.
_ The efficiency of the method is determined by the fineness of
the perforations through which the gases are introduced into
the liquid. In the method described the gases are introduced
in the form of a fine spray, which permits a more thorough mixing
in the solution. The introduction of the gases in the form of
large bubbles was found to be inefficient and unsatisfactory in
preliminary tests. The use of distilled water has been found
more desirable than tap water, especially since the latter may
contain sufficient salts or alkalies to effect a partial precipitation
of the colloidal sulphur as it is formed.

The colloidal sulphur solution thus prepared has an acid
reaction slightly beyond the acid range of available indicators,
Ри 1.0. Wackenroder (46) found that some H.SO, was formed
during the process in the presence of oxygen. It was later found
difficult to free the mixture entirely of SO», and it was evident
that traces were left in the solution. The solution was found to
be injurious to the foliage of plants; therefore, it was necessary to
adjust the reaction, or treat the solution in some way before it
could be used as a spray.

The weak alkalies used for adjusting the colloidal sulphur

1925]
TISDALE—TOXICITY OF COLLOIDAL SULPHUR 397

solution prepared by the hypo-H;SO, method, namely, М№а:СО;
and the Na;HPO,, caused precipitation of the colloidal sulphur
when freshly prepared by this method. Similar results were also
produced by the introduction of glue or gelatin. However,
promising results were obtained by aerating the mixture for 48
hours and then adjusting the reaction to Pg 4.2 with a weak
solution of Na,HPO,. The resulting solution remained stable
for 3 months, after which time the colloidal sulphur was precipi-
tated. This material was found to be very toxic to fungi when
first prepared. It proved effective for control of carnation rust
in the greenhouse.

Later the colloidal sulphur was coagulated with NaCl and
allowed to settle out of solution. The method was as follows:
A saturated solution of NaCl was added to the freshly prepared
colloidal sulphur suspension until it changed from a bright to
a pale yellow color. The mixture was allowed to set two hours,
after which time the coagulum had settled out of suspension.
The supernatant liquid was then siphoned off and the coagulum
washed with water and allowed to settle again. Тһе supernatant
liquid was again siphoned off. The sulphur is left in the form
of a thin paste, containing about 70 per cent water and a little
NaCl. The sulphur readily assumes the colloidal state when
further quantities of water are added. Selmi and Sobrero (’50)
found that colloidal sulphur prepared from SO: and H;S could be
coagulated by a neutral sodium salt, and again be emulsified in
water. They state that when potassium salts are used the pre-
cipitated sulphur completely loses the property of emulsifying
in water.

When the sulphur paste (prepared by the above method) is
resuspended in water, shortly after it has been coagulated,
the suspension remains stable over a long period of time. In
fact, such a suspension was kept in the laboratory for 8 months,
and after this there was still little or no settling of the sulphur.
However, if the coagulum is not taken up in water, the sulphur
begins to crystallize in about 1 month and soon loses its colloidal
property. The paste form would be much more desirable for
practical use. "Therefore further investigations should be made
to determine if it can be treated in such way as to prevent crystal-
lization.

4

[Vor. 12
398 ANNALS OF THE MISSOURI BOTANICAL GARDEN

The results on the toxicity tests of this form of colloidal
sulphur are discussed on subsequent pages.

THE TOXICITY OF CERTAIN FORMS OF COLLOIDAL SULPHUR IN
RELATION TO THE HYDROGEN-ION CONCENTRATION

Since it was shown by Young (’22) that the hydrogen-ion
concentration has a marked influence upon the toxicity of
certain forms of colloidal sulphur, it was thought important to
determine if this influence may be exhibited by other forms of
these colloidal preparations. A colloidal sulphur solution was
prepared from SO, and HS as already described, with modifica-
tions as follows: After the colloidal sulphur had been coagulated
with NaCl, the solution was then centrifuged for 15 minutes at
1500 revolutions per minute. The supernatant liquid was then
decanted. Upon drying and weighing, the coagulum was found
to contain 50 per cent of sulphur.

One gm. of the coagulum was weighed out and taken up in
100 сс. of distilled water. This gave a dilution of sulphur of
0.5 gm. in 100 cc., or 1 to 200. When freshly made, the reaction
of the solution was Pg 4.0; after standing in a closed container
for 1 week the Py had changed to 2.2. The freshly made sulphur
suspension also changed the hydrogen-ion concentrations of the
culture solutions. "Therefore it was found necessary to adjust
the sulphur suspensions to the range of hydrogen-ion concentra-
tions corresponding to the particular hydrogen-ion concentration
of the culture solution, which was done as follows: Fifty-cc.
quantities of the 1 to 200 colloidal sulphur suspension of Ри 4.0
were measured into each of 11 clean glass beakers. The suspen-
sions were then adjusted toward and into the alkaline range with
an increasing number of drops of M/10 Ма„НРО, solution,
and into the acid range with M/10 H;PO, solution. This gave a
range of hydrogen-ion concentrations as follows: Pg 2.4, 3.4,
4.2, 5.0, 5.4, 5.8, 6.2, 6.8, 7.4, and 84. The colloidal sulphur
suspension thus prepared did not change the hydrogen-ion
concentrations of the culture solution during the 18-hour period
of incubation of the cultures. The stability of the sulphur
suspensions of different hydrogen-ion concentrations will be
discussed later.

1925]
TISDALE——TOXICITY OF COLLOIDAL SULPHUR 399

In later work it was found that if the coagulation, centri-
fuging, and peptizing with water is repeated 3 times, the hydro-
gen-ion concentration of the resulting colloidal sulphur suspen-
sion does not change even on prolonged standing. Likewise,
such a suspension does not evert the hydrogen-ion concentration
of the culture solution. This is probably due to the buffer action
of the phosphate which was added.

The culture solution used for making the toxicity studies was
the slightly buffered mannite-H;PO,-NaOH mixture prepared
according to the method already described. Twenty test-tubes
were provided with pipettes that extended through the cork

foe

= Manriife - НУР Мады
$ Buffered Mixture.
ES
S60
5
EN
Q
So
* :
S5
М
e M CAU ad
Pa 2 3 4 5 6 7 8
1. Germination spores Botrytis ci in hanging-drop cultures
toxic action of certain forms of colloidal sulphur; prepared by method 1— - — -;
by method 1, ———i b; 2 -----; соп-

trol, without sulph

stopper to the bottom of the tubes, according to the method
employed by Young (22). 'This constituted a duplicate series
of 10 each, and 10 cc. each of the slightly buffered solutions were
added to the tubes so that each tube represented a particular

[Vor. 12
400 ANNALS OF THE MISSOURI BOTANICAL GARDEN

hydrogen-ion concentration. The colloidal sulphur suspensions
were added to one series, with accurately calibrated pipettes,
in quantities to make the dilutions indicated in table пт.

The hanging-drop cultures were prepared in closed-ring cells
according to the method already described, and were incubated
at 23° C. After germination of 18 hours, counts were made.
The results of these tests are shown in table пт, also in figs. 1-4.

/40

А

Germination,
Q

Mannite-Hz Po- NaOH.

