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University of California Publications in

BOTANY

« • •

VOLUME VII

1916-1922

EDITOR

WILLIAM ALBERT SETCHELL

UNIVERSITY OF CALIFORNIA PRESS
BERKELEY, CALIFORNIA

-■7

CONTENTS

PAGE

No. 1. Notes on the Californian Species of Trillium L., by Thomas Harper

Goodspeed and Robert Percy Brandt 1-24

I. A Report of the General Results of Field and Garden Studies,

1911-1916 1

No. 2. Notes on the Californian Species of Trillium L., by Thomas Harper

Goodspeed and Robert Percy Brandt 25-38

II. The Nature and Occurrence of Undeveloped Flowers 25

No. 3. Notes on the Californian Species of Trillium L., by Robert Percy

Brandt 39-68

III. Seasonal Changes in Trillium Species with Special Reference

to the Reproductive Tissues 39

No. 4. Notes on the Californian Species of Trillium L., by Thomas Harper

Goodspeed 69-100

IV. Teratological Variations of Trillium Sessile var. giganteum

H. & A 69

No. 5. A Preliminary List of the Uredinales of California, by Walter C.

Blasdale 101-157

I. Introductory statement 101

II. The need of experimental culture work 102

III. Plan of the present paper 103

IV. Acknowledgments 104

V. General features of the Uredinales 104

VI. List of species '. 105

VII. Index to the species of Uredinales 150

VIII. Index to the species of host plants 153

Nos. 6-8. A Rubber Plant Survey of Western North America, by Harvey

Monroe Hall and Thomas Harper Goodspeed 159-278

6. Chrysothamnus nauseosus and its Varieties, by Harvey Monroe

Hall 159-181

I. Introduction and acknowledgments 159

II. Key to the varieties of Chrysothamnus nauseosus 163

III. Synopsis of the varieties of Chrysothamnus nauseosus 165

IV. Index of specific and varietal names 181

7. Chrysil, a New Rubber from Chrysothamnus nau^eosu^, by Harvey
Monroe Hall and Thomas Harper Goodspeed 183-264

I. Object and scope of the investigation 184

II. Acknowledgments 186

III. Nature and properties of Chrysil 188

IV. Botanical classification; the names of the plants 189

V. Chrysothamnus nauseosus: habit, flowering, rate of growth,

etc 191

VI. Distribution and habitats of the various forms 194

VII. Estimates by districts of the amount of Chrysil available

in western North America 197

a. District 1. East Central California and adjacent

Nevada 198

b. District 2. Mojave Desert, California 200

c. District 3. Northeastern CaUfornia and adjacent

Nevada and Oregon 201

d. DLstrict 4. West central Nevada 202

e. District 5. Northern and central Nevada 204

/. District 6. Utah 207

g. District 7. Colorado 208

h. Estimates not included in the above districts 209

i. Conclusions as to the quantity of rubber obtainable 209

[i]

PAGE

\III. Mctliods of {l(>t(Htinn the presence of rubber and deter-

iiuninn its amount '•^1^^

a. Microscopical nu'thods '-^10

b. Chemical analysis 216

IX. Hcsultsof tlic chemical analyses and microscopical examina-
tions; talmlation of percentage of rubber in each variety 226

X. Distribution of rubtuT in the plant 234

(;. Hcfiional distribution in the ])lant 234

6. iSiM'cific distribution in mature tissues 239

XI. Factors influencing rubber content 243

a. Variation with the botanical variety. 243

b. Variation due to environment 24.'5

c. .Sea.sonal variation 246

Xll. Methods of harvesting: season, age, depth of cutting, etc... 248

XIII. Possibilities of Chrysothamnus as a cultivated plant 2.51

XIV. Cultural requirements 2.5.5

XV. Summary 2.57

8. The Occurrence of Rubber in Certain West American Shrubs, by

Harvey Monroe Hall and Thomas Harper Ctoodsjjeed 265-278

I. Chrysothamnus (exclusive of C. iiauseonu.s) and Haplopappus 265

II. Regional distribution of rubber in Haplopappus 274

III. Species in which no rubber is found 276

No. 9. Phycological Contributions. I, by William Albert Setchell and

5jathaniel Lyon ( Jardner ' 279-324

Xo. 10. Plantac Mexicanae Purpusianae. X, bv Townshcnd Stith Bran-

degee ' 325-.331

Xo. 11. Phycological Contributions. II to VI, by William Albert Setchell and

Nathaniel Lyon Gardner 333-426

II. New Species of Mj'rionema 334

III. New Species of Compsonema 353

IV. New Species of Hecatonema 377

V. New Species of Pylaiella and Streblonema 385

VI. New Species of Ectocarpus 403

No. 12. Notes on Pacific Coast Algae, by Carl Skottsberg 427-436

1 1 . On t h(> Calif ornian "Delesseria quercifnlin " 427

No. 13. Undescribcd Plants Mostly from Baja California, by Ivan Murray

Johnston 437-446

No. 14. The Morphology, Development, and Economic Aspects of Schizo-

phyllum commune Fries, by Frederick Monroe Essig 447-498

I. Introduction '. 448

II. Material and Technique 449

III. Morphology 451

1. General characteristics of the sporophores 451

2. Description of the mature sporophore 4.53

3. Microscopic structure 454

IV. Growth of the sporophore 458

1. Development in general 458

2. Origin and development of the "gills" 461

3. Taxonomic interpretation of the structure and devel-

opment of the sporophores 463

V. Economic aspects 464

1. Geographical distribution 464

2. List of host plants 46.5

3. Extreme hardiness of the fungus 4()()

4. Relation of the mycehum to cells of dead wood 467

5. Growths upon fresh wood and living trees 46<S

6. Methods of infection under natural conditions 470

7. Association with other wood decay fungi 471

VI. Summary and conclusion 472

VII. Acknowledgments 473

Literature cited 473

Explanation of plates 478

Index 499

[ii]

UNIVERSITY OF CALIFORNIA PUBLICATIONS

IN

BOTANY

Vol. 7, No. 1, pp. 1-24, plates 1-4

October 3, 1916

NOTES ON THE CALIFORNIAN SPECIES OF

TBILLIUM L.

I. A REPORT OF THE GENERAL RESULTS OF FIELD AND
GARDEN STUDIES, 1911-1916

BY

THOMAS HAEPEE GOODSPEED and EOBEET PEECY BEANDT

For the past six years a study of the species of Trillium peculiar
especially to Western Middle California has been carried on under
the general supervision of the senior author with a number of assist-
ants. The investigation was originally undertaken at the suggestion
of Professor W. L. Jepson, who pointed out the remarkable range
of color variation in Tnlliuni sessile var. giganteum H. & A. and
suggested that it might be possible to segregate color forms geograph-
ically. It was also suggested that garden studies might throw light
upon their origin. Interest at the start thus centered about the range
and origin of color variation in T. sessile var. giganteum ; the extent
of the investigation necessary in this connection has brought up a
large number of problems seemingly of equal or greater interest.
Indeed, it has been found necessary to extend the investigation to
include comparative studies of T. ovatum Pursh. The present report
aims to treat very briefly of the sj^stematic position of the various
species of Trillium native to California and to include a description
of the general results of field and garden studies primarily upon
T. sessile var. giganteum.

Field studies of var. giganteum have covered rather thoroughly
the territory which is fairly definitely limited by the term Western
Middle California.^ That portion of the Coast Range which borders
upon San Francisco Bay on tlie east and may be spoken of as the

1 Jepson, W. L., Flora of Western Middle California, San Francisco, 1911.

2 Univcrsilj/ of Califnniid I'lihlications in Botany [Vol.7

]iorkeloy and Oakland Hills lias, by reason of its accessibility, received
principal attention. Exten.sive .studies liave, however, been carried
on to the north in .\apa Valley (Napa County), in Southern ]\lendo-
cino County and in ^Tarin County. To the south certain stations
in Monlcicy ('onnty. Santa Cruz County and San ]\Iateo County have
received atti'ution and ratluT thorough study. Our own field studies
have Ironi time to time been supplemented by examination of plants
sent in by a considerable number of correspondents. Collections by
the hit t IT have added Sonoma County to the above list and have
furnished valuable comparative material from stations in the .same
general regions from which our own collections have come. Beyond
the range indicated above, some study has been made at stations near
Dutch Flat. I'lacer County (3500 ft.) and at Crockers, Tuolumne
County (4500 ft.).

Two seasons, a dry and a wet season, comprise the range of
climatic conditions peculiar to that portion of California with which
we here are dealing. In general the month of January, which includes
the period of greatest precipitation, is the coldest, whereas during
August, the driest month, highest temperature conditions usually
prevail. The two tables below, Table 1 for San Francisco and Table 2
for Berkeley, give the approximate average rainfall in inches month
by month in the region surrounding San Francisco Bay. As may

Table 1

Jan.

Fob.

Mar.

Apr.

May

June July

Aug.

Sept.

Oct.

Nov.

Dee.

4.89

3..50

3.34

1.64

0.73

0.16 0.02
Table 2

0.02

0.32

1.00

2. .55

4.51

Jan.

Feb.

Mar.

Apr.

May

June July

Aug.

Sept.

Oct.

Nov.

Dec.

5.81

4.1.5

4.95

1.49

1.15

0.22 0.02

0.04

0.57

1.48

2..54

4.20

be noted in the above ta])l('s, approximately three-fourths of the annual
rainfall occurs in the months from December to March inclusive.
The figures for July and August refer to heavy mists and slight July
rains, which sometimes occur. Precipitation in practically all cases
consists of storms of one to thirty or more days' duration, interspersed
with periods of clear, dry weather. Precipitation in spring decreases
as the storms become shorter and less severe and the intervening
periods of clear weather increase in length. In the northern part
of California the rainfall is much heavier than in the central part,
and very appreciable precipitation may occur in July. Such con-
ditions are also characteristic of the Sierra Nevada. In Southern

1916] Goodspeed-Brandt : Notes on Trillium 3

California, on the other hand, the rainy season becomes shorter, and
the lessened precipitation consists of a few drenching rains of short
duration. In addition to the actual precipitation, sea fogs more
or less relieve the drought of summer along the Coast. With the
exception of the Sierra Nevada, the state is free from severe cold
in winter. Under the heavy forests along the northern coast and
even under chapparal in the south, frost rarely occurs.

In the Sierra Nevada and along the northern coast high evergreen
forest prevails, with low trees and chapparal on high, dry ridges
where the soil is shallow and poor. During heavy summer fogs the
trees in the coast forest drip very freely. Much of this water is
retained, so that the soil is visibly moist at the surface as late as the
first of August. Such plants as can endure shade are found here.
Trillium ovaUmi Pursh. is one of these. It grows on flats and on
both northerly and southerly slopes wherever the soil is sufficiently
moist and fertile. It does not grow in the dry chapparal nor on those
hot mountain tops where the trees are tall but widely separated. In
recently cleared or burned-over areas this plant grows luxuriantly,
sending up two shoots a season from a single rootstock and prac-
tically always maturing fruits and viable seeds. It does not multiply
in such localities, however, because of the use of the seeds for food
by animals and birds and because the seeds which are dropped are
unable to germinate on account of the extremely dry condition of
the surface soil. Toward the inner boundary of the redwood forest,
where the influence of fogs is lessened, the forest tends to retire to
northerly slopes. Trillium ovatum is found only well within the
limits of the forest in such locations. Still farther inland the species
disappears, to reappear in the northern Sierras.

In certain localities in the Redwood Belt which are exposed to
dry winds or have soil too shallow for large trees, the forest is replaced
by grassland and thickets of deciduous shrubs and large herbs.
Trillium sessile var. giganteum is peculiar to such associations. This
species may also be found near the edges of groves of redwood but
never in the dense shade furnished by a close stand of these trees.
It also occurs beyond the limits of shrub formations, but rarely in
the drier grassy areas.

In Central California the prevailing formation is grassland on
southerly exposures, and deciduous shrubbery on northerly slopes with
occasional groups of low-growing deciduous or thick-leaved evergreen
trees. Such trees and shrubs also form a fringe along streams. Large

■1 I'liin rsil If of ('(ilifuniid I'lihJicaiions in Botany [Vol. 7

pereniii.-il hci'lis .-n-c round (•(i\ci'iii<j- the <_m-()uiii1 iiii<lcr llicse trees and
shrubs. Trilliitni sessile var. (jiyanh iiin ^m-ows in <fi'c;i1 ahimdanee
ill lliis rog:ion, l)\i1 (lisapjx'ars some eight miles east of S;iii Francisco
liny, reappearing in Ihe middle Sici-i-;! Nevada. In addition to the
prevailing format ioii.s noted abovr. I'cdwood forest occurs in unusually
moist canons of CentT-nl Cnlifoi-nia, such as those on Mount Tamalpais
and in San Ticandro ('anon. Trillinm ovahdii regularly appears in
these localitii's. soiucl iincs in great abundance, while T. sessile var.
gignnteiim occurs only on the edges of these groves.

Three species of Trillium are with certainty to be distinguished
as occurring in California. The total range of all the west coast species
extends at least from Washington south to San Luis Obispo County,
California. The pedunculate species of California are two — T. ovatum
Pursh.. and T. nivale Watson. The latter species is not too well
known but is reported as characterized by a short, slender habit,
petioled leaves, long pedunculate flowers and in general to be "allied
to T. nivale Riddell, which it much resembles in habit." This species
we have not collected and is not to be dealt with in what follows.
Trillium ovaUim is sometimes referred to as the west coast form of
T. grandiftorum (]\Iichx.) Salisb., from which, however, it is in fact
distinctly set apart. In height ovafum ranges from 6 to 30 cm. from
the surface of the ground, the height of the great majority of the
plants falling between 20 and 26 cm. Where the natural cover has
disappeared, as in burned-over areas, the west coast species of Trillium
are greatly reduced in height but vegetative and floral organs are
not diminished as to size. In general, height of plants seems scarcely
worthy of notice in this genus where the character can be varied at
will. This i)oint is emphasized in the reaction shown by the ])lants
figured in plate 1, figure 2. All the four plants were kept under a
shed which was open at one side. The two middle plants were grown
near the open side and were fully and on all sides almost equally
illuminated, while the two plants at either end were grown toward
the back of the shed in greater shade and illuminated from one side
only. During the year preceding and in the year following when
all were growing under the same conditions their height was closely
corresponding. The above is mentioned here not because the
observed reaction is other than the one to be anticipated, but because
the difference in height and habit of the two groups is out of all
proportion to the relatively slight difference in growing conditions,
in other words, plants of Trillium species appear to be so markedly

l^l*^] Goodspeed^Brandt: Notes on Trillium 5

affected by variation in illumination that height, general habit and,
as can be seen, the presence or absence of a slight petiole, are not
to be taken as reliable criteria for species differentiated.

T. ovatum is thoroughly described by Jepson,- and only a few
points are worthy of special mention. In color of flower this species
is characteristically a chalky white and the lack of variation in color
is striking in connection with the extreme variability of T. sessile
var. giganteum in this character. After anthesis and while fading
the petals become distinctly pigmented to such an extent in many
cases that the petals may become deep rose-red as they Avither.
Herbarium specimens of this and other white-flowered species seem
to show a considerable variation as to the presence or absence of this
pigmented condition. According to our preliminary observations on
this matter, it appears that a plant collected before pollination or
fertilization may not exhibit the pigmented condition when dried,
but that when collected after pollination or fertilization the color will
appear on drying even though not apparent when collected.

Turning now to T. sessile var. giganteum, there seems to be no
doubt that this Californian plant represents at least a very distinct
modification of the eastern T. sessile. The difference in size between
the eastern and western forms is very striking and it may ultimately
seem desirable to accept the western form as a distinct species under
the name T. giganteum (H. & A.) Heller. T. sessile var. giganteum
is, again, fully described by Jepson,^ but a number of points,
especially with reference to its variability in certain characters, are
worthy of note. Characteristic individuals of the Californian variety
of T. sessile just coming into full flower are shown in plate 1, figure
1, and in plate 3, figure 6, a vigorous and characteristic plant fully
in flower. The very considerable size of all characters may be seen
by reference to plates 1 to 3. As noted below, we have undertaken
a rather extensive statistical study of the size relations of vegetative
and floral characters and it is hoped that measurements of the
eastern T. sessile can also be obtained. Since our data are not
entirely complete and we do not know the exact limits of variation
of corresponding characters peculiar to eastern species, it seems best
to discuss only in very general fashion the variation in size of the
leaves and flower parts of T. sessile var. giganteum.

At the start it might be noted that for all characters the limits

2 Op. cit., p. 108.

3 Op. cit., p. 109; cf. Garden and Forest, vol. 3, p. 320, 1800.

6 University of California Publications in Botany [^ol. 7

of size variation of the eastern and western forms of T. sessile prob-
ably overlap slightly, but also that the average individual of the
eastern form is throughout strikingly diminutive as compared with
its Californian variety. Tluis, with reference to length of petal,
a very small percentage of the more than five hundred flowers meas-
ured have fallen below 4 cm., while 6 to 7 cm. represents perhaps the
average length and many measurements show a length of over 8 cm.
The relatively few specimens of the eastern sessile examined give

4 cm. as the maximum and 1.3 cm. as the minimum length of petal.
One other size difference might be mentioned, i.e., leaf size. Accord-
ing to our limited observation, the eastern species bears leaves not
over 8 cm. long and always longer than broad, while the smallest
specimen measured showed leaves only 4.2 cm. in length. Possibly the
largest leaf of the western form observed was 17.5 cm. in length
and its greatest width was 19.5 cm. An excess of width over length
of leaf is not unusual and may even be characteristic of tlic ])lants
from certain localities. The minimum length of leaf approximates

5 cm. in one specimen. In this matter of leaf size we are referring
only to normally flowering plants of mature structure. It may in
general be said that T. sessile var. giganteum exhibits a most remark-
able range of variation as to the size of all characters both vegetative
and floral. This situation is illustrated in plate 2, figures 3 and 4,
and is sufficiently striking to make unnecessary further comment here.
Variations in form and shape of corresponding characters are brought
out in these same plates. With reference to this matter of form and
shape variation we again prefer to postpone the presentation of the
complete data until it may be possible to consider together in full
the relation of size, shape and color forms peculiar to definite localities.
That the range of variation in shape and form of vegetative and
floral organs is as great as their size differences should be sufficiently
apparent from the specimens shown in plates 3 and 4. An extreme
variation with respect to width of petal is the {T. sessile) variety
angustipetalum of Torrey. This extremely narrow-petalled form
represents simply a point in the very wide range of variation in petal
width peculiar to the species in California, although it may, as
reported, be characteristic of certain definite localities — i.e., the
Sierra Nevada — though on plants growing at 3500 feet near Dutch
Flat, Placer County, the petals were rather broadly ovate.

With reference to the nature of the apparent gigantism of T.
sessile var. giganteum we have some more or less fragmentary cyto-

1916] Goodspeed-Branclt : Notes on Trillium 7

logical data. At the suggestion of Professor R. Ruggles Gates we
attempted to determine the relative cell size and the relative
chromosome number and chromosome size of the eastern and Cali-
fornia varieties of T. sessile and to make the same comparative studies
in the case of T. grandiffornm and the rather closely allied T. ovatum.
This investigation is still in progress, but it appears desirable to
mention such results as are at hand. It seems certain that there
is no doubling of the chromosome number, as in Oenothera gigas,
giant races of Primula sinensis, Spiranthcs cernua, etc., in the
California variety of T. sessile, nor is there any distinction in
chromosome size or constitution between the eastern and Californian
varieties of this species. Similarly we have been unable to note
any increase of cell size in the Californian as compared with the
eastern T. sessile, though the measurements made are too few to allow
of a definite conclusion in this connection. Trillium grandiflorum
and T. ovatum are not set apart by conspicuous size differences and
in cytological details they correspond rather closely one to the other.
The chromosome number in T. sessile var. giganteum was determined
by a study of root tips and of the maturation divisions of the micro-
spore mother-cells, while the tentative conclusions as to comparative
cell size are based upon measurements of cells of the root tip.

As stated above, the entire investigation in progress dealing with
the Californian species of Trillium had its origin in an effort to map
the distribution of particular color varieties of T. sessile var.
giganteum and to trace their relationships. Our field studies have
led us to distinguish three main color types within the Californian
form of sessile. First, a white-flowered form which is peculiar to
the northern portion of our district and which only in isolated cases
occurs elsewhere. So far as color of flower is concerned and also
with respect to some other characters, this form corresponds to the
species described as Trillium chloropetalum by Howell. We have
studied it most thoroughly in Napa Valley, Napa County. In caiions
of the hills on the sides of the valley and in the nearby Napa Range
at an altitude of 2000 feet this pigmentless form occurs in rather
great abundance. The petals are dull white, somewhat greenish at
the bases, and the flower parts within are light green throughout
with the exception of the anther coats, which are a light yellow.
This form has been sent in from Sonoma, Sonoma County, and appar-
ently occurs also in Lake County. White-flowered individuals only
occur near Dutch Flat, Placer County, and may be characteristic of

8 University of Califcnild I'lihlirdlidiis in liolaiin [Vol. <

the Sierras at altitudes of from 3000 to 4500 feet. On :\rount St.
Helena, however, a ])eeiiliarly deep garnet-petalled form is reported
and it is very desirable that field studies be made at tlie head of
Napa Valley on 1lie line along which 1h<' 1\vo eolor types come into
contact. A distinelly wliile-petalled form is characteristic of Wash-
ington, Oregon and of nortliern California, and Napa Valley appears
to represent the southern limit of its range. In the main all the
northern white-petalled plants correspond to those in Napa Valley
in showing no pigmentation of flower i)ai'ts, although many plants
from southern Mendocino County with pure white petals show
evident traces of deep red pigmentation on the stamens and pistil.
Isolated individuals which bear flowers entirely colorless have at times
been found south of Napa Valley, but they seem best considered as
representing a stage in the transition from undeveloped to normal
flower which will be discussed in the next paper of this series.

The second distinct color form is one involving variation abont
a greenish*yellow shade of petal color and is found in the outer
Coast Range from Sonoma County south to Monterey County. Plants
with green, greenish-yellow, dull orange or reddish-brown petals were
first found on Point Reyes peninsula, which is perhaps the northern
limit of this color form. It is found predominating in eaiions at
the base and south of Mount Tamalpais in Marin County and at a
number of stations in Santa Cruz County. Within this color form
there is a striking absence of white or very light green petalled in-
dividuals, though deep garnet flowers are not infrequently found.
Among the lighter-colored flowers of this form a brilliant viridine
yellow is not an uncommon shade, while in the green-flowered forms
the leaves, sepals and petals are indistinguishable so far as eolor is
concerned. In plate 2, figure 3, flowers of Marin County representative
of this second color form are shown. The extremes w^ere included for
comparison and no attempt was made to indicate the predominance
of the lighter color shades. Plate 2, figure 4, from a photograph of
dried specimens and included to exhibit size variation, indicates
nothing as to the range of color forms to be found in the third eolor
form described below and to which the original plants belonged.
Throughout the range of variation in coloi- of this second form, the
stamens and i)istils uniformly show a greater or lesser amount of
pigmentation.

The third color form is in general peculiar to caiion.s of the hills
sui-rounding San Fi-ancisco P)ay on the east and southwest — i.e.,

1916] Goodspeed-Bmndt : Xotes on Trillium 9

San Francisco County, the western edge of Contra Costa and Alameda
counties and northern San Mateo County. The following points seem,
in general, to characterize the plants from the region so far as petal
color is concerned. First, the strictly pigmentless northern form is
not represented; second, the yellow-green color form is not found;
and thus, third, every flower is characterized by the presence of red
pigmentation on some of its parts. We have here a most remarkable
series of color variations, from flowers with petals white to the base
but with deep red stamens and pistil to flowers with all the flower
parts except anther sacs and stigmatic surfaces of such a deep garnet
shade as to appear black (cf. plate 3, figure 6). As has been noted,
every significant fiower bears some pigmented parts. Delicate rose
pink, violet and even bluish shades are common in this color form.
The origin of these three rather sharply distinguishable color
forms and their relation to one another is in doubt. The extremely
wide variation of the last-mentioned form has suggested original
hybridization between the white-flowered northern form and the
more heavily pigmented southern forms. There is, however, little
doubt that crossing between the plants which at present make up
the series does not take place and probably in but very few instances
does anything but asexual reproduction increase the number of indi-
viduals because of the almost universal occurrence of partial or
complete sterility. This wide range of variation in form three may,
perhaps, be looked upon as representing normal variation from a
pigmented type. Our garden studies of the three color forms give
no evidence of variation in color of flowers produced from the same
rootstock in different years, and the results of the field studies indicate
no variation among the plants vegetatively produced as offsets from
a parent rootstock nor variation in color as compared with the parent
plant. The complete sterility, indicated by the entire lack of seed
production, of the plants in the garden cultures has made it impossible
to cross widely separated color forms. We have, as will be noted
below, secured seeds of a number of color types of form three which
have proved to be viable. There is thus a possibility of obtaining
data on the amount of variability of the progeny of a given color form.
The three color forms and their varieties taken together may be
thought of as the result (1) of an absence of any cell sap pigment
or chromatophore color, (2) the presence of anthocyanin derivatives
in cell saps of various reactions which may (3) be present in con-
nection with red, orange or yellow chromatophores or (4) green

lU Univcrsilij of Calif urn ia Publications in Botany [Vol. 7

cliloniplast coloration alone An investigation from this p<iint of view
ol' the nalni'i' ni' tin- vai'ions coloi's concerned is in ])rogress.

These color forms which seem to be rather strikingly peculiar to
certain definite localities might appear superficially to correspond
to one anotlit'i' in cliai'acli'rs which are of more significance from the
point of view of specific differentiation — i.e., size and form characters
of vegetative and fioi-al organs. We feel rather certain, however,
that more exact (juantitative studies will, in some cases at least,
reveal correlation between color and other characters, the sura total
giving sufficient differentiation fi-om all other forms possibly to
indicate specific definition and to warrant specific description. With
this in mind we have measured all significant structures whenever
collections have been made. The criteria of specific differentiation
in the genus Trillium are not such that simple size differences of
vegetative or floral organs, or even proof that the correlation in size
of organs is different in the various color forms, will give convincing
evidence on relationships. Such evidence will depend upon demon-
stration of shape and form differences, and thus it is desirable to
find a method according to which the measurements of size of organs
can be used in expressing their shape. It is also highly desirable
that the tentative use of the combination T. sessile var. giganteum
which later on may seem best changed to T. giganteum, both repre-
senting efforts to recognize the size distinction between eastern and
Californian sessile forms, be placed upon a demonstrable basis. It
is thus planned to compare the size differences of these geographically
distinct sessile species, and until this is done we do not propose, as
noted above, to present the statistical data at hand.

Rootstocks of T. sessile var. giganteum were first brought into
the University of California Botanical Garden in the spring of 1911.
In February, 1916, the total number of rootstocks under observation
was nearly 150. We have data on sixty plants over a period of six
years and on forty plants over a period of four or five years, while
the remainder were introduced into the cultures after 1913. The
various localities where, as mentioned above, field studies were made
are practically all i'ej)resented by rootstocks in the Garden. In May,
1914, we secured rootstocks of T. sessile L. from New Lennox, Illinois,
and rootstocks of T. grandiflorum (Michx.) Salisb. from Sawyer.
Michigan. During the past two growing seasons we have had a
few flowering and a number of flowerless shoots from these rootstocks.
From time to time a few rootstocks of T. ovatum Pursh. have been

1916] Goodspeed-Brandt : Notes on Trillium 11

iutrodiiced into tlie cultures, but we have experienced considerable
difficulty in keeping them alive.

The rootstocks have, in practically all cases, been grown outdoors
in pots. It was impossible even to approximate the soil or environ-
mental conditions from which the different rootstocks had been taken
and all were grown in rather light garden soil and under identical
cultural conditions. The rootstocks have been repotted yearly,
usually in August. The conditions under which the garden cultures
have been maintained are the best at our command, but are certainly
not the most favorable and may even be the determining cause of
certain observed conditions to be described below.

Perhaps the most significant fact concerning the nature of the
species of Trillium under observation through field and garden
studies has to do with the occurrence of undeveloped flowers in those
plants which appear to be completely flowerless. This matter is to
be taken up separately in the second paper of this series and here
it is not necessary to do more than mention the points established.
Where a considerable number of Trillium plants grow together in a
relatively limited area, a glance reveals the fact that some are in
flower and that other plants, perhaps by far the larger number,
appear to be entirely without flowers, while a few exhibit small,
unopened buds in the center of the whorl of leaves. It has, however,
been found that every plant contains a flower of a greater or lesser
degree of differentiation or development. At one extreme are entirely
undifferentiated primordia of microscopic size and at the other are
the buds mentioned above which never open and in which the petals
are always white and early withering, the pistil suppressed or the
ovary lacking ovules, and the anthers without functional pollen.
Between these two extremes we have found all intermediate stages.
Study of the garden cultures has aided in pointing out the significance
of this situation. In only a few instances in the cultures has it been
possible to keep the shoots produced from the rootstocks in a normally
flowering condition from year to year. Thus, though the great
majority of the 150 rootstocks under observation bore a flowering
shoot or shoots when collected or during the growing season previous
to collection, we have at present scarcely a half dozen rootstocks
producing flowering shoots. Usually in the year following collection
the shoot or shoots produced contain undeveloped flowers.

Our field studies have in part been concerned with an investiga-
tion of the various degrees of sterility exhibited by var. giganteum,

12 University of California Pnhlications in Botany [^o^. 7

a condition in i-atlicr strikinf? contrast to lliat exhibited hy T. ovatum
whieli almost witlioiit cxccpi ion produces an abundance of seed. In
California it is diftieult. unless llie plants are staked in the spring
wlien in leaf and flowei'. to liml tlieir drying remnants when they
are covered witli oilier larger species wliich have dried down over
them. Tt has, iiowever, been apparent throughont our field and
garden studies that many ]>lants of sessile do not bear fruit either
in a given season or during a number of seasons. Indeed, for some
years we did not succeed in finding a single fruit containing seeds,
though many plants known to have ])roduced apparently normal
flowei-s were under observation in the tield as they ripened down. Fur-
tliei'. many other- plants examined near tlie close of their active sea>;on
above ground were found to mature only a shrunken, undeveloped
pistil provided with rudimentary ovules, though these same plants had
been strong in appearance and vigorous in growth and at anthesis
were provided with pollen and ovules normal in appearance. Again,
there are still other plants in the flowers of which at anthesis the
pistil is entirely undeveloped. The undeveloped flowers referred to
above represent the extreme case of sterility accompanied by the lack
of development of all the flower parts. Finally, the almost complete
absence of seedlings and young plants, other than those derived
vegetatively from older rootstocks, added evidence in support of the
contention that lack of seed production was the rule. These earlier
observations were confined to stations in the Berkeley hills which by
reason of accessibility were favorable for following closely the more
obscure periods in the annual life-cycle of the plants. The white-
flowered form of var. giganteum characteristic of the northern coast
counties has moi-e recently been found to seed rather profusely in
certain localities or under certain conditions. More recently, also,
a number of i)laiits of sessile in the Berkeley hills have been found
to give viable seed, particularly favorable seasons climatically being
perhaps responsible. In general, however, it can be said that sessile,
in contrast to ovatum, is an extremely uncertain producer of seed
and that a very considerable number of individuals are not capable
of seed production over a course of a number of years.

The fact, as noted above, that plants which apparently produced
normal sexual elements failed to form fruit led to an experimental
and cytological investigation of the causes of sterility in such cases.
In the garden a considerable number of flowers were self-pollinated
at different periods before and after anthesis. Cross-pollinations,

1916] Goodspeed-Brandi : Notes on Trillium 13

also, were made between plants of different flower color and many-
attempts were made in which pollen of ovatum was used on sessile.
Pollen of sessile will give a fairly high percentage of germination in
tap water. Still it seemed possible that the erect, open flower of sessile
borne on a plant which usually grows in open thickets might be
subjected to too excessive moisture conditions at pollination and thus
flowers in the field and in the garden cultures were protected in
various ways during and for a time after pollination. The lack of
any positive results following these efforts seems to indicate that
moisture conditions are not the determining factors in this connection.
It seemed possible that in the rapid growth of the vegetative portions
of the shoot the food reserves of the rootstock were depleted
sufficiently to make it impossible for the final stages in the maturing
of the sex cells to take place normally. With this possibility in mind
we cut off one, two, or all three of the leaves when still undeveloped.
This treatment was of no value in increasing the amount of normally
matured reproductive tissue but, on the other hand, did not appear
to interfere with the growth of the stem or the increase in size of
the flower bud. The rootstock was, in other cases, cut away from
its growing apex at different stages of development of the floAver
shoot, with the thought that such shock might provide a stimulus to
more normal fruiting. In such cases over two-thirds of the rootstock
was removed and the plant consisted of nothing but the aerial shoot
rising from a small disc of tlie rootstock. The desired effect was not
produced by this treatment but, on the other hand, in only a few
cases did the parts of the plant above ground show any effects of
the mutilation of parts below ground. Etherization experiments with
rootstocks were attempted during the dormant season with the thought
that an enforced re.st period at this stage might result in the pro-
duction of more highly fertile flowers. The spores are being
matured throughout the dry season when available moisture is at
a minimum and it seemed possible that the maturation of the sex
cells might be more successfully carried through if the buds in the
rootstock crowns were forced to remain inactive until the opening
of the wet season. On the other hand, there is, after etherization,
often a stimulation to abnormally rapid growth and early develop-
ment of flowers. A considerable number of rootstocks were treated
but no apparent effect in either direction was noted.

As mentioned below, a rootstock may be divided many times and

14 University of California Publications in Botany [^ol. 7

segments under 5 mni. in tliickness Avill produce shoots and increase
in size normally from year to year. We attempted to graft all
sizes of rootstock segments of sessile onto one another and to unite
portions of rootstocks of this species and of ovatum. A possible
increase in vegetative vigor of the grafted rootstock was anticipated.
It was also hoped that, in the case of interspecific graft unions, an
offset might be produced at the point of contact. A few successful
unions were obtained in both cases, but no visible effects could be
noted during the first year in the case of the shoots which were
normally produced from the crowns of the treated rootstocks. No
offsets were formed during this first year, and the experiments were
terminated because it was not possible to give the grafted rootstocks
sufficient care to prevent the drying out of the graft unions during
the dry season.

Seeds of T. sessile var. giganteum have from time to time been
obtained, both of the white-flowered form of the northern counties
and also from plants in tlie Berkeley hills. Up to this past year it
has not been possible to secure germination of any of this seed,
although in all cases it appeared to be entirely normally matured.
The seed coverings were broken and softened by mechanical means
and by the use of sulphuric acid and were subjected to a variety
of growing conditions without securing a single germination.
Recently seeds from plants growing in the Berkeley hills have germ-
inated over 60 per cent. Apparently seed of the western sessile will
not germinate for eighteen months or longer if placed in the ground
as soon as mature, but if allowed a period of six to twelve months
or more for after-ripening will then germinate almost at once Avhen
placed under ordinary germinating conditions. The failure of our
earlier attempts at germination of this seed was due to a termination
of the experiment long before germination could have taken place.
Further experiments are in progress to confirm and amplify these
preliminary observations. Various stages of development in seed-
lings of tlie first year are shown in plate 4. The third and fifth
seedling from the left show the seed coats still attached.

Nothing has been more striking throughout our field and garden
observations than the extreme development of vegetative propagation
in T. sessile var. giganteum, a condition which appears to be
accompanied by or a result of a highly defective power of sexual
reproduction. T. ovatum, on the other hand, very rarely forms a new

1916] Goodspeed-Brandt : Notes on Trillium 15

individual asexually and is correspondingly a consistent producer of
large numbers of viable seeds. In plate 3. figure 5. is sho^^•n the
extreme expression of this vegetative method of propagation peculiar
to the western sessile. Fourteen plants, three of which do not appear
in the photograph, were produced by the parent rootstock by lateral
budding to form offsets. In the mass of rootstocks shown it was
possible to trace in many cases the original connection between the
rootstocks, since the older partially decayed portions of the rootstocks
were held in position by the heavily matted roots. The difference
in the size and thus in the age of the different offsets, the fact that
the whole mass was very compact, that the offsets were quite
symmetrically arranged about the parent rootstock in the center, the
obvious connection between the offsets and the parent and finally
the absence of seedlings in the neighborhood seems to leave no room for
doubt that this group shown in plate 3, figure 5, is the result solely of
asexual reproduction. Further than this we have noted in the
field a large number of rootstocks with four to six small, recently
produced offsets still attached to the parent and other larger plants
nearby which very apparently had been derived from this same
parent. The production of small offsets is shown on the rootstock in
plate 3, figure 6. The appearance of the plant there illustrated brings
up another matter which is of importance in this connection. Just as
T. sessile var. giganteum produces new plants asexually in consider-
able numbers, so from the "crown," or covering of the growing
point, of each rootstock more than one shoot is often sent up annually.
In plate 3. figure 6, the parent rootstock is seen to have produced at
least five vigorous flowering shoots from buds formed during the dor-
mant season at the bases of the scale-like, sheathing leaves of the crown.
T. ovatum, on the other hand, rarely sends up more than a single
shoot from a rootstock, just as it almost never produces new indi-
viduals asexually. The morpohology of the rootstock and especially
the structure of the growing apex is to be taken up in detail in a
forthcoming paper of this series.

In the garden cultures we have undertaken a study of the nature
of the teratological variations which are of such frequent occurrence
in the west coast sessile. In our cultures three different rootstocks
have produced four-leaved shoots in some one given year, the
abnormality never having appeared before or thereafter in shoots
from the same rootstocks. Fasciated shoots appear, however, in some

16 University of California Publications in Bohnni [Vol. 7

eases lo lie i-cprodueed year after year from llic same rootstock, as
does tlie peculiai- altiioniialil y which consists of the production of
small 1mi1 fully ilcvrloiicd pistils witliiii tlir ])lacentation of the legiti-
iiialc ovary. Two cases of |)istillo(ly of the j)erianth, a case in which
sepals are reduced to scales, and many cases of pctalody of sepals
and stamens and duplication of parts have been noted in the field
and the rootstoclcs bi-ouprht under observation in the garden.

SUMMARY

Following a description of the Calif ornian species of Trillium —
T. scssilf vai'. (jigantcum, T. ovatum and T. nivale — the following
points wci-c bi'ouii'ht uj) and discussed more or less in detail. They
represent llic i-esults of field and garden studies and in many cases
have been mentioned in this first pai)er of the contemplated series
dealing with the Californian species of Trillium, simply as prelim-
inary statements of important problems which are being separately
attacked in considerable detail.

1. The extent of the size differences between the eastern T. sessile
L. and its Californian variety giganteum H. & A.

2. The wide range of variation in flower color characteristic of
the Californian variety of 2\ sessile.

3. The occurrence of a pure white, a yellow-green and a heavily
l^igmented color form and a definition of their range of distribution.

4. The high degree of sterility exhibited by the Californian T.
sessile and the lack of any appreciable degree of sterility in T.
ovatum.

5. The importance of asexual reproduction in T. sessile var.
giganteum and the almost entire absence of the same in T. ovatum.

6. The yearly recurrence or lack of recurrence of teratological
variations in T. srssile var. giganteum.

Transmitted April 6, 1916.

PLATE 1

Fig. 1. Two plants of T. sessile var. giganieum with the soil exposed to
show their depth of insertion, the nature of the rootstock, amount of root
system and general vigor. The flowers are just on the point of opening.

Fig. 2. Plants from four difl'erent rootstocks of T. sessile var. gigantcum
to show the effect of reduced and unilateral illumination on height and petiole
formation. The two plants in the center were at the front of a shed while
the other two plants were three feet back from the open front.

[18]

UNIV. CALIF. PUBL. BOT, VOL. 7

[GOODSPEED-BRANDT] PLATE

Fiujnre 1

Fiarui'e 2

PLATE 2

Fig. 3. Showing the extremes of variation in size and pigmentation of
leaves and flower parts. Plants of the yellow-green color form of T. sessile
var. giganieum from Camp Taylor, Marin County.

Fig. 4. Showing variation in size and shape of flower parts in T. sessile
var. giganieum. From herbarium specimens, plants of the Berkeley Hills.

[20]

UNIV. CALIF. PUBL. BOT. VOL. 7

[GOODSPEED-BRANDT] PLATE 2

il4l

I

/' A

A

Fisfure 3

V

Fiuuu' 4

PLATE 3

Fig. 5. Showing possible extent of vegetative reproduction in T. sessile
var. giganteum. Plants from Felton, Santa Cruz County.

Fig. 6. Showing heavily pigmented flower parts, the formation of young
offsets and the number of shoots often produced from a single rootstock
crown. Plant of T. sessile var. giganteum from the Berkeley Hills.

[22]

00.

ri

<

o

>

cz

CD

o

<

O

o
o
o
o

T)

X

"^

PLATE 4

Fig. 7. Seedlings of T. sessile var. giganteum. Both the third and the last
seedling from the left shows the seed coat still attached.

[24]

UNIV. CALIF. PUBL. BOT. VOL. 7

[GOODSPEED-BRANDl] PLATE 4

CD

/

•V^

UNIVERSITY OF CALIFORNIA PUBLICATIONS

IN

BOTANY

Vol. 7, No. 2, pp. 25-38, plates 5-6 October 4, 1916

NOTES ON THE CALIFORNIAN SPECIES OF

TBILLIUM L.

II. THE NATURE AND OCCURRENCE OF UNDEVELOPED

FLOWERS

BY
THOMAS HAEPER GOODSPEED and EGBERT PERCY BRANDT

In the first paper of this series (Goodspeed and Brandt, 1916)
reference was made to the fact that when one examines in the field
an assemblage apparently of any species of Trillium, a considerable
number of the plants npon superficial examination appear to contain
no floral parts within the whorl of leaves, but upon more careful
examination are found to be provided with more or less highly
differentiated floral primordia, making up what will be spoken of
throughout as "undeveloped" flowers. Plates 5 and 6 show various
types of these undeveloped flowers. Table 1, below, consists of
measurements of the length of the various structures found in a
number of undeveloped flowers of different stages of complexity,
the structures measured being named in some cases rather arbitrarily
and in general for convenience as though corresponding in position
to the distinct floral whorls that make up the normal flower of
Trillium. From a very large number of specimens examined we are
led to make the rather sweeping statement that every plant of T.
sessile var. giganteum H. & A. and T. ovatum Pursh., the two species
peculiar to Western ^Middle California, contains floral organs which
are either complete in number and differentiation, giving a normal
flower, or are reduced in number and differentiation to give various
types of undeveloped flowers. In addition, all the apparently flower-
less plants of T. sessile L. from Illinois and of T. grandifiorum

2(i University of California Puhlications in Botany [^■o^- 7

(Michx.) Salisb. from Midii^Mii which we have examined have been
found to contain floi'al iiidinicnts. In addition to the brief report
of Smith, noted below, we have been unable to find mention made of
these facts, ami tln' ju-cscnt communication is ])r('liminary in the
sense that the results of a luunber of years" investigatioji herein
reported have (1on(> little more than point out the lines along which
the problem can be attacked. A rather large amount of data based
upon observation in the field and upon tlie results of garden cultures
makes it possible to establish certain facts with some definiteness
and it thus seems best to submit now the evidence at hand.

Miss A. S. Smith (1896) in collections nmde on April 5, 1896,
near Ithaca, New York, examined fifteen buds, probably still in the
rootstock '* crowns, " which were starting their growth in the par-
tially frozen soil. Only one of the fifteen buds was maturing a normal
flower and the remainder apparently corresponded to those figured
in i)late 6. ]\Iiss Smith cut sections in celloidin and reports that

TABLE 1

Length in mm. of the structures making up various sizes of undeveloped
flowers in TriUium sessile var. giganteum

Petals Stamens Pistil

22.0 15.0 8.0

22.0 12.0 7.0

21.0 18.0 8.0

20.0 16.0 7.0

17.0 12.0 7.0

15.0 10.0 7.0

11.0 10.0 5.0

10.0 9.0 6.0

10.0 9.0 3.0

8.0 7.0 3.0

8.0 6.0 3.0

7.0 6.0 1.0

3.0 2.0 1.0

2.0 1.0 0.3

1.6 0.8 0.3

1.2 " 1.2 0.6

0.6

0.4

0.4

0.1
0.1

0.1

Number

Sepals

1

24.0

2

23.0

3

22,0

4

22.0

5

18.0

6

16.0

7

12.0

8

11.0

9

10.0

10

9.0

11

8.0

12

7.0

13

3.0

14

2.0

15

1.7

Ifi

1.7

37

0.8

18

0.6

19

0.5

20

0.4

21

0.3

22

0.2

23

0.2

24

0.1

25

0.1

1916] Goodspeed-Brandt : Notes on Trillium 27

structures representing sepals were often normally matured but
within them a ''confused mass of cells" occurred. A much larger
collection was made in the same region on April 10 of the same
year and sixty plants were found to contain "abortive" flower buds,
concerning which she says: "Usually the rudiments of a perianth
could be distingiiished either as a white speck or as very evident
floral leaves sometimes 2 to 3 mm. in length, but withered and
abortive." Miss Smith suggests that "truly sterile" plants might
have been present, but she found none without at least rudimentary
floral structures.

The drawings shown in plates 5 and 6 represent median longi-
tudinal sections through the center of the stem apex which in the
apparently flowerless shoots is seen as a cuplike depression, on the
rim of which the almost completely sessile leaves are borne. The
drawings were made from material sectioned in paraffin ; while
undeveloped flowers of the size shown in plate 6 might be distin-
guished by the naked eye as minute, white, withered structures in
the center of the cuplike depression, those shown in plate 5 would
escape notice without magnification. The complete range of variation
in size and differentiation of parts given in table 1 is not paralleled
in the drawings, wherein only smaller and more rudimentary floral
structures are represented. It is important to note in table 1 that
a complete series can be obtained, from undeveloped flowers consist-
ing of such minute primordia as these under numbers 17 to 25 to
those which are really perfectly matured but functionless buds illus-
trated under numbers 1 to 3. As mentioned above, we have in this
table used the terms petals, sepals, etc., simply for convenience of
description in giving the measurements of the structures found.
Thus in the case of the smaller group, from numbers 13 to 25, the
separation into circles of such structures as are shown in the drawings
in plate 5 is a rather arbitrary matter. It must be said, however,
that, in dissecting under magnification even these minute organs, one
is impressed with the fact that the structures present are arranged in
some more or less definite order and inserted at various points upon
the reduced receptacle. Certainly in the case of the group of
intermediate size, numbers 7 to 12, there is cause for assigning groups
of structures to given circles, while in the largest group, numbers
1 to 6. the characteristic floral whorls are perfectly represented. It
might also be noted that measurements are at hand which together
would form a still more continuous series in which the size of the

28 University of ('alifoniid I'uhJications i)t Botany [^^o^- '''

structiiri's would vary ().") mm. or less l)('t\v('en different individuals.
The series jjiveu in tabU' 1 is suffieiently eomidete lo bring out the
points wliieli seem of most signifieance.

If we lurn now to a description of the ai)pearance of these various
sizes of undevelopeil flowers. Hie more highly differentiated form —
numbiTs 1 lo () — might lirsl bi- mentioned. As stated above, these
larger undeveloped flowei-s ai'e really small, fully formed flower buds.
They are (piite conspicuous, standing in the center of the whorl of
leaves, with while, transparent, parehmeiit-like .sepals and petals.
Such "buds" never open and I'apidly dry down as the leaves are
fully expanded. The stamens are usually perfect in appearance with
anthers fully matured, 1 hough they are very slender and perfectly
white except along the pollen sacs, which are sometimes a very light
j^ellow in color. Pollen is often present in such cases with every grain
shriveled and functionless, while in other cases sporogenons tissue
apparently is suppressed entirely in the development of the anthers.
The ])istil in these lai-gest undeveloped flowers is always functionless
and often entirely wanting. Where if does occur the pistil is very
nuR'li reduced, with diminutive stylar tissue and with no ovules or
structures suggesting them in appearance. On T. ovatum. a
pedunculate species, such large undeveloped flowers at times are to
be seen and in these cases a peduncle is present proportional in size
to that of the bud. At times, however, this proportion is not main-
tained, and the small, white bud appears to be sessile, so short is
the peduncle upon which if is borne. There is a more or less abrupt
transition from the early withering, functionless bud that never
opens, although its parts are rather fully and characteristically
matured, to the small undeveloped flowers in which all the parts are
similar in appearance though differing greatly in size so far as the
different circles of perianth segments are concerned. As stated
above, the pistil even in the largest undeveloped flowers is often
suppressed, the sepals and petals are of the same color, texture and
size, and the remainder of the functionless bud consists of narrow,
delicate anther-bearing structures. If is in the differentiation of these
stamens of more or less characteristic appearance and sfi-uctui-e that
file rafhei- abinipt fi'ansition occurs. Thus all the smaller undeveloped
flowers consist of nothiug more than minute, white, di-y perianth
segments which are, in all but those of the smallest size, arranged
rather definitely in separate whorls. As shown in ]ilate 5, figure 2,
the smallest uTidevelojied flowers in some cases show a union of the

1916] Goodspeed-Braiidt : Xotcs o)i Tyilliiim 29

individual segments to form an enclosed structure which suggests a
carpel.

TABLE 2

Production of normal flowering shoots (marked x) and of shoots with undevel-
ojied flowers (marked -) during six years in garden cultures

1915 1916 Place of collection

- — Point Reyes

- - Point Eeyes

- - Point Eeyes

- X Point Reyes

- - Berkeley Hills

- - Berkeley Hills
Berkeley Hills

-* X Berkeley Hills

- - Berkeley Hills

- - Berkeley Hills

- X Berkeley Hills

- - Point Reyes
Berkeley Hills

- - Berkeley Hills

- - Berkeley Hills

- - Monterey Peninsula

- - Monterey Peninsula
- Napa Range, 2000 ft.

- - Crystal Springs, San Mateo Co.

- - Monterey Peninsula

- - Monterey Peninsula
88 _____ Berkeley Hills

93 X X X - - Gualala, Mendocino Co.

*irndeveloped flowers of the largest size and most highly difl'erentiated —
i.e., buds of almost normal size which never open and wither early.

In connection with an investigation of the nature of thes(3
undeveloped flowers we have been concerned with a study of their
occurrence both in the field and in garden cultures. As a result we
are prepared to make the following general statement : A rootstock
bearing a normally flowering shoot in a given year may the following
year produce a shoot bearing an undeveloped flower of larger or
smaller size, while the reverse sequence of events rarely takes place.
This point is emphasized in table 2, in which a cross indicates the
production by a rootstock of a shoot or shoots with normal flowers
and a dash indicates the presence of an undeveloped flower. A large
percentage of the more than 150 rootstocks grown in our garden cul-
tures bore normally flowering shoots when collected, or were known

Rootstock

number

1911

1912

1913

191

1

X

X

-"

-

2
3

X

X

—

—

4
5

X

—

—

—

7

X

X

-

-

9

?

X

-

-

11

_*

_*

_*

-'

29

X

X

-

-

31

-

-

-

-

39

X

X

-

-

4.5

X

X

X

-

48

X

X

-

-

55

-

-

-

-

57

-

-

-

69

X

-

-

70

X

-

-

71

X

X

X

1 1

X

-

-

81

X

X

X

83

X

X

X

30 University of California ruhUcations in Botany [Vol. 7

to have borne sncli shoots duriug the season previous to coHcetion.
Diii'iiiir the present season (1916) only six rootstoeks produced such
normally flowering shoots ami 1lic i riiiaiiidri- pi'odueed shoots with
undeveloped flowers. Table 2 gives the history of twenty-three of the
rootstoeks in the garden cultures and imlndes three of the six normally
flowering plants of the 1016 season. The other three flowering plants
were fmin rootstoeks wlii--li have been under observation during two
seasons only.

In table 2 characteristic eases were chosen, and a wide range of
original habitat of the rootstoeks was selected to emphasize the fact
that the production of undeveloped flowers is peculiar to all root-
stocks, irrespective of the soil or climatic conditions under which
they were originally growing. Further, the instances cited include
nearly all of the rootstoeks which produced normally flowering shoots
for more than a single season following collection. Rootstoeks 3, 31,
55, 57 and 88 exhibit the extreme condition in that when collected
they bore shoots with undeveloped flowers and have continued to
produce only such shoots thereafter. Rootstoeks 5, 69, 70 and 77
were collected with normally flowering shoots and the following year
and thereafter produced shoots with undeveloped flowers only.
Rootstoeks 1. 2. 7. 29 and 48 and rootstoeks 45. 71, 81. 83 and
93 continued to produce normally flowering plants for respec-
tively two and three years following collection and thereafter
have produced plants with undeveloped flowers only. There is,
then, no doubt that the change from perfectly normal flower
to undeveloped flower can take place in a single season and that
when this change has once taken place it may, in the majority of
cases, be expected to reappear in succeeding years more or less
indefinitely. These points have many times been confirmed by field
observation during the past six years.

The histories of rootstoeks 4, 11 and 39 are obviously of importance
in this connection. Here we have cases in which rootstoeks have
produced only plants with undeveloped flowers for three successive
seasons or more and then in a single season have produced plants
with large, perfectly normal flowers. In the case of rootstock 4 the
flower produced was the first one ever seen from this rootstock,
which was originally collected on Point Reyes peninsula, and it is
interesting to note that this flower, produced after six years of growth
under different conditions of soil and climate, corresponds exactly in
color and other characters (cf. Goodspeed and Brandt, 1916) to

1916] Goodspeed-Brcuidt : Notes on Trillium 31

those found during the same season in their native habitat. The
evidence furnished by rootstock 11 and a number of others seemed
for a time to indicate that there was each year an increase in the
size and differentiation of the undeveloped flowers, and it was thought
that this increase in size and differentiation would finally result in
the production of a normally flowering shoot. As indicated, however,
rootstock 11 produced a highly rudimentary undeveloped flower in
1915 after producing undeveloped flowers of larger size for a number
of years. In general it seems plain that there is no gradual transition
from undeveloped to normal flowers, but that a rootstock may produce
a diminutive undeveloped flower in one season and a perfectly normal
flower in the following season, just as the same rootstock might
conceivably in a third season produce again a diminutive, undeveloped
flower.

A further point must be noted in this connection. An offset
produced from a parent rootstock never forms anything but an
undeveloped flower in the first season in which it bears a shoot, and
it is doubtful whether it produces a normally flowering shoot for a
considerable number of years thereafter. A seedling rootstock has
apparently the same history in this regard as the young offset. We
are thus concerned with two rather different situations in connection
with the occurrence of undeveloped flowers in Trillium species:
first, the number of years of growth necessary in the case of offsets
and seedlings for the production of a normal flower, and second,
the number of years that must intervene, after a normally flowering
rootstock has reverted to the condition of producing undeveloped
flowers before a normal flower can again be matured. With reference
to the first point, we have seen small offset rootstocks which when
collected gave evidence of at least five years previous growth, pro-
duce only undeveloped flowers for six years in the garden cultures.
As might perhaps be expected, it has been determined that seedling
rootstocks in the field remain even longer in the undeveloped flower
condition than do offsets from old, vigorous rootstocks. Concerning
the second point above, we have, as can be seen in table 2, only
one instance in the garden cultures — i.e., rootstock 39 — in which a
rootstock produced normal flowers, reverted to production of unde-
veloped flowers, and again after three years produced a normally
flowering shoot. In the great majority of cases normally flowering
rootstocks introduced into the garden cultures five or six years ago
have thereafter borne only shoots with undeveloped flowers.

32 Ciiircrsitij of California I'nhUcations in Botany I^'Ol. 7

The problem of llic occunvncc of undeveloped flowers and its
cause is furllici- coinpliciilrd by the t'jict that in Trillium sessile var.
giga)ihinit a vi^'oi-ous roolstock may i)!'(»ilii<-c t'oiii- or five shoots in
a siufi'le season IVom its ^r()\viii<i- apex and all of these may bear
noi-mal flo\vei-s. all iiia.\- bear undeveloped fiowei's, or one or two
shoots may bcai- normal flowers and the I'emaintU-r bear undeveloped
flowers, this laltei- condition being by far the most eonnnon. The
moriiholojry of the growing apex of the rootstoek, whieh is covered
by heavy, i)roteetive, sheathing scales, will be dealt with in the
following ]iaper of this series and only certain points need to be
mentioned here. There appears to be a critical stage during the active
period below ground in the develoi)ment of the buds at the bases of
the sheaths of the I'ootstock "crown" such that if a given bud has
not attained a fair degree of development by tlie first of July the
growth of that bud is arrested and it may shortly thereafter die and
shrivel up. It fui'ther appears that from a bud whieh has reached
a somewhat more advanced but not tlie maximum development at
the critical period a shoot with an undeveloped flower will appear
the following spring. In other words there are three possibilities
Avith reference to development of the buds following the critical
period. First, the development of a given bud may be insufficient
at the critical period and thus its growth ceases and it dies shortly
thereafter. Second, a given bud may have reached the maximum
degree of development possible at the critical period and thus suc-
cessfully passes through this period and produces a normally flowering
shoot the following spring. Third, a given bud may have reached
such a stage of developmeiit at the critical period that fui-fher
vegetative development is possible but the development of the floral
primoi'dia is not sufficiently advanced and growth of these structures
is arrested or goes on very slowly to ])roduce an undeveloped flower
the following season. We nuiy think of an intermediate stage between
the last two, in which development is sufficient to allow of the
formation of almost normal reproductive structures, the actual
reprodui'tive tissue failing of normal development.

The occurrence of a critical period is readily demonstrable. It
appears to be connected with the shifting of a j)ortinu of the meta-
l)()lie activity of the growing region, jii-eviously de\dted to the
fornuition of vegetative structures and floral primordia, to the
tlevelopmenf of reproductive tissue whieh takes place vigorously from
July on to the jx-riod above ground. The oeeuii-ence of undeveloped

1916] Goodspeed-Brandi: Notes on Trillium 33

flowers is thus tentatively assigned to a lack of sufficient development
of flower parts at the period when the products of metabolic activity
stored in the rootstock are for a time shifted almost wholly to the
reproductive tissues for their rapid development. A seedling root-
stock or young offset is thus to be thought of as capable of maintaining
nothing more than a vegetative existence above ground, a balance of
available food properly to mature floral organs being accumulated
in small amounts each year until after a variable but usually con-
siderable number of years a normal in place of an undeveloped flower
is produced. This explanation apparently covers such a case as that
of rootstock 39 in table 2, since a rootstock which has successfully
produced shoots should, other things being equal, be successful in
passing from the condition of undeveloped to normal flower more
rapidly than a seedling rootstock or offset which has never succeeded
in producing a normal flower. The abrupt transition from normal to
undeveloped flower production may be taken to be due similarly to
a disturbance of metabolic relations, the most obvious of which is
the taking up of a normally flowering rootstock during the height
of its active period above ground. Such treatment almost invariably
results in the production by it of an undeveloped flower in the follow-
ing season.

In general, of course, the production of undeveloped flowers may
be taken as an extreme instance of the sterility commonly seen in
normally flowering plants of T. sessile var. giganteum, which produce
very little viable seed and in which the pollen may be highly defective
or the development of the pistil entirely suppressed. That the sterility
found in normally flowering plants and expressed in its extreme form
in the production of undeveloped flowers may be assigned to condi-
tions of nutrition and available food-supply is at least suggested by
a comparison of the reproductive methods of two distinct Californian
species — T. sessile var. giganteum and T. ovatum. In the former,
as we have seen, undeveloped flower production and sterility in
normal flowers is common and here vegetative reproduction is im-
portant and very vigorously carried on (cf. Goodspeed and Brandt,
1916). In T. ovatum, on the other hand, viable seed is formed in
a majority of the normal flowers ; and undeveloped flowers, while
common, are not produced in such amount, and finally we have hardly
succeeded in finding a single instance of vegetative reproduction by
offsets.

A variety of experiments are contemplated in an attempt to gain

3-1 University of ('(iJifnnila T'uJilicafions In Bofcnnj L^^o'- 7

positive evidciR'c as to \hv cause of uiidcvelopc'd flowci- i)rocluctioii
and the conditions both upon which it is dependent i'or continued
appearance in the successive slioots from a rootstock and those which
are responsible for the change back to the condition of normal flower.

Transmitted April 6, 1916.

LITERATURE CITED

GOODSPEED, T. II. and liRAXDT, K. P.

1916. Notes on the Califoruian species of Trillium L. I. A report of the
•general results of field and garden studies, 1911-1916. Univ. Calif.
Publ. Jiotaiiy, vol. 7, no. 1.

Smith, Arma S.

1896. Abortive flower buds of Trillium. Bot. Gaz., vol. 22, ]>. 402.

PLATE 5

Figures 1 to 6
Longitudinal fsections through the apices of shoots of Trillium sessile var.
giganteum, showing the uature of undevelojied flowers of different degrees of
complexity.

[36]

UNIV. CALIF. PUBl.. BOT, VOL, 7

[GOODSPEED-BRANDT] PLATE 5

PLATE 6

Fijfures 7 and 8
Longitudinal sections through the apices of shoots of Trillium sessile var.
giganteum, showing rather highly differentiated types of undeveloped flowers.

[38]

UNIV. CALIF. PUBL, BOT. VOL, 7

[GOODSPEED-BRANDT] PLATE 6

UNIVERSITY OF CALIFORNIA PUBLICATIONS

IN

BOTANY

Vol. 7, No. 3, pp. 39-68, plates 7-10 December 9, 1916

NOTES ON THE CALIFORNIAN SPECIES OF

TBILLIUM L.

III. SEASONAL CHANGES IN TRILLIUM SPECIES WITH ^.t'***"'^

SPECIAL REFERENCE TO THE REPRODUCTIVE •*«'A/v#<

TISSUES

BY

EGBERT PERCY BRANDT

CONTENTS

PAGE

I. Introduction 39

II. The subterranean organs of Trillium species 42

III. The active period — February to July 46

1. Above ground 46

a. Trillium sessile var. giganteum 46

b. T. ovatum 49

2. Below ground 49

a. T. sessile var. giganteum ". 49

6. T. ovatum 51

IV. The dormant period — August to January 52

a. T. sessile var. giganteum 52

6. T. ovatum 57

V. Summary 57

VI. Literature cited 60

VII. Explanation of plates 62

I. Introduction

The variety of problems presented has made a study of the species
of Trillium native in the region immediately surrounding San Fran-
cisco Bay of unusual interest and importance. The first two of the

40 University of California Puhlications in Botany [Vol. 7

studies in this series (Gocxlspeed and Brandt, 1916, a and h) were
concerned witli a general description of tlie Californian species of
Trillium, with <i hi-icf inciilioii of cci-liiin points in their lifc-liistory
and a discussion of the origin, nature and significance of the widely
occurring undeveloped type of flower. The present paper, the third
in the series, is particularly concerned with the condition in which
these western species pass their resting period, a period apparently
determined by available moisture. The matter seemed of special value
since information is furnislied by the literature concerning the con-
dition of eastern species during a corresponding dormant period in
which both moisture and temperature relations ai"e seemingly con-
cerned.

Chamberlain (1898) reports that spring-flowering perennial plants
either pass the wdnter in the spore mother-cell stage or mature their
spores before the winter season, but with the data at hand was unable
to decide which was the more general condition (cf. PfeifYer, 1912).
Among those plants which pass the winter in the spore mother-cell
stage he includes Trillium species, on the basis of the report of Miss
A. M. Smith (1896), who found that the pollen mother-cells were "in
the early stages of division" in a specimen dug from beneath the
frozen soil on April 5. She found the "later stages of division" and
pollen fully matured on April 15, when numerous plants were ap-
pearing above ground (cf. Gates, 1908, p. 46). Atkinson (1899),
working on T. grandiflorum (Michx.) Salisb. in the vicinity of Ithaca,
New York, where Smith had previously worked, reports that in all
plants he examined the pollen mother-cells were formed by the be-
ginning of winter, and that in open woods on rather high ground,
protected from north and west winds, pollen was often mature in
September. In cold ravines, on the other hand, plants entered into
the resting period in the pollen mother-cell stage. Late in February
and early in ]\Iarch, when the w^eather is relatively warm, formation
of pollen begins. The report of Ernst (1902) dealing with the con-
dition of Paris quadrifolia L. and Trillium grandiflorum tends to con-
firm that of Smith, noted above. He states that micro- and megaspore
mother-cells become plainly distinguishable toward the beginning of
the growing season, late in February or early in IMarch. Chamberlain
(1898) also reports four potential magaspores fully formed in April
on a plant of T. recurvatum Beck, not more than 5 cm. high. The
above reports show that Trillium species native in the Eastern United
States and cultivated in gardens in Northern Germany usually mature

1916] Brandt: Notes on Trillium 41

their spores in the early part of their growing season before or shortly
after the young shoots appear above ground. Miss Alden (1912) gives
a review of the rather meager literature on this general subject in
connection with her pertinent study of stages in the seasonal develop-
ment of Uvidaria sessilifolia. Howard (1915) has made a series of
interesting experiments in an effort to break the summer rest period
of bulbous plants.

With the object, then, of finding out how nearly the behavior of
our forms of Trillium in the resting season agreed with that of the
eastern forms, collections and observations were begun in the autumn
of 1913. Collecting was begun while the plants were in the pollen
mother-cell stage, late in September, and was discontinued in April,
when most of the plants were going out of flower. Collecting was
resumed in August, 1914, and continued until November 10, 1915.

i\Iost of the material was collected in the hills near Berkeley, on
the eastern side of San Francisco Bay, but from the latter part of
May to the early part of August, 1915, T. sessile var. giganteum H.

6 A. was collected at Point Arena, Mendocino County. From time to
time T. ovatum Pursh was collected for purposes of comparison, at
Mill Valley, Marin County, and at Gualala, Mendocino County.

Up to the autumn of 1915 all specimens were trimmed and killed
in the field to avoid wilting. In these last collections, made for the
express purpose of securing a large supply of material showing chro-
mosome reduction in the pollen mother-cells, the material was brought
into the laboratory and examined before killing, suitable precautions
having been taken to keep it fresh. The following killing solutions
were used: Bouin's solution, Zenker's mixture, Flemming's fluid
(strong), and Farmer's absolute alcohol-acetic acid solution. The last
two gave the best results.

All material was imbedded in paraffin. Anthers were sectioned

7 to lOfx thick, ovaries 7 to 25fx, according to the stage of development
of the ovule, and young stems just forming lateral organs were sec-
tioned 10 or 12/x thick. The stains used were aqueous safranin and
Delafield 's haematoxylon in combination or alone ; safranin and ' ' light
green"; the safranin, gentian-violet, orange G. combination; and
Haidenhain's iron alum haematoxylon. The last stain was particu-
larly useful in studies of chromosomes, while the most generally suc-
cessful was the safranin ' ' light green ' ' combination.

42 Universifji of Calif oruut Piihlicdfidiis in Botany [Vol.7

II. The Subterranean Organs of Trillium Species

The stoi'agc sti'uctui'c of TviUinm consists of a stout subterranean
stem provided willi a iclatively large terminal ImuI, or "crown," and
perennial roots. In shape the rootstock of T. sessile var. (liganteum
is usually nearly cylindrical, but tapei-s slightly to one or both ends.
It varies in length from 3 to 8 cm. and in diameter from 2 to 5 cm. or
more. The rootstock of T. ovatum is more slender, varying in length
from 2 to 4 cm. and in diameter from 5 to 15 mm. (cf. Rimbach, 1902).
In both species the rootstock is either erect or horizontal. When the
rootstock is buried deeply it usually stands erect, as Rimbach {loc. cit.)
states for T. ovnlioii. Often, as noted by Rimbach, the rootstock of
T. ovatum may gi'ow sharply downward at the crown end, and thus
the erect terminal hud makes an acute angle with it.

Since the rootstock in general grows only enough to make room
on its surface for the bases of bud sheaths and axillary structures, the
annual growth is slow, probably never more than two centimeters.
Rimbach states that the rootstock of T. ovatum does not grow more
than 1 mm. per year, but I have seen specimens that showed indication
of an annual growth of 2 mm. or more. The scars on the surface left
by the decay of the bases of aerial shoots furnish a good indication
of the amount of annual growth, but when a plant sends up several
shoots per year one must depend more upon the slight annual con-
strictions of the rootstock, which are often readily discernible. As
growth goes on at the "crown" end of the rootstock the base rots
away, generally leaving a smooth scar ; the age of tlie living portion
of the rootstock does not thus necessarily indicate the actual age of
the plant. Rimbach further states that the rootstock of T. ovatum
sometimes contains the living product of twenty to forty years" growth,
but that is far too great an estimate. Carl Purdy, the well-known
horticultural authority on California Liliaceac, informs me that in his
experience Trillium rootstocks seldom persist entire for more than
fifteen years even in well drained gravelly soil, and in ordinary garden
soil the rootstock of T. sessile var. giganteuni contains the living pro-
duct of less than five years' growth. It would seem that Rimbach,
who collected his material on the sloi)es of JMount Tamalpais, I\Iarin
County, must have based his conclusions as to the persistence of root-
stocks upon the rate at which young rootstocks grow during their first
four or five years. The evidence furnished by my own collections
made in the same locality and elsewhere confirms the statements of

1916] Brandt: Notes on Trillium 43

Purely mentioned above. My determination of the age of a given
rootstock is made by counting the sears left by the decayed bases of
aerial shoots and noting the slight annual constructions which usually
can be seen on the surface of the rootstock.

Roots grow out from the lower side of a horizontal rootstock, but
when the rootstock stands erect the roots grow out equallj' from all
sides (cf. Rimbach, loc. cit.). Rinibach states that T. ovatuni produces
two or three roots per year, and that they live eight or ten years, so
that the plant may have as many as twenty roots, but he probably
overestimates the age of the roots just as he does that of the rootstock.
T. sessile var. giganteum sends out three to five or more roots from
near the base of the terminal bud or "crown" at about the time the
young shoots appear above ground. These attain a length of 15 to
20 cm. or more the first year. During the second year they increase
their length still more and send out fairly numerous branches of the
first order. They apparently live three or four years, or even longer,
but probably not so long as the portion of the rootstock which bears
them. When the roots have attained a considerable length, they begin
to contract, the surface of the roots becoming noticeably wrinkled as
they do so. This contraction draws the rootstock down into the ground.
Since only the young roots contract, the pull is exerted only at the
growing end of the rootstock. If the ground is hard this will cause
the young rootstock, upright in the young seedling stage, to become
inclined from the perpendicular. As the rootstock becomes inclined it
develops a tendency to send out roots only from the lower side, thereby
eventually becoming horizontal or inverted. Evidence of this gradual
change of position of the rootstock is furnished by the slight arching
so common in that of T. ovatum. On the other hand, the rootstock
tends of itself to assume an upright position, so that in loose soil up-
right rootstocks may be found. When the rootstock is erect, roots,
as noted above, grow out equally from all sides, and thus enable it to
maintain an upright position.

As a result of this tendency of the rootstock to grow erect and to
send out roots more nearly in all directions the more nearly erect it
becomes, we find most of the rootstocks of T. sessile var. giganteum
and the stronger ones of T. ovatum growing erect or curving upwards.
Rimbach states that the roots of T. ovatum, which attain a total length
of 20 to 30 cm., reduce their length about 1 cm., becoming wrinkled
for a distance of 3 or •! cm. in doing so. The roots of T. sessile var.
giganteum show this same surface wrinkling. Photographs of root-

44

Universitij of Calif oruut Publications iti Botainj [Voi,.

stocks are included in a previous report (cf. Goodspeed and lirandt,
]i)16, fl. pis. 1 and 5).

Tile tcniiiiial hutl, or "ci'own,"' suggests a large aci-ial winln' hud
as much as it suggests tlie bulb of a lily. It is more or less elongated
and conical, w itli a diameter somewhat less than that of the rootstock.
Tlic sheathing scales, each of which completely enwraps all the parts
inserted above it, arc Ihin and tougli, adapted to i)rotective purposes

i h. %, h, Xj J, h, h', h i; A '-6. J.

Fig. 1

rather tiian serving as stareh-storing tissue. Figure 1 above illns-
trates the structure of a strong "crown" early in September, .s^, is
the outermost sheathing scale, not quite reaching to the top of the
crown, brown and dry ; bearing in its axil a spongy, scale-like body, .r^,
wliich in turn has an aborted stem-bud. h^. in a socket in its l)ase. s._,
is a large living sheath which envelops the crown, folding completely
over it bnt not fusing along its nuirgins. The bud h. is the largest in
this crown, it has grown up out of the "axillary sheath or scale-like
body" and is pressing it back against the next protective sheatli. It

1910] Brandt: Notes on Trillium 45

will be noticed that the leaves greatly surpass the sepals and are folded
several layers thick around the flower-bud and the anthers at this
time practically fill the space in the center of it. Vascular bundles
are plainly distinguishable in the young stem. &., is the second normal
bud, which is shown entire. As in ho, the young foliage leaves com-
pletely enwrap the flower bud. The size and shape of the "axillary
sheath" is illustrated by x^. Sheath s^, as will be noted, is consider-
ably shorter than bud 63, so bud h^ would naturally be much smaller
than h.,. Only the position that bud h^ would occupy can be indicated.
Bud h\ will be noted under a very short, young sheath. This bud if
shriveled up would be a trifle smaller than h^, and perhaps a little less
advanced. It is not a bud for next season, but for the season following,
when it will fail of development in the same way as &i. b'2 is barely
distinguishable, and at a the apex of the rootstock is covered by a
few young sheaths which are merely arched and have no buds, only
"axillary sheaths," in their axils.

It is important to note the abortion of h^ and the possible abortion
of h\. Apparently all the buds from fe. and inward were laid down
in the preceding active season, but h\ and 6'._, were arrested in devel-
opment so that they could not become shoots during the next season.
There is apparently a critical period during which further develop-
ment in the younger buds is temporarily suspended. Examination
of a crown late in May seemed to show that there is a selective elimi-
nation relative to the amount of differentiation the various buds ex-
hibit. Thus, as a general rule, the bud which fails during this critical
period to meet the requirements of sufficient growth dies and shrivels
up later on. In very strong plants of T. sessile var. giganteum, those
which send up four or more shoots in a season, the largest arrested
bud develops into a shoot, usually smaller than the one next within.
In very large plants this outer shoot may show increase in number
of parts or may even become fasciated. A report upon numerous
teratological forms peculiar to T. sessile var. giganteum is in progress
of preparation.

The "crown" of T. ovatum is very similar in structure to that of
T. sessile var. giganteum. Usually not over two buds are formed each
year instead of two or more, as in the latter species. Tlie first of these
fails to develop. When the plant is given an unusual amount of light
both buds develop into strong, normal shoots. Because of its habit of
forming two buds a year, one may often find T. ovatum in partially
cleared forest areas regularly sending up two shoots from one root-

46 I' nii'( r.silij of Calif orn'ui l*uhUcations in Botunij [Vol. 7

stock. Plants with three or four slioots are always in ihc minority,
whereas there seems to hv no limit to the inmihei- of shoots that a
single rootstock of T. sessile var. giganteum may send up, eight having
been seen in one specimen.

In addition to tlie ])ai'ts eninneraled al)Ove, the i-ootstock of T.
sessile var. giganteum very frequently has one oi- more young root-
stocks growing out from it as lateral branches or ''offsets." These
increase their diameter from year to year, build up their own root
systems, and by decay of the older portions of the rootstock soon be-
come independent plants. In this way a single plant often in time
produces a clumj) of a dozen or more. On the other hand, T. ovaium
very rarely produces an otTset (cf. Kerner and Oliver, 1895, pp. 452-
454). The general significance of vegetative reproduction in Trillium
species has been fully discussed in a previous report (Goodspeed and
Brandt, 1916, o).

Foerste (1891) states that the Eastern or type form of T. sessile
L. is characterized by a horizontal rootstock, and refers to the orien-
tation of the rootstock by the contraction of the roots. He also states
that in early wdnter there are buds present in the rootstock crow'n for
tw^o active seasons in advance.

III. The Active Period, February to July

1. ABOVE GROUND
a. Trillium sessile var. giganteum

In a previous report (1916, a) we have fully outlined the climatic
conditions under which plants in western middle California pass their
annual life-cycles. A number of the points there emphasized are of
importance in the following discussion and will be briefly referred to.

Trillium sessile var. giganteum appears above ground in the latter
part of January and in February. This species has withered and in
almost all cases disappeared by the end of July. For convenience 1
have chosen to regard this period — February to July — which includes
the life of the plant above ground as equivalent to th(^ ''active" period.
It is of course obvious that the active period of growth and flowering
occupies a considerably shorter interval. How-ever, the period chosen
covers the interval fi-om the appearance of the green shoot to the
shedding of seed, and therefore marks the life-cycle above ground.

When the young shoot first appears above ground the leaves are
tightly rolled into a cone about lln^ flower-bud. In some plants the

1916] Brandt: Notes on Trillium 47

leaves are imbricated in the bud, in others convohite, the species ap-
parently not adhering to one arrangement. The characteristic colors
of the various aerial parts are variable in the time of their appearance.
As the shoot emerges, the leaves and sepals rapidly change from deep
yellow to green, while the petals attain their full coloration only -when
entirely exposed to the light and free from the enveloping sepals and
leaves. The stamens are heavily pigmented, showing almost their full
coloration a short time before the flower unfolds.

From the first appearance of the tip of the shoot above ground
to the complete emerging of the tightly rolled cone of leaxes approxi-
mately a week intervenes. Thereafter growth is rapid until the
maximum length of stem is reached. Measurements made on devel-
oping plants in the University of California Botanical Garden, and
field notes, provide the basis for this statement. Almost with the
start of this rapid growth to increase the length of the stem a loosening
of the cone of leaves is evident, and as growth proceeds the unfolding
of the leaves is sufficiently rapid so that they are completely flattened
shortly after the maximum height is attained. This gradual unfolding
and flattening of the leaves is accompanied by a sufficient increase in
size so that they are nearly mature, so far as growth and expansion
are concerned, when the stem has attained its full height. In general
it might be said that this period of rapid growth occupies, in average
cases, a little more than two weeks.

The significant stages in the development of the flower take place
subsequent to the complete unfolding of the leaves. The sepals are
practically full size at this time and are pushed apart by the rapid
growth of the petals, which are hardly in sight within the sepals when
the leaves first unfold. The petals are fully mature and the flower
unfolds approximately three weeks after the shoot first appears.
Owing to the method of dehiscence of the pollen sacs, anthesis may
follow immediately or after an interval of more than a week, according
to atmospheric conditions. Dehiscence is accomplished by the mere
straightening of the walls of the locules, caused by the epidermis
contracting more than the endothecium. The locules may remain
distinct until anthesis, in which case a zone of small-celled tissue is
developed between the locules (see pi. 10, fig. 2), or they may become
fused into one by the disintegration of the parenchyma between and
back of them, but this is not essential to dehiscence. It can be shown
that anthers from unopened flower-buds may dehisce within an hour
in the dry air of the laboratory.

48

Universil ij of Califontld I'uhlications in Botany [Vol.

Whi'ii tlu' flower first opens the stout styles are erect and closely
pressed together. Gradually they roll back at the tips and also spread
apart at otlier points below, l)ut never spread apart very widely. The
nuirkedly i)ai)illose surface of tlie stigma is veiy undulate. This
probably aitls in pollination in two ways, by increasing the surface
area and by providing hollows and pockets in wliidi ])ollen may lodge.

Tlie Muthei's sIkuv more (ir less tendency to curve ovei- tlie stigma,
sometimes foi'ining a close cone over it. rdth()U<rh in other eases
they are somewhat divergent. As they open introrsely self-pollination
is in most cases inevitable. The flowei- is larcly visiled by insects
(cf. Carter, 1892).

Fiu'.

In from three to six weeks the petals and stamens wither, but do
not fall. Sepals and foliage leaves persist until the fruit ripens, or
the leaves may perish earlier. As has already been stated, the fruit
ripens late in July. It varies in sliape from elongate-oval to slightly
spheroid, 15 to 35 mm. long by 12 to 34 mm. thick ; is terete at the base,
slightly six-angled above, with the persistent bases of the styles form-
ing three short beaks. The color varies from pale green without a
puiplisli tinge to red or purple. The fruit dehisces by breaking away
at the base and teai'ing up one side (fig. 2). The seeds are usually
luimerous, sometimes numbering nearly 200, are 4 iiiiii. loni;: by 2 mm.
thick, oval, light brown: witli fleshy white aril larger than tiie seed,
developed from the funiculus (fig. 2). This ai'il is greedily eaten by
ants (Robertson, 1897, p. 288). So far as T am aware, this is the fii-.st
complete description of the fruit of the western ally of T. sessile L.

391(5] Brandt: Notes on Trillium 49

h. Trillium ovatum

Practically all the points noted above in the description of the
active period above ground peculiar to Trillium sessile var. giganteum
hold in modified form for T. ovatum. It might be recalled that T.
ovatum is a pedunculate species in contrast to T. sessile var. gigan-
teum, which is sessile, and that previous to anthesis the flower is white.
Growth is much more rapid, the shoot attaining full height in a week.
The flower opens before the leaves are fully spread and the petals
have faded and fallen by the time the plant has been out of the ground
a month. The species is distinctly protandrous and is doubtless ento-
mophilous, as Robertson (1896) states is the case in T. grandiflorum
and T. erecium. Little need be said about the fruit, which usually
ripens in May, except that dehiscence is septicidal, beginning at the
apex and extending to the base. The seeds, which are smaller and
darker brown than those of T. sessile var. giganteum, sometimes num-
ber more than 300. They also are myrmecophilous, as Robertson
(1897) reports for T. recurvatum^

2. BELOW GROUND
a. Trillium sessile var. giganteum

With the beginning of growth in February the bud-sheaths lengthen
sufficiently to appear a few centimeters above ground and protect the
young leaves as they push up through the soil. They are separated
by the rapid gi'owth of the young shoots and gradually disappear.
The axillary "sheaths," or spongy bodies at the bases of the main
sheaths, lengthen but little, but appear to remain alive quite late in
the flowering season. At this same time the apex of the rootstock
begins to grow and to lay down new stem-buds while the young buds
whose growth was suspended resume development, with the exception
of the first bud, which now shrivels up, as previously stated.

The young stem is at first a short cylinder of undifferentiated
parenchyma. Soon the lateral structures make their appearance in
acropetal succession. The young axis is produced beyond the point
of insertion of foliage primordia so that the young sepals slightly
overtop them at first (pi. 7, figs. 2 and 5). During early development
the floral organs apparently grow more rapidly than the foliage leaves,
so that a bud examined in ]\Iay would give one the impression that the
sepals begin their growth before the foliage leaves.

By May 1, in at least the most advanced bud, sepals and petals
have completely closed over the parts within (pi. 7, figs. 3 and 6).

50 L'liln rsil jj oj Calijornia J 'iib Lea I ions in Botanij ! '^'"'- '

At this lime llic youii^ flower-luul averages 2 to 8 iiiin. in Icngtli, the
size of the 1)11(1 and the (h'gi-ce of (h'V('h)i)ni('ii1 a|i|)ai-cii1 ly varxiiig
considerably, according to llic strength of Ilic planl. The aiillici's by
this date practically fill llic iiitcrioi- of llic Howcr-lmd. In general
the second bud in llie "crown" — llic first, as noted above, being func-
tionless — is the largest and most highly differentiated, each successive
bud thereafter being two weeks or more behind its immediate pre-
decessor.

What appears to be the critical period in development at which
the ajiex of the rootstock ceases to grow and llic buds nearest it are
arrested in their development occurs late in May. Buds seem to
gather momentum in their development, so that those which have their
])i-imordia well advanced before this critical period go on developing,
while in the case of those which are not sufficiently advanced further
development is suspended until the following year. As has been stated
above, when the development of a bud which has formed its primordia
is sharply arrested the result is usually fatal, still younger buds (fig. 1,
&'o) not so M^ell differentiated are usually able to resume growth upon
the arrival of the next active season. Small plants that form only
two buds and send up one shoot a year thus lay down their buds nearly
two years in advance. On the other hand, those plants which send
up four or more shoots a year begin to form some of their buds one
season and the others two seasons before they are to appear above
ground.

In June and July, following the period of suspended activity, the
entire terminal bud of the rootstock, or crown, is growing with maxi-
mum rapidity. All parts are increasing rapidly in size at this time,
but most noticeable is the renewed activity of the young foliage leaves.
They increase in size more rapidly than the young flower-buds and
overtop them by the latter part of July (pi. 7, figs. 4 and 7). There
is still a striking inequality in size and in degree of development among
the various buds, but by the middle of July, or earlier, the youngest
or innermost bud for next season is larger and better differentiated
than the oldest or outermost was in May. Sometimes when only two
buds are developing for next season they are nearly ecpial in size
and in degree of differentiation, and when the outermost — usually
functionless — bud (cf. fig. 1, />,) develoi)S in addition to the otlicrs it
is very likely to be smaller than its imniediate successoi'. It seems
that the checking of development of a bud as far advanced as this
outermost bud always is at the critical period has a niueli more per-

1916] Brandt: Notes on Trillium 51

manent effect than the checking of a bud less advanced. This outer-
most bud, in the relatively few cases when it thus actually develops,
tends to produce a smaller flower and seems less likely to produce a
fruit than the next. This statement is based upon field observations.
Because of the great increase in size of the buds for the next season
it becomes very easy from this time on to distinguish those whose
development has been arrested.

Somatic divisions are taking place in all parts of the young buds,
but more especially in the anthers. From the middle of May to the
middle of June the archesporial cells of the anthers are cut off (pi. 7,
fig. 8). In an anther of T. sessile var. giganteum there are usually
four rows of archesporial cells. These closely resemble the adjacent
cells, but may be recognized by their larger size, radial elongation, and
position immediately under the epidermis. They soon begin to divide
to form sporogenous and parietal tissue. As development progresses
the differences between sporogenous and parietal cells become more
and more noticeable. All the parietal cells which are arranged in
layers are greatly flattened as a result of periclinal division, their
cytoplasmic content is not very abundant and their nuclei are rela-
tively small but rich in chromatin. The sporogenous cells, on the
other hand, are not arranged in layers and have no definite form.
They are non-vacuolated and their cytoplasm shows a reaction to stains
different from that of the cytoplasm in other cells of the anther. Their
nuclei are large, but rather less densely filled with chromatin than
those of the parietal cells. There is therefore a great difference in
appearance between sporogenous and parietal cells, but the tapetum
is not yet cut off. Late in July the ovules begin to make their ap-
pearance as slight protuberances of the placentae, but their arche-
sporial cells do not appear until later in the year.

h. Trillium ovatum
Practically the same series of phenomena reported above for T.
sessile var. giganteum- occurs during this active period below ground in
T. ovatum, but a few exceptions should be noted. Mention has been
made of the fact that fewer buds are formed in T. ovatum, the number
being limited to two in average cases. It might here be noted that
the peduncle becomes noticeable before the flower-bud has attained
one-fifth of its mature length. The most noticeable difference is in
the rate of development of the floral organs. In some specimens
young pollen grains are present before the last of July. This very

52 UniversH !J of CdJifornia Puhlicatiutis in Bofaini [Vol. 7

early ai)pe'arane(' of i)ollcii in 7'. ovatain has a i)ai'all('l in tliosc plants
of T. grandifiorum, reported hy Atkinson (liSi)9), whose pollen was
fully matured in September. It is also interesting to note here two
statements which have been made with regai-d to the size of the flower-
bud in Eastern species of Trillium the season before flowering. Foerste
(1892) states that the largest bud of T. erythrocarpum Michx. which
he collected in (.-onnecticut in August was 5.5 mm. long, quite com-
parable in relative size to buds of T. sessile var. giganteum in June.
Smith (1896) states that a flower-bud of T. grandi/loriiin found in
central New York on July 11 was 2 luni. long witli anthers 1.7 nun.,
such a bud as one can find in T. sessile var. giganteum early in ]May.

IV. The Dormant Period, Augitst to January

a. Trillin))! sessile var. giganteum

The season from August to January inclusive is to be called the
dormant season, in the present paper. The terra "dornuint season"'
is again used here simply for convenience, just as the term "active
season" is used to designate the season from January to J\dy inclusive.
Many plants, as previously noted, cease their activity above ground
in May or earlier, while, on the other liand, others are appearing above
ground in November or earlier. Generally speaking, however, the
season from August to January inclusive is the season in which 7'.
sessile var. giganteum, growing in the vicinity of San Francisco Bay,
has no green leaves above ground.

This season may be divided into two parts, a season of decreased
activity and a season of increased activity. The first division con-
tinues until the beginning of the rainy season, late in September or
early in October. As noted above, the period May to July is one of
active and continued growth, resulting in the differentiating out of
the floral organs. During the period August to late September, the
dryest season of the year, development is practically at a standstill,
while with the opening of the rainy season growth becomes even more
rapid than during the period May to July. The period of decreased
activity is, however, characterized by the cutting off of the pollen
mother-cells and a resting stage preliminary to their rounding up and
entering upon the prophases of the meiotic divisions. The last sonmtic
divisions in the sporogenous tissues, giving rise to the full number of
pollen mother-cells, are completed in the latter part of August and

1916] Brandt: Notes on Trillium 53

early part of September. The anthers are at this time abnost fully
grown and nearly fill the interior of the flower-bud. The pollen-sacs,
which are still nearly colorless, are also very slender, owing to the
fact that the pollen mother-cells are not rounded off. They are very
angular in outline and, when crushed out of a living anther, cohere
firmly in a mass. The cytoplasm is dense and the nuclei are verj^
large. From one to three nucleoli are present and the open reticulum
is composed of thinly scattered chromatin granules (pi. 8, fig. 1).
The tapetum is definitely cut off, but its cells have not as yet become
so greatly enlarged as they appear in a mature anther, nor have their
nuclei divided.

The pollen mother-cells continue in the resting stage for four
weeks or longer before rounding off and preparing to divide. At
about the middle of September or early in October, when the rainy
season begins and the weather becomes cooler and the soil contains
some moisture, the underground organs of Trillium sessile var.
giganteum begin to show increased activity. This increase is manifest
first in a lengthening of the protective sheaths of the crown, and then
in an increase in size in all the parts of the stem buds for the next
active season. There is a very noticeable change in the appearance
of the anthers. Their length does not increase very appreciably, but
the pollen-sacs begin to increase rapidly in diameter and their color
begins to change from nearly white to pale yellow — the color of pollen-
sacs in which the reduction divisions are taking place — and finally,
as the pollen attains maturity the color becomes deep yellow. When
examined under the microscope the pollen mother-cells are found to
be rapidly increasing in size and their nuclei show a more than pro-
portionate increase (pi. 8, fig. 1). In some locules pollen mother-cells
are in synapsis, while in others, even in the same anther, the chromatin
is still distributed in the form of rather large granules throughout the
nuclear cavity. These "resting" nuclei are fully as large in diameter
as are those in synapsis (see pi. 8, figs. 1 and 2). I have not found
such an increase in size of the nuclear cavity during synapsis in
Trillium as Lawson (1911) reports in Smilacina.

The statement has already been made that in the same flower-bud,
or even in the same anther, in T. sessile var. giganteum the pollen
mother-cells in one locule may be in synapsis, while in another the
chromatin still forms an open reticulum. It may be well here to state
that the period during which the meiotic divisions occur extends over
two months — October to November — a condition in striking contrast

54 Univcrsitjf of ('(ilifornia Pii])Iirat{o)i.^ in Botcnn/ [Vol. 7

to that which Atkinson (1899, p. 10) found in T. grandifionim, where
"the period during which tlic division of the pollen mother-cells takes
place extends over seven or cigiit months." As in T. grandifionim.
Ilic time of division of the pollen mother-cells vai-ies with the season,
the location, and the degree of development of the other parts in the
individual. As noted above in the case of large i)lants wliidi produce
a numl)er of buds during the active season below ground, any two buds
may be strikingly different in the amount of development wliicli they
exhibit during the month of May. By October oi- November this
discrepancy in development appears to have been made up and the
reduction division takes place much more nearly simultaneously in the
various flowers of a rootstock than inii>'ht have been anticipated. It
seems clear that approximately six weeks elapse in average cases be-
tween the synaptic condition and the cutting off of the generative
nuclei. Of this period of six weeks, the first three include the stages
intervening between synapsis and the homotypic division and the
remainder are occupied by the stages between the homotypic divisions
and the cutting off of the generative nuclei.

Mention has already been made of the rapid growth of the pollen
mother-cells and their nuclei previous to synapsis. The locule also
may often increase so rapidly in size that the pollen mother-cells come
to lie in a mass in the middle of a large cavity. During synapsis the
chromatin often becomes so completely contracted that all semblance
of structure disappears. It is during this stage apparently that the
nuclei of many pollen mother-cells degenerate, cytoplasm and cell-
walls disappearing shortly thereafter. During the later prophases of
the heterotypic division the nuclear membrane seems to disappear and
the pollen mother-cells round up. Atkinson (1899, p. 12) states that
the double nature of the broad chromatin band is indicated by the
presence of a double row of dense bodies. In the material at hand
it was not possible to distinguish many granules in the bivalent chro-
mosomes because of thickness and density of the latter, but evidence
of their double nature was furnished by the occasional appearance of
a longitudinal split while they were in the e(|uatorial plate stage. In
the metaphase the chromosomes are arranged in a nearly flat equa-
torial plate as Vs whose free ends stand out around the spindle (pi. 9.
fig. 1), as Ernst (1902) reports for the megaspore mother-cell in Paris
quadrifolia and Trillium grandiflorvm. The s])in(lle fibers are at-
tached to the chromosomes at the apices of the \'s, the chromatin
becoming drawn out thin at the point of attachment, as Atkinson

1916] Brandt: Notes on Trillium 55

reports for the heterotypic mitosis in T. grandiflorum, and as Gregoire
(1912) reports for somatic mitoses in the same species and in Galtonia.
As the chromosomes migrate to the poles they gradually divide and
before the cross-wall is formed in the equatorial region twelve sep-
arate chromosomes may be found in each polar group. They are much
more slender than the bivalent chromosomes, are irregular in outline,
and a single row of granules is plainly discernible. The spindle fibers
disappear and a cross-wall is formed before the homotypic spindle
fibers appear, but a true resting stage was not found, such as Ernst
(1902) reported to intervene between the heterotypic and homotypic
divisions in the megaspore mother-cell of Paris. My results seem to
confirm those of Atkinson in this matter. The homotypic division
takes place so rapidly that I have been unable to obtain preparations
showing stages earlier than the polar groups of six chromosomes each.
Soon after the homotypic division has been completed cross-walls are
laid down to form the tetrad.

The tetrads soon break up into very angular young pollen grains
whose non-vacuolate cytoplasm appears to be very compact. Soon the
pollen grains begin to round out, but at no time in their development
does their cytoplasm become vacuolated, as Dorsey (1914) reports is
the case in Vitis species. As stated above, in approximately three
weeks after the homotypic division many of the pollen grains have
attained maturity, the exine has its characteristic markings, and the
nuclei are dividing. The microspore nuclei do not divide with the
uniformity in regard to point of time characteristic of pollen mother-
cells, and only a few division figures may be found at any one time in
a given locule (pi. 10, fig. 1). Many pollen grains in this species
never round up properly and many others are less than the normal size.

The cells of the tapetum attain their maximum size when the pollen
mother-cells are in the tetrad stage. Their nuclei are very large and
very rich in chromatin. When they divide, the chromosomes — having
apparently the full somatic number — form a very compact division
figure. Variations from the binucleate condition have not been found.
As the young pollen grains mature the cells of the tapetum gradually
lose their contents and become flattened against the outer layers.
Pollen sterility is often anticipated in an abnormal condition of the
tapetum, which may become separate from adjacent parietal layers or
may begin to degenerate before the pollen mother-cells divide. Other
tissues of the anthers also degenerate. Most frequent is the degener-
ation of the tissue between adjacent locules, which may include the

56 Univcrsil !/ of ('(ilifdniid I'lihlicdfidiis in liohnn/ fVoi,. 7

epidermis and eventually aid in deliiseeiiee. As tlie tapelum degen-
erates anther sap appears and sometimes is very abundant when the
pollen grains are about half grown, hut disappears soon after the
division of the microspore nucleus. From tlic early part of December
to anthesis, late in February, no changes take ])lace in the appearance
of normal i)ollen. During this period, however, the cells of the endo-
thecium are increasing theii- radial diametei- and develop radial thick-
enings on their walls (pi. I), fig. 2). The method of dehiscence of the
auther has been described above.

At the time when the pollen mother-cells are dividing the o\^iles
already have the nucellus and the integuments well differentiated
(cf. Coulter and Chamberlain, 1903, p. 52). The megaspore mother-
cell is discernible in the third layer of cells immediately under the
apex of the nucellus. It may be recognized by its slightly increased
size, nearly square shape, and large nucleus with rather open retic-
ulum (pi. 7, fig. 9). Nucellus and integuments grow with gradually
increasing rapidity until the arrival of the flowering season, but the
megaspore mother-cell changes but little in size and appearance at the
time the shoot appears above ground (cf. Alden. 1912. p. 448). As
the stem and leaves begin their rapid growth above ground the mega-
spore mother-cell begins to grow rapidly and take on an elliptical
form. When the leaves are becoming fairly well unfolded the hetero-
typic division occurs (pi. 7, fig. 10). Sufficient material was not ob-
tained for a critical study of this stage and the succeeding stages in
the formation of the embryo-sac, but from the fact that the heterotypic
division takes place after the shoot has appeared above ground and
that a large embryo-sac is present when the flower unfolds it seems
safe to assume that the female gametophyte in T. sessile var. giganieum
follows practically the same course of development as Ernst describes
for the female gametophyte in T. grandiflorum.

Actual confirmation of Ernst's conclusions can be made only with
regard to the earlier stages in the maturation of the female gameto-
phyte. Thus I have seen the formation of the cross-wall following the
first or reduction division of the megaspore mother-cell (cf. pi. 1,
fig. 10). This wall is curved and arches upward as the inner cell
develops at the expense of the outer (cf. Ernst, 1902, p. 14, ])1. 5, figs.
143 and 144). The inner, persistent cell enlarges, takes on an oval
form and becomes the young embryo-sac (cf. Heatley, 1916). During
its increase in size free nuclear division takes place to produce first a
binucleate embryo-sac. Ernst reports that the young embryo-sac con-

1916] Brandt: Notes on Trillium 57

tinues in this binucleate condition for two weeks or more {loc. cit.,
p. 20).

h. Trillium ovatum
As noted above, T. ovatum matures its pollen earlier in the season
than T. sessile var. giganteum. In general the pollen mother-cells
divide during August and September, and pollen is apparently mature
in all plants by the first of November. By the latter part of December
nucellus and integuments apparently have attained nearly their mature
size, and it seems probable that growth of the female gametophyte
begins before the shoot appears above ground. As in T. grandiflorum,
the megaspore mother-cell in T. ovatum^ becomes centrally located in
the nucellus.

V. Summary

1. The underground structures of a Trillium plant consist of a
stout subterranean stem, or rootstock, bearing perennial roots and a
large terminal bud. As the rootstock adds live tissue at its growing
end from year to year the basal end decays. Thus at any one time
the rootstock, instead of representing the actual age of the plant, rep-
resents only a portion of its age, three to fifteen years, according to the
rate of decay.

From two to five or more roots are sent out each season, according
to the size and vigor of the plant, and live from two to ten years. By
contraction they draw the rootstock down into the ground, and, usually
in the case of T. ovatum Pursh, sometimes in T. sessile var. giganteum
H. & A., orient it in a horizontal direction.

The terminal bud in September contains not only the young stem-
buds for the next growing season but also two or more young stem-
buds for the second season in advance. Usually the first of these
stem-buds, which already has cut off the primordia of lateral organs,
fails to develop farther. Each stem-bud is formed in the base of a
fleshy scale-like body which in turn arises in the axil of a bud-scale
or sheath. At least a rudimentary stem-bud is found in the axil of
every bud-scale.

The rootstock of T. sessile var. giganteum bears lateral branches or
offsets, which in time become separated by decay of tissue from the
parent plant, thereby affording a ready means of vegetative repro-
duction. Such offsets are rarely found in T. ovatum.

58 University of Cdlifoniid I'uhllcaft'nuft in Bofroiij [Vol.7

2. Tile Mclivo season in T. sessile vai-. <ji<j(iitUtiin lasts fi'oiu Feb-
ruary 1o July. In the vicinity of San Francisco Bay, however, unfa-
vorable atmospheric and soil conditions cause a majority of plants to
end their active season prematurely. On the other hand, in T. ovatum
the active season is greatly i)rolonged. The former species ripens its
fruit in July, the latter in ]\Iay.

3. During the active season the young stem-bud, or buds for the
next year whose activity had been suspended late in the previous
active season, begin to grow anew and differentiate various parts. At
the same time additional stem-buds are laid down of which some may
develop into buds for the next season. Tliose laid down last, however,
and therefore least differentiated, are checked in May, so that they
cannot develop farther until the next active season arrives.

4. In T. sessile var. giganteum a plate consisting of four rows of
archesporial cells is cut off in each locule of the anther in May. Late
in August or in September the tapetum is cut off.

5. Tlie dormant season in T. sessile var. giganteum lasts from
August to January inclusive. However, activity is renewed in the
underground organs when the rainy season begins, in September or
October.

6. The pollen mother-cells divide after the parts of the plant have
begun to show renewed activity in growth. If a plant has several
stem-buds for next season their pollen mother-cells divide at different
times. Other causes affect the time when the meiotic divisions occur,
but it may be said that in general the pollen mother-cells of T. sessile
var. giganteum divide in October and November. The various phases
of the two divisions, including synapsis, occupy not more than three
weeks, and the pollen grains are cutting off generative nuclei in three
weeks after the completion of the homotypic division. ^Mature pollen
may be found in November and December. From that time onward
no changes are taking place in the anthers except the development of
the mechanism of dehiscence. In many pollen mother-cells the nuclei
degenerate in synapsis or before, and many pollen grains fail to
develop properly. In T. ovatum the pollen mother-cells divide in
August and September and practically all i)ollen is mature by
November 1.

7. The megaspore mother-cells are differentiated just before the
microspore mother-cells divide and change but little in appearance in
T. sessile var. giganteum before the young shoot appears above ground.
The megaspore mother-cell divides and the female gametophyte de-

1916] Brandt: Notes on Trillium 59

velops while the young shoot is growing most actively. At the time
of anthesis the embryo-sac is mature. In T. ovatiim the large size of
the ovules in midwinter and the rapidity with which the plant comes
into flower in spring indicate that the megaspore mother-cell divides
before the young shoot has appeared above ground.

8. In Trillium grandifforum in the eastern United States and in
northern Germany the dormant period is approximately equal in
length to the dormant period in T. sessile var. giganteum in the vicinity
of San Francisco Bay. According to Atkinson, in T. grandiflorum
the pollen mother-cells are cut off and pollen may even be fully ma-
tured at the beginning of the dormant period, whereas in T. sessile
var. giganteum pollen mother-cells are cut off wdthin the dormant
period. In most plants of T. grandiflorum, however, maturation of
pollen occurs after the subterranean parts have begun to show indi-
cations of renewed growth. The same is true of T. sessile var. gigan-
teum. In T. sessile var. giganteum, just as in T. grandiflorum and
T. recurvatum, the megaspore mother-cell divides after the young shoot
has appeared above ground. In T. ovatum, on the other hand, pollen
is usually mature before the close of the prolonged active season, and
the megaspore mother-cell is apparently ready to divide two months
before the shoot appears above ground.

I wish to thank Dr. T. H. Goodspeed, under whose direction the
work has been carried on, for his constant and valuable assistance in
the preparation of this paper. I also wish, among others who have
assisted in its preparation, especially to thank Professor W. A. Setchell.
I am, further, indebted to Professor R. Ruggles Gates for a number of
helpful suggestions and criticisms.

Transmitted April 21, 1916.

60 Unii'rrsih/ of ('(iJifor)iia PuhlicatiovK in Botan}! | Voi- 7

VI. Literature Cited

Alden, I.

1912. A contribution to the life-history of Uinktria sessilifolia, Bull. Tor.
Bot. (Mul). vol. :i!), p. 4:59.
Atkinson, Geo. F.

1899. Studies on reduction in plants, Bot. Gaz., \o\. 28, p. 1.
B.\ILEY. L. H.

189(5. The Survival of the Unlike, New York.

1902. Encycloiiedia of American Horticulture, vol. 4, pp. 1853-56. New-

York.
Britton, N. L., and Brown, H. C.

191.3. Illustrated Flora of the Northern States and Canada, vol. 1, pp. 52.3-
526. New Y^ork.
Carter, A.

1892. Notes on Pollination, Bot. Gaz., vol. 17, p. 19.
Chamberlain, C. J.

1898. Winter characteristics of certain sporangia, Bot. Gaz., vol. 2.j, ].. 124.
Coulter, J. M., and Chamberlain, C. J.

1903. Morphology of the Angiosperms. New York.
Davis, Charles A.

1897. TriJliiDv f/raiidiflonmi. its variations normal and teratologieal, Bot.
Gaz., vol. 24, p. 187.
Dorset, M. J.

1914. Pollen development in the grape with special reference to sterility,
Univ. Minn. Agr. Exp. Sta. Bull. 144.
Ernst, A.

1902. Chromosomen reduction, Entwickelung des Embryosaekes, und Be-
fruchtung bei Paris qvadrifolia L., und Trillium firaiidifonim Salisb.,
Flora, vol. 91, pp. 1-46, T. I-VI.
FoERSTE, Aug. F.

1891. Abnormal phyllotactic conditions as shown by the leaves or flowers

of certain plants, Bot. Gaz., vol. 16, pp. 159-166, especiallv pp.
16.3-164.

1892. On the relations of certain fall to sjiring lilossoming plants, Bot. Gaz.,

vol. 17, ]ip. 1-S.
Gates, F. C.

1908. Comparison of the season at Chicago and Urbana, Illinois, Anier. Bot.,
vol. 14, p. 45.
Goodspeed, T. H., and Brandt, E. P.

1916a. Notes on the Californian species of Trillium Tj.. 1, T'niv. Calif, l^ubl.

Bot., vol. 7, no. 1.
1916&. Notes on the Californian species of TriUinm L.. Tl, ibid., vol. 7, no. 2.
Gray, Asa

1887. S(diool and field book of botany, ]i. 440, revised edition. New York.
Greene, E. L.

1894. Manual of the Bay Region. San Francisco.
Grf.ooire, Victor

1911-12. Les phenomenes de la metaphase et de I'anaphase dans la caryocines
somatique, Ann. Soc. Sclent, de Bru.xelles, nos. 3 and 4, p. 339.

1916] Brandt: Notes on Trillium 61

Heatley, Margaret

Un6. A study of the life-history of Trillium cermium L., Bot. Gaz., vol. 61,
p. 425.
Howard, W. L.

1915. An experimental study of the rest period in plants — the summer rest
period of bulbs and herbaceous perennials, Missouri Agr. Exp. Sta.
Res. Bull., no. 16.
Jepsox. W. L.

1911. Flora of Western Middle California. San Francisco.
Kerxer and Oliver

1895. Natural history of plants, vol. 2, pp. 452-454. New York.
Lawsox, a. Axstruther

1911. The phase of the nucleus known as synapsis, Trans. Roy. Soc, Edin-
burgh, vol. 42, pt. Ill, no. 20.

LUDWIG F.

1900. On self -sterility, Jour. Roy. Hort. Soc, London, vol. 24, pp. 214-17.
Pacific Coast Geographical Survey, Botany, vol. 2, p. 181. Cambridge, Mass.,

1880.
Pfeiffer, Waxda ~Sl.

1912. The morphology of Leiteneria floridana, Bot. Gaz., vol. 58, p. 189.
Rattax, Volny

1888. California flora and "West Coast botany. San Francisco.

RiMBACH, A.

1900. Physiological observations on some perennial herbs, Bot. Gaz., vol. 30.

p. 171.
1902. Physiological observations on the subterranean organs of some Cali-
fornian Liliaceae, ihid.. vol. 33, p. 401.
RoBERTSox, Charles

1896. Flowers and insects, Bot. Gaz., vol. 21, p. 266.

1897. Seed crests and myrmecophilous seed dissemination in certain plants,
ihid., vol. 23, p. 288.

Smith, Miss A. M.

1896. Abortive flower-buds of TriUiitm. Bot. Gaz., vol. 22, pp. 402-3.

\'1I. Hxi'LAXATION OF PlATES

PLATE 7

Stem-biuls of TriUinm .<.<'.s.s/7r var. (jif/anteum from the rootstock crown in
various stages of development. iSemi-diagrammatic.

Fig. 1. Longitudinal section of bud collected in November, primordia not
all present, still very rudimentary, fifteen months before its active season above
ground commences.

Figs. 2 and 5. Bud in February: figure 5, entire, figure 2 in longitudinal
section. Primordia more highly differentiated than in November. The sepals
slightly surpass the foliage leaves.

Figs. 3 and 6. Bud collected in May: figure 6 entire and figure 3 in longi-
tudinal section. The foliage leaves are seen to be shorter than the sepals.
Sepals and petals now include the stamens and carpels. At about this time the
archesporial cells are cut off in the young anthers.

Figs. 4 and 7. Bud as it w^ould appear late in July: figure 7 eutire and
figure 4 in longitudinal section. The foliage leaves are now" growing rapidly
and equal or slightly surpass the sepals in size. As in figure 3, the maturing
anthers practically fill the interior of the fiower bud. In T. ovatum the flower
bud at this season would be elevated in a short peduncle. For comparative
sizes of the different parts of the stem-buds in September, see figure 1, page 44,
h,, b, .

Fig. 8. Portion of cross-section of a young anther showing archesporial
cells just cut off. Collected May 29, 1915.

Fig. 9. Portion of young ovule showing the niegaspore mother-cell. Col-
lected October 7, 1914.

Fig. 10. First division of megaspore mother-cell. Collected February 26,
1915.

[62]

UNIV. CALIF. PUBL. BOT. VOL. 7

[BRANDT] PLATE 7

8

10

PLATE 8

Fig. 1. Youug polliMi luotlier-cells in the resting condition inmietliately
preceding synapsis.

Fig. 2. Young pollen mother-t*ells in synaptic condition.

Both conditions were foinul in different anthers of the same flower laid
collected November 26, 1914.

[64]

UNIV. CALIF. PUBL, BOT, VOL. 7

[BRANDT] PLATE 8

Fia-. 1

\

PLATE 9

Fig. 1. Pollen mother-cells iu the metaphase of the heterotypic division,
both polar and longitudinal views of the spindle. The reduced number of chro-
mosomes is clearly discernible in at least one of the division figures. Collected
October 21, 1914.

Fig. 2. Portion of a mature loeule showing parietal tissue and a few mature
pollen grains. Collected March 11, 1916.

[66]

UNIV. CALIF. PUBL. BOT. VOL. 7

^';

[BRANDT] PLATE 9

- 1

Iff.

PLATE 10

Fig. 1. Germination of the mierospore. Collected November 5, 1913.

Fig. 2. Adjacent walls of two locules of a mature anther just before de-
hiscence. A layer of small cells has been formed which cuts across the walls of
both locules to assist in effecting dehiscence. Collected March 7, 1915.

|68J

UNIV. CALIF. PUBL. BOT, VOL. 7

[BRANDT] PLATE 10

Fig. 1

Fig. 2

UNIVERSITY OF CALIFORNIA PUBLICATIONS

IN

BOTANY

Vol. 7, No. 4, pp. 69-100, plates 1 1-17 January 17, 1917

NOTES ON THE CALIFOENIAX SPECIES OF

TRILLIUM L.

IV— TERATOLOGIC AL VARIATIONS OF TRILLIUM SESSILE

VAR. GIGANTEUM H. & A.

BY
THOMAS HAEPER GOODSPEED

The genus Trillium has long been recognized as unusually prone
to the production of teratological forms, and the occurrence of such
abnormalities has from time to time been noted for practically all the
more common and well-defined species. It is the purpose of the
present note to gather together a number of the references to tera-
tology in the genus, and particularly to describe the nature of a
number of abnormal forms found in the Californian representative
of T. sessile L.

A rather thorough examination of the available literature has re-
sulted in the accumulation of a considerable number of references to
Trillium teratology. Since a compilation of the literature on the
subject has not, to my knowledge, been attempted, and especially since
many observers have published accounts of abnormalities apparently
without knowing that similar specimens have a number of times been
described elsewhere, it has seemed best to bring together certain of
the somewhat scattered references and to arrange them under the
species to which they severally refer. No claim is made for the
completeness of the list submitted below, since often in the older liter-
ature only the briefest notices are given to teratological phenomena.
It seems probable, however, that most of the more important contri-
butions have been examined.

IBff>

70 TJnivcrsihj of California Pnhlications in Bolamj | Voi.. 7

Trillmm gTandiflorum uMichx.) Salisb.
This six'cics jippcars to he llic most hi<i^hly vari;ihk' of the nine
hero noted, as is indicated hy llie predominating numl)er oi" references
in the literature on teratology in the genus. Hritclu^r ni)02) lias
deserihed at some length the extraordinary sei'ies of aljnormal forms
wliieli occur near Syracuse. N(nv Yoi-k. He includes measurements
of such characters as "length of stem," "length of petiole," "size
of leaf blade," "length of peduncle," etc., together with notes upon
the more highly abnormal variations observed. Flower color was
found to vary for the petals from "typical white or piiil<, tlii-ough
white with green center stripe, to solid green." The following is
Britcher'.s condensed tabulation of the measurements on variation in
size of parts {loc. oil., \^. 106) :

Lengtli of plant stem varies from U mm. to 340 mm.

Leugth of petiole 0 160

Width of leaf blade 22 124

Length of leaf blade 30 157

Length of peduncle 2 220

Length of sepal stem 0 44

Width of sepal blade 9 37

Length of sepal blade 26 78

Length of petal stem 0 64

Width of petal blade 8 50

Length of petal blade 18 80

Length of filament 1 34

Length of ovary 1 30

Length of ovary stalk 0 23

Length of style 2 27

Length of anther 0 20

Atkinson (1905) notes two teratological modifications. In the first
case the flower shows the "compound pistil expanded into three leaf-
like members." This in general corresponds to Britcher's plant 128:
"The place of the ovary is taken by three leaflike parts with stems
10 mm. and blades 5 by 14 nnn. in size. Within this circle are two
pollen-bearing stamens with filaments 4 and 7 nun. long and anthers
5 and 10 mm. long" (p. 174). Atkinson's second abnormality is not
noted by Britcher. who does not definitely describe any instance in
which all the parts of the perianth are strictly foliaceous or in which
a dui)lication of parts occurs. In this second case "the nine parts of
the perianth are green and the outei- whorls of stamens are expanded
into petal-like members." Prom the illustration there is obviously
an extra whorl of perianth segments; the outer whorl of expanded

wn] Goodspeed : Notes on Trillium 71

stamens are anther-bearing along their margins, while the inner whorl
of stamens and the pistil are normal.

Bishop (1902) gives cases included under Britcher's plant 103
(p. 178) and others in which the peduncle is longer than the stem
below the leaves. He refers to other cases showing an extreme length
of petiole (Britcher's plant 109, and others, p. 189), and states: "The
green-petalled variety is common . . . ^Multiplication of organs is com-
mon, in one case extending not only to the petaloid and bractlike
forms, but also to the leaves." Davis (1897) reports the examination
of a large number of abnormal specimens, among which "the simplest
departure was markedly the presence of green stripes in the petals,"
adding that "this striping was accompanied by a lengthening of the
petioles and degeneration of the pistil." He found over fifty indi-
viduals in which the leaves were reduced to bracts or had entirely dis-
appeared (cf. Britcher, plant 149a, p. 174) and notes that "in such
forms the stamens are the most stable of the organs of the flower,
only a few reversions to the leaf type occurring, while the pistil was
usually sterile, rarely containing ovules, frequently being reduced to
the leaf form and sometimes containing well-marked whorls of leaves. ' '
Dickson (1897) describes, from Hamilton, Canada, a long-petioled
specimen corresponding to a number of the plants described and fig-
ured by Britcher (cf. p. 189). Dudley (1886, p. 99) speaks of
"double-flowered specimens" that "have about fourteen parts to the
perianth." Gary (1905) describes the long-petioled and green-
flowered forms noted by Britcher and mentions a plant tetramerous
throughout. ]\frs. Goodrich (cf. Meehans' Monthly, vol. 8, p. 182,
1897) reports the common variations found by Britcher from the
same locality in which his collections were made.

Gray (1875a) mentions a long-petioled form with green, striped
petals and a plant with similar petals, no leaves and having sepals
which were "more foliaceous than is usual and are fully two inches
long, quite equalling the petals." These plants were from St. Louis,
IMichigan, and compare to individuals found later by Britcher. Gray
(1875&) notes still other forms described by Britcher and says that
they suggest large specimens of T. nivale because of their long-petioled
leaves, but that they are sharply distinguished from this latter species
by "the acute base and acuminate apex of every leaf." Hankensen
(1870) describes plants of the Syracuse type from Newark, New York,
and lays emphasis upon the fact that plants without leaves apparently
have larger, foliaceous sepals. In this connection, attention might be

72 Universih/ of CaJifnrii'xi riihlicdlimis in Bolanjj | Vol. 7

called to an ohscrvatioii by Peek (1885) who. in describing the usual
long-petioled, green-striped variation, says that "the specimens indi-
cate a coincidence between the petioles, peduncles and green color of
petals. Generally the longer petioles are accompanied by longer
peduncles and broader green stripes on the petals. The coincidence
between form and structure is tiuly remarkable."

Holzinger (1901) figures a plant in which the leaves are normal
but the flower consists- of six whorls of three green, foliaceous perianth
segments each. Osband (1894) describes an unusual plant, the flower
of which exhibited two whorls of three normal sepals each ; similarly
two whorls of petals, the outermost being striped or w'hite; and three
stamens, of which only one was normal, one being "expanded into a
half-sized petal" and anther-bearing, and the third sterile. The
ovary is reported as normal in size but "near the top of one of the
carpels arises an outgrowth about half an inch long, white, doubled
together and drawn over at the top like a hood." On the edges of
this outgrowth there were "two pollen-bearing lines about an eighth
of an inch long. ' '

Pollard (1901) reprints the figure and description of a plant noted
by Mrs. Kellerman (1893). The plant bore a conspicuous double
flower, and the double condition reappeared for some years from the
original rootstock, w'hich was brought into a garden. The flowers
were white in color and varied in number of whorls of petals from
nine to thirteen. The reproductive tissues were almost completely
suppressed. Smith (1879) comments upon the number of abnormal
individuals and their wade distribution in ^lichigan. Nothing is added
to the types noted by Britcher, but the statement is made that in
abnormal plants "the ovules are sometimes twelve in number, but
generally less, often none, or only one or two, the cavity being filled
by an enlarged placenta." It might be mentioned in this connection
that rootstocks of normally fruiting plants of TiilJium sessile var.
giganteum H. & A. when brought under greenhouse conditions have
apparently no dormant period (cf. Brandt. 1916), and each of the
flowering shoots matures a large ovary which contains only a spongy
mass of placental tissue. With regard to the prevalence of abnormal
individuals in ^lichigan, J\leehan (18946) reports that many ])laiits are
found with "four leaves, three sepals, and four petals, a less number
with four leaves, three sepals, and five petals. ' ' Since these forms are
found only at the end of the season, a causal relation is suggested
between lateness of flowering and abnormal variation.

1917] Goodspeed: Notes on Trillium 73

Stevens (1894) notes the case of a flower in which ''a stamen and
a petal were united edge to edge, one anther cell being obliterated by
the union." The plant was otherwise normal. Victorin (1904) de-
scribes two plants apparently tetramerous throughout. In one flower
a petal was anther-bearing. A flower, presumably of 1\ grandiflorum,
is mentioned by Meehan (1894a) consisting of "eighteen beautiful
white petals." A normal whorl of leaves is spoken of and it is prob-
able that the sepals were also normal (ef. Meehan, 1894&, who describes
a "very perfect eighteen-petalled flower"). The rootstock apparently
produced abnormally flowering shoots for three years after being
transplanted into a garden. Peck (1888) notes two plants in which
"not only petals but also sepals were wholly white."

Trillium erectum L.

Andrews (1906) briefly mentions an abnormal individual of this
species, the flower of which showed flve petals, four stamens, and two
styles, while the outer whorls were normal. Deane (1910) notes a
plant with a single whorl of four leaves, five sepals partially petaloid,
four petals, eight stamens, one of which was double, and an eight-
winged ovary. Bishop (1902) refers to "a few instances where the
whorls are repeated but none with the lengthening of leaf and flower
stems" so characteristic of the extreme variations of T. gmndiflorutn.
Clute (1895) reports a flower of this species with four petals — each
edged by a pollen-bearing anther — and flve stamens. James (1884)
describes a teratological specimen of the following type : four sepals,
four petals, eight stamens, "four pistils and a four-celled ovary,"
whorl of leaves normal, a fourth leaf on the peduncle halfway between
flower and leaves, and the sepals wholly or partially petaloid.

The remainder of the pertinent references to T. erectum which
have been found deal with the color of the flower and especially the
occurrence and origin of the var. album Pursh. Gray (1908, p. 293)
describes the petal color of erectum as "brown-purple, or often white
or greenish or pinkish." Thus a considerable range of color-forms
seems well recognized. Hall (1870a) comments upon the occurrence
of the normal "purple" form associated with the less strongly pig-
mented forms. He says: "I have watched individual plants of this
variety (var. album) as closely as possible, and seen the regular purple
flower in their place the next year; and have also seen the var. album
flower in places when the year before a purple flower had ai)peared.
T have noticed, too. that, as a rule, the var. albu))i is a starveling:

74 Vniversiln of ('(ilifoniid /'liblicatioiis in Botany [Vol.7

rarely as large or as healthy as the pui'ple i)ljni1s in the midst of which
it grows. Also I ]i;iv(> ivirdy found a var. album away from the
company of the normal plant, and never more than three or four
plants together, though thi-y are quite common." Elsewhere (18706)
he notes the occurrence of a plant with "creamy green" colored petals
for a number of ycni-s after l)(Mng transplanted to a garden. Gary
(1905) mentions a phint with cream-colored petals and red ovary.
and Horsford (1890) reports a considerable local variation in the
color forms of this species. Thus, from Pennsylvania comes "the true
var. album with pure M'liite flowers as white as those of T. grandi-
ftorum. Indeed, a casual observer would take them for yrandifJornm.'"
A shade intermediate between the "pure white and the typical
T. erect um" and a form in which tlie "petals are of a yellowish shade
M'ith a sprinkling of purple" are found in Connecticut. "Botli forms
of the variety and the true purple type" occur together. Clute (1908)
gives some evidence in this connection. He says that "in eastern
America it is almost invariably dark purplish red in color; in the west
it is invariably pure white without even a tinge of red." Plants of
the red form sent to him from New Britain, Connecticut, came up in
his garden (Joliet, Illinois) the following spring bearing, with one
exception, pure white flowers, and in this one instance there was only
a "trace of red in the stamens." I have been unable to learn more
as to the later history of these plants. Clute (1907) also describes a
"yellow trillium" in which the "inner tliree leaves of the perianth
were wholly a pure sulphur yellow ; the outer leaves of the perianth
were pale green below, but as bright a yellow above as the inner part
of the perianth." The species is not noted, but the statement is made
that the ])lant "grew amongst a larger number of specimens of T.
grandiflorum." The remark is made that "seeds from an aberrant
plant will usually produce the same form." In seeking an explana-
tion of the origin of vai'. dllnim, it seems jiossible tliat in general
it may represent a type of "undeveloped ilower" such as is com-
mon in the sessile-flowered Califoi'uian species (cf. Goodspeed and
Brandt. 1916?;), although the occurrence of intermediate shades be-
tween white and red might ratliei' l)e taken to preclude such an
interpretation. Certainly conti'dllcd breeding experiments are very
necessary in this comiection.

Strictly teratological variations in <-nvluiit seem to consist solely
of vai'ious degrees of doubling without accompanying malformations
of a striking character.

1917] Goodspeed: Notes on Trillmm 75

Trillium sessile L.

Andrews (1906) describes two abnormal plants of this species.
One bore fourteen "floral leaves," and it was noted that "no trace of
the usual stamens or pistil was present, all the floral organs being
completelj^ transformed into floral leaves." The other plant had four
leaves, three (?) sepals, "four large partly-greenish petals, six small
stamens and four styles." Foerste (1891 and 1894) in two articles
discusses the phylotaxy of a number of teratological individuals, de-
scribes the effort of the typically trimerous plant to accommodate itself
to the dimerous conditions, and comments upon the ability of one
abnormal individual to maintain the quaternate arrangement with
only six floral envelopes. He mentions five cases of doubling in this
species. One plant was four-parted throughout, the whorls being
normally oriented. A second plant bore four leaves, two sepals, four
petals, two of which were partially sepaloid, and six stamens. Another
plant is described as being four-parted, except that one petal is missing
and but six stamens (all double) are present. A fourth plant was
more highly abnormal and irregular, bearing five leaves, two sepals,
three petals, seven stamens, and a four-celled ovary. Hopkins (1902)
describes three teratological shoots, all from the same rootstock. One
showed three whorls of three leaves each, four petals, no sepals, five
stamens, three styles and stigmas, and the ovary six-angled ; another,
two whorls of three leaves each, no sepals, six petals, seven stamens,
four styles and stigmas, and the ovary eight-angled; while the third
corresponded to this last, except that nine stamens were present and
there were but two styles and stigmas. The author assigns the con-
dition of this teratological individual to over-nutrition. Morris (1902)
notes the fact that the production of single, long-petioled leaves from
the rootstock has been widely mentioned in general descriptions of the
genus. He describes and figures a rootstock of sessile which bears two
normal, flowering shoots from the terminal rootstock crown and in
addition two small lateral offsets, each bearing a single one of these
long-petioled leaves. As stated by the author, "these leaves sprang,
each from the node nearest the apices of two tiny branches from the
main rootstock." Hankensen {lac. cit.) also mentions plants of grandi-
flot'um with a single radical leaf.

This question of the production of single, long-petioled leaves from
the rootstock of Trillium is one which has throughout been of con-
siderable interest in our studies upon the Calif ornian species. As

76 Univrrsihf of California l'uh!lrafio))s in Bolainj [Vm.. 7

elsewhere noted in this series of papers, tl)e (piite eliaraeleristie ster-
ility of var. giganteum seems to be compensated for by very active
vegetative reproduction ])y offsets of the rootstock. In this connec-
tion it may l)e recaHed that oval urn, a heavily fruiting species,
practical!}' never n'i)rodiices vegetatively. A single rootstock of T.
sessile var. giganteum may exhibit from three to six normally tlowering
or normal flowerless shoots from the rootstock crown and an ecpial
number of offsets of varying ages which may bear either a single, simple,
long-petioled leaf, or a small but normally three-parted whorl of
leaves, or, moi-e rarely, two opposite leaves. Plate 11 shows two root-
stocks, one of which bears a two-leaved shoot from the crown, and
the other a single, long-petioled leaf from the same position on the
rootstock and in addition a similar leaf from an offset. These forms
occur very commonly in California, and the problems connected with
their origin and the mode of transition from the simple one-leaved to
the three-leaved condition are being investigated in connection with
those brought up by the occurrence of undeveloped flowers.

Trillium undulatum Willd.

Beattie (1905. p. 40) comments upon the large proportion of two-
stemmed plants of this species in Rhode Island and notes an instance
of phyllody in one shoot of a two-stemmed individual, while the other
shoot apparently was entirely normal. Peck (1878) describes from
Oswego, New York, a plant in which there was almost perfect doubling
of the number of parts in whorls of leaves and flower parts.

Deane (1908a, &, and 1910) describes a number of interesting tera-
tological specimens and gives important evidence as to the recurrence
of such variations in succeeding years. Three abnormal forms in the
Gray Herbarium are described. The first Avas apparently normal
except that the sepals were greatly enlarged and foliaceous, somewhat
corresponding to the forms noted by Beattie (loc. cit.), and the
second was octamerous throughout except that there were seven
stamens. Deane re-examined the "polymerous" specimen described
by Gray (1878) and found it to consist of eight sepals, one Avith a
"white petaloid growth on one side," eight petals, "at least twenty
stamens," six carpellary divisions of the ovary, and seven leaves, one
of which was double. Another plant is described from INIaine Avhich
bore four leaves, three sepals, three petals, six stamens, one more or
less petaloid, and a one-celled ovary. All parts were normal in shape

1917] Goodspeed : Notes on Trillium 77

and size. Of special interest is the description of the recurrence dur-
ing two seasons, following the one during which the plants came under
observation, of a group of teratological individuals. In general, the
plants strikingly resembled the remarkable specimens of sessile de-
scribed by Hopkins {loc. cit.). The station is Squam Lake, Holder-
ness. New Hampshire, and there seems to be no doubt but that cor-
responding abnormal variations were produced by the same rootstock
or rootstocks over a period of three seasons.

Trillium cernuum L.

I have found but two references to teratology in this species. Owen
(1894) describes two specimens from New Hampshire. One showed
a normal whorl of three leaves and above it ''three whorls of three
leaves each," the sepals larger than common, petals with a white stripe
running dowTi the center and a green stripe on each edge and one of
them two-parted, four stigmas. The other specimen exhibited a
"rosette of two whorls, a third abnormal in this specimen also, but
lifted one-half inch above the others to the base of the flower." Tracy
gives the following description of an abnormal individual : ' ' One petal
and two sepals of the ordinarj^ form and color, while the third sepal
has been replaced by a perfect leaf, and the other two petals have a
green stripe through the center."

Trillium nivale Riddell.

Andrews (1906) reports that in this species "some slight deviations
have been observed ... in the way of a union of the floral parts."

Trillium recurvatum Beck.

Andrews (loc. cit.) notes a plant with twenty-three "floral leaves"
in which no trace of the "usual stamens or pistil was present, all the
floral organs being completely transformed into floral leaves which
were considerably larger (with the exception of the central ones) than
the same parts in normal flowers growing near them. ' '

Trillium ovatum Pursh. and T. sessile var. giganteum II. & A.

In these two western species teratological variations have on only
two occasions been described, so far as I have been able to determine.
In thousands of specimens of T. ovatum which have come under
observation in our studies, no abnormalities have been found. Deane
(1911). however, describes an interesting plant from Lake Cushman.

78 Universih/ of Cnlifonila P\(blications in Botany [Vol. 7

Wa.shiii^ton. The tlircr leaves are normal but there; ai"e twenty-
four "petals," eighteen of them in "regular alternating whorls of
three, the first, third, and fifth, and the second, fourth, and sixth
whorls being superposed upon each other," while "the six petals in
the center of the flower form a partially closed cluster." The largest
of the six "petals" was 1.8 cm. in length and nnich "crumpled."

In 7'. sessile var. giganteum we have noted a eonsiderable number
of teratologieal variations, the following description o\' wliirli, with
notes on their recurrence from the rootstoek, represents the principal
purpose of thi.s paper. Miss Eastwood (1896) has given the only de-
scription with which I am acquainted, of abnormalities in this western
ally of T. sessile L. She reports on "three abnormal specimens of
white-flowered Trillium sessile var. Californicum found in the San
Bruno Hills of San Mateo County." The first specimen was four-
parted throughout, "even the ovary," and the stamens were eight in
number. The second bore six leaves, six outer divisions of the peri-
anth and five inner, ten stamens, and a six-celled ovary. The third
was normal except that "one of the outer divisions of the perianth
was a true leaf." She states that "this species is exceedingly variable
in the color, shape and size of its flowers; but such abnormal forms
are rare."

I wish in what follows to describe the abnormal shoots produced
by seventeen different rootstocks of T. sessile var. giganteum. Of these
seventeen rootstocks, twelve have been or are at present under observa-
tion in garden cultures.

1. Collected in Berkeley Hills, Berkeley, California, by Dr. R. P.
Brandt ; rootstoek not preserved. One shoot normal in every respect
except the condition of tlie stamens. These were six in number, but
each was flattened and the connective expanded to form structures very
similar in appearance to a corresponding condition in T. grandiflorum
described and figured by Atkinson (loc. cH.). In hi/a' the iiulividiiMl
stamens varied from 32 X 5 mm. to 32 X 8 mm., the width measure-
ment being taken across the widest portion of the expanded stamen.
Anther cells containing apparently noi'nially matured pollen were
present on both edges of each stamen. Partial pistillody of all the
stamens also occurred here, in that ridges of lighter-eoloi-ed tissue
stood out from the inner surfaces of the expanded connective and this
expanded connective was inflated to a considerable degree. These
ridges of tissue undcmbtedly represent stylar and stigmatic .surfaces,
although theii- lila-ncss to such structures of the j^istil is not as

1917] Goodspeed: Notes on Trillium 79

striking as is the case of pistollody of the perianth shown in plate 15,
figure 1. No trace of ovules was to be found in the inflated regions
of the expanded connective.

2. Collected as 1 ; rootstock not preserved. Normal in every way
except that one petal was much curved and bore a shrunken and
empty anther cell along half its inner, curved edge.

3. Collected in the Berkeley Hills; the rootstock not preserved.
This plant showed four leaves, averaging 103 mm. in length and
95 mm. in greatest width ; four sepals, averaging 46 X 13 mm. ; four
petals, of which three were normal and averaged 65 X 16 mm., and
the fourth was curved and anther-bearing as in number 1 above ; seven
stamens, and four styles and stigmas. The leaves and flower parts
were regular and, as indicated, the flower was practically symmetrical.

4. First collected in the Berkeley Hills by Dr. Brandt in 1915,
marked and collected again in 1916 ; the rootstock not preserved. In
1915 this rootstock bore at least two shoots. When this plant was first
noted a large collection of mature ovules was being made for cyto-
logical purposes; and when the pistils of the shoots from this root-
stock were cut open, supernumerary pistils were found within the
ovary at the base. This condition was apparently characteristic of
the pistils of all the shoots from this rootstock, and the structures
themselves, in the case of two of the shoots, were preserved in fluid.
One of these intracarpellary pistils is shown in plate 17. d. The size
of the main carpellary structure was not perceptibly increased and it
was normally provided with ovules on parietal placentae while the
small included pistils of the type shown in the figure occurred at the
base, imbedded in a mass of spongj' proliferations of the placental
tissue. As will be seen, ovules were present on these intracarpellary
pistils, although borne on their surfaces, while the tissues of the ovary
within were undifferentiated.

Plants from this rootstock collected the following season (1916)
showed merely a trace of the same condition, but distinctly enough
to indicate that it is subject to recurrence from this rootstock.

5. Collected in the Berkeley Hills, 1914; rootstock not preserved.
Numbers 5 and 6 represent nearly identical instances of pistillody in
which the petals are o\aile-bearing. Number 5 has been elsewhere
described (Goodspeed. 1916) and the nature of the abnormal condi-
tion is shown in plate 15, figure 2, and in plate 17, a. The description
given in the case of number 6 below will, with modification there noted,
hold for this plant also.

80 Universih/ of ('olifor)tin I'lihlirations in Botany [Vol. 7

6. Collected in Marin County, 191G; rootstock in pot 4G oi" the
T'rillium cultures in the University of California Botanical Garden.
This rootstock bore two tlowering .slioots, one of which was normal in
every respect; the condition of the other is shown in phitc 1."). (i<^ure 1.
As will be noted, (he leaves were normal in number and they were
also normal as to shape and si/c. while the three sepals were partially
petaloid. There were six stamens, all normally provided with the long
filaments characteristic of plants from tliis locality. The pistil was
also normally formed and ovule-bearing. The striking teratological
condition is seen in the petals. They were reduced below the sepals
in size, and along their rolled edges ovules were bor-ne. while modified
stylar and stigmatic tissue was present along the inner surfaces. The
petals of nvimber 5 above were identical except that no trace of stylar
and stigmatic tissue was found. The cross-section of the modified
petal of number 5 shown in plate 17. a, is applicable as de-scriptixc of
the condition found here (cf. ITalsted. 1891).

7. Collected in the Berkeley Hills by Dr. Brandt and Professor
R. R. Gates, 1916; rootstock in pot 40. This rootstock bore six shoots,
two of which were entirely normal and four exhibited abnormal modi-
fications in the way of a distinctly fasciated condition and a more or
less corresponding increase in number of parts in each whorl in all
but one case.

In all four abnormal shoots, the stem was quite strongly flattened
and often distinctly ridged. In one case a small sixth leaf was inserted
beloM" the main whorl of five leaves and from it a ridge or petiole ran
down the entire length of the stem.

Two shoots bore five leaves, four sepals, four petals, eight stamens,
and from four to six styles and stigmas, while the ovary was usually
composed of one large cell with from six to eight parietal placentae.
A third abnormal shoot was similar with the exception of the presence
of a sixth leaf noted above. There were various more or h^ss unim-
portant modifications of organs, such as the union of two stamens, a
partially petaloid stamen, a petal curved and anther-bearing along one
edge. In one case there appeared to l)e two distinct whorls of leaves
distinguished by size as well as heiglit of insertion. One df these
three shoots is figured in plate 14. figure 2, and the natui-c of the
justils is shown in ])late 17. & and c.

The fourtii ahnormal shoot of tliis rootstock is figured in plate
l;i At the toj), along the i-idgc formed by the mucli-fiattened
stem, fourteen leaves were borne, and just bi'low a complete three-

1917] Goodspeed : Notes on Trillium 81

parted whorl of leaves corresponding in shape and provided with a
small undeveloped flower in the center (cf. Goodspeed and Brandt,
19166). Within the leaves on the ridge at the top of the stem were
a series of five such undeveloped flowers. The leaf measurements show
that there were five distinctly larger leaves, and these made up a some-
what irregular outer whorl. Some years ago another plant of almost
exactly this description was found in the Berkeley Hills, but the
rootstock unfortunately was not kept under observation.

8. Collected in the Berkeley Hills, 1916, by Dr. Brandt ; rootstock
in pot 47. This rootstock produced six flowering shoots, five of which
were normal throughout. The sixth was again a fasciated shoot with
flattened stem bearing two whorls of three leaves each, the leaves of
the outer whorl being somewhat the larger. There were also four
sepals, all of them f oliaceous ; five petals, nine stamens, three long and
six short ; six styles and stigmas, and a single large-celled ovary with
six parietal placentae.

9. Collected in Marin County, 1916; rootstock in pot 11. This
plant is figured in plate 12. As will be seen, the rootstock produced
two shoots, both of which bore a single four-parted whorl of leaves
with an undeveloped flower. The other shoot is arranged on the plan
of 5, with ten stamens and a one-celled ovary with five placental
surfaces.

