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http://www.archive.org/details/sSbotproceedingsO2call

PROCEEDINGS

CALIFORNIA ACADEMY OF SCIENCES

THIRD SERIES

Se Ge AN ay
Vor, Lt

1900-1904

SAN FRANCISCO
PUBLISHED BY THE ACADEMY

1904

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ZePe-
kee
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672950
Eee

CONTENTS, OF VOLUME: Ii:
PLATES I-X XXII.

PAGE
SUR fOr re otc ieee opnsineta cl te staan ai Giste eicypiars oats dipisis vicum'e ake skis! isyae,e Slaten i
(CLOY THEY oI EC Re Wire ans nt Ae ear re Ag DR ie at EO ad oN SEA ili

No. 1. Nitophylla of California. oe and distribution. ma Chas.
Palmer Nott. (Plates I-IX) . Hees je anaes I
(Published August 3, 1900.)

No. 2. The Development of the Karyokinetic Spindle in the Pollen-
Mother - Cells of Lavatera. By Edith Sumner Byxbee.
(Plates X=—XAIID))s. o.25 25.0 ck ieee ae GLA ee 6 63
(Published October 31, 1900.)

No. 3. Studies on the Coast Redwood, Sequoia sempervirens Endl.
By George James) Peirce., ((Plate XIV 5.5.) eae. ols sierde- 83
(Published April 4, 1901.)

No. 4. A Revision of the Genus Calochortus. By Carl Purdy. (Plates
DENVED-4 DO ao SEE REIS Ae eters + 2 SU Ada ae an 107
(Published December 14, 1901.)

No. 5. A Group of Western American Solanums. By S. B. Parish... 159
(Published October 23, 1901.)

No. 6. An Account of the Species of Porphyra Found on the Pacific
Coast of North America. By Henri T.-A. Hus. (Plates
DBS, |) WO AER OLA mn ne Si OTE Uk AG A 173
(Published January 4, 1902.) ‘

No. 7. Some new Species of Pacific Coast Ribes. By Alice Eastwood.
(Plates XEXTTISX XIV) vais ose e cae 2 ec ce cares ene eee ie 241
(Published April 14, 1902.)
No. 8. Cell Studies. I. Spindle Formation in Agave. By W. J. V.
@sterhortaye (blates xOGV— NeONOVAMI ie crates elisiayelsteier ciereisieeh sie 255
(Published May 5, 1902.)

No. 9. New Species from the Sierra Nevada Mountains of California.
Biymaice MaStwOOG) acon: sects sha titae aa cedave sl overefots a mleiet bios 285
(Published June 3, 1902.)

~No. 10. The Root-tubercles of Bur Clover (Medicago denticulata
Willd.) and of some other Leguminous Plants. By George
amestheince:) (Plate, Rex Le) Mier csp aleycislaier vets neice © « te 295
(Published June 21, 1902.)
No. 11. Spindle Formation in the Pollen-Mother-Cells of Cassia to-
mentosa L. By Henri T. A. Hus. (Plates XXX-XXXII) 329
(Published April 30, 1904.)

April 30, 1904.

; ie Derr,

NITOPHYLLA OF CALIFORNIA. DESCRIPTION

AND DISTRIBUTION :!

BY CHARLES PALMER NOTT.

CONTENTS.
Piates I-IX.

PAGE
TARAUNT CRO DUCTION Het 35 oas oe acetate ei tee en ciitim erin stesiaaone atte g 2
SR EEAISROR WH cesta d ote his ees eelee Geet ees Ns aerate orsie au e vate be 2
MING HMERAT IOESCRIPEION ©: canis cede aa ee oso 6 cisco um cess 4, nice % 5
SyvNOPSIS OF GENERIC (CHARACTERS) » 6.-[5.0)25 101452) sles <0: cei s 5

REMARKS ON SOME GENERIC CHARACTERS AS ILLUSTRATED
BY, CAMIFORNIANS SPECIES Sache pst istiae oo = sicle<ieic'a\'s 7
LRE PROSEGALE ET OME: Fen ele terckeri to siete eed = 5) isi wena a's 7
DR LUCHICE DF} NIOUSETUEUI Ds Diets Ge taint fale e's) ticie ep) sis sale 2 2 7
ONSITE Dede OES re terse, cheb hayes akg sro rs a ele ate 8
(DF SROOLS) Cte AGMODOMOMS sae velapis aicie\aieie m1 2's) 2 Sia vie cla 22 9
IMM By AAG d LAG TOR yaa OOOO OCD HEC TOUIAIG ae OOO E SC 9
SETS LIE oie 5 SEMEL a: 2a MRS WO Gok es Oe i RMA eR Ue 9
SDE LID DEPP PIC HIE A Hee oar ik la che) ale 5:20) Ber orud ayia cieioleteini oigts 9
Form and Branching as Specific Characters........... IO
IS ARE A OO Dee ne ROC oa aoe noice Sch a neti sf)
STO KATO e A ed WEES EID COOLS CIE RRO Ue iGn clone DOES 10
INOUE SAPD ars Poa et etote eV Re Le eal ete See state oR Nate Se teks Tat
VOUS let Naat ont ee lae elatelal alae a ene lara A neleve s eieiae ey eats II
Variable and Fixed. CROCCO S. 5.2 ids Say LAR oe II
PA DETOU Ysera hou tdel takeoigra eas assis epee tae «Mele BLE 12
SORT, du he etre sale NaS ieieh nae Taceee ines, Su et eens aie eel aes 12
PR BLOC IOs ee iaicie) os niet aes ores Ste Sea ee ee 13
MAIMEKIAIa ANd» CYSIOCAT PS tn ceijomn cide sees ise 14
U PAT ISAAC A OCTETS ee CTA ROP PERE OOPS RAAT OPIN eae Se 14
Geographical Distribution of the Genus .......... 2.4. 14
LSAT ET ME BY Sot SE AN PD ey ASS ea, VELL ode OS 15
IV. DESCRIPTION AND DISTRIBUTION OF SPECIES.............0.-206- 15
Kry TO THE CALIFORNIAN SPECIES OF NITOPHYLLUM...... 15
DV URAICUSORTOMUN cierto Sse, ale ssi soa) sila AOS, Des Ae) OF ier 32
ELD BCY OIIGHEN AAA sive, oho 70 Blais snare 6p tin Mis ie ey ate Pood 29
UA IUIEDIOOLEIIE So Se ALI hk Ra Hale Santee ae See ae Baha Zi}
Dy EOURLOUNES «Cried cha Gala loln ele afs arakw. 4 3 16he be iafa sheen othe old 39
GTEC ENOLLAIULERTE soca aoe rhein 8 sarniahe, A eING Se Ponte eRe SA 34
UNE POETS SOMITIEDIE iid, fc) oA oha1 2a RO Wola Me opwaid a Sete skies eo. 16
LT PCO GAL 5 Mie arta soe a Ie SS OR o ede eae ey eR sion 22
I OLA CTO ROO G AAO EOE AIO OAC one 26
ISN ONE LEE OSE AD IA SECT COO Ten OU oe ae 24
VEE SPCED LOUIE he to, rae noe Rhys. ota a erase cata a. ys eoban sian e 21
DAREH OF MISTRIBUTION, OF) SPHOIES 5 5.-25)5 552 catenins oye 4 eis 43
LITERATURE AND EXSICCAT# CITED IN THIS PAPER............ 0.0005 45
EXPEANATION OR MPEATES./0. Heo 52 {cissiersS yey veined osledeidais Hoktoe bee wees 46

1 Contributions from the Botanical Laboratories of the University of California, No. ro.
{r] July 24, 1900,

2 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

I. INTRODUCTION.

Our knowledge of the genus /Vtophyllum Grev. as rep-
resented upon the coast of California and adjacent shores
has been hitherto of a fragmentary and limited nature.
The number of species found in the region named, the
points of their occurrence, and their identity with the forms
already known, or their title to recognition as distinct spe-
cies, have been for some time largely matters of conjecture.
With a view of providing a more connected account of
these species, so far as data could be obtained, the follow-
ing descriptions and accompanying notes upon distribution
are presented.

II. HustTory.

The first notice of Californian /Vetophylla, so far as the
writer is aware, was made by W. H. Harvey (1858, Pt. II,
p- 104, Supp., p. 128), who mentions the species of /Vito-
phyllum then known upon the western coast of North
America. Two forms, JV. fryeanum and JV. laceratum (LV.
violaceum )', were mentioned as occurring at Golden Gate,
San Francisco Bay, California. Still later, in a ‘‘ Notice
of a Collection of Algz made on the Northwest Coast of
North America, chiefly at Vancouver’s Island, by David
Lyall, in the years 1859-61.’ Harvey (1862, p. 170)
gives the original description of Hymenena latissima (LV.
latissimum ), specimens of which were dredged, or found
adrift in Esquimault Harbor, Vancouver’s Island, B. C.

The next mention of Californian /V:tophyl/a is found in
the ‘‘Bidrag till Florideernes Systematik” of J. G. Agardh
(1871, p. 49), where reference is made to Harvey’s species
Hymenena latissima Harv. (LV. latissimum ).

A noteworthy addition to the number of North American
species was made when Professor W. G. Farlow (1875,
p- 365) published a ‘‘List of the Marine Alge of the United
States, with Notes of New and Imperfectly Known Species.’’

1 The names in parentheses are those applied to the species as recognized in this
paper, where the quoted name differs.

Bot.—Vot. II.] MOZT7—CALIFORNIAN NITOPHYLLA. 3

In this paper were enumerated the following six species:
N. (Neuroglossum) andersonit J. Ag. (lV. andersonta-
num); lV. fissum J. Ag. (iV. ruprechtianum); NV. fryeanum
Harv.; JV. laceratum Grev. (lV. violaceum); LV. latissimum
J. Ag.; V. ruprechtianum J. Ag.

In the following year Farlow (1876, p. 695) published a
second list, which included all the above mentioned species
with the addition of /V. areolatum D. C. Eaton (LV. latis-
stmum) and JV. spectabile D. C. Eaton.

The most comprehensive account of /Vitophyllum yet pre-
sented was that contained in the Epicrisis Floridearum of
J. G. Agardh (1876, pp. 446-472, 698-701). This account
of the genus included all of the species then known to
occur on the west coast of North America, with the excep-
tion of JV. spectabéle D. C. Eaton. To the forms men-
tioned by Harvey and Farlow were now added JV. multz-
lobum J. Ag., WV. violaceum J. Ag., and VV. flabelligerum
J. Ag. (LV. ruprechtianum).

Farlow (1877, pp. 238, 245), in a paper discussing some
algz new to the United States, comments on some of the
species mentioned by him in the two papers of 1875-76,
and also gives the original description of JV. spectabile
D. C. Eaton.

Between 1877 and 1898, the literature pertaining to Cali-
fornian /Vtophyl/a consists of lists of the forms occurring
on the coast, with the exception of Hervey’s ‘‘ Sea Mosses”’
(1881), a popular work, in which some of the character-
istics of color, size and venation of the JVztophylla are
described. Several other writers should here be noticed.
Moye. Anderson (1897, p. 224), MA. Howe (1893,
p. 67), Daniel Cleveland, and A. J. McClatchie (1897, p.
358) have published lists of coast forms. No additions
were made to the number of west coast forms by any of
these writers except the last named, whose paper contains
the first mention of the occurrence of JV. uncznatum J. Ag.
upon the Californian coast.

The latest and most complete statement in regard to west
coast /Vztophylla is found in the volume published by J. G.
Agardh (1898).

4 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

According to Agardh twelve species of JVtophyllum and
two species of /Vewrog/ossum are assigned to the west coast
of North America. The complete list is as follows, viz.:
NV. farlowianum J. Ag. (lV. ruprechtianum); N. flabell-
gerum J. Ag. (NV. ruprechtianum); N. fryeanum Harv. (LV.
fryeanum); LV. latissimum J. Ag.; V. macroglossum J. Ag.
(LV. latissimum); WN. multilobum J. Ag.; WV. marginatum
J. Ag. (LV. ruprechtianum); N. ruprechtianum J. Ag.; NV.
spectabile Eaton.; JV. stenoglossum J. Ag. (lV. violaceum);
LV. uncinatum J. Ag.; LV. violaceum J. Ag.; Neuroglossum
andersonianum J. Ag. (Vit. andersonianum); Neuroglos-
sum lobuliferum J. Ag. (Vit. violaceum ?).

Agardh, basing his distinction between species upon dif-
ferences in color, texture, form of frond, and position of
cystocarps, as well as upon the more reliable characters,
such as shape and position of sori, and venation of frond,
regards all these forms established by him as valid species.