"n Buttered Mixture.
©
x
©
2
Re
dest ee fs e.
p x e шай E = — жо -
Py 2 3 4 J 6 7 $
Fig. 2. Germination of spores of Gloeosporium venetum in hanging-drop cultures:
toxic action of colloidal sulphur prepared by method 1 — - —; control, without
sulphur —————.,

The results show that the concentration of the sulphur em-
ployed in the culture solutions was a very important factor.
The organisms tested vary greatly in their resistance to the
toxicity of this form of colloidal sulphur, M. acrosporium sarcinae-
forme being very resistant, while extremely weak dilutions are
sufficient to kill Sclerotinia cinerea. In general, this form of
sulphur i 1s most toxic toward, and in, the alkaline range of hydro-
gen-ion concentrations. The highest percentage of germination
was in most cases at Ри 3.4. Beyond this point toward the

1925]
TISDALE—TOXICITY OF COLLOIDAL SULPHUR 401

alkaline range there was a gradual decrease to the point Pg 5.8,
after which there is a sharp decrease.

The influence of the hydrogen-ion concentration upon the
toxicity of this form of colloidal sulphur differs from the results
obtained by Young (22) with his hydrophilic colloidal sulphur.
In order to make a comparison by the same method, hydrophilic

дд
To
$
v PTannie-H, РО МОН
PA Buffered Mixture
Q
ie
S
$
+
Pao
T Set a meer z=
DPE ong Vania,
Pu 2 3 E 23 6 7 y
Fig. 3. Germination of spores of Sclerotinia cinerea in engi tir cultures
toxic action of certain forms of colloidal sulphur; т, by method 1 —-—;
prepared by method 2 - - - - - - ; control, without sulphur ———————.

colloidal sulphur was prepared according to the method of Young
(22) which was as follows: Fifty gms. of pure crystalline sodium
thiosulphate were dissolved in 30 cc. of distilled water; 70 gms.
of concentrated H,SO,, sp. gr. 1.84, arsenic-free, were weighed
into a glass cylinder of 300 cc. capacity. The cylinder was
placed in a vessel of cold water, and the saturated solution of
sodium thiosulphate added very slowly with occasional stirring.
The mixture was then allowed to cool, and 30 cc. of distilled water
were added. The preparation was then placed in a water bath
and warmed to 80? C. for 10 minutes and filtered through glass

[Vor. 12
402 ANNALS OF THE MISSOURI BOTANICAL GARDEN

wool to remove insoluble sulphur. After the mixture was allowed
to cool, concentrated NaCl was added and the mixture centrifuged
for 30 minutes at 1500 revolutions per minute. The coagulum
was then peptized with 100 cc. of distilled water and the insoluble
sulphur centrifuged out. The peptized sulphur was then treated
3 times with 25 cc. of saturated NaCl and finally peptized by
adding 1 gm. of the coagulum to 100 cc. distilled water. The
solution had a Pg of 4.2 ,which does not change upon standing.

100
| 80 Mannite- l PO-No0H
$ Buffered Mixture
E
Ф bo
3
N
A
Q ps A x \
Cs o
N
$ \
v `
e 20 Br Ss X a
а ~. ес
———— я
Рн 2 ее. ГА

Fig. 4. Germination of spores of Ustilago Hordei in hanging-drop cultures:
toxic action of certain forms of colloidal sulphur; prepared by method 1- — - "i
prepared by method 1, crystallized — — — ; prepared by method 2 ----- H
control, without sulphur ————,

By means of an accurately calibrated pipette, aliquot parts
were measured into each of 10 tubes containing the slightly
buffered solution, to make the dilutions indicated in table ш.
The hanging-drop cultures were made in exactly the same way
as described for the other forms of colloidal sulphur, and germina-
tion counts were made after an incubation interval of 18 hours
at 23° С. The results are shown in table тп and in figs. 1, 3, and 4.

1925]

TISDALE—TOXICITY OF COLLOIDAL SULPHUR

S5 [95.15 гого ооо 0000-1 | = роцзәуу Aq poredoxq

I 5 Y L 8 rat гта 9 I 0007-1

0 0 0 0 0 0 5 0 0 0 000°%-1 Co umuodsoooto
7$ | OF It СЪ 13 79 63 53 8? L6 гацав зпоцулм *[o13u07)

+ | 6 OF рор ултты —Tw aE | 000‘0I-1 І pom 4qperwdenp| падззок)

2 | er || |o | v9 | s2 | — | eo | е | oF зачатие зпоцум “толупогу | 294213039100)

н тан

о |3 * 198 |1 |. 99 | — | WW. | 181 8 1 UNT q te rf yen a net Shoe

о | 0 0 p от | 61 | — | O | 2 | er | 000-1 I poy Aq poredoiq

0 9 eT aise | Vj тт гити | ydys зпоцзм '[orjuo)

0 0 0 0 0 0 0 0 0 0 0002-1 € pomo Aq peredo1q

0 0 0 I 5 3 6 E | » 13 000'c-1 т PON Aq рэзедэз | my 89404

0 c 8 ге m тїчаүпв почта ‘101305)
PSI PL 89; SO Se | FE POG Set re vt жы ышы ы ао

(и) чотувлупәәпоә uor-uoo1pAgr Jo поп] 7 v.
(SLSAL 9-2 ЯО HOVH WAV) NOLLVNINUAD
ЯО WHOVLNHOUHWd JLNWSHNdWHH БЯНПОІЯ AHL 'SNOISNWd$8208 SAOHd'I108 NIVLUAO AO ALIOIXOL

III WIAVL

TABLE III (Cont'd.)
-i ti P
GG | Fori ог оома sulphur Dieten: of Hydrogen-ion concentration (Pa)
2.4 | 3.4 | 4.2 | 5.0 | 5.4 | 5.8 | 6.2 | 6.8 | 7.4 | 8.4
M porium Control, without sulphur 17 90 94 97 94 93 93 90 83 | 75
sarcinaeforme! Prepared by Method 1 1-500 м ммм | 18 | 19 | 30
Control, without sulphur 53 56 59 57 56 50 28 20 2 0
è -10,000 0 `0 0 0 0 0 0 0 0 0
Sclerotinia | Prepared by Method 1
— á 1-20,000 1 7 6 81 4 2 1 1 0| 0
Prepared by Method 2 Aa оа ете [э з тее Тоо 10
Control, without sulphur 71 72 73 — 68 44 40 39 23 | 14
Prepared by Method 1 1-1,000 6 8 T — 7 6 5 0 0 0
Ustilago
Hordei Prepared by Method 1, ;
sulphur crystallized 1-1,000 39 49 20 — 14 — 16 — —
Prepared by Method 2 1-2,000 22 18 15 — 11 12 24 3 0

Nadav) 'IVOINVLO&S THNOSSIN AHL AO S'IVNNV ҒОР

ZI тод]

1925]
TISDALE— TOXICITY OF COLLOIDAL SULPHUR 405

In the table and figures just referred to, the form of colloidal
sulphur is indicated according to the method of its preparation.
Thus, Method I refers to the colloidal sulphur prepared by the
SO.-H,S process, and Method 2 refers to the hydrophilic form
of colloidal sulphur prepared with sodium thiosulphate and
Н,ЗО..