10. Collected in Marin County, 1916; rootstock in pot 45. This
rather unusual individual is shown in plate 14, figure 1. Its striking
abnormality consists in the condition of the sepals, which are reduced
to shriveled, scalelike structures. The petals were, also, of a peculiar
shade of yellowish-green, very brilliantly shining, while the ovary was
white and the stamens light violet in color.

The following individuals have been under observation for a year
or longer in the garden cultures :

11. Collected in I\Iarin County, 1911 ; under observation for five
years :

1911 — Rootstock collected with two entirely normal flowering

shoots.
1912 — Two shoots produced. One shoot entirely normal and the

other bearing four leaves, three sepals, four petals, and

six stamens, one of which was connected with a petal.
1913 to 1916 — One or two entirely normal shoots produced each

vear.

82 Uniucrsilij uf (.'alifoniia I'libluations in Boianxj I Vui.. 7

12. Rootstock colleetod in lf)ll ; until iDKi produced two normal
shoots each year wilh uiideveh)ped flowers. In 1916 there appeared
three very small shoots, one of which bore four haxcs aud an unde-
veloped flower, while Ihc ollici- Iwo shoots were normal.

13. Collected in IDIO; until 1!)14 produced miiiiial shoots. In 1914
the two shoots figured in plale Ki. fi<fur(» 2, both bore large undeveloped
flowers. On the shoot shown in the right in the figure, one leaf was
diminutive in size, thin and transparent, and rather tightly curled
about the flower. It was, further, inserted within the leaf whorl.
In the figure it is shown partially folded back to display the flower.
In the two following seasons the shoots from this rootstock were
entirely normal.

14. Collected in Marin County in 1911, this rootstock had produced
five offsets, which were separated from the parent rootstock and
planted in separate pots. The parent and the offset rootstocks have
since that time borne normal shoots with the exception of one of the
offsets, which in 1912 produced the two shoots seen in plate 16, figure
1. Since 1912 this rootstock has given rise to normal plants.

15. AYhen collected in the Berkeley Hills in 1912, this rootstock
bore two shoots — one normal and in flower, and the other with a
flattened stem, six leaves, and a small undeveloped flower. In 1918
and 1914 two normally formed whorls of leaves were borne with un-
developed flowers. No shoots were produced thereafter.

16. Collected by Dr. Brandt in 1915; rootstock in pot 149. There
is no record as to place of collection or as to condition of plant, except
that it was highly abnormal. In 1916 this rootstock produced one
shoot with three leaves, three large foliaceous sepals, five petals, and
three stamens. One stamen was normal in shape and size, but was
strongly bent inward for a distance of 3 mm. below the tip. Another
stamen was petaloid and anther-bearing along only one edge, and
the third was distinctly double.

17. Collected in 1915; rootstock in pot 150. Again there is no
record except that the plant was abnormal in number of parts. In
1916 this rootstock produced six shoots. Two of them were normally
flowering; two were likewise normal, but provided with undeveloped
flowers; one bore four leaves, and the sixth seven leaves, with un-
developed flowers in both cases.

if^i7] Goodspeed : Notes on Trillium 83

DISCUSSION

There seems to be little doubt that within the Eastern species
grandiflorum there is a widely distributed form or variety which ex-
hibits correspondingh' rather marked variations in the various locali-
ties from which it has been reported. This form, sometimes referred
to as var. variegatum Pk. (cf. Peck, 1888), is characterized by long-
petioled leaves, striped perianth segments, and often an elongation
of the peducle. Apparently no other American species possesses any
similarly well-marked and widely distributed form or variety based
upon abnormal structural variations. Color variations in most of
the species are striking and well recognized but probabh' represent,
with possibly one or two exceptions, extremes of the normal fluctuating
variation of a highly variable genus. Although var. giganteum con-
tains no characteristic form of extreme or teratologieal structural vari-
ation, the extent and intensity of such variation seems remarkable.
That practically all organs of the flower may exhibit duplication of
parts, suppression or extreme modification and metamorphosis, has
been indicated in the description of specimens above.

The question of the recurrence from the rootstock and, in the case
of perennial plants in general, of abnormal shoots, and the more im-
portant question as to the origin of such inherently abnormal root-
stocks, are of maximum interest and significance. The genus Trillium
would seem to furnish rather imusually valuable and readily investi-
gated material for such studies. As yet, however, the evidence at
hand, if not contradictory, is at least highly fragmentary and in gen-
eral rather unsatisfactory so far as this genus is concerned. The
observations on Eastern species, especially those of Deane on un-
dulatum, indicate that, in the case of well-defined and extensive tera-
tologieal variation, the abnormal condition may be expected to recur
year after year in the shoots produced by a given, inherently abnormal
rootstock. Our garden cultures of var. giganteum seem to suggest
much the same condition, although the more important evidence will
come from the results of the next few years. There is no doubt, how-
ever, that such apparently relatively simple variations in structure as
the change from the trimerous to the tetramerous condition do not
recur in succeeding years from the same rootstock. Our cultures
appear to indicate that such changes may be environmentally deter-
mined, a fact further suggested by the ease with which many of the

84 U)iu'rrsif!i of ('oJifor)iia ri(J)li(afio)is in liofainj | Vol. 7

most fniulaiiu'iital oiiloyciictiL' processes of tlie plant can be modified
l)v ciillural iiiHucnces. Field studies and garden eultiires, however,
do not support tlie suggestion, elsewhere rioted, that these transitory
or the other more deeply seated teratological variations are connected
with the time oi' appearance of the shoot during the growing season.

An inherently abnormal rootstock, which may be expected always
to produce some abnormal shoots each season, must have had its origin
in a seed, since there is no evidence, on the one hand, that variations
induced in rootstocks l)y environmental ccmditions will recui- or, on
the other, that the individuals produced by offsets (asexual reproduc-
tion) ditfi'r in any way from the shoots formed from the parent root-
stock. This latter jooint has been thoroughly investigated in seeking
an explanation for the origin of the remarkable range of color forms
found in var. gigaiiieum. It is unfortunate that no one has grown
the progen.y of a teratological individual. It is apparent that among
the Eastern species certain of the abnormal individuals observed do
produce a little seed, and this is, I judge, especialW true of T. grandi-
florum var. variegatum. It is highly desirable that someone determine
the proportional occurrence of the variety and the normal form from
the seed of var. variegatimv. Unfortunately var. giganteum, even in
the case of entirely normal lowers, only rarely produces viable seed,
and thus it is probable that only the question of the recurrence of
teratological variations from the rootstock can be investigated.

In general the initial production of the seeds giving rise to tera-
tological rootstocks can apparently be explained most readily on the
assumption that the parent plant was in a basically heterozygous con-
dition. The occurrence of abnormalities following hybridization,
especially in the case of wide crosses, is well recognized. A case some-
what in point is that of species Iwbridization in the genus Nicotiana,
in whi<'h the back-cross N. sylvestris X the F^ hybrid N. tahacum var.
macroplujlla X A. sylvestris yields a rather high percentage of very
abnormal individuals. In this connection it might be noted that the
wide range of color forms in T. sessile var. gigantciim seems best ex-
plained on the assumption of initial crossing of widelj^ differing forms,
an assumption perhaps borne out by the rather widespread sterility
in this variety. Finally, it should be borne in mind in connection with
this latter point that T. ovatum, which exhibits a relatively narrow
range of variability in flower color and in form and size characters
and rarely gives rise to teratological indi\i(luals. jtroduees an abund-
*ance of viable seed.

1917] Goodspeed : Notes on Trillium 85

SUMMARY

1. An effort has been made to tabulate the more important refer-
ences in the literature to teratological variations found in the following
species of Trillium: grandifloriim, erectum, sessile, luidulatum,
cernimm, nivale, recurvatum, ovatum, and sessile var. giganteum.

2. Seventeen teratological variations of T. sessile var. giganteum
are described.

3. Evidence is given to show that such relatively superficial vari-
ations as the change from the trimerous to the tetramerous condition
do not recur in succeeding years from the same rootstock, while more
profound structural variations may so recur.

4. It is suggested that the origin of such recurring teratological
variations may be due to the heterozygous condition of an original
parent plant.

Transmitted September 8, 1916.

86 Uliivcrsilii (if Cdlifoniid riihlicftlions in liohmn |Vi)i.. 7

LITERATURE CITED
Andrews, F. M.

15)06. Some monstrosities in 'J'rillium. I'lant World, vol. It, p. lui.
Atkinson, G. F.

litU.1. A college text-hook of l)otany (ed. 2, New York), PI'- 321-322.
Beattie. F. S.

1905. Remarks on Rhode Jsland plants. Rliodora, vol. 7, ]>. '.V.K
Bishop, I. T.

1902. Variations in TriUium flowers. Plant World, vol. ."), ji. II.
Brandt, R. P.

1916. Notes on the Californian species of Trilliitm L. J 11. I'niv. Calif.
Publ. Bot., vol. 7, no. 3, 2». 31.
Britcher, H. W.

1902. Variation in Trillium gravdi forum Salisb. Me. Agric. Kxji. Stat.,
Bull. 86.
Clute, W. N.

1895. A malformed TriUiiim. Meehan 's Monthly, vol. 5, \\. S-l.
(Clute, W. N.)

1907. A yellow Trillium. Amer. Bot., vol. 12, p. 83, 1907.

1908. A remarkable change of color in Trillium. Ibid., vol. 14, p. 33.
Davis, C. A.

1897. Trillium grandiftorum (Michx.) Salisb.; its variations normal and
teratological. Reviewed in Bot. Gaz., vol. 24, p. 187.
Deane, W.

1908a. Teratological forms of Trillium undulaium. Rhodora, vol. H), p. 21.
1908&. More teratological forms of Trillium undulaium. Ibid., p. 214.

1910. Teratology in Trillium. Ibid., vol. 12, p. 163.

1911. Teratology in Trillium ovatum Pursh. Ibid., vol. 13, p. 189.
Dickson, J. M.

1897. Meehans' Monthly, vol. 8, p. 85.
Dudley, W. R.

1886. The Cayuga flora. 1886, p. 99. Quoted from Deane (1911).
Eastwood, Alice.

1896. Trillium sessile. Erythea, vol. 4, p. 71.
Foerste, a. F.

1891. Abnormal jdiyllotactic conditions as shown by the leaves or flowers
of certain plants. Bot. Gaz., vol. 16, p. 159.

1894. Notes on dedoublement. Ibid., vol. 19, p. 460.
(!arv, L. B.

1905. Variation in TriUium. Plant World, vol. 8, p. 257.
(Goodrich, Mrs. L. L.)

1897. Trillium variations. Meehans' Monthly, vol. 8, p. 182.
Goodspeed, T. H.

1916. Pistillody of the perianth in Trillium scisile var. giganteum. Madrofia,

vol. 1, no. 1, p. 19.
Goodspeed, T. H., and Brandt, R. P.

1916a. Notes on the California species of Trillium Ij. T. T^niv. Calif. I'ubl.

Bot., vol. 7, no. 1, p. 1.
1916&. Notes on the Californian species of Trillium L. II. Ibid., vol. 6,

no. 2, p. 25.

1917] Goodspeed: Notes on Trillium 87

Gray, Asa.

1875a. Two remarkable forms of Trillium. Bull. Tor. Bot. Club, vol. 6, p. 272.

1875b. Sporting Trillium grandiflorum. Ibid., p. 277.

1878. Jour, of Science, ser. 3, vol. 15, p. 153.

1908. New manual of botany (ed. 7, New York).
Hall, I. H.

1870a. Trillium erectum L., var. album Pursli. Bull. Tor. Bot. Club., vol. 1,
p. 21.

1870b. Trillium erectum L. Ibid., p. 36.
Halsted, B. D.

1891. Intracarpellary pistils and other floral derangements. Ibid., vol. 18,
p. 246.
Hankenson, E. L.

1870. Monstrosities of Trillium grandiflorum. Ibid., vol. 1, p. 21.
HOLZINGER, J. M.

1901. A green Trillium. Plant World, vol. 4, p. 132.
Hopkins, L. S.

1902. A rare freak of the Trillium. Ibid., vol. 5, p. 182.
HORSFORD, F. H.

1890. Garden and Forest, vol. 3, p. 240.
James, J. F.

1883. Bull. Tor. Bot. Club, vol. 10, p. 57.

1884. Bot. Gaz., vol. 9, p. 113.
Kellerman, Mrs. W. A.

1893. Jour. Cinn. Soc. Nat. Hist., vol. 16, p. 49.
(Meehan, Thomas.)

1894a. Double Trillium. Meehans ' Monthly, vol. 4, p. 70.
1894b. Trillium grandiflorum. Ibid., p. 134.
Morris, E. L.

1902. "Occasional" leaves of Trillium. Plant World, vol. 5, p. 92.

1903. Abnormal Trilliums. Ibid., vol. 6, p. 87.
Osband, L. a.

1894. Abnormal plant growths. Amer. Nat., vol. 28, p. 706.
Owen, M. L.

1894. Trillium cernuum L. Bot. Gaz., vol. 19, p. 337.
Peck, C. H.

1878. Report of the Botanist. 31st Ann. Kept. N. Y. State Mus., p. 53.

1885. Report of the Botanist. 38th Ann. Rept. N. Y. State Mus., p. 108.
1888. Report of the Botanist. 42nd Ann. Rept. N. Y. State Mus., p. 132.

Pollard, C. L.

1901. Double TriUinms. Plant World, vol. 4. p. 213.
Smith, E. T.

1879. A Michigan TriUium. Bot. Gaz., vol. 4, p. 180.
Stevens, F. L.

1894. Teratology. Bull. Tor. Bot. Club, vol. 21, p. 490.
Tracy, Mrs. C. T.

1883. Abnormal flowers. Bull. Tor. Bot. Club, vol. 10, p. 71.
Victorin, Fr. Marie.

1914. line variation meristique remarquable du "Trillium grnndifiorum."
Le Nat. Canadien, vol. 40 (ser. 2, vol. 20), p. 1]3.
Wright, S. H.

1879. Bot. Gaz., vol. 4, p. 232.

PLATE ]1

Entire plants of Trillium sessile var. fjitjanteum, showing single and two-
leaved conditions.

88]

UNIV. CALIF. PUBL. BOT. VOL.

[GOODSPEED] PLATE II

PLATE lii
Two ahiiormal shoots and the rootstock described under 9, on page 81.

90

UNIV. CALIF. PUBL. BOT. VOL, 7

[GOODSPEED] PLATE 12

15U

i '■

PLATE 13
One of the abnormal shoots of the rootstoek described under 7 on page 80.

[92]

UNIV. CALIF. PUBL. BOT. VOL. 7

[GOODSPEED] PLATE 13

PLATE 14

Fig. 1. One of the abnormal shoots of the rootstock described under 10 on
page 81.

Fig. 2. Abnormal shoot of the rootstock described under 7 on page 80.

[94]

<

c:

CD

o

o

o
o
o

a

CO
"D
m
m
O

"D
|—
3>
H
rri

PLATE 1.')

Fiji'. L Abnormal flower of the shoot from the rootstock described under
fi oil page 80.

Fig'. 2. The same tyi>e of tlower of tlie shoot from the rootstock described
under 5 on page 79.

I f)6 1

UNIV. CALIF, PUBL. BOT. VOL. 7

[GOODSPEED] PLATE 15

'Ih

f

Fig. 1

Fm. 2

PLATE 16

Fig. 1. One normal and one abnormal shoot from tlie rootstock described
under 14 on page S2.

Fig. 2. One normal and one abnormal shoot from the rootstock described
under 11 on page 81.

5)S

>

d

CD

O

o

o
o
o
o

"0

PLATE 17

«. A cross-section of an ovule-bearing petal described under rootstocks 5
and 6 on pages 79 and SO.

b. View of the abnormal jiistil and increased number of stamens cliaracter-
istic of the abnormal shoots of the rootstock described under 7 on page 80. X 2.

c. Pistil of one of these same abnormal shoots showing the ovary opened
longitudinally. X 2. ,

d. One of the intracarpellary pistils of the flower from tlie rootstock de-
scribed under 4 on page 79. X S.

[100]

UNIV. CALIF. PUBL, BOT. VOL. 7

[GOODSPEED] PLATE 17

UNIVERSITY OF CALIFORNIA PUBLICATIONS

IN

BOTANY

Vol. 7, No. 5, pp. 101-157 August 14, 1919

A PRELIMINARY LIST OF THE UREDINALES

OF CALIFORNIA

BY *- •

WALTEE C. BLASDALE

CONTENTS

PAGE

I. Introductory statement 101

II. The need of experimental culture work 102

III. Plan of the present paper 103

IV. Acknowledgments 104

V. General features of the Uredinales 104

VI. List of species 105

VII. Index to the species of Uredinales 150

VIII. Index to the species of host plants 153

I. INTRODUCTORY STATEMENT

The study of the Uredinales of California began with the pioneer
work of Dr. H. W. Harkness and Justian P. Moore, who in 1880
published a Catalogue of Pacific Coast Fungi, which enumerates fifty-
five species belonging to this group. Unfortunately this list does not
give the specific names of a large number of the host plants referred
to and also the localities are often very indefinite, which makes it diffi-
cult to correlate this list with the work of subsequent collectors. Some-
what later Dr. Harkness, either in conjunction with M. C. Cooke and
J. B. Ellis, or independently, published in Grevillea and in the Bulle-
tin of the California Academy of Sciences a number of new species
including many Uredinales. The extensive collections of Pacific Coast
fungi which Dr. Harkness had presented to the California Academy
of Sciences before his death were destroyed in the great fire of 1906,
with the exception of the types of new species discovered by him,
which fortunately were preserved through the foresight of Miss Alice
Eastwood, the present Curator of the Herbarium.

102 University of Calif orm<i PuhlicaUans in Botany [Vol. 7

In 1882 Ml. E. \\. I), llohvay began work on the rust flora of
California by an extended exploration of King's River Canon and
vicinity, wliii-li lirought to liglit a large number of species, some of
which were })ul)lislied as new by Dietel and Holway, especially in
Erythca and Ucdwiyia. With tlie aid of Mr. Holway this work was
continued by the writer, and collections were made in the region about
San Francisco Bay (1893 and later), Shasta and Siskyou counties
(1894), Tuolumne and Calaveras counties (1895), Lake Tahoe (1897),
and Mendocino and Humboldt counties (1896 and 1899). Contempo-
raneously and in the succeeding years a large number of collectors
did work in other regions. Of these especial mention should be made
of A. J. McChitcliie (Pasadena), S. B. Parish (San Bernardino),
J. Burtt Davy (Berkeley), W. R. Dudley, and C. H. Thompson (Stan-
ford University). In more recent years extensive collections have
been made by Ellsworth Bethel, Carl T. Baker, David Griffiths, E. E.
Heller, Dr. Meinicke of the Federal Forest Service, and Professor
Home of the Agricultural Experiment Station of the University of
California.

IT. THE NEED OF EXPERIMENTAL CULTURE WORK

Although much more extended collections and critical studies in
both field and laboratory must be made before our knowledge of the
California representatives of this group of fungi can be considered
either exact or complete, it is believed that an attempt to summarize
the information now available will be of assistance to those working
on the rust flora of the state and will give at least an approximate
idea of its extent and content. The need of systematic culture experi-
ments designed to establish with certainty the life histories of those
species which are known to be heteroecious is a pressing one. The
results obtained in other regions of the United States, especially
by Arthur and his collaborators at Purdue Universitj^ suggest
the species of host plants probably concerned in the life histories of
many of the Pacific Coast species ; but the differences in the climatic
conditions of the two regions and of the species of host plants found
in them make it extremely desirable to confirm these suggestions by
actual experiments. It is to be regretted that owing largely to the
lack of such data it has been found necessary to include in the list
here presented a considerable number of form species, that is, aecial
and uredineal forms not definitely correlated with the mature forms

1919] Blasdale: Uredinales of California 103

necessarj^ for their identification. Apparently an unusually large
number of Pacific Coast forms winter over, by means of uredineal
spores, and produce telia only occasionally and in very small amounts
or fail entirely to do so.

Culture experiments are also greatly needed for the purpose of
ascertaining the specific identity of many of those species which
develop on closely related host plants, such, for instance, as the forms
of Puccinia which are found on the different genera of the Onagraceae
and Compositae. Such experiments in other regions have shown great
variability as to the closeness of the adaptation between the rust
and its host. In some instances, as illustrated by the classical work
of Eriksson and Hennings with the grain rusts, this relation is an
extremely narrow one and it becomes clearly necessary to recognize
species or subspecies based upon purely physiological distinctions.
It has been shown by Bethel, on the other hand, that certain species,
such as Puccinia suhnitens and P. stipae, produce their aecia on hosts
belonging to genera representing widely different natural families,
and it is not improbable that too much emphasis has been placed upon
the host plant as a criterion in ascertaining the specific limitations of
the rusts.

III. PLAN OF THE PRESENT PAPER

As one of the chief objects of the present publication is to assist
collectors in naming new collections and to tabulate all the known
forms and their accompanying host plants, the different genera, which
can usually be readily recognized if the mature form is at hand, have
been arranged alphabetically, and where a large number of species
are represented those species which occur on host plants belonging to
the same natural family have been grouped together and the groups
arranged according to the sequence of families found in Jepson's
Flora of Middle Western California.

The references cited are designed merely to identify with certainty
the form referred to and to indicate where a description of it can be
found. Only those synonyms have been given which are necessarj^ to
show where certain species, whose validity is not accepted, belong in
the list.

The data represented by the list is based almost wholly upon speci-
mens in the herbarium of the writer, and the name of the collector
of each specimen cited is given in parenthesis except where the col-
lection is that of the writer, in which case it is omitted.

104 University of California Piiblications in Botany [^ou 7

IV. ACKNOWLEDGMENTS

The completeness of the sui^ej' of the rust flora of California here
presented is in a large measure due to the assistance of the collectors
whose names have been cited, who have furnished specimens or have
given additional data regarding them. Especial obligations are due
Mr. Ellsworth Bethel for a large amount of information regarding the
heteroecious species, and for extended collections of grass and sedge
inhabiting species from the southern part of the state both of which are
poorly represented in most of the collections which have been made
up to tlie present time. I desire also to express my thanks to Pro-
fessor J. C. Arthur, who read over the entire list of species here
presented and made many valuable suggestions. It will also be
()l)vi<)us that much help has been derived from such portions of
Arthur's arrangement of the North American species of Uredinales
{North American Flora, vol. 7) as have been published, of the critical
work of Holway in that portion of his North American Vredineae
which has been published, and Jackson's Uredinales of Oregon, pub-
lished in the Memoirs of the Brooklyn Botanic Garden.

V. GENERAL FEATURES OF THE UREDINALES

The Uredinales represent a group of fungi whose vegetative parts
consist of a much branched and septate mycelium, which develops in
the leaves, branches, and fruits of certain seed producing plants and
ferns and abstracts nourishment from the cells of the surrounding
tissues by means of haustoria. The reproductive parts are represented
by five different types of spores, one or more of which are frequently
lacking and which differ greatly as to form and size as well in the
mode of production and germination. The different spore forms are
the following :

Spermatia, designated by the symbol 0, are produced in sub-
globose or flask-shaped spcrmogonia and are minute hyaline bodies,
which can be made to germinate but do not appear to perform any
important function in the life history. They usually appear with
or slightly before the aecia.

Aeciospores, designated by the symbol I, are produced in cup-
sha]ied or cylindrical aecia, which have a more or less well developed
peridium, or in masses, frequentlj^ surrounded by paraphyses, in
which a peridium is lacking. They are always one-celled and are
formed by the successive abstriction of the ends of closely packed

1919] Blasdale: Uredmales of California 105

liyphae. They have a hyaline epispore and a number of germ pores
but rapidl}^ lose their ability to germinate.

Urediniospores designated by the symbol II, are usually produced
in subepidermal sori and ultimately burst through the epidermis as
light brown or yellow pulverulent masses. They are produced singly
at the ends of hyphae, are always one-celled, either globose, elliptical
or pyriform, possess several germ pores and are either echinulate or
tuberculate.

Teliospores, designated by the symbol III, are formed like the
urediniospores, but sometimes appear singly, either in or on the tissues
of the host. They usually possess a definite pedicel and may be
fascicled or compacted into a crust or a cylindrical column. Each
spore may consist of a single cell or of as many as twenty cells;
each cell usually possesses a single germ pore. They possess a thick
epispore, which is dark brown or black in color. On germination
they produce a thick promycelial tube, which ultimately produces four
cells, each of which develops a single basidia and sporidium. When
the sporidia germinate they produce short tubes which are able to
penetrate the tissues of the proper host plant. The formation of
basidia is one of the most distinguishing characteristics of the group.

Many species show Avell characterized alternation of generations
on host plants belonging to the same species (autoecious) or on host
plants belonging to different species (heteroecious).

VI. LIST OF SPECIES

. The genera are arranged alj)habetically, and the species numbered
consecutively throughout the paper.

AECIDIUM

Under this form genus are grouped those aecial forms which have
a true peridium and for which no other stage has as yet been recog-
nized.

1. Aecidium Collinsiae Ell. and Ev.

Bull. Washb. Lab., vol. 1, p. 4, 1884.
On Collinsia hicolor Benth., Berkeley and Ukiah.

2. Aecidium Graebnerianiim Hennings

Hedwigia, vol. 37, p. 273, 1898.
On Hdbcnariu dilatata (Pursh.) Hook., Death Valley (Coville) ;
on H. leucostachys Wats., Mount Eddy (Copeland).

lOG Universifij of CaUfornm PubUcatimis in Botany [Vol. 7

3. Aecidium Triglochinis Diet, and ITolw.

Erythca, vol. 7, j.. i>S, 1899.

Oil Triglochin conciinid Davy, Lassen County, type collection
(Davy).

4. Aecidium Valerianellae (Biv.) Bernh.

Stirp. Ear. Sci., vol. 4, p. 28.
On Vahrianflla coiigesta Lindr., ]\Iill Valley and Applegate.

BAEODROMUS arth.
Ann. Mycologici, vol. 3, p. 19, 1905.

Cycle of development includes pj^cnia and telia only. Telia sub-
epidermal but erumpent ; teliospores one-celled, ellipsoid, produced in
chains but compacted into masses.

5. Baeodromus californicus Arthur. Ill
Ann. Mj'cologici, vol. 3, p. 19, 1905.

On Senecio Doiiglasii DC, Lytle Creek, San Bernardino Mountains
(Parish).

BUBAKIA ARTH.
Result. Sci. Cong. Bot. Yienne, p. 338, 1906.

Uredinia erumpent and pulverulent; urediniospores borne singly
on pedicels. Telia subepidermal and indehiscent ; teliospores one-
celled, compacted into dense masses.

6. Bubakia Crotonis (Cke.) De Toni. II, III
Grevillea, vol. 6, p. 137, 1878.

On Croton californicus Miill. Arg., San Francisco, Long Beach
(Bethel).

CALYPTOSPORA kuhn
Hedwigia, vol. 8, p. 81, 1869.

Aecia erumpent, cj'lindrical with thin-walled peridium. Telia
forming compact layers, indehiscent. Teliospores ellipsoid, usually
four-celled.

7. Calyptospora columnaris (Alb. and Schw.) Kiihn. 0, I, III
The aecial stage has been shown hy Winter (Hedwigia, vol. 26,

p. 28, 1887) to develop on a large number of species of Ahics, includ-
ing A. magnifica Murray and A. concolor Lindl. Arthur (Mj'cologia,

1919] Blasdale: TJredinales of California 107

vol. 2, p. 231, 1910) also obtained aecia on potted plants of A. Fraseri
(Pursli.) Lindl., which he had infected with C. cohimnaris from Nova
Scotia, and Professor Frazier found aecia identical with those obtained
by Arthur on Abies halsamea at Picto, Nova Scotia. In California
Meinicke has collected similar aecia on Abies concolor Lindl. at Lake
Talioe, and on A. rnagnifica Murraj^ at Heckel's, Lassen County.

Telia on Vaccinium ovatum Pursh. and V. parvifolium Smith,
Santa Cruz, Marin, Mendocino and Humboldt counties; on V. mem-
hranaceum Pursh., Tahoe National Forest (Meinicke) ; on V. Chandleri
Jepson. Mount Eddy (Copeland).

COLEOSPOEIUM lev.
Ann. sci. nat. I, Bot., vol. 8, p. 373, 1847.
Aecia definite, erumpent ; peridium colorless. Uredinia erumpent,
pulverulent, spores in chains. Telia waxy, indefinite, liberated by
disintegration of host nnh'. Heteroecious.

8. Coleospormm Bletiae Diet. II, III

Hedvi-igia, vol. 37, p. 218, 1898.
On Phajus Wallichii Lindl. Reported by Arthur (iY. A. Flora,
vol. 7, p. 86) on plants imported from Japan, but not found by the
writer although frequent inquiries were made for it.

-9. Coleosporium Campanulae (Pers.) Lev. 0, I, II, III

Syn. Fung., p. 217, 1801; Ann. Sei. Nat., ser. 3, Bot., vol. 8, p. 373, 1847.

The aecial stage not known from California, but found on the
Atlantic Coast on Pimis rigida and P. virginiana.

Uredinia and telia on Campanula persicifolia Linn., San Francisco.
Probabl}' a recent introduction as it was first noted by the writer in
1915 on the Exposition grounds.

10. Coleosporium Madiae, Cooke. II, III

Grevillea, vol. 7, p. 102, 1879; Sydow, Ann. Mycologiei, vol. 2, p. 30, 1904.

Aecial stage probably represented by Peridermium californicum
Arth. and Kern (see no. 35).

Uredinia and telia on Madia capitata Nutt., 31. anomala Greene,
M. dissitiflora (Nutt.) T. and G., 31. elegans Don., 31. sativa Molina,
M. Nutt alia Gray, Centromadia pungens (H. and A.) Greene, and
Zonanthis corymb osa (DC) Greene. Very common throughout the
central portion of the state.

108 Universitj/ of California Puhlicatians in Botany [Vol. 7

11. Coleosporium Solidaginis (Scliw.) Thiim. 0, I, II, III

Bull. Torr. Club, vol. 6, p. 216, 1878.

Tlie aeeial stage probably represented by Peridermium montanum
Arth. and Kern, which is reported from California by Hedgecock
(Phytopathology, vol. 6, p. 64, 1916).

Urediuia common in the bay region on Aster radulinus Gray, A.
Menziesii Lindl., A. chilensis Nee., and Solidago calif ornica Nutt. On
Aster sp., Yosemite Vallej^ (Bethel) ; on Solidago confinis Gray, Pasa-
dena (McClatchie).

CRONARTIUM fries
Observ. Myc, vol. 1, p. 220, 1815.

Aecia erumpent, inflated, usnally forming galls. Uredinial spores
born singly on pedicels. Telia erumpent, scattered ; teliospores formed
in chains which adhere to form a cylindrical column, one-celled,
usuallj^ fusiform. Heteroecious.

12. Cronartium coleosporioides (Diet, and Holw.) Arthur. 0, I,

II, III

Erythea, vol. 1, p. 247; N. A. Flora, vol. 7, p. 123, 1907.

The aeeial stage {Peridermium Harknessii Moore) produces spher-
oid galls on Pinus contorta Dougl., P. Sahiniana Dougl. and P.
ponderosa Dougl. Widely distributed throughout the Sierras, also
at Mount Diablo (Blasdale and Bethel).

Uredinia and telia on Castilleja foUolosa H. and A., Berkeley (type
of Uredo coleosporiodes Diet, and Holw.), also Mill Valley (Bethel) :
on C. latifolia H. and A., and C. WigJitii Elmer, Moss Beach (Mrs.
Brandegee) ; on C. miniata Dougl., Lassen National Forest (Meinicke) ;
on C. sp., Lake Tahoe (Bethel) ; on C. Doiiglasii Benth., San Mateo
County (Copeland), on C. Martini Abrams, San Diego (Bethel).

13. Cronartium filamentosum (Peck) Hedgcock. 0, I, II, III

Bot. Gaz., vol. 7, p. 56, 1882; Phytopathology, voL 2, p. 176, 1912.

The aeeial stage produces slight hyperthrophy of branches on
Pinus ponderosa Dougl. {Peridermium filamentosum Peck), Lake
Tahoe; on P. contorta Dougl., Lake Tahoe, Plumas County (Meinicke).

Telia on Castilleja miniata Dougl., and according to Hedgcock not
distinguishable morphologicallj^ from those of Cronartium coleo-
sporioides, Lake Tahoe.

1919] Blasdale: Uredmales of Calif ornm 109

14. Cronartium pyriforme (Peck) Hedg. and Long. 0, I, II, III

Bull. Torr. Club, vol. 6, p. 13, 1875, and vol. 11, p. 50, 1884.

Aecia on Pinus ponderosa Dougl, {Peridermium pyriforme Peck)
Rocky Gulch, Siskiyou County (Meinicke).

Uredinia and telia on Comiandra umbellata (L) Nutt. {Cronartium
Comandrae Peck), Shasta Springs. The genetic connection between
these forms was shown by Hedgcock and Long (Bull. 247, U. S. Dept.
Agr., 1914), also by Kirkwood (Phytopathology, vol. 5, p. 233, 1915).

15. Cronartium Quercus (Brond.) Schrot. 0, I, II, III

Saee. Michelia, vol. 2, p. 308, 1881.

The aecial stage (Peridermium Harknessii Moore) produces
globose galls often of large size on branches of Pinus radiata Don, and
is sometimes very destructive. Also on P. attenuata Lemmon, P. muri-
cata Don and P. ponderosa Dougl. in the central Coast Ranges.

Uredinia and more rarely telia on Quercus agrifolia Nee., Q. Kel-
logii Newb. and Pasania densifiora Oerst., Gilroy, Mount Diablo,
Monterey (Meinicke), Mill Valley (Bethel), Santa Barbara (Bethel) ;
on Q. dumosa Nutt., Pasadena (McClatchie).

There seems to be no essential difference between the eastern and
western forms of this species, although culture experiments are needed
to prove this conclusively. Meinicke (Phytopathology, vol. 6, p. 225,
1916) has shown that aecial spores from Pinus radiata could be made
to reproduce aecia on the same host without an intervening stage;
also that the fungus winters over in the uredinial stage on the leaves
of Quercus agrifolia.

ERIOSPORANGIUM (bertero) lev.
Ann. Sci. Nat., ser. 3, vol. 5, p. 269, 1846.
Aecia usually indefinite, peridium usually fragile. L'redinia
definite, without parapliyses. Teliospores two-celled, usually pale or
colorless.

16. Eriosporangium evadens (Hark.) Arthur. 0, I, II, III

Bull. Calif. Acad. Sci., no. 1, p. 34, 1884; Arthur, Result. Sci. Cong. Bot.
Vienne, p. 343, 1906.

The aecial stage is probably Coleosporium Baccharidis Cooke
(Grevillea, vol. 9, p. 7, 1880), which produces galls often of large size
on the branches of Baccharis consanguinea Greene and B. piliduris
DC, similar to those of the bark-inhabiting species of Peridermium.

110 University of California Piihlication^ in Botany [^ol. 7

TJredinia and telia occur on tlic leaves of the same hosts, but as the
sori are small they are easily overlooked. The genetic connection
between the two forms has not been proven; but the frequency with
which they are associated makes it probable that they are so related.
It is common tliroujjrliout the bay region and in southern California.

17. Eriosporangiumpunctato-striatum (Dietel and Neger) Arthur

Bot. Jahrb., vol. 22, p. 357, 1896; N. A. Flora, vol. 7, p. 213, 1912.

Aecia on stems and leaves on Baccharis viminea DC, but does not
produce conspicuous galls.

Uredinia and telia on the same host (type of Fuccinia BaccJiaridis
Diet, and Holway), Santa Rosa, Los Gatos, Pasadena (McClatchie),
San Bernardino (Parish), Paso Eobles (Bethel), San Ysidro (Bar-
tholomew), Lakeside (Bethel).

GYMNOSPORANGIUM hedw.
Flora frangaise, vol. 2, p. 216, 1805.

Aecia usually cylindrical but rarely cupulate ; peridia membranous,
the cells imbricate. Uredinia with one exception lacking. Telia
naked, gelatinous, and somewhat elastic when mature ; teliospores two-
celled, sometimes three- to five-celled, by transverse septa, pedicels
hyaline, usually long.

18. Gymnosporangium Blasdaleanum (Diet, and Holw.) Kern
I, III

Erythea, vol. 3, p. 77, 1895; Kern, Bull. N. Y. Bot. Gard., vol. 7, p. 437,
1911.

Aecia on Amelanchier alnifolia Nutt. and Crategus rivularis Nutt.,
Sisson and Shasta Springs, type collection; on Amelanchier alnifolia
Nutt., Hoopa Valley, Plumas County (Meinicke), Yosemite Valley
and Lake Tahoe (Bethel), on Amelanchier pallida Greene and Cra-
tegus Doiujlasii Lindl., northern California (Kern.). This has the
well developed peridium of a typical aecidium.

Telia on Lihocedrus decurrens Torr. Sisson, Hoopa Valley, Sis-
kiyou County (Copeland), Potter Vallej^ (Purpus), Yosemite Valley
and Lake Tahoe (Bethel) ; "widely distributed throughout the north-
ern part of the state" (Meinicke). It sometimes produces witches'
brooms of some size, but the spore masses, which appear as brown
cushions on the smaller twigs and leaves, are not conspicuous.

1919] Blasdale: Uredmales of California 111

The genetic connection between the two stages was first established
by Arthur (Mycologia, vol. 1, p. 252, 1909, vol. 4, p. 57, 1912), who
succeeded in growing aecia on Crategus Pringlii Sarg. from telia
obtained from Corvallis, Oregon. From the latter state it is reported
to be a rather serious pest by O'Gara (Science, vol. 39, p. 60, 1914),
and by Jackson (Phytopathology, vol. 4, pp. 41, 261), attacking
especially pears, quinces, and certain varieties of apples.

19. Gymnosporangium Harknessianum (Ell. and Ev.) Kern. I,

III

Bull. N. Y. Bot. Gard., vol. 7, p. 441, 1911.

Aecia on Amelanchier alnifolia Nutt., Klamath River (Harkness),
Plumas County (Meinicke).

What is believed to be the telial stage of this species has been col-
lected by Meinicke on Juniperus occidentalis Hook., near Clio, Plumas
County.

20. Gymnosporangium koreaense (P. Hennings) Jackson. I, III

Jour. Agr. Ees., vol. 5, p. 1006, 1916.

Aecia on leaves of an ornamental species of Pyriis from a nursery
near Oakland (D. P. T. MacDonald), but not otherwise reported from
the state.

Telia not reported from California, but found in Oregon on Juni-
perus chinensis Lindl. (Jackson).

HYALOPSORA magn.

Ber. deut. bot. Ges., vol. 19, p. 582, 1901.

Aecia without peridium not conspicuous, irregularly dehiscent;
aeciospores borne singly on pedicels, colorless. Uredinia with globoid
peridium. Telia scattered, indehiscent; teliospores globoid, usually
four-celled, colorless, wall thin and smooth.

21. Hyalopsora Cheilanthis (Peck) Arthur. II

Bull. Torr. Club, vol. 10, p. 62, 1883; Arthur, N. A. Flora, vol. 7, p. 113,
1907.

On Pellaea a7idror)\edafolia Kaulf., San Gabriel Caiion (Leroy
Abrams) ; on Ceratoptcris triangularis (Kaulf.) Underwood, Pasa-
dena, type of JJredo pasadenae Sydow (McClatchie), Mount Tamal-
pais; Julian, San Diego County (Bethel).

112 University of California Puhlicati&ns in Botany [Vol. 7

22. Hyalopsora laeviscula (Diet, and llolw.) Arthur. II, III

Erythea, vol. 2, p. 127, 1894; Arthur, N. A. Flora, vol. 7, p. 113, 1907.
On Polypodia})! calif orotic urn Kaulf., Mount Tamalpais, type col-
lections.

23. Hyalopsora Polypodii (DC) Magnus. I, II, III

Fl. franc, vol. 6, p. 81, 1815 j Magnus, Ber. deut. bot. Ges., vol. 19, p. 582,
1901.

On FiUx fragilis (L) Underwood, Humboldt Count}^, King's River
Caiion (Holway).

KUEHNEOLA magn.
Bot. Ccntralbl., vol. 7-4, p. 169, 1898.
Uredinia definite, erumpent, usually with paraphrases; uredinio-
spores borne singly on pedicels, usually pale or colorless; telia erum-
pent, without paraphyses; teliospores three- to seven-celled, with
transverse septa.

24. Kuehneola uredinis (Link) Arthur. 0, II, III

Willd. Sp. Plant, vol. 6, p. 123, 1824; Arthur, N. A. Flora, vol. 7, p. 186,
1912.

On Ruhiis vitifolius Cham, and Schlecht., reported by Arthur from
California but not seen by the writer.

KUNKELIA ARTHUR
Bot. Gaz., vol. 63, p. 504, 1917.

This genus is based upon the fact, first shown by Kunkel (Bull.
Torr. Bot. Club, vol. 40, p. 361, 1913, Am. Jour. Bot., vol. 1, p. 37,
1914), that the entire life cycle consists of a caeomoid stage associated
with pycnia, the spores of which germinate like teliospores. It is not
distinguishable morphologically from the aecial stage of the genus
Gymnoconia, but the aecial spores of the genus last named germinate
like true aecia spores, and there is an alternate stage not distinguish-
able from the telia of the genus Puccinia.

25. Kunkelia nitens (Schwein) Arthur. 0. I
Bot. Gaz., vol. 63, p. 504, 1917.

On Ruhvs vitifolius Cham, and Schl., Santa Rosa, on cultivated
blackberry. Orange, Berkeley, Chico (Copeland), Glendora (Baker),
Long Beach (Bethel). This is the widely distributed "orange rust"
of the lilackberry formerh- known as Caeo^na nitens.

1919] Blasdale: Uredinales of California 113

26. Kunkelia Rosae-gymnocarpae (Dietel) Arthur

Hedwigia, vol. 44, p. 334, 1905; Arthur, Bot. Gaz., vol. 42, p. 505, 1917.
On Rosa gymnocarpa Nutt., Amador County, type collection (Han-
sen), Plumas County (Meinicke), Paso Robles and Yosemite Valley
(Bethel). Very little is known about the life history of this rust, but
its similarity to the preceding species makes it probable that it is also
a short-cycle form. It appears very early in the spring and produces
hypertrophy and ultimate death of the affected shoots.

MELAMPSORA cast.
Observ., vol. 2, p. 18, 1843.
Aecia erumpent, peridia and pycnia wanting. Aeciospores globose
with colorless walls. Uredinia erumpent, pulverulent, borne singly on
pedicels. Telia in waxy layers, indehiscent; teliospores one-celled,
prismatic or ellipsoid.

27. Melampsora albertensis Arthur. 0, I, II, III

Bull. Torr. Club, vol. 33, p. 517, 1906.

Aecia not reported from California but found in British Columbia
and Colorado on Pseudotsuga niucronata (Raf.) Sudw. {Ca&oma
occidentalis Arthur).

Uredinia and telia on Populiis tremuloides Michx., Dunsmuir (E. L.
Smith) ; on P. trichocarpa T. and G., Yosemite Valley (Bethel). The
genetic connection between these forms was established by Arthur
(Mycologia, vol. 4, pp. 29, 58, 1912).

28. Melampsora Bigelowii Thiim. 0, I, II, III

Mitth. Forsty. Vers. Oest., vol. 2, p. 37, 1879.

The aecial stage {Caeonia Bigelowii Thum. Arth.) not reported
from California but found elsewhere on Larix Lyalii Pari.

Uredinia and telia on Salix lasiandra Benth. and S. lasiolepis
Benth., Berkeley; on 8. laevigata Bebb and 8. exigua Nutt., Long
Beach (Bethel). Extremely common and widely distributed.

29. Melampsora Lini (Pers.) Desmaz. 0. I, II, III

Syn. Fungi, p. 216, 1901; Desmaz, PI. Crypt., fasc. 41, no. 2049, 1850.
On Linnm micranthiim Graj^ Sisson; on L. digynum Gray,
Amador County (Hansen). Also reported by Arthur (N. A. Flora,
vol. 7, p. 102) ; on L. congesUim Gray, L. Leivisii Pursh. ; L. Brewcri
Gray, and L. drymarioides Curran. This species was shown to be
autoecious by Arthur (Jour, of Mycol., vol. 13, p. 207, 1907).

114 University of Calif oniia PuhUcaticns in Botany [Vol.7

30. Melampsora occidentalis Jackson. II, 111

Pliytoi)atliolog5', vol. 7, p. 354, 1917.
On Fop III US trichocarpa T. and G., Siskiyou, Trinity, and Shasta
counties (Meinicke) ; on P. Frcmontii Wats., San Bernardino (Par-
ish). These specimens were formerly referred to Melampsora Medusae
Thiini., from which the species named above differs in a number of
minor characters. Still another specimen collected b}^ Bethel on
Populus alha L. at San Jose, which has also been referred to M.
Medusae, has not been satisfactorily named.

MELAMPSOROPSIS (schroet) abthub
Result. Sei. Coug. Bot. Vienne, p. 338, 1908.
Aecia erumpent but laterally flattened ; peridium firm. Aecio-
spores ellipsoid to globoid, walls colorless. Uredinia erumpent, pul-
verulent. Telia waxy; teliospores in chains, one-celled, oblong or
cuboid, walls colorless.

31. Melampsoropsis Piperiana Arthur. II

N. A. Flora, vol. 7, p. 120, 1907.
On Khododendron calif ornicum Hook, Humboldt County.

32. Melampsoropsis Pyrolae (DC) Arthur. 0, I, II, III

Fl. franc, vol. (5, p. 99, 1815; Arthur, Result. Sci. Cong. Bot. Vienne,
p. 338, 1906.

The aecial stage not rejDorted from California but collected by
Hedgcock on Picea Engelmanii Parry. The genetic connection of this
species with Peridernmmi conorum Piceae was shown by Fraser
(Mycologia, vol. 4, p. 183, 1912).

On Pyrola uliginosa Torr., Sisson (Holway).

NEORAVENELIA long
Bot. Gaz., vol. 35, p. 131, 1903.

Aecia erumpent, without peridium ; aeciospores in chains, wall
colored. Uredinia erumpent ; urediniospores borne singly on pedicels,
wall colored. Telia erumpent; teliospores like those of Ravenelia.

33. Neoravenelia Holwayi (Dietel) Long. 0, I, II, III

Hedwigia, vol. 33, p. 61, 1894; Bot. Gaz., vol. 35, p. 131, 1903.

On Prosopis jidi flora (Sw.) DC, San Bernardino County (Mc-
Clatchie).

1919] Blasdale: Uredinales of California il5

NYSSOPSORA AKTH.
Eesult. Sci. Cong. Bot. Vienne, p. 342, 1906.
Telia erumpent; teliospores three-celled by oblique partitions, lat-
erally flattened ; walls deeply colored, spinous.

34. Nyssopsora echinata (Lev.) Arth. Ill

Ann. Sei. Nat., ser. 3, Bot., vol. 9, p. 247, 1848; Arthur, op. cit.

On Oenanthe calif ornica Wats., San Francisco, Santa Cruz (Thom-
son) ; on Selinum paciftcum Wats., Santa Rosa and Hoopa Valley.

PERIDERMIUM

Under this form genus are grouped those aecial forms for which
no other stages are known and which resemble the aecia of Coleo-
sporium and CronarUum.

35. Peridermium californicum Arth. and Kern
Mucologia, vol. 6, p. 118, 1914.

On Pinns radiaia Don., Monterey (Bethel). This is believed to be
the aecial stage of Coleosporium Madiae by Meinicke, but the data
upon which this conclusion is based has not yet been published.

36. Peridermium Ephedras Cooke

Indian Forester, vol. 3, p. 95, 1877; Arthur and Kern, Bull. Torr. Club,
vol. 33, p. 420, 1906.

On Ephedra calif ornica Wats., San Diego (Bethel), Riverside
County (Parish). Conspicuous on account of its large, orange colored
spore masses; see Standley (Plant World, vol. 13, p. 61, 1909).

PHRAGMIDIUM likk.
Ges. nat. Freunde, Berlin, vol. 7, p. 30, 1915.
Aecia erumpent, usually indefinite and surrounded by paraphyses.
Aeciospores in chains, globoid, with colorless walls. Uredinia erum-
pent, usually with paraphyses ; urediniospores borne singly on pedicles.
Telia erumpent, usually definite and without paraphyses; teliospores
two-celled or more by transverse septa with two or more lateral pores.

37. Phragmidium Andersoni Shear. I, II, III

Bull. Torr. Club, vol. 29, p. 453, 1902.
Qn Potentilla fruticosa L., Mount Eddy, Siskiyou County (Cope-
land).

116 Umversitij of California Puhlimtions in Botany [Vol.7

38. Phragmidium disciflorum (Tock-j James. 0, I, 11, III

Funyi Mt'ikl., vol. 1, p. 16, 1790; James. Contr. U. S. Nat. Herb., vol. 3,
p. 276, 1895.
On cultivated roses of the hybrid perpetual class, common through-
out the state.

39. Phragmidium imitans Arthur. I, II, III

N. A. Flora, vol. 7, p. 165, 1912.
On Rnhiis leucodermis Dough, Sisson, Shasta Springs, Yosemite
Valley and Rionido; on cultivated varieties of raspberry rather
frequent and sometimes destructive.

40. Phrag"midium Ivesiae Sydow. I, IT. Ill

Auu. Mycologici, vol. 1, p. 329, 1903.
On PotentUla Blaschkeana Turez., Sisson, Lake Tahoe, Lassen
County (Davy) ; on P. glandulosa Lindl., Modoc County (Meinicke) ;
on P. Hallii Rydb., Lake Tahoe (Holway) ; on P. Elmeri Rydb., Don-
ner Lake (Heller).

41. Phragmidium montivagum Arth. 0, I, II, III
Torreya, vol. 9, p. 24, 1909.

On Rosa gymnocarpa Nutt., Pine Ridge, Fresno County (Hall and

Chandler).

42. Phragmidium occidentale Arthur. 0, I, II, III

Greene, Plantae Bakerianae, vol. 2, p. 3, 1901.
On Ruhiis parviflorus Nutt., Shasta Springs, Los Gatos, Berkeley,
Mount Tamalpais, Rionido, Yosemite Valley, and Lake Tahoe.

43. Phragmidium Rosae-californicae Diet. I, II, III

Hedwigia, vol. 44, pp. 125, 333, 1904.
On Rosa californica Cham, and Schl., R. gymnocarpa Nutt., R.
sononiensis Greene, and R. minutifolia Englm. Of frequent occur-
rence throughout the Coast Ranges. The aecia produce hypertrophy
of the young shoots early in the season ; later they appear on the
mature leaves, which they injure but little.

PILEOLARIA CAST.
Obs., vol. 1, p. 22, 1842.
Uredinia erumpent, with paraphyses; urediniospores borne singh'-
on pedicels, walls colored. Telia erumpent, definite ; teliospores borne
on pedicels, one-celled, flattened laterally, wall deeplj^ colored, very
verrucose.

1919] Blasdale: Uredinales of California 117

44. Pileolaria Toxicodendri (Berk, and Rav.) Arthur. 0, II, III

Grevillea, vol. 3, p. 56, 1874; Arthur, N. A. Flora, vol. 7, p. 147, 1907.
On Rhus diversiloha T. and G., common in the Coast Ranges, Cata-
lina Island (Bartholomew).

POLYTHELIS abthur

Result. Sci. Cong. Bot. Vienne, p. 341, 1906.
Telia erumpent, definite, without peridium. Teliospores forming
heads attached by fragile pedicels to a common stalk, which is incon-
spicuous; two-celled by a transverse septum, easily separating at the
septum.

45. Polythelis Thalictri (Chev.) Arthur. 0, III

Chev., Fl. Paris., vol. 1, p. 417, 1826; Arthur, Result. Sci. Cong. Bot.
Vienne, p. 341, 1906.

On Thalictrum Fendleri Englm., Lake Tahoe.

PUCCINIA PERS.
Sp. PI., vol. 6, p. 67, 1825.
Aecia erumpent, cupulate, or c^dindrical; aeciospores in chains,
globoid or ellipsoid, wall colorless. Uredinia erumpent definite, with-
out peridium; urediniospores borne on pedicels, wall colored, echinu-
late or verrucose. Telia erumpent, sometimes long covered by the
epidermis; teliospores two-celled; wall colored, with a single pore to
each cell.

Telia on Graminaceae

46. Puccinia abundans (Pk.) Jackson. I, III

Mem. Brooklyn Bot. Card., vol. 1, p. 229, 1918.
The aecial stage of this species {Aecidium abundans Pk.) is occasion-
ally found on Symphoricarpos racemosus Michx., Berkeley, Santa
Rosa.

Telia not definitely reported from California but to be expected
on species of Festuca.

47. Puccinia adspersa Diet, and Holw. II, III

Erythea, vol. 3, p. 81, 1895.
On unknown grass, Modoc County (F. P. Nutting). This species
seems to differ but little if at all from Puccinia Clenmtidis (DC)
Lagerh.

118 Vniversitij of California Piiblications in Botany [Vol. 7

48. Puccinia anomala Rostr. I, J 1 , 1 H

Thiim, Mye. Univ., uo. 831.
Aecia, not reported from California, but in Russia on species of
Ornithogaliim (INIycol. Cent., vol. 4, p. 70, 1914).

Uredinia and telia on Ilordcum vulgare L., Davis (L. R. Jones).

49. Puccinia Aristidae Tracy. II, III
Jour, of Mycol., vol. 9, p. 281, 1893.

On Aristida bromoides, "Waterman's Hot Springs, San Bernardino
County (Parish).

50. Puccinia Hordei Fuckel. II, III

Jahrb. Ver. Nat. Nassau, vol. 15, p. 16, 1860.
Uredinia and telia on Horde um murinum L., Berkeley, San Diego,
and Long Beach (Bethel) ; on H. Gussonianum Pari., Los Angeles
(Bethel).

51. Puccinia Clematidis DC, Lagerh. I, II, III

Troniso Mus. Parsh., vol. 17, p. 47, 1895.

Aecia on Clematis lasiantha Nutt. and C. ligusticifolia Nutt. {Aeci-
dium Clematidis DC), Berkeley, Atlas, Napa County, and Sisson,
Mount Wilson (Bethel) ; on Aquilegia trunoata F. and M. {Aecidium
Aquilegiae Pers.), Berkeley, Acorn, Humboldt County; on Clematis
pauciflora Nutt., San Diego County (LeRoy Abrams) ; on TJialictrum
Fendleri, Lake Tahoe.

Uredinia and telia on Bromus villosus Forst., Berkeley, Ukiah,
Ontario, San Bernardino, Long Beach (Bethel) ; on B. hordaceus L.,
San Diego (Bethel) ; on B. carinatus H. and A., Catalina Island
(Bethel) ; on B. marginatus Nee., Willits (Hitchcock) ; on Elymus
triticoides Buckl. Catalina Island (Bethel) ; on E. condensatus Presl.
(Bethel); on E. glaucus Buckl., Campbell (H. B. Humphrey), and
near Mount Shasta (E. Palmer).

52. Puccinia Cynodontis Desm. II, III

Exsiccati, vol. 3, no. 655.

Uredinia on Cynodon dactylon Pers., Sacramento, Anaheim; "very
common throughout southern California" (Bethel). Telia produced
only during the winter season.

1919] Blasdale: Uredinales of California 119

53. Puccinia Epicampis Arthur. II, III

Bull. Torr. Bot. Club, vol. 33, p. 662, 1901.
On Epicanipes ringcns Benth., San Diego (Bethel).

54. Puccinia Fendleri (Tracy and Earle) Jackson. I, III

Mem. Brooklyn Bot. Gard., vol. 1, p. 246, 1918.

Aecia {Aecidium Fendleri Tracy and Earle) on Berheris pinnata
Lag., Berkeley. Collected but once and but poorly developed. This
was formerly taken to be the aecial stage of Uropyxis sanguineu
(Erythea, vol. 3, p. 131, 1895) and its specific identity is still an open
question.

Telia not reported from California but to be expected on species of
Koeleria.

55. Puccinia glumarum (Schmidt) Erikss. and Ilenn. II, III

Zeitschr. f. Pflanzenkr., vol. 4, p. 1917, 1894.
On Hordeum murinum L., near Tehachipi (A. G. Johnson). This
is one of the most destructive grain rusts in Europe but apparently
it is not common in California. See Humphrey and Johnson, Phj^to-
path., vol. 6, p. 80, 1916.

56. Puccinia graminella (Spez.) Diet, and Holw. I, III

Erythea, vol. 3, p. 80, 1895.
On Stipu eminens Cav., Berkeley. This is the only rust which
produces its aecia on a grass ; the aecia are easily overlooked.

57. Puccinia Holcina Erikss. II, III

Ann. Sci. Nat., ser. 8, vol. 9, p. 274, 1899.
Uredinia on Notholcus lanahis (L) Nash, extremely common in
the bay region, Humboldt County, southern California (Bethel).

58. Puccinia interveniens (Pk.) Bethel comb. nov. 0, I, III

Pk. Bull. Torr. Bot. Club, vol. 29, p. 74, 1883.
Aecia (Aecidium roestilioides Ell. and Ev.) on Sidalcea malvaeflora
(Moc. and Sesse) Gray, Oakland Hills, Mount Tamalpais, San Fran-
cisco, Pasadena (McClatchie), San Diego, Monterey, and Julian
(Bethel) ; on 8. asprella Greene, Butte County (Heller) ; on Mal-
vastrum Thurheri Gray, San Diego and San Jacinto (Bethel) ; on
M. amhigua, Victorville and Granite Mountain, near Julian (Bethel).

120 University of Calif ornia Publications in Botany [Vol.7

Telia {Puccinia Burnettii Griff.) an Stipa pulchra Hitch., Santa
Cniz (Griffiths) ; on S. speciosa, Victorville and Granite Mountain
(Bethel) ; on 8. sp., Mill Valley and San Diego (Bethel) ; on Oryzopsis
hyuienoidcs Ricker, Victorville (Bethel).

The connection between these forms has been established by Bethel
by repeated cultures with both telia and aecia, the first being made
in Colorado in 1912. His numerous collections of both aecia and telia
in 1916 and 1918 in southern California, not far from the type locality
of Roestelia interveniens establish the validity of the combination
given above. The aecia have xqyj short peridia, split almost to the
base, as in the form genus Roestilia; the telia form sori from one to
three inches in length, especially on the sheaths, which are tardy in
rupturing.

59. Puccinia luxuriosa Sydow. II, III

Monogr. Uml., vol. 1, p. 812, 1904.
On Sporoholus airoides Torr., Lakeside, Riverside County (Bethel).

60. Puccinia Majanthae (Sehrum.) Arth. and Holw. 0, I, II, III

Bull. Lab. Nat. Hist. Iowa, vol. 5, p. 188, 1901.

Aecia not reported from California but elsewhere on species of
Polygonatuni and Convallaria.

Uredinia and telia on Phalaris minor Retx., Long Beach (Bethel)
and Palo Alto (Thomson) ; on P. calif oryiica H. and A., Montara
Point, San Mateo County (Copeland).

61. Puccinia montanensis Ellis. I, II, III

Jour, of Mycol., vol. 7, p. 274, 1883.

Aecia on IlydropJiyllum capitatmn Dougl. and H. occidentale Gray
{Aecidiuni Hydrophyllae Peck.), Humboldt and Mendocino counties,
Amador County (Hansen) ; on Phacelia calif ornica Cham, and P.
nemoralis Greene {Aecidinm Phaceliae Peck), Berkeley; on Phacelia
tanacetifolia Benth., Carmel.

Telia on Hordenm nodosum L., Catalina Island (Bethel),

62. Puccinia Poarum Niels. II, III

Bot. Tids., vol. 2, p. 26, 1877.

Uredinia on Poa annua L., and P. pratensis L., Berkeley, Long
Beach (Bethel).

Telia not reported from California.

1919] Blasdale: Uredmales of California 121

63. Puccinia poculiformis (Jacq.) Wettst.

Verhl. Zool. Ges. A^ienue, vol. 35, p. 544, 1885.

Aecia on species of Berberis, but not reported from California.

Uredinia and telia on Avena fatua L., A. harhata Brot., and A.
sativa L. ; on Hordeum murinum L., H. Gusso-nianum Pari, and H.
sativum Jessen; on Phleum pratense L. ; on Agrostis alha L. and A.
diegoensis Yas. ; on Elymus condensatus Presl. ; on Lamarckiu aurea
Moench; on Dactylis glomerata L. ; on Panicularia pauciflora (Presl.)
Kze. Frequent throughout the state. This is the "black stem rust"
found especially on the culms of oats, and in some seasons decidedly
destructive. It includes several well defined biological forms, which
are discussed by Stakeman and Piemeisel (Phytopathology, vol. 6,
p. 99, 1918).

61. Puccinia procera Diet, and Holw. II, III

Erythea, vol. 1, p. 249, 1893.
On Elymiis condensatus Presl., Pasadena (McClatchie), Berkeley,
Long Beach, and San Diego (Bethel).

65. Puccinia purpurea Cke. II, III

Grevillea, vol. 5, p. 15, 1879.
On Andropogon sorghum var. halep&nsis L., Gilroy, Santa Ana
(Bethel).

66. Puccinia Rhamni (Pers.) Wettst. I, II, III

Ver. zool.-bot. Ges. Wien, p. 544, 1885.

Aecia not reported from California, but to be looked for on species
of Rhamnus.

Uredinia and telia on Polypogon monspeliensis Desf., Berkeley,
Humboldt County, Ontario, Santa Barbara ; on Avena harlata Brot.,
San Diego (Bethel) ; on Lamarckia aurea Moench, Los Angeles
(Bethel) on Lolium perenne L., and L. nmltiflorum Lam., Long Beach
(Bethel).

67. Puccinia sejuncta Sydow. 0, I, II, III

Ann. Mycologici, vol. 1, p. 326, 1903.
Aecia on Hieracium sp., Sisson (Holway).

68. Puccinia Sorghi Schw. II, III

Trans. Am. Philos. Soc, ser. 2, vol. 4, p. 295, 1832.
On Zea mays L., frequent but not destructive.

122 University of California Puhlications in Botany [Vol. 7

69. Puccinia Stipae (Opiz.) Arth. 0, I, II, III

Bull. Iowa Agri. Coll., p. 160, 1884,

Aecia not definitely reported from California, but this stage is
known to occur on nine different cliicoriaceous and carduaceous genera
(Bethel, Phytopathology, vol. 6, p. 99, 1916).

Uredinia and telia on Stipa setigera Presl., Berkeley; on S. pulchra
Hitch., San Diego, Coronado Island, and Long Beach (Bethel) ; on
8. eminens Andersoni Vassey, San Diego (Bethel) ; on Oryzopsis
hymenoides Ricker, Victorville (Bethel).

70. Puccinia subnitens Dietel. I, III

Erythea, vol. 3, p. 81, 1895.

Aecia on Spcrgidaria Clevelandii (Greene) Robinson, Long Beach;
on 8. macrotheca (Hornem) Heynh ; on Thely podium laciniatum
Endl. and Heliotr opium curassavicwn L., Lassen County (Davy) ; on
Ahronia pinetorum Jepson, San Jacinto (Bethel) ; on Isomeris arhorea
Nutt., San Diego (Jones). Shown by Bethel (Phytopathology, vol. 7,
p. 92, 1917) to develop on eighty-five different species of host plants,
representing fift^'-two genera and nineteen families.

Telia on Distichlis spicata Greene, Redwood City, "common from
Los Angeles to San Diego," San Jacinto (Bethel).

71. Puccinia Triticina Erikss. II, III

Ann. Sci. Nat., ser. 8, Bot., vol. 9, p. 270, 1899.
Frequent on certain varieties of wheat, Berkeley, San Francisco,
Anaheim, Long Beach (Bethel). It is closely related to P. Clefriatidis.

Telia on Cyperaceae

72. Puccinia Asterum (Schw.) Kern. I, II, III

Mycologia, vol. 9, p. 224, 1917.
Aecia on Aster sp.. Lake Tahoe and Calaveras Co., Seabright (]\Irs.
Clemens) ; on A. occidentaUs, Yosemite Vallej^ (Bethel),

Telia on Carex sp.. Lake Tahoe, Seabright (Mrs. demons).

73. Puccinia atro-fusca (Dudley and Thomson) Ilohvay. II, III
Jour, of Mycol., vol. 10, pp. 55, 228, 1904.

On Carex Douglasii Bott. and C. usta Bailey, San Bernardino
County (Dudle}^ and Thompson). This species is peculiar in that it
produces large numbers of amphispores, and honeo was first placed
in the genus Uromyces.

1919] ^ Blasdale: Vredmales of California 123

74. Puccinia canaliculata (Schw.) Lagerh. 0, I, II, III

Tromso Mus. Aarsh., vol. 17, p. 51, 1894.

Aecia on Xanthium Canadense Mill, Lakeside (Bethel).
Telia on Cypertis esculentus L., Long Beach and San Bernardino
(Bethel).

75. Puccinia Caricis (Schum.) Eebent. 0, I, II, III

Fl. neomarch, p. 356, 1804.

Aecia on Urtica gracilis holosericea Jepson, Berkeley.
Uredinia and telia on Carex ohnupta Bailey, Berkeley.

76. Puccinia Eleocharidis Arthur. Ill

Prelim. List Iowa Ured., p. 158, 1884.

On EleocJiaris montana (H. B. K.) R. and S., Long Beach
(Bethel).

77. Puccinia Grossulariae (Schum.) Lager. 0, I, II, III

Aecia on Ribes divaricatum Dougl., Berkeley, San Francisco,
Sisson.

Telia not reported in California but probably present on species
of Carex.

78. Puccinia McClatchieana Diet, and Holw. II, III

Erythea, vol. 2, p. 127, 1895.

On Scirpus microcarpus Presl., San Francisco, Pasadena (Mc-
Clatchie).

79. Puccinia obtecta Peck. II, III

Bull. Buffalo Soc. Nat. Hist., vol. 1, p. 66, 1873.

On Scirpus lacustris occidentalis Wats., Sacramento; on S. cali-
fornicus (Meyer) Britton and S. Olneyi Gray, Long Beach (Bethel) ;
on 8. americanus Pers., Escondido (Bartholemew).

80. Puccinia patruelis Arthur. I, II, III

Mycologia, vol. 1, p. 245, 1909.

Telia not reported in California, but probably present on species
of Agoseris and Crepis.

Uredinia and telia on Carex niarcida Boott., San Bernardino
(Bethel and Parish),

124: TJrviversity of California Puhlicatians in Botany [Vol.7

81. Puccinia Peckii (De Toni) Kell. 1, 111

Jour, of Mycol., vol. 8, p. 20, 1902.

Aecia on Onagra Ilookcri (T. and G.) Small, Yosemite Valley,
Giant Forest, King's River Caiion (Holway), San Bernardino
(Bethel).

Uredinia and telia on Carex siccata Dewey, Yosemite Valley
(Bethel).

Telia on Liliaceae

82. Puccinia Alliorum (DC) Corda. I, II, III

Fl. frauc, vol. 6, p. 82, 1815; Corda leones, vol. 4, p. 12, 1840.

On Allium serratum Wats, type of Puccinia Blasdalei Diet, and
Holw., Antioch and Mount Diablo; on A. falcifolium II. and A.,
Benicia (Bigelow).

Unlike the European P. Alliorum I and III are produced on the
same plants.

83. Puccinia Asparagi DC. 0, I, II, III

Fl. franc, vol. 2, p. .595, 1805.
On Asparagus officinalis L. Common throughout the state, and
the source of large losses to growers. The life history and methods
of combating it have been exhaustively studied by Professor R. E.
Smith, Calif. Agr. Exp. Sta., Bulls. 165, 172.

84. Puccinia Calochorti Peck. I, III

Bot. Gaz., vol. 6, p. 228, 1881.
On Calochortiis alhns Dougl., Hoopa Valley; on C. vemistus
Benth., C. Maweanus Leichtl., Plumas County (Home) ; on C. hiscep-
trum Wats., King's River Canon, type of Puccinia Holway i Diet.
(Holway) ; on C. elegans Pursh., type of Puccinia anachorita Ell. and
Hark. (Harkness).

85. Puccinia granulispora Ell. and Gall. I, II. HI

Bull. Torn Club, vol. 22, p. 61, 1895.
On Allium precox Brandcgee, San Bernardino (Parish).

86. Puccinia mesomegala lierk. and Cooke. Ill

25th Ann. Rept. N. Y. State Mus., p. Ill, 1873.
On Clintonia uniflora (Menzies) Knuth, Amador County (Han-
sen), and Siskiyou County (Meinicke).

1919] BlasdaJe: Uredinales of Calif or nia 125

87. Puccinia Moreniana Dudley and Thomson. Ill
Jour, of Mycol., vol. 10, p. 53, 1904.

On Brodiaea capitata Benth., Searsville Lake, San Mateo County
(Dudley and Thompson).

88. Puccinia nodosa Ell. and Hark. II, III

Bull. Calif. Acad. Sci., vol. 1, p. 27, 1884.
On Brodiaea capitata Benth., Berkeley, Ukiali.

89. Puccinia Veratri Duby. I, II, III

Bot. Gall., vol. 2, p. 890, 1830.

Aecia not definitely reported from California, but known to
develop on species of Epilohium.

Uredinia and telia on Vcratrum caUfornicum Durand, Placer
County, Lake Tahoe, Yosemite Valley (Bethel), Modoc County (F. P.
Nutting).

Telia on Iridaceae

90. Puccinia Iridis (DC) Wallr. II, III

Eneyc, vol. 8, p. 224, 1808; Rabh. Krypt. Flora, vol. 1, p. 23, 1844.
Uredinia on Iris longipetala Herb., /. Douglasiana Herb., /. mis-
souriensis Nutt., common in the Coast Ranges. Occasionally found on
certain cultivated varieties of /. germanica, I. xiphium L., I. al-ata
Poir., and I. pumila. Telia ver}^ rare.

Telia ;>n Santalaceae
9L Puccinia Comandrae Peck. Ill

Bull. Torr. Club, vol. 11, p. 49, 1884.
0:^ Comandra umbellata (L) Nutt., Shasta Springs, Fresno County
(Holway).

Telia on Aristolochiaceae

92. Puccinia Asarina Kze. Ill

Kunze and Schmidt, Mykol., vol. 1, p. 70, 1817.
On Asarum Lemmoni Wats., Sisson, King's River Canon (Hol-
way) ; on A. caudatum Lindl., Olema.

93. Puccinia Cynanchi Lagerh. Ill

Biol. Soc. Brot., p. 129, 1889.
On Funastrum hirtellum (Gray) Schltr., Palm Springs (Parish).

126 University of California Piibli cations in Botany ["Vol. 7

Telia on Polygonaceae

94. Puccinia Acetosae (Schum.) Koern. 11, 111

Hechv., vol. 15, p. 184, 1876. .
On Runiex occidentale Wats., and R. acetosella L., Berkeley; on
R. persicarioides L., Long Beach (Bethel).

95. Puccinia amphispilusa Diet, and Holw. II, III

Erythea, vol. 3, p. 79, 1895.
On Polygonum Davisiae Brewer, Lake Tahoe; on P. Newherryi,
Lassen County, type collection (F. P. Nutting).

96. Puccinia Bistortae (Str.) DC. II, III

n. franc, vol. 6, p. 61, 1815.

On Polygonum histortaides (Pursh.) Small, Sherwood, Mendocino
County, King's River Caiion (Holway).

97. Puccinia Polygoni-amphibii Pers. 0, I, II, III

Syn. Method. Fung., p. 227, 1801.

Aecia not reported from California but found in Europe on species
of Geranium. Uredinia and telia on Polygonum Muhlenlyergii Wats.,
San Francisco, Los Gatos, Victorville (Bethel) ; on P. lapathifolium
L., San Bernardino (Parish) ; on P. acre H. B. K., Berkelej^, San
Mateo Countj^ (Baker) ; on P. amphibium L., Palo Alto (Scherfee) ;
on P. hydropiperoides Michx., Los Angeles County (LeRoy Abrams).

98. Puccinia punctiformis Diet, and Holw. II, III

Erythea, vol. 2, p. 128, 1894.

On Rumex salicifolius Weinn., Berkeley, tj'pe collection ; on R.
hym.enosepalus Torr., Berkeley, in cultivation.

Telia on Chenopodiaceae
99. Puccinia Dondiae Arthur. II, III

Bull. Torr. Bot. Club, vol. 42, p. 592, 1915.

On Dondiu intermedia (Wats.) Heller, San Diego (IMarcus E.
Jones).

1919] Blasdale: Uredinales of California 127

Telia on Cruciferae

100. Puccinia Holboellii (Hornem) Rostr. Ill

Fl. Danica, vol. 37, p. 11, 1840; Eostr. Fungi Groen., p. 34, 1886.
On Arabis HolhoeUi Hornem, Yosemite Valley, Pasadena, type of
Puccinia palefaciens Diet, and Holw. (McClatchie), San Diego County
(Bethel) ; on A. Breweri Wats., Mount Diablo (Bioletti) ; on A. arcuta
Gray, King's River Canon (Holway) ; on A. pere^mans Wats., San
Bernardino (Parish).

Telia on Ranunculaceae

101. Puccinia Delphinii Diet, and Holw. Ill

Hedwigia, vol. 32, p. 29, 1893.
On Delphinium sp.. King's River Canon (Holway).

102. Puccinia gemella Diet, and Holw. Ill

Sydow, Monogr. Ured., vol. 1, p. 541, 1903.
On Caltha Howellii Greene, reported from California by Holway
(N. A. Ured., vol. 1, p. 6).

Telia on Saxifragaceae

103. Puccinia Heucherae (Schw.) Diet. Ill

Ber. deut. bot. Ges., vol. 9, p. 42, 1892.
On Heuchera micrantha Dougl. {Puccinia congregata Hark.), fre-
quent in the Coast Ranges; on Tellima grandiflora (Pursh.) Dougl.,
Shasta Springs; on Tellima affinis, Mount Tamalpais (R. S. Gray).

Telia on Crassulaceae

104. Puccinia Rhodiolae B. and Br. HI

Ann. Mag. Nat. Hist., ser. 2, vol. 5, p. 452, 1850.
On Sediwi sp.. King's River Caiion (Holway).

Telia on Rhamnaceae

105. Puccinia Mesnieriana Thiim. Ill

Myeotheca Universalis, vol. 9, p. 834, 1877.
On RJiamnus crocea Nutt., Ukiah, Napa County, Pasadena (Mc-
Clatchie) ; Fresno County (Holway), San Diego (Parish), Santa
Barbara, and Yosemite Valley (Bethel), Mount Tamalpais (Hark-
ness), type of Puccinia digitata Ell. and Hark.; on R. ilicifolius
Kellogg, Mount Diablo; on B. imularis Kellogg, Claremont (Baker).

128 University of California- Publications in Botany [Vol. 7

Telia on Malvaceae

106. Puccinia lobata B. and C. Ill

Grevillca, vol. 3, p. 54, 1874.

On Sida hederacea (Dougl.) Torr., Holtville, Imperial County,
Long Boach (Bethel),

107. Puccinia Malvacearum Bertero. Ill

Gay, Hist, de Cliille, vol. 8, p. 43, 1852.
On Malva horealis Wallm., 31. parviflora L. and Althea rosea Cav.,
common throughout the state; on Sida hederacea (Dougl.) Torr., Sui-
sun (Dav}'). At the Botanic Garden of the University of California
has been found on Malva moschata Linn., M. sylvestris L., M. crispa
L., M. oxyloha Boiss., Lavatera maritima Gouan, L. arhorea L., L.
sylvestris Brot., L. pleviea Sims, and L. assurgentifolia Kell.

108. Puccinia Sherardiana Koern. Ill

Hedwigia, voL 16, p. 19, 1877.
On Malvastrum Thurheri Gray, Fresno County (Holway), San
Diego, and San Jacinto (Bethel) ; on M. splendidum Kellogg, Clare-
mont (Baker) ; on Sidalcea spicata Greene, Lake Tahoe (Bethel).

109. Puccinia Sphaeralceae Ell. and Ev. I, III

Am. Naturalist, p. 428, 1879.
On Sidalcea sp., Humboldt County.

Telia on Violaceae
no. Puccinia effusa Diet, and Hohv. 0, I, III

Erythea, vol. 3, p. 81, 1895.
On Viola lohata Benth., Dunsmuir (Holway).

111. Puccinia Violae (Schum.) DC. 0, I, II, III

PI. Saell., vol. 3, p. 224, 1803; FL franc, vol. 6, p. 62, 1915.
On Viola nephrophylla Greene and V. adunca longipes Sisson ; on
V. fjlahella Nutt., Sherwood ; on V. ocellata T. and G., Ukiah.

Telia on Onagraceae

112. Puccinia Circaeae Pers. HI

Tent. Disp. Moth. Fung., p. 39, 1797.

On Circaea pacifica Aschers and Magnus, Sisson and Weitchpee,
Trinity County.

l^^9] Blasdale: TJredinales of California 129

113. Puccinia Epilobii-tetragoni (DC) Wint. 0, I, II, III

Eabh. Krypt. Flora, vol. 1, p. 214, 188-i.
On Epilohium franciscanum Barbey, San Francisco, and Olema;
on E. paniculatum Nutt., Rionido and Los Gatos, 'Pasadena (Mc-
Clatchie), San Mateo County (Thomson); on E. sp., King's River
Caiion, type of Puccinia intermedia Diet, and Holw.

114. Puccinia Gayophyti Billings. I, II, III

Bot. Gaz., vol. 7, p. 56, 1882.
On GayopJujtum diffusum. T. and G., Sisson, Calaveras Big Trees
(Dudley) ; on G. pumilum Wats., Mount Eddy (Heller) ; on G. ramos-
si'Ssimum T. and G. (Harkness).

115. Puccinia heterantha Ell. and Ev. 0, I, II, III

Erythea, vol. 1, p. 204, 1893.

On Taraxia ovata (Nutt.) Small, Berkelej^ and Atlas, Napa
County; on T. graciliflora (H. and A.) Small (A. Kellogg and W. G.
W. Harford). The aecia appear very early in the spring and cause
hypertrophy and ultimate destruction of the first leaves. The uredinia
and telia appear later and are less destructive. The genetic relation
between the two stages was shown by the writer (Rep. Calif. Exp.
Sta., p. 227, 1894).

116. Puccinia Ludwigiae (Ell. and Ev.) Holw. 0, I, HI

Proc. Phil. Acad. Sci., p. 153, 1893; Holway, N. A. Ured., vol. 1, p. 72,
1907.

On Ludwigia natans Ellis, San Bernardino (Parish).

117. Puccinia Oenotherae Vize. II, HI

Grevillea, vol. 5, p. 109, 1877.
On Clarkia elegans Dougl., C. concinna (F. and M.) Greene, and
C. rhomhoidea Dougl., type of Puccinia Clarkiae Peck. On Godetm
ruMcunda Lindl., G. amoena Don., G. hiloha (Durand) Wats., G.
grandiflora Lindl. On Boisduvalia densifJora (Lindl.) Wats, and
B. sparsiflora Heller, type of Puccinia BoisduvaUae Peck. On Eulohus
californicus Nutt., type of Puccinia Eulohi Diet, and Holw. On
Oenothera strigidosa T. and G. and 0. ganraeflora T, and G. On
Sphaerostigma spirale (Lehm) Walp., S. hirtella (Greene) Small, S.
viridescens (Lehm) Walp., S. micrantha (Hornem) Walp., and S.
Veitchianum (Hook) Small. Widely distributed and of frequent
occurrence.

130 University of California Pnhlications in Botany [Vol, 7

118. Puccinia Zauschneriae Sydow. I, II, III

Monogr, Ured., vol. 1, p. 436, 1906.

On Zauschneria calif ornica Presl., common in the bay region and
Coast Ranges. The aeeial stage produces witches' brooms which soon
wither away.

Uredinia and telia appear much later and are scarcely distinguish-
able from those of Puccinia Oneotherae. This species together with
the fonns included under numbers 113, 114, and 115 are considered
forms of P. Oenotherae by G. R. Bisby (Am. Jour. Bot., vol. 3, p. 527,
1916).

Telia on Umbelliferae

119. Puccinia asperior Ell. and Ev. I, III

Bull. Waslib. Lab., vol. 1, p. 3, 1884.
On Lcptotacnia dissecta Nutt., Ukiah.

120. Puccinia Cicutae Lasch. II, III

Klotzsch Herb. Mycol., no. 787, 1845,
On Cicuta Bolanderi Wats., Suisun, Pajaro (Thompson) ; on C.
virosa var. calif ornica C, and R,, San Francisco; on C. occidentalis
Dough, Sisson.

121. Puccinia Cymopteri Diet, and Holw. I, III

Bot. Goz., vol. 18, p. 255, 1893.

On Cymopterus teribinthium (Hook) T. and G., King's River
Canon (Holway).

122. Puccinia Ellisi De-Toni. II, III

Saccardo, Syllogi Fungorum, vol. 7, p. 651, 1888.

On Angelica tomentosa "Wats., type of Puccinia Bakeriana Arthur,
Palo Alto (Baker).

123. Puccinia Hydrocotyle (Link) Cke. II, III

Spec. Plant., vol. 6, p. 22, 1825; Grevillea, vol. 9, p. 14, 1880.
On Hydrocotyle prolifera Kell., San Mateo County (Thompson),
San Bernardino (Parish), Pacific Grove (Mrs. Clemens).

124. Puccinia Jonesii Peck. I, III
Bot. Gaz., vol. 6, p. 226, 1881.

On Peucedanum dasycarpuni T. and G., Mount Diablo, Palo Alto
(Thompson), on P. Ilassci C. and R., Atlas, Napa County.

1919] Blasdale: Uredinales of Calif omia 131

125. Puccinia Lindrothii Sjdow. I, III

Acta Soe. Fauna et Flora Fenniea, vol. 22, p. 62, 1902.
On Drudeophytum Hartwegii (Gray) C. and R., Berkeley; on
Velaea arguia (F. and Q.) C. and R., Mount Lowe (Bethel).

126. Puccinia Osmorrhizae (Peck) Cke. and Peck. 0, I, II, III

Kep. N. Y. state Mus., vol. 24, p. 92, 1872; vol. 29, p. 73, 1878.
On Osmorrhiza nuda Torr. and 0. occidentalis (Nutt.) Torr., com-
mon in the Coast Ranges, Calaveras County (Dudley), King's River
Canon (Holway) ; on 0. hrevipes, Sonoma County (Heller).

Telia on Primulaceae

127. Puccinia melanconioides Ell. and Hark. I, II, III

Bull. Calif. Acad. Sci., no. 1, p. 27, 1884.
On Dodecatheon Hendersonii Gray, common throughout the state ;
on D. Jeffreyi Van Houtte, San Bernardino Mountains (Parish).

Telia on Gentianaceae

128. Puccinia Gentianae (Str.) Link. II, III

Spec. Plant, vol. 2, p. 73, 1824.
On Gentiana oregana Engim., Mount Tamalpais; on G. Menziesii
Griseb., Mendocino County (Davy).

Telia on Convolvulaceae

129. Puccinia Convolvuli (Pers.) Cast. I, II, III

Cat. PI. Marseilles, p. 202, 1845.
On Convolvulus luteolus Gray and C. suhacaulis Gray, common
throughout the bay region; on C. occidentalis Gray, Pasadena (Mc-
Clatchie).

130. Puccinia Cressae (DC) Lagh. I, II, III

Biol. Soc. Brot., p. 131, 1889.
On Cressa cretica L., Napa County, Calaveras County, Lassen
County (Davy), Catalina Island (McClatchie), Mountain View
(Thompson), southern California (Bethel).

131. Puccinia Dichondrae Mont. I, III

Gay, Fl. Cliil., vol. 8, p. 46, 1853; also Syll. Crypt., p. 313, 1856.
On Dichondra repens, Forst., San Diego and Orange County
(M. E. Jones).

132 University uf California Publications in Botany [Vol.7

Telia on Polemoxiaceae
MV2. Puccinia Giliae Hark. II, III

Bull. Calif. Acjul. Sei., no. 1, p. 34, 1884.

On Linanthus ciliatus (Benth.) Greene, Mount Diablo (Ilark-
ness) ; on Collomin grandiflora Dougl., Sisson ; on Giliu giliaides
(Benth.) Greene, Tnolumne County; on G. capitata Dougl., Klamath
River, Humboldt County (H. P. Chandler) ; on Navarretia atracty-
laides (Benth.), H. and A., Pasadena (McClatehie).

133. Puccinia plumbaria Peek. I, III

Bot. Gaz., vol. (5, p. 238, 1881.
On Microsteris gracilis (Dougl.) Greene, Mendocino County; on
Linanthus ciliatus (Benth.) Greene, Calaveras County; on Gilia
californica Benth., Snow Mountain, type of Puccinia gilicola Hen-
nings (Purpus).

Telia on Boraginaceae
13-i. Puccinia Cryptanthis Diet, and Holw. II, III

Erythea, vol. 1, p. 249, 1893.

On CryptantJie Torreyana (Gray) Greene, Fresno County (Hol-
way) ; on C. flaccida (Lehm) Greene, Fresno County (Holway).

Telia on Labiateae

135. Puccinia distorta Holway. II, III

Auu. Mycol., vol. 3, p. 20, 1905.
On Hyptis Emoryi Torr., Palm Springs (Parish).

136. Puccinia mellifera Diet, and Holw. I, III

Erythea, vol. 1, p. 25, 1893.

On Salvia mellifera Benth., Pasadena (McClatehie), San Jacinto
and Santa Barbara (Bethel) ; Carmel (Mrs. Clemens) ; on Audiberiia
Palmeri Gray, San Jacinto (Bethel).

137. Puccinia Menthae Pers. I, II, III

Synopsis Fung., p. 227, 1801.

On Mentha canadensis L., Suisun, San Bernardino (Parish), San
Ysidero (Bartholomew) ; on M. sativa L., Pajaro (Thompson) ; on
Micronieria Chamissonis (Benth.) Greene, common in the bay region,
type of Puccinia Micromeriae Dudley and Thompson. On Monardella

1919] Blasdale: Uredinales of Calif ornia ■ 133

villosa Benth., common in the Coast ranges, type of Puccmia Monar-
dellae Dudley and Thompson ; on M. odoratissimum Benth., Lake
Tahoe; on M. lanceolata Gray, Grass Valley (C. J. Wright) ; on M.
viridis Jepson, Napa County; on M. micrantha Gray, San Bernardino

(Parish).

Telia on Solanaceae

138. Puccinia Chamaesarachae Sydow. I, III

Ell. and Ev., N. A. Fungi, no. 1476; SydoAv, Monogr. Ured., vol. 1, p.
263, 1902.

On Chaniaesarache nana Gray, Truckee (Bethel).

139. Puccinia globosipes Peck. II, III

Bull. Torr. Club, vol. 12, p. 34, 1885.
On Lycium calif ornicum Nutt., San Pedro (McClatchie) ; on L.
Andersoni Wrightii Gray, San Bernardino County (Parish),

Telia on Scrophulariaceae

140. Puccinia Adenostegiae Arthur. Ill

Bull. Torr. Club, vol. 29, p. 231, 1902.
On Adenostegia pilosa Greene, Palo Alto (LeRoy Abrams) ; on A.
rigida Jepson, San Diego County (Mrs. Brandegee).

141. Puccinia Antirrhini Diet, and Holw. II, III

Hedwigia, vol. 36, p. 298, 1899.
On Antirrhinum ma jus L., common in the bay region and south-
em California; on A. Nutt all ianum Benth., San Bernardino (Parish) ;
on A. virga Gray, Ukiah (Heller) and Lake County (Jepson). The
writer has also found it possible to infect with this species plants of
the native A. vagans Gray, and the exotic A. assurgens, A. nudicum,
Linaria amethystina, L. delphinifolia, L. Upartita, L. melanthera, and
L. marocanna. The "snapdragon rust" has been a troublesome pest
in California for many years, and is reported to have appeared in the
neighborhood of Chicago, in Ohio, and Indiana {Phytopathology,
vol. 4, p. 400, 1914). It was first found by the writer at San Leandro
in 1896.

142. Puccinia Castillejae (Diet, and Holw.) Arthur, comb. nov.

IL III

Erythea, vol. 1, p. 247, 1893.
Uredinia {Uredo Castilleiae Diet, and Holw.) amphigenous, scat-
tered, round or oval, 0.5 mm. across, early naked, pulverulent, light

134 University of Calif ornia Publications in Botmiy {\^oh. 7

cinnamon brown, ruptured epidermis, not evident; urediniospores
broadly ellipsoid or globoid, 16-24 by 20-29/x; wall golden brown,
thick, 2-3/x, moderately to closely eehinulate, the pores 3, equatorial.

Telia not soon; toliosporos broadly ellipsoid to ovoid, 18-23 by
23-29/x, rounded above, slightly narrowed below, slightly or not con-
stricted at the septmu; wall chestnut to dark golden brown, 2-3/x
thick, moderately thickened at the apex, 5 to 7/*, smooth; pedicel
colorless, short, fragile.

On Castdleja foliolosa IT. and A., Berkeley, Coahuila Valley
(Bethel).

143. Puccinia Cordylanthi Blasdalo. sp. nov. II, III

Sori containing both urediniospores and teliospores, scattered,
minute, rarely confluent, erumpent ; urediospores globose or ellipsoid,
cinnamon brown, with two or three germ-spores; teliospores oblong,
not clavate nor constricted at the septum, apex not thickened ; 15-24//,
by 29-41/*, brown, pedicel short and hyaline.

On stems and leaves of Cordylanthes filifolia Nutt., collected by
E. Bethel at Cajon Pass, San Bernardino County.

This species is more closely related to P. Adeiiostegi-ae Arth. than
to P. Anthirrhini Diet, and Holw. ; it differs from the latter in the
characters of the sori, and from both in the form and size of the telio-
spores. These differences are shown in the figures given below, which
represent the results of the measurement of forty mature spores of
each of the three species.

Average measurement Extreme measurement

Puccinia Antirrhini 21.9 by 51.37/c 16-26 by 41-64^

Puccinia Adenostegiae 21.75 by 42.33;a 18-29 by 32-49^

Puccinia Cordylanthi 20.68 by 35.13/t 15-24 by 29-41u

144. Puccinia Palmeri Diet, and IIolw. Ill
Erythea, vol. 7, p. 98, 1899.

On Penstemon canfertus DougL, Lake Tahoe; on P. Newhcrryi
Graj', Alpine County (Hansen).

145. Puccinia Pentastemonis Peck. Ill

Bull. Torr. Club, vol. 12, p. 35, 1885.
On Pentstemon Roezli Eegel., Cisco (L. S. Smith) ; on P. deustus
Douglas, Sloate, Plumas County (Home) ; on P. azurcns Benth.,
Kewville (Heller) ; on P. Bridgesii Gray, King's River Canon (Hol-
way).

1919] Blasdale: Uredinales of California 135

146. Piiccinia rufescens Diet, and Holw. I, III

Bot. Gaz., vol. 18, p. 253, 1893.
On Pedicularis seniiharhatus Gray, Lake Talioe, King's River
Gallon (Holway), Mount San Antonio (McClatchie).

147. Puccinia Wulfeniae Diet, and Holw. Ill

Erythea, vol. 3, p. 79, 1895.
On Synthyris rotundifolia Gray, Ukiali.

Telia on Rubiaceae

148. Puccinia punctata Link. 0, I, II, III

Obs. Myc, in Ges. naturf. Freunde, Berlin, vol. 2, p. 30, 1816.
On Galium aparine L., Berkeley; on G. triflorum Michx., Sisson
and Shasta Springs, type of Puccinia chondroderma Lindr.

149. Puccinia rubifaciens Johans. Ill

Bot. Centralbl., vol. 28, p. 394, 1888.
On Galium calif omicv.m 11. and A., King's River Cailon (Holway).

Telia on Caprifoliaceae

150. Puccinia Symphoricarpi Hark. Ill

BuU. Calif. Acad. Sci., vol. 1, p. 35, 1884.
On Symphoricarpos racemosus Michx., very common in the bay
region and the Coast Ranges.

Telia on Compositae

151. Puccinia Absinthi DC. II, III

n. franc, vol. 6, p. 56, 1815,
On Artemisia heterophylla Nutt., Berkeley, Los Gatos, Napa and
Rionido; on A. tridentata Nutt., Lassen County (Davy) and Cajon
Pass (Bethel) ; on A. Suksdorfii Piper, Los Gatos (Heller) ; on A.
dracuncidoides Pursh., San Jacinto (Bethel) ; on A. elatior (T. and
G.) Rydberg, Visalia (Holway) ; on A. Douglasiana Bess., Santa Bar-
bara (A. D. E. Elmer). This species is frequently included under
P. Tanaceti DC, from which it differs but little.

152. Puccinia Asteris Duby. HI

Bot. Gall., vol. 2, p. 888, 1830.
On Aster chilensis Nee., Berkeley; on A. yoscmitanus Greene,
Yosemite Yallej^

136 University of California Puhlications in Botany [Vol.7

153. Puccinia Balsamorrhizae Peck. II, III

Bull. Ton-. Cnub, vol. 11, 49, 188-i.
On Behamorrhiza sagittata Nutt., Lake Tahoc, Amador County
(Hansen), Lassen Count}' (F. P. Nutting); on B. dcltoidca Nutt.,
King's River Cafion (Holway).

154. Puccinia Chrysanthemi Rose. II, III
Bull. See. Mycol.,' Franc, vol. 17, p. 92, 1900.

On Chrysanthemum indicum DC, frequeiit throughout the state
and sometimes a serious pest. Telia found but once in southern Cali-
fornia by Bethel. This species is now widely distributed over Europe
and North America. It was first observed by the writer at Berkeley
in 1910.

155. Puccinia Cirsii Lasch. II, III

Rabh. Fungi Europ, No. 89, 1859.

On Cirsium edule Nutt. and C. lanceolatum (L) Scop., Berkeley;
on C. undidatum (Nutt.) Spring, Antioch ; on C. Breweri (Gray)
Jepson, King's River Canon, type of Pucciiiin calif ornica Dietel and
Holw. (Holway); on C. occidentale (Nutt.), Jepson, San Francisco;
on C. Driimmondii acaulescens (Gr.) Cov., King's River Cafion (Hol-
way) ; on C. quercetorum (Gray) Jepson, Santa Cruz (Thompson).

156. Puccinia conferta Diet, and Holw. HI
Erythea, vol. 1, p. 250, 1893.

On Artemisia heterophylla Nutt., King's River Cafion, type collec-
tion (Holway), Ukiah (Holway and Blasdale) ; on A. calif ornica
Less., Pasadena, type of Puccinia recondita Dietel (McClatchie),

157. Puccinia crepidicola Sydow. II. Ill

Oestr. bot. Zeitsehr., vol. 51, p. 17, 1901.

On Crepis pleurocarpa Greene, Mount Eddy, Siskiyou County
(C. F. Baker).

158. Puccinia Crepidis-acuminatae Sydow. II. Ill

Oestr. bot. Zeitsehr., vol. 51, p. 27, 1901.

On leaves and stems of Crepis acuminata Nutt., King's River
Canon (Holway) ; on C. pleurocarpa, Siskiyou and Trinity- counties
(G. D. Butler).

1919] Blasdale: Uredinales of California 137

159. Puccia Franseriae Sydw. II, III

Ann. Mycol., vol. 1, p. 326, 1903.
On Franseria dumosa Gray, Palm Springs.

160. Puccinia Grindeliae Peck. Ill

Bot. Gaz., voL 4, p. 127, 1879.
On Grindeliu sp., Julian, San Diego County (Bethel).

161. Puccinia Harknessii Vize. II, III

Grevillea, vol. 7, p. 11, 1878.
On Lygodesmia spinosa Nutt., Lassen County (Davy) ; on Ptiloria
exigua Nutt., San Bernardino County (Parish).

162. Puccinia Helianthellae (Peck) Arthur. II, III

Bull. Torr. Club, vol. 31, p. 4, 1904.
On Helianthella nevadensis Greene, Nevada County (Heller) ; on
H. calif ornica Gray, Nevada County (M, E. Jones).

163. Puccinia Helianthi Schw. II, III

Syn. Fung. Carol., p. 73, 1822.
On Helianthus annuus L., frequent throughout the state; on H.
lenUcularis Dough, Fresno County (Holway) and Pasadena (Mc-
Clatchie) ; on H. delilis Nutt, in cultivation, Berkeley.

164. Puccinia Hemizoniae Ell. and Tracy. II, III

Jour, of Mycol., vol. 7, p. 43, 1891.
On Hemizaniu luzulaefolia DC, Berkeley and Santa Rosa; on E.
Clevelandii Greene, Santa Rosa ; on H. citrina Greene, Marin County
(Heller) ; on Lagophylla congesta Greene, Berkeley, type of Puccinia
LagophyUae Diet, and Holw.

165. Puccinia Hypochoeridis Oud. II, III

Nederl. Kruidk. Archief., (2), vol. 1, p. 175, 1872.
On Hypochaeris glabra L., Berkeley, San Francisco, Mount Diablo,
Monterey (Bethel).

166. Puccinia intermixta Peck. I. Ill

Bot. Gaz., vol. 4, p. 231, 1879.
On Iva axillaris Pursh., Lassen County (DavjO-

138 University of Calif ornia Publications in Botany [Vol.7

107. Puccinia investita Sclnv. I, III

N. A. Fungi, do. 2932; Peek, Rep. X. Y. St. Mus., p. 117, 1872.

On Gnaphalium chilense Spreng, San Francisco; on G. sp., Mount
Tamalpais (Bethel).

168. Puccinia Millefolii Fuckl. Ill

Syiiib. Myc, vol. 5J, 1869.
On Achillea millefolium L., San Francisco (Bethel).

169. Puccinia splendens Vizc ITT
Grevillea, vol. 7, p. 11, 1878.

On Ilymenoclea salsola T. and G., Mojave Desert (Parish) ; on
H. monoejyra T. and G. (Griffiths).

170. Puccinia Stephanomeriae Sydow. IT, HI

Mouogr. Urod., vol. 1, p. 117, 1904.

On Stephanomeria lactucina Gray, Sisson; on 8. chicoriaceum
Gray, Pasadena (McClatchie), Mount Wilson (Bethel); on ^S'. runci-
nata, Victorville (Bethel) ; on Ptiloria carduacea, Claremont (Baker).

171. Puccinia Taraxaci (Rebent) Plowr. II, III

Plo^vr. British Ured. and Ustilig., p. 186, 1889.
On Taraxacwm Taraxacum (L), Karst., Berkeley and Sisson.

172. Puccinia Troximontis Peek. II. HI

Bot. Gaz., vol. 6, p. 227, 1881.

On Agoseris plehiea Greene, Santa Rosa, Berkeley and San Fran-
cisco; on A. larhellata, Mount Eddy (Copeland) ; on A. hirsuta
(Hook) Greene, Berkeley.

173. Puccinia variolans Hark. HI

Bull. Calif. Acad. Sci., no. 1, p. 15, 1884.

On Aplopappus squarrosus H. and A., Pasadena (McClatchie) ; on
Tetradymia glahrata Gray, Lassen County (Da\y).

174. Puccinia Wyethiae (Peck) Ell. and Ev. IT. TIT

Bot. Gaz., vol. 7, p. 46, 1882; Ell. and Ev., N. A. Fungi, no. 2987.

On Wyethia anejustifolia Nutt., Berkeley and Ukiah, also Los
Angeles, Long Beach, and San Ysidro (Bethel).

1919] Blasdale: Uredinales of Calif ornm 139

175. Puccinia Xanthii Schw. Ill

Syn. Fung. Carol, p. 73, 1822.
On Xanthium canadense Mill., San Francisco, Suisuu, Fresno
County (Holway), Pasadena (McClatchie), San Bernardino (Parish),
Los Angeles, and Long Beach (Bethel).

PUCCINIASTRUM otth.
Mitth. nat. Ges. Bern, p. 71, 1861.
Aecial stage similar to that of the genus Coleosporiuni', on the
leaves of species of Pin us and Ahies. Urediniospores borne singly
on pedicels, walls colorless. Telia indehiscent in layers in or beneath
the epidermis. Teliospores usually prismatic, two- to four-celled.

176. Pucciniastrmn Goodyerae (Tranz.) Arthur. II

Arthur, N. A. Flora, vol. 7, p. 105, 1907.
On Goody era 3Ienziesii Lindl., Mount Eddy (Copeland).

177. Pucciniastrum pustulatum (Pers.) Dietel. 0, I, II, III

Dietel, in Engler and Prant., Natiirl. Pflanzenfam., vol. 1, p. 47, 1897.

The aecial stage not definitely known from California, but shown
by Frazier (Mycologia, vol. 4, p. 175, 1912) to develop on Abies hal-
samea (L) Mill.

Uredinia and telia on Epilohium franciscanum Barbey ; on E. aden-
ocaulon Treal., E. coloratum Muhl., E. Kolosericeum Treal., and E.
calif ornicu'm Haussek. Common in the Coast Ranges. Uredinia found
throughout the year.

178. Pucciniastrum P5rrolae (Pers.) Dietel. II, III

Gmel. Syst. Xat., vol. 2, p. 1474; Dietel, in Engler and Prant., Natiirl.
Pflanzenfam., vol. 1, p. 47, 1897.

On Pyrola picta Smith and P. secunda L., Sisson and Mount Tal-

lac, Plumas County (Home) ; on Chimaphila umhellata (L) Nutt.,

Siskiyou County (Meinicke).

179. Pucciniastrum sparsum (Wint.) Ed. Fischer. II, III

Eabh. Krypt. Flora, vol. 1, p. 245, 1881 ; Beitrag. Krypt. Sehweiz., vol. 2,
p. 469, 1904.

On Arbutus MenziesU Pursh., Mount Tamalpais, type of TJredo

Arhuii Diet, and Holw. ; on Arctostaphylos patula Greene, Sisson, tj'pe

of TJredo Copel^ndi Sydow; on A. Hookcri Don., Antonio Station,

Santa Barbara County (Brandegee).

140 University of Calif orma Puhlicatians in Botany [Vol.7

RAVENELIA uerk.
Card. Chron., p. 132, 1853.
Uredinia erumpent, without peridiuin ; iirodiniospores borne singly
on pedicels, wall colored. Teliospores fascicled on compound stalks,
one- or two-celled, forming heads bordered by hyaline cysts,

ISO. Ravenelia arizonica Ell. and Ev. II, III

Bull. Torr. Club, vol. 22, p. 363, 1895.
On Prosopis juliflora (Sw.) DC, San Diego (Bethel).

181. Ravenelia versatilis (Peck) Diet. II, III

Hcdwigia, vol. 33, p. 368, 1894.
On Acacia Gregii Gray, San Bernardino (Toumey), Banning
(Parish).

TRANZSCHELIA arth.

Result. Sc'i. Cong. Bot. Vienue, p. 340, 1906.

Aecia cylindrical ; aeciospores globoid ; wall colored. Uredinia
erumpent ; urediniospores borne singly on pedicels mixed with para-
pliyses. Telia erumpent, pulverulent; teliospores attached to a com-
mon stalk by short inconspicuous pedicels.

182. Tranzschelia punctata (Pers.) Arth. 0, I. II, III

Ann. Bot. Usteri, vol. 20, p. 135, 1796; Arthur, op. cit.

Aecia not known from California but found on various species of
Hepatica, A^iemone and Thalictrum {Aecidiiim piuictatum Pers.) in
the eastern United States.

Uredinia and more rarely telia on cultivated peach, plum, prune,
almond, and apricot, especially in the southern part of the state, but
widely distributed. This is the "prune rust" which is sometimes the
source of considerable losses to fruit growers.

UREDINOPSIS JIAGN.

Atti Cong. Bot. Geneva, p. 167, 1893.

Aecia similar to those of Coleosporium, found on leaves of species
of Ahies. Uredinia larger and more conspicuous than the aecia, the
agglutinated spores ejected from the delicate peridium in a long
mucilaginous filament. Telia indehiscent; teliospores four-celled, with
thin wall.

1919] Blasdale: TJredhmles of Calif orrda 141

383. Uredinopsis Copelandii Sydow. I, II, III

Ann. Mycologici, vol. 2, p. 34, 1904.

Aecia not definitely reported from California, but elsewhere on
species of Abies.

Uredinia and telia on Athyrium cyclosorum Kupr., Sisson (Cope-
land).

184. Uredinopsis Pteridis Diet, and Holw. I, II, III

Ber. d. deut. bot. GeselL, Bd. 13, p. 331, 1895.

Aecia {Peridermium pseudo-halsameum Arthur and Kern) on
AMes grandis Lindl., Eureka. Reported by Hedgcock (Mycologia,
vol. 4, p. 141, 1912) on A. lasiocarpa Nutt., and A. nohilis Lindl.

Uredinia and telia on Pteris aquilinum piibescens Underwood.
Sisson , Lake Tahoe, San Francisco, Mount Tamalpais, Pasadena
(McClatchie), Long Beach (Bethel). For culture experiments relat-
ing to this form see "Weir and Hubert (Am. Jour. Bot., vol. 4, p. 328,
1917).

UEEDO

Under this form genus are grouped the uredinial stages of certain
species for which the mature stages are unknown.

185. Uredo Acaenae Ell. and Ev.

N. A. Fungi, no. 3150, 1894.
On Acae'n-a tridactyla PresL, frequent in the bay region. Arthur
(N. A. Flora, vol. 7, p. 174) takes this to be uredinia of Phragmidium
Ivesiae Sydow.

186. Uredo Gaillardiae Diet, and Holw.

Erythea, vol. 7, p. 98, 1899.
On Gaillardia aristata Pursh, Dunsmuir (Holway).

187. Uredo Nicotianae Arthur, sp. nov.

Uredinia mostly hypophyllus, scattered upon discolored areas 1 to
1.5 cm. across, round, 0.2 to 0.5 mm. in diameter, early naked, applan-
ate, subepidermal, pulverulent, pale cinnamon to whitish, ruptured
epidermis evident ; urediniospores globoid to broadly ellipsoid, 23 to
26/x by 24 to 32/a; wall colorless, 1.5 to 5/* thick, closely and coarsely
verrucose, the pores obscure.* Collected in small amount at Rionido
on Nicotiana Bigelowii "Wats. The spores resemble those of a Coleo-
sporium, although not in well defined chains.

This description furnished by Professor J. C. Arthur.

142 University of Calif ornia Ptihlications in Botany [Vol. 7

188. Uredo Phoradendri Jackson

Mem. Brooklyn Bot. Gard., vol. 1, p. 285, 1918.
On Phoradcndron longispicum Trelease, Chico (C. C. Thomas).

189. Uredo Sphacelicola Diet, and ITohv.

Erythea, vol. 1, p. 248, 1893.
On Sphaccle calycina Benth., Mount Tamalpais.

UROMYCES UNGER

Exanth. Pfl., p. 277, 1833.

Accia eriimpent, cupulate, or cylindrical ; aeciospores in chains
globoid or ellipsoid, often angular. Uredinia definite and without
paraphyses ; urediniospores borne singly on pedicels ; wall colored,
usuall}" echinulate. Telia erumpent or long-covered by the epidermis ;
teliospores one-celled, wall firm, deeply colored, thick.

Telia on Graminaceae

190. Uromyces Jacksonii Arth. and Fromme. II, III
Torreya, vol. 15, p. 2G0, 1915.

On Agrostis pallens Trin., Atlas, Napa County; on Hordeum nodo-
sum L, San Mateo County (Copeland).

191. Uromyces Peckianus Farlow\ II, III

Proc. Am. Acad. Arts and Sci., p. 76, 1883.

Aecia on Salicornia amhigua Michx., Chenopodinm album L.,
C. murali L., and Atriplex sp., Monterey (Bethel).

Uredinia and telia on Distychlis spicata Greene, Long Beach
(Bethel), Femdale (Davy and Blasdale). Probably common along
the coast.

Telia on Cyperaceae

192. Uromyces Scirpi (Cast.) Burr. 0, I, II, III

Cat. PI. Marseilles, p. 214, 1845; Bot. Gaz., vol. 9, p. 188, 1884.

Aecia on Oenanthe calif ornica Wats., central California.

Uredinia and telia on Scirpus pacificus Britton, and S. calif ornicus
(Mey.) Britt, Long Beach (Bethel) ; on 8. paludosiis Nels., Escondido
(Bartholomew).

1919] Blasdale: Uredinales of California 143

Telia on Juncaceae

193. Uromyces Junci (Desmaz) Tnl. 0, I, II, III

Ann. Sci. Nat., ser. 4, Bot., vol. 2, p. 148, 1854.

Aecia on Ambrosia psilostachya DC, San Jacinto and Coahuila
Valley (Bethel).

Uredinia and telia on Juncus halticus "VVilld., Long Beach, Cajon
Pass, and Victorville (Bethel), Mohave C. (Parish) ; Catalina Island
(Bethel); on J. leseurii Boland, Sunol (Home), Pasadena (Mc-
Clatchie) ; on /. textUis Buch., Cajon Pass (Bethel), Pasadena
(McClatchie), San Bernardino County (Parish) ; on J. patens Mey.,
San Mateo County (Copeland) ; on /. Mexicanus Willd., San Diego
County (Chandler).

194. Uromyces Junci-effusi Sydow. II, III

Monogr. Ured., vol. 2, p. 290, 1910.

On Juncus Xiphioides E. Meyer, Ukiah and Mill Valley; on /,
phaeocephalus Engelm., Long Beach (Bethel) ; on /. nevadensis Wats.

Telia on Liliaceae

195. Uromyces aureus Diet, and Hohv. I, III

Hedwigia, vol. 32, p. 30, 1893.
On Allium validum Wats., King's River Caiion, type collection
(Hoi way).

196. Uromyces bicolor Ellis. 0, I, II, III

Contr. U. S. Xat. Herb., vol. 4, p. 231, 1893; Bull. Torr. Club, vol. 24,
p. 282, 1897.

On Allium, unifolium Kell., Berkeley (type collection for JJromijces
aterrimus Diet, and Holw.) and Sherwood, Mendocino County; on
A. validum Wats., Sisson and Lake Tahoe.

197. Uromyces Brodieae Ell. and Hark. I, III

Bull. Calif. Acad. Sci., no. 1, p. 28, 1884.
On Brodiaea oapitata, Berkeley, Mount Diablo, Yosemite Valley.

198. Uromyces Chlorogali Diet, and Hohv. I, II. Ill
Erythea, vol. 1, p. 246, 1893.

On Chlorogalum pomeridianum (Ker.) Knuth, Berkeley, ]\Iount
Diablo, Atlas, Napa County, Palo Alto (Thompson).

144 TJniverdty of California Puhlications in Botany [Vol. 7

199. Uromyces Lilii Clinton. 0, I, IT, III

27th Ik'pt. N. y. State Mus., p. 103, 1875.
On Lilium columhiamim Hansen, Sisson and King's River Canon,
type of Uramyces Ilolwayi Lagh. (Ilolway) ; on L. ruhescens Wats.,
Shasta Springs ; on L. parvum Kellogg, Lake Tahoe ; on L. Washing-
tonianum Kell., Forest, Lassen County (F. P. Nutting).

200. Uromyces Zygadeni Peck. 0, I, II, III
Bot. Gaz., vol. 6, p. 239, 1881.

On Zygadenus Fremonti Torr., Pasadena (McClatchie), San
Mateo County (Thompson). On Z. sp., Santa Monica Mountains
(Holway).

Telia on Polygonaceae

20L Uromyces intricatus Cooke. 0, I, II, III
Grevillea, vol. 7, p. 3, 1878.

On species of Eriogonum {Uromyces Eriogoni Ell. and Hark.)
and Chorizanthe {Uromyces Chorizanthis Ell. and Hark), very com-
mon throughout the state. Arthur (N. A. Flora, vol. 7, p. 245) reports
it from Eriogonum cernum Nutt., E. dumosum Greene, E. elongatum
Benth., E. fascicidatum Benth., E. latifoliiim Smith, E. nudum Dougl.,
E. parvifolium Smith, E. vimineum Dougl., E. virgatum Benth.,
Chorizanthe cuspidata Wats., C. rohusta Parry, and C. pungens Benth.

202. Uromyces Polygoni (Pers.) Fuckl. I, II, III

Symb. Mueol., p.-G-i, 1869.

Aecia very rare but collected by Bethel on Polygonum aviculare L.
at Long Beach and San Jacinto.

Uredinia and telia very common througliout the state on the same
host; on P. erectum L., Escondido (Bartholomew).

Telia on Chenopodiaceae

203. Uromyces Betae (Pers.) Lev. 0, I, II, III

Pers. Syn. Fung., p. 220, 1801; Ann. Sci. Nat., ser. 3, Bot., vol. 8, p. 375,
1847.

Aecia not reported from California but found in Europe and
Australia.

Uredinia very common throughout the state on Beta vulgaris L.
and B. chicla.

Telia found occasionally, specially upon seed-bearing plants.

1919] Blasdale: Uredmales of California 145

204. Uromyces Chenopodii (Duby) Schroet. I, II, III

Duby., Bot. gall., vol. 2, p. 899, 1830; Schroeter, Kunze, Fung. Sel.,
no. 214, 1880.

Aecia, uredinia, and telia on leaves and stems of Dondia multiflora

(Torr.) Heller, Long Beach, San Diego, and Point .Firmin (Bethel) ;

on D. calif ornica (Wats.) Heller, Laguna Beach (H. W. Fawcett).

Telia on Portulaceae

205. Uromyces Spragueae Hark. I, III

Bull. Calif. Acad. Sci., vol. 1, p. 44, 1884.

On Calypiridium umhcllatum Torr. Greene, Lake Tahoe, Yosemite
Valley, Alpine County (Hansen), Plumas County (Meinicke), King's
River Caiion (Holway).

Telia on Caryophyllaceae

206. Uromyces Caryophyllina (Schrank) Wint. 0, I, II, III

Sch. Baier. Fl., vol. 2, p. 666, 1789; Eab. Krypt. Flora, vol. 1, p. 149,
1881.
Uredinia and telia frequent throughout the state on certain varie-
ties of Dianthus caryophylliis L., and sometimes the cause of large
losses to florists. It was introduced into the eastern states about 1860
and first noted by the writer in California in 1896.

207. Uromyces pulchellus Ell. and Ev. II, HI

Bull. Torr. Club, vol. 22, p. 57, 1895.
On Silene Douglasii Hook., Lake Tahoe.

Telia on Ranunculaceae

208. Uromyces Aconiti-lycoctoni (DC) Wint. I, II, III

Kabh. Krypt. Flora, vol. 1, p. 153, 1884.
On Aconitum coliimhianum Nutt., Sisson.

209. Uromyces Jonesii Peck. II, III

Bot. Gaz., vol. 7, p. 45, 1882.
On Eanunculus flammula L., King's River Canon (Holway).

Telia on Leguminoseae

210. Uromyces abbreviatus Arthur. Ill

Bull. Torr. Ciuh, vol. 42, p. 587, 1915.
On Psoralea pJujsoides Dougl., Calistoga (Harkness), Pinehurst
(Bethel).

146 University of Calif ornm Puhlications in Botany ['^^oi,. 7

I'll. Uromyces albus Diet, and IIolw. 1. Ill

llodwigia, vol. 3(j, p. 297, 1897.
On Vici.a americana Mulil., Crocker's Ranch, Mariposa County;
on V. americana truncata Brewer, Sloate, Plumas County (Home) ;
on V. calif ornica Greene, Cuyamaca INIountains (LeRoy Abrams), San
Diego County (Bethel).

212. Uromyces appendiculatus (Pers.) Link. 0, I, II, III

Ann. Bot. Usteri, vol. 15, p. 16, 1795; Link, Observ., vol. 2, p. 26, 1816.

On Phaseolus vulgaris L. and P. lunatus L., frequent, especially
in the southern part of the state, but not destructive.

213. Uromyces Fabae (Pers.) De Barv. 0, I, II, III

Neues Mag. Bot., vol. 1, p. 93, 1794; Ann. Sci. Nat., Bot., vol. 20, p. 80,
1863.

On Lathyrus Bolanderi Wats., L. Jepsonii Greene, L. violaceus

Greene and L. Torrcyi Gray, common in the bay region; on L. sul-

fureus Brewer, Plumas County (Home).

214. Uromyces fallens (Desmaz) Kern

Plant Krypt., 1325; Kern, Phytopath., vol. 1, p. 6, 1911.
On Trifolium prate^ise L., of frequent occurrence in both the north-
ern and southern portions of the state.

215. Uromyces Glycyrrhizae (Rabh.) Magn. II, III

Ber. cleutsch. bot. Ges., vol. 8, p. 383, 1890.
On Glycyrrhiza lepidota glutinosa Wats., Tuolumne County, Wal-
nut Grove, Cloverdale (Heller), Victorville (Bethel).

21 G. Uromyces Lupini B. and C. 0. I. II. Ill

Proc. Am. Acad. Arts and Sci., vol. 4, p. 127, 1858.
On Lupinns formosu-s hridgesii Green, L. alhifrotis Benth. and
L. Chamissonis Esch., frequent in the bay region; on L. rivularis
Dough, San Bernardino County (Parish) ; on L. Douglasii Agardh,
King's River Caflon (Holway) ; on L. latifoUus Agardh., Santa Cruz
County ( Thompson ) .

217. Uromyces Medicaginis Pass. II, III

Thiini, Herb. Myc. Oecon., p. 155, 1874.
On Medicago lupulina L., Berkeley and Humboldt County; on
M. sativa L., common, especially in the southern part of the state.

1919] Blasdale: Uredinales of California 147

218. Uromyces oblongus Yize. I, III

Grevillea, vol. 5, p. 110, 1877.

On Trifolium variegatum Nutt., T. gradient iim T. and G., T.
microcephalum Pursh., T. microdon H. and A., T. diiMum Sibth,, T.
roscidum Greene, T. ciliolatum Bentli., T. depauperatum Desv., T.
albopurpureum T. and G., T. Macraei H. and A., T. tridentation
Lindl., T. stcnopJiyllum Nutt., T. oliganthum Steud. Of frequent
occurrence in the Coast Eanges and in the Sierras.

219. Uromyces occidentalis Diet. II, III

Hedwigia, vol. 42, p. 98, 1903.

On Lupinus leptophyllus Benth., Sisson; on L. latifolius Agardh,
Boulder Creek and Berkeley.

220. Uromyces punctatus Schrot. 0, I, II, III

Abh. scliles. Ges., vol. 48, p. 10, 1870.

Aecia on species of EupJiorltia but not reported from America.

Uredinia and rarely telia on Astragalus Memiesii Gray, San
Francisco; on A. lentiginosus Dougl., Kern County (Davy) ; on
A. leucopsis Torr., Long Beach (Bethel) ; on A. Preussii Gray, Indio
(E. A. BessejO ; on A. Purshii Dougl., Hornbrock (Copeland) ; on
A. pycnostachys Gray, San Mateo County (Baker) ; on Lotus erio-
phorus Greene, San Francisco (M. E. Jones) ; on L. glaher (Yogel)
Greene, Long Beach (Bethel).

221. Uromyces Trifolii (Hedw.) Lev. I, II. Ill

An. Sci. Nat., ser. 3, Bot., vol. 8, p. 371, 1847.

On Trifolium repens L., frequent in both the northern and south-
ern part of the state, but not destructive.

Telia on Euphorbiaceae
222. Uromyces proeminens (DC) Pass. 0, I, II, III

Fl. franc, vol. 2, p. 235, 1805 ; Rabh. Krypt. Flora Europ., p. 1795, 1873.
Aecia {Aecidium Eup}iorl)i.ae-hypericifoliae Schw.) on Euphorhiu
serpyUifolia Pers., Berkeley, Olema, San Francisco, Amador County
(Hansen), King's River Canon .(Holway), Catalina Island (Mc-
Clatchie) ; on E. polycarpa Benth., Mojave County (Parish) ; on E.
albomarginata T. and G., Inglewood (LeRoy Abrams).

148 University of Calif orni<b Puhlications in Botany [Vol. 7

Telia on IIypericaceab
228. Uromyces Hyperici-frondosi (Schw.) Arthur. 0, I, II. Ill

Sclnv. mat. Ges., Leipzij^, vol. ]. p. 68, 1822; Arthur, Bull. Minnesota,
Acad. Sci., vol. 2, p. 1.5, 1883.

On Ilypericum anagaloidcs Ch. and Sch., San Francisco, Olema,
Mendocino County; on H. Scouleri Coulter, Amador County (Han-
sen).

Telia on Primulaceae

224. Uromyces nevadensis Hark. Ill

Bull. Calif. Acad. Sci., vol. 1, p. 36, 188-i.
On Primula suffrutescens Gray, Lake Tahoe (Ilarkness). This
collection was probably made in Nevada but very near the California
boundary.

225. Uromyces Armeriae (Schl.) Lev. I. II. Ill

Ann. Sci. Nat., ser. 3, Bot., vol. 8, p. 375, 1847.
On Armeria vulgaris Willd., San Francisco, Pacific Grove (Hel-
ler).

Telia on Plumbaginacae

226. Uromyces Limonii (DC) Lev. I, II, III

Fl. franc, vol. 2, p. 195, 1805; Lev., Diet, d'hist. nat., p. 19, 1840.
On Statice Limoniuin L. var. calif ornicum Gray, San Francisco,
Belmont, Long Beach (Bethel).

Telia on Acanthaceae

227. Uromyces Ruelliae Holway. II, III

Ann. Myeol., vol. 2, p. 394, 1904.
On Beloperone californica Benth., western edge of the Colorado
Desert, type of Uredo Beloperonis Arthur (Parish).

Telia on Compositae

228. Uromyces Bidentis Lagh. II, III

Bull. Soe. Mycol. France, p. 213, 1895.
On Bidens pilosa L., Orange; on B. expansa Greene, Long Beach
(Bethel).

UROPYXIS SCHROT
Hedwigia, vol. 14, p. 165, 1875.
Uredinia erumpent, usually wit-h paraphyses ; urediniospores borne
singly on pedicels. Telia erumpent ; teliospores two-celled by a trans-
verse septum; inner wall firm and deeply colored, outer wall gelatin-
ous, thick.

1919] Blasclale: Uredinales of California 149

229. Uropyxis Amorphae (Curt.) Schroet. II, III

Am. Jour. Ai-ts and Sci., (2), vol. 6, p. 353, 1848; Hedwigia, vol. 14,
p. 165.

On Amorpha calif ornica Nutt., Mount Tamalpais and Los Gatos,
Catalina Island (Bethel).

230. Uropjrxis sanguinea (Peek) Arthur. II, III

Bot. Gaz., vol. 4, p. 128, 1879; N. A. Flora, vol. 7, p. 155.
On Berheris pinnata Leg., Berkeley ; on B. nervosa Pursh., Hum-
boldt Count}' ; on B. repens Lindl., Calaveras and Tuolumne counties.

ADDENDA

The following represent important additions to the foregoing list
the data for which were received too late for inclusion in the proper
sequence.

231. Cronartium occidentale Hedgcock, Bethel, and Hunt. 0, I,

II, III

Jour. Agric. Eesearch, vol. 14, p. 411, 1918.

Aecial stage not as yet reported from California but found in
Colorado, Utah, New Mexico, and Arizona on species of Pimis belong-
ing to the piiion group.

Uredinia and telia on Rihes tenuiflormn Lindl., "abundant along
river wash near Monrovia" (Bethel). It was first reported from the
same locality by "W. M. Phillipson.

This record is of especial interest on account of the similarity of
the uredinial and telial stages to those of the white pine blister rust,
the aecial stage of which is very destructive to pines of the five-leaved
group.

232. Gymnosporangium inconspicimm Kern. 0, I, III

Bull. Torr. Club, voL 34, p. 461, 1907.
Aecia {Roestilia Harknessianoides Kern) on Amelaiiehier sp.
Telia on Juniperxis occidentalis Hook. Both forms collected by
Bethel in Bear River Valley, San Bernardino Mountains.

233. Melampsora monticola Mains. II, III

Phytopathology, vol. 7, p. 103, 1917.
Uredinia on Euphorbia sp., San Jose Canon, near Carmel (Hol-
way) ; on E. leptosera Engelm., Pacific Grove (Mrs, Clemens).

150 University of Calif orni<i Piiblicatimis in Botany [Vol. 7

234. Puccinia Andropogonis Schw. 0, I, II, III

Trans. Am. I'hil. Woe, II, vol. 4, p. 295, 1834.

Aecia on Poistenion Menziesii Hook., Yosemite Valley and Lake
Taboo.

Telia not reported from California, but to be expected on species
of Andropogon.

235. Puccinia Toumeyi Sidow. Ill
Saccardo, Sylloge Fung., vol. 16, p. 299, 1902.

On Penstemon spectabilis Thurber, Mount Lowe.

236. Puccinia yosemitana Blasdale, sp. nov. I, III

Aecia usually solitary and widely scattered, cylindrical, about
0.2 mm. wide and 2.0 mm. long, orange yellow ; pseudoperidia of thick-
walled cells with deeply-colored contents, finally dehiscing by disinte-
gration at apex.

Telia associated with aecia, amphigenous, scattered but occasionally
confluent, about 0.5 mm. long, elliptical, black, pulverulent, early
naked ; teliospores ellipsoid or oblong, rounded and but slightly thick-
ened at apex, barely constricted at septum, smooth, 32-38 by 20-22/jt ;
pedicel thick, persistent, often 80/x long.

On Grilia pungens var. Hookeri Gray, found on trail from Yosemite
Valley to top of Yosemite Falls.

237. Pucciniastrum Galii Fischer

Ured. d. Schweitz, p. 471, 1904.
Uredinia on Galiiim trifloruni Michx. Yosemite Valley.
3. Aecedium Trigochinis Diet, and Holway

It has be^n shown very recently by Bethel that this is one of the
many forms representing the aecial stage of Puccinia suhmitens.

67. Puccinia sejuncta Sydow.

It is now known that the form referred to under this number is
Aeciddum Colunibiense Ell. and Ev. (Erythea, vol. 1, p. 206, 1893).

1919]

Blasdale: Uredinales of California

151

VII. INDEX TO SPECIES OF UREDINALES

Aecidium Aquilegiae Pers., 51.
aematidis DC, 51.
Collinsiae Ell. and Ev., 1.
Columbiense Ell. and Ev., 67.
Euphorbiae-hypericifoliae Schw.,

222.
Fendleri Tracy and Earl, 54.
Graebnerianum Hennings, 2.
Hydrophillae Peck, 61.
punctatum Pers., 182.
Phaceliae Peck, 61.
roestilioides Ell. and Ev., 58.
Triglochinis Diet and Holw., 3.
Valerianellae (Biv.) Bernli., 4.
Baeodromus californicus Arth., 5.
Bubackia Crotonis (Cke.) De Toni, 6.
Caeoma Bigelowii Thiim, 28.
nitens, 25.

occidentalis Arth., 27.
Calyptospora columnaris (Alb. and

Schw.) Kuhn, 7.
Coleosporium Baccharidis Cooke, 16.
Bletiae Diet., 8.
Campanulae (Pers.) Lev., 9.
Madiae Cooke, 10, 35.
Solidaginis (Schw.) Thiim, 11.
Cronartium occidentale Hedg.,
Bethel and Hunt, 231.
coleosporioides (Diet, and Holw.)

Arth., 12, 13.
Comandrae Peck, 14.
filamentosum (Peck), Hedgcock, 13.
pyriforme (Peck) Hedg. and Long,

14.
Qiiercus (Brond) Schroet, 15.
Eriosporangium evadens (Hark.)
Arth., 16.
punctato-striatum (Diet, and
Neger.) Arth., 17.
Gymnosporangium Blasdaleanum
(Diet, and Holw.) Kern, 18.
Harknessianum (Ell. and Ev.)

Kern, 19.
inconspicuum Kern, 232.
Koreaense (Hennings) Jackson, 20.
Hyalopsora Cheilanthis (Peck)
Arth., 21.
laeviscula (Diet, and Holw.) Arth.,

22.
Polypodii (DC) Magnus, 23.

Kuehneola uredinis (Link) Arth., 24.
Kunkelia nitens (Schw.) Arth., 25.

Eosae-gyninocarpae (Diet.) Arth.,
26.
Melampsora albertensis Arth., 27.

Bigelowii Thiim, 28.

Lini (Schum) Desmaz., 29.

Medusae Thiim, 30.

nionticola Mains, 233.

occidentalis Jackson, 30.
Melampsoropsis Piperiana Arth., 31.

Pyrolae (DO) Arth., 32.
Neoravenelia Holwayi (Diet.) Long,

33.
Nyssopsora echinata (Lev.) Arth., 34.
Peridermium californicum Arth. and
Kern, 35.

conorum Piceae, 32.

Ephedrae Cooke, 36.

filamentosum Peck, 13.

Harknessii Moore, 12, 15.

montanum Arthur and Kern, 11.

pseudo-balsameum (Diet, and
Holw.) Arth. and Kern, 184.

pyriforme Peck, 14.
Phragmidiuni Andersoni Shear, 37.

disciflorum (Tode) James, 38.

imitans Arth., 39.

Ivesiae Sydow, 40, 185.

montivagum Arth., 41.

occidentale Arth., 42.

Eosae-californicae Diet., 43.
Pileolaria Toxicondendri (Berk, and

Rav.) Arth., 44.
Polythelis Thalictri (Chev.) Arth.,

45.
Puecinia Absinthi DC, 151.

abundans (Pk.) Jackson, 46.

Aristidae Tracy, 49.

Acetosae (Schum.) Koern, 94.

Adenostegiae Arth., 140, 143.

adspersa Diet, and Holw., 47.

Alliorum (DC) Corda, 82.

amphispilusa Diet, and Holw., 95.

anachorita Ell. and Hark., 84.

Andropogonis Sclnv., 234.

anomala Rostr., 48.

Antirrhini Diet, and Holw., 141,
143.

Asarina Kze., 92.

152

University of California Publications in Botany [^^ol. 7

Asparagi DC, 83.
asperior Ell. and Ev., 119.
Asteris Duby, 152.
Asterum (Schw.) Kern, 72.
atro-fusca (Dudley and Thompson)

llolway, 73.
Baccharidis Diet, and Hohv., 17.
Bakeriana Arthur, 122.
B;ilsamorrhizae Peck, 153.
Blasdalei, Diet, and Holw., 82.
bistortae (Str.) DC, 96.
Boisduvaliae Peck, 117.
Burnettii Griff., 58.
californica Diet and Holw., 155.
Calochorti Peck, 84.
caniculata, (Schw.) Lagerh., 74.
Caricis (Schum) Kebent., 75.
Castillejae (Diet, and Holw.)

Arthur, 142.
Chamaesarache Sydow, 138.
chondroderma Lindr., 148.
Chrysanthemi Koze, 154.
Cicutae Lasch., 120.
Circeae Pers., 112.
Cirsii Lasch., 155.
Clarkiae Peck, 117.
Comandrae Peck, 91.
congregata Hark., 103.
conferta Diet, and Holw., 156.
Convolvuli (Pers.) Cast., 129.
Cordylanthi Blasdale, 143.
Crepidis-accuminata Sydow, 158.
crepidicola Sydow, 157.
Cressae (DC) Lagerh., 130.
Cryptanthis Diet, and Holw., 134.
Cyanchi Lagerh., 93.
Cymopteri Diet, and Holw., 121.
Cynodontis Desm., 52.
Delphinii Diet, and Holw., 101.
Dichondrae Mont., 131.
digitata Ell. and Hark., 105.
distorta Holway, 135.
Dondiae Arth., 99.
effusa Diet, and Holw., 110.
Eleocharidis Arth., 76.
Ellisii De-Toni, 122.
Epicampis Arth., 53.
Epilobi-totragoni (DC) Wint., 113.
Eulobi Diet, and IIolw., 117.
Fendleri (Tracy and Earl) Jackson,

54.
Franseriae Sydow, 159.

Gayophiti Billings, 114.

gemella Diet, and Holw., 102.

Gentianae (Str.) Link., 128.

Giliae Hark., 132.

gilieola Hennings, 133.

globosipes Peck, 139.

glumarum (Schum.) Eriks. and

Honn., 55.
graminclla (Spez.) Diet, and Holw.,

56.
granulispora Ell. and Gall., 85.
Grindeliae Peck, 160.
Grossulariae (Schum.) Lagerh., 77.
Harknessii Vize, 161.
Helianthellae (Peck) Arth., 162.
Helianthi Schw., 16;?.
Hemizoniae Ell. and Tracy, 164.
Heucherae (Schw.) Diet., 103.
heterantha Ell. and Ev., 115.
Holboelii (Hornem) Rostr., 100.
Holcina Erikss., 57.
Holwayi Diet., 84.
Hordei Fuckel, 50.
Hydrocotyle (Link) Cke., 123.
Hypochaeridis Oud., 165.
intermedia Diet, and Holw., 113.
intermixta Peck, 166.
interveniens (Pk.) Bethel, 58.
investita Schw., 167.
Iridis (DC) Wallr., 90.
Jonesii Peck, 124.
Lagophyllae Diet, and Holw., 164.
Lindrothii Sydow, 125.
lobata B. and C, 106.
Ludwigiae (Ell. and Ev.) Holw.,

116.
luxuriosa Sydow, 59.
McClatchieana Diet, and Holw., 78.
Majanthae (Schum.) Arth. and

Holw., 60.
Malvacearum Bertero, 107.
melanconioides Ell. and Hark., 127.
mellifera Diet, and Holw., 136.
Menthae Pers., 137.
Mesnieriana Thiim., 105.
mesomegala Berk, and Cooke, 86.
Micromeriae D. and F., 137.
millefolii Fuck., 168.
Monardellae D. and F., 137.
montanensis Ellis, 61.
Moreniana Dudley and Thompson,

87.

1919]

Blasdale: Uredinales of California

153

nodosa Ell. and Hark., 88.
obtecta Peck, 79.
Oenotherae Vize, 117.
Osmorrhizae (Peck) Cke. and Peck,

126.
palefaciens Diet, and Holw., 100.
Palmeri (Anderson) Diet, and

Holw., 144.
patruelis Arth., 80.
Peckii (De-Toni) KelL, 81.
Pentastemonis Peck, 145.
plumbaria Peck, 133.
Poarum Niels., 62.
poculiformis, 63.
Polygoui-amphibii Pers., 97.
procera Diet, and Holw., 64.
punctata Link., 148.
punctiformis Diet, and Holw., 98.
purpurea Cke., 65.
recondita Dietel, 156.
Rhamni (Pers.) Wettst., 66.
Ehodiolae B. and Br., 104.
rubifaciens Johans., 149.
rufescens Diet, and Holw., 146.
sejuncta Sydow, 67.
Sherardiana Koern., 108.
Sorghii Schw., 68.
Sphaeralceae Ell. and Ev., 109.
splendens Vize, 169.
Stephanomeriae Sydow, 170.
Stipae (Opiz.) Arth., 69.
subniteus Dietel, 70.
Symphoricarpi Hark., 150.
Taraxici (Eebent.) Plowr., 171.
Toumeyi Sydow, 235.
Triticina Erikss., 71.
Troximontis Peck, 172.
variolans Hark., 173.
Veratri Duby, 89.
Violae (Schum.) DC, 111.
Wulfeniae Diet, and Holw., 147.
Wyethiae (Peck) Ell. and Ev., 174.
Xanthii (Schw.), 175.
yosemitana Blasdale, 236.
Zauschneriae Sydow, 118.
Pucciniastrum Goodyeriae (Tranz.)
Arth., 176.
Galii Fischer, 237.
pustulatum (Pers.) Dietel, 177.
Pyrolae (Pers.) Diet., 178.
sparsum (Wint.) Fischer, 179.

Ravenelia arizonica Ell. and Ev., 180.

versatilis (Peck) Diet., 181.
Roestelia interveniens, 58.

Harknessianoides Kern, 232.
Tranzschelia punctata (Pers.) Arth.,

182.
Uredinopsis Copelandi Sydow, 183.
Uredinopsis Pteridis Diet, and Holw.,

184.
Uredo Acanae Ell. and Ev., 185.

Arbuti Diet, and Holw., 179.

Beloperonis Arthur, 227.

Castilleiae Diet, and Holw., 142.

Coleosporioides Diet, and Holw., 12.

Copelandi, 179.

Gaillardiae Diet, and Holw., 186,

Nicotianae Arthur, 187.

pasadenae Sydow, 21.

Phoradendri Jackson, 188.

Sphacelicola Diet, and Holw., 189.
Uromyces abbreviatus Arth., 210.

Aconoti-lycoctoni (DC) Wint., 208.

albus Diet, and Holw., 211.

appendiculatus (Pers.) Link, 212.

Armeriae (Schl.) Lev., 225.

aterrimus Diet, and Holw., 196.

aureus Diet, and Holw., 195.

Betae (Pers.) Lev., 203.

bicolor Ellis, 196.

Bidentis Lagerh., 228.

Brodieae EU. and Hark., 197.

Caryophyllina (Schrank) Witt.,
206.

Chenopodii (Duby.) Schr., 204.

Chlorogali Diet, and Holw., 198.

Chorizanthis Ell. and Hark., 201.

Eriogoni Ell. and Hark., 201.

Fabae (Pers.) Dby., 213.

fallens (Desmaz.) Kern, 214.

Glycyrrhizae (Rabh.) Magnus, 215.

Holwayi Lagh., 199.

Hyperici-frondosi (Schw.) Arth.,
223.

intricatus Cooke, 201.

Jaeksoni Arth. and Fromme, 190.

Jonesii Peck, 209.

Junci (Desmaz.) Tul., 193.

Junci-effusi Sydow, 194.

Lilii Clinton, 199.

Limouii (DC) Lev., 226.

Lupini Berk, and Cook, 216.

Medicaginis Pass., 217.

154 University 'of California Publications in Botany [Vol. 7

nevadensis Hark., 224.
oblongua Vize, 218.
occidentalis Diet., 219.
Peckianus Farlow, 191.
rolygoni (Pers.) Fuckl., 202.
promiuens (DC) Pass., 222.
pulchellus Ell. and Ev., 207.
punctatus Schroet., 220.

Euelliae Holw., 227.
Scirpi (Cast.) Burrill, 192.
Spragueae Hark., 205.
Trifolii (Hedw.) Lev., 221.
Zygadeni Peck, 200.
Uropyxis Amorphae (Curt.) Schroet.,
229,
sanguinea (Peck) Arth., 230, 60.