It should be observed, however, in regard to these forms,
that the descriptions are in many cases, as admitted by the
author himself, drawn up from fragmentary or imperfect
specimens. Further, the characteristics which are em-
ployed to a considerable extent by Agardh as distinctions
between species, and even between subsections, are vari-
able to a marked degree. Observation of a considerable
range of forms by the writer has led to the conclusion that
sufficient allowance has not been made for the variations
which require that great freedom should be used in defining
the boundaries of species occurring on the Californian coast.
A distinction should be made between the more variable
characters such as color, texture, form of frond, and posi-
tion of the cystocarps, and the less variable characters, such
as shape and position of sori, and venation of frond.

After a careful examination of the descriptions of the
species as formulated by Agardh, and further study and
comparison of the variations of the plants themselves, the
writer is obliged to conclude that the degree of elasticity
which seems desirable in considering Californian forms has
not been permitted in regulating the limits of a species, or

Bot.—Vot. II.] MOT7—CALIFORNIAN NITOPHYLLA. 5

else access has been had to material which has not come
under the writer’s observation. A further consideration of
these forms will be found in the remarks upon species in a
later portion of this paper.

The writings of the three authors mentioned above, viz.,
Harvey, Agardh, and Farlow, constitute the important liter-
ature upon Californian /Vztophylla, and their work alone
will be considered in the further discussion of the species.

III. GENERAL DESCRIPTION.

The fuller discussion of the species of /Vetophyllum of the
Californian coast will be advanced by some special treat-
ment of the prominent morphological characters and geo-
graphical distribution of the genus itself, for the sake of the
increased light thrown by such treatment upon like points
in connection with the forms to be hereafter discussed.

The following synopsis exhibits the principal characters
of this genus and will be followed by a discussion of some
special features of morphology and distribution, as illus-
trated by Californian species.

SYNOPSIS OF GENERIC CHARACTERS.

Frond either erect or exhibiting a prostrate and an erect
portion.

Prostrate frond creeping, linear, or irregularly expanded
or membranous, occasionally with midrib, nerves, or veins;
with or without rhizoids; variously lobed, divided or
branched, occasionally proliferating; branches rising at
intervals into erect fronds; margin entire, serrate, dentate,
crenate, undulate, or lobed; sometimes forming offshoots
and innovations.

Erect fronds rising from holdfast or prostrate frond,
singly or several together; sessile, subsessile, or stalked;
flat and linear, or membranous, variously lobed, divided,
forked, segmented, and branched; frequently proliferating ;

6 CALIFORNIA ACADEMY OF SCIENCES. [PRoC. 3D SER.

with or without midrib, nerves or veins; with margin
entire, serrate, dentate, crenate, undulate, orlobed. Branch-
ing subdichotomous, subpinnate, subpalmate, or palmate,
with branches or segments entire, linear or expanded,
sometimes much prolonged. Stalk linear, flat, with or
without distinct midrib; frequently becoming thickened and
cylindrical through wearing away of margin of frond and
renewed growth of remaining portion; frequently twisted
by wave action; often persistent and freely proliferating.

Midrib, when present, usually conspicuous, narrow or
wide, simple below, sometames branched above, sometimes
evanescent or dividing into flabellate or anastomosing
nerves, frequently becoming thickened, stout and persist-
ent, freely proliferating. Nerves usually conspicuous, occu-
pying body of frond, margin, or apices, usually branching
‘freely, flabellate, free, or anastomosing, sometimes dividing
into minute and inconspicuous veins. Veins inconspicuous
or microscopic, occupying body of frond, margin or apices,
simple or branching, flabellate, free, or anastomosing, com-
monly evanescent and indistinguishable from ordinary tis-
sue of frond.

Sporangia found on both surfaces of the frond, usually
in locally thickened portions, in sori of varied shape, con-
taining tripartite tetraspores. Sori minute and scattered
over the entire surface of the thallus, or large, forming
orbicular patches disposed irregularly over the surface; or
linear marginal patches; or lines arranged radially along
the margin; or borne on marginal or surface proliferations
of varying size.

Antheridia developed from the superficial cells of the
thallus, forming whitish patches scattered over the surface
of the frond, the latter frequently becoming rugose.

Cystocarps scattered over both surfaces of the frond, or
arranged along the margin, or borne on marginal prolifera-
tions, usually large, projecting beyond the surface of the
thallus, opening by a carpostome.

Bot.—Vou. Il.] MWOTT—CALIFORNIAN NITOPHYLLA. 7

REMARKS ON SOME GENERIC CHARACTERS, AS ILLUS-
TRATED BY CALIFORNIAN SPECIES.

The Prostrate Frond.—The prostrate creeping frond
possessed by many species of Vitophyllum deserves special
consideration. Agardh makes some use of this character
in separating the genus into subsections, but the importance
of this structure to the plant and the extent to which it may
be developed have not, so far as it has been possible to
learn, been very fully demonstrated.

Influence of Substratum.—The character of the sub-
stratum upon which the plant is located and the extent to
which it is exposed to the dashing, drawing, or swirling
force of the waves affect both the amount of growth and
the shape of the prostrate frond.

The various species of /Vzfophyllum occur in a variety of
situations, from about mid-tide mark outward and down-
ward to deep water. They grow in some cases upon the
piles of wharves, where the prostrate frond must take advan-
tage of cracks in the wood or roughenings of the surface to
secure a foothold. Other species are found upon bare rock-
surfaces, exposed to the dash of breakers. The most com-
mon situation is that of those species which inhabit sheltered
rock crevices or pools surrounded by rocks which protect
them from the force of the waves. In such spots there will
usually be found upon the rocks a rich growth of Bryozoa
and Porifera, whose sponge-like substance affords an excel-
lent foothold for the plants and is conducive to a free devel-
opment of the prostrate frond. Other alge, notably the
Corallines, afford by reason of their jointed structure and
rough surface, excellent habitats. In general, the charac-
ters of the prostrate frond correspond to those of the erect
portion, but there are marked exceptions. JV. datissimum,
for instance, possesses a membranous, broadly divided,
erect frond, while that of /V. andersontanum is very much
branched. Two forms in the same genus could hardly
seem more widely different, yet the prostrate frond of the
two species is very much alike where growing under

8 CALIFORNIA ACADEMY OF SCIENCES. [PRroc. 3D SER.

approximately similar conditions of substratum. and wave
action. Under other conditions the prostrate frond of JV.
Jatissimum may become widely linear and irregularly lobed,
lose the toothed margin which forms one of the points of
resemblance to JV. andersonzanum, and acquire a midrib and
nerves, in this case resembling the prostrate frond of JV.
ruprechtianum. Again, /V. multilobum, which as a rule
selects sharply inclined or vertical rock-surfaces as a habi-
tat, forms by means of its prostrate frond orbicular patches
composed of the closely overlapping, broadly lobed, and
membranous portions of the creeping, prostrate frond. The
room afforded on the bare rock apparently favors the radial
development here exhibited; while the necessity for secur-
ing adequate thickness and firmness to meet the dash of the
waves has led to the close overlapping or dovetailing of the
various divisions of the prostrate frond.

It has already been remarked that the prostrate part of
the plant resembles in many respects the erect portion which
rises from it. This statement usually holds good with
regard to the shape and branching, but does not apply to
the venation. As a rule, a midrib or nerves are lacking
in the prostrate frond of those species whose erect fronds
are provided with such structures.

Formation of Rhizoids.—A number of species also exhibit
a response apparently to the stimulus of contact, by send-
ing short processes or rhizoids from the surface of the
frond to the substratum to which they adhere. These pro-
cesses have been observed on the under surface of the
prostrate frond of JV. ruprechtianum, NV. violaceum, LV.
multilobum, IV. harveyanum and JV. corallinarum, and are
recorded for several other species. Still more remarkable
is the instance observed of the formation of these rhizoids
in the case of a plant of JV. vzolaccum. This specimen had
wrapped itself around portions of the thick frond of Przo-
nites lanceolata, to whose surface numerous processes sent
forth from the surface of the Vztophyl/um in contact with
the Przonztzs had attached themselves.

Bot.—Vot. II.] MOTT—CALIFORNIAN NITOPHYLLA. 9

Offshoots and Innovations.—A point of further interest in
connection with the development of the prostrate frond is
found in the formation of offshoots and innovations. In the
first case, slender branches may arise from the margin of
the older portion of the frond. These grow, secure attach-
ment for themselves, and separate from the parent frond,
and later give rise in turn to erect fronds. In the second
case, by the growth and branching of the prostrate frond,
an extended structure is produced, the ramifications of
which become separated from each other by the decay and
disappearance of the older portions, thus forming innova-
tions in a manner similar to the process occurring in the
Bryophytes.

The Erect Frond.—The erect part of the plant commonly
rises singly from the prostrate portion, but occasionally the
fronds are clustered together, as in JV. mz/tilobum, where
they are grouped in the middle of the orbicular patch
formed by the prostrate frond.

Size.—The height and breadth of the erect frond is an
extremely variable character. JV. corallinarum, occurring
upon Corallina chilensts, does not reach a height of 2 cm.;
while JV. spectabile is reported by D. C. Eaton as reaching
50-60 cm.

Shape and Branching.—Great diversity exists in the
shape and branching of the erect frond. Some species are
broadly membranous, and but slightly lobed or divided.
Good examples of this type are JV. spectabile, NV. latissomum
and JV. fryeanum. At the other extreme may be placed
such a finely dissected and abundantly branched species as
LV. andersonianum. Between these two opposing types are
found all gradations of frond division and arrangement of
branches. It is not uncommon to find in one and the same
species forms exhibiting a tendency to become broadly
membranous, or very much divided and branched. For
instance, in lV. ruprechttanum, there seems to be a ten-
dency toward the flabellate or expanded type of frond,
though the typical specimens of the species are character-
ized by division of the frond’ into linear, much prolonged

IO CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

branches. JV. violaceum, on the other hand, varies in the
direction of the finely dissected type of frond. JV. fryeanum
is a membranous form which often becomes narrowly lacin-
iate, with the segments much prolonged. V. andersont-
anum, which has been instanced as an example of a finely
dissected frond, becomes, when growing in quiet water, very
broadly linear, with the amount of branching largely reduced,
and the expanded branches very regularly arranged.

Form and Branching as Specific Characters.—Some allu-
sion has been made in the foregoing pages to the value of
such characters as form and branching of the erect frond
for distinctive purposes in describing a species. The ques-
tion was raised then as to the advisability of making too
narrow limits for a species upon such distinctions as form
and branching, without regard to the effect of environ-
mental factors. ‘The remarkable variations in these two
respects existing within the limits of a single species empha-
size this fact, and further call attention to the influence of
environmental relations upon form.

Stalk.—The stalk, which characterizes many species,
varies in length, width and thickness. The plant, by vari-
ations in the length of the stalk, may be sessile, subsessile,
or long-stalked. In some forms the stalk is narrow and
somewhat thickened; in others it is furnished with a thin,
expanded margin. It may vary in thickness from a few
cell-layers to a thick, almost fleshy tissue. A midrib, when
present, usually runs through the median portion of the
stalk. Increase in thickness may take place through a growth
of the superficial cells, or through an increase in the number
of cells composing the central layer. Frequently the thin
margin becomes worn away, and this is accompanied by an
increase in thickness of the median portion of the frond, so
that the stalk becomes cylindrical.

Midrib.—The midrib presents considerable variation in
form and extent. In some species it may be distinguished
only as a slight thickening of the median portion of the
stalk or frond, while in others it is more highly differenti-
ated, appearing as a ridge of considerable prominence. It

Bor.—VOL. II.] NOTT—CALIFORNIAN NITOPHYLLA. II

may be either simple and unbranched, as is commonly the
case in JV. multilobum, or it may become considerably
divided, as in WV. ruprechtianum. Frequently it does not
extend in the frond beyond the upper portion of the stalk
or lower segments. In other cases its ramifications reach
out into the branches almost to their tip, and there evan-
esce, or divide into nerves.

Nerves.—These structures are a characteristic feature of
the frond in several species, reaching a high degree of
development in some forms. In JV. /atiss¢mum no midrib is
present, but the large membranous frond is supported by
a network of intersecting nerves and veins of considerable
prominence. The other species are not distinguished by
such a full development of these structures. Usually the
nerves are limited to the outer margins or apices of the
frond, where they become flabellate or anastomose freely
with each other.

Veins.—More or less conspicuous veins constitute a note-
worthy structural element in some species, especially in JV.
ruprechtianum and JV. violaceum. In these plants, particu-
larly where the frond is at all flabellate, a rich development
of the finer venation may be seen, whose ramifications
extend in a flabellate fashion throughout the frond, or, in
some cases, anastomose with one another, finally becoming
free.

In WV. fryeanum, NV. uncinatum, and LV. corallinarum the
midrib and conspicuous veins are entirely wanting and the
only trace of venation is seen in the microscopic veins
which characterize these species. These minute structures
are frequently very delicate and invisible to the naked eye.
They extend as a rule throughout the frond, branching
freely or anastomosing. In JV. fryeanum, however, the del-
icate veins, in nearly every case, become somewhat stouter
toward the base of the erect frond, where they form a more
or less conspicuous fan-shaped area. A single species, JV.
speciabile, is destitute of any sort of veins.