The results of the toxicity tests of the two forms of colloidal
sulphur show that Young's hydrophilic form is much more
toxic to the organisms used, as it required much smaller quantities
of this form to inhibit germination of spores. This toxicity
corresponds to the size of the sulphur particles. It has already
been indicated that the form prepared by the SO.-H.S method
is largely the hydrophobic form with probably a small amount
of the hydrophilic form.

The influence of the hydrogen-ion concentration upon the
hydrophilie colloidal sulphur was found to be essentially the
same as that recorded by Young (22). Upon examining the
culture tubes containing the hydrophilic colloidal sulphur, it
was also found that settling out was rapidly increased as the
Py increased beyond Py, 5.4. However, this does not explain
the difference in toxicity of the two forms. It appears logical
that if the toxic substance, exhibited by the colloidal sulphur
prepared by the SO.-H:S method, is pentathionic acid (“truly
‘soluble’ sulphur") it would be precipitated at a higher Ри than
5.4, as has been shown to be the case with hydrophilic colloidal
sulphur. Further proof that pentathionic acid would be precipi-
tated, particularly in the more alkaline solution, is shown by the
investigations of Wackenroder (46), Shaw (’83), Débus (88),
and others. "Therefore, it is shown conclusively that the toxie
substance exhibited by the form of colloidal sulphur prepared
from SO, and HS possesses properties different from the toxic
substance exhibited by the hydrophilie form, prepared according
to the method of Young. From the evidence, the latter is, in all
probability, pentathionic acid.

There were some indications that the toxic substance exhibited
by the colloidal sulphur prepared from SO, and H;S may be
destroyed by repeating the coagulation, centrifuging, etc., several
times. This remains to be established by more detailed study.

[Vor. 12
406 ANNALS OF THE MISSOURI BOTANICAL GARDEN

It was later found that when colloidal sulphur was prepared
by the SO.-H.S method, coagulated with NaCl, and kept for
some time in coagulated form, the sulphur soon crystallized and
would no longer remain in suspension when water was added.
Suspensions were made of this crystallized sulphur and tests
were made in the usual way with Botrytis cinerea and Ustilago
Hordei. The results of these tests, in table пт, show that the
toxie substance has been destroyed to a large extent.

THE INFLUENCE OF THE HYDROGEN-ION CONCENTRATION UPON

THE STABILITY OF COLLOIDAL SULPHUR SUSPENSIONS, AND

THE RESULTING EFFECT UPON THE TOXICITY

Since it was shown in the toxicity tests that the form of colloidal
sulphur prepared by the reaction of SO: and HS exhibits its
greatest toxicity at a higher Р. than 4.2, it was considered
important to determine whether the hydrogen-ion concentration
has any influence upon the stability of the suspension. A colloidal
sulphur dispersion was prepared by the SO.-H.S method, coagu-
lated with NaCl and centrifuged. The coagulum was then
weighed and taken up with distilled water, in the proportion of
1 gm. of sulphur to 200 cc. of water. Fifty-cc. portions of the
suspension were measured into each of 11 beakers. The Ри of
the suspension was 4.2. The suspensions were then adjusted
towards, and into, the alkaline range with M/10 Na;HPO,, and
on the acid side with M/10 Н,РО,. The resulting preparations
had the following range of hydrogen-ion concentrations: Pg
beyond the acid range, 2.4, 3.2, 4.2, 5.0, 5.4, 5.8, 6.2, 6.8, 7.4,
and 8.4. The acid solution was adjusted slightly beyond the
acid range of available indicators. Portions of these solutions
were transferred to clean test-tubes, and the tubes were closed
with corks. The remaining portions were used in the subsequent
experiment.

The suspensions showed a very striking range in stability.
Settling out of the sulphur was increased rapidly as the Pg
increased beyond 5.8. A photograph of the tubes was made
after 8 months and is shown in pl. 20. The stability of the prepara-
tions was as follows: Рн 8.4 began to aggregate and settle in
about 1 week and had settled out completely in 2 weeks. Pg 7.4

1925]
TISDALE—— TOXICITY OF COLLOIDAL SULPHUR 407

settled out in 3 weeks. Pg 6.8 settled out in 4 weeks. Pa 6.2
had settled out to a large extent after 7 weeks. Pg 5.8 changed
from a bright yellow to cream-white in about 8 weeks, with some
settling out. After 8 months this solution had largely settled, but
the photograph does not bring this out clearly because of sulphur
particles sticking to the sides of the tubes, which could not be
removed without agitation of the solution. The acid solution
(that is, the solution which was adjusted beyond the acid range of
available indicators) began to settle in about 1 month and con-
tinued slowly. There was still some sulphur in the suspension
after 8 months, as shown in pl. 20. No great difference was
manifest in the stability of the preparations represented by
Py 2.4, 3.2, 4.2, 5.0, and 5.4. There was some sediment in all
the tubes after 8 months, but the range from Ри 4.2 to 5.4
shows greatest stability.

The tubes were shaken, and 0.5 cc. were taken from each
tube and transferred to tubes containing 10 cc. of the slightly
buffered solution of like hydrogen-ion concentration. This gave
a final concentration of sulphur of 1 to 4,000 which is weaker
than that used for other toxicity tests. Spore suspensions of
Botrytis cinerea were made from the sulphur suspensions, and
germination tests were made in the usual way in closed-ring cells.
The results are shown in table Iv.

TABLE IV

TOXICITY TESTS WITH CERTAIN FORMS OF COLLOIDAL SULPHUR. THE
FIGURES INDICATE PERCENTAGE GERMINATION

Hydrogen-ion concentration (Px)

Organism | Form of sulphur
24|3.2 | 4.2 | 5.4 | 5.8 | 6.2 | 6.8 7.4

Control, without
DU eR 72 | 83 | 80 | 75 | 57 | 47 | 18 6

Botrytis [Colloidal sulphur
cinerea adjusted to Ри
range indicated
after

8 months 24 | 60 | 67 | 47 | 62 | 39 | 29 | 10

The results of this test show that when the stability of the

sulphur is destroyed by a higher P, than 5.4, the toxic
is likewise destroyed.. In this case there was still some sulphur

[Vor. 12
408 ANNALS OF THE MISSOURI BOTANICAL GARDEN

in suspension at Pg 5.8. However, it did not exhibit any toxicity
to Botrytis cinerea. The suspensions referred to as Pg 2.4, 3.2,
4.2, and 5.4 are all toxic even with the very weak concentration
of sulphur used.

THE TOXICITY OF VOLATILE PRODUCTS OF COLLOIDAL SULPHUR

Since it was demonstrated by Young (22) that hydrophilic
colloidal sulphur exhibited the usual degree of toxicity when
the spores were not in direct contact with the sulphur, it was
thought important to determine whether the toxic substance of
colloidal sulphur made by the SO.-H.S method would be liberated
to the same extent. To test this point, portions of the freshly
prepared sulphur suspensions, employed in the preceding experi-
ment, were used. These suspensions had been adjusted, as
indicated, to the hydrogen-ion concentration range Pa 2.4-8.4.