VIII. INDEX TO HOST PLANTS

Abies balsamea, 7, 177; concolor, 7;

Fraseri, 7; grandis, 184; lasiocarpa-

184; magnifiea, 7; nobilis, 184.
Abronia piuetorum, 70.
Acacia Gregii, 181.
Acaena tridactyla, 185.
Achillea millefolium, 168.
Acoiiitum colunibianum, 208.
Adenostegia pilosa, 140; rigida, 140.
Agoseris barbellata, 172; hirsuta,

172; plebiea, 172.
Agropyron Parishii, 51.
Agrostis alba, 63; diegoensis, 63;

pallens, 190.
Allium falcifolium, 82; precox, 85;

serratum, 82; unifolium, 196; vali-

dum, 195, 196.
Althea rosea, 107.
Ambrosia psilostachya, 193.
Amelanchier alnifolia, 18, 19; pallida,

18, 232.
Amorpha californica, 229.
Andropogon sorghum var. halypensis,

65.
Angelica tomentosa, 122.
Antirrhinum assurgens, 141; majus,

141; nudicum, 141; Nuttalianum,

141; vagans, 141; virga, 141.
Aplopappus squarrosus, 173.
Aquilegia truncata, 51.
Arabis arcuta, 100; Breweri, 100;

Holboelii, TOO; palefaeieus, 100;

perennans, TOO.
Arbutus Meuziesii, 179.
Arctostaphylos Hookeri, 179; patula,

179.
Aristida bromoides, 49.
Armeria viilgaris, 225.
Artemisia californica, 156; Douglas-

iana, 151; dracunculoides, 151;

elatior, 151; heterophylla, 151, 156;

Suksdorfii, 151; tridentata, 151.
Asarum caudatum, 92; Lemonii, 92.
Asparagus officinalis, 83.
Aster Chilensis, 11, 152; Menziesii,

11; occidentalis, 72; radulinus, 11;

sp., 11, 72.
Astragalus leucopsis, 220; lentigino-

sus, 220; Menziesii, 220; Preussii,

220; Purshii, 220; pychnostachys,

220.
Atliyrium eyclosorum, 183.
Atriplex sp., 191.
Audibertia Palmeri, 136.
Avena barbata, 63, 66 ; f atua, 63 ;

sativa, 63.
Baccharis consanguinea, 16; pilularis,

16; viminea, 17.
Balaamorrhiza deltoidea, 153; sagit-

tata, 153.
Beloperone californica, 227.
Berberis nervosa, 230; pinnata, 54,

230; repens, 230.
Beta chicla, 203; vulgaris, 203.
Bidens expansa, 228; pilosa, 228.
Boisduvalia densiflora, 117; sparsi-

flora, 117.
Brodiaea capitata, 87, 88, 197.
Bromus carinatus, 51; hordaeeus, 51;

marginatus, 51; villosus, 51.
Calochortus albus, 84; bisceptrum,

84; elegans, 84; maweanus, 84;

venustus, 84.
Caltha Howellii, 102.
Calylptridium umbellatum, 205.
Campanula persicifolia, 9.
Carex Douglasii, 73; marcida, 80;

obnupta, 75 ; siccata, 81 ; usta, 73 ;

sp., 72.
Castilleja douglasii, 12; foliolosa, 12,

142; latifolia, 12; Martini, 12; min-

iata, 12, 13 ; Wightii, 12.

1919]

Blasdale: Uredinales of California,

155

Centromadia pungens, 10.
Ceratopteris triangularis, 21.
Chamaesarache nana, 138.
Chenopodium album, 191 ; murale, 191.
Chimaphila unibellata, 178.
Chlorogalum pomeridianum, 198.
Chorizanthe euspidata, 201; pungens,

201; robusta, 201.
Chrysanthemum indicum, 154.
Cicuta Bolanderi, 120; occidentalis,

120; virosa calif ornica, 120.
Circaea pacifica, 112.
Cirsiujn Breweri, 155; Drummondii

acaulescens, 155; edule, 155; lan-

ceolatum, 155 ; occidentale, 155 ;

quercetorun, 155; undulatum, 155.
Clarkia elegans, 117; concinna, 117;

rhomboidea, 117.
Clematis lasiantha, 51; ligusticifolia,

51 ; pauciflora, 51.
Clintonia uniflora, 86.
Collinsia bicolor, 1.
Collomia grandiflora, 132.
Comandra umbellata, 14, 91.
Convolvulus occidentalis, 129; luteo-

lus, 129; subacaulis, 129.
Cordylanthes filifolia, 143.
Crategus Douglasii, 18; rivularis, 18;

Pringlii, 18.
Crepis acuminata, 158; pleurocarpa,

158, 157.
Cressa cretica, 130.
Croton californicus, 6.
Cryptanthe flaccida, 134; Torreyana,

134; terebinthinum, 152.
Cymopterus terebinthinum, 121.
Cynodon dactylon, 52.
Cyperus eseulentus, 74,
Dactylis glomerata, 63.
Delphinium sp., 101.
Dianthus caryophyllus, 206.
Dichondra repens, 131.
Distychlis spicata, 70, 191,
Dodecatheon Hendersonii, 127; Jeff-

reyi, 127.
Dondia calif ornica, 204; intermedia,

99; multiflora, 204.
Drudeophytum Hartwegii, 125.
Eleocharis montana, 76.
Elymus condensatus, 51, 63, 64; glau-

cu3, 51; triticoides, 51.
Epicampis ringens, 53,

Ephedra calif ornica, 36.
Epilobium adenocaulon, 177; calif or-
• nicum, 177; coloratura, 177; fran-

ciscanum, 113, 177; holosericeum,

177; paniculatum, 113; sp., 89, 113.

Eriogonum cernum, 201; dumosum,

201; elongatum, 201; fasciculatum,

201; latifolium, 201; nudum, 201;

parvifolium, 201; vimineum, 201;

virgatum, 201.
Eulobus californicus, 117.
Euphorbia albomarginata, 222; poly-

carpa, 222; serpyllifolia, 222, 233.
Filix fragilis, 23.
Franseria dumosa, 159.
Funastrum hirtellum, 93.
Gaillardia aristata, 186.
Galium aparine, 148; calif ornicum,

149; triflorum, 148, 237.
Gayophytum diffusum, 114; pumilum,

114; ramosissimum, 114.
Gentiana Menziesii, 128; oregana,

128.
Gilia calif ornica, 133; capitata, 132;

gilioides, 132; pungens, 236.
Glycyrrhiza lepidota glutinosa, 215.
Godetia amoena, 117; biloba, 117;

grandiflora, 117; rubicunda, 117.
Goodyera Menziesii, 176.
Gnaphalium chilense, 167.
Grindelia sp., 160,

Habenaria dilatata, 2 ; leucostachys, 2.
Helianthella californica, 162 ; neva-

densis, 162.
Helianthus annus, 163; debilis, 163;

lenticularis, 163.
Heliotropium curassavicum, 70.
Hemizonia citrina, 164; Clevelandi,

164; luzulaefolia, 164.
Heuchera micrantha, 103.
Hieracium sp., 67.
Hordeum Gussonianum, 50, 63 ; mur-

inum, 50, 55, 63; nodosum, 61, 190;

sativum, 63.
Hydrocotyle prolifera, 123.
Hydrophyllum capitatum, 61; occi-
dentale, 61.
Hymenoclea monogyra, 169; salsola,

169.
Hypericum anagaloides, 223; Scouleri,

223.
Hypochaeris glabra, 165.

156

University uf California Pubh-cations in Botcvny [Vol. 7

Hj'ptis Enioryi, 135.

Iris alata, 90; Doufjlasiaiia. 90; fjer-

manica, 90; longipctala, 90; mis-

souriensis, 90; pumila, 90; xiph-

ium, 90.
Isomoris arboroa, 70.
Iva axillaris, 16G.
Juncus balticus, 193; losurii, 193

mexieanus, 193; nevadensis, 194

patens, 193; phaeoccphahis, 194

robustus, 193; textilis, 193; xiphi-

oides, 194.
Juniperus chinensis, 20; occidentalis,

19, 232.
Lagophylla congesta, 164.
Lamarckia aurea, 63, 66.
Larix lyalii, 29.
Lathyrus Bolanderi, 213; Jepsonii,

213; sulpluireus, 213; Torreyi, 213;

violaceus, 213.
Lavatera arborea, 107; assurgenti-

folia, 107; niaritima, 107; plevioa,

107; sylvestris, 107.
Leptotenia dissecta, 119.
Libocedrus decurrens, 18.
Liliiim columbianiim, 199; parv^um,

199; rubescens, 199; Washington-

ianum, 199.
Linanthus ciliatus, 132, 133.
Linaria amethystina, 141; bipartita,

141; delphinifolia, 141; melan-

thera, 141; marocanna, 141.
Linum Breweri, 29; congestiim, 29;

digynum, 29; drymarioides, 29;

Lewesii, 29; micranthum, 29.
Lolium multiflorum, 66; perenne, 66.
Lotus eriophorus, 220; glaber, 220.
Ludwigia natans, 116.
Lupinus albifrons, 216; Chamissonis,

216, Douglasii, 216; formosus

Bridgesii, 216; latifolius, 216,

219; leptophyllus, 219; rivularis,

216.
Lycium Andersoni Wrightii, 139;

oaliforniciim, 139.
Lygodesmia spinosa, 161.
Madia anomala, 10; capitata, 10;

dissitiflora, 10; elegans, 10; Nut-

talii, 10; sativa, 10.
Malva borealis, 107; crispa, 107;

moschata, 107; oxyloba, 107; par-

viflora, 107; sylvestris, 107.

Malvastnim ambigua, 58; splendidum,

108; Thurbcri, 58, 108.
Mcdicago lupuliua, 217; sativa, 217.
Mentha canadensis, 137; sativa, 137.
Micromeria Chamissonis, 137.
Microsteris gracilis, 133.
Monardella lanceolata, 137; micran-

tha, 137; odoratissimum, 137; viri-

dis, 137; villosa, 137.
Navarretia atractyloides, 132.
Nicotiana Bigclowii, 187.
Notholcus lanatus, 57.
Oenanthe californica, 34, 192.
Oenothera gauraeflora, 117; stringu-

losa, 117.
Onagra Hookeri, 81.
Oryzopsis hymenoides, 58, 69.
Osmorrhiza brevipes, 126; nuda, 126;

occidentalis, 126.
Panicularia pauciflora, 63.
Pasania densiflora, 15.
Pediciilaris semibarbata, 146.
Pellaea andromedafolia, 21.
Penstemon azureus, 145; Bridgesii,

145; confertus, 144; deustus, 145;

Menziesii, 234; Newberryi, 144;

Eoezli, 145; spectabilis, 235.
Peucedanum dasycarpum, 124; Has-

sei, 124.
Phacelia californica, 61 ; memoralis,

61; tanacetifolia, 61.
Phajus Wallichii, 8.
Phalaris californica, 60; minor, 60.
Phaseolus lunatus, 212; vulgaris, 212.
Phleum pratense, 63.
Phoradendron longispicum, 188.
Pinus attenuata, 15 ; contorta, 12, 13,

15; muricata, 12; ponderosa, 13, 14,

15, 12; radiata, 15, 35; Sabiniana,

12.
Poa annua, 62; pratensis, 62, sp., 50.
Polygonum acre, 97; amphibium, 97;

aviculare, 202; bistorta, 96; Davis-

iae, 95 ; erectum, 202 ; hydropiper-

oides, 97 ; 1 a p a t h i f o 1 i u ni, 97 ;

Muhlenbergii, 97; Newberryi, 95,
Polypodium californicum, 22.
Polypogon monspeliensis, 66.
Populus alba, 30; Frcmontii, 30;

tremuloides, 27; trichocarpa, 27,

30.

1919]

Blasdale: Uredinales of California

lOi

Potentilla Blaschkeana, 40; Elmeri,

40; fruticosa, 37; glandulosa, 40;

Hallii, 40.
Primula suffretesceus, 224.
Prosopis juliflora, 33, 180.
Prunus sp., 113.
Pseudotsuga mucronata, 27.
Psoralia physoides, 210.
Pterdis aquilinum pubescens, 184.
Ptiloria exigua, 161; carduacea, 170.
Pyrola pieta, 178; secunda, 178; uli-

ginosa, 32.
Pyrus sp., 20.
Quercus agrif olia, 15 ; dumosa, 15 ;

Kellogii, 15.
Eanimculus calif ornicus, 92; flam-

mula, 209; sp., 50.
Ehamnus crocea, 105; ilicifolius,

105; insularis, 105.
Rhododendron californicum, 31.
Rhus diversiloba, 44.
Ribes divarieatum, 77.

tenuifiorum Lindl., 231.
Rosa gymnocarpa, 26, 41, 43; califor-

nica, 43 ; minutif olia, 43 ; sonomen-

sis, 43 ; sp., 107.
Rubus leucodermis, 39; parviflorus,

42; vitifolius, 24, 25; sp., 108, 104.
Rumex aeetosella, 94; hymenoceph-

alus, 98; occidentale, 94; persi-

carioides, 94; salicifolius, 98.
Salicornia ambigua, 191.
Salix exigua, 28; laevigata, 28; lasi-

andra, 28; lasiolepis, 28.
Salvia mellifera, 136.
Scirpus americanus. 79; calif ornicus,

79, 192 ; paludosus, 192 ; lacustris

occidentalis, 79 ; microcarpus, 78 ;

Olneyi, 79; pacificus, 192.
Sedum sp., 104.
Selinum pacificum, 34,
Senecio Douglasii, 5.
Sida hederacea, 106, 107.
Sidalcea asprella, 58; malvaeflora, 58;

spicata, 108; sp., 109.
Silene Douglasii, 207.
Solidago californica, 11 ; confinis, 11.

Spergularia Clevelandii, 70; macro-

theca, 70.
Sphacele calycina, 189.
Sphaerostigma hirtella, 117; micran-

tha, 117; spirale, 117; viridescens,

117; Veitchianum, 117.
Sporobolus airoides, 59.
Statice linionium californicum, 226.
Stephanomeria chicoriaceum, 170;

lactucina, 170; runcinata, 170.
Stipa eminens, 56, 69; pulchra, 58,

i59 ; setigera, 69 ; speciosa, 58.
Sjmphoricarpos racemosus, 46, 150.
Synthyris rotundifolia, 147.
Taraxia ovata, 115; graciliflora, 115.
Taraxacum Taraxacum. 171.
Tellima affinis, 103; grandiflora, 103.
Tetradymia glabrata, 173.
Tlialictrum Tendleri, 45, 51.
Thelypodium lacinatum, 70.
Trifolium albopurpureum, 218; cilio-

latum, 218; depauperatum, 218;

dubium, 218; gracilentum, 218;

Macraei, 218; microcephalum, 218;

microdon, 218; oliganthum, 218;

pratense, 214; repens, 221; ros-

cidum, 218; stenophyllum, 218;

tridentatum, 218; variegatum, 218.
Triglochin concinna, 3.
Urtica gracilis, 75.
Vaccinium Chandleri, 7; membrana-

eeum, 7; ovatum, 7; parvifolium, 7.
Velaea arguta, 125.
Valerianella congesta, 4.
Veratrum californicum, 89.
Vicia americana, 211; californica,

211; truncata, 211.
Viola adunca longipes, 111; glabella,

111; lobata, 110; nephrophylla,

111; ocellata, 110.
Xanthinum canadense, 74, 175.
Wyethia angustifolia, 174.
Zauschneria californica, 118.
Zea mays, 68.
Zonanthis corjTnbosa, 10.
Zygadenus Fremonti, 200.

A RUBBER PLANT SURVEY

OF

WESTERN NORTH AMERICA

I. CHRYSOTHAMNUS NAUSEOSUS AND ITS VARIETIES

HARVEY MONROE HALL

11. CHRYSIL, A NEW RUBBER FROM CHRYSOTHAMNUS
NAUSEOSUS

HARVEY MONROE HALL and THOMAS HARPER GOODSPEED

III. THE OCCURRENCE OF RUBBER IN CERTAIN
WEST AMERICAN SHRUBS

HARVEY MONROE HALL and THOMAS HARPER GOODSPEED

UNIVERSITY OF CALIFORNIA PUBLICATIONS

IN

BOTANY

Vol. 7, No. 6, pp. 159-181 . November 7, 1919

o~>

I. CHRYSOTHAMNUS NAUSEOSUS AND ITS

VARIETIES

HARVEY MONROE HALL ,'2 .
QAHDEN

CONTENTS

PAGE

I. Tntioduction and acknowledgments 159

IL Key to the varieties of Chrysothamnus nauseosun 163

III. Synopsis of the varieties of Chrysothamnus nauseosus 165

IV. Index of specific and varietal names 181

I. INTRODUCTION AND ACKNOWLEDGMENTS

In the course of an investigation of Chrysothamnus as a possible
source of rubber, it has been found necessary to give considerable
attention to the botanical relationships of the species and varieties of
this genus of shrubs. Even before that investigation was begun in
1917, the author had undertaken a revision of the group and although
the preparation of this has been delayed by the rubber studies it is
intended now to carry it forward to completion. In the meantime,
however, it is necessary that the forms discussed in the rubber report
be properly defined and that those not hitherto recognized be named
and described. These are the rea.sons for tlie publication of this
preliminary paper.

Chrysothamnns is a genus of the Aster Tribe of the Compositae
and belongs to what is often called the Haplopappus group. From
related genera it differs chiefly in the more cylindraceous involucre,
the bracts of which are usually more or less keeled and tend to fall
into five vertical ranks. Although these characters are too variable
to give to the genus that sharpness of delimitation that one would like,
yet there can be no question as to its homogeneity. It represents what

160 I'liiiursili/ of California I'liblicutiotis in Botanij | Voi- 7

is pcrluips till' most sat isfiH'tory result ot" all of the iiiiincroiis a1t(iii|)ts
to set up genera in this <iroii|) where generic limits are admittedly
most diffieult to estahlish.

Within the genus we find a multitude of natural forms of greatei-
or li'ss siguitieanee. This has led botanists greatly to increase tlie
num])er of si)eeies and vai'ieties, so that not less than one hundred
and thirt<'en of these have been described and this numbei- could l)e
multij)lied several times through more extensive field studies and
witliout the employment of characters other than those alreadx' pro-
posed for the basis of species. This is partly because these single
characters have been vai"iously uniti'd in nature to form a largt'
nuiid)er of combinations, but it is also because of the extreme varia-
bility of some of the chai'acters themselves. The ninnerous forms of
Chrijsothainims may be assembled into five natui'al groups as follows:

7\EV TO TITP: RECTTOXS of CHFYSOTHAMiXCS

Tr('rl)aije not it'siiious-pum-tatc: leaves oblauceolate to nar-
rowly linear or terete.
Heads in leafj' spike-like or raceme-like clusters: outer
bracts of the involucre prolonged into a slender

herbaceous tij) or a])pendage Section 1. Parryani

(Principal species: Parri/i, Hownrdi.)
Heads cymose, paniculate, or solitary at the ends of the
branches: bracts obtuse to acuminate, devoid of
herbaceous tip.
Twigs not tomentose, either smooth and glabrous or
only scabrid or puberulent: style-appendages from
included to long-exserted.
Bracts strongly keeled, in five very distinct vertical

ridges Section 2. Pui.chem.i

(Principal species: pulehellus, depressii,i.)
Bracts scarcely keeled and the vertical ridges obscure. .Section :!. Tvpici
(Principal sj)ecies: <ilhidiin, (irreiici. (iroinincns.
Vasciji, viscidifioius. )
Twigs closely covered with a pannose tomentum which
usually persists for several years: style-appendages
mostly longei- than tlie stigmatic portion and long-
exserted Section 4. Xaiseosi

(Single species: dohscosu^s.^)

Herbage resinous-punctate: leaves terete Section .">. I'i'sctati

(Principal species: teretifolius, paniculatus.)

Of the at)ove five grou|)s rubber has been found in more than
minute quantities only in the NauseosiaLwd I'unctati. Since the latter

1 It is possilde tiiat C. formosu^ Greene and C. turbiiintu.f (M. E. Jones)
Rydb. constitute additional species of this grou]i. See p. ISO.

1919] Hall: Chrysotha})iiius tiauseosus and Its Varieties 161

of these groups comprises only two species, and since these two species
are alreach' well defined in the literature and are not inclined to
break up into numerous variants in nature, it is at the present time
necessary to consider only the nauseosus group.

Taking Chrysothatiums nauseosus as comprising all of the forms
of the Nauseosi, as indicated in the above key. we find its best dis-
tinguishbig character to be the presence of a most remarkable pannose
tomentum on the young twigs. It is fully recognized that pubescence
characters often furnish but a very treacherous basis for constant
species, yet in this instance the nature of the -pubescence is so unlike
that in any other species of the genus, and it is so obviously correlated
with habit and other minor characters that it seems certainly to indi-
cate a natural group. This felt-like covering to the twigs is more or
less infiltrated with a resinous substance. In some forms the surface
is loose and fluffy, the tomentum thus more or less completely masking
the striae of the stems. In these forms the surface is usually light
gray or dull white or even almost pure white, and the loose tomentum
extends even to the involucres. The original C. )iauseosus is one of
these forms. In other varieties the surface of the tomentum is firm
and smooth, thus revealing the striae. In these forms the twigs are
dull to bright yellowish green in color or rarely somewhat whitish
and the involucres are mostly glabrous.

Within what is here included under C. nauseosus, previous writers
have described forty forms, all but six of which have been accorded
specific rank at one time or another. In the present paper two new
varieties are added. Nothing can be more certain than that these
forty-two attempts to recognize species and varieties do not by any
means exhaust the resources of the group. Everj'- autumnal excursion
into a new district brings to light one or more forms not previously
described. The only limits set to the number of new species or
varieties whicli might be set up lie in one's ability to visit all parts
of the field during the flowering period and the failure or disin-
clination to recognize minute variations. Yet the systematist should
include in his ultimate object not only the recognition of this multi-
tude of forms but also their proper arrangement in a scheme which
will display their natural relationships. Since this would entail an
enormous amount of detailed labor, including extensive experiments,
and since the results, even if attainable, would be of but little practical
value at the present time, the writer has satisfied himself with the
acceptance of twenty-two forms, all of which are treated as varieties.

162 Univcrsify of California. Puhlications in Botany \yoh.7

This is believed to provide for all of the principal forms, and it is
quite certain tliat eacli of the described varieties is a natural unit,
althoufrh in most cases it is itself made up of still smaller variants.

Altliougii tlu' relationships between the varieties accepted in this
paper are expressed somewhat in the arranj^ement, and in the form
of brief notes from time to time, yet our actual knowledge of these
matters is so slight tliat it seems unwise to attempt an expression of
it through an elaborate system of polynomials. Some of the varieties
are based upon characters which are at least partially heritabl(% while
others are doubtless ecologic forms. In some cases two varieties
occupying the same territory may be easily distinguished by a number
of characters which at that locality do not overlap in their variations;
yet as forms from other places are brought into comparison, it is
discovered that these characters are so variable that they cannot be
used for the recognition of species in the broad sense. It seems almost
certain that after all of the forms shall have been assembled it will
be found that no two of the characters thus far proposed for the
separation of "species" in this group wall be found to vary in unison,
and furthermore that there is not a single one of them that does not
vary by imperceptible degrees. Therefore we need not be surprised
ultimately to find as man}- forms as there are possible combinations
of characters and to have, in addition, numerous forms which cannot
be satisfactorily^ placed because of the intermediate nature of one
or more of the characters. It is thus seen to be impossible to accord
specific rank to any of the units of this exceedingly complex and
variable assemblage of forms unless one is willing to accept a species
concept so narrow that its usefulness would be lost because of the
impossibility of definite application.

If some readers object to the use of even trinomials we woidd
suggest that the binomial indicating the inclusive major species,
that is, Chrysothamniis nauscosus in this case, is all that need be
used in most instances, while in any special paper on the group the
varietal name alone may be used without confusion, since the generic
and specific names are understood. This practice will be followed
in the report on Chrysothamnits as a possible source of rubber, in
which paper the actual w^orking out of the method may be observed.

In preparing the following synopsis, the presence or absence of
hairs on the involucre is taken as the principal character, notwith-
standing its variable nature. This is done because it comes the nearest
to assembling the forms into what seem to bi' natural groups, or sub-

1919] Hall: Chrysotlnnnnus nmiseosus and Its Varieties 163

species. Every other character tried, such as size or shape of the
leaves or of the parts of the involucre or flower, the nature of the
pappus or style-branches, and the pubescence of the akenes, have all
been found to be even more variable and not to serve well as a basis
for a natural classification. Moreover, the pubescent involucre is
nearly always associated with a white or grajdsh appearance of the
foliage so that the use of this character divides the whole species into
two series of varieties, which series may, for the most part, be recog-
nized by their general appearance.

After the foregoing remarks as to the variability of characters in
this species it will perhaps be understood that one cannot expect to
use the key to varieties with entire satisfaction. The large number
of intermediate forms and the numerous permutations of characters
render it impossible to prepare a key whereby one may definitely
identify every specimen that comes to hand. It is believed, how-
ever, that with the exception of one or two little-known forms of
eastern Oregon and Washington, all of the major variants have been
accounted for.

This study has been greatly facilitated through the assistance of
a considerable number of people but since these will be specifically
mentioned in the report on the rubber plant investigations the list
need not be given here. AcknowledgTuent should be made at this
place, however, of the kindness of Dr. B. L. Robinson, of the Gray
Herbarium of Harvard University, and also of that of Dr. P. A.
Rj-dberg, of the New York Botanical Garden, in permitting the
author to examine certain type specimens in their charge. Further-
more, Dr. Aven Nelson has made available the rich collections of the
Rocky Mountain Herbarium at the University of Wyoming.

II. KEY TO THE VARIETIES OF CHRYSOTHAMNUS

NAUSE0SU8-

Series A. The Gray Forms
Involucres tomentulose (at least the short outer bracts) to densely woolly:
foliage mostly gray or even white with a rather copious pubescence, this either fine
and close or floccose-tomentose (least pronounced in oreophil'us and speciosus).

Akenes densely strigose.

Tomentum loose, copious, and nearly pure white on twigs and leaves, extending
to the iuvolucies: corolla-tube nearly always arachnoid-pubescent.

-See also doubtful forms on p. 180.

104 riiirn-sih/ of CaUfoviiid I'lihllcdtioiis in BiAinnj I V<'i.. 7

Inner bracts of iiivnliicrc ])l;iiiily tomciitosc oi' if iiciirly ^lahioiis \\\v bracts
then acute.
Corolla-lolics liniccolnti'. 1 to 12 iiini. loiiy : iiivdiucral bracts acute (at least

in part) !• iKiuscnsus s. str.

Corolla hilx's sliort-ovate, less than 1 mm. lon<i: bracts all very obtuse.

2. Iiololeucus
Inner bracts smooth ami {rlabrous or marly so, very obtuse .....3. lot isqiui mens
Tomentuni close, compact, smooth, and usually >irayish on twig-s and leaves:
corolla-tube jjlabrous or puberulent. not arachnoid-pube.scent.

Leaves A to (5 nun. wide: bracts very obtuse 4. salicifolius

Leaves less than :> mm. wide: bracts various but mostly acute.

Bracts abruptly acute. Far western fi. nrridnifuli.s

Bracts "gradually acute or somewhat obtuse.
Involucre 7 to 9 or rarely 10 mm. hi<);h.

Shrubs normally .") to 20 dm. hiyh: corcdla 8 to 10 mm. loun or rarely

only 7 mm. in {itiapJuiUidcs {latisquainrus of Arizona and New

Mexico, with broad and very obtuse and smooth inner bracts might

be sought here ).

Leaves of flowering twigs 1 to '.', cm. long, not numerous: herbage

usually fragrant ='>. giuiphaUxhs

Leaves of flowering twigs mostly over 3 cm. long, often crowded:

herbage not fragrant 7. specios'm

Shiubs only 2 to H dm. high: corolla H to 8 mm. long (very rarely
9 mm.).
Leaves erect or ascending: bracts all tonientulose or ciliate..8. frigidus
Leaves spreading or recurved : oidy the outer bracts tomentulose.

9. plattensis

Involucre 11 to 12 mm. or rarely only 10 mm. high 10. bernardinu/i

Akenes glabrous.

Bracts of the involucie acuminate 11. Bigelovii

Bracts of the invohuic obtuse 12. glareosus

Series B. The Green Forms

Involucres perfectly glabrous although .sometimes viscidulous or glaudular:
foliage mostly greenish, the tomentum rather s])arse or wanting on mature leaves.

Akenes glabrous 1 3. Irinsperm us

Akenes densely strigo.se.

Leaves linear, mostly more than 1 mm. wide,-"! mostly 3 to ."-nerved e.xcept in
bfnuirdinu,s.
Involucre 6 to 8 mm. long: corolla (i to 9 mm. long; lobes 1 mm. or less

long 14. graveolenft

Tiivoluric 9 mm. or nnire long: coiolla 9 to 10 mm. long; hdies 1 mm. or more
long. Western foi'ms.
Heads 8 to 10-tlowercd : leaves flat or twisted, 2 to •") nnii. wide.

15. calif ornicus
Hea<ls .j-flowered (rarely 6-flowered) : leaves often conduplicate, 1 to 2 mm.

wide 10. beniardinus

Leaves linear-filiform or very narrowly linear, mostly 1 mm. or less wide (except
in occidentalifi) , 1 -nerved.

'■'■ Care must be exercised in making this measurement since the leaves are often
longitudinally folded. The figures given are for the total width of the flat leaf.

1919J Hall: Chrysothamnus naiiseosus and Its Varieties 165

Bracts of the iuvoliK-re abruptly pointed or some ouly acute. Far western.

Tip of the bract short, erect 6. occidentalis

Tip of the bract about 1 mm. long, very slemler, recurved.. -.16. renimiiwsu.s
Bracts obtuse to acute, not abruptly pointed.
Corolla-lobes glabrous.

Involucre 6 to 9 mm. long, not sharply angled.

Bracts ciliate, the outer often slightly hairy also on back, flat, thin,

scarcely keeled 17. oreopJiihts

Bracts not ciliate, all perfectly glabrous, concave, somewhat keeled.
Corolla 7 to 8.'.j mm. long; lobes under 2 mm.: twigs ami foliage
slender.
Lobes of corolla O.-j to 1 mm. long, rarely more : inflorescence
typically rounded to pyramidal. Rocky Mountain states.

18. pinifoUus
Lobes of corolla 1 to 2 mm. long: inflorescence typically cylindric

to pyramidal. Great Basin 19. coiislmiUs

Corolla 8 to 10 mm. long; lobes 1.7 to 2.5 mm. long: twigs an<l

foliage stouter. Southwestern 20. viridulus

Involucre 9 to 10 mm. long, shai-ply 5-angled, the strongly keeled bracts.

in very distinct vertical rows 21. mohavensis

Corolladobes sparsely long-hairj' in the bud: shrub nearly leafless.

22. junceus

III. SYNOPSIS OF THE VARIETIES OF CHRYSOTHAMNUS

NAUSEOSUS

1. Chrysothamnus nauseosus (Pall.) Britt. {sensu strietii)
in Britt and Br., 111. FL, vol. 3, p. 326, 1898.

Chrysocoma nauscusa Pall., in Pursh, Fl., vol. 2. p. .117, 1814.
Chri/sotlunrmus speciosus albicaulis Nutt., Trans. Am. Philos. Soc, ser. 2,

vol. 7, p. 324, 1841.
Bigcloria graveolens albicaulis Gray, Proc. Am. Acad., vol. 8, p. 64.j, 1873.

Shrub 5 to 10 dm. hig-h, with numerous erect or ascending- twigs,
leafy to the top, permanently white-tomentose throughout ; twigs not
evidently striate : leaves narrowly linear, 3 to 6 cm. long, 0.5 to 1.5
mm. wide : inflorescence a rather loose round-topped cyme : involucre
7 to 8 mm. high ; bracts mostly acute, plainly keeled, in 5 distinct
vertical ranks, white-woolly but not ciliate: corolla 7.5 to 9 mm. long;
tube perhaps always cobwebby with long weak liairs; lobes lanceolate,
1 to 2 mm. long.

This type form of the species is much less common than numy of
its varieties. It occurs from Utah to Oregon and ])r()bal)ly north to
British Columbia and J\Iontana, inhabiting well draiiK^l soil with
little or no alkali. In the typical form, as understood by the writer,
the leaves are very narrow and, like the twigs, are entirely covered
with a white flocculent tomentum, while the corolla-tube is conspicu-

166 UnivcrsUu of California Publications in Botany fVoi,. 7

ously araclinoid-pubcscciil. Sut'li arr .spi'ciiufns gatluTod by Leiberg
in eastern Washington uiulcr no. 884 and others by Marcus E. Jones
at Marysvah% Utah, uiidcf no. 5968. In northern Mono County, Cali-
t'ornia. is encountered a form in vvliicli the leaves are 2 to 3 mm. wide
and the inflorescence is more compact and rounded (H. M. H., no.
468), but in the pubescence of the corolla-tube and in all other
eiiaracters than those mentioned it is plainly nauseosus. Nearly
identical are specimens from Truckee, California (Ileller, no. 7192).
Another divergence is indicated by specimens with the broad leaves
and other characters of the Mono and Truckee collections just cited
but with a corolla-tube which is only crisp-pubescent as in most
varieties. A recognition of these forms would lead only to confusion
since further field studies would doubtless reveal still other divisions
that might be made. They are therefore retained in this paper as
only trivial variants of nauseosus. C. orthophyUus Greene, Pitt.,
vol. 5, p. 62, 1902, known only from Plumas County, California, is
described as less than a foot high and with the tips of the corolla-
lobes as well as the tube long-villous.

2. Chrysothamnus nauseosus var. hololeucus (Gray) comb. nov.

liigclovia (/raveolens var. Jiolulcuca Gray, Proc. Am. Acad., vol. 8, p. 64-1,
1873.

Shrub 6 to 18 dm. high and fully as broad, closely branched and
usuall}' of rounded outline, leafy throughout : herbage exceedingly
fragrant, densely clothed with a nearly pure white tomentum which
completely masks the striae of the stem and extends even to the
involucres : leaves 1 to 3 cm. long, about 1 but sometimes nearly 2 mm.
wide : inflorescence a rounded often compact cyme : involucre 6 to
7 mm. high ; bracts very obtuse, plainly keeled and in 5 distinct ver-
tical ranks, woolly but not ciliatc: corolla 6.5 to 8 mm. long; tube
more or less cobwebby with loose hairs or these occasionally wanting;
lobes ovate, acute, strictly erect, 0.5 to 1 mm. long.

The type of hololeucus came from Owens Valley, California,
whence we now have collections from a number of stations. What
appears to be the same form occurs as far south as Antelope Valley
and very good hololeucus grows as far north as Pyramid Lake, Nevada,
but it does not have a wide east-and-west range. It is confined to loose
gravelly or sandy well drained slopes and even in such places it never
forms pure stands but occurs as scattered individuals among bushes
of other sorts, commonly C. n. gnaph^lodes and Artemisia iridentata.
The plants are so white in comparison with these other shrubs that
they can be distinguished in the field without difficultv.

1919 J Hall: Chrysothamniis nauseosus and Its Varieties 167

3. Chrysothamnus nauseosus var. latisquameus (Gray) comb. nov.

Bigelovia graveolens latisquamea Gray, Proc. Am. Acad., vol. 8, p. 64-3,
1873.

Shrub tall (perhaps rounded at top), leafy to summit: herbage
white with a rather loose tomentum : leaves 2 to 5 cm. long, less than
1 mm. wide : inflorescence a loosely branched rounded compound
cyme: involucre about 8 mm. high (7 to 9 mm.); bracts carinate,
about 4 in each of the 5 distinct vertical rows, outermost ones tomen-
tulose, inner ones very obtuse and usually glabrous : corolla about
9 mm. long ; tube short-pubescent or glabrous ; lobes ovate or short-
lanceolate, 0.4 to 0.9 mm. long, erect.

A common variety of New Mexico and Arizona. Bigelovia graveo-
lens appendiculata Eastw., Proc. Calif. Acad. Sci., ser. 3, vol. 1, p. 74,
1897, from the White Sands of New Mexico, is an abnormal form
with one to four linear appendages on the corolla-tube. The tube is
arachnoid-pubescent and the lobes are only about 0.4 mm. long, both
of which characters suggest an affinity with hololeucus. C. arizonicus
Greene, Pitt., vol. 4, p. 42, 1899, is a form with the corolla-tube ' ' cleft
rather deeply. ' ' The type is from the Santa Rita Mountains.

4. Chrysothamnus nauseosus var. salicifolius (Rydb.) comb. nov.

Chrysothamnus salicifolius Rydb., Bull. Torrey Club, vol. 37, p. 130, 1910.

Shrub 3 to 10 dm. high, with erect branches: leaves 3-nerved, 4
to 8 cm. long, 3 to 6 mm. wide, minutely tomentulose : inflorescence
cymose, dense : involucre 7 to 8 mm. high ; outer bracts slightly tomen-
tulose, the inner glabrous and very obtuse : corolla about 10 mm.
long ; lobes 1.5 to 2 mm. long.

Apparently a rare variety and confined to Utah ; known to the
writer from only two collections, namely, Strawberry Valley, at 7000
feet altitude, F. E. Leonard, no. 288 (type), and near Salt Lake City,
A. 0. Garrett, no. 2455.

5. Chrysothamnus nauseosus var. gnaphalodes (Greene) comb. nov.

Chrysotliamnus speciosus var. gnaphaJodes Greene, Eryth., vol. 3, p. 110,
1895.

Shrubs 5 to 15 or rarely 25 dm. high, usually globoid in outline,
with very many short twiggy branches, not densely leafy: herbage
very fragrant, gray with a closely packed tomentum, the twigs
obscurely striate : leaves 2 to 4 mm. long, 1 mm, or less wide, those of
the end twigs only 1 to 3 cm. long and often recurved : inflorescence
a rounded cyme terminating each of the twigs : involucre about 7 mm.
high ; bracts rather obtuse, keeled, in 5 distinct vertical rows, tomen-
tose, not ciliate : corolla 7 to 8 mm. long ; tube sparsely pubescent with
, short crisp or rigid hairs ; lobes ovate, acute, erect or even eonnivent
around the stamen-tube, 0.5 to 1 mm. long.

168 Universih/ of Califoniia I'lihJicdiioiin in Botany [Voi-. 7

This is till' roimiKiii ^ray \ari('ty ni' ^ji-a\cll\- or saiidx" iioii-alkaliiic
slopes and lunches of western Xeva(ht and eastern ( 'alit'oniia fi-oiii
Pyramid Lake south to the Cajon, Sole(hid, and Tehaehapi i)asses, and
to Arizona. It lieh)n<:s I'spceiallx to the h)os(" soil of well drained sloi)os
suri'ounding valleys the alkaline liottonis of which ai-e oeeiipifd by
riridnlns, and often forms belts of considerable extent. It is the
most common shrub on land from whicli saj>ebnisli has been burned
or otherwise cleared, and is subclimax to that abundant shrub.

(). Chrysothamnus nauseosus vai'. occidentalis (Greene) Hall
I'niv. Calif. Publ. Hot., vol. :{. !>. (iO, 1907.
Chrysdt lid limns caiifoniicus occichiitdlis Grociw, VA-yih.. vitl. ?,. p. 112, 1895.

Shrub probably rathei' low, with numerous short slender erect
bi'auches, the leafy stems g-ray-tomentose but not loosely or fioeeulently
so: leaves narrowly linear, 4 cm. or less long, mostly less than 1 mm.
wide, but occasionally up to 2 mm., 1-nerved, tomentulose : intior-
eseence compactly eymose, rounded, 2 to 5 cm. across: involucre 7 to
9 nnn. high; at least some of the bracts abruptly acute or cuspidate,
the outer ones more or less glandular-pubei-ulent : corolla 8 to 9 mm.
long; lobes lanceolate-linear, 1.7 to 2.5 mm. long.

The distribution of this variety was originally stated by Greene to
be "In the Coast Range, from Humboldt County (California) south-
ward."" ]iater, this same author stated it as "Kern and Santa Bar-
bara counties."^ This restriction in the adoi)ted range was perhaps
due to the fact that certain specimens from Hum])oldt and oth(M-
northern counties do not meet the recpiii-ements of the description as
well as those from further south. The cuspidate bracts and long
corolla-lobes, together with the habit (especially- the small compact
rounded inflorescence), are here taken as the most satisfactory char-
acters for the variety. Accepting this definition, we find fairly typical
collections from the dry inner north Coast Ranges and from the
southern Sierra Nevada and San Bernardino ^Moiuitains. North and
east of Trinity County it apparc^ntly passes into spirit/sKs, from wliich
it scarcely differs save in the more nearly glabrous and abruptly
l)ointed bi'acts. At its southei'innost stations it meets and perhaps
merges into bernardi)ii(s.

The following are tak(Mi as typical : near sunnnit of South VoUa
Bolla, Trinity(?) County, October, 1916. Meri-iam and Bailey; Little
Kern River, Tulare ('(rnnty. C. A. Purpus, no. 2040; Barton Flats,
in the San Bernardino ^Mountains, Mrs. Wildei-, no. 597 (corolla up

4 Greene, Fl. Vr.. 18J)7. ]>. :'.(i!).

1919] Hall: Chrijsothamnus nmiseosus and Its Varieties 169

to 10 mm. long). C. iorinosus Greene, Pitt., vol. 5, p. 63, 1902, is
aparently a form intermediate between this and speciosus. It is
described as having tortuous flowering branches, nearly filiform more
or less spreading leaves which are also tortuous, and rather pungenth'
acute bracts. The types came from Plumas County- and Mount Shasta,
California.

7. Chrysothamnus nauseosus var. speciosus (Xutt.) comb. nov.

Chrysothamnus speciosus Nutt., Trans. Am. Pliilos. Soc, ser. 2, vol. 7, p. 323,
1841.

Shrub commonly 6 to 20 dm. high, broad and rounded, leafy to
summit : twigs greenish white, the tomentum comparatively thin and
smooth : leaves 2 to 6 cm. long, typically about 1 mm. wide but varying
to 3 mm., usually erect or ascending, becoming dense towards the
inflorescences and there scarcely reduced in size, gray, tomentose or
the tomentum partly deciduous and the foliage then greenish : inflor-
escence a round-topped or somewhat elongated cyme, commonly loose :
involucre 7 to 10 mm. high ; bracts 3 or 4 in each row, acute, concave,
tomentulose on the back, not ciliate : corolla 8 to 10 mm. long ; tube
sparsely puberulent or glabrous ; lobes 0.8 to 2 mm. long.

The above description is drawn to include a number of forms.
In this broad sense speciosus has a range from Idaho and Utah to
California and Washington. It is a variety belonging to sandy slopes
and benches with little or no alkali. The type came from "the Rocky
^Mountain plains, near Lewis River" and was described as having
narrow, linear, acute, more or less tomentose leaves and heads in
dense, conglomerate, terminal clusters. A form which answers to
this but with a slightly elongated and loose inflorescence is common
from northern Utah to eastern Oregon and northern California where,
through the nearly glabrous character of the involucre it passes into
occidentalis. Further south in eastern California and western Nevada
the involucre becomes more compact and nicely rounded but in this
form the comparatively rigid leaves are 2 to 3 mm. wide and vary
from gray to yellowish green. This type was apparently included by
Dr. Greene in his californicus. In the absence of better characters
and with our scant knowledge of the real speciosus it seems unwise
to segregate these forms more definitely at present, although they are
more striking than many of the Rocky ^Mountain varieties accepted
in this paper. C. pulcJurritnus, A. Nels., Bot. Gaz., vol. 28, p. 370,
1899, is a form of the high plains of the Rocky Mountain states,
especially in moist soil. All of the characters used to differentiate
it from speciosus are variable and no two of tliera vary in unison, but

170 Univcrsitij of California Publications in Botany [Vol. 7

the more nearly jjlabrous nature of the involucre seems to be the most
important. As pointed out by Dr. Nelson, it also resembles graveolens,
but the narrower leaves are constantly one-nerved and the involucre
is not absolutely jrlabrous. C pulcherrimus fasci-culatus A. Nels.,
I.e., is described as having numerous short branchlets and luimerous
rigid leaves only 2 to )} cm. long. It has been collected at Boulder
Creek and at Creston, botii of these localities being in Wyoming.

8. Chrysothamnus nauseosus vai-. frigidus (Greene) comb. nov.

Chrysothamnus frigidan Cirecue, Eryth., vol. 3, p. 112, 1895.

Sub-shrub woody only at the base, often nearly prostrate, 2 to
6 dm. high, with mostly erect stems, very leafy: twigs whitish with a
smooth close tomentum : leaves 2 to 5 cm. long, 1 to 1.5 mm. wide,
mostly erect or ascending, white-tomentose : inflorescence cyraose or
elongated : involucre about 7 mm. high ; bracts tomentulose and some-
what glandular, ciliolate at least at summit: corolla 6 to 7 mm. long;
tube puberulent; lobes linear-lanceolate, 1.2 to 1.5 mm. long.

This variety belongs to the elevated bleak plains of the Rocky
Mountain states, is especially common on the Wyoming plateaus, and
is said to occur as far north as Alberta. In one collection (Aven
Nelson, no. 2787, from Laramie) the corolla-tube is well provided
with a long loose cobwebby pubescence, just as in the otherwise very
ditferent nmiseosus proper and in Jiololeucus. C. frigidus concolor
A. Nels., Bot. Gaz., vol. 28, p. 371, 1899, is a form w^ith yellowish
green herbage and somewhat elongated inflorescences. It grows in
sandy more or less alkaline soil in Wyoming and Montana. C. pallidus
A. Nels., I.e., p. 372, is a form also allied to frigidus and perhaps to
be united with it. The twigs are less leafy except near the top, where
they are shorter and more crowded ; the herbage is nearly wiiite with
a close persistent tomentum. It inhabits alkaline soil in Wyoming
and northern Colorado.

9. Chrysothamnus nauseosus var. plattensis (Greene) comb. nov.

Chrysatlianuius spcciosus{J') plnttensis Greene, Eryth., vol. 3, p. Ill, 1895.

Characters as given for frigidus except that the narrower leaves
are loosely spreading or even recurved and that oidy the outer bracts
are tomentulose.

A form of the alkaline plains along the eastern base of tiie Rocky
Mountains. The type was described as having rather densely woolly-
eiliate bracts but in most collections the bracts are onlv obscurely

1919] Hall: Chrysothamnus nmiseosus and Its Varieties 171

ciliate. Since also the direction assumed by the leaves is not constant,
this variety is exceedingly unstable and probably represents an
ecologic form.

10. Chrysothamnus nauseosus var. bernardinus var. nov.

Shrub leafy to the summit : annual twigs strict, about 3 dm. long,
coated with a smooth gray pannose tomentum, not striate : leaves
ascending, linear, 3 to 5 cm. long, 1 to 2 mm. wide, mostly condupli-
cate, 1-nerved, very acute, often recurved at tip, green and scabro-
puberulent : inflorescence a loose rounded cyme, 3 to 5 cm. across,
the lower branches 2 to 6 cm. long: involucre 5-flowered, cylindric-
turbinate, about 12 mm. high ; bracts in well-defined ranks, 4 in each
rank, keeled, thin, stramineous, lanceolate (the short outermost ones
ovate), sharply acuminate, glabrous or slightly erose-ciliate or the
outer ones obscurely puberulent : corolla 9 to 10 mm. long ; tube
sparsely crisp-puberulent ; lobes linear-lanceolate, acute, 1.8 to 2 mm.
long: anther-tips acute, about 0.5 mm. long: style-branches about
4 mm. long, stigmatic about one-half their length : akenes densely
sericeous-pubescent: pappus soft, about 8 mm. long, becoming pink.

Frutex undique foliosus: ramulis horotinis strictis non striatis,
ca. 3 dm. longis, tomento pannoso cinereo vestitis : foliis ascendentibus,
linearibus, 3-5 cm. longis, 1-2 mm. latis, plerumque complicatis, 1-
nerviis, acutissimis, apice saepe recurvatis, viridis, scabro-puberulis :
inflorescentia cymam rotundam laxifloram formante, 3-5 cm. latam,
ramulis inferioribus 2-6 cm. longis: involucris 5-floris, cylindrato-
turbinatis, ca. 12 mm. altis, squamis distincte seriatis in utraque serieri
verticali 4, carinatis, tenuibus, stramineis, lanceolatis (exterioribus
brevioribus ovatis), acuminatis, glabris, nunc parce eroso-ciliatis nunc
exterioribus obscure puberulis : corolla 9-10 mm. longa ; tubo parum
crispo-puberulo ; lobis lineari-lanceolatis, acutis, 1.8-2 mm. longis :
antheris apice acutis, ca. 0.5 mm. longis : styli ramis ca. 4 mm. longis,
parte stigmatice duplo longioribus : achaeniis dense sericeo-pubesceuti-
bus : pappi setis tenuibus, ca. 8 mm. longis, colore deinde roseis.

Dry, open hillside at 7400 feet altitude, Bluff Lake, San Bernar-
dino Mountains, California, September 2, 1905, Joseph Grinnell
(Herb. Univ. Calif, no. 149208), type. As far as known, this variety
is confined to the San Bernardino and San Antonio mountains of
southern California. The specimens cited below exhibit the follow-
ing variations : cyme up to 10 cm. in diameter and the lower branches
10 cm. long: involucre 10 to 13 mm. high, occasionally six-flowered.
Because of the frequent longitudinal folding of the leaves, these
structures often appear much narrower than they really are. This
applies to the type as well as to most of the other specimens cited.

In addition to the type, the following collections apparently belong
here : Little Bear Valley, San Bernardino Mountains, Mrs. C. M.
Wilder, no. 740; Little Green Valley, San Bernardino Mountains,

ITl! (' tiir( rsil 1/ of Cnlifoniia I'uhlicdtioiis in Iloinini | \'<)i.. 7

(Jt'o. H. Hall. iHi. l!4; Pine Mdiiiitaiii i\idfi:o, San Antonio MoiiiitMins,
I. M. .Joliiisloii. MO. K).")!*: rockv hillsides of Ixoinul \'allt'y. San
.lacinto Mountain al !»()()() lo D.IOO feet altitndc, II. M. Hall, no. ::;41.
In tliis last lui'nl ioMid collect ion tin' coni|)act cynus ai'c only .'^ to 4 cm.
aci-oss, the anther-tips are 1 nun. or iiioi'e lonji' and vei'y slendei", the
stylc-hi'anchcs are fully exserted and 4 to 6 nnn. lon<>-. A detailed
study of fresh material may dcMiionsti'ate that it is a variety di.stinet
from ht rnarcJiiius, bnt the s])eeimens at iiand are too nieaj^re to justify
this.

The variet\' b( riuirdinus is somewhat like (jccl<l( ntdlis hut with a
larger and loo.ser intloreseenee, lonjjer involucres and flowers, and
thinner tegules which are gradually acuminate instead of abruptly
jiointed. In most of its characters, as well as in the appearance of
herbarium specimens it almost du])licates th(^ Rocky ^Mountain piil-
rh< rriiHiis, here referred to var. sp(ci()siis, but the involucre is longer
and the moi-e sli'ongly keeled bracts are acinninate instead of merely
acute. Moreover the geographic isolation of l)( r)i(ir<ll)iiis argues for
its recognition as a variety distinct from pulchcrriiiius.

11. Chrysothamnus nauseosus var. Bigelovii (Gray) comb. nov.

Linosyris {Chriisathdninus} Bi(f<iovii Gray, Pacif. E. E. Eopt.. vol. 4, no. 4.
p. 98, 18.j7.

Shrub densely branched, 3 to 10 dm. liigh : twigs siiort, leafy,
yellowish green v.'ith a closely ])acked tomentiun : leaves linear-filiform,
1 to 3 cm. long, tomentulose when young: inflorescence a lax few-
headed cyme terminating each of the branches: involucre 10 to 12
mm. high ; bracts 4 or 5 in each row, tomentulose and sometimes
ciliate, attenuate, spi'cading in age: corolla 8 to 9 mm. long: lobes
ovate, erect, 0.5 to O.S mm. long: akenes glabrous.

The variety HujiJovii is a form from the vei-y dry plains and hills
of the southern Rocky ^lountain region and southward to Texas. It
has been classified near to C. piilch<lhis but the brittle striate twigs
and divaricate branching of that sjiecies indicate for it an entii'cly
different origin. Moreover, the loose invohiei-es of Biyelovil, with
bracts in oidy obscurely vertical ranks, are very unlike those of the
Pulchdli. It is plaiidy a variant of nauseosus, and its nearest affinity
will probably be found in vai'. Jeiosperiiixs, from which it differs
chiefly in the very pubescent and acute bracts. Specimens gathered
in southea-stern Utah by Rydberg and Garrett (nos. 94.']5, 9940) are
intermediate, having the essentially glabrous involucres of l('iosp( rnms
but with bracts whicli are almost as sharp as in Iii(/eh>i'ii.

1919J Hall: Chrysothanuius nmiscosus and Its Varieties 173

12. Chrysothamnus nauseosus var.glareosus (M.E.. Junes) comb. nov.

Bigelovia glarcosa M. E. Jouos, Zoe, vol. 2. p. 247, 1891.

Shrub many-stemmed, said to be only about 3 dm. high : leaves
broadly linear, slightly widened above, plane : inflorescence cymose :
bracts about 4 in each vertical row, somewhat keeled, obtuse, sparsely
erose-ciliate, scurfy-tomentulose : corolla about 12 mm. long ; lobes
linear-lanceolate : akenes glabrous.

This variety is apparently restricted to central and southern Utah,
where it grows on gravelly mesas. It was originally compared with
leiospermus, from which it differs in the tomentulose involucre and
other characters.

13. Chrysothamnus nauseosus var. leiospermus (Gray) comb. nov.

Bigelovia leiosperma Gray, Syn. Fl., vol. 1, part 2, p. 139, 1884.

Shrub low (3 to 12 dm. high), with numerous short erect twigs,
these either moderately leafy or almost naked : twigs yellowish green
with a verA' close tomentum : leaves filiform or nearly so, acute, mostly
0.5 to 2 or 3 cm. long, essentially glabrous : lieads in close terminal
cymes of 2 or 3 cm. diameter : involucre 6 to 8 mm. high : bracts
linear-oblong except the short ovate outer ones, obtuse, glabrous :
corolla 5 to 8 mm. long, the tube very obscurely pubescent (or
glabrous?) ; lobes ovate, erect, glabrous, 0.5 mm. or less long: akenes
completely glabrous in the typical form but often sparsely pubescent,
especially along the prominent nerves.

This form inhabits the most arid portions of the Great Basin,
growing mostly on very dry exposed hillsides or in dry rocky stream-
ways. The only collections seen by the writer are from Clear Creek
Caiion, Utah, Candelaria in western Nevada, and Caliente in eastern
Nevada. The Clear Creek specimens formed the basis of Bigelovia
leiosperma var. ahhreviata M. E. Jones, Proc. Calif. Acad. Sci., ser.
2, vol. 5, p. 693, 1895, a form with scant tomentum and leaves 1 em.
or less long. The Caliente specimens (H. M. H., nos. 10791, 10795)
represent an extremely xerophile type of very dry exposed hillsides
in the Lower Sonoran Zone. Their stems are green and rush-like,
and the leaves reduced to scales about 3 mm. long, or rarely developed
and then about 1 cm. long. In one of the Caliente specimens the
akenes are very sparsely pubescent on the edges ; in another from the
same station they are sparsely pubescent also on the intervals. This
indicates that the absence of pubescence cannot be satisfactorily used
as a specific character. The relation.ships of leiospermus are probably
with Bigelovii.

174 Univcrsitjj of California Publications in Botany [Vtn,. 7

14. Chrysothamnus nauseosus var. graveolens (Nutt.) Piper.
C'oiitr. i:. S. Nat. Ik-rb., vol. 11, p. f).-)!). 1906.

Chrysocoiiui (/raveoleiis Nutt., Gen., vol. 2, p. 1'Mi, 1818.

Bigelovia (/rarcoh-ns phthrata Gray. Proc. Am. Acad., vol. 8, p. CA'), 1873.

Shrub robust, leafy to summit : twigs yellowish green to nearly
white, more or less striate, the tomentum compact and smooth : leaves
broadly linear, 1 to 8 mm. wide, mostly 3- oi- 5-n('rv('d, impuiictate,
smooth and green but often slightly tomentvdose, especially beneath :
intiorescence a round or flat-topped cyme, fastigiate, the heads
crowded: involucre 6 to 9 mm. high, glabrous: corolla 6 to 9 (rarely
10) mm. long; lobes 0.5 to 1 or rarely 1.6 mm. long, erect.

There can be little doubt that the original of Nuttall's Chrysoconm
graveolens is the plant here described, for in the brief description are
mentioned the linear, three-nerved, smooth leaves ; the corymboselj'
fastigiate and crowded "flowers"; and the smooth, five-flowered,
angular "calix." This is the common form in the easterly part of
the range of the genus, especially in Wyoming, Colorado, and
northern New Mexico. C. virens Greene, Pitt., vol. 5, p. 61, 1902,
apparently is not to be retained even in the most subordinate rank.
From the description it seems to be very near to graveolens but it is
perhaps greener, the involucre only one-half as long as the flowers,
and the bracts somewhat triangular. These characters are all
extremely variable. Although the writer has searched for it at the
type locality (Canon City, Colorado) and in surrounding districts,
DO specimens were found that could be satisfactorily separated from
graveolens. C. laetevirens Greene, I.e., described from material
gathered at Grand Junction, Colorado, has been studied at its type
locality. Apparently there is no reason for considering it more than
a light-green form or state of graveolens.

Id. Chrysothamnus nauseosus var. calif ornicus (Greene) comb. nov.

Chrysothamnus calif ornicus Greene, Eryth., vol. 3, p. Ill, 1895, in part.

Shrub low and leafy-stemmed : twigs white with appressed tomen-
tum, leafy to the summit: leaves broad (2 to 5 mm.), narrowly
oblanceolate, 3-nerved, scarcely tomentulose but green and densely
glandular: inflorescence minutely glandular, consisting of short ter-
minal cymes: involucre 9 to 10 mm. high, 8 to 10-flowered ; bracts
thin, loose, only 2 or 3 in each vertical row, all lanceolate, attenuate,
not tomentose but rather densely glandular: corolla about 10 mm,
long; lobes 1.2 to 1.5 nnn. long; tube puberulent (not arachnoid).

1919J Hall: ChrysothcDinius ^aiisfosus (uid lis Vnrictirs 175

As here restricted, calif ornicus is a rare siib-shriib oi' the high
Sierra Nevada. Oiir description is drawn from Bolander's no. 6137
from Mono Pass at 9000 to 10,000 feet altitude. In the original
description Greene combined with this a taller form with tomentulose
leaves and inflorescence and 5-fiowered he.ads, which form belongs to
lower altitudes and is included in speciosus of the present synopsis.

16. Chrysothamnus nauseosus var. ceruminosus (l)ur. & Ililg.)

comb. nov.

Liiiosiiru'i (■( ) uiniiiosd J)iir. & Hilg., Pac. R. Rept., vol. ."), part iS. p. 9,
pi. 6, 1836.

Shrub 5 to 12 dm. high, fastigiately branched, the stems yellowish
green with a compact tomentum : leaves linear-filiform, 1 to 3 cm.
long, tomentulose : inflorescence compactl.y cymose, rounded, 2 to 3 cm.
across: involucre 7 to 8 mm. high; bracts thin, yellowish, abruptly
narrowed to a filiform recurved mucro about 1 mm. long, glabrous
and glutinous or apparently somewhat puberulent : corolla about 6.5
mm. long; lobes 1 to 1.7 mm. long.

Known from oul}' two collections, namely, the type collection by
Dr. Heermann somewhere near Tejon Pass, California, and one made
by Mrs. Spencer in October, 1917, at Hesperia, a station on the Mojave
Desert about ninety miles east of Tejon Pass.

17. Chrysothamnus nauseosus vai-. creophilus (A. Nels.) comb. nnv.

Chrysotliamiius orcophilus A. Ncls., Bot. Gaz., vol. 28, p. 'M'k 1899.

Shrub described as 2 to 4 dm. high with erect stems, very leafy
to the top : twigs .yellowish green, the tomentum thin and smooth :
leaves 3 to 7 cm. long, about 1 mm. wide, strictly erect, the upper
ones crowded and not reduced, all tomentulose at least when young-
but green : inflorescence composed of numerous small cymes in a more
or less elongated but round-topped thyrse : involucre 7 to 8 mm. high ;
bracts carinate, acute, sparingly tomentulose, ciliolate : corolla 7 to
10 mm. long; tube crisp-pubescent; lobes linear-lanceolate, 1.3 to
2 mm. long.

This is an apparently rare plant of saline soils in Wyoming
and Idaho. Its characters indicate that it may be a derivative of
graveolens differing chiefly in the narrower leaves and much longer
corolla-lobes. In the latter respect it resembles consi))uIis, into which
more westerly form it probably intergrades. The more numerous and
wider strictly erect upper leaves and the somewhat flat ciliate bracts
may, however, serve to distinguish it from co)islniiIis.

176 University of California Puhlicatimis in Botany [Vol. 7

18. Chrysothamnus nauseosus vniv pinifolius (Greene) comb. nov.

Clirijsolliamnus pinlfulius Grccue, I'itt., vol. 5, p. GO, 1902.

Slirub of medium size, with slender, very leafy stems: leaves linear-
filiform, 1 mm. or less wide, 1-nerved, green to grayish puberulent,
often densely toniciilulose beneath: infloreseenee a thyrsoid i)ani('le of
roundi'd or pyramidal outline, varying to sliort-oblong : involucre
about 7 mm. long, glabrous : corolla 6 to 9 mm. long ; lobes 0.5 to 1 mm.
long.

•o*

Most abundant in Colorado but extending into adjacent states ;
replaced further west by co^isimiUs. The characters separating this
variety from graveolens seem to be correlated with the more vigorous
growth of the latter and intermediate forms are common. C. patens
Rydb., Bull. Torrey Club, vol. 31, p. 652, 1904, is a form of pini-
folius with spreading and more or less falcate leaves, but this character
is widely variable on single plants and moreover it does not vary in
unison with otlier characters. The corolla-lobes in pinifolius are
described as about 0.5 mm. long; those of patens as about 1 mm. long.
However the lobes are not infrequently as much as 1 mm. long and
occasionally even 1.4 nnn. in plants otherwise typical of pinifolius.
C. con finis Greene, Pitt., vol. 5, p. 62, 1902, of New Mexico, is scarcely
separable from pinifolius. Its best characters lie in the rather obtuse
and short-ciliate bracts, but even in pinifolius at least a portion of
the bracts aire more or less ciliate. C. falcatvs Greene, I.e., based
upon specimens collected on the plains about Grand Junction, Colo-
rado, is pinifolius witli the lower face of the leaves white-tomentose,
as is commonly the case. Dr. Greene used also the curvature of the
leaves and the angle made by them with the stem as differentiating
criteria but field studies made by the author at Grand Junction
indicate that such characters are much too variable, even in individual
plants, to be of use. This is indicated by sheets nos. 203083 and
203177 at the Universitj- of California, under graveolens.

10. Chrysothamnus nauseosus \;\r. consimilis (Greene) comb. nov.

Chrysothamnus consimilis Greene, Pitt., vol. 5, p. fiO, 1902.

Shrub of medium or large size, the slender erect twigs very leafy:
leaves a.scending or spreading, nearly filiform, less than 1 mm. wide,
l-ncT'ved. somewhat resinous and eaneseently tomentulose to neai'ly
glabrous: inflorescence an elongated ]iyramidial or cylindrie thyrsus:
involucre 7 to mostly 8 or 9 mm. high; bracts not in distinct vertical
rows, glabrous: corolla 7.3 to 8.3 mm. long; lobes 1 to 2 nun. long,
spreading in age.

1919] Hall: Chrysothamnus natiseosus and Its Varieties 177

This is the common Chrysothamnus on alkaline flats of the Great
Basin, especiallj^ in northern Nevada, whence it extends to Arizona,
California, Oregon, and Idaho. In western Utah also it is the common
alkali-flat form but easterly it merges both in characters and distri-
bution into pinifolius. It may usually be distinguished from this
form by the longer, more spreading corolla-lobes, and sometimes by
the more elongated inflorescence. To the southwest of its range it
passes into viriduliis, as indicated under that variety. C. angnstus
Greene, Pitt., vol. 5, p. 64, 1902, described from specimens collected
in northeastern California, is exactly the same. It was said to be
distinguishable by its canescent "woolliness," but copious material
from both tj^pe localities indicates that this character can be correlated
neither with other characters nor with geographic distribution, and
the amount of pubescence is of course variable. These remarks apply
equally well to leaf-length, and no other differentiating characters
appear in the descriptions.

20. Chrysothamnus nauseosus var. viridulus var. nov.

Shrub robust, green, about 1.5 m. high, leafy to the summit :
twigs densely covered with pannose yellowish green tomentum, rather
stout, striate : herbage malodorous : leaves at first erect or ascending,
later inclined to droop, narrowly linear, 3 to 5 cm. long, about 1 mm.
wide, acute, 1-nerved, channeled above, green but tomentulose on
both sides : inflorescence a pyramidal to nearly globose, thyrse : in-
volucre 6 to 7 mm. high, glabrous but viscid with a resinous exudation ;
bracts 3 or 4 in each of the 5 vertical ranks, carinate, the outer ones
acute, the innermost ones obtuse : flowers, 5 : corolla 8.3 to 10 mm.
long; tube glabrous or sparsely puberulent, passing gradually into
the throat; lobes linear, acute, 2 to 2.5 mm. long, 0.4 mm. wide,
recurving in age : anthers 3 to 3.3 mm. long, the tip 0.42 to 0.68 mm.
long : akenes densely sericeous : pappus about 7.5 mm. long.

Frutex validus, viridis, ca. 15 dm. altis, undique foliosus : ramulis
tomento luteolo-viridi dense vestitis, paullo robustis striatis, odore
injucundis : foliis primum erectis vel ascendentibus, delude flaccescere
inclinantibus, anguste linearibus, 3-5 cm. longis, ca. 1 mm. latis,
aeutis, 1-nerviis, supra impressis, viridulis, utrinque tomentulosis :
inflorescentia thyrsum globoso-pyramidatum formante : involucro 6-7
mm. alto, glabro, resinoso-viscido : squamarum seriebus verticalibus 5,
in utraque serieri 3^, carinatis, exterioribus aeutis, interioribus
obtusis : floribus 5 : corolla 8.3-10 mm. longa, tubo glabro parce
puberulo, infundibuliformi, lobis linearibus, aeutis, 2-2.5 mm. longis,
0.4 mm. latis, deinde recurvatis : antheris 3-3.3 mm. longis muorone
0.42-0.68 mm. longo : achaeniis dense sericeis : pappo ca. 7.5 mm.
longo.

■&^

Benton, Mono County, California, on sandy alkaline flats with
Distichlis, in the Upper Sonoran Zone, at 5640 feet altitude, Novem-

178 Univcrsllii of Californ'in I'liblicdtloiis i)i Hotaiiij | V'<>i.. 7

b.T :]. 1917, 11. :\I. 11.. n... 10642 (ITcrl). Tiiiv. Calif, no. 203068), type.
This is the coiiiiiioii fonn on alkaline fiats in soutiici'n Mono County
and in Inyo County. California, and in wcslci'ii Esmeralda County,
Nevada. al1liou;:li there ai'e many slii^lit variations from tin- type as
dcscrilnd above. Almost evciy valley exhibits forms not exactly like
those in any other. The variations are chiefly in liabit, ])ubescence.
leafiness, size and shaj)e of iiiHorescenee, shape of biaets, and Icnjrth
of corolla-lobes. The plants range in height from a few dm. to nearly
3 m., but are always taller than broad unless abnormal ; the corollas
vary from 8 to 10 n]]n. in total length ; the corolla-lobes are seldom
shorter than 2 mm., yet in two eolh ctions there are some flowers with
lobes oidy 1.7 mm. long; although the iiivolucres of the ty])e are only
6 to 7 mm. long, they vary in other specimens to 8 or even 9 mm. ;
the mature papus is 7 to 9 mm. long. A dwarf form of the alkali
flats of Antelope A'alle\- is referred here provisionally. It has flexuous
stems, short rounded inflorescences and exceptionally small flowers
(eommoul}' 6 or 7 in a head), but the flowers, although reduced in size,
have the narrow elongated and spreading lobes of viriduluft. More
nearly typical specimens have been gathered in the San Antonio
Mountains at an altitude of 8000 feet, I. ^I. Johnston, no. 1706.

As to relationships, viridulus is probably a southwestern derivative
of cojisimilis (or vice versa), from which it ditfers in the larger
corollas with longer lobes, the thicker, more robust and rigid twigs
and leaves, the heavier and more rounded inflorescence, and the
stronger odor of the herbage. These characters are far from constant
at all stations. The length of corolla-lobe is the most satisfactory.
Of twenty-seven collections taken throughout the established range
of the variety, ojdy five have corolla-lobes 2 mm. or less long ; of
twenty-two collections from the range of coif^iiiiiJifi none exhibit
corolla-lobes of over 2 mm. in length : where the ranges meet as
ai'ound ]\Iono Lake and at Sodaville, Nevada, intermediate sizes are
frecpient and. here the other differentiating characters also intergrade.
The very long corolla-lobes serve as a certain means of distinguishing
viridulus from all other varieties except consiniilis, occid( ntalis,
htriKirdiuK.'i, and an occasional mohavensis.

1919] Hall: Chrysothamnus )wuseosus and Its Varieties 179

21. Chrysothamnus nauseosus var. mohavensis (Greene) comb. iiov.

Bigeloviu mvliavensis Greene, in Gray, Syn. Fl., vol. 1, part 2, p. 138, 1884.
Clniisnthavinns mohavensis Greene, Eryth., vol. 3, p. 113, 1895.

Shrub of medium or large size, with many erect or ascending
branches which are often nearly leafless and rush-like: leaves filiform,
very acute, 1-nerved, nearly glabrous : inflorescence a rounded or some-
what elongated thyrse : involucre narrow, 9 to 10 mm. long, sharply
5-angled; bracts obtuse to acute, in very distinct vertical ranks,
glabrous: corolla 8 to 10 mm. long; lobes 1.5 to 1.8 mm. long, spread
ing.

In this variety we have apparently a southern derivative of viridu-
lus. It belongs to higher ground where the drainage is better and the
soil not obviously alkaline. It skirts the westerly side of Owens Valley,
California, extending southward to the slopes west of Antelope Valley
and swings around the latter as far as the desert slopes of the San
Bernardino Mountains. It occurs also at an isolated station on
Mt. Hamilton, far out of its general known range, but the specimens
at that station seem entirely typical and Dr. Greene, the first to detect
the form, himself admitted them as mohavensis. The species will
probably be found elsewhere along the hot inner South Coast ranges,
a little-explored district where many species of the soutlicrii deserts
extend their ranges northward.

Although first described as sparsely leafy or leafless this character
cannot be relied upon since the tendency toward an early dropping
of the leaves is common to the whole group. It is true, liowever, that
mohavensis is more commonly leafless or nearly so. Tlie naked wand-
like branches are sometimes much elongated, in one case measuring
7.5 dm. without leaf or branch (11. M. H., no. 10570). The best
characters lie in the involucre, which is mostly longer than in the
other forms. Although the longest involucres of virididus slightly
exceed the shortest ones of mohavensis, yet tlie average of the former
is about 7 mm., of the latter about 9 mm. The l)racts are distinctly
five-ranked and carinate, so that the involucre is sharply five-angled.
They are somewhat obtuse but in some specimens which are referable
here by all other characters the bracts are decidedly acute (II. M. H.,
nos. 9894, 10570, 10611, and Hall and Babcock, no. 5090).

As indicative of the gradation between mohavensis and viridulus
may be cited a specimen from Oak Creek, along the west side of
Owens Valley (H. M. H., no. 10611). The habit is that of mohavensis
althouuh some of the twigs were leafy when gathered on October 30.

180 Universifij of Califoriiiu I'lihlications in Butany [Vol.7

The togiiles are in very sharply defined vertical ranks, as in tliat
variety, but tliey are even more acute than in viridulus; and the
corn]la-h)l)(s arc exactly intermediate in len<ilh, measuring: 2.1 mm.

22. Chrysothamnus nauseosus var. junceus (Greene) condj. no v.

Bigelovia Juiuua (ireciic, Bot. Gaz., vol. (J, p. 18-4, 1881.
ClniisoUuimiius junccus Greene, Erytli., vol. 3, p. 113, 1895.

Shrub strict, fastif»'iately nnich branched, with slender rush-like
mostl}' leafless yellowish green branches : leaves linear-flliform : inflor-
escence fastigiate-C3^mose : involucre about 10 mm. high, glabrous;
bracts acute, 5 in each of the distinct vertical rows : corolla about
9 mm. long; tube pubescent but not arachnoid; lobes about 1.5 mm.
long, externally beset with long delicate hairs.

This is a little-known shrub of eastern Arizona. It is described
by Greene as "cinereous"; by Gray as "minutely canescent", but
the twigs ill the t^'pe have the usual pannose tomentum and all
other characters indicate that it belongs to the present group. The
pubescence of the corolla-lobes should be distinctive if constant.

FORMS NOT DEFINITELY PLACED

The following named species are all very close to Chrysothanimis
nauseosus and at least some of them should doubtless be included in
that species when taken in its broadest sense. However, they are
not sufficiently well known to the writer to justify him in passing
upon them at present.

Chrysothamnus formosus Greene, Pitt., vol. 4, p. 41, 1899. A low
white shrub with narrow spreading foliage and wholly glabrous
involucres with 6 or 7 bracts in each vertical row. Described from
immature plants gathered "in the neighborhood of a mineral spring
among the hills a few miles southwest from Grand Junction, Colo-
rado."

Chrysothammis Macounii Greene, Pitt., vol. 5, p. 63, 1902. Said
by Dr. Greene to resemble plattensis and like it a low plant with
white-t(mientose twigs and very narrow spreading leaves. The type
was from near Lytton, British Columbia.

Chrysothamnus tnoqumnus Greene, I.e., p. 60. Described from
imperfect specimens but seemingly a variety of nauseosus, perhaps
one of those with glabrous akenes.

Chrysothamnus turhinatus Rydb., Fl. Rocky Mts., p. 859, 1917
(Bif/rlovia turhinata M. E. Jones, Proc. Calif. Acad. Sci., ser. 2, vol. 5,

1919] Hall: Chrysothamnus nauseosus and Its Varieties

181

p. 691, 1895). Said to be next to junceus and with the uppermost
leaves reduced as in that form but with glabrous corollas. The bracts
are five or six in each row and all obtuse or only apiculate. The type
locality is Canaan Ranch, Utah.

IV. INDEX OF SPECIFIC AND VARIETAL NAMES

The numbers given in this index refer to the accepted variety in
the foregoing synopsis, under which the name will be found.

abbreviatus, 13
albicaulis, 1
angustus, 19
appendiculatus, 3
arizouicus, 3
beniardinus, 10
Bigelovii, 11
californicus, 15
ceruminosus, 16
concolor, 8
confiuis, 18
consimilis, 19
falcatus, 18
fasciculatus, 7
formosus, see p. 180
frigidus, 8
glabrata, 1-4
glareosus, 12
gnaphalodes, 5
graveolens, 14
hololeucus, 2
junceus, 22

laetevirens, 14
latisquameus, 3
leiospermus, 13.
Macounii, see p. 180
moliavensis, 21
moquianus, see p. 180
nauseosus (s. str.), 1
occidentalis, 6
oreophilus, 17
orthophyllus, 1
pallidus, 8
patens, 18
pulcherrimus, 7
pinifolius, 18
plattensis, 9
salieifolius, 4
speciosus, 7
tortuosus, 6
turbiuatus, see p. 180
virens, 14
viridulus, 20

UNIVERSITY OF CALIFORNIA PUBLICATIONS

IN

BOTANY

Vol. 7, No. 7, pp. 183-264, plates 18-20, 6 figures in text November 7, 1919

IT. CHKYSIL, A NEW liUBBER FROM
CHRYSOTHAMXrS NArSEOSUS

BY
HARVEY MONROE HALL axd THOMAS HARPER GOODSPEED

CONTENTS

PAGE

I. Object and scope of the investigation 184

II. Acknowledgments 186

III. Nature and prop':^rties of Chr_ysil 188

IV. Botanical classification; the names of the plants 189

V. Chrysothamnns nau.'ieasu!^: habit, flowering, rate of growth, etc 191

VI. Distribution and habitats of the various forms 194

VII. Estimates by districts of the amount of Chrvsil available in western

North America 107

a. District 1. East central California and adjacent Nevada 198

b. District 2. Mojave Desert, California 200

c. District 3. Northeastern California and adjacent Nevada and

Oregon 201

d. District 4. West central Nevada 202

e. District .5. Northern and central Nevada 204

/. District 6. Utah 207

g. District 7. Colorado 208

h. Estimates not included in the above districts 209

i. Conclusions as to the quantity of rubber obtainable 209

VIII. Methods of detecting the presence of rubber and determining its nmnunt . 210

a. Microscopical methods 210

b. Chemical analysis 216

IX. Results of the chemical analyses and microscoi)ical examinations; tal)ula-

tion of percentage of rubber in each variety 226

X. Distribution of rubber in the plant 234

a. Regional distribution in the plant 234

b. Specific distribution in mature tissues 239

XI. Factors influencing rubber content 243

a. Variation with the botanical variety 243

b. Variation due to environment 245

c. Seasonal variation 246

XII. Methods of harvesting: season, age, depth of cutting, etc 248

XIII. Possibilities of Chrijsnthamnus as a cultivated plant 251

XIV. Cultural requirements 2.55

XV. Summary 257

184 University of California Publications in Botany [Vol. 7

T. OBJECT AND SCOPE OP^ THE INVESTIGATION^

The investigation here reported upon was instituted for the pur-
pose of locating: a supply of rubber which it was thought might exist
in certain native West American shrubs. Of all the various species
originally considered, those belonging to the genus Chrysothamniis,
commonly known as Rabbit-brush, seemed to be the most promising,
and consequently they have received the most attention.

This work was undertaken immediately after the entrj^ of the
United States into the recent war and had for its incentive a considera-
tion of the following facts : that rubber is absolutely essential to mod-
ern warfare ; that it is the only war essential not now produced in
this country ; and that, therefore, we as a warring nation would be
seriously handicapped in case the enemy should be able to carry out
his threats and put a stop to our oversea commerce. This danger of
interference with our importation .of rubber has now been happily
averted but since no one is able to state with certainty that it may not
recur, the location of a native supply is still a matter of national con-
cern. Our studies have therefore been continued somewhat beyond
the duration of the war and have resulted, as indicated in the follow-
ing pages, in the finding of rubber of good quality in some of our
western shrubs. If it were all assembled the total amount (estimated
to be over 300,000,000 pounds) would be considerable, but the per-
centage content of the plants is too small to warrant harvesting except
under stress of national emergency. The very fact that the shrub is
not rich in rubber may have its advantages when the matter is viewed
from the standpoint of the nation's needs. It means that the rubber
will not be subjected to commercial exploitation and will thus be pre-
served as an emergency supply to be drawn upon only in case we reach
such straits that its utilization becomes necessary notwithstanding the
high cost of harvesting.

The very best protection against the possibility of enemy interfer-
ence with our supply would be the development of a permanent rubber-
producing industry in this country. Aside from the manufacture of
synthetic rubber on a commercially profitable scale — a desideratum
which does not seem to give promise of early realization — our only

1 This investigation was carried out t-hiefly by nicniberJi of the Department of
Botany of the University of California and was one of tlie jnojects of the Sub-
committee on Botany of tlie Committee on Scientific Researcli of tlie State Council
of Defense of California.

1919] Hall-Goodspecd: Chrysil 185

hope for this lies iu the discovery and iraprovement of plants which
can be grown in this country and which will be sufficiently productive
to warrant their cultivation in competition with imported rubbers.
One such species has already been introduced — Mexican Guayule
(Parthenium argentatum), now being grown to a limited extent in
southern Arizona. There is no assurance, however, th^t this plant
can be profitably grown over a sufficiently large area to supply the
country's needs in time of ^var, nor are we certain that it is the
best one to be used. It is desirable that all promising sorts should
be carefully investigated and their possibilities determined. These
reasons have led to a broadening of the scope of our inquiry to include
a preliminary study of the cultural possibilities of the plants under
consideration.

This extension of the scope of the investigation has emphasized our
deficiencies in time and facilities for the satisfactory prosecution of the
work. Since it has been carried on thus far as a purely war emergency
matter, the aim has been to assemble our information as rapidly as
possible, and we have not permitted ourselves to become unduly en-
grossed in any phase of the work that seemed to require a long period
of study or experimentation. The authors regret this keenly, since it
means that the report here presented must be very incomplete ; that
many fundamental questions regarding both the scientific and practi-
cal aspects of the subject must go unanswered. It seems our duty,
however, to place on record such data as we have, including particu-
larly such information as will assist those who may interest them-
selves in a further search for rubber-bearing plants. Methods of car-
rying out the histological examinations and chemical analyses are for
the same reason given in detail though they may prove neither novel
nor instructive to the trained botanist and chemist. It is hoped that
this report will serve as a basis for more intensive researches, either
by ourselves or by others, when conditions are more favorable and that
it will point the way to special investigations that have to do with the
formation and occurrence of rubber in these West American shrubs
and its possible utilization by man.

The preliminary studies here described have been in progress for
about two years but the work was carried on only during vacation
periods and in such additional time as could be spared from regular
university duties. In addition we have profited by the generous assist-
ance of numerous co-workers to whom acknowledgment is made below.

186 Uuivf rsil !/ of Cal ijOniid V uhiic(iti(ni<; In l>i)tinni |\(>i,. 7

Till' (liscovci'v of rubber in Chrt/sotlianDius, wliicli wms made about

fifteen years ago, has been described by us in a icecnt [)ai)er-' as

follows :

Tlic I'lioii'c (if (']ir!i.s<>th(int)nis ;\Ui\ iclatcd "rcnora as tin- |il:ints fii-st to bf
iiivestiyati'il was tlio result of ;i proliniiiiary cxaniiiiatioii iii.nli' in IDOI. In
8i'i)t('iiib('r of tliat year tlie late Jmlgp A. V. Davidson, of I nilriMinliMicc, Inyo
County, California, sent some twijrs to tlic i)c|iaitincnt of Botany for idciitidcation.
with tlu' information that tho Indians jircpiircd from tlie ])lant a sort of ''t^um''
Avhitdi they chewed. The plant was a sj)eeies of Clirysotlianiiius of the niavcolens
grouj). Further samples were submitted at our recpiest, and in Oetolx'r, 190-^, a
preliminary idiemi<-al examination of them was made by I'rofessor G. H. Colby,
of the California Kxjieriment (Station. This examination indicated the presence
of rubber, but not in sufficient amount to warrant further investijration. A report
to this effect was made public in the press and as a result some further examin-
ations were niaiie by at least one commercial rubber conii)any. The matter was
soon drojiped. however. It is probable that the plants used in tliis commercial
examination were of an entirely different s})ecies from those now bein<r examined.

The examination of 1!)04 was followed with field work b,\- the senior
author in Inyo County in 1!)06, but no further ehenueal examinations
were made. Professor Marcus E. Jones informs us that steps were
taken about 1908 to erect a rubber-extraction plant at Salida, Colo-
rado, the intention being to use Rabbit-brush but the plan fell through.
He also tells us that as early as 1878 the Indians near St. George,
I^tah, taught some Mormon boys how to [)repare nd)ber by mastica-
tion of the inner bark of these plants. It is evident that the Indians
have long made use of the rubber as a chewing gum, but we have
been unable to learn of any scientific study of the plants as rubber
pi-odue( rs up to the time of our preliminary work in 1904. '■

II. ACKNOWLEDGMENTS

Throughout the course of the investigation the autliois have had
the valual)le co-operation of a considerable nundx'i' of workers, some
of whom have given largely of theii- tinu' without reiiuinei-ation. In
the eai'ly stages of the work the chemical analyses were made by Pi"o-
fessor Paul L. Ilibbard. in the laboratory of the Division of Agricul-
tural ('JKiiiistiy of the California Exeriment Station and the staff of
that laboratoi'v, under the direction of Pi'ofessor .1. S. I'>urd, has given

^Science, n.s., vol. 47, ji. 4.12 (May 10, 1918).

^5 Since the above was written word has come to us that a factory w^as in ojut-
ation at Jluraiifjo, Colorado, as early as 190.') and that the company in charge of
it actually ]daced ujion the market i-ubber made from ' • h'abldt weed. ' ' How-
ever, the results of fiuther in<|uiiy indicat*' almost with ceitainty that the plant
used was not Chriisothanuiax, but the "Colorado Rubber IMant'' {II i/inrnojijs
floribiDula ^itilis).

1919] Hallr-Goodspcrd: ('hrijsil 187

invaluable aid in the establishment and operation of our own labora-
tory', where the later analyses have been made by the junior author
and Miss Mildred Crane. Dr. David Spence, Chairman of the
Subcommittee on Rubber and Allied Substances of the National Re-
search Council, has superintended the extraction and vulcanization of
rubber from sample shrubs sent him and has given opinions from time
to time regarding our metliods and the quality of the product. Dr.
W. B. McCallum, and Professor Francis E. Lloyd, both recognized ex-
perts on Guayule production, have made many valuable suggestions.
Mr. E. C. McCarty of the University of California made a field trip of
two weeks' duration into central Nevada for the purpose of collecting
samples and making estimates as to the occurrence and distribution of
certain species and Mr. J. R. Bruff, of the same institution, spent two
weeks on such work in northern California and Nevada and southeast-
ern Oregon. Professor Marcus E. Jones, of Salt Lake City, has per-
formed a similar service in Utah. He spent a total of twenty days in
the field and in addition has contributed freely from his very de-
tailed knowledge of the botany and distribution of the shrubs through-
out the Great Basin Area. Through the courtesy of Dr. Frederic E.
Clements, of the Carnegie Institution of AVashington, the senior au-
thor, was enabled to spend a month in field studies extending from
Oregon to New Mexico. It is a pleasure to acknowledge also the as-
sistance of Professor W. A. Setchell, head of the Department of Bot-
any of the University of California, who has supported the work
throughout both by his sympathy and advice and by making it pos-
sible for the authors to give to it a larger portion of their time than
is ordinarily available for investigational work. The laboratories and
herbarium of the Department of Botany have been freely used and
much of the expense has been borne by the University. Grants have
also been made, more especially for field expenses, by the Committee
on Scientific Research of the State Council of Defense of California.
The list of friends who have sent information and samples, often
at much trouble, includes the following: Mrs. Sidney Armer, Berke-
ley; Mr. W. W. Blakeslee, U. S. Forest Service; Mr. Fred E. Burlew,
Los Angeles; Miss Ethel J. Case, Spokane; Mr. John Dondero, Mono
Lake; Mrs. Roxana S. Ferris, Stanford University; Mr. M. French
Oilman, Banning; Mr. Benj. J. Holfner, U. S. Forest Service; Mr.
Edmund C. Jaeger, Palm Springs; Mr. Geo. C. Larsen, U. S. Purest
Service; Mr. AV. M. Maule, U. S. Forest Service; Mr. James W. Mc-
Gow^an, U. S. Forest Service; Mr. Alexander McQueen, IT. S. Forest

188 TJniversiiy of California Puhlications in Botani/ [Vol. 7

Service; Mr. Geo. E. Moore, U. S. Forest Service; Mr. R. C. North,
IMcGill, Nevada ; Mr. S. B. Parish, San Bernardino ; Mr. George Park,
U. S. Forest Service ; ]\Ir. W. O. Sander, Center, Colorado ; Mr. Ta S.
Smith, U. S. Forest Service; Mr. O. P. Stilwell, Ogden, Utah; .Mr.
J. Sutcliff, Sntcliff, Nevada ; Professor J. J. Thornber, University of
Arizona; Mr. H. B. Way, Montello, Nevada; Professor A. 0. Weese,
Universitj^ of New Mexico; ]\Ir. Kirk Whited, Redmond, Oregon.

For courtesies extended in connection with the field experiments,
chiefly through the granting of the use of land and assistance in fenc-
ing, we are indebted to Messrs. George R. Shuey, Independence, Cali-
fornia; W. H. Davis, Benton, California; and Eugene L. Dutertre,
Golconda, Nevada.

To all of those who have assisted in the work, the authors tender
their sincere thanks.

III. NATURE AND PROPERTIES OF CHRYSIL

Chrysil is the name here proposed for the rubber prepared from
any form of Chrysothamnus nauseosus. It was suggested by Dr.
Frederic E. Clements and is selected because of its euphony and
brevity as well as for its suggestion of the botanical name of the plants
from which the rubber is obtained. "Chrysothamnus'' in turn, is
derived from two Greek roots signifying "golden" and "wood."

The first samples of Chrysil were prepared for us by mastication
from shrub of the viridulus form by Paiute Indians at Benton, Cali-
fornia. These pieces were in the form of cylinders three-fourths to
one inch long by about three-eighths of an inch in diameter. They
included, in addition to the rubber, small amounts of resin, fiber, and
other impurities. They were brown in color, firm, scarcely if at all
tacky, and seemed to possess considerable strength and elasticity. Two
experts accustomed to the handling of crude rubbers declared them to
be "of good quality and considerably better than rubber prepared
from Guayule." These samples were exposed to air and liglit and
underwent a gradual change, so that at tlie end of a year they were
black in color and quite tacky on the surface, these changes doubtless
being due to impurities.

Twenty-five pounds of viridulus shrub were gathered from the flats
at Benton and s]iii)ped to Dr. David Spence, Chairman of tlie
Subcommittee on Rubber and Allied Substances of the National Re-
search Council, with the request that he prepare rubber from it and

1913] Hall-Goodspced: Chrysil 189

report on the yield and the quality of the product. The samples of
shrub consisted of plants cut off a short distance below the surface of
the soil and with all growth of less than three j-ears removed ; in other
words, they consisted of those portions which would presumably be
utilized in commercial operations. Dr. Spence very kindly undertook
to make the examinations, and reports as follows regarding the prep-
aration of the rubber and its properties :

In preparing the samples for analysis the woody ones were ground up on a
pair of corrugated rolls, or a " cracker, ' ' and the bark samples were ground up
and sheeted out on a smooth roll. It is interesting to note that the bark forms a
smooth sheet on the mill.

.Each of the four samples was placed in a bottle, covered with 100% benzol and
shaken one hour. It was then allowed to stand twelve hours and the liquor drained
off. This washing was repeated twice. After allowing the small dirt to settle
out, the three combined washes were distilled to a small bulk, 400 to 500 cc. The
ruljber was coagulated from this concentrated solution by the addition of pure
methyl alcohol. The coagulated mass of rubber was kneaded in fresh methyl
alcohol and vacuum dried at 95 °C. The dried rubber is dark brown in color, has
very little tackiness and considerable elasticity. When the solvents are removed
by boiling in water, a very tacky rubber results.

A larger sample of bark from both stems and branches was extracted and the
rubber obtained was compounded as follows : rubber 100 g., zinc oxide 100 g., ac-
celerator 3 g., sulfur 6 g. This compound was then divided into five portions and
these W'Cre vulcanized for thirty minutes, sixty minutes, ninety minutes, two hours,
and three hours, respectively. Forty pounds of pressure was applied in each case.
The sixty minute treatment under forty pounds was full cure.

These samples* indicate that the rubber vulcanizes readily and gives a product
of a very fair quality. I have no hesitation in committing myself as to its value
as far as it is possible to estimate this from such small samples as I was able to ob-
tain. In order to report on the exact commercial value of this rubber it would be
necessary to make many more tests and much more exhaustive ones, but from the
small samples I sent you I can already safely say that the rubber is of high
grade and average quality. It is not as good as the best fine Para, but it is a
great deal better than most Africans or low grade rubbers. The results would
indicate that the bark of shrub No. 64 (that is viridiilus) might well be investi-
gated more extensively as a possible source of crude rubber existent in this country.

IV. BOTANICAL CLASSIFICATION; THE NAMES OF THE

PLANTS

The plants in which rubber has been found during the course of
this investigation all belong to two closely related genera of the Com-
positae, namely Chrysothamnus and Haplopappus. The latter is
apparently of little importance, for although rubber has now been

i The five samples referred to by Dr. Spence are in the form of circular discs
one-fourth inch thick by one and one-fourth inches in diameter. They are pre-
served in the Botanical Museum of the University of California.

190 Viiiiu rsilji of Califoriiid /'t(h1icatio)is in lU/hnnj | \'<m.. 7

found ill tell of its species and in ralluT ]i\rgo amounts in two of these,
vol the iiijiii-])ereenta<»'e species are small ]>Iants. This <i-enns will he
dealt with elsewiiere.''

The otln'i', tliat is, Cliri/solliarnints, is a West American genus of
about sixteen iiiajor species, but these ineluch' so many variations that
sonic botanists would recognize nioic than one hundred specific segre-
gates. Eleven of the major si)ecies of Clirysolhainiius have now been
examined and rubber found in tive of them. One of these, C. nausrosus,
is of outstanding promise; the othei- four are discussed further on.''

Botanists are far from agreement as to the limitation of species in
the genus, but for pi-actical purposes it seems best to adopt a broad
species concejit and to recognize a considerabh^ number of varieties
under each of the sj)eeies. According to this ai-rangement Chrijso-
tlidiiuuis )i(iHseosus constitutes a nuijor species with twenty-two
varieties. Twelve of these have been examined for rubber and it was
found to be present in all of them. The complete botanical designa-
tion of each variety may be expressed by a trinomial, for example,
Chrysothammis nanscdsns var. gravcolciis; ('ItrysotlianiiiKs nauseosns
var. viriduhis, etc., but for simplicity we may be permitted, in a special
paper such as the present one, to omit the generic and specific names
and refer to the various forms only by their varietal names. We shall,
therefore, use such detached terms as graveolms and viridulus to ex-
press our concept of the respective varieties belonging to the very com-
plex species known as ('hr^isofhoiinnis nanseosus. It is ho])ed that this
simplification will appeal to the non-botanical reader, who often has
little time for and less interest in an involved taxonomic terminology
and that it will, at the same time, meet the needs of the professional
botanist. In some cases the name of a species itself will be used in-
dependently of its generic name. The exact taxonomic position of any
variety of (y'krysotlium)nis nauseosits may be determined by reference
to pages 15<) to 181.

The correct common name of ('IirysotJianDius, and more particu-
larly of C. nauseosns, and its varieties is Rabbit-brush, a name which
has been in use both among non-botanists and in botanical litiM-ature
for at least twenty-five years. Through erroneous identification it has
been frequently ap])lied to other plants. In two ])uhlieations it has
been used indiscriminately" for species of Chrijsotli'iiiuitis, T( I nid i/miii.

•■•' See pp. 268-274,
« See pp. 265-268.
7 X. Am. Fauna, no. 3.") (1913), pp. 28, 31. 37; no. 42 (1917), pp. 79. 81.

1 ^»1 9] HaU-Goodspecd : ( 'h rijsU ^ 91

and Gutirrrrzia and in a recent Experiment Station bulletin it has
been adopted for 7'etradijinia glahrata.^ But on grounds of usage as
well as priority the term " Rabbit -brush ' " should be used only for
species of (linjsotltamnHH. It is defined in this sense in three of the
leading dictionaries used in America and also in some designed more
especially for use in Europe. "Rabbit-brush" is given as the com-
mon name for (Jlirysothamnus in at least fifteen floras and technical
reports and, so far as we can discover, it has never been defined either
in dictionaries or in floras as belonging to any otlier plant. While
there is considerable misapplication of the term by stockmen and other
residents of the AVest, yet it is much more frequently applied to these
plants than to any others. The name "Golden Bush" is used some-
what in the Rocky Mountain states. The undiscriminating sometimes
fail to distinguish between these plants and Sage-brush, but the true
Sage-brush {Artemisia tridentata) is a very ditt'erent plant and maybe
recognized by the leaves, which are, for the most part, three-toothed at
the summit, whereas in the Rabbit-brush they are always entire.

The Paiute name for any sort of Chrysothamnus from which rubber
is prepared is tsigupi (pronounced tse-gdo-pee) ; or if the plant grows
in sandy soil it is called trha-tsigupi. It is possible that the older In-
dians restricted the use of tcha-fsigupi to some particular variety, but
the younger generation is not so discriminating. Dr. A. L. Kroeber,
who has indicated for us the proper spelling of these words, says that
teha means "sun." According to Dr. David P. Barrows, the Coahuilla
Indians of southern California call the graveolens form tes-i-nit and
prepare from its twigs a tea taken for coughs and pains in the chest.

V. CHRYSOTHAMNUS NAUSEOSUS: HABIT, FLOWERING,

RATE OB^ GROWTH, ETC.

All of the species of (Jhi-ysotliamnus are shrubs. Some are mere
dwarfs but those which interest us as possible rubber ])rodueers are
usually of good size, measuring three to eight feet high, and about as
broad. The rubber is present for the most part in those stem parts
which are three years old or more, and these portions in average ma-
ture plants of the more important varieties {viriduhis, hololeiicus, etc.)
will weigh fi-om five to fifteen pounds (2.3 to 6.8 kg.). An exceptionally
large plant found near Lone Pine, California, weighed sixty pounds ex-

« Nevada Exporinieiit Statiou, Bull. 95 (July, 1918j, pp. 2, 7.

192 University of Calif ornia Puhlications in Botany [Vol. 7

elusive of the twigs, and shrubs weighing twenty to forty pounds are
not rare. For purposes of estimate, however, much smaller figures
must be used. This is partly because the plants reach the maximum
size only under favorable conditions and partly because they are fre-
quently burned or cut off near the base, after which new stems shoot
up only to be again destroyed before reaching maturity. After weigh-
ing numerous samples and after making thousands of estimates in the
field tlie authors believe six pounds to be a fair average weight of the
woody portions of the vindulus variety as it grows in eastern Califor-
nia and western Nevada. This includes the root to a depth of only
four inches (10 cm.). This estimate does not include young seedlings
nor young second growth.

On a similar basis it is estimated that individuals of consimilis, the
common variety in northeastern California, Nevada, and western
Utali, will average five pounds, while variety pinifolius of Utah,
southern Colorado, etc., will average four or five pounds.

The rubber producing kinds of Rabbit-brush all grow from deep
taproots which have normally but few main laterals. There are usually
several trunks from a single base and these are clothed in age wuth
loose, fibrous brown bark which peels off in strips. Straight clear
stems are the exception. The branches are usually numerous; they
often exhibit many distortions and irregularities and are of unequal
size. This has a bearing upon the possibility of decortication by ma-
chinery, a point of considerable practical importance. The rubber is
carried chiefly in the inner bark and in the very outermost portion of
the wood, so that decortication would greatly reduce the bulk and
weight of the material to be extracted. The irregularities just men-
tioned would presumably render decortication by machinery imprac-
ticable as far as wild shrub is concerned. If the plants are brought
under cultivation they would be more regular and uniform in their
growth and the separation of the outer layers might then become
feasible. According to Dr. D. Spence the bark could be very easily
separated from the wood on a large scale even in irregularly shaped
plants by soaking in hot, dilute, caustic soda solution. The young
shoots are long, straight, and erect in viridulus; similar l)ut shorter
in graveolens and consiniilis; yery much branched and twiggy in
(jnaphalodes and hololeticns. In all of these forms they are covered
with a closely packed gray, green, or white felt-like tomentuin which
is deciduous only after several years. The narrow entire leaves are
rather sparse and may be either green, as in consi))iih\'i and gravrolens,

1919] Hall-Goodspeed; Chrysil 193

dull green as in viridulus, or gray as in gnaphalodes. In typical
nauseosus and in hololeuciis the foliage is entirely covered by a beauti-
ful soft and white woolly tomentum.

The flowers are yellow, very showy, and grow in heads arranged
in dome-shaped to oblong clusters terminating the stems. The plants
are therefore highly ornamental during the flowering season, which
extends from August to October. At that time the large round-topped
shrubs are crowned with a profusion of golden-yellow flower-
clusters. As winter advances the flowers wither and fall, the foliage
becomes more and more sparse, and finally only the naked gray or
dull green twigs and empty involucres remain. The young twigs then
die back for a considerable distance. But soon after the first winter
rains or snows new shoots, springing from lateral buds well down on
the year-old wood, form a new top to the plant. Since the die-back
of the annual shoots does not extend to the base the general height, or
level, of the old wood is increased each year. At the same time there
is a thickening of all of the older stems through the addition of another
annual layer of wood. The flowers of Chrysothamnus, which are very
small and assembled into heads, as in all the Compositae, are much
visited by honey bees and other insects. This assures cross-pollination
and there is no evidence that the flowers are self-fertile. Seeds are
set in abundance, and they show a high percentage of viability. These
characters enable the plants to occupy quickly any suitable area that
has been cleared of other brush.

The herbage of Chrysothamnus possesses a peculiar odor v/hich
varies somewhat with the different forms. The name nauseosus indi-
cates that the type of that species was disagreeably scented but this
we have not been able to verify. In viridulus, however, the green
stems when broken emit a very strong, disagreeable odor, especially
penetrating if they are burned in a closed room. On the other hand,
when the twigs and foliage of gnaphalodes and hololeucus are broken
in the hand the odor is exceedingly pleasant, suggesting a combina-
tion of tropical fruits and berries. It seems probable that this is due
to the presence of a volatile oil which might have commercial possi-
bilities if properly exploited, more especially as a by-product in case
the plants are brought under cultivation for their Chrysil.

The age of full-sized shrubs is quite variable, depending upon the
conditions under which the shrubs have grown and also possibly upon
the botanical variety concerned. An estimate based upon counts of
annual rings, places the average age of the more important kinds.

li)4 ( iiir<rsilii of ('(tl ifufiiid /' iihl iciit talis iit llottnni |\<ii.. 7

siicli ;is riridiiliis and itnisimilis. at about ('i<jlit Ni-ai's. Tliis is for
plants w iMoliiii;^- five of six pounds and ^i-owin^ in oi'dinai'v alkaliiu*
soil \villn>n1 at1(nlion or dist urliancr liy man. Plants seldom reach
full size lici'oic tlicy ai-c five ycai-s old and tlicy often increase in
weiji'lit up to tru xcai's and p'M'lia|)s lonti'ci'.

VI. I)lSTKl]^>rT10X AND HABITATS OF THE \AH10US

FORMS

Rubbef pi'odiK'iiig vai-ieties of wild Rabbit-])rusli are widely dis-
ffibuted in western North Amei'iea. They belon*"' to the Lower and
I'ppi'i- Sonoi'an life zones with slight extensions into the Transition,
and ran<ie in altitnde from about sea level in some of the desert basins,
to 8000 feet in the southern Colorado mountains. Some varieties of
('Jirysoiliamiius occur at even higher altitudes but they have not been
examined as to their I'ubber content. The ])lants are most abundant
and of maxiiinnn size in the Great l^asin area. l)eeoming moi'e and more
scattered and aji]>arently diminishing in tlieii' percentage of rubber
as we pass outwai'd from this center of disti'ibution. The northern
limits are reached in Britisli Tohnnbia, Alberta, and Saskatchewan,
but no detailed studies have been made of ])lants from those districts.
They probably contain rubber in only limited (piantity and the areas
covered by the shrub are not extensive. Oui* northern tier of .states
supports some Chrijsothamnus but only in the more arid portions such
as southeastei'ii Washington and southern Idaho aiul Montana. The
most northerly points from which we have taken sami)les for analysis
are in eastern Oregon (Redmond, Burns, ete.^. eastern Washington
(Spokane), southern Idaho, and southern Wyoming (Rawlins, Lara-
mie). The best sami)les carried only -5 \wv cent of I'ubber and most of
them ran less than two jx'i- cent. This low yield uuiy be due to the
habitat oi- it niay be that a larger series of examinations would show
that we happened to select oidy low-gr;nle indixidu.ds and that the
average is really as high as in districts lyijig farthei- to the south.

The eastei'Iy limits of the geinis are reached in South Dakota and
western Nebraska : the southerly limits in western Texas, southern New
Mexico, and southeni Arizona, with some ])ossible extensions into
Mexico, or at least into Lower California. Analyses have not been
made of plants fi-om these outlying (listi"icts but the indications are
that they ai'e not good rul)l)ci' |)i-o(lueers. On the Pacific Coa.st we find

1919]

Hall-Goodsppfd : ( 7/ ri/sil

195

scattered groups of the plants as far west as the Coast Range moun-
tains, for example, San Benito County, Mt. Hamilton, Lake County,
Trinity County, etc. Four samples of moharensis from Mt. Hamilton
averaged only 0.53 per cent of Chrysil.

r*;.** ^M.^^^.

'^??*

Fig. 1. ChrysoflKuiutiis iKtuseosiis var. riridulufi. Plant no. 468. Seven feet
liigh; growing in alkaline soil with Distichlis spicata and Ira axillaris. Artemisia
tridentata in background. (loat Ranch, north of Mono Lake, California.

Coming now to the more important areas centering around the
Great Basin, we find that some of the densest stands and also the
largest plants are found in the alkaline deserts of Nevada and eastern
California. There are some large and important areas of the brush
in southern Colorado and in Utah, but the plants in these states are not
as good rubber producers as those farther west. Throughout the whole

196 University of California, Puhlications in Botany V^oh. 7

area Rabbit-brush is confined to certain definite tracts ranging in
size from a few square feet to about 800,000 acres. It is prevented
from growing in other places, sometimes by unfavorable soil, moisture,
or climatic condition ; sometimes by the more aggressive habits of other
species with which it is obliged to compete. For example, when the soil
is very strongly alkaline it is either devoid of vegetation or supports
such strongly alkali-tolerant species as Grease-wood (Sarcohatus),
Pickle-weed {Salicornia) , Sea Blite {Suaeda), and Spirostachys.

Where the alkali is somewhat less in amount but still too abundant
for upland plants certain varieties of Rabbit-brush, particularly the
green forms such as viridulus, graveolen<f, and consimilis become dom-
inant and with Salt-grass (Distichlis) , Poverty -weed (Iva axillaris),
and similar plants cover areas of considerable extent. Here are found
the most extensive and important stands of rubber-producing shrub
and the percentage of rubber is fairly higli. Tlie soil is somewhat
moist but too strongly alkaline for standard agricultural crops and
since the native grasses and other forage and browse plants are of but
little use to stockmen the value of the land is very slight. The removal
of the Rabbit-brush would favor the growth of grazing plants, as has
been demonstrated by our experiments. However, the brush would
again assert itself in a few years unless held in check. If it is found
practicable to grow Rabbit-brush for its rubber, these extensive tracts
could be had at a very low price and without serious interference to
any established industry.

Passing upwards from the alkaline flats just described one often
comes to better drained and scarcely alkaline slopes. Here the Sage-
brush {Artemisia tridcntata) is usually dominant and forms the ex-
tensive brush-lands of the Great Basin area. It mixes more or less
with the Rabbit-brush varieties mentioned above, but it is less alkali-
tolerant and so yields to them on the lowlands. On the slopes and
higher plains and especially in loose soil, or along dry water-ways,
the Sage-brush is often mixed with or replaced by another series of
varieties of Chrysothamnus. These are the gray forms, particularly
nmiseosus, speciosiis, gnaphalodcs, hololeucus, and frigidus. The
green and the gra}^ forms intermingle slightly where they meet in soils
of moderate alkalinity and moisture, but the latter are much more
intimatel.y associated with Sage-brush. In the terminology of the
ecologist, the Sage-brush is climax, the gray Rabbit-brush is subclimax.
In other words if the former is disturbed b}' fire, trampling, washout,
etc., it is replaced by the Rabbit-brush, which, however, is in turn

1919] Hall-Goodspecd: Chrysil 197

slowly crowded out by a new generation of the more aggressive Sage-
brush. The result is that the gray forms are much more scattered
than the green ones and almost never form pure stands of any great
extent. Thej^ are sufficiently abundant in some places to yield a
supplementary supply of rubber in case the green forms are harvested
from near-b}- areas, but they should not be considered in making esti-
mates of the total amount of native rubber available. Their chief
interest lies in the fact that they are very erratic as rubber producers
and might therefore furnish starting points in case experiments were
undertaken in the breeding or selection of superior strains.

VII. ESTIMATES BY DISTRICTS OF THE AMOUNT OF
CHRYSIL AVAILABLE IN WESTERN NORTH AMERICA

Although field studies on the occurrence and distribution of rubber-
bearing shrubs have been carried on in all of the western states, actual
estimates have been made for only a few of the areas upon which they
are known to grow. The sudden termination of the war has removed
the immediate need of a complete survey, but if an emergencj^ call
is ever made the work thus far completed will furnish a starting
point for a definite estimate of the total amount obtainable as well
as indicate the localities where it may probably be found in the largest
quantities. Somewhat detailed estimates for the areas examined are
therefore presented by districts.

This phase of the work has been carried on chiefly by the senior
author, who has himself visited a majority of the areas reported upon
and who assumes full responsibility for the estimates here given. His
studies have been supplemented by those of a number of other botanists
as mentioned on page 186.

The estimates were prepared in the following manner. Districts
where the brush grows abundantly were visited and traversed when
possible in several directions. Data thus accumulated, supplemented
by such information as could be obtained from reliable residents,
furnished rough estimates of the acreage covered, the percentage of
vacant land within the area, and the ratio of Rabbit-brush to other
shrubs. Small quadrats were then selected and the number of plants
per acre determined. Average samples were weighed or at least
estimated as to weight. From these data the total tonnage of shrub
has been calculated. A sufficient number of samples has been analyzed

198 Viiirf rsili/ of Califorxiit ruhllcdtiniis hi TiotiDiu ' I Voi,. 7

I'ro'ii widely st'pai'alcd <i('()<ii'a|)liic afciis, and ol' the many l)otaiiical
varieties to fnriiisli sonietliiiii: of ;i basis for the ealeidation of tlie
total amount of cnlilie!- pi'esent on any jrivoii area, tlie total tonnajie
liavin<i' been di'tei'niined in the manner just described. AVhile these
results are fai" from satisfactoi'X' beeause of the mea<>er data upon
which they are iiaserl they will fui'nish at least a b('<riniiin^ in case
move extensive iiivestifi'ations are inaufjurated, and will indicate the
best locations for rubber mills in case the p-overnment is ever oljji^cd
to resoi't to Rabbit -biMish as a soui'ce of rubbei- in time of emero-eney.
The areas indicated in the following' list include, we beliexc, most
of those where Rabbit-bi'ush is abundant, yet it is (piite certain that
at least a few larj^e areas have escaped our hasty held survey. It is
well know n that, in addition, there is a multitude of small fields of the
shi'ub and also that it often grows scatteringly whei'c other species are
dominant. However, because of the gi-eat expense of assembling shrub
from small scattered fields, this source is not included in the estimates.

a. DISTRICT I— EAST CENTKAL CALIFORNIA ANF) A1).)A( K.\T

NEVADA

This district comprises Tnyo and Mono counties in California, and
an extension into Esmeralda County, Nevada, chiefly to include Fish
J^ake Valley. In it are found large areas of shrub carrying over ;! per
cent of rubber, although the total area of shrub is much smaller than
in some of the other districts where the percentage content is less.
The chief variety included in the estimate is viridnlus, which is larger,
on the average, than any of the others. It occurs chiefly on the alka-
line valley bottoms but ranges sparingly to altitudes of 7200 feet in
alkaline soil. Mixing sonunvhat with it around the borders of the val-
leys, where the soil is better drained and less alkaline, are considerable
quantites of gnaphalodcs, Jiololciicus. and mohavrusis. These extend
well uj) the foothill slopes but always as mixtures in the Sage-brush
climax — never as pure stands. North and east of Mono Lake, virididiis
is replaced to some extent by coHsintilis. Thei'c is also a supi)leMientary
supply, furnished by th(> very considerable amount of Ch ri/snllidmuiis
teretifolius, a low shrub carrying about 2.7 pei- cent iut)bci': but since
the plants are somt>what scattered it does not enter into our estinuites
(see p. 266).

Some of the l)est shrub in this district is found in Owens N'allcy.
where it grows on most of the flats along the i-iver; in an irregular
strij) along the Los Angeles aqueduct ; and on isolated areas of from

1919]

Hall-Goodspeed : (Jh rysil

im

one-half to several square miles in extent. Some of the largest plants
seen anywhere grow near the aqueduct (but not irrigated from it) a
few miles north of Lone Pine. The district around Keeler has not
been examined, nor has Long Valley, although both are reported to
have some Rabbit-brush. A considerable area in the lower part of
Deep Spring Valley is also said to be covered with Rabbit-brush of the

Pig. 2. Chijisdiluinnuis iiauseosus var. riiiduhts, iiuiiolino- with var. f/iiaph-
(tJodcs. Experinieutal tract. Lone Pine, California. Soil moderately alkaline,
much trampled. Sierra Nevada Mountains in background; Mount Whitney at
ritjlit of center.

riridulKs variety. If liarvcstcd, ])r()l)ably the best place to take it for
milling would be Owens A^alley. Small lots are also obtainable to the
northeast of Owens Valley, particularly in the vicinity of Benton,
whence come our richest samples of inriduhis, and in Chidago Caiion,
etc. Our estimate of the total amount of rubber in Owens Valley and
contiguous territory is 300,000 pounds.

Fish Lake Valley, in western Esmeralda County, Nevada, is in-
cluded in District 1 since the size, habit, and variety of the shrub is
similar to that around Mono Lake and in Owens Valley. The exact
botanical classification of the plants, however, is not satisfactory. They

200 University of California Puhlications in Botany [Vol. 7

grow in varying degrees of density over most of the valley bottom
except wliore farming operations have interferred. An estimate based
upon a hurried visit in 1918 and upon data supplied by Forest Ranger
George Parke places the total amount of rubber at 2,280,000 pounds.
Extensive areas of Rabbit-brush have been located in the vicinity
of Mono Lake, but our analyses indicate that it is not so rich as that
from Owens Valley and Benton. The principal varieties represented
are viridulus and consimilis. There are about ten square miles of
fairly large plants to the north and northeast of the lake in a nearly
pure stand. On the south and southwest sides it is mixed with Sage-
brush over an area of perhaps sixteen square miles. In Adobe Valley,
which lies southeast of the lake, we find it in good formation from the
Adobe Hills to the River Spring Ranch and in a narrower belt as far
as Dutch Pete's while there are perhaps a thousand acres of it in a
belt extending from Gaspipe to Indian Spring. By using the methods
indicated above, we have estimated that these areas centering around
]\Iono Lake would yield about 700,000 pounds of rubber.

The estimated total amount of rubber obtainable from District 1 is
3,280,000 pounds.

1). DISTRICT 2— MOJAVE DESERT, CALIFORNIA

This might almost as well be called the "Antelope Valley District"
since it centers around this westerly arm of the Mojave Desert. It is
not very promising as a rubber producer because of the large number
of varieties represented and the consequent fluctiiation in rubber con-
tent. The largest continuous area of shrub is a belt of gnaphalodes
about two miles wide and perhaps thirty miles long, which extends
diagonall}' across the valley in a northwesterly direction from near
Palmdale to a point west of Rosamond. The shrubs are below medium
size, the woody portion weighing on the average about three pounds,
and there are about 50,000 plants per square mile. A similar belt is
reported still farther west. These belts are undoubtedly due in large
part to the burning off or clearing of the Sage-brush in the remote
past and represent one stage, in the succession whicli will ultimately
bring this Sage-brush back as the dominant climax species. Analyses
indicate that this shrub carries an average of two per cent of rubber.
Smaller areas of gnaphalodes are scattered all around the borders of
Antelope Valley and even to Tehachapi Pass and northward to Owens
Lake, mixing slightly along its upper limits with moliavensis, but the
lower, more alkaline lands support a scattered growth of what ap-

1919] Hall-Goodspecd: CJinjsil 201

pears to be a dwarf form of viridulus. Most of these small plants
carry less than two or three per cent of rubber and are of but little
practical importance.

C. teretifoUus, carrying about 2.7 per cent of rubber, grows sparse-
ly on the Rosamond Hills but may be abundant on the mountains
to the west of Antelope Valley. Around the borders of the Mojave
Desert, especially toward the west, are scattered stands of two other
low-grade rubber shrubs, Haplopappus monactis and H. lineanfolius,
but these are of little value.

The amount of rubber in the shrub which we have located in Dis-
trict 2, taking no account of the species mentioned in the preceding
paragraph, is estimated at 400,000 pounds.

c. DISTEICT 3— NOETHEASTERN CALIFORNIA AND ADJACENT

NEVADA AND OREGON

The principal areas thus far located in this district are on the alka-
line plains of eastern Lassen and Modoc counties, California, where
Rabbit-brush, in the consimilis form, makes pure stands over many
thousands of acres. The percentage of rubber in these plants is ap-
parently quite uniform and runs a little more than two per cent. There
are about two square miles east of Honey Lake where the plants aver-
age sixteen feet apart and two representative samples weighed ten and
twenty-eight pounds, respectively. In the vicinity of Karlo we find
about seven square miles covered by a nearly pure stand of the shrub ;
an area of about the same size occurs on Painter 's Flat ; and still an-
other to the west of Madeline. The plants on these tracts are often
smaller than in the Honey Lake Valley (two taken near Karlo weighed
under two pounds each for the woody portion), but when small they
grow closer together. There is an area of 1000 acres south from
Ravendale, covered with medium sized shrub. About twenty square
miles have been located by Mr. L. S. Smith in northeastern Modoc
Countj^, where the Rabbit-brush comprises about 25 per cent of the
vegetation, but it occurs chiefly in one of the gray-foliaged varieties
and is therefore poor in rubber. There are some unexamined areas of
consimilis west of Alturas, several sections of the gray form in Butte
Valley and around Klamath Lake, and some extensive areas of con-
similis on the Klamath River drainage of north-central Siskiyou
County, California.

In southeastern Oregon, the Rabbit-brush was found by Mr. J. R.
Bruff to exist only in small stands. The largest area located is in

202 (' n ii'i rsil 1/ of Ciilifoniiii I'lihlicdtifjiis in /Sohnni 1 \ mi.. 7

lUitzcii \';illcy. where tliei-e ;ire peflui|)s Iwehc s(|iifire miles of cov-
siinilis. and in ('atldw \'alley. where thei'e is a soiiiewliat sniaUei' area
of the same t'ofm. SUulies made hy the seinof anthof fi'om IJurns to
Rend and northward indicate that the phuits there gi'ow too si)ars(»ly
to be of interest anil that tlie percentage content is not nnifoi'm. Re-
ports of hirjre ai'cas at l'>nft"alo Meadows and on the I5hici< Rock Desert
of Nevaihi wci-e fonnd to l)e ei-roneons and it seems that there is hnt
litth' of the shrnh in the nort hwcstei-n corner of that state. It is in-
teresting to note, however, that two sanij)h's of tlie variety ro))simiHs
gathered along tlie edges of the alkali flats neai' (lerlach yielded tlie
highest I'etnrns of all of the samples taken of ('hri/si)fli(iiinnis ti(iHS(t>-
SKS. They analyzed 4.71 and 0.57 ])er cent, respectively, of pni-e
rubber.

The most impoi'tant areas in tliis district are to he fonnd, there-
fore- in eastern Lassen County, California, and in IMitzen and Catlow
valleys of southei-n Oregon. The total amount of rubber in the shiMibs
thns fai' locati-d in District ?> is estimated at abont 1 ,000, ()()() ponnds.

,1. DISTRICT 4— WEST CENTRAL NEVADA
This small disti'ict lies wholly in Nevada and is between the C'ali-
fornia boundary and a line connecting Hazen and Tonopab. (^n the
north it is limited by tlie line of the Southern Pacific Railway; on the
south, liy an east-and-west line passing through Tonopab.

The only ^■ariety of ('liri/solh<ninn(s to be considered here as a
possible source of rul)ber is (■())isinulis. but this grows in abundance in
valley bottoms whei-e the soil is somewhat sandy, slightly moist, and.
for the most part, too alkaline for Sage-brush. Around the edges of
such areas, and rnnning back into the hills, it is not uncommon to
find otliei' varieties, especially those of the gi'ay types like (/risi iis.
nauseosiis. and sixciosiis, liut these occur as impurities in the Sage-
bi'iisli consociation and ;ire therefoi-e too scattered to be of coiise(|iience
as a possible source of rubber.

The consimiiis foi'iu varies much in size and habit, deixmding in
these characters upon soil and clinuitic conditions. Excejitionally large
])lants were encounterc^d on the flats at the head of Walker Lake.
These wei'c six feet high and almost tree-like, with ti-unks five inches
thick and over a foot in length to the first nuiin branches. Such plants
have a weight of atioiit thii'ty ])ounds, exclusive of twigs. The aver-
age weight of samples, howevt'r, taken from along Walkej- River, north
of the lake is al)0ut ten i)oun(ls, while the average for the whole dis-

nini!

IldU-ddodspci d : ( 7/ ///.v/V

20:]

trict is nearer six pounds. The rubber content of seven samples taken
at widely sejia rated stations in the district averages 2.0 per cent and
microscopic examinations of others indicate that th(>re is not a wide
fluctuation from this mean.

Fig. 3. ChrifSotJuimnus nauscosus var. consimilis. IMant no. 21(5, Walker Lake,
Nevada. Root two ami three-eighths inches thick ; trunk five inches thick at base
of first branch; total height six feet; estimated ^veight about thirty pounds. This
is an exceptionally large plant.

The southernmost areas of Rabbit-brush within this district are in
Esmeralda County and lie along the Tonopah and Goldfield Railroad
from Redlich nearly to Millers, one of them extending as a belt along
the westerly side of the San Antonio Wash. Small areas of large brush
were found at the upper end of Walker Lake, but mucli more import-

204 University of California Publications in Botany [Vol. 7

ant is the irregular belt which skirts the river on both sides from a
point about nine miles above Schurz nearly to Wabuska, a distance of
eighteen mih-s. This belt will average three-eighths of a mile in width ;
the plants are ten to twelve feet apart and have an average weight
of about ten pounds. Forest Supervisor W. M. Maule reports about
fifty square miles of shrub from Whiskey Flat north to Hawthorne;
also about thirty-five square miles between Sweetwater and Fletchers.
These areas are in western Mineral County. We have not ourselves
visited them and liave not had an opportunity to examine samples.

The Carson Valley and contiguous areas support about one-hun-
dred and fifty square miles of Rabbit-brush of the consimilis form.
Several other forms are present, some of them apparently devoid of
rubber, but the}^ grow scatteringly around the borders of the main
bodies and extend up over the slopes as impurities in the sage-brush.
It is on the lower, more nearly level, and alkaline flats of the valley
that the best stands of consimilis are found. Here it often grows to
the exclusion -of other shrubs, and almost completely covers the ground
with plants weighing three to five pounds.

According to our estimates, District 4 should yield at least 7,680,000
pounds of rubber, of which nearly one-half is in or near the Carson
Valley. It is probable that further exploration would result in a con-
siderable addition to this estimate.

e. DISTEICT 5— NOETHERN AND CENTRAL NEVADA

By E. C. McCarty

This district comprises the counties of Humboldt, Leander, Eureka,
Elko, White Pine, and a part of Nye County. In view of the many
areas in this section which are favorable to the growth of Chrysotham-
nus, it would without doubt prove to be an important one in the pro-
duction of an emergency supply of raw rubber. The long valley of
the Humboldt River, extending through the counties of Elko, Eureka,
Leander, and Humbolt, has many tributary valleys extending both to
the north and to the south, and ranging from comparatively short
valleys to valleys of more than fifty miles, the Reese River Valley be-
ing one of the longer ones.

The Reese River Valley, which is perhaps typical, is half a mile
wide at Caiion, and more than fifteen miles wide at Austin. It extends
the entire length of Leander County and into Nye County. Tht' moun-
tain range is continuous on either side of the valley and is marked at

1919] Hall-Goodspeed: Chrysil 205

frequent intervals by draws and gulches from which flow the streams
tributary to the Reese River. This river has an interrupted flow
through the greater part of the valley though it is itself a very small
stream. The floor of the valley is comparatively level, and rises grad-
ually to the mountains on either side. The altitude, which is four thou-
sand five hundred feet at Battle Mountain, increases at the southerly
end to upward of six thousand feet. The soil is alkaline and is of the
type usually found in arid regions. The precipitation is meager ; the
greater portion probably occurs in the fall and winter. The summers
are comparatively long and the number of days of sunshine in one year
probably approaches three hundred. The coldest temperature experi-
enced by the writer while in this valley during the month of December,
1918, was -15° F.

The character of both soil and climate permits the growth of only
drought-resistant vegetation. Generally speaking, the lowest lands
of the valley floor are occupied by Chrysothamnus, while the more ele-
vated portions are covered for the most part with Artemisia. Much
of the lowlands is covered with pure stands of Chrysothamnus. Again
large tracts occur in which this shrub represents fifty per cent of the
entire stand. Along the river bottom the dominant shrub is Chryso-
thamnus, but in many places in the immediate vicinity of the stream
the soil appears to be fairly moist in character, and is occupied by
willows. In such sections, which are comparatively few, Chrysotham-
nus occupies the zone next to the willows and on either side of the
stream. At many places in the valley the brush, in stands more or less
pure, extends into the draws and gulches. Where the Chrysothamnus
forms occur to the practical exclusion of other types, the stands are
quite dense, the plants averaging from four to ten feet apart. It is
believed that the pure stands occupj^ those parts of the valley in which
the physiological drought of the soil reaches the maximum.

The pure stands of Chrysothamnus throughout the entire district
are for the most part of the consimilis form. While there are some
comparatively small areas of the gray varieties, these seldom occur
in pure stands- and none of this type are included in the estimate for
the district. The size of the green brush is variable. Some of the
plants are quite large, the height being three to eight feet. While
some of the larger ones would weigh as much as twenty-five pounds,
it is not difficult to find others that would weigh still more. The
average, however, is much lower, and the weights employed for the
purpose of the estimates in this report are between two and seven

206 r iiiff rsilif of Califoniid I'lihlicatioris in Jidfaiiji | Voi.. 7

poiuids. Tlicsf fifj^iii'fs. liowcvcr, ;ii'<' l)asc<l on llic wcijjlit of a small
j)Ortioii ())' the I'ool. and ot" tlic siciiis upwai'ds (if four ycai's (A' a^c
Slidiild it licconic (icsii'atilc 1<t use tin' younger wood the avcrajic woijjht
would he increased hy one to two j)Ouiids. It is probable that the
amotiiit of bnisli now available will I'eniain constant for a nunil)('r of
years, sinee tlic cliaraetci' of tlie soil and climate is moi'e favoi'alile to
stock raising' than to a;ii'icult nre.

While many of the jtoints in this district jiavc been visitecl and ai'e
known to contain lai'iie ti'acts of Cli ri/sotlKtm n tis. reliable information
lias been recei\-ed in otliei' ways relative to tile occnrrence of lai'^e
stands in those sections which were not \isitei|. The character of this
information and the source whence it has come wai'i-ant the inclusion
of th(>se sm'ei'al ai'cas in tlu^ estimate of available brush in the district.
On tlie other hand the coiubiniHl areas reported on and includeil in the
estimate are but a part of the entire territory comprising: the district.
In \iew of the similai'it>' in tojiograjihy, in climate, ami in soil condi-
tions pi'cvailing thi'oughout this entire region it is believed that a more
intensive survey of the district might reveal still other ti'acts of tlu'
shrul) which in the aggi"egate would ecjual those included in this rcpoi't.

The principal areas thus far located are the following:

Elko ("oinity: Ten Mile Flat: North Fork. Hmuboldt River Val-
ley and tributary valleys; along the main line, Southern Pacific and
Western Pacific railroads; Ruby Valley; along the Nevada Northern
Railroad. Cobre to Goshute.

Eureka County: Along the main line, Southern Pacific and West-
ern Pacific I'aili'oads; Antelope Valley.

Humboldt County: Areas lying 6 and 15 miles, respectively, north-
cast of Winnemucca ; Rebel Creek; Quinn River Valley.

Leander and Nye counties: Along the main line of Southern
Pacific and Western Paciiic railroads; Reese River \alley : Big Smoky
Valleys (north v]u\) ; Monitor A^allex- : Fish Spring.

White Pine County : Uutte Valley ; Long Valley ; Newai-k Valley ;
S|)ring \'allcy ; St(^])toe \'alley: White Rivei' \'alley.

The total weight of Rabbit-biiisli known to exist in this district has
been placed at 1,250, 000, 000 jxiunds. Twenty-thi-ee samjdes have
been gathered and analyz(Ml. The rubi)er content of these samples
vai'ies from 0.4o to 8.86 per cent, with an average of l.Si) ])er cent.
On the basis of these figui'es the total available su|)|)l\- thus far located

'■' It is r(>p()it<'<l tli;it ('hi!i>:i)lli<innti(.s is juiiseiious to laiijic stork iluiiii^ tlio
winter season.

1919] Hall-Good speed: Chnjsil 207

would be 23,700,000 pounds of pure rubber. It must be borne in
mind that these estimates are based on the weight of plants after the
removal of the foliage and of all stems under four years of age.

/. DISTRICT (5— UTAH

Within the state of Utah, Rabbit -brush in several forms occurs
over extensive areas, especially around the alkaline valleys of the cen-
tral and southeasterly parts of the state. Much information as to the
location of shrub in quantity has been assembled by Professor Marcus
E. Jonf s, but an examination of the samples supplied by him indicates
that the rubber content is less than that of plants from Nevada and
California. The analyses we have made of his plants average only 1.12
per cent for the consiniilis variety, which is by far the most abundant
one, but this is perhaps too low an average for all of the Utah shrub,
since three samples of consimilis collected by ourselves near Stockton
carried 1.31, 2.46, and 2.84 per cent of rubber, respectively. What-
ever the exact average may be, it certainly is low and renders unneces-
sary the publication of detailed information as to distribution. The
largest areas of shrub in Utah lie along the alkaline flats of Sevier
Valley, Tooele Valley (and south to beyond Vernon), Tintic Valley,
Rush Valley, Utah Valley, Juab Valley, and the valleys of the Price
and Green rivers. B}' far the largest stand of shrub is along the
Sevier River, where Professor Jones finds a strip about one-f(mrth
mile Mnde by one hundred and fifteen miles long, extending from
Panguitch north to Fayette, and he describes other large areas in that
valley. In the vallej's of northeastern Utah, such as the Uinta Basin,
Chrijsothamnus occurs only in small, isolated patches and is of no
importance in preparing estimates. In northwestern I^tah, the varie-
ties present are those which carry but little if any rubber and tliere
are no extensive areas even of these. To the southwest, however, there
are some stands of consim.ilis that should be taken into account. There
is one area of at least 20,000 acres along the line of the Salt Lake Rail-
road and farther south than any of those mentioned above. Tlie
shrubs are closely placed but quite small, the average weight being
estimated at scarcely over one pound. A sample taken at Milford
yielded 1.24 per cent of rubber. A single sample of ('. tiirhinatus
gathered on the plains near Lund analyzed 4.88 per cent of rubber.
This species is too scarce to be of impoitance save as a starting point
in breeding experiments.

208 VnivtrsUy of Califoriiiu I'lihlications in llolatiu l^'"'-"

According to Professor Jones, the avernge wciglit of Utali plants,
after cntting off tlie twigs, is about fifteen pounds, but in ordei- to be
pci-fcetly safe \v<' have taken only t<'ii pounds as the weight in uiaking
up oui" estimates.

Fi'om the dala indicated ;d)Ove and taking 1.12 per cent as the
jiverage eonlent. we estimate tlie total amount of rubber present in
('liri/sofhanDius in t'tah to be not less than 20,000,000 pounds.

(J. DISTRICT 7— (COLORADO

Our explorations in this state luive been far from exhaustive, con-
sisting Duly of one visit to the northwestern part, a north-and-south
journey along the easterly base of the Rocky Mountains, a visit to the
San Luis Valley, and notes and samples taken along the line of the
Denver & Rio Grande Railroad. The only important area of (lirijso-
thamnus that we have located is in the San Luis Valley of south cen-
tral Colorado, where we estimate, from observation and from rei)oi-ts
of reliable residents, that not less than S()(),000 acres are fairly well
covered with (Uiri/sofliamii us of the pinifolins variety. Over much of
this area the plants stand about ten feet apart, that is, tliere are about
four hundred and thirty-five i)lants to the acre. Since Grease-wood,
or Cliico {Sarco'batus vermicid'atus) and other impurities occur in
some portions of the valley, we have taken only one-half of this
number as a basis for estimates. The plants are of medium size, the
average weight after the removal of the non-rubber-bearing jiortions
being perhaps five pounds. Two samples taken at Alamosa yielded
3.41 per cent and 1.11 per cent, respectively. A third, sent fr(mi near
Center by Mr. Wm. 0. Sander, analyzed 3.59 per cent. This makes an
average of 2.70 per cent of pure rubber. On this basis the total amount
of rubber present in this one valley would l)e about 24, 300, 000 pounds.
This is probably the largest single body of shrub in Colorado, if not
in the whole West. Smaller but otherwise siinilar areas doid)tless occur
in some of tlie unvisited river valleys of the southwestern i^u't of the
state. ^'^ Along the Gunnison we found only small detached areas of
Rabbit-brush and the same is true of the valley of the Yam[)a, in north-
western Colorado.

The most easterly stations at which we have found rubbei'-bearing
shndis lie along the east base of the liockx' ^iountains from Fort Col-
lins and Denvei' to Trinidad. Five samples of grovcoJnia from these

1" See }>. ISO. footiioto,

1919] Hall-Goodspccd : Ckrijsil 209

places averaged less than 1 per cent of rubber and nowhere did we find
other than small groups of the plants. A single sample of frigidus
from Laramie, Wyoming, was found to carry 1.86 per cent of rul)ber.
A sample of pinifolius from Salida yielded 4 per cent.

Our estimate of the total amount of rubber present in Chnjso-
thamnus in Colorado is based entirely upon Avhat was found in the San
Luis Valley and is therefore, as indicated above, about 24,300,000
pounds.

//. ESTIMATES NOT IXCLUDED IX THE ABOVE DISTRICTS

There are many districts in addition to those just enumerated
that will need to be explored before a final statement as to the occur-
rence of rubber-bearing shrubs can be made. Large bodies of these
plants might be found in any of the western states. The most promis-
ing of these, in addition to those already specifically mentioned, is per-
haps Wj-oming. It is well known that this state supports large areas
of Kabbit-brush but aside from a few samples taken along the southern
border Ave have made no study of its rubber possibilities. "We have
been told also of considerable areas in southern IMontana, in Idaho,
and in eastern Washington, but judging from the varieti(^s there pres-
ent the percentage of rubber in the plants is very low. In Nevada,
which we have eovei'ed as well as any other state, there are still some
promising unexplored fields. Chrysothamnns occurs in a number of
species and varieties in New Mexico, according to good authority,^ ^ but
we have scarcely considered that state. Professor A. 0. Weese reports
a great deal in the region around Albuquerque. Arizona is likewise a
state of possibilities, but the areas of Rabbit-brush to be found there
are probably quite limited in extent. Tliese two southerly states should
be carefully explored in case a search is made for new strains of high
quality, since a considerable number of species and variants not yet
examined are known to occur and tlie climatic conditions are such as
would be expected to result in the formation of rubber.

(. CONCLUSIOXS AS TO THE QUANTITY OF EUBBER OBTAINABLE
As previously noted, the explorations in search of rubber plants
have not extended to all parts of the West, nor, indeed, has any single

11 Vernon Bailev. Life Zones and Crop Zones of New Mexico. N. Am. Fauna,
no. 3.5 (1913), pp. "28. 31, 37.

Wooton and Stamllev, Flora of New Mexico. Contril). T^. S. Nat. Herl)., vol.
19 (1915), pp. 660-(563.*

210 Unir< r.slhj of California ruhlicaiions in Botanij [Vou 7

stati' hccii tlioi'onjjlily cxainincd. Tlic seven districts reported upon,
liowever, furnish some indieatioii of tlie total amount of rubber tlial
mi^lit be obtained in case of need. The estimated amount alread.N- lo-
cal e(l by districts is as follows:

Pounds

J)istik-t ] — East Central Calit'oniia and adjai-cnt >.'i'va(la 3,280,000

District 2— Mojave Desert, California 400,000

District 3 — Northeastern California and adjacent Ncvaila and Oregon 1.000,000

District 4— West Central Nevada 7,(i80.000

District 5— Northern and Central Nevada 23,700,000

District (5— Utah 20,000,000

District 7— Colorado 24,300,000

80,360,000

Taking this total of approximately 80,000, 000 ])()uii(ls as represent-
ing the ])robable amount of pure rubl)er present in the shrub located,
it is believed that this estimate could be safely increas(d by 50
per cent by allowing for the presence of other areas within these dis-
tricts, althoufih we have not been able to visit them. After this has
been done, we have still to account for several whole states as well as
several fractions of states lying outside the seven districts noted above.
We hesitate to venture a guess as to wliat tliese might yield but it
seems i)robable that the total amount in all of the western states is
not less than 300.000,000 pounds.

VIII. METHODS OF DETECTING THE PRESENCE OF
RUBBER AND DETERMINING ITS AMOUNT

(I. :\IICROSCOPICAL METHODS

Microscopical examinations of Chrysothamnns and related genera
were at first undertaken as a short cut method of determining whether
or not a given sample of shrub contained rubber. In addition we liave
attempted, with some success, to emjiloy the microscopical method for
estimating the relative amounts of rubber in the different samples. At
the beginning of the investigations we were forced to rely ui)on this
method almost entirely since facilities could not be secured for nuiking
ourselves the many chemical analyses which luivc since been carried
through. Chemical analysis is, of course, the only method of making
both the oi'iginal determination and of obtaining an estimate of rubber
present which is at once thoroughly dependable and i-eally significant.

1919] Hall-Goodspccd: Chnjsil 211

The microscopical method, in brief, consists in the cutting of cross
sections of a mature portion of the sample and the making therefrom,
after the action of suitable solvents, stains and mounting media, of a
preparation in which the rubber, if present, may be seen under the
microscope. As will be shown in the detailed description given below,
the process of making such a preparation is simple and may be suf-
ficiently well controlled to yield results that will be fairly uniform and
reliable.

As in Guayule, the rubber in Clirysothamnus and Haplopappus
is present within the individual cells of the plant, and is not a latex
rubber. Like Guayule again it is found principally in the parenchy-
matous elements of the cortex ; indeed it may occur in any undifferen-
tiated elements lying without the cambium or, in other words, in what
is often spoken of as the "soft" or "inner bark." It may also be
noted here that rubber does not appear to be laid down during the
first year of growth of a tissue and indeed, unless present in large
amount, is not readily detected by the histological method in por-
tions of the plant less than three or four years old. A section of a
plant taken slightly above the soil line will exhibit tissues containing
practically maximum quantities of rubber. In investigating new and
untried species by the microscopical method it is wise to make sections
from well down the root as well as at the soil line. This is indicated
by our results on Haplopappus mentioned in the third part of this
report. In certain species of that genus practically no rubber was
found in sections of stem tissue, while quite appreciable amounts were
shown in sections taken well down the main root. We have found
in Chrijsothaninus that maximum quantities are borne by the stem,
although for a limited distance below the soil line the root may bear
an almost equivalent amount.