Variable and Fixed Characters.—It has been said in a
preceding portion of this paper that stress should be laid

12 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

upon the greater value of certain characters as specific
distinctions and the less value of others. The more variable
characters there mentioned have been sufficiently discussed,
and the less variable, more important specific distinctions,
based upon the character of the venation and shape and
position of the sori, may here be treated.

Venation.—The Californian Vztophylla may be separated
into groups, distinguished from one another by the charac-
ter of the venation. In that group which includes the
greater number of forms, the species possess a midrib,
nerves, and conspicuous veins, developed to a greater or
less extent according to the species. Some forms exhibit
only the midrib, which passes over at once into the undiffer-
entiated frond as in /V. andersonianum. Others, again, are
provided with a midrib which divides into more or less con-
spicuous nerves, the latter passing again into the ordinary
frond, ¢. g., WV. harveyanum and LV. multilobum. Still
other forms, such as JV. vzolaceum and JV. ruprechtianum,
show a full development of midrib, nerves, and veins, the
last named structures usually conspicuous, and either anasto-
mosing or remaining free and flabellate. A second group
consists of forms in which no midrib is present, but the sur-
face of the frond is marked by a network of reticulate
nerves and veins, as in JV. /atisstmum. 'The forms of the
third group, including VV. fryeanum, NV. uncinatum and JV.
corallinarum, are destitute of midrib and nerves, and are
provided with scarcely perceptible, usually microscopic
veins, which either anastomose with one another, or remain
free. A fourth group comprises forms which wholly lack
venation of any kind whatsoever, as ¢. 9., lV. spectabile.

Sort.—The sporangia of JVitophyllum are gathered
together into sori of varying shape and size. These latter
structures may be employed as reliable specific distinctions
in discriminating between species. The Californian plants,
taken as a whole, show a considerable range of forms as
regards the shape of the sorus, and likewise a considerable
variety in its position. Within the limits of certain species,

e. g., IV. violaceum and JV. ruprechtianum, these variations

Bot.—Vou. II.} MNO77—CALIFORNIAN NITOPHYLLA. 53

are remarkable. Notwithstanding this fact, the variations
can be so expressed in terms of size, shape and position
that clear distinctions can be drawn between species. In
some cases the sporangia occur in narrow lines of varying
width arranged in a flabellate fashion on the upper divisions
of the frond, extending perhaps from its median portion to
the margin—their regular position in /V. harveyanum and a
disposition frequently seen in JV. ruprechtcanum—perhaps
only found as a fringe just within the margin, as often
observed in JV. ruprechtcanum. Other forms have the sori
in linear strips or patches along the margin of the frond, an
arrangement best seen in /V. vzolaceum. In another spe-
cies, JV. mu/tzlobum, the sori are irregular in shape, rounded
or linear-elliptical, with their longer dimensions extending
transversely across the frond upon whose upper segments
they are borne. The sori occur in other instances as
rounded or elliptical patches borne either singly upon distal
lobes, as in /V. andersontanum, or scattered irregularly over
the surface, as in JV. spectabile, sometimes displaying a ten-
dency to arrange themselves into lines, as is well shown by
LV. fryeanum. In yet other plants the sori are minute,
rounded structures, densely aggregated between the anas-
tomosing nerves and veins of the frond, e. o., LV. datissemum.

Proliferation.—The capacity for proliferation possessed
by some species is very great. To such an extent does
this phenomenon take place, that specimens are often
found in which the primary frond has remained compara-
tively undivided, and has produced from its margin numer-
ous proliferations which exceed in size the primary frond
itself. The favorite point for the production of prolifera-
tions is along the margin of the frond, especially after this
has been weathered by the waves. In such cases the
original frond may be reduced to a narrow strip, which will
produce proliferations surpassing in size the original portion
of the plant. In some species, particularly in LV. ruprechtz-
anum, the frond becomes reduced to the midrib; this per-
sists for a longer or shorter period, and proliferates very
freely, so that by this process the plant practically becomes

14 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

perennial. Some species, again, notably JV. vzolaceum and
LV. ruprechttanum, bear marginal proliferations in great
numbers, upon which appear the sporangia and cystocarps.

Antheridia and Cystocarps.—The antheridia and cysto-
carps need be mentioned only in a cursory way, inasmuch
as they take no prominent part in the identification of the
species. The antheridia first make their appearance as
whitish patches distributed rather evenly over the surface
of the frond. As they mature, the frond becomes wrinkled
or rugose to a marked degree. Antheridial plants have
now been found in three species, viz., VV. fryeanum, JV.
latissimum and lV. spectabtle, in considerable abundance.
The cystocarps are conspicuous structures, making their
appearance irregularly over the surface. In some species
the most weathered or reduced plants seem to be favored in
the production of cystocarps; and in this case the latter
sometimes occur in clusters of proliferations produced from
the surface or worn margins of the frond.

Parasites.—In only one instance have the plants of /Véto-
phyllum been found to harbor parasites. As a rule, the
species are quite free from epiphytes and parasites.

Gonimophyllum buffhamz Batters has been found by the
writer (1897, p. 81) upon JV. ruprechtianum. The plants are
a pale pink color, contrasting strongly with the deep lake red
of the host plant. The parasite seems to prefer the lower.
portions of the frond as its habitat, occurring on the /Vto-
phyllum near its base much more frequently than elsewhere.
It grows in patches, extending in some cases an inch in
breadth, while the individual fronds reach a height of ten
millimeters. So far, Gonzmophyllum has been found only
upon the tetrasporic JV. ruprechtianum.

Agardh (1898, p. 88) is of the opinion that Gonzmophyl-
Zum is a monstrous form of /V. daceratum. He suggests
further that JV. /aceratum occasionally may be hermaphro-
ditic, contrary to the normal course of development, and that
this monstrous form may be the structure concerned in the
phenomenon.

Geographical Distribution of the Genus.—The genus
LNitophyllum is distributed universally throughout the oceans

Bor.—Vo.. II.]} MWOTT—CALIFORNIAN NITOPAYVLLA. ES

of the globe. It has been found in the Arctic and Antarc-
tic oceans, along the shores of the continents bordering
upon the Atlantic and the Pacific, in the Mediterranean Sea
and the Indian ocean, and in the waters of Australia. The
coast of the British Islands is particularly rich in species,
while some of the finest examples of the genus come from
Australia.

About seventy species are known, of which number ten
occur on the west coast of North America. Of the ten
species thus credited, eight are limited to California or
neighboring shores. A ninth species, /V. wncenatum, is one
of the oldest known forms, and is more widely distributed,
being found in the Adriatic Sea, on the coast of Australia,
in California, on the shores of Europe, in the upper Atlan-
tic and Mediterranean Sea, and in New Zealand waters.
The tenth species, V. harveyanum, is an inhabitant of New
Zealand coasts.

flabitat.—The species of /Vztophyllum occur on other
alge and on Phyllospadix; on bare rocks exposed to the
dash of the waves or protected from their violence; or upon
rock surfaces which have become thickly coated with Cor-
allines or various Porifera and Bryozoa; or upon the piles
of wharves on the surface of the wood. The plants range
from the littoral zone between tide marks to the sublittoral
zone and have been dredged at a depth of 12-15 fathoms.

IV. DESCRIPTION AND DISTRIBUTION OF SPECIES.

Kry TO THE CALIFORNIAN SPECIES OF NITOPHYLLUM.

Frond provided with midrib only, or with midrib and nerves, or midrib,
nerves, and usually conspicuous veins.
Frond with midrib only, margin of frond serrate or dentate, sori in
rounded patches on distal lobes of frond.
NN. andersonianum (32).
Frond with midrib and nerves, but without veins.
Sori forming flabellate lines on upper segments of frond.
N. harveyanum (29).
Sori forming transverse patches on upper segments of frond.
N. multilobum (27).

16 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

Frond with midrib, nerves and usually conspicuous veins.

Color a pale violet through bright violet red to purple violet ;
thin and papery in texture; sori forming narrow flabellate lines
on upper divisions of frond, or linear patches along margin,
or rounded patches borne on the crispate margin or on mar-
ginal proliferations, or appearing in both lines and patches.

N. violaceum (39).

Color a bright red to blackish red; firm and leathery in texture ;
sori forming wide flabellate lines on upper divisions of frond,
or rounded patches on marginal or surface proliferations, or
appearing in both lines and patches... NV. ruprechtianum (34).

Frond without midrib, but provided with network of conspicuous nerves and
VEINSES sheet Baie 2 ete cle n nd bei eee sv etaes eee NV. latissimum (16).
Frond without midrib, nerves, or conspicuous veins, but provided with
microscopic veins.

Frond ample, erect, membranous below, branching above; sori form-

ing elliptical patches scattered over surface of frond.
N. fryveanum (22).
Frond erect, branching throughout, apices of ultimate branches re-
curved or hooked ...........02057 s2..2:-s 4. LV: UNCINAMII, (26):
Frond minute, creeping, ultimate lobes becoming free; sori forming
rounded patches on free lobes; on Corallina chilensis (and

other alece:?):. es. enigercce secon eee NV. corallinarum (24).
Frond destitute of veins, membranous; sori forming elliptical patches scat-
terediover sutiace ob irond.. 2) seca eee N. spectabile (21).

N. latissimum 7. Ag.

Epicrisis Floridearum, Contin. Spec. Gen. et Ord. Alg., 1876, p. 464.

Hymenena latissima Harvey, W. H., Proc. Linn. Soc., (Botany) Vol. VI,
1862, p. 170.

N. latissimum AGARDH, J. G., Bidrag Florid. Sys., 1871, p. 49. FARLow,
W. G., Proc. Amer. Acad. Arts and Sci., Vol. X, 1875, p. 365; Report
U. S. Fish Comm. for 1875, p. 695, 1876.

N. areolatum Eaton, D. C., Farlow, l. c., p. 695.

N. latissimum AGARDH, J. G., Epicrisis Florid., Contin. Spec. Gen. et Ord.
Alg., 1876, pp. 464 and 699. FARLow, W. G., Proc. Amer. Acad. Arts
and Sci., Vol. XII, 1877, p. 238. FARLow, W. G., ANDERSON, C. L.,
and Eaton, D. C., Algze Amer.-Bor. Exsiccatz, No. 68, 1878. HERVEY,
A. B., Sea Mosses, 1881, p. 175. ANDERSON, C. L., Zoe, Vol. II, 1891,
p. 224. Howe, M. A., Erythea, Vol. I, 1893, p. 68. McCCLATCHIE, A.
Jj., Proc. So: Cal. Acad: Sci., Vol..1,: 1897, p. 358. Norr, C22
Phyc. Bor.-Amer., CoLLtns, F. S., HoLpen, I., and SETCHELL, W. A.,
Fasc. VII; No. 335, 1897. TILDEN, J. E., American Algz, Century III,
No. 212, 1898. AGARDH, J. G., Contin. Spec. Gen. et Ord. Alg., Vol.
lil, Pt 3; 1898; p- 83:

N. macroglossum AGARDH, J. G., l. c., p. 84.

Bot.—VOL. II.] NOTT—CALIFORNIAN NITOPHYLLA., uy |

Synopsis.—Frond both prostrate and erect. Prostrate frond creeping,
slender, linear, much branched; without rhizoids, midrib, nerves, or veins;
margin serrate or toothed.

Erect frond subsessile or shortly stalked, flat, membranous; without mid-
rib, but with numerous, conspicuous nerves and veins; palmately divided or
segmented, frequently proliferating; with margin usually entire, occasionally
minutely serrate. Segments linear or wedge-shaped, usually expanded and
lobed at apices. Stalk, when present, usually short, occasionally longer,
formed by the wearing away of frond on margins at base and subsequent
thickening of nerves and remaining portion of frond. Nerves and veins
numerous, conspicuous, extending over almost entire surface of frond, evan-
escent in outer lobes, branching and anastomosing, frond thus conspicuously
areolate.

Proliferations usually numerous, especially upon more reduced _ portions of
frond, membranous, segmented or lobed, nerved and veined.

Sporangia in minute sori scattered over both surfaces of the frond between
the nerves and veins, in the conspicuous areoles.

Antheridia forming dull whitish patches over the entire surface, the latter
becoming ridged, indented, or rugose.

Cystocarps numerous, large, irregularly disposed over both surfaces, pro-
jecting beyond the surface. 5

Femarks on the Species.—The color of LV. latessimum
does not change to any extent when dried. In the living
state the tint is a deep, rich, lake red, becoming a shade
darker in the dried condition. Plants frequently occur
which attain a length of 35-40 cm., with the frond divided
into numerous segments from 3-5 cm. in width. The spe-
cies is one of the handsomest and most luxuriant in habit
of any of the forms inhabiting the coast.