The organisms used were Botrytis Alii and Ustilago Hordei.
The hanging-drop cultures were made in the usual way, with
modifications as follows: A small quantity (one-tenth cc.) of
each sulphur suspension was placed in the bottom of each cell.
The spores were suspended in the slightly buffered mixture
without sulphur. In this way the spores were not in direct
contact with the sulphur particles, being separated by the height
of the cell, which was 8 mm. The cultures were incubated at
23^ C. and germination counts made at the end of 18 hours.
The results are given in table v.

TABLE V
TOXICITY OF VOLATILE PRODUCTS FROM CERTAIN COLLOIDAL SULPHUR
SUSPENSIONS. THE FIGURES REPRESENT PERCENTAGE
GERMINATION

— —

Hydrogen-ion concentration
Organism | Form of sulphur +

2.4 | 3.2 | 4.2 | 5.4 | 5.8 | 6.2 | 6.8 | 7.4

Botrytis grec — 59 | 81 | 82 | 70 | 51
à ur 51 | 38 | 8
Ahi Colloidal sulphur in
bottom of cells 5 7 6 4 0 0 0
Control, without
Ustilago 71 | 72 | 73 | 68 | 44 | 40 | 39 | 23

ur
Hordei |Colloidal sulphur in|
bottom of cells

1925]
TISDALE—TOXICITY OF COLLOIDAL SULPHUR 409

The results in this table are similar to those recorded in table
ш. The colloidal sulphur exhibits the usual degree of toxicity,
that is, the toxicity was greatest beyond Py 4.2, with freshly
prepared suspensions.

THE EFFECT OF DRYING AND AERATING UPON THE TOXICITY
OF COLLOIDAL SULPHUR

Since some growers prefer to use sulphur dusts, rather than
spray solutions, a test was made to determine whether colloidal
sulphur could be dried without destroying its stability and toxic
property. A colloidal sulphur suspension was prepared by the
SO.-H.S method, was coagulated with NaCl, and centrifuged.
One gm. of the coagulum was weighed on to a clean watch-glass,
spread over the surface, and allowed to dry thoroughly at room
temperature. After 5 days the sulphur was in the form of a very
fine powder. The dry sulphur was taken up with 200 сс. of
distilled water. The sulphur assumed the colloidal state readily
and the resulting suspension had a reaction of Py 5.0. Toxicity
tests were made in the usual Way, and the results are recorded
in table ут.

Two series of wash bottles were arranged to test the effect
of aeration upon the stability and toxicity of colloidal sulphur.
Freshly made colloidal sulphur preparations containing 1 gm. of
sulphur in 200 ec. of water and having a reaction of Py 4.0 were
transferred to the wash bottles. Series No. 1 was aerated con-
tinuously for 5 days. Air from which the oxygen was removed
with alkaline pyrogallol' was passed through the duplicate series
No. 2.

After 5 days the reaction of each sulphur suspension was
tested and found to be Ру 1.4 in Series No. 1, and Py 1.8 in Series
No. 2. The hydrogen-ion concentrations were adjusted to Py
5.0 with M/10 Na;HPO,, and toxicity tests were made in closed-
ring cells in the usual way. For a comparison a freshly made
solution of colloidal sulphur was also tested. Spores of Sclerotinia
cinerea and Botrytis Alii were used as indicators of the toxicity.
The results are recorded in table ут. In the table the dilution
of sulphur used was 1-20,000 for Sclerotinia cinerea and 1—2,000
for Botrytis Alii.

1 One part pyrogallol, 5 parts NaOH, and 30 parts H;O.

[Vor. 12
410 ANNALS OF THE MISSOURI BOTANICAL GARDEN

TABLE VI
TOXICITY OF PRE-DRIED AND OF AERATED SULPHUR SUSPENSIONS. THE
FIGURES REPRESENT PERCENTAGE OF GERMINATION (AVERAGE
OF DUPLICATE CULTURES MADE AT DIFFERENT TIMES)

Treatment of sulphur
Organism
Control, with-| Freshly Air-dried Aerated Aerated
out sulphur prepared +0, —0,
Botrytis Alii 71 15 10 2 8
Sclerotinia
cinerea 57 8 12 11 7

The results show that drying and continuous aeration for 5
days has no appreciable effect upon the toxicity of this form of
colloidal sulphur. Further tests will be required to determine
whether there is any influence from more protracted periods of
drying and aerating. Likewise, further investigations are neces-
sary to determine what acid or acids are responsible for changing
the Р. of the colloidal sulphur solution during the aeration.
It was found that the aeration of sulphur “flour” resulted in a
change in reaction toward the acid range within a very short
time.

THE INFLUENCE OF TEMPERATURE UPON THE TOXICITY OF

COLLOIDAL SULPHUR .

It has been shown by Doran (22) that the fungicidal value
of sulphur inereases with the rise of temperature. Butler (23)
found that the temperature was an important factor in controlling
snapdragon rust by the use of sulphur dust. Below 15? C. sulphur
was found to be ineffective.

An experiment was arranged to test the influence of a restricted
temperature range upon the toxicity of a colloidal sulphur
preparation. A suspension containing 1 gm. of colloidal sulphur
(SO;-H58 form) in 200 сс. of water was freshly prepared. The
reaction of this suspension was Py 4.0. A dilution of 1 to 8,000 was
made from this suspension in the slightly buffered solution
Py 4.2. This concentration of sulphur was known not to inhibit
germination of spores at ordinary temperature. "Toxicity tests
were made with spores of Botrytis cinerea at the temperatures
18°, 22°, 24°, and 28* C. Controls without sulphur were also

1925]
TISDALE—TOXICITY OF COLLOIDAL SULPHUR 411

run at the same time. These tests were made in duplicate and
conducted at two different times. The results are shown in
table vir.

TABLE VII

THE EFFECT OF TEMPERATURE UPON THE TOXICITY OF A COLLOIDAL
SULPHUR SUSPENSION. THE FIGURES REPRESENT PERCENTAGE
OF GERMINATIO

Temperature ? C.
Organism Form of sulphur
19 | 22 24 | 28
Control 67 | 70 81 | 40
Botrytis cinerea
Colloidal sulphur is | 22 "WW 1 3

Toxicity was exhibited at the lowest temperature tested,
though the higher temperatures increased the toxicity to a
considerable extent. It would be of importance to test the
effect of a wider range of temperatures and with a variety of
organisms.

PRACTICAL APPLICATION

After it was found that colloidal sulphur prepared by the
methods herein described was effective in inhibiting the germina-
tion of spores of certain economic fungi, as usually tested in the
laboratory, it was next necessary to demonstrate the fungicidal
value of these materials by practical field applications. Ar-
rangements were made with Dr. L. F. Nickell of the Monsanto
Chemical Works, East St. Louis, Ill, for the use of equipment
to manufacture a sufficient quantity of materials for field tests.

The materials were prepared according to methods already
described, with certain modifications to suit the available equip-
ment. A 500-gallon wooden tub, fitted with a steam coil, agitat-
ors, cover, and a flue provided with a steam jet were used for the

purpose.