In selecting and removing the material to be sectioned care must
be exercised to retain the bark. This is often diificiilt, as the dry
corky tissues readily split off, together with the living cortex, along
the delicate cambium region, from the hard, woody cylinder. When
the tissues are stripped off in this way it is useless to attempt the cut-
ting of sections, since practically the entire rubber-bearing cortical
area will be missing. It has been our practice to cut out the desired
piece of mature woody tissue^- and allow it to soak in water for twenty-
four hours before sectioning. This treatment softens all the tissues and

1^ We liave found that a c-opino; saw can be very successfully employed to cut
out the piece of stem to be sectioned.

212 r in'n rsil !i of (\il ifoniid I'lihlicdl laiis in lloidiiii | N'm,. 7

Toiidors iiinvo simple tlic lijiiidliiii;' of Ihc iiii|Hit't;iii1 l)tii'ky clenieiits.
The ad ii;il cross srcl ions jii'c iii.kIc with ;i sectioJi kiiifi' or h<';i vy-hnckcd
I'lizoi" iiol hoMow Lirouiid. 'The piece of iiuiterird may he h''hl in the
iiiij^'crs i'oi' ciiltiii^ or in a so-caUed hand oi' well microtome. One of
lis lias devised a modification of this 1\'pe of ndcrotome which was
nsed in these investijiations.'" A slidin<i' microtome has also hecii nsi'd.
Successful sections can he cut only with an extremely sharp instrument
and only a few sections can he cut heforc sliar])eninf; must he repeated.
Smoothing' the surface to he cut with a sluiip ])Ocket knife is desirable
and saves toi- a lime the liner edge of the sectioning razor. 11' interest
centers enlii'ely upon the (piestion of the pri'sence or absence of rubber,
it is not essential that the sections cut he paiiicuhirlx- thin nor that
they he taken exactly at right angles to the long axis of the sample. In
our work details of structui'e and other matters were often of special
im])ortance for which reason nW sections were cut less than .lO/j, in
thickness and were carefully oriented.

As noted by Lloyd^* rubber when jireseut in large amount is. after
some practice, rather readily detected in fresh or unstained sections
of non-latex rtd^her ])lants. When present in smallei- quantities it is
often indistinguishable fi'om the protoi)lasmic matrix of the cell or is
confused with accumulations of oil or resin in the tissues. According
to oui- histological method oils and resins were dissolved out because of
tlie i-elatively small amount of ruhhcr usually present and especially
as a j)reliminary to the use of a stain which was not definitive for
rubber i]i the pi-esence of these other sid)stances. As a solvent, acetone
was used exclusive]}', the sections being placed in a test tube half full
of acetone and allowed to stand in a water bath at 60° C. for from
fifteen to thii-ty minutes. For staining the sections we at first tried
alkanin,'-' hut Sudan III was early found to be more satisfactory and
was therefore employed in the great majority of the histological exam-
inations. Our Sudan TTT was made \\\) with glyceiine according to
Stevens' fonnula."' This stain imparts a brilliant seai'let coloi- to fats.
iTsins and i-uhbers as they occur in the cell. Wy the ])revious acetone
extraction fats and resins had been dissolved fi'oni the cell contents
hut rublxM- was left ])ractically uiuiffected and if i)i'csent took up the
stain. The stain was allowed to act for eic-htecn hours, after which the

'•'' (t()()ils]><'<i|, T. ]I.. Modification ol' liaml iiiiciotdiiic liot. (iaz., vol. ()(>
(1918), p. 5:54.

i-t Lloyd, F. E., Giiayulr. Canicoi,. Inst. I'lil.l. no. I .S9 (1911). y. 17(!.

i-^'Cf. Stevens, \V. (\. Plant anatom.v. i'.d cd. (191(;). p. 29:!.

!'•• L.C., ]). .*}:57.

191«I Hall-Goodspeed: Chrysil 218

excess was washed from the sections with 50 per cent alcohol. The
sections were then monnted in glycerine and in some cases were ringed.

Some search has been made for a staining method which would be
strictly definitive for rubber and which would result in imparting
such a chai-aetei'istic color to the rubber cell-inclusions as would serve
to distinguisli them definitely from all other inclusions. The desira-
bility of securing such a staining method is seen in tlie fact noted
above, that alkanin and Sudan III are in histological practice used to
indicate the presence of resins and fats, respectively, and thus their
ability to stain rubbei' in the cell is in a sense more or less fortuitous.
The fact that alkalin is a specific stain for resins might suggest a cor-
responding staining reaction in the case of rubber. As far as Sudan
III is concerned, however, we are employing a stain used in animal
histology to give definition to globules of fat occuri-ing in the cell.
Without going into the question of the chemisti-y and differential stain-
ing of the fats, fatty acids, and lipoids, it may be noted that the stain-
ing of fat globules by Sudan III is taken to indicate that this stain is
soluble in the contents of the fat globule whereas it is not solnblc in
the other constituents of the cell which, therefore, remain unstained.
It is somewhat difficult with this explanation of tlie characteristic
staining reaction of Sudan III in mind, to understand its action in
the case of rubber. This matter is mentioned simply to call attention
to the possible theoretical interest attached to the problem of the
staining of rubber inclusions in the cell.

We have, further, been interested in this matter because cell in-
clusions have been consistently found in clilorophyllous tissue of both
stems and leaves which, with Sudan III, stained as rubber, but which
were difficult to differentiate witli this stain from tlie residue of the
protoplast witli its included chloroplasts, which does not in such tissiii'
entirelj' disappear after acetone extraction and which stains to some
extent with Sudan III. In this connection it should be stated that in
other tissues also it is difficult to distinguish between the protoplasmic
matrices of the c(^lls wliieh may be stained with Sudan III and the
rubber inclusions wliieh may oi' nuiy not be present. Thus in the
tables which follow it is in some cases possible that the plants which
on the basis of "microscopical examination" iwo stated to contain
"traces" of rubber may have shown only stained cell inclusions which
could not loositively be identified as rubber.

We have made some preliminaiy investigation of the effects of a
variety of stains. A luimber of staining combinations were attempted,

214 UniversHij of California Puhlications in Botanii r^'*^"-"

especially in llie case of sections involving- cliloroi)liyllous tissue, to
stain the i)rot()|)lasmie matrix of tlic cell and follow with Sudan III.
It appears that a heavy staining willi Orange G. followed hy Sudan
III may he a useful eonihination in this connection. Of a number of
stains ordinai'ily employed for the diiferentiation of the elements of
woody tissue one at least gave some indication of being of interest.
Sections were placed in Delafield's haeniatoxylin for twelve hours,
partially destained and placed over night in Sudan III. In the result-
ing preparation the rubber inclusions wei-e stained scarlet by Sudan
III while in llie center of each was a single spot of dark purple indi-
cating, apparently, that each i-ubbei- globule contains a |ttotoplasmie
"nucleus'' around which it is built up. Results of this sort indicate
a field for further investigation.

It occurred to us, also, that vulcanization of the rubber in situ
might be possible, and thus giving specific differentiation under the
microscope to rubber inclusions in the cell. After suitable dehydra-
tion, sections were placed in the cold in sulfur chloride (in carbon
tetraeholoride) or were allowed to stand in the fumes of such a solu-
tion. The results of a number of preliminary experiments indicated
that vulcanization of the rubber inclusions in the cells w^as possible.

The following outline of the process of making preparations of rub-
ber-containing tissue described in some detail above may be of assist-
ance to those who may desire to make histological examinations for a
similar purpose. Attention must be called to tlie fact that nothing
original is claimed for this process, nor will it necessarily prove suc-
cessful in all its details for other species.

It is assumed that a woody plant is under investigation and that
sections have been cut from a sample of mature tissue.

(1) Sections from water to 95 per cent aleoliol : .") minutes.

(2) Boiling acetone; 15 to oO minutes.

(3) Sudan III; 18 hours.

(4) AVash off excess of stain in 50 i)er cent alcohol ; as rapidly as
])Ossible.

(5) [Mount in pure glycerine.

A preparation made in this manner from material cut from a rela-
tively higii percentage plant of Clirysotlunnnns nauseosus will show
cell inclusions of rubber stained a brilliant scarlet. At a magnification
of 150 diameters the parenchymatous elements of the cortex and es-
pecially the broad wedge-shaped cortical extensions of the i)rimary

1919] Hall-Goodspeed: Chnjsil 215

rays will appear to be solidly filled with staiuable material. In the
vouuffer corkv lavers of the inner bark stainable cell inclnsions will
probably be nnmerous. Nearer the periphery a tissue which is often
intensely stained will usually be found, with greater magnification, to
consist largely of much crushed bark cells the suberized walls of which
take up the stain readily. Within the cell the rubber occurs in globular
form or in a more or less diffused state filling the entire cell cavity.
The globules may be large, one or two to a cell, or small and numerous
in each cell (cf. plate 18, figs. 4 and 5). Only this brief description
will be made here of the appearance of a typical preparation made
according to the process detailed above. The subject of the occurrence
of rubber in the plant and in the cell is treated of in some detail
elsewhere (cf. p. 234).

As mentioned in an earlier paragraph, we have attempted with
some success to use the microscopical method for estimating the rela-
tive amounts of rubber in the various samples examined. A purely
arbitrary scale of values was adopted and relative rubber content de-
termined in each case by comparison with other preparations selected
as representing high, low, and medium rubber content. For such
comparisons a comparison-ocular was found to be useful. Tniformity
in the quality and quantity of the illumination is quite important
and artificial light was therefore used throughout. As soon as it
became possible to obtain large numbers of chemical analyses the
histological method of estimation became largely superfluous. How-
ever, in almost all cases it was resorted to prior to the chemical analysis
and a rough estimate made. Often when the rubber content of a sam-
ple appeared to be quite low no chemical analysis was made, and a
considerable amount of time and labor was thus saved. Throughout
the investigation we continued to examine microscopically numbers of
samples of doubtfully valuable species which were suspected by others
of containing rubber or which were collected by ourselves in order that
no possible source of rubber accessible to us should be overlooked.

We have found the microscopical method invaluable in many phases
of this investigation. The ([uestion as to the place and time of the ori-
gin of rubber in the plant can only be answered by employing this
method. Furthermore, supplemented by chemical analysis it gives
important evidence concerning the parts of the plant which carry rub-
ber and their richness relative to age and location. Ultimately the sea-
son and method of harvesting such a rubber crop will be determined
largely on the basis of information so derived.

21() I' iiii'i rsil 11 of Calif oniiii I'tihlicdl l(His in liotdtiii |\'ii,.

h. (IIK.MICAL ANALYSIS

Tlic (>ri<iiiiiil clKMiiical aiuilysis ot" CJirnsofluiin n us was nijulc in
Oetolx'i'. IfH)."), liy Profcssoi- (i. K. ('olhy of tlu' ( ';ilif()riii;i ^^xpci-iinciit
Station. Al llir st;ir1 of onr investigations in l!tl7 ;i few inuilyscs
were very kindly made lor us by i*i-of('SSor P. I>. llililiartl of llic sauH'
station. Tlic gvi-ni majoi-ity of tlie analyses listed in ilie various tables
wbieli follow were inad<' under ibe dii'i'dion of tlie junioi- autbor, in
wbose biboratoi-y Ibe iieeessary gi-inding and extraction apparatus was
set u|).

Tbe i)ai"liLMdai' iiietbod of ebeinieal aual\sis wbieb we ba\'e almost
oxeliisively em))loy(Hl consists in tbe tborouoii exti'aetion of a finely
ground sani))b' of nudei'ial. first wilb acetone and second witli benzene.
'I'be acetone e.xti'act is taken to contain approximately all resins, fats,
and sindlar bodies; tbe benzene exti'act to contain tbe I'ubbei'. A de-
tailed scbedule of tlic wliole proccss is giveu below for tbe convenience
and assistance of those wlio may care to make use of this method or
some improvement upon it. It is to be understood that the ])eriods of
extraction as well as the length of the periods of di-ying to constant
weight were definitely determined after a large numbei- of ])reliminai'y
efforts. The few early extractions were made with one of the numerous
modifications of the Soxhlet apparatus, while for tbe great majority
the Bailey-Walker extractor was employed. Tbe acetoiu^ extraction
was made over the water-bath, tbe benzene extraction on electi'ic hot
plates. The method may be sunnnarizi d as follows:

(1) Five-gi'am sample extracted .'! bours. boiling acetone.

(2) Acetone flask dried S hours, cooled in desiccator and weigln d.
(8) ^laterial in cxti-action tbiiuble or sipbon tube dried, placed in

a second flask, and subjected to action of boiling benzene
for '.-> bours.
(-1-) Flask containing benzene extract di'ied 4 bours. coob d in
desiccatoi". and weigbed.

Two layei's of filler papei- were placed at the liottom of tbe siphon
tube, as well as a plug of cotton. So complete a [)roteetion was this ar-
raiigement against the coming o\'er of partiides of tbe sample that fil-
tration of th(^ solution at the completion of the exti'act ion was iu)t nec-
essary. Similarly, a plug of cotton placed upon tbe top of tbe ma-
tei'ial in tln' sipbon tube obviated ;iii\- dangei' of an o\-ertlowing of

1919] llall-Ooodspced : Chrusil 217

the finely divided sample, especially as the bottom of tlie condenser
pressed down on this cotton plug during the course of the extraction.

In almost all cases the first of the acetone-soluble substances ex-
tracted were strongly colored and turned the solvent a dark greenish
brown as soon as the extraction commenced. Thereafter the liquid
passing through the siphon tube was practically colorless. The benzene
extract was in practically all cases entirely without color. The acetone
extract after drying was very dark brown in color and strongly
odorous. The benzene extract when dry was on the other hand
colored light brown or yellowish, was almost entirely odorless, and
was distinctly rubber-like in consistency.

There are, of course, a variety of other methods of analysis whicli
might possibly- have successfully replaced the one wliieli we have
employed throughout. For the sake of comparison we made trial of
another method, according to which the sample is first cxtraetrd
thoroughly with gasoline, then treated with strong sodium liydroxide,
then filtered, and the residue, finally, extracted with carbon tetra-
chloride. This method is far more time-consuming than llie one which
we have used and appeared to offer no real advantages as compared
with it. In the extraction of Guaj'ule shrul) on a large scale gasoline
is sometimes used as the solvent and acetone as a precipitant. The
recover^' of these reagents used in large (juantity in such a process
is a complicated problem l)ut it can be accomplished with very slight
losses. Moisture determinations were made in most cases.

Tlic following table indicates something as to the relative accuracy
of our chemical analyses. Duplicate or triplicate analyses were often
run ill cases where comparisons were to be made with analyses of other
parts of the same plant or with equivalent portions of a given indi-
vidual collected on various different dates. The figures given are
selected as being representative. They indicate a relative degree of
accuracy for our analyses, sufficient at least to give an estimate of
the average amount of rubber contained in tlu; various species witli
which we have been concerned.

218

Unlvcrsil If of Californiit PuhVications i)) Botnny T^'oi,. 7

T.\Bi,E 1. — The Kestlts ok a Numuek of T)rpr,TCATE Axat.yses

Spccirs or v:iriot.\'

157 Haplopapi)U.s ericoicliv

20.'? C. n. Kiiaphalodes*

2()(') Haplopappus nanus

220 C. n. consimilis

240 C. n. speciosus

255 C. n. speciosus

257 C. n. consimilis

290 C II. pinifolius

"■ 111 tliis and followiiio'
('hnjHotha m nun na uscosiis.

Place of Collection

San Francisco, Calif.
San Francisco, Calif.
Koarsargo, Inyo Co., Calif.
KearsarKf, Inyo Co., Calif.
Near Benton, Calif.
Near Benton, Calif.
Near Wabuska, Nev.
Near Wabuska, Nev.
Near Weed, Calif.
Near Weed, Calif.
East of H;immett, Idaho
East of Haminett, Idaho
East of Hammett, Idaho
East of Hammett, Irlaho
East of Hammett, Idaho
East of Hammett, Idaho
East of Hammett, Idaho
Shoshone Falls, Idaho
Shoshone Falls, Idaho
Alamosa, Colo.
Alamosa, Colo,
Alamosa, Colo.

tables tlic initial letters

Date of .Acptoru' Honzonp

Coljpction Kxtriict Extijirt

1918 Per cent. Per cent.

July 27

0.07

2.10

July 27

4.99

1.89

Jan. 28

2.81

1 45

Jan. 2S

2 50

1.40

Fob. 19

9.57

0.72

Feb. 19

10. n

0.88

Feb. 27

3.09

2 75

Feb. 27

3.()2

2.95

June t)

2.90

1.48

June 6

2.43

1.52

June 22

4.59

2.07

June 22

4 . 05

2.50

Jime 22

5.19

2.49

June 22

4 . 09

2.70

June 22

4 10

2 52

June 22

4.00

2.60

June 22

4.20

2.58

June 23

3.90

2.63

June 23

3.51

2.50

Aug. 31

4.27

3.41

Aug. 31

4.70

3.40

Aug. 31

4.23

3.59

C. u. are

used to

indicate

The reduction by maceration and grinding of woody plants for
detailed chemical analysis of the contents of their tissnes is at best
attended with some difficulty. The difficulties were somewhat mag-
nified in the material with which we worked, since the wood of most
species of ChrysothniiniHs is quite hard and since, also, it was necessary
in most cases to grind green shruh.

Each plant to be analyzed was trimmed with care. Since it seemed
clear tliat for all jiractical purposes the older ])ortions of the main
stem contained maximum (juantities of rubber, all the younger shoots
and most of the root was eliminated previous to grinding. Figure 4
shows portions of a representative jilant of Chrijsothiunnus )i(iiisrosus
var. viriduJus trimmed in this way and ready for grinding.

Usually such large plants were cut up into a number of e([uivaleiit
portions, alternate pieces being used for grinding. In s(mie cases
where the plants were unusually large the trimmed mass was split
longitudinally into portions as nearly equivalent as possible. In this
connection may be mentioned th(^ difficulty of {performing this opera-
tion successfully. It is evident that the including of "bark" belonging

1919]

Hall-Goodspecd : Chrysil

219

?'

■ '/

Vi^

1^

iftii

Jifi

*

*i«

Fig. 4. Portions of a representative plant of Chrysothamnus nauseosus var.
viridulus trimmed and ready for grinding.

'2-i) r iiir( rsil 1/ of CdlifaniKt I'lihllcal Ions in liotaiuj | \ <ii.. 7

1() one portion ;iloii<; witli till' |)oi'tioii to 1)(' yi'ouiid will decidedly
affect the nililiec content of the ^ronnd saiiiple since the j'lvater pai't
of the rnlilu'i' is contained in the e\ti-a-cainbial elements of the stem.
To test this niattei- \vc I'an a few analyses of ground matcTial from
both pofti(His of a nninbei- of |)lants. The results are o-iveii in the
following' tatth'.

Tab[,e 2. — Vakiation in Rubber Content Due to Unequal Separation, Shown

BY Analyses of Both Halves of Large 1'lants which had

BEEN Split Lonoitudinally

Date of Aeetone Benzene

Cullortion Extract Extract

Species Place of Collection 19IH Percent. Percent.

4;") r. n. consimilis Adobe Valley, Calif. Sept. 19 ;i.70 2.55

Adobe Valley, Calif. Sept. 19 :j.75 2.44*

Adobe Valley, Calif. Sept. 19 :j.21 2.81

457 (^. n. consimilis Gaspipe, Calif. Sept. 19 ;i.66 1.95

Gaspipe, Calif. Sept. 19 2.97 2.07

lt)7 C. u. viridulus North of Mono Lake, Calif. Sept. 22 3.71 1.74

North of Mono Lake, Calif. Sept. 22 2 . 2S 2 . 47
*■ r)u])li<'ati' analysis.

These figures imiicate that care must be exercised in splitting off
jxtitions of a plant for grinding- and analysis. As noted above it has
been almost uniformly our practice to cut u]) larger plants into trans-
verse sections and grind alternate pieces.

For the first rough crusliing or grinding a small power di-iven
feed mill ])roved satisfactory. Tbc material was run thi'ongh this
inill a uiimbei' of times, the grinding phites being continually brought
neai'cr together until the maximum degree of fineness was attained.
At this stage practically all tlu> material could be passed through a
lO-inesh sieve. For the final grinding a hand-operated No. 0 Enter-
l)rise coffee mill pi'oved successful. The final i-esult of this last
grinding was the |)r()(luction of material that would pass through a
24-mesh sieve.

We have attem|)ted with some success to substitute the action of
a pebbh' mill for the greater part of tlu' final grinding.^' Since the
rubber is held within the walls of tlie individual cells and since, even
when put into solution by benzene, these confining walls must serve
to binder outward diffusion of the rubbei- to some extent, it seemed

1" A niodifieatioii of a small Alihe pel)l)l(' mill (''siiij^lc sjKH-inun mill'') was
used. Mention mijiflit liere ho made of the fact that there are on the market a
nund>er of ])ower driven grinding machines which wonlil uiidouljtedlv ])erfonn an
iidtial reduction to 10-mosh nmcli more rapidly and i-veidy than the apparatus
descrilii'il almve.

1919]

Hall-Goodspcrd : Ch rysil

221

clear that previous to extraction the finest possible subdivision of the
plant material was desirable. Preliminary tests indicated that after
prolonged grinding in the pebble mill a greater reduction in size of
particles was attained tliau by our usual final grinding in the coffee
mill. Examination of tlie pebbles at the end of their action showed
particles of rubber as well as woody and fibrous tissue adhering to
their surfaces. We therefore made a number of analyses to determine,
if possible, just what proportion of the rubber was left behind on the
pebbles after pebble mill grinding. The following table indicates the
relation in each case between the amount of rubber extracted from a
sample of the ground material as it came out of the pebble mill and
the amount extracted from another portion of the same rough ground
material which had been reduced in the coffee mill. Attention should
be called to the fact that no rubber was found to adhere to the
metal grinding plates of the feed mill or coffee mill and that both
were taken apart and l)ruslird out after each grinding, the material
remaining in the mills being added to the ground substance which
had gone through them. Tn a few cases material was ground in a
pebble mill operated in the Food and Drug Laboratory of the Uni-
versity of California. This mill was larger than ours and the product
was somewhat more finely divided than the product of our pebble
mill.

Table 3. — Influence Upon the Results of Axalysls of Final Grinding in a

Coffee Mill and in a Pebble Mill

Species

Acetone
Extract
Per pent.

Benzene
Extract
Per cent.

R(

?iiiarks

S7

C.

n.

gnaphalodes

2.78

0.59

Coffee mill.

3.07

0.46

Pebble mill.

4 hours.

133

C.

n.

viridulus

4.12

3.64

Coffee mill.

4.60

4.06

Pebble mill.

4 hours.

134

(\

n.

viridiilus

3.73

2. IS

Coffee mill.

3.93

2 14

Pebble mill.

4 hours.

135

C.

n.

viridulus

4.o7

1.30

Coffee mill.

3.69

1.29

Pebble mUl.

6 hours.

4.07

1.22

Pebble mill (Food & Drug Lai).)

140

C.

n.

c'onsimilis

4.76

3.86

Coffee mill.

5.50

3.89

Pebble mill.

4 hours.

144

C.

n.

consimilis

3.75

0 81

Coffee mill.

3.74

0.79

Pebble mill.

4 hours.

146

C.

n.

consiinilis

3.69

1.74

Coffee mill.

3.54

1 . 54

Pebble mill.

4 hours.

222

University of California PuhIicatio)is in Tioianij [Vol. 7

T.\BLE :i. — (Continued)

Species
149 Haplopappus cricoides

215 C. n. gnaphalodes

227 C. n. consimilis

229 C. n. speciosus

.Acetone
Extract
VcT cent.

9.17

7.88

7.47
5.53

4.29
4.29

98
30

3 . 55
4.83
4.72

Henzene
Extract
Per cent.

3.92

2.48

3 11
2 . 50

2.57
1.92

2.14
2 12

2.49
2.37
2,07

Remarks

C"offoo mill.
Pohhlf mill.

Coffee mill.
Pebble mill.

Coffee mill.
Pebble mill.

Coffee mill.
Pebble mill.

Coffee mill.
Pebble mill.

4 hours.

4 hours.

4 hours.

4 hours.

G hours.

Pebble mill (Food & Drug Lab.)

Although, as in all such cases, a far larger number of analyses is
necessary to give entirely reliable evidence it appears from the figures
given above that the product of the pebble mill, irrespective of the
fact that some rubber is kept behind on the pebbles, yields approxi-
mately as large an amount of rubber as does the product of the coffee
mill. The results given in the following table leave no room for doubt
that rubber does adhere to the pebbles.

Table 4. — Evidence that Eubber Adheres to the Pebble.s avhen Material is

Ground in a Pebble Mill

Species

206 Haplopappus nanus

Acetone
Extract
Per cent.

Benzene
Extract
Per cent.

Remarks

9.57

6.72

Coffee mill.

8.71

6.22

Pebble mill. 4 hours.

8.88

8.51

Pebble mill. 4 hours. Pebbles
washed in benzene and solu-
tion filtered. Residue (2gm.)
added to ground material
(3gm.) and whole extracted
with filtrate.

Tn view of these results the conclusion seems inevitable that the
degree of fineness of the ground material is an important factor in
the extraction of rubber from non-latex rubber-bearing plants accord-
ing to our method of chemical analysis. Apparently the added
fineness of the product of the pebble mill facilitates the thorough
extraction of the rubber to such an extent that the amount of rubber
added by this more nearh^ complete extraction compensates for the
amount retained upon the pebbles. In other words if over 2 per cent
of ni])])er is lidd on the pebbles, as is indicated by the results given
in table 4. one might expect that the benzene extract from pebble mill
mati'i-iiil would lie approxinuitcly 2 per cent less than that from coifee

'■*!■• I Hall-Goodspevd : ( 'h rijsil 223

mill material. In fact, however, these two analyses were in most cases
approximately the same and this result is assumed to depend upon
the fact tluit the finer division of the product of the pebble mill allows
tlie extraction of 2 per cent more rubber than was possible in the
case of the coarser product of the coffee mill.

With these facts in miiul we decided to make the final grinding
of all our samples in tlie coffee mill since the matter of removing the
rubber adhering to the pebbles of the pebble mill is somewhat difficult
and time-consuming. This was done as a matter of practicability not-
withstanding the knowledge that the results would be too low to
express the actual percentages present. The discrepancy, liowev(M',
cannot be anywhere near the 2 per cent mentioned since tliat referred
to a plant in which the rubber content is much higher than the average,
and one moreover in which the very resinous material adhered to the
pebbles more persistenth' than in most cases. It is believed that the
amount of rubber remaining in samples of Chrysothamnus nauseosus
after grinding in a coffee mill and extracting by our method seldom
if ever api^roaches 1 per cent of the original sample.

In a preliminary way, at least, we have obtained evidence as to the
effect of storage of the ground material upon rubber content. At
the start of our investigations it was assumed that if for any reason a
plant was not analyzed almost immediately after collection a deteriora-
tion in content and quality of the contained rubber would soon render
the result of its anah'sis of doubtful value. Undoubtedly the exposure
of shrub to drj'ing and weathering out of doors will bring about such
deterioration in the course of time (cf. Lloj'd, I.e., p. 10). As has
been noted above (p. 220) it is difficult so to divide a plant for analysis
that the halves or apparently equivalent portions will give closely
corresponding analyses. For this reason we did not attempt, in
seeking evidence as to the influence of storage upon rubber content,
to analyze a portion of a plant and after subsequent storage for some
months analyze an apparently e(iuivalent portion represented by the
remainder of the same plant. Kather the residue, after the first
analysis, of the ground material of a plant was stored in a stoppered
bottle and analyzed after a time. We have assumed that the amount
of deterioration in sucli a finely ground sample after storage approxi-
mates that which might be expected to occur within tlie tissues of
an entire plant after a corresponding period of time. The table whicli
follows details the results of a number of analyses of ground material
shortly after the original dates of collection and after periods of
storage varying from five to ten months.

094.

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l»l»] Hall-Goodspeed : Chrysil 225

It seems clear from the above figures that there is little to be feared
in the way of loss of rubber by the storage for some months of the
ground material of such rubber bearing plants as those whicli we
have investigated. It is assumed that a similar statement can be made
for plants in their original condition when stored in the laboratory
or mill under more or less uniform conditions of temperature and
moisture.

Attention miglit be called to the fact that in table 5 it appears
that the content of acetone-soluble substances in general shows a some-
what greater diminution in amount following storage than does the
rubber content. Further experiments along this general line are
planned since evidence may, seemingly, be obtained regarding the
relation between resins as well as other acetone soluble substances
and rubber. Indeed other analyses of stored material, not so well
controlled and therefore not included in table 5, indicate that accom-
panying the loss of resins, etc., there is an actual increase in rubber
after storage.

It had been our original intention to include in the various tables
which follow the date of analysis as well as the date of collection.
Since the period elapsing between these two dates seems to have little
or no influence upon the rubber content of a plant the former date
was eliminated. It should be said, however, that every effort was
made to analyze the various plants as soon as possible after they were
brought into the laboratory. In the majority of cases less than a
month or six weeks intervened between collection and analysis.

226 I' iih'i rsihj of ('(illfoni id I'lihl ic<it itnis^ in /ioftnii/ [\<)i.. *

IX. HEsn/rs OF THE CHHMICAL analyses and MK'RO-

SCOrU'AL EXAMINATIONS; TAP.FLATION OK PER-

CENTACJE OF Hl'RRER IN EACH VARIETY

The methods eiui)h)y('(l in inakiii<i- Ihe chcniical analyses and
inieroseoi)ieaI exaininations iiave Ix'cn detailed elsewhere (cf. p. 1210).
The following tables give the residts of sueh analyses and examina-
tions as wei'i' made on individnal plants of a nnndici" of varieties of
C. nauscusus. As in pfcvions tables the eollectioji number is iiiehuled
for the sake of eompleteiiess and foi" future referenee. Tlic date of
analysis is not given because of eonsiderations uicntioiicd al)ov('. The
"X" following certain of the jiercentages indicates tliat tiiey ai'e not
based upon dry weight. In all other eases moisture determinations
were made, and the figures are based upon dry weight.

The terms used in describing the amount of rubber determined by
microscopical examination are only relative and are derived from a
more elaborate classification used during tlie course of the investi-
gations.

In the reports on the chemical analyses, the first column, or
"acetone extract" is taken to represent the percentage of resins, fats,
waxes, etc., in the sample while the second column indicates the per
centage of pure rubber, or C-hrysil. As indicated on page 223, the
figures in this second column are probably too low by a fraction of
1 per cent in each case owing to the difficulty of obtaining the proper
degree of fineness in grinding the samples.

A summary of these tables is given on page 244.

We are under great obligation to Miss Mildred Crane who under-
took the carrying through of the majority of the chemical anah'ses
and assisted during the entire investigation in a variety of ways.

Table (i. — Resi'lts of Chemical Analyses and Microscopical Kxamixations
OK Jndiviihai- Plants of Twelve Varieties of Clirysofltomnu.'i lunisrosu.s

Chrysothamnus nauseosus (schku sfrictu)

Microscopical Examinations

Place of Collection

251 Xeai' Harney, Ore.
46.3 Mono Lake, Calif.

Dato of

Estiiiiatci

Colliction

AlllDUIlt

.lime lit, ims

Xothinj

Sept. 20. 19 IS

Traces

1^1^] Hall-Goodspred: Chrysil 227

Chrysothamnus nauseosus var. consimiUs
Che.mk AL Analyses

v,^ n 1'.^ .*l n .1 Li .

Place of Collection.

V x-S j"V 1 J I O III .^

Date of
Collection

.\cetone
Extract
Per cent.

Benzene
Extract
Per cent.

16

Deeth, Nev.

June 17, 1917

3.10

2.30

139

Golconda, Nev.

Mar. 4, 1918

4 72

2.21

140

Golconda, Nev.

Mar. 4, 1918

4.76

3.86

141

Golconda, Nev.

Mar. 4, 1918

5.12

3.71

142

Golconda, Nev.

Mar. 4, 1918

4.70

2.27

143

Golconda, Nev.

Mar. 4, 1918

3.85

1.07

144

Golconda, Nev.

Mar. 4, 1918

3.75

0.81

145

Golconda, Nev.

Mar. 4, 1918

3.59

2.12

146

Golconda, Nev.

Mar. 4, 1918

3.69

1.74

208

Near Blair Junction, Nev.

Feb. 24, 1918

5.58

2.90

210

Near Miller.s, Nev.

Feb. 24, 1918

3.67

4.17

214

Schurz, Nev.

Feb. 26, 1918

4.32

0.99

216

Schurz, Nev.

Feb. 26, 1918

3.62

0.65

217

Walker River, Nev.

Feb. 27, 1918

4.77

1.27

218

Walker River. Nev.

Feb. 27, 1918

3.79

2.17

219

Walker River, Nev.

Feb. 27, 1918

5.06

1.03

220

Near Wabuska, Nev.

Feb. 27, 1918

3.69

2.75

227

Karlo, Modoc Co., Calif.

Mar. 7, 1918

3.98

2.14

231

Karlo, Modoc Co., Calif.

Mar. 8, 1918

4.26

2.23

232

Honey Lake Valley, Calif.

Mar. 9, 1918

5.17

2.43

241

Near Weed, Calif.

June 6, 1918

3.25

1.02

250

Near Burns, Ore.

June 19, 1918

2.55

0.89

253

Near Ontario, Ore.

June 20, 1918

3.57

3 . 14

254

Southeast of Mt. Home, Idaho

June 20, 1918

2.50

0.46^

257

Shoshone Falls, Idaho

June 23, 1918

3.90

2.63

402

Stockton Lake, Utah

Sept. 3, 1918

2.28

2.46

403

Stockton Lake, Utah

Sept. 3, 1918

3.14

1.31

404

Stockton, Utah

Sept. 3, 1918

3.91

2.84

408

Milford, Utah

Sept. 4, 1918

4.64

1.25

430

Goldfield, Nev.

Sept. 9, 1918

1.65

1.69

455

Adobe Valley, Calif.

Sept. 19, 1918

3.70

2.55

456

Adobe Valley, Calif.

Sept. 19, 1918

4.77

2.84

457

Gaspipe, Calif.

Sept. 10, 1918

3.66

1.95

472

Near Mountain House, Nev.

Sept. 22, 1918

3.44

1.26

473

Carson, Nev.

Sept. 23, 1918

2.08

2.85

474

Carson, Nev.

Sept. 23, 1918

2.70

2.30

475

North side of Carson River, Nev.

Sept. 23, 1918

4.62

2.42

476

South side of Carson River, Nev.

Sept. 23, 1918

3.14

0.59

477

Southeast of Gardnerville, Nev.

Sept. 23, 1918

3.75

0.83

478

South of Carson, Nev.

Sept. 23, 1918

2.82

3.54

479

East of Carson, Nev.

Sept. 23, 1918

3.12

3.79

480

Near Dayton, Nev.

Sept. 23, 191S

3 .92

1.97

507

Stockton Lake, Utah

Sept. 25, 1918

1.74

0.20

514

Cedar Valley, Utah

Oct. 12, 1918

2.43

0.15

519

Muskrat Springs, I'tah

Oct. 24, 1918

2.73

0.48

520

Near Josepha, Utah

Oct. 24, 1918

3.16

1 23

522

Tooele Valley, Utah

Oct. 24, 1918

3.47

0.62

523

Vernon, Utah

Oct. 24, 1918

2.64

0.67^

524

Utah Lake, Utah

Oct. 30, 1918

2.69

1.63

527

Sanpete Valley, Utah

Nov. 12, 1918

3.00

1.25^

228

(' iiir< rsil 11 of (^(tliforiiiii riil>llc(it

Id IIS III

llitiiiinj [Vol. 7

Chemical Analyses

(CJo.ntinukd)

A Oft one

Henzene

Date of

lOxtract

Extract

Plaoc of Colli'ctioii

Colicftioii

Per cent. 1

Per cent .

527a

Sanpete Valley, Utah

Nov. 12, 191S

2.46

0 . 39^

528

Redmond, Utah

Nov. 12, 1918

3.10

3 . 48^

528ii

HedmoiKJ, Utah

Nov. 12, 191,s

2 33

1 77^

530

Circle City, ITtah

Nov. 12, 1918

3.04

1.26^

532

Panguitch, Utah

Nov. 12, 1918

2.90

2 OO'^

533

Mona, Utah

Nov. 12, 1918

2 . 52

0 . 82>^

534

C.oshon, Utali

Nov. 18, 191S

2 80

0.81'^

(500

Colconda, Nev.

Dec. 23, 1918

3.30

1 . 51

602

Watts, Nev.

Dec. 24, 1918

3.27

1.58

603

lieese Jliver Valley, Nev.

Dec. 25, 1918

3.72

1.07

604

Battle Mountain, Nev.

Dec. 28, 1918

3.41

2,98

605

Near Ely, Nev.

Dec. 30, 1918

2.98

0.54

606

Near Ely, Nev.

Dec. 30, 1918

3.04

1.84

607

Near McCill, Nev.

Dec. 31, 1918

3.80

1.15

608

Near McCill, Nev.

Dec. 31, 1918

3.04

1.04

609

Shaft or, Nev.

Jan. 1, 1919

5.06

3.04

610

Shaftcr, Nev.

Jan. L 1919

4.15

1.21

611

Lamoille, Nev.

Jan. 2, 1919

3 . 13

1.08

613

Near Elko, Nev.

Jan. 2, 1919

3.06

0.43

614

North fork of Humboldt River, Nev.

Jan. 3, 1919

3 . 42

1.07

615

North fork of Humboldt River, Nev.

Jan. 3. 1919

3.36

0.93

650

Near Macdoel, Siskiyou Co., Calif.

Dec. 25, 1918

3.35

1.57

651

Blitzen Valley, Ore.

Dec. 27, 1918

2.91

0.78

652

Catlow Valley, Ore.

Dec. 29, 1918

3.44

2.06

653

Warner Valley, Ore.

Dec. 29, 1918

4.00

1.68

654

Warner Valley, Ore.

Dec. 29, 1918

3 25

0.84

655

Warner Valley, Ore.

Dec. 29, 1918

2 . 95

1.43

656

(^uinn River Valley, Nev.

Jan. 3, 1919

4.79

3.36

657

Quiim River Valley, Nev.

Jan. 3, 1919

3 . 64

2,83

659

Near Gerlach, Nev.

Jan. 3, 1919

4.23

4.71

660

Near Gerlach, Nev.

Jan. 3, 1919

3.08

6.57

Microscopical Examination's

Date of

Estimated

Place of Collection

Collection

Amount

11

Pyramid Lake Road, Nev.

June 14, 1917

Fair

26

Coldfield, \ev.

Aug. 10, 1917

Fair

32

Pyramid Lake Road, Nev.

Aug. 16, 1917

Fair

159

Near Magdalena, New Mex.

July 22, 1918

Poor

233

Honey Lake Valley, Calif.

Mar. 9, 1918

Fair

252

Near Westfall, Ore.

June 20, 1918

Traces

25S

Southeast of Burley, Idaho

June 24, 1918

Traces

260

Southeast of Albion, Idaho

June 24, 1918

Traces

261

Southeast of Snowville, Utah

June 25, 191 >

Poor

263

Near Duchesne, Utah

June 30, 1918

Traces

417

Caliente, Nev.

Sept. 6, 1918

Traces

502

Grantsville, Utah

Sept. 25, 1918

Traces

505

Stockton Lake, Utah

Sept. 25, 1918

Nothing

506

Stockton Lake, Utah

Sept. 25, 1918

Traces

513

Cedar Valley, Utah

Oct. 12, 1918

Poor

521

Near Josepha, Utah

Oct. 24, 1918

Poor

525

Near Jericho, Utah

Oct. 30, 1918

Traces

526

Dcjremus, Utah

Oct. 30, 1918

Poor

1910]

Hall-Goodspeed : Ch rysil

229

Chrysothamnus nauseosus var. frigidus
Chemical Analysis

Place of Collection

271 Laramie, Wyoming

Date of
Collection

Acetone lienzene
Extract Extract
Per cent. Per cent.

July 7, 191S 16.60 1.S6

Microscopical ICxamination

Place of Collection

262 Near Rock Hill, Utah

Date of
Collection

Estimated
Amount

June 30, 1918 Traces

Chrysothamnus nauseosus var. gnaphalodes
Chemical Analyses

Place of Collection

48 West of Lancaster, Calif.

72 Benton, Calif.

87 West of Lancaster, Calif.

203 Kearsarge, Inyo Co., Calif.

215 Schurz, Nev.

447 Benton, Calif.

448 Benton, Calif.

449 Benton, Calif.

561 Basalt, Nev.

562 Basalt, Nev.

577 Near Barstow, Calif.

578 Near Barstow, Calif.

658 Quinn River Valley, Nev.

711 Near Victorville, Calif.

712 Near Victorville, Calif.

778 West of Lancaster, Calif.

779 West of Lancaster, Calif.

780 West of Lancaster, Calif.

Date (
Collect:

if

ion

Acetone
Extract
ppr cent.

Benzene
Extract
Per cent.

Oct. 27,

1917

3.10

1.90

Nov. 5,

1917

4.10

3.60'^'

Jan. 19,

1918

2.78

0.59

Jan. 28,

191S

2.81

1.45

Feb. 26,

1918

4.29

2.57

Sept. 16,

1918

3.66

2.66

Sept. 16,

1918

3 . 55

3.58

Sept. 16,

1918

2.39

1 . 85

Dec. 21,

1918

3 67

0.94^

Dec. 21,

1918

2,88

0.95^

Dec. 24,

1918

2.12

0.26^

Dec. 24,

1918

2.71

0.88

Jan. 1,

1918

3.30

1.88

Dec. 26,

1918

2.27

0.96

Dec. 26,

1918

2.96

1.80

Mar. 13,

1919

2.72

1.23

Mar. 13,

1919

2.30

1.20

Mar. 13,

1919

2.40

0.76

Microscopical Examinations

Place of Collection

13 Pyramid Lake Road, Nev.

35 Pyramid Lake, Nev.

41 Mojave, Calif.

42 Mojave, Calif.
49 Lancaster, Calif.

51 West of Owens Lake, Calif.

53 Kearsarge, Inyo Co., Calif.

59 Independence, Calif.

63 West of Bishop, Calif.

481 West of Mound House, Nev.

482 Pyramid Lake, Nev.

Date of

Estimated

Collection

Amount

June 14, 1917

Fair

Aug. 16, 1917

Good

Oct. 25. 1917

Traces

Oct. 25, 1917

Poor

Oct. 28, 1917

Poor

Oct. 29, 1917

Traces

Oct. 29, 1917

Good

Oct. 31, 1917

Good

Nov. 1, 1917

Poor

Sept. 23, 1918

Fair

Sept. 24, 1918

Nothing

2:50

riiir( rsilfi of Cdliforiiid riihlicdtioHH in liotdiiii

Vol. 7

Chrysothamnus nauseosus \;u. graveolens

ClIKMICAI. AnALVSKS

Place of Cdllrction

272 Ft. Collins, Colo.

278 Trinidad, Colo.

279 Trinidad, Colo.

250 Trinidad, Colo.

251 Trinidad, Colo.

288 Pnrfiatoir River, C'olo.

29G Cimarron, Colo.

299 Near Grand .Imiction, Colo.

411 Near Cedar City, I'tali

516 C.rcen l{ivcr. I'tah

Date of
Collection

July 7, 1918

Aufi. 11, 1918

Auii. 11, 1918

Aus. 8, 19 IS

Ausr. 11, 191,S

Aim;, l.i, 1918

Sept. 1, 1918

Sept. 1, 191S

Sept. .■), 1918

Oct. 18, 1918

.Acetone
Extract
Per cent.

2 . 84

2.39

2 . f).-.

:i.02

2.88

:m:!

3.1.-)

2.0:;

2.0 5

2 . 88

Benzene

Extract

Per cent.

0 t)7^'

0.07

1.03

0.68

0 . 69

0.36

0.21

3 19

0 . 49

0 . 87

MlCROSCOI'ICAL Ex.\MINATIONS

Place of Collection

266 Vernal, l^tali

288 Canon City, Colo.

297 Grand Junction, Colo.

298 Grand Junction, Colo.

401 Near Grand Junction, Colo.

412 Near Cedar City, Utah

497 Garden of the Gods, Colo.

Date of
Collection

July 1, 1918

Aug. 6, 1918

Rei)t. 1, 1918

Sept. 1, 1918

Sept. 2, 1918

Sept. o, 1918

July — 1918

Estimated
Amount

Trace.s

Trace.s

Traces

Traces

Traces

Poor

Poor

Chrysothamnus nauseosus var. hololeucus
Chemical .\\ai,ysks

Place of Collection

62 West of Bishop, Calii.

451 Benton, Calif.

452 Benton, Calif.

453 Benton, Calif.

Date
Collect

of
ion

.Acetone

Extract

Per cent.

Benzene

Ext 1 act

Per cent.

Nov. 1,

1917

4.00

4.10

Sept. 16,

1918

2.87

1.03

Sept. 16,

1918

2.98

3.98

S(>pt. 16,

1918

3.14

2 22

Mu'HOSroi'K'.VL FiXA.\II\.\TIOXS

Place of Collection

12 Pyramid Lake Road, Nev.

3 1 Pyramid Lake, Nev.

58 Independence, Calif.

65 Benton, Calif.

91 Near Palmdale. Calif.

483 Pvramid Lake, Nev.

Date
Collect

of
ion

Estiniatet
Amount

June 14,

1917

Traces

.\n<i. 17,

1917

Fair

Oct. 31,

1917

Poor

Nov. 5,

1917

Poor

Jan. 19,

1918

Poor

Sept. 24,

1918

Fair

1919]

Hall-Goodsprcd : ( li rysil

231

Chrysothamnus nauseosus var. leiospermus
Che.viical Analyses

Place of Collection
418 Near Caliente, Nev.
529 Joseph City, Utah

Place of Collection

78 Near Candelaria, Nev.
415 East of Caliente, Nev.

Date of
Collection

Acetone Benzene

Extract Extract

Per cent. Per cent.

Sept. 6, 1918

2.82 0.84

Nov. 12, 1918

2.41 1.17^

NATIONS

Date of
Collection

Estimated
Amount

Nov. 8. 1917

Traces

Sept. 6, 1918 Traces

Chrysothamnus nauseosus \ ar. mohavensis
Chemical Analyses

Place of Collection

550 Mt. Hamilton, Calif.

551 Mt. Hamilton, Calif.

552 Mt. Hamilton, Calif.

553 Mt. Hamilton, Calif.

Place of Collection

40 Mojave, Calif.

43 Southwest of Mojave, Calif.

44 Southwest of Mojave, Calif.
92 Near Palmdale, Calif.

Date of
Collection

.Acetone
Extract
Per cent.

Benzene

Extract

Per cent.

Dec. 15, 1918

2.05

0.30^

Dec. 15, 1918

2.90

0.30^

Dec. 15, 1918

2.79

1.08^

Dec. 15, 1918

2.15

0.44^

nations

Date of
Collection

Estimated
.\niount

Oct. 25, 1917

Nothin:

tT

Oct. 25, 1917

Poor

Oct. 25, 1917

Traces

Jan. 19, 1918

Poor

Chrysothamnus nauseosus var. occidentalis
Chemical Analyses

Place of Collection

775 Near Tejon Pa.ss, Calif.

776 Near Tejon Pass, Calif.

777 Tejon Pass, Calif.

Date of
Collection

Mar. 13, 1919
Mar. 13, 1919
Mar. 13, 1919

.\cetone
Extract
Per cent.

2.78
2.84

1.88

Benzene
Extract
Per cent.

0 95
0.71

1 54

290
291
293
294

788

Chrysothamnus nauseosus var. pinifolius
Chemical Analyses

Place of Collection
Alamosa, Colo.
Alamosa, Colo.
Salida, Colo.
Salida, Colo.

San Luis Valley, Colo.

MiCUO.SCOI'ICAL EXA.MINATIOX

Place of Collection

265 East of Roosevelt, Colo.

Date of
Collection

.Acetone
lOxtraet
Per cent.

Benzene
Extract
Per cent.

Aug. 31, 1918

4,27

3.41

Aug. 31, 1918

4.81

1.11

Aug. 3, 1918

4.05

3.98

Aug. 31, 1918

4.00

2.65^

Mar. 15, 1919

3.62

3 . 59

NATION

Date of
Collection

F]stimate(
.\ mount

1

June 30, 1918

Traces

•2:i-2

U)iivcrsit !l of Califonild I'lihlicdf iaiis In liotmni 1^

(II,. (

229
240
24")
248
255
270
40')
406
407
402
405
471
495
498
557
559
758

Chrysothamnus naiiseosus \;ii. speciosus

CmCMK .\I. .V\" ALYSICS

I'liiro iif C'olloction

South of Likely, Modoc Co., Calif.
X(>:ir \^'oed, Calif.
Antelope, Ore.
Near Burns, Ore.

It,

bho

East of Haninie
Rawlins, Wyo.
Stockton, Colo.
Stockton, Colo.
Stockton, Colo.
Warren Creek, Calif.
Mono Lake, Calif.
Xear State Lino Lake, Nev.
lledmond. Ore.
Lolo, Mont.
Keno, Nev.
Near Reno, Nev.
Spokane, Wash.

Dat<> .

r.f
ion

.\((^tono

I'.xtniot

I'cr ociit.

BrIIZCIK'

Extiart
Per cent.

Mar. S,

191S

8.55

2.49

June 6,

1918

2 48

1 . 52

June 16,

1918

8 . 88

2.77

June 19,

1918

8 . 10

0.40

June 22,

1918

4.16

2.52

July 8,

1918

8.70

1 . 80^'

Sept. 8,

1918

2.87

0.52

Sept. 8,

1918

2.39

0.44

Sept. 8,

1918

8.09

0.38

Sept. 20,

1918

8.98

0.58

Sept. 20,

1918

8.48

1 . 55

Sept. 22,

1918

8 . 87

0.49

Sept. 29,

1918

4.02

0.78

Dec. 4,

1918

2.75

0.77^^

Dec. 20.

1918

8.43

2.81'^

Dec. 20.

1918

2.81

0 . i\'l^

Feb. IS,

1919

8 . 49

OK)

Microscopical Examixations

Place of Collection

6 Truckee, Calif.

10 Boca, Calif.

17 Ogden, Utah

20 Ogden, Utah

21 Ogden, Utah
25 Coldfield, Nev.
27 Reno, Nev.

87 Truckee, Calif.

89 Reno, Nev.

150 Redmond, Ore.

161 Albuquerque, New Mc.x.

162 Mofloc Co., Calif.
168 Modoc Co., Calif.
164 Modoc Co., Calif.

244 South of Roseburg, Ore.

246 Near Antelope, Ore.

249 Near Burns, Ore.

264 Near Duchesne, Utah

269 Southwest of Rawlins, Wj'o.

435 Lida, Nev.

464 Mono Lake, Calif.

488 Reno, Nev.

501 Cirantsville, Utah

503 Grantsville, Utah

511 Near Salt Lake, IJt^ih

512 N{!ar Salt Lake, Utah

Date

of

Estimated

Collect

ion

Amount

June 6,

1917

Traces

June 18,

1917

Nothing

June 19,

1917

Traces

Aug. 7,

1917

Traces

Aug. 7,

1917

Traces

Aug. 10,

1917

Poor

Aug. 14,

1917

Poor

Aug. 17,

1917

Nothing.

Aug. 14,

1917

Poor

May — ,

1918

Poor

July 29,

1918

Poor

Nov. — ,

1918

Poor

Nov. — ,

1918

Fair

Nov. — ,

1918

Fair

June 8,

1918

Nothing

June 16,

1918

Traces

June 19,

1918

Nothing

June 30,

1918

Traces

July 7,

1918

Traces

Sept. 10,

1918

Traces

Sept. 20,

1918

Nothing

Sept. 25,

1918

Poor

Sept. 25,

1918

Poor

Sept. 25,

1918

Poor

Oct. 7,

1918

Poor

Oct. 7,

1918

Poor

Utl91

Hall-Goodspeed : Ch rysil

233

Chrysothamnus nauseosus var. vlridulus
Chemical Analyses

54
64
80
133
134
211
212
428
434
439
441
442
443
444
445
446
450
459
467
468
563
565
570
571
572
574
576
590
704
705
706
718
748
749
752
755

47
50
55
56
57
60
79
84
88

Place of Collection
Koarsargo, Inyo Co., Calif.
Benton, Calif.
Near Benton, Calif.
Benton, Calif.
Near Benton, Calif.
Fish Lake Valley, Nev.
Near Oasis, Nev.
Beatty, Nev.
Lida, Nev.

Near Deep Spring Valley, Calif.
Benton, Calif.
Benton, Calif.
Benton, Calif.
Benton, Calif.
Benton, Calif.
Benton, Calif.
Benton, Calif.
Near Mono Lake, Calif.
Near Mono Lake, Calif.
Near Mono Lake, Calif.
Near Benton, Calif.
Benton, Calif.
Benton, Calif.
Near Benton. Calif.
Benton, Calif.
Near Benton, Calif.
Near Benton, Calif.
Near Victorville, Calif.
Cushenberry Spring, Calif.
Cushenberry Spring, Calif.
Cushenberry Spring, Calif.
Hesperia, Calif.
Near Barstow, Calif.
Near Benton, Calif.
Mono Lake, Calif.
Mono Lake, Calif.

Microscopical Examin.\tions

Place of Collection
Near Rosamond. Calif.
West of Owens Lake, Calif.
Kearsarge, Inyo Co., Calif.
Kearsarge, Inyo Co., Calif.
Kearsarge, Inyo Co., Calif.
West of Bishop, Calif.
Near Mina, Nev.
Laws, Inyo Co., Calif.
West of Lancaster, Calif.

Date
Collecf

of
ion

.\cetone

Extract

Per cent.

Benzene

Extract

Per cent.

Oct. 29

1917

3.90

4.84^

Nov. 3

1917

3.74

5.56^^

Nov. 10

, 1917

3.56

2.00

Feb. 15

1918

4.12

3.64^^

Feb. 17

1918

3 . 73

2.18

Feb. 25

1918

5.65

2.00

Feb. 25

1918

5.02

0.96

Sept. 8

1918

2.40

2 . 70

Sept. 10

1918

2.57

1.67

Sept. 12

1918

2 22

2.39

Sept. 16

1918

2.80

2 31

Sept. 16

1918

3 . 18

3.97

Sept. 16

1918

2,52

3.11

Sept. 16

1918

2.60

3 . 83

Sept. 16

1918

3.13

5,29

Sept. 16

1918

3.61

4.16

Sept. 16

1918

2.50

2.09

Sept. 19

1918

3.64

1.79

Sept. 22

1918

3.71

1.74^^

Sept. 22

1918

2.88

0,65

Dec. 21

1918

3.38

0,52

Dec. 21

1918

2.64

0,58

Dec. 21

1918

2.92

1.09^

Dec. 21

1918

3.49

4,45^^

Dec. 21

1918

3.04

4.00^

Dec. 23

1918

2.61

2.00^

Dec. 23

1918

6.85

3.93^^'

Dec. 25

1918

2.70

2.43

Dec. 25

1918

3.21

1.02

Dec. 25

1918

2.13

1.30

Dec. 25

1918

2.08

1.34

Dec. 26

1918

1.94

0,44

Dec. 30

1918

3.71

3.35

Dec. 23

1918

5.79

1 . 32

Feb. 1

1919

3 47

3.75

Feb. 1

1919

3.74

2.26

V.\TIONS

Date
Collect

of
ion

Estimated
Amount

Oct. 27,

1917

Good

Oct. 29,

1917

Fair

Oct. 29,

1917

Cxood

Oct. 29,

1917

Poor

Oct. 29

1917

Good

Nov. 1

1917

Good

Nov. 9,

1917

Good

Nov. 13,

1917

Fair

Jan. 19

1918

Poor

234 Vn'wcrsil !i of Colifoniia riihlicdf ions In fiohnni | N oi,. 7

X. DlSTKir.l'TlOX OF UriiP.EH IX TIIK PLAXT
</. KFXaONAL DISTRIBUTION iX THK i'LAXT

Since our iiuiiii ciuleavors have l)i'('ii dirceted to an examination
of ;i laf<:e nunil)ei' of individual plants of a iiiiiiihei' of species of
Chri/sothainnKs and otlici- <>'enera, no <ii'cat attention lias been <>;iven
to many interesting and perhaps ultimately important details which
have to do w itii anatomieal peculiarities and with tlie ori<iin and occur-
rence of rul)l)er, whieh. under other circumstances, mijjht well have
concei'ned us. Intleed. various causes have resniti'd in an accumula-
tion of studies much less detailed concernino- these matter.s than was
ori<»inalIy anticipated. Again, since our interest lay pi-imarily in
discovering the average rubber content of the main rubber bearing
tissues and since it was early seen that the latter were peculiar to the
older parts of the stem, the younger portions of both stem and root,
the study of which is necessary for a thorough elucidation of the
problems of the origin and occurrence of rubber, were not usuall}'
collected in the Held and wei'e thus not available for stiuly in many
cases. At the present time, however, we are prepared to make certain
general statements concerning the distribution of rubber in ttie plants
with which we have been concerned, and leave for further description
a number of matters which are still under investigation. Particularly,
we aim in what follows to interpret for such persons as may concern
themselves with histological examinations of plants suspected of con-
taining rubber, the appearance of the sections wdiich they will be
likely to obtain and to indicate to them the nature and distribution
of the tissues in which rubber may be expected to occur. In this
endeavor we will emphasize the results of our histological studies, but
will also draw upon those derived from chemical analysis.

Attention has already been called to the fact that as in (hiayule,
so in Chrjjsothamnus we are dealing with a non-latex rubber which is
found deposited as such within the individual cells of certain portions
of the plant body. It may be noted in passing that many persons,
<'ven at the present time, are not acquainted with the nature of the
oeeui'rence of r(d)ber in (;ua\ule, noi' with the consetjuent processes
of extraction, tlie latter differing so decidedly fi'om those employed
ill the case of plants which bear latex-rubber. Since identical or simi-
lar i>rocesses will undoubtedly be employed should ("hi-ysil at any
time prove of sufficient importance to warrant its extraction on a

1919]

Hdll-Goochpccd : Ch rysil

285

commercial scale, it may not be out of place to refer tlie reader to the
description of the commercial process for the extraction of Guayule,
given by Lloyd. ^ -

With reference to the relative amounts and the general distribu-
tion of rubber in the various parts of the plant body certain points
seem rather well established. This is an important matter since
persons plaiming to extract Chrysil on a commercial scale would at
the start wish to know which portions of the plant contain maximum
quantities of rubber and should be harvested and which portions
should be discarded as containing negligible quantities. The follow-
ing table contains the results of such analyses as we have at hand
bearing upon this matter.

Table 7.-

Collection
Number

-Results of Analyses Made to Indicate the Relative Ajiouxts of
Rubber Borne by the VARiors Parts of the Plant
Body — Chnjsothamnus

Variety

144 (1) 'C. n. consimilis

(2)

(3)
(4)

Acetone

Extract

Per cent.

3 . 7.5
3.. 59

3.69

4.64

752 (1) 2C. n. viridulus 3.47

(2) 11.80

(3) 22.57
755 (1) -C. n. viridulus 3.74

(2) 15.10

(3) 19.80
620 (1) 'C. n. consimilis 2,89

(2) 3.. 54

(3) 2.99

(4) 2.89
(.5) 3.46

(6) 2.79

(7) 2.97

(8) 3.24

(9) 3.16

(10) 3.43

(11) 2.79

(12) 2.86

1 Collected, Golconda, Nev., Mar. 4, 1918.
:i Collected, Mono Lake, Feb. 1, 1919.
3 Collected, Ely, Nev., Feb., 1919.

Benzene
Extract
Per cent. Remarks

0 . 81 First 10 cm. up from end of root

2.12 10 cm. section, from first rootlets to soil
line

1 . 74 Main branches, 4 to S years old at bases

and 30 cm. long
1.85 Tops of branches, 3 to 4 years old a'
bases, 2 years old at tops
At soil line
2 year old shoots
Tops, 1 year old
At r^oil line
2 year old shoots
Tops, 1 year old
End of root, 10 cm.
Ne.xt 10 cm. up root
Third 10 cm. up root
Fourth 10 cm. up root
Soil line

First 7 cm. up from soil line
Second section up from soil line, 8 cm.
Third section up from soil line, 8 yrs. old
Fourth section, 7 years old
Fifth section, 6 years old
Sixth section, 5 years old
Seventh section, 4 years old

3.75
1.70
0.42
2.26
1.63
0.74
0.13
0.28
0.37
0.43
3.03
1.54
1.91
2.07
2.20
1.91
2.20
2.77

i« L.t

8.

2:](i

Universih/ of CaUfornia I'uhlicaiions in Botanij \^'oh.l

Table 7. — {Conlinuid)

Collection
Number

(13)
(14)
(15)
(16)
622 (1) ''C.

(2)

(3)

(4)

(5)

(6)

(7)

(8)

(9)
(10)

(11)
(12)
(13)

\'ai'ioty

n. consiniilis

Acetone

Kxtrapt
l^or cent.

2. SO

6.12
13.79
20.33
4.72
4 . 59
4.20
5.49
2.15
2.15
3.42
2.65
3.29
4.52
5.52
10.72
2.12

Benzene

Extract
Per cent.

1

1.
1
0.

43
.03
27
.01
. 3()
0.35
0.4S
0.96
1.56
2 . 36
2.12
1.88
1.64
2.39
0.96
0.83
0.86

3 Collected, Ely, Nev. Feb., 1919.

lU'riiarkrs

Eif;fith section, 3 years old

Shoots, 2 y(>ar,s old

Shoots, 1 year old

Tips, current year

End of root, 30 cm.

Next 10 cm. up root

Next 10 cm. up root

Next 10 cm. up root

Next 10 cm. up root

Soil line

Main branches, 5 to 0 years old.

Main branches, 4 to 5 years old

Main branches, 3 to 4 years old

Main branches, 2 to 3 years old

Shoots, 2 years old

Shoots, 1 year old

Tips, current year.

P\)r convenience of discussion the table above ha.s been differently
arranjred and somewhat condensed in the table which follows:

Table 8. — Thk Eelative Amounts of Rubber Found in Various Portions of
THE Root and in Portions of the Stem of Various Ages

Collection Number

144 ,

Va\(\ of root

0.81

Koot up to soil line (av.)

Soil line

2.12

Base of main stems, 0 years

old and

older

1.74

Stems, 4 to 5 years old

Stems, 3 to 4 years old

1 . 85

Branches, 2 to 3 years old

Braiu!h(>s, 2 years old

Branches, 1 year old

'i'oiis, current year

7o2

.75

1.70
0.42

755

2 . 2r)

1.63

0.74

620

622

0.13

0.36

0.32

0.83

:', . o:!

2.36

1.93

2 . 12

2.48

1.88

2 . 60

1.64

2 . 03

2.39

1.63

0.96

1.27

0.83

1.01

0 . SC)

These data indicate that in Chryaoihamnui^ the root proper
(below a point from six to ei^ht inches below the soil line) contains
little or no rubber. In addition to the two species for which analyses
are given in tables 7 and 8 we have analyses comparing the rubber
content of stem and root in the case of gnaphalodea and pinifolius
which give rcsnlts closely similar to those above. Attention should
ht'iT bt' called to the fact, discussed in detail elsewhere (p. 274) that

1919

Hall-Goodspced : Chrysil

237

Fig. '). A large ])laiit of Cltrii.sotluunnu.s
iiauKeosax var. rtriduhis. 1 indicates the as-
sumed soil-line; 2 the upper limits of the
iriain trunk; o the approximate up[>er limits
of the three- to four-vear-oM wood.

in certain species of Hap-
lopappus rubber is con-
fined to the root or occurs
in approximately equal
amounts in root and stem.
As far as stem struc-
tures are concerned it is
more difficult to indicate a
sharp line of demarcation
between those which are
significant and those which
are not significant as far
as rubber content is con-
cerned. Figure 5 shows a
large plant of vlridiilus
(no. 565) and serves to
indicate the characteristic
method of branching and
the general configuration
of plants of this and other
varieties which we have ex-
amined.

The assumed soil line,
the upper limits of the
main trunk, and the ap-
proximate upper limits of
the three- to four-year-old
wood are shown. The re-
sults listed in tables 7 and
8 suggest that only such
parts of the plant as lie
between 1 and 3 contain
sufficient rubber to war-
rant their harvesting if
the plants were used as a
commercial source of rub-
ber.

In the various tables
above and in those which
follow, analyses of portions

2oH r iiin rslh/ of (\ilifor)il(i I'lihllrafioiia i)i Botanii [Vol.7

(if phiiils t'lniiish the dala on the liasis of wliicli ;i ('crUnii |)('i'ct'iita<i('
nihlici- coiilciit is assi<iiic(l 1o a ^ivcii iiHlivi<liial. Fij»iii'(' 4 sliows
plants trinmicd for liriiidiii^' and analysis and it is ajiparent that
what is h'ft of the |)hiuts I'cprescnt areas i-ou<^hly ('(luivah'nt to the
portion lictwccn 1 and '1 in Haui'c ').

It appiars from tal»hs 7 and S tliat inaxinium (piantitics of rnlil)i'r
occur at the soil line. In the ease of futni'e examinations of plants to
li'd at the per cent of contained rulihei' it would appear that this
hasal poi'tion of the nuun ti'unk alone iiiijiht funnsh a characteristic
sample and that the whole ai'(>a between 1 and :{ ( ti^-. .") ) need not
he handled. In other words analysis of a soil line sanijile of such a
plaid as 620 (table 7, 620 (5) ) indicated that approximately 3 per cent
of rubber was ])resent and it is held that this fi<:ui'e uuiy safely be
taken as a fair a])proximation of the per cent borne liy the entire ])lant
exclusive of the I'oot and the shoots and twig's less than three or four
years old. At first sijiht it would appear that this figure is too high
since table 7 indicates that an average percentage content of segments
(o) to (13) of 620 would ajiproximate 2 rather than 3 per cent. In
this connection, however, the relative weights of the areas between
1 and 2 (cf. fig. o) and between 2 and 3 must be taken into account.
A lunnber of weighings indicate that the weight of that portion
between 2 and 3 is approximately 30 per cent of the total weight of
the region between 1 and 3. This total weight will roughly approxi-
mate 1500 grams. It appears from the data given that if the mai)i
trunk bears in the neighborhood of 3 per cent the branches up to
the limits of the three to four year old wood will bear about 2 per cent.
With this assumption in mind the following relation ajipears to hold
good :

Wcifrlit 1 to 2 = n.in gm., -which at 3% = 34..")0 gm. rublicr.
Wciglit 2 to 3 = 4."iO gni., wliit'h at 2% =^ 9.00 gni. rubber.

Total wciglit = l.")00 gm., total rubber = 43.50 gin.
Total weight = l.'OO gm. at 3% =; 4.5.00 gm. rubber.
Total weight = I.jOO gm. at 2% = 30.00 gm. rubber.

Based upon such rotigh estimates as these it seems clear that, due
to the greater weight of the main trunk as compared with the older
blanches, a report of per cent of contained ridiber based upon an
analysis of portions of the i)lant near the soil line would not give a
wholly errone()ns estimate of the amount of rubber to be obtained
fi'om a plant harvested so that jiarts from 1 to 3 (tig. .")) alone were
I'etaiucd.

I9if»j Hall-Goodspecd: ('lirijsil 239

As stated above we liave been unable to secure a sufficiently large
number of analyses of the younger and older portions of plants to
•make our conclusions as to regional distribution of rubber entirely
authoritative. The surprising decrease in rubber content a short
distance above the soil line is perplexing as is the succeeding increase
in the upper portion of the main trunk and branches. Elimination
of the primar.y cortex by peridermal activity in the older trunk
elements is obviously not an adequate explanation since at the soil
line maximum elimination of cortical tissue has taken place. Indeed,
cork formation with the accompanying distintegration of rubber cut
out thereby has progressed to a decided extent before the third
year of growth. The results obtained suggest, among other things,
a translocation of rubber or its derivatives. Such implications deserve
nothing more than a mere mention when the data are so fragmentary.

h. SPECIFIC DISTEIBUTION IN MATURE TISSUES

In an earlier paragraph it was stated that for making a rough
estimate under the microscope of the amount of rubber borne by a
given plant, a piece of mature tissue was cut from the main stem
axis. For convenience we may use as the basis for a description
of the tissues in which rubber, if present, may be expected to occur,
the accompanying photomicrographs and photographs of stained
preparations. As will be noted in plates — and — the stained areas
are for the most part extracambial although the ra^'s are also con-
spicuously stained. Anah'ses of one plant indicate that the amount
of rubber in the cortex as compared with that in the woody cylinder
is in the ratio of approximately six to one.

Not all the stained areas without the cambium are rubber bearing,
however. Sudan Til stains intensely the walls of corky tissues and
the superficial portions of the stained material outside the woody
cj'linder represent cork and bark. Within this outermost layer the
broad wedge-shaped extensions of the primary rays are most conspicu-
ousl}^ stained. In a plant bearing from 4 to 5 per cent of rubber all
the cells of these ray extensions are filled with rubber inclusions.
Indeed in microscopical examinations of Chrysoihamnus these, areas
should first be examined for the presence of rubber as, uniformly, in
sections of stems over two 3'ears old, stainable rubber inclusions, if
present at all, will here be found.

Bands of stained tissue will be seen connecting tangentially the
cortical ray extensions. These represent the parench^ymatous elements

240 f'iiii'( rsil 11 (if (\il ifonilii I'lihllcaf ions in Ihildinj | Voi,. 7

of {he cortex and iinii-ruiict ional plilociii iiicrv niciits. 'I'lii- t'mictioiuil
pliloi'iu aiH'as ai'c also nil)l)('i' l)t'ai'iiiji' altlioujili in tliis case tlic
protoplasmic matrices of the cells often stain in simulation of rul)ber
inclusions. The sieve tubes and companion cells are (piite small in
eross-seetional diametei- and oidy in longitudinal sections can rubber
lildbides be identitied within them. The ])hloein parenchyma bears
the lar<»'est amount of ruitber in the phloem ai'ea and even in cross-
sectional view thei'e is no difficulty in identifA-ing the stained inclu-
sions as such.

The candiium aj^pears consistently to bear rubber inclusions in
stems moi'e than one year old. The rubber is in tlie form of small
globules which seem to increase in innnber during cambial activity.
As an annual cand)ium increment towai'd the j)eriphery begins to
become diffVi-eutiated into characteristic phloem elements these snudl
globules tend to agglomerate. Intracambially. however, the partially
differentiated xylem elements contain no rubber inclusions although
originating from the same nd)ber bearing cambinm which passes
on its rubber inclusions to the developing phloem increments. This
situation might furnish a starting poini for investigations on the
chemical or physical constitution of the j^rotoplast which is specific
for the differentiation of rubber inclusions or essential for the perma-
nence and persistence of such inclusions. In none of the mature xylem
elements, parenchymatous or otherwise, have rubber inclusions been
found.

Under the microscope one of the most conspicuous features which
attracts attention when preparations such as those shown in plates
18 and 1!) ai-e examined is the increase in I'ubber deposition at certain
l^oints along the rays. Although this matter is not well brought out
in all the photographs it may still be seen that at points correspond-
ing to the spring wood of each annual xjdem increment the amount
of rubber is less than at those points along the rays which correspond
to the summei" or fall wood. This situation will be mentioned (dse-
where in eomiectif)!! with a discussion of seasonal vai'iation in i-ubl)er
content.

Plates IS and Ifj (exclusive of figures 4 and 5) show cross sections
of stems of vaiious ages. The stem of the current year shown in
j)late 18, figuie 1, is included to call attention to the deep staining of
the chlorenchyma, the absence of stainable substance in cortex and
rays, and es])ecially the large (piantity of stained ru])bei' in the xylem
intrust ion of an axillary bud. The three year old stem in plate 18,

191«] Hall-Goods peed: Chnjsil 241

figure 3, illustrates the statements made above as to the conspicuous
staining of the cortical ray extensions and the presence of rubber in
the phloem and rays. It will be noted that the portions of the rays
lying in the first annual increment contain relatively negligible
amounts of rubber as compared with the portions of the rays lying
in the second and third year xylem. The line " C " marks the approx-
imate limits of the living cortex.

Figure 2 in plate 18 is included to show the extent of the rubber
bearing tissues and of the non-rubber bearing cork and barky
elements. The "A" line indicates roughly the external limits of the
rubber bearing cortex. Cork has formed below this line but the
primary cortical tissues cut out thereby show rubber not yet entirely
disintegrated. The "B"-line points to the tangential strip in which
maximum (quantities of rubber occurring in the cortical parenchyma
and in the cortical ray extensions may ordinarily be expected to
occur. Figure 2 is a cross section of a five year old stem.

In plate 19 the two cross sections give evidence as to the distribution
of rubber in mature stems. Figure 1 was cut from a six year old
stem and figure 2 from a nine year old stem. The greater portion
of the bark and inner cork is absent in figure 1 and it will be seen that
the cortical ray extensions filled with rubber extend outward for a
long distance. The broad ray extension on the extreme left in figure 1
is characteristic of many of the cortical extensions of the primary
rays and its cells are densely packed with rubber.

Figure 2 in plate 18 shows a particularly thin section from which
in places the rubber inclusions have fallen or have been washed out.
Line "A" indicates roughly the limits of the rubber bearing tissues;
those external being composed largely of cork and bark.

We have found that rubber is present in the cell either in what
may be spoken of as a '' diffused "" state or in the form of globules.
In the difl'used state the entire cell cavity is filled with a mass of
stained substance which appears to be somewhat homogeneous. This
condition is shown in certain of the cells in plate 18, figure 4, the
spherical globules representing detached masses of rubber which have
rounded up when free from the cell wall. When present in globular
form, one or more large spherical globules partially fill the cell cavity
or a number of smaller definitely spherical globules are distributed
without apparent arrangement within it. Spherical globules in situ
are shown in a number of the cells in plate 18, figure 5. For a time
it seemed possible that the form in which the rubber occurred in the

242

I'liliu rsil !i (if Cuiiforii'ui /'iil)lic(iiii>iis in liotdiu) [Voi,. 7

coll niifilit serve as one of a iiuiulx'i- of bases for species dilfereiitiatioii
in Chriisofhdiiniiis. Cei'taiiily ('. hrrf ifolius (cf. |). 266) consistently
exhibits I'libber only in the foi-ni of lai'<:-e spliei'ical <i,-lobnles which
usually occnr sinj^ly and almost exclusively in the cells of the cortical

y^^

Vig. *i. Portion of a cross-section ot" a mature loaf of ('liriisotluuuiius iiinisro»us
var. ronsiniilis. I'latit <rro\vii in tlio University of Califoi'iiia Botanical (iarden.
J-Jcrkclcy. Plioto-niici-ogi'apli.

parenchyma. In the ikihscusus group a number of <»lobules often are
present in the cells of younjii: rubber bearing tissues but thej' tend
to a<i;j?l()merate with age so that the corresponding tissues when mature
show few if any globular rubber inclusions, this substance being
present in the "difTnsed" condition. Such a condition does not obtain

1^19] Hull-G Godspeed: Chnjsil 243

for teretifolius, globules only being found both in younger and older
elements. The cross section of a leaf of a plant grown in the
University of California Botanical Garden is shown in figure 6. The
dark stained rubber globules in the palisade cells are conspicuous.
Lloyd^^" reports that a single large globule of rubber was character-
istic of each palisade cell in certain portions of the leaf in Guayule.
A corresponding condition holds for Chrysothaninus although in some
instances it appears that more than one globule may occur in a single
palisade cell. Various problems dealing with the origin of rubber and
its distribution in younger tissues have been investigated in a prelim-
inary way, at least. The results of these studies will be published in
the near future as the fourth paper of the present series.

XI. FACTORS INFLUENCING RUBBER CONTENT

a. VAEIATIOX WITH THE BOTANICAL VARIETY

The results of our studies, more particularly the chemical and
microscopical examinations, indicate that rubber is present in only a
few of the major species of Chrysothamnns. The species examined
and in which it was not found with certainty are : C. Greenei, C.
Howardi, C. nevadensis, C. Parryi, C. hiimilis, C. piiherulus, and
C. viscidiflorus. In C. liuifolius, which is perhaps only a varietj^ of
viscidiflorus, it was found to the extent of 1 per cent in one sample
but was lacking- in another. The species of Chrysothamnns now
known to yield rubber are (\ nauseosns, ('. inrhinatus, ('. trretifolins.
and C paniculafus. The last three of these are discussed elsewhere
since they are of minor importance and the product is not Chrysil.^"

Coming now to a consideration of C. nanseosns, we find that this
variable and widespread species breaks up into about twenty-two
varieties. It may be predicted with reasonable certainty that Chrysil
will be found in all of these twenty-two forms. This statement is
based upon the fact that of the thirteen thus far investigated every
one has yielded a greater or less amount of this substance. It might
be assumed, a priori, that certain of these botanical varieties would
carry a consistently higher percentage than others. This assumption
is borne out to some extent by the analyses, as is indicated in thi'
following summary of the analytical tables published in detail on
pages 226 to 233.

18" L.C., p. 184.
i:i See pp. 265-268.

244

Univrrslhj of Califoniid I'lililicatioiis in Botanji f^'"'-

SuMMAitv or 'I'ahm: (i

\;irict>

1

oons'inilis

2

viridulus

3

Hn:ii)h;ilo(los

4

spcoiosiis

o

Kr;ivooI(Mis

G

pinifolius

7

hololoucus

8

inohavensis

9

occidental is

10

loiospermus

11

frisi'lus

12 naiisoosus (typical)

Viimbor of
samples
analyzed

Hiihhcr ill
poorest
sample

Per cent.

Uiil)l)iM- ill

best

sample

I'irr cent.

.\verane

of all
samples
Per cent

m*

0.39

6 . 57

1.97*

■.m

0.44

5.56

2.52

1S

0.26

3 60

1.61

17

0.16

2 77

1.18

10

0.07

3.19

0.83

5

1.11

3.9S

2.95

4

1.05

4.1)

2.83

4

0.30

1.08

0.53

3

0.71

1.54

1.07

2

0.,S4

1 17

1.00

1

1.86

2

(inicro.sc()i)ical exam-

ination only

)

traces

arc here f)initto(l since the saiiir

>le.s were imp

erfect.

Ill making' use of tliis table it is bettor to take into account only
the first five varieties since the others are not represented by a suf-
ficiently lar<i:e number of samples to rtnider the rc^sults dependable.
''Individual variation'' is an important factor and it is only after a
laro:e number of samples from widely separated localities have been
analyzed that one is justified in drawing conclusions.

Taking tlicii only the first five varieties it would seem tliat tlu-y
would stand, as to rubber content, in the order of virkhdus, consimilis,
rjnaphalodfs, speciosus, (jravcolens. While tliis may be of significance,
it is also to be noted that the first two are inhabitants of alkaline flats,
the next two belong to non-alkaline slopes, and the last (as far as our
samples are concerned) to only moderately if at all alkaline soil. It
may be. tlierefore. tliat an apparent parallel between r\d)ber content
and botanical varieties is. in fact, due to environmental factors.

While, as just shown, tlici'i' is no direct evidence that the botanical
varieties represent hereditary units, eaeli witli a dift'erent capacity for
iiil)l)rr pi'odnetion, tliere is nothing, on tlic other hand, to indicate
that eacli vari<'ty may not itself be composed of several or even
numerous biotypes. If this is the ease, Ihen tlie separation of these
tlirough selection might lead to the discovery of a superior strain.
The notable fluctuation in the rubber content of any one variety might
seem to indicate this possibility but the alternative as to the influence
of I'Mvii'diiincnt must also be kept in mind.

I9ii^] Hall-Goodspeed: Chrysil 245

h. VABIATIOX DUE TO ENVIRONMENT

Attempts have been made to correlate the differences in the rubber
content of individual plants with various factors of the environment.
This has not led, as yet, to positive conclusions. One reason for the
failure to obtain definite results is the low percentage in even the best
plants, so that auA* slight variation in the selection, preparation, or
analysis of any two samples might more than offset any original differ-
ence in their composition. Only through extensive experimental work,
can conclusive results be hoped for.

Notwithstanding the difficulties just mentioned, a few observations
may be permitted. The water content of the soil is often considered
to exert an influence on rubber deposition in plants. Specimens of a
single variet}' of Chrysothamnus }wuseosus selected to determine this
point did not var}" widely in the amount of Chrj'sil present. There
was, if anything, a slight balance in favor of those growing on moist
banks but tliis difference was so slight that it is probably of no sig-
nificance. However, when different varieties are considered, and only
the average of a large number of samples taken into account, it is
found that those varieties which inhabit the moist and poorly drained
vallev bottoms contain rubber in larger amounts than do the varieties
of the \ery arid and well drained surrounding slopes.

This is indicated by the table on page 244. The varieties there
numbered 1, 2, and 6 are lowland forms and these run higher in rubber
content than do numbers 3, 4, 8, 9, 10, 11, and 12, which are upland
forms. The single apparent exception to this rule is number 7, but
this may be due to the small number of samples analyzed for tliat
variety.

But. while the formation of rubber thus seems to be favored by
the more abmidant moisture of the lowlands, another factor may be
the real cause of the increa.se in percentage, namely, the presence of
alkali. It is not impossible that the alkali acts indirectly through the
water relation. It is well known that its presence reduces the amount
of water available to plants, and it is conceivable that by thus lowering
the chresard it causes a deficiency in physiological moisture which is
even greater than the deficiency on the surrounding slopes. However
this may be, it is certain that the best rubber producing varieties of
Chrtjsothamnus nausposus are those of the alkaline valley bottoms.

As to temperature, it seems unlikely that this factor exerts an
influence on rubber formation except as it acts through other agencies.

246 Univcrsih/ of Call for iiia Piihlicathns i)i Botmnj [Vfn,. 7

Tlit'iv is no sIriUiiig tlilTcrciico between the pci-ccntii^cs found in the
plants from the hot desert valleys of California and tlie cold niounlain
valleys of ('olora(ht. 'i'he apparent i'unnin<i down of rubber content
as one ai)proaehes the limits of distribution for tlie s])ecios is j)erhai)s
to be accounted for on otliei- <i'rounds. [f temporatnro were the con-
trollinfi: factoi" it would be dit^eult to explain the unifoi-ndy low
rubber content of plants from the warm Coast ranfres of California
and also of the cool plateaus at the eastern base of the Rockies.

The effect of wounds and of pruning- may be a matter of much
importance in case the ))lants are ever brouglit under cultivation.
The wild shi-ubs are often riddled l)y the attacks of larvae and l)eetles,
but their influence, if any, is not known. Experiments now in prog-
i-ess have ali'cady determined that the plants may be pruned back
without injury and that this greatly increases the number and W'cight
of the rubbei' bearing stems. The possible effect upon the formation
of rubbei" will be determined later. (See plate 20.)

('. SEASONAL VAKIATION

In the case of Guayule a variety of evidence points to the fact
that there exists a sti'iking seasonal variation in rubber content. Thus,
it has been found that during the active season of growth onl\' ncgli-
gibl( (piantities of rubber are deposited while during the following
resting period rubb(>r nuikes its appearance in the tissues. These two
periods are correlated with the duration of the rainy season in the
desert region where Guaj'ule grows wild. During the rains growth
is initiated and continues for a time. Shortly after the start of the
dry season active vegetative growth ceases and rubber then begins
to be deposited in the recently formed tissues.

With these facts in mind some experiments were made to determine
whether a similar situation holds for rubber deposition in Chryso-
tharnnus. Table 9 gives the results of these experiments. On Sep-
tember 16, 1918, i)()rtions of mature tissues were cut from thirteen
plants representing three different varieties of naiiscosiis growing in
a small tract near Benton, California. Notes and photographs nuide
it possible on December 22, 1918, to remove from the same i)lants
portions ai)j)arently etpiivalent to those which had been secured in
September. Thei-e was no evidence that the removal of the first
portion had in any way imjiaired the i-enutinder of the i)lant or im-
peded its normal develo]>ment. The Se|iteml)er date is taken to
represent the close ttf the growing period at which time the flowers

1919]

Hull-Goodspcecl : Ch rysil

247

are produced. By the end of December the resting period is prac-
tically at an end since shortly thereafter the early spring growing
season begins.

Table 9. — Seasonal ^'ARIATION in Kubber

ChrysotJuimnus na

(September 16, 1918, represents the close
ber 22, 1918, the close of the resting season.)

Content of Three Varieties of
of the growing season and Decern-

Acetone

Benzene

Place of

Date of

Extract

Extract

Variety

Collection

Collection

Per cent.

Per cent.

441

C.

n.

viridulus

Benton, Calif.

Sept. 16, 191S

2.80

2.31

Benton, Calif.

Dec. 22, 1918

3.16

2.86

442

C.

n.

viridulus

Benton, Calif.

Sept. 16, 1918

3.18

3.97

Benton, Calif.

Dec. 22, 1918

3.58

3.77

443

C.

n.

viridulus

Benton, Calif.

Sept. 16, 1918

2, 52

3.11

Benton, Calif.

Sept. 16, 1918

2.18

3.27*

Benton, Calif.

Dec. 22, 1918

2.82

3.06

Benton, Calif.

Dec. 22, 1918

2.97

2.95*

444

C.

n.

viridulus

Benton, Calif.

Sept. 16, 1918

2.60

3.83

Benton, Calif.

Dec. 22, 1918

2.80

2.63

445

C.

n.

viridulus

Benton, Calif.

Sept. 16, 1918

3.13

5.29

Benton, Calif.

Dec. 22, 1918

3 . 13

3.49

446

C.

n.

viridulus

Benton, Calif.

Sept. 16, 1918

3.61

4.16

Benton, Calif.

Dec. 22, 1918

3.87

3.97

447

C.

n.

gnaphalodes

Benton, CaHf.

Sept. 16, 1918

3.66

2.66

Benton, Calif.

Dec. 22, 1918

3.43

1.52

448

C.

n.

gnaphalodes

Benton, Calif.

Sept. 16, 1918

3.. 55

3.58

Benton, Calif.

Dec. 22, 1918

4.76

3.95

449

C.

n.

gnaphalodes

Benton, Calif.

Sept. 16, 1918

2.39

1.85

Benton, Cahf.

Dec. 22, 1918

2.79

0.99

450

C.

n.

gnaphalodes

Benton, Calif.

Sept. 16, 1918

2.50

2.09

Benton, Calif.

Dec. 22, 1918

2.63

1.65

451

C.

n.

, hololeucus

Benton, Calif.

Sept. 16, 1918

2.87

1.03

Benton, Calif.

Dec. 22, 1918

3.44

0.79

452

C.

n

. hololeucus

Benton, Calif.

Sept. 16, 1918

2.98

3.98

Benton, Calif.

Dec. 22, 1918

4.77

4.06

453

C.

n

. hololeucus

Benton, Calif.

Sept. 16, 1918

3.14

2.22

Benton, Calif.

Dec. 22, 1918

3.98

2.12

♦Duplicate analysis.

It seems probable from the results listed above that during the rest-
ing period the resins and other acetone-soluble substances increase in
amount while the amount of rubber is diminished. This situation is
unexpected in view of the reverse condition described above for
Guayule. It is, also, of particular interest when the influence of stor-
age of shrub upon resin and rubber content is recalled (cf. p. 224).
In the case of storage the per cent of resins was considerably reduced
along with a slight reduction in per cent of rubber. Obviously there

248 Universliji of Calif ornm I'Khlicatiotis in Bota)nf \\'o\..7

is need of fiii'llu'i- oxpoi'iiiiciits mikI ;iiialyses to continn tlic I'csiilis of
tln' iiioi-c ur less prcliniiiiary experiments tabulated above. Iiub'cd a
few ex('e|)tioiis to our f^eiiora] conelusion appear in labU" 9 and while
not mmii'rous eiiou<^li to att'eet it materially eertaiidy cast some doubt.
n|>oii it. It is entir<'ly possiltle that in selectiufr tlie two dates for
i-olleetion of sam|)l('s the critical jx-riod may liave been overlooked.
Thus, if a sample had been taken in August or earliei" as well as in
Septemlx'i' a decided iiicrease in ?-nbber content mij»ht have been
noted for the lattci" date. This would have meant that tlie flowering
period in Chnjsothamnvs represents, in terms of seasonal variation
in rubber content as observed for Guayule. not tlie eiul of the grov^'ing
])eriod but the beginning' of the resting period tlui'ing the start of
which maximum deposition of rubber takes place very rapidly.-'' Had
such collections been inade and such results obtained no light wo\d(l.
however, have been thrown upon the seeming disappearance durin>i-
the i-esting period of a certain proportion of the nd)ber present within
a month of its start.

XI!. METHODS OF HARVESTING: SEASON. AGE, DEPTH

OF CUTTING. ETC.

The methods of harvesting have not been woiked out in detail since
it has not been demonstrated that Chrysothannnis can be utilized on a
commercial scale. However, our preliminary studies have indicated
certain general principles to be observed in case rubber is ever prepared
from the plants and they are here set forth as follows.

The question of the distribution of rubl)er in the various plant or-
gans and the extent of its occurrence in stems of various ages is dis-
cussed elsewhere (cf. p. 234). It may here be said that the amount of
rubber in i)arts less tluui three years old is relatively small. From the
thii'd year onward for one oi- two years the amount increases rather
I'apidly. After the fifth or sixth year the (piantity of i'id)ber ])i'esent
remains more or less constant. The cause of this constancy in amount
depends upon certain facts taken up in detail elsewhei-e. In the first
place, the extracambial tissues ai'c almost the exclusive i-egions of lub-
ber deposition. Secondly, when cork fornuition takes place there is.
each yeai". an amount of nd)ber containing cortical tissue cut off whieli
roughly appj'oxinuites the amount of new cortieaj tissue aniuudly

-" The microscopical evidence (cf. j). 240) appears to give a partial confirinatioii
of such a supposition.

1 i'l ^M UaU-Goodspofd : ( 1> rysil 249

laid down. Finally, with the appearance of the phellogen deep in the
cortex and the production by it of a few layers of cork the rubber in
the cells external thereto begins to degenerate and soon loses its char-
acteristic staining capacity. It is true, of course, that the annual in-
crements added intracambially to the primary medullary rays and
which in part are devoted to the origination of secondary rays in-
creases each year the number of cells in the woody cylinder available
for rubber deposition. There is in this case, obviously, no such annual
reduction in the amount of rubber bearing tissues as is characteristic
of the cortex. The amount of new rubber bearing tissue added each
year to the existing ray material is so small and the quantity of rub-
ber present in the individual ray cells is so relatively diminutive, that
this source of increase in the rubber content of old as compared with
younger stems is negligible.

The problem of a seasonal or periodic deposition will re(|uire for
its final solution a much more detailed study than we liave been able
to give to it. The experiments mentioned on page 246 seem to indicate
that the maximum amount of rubber is present near the close of the
growing season. If these results are confirmed they will have a
directive influence in determining the season at which harvesting
should take place. This probably would not be a determining factor
in selecting the season for harvesting wild shrub, but it might mean
the difference between financial success and failure when the handling
of cultivated plants is under consideration.

It would follow from the foregoing that old plants would be the
best rubber producers. It is evident that as the plant becomes over-
mature decay sets in and reduces the weight of the rubber earrjang
portions. This is borne out to some extent by the results of our
analyses, which indicate that the average percentage composition is
about the same for large old plants as for medium sized plants of
moderate age. Young plants, say four years old or less, are almo.st
always low in percentage content, a direct result of the fact that
deposition is small until about the third year and the ratio of rubber
bearing tissue to the whole plant is therefore low. This discrepancy
gradually disappears with age. It is quite certain that shrub which
has reached maturity, that is, six to ten years for the viridiihis form.
bears its maximum quantity of rubber.

The depth at which the plants may be cut in harvesting without de-
stroying the power of regeneration is a matter which has received con-
siderable attention because of its practical bearings. Histological and

^oO Univrrsihj of California Puhlicationa in Botdini | Vol. 7

chemical analyses of Ihc loot indicate that it carries its highest per-
centage of i-iibher at about liic siii-face of the soil and that the per-
centage i-apidly tliininishes downward, iiiiti] at an avei-age depth of
al)out four inches the amount present is so small that the remainder of
the root is of no value. In harvesting, it would therefore be desirable
to make the cut about four inches below the surface. Our ('Xjx'riments,
performed on over three hundred plants of three leading varieties,
have definitely determined, however, that if this is done, or if the
root is cut oft' anywhere below its junction with the stem, the portion
remaining in the soil will die. The obvious explanation of this phe-
nomenon is thai, as in most plants, new shoots arise only from stem
tissue. In all cases where the stems were cut off just above the soil
surface the stumps promptly sent up an abundant growth of new
shoots. In one case a diagonal cut was made just at the surface so that
about an inch of stem was left on one side of the stump, while on the
opposite side the cut extended to about an inch below the top of the
root. After a lapse of six mouths there was a copious growth of shoots
two feet long from the higher side of the stump, none at all from the
lower.

The line of demarcation between stem and root is not a sharp one.
A microscopic examination for dormant buds is, of conrse, impractical
in field woi-k ; the bark is so similar on the lower part of the stem and
tlic upper part of th(^ root that its texture and color furnish no clue;
and the soil level cannot be accepted as a criterion since it is easily
modified by disturbing agencies such as erosion, deposition, accumula-
tion of vegetable matter, and the activities of rodents. In most cases,
a cut made just at the first crotch, which is usually also at the original
soil surface, will safely provide for regeneration but much experience
will be necessary if one desires to obtain the maximum amount of rub-
bei- and at the sauK^ time be assured of new growth without replanting.
In case the wild shrub is ever utilized for an emergency supply of
rubber it will probably be best to make the cut at about six inches be-
low the surface, thus sacrificing the roots for the sake of the rubbei"
in tlicir upper portions; on the other hand, if Chrysotlunnniis is grown
as a field crop it will probably be found profitable to provide for vege-
tative reproduction, either by leaving an inch or so of stem oi- by leav-
ing portions of several of the lower branches as a foundation for a
lai'ger and bi-oader plant.

The bulk of the shrub may be materially reduced without much
loss of rubbei- l)y cutting off and discarding all growtli less than thi'ee

li'i^J Hall-Goodsp<'((l: Chnjsil 251

years old. Wood which is just in its second year may or may not be
worth preserving, depending upon methods of milling, local conditions,
and the value of the rubber. The age is easily determined in young
stems by counting the growth rings. The color and texture of the
bark may also be used as a guide by one familiar with the plant. The
removal of all growth of less than three years will take with it all of
the leaves, but these, like the twigs themselves, carry only traces of
rubber.

XIII. POSSIBILITIES OF CHRYSOTHAMNUS AS A
CULTIVATED PLANT

It is eminently desirable that a portion of the rubber consumed in
the United States should be produced within our own borders. It is
the only important commodity essential to modern warfare which we
have not yet learned to produce. If tlie industry of rubber growing
Avere once established, even though it yielded only a fraction of our
normal needs, we could, through economy in use and through govern-
mental encouragement in war time, render ourselves independent of
other nations, who might refuse to supply our needs or who might be
unable to transport their products across the seas.

These same arguments can be advanced for the support of investi-
gations looking to the production of synthetic rubber and there is no
doubt that this line of work should also be encouraged. It is even pos-
sible that the synthetic product will in time replace that from the
plantations. We are not in a position to forecast what the future may
bring forth in this connection, but the opinion of those best capable of
judging seems to be that high grade synthetic rubber in large quan-
tities is something which we need not expect for a long time to come,
if at all. In the meantime, dependence must be placed upon importa-
tions unless we can discover some commercially profitable method of
growing rubber plants in our own country. The list of plants to be
considered in this connection is a long one. It includes various exotics
some of which are now grown in foreign countries for their rubber but
none of which have been given a conclusive trial here. Certain of our
native latex bearing plants, such as the milkweeds, spurges, dogbanes,
etc., are now under investigation by the Carnegie Institution of Wash-
ington with some promise of success.-^ The Pinguay, or Colorado

21 Caruegie lust., Wash. Year-book no. 17 (1918), p. 297.

2")2 JJniversihi of CdJifoniia f'lihlicdfiotis in Jiotdtn/ |\in.. 7

Ivubbcr Plant {II ijiik no.ijjs j{urihiiit(Ui i(tilis) lias hccii iiiisiu^eessfully
tried and the (Juayidc a shrub native to oui- soutlurn boidcrs and to
Mexico, is now iukUt ti'ial in Arizona. Since none of these rubber
])lants luis as yet been ajjrieulturally established in the Tnited States
it would seem worth while to yive attention also to (Uiri/sodKim iiu.s as
a possible plant to t)e brou<:ht under cultivation t'oi- its luliher.

If a further incentive is needed for the study of rubber plants
suitable to our conditions it may be found in the extent of uinised and
apparently unusable lands that are still to be found in our western
states. The inti-oduction of new af?ricultui'al industries for the i)roper
utilization of these practically idle lands is one of the most important
problems that now confront the people of the arid West. While we
hesitate to predict that they will ever l)e used foi- the pi'o(hiction of
rubber, yet that this is within the realm of i)Ossibility is evidenced by
the planting of a considerable acreage to Guayule in southern Aiizona
in 1918 by one of the leading rubber companies.

Guayule is a small ^Mexican shrub belonging to the same botanical
family as th(> Kabbit-brush, although not vei-y closely related to it.
The similarity between the two is such that a consideration of the
Guavule situation mav furnish some evidence as to the possible worth
of Chrysothamnus. The wild Guayule yields a "rubber" which
had been imported into the United States in considerable (luantity--
hefore the political troubles in Mexico interfered with the industry.
The supply of wild shrub was constantly on the wane, which \vd to
extensive experiments in propagation, selection, breeding, and culti-
vation. The wild plants carry an average of 10 per cent of "(hiayule
gum," as it is called, but when they were brought under cultivation
it was found that some plants yielded only 2 or 3 per cent, and in some
cases less than 1 per cent, of pure rubber. The application of .scien-
tific methods of breeding and selection, together with the control of
environmental conditions of growth have now brought the yield to as
high as twenty-seven per cent in a f(>w plants. We understand that
these high percentage strains are not suitable for iield gi-owing Imt
that those selected for j)lanting on a large scale yiehl about tifteen to
seventeen per cent of rubber in their fifth or sixth yeai".

These experiments with Guayule extended ovei- a ten-year ])ei-iod
and ai-e said to have cost one company alone in the neighborhood of
$r)0(),0()0. Their promise, however, may be inferred from the present
intention of the ccmipany to undertake planting in Arizona on a large

;!2 (19,000,000 poniuls in 1911.)

1 !' 1 ^ 1 Hall-Good speed : Ch rysil 253

scale. Whether or not Chrijsothcuiinus eould be "improved"' to the
same extent is a question which we are unable to answer, for the two
plants are so different in their life histories that methods used success-
fully with one might entirely' fail with the other. Nevertheless it seems
logical that the possibilities of both should be tested out before either
one is exclusively selected for cultivation, since each possesses certain
advantages which the other does not have. It is in order to draw a
comparision between the two that the history of the introduction of
Guayule into cultivation has been briefly outlined above.

The great superiority of Guayule lies in its high rubber content.
As stated above, this averages ten per cent for wild plants as against
an average of only 2.5 per cent for the I'iriduhis form of Chrysotham-
niis. These figures are based upon pure dry rubber and dry shrub.
Guayule; moreover, has been extensively studied : its habits, and its
response to treatment, are at least somewhat understood, and the work
thus far done upon it has resulted in the development of a fifteen to
seventeen per cent strain. Tlie improvement of Chrijsofhamnus could
undoubtedly be brought about, at least to some extent, by selection
and ])reeding. However, since it is presunmbl>' a cross fertilized
plant, the separation of .superior strains would be a more difficult mat-
ter than in Guayule. A starting point is indicated in that certain in-
dividuals of the variety consimiUs actually ran as high as 6.7 per cent.

Guayule has a still further advantage in that the methods of field
management, of milling, and of marketing are also established and the
product is well known to the rubber trade. Here, however, the ad-
vantages of Guayule over ChrijsotJiamnHs seem to end.

As an offset against the above more favorable attributes of Guayule.
and especially the higher rubber content, we find a number of points in
favor of Chrysothdnuius. These are here set down in order that the
^Mexican plant may be used to some extent as a gauge in measuring the
possibilities of tlie other one.

(1) Chrysofhdiiiiius is a larger plant than Guayule, tlie wild shrubs
averaging six {)ounds of rubber carrying wood, whereas the plants of
the latter weigli one-half to three ]iounds, rarely attaining to six
pounds.

(2j Chrysutliaiinius is a native of the western I'liited States and
is therefore adapted to our conditions. It grows where the tempera-
ture falls to zero and probably much lower, since it ranges up the
mountains to over 7000 feet altitude. In the consimilis form it is
known to grow over large areas where the temperature not infrequently

1254 J^iiiri rsHi/ of Califoniid P\<hJ'\r(it\(nis in liutain/ r^'<"--7

n'iK'lics — 20^ h\ (iuayulc on the (itlii'i- hiind. is Too tciuU'r i'of any
but oui- uaniici' valleys and i'\cn there its cultivation is apparently at-
tended with soiiu' I'isk. Its intr(xluction into this country is an exten-
sion of its natural range, whei'eas the cultivation of (.'hrysothamnus
would be merely the utilization of a native plant adajtted by nature to
our region.

(3) Chri/sotJiamniis is verj' resistant to alkali, often growing on
soils too alkaline for any ordinary agricultural ci-op. It could there-
tVti'e be grown on land not now utilized and which could V)e obtained
for a very moderate price. Guayule is sensitive to alkali.

(4) The water ri^piirement of Chrijsotliannius is considerably less
than that of Guayule. It could certainly be grown without irrigation
on the cheap lands mentioned in paragraph 3.

(5) The })i-oduct is superior to that of Guayule (see ]). 188). It
would bring a higher price in the market and if need arose would
more nearly replace the tine imported rubbers in oui' industries.

(6) Chrysothanniiis is more easily and cheaply propagated, both
from seed, which form in abundance and are easily germinated, and
vegetatively. Both old plants and seedlings may be transplanted with-
out loss.

(7) Certain cultural practices have been developed in connection
with Guayule that result in an increased yield of i-ubber, and methods
of operation have been devised that permit of cheap handling through
the use of machinery, so that the crop is said to be "machine-grown."
There is no reason to suppose that similar practices and methods cannot
be perfected for ('Jirijsothannius. The larger size of the shrubs and
tlieir habit of throwing up numerous new shoots when cut back indi-
cate^ the possibility of increasing the tonnage of rubber carrying shrub
by the cutting back of the tops (see pi. 20). Experimental work along
these lines has not yet progressed to a stage where estimates would be
warranted, but it is believed that a method can be worked out that
will multiply by a considerable factor the tonnage of shrub per aere-
3'ear. It is possible that such mani]nilation may at the same time
favor an increased formation of rubber in the tissues.

It would seem, therefore, that the possibilities of ('hrj/soflidnniKs as
a cultivated rubber plant should be looked into more closely. Any
spe<'ies which gives even the least promise should be thoroughly studied
in ortler to find, if possible, a new crop that will turn our waste lands
into productive fields and at the sanu^ time safeguard the nation against
a possible deficiency in rubber during critical periods. These investi-

1^1^] Hall-Goodspc( (J : Chnjsil 255

gations, as applied to Chnjsotliamnus. should be along several diverse
lines, yet all leading to the same objective. They should include the
following : detailed studies of wild plants for the purpose of discov-
ering, if possible, better varieties or strains than we now have ; garden
experimentation in breeding and selection ; modifications in environ-
ment, especially changes in the water relation, and the effect of methods
of cultivation ; the effect of varying amounts of alkaline salts on rubber
deposition ; and, finally, pruning and other experiments designed to
test the possibility of increasing the amount of rubber bearing tissue
and also of increasing the percentage in the tissues.

XIV. CULTURAL REQUIREMENTS

In case Chrysothamnus is brought under cultivation, its cultural
requirements would need to be looked into very closely. At present,
our knowledge of these is based upon observations of the wild shrub
and upon a limited number of experiments in transplanting and in
growing from seed.

These plants are not exacting as to their climatic requirements.
The viriduliis and consimilis forms grow readily from the lowest to the
highest limits of the Upper Sonoran Life Zone and exceed these limits
slightly in both directions. Expressed in another way, they range from
fairly hot interior valleys, where the temperature runs to 110° F. in
summer, to over 7000 feet in the mountains where snow falls to a
depth of several feet and the temperature drops to — 20° F. in winter.
No difficulty has been experienced in growing these varieties in the
botanical garden at Berkeley, California, both from seed and from
root transplants, but it is doubtful if they will grow here as rapidly
or as large as they do under conditions of greater summer heat. Trans-
planted seedlings are also growing at the Citrus Experiment Station
at Riverside, but they are not at all vigorous. It is not known whether
this is due to the very unfavorable climatic conditions that prevailed
when thev were set out in the field or to some other factor. Moreover,
Riverside is in the Lower Sonoran Life Zone and therefore perhaps too
hot in summer for these Upper Sonoran plants. While the exact
geographic limits for the cultivation of Chrysothamnus will need to be
determined with some accuracy it seems reasonably safe to say that, as
far as climate is concerned, it may be grown in any of the larger in-
terior valleys of the West except perhaps those of the extreme north
and the very hot Lower Sonoran valleys of the extreme south.

'256 r)ii}'< r.<iihi of Califorxia PnhJlcatutUR in Botanij [Vou7

The iiioistui'c and soil ivMiiiii-cinciits have not l)('fii can'fully
determined. As indicated above tlie most promising- foi-ins grow
(111 valley bottoms where there is a moderate amonnt of moisture
and wliei-e the soil is moderately to stT^Jiigly alkaline. Salt-grass {Dis-
tichlis) is so eonnnon a concommitant of vivid nlns and consimilis that
its presence undoubtedly indicates favorable soil conditions. That the
l)lants can be grown with a minimum of i)recipitation is indicated by
tiieir abundance in places where the precipitation drops to below two
inches in some years, but in these places it flourishes only on valley
bottoms. The soil is always more or less sandy, but the plants grow
readily, or at least make a satisfactory start, in the heavy clay of the
botanical garden at Berkeley.

An ecologic consideration always to be kept in mind is that Chrijso-
thamnus is not a strong competitor among plants. It is easily crowded
out by other species, especially by Grease-wood if the alkali content
of the soil is high and by Sage-brush if the alkalinity is slight, so that
the area at present covered by it could be greatly extended through the
removal of these more aggressive shrubs. That this is true as regards
Sage-brush is attested by the experience of settlers who find that Rab-
bit brush (especially the gray forms, e.g., ynaphalodes) almost always
takes immediate possession of the ground after the removal of the
Sage-brush. This leads to the conclusion that the present occurrence
of Rabbit-brush does not by any means represent the total area suited
to its growth. It also indicates one rather simple method of extending
the growth of the shrub, namely by removing its competitors.

In case the cultivation of ('hrysothamnus is ever made a commercial
success, the first plantations will presumably be located where the best
varieties now grow wild. Aside from the assurance that such loca-
tions would provide the climatic, soil, and otlier requirements for suc-
cessful growth, this would enable the management to utilize the wild
shrub as a supplementary supply of crude rubber, although it would,
of course, carry a lower percentage than the improved cultivated
strains.

It is perhaps premature even to suggest districts where plantations
might be located but the mention of a few may be not without interest.
In Owens Valley, California, for example, it happens that the city of
Los Angeles owns nearly 200,000 acres of land, partly covered with
Clirijsothamnus and not at present utilized except to a very limited
extent for grazing i)uri)oses. It is estimated that at least one half of
this is suital)le to the growth of C hrysothamnus of the virkhilus form.

1919] Hall-Goodspced: Chnjsil 257

Similarly, large suitable areas now partly covered with Rabbit-brush
are located in Fish Lake Valley, Big Smoky Valley, Walker River
Valley, near Mono Lake, and elsewhere in Nevada and eastern Cali-
fornia. In Utah and in Colorado (especially the San Luis Valley)
there are extensive tracts of practically waste land where the varieties
consimilis and pinifolius are abundant. Finally, the Sacramento and
San Joaquin valleys of California include large acreages of practically
unused alkali lands where, unless the summer heat is too great,
Chrysothamnns could probably be grown.

Chrysothamnus is thus seen to be a hardy plant capable of growing
under a considerable range of climatic and soil conditions. It is ap-
parently best suited to the alkaline plains of the Great Basin, especially
if the soil is somewhat sandy. This fact, together with certain other
practical considerations, would indicate that if the plants are brought
under cultivation the first plantations should be located in these in-
terior districts, some of the most promising of which are those indicated
above. Zonal position should be taken into account since it seems un-
likely that the best sorts of Rabbit-brush can be grown to advantage
outside of the Upper Sonoran Life Zone.

XV. SUMMARY

(1) It was the object of this investigation to discover a supply
of rubber in native North American shrubs which might be used in
time of war, thus rendering the nation to some extent independent
of overseas importations of this substance. The studies were extended
to include a preliminary inquiry into the possibility of bringing the
plants under cultivation. Chrysothamnus or Rabbit-brush, was the
genus of shrubs especially studied.

(2) The work was fostered by the Committee on Scientific Research
of the State Council of Defense for California and by the University
of California. It was furthered through the aid of a considerable
number of correspondents.

(3) Chrysil is the name adopted for the particular kind of rubber
found in Chrysothamnus nmiseosus. It is a rubber of high grade and
vulcanizes without difficult}'.

(4) Rubber was found in two closely related genera of shrubs,
namely, Chrysothamnus and Haplopappus. Onh- the former yields

258 Universihj of California Publications in Botany [Vol.7

Clirysil. The most iiuportaiit spet'ies of Chrysothnninns is C. iiauseu-
SHS, under wliidi are rccofi'iiized twenty-two varieties. Twelve of
these vai'ieties have been examined and fulilier found in all of 1liem
although individual jilants may be devoid of rubber.

(51 ChnjsothdniHiis nau^eosus is a large shrub, the average weight
being from four to six pounds. Tt grows readily from seed and reaelies
nuitui'ity in from six to eight years.

(6) Tlie speeies is widely disti'ibutid in western Xortli America,
often iidialnting alkaline flats.

(7) The largest stands of Chrysil-bearing shi'ub are in ("oloi-ado,
Nevada, and T'tali. The ])lants with the highest rubber content are
from Nevada and California. The total amount of I'ubber pi-esent
in \\ild shrub is estimated at over 300,000,000 iiounds.

(8) The rubb(^i- oeeurs in the individual cells, and is not a latex-
rubber. Its place of occurrence in the i)lant has been rather definitely
located. Methods have been developed for its detection by micro-
scopical examination as well as by cbemical analysis. A'arious experi-
ments indicate that the chemical methods adopted are reasonably
accurate ; that great care must be exercised in the preparation of
samples for anal^'sis; and that samples do not deteriorate when stored
for periods of several months.

(9) The results of one hundred and eighty chemical analyses and
eighty microscopical examinations are tabulated. These indicate an
average of 2.83 per cent of rubber in the variety hololeucus, 2.69 per
cent in the variety pinifolius, 2.52 per cent in the variety viridulus,
1.97 per cent in the variety consimilis, and lower percentages in other
varieties. The low-percentage varieties are all comparatively uncom-
mon. The highest absolute percentage was found in a ])lant of
consimHis collected near Gerlach, Nevada, which analyzed 6.57 ])er cent
of pure rubber. The second-highest was a ])lant of viridulus from
Benton, California, wiiieh \ielded 5.56 per cent.

(10) Chrysil occurs in the plant in gi-eatest amount at about the
.soil line. In the root it is present in oidy the upper i)art. In young-
twigs and leaves it occurs in onl_\- snudl amounts. The richest tissues
are found in the cortex and the medullary rays, the former carrying
much moi-e tiuin the latter.

(11) The amount of rubber varies with the botaiueal variety: the
best varieties being those which inhabit alkaline soils.

1^11^1] Ilall-Goodsperd: Cliri/.sil 259

(12) 111 harvesting wild shrub the whole plaiit should be taken,
including four inches of root. Bulk may be reduced by removing
the twigs. Cultivated plants might be so cut as to leave the base of
the stem for regeneration,

(13) Because of the desirability of establishing the rubber-growing
industry- in the United States on waste lands Chrysothamnus and other
plants should be further investigated with a view to making their
cultivation a financial success.

(14) Its cultural requirements are such that Chrysothamnus could
be grown on many of the alkaline plains of the West without irriga-
tion; certain varieties endure winter temperature of — 20° F. ; others
would withstand summer temperatures obtaining anywhere in the
western states except possibly in the hottest valleys.

PLATE 18

Fig. 1. Photomicrograi)!! of a portiou of a cioss-scetion of a twig of the
current year — viridulits. Stained films in a few cells of the ]iith, absence of
rubber in rays and in cortical parenchyma, accumulation of rubber in the bud
intrusion. The chloreuchynia is heavily stained.

Fig. 2. Photograph of a portion of a cross-section of a five-year-old stem —

I'iriduJus.

a. The outer limits of the rubber-bearing cortex.

h. The tangential zone of maximum, cortical, rubber deposition.

Fig. 3. Photograph of a portiou of a three-year-old shoot — liiidulus. 2s'o
rubber in the pith.

c. The outer limits of the rubbear-bearing cortex.

Figs. 4 and .">. Photomicrographs showing nature of occinrence of rubber in
the cell — gnapJialodes.

L'<)0

UNIV. CALIF. PUBL. BOT. VOL. 7

[HALL AND GOODSPEED ] PLATE 18

PLATE 19

Fig. 1. PhotoniicrogTapli of a portiou of a c-ioss-section of a six-year cili I
stem — sprciosus — sliowiii<; the afcuimilatioii of iiihlier in the rays and their wide
i-ortical exteiision.s.

Fig. 2. Pliotograph of a portiou of a particularly thiu cross-section of a uiiie-
year-old stem — viridulu,^.

(I. Tlie outer limits of the significant rubber-bearing coi'tex.

[ 2<32 ]

UNIV. CALIF. PUBL. BOT. VOL. 7

[HALL AND GOODSPEED ] PLATE 19

PLATE 20

('liriit>ot]uim)ni-s nauncofius var. viriduJus. Plant H-3. To illustrate incthod
for increasing weight of rubber-bearing parts.

Fig. 1. Tlie wild plant as it appeared February l(i, 1918. Height 5i/^ feet,
spread 5 feet.

Fig. 2. The same plant after pruning, February 16, 1918. Greatest height 3^^
feet. The cut brant-hes are 276 in number, one to three years old, and average
one half inch thick.

Fig. 3. The same plant September 16, 1918. Height 4 feet 3 inches, spread
■4 feet 6 inches. The number of branches has increased to 640. This increase is
somewhat greater than in other plants subjected to the same treatment.

The black stake in figs. 1 and 2 is 40 cm. high. The short stake in all three
figures is 4 inches high.

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UNIVERSITY OF CALIFORNIA PUBLICATIONS

IN

BOTANY

Vol. 7, No. 8, pp. 265-278, 2 figures in text November 7, 1919

ITL THE OCCURRENCE OF RUBBER IN
CERTAIN WEST A^EERICAN SHRUBS

BY

Harvey Monroe Hall and Thomas Harper Goodspeed

CONTENTS

PAGE

I. Chriisothamniis (exclusive of C. nauseosu^) and HapJopappus 265

11. Reg-ional rlistribution of rubber in Haplopappus 274

III. Species iu which no ruV)ber is found 276

I. CHRYSOTHAMNUS (EXCLUSIVE OF C. NAU8E0SU8)

AND HAPLOPAPPUS
In a foregoing report attention has been paid almost exclusively to
rubber as it occurs in the numerous varieties of Chrysothamnus naiis-
cosns. In the course of the investigation we have also examined as
many of the botanically related species and genera as were readily
obtainable as well as some species of shrubs not closely related to
Chri/sothannius. The total number of species of woody plants exam-
ined is seventy-nine. As a result of these examinations rubber is now
known to occur in four other species of ChrijsotJiammis and in ten
species of Haplopappus. In no case, however, were we able to detect
rubber in any plant other than those showing a close botanical affinity
to Chrysothcnuius. Those in which rubber was found are indicated
below. In the tables the percentage of pure rubber is indicated by
the benzene extract. All percentages are based upon dry shrub except
where "x" indicates that no moisture determination was made.

Chrysothamnus turbinatus (IM. E. Jones) Rydb.

The general habit and appearance of this species is. very much
like that of certain varieties of C. nauseosus. Although it differs from
that species in what appear to be important technical characters, such
as the elongated, columnar involucre, it is not unlikely that further

J66

Universihj of California Publications in Botaii}) [yo^i..!

study will reduce it to a variety of nauseosus. The plants are two
to foui- feet liigii. i-ouud-topped, and very twigi^y. They grow on
saiid\- mounds of strongly alkaline clay fiats. The species has been
found only in Utah, and even there it is (piite I'ai-e. A single sample
(410) from the alkaline flats just east of Luntl analyzed 4.88 per cent
of rubbei". Since the species is so close to Haasrosiis botanically it is
probable that its rubber is also similar and should be referred to as
Chrysil.

Fig. 1. Cliriisotlxuniins h'rclifolin.s growing aiiiony the locks at JJeiiton Hot
Springs, California. J'laiits average 1% feet high.

Chrysothamnus teretifolius (Dur. and liilg.) Hall.
This is a low, broadly branched, woody plant, sometimes as much
as six feet high and fully as broad, but usually mueli smaller (cf.
fig. 1). Average plants will weigh one to three pounds. The lai-gest
we have seen had several trunks each eight inches thick and the whole
]>lant had an estimated weight of twenty pounds. Unlike the genuine
species of Ckrysothainnas this one has a veiy resinous herbage and
because of this it is placed in tlu^ genus Ericameria by some botanists.
It grows on gravelly or ston\ iiillsides, often in rocky canon bottoms.

1919] Hall-Goodspeed : Ruhher in West American Shrubs 267

and although it never forms pure stands of any great extent yet a
considerable supply could be assembled from some of the desert ranges.
The distribution of t( rctifolius is from the easterly slope of the Sierra
Nevada and Tehachapi mountains to central Nevada and possibly
Arizona. The largest stands we have seen are in the cafions on both
slopes of the White and Inyo ranges in eastern California ; there is
also a considerable quantity on the mountains to the west of Antelope
Yalley, California.

Table 1 — ^Chemical Axalyses

Place of Collection

Date of
Collection

Acetone

Extract

Per cent.

Benzene
Extract
Per cent.

45

Near Rosamond, Calif.

Oct. 26, 1917

5.46

2.80

436

Lida, Nev.

Sept. 10, 1918

2.85

1.67

440

West of Deep Spring Valley, Calif.

Sept. 12, 1918

2.22

2.48

575

Near Benton, Calif.

Dec. 23, 1918

4.61

4.51^

592

Near Victorville, Calif.

Microscopical

Dec. 25, 1918

EXAMIXATIONS

4.93

2.04>^

Place of Collection

Date of
Collection

Estimated
Amount

61

Near Bishop, Calif.

Nov. 1, 1917

Fair

67

Benton Hills, Calif.

Nov. 4, 1917

Fair

76

Near Candelaria, Nev.

Oct. 8, 1917

Poor

82

Near Benton, Calif.

Nov. 11, 1917

Good

83

Silver Canon, Inyo Mts., Calif.

Nov. 12, 1917

Fair

85

Near Owenyo, Inyo Co., Calif.

Nov. 14, 1917

Fair

Chrysothamnus paniculatus (Gray) Hall.

The plants of this species are on the average taller and less
spreading than C. tcretifoUus but they have a similarly glandular-
dotted foliage and the two are very close botauically. It grows on the
southerly part of the Mojave Desert and from Whitewater, on the
west side of the Colorado Desert, east to Arizona but it is nowhere
abundant. Six samples have been examined, with the following
results.

Table 2 — Chemic.\l Analyses

Place of Collection

579 North of Barstow, Calif.

580 North of Barstow, Calif.

Date of
Collection

Dec. 24, 1918

-Acetone Benzene

Extract Extract

Per cent. Per cent.

2.73 1.20^

Dec. 24, 1918 4.10 3.24^

Microscopical Ex.\mixatioxs

Date of Estimated

Place of Collection Collection Amount

96 Cabazon, Calif. Jan. 22, 1918 Fair

98 West of Whitewater, Calif. Jan. 22, 1918 Fair

99 North side of San Jacinto Mountain, Calif. Jan. 22, 1918 Fair
581 North of Barstow, Calif. Dec. 24, 1918 Poor

268 University of California Puhlicotions in Botanjj [Vol. 7

Chrysothamnus linifolius rji-oovic.

This is perliaps only a variety of ('. viscidi/l'iriis l)iit it (litlfcrs from
all otlu'i' forms of that cxceodinwly vai'iablc species in tiie taller and
more robust habit (the woody stems are sometimes eight feet hig^h), in
the broadly linear leaves, and in the greenish thiekened tips of the in-
volucral bracts. Its ecologie liabitat is also ditferent, since it grows
only in alkaline soil, whereas the forms of viscidi/fonis are confined to
well drained, non-alkaline slopes and benches. Tliis may account for
the occurrence of rubber in linifolius, and its absence in viscid iflonts as
well as in two othei- very closely related upland species, namely,
linmilis and piih( ndtis. This is in accord with what has been found in
C. nauseosHs, in which species, as noted elsewhere,^ the varieties carrj'-
ing the most rubber are all inhabitants of alkaline soils.

Only two samples of linifolius have been examined. One (4()())
from an alkaline flat near Grand Junction, Colorado, yielded on an-
alysis 1 per cent of rubber. The other (515), which was gathered at
Green River, Utah, was not analyzed. A microscopic examination failed
to detect any rubber in it.

Haplopappus nanus D. C. Eaton

The stems of this species are very low, one foot or less in height,
much gnarled and branched and with numerous short leafy twigs.
They have a thick, corky bark and a considerable portion of the wood
is dead in many of the plants. The foliage is deep green and (juite
resinous. Average plants will weigh about one pound l)ut it is usually
impossible to obtain the whole plant since the roots are tightly held by
the rocks in which they grow. The species inhabits rocky ledges and
outcroppings and may be expected on almost any of the mountaiii
ranges of the Great Basin area. It is known to grow in such situations
from Mono County, California, to southwestern Utah and north to
Washington. Wherever we have found it during tlie course of these
investigations it was growing in only limited quantities and there are
probably no considerable stands of it anywhei-e. (See fig. 2.)

The percentage of rubber in Haplopappus nanus is Ihc highest
kn(jwn for any native American shrub except Guayule, our analyses,
four in number, indicating that it runs from 6 to 10 per cent of the dry
weight of the entire plant. The (piality of the product would i)i'obably
be inferior to that of the viridulus form of Chrysothamnus because of
the large amount of resins present. A sample was worked out mechan-

iSee p. 245.

1919] Ilall-Goodspeed : Ruhher in West American Shrub!-

269

ically, separated in water, and found to be rather soft and dark colored.
It was not subjected to detailed examination. The recovery of any
considerable amount of rubber from the wild shrub is practically im-
possible because of the verj' limited known supply. It is conceivable
that under cultivation it might grow to good size and at the same time
retain its fairly high rubber content and that both the size of plant and
percentage of rubber could then be increased by selection or breeding,
but the outlook for this is not promising.

Six samples have been examined and five of them are reported on
below. A considerable variation in the rubber content is to be expected

since much depends upon the amount
of dead wood present in the sample.
Usually this is considerable since the
plants are very old and grow under
(^xtremely adverse conditions. The
analyses are for the whole plant
(after the obviously dead branches
had been removed) except that only
tli(^ upper portion of the root is in-
cluded. One other sample (438,
from near Deep Spring Valley, Cali-
fornia) was analyzed with the result
that the amount of the benzene ex-
tract was negligible. This is omitted
from the table since the plant was
not in a condition to permit of a
positive identification. It may be-
long to some other species or even
to another genus.

Fig. 2. Haplopappus nanus,

plaut no. 200. Beutoii Hills

California

illustrated. IV2 feet.

Total height as

Table 3 — Chemical Analyses

Place of Collection

Date of
Collection

Acetone

Extract

Per cent.

Benzene
Extract
Per cent

71

Near Benton, Calif.

Nov. 4, 1917

7.85*

9.46*

206

Near Benton, Calif.

Feb. 19, 1918

9.57

6.72

256

Shoshone Falls, Idaho

June 2.3, 1918

5.86

4.61

416

Caliente, Nev.

Sept. 6, 1918

4.49

8.42

Microscopical

Collection

Ex

\.\nNATIOX

Date of
Collection

Estimated
.Amount

75

Near Candelaria, Nev.

Oct. 8, 1917

Good

*Based on air-dried sample analyzed by Professor P. L. Hibbard; if perfectly dry the benzene extract
would probably be between 9.fi and 10 per cent.

1*70 [Inivet'silji of Calif orum riihlicdfitnis in Botdinj | \'ol. 7

Haplopappus cervinus Wats,
'riiis is a low sliful). scai'ccly a fool lii^li, with rcsiiious-punctat"'
leaves. It grows in the t'ootliill eaMons of westei'ii Utali and is espe-
cially eoininon along the east side of Salt Lake \'allev, where, however,
it does not grow in continuous belts. The ijuality of the rubber is
probably similai' to that of Haplopappus iuihks. of which rrrvinus is
pei-haps only a variety. The above-ground portions of four samples
submitted by Pi'ofessoi- Marcus E. Jones have been aiuil\ze(l with the
following result.

Table 4 — Chemical Analyses

Place of Collection
.'in Near Salt Lake, Utah
oo.") Parleys Canon, Utah
53()a Parleys Canon, Utah
537 Parleys Caiion, Utah

Haplopappus ericoides (Less.) H. & A.

This iieatlier-like shrub grows one to five feet bigh. The main stems
are one to three inches tbick at base, eitber erect or decumbent, and
emit numerous erect branchlets densely clotbed with sbort narrow
leaves, the whole plant thus resembling heather. The herbage is green
and resinous. The plants grow in sandy soil and are most abundant
as well as of greatest size in the sand dunes along the seashore, where
they sometimes form extensive colonies. The species extends geographi-
cally along the California coast from southern Los Angeles County to
San Francisco. The distribution of rubber in this plant is peculiar, if
a sample gathered in tlie sand dunes near Moss Beach, south of San
Francisco, is characteristic of the species. Only traces of rubber were
found in the stems of this plant but in the root, which was analyzed in
sections, the j)ercentage gradually increased until at a d('])t]i of two
feet it was 3.92.

Date of
Collection

Acetone
Extract
Per cent.

Benzene

Extract

Per cent.

Oct. 7, 1918

4.63

4.03

Dec. 22, 1918

5 . 58

2.28

Dec. 22, 1918

5.68

2 21

Dec. 22, 1918

5.54

2.90^^

1919] Hall-Goodspeed : Ruhher in West American Shruhs 271

Table 5—

-Ch

emical

Analyses

Place of
Collection

Date of
Collection

Acetone Benzene
Extract Extract
Per cent. Per cent.

Remarks

93

Duarte, Calif.

Jan.

20,

1918

5.49

0.49

Base of stem

149

Moss Beach, Calif.

May

23,

1918

9.17

3.92

End of root; 50

cm. below soil

line

Moss Beach, Calif.

May 23,

1918

7.47

3.11

Next 20 cm. up

root

Moss Beach, Calif.

May

23,

1918

5.16

1.67

Next 20 cm. up

root

Moss Beach, Calif.

May

23,

1918

5.49

0.58

10 cm. below soil

line

157

San Francisco,

Calif.

July

27,

1918

6.67

2.10

Root; piece 50 cm.
below soil line

San Francisco,

Calif.

July

27,

1918

5.14

0.75

Root; piece 10 cm.
below soil line

San Francisco,

Calif.

July

27,

1918

9.10

0.19

Base of stem

743

Duarte, Calif.

Jan.

^f

1919

3.72

0.13

Base of stem

Duarte, Calif.

Jan.

^}

1919

7.51

0.25

Twigs

Duarte, Calif.

Jan.

'■)

1919

3.95

0.90

Entire root

744

Duarte, Calif.

Jan.

■*•?

1919

3.11

0.27

Stem

745

Duarte, Calif.

Jan.

^j

1919

4.07

0.38

Stem

Duarte, Calif.

Jan.

•*■»

1919

4.81

1.97

Root

-

Duarte, Calif.

Jan.

^)

1919

3.32

0.19

Tops; 3 years old
and younger

Microscopical Examinations

Place of Collection

Date of
Collection

Estimated
Amount

95

Claremont, Calif.

Jan. 20,

, 1918

Poor

137

San Francisco,

CaUf.

Mar. 28,

, 1918

Poor

138

San Francisco,

Calif.

Mar. 28,

, 1918

Poor

156

San Francisco,

Calif.

July 27,

1918

Poor

Haplopappus Palmeri Gray.

In habit and general appearance this species is much like //.
ericoides except that the plants are usually smaller. It belongs to
southern California and is especially common on the plains and lower
footliills of the coastal slopes. It is absent from the sand dunes along
the coast. One sample (725) from Riverside was found to contain
0.91 per cent of rubber. Another (721), also from Riverside, and
one (201) from Colton were examined microscopically but no rubber
could be detected in either of them.

27'-'

I'tiirrralh/ of CaliforuM Publications in Botanjj [Vol.7

Haplopappus pinifolius Gray.
Til is soutluM'ii Califoi'iiia sijccics is much like //. (ricoides. One
c'olh'ftion (129) made near San Bernardino, was examined micro-
seopieally, and found to eontain hut a small amount of iiii>l)er. A
second samjile was taken in San FciMiatido \'all('y and fduiid to con-
tain 1.(11 pel' cent in the hasal portion of the stem.

Haplopappus monactis rJray.

This is a good sized shruh, commonly two or three feet high,
sometimes six feet high and with a trunk diameter of foni- inches;
in one case there were three principal branches, each one to tliree
inclu's thick. The weight is estimated at five pounds foi- average
plants; thirty pounds for exceptionally large ones. The species is
most abundant around the westerly extensions of the Mojave Desert
but it grows at many places along the desert borders and east into
southern Nevada; also to a very limited extent on the coastal slope
of southern California. One large shrub (90) was collected near
Elizabeth Lake and examined microscopically. It appeared to con-
tain a fair amount of rubber. A much smaller one (713) from near
Hesperia Avas analyzed and found to contain only 0.38 per cent of
rultber.

Haplopappus laricifolius Gray.

Very close botanically to //. nianacti'S, this species is also like it
in habit, general appearance, and rid)ber content. It grows in south-
ern Arizona, but apparently it is not abundant. Five samples have
been examined with the result indicated below.

Table (i — Ohe\iic.\l A>f\LYSEs

Place <if Collection

760 Near Tucson, Ariz.
783* Santa Catalina Mts., Ariz.
783t Santa Catalina Mts., Ariz.
786* Tucson Mts., Ariz.
78fit Tucson Mts., Ariz.

* Stem,
i' Root.

Date of
Collection

.■\cetone
Extract
Per cent.

Benzene
Pjxtract
Per cent.

Feb. 18, 191!)

;5.44

.5 16

Mar. 16, 1919

.5.44

2.32

Mar. 16, 1919

3.62

3.37

Mar. 23, 1919

3.94

2.01

Mar. 2:^, 1919

3 33

2. 87

MiCRO.SCOI'ICAI. KXAMIN'.VTION

Place of Collection

1.58 Near Tucson, .\ri/..

Date of
Collection

.Julv 27, 191.S

Estimated
.Vniount

Fair

1919] Hall-Goodspeed : Ruhher i)i West American Shrubs 273

Haplopappus arborescens (Gray)-
Althoiigh this is one of the largest species of the genus it is not
really arborescent, as its specific name would seem to indicate. How-
ever, the woody stems are often several inches thick at the base and
the shrubs are commonly three to five feet high. The species is com-
mon in the Coast Ranges of middle California and occurs also in the
foothills of the Sierra Nevada. Two specimens (151, 152), both from
one station on ]\It. Tamalpais, showed only traces of rubber on micro-
scopical examination.

Haplopappus brachylepis (Crray)^'
The geographic distribution of H. hrachylepiji is restricted to a
small area in southwestern San Diego County, California, and to
northern Lower California. The species is a large shrub, commonly
three to six feet high, with brittle stems. It grows in the arid chapar-
ral belt, usually on stony hillsides. Four samples taken in San Diego
Countj^ near the borders of Lower California, were examined with
results as indicated below.

Table 7 — Chemical Analyses

Place of Collection

Date of
Collection

Acetone Benzene
Extract Extract
Per cent. Per cent.

728

El Campo,

Calif.

Dec. 29, 1918

13.18 0.81

730

El Campo,

Calif.

Microscopical

Dec. 29, 1918
Examinations

10.69 0.16

Place of Collection

Date of
Collection

E.stimated
Amount

727

El Campo,

Cahf.

Dec. 27, 1918

Nothing

729

El Campo,

Calif.

Dec. 29, 1918

Traces

Haplopappus linearifolius DC.

This is an erect shrub two to live feet high, with stems commonly
one to four inches thick, and resinous herbage. It grows from Lake
County, in the Coast ranges, south into Lower California and east
to southern Utah and Arizona. It is perhaps most plentiful around
the borders of Antelope Valley, California. There are two forms.
Typical linearifolius grows in the Coast ranges. The other form,
distinguished by its shorter leaves and smaller heads, replaces it in
the desert country and is known as var. inferior (Coville) M. E. Jones.

- Ha i>h>i)(i Pints arborescens (Gray) H. M. Hall, comb. iiov. Jyi'/fo.sv/'w arhor-
iHccns Gray, Bot. Mex. Bound. (1859), p. 79.

'^Haplopappus hrachiilepis (Gray) H. M. Hall, comb. uov. Bigelovia brachy-
lepis Gray, Bot. Calif., vol. 1 (1876), p. 614.

•J74

I'nivcrsil ij <ij' California Puhllcdt iaiis in liotdiii/ [Vol. 7

111 tlir followinj^' labuhit ion of cxaniiimt ions, only nos. 492 and 493
fi'|)i-cscn1 1\j)ical liii((irifolii(s : tlic remainder are of llio interior
\i\y\v\y.

'IauU'; S — CiiE.\n(Ai, Axai.vsks

Pl.ice of Collection

492 Soutli of Antioch, Cnlif.

493 South of .Vntioch, Calif.

701 Mojave Desert, Calif.

702 Mojave Desert, Calif.

703 Mojave Desert, Calif.
714 Near Hesperia, Calif.

Dafr of
Collection

Acetone

Kxtract

Per cent.

lienzene

Kxtract

Per cent.

Oet.

12, 191S

2.47

0.30

Oct.

12, 191S

1 . 53

(J 03

Dec.

25, 1918

3.15

1.21

Dec.

25, 1918

3.32

1 . 32

Dec.

25, 1918

3 . 13

1,19

Dec.

2G, 1918

2.78

0.45

.MicKo.scopicAL Examination

Place of Collection
89 Del Sur, .\nt elope \^>>lley, (^alif.

Date of
Collection

.Ian. 19, 1918

Estimated
.Amount

Poor

II. REGIONAL DISTRIBUTION OF RUBBER IN

HAPLOPAPFVS

We liave sliown that Chrj^sil is peculiar to tlie ])art of the plant
above ground, analyses of root and stem indieatinji- that below a point
approximateh' 10 cm. beneath the soil line rubber occurs in negligible
(quantities even if the parts above are relatively rich in this substance.
In tlie ease of at least two species of Haplopappus a different distri-
bution obtains as is indicated by the results given in the following
table.

1919] Hall-Goodspeed : Ruhher in West American Shrubs 275

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276 Unii'( rsitij of Culij'ornia I'liblicdiioiis in Botany | \<h.. 7

It seems eleai* from llic results iiichuli'd in tlic aliove table that
tlic root areas of the two species of Ilaplopappus examined contain
ai)i))'ecialih' (|uaiitities of i-ul)l)er. In the case of //. ericoides the
parts above tin' soil line are strikingly detieient in rubber as compared
with the roots. Indeed, in the extremities of the root the percentage
is often highest and decreases as one passes upward towards the soil
line. The regional distribution in ericoides is, thus, the reverse of
that found in the various varieties of Chrysothannitis.

In //. tuiiiKS, on the other hand, stem and root aj)pear to Ix-ai'
ecjual (luantities of rubber. The evidence here is o])vi()usly fragmen-
tary but the above statement holds rather strictly for 206, and in the
case of 256 the stem probably bears a maximum (juantity for this
plant and it might be assumed that the root, if anal^'zed, would show
an approximately equal amount.

It appears, then, that three types of regional distribution of con-
tained rubber are shown by the plants which we have examined in
the course of the entire investigation. In the first place, the maxinnnn
quantity is borne by the stem. This is peculiar to the distribution of
Chr^-sil. In the second place, root and stem bear equal amounts.
This condition obtains in Haplopappiis nauus. Finally, we have the
case in which the root carries almost the entire amount of rubber
borne by the plant, //. ericoides. This seems a rather remarkable
situation and one which would repay further investigation. In par-
ticular it would be important to examine in detail the distribution
of rubber in the other species of Ilaplopappus not investigated in
this particular connection by us.

111. SPECIES IN WHICH NO KUBBER WAS FOUND

In the course of the investigation reported on in the foregoing
pages a miscellaneous collection of plants was made as opportunity
offered and these have been examined histologically for rubber con-
tent wdth the result that rubber was located in certain cases but only
in species of Chrysoihammis or of related genera. This occurrence
has been noted in the preceding list but it now remains to enumerate
those species in which no rubber could be detected with certainty.
This list is given as it may be of some service in case a more extensive
survey is undertaken. As in the preceding report, the numbers refer
to field notes and have been assigned merely for convenience of refer-
ence. Tin- locality imiitioned in each case is the jilace of collection.

1919] Hall-Goodspeed : Riihher in West American Shrubs 277

I. COMPOSITAE

700. Acamptopappus sphaerocephalus. Near Hesperia, California.

491. Artemisia californica. Antioeh, California.

J 9. Artemisia filifolia. Denver, Colorado.