NV. latissimum is characterized by its broadly expanded
fronds, divided into large, rounded lobes and segments, and
by the extensive network of prominent nerves and veins
covering the surface. This conspicuous network gives to
the surface a markedly areolated appearance, which serves
to distinguish the plants from all others of the genus as
represented on Californian shores. The frond frequently
proliferates along the margin, thus giving rise to numerous
ovate or lanceolate proliferations, which possess a well
marked venation and often produce sporangia and cysto-
carps. ° Proliferations likewise appear freely from reduced
portions of the frond.

(2) : ; July 26, 1900.

18 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

The creeping, much branched, claw-like, prostrate frond
possessed by JV. /atzsstmum is an excellent adaptation for
the purpose of enabling the plant to retain its foothold upon
the rough and seamy piles and rock surfaces which are its
habitat. Very little variation has been observed in the
prostrate frond; and it seems to be a well defined and spec-
ially differentiated structure.

Tetrasporic plants are of the most common occurrence.
Sporangia are produced very abundantly over the entire
surface, between the nerves and veins, thus adding to the
distinctive areolate aspect of the plant. The sori are minute
and crowded together, giving the impression of large sori
completely filling the areoles. Antheridial specimens have
been observed in but one locality. They reacha large size,
being among the most magnificent examples of the species.
In the early stages of development the antheridia form pale,
whitish patches. Later, they become more evident, and
when well developed, cause the entire frond to appear dull
reddish white. The surface of the frond is then decidedly
rugose.

A comparison of JV. dat¢sscmum with a specimen of JV.
hillie Grev., distributed by Le Jolis in Algues Marines de
Cherbourg, No. 215 (in herbarium W. A. Setchell), shows
a striking similarity between the two species in texture,
venation, and character of lobes or branches. A series of
specimens of JV. Az//ze@ in the herbarium of Professor W. G.
Farlow shows that this plant varies greatly as to prominence
of veins. Careful comparisonof JV. /atrsstmum with strongly
veined specimens of /V. Az/ii@ brings out a strong resem-
blance between the two. With the material in hand, how-
ever, it is not advisable to do more than point out the possi-
bility that the two species may be identical.

It is desirable to call attention at this point to the latest
views of Agardh (1876, pp. 464 and 699, 1898, pp. 83-85)
concerning iV. Jattssemum.

According to Agardh, JV. /atisstmum is a form which was
first collected by David Lyall at Vancouver Island, B. C.,
and described by Harvey (1862, p. 170) under the name

50

Fig.

. LO.

= ok Tre

12.

CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

EXPLANATION OF PLATE III.

Nitophyllum fryeanum,

Cystocarpic plant, showing disposition of cystocarps and segments
of frond. One-third natural size.

Tetrasporic plant, showing palmate frond with much prolonged
segments, arrangement of sori and delicate veins. One-third
natural size.

Tetrasporic plant, showing venation and attenuate segments of
frond. One-third natural size.

Cystocarpic plant, showing pinnately arranged segments of frond.
One-third natural size.

Nitophyllum corallinarum.

Tetrasporic plant, showing frond creeping over branches of Coral-
lina chilensis, and sori borne on free, erect fronds, é. g., at x
and x’. One-half natural size.

Nitophyllum multilobum.

Tetrasporic plant, showing habit and characteristic transverse sori.
One-half natural size.

Nitophylium uncinatum.

Sterile plant, showing character of branching and characteristic
recurved apices of branches. One-half natural size.

5}
4
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DA

S
L

OPHYLLUA

7
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Figs. 6-9.

ap

j

Po

Fie.10. NITOPHYLLUM

in)

52

Fig.

5 ES:
g. 14.

16.

Ey:

CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

EXPLANATION OF PLATE IV.

Nitophyllum harveyanum.

Slender form, tetrasporic, showing habit. One-half natural size.
Robust form, showing linear sori and flabellately expanded habit.
One-half natural size.

Nitophyllum ruprechtianum.

Prostrate frond of the stout membranous type. One-half natural
size.
Nitophyllum fryeanum.

Prostrate frond of the delicate membranous type, showing rhizoids
as minute processes on surface. One-half natural size.

Nitophyllum andersonianum.

Prostrate frond, of the freely branching type, with portions of the
bases of erect fronds. One-half natural size.

Fig.l5. N

i

TOPHY

IM HARVEYANUM.

RUPRECHTIANLIM

PHOTO -TITH. BRITTON & REY, 5.F

Ct RYEANUM

FHYLLUM ANUERSONIANUM

DP Tisaltyt

a. ®

Stee

- 19:

. 20.

. 21.

CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

EXPLANATION OF PLATE V.

Nitophyllum andersonianum.

Robust form, sterile, showing usual character of branching, with
branches flabellately expanded at apices. One-third natural size.

Broad membranous form of quiet waters, sterile. One-third natural
size.

Slender form, sterile, showing fine dissection of frond. One-third
natural size.

Transition form, tetrasporic, much branched, showing tendency to
becomeSmembranous. One-third natural size.

ER
SONI
TIAN
IL]
M

NU

Nene
Whe
PHY
TYLLUM
Al

*
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Mie:
Sor ty |

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56

Fig.
Fig.

Fig.

Fig.

CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

EXPLANATION OF PLATE VI.

Nitophyllum ruprechtianum.

Expanded, membranous frond, sterile. One-fifth natural size.

Examples of weathered and proliferating plants. One-fifth natural
size.

Slender, much prolonged type, minutely proliferating along margin,
sori on proliferations. One-fifth natural size.

Typical plant, showing stalk, branching, venation, and proliferations,
with sori on proliferations. One-fifth natural size.

—

i.
_

he
_-+%
i

58 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

EXPLANATION OF PLATE VII.

Nitophyllum ruprechtianum.

Fig. 26. Typical plant, showing stalk, branching, venation, proliferations,
and sori on proliferations. One-third natural size.

Fig. 27. Portion of frond, showing flabellately disposed sori. One-third

natural size.

Fig. 28. Same, showing sori (linear) confined to apices of branches. One-
third natural size.

Fig. 29. Weathered frond, cystocarpic, with prostrate frond. One-third
natural size.

Fig. 30. Portion of frond, showing linear sori confined to ends of branches
and rounded sori on marginal proliferations. One-third natural
size.

60

Wane

-030:
See

CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

EXPLANATION OF PLATE VIII.

Nitophyllum violaceum.

Intermediate form, showing segments of frond, with crispate mar-
gin and marginal sori. One-fifth natural size.

Typical form, showing slender, linear branches, marginal prolifer-
ations, and sori forming linear patches along margin or rounded .
patches on the proliferations. One-fifth natural size.

Intermediate form, with branches flabellately expanded at ends;
sori as in preceding figure. One-fifth natural size.

Robust, membranous form, sterile. One-fifth natural size.

Reduced and weathered form, the conspicuous sori placed singly
or together along the margin, or borne on proliferations. One-
fifth natural size.

Weathered form, as in preceding figure. One-fifth natural size.

Finely dissected form. One-fifth natural size.

VITTON & REY, £F

“ITH

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Fig.

Fig.

ig. 38.

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45.

. 46.

CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

EXPLANATION OF PLATE IX.

Nitophyllum ruprechtianum.

Portion of tetrasporic plant, showing rather wide, linear, flabellately
disposed sori. Four-fifths natural size.

Same, showing transition from linear, flabellate sori to rounded sori
borne on marginal proliferations. Four-fifths natural size.

Same, showing sori borne only on marginal proliferations. Four-
fifths natural size.

Nitophyllum violaceum.

Portion of tetrasporic plant, showing narrow, linear, flabellately
disposed sori. Four-fifths natural size.

Same, showing transition from linear, flabellate sori to rounded,
inframarginal sori. Four-fifths natural size.

Same, showing rounded inframarginal sori. Four-fifths natural
size.

Nitophyllum latissimum.

Detail of tetrasporic sori, showing venation, and areoles occupied
by minute sori. Four-fifths natural size.

Nitophyllum andersonianum.

Portion of tetrasporic plant, showing sori as rounded or elliptical
patches on distal segments of frond. Four-fifths natural size.

Nitophyllum multilobum.

Portion of plant, showing characteristic transverse sori. Four-fifths
natural size.

ATTITRE

LAAN

UPRECHTI

titt

Bor.—Vou. II.} MO7T7—CALIFORNIAN NITOPHYLLA. 27

linear and extend in some cases a considerable distance
along the margin. When compared with JV. harveyanum,
which also bears linear sori arranged likewise in flabellate
fashion, it is seen that, while the resemblance in the pro-
duction of the sori is strong, /V. harveyanum possesses a
darker purplish tint, is not so robust in habit, and is desti-
tute of such a well developed system of venation as belongs
to LV. ruprechtianum.

The agreement between JV. ruprechtianum and JV. viola-
ceum in the production of sori and venation is much more
marked, yet the two can be distinguished by the character-
istic violet hue and papery texture of JV. vzolaceum, in con-
trast to the dull red color and leathery texture of JV.
ruprechtianum.

The same variety of position as is seen in the case of the
sori may also be observed in the cystocarps. These struct-
ures may be borne by JV. ruprechtianum, either upon the
surface, when they are scattered at irregular intervals over
the entire surface or form a border just within the margin,
or they may be borne singly or several together on surface
or marginal proliferations.

The existence of such numerous transition forms between
the two extremes of surface and marginal production of the
sori, a fact clearly established by a careful examination of
material from a long extent of coast, points to the conclu-
sion that within the limits of the species known as JV.
ruprechtianum it is possible to include a wide range of
forms characterized by the features already pointed out,
and that sufficient grounds do not exist for the establish-
ment of several species among which these forms may be
distributed.

The evidence afforded by these transition forms should,
therefore, be employed in examining the species estab-
lished by Agardh (1876, p. 699; 1898, pp. 93-96), viz.,
NV. flabelligerum, N. marginatum and JV. farlowianum,
which,-so far as the writer can determine from the de-
scriptions, have been split off from JV. ruprechtianum.
The characterization of these three species is made to

38 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

depend upon differences in texture, branching, venation
and sori. In the foregoing pages an attempt has been
made to show the amount of variation possible in these
characters, and to emphasize their relative value for spe-
cific purposes. The numerous collections made between
widely distant limits along the coast at various points and at
different seasons of the year, as well as all the available
material at hand in herbaria, have failed to yield forms
which could not be referred to JV. ruprechttanum as defined
in the foregoing account of the species.

A word with respect to the general shape and segmenta-
tion of the frond may be added in regard to the variation
existing in JV. ruprechtianum. Forms may be found that
tend to become elongated, with much prolonged, narrow, and
acute segments. Near to these may be placed forms like-
wise with the segments prolonged, but flabellately expanded,
and with rounded apices. The tendency to expansion is
seen to best advantage, finally, in plants that become divided
into a few broadly wedge-shaped segments, with these in
turn somewhat deeply lobed, with the lobes rounded as in
the flabellate type.

Flabitat.—V. ruprechtianum especially occurs on rocks
usually covered with Corallines or Bryozoa and Porifera,
among which the prostrate frond attains a rich develop-
ment, in deep rock pools on gently sloping shores, littoral
to sublittoral zones.

Distribution.—Along the coast from San Diego, Califor-
nia, northward to Port Orchard, Washington.

Localities. —San Diego! (D. Cleveland); Point Loma!
(Miss Minnie Reed); La Holla! (Miss Minnie Reed);
San Pedro! (Mrs. S. P. Monks); Santa Barbara! (Dr.
and Mrs. L. M. Dimmick; Mrs. S. P. Cooper); shore of
San Luis Obispo County! (Mrs. R. W. Summers); Port
Harford! (W. A. Setchell); Avila Beach! (Miss Mabel
Miles); San Simeon! (E. Palmer); Carmel Bay! (C. P.
Nott); Pacific Grove! (M. A. Howe; Mrs. J. M. Weeks;
C.. P.. Nott); Santa. Craz! ((Mrs. Boston; C. L. Ander
son; Mrs. J. M. Weeks); Duxbury Reef! (W. A. Setchell) ;

Bor.—VOL. II.] NOTT—CALIFORNIAN NITOPAVLLA. 39

Dillon’s Beach! (W. A. Setchell); Fort Ross! (W. A.
Setchell; C. P. Nott); Port Orchard, Washington! (J. E.
Tilden).

Nitophyllum violaceum 7. Ag.

Epicrisis Floridearum, Contin. Spec. Gen. et Ord. Alg., 1876, p. 700.

Nitophyllum laceratum Harvey, W. H., Ner. Bor.-Amer., Vol. II, 1858, p.
Io4. FARLOw, W.G., Proc. Amer. Acad. Arts and Sci., Vol. X, 1875,
p. 365; Report U. S. Fish Comm. for 1875, p. 695, 1876.