The hypo-H;SO method was necessarily modified, since the
concentrated H.SO, could not be used in the tub because of the
iron agitator and steam coil. The method used was as follows:
Seven hundred pounds of hypo were dissolved in 75 gallons of
water and the solution warmed with the steam coil to 50^ C.
Ten pounds of glue were dissolved by means of steam and this

[Vor. 12
412 ANNALS OF THE MISSOURI BOTANICAL GARDEN

was added to the hypo solution. One hundred pounds of com-
mercial H;,30,, sp. gr. 1.84, were added slowly by means of a
siphon. The mixture was then adjusted to Ру 4.2 with sulfocide,
1 part to 5 parts of water. The agitator was kept in motion
during the entire process.

The lime sulphur method was slightly modified as follows:
Commercial lime sulphur solution was diluted with 4 parts of
water and the mixture warmed to 40? C. Ten pounds of glue,
previously dissolved by means of steam, were added for each
100 gallons of the diluted lime sulphur. The reaction of the lime
sulphur mixture was then adjusted to Py 4.2 with НМО,, 1 part
to 20 parts of water. The agitator was kept in motion throughout
the process. It was originally planned to allow the mixture to
set over night with the agitator in motion in order to free the
mixture of H;S. Due to the short interval of time to which the
writer was restricted in the use of the apparatus, another and
more rapid method was used to eliminate the H.S. Therefore,
the mixture was heated to 70? C. and aerated several hours by
means of an air pump. This aeration and heating unfortunately
destroyed the stability of the solution to a considerable extent.
Additional quantities of the mixture were prepared in a similar
way by using 1 to 20 H;SO..

The materials thus prepared were sent to the Missouri Agri-
cultural Experiment Station, and to Adams and Chester Counties,
Pennsylvania, to be used in tests being made for the control of
apple diseases. Mr. R. C. Walton, of the Pennsylvania Agri-
cultural Experiment Station, had charge of the field tests in
Pennsylvania.

Due to the seasonal conditions very little disease appeared in
the apple orchard at the Missouri Station. Therefore, no com-
parative results of value were obtained.

The results of the tests conducted in the commercial orchards
of Pennsylvania were very favorable, particularly in the case of
the preparation made from lime sulphur. These materials gave
very efficient control of apple scab, a disease which was severe
in Pennsylvania during that season. The details of these results
will presumably be published separately (Walton, ’26). The
injury to foliage and fruit was a minimum in the plots sprayed

1925]
TISDALE— TOXICITY OF COLLOIDAL SULPHUR 413

with these materials. The writer observed these plots in the
late summer, and it was then evident that the foliage of the trees
sprayed with the mixture prepared from lime sulphur and HNO;
was of a much darker green color. Furthermore, the growers
found the use of these materials satisfactory because of the
ease of handling and applying.

Colloidal sulphur was made in the laboratory by the SO;-H,S
method for field tests. The method used was the same as has
already been described except that a 10-gallon glass carboy was
used as a reaction tank for the SO, and НЗ. Large earthenware
jars (especially made for the Chemical Department, Washington
University) were used for НЗ generators. The product produced
by this method was sent to the Michigan, Missouri, and New
York Agricultural Experiment Stations, and to Adams County
and Chester County, Pennsylvania, for field tests.

'The results of these tests were not so promising as was antici-
pated, due probably to two factors: (1) The dilution of the sulphur
used was probably too great; and (2), as has already been indi-
cated, the sulphur crystallized in a short time after it was pre-
pared. The technique of large-scale preparation, therefore,
requires further detailed study.

CONCLUSIONS

1. Methods have been devised for preparing promising col-
loidal sulphur suspensions for use in praetical spraying.

2. The stability of colloidal sulphur preparations was found
to depend upon the method of mixing the chemicals, the tempera-
ture of solutions, and upon the introduction of protective colloids.

3. Of the emulsoid colloids that it has been practicable to
employ, glue has proved to be the most effective in preserving
the stability of colloidal sulphur suspensions.

4. The use of weak alkalies in adjusting suspensions prepared
from hypo and H;SO, was found necessary; and for this purpose
М№а,СО, and Na;HPO, gave the best results.

5. In preparing colloidal sulphur from lime sulphur, weak,
rather than strong, solutions of HNO; and H,SO, were more
satisfactory. !

6. The colloidal sulphur mixtures prepared by the meth

[Vor. 12
414 ANNALS OF THE MISSOURI BOTANICAL GARDEN
Ра

devised in the course of this work were found to be toxic to all
forms of fungi tested under laboratory conditions.

7. Greenhouse tests with a variety of plants have shown that
the products spread well and that they adhere to the foliage;
at the same time they do not cause injury under the conditions

ted.

8. Colloidal sulphur freshly prepared from SO, and H.S was
effective for the control of carnation rust in the greenhouse.

9. It would seem to be demonstrated that the toxicity of
different forms of colloidal sulphur may be ascribed to different
toxic substances. The properties of the toxic substances exhibited
by colloidal sulphur prepared from SO, and Н,8 are different
from the toxic substance (pentathionic acid) exhibited by Young’s
hydrophilic colloidal sulphur.

10. There is a gradual increase in toxicity of colloidal sulphur
formed from SO, and H-S, beginning at Ру 4.2 and continuing to
Ри 5.4, while at higher Py values there is a rapid increase. Young's
hydrophilie colloidal sulphur exhibits the greatest toxicity in the
range P4 4.2 to 5.4.

11. Colloidal sulphur prepared from SO, and HS in paste
form will crystallize in about 1 month. This crystallization
destroys the toxic property.

12. Colloidal sulphur preparations have greatest stability
between Pg 3.0 and 5.4. The loss of stability is accompanied
by a destruction of the toxic property.

13. The toxic constituent of colloidal sulphur, when freshly
prepared from SO, and H-S, is slightly volatile and is liberated
most rapidly at a higher Ру than 4.2.

14. Desiccation and aeration do not destroy the stability or
the toxic component of colloidal sulphur prepared from SO;
and H.S.

15. Colloidal sulphur prepared from SO, and HS is most toxic
at higher temperatures.

16. When manufactured for field spraying tests the methods
for preparing colloidal sulphur were modified to suit the equip-
ment available. Prepared in this way, certain of the products
б excellent control of apple всаЬ during the summer of

1925]
TISDALE—TOXICITY OF COLLOIDAL SULPHUR 415

The writer takes great pleasure in extending thanks for many
suggestions and criticisms to Dr. B. M. Duggar, who directed the
work; to Dr. H. C. Young, of the Ohio Agricultural Experiment
Station for many suggestions; to Dr. George T. Moore, for the
privileges and facilities of the Missouri Botanical Garden; and to
the Crop Protection Institute for the necessary funds and mate-

8.
Graduate Laboratory, Missouri Botanical Garden.

BIBLIOGRAPHY

qt eei > аа (1798). Observations sur l'hydrogéne sulfré. Ann. de Chim.
25: 233-27

Anania Е. нт Insecticides, fungicides, and weed killers. 431 рр. (see pp.
44—46). London, 1913. (Translated by D. Grant.)

Brinley, F. J. (23). а and properties of colloidal arsenate of lead.
Jour. Agr. Res. 26: 373-374.