460. Artemisia Bothroclcii. Tioga Pass, California.

8. Artemisia trideiitata. Truckee, California.
589. Baccharis glutinosa. Barstow, California.
747. Bacoharis pilulari-s. Near Ventura, California.

97. Baccliaris sergiloides. Near Banning, California.
7.51. Bebbia juncea. Palm Springs, California.

595. BricleUia atractyloides. Near Vietorville, California.

596. Brickellia desertorum. Near Vietorville, California.
513a. Brickellia microphyUa. Near Salt Lake, Utah,
409. Chrysothamnu-s Greenei. Lund, Utah.

285. Clirysothamnus Hoivardi. Near Walsenburg; also no. 287, Canon City,

and no. 292, Villa Grove; all in Colorado.
458. Chrysothamnus nevadensis. Mono Mills, California.
18. Chrysothamnus Parryi. Teller Lake, Colorado.

9. Chrysothamnus humilis. Truekee Valley, California.

14. Chrysothamnus puberulus. Pyramid Lake, Nevada; also 23, Tonopah,
Nevada; also 28, 29, Keno, Nevada; and 259, southeastern Idaho.
7. Chrysothamnus viscidiflorus var. tortifoUxis. Truckee Valley, Califor-
nia; also 30, Reno, Nevada.
746. Corethrogyne filaginifolia. Near Ventura, California.

763. Dysodia porophylloides. Palm Springs, California.
736. Encelia californica. Near San Diego, California.

598. Encelia farinosa. Near Vietorville, California; also 722, Riverside,
California.

68. Ericameria cuneata spatulaia. Benton, Mono County, California; also
near Vietorville, California.

86. Franseria dumosa. Chvenyo, California; also 587, Barstow, California.

594. GnaphaUum bicolor. Near Vietorville, California.

490. Gutierrezia californica. Antioeh, California.

46. Guiterrezia lucida. Rosamond, California; also 586, Barstow, California.

558. Gutierrezia Sarothrae. Near Reno, Nevada.

38. Eaplopappus Bloomeri. Truekee, California.

273. Haplopappus Fremonti. Laramie, Wyoming.

735. Hazardin squarrosa. El Campo, San Diego County, California.

764. Mofmeisteria pluriseta. Palm Springs, California.

73. Hymenoclea salsola. Mina, Nevada; also 585, Barstow, California.

761. Isocoma cor onopif alia. Near Tucson, Arizona.

160. Isocoma Harttcegi. Near Tucson, Arizona; also 762, from Tucson.

427. Isocoma veneta acradenia. Las Vegas, Nevada; also 599, Vietorville,
California.

94. Lepidospartum squamatum. Duarte, California.

242. Ficradenia Lemmoni. Near Weed, California.

560. Tetradymia cancscens. Near Reno, Nevada; also 433, Goldfield, Nevada.

278 Vnivcrsilij of Calif orn'm J'ublicatioiis in Botany |\«>i,. 7

II. MISCELLANEOUS FAMILIES

741. Astraffohts leucopsis (Leguminosae). Near San Diego, Califoruia.
42.J. AtripJex cancsccns (Chrnopoduiccac). Las Vegas, Nevada.

24. Atrli>lcx coiifcrtifolia {Chenopodiocccn). Tonopah, Nevada; also 582,

Barstow, California.

740. Cncoridium dumosum (Butaccae). Near San Diego, California.

31. Ephedra nevadcnsi^ {Gnctaceae). Eeno, Nevada.

707. Eriodicti/on calif ornicum (HydrophyUaccae) . Mojave Desert, California.

432. Eriogonum fasciculatum var. (Polygomiceae). Goldfield, Nevada.

494. Erlogonum nwlum (Polygonaccac). Mt. Diablo, California.

74. Eurotia lanata (Chcnopodiaceac). Mina, Nevada.

734. Frascra Parryi (Gcntiaimceae). El Campo, San Diego County, Calif oi'-

nia.

420. Glycyrrhiza Icpidota (Leguminosae). Las Vegas, Nevada.

;j97. Isomeris arborea {Capparidaccuc). Near Victorville, California.

584. Larrea divaricata {Zygophyllaceae) . Near Barstow, Califoruia.

429. Lepidium Fremonti (Cruciferae). Indian Spring, Nevada.

583. Lycium Cooperi (Sokm-aceae). Near Barstow, California.

422. Olneya tesota {Leguminosae). Las Vegas, Nevada.

423. Prosopis juliflora (Leguminosae) . Las Vegas, Nevada.

424. Prosopis pubescens (Leguminosae). Las Vegas, Nevada.

742. Phamnius crocea (Phamnaceae). Near San Diego, California.
739. Ehu^ integrifolia (Anacardiaceae) . Near San Diego, Califoruia.

732. Plius ovata (Bliamnaceae) . El Campo, San Diego County, Califoruia.
420. Ehus trilobata (Anacardiaceae). Las Vegas, Nevada; also 733, El

Campo, California.
15. Sarcobatus vermiculatiis (Chenopodiaccae). Near Pyramid Lake,

Nevada; also 33, same locality.
738. Simmondsia calif or iiica (Buxaceae). Near San Diego, California.

UNIVERSITY OF CALIFORNIA PUBLICATIONS

IN

BOTANY

Vol. 7, No. 9, pp. 279-324, plates 21-31 July 8, 1920

PHYCOLOGICAL CONTRIBUTIONS library

BY

WILLIAM ALBERT SETCHELL and NATHANIEL LYON GAEDNER

Hormiscia doliifera sp. nov.

Filamentis 3-4 cm. longis, dum juvenis prope cylindicis, ad basim
tantummodo attenuatis, rhizoideis extramatricalibiis e segmentis panels
infernis oriundis adjunctis, atroviridibus ; segmentis fertilibus 80-130ju
usque ad 184/x diam., 0.75-1.25-plo diam., longioribus, doliiformibus,
parietibus tenuibus, 5-7 /x erassis, hyalinis, honiogeneis ; ehromatophoris
tenuibus fenestratis taeniaeformibus, parietalibus, pyrenoideis parvis
numerosis.

Filaments 3-4 cm. long, nearly cylindrical throughout when young,
tapering only at the base, attached by extramatrical rhizoids from a
few of the lower segments ; color dark green ; fertile segments 80-1 30fi,
up to 184|Li diam., 0.75-1.25 times as long, doliiform, with thin, 5-7ju,
thick, hyaline, homogeneous walls ; chromatophore a thin fenestrate
parietal band, with numerous small pyrenoids.

Growing on rocks in the upper littoral belt. South side of the
Golden Gate, San Francisco, California. July, 1918. Type no. 4462,
Gardner.

Hormiscia doliifera resembles most closely Urospora Hartzii Eosen-
vinge (1893, p. 922) and V. in cmssaia Kjellman (1897, p. 7). From
each of these species it differs in having filaments of larger diameter,
in having more uniformly swollen, sometimes almost spherical, fertile
segments, and in having the segments more nearly uniform in length,
averaging a little less than quadrate. From Z7. incrassata it differs
also in its strictly extramatrical rhizoids. It approaches also the
little known V. crassa Rosenv., but its segments seem never so short
as represented for that species. It is much too slender for Hormiscia
collahens (i.e., up to 450/j,), as indicated by Batters (1894, p. 114).
CNj The filaments are decidedly larger than any dimensions given for

280 University of California Puhlications in Botany [Vol. 7

If. pcnicilUformis (Rotli) Fries and the chromatophore is thinner,
usually more coarsely reticulate, and with many more and much
smaller pyrenoids.

Spongomorpha Mertensii (Rupr.) comb. nov.

Conferva Mirteiisii Ruproclit, Tange, 18.51, p. 40.3.
Conferva viminea Ruprecht, lor. cit. (fide Yendo, 1916, p. 246).
Cladophora Mertensii De-Toni, Syll. Alg., vol. 1, 1889, p. .317.
Cladophora viminea De-Toni, loc. cit., p. 318.

Spongomorpfm arcia var. limiianea Collins, Green Alg. N. A., Suppl. I, 1912,
! p. 97, in Collins, Holden and Setchell, Phyc. Bor.-Amer. (Exsicc), no.

. 1736.

Acrosiphonia Mertensii (Rupr.) Yendo, Notes on algae new to Japan V,
1916, p. 246.

It seems best to follow Yendo {loc. cit.) in considering the Conferva
Mertensii Rupr. distinct, and we refer here, for the present at least,
the Spongomorpha arcta var. limitanca Collins. The species, as we
understand it, has no spiny branches or branchlets, but usually shows
circinate, simph' or compound, branchlets. Since we prefer Spongo-
morpha to Acrosiphonia, we are compelled to establish the new combi-
nation given above.

Capsosiphon fulvescens (Ag.) comb. nov.

Viva fulvescens Agardh, Sp. Alg., 1822, p. 420.

Ilea fulvescens J. Agardh, Till Alg. Syst., pt. 3, 1882, p. 115, pi. 4, figs.

95-99.
Viva aureola Agardh, Icon. Alg. Eur., 1829, pi. 29.
Capsosiphon aureolum Gobi, Ber. Alg. Forsch. im Finn. Meerbus., 1877, etc.,

1879, p. 88.

We feel compelled to accept the dictum of J. G. Agardh {loc cit.)
that the Ulva fulvescens of C. A. Agardh is only a younger condition
of Ulva aureola C. A. Agardh. Since it seems to us, also, that the
proper generic designation is Capsosiphon, the new combination ])ro-
posed above is rendered necessary.

Enteromorpha gToenlandica (J. Ag.) comb. nov.

Monosiroma fjrocnlandicum J. Agardh, Till Alg. Syst., i)t. IN, 1SS2, p. 107,
pi. 3, figs. 80-83; Collins, Green Alg. N. A.. 1909. p. 208; Saunders,
Alg. Harriinan Exp., 1901, p. 410; Setchell iiii.i CanliuT. Alg. N. W.
Anier., 1903, p. 208.

1920] Setchell-Gardner : Phycological Contributions 281

Enteromorpha groenlandica has always been puzzling as to its
proper placing. While technically it may seem to belong to the genus
Monostroma, under which it was originally described, more properly
than to any other genus of the Ulvaceae, yet its slender filiform habit
certainly more closely resembles that of some species of Enteromorpha.
From Enteromorpha, however, it differs in not having its cells set
sufficiently closely together to be parenchymatous in appearance. It
is at first solid, becoming hollow only late, but never rupturing longi-
tudinally and opening out into a membrane as do the characteristic
species of Monostroma. Certain species of Enteromorpha show a
tendency towards abundance of intercellular jelly at times, while
certain species of Monostroma are parenchymatous. It seems best to
us, therefore, to transfer this species to Enteromorpha.

Monostroma areolatum sp. nov.

Plate 30 and plate 31, figure 2

Thallo delicatissimo, lubrico, 20-35 cm. alto, sessili, juveno saccato,
mox fisso et lobos late ovatos aut obovatos, undulatos, plicatos et
crispatos formanti, dilute viridi ; membrana distincte minuteque areo-
lata, 9-12/A crassa; cellulis rotunde angulatis, 6-7/a crassis in sectione
transversali subsphericis, in areolis quibusque aggregatis.

Frond very delicate, lubricous, 20-35 cm. high, sessile, saccate when
young, soon splitting and forming numerous, long, broadly ovate, or
obovate, undulate, plicate and crisped lobes, pale green ; membrane
distinctly and finely areolate, 9-12/^ thick; cells with rounded angles,
6-7/x diam., subspherical in cross-section, grouped within each areole.

Growing on Zostera in quiet waters. Sitka, Alaska. Type no.
3924, Gardner.

This species of Monostroma is exceedingly beautiful and is among
the most delicate and flaccid of the genus. The frond remains saccate
for only a brief period, attaining a height of only a millimeter or two.
The sack then breaks and the membrane spreads out at once, early
developing small lobes. Finally a few primary lobes are established
and these develop numerous secondary lobes. The gro\\'th on the
whole margin greatly exceeds that of the interior, and this results in
the production of a great number of folds, making the margin very
much crisped. In the thickness of the frond and shape of the cells
M. areolatum closelj^ approximates M. zostericola Tilden. The cells
of the latter are, however, more angular and more closely placed and
the frond is not divided into areolae. There is a marked difference

282 University of California Puhlicutions in Botany [Vol. 7

in the size of these two species, jis well as in their methods of develop-
ment. M. zostericola is diminutive, remains saccate for some time, and
splits longitudinally, forming several lobes broadening outward. M.
areolaium verj'^ closely resembles the genus Prasiola in the grouping
of the cells as seen in surface view.

Ulva stenophylla sp. nov.

I'late 26, figure 2, and plate 29

Thallo siiii])lici, lineari-lanceolato, ad basim in stipitem brevissi-
mum, coinplanatum, euneatum longe attenuato, 5-8 dm. longo. 5-10
cm. lato. medio piano, marginibus undulatis ; raembrana 60-1 10/x
crasso, cellulis e superticie quadratis, 14— 20/<, diam. in sectione trans-
versali, 1.5-2-plo diam. longioribus; chromatoi)horis tenuibus, parie-
talibus, omnino aut i)artim cellulam tegentibus ; pyrenoidibus nuUis.

Frond simple, linear-lanceolate, tapering abruptly at the base to
a very short, flattened, euneate stipe, 5-8 dm. high, 5-10 cm. wide,
jdane in the middle with undulate margins; membrane 60-110/x thick;
cells squarish in surface view, 14-20/1, diam., 1.5-2 times as long as
tlie diameter in section, chromatophore a thin, parietal layer, covering
a part or the whole of the cell ; pyrenoids absent.

Growing on rocks in the lower littoral belt. Central California.
Type no. 5445 (Herb. Univ. Calif., no. 98511), Setchell.

The plants described under this name are quite distinct from the
other species of Ulva, in shape, in texture and in anatomical details.
They are dark green, tough and harsh to the touch. The usually
simple, long, lanceolate shape serves to distinguish them from other
species at a glance.

Ulva vexata sp. nov.

Plate 22, figures 4-7

. Thallo parvo, simplici, rigido, lineari ad oblanceolato aut spatulato,
piano aut parce undulato, plus minusve bullato, basi cuneato stipite
parvo et solido, 1-3 cm. longo, 3-10 mm. lato, atroviridi, siccitate atro;
membrana 45-55/1, usque ad lOO/x crassa, cellulis verticaliter elongatis,
11-15/1 usque ad 18/t longis, 3.5-5/t crassis, parietibus crassis et e
superticie angulos obtusos ostendentibus ; chromatophoris cellulas im-
l)lentibus: ))yrenoi(libus nullis.

Fi-oiid small, unbranehed, rigid, linear to ol)hinceolate or si)atulate,
plane or slightly undulate, more or less buUate with euneate base and
small, solid stipe, 1-3 cm. long, 3-10 mm. wide, dark green, black on
drying; membrane 45-55/i, up to lOO/i thick, cells vertically elongated,
11-15/1, U]) to 18/t long, 3.5-5/x wide, with thick walls and very blunt
angles in surface view ; chromatophore filling the cell, pyrenoids absent.

1920] Setchell-Gardner : Phycological Contributions 283

Growing on rocks along high-tide level. In the vicinity of San
Francisco, California. Type no. 4444, Gardner.

TJlva califarnica Reed, Two ascomycetous fungi, etc., 1902, p. 149
(not of Wille).

Ulva vcxata has been observed only in the vicinity of San Fran-
cisco, as mentioned above, where it grows in considerable profusion.
It seems quite probable that it may be much more widely distributed
both north and south of San Francisco. It might be suspected of
being a malformation due to the parasite always found more or less
infesting it, but the size and proportions of the cells of the less para-
sitized portions seem to mark it as a distinct species.

Ulva angusta sp. nov.

Plate 27 and plate 31, figure 1

Thallo simplici aut rarissime lobato, lanceolato aut oblanceolato.
8-15 em. longo, 0.5-1.5 cm. lato, 35-45/i, (interdum prope SS/x) crasso,
basi aut longe abrupto ad stipitem delicatum solidumque attenuato,
disco affixo, dilute viridi, marginibus fere planis usque ad erispatissi-
mis ; cellulis e superficie 3-6 lateralibus angulis rotundis, 5-12fx diam.
seetione transversali quadratis ad 1.5-plo longioribus, angulis rotundis;
chromatophoris dimidium externum cellularum implentibus; pyrenoi-
dibus singulis.

Frond simple or very rarely lobed, lanceolate to oblanceolate, 8-15
cm. long, 0.5-1.5 cm. wide, 35-45/i, thick (occasionally about 53,u),
tapering either gradually or abruptly at the base to a delicate, solid
stipe with discoid holdfast, color of fronds pale green, margins varying
from almost plane to very much crisped ; cells in surface view 3-6
sided, with rounded angles, 5-12/x, diam. in section, quadrate to one
and a half times longer than broad, with rounded angles ; chromato-
phore filling the outer half of the cell ; pyrenoid single.

Growing in shallow pools along high-tide level. Moss Beach, San
Mateo County, California, April, 1919. Type no. 4430, Gardner.

"We find at several places along the coast of central California a
rather short and narrow Ulva which does not seem to belong to any
of the hitherto described species. We have felt compelled, therefore,
to give it a name. It resembles the Phycoseris lapathifolia of Kuetzing
(1856, pi. 25), but is shorter and narrower. It also resembles, even
more closely, Kuetzing 's figure of Phycoseris Lima (1856, pi. 16, fig. 1 ) ,
but is a smaller plant than that also. The short, flattened stipe is solid.
The narrow blade varies from plane to undulate or even crisply
ruffled on the margins. The cells are oblong or rounded in section,
each provided with a more or less distinct wall. Although we have
only recently become acquainted with it, this seems to be a vernal
species. It has been observed in fertile condition in April.

284 University of California ruhlicutions in Botany [Vou7

Ulva lobata (Kuctz.) comb. iiov.

rhiicoscris lobata Kuetzing, Spec. Alg., 1849, ip. 477, Tab. Phyc, vol. 6,

1856, p. 10, pi. 27.
riva fasciata f. lobata Setehell, in Collins, Holiion and Setehell, I'liyc

Bor.-Anier. (Exsicc), no. 863; Collins, Green Alg. N. A., 1909, p. 216.

Among the Ulvac of tlie Calif orniaii coast is one of moderate size
(up to 30 cm. or more long, and up to 15 cm. or more broad) which
is distinct in general appearance. It is attenuate at the crisped base,
broadening above and usually lobed or divided into several broad
divisions. The margins are either plane or slightly undulate. Like
U. expansa it is thicker iii the center with palisade-like cells (in
section) and thinner on the margins where the cells are nearly square
(in section). It bears a striking likeness in every way to Kuetzing's
figure (1856, pi. 27) of his Phycoseris lohata from Chili. We have,
therefore, referred it to his species with some doubt.

Ulva lohata belongs to the same group of species as U. expansa,
but is generally firmer in substance, slightly thicker, never reaches a
great size, and is less deeply or conspicuously ruffled. It is well
represented by the specimens distributed in the Phycotheca Boreali-
Americana (under no. 863).

The most typical plants are those of the central Californian coast
(San Francisco to Monterey). We have referred here, also, one plant
from southern California, but with some doubt.

Ulva expansa (Setehell) comb. nov.

Frond ample, pale green, orbicular or broadly elongated, margin
deeply ruffled ; frond 60-70/* thick in the middle, 38-45/* on the mar-
gins; cells, in section, vertically elongated in the middle of the frond
(up to 28-30/* long, 10-12/* wide), nearly square in the margins.

Growing on rocks in the lower littoral belt. Puget Sound, Wash-
ington, to Mexico (La Paz).

Ulva fasciata f. expansa Setehell, in Collins, Ilolden and Setehell,
Phyc. Bor.-Araer. (Exsicc), no. LXXVII ; Collins, Green Alg. N. A.,
1009, p. 216.

We find along the coast of central California a broad species of
Ulva, often also long, something like Ulva latissima in ai)pearance,
yet of a more vivid green color, thicker in the center of the frond and
with distinct, broad, ruffled margins. The cells of the thicker center
of the frond are distinctly palisade-like in section, while on the thinner

1920] Seichell-Gardner : Phycological Contrihutians 285

margins they are nearly square. A younger specimen of this plant
was distributed by one of us as Ulva fasciata f. expansa (Phyc. Bor.-
Amer., no. LXXVII), but it has seemed, on further study, to belong
neither to Ulva fasciata Delile nor to the Ulva fasciata f. taeniata also
distributed by one of us (Phyc. Bor.-Amer., no. 809), but described
later on in this account as Ulva taeniata. We, therefore, describe it
as an independent species under the name of Ulva expansa.

Ulva expansa, so far as we have observed it, remains attached only
for a short time. It soon becomes free and floats or drifts, increasing
in size, becoming at times at least 3 M. long and varying in width
from 18 cm. to 75 cm. In form and structure it differs from Ulva
latissima and from all the other species of Ulva of our coasts. It
comes nearest to Ulva fencstrata, as we have described that species,
but is little, if at all, perforate. Plants of what appears to be the
same species have been found in the Puget Sound region and Howe
(1911, p. 490) is inclined to credit here some from La Paz, Mexico.

Ulva dactylifera sp. nov.

Plate 26, figure 1

Thallo sessili aut stipite brevi suffulto; parte basali obovata aut
reniformi, crispatissimo, 2-4 cm. alto, e margine superiore 1-6 lobos
aut lacinias, simplices aut sparse ramosos et costam definitam osten-
dentes et marginibus crispatissimis instructos producente, 5-15 cm.
high, 0.5-1.5 cm. wide; membrana basali ad marginem 50/x, ad medium
usque ad 100/x crassa, cellulis e superficie 16-20/x diam. sectione trans-
versali quadratis usque ad 2-plo diam, longioribus; membrana laci-
niarum marginibus 40-50/* crassa, medio usque ad 190/a crassa, cellulis
e superficie 12-16/t diam., sectione transversali quadratis usque ad
5-plo diam. longiores; chromatophoris dimidium externum cellularum
implentibus.

Frond sessile or with a very short stipe ; basal portion orbicular
or reniform, much crisped. 2-4 cm. high, giving rise from the upper
margin to 1-6 lanceolate, simple or occasionally branched lobes or
laeiniae with plane midrib, and much crisped margins, 5-15 cm. high,
0.5-1.5 em. wide ; membrane of basal portion 50/a thick at the margin,
up to 100/x thick in the middle, with cells 16-20/i. diam. in the surface
view, quadrate to 2 times as long as wide in section, membrane of the
laeiniae 40-50/t thick on margin, up to 190/i thick in the middle, with
cells 12-16/1 diam. in surface view, quadrate to 5 times as long as wide
in section ; chromatophore filling the outer half of the cell.

On exposed rocks, uppermost littoral belt. Southern California
to Mexico (San Roque?). Type no. 3988, Gardner.

"We have along the Californian coast two species related to Ulva
fasciata, neither of which seems to be exactly like the Mediterranean

286 University of California Publications in Botany [Vol.7

species. Both are characterized by long, narrow fronds or laciniae,
much thicker along the middle and with thinner, very much crisped
margins. One of these, Ulva dactylifera, possesses a comparatively
broad, though sliort, undivided basal portion from which arise the
several, narrow, elongated, crisped laciniae. Neither the basal portion
nor the laciniae show distinctly toothed margins. The other species,
Ulva taeniata, is either simple, long, slender, plane and dentate below,
but with crisped margins above, or divided to the very disk itself into
two or three such divisions. The "midrib" portions differ slightly
in thickness in tlie two species and the cells of the "midribs" differ
in ])roportions.

Ulva dactylifcra has been distributed under no. 221 & (sub "Ulva
fasciata") of tlie Phycotheca Boreali-Araericana. Unfortunately the
plants under this number are not uniform. We have examined no.
2216 in two copies. In one the plant is certainly, although not
typically, U. dactylifcra. In the other it seems rather to be a form
of Ulva Lactuca.

Ulva dactylifcra is nearest to U. fasciata f. costata Howe (1914,
p. 20, pis. 1, 2, figs. 10-23), but differs as to the basal portion, thick-
ness, and possibly also in proportions of cells. It differs from U.
fasciata Delile, so far as descriptions and figures indicate, in branch-
ing, in ruffling, and probably in thickness. It is a very much thinner
plant than U. ncmatoidca Bory, judging from the dimensions given
by Bornet (1892, p. 36 or 196).

Ulva taeniata (Setchell) comb. nov.

Plate 28

Frond elongated, up to 1 to 2 M. long, simple or split to the very
base into long, narrow segments, plane below and coarsely dentate,
densely crisped and ruffled on the margins above, with a plane, thiekei-
midrib ; membrane up to 140/x thick as to the midrib, and down to 40/i
thick on the margins; cells of the midrib vertically elongated, in
section, up to two and one-half times as high as broad, l)ut becoming
nearly square towards the margins.

On rocks in the lowermost littoral or upper sublittoral belts.
Central California (Tomales Bay to Monterey.)

Ulva fasciata f. taeniata Setchell, in Collins, Holden and Setchell,
Phyc. Bor.-Amer. (Exsicc), no. 862; Collins, Green Alg. N. A., 1909,
p. 216.

1920] Setchell-Gardner : Phycological Contributians 287

Ulva taeniata has been found thus far on the coast of central
California only, while TJ. dactylifera has been found only on that of
southern California. The differences between the two have been
enumerated under the latter species. From U. fasciata f. costata
Howe it differs particularly in its basal portion. Ulva fasciata Delile
seems to be a species nearly if not absolutely plane, while U. taeniata
is always crisply ruffled. No. 862 of the Phycotheca Boreali- Americana
represents this species very well.

Prasiola

There are eight species of Prasiola at present accredited to the
territory of the Pacific Coast of North America. One of these,
Prasiola crispa (Lightf.) Menegh., is, we believe, strictly terrestrial,
at least so far as our territory is concerned. Another, P. calophylla
(Carmich.) Menegh., is also terrestrial, but found with us in a marine
locality. Three species, P. horealis Reed, P. meridionalis sp. nov., and
P. delicota sp. nov., are strictly marine. One species, P. Gardneri
Collins, has been removed to the genus Merismopedia (M. Gardneri
Setehell), while three species, P. fluviatilis (Sommerf.) Aresch., P.
mexicana J. Ag., and P. nevadensis sp. nov., are inhabitants of cold
fresh-water streams.

Prasiola mexicana J. Ag. (1847, p. 6) is based upon specimens
collected in Mexico by Professor Liebmann of Copenhagen. The type
locality is Santa Maria Alpatlachna in Mexico. The habitat is
turbulent mountain streams. The habit is that of a broad plant,
umbilicate at the base and laciniate and irregularly crispate. Agardh
simply adds that the areolae are regularly quaternate. No mention
is made of the presence of intercellular lines, "viae inter stitiales," in
this first description. Jessen (1848, pp. 19, 20, pi. 1, figs. 17-20) gave
a more ample description and illustrations from materials and infor-
mation provided by Liebmann. Lagerstedt (1869, p. 26) redeseribed
the species, but drew his information largely, if not entirely, from
Bolivian specimens. J. G. Agardh (1882, p. 84) added nothing in his
final review of the genus Prasiola.

Thus far the specimens from our mountain streams have all been
referred to Prasiola mexicana J. Ag. (1847, p. 6), a species with broad
fronds, umbilicate or nearly so at the base, and arising from a disk,
possibly later developing secondary rhizoidal outgrowths of attach-
ment on the decumbent base. The color is dark green and the laciniae

288 University of California Publicati-ons in Botany [Vol.7

have crisped oi' nifflcd edges. This description is taken largely from
Jessen (1848, p. 19, \)\. 1, figs. 17-20), who, as noted above, received
his materials from the type collection directly from Liebmann, the
oi-iginal collector. Unfortunately we have not been able to examine
any of the type material and, consequently, must judge our plants
entirely from the point of view of the descriptions and figures. From
tliis point of view it seems safe to refer most of the various specimens
accessible to us to P. mexieana.

Under Prasiola mexieana have thus been referred specimens from
Montana (Tilden, Amer. Alg., no. 129), Wyoming (Tilden, loc. cit.,
no. 555«) and Oregon (Tilden, loe. cit., no. 5556, and Collins, Holden
and Setchell, Phyc. Bor.-Amer., no. 1186). The species is represented
in the Herbarium of the University of California by specimens from
"Washington (no. 132922, collected by Professor T. C. Frye in the
Ehoha River in the Olympic Mountains), from Oregon (no. 98182,
from McCleary Canyon near Portland, collected by A. S. Foster ; nos.
98181 and 98183, collected near Eugene and near Forest Grove re-
spectively by Professor A. R. Sweetser), while in California it has
been collected in Crane Creek above El Portal at an altitude of 4000-
4500 feet (Gardner, no. 4466). It is reported by Wolle (1887, p. 107,
pi. 91, fig. 24) from Colorado, where it was collected by T. S. Bran-
degee. It has also been collected in the Diamond range of mountains
in eastern Nevada, where it was found by Sereno Watson and de-
scribed by Horatio C. Wood (1871, p. 415, 1873, p. 182) under the
name of Viva merismopedioides. A portion of the type (or cotype)
has been examined and there seems little question as to the identity
of Wood's species and those we are assigning to Prasiola mexieana
J. Ag. The fragment of the type examined and from which our
figures (cf. pi. 21, figs. 8-10) are drawn for comparison was communi-
cated to us by Frank S. Collins, having been received by him from
the United States National Herbarium. Collins (1903, p. 17) placed
the Viva merismopedioides Wood under Manostroma quaternarium,
but later, as he writes us, made a second examination of the material
and found something seemingly different from what he had examined
previously and upon which he had based his oj)inion. The fragment
examined by us is clearly Prasiola mexieana as we understand that
species. The crass-section of Wood's plant, however, differs from the
description of Wood in that the cells (cf. Wood, 1873, p. 182) are not
in a single layer, but in two to several, as a glance at our illustrations
(ef. 1)1. 21, figs. 9, 10) will show.

1920] Setchell-Gardrier : Phycologieal Contributions 289

In connection with cross-sections of Prasiola mexicana, it may be
said that the various specimens agree very well as to the number and
arrangement of cells but differ somewhat in width of section. In
width the sections vary from 24/x to 52/x, but the increase in thickness
is more or less associated with the development of the so-called aplauo-
spores. "We find these arising in greater numbers from a single cell
than any description thus far indicates for any species of Prasiola.
Lagerheim has described and figured them for Prasiola mexicana var.
quitensis Lagerheim (1892, pp. 370-372, pi. 20, figs. 15-22) . He found
either a single layer of tetrads {loc. cit., fig. 18) or two layers of tetrads
(loc. cit., fig. 20) in the membranes of his variety. In our specimens
we find the double row of tetrads (cf. pi. 21, figs. 3-7), and also double
rows of tetrads intermingled with groups where the horizontal divisions
have proceeded farther, as they do in antheridial formation in certain
species of Porphyra. The majority of sections show the two kinds
(or stages in development) of these bodies intermingled. We feel it
necessary to consider these bodies as aplanospores until further investi-
gation can shed more light as to their fate, but the resemblance
between them and the species of Porphyra, as well as the resemblances
in cell structure, particularly as regards the shape and position of
the chromatophore, suggests strongly the possibility of a nearer re-
lation to the Bangiales than to the Ulvales and a remote ( ?) possibility
that we may be dealing here with antheridia and a very simple type
of cystocarp.

Comparing our specimens with the descriptions and figures of
Agardh and Jessen, we believe them to be true Prasiola mexicana, and
comparing them with Lagerheim 's descriptions and figures we are
inclined to agree with him that the Equador plant is different, possibly
even specifically.

One of us (Setchell) has found a Prasiola in Crane Creek (Mari-
posa County, California), between Big Meadow and MeCauley's, which
differs so much from P. mexicana that it seems desirable to describe
it as new. It diff'ers from P. mexicana in habit, color, and structure.
It is always elongated (cf. pi. 21, figs. 11-13), while P. mexicana is
broad and umbilicate. In color it is light, almost yellow-green, instead
of the darker green characteristic of P. mexicana. In surface view
(cf. pi. 21, figs. 14^16) the smaller groups of cells are more distinctly
placed and are further marked off into areas by delicate lines (or
borders) in the enclosing jelly. The sections (cf. pi. 21, figs. 17, 18)
are as broad as the narrower sections of P. mexicana, being 11 fj. to 25ju,

290 University of California FuhUcations in Botany [Vol. 7

or 26/u, wide. The aplanospores form a simple series of tetrads onlj'
and show no disposition to proceed further. On account of these
differences it seems best to describe these plants as belonging to a
new species, Prasiola nevadensis. It is to be recorded and is of
decided significance in Ibis connection that the other one of us
(Gardner) has collected characteristic Prasiola mexicana in the same
stretch of the same stream, at the same season but in a different year.
It may be, therefore, that our new species may be found to be included
among the stages in the development of, or states of, P. mexicana.
It seems so distinct, however, that we venture to describe it as new
and specimens will be distributed in a fascicle of the Phycotheca
Boreali- Americana soon to be issued.

Prasiola nevadensis comes much nearer to P. fluviatilis (Sonnuerf.)
Aresch. than to P. mexicana. It differs from P. fluviatilis so far as
we may determine from descriptions, figures and scanty specimens
available to us, in being broader and not tapering regularly from
above toward, the base. We do not know much about the structure of
P. fluviatilis, but suspect that the general character and arrangement
of cells may closely resemble those of our P. nevadensis. Our plant
may be closer to P. fluviatilis var. Hausmanni Grun., in that it is a
broader species than typical P. fluviatilis, but further comparison is
impossible. We append the following diagnosis:

Prasiola nevadensis sp. nov.

Plate 21, figures 11-18

Thallis gregariis, angustis, 2-3 cm. longis, 0.5-1 cm. latis, oblongis
usque ad oblaneeolatis, e basibus latis oriundis, primo disco minuto,
aetate provecta, rhizoidibus e basi decumbente emissis affixis, dilute
viridibus, marginibus undulatis et crispatissimis ; membrana l-i~25/i.
crassa; eellulis in gregibus parvis e superficie visis quaternis, rotundis
aut augulatis, laxe positis et in areolas indistinctas segregatis, in
sectione transversal! rotundis usque ad valliformibus ; viis intersti-
tialibus distinctis nullis; aplanosporis in serie singula, quaternis.

Tlialli gregarious, narrow, 2-3 cm. long, 0.5-1 cm. wide, oblong to
oblanceolate, from a broad base, attached at first by a small disk, later
by rhizoids from the basal portion, pale green, witli margins undulate
and very much crisped ; membrane 14-25/x thick ; cells in small groups
of tetrads as seen from the surface, rounded or angular, loosely ])]aced,
forming indistinct areoles, but without interstitial lines, cells in trans-
verse section rounded to elongated vertically; aplanospores (?) in a
single layer of tetrads.

In rapid waters of Crane Creek, Foresta townsite, Mariposa
County, California, at 3500^000 feet altitude. Type no. 6507,
Setchell.

1920] Setchell-Gardner : Phycological Contributions 291

Prasiola meridionalis sp. nov.

Plate 25, figure 2

Thallo usque ad 7 mm. alto, stipiti brevi latoque in laminam latam,
eordatam et rosulatam aut cucullatam abeunte, sordide viridi ; cellulis
neque in areolis distinctis ordinatis neque per vias interstitiales sepa-
ratis ; membrana 40-45/; crassa, in seetione transversali cellulas 14-18ju,
altas et 7-8/u, latas marginibus latis hyalinisque (usque ad 13/x erassis)
ostendente ; akinetis dispersis, magnis, parietibus erassis ; aplano-
sporis probabiliter 128-512 (4X4X8 aut 8X8X8) in aplanospo-
rangio singulo.

Frond up to 7 mm. high, with short and broad stipe, soon expanded
into a broad, cordate, rosulate or cucullate blade, dirty green ; cells
neither arranged in distinct areolae nor separated by interstitial lines ;
membrane 40-45/a thick, section showing cells 14-1 8jii high and 7-8/a
wide with broad hyaline margins (up to ISfx thick) ; akinetes scattered,
large, thick walled; aplanospores probably 128-512 (4X4X8 or
8X8X8) from a single aplanosporangium.

On exposed rocks or rocky islets above the high-water mark but
exposed to the force of the waves. Washington (Friday Harbor
and Neah Bay) to central California (entrance to Tomales Bay).
Type no. 3824, Gardner.

The specimens taken as the type of this species were collected by
one of us (Gardner) at Neah Bay, Washington. We are also inclined
to refer here specimens collected at "Minnesota Reef" at Friday
Harbor, Washington, and at the entrance to Tomales Bay, California.
In the last two localities the species is associated with Gayella con-
stncta. Cultures, however, strongly indicate the independence of the
two plants from one another.

Prasiola meridionalis comes near to P. horealis Reed, but the frond
of the latter is areolate and Avith more or less distinct intercellular
lines. P. horealis is, so far as found, infested with a fungus
{Guignardia alaskana Reed), while none of the three collections of
P. meridionalis shows any trace of such a parasite.

Prasiola delicata sp. nov.

Plate 22, figure Za-f, and plate 25, figure 1, and plate 24, figure 8

Thallo 1-1.5 mm. alto, lato et breviter stipitato, in laminam late
oblongam aut eordatam expanso, marginibus crispatis et involutis,
atro-cyaneoviridi ; membrana 17-20/a crassa, cellulis neque in areolas
distinctas ordinatis neque per vias interstitiales separatis; akinetis
non visis; aplanosporis usque ad 512 e cellula singula (8X8X8) sed
frequenter tantum 128; cellulis valliformibus, in seetione transversali
verticaliter 10-12/x, 2-plo aut ultra diam. longioribus.

292 University of California Publications in Botany [Vol.7

Frond 1-1.5 mm. high, broad and shortly stipitate, expanding
directly and abruptly to broadly oblong oi- cordate, margins crisped
and inrollcd, dark bluisli green; mciubrane 17-20/x tliiek, cells not
groui)ed into distinct areolae and not separated by interstitial lines;
akinetes not seen; aplanospores up to 512 from a single cell (8X8
X 8), but often only 128; cells palisade-like and 10-12/x in vertical
diameter, in section twice or more times as high as broad.

Growing at or near the upper tide mark on rocky islets. Sitka,
Alaska. Type no. 3981, Gardner.

Prasiola delicata has a decidedly thinner membrane than any other
of our marine species, shows little areolation, and has a larger number
of aplanospores formed within a single aplanosporangium. These
characters seem to indicate its just claim to be considered a distinct
species.

Entocladia cingens sp. nov.

Plate 23, figure 7

Thallo textum pseudoparenchymaticum filamenta hospitis cingens
et intus membranam, sed filamenta marginalia ad longitutlinem hos-
pitis parallela ostendens mox formante ; cellulis centralibus fere isodia-
metricis, 5-8yu, diam., tardiore augescentibus et in sporangia trans-
formatis ; cellulis filamentorum liberorum marginalium 3-4/j, diam.,
2-3-])lo diam. longioribus, terminalibus longis, conicis.

Thallus early forming a pseudoparenchymatous tissue surrounding
the filaments of the host within the membrane, having a few marginal
filaments extending parallel with the long diameter of the host ; cells
in the center of the thallus nearly isodiametric, 5-8/x diam., enlarging
later to form sporangia ; cells of the free marginal filaments 3-4;a diam.,
2-3 times as long as the diameter, terminal cells long, conical.

Growing within the membrane of Chaetomorpha calif arnica Wille.
Southern California (Ocean Beach, near San Diego), December. Type
no. 3528fl, Gardner.

The plants of this species seem to be nearing maturity in December,
since a few empty cells in the center of the thallus were found from
which reproductive bodies probably had escaped. Aside from this
condition, nothing further is known of its method of reproduction.

Entocladia cingens is placed in this genus on account of the re-
semblance of the vegetative development to that of the type species,
E. viriclis Reinke, and because it has the same endophptic habit as
has that species. It differs from E. viridis in having the branching
filaments more closely coalescent, the enlarging cells in the main ])ai-t
of the thallus soon forming a pseudoparenchymatous tissue, leaving
only a few free marginal filaments.

1920] S&tchell-Gardner : Phycological Contributions 293

E. viridis, E. codicola, and E. cingens form a well connected series,
using the vegetative characters as a basis. The first named species
has a rather wide spreading thallus, composed of relatively sparsely
branching filaments, scarcely, if at all, coalescing in the center. In
the second the filaments coalesce freely in the center so that at least
half of the thallus is formed into a pseudoparenchymatous tissue at
the time of reproduction, but leaving an abundance of free branching
marginal filaments. The thallus of the third is almost wholly trans-
formed into a pseudoparenchymatous tissue at maturity leaving only
a few free marginal filaments.

Entocladia codicola sp. nov.

Plate 24, figures la, 7b

Filamentis laete viridibus, profuse ramosis, maturitate stratum
continuum centrale ad peripheriam terminos liberos et attenuatos ex-
hibens formantibus ; cellulis juvenis 3—1^ diam., 1-2.5-plo diam. longi-
oribus, terminantibus gracilibus et conicis; cellulis thalli centralibus
5— 8/x diam. ; pyrenoidibus singulis ; generatione ignota.

Filaments light green, branching profusely, at maturity forming
a continuous layer in the center of the mass with many tapering free
ends around the margin ; young cells 3-4/i, diam., 1-2.5 times as long,
terminal cells slender and conical ; cells in the center of the thallus
5-8/i. diam. ; pyrenoid single ; reproduction unknown.

Growing in the membrane, at the tips of the utricles of C odium
fragile. Central and southern California. Type no. 4121, Gardner.

Entocladia codicola seems closely related to Entocladia viridis
Keinke (1879, p. 476, pi. VI, figs. 6-9), found growing in the mem-
brane of Derhesia; but it is a larger plant with the filaments much
more compact in the center, forming in fact a pseudoparenchymatous
disk with free filaments around the margin. The cells are shorter
than those of E. viridis, some being even shorter than the diameter.
In the pseudoparenchymatous character of the center of the disk-like
frond it resembles Entocladia Flustrae Reinke (1888, p. 241, nomen
nudum, 1889, p. 31, pi. XXIV, 1889a, p. 86), but the dimensions given
for that species are in general greater than those in ours. Repro-
ductive bodies have been observed in the cells of the central portion
of the disk in E. codicola, but the nature of these, their method of
escape, and their subsequent behavior have not been determined. Until
more is known concerning these later phases of the plant, its proper
placing must remain somewhat in doubt. It is here ])rovisionally
placed with Entocladia on account of its endophytic habit of growth.

294 Vniversity of California Puhlications in Botany ['^'ol. 7

rather than with Epicladia, which has the habit of gro^\•ing on the
outside of the liost. Tliis habit of growth seems to be the only one
by wliicli tlie two genera are distinguished as far as the diagnoses
reveal. Little, however, is known concerning the reproduction in
Epicladia, and uiilil the matter can be cleared up it can have but
little claim to generic distinction. Reinke expressed doubt as to the
validity of the genus when he diagnosed it (1889). Collins (1909)
has retained both genera, and under Endoderma (Entocladia) has
included two species, viz., E. pithophorae West and E. polyniorpkum
West, which are epiphytic, and thus, as he remarks {loc. cit., p. 280),
"connects Endoderma with Epicladia, but the filaments do not unite
to form a definite disk."

Entocladia codicola seems to be confined to the coast of California
and to the above mentioned host plant, at least, examination of con-
siderable material of different species of Codium in different localities
ranging from Sitka, Alaska, to southern California has not revealed
its presence elsewhere.

Internoretia gen. nov.

Thallus endophyticus, e filamentis profuse ramosis, primo cellulis
in serie simpliei per divisiones apicales verticaliter ad longitudinem
posit as augescentibus, tardiore per divisiones obliquas et longitudi-
nales, filamenta cylindrica cellulis numerosis composita formantibus ;
filamenta ramos ad angulos rectos emittentia, ramis anastomosantibus ;
chromatoi)hora parietalis, pyrenoidibus singulis ; generatio ignota.

Tliallus endophytic, consisting of profusely branched filaments, at
first of a single series of cells increasing by apical divisions perpen-
dicular to the long diameter, but later by oblique and longitudinal
divisions, building up cylindrical threads composed of numerous cells
in cross diameter ; branching at right angles, anastomosing, forming
a network ; chromatophore parietal, with one pyrenoid ; reproduction
unknown.

The genus Internoretia is proposed for a peculiar endophyte found
by Professor T. C. Frye, growing within the membranes of Parphyra
Naiadum. Its reproduction not having been determined, it is among
the numerous form-genera of uncertain position and placed provision-
ally among the Chaetophoraceae. It resembles Pseudodictyon Gardner
and Zygomitus B. and F. From the former it differs in forming solid
filaments several cells in thickness. From Zygomitus, Internoretia
differs in the greater regularity of its solid portions and in tlic more
uniform network brought about by the regular giving off of branches
at right angles.

1920] Setchell-G<irdner : Phycological Contrihutians 295

Internoretia Fryeana sp. nov.

Plate 23, figures 3-6

Cellulis filamentorum terminaliuni 3.5-5/* diam., 3-5-plo diam.
longioribiis, apicalibus conicis ; cellulis partium thallorum vetustiorum
isodiametricis, angularibus ; aliter ut in generi.

Cells of the terminal filaments 3.5-5/a diam., 3-5 times as long,
apical cell conical ; cells of the older part of the thallus isodiametric,
angular; otherwise as the genus.

Growing within the membrane of Porphyra Naiadum. Friday
Harbor, Washington, July. Type no. 4260, Gardner.

This most interesting little plant is as yet known only from the
collections and observations of Professor T. C. Frye. It generally
occurs in such abundance as to discolor the host plant. In some years
it is very common, discoloring most of the plants of Porphyra in the
neighborhood of the Puget Sound Marine Station, while in other years
it is difficult to find any plants at all.

Pseudulvella prostrata (Gardner) comb. nov.

Growing on the basal portion of Iridaea laminarioides. Central
California (Lands End, San Francisco).

Ulvella prostrata Gardner, New Chlorophj^ceae, 1909, p. 373, pi.
14, figs. 1, 2 ; Collins, Green Alg. N. A., 1909, p. 287 ; Collins, Holden
and Setchell, Phyc. Bor.-Amer. (Exsicc), no. 1629.

Recent observation has revealed the presence of a pyrenoid in each
cell of the plant under consideration, a character not belonging to the
genus Ulvella. This character militates against placing it in that
genus. As found growing on Iridaea, it has no penetrating rhizoids,
thus differing in this respect from the genus Pseudopringsheimia, to
which it seems closely related. Nothing has yet been learned regarding
reproduction. Considered in the light of our present knowledge,
Pseudulvella seems to be the genus most appropriate for its reception.

Pseudulvella applanata sp. nov.

Thallo tenui parenchymatico, per incrementum marginale ad mm.
plura augescente, laevi et nitido, 45-55;u, crasso, laete viridi ; cellulis
in series verticales, proxime regulares positis, proxime isodiaraeitricis,
acute angulatis, 6-7. 5/a diam. ; chromatophoris parietem cellularum
tegentibus, pyreuoidibus singulis; zoosporangiis (?) e cellulis super-
ficialibus leviter transformatis oriundis; generatione ignota.

Thallus thin, parenchymatous, spreading by marginal growth, ex-
panded to several mm. in diam., smooth and glossy, 45-55/a thick ;
color grass green ; cells in fairly regular vertical rows nearly isodia-

296 Vniversity of California Publications in Botany [Vol. 7

mcti-ic. sliarply aiij^'Icd, 6-7.5/1 diam. ; chroinatopliore covering the cell
wall provider with one pyi'ciioid ; zoosporangia (?) slightly modified
siii'face cells; reproduction unknown.

Growing on the shells of Littorina planaxis Nutt. Central Cali-
fornia. Type no. ;3597, Gardner.

Littorina planaxis Nutt. is very abundant in small tide pools and
in moist places along high-tide level from Sitka to San Diego.
Pseudulvclla applanata has been studied only on material found along
the coast of central California, but it is presumed to have a much
wider distribution.

Its presence on the host is readily recognizable by the expanded,
grass green, glossy appearance of the surface of the shell. Micro-
scopically it may readily be distinguished from all other described
species of the genus by its very small, closely compact, parenchymatous
cells, and the seeming absence of radiating filaments composing the
basal layer. It spreads over the host by tangential and by radial
divisions of the peripheral cells, at least the process can thus be de-
scribed when the plant is of considerable size. It probably starts on
the very young host, and doubtless many plants early coalesce to
form a confluent thallus. We have not been able to ascertain the
nature of the early developmental stages, although very small shells
have been examined.

Reproductive bodies have been seen to escape from the surface
cells. Whether these are zoospores or gametes, the number of cilia
they possess and their behavior after escaping are subjects for further
investigation.

The three species of Pseudulvella known from the M^estern coast
of North America may be arranged, so far as the basal layer is con-
cerned, in the following sequence : P. prostrata, with distinctly
radiating basal filaments which branch rather frequently and which
are comparatively loosely coalescent ; P. cmisociata, with indistinctly
radiating ba.sal filaments closely coalescent; and P. applanata with a
parenchymatous basal layer.

Pseudulvella consociata sp. nov.

Plate 24, figures 4-6

Thallo magnitudine et peripheria irregulari, 100-1 40/a crasso, per
filamcnta ii'rcgularia et radiantia augeseente ; filamentis mox coale-
scentil)us <'t iJai'fnchymaticis, atroviridi ; filamentis erectis dense ad-
haerentibus. T-lO/x diam., cellulis prope cylindricis, forma leviter
irregularibus, 1-2-plo diam. longioribus; zoosporangiis (?) terminal-
ibus. ])yrifoi'ini])us us<|ue ad sphaericis, 8 zoosporas emittentibus.

1920] Setchell-Gardner : Phycological Contributions 297

Thallus irregular in size and in outline, 100-140ja thick, increasing
by irregular and obscurely radiating filaments early coalescing and
becoming parenchymatous ; color dark green ; erect filaments firmly
coalescent, 7-10/x diam., cells nearly cylindrical, slightly irregular in
form, 1-2 times as long as the diameter; zoosporangia (?) terminal,
pyriform to spherical, producing 8 zoospores.

Growing on the shells of Ilyanassa ohsoleta Say. Central Cali-
fornia (Bay Farm Island, Alameda). Type no. 4268, Gardner.

The shells of Ilyanassa ohsoleta Say. were introduced some years
ago along with oysters from the Atlantic coast of North America, and
possibly the plant here described was introduced with the host.

A comparison of this species with others will be found included in
the discussion under P. applanata.

Pseudopringsheimia apiculata sp. nov.

Plate 22, figures 1, 2

Thallis minutis, 145-1 60/i, crassis, dum solitariis hemisphaericis, sed
frequenter dense gregariis et stratum continuum 2-3 mm. diam. for-
mantibus, laete viridibus; filamentis erectis 8-12jli diam., 9-12 cellulis
cylindricis aut leviter tumidis compositis ; zoosporangiis (?) 8 zoosporas
emittentibus, terminalibus, leviter tumidis, convexis usque ad conspicue
apiculatis ; zoosporis (?) 4-ciliatis.

Thallus minute, 145-1 GO/x thick, hemispherical when alone, but
often with many crowded closely together forming a continuous
stratum 2-3 mm. diam. ; color bright green ; erect filaments 8-12/x diam.,
composed of 9-12 cylindrical or slightly swollen cells ; zoosporangia
( ?) producing 8 zoospores, terminal, slightly swollen, varying from
convex to decidedly apiculate; zoospores ( ?) 4-ciliated.

Growing on the rhachis and the cysts of Egregia Menziesii.
Central California. Type no. 4361, Gardner.

Psendopnngsheimia apiculata is closely related to P. confluens
(Rosenv.) Wille. The most conspicuous difference is to be found in
the shape and size of the zoosporangia, if the terminal reproductive
cells are to be designated as such. Those of P. confluens are long and
comparatively narrow, and produce 30 to 40 zoospores, while in P.
apiculata they are shorter, somewhat swollen, mostly with a pro-
nounced terminal projection, and produce about 8 zoospores. These
reproductive bodies are very small and it is exceedingly difficult to
determine their number of cilia. On one occasion four cilia were
observed, but the reproductive bodies seemed a little larger and some-
what more irregular in form than the average. These may have been
the zygotes which had been formed by the fusion of 2-ciliated gametes
and which had not vet come to rest.

298 University of California Puhlicxitions in Botany 1^'"^. 7

Gomontia polyrhiza (Lagerh.) B. and F.
Plate 24, figure 1

"Sporangia" irregularly and broadly clavate to nearly cylindrical,
up to 150/A diain., and 240/x long, producing usually several blunt, at
times sliglitly l)i;iii('licd, rhizoids at the smaller end.

Growing in clam shells. Neah Bay, Washington.

Bornet and Flahault, Note sur deux nouveaux genres d'algues
perforantes, 1888, pp. 161-163 (as to combination only). Codiolum
polyrkizum n. sp. Lagerheim (at least in greater part).

The above description is taken in part from the original of Lager-
heim and in jiart from the material collected at Neah Bay. The
material from which Lagerheim drew his description was apparently
in the sporangial stage exclusively, at least he did not recognize a
sterile or vegetative stage. The Neah Bay material, collected in May,
is likewise in a reproductive stage, or if the vegetative stage is present
it and the sporangial stage could not be identified as belonging to the
same species, hence the incompleteness of the description.

The sporangia approximate so closely to the figures (especially
figs. 10, 11) and the description of Lagerheim as to make it sufficiently
safe to ally our plant with his and to keep it distinct from the G.
polyrhiza of Bornet and Flahault (G. Bornetii S. and G.).

The filaments of this species have been examined by us in a
specimen distributed by Reinbold from Kiel. Beinbold's specimens
have "sporangia" largely of the Codiohim-type, both old and young,
but is also has an occasional "sporangium" of the Acarid-type
(apparently good G. Bornetii). Since Beinbold's locality is not far
distant from Lagerheim 's type locality, it seems extremely probable
that his plant is true G. polyrhiza. The filaments in Beinbold's speci-
mens are so close to those of G. Bornetii as figured by Bornet and
Flahault {loc. cit.) as to be indistinguishable.

Gomontia Bornetii nom. nov.

Horizontal filaments irregular, much branched, erect filaments with
clavate ends, less bi-anched ; cells 4-12fi, most frequently 6/x diam.,
15-5;V long, cylindrical to more or less swollen and crooked :
"sporangia" variable and irregular in form, 8-125/i, wide, 150-200/x
long, having numerous, mostly simple rhizoids arising principally on
one side ])ut occasionally ])romiscuously scattered all over tlie spo-
rangia ; zoospores of two sorts, one 3.5/t wide and 5/i. long, the other
5-6|u, wide and 10-1 2/i, long, development unknown; aplanospores 4fi
diam.

1920] S&tchell-Gardner : Phycological Ccnitrihutians 299

Growing in clam shells. Neah Bay, Washington.

Gomontia polyriiiza Bornet and Flahault, Notes sur deux nouveaux
genres d'algues perforantes, 1888, pp. 161-163 (not Codiolum poly-
riiiza Lagerheim),

Bornet and Flahault (1889, p. 9) distinctly state that the greatest
dimensions of the "sporangia" in their specimens are 120/x for height
and 75/A for width, and mention that Lagerheim found "sporangia"
in the specimens up to 240/a in height and 60/i, in breadth. We judge,
therefore, that the Codiolum-type of "sporangium" which Lager-
heim figures (1885, pi. 28, figs. 10, 11 in particular) and describes
{'^plerumque plus minus elongatis," loc. cit., p. 22) was not to be
found in the French material and certainly is not illustrated by
Bornet and Flahault, unless figure 9 on plate 7 may represent it.
The type of "sporangium" illustrated by Bornet and Flahault (1889,
pis. 7, 8) belongs to the shorter and broader type, the Acarid-type as
it may be called, and has blunt, simple or slightly branched rhizoids.
Lagerheim (loc. cit., pi. 28, figs. 7, 8, 12, 13) has also figured "spo-
rangia" of the Acarid-type and probably found a mixture of species
in the shells he examined. Since, however, he emphasizes the elon-
gated, or Codiolum, type of sporangia, it seems best to reserve his
specific name for the species with the Codiolum-type of "sporangium"
and assign the new specific name {Bornetii) to the species having the
Acarid-type of "sporangium" and with blunt, rather stout, simple
or, at most, slightly branched rhizoids.

The filaments of G. Bornetii are well represented by Bornet and
Flahault (1889, pi. 6, figs. 1-8) and by their usually large number of
short, blunt or almost bulbously enlarged branchlets and their compact
massing make a characteristic appearance after decalcification. They
are very similar to those of G. polyrMza, as far as we can determine,
but someAvhat different from those of G. kabrorhiza, although this
difference is not readily described.

While we find what seems referable to G. Bornetii in the Puget
Sound region and that of central California, we desire more abundant
and more decisive material before we can determine this.

Gomontia habrorhiza sp. nov.

Plate 24, figures 2, 3a, 3b, 3c

Filamentis repetite et irregulariter ramosis; cellulis forma magni-
tudineque maxime variabilibus, typice cylindrieis, 4-7/a diam., 2-8-plo
diam. longioribus, chromatophoris pyrenoidibus destitutis, eellulam
totam implentibus; "sproangiis" (gametangiis ? aut aplanosporan-
giis ?) angustis aut latis, obtuse conicis, 50-70^ altis, 25-60//, latis, in

300 University of California Piihlicatimis in Botany [Vol.7

latere inferno i']ii/.oi(lil)us imiltis gracilliinis, attenuatis et dendritice
raniosis in<luti.s; gciu'ratioiu' ignota.

Filaiucnts repeatedly and ii'regulai-ly branched; cells very variable
in form and size, typically cylindrical, 4-7 fx. diam., 2-8 times as long;
elu'oiiiatopliore without jjyi-enoids. filling the cell; "sj)Oi'angia " narrow
to witle, bluntly conical, oO-TO/i. high, 25-60/a wide, developing many
very slender, attenuate, dendritically branched rhizoids from th(* lower
side ; reproduction unknown.

Growing on dead clam shells. Neah Bay, "Washington. Type no.
3825, Gardner.

In a Gomontia inhabiting certain shells from Neah Bay we have
found all the "sporangia" of the Acarid-type and with the processes
or rhizoids slender, branched, and attenuated to a jioint. The "spo-
rangia" seem so distinct from those of G. Bornctii that we de.scribe
the plant possessing them as new. In some shells we have found the
"sporangia" of this species intermingled with others. The vegetative
filaments of G. hahrorhiza seem less entangled and slightly larger than
those of either G. polyrhiza or G. Bornetii.

Gomontia caudata sp. nov.

Plate 23, figures 1, 2

Filamentis brevibus, parce ramosis; cellulis 5.5-6.5^ diam., 2-12-
plo diam. longioribus; chromatophoris cellulas terminales et "spo-
rangia" juvena iinplentibus, in cellulis vetustioribus interruptis;
pyrenoidibus inconspicuis ; "sporangiis" (gametangiis ? aut aplano-
sporangiis ?) clavatis, 50-70//, latis, 160-200/x longis, inferne ad rhizoide
singula attenuatis, parietibus maturitate crassis, hyalinis, homogeneis ;
rhizoidibus saepe maxime incrassatis striatisque.

Filaments short, sparsely branched ; cells 5.5-6.5/a diam., 2-12 times
as long; chromatophore covering the terminal cells and young "spo-
rangia," broken in the older cells; pyrenoids inconspicuous: "spo-
rangia" clavate, 50-70/^ diam., 160-200/x long, tapering to a single
rhizoid below with thick, hyaline, homogeneous wall at maturity ;
rhizoid often becoming much thickened and striated.

Growing in shells of Mytilus californicus Conr. Neah Ba}-, Wash-
ington. Type no. 3825, Gardner.

"We have found in shells of the larger edible mussel of our coast a
Gomontia with filaments seemingly less abundantly branched and
"sporangia" (aplanosporangia ?) with very thick walls and a .single
long rhizoid (cf. pi. 23, figs. 1, 2). These "sporangia"" bear a certain
resemblance to the "cells" figured by Lagerheim (1885, pi. 28, figs.
4, 6), but are, at least, thicker walled. The fact which seemed to
indicate distinctness was that only this ty])(' of "sporangium" was
found in the shells examined.

1920] Setchell-Gardner : Phycological Contributions 301

LITERATURE CITED

Agardh, C. a.

1822. Species algarum rite coguitae cum synonymis, differentiis specificis

et descriptionibus succinctis. Vol. I, pt. II, pp. 269-531.
1829. Icones algarum europaearum. Representation d 'algues europeennes
suivi de celle des espeees exotiques les plus remarquables recem-
ment deeouvertes. Vol. 3, pis. 21-30. Leipzig.

Agardh, J. G.

1847. Nya Alger fran Mexico. Oefvers. af Kongl. Vet.-Akad., Forhandl.

Arg. 4, no. 1. Stockholm. Often quoted as "Alg. Lieb."
1882. Till Algernes systematik, Nya bidrag. Tredje afdelningen, vol. 19.

Lunds Univ. Arssk.

Batters, E. A. L.

1894. New or critical British algae. Grevillea, vol. 22, p. 114.

BORXET, E.

1892. Les algues de P.-K.-A. Schousboe, reeoltees au Maroc & dans la
Mediterranee de 1815 a 1829. Mem. Soe. Nat. Sci. Nat. et Math,
de Cherbourg, vol. 28, p. 165. Paris.

BoRXET, E., and Flahault, C.

1888. Note sur deux nouveaux genres d 'algues perforautes. Journal de

Botanique. Vol. 2, p. 161.

1889. Sur quelques plantes vivant dans le teste calcaire des mollusques.

Bull. Soc. Bot. de France, vol. 36, p. cxlvii, pis. 6-12.

Collins, F. S.

1903. The Ulvaceae of North America. Rhodora, vol. 5, p. 1, pis. 41-43.
1909. The green algae of North America. Tufts College Studies, vol. 2,

no. 3, Scientific Series. Mass.
1912. The green algae of North America, Supplement I. Tufts College

Studies, vol. 3, no. 2, Scientific Series. Mass.

Collins, F. S., Holdex, L, and Setchell, W. A.

Phycotheca Boreali-Americana. (Exsicc). Maiden, Mass. Fascicles
18, 24, 35, C.

Gardner, N. L.

1909. New Chlorophyceae from California. Univ. Calif. Publ. Bot., vol. 3,
pp. 371-375, pi. 14.

Gobi, C.

1879. Bericht ueber die algologischen Forschungen im finnisehen Meerbusen
im Sommer 1877 ausgefiihrt. St. Petersb. Gesellsch. d. Naturf.,
vol. 10, p. 83 (Obshchestvo estestvoispytatelei).
Howe, M. A.

1911. Phycological studies V. Some marine algae of Lower California,

Mexico. Bull. Terr. Bot. Club, vol. 38, pp. 489-514. pis. 27-34.
1914. The marine algae of Peru. Mem. Torr. Bot. Club, vol. 15, pp. 1-185,
pis. 1-66.

302 University of California Publications in Botany [Vol. 7

Jessen, C. F. G.

1848. Prasiolae generis alsaruiii nionographia. "Dissertatio inauguralis
botanica. " Kilia (Kiliae).

Kjellmax, F. R.

1897. Blastophy.sa potymornJia och Urospora incrassata. Tva uya Chloro-

jihyceer fran sveriges vestra kust. Bihang till Kongl. Sv. Vet.-

Akad. Handl., vol. 215, af<l. li, uo. 9. Stockholm.

KUETZING, F. T.

1S49. Species Algarum. Leipzig.
185(5. Tabulae Phycologicae. Vol. 6.

Lagerheim, G.

1885. Codiolum pohjrhizum n. sp. Ett bidrag till kannedomen cm slagtet
Codiolum A. Br. Oefversigt af Kongl. Vet.-Akad. Forhand., no. 8,
p. 21, pi. 28. Stockholm.

1892. Ueber die Fortpllanzung von Frasiola (Ag.) Menegh. Berichte Deut.

Botan. Gesellseh., vol. 10, p. 366.

Lagerstedt, N. G. W.

1869. Om algslagtet Prasiola. Forsok till en monograph! Akademisk afhand-
ling. Upsala.

Reed, Minnie

1902. Two ascomycetous fungi parasitic on marine algae. Univ. Calif.

Publ. Bot., vol. 1, pp. 141-161, pis. 15, 16.

Reinke, J.

1879. Zwei parasitische Algen. Botan. Zeitung, vol. 37, p. 473, pi. VI.

1888. Einige neue braune und griine Algen der Kieler Bucht. Berichte der

deutsch. Botan. Gesellseh., vol. 6, p. 240.

1889. Atlas deutscher Meeresalgen. Ft. I. Berlin.

1889«. Algenflora der westlichen Ostsee deutschen Antheils. Sexter Bericht
der Konimission zur wissenschaftlichen Untersuehung der deutschen
Meere in Kiel. I Heft. Berlin.

Rosen vinge, L. K.

1893. Gronlands Havalger. Meddelelser om Gronland, vol. 3, pp. 765-981.

Copenhagen.

Ruprecht, F. J.

1851. Tange des Ochotskischen Meeres. Middendorflf 's siberische Reise,
vol. 1, pt. 2, lieferung 2, p. 193.

Saunders, De A.

1901. Papers from the Harriman Alaska Expedition, 25, The Algae. Proc.
Wash. Acad. Sci., vol. 3, pp. 391-486. Washington.

Setchell, W. a., and Gardner, N. L.

1903. Algae of Northwestern America. Univ. Calif. Publ. Bot., vol. 1, uo. 3,

pp. 165-418, pis. 17-27.

TiLDEN, Josephine E.

American Algae (Exsicc). Centuries IT, A'T.

WiLLE, N.

1909. Conjugatae und Chlorophyceae, in Engler and Prantl, Natiirl. Pflan-
zenfam., Nachtrage zum 1 Th., 2 Abt.
WOLLE, F.

1887. Fresh-water algae of the United States, etc. Bcthieliiiiii, Pa.

1920] Setchell-Gardner : Phycological Contributions 303

Wood, H. C.

1871. Algae, in Watson, in United States geological exploration of the

fortieth parallel. P. 415. Washington.

1872. A contribution to the history of the fresh-water algae of North

America. Smithsonian contributions to knowledge, 241, vol. 19.
Washington*

(The contribution was accepted for publication in Februar}-,
1872, but judging from the imprints of the various folders the
printing began in November, 1871, and was completed in October,
1872. Just (vol. 1, p. 2) gives the date of publication as 1873, and
vol. 19 of the Smithsonian Contributions is dated 1874. We are
using the date 1872 as seemingly the most authoritative one.)
Yendo, K.

1916. Notes on algae new to Japan V. The Bot. Mag., vol. 30, p. 243.
Tokyo.

PLATE 21
Prasiola mexicana J. Ag.

Ih ill. Tniv. Calif., no. 911S1, collected near Eugene, Oregon, Vjy Professor
A. R. Sweetser, May, 190:5.

Fig. 1. Habit of typical plant. X 0.5.

Fig. 2. Surface view of thallus lobe. X 375.

Fig. 3. Transverse section showing double layer of tetrad-type of aplaiio-
spore (?) formation. X 375.

Fig. 4. Similar section showing both tetra<l and "authcridial " typo of
aplanospore (?) formation. X 375.

Fig. 5. Similar section to that of figure 3, but less regular. X 375.

Herb. Univ. Calif., no. 91182, collected near Portland, Oregon, by H. A.
Foster.

Fig. (). Transverse section showing single layer of tetrad-type of aplano-
spore (?) formation. X .375.

Fig. 7. Transverse section showing "antheridial" type of aplanospore (?)
formation. X 375.

No. 1545, U. S. Geol. Surv., collected in the Diamond Range, Nevada, by
Sereno Watson and type collection of Ulva merismopedioides Wood.

V'ig. 8. Surface view of lobe of thallus. X 375.

Fig. 9. Transerve section showing double layer of tetrad-tyi^e of aplano-
spore (?) formation. X 375.

Fig. 10. Similar section showing "antheridial" type of aplanospore (?)
formation. X 375.

Prasiola nevadensis S. and G.

No. 6507, collected in Crane Creek, Foresta townsite, Mariposa County,
California, by W. A. Setchell, June, 1914.

Fig. 11. Habit of young plant. X 0.5.

Fig. 12. Habit of young plant. X 0.5.

Fig. 13. Habit of older plant. X 1.

Fig. 14. Surface view of lobe of thallus. X 375.

Fig. 15. Surface view of lobe of thallus. X 375.

Fig. 16. Surface view of portion of thallus. X 375.

Fig. 17. Transverse section showing cells probably on the way to aplano-
spore (?) formation. X 375.

Fig. 18. Similar section showing tetrad-type of aplanospore ( ?) formation.
X 375.

All the figures of this plate were drawn by Dr. Helen M. CJilkcy under the
direction of W. A. Setchell.

[304]

UNIV. CALIF. PUBL. EOT. VOL. 7

[ SETCHELL-GARDNER ] PLATE 21

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PLATE 22

Pseudopringsheimia apiculata S. and G.

Fig. 1. A section through the thallus of a mature plant perpendicular to
the host. X 250.

Fig. 2. A section through the thallus of a young plant. X 250.

Prasiola delicata S. and G.
Fig. 3. a-f, series of different forms of plants. X 10.

Ulva vexata S. and G.

Fig. 4. A group of plants showing different shapes and sizes. X 1.
Fig. 5. A group of mature plants showing the presence of the parasitic
fungus Guignardia Ulvae Reed. X 3. Taken from Reed, 1902, pi. 15, fig. 1.

Fig. 6. A cross-section showing the presence of fungal hyphae in the
medulla. X 250.

Fig. 7. A surface view. X 250.

[306]

UNIV. CALIF. PUBL. BOX. VOL. 7

[ SETCHELL-GARDNER ] PLATE 22

wmsbMOTs?

3MM|W®f

yj

Wfk

PLATE 23

Gomontia caudata S. and G.

Fig. 1. Two pieces of filaments. X 400.

Fig. 2. ad, different stages and forms of "sporangia." X 400.

Internoretia Fryeana S. and G.

Fig. 3. A surface view of the host plant showing the method of perme-
ation and branching of a few terminal filaments. X 375.

Fig. 4. A stage slightly in advance of figure 3 showing cell divisions in
planes parallel to the long diameter of the cells. X 375.

Fig. 5. A stage in development nearing maturity. X 375.

Fig. 6. A cross-section of the host cutting the filaments of Internoretia at
right angles to their long diameter. X 375.

Entocladia cingens S. and G.

Fig. 7. A plant growing in the membrane of Chaetomorpha californica and
nearing maturity. X 250.

[308]

UNIV. CALIF. PUBL. BOT. VOL. 7

[ SETCHELL-GARDNER ] PLATE 23

Or

0

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PLATE 24

Gomontia poJyrkiza (Lagerh.) B. and F.

Fig. 1. A group of three "sporangia," the two larger nearing maturity.
X 175.

Gomontia habrorJiiza S. and G.

Fig. 2. A young thallus. X 375.

Fig. 3. a-e, illustrating three forms of ' ' sporangia. ' '

PseudulveUa consociata S. and G.

Fig. 4. A surface view of a young thallus. X 375.

Fig. 5. A section of a mature thallus. X 375.

Fig. 6. A vertical filament near the surface of a young thallus showing

branching. X 225.

Entocladia codicoJa S. and G.

Fig. 7. (a) A young thallus, showing the method of branching of the
filaments and their radiating from a center. X 125.

Fig. 7. (6) A mature thallus with sporangia in the center. X 125.

Prasiola delicata S. and G.
Fig. 8. A surface view showing typical arrangement of cells. X 500.

[310]

UNIV. CALIF, PUBL. BOT. VOL. 7

[ SETCHELL-GARDNER ] PLATE 24

><s

8

6

^/^

PLATE 25

Prasiola delicata S. and G.

Fig. 1. A mierophotograph of a marginal segment, surface view showing
the arrangement of the vegetative cells. X 442.

Prasiola meridionalis S. and G.

Fig. 2. A mierophotograph of a portion of the surface, showing vegetative
cells and interspersed aplanospores (?). X 442.

[312]

UNIV. CALIF. PUBL. EOT. VOL. 7

[ SETCHELL-GARDNER ] PLATE 25

// c

I'LATE 26

Ulva dactyJifcra S. and G.

Fig. 1. Photograph of a whole plant, the type specimen, with the exception
of a jjortiou of the base. X 0.75.

Ulva stenophylla S. and G.

Fig. 2. A microphotograph of a portion of the surface, showing the rounded
angles and relatively thi( k walls of the cells. X 442.

[314]

UNIV. CALIF. PUBL. EOT. VOL. 7

[ SETCHELL-GARDNER ] PLATE 26

!

.^

X

V

PLATE 27
Ulva angusta S. and G.
A photograph of a group of plants, the type specimen. X 1.

31(1 J

UNIV. CALIF, PUBL. BOX. VOL. 7

[ SETCHELL-GARDNER ] PLATE 27

PLATE 28

Uha tacniata (Sotehel!) S. and G.

A photograph of a whole dried plant, showing the extreme crisped nature
and the dentate margins at the base. X 0.3.

[318]

UNIV. CALIF. PUBL. BOT. VOL. 7

[ SETCHELL-GARDNER ] PLATE 28

PLATE 29
Ulva stenophylla S. and G.
A photograph of the whole plant, the type specimen. X 0.3.

[320]

UNIV. CALIF. PUBL. EOT. VOL. 7 [ SETCHELL-GARDNER ] PLATE 29

PLATE 30
Monostroma areolatum S. and G.
A photograph of a whole dried plant, the type specimen. X 0.5.

[322]

UNIV. CALIF, PUBL. BOT. VOL. 7

[ SETCHELL-GARDNER ] PLATE 30

PLATE 31
Ulva angusta S. and G.
Fig. 1. A microphotograph of a part of the surface. X 442.

Monostroma areolatum S. and G.

Fig. 2. A microphotograph of a part of the surface. X 442.

The photographic work was all done by W. C. Matthews except plate 30,
which was done by E. W. Merrill. All the figures of plate 21, all but figure 5
of jilate 22 and figures 2, 3, 6, 7, and 8 of plate 24 were drawn by Dr. Helen M.
Gilkej'. All of plate 23 and figures 1, 4, and 5 of plate 24 were drawn by Miss
Almeda H. Nordyke.

[324]

UNIV. CALIF. PUBL. BOT. VOL. 7

f SETCHELL-GARDNER 1 PLATE 31

UNIVERSITY OF CALIFORNIA PUBLICATIONS

IN

BOTANY

Vol. 7, No. 10, pp. 325-331 December 29, 1920

PLANTAE MEXICANAE PURPUSIANAE, X

BY

TOWNSHEND STITH BEANDEGEE

LIBRARY

NEW YtfkX

. botanscal

QARijife;^

Hechtia glabra, sp. no v.

Undique glabra : scapo alto, pimiato ; ramulis patentibus multifloris,
inferioribiis ca. 2.3 dm. longis, 2-3 cm. distantibus : foliis gradatim
attenuatis margine siniiatis, spinis adscendentibus ca. 2 mm. longis
praeditis, supra nitentibus subtus multo lineati.s : bracteolis capsulas
subtendentibus parvis : capsnlis plerumque sessilibus interdnm pedi-
cellis ca. 7mm. longis; sepalorum tubo ca. 1mm. longo, lobis ovato-
lanceolatis ; petalis lanceolatis planis ca. 3 mm. longis quam sepalis
dnplo longioribns, basi sepalorum tubo adnatis ; capsulis ca. 5 mm.
longis apice stylos ca. 1.5 mm. longas recurvos gerentibus. Specimen
frugiferens solum visum.

The specimens are fragmentary. The upper parts of the leaves,
the only parts collected, are 5 dm. long. The panicle seems to be
pinnate but may be bipinnate and is at least 5 dm. high. The attach-
ment of the pedicellate capsules to their pedicels forms a right angle
and the pedicel is closely appre.ssed to the stem. Collected at Barranca
de Panoaya, Vera Cruz. No. 8506. Type, Herb. Univ. Calif. No.
204966.

Hechtia Purpusii, sp. nov.

Foliis lanceolatis, e basi ad apicem gradatim attenuatis, basi ca.
1.8 cm. latis, usque ad 20 cm. longis, utrinque lineatim peradpresse
lepidotis, pallidis, margine spinis minutissimis numerosis praeditis:
scapo 5-6 dm. alto, glabro : inflorescentia tripinnatim laxe paniculata,
multitlora, ramulis usque ad 13 cm. longi.s, pedicellis ca. 3 mm. longis,
quam bracteolis lineari-laneeolatis duplo longioribns; petalis oblongis
ca. 2 mm. longis, albis obtusis ; ovario glabro.

.— This Eccliiia differs from other species of the genus in having

cr> leaves of a thin texture margined with inimerous minute spines.
Collected in Barranca de Tenampa, Vera Cruz, growing on steep rocks.
^ No. 8420. Type, Herb. Univ. Calif. No. 200824.

326 Universitij of California Piihlicatio IIS ill Botany [Vol.7

Ocotea pyramidata lilake, s]). iiov.

Ai'hoi- -riMiulis; ranmli iii<;i-('scciitcs sparse ])ul)('nili <i-hil)rali snlcati;
t'.iiia ()l)l(iiip)-('llii)1ica lO-lSciii. loiiua 4-7.5 cm. lata obtusa i)asi
cuncato-roliiiKlala pcrgainriitacea sui»ra glabra lucida laxo ])r()miiml()-
reticulata subtns ol)s('uriiis viridia in costa et venis 8-10 prominentibns
et sparse in vennlis i)ubenda in axillis non barbata laxe proiniiiulo-
rcti.-ubita, in pctiolis sparse pnbernlis 1-2.5 em. longis; panieuhie
axilhires e ])asi ramosae pyramidatae snl)sessiles 0-12 em. longae ca.
6 em. latae sordide pnbernlae; pedieelli 1-3 mm. longi pnbernli ; ealyx
3 mm. longns snbglaber, segmcntis ovalibns .su])ae(|nalibns rotundatis;
slamiiia ser. I. 2.2 mm. longa glabra, filamentis antheras leyiter
emarginatas aeqnantibns, loeellis introrsis per paria snperpositis;
stamina ser. II. similia; stamina ser. III. basi glandulis binis magnis
tilamenti dimidinm aerjnantibiis sessilil)us donata, loeellis inferioribns
extrorso-laleralibns snperioribns intror.so-lateralibus; staminodia nulla ;
ovarium sterile subulatum glabrum 2 mm. longum.

This does not seem closely related to any other species known from
]\rexico. Type in the U. S. National Plerbarium, No. 1039029. collected
at Zacuapan, Vera Cruz. November. 1919 ; Purpus No. 8456.

Bauhinia {Casparia) jucunda, sp. nov.

Ramis glabris cinereis : foliis membranaeeis glabris 5-nerviis. laete
viridibus; foliolis ca. 5.5 cm. longis fere usque ad medium connatis,
partibus liberis ca. 2 cm. latis, gradatim in apices obtusos ca. 4 mm.
latos angustatis, apicibus ca. 6cm. separatis; petiolis ca. 2cm. longis;
stipulis parvis cito deciduis : calycis tubo ca. 7 mm. longo. limbo deinde
spathaceo : petalis linearibus tomentosis quam stamine perfecto brevi-
oribus. Fructus deest.

Only one specimen was collected. The thin, bright green, divaricate
leaves resemble those of some Passiflorae. Collected in Barranca de
Panoaya. Vera Cruz. No. 8535. Type, Herb. Cniv. Calif. No. 204997.

Indigofera acasonicae, sp. nov.

Frutieosa. ereeta : ramis cinereis pilis brevibus : foliis ereetis ses-
silibus vel petiolis uscjue ad 1cm. longis; foliolis 3-5 euneiformibus,
1.5-2.3 mm. longis, prope apicem ca. 5 mm. latis, apice rotundatis
mucronatis, ntrinqne breviter pilosis pilis medifixis; stipulis ovato-
lanceolatis: raeemis axillaribus ca. 2 cm. longis ca. 10-floris; bracteolis
parvis ovato-aeuminatis (|uam pedicellis duple longioribus; calycis
lobis quam tubo longioribus, ovato-aeuminatis ; legumine tereti ca.
2.5 cm. longo, pedicello curvato deflexo, ad raebin ramosrpie arte
adpresso.

1920] Brandegee: Plantae Mexicanae Purpimanae 327

The leaves are not spreading in the manner common to other species'
of the genus but stand upright close to the branches. The fruiting
racemes and deflexed legumes are closely appressed to the stem and
branches. Collected at Acasonica, Vera Cruz. No. 8510. Type,
Herb. Univ. Calif. No. 204971.

Eriosema nigropunctatum, sp. nov.

Erectum : caulibus pubescentibus ca. 2.3 dm. altis : foliis trifolio-
latis petiolis usque ad 2.3 cm. longis, foliolis lateralibus fere sessilibus,
foliolo terminali petiolulo ca. 8 mm. longo ; foliolis late ovatis longe
acuminatis. basi truncatis. subtus dense nigropunctatis, ad venas
pubescentibus. ca. 2.7 cm. longis, 2.2 cm. latis; stipulis lanceolatis
ciliatis : pedunculis axillaribus paucifloris ca. 5 cm. longis ; floribus ca.
6 mm. longis. stamine vexillari liberi; legumine oblongo ca. 2 cm.
longo, 2-spermo paululum hirsuto apice mucronato ; semine oblique
transverso funiculo ad extremitam liili linearis affixo.

The ovate leaves resinose punctate beneath give to this plant the
appearance of a Rhynchosia, but the position of the seed and its funicle
is that of Eriosema. Collected at Palmilla near Zacuapan, Vera Cruz.
No. 8386. Type, Herb. Univ. Calif: No. 200794.

Esenbeckia ovata, sp. nov.

Foliis digitatim trifoliolatis. foliolis glabris. ovatis, apice obtusis-
simis vel retusis, e medio in petiolum breve gradatim longe attmuatis.
pellucido-punctatis, usque ad 13 em. longis, 9 cm. latis ; petiolo puberulo
ca. 2 cm. longo : paniculis ca. 10 cm. longis, puberulis multifloris ;
petalis 5 imbricatis pellucido-punctatis; ovario 5-loculari tuberculato.
Fructus deest.

The leaves are broadly ovate or obovate, very obtuse and often
emarginate ; from the middle they are gradually attenuate into a short
petiolule. The petiolule of the usually larger terminal is longer than
the short ones of the lateral leaflets. Collected at Acasonica, Vera
Cruz. No. 8419. Type, Herb. Univ. Calif. No. 200823.

Euphorbia consoquitlae, sp nov.

Fruticosa, ramosa: caulibus tenuibus plus quam 6dm. altis: foliis
sparsis cito delapsis, cicatricibus pubescentibus : cymis in panieulam
ca. 2 cm. longam dispositis, bracteosis, bracteis vel foliis parvis anguste
ovatis vel lanceolatis usque ad 1 cm. longis; involucris paullo pube-
scentibus, axillaribus vel terminalibus, campanulatis; pedicellis ca.

328 University of California PuUications in Botanij [Vol. 7

1 cm. loiif^is; Idhis 4 obloiigis. t^Haiidiilis 4 planis orbicnlaribiis, appen-
(licc iis iiiiilto loM^iori, ca. 2 nun. loii^'a, apice ca. 1.5 mm. lata, basi
an^-us1i(n-i; l)i-acti'olis inter fiores mascnlos pubescent i bus; stylis t'ere
us(iiir ;i(l h.isiii (livisis; capsulae pedicellis ca. 3mm. longis, seminibns
globosis cinci-cis Inlx'i-culatis.

The si)ecimeii.s with the exception of a few leaf-like bracts of the
panicle are leafless. The stems are slender and branched. Collected
near Conso(initla, Vera Cruz. No. 8366. Type, Herb. Univ. Calif.
No. 204969.

Jatropha longepedunculata, sp. nov.

Jairopha wrr/i.s L. v;ir. longepedunculuta Brandg., Univ. Calif. Publ. Bot.,
vol. n, J). 368.

Arborescens ad 5 m. alta, undique estimulosa : f oliis latioribus quam
longis usque ad 20 cm. latis, 14 cm. longis, profunde 3-5-lobatis, ba.si
cordatis, lobis grosse dentatis; petiolis ad 20cm. longis: pedunculis
ad 23 cm. longis apice tiores in cymas dichotome ramosas dispositis
gerentibus; floribus feraineis petalis 5 liberis, ca. 5 mm. longis, sepalis
5 parvis connatis; stylis liberis apice laceratis; floribus maseulis
femineis similil)us. staminibus 2-verticillatis, exterioribus lilieris;
capsulis ca. 7 nnn. longis, seminibus carunculatis.

This plant was wrongly referred to Jatropha wrens L., from which
it differs in its separated petals, its minute, cupulate, dentate calyx,
lack of stinging bristles, and other characters. Collected, at Barranca
de Panoaya and Zacuapan, Vera Cruz. Nos. 7639, 7511, 8440.

Ayenia Purpusii, sp. nov.

Fruticulus : ramis juvenibus stellato-pubescentibus : foliis late
ovatis longe acuminatis, margine serratis, basi truncatis vel subeordatis.
supra glabris subtus stellato-pubescentibus uscjue ad 10 cm. longis,
9 cm. latis, petiolis ea. 3 em. longis, stipulis ca. 5 mm. longis: floribus
purpureis unibellatis, pedunculis ad axillas fasciculatis inaequalibus
quam petiolis brevioribus; sepalis ovato-acutis ca. 1.5 mm. longis;
petalorum laminis 2-partitis. lobis laciniatis, unguiculis filiformibus;
ovario sessili villoso, stigmate capitato; fructu immaturo processibus
linearibus ca. 2 mm. longis instructis.

The specimens appear to be branches of a bush. The young fruit
is covered with linear processes and is not muricate. Collected near
Zacuapan, Vera Cruz. No. 8378. Type, Herb. Univ. Calif. No. 200783.

1920] Brandegee: Plantae Mexicanae Purpiisianae 329

Maba Purpusii, sp. no v.

Frueticulus ? racemosus : ramis vetiistis glabris brunneis, junior-
ibns piibescentibus : foliis obovatis apice anguste obtiisis basi cuneatis,
margine integris minute ciliatis, utrinqne fere glabris, ca. 11.5 cm.
longis, 4.2 cm. latis ; petiolis ca. 3 mm. longis : floribus plerumque
solitariis axillaribus, pedicellis ca. 2 mm. longis ; calyce extns piibes-
centi campanulato, ca. 1 cm. longo, usque ad medium trifido, lobis
obtuse deltoideis; corolla tubulosa e calyce paululum exserta, apice
3-lobata, densissime f ulvo-hirsuta ; ovario hirsuto 6-loculare loculis
1-ovulatis. Fructus et flores masculi desunt.

Collected in Barranca de Panoaya, Vera Cruz. No. 8516. Type,
Herb. Univ. Calif. No. 204975.

•^ Sideroxylon campestre, sp. nov.

Arbor parva : ramis rugosis glabris : foliis ovato-oblongis utrinque
glabris basi longe cuneatis, apice acutis, usque ad 18 cm. longis, 4 cm.
latis. subtus venis prominulis; petiolis puberulis, ca. 2 cm. longis:
floribus uumerosis; pedicellis inter folios faseiculatis puberulis, ca.
1 cm. longis ; segmentis calycis 6 imbricatis orbicularibus, paululum
puberulis, ca. 6 mm. longis, exterioribus paulo minoribus ; corolla
tubulosa calycem paululum superanti, ad medium 6-lobata. lobis
oblongis apice retusis, ca, 4 mm. longis ; staminibus ad basin loborum
affixis, filamentis longitudine antheris oblongis acutis aequalibus, ca.
3 mm. longis; staminodiis linearibus integris, staminibus aequalibus, in
eadem serie alternis; ovario hirsuto 6-loculari. Baccae desunt.

The leaves with the fascicles of flowers are crowded about the ends
of the branches. Collected at Zacuapan, Vera Cruz. No. 8545. Type,
Herb. Univ. Calif. No. 204005.

Dictyanthus prostratus, sp. nov.

Hirsutus : caulibus pilis patentibus vel reflexis, prostrtais, basi
lignosis, usque ad 7 dm. longis : foliis late cordato-ovatis acuminatis,
utrinque hirsutis, ca. 2.2 em. longis, 2 cm. latis ; petiolis ca. 1 cm.
longis; cymis 2-floris in una axilla, pedunculis quam petiolis multo
brevioribus ; calycis lobis ovato-acuminatis, ca. 3 mm. longis ; corolla
late campanulata atropurpurea reticulata, ca. 7 mm. louga, 6 mm. lata,
lobis ovato-acuminatis. Folliculi desunt.

This species resembles closely D. parviflorus Hemsley, but the
corona is very different. In addition to the five narrow lamellate
scales adnate to the corolla there are five minute acute scales attached
to the middle of the gynostegium representing an inner corona.
Growing over rocks near Acasonica, Vera Cruz. No. 8411. Type,
Herb. Univ. Calif. No. 200816.

330 University of California Puhiications in Botany [Vol.7

Polystemma rupestre, sp. iiov.

Volnl)il(': cimliliiis liii-sutis plus (luam 8 dm. longis: foliis ovatu-
acuiiiiiialis. l»asi siiiii ohloiiyo apcrto, su[)ra liirsutulis, subtus ad venas
liirsiilis. iis(|uc ad S cm. loiigis, 8.3 cm. latis; petiolis ca. 2.5 cm. longis:
cymis ca. 8 cm. loiiuis. pauciHori.s : calyce liirsntis fere ad l)asin
iVlobatis. lobis laiiceolalis ca. 8 mm. loiigis, basi ca. 2 mm. latis : corolla
cami)aiiidata ca. 2 cm. longa, intus reticulata, alte lobata, lobis laneeo-
latis ca. 1 cm. longis; coronae sqiiamis gyiiostegiiim affixis, quorum
f) ligulalis pi'orimdc 3-lol)atis erectis. ad utrin(ine latus vel dorsum
lobi.s tiliformibus apicc glaudulam parvam gercntibus ct ligulas multo
supcrantibus; stigmate verticc piano ; staminura tubo brevi; polliniis
iis /'. r/nf////o/-i similibus ; folliculis juvenibu.s longe fusiformibus,
Icvibus.

Collected at Barranca de Panoaya, Vera Cruz, growing on "rocky
slopes." No. 8451. Type, Herb. Univ. Calif. 200855.

Pachystelma gen. nor. Asclepiadacearum

Calyx alte 5-fidus. Corolla cyathiformis 5-lobata. Coronae
squamae 5, partim corollae adnatae, earno.sae, tumidae, apice obtusae.
Gynostegium sessile, filamentorum tubo nullo. Pollinia in quoque
loculo solitaria fere pendula. Stigma vcrtice depressum, angulis valde
prominentibus. Cymae paucifiorae in una axilla pedunculatae. Flores
majusculi.

Pachystelma cordatum, sp. nov.

Volubile, frutescens : caulibus liirsutis pilis patentibus : foliis ovato-
acuminatis basi cordatis sinu aperto, usque ad 7 cm. longis. 5 cm. latis,
utriuque paulo breviterque hirsutis: cymis 2-8-ti()ris, pedunculis ca.
1cm. longis; calyce alte 5-lobata, lobis ovato-lanceolatis margine
ciliatis; corolla usque ad medium 5-lobata, ca. 6 mm. alta, 8 mm. lata;
coronae squami.s usque ad medium corollae adnatis, carnosis, crassis,
parte inferior! adnata quam parte superiori libera duplo crassa.
Folliculi desunt.

The long stems twining about themselves appear to have been
prostrate. The dry cup-shaped corolla is chartaceous and light colored.
The scales of the corona slightly surpass the stigma, they are erect,
rigid, thick, and about 1 mm. wide, at the apex are somewhat flattened
and obtuse ; the lower half adnate to the corolla is very much thickened.
The anthers at the angles of the depressed stigmatic part are very
prominent. Collected at Acasonica, Vera Cruz. No. 8008. Type,
Herb. Univ. Calif. No. 204968.

1920] Brandegce: Plantae Mexicanae Purpitsianae 331

Cynanchum racemosum, sp. nov.

Voliibile : eaulibus pubeseeiitibus : foliis ovato-acnminatis, ba.si
c'ordatis sinu aperto. lobis rotimdatis. utrinque fere glabris, majoribus
6 cm. longis, 4 cm. latis; petiolis 1.3-2.3 cm. longis: floribus circa
apices pedimculorum ca. 5 cm. longorum, breviter racemosis; pedicellis
ca. 4 mm. longis; calycis segmentis ovato-acntis ea. 3 mm. longis
corollam campanulatum superantilnis ; polliniis pendulis ; corona mem-
branacea cyathiformi alte 5-loba, lobis dentatis. ima basi tubi staminei
afifixa ; stigmate vertice piano.

Collected on rocky plains near Acasonica, Vera Cruz. No. 8378.
Type, Herb. Univ. Calif. No. 200783.

Solanum molestum, sp. nov.

Suffrutescens : ramis stellato-pubescentibns simul multis aculeis
rectis subiilatis patentibus armatis: foliis ovatis acutis, basi cordatis.
margine integris vel sinnatis. plerumqne solitariis. supra parce stellato-
pubescentibus olivaceo-viridibu.s, subtus dense albo-stellato-pube-
scentibus. usque ad 2 dm. longis. 12 cm. latis; petiolis 3-7 cm. longis.
dense stellato-pubescentibns interdum aculeos paucos gerentibus :
floribus lateralibus. pedunculis unifloris 1-3 fasciculatis stellato-
pubescentibns, ca. 1.5cm. longis; calyce alte 5-lobato, lobis ovato-
acutis. utrinque stellato-pubescentibus deinde reflexis, ca. 7 mm. longis ;
corolla calycis aequali usque ad basin 5-partitis; lobis oblongis apice
obtusis, extus dense stellato-pubescentibus, intus glabris brunneis;
antheris brunneis sessilibus apice poris dehiscentibus ; ovario setoso;
bacca densissime fulvo-setosa, diametro ca. 1.7 cm.

The fruit is covered with long yellow bristles and is more con-
spicuous than the colorless flowers. Collected in Barranca de Panoaya,
Vera Cruz. No. 8526. Type. Herb. Univ. Calif. No. 204992.

Viguiera pauciflora, sp. nov.

Herbacea : eaulibus strictis, pilosis plus quam 4 dm. altis : foliis
oppositis ovato-lanceolatis basi in petiolum ca. 6 mm. longTim cuneatis,
ca. 3.5 cm. longis. 1 cm. latis, utrinque paululum i)ilosis, margine
serratulis : capitulis plerumque 1 caules terminantil)us. quando 2
pedunculis ca. 14 cm. longis, diametro ca. 8 mm. ; phyllariis 3-seriatis
induratis ovato-acuminatis appresso-pilosis ; radiis luteis ca. 7 mm.
longis; pappi aristis 2; aquamellis fimbriatis; paleis glabris.

Stems slender, mostly bearing a single head. The specimens do not
show the basal part of the plant. Collected on plains near Acasonica,
Vera Cruz. No. 8375. Type, Herb. Univ. Calif. No. 200781.

Transmitted December 10, 1920.

UNIVERSITY OF CALIFORNIA PUBLICATIONS

IN

BOTANY

Vol. 7, No. 1 1, pp. 333-426, plates 32-49 May 16, 1922

PHYCOLOGICAL CONTRIBUTIONS

II TO VI

BY

WILLIAM ALBERT SETCHELL

AND

NATHANIEL LYON GARDNER

CONTENTS

PAGES

II.* New Species of Myrionema 334

III. New Species of Compsonema 353

IV. New Species of Hecatonema 377

V. New Species of Pylaiella and Streblonema 385

VI. New Species of Eetocarpus 403

* Phycological Contributions I appeared as No. 9 in the present volume of
this series.

334 Vniversily of California 'Publications in Botany [Vol.7

II. NEW SPECIES OF MYRIONEMA

Introduction

Critical study of an extensive series of material of small filamentous
species of Melanophyceae, mostly epiphytic upon larger Melanophyceae,
but in part epiphytic upon other algae and upon eef-grass, has brought
to light a rather surprisingly large number of little known or wholly
new species and forms on tlie Pacific coast of North America. Among
these is a series of similar organisms which we have grouped together
under the genus Myrionema, an account of which is presented in tliis
paper.

We are recognizing as the principal characters of this genus :
(1) a monostromatic basal disk, composed of closely crowded, short
celled filaments radiating from a common center, starting from a single
cell, each filament having terminal growth, and branching by radial
bifurcation of the terminal cell; (2) springing outward from practi-
cally all of the cells of the basal disk, except at the margin, are erect
filaments all of which may be fertile, bearing either zoosporangia or
gametangia, or a part of which may remain sterile; (3) the loculi of
the gametangia are uniseriate.

MjTrionema primarium sp. nov.

Plate 34, figure 12

Frondibus pulvinulos orbiculares, 0.5-1.25 mm. diam., formantibus;
parte prostrata filamentis regulariter radiantibus arete compactis.
formante ; filamentis erectis eramosis, dense stipatis, e cellula quaque
filamentorum repentium oriendis; pilis veris numerosis, per medium
frondis promisque sparsis; cellulis filamentorum erectorum 4-4.5(1
diam., inferne quadratis. cellula apicali juvena 2-3-plo longioribus;
cellulis pilorum verorum 4-5/x diam., 4-6-plo longioribus; zoosporangiis
ignotis; gametangiis in filamentis repentibus sessilibus per transfor-
matione filamenti toti quisque erecti oriendis, maxime stipatis, in medio
frondibus longitudine equalibus, prope niargines breviorilms. eylin-
dricis, 55-56//, longis. 5-()ju, latis, obtusis nut apice leviter attenuatis.
cellula apicali acute conica.

Growing on the outer end of the young blade of Costaria costata
in the lower littoral and the upper sublittoral belts. Oregon (Coos
Bay) to central California. Type, Gardner, no. 2764 (Herb. Univ.
Calif., no. 207012), mouth of Coos Bay, May.

1922] Setchell-Gardner : Phycological Coninhutions 335

We are considering Myrionema primarium to be a typical repre-
sentative of a group whose members are the most primitive of the
genus as considered from the standpoint of differentiation. The
prostrate basal layer is composed of long, regularly radiating filaments
with apical growth. Kadial divisions of apical cells occur just often
enough to completely occupy all of the space between each other as
the plant continues to increase in diameter and in circumference. The
branching is always dichotomous, and is accompanied by widening of
the apical cell, and the establishment of two growing regions on
opposite corners, which are subsequently separated from the remainder
of the cell by a wall, thus establishing two equal branches. Beginning
in the center, each cell successively toward the periphery gives rise by
horizontal divisions to erect filaments of nearly equal length, the only
exception or modification being that some cells give rise to long fila-
ments, the so-called true hairs, with the meristem at the base, and the
outer cells long and colorless. All other erect filaments are trans-
formed into gametangia. Only the cells of the basal filaments remain
sterile, or probably a few short erect filaments at the periphery of the
frond may never come to maturity. Thus starting with a single cell,
the maximum of reproductive cells arises in this group. Starting with
such forms the course of evolution seems to have been in the direction
of the sterilization of tissue.

Myrionema primarium f. acuminatum forma nov.

Plate 32, figure 9

Frondibus microscopicis, inter algas parvas alteras crescentibus ;
filamentis erectis pro parte sterilibus. SO-lOO/x longis, cellulis leviter
doliif ormibus ; gametangiis 4r-5jx diam., acuminatis, pro parte brevi-
pedicellatis.

Growing on Macrocystis pyrifcra. Carmel Bay, Monterey County,
California. Type, Gardner, no. 3110& (Herb. Univ. Calif., no. 207014),
December.

There is but a slight difference between M. primarium and forma
acuminatum. It is worthy of note that many gametangia in the center
of the thallus are short pedicellate (plate 32, fig. 9), and that a few,
about one in twenty-five, of the erect filaments continue to grow, attain-
ing a length of two to three times as long as the gametangia.

336 University of California Publications in Botany [Vol.

Myrionema minutissimum sp. nov.

Plate 34, figures 1-3

Froiulil)us pulviiiulos iiicoiispicnos. 0.75-1.5 mm. diam.. for-
maiilibus; parte prostrata filamentis dense compactis, re^ulariter
radiantilnis eomposita; filamentis erectis dense stipatis, eylindricis,
omnibus fruetiferis; pilis veris ignotis; eellulis filamentorum repentium
4/u, diam., in centre frondis, quadratis, ad margines leviter longioribus
quam latis; eellulis filamentorum erectorum sirailibus iis repentium;
zoosporangiis et "aseocvstis" ignotis; gametangiis pedieellis 4-8-
cellulis eompositis sufifultis, eylindricis, dense stipatis, 34-40|Li longis,
4-4.5/i, latis.

Growing on the cysts of Egrcgia Menziesii. Cast ashore near the
"ClifY House," San Francisco, California. Type, Gardner, no. 4123&
(Herb. Univ. Calif., no. 207020), November.

Myrionema minutissimum may be distinguished by the absence of
sterile erect filaments and hairs, and by the small sized, very densely
crowded pedicellate gametangia. Thus far no "ascocysts" nor zoo-
sporangia have been observed. In its phylogenetic relationship it would
seem to stand equally close to M. corunnae and M. foecundum, differ-
ing from each, however, in details of measurements. The erect fila-
ments are of the same diameter at their bases as the length of the cells
in the creeping filaments. Since the lower part of each remains sterile,
it gives the mass of cells the appearance of being parenchymatous.

Myrionema foecundum f. simplicissimum forma nov.

Plate 33, figures 9', 10

Frondibus pulvinulos orbiculares, 0.5-1.5 mm. diam. formantibus;
parte prostrata filamentis regulariter radiantibus et arete adhaeran-
tibus augmentatione ramificationeque apicali eomposita ; filamentis
erectis omnibus fruetiferis; pilis veris ignotis; eellulis filamentorum
repentium 3-4. 5|a diam., medio quadratis, prope marginem 2-plo
longioribus; zoosporangiis ignotis; "ascoeystis" raris usque ad numer-
osis, elavatis 28-34ju, altis; gametangiis transformatione filamenti omnis
quisque erectiis marginis frondis exceptis oriendis, 30-40/x altis. 7-9/i
latis.

Growing on the sporophylls of Nereocystis Luetkcana. Puget
Sound, Washington, to central California. Type, Gardner, no. 4316
(Herb. Univ. Calif., no. 207022), Moss Beach, San Mateo County,
California, July.

We have not seen either the type or any other authentic material
of Phycocelis foccunda Stroemfelt. Neither have we anv authentic

1922] Setchell-Gardner : Phycological Contributions 337

measurements of the original. De-Toni (1895, p. 582) gives 35/* to 70/x
as the height and 7,/x to 12/x as the width of the gametangia. In height
of gametangia our form approximates to forma seriata Reinke (1889,
p. 46) , but has no true hairs so far as we have ever been able to observe
from the examination of many specimens from different localities.

At times the individuals are very numerous on the host, so much so
as to interfere with each other's growth. In one collection (Gardner,
no. 4651) they are completely confluent over several square inches, and
the outline of the individuals can be detected only by staining and hy
microscopic examination.

Myrionema foecundum f. subulatum forma nov.

Plate 33, figures 1-3

Filamentis basalibus per ramulos numerosos, subulatos, 2-4 cellulis
compositos, inter cellulas hospitis penetrantes annectis; zoosporangiis,
"ascocystis" et pilis veris ignotis.

Growing on the sporophylls of Nereocystis Luctkeana. Near the
"Cliff House," San Francisco, California. Type, Gardner, no. 4651
(Herb. Univ. Calif., no. 207028), November.

Forma subulatum is very similar to forma simplicissimiim. The
gametangia average slightly larger, and are usually more blunt. There
are no so-called "ascocysts" present. The chief distinction is the
presence in this form of numerous awl-shaped rhizoids, found in no
other form of the species, so far as we have been able to ascertain.

Myrionema foecundum f . ramulosum forma nov.
Plate 33, figure 8

Frondibus pulvinulos parvos, orbiculares, 1-3 mm. diam. forman-
tibus ; filamentis erectis superne sparse ramosis, 55-65/a longis ; cellulis
filamentorum erectorum 5.5-6.5/t diam. quadratis, cellulis terminalibus
frequenter 2-3-plo longioribus quam latis; zoosporangiis, "asco-
cystiis, ' ' et pilis veris ignotis ; filamentis erectis omnibus in gametangia
transformatis.

Growing on the pneumatocysts of Nereocystis Luetkeana. Friday
Harbor, San Juan Island, Washington. Type, Gardner, no. 4065
(Herb. Univ. Calif., no. 207018), August.

This form of 31. foecundum differs from the other forms of the
species in having secund "proliferations" near to the outer ends of-
many of the gametangia.

338 Vniversiti) of California Puhlications in Botany ["^^o^'

Myrionema foecundum f. divergens forma iiov.

Plate 34, figures 8, 9

Froiulibus pulvinulos orbiculares, 1.5-3.5 mm. diam. formantibns;
parte ha.sali filaiiK'iitis dense eompactis. eoiijnnctim et ad siipcrficiem
hospitis aivte adhaereiile ; filamentis sterilibus ereetis exij;iis, cramosis,
obtusis, promisqne inter gametangiis sparsis, 300-400/x, longis; pilis
veris denntibus; "ascocystis" niimerosis, inter, et brevioribns qnam,
prametancfiis. sparsis. cylindrieis nsqiie ad elavatis; eollnlis filamen-
tornm repentium 5-Gja diam., 1.5-2-plo longioril)us; ccllulis filamcn-
torum erectorum 8-9/x diam.. 1.5-3-plo longioribus, cylindrieis, non
constrietis. zoosporangiis ignotis; gametangiis forma varialiilibus,
cylindrieis. eylindro-eonicis usque ad fnsiformibiis, 50-60/x longis.
7.5-8.5;u, latis; loculis 1-2-seriatis.

Growing on the sporophylls of Nereocystis Luetkeana. Pacific
Grove. California. Type, Gardner, no. 4511 (Herb. T^niv. Calif., no.
207024), December.

This form of 3/. foecundum manifests some slight differentiations
not present in any other form of the species. Scattered promiscuously
over the frond are erect filaments two to three times as long as the
gametaugia. The "ascoeysts" are abundant. Some irregularities are
also shown in the creeping filaments. Typically the branching seems
to take place by the splitting of the apical cell, but it seems that
frequently one of the dichotomy fails to develop till later, producing
the appearance of lateral branching. There is also an occasional
oblique or longitudinal division of a cell of the creeping filaments
perpendicular to the host.

M5nnonema foecundum f. majus forma nov.

Plate 32, figure 7

Frondibus pulvinulos orbiculares, 1-2 mm. diam.. superficie super-
iore convexa ; parte prostrata filamentis arete adhaerantibus. regu-
lariter radiantibus composita : filamentis ereetis dense stipatis, sim-
plicibus, cellulis cylindrieis, 5-7ju, diam. ; pilis veris sparsis, per f rondem
promiscue distributis, basim meristematicis, vaginatis, 400-500// longis,
cellulis 6// diam. apice, 5-8-plo longioribus; "ascocystis" in filamentis
prostratis ses.silibus, cylindrieis, apice leviter distentis, 45-55/i, longis,
8-12yLt latis, nunc in fronde tota sparsis nunc in gregibus isolatis;
znosporangiis ignotis; gametangiis m filamentis basalibus se.ssilibus,
cylindrieis, 60-80ju, longis, 6-8/i, latis, per transformationem quisque toti
filamenti oriendis, dissepimentis horizontalibus aut obliquis.

Growing on Laminaria ephemera Setchell. Port Renfrew, Van-
couver Island, Neah Bay, Puget Sound (Tacoma) and central Cali-
fornia (Carmel Bay). Type, Gardner, no. 2910 (Herb. Univ. Calif.,
no. 207013^. Carmel Bay, California, May.

1922] Setchell-Gardner : Phycological Contributions 339

This small epiphyte seems to be restricted in its habitat to the
above named host. The only known localities in which the host has
been observed are those mentioned above. The material from each
locality is richly infested by this epiphyte. The host, richly infested,
was distributed by Miss Josephine Tilden in her American Algae
(Exsicc), no. 609, under Eenfrewia parvula Griggs.

It seems very closely related to Mi/rionema foecundum f . simplicis-
sivmm. The differences are in the larger measurements of all of the
parts, the presence of numerous, larger, so-called ascocysts, occurring
in groups (see plate 32, figure 7), or scattered among the gametangia,
in the presence of well developed hairs, and in the more convex upper
surface of the mass of erect filaments.

Myrionema corunnae f. uniforme forma nov.

Plate 33, figures 4, 5

Frondibus pulvinulos irregulares 0.5-1.5 mm. diam. formant-
ibus; filamentis repentibus stratum compactum augmentatione
peripherica efficientibus ; pilis veris deuntibus ; filamentis erectis
eramosis. apice leviter attenuatis, SO-lOO/x (usque ad 120ju,) altis;
cellulis filamentorum erectorum 4.5-5.5/a diam., 1-1.5-plo longioribus;
"ascocystis" sparsis, clavatis; gametangiis stratum solidum com-
pactum, ordine palorum, centro altitudine aequalibus, ad margines
gradatim diminuendis, plerumque sessilibus, nunc pedicellis 1-3 cell-
ulis compositis suffultis, SO-lOO^u, altis, 6-6. 5/x latis; loculis uniseriatis.

Growing on the blades of Costaria costata, Liminaria Sinclairii, and
Alariu marginata. Central California (Moss Beach in San i\Iateo
County, Fort Point in San Francisco). Type, Gardner, no. 4473
(Herb. Univ. Calif., no. 207023), San Francisco, July.

Forma uniforme differs from the species as described and figured
by Sauvageau (1897, pp. 77-82, repr., figs. 14 A-F) in having no hair
filaments, in having no branched gametangia, in having "ascocysts,"
and in having slightly different dimensions.

There are some differences in the size of the plants we have found
growing on different hosts in central California. The plants chosen
as the type of the form grow on the blades of Alaria marginata at
Fort Point, San Francisco. Very generally, this species of Alaria has
delicate grooves obliquely radiating from the midrib to the margin.
The presence of these grooves is often accentuated by the growth of
this minute Myrionema, usually so numerous as to be confluent, so that
the individuals cannot be detected except by microscopic examination.

340 University of California Publications in Botany [Vol. 7

Myrionema corunnae f. angulatum forma nov.

I'late 33, figures 6, 7

Prondibn.s pulviiinlos microscopicos linea extoriore irregularibus,
ill hopites inter algas alteras parvas, f omiantibus ; filamontis repen-
tibus sub conditiones augmentation is liberae radiantibus; filamcntis
erectis eramosis. dense stipatis, 130-140|U. longis; pilis veris dcuntilius;
eellulis filamentorum repentium 4-4. 5/x diam., plerum(|ue quadratis;
cellulis filamentorum ereetorum cylindricis, 4-4.5/x, diam., 1.5-3-plo
longioribus; zoosporangiis et "ascocystis" ignotis; gametangiis cylin-
drieis, apiee leviter attenuatis, in pedicellis cellulis 6-10 compositis
suffultis, 35-45/A (usque ad OO/x) lougis, 4-6. 5/i. latis, septorum multis
obliquis.

Growing on the cysts of Egregia Menziesii. Cast ashore near the
"Cliff House," San Francisco, California. Type, Gardner, no. 3471&
(Herb. I'niv. Calif., no. 207015), August.

This form is to be distinguished from the species as described by
Sauvageau (1897, pp. 77-82, repr.) by the absence of hairs, the presence
of sterile erect filaments, the gametangia mostly on longer pedicels and
by the very small .size of the loculi, often produced by perpendicular
and slanting walls. In part, the gametangia seem pluriseriate, caused
by the division of some of the original cells into small, angular loculi.

Myrionema corunnae f. sterile forma uov.
Plate 33, figures 13-] 5

Frondibus pulvinulos orbiculares 1.5-3 mm. diam. formantibus;
filamentis erectis rectis. simplicibus. cylindricis, obtusis, 55-65/a longis;
pilis veris et "ascocystiis" deuntibus; gametangiis cylindricis. obtusis,
in pedicellis cellulis 3-5 compositis suffultis. raro sessilibus, 55-65^
longis, 4.5-5. 5;a latis; gametangiis et filamentis erectis sterilibus e
centre ad margines frondis longitudine regulariter gradatimque
diminuendis.

Growing on the pneumatocysts of Xereocystis Luetl-eana. Cast
ashore. Carmel Bay, Monterey County, California. Type, Gardner,
no. 4557 (Herb. Univ. Calif., no. 207027), December.

This form is to be distinguished from the species as described by
Sauvageau (1897. pp. 77-82, repr.) by the presence of numerous sterile
erect filaments interspersed among the gametangia, about one-fourth
to one-fifth as many as the gametangia, in the absence of hairs, and in
the very gradual diminution in the length of the sterile filaments and
the gametangia from the center to the circumference of the fronds.
The sterile erect filaments are of the same length as the gametangia.

i922] Setchell-Gardner : PJujcological Contnhutions 341

Mjrrionema balticum f. pedicellatum forma nov.

Plate 33, figures 11, 12

Frondibiis pulvimilos mieroseopieos 150-400/x (usque ad 800/x)
diam., linea exteriore maxime irregiilares formantibiis ; filamentis
ereetis pro parte brevi pilif eris, 140-160^ longis ; pilis veris ignotis ;
zoosporangiis( ?) angiiste clavatis, in filamentis repentibiis sessilibus
aiit in filamentis ereetis terminalibiis ; gametangiis in pedicellis cellulis
10-15^u compositis suffnltis terminalibus aut sessilibus, usque ad 150/*
longis.

Growing on the outer ends of the leaves of Phyllospadix sp. Point
Carmel, Monterey County, California. Type, Setchell, no. 5417 (Herb.
Univ. Calif., no. 207030), June.

This form differs from the species in the absence of hairs, in having
a few very short-piliferous, vegetative filaments, and in having the
gametangia on long pedicels.

In this form, as in many others, the correct interpretation of the
zoosporangia mentioned above in the diagnosis is an open question.
They are filled with dense cell contents, and, although none have been
observed to produce zoospores, they appear as though they might later.
Whatever their nature may finally prove to be. they represent another
character in which this form differs from the species. Reinke neither
figures nor describes them in Ascocyclus halticus. They appear on
the same individuals with the gametangia or on different individuals.

Myrionema balticum f . californicum forma nov.

Frondibus pulviniTlos microscopicos. 0.5-0.75 mm. diam. ; parte
prostrata filamentis relative rectis, regulariter radiantibus composita ;
filamentis ereetis simplicibus. et superne et infeme leviter attenuatis,
obtusis. 75-125/A longis; pilis veris 0.5-1 mm. longis; cellulis fila-
mentorum prostratorum 6-7/* diam., 1-1.5-plo longioribus: cellulis
filamentorum erectorum 7.5-10/i diam., quadratis; cellulis pilorum
verorum 4-5. 5/t diam., superne 6-10-plo. longioribus, basi 3-5-plo
longioribus, sed cellulis paucis basim superne remotis quadratis;
chromatophoris taeniatis irregulariter interruptis ^ compositis ; zoo-
sporangiis ignotis; gametangiis pedicellis 1-3 cellulis compositis
suffultis, aut e filamentis repentibus aut e basibus filamentorum erec-
torum lateralibus oriendis, cylindricis, 90-130/* longis, 7-9/* latis.

Growing on the outer ends of the leaves of Phyllospadix sp. Point
Carmel, Monterey County. California. Type, Setchell, no. 5439 (Herb.
Univ. Calif., no. 207031), June.

We have no authentic measurements of Ascocyclus halticus Reinke.
He does not mention the size of the gametangia either in the Atlas

342 TJniversitij of California Publications in Botany l^^ou 7

(1889, })!. 16, figs. 1-4), where he first describes and figures it. or in
the Algenfiora (1889a, p. 45) where he describes it again. Foslie
(1894, p. 17, repr.) made the combination Myrionema (Phycocelis)
halticum, but he gave no measurements for any of its parts. De-Toni
(1895, p. 581) lists the plant as Phycocelis(f) haltica (Reinke) Foslie,
but gives no additional information concerning the size of the game-
tangia. Kylin (1907, p. 35) lists a plant under Myrionema halticum
(Rke.) Fosl. which he found growing on Delesseria sangiiinea at 15 M.
depth. lie states that the assimilation filaments are 100-125)li long
and 4-6/x broad, but gives no measurements for the gametangia.

The plant which we have diagnosed here is doubtless very closely
related to Reinke 's Ascocyclus halticus. It may be a distinct species,
but we are listing it as a form of halticus until we can obtain more
knowledge of its variation, and more authentic information concerning
the type material of halticus. Tlie particular distinguishing character
is the very frequent occurrence of lateral gametangia arising very
close to the base of the vegetative filaments.

Myrionema globosum f. affine forma nov. .

Plate 33, figure 16

Frondibus microscopicis, 200-300/i, diam., linea exteriore irrecru-
laribus, algis alteris parvis immixtis; filamentis prostratis multo dis-
tortis, libera ramosis; filamentis erectis fasciculato-ramosis, 100-140//
longis, superne attenuatis, pro parte piliferis; pilis veris 300-400/^
longis; cellulis filamentorum prostratorum 4-6/i diam., forma varia-
bilibus; cellulis filamentorum erectorum cylindrieis, 4-5ju, diam.; zoo-
sporangiis ignotis; gametangiis cylindrieis, 35-45/x longis, 4— 4.5ju, latis,
in filamentis erectis sessilibus lateralibusque aut raro terminalibus;
loeulis uniseriatis.

Growing on the leaves of eel-grass. Sitka, Alaska, to central Cali-
fornia (Point Carmel. Monterey County). Type, Setchell. no. 5439a
(Herb. Univ. Calif., no. 207032), Point Carmel, June.

"We have no measurements in detail for the parts of Myrionema
glohosnm, this seemingly distinct species. Reinke (1889a, p. 46) states
that the plants are one-half to one millimeter in diameter. Our plants
seem much smaller, in general. It is difficult to make out the size on
account of their being so intimately intermixed with other small algae.
This is the case in both collections of plants w^hicli we have examined,
one from Sitka, Alaska, and the other from Point Carmel, California.
We are placing it as a form of M. glohosnm (Reinke) Foslie, awaiting
knowledge of further details concerning the type.

1922] Setchell-Gardner : Phycological Contributions 343

Myrionema compsonematoides sp. nov.

Plate 34, figure 10

Frondibus pulvinulos microscopicos linea exteriore irregulares,
200-400jU, diam. f ormantibus ; parte prostrata filamentis distortis com-
posita; filamentis erectis cyliiidricis, simplicibus TO-SO/a altis, pilis
veris ignotis; cellulis filamentorum repentium forma magnitudineque
aliquantum irregularibus, 4-6ju, diam. ; cellulis filamentorum ereetorum
cylindricis, 5-1 fx diam., quadratis; "ascocystis" sparses, terminalibus
aut prope basim filamentorum ereetorum lateralibus; gametangiis
fusiformibus usque ad clavatis, sessiJibus aut brevi-pedicellatis, 50-65/x
longis, 7-10/x latis; loculis inferne uniseriatis, superne biseriatis.

Growing on the blade of Laminaria complanata. Friday Harbor,
Washington. Type, Gardner, no. 4118 (Herb. Univ. Calif., no. 207019),
August.

Myrionema compsonematoides seems to be a modification of M.
foecundum. There is a slight sterilization of tissue in that a few of the
erect filaments remain sterile and develop slightly beyond the game-
tangia, and in that a part of the gametangia are pedicellate. It seems
to tend toward Compsonema in these respects, and in having biseriate
gametangia in part.

Myrionema hecatonematoides sp. nov.

Plate 34, figure 11

Frondibus pulvinulos tenues dilute fuscos, linea exteriore irregu-
lares, usqve ad 6 mm. diam. f ormantibus; parte prostrata filamentis
distortis, irregulariter radiantibus composita, plerumque monostro-
matica, pro parte distromatiea, inferne ramulos radiciformes pene-
trantes, paxilliformes, paucos emittente; filamentis erectis simplicibus,
cylindricis, 240-260/1, longis; pilis veris deuntibus; cellulis filamen-
torum ereetorum 6.5-7.5ju, diam., 2-3-plo longioribus; zoosporangiis
ignotis ; gametangiis sessilibus aut in pedicellis 1-2-cellulis compositis
e filamentis repentibus oriundis suffultis, cylindricis usque ad fusi-
formibus, 45-55/A longis, 7-8. 5ja latis ; loculis plerumque uniseriatis, in
parte latiore gametangiorum biseriatis.

Growing on the pneumatocysts of Nereocystis Luetkeana. Pacific
Grove, California. Type, Gardner, no. 4533 (Herb. Univ. Calif., no.
207025), December.

As indicated by the partial distromatic base, the slight tendency to
biseriate loculi in the gametangia and the sterilization of a part of the
erect filaments, this species of Myrionema seems to be differentiating
in the direction of the genus Hecatonema. We are retaining it in the
genus Myrionema because of a preponderance in number of uniseriate
loculi and monostromatic filaments in the prostrate portion.

344 University of California Publications in Botany [You

Myrionema atteniiatum sp. no v.

Plate 34, figures 6, 7

Froiidilius strata mioroscopica pins miinisve oonflncntia in snpcr-
ficic liospitis formantilnis; ])arte prostrata filamcntis. reo^ulariter
radianlibus coniiiosita ; (ilanicntis criM-tis siinplicil)ns supcrne basinuine
leviter attenuatis, 180-220/a longis, pilis veris deuntibus; cell.nlis
filanientornm repontium 4.5-5.5/i. diam.. proximo qnadratis; ccllnlis
filamentornm ereetorum 6-7/a diam., lonj,ntu(lin(' variabilibus; zoo-
spoi-anjriis ignotis; gametangiis sparsis, cylindricis usque ad leviter
fusiformibus in pedieellis longiorilms aut brevioribus, 30-45/x longis,
4.5-6;u, latis.

Growing on the stipes of Macrocystis pyrifera, Laminaria Farlowii,
and on the sterile base of Gigartina radula f. Carmel Bay, Monterey
County, California. Type, Gardner, no. 4685a (Herb. Univ. Calif.,
no. 207029), December.

It seems that sterilization has proceeded in this species of Myrio-
nema until about two-thirds of the erect filaments no longer produce
gametangia, and these filaments are several times longer than the
gametangia. Tlie cells are slightly doliiform in some collections.

Myrionema attenuatum f. doliiforme forma nov.

Plate 34, figures 4, 5

Frondibus floeeulos parvos, irregulares, inter algas alteras epi-
phytieas formantibus; filamentis erectis usque ad 275ia longis ; cellulis
filamentorum ereetorum quadratis usque ad 8-plo longioribus quara
latis hand dubie doliiformibus ; gametangiis nunc parvis et in fila-
mentis longis erectis terminalibus nunc ut in forma species typica.

Growing on the stipe of Macrocystis pyrifera. Carmel Bay,
IMonterey County, California. Type, Gardner, no. 4540a (Herb.
Univ. Calif., no. 207026), December.

The chief distinction between this form and the species, M. attenu-
alum, is the decided doliiform character of the erect filaments. Small
terminal gametangia appear very frequently on the long, erect fila-
ments.

Myrionema phyllophilum sp. nov.

Plate 32, figures 1-5

Frondibus pulvinulos plus minusve orhiculares, 400-800;u, diam.
formantibus; parte repente filamentis rectis e centro radiantibus. dense
compactis composita ; filamentis erectis simplicibus, cylindricis. basim
constrictis, 11()-130/a longis, pro parte superne leviter attenuatis
piliferisque; pilis veris inter filamenta gametangia<iue promiscue

1922] Setchell-Gardner : Phycological Contributions 345

sparsis; cellulis filamentoruni repentium cylindrieis, 4-5/a diam.,
2-4-plo longioribus; cellulis filamentorum erectorum 8-9/i, diam.,
quadratis usque ad 2.5-plo longioribus; cellulis pilorum verorum basim
quadratis, superne 5-6/i, diam. et usque ad 20-plo longioribus; zoo-
sporangiis( ?) late clavatis, sessilibus aut in pedicellis longioribus aut
brevibus suffultis, 50-70/i, longis, 14-18/i, superne latis; gametangiis
cylindrieis, plerumque in pedicellis 1-2-cellulis compositis suffultis in
filamentis repentibus aut rariore in basi filamentorum erectorum
lateralibus. obtusis, 90-1.30/x longis, 9-llfx latis ; loeulis uniseriatis.

Growing on the outer ends of the blades of eel-grass, in the lower
littoral and upper sublittoral belts. Sitka, Alaska. Type. Gardner,
no. 3969 (Herb. Univ. Calif., no. 207017), July.

In this species about one-third of the erect filaments remain sterile,
and extend beyond the gametangia. The cells in the creeping filaments
are relatively long. The filaments are closely crowded, but distinct and
readily separable. All the erect filaments are abruptly narrowed at
the base. On account of the long creeping cells, the erect filaments are
very much less crowded than is usually the case in most Mtjrionemas.
The erect filaments do not arise successively toward the margin of the
nearly mature plants. Cells here and there develop an erect filament
to the length of several cells before the intervening cells show signs
of farther growth. The gametangia are typical of the genus, with
uniseriate loculi and with mostly horizontal, cross cell-walls. The
zoosporangia( ?) are fairly abundant, and are either with the game-
tangia on the same individual or on separate individuals. We question
these structures as being functional since none of them has been
observed to produce zoospores. They have the appearance of being
abortive organs, either zoosporangia or gametangia, more likely the
latter, and are probably like the organs observed by ]\Iagnus which
induced him to establish the genus Ascocydus.

Myrioenma setiferum sp. nov.

Plate 32, figure 8

Frondibus flocculos aut pulviuulos parvos. densos, irregulares,
0.5-0.75 mm. diam. f ormantibus ; parte prostrata filamentis regulariter
radiantibus, dense compactis, cellulis -1— 5/x diam., prope margines
1.5-3-plo longioribus composita; pilis veris ignotis; filamentis erectis
numerosis. simplicibus. apice leviter attenuatis, longe-piliferis, 700-
800/i longis; cellulis filamentorum erectorum cylindrieis, 4— 5/x diam.,
2-3-plo longioribus; zoosporangiis late clavatis, 80-90ja longis, 18-24/x
latis; gametangiis cylindrieis. 150-200;Lt longis, 7-8. 5/a latis, sessilibus
aut in pedicellis curtis in filamentis erectis aut raro in filamentis
repentibus suffultis ; loeulis uniseriatis.

346 University of California Publications in Botany [You i

Growing on the outer ends of the leaves of eel-grass, in company
witli other small algae, in the lower littoral belt. Sitka, Alaska. Type,
Gardner, no. 89G8c (Herb. Univ. Calif., no. 207016), July.

M\irwncm,a sctiferum is undoubtedly very closely related to M.
phyllophihun on the one hand, and to M. foecundum f. californicum
on the other. From the former, it differs in having longer and nar-
rower erect filaments with longer cells and in having much longer and
narrower gametangia which are more frequently borne laterally on the
erect filaments, and often fasciculately branched from a short pedicel.
From the latter, it differs in having functional zoosporangia, one of
the few instances in which we have actually seen the undoubted
zoospores, in having longer and setiferous erect filaments, and longer
and slightly narrower gametangia.

Myrionema obscurum sp. no v.

Plate 32, figure 6

Frondibus microscopicis, supra superfieiem hospitis irregulariter
dispersis; filamentis repentibus multo ramosis, maxime compaetis et
contortis, stratum pseudo-parenchyinaticum formantibus ; filamentis
erectis sparsis, simplicibus, 40-70/x altis; pilis veris ignotis; eellulis
filamentorum repentium forma magnitudineque maxime irregularibus,
4.5-10/A diam., eellulis filamentorum erectorum cylindricis, 6.5-7.5/x
diam., exacte aut fere quadratis; zoosporangiis numerisissimis. in
filamentis repentibus, sessilibus, ovoideis usque ad late clavatis, 45-65/*
longis, 18-30/i, latis; gametangiis ignotis.

Growing on the blade of Costaria costata. Moss Beach, San ^Mateo
County, California. Type, Gardner, no. 4315 (Herb. Univ. Calif.,
no. 207021), July.

Myrionema obscurum seems decidedly distinct from all other species
of the genus. It seems to be a near relative of M. strangulans Grev.
The basal layer is very compact, and adheres very firmly to the host,
following closely the irregularities on its surface. We are unable to
make out whether or not the epiphyte causes the death of tlie cells of
tlic liost, which seems to be quite frequent in all of the plants we have
observed. Tlie creeping filaments push among the dead cells of the
host at times. The zoosporangia are numerous and all sessile.

1922] Setchell-Gardner : Phycological Contributions 347

LITERATURE CITED

De-Toxi, J. B. (G. B.)

1895. Sylloge Algarum, vol. 3, Fucoideae.

FOSLIE, M.

1894. New or critical Norwegian Algae. Det. Kgl. norske Videnskabers
Selskabs. Skrifter. Trondlijem.

Kylin, H.

1907. Studien ueber die Algenflora der schwedischen Westkiiste. Akad-
emische Abhandlung. Upsala.

Eeinke, J.

1889. Atlas deutscher Meeresalgen, Part I. Berlin.

1889a. Algenflora der westlichen Ostsee deutschen Antheils. Sexter Bericht

der Kommission zur wissenschaftlichen untersuchung der deutschen

Meere in Kiel. I. Heft. Berlin.

Sauvageau, C.

1897. Sur quelques Myrionemacees. Ann. sci. nat. 8 ser., Bot., vol. 5, pp. 161-

288.

PLATE 32

Myrioncma phyllophilum S. and G.

Fig. 1. A single creeping filament near the margin of a frond showing
zoosporangia(?) and 3'oung erect filaments. X 2.j0.

Fig. '2. The same as figure 1, but illustrating the gametangia. X 250.

Fig. 3. A fragment from the center of a mature plant showing a true hair,
mature erect filaments with terminal hairs, and gametangia. X 125.

Fig. 4. The same as figure 3, but illustrating sessile zoosporangia( ?). X 125.

Fig. 5. A segment of the basal portion showing the coni])aet nature of the
radiating filaments. X 125.

Myrionema obscurum S. and G.
Fig. 6. A few fragments of typical plants. X 125.

Myrionema foecundiim f. majus S. and G.

Fig. 7. A terminal part of a creeping filament showing a group of so-called
ascocysts, true hairs, and typical gametangia. X 250.

Myrionema setiferum S. and G.
Fig. 8. A fragment of a tyi)ieal jilant. X 125.

Myrionema primarium f. acuminatum S. and G.
Fig. 9. Fragments taken from different parts of a plant. X 250.

[348]

UNIV. CALIF, PUBL. BOT, VOL. 7

[SETCHELL-GARDNER ] PLATE 32

10QQpOQ^2QmQ

CO^P^^^^W

PLATE 33
Myrionema foecundum f. subulatum S. and G.

FifT. 1. A section through a plant and its host perpendicular to the surface,
tnlvoii near tlio iiiarLriii. X 125.

Fig. 2. Same as figure 1. X '2.~)0.

Fig. 3. A segment of the base at the margin on the surface of the host.
X 125.

Myrionema corunnae f. uniforme S. and G.

Fig. 4. Three fragments of plants showing the base and typical gametangia.
X 250.

Fig. 5. Segments of the base, diagrammatic.

Myrionema corunnae f. angulatum S. and G.

Fig. 6. A part of a frond taken near the center of the frond. X 250.
Fig. 7. A part of the frond taken near the circumference of the frond.
X 250.

Myrionema foecundum f. ramulosum S. and G.

Fig. 8. Two fragments of fronds showing the base and tj'pical gametangia.
X 125.

Myrionema foecundum f. simplicissimum S. and G.

Fig. 9. A section through a frond near its margin, and its host perpendicular
to the surface showing the base, the sessile gametangia and the so-called asco-
cysts. X 125.

Fig. 10. A section showing the "ascoeysts" grouped. X 250.

Myrionema haltieum f. pedicellatum S. and G.

Fig. 11. A group of typical gametangia and erect filaments from the basal
layer. X 125.

Fig. 12. Same as figure 10 with zoosporangia( ?). X 125.

Myrionema corunnae f. sterile S. and G.

Fig. 13. A diagrammatic section to show tlie gradual diminution in the
length of the erect filaments from the center to the circumference of the frond.

Fig. 14. A section at the margin of the frond. X 250.

Fig. 15. A section through the center of the frond showing mature game-
tangia and interspersed sterile filaments. X 250.

Myrionema globosum f. afme S. and G.
Fig. 16. A fragment of a typical plant. X 125.

[350]

UNIV. CALIF. PUBL. BOT. VOL. 7

[SETCHELL-GARDNER ] PLATE 33

PLATE 34
Myroinema minutissimum S. and G.

Fig. 1. Two groups of gametangia, the left, younger, and the right, older.
X 125.

Fig. 2. A segment of the base showing the typical splitting of the terminal
cells. X 250.

Fig. 3. Small groups of gametangia showing different lengths of pedicels.
X 250.

Myrionema attenuatum f. doUiformum S. and G.

Fig. 4. A fragment showing relatively large erect sterile filaments and
relatively small and few gametangia. X 250.

Fig. 5. A fragment showing the ojiposite condition to that of figure 4.
X 250.

Myrionema attenuatum S. and G.

Fig. 6. A fragment taken near the margin of the frond. X 250.
Fig. 7. A fragment taken near the center of the. frond. X 250,

Myrionema foecundum f. divergens S. and G.

Fig. 8. Sections taken from four different places in the frond. X 125.
Fig. 9. A segment of the base at the margin of the frond. X 250.

Myrionema compsonemat aides S. and G.

Fig. 10. A section of a characteristic frond showing the erect sterile fila-
ments, gametangia, and "ascocysts. " X 125.

Myrionema Jiecatonematoides S. and G.

Fig. 11. Five fragments showing the nature of the base, the shapes, sizes,
and positions of the gametangia, and the erect filaments. X 125.

Myrionema primarium S. and G.

Fig. 12. A section through a typical frond and its host perpendicular to the
latter. X 250.

[352]

UNIV. CALIF. PUBL. BOT. VOL. 7

[SETCHELL-GARDNER ] PLATE 34

1922] Setchell-Gardner : Phycological Contributions 353

III. NEW SPECIES OF COMPSONEMA

Introduction

The genus Compsonema was established by Kuekuck. We have
accepted the genus to include forms which, like Myrionema, start
from a single cell that, by divisions, soon originates a series of closely
crowded, radiating filaments, forming a monostromatic basal layer
more or less circular in outline ; and which, unlike Myrionema, produce
on erect filaments numerous gametangia that develop pluriseriate
loculi. The genus is very closely related to the genus Myrionema on
one side and to Hecatonema on the other. No species of this genus
have previously been reported from the Pacific Coast of North America.

Compsonema streblonematoides sp. nov.

Plate 35, figure 4

Frondibus pulvinulos tenues, orbiculares, 3-5 mm. diam. for-
mantibus; parte prostrata filamentis irregularitates hospitis arete
adhaerantibus composita, inferne ramulos radiciformes inter cellulas
hospitis usque ad 300-350/* profunditate penetrantes, superne filamenta
sterilia gametangiaque emittente; filamentis erectis 140-160/x longis;
cellulis filamentorum erectorum cylindricis, 8.5-10;a diam.. 1.5-3-plo
longioribus; zoosporangiis ignotis; gametangiis plerumque cylindricis,
brevi-pedicellatis, 55-70/* longis, ll-ll/t latis; loculis plerumque
2-seriatis.

Growing on the pneumatocysts of Nereocystis Luetkeana. IMouth
of Tomales Bay, Marin County, California. Type, Gardner, no. 3442a
(Herb. Univ. Calif., no. 207035), August.

Compsonema strehlonematoides, like a Strehlonema, has a large
part of the vegetative portion of the frond within the host-plant, the
penetrating filaments extending into the host two to three times as far
as the length of the external vegetative filaments. The whole frond,
however, is the result of the growth of a single cell and forms a compact
monostromatic basal disk. In the character of the gametangia it is
close to a Myrionema, but in a very large majority of cases the loculi
are biseriate. i

354 University of California Publications in Botany [Vol. 7

Compsonema intricatum sp. nov.

Plate 35, figures 1-3

Frondilnis 1.5-2.5 mm. nltis, plus minusve confluentibus, stratum
cojitimium volntimim iiulcfiiiitc cxpaiisum foi-maiitibus; filanicntis
(•('pciitibns iinmcro.sis. i)i-ol'use ramosis, tlcetuantibus; lilameutis circtis
mimerosis, ant simi)li('ibus aut basim ramis paucis brevibus instructis,
rcctis. apice leviter atteniiatis, pro parte pilif eris ; pilis veris demitibus ;
ccllulis filaiiiciiloniin i-0])oiitiiiiii f(irma masrnitiidinof|no irrcfrnbiribiis;
ecllulis lilameiitorum ercctorum cylindrieis, 8-!)/a diain., 1-2-plo lon^:!-
oribus, ad basim filamentonim leviter angiistioribus; ehromatopboris
taeniatis; zoosporangiis ovoideis usque ad ellipsoideis, in filamcntis
brovil)u.s 0 filamoutis ropontilius orinndis torminalil)us, in fihmicntis
erectis prope basim jx'dicellatis scssilibusve, laleralibus, 25-33/x loiigis,
18-22fi latis; gamotangiis in filamentis erectis brevibus e filamentis
repentibus orieundis terminalibus, in filamentis erectis ad basim sessil-
ibus p(>diepllatisve lateralibns, aut in filamentis primariis ereetis intor-
calaribus, 80-120/* (usque ad 175/*) longis, 10-14/1 latis, iis terminalibus
interealaribusque usque ad 600/* longis; loeulis plerumque 2-seriatis.

Growing on Funis furcatus f. Carmel Bay, Monterey County,
California. Type, Gardner, no. 2893 (Herb. Univ. Calif., no. 207033),
May.

Compsonema intricatum bas diverged very far from a typical
Compsonema. The size of a single individual at maturity cannot be
ascertained, since the creeping filaments, and to some extent the erect
filaments, are thoroughly and inextricably intertwined, forming con-
tinuous velvety strata on the surface of the host for several inches in
expanse. The gametangia have, to a considerable degree, moved to
the lateral position, the majority, however, remain terminal on long
pedicels from the basal filaments. The lateral position is characteristic
of Ectocarpus. The intercalary gametangia are typical of the genus
Pylaiella. The creejoing filaments are very numerous, although they
do not radiate regularly side by side, thus forming a regular circular
disk typical of the genus Myrionem,a. The zoosporangia( ?) are
numerous and develop with the gametangia on the same individual.
It is exceedingly doubtful whether or not these structures are func-
tional. At the stage in which the material was found, they were
practically all of the same size, and some of the accompanying game-
tangia were empty. There is no indication of the formation of
zoospores. They are filled with what seems to be densely crowded,
angular ehromatophores. The apical end is slightly beaked, and lli(>
outer wall dissolved, except a thin inner membrane. Their position
on the .same plants with gametangia. the peculiar oj)cn beak, and the

1922] Setchell-Gardner : Pliycologkal Contributions 355

fact that they all seem to be of the same age lead us to suspect that
they are some sort of abnormalities, or pathological conditions of the
gametangia, which will require much more investigation to interpret.
We are retaining this in the genus Compsonema on account of the
prevailing simplicity of the erect filaments and the abundance of
creeping filaments, practically every cell of which gives rise to an erect
filament.