Nitophyllum violaceum AGARDH, J. G., Epicrisis Floridearum, Contin. Spec.
Gen. et Ord. Alg., 1876, p. 700. FARLOw, W. G., Proc. Amer. Acad.
Arts and Sci., Vol. XII, 1877, p. 238. Herrvey, A. B., Sea Mosses,
188i, p. 180. ANDERSON, GL Ss Zoey Vol EH. 1891; p. 224. Howe,
M.A., Erythea, Vol. I, 1893, p. 68. .McCratcuir, A. J., Proc:.So. Cal.
Acad. Sci., Vol. I, 1897, p. 358. Nott, C. P., in Phyc. Bor.-Amer.,
Co uins, F.S., HOLDEN, I., and SETCHELL, W. A., Fasc. VIII, No. 389,
1897. AGARDH, J. G., Contin., Spec. Gen. et Ord. Alg., Vol. III, Pt. 3,
1898, p. 91.

Nitophyllum stenoglossum AGARDH, J. G., loc. cit., p. 92.

Neuroglossum lobuliferum ? AGARDH, J. G., loc. cit., p. 121.

Nitophyllum violaceum formum crispulum SETCHELL, Phyk. Bor.-Amer.
Couns, F. S., HoLtpsen, I., and SETCcHELL, W. A., Fasc. XIV, No.

694, 1900.

Synopsis.—Frond both prostrate and erect. Prostrate frond creeping,
broadly linear, or membranous, with rhizoids, branching, without midrib or
nerves; margin toothed or laciniate.

Erect frond stalked, flat, linear, occasionally with midrib, with flabellate
nerves; subdichotomously or subpalmately divided into numerous segments,
in some cases finely laciniate, frequently proliferating; margin entire, finely
serrate, crispate, or toothed; segments or branches in some cases broadly
obcuneate, in other cases narrow, becoming broadly linear, much prolonged
and flabellately expanded at apices or remaining linear. Stalk short, soon
merging into the frond, usually without midrib, but with flabellate, sometimes
anastomosing nerves extending into the branches; or long, somewhat nar-
row and thickened, almost cylindrical, and again merging into the flabellately
nerved branches. Nerves not very conspicuous, usually extending through-
out the frond from base nearly to apex, becoming divided into flabellate,
frequently anastomosing veins, the latter soon evanescent.

Sporangia in narrow lines flabellately disposed, occasionally confluent, on
upper segments of frond, or placed singly or in clusters along the margin, or
upon marginal sporophylls, the latter appearing at intervals or in a dense
fringe along the margin. Antheridia as yet unobserved. Cystocarps large,
projecting, irregularly disposed over both surfaces, or submarginal, or upon
marginal proliferations.

Peemarks on the Species.—The color of V. violaceum
varies from a pale violet through bright violet red to purple
violet, in both the living and dried states. The plants attain

40 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

a height of 20-25 cm. Though not so robust in habit as
NV. ruprechtianum, still some specimens of JV. vzolaceum
are vigorous in their growth, branching freely and forming
large, handsome plants. The plant is papery or parchment-
like in texture, rather brittle when dried, and does not
adhere well to paper.

NV. violaceum, as before remarked, agrees with JV.
ruprechttanum in many morphological details, such as char-
acter of branching, general shape and position of sori, and
venation; but it may be distinguished from that species by
its different color and texture, and the minute differences
in size and shape of the sori.

An extremely wide range of variation may be seen in
the amount of dissection which the frond of JV. v7olaceum
undergoes. On the one hand, there is found a form in
which the frond becomes divided from the base into a great
number of slender, much prolonged branches, which divide
again and again, until finally the apices of the ultimate
branches are prolonged in a flabellate fashion sufficiently to
show the specific characters of venation and color. On
the other hand, there occur forms rather broadly membra-
nous at base, that divide into a few broadly obcuneate seg-
ments cleft from the outer edge into narrower portions
which are prolonged into lobes, again exhibiting the charac-
ters of the species. Between these extreme types may be
found intermediate forms that in some cases vary toward
the finely dissected frond, in others, toward the broadly
membranous frond. Stunted and weathered plants also
occur in considerable numbers in late winter and spring.
The frond in these instances is occasionally thickened and
rather fleshy, while the sori are gathered into clusters of mar-
ginal proliferations or remnants of the former margin. Pro-
liferation takes place, though not to the same extent as in
LV. ruprechttanum. The proliferations are, as a rule, small,
and of varying size. ‘They usually bear sori.

In common with JV. ruprechtianum, LV. violaceum chal-
lenges attention by reason of the peculiarities of its sori,
which exhibit again the same range of variation in regard

4

Bot.—Vou. II.]. MWOT7T7—CALIFORNIAN NITOPHYLLA. 41

to their shape and position that was observed in JV. ruprecht-
zanum. There are to be found forms with the sori
arranged in flabellate, occasionally confluent lines, differ-
ing, however, from JV. ruprechtianum in the relative nar-
rowness of the lines. Numerous transition forms occur
which combine the flabellate linear sori with rounded or
linear patches upon the margin. Finally, frequent instances
may be found of plants bearing the sori as rounded or
semicircular patches along the margin, or upon marginal
proliferations.

The same conclusion that was reached in regard to the
limits of the species in the case of JV. ruprechtianum may
be applied to /V. vzolaceum, for here almost exactly similar
conditions exist as to specific characters. The variations
are in similar directions. Extended study of the forms from
numerous localities has shown that they may be referred
along the lines there laid down, which leads to the conclu-
sion that /V. vzolaceum is a species with limits sufficiently
wide to include all the forms common to the coast.

The two species established by Agardh (1898, pp. 92 and
121), viz., lV. stenoglossum and Veuroglossum lobuliferum,
seem, therefore, to the writer, as nearly as can be deter-
mined from the descriptions, to be forms of JV. vzolaceum.

Hlabitat.—On rocks covered with Corallines, Bryozoa
and Porifera, in sheltered situations or in rock pools on
gently sloping shores, and on piles of wharves, in the
littoral and sublittoral zones.

Distribution.—Along the coast from San Pedro north-
ward to Fort Ross.

Localities.—San Pedro! (Mrs. Lawrence; Mrs. 8S. C. .
Purdy); White’s Point! (A. J. McClatchie); Santa Bar-
paca!) (Dr.-L. N.Dimmick; Mrs: *S: P. Cooper); San
Simeon! (E. Palmer); Carmel Bay! (C. P. Nott); Pacific
Grove! (M. A. Howe; C. P. Nott); Santa Cruz! (Dr. C.
L. Anderson); Land’s End, San Francisco! (C. P. Nott);
Fort Point, San Francisco! (M. A. Howe; W. A. Setchell;
C.P. Nott); North Beach, San Francisco! (W. A. Setchell;
C. P. Nott); Golden Gate, San Francisco Bay (Berggren,

42 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

fide J. G. Agardh, Epicrisis Floridearum, 1876, p. 700);
San Francisco! (G. W. Lichtenthaler); Lime Point, San
Francisco Bay! (C. P. Nott); Duxbury Reef! (W. A.
Setchell); Fort Ross! (C. P. Nott).

By way of summarizing some of the features mentioned
in the foregoing account, the following table, showing the
distribution of species within Californian limits, is incorpo-
rated. It will be seen that ten species occur on the coast.
These have now been reported from twenty-eight localities,
ranging from San Diego northward to Puget Sound, Wash-
ington, and points on Vancouver Island, B. C., embracing
twenty degrees of latitude and fourteen hundred miles of
coast line. It may be pointed out that there is here a
coastal distribution equal to that of the Atlantic shores of
Europe and the Mediterranean together.

In the light of the evidence collected from a wide range
of material in field and herbarium, the ten species recog-
nized may be considered valid until more extended com-
parison with European specimens proves their identity with
previously described species.

a

Bot.—Vou. II.} MO7T7—CALIFORNIAN NITOPAYLLA.

TABLE OF DISTRIBUTION OF SPECIES.

(San Diego, Calif., northward to Esquimault Bay, B. C.)

|
|
|
|
|
|

“UWN~DULIUN * AT |
|
"Uu6Nn991D) 020 * AT |

e(e| =| s|s)5[=| =
So eel Gas ine lide ses |e. fe
S$ S Se ilies SAN Poe see | SE
Sails ees eee | ee |
= = = ~ 3 3 Spy se
S |

Sat NEC Ose Je. nae oes * | cart

Rointg eOma etree, «cae *

| We'] C0) | Ra a #

Samp meanOne asee ee les % *

Wihite:s"POnt. = 2.4 sss.

Satta Monica he jas ss

Sailtaebanbabas- sine eee a: a 2 7

San Luis Obispo County.) * _ =

One larorden cbse *

Jain el BRYa 1 ag ae a gree Ne Ss

Sany Simeon. sas cs). - %

CarmelsBay asacecciccsts ze = #

Pacific Grove..........-. * He Se

Santay @ruZeate set a aciec cc * * % = * *

iLeiivelie IDG b AeAaadoee ner * %

oni hOitit. cs. eoa cis a - #

GoldeniGates... 6.2.02. we |x *

INofth) Beachinn. occ. .

SanVHranGisCO: «asses <0 a % x

WME BOIMtS + osc sscie cee * *

Duxbury Reef...... a ehetass * :

Willenssipeacht ee ce ee. 3 oS

OnE ROSS Usetls ss cesio nse. * * * *

Cape Mendocino........ 2

Klatsop, Oregon........ *

Port Orchard, Wash..... % *

Puget Sound, Wash..... * |

Esquimault Bay, B. C.... *

44 CALIFORNIA ACADEMY OF SCIENCES. [PRoC. 3D SER.

The writer is deeply indebted to Professor W. A. Setch-
ell, upon whose suggestion this paper was undertaken, not
only for collections of material and the use of his private
herbarium, but more especially for his discriminating advice
and helpful suggestions throughout the entire work, as well
as for a comparison of some coast forms with specimens
in the herbarium of Professor W. G. Farlow. Further
acknowledgment is made to Professor Farlow, who kindly
permitted an examination of the /V/ophylia in his collection,
and to Mr. F. S. Collins and Dr. C. L. Anderson, both of
whom courteously forwarded to the writer all the /Vzto-
pAylla contained in their herbaria. Acknowledgment should
also be made to Mr. Collins for the specimen from which
the description of JV. corallinarum was drawn up. To Mrs.
J. M. Weeks the writer is especially indebted for collections
which permitted a much more comprehensive account of
certain species, and also for the privilege of examining
specimens in her herbarium. The herbarium of the Uni-
versity of California was consulted frequently in the prepa-
ration of this paper. Special acknowledgment is due to
Professors O. P. Jenkins and G. C. Price for their courtesy
in extending to the writer during several seasons the privi-
leges of the Hopkin’s Seaside Laboratory at Pacific Grove,
California, thus facilitating the examination of a large
amount of living material, at different times in the year.
UNIVERSITY OF CALIFORNIA,

BERKELEY,
June, 1900.

Bot.—Vot. II.] MWO7TT—CALIFORNIAN NITOPAYLLA. 45

LITERATURE AND EXSICCATZ: CITED IN THIS PAPER.

1852.
1871.

1876.
1898.
1878.
1891.
1877.
1875.

1876.
1877.

1858.
1862.

1889.
1881.
1893.

1889.
1897.
1899.
1897a.
18976.

1897¢.
1897.

1900.
1898.

AGARDH, J. G. Spec. Gen. et Ord. Algarum, Vol. II, Pt. 2.

Bidrag till Florideernes Systematik. Lund’s Universitets

Arsskrife, Tome VIII.

Epicrisis Floridearum. Contin. Spec. Gen. et Ord. Algarum.

Contin. Spec. Gen. et Ord, Algarum, Vol. III, Pt. 3.

Algzee Americanz-Borealis Exsiccate. FARLOw, W. G., ANDERSON,
Cre vandee Aron yy Ge

ANDERSON, C. L. List of California Marine Algz. Zoe, Vol. II,
No. 3.

Eaton, D. C., in Farrow. Proc. Amer. Acad. Arts and Sci.,
Vol. XII, p. 245.

FarLow, W.G. List of the Marine Algz of the United States, etc.
Proc. Amer. Acad. Arts and Sct., Vol. X.

Id. Report U.S. Fish Comm. for 1875. ,

On Some Algze New to the United States. Proc. Amer. Acad.
Arts and Sct., Vol. X11, No. 20.

Harvey, W. H. Nereis Boreali-Americana.

Notice of a Collection of Algze made on the Northwest Coast of
North America, chiefly at Vancouver’s Island, by David Lyall, in
the years 1859-61. Proc. Linn. Soc., (Botany), Vol. VI.

Hauck und Ricuter. /hykotheca Universalis, Fasc. VII.

Hervey, A. B. Sea Mosses. ;

Howe, M. A. A Month on the Shores of Monterey Bay. Avythea,
WolSIs INow2:

KJELLMAN, F. R. Om _ Beringshafvets Algflora. XK. Svenska
Vetensk.-Akad. Handl., Tome XXIII, No. 8.