Butler, О. (23). Experiments on the field control of snapdragon rust together
with a description of a method for the control of the disease in greenhouses. N. H.
Agr. Exp. Sta., Tech. Bul. 22: 1-14. f. 1-2. 1923.

Clark, W. M. (20). The determination of hydrogen-ions. 317 pp. pl. 1-2.
f. 1-38. Baltimore, 1920.

Débus, H. (’88). Chemical investigation of Wackenroder’s solution and explana-
tion of the formation of its constituents. Jour. Chem. Soc. 53: 278-357. 1888.

Doran, W. L. (’22). Laboratory studies of the toxicity of some sulphur fungicides.
N. H. Agr. Exp. Sta., Tech. Bul. 19: 1-11.

Duggar, B. M. “Bevery, J. W., and Schmitz, H. (17). Studies in the physiology
of fungi. V. The growth ve certain fungi in plant decoctions. Ann. Mo. Bot.

Galloway, B. T. (92). "oi work of the year in treating plant diseases. U.
S. Sec. Agr. Rept. 1892: 216-234. 1892.
Gillette, C. P. (90). Experiments with arsenites. Iowa Agr. Exp. Sta. Bul. 10:
401—420. 1890
Jones, P. R. (19). Combined insecticides and ee (U. 8. Patent No.
1291013). U. S. Patent Off., Off. Gaz. 258: 264. 1919.
Karrer, J. L., and Webb, R. У. (20). venerdi curves of certain liquid culture
media. Ann. Mo. Bot. Gard. 7: 299-305. f. 1-6. 1920.
Lewes, V. (81). On ИКБ acid. Chem. Soc. Jour. 39: 68-76. 1881.
Lovett, A. L. (18). Spreaders for arsenate sprays. Jour. Econ. Ent. 11: 66-69.
1918.
——_ de Insectieide investigations. Ore. Agr. Exp. Sta. Bul. 169:
1-55. f. 1-11.
— Е n, vow umet ee N. Y. Agr. Exp.

15: 543-544.
Moore, W. rtg елаз a adherence of arsenical sprays. Minn. Agr. Exp.
Sta., Tech. Bul. 2: 1-50.
Odén, S. (13). Der M Schwefel. R. Soc. Sci. Ups, Nova Acta, IV.
3*: 1-193. f 1-22. 1913.

[Vor. 12
416 ANNALS OF THE MISSOURI BOTANICAL GARDEN

Parker, J. R. (11). The use of soap to retard the settling of certain arsenicals.
Mont. Agr. Exp. Sta. Bul. 86: 35-45. 1911.
Raffo, M. (08). Ueber kolloiden Schwefel. Koll. Zeitschr. 2: 358-360. 1908.
msay, A. A., and Cooke, W. W. (22). The preparation of colloidal sulphur.
Agr. Gaz. New South Wales 33: 819-823. 1922

Riesenfeld, E. H. von, und Feld, G. W. ( 21). Polythionsauren und Polythionate.
Zeitschr. f. anorg. u. allgem. Chem. 119: 225-270.

Safro, V. I. C13). Ар investigation of аон и its causes and pre-
owes с Agr. Exp. Sta., Res. Bul. 2: 1-32. 1913.

У. M. (08). Self-boiled lime-sulphur mixture as a promising fungicide.
X S. Do t. Agr., Bur. Pl. Ind. Circ. 1: 1-18. f. 1-2. 1908.

Selmi, Fr. (52). Sur le soufre pseudosoluble, sa pseudosolution et le soufre mou,
Jour. f. prakt. Chem. 57: 49-57. 1852.

— — — —., et Sobrero, A. (50). Sur les produits de la décomposition des acides
sulfhydrique « gres au sein del'eau. Ann. de Chim. et de Phys. 28: 210-214.
1850. (Cited by Odén, ’13).

Shaw, " Msg On the preparation of pentathionates. Chem. Soc. Jour. 43:
351-355.

Spring, cr (82) Ueber die Nichtexistenz der Pentathionsaure. Lieb. Ann. d.
em. 29-363. 1882.

Stewart, J. P. (12). Preparation and use of the concentrated lime-sulphur spray.
Pa. Agr. Exp. Sta. Bul. 115: 1-23. f. 1-3. 1912.

Surface, H. A. ('05). Calendar for 1905 for treating insect pest ] plant diseases.
Div. Zool. Month. Bul. 3: 8-30. 1905.

Swingle, W. T. ('94). An improved method of making Bordeaux mixture. Jour.
Myc. 7: 365-371. 1894.
РИК W. W. (21). The chemistry of colloids. 332 рр. f. 1-22. (2nd ed.).

ndon, з

Van Slyke, L. L., Bosworth, А. W., and Hedges, C. С. (10). Chemical investi-
gations of best conditis for making the lime sulphur wash. N. Y. Agr. Exp. Sta.
Bul. 329: 405-449. pl.1. 1910.

—, amd Они FK. А, (0D. um — of commercial soaps in
relation to spraying. Ibid. 257: 427-438.

Vermorel, V., et Dantony, E. (10). s pes généraux qui doivent présider
à l'établissement des formules insecticides. Са Rend. Acad. Paris 151: 1144—

46. 1910.

Бана Н. (46). Ueber eine neue Saure des Schwefels. Archiv. d. Pharm.
47: 272-288.

иены, € Bestimmung des Sauerstoffgehalts der Pentathionsaure.
Ibid. 48: 140-151. 1846.

Agr. Exp. Sta. Bul. 288: 101-137. f. 61-69. 1910.
Walton, В. C. (26). Spraying experiments conducted in Adams and Chester
Pise meia with sulphur materials furnished by the Crop Protection
it

be published.)
Washburn, F. L. no Practical work with the codling moth and with a com-
bined insecticide and fungicide. Ore. Ore. Agr. Exp. Sta., Bul. 10: 1-34. 1891.

1925]
TISDALE—TOXICITY OF COLLOIDAL SULPHUR 417

Webb, К. W. (21). Studies in the pire of the fungi. XV. Germination of
the spores of certain fungi in relation to hydrogen-ion concentration, Ann. Mo.

Yothers, W. W., sd Winston, J. R. (24). Mixing emulsified mineral lubricating
oils with dogmai waters and lime-sulphur solutions. U. S. Dept. Agr. Bul. 1217:
1-5. 1924.

Young, H. z (22). The toxic property of sulphur. Ann. Mo. Bot. Gard. 9:
403-435. f. 1

‚ (25). Colloidal sulphur as a spray material. Ibid. 12: 133-143.
1925.

Zsigmondy, В. (09). Colloids and the ultramicroscope. 245 pp. pl. 1-4. f. 1-7.

New York, 1909. (Translated by J. Alexander.)