Compsonema fructuosum sp. nov.
Plate 36, figure 2

Frondibus pulvinulos orbiculares aut leviter irregulares, 4-5 mm.
diam. formantibus; parte prostrata filamentis tortuosis repentibus
irregularitates hospitis superficies arete adhaerantibus ; filamentis
erectis simplicibus, dense stipatis, 190-230ju, longis, cellulis cylindricis.
8-9/x diam., inferne 2-4-plo, superne 1-2-plo longioribus; chromato-
■phoris taeniatis; pilis veris deuntibus; zoosporangiis sparsis, late
clavatis, pedicellatis brevibus e filamentis repentibus oriundis suifultis,
55-65/x (usque ad lOO^a) longis. 22-28/* latis; gametangiis in filamentis
erectis terminalibus, cylindricis usque ad leviter fusiformibus, obtusis,
80-120/* longis, 12-16/* latis; loculis 2— 4-seriatis.

Growing on the pneumatocysts of Nereocystis Luetkeana. Tomales
Bay, Marin County, California. Type, Gardner, no. 3442 (Herb.
Univ. Calif., no. 207034), August.

Compsonema fructuosum, departs from the type of the genus C.
gracile Kuckuck (1899, p. 90, pi. VI (12), figs. 6-9) in having a
different host, in having no hairs, in having fewer and shorter sterile
filaments, in having more gametangia and in having differences in
details of measurements. In the abundance and position of the fruit
and in the definite circular thallus with compact monostromatic basal
layer of filaments, it approaches very closely to a typical Myrionenm,
but the multiseriate gametangia represent a stage in development
which we have not admitted into that genus. The gametangia occupy
a definite zone on the outer ends of the erect filaments. Practically
all of the erect filaments bear gametangia.

At times the basal layer seems to be distromatic, or even poly-
stromatic, but careful investigation shows that this appearance is
occasioned by the overlapping creeping filaments which follow the very
irregular surface of the host, due to the death of its surface layer of
cells, probably before the epiphyte took possession. The zoosporangia
are very sparse, but specimens have been seen which show undoubted
zoospores.

356 University of California Puhlications in Botany [Vol.7

Compsonema pusillum sp. nov.

Plate 37, figure 3

Frontlil)iis jnilvinnlos tenues dilute fuscos, linea exteriore irregu-
lares, 3-4 nun. <liam. fonnantibus; parte prostrata filamoutis tortuosis,
raniosis. irrc<rularitatos superficies hospitis arete adhaerante e latere
inlVro lilanicnta pauca. brevia, et inter eellulas hospitis penetrantia
emittente ; filamentis erectis simplicibus, cylindricis, 270-300/x longis ;
cellulis filanieutorum repentium maxime forma mafruitudineque irre^i-
larilms; cellulis filanieutorum erectorum cylindricis, 6-1 fx diam., 3-5-
plo lougioribu.s; cln-oniatophoris leviter interrupto-taeniatis ; zoo-
sporangiis sphaericis usque ad ovoideis, sessilibus aut pedicellis brevi-
bus e filamentis repeutibus oriundis sufiFultis, 18-25/x (usque ad 40/x)
lonjris. in-22/A latis; gametanj^riis cylindrico-couicis usque ad obtuse-
fusiformibus, sessilibus aut pedicellis brevibus e filamentis repentibus
oriundis sufiFultis, 40-60/^ longis, 10-12/x latis; loculis 1-2-seriatis.

Growing on tlie pneumatocysts of Nereocystis Luetkeana. Carmel
Bay, :\Iouterey County, California. Type, Gardner, no. 3582 (Herb.
T^niv. Calif., no. 207036), January.

Compsonema pusillum seems to overlap each of several genera. Its
zoosporangia, gametangia. and penetrating filaments ally it with
Strchlonema. Sauvageau (1897) has figured Myrioncma with rhizoids.
The gametangia are very small and have about as many uniseriate as
biseriate loculi. It might be considered a very minute creeping
Ectocarpus. We have placed it in the genus Compsonema on account
of the scarcity of fruit in ])roportion to the sterile tissue, and on
account of the monostromatic base, believing that the penetrating
filaments are more or less accidental on account of the injured con-
dition of the surface layer of cells of the host.

Compsonema speciosum f. piliferum f. nov.

Plate 38, figures 1, 2

Frondibus pulvinulos linea exteriore plus minusve orbiculares,
5-8 mm. diam. formantibus; filamentis repentibus maxime irregulariter
et profuse ramosis. irregularitates superficies hospitis arete alhaeran-
tibus. interdum ramis brevibus inter eellulas incolunies hospitis pene-
trantibus; filamentis erectis numerosis, simplicibus aut ramos breves
secundo.s, fructiferos aut interdum ramum piliferum emittentibus,
500-800;a longis, ba.sim attenuatis, superne piliferis; cellulis filanieu-
torum repentium forma magnitudiueque maxime irregularibus; cellulis
filamentorum erectorum cylindricis, basim 6-7.5jit diam., in parte
latissima ^-Vlfi diam., usque ad 15-plo longioribus; chromatophoris
taeniatis. in cellula quaque panels ; zoosporangiis obovoideis, ^o-I^o/a
longis, 28-32/A latis, raris; gametangiis filamentis repentibus oriundis

1922] Setchell-Gardner : Phycological Conti-thutions 357

sessilibus ant in pedicellis longioribus brevioribnsve, in filamentis
erectis secnndis, sessilibus pedicellatisve, aut rariore intercalaribus,
gametangiis terminalibus 60-100/a longis, 14-18/a latis, gametangiis
secnndis variabilibus, parvioribus.

Growing on the pneumatocysts of Nereocystis Luetkeana. ]\Ioclips,
Washington. Type, Gardner, no. 3812 (Herb. Univ. Calif., no.
207037), May.

Compsonema speciosum f. piliferum seems undoubtedly to be very
closely related to Myrionema speciosum Borgesen [1902, pp. 421-424,
which is the Hecatonema speciosum (Borgesen) Cotton, 1912, p. 15
(122) and the Hecatonema diffusum Kylin (cf. Cotton, loc. cit.)],
at least the two sets of plants agree very closely in general habit and
structure as described by Borgesen. Judging' alone from the descrip-
tion and figures the chief differences are that our plant has short
penetrating rhizoids, many terminal and no lateral hairs, and larger
gametangia. We are unable to say that the plant would produce
rhizoids on an uninjured host. The surface cells of the host are
destroyed under the epiphyte, and it is an open question whether the
penetrating rhizoids are or are not the cause of their death. Borgesen
does not give the dimensions of the whole plant, but he states that the
plants form short dense mats on the coneeptacles of Himanthalia lorea,
which would indicate that they were quite small. Our plants form
circular cushions or mats 5-8 mm. in diameter. Typical zoosporangia
were found at the ba»se of the erect j&laments. The lateral secund
gametangia standing nearly perpendicular to the filaments of which
the cell at the base in the filament becomes a part, is a character not
found in Compsonema gracilis Kuckuck.

Compsonema sporangiiferum sp. nov.

Plate 36, figures 3-8

Frondibus pulvinulos orbiculares, 1-4 mm. diam. formantibus ;
parte prostrata filamentis compactis, irregularibus, plus minusve con-
tortis, irregularitates superficies hospitis arete adhaerantibus, mono-
stromatica aut pro parte distromatica composita; filamentis erectis
300-700/x longis, simplicibus aut superne ramis panels, brevibus,
plerumque secnndis, fructiferis indutis; cellulis filamentorum erec-
torum basim 6-8/t diam., 2-3-plo longioribus, apice ll-13ju, diam.,
1-1.5-plo longioribus, cellulis multis in parte superiore filamentorum
1-2-plo longitudinaliter divisis; chromatophoris taeniatis. in eellula
quaque paueis; zoosporangiis numerosissimis, forma magnitudineque
maxime variabilibus, anguste ellipsoideis, clavatis, usque ad late
ovoideis, 40-130/a longis, 20-35/* latis, sessilibus aut pedicellis brevibus

358 UniversHy of California Puhlications in Botany [Vol. 7

0 filninciitis reii('i)til)us oriniulis siifTiiHls. ant in filMmeiitis erectis
lalcralihus tcniiiiialihiisvc ; f^ainctaiiffiis raris, cylindricis, in filamentis
reiiciitihus scssilihus hn'vi-ix'diccllatisve ant raro in filamentis erectis
Irniiin.ilihus Inhereulatisqne, 8()-]30/a longis, 11-15/i, latis.

(Jrowinj,'- on tiie i)neuiiiatuey.st of Nereocystis Luetkeana. Neah
Bay, Washington. Type, Gardner, no. 3859 (Herb. Univ. Calif., no.
207038). :\ray.

Tlie unusual condition in Compsonema prevails in C. sporangii-
fcnun. The gametangia seem to be of rare occurrence. Examination
of specimens of different sizes from different parts of the host revealed
the presence of a large number of zoosporangia in all stages of develop-
ment, man>' I)eing empty. The gametangia are prevailingly near tlie
base, although occasionally a complex one appears terminal on an erect
lilaiiii'ut. A very common character is the division of the cells in the
u|)|)ci' i)arts of the filaments into 2-4 parts by longitudinal walls, pro-
ducing a slight clavate condition in such filaments. AVe do not know,
at present, tlie morphological significance of this condition. The
zoosporangia and the gametangia, so far as observed, develop on distinct
individuals.

Compsonema sessile sp. nov.

Plate 39, figure 6

Frondibus pulvinulos inconspicuos usque ad 1.5 mm. diam., in linea
exteriore orbiculares usque ad irregulares formantibus; filamentis
repentibus contortis, dense compactis, hospitem arete adhaerantibus,
ramulis radiciformibus deuntibus; filamentis erectis sparsis 20-2-!:,u
altis; pilis veris deuntibus: collulis filamentorum repentium foimia
magnitudineque irregularibus, 9-12/x diam.; chromatophoris taeniatis
in cellula quaque singulis; zoosporangiis ignotis; gametangiis numer-
osis, in filamentis repentibus sessilibus, conicis usque ad fusiformibus,
16-22/i longis. ll-14ju, latis, parietibus multorum loculorum obliquis.

Growing on the lamina of HedophyUum sessile near the outer end.
Neah liay, Washington. Type, Gardner, no. 3866 (Herb. Univ. Calif.,
no. 207039), May.

Compsonema sessile represents the extreme of the genus in the
direction of simplicity. There are no hair filaments, and it is doubtful
if any of the erect filaments remain sterile. At times even the cells of
the basal layer seem to become a part of the gametangia, at least more
than one cell in the basal layer may be at the base of a single game-
tangia. There would seem to be only a single modification of a
Myrianema of the simplest type to produce this species, viz., the
change in the type of gametangium from uniseriate to ])luriseriate. If

1922] Setchell-Gardncr : Fhycological Contributions 359

we hold to the principal distinction between Compsonema and Myrio-
nema, it will be necessary to ally this species with the former. It was
found growing in company with Streblonema aecidioides f. pacificiim
S. and G., which gave to the host a roughened appearance, otherwise
it probably would not be noticeable without microscopic examination.

Compsonema tenue sp. nov.

Plate 37, figure 6

Frondibus microscopicis, stratum plus minusve continuum in
hospite inter algas parvas alteras formantibus; filamentis repentibus
numerosis, ramosis, irregularitates superfices hospitis arete adhaeran-
tibus; filamentis erectis sparsis, simplicibus, apice basimque leviter
attenuatis, SO-lOO^u, altis; pilis veris sparsis; cellulis filamentorum
repentium forma magnitudineque irregularibus ; cellulis filamentorum
erectorum cylindricis, non constrictis, 7-8/x diam., 1.5-2.5-plo longior-
ibus; cellulis filamentorum piliferorum cylindricis, 5.5-6.5^ diam.,
inferne exacte aut fere quadratis, superne 7-10-plo longioribus; zoo-
sporangiis( ?) clavatis, 24-28ju, longis, 8-1 0/t latis. sparsis; gametangiis
fusiformibus usque ad irregulariter conicis, in filamentis repentibus
sessilibus, numerosis, 22-28ja longis, 7-11/x latis; loculis plerumque
2-seriatis.

Growing on the fruiting fronds of Cystopliyllum geminatum.
Sitka, Alaska. Type, Gardner, no. 3972 (Herb. Univ. Calif., no.
207040), July.

Compsonema teyiue, though scarcely typical of the genus, seems to
possess affinities as closely allied to Compsonema as to any other
genus, and hence we are placing it here for the present.

Compsonema nummuloides sp. nov.

Plate 35, figures 5, 6

Frondibus pulvinulos tenues orbiculares, 7-12 mm. diam. forman-
tibus; parte prostrata filamentis radiantibus, multo crispatis, dense
compactis, irregularitates superficies hospitis arete adhaerantibus
composita; filamentis erectis simplicibus, numerosis, 300-400ju, longis,
apice basimque leviter attenuatis; pilis veris deuntibus; cellulis fila-
mentorum repentium forma magnitudineque multo irregularibus;
cellulis filamentorum erectorum cylindricis, basim 6-7ju. diam., 3-8-plo
longioribus, in parte latissima 9-1 I/a diam., cellula apicali quadrata
6-7/x diam., 3-8-plo longiore ; zoosporangiis ignotis; gametangiis
plerumque sessilibus aut brevi-pedicellatis, nunc in filamentis erectis
terminalibus, 70-90/a (usque ad 140/a) longis, ll-15ju, latis; loculis
plerumque 2-seriatis.

Growing on the pneumatocysts of Nereocystis Luetkeana. IVIoss
Beach, San Mateo County, California. Type, Gardner, no. 4318 (Herb.
Univ. Calif., no. 207041), July.

360 UnivcrsiUj of California Publications in Botany [Vol. 7

Compsonema nummuloidcs forms tliin, liy:lit brown cushions on the
pnenniatoeysts of the host. The whole cushion is a single plant,
cin-iilar in ontlino and attaining a diameter of twelve or more milli-
nu'tcrs, although the majority of the plants are much smaller. Like
a typical Mjjrionnno the plant starts from a single cell and by divisions
forms a series of filaments radiating in all directions with apical
growth and dichotomous branching, by the splitting of the apical cell,
thus forming a continuous monostromatic layer of cells. Beginning
in the center, practically every cell in each radiating filament of the
basal layer gives rise successively to an erect filament, about two- thirds
of which are fructiferous, the others remaining sterile. A very large
majority of the gametangia are sessile or on short pedicels, thus
occupying a zone near the creeping filaments, a Myrionematoid char-
acter, but the remainder of the filaments continue to grow and attain
a length of 300-400ju,. Some of these filaments are terminated by
relatively short gametangia. This is one of several species with this
general method of development which has been found growing only
on the above mentioned host, differing from each other and from the
type of the genus, T. gracile Kuckuck, in the size of the plant as a
whole, in details of dimensions of their parts, in the presence or absence
of zoosporangia and hairs, in the position of the gametangia, and
in the relative amount of sterile filaments. It seems to form a
fairly compact group ])ut with overlappings, however, in the genera
Strehlonema, Myrionema, and Hecatonema.

Compsonema fasciculatum sp. nov.

Plate 38, figures 7-9

Frondibus flocculos micro.scopicos plus minusve confluentes for-
mantibus; filamentis repentibus tortuosis. hospitem sine ramulis
radiciformibus adhaerantibus; filamentis erectis prope basim ramosis.
90-130/x longis, basim apiceque attenuatis, non piliferis, ad dissepi-
mentaconstrictis; pilis verisdeuntibus; cellulis filamcntornm ei-ectorum
in parte latissima, 10-1 4/x, diam.. 1-2-plo longioribus; zoosporangiis
ignotis; gametangiis forma moderate irregnlaribus. conicis usque ad
fusiformibus, 35^5/x longis, 12-18/a latis; pleruiiKinc lateralibus. brevi-
pedicellatis.

Growing on Gigartina radida f. Pacific Grove, California. Type,
Gardner, no. 4503 (Herb. Univ. Calif., no. 207042), December.

This species is allied here rather than with Edocarpus because of
its extremely small size and its creeping, fasciculate habit.

1922] SetcJiell-Gardner : Phycological Contributions 361

Compsonema myrionematoides sp. nov.

Plate 36, figure 1

Frondibus pulvinulos parvos, orbiculares usque ad irregulares,
1-2.5 mm. diam. f ormantibus ; parte prostrata arete monostromatica,
filamentis dense compactis, tortuosis, radiantibis composita ; filamentis
erectis simplieibus. eylindricis, 110-130/x longis; pilis veris deuntibus;
eellulis filameiitorum repentium forma eylindricis usque ad irregular-
ibus; eellulis erectorum eylindricis, 7.5-8.5/* diam., 1-2-plo longioribus,
terminalibis plerumque plurimo longioribus ; chromatophoris leviter
interrupto-taeniatis, in quaque cellula singulis; zoosporangiis( ?) late
clavatis sessilibus aut brevi-pedicellatis e eellulis filamentorum repen-
tium oriundis, 35-50/a longis, 20-28/* latis ; gametangiis numerosis,
eylindricis usque ad fnsiformibus sessilibus aut brevi-pedicellatis e
eellulis filamentorum repentium orundis, 50-70/t (usque ad 100/t)
longis, 9-12/t latis.

Growing on the stipe of Nereocystis Luetkeana. Pacific Grove,
California. Type, Gardner, no. 4510 (Herb. Univ. Calif., no. 207043),
December.

There is but little choice as to the generic position of this small
alga. It forms small brown tufts or cushions of loose filaments. Very
frequently the gametangia are wider in the middle than at the ends and
have decidedly biseriate loculi. Because of the more or less loose tufted
character of the erect filaments and the partially biseriate gametangia
we are placing it with the genus Compsonema.

Compsonema secundum sp. nov.

Plate 37, figures 1, 2

Frondibus parvissinus, plerumque plus minusve eonfluentibus ;
parte prostrata monostromatica, filamentis crispatissimis, ramosis com-
posita; eellulis filamentorum repentium forma irregularibus, 7-8,/i
diam., longitudine variabilibus; filamentis erectis usque ad 1.25 mm.
altis, simplieibus aut raro ramulis paucis brevibus fructiferis in-
structis; eellulis filamentorum erectorum eylindricis, basim 5. 5-6. 5/*
diam., 4.5-6-plo. longioribus. in parte latissima 9.5-10.5/x diam., eellulis
terminalibus usque ad 9-plo longioribus quam latis; chromatophoris
taeniatis; zoosporangiis late clavatis, 60-90/t longis, 22-28/t latis.
sessilibus aut brevi-pedicellatis e eellulis filamentorum repentium
oriundis; gametangiis maxime variabilibus, raro pedicellis brevibus e
filamentis repentibus oriundis sufi'ultis, interdum per transformatione
partis superioris filament! erecti longi extensis, nunc .simplieibus, nunc
prolifieationes numerosas laterales, breves plerumque secundas, raro
oppositas producentibus, usque ad 400/*. longis, ll-15/i latis, obtusis,
clavatis.

362 University of California Publications in Botany [^^«i- "

Growinj; on tlic i)n('uinatocyst of Nereocystis Luetkeana. West
coast of Wasliin}?toii (Moclips) to central California (Carmel Bay).
Type. Gardner, no. 4547 (Herb. Univ. Calif., no. 207044), Carmel Bay,
California, December.

Like Compsonema ramulosum, C. secundum has gametangia occupy-
ing a variety of positions and assuming a diversity of forms. The
form of gamentangiiim which marks the species is the long terminal
form which develops numerous, short, blunt-conical, seriate, secund,
sessile gametangia, thus forming a complex composed of these game-
tangia and the metamorphosed cells in the upper part of tlic filament.
The end of the filament becomes reflexed. at times almost scorpioid.
The specimen from ]\foclips, Washington, was taken in IMay and tlie
gametangia were practicality all empty. The other parts of tlio plants
manifested signs of old age conditions. They seem to belong to this
species, but the filaments have a considerably greater diameter. Typical
zoosporangia with well formed zoospores have been observed in tliis
collection.

Compsonema ramulosum s]i. nov.

Plate 39, figures 1-5

Frondibus pulvinulus orbicnlares, 5-7 mm. diam. formantilms;
parte prostrata monostromatiea, filamentis plus minusve contortis et
irregulariter ramosis, radiantibus composita ; filamentis erectis pro
parte piliferis simplicibus aut ramos breves plerumque fructiferos,
secundos et interdnin ])ilos ferentibus, basim leviter attenuatis; ccllnlis
filamentorum rejK'nliuni 6-7/a diam., (juadratis aut leviter longioribus
quam latis; cellulis filamentorum erectorum basim 6-7. 5/a diam.,
superne leviter latioribus, 1-3.5-plo longioribus ; cellulis ramorum
pilornmque 4-6/x diam.; chromatophoris taeniatis. plus minusve inter-
rui)tis angnhiribusque; zoosporangiis( ?) late clavatis, terminalibus
pedicellis longioribus bervioribusve suffultis aut prope apices fila-
mentorum erectorum lateralibus, secundis aut verticillatis; gametangiis
forma, magnitudine et positione maxime variabilibus, in filamentis
repentibus sessilibus ])edicellatisve, in filamentis erectis lateralibus. aut
in filamentis primariis aut in ramulis lateralibus brevibus intercalar-
ibus terminalibusve. clavatis, cylindricis, usque ad cylindrico-conicis,
usque ad 14()/i. longis, 16/^ latis; loculis pl(>runuiue 2-seriatis.

Growing on the pneumatocysts of Nereocystis Luetkeana. Carmel
Bay, Monterey County, California. Type, Gardner, no. 4549 (Herb.
Univ. Calif., no. 207045), December.

There exists in this species extreme variation in tlu> form, size, and
position of the gametangia. Some are sessile on the creeping filaments,
though they are more frequently to be found on the pedicels up to

1922] Setchell-Gardner : Phijcological Contributions 363

fifteen cells long. At times sessile forms are to be found on the erect
filaments about two-thirds of the way up to the apex, but more gener-
ally they are terminal on small lateral ramuli on the erect filaments
arising from the middle to near the apices. Rarely there are small
lateral forms on the ramuli. Finally, they may rarely be terminal or
intercalary and seriate on the erect filaments although the erect
filaments are usually piliferous. The terminal forms on pedicels, and
particularly on the main erect filaments, are often quite blunt, even
clavate. Others on the prostrate filaments are narrow and sharply
attenuated, at times terminated by a sterile pointed filament, in effect
intercalary. Many of the gametangia, terminal on the ramuli, are
composed of a single series of loculi. Most of the others are two or
more seriate. The hair filaments do not seem to be abundant. They
arise laterally on the main filaments, usually above the center. The
meristem in these hairs is at the base which is surrounded by a sheath
similar to that described by Sauvageau (1897, p. 47) for Myrionema.
The ramuli are decidedly curved upwards.

The species is similar to Myrionema speciosum from the Faeroes,
described by Borgesen (1902, p. 421). Our plant is more slender in
all of its parts, has much longer gametangia, and possesses, in addition
to the sessile, secund gametangia figured by Borgesen, occasional inter-
calary ones and many terminal ones on the numerous clustered ramuli.
These ramuli are in turn occasionally branched, usually arise in
groups, sometimes whorled, two or three arising from the same cell, in
other instances they are secund.

TliQ species often grows in association with other members of the
Myrionemataceae and the Ectocarpaceae. As a rule, the main erect
filaments do not enter into the formation of gametangia, but give rise
either to sessile gametangia or to the ramuli. No zoospores have been
observed in the so-called zoosporangia. We suspect that these may
represent pathological conditions.

Compsonema serpens sp. nov.

Plate 39, figure 7

Frondibus stratum plus minusve expansum in superficie hospitis
formantibus; parte repente filamentis multo contortis, irregulariter
ramosis composita ; filamentis erectis simplicibus aut raro superne
ramis paucis subulatis indutis, interne leviter attenuatis, superne
gradatim attenuatis, non piliferis, 375-425/i, longis; pilis veris deunt-
ibus; cellulis filamentorum repentium cylindricis ad dissepimenta
usque ad irregularibus, 10-13/* diam., 0.5-3-plo pongioribus; cellulis

364 University of California rnhlications in Botany [Vol. 7

filamentorum eroetorum evlindricis non constrictis. basi 5.5-8. 5/x diam.,
in part(^ latioro. lO-17/i diam., 1.2r)-2.5-plo loiifriorilm.s; chromatophoris
tat'iiiatis, in t-t'Ilulis juvt-nis dense aggrcgatis, in celhili.s scniorihus
numerosi.ssimis et sogregatis ; zoosporangiis ignotis ; gametangiis cylin-
drieo-eonit'is, in pediccllis longiorilms brevioribnsve e filamentis repent-
ibus nrinndis tcrniinalibns. rard in filamentis longis crcctis terminal-
ibns. ()()-! ;i(V longi.s, 18-28^ latis.

Growing on the sterile base of Gigartina radula f., in company with
several other small algae. Cypress Point, Monterey County, Cali-
fornia. Type, Gardner, no. 4684 (Herb. Univ. Calif., no. 207046),
December.

This diminutive member of the Melanophyceae is on the border line
between Compsonema and Ectocarpus. Our comprehension of these
two genera, so far as the vegetative portion is concerned, is that a
typical Compsonema, starting from a single cell, develops an extensive,
creeping, attaching mass of filaments, radiating in all directions from
the origin, which later produces very numerous erect filaments, some
of which may remain sterile and others become fructiferous ; and that
a typical Ectocarpus, starting likewise from a single cell, develops a
rather insignificant series of attaching filaments, more or less rhizoidal
in nature, and a relatively extensive system of erect filaments finally
producing the fruit. Starting with this conception concerning the
two genera, we are assuming that differentiation has proceeded from
Compsonema in the direction of the reduction of the creeping portion,
to a more extensive development of the erect portion, finally resulting
in an Ectocarpus.

The species under consideration has a relatively extensive system
of creeping filaments but very much less so than that of a typical
Compsonema. The creeping filaments do not form a solid disk, but
those from different plants are so intertwined that it is impossible to
determine the limits of a distinct individual. In tliis character it
resembles an Ectocarpus. The gametangia are like those of a. typical
Ectocarpus, but since they are mostly .short pedicellate and spring
directly from the creeping filaments, as is the case in a typical
Compsonema, and because of the relatively extensive system of creep-
ing filaments, we have placed it in the latter genus.

1922] Setchell-Gardner: PJiycological Contnhutions 365

Compsonema coniferum sp. nov.

Plate 38, figure 3

Frondibus microscopicis, plus minusve confluentibus, stratum
velutinum in superficie hospitis algis alteris minutis immixtis for-
mantibus; parte prastrata filamentis profuse ramosis. multo contortis
composita; filamentis erectis numerosis. simplicibus, apice basimque
leviter attenuatis, non piliferis, 150-200jU, altis; pilis veris deuntibus;
cellulis filamentorum repentium forma magnitudineque irregularibus,
6-7. 5/A latis, 9-12/i. longis ; cellulis filamentorum erectorum cylindricis
usque ad leviter doliiformibus in parte latiore, 8-10^ diam., 1-1.5-plo
longioribus ; chromatophoris taeniatis ; zoosporangiis ignotis ; game-
tangiis sessilibus aut in pedicellis brevibus e filamentis repentibus
oriundis, angusta conicis, 46-65/* longis, 15-18/a latis.

Growing on the sterile base of Gigartina radula f . Cypress Point,
Monterey County, California. Type, Gardner, no. 4684a (Herb.
Univ. Calif., no. 207047), December.

Compsonema coniferum seems to be a very close relative of C
fasciculatum, found growing on the same host. The fronds are more
extensive, unbranched, and have larger gametangia. We consider this
species of Coi7ipsonema a very near approach to an Ectocarpus of
extremely small size. We are inclined toward the genus Compsonema
as a more appropriate genus for this species than Ectocarpus on
account of its extensive creeping habit and prolific production from
the creeping filaments of nearly sessile gametangia and many sterile
erect filaments. It is to be found quite commonly associated on the
same host with several other species of ]\Iyrionemataceae.

Compsonema dubium sp. nov.
Plate 38, figure 6

Frondibus diminutivibus, flocculos parvos inter algas micro-
scopieas alteras in hospite formantibus; parte prostrata filamentis
multo contortis, ramosis, repentibus composita ; filamentis erectis
simplicibus. totaliter prope exacte cylindricis, 275-350/t longis. non
piliferis ; pilis veris deuntibus ; cellulis filamentorum repentium forma
magnitudineque maxime variabilibus ; cellulis filamentorum erectorum
cylindricis, medio 7-9/t diam., superne inferneque minoribus, 2-3-plo
longioribus; zoosporangiis( ?) ellipsoideis, terminalibus, brevi-pedi-
cellatis 16-22/it longis, 12-15/x latis ; gametangiis cylindricis usque ad
leviter fusiformibus, in pedicellis longioribiLS brevioribusve filamentis
repentibus oriundis, terminalibus, raro in filamentis longis erectis
terminalibus, 50-75/a longis, 8-11/x latis; loculis plerumque 2-seriatis.

366 University of California Puhlications in Botany [Vol.7

Growing on the sterile base of Gigartina radula f. Cypress Point,
Monterey County, California. Type, Gardner, no. 4684d (Herb.
Univ. Calif., no. 207048), December.

CompsuHona diihium, like C. coniferum and C. serpens, has close
affinities with small members of the genus Ectocarpus. Its relatively
long and narrow gametangia distinguish it from the two species here
mentioned, with which it is frequently associated.

Compsonema secundum f. terminale f. nov.
Plate 37, figures 4, 5

Frondibus stratum indefinite expansum, velutinum, per filamenta
niaxime distorta. ramosa, repentia, per ramulos radiciformes. numcr-
osis, penetrantes. e superficie infera oriondes affixum formantibus;
filamentis erectis 700-800)u, (usque ad 1 mm.) longis, simplicibus aut
ramulos paucos. breves, secundos, fruetiferos produccntibus. rectis,
basim leviter attcnuatis, superne cylindrieis; cclhili.s filamcntorum
repentinm et ramuloi-um radiciformium irregular-ibus, maunitudine
variabilibus; cellulis filamentorum erectorum cylindrieis, ad dissepi-
menta non constrictis a 6-8/x diam., liasi 2-3.5-plo, superne 0.5-2-plo,
longioribus; chromatophoris irregulariter taeniatis, in cellula quaque
paucis; zoosporangiis( ?) late clavatis, c filamentis repentibus oriundis,
50-60/i longis. 18-22^ latis, sessilibus, brevi-pedicellatisve, aut ter-
minalibus seriatisque. proliferationibus lateralibus secundisque indutis ;
gamctangiis e filamentis repentibu.s oriundis, sessilibus aut longe-
])edicellatis, 90-1.1 0/x longis, 15-1 8/.i latis, proxime cylindrieis aut in
filamentis erectis terminalibus, clavatis, frequenter prolongationibus
tubereulatis, secundis et magnitudine variabilibus.

Growing on the pneumatocysts of Nereocystis Luctkeana. Central
California (San Francisco and Pacific Grove). Type, Gardner, no.
4690 (Herb. Univ. Calif., no. 207049), Pacific Grove, December.

This form of Compsonema is to be distinguished by the great pre-
dominance of relatively short terminal gametangia, and manj^ rela-
tively large, lateral, seeund, sessile gametangia which are curved up-
wards. There are a few sessile or short pedicellate examples at the
base and a few lateral forms on short ramuli. Specimens taken in
December had the larger number of gametangia empty.

1922] Setchell-Gardner : Phycological Contributions 3G7

LITERATURE CITED

BORGESEN, r.

1902. The marine algae of the Faeroes, in Warming, Botany of the Faeroes,
part 2, p. 339. Copenhagen.

Cotton, A. D.

1912. Marine algae, in Clare Island Survey, part 15. Proc. Eoyal Irish
Acad., vol. 31.

KUCKUCK, P.

1899. Beitrage zur Kenntniss der Meeresalgen, in Wissenschaftliche Meeres-
untersuehungen herausgegeben von der Kommission zur deutsehen
Meere in Kiel und der biologischen Anstalt auf Helgoland. Neue
folge. Dritter Band. Abteiiung Helgoland. Heft 1. Kiel und
Leipzig.
Kylin, H.

1907. Studien iiber die Algenflora der schwedisehen Westkiiste. Akadem-
isch. Abhandlung. Upsala.

Sauvageau, C.

1897. Sur quelques Myrionemacees. Ann. des. Sci. Nat., 8 ser., Bot., pp. 161-

288.

PLATE 35

Compsonema intricattim S. and G.

Fig. 1. A diagrammatic illustration of a part of a plant.

Fig. 2. A part of a plant showing a variety of forms of gametangia, lateral
and pedicellate or sessile, terminal, and intercalary; also numerous zoospo-
rangia(?) on the same plant and also, in some cases, on the same filament. X 125.

Fig. 3. A small part of a plant showing the acute erect filaments, and
pedicellate gametangia and zoosporangia. X 225.

Compsonema streMonematoides S. and G.

Fig. 4. A part of a section through a plant and its underlying host, showing
a ba.sal filament on the surface and rhizoidal filaments penetrating into the host,
and erect filaments and gametangia above. X 250.

Compsonema nummuloides S. and G.

Fig. 5. A segment of the sterile creeping filaments showing the method of
branching. X 125.

Fig. 6. Eight fragments of plants showing the various shapes and positions
of the gametangia and the character of the erect filaments. X 125

[368]

UNIV. CALIF. PUBL. BOT. VOL. 7

[SETCHELL-GARDNER ] PLATE 35

PLATE 36

Compsonevia myrionematoides S. and G.

Fig. 1. Six fragments of plants showing the character and position of the
gametangia, the basal filaments and the erect sterile filaments. X 125.

Compsonema fructuosum S. and G.

Fig. 2. A small part of a frond showing some abnormal developments of
gametangia. Two are terminal on lateral branches. Two have developed in
one old gametangium. One is terminal on a long pedicel growing up through
an empty gametangium. X 2.50.

Compsonema sporangiifenim S. and G.

Fig. .3. A few fragments of erect filaments showing unusual terminal game-
tangia and terminal zoosporangia. X 125.

Fig. 4. A group of young gametangia and erect filaments. X 12.5.

Fig. 5. A fragment of a plant showing zoosporangia in various positions,
some sessile on the basal filaments, some sessile and lateral and some terminal
on the erect filaments. X 125.

Figs. 6, 7. Fragments of filaments showing lateral secund branching near
the apices. X 125.

Fig. 8. Fragments showing positions of well developed zoosporangia. X 125.

[370 J

UNIV. CALIF. PUBL. BOT. VOL. 7

[SETCHELL-GARDNER ] PLATE 36

PLATE 37

Compsonema semmdum S. and G.

Fig. 1. Three fragments of plants showing small gametangia on short
pedicels from a basal filament and complex terminal gametangia with mostly
secund lateral developments. X 125.

Fig, 2. A fragment of a plant showing a modification in which the game-
tangia are mostly terminal and erect, and only rarely possessing lateral pro-
tuberances. X 125.

Compsonema pusillum S. and G.

Fig. 3. Three fragments of plants showing shapes and positions of zoospo-
rangia and gametangia. Character of the creeping and of the erect filaments
and a few branches pushing between the surface cells of the host. X 125.

Compsonema secundum f. terminate S. and G.

Fig. 4. A fragment of a plant showing the characteristic terminal and large
lateral gametangia. X 125.

Fig. 5. A filament which seems to be producing zoosporangia below an
empty gametangium. X 125.

Compsonema tenue S. and G.

Fig. 6. Fragments of plants showing erect filaments, true hairs and zoospo-
rangia ( ?) and gametangia sessile on the creeping filaments. X 250.

[372]

UNIV. CALIF. PUBL. BOT. VOL. 7

[SETCHELL-GARDNER ] PLATE 37

PLATE 38
Compsonema speciosum f. piliferum S. and G.

Fig. 1. A fragment of a plant showing well developed rhizoids from the
creeping filament, various shapes and positions of the gametangia and terminal
hairs. X 125.

Fig. 2. A fragment showing a well developed zoosporangium with included
zoospores. X 125.

Compsonema coniferum S. and G.

Fig. 3. Fragments of a frond, showing the characteristic shapes, sizes, and
positions of the gametangia, and the character of the erect filaments. X 125.

Compsonema fructuosum S. and G.

Fig. 4. A fragment of a frond showing the character of the erect filaments
and a well developed zoosporangium. X 125.

Fig. 5. A group of characteristic gametangia. X 125.

Compsonema dubium S. and G.
Fig. 6. Fragments of typical plants bearing gametangia. X 250.

Compsonema fasciculaium S. and G.

Figs. 7-9. Fragments of plants showing the positions, shapes, and sizes of
the gametangia, the branching erect filaments and the tortuous character of the
creeping filaments. X 250.

[374]

UNIV. CALIF. PUBL, BOT. VOL. 7

[SETCHELL-GARDNER ] PLATE 33

PLATE 39

Compsonema ramulosum S. and G.

Figs. 1, 3, 4, 5. Fragments of plants showing the character of the creeping
filaments, the abundance of erect filaments, the method of their branching, and
the great variety in shapes, sizes, and position of gametangia. X 125.

Fig. 2. A fragment showing zoosporangia(?). X 125.

Compsonema sessile S. and G.
Fig. 6. A fragment of a typical plant sitting upon its host. X 250.

Compsonema serpens S. and G.
Fig. 7. Two fragments of typical plants. X 125.

[376]

UNIV. CALIF. PUBL. BOT. VOL. 7 [SETCHELL-GARDNER ] PLATE 39

1922] Setchell-Gardner : Phycological Contributions 377

IV. NEW SPECIES OF HECATONEMA

Introduction

The genus Hecatonema was established by Sauvagean (1897,
p. 249). We have in the Herbarium of the University of California
numerous collections of small epiphytic plants found upon Nereocystis
Luetkea^ia in various localities on the Pacific Coast, which we have
placed in the genus Hecatonema. This genus, as we interpret it, is
quite similar to Compsonema and to Myrionema, but differs from each
in having a distromatic base. Of the two, it seems more nearly akin
to Compsonema than to Myrionema in that it develops gametangia
with pluriseriate loeuli instead of uniseriate, as is the ease in Myrio-
nema.

There have been no species of this genus reported previously on the
Pacific Coast of North America. This study is but a preliminary one
and it is hoped that it may stimulate further investigation of this genus
as well as of the other closely related genera.

Hecatonema variabile sp. nov.

Plate 41, figures 1-12

Frondibus pulvinulos orbiculares, 4-7 mm. diam. formantibus ;
parte prostrata filamentis radiantibus subapicaliter ramosis, per
ramulos radiciformes numeros, breves, 1-3-cellulates adhaerantibus
composita ; filamentis erectis non dense stipatis. non e centro ad peri-
pheriam seriater evolvatis, simplicibus. 400-500/* longis; pilis veris
ignotis; cellulis filamentorum repentium forma irregularibus. 7-9/4
diam., margine 1-3-plo longioribus ; cellulis filamentorum ereetorum
basim 4.5-5/a diam., 2-5-plo longioribus, apice S-lOju, diam., 1-3-plo
longioribus; zoosporangiis( ?) late clavatis in filamentLs prostratis
sessilibus brevi-pedicellatisve raro in filamentis erectis terminalibus
lateralibusve, 50-65/i, longis, 20-24/i latis; gamentangiis cylindricis,
apice leviter attenuatis. in filamentis sessilibus brevi-pedicellatisve,
raro in filamentis erectis terminalibus, 70-120/x longis, 9-12/* latis;
loeuli 1-2-seriatis.

Growing on the pneumatocysts of Nereocystis Luetkcana. Carmel
Bay, Monterey County, California. Type, Gardner, no. 3109 (Herb.
Univ. Calif., no. 207050), December.

From the standpoint of the structure of the gametangia, the type
of this species of Hecatonema could scarcely be separated from a

378 University of California Publications in Botany [Vol. 7

Myrioncma such as M. phyllophyhim S. and G. They, in part, possess
unisori;itc loculi. Imt have many perpendicular and slanting walls,
iii;il<ing a partial biseriate condition. The basal layer, particularly in
the center of the thallus, is distinctly distromatic, developing numerous
peg-like rhizoids, which serve to anchor it more firmly to the host, and,
for the most part at least, the branching of the filaments is subterminal,
whereas that of a true Myrionema is terminal and brought about by
the splitting of the apical cell.

The size of the plants as a whole varies considerably. The material
of the type varies in this respect from 4 mm. to 7 mm. in diameter.
In some collections of specimens which we have included under the
species, the specimens are as small as 2 mm. in diameter, in others they
are up to 10 mm. in diameter. The length of the erect filaments
varies from 150/x. to 500/*. JMature gametangia vary from 55/* to 120/a
long. The relative number of gametangia terminal on the erect fila-
ments, as well as their size and complexity, is exceedingly variable in
different collections. Two or three cells only may metamorphose in
some filaments, in others eight or ten. Occasionally small lateral out-
growths from these metamorphosed cells may occur. The number of
erect filaments in proportion to the number of gametangia is a char-
acter which seems to be very unstable. In some collections they exceed
the number of gametangia almost two to one, in others they do not
occur more than in proportion of one to five. Tliej' are very irregular
in origin along the radiating creeping filaments. Some near the
margin of the frond may be nearing maturity before others near the
frond have started (pi. 41, fig. 5).

Hecatonema clavatum sp. nov.

Plate 40, figures 1-4

Frondibus pulvinulos orbiculares, 2-3 mm. diam., margine lato
filamentis ereetis destituta formantibus; parte prostrata per rnmnlos
radii-iformes nunierosos. breves, subulatosque adhaerente. et filamentis
regulariter radiantibus, dense compactis composita ; filamentis ereetis
plerumque clavatis simplieibus, 190-210;u, longis; cellulis filamentorum
repentium 5-6^ diam.. 1.5-2-plo longioribus; cellulis filameiiloi'um
erectorum inferne eylindrici.s. 5.5-6.5/* diam., 2-2.5-plo longioribus.
superne ad discepimenta leviter constrietis, 8-11/* diam., 1.5-plo
longioribus; zoosporangiis( ?) in filamentis repentibus. sessilibus sub-
sessilibnsve, clavatis, 60-80/* longis, 10-14/* latis; gametangiis cylin-
dricis us(|ue ad leviter fusiformibus. in filamentis repentibus .sessilibus
brevi-pedicellatisve, 55-85/* (usque ad 110/*) longis, 8-12/* latis; loculis
plerumque 2-seriatis.

1922] Setchell^Gardner : Phycological Contributions 379

Growing on the pneiimatocysts of Nereocystis Luetkeana. IMoutli
of Tomales Bay, Marin County, California. Type, Gardner, no. 3456
(Herb. Univ. Calif., no. 207051), May.

The structures designated zoosporangia( ?) in the above diagnosis
of this species were found interspersed among the gametangia. No
indication of the formation of zoospores has been observed, which leads
to the suspicion that they may be abortive gametangia. The terminal
cells of the erect filaments are often longer than the cells below it, and
in these filaments in which the cells are divided lengthwise, producing
the clavate condition, the terminal cell degenerates in a similar manner
as do the hairs in other species. The longitudinal division of cells in
the upper parts of the filaments is undoubtedly connected with the
formation of the short, lateral, often secund, gametangia characteristic
of nearly related species in Hecatonema and in Compsonema, for in a
few instances such gametangia were observed in old specimens in which
the gametes had. to a large degree, escaped from the characteristic
gametangia.

Hecatonema Lawsonii sp. uov.
Plate 40, figures 5-7

Frondibus pulvinulos microscopicos, 250-500;a diam., linea exteriore
irregulares formantibus; parte prostrata filamentis maxime contortis,
profuse ramosis, subterminaliter furcatis composita; filamentis erectis
stratum compactum valliforme filamenta pilifera numerosa sparsa
includens formantibus, 0.75-1.5 mm. longis; cellulis filamentorum
repentium radianter tangentialiterque divisis, in centro thalli pseudo-
parenchyma formantibus, forma irregularibus, 4-5ju, diam. ; cellulis
filamentorum piliferorum basi quadratis et vaginatis, superne usque
ad 25-plo longioribus quam latis; zoosporangiis ignotis; gametangiis
cylindricis usque ad leviter fusiformibus, 25-30;u, longis, 5-6. 5/^ latis;
loculis multis per parieties longitudinales et obliquos formatis.

Growing on the sporophylls of Nereocystis Luetkeana. Uyak Bay,
Alaska. Type, Setchell and Lawson, no. 5131 (Herb. Univ. Calif.,
no. 99357), August.

Myrionema vulgar e Collins, Holden and Setchell, Phyc. Bor.-Amer.
(Exsicc), no. 924 (non Thuret). Myrionema strangulans Setchell and
Gardner, Alg. N. W. Amer., 1903, p. 249 (non Grev.).

This plant superficially resembles very closely Myrionema foecun-
dum f. simplicissimum S. and G. Microscopic examination, however,
reveals several important differences. The creeping filaments are
unique. The branches are very numerous, come off at wide angles and
seem, for the most part at least, to be subterminal, or if the terminal

380 University of California Publications in Botany [Vol. 7

cell splits, one of the dichotomy very frequently fails to develop till
much later. IMany of the cells in the center, and even toward the
periphery, divide radially and perpendicular to the surface of the
host, forming a pseudoparenchymatous layer. Thus this species,
strictly speaking, cannot be said to be distromatic, since the distromatic
condition of a frond is really brought about by radial divisions of the
cells of filaments, but parallel to the surface of the host. The character
of the gametangia and of the creeping filaments will not permit of its
being placed with Myrionema vulgare Thuret as further described and
figured by Sauvageau (1897, p. 186, et seq.). On account of the radial
divisions of the creeping filaments and the prevailing biseriate con-
dition of most of the gametangia, we are placing this species with the
genus Hecatonema and dedicating it to Professor A. A. Lawson, one
of the collectors.

LITERATURE CITED

Sauvageau, C.

1897. Sur quelques Myrionemacees. Ann. sci nat., 8 ser., Bot., vol. 5, pp. 161-

288.

Setchell, W. a., and Gardner, N. L.

1903. Algae of Northwestern America. Univ. Calif., Publ. Bot., vol. 1,
pp. 165-418, pis. 17-27.

PLATE 40

Hecatonema clavatum S. and G.

Fig. 1. A section through a typical mature plant showing penetrating
rhizoids from the basal layer, the clavate, sterile, erect filaments, and typical
gametangia. X 250.

Fig. 2. The same as figure 1, but showing " ascocysts. " X 250.
Fig. 3. A small fragment without rhizoids. X 125.
Fig. 4. A segment of the base at its margin. X 250.

Hecatonema Latvsonii S. and G.

Fig. 5. Two segments of the base showing frequent radial divisions of the
cells. X 250.

Fig. 6. A young plant before any erect filaments have begun to develop.
X 250.

Fig. 7. Typical gametangia and true hairs. X 250.

[382]

UNIV. CALIF. PUBL, BOX. VOL. 7

[SETCHELL-GARDNER ] PLATE 40

PLATE 41

Hecatonema variaMle S. and G.

Fig. 1. Five fragments of plants taken near the margin of the frond. X 125.
Fig. 2. Three fragments, older than figure 1, showing secondary growth
within the empty gametangia. X 125.

Fig. 3. A group of sessile gametangia. X 125.

Fig. 4. Terminal part of a creeping filament seemingly showing the early
stages in the development of the zoosporangia. X 125.

Fig. 5. A part of a creeping filament showing the heterogeneous develop-
ment of erect filaments. X 125.

Figs. 6, 7, 8, 9. Typical fragments of mature fronds. X 125.
Figs. 10, 11. Fragments with zoosporangia, X 125.
Fig. 12. A segment of the base. X 125.

[384]

UNIV. CALIF. PUBL. BOT. VOL. 7 [SETCHELL-GARDNER ] PLATE 41

1922] Setchell-Gardner : Phycological Contributions 385

y. NEW SPECIES OF PYLAIELLA AND
STREBLONEMA

Pylaiella tenella sp. nov.
Plate 42, figures 9-11

Frondibus late diffiisis, simplicibus, 0.5-0.75 mm. altis, siiperne
longe-attenuatis, non piliferis, per filamenta curta, ramosa, contortaqne
affixis; eellulis 7-10/* diam., 1-2.5-plo. longioribiis quam latis; chro-
matophoris juvenis, singulis, intemipto-taeniatis, cellulam fere im-
plentibus, ultimo in frnstis distinctis separantibus ; zoosporangiis
longe-catenatis snbterminalibus. eellulis numerosis assimilatoribus in
seriebus per divisiones longitudinales zoosporangiis bina aut quaterna
f ormantibus ; gametangiis subterminalibus.

Growing on Plcurophycus Gardneri Setchell and Saunders, the
plants forming minute tufts, which later become confluent. Neah Bay,
near Cape Flattery. AVashington. Type, Gardner, no. 3818 (Herb.
Univ. Calif., no. 207010), May.

This species of Pylaiella is the most diminutive of all the known
species of the genus, rarely attaining the length of one millimeter, its
nearest rival, in this respect, being P. nana of Kjellman, from the
Norwegian Polar Sea. It differs from that species in size, being only
about one-half as large, in not being branched, in having subterminal
gametangia in a long series, instead of terminal branched ones as
described and figured by Kjellman for P. nana and in having double
and quadruple zoosporangia formed by longitudinal and cross divisions
of fructiferous cells. We have also observed this condition highly
developed in forms of P. littoralis. Double zoosporangia have been
observed by Borgesen (1920. p. 433) in P. fulvescens (Schousb.)
Bornet from the Danish West Indies. It evidently belongs to the
Panthocarpus group of the genus as established by Skottsberg (1915,
p. 158). The gametangia are formed by the transformation of vege-
tative cells, usually beginning relatively few cells back of the apices
of the filaments, a considerable number of cells transforming simul-
taneously. The transformation continues in both directions until one-
half to three-fourths of the filament is converted into gametangia.
They vary greatly in size, some producing as many as thirty-two loculi
from a single assimilating cell. They may be continuous or discon-
tinuous, certain assimilating cells in the series not being transformed.
The cell walls in the formation of loculi frequently are quite oblique.

386 University of California Publications in Botany [Vol.7

Zoosporangia occupy very niiu-li the same position in the filaments as
tlie irametangia. Tliey are subterminal, long-catenate, but seem to be
i"ornied more nearly simultaneously than are the gametangia.

These plants are quite abundant on the blade of the host-plant at
Neah Bay. We have not observed it growing elsewhere, although the
host is abundant in the vicinity of the Strait of Juan de Fuca. They
are, however, very inconspicuous, forming small, more or less con-
tinuous expansions, barely recognizable when the host plant is wet.

Pylaiella unilateralis sp. nov.

Plate 42, figures 6-8

Filamentis erectis per filamenta repentia, irregularia, plus minusve
ramosa afifixis, floccosis. leviter clavatis, 4-7 mm. altis, basi 19-22/x
diam., apieali 28-32/a diam., ramis in feme plerumque alternis spar-
sisque, superne secundis, e cellulis 1-2-plo longitudinaliter divisis
oriendis; cellulis 2-2.5-plo longioribus quam latis, basi cylindricis,
superne plus minusve quadratis doliiformibusque ; chromatophoris
tenuibus, irregulariter taeniatis et plus minusve connectis; zoospor-
angiis eodem tempore maturantibus, 7-12-catenatis, in ramulis plerum-
que terminalibus, sphaeroideis, 24-28/a diam. ; gametangiis ignotis.

Growing on rocks in shallow pools, near high-tide limit. Sunset
Beach, near the mouth of Coos Bay, Oregon. Type, Gardner, no. 2748
(Herb. Univ. Calif., no. 207011), May, 1914.

The secund branching, together with the longitudinal divisions of
the cells from which the secund branches arise forming a poly-
siphonous region are the chief distinguishing characteristics of this
species. The branching below is sparse, alternate or very rarely
opposite, and the branches are relatively long and attenuated. These
branches may give rise to terminal zoosporangia, or to a few short
alternate ramuli which in turn produce the zoosporangia. ]\Iany of
the main filaments become slightly enlarged toward their outer ends,
the cells becoming nearly quadrate, their walls thickened, and their
contents very dense. An average of about thirty-five cells, though
frequently as many as sixty-five, are thus transformed. A few cells
of the ends of the filaments are not transformed and soon disintegrate,
thiLs producing a distinctly clavate filament. The region of trans-
fonnation seems to be a new meristem of a peculiar nature. ]\Iauy
of the cells divide once or, less frequently, twice, by longitudinal planes
as a rule, but occasionally the dividing planes are at right angles
separating the cell into four parts. The unique feature resulting from
these divisions is the lack of further growth of the cells. Usually one

1922] Setchell-Gardner: Phycological Contributions 387

of the cells resulting from longitudinal division gives rise to a branch,
shorter or longer, but similar to branches arising in the regular
manner. The branches are typically secund, but may rarely come from
the opposite side of the filament. ]\Iany of the main filaments become
much curved in this branching region.

Streblonema Porphjnrae sp. nov.

Plate 44, figure 6

Frondibus plerumque endophyticis ; filamentis prostratis maxime
tortuosis, inter parietes cellularum hospitis penetrantibus, copiose
ramosis; filamentis erectis ultra superficiem hospitis leviter protrud-
entibus, passim ramosis; filamentis piliferis ignotis; cellulis fila-
mentorum repentium 3-4|U, diam., forma irregularibus ; zoosporangiis
ignotis; gametangiis filamenta erecta terminantibus, ultra superficiem
hospitis leviter protrudentibus, forma fusiformibus ad irregularibus,
25-35jU, longis, S-S/x latis.

Growing in the parenchymatous base of Porphyra naiaduni on eel-
grass. Pacific Grove, California. Type, Gardner, no. 4686 (Herb.
Univ. Calif., no. 207008), December.

Strehloncma Porphyrae is an exceedingly diminutive, though appar-
ently distinct, species, confined, so far as we know at present, to the
cushion-like bases of Porphyra naiadum. It ramifies deep into the
tissue, apparently not entering the cells.

Streblonema myrionematoides sp. nov.

Plate 44, figure 7

Frondibus microscopicis ; partibus prostratis exiguissime evolvatis,
inter srata duo aut tria externa cellularum hospitis leviter penetrant-
ibus; filamentis erectis plus minusve ad superficiem hospitis fascieu-
lato-ramosis ; 65-80/a longis, plerumque fructuosis. pilis exiguis; cellulis
filamentorum penetrantium 4-5/t diam., forma irregularibus; zoo-
sporangiis ignotis; gametangiis numerosis, cylindricis, 50-65;U longis,
4.5-6.5/x latis ; loculis uniseriatis.

Growing on the blade of Laniinaria Andersonii. Moss Beach. San
Mateo County, California. Type, Gardner, no. 4622 (Herb. Univ.
Calif., no. 207005), July.

It is an extremely perplexing problem to decide upon the generic
position of forms such as the one we have named and described above.
It has close affinities with the three genera, Streblonema, Ectocarpus,
and Myrionema. The erect fronds exterior to the host and more or less
branched, with relatively scanty attaching portions, constitute the

388 University of California Publications in Botany [^''ol. 7

vegetative portion of iin Ectocarpus of microscopic size. The game-
tangia are strict I \- those of a typical Myrionema, but it lacks the disk-
shaped basal layer of filaments spreading out on the surface of the
host, characteristic of that genus. The extremely reduced character
of the penetrating portion does not speak well for the genus Streblo-
ncma. On the whole, we feel that with our present criteria for these
genera it Ix'st agrees with the characters of the genus Streblonema,
M'here we are placing it. It penetrates the uninjured host, but only
to a slight depth. The plants, however, are usually so congested that
their growth soon crowds the surface layer of the host cells to such an
extent that they die and disintegrate, there being no evidence that the
associate penetrates them and absorbs their material. Tlic palisade-like
stratum of gametangia suggests very strongly the Myrionema character
which is the reason for the specific name.

Streblonema penetrale sp. nov.

Plate 44, figures 3, 4

Frondibus stratum continuum pulvijiatumque forma magnitudine-
que indefinitum supra stipitem hospitis formantibus ; parte penetrante,
filamentis parce ramosis in hospitem comparate profundi penetrantibus
et ad superficiem hospitis ]ierpondipularibus composita ; filamentis
erectis ad superficiem hospitis fasciculato-ramosis, 70-125/x longis,
supra leviter attenuatis, non piliferis; cellulis fllamentorum pene-
trantium cylindricis ad irregularibus, 6.5-8ja diam., 3-5-plo longior-
ibus; cellulis fllamentorum erectorum cylindricis ad leviter doliiform-
ibus, 6.5-8/A diam., 1.5-2.5-plo longioribus; chromatophoris taeniatis;
zoosporangiis ignoti.s; gametangiis cylindricis usque ad obtuse-fusi-
formibus. plus minusve irregularibus, 30-40/x longis, S-llju, latis ; loculis
uniseriatis.

Growing on the stipes of nesperopliycus Harveyamis. Pacific
Grove, California. Type, Gardner, no. 4677 (Herb. Univ. Calif., no.
207007), December.

Streblonema penetrale, like S. my rionemat aides above, is a difficult
species to classify. In its method of development and general struc-
ture it approximates to 8. myrionematoides closely. The penetrating
portion is much more highly developed than in that species, extending
into the host among the cells to a depth of four or five times as great
as the part which extends beyond the surface. The size of all the parts
is, in general, greater than in 8. myrionematoides. No horizontal
filaments are present. The penetrating filaments are mostly perpen-
dicular to the surface.

1922] Setchell-Gardner : Phycological Contributions 389

Streblonema vorax, sp. nov.

Plate 44, figures 1, 2

Frondibus microscopicis, filamentis prostratis profuse ramosis,
maxime tortuosis, inter cellulas epidermales hospitis penetrantibus, in
interiore hospitis diffundentibus et celhilarnm parenchymaticarum
parietes dissolventibus, easdem complentibus. et earam partem inter-
iorem devorantibus ; cellulis 6-10/* longis, 5-8/x latis ; filamentis erectis
400-600/* longis, 6.5-8/* diam., superne in pilis attenuatis, ad aut prope
snperficiem hospitis fasciculato-ramosis, massam compactam celhilarum
formantibiis ; zoosporangiis late clavatis, 60-100/* longis, 15-30/* latis;
gametangiis niimerosis, lateralibus, sessilibus aut pedicellis brevibus
suffultis, anguste cylindricis, obtusis, 40-70/* (ad 100/*) longis. 7-9/*
latis; loculis plerumque uniseriatis.

Growing on the outer ends of the leaves of eel-grass, in the lower
littoral and upper sublittoral belts. Sitka, Alaska. Type, Gardner,
no. 3968& (Herb. Univ. Calif., no. 207003), July.

Of all the species of Streblonema thus far discovered on our coast,
S. vorax is the most destructive to the host. Although the penetrating
filaments have abundance of ehromatophores, the plants seem to be in
a large degree parasitic. The cells of the host are closely compacted,
and have thick walls, yet they are devoured in large quantities. We
suggest the possibility of the secretion of enzymes with digestive power
which act upon the cell wall and its protoplasm, after which they are
absorbed. This plant is found in company with several other small
Melanophyceae, Chlorophyceae, and Rhodophyceae, none of which
penetrate the host.

Streblomena scabiosum sp. nov.
Plate 44, figure 5

Frondibus pustulas rotimdas aut ellipticas magnitudine indefinatas
in superficie hospitis formantibus; partibus penetrantibus filamentis
copiose ramosis primo inter cellulas penetrantibus et eas necantibus,
ultimo interiorem cellularum occupantibus et eversionem totam effici-
entibus; filamentis erectis supra superficiem hospitis extendentibus,
aut simplicibus aut plus minusve basi fasciculato-ramosis. 50-80/*
altis; cellulis filamentorum repentium forma magnitudineque maxime
irregularibus ; cellulis filamentorum erectorum cylindricis. 4.5-5.5/i
diam., 1-2.5-plo longioribus; chromatophoris taeniatis; zoosporangiis
ignotis ; gametangiis cylindricis, sessilibus aut brevi-pedicellatis, supra
superficiem hospitis extendentibus, 40-60/* longis, 4.5-6/* latis.

Growing on the stipe of Nereocystis Luetkeana. Cast ashore near
the "Cliff House," San Francisco, California. Type, Gardner, no.
4628 (Herb. Univ. Calif., no. 207006), August.

390 University of Calif orma Publications in Botany [Vol. 7

Strchlonona scabiosum forms iimrkcd scrofulous-like areas on the
lower part of the stipe of the host. It has a disastrous effect upon the
hast, destroying its cells as far as it penetrates, and seems constantly
to spread by new infections around the margin of the affected area.
The cause of the death of the cells of the host has not been investi-
gated. In habit of growth and general form the species seems related
to S. myrioncmatoides and to S. penetrale.

Streblonema evag'atum sp. nov.

Plate 42, figures 1-5

Frondibus maculas orbiculares 1-2 cm. diam. formantibus; fila-
mentis repentibus irregularibus, mvilte ramosis, inter bases sporangi-
ornin hospitis penetrantibus: filamentis erectis fasciculato- ramosis,
plerunique in liospite subniersis, ad apicem basimque levissime attenu-
atis, 190-230ju, altis; eellulis filamentorum repentium 3.5-4/a diam.;
cellulis filamentorum erectorum basim 4jU, diam., 2-3.5-plo longioribus,
in parte latissimo 6.5-7.5/i, diam., 1-2-plo longioribus, eylindrieis, ad
dissej)imcnta loviter constrictis; chromatophoris taeniatis, in cellula
quaque 1-2; zoosporangiis ignotis; gametangiis numerosis, eylindrieis,
lateral ibus, sessilibus aut in pedieellis eurtis suffultis, 65-80jli longis,
5.5-6. 5yu, latis; loculis uniseriatis.

Growing in the blades of Laminaria Andersonii, upper sublittoral
belt. Cypress Point, Monterey County, California. Type; Gardner,
no. 4688 (Herb. Univ. Calif., no. 207009), December.

This species has a habit of growth very similar to that of Streblo-
nema pacificum, but the plants cover much more extensive areas which
are quite certainly not infested by a single plant, as may be the case
in that species. Their presence seems to affect the host materially, to
a large extent destroying its sporangia and causing discoloration. It
is possibly partially parasitic.

Streblonema rugosum sp. nov.

Plate 43, figures 5-7

Frondibus microscopicis in superficie hospitis areas rugosas extensas
offifiontibus ; filamentis repentibus inter cellulas superficiales hospitis
pciK'trantibus et filamenta erecta, faseiculato-ramosa, fructifera et
j)ilifera emittentibus ; cellulis filamentorum et erectorum et repentium
forma magnitudineque irregularibus; chromatophoris taeniatis; zoo-
sporangiis ignotis; gametangiis obtuse fusiformibus, in filamentis
erectis pluriiiiis terminalibus, 18-24/i, longis, 5-6. 5/a latis; loculi uni-
•seriatis, in serie longitudinali quaque 4-6.

1922] Setchell-Gardner : PJiycological Contributions 391

Growing on the blade of Alaria tenuifolia. Friday Harbor, "Wash-
ington. Type, Gardner, no. 4041 (Herb. Univ. Calif., no. 207004),
July.

This species, like Strehlonema myrionematoides and S. scabiosum,
has the larger part outside of the host. It penetrates to a considerable
depth among the cells of the uninjured host, later apparently mechanic-
ally causing the death of a few surface cells and an abnormal gro^rth
of others, giving to the surface a rugose appearance. The plants seem
to associate in small groups of indefinite shape and size, often con-
fluent and, although small, can readily be detected by the peculiar
appearance they give to the host.

Streblonema transfixum sp. nov.

Frondibus maculas 5-8 mm. diam. f ormantibus ; filamentis repent-
ibus inter cellulas hospitis profunde penetrantibus, distortis, irregu-
lariter ramosis, 4— Sju, diam., filamenta dispersa, curta erecta, eramosa,
e superficie hospitis protusa et a gametangiis terminata emittentibus ;
cellulis filameutorum erectorum cylindricis, 7-9//, diam., 0.75-1. 5-plo
longioribus; chromatophoris taeniatis, parietem cellulae proxime
tegentibus; zoosporangiis ignotis; gametangiis cylindrico-conicis,
obtusis, 40-60ju, longis, 8-12/* latis; loculis 1-2-seriate.

Growing on Desmarestia ligtdata f. Jierhacea. San Pedro, Cali-
fornia. Type, Gardner, no. 1992a (Herb. Univ. Calif., no. 207001),
September.

Strehlonema trans fixum, here described, forms definite, discolored
areas on the surface of the host, readily detectable. The species is
described from dried material. The cells of the host are much collapsed
and do not straighten out when soaked and boiled, hence the details
of the species are not so thoroughly and satisfactorily presented as is
desirable. The plants penetrate among the cells of the host and seem-
ingly pass clear through it. Just how much constitutes a single plant
cannot at present be made out.

Streblonema corymbifenim sp. nov.

Plate 43, figure 8

Frondibus microscopicis, filamentis irregulariter alterne ramosis
et inter cellulis hospitis penetrantibus compositis, ramulis fructiferis
plane aggregatis, fasciculos corymbiformes prope superficiem hospitis
formantibus; cellulis plerumque cylindricis, pro parte irregularibus,
4— 5/A diam., 1.5-4-plo longioribus; chromatophoris taeniatis, parietem
cellulae non tegentibus; zoosporangiis ignotis; gametangiis cylindricis

302 rnliursifn of (UiJifornia J'uhlications in Bofaiiii [Vou 7

us(|U(' iul Icvitci- riisiroriiiihiis, olitiisis, in pedieellis ciirtis suifiiltis
a^irrcirjitis aiit in filanientis repentibus prope snperficiem hospitis
sessilibus, 2'->-'A')fi longis, 4.5-5.5/a latis; loculis uniseriatis, septis
|)lciMiin(|U(' (ililir|nis.

Growing on (Uunagloia Andersonii (Farlow) S. and G., in company
witli Slrcbloncma anonialum and S. Johnstonae. San Pedro, Cali-
fornia. Type. :\Irs. II. D. Johnston, no. llGa (Herb. Univ. Calif.,
no. 94663), August.

While studying the material of Nemalion Andersonii Farlow. col-
lected by ]\Irs. II. 1). Johnston at San Pedro, California, in 1899 and
deposited in the Ilerbariuiu of the University of California under
no. 94663. some specimens of p]ctocarpaceae were incidentally observed.
^Yllen we came to study our material of this family, these specimens
were given careful scrutiny with the result that three species new to
science were detected on a single specimen of the host, viz., Strehlonema
corymhiferum, S. anomalum, and S. Johnstonae, all of which are pre-
sented for the first time in this paper.

The habit of each and the size of all the parts are so characteristic
that, notwithstanding their intimate association on the same host, they
can readih^ be distinguished. S. corymhiferum is the most delicate of
the three. The very frequent grouping of the gametangia into corymb-
like clusters is one of the most prominent distinguishing characters of
the species. Other species of this genus are as small or even smaller.
In size of parts, somewhat in habit and in habitat, it resembles S.
mimitissimum Saunders, found growing at Sitka. Alaska, on "Lieh-
mnnnia sp." (Saunders, 1901, p. 416).

Streblonema anomalum sp. nov.

Plate 43, figures 1-3

Frondibus microscopicis, plus miinisve confluentibus; filanientis
partis prostratae inter cellulas hospitis penetrantibus moderate et
irregulariter ramosis. multe contortis, usque ad 16/x diam. ; filanientis
erectis siniplicilms. plerumque cylindricis, pro parte olitusis et pro
parte pilifcris, 200-250/^ longis, ultra superficiem hospitis exten-
dentibus; filamentis piliferis sparsis, e filanientis repentibus oriundis;
cellulis filamentorum repentium plus minusve in partibus senilibus
1-2-plo lougitudiualiter divisis. filamentum polysiphoiioum produeent-
ibus. 18-24/j. diaiii.; cellulis alteris I'orma leviter irregularihus. 8-1 Iju,
diam.. admodum ad dissepimenta leviter constrictis; cellulis fila-
mentorum ereetorum cylindricis. 8-9ju, diam., quadratis; chromato-
phoris singulis, taeniatis, in filanientis ereetis continuis. in filamentis
repent ilms i)lus minusve interruptis; zoosporangiis ellipsoideis, ovoideis

1922] Setchell-Gardner : Phycological Contributions 393

aut proxime sphaericis, 40-60ju. longis, 28-35/a latis, in filamentis
repentibiis sessilibus; garaetangiis cylindrico-conicis, plerumque
obtusis, 50-lOOfi longis, 10-16ju. latis, sessilibus ant in filamentis
repentibus longe-pedicellatis singulis aut seeundis.

Growing in Cumagloia Andersonii (Farlow) S. and G. in company
with StreUonema corymhif erum and S. Johnstonae. San Pedro, Cali-
fornia. Type, Mrs. H. D. Johnston, no. 115& (Herb. Univ. Calif.,
no. 94663), August.

StreUonema fasciculatum Saunders, Phyc. j\Iem., 1898, p. 148 (not
of Thuret).

Examination of a small piece of authentic material of S. fascicu-
latum Saunders collected at San Pedro, California, in August, 1896,
revealed the presence of specimens of the same three species mentioned
under S. corymhif erum of this paper, collected by ]\Irs. Johnston at
the same place and in the same month. S. anomalum is undoubtedly
the one to which Saunders refers as *S'. fasciculatum Thuret. It does
not seem, however, to be this species, if we have a correct conception
of what S. fasciculatum Thuret includes. This species was published
in Le Jolis, Algues marines de Cherbourg, no. 100, and in Liste, 1863,
p. 73. Thuret quotes as a synonym, S. voluhilis Pringsheim (Beitr.
Morph. Meeres Alg., p. 13, plate 3, fig. B, read in 1862) . Pringsheim 's
figure of voluhilis shows the gametangia fasciculately branched, and he
states that this charatcer is one of the chief characters of the genus
Strehldnema. This figure has been much quoted since. He gives no
measurements of the parts, neither does Thuret.

Reinke (Algenfl., 1889. p. 41) recognizes Strehlonema as a sub-
genus of Ectocarpus. He recognizes Pringsheim 's plant, quoting the
above figure, but since voluhilis was already occupied by Crouan (1867,
p. 161) and fasciculatum was occupied by Harvey (Phyc. Brit., pi.
273), he renamed the plant, calling it Pringsheimii. Hauck (1884,
p. 323) seems to have been the first to give measurements of the
various parts of the plant. He lists it under Strehlonema, cites the
above literature of Thuret, and quotes S. voluhilis Pringsheim. Con-
sidering Pringsheim 's plant, recognized by Reinke and Hauck, as
being the same as Thuret 's, and taking Pringsheim 's figure and
Hauck 's measurements as being correct for Thuret 's S. fasciculatum,
then our plant is distinct and undescribed.

There is a little doubt at present in our minds whether the plant
we are here describing as Strehlonema anomalum. is one or two species.
We do not find the filaments bearing the zoosporangia in the type

394 Univcrsify of California Puhlications in Botany [Vui,. 7

material examined to be like those figured by Saunders. His figures
sliow tlh' main filaments as being like those of all the known Sfrehlo-
nrmas, monosiphonous, while those in both collections of material which
we have examined are uniformly polysiphonous, that is, the cells of the
main central parts of the thallus are all divided once or twice length-
wise. This never takes place in the plants which bear gametangia and
which are intimately associated with them.

The measurements, method of branching of the two sets of plants,
and tlu'ir chromatophore characters are practically the same. At
least two interpretations of this polysiphonous phenomenon appeal to
us. They may represent a polymorphic state, in which a nonsexual
plant differs in form from a sexual plant of the same species, a
unique condition in the genus Strehlonema, or the polysiphonous con-
dition of the main filaments, found only in the nonsexual plants, may
represent a character belonging to a wholly different genus, and hence
it is a new species of that genus. Until a more extensive study of fresh
material can be made, we feel that it is best to take the former view,
and place it in a new and polymorphic species of Strehloncma. We
have amended the family Ectocarpaceae to include species with this
poh'siphonous condition.

Streblonema Johnstonae sp. nov.

Plate 43, figure 4

Frondibus microscopicis ; filamentis repentibus moderate ramosis.
ramis alternis aut oppositis, raro leviter secundis ; filamentis erectis
plerumque simplicibus, apice basimque leviter attenuatis, supra super-
ficiem hospitis extendentibus, pro parte, brevi-pilif eris ; cellulis fila-
mentorum primariorum repentium forma plus minusve variabilibus,
plerumque doliiformibus, 12-18/ji diam., 2.5-5-plo longioribus; cellulis
filamentorum erectorum pro parte latissima usque ad 24/a diam., fere
cylindricis, ad dissepimenta constrictis ; chromatophoris tenuibus,
parietalibus. in cellula quaque singulis, parietes cellularum fere
tegentibus; zoosporangiis ignotis; gametangiis conicis usque ad in-
aequaliter fusiformibus, 90-130/x longis, 28-36/j, latis, plerumque brevi-
pedicellatis in filamentis prope superficiem hospitis repentibus positis.

Growing in Cumagloia Andersonii (Farlow) S. and G. in company
with Streblonema corymhiferum and S. anomalum. San Pedro, Cali-
fornia. Type, Mrs. H. D. Johnston, no. 115c (Herb. Univ. Calif.,
no. 04663), August.

Of the three plants found ramifying among the filaments of the
host mentioned above, Strehlonema Johnstonae is the most robust in all

1922] Setcliell-Gardner : Phycological Contributions 395

of its parts. It can readily be distinguished from the other two species
with which it is associated by the large size of the gametangia. No
zoosporangia have been observed. We take pleasure in dedicating this
species to ]Mrs. H. D. Jolmston, who collected the host and donated it
to the Herbarium of the University of California with several other
interesting forms from San Pedro, California.

Streblonema aecidioides f. pacificum forma nov.

Plate 44, figures 8, 9

Frondibus microscopicis, supra superficiem hospitis ut maculas
elevatas 75-150/i, diam. notatis ; parte vegetativa stratum plus minusve
parenchymaticnm exigue infra stratum superficiale hospitis positum
et infra filamenta pauca radiciformia in hospitem profundiore pene-
trantia emittens formantibus; filamentis erectis omnibus fructiferis
filamentis paucis, piliferis in centre frondis positis exceptis; cellulis
filamentorum piliferorum 4— 5.5/x diam.. inferne quadratis, superne
5-8-plo longioribus, evaginatis; zoosporangiis( ?) anguste clavatis,
sessilibus, 22-28/i, longis. apice 8-12/i, latis; gametangiis numerosis,
dense aggregatis, eylindricis, in strato prostrato sessilibus, 45-55|U,
longis, 5-6.5|Li latis ; loculis uniseriatis.

Growing within the lamina of HedopJiyllum sessile (Aresch.)
Setchell, near the outer end. Neah Bay, Washington. Type, Gardner,
no. 3866a (Herb. Univ. Calif., no. 207002), May.

Streblonema aecidioides f. pacificum seems very closely related to
the Ectocarpus aecidioides of Rosenvinge (1893, p. 894), found grow-
ing in Greenland on Laminaria longicruris and L. Groenlandica. It
differs only in minor details as to the dimensions of the parts. Regard-
ing the zoosporangia we have to speak with uncertainty. As figured
by Rosenvinge, the Greenland plant has them well developed and pro-
ducing zoospores. They are in distinct "aecidia" and apparently on
distinct nonsexual plants. In our species they likewise appear to be
on nonsexual plants. In ours there is no indication of the production
of zoospores. They are possibly too young, or possibly they are
abortive organs so commonly met with in various other genera on our
coast, the nature of which is still an open question. This form seems
to enter the host from the surface and after penetrating to the second
layer of cells spreads out horizontalh^ between the surface layer and
the second layer. Later, filaments arise from the under side of this
layer and penetrate among the cells of the host, apparently never
entering them. Finally from the upper surface each cell in the central
region of the layer gives rise to a filament and the mass acting together

396 Univrrsil!/ of California I'ublicaiions in Botany [Vol.7

lifts up llic surfnci' layer oi" cells of the host, forming a small blister
which finall\ ruptures, as in the case of Rosenvinge's plants. The
erect filaments are almost simultaneously transformed into gametangia,
exeej)! a few in llic center which develop into hairs.

P^slie (1894. j). 167, 23 in reprint) describes forms of the same
species found growing on Laminaria saccharina at Kjelvik and at
Lyngii near Tronis0, Norway. These he listed under Ectocarpus
(Strcblonema) accicUoides Rosen vinge. The measurements of the
parts of his forms average, in general, a little larger than those of
Rosenvinge. De-Toni (1895, p. 577) cites these Arctic plants under
Strcblonema aecidioides Rosenv. Foslie gives 80/i, as the extreme
length of the gametangia.

Streblonema investiens (Collins) comb. nov.

Fronds occupying indefinite areas on the host ; creeping filaments
irregularly branched, often curvdng outward and bearing on the out-
side short, simple, or sparsely branched filaments ; hairs sparse ; cells
of creeping filaments o-S/x diam., 1-2 (up to 4) times as long, swollen,
or cylindrical; cells of . the ramuli Gfi diam., 1-2 times as long; cells
of the hairs S/x diam. ; chromatophores discoid, small, several in a cell ;
zoosporangia ovoid, sessile or on 1-celled pedicels on both the creeping
filaments and the ramuli, 20/x long, 15fx broad ; gametangia cylindrical,
25-40/A long. 8-1 0/i, broad; loculi mostly uniseriate, gametangia and
zoosporangia growing on the same plant.

Growing in the fronds of HelmintJiocladia calvadosii (Lamour.)
Setchell. San Pedro, California. July.

Strepsithalia investiens Collins, in Collins, Holden, and Setchell,
Phyc. Bor.-Amer. (Exsicc), no. 738,

It seems that the chief distinction between the genera Strepsithalia
Sauvageau and Streblonema Derb. and Sol. is the secretion by Strepsi-
thalia of a rather copious gelatinous sheath investing the entire plant,
particularly the exposed portions, the ramuli. Since we are not able
to demonstrate the presence of such a sheath, even to the slightest
degree, we have thought it best to place Collins' Strepsithalia investiens
in the genus Streblonema.

1922] Setchell Gardner: Phycological Contributions 397

LITERATURE CITED

BORGESEN, F.

1920. The marine algae of the Danish West Indies, vol. 2, Ehodophyceae,
with addenda to the Clilorophyceae, Phaeopliyceae, and Ehodo-
phyceae. Copenhagen.
Crouan, p. L., and H. M.

1867. Florule du Finistere, conteuant les descriptions de 360 especes nouvelles
de sporogames de nombreuses observationes et une synonymic des
plantes cellulaires et vasculares qui croissent spontanement dans
ce department. Paris.
De-Toni, J. B. (or G. B.)

1895. Sylloge x\lgarum, vol. 3, Fueoideae.
FOSLIE, M.

1894. New or critical Norwegian algae. Det. Kgl. norske Vidensk. Selsk.
Skrifter., pp. 144-175 (1-31 in reprint). Trondhjem.
Harvey, W. H.

1851. Phycologia Britannica, vol. 3. London.
Hauck, F.

1884. Die Meeresalgen Deutschlands imd Oesterreichs, in Dr. L. Eaben-
horst's Kryptogamen-Flora von Deutschland, Oesterreich und der
Schweiz, vol. 2, 8 Lieferung, p. 316. Lei]izig.
Le Jolis, a.

1863. Liste des algues marines de Cherbourg. Mem. Soc. Imp. Sci. Nat. de
Cherbourg, vol. 10. Paris.
Algues marines de Cherbourg (Exsicc), fase. 5. Paris.
Pringsheim, N.

1862. Beitrage zur Morphologic der Meeresalgen. Physikal. Abhandl. der
Kon. Akad. d. Wissensch. Berlin.
Eeinke, J.

1889. Algenflora der westliehen Ostsee deutschen Antheils. Sexter Bericht
der Commission zur wissenschaftlichen Untersuchung der deutschen
Meere in Kiel. I Heft. Berlin.
Eosenvinge, L. K.

1893. Gronlands Havalger. Meddelelser om Gronland, vol. 3, ]ip. 76o-9'81.
Copenhagen.
Saunders, De A.

1898. Phycological Memoirs. Proc. Calif. Acad. Sci., ser. 3., Bot., vol. 1,

pp. 147-168, pis. 12-32.
1901. Papers from the Harriman Alaska Expedition, 25, The Algae. Proc.
Wash. Acad. Sci., vol. 3, pp. 391-486. Washington.
Skottsberg, C.

1915. Notes on Pacific Coast Algae. I, PyllaielJa PosteJsiae, n. sp., a new
type in the genus PylaieUa. Univ. Calif. Publ. Bot., vol. 6, no. 6,
pp. 153-164, pis. 17-19,

PLATE 42
Streblonema evagatum S. and G.
Figs. 1, 2, 4. Fragments of plants freed from their host. X 250.
Fig. 3. A diagrammatic section.
Fig. 5. A young plant penetrating among the sporangia of the host. X 125.

PylaieUa unilateraJis S. and G.

Fig. 6. Diagrammatic sketch of a small group of plants.

Fig. 7. A filament showing method of branching and characteristic inter-
calary zoosporangia. X 125.

Fig. 8. A small fragment showing longitudinal divisions of the cells of the
main filament in the region of branching. X 250.

FylaicUa tenella S. and G.

Fig. 9. A fragment of a filament showing gametangia, and a fragment to
the right of it showing ehromatophores. X 500.

Fig. 10. A fragment showing zoosporangia. X 500.

Fig. 11. A fragment showing seriate intercalary^ gametangia. X 125.

[398]

UNIV, CALIF. PUBL, BOT. VOL, 7

[SETCHELL-GARDNER ] PLATE 42

PLATE 43
Strehlonema anomalum S. and G.

Fig. 1. A part of a plant freed from its host bearing gametangia. X 250.

Figs. 2, 3. Fragments of plants bearing zoosporangia, some of which contain
mature zoospores. The cells of the main filaments are divided longitudinally.
X 250.

Strehlonema JoJinstonae S. and G.

Fig. 4. Parts of plants showing characteristic creeping and erect filaments
and gametangia. X 125.

Strehlonema rugosum S. and G.

Fig. 5. Section through the host perpendicular to its surface, showing the
papillate character caused by the presence of Strehlonema. Diagrammatic.
Fig. 6. A surface view of figure 5. Diagrammatic.

Fig. 7. A section showing the structure of the Strehlonema plants and their
relation to the host. X 250.

Strehlonema corymhiferum S. and G.
Fig. 8. Fragments of typical plants freed from their host. X 250.

[400]

UNIV. CALIF. PUBL. BOT. VOL. 7

[SETCHELL-GARDNER ] PLATE 43

PLATE 44
Streblonema vorax S. and G.

Fig. 1. A section through the host showing the presence of the parasite and
its effect upon the host. X 125.

Fig. 2. A fragment of a plant showing the tortuous creeping filament, a
true hair, and several typical gametangia. X 125.

Streblonema penetrate S. and G.

Fig. 3. A section through the host perpendicular to its surface showing the
apparent disorganizing effect of the penetrating portion of the Streblonema.
X 125.

Fig. 4. Individual plants separated from the host. X 250.

Streblonema scabiosum S. and G.

Fig. 5. A section through the host perpendicular to its surface showing the
relation of the two plants. X 250.

Streblonema Porphyrae S. and G.

Fig. 6. A section showing the relation of this Streblonema to its host.
X 125.

Streblonema myrionematoides S. and G.
Fig. 7. Showing plants in various stages of development, X 250.

Streblonema aecidioides f. pacificum S. and G.

Fig. 8. A section tlirough an "aeoidium" and perpendicular to the surface
of the host, showing the structure and relations of the two i)lants. The section
seemingly represents young zoosporangia. X 250.

Fig. 9. The same as figure 8, but of a plant bearing gametangia. X 250.

[402]

UNIV. CALIF. PUBL, BOT, VOL. 7

[SETCHELL-GARDNER ] PLATE 44

1922] SetcJiell-Gardner: Phycological Contributions 403

VI. NEW SPECIES OF ECTOCARPUS

Since the publication of our account of the Algae of Northwestern
America (1903), more or less extensive search and study has been
carried on in that same region as well as farther south along the whole
coast of North America, with the result that a considerable number of
forms of Ectocarpus have been discovered and much more data on
previously known forms have been brought to light, necessitating some
changes in our previous account and particularly necessitating the
founding of several new and quite distinct species. We do not imagine
that the field has been, by any means, thoroughly investigated, but we
are publishing these findings up to date hoping to stimulate further
investigation. Doubtless there are many more undiscovered species
and certainly much remains to be worked out regarding life-histories of
the various kno'RTi species. The "megasporangia," "meiosporangia,"
and "antheridia" of Sauvageau (1896, 1896a, 18966, 1896c) offer
further and attractive subjects for investigation in the reproductive
phase of this group. The nature of the "zoosporangia" or "unilocular
sporangia" and the position of the plants which bear them in the life
cycle is a matter concerning which but little is known, and which
deserves critical study.

For the sake of uniformity in the method of grouping, in our
Pacific coast algological studies, we have found it necessary to name
a new Order for use in our forthcoming Part III, the IMelanophyceae,
of our 'Marine Algae of the Pacific Coast of North America." The
following diagnosis, here published for the first time, sets forth our
conception of the group.

Ectocarpales nom. nov.

Confervoid (monosiphonous) or solid Phaeosporeae of varying
dimensions, habit, and complexity, typically possessing unilocular
zoosporangia and plurilocular gametangia and with no portion of the
cell membranes turning black with can de Javelle; growth in length
strictly apical, subapical ("trichothallic"), or more or less inter-
mediate between typical forms of either ; both unilocular zoosporangia
and plurilocular gametangia rarely on the same individual in some
species, but, most commonly, borne on different individuals, thus point-
ing to an alternation of generations, reduction division taking place in
the primary nucleus of the unilocular zoosporangia, at least in some
species; gametophyte and sporophyte practically indistinguishable as
to size and complexity.

404 Umversity of California rublications in Botany IVol. 7

Eciocarpaccae Oltniaiins, .Morpli. uiul liiol. der Alg., vol. 1, 3904,
p. H50 (pro parte majore).

It seems more consistent with the present usage in other subclasses
of the Thallophyta, to consider the extended Ectoearpaceae of Olt-
manns as an Order rather than as a Family. While the families under
it jiresent a very considerable variety in the details of structural
dilTerenees, they have in common the possession of plurilocular game-
tangia and an identity in size and sliiictui-c of \n>\\\ gametophyte and
sporophyte. The Eetoearpales are closely related to the Cutleriales. 1)ut
in the latter there are some decided differences between the two
generations. From the Sphacelariales, the Eetoearpales differ in the
absence of the conspicuous apical cell and the very different cell wall.
The cell wall, or the older layers of it. changes to black in the
Sphacelariales when treated with ran de Javelle, but does not show
this reaction in any of the Eetoearpales. The LaniiiKiriales are to be
distinguished from all the other orders of the Phaeosporeae by their
intercalary region of longitudinal growth in connection with their
microscopic gametophyte. The existence of a reduced gametopliyte
in the Dictyosiphonales separates them also from the Eetoearpales.

Ectocarpus acutus nom. nov.

Plate 48, figures 36-39, and Plate 49, figures 40, 41

Frondibus 5-9 cm. altis, siccitate saturate fuscis, vivo olivario-
viridibus. phimosis; filamentis erectis, inferne plus rainusve intricatis
fasciculatisque. superne liberis, profuse ramosis. filamentis primariis
corticatis ; ramis plerumque alternis. strietis ; ramulis ultimis plerumque
secundis. acute sul)ulatis; cellulis leviter doliiformibus, 40-()0<x diam.,
0.4-2-plo longioribus quam latis (\ailgo brevioribus quam latis) ;
chromatophoris crassis, irregulariter ramosis, in cellula quaque paucis,
pyrenoideas (plures) exhibentibus; zoosporangiis ignotis: gametangiis
numerosis et regulariter dispositis. cyliiulrieo-conicis. lOO-laOjn (usque
ad 230/x) longis, 20-35/x latis.

Growing, for the most part at least, on larger ^Melanophyceae.
Puget Sound to central California (Carmel).

Ectocarpus penicillatus Saunders, Phyc. Mem., 1889, p. 155. pi. 21,
figs. 3, 4 (not of Kjellm.). Ectocarpus confervoicles f. acuminatus
Collins, in Setchell and Gardner, Alg. N.W. Amer.. 1903, p. 237, iMar.
Alg. Vancouver Is., 1913, p. 106; Collins, Holden. and Setchell, Phyc.
Bor.-Amer. (Exsicc), nos. 1033, 1127.

The type of the species is no. 2886, Gardner (Herb. I'niv. Calif.,
no. 206991), collected at Carmel. California, in May, 1915, and grow-

1922] Setchell^Gardner : Phycological Contributions 405

ing on a form of '' Desmarestia ligiUata." It seems to be the same as
the plant figured (and described) by Saunders {loc. cit.) under
Ectocarpus penicillatus. It is undoubtedly the same plant as that
described by F. S. Collins under the name E. confervoides t acumi-
natus which was founded on no. 235, Gardner (Herb. Univ. Calif.,
no. 99022) collected at Whidby Island, "Washington, growing on a
broad form of Desmarestia. Its affinities are closely with Ectocarpus
confervoides f. typicus, but it has shorter cells, sharper ramuli, de-
cidedly constricted filaments, with the gametangia more inclined
toward cylindrical than is general in that form. The chromatophores
are more distinctly band-shaped and more separated from one another
than in E. confervoides.

A few small but profusely fruiting specimens of plants seemingly
of this species have been found growing on Mesogloia Andersonii at
Neah Bay, Washington (Gardner, no. 3817). These were attached to
the host by means of penetrating rhizoidal filaments, particularly by
the descending corticating filaments. The specimens difi'er from the
typical form in that they are smaller, the gametangia are shorter and
blunter, and on separate individuals occur seriate zoosporangia( ?),
some of which are divided longitudinally into four loculi. Further
study of more material will be necessary to determine the status of
this form. For the present we are placing it with E. acutus. The
gametangia are represented on plate 49, figure 41, and the zoosporangia
on plate 49, figure 40. The zoosporangia are very numerous, and they
resemble in form those figured by Sauvageau (1896&, p. 33, reprint)
for Ectocarpus virescens. They, however, have a glistening appear-
ance, as though they were abortive or in a pathological condition, while
those represented by Sauvageau are producing zoospores. This con-
dition in which we find ours is quite common among our Pacific Coast
species of Ectocarpus. AVe have not yet seen a single specimen in
which the so-called seriate zoosporangia show any indication of pro-
ducing zoospores.

Ectocarpus afSnis sp. nov.

Plate 46, figures ]6, 17

Frondibus minutissimis, 500-700/a altis, ad hospitem per filamenta
rhizoidea pauca ramosa, decolorata, implicata affixis ; filamentis ereetis
sparse ramosis. superne longe attenuatis, pilif eris ; ramis alternis aut
oppositis ; cellulis filamentorum primariorum cylindricis, basim 24-30ju,
diam., apice 9-11/x, in longitudine per filamentum totura variabilibus,
0.75-3-plo longioribus quam latis ; chromatophoris parvis. tenuibus, in

40n T^)}ivfrsihi of California Publications in BoUuty [Vol.7

eelhilis juvcnis, anfrulato-laminacformilms, in cellnlis vetustioribns
fere re^ulariter discoideis; zoosporanfjjiis ifrnotis: gametangiis lateral-
ibus. plcruiiKine sessilibiis. iiitcnlum l-cellnlato pcdiccllatis. solitariis,
se.Miiulis aul ad ccllulaiu sin;iulaiii verticillatis, oI)tnsc-conicis ns(iue
ad spliacroidcis. 28-8V londs, 25-28/x latis, per filamentum totam
iis(|\ic ad pilos Irnniiiales seriatis.

Growing' on CallUhamnion sp. Sitka. Alaska. T3'pe, Gardner, no.
3961 (Herb. Univ. Calif., no. 206998). July.

Two eollections from Sitka, Alaska, but both made on the same day,
show a small Ecfocarpus growing on a species of CaUithamnion which
is so closely related to E. ovatus Kjellman (1877. p. 35) that perhaps
it may seem necessary at some time to refer it to that species. The
plants ai-e, however, less stout than those of Kjellman and witli more
nearly spherical gametangia. For these reasons and because of its
geographic remoteness and the somewhat different elimatic conditions,
we feel that it is desirable to describe the Alaskan plant as a distinct,
but closely related species.

Ectocarpus chantransioides sp. nov.

T'late 48, figures 28-31

Frondibus e fila mentis contortis repentibu.sque oriendis. pulvinulos
densos hemisphaericosque, 4-8 mm. altos formantibus ; ramis profusis,
inferne alternis. superne plerumque secundis; filamentis primariis
ramisque non attenuatis, apice crescentibus ; ; eellulis 8-1 0/a diam.,
inferne 2-3-plo longioribus quam latis. superne quadratis; chromato-
phoris taeniatis; zoosporangiis ignotis; gametangiis plerumque ses-
silibus. aut brevi pedicellatis. anguste cylindrieo-conicis, 80-110^1/. longis,
basim 16-20jli latis.

Growing on boulders in the lower littoral belt. Three miles north-
west of Santa ]\lonica. California. Type. Gardner, no. 2523 (Herb.
Univ. Calif., no. 206988), November.

The habit of Ectocarpus ch ant ransio ides distinguishes it at once
from all other species of the genus with the exception of E. hemi-
sphericus Saunders. The latter species is alw^ays found, so far as our
knowledge is concerned, epiphytic on Fucaceae, while the former
species is confined to rocks. Ectocarpus chantransioides has also more
slender filaments than has E. hemisphericns, not at all tapering, and
has distinct apical growth. The gametangia are differently shaped,
being longer and more slender. It is therefore very distinct even from
E. hcmiphcricus. It resembles the genus Choristocarpus of the family
Choristocarpaceae, as diagnosed by Kjellman (1897. pp. 190, 191^, in

1922] • Setchell-Gardner : Phycological Contributions 407

having apical growth. The growth in length of the erect filaments
in Choristocarpaceae is by the division of the terminal cell only,
whereas in E. cliantransioides the meristem extends over a number of
cells at the outer or apical end of the filaments. These cells, some
10 to 15 in number, are much richer in cell contents, the terminal cell
being the richest of all in the series. This is a very unusual condition
for an Ectocarpus. The nearly uniform diameter throughout of the
erect filaments and their method of branching resemble to a remarkable
degree those characters found in Chantransia. The gametangia are
typically those of Ectocarpus, and notwithstanding the other rather
unusual characters, it seems to be most closely related to that genus,
but a very distinct species.

Ectocarpus commensalis sp. nov.

Plate 48, figures 32-35

Frondibus 200-400/* altis, dense fasciculatis. per filamenta rhizoidea
intricata, parce ramosa inter utriculos et usque inter filamenta medul-
laria hospitis penetrantia affixis ; filamentis erectis prope basim alterne
ramosis, superne simplicibus et leviter attenuatis. non pilif eris ; cellulis
filamentorum erectorum cylindricis, non constrictis, basim 12-18/i,
diam.", 1.5-2.5-plo longioribus quam latis; chromatophoris taeniatis, in
cellula quaque paucis; zoosporangiis ignotis; gametangiis obtuse fusi-
formibus usque ad cylindrico-conicis, brevi-pedieellatis. prope basim
filamentorum plerumque lateralibus, raro terminalibus, 60-lOOju, longis,
15-20/x latis.

Growing on Codium Setchellii Gardner, Carmel Bay, and on
Codium fragile (Suringar) Hariot, Pacific Grove. Monterey County,
California. Type, Gardner, no. 3319 (Herb. Univ. Calif., no. 206994),
May.

This minute species is one of several occurring on species of Codium,
both on this coast and on that of southwestern Europe. The colorless
rhizoidal filaments descend into the substance of the host even to the
central (or basal) medullary tissues. The smaller diameter of the
erect filaments and the more narrowly conical or fusiform gametangia
readily distinguish this species from the others.

Ectocarpus eramosus sp. nov.

Plate 47, figures 18-23

Frondibus diffusis aut flocculosis. 1-3 mm. altis, per filamenta
rhizoidea irregulariter ramosa inter utriculos hospitis profunde pene-
trantes affixis; filamentis erectis eramosis aut raro ramos breves.

408 University of California Puhlications in Botany T^'ol. 7

divorf^'ciitcs vnlj,'o in ^MiiU'tan^nis tLTiiiinantcs t'lnitteutibus, basini et
apicc It'vitor attfimatis, noii piliferis; ccllulis filamentorum maturorum
28-40/1 (liaiii.. 0.5-1.25-plo longioribus quam latis, cylindricis, levissime
coiistrictis; chroiniitoplioris coinparate crassis. dense confertis. irretjii-
lariler laeiiiatis. sine pyrenoideis; zoosporangiis ignotis; gametangiis
iiiaxime niagnitudine foniiaqne variabilibus ant lateralibus et per
filainenta tota alternis ant interdnm in filamentis primariis terminal-
ilms et seriatis ])leniin(|ne pedicellis paneicellnlatis, lateralibus cylin-
drieis ns(|ue ad eylindrieo-eonicis, 150-280/a longis, 26-36/x latis, ter-
niinalilnis seriatis, 700-900/a longis, 15-20/a latis.

Growing on C odium fragile in the lower littoral belt. Near the
entrance to Tomales Bay, Marin Connty, California. Type, Gardner,
no. 3448& (Herb. Univ. Calif., no. 206995), August.

This very small species has the general appearance of a form of
Ectocarpus confervoides, but the cells are short and the filaments very
slightly constricted at the partitions. It seems, in these respects, nearer
to E. acutus and E. corticidatus, possibly bearing something of the
same relation to these species that the dwarf forms of E. confervoides
do to the typical form. Ectocarpus eramosus, however, is not readily
to be referred as a dwarf form of either E. acutus or E. corticulatus
and is consequently to be kept separate, at least for the present.

Ectocarpus flagelliferus sp. nov.

Plate 47, figures 2.5-27

Frondibns dense floccosis, 3-5 mm. altis, per filamenta pauca
repentia et rhizoidea e cellulis inferis descendentia, decolorata affixis;
filamentis ereetis inferne simplieibus, superne sparse et alterne ramosis,
superne longe attenuatis, inferne abrupte contractis, juvenis piliferis;
cellulis filamentorum primariorum leviter doliiformibus, 25-35/a diam.,
0.25-1.5-plo longioribus quam latis; chromatophoris parvis, irregu-
lariter taeniatis ant lamiiiaeformibus. pyrenoidis deniitibns: zoospor-
angiis ignotis; gametangiis magnitudine formaque variabilibus fusi-
formibus usque ad cylindrico-conicis, plerumque in pedicellis longis
lateralibus, interdum in filamentis curtis erectisque terminalibus, aut
in filamentis primariis longe-seriato-interealarilms, lateralibus 125-
200fjL longis, 28-40yx latis, terminalibus et intercalaribus usque ad
1.5 mm. longis.

Growing on eel-grass. Sitka, Alaska. Type, Gardner, no. 3970
(Herb. T^niv. Calif., no. 206999), July.

Thr species we have named Ectocarpus flagelliferus, while re-
sembling superficially some of the shortest plants of Ectocarpus
confervoides f. variahilis, presents several peculiarities whicli are
characteristic. The pbints of this species are attached, so far as the

1922] Setchell-Gardner : Phycological Contributions 409

adult specimens we have for study are concerned, by rhizoidal fila-
ments which originate from several of the lower cells of the erect
filaments and form complex basal masses which almost completely
obscure the creeping filaments. Neither the rhizoidal filaments nor
the creeping filaments, however, penetrate the host. The chromato-
phores are small and irregular in outline, seeming to be short bands
rather than regular disks. They are generally so closely placed in
the younger cells as to seem almost like a continuous band, but are
separate in the older cells with slender processes almost connecting
them to one another. The cells are short and the erect filaments and
their branches extend out into long hairs like whip-lashes. The lateral
gametangia are variable in shape, arranged much as in Ectocarpus
confer voides f. variabilis, but are of different dimensions. Besides the
characteristic Ectocarpus-iy\)e of gametangia which are lateral, there
occur terminal seriate gametangia of the Pylaiella-type which reach an
extreme length of 1.5 mm.

Ectocarpus flocculiformis sp. nov.

Plate 47, figure 24

Frondibus dense caespitosis, 0.75-1.5 mm. altis, per filamenta inter-
texta, penetrantia, rhizoidea affixis ; filamentis erectis basi plus minusve
furcatis, superne simplicibus aut in plantis, maximis, ramulis paucis,
Curtis, alternisque instructis, basi leviter attenuatis, superne longe
attenuatis, non piliferis; cellulis cylindricis ad leviter doliiformibus,
20-25/x diam., 1-2.5-plo longioribus quam latis ; chromatophoris numer-
osis, in cellulis junioribus angulato-discoideis, in vetustioribus rotun-
datis; zoosporangiis ellipsoideis, 56-66/x longis, 35-40jU, latis, prope
basim filamentorum erectorum in pedicellis curtis suffultis; game-
tangiis ovoideis ad ellipsoideis, numerosis, sessilibus aut brevi-pedicel-
latis, prope basim filamentorum erectorum lateralibus, 95-110/x (usque
ad IdOfi) longis, 30-45ja latis.

Growing on Codiuni fragile, the rhizoidal filaments penetrating
deeply into the host, among and beyond the utricles: La Jolla, Cali-
fornia. Type, Gardner, no. 3540 (Herb. Univ. Calif., no. 206996),
December.

The nearest relative of E. floccidiformis apparently is E. cylindricus
f. codiophilus. One marked difference between it and any of the forms
of E. cylindricus is in the shape and size of the gametangia. This
difference, along with others of less importance perhaps, though appar-
ently constant, seems to be sufficient to warrant keeping it separate.

410 University of California PuUications in Botany [Vol. 7

Ectocarpus fructuosus sp. nov.

IMate 45, figures 1-4

Fntmlihus iloccosis, i)roi"iise et altenie ramosis, usque ad 2.5 cm,
altis. jKT lilaincnta copiosa, repentia et comparate brevia affixis: fila-
nu'iitis orcctis ])asi repetite furcatis, ramos longos ramulis ordinuiii
dunniiii IriuiiKiuc l)r('vil)ns o1)tusis(|Uc ubi(|Uo eircumdantcs prndue-
ontibus; ccHuIis liliiiiR'ntorum primariurum orectorum cylindricis
usque ad leviter doliiformibus et parte ad dissepimenta constrictis,
inforne 18-25/Lt diam., 2-4-plo longioribus quam latis, superne leviter
latioribus brevioribusque ; zoosporaugiis iguotis: ofametangiis numer-
osis, hite couiois. sessilibus aut 1-3-cellulato pedieellatis, 50-7()/x longis,
basi 25-35/A latis.

Growing on the pneumatocyst of Nereocystis Luetkeana. Moss
Beach. San M-Atoo County, California. Type, Gardner, no. 4568 fHerb.
Univ. Calif., no. 207000), April.

Ectocarpus fructuosus comes within the E. confervoides group, but
seems amply distinct from any known forms to constitute a species.
This species is based upon the character of the branches, viz. ; numerous
long branches, producing throughout a great abundance of short,
rather blunt ramuli, and upon the very numerous, rather short and
blunt, predominatingly sessile gametangia. The species seems to be
rare.

Ectocarpus granulosoides sp. nov.

Plate 45, figures 7, 8

Frondibus 2-3 cm. altis. profuse ramosis ; filamentis primariis sub-
dichotome ramosis, ramis omnibus basi abrupte attenuatis, inferne
eorticulatis ; filamentis secundariis plerumque alternis, partim secun-
dis, strietis, longe attenuatis, acutis, non piliferis, ramulis ultimis
plerumque secimdis, aeutis; cellulis filamentorum primariorum 70-SOju.
diam., 0.5-1-plo longioribus quam latis, filametorum secundariorum
30-40/1 diam., 0.3-1-plo longioribus quam latis, ramulorum ultimorum
14-20/x diam., 0.3-1 .5-plo longioril)us (|u;im latis, omni])us ad dissey)i-
menta leviter constrictis; ehromatophoris numerosis. in cellulis vctusti-
oribus discoideis,in cellulis ramulorum irregulariter angulosis; cellulis
filamentorum cortieantum 7-lOju, diam.. 3-4-plo longioribus quam latis ;
zoosporangiis ignotis; gametangiis in latere superiore ramuhu'um
ultimorum pcnultimorumque seeundis, sessilibus, subfusiformibus,
asymmetricis, 40-60/i longis, 12-20/1 latis.

Growing on rocks(?). San Pedro, California. Type, Setchell,
no. lloGa, December, 1895.

The very distinct species we have described under the name of
Ectocarpus yranulosoides has the habit of a small E. gramdosus and

1922] Setchell-Gardner : Phycological Contributions 411

gametangia of the same general type as found in that species, but con-
siderably smaller. The branching, however, is never opposite, and the
acute ramuli gradually attenuated upward recall those of E. acutus,
from which our species is amply distinct in both chromatophore and
characters of its gametangia. The branches, particularly the larger,
are very suddenly and considerably attenuated at the base, giving the
species a striking characteristic of its own, at least within the group
of species with discoid chromatophores.

Ectocarpus Mesogloiae sp. nov.

Plate 45, figures 5, 6

Frondibus minutis, 0.75-1.5 mm. altis, per filamenta copiosa, dense
intertexta ramosa, rhizoidea, inter callulas hospitis penetrantia affixis ;
filamentis erectis sparse alterneque ramosis ; filamentis primariis et
ramulis superne longe attenuatis, acutissimis, non piliferis; cellulis
cylindricis, ad dissepimenta leviter constrictis, basi 15-18/a diam.,
1-2-plo longioribus quam latis, apicibus filamentorum 4- 6/a diam. ;
chromatophoris tenuibus, irregulariter taeniatis, parietes cellularum
f ere tegentibus ; zoosporangiis ignotis; gametangiis anguste cylindrico-
conicis, 120-160|U, (usque ad 210./x) longis, 18-22/x latis, brevi-pedicel-
latis, raro sessilibus.

Growing on Mesogloia Andersonii. Carmel Bay, Monterey County,
California. Type, Gardner, no. 2865a (Herb. Univ. Calif., no. 206989),
May, 1915.

The general characters of this diminutive species ally it with the
E. confervoides group. "We have deemed it best to consider it a distinct
species on account of the small dimensions of all of its parts and its
penetrating habit. It has not been seen on any other host except the
one mentioned above.

Ectocarpus Saundersii sp. nov.

Frondibus floccos parvos 2-5 mm. altos formantibus, per filamenta
copiosa, ramosa, in superficiem hospitis repentia affixis : ramis alternis ;
ramis primariis longis divergentibusque ramulis ultimis sparsis.
brevibus, acuminatis ; cellulis filamentorum primariorum 25-40/i, diam.,
inferne quadratis, superne brevioribus, ad dissepimenta leviter con-
strictis; chromatophoris numerosis, discoideis; zoosporangiis breviter
pedicellatis aut intercalaribus, globosis, ad 30/x diam. ; gametangiis
cylindricis usque ad ovoideis. obtusis aut acuminatis, lateralibus.
pedicellis longioribus aut brevioribus suffultis, 70-150/^ longis, 25-50/x
latis.

41"J rnivcrsH;/ of California Publications in Botany [Vol.7

Growiiiti: on Fu(i(s. in llic middle of the littoral belt. Pacific Grove,
California.

Erfocarpns ))ara(lo.rus var. pacificus Saunders, Pliyc. ^lein.. 1898,
p. 152, 1)1. 18, fifjfs. 4-7; Collins, Ilolden, and Setchell, Phyc. Bor.-Amer.
(Exsiec), no. 530.

Aofordinj? to Saunder.s, hi.s plant, the vai'. pacificus, differs from
the type of E. paradoxus IMont., in lacking any manifestation of
opposite branching as well as in ha^dng the gametangia longer and
more pointed. In these respects and also in that it is a shorter plant,
it seem.s to us to be siifificiently distinct to be separated specifically
from tlie type, and to be difTerent from any other species known to us.
We are consequently dedicating it to its discoverer.

Ectocarpus simulans sp. nov.
Plate 45, figures 9-11

Frondibus 1-2 nun. altis. floccosis; filamentis prostratis tortuosis,
discretis ; filamentis erectis simplieilms. superne leviter attenuatis, non
piliferis; cellulis cylindrieis. ad dissepimenta non constrictis, medio
11-13/A diam., 1-2.5-plo longioribus quam latis: chromatophoris taeni-
atis. parietes cellularum prope exacte tegentibns, pancis; zoosporangiis
ignotis; gametangiis lateralibus aut interdum terminalibns. sessilibus.
anguste ellipsoideis. obtusis, plerumque leviter curvatis, 55-65/x longis,
15-20/A latis.

Growing on Chaetomorpha aerea. Cypress Point. Monterey County,
California. Type, Gardner, no. 3087 (Herb. Univ. Calif., no. 206993).
December.

The present species resembles, in general, certain forms of Ecto-
carpus pusillus Griffiths (cf. Sauvagean, 1895), but the erect filaments
are much more slender, usually never branched, and with much smaller
gametangia. It forms tufts on Chaetomorpha aerea, whose filaments
are encircled by the prostrate filaments of the Ectocarpus which form
small cushions in their growth over one another. The gametangia are
rather blunt and often more or less curved.

Ectocarpus socialis sp. nov.
Plate 46, figures 12, 13

Frondibus eaespitosis, 1-3 mm. altis, per filamenta rhizoidea dense
intrieata et penetrantia affixis; filamentis erectis, ad superficiem
ho.spitis. repetite fureatis. superne simplieihns aut alternate-ramnsis,
longe attenuates et piliferis; cellulis cylindrieis uscpie ad leviter dolii-

1922] Setchell-Gardner : Phycological Contributions 413

formibus, non constrictis, 22-28/x cliam., inferne 1-2-plo, superne
1-2-plo longioribus qiiam latis; chromatophoris parvis, in cellula
quaque paucis, cliscoideis ; zoosporaiigiis ellipsoideis, erectis, lateralibus,
alternis 1-2-cellulato-pedicellatis, interdum prope basim filamentorura
erectoriim sessilibus, 60-95/x longis, 35-50/^, latis; gametangiis fusi-
formibiis, sparsis, partes superiores filamentornm erectoriim occu-
pantibus, sessilibus aut 1-2-cellulato-pedicellatis, alternis 70-85/x,
longis, 25-30^ latis.

Growing on Codium fragile (Suringar) Hariot, in companv M-ith
Ectocarpus glohifer Kuetz. and with various species of Myxophyceae
and Rhodophyceae. Redondo, California. Type, Gardner, no. 1947&
(Herb. Univ. Calif., no. 206987), March.

Ectocarpus socialis seems to be a relative of the E. cylindricus
group. The chief distinctions are in the shape, size, and position of
the gametangia.

Ectocarpus Taeniae sp. nov.

Plate 46, figure 15

Frondibus diffusis, 0.5-1.5 mm. altis, per filamenta superfieialia
repentia. profuse ramosa affixis; filamentis erectis simplicibus, basim
le^nter attenuatis. superne longe attenuatis. piliferis; cellulis. fila-
mentorum repentium cylindricis, 4-6jU, diam., filamentorum erectorum
8-lOju, diam.. inferne quadratis, apicibus 4-6-plo longioribus quam
latis; chromatophoris eurtis, comparate crassis. irregulariter taeniatis;
zoosporangiis ignotis ; gametangiis anguste ellipsoideis. interdum lente
curvatis, in filamentis repentibus et erectis plerumque sessilibus, 20-28ju
(usque ad -iOix) longis, 15-20/* latis.

Growing on Taonia Lennebackerae. San Pedro, California. Type,
Gardner, no. 3561a (Herb. Univ. Calif., no. 206997), December.

This species forms rather diffuse and even layers, mostly along the
margins of Taonia, over w^hose surface the creeping filaments spread
and give rise to gametangia or to erect filaments not over 1.5 mm.
high. In many ways it seems like a miniature Ectocarpus Padinae
(Buffh.) Sauvageau (1897, p. 30 et seq.). Our species, however, is
only about half as tall, the cells of the erect filaments about half as
broad, and the gametangia less than half as large as those of E.
Padinae. We have found gametangia only of the "antheridium" type.
Ectocarpus Padinae has its creeping filaments entirely endophytic
while E. Taoniae has them entirely epiphytic.

41'4 University of California Publications in Botany [Vol. 7

Ectocarpus confervoides f. parvus (Saunders) comb. nov.

Fronds <j:rc^Mri()ii.s. r<»i-niin<^ tufted or extended masses. 1-2 em.
liiixh. more or le.ss alternately branched; erect filaments 21-30/x broad,
.slifxhtly constricted at the joints, much attenuated at the tips or end-
injr in hairs; cells 1-3 times as long as broad below, shorter above;
chroiiiatophoi-es irrefruhii'ly band-shaped; zoosporanfiia ovoid or ellip-
.soid, .35-5;V long, 20-27/x broad, usually sessile, sometimes on the same
filament with the gametangia; gametangia narrowly lanceolate-conical,
gradually long-attenuated above, 120-400ju, long, 20-27/a broad, borne
on a longer or shorter pedicel, occasionally seriate and intercalary
(a,s in rylaidla).

On sand-covered rocks (type!), logs, or on the larger Melano-
phyceae. Central California (near San Francisco) and southern
California (San Pedro, the type locality, and San Diego).

Ectocarpus siliculosus parvus Saunders, Phyc. Mem., 1898. p. 153,
pi. 22. figs. 1-9.

The f. parvus, as defined above, presents certain resemblances to
both Ectocarpus siliculosus and E. confervoides. Its slender elongated
gametangia resemble those of E. siliculosus and much might be said
in favor of the opinion of Saunders in making it a "variety" of that
species. On the other hand, very few, or none, of the typical game-
tangia end in a hair and we have therefore preferred to place it rather
with E. confervoides. There are often found in this form, as well as
in forms of E. siliculosis, very curious, elongated, intercalary game-
tangia which seem to be seriate in that \\\ey open laterally in sections,
as do those of Pylaiclla. Such gametangia are very puzzling, but may
possibly be regarded as indicating hybridization between Pylaiella and
Ectocarpus or a mutation of some sort. Speculation, however, can
scarcely explain such anomalies. The majority of the gametangia are
typically of Ectocarpus and while ajiproaching in shape those of E.
silicidosus, are less slender and less elongated than is typical for that
species and are not ordinarily prolonged into terminal hairs.

So far as we may judge from the scanty specimen in our copy. no.
358 of Tilden's American Algae (from Pacific Grove) belongs rather
under f. parvus than f. variahilis, although the determination as f.
variahili.s is attributed to Saunders.

1922] Setchell-Gardyier : Phycological Contributions 415

Ectocarpus cylindricus f. tj^icus nom. nov.

Fronds diffused, 1-2 mm. high ; creeping filaments superficial ; erect
filaments very sparingly branched above, several arising from the same
creeping filament, tapering slightly at the base and apex ; cells of the
creeping filaments 16-20/i, diam. ; cells of erect filaments cylindrical,
slightly constricted at the cross-walls, 2-3 times as long as broad below
and at the apex, 0.5-1 times in the middle of the filament ; gametangia
mostly on the middle and upper parts of the filaments, mostly alternate.

Growing on Egregia Menziesii, Halidrys dioica and C]ifitose'ra
osmundacea. The type locality is Pacific Grove, California.

Ectocarpus cylindricus f. codiophilus f. nov.

Plate 46, figure 14, and Plate 49, figures 42-4.5

Frondibus dense caespitosis, 3-5 mm. altis, per filamenta rhizoidea
dense intricata. hospitem penetrantibus ; filamentis erectis prope super-
ficiem anastomosantibus et furcatis, superne eramosis, piliferis, game-
tangiis plerumque prope basim filamentorum erectorum sitis.

Growing on C odium fragile (Suringar) Hariot and C odium Set-
cliellii Gardner. Carmel Bay, Monterey County, California, and La
Jolla. San Diego County, California.* Type, Gardner, no. 3510a (Herb.
Univ. Calif., no. 206992), December.

This form differs from f. typicus in having penetrating rhizoidal
filaments and in that the gametangia are gathered in a dense zone at
or near the base of the erect filaments instead of being scattered.

Ectocarpus cylindricus f. acmaeophilus f. nov.

Plate 49, figure 46

Frondibus floccosis, 7-10 mm. altis; filamentis repentibus super-
ficialibus, filamentis erectis eramosis, diametro in parte quaque
similibus ; zoosporangiis ignotis ; gametangiis usque ad 270/x longis,
plerumque in parte superiore filamentorum erectorum oppositis.

Growing on Acmaea sp. Carmel Bay, Monterey County, California.
Type, Gardner, no. 2881 (Herb. Univ. Calif., no. 206990), May.

This form seems to be rare, at least but few specimens have been
detected up to the present time. It differs from f. typicus in being
practically unbranched, except the fructiferous branches. Where
branches occur, they seem to arise as the result of an injury to the
main filament. It differs further in that the erect filaments are more
nearly cylindrical throughout, and the gametangia more often opposite
and generally longer and proportionally narrower.

416 Universiiy of Calif ornia Publications in Botany [^'ol. 7

Ectocarpus siliciilosus I", subulatus (Kuetz.) comb. nov.

Fronds 5-25 cm. \ug\\. litilit yellow, fleecy, ]iiuch bi-andicd, not
const ricfcd at the joints; l)ranc]ies long, attennated above, many end-
inir in a loiiir hair; ('rjls of the main filaments 30-36/x, broad. 1-1.25
times as lon^' as i)road. zoos|)oi-aii^ia nnknovvn ; gametan^na elongated
snbnlate-ovoid, some stouter, some more slender, 200-GU()/^ long. 12-48|u,
l)n)a(i. the upper (and as a rule more slender) usually terminating in
a hail', on a 2-10 or 12 celled pedicel.

In brackish pools, on sticks or grasses. Central California (San
Francisco Bay).

Ectocarpus confervoidcs f. subulahis Collins, llolden, and Setehell.
Pliyc. Bor.-Amer. (Exsicc), no. 1231. Ectocarpus suhulatus Kuetzing,
Spec. Alg., 1849, p. 454, Tab. Pbyc, vol. 5, 1885. p. 19, pi. 61, fig. II.
Ectocarpus confervoides Hauck, Meeresalg., 1884, p. 331 (excl.
synonomy). Ectocarpus amphihius Harvey, Phyc, Brit., vol. 2, 1848,
pl. 183, Ner. Bor.-Amer.. part III, 1858, p. 125.

Our plant seems to agree very well Avith the figures and description
of Ectocarpus amphihius Harvey and also, although not so perfectly,
with E. suhulatus Kuetz. They are both brackish water forms, as are
our plants. Hauck refers to his var. suhulatus also Ectocarpus drapar-
naldiaeformis Kuetz. and E. macrocera^ Kuetz. Judging from Kuetz-
ing's illustrations, these two species may be forms of E. siliculosus
but are not to be included under f, suhulatus.

19--] Setchell-Gardner : Phycological Contributions 417

LITERATURE CITED

Collins, F. S.

1913. The marine algae of Vancouver Island. Canada Geological Survey,
Victoria Memorial Museum. Bulletin no. 1, pp. 9'9-137. Victoria,
B.C.
Collins, F. S., Holden. I., and Setchell, W. A.

Phycotheca Boreali- Americana (Exsicc), fascicles 11, 21, 23, 25.
Maiden, Mass.

Harvey, W. H.

18.58. Nereis Boreali-Amerieana. Part 3, Chlorospermeae.

1848. Phycologia Britannica, vol. 2, pi. 183. London.

Hauck, F.

1884. Die Meeresalgen Deutschlands und Oesterreichs, in Rabenhorst's
Kryptogamen-Flora von Deutschland, Oesterreich und der Schweitz,
vol. 2, lief. 8.

KUETZING, F. T.

1849. Species Algarum. Leipzig.
1855. Tabulae Phycologicae, vol. 5.

Kjellman, F. R.

1877. Om Spetsbergens marina Klorofyllforande Thallophyter II. Bihang
till K. Sv. Vet.-Akad. Handl., vol. 4, no. 6. Stockholm.

1897. Choristocarpaceae, in Engler and Prantl, Natiirl. Pflanzenfam., 1 Th.,

2 Abt. Leipzig.

Saunders, De A.

1898. Phycological Memoirs. Proc. Calif. Acad. Sci., (3), Bot., vol. 1, pp.

147-168, pis. 12-32. San Francisco.

Sauvageau, C.

1895. Note sur I'Ecfocarinis pusiUus Griffiths. Jour, de Bot., vol. 9.

1896. Remarques sur la reproduction des Pheosporees et en particulier des

Ectocarpus. Ann. des Sci. Nat., 8 ser., Bot., vol. 2, pp. 223-274.
1896a. Sur la nature des sporanges en chapelet de 1' "Ectocarpus confer-

voides." Jour, de Bot., vol. 10.
1896b. Sur 1' "Ectocarpus virescens" Thuret, et ses deux sortes de sporanges

pluriloculaires. Jour, de Bot., vol. 10.
1896-7r. Observations relatives a la sexualite des Pheosporees. Jour, de Bot.,

vols. 10, 11.
Setchell, W. A., and Gardner, N. L.

1903. Algae of Northwestern America. Univ. Calif. Publ. Bot., vol. 1, pp.

165-418, pis. 17-27.

PLATE 45
Ectocarpus fructuosus S. and G.

Fig. 1. Diagrammatic illustration of whole plant.

Fig. 2. Part of a plant showing positions and various shapes and sizes of
the gametangia. X 175.

Figs. 3, 4. Terminal ramuli showing variations in shape of cells and some-
what reduced gametangia. X 60.

Ectocarpus Mesogloiae S. and G.

Fig. 5. A single gametangium and a piece of terminal ramulus illustrating
chromatophores and cells. X 200.

Fig. 6. A group of plants showing rhizoids, character of branching, relative
shapes and sizes of gametangia. X 40.

Ectocarpus gramdosoides S. and G.

Fig. 7. A group of terminal ramuli showing character of branching, shapes
of cells, positions and shapes of gametangia. X 125.

Fig. 8. A few cells of the main filament showing the shapes of cells and
of chromatophores. X 125.

Ectocarpus simulans S. and G.

Fig. 9. Diagrammatic illustration of groups of plants on their host.

Fig. 10. Parts of plants showing shapes and positions of gametangia.
X 125.

Fig. 11. A group of cells of the main filament showing chromatophores.
X 250.

[418]

UNIV. CALIF, PUBL. BOT. VOL, 7

[SETCt-IELL-GARDNER ] PLATE 45

PLATE 46
Ectocarpus socialis S. and G.

Fig. ]2. A small group of plants showing rhizoids, a few cells with chro-
matophores, and the positions and shapes of the zoosporangia. X 125.

Fig. 13. A selection of parts of filaments showing the positions and relative
shapes and sizes of the gametangia. X 125.

Ectocarpus cylindricus f. codiophilus S. and G.

Fig. 14. A group of young plants showing both zoosporangia and game-
tangia, the two parts of ramuli on the left representing these organs on the
same plant. X 60.

Ectocarpus Taoniae S. and G.

Fig. 15. A portion of a creeping filament showing erect filaments and the
positions and relative shapes and sizes of the gametangia on both the creeping
and the erect filaments. X 250.

Ectocarpus affinis S. and G.
Fig. 16. Diagrammatic representation of a small group of plants on their host.
Fig. 17. Portions of filaments illustrating positions, relative shapes and
sizes of gametangia. X 225.

[420]

UNIV. CALIF. PUBL. BOT. VOL, 7

[SETCHELL-GARDNER ] PLATE 46

^\ >^U

Fig.

18,

and sizes.

Fig.

19,

Fig.

20.

PLATE 47

Ectocarpus eramosus S. and G.

A group of typical gametangia showing the range of relative shapes
X 60.
Part of a filament showing an intercalary gametangium. X 60.
A part of a filament near its outer end, showing variation in shapes
and sizes of the cells and scattered dwarf gametangia. X 60.

Fig. 21. The basal parts of a group of erect filaments showing long rhizoidal
filaments. X 60.

Fig. 22. A filament showing two gametangia with long pedicels. X 60.
Fig. 23. Terminal parts of filaments, one showing chromatophorcs. X 60.

Ectocarpus flocculiformis S. and G.

Fig. 24. A group of filaments anastomosing at the base and developing
numerous rhizoids and characteristic gametangia. X 125.

Ectocarpus flagelUferus S. and G.
Fig. 25. Showing an intercalary gametangium. X 125.
Fig. 26. Showing normal gametangia and chromatophorcs. X 125.
Fig. 27. Showing basal parts of filaments. X 125.

[422]

UNIV. CALIF. PUBL. BOT. VOL. 7

[SETCHELL-GARDNER ] PLATE 47

PLATE 48
Ectocarpus chantransioides S. and G.

Figs. 28, 29. Characteristic terminal ramuli with characteristic gametangia.
X loO.

Fig. 30. Diagrams showing the pulvinate habit of growth. One in the upper
right-hand corner rejiresents a vertical section view.

Fig. 31. Illustrating the creeping filaments. X 150.

Ectocarpus commensalis S. and G.

Fig. 32. A grouj) of plants with relatively long gametangia. X 50.

Fig. 33. A single typical gametangium and parts of filaments showing
chromatophores. X 225.

Fig. 34. A group of young filaments with long rhizoids.

Fig. 35. A group of plants showing their relation to the host. Diagrammatic.

Ectocarpus acutus S. and G.

Fig. 36. A diagrammatic illustration showing method of branching and tlie
shape and position of the gametangia.

Fig. 37. A group of gametangia showing some variations in form and size.
X 125.

Fig. 38. A piece of a filament showing shapes of cells and chromatophores.
X 125.

Fig. 39. A few terminal acute ramuli. X 125.

[424]

UNIV, CALIF. PUBL. BOT. VOL. 7

[SETCHELL-GARDNER ] PLATE 48

PLATE 49

Ectocarpus acutus S. and G.

rig. 40. A group of terminal ramuli bearing numerous complex zoospo-
rangia(?). X 125.

Fig. 41. A group of typical gametangia. X 125.

Ectocarpus cylindricus f. codiophilus S. and G.

Fig. 42. Piece of a filament bearing zoosporangia. X 125.
Fig. 4,3. Piece of a filament bearing opposite typical gametangia. X 125.
Fig. 44. Piece of a terminal filament bearing zoosporangia. X 125.
Fig. 45. A group of plants showing the typical crowded basal gametangia.
X 65.

Ectocarpus cylindricus f. acmaeopliilus S. and G.

Fig. 46. Piece of a typical filament bearing opposite sessile gametangia.

X DO.

[426]

UNIV. CALIF. PUBL. BOX. VOL. 7 [SETCHELL-GARDNER ] PLATE 49

UNIVERSITY OF CALIFORNIA PUBLICATIONS

IN

BOTANY

Vol. 7, No. 12, pp. 427-436, plate 50 June 16, 1922

LWmAWf

NOTES ON PACIFIC COAST ALGAE
II. ON THE CALIFORNIAN "DELESSERIA QUERCIFOLIA"

BY

GAEL SKOTTSBERG

During my visit to California in 1913 Professor W. A. Setchell
introduced me to a red alga called Delesseria (or Schizmieura) querci-
folia by American algologists and identified by them with a plant
described by Bory in Voyage sur "La Coquille, " Cryptogamie, p. 136,
table 18, fig. 1, from the shores of subantarctic America. However,
it appeared to me that it was different from this latter, which I had
collected at various occasions on the coasts of Tierra del Fuego and
the Falkland Islands, where it is one of the commoner species. When
preparing the paper on my collection of 1902 I compared my material
of Bory's species with a specimen from California in Herb. Stockholm,
and stated that the latter belongs to a different yet undescribed species
(see Kylin and Skottsberg, "Zur Kenntnis der subantarktischen und
antarktischen Meeresalgen II," p. 94, Wiss. Ergebn. der schwed. Siid-
polar-Exp., IV:15, 1919).

In order to make a more careful comparison between the two species,
I asked Professor Setchell for some material from California, and he
most willingly sent me a good set of specimens with all kinds of repro-
ductive bodies, for which I give him my best thanks. As all the
material is dry, it has not been possible to enter into histological details
regarding the formation of spermatia, cystocarps, or tetraspores, but
this does not, I dare say, prevent us from arriving at a safe conclusion
as to the systematic position.

Both species agree in most characters and are nearly related
to each other, but well separated from other species. In both the
frond consists of an oblong, short, stipitate lamina of ordinary size,
with more or less sinuate margins, monostromatic except for the costa
and nerves; as usual, the parts where tetraspores, etc., are formed

428 University of California Puhlications in Botany [Vol.7

become polystromatic. The eosta is very conspicuous and of the same
anatomical structure in both, it emits opposite, nearly straight to irreg-
ularly bent, distinct nerves. Tlie ramification from the lamina is
marginal, solitary segments growing out to form new blades. When
the lamina has worn away numerous shortly stipitate proliferations
grow out from the edges of the old, flattened costa. The hapter is a
small disc.

The cystoearps are scattered between the nerves, and the same
position is occupied by the spermatangia and tetrasporic sori, the
latter often occupying the entire space between the costa and the
margin of the frond, only leaving the nerves free. Further, both
species have exactly the same mode of apical growth (see below).

The Calif ornian plant differs from the true D. quercifoUa in the
following characters : The frond is more oblong, Ungulate, and, except
for the segments growing out to form branches, entire, or only slightly
undulate and denticulate. The frond of D. quercifoUa Bory is more
broadly obovate in outline, more or less deeply sinuate-dentate, young
intact specimens recalling the leaves of the common European Quercus
rohur and very similar to D. sinioosa of the northern coasts. The
anatomical structure is the same in all these species. To judge from
the material examined there is some little difference in the costa
between the Californian species and D. quercifoUa. In the former
(fig. 1) the central lamella consists of one layer of very large cells,
corresponding to the monostromatic lamina; and even the innermost
cortical cells are much smaller, each cell, on the cross-section, sup-
porting one or two rows of radially arranged cells. In D. quercifoUa
there generally are as much as five layers of large cells, because the
inner cortical strata are more similar to the central lamella and con-
trasting with the outer radial rows of small cells, but in other cases
this structure has proved to be submitted to some variation, so we
should not lay very great weight upon this difference.

Anyhow we have to do with two distinct species belonging to the
same genus. D. quercifoUa Bory is sometimes called Schizoneura
quercifoUa (Bory) J. Ag. and, if we follow Agardh, it lies near at
hand to describe the other as a new species of Schizoneura. But if
we advance deeper into this matter we shall find that Agardh 's genus
is a mixtum compositum, and that we cannot arrive at a solution unless
entering upon the history of the genus Delesseria and the systematic
value of some of the genera created by Agardh in Sp. Alg. Ill :3
(1898).

1922] Skottsberg: Notes on Pacific Coast Algae 429

Ruprecht gave a summary of the history of nomenclature in
"Tange des Ochotsch. Meeres, " p. 250. When Lamouroux established
the genus Delesseria there was an older name for it, Hydrolapatha
(-urn) Stackhouse, pp. 54, 67, Tout, marino-crypt. II (1809). Stack-
house listed six species, among them H. sanguinea and H. sinuosa.
But the name Delesseria continued to be generally used, and this
caused the Brussels Congress, 1910, to put it on the list of "genera
conservanda" and to reject Hydrolapatha in spite of being older. I
think we had better agree to this proposal. D. sanguinea, mentioned
first under Delesseria by Lamouroux, should be regarded as the type
of the genus.

In Phycol. Gener. (1843) Kiitzing had established two new genera,
Phycodrys, based on D. sinuosa, and Hypoglossum, with H. Wood-
wardii (D. hypoglossum) as type. In Sp. Alg. (1849) the same sub-
division is retained. Here we need not occupy ourselves further with
Kiitzing 's system. Schmitz (Rhodophyceae in Engler and Prantl's
Natiirl. Pflanzenfam.) rejected both Phycodrys and Hypoglossum, but
at the same time expressed the view that the genus Delesseria ought
to become split up, though the time when this might be done in a
proper manner had not yet arrived.

In Sp. Alg. 111:1 (Epicrisis systematis floridearum, 1876) J. G.
Agardh divided Delesseria into thirteen sections: Schizoneiira was
one of these, characterized by the lamina being "subvage laciniato-
partita" and by the tetraspores situated on the main frond between
the nerves. Sect. Phycodrys had "frondes sinuato-pinnatifidae " and
sori along the costa or in the tips of the segments, or in separate leaf-
lets. D. sanguinea, the typical species of the genus, was excluded from
the order and brought to the Rhodymeniaceae as a monotypic genus
Hydrolapathum. During the whole of his life Agardh firmly adhered
to this peculiar idea. In the monograph on the Delesseriaceae, Sp.
Alg. 111:3 (1898), the sections of 1876 were given generic rank. The
genus Schizoneu^ra is now characterized mainly by the formation of
tetrasporangia : the sori are called "gemini" and "oppositi," while
they are called "singuli" in other genera, where two sporangia, accord-
ing to Agardh, were not formed opposite to each other on the two pages
of the frond. This was the case, f . i., in Delesseria in Agardh 's sense :
this name is applied to his former section Phycodrys. I cannot see
this point of difference, as sections through the sporophylls of D.
sinuosa show a bilateral development of the sori just as in Schizoneiira
quercifolia. Further, the anatomical structure and mode of growth

430 Umvcrsitij of California Puhlications in Botany [Vol.7

are identical in both. But Schizoneura comprises several species. S.
stihcostata, J. Ag., is mentioned first and should, perhaps, be regarded
as tlie type : in any case, quercifolia was declared by Agardh to be a
non-typical memlx'i- of the genus. I have examined part of the original
material of -S'. snhcostata, coll. by Schousboe. Here the costa does not
reach to the top of the frond, the top-cell soon loses its leading position
and can hardly be traced in- the older lamina, stray marginal cells
become centers of action, soon replaced by others ; we find conditions
characteristic of Nitophyllum, and the anatomical structure, with little
or no difference between the central lamella and the cortex, is quite
nitophylloid. This may also be seen from Agardh 's illustrations on
plate 26 in " Florid eernes morfologi" (K. Sv. Vetensk. Akad. Handl.
15:6, 1879) ; the figure of a growing apex is rather idealized. Of 8.
Davisii (Hook. fil. et Harv.) J. Ag., I have examined Hooker's type
(Herb. Kew). Tliis is still more like a Nitophyllum, as even in very
small proliferations no active top-cell is present, the growth being
marginal and intercalary. But in S. quercifolia the costa can be fol-
lowed right through to a typical, active top-cell, just as in D. sinuosa.
The nerves in *S'. suhcostata, Davisii, etc., are not opposite, but gener-
ally very irregular. There are hardly any nerves exactly correspond-
ing to the regular opposite ones in S. quercifolia, which must be
excluded from Agardh 's genus.

In the "Nachtrag" to Schmitz' Rhodophyceae, by N. Svedelius
(1911), some of Agardh 's new genera were recognized and others,
among them Schizoneura, rejected. So if we follow Schmitz and
Svedelius, Delesseria comprises sangiiinea, simcosa, quercifolia,
hypoylossum, etc. Of course neither of those authors was satisfied
with this arrangement; only, as Svedelius expresses himself, as long
as our knowledge of the various types is so insufficient as it is now,
it seems better to regard some of Agardh 's genera as divisions of
Delesseria. Still, I think that Nienburg's studies, also quoted by
Svedelius, permit us to reestablish Kiitzing's genus Phycodrys, with
Ph. sinuosa as type.

Tlie mode of growth in the apex of the Delesseriaceae has been
studied b}' various authors, including Nageli, Wille, Oltmanns.
Lately Nienburg made a critical examination of numerous
species and published his results under the title ' ' Zur Keimungs-und
Waehstimisgeschichte der Delesseriaeceen" (Bot. Zeitung, 66, 1908).
Of species of Delesseria (taken in its widest sense) he examined san-
guinea, sinuosa, Lyallii, alata, and hypoglossum. He distinguishes

1922] Skottsherg: Notes on Pacific Coast Algae 431

two types of apical growth, the hypoglossum-tyipe and the sinuosa-
type. The difference between them, as described and illustrated by
Nienburg, may be summarized as follows. Both have a central axis
with a primary top-cell and regular opposite branches, each ending in
a secondary top-cell. In the liypoglossum-type these branches of 1.
order carry, on their external side, branches of 2. order, each ending
in a tertiary top-cell (fig. 2). In the sinuosa-tyipe such top-cells do
not become developed : the cells of the branches of 1. order are divided
by length and cross-walls, but we do not find the regular arrangement
of branches second order as in the other type. Thus the appearance
of the growing apex becomes quite different in the two eases. To the
Jiypoglossum-type also belong D. sanguinea and alata, to the sinuosa-
type, D. Lyallii.

It was easy to see that D. quercifolia Bory as well as the Californian
species belong to the sinuosa-type. But when it came to the analysis
of the mode of development I could not bring my results in accordance
with Nienburg 's scheme. Anybody comparing my figure 3 with his
figures 16 and 44 will discover the difference. And as the plasma
connections designed by the writer were observed on alcohol material,
while Nienburg 's analytical scheme is a construction, I concluded that
the latter was not correct, or that, after all, D. quercifolia differed
from the sinuosa-type. I had a discussion on this matter with Pro-
fessor H. Kylin, who has occupied himself with an examination of
D. sinuosa. He kindly informed me that Nienburg 's explanation of
the growth is partly incorrect and that, in fact, the development is the
same in sinuosa and quercifolia. Nienburg 's types will stand, their
systematic importance will prove to be very great, but the main point
of difference seems to have escaped his full attention. In the hypo-
glossum-type, the cells of the main axis keep pace with the develop-
ment of the branches of 1. and 2. orders by getting elongated, while,
in the simiosa-type, they become divided by intercalary cross-walls
(a well-known fact), and the intercalary cells thus formed develop
short, few-celled branches which become inserted between the branches
of 1. order. This system of intercalary branchlets is dotted in figure
3. The same mode of growth is repeated in the branches of 1. order.
Thus it is easy to understand why the beautiful cellular architecture
of a Z>. hypoglossum is lost in D. sinuosa or quercifolia. In the Cali-
fornian species, owing to the dry material, no plasma connections were
distinctly visible, but a comparison between figures 3 and 4 will show
that the development is the same in both cases.