McCratcuir, A. J. Proc. So. Cal. Acad. Sct., Vol. I.

NoRDHAUSEN, M. Zur Anatomie und Physiologie einiger ranken-
tragender Meeresalgen. PringSheim’s Jahrbiicher f. wiss. Botanik,
Band XXXIV, Heft. 2.

Nott, C. P. Some Parasitic Florideze of the Californian Coast.
Erythea, Vol. V, No. 9..

In Phykotheca Borealis-Americane. COLuins, F. S., HOLDEN,

I. and SETCHELL, W. A. Fasc. VII.

kG | ase: WOE

Phykotheca Borealis-Americane. Co.uins, F. S., HOLDEN, I. and
SETCHELL, Wi At Fasc: Ville

Ine; Fase. XIV:

TILDEN, J. E. American Algz. Century ITI.

46 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

EXPLANATION OF PLATE I.

Nitophyllum latissimum.

Fig. 1. Tetrasporic plant of late spring, with broad, flat veins and large
areoles. One-third natural size.

Fig. 2. Cystocarpic plant, otherwise as in fig. 1. One-third natural size.

Fig. 3. Antheridial plant, mature, showing in the lower third the character-
istic wrinkled surface of the antheridial plant. One-third natural
size.

i
ii

Dixie vee
i@ Avena i

ee |

ue ry 7 of. :
" apy i 4 ay a! ME As |

Pe Vu ‘| ~

48 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

EXPLANATION OF PLATE II.

Nitophyllum latissimum.

Fig. 4. Typical cystocarpic plant, showing portion of prostrate frond, vena-
tion, proliferation, cystocarps, and a portion of the creeping,
claw-like prostrate frond. One-third natural size.

Nitophyllum spectabrle.
Fig. 5. Typical tetrasporic plant, showing prostrate frond, young and

mature erect fronds, and position and shape of sori. One-third
natural size.

ATISSIMUM.

A

A I

T
Ll

Fig.4. NITOFHYLLI

> *

=

Bor.—VOL. II.] NOTT—CALIFORNIAN NITOPHYLLA. 19

Hymenena Jatissima. A similar form was received some-
what later by Agardh from Golden Gate, San Francisco
Bay, collected by Berggren. Further than this, corre-
sponding forms were issued in the Ale. Amer.-Bor. Exsicc.
(1878, No. 68), collected by Dr. C. L. Anderson, at
Santa Cruz.

Again, according to Agardh, there is still another form,
viz., JV. macroglossum, which is referred to Californian
shores. This plant was collected originally by Berggren
also at the Golden Gate, and at first was included by Agardh
in JV. latissimum. Specimens collected by the writer (1897,
Fasc. VII, No. 335) at'the type locality are said by Agardh
to be identical with the plant first referred by him to JV.
Jatissimum and now included under JV. macroglossum.

The writer has now examined in the field and herbarium a
wide range of plants comprehended under these two species
of Agardh. Specimens, collected at numerous localities
and at various times in the year along the coast from San
Pedro northward to Puget Sound entrance, provide a fair
amount of material as a basis for estimating Agardh’s spe-
cies. An especially abundant series of forms was collected
by the writer between January and May at Golden Gate,
San Francisco Bay, the type locality for Agardh’s V. mac-
reglossum. This collection included numerous specimens
of both the species above referred -to as established by
Agardh.

A discriminating examination of Agardh’s descriptions
shows, as far as the writer can determine, that JV. Jatisstmum
differs from JV. macroglossum in being of a paler color,
possibly less luxuriant in habit, with by no means such a
well developed system of venation. The nerves and veins
anastomose less freely, project above the surface of the
frond less prominently, or not at all, as is more commonly
the case, and soon evanesce into the frond. The areoles
are larger and more conspicuous by reason of the broad,
flat aspect of the nerves and veins. The sori are fewer in
number and are most numerous about the borders of the
areoles.

20 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

The range of forms examined by the writer with these
points in mind exhibits, it is true, the structural differences
pointed out by Agardh. But further, it should be observed
that the species here in question, as field studies show, is
apparently aform occurring between December and August,
reaching its finest development in February and March, and
at that time exhibiting the characters of /V. macroglossum.
The forms collected from April to August show the charac-
ters of JV. datrssimum, with frequent occurrence of transi-
tion forms between the two species.

The evidence collected by the writer in field and herba-
rium points to the conclusion that JV. macroglossum and JV.
Jatissimum are seasonal variations of the same species, that
is, JV. latiss¢mum.

Habitat.—On piles of docks and wharves, or on sloping
rock surfaces where the wave action is not violent; lower
littoral to sublittoral zone.

Distribution.—At various points along the coast from
San Pedro to Vancouver.

Localities. —San Pedro! (Mrs. S. P. Monks); Santa
Barbara! (Dr. and Mrs. L. M. Dimmick; Mrs. S27
Cooper); shores of San Luis Obispo County! (Mrs. R. W.
Summers); Pacific Grove! (M. A. Howe; Mrs. J: XE
Weeks; ‘C: P. Nott);\Santa Cruz! (Dr. C;. L. Andersong
Mrs. J. M. Weeks); San Francisco! (G._W. Lichten-
thaler); San Francisco Bay entrance (Golden Gate) at
Fort,.Pomt! (M.A. Howe;:W. A. Setchells: W. yj
Osterhout; C. P. Nott); at Lime Point! (C. P. Nott); Fort
Ross! (W. A. Setchell); Klatsop, Oregon, (L. F. Hen-
derson, in Herb.; W. G. Farlow, fide W. A. Setchell) ;
Port Orchard, Washington! (J. E. Tilden); Esquimault
Bay, Vancouver Island, B. C. (Harvey, in Proc. Linn.
Soc., (Botany) -Vol. VI, 1862, p. 170; Farlow, Proc.
Amer. Acad. Arts and Sci., Vol. X, 1875, p. 365); Puget
Sound! (N. L. Gardner).

Bot.—Vou. II.]} MWO77—CALIFORNIAN NITOPHYLLA. 21

Nitophyllum spectabile DV. C. Eaton.

In FarLow, Proc. Amer. Acad. Arts and Sci., Vol. XII, 1877, p. 245.

Nitophyllum spectabile Farntow, W. G., Report U. S. Fish Comm. for 1875,
p. 695, 1876; Proc. Amer. Acad. Arts and Sci., Vol. XII, 1877, p. 238.
Eaton, D. C., in Farlow, |. c., p. 245. FARtow, W. G., ANDERSON,
C. L., and Eaton, D. C., Algze Amer.-Bor. Exsiccatz., No. 67, 1878.
Hervey, A. B., Sea Mosses, 1881, p. 174. ANDERSON, C. L., Zoe, Vol.
II, 1891,.p. 224. AGARDH, J. G., Contin. Spec. Gen. et Ord. Alg., Vol.
IIt, Pt. 3, 1898, p. 43-

Synopsis.—Frond both prostrate and erect. Prostrate frond thin, linear,
creeping, destitute of venation; becoming thickened when weathered; branch-
ing irregularly; branches rising into erect fronds.

Erect frond sessile or subsessile, flat, membranous; destitute of venation;
irregularly oblong, deeply pinnately lobed, occasionally palmately segmented,
sometimes Aeoleratine: margin entire, sinuate, or lobed. Segments linear,
lanceolate, ovate, or cuneate, frequently deeply lobed at apices.

Sporangia in elliptical sori, disposed at nearly regular intervals over both
surfaces of the frond. Antheridia in whitish patches over entire surface of
frond, giving to latter an areolate aspect. Cystocarps numerous, conspicu-
ous, irregularly disposed over both surfaces, projecting beyond the surface.

Remarks on the Species.—This plant retains, when dried,
the bright, rosy red hue which characterizes it in the living
state. The species is said by Professor Eaton, who estab-
lished it, to reach a length of 50-60 cm. It is one of the
largest and finest species of the coast. The general aspect
of the frond is much like that of VV. fryeanum. Kt differs
from that form, however, in not possessing any kind of
venation.

Comparison of JV. Eire with JV. ruthenicum (P. & R.)
Kjell. aroused a suspicion that the two forms might be iden-
tical. A more. careful examination showed that, in speci-
mens of JV. ruthenicum received from Professor Kjellman,
the plants were ‘‘obsoletely veined below,’’ as is stated in
the description of V. ruthenzcum (1889, p. 25, Pl. I, figs.
II-12), and as is shown in the figure. JV. sfectabzle, on
the contrary, is totally destitute of venation. All the evi-
dence at hand demonstrates that JV. spectadzle TSENG is
a distinct species.

The writer is indebted to Professor W. A. Setchell for
the following note upon JV. spectabzle, through the courtesy
of Professor W. G. Farlow, who kindly permitted an exam-
ination of his specimens of /Vitophyl/a in connection with

22 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

the preparation of this paper. Professor Setchell says,
‘‘Specimens in herbarium of Professor Farlow (ex. herb.
Acad. Petrop.) labelled ‘Ag/aophyllum ruthenicum, Exp.
Lutk. ad litora Americane borealis-occidentalis, Ross’ are
young (about an inch high, with no fruit present), and
might be the young plants of JV. spectabile. In the herba-
rium of Farlow are also several specimens from St. Paul
Islands, in Behring Sea (legit White), which might be JV.
spectabile.”’

The account of this species was advanced to a great
extent by examination of a large collection of material from
Mrs. J. M. Weeks, of Santa Cruz, Calif., who made an
especial effort to secure antheridial, tetrasporic, and cysto-
carpic plants. The material thus obtained confirms the
conclusion that /V. spectadz/e is entitled to rank as a distinct
species, as established by Professor D. C. Eaton.

Hlabitat.—On rocks ? or other algz, sublittoral to elittoral
zone. Dredged in 12-15 fathoms, Monterey Bay, Calif.

Distribution.—Along the coast from Santa Monica north-
ward to Santa Cruz. |

Localities. —Santa Monica! (Miss S. P. Monks); Paci-
fic Grove, in Monterey Bay! (C. P. Nott); Santa Cruz!
(Dr. C. L. Anderson, fde Eaton in Farlow, Proc. Amer.
Acad. Arts and Sci.,-Vol. XII, 1877, p. 245; Mrs. Joe
Weeks).

Nitophyllum fryeanum /arlow.

Algze Amer.-Bor. Exsiccatz, No. 69, 1878.

Nitophyllum fryeanum Harvey, W. H., Ner. Bor.-Amer., Supp., 1858, p.
128? (See remarks on species). FARLOw, W. G., Proc. Amer. Acad.
Arts and Sci., Vol. X, 1875, p. 365; Report U. S. Fish Comm. for 1875,
p. 695, 1876. FarLow, W. G., ANDERSON, C. L., and Eaton, D. C.,
Algz Amer.-Bor. Exsiccatz, No. 69, 1878. HERVEY, A. B., Sea Mosses,
1881, p. 176. ANDERSON, C. L., Zoe, Vol. II, 1891, p..224. Howe,
M. A., Erythea, Vol. I, 1893, p. 68. McCratrcuie, A. J., Proc. So. Cal.
Acad. Sci., Vol. I, 1897, p. 358. AGARDH, J. G., Contin. Spec. Gen. et
Ord. Alg., Vol. III, Pt. 3, 1898, p. 74.

Synopsis.—Frond both prostrate and erect. Prostrate frond much reduced,
flat, membranous, lobed, without venation or rhizoids.

Erect frond sessile or subsessile, flat, membranous, with microscopic veins;
»-~dichotomously or palmately branched or segmented; margin entire,

Bot.—VoL. II.] MOT7—CALIFORNIAN NITOPHYLLA. 23

serrate or toothed. Segments linear, frequently prolonged, occasionally
expanded and lobed at apices. Veins not numerous, extending through the
frond, branching and anastomosing.

Proliferations minute or wanting, appearing along the margin of the frond.

Sporangia in small elliptical sori scattered over the entire frond, tending to
become arranged into lines. Antheridia in whitish areolate patches scattered
over surface. Cystocarps conspicuous, irregularly disposed ovet both sur-
faces, projecting beyond the surface.

ftemarks on the Species.—The color of WV. fryeanum is
very attractive, being a bright rosy red in both the living
and dried states. Some of the plants reach a height of 15
cm. The size and more especially the shape is subject to
considerable variation. The frond may be short and deeply
lobed, or long and branching, with the branches consider-
ably prolonged and linear.

NV. fryeanum is noteworthy as being one of the three
forms of the coast which possess delicate, microscopic
veins. With the exception of this character, JV. Sryeanum
and JV. sfectabile have many points in common. The last
named species, however, is destitute of any kind of
venation.