[Vor. 12, 1925]
418 ANNALS OF THE MISSOURI BOTANICAL GARDEN

EXPLANATION OF PLATE

PLATE 20

Effect of the hydrogen-ion concentration (Pq) upon the stability of colloidal
sulphur solution.

ed
z
в
ew
-
=
22
H
<
а
—
©
—
-
-
о
d
—
<
а
=

Oc ЗУД €c6I ‘ZI "10A "auvo чоя ‘ом "'NNV

= 4

GENERAL INDEX TO VOLUME XII

New е names of plants and the final ise wt of new combinations are

rinted in bold face

in o
A

— (Peniophora), 304
ida, 226

omnis ( (Peniophora), 366
ЖО Treubii, plastids of, 156, 175,

(Peniophora), 231
Allescheri (Gloeo ), 301
Allescheri (Kneiffia),
Allescheri (Peniophora), 301
alutaria (Peniophora)
Amide-nitrogen,
Amino-nitrogen, 369
Ammonia-nitrogen,
amoena (Peniophora), 276
Apparatus used in вора: co etermina-
n experiments, for the evaporation

кар es under vacuum,
мае ect of juices of, on pathogen-

icity of tobacco mosaic virus,

Apple scab, effect of sulphur treatment
on, 136, 412

arachnoidea OE нн

tea (Penio UR ‚ 346
Arisaema Pr um, tide of, 157,

juices of various p
er

0, 200
” effect of pokeweed
juices on.

‚36
asperipilaie (Peniophora), 20
— oe epiphylla, 24

( » 311
уени лог (Knei 311
aurantiaca (Раоа) 310
aurantiacum ( um), 311

Ann. Mo. Bor. Garp., Vor. 12, 1925

ace type; synonyms and page n
and plates, эа yo and previously published scientifio names and all ide: matter,
rdinary type.

rs having reference to figures

B

Bacterium одено effect of poke-
weed juice on, 363

borealis (Peni entum). №

Botrytis Alii, Me of d sulphur

certain forms

suspensions
Botrytis cinerea. eae o
of colloidal sulph

phur
Brunella ар) Т 124
Burkei орана) 0
Burt e. Thelephoraceae of
P North Ру, V, 218
Burtii (Peniophora), 278

C

смт (Peniophora), 353
m (Corticium), 353

cana (Peniophora), 227
(Peni и 296

stals obtained by ad ition с of “4

— (Решорһога), 854

carneum (Corticium), 354
carnosa (Peniophora), 325

Carotin, 178, 184

Carotinoids in fruits, ых studies
with special reference to, 145

on, 1
cinerea ic Corticium (Thelephora), 348
cinerea (K

cinerea (Peniophora), 348

(419)

420

cinereum (Corticium), 3

citrinella (Peniophora), 3 327
citrinellum (Corticium),

Citrullus "or 157, 0 pigment of, 166, 175;

plastids = -
ere з va (Peniophora), 253
neo-fulv у (Phlebia),
Colletotrichum of tolli ecl of S
tain anig sulphur о
389; toxicity ‘of

oxicity of,

pract tical application of, 411;
toxicity of volatile products of,
col eniophora

а method for the determina-
tion relative amounts of pigment

convolvens (Peniophora), 355

mentosum, 320; fumigatum,
348; giganteum, 216; glabrum, 214; in-
carnatum, 308; incons m, 221 :
laeve, 257; laevigatum, 338 ў

322;
29; subsulphureum 329; rues 317;
dn 339; ранын. 264; violaceo-
; viticolum. ‚ 322

pee effect of of juices of, on oe рее.

NS ng Mee порот), 280
тв phaenop: е ob-
tained by addition of ef of pe Si potash,
206; Bs) an rm; m ripe fruit, 157, 202
QUEM (Peniophora), Lid
crocicreas (

D

Daucus Caro 208;
171; natural SER cote apad
), 944

(Peniophors), 344
Determination of total
see Merge сина: in plants

[Vor. 12

ANNALS OF THE MISSOURI BOTANICAL GARDEN

Deven у matbad of nitrogen determina-
dis iseiformis var. borealis (Peniophora),
Dracaena Godseffiana, plastids of,157, 198

ne Karrer Arm-

uggar, B. M., and Joan
strong. The effect c of. treating the
ne = соран mosaie with the
f various plants,
бири: Мусы ы 298
Е

Eichlerianum (Corticium А pe

Mono ograph of
ab ar us Monardella, 1; rns on

South American Labiatae

alcoholic жив. 210; т of, 157,
exigua ыо рад 224
(Penioph ora), 239

F

farinosa (Knei
Farlow (Peuophora), 343
filamentosa icem 320

gicide i cre - 'а, 381
fusca. (Peniop ora), 244
usco-marginata (Peniophora), 335
G
Бахи (Peniophora), 222
um, effect of juices x on patho-
genicity of tobacco m virus, 362
Pons (Peniophora), 216
gigantea

ion гЬ. -

n germination of, 394

1925]

INDEX

Grapes, results of sulphur experiments
on, 142

guttulifera орана) 28

guttuliferu Lee end. 247

Glycine, 369

hete рое, 293

hiulos TP ves Aem ip A

Howard, E. Pigme ent studies
with ее reference to carotinoi
in fruits.

humifaciens (Penio phora), 225

Hydrogen-ion concentration: of various
juices, effec
virus

o, 398, 399, 400,
401, vli acd of аро stability
of Colloidal sul
Hydrogen prin ied and id acres а.
Dui a Ж colloidal sulphur by
Hypocknus M fo PES 229; Sambuci,
y
233; sulphurinus, 324

I
incarnata ( er чела сэ hae
incarnata (Kneiffia), 308
' incarnata (Penio hora), 307
incarnata (T.
incarnatum Corticiu m), 308
inconspicua (Peniophora), 2 221
$

ога),
tato, “effect of juices of, on
pathogenicity of tobacco mosaic virus,

Pia ophora),
isabellina oom чун A

J

n weed, effect of Mu tf on
poe ac P of tobacco m
Juniperi (Xerocarpus), 338

Karstenii (Peniophora), 254, 286
Karstenii ом, )

enio) ), 296
Kneiffia, 213; A 301; 7
311; ci 348; 261; fari-
226; incar 4 7
338; is, 257; ispora, >
ирон, 214; ү Ко 318; sub-
alutacea, 288; tenuis, 317; velutina,

421

L
laeve (Corticium), 257
1 A be ho xw
aevi eniophora),
pota. (Theleph

Lime sulphur, 134; preparation of col-
Pera sulphur from, 390; toxicity of,

limonia (Peniophora
livida зен iophora na xx

pora

longispora р 229

longisporus (Hypochnus), 229

Lonicera sp., color pigment in, 170;
plastids of, 157, 204, 206

ludo viciana (Peniophora), 244

color changes dur-
ing ripening, 187; 7; plastids of, 156, 200,

Lycopersicin, 159, 175, 178

tion of, 389, of tated
phur on germination of, 394;
toxicity of certain sulphur supensions

serrata, 48; thymifolia, 44; undulata,

76; villosa, . 46; viminea, 72: viridis, 55
(Peniopho ra), 238

Marrubium, 122; hamatum, 122; vul-

gare,
martiana борна), 330
martianum (Corticiu: 330
medioburiensis (Peniophora), 328
mexicana Verse mg a), 2
miniata (Peniophora), LN

hora), 277

Monar = fistulosa, 2; qt unct ata,
onardella, monograph of the genus, E
diagran: of relatio: i

2 78, 104, 106; б; Einer b5; epil

74, var. 4; "exilis. te
Pe ME 487 glabra. 59; pest 62.
— „6 һу leuca, 53, 100; in-
grata, 6: ucrata, 46; ata, 54,
100; 'Ianceolata 79 104, var. пуз:

cep

fera 84: ledifolia, 55; аи са 90,

106; ; linoides, 71, 102 „subsp. eulinoides,
bsp. stricta,

А › 96; macrantha, 35,
аг , 36, var. nana, 36, у
pinetorum, 37, var. tenuiflora, 35;
modocensis, 62; mollis, 48; montana, 2;
38, var Орос. 96, ver, МО

var. iphon, pa Ma var. ars
flora, Si р 52;
odoratisaima, 57 57, ' 102,
subsp. y“ * 4, 102, S bsp.