432 Umversity of CaUfornin I'lihlicatioiis in Botany [Vol.7

I have exaniiiK'd a great iiuinber oi' species belonging to the present
tribe Delesserieae as to the mode of growth, anatomical structure,
position of cystocarps and sori, and I believe that I can refer every
species to one of the two principal types. There are of course many
I have not seen or of which the material was less satisfactory. In all
the species of simtosa-growth. the cells of the main axis, as told above,
get divided by cross-walls, so that the central lamella of the costa is
formed by comparatively short cells. A cortex is formed by tangential
and radial divisions, the cells being arranged in a regular manner
(fig. 1) recalling the cortex in Nitophylluni, though tlie difference
between a "medulla" and a "cortex" is more evident than in the
latter genus. \n Hie hypolossum-tyipe, the costal cell-rows are formed
by very long cells: these cut off cortical cells, that divide by radial
walls, the cells thus formed get stretched in longitudinal direction,
are again divided by a tangential wall, and so forth, so that we observe,
on a length section, very long tubes in the center and gradually shorter
cells toward the surface. Even in a cross-section of a young costa
the radial arrangement is much less distinct than in the other type,
and later the difference becomes more marked. The long tubes make
the costa a rather weak structure, which is strengthened by the devel-
opment of numerous narrow hyphae (fig. 5). I have not found such
hyphae in any species belonging to the sinuosa-type. Those different
kinds of anatomical structure have been described and figured before
(see Wille, "Beitrage zur Entwickelungsgesehichte der physiologischen
GeW'Cbesysteme bei einigen Florideen" — Nova Acta der K. Leop.-
Carol. Akad. LII [1887]), but they were not, as far as I am aware,
combined with the different modes of apical growth. It should be
mentioned that, among species belonging to the h ypoglossum-type, I
have found two with non-typical anatomy, D. Montagneana J. Ag.
and B. ricscifolia Ag., both referred to the genus Apoglossum by J. G.
Agardh. In these small species hardly any pyphae at all get developed,
the differences between "medulla" and "cortex" are marked, but the
radial arrangement of cells less distinct than in sinuosa, etc. I may
add that the larger species, A. decipiens J. Ag., has typical hypoglos-
sum-strncture in the costa. In spite of these apparent exceptions, the
anatomical distinction between the two types is obvious.

If Delesseria sanguinea be recognized as the type of the genus, D.
sinuosa must of course be excluded and Phycodrys of Kiitzing be
restored. We have seen that Scliizoneura quercifolia differs from other
members of this dubious genus while it agrees with Phycodrys sinuosa

1^22] Skottsherg: Notes on Pacific Coast Algae 433

(Good, et Wood.) Kiitz. in essential characters. The principal dif-
ference lies in the position of the tetraspores. In the latter these
generally are developed in minute marginal leaflets forming a fringe
to the old frond, but they sometimes occur along the margins of the
main frond (see Kiitzing, Sp. Alg., p. 874). I do not think that we
can attribute any great importance to this difference, which, in this
case, seems to be of a more biological nature. The cj^stocarps have
the same position in both. Schizoneura quercifolia will receive the
name Phycodrys quercifolia (Bory) Skottsb., and the Californian
species is distinguished as

Phycodrys Setchellii, nov. spec.

Syn. Delesseria quercifolia auctt. quoad plantam califoruicam, non
Bory.

Frons primo simplex, oblonga, lingulata, brevissime stipitata, inte-
gra sed margine plus minusve sinuata et denticulata, mox ramificatione
marginali parce et irregulariter laciniata, eximie costata et opposite
venosa, serius e costa denudata stipitem elongatum formante prolifera.
Crescentia apicalis nee non structura anatomica pro genere typica.
Spermatangia et tetrasporangia soros inter venas formantia, cysto-
carpia eodem loco adspersa. Color in sicco sat obscure ruber. — Exs. :
Algae exsicc. Amer. Bor. num. 65. — -Hab. ad oras Californiae, cum
cystocarpiis et spermatangiis in cum tetrasporangia in lecta.

The largest specimen at hand measures about 15 cm. ; simple fronds
and marginal proliferations are up to 10 cm. long and 3 cm. broad,
with their greatest width generally a little above the middle. In old
specimens, the lower part of the primary costa forms a conspicuous
stipe, and the older proliferations are distinctly pedicellate.

Botanic Garden, Gothenbueg, Sweden,
October, 1921.

434 University of California PuUications in Botany [Vol. 7

BIBLIOGRAPHY

Agardh, J. G.

1898. Species Algarum, vol. 111:3, 1876.
1876. Species Algarum, vol. 111:1.

1879. Floridoenies Morphilogi. Kongl. Sv. Vetensk.-Akad. Handl., vol. 15,
no. 6.

BoRY, DE Saint Vincent

1828. Voyage "La Coquille," Botanique, Cryptogamie, p. 136, table 18,
isg. 1.

KUETZING, F. T.

1843. Phycologia Generalis.
1849. Species Algarum.

Kylin, H., and Skottsbekg, C.

1919. Zur Kenntniss der subantarktischen und antarktischen Meeresalgen
II, p. 94. Wiss. Ergebn. der Sell wed. Siidpolar-Exp., IV: 15.

Lamouboux, J. V. F.
1813. Essai Thalass.

NiENBERG, W.

1908. Zur Keimungs- und Wachstumsgeschichte der Delesseriaecen. Bot.
Zeit., vol. 66.

Schmitz, F.

1897. Rhodophyceae in Engler and Prantl's Naturl. Pflanzenfam., vol. I, 2.

Skottsberg, C.

1915. Notes on Pacific Coast Algae. I, Pylaiella Postelsiae, n. sp., a new type
in the genus Fylaiclla. Univ. Calif. Publ. Bot., vol. 6, pp. 153-164,
pis. 17-19.

Stackhouse, J.

1809. Tentamen mariuo-cryptogamicum.

SVEDELIUS, N.

1911. Ehodophyceae in Engler and Prantl, Naturl. Pflanzenfam., Nachtrage
zum 1 Th., 2 Abt.

EXPLANATION OF PLATE 50

Fig. 1. Cross-section through tlie costa of Phycodrys Setchellii.

Fig. 2. Growing apex of Hypoglossum Wodwardii.

Fig. 3. Growing apex of Phycodrys quercifoUa. Intercalary cells and branch-
lets of the main axis dotted.

Fig. 4. Growing apex of Phycodrys Setchellii. In the lower part the forma-
tion of cortical cells has begun.

Fig. 5. Cross-section through the stipe of Hypoglossum Woodwardii.

All figures drawn by the author at an enlargement of c. 700, and reduced to
half size.

[436]

UNIV. CALIF. PUBL. BOT. VOL. 7

[SKOTTSBERG] PLATE 50

UNIVERSITY OF CALIFORNIA PUBLICATIONS

IN

BOTANY

Vol. 7, No. 13, pp. 437-446 August 7, 1922

• ■ A K t

, YORK.
b../rAN5CAL
UA!<t>BN

UNDESCRIBED PLANTS MOSTLY FROM
BAJA CALIFORNIA

BY
IVAN MUEEAY JOHNSTON

Ephedra peninsiilaris sp. nov.

Fniticulus intricatus superne dense ramosus, ca. 8 dm. altus ;
ramis tenuibus, patentibus, numerosis, pallide viridibus; squamis 2,
1.0-1.7 mm. longis, Mi-% eonnatis. apice late triangularis vel obtusis,
basi plerumque brunneis incrassatis ; spiculis axillaribus. solitariis vel
geminatis; spicula mascula ovato-oblonga, 5-6 mm. longa; pedicellis
0.1 mm. longis; bracteis 3-4 vertieillastris. orbicularibus ; perianthio
obovato plernmqne exerto; staminibus distincte exertis; antheris 5-7,.
stipitatis vel subsessilibus ; spiculis f emineis acute ovatis, 6-8 mm.
longis ; bracteis frutescentibus ovato-orbicularibus, sessilibus, margine
seariosis; fructu multo exserto, solitario, triquetro, glabro, castaneo,
5-6 mm. longo.

^to^

Type. — Magdalena Island, Baja California, January 12, 1889,
T. S. Brandegee. Sheet no. 119069 in Herb. Univ. Calif.

This is the common and widely distributed Ephedra of Baja Cali-
fornia, which ranges over the peninsula from the cape region north-
ward at least to Calmalli and Cedros Island. The only other Ephedra
that I have seen from the peninsula is the three-bracted E. californica,
which occurs in the northernmost parts of the territory. I have
examined material representing the new species collected by Purpus
at Las Animas (269) and Calmalli (6), by Brandegee at INIagdalena
Bay and Calmalli, by Anthony on Cedros Island (281), and by myself
on Coronados Island in the Gulf of California (3757). Brandegee
(Proc. Calif. Acad. Sci., II, 2:205. 1899) reports the plant as E.
nevadensis; but the ranges of E. nevadensis and E. peninsularis are
separated by a broad geographical hiatus, and, furthermore, the two
species are quite distinct morphologically. The latter species differs
from E. nevadensis in its more slender and limber stems, shorter and

438 University of California Publications in Botany [Vol. 7

more united bi-aets, and fewer scaled aments. In the development of
the leaf st-ales the new species is remarkably like E. pedunculata, but
that species differs in its long reclining stems, long pedunculate
ovulate aments, and in its far removed range. The short, much united
leaf scale is a character which will distinguish E. peninsularis from
most North American species.

Stenophyllus nesioticus sp. nov.

Annuus multieaulis ; foliis filiformibus, 2-6 cm. longis, minus quam
1 mm. latis, glabris, suleatis, scabridis, quam culmo multo brevioribus,
vaginaruni faucibus pilosis; culmis tenuibus, levibus, numerosis,
obscure trigonis, prope apicem suleatis, 1-G dm. altis ; involucri
braeteis inconspieuis, quam spiculis plerumque brevioribus, 4—12 mm.
longis, exterioribus 2 interiores multo superantibus, anguste dilatatis;
spicularum braeteis ovato-lanceolatis, 3-6 in capitulum densum sessile,
4-6 mm. longum, 8-12-florum congestis, badiis; glumis late ovatis,
25-30 mm. longis, 3-ner\nis, extus tenuiter scabrido-pubescentibus,
margine plerumque fimbriatis; glumis inferioribus mucronatis;
staminibus 3; antheris lineari-laneeolatis, ca. 1.7 mm. longis, apice
mucronatis, loculis basi pilosis ; stylo trifido, quam ackaenium maturum
triplo longiori; nucis obovatis, ca. 1 mm. longis, albis, trigonis; stylis
coronatis, angulis levibus prominentibus, faciebus transverse rugosis.

Type. — San Benedicto Island, Revillagigedo Islands, ]May-Juue,
1897, A. W. Anthony 317. Sheet no. 201076 in Herb. Univ. Calif.

A species most nearly related to S. Warei Britt. (Tsolepis Ward
Torr.) of Florida, and to S. Selloiviana n. comb. (75. Scllowiana Kunth,
Enum. 2:208. 1837) of Brazil. From S. Warei (cf. Clarke. Ill,
Cyperac. t. 45, figs. 4-7. 1909) it differs in its darker and fewer
spikelets, narrower scales, and longer fringed involucral bracts; from
S. Selloiviana it differs in its fewer spikelets, and longer involucral
bracts which are neither obtuse nor ovate-oblong. From descriptions
it appears to be near some exotic forms, notably F. cinnamoneus
(Clarke) of Africa (cf. Clarke in Thiselton-Dyer, Fl. Trop. Africa,
8:432. 1902). The new species is apparently restricted to the Revil-
lagigedo Islands, which lie three hundred to four hundred miles
southerly from the tip of Baja California. I have seen material,
collected by Anthony and by Barkelew, from San Benedicto, Clarion,
and Socorro islands. The plant is mentioned as " Fimbristylis sp."
by Brandegee (Zoe, 5:27-28. 1900) and by Vasey and Rose (Proc.
U. S. Nat Mus., 13:145-149. 1891).

1922] Johnston: Undescrihed Plants Mostly from Baja California 439

Viscainoa geniculata var. pinnata var. nov.

Folia pinnis 3-5 pinnatis.

Type. — San Eaimonda Creek, Baja California, April, 1889,
Brandecjee. Sheet no. 109442 in Herb. Univ. Calif.

Viscainoa genictdata is so constant throughout most of its range
that this very pronounced variant deserves some recognition. It has
been detected in only a single locality and perhaps represents only a
local race. It is of interest as its leaves are similar to those of the
near related genus Chitonia. The type collection made at San
Raimonda Creek has definitely pinnate leaves which have a rhachis
1-3 cm. long ; and 3-5, distinct, spaced, oblong or lance-oblong pinnae
12-25 mm. long. A tendency toward the variety is found in Orcutt
1344 from Rosario (Herb. Univ. Calif, no. 109445), which has a
vigorous shoot on which two leaves are parted, although the rest of
the collection has its leaves entire.

Condalia Parryi var. microphylla var. nov.

Folia parva, minus quam 1 cm. longa ; f ructu quam eo specie!
minori.

Type. — Las Huevitas, Baja California, May 19, 1889, Brandegee.
Sheet no. 80099 in Herb. Univ. Calif.

Collections made on Cedros Island by Palmer (752) and Brandegee,
and one made at Las Huevitas by Brandegee are similar to one another,
but differ from the forms of C. Parryi which occur in Alta California
in their conspicuoush^ smaller leaves, stouter habit, and slightly
smaller fruit.

Euphorbia dentosa nom. nov.

Fuphoriia setiloia dentata Engelm. ; Boiss. in DC, Prodr. 1.5':44. 1862.

Annual; stems very slender. 1-3 dm. long, villous, internodes 15-
30 mm. long; leaves opposite, obliquely ovate, rather thin, glabrous or
sparsely villous, finely serrate, light green above, glaucous below, 5-
14 mm. long, 4-12 mm. wide, petiol&s 1-2 mm. long, stipules subulate ;
involucres solitary" or in close clusters, long villous, about 0.9 mm. high,
on peduncles 0.5-2.0 mm. long ; glands brownish ; appendages white,
lacerate, 0.3 mm. long. 1.0 mm. wide; seeds ashy, prismatic, 1 mm.
long, faces transversely wrinkled.

The plant described briefly above has been referred to as a variety
of E. setiloha, but it seems quite distinct from that species, differing in

440 Vnwersity of California Publications in Botany [Vol. 7

habit, in its much larger serrate leaves, and in its southern range. The
varietal name used by Engelmann is already employed in Euphorbia
as a specific name, so that it has been necessary to propose a new one.
The species is based on material collected at Cape San Lucas by
Xantus. I have studied, and the above diagnosis is based upon col-
lections made by Palmer (34) at La Paz, Grabendorfer at San Jase
del Cabo, and Brandegee at San Felipe; all from the cape region of
Baja California.

Euphorbia peninsularis sp. nov.

Perennis; caulibus 1-4 dm. longis. ut videtur ascendentibus vel
prostratis. brunnois, laxe dichotomeque ramosis. internodiis 20-45 mm.
longis, juvenibus dense tomentosis, vetustis praesertim ad nodes
sparsim canescenti-pubescentibus; foliis oppositis, integris subsessil-
ibus, rhomboideis, subfuscis, supra pallidioribus, apice obtusis, prope
basis latioribus, 1.5-2.7 mm. longis, 5-12 mm. latis, sparsim breviter-
que villosis ; petiolis 0.1 mm. longis ; stipulis lineari-subulatis, eiliatis ;
involueris in glomerulis foliosis ramos breves terminantibus, ca. 1.5 mm.
altis, villosis; pedunculis 0.5-1.0 mm. longis; glandulis 4, transverse
oblongis. brunneis, appendicibus petaloideis. albis, conspicuis, 1-2 mm.
longis, 2-3 mm. latis ; capsu-la ca. 2 mm. longa, pubescenti ; seminibus
ovatis, angulis 4 rotundatis. faciebus transverse lateque rugosis, 1.0-
1.3 mm. longis, 0.8-0.9 mm. latis.

Type. — San Jose del Cabo, Baja California, January, 1901, Purpus
325. Sheet no. 178322 in Herb. Univ. Calif.

The species here described is very distinct and appears to have no
immediate relatives. Its large oblongish leaves at first glance suggest
the species of the section Hypericifoliae, but its real relationships are
in the section Chamaesyce. It may possibly be related to E. leuco-
phylla, but that species, while resembling it much in structure and
disposition of the involucres, as well as in general habit, has smaller
rounded ovate leaves with conspicuously dentate margins. The out-
standing features of the plant are its large entire brownish subsessile
leaves and tomentose young stems. Besides the type, I have seen an
identical plant collected at the type locality by Brandegee (535).

Euphorbia podagrica sp. nov.

Annua ; caulibus prostratis, glabratis, 10-15 cm. longis, internodiis
6-10 mm. longis. nodis turgidis; foliis numerosis, oppositis. oblique
ovatis, integris, glal)ris vel paullo villosis, lamina 2-4 mm. longa et
1.5-3.0 mm. lata, petiolis ca. 0.7 mm. longis; sti])ulis eonnatis oeream
formantibus caule crescente laceratis; stipularum lobis obtusis vel
rudimentariis, apice plerumque villosis; involueris in racemis foliosis,

1922] Johnston: Undescrihed Plants Mostly fr&ni Baja California 441

ea. 6 mm. longis, anguste turbinatis, sparsim villosis, lobis 5 parvis;
glandulis 4, bnmneis, exappendulatis ; pedunculis validis, 0.2-0.5 mm.
longis; capsula glabra, 12-15 mm. longa, acute angulata; seminibus
prismaticis, levibus vel paullo rugosis, ca. 1 mm. longis.

Type. — Washes at Gold Mountain, Nevada, 1898, Purpus 6437.
Sheet no. 110920 in Herb. Univ. Calif.

Besides the type, I have seen collections of this species from ATil-
mont, Arizona (TJwrnher 341), from near Holtville, California
(Parish 8087), and from Imperial County, California (Wales 8).
The plants of the Californian collections are villous throughout, but
those of the other collections are glabrate. The outstanding characters
of E. podagrica are its annual prostrate habit, knobby stem nodes,
entire leaves, and small involucres with reduced unappenclaged glands.
It seems to come nearest to E. glyptosperma, but that species has
oblong serrate leaves, unthickened stem nodes, and seeds differently
roughened.

Securinega capensis sp. nov.

Fruticulus laxe ramosus, dioicus, ca. 1 m. altus ; caulibus rigidis,
cinereis praesertim ad ramos juvenes; ramis vetustis breviter calcar-
atis, foliis oblanceolatis, plerumque fasciculatis, 7-15 mm. longis,
3-4 mm. latis. glabris, subtus manifeste costatis, obscure venosisque ;
petiolis ca. 1 mm. longis; floribus masculis 2-3-glomeratis ; pedunculis
strigosis, ca. 1 mm. longis; pedicellis glabris, 2.5-5.0 mm. longis;
sepalis 4-5, ovatis; staminibus 5, ca. 2 mm. longis, circa discum
lobatum positis ; floribus femineis paucis, pedicellis brevibus validis ;
f ructibus juvenibus strigoso-canescentibus ; stigmatibus 3, reeurvis,
1 mm. longis, apice dilatatis complanatisque : ovario trilocnlari, loculis
interdum biovulatis; fructu brunneo, tenuiter pubescenti, manifeste
lobato, diametro ca. 9 mm., globoso-oblongo, pedicellis 3-6 mm. longis.

Type. — West side of cape region, Baja California, October 22, 1893,
Brandegee. Sheet no. 110393 in Herb. Univ. Calif.

Securinega capensis is a near relative of *S^. fasciculata comb. nov.
(Bernardia fasciculata AVats., Proc. Am. Acad., 18:153. 1883) and
has been referred to that species by Brandegee (Zoe, 4 :405. 1894) . It
differs, however, in the size of its leaves and in the shape and size of
its fruit. The new species is known only from the cape region of
Baja California,whence I have seen Brandegee 536 from San Jose del
Cabo, and Brandegee 's collection of October, 1893, from tlie "west
side of the cape region." The congener S. fasciculata has long
remained under Bernardia where AVatson first placed it with a query.
But there is no doubt that this past treatment is unsatisfactory, for

442 University of California Puhlications in Botmiy [Vol. 7

the plant has two ovules in eaeli cell of the ovarj', and so belongs to
the Phyllanthoideae (as Brandegee has pointed out), and not among
the Crotouoideae.

Tetracoccus Ilallii Braudg. (Zoe, 5:229. .1906) is very close to
8. fasciculata and I am in some doubt as to its distinctness. The
association of T. Hallii with the very different T. dioicus does violence
to natural relationships, for not only do the plants differ in the
number of cells to the ovary, but in sepal number and shape, in
pubescence, and in shape and position of leaves. The opposite leaves
in Tetracoccus dioicns clearly show a relationship to the alliance
called the Toxieodendrinae by Pax (E. & P., Nat. Ptianzenf., 3^31.
1896), whereas the characters of T. Hallii evidently relate it to
Phyllanthinae as defined by Pax (op. cit., 17). If one is to accept
the character evaluations maintained by the master workers in the
Euphorbiaceae, it is impossible to leaA^e T. Hallii congeneric with
T. (Jim'c}(s. Among the described genera the plant seems to fit better
in Securingea. and so it seems best to call it ;S'. Hallii n. comb.
Securinega Hallii, S. fasciculata, and S. capeTisis form a small natural
group in the section Fluggea, and though suggesting S. ramiflora
probably are to be associated with S. aciclothamnus : the relationships,
however, are not close, and the three species differ from other
Securinegae in habit and caruncular seeds. The following key con-
trasts the characters of Tetracoccus dioicus and the three Securinega
species discussed.

Ovary 4-celled; sepals on female flowers linear, becoming 3-5 mm. long; leaves

linear, opposite; plant glabrous throughout Tetracoccus dioicus

Ovary 3-celled; sepals on female flowers triangular or ovate, becoming 2 mm.
long; leaves spatulate or oblanceolate, alternate; young branches and fruit
pubescent
Fruit globose-oblong, rather strongly lobed, becoming 9 mm. in diameter,

flattened at base; leaf blades 7-15 mm. long, 3-4 mm. wide

Securingea capensis
Fruit oblong, weakly lobed, becoming 5 mm. in diameter, sunken in deeply
at base; leaf blades 9' mm. long or less, becoming 3 mm. wide

Sepals 6; male flowers on pedicels 5 mm. long, Californian

Securingea Hallii

Sepals 5; male flowers on pedicels 2 mm. long, Mexican

Securinega fasciculata

1922] Joh^istmi: Vndescrihed Plants Mostly from Baja California 443

Mentzelia hirsutissima var. nesiotes var. nov.

Corolla aurantiaea ; f oliis 1-2 em. latis, crasso-dentatis.

Type.— San Benito Island, March 28, 1897, T. S. Brandegee. Sheet
no. 138558 in Herb. Univ. Calif.

Typical 31. hirsutissima Wats, is known only from the type locality
on Angel de la Guardia Island, where in 1921 I re-collected it in a
small-flowered form. The species differs from M. involucrata in its
more narrow green, instead of broad scarious, bracts. There are two
varieties of M. hirsutissima, one from the peninsula of Baja California,
and the other, the one here described, from the west-coast islands. The
first variety may be called M. hirsutissima stenophylla n. comb. (M.
stenopliylla Urb. & Gilg, Nov. Act. D. Akad., 76:80. 1900). In the
species and in the variety nesiotes, the middle lobe of the anthers
equals or exceeds the lateral lobes, but in the variety stenophylla the
middle lobe is greatly elongated and twice the length of the lateral
lobes. The variety stenophylla ranges over the northern third of Baja
California. The variety nesiotes is kno"\^Ti only from San Benito,
Natividad, and Cedros islands, from which it has been reported as
M. involucrata (Zoe, 5:22-24. 1900) ; it is characterized by its orange
corolla, and not very thick coarsely toothed leaves. Possibly the
variety nesiotes is the basis of the report of M. triciispis from Cedros
Island (Urb. & Gilg, op. cit., 80), but Palmer 712, cited as the basis of
that report, is determined as M. adherens by Vasey and Rose (Contr.
U. S. Nat. Herb., 1:16. 1890).

Mentzelia involucrata var. megalantha var. nov.

Fetalis 3.5-4.5 cm. longis.

Type. — Near Salton. Riverside County, California, April 21, 1905,
Hall 5839. Sheet no. 68744 in Herb. Univ. Calif.

The species is characterized by its large, white, scarious, green-
marginecl floral bracts. In California it is most common in the
Colorado Desert from which it extends eastward into Arizona. The
species occurs in two forms, which are sharply separated by their
flower size. The typical form of the species is the small flowered plant,
with petals 1.5-2.5 cm. long, for in the original description Watson
gives the length of the petals as " an inch. ' ' The flowers in the variety
megalantha are very large and much resemble those of M. laevicaulis.
The variety ocurs only in the Colorado Desert, where, judging from
the relative amounts of material, it is less common than the species.

444 University of California Puhlications in Botany [Vol. 7

Mentzelia tricuspis var. brevicornuta var. nov.

Lobis latoralibns antherarum oblongis, rotnndatis, minus 0.5 mm.
latis.

Type. — Barstow, San Bernardino County, California, ^lay 14, 1903,
K. Brandegce. Sheet no. 108381 in Herb. Univ. Calif.

The species is unique in the section Bicuspidaria because of its
pedicellate flowers and petiolate leaves and bracts. It ranges from
Utah and Arizona into the eastern part of the Mohave Desert in Cali-
fornia. The variety brevicornuta is the form which the species assumes
near its western limit in the vicinity of Barstow. In typical tricuspis
the lobes of the filaments are linear and 2 mm. long at least ; but in the
new variety the lobes are short oblong and only about 0.5 mm. long.
Mentzelia tricuspis has been frequently misidentified as M. reflexa
Cov., no doubt because its fruits are reflexed and appear much as those
illustrated in Coville's (Contr. U. S. Nat; Herb., 4:108, t 9. 1893)
plate of reflexa. Mentzelia reflexa is not even closely related to M.
tricuspis and can be recognized by its smaller corollas and unlobed
filaments.

The species and varieties of ]\Ientzelia which make up the section
Bicuspidaria may be distinguished by the following key.

Flowers pedicellate; upper leaves and floral bracts petiolate

Lateral lobes of anthers linear, acute, 2 mm. long or more M. tricuspis

Lateral lobes of anthers oblong, rounded, about 0..5 mm. long

M. t. brevicornuta
Flowers, upper leaves, and floral bracts sessile

Floral bracts white and scarious, with green midrib and margin

Petals small, 1.5-2..5 mm. long M. involucrata

Petals large, 3.5-4..5 mm. long M. i. megalantha

Floral bracts green, not at all scarious

Middle lobe of anthers equalling or shorter than lateral ones

Corolla pale yellow; leaves 7-] 2 mm. wide, thickish, deeply lobed

M. hirsutissima
Corolla orange-yellow, leaves 10-20 mm. wide, thinner, coarsely toothed

M. h. nesiotes
Middle lobe of anthers twice the length of lateral ones....M. h. stenophylla

Crjrptantha inaequata sp. nov.

3—4 dm. alta, caulibus ramis laxe ramosis; foliis planis, lineari-
oblaneeolatis, 2-4 cm. longis; inflorescentia laxe racemosa. 4-12 cm.
loiiga ; calyce fructifero 2.5-3.0 mm. longo, lobo abaxillari maximo
hispidissimoque ; pedicellis brevibus denum 0.1 mm. longis; nuculis 4,
lieteromorphis. subtrigono-ovatis, acute marginatis; nuce larga quam
gynobasi longiore, 1.7 mm. longa, sulco supra medium clauso inferne
dilatato areolam triangulam formante.

1922] Johnston: Undescribed Plants Mostly from Baja California 445

Tifpe. — Pleasant Cailon, Panamint IMountains, California, ]\Iay 10,
1906, Hall & Chandler 6925. Sheet no. 100917 in Herb. Univ. Calif.

This is a species most nearly related to C. racemosa, but differing
from it in its sessile or very short pedicellate calyx, and in the shorter
lived herbaceous stems. Excluding C. pterocarya, it differs with C.
racemosa from all other winged fruited species in having hetero-
morphous nutlets. The plant is known only from two collections in
the eastern part of the Mohave Desert of California.

Cryptantha maritima var. pilosa var. nov.

Specie! similis sed calyce pilis longe patentibus praedito.

Type. — On stony ridges at Los Angeles Bay, Baja California, 1887,
Palmer 551.

This well marked variant, characterized by its pilose calyx lobes, is
frequent in the deserts of southeastern California. From Logan,
Nevada, where collected by Kennedy, it ranges south to Santa Agueda
{Palmer 2142) in Baja California.

Cryptantha racemosa var. lignosa var. nov.

Fruticosa, laxe ramosa, 2-4 dm. alta; ramulis floriferissimis ; in-
florescentia laxe racemoso-paniculata, per omnes partes plus minus
aequaliter distributa.

Type. — Panamint Cafion, Inyo County, California, ]\Iay 15, 1906,
Ball & Chandler 7034. Sheet no. 100892 in Herb. Univ. Calif.

Typical C. racemosa, which ranges south into Baja California from
the southern part of the Colorado Desert, is replaced in the Mohave
Desert and in the northern part of the Colorado Desert by a slender
diffusely branched very fioriferous form, which I am here naming
variety lignosa. The new variety differs from the species in its loosely
racemose-paniculate, instead of racemose, inflorescence, and in having
its flowers distributed more or less completely through the plant and
not confined to a terminal cluster or a peripheral belt. Piper appar-
ently recognized the two forms of C. racemosa, but in attempting to
name them described the wrong plant; his C. suffruticosa (Proc, Biol.
Soc. Wash., 32:42. 1919) being typical C. racemosa.

446 Universihj of California Publications in Botany [Vol.7

Houstonia australis sp. no v.

Poronnis grlabra ; ('jiiilil)us e radieo multis. erectis. slmplicibiis vel
iiitci'duni ramo iino, tei-ctibiis, 80-45 cm. altis ; folii.s lincaribns,
acuiniiiatis, 25-45 iiiiii. loiigis, 1-2 mm. latis, valde costatis, oi)positis;
petiolis attenuatis; stipulis rainiitis, 2-4, setiferis; corolla hypercrateri-
infundibiiliformi, 7-9 mm. longa lit videtiir nifa, tubo 3-5 mm. longo
faucibns 1.5 mm. longis, tnbnlatis vel tubiilato- infimdibidiformibus
lobis obloiigis vel ovato-oblongis, 2.5-3.0 mm. longis, intns villosis
antheris exertis; stylo incluso; hypanthio 0.5-0.7 mm. alto, quando
frntescente fere 2 mm. alto ; capsula globosa, 4^ infera, diametro 2 mm. ;
seminibus 0.3-0.6 mm. longis, ovatLs, concavo-convexis, scrobienlatis,
bnmneis.

Type. — Binorama, Baja California, September 27, 1899. T. S.
Brandegee. Sheet no. 201101 in Herb. Univ. Calif.

Houstonia australis i.s known only from the mountains of the cape
region of Baja California whence it has been reported by Brandegee
as H. hreuipes. The nearest relative of H. aaistralis is H. a ng list i folia,
which has a far removed range, oblong instead of globose fruits, and
smaller corollas which vary between 4 and 7 mm. instead of between
7-9 mm. in length. Tloustonia hrevipes, which is lacking in the cape
region, has larger (9-15 mm. long) flow^ers, and narrower (under
instead of over a mm. wide) sessile rather than short petiolate leaves.

UNIVERSITY OF CALIFORNIA PUBLICATIONS

IN

BOTANY

Vol. 7, No. 14, pp. 447-498, plates 51-61 August 11, 1922

LPJRARY
NEW YORK
BOTANICAL

UAKDEN

THE MORPHOLOGY, DEVELOPMENT, AND
ECONOMIC ASPECTS OF SCHIZOPHYLLUM

COMMUNE FRIES

BY

FEEDEEICK MONROE ESSIG

CONTENTS

PAGE

I. Introduction 448

II. Material and Technique 449

III. Morphology 451

1. General characteristics of the sporophores .■ 451

2. Description of the mature sporophore 453

3. Microscopic structure 454

IV. Growth of the sporophore 458

i. Development in general 458

2. Origin and development of the "gills" 461

3. Taxonomic interpretation of the structure and development of

the sporophores 463

V. Economic aspects 464

1. Geographical distribution 464

2. List of host plants 465

3. Extreme hardiness of the fungus 466

4. Relation of the mycelium to cells of dead wood 467

5. Growths upon fresh wood and living trees 468

6. Methods of infection under natural conditions 470

7. Association with other wood decay fungi 471

VI. Summary and conclusion 472

VII. Acknowledgments 473

Literature cited 473

Explanation of plates 478

448 Vmversity of Calif ornm Pudlicati&ns in Botany ["^ol. 7

I. INTRODUCTION

The peculiar split character of the gills of Scliizopliylluin called
the attention of botanists to this genus in comparatively early times.
Mention of this fungus appeared in Dillenius' "Catalogus plantarum
sponte circa Gissam nascentium" more than 200 years ago. Since
that time (1719) it has been frequently collected and described, and
in the last quarter-centurj^ has been reported to be of considerable
economic importance, but the literature upon the common Schizo-
phyllum is singularly fragmentary and incomplete. Early mention
was in the form of collection notes or brief descriptions of the dried
sporophores. Many of these appeared in print between the time of
Dillenius and the latter part of the 19th century.

In 1884 W. G. Smith found Schizophyllum growing upon ensilage.
This is the earliest mention of its economic importance in available
literature. Since this article appeared Schizophyllum has been
reported as parasitic upon a large variety of hosts, but papers relating
to the exact nature of the parasitism are not to be found.

There is also very little published concerning the microscopic struc-
ture of the sporophores or vegetative hypliae. A few sentences and
one figure in Duller 's "Researches on Fungi" (1909) give an inkling
as to the structure of the hymenium and hyphae composing the gills.
No mention or illustration of contained protoplasmic structure has
been found. Rumbold (1910) described and featured the walls of
the vegetative hyphae, but did not mention the cell contents.

The literature upon the morphology of the sporophores, however,
is much more complete than that upon the microscopic structure.
Early descriptions were mostly confined to the structure of desiccated
specimens. Hasselbring in 1907 called attention to the development
of the sporophores and the peculiar relationship of the hymenophore
to the pileus and stipe. Buller (1909) gave a full description of the
mature sporophores, origin of the secondary lamellae, marginal split-
ting, and incurving of the lamellar plates. Adams (1918) described
the origin and development of the lamellae. The phenomena he
describes, however, do not agree with the phenomena displayed by
sporophores growing in their normal habitat on the University of
California campus.

1922] Essig: Morpholagy of Schizophyllum commune Fries 449

This paper is presented in an effort to give a more accurate and
complete description of the sporophores; a description of the micro-
scopic structure of the hyphae which go to make up the sporophores
and the vegetative mycelium ; an account of the origin and develop-
ment of the gills as found in specimens growing in the field at Berkeley,
California ; and a report on the economic aspects of the fungus.

According to Saccardo (1887-1895), there are twelve species of
Schizophyllum, all of them being tropical or subtropical except S.
commune Fries, which is distributed throughout the northern and
southern temperate zones. Hennings (1898) stated that all of the
twelve forms described probably constituted only two or three distinct
species, and this view seems to the writer to be more nearly correct.
The original work reported in this paper is confined entirely to ScJiizo-
phyllum commune Fries,^ but in the table of geographical distribution
and the list of host plants reference is made to the genus as a whole,
as it is extremely difficult to distinguish between the one species of the
temperate zones and the several tropical species described.

II. MATERIAL AND TECHNIQUE

The sporophores studied came from three sources:

1. Decayed wood in the field.

2. Decayed wood kept in moist chambers in the laboratory.

3. Specimens from the Herbarium of the University of California.
The specimens obtained from the field were collected over a period

extending from September, 1917, to February, 1920. They were
found growing under natural conditions upon the wood or bark of
Acacia sp., Quercus agrifoUa, and, UmhellulaHa calif ornica. In all
several hundred specimens have been collected, ranging in age from
apparently a few hours to two years, and in size from less than 1 mm.
to 5 or 6 cm. in diameter. More than a hundred specimens of very
young sporophores were obtained during the autumn of 1919 from a
single log of Vmhellulana partly imbedded in gravel along a small
creek near the laboratory.

As an experiment short sections of small trunks and branches of
trees infected with Schizophyllum were placed in shallow pans of water
and covered with bell jars. Air was admitted by supporting the bell

1 For a complete list of synonyms see Greville (1824) and Murrill (1915).
The most important are Agaricus alneus Linnaeus, SchizophyUum aJneum Schroeter,
SchizophyJJus alneus Murrill.

450 Umversity of California PuhUcaiions in Botmiy [Vol. 7

jars so tliat the lower edge did not quite touch the surface of the water.
Under these conditions sporophores could be induced to grow through-
out the year, new ones appearing successively as the mature specimens
were removed. In this way the development of the sporophores could
be watched from day to day and compared with conditions found in
the field. It might be stated here that at no time was there any essen-
tial difference between specimens found in the field and those grown
in moist chambers in the laboratory. The moist chambers also fur-
nished an excellent opportunity for the study of regeneration phe-
nomena.

In the third source of material, the Herbarium of the University
of California, specimens from many localities in California are pre-
served, as well as some from Whidbey Island, Washington ; from
Ontario, Canada ; and from France. These sporophores were used for
comparison with respect to general morphological characters.

In the matter of technique no claim is made for originality. It
was found at the beginning of th€ work that the processes used in the
treatment of the fleshy fungi did not give satisfactory results when
applied to Schizophyllum. After experimenting with many methods
those outlined below were found to be the best suited for use with this
fungus.

The killing and fixing agent used for the young sporophores was a
70 per cent solution of alcohol with 6 c.c. of commercial formalin
added to each 100 c.c. of the alcoholic solution. This not only killed
and fixed the material but preserved it indefinitely. Spores were
caught in a film of albumen fixative on a slide and fixed in 100 per cent
alcohol, thus hardening the fixative and fastening the spores to the
slide.

Most of the sectioning was done on a rotary microtome, the material
being frozen in a solution of gum arable (compare Gardner, 1917).
Some of the specimens were imbedded and sectioned in paraffin, but
this method did not give good results, as the material became hard and
difficult to section. The sporophores for the study of early develop-
mental stages were sectioned individually, and all the sections from a
single specimen were preserved in a vial. From these vials the sec-
tions were poured into shallow dishes where all could be seen. Only
those sections cut at or near the median plane were selected for mount-
ing. Albumen fixative was used to fasten the sections on tlie slide.
Great care was used in orienting the imbedded sporophores before
sectioning, as oblique sections through the revolute hymenial margins

1922] Essig: Morphology of Schizophyllum commune Fries 451

lead to entirely erroneous interpretations of the structure of the sporo-
phore. Thus sections cut obliquely through specimens with well devel-
oped "gills" which can be seen unmistakably with the unaided eye
appear as though the hymenium lined a series of chambers, or as if
the "gills" in the middle of the section were normal, with "hymenial
chambers" at each edge. Certain sections cut in this manner corre-
sponded in many ways to figures given by Adams (1918). Material
for young developmental stages was sectioned from 20 to 25fji in thick-
ness so that the sections would remain entire. For eytological work
the sections were cut 5 and 10 micra thick. Sections as thin as 5 micra
will not hold together well, so that, for the finer sti"ucture, it was neces-
sary to use only fragments.

Flemming's triple stain was used for some sections, but was not so
satisfactory as safranin alone. Using a two-minute period in a 3 per
cent solution of safranin in 50 per cent alcohol and washing out rap-
idly, a fair differentiation was obtained. The nuclei stain deep red,
the cytoplasm a very light pink, and the cell walls an intermediate
shade. All efforts to make the nuclei stand out more clearly by coun-
terstaining resulted in failure. For staining spores, a 48-hour period
was needed, as in a shorter period the stain would be almost entirely
removed in the washing-out and dehydrating operations preceding
clearing and mounting. The sections and spores were cleared in xylol
and mounted in Canada balsam.

III. MORPHOLOGY

1. GENERAL CHAEACTERISTICS OF THE SPOROPHORES

Schizophyllum is distinctly a xerophyte. The sporophores are
found in either of two conditions :

1. In dry weather the sporophores are desiccated, hard, and some-
what brittle (fig. 1, pi. 51). The margin of the pileus is curved inward,
decreasing the width of the sporophore about 25 per cent. Each
hymenial plate is incurved on the side toward the hymenium. The
hymenial surface is hidden and protected. Only the villous sterile
surface of the hymenial plates can be seen from the lower side. There
is no discharge of spores. This is an inactive period.

2. In moist weather the sporophores take up water and become
flexible and leathery in consistency. The pileus margin is only slightly
curved downward. The gill plates unroll and extend vertically down-
ward, or nearly so (fig. 2, pi. 51). Spore discharge begins about an

452 University of California PiihUcations in Botany [Vol. 7

hour after the sporophores are moistened, and continues for a maxi-
iniiin period of about two weeks. However, under natural conditions
liie period is usually sliorter, as the spores cease to fall as soon as the
sporophores become dry. Growth takes place, the most actively grow-
ing region being at the pileus margin and at the edge of each gill plate.
Specimens are naturally most commonly collected when in a dry
condition.

The form of the fruit bodies varies greatly, depending in great
part upon the position of the surface of the substratum. Thus the
sporophores growing upon the under surface of a piece of wood are
quite different in form from those found upon the upper surface, and
those growing from a vertical surface differ from both of the preced-
ing ; but for every one of these three positions the form is fairly con-
stant.

The diversity of form is due to the peculiar organization of the
sporophore, the stipe being attached to the upper surface of the pileus,
with the hymenophore upon the opposite side of the pileus away from
the stipe. Thus Schizophyllum differs from all other stipitate members
of the Agaricaceae so far studied. This difference was conclusively
demonstrated by Hasselbring (1907), who grew the sporophores upon
a klinostat. He showed that, when not influenced by the force of
gravity, the stipe was always attached near the center of the pileus,
but on the opposite side from the ' ' gills. ' ' This unusual organization,
according to De Bary (1887), is also possessed by Cyphella, a member
of the Thellephoraceae.

The form which Hasselbring found is the one assumed by fruit
bodies growing in nature from the under surface of the substratum.
There is a stipe, which is usually short, attached to the center of the
upper surface of tlie pileus. The sporophore is shaped like a broad
funnel or bell, the hymenium lining the inner, and also the lower, sur-
face (fig. 1, pi. 52), On a vertical surface the form of the sporophores
depends in great part upon the length of the stipe (fig. 2, pi. 52). If
this structure is short, it is attached to the pileus near the edge, and
the sporophore is ear-shaped. If the stipe is long, it may be curved
down\vard at the outer end and be attached to the pileus near the
center, as in specimens grown upon an under surface. Then the shape
is that of a curved trumpet. In the ear-shaped forms the stipe is so
short that it cannot curve downward, so that the upper edge of the
young sporophore is stimulated by gravity (as shown by Hasselbring)
to grow more rapidly than the lower edge. Thus the hymenium is

1922] Essig: Morphology of Schizophyllum convmune Fries 453

brought into a more advantageous position for spore discharge. On
an upper surface the stipe is attached to the pileus at the very edge
(fig. 3, pi. 52). The lower edge of the young trumpet-shaped sporo-
phore never develops. The gill plates radiate outward from a place
near the attachment of the stipe.

The sporophores are borne singly or in groups. The groups may
contain from a few to several scores of specimens attached to each
other at the base of the stipe. Sometimes more than one sporophore
grows upon a single stipe, but such an occurrence is rare. Only cer-
tain members of a group reach maturity, a large percentage never
developing beyond a very early stage.

2. DESCRIPTION OF THE MATURE SPOROPHORE

The shape of the individual sporophore varies from broadly bell-
shaped with a centrally attached stipe to ear- or racket-shaped with
the stipe attached to the edge of the pileus. The edge of the pileus
may be entire or more or less deeply lobed. If the lobes are large and
deep, they may have secondary lobing. The size of the mature sporo-
phores ranges from 2 mm. to 5 cm. in length and from 3 mm. to 6 cm.
in width.

The color of the pileus may be silvery or velvety white, gray, or
cream colored. The sterile surfaces of the hymenial plates are a dark
gray with often a purplish tint. The hymenium is a shiny brownish-
gray. There is a considerable variation in the color of all parts of
the fruit bodies, depending upon the age and whether they are w^et
or dry.

A stipe is usually, but not always, present. Its presence and length
depend upon the amount of moisture in the substratum and atmo-
sphere at the early stages of growth, a maximum amount of moisture
inducing a greater growth in length. The length varies from 1 mm.
to 2 or 3 cm. The form is cylindrical. The stipe rarely attains a
width of more than half a centimeter.

The pileus is covered with a dense mass of thick-walled hyphae.
If these hyphae are vertical and remain free from each other, the
surface appears velvety. If they are agglutinated into groups at the
upper ends, the surface is rough and scurfy. If the upper ends form
a horizontal layer, the surface has a silvery sheen. The depth of the
hyphal covering varies from 1.5 to 2.5 mm. The pileus is made up
of thick-walled, septate hyphae. These are closely packed together,

454 Vmvcrdty of California Puhlioatwns in Botaniy [^^oi- 7

but do not in any case constitute a pseudoparenchymatous tissue. The
upper layer of the pileus flesh is pigmented brown.

The lamellar plates extend downward from the pileus (fig. 22, pi.
52). They are arranged in pairs; each pair, with the sterile, hairy
surfaces together, giving the appearance of a lamella. Each plate,
however, is independent of the other to a certain degree, and may
varj^ from it in size and shape. A hymenial plate may attain a depth
of 3 mm., the depth depending entirely upon the number and length
of the growing periods following the origin of the plate, for growth is
continuous throughout the duration of favorable conditions. Each
plate exhibits a growth region at the margin, which, after the earliest
stages, is continuous with and similar to the margin of the pileus.
Since the plates arise successively in pairs, a great many different
ages and sizes may be found in the same mature sporophore.

The hymeniuni either covers a much divided single area or is
separated into several different areas in the same sporophore, as in
plate 53. The elements in the hymenial layer are closely crowded
together, and in old specimens tightly adhere to each other, so that
a large area of the hymenium may be removed from the subhymenial
layer without separating the basidia from one another.

3. MICROSCOPIC STRUCTURE

The study of the microscopic structure of Schizophyllum presents
considerable difficulty. Upon dehydration the sporophores become
hard and brittle, so that the paraffin method of obtaining sections is
impracticable. When, by using other methods, sections are obtained,
it is found that the hyphal walls are thick, the segments extremely
long, and that the nuclei are small and difficult to differentiate by
staining.

The vegetative hyphae commonly branch, but not with great fre-
quency. The branching rarely occurs at or near a septum, but usually
takes place about the middle of a segment (fig. 1, pi. 54). The hyphae
are of two sizes, one having a diameter of 3 to 5/^, and the other being
only about 1 or 2ju, wide. The finer hyphae are particularly abundant
w^hen the mycelium is grown upon artificial media, but are also found
to some extent in wood. The hyphae are often covered with small
tubercles, as was described by Brefeld (1889) in S. lohatum and Miss
Rumbold (1910) in S. commune. This seems to be a distinctive char-
acter. The function of these lateral projections (fig. 2, pi. 54) is not

1922] Essig: Morphology of Schizophyllum commune Fries 455

apparent from their structure. It is possible that they aid in the
absorption of food materials, since they closely resemble haustoria in
shape and are found only on those hyphae which are purely vegetative.
Clamp connections (fig. 1, pi. 54) are found at more than half of the
septa in actively growing mycelium. They have been observed upon
the hyphae of members of the Hymenomycetes since the earliest studies
of their cell structure. Though they occur in a great number of fungi
in this group, their exact function is not understood. Harper (1902)
suggested that they possibly facilitated the exchange of food materials
between segments, but just how this is accomplished is not clear. The
length of the segments varies from about 30/a to more than 200/*, the
usual length being about 80/*. The thickness of the wall varies from
about 0.1 to 0.5ju,. In rapidly growing hyphae there are few vacuoles
and these are small. The protoplasm is of fine granular structure and
very homogeneous. As the mycelium becomes older the vacuoles
enlarge, oil droplets are formed, and many of the cells collapse. There
are two nuclei to a segment (fig. 1, pi. 54). They are small and spher-
ical, about 0.3 to 0.5/A in diameter. Their structure is granular. No
nucleoli have been seen. The nuclei are usually found about 10 to 20/1
apart near the center of a segment. These compare very w^ell with
the nuclei in the vegetative hyphae of Hypochniis subtilis (Harper,
1902, fig. 1, pi. 1) . Maire (1900) found but one nucleus in the "cells"
of the mycelium of Coprinus radiatus.

The hyphae which compose a sporophore are of several different
types. One kind includes those which form the hairy covering of the
pileus and sterile surface of the hymenial plates; another makes up
the pileus and tramal structure, and the third forms the subhymenial
layer. Again, each of these types varies somewhat according to the
age and state of development of the sporophore.

The hyphae which cover the pileus and sterile surfaces of the
hymenial plates are composed in nearly all cases of but a single seg-
ment, which may be as long as 3 mm. These hyphae are of a fairly
uniform size and length (fig. 3, pi, 54). They are irregularly curved
and tangled together. The walls are in the younger stages fairly thin,
but as development proceeds they gradually thicken until in very old
specimens the lumen has almost entirely disappeared. Two nuclei
are present in each hypha of the hairy covering. They are similar in
size and shape to those of the vegetative hyphae. They are found
regularly near the base of the segment.

456 Vmverdty of Californm Publications in Botany [Vol. 7

The filaments which constitute the solid portion of the sporophore
arc ill tlioir vonngrcr stages similar to the larger hyphae which make
np the vegetative mycelium (fig. 4, pi. 54). As the fruit bodies become
older the segments lengthen, and the walls thicken until they are about
eipial in thickness to the width of the lumen (fig. 5, pi. 53). When
the microscope is focused up and down upon thick sections cut trans-
versely across the hyphae it is seen that the filaments are loosely
coiled in a fairly regular spiral, some turning clockwise and others
counter-clockwise. In old sporophores the hyphae adhere tightly to
each other wherever they are in contact. In the earlier thin-walled
state the segments are filled with cytoplasm and have two typical,
small nuclei. This is precisely the situation found by Harper (1902)
in Coprinus ephemerus and Hypochnus suhtilis. Clamp connections
are numerous, but no spine-like tubercles appear. The protoplasmic
contents of the thick-walled hyphae of older sporophores are masked
by the walls to such a degree that the number of nuclei present can-
not be determined. In Coprinus ephemerus (Harper, 1902) there are
many nuclei in the old "cells" of the pileus and stipe. Maire (1900)
found the same to be true in a large number of the fleshy Agaricaceae
which he had examined.

The subhymenial layer is composed of hyphae which are thin-
walled, and which retain the characters displayed by all the hyphae
in their earlier state. Branching is common, and clamp connections
are plentiful. There are two nuclei to each segment. The segments
are rich in cytoplasm.

The basidia are borne at the ends of thin-walled hyphae. They
are only slightly larger in diameter than the hyphae which bear them.
All the elements in the hymenium are similar (fig. 6, pi. 54), that is
there is no distinction between potential basidia and paraphyses.
Mature basidia project beyond the hymenial surface. The basidia
come to maturity in succession, only a few in a relatively large area
being found with spores attached at any one time. The basidia meas-
ure 5 by 20/x. Each has four long, slender sterigmata and bears four
spores. Immature basidia display two nuclei. No fusion of these
nuclei has been observed, but in later stages four have been seen. It
is therefore presumed that the usual fusion and two successive divi-
sions, as described by Wager (1893), Maire (1900), and Harper
(1902), have taken place.

The spores en masse are white. When seen with the microscope
they are nearly hyaline and of an olive green shade. There is a con-

1922] Esdg: Morphology of Schizophyllum commune Fries 457

siderable difference of opinion in the literature as regards the shape
and size of the spores. The early writers, such as Fries (1821), Cooke
(1871), and Saccardo (1887), stated that they Avere subglobose, about
2.5ju, in diameter. Morgan (1890) called attention to the fact that
the spores he had been examining averaged 5-6 by 2.5/x, and won-
dered if a mistake had been made, or if his were possibly a different
species. The possibility of a different species seems unlikely, for later
Hennings (1898) and Rumbold (1910) in Europe and Murrill (1915)
in America have found them to be oblong, at least twice as long as
broad. It is possible that the globular bodies supposed by the early
writers to be spores were nothing but the peculiar structures which
are shed by the dried sporophores when first they are wetted to obtain
a spore print. These bodies and no spores are dropped by old her-
barium specimens, which have lost their vitality, when they are mois-
tened. The spores occasionally possess small vacuoles. No oil drop-
lets have been demonstrated. The wall is thin. The spores are densely
filled with protoplasm. At shedding time two nuclei are present
(fig. 7, pi. 54).

The number of nuclei in the spores of the Hymenoraycetes so far
examined is either one or two. One was found in the spore of Hypoch-
nus suhtilis (Harper, 1902), Amanita vaginata, Tricholoma virgatum,
and Cantharellus infundihuliformis (Rosenvinge, 1886), while two
were found in Craterellus cornucopioides, Clavaria vermictctaris, Bole-
tus edulis, and 5. variegatus (Rosenvinge, 1886). Maire (1900), after
studying some thirty species, stated that there might be either one or
two nuclei in a spore. In the case of two nuclei, the single nucleus
divides as soon as it enters the spore from the basidium, instead of
just preceding the first segmentation during spore germination.

Spores germinate readily in water and in a great variety of culture
media. The spores first swell to nearly twice the normal size ; then a
germ tube appears at either one or both ends (fig. 8, pi. 54). The
width of the germ tube often approximates that of the spore, and as
a result the identity of the spore may soon be lost. The length that
the tube attains prior to segmentation depends to a certain extent
upon the nature of the culture medium, segmentation occurring earlier
when the medium is rich in food materials. In tap water growth
ceases about the time the first septum and branch appear. Branching
may occur either before or after the first septum is laid down, but it
usually occurs about that time.

458 University of Calif ornm Puhlications in Botany [^^o^^- 7

IV. GROWTH OF THE SPOROPHORE

1. DEVELOPMENT IN GENERAL

The development of the sporophores was early looked to for an
explanation of the peculiar pairs of hymenial plates which charac-
terize SchizophyUum. Fries (1821) believed that they arose as ordi-
nary gills and were split by drying. This view was again brought
forward by Fayod (1889), and still later by Buller (1909). Hoffman
(1860) believed the sporophore to be divided into a series of lamellar
systems, considering all the secondary gills to belong to the primary
hymenial plates which enclosed them. His view was adopted by Winter
(1884).

Adams recently (1918) made the statement that each lamella con-
sists "of the adjacent walls of two gill cavities which originate endo-
genously as tubes in the substance of the carpophore. The gill cavities
(tubes) split along their lower edges and lamellae are thus completed."

In Hasselbring's (1907) paper on "Gravity as a Form-Stimulus
in Fungi" appears the statement that "they [the young sporophores
of S. commune] appear as small outgrowths resembling simple forms
of Clavana, and attain a length of about one centimeter. Early in
their development a cup-like depression appears at the summit, and
within this the rudimentary lamellae are formed, radiating from the
center. ' '

Two distinctly opposed views have, then, been advanced by Hassel-
bring and Adams. Adams held that the "gills" originated endogen-
ously as the sides of horizontal tubes which later ruptured at the
lower edge and exposed the hymenium, while Hasselbring claimed that
the "gills" arose exogenously upon the surface of an apical cup-like
depression. It seems unlikely that Buller and the previous writers
had access to sporophores in the first stages of development, for they
made no statements concerning the early appearance of tlie fruit
bodies. Both Adams and Hasselbring, however, grew the sporophores
through all their stages in the laboratory. Adams grew his in flasks
on agar media from "immature carpophores collected in the field."
Hasselbring caused them to grow from ' ' pieces of a maple branch con-
taining the mycelium — placed on a klinostat."

The writer has had an opportunity to study the origin and devel-
opment of the sporophores both on a log of Vmhcllulai-ia caUfornica

1922] Es^ig: Marphology of Schizophyllum cammune Fries 459

(California Bay) in the field near the laboratory where more than
a hundred sporophores have grown during the autumn and winter of
1919-1920; and in the laboratory where scores of sporophores have
appeared upon the wood of Acacia, Quercus, and Umhellularm in moist
chambers. Dozens of the fruit bodies have been sectioned, either longi-
tudinally or transversely.

No phenomena in the course of the development of the sporophores
have been observed that in any way approximated those described by
Adams (1918) for Schizophyllum commune. By cutting oblique sec-
tions through small mature specimens his figures 2 to 7 in plate 9
may be imitated with fair accuracy, but these sections cannot permit
of such an interpretation as he has given for his sections. The crena-
tures which he shows in figures 2 to 5, plate 9, are entirely absent in
all of the 160 or more young specimens the writer has examined. The
sporophores which grew here in the laboratory and in the field devel-
oped much as was described by Hasselbring (1907).

The fruit bodies appear first as small, loose tufts on the substratum.
These develop into small white woolly projections either short and
hemispherical or prolonged into horn-like structures, "resembling
simple forms of Clavaria" (fig. 2, pi. 56). The end is either rounded
or conical. These small bodies are covered with loose tangled hyphae.
Later the apex becomes smooth, slightly darker in color, and covered
with shorter hyphae.

Next a single pore appears at the apex (fig. 3, pi. 56), as Hassel-
bring (1907) found. Early stages which show the origin of this pore
have been difficult to distinguish, as it develops within a few hours
after the formation of the buttons or horns, and the loose hairs at the
apex screen its first appearance. Longitudinal sections at this stage
show first a differentiation of the hyphae just behind the apex (fig. 1,
pi. 59). This region stains more deeply than the remainder of the
section. The growth at this place is accelerated, and, as increase in
size takes place behind the apex, the hyphae at the tip are pulled apart
(fig. 2, pi. 59). The hyphae beneath the rupture form a palisade layer
which extends laterally into a plane surface (fig. 3, pi. 59). The
growth then becomes more rapid at the edge of the layer, producing
it outward into a saucer-shaped and later a cup-shaped depression.
In this stage it resembles a small sporophore of a Peziza (fig. 4,
pi. 59).

In all cases observed by the writer the so-called lamellae have
originated upon the surface of this apical cavity (figs. 4-14, pi. 52),

460 University of Calif ornia Puhlications in Botany [^'ol. 7

which surface constitutes the hymenium primordium. This is exactly
as described by Ilasselbring (1907), but he did not go into detail
concerning tlie i^laeenient or development of the hymenial plates.
The i)ore attains a width of from 1.5 to 2 mm. before the appearance
of the first pair of plates. Dozens of specimens in the "apical depres-
sion " or " peziza ' ' stage have been observed where there was no indi-
cation of lamellae. By splitting the specimen in half longitudinally
the entire surface of the hymenium primordium may be examined
with a hand lens. Microscopical examination of prepared sections
fails to disclose any indication of closed chambers or of "gills" before
the "lamellae" are plainly visible upon the surface of the hymenium
priniordium.

The placement of the "gills" may be easily observed in actively
growing moist specimens (figs. 4-14, pi. 52). They arise as short,
isolated ridges upon the surface of the hymenium primordium. The
primary ridges arise successively from a point beneath the attachment
of the stipe, and grow outward in a radial direction until they finally
unite with the edge of the pileus. The secondary "gills" originate
between the gills already developed, but do not extend so near to the
stipe as do those already formed. They occupy, as Buller (1909)
noted, an isolated, subterminal position within the interlamellar space
in wliich they have been formed. As growth proceeds, however, the
distal ends gradually approach the pileus margin and eventually unite
with it, as do the primary ridges.

Soon after a "gill" unites with the pileus margin, the pileus
becomes split in from the edge, though often this is not disclosed
upon the dorsal surface because of the hyphal covering. This mar-
ginal splitting is doubtless to some extent hygroscopic, as specimens
kept moist from the first are split only slightly, while those subjected
to alternate wetting and drying are split farther toward the stipe ends
of the "gills," dividing the pileus as well as the hymenium into narrow
finger-like projections, the crenatures of Buller (1909).

Occasionally there is an unusual placement of certain "gills."
Sometimes they arise at an angle to the radial direction. In this case
they frequently remain short and isolated. Often there is a consider-
able sterile area between the two hymenial plates (fig. 1, pi. 60). This
is very commonly found at the stipe end of "gills" in lateral sporo-
phores. Rarely the first few primary gills formed unite with each
other and the pileus margin to divide the hymenium primordium into
several separate areas, in which the secondary "gills" later are formed
(fig. 15, pi. 52).

1922] Essig: Morphology of Schizophyllum commune Fries 461

Upon a superficial examination the "gills" appear to be much
branched. This is especially noticeable in dried specimens (fig. 1,
pi. 51). The secondary "gills" are not attached to the primary ones,
however, but fit in between and beneath them. Branching occurs but
rarely, and is the result of the anastomosing of two "gills" in the
early stages of development.

2. OEIGTN AND DEVELOPMENT OF THE ''GILLS"

The origin and development of the lamellae has been studied in a
fairly large number of members of the Agaricaceae. In all of these
Atkinson (1916) recognized two general types. In the first, which
he called the " Agaricus" type, the hyphae form a palisade layer at
the roof of a well-developed annular cavity which appears on the
under side of the j^ileus ; and from this palisade layer, which is the
hymenium primordium, the lamellae grow downward into the cavity.
In the second, or "Amanita" type, the gills originate as bars radiat-
ing out from the stipe to the under surface of the pileus. The origin
and development of the "lamellae" in Schizophyllum commune is
entirely distinct from either of the above types. The gills grow out-
ward from a palisade layer which forms the lining of a single apical
depression or cup.

The origin of a "gill" is evidenced in cross-sections in two ways:
either a split appears in the palisade layer and the edges grow out-
ward (fig. 6, pi. 56) ; or a small area of the palisade layer becomes
loosened, grows outward a short distance, and then splits in the middle
to a point beneath the original primordial layer (fig. 9, pi. 56). In
both cases the growth continues in the same manner. The hyphae
beneath the edge of the hymenium on each side of the split grow out-
ward rapidly and cause the hymenial edges to turn downward, and
by marginal growth a pair of hymenial plates are soon formed (figs.
1-4, pi. 57). Growth continues at the edges of these plates through-
out the life of the sporophore, so that in very old fruit bodies some
gill plates may be comparatively deep,

Buller (1909), noting the fact that, in cross-sections of the mature
specimens, the tramal layer was split to different depths, thought that
the "gills" arose entire and were later split due to hygroscopic ten-
sions. This theory had been earlier advanced by Fayod (1889), who
claimed that specimens grown under water have entire "gills." In
attempting to demonstrate Fayod 's statement it was found that sporo-
phores grow with difficulty under water and decay after a few days.

462 University of California, Puhlications in Botany [Vol. 7

Those "gills" which did arise under these conditions, however, showed
no departure from the process as described by the writer above. In
all cases observed the plates have been separated at the edges from
the very first. In later stages they may be separated only a short dis-
tance toward the pileus, or they may be split to any depth in the
tramal hyi)hae or pileus, or even completely through the flesh of the
pileus. Each "gill," after it unites with the pileus margin, is split
more deeply at the outer end, and the depth decreases toward the stipe
end, as does likewise the size of the "gill."

The si)litting or loosening of the hymenium to permit the origin
of the paired hymenial plates is due to the same tensions Avhich cause
the "gills" themselves later to be split to different depths. The addi-
tion of new elements to the hymenial layer does not keep pace with
the growth of the hyphae beneath the hymenium. When the hyme-
nium has attained a certain width the tension upon the closely crowded
elements of the hymenial layer is so great that it is either split longi-
tudinally near the center of the area, or the palisade elements are
loosened, grow outward a short distance, and then split. Likewise
the gill plates are split apart to different depths due to the tensions
set up by these differences in the rate of growth at different regions
of the sporophore. The most rapidly growing region is at the margin
of the pileus, and, while growth may and actually does take place
throughout the sporophore, it decreases in rapidity from the periphery
to the place of attachment of the stipe. Thus the gill plates are
longer at the periphery, but the difference between the rates of growth
of the hymenium and subhymenial and tramal layers is just as pro-
nounced. As a result the hymenial margins are incurved more at the
margin of the pileus and the gill plates are gradually drawn apart.

In some young sporophores the two hymenial margins are separated
by a considerable layer of sterile surface which is level with the hyme-
nium (fig. 1, pi. 60). In this case there is not even a resemblance to
gills.

From the evidence at hand it seems that the so-called "gills" of
Schizophylluni commune Fries are such by analogy only, being act-
ually two adjacent edges of hymenial areas which arise together, but
which become continuous with and are homologous to the margin of
the pileus. They increase the area of the hymenium as do gills, and,
when in a moist condition, look much like typical gills ; but each plate
is independent of the other from the first, increasing in size by mar-
ginal growth.

1922] Essig: Morphology of Schizophyllum commune Fries 463

3. TAXONOMIC INTEEPEETATION OF THE STRUCTURE AND
DEVELOPMENT OF THE SPOROPHORES

It was early observed that while Schizophyllum was presumed to
possess gills, the structures were unique among the members of the
Agaricaceae. Consequently considerable difficulty has been experi-
enced among systematists in associating this genus with the other
members of the family from which it is so distinct.

Fayod (1889) believed it to be precisely like Pamis, except that
in Panus the gills were entire. He classified them under the tribe
Panoides, and noted a similarity in the geographical distribution of
the two genera. Hennings (1898), following Saccardo (1887-1895),
placed Schizophyllum in the tribe Schizophylleae with Rhacophyllus,
Oudemansiella, and Pterophyllus, but stated that the other genera do
not seem to belong to this group. Murrill (1915) put it in the tribe
Agariceae and subtribe Lepiotanae along with Marasmius, Lepiota, and
other white-spored members of the Agaricaceae.

All these writers have based their classification upon the assump-
tion that the hymenium in Schizophyllum is borne upon the surface
of lamellae. Since this is not the case — for the hymenophore while
quite complex in structure bears a smooth h^Tnenium — the fungus
should be placed in the family Thellephoraceae. In this family it
resembles Cyphella in the organization of the sporophores, for, in this
genus, according to De Bary (1887), the hymenium lines the inner
and lower surfaces of a funnel-shaped sporophore, the stipe being
attached to the opposite side of the pileus from the hymenium (fig. 16,
pi. 52). The early stages in the development of Schizophyllum com-
mune and Stereum hirmitum,, likewise, have much in common. In the
later stages, however, the hymenium of Stereum hirsutum remains
entire, and bears no resemblance to the much divided hymenium
of Schizophyllum commune. Only in the genus Cladoderris is there
anything comparable to the hymenial plates of Schizophyllum. Clado-
derns is somewhat similar to Stereum., but differs in possessing radiat-
ing, branched ribs upon the hymenial surface. Some species of Clado-
derris have the pileus margin much incised. The representatives of
the genus Cladoderris are chiefly tropical, and are known to the writer
only through descriptions and illustrations. From these it seems that
there is only a difference in degree between the splitting of the pileus
margin in sporophores of Cladoderris infundihuUformis Fries (cf.
Hennings, 1898) and the marginal division in fruit bodies of Schizo-
phyllum commune.

464

University of California Publications in Botany [Vol. 7

V. ECONOMIC ASPECTS

1. GEOGRAPHICAL DISTRIBUTION

In a discussion of the economic importance of Schizophijllum it
seems well to go into detail concerning its distribution throughout
the world, as any consideration of the amount of damage done must
take into account both its distribution and its abundance in any dis-
trict. Fortunately a great number of statements have been published
concerning its collection in various places. In the table below are
given by continents the countries or regions in which Schizophyllum
has been collected or reported, the authority, and the date of publi-
cation of the article. The table is representative rather than exhaus-
tive, as only one reference to a locality has been included.

TABLE OP^ GEOGRAPHICAL DISTRIBUTION

Continent

Place

Authority

Date

Eurasia

England

Cheesman, W. N.

1904

Scotland

Paterson, R. H.

1877

Sweden

Linnaeus, C.

1753

Germany

Hennings, P.

1898

France

Gueguen, F.

1901

Italy

Archangeli, G.

1887

Central Asia

Sorokine, N.

1890

China

Roumeguere, C.

1879

Ceylon

Berkeley, M. J., and Broome, C. E.

1871

Africa

Tripoli

Baroni, E.

1892

Abyssinia

Saccardo, P. A.

1891

Cape of Good Hope

Berkeley, M. J.

1876

Africa

Hennings, P.

1891

North America Canada

Dearness, J.

1896

Eastern United States

Atkinson, G. F.

1901

Middle Western U. S.

Heald, F. D.

1906

Oregon

Griffin, F. L.

1911

California

Smith, R. E., and E.

H.

1911

Mexico

Patouillard, N.

1887

West Indies

Massee, G.

1892

South America Brazil

Averna-Sacca, R.

1916

Australia

Australia

MacAlpine, D.

1902

New Zealand

Buchanan, J.

1874

The sporophores of the Schizophyllum in the field are usually small
and inconspicuous. "When found they are usually in large numbers
within a small area on a log or tree. Only an occasional tree or log
displays them in this region (California). Heald (1906) reported
that every tree of a small orchard of cherries in Nebraska was infected
with the fungus, but this seems to be an extreme case.

1922] Essig: MorpJiology of Schizophyllum commune Fries 465

2. LIST OF HOST PLANTS

A large number of plants have been mentioned in scientific liter-
ature as hosts of the sporophores of Schizophyllum. It is not clear in
most of these references whether the specimens were found upon living
or dead plants. The distinction is of some importance in the consid-
eration of the economic aspects of the fungus. The information in the
following table has been obtained from publications, from the spoken
word of collectors whom the writer has been fortunate enough to meet,
from specimens in the Herbarium of the University of California, and
from observations in the field. Dates are given for references to
publications only.

TABLE OF HOST PLANTS

Family

Name

Authority

Date

Pinaceae

Yellow Pine

(Pinus ponderosa)

Stilhnger, C. R.

Western Hemlock

Stillinger, C. R.

(Tsuga heterophjlla)

Gramineae

Sugar Cane

(Saccharum officinarum)

Ray, J.

1896

Bamboo {Bamhusa sp.)

Home, W. T.

Palmaceae

Royal Palm (Oreodoxa regia)

Home, W. T.

Juglandaceae

English Walnut (Juglans sp.)

Smith, R. E., and E. H.

1911

Hickory {Hicoria sp.)

Dearness, J.

1S96

Betulaceae

Alder {Alnus sp.)

Linnaeus, C.

1783

Birch {Betula sp.)

Adams, J. F.

.1918

Fagaceae

Beech (Fagus sp.)

Rumbold, C.

1910

Oak (Quercus serrata)

Roumeguere, C.

1879

Chestnut (Castanea sp.)

Stevens, F. L.

1913

Moraceae

Mulberry (Morus sp.)

Prillieux and Delacroix

1893

Lauraceae

California Baytree

( Umhelhdaria calif ornica)

Brown, V. S.

Rutaceae

Oranges and Lemons

(Citrus sp.)

Smith, R. E., and E. H.

1911

Tiliaceae

Linden (Tilia sp.)

Hennings, P.

1898

Aceraceae

Maple (Acer sp.)

Hasselbring, H.

1907

Hippoeastanaceae

Horse Chestnut

(Aesculus Hippocastanum)

Gueguen, F.

1901

Rosaceae

Apple (Pyrus malus)

Fulton, H. R.

1912

Pear (Pyrus communis)

Baroni, E.

1892

Cherry (Primus cerasus)

Griffin, F. L.

1911

Peach (Prunus persica)

Camp, A. F.

Almond (Prunus comrtiunis)

Kellogg, E. S.

Leguminosae

Pterocarpus indicus

Kew Bull. Misc. Inf.

1910

Acacia (Acacia sp.)

Seen in the field

Hardy Catalpa

Stevens, N. E.

1912

(Catalpa speciosa)

Rubiaceae

Coffee (Coffea sp.)

Averna-Sacca, R.

1916

466 Uwiverdty of Calif ornia Puhlicatio-ns in Botany [Vol.7

3. EXTREME HARDINESS OF THE FUNGUS

SchizophyUum is able to persist under very adverse circumstances.
Its unus\ial vitality is displayed in three ways: (1) the sporophores
are able to endure long periods of drought; (2) the mycelium can
grow upon almost any moist organic substance; and (3) the sporo-
phores possess the ability to regenerate lost parts.

BuUer in 1909 called attention to the long period over which the
sporophores can retain their vitality. He stated that "whilst in the
dried condition a fruit body can retain its vitality for at least two
years, and, with intermittent revivals, for at least three years. ' ' Later
(1912) he and Cameron found that the fruit bodies could endure
sudden changes of temperature, suspension in a vacuum, extreme cold,
or a long period in darkness. In this respect they resemble certain
seeds and mold spores.

A mycelium produced either from spores or pieces of sporophores
will grow upon a whole series of substances. Some of the materials
upon which the hyphae grow well are such starchy media as potato
tubers, corn meal, rice, "Cream of Wheat," and lima beans; sugary
media such as beets, prune juice, and grapes; upon agar and gelatin
nutritive media ; and upon dung, a wood decoction, or dead leaves.
Kellogg (1915) grew the fungus from spore to spore, or through all
of its life history, upon sterilized potato plugs in glass flasks, showing
that Schizophyllum can exist in an entirely saprophytic condition.
Earlier Rumbold (1910) had produced sporophores on bread from
spore cultures, but, although these bore basidia on a definite hymenium,
no spores developed.

Experiments have been carried on to determine to what extent the
sporophores can regenerate lost parts. The fruit bodies studied were
grown upon blocks of Acacia wood kept in moist chambers in the
laboratory. In one case about one-half of the pileus at the distal ends
of mature sporophores was removed by cutting with a sharp knife.
Some of the specimens were left in the original position, and others
were inverted by reversing the position of the blocks of wood upon
which they were growing. Where the mutilated sporophores were
left in fjosition growth ceased at the cut edge, but continued in a
normal manner, though rather slowly, at the pileus margins to the
side. Most of the inverted specimens ceased growth altogether. In
one case, however, in a few days the margin at one side began to turn
and grow outward in a horizontal direction with the hymenium facing

1922] Essig: MorpJiology of Schizophylliim cmnmune Fries 467

downward. At the end of 17 days a normal sporophore about 1 cm.
in diameter had developed (fig. 2, pi. 60).

In another experiment the entire hymenophore was removed by
cutting across the stipe at the distal end. Specimens left in the orig-
inal position produced new hymenophores in one of two methods. If
the stipe was small at the cut end, only one new sporophore, as a rule,
developed by the growth of hyphae out through the cut end of the
stipe (fig. 3, pi. 60). This developed in the usual way. If the cut
end of the stipe was of considerable area, several small sporophores
developed. These grew in the usual way except that the hymenial
plates arose in position with respect to the old stipe and not as though
the separate sporophores were distinct individuals (fig. 18^, pi. 52).
The sporophores more advantageously placed, that is, at the upper
edge of the stipe, grew more rapidly and became much larger than
those at the sides. The sporophores might be cut away to within a
millimeter of the base of the stipe and still a new sporophore would
develop upon the cut end. The specimens used were mature and were
shedding spores, but were comparatively young. Thus the sporo-
phores have, at least while still young and fresh, the ability to regen-
erate practically the entire body.

Stipes with the entire hymenophore removed and in an inverted
position in all cases produced either one or several small sporophores
upon the cut end, but growth soon ceased. Sections through these
specimens showed that they had stopped growing either in the
"peziza" stage or after one or two hymenial plates had been formed
(figs. 19-20, pi. 52). The inability" of the sporophores to develop
further in an inverted position is doubtless due to their lack of power
to change the polarity of the different parts with respect to the
reaction to the force of gravity. That gravity is the form-stimulus
was clearly demonstrated by Hasselbring (1907).

4. EELATION OF THE MYCELIUM TO CELLS OF DEAD WOOD

Upon sectioning dead wood it is found that there is an unexpected
paucity of mycelium in the tissues infected by Schizophyllum. Wood
brought in from the field in a dry condition covered with the sporo-
phores may be sectioned and fail to display any mycelium in a large
percentage of the sections. In some, however, a few hyphae can be
seen.

The hyphae of this fungus are shown by sections to be confined
almost entirely to the tracheae of the wood (pi. 61). In some ducts

468 University of California Publications in Botany [Vol.7

there may be only one or two, but in others the lumen may be almost
filled. In all cases the mycelium varies considerably in size, the walls
are thin, and branching is infrequent. The lateral tubercles are
present upon tiie walls of some of the hyphae.

Wherever the mycelium is found in wood in earlier stages of decay
there is present a series of small, globular masses of a brown exudate.
The mycelium of Sdiizophyllum growing upon artificial media pro-
duces a like substance, so that found in the wood is probably pro-
duced by the hyphae. In certain regions the droplets are so numerous
that the wood is discolored. They account for the black or dark brown
layers often seen near the edge of the decayed areas. In regions of
advanced decaj^ they have entirely disappeared.

Cross-sections through the limb of a living tree which was infested
with the fungus in a narrow area along one side of the limb showed
that the infected area extended in a radial direction to the center of
the limb. Some pieces of this limb were placed in moist chambers.
In a few days tufts of hyphae grew out of the wood at the edges of
the infected area near the living wood and only a few scattering threads
could be seen in the central part of the discolored tissues. Thin sec-
tions also disclose the fact that the greater part of the vegetative
mycelium is near the living wood in partially killed limbs or trunks
of trees.

The decay is marked at first by a darkening of the tissues. There
are dark brown or black layers near the edge of the darkened areas.
At later stages delignification sets in, and the decayed areas become
straw-colored. The cell walls become softened, but retain their struc-
ture for a long period. The mycelium of Sdiizophyllum is frequently
found in areas of advanced decay along with the hyphae of other
fungi, and it is difficult to determine how much of the decay is due to
the work of ScJiizophyllum commune alone.

5. GROWTH UPON FRESH WOOD AND LIVING TISSUES

Freshly cut pieces of Acacia wood were placed in moist chambers
and spores of Schizophyllum were planted upon different tissues.
Acacia wood was chosen because sporophores are found in abundance
upon tlic df-ad wood of this tree. The pieces of wood were kept moist
enough to cause the spores to germinate. The experiment was carried
on for two months. At the end of that time it was found that the

1922] Essig: Morphology of Schizophyllum commune Fries 469

hyphae had not penetrated through the fresh, uninjured bark, or cor-
tical tissues, or through the wood tissues in a lateral direction, but had
grown through the wood in the direction of the tracheae.

Attempts to prove the parasitism of this organism were made by
Gueguen (1901) and Fulton (1912) with negative results. Rumbold
(1910) stated that Tuzson (1905), (whose paper is not available) grew
the fungus upon fresh (frais) beech wood. Kellogg (1915) was not
able to demonstrate the mycelium of Schizophyllum in the inoculations
made upon fruit trees.

In my inoculation experiments young fruit trees of apple, pear,
and plum were used. Inoculations were made from agar plates of
pure cultures of the mycelium. The limbs of these trees were either
split through the center or cut into from the surface to varying depths.
A sterile knife was used in the incisions. The mycelium Math sub-
stratum was transferred to the cut or slit surface and the wound was
tied up with string and covered with waxed paper, a layer of wet
absorbent cotton, and another layer of waxed paper. The purpose of
the wet cotton and w^axed paper was to prevent the drying out of the
exposed surfaces. At intervals of one week after the time of inocu-
lation certain limbs were removed, examined, and sections made to
determine if there had been any growth of the mycelium into the
living tissues. In most cases no trace of the mycelium could be found
in the wood. In two or three branches there was a darkening of the
surface exposed to the mycelium, and in the vessels the typical exudate
which is produced by the mycelium, but so little mycelium was found
in the ducts that the growth could not be identified, with certainty,
as Schizophyllum. In only one branch was there an unmistakable
infection. This was in a limb of a plum tree cut off three months
after inoculation. The mycelium had penetrated to a maximum depth
of 3 mm. and the infected area was about 4 cm. long and 1 cm. in
width. Many hyphae were present in the vessels, and a few could be
seen in the medullary ray cells. There was the characteristic darkened
layer at the edge of the infection about 0.5 mm. thick. From the
results of previous investigations and these experiments it is evident
that the living woody tissues can be penetrated and killed by the
mycelium of Schizophyllum, but that this process takes place slowly
and with difficulty.

470 University of California Puhlications in Botany [Vol. 7

6. METHODS OF INFECTION UNDER NATURAL CONDITIONS

There have been many expressions of opinion in published papers
concerning the manner in which living trees become infected with tlie
mycelium of Schizophyllum. Infection takes place in three more or
less distinct ways: (1) by entering through surfaces exposed by
mechanical injury, (2) by attacking parts of trees weakened through
certain physiological causes, and (3) by gaining admission through
tissues first injured or killed by other organisms.

As regards mechanical injury, in Stevens and Hall (1910) it is
stated that "apparently this disease starts in roots injured by tools
during cultivation." Professor Home of the University of California
has observed an infection beneath an apple tree graft (fig. 1, pi. 58)
that had not been properly sealed with wax. An infection at a crotch
split in a peach tree (fig. 2, pi. 58) was reported by A. F. Camp, a
student in the University of California. Any woody part exposed
by injury forms a possible place of entrance for the fungus.

"Weakening of the trees by excess water, or lack of proper drain-
age, was decided by Guegen (1901) to be a contributory cause of
infection in horse chestnut trees. Stone (1910) found that sun scald
and scorch of maple trees was followed by Schizophyllum and other
fungi. The writer has seen the sunburned parts of California Bay
trees covered with sporophores. It is evident that trees weakened by
certain physiological agents fall prey to this fungus, and probably
any loss of vitality on the part of the tree makes it susceptible to the
attacks of Schizophyllum.

Wilson (1912) found that the sporophores issued through the
burrows made by the shot hole borer {Xylehorus dispar Fabricus).
Griffin (1911) stated that cherry trees weakened by bacterial gum-
mosis are frequently attacked and killed by Schizophyllum commune.

No evidence has been brought forward in available literature to
show that Schizophyllum is able to infect healthy trees, or those not
injured or weakened in some way. It seems likely that infection can
be prevented by using care in cultivation, by painting wounds made
in pruning, by preventing crotch splitting, by protecting the trees from
sunburning, and by keeping them free from other diseases.

1922] Essig: Morphology of Schizophylluni commaine Fries 471

7. ASSOCIATION WITH OTHER WOOD DECAY FUNGI

Fulton (1912) found that in an apple collar rot which he described
Schizophyllum commune was present along with two other organisms.

Stone (1910) found in sun scald and scorch of maples that it was
followed first by a canker fungus [Nectna dnnaharina) , and then by
ScJiizophylUim and Polystictiis.

In the case of the apple graft mentioned above, the writer took
the branch, sawed it into sections, and placed some of the sections in
a moist chamber. From the decayed area a mass of mycelium appeared
which was not of Schizophyllum, but as it has not yet produced any
fruit bodies it cannot be identified.

Observations show that the fruit bodies of Schizophyllum. are pro-
duced in a comparatively short time after inoculation has taken place.
I have noted several times in the field that this fungus is the first to
appear upon uprooted trees. Later, sporophores of Polystictus,
Polyporus, Tremella, Hydnum, and other fungi are produced. ]\Iost
of the pieces of wood I have collected in the field, which have only
sporophores of Schizophyllum upon them, when placed in moist cham-
bers long enough will produce sporophores of Hydnum or Polystictus,
or both, long after a great number of fresh Schizophyllum fruit bodies
have been formed. Since the hyphae of these other fungi were in
the wood along with that of Schizophyllum, it is apparent that the
latter fungus develops sporophores in a much shorter time than the
others present. Schizophyllum, forming fruit bodies first upon a dis-
eased tree, is naturally accused of being the parasite causing the dam-
age. The writer believes that much or even most of the injury to trees
attributed to this fungus is actually caused by the fungi so often
associated with it, such as Polystictus versicolor, which is beyond doubt
a parasite.'

2 Proof of the parasitism of P. versicolor was given by W. W. Thomas in 1916
at the University of California in a thesis submitted for a Master's degree.
Mr. Thomas inoculated living trees with positive results as regards infection.

472 University of California Fnhlications in Botany [Vol. 7

VI. SUMMARY AND CONCLUSION

In summarizing tlio morphology and development of Schizophyllum
it may be said that:

1. The sporophores vary greatly as to form and shape.

2. The segments of the mycelium and sporophores and spores are
regularly binucleate.

3. The fungus develops its sporophores as does no other member
of the Hymenorayeetes so far studied, the hymenium primordium aris-
ing in an apical cavity.

4. The "gills" arise upon the surface of an apical depression due
to tensions set up by unequal rates of growth.

5. The "lamellae" are such by analogy only, being the edges of
smooth hj'menial areas, and therefore Schizophyllum belongs in the
family Thellephoraceae.

As regards the economic aspects of Schizophyllum, it has been
shown that :

1. Members of this genus are found throughout the tropical and
temperate zones of the world.

2. They live upon a great number of woody plants, both in the
Dicotyledonae, Monocotyledonae, and Gymnospermae.

3. The fungus possesses unusual vitality.

4. The mycelium is found only in small amount in infected wood.

5. It can grow upon fresh wood, and, under very favorable condi-
tions, living wood.

6. Natural infection takes place through some injured or Aveakened
part of the tree.

7. The fungus is usually associated with other parasitic fungi,
which probably do most of the damage attributed to Schizophyllum,
but escape attention due to the longer time necessary for them to
produce fruit bodies.

1922] Essig: MorpJiology of Schizophyllum commune Fries 473

VII. ACKNOWLEDGMENTS

For helpful advice and criticism in the preparation of this paper
the writer wishes to thank and give credit to Dr. W. A. Setchell, Dr.
T. H. Goodspeed, and Professor W. T. Home. Mr. C. R. Stillinger
of the Bureau of Plant Industry, United States Department of Agri-
culture, has given valuable information concerning certain phases of
the work. In the collection of specimens aid has been given by Mr.
A. F. Camp and others.

LITERATURE CITED

Adams, J. F.

1918. Origin and development of the lamellae in Schizophyllum commune.
Memoirs of the Torrey Botanical Club, vol. 17, pp. 326-333, pi. 9.

Archangeli, G.

1887. Sopra alcune erittogame raccolte nel Piceno e nell 'Abruzzo. P. V.
Pisa, vol. 5, pp. 243-246.

Atkinson, G. F.

1901. Studies of American Fungi. Mushrooms, edible, poisonous, etc.

Ithaca.
1916. Origin and development of the lamellae in Coprinus. Botanical

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Averna-Sacca, E.

1916. Fungous diseases of coffee. Boletim de Agricultura (Sao Paulo,
Brazil), ser. 17, no. 10, pp. 790-840, figs. 49; no. 11, pp. 878-922,
fig. 46.
Baroni, E.

1892. Cryptogams from Tripoli. BoUettino della Societa Botaniea Italiana.
Florence. Pp. 239-243.

Berkeley, M. ,1.

1874. Enumeration of the fungi collected during the expedition of H. M. S.

"Challenger," Feb. -Aug., 1873. Journal of the Linnean Society,

pp. 350-354.

1876. Fungi collected at Cape of Good Hope, etc. Journal of Botany,
pp. 173-176.

Berkeley, M. J., & Broome, C. E.

1871. Enumeration of the fungi of Ceylon. Journal of the Linnean Society,
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Brefeld, O.

1889. Untersuchungen aus dem Gesammtgebiete der Mykologie, vol. 8, pp.
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Buchanan, J.

1874. Notes on the flora of the Province of Wellington, etc. Transactions
and Proceedings of the New Zealand Institute, vol. 6, pp. 210-235.

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BULLER, A. H. R.

1909. Eesearches on fungi. Loudon.

BuLLER, A. H. R., & Cameron, A. T.

1912. On the temporary suspension of vitality in the fruit bodies of certain
HynienoniyfCtes. Proceedings and Transactions of the Royal So-
ciety of Canada, ser. 3, vol. 6, sec. IV, pp. 7:3-78.

Cheesman, W. N.

1901. Schizophyllum commune in East Yorkshire. Naturalist, vol. l."], pp.
283-294.

Cooke, M. C.

1871. Handbook of British fungi. Vol. 1. London.

De Bary, A.

1887. Morphology and biology of the fungi, mycetozoa, and bacteria. Eng-
lish ed., London.

Dearxess, J.

1896. Ellis, J. B., & Everhard, B. M. Fungi Coluniljiaiii, vol. 9, no. 803.
(Exsicc.)

Dillenius, J. J.

1719. Catalogus plantarum sponte circa Gissam nascentium. Frankfort. (Cf.
Greville (1824) and p. 70 in Pritzel's (1851) "Thesaurus.")

Fayod, M. V.

1889. Prodrome d 'une histoire naturelle des Agaricines. Annales des Sci-
ences Naturelles, ser. 7, Botanique, vol. 9, pp. 331-333.
Fries, E.

1821. Systenia mycologicum.

Fulton, H. R.

1912. Apple collar rot. Pennsylvania Station Report, pp. 251-253.

Gardner, N. L.

1917. A freezing device for the rotary microtome. Botanical Gazette, vol.
63, pp. 236-239, fig. 1.

Greville, R. K

1824. Scottish cryptogamic flora, vol. 2, fasc. 61.
Griffin, F. L.

1911. A bacterial gummosis of cherries. Abstract in Science, n. ser., vol. 34,
no. 879, pp. 615, 616.
Gueguen, F.

1901. Le Schizophyllum commune, parasite du Marronier d ' Inde. Bulletin

de la Societe Mycologique de France, vol. 17, pp. 283-298.
Hard, :M. E.

1908. The mushroom, edible and otherwise. Columbus, Ohio.
Harper, R. A.

1902. Binucleate cells in certain Hymenomycetes. Botanical Gazette, vol.

33, no. 1, pp. 1-25, pi. 1.

Has.selbring, H.

1907. Gravity as a form-stimulus in fungi. Botanical Gazette, vol. 43, pp.
251-258, figs. 1-3.
Heald, F. D.

1906. New and little known plant diseases in Nebraska. Abstract in Sci-
ence, n. ser., vol. 23, no. 590, p. 624.

1922] Essig: Morphology of Schizophyllum commune Fries 475

Hennings, p.

1891. Fungi africani. Eng. Jahr., vol. 14, pp. 337-373.

1898. Hymenomycetineae. Engler and Prantl, Die naturliehen Pflanzenfa-
milien, vol. li**, p. 221.

Hoffman, H.

1860. Beitrage zur Entwdckelungsgeschichte und Anatomic der Agaricinen.
Botanisehe Zeitung, vol. 18, pp. 389-397, pi. 13.

Kellogg, E. S.

1915. Studies on SchizophyUum. (Unpublished manuscript in Library of
Plant Pathology Division, University of California.)

Linnaeus, C.

1753. Species plantarum.

MacAlpine, D.

1902. Fungous diseases of stone-fruit trees in Australia and their treatment.
Melbourne.
Maire, E.

1900. Sur la cytologic des Hymenomycetes. Comptes Eendus, Soc. de
Biologie, Paris, vol. 131, p. 121.
Massee, G.

1892. Some West Indian fungi. Journal of Botany, vol. 30, pp. 161-164,

pi. 321-323.

Morgan, A. P.

1890. Mycological observations. LI. Botanical Gazette, vol. 15, pp. 84-86.

MURRILL, W. A.

1915. Agaricaceae. North American Flora, vol. 9, no. 4, pp. 237, 238.

Paterson, E. H.

1877. Exotic fungi of Scotland. Grevillea, vol. 5, p. 112.

Patouillard, N.

1887. Note sur quelques champignons de I'herbier du Museum d'histoire
naturelle de Paris. Journal de Botanique, vol. 1, pp. 169-171.

Prillieux, E., & Delacroix, G.

1893. Diseases of the Mulberry. Bulletin du Ministre de I'Agricole, vol. 12,

no. 5, pp. 452-472.

Pritzel, G. a.

1851. Thesaurus literaturae botanieae omnium gentium inde a rerum botani-
carum initiis ad nostra usque tempora, quindecim millia operum
recensens. Leipzig.
Eay, J.

1896. On the diseases of the sugar cane. Bulletin de la Societe Mycologique
de France, pp. 139-143.

Eosenvinge, K.

1886. Sur les noyaux des Hymenomycetes. Annales des Sciences Naturelles,
ser. 7, Botanique, vol. 3, pp. 75-93, pi. 1.

Eoumeguere, C.

1879. Chronique mycologique. Eevue Mycologique, vol. 1, pp. 145, 146,
150-154, pis. 2-4.

EUMBOLD, C.

1910. Les champignons destructeurs du bois. Annales de la Science Agro-
nomique, ser. 3, vol. 5, no. 6, pp. 402-404, fig. 25.

476 V nwcrsii II of California Puhlications in Botany [Vol.7

Saccardo, p. a.

1887. Sylloge fungorum omnium hucusque cognitorum, vol. 5, pp. 654-656.
1891. Hid., vol. 9, p. 81.
1895. Ihid., vol. 11, p. 42.
1899. Ihid., vol. 14, p. 123.
litll. Ihid., vol. 20, pp. 750, 751.

SCHROETER, J.

1889. Die Pilze Sclilesiens, vol. 3, fasc. 553.

Smith, E. E. & E. H.

1911. California plant diseases. California Agricultural Experiment Sta-

tion, Bulletin no. 218, pp. 1088, 1089, 1140, 1173, figs. 16, 65.

Smith, W. G.

1884. Fungus on ensilage. Garden Chronicle II, vol. 22, p. 405.

SOROKINE, N.

1889--1890. Cryptogamique flora de central Asia. Eevue Mycologique, vols.
11, 12.

Stevens, F. L.

1913. The fungi which cause plant disease. New York.

Stevens, F. L., & Hall.

1910. Diseases of economic plants. New York.
Stevens, N. E.

1912. Wood rots of the hardy catalpa. Phytopathology, vol. 2, no. 3, pp.

114-119.
Stone, G. E.

1910. Shade tree troubles. Eeport of the Massachusetts Agricultural Ex-
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Tuzsox, J.

1905. Anatomische und mykologische Untersuchungen uber die Zersetzung

und Konservierung des Eotbuchenholzes. Berlin.
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1893. On the nuclear divisions in the Hymenomycetes. Annals of Botany,

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Wilson, H. F.

1913. The shot hole borer of the Northwest; or the pear blight beetle of

the East. Biennial Crop Pest and Horticultural Eeport, 1911-1912,
Oregon Agricultural College Experiment Station, pp. 97-107, fig. 1.
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1884. Die Pilze Deutschlands, Oesterreichs und der Schweiz. Eabenhorst,
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Bulletin of Miscellaneous Information, no. 3, pp. 95, 96.

EXPLANATION OF PLATES

PLATE 51

Fig. 1. Group of sporophores in a dry condition, viewed from the lower
side. 2 diameters.

Fig. 2. Group of sporophores in a moist condition, as seen from below.
Some of the smaller specimens at the base of the group have been cut away.
iy2 diameters.

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UNIV. CALIF. PUBL. BOT, VOL. 7

[ ESSIG ] PLATE 51

2

PLATE 52

Fig. 1. Form of sporophores growing upon an under surface, a, side view
of fruit bodies iu position, b, the hyinenium of a typical fruit body viewed
from below, showing the gill plates radiating outward from a region near the
center of the hymenium. ^A natural size.

Fig. 2. Form of fruit bodies growing upon a vertical surface, a, three
sporophores in position, i, arrangement of the lamellar plates in a sporophore
grown upon a vertical substratum. % natural size.

Fig. 3. Form of fruit bodies growing on an upper surface, a, position
assumed by sporophores upon the upper surface of a limb, b, typical arrange-
ment of the lamellar plates. % natural size.

Figs. 4-15. Views of young actively growing fruit bodies from the under
side to show the origin and placement of the lamellar plates or ridges. All 5
diameters. Fig. 4, a sporophore with a smooth hymenium primordium just prior
to the formation of lamellar ridges. Fig. 5, a fruit body with a single lamellar
ridge. Fig. 6, a single lamellar ridge with the sides so far separated by sterile
hyphae as to become merely edges of the hymenium primordium. Fig. 7, a
specimen with four newly formed lamellar ridges. Figs. 8-14, six sporophores
with some isolated ridges and some extending to the edge of the pileus. Fig. 15,
a lateral sporophore with the lamellar ridges widening into sterile areas towards
the base.

Fig. 16. Sporophore of CypheJIa Urhani. 3 diameters. (After Engler and
Prantl.)

Figs. 17-18. Eegeneration in position, a, a small sporophore showing with
a broken line where the hymenophore was removed, b, face view of the hymeno-
phore removed. 18c, side view of the sporophore after seven days, d, face view
of the same, showing five small sporophores formed upon the cut end of the old
stipe. All lYo diameters.

Fig. 19. Regeneration inverted, o, a small fruit body shoAving where the
end was cut away, d, a median section of the same thirty days later. Only one
pair of lamellar plates was formed. 1% diameters.

Fig. 20. Eegeneration inverted, c, a small sporophore showing where the
end was removed, d, the same a week after the apex had been cut away, show-
ing three very small sporophores formed upon the cut end of the inverted stipe.
These developed no further. P/^ diameters.

Fig. 21. A transverse section through a sporophore when in a dry state,
showing the revolute lamellar plates, and their arrangement. 5 diameters.

Fig. 22. A transverse section through the same sporophore represented in
fig. 21, but drawn while the sporophore was in a moist condition. 5 diameters.

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UNIV. CALIF. PUBL, BOT. VOL. 7

[ ESSIG ] PLATE 52

t—

U^^JS.

V

19

>

20

22

PLATE 53

A sporophore in a semi-moist condition, as seen from below. The hymenium
is shown to be divided into four areas, three of which are much smaller than
the fourth. Several newly formed isolated pairs of lamellar plates can be seen,
as well as three very small ridges near the center of the hymenophore which
arose in an early development stage but remained isolated. 10 diameters.

[482]

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[ ESSIG ] PLATE 53

PLATE 54

Fig. 1. Part of a typical vegetative hypha to show the structural charac-
ters. Clamp connections are shown at each septum. The nuclei are near the
center of each cell. 1500 diameters.

Fig. 2. Portion of a vegetative hypha covered with lateral projections.
1500 diameters.

Fig. 3. Hyphae which form the hairy covering of the pileus. They are
usually unicellular, the nuclei being near the base of the cell. In this drawing
a portion of each cell 1.4 mm. long was left out. This would represent a distance
of about 210 cm. o6 the drawing paper. 1500 diameters.

Fig. 4. Portion of a longitudinal section through the flesh of a very young
sporophore to show the hyphal structure. 1500 diameters.

Fig. 5. Same as in fig. 4, except that the section was taken from an old
sporophore. . 1500 diameters.

Fig. 6. Portion of the hymenium, showing one mature and one immature
basidium. 1500 diameters.

Fig. 7. Six typical spores, stained in safranin and gentian violet to show
the nuclei. 2500 diameters.

Fig. 8. Germinating spores in various stages of development after being
in distilled water for 48 hours. Stained in safranin. 1500 diameters.

[484]

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[ ESSIG ] PLATE 54

8

PLATE 55

Fig. 1. Longitiuliiial inediaii section through a young fruit body which is
Ijractically undifferentiated. There is a region near the apex which stains
more deeply and is the seat of later developments. 25 diameters.

Fig. 2. Similar to fig. 1, but a later stage, showing the first appearance
of the apical cavity. 25 diameters.

Fig. 3. The apical cavity is still further developed. 25 diameters.

Fig. 4. Beginning of the palisade layer. 25 diameters.

Fig. 5. Further development of the palisade layer. 25 diameters.

Fig. C. Maximum growth of the palisade layer in a plane surface. 25 diam-
eters.

Fig. 7. The palisade layer (hymenium priniordiuni) produced into a eon-
cave surface by the outward growth of the edges Of the pileus. 25 diameters.

Fig. 8. Median longitudinal section through a small sporophore formed
upon the cut end of a stipe. 18 diameters.

[483]

U^ilV. CALIF. PUBL. BOX. VOL. 7

[ ES.SIG ] PLATE 55

4

8

PLATE 56

Fig. 1. A piece of wood removed from the surface of an UmhcUularia log,
showing a number of very small sporophores in the earliest stage of development.
Surface and side views, 2% diameters.

Fig. 2. Shapes assumed by very young undifferentiated fruit bodies. 2^^
diameters.

Fig. 3. A group of young fruit bodies in position upon a piece of TJmhel-
lularia bark. Five of the specimens are in the "apical cavity" stage, one is
still undifferentiated, and another (at the extreme right) has already developed
lamellar plates. 3% diameters.

Fig. 4. Tangential section of Acacia wood to show the mycelium of Schizo-
phyUum commune in a duet. About 125 diameters.

Fig. 5. Eadial section of UmheUularia wood to show the hyphae of Schizo-
phyllum in the ducts. About 125 diameters.

Fig. 6. Splitting of the palisade layer previous to the formation of a
lamellar ridge. 250 diameters.

Fig. 7. Outward growth of the palisade layer at the sides of a split. 250
diameters.

Fig. 8. Loosening and outward growth of the elements of the palisade
layer to form a lamellar ridge. 250 diameters.

Fig. 9. Cross-section of the palisade layer to show a very early stage in
the origin of a lamellar ridge. 250 diameters.

[488]

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[ ESSIG ] PLATE 56

oAUUUD'o

opoO

ZDO{

8

PLATE 57

Fig. 1. A transverse section across a lamellar ridge in a very early stage
of development. 448 diameters.

Fig. 2. A cross-section through a lamellar ridge divided down in the middle
to form two small lamellar plates. 448 diameters.

Fig. 3. Another lamellar ridge in a stage of development similar to that
shown in fig. 2. 448 diameters.

Fig. 4. A lamellar ridge whose halves are split apart deeply into the pileus
flesh. 448 diameters.

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[ ESSIG ] PLATE 57

3

PLATE 58

Fig. 1. Sections of apple limbs with the wood rotted by fuugi penetrating
through an improperly sealed graft. Sporophores of Schisophyllum appeared
upon the bark of the tree beneath the graft about a year after the grafting
was done. See the text for further explanation. Yz natural size.

Fig. 2. Sporophores of ScMzophyllum commune in a dry condition upon the
bark of a living peach tree at the side of a crotch split. The lower end of the
split can be seen on the right side of the trunk. % natural size.

[492]

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[ ESSIG ] PLATE 58

„iffil

PLATE 59

Fig. 1. A median longitudinal section through a very small undifferentiated
fruit body. See fig. 1, pi. 54. 55 diameters.

Fig. 2. A median longitudinal section of a sporophore showing the breaking
in of the apical cavity. 55 diameters.

Fig. 3. Photomicrograph of the section dra\\Ti in fig. 6, pi. 54. 55 diameters.

Fig. 4. A median longitudinal section through a fruit body with the apical
cavity fully developed. 55 diameters.

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[ ESSIG ] PLATE 59

3

4

PLATE 60

Fig. 1. A widened lamellar ridge in cross-section, showing the sterile area
between the edges of the hymenium. 75 diameters.

Fig. 2. A sporophore developed from a single lobe of a mutilated inverted
specimen. For a further description see the text. 2 diameters.

Fig. 3. A median longitudinal section through a regenerated sporophore.
See text for further explanation. 75 diameters.

[496]

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[ ESSIG ] PLATE 60

PLATE 61

A radial section through Umhellularia wood with the hyphae of SehisophyUum
commune in the tracheae. 350 diameters.

[498]

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[ ESSIG J PLATE 61

INDEX

Univ. Catjf. Publ. Bot., Vol. 7

Titles of papers and names of new systematic units in boldface.
Acknowledgment of autliors cited is made for each paper under the heading
"Literature Cited."

Acacia, host for Schizophyllum com-
mune, 449, 459, 468, fig. opp. 488.

Acamptopappus sphaerocephalus, 277.

Acer (maple), host for Schizophyllum,
465.

Acknowledgments, 59, 104, 163, 186-
188, 473.

Acmaea, host for Ectocarpus cylin-
dricus f. acmaeophilus, 415.

Acrosiphonia Mertensii, 280.

Aesculus Hippocastanum (horse chest-
nut), host for Schizophyllum,
465.

"Agaricus" type, in origin of
lamellae, 461.

Alaria marginata, host for Myrio-
nema corunnae, 339.
tenuifolia, host for Streblonema
rugosum, 391.

Algae, Pacific Coast, notes on, 427-
436. See also Phycological Con-
tributions.

Alkali, tolerance of Chrysothamnus
to certain amounts of, 196, 254,
257.

Alnus (alder), host for Schizophyl-
lum, 465.

Amanita vaginata, 457; amanita type
in origin of lamellae, 461.

Apoglossum, 432.
decipiens, 432.

Artemisia californica, 277.

filifolia, 277.

Eothrockii, 277.

tridentata (sage-brush), 166, dis-
tinguished from Eabbit-brush,
191, 195 (fig.), 196, 256; no
rubber content in, 277.

Ascocyclus balticus, 341, 342.
Astragalus leucopsis, 278.
Atriplex canescens, 278.

confertifolia, 278.
Ayenia Purpusii, 328.
Baccharis glutinosa, 277.

pilularis, 277.

sergiloides, 277.
Bacterial gummosis, agent in fungus

attack, 470.
Bailey, V., cited, 209.

Baja California, undescribed plants
from, 437-446.

Bambusa (bamboo), host for Schizo-
jjhyllum, 465.

Bauhinia jucunda, 326.

Bebbia juncea, 277.

Bernardia fasciculata, 441.

Betula (birch), host for Schizophyl-
lum, 465.

Bigelovia glareosa, 173.
graveolens albicaulis, 165.

appendiculata, 167.

glabrata, 174.

hololeuca, 166.

latisquamea, 167.
juncea, 180.
leisoperma, 173.

abbreviata, 173.
mohavensis, 179.
turbinata, 180.

Blasdale, W. C, 101-157.

Boletus edulis, 457.
variegatus, 457.

Brandegee, T. S., 325-331.

Brandt, E. P., 39-68.

Brandt, E. P., and Goodspeed, T. H.,
1-24, 25-38.

Brickellia atractyloides, 277.

desertorum, 277.

microphylla, 277.
California, western middle, climatic
conditions in, 2-3.

Callithamnion, host for Ectocarpus

affinis, 406.
Cantharellus infundibuliformis, 457.

Capsosiphon aureolum, 280.

fulvescens, 280.
Carnegie Institution of Washington,
rubber-plant investigations, 251.
Castanea (chestnut), host for Schizo-
phyllum, 465.
Catalpa speciosa (hardy catalpa),

host for Schizophyllum, 465.
Chaetomorpha aerea, host for Ecto-
carpus simulans, 412.
californica, host for Entocladia
cingens, 292, fig. opp. 308.
Choristocarpus, resemblance to Ecto-
carpus chantransioides, 406, 407.

[499]

Index

Chrysil, a New Rubber from Chryso-
thamnus nauseosus, 183-:264; deti-
iiitioii of, LSS; first experiments
with, 1S8-1.S9; estimated possible
product of in western North
America, 197-210; determination
of presence of, microscopicallv,
210-215, chemically, 216-225,
with percentages, 226-2315; dis-
tribution in jilant, 2.'54-24;{; con-
tent of in different varieties,
243-244; effect of environment
on, 245 ; seasonal variation affect-
ing, 246-248; harvesting of, 248-
251; possibilities of product of,
in Chrysothannnis, 251, compared
with Guayule, 252-255.

Chrysocoma graveolens, 174.
laetevirens, 174.
nauseosa, 165.
virens, 174.

Chrysothamnus, as possible source
of rubber, 159, 184, 251-255, see
Chrysil; key to sections of, 160;
vegetative reproduction of, 250;
culture requirements, 255-257.
angustus, 177.
arizonicus, 167.

californicus occidentalis, 168, 169.
confinis, 176.
consimilis, 176.
falcatus, 176.
formosus, 180.
frigidus concolor, 170.
Greenei, 243, 277.
Howardi, 243, 277.
humilis, 243, 277.
junceus, 180, 181.
linifolius, 243, 268.
Macounii, 180.
mohavensis, 179.
nioquinaus, 180.

nauseosus, distinguishing character
of, 161; key to varieties of:
gray forms, 163-164, green
forms, 164-165; synopsis of
varieties of, 16.5-181; index to
specific and varietal names of,
181; habits and habitats, 191-
197; as source of rubber, see
Chrysil.
bemardinus, 171.
Bigelovii, 172.
caUfomicus, 174.
ceruminosus, 175.
consimilis, 176-177, 178, 192,

194, 200, 203 (fig.), 227.
frigidus, 170, 196, 209, 229.
glareosus, 173.

gnaphalodes, 166, 167-168, 192,
193, 196, 198, 199 (fig.), 229,
fig. opp. 260.

graveolens, 174, 190, 191, 230.

griseus, 202.

hololeucus, 166, 191, 192, 193.

junceus, 180.

latisquameus, 167.

leiospermus, 172, 173, 231, 244.

mohavensis, 179, 195, 198, 200,
231.

occidentaUs, 168, 231.

oreophilus, 175.

pinifoUus, 176, 192, 208, 209, 218,
231.

plattensis, 170.

salicifolius, 167.

speciosus, 169, 196, 202, 232, fig.
ojip. 262.

viridulus, 177, 188, 190, 191, 233,
fig. opp. 260, 262, 264.
nevadensis, 243, 277.
oreophilus, 175.
orthophyllus, 166.
pallidus, 170.
paniculatus, 243, 267.
Parryi, 243, 277.
patens, 176.
pinifolius, 176.
puberulus, 243, 277.
pulcherrimus, 169.
salicifolius, 167.
speciosus, 169.

albicaulis, 165.

gnaphalodes, 167.

plattensis, 170.
teretifolius, 198, 201, 242, 243, 266-

267.
tortuosus, 169.

turbinatus, 180, 207, 243, 265-266.
viscidiflorus, 243, 277.

Chrysothamnus nauseosus and its
Varieties, 159-181.

Citrus (oranges and lemons), hosts
for Schizophyllum, 465.

Cladoderris, compared to Schizo-
phyllum and Stereum, 463.
infundibuliformis, 463.

Cladophora Mertensii, 280.
viminea, 280.

Clavaria vermicularis, 457, 458.

Cneoridium dumosum, 278.

Codium fragile, host for Entocladia
codicola, 293, 294, and for species
of Ectocarpus, 407, 408, 409, 413,
415.
Setchellii, host for species of Ecto-
carpus, 407, 415.

Codiolum polyrhizum, 298, 299.

Coffea (coffee), host for Schizophyl-
lum, 465.

Colorado rubber plant. See Hymen-
oxys tloribunda utilis.

[500]

Index

Compsonema, new species of, 353-

376; relationship with Myrio-

nema, 353, and with Hecato-

nema, 377.

coniferum, 365, 366, fig. opp. 374.

dubium, 365-366, fig. opp. 374.

fasciculatum, 360, 365, fig. opp.

374.
fructuosum, 355, fig. opp. 370, 374.
gracile, 355, 360.
gracilis, 357.

intricatum, 354-355, fig. opp. 368.
myrionematoides, 361, fig. opp. 370.
nummuloides, 359-360, fig. opp.

368.
pusillum, 356, fig. opp. 372.
ramulosum, 362-363, fig. opp. 376.
secundum, 361-362, fig. opp. 372.
f. tenninale, 366, fig. opp. 372.
serpens, 363-364, 366, fig. opp. 376.
sessile, 358-359, fig. opp. 376.
speciosum f. piliferum, 356-357, fig.

opp. 374.
sporangiiferum, 357-358, fig. opp.

370.
streblonematoides, 353, fig. opp.

368.
tenue, 359, fig. opp. 372.
Condalia Parryi microphylla, 439.
Conferva Mertensii, 280.

viminea, 280.
Coprinus ephemerus, 456.

radiatus, 455.
Corethrogyne filaginifolia, 277.
Costaria costata, host for Myrionema
primarium, 334, M. corunnae,
339, and M. obscurum, 346.
Craterellus cornucopioides, 457.
Cryptantha inaequata, 444-445.
maritima pilosa, 445.
pterocarya, 445.
racemosa Ugnosa, 445.
suffruticosa, 445.
Cumagloia Andersonii,- host for
Streblonema corymbiferum, 392,
S. anomalum, 392, and S. John-
stonae, 394.
Cutleriales, 404.
Cynanchum racemosum, 331.
Cyphella, 452, 463.

Urbani, sporophore in, fig. opp. 480.
Cystophyllum geminatum, host for

Compsonema tenue, 359.
Delesseria, 430, 431.
alata, 430, 431.

Hvpoglossum, 429, 430, 431, 432.
Lyallii, 430, 431.
Montagneana, 432.
quercifolia, Californian, notes on,
427-436; characters of, differ-
ing from true type, 428; differ-
ence between D. Hypoglossum
and D. sinuosa, 431, 432.

ruscifolia, 432.

sanguinea, 342, 429, 430, 432.
sinuosa, 428, 429, 430, 431, 432.
Derbesia, host for Entocladia viridis,

293.

Desmarestia ligulata f. herbacea,

host for Streblonema transfixum,

391, for Ectocarpus acutus, 405.

Distichlis spicata (salt-grass), 195

(fig.), 196, 256.
Dictyanthus parviflorus, 329.
prostratus, 329.

Dictyosiphonales, 404.
Distichlis, 177.
Dysodia porophylloides, 277.
Ectocarpales, 403-404.
Ectocarpus, similarity with Compso-
nema, 354, 356, 360, 364, 365,
366, with Streblonema and Myrio-
nema, 387, 388, 393; new species
of, 403-426.

acutus, 404-405, 408, 411, figs. opp.
424, 426.

aeeidioides, 395.

affinis, 405-406, figs. opp. 420.

amphibius, 416.

chantransioides, 406-407, figs. opp.
424.

conunensalis, 407, figs. opp. 424.

confervoides, 408, 410, 411, 414,416.
f. acuminatus, 404.
f. parvus, 414.
f. subulatus, 416.
f. typicus, 405, 415.
f. variabilis, 408, 409, 414.

corticulatus, 408.

cylindricus, 413.

f. acmaeophilus, 415, fig. opp. 426.
f. codiophilus, 409, 415, figs. opp.
420, 426.

draparnaldiaeformis, 416.

eramosus, 407-408, figs. opp. 422.

flagelUferus, 408-409, figs. opp. 422.

flocculiformis, 409, fig. opp. 422.

fructuosus, 410, figs. opp. 418.

globifer, 413.

granulosoides, 410—411, fig. opp.
418.

granulosus, 410.

hemisphericus, 406.

macroeeras, 416.

Mesogloiae, 411, figs. opp. 418.

ovatus, 406.

Padinae, 413.

paradoxus pacificus, 412.

penicillatus, 404, 405.

pusillus, 412.

Saundersii, 411^12.

siliculosus, 414.
f. parvus, 414.
f. subulatus, 416.

simulans, 412, figs. opp. 418.

[501]

Index

socialis, 412^13, figs. opp. 420.
suhulatiis, 41().
Taeniae, 413, fig. opp. 420.
virescens, 405.
Eel-grass, host for Myrionomn globo-
sum, 342, for M. i)hyH()pliiliini,
345, for M. sotifenim, 345; for
Strobloneiiia vorax, 389; for Ecto-
carpus flagelliforus, 408.
Egrcgia Monzicsii, liost for Pseiulo-
pringshciniia apiculata, 297; for
Myrionema niimitissiiiuim, 336,
for M. corunuae, 340.
Encelia californica, 277.

farinosa, 277.
Endoderma pithophorae, 294.

])olyiii()rp]iiim, 2!>4.
Enteromorpha groenlandica, 280-281.
Entocladia cingens, 292-293, fig. opp.
308.
codicola, 293-294, fig. opp. 310.
Flustrae, 293.
viridis, 292, 293.
Ephedra californica, 437.
nevadensis, 278, 437.
pedunculata, 438.
peninsiUaris, 437-438.
Epieladia, 294.

Ericameria cuneata spatulata, 277.
Eriodictyon californicum, 278.
Eriogonum fasciculatum, 278.

nudum, 278.
Eriosema nigropunctatum, 327.
Esenbeckia ovata, 327.
Essig, F. M., 447-498.
Euphorbia consoqnitlae, 327-328.
dentosa. 439-440.
giyptospernia, 441.
leucophylla, 440.
peninsuiaris, 440.
podagrica, 440-441.
setiloba dentata, 439.
Eurotia lanata, 278.
Fagus (beech), host for Schizophyl-

lum, 465, 469.
Fimbristylis ciniiamoneus, 438.
Franscria dumosa, 277.
Frasera Parryi, 278.
Fucus, host fr Ectocarpus Saundersii,
412.
furoatus, host for Compsonema in-
ti-icatum, 354.
Fungi. Sec Uredinales; Schizophyl-

lum commune.
Galtonia, somatic mitosis in, 55.
Cardncr, N. L., and SctcheJl, W. A.,

279-324, 333-426.
Gayclla constricta, 291.
Gigartina radula, host for Myrionema
attcnuatum, 344, Compsonema
fasciculatum, 360, C. serpens, 364,
C. coniferum, 365, and C. dubium,
366.
Glycyrrhiza lepidota, 278.

Gnaphalium bicolor, 277.
Gohlen Bush" (Chrysothamnus), 191.
Gomontia Bometii, 298-299, 300.
caudata, 3()(i, fig. opp. 308.
habrorhiza, 299-300, fig. opp. 310.
polyrhiza, 298, 299, 300, fig. opp.
310.
Goodspeed, T. II., 69-100.
Goodspeed, T. H., and Bran.lt, K. P.,

1-24, 25-38.
Goodspeed, T. PL, and Hall, IT. M.,

183-264, 265-278.
Guayule, Mexican (Parthenium argen-
tatum), possible rubber produc-
ing plant, 185, 252; comparison
with Chrysothamnus, 211, 253-
254; seasonal variation in rubber
content, 246; Arizona plantings
of, 252.

Guignardia alaskana, fungus parasite
on Prasiola borealis, 291.
Ulvae, fig. opp. 306.
Gutierrezia californica, 277.
lucida, 277.
Sarothrae, 277.
Hall, H. M., 159-181.
Hall, H. M., and Goodspeed, T. IL,

1S3-264, 265-278.
Haplopappus, group of Aster tribe of
Compositae, 159, 189; comparison
w-ith Guayule and Chrysotham-
nus in rubber content, 211.
arborescens, 273.
Bloomeri, 277.
brachylepis, 273,
cervinus, 270.
ericoides, 218, 222, 270-271, 275,

276.
Fremonti, 277.
laricifolius, 272.
lincarifolius, 201, 273-274.
monactis, 201, 272.
nanus, 218, 222, 224, 268-269 (fig.),

275, 276.
Palmeri, 271.
pinifolius, 272.
Hazardia squarrosa, 277.
Hecatonema, 353, 360; new species of,
377-384; closely related to Comp-
sonema and Myrionema, 353, 377,
379.
clavatum, 378-379, figs. opp. 382.
diffusum, 357.

Lawsonii, 379-380, figs. opp. 382.
speciosum, 357.
variabile. 377. figs. opp. 384.
Ileclitia glabra, 325.

Purpusii, 325.
Iledoijliyllum sessile, host for Comp-
sonema sessile, 358; for Strcblo-
iiema aecidioides, 395.
llelmintliocladia calvadosii, host for
Streblonema investiens, 396.

[502]

Index

Hesperophycus Harveyanus, host for

Streblonema penetrale, 388.
Hicoria (hickoiy), host for Schizo-

pliylhim, 465.
Himanthalia lorea, 357.
Hofmeisteria pluriseta, 277.
Hoi-misfia collabens, 279.
doliifera, 279.
penicilliformis, 280.
Houstonia ang-ustifolia, 446.
australis, 446.
brevipes, 446.
Hydnum, 471.

Hydrolapatha (-inn), older name for
Delesseria, 429.
sangiiinea, 429.
sinuosa, 429.
Hymenoelea salsola, 277.
Hymenoxys floribiinda utilis (Pinguay
or Colorado rubber plant), 186
(footnote), 252.
Hypochnns subtilis, 455, 456, 457.
Hypoglossum, 429.

Woodwardii, 429, figs. opp. 436.
Ilea fulvespens, 280.
Ilyanassa obsoleta, host for Pseudul-

vella consociata, 297.
Indigofera acasonicae, 326-327.
Intemor©tia, 294.

Fryeana, 295, fig. opp. 308.
Iridaea laminarioides, host for Pseu-
dnlvella prostrata, 295.
' Isocoma coronopifolia, 277.
Hartwegi, 277.
veneta acradenia, 277.
Isolepis Warei, 438.

Sellowiana, 438.
Tsomeris arborea, 278.
Iva axillaris (povertv-Aveed), 195

(fig.), 196.
Jatroplia longipedunciilata, 328.

urens, 328.
Johnston, I. M., 437-446.
Juglans (English walnut), host for

Schizophyllum, 465.
Laminaria Andersonii, host for Streb-
lonema, 387, 390.

complanata, 343.

ephemera, 338.

Farlowii, 344.

groenlandica, 395.

longicniris, 395.

saeeharina, 396.

Sinclairii, 339.
Laminariales, 404.
Larrea divaricata, 278.
Lepidium Fremonti, 278.
Lepidospartum squamatum, 277.
Lepiota, 463.
Liebmannia, host for Streblonema

minutissimum, 392.
Linosyris Bigelovii, 172.

ceruminosa, 175.

Literature cited on: algae, 301-303,

347, 367, 380, 397, 417, 434;

fungus, 473-i76; Trillium, 34,

60-61, 86-87.

Littorina planaxis, host for Pseudul-

vella applanata, 296.
Lloyd, F. E., cited, 212.
Lycium Cooperi, 278.
Maba Purpusii, 329.
Macrocystis pyrif era, host for Myrio-
nema primarium, 335, and for M.
attenuatum, 344.
Marasmius, 463.

McCarty, E. C, estimate by, of Chrysil
available in northern and south-
ern Nevada, 204-207.
Mentzelia adherens, 443.
hirsutissima, 444.
nesiotes, 443, 444.
stenophyUa, 443, 444.
involucrata, 443, 444.

megalantha, 443, 444.
laevicaulis, 443.
reflexa, 444.
stenophylla, 443.
tricuspis, 443, 444.
brevicomuta, 444.
Mesogloia Andersonii, host for species

of Ectocai-pus, 405, 411.
Monostroma areolatum, 281-282, fig.
opp. 322, 324.
groenlandicum, 280.
quaternarium, 288.
zostericola, 281, 282.
Morphology, Dovelopment and Eco-
nomic Aspects of Schizophyllum
commune Fries, 447-498.
Morus (mulberr^^), host for Schizo-
phyllum, 465.
Myrionema, principal characters of,
334; new species of, 334-352; re-
lated to Compsonema, 353, 354,
355; to Hecatonema, 377, 378,
Ijassim, and to Streblonema, 387,
388.
attenuatum f. doliiforme, 344, fig.

opp. 352.
balticum f. califomicum, 341-342.

f. pediceUatum, 341, fig. opp. 350.
compsonematoides, 343, fig. opp.

352.
corunnae, 336.

f. angulatum, 340, fig. opp. 350.
f. sterile, 340, fig. opp. 350.
f. uniforme, 339, fig. opp. 350.
foecundum, 336, 343.
f. californicum, 346.
f. divergens, 338, fig. opp. 352.
f. majus, 338-339, fig. opp. 348.
f. ramulosmn, 337, fig. opp. 350.
f. simplicissimum, 336-337, 339,

fig. opp. 350, 379.
f. subulatum, 337, fig. opp. 350.

[503]

Index

plobosum f. aiiine, 342, fig. opp. SHO.
hecatonematoides, IU3, fig. opp. S.l'J.
minutissimiun, ;{3(), fig. opp. 3;')2.
obscurum, IU(), fig. ojip. 348.
phyllophilum, 344-345, 346, fig. opp.

■MS, -AJH.
primariiun, 334-33.1, fig. opp. 3;";2.
setifenim, 34ii-34(i, fig. opp. 348.
sitociosnni, 357, 3()3.
straiigulans, 346, 379.
vulgaio, 379, 380.
Mytilus californieus, host for Gomon-

tia cauflata, 300.
Neetria cinnabaiina (canker fungus),

471.
Nomalion AndtM-sonii, 392.
Nereocystie Luotkcana, host for
species of Myrionema, 336, 337,
340, 343; for Coinpsonema, 353,
3.55, 356, 357, 358, 3.59, 361, 362,
366; for Hecatonema, 377, 379;
for Streblonema, 389; for Ecto-
carpus, 410.
New Species of Compsonema, 353-

376.
New Species of Hecatonema, 377-384.
New Species of Myrionema, 334-352.
New Species of Pylaiella and Streb-
lonema, 38.')— 402.
Nicotiaiia. abnormalities in species

liybridization of, 84.
Xitophylliim, 430, 432.
Notes on the Califomian Species of
Trillium L.

I. A Report of the General Results

of Field and Garden Studies,
1911-1916, 1-24.

II. Nature and Occurrence of Un-

developed Flowers, 25-38.

III. Seasonal Changes in Trillium
Species with Special Reference
to the Reproductive Tissues,
39-68.

TV. Teratological Variations of
Trillium Sessile var. Gigan-
teum, H. & A., 69-1 00.
Notes on Pacific Coast Algae.

II. On the Califomian "Delesseria
Quercifolia," 427 -43().
Occurrence of Rubber in Certain
West American Shrubs, 265-278.
Ocotea pyramidata, 320.
Oenothera gigas, 7.
Olneya tesota, 278.
Oreodoxa regia (royal palm), host

for .Schizojdiyllum, 465.
Oiideiiiansiolla, 4()3.
Pachystelma Asclepiadacearum, 330.

cordatum, 330.
Panus, 403.

Paris quadrifolia, 40, .54, 55.
Partheiiiuni argentatuni. See Guayule.

Peziza, sporophore of, 459.
Plivcocelis foecunda, 336.
Phycodiys, 429, 430, 432.

quercifolia, 433, fig. opp. 436.
Setchellii, 433, figs. opp. 436.
siimosa, 430, 432.
Phycological Contributions. I, 279-

324. II-VI, 333^20.
Phvcoseris lapathifolia, resemblance
" to Ulva angusta, 283.
Liuza, 283.
lobata, 284.
Phyllospadix, host for Myrionema

l)alticun), 341.
Picradenia Lemmoni, 277.
Pinus ponderosa (yellow pine), host

for Schizophyllum, 465.
Plantae Mexicanae Purpusianae. X,

325-331.
Pleurophycus Gardneri, host for

Pyla'iella tenella, 385.
Plum, host for Schizophyllum, ex-
periments ■wnth, 469.
Polyponis, 471.
Polystenima rupestre, 330.

viridiflori, 330.
Polystictus, 471.

versicolor, 471.
Porphyra, 289.

Naiadum, host for Internoretia,
294, 295, and for Streblonema,
387.
Poverty-weed. See Iva axillaris.
Prasiola, resemblance to Monostroma
areolatum, 282; eight Pacific
Coast species of, 287. •
borealis, 287, 291.
calophylla, 287.
crispa, 287.
dellcata, 287, 291-292, figs. opp.

306, 310, 312.
fluviatilis, 287, 290.
Gardneri, 287.

meridionalis, 287, 291, fig. opp. 312.
nierisinopedioides, 288.
mexicana, 287-290, figs. opp. 304.
nevadensls, 287, 290, figs. opp. 304.
Preliminary List of the XJredinales

of California, 101-157.
Primula sinensis, 7.
Prosopis juliflora, 278.

jiubescens, 278.
Pruning, possible effect of on rubber

content, 246.
Prunus cerasus (cherry), host for
Schizopliylhim, 465, 470.
communis (almond), 465.
persica (peach), 465, fig. opp. 492.
Pseudodictyon, 294.
Pseud opringsh elm ia, 295.
apiculata, 2it7, figs. opp. 306.
confluen.s, 297.

[504]

Index

Pseudulvella applanata, 295-296.
consociata, 296-297, fig. opp. 310.
prostrata, 295, 296.
Pterocarpus indicus, host for Schizo-

phyllum, 465.
Pterophyllus, 463.

Pylaiella, 354, 409, 414; new species
of, 385.
fulvescens, 385.
littoralis, 385.
nana, 38.5.

teneUa, 385-386, figs. opp. 398.
imilateralis, 386-387, figs. opp. 398.
Pyrus communis (pear), 465.

mains (apple), host for Sehizophyl-
lum, 465, 469. 471, fig. opp. 492.
phyllum, 465, 469, 471, fig. opp. 492.
Quercus agrifolia, host for Schizo-
phyllum, 449, 459.
robur, 428.
serrata (oak), 465.
Eabbit-brush, 184, 190-191, 192, 256.

See Chrj'sothamnus.
Eenfrewia parvula, 339.
Ehacophyllus, 463.
Ehamnus crocea, 2^8.
Ehus integrifolia, 278.
ovata, 278.
trilobata, 278.
Eubber, in species of Chrysothamnus
otiier than nauseosus, 265-268;
in Haplopappus, 268-276; plants
lacking in, 276-277. See also
Chrysil; Chiysothamnus.
Eubber Plant Survey of Western

North. America, 159-278.
Eust flora, 102, 103. See Uredinales.
Sacchanim officinanini (sugar cane),

host for Schizophyllum, 465.
Sage-brush. See Artemisia tridentata.
Salt-grass. See Distichlis.
Sarcobatus vermiculatus (grease-
wood), 208, 278.
Schizoneura, 427, 428, 429, passim.
See Delesseria.
quercifolia, 428, 429, 430, 432, 433.
subcostata, 430.
Schizophyllum commune Fries, Mor-
phology, Development, and Eco-
nomic Aspects of, 447-498 ; sporo-
phores of, 449-463; technique
best suited, 450-451, 469; origin
of "gills" of, 461-462; extreme
vitality of, 466-467; effect of on
host, 467-469; methods of infec-
tion of, 470; preventive measures
against, 470.
Securinega acidothamnus, 442.
capensis, 441-442.
fasciculata, 441, 442.
Hallii, 442.
ramiflora, 442.
Setchell, W. A., and Gardner, X. L.,
279-324, 333-426.

[505]

Sideroxylon campestre, 329.
Simmondsia californica, 278.
Skottsberg, C, 427-436.
Smilacina, synapsis in, 53.
Solanum molestum, 331.
Sphacelariales, 404.
Spiranthes cemua, 7. ■
Spongomorpha arcta var. limitanea,
280.
Mertensii, 280.
Sporophores, in Schizophyllum com-
mune, 448, 449, 450, lyassim, figs,
opp. 478-498; technique in ex-
periments with, 450.
Stenophyllus nesloticus, 438.
Sellowiana, 438.
Warei, 438.
Stereum hirsutuni, 463.
Stevens, W. C, cited, 212.
Streblonema, 353, 356, 360; new
species of, 385.
aecidioides f. pacificum, 359, 395-

396, figs. opp. 402.
anomalum, 392-394, figs. opp. 400.
corymbiferum, 391-392, 393, 394,

fig. opp. 400.
evagatum, 390, figs. opp. 398.
faseiculatum, 393.
investiens, 396.
Johnstonae, 392, 393, 394-395, fig.

opp. 400.
niinutissimum, 392.
myrionematoides, 387-388, 390, 391,

fig. opp. 402.
pacificum, 390.

penetrale, 388, 390, figs. opp. 402.
Porphyrae, 387, fig. opp. 402.
Pringsheimii, 393.
rugosum, 390-391, figs. opp. 400.
scabiosum, 389-390, 391, fig. opp,

402.
transfixum, 391.
volubilis, 393.
vorax, 389, figs. opp. 402.
Strepsithalia investiens, 396.
Taonia Lennebackerae, host for Ecto-

carpus Taoniae, 413.
Teba-tsigupi. See Tsigupi.
Tetracoccus Hallii, 442.

dioicus, contrasted with species of
Securinega, 442.
Tetradymia canescens, 277.

glabrata, 191.
Tilia (linden), host for Schizophyl-
lum, 465.
Tremella, 471.
Tricholoma virgatum, 457.
Trillium, the Califomian Species of,
Notes on, 1-100; habitat, 2-4;
distinctive California species, 4-
5; undeveloped flowers of, nature
and occurrence, 11, 25-38; sea-
sonal changes in, with reference
to reproductive tissues, 39-68;

Index

subterrancnn organs of, 42;

active period, 40-52; dormant

period, o2-57; toratological forms

of, 09-100.
cernuum, 77.
chloropctalum, 7.
erectum, 73-74.
erythrocarpum, 52.
grandiriorum, 4, 7, 10, 25, 40, 52,

54, 55, 50, 59, 70-73, jfassim.
nivale, 4, 16, 71, 77.
ovatum, 1, 3, 4; lack of color vari-
ation in, 5; viable seeds in, 12,

14, 15, 33; contrasted with T.
sessile var. giganteum, 12, 14-

15, 33, 42, 43, 45, 46, 49, 51, 57,
59; undeveloped flowers in, 28;
teratological variations in, 77-
78.

recurvatum, 40, 77.

sessile, teratological variations in,
75.

var. angustipetalum, 6.
var. giganteum, color variation
in, 1, 7-10, fig. opp. 20; habi-
tat, 2^, fig. opp. 18; com-
parison with eastern form,
5; size variation in, 5-6, fig.
opp. 20, with reference to
gigantism, 7 ; experimental
culture, rootstocks, 10-11,
42, 43, 46, 49-51, figs. opp.
62; grafts, 14; seeds, 14, fig.
opp. 24; sterility, 11-12, 33;
vegetative propagation, 14-

15, 46, fig. opp. 22, 88; tera-
tological variations in, 15—

16, 77-82, figs. opp. 90-100;
undeveloped flowers in, 25,
32, 33, figs. opp. 36, 38;
growth of roots of, 43; con-
traction of, 43 ; crown, 44-
45; offsets, 46; growth above
ground: leaves, 46, flower,
47^8, fruit, 48, dehiscence
of, 47, 48; dormant period
of, 52, 58; pollen, cutting off
of, 52, sporogenous tissues,
52; resting stage, 53; synap-
sis, 53, 54, 58, fig. opp. 64 ;
chromosomes during meta-
phase, 54; tetrads, 55; tape-

turn, 55; development of
ovules, 56 ; mogasporo mother-
cell, 56, 58, figs. opp. 62, 66,
68.
undulatum, 76-77, 83.

Tsigupi, Paiute for Chrysothamnus,
191.

Tsuga heterophylla (western hem-
lock), host for Schizophyllum,
46.5.

Ulva angusta, 283, fig. opp. 316, 324.
aureola, 280.
californiea, 283.
dactylifera, 285-286, 287, fig. opp.

314.
expansa, 284-285.
fasciata, 284, 285, 286, 287.
fenestrata, 285.
fulvescens, 280.
Lactuca, 286.
latissima, 284.
lobata, 284.
noniatoidea, 286.

stenophylla, 282, figs. opp. 314, 320.
taeniata, 285, 286-287, pi. opp. 318.
vexata, 282-283, fig. opp. 206.
Ulvella, 295.

Umbellularia californiea (California

baytree), host for Schizophyllum,

449, 458, 459, 465, 470, figs. opp.

488, 498.

Undescribed. Plants Mostly from

Baja California, 437-446.
Urcdinales, of California, preliminary
list of, 101-157; morphology of,
104-105; index to species of,
151-154; index to host plants,
154-157.
Urospora crassa, 279.
Hartzii, 279.
incrassata, 279.
Uvularia sessilifolia, 41.
Viguiera pauciflora, 331.
Viscainoa geniculata var. pinnata,

439.
Wooton and Standley, cited, 209.
Xyleborus dispar, shot hole borer,

agent in fungus attack, 470.
Zostera, host for Monostroma, 281.
Zygomitus, 294.

EREATA

Page 277, line 14 from bottom. For Guiterrezia read Gutierrezia.
Page 443, line 6 from bottom. Bead: "cystocarpiis et spermatangiis in hieme,
cum tetrasporangiis in hieme et vere lecta."

[506]

UNIVERSITY OF CALIFORNIA PUBLICATIONS

IN

BOTANY

Vol. 7, Nos. 6-7-8, pp. 159-278, pis. 18-20, 8 figs, in text. November 7, 1919

A RUBBER PLANT SURVEY

OF

WESTERN NORTH AMERICA

BY

HARVEY MONROE HALL

AND

THOMAS HARPER GOODSPEED

UNIVERSITY OF CALIFORNIA PRESS
BERKELEY

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Physiology, and Zoology.

BOTANY. — W. A. Setchell, Editor. Price per volume, $3.50; beginning with vol. 5, $5.00.
Volumes I (pp. 418), II (pp. 360), III (pp. 400), IV (pp. 379), completed. Volumes
V, VI and Vn In progress.

Cited as Univ. Calif. Publ. Bot.

Volume 1, 1902-1903, 418 pages, with 27 plates $3.50

Volume 2, 1904-1907, 360 pages, with 29 plates 3.50

Volume 3, 1907-1909, 400 pages, with 14 plates ..-. „ 3.50

VoL 4. 1910-1912.

1. Studies in Ornamental Trees and Shnibs, by Harvey Monroe HalL Pp. 1-74,

plates 1-11, 15 text figures. March, 1910 75

2. Gracilariophila, a New Parasite on Gracilaria confervoides, by Harriet L.

Wilson. Pp. 75-84, plates 12-13. May, 1910 10

3. Plantae Mexlcauae Furpusianae, II, by T. S. Brandegee. Pp. 85-95. May,

1910 „..,_._„... „ 10

4. Leuvenia, a New Genus of Flagellates, by N. L. Gardner. Pp. 97-106, plate

14. May, 1910 .10

5. The Genus Sphaerosoma, b William Albert SetchelL Pp. 107-120, plate

15. May, 1910 _ „ IB

6. Variations in Nuclear Extrusion Among the Fucaceae, by Nathaniel Lyon

Gardner. Pp. 121-136, plates 16-17. August, 1910 15

7. The Nature of the Carpostomes in the Cystocarp of Ahnfeldtia gigartinoides,

by Ada Sara McFadden. Pp. 137-142, plate 18. February, 1911 06

8. On a Celacodasya from Southern California, by Mabel Effle McFaddMi.

Pp. 143-150, plate 19. February, 1911 05

9. Fructification of Macrocystis, by Edna Juanlta Hoffman. Pp. 151-158, plate

20. February 1911 05

10. Erythrophyllum delesserioides J. Ag., by Wilfred Charles Twlss. Pp. 159-

176, plates 21-24. March, 1911 15

11. Plantae Mexlcanae Purpusianae, m, by T. S. Brandegee. Pp. 177-194.

July, 1911 _. - „ „ 15

12. New and Noteworthy California Plants, I, by Harvey Monroe HalL Pp.

195-208. March, 1912 15

13. Die Hydrophyllaceen der Sierra Nevada, by August Brand. Pp. 209-227.

March, 1912 _ .20

14. Algae Novae et Minus Cognltae, I, by William Albert Setchell. Pp. 229-268,

plates 25-31. May, 1912 40

15. Plantae Mexlcanae Purpusianae, IV, by Townshend Stlth Brandegee. Pp.

269-281. June, 1912 _ _ IB

16. Comparative Development of the Cystocarps of Antithamnion and Prionitis,

by Lyman Luther Daines. Pp. 283-302, plates 32-34. March, 1913 20

17. Fungus Galls on Cy/stoseira and Halidrys, by Lulu May Estee. Pp. 305-316,

plate 35. March, 1913 10

18. New Fucaceae, by Nathaniel Lyon Gardner. Pp. 817-374, plates 36-53.

April, 1913 „ 75

19. Plantae Mexicanao Purpusianae, V, by Townshend Stlth Brandegee. Pp.

375-388. June, 1913 „ _ „ _ .IB

Index, pp. 889-397.