It is a matter of some doubt whether Harvey’s (1858,
Supp., p. 128) original description of this plant does not
better apply to JV. mu/tlobum. His mention of a lobed and
crenulate margin does not seem to hold good for WV. frye-
anum. Harvey’s name, however, was applied to the speci-
mens issued in the Alg. Amer.-Bor. Exsiccatez (1878, No.
69) and Agardh (1898, p. 74) also retains Harvey’s name,
with an additional reference to the specimens above men-
tioned. It seems advisable, therefore, to retain this name
for the plant here dealt with, which is identical with that
published in the Alg. Amer.-Bor. Exsiccate.

There were reasons for supposing that VV. fryeanum
might be referred to JV. ruthenicum (P. & R.) Kjellman
(1889, p. 25, Pl. I, figs. 11-12), with which it agrees to a
certain extent in the characters of venation and sori. Care-
fully selected specimens were forwarded to Professor Kjell-
man, who replied that the two species were not identical.
The plants exchanged with Kjellman for purposes of

24 CALIFORNIA ACADEMY OF SCIENCES. [PRroc. 3D SER.

comparison, while not wholly inducing the writer to accept
Kjellman’s conclusion as to the non-identity of the two spe-
cies, yet do not furnish sufficient reason for declaring them
identical. Until a more extended comparison can be made
of a wide range of forms, the writer prefers to leave the
species as established in the Alg. Amer.-Bor. Exsiccate
(1878, No. 69). ;

The writer further takes this opportunity to express his
obligations to Mrs. J. M. Weeks for material of 1V. fryeanum
collected by her, which permitted a careful study of anther-
idial, tetrasporic and cystocarpic plants.

Habitat.—No reliable data are at hand concerning the
habitat of this species. Some twenty-five specimens seen
by the writer were all washed ashore from deep water.
The plant presumably occurs upon rocks, and other alge,
in the lower sublittoral, and perhaps elittoral zone.

Distribution.—Known to occur with certainty at but two
localities on the Californian coast. At Golden Gate, San
Francisco Bay, it apparently has not been collected since
the first specimen, if indeed it was identical, was secured
by A. D. Frye and forwarded to Harvey.

Localities.—Pebble Beach, Monterey Peninsula! (Miss
Bayles); Santa Cruz! (Dr. C. L. Anderson; Mrs. J. M.
Weeks); Golden Gate, San Francisco Bay? (A. D. Frye,
fide Harvey, Ner. Bor.-Amer., Supp., 1858, p. 128).

Nitophyllum corallinarum, sp. nov.

Synopsis.—Frond both prostrate and erect. Prostrate frond creeping, flat,
membranous, with microscopic veins and with rhizoids; lobed and branching,
with branches becoming erect at intervals, margin entire.

Erect frond subsessile, shortly stalked, flat, membranous, with microscopic
veins; Ovate-spatulate to elliptical, two to three times longer than broad; sub-
dichotomously lobed or divided, margin entire. Segments minute, ovate,
oblong or cuneate. Stalk very short, narrowly linear or cylindrical, passing
into a midrib, the latter extending throughout frond, usually branching and
free.

Sporangia large, prominent, in sori of varying shape and size, solitary or
clustered on body of frond or its segments.

Femarks on the Spectes.—The form from which the syn-
opsis of this species is drawn up was obtained at San Diego
by Mrs. E. Snyder, and sent to the writer by Mr. F. S.

Bot.—Vou. II.] MOTT—CALIFORNIAN NITOPHYLLA. 25

Collins. It is the only specimen which, to the writer’s
knowledge, has thus far been seen on this coast.

NV. corallinarum isa very minute plant in comparison with
the other Vetophylla of the coast. Corallina chilensts, upon
which the Vetophyllum grows, attains a height between 8
and 12 cm., and in the lower portion a width of 5 cm. or
more. The branches of the Coralline are arranged pin-
nately along the main axis from base to apex of the plant.
The general outline is very regularly fan-shaped. The epi-
phytic W7tophyllum extends, by means of its prostrate frond,
over the entire surface of the Coralline, to which it adheres
firmly by means of the rhizoidal processes that are produced
abundantly from the surface in contact with the Coralline.
From the prostrate frond rise at regular intervals erect
branches which are shortly stalked and expand into ovate-
spatulate or elliptical fronds, which may reach a height of 7
mm. and a width of 3 mm. The color is a rosy red to dull
carmine. So completely is the Coralline enveloped by the
Nitophyllum, that the natural color of the former, as well as
its jointed structure, is very much obscured. The.thin and
delicate frond of JV. corallinarum presents the tessellated
surface characteristic of the genus. Throughout both the
prostrate and erect fronds may be distinguished minute
microscopic veins, which branch more or less freely and
remain free.

The sporangia are prominent and large, considering the
general minute size of the plant. They occur in clusters
rather than in sori of definite shape, and are borne upon the
body of the erect frond or on its segments. Tetraspores
of unusually large size are formed in the sporangia.

This plant may ultimately be found to be identical with
some species of Europe. Such a determination must be
left, however, to future observation upon more abundant
material. .

flabitat.—Epiphytic on Corallina chilensis.

Distribution.—Concerning the distribution of /V. coralli-
narum little can be said. The Coralline upon which it
grows is found along the coast from San Diego northward

26 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

to Fort Ross, but no observations are known to the writer
on the occurrence of JV. corallinarum at any other place
than the type locality.

Locality.x—San Diego! (Mrs. E. Snyder in herb. F. 8S.
Collins).

Nitophyllum uncinatum ¥. Ag.

Spec. Gen. et Ord. Alg., Vol. II, Pt. 2, 1852, p. 654.

Nitophyllum uncinatum MCCLaATcHIE, A. J., Proc. So. Cal. Acad. Sci., Vol.
I, 1897, p. 358; also in Phyk. Bor.-Amer. Co .tuins, F.S., Holden, I.,
and SETCHELL, W. A., Fasc. VII, No. 337, 1897. AGARDH, J. G.,
Contin. Spec. Gen. et Ord. Alg., Vol. III, Pt. 3, 1898, p. 65.

Synopsis.—Frond both prostrate and erect. Prostrate frond creeping,
narrow, linear, thin and membranous; frequently weathered, reduced, and
thickened along median portion, with microscopic veins, and with rhizoids;
margin entire, or toothed; branching, branches rising into erect fronds at
intervals.

Erect frond sessile or subsessile, flat and linear, thin and delicate, some-
times thickened in median portion, with microscopic veins; branching sub-
dichotomously from the base upwards, with margin entire, or occasionally
toothed. Branches linear, occasionally expanded, or acuminate, frequently
recurved or hooked at apices. Veins microscopic, extending throughout
frond, occasionally branching and anastomosing.

‘* Sporangia in solitary disc-like sori, on the upper branches ”’ or ‘‘in round-
ed sori’’ on the outer branches. (See Remarks on Species.) Antheridia as
yet unobserved. Cystocarps minute, marginal or submarginal, produced at
infrequent intervals, projecting slightly beyond surface.!

Remarks on the Species.—lV. uncinatum has a bright,
rosy red tint when alive, usually changing to a dull purplish
or brownish red when dried. The fronds may attain a
length of 10-15 cm. The plant is one of the more delicate
species of the coast, as may be seen in the thin and mem-
branous character of the frond. Throughout the narrow,
linear segments extend microscopic veins, which, with the
numerous recurved or hooked apices of the branches, may
be regarded as the prominent morphological characters.’

1 Account of cystocarps from specimen in Hauck und Richter, Phykotheca Univer-
salis. Fasc. VII, No. 306, 1889.

2 Nordhausen (Pringsheim’s Jahrbiicher f. Wiss. Botanik, Band XXXIV, Heft 2, 1899
p- 263) finds that the hooked apices of the branches of N. unctuatum serve as climbing
organs.

Bot.—VOL. II.]}] MOTT—CALIFORNIAN NITOPHYVLLA. 27

No fruiting specimens of JV. wuctnatum, so far as can be
ascertained, have been reported from Californian shores.
The plant seems to be an exclusively southern form on this
coast, having been collected only at San Diego and San
Pedro, in southern California. From these two localities
numerous and abundant collections have been taken, none
of which, however, have revealed fruiting specimens. There
is a strong probability that the plant propagates itself largely,
if not entirely, by vegetative means. It occurs commonly
on Phyllospadix, in quiet water, conditions of substratum
which would favor the active development of the prostrate
frond. Its local abundance is shown by its occurrence in
such quantities as sometimes to clog the nets of fishermen.

The description of the sori, as given in the synopsis of
the species, is taken from Agardh (1852, p. 654, 1876, p.
465, 1898, p. 65). The account of the cystocarp is based
upon an examination of the specimen issued in the Phyko-
theca Universalis (see note under synopsis of species).
There seems to be little doubt that the species of this coast
is identical with the European plant.

flabitat.—In quiet water, on other alge, and on
Phyllospadix.

Nisiribution.—V. uncinatum is a cosmopolitan species,
limited in its local distribution, having been found at but
two points on the coast.

Localities.—San Diego! (Herb., F. S. Collins); San
Pedro! (Mrs. E. A. Lawrence; A. J. McClatchie; Mrs.
S. C. Purdy; W. A. Setchell).

Nitophyllum multilobum 7. Ag.

Epicrisis Floridearum, Contin. Spec. Gen. et Ord. Alg., 1876, p. 698.

Nitophyllum multilobum AGARDH, J. G., Epicrisis Floridearum, Contin.
Spec. Gen. et Ord. Alg., 1876, p. 698. Fartow, W. G., Proc. Amer.
Acad. Arts and Sci., Vol. XII, 1877, p. 238; ANDERSON, C. L., Zoe,
Vol. II, 1891, p. 224. Coxruins, F. S., HoLpEn, I., and SETCHELL, W.
A., Phyc. Bor.-Amer., Fasc. VII, No. 336, 1897. AGARDH, J. G., Con-
tin. Spec. Gen. et Ord. Alg., Vol. III, Pt. 3, 1898, p. 45.

28 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

Synopsis.—Frond both prostrate and erect. Prostrate frond creeping,
irregularly expanded and membranous, sometimes a rounded expansion,
sometimes irregularly branched or lobed; with rhizoids; without midrib,
nerves or veins; margin entire, sinuous, or somewhat lobed.

Erect frond slightly stalked, flat and linear, with distinct midrib; simple
below, subdichotomously or subpalmately segmented, not proliferating; mar-
gin sinuous, toothed or incised. Segments decidedly bullose, usually obtusely
lobed, with sinuous or toothed margin. Stalk short, linear, flat, occasionally
becoming thickened. Midrib well developed, rather wide, frequently
branched, soon evanescent. .

Sporangia formed in large, irregularly oblong, frequently lobed or confluent
sori, transversely placed upon the segments of the frond. Antheridia as yet
unobserved. Cystocarps few, large, conspicuous, scattered over both
surfaces.

Remarks on the Species.—In the fresh state V. mudtz-
lobum is a dark red to dull carmine, becoming a burnt car-
mine to blackish red when dried. The plant is a dwarf
one, rarely reaching a height of 9 cm.

The predominant characters of /V. mu/t:lobum are seen
in the rather prominent development of the midrib, in the
bullose aspect of the frond, and in the peculiar transverse
sori. The first named.structure is confined to the lower
portion of the frond, where it is visible as a definite thick-
ening of the median part, though it does not project promi-
nently above the surface. At its upper extremity it fre-
quently branches, and the resulting portions evanesce very
soon into the-ordinary tissue of the frond. The bullose
frond of LV. multilobum is an important feature in the,
appearance of the tetrasporic plant. Usually the surface of
the segments which form the upper portion of the plant
exhibits this trait. Here the surface is alternately raised and
depressed, while the margin becomes crinkled and lobed.
The sori, together with the bullose aspect, furnish the most
certain means of identifying the species. No other plant
of the coast possesses such a characteristic feature as these
transverse sori, usually produced in great abundance on the
segments of the frond.

The characteristic transverse sori serve to distinguish JV.
multilobum from LV. harveyanum, with which this plant other-
wise has several points in common. In JV. harveyanum,
however, the sori form flabellate lines on the segments of

Bot.—Vor. I1.] MOZTT—CALIFORNIAN NITOPHYLLA, 29

the frond. In color, the two species are much the same.
The midrib of V. harveyanum is usually more pronounced
than that of JV. multilobum. . harveyanum is much slen-
derer than JV. mu/tzlobum, and may attain a height three or
four times that of the latter.

flabitat.—On bare rock surfaces or on rocks coated with
Corallines, from high water mark to the sublittoral zone.

Distribution.—Limited at the present time to the Cali-
fornian coast. Has now been reported from Carmel Bay
northward to Cape Mendocino. Apparently a northern
form.

Localities.—Carmel Bay! (C. P. Nott); Pacific Grove!
(CC, P. Nott); SantacCruz!: (Mrs: J. M. Weeks); uand’s
End, San Francisco! (W. A. Setchell; C. P. Nott); Golden
Gate, San Francisco Bay (Berggren, fide J. Agardh, Epi-
crisis Floridearum, 1876, p. 698; W. A. Setchell); Lime
Point, San Francisco Bay! (C. P. Nott); Dillon’s Beach
(W. A. Setchell); Fort Ross! (C. P. Nott); Cape Men-
docino (C. G. Pringle, in herb., W. G. Farlow, jide W. A.
Setchell).