га-

tissima
94, 102, subsp. pallida, 6 102,
subsp. E us Fue Bh, 108 108, "Subs
» 68, 9 a um atissima, 59;

ei
106; C 62; reflexa, 50 ‘saxicole
М sanguinea, 79; 'saxico
; Sheltoni, 50; subserrata,
ymifolia, 44, 98; tomentosa, 48;
Pei s 37; ore 76, 104, var
"m villosa, 98, смо.

[Vor. 12

ANNALS OF THE MISSOURI BOTANICAL GARDEN

glabella, 50, var. leptosiphon, 36, 48;
viminea, 72; viridis, 55, 100

mon 237
Mosaie, tobacco, the effect of treating
the Boe of, with the juices of various

p
bnc (Peniophora), 299
mutatum (Corticium), 299

N

ee ый
оди і tion of total, in
plan Tene solutions, 367
nuda (Peniopho 345
nuda (T. ), 345
nudum (Comoe) 345

о

ochracea (Peniophora), 345
ochroleucum aru), ai
odontioides (Peni ophora), 223
odorata (Peniophora), 35

E nerochaete), 286, 289
odoratum (Stereum), 289

P

аса eus of sulphur experiments

Peckii (Peulonhese): 291

Peniophora, 213; admirabilis, 304;
Aegerita, 226; aemulans, 308; amnis,
266; alba, 297; albofarcta, 228
straminea, 305; albugo, 231; mom
231; Allescheri, 301; alutaria, 332;

amoena,

35;
gantea, 216; EN

ч
lerin 205. leprosa, 254 254; lilacina,

1925]

INDEX

limonia, 275; livida, 239; longispo
229; 1 udoviciana, 244; а

ella, 298; tenuis, 317; tephra,
237;

Thujae 236; typhioola, 319; re i
320; velutina, 250;
305; 285; viola-

ta,
ceo-livida, 347; LION 322; Weiri,

pertenue ue (Corticium), 315
M (Peniophom) 315
Fannia raia, 288 280 ), 273
p. orescens ophora),
Phlebia coccineo-fulva, 223
di (Peniophora), 241
t studies special reference
to carotinoids inf red. s 145
piliseta (Penio ), 242
pilosa Peniophora), 291

AP TR

в effect of j pm v) on patho-
genicity of tobacco mosaic virus, 359

radicatum inoue h 320
re а.
nitrogen т plants and plant

423

solutions: & comparison of methods
with modifications, 367

Ravenelii _(Peniophora) 269

rhodella (Peniophora), 254

HE
9i
an;
:

X пепі ів, 1 d

rugosa, color pigment in, 168;
plastids of, 155, 196 Н
Ко yog nus, 120; chilensis, 120; officinalis,
ate sob (Кш) 370 270
meguerit (Knei,
Roumeguerii ы, 270

S

Sacchari (Peniophora), 3
Salicylic-thiosulphate оа of nit
determination, 367; cation of,

eff
айк oidal sulphur on germina-
tion ion o ж. = lime-sulphur ve
on n of, 394; city of
certain Br ODIO
era Aegerita

а Cleon), 250

Sheari (Peniophora),
simile ns o 30026
similis (Peniophora
Sodium thiosulphate and sulphuric acid,
preparation of colloidal hur from,
camara, plastids of, 156,

ls of, 10 obtained by

NE AD |
addition of alcoholic potash, 206;
plastids of, 156, 202; pigment in, 169

424

sordida (Peniophora), 280
sordidum (Corticium), 280
Spectroscopic analysis of fruits, 176

т к, colloidal sulphur as а,

340; inum, 341
stratosa ( (Peniophora), 333
balutacea (Peniophora), 288
hal actum), pc
bapiculata (Peniophora), 280

(Cort " 322

Dcremea но ra), 30

oe (Peniophora), 250

ит
моги. (Peniophora), 329
C m), 329

е,
381; colloidal,
1^ 382; precipi-
m peus paratio: on of, 134
Sulphur ioxide and RAE sulphide,
preparation of colloidal sulphur by

means of, 395
Sulphuric acid and sodium thiosulphate,
preparation of colloidal sulphur from,

HUNE
5 3

Bre

sulphurina торлог), 8204
phurina [nir
rie yrs (Hypochnus j, 324
of juices of, on

tato, effect
"pathogenicity of tobacco mosaic virus,

324

F
—
Taxodii 'Peniop ora), 306
Temperature, uence of, upon the
toxicity of colloidal sulphur, 410;
relation of, to ripening of fruits, 186
(Peniophora), 298
tenue (Corticium), 317
е ( ium), 317
tenuis (Kneiffia), 317
tenuis (Peniophora), 317
tephra (Penio ), 339
tephrum (Corticium), 339
Teucri irm má 215; Cavenel-
; bicolor, av
lesii, 115; Chamaedrifolium, 112; chi-
lense, 115; к. 111, 5
, subsp.
» 113, 130, var. leucanth:
114; cubense, 1 3; Grisebachii, 113; het-

nudicaule, 11
tendi, 1 19, f 182; parti-
tem, 119 е in

ANNALS OF THE MISSOURI BOTANICAL GARDE

[Vor. 12, 1925]
N

Conticium, 348;
gantea, vate eet incarnata, 08; я
; laevis, 257; lilacina, 348; miniata,
277; nuda, a, 313; é
233; sanguinea, 274; sera, 233; velutina,
64; ; viticola, 322

me of North ‘America, The,

Thujae (Сеш һога), 28
E. ода!
укай апа Fagor id 1

ibtd.

the
wid
omatoes, "colorimetric method for de-

te rmination f relative amounts o
pigment in

Tomentella sulph

typhicola [Аучы Ачы 519

U

umbrinum рч), 341
unicolor (Pen a), 3
Ustilago Hordei, effect i lime sulphur

solutions on germination of, sin]
toxicity of certain pan pt
ms to, 404
V
velutina (Kneiffia),
Rk gore (Peony, 24

velutinum (Corticium

pigment in.
olaceo- livida qc я
olaceo-livida o oH

vi
vi

vi

viticola

viti ), ba
viticolum (Corticium), 322

Weiri (Peniophora), 342

peri, 338; subsulphureus,

X :
Yo Н. C. Colloidal sulphur as a
"i" material, 133

Ducem e 173, 178
erocarpus J’
329

zonata (Peniophora), 245