XJNIVEESITT OF OALIFOBNIA PUBLICATIONS— (Continued)

•« 6. 1912-.

1 Studies in Nicotiana, I, by William Albert SetcbeU. Pp. 1-86. December,

1912 " — 1-25

2 Quantitative Studies of Inheritance in Nicotiana Hybrids, I, by Tbomaa

Harper Ooodspeed. Pp. 87-168. December, 1912 ^ — 1.00

3 Quantitative Studies of Inheritance in Nicotiana Hybrids, II, by Thomas

' Harper Goodspeed. Pp. 169-188. January, 1913 .20

4. On the Partial Sterility of Nicotiana Hybrids made with N. sylvestris as a

Parent, by Thomas Harper Goodspeed. Pp. 189-198. March, 1913 10

5. Notes on the Germination of Tobacco Seed, I, by Thomas Harper Goodspeed.

Pp. 199-222. May, 1913 - - 25

6 Quantitative Studies of Inheritance in Nicotiana Hybrids, HI, by Thomas

Harper Goodspeed. Pp. 223-231. April, 1915 - 10

7. Notes on the Germination of Tobacco Seed, II, by Thomas Harper Good-
speed. Pp. 233-248. June, 1915 ...„ -- - - - 1&

8 Parthenogenesis, Parthenocarpy and Phenospermy in Nicotiana, by Thomas

Harper Goodspeed. Pp. 249-272, plate 35. July. 1915 ~ .25

9 On the Partial Sterility of Nicotiana Hybrids made with N. sylvestris as a

Parent, II, by T. H. Goodspeed and A. H. Ayres. Pp. 273-292, plate 38.
October, 1916 - - - —- '^^

10 On the Partial Sterility of Nicotiana Hybrids made with N. sylvestris as a

Parent IIL An Account of the Mode of Floral Abscission in the F^ Species
Hybrids, by T. H. Goodspeed and J. N. Kendall. Pp. 293-299. November,
1916 -- '^°

11 The Nature of the F, Species Hybrids between Nicotiana sylvestris and

Varieties of Nicotiana Tabacum, with Special Eeference to the Conception
of Eeaction System Contrasts in Heredity, by T. H. Goodspeed and R. E.
Clausen. Pp. 301-346, plates 37-48. January, 1917 45

12 Abscission of Flowers and Fruits in the Solanaceae, with Special Eeference

to Nicotiana, by John N. Kendall. Pp. 347-428, 10 text figures, plates 49-

53. March, 1918 - - -^

13. Controlled Pollination in Nicotiana, by Thomas Harper Goodspeed and Plrie

Davidson. Pp. 429-434. August, 1918 - l"

14. An Apparatus for Flower Measurement, by T. H. Goodspeed and B. E.

Clausen. Pp. 435-437, plate 54, 1 figure in text. September 25, 1918 .05
15 Note on the Effects of lUuminating Gas and its Constituents in Causing

Abscission of Flowers in Nicotiana and Citrus, by T. H. Goodspeed, J. M.

McGee and R. W. Hodgson. Pp. 439-450. December, 1918 ...._ 15

16. Notes on the Germination of Tobacco Seed, III. Note on the Eelation of

Light and Darkness to Germination, by T. Harper Goodspeed. Pp. 451-

455. April, 1919

k 6. 1914-. ■ -

1. Parasitic Florldeae, by William Albert SetchelL Pp. 1-34, plates 1-6.

April, 1914 "-■

2. Phytomorula regularis, a Symmetrical Protophyte Related to Coelastrum, by

ChaBles Atwood Kofoid. Pp. 35-40, plate 7. April, 1914 „ - .oo

3. Variation in Oenothera <yvata, by Katherine Layne Brandegee. Pp. 41-50,

plates 8-9. June, 1914 . — - - — ~ '^^

4. Plantae Mexlcanae Purpusianae, VI, by Townshend Stith Brandegee. Pp.

51-77. July, 1914 ^

5. The Scinaia Assemblage, by William Albert Setchell. Pp. 79-152, plates

10-12. October, 1914 .— - - " '°

6. Notes on Pacific Coast Algae, I, Pylaiella Fostelsiae, n. sp., a New Type In

the Genus Fylaiella, by Carl Skottsberg. Pp. 153-164, plates 17-19. May,
1915 „..-- ■"-■ '^^

7. New and Noteworthy OaUfomlan Plants, II, by Harvey Monroe Hall. Pp.

165-176, plate 20. October, 1915 — ^°

8. Plantae Mexlcanae Purpusianae, VTI, by Townshend StIth Brandegee. Pp.

177-197. October, 1915 _ "

05

36

UNIVEESITY OF OAIiITORNTA PUBLICATIONS— (Continued)

9. Floral Relations Among the Galapagos Islands, by A. L. Kroeber. Pp
199-220. March, 1916 ._

10. The Comparative Histology of Certain CaUfomian Boletaceae, by Harry 8.

Yates. Pp. 221-274, plates 21-25. February, 1916 '.

11. A Revision of the Tuberales of California, by Helen Margaret Oilkey. Pp

275-356, plates 26-30. March, 1916 ._

12. Species Novae vel Minus Cognitae, by T. S. Brandegee. Pp. 357-361. May

1916 „ _ ''

IS. Planta© Mexicanae Purpusianae, Vin, by Townshend Stith Brandegee Pd
363-375. March, 1917 „ _ '_

14. New Pacific Coast Marine Algae, I, by Nathaniel Lyon Gardner. Pp. 377.

416, plates 31-35. June, 1917 ^

15. An Account of the Mode of Foliar Abscission in Citrus, by Robert W.

Hodgson. Pp. 417-428, 3 text figures. February, 1918 _

16. New Pacific Coast Marine Algae, II, by Nathaniel Lyon Gardner. Pp. 429-

464, plates 36-37. July, 1918 _

17. New Pacific Coast Marine Algae, HI, by Nathaniel Lyon Gardner. Pp. 455-

486, plates 38-41. December, 1918 _ _

18. New Pacific Coast Marine Algae IV, by Nathaniel Lyon Gardner. Pp. 487-

496, plate 42. January, 1919

19. Plantae Mexicanae Purpusianae, IX, by Townshend Stith Brandegee. Pp.

497-504. November, 1919

Vol 7. 1916-.

1. Notes on the Califomlan Species of Trillium L. I, A Report of the General

Results of Field and Garden Studies, 1911-1916, by Thomas Harper Good,
speed and Robert Percy Brandt. Pp. 1-24, plates 1-4. October, 1916 ^

2. Notes on the CaHfomian Species of Trillium L. II, The Nature and Occur-

rence of Undeveloped Flowers, by Thomas Harper Goodspeed and Robert

Percy Brandt. Pp. 25-38, plates 5-6. October, 1916

8. Notes on the Califomlan Species of Trillium L. in. Seasonal Changes In
Trillium Species with Special Reference to the Reproductive Tissues, by
Robert Percy Brandt. Pp. 39-68, plates 7-10. December, 1916

4. Notes on the Califomlan Species of Trillium L. IV, Teratologlcal Varia-

tions of Trillium /tcssile var. giganteum H. & A., by Thomas Harper Good-
speed. Pp. 69-100, plates 11-17. January, 1917 ...._ „

5. A Preliminary List of the Uredlnales of California, by Walter 0. Blasdale.

Pp. 101-157. August, 1919

6, 7, 8. A Rubber Plant Survey of Westem North America. I. Chrysothamnus
nameosus and its Varieties, by Harvey Monroe Hall. n. Chrysil, a New
Rubber from Chrysothamnus nauseosus, by Harvey Monroe Hall and
Thomas Harper Goodspeed. IIL The Occurrence of Rubber in Certain
West American Slirubs, by Harvey Monroe Hall and Thomas Harper
Goodspeed. Pp. 159-279, plates 18-20, 8 figures in text. November, 1919.

UNIVERSITY OF CALIFORNIA PUBLICATIONS

IN

BOTANY

Vol. 7, No. 12, pp. 427-436, plate 50 June 16, 1922

NOTES ON PACIFIC COAST ALGAE

II. ON THE CALIFORNIAN "DELESSERIA
QUERCIFOLIA"

BY

CARL SKOTTSBERG

UNIVERSITY OF CALIFORNIA PRESS
BERKELEY, CALIFORNIA

UNIVERSITY OF CALIFOENIA PUBLICATIONS

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CALIFORNIA, U.S.A.

WILLIAM WESLEY & SONS, LONDON

Agent for the series in American Archaeology and Ethnology, Botany, Geology,

Physiology, and Zoology.

BOTAITY. — W. A. Setchell, Editor. Price per volume, $3.50 (vol. 5, $6.00) ; beginning "with
volume 6, $5.00. Volumes I, II, HI, IV, V, and IX completed. Volumes VI, vn,
Vm, and X In progress.

Cited as Univ. Calif. Publ. Bot.
Vol. 5. 1912-1922.

1. Studies in Nicotiana. I, by William Albert Setchell. Pp. 1-86. December,

1912 ._ $1.25

2. Quantitative Studies of Liheritance In Nicotiana Hybrids. I, by Thomas

Harper Goodspeed. Pp. 87-168, plates 1-28. December, 1912 „ 1.00

3. Quantitative Studies of Inheritance In Nicotiana Hybrids. II, by Thomas

Harper Goodspeed. Pp. 169-188, plates 29-34. January, 1913 20

4. On the Partial Sterility of Nicotiana Hybrids njade with N. sylvestris as a

Parent, by Thomas Harper Goodspeed. Pp. 189-198. March, 1913 10

5. Notes on the Germination of Tobacco Seed, I, by Thomas Harper Good-

speed. Pp. 199-222. May, 1913 „ _ „. .25

6. Quantitative Studies of Inheritance in Nicotiana Hybrids. HE, by Thomas

Harper Goodspeed. Pp. 223-231. April, 1915 10

7. Notes on the Germination of Tobacco Seed. II, by Thomas Harper Good-

speed. Pp. 233-248. June, 1915 „.. _ .15

8. Parthenogenesis, Parthenocarpy and Phenospermy In Nicotiana, by Thomas

Harper Goodspeed. Pp. 249-272, plate 35. July, 1915 „ .25

9. On the Partial Sterility of Nicotiana Hybrids made with N. sylvestris as a

Parent. II, by T. H. Goodspeed and A. H. Ayres. Pp. 273-292, plate 36.
October, 1916 „.. „ 20

10. On the Partial Sterility of Nicotiana Hybrids made with N. sylvestris as a

Parent. Ill, An Account of the Mode of Floral Abscission in the Fj Species
Hybrids, by T. H. Goodspeed and J. N. Kendall. Pp. 293-299. November,
1916 ~ ,05

11. The Nature of the F^ Species Hybrids between Nicotiana sylvestris and

Varieties of Nicotiana Tabacum, with Special Reference to the Conception
of Reaction System Contrasts in Heredity, by T. H. Goodspeed and R. E.
Clausen. Pp. 301-346, plates 37-48. January, 1917 45

12. Abscission of Flowers and Fruits in the Solanaceae, with Special Reference

to Nicotiana, by John N. Kendall. Pp. 347-428, 10 text figures, plates 49-

53. March, 1918 .....: 85

13. Controlled Pollination in Nicotiana, by Thomas Harper Goodspeed and Pirie

Davidson. Pp. 429-434. August, 1918 10

14. An Apparatus for Flower Measurement, by T. H. Goodspeed and R. E.

Clausen. Pp. 435-437, plate 54, 1 figure in text. September, 1918 05

15. Note on the Effects of Illuminating Gas and Its Constituents in Causing

Abscission of Flowers in Nicotiana and Citrus, by T. H. Goodspeed, J. M.
McGee and R. W. Hodgson. Pp. 439-450, December, 1918 J.0

16. Notes on the Germination of Tobacco Seed. HI, Note on the Relation of

Light and Darkness to Germination, by T. Harper Goodspeed. Pp. 451-

455. April, 1919 „ ~ 05

17. Inheritance in Nicotiana Tabacum. I, A Report on the Results of Crossing

Certain Varieties, by WUUam Albert Setchell, Thomas Harper Goodspeed,
and Roy Elwood Clausen. Pp. 457-582, 2 figures in text, plates 55-85.
April, 1922 1.76

UNIVERSITY OF CAUFOENIA PUBLICATIONS— (Continued)

Vol. 6, 1914-1919.

1. Parasitic Florideae, "by William Albert Setchell. Pp. 1-34, plates 1-6.

April, 1914 „ „ __ „ 35

2. Phytomorula reguJaris, a Symmetrical Protophyte Related to Coelastrum, by

Charles Atwood Kofoid. Pp. 35-40, plate 7. April, 1914 05

3. Variation in Oenothera ovata, by Blatherine Layne Brandegee. Pp. 41-50,

plates 8-9. June, 1914 10

4. Plantae Mexicanae Purpusianae. VI, by Townshend Stitb Brandegee. Pp.

51-77. July, 1914 „ _..... .25

5. Tbe Scinaia Assemblage, by William Albert Setchell. Pp. 79-152, plates

10-16. October, 1914 ...„..„ _... .75

6. Notes on Pacific Coast Algae. I, FylaieUa Postetsiae, n. sp., a New Type in

the Genus FylaieUa, by Carl Skottsberg. Pp. 153-164, plates 17-19. May,
1915 ....:..: „ _ „. „. 15

7. New and Noteworthy Calif omian Plants. II, by Harvey Monroe Hall. Pp.

165-176, plate 20. October, 1915 _ „. .._ .15

8. Plantae Mexicanae Purpusianae. VII, by Townshend Stith Brandegee. Pp.

177-197. October, 1915 _ .._ _ _ 25

9. Floral Relations among the Galapagos Islands, by A. L. Kroeber. Pp.

199-220, March, 1916 „.. _ 20

10. The Comparative Histology of Certain CaJifomian Boletaceae, by Harry S.

Yates. Pp. 221-274, plates 21-25. February, 1916 50

11. A Revision of the Tuberales of California, by Helen Margaret Giltey. Pp.

275-356, plates 26-30. March, 1916 .,....._.. _ .80

12. Species Novae vel Minus Cognitae, by T. S. Brandegee. Pp. 357-361. May,

1916 .„ ..„ nrrnn: „ - „ 05

13. Plantae Mexicanae Purpusianae. VIII, by Townshend Stith Brandegee.

Pp. 263-375. March, 1917 ,.„........_ 15

14. New Pacific Coast Marine Algae. I, by Nathaniel Lyon Gardner. Pp. 377-

416, plates 31-35. June, 1917 .40

15. An Account of the Mode of Foliar Abscission in Citrus, by Robert W.

Hodgson. Pp. 417-428, 3 text figures. February, 1918 ....;. ,..„ 10

16. New Pacific Coast Marine Algae, II, by Nathaniel Lyon Gardner. Pp. 429-

454, plates 36-37, July, 1918 „ 25

17. New Pacific Coast Marine Algae, in, by Nathaniel Lyon Gardner, Pp.

455-486, plates 38-41. December, 1918 „ .35

18. New Pacific Coast Marine Algae. IV, by Nathaniel Lyon Gardner. Pp.

487-496, plate 42. January, 1919 15

19. Plantae Mexicanae Purpusianae. IX, by Townshend Stith Brandegee. Pp.

497-504. November, 1919 „ 05

Index in preparation.

Vol. 7. 1916-.

1. Notes on the Calif omian Species of TriUium L. I, A Report of the General

Results of Field and Garden Studies, 1911-1916, by Tbomas Harper Good-
speed and Robert Percy Brandt. Pp. 1-24, plates 1-4. October, 1916 25

2, Notes on the Califomian Species of TriUium L. II, The Nature and Occur-

rence of Undeveloped Flowers, by Thomas Harper Goodspeed and Robert
Percy Brandt. Pp. 25-38, plates 5-6. October, 1916 .15

3, Notes on the Califomian Species of TriUium L. Ill, Seasonal Changes in

TriUium Species with Special Reference to the Reproductive Tissues, by
Robert Percy Brandt. Pp. 39-68, plates 7-10. December, 1916 _. .30

4. Notes on the Califomian Species of TriUium L. IV, Teratological Varia-

tions of TriUium sessile var. giganteum H. & A., by Thomas Harper Good-
speed. Pp. 69-100, plates 11-17. January, 1917 _ .30

UNIVEESITY OF CALIFORNIA PUBLICATIONS— (Continued)

5. A Preliminary List of the Urcdinales of California, by Walter C. Blasdale.

Pp. 101-157. August, 1919 _ - JBO

6, 7, 8. A Rubber Plant Survey of Western North America. I. Chrysothamnus
nauseosus and Its Varieties, by Harvey Monroe Hall. II. Chrysil, a New
Rubber from Chrysothamnus nauseosus, by Harvey Monroe HaU and
Thomas Harper Goodspeed. III. The Occurrence of Rubber in Certain
West American Shrubs, by Harvey Monroe Hall and Thomas Harper
Goodspeed. Pp. 159-278, plates 18-20, 8 figures in text. November, 1919. 1.25

9. Phycological Contributions. I, by William Albert Setchell and Nathaniel

Lyon Gardner. Pp. 279-324, plates 21-^1. April, 1920 50

10. Plantae Mexicanae Purpusianae. X, by Townshend Stith Brandegee. Pp.

325-331. December, 1920 ...„ 10

11. Phycological Contributions II to VL New Species of: II. Myrionema;

in. Compsonema; IV. Hecatonema; V. Pylaiella and Streblonema; VL
Ectocarpus. By William Albert Setchell and Natbaniel Lyon Gardner.
Pp. 333-426, plates 32-49. May, 1922 1.50

12. Notes on Pacific Coast Algae. II. On the Calif omian "Delesseria Querci-

folia^" by Carl Skottsberg. Pp. 427-436, plate 50. June, 1922 25

VcL 8. 1919-.

1. The Marine Algae of the Pacific Coast of Nortb America. Part L

Myxophyceae, by William Albert Setchell and Nathaniel Lyon Gardner.

Pp. 1-138, plates 1-8. November, 1919 $1.50

2. The Marine Algae of the Pacific Coast of North America. Part 11.

Chlorophyceae, by William Albert Setchell and Nathaniel Lyon Gardner.

Pp. 139-374, plates 9-33. July, 1920 „ :. 2.75

Vol. 9. A Report upon the Boreal Flora of the Sierra Nevada of California, by Frank

Jason Smiley. Pp. 1-423, plates 1-7. October, 1921 6.00

Vol. 10. 1922-.

1. The Genus Fucus on the Pacific Coast of North America, by Nathaniel Lyon

Gardner. Pp. 1-180, plates 1-60. April, 1922 2.25

UNIVERSITY OF CALIFORNIA PUBLICATIONS

IN ]

BOTANY

Vol. 7, No. 13, pp. 437-446 August 7, 1922

UNDESCRIBED PLANTS MOSTLY FROM

BAJA CALIFORNIA

BY
IVAN MURRAY JOHNSTON

UNIVERSITY OF CALIFORNIA PRESS
BERKELEY, CALIFORNIA

XJNIVERSITY OF CALIFORNIA PUBLICATIONS

Note. — The University of California Publications are offered in exchange for the pnbll>
cations of learned societies and institutions, iiniverslties and libraries. Complete lists of
all the publications of the University will be sent upon request. For sample copies, lists
of publications and other information, address the MANAGER OF THE UNIVERSITY
PRESS, BERKELEY, CALIFORNIA, U. S. A. All matter sent in exchange should be
addressed to THE EXCHANGE DEPARTMENT, UNIVERSITY LIBRARY, BERKELEY,
CALIFORNIA, U. S. A.

WILLIAM WESLEY & SONS, LONDON

Agent for the series In American Archaeology and Ethnology, Botany, Geology,

Physiology, and Zoology.

BOTANY.— W. A. Setchell, Editor, Price per volume, $3.50 (vol. 5, $6.00) ; beginning with
volume 6, $5.00. Volumes I, II, III, IV, V, VI, VII, and IX completed. Volumes
Vni, X, and XI in progress.

Cited as Univ. Calif. Publ. Bot.
Vol. 5. 1912-1922.

1. Studies in Nicotiana. 1, by William Albert Setchell. Pp. 1-86. December,

1912 ....„ „ $1.25

2. Quantitative Studies of Inheritance in Nicotiana Hybrids. I, by Thomas

Harper Goodspeed. Pp. 87-168, plates 1-28. December, 1912 1.00

3. Quantitative Studies of Inheritance tn Nicotiana Hybrids. II, by Thomas

Harper Goodspeed. Pp. 169-188, plates 29-34. January, 1913 20

4. On the Partial Sterility of Nicotiana Hybrids made with N. sylvestris as a

Parent, by Thomas Harper Goodspeed. Pp. 189-198. March, 1913 „ .10

5. Notes on the Genjaination of Tobacco Seed. I, by Thomas Harper Good-

speed. Pp. 199-222. May, 1913 _. .25

6. Quantitative Studies of Inheritance in Nicotiana Hybrids, in, by Thomas

Harper Goodspeed. Pp. 223-231. April, 1Q15 10

7. Notes on the Germination of Tobacco Seed. II, by Thomas Harper Good-

speed. Pp. 233-248. June, 1915 „ _ — .15

8. Parthenogenesis, Parthenocarpy and Phenospermy in Nicotiana, by Thomas

Harper Goodspeed. Pp. 249-272, plate 35. July, 1915 „ ~ .25

9. On the Partial Sterility of Nicotiana Eyirids made with N. sylvestris as a

Parent. II, by T. H. Goodspeed and A. H. Ayres. Pp. 273-292, plate 36.
October, 1916 : 20

10. On the Partial Sterility of Nicotiana Hybrids made with N. sylvestris as a

Parent. Ill, An Account of the Mode of Floral Abscission in the Fj Species
Hybrids, by T. H. Goodspeed and J. N. KendaU. Pp. 293-299. November,
1916 - — 05

11. The Nature of the Fj Species Hybrids between Nicotiana sylvestris and

Varieties of Nicotiana Tahacum, with Special Reference to the Conception
of Reaction System Contrasts in Heredity, by T. H. Goodspeed and K. E.
Clausen. Pp. 301-346, plates 37-48. January, 1917 45

12. Abscission of Flowers and Fruits in the Solanaceae, with Special Reference

to Nicotiana, by John N. Kendall. Pp. 347-428, 10 text figures, plates 49-

53. March, 1918 ...„ , 85

13. Controlled Pollination in Nicotiana, by Thomas Harper Goodspeed and Pirl©

Davidson. Pp. 429-434. August, 1918 J.0

14. An Apparatus for Flower Measurement, by T. H. Goodspeed and R. E.

Clausen. Pp. 435-437, plate 54, 1 figure in text. September, 1918 05

15. Note on the Effects of Illuminating Gas and Its Constituents in Causing

Abscission of Flowers in Nicotiana and Citrus, by T. H. Goodspeed, J. M.
McGee and R. W. Hodgson. Pp. 439-450. December, 1918 _ J.0

16. Notes on the Germination of Tobacco Seed. HI, Note on the Relation of

Light and Darkness to Germination, by T. Harper Goodspeed. Pp. 451-

455. April, 1919 _ „ - — .05

17. Inheritance in Nicotiana Tahacum. I, A Report on the Results of Crossing

Certain Varieties, by William Albert Setchell, Thomas Harper Goodspeed,
and Roy Elwood Clausen. Pp. 457-582, 2 figures in text, plates 55-85.
April, 1922 1.75

UNIVEESITY OF CAUFOKNIA PUBLICATIONS— (Continited)

Vol. 6. 1914-1919. V

1. Parasitic Florideae, by William Albert Setchell. Pp. 1-34, plates 1-6.

April, 1914 _ 35

2. Fhytomorula regularis, a Symmetrical Protophyte Belated to Coelastrum, by

Cbaries Atwood Kofoid. Pp. 35-40, plate 7. April, 1914 05

3. Variation in Oenothera ovata, by Eatherine Layne Brandegee. Pp. 41-50,

plates 8-9. June, 1914 : 10

4. Plant^ Mexicanae Purpusianae. VI, by Townsbend Stith Brandegee, Pp.

51-77. JiUy, 1914 „ _ 25

5. The Scinaia Assemblage, by William Albert Setchell. Pp. 79-152, plates

10-16. October, 1914 _ .75

6. Notes on Pacific Coast Algae. I, PylaieUa Postelsiae, n. sp., a New Type in

the Genus PylaieUa, by Carl Skottsberg. Pp. 153-164, plates 17-19. May,

1915 _ „ .15

7. New and Noteworthy Calif omian Plants. II, by Harvey Monroe Hall. Pp.

165-176, plate 20. October, 1915 — _..„ 15

8. Plantae Mexicanae Purpusianae. VII, by Townshend Stith Brandegee. Pp. /

177-197. October, 1915 „ „ _ 25

9. Floral Relations among the Galapagos Islands, by A. L. Kroeber. Pp.

199^220. March, 1916 20

10. The Comparative Histology of Certain Califomian Boletaceae, by Harry S.

Yates. Pp. 221-274, plates 21-25. February, 1916 50

11. A Eevision of the Tuberales of California, by Helen Margaret Gilkey. Pp.

275-356, plates 26-30. March, 1916 „ „ 80

12. Species Novae vel Minus Cognitae, by T. S. Brandegee. Pp. 357-361. May,

1916 „ 05

13. Plantae Mexicanae Purpusianae. VIII, by Townshend Stith Brandegee.

Pp. 263-375. March, 1917 _.._ „ 15

14. New Pacific Coast Marine Algae. I, by Nathaniel Lyon Gardner. Pp. 377-

416, plates 31-35. June, 1917 _... .40

15. An Account of the Mode of Foliar Abscission in Citrus, by Eoberi; W.

Hodgson. Pp. 417-428, 3 text figures. February, 1918 10

16. New Pacific Coast Marine Algae. II, by Nathaniel Lyon Gardner. Pp. 429-

454, plates 36-37. July, 1918 „ 25

17. New Pacific Coast Marine Algae, m, by Nathaniel Lyon Gardner. Pp.

455-486, plates 38-41. December, 1918 .85

18. New Pacific Coast Marine Algae. IV, by Nathaniel Lyon Gardner. Pp.

487-496, plate 42. January, 1919 15

19. Plantae Mexicanae Purpusianae. IX, by Townshend Stith Brandegee. Pp.

497-504. November, 1919 ..._ _ 05

Index in preparation.

Vol. 7. 1916-1922.

1. Notes on the Califomian Species of Trillium L. I, A Report of the General

Results of Field and Garden Studies, 1911-1916, by Thomas Harper Good-
speed and Robert Percy Brandt. Pp. 1-24, plates 1-4. October, 1916 .25

2. Notes on the Califomian Species of Trillium L. II, The Nature and Occur-

rence of Undeveloped Flowers, by Thomas Harper Goodspeed and Robert
Percy Brandt. Pp. 25-38, plates 5-6. October, 1916 .15

3. Notes on the Califomian Species of Trillium L. Ill, Seasonal Changes in

Trillium Species with Special Reference to the Reproductive Tissues, by
Robert Percy Brandt. Pp. 39-68, plates 7-10. December, 1916 _. .30

4. Notes on the Califomian Species of Trillium L. IV, Teratological Varia-

tions of Trillium sessile var. giganteum H. & A., by Thomas Harper Good-
speed. Pp. 69-100, plates 11-17. January, 1917 .80

ITNIVEESITY OF CAUFOENIA PUBLICATIONS— (Continued)

5. A Preliminary List of the Uredinales of California, by Walter C. Blasdale.

Pp. 101-157. August, 1919 £0

6, 7, 8. A Eubber Plant Survey of Western North Air.erica. I. Chrysothamnus
nmiseosu-s and Its Varieties, by Harvey Monroe Hall. II. Chrysil, a New
Eubber from Chrysothamnus nauaeosus, by Harvey Monroe Hall and
Thomas Harper Goodspeed. III. The Occurrence of Eubber in Certain
West American Shrubs, by Harvey Monroe Hall and Thomas Harper
Goodspeed. Pp. 159-278, plates 18-20, 8 figures in text. November, 1919. 1.25

9. Phycologlcal Contributions. I, by WiUiara Albert Setchell and Nathaniel

Lyon Gardner. Pp. 279-324, plates 21^1. April, 1920 50

10. Plantae Mexicanae Purpusianae. X, by Townshend Stith Brandegee. Pp.

325-331. December, 1920 ..._ _ - J.0

11. Phycologlcal Contributions n to VI. New Species of: II. Myrionema;

in. Compsoneraa; IV. Hecatonema; V. Pylaiella and Streblonema; VI.
Ectocarpus. By WiUiam Albert Setchell and Nathaniel Lyon Gardner.
Pp. 333-426, plates 32-49. May, 1922 1.50

12. Notes on Pacific Coast Algae. U. On the Calif omian "Delesseria Querci-

foUa," by Carl Skottsberg. Pp. 427-436, plate 50. June, 1922 25

13. Undescribed plants mostly from Baja California, by Ivan Murray Johnston.

Pp. 437-446. August, 1922 25

14. Morphology, Development, and Economic Aspects of ScliizophiiUum com-

mune Fries, by Frederick Monroe Essig. Pp. 447-498, plates 51-61.

August, 1922 80

Index in preparation.

VcL 8. 1919-.

1. The Marine Algae of the Pacific Coast of North America. Part L

Myxophyceae, by William Albert Setchell and Nathaniel Lyon Gardner.

Pp. 1-138, plates 1-8. November, 1919 _ $1.50

2. The Marine Algae of the Pacific Coast of North America. Part IL

Chlorophyceae, by William Albert Setchell and Nathaniel Lyon Gardner.

Pp. 139-374, plates 9-33. July, 1920 _ 2.75

VoL 9. A Eeport upon the Boreal Flora of the Sierra Nevada of California^ by Frank

Jason Smiley. Pp. 1-423, plates 1-7. October, 1921 _ _.. 5.00

VoL 10. 1922-.

1. The Genus Fucus on the Pacific Coast of North America, by Nathaniel Lyon

Gardner. Pp. 1-180, plates 1-60. April, 1922 2.25

UNIVERSITY OF CALIFORNIA PUBLICATIONS

IN

BOTANY

Vol. 7, No. 14, pp. 447-498, plates 51-61 August 11, 1922

THE MORPHOLOGY, DEVELOPMENT, AND
ECONOMIC ASPECTS OF SCHIZOPHYLLUM

COMMUNE FRIES

BY
FREDERICK MONROE ESSIG

UNIVERSITY OF CALIFORNIA PRESS
BERKELEY, CALIFORNIA

UNIVEESITY OF CALIFORNIA PUBLICATIONS

Note. — The University of California Publications are offered in exchange for the publl.
cations of learned societies and Institutions, luiiyersities and libraries. Complete lists of
all the publications of the University will be sent upon request. For sample copies, llsta
of publications and other information, address the MANAGER OF THE UNIVEESITY
PRESS, BERKELEY, CALIFOENIA, U. S. A. AH matter sent in exchange should be
addressed to THE EXCHANGE DEPARTMENT, UNIVERSITY LIBRARY, BERKELEY,
CALIFORNIA, U.S.A.

WILLIAM WESLEY & SONS, LONDON

Agent for the series in American Archaeology and Ethnology, Botany, Geology,

Physiology, and Zoology.

BOTANY.— W. A. Setchell, Editor. Price per volume, $3.50 (vol. 5, $6.00) ; beginning with
volume 6, $5.00. Volumes I, 11, HI, IV, V, VI, VII, and IX completed. Volumei
Vni, X, and XI in progress.

Cited as Univ. Calif. Publ. Bot.
VoL 5. 1912-1922.

1. Studies in Nicotiana. I, by William Albert Setchell. Pp. 1-86. December,

1912 _ _ $1.25

2. Quantitative Studies of Inheritance in Nicotiana Hybrids. I, by Thomas

Harper Goodspeed. Pp. 87-168, plates 1-28. December, 1912 _ 1.00

S. Quantitative Studies of Inheritance in Nicotiana Hybrids, n, by Thomas

Harper Goodspeed. Pp. 169-188, plates 29-34. January, 1913 ...„ 20

4. On the Partial Sterility of Nicotiana Hybrids made with N. sylvestris as a

Parent, by Thomas Harper Goodspeed. Pp. 189-198. March, 1913 10

5. Notes on the Germination of Tobacco Seed. I, by Thomas Harper Good-

speed. Pp. 199-222. May, 1913 25

.6. Quantitative Studies of Inheritance in Nicotiana Hybrids, m, by Thomas

Harper Goodspeed. Pp. 223-231. April, 1915 „ _ 10

7. Notes on the Grenntnation of Tobacco Seed. U, by Thomas Harper Good-

speed. Pp. 233-248. June, 1915 ..._ - _ .15

8. Parthenogenesis, Parthenocarpy and Phenospermy in Nicotiana, by Thomaa

Harper Goodspeed. Pp. 249-272, plate 35. July, 1915 ..._ „ 25

9. On the Partial Sterility of Nicotiana Hybrids made with N. sylvestris as a

Parent. II, by T. H. Goodspeed and-A. H. Ayres. Pp. 273-292, plate 36.
October, 1916 „ 20

10. On the Partial Sterility of Nicotiana Hybrids made with N. sylvestris as a

Parent. Ill, An Account of the Mode of Floral Abscission In the Fi Species
Hybrids, by T. H. Goodspeed and J. N. Kendall. Pp. 293-299. November,
1916 ..._..^- - _ .05

11. The Nature of the Fi Species Hybrids between Nicotiana sylvestris and

Varieties of Nicotiana Tabacum, with Special Reference to the Conception
of Reaction System Contrasts In Heredity, by T. H. Goodspeed and R. E.
Clausen. Pp. 301-346, plates 37-48. January, 1917 45

12. Abscission of Flowers and Fruits in the Solanaceae, with Special Reference

to Nicotiana, by John N. Kendall. Pp. 347-428, 10 text figures, plates 49-

53. March, 1918 _ 85

13. Controlled Pollination In Nicotiana, by Thomas Harper Goodspeed and Pirle

Davidson. Pp. 429-434. August, 1918 _ „. J.0

14. An Apparatus for Flower Measurement, by T. H. Goodspeed and R. B.

Clausen. Pp. 435-437, plate 54, 1 figure in text. September, 1918 05

15. Note on the Effects of niuminatlng Gas and Its Constituents in Causing

Abscission of Flowers in Nicotiana and Citrus, by T. H. Goodspeed, J. M.
McGee and R. W. Hodgson. Pp. 439-450. December, 1918 ..._ _ _ J.0

16. Notes on the Germination of Tobacco Seed, m. Note on the Relation of

Light and Darkness to Germination, by T. Haarper Goodspeed. Pp. 451-

455. April, 1919 .._ „ — .05

17. Inheritance in Nicotiana Tabacum. I, A Report on the Results of Crossing

Certain Varieties, by William Albert Setchell, Thomas Harper Goodspeed,
and Roy Elwood Clausen. Pp. 457-582, 2 figures in text, plates 55-85.
April, 1922 1.76

UNIVEESITY OF CAUFOENIA PUBLICATIONS— (Continiied) "-

Vol. 6. 1914-1919.

1. Parasitic Florideae, lay William Albert Setchell. Pp. 1-34, plates 1-6.

April, 1914 „ .„ 35

2. PhytomoruJa regularis, a Symmetrical Protophyte Related to Coelastrum, by

Charles Atwood Kofoid. Pp. 35-40, plate 7. April, 1914 „ 05

3. Variation in Oenothera ovata, by Katherine Layne Brandegee. Pp. 41-50,

plates 8-9, June, 1914 „ .10

4. Plantae Mesicanae Purpusianae. VI, by Townshend Stith Brandegee. Pp.

51-77. July, 1914 „ _ _ SS

5. The Scinaia Assemblage, by William Albert Setchell. Pp. 79-152, plates

10-16. October, 1914 .75

6. Notes on Pacific Coast Algae. I, Pylaiella Postelsiae, n. sp., a New Type in

the Genus Pylaiella, by Carl Skottsberg. Pp. 153-164, plates 17-19. May,

1915 „ „ 15

7. New and Noteworthy Calif ornian Plants. II, by Harvey Monroe Hall. Pp.

165-176, plate 20. October, 1915 _ _..„ 15

8. Plantae Mexicanae Purpusianae. VII, by Townshend Stith Brandegee. Pp.

177-197. October, 1915 „ .._ „ _ _ 25

9. Floral Relations among the Galapagos Islands, by A. L. Kroeber. Pp.

199^220. March, 1916 „ 20

10. The Comparative Histology of Certain CaUfomian Boletaceae, by Harry S.

Yates. Pp. 221-274, plates 21-25. February, 1916 50

11. A Revision of the Tnberales of California, by Helen Margaret Gilkey. Pp.

275-356, plates 26-30. March, 1916 _ , _ 80

12. Species Novae vel lilinus Cognitae, by T. S. Brandegee. Pp. 357-361. May,

1916 , 05

13. Plantae Mericana« Purpusianae. Vin, by Townshend Stith Srandegee.

Pp. 263-375. March, 1917 _ „. .15

14. New Pacific Coast Marine Algae. I, by Nathaniel Lyon Gardner. Pp. 377-

416, plates 31-35. June, 1917 „ .40

15. An Account of the Mode of Foliar Abscission in Citrus, by Robert W.

Hodgson. Pp. 417-428, 3 text figures. February, 1918 ....„ 10

16. New Pacific Coast Marine Algae. II, by Nathaniel Lyon Gardner. Pp. 429-

454, plates 36-37. July, 1918 „ .„ 25

17. New Pacific Coast Marine Algae, m, by Nathaniel Lyon Gardner. Pp.

455-486, plates 38-41. December, 1918 .35

18. New Pacific Coast Marine Algae. IV, by Nathaniel Lyon Gardner. Pp.

487-496, plate 42. January, 1919 ..._ 15

19. Plantae Mexicanae Purpusianae. IX, by Townshend Stith Brandegee. Pp.

497-504. November, 1919 „ „ ..„ „ 05

Index in preparation.

Vol. 7. 1916-1922.

1. Notes on the Calif omlan Species of Trillium L. I, A Report of the General

Results of Field and Garden Studies, 1911-1916, by Thomas Harper Good-
speed and Robert Percy Brandt. Pp. 1-24, plates 1-4. October, 1916 .25

2. Notes on the Califomian Species of Trillium L. II, The Nature and Occur-

rence of Undeveloped Flowers, by Thomas Harper Goodspeed and Robert
Percy Brandt. Pp. 25-38, plates 5-6. October, 1916 _. .15

S. Notes on the Califomian Species of Trillvum L. m. Seasonal Changes in
Trillium Species with Special Reference to the Reproductive Tissues, by
Robert Percy Brandt. Pp. 39-68, plates 7-10. December, 1916 _. .30

4. Notes on the Califomian Species of Trillvum L. IV, Teratological Varia-
tions of Trillium sessile var. giganteum H, & A., by Thomas Harper Good-
speed. Pp. 69.-100, plates 11-17. Jaunary, 1917 .30

UNTVEESrTT OF CALIF OIINIA PUBLICATIONS— (Continued)

5. A Preliminary List of the Uredinales of California, by Walter C. Blasdale.

Pp. 101-157. August, 1919 - J50

6, 7, 8. A Rubber Plant Survey of Western North America. L Chrysothamnus
nauf^eosus and Its Varieties, by Harvey Monroe Hall. II. Chrysil, a New
Rubber from Chrysothamnus nauaeosus, by Harvey Monroe Hall and
Thomas Harper Goodspeed. III. The Occurrence of Rubber in Certain
West American Shrubs, by Harvey Monroe Hall and Thomas Harper
Goodspeed. Pp. 159-278, plates 18-20, 8 figures in text. November, 1919. 1.25

9. Phycological Contributions. I, by WiUlam Albert Setchell and Nathaniel

Lyon Gardner. Pp. 279-324, plates 21-51. AprO, 1920 50

10. Plantae Mexlcanae Purpusianae. X, by Townshend Stith Brandegee. Pp.

325-331. December, 1920 ...„ „ JO

11. Phycological Contributions II to VI. New Species of: II. Myrionema;

m. Compsonema; IV. Hecatonema; V. Pylaiella and Streblonema; VI.
Ectocarpus. By William Albert SetcheU and Nathaniel Lyon Gardner.
Pp. 333-426, plates 32-49. May, 1922 „ 1.50

12. Notes on Pacific Coast Algae. II. On the Calif omian "Delesseria Querci-

folia>" by Carl Skottsberg. Pp. 427-436, plate 50. June, 1922 25

13. XJndescribed plants mostly from Baja CaUfoipnia, by.Ivan Murray Johnston.

Pp. 437-446. August, 1922 .;.:.„.J.i,„4...i...„ 25

14. Morphology, Development, and Economic Aspects of Schizophylluvi com-

mune Fries, by Frederick Monroe Essig. ^ Pp. 447-498, pla,tes 51-61.

August, 1922 ; 80

Index in preparation.

VcL 8. 1919-.

1. The Marine Algae of the Pacific Coast of North America. Part L

Myxophyceae, by WUliam Albert Setchell and Nathaniel Lyon Gardner.

Pp. 1-138, plates 1-8. November, 1919 „ $1.50

2. The Marine Algae of the Pacific Coast of North America. Part IL

Chlorophyceae, by William Albert Setchell and Nathaniel Lyon Gardner.

Pp. 139-374, plates 9-33. July, 1920 „ 2.75

Vol. 9. A Report upon the Boreal Flora of the Sierra Nevada of CaUfomia, by Frank

Jason Smiley. Pp. 1-423, plates 1-7. October, 1921 _ 5.00

Vol. 10. 1922-.

1. The Genus Fucus on the Pacific Coast of North America, by Nathaniel Lyon

Gardner. Pp. 1-180, plates 1-60. April, 1922 2.25

Vol. 11. 1922-.

1. Interspecific Hybridization in Nicotiana. I. On the Results of Backcrossing
the F, Sylvestris-T ahacum. Hybrids to Sylvestris, by Thomas Harper Good-
speed and Roy Elwood Clausen. Pp. 1-30. August, 1922 45

UNIVERSITY OF CALIFORNIA PUBLICATIONS

IN

BOTANY

Vol. 7, No. 9, pp. 279-324, plates 21-31 July, 8, 1920

\

PHYCOLOGICAL CONTRIBUTIONS

I

BY

WILLIAM ALBERT SETCHELL

AND
NATHANIEL LYON GARDNER

UNIVERSITY OF CALIFORNIA PRESS
BERKELEY

UNIVERSITY OF CAIiIFORNIA PUBLICATIONS

II-Qte. — The University of California Publications are offered in exchange for the publl-
catlonfl of learned societies and institutions, universities and libraries. Complete lists of
all the publications of the University will be sent upon request. For sample copies, lists
of publications and other information, address the MANAGER OF THE UNIVERSITY
PRESS, BERKELEY, CALIFORNIA, U. S. A. All matter sent in exchange should b«
addressed to THE EXCHANGE DEPARTMENT, UNIVERSITY LIBRARY, BERKELEY,
CALIFORNIA, U.S.A.

WILLIAM WESLEY & SONS, LONDON
Agent for the series in American Archaeology and Ethnology, Botany, Geology,

Physiology, and Zoology.

BOTANY. — W. A. Setchell, Editor. Price per volume, '?3.50; beginning with vol. 5, $5.00.
Volumes I (pp. 418), II (pp. 360), m (pp. 400), IV (pp. 379), completed. Volumes
V, VI, VII, and VIII in progress.

Cited as Univ. Calif. Publ. Bot.

Volume 1, 1902-1903, 418 pages, with 27 plates $3.50

Volume 2, 1904-1907, 360 pages, with 29 plates _ „ 3.50

Volume 3, 1907-1909, 400 pages, with 14 plates _ 3.50

Vol 4. 1910-1912.

1. Studies in Ornamental Trees and Shrubs, by Harvey Monroe HalL Pp. 1-74,

plates 1-11, 15 text figures. March, 1910 _ .75

2. GracUariophUa, a New Parasite on Gracilaria confervoides, by Harriet L.

Wilson. Pp. 75-84, plates 12-13. May, 1910 ...„ .10

3. Plantae Mexicanae Purpusianae, II, by T. S. Brandegee. Pp. 85-95. May,

1910 : - 10

4. Leuvenia, a New (Jenus of Flagellates, by N. L. Gardner. Pp. 97-106, plate

14. May, 1910 _ _ .10

5. The Genus Sphnerosoma, by William Albert Setchell. Pp. 107-120, plate

15. May, 1910 ..„ _ _ _ .15

6. Variations in Nuclear Extrusion Among the Fucaceae, by Nathaniel Lyon

Gardner. Pp. 121-136, plates 16-17. August, 1910 15

7. The Nature of the Carpostomes In the Cystocarp of Ahnfeldtia gigartinoides,

by Ada Sara McFadden. Pp. 137-142, plate 18. February, 1911 ..„ 05

8. On a Celacodasya from Southern California, by Mabel Eflle McFadden.

Pp. 143-150, plate 19. February, 1911 _ „ .05

9. Fructification of Macrocystis, by Edna Juanita Hoffman. Pp. 151-158, plate

20. February 1911 .05

10. Erythrophyllum delesserioides J. Ag., by Wilfred Charles Twiss. Pp. 159-

176, plates 21-24. March, 1911 _ .16

11. Plantae Mexicanae Purpusianae, in, by T. S. Brandegee. Pp. 177-194.

12. New and Noteworthy California Plants, I, by Harvey Monroe HalL Pp.

195-208, March, 1912 _ _„ .16

18. Die Hydrophyllaceen der Sierra Nevada, by August Brand. Pp. 209-227.

March, 1912 „ „ _. .20

14. Algae Novae et Minus Cognitae, I, by William Albert Setchell. Pp. 229-268,

plates 25-31. May, 1912 :..... 40

15. Plantae Mexicanae Purpusianae, IV, by Townshend Stith Brandegee. Pp.

269-281. June, 1912 _ „ „ =16

16. Comparative Development of the Cystocarps of Antithamnion and Prionitis,

by L3rman Luther Daines. Pp. 283-302, plates 32-34. March, 1913 _ .20

17. Fungus Galls on Cystoseira and Halidrys, by Lulu May Estee. Pp. 305-316,

plate 35. March, 1913 _ .^..„ ...^-....^ .10

18. New Fucaceae, by Nathaniel Lyon Gardner. Pp. 817-374, plates 36-53.

April, 1913 „ _ 75

19. Plantae Mexicanae Purpusianae, V, by Townshend Stith Brandegee. Pp.

875-388. June, 1913 .16

Index, pp. 389-397.

UNIVEESITY OF CALIFORNIA PUBLICATIONS— (Continued)

VOL 6. 1912-.

1. Studies in Nicotiana, I, by William All}ert SetchelL Pp. 1-86. December,

2. Quantitative Studies of Inheritance in Nicotiana Hybrids, I, by Tbomaa

Harper Goodspeed. Pp. 87-168. December, 1912 „ 1.00

3. Quantitative Studies of Inheritance in Nicotiana Hybrids, H, by Thomas

Harper Goodspeed. Pp. 169-188. January, 1913 ...„ _ __ ,.20

4. On the Partial Sterility of Nicotiana Hybrids made with N. sylvestris as a

Parent, by Thomas Harper Goodspeed. Pp. 189-198. March, 1913 .10

5. Notes on the Germination of Tobacco Seed, I, by Thomas Harper Goodspeed.

Pp. 199-222. May, 1913 . _ .25

6. Quantitative Studies of Inheritance in Nicotiana Hybrids, m, by Thomas

Harper Goodspeed. Pp. 223-231. April, 1915 _ _ .10

7. Notes on the Germination of Tobacco Seed, II, by Thomas Harper Good-

speed. Pp. 233-248. June, 1915 „ „ .15

8. Parthenogenesis, Parthenocarpy and Phenospermy In Nicotiana, by Thomas

Harper Goodspeed. Pp. 249-272, plate 35. July, 1915 . — „ „ .26

9. On the Partial Sterility of Nicotiana Hybrids made 'VTlth N. sylvestris as a

Parent, II, by T. H. Goodspeed and A. H. Ayres. Pp. 273-292, plate 36.
October, 1916 20

10. On the Partial Sterility of Nicotiana Hybrids made with N. sylvestris as a

Parent, III, An Account of the Mode of Floral Abscission in the F, Species
Hybrids, by T. H. Goodspeed and J. N. Kendall. Pp. 293-299. November,
1916 „ - _ .05

11. The Nature of the Fi Species Hybrids between Nicotiana sylvestris and

Varieties of Nicotiana Tdbacum, with Special Reference to the Conception
of Reaction System Contrasts in Heredity, by T. H. Goodspeed and R. E.
Clausen. Pp. 301-346, plates 37-48. January, 1917 45

12. Abscission of Flowers and Fruits In the Solanaceae, with Special Reference

to Nicotiana, by John N. Kendall. Pp. 347-428, 10 text figures, plates 49-

53. March, 1918 „ „ „ .85

IS. Controlled Pollination in Nicotiana, by Thomas Harper Goodspeed and Pirie

Davidson. Pp. 429-434. August, 1918 _ „ .10

"^ 14. An Apparatus for Flower Measurement, by T. H. Goodspeed and R. E.

Clausen. Pp. 435-437, plate 54, 1 figure in text. September 25, 1918 .05

15. Note on the Effects of Illuminating Gas and its Constituents in Causing

Abscission of Flowers in Nicotiana and Citrus, by T. H. Goodspeed, J. M.
McGee and R. W. Hodgson. Pp. 439-450. December, 1918 ....„ 15

16. Notes on the Germination of Tobacco Seed, III. Note on the Relation of

Light and Darkness to Germination, by T. Harper Goodspeed. Pp. 451-

455. April, 1919 „ 05

VoL 6. 1914-.

1. Parasitic Florideae, by William Albert Setchell. Pp. 1-34, plates 1-6.

April, 1914 _ _ .35

2. Phytomorula regularis, a Symmetrical Protophyte Related to Coelastrum, by

Charles Atwood Kofoid. Pp. 35-40, plate 7. April, 1914 ™ .05

S. Variation in Oenothera ovata, by Katherine Layne Brandegee. Pp. 41-50,

plates 8-9. June, 1914 ...„ _ _ .10

4. Plantae Mexicanae Purpusianae, VI, by Townshend Stith Brandegee. Pp.

51-77. July, 1914 „ 25

5. The Scinma Assemblage, by William Albert Setchell. Pp. 79-152, plates

10-12. October, 1914 _ ~ .75

6. Notes on Pacific Coast Algae, I, Pylaiella Posielsiae, n. sp., a New Type In

the Genus Pylaiella, by Carl Skottsberg. Pp. 153-164, plates 17-19. May,
1915 „ . . s _ .16

7. New and Noteworthy CaTlfomian Plants, H, by Harvey Monroe Hall. Pp.

165-176, plate 20. October, 1915 _ .15

8. Plantae Mexicanae Purpusianae, VII, by Townshend Stith Brandegee. Pp.

177-197. October, 1915 „ ^

UNIVEBSITY or CALIFOENIA PUBLICATIONS— (Continued)

9. rioral Eolations Among the Galapagos Islands, by A. L. Kroeber. Pp.

199-220, March, 1916 .20

10. The Comparative Histology of Certain Oallfomlan Boletaceae, by Harry S.

Yates, Pp, 221-274, plates 21-25. Pebrtiary, 1916 £0

11. A Eevision of the Tuberales of California, by Helen Margaret Qllkcy. Pp.

275-356, plates 26-30, March, 1916 80

12. Species Novae vel Minus Cognitae, by T. S, Brandegee. Pp. 357-361. May,

1916 - .-. .05

13. Plantae Mexicanae Purpusianae, Vm, by Townshend Stith Brandegee. Pp.

363-375. March, 1917 _ _ .16

14. New Pacific Coast Marine Algae, I, by Nathaniel Lyon Gardner. Pp. 377-

416, plates 31-35, June, 1917 „ _ .40

15. An Account of the Mode of Foliar Abscission in Citrus, by Robert W.

Hodgson, Pp. 417-428, 3 text figures. February, 1918 .10

16. New Pacific Coast Marine Algae, II, by Nathaniel Lyon Gardner. Pp, 429-

454, plates 36-37. July, 1918 :. 25

17. New Pacific Coast Marine Algae, HI, by Nathaniel Lyon Gardner. Pp, 455-

486, plates 38-41, December, 1918 „ _ _ _ .36

18. New Pacific Coast Marine Algae IV, by Nathaniel Lyon Gardner. Pp, 487-

496, plate 42. January, 1919 _ „ .15

19. Plantae Mexicanae Purpusianae, IX, by Townshend Stith Brandegee, Pp,

497-504. November, 1919 _ 10

Vol 7. 1916-.

1. Notes on the Califomlan Species of Trillium L. I, A Eeport of the General

Results of Field and Garden Studies, 1911-1916, by Thomas Harper Good-
speed and Robert Percy Brandt. Pp. 1-24, plates 1-4. October, 1916 25

2. Notes on the Califomlan Species of Trillium L, II, The Nature and Occur-

rence of Undeveloped Flowers, by Thomas Harper Goodspeed and Robert
Percy Brandt. Pp. 25-38, plates 5-6. October, 1916 „ 15

3. Notes on the Callfomian Species of Trillium L. m, Seasonal Changes In

Trillium Species with Special Reference to the Reproductive Tissues, by
Robert Percy Brandt. Pp. 39-68, plates 7-10. December, 1916 .30

4. Notes on the Califomlan Species of Trillium L. IV, Teratologlcal Varia-

tions of Trillium sessile var. giganteum H. & A,, by Thomas Harper Good-
speed. Pp, 69-100, plates 11-17. January, 1917 „. .30

5. A Preliminary List of the Uredinales of California, by Walter C. Blasdale.

Pp, 101-157. August, 1919 50

6, 7, 8, A Rubber Plant Survey of Western North America. I. Chrysothamnw
7iauseosus and Its Varieties, by Harvey Monroe HaU, n. Chrysil, a New
Rubber from Chrysothamnus nauseosus, by Harvey Monroe Hall and
Thomas Harper Goodspeed, III. The Occurrence of Rubber in Certain
West American Shrubs, by Harvey Monroe Hall and Thomas Harper
Goodspeed, Pp. 159-278, plates 18-20, 8 figures in text. November, 1919. 1.25

9. Phycological Contributions. I, by William Albert Setchell and Nathaniel

Lyon Gardner, Pp, 279-324, plates 21-31. April, 1920 50

Vol. 8, 1919-

1. The Marine Algae of the Pacific Coast of North America, Part I,
Myrxophyceae, by William Albert Setchell and Nathaniel Lyon Gardner,
Pp. 1-138, plates 1-8, November, 1919 $1,50

UNIVERSITY OF CALIFORNIA PUBLICATIONS

IN

BOTANY

VoL 7, No. 10, pp. 325-331 December 29, 1920

PLANTAE MEXICANAE PURPUSIANAE, X

BY
TOWNSHEND STITH BRANDEGEE

UNIVERSITY OF CALIFORNIA PRESS
BERKELEY

UNIVERSITY OF CALITOENIA PUBLICATIONS

Note. — Tlie University of California Publications are offered in exchange for the publi-
cations of learned societies and institutions, universities and libraries. Complete lists of
all the publications of the University will be sent upon request. For sample copies, lists
of publications and other information, address the MANAGER OF THE UNIVERSITY
PRESS, BERKELEY, CALIFORNIA, U. S. A. All matter sent in exchange should be
addressed to THE EXCHANGE DEPARTMENT, UNIVERSITY LIBRARY, BERKELEY,
CALIFORNIA, U.S.A.

AVTLLTAM AVBSLElT &'sONS, LONDON
Agent for the series in American Archaeology and Ethnology, Botany, Geology,

Physiology, and Zoology.

BOTANY. — W. A. Setchell, Editor. Price per volume, $3.50; beginning with vol. 5, $5.00.
Volumes I, II, HI, IV, and IX completed. Volumes V, VI, VII, and VIII in progress.

Cited as Univ. Calif. Publ. Bot.

Vol. 5. 1912-.

1. Studies in Nicotiana. I, by William Albert Setchell. Pp. 1-86. December,

1912 ..., $1.25

2. Quantitative Studies of Inheritance in Nicotiana Hybrids. I, by Thomas

Harper Goodspeed. Pp. 87-168. December, 1912 1.00

3. Quantitative Studies of Inheritance in NicotianaHy'bTids. II, by Thomas

Harper Goodspeed. Pp. 169-188. January, 1913 20

4. On the Partial Sterility of Nicotiana Hybrids njade with N. sylvestris as a

Parent, by Thomas Harper Goodspeed. Pp. 189-198. March, 1913 10

5. Notes on the Germination of Tobacco Seed. I, by Thomas Harper Good-

speed. Pp. 199-222. May, 1913 25

6. Quantitative Studies of Liheritance in Nicotiana Hybrids, in, by Thomas

Harper Goodspeed. Pp. 223-231. April, 1915 10

7. Notes on the Germination of Tobacco Seed. II, by Thomas Harper Good-

speed. Pp. 233-2i8. June, 1915 15

8. Parthenogenesis, Parthenocarpy and Phenospermy in Nicotiana; by Thomas

Harper Goodspeed. Pp. 249-272, plate 35. July, 1915 .25

9. On the Partial Sterility of Nicotiana Hyhrirls made with N. sylvestris as a

Parent. II, by T. H. Goodspeed and A. H. Ayres. Pp. 273-292, plate 36.
October, 1916 20

10. On the Partial Sterility of Nicotiana Hybrids made with N. sylvestris as a

Parent. Ill, An Account of the Mode of Floral Abscission in the F, Species
Hybrids, by T. H. Goodspeed and J. N. Kendall. Pp. 293-299. November,
1916 .....:. 05

11. The Nature of the Fj Species Hybrids between Nicotiana sylvestris and

Varieties of Nicotiana Tabacum, with Special Reference to the Conception
of Reaction System Contrasts in Heredity, by T. H. Goodspeed and R. E.
Clausen. Pp. 301-346, plates 37-48, January, 1917 45

12. Abscission of Flowers and Fruits in the Solanacea«, with Special Reference

to Nicotiana, by John N. Kendall. Pp. 347-428, 10 text figures, plates 49-

53. March, 1918 : 85

13. Controlled Pollination in Nieotiana, "by Thomas Harper Goodspeed and Pirie

Davidson. Pp. 429-434. August, 1918 .10

14. An Apparatus for Flower Measurement, by T. H. Goodspeed and R. E.

Clausen. Pp. 435-437, plate 54, 1 figure in text. September, 1918 05

15. Note on the Effects of Illuminating Gas and its Constituents in Causing

Abscission of Flowers in Nicotiana and Citrus, by T. H. Goodspeed, J. M.
McGee and R. W. Hodgson. Pp. 439-450. December, 1918 15

16. Notes on the Germination of Tobacco Seed. HI, Note on the Relation of

Light and Darkness to Germination, by T. Harper Goodspeed. Pp. 451-

455. April, 1919 05

UNIVERSITY OF CALIFORNIA PUBLICATIONS— (Continued)

Vol. 6. 1914-.

1. Parasitic Florideae, by WiUiam Albert Setchell. Pp. 1-34, plates 1-6.

April, 1914 35

2. Fhiftomorula regularis. a Symmetrical Protophyte Related to Coelastnim,'bY

Charles Atwood Kofoid. Pp. 35-40, plate 7. April, 1914 05

3. Variation in Oenothera ovata, by Katherine Layne Erandegee. Pp. 41-50,

plates 8-9, June, 1914 : 10

4. Plantae Mexicanae Purpusianae. VI, by Townshend Stith Erandegee. Pp.

51-77. July, 1914 - 25

5. The Scinaia Assemblage, by William Albert Setchell. Pp. 79-152, plates

10-12. October, 1914 ..., 75

6. Notes on Pacific Coast Algae. I, Pylaiella Postelsiae, n. sp., a New Type in

the Genus Pylaiella, by Carl Skottsberg. Pp. 153-164, plates 17-19. May,
1915 ;... 15

7. New and Noteworthy Calif ornian Plants. II, by Harvey Monroe Hall. Pp.

165-176, plate 20. October, 1915 ....: 15

8. Plantae Mexicanae Purpusianae. VII, by Townshend Stith Erandegee. Pp.

177-197. October, 1915 .- 25

9. Floral Relations among the Galapagos Islands, by A. L. Kroeber. Pp.

199^220. March, 1916 , 20

10. The Comparative Histology of Certain Califomian Boletaceae, by Harry S.

Yates. Pp. 221-274, plates 21-25. February, 1916 .50

11. A Revision of the Tuberales of California, by Helen Margaret GUkey. Pp.

275-356, plates 26-30. March, 1916 80

12. Species Novae vel Minus Cognitae, by T. S. Erandegee. Pp. 357-361. May,

1916 .- .05

13. Plantae Mexicanae Purpusianae. Vin, by Townshend Stith Erandegee.

Pp. 263-375. March, 1917 15

14. New Pacific Coast Marine Algae. I, by Nathaniel Lyon Gardner. Pp. 377-

416, plates 31-35. June, 1917 40

15. An Account of the Mode of Foliar Abscission in Citrus, by Robert W.

Hodgson. Pp. 417-428, 3 text figures. February, 1918 10

16. New Pacific Coast Marine Algae. II, by Nathaniel Lyon Gardner. Pp. 429-

454, plates 36-37. July, 1918 25

17. New Pacific Coast Marine Algae. Ill, by Nathaniel Lyon Gardner. Pp.

455-486, plates 38-41. December, 1918 35

18. New Pacific Coast Marine Algae. IV, by Nathaniel Lyon Gardner. Pp.

487-496, plate 42. January, 1919 15

19. Plantae Mexicanae Purpusianae. IX, by Townshend Stith Erandegee. Pp.

497-504. November, 1919 10

Vol. 7. 1916-.

1. Notes on the Californian Species of TrMlium L. I, A Report of the General

Results of Field and Garden Studies, 1911-1916, by Thomas Harper Good-
speed and Robert Percy Erandt. Pp. 1-24, plates 1-4. October, 1916 25

2. Notes on the Califomian Species ot TriUmm L. II, The Nature and Occur-

rence of Undeveloped Flowers, by Thomas Harper Goodspeed and Robert
Percy Brandt. Pp. 25-38, plates 5-6. October, 1916 15

3. Notes on the Californian Species of Trillium L. Ill, Seasonal Changes in

Trillium Species with Special Reference to the Reproductive Tissues, by
Robert Percy Brandt. Pp. 39-68, plates 7-10. December, 1916 30

4. Notes on the Califomian Species of Trillium L. TV, Teratological Varia-

tions of TriUium sessile var. giganteum H. & A., by Thomas Harper Good-
speed. Pp. 69-100, plates 11-17. January, 1917 30

5. A Preliminary List of the Uredinales of California, by Walter C. Elasdale.

Pp. 101-157. August^ 1919 50

UNIVERSITY or CALirOENIA PUBLICATIONS— (Continued)

6, 7, 8, A Rubber Plant Survey of Western North America. I. Chrysathammis
iiavficosu^ and Its Varieties, by Harvey Monroe Hall. II. Chrysil, a New
Rubber from Cltri/^othamnus nuu.sco.'<us, by Harvey Monroe Kail and
Thomas Harper Goodspeed. III. The Occurrence of Rubber in Certain
West American Shrubs, by Harvey Monroe Hall and Thomas Harper
Goodspeed. Pp. 159-278, plates 18-20, 8 figures in text. November, 1919. 1,25

9. Phycological Contributions. I, by William Albert Setchell and Nathaniel

Lyon Gardner. Pp. 279-324, plates 21-31. April, 1920 50

10. Plantae Mexicanae Purpusianae, X, by Townshend Stith Brandegee. Pp.

325-331, December, 1920 10

VoL 8, 1919-.

1. The Marine Algae of the Pacific Coast of North America. Part L

Myxophyceae, by William Albert Setchell and Nathaniel Lyon Gardner. ,
Pp. 1-138, plates 1-8. November, 1919 'fl.50

2. The Marine Algae of the Pacific Coast of North America. Part II.

Chlorophyceae, by William Albert Setchell and Nathaniel Lyon Gardner.

Pp. 139-374, plates 9-33. July, 1920 2.75

Vol. 9. A Report upon the Boreal Flora of the Sierra Nevada of Calif omia^ by Frank

Jason Smiley. Pp. 1-423, plates 1-7. January, 1921 5.00

UNIVERSITY OF CALIFORNIA PUBLICATIONS

IN

BOTANY

Vol. 7, No. 11, pp. 333-426, plates 32-49 May 16, 1922

PHYCOLOGICAL CONTRIBUTIONS

II TO VI

BY
WILLIAM ALBERT SETCHELL

AND

NATHANIEL LYON GARDNER

UNIVERSITY OF CALIFORNIA PRESS
BERKELEY, CALIFORNIA

UNIVERSITY OF CALIFORNIA PUBLICATIONS

!

't

Note. — The University of California Publications are offered In exchange for the publl- i
cations of learned societies and institutions, universities and libraries. Complete lists of
all the publications of the University will be sent upon request. For sample copies, llata i
of publications and other information, address the MANAGER OF THE UNIVERSITY '
PRESS, BERKELEY, CALIFORNIA, U. S. A. All matter sent in exchange should be
addressed to THE EXCHANQE DEPARTMENT, UNIVERSITY LIBRARY, BERKELEY
CALIFORNIA, U. S. A. ^

WILLIAM WESLEY & SONS, LONDON

Agent for the series in American Archaeology and Ethnology, Botany, Geology,

Physiology, and Zoology.

BOTANY.— W. A. Setchell, Editor. Price per volume, $3.50 (vol. 5, $6.00) ; beginning with
volume 6, $5.00. Volumes I, II, HI, IV, V, and IX completed. Volumes VI, VII,
VIII, and X in progress.

Cited as Univ. Calif. Publ. Bot.
Vol. 5. 1912-1922.

1. Studies in Nicotiana. 1, by William Albert Setchell. Pp. 1-86. December,

1912 $1,25

2. Quantitative Studies of Inheritance in Nicotiana Hybrids. I, by Thomas I

Harper Goodspeed. Pp. 87-168, plates 1-28. December, 1912 1,00 '

3. Quantitative Studies of Inheritance in Nicotiana Hybrids, n, by Thomas

Harper Goodspeed. Pp. 169-188, plates 29-34. January, 1913 ..._ 20

4. On the Partial Sterility of Nicotiana Hybrids njade with N. sylvestris as a

Parent, by Thomas Harper Goodspeed. Pp. 189-198. March, 1913 10

5. Notes on the Germination of Tobacco Seed. I, by Thomas Harper Good-

speed. Pp. 199-222. May, 1913 25

6. Quantitative Studies of Inheritance in Nicotiana Hybrids, III, by Thom.a3

Harper Goodspeed, Pp. 223-231, April, 1915 10

7. Notes on the Germination of Tobacco Seed. II, by Thomas Harper (Jood-

speed. Pp. 233-248. June, 1915 .;,....^ .15

8. Parthenogenesis, Parthenocarpy and Phenospermy in Nicotiana, by Thomas

Harper Goodspeed. Pp. 249-272, plate 35. July, 1915 25

9. On the Partial Sterility of Nicotiana Hybrids made with N. sylvestris as a

Parent. II, by T. H. Goodspeed and A, H, A3rres. Pp, 273-292, plate 36.
October, 1916 20

10. On the Partial Sterility of Nicotiana Hybrids made with N. sylvestris as a

Parent, in. An Account of the Mode of Floral Abscission in the F, Species
Hybrids, by T. H, Goodspeed and J. N. Kendall. Pp. 293-299. November,
1916 - 05

11. The Nature of the Fi Species Hybrids between Nicotiana sylvestris and

Varieties of Nicotiana Tahacum, with Special Reference to the Conception
of Reaction System Contrasts in Heredity, by T, H. Goodspeed and R, E.
Clausen. Pp. 301-346, plates 37-48. January, 1917 45

12. Abscission of Flowers and Fruits in the Solanaceae, with Special Reference

to Nieotiana, by John N. Kendall. Pp, 347-428, 10 text figures, plates 49-

53, March, 1918 85

13. Controlled Pollination in Nicotiana, by Thomas Harper Goodspeed and Pirle

Davidson. Pp. 429-434, August, 1918 10

14. An Apparatus for Flower Measurement, by T. H. Goodspeed and B. B.

Clausen. Pp. 435-437, plate 54, 1 figure in text. September, 1918 .....'.^J^ .05

15. Note on the Effects of Illuminating Gas and Its Constituents in Causing

Abscission of Flowers in Nicotiana and Citrus, by T, H. Goodspeed, J. M.
McGee and B. W. Hodgson. Pp. 439-450. December, 1918 _ 10

16. Notes on the Germination of Tobacco Seed. Ill, Note on the Relation of

Light and Darkness to Germination, by T. Harper Goodspeed. Pp. 451-

455. April, 1919 - 05^

17. Inheritance in Nicotiana Tahacum. I, A Report on the Results of Crossing

Certain Varieties, by William Albert Setchell, Thomas Harper Goodspeed,
and Roy Elwood Clausen. Pp. 457-582, 2 figures in text, plates 55-85.

April, 1922 1.76

Index in press.

UNIVEESITY OF CALIFORNIA PUBLICATIONS— (Contintied)

Vol. 6. 1914-1919.

1. Parasitic Florideae, by William Albert Setchell. Pp. 1-34, plates 1-6.

April, 1914 „ ^ .85

2. Phytomorula regularis, a Symmetrical Protopbyte Related to Coelastrum, by

Charles Atwood Kofoid. Pp. 35-40, plate 7. April, 1914 05

3. Variation in Oenothera ovata, by Katherine Layne Brandegee. Pp. 41-50,

plates 8-9. June, 1914 10

4. Plantae Mexicanae Purpusianae. VI, by Townshend Stitb Brandegee. Pp.

51-77. July, 1914 „ 25

5. Tbe Scinaia Assemblage, by William Albert Setchell. Pp. 79-152, plates

10-16. October, 1914 „ „ .75

6. Notes on Pacific Coast Algae. I, Pylaiella Postelsiae, n. sp., a New Type in

the Genus PylaieUa, by Carl Skottsberg. Pp. 153-164, plates 17-19. May,

1915 _ „ .15

7. New and Noteworthy Californian Plants. II, by Harvey Monroe Hall. Pp.

165-176, plate 20. October, 1915 ...; 15

8. Plantae Mexicanae Purpusianae. VII, by Townshend Stith Brandegee. Pp.

177-197. October, 1915 „ „ ,25

9. Floral Relations among the Galapagos Islands, by A, L. Kroeber. Pp.

199-^220. March, 1916 .20

10. The Comparative Histology of Certain Californian Boletaceae, by Harry S.

Yates. Pp. 221-274, plates 21-25. February, 1916 .50

11. A Revision of the Tuberales of California, by Helen Margaret Gilkey. Pp.

275-356, plates 26-30. March, 1916 _ 80

12. Species Novae vel Minus Cognitae, by T. S. Brandegee. Pp. 357-361. May,

1916 05

13. Plantae Mexicanae Purpusianae. VIII, by Townshend Stith Brandegee.

Pp. 263-375. March, 1917 15

14. New Pacific Coast Marine Algae. I, by Nathaniel Lyon Gardner. Pp. 377-

416, plates 31-35. June, 1917 _ _... .40

15. An Account of the Mode of Foliar Abscission in Citrus, by Robert W.

Hodgson. Pp. 417-428, 3 text figures. February, 1918 10

16. New Pacific Coast Marine Algae. II, by Nathaniel Lyon Gardner. Pp. 429-

454, plates 36-37. July, 1918 25

17. New Pacific Coast Marine Algae, m, by Nathaniel Lyon Gardner, Pp.

455-486, plates 38-41. December, 1918 „ .36

18. New Pacific Coast Marine Algae. IV, by Nathaniel Lyon Gardner. Pp.

487-496, plate 42. January, 1919 15

19. Plantae Mexicanae Purpusianae, IX, by Townshend Stith Brandegee. Pp.

497-504. November, 1919 05

Index in preparation.

Vol. 7. 1916-.

1. Notes on the Californian Species of Trillium L. I, A Report of the General

Results of Field and Garden Studies, 1911-1916, by Thomas Harper Good-
speed and Robert Percy Brandt. Pp. 1-24, plates 1-4. October, 1916 25

2. Notes on the CaUfomian Species of Trillium L. II, The Nature and Occur-

rence of Undeveloped Flowers, by Thomas Harper Goodspeed and Robert
Percy Brandt. Pp. 25-38, plates 5-6. October, 1916 ..._ _. .15

S. Notes on the Californian Species of Trillium L. Ill, Seasonal Changes in
Trillium Species with Special Reference to the Reproductive Tissues, by
Robert Percy Brandt. Pp. 39-68, plates 7-10. December, 1916 „. .30

4. Notes on the Californian Species of Trillium L. IV, Teratological Varia-
tions of Trillium sessile var. giganteum H. & A., by Thomas Harper Good-
speed. Pp. 69-100, plates 11-17. January, 1917 _ „ _ .. .30

UNIVEESiry OF CALIFORNIA PUBLICATIONS— (Continued)

5. A Preliminary List of the Uredinales of California, by Walter C. Blasdale.

Pp. 101-157. August, 1919 JBO

6, 7, 8. A Rubber Plant Survey of Western Nortb America. I. Chry.iothamnus
natuteosus and Its Varieties, by Harvey Monroe Hall. II. Chrysil, a New
Rubber from Chnjsothamnus nauaeosus, by Harvey Monroe Hall and
Thomas Harper Goodspeed. III. The Occurrence of Rubber in Certain
West American Shrubs, by Harvey Monroe Hall and Thomas Harper
Goodspeed. Pp. 159-278, plates 18-20, 8 figures in text. November, 1919. 1.25

9. Phycological Contributions. I, by William Albert Setchell and Nathaniel

Lyon Gardner. Pp. 279-324, plates 21^1. April, 1920 60

10. Plantae Mexicanae Purpusianae. X, by Townshend Stith Brandegee. Pp.

325-331. December, 1920 J.0

11. Phycological Contributions II to VI. New Species of: 11. Myrionema;

m. Compsonema; IV. Hecatonema; V. Pylaiella and Streblonema; VI.
Ectocarpus. By William Albert SetcheU and Nathaniel Lyon Gardner.
Pp. 333-426, plates 32-49. May, 1922 1.50

VcL 8. 1919-.

1. The Marine Algae of the Pacific Coast of North America, Part L

M3rxophyceae, by William Albert Setchell and Nathaniel Lyon Gardner.

Pp. 1-138, plates 1-8. November, 1919 _ $1.50

2. The Marine Algae of the Pacific Coast of North America. Part 11.

Chlorophycea«, by William Albert Setchell and Nathaniel Lyon Gardner.

Pp. 139-374, plates 9-33. July, 1920 „ 2.75

Vol. 9. A Report upon the Boreal Flora of the Sierra Nevada of California, by Frajik

Jason Smiley. Pp. 1-423, plates 1-7. October, 1921 „ 5.00

Vol. 10. 1922-.

1. The Genus Fucus on the Pacific Coast of North America, by Nathaniel Lyon

Gardner. Pp. 1-180, plates 1-60. April, 1922 2.25

UNIVERSITY OF CALIFORNIA PUBLICATIONS

IN

BOTANY

Vol. 7, No. 4, pp. 69-100, plates 11-17 January 17, 1917

NOTES ON THE CALIFORNIAN SPECIES OF

TRILLIUM L.

IV. TERATOLOGIC AL VARIATIONS OF TRILLIUM
SESSILE VAR. GIGANTEUM H. & A.

BY

THOMAS HARPER GOODSPEED

UNIVERSITY OF CALIFORNIA PRESS
BERKELEY

UNIVERSITY OF CALIFORNIA PUBLICATIONS

Note. — The University of California Publications are offered in exchange for the pnbll-
catlonB of learned societies and Institutious, universities and libraries. Complete lists of
all the publications of the University will be sent upon request. For sample copies, lists of
pabiicatlons and other iuformation, address the Manager of the University Press, Berkeley,
California, U. 8. A. All matter sent in exchange should be addressed to The &zchang«
Department, University Library, Berkeley, Caliloroia, U. S. A.

OTTO HARRASOWITZ
LEIPZIG
Agent for the series in American Arch-
aeology and Ethnology, Classical Philology,
Education, Modern Philology, Philosophy,
Psychology.

E. FEIEDLANDEE & SOHN
BERLIN
Agent for the series in American Arch-
aeology and Ethnology, Botany, Geography,
Geology, Mathematics, Pathology, Physi-
ology, Zoology, and Memoirs.

BOTANY.— W. A. Setchell, Editor. Price per volume, $3.50. Volumes I (pp. 418), II (pp
360), m (pp. 400), IV (pp. 379), completed. Vols. V and VI in progreaa.

Cited as Univ. Calif. PubL Bot.

VoL 1. 1. A Botanical Survey of San Jacinto Mountain, by Harvey Monroe

HalL Pp. 1-140; plates 1-14. Jane, 1902 11.00

2. Two new Ascomycetous Fungi Parasitic on Marine Alj^ae, by Minnie

Eoed. Pp. 141-164; plates 15-16. November, 1902 .25

S. Algae of Northwestern America, by William Albert Setchell and Na-
thaniel Lyon Gardner. Pp. 165-418; plates 17-27. March, 1903 8.2S

VoL 2. 1. A Eeview of Califomian Polemoniaceae, by Jeasie Milliken. Pp. 1-

71; plates 1-11. May, 1904 „ „ .76

2. Contributions to Cytological Technique, by W. J. V. Osterhout. Pp.

73-90; 5 text-figures. June, 1904 „ .25

5. Limu, by William Albert SetchelL Pp. 91-113. April, 1905 .25

4. Post-Embryonal Stages of the Laminarlaceao, by William Albert

Setchell. Pp. 115-138; plates 13-14. April, 1905 „ ~ J26

6. Eegeneration among Kelps, by William Albert SetchelL Pp. 139-168;

plates 15-17. July, 1905 „ _ .80

6. A New Genus of Aiscomycetous Fungi, by Nathaniel Lyon Gardner.

Pp. 169-180; plate 18. July, 1905 _.. 06

7. Teratology in the Flowers of some Califomian WiUows, by William

Warner Mott. Pp. 181-226; plates 16-20. December, 1905 .60

8. 9, 10, 11. (In one cover.) The Resistance of Certain Marine Algae to

Changes in Osmotic Pressure and Temperaturs. The Edle of 0»-
motic Pressure in Marine Plants. On the Importance of Physiolog-
ically Balanced Solutions for Plants. The Antitoxic Action of
Potassium on Magnesium. By W. J. V. Osterhout. Pp. 227-238.
March, 1906 ._ _.._ .26

12. Cjrtologlcal Studies In Cyanophyceae, by Nathaniel Lyon Gardner.

Pp. 237-296; plates 21-26. November, 1906 „ _ 1.00

IS. On a Small Collection of Mosses from Alaska, by J. Cardot and T.

Th6riot. Pp. 297-S08; plates 27-28. December, 1906 JO

14. Some Unreported Alaskan Sphagna, togsther with a Sunw.ary of the

Cryptogamlc Work of the University of California Botanical Ex-
pedition to Alaska In 1899, by William Albert SetchelL Pp. 309-
315. September, 1907 _ _ M

15. On Nutrient and Balanced Solutions, by W. J. V. Osterhout. Pp. 817-

313. October, 1907 ..... .08

16. A Synopsis of the North American Godetlas, by Willis T-^nn Jepson.

Pp. 319-354; plate 29. December, 1907 40

Index, pp. 355-360.

VoL ». 1907-1909.

L Composltae of Southern California, by Harvey Monroe Hall. Pp. 1-

302; plates 1-3, with a map. December, 1907 8.00

5. The Origin, Structure, and Function of the Polar Cap* In FmQacina

ampleriiYiulis Nutt., by H. D. Densmoro. Pp. 803-330; plates 4-8.

December. 1908 _ _ _ .55

t. 4. (In one cover.) The Value of Sodlnm to Plants by Season of Its
Protective Action. On the Effects of Certain Poisonous Gases on
Plants. By W. J. V. Osterhout. Pp. 331-340. Juub. 1908 JO

6. Contributions to the Knowledge of the California Species of Omsta- ^

ceoTis Corallines. I. by Maurice Barstow Nichols. Pp. S41-S48; \"v
plate 9. December, 1908 M

UNTVEaSITT OF CAUTOSNIA PTTBUOATIONS— (Continued)

6. Contrtbntlons te the Knowledge of the California Species of Cmsta-
ceous Corallines, n. by Maurice Barstow Nichols. Pp. S49-S70;

plates 10-13. April. 1909 ._ _ J5

?, New Chlorophyceae from California, by Nathaniel Lyon Gardner. Pp.

371-375; plate 14. April. 1909 _ ^ JLO

8. Flantae Mexicanae Purpusianae, I, by T. S. Brandegea. Pp. 377-396.

May, 1909 , „.„...... „ .15

Index, pp. S97-400,
Vol 4. 1910-1912.

1. Stuuies In Ornamental Treea and Shmba, by Harvey Mtmroe HaU. Pp.

1-74; plates 1-11; 15 text-fig\ires. March, 1910 „ ^ .75

2. GracilartopiiUa, a New Parasite on GrartUirio confervoides, by Harriet

L. V7ilson. Pp. 75-84; plates 12-13. May, 1910 „ „ .10

8. Plantae Mexicanae Pnrpuaianae, II, by T. 8. Brandegee. Pp. 85-95.

May. 1910 _ _.... „ AO

4. Leuvenia, a New Genua of Flagellates, by N. L. Gardner. Pp. 97-106;

plate 14. May, 1910 „... , JIO

5. The Genus Sphaerosoma, by William Albert Setchell. Pp. 107-120;

plate 15. May, 1910 „... .15

6. Variations in Nrcloar Extrcsion Among the Fncaceae, by Nathaniel

Lyon Gardner. Pp. 121 136; plates 16-17. Angnst, 1910 4.8

7. The Nature of the Carpostomes In the Cyatocarp of Ahnfeldtut gigarti-

no'des, by Ada Sara McFadden. Pp. 137-142; plate 18. Febmary,
1911 .' .....„..:. ._.. .05

8. On a Colacodasya from Southern California, bj Mabel Bffle McFadden.

Pp. 143 150; plate 19. February, 1911 _ _ _ .08

9. Fmctiflcatlon of Macrocystia, by Edna Juanlta Hoffman. Pp. 151-158;

plate 20. February, 1911 _ _ .05

10. Erpthrophytlvm drlesxrrioides J. Ag., by Wilfred Charles Twlas. Pp.

159-176; plates 21-24. March, 1911 „ 0.5

11. Plantae Mexicanae Purpusianae, III, by T. S. Brandeged. Pp. 177-194.

July, 191 1 ....„.:_ .._„ — 45

12. New and Noteworthy Calif omian Plants, I, by Harvey Monroe HalL

Pp. 195-208. March, 1912 ..._.. , 45

18. Die Hydrophyllaceen der Sierra Nevada, by August Brand. Pp. 209-

227. March, 1912 _ J20

14. Algae Novae et Minns Cognitae. I, by William Albert SetchelL Pp.

229-268; plates 25-31. May, 1912 _ - - .40

15. Plantae Mexicanae Purpnaianae, IV, by Townshend Stith Brandegee.

Pp. 269-281. Jane, 1912 ..-, ... J.8

16. Comparative Development of the Cystocarps of Avtithnmnion and

Prioniti^. by Lyman Luther Daines. Pp. 283-302; plates 32-34.
March, 1913 - „ „ SX)

17. Fungus Galls on Cystoseira and Halidrys. by Lulu May Estee. Pp. SOS-

SIB; plate 35. March, 1913 „ - .10

18. New Fucaceac, by Nathaniel Lyon Gardner. Pp. 317-374; plates 36-

53. April, 1913 - .- 76

19. Plantae Mexicanae Purpusianae, V, by Townshend Stith Brandegee.

Pp. 375-388. June. 1913 A8

Index, pp. 389-397.
Vol 5. 19 12-.

1. Studies in Nirotiaim, I, by William Albert Setchell. Pp. 1-86. De-

cember, 1912 ...~ _. _ - 1.2B

2. Quantitative Studies of Inheritance in NicoHana Hybrids, I, by Thomaa

Harper Good speed. Pp. 87-168. December, 1912 1.00

5. Quantitative Studies of Inheritance in Nicotiana Hybrids, n, by

Thomas Harper Goodspeed. Pp. 169-188. January, 1913 _. .20

4. On the Partial Sterility of Nicotiana Hybrids made with ff. Sylvesiris
as a Parent, by Thomas Harper Goodspeed. Pp. 189-198. March,
1913 _ - 40

6. Notes on the Germination of Tobacco Seed, I, by Thomas Harper Good-

speed. Pp. 199-222. May, 1913 _ .88

6. Quantitative Studies of Inheritance in Nicotiana Hybrids, m, by

Thomas Harper Goodspeed. Pp. 223-231. April, 1915 „ .10

7. Notes on the Germination of Tobacco Seed, II, by Thomas Harper

Goodspeed. Pp. 233-248. June, 1915 - .15

8. Parthenogenesis, Parthenocarpy and Phenospermy in Nicotiana, by

Thomas Harper Goodspeed. Pp. 249-272, plate 35. July, 1915 ~. J2S

UNTVEESITT OF CALIFORNIA PUBLICATIONS— (Oon tinned)

9. On the Partial Sterility of Nicotiana Hybrids made with N. sijlvestris
as a Parent. II, by T. H. Goodspeed aud A. H. Ayres. Pp. 273-292,
plate 36. October, 1916 „ JZO

10. On the Partial Sterility of Nicotiana Hybrids made with A^. sylvestris

as a Parent. III. An Account of the Mode of Floral Abscission in
the F, Species Hybrids, by T. H. Goodspeed and J. N. KendalL
Pp. 293-299. November, 1916 05

11. The Nature of the F, Species Hybrids between Nicotiana sylvestris

and Varieties of Nicotiana tabucum, with Special Reference to the
Conception of Reaction System Contrasts in Heredity, by T. H.
Goodspeed and R. E, Clausen. Pp. 301-346, plates 37-48. Janu-
ary, 1917 4S

Vol 6. 1914-

1. Parasitic Florideae, I, by William Albert Setchell. Pp. 1-34, plates

1 6. April, 1914 „ _ JS6

2. Phytomomla Tegulnris, a Symmetrical Protophyte Related to Coelag-

trum, by Charles Atwood Kofoid. Pp. 35-40, plate 7. April, 1914, .06
S. Variation in Oenothera ovata, by Katherine Layne Brandegee. Pp. 41-

50, plates 8-9. June, 1914 _ J.0

4. Plantae Mexicanae Piirpusianae, VI, by Townshend Stith Brandegee.

Pp. 51-77. July, 1914 „ J2S

5. The Scinain Assemblage, by William Albert Setchell. Pp. 79-152,

plates 10-16. October, 1914 „ „ _ .76

6. Notes on Pacific Coast Algae. I. Pylaiella Postelsiae, n. sp., a New

Type In the Genus Pylaiella, by Carl Skottsberg. Pp. 153-164, plates
17-19. May. 1915 _ .15

7. New and Noteworthy Califomian Plants, II, by Harvey Monroe HalL

Pp. 165-176, plate 20. October, 1915 „ „ .15

8. Plantae Mexicanae Purpnsianae VII, by Townshend Stith Brandegee.

Pp. 177-197. October, 1915 .25

9. Floral Relations Among the Galapagos Islands, by A. L. Ejroeber.

Pp. 199-220. March, 1916 „ .20

10. The Comparative Histology of Certain Califomian Boletaceae, by

Harry S. Yates. Pp. 221-274, plates 21-25. February, 1916 .60

11. A Revision of the Tuberales of California, by Helen Margaret Gilkey.

Pp. 275-356, plates 26-30. March, 1916 .80

12. Species Novae vel Minus Cognltae, uy T. S. Brandegee. Pp. 357-361.

May, 1916 _ 05

VoL 7. 1916-

1. Notes on the Califomian Species of Trillium L. I. A Report of the

General Results of Field and Garden Studies, 1911-1916, by Thomas
Harper Goodspeed and Robert Percy Brandt. Pp. 1-24, plates 1-4.
October, 1916 .25

2. Notes on the Califomian Species of Trillium L. II. The Nature and

Occurrence of Undeveloped Flowers, by Thomas Harper Goodspeed

and Robert Percy Brandt. Pp. 25-38, plates 5-6. October, 1916 ... J.5

3. Notes on the Califomian Species of Trillium L. HI. Seasonal Changes

in Trillium Species with Special Reference to the Reproductive Tis-
sues, by Robert Percy Brandt. Pp. 39-68, plates 7-10. December,
1916 30

4. Notes on the Califomian Species of Trillium L. IV. Teratologlcal

Variations of Trillium sessile var. qiganteum H. & A., by Thomas
Harper Goodspeed. Pp. 69-100, plates 11-17. January, 1917 30

UNIVERSITY OF CALIFORNIA PUBLICATIONS

IN

BOTANY

Vol. 7, No. 5, pp. 101-157 August 14, J919

A PRELIMINARY LIST OF THE
UREDINALES OF CALIFORNIA

BY
WALTER C. BLASDALE

UNIVERSITY OF CALIFORNIA PRESS
BERKELEY

UNIVERSITY OF CALIFORNIA PUBLICATIONS

Note. — The University of California Publications are offered in exchange for the publi-
cations of learned societies and institutions, universities and libraries. Complete lists of
all the publications of the University will be sent upon request. For sample copies, lists
of publications and other information, address the MANAGER OF THE UNIVERSITY
PRESS, BERKELEY, CALIFORNIA, U. S. A. AU matter sent in exchange should be
addressed to THE EXCHANGE DEPARTMENT, UlJnVERSITY LIBRARY, BERKELEY,
CALIFORNIA, U. S, A.

WILLIAM WESLEY & SONS, LONDON

Agent for the series in American Archaeology and Ethnology, Botany, Geology,

Physiology, and Zoology.

BOTANY. — W. A. Setchell, Editor. Price per volume, ?8.50; beginning with vol. 5, $5.00.
Volumes I (pp. 418), II (pp. 360), III (pp. 400), IV (pp. 379), completed. Volumes
V, VI and VII in progress.

Cited as Univ. Calif. Publ. Bot.

Volume 1, 1902-1903, 418 pages, with 27 plates $3.50

Volume 2, 19C4-1907, 360 pages, with 29 plates 3.50

Volume 3, 1907-1909, 400 pages, with 14 plates 3.50

Vol. 4. 1910-1912.

1. Studies in Ornamental Trees and Shrubs, by Harvey Monroe Hall. Pp. 1-74,

plates 1-11, 15 text figures. March, 1910 75

2. Gracilariophila, a New Parasite on Graoilaria conferv aides, by Harriet L.

Wilson. Pp. 75-84, plates 12-13. May, 1910 „ 10

3. Plantae Mexlcanae Purpusianae, II, by T. S. Brandegee. Pp. 85-95. May,

1910 ; 10

4. Leuvenia, a New Genus of Flagellates, by N. L. Gardner. Pp. 97-106, plate

14. May, 1910 „ .10

5. The Genus Sphaerosoma, b William Albert Setchell. Pp. 107-120, plate

15. May, 1910 15

6. Variations in Nuclear Extrusion Among the Fucaceae, by Nathaniel Lyon

Gardner. Pp. 121-136, plates 16-17. August, 1910 „ 15

7. The Nature of the Carpostomes in the Cystocarp of Ahnfeldtia gigartinoides,

by Ada Sara McFadden. Pp. 137-142, plate 18. February, 1911 05

8. On a Colacodasya from Southern California, by Mabel Efi&e McFadden.

Pp. 143-150, plate 19. February, 1911 05

9. Fructification of Macrocystis, by Edna Juanita Hoffman. Pp. 151-158, plate

20. February 1911 05

10. Erythrophyllum delesserioides J. Ag., by Wilfred Charles Twiss. Pp. 159-

176, plates 21-24. March, 1911 15

11. Plantae Mexicanae Purpusianae, m, by T. S. Brandegee. Pp. 177-194.

July, 1911 __ 15

12. New and Noteworthy California Plants, I, by Harvey Monroe Hall. Pp.

195-208. March, 1912 ; 15

13. Die Hydrophyllaceen der Sierra Nevada, by August Brand. Pp. 209-227.

March, 1912 20

14. Algae Novae et Minus Cognitae, I, by William Albert Setchell. Pp. 229-268,

plates 25-31. May, 1912 40

15. Plantae Mexicanae Purpusianae, IV, by Townshend Stith Brandegee. Pp.

269-281. June, 1912 15

16. Comparative Development of the Cystocarps of Antiihamnion and Prionitis,

by Lyman Luther Daines. Pp. 283-302, plates 32-34. March, 1913 20

17. Fungus Galls on Cystoseira and Ealidrys, by Lulu May Estee. Pp. 305-316,

plate 35. March, 1913 .....^: .„;„..„ „ 10

18. New Fucaceae, by Nathaniel Lyon Gardner. Pp. 317-374, plates 36-53.

April, 1913 75

19. Plantae Mexlcanao Purpusianae, V, by Townshend Stith Brandegee. Pp.

375-388. June, 1913 „ _ 15

Index, pp. 389-397.

UNIVEESITY OF CALIFORNIA PUBLICATIONS— (Continued)

Vol. 5. 1912-.

1. Studies in Nicotiana, I, Ijy William Albert Setchell. Pp. 1-86. December,

1912 ' 1.25

2. Quantitative Studies of Inheritance in Nicotiana Hybrids, I, by Thomas

Harper Goodspeed. Pp. 87-168. December, 1912 1.00

3. Quantitative Studies of Inheritance in Nicotiana Hybrids, II, by Thomas

Harper Goodspeed. Pp. 169-188. January, 1913 20

4. On the Partial Sterility of Nicotiana Hybrids made with N. sylvestris as a

Parent, by Thomas Harper Goodspeed. Pp. 189-198. March, 1913 10

5. Notes on the Germination of Tobacco Seed, I, by Thomas Harper Goodspeed.

Pp. 199-222. May, 1913 25

6. Quantitative Studies of Inheritance in Nicotiana Hybrids, III, by Thomas

Harper Goodspeed. Pp. 223-231. April, 1915 10

7. Notes on the Germination of Tobacco Seed, II, by Thomas Harper Good-

speed. Pp. 233-248. June, 1915 15

8. Parthenogenesis, Parthenocarpy and Phenospermy in Nicotiana, by Thomas

Harper Goodspeed. Pp. 249-272, plate 35. July, 1915 25

9. On the Partial Sterility of Nicotiarux Hybrids made with N. sylvestris as a

Parent, II, by T. H. Goodspeed and A. H. Ayres. Pp. 273-292, plate 36.
October, 1916 „... .20

10. On the Partial Sterility of Nicotiana Hybrids made with N. sylvestris as a

Parent, III, An Account of the Mode of Floral Abscission in the Fj Species
Hybrids, by T. H. Goodspeed and J. N. Kendall. Pp. 293-299. November,
1916 _ „ 05

11. The Nature of the Fj Species Hybrids between Nicotiana sylvestris and

Varieties of Nicotiana Tdbacum, with Special Reference to the Conception
of Eeaction System Contrasts in Heredity, by T. H. Goodspeed and E. E.
Clausen. Pp. 301-346, plates 37-48. January, 1917 45

12. Abscission of Flowers and Fruits in the Solanaceae, with Special Reference

to Nicotiana, by John N. Kendall. Pp. 347-428, 10 test figures, plates 49-

53. March, 1918 85

13. Controlled Pollination in Nicotiana, by Thomas Harper Goodspeed and Pirie

Davidson. Pp. 429-434. August, 1918 „. 10

14. An Apparatus for Flower Measurement, by T. H. Goodspeed and R. E.

Clausen. Pp. 435-437, plate 54, 1 figure in text. September 25, 1918 .05

15. Note on the Effects of Illuminating Gas and its Constituents in Causing

Abscission of Flowers in Nicotiana and Citrus, by T. H. Goodspeed, J. M.
McG-ee and R. W. Hodgson. Pp. 439-450. December, 1918 :...: .15

16. Notes on the Germination of Tobacco Seed, III. Note on the Relation of

Light and Darkness to Germination, by T. Harper Goodspeed. Pp. 451-

455. April, 1919 05

Vol. 6. 1914-.

1. Parasitic Florideae, by William Albert Setchell. Pp. 1-34, plates 1-6.

April, 1914 „. .35

2. PJiytomorula regnlaris, a Symmetrical Protophyte Related to Coelastrum, by

Charles Atwood Kofoid. Pp. 35-40, plate 7. April, 1914 05

3. Variation in Oenothera O'vata, by Katherine Lasme Brandegee. Pp. 41-50,

plates 8-9. June, 1914 10

4. Plantae Mexicanae Purpusianae, VI, by Townshend Stith Brandegee. Pp.

51-77. July, 1914 25

5. The Scinaia Assemblage, by William Albert Setchell. Pp. 79-152, plates

10-12. October, 1914 .75

6. Notes on Pacific Coast Algae, I, Pylaiella Postelsiae, n. sp., a, New Type in

the Genus Pylaiella, by Carl Skottsberg. Pp. 153-164, plates 17-19. May,
1915 15

7. New and Noteworthy Califomian Plants, II, by Harvey Monroe Hall. Pp.

165-176, plate 20. October, 1915 15

8. Plantae Mexicanae Purpusianae, VII, by Townshend Stith Brandegee. Pp.

177-197. October, 1915 _ 25

UNIVERSITY OF CALIFORNIA PUBLICATIONS— (Continued)

9. Floral Relations Among the Galapagos Islands, by A. L. Kroeber. Pp.

199-220. March, 1916 _ 20

10. The Comparative Histology of Certain Califomian Boletaceae, by Harry S.

Yates. Pp. 221-274, plates 21-25. February, 1916 .50

11. A Revision of the Tuberales of California, by Helen Margaret Gilkey. Pp.

275-356, plates 26-30. March, 1916 80

12. Species Novae vel Dlinus Cognitae, by T. S. Brandegee. Pp. 357-361. May,

1916 05

13. Plantae Mexicanae Purpusianae, Vin, by Townshend Stith. Brandegee. Pp.

363-375. March, 1917 .;.:.....*... 15

14. New Pacific Coast Marine Algae, I, by Nathaniel Lyon Gardner. Pp. 377-

416, plates 31-35. June, 1917 40

15. An Account of the Mode of Foliar Abscission in Citrus, by Robert W.

Hodgson. Pp. 417-428, 3 text figures. February, 1918 10

16. New Pacific Coast Marine Algae, II, by Nathaniel Lyon Gardner. Pp. 429-

454, plates 36-37. July, 1918 , : 25

17. New Pacific Coast Marine Algae, HI, by Nathaniel Lyon Gardner. Pp. 455-

486, plates 38-41. December, 1918 „ „ 85

18. New Pacific Coast Marine Algae IV, by Nathaniel Lyon Gardner. Pp. 487-

496, plate 42. January, 1919 15

Vol. 7. 1916-.

1. Notes on the Califomian Species of Trillnim L. I, A Report of the General

Results of Field and Garden Studies, 1911-1916, by Thomas Harper Good-
speed and Robert Percy Brandt. Pp. 1-24, plates 1-4. October, 1916 25

2. Notes on the Califomian Species of Trillium L. II, The Nature and Occur-

rence of Undeveloped Flowers, by Thomas Harper Goodspeed and Robert
Percy Brandt. Pp. 25-38, plates 5-6. October, 1916 15

3. Notes on the Califomian Species of Trillium L. Ill, Seasonal Changes in

Trillium Species with Special Reference to the Reproductive Tissues, by
Robert Percy Brandt. Pp. 39-68, plates 7-10. December, 1916 30

4. Notes on the Califomian Species of Trillium L. IV, Teratological Varia-

tions of Trillium sessile var. giganteum H. & A,, by Thomas Harper Good-
speed. Pp. 69-100, plates 11-17. January, 1917 .30

5. A Preliminary List of the Uredinales of California, by Walter C. Blasdale.

Pp. 101-157. August, 1919 50

UNIVERSITY OF CALIFORNIA PUBLICATIONS

IN

BOTANY

Vol. 7, No. 2, pp. 25-38, plates 5-6 October 4, 1916

NOTES ON THE CALIFORNIAN SPECIES OF

TRILLIUM L.

II. THE NATURE AND OCCURRENCE OF
UNDEVELOPED FLOWERS

BY

THOMAS HARPER GOODSPEED and ROBERT PERCY BRANDT

UNIVERSITY OF CALIFORNIA PRESS
BERKELEY

UNIVERSITY OF CALIFORNIA PUBLICATIONS

Note. — The University of California Publications are offered in exchange for the pabll-
catlona of learned societies and institutions, universities and libraries. Complete lists of
all the publications of the University will be sent upon request. For sample copies, lists of
publications and other information, address the Manager of the University Press, Berkeley,
California, U. S. A. All matter sent in exchange should be addressed to The Exchange
Department, University Library, Berkeley, California, U. S. A.

OTTO HAEEASOWITZ E. FEIEDLANDER & SOHN
LEIPZIG BERLIN
Agent for the series in American Arch- Agent for the series in American Arch-
aeology and Ethnology, Classical Philology, aeology and Ethnology, Botany, Geography,
Education, Modern Philology, Philosophy, Geology, Mathematics, Pathology, Phyai-
Psychology. ology. Zoology, and Memoirs.

BOTANY.—W. A. Betchell, Editor. Price per volume, ^3.50. Volumes I (pp. 418), It (pi^
360), III (pp. 400), IV (pp. 379), completed. Vols. V and VI in progress.

Cited as Univ. Calif. PubL Bot.

VoL L 1. A Botanical Survey of San Jacinto Mountain, by Harvey Monroe

EaU. Pp. 1-140; plates 1-14. June, 1902 fl.00

2. Two new Ascomycetoua Fungi Parasitic on Marine Algae, by Minnie

Eoed. Pp. 141-164; plates 15-16. November, 1902 _. J25

3. Algae of Northwestern America, by William Albert Setchell and Na-

thaniel Lyon Gardner. Pp. 165-418; plates 17-27. March, 1903 — 2.2S

VoL jS. 1. A Eevlew of Oalifomlan Polemoniaceao, by Jossle Milliken, Pp. 1-

71; plates 1-11. May, 1904 .76

2. Contributiona to Cytological Technique, by W. J. V. Osterhout. Pp.

73-90; 5 text-figures. June, 1904 :. .25

3. Llmu, by William Albert Setchell. Pp. 91-113. April, 1905 — .25

4. Post-Embryonal Stages of the Laminariaceaa, by William Albert

SetcheU. Pp. 115-138; plates 13-14. April, 1905 .._ - — .26

5. Regeneration among Kelps, by William Albert Setchell. Pp. 139-168;

plates 15-17. July, 1905 — .30

6. A New Genus of Ascomycetoua Fungi, by Nathaniel Lyon Gardner.

Pp. 169-180; plate 18. July, 1905 _ - 15

7. Teratology in the Flowers of Bome Caltfomian Willows, by WlUiam

Warner Mott. Pp. 181-226; plates 16-20. December, 1905 _ .50

8. 9, 10, 11. (In one cover.) The Resistance of Certain Marino Algae to

Changes in Osmotic Pressure and Temperature. The E61e of Os-
motic Pressure in Marine Plants. On the Importance of Physiolog-
ically Balanced Solutions for Plants. The Antitoxic Action of
Potassium on Magnesium. By W. J. V. Osterhout. Pp. 227-236.

March, 1906 . .28

12. Cytological Studies in Cyanophyccae, by Nathaniel Lyon Gardner.

Pp. 237-206; plates 21-26. November, 1906 LOO

15. On a Small Collection of Mosses from Alaska, by J. Cardot and T.

Th6riot. Pp. 297-308; plates 27-28. December, 1906 M

14. Some Unreported Alaskan Sphagna, together with a Summary of the
Oryptogamlc Work of the University of Califomia Botanical Ex-
pedition to Alaska in 1899, by William Albert SetchelL Pp. 309-
315. September, 1907 .06

16. On Nutrient and Balanced Solutions, by W. J. V. Osterhout. Pp. 817-

318. October, 1907 - — .06

16. A Synopsis of the North American Godetias, by WlUifl Linn Jepson.

Pp. S19-364; plate 29. December, 1907 M

Index, pp. 355-360.

VoL 9. 1907-1909.

1. Compositao of Southern Califomia, by Harvey Monroe Hall. Pp. 1-

302; plates 1-3, with a map. December, 1907 8.00

t. The Origin, Structure, and Function of the Polar Caps La SmiUicina

amplexieaulis Nutt., by H. D. Densmorc. Pp. 303-330; plates 4-8.

December, 1908 _ 38

t. 4. (In one cover.) The Value of Sodium to Plants by Eeaoon of Its

Protective Action. On the Effects of Certain Poisonous Gases on

Plants. By W. J. V. Osterhout. Pp. 831-340. June, 1908 .10

B. Contributions to the Knowledge of the Califomia Species of Oruata-

cooBS Corallines. L by Maurice Barstow Nlchola. Pp. 341-348;

plat* 9. December, 1908 .1*

XJNIVBSSITT OP OAUFOENIA PUBLICATIONS— (Oontinuod)

6. Contribntioiui to the Knowledge of the California Species of Onut»-

ceous Coralilncs. U. by Maurice Barstow Nichols. Pp. S49-370;
plates 10-13. April, 1909 45

7. New Ohlorophyceae from California, "by Nathaniel Lyon Gardner. Pp.

371-375; plate 14. April, 1909 _ .„_ .10

8. Plantae Mexicanae Purpusianae, I, by T. S. Brandegee. Pp. 377-396.

May, 1909 _. ^ ,_... .15

Index, pp. S97-400.
Vol 4. 1910-1912.

1. Studies in Ornamental Trees and Shrubs, by Harrey Monroe HalL Pp.

i-7i; plates 1-11; 15 text-figures. March, 1910 75

2. Graeilarlophlla, a New Parasite on Gracilaria confervoides, by Harriet

L. Wilson. Pp. 75-84; plates 12-13. May, 1910 „ .10

8. Plantae Mexicanae Purpusianae, II, by T. S. Brandegee. Pp. 85-95.

May, 1910 ._ _ _... . JO

4. Leuvenia, a New Genus of Flagellates, by N. L. Gardner. Pp. 97-106;

plate 14. May, 1910 SO

5. The Genus Sphaerosoma, by William Albert Setchell. Pp. 107-120;

plate 15. May, 1910 _ .16

6. Variations in Nuclear Extrusion Among the Fncaceae, by Nathaniel

Lyon Gardner. Pp. 121-136; plates 16-17. August, 1910 J.6

7. The Nature of the Carpostomes in the Cystocarp of Ahnfeldtia gigarii-

noides, by Ada Sara McFadden. Pp. 137-142; plate 18. February,
1911 „ „ .05

8. On a Colacodasya from Southern Callfcmla, by Mabel Effle McFadden.

Pp. 143-150; plate 19. February, 1911 _....^.„ _ > .05

9. Fructification of Macrocystis, by Edna Juanita Hoffman. Pp. 151-158;

plate 20. February, 1911 - _ - .06

10. ErythropTiyllum delesserioides J. Ag., by Wilfred Oharleg Twlss. Pp.

159-176; plates 21-24. March, 1911 J.5

11. Plantae Mexicanae Purptmianae, HI, by T. S. Brandegee. Pp. 177-194,

July, 1911 _ _ .16

12. New and Noteworthy Califomiaa Plants, I, by Harvey Monroe HalL

Pp. 195-208. March, 1912 ._ _ .15

IS. Die Hydrophyliaceen der Sierra Nevada, by August Brand. Pp. 209-

227. March, 1912 - _ .20

14. Algae Novae et Minus Cognltae, I, by William Albert SetcheU. Pp.

229-268; plates 25-31. May, 1912 .„ .-... — .40

15. Plantae Mexicanae Purpusianae, IV, by Townshend Stith Brandegee.

Pp. 269-281. June, 1912 .15

16. Comparative Development of the Cystocarps of AntitJuimnion and

Prion tf is, by Lyman Luther Daines. Pp. 283-302; plates 32-34.
March, 1913 „ ~ .20

17. Fungus Galls on Cystoseira and Ealidrys. by Lulu May Estee. Pp. 305-

316; plate 35. March, 1913 . „ 10

18. New Fucaceae, by Nathaniel Lyon Gardner. Pp. 317-374; plates 36-

53. April, 1913 - „ - — .75

19. Plantae Mexicanae Purpusianae, V, by Townshend Stith Brandegee.

Pp. 375-388. June, 1913 _ - .16

Index, pp. 389-397.
Vol. 5. 1912-.

1. Stiidies in Nicotiana, I, by William Albert Setchell. Pp. 1-86. De-

cember, 1912 „ _ « - « — 1.25

2. Quantitative Studies of Inheritance in Nicotiana Hybrids, I, by Thomas

Harper Goodspeed. Pp. 87-168. December, 1912 1.00

S. Quantitative Studies of Inheritance in Nicotiana Hybrids, n, by

Thomas Harper Goodspeed. Pp. 169-188. January, 1913 _ .20

4. On the Partial Sterility of Nicotiana Hybrids made with N. Sylvestris

as a Parent, by Thomas Harper Goodspeed. Pp. 189-198. March,

1913 _ _ ~ _ - - .10

5. Notes on the Germination of Tobacco Seed, I, by Thomas Harper Good-

speed. Pp. 199-222. May, 1913 .25

6. Quantitative Studies of Inheritance in Nicotiana Hybrids, m, by

Thomas Harper Goodspeed. Pp. 223-231. April, 1915 10

7. Notes on the Germination of Tobacco Seed, n, by Thomas 'Harper

Goodspeed. Pp. 233-248. June, 1915 15

8. Parthenogenesis, Parthenocarpy and Phenospermy in Nicotiana, by

Thomas Harper Goodspeed. Pp. 249-272, plate 35. July, 1915 25

9. On the Partial Sterility of Nicotiana Hybrids made with A'', sylvestris

as a Parent. II, by T. H. Goodspeed and A. H. Ayres. Pp. 273-292,
plate 36. October, 1916 20

UNIVEESITY OF OALIFOENIA PUBLICATIONS— (Continued)

Vol. 6. 19 U-

1. Parasitic Florideae, I, by WlUiam Albert Eetchell. Pp. 1-34, plates

1^. April, 1914 „ „ JS5

2. Phytiimorula rcfiuJaris, a Symmetrical Protopbyte Related to Codas-

\rum, by Charles Atwood Kofoid. Pp. 35-40, plate 7. April, 1914. .06

3. Variation in Oenothera ovata, by Katharine Layne Brandegee. Pp. 41-

50, plates 8-9. June, 1914 „ JLO

4. Plantae Mexicanae Purpusianae, VI, by Townshend Stith Brandegee.

Pp. 51-77. July, 1914 „ _ sa

5. The Scinaia Assemblage, by William Albert Setchell, Pp. 79-152,

plates 10-16. October, 1914 7B

6. Notes on Pacific Coast Algae. I. Tylaiella Postelsiae, n. sp., a New

Type in the Genus Pylaiella, by Carl Skottsberg. Pp. 153-164, plates
17-19. May, 1915 „ 15

7. New and Noteworthy Califomian Plants, II, by Harvey Monroe Hall.

Pp. 165-176, plate 20. October, 1915 15

8. Plantae Mexicanae Purpusianae VII, by Townshend Stith Brandegee.

Pp. 177-197. October, 1915 _,.:......^..,_. „. .25

9. Floral Relations Among the Galapagos Islands, by A. L. Kroeber.

Pp. 199-220. March, 1916 20

10. The Comparative Histology of Certain Califomian Boletaceae, by

Harry S. Yates. Pp. 221-274, plates 21-25. February, 1916 50

11. A Revision of the Tuberales of California, by Helen Margaret Gilkey.

Pp. 275-356, plates 26-30. March, 1916 80

12. Species Novae vel Minus Cognitae, by T. S. Brandegee. Pp. 357-361.

May, 1916 05

Vol. 7. 1916-

1. Notes on the Califomian Species of Trillium L. I. A Report of the

General Results of Field and Garden Studies, 1911-1916, by Thomas
Harper Goodspeed and Robert Percy Brandt. Pp. 1-24, plates 1-4.
October, 1916 25

2. Notes on the Califomian Species of TriUium L. II. The Nature and

Occurrence of Undeveloped Flowers, by Thomas Harper Goodspeed

and Robert Percy Brandt. Pp. 25-38, plates 5-6. October, 1916 ... .15

UNIVERSITY OF CALIFORNIA PUBLICATIONS

IN

BOTANY

Vol. 7, No. 3, pp. 39-68, plates 7-10 December 9, 1916

NOTES ON THE CALIFORNIAN SPECIES OF

TRILLIUM L.

III. SEASONAL CHANGES IN TRILLIUM SPECIES WITH
SPECIAL REFERENCE TO THE REPRODUCTIVE

TISSUES

BY

ROBERT PERCY BRANDT

UNIVERSITY OF CALIFORNIA PRESS
BERKELEY

UNIVERSITY OF CALIFORNIA PUBLICATIONS

Kote. — The University of California Publications are ofifered in exchange for the pnhU-
eatlona of leamod societies and institutions, universities and libraries. Complete listi of
all the publications of the University will be sent upon request. For sample copieii, lists of
publications and other Information, address the Manager of the University Press, Berkeley,
Oallfomla, U. B. A. All matter sent in exchange should be addressed to The Exchange
Department, University Library, Berkeley, California, U. 8. A.

OTTO HAREASOWITZ
LEIPZIG
Agent for the series in American Arch-
aeology and Ethnology, Classical Philology,
Education, Modern Philology, Philosophy,
Psychology.

E. FEIEDLANDEE & SOHN
BEELIN
Agent for the series in American Arch-
aeology and Ethnology, Botany, Geography,
Geology, Mathematics, Pathology, Physi-
ology, Zoology, and Memoirs.

BOTAIT?.— W. A. Setchell, Editor. Price per voliune, $3.50. Volnmes I (pp. 418), n (ppi
860), in (pp. 400), rv (pp. S79), completed. Vols. V and VI in progress.

Cited as Univ. Calif. Publ. Bot.

VoL 1. LA Botanical Snrvey of San Jacinto Mountain, by Harvey Monroe

HalL Pp. 1-140; plates 1-14. June, 1902 $1.00

2. Two new Ascoraycetous Fungi Parasitic on Marine Algae, by Minnie

Eoed. Pp. 141-164; plates 15-16. November, 1902 J2S

5. Algae of Northwestern America, by William Albert Setchell and Nl^

thaniel Lyon Gardner. Pp. 165-418; plates 17-27. March, 1903 2J2S

VoL 2. LA Bevlew of Callfomian Polemonlaceae, by Jessie Milliken. Pp. 1-

71; plates 1-11. May, 1904 .76

2. Contributions to Cytologlcal Technique, by W. J. V. Osterhout. Pp.

73-90; 5 text-figures. June, 1904 ' .25

3. Llmu, by WiUiam Albert Setchell. Pp. 91-113. April, 1905 J25

4. Post-Embryonal Stages of the Laminariaceao, by William Albert

SetcheU. Pp. 115-138; plates 13-14. April, 1905 .25

6. Beg'ineratlon among Kelps, by Willi am Albert Setchell. Pp. 139-168;

plates 15-17. July, 1905 „ JO

6. A New Genus of Ascomycetous Fungi, by Nathaniel Lyon Gardner.

Pp. 169-180; plate 18. July, 1905 _. „ -^ .15

7. Teratology In the Flowers of some Callfomian Willows, by William

Warner Mott. Pp. 181-226; plates 16-20. December, 1905 .60

8. 9, 10, 11. (In one cover.) The Eeslstance of Certain Marine Algae to

Changes in Osmotic Pressure and Temperatura. The E61o of Oo-
motlc Pressure in Marine Plants. On the Importance of Physiolog-
ically Balanced Solutions for Plants. The Antitoxic Action of
Potassium on Magnesium. By W. J. V. Osterhout. Pp. 227-238.
Idarch, 1906 _ .28

12. Cytologlcal Studies in Cyanopbyceae, by Nathaniel Lyon Gardner.

Pp. 237-296: plates 21-26. November, 1906 _ LOO

18. On a Small Collection of Mosses from Alaska, by J. Cardot and T.

Th6riot. Pp. 297-308; plates 27-^. December, 1906 „ JO

14. Some Unreported Alaskan Sphagna, together with a Summary of the
Cryptogamlc Work of the University of California Botanical Ex-
pedition to Alaska In 1899, by William Albert SetchelL Pp. 309-
315. September, 1907 „ .06

16. On Nutrient and Balanced Solutions, by W. J. V. Osterhout. Pp. S17-

318. October, 1907 . M

16. A Synopsis of the North American Godetias, by Willis T.inp Jepson.

Pp. 319-354; plate 29. December, 1907 M

Index, pp. 355-360.

Vol t. 1907-1909.

L Compogltae of Southern California, by Harvey Monroe H^^? Pp. 1-

302; plates 1-3, with a map. December, 1907 S.00

t. The Origin, Structure, and Function of the Polar Caps in SmUacina

amplexicuulis Nutt^ by H. D. Densmore. Pp. 303-330; plates 4-8.

December, 1908 _ _ .88

I. 4. (In one cover.) The Value of Sodium to Plants by Season of Ito

Protective Action. On the Effects of Certain Poisonous Gases on

Plants. By W. J. V. Osterhout. Pp. 331-340. June, 1908 — M

6. Contributions to the Knowledge of the California Species of Crusta-

ceons Corallines. I. by Maurice Barstow Nichols. Pp. 841-348;

plate 9. December, 1908 JO

UNTTESSITT OF OALIFOSNIA FUBLIOATIONS— (Continued)

6. Contributions to the Knowledge of the Callfomla Species of Crusta-

ceous Corallines, n. by Maurice Barstow Nichols. Pp. 849-870;
plates 10-13, April, 1909 ..- JL6

7. New Ohlorophyceae from California, by Nathaniel Lyon Gardner. Pp.

371-375; plate 14. April, 1909 M

8. Plantae Mexicanae Purpusianae, I, by T. S. Brandegee. Pp. 377-396.

May, 1909 _ „ .15

Index, pp. 397-400.
▼oL 4. 1010-1912.

1. Studies in Ornamental Trees and Shrubs, by Harrey Monroe HalL Pp.

1-74; plates 1-11; 15 text-figures. March, 1910 .76

2. Gracllarlophila, a New Parasite on Gractlaria confervoidet, by Harriet

L. Wilson. Pp. 75-84; plates 12-13. May, 1910 ^ .10

8. Plantae Mexicanae Purpusianae, U, by T. S. Brandegee. Pp. 85-95.

May, 1910 .10

4. Leuvenia, a New Genus of Flagellates, by N. L. Gardner. Pp. 97-106;

plate 14. May, 1910 „ ..„ J.0

5. The Genus Sphaerosoma, by William Albert Setchell. Pp. 107-120;

plate 15. May, 1910 — . .15

6. Variations in Nucloar Extrusion Among the Fucaceae, by Nathaniel

Lyon Gardner. Pp. 121-136; plates 16-17. August, 1910 J.6

7. The Nature of the Carpostomes in the Cystocarp of Ahnfeldtia gigarti-

noides, by Ada Sara McFadden. Pp. 137-142; plate 18. February,
1911 ~ JOS

8. On a Colacodasya from Southern California, by Mabel Effie McFadden.

Pp. 143-150; plate 19. February, 1911 .05

9. Fructification of Macrocystls, by Edna Juanita Hoffman. Pp. 151-158;

plate ^0. February, 1911 _„ _ .05

10. Erythrophyllum delesserioides J. Ag., by Wilfred Charles Twiss. Pp.

159-176; plates 21-24. March, 1911 ,.._ .15

11. Plantae Mexicanae Purpusianae, m, by T. S. Brandegee. Pp. 177-194.

July, 1911 _ .„ „. -..-...._ _ — J,6

12. New and Noteworthy Calif omian Plants, I, by Harvey Monroe HalL

Pp. 195-208, March, 1912 „ _ „ .15

13. Die Hydrophyllaceen der Sierra Nevada, by August Brand. Pp. 209-

227. March, 1912 „ - .20

14. Algae Novae et Minus Cognltae, L by WiHiam Albert SetchelL Pp.

229-268; plates 25-31. May, 1912 - - M

15. Plantae Mexicanae Purpusianae, IV, by Townshend Stith Brandegee.

Pp. 269-281. June, 1912 - _ ~ IB

16. Comparative Development of the Cystocarps of Antithamnion and

Frionitis, by Lyman Luther Daines. Pp. 283-302; plates 32-34.
March, 1913 „ ..™ — - ^

17. Fungus Galls on Cystoseira and Ealidrys. by Lulu May Estee. Pp. 305-

316; plate 35. March, 1913 _ 10

18. New Fucaceae, by Nathaniel Lyon Gardner. Pp. 317-374; plates 36-

53. April, 1913 ..„ „....„ - .75

19. Plantae Mexicanae Purpusianae, V, by Townshend Stith Brandegee.

Index, pp. 389-397.
Vol 6. 1912-.

1. Studies in Nicotiana, I, by William Albert Setchell. Pp. 1-86. De-

cember, 1912 - ~ — 1.25

2. Quantitative Studies of Inheritance in Nicotiana Hybrids, I, by Thomas

Harper Goodspeed, Pp. 87-168. December, 1912 — 1.00

8. Quantitative Studies of Inheritance in Nicotiana Hybiids, n, by

Thomas Harper Goodspeed. Pp. 169-188. January, 1913 „ __ .20

4. On the Partial Sterility of Nicotiana Hybrids made with N. Sylvestrit
as a Parent, by Thomas Harper Goodspeed. Pp. 189-198. March,

1913 „ „... ~ .10

6. Notes on the Germination of Tobacco Seed, I, by Thomas Harper Good-
speed. Pp. 199-222. May, 1913 _ _ .26

6. Quantitative Studies of Inheritance in Nicotiana Hybrids, HI, by

Thomas Harper Goodspeed. Pp. 223-231. April, 1915 .10

7. Notes on the Germination of Tobacco Seed, U, by Thomas Harper

Goodspeed. Pp. 233-248. June, 1915 ~ 16

8. Parthenogenesis, Parthenocarpy and Phenospermy in Nicotiana, by

Thomas Harper Goodspeed. Pp. 249-272, plate 35. July, 1915 ~ .25

UNIVEaSITY OF OAUFOENIA PUBLIOATIONS— (Continued)

9. On the Partial Sterility of Nicotiana Hybrids made with N. sylvestris
as a Parent. II, by T. H. Goodspeed and A. H. Ayres. Pp. 273-292,

plate 36. October, 1916 J20

10. On the Partial Sterility of Nicotiana Hybrids made with N. sylvestris
as a Parent. III. An Account of the Mode of Floral Abscission in
the F, Species Hybrids, by T. H. Goodspeed and J. N. KendalL
Pp. 293-299. November, 1916 .06

Vol 6. 1914-

1. Parasitic Florldeae, I, by William Albert Setcbell. Pp. 1-34, plates

2. Phytomorula regularis, a Symmetrical Protophyte Belated to Coehu-

trum, by Charles Atwood Kofoid. Pp. 36-40, plate 7. April, 1914, .06
8. Variation in Oenothera ovata, by Eatherlne Layne Brandegee. Pp. 41-

50, plates 8-9. June, 1914 „ _ J.0

4. Plantae Mexicanae Piirpusianae, VI, by Townshend Stith Brandegee.

Pp. 51-77. July, 1914 ~ SB

6. The Scinaia Assemblage, by William Albert Setchell. Pp. 79-152,

plates 10-16. October, 1914 .76

6. Notes on Pacific Coast Algae. I. Pylaiella Postelsiae, n. sp., a New

Type in the Genus Pylaiella, by Carl Skottsberg. Pp. 153-164, plates
17-19. May, 1915 „ „ _ — .15

7. New and Noteworthy Calif omian Plants, n, by Harvey Monroe HalL

Pp. 165-176, plate 20. October, 1915 - .16

8. Plantae Mexicanae Purpusianae VII, by Townshend Stith Brandegee.

Pp. 177-197. October, 1915 _ - .25

9. Floral Relations Among the Galapagos Islands, by A. L. Kroeber.

Pp. 199-220. March, 1916 .20

10. The Comparative Histology of Certain Califomian Boletaceae, by

Harry S. Yates. Pp. 221-274, plates 21-25. February, 1916 — .50

11. A Revision of the TuberaJes of California, by Helen Margaret Gilkey.

Pp. 275-356, plates 26-30. March, 1916 _ .80

12. Species Novae vel Minus Cognitae, uy T. S. Brandegee. Pp. 357-361.

May, 1916 05

Vol. 7. 1916-

1. Notes on the Califomian Species of Trillium L. I. A Report of the

General Results of Field aud Garden Studies, 1911-1916, by Thomas
Harper Goodspeed and Robert Percy Brandt. Pp. 1-24, plates 1-4.
October, 1916 25

2. Notes on the Califomian Species of Trillium L. IL The Nature and

Occurrence of Undeveloped Flowers, by Thomas Harper Goodspeed

and Robert Percy Brandt. Pp. 25-38, plates 5-6. October, 1916 ... .15

3. Notes on the Califomian Species of Trillium L. III. Seasonal Changes

in Trillium Species with Special Reference to the Reproductive Tis-
sues, by Robert Percy Brandt. Pp. 39-68, plates 7-10. December,
1916 .- 30

UNIVERSITY OF CALIFORNIA PUBLICATIONS

tN

BOTANY

Vol. 7, No. 1, pp. 1-24, plates 1-4 October 3, 1916

NOTES ON THE CALIFORNIAN SPECIES OF

TRILLIUM L.

I. A REPORT OF THE GENERAL RESULTS OF FIELD
AND GARDEN STUDIES, 1911-1916

BY

THOMAS HARPER GOODSPEED and ROBERT PERCY BRANDT

UNIVERSITY OF CALIFORNIA PRESS
BERKELEY

UNIVERSITY OF CALIFORNIA PUBLICATIONS

Note. — The TTnlverslty of California Publications are offered in exchange for the pabll-
eatlona of learned societies and institutions, universities and libraries. Complete lists of
all the publications of the University will be sent upon request. For sample copies, lists of
publications and other information, address the Manager of the University Press, Berkeley,
O&llfomla, U. S. A. All matter sent in exchange should be addressed to The Bxchango
Department, University Library, Berkeley, California, U. S. A.

OTTO HAERASOWITZ B. FRIEDLANDEE & SOHN
LEIPZIG BERLIN
Agent for the series in American Arch- Agent for the series in American Arch-
aeology and Ethnology, Classical Philology, aeology and Ethnology, Botany, Geography,
Education, Modern Philology, Philosophy, Geology, Mathematics, Pathology, Physi-
Psychology. ology. Zoology, and Memoirs.

BOTAKT.-— W. A. Setchell, Editor. Price per volume, $3.60. Volmnes I (pp. 418), II (ppi
360), III (pp. 400), IV (pp. 379), completed. Vols. V and VI In progress.

Cited as Univ. Calif. Pnbl. Bot.

Vol. 1. 1. A Eotanical Enrvey of San Jacinto Mountain, by Harvey Monroe

Hall. Pp. 1-140; plates 1-14. June, 1902 ILOO

5. Two new Ascomycetous Fungi Parasitic on Marine Algae, by Mlnde

Eced. Pp. 141-164; plates 15-16. November, 1902 - J25

8. Algae of Northwestern America, by William Albert Setchell and Na-
thaniel Lyon Gardner. Pp. 165-418; plates 17-27. March, 1903 Z2S

Vol 2. 1. A Bevlew of Califomian Polemonlaceae, by Jessie MlUiken. Pp. 1-

71; plates 1-11. May, 1904 .75

2. Contributions to Cytological Technique, by W. J. V. Osterhout. Pp.

73-90; 5 text-figurea. June, 1904 J25

3. LImu, by William Albert SetcheU. Pp. 91-113. April, 1905 .26

4. Post-Embryonal Stages of the Laminariaceas, by WtUiam Albert

Setchell. Pp. 115-138; plates 13-14. April, 1905 26

6. Begeneration among Kelps, by William Albert Setchell. Pp. 139-168;

plates 15-17. July, 1905 » .30

6. A Now Genus of Aacomycetous Fungi, by Nathaniel Lyon Gardner.

Pp. 169-180; plate 18. July, 1905 „ „.. .15

7. Teratology in the Flowers of some Califomian Willows, by William

Warner Mott. Pp. 181-226; plates 16-20. December, 1905 „ 50

8. 9, 10, 11. (In one cover.) The Resistance of Certain Marine Algae to

Changes in Osmotic Pressure and Temperaturs. The ESle of Os-
motic Pressure in Marine Plants. On the Importance of Physiolog-
ically Balanced Solutions for Plants. The Antitoxic Action of
Potassium on Magnesium. By W. J. V. Osterhout. Pp. 227-238.

March, 1906 „ _ J25

12. Cytological Studies in Cyanophyceae, by Nathaniel Lyoa Gardner.

Pp. 237-296; plates 21-26. November, 1906 1.00

15. On a Small Collection of Mosses from Alaska, by J. Cardot and T.

Th6riot. Pp. 297-308; plates 27-28. December, 1906 J.0

14. Some Unreported Alaskan Sphagna, together with a Siuuai&ry of tho
CryT)togamic Work of the University of California Botanical Ex-
pedition to Alaska In 1899, by WlUiam Albert SetchelL Pp. 309-

315. September, 1907 .08

IB. On Nutrient and Balanced Solutions, by W. J. V. Osterhout. Pp. 317-

318. October, 1907 _ .06

16. A Synopsis of the North American Godetlas, by Wllllfl Linn Jepson.

Pp. 319-354; plate 29. December, 1907 .40

Index, pp. 355-360.

VOL 8. 1907-1009.

1. Composltae of Southern California, by Harvey Monroe Hall. Pp. 1-

302; plates 1-3, with a map. December, 1907 1.00

2. The Origin, Structure, and Function of the Polar Caps In SmQacina

amplexxcaulig Nutt., by H. D. Denamore. Pp. 303-330; plates 4-8.

December, 1908 35

8. 4. (In one cover.) The Value of Sodlnm to Plants by Eeaaon of Its

Protective Action. On the Effects of Certain Poisonons Gases on

Plants. By W. J. V. Osterhout. Pp. 331-340. June, 1908 JO

6. Contributions to the Knowledge of the California Species of Orusta-

ceouB Corallines. I. by Maurice Barstow Nichols. Pp. S41-348;

plate 8. December, 1908 .10

UNIVUaSITY OP OAUFOENIA PUBLICATIONS— (Oontinued)

6. Contributions to the Knowledge of the California Species of Crusta-

ceous Coraliine». n. by Maurice Barstow Nichols. Pp. 349-370;
plates 10-18. April, 1909 .„ „ „ .16

7. New Chlorophyceae from California, by Nathaniel Lyon Gardner. Pp.

371-375; plate 14. April, 1909 ,._ „..;.... .10

8. Plantae Mexicanae Purpusianae, I, by T. S. Brandegee. Pp. 377-396.

May, 1909 -....,- .::. :...... 15

Index, pp. 397-400.
Vol 4. 1910-1912,

1. Studies In Ornamental Trees and Shrubs, by Harvey Monroe H&IL Pp.

1-74; plates 1-11; 15 text-figures. March, 1910 .75

2. Qracilariophila, a New Parasite on Gracilaria confervoides, by Harriet

L. Wilson. Pp. 75-84; plates 12-13. May, 1910 „ .10

3. Plantae Mexicanae Purpusianae, II, by T. S. Brandegee. Pp. 85-05.

May, 1910 ..„.. -... .,._ _..... _ _ .lo

4. Leuvenia, a New Genus of Flagellates, by N. L. Gardner. Pp. 97-106;

plate 14. May, 1910 _.. .10

6. The Genus Sphaetosoma, by WUUam Albert Setchell. Pp. 107-120;

plate 15. May, 1910 15

6. Variations in Nuclear Extrusion Among tbe Fncaceae, by Nathaniel

Lyon Gardner. Pp. 121-136; plates 16-17. August, 1910 __ .18

7. The Nature of the Carpostomes in the Cystocarp of Ahnfeldtia gigarti-

noides, by Ada Sara McPadden. Pp. 137-142; plate 18. February,
1911 _._ .05

8. On a Colacodasya from Southern California, by Mabel Eflie McFadden.

Pp. 143-150; plate 19. February, 1911 _ _ .OS

9. Fructification of Macrocystis, by Edna Juanita Hoffman. Pp. 151-158;

plate 20. February, 1911 „ .05

10. ErythropTiyllum delesserioides J. Ag., by Wilfred Charles Twiss. Pp.

159-176; plates 21-24. March, 1911 S5

11. Plantae Mexicanae Purpusianae, III, by T. S. Brandegee. Pp. 177-194.

July, 1911 _ .15

12. New and Noteworthy Calif omian Plants, I, by Harvey Monroe HalL

Pp. 195-208. March, 1912 _........_ „ — J.5

IS. Die Hydrophyllaceen der Sierra Nevada, by August Brand. Pp. 209-

227. March, 1912 ,.......- ...„ 20

14. Algae Novae et Minus Cognitae, I, by William Albert Setchell. Pp.

229-268; plates 25-31. May, 1912 ™ .40

15. Plantae Mexicanae Purpusianae, IV, by Townshend Stith Brandegee.

Pp. 269-281. June, 1912 _ 16

16. Comparative Development of the Cystocarps of Antithamnion and

Prionitis, by Lyman Luther Daines. Pp. 283-302; plates 32-34.
March, 1913 .20

17. Fungus Galls on Cystoseira and Halidrys, by Lulu May Estee. Pp. 305-

316; plate 35. March, 1913 ..„ 10

18. New Fucaceae, by Nathaniel Lyon Gardner. Pp. 317-374; plates 36-

53. April, 1913 - .75

19. Plantae Mexicanae Purpusianae, V, by Townshend Stith Brandegee.

Pp. 375-388. June, 1913 „ 16

Inde?, pp. 389-397.
Vol. B. 1912-.

1. Studies in Nicotiana, I, by William Albert Setchell, Pp. 1-86. De-

cember, 1912 _ 1.28

2. Quantitative Studies of Inheritance in Nicotiana Hybrids, I, by Thomas

Harper Goodspeed. Pp. 87-168. December, 1912 1.00

8. Quantitative Studies of Inheritance in Nicotiana Hybrids, n, by

Thomas Harper Goodspeed. Pp. 169-188. January, 1913 _. .20

4. On the Partial Sterility of Nicotiana Hybrids made with N. Sylvestri^

as a Parent, by Thomas Harper Goodspeed. Pp. 189-198. March,

1913 _ .10

5. Notes on the Germination of Tobacco Seed, I, by Thomas Harper Good-

speed. Pp, 199-222. May, 1913 25

6. Quantitative Studies of Inheritance in Nicotiana Hybrids, III, by

Thomas Harper Goodspeed. Pp. 223-231, April, 1915 10

7. Notes on the Germination of Tobacco Seed, II, by Thoraaa Harper

Goodspeed. Pp, 233-248. June, 1915 15

8. Parthenogenesis, Parthenocarpy and Phenospermy in Nicotiana, by

Thomas Harper Goodspeed. Pp. 249-272, plate 35. July, 1915.„ 25

9. On the Partial Sterility of Nicotiana Hybrids made with N. siflvestris

as a Parent. II, by T. H. Goodspeed and A, H, Ayres, Pp, 273-292,
plate 36, October, 1916 20

UNTVERSITY OF OALIFOaNIA PUBLICATIONS— (Oontinned)

Vol. 6. 1914-

1. Parasitic Florideae, I, by V/illiara Albert Setchell. Pp. l-3i, plates

1 C. April, 1914 ™ _„ ^

2. Fhytomorula regularis, a Symmetrical Protophyte Eelated to Codas-

trum, by Charles Atwood Kofoid. Pp. 35-40, plate 7. April, 1914. .06

3. Variation in Oenothera ovata, by Katherine Layne Brandegee. Pp. 41-

50, plates 8-9. June, 1914 _ __ .10

4. Plantae Mexicanae Purpusianae, VI, by Towushend Stith Brandegee.

Pp. 51-77. July, 1914 .26

5. The Scinaia Assemblage, by WiUiam Albert Setchell. Pp. 79-152,

plates 10-16. October, 1914 - _ .75

6. Notes on Pacific Coast Algae. I. Pylaiella Postelsiae, n. sp., a New

Type In the Genus Pylaiella, by Carl Skottsberg. Pp. 153-164, platea
17-19. May, 1915 — _ .15

7. New and Noteworthy Califomian Plants, U, by Harvey Monroe Hall.

Pp. 165-176, plate 20. October, 1915 ._,............i.*...^^^.. .16

8. Plantae Mexicanae Purpusianae VII, by Townshend Stith Brandegee.

Pp. 177-197. October, 1915 25

9. Floral Eelations Among the Galapagos Islands, by A. L. Kroeber.

Pp. 199-220. March, 1916 _ 20

10. The Comparative Histology of Certain Califomian Boletaceae, by

Harry S. Yates. Pp. 221-274, plates 21-25. February, 1916 „ 50

11. A Revision of the Tuberales of California, by Helen Margaret Gilkey.

Pp. 275-356, plates 26-30. March, 1916 80

12. Species Novae vel Minus Cognitae, by T. S. Brandegee. Pp. 357-361.

May, 1916 05

Vol. 7. 1916-

1. Notes on the Califomian Species of Trillium L. I. A Report of the

General Results of Field and Garden Studies, 1911-1916, by Thomas
Harper Goodspeed and Robert Percy Brandt. Pp. 1-24, plates 1-4.
October, 1916 , 25

2. Notes on the Califomian Species of -Trillium L. II. The Nature and

Occurrence of Undeveloped Flowers, by Thomas Harper Goodspeed

and Robert Percy Brandt. Pp. 25-38, plates 5-6. October, 1916 ... .15

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