Nitophyllum harveyanum 7. Ag.

Epicrisis Floridearum, Contin. Spec. Gen. et Ord. Alg., 1876, p. 462.

Nitophyllum harveyanum J. Ac., Phyk. Bor.-Amer. CoL.ins, F. 5.,
Ho pen, I., and SETCHELL, W. A., Fasc. XIV, No. 693, Igo0.

Synopsis.—Frond both prostrate and erect. Prostrate frond creeping,
linear, flat; without rhizoids, destitute of midrib, and not proliferating;
branching, branches becoming erect at intervals; margin entire, serrate, or
somewhat laciniate.

Erect frond stalked, flat, linear; with midrib and flabellate nerves; branch-
ing, rarely proliferating, margin entire, or somewhat laciniate. Branches
palmate or subpalmate, linear or becoming expanded, occasionally lobed or
cleft. Stalk flat, linear, with distinct midrib, becoming thickened and cylin-
drical through wearing away of margin and renewed growth of median por-
tion. Midrib narrow, conspicuous, branching above, becoming divided into
flabellate nerves, the latter conspicuous, branching wager remaining free and
flabellate. Veins minute or wanting.

Sporangia in linear sori extending flabellately from the nerves to the mar-
gin of the frond. Antheridia as yet unobserved. Cystocarps large, promi-
nent, irregularly disposed, projecting beyond the surface.

30 CALIFORNIA ACADEMY OF SCIENCES. [PRroc. 3D SER.

Remarks on the Species.—N. harveyanum varies in color
in the living state from deep salmon red to dull carmine,
becoming purplish to blackish red when dried. The plant
may reach a height of 20 cm., but is as a rule 6-10 cm.

The prostrate frond in JV. harveyanum does not develop
so extensively as in other species. It is destitute of midrib
and nerves, and is not specially thickened. JV. harveyanum
is a sparingly branched form as regards its erect frond and
the whole plant is rather stiff and unyielding, even when
freshly taken from the water. The midrib is conspicuous,
tapering slightly towards its upper extremity, and rather
suddenly beccming divided into flabellate nerves. The
margin in the lower portions of the frond and upon the stalk
frequently wears away, the remaining median portion then
becoming thickened and cylindrical. In the branches the
margin occasionally is serrate or laciniate.

LV. harveyanum was first collected on this coast at Land’s
End, San Francisco, by Professor W. A. Setchell. It grew
in company with JV. mz/tilobum, to which, at this locality,
it bears some resemblance, on account of its size and vena-
tion. Professor Setchell, however, upon noting the non-
bullose character of the frond and the flabellate arrangement
of the sori, so different from the transverse sori of /V. muz/-
tilobum, concluded that the plant was a distinct species.
The writer, when examining the plant in connection with
other material secured by him at Fort Ross, was of the
opinion that it must be the plant described by Agardh (1876,
p- 699) under the name JV. flabelligerum, although pre-
viously the conclusion had been reached that Agardh’s JV.
flabelligerum was but a form of JV. ruprechtianum.

Sufficient comment has been made already upon the dis-
tinctions to be drawn between JV. harveyanum and JV. mul-
tilobum. It is desirable, however, to point out here some
of the differences existing between JV. harveyanum and cer-
tain forms of JV. ruprechttanum. There is enough of simi-
larity between certain variations of the latter species and
LV. harveyanum to give reason for the suspicion that the
two are identical. The examination of a good range of

Bot.—Vov. II.]/ MO7TT—CALIFORNIAN NITOPHYLLA. 31

specimens of JV. ruprechtianum soon brings to light the
variation in that species, however, and helps to establish its
non-identity with JV. harveyanum.

Certain forms of /V. ruprechtianum exhibit a pronounced
dark purplish red tinge, both in the fresh and dried states,
and are somewhat stiff and brittle. The segments of the
frond are narrower, and more or less prolonged. Such
forms almost invariably bear flabellately arranged, linear
sori, and on the whole, present the distinctive characters of
LV. harveyanum. Between such an extreme variation as
this and the typical /V. ruprechtianum, however, there may
be found every gradation in color, form of segments, and
position and shape of sori, which are discussed more in
detail under JV. ruprechtianum. The form in question,
however, usually retains enough of the distinctive color and
venation of JV. ruprechiianum to enable it to be recognized.

The writer is further indebted to Professor Setchell for a
comparison made by him between specimens of JV. harvey-
anum in the herbarium of Professor Farlow, and _ plants
from this coast. The specimens in Professor Farlow’s
herbarium are from New Zealand, and are designated as
LV. harveyanum by Agardh. The resemblance in habit
between these and plants collected by the writer at Fort
Ross is very striking, both in the tetrasporic and cystocarpic
plants. Judging from Agardh’s description and from this
comparison of specimens, there seems to be good reason for
keeping this species under VV. harveyanum.

Habitat.—NV. harveyanum is found most frequently upon
very much exposed rock surfaces which are bare or coated
with Corallines, at extreme low tide-mark in the littoral and
sublittoral zones.

Distribution.—Along the coast from Santa Cruz north-
ward to Puget Sound. Apparently a northern form.

Localities.—Santa Cruz! (Dr. C. L. Anderson); San
Francisco! (G. W. Lichtenthaler) ; Land’s End, San Fran-
cisco! (W. A. Setchell; C. P. Nott); Duxbury Reef! (W.
A. Setchell); Fort Ross! (C. P. Nott); Puget Sound!
(Thomas Stratton).

32 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

Nitophyllum andersonianum 7. Ag.

Epicrisis Floridearum, Contin. Spec. Gen. et Ord. Alg., 1876, p. 474.!

Nitophyllum (Neuroglossum) andersonit FARLOow, W. G., Proc. Amer.
Acad. Arts and Sci., Vol. X, 1875, p. 365; Report U. S. Fish Comm. for
1875, p. 696, 1876. ,

Neuroglossum andersonianum AGARDH, J. G., Epicrisis Floridearum, Con-
tin. Spec. Gen. et Ord. Alg., 1876, p. 474.

Nitophyllum andersonti Hervey, A. B., Sea Mosses, 1881, p. 177.

Nitophyllum (Neuroglossum) andersonii ANDERSON, C. L., Zoe, Vol. II
1891, p. 224.

Nitophyllum andersonit Howe, M. A., Erythea, Vol. I, 1893, p. 68. Mc-
CLaTcHIE, A. J., Proc. So. Cal. Acad. Sci., Vol. I, 1897, p. 358.

Neuroglossum andersonianum AGARDH, J. G., Contin. Spec. Gen. et Ord.
Alg., Vol. III, Pt. 3, 1898, p. 122.

Synopsis.—Frond both prostrate and erect. Prostrate frond creeping,
slender, linear, without rhizoids; branching irregularly, occasionally prolifer-
ating; without midrib, nerves, or veins. Margin beset at regular intervals
with spine-like, sometimes recurved, pinnate teeth. Branches becoming
erect at intervals, expanding into erect fronds.

Erect frond shortly stalked, linear, flat, simple below, branching above,
with midrib; margin serrate, dentate, or beset with numerous spine-like,
pinnate teeth. Branches subpinnately arranged, linear, or alternate at base
and expanding at their apices, usually much prolonged. Stalk linear, flat,
with definite midrib and thin margin, the margin sometimes disappearing and
the median portion becoming thickened, almost cylindrical. Midrib of vary-
ing width, becoming prominent, in some cases thickened, almost cylindrical,
branching and evanescent in upper portions of frond.

Sporangia in rounded sori, the latter usually large and conspicuous, at the
apices of the upper, sometimes expanded, branches. Antheridia and cysto-
carps so far unobserved.

Pemarks on the Species.—The color of /V. andersont-
anum varies from bright red to dull carmine when alive,
becoming a burnt carmine in the dried specimens. More
often the plant has the darker hue mentioned above. The
frond may attain a height of 20 cm.

The prostrate frond is commonly slender and much
branched, showing much similarity to the corresponding
portion of /V. Jatéssimum. It is destitute of midrib and
nerves, and seldom becomes thickened or broadly linear.
The erect frond branches freely, while its divisions exhibit
considerable variation in width. In some plants they are
very slender, linear, and much divided or branched. In

1 This plant was here for the first time described. It had, in 1875, been mentioned by
Farlow (cf. citations) under the name Nitophylinm (Neuroglossum andersonit) J. Ag. Ws,

Bot.—Vot. II.] MO7T—CALIFORNIAN NITOPHYLLA. 23

others the segments are quite broad and very regularly pin-
nately arranged. The cause of this variation apparently
may be found in the environment. When exposed to vio-
lent wave action the fronds become extensively branched.
In comparatively quiet waters, on the other hand, the ex-
panded frond reaches its widest development. The pre-
dominant characteristic of V. andersonianum is the produc-
tion, along the margin of both the prostrate and erect frond,
of numerous pinnately arranged, spine-like, minute projec-
tions or teeth. The midrib, in its normal state, is the
slightly thickened median portion of the frond, due to an
increase in size of the cells of the central layer. Unless
the frond is stimulated to further growth by injury, the
midrib remains in this state, and, on reaching the upper
branches of the frond, soon evanesces. Under the process
of weathering, however, the margin becomes worn away.
This seems to incite the cells of the median portion to
renewed growth, with the result that the stalk and definite
midrib become thickened and almost cylindrical. This spe-
cies does not commonly proliferate.

The tetrasporangia are as yet the only reproductive
structures observed. The antheridia and cystocarps have
not, so far as can be learned, been seen in the species.

It seems to the writer desirable to allow this species to
remain under Vtophyllum rather than to assign it to Veuro-
glossum. The habit, sori, and inner structure furnish evi-
dence for thus placing it. Since it was first reported from
this coast, doubt seems to have existed in the minds of writ-
ers as to whether it was a (Vétophyllum or a Neuroglossum.
The synopses already in existence of the genus Neuroglos-
sum and of the present species do not at all permit a defi-
nite idea to be gained of the internal structure, and widely
varying statements exist as to the position of the sori. But
the careful study of an extensive range of specimens has
served to establish the writer’s belief that for the reasons
above mentioned the species can be referred to Witophyllum
until a comparison can be made with type specimens.

(3) July 28, 1900,

34 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

Habitat.—On other alge and on rocks covered with Porif-
era and Bryozoa at low tide-mark in the littoral and sub-
littoral regions.

Distribution.—lV. andersonianum has now been observed
along the coast from San Pedro northward to Carmel Bay.
It has never been recorded elsewhere than from the coast of
California.

Localities.—San Pedro! (Mrs. E. A. Lawrence); Santa
Barbaral (Dr. L. N: -Dimimick; Mrs. S. P; (Coeoperje
shores of San Luis Obispo County! (Mrs. R. W. Sum-
mers); Carmel Bay! (C. P. Nott); Pacific Grove! (Mrs.
J. M. Weeks; M. A. Howe); Santa Cruz! (Dr. C. L.
Anderson, Mrs. Boardman).

Nitophyllum ruprechtianum 7. Ag.

Bidrag till Florideernes Systematik. Lunds Universitets Arsskrift., Tome
VIII, 1871, p. 51.

Nitophyllum ruprechtianum Far.tow, W. G., Proc. Amer. Acad. Arts and
Sci., Vol. X, 1875, p. 365; Report U. S. Fish Comm. for 1875, p. 696,
1876. AGARDH, J. G., Epicrisis Floridearum, Contin. Spec. Gen. et Ord.
Alg., 1876, p. 470.

Nitophyllum flabelligerum AGARDH, J. G., loc. cit., p. 699.

Nitophyllum ruprechtianum Hervey, A. B., Sea Mosses, 1881, p. 178.
ANDERSON, C. L., Zoe, Vol. II, 1891, p. 223. Howsz, M. A., Ery-
thea, Vol. I, 1893, p. 68. McCtatcnis, A. J., Proc. So. Cal. Acad.
Sci., Vol. I, 1897, p. 358. TILDEN, J. E., American Algz, Century III,
No. 213, 1898. AGARDH, J. G., Contin. Spec. Gen. et Ord. Alg., Vol.
Pi bt 35. 1895, p94:

Nitophyllum marginatum AGARDH, J. G., loc. cit., p. 93.

Nitophyllum farlowianum AGARDH, J. G., loc. cit., p. 95.

Synopsis.—Frond both prostrate and erect. Prostrate frond creeping, flat,
narrowly membranous, or linear; with rhizoids; frequently proliferating,
lobed or branching; often provided with midrib and nerves; margin entire,
cuneate, or lobed; frequently forming offshoots or innovations.

Erect frond stalked, with midrib, nerves, and veins; branching, very often
proliferating; margin entire, cuneate, undulate, crispulate, laciniate or lobed,
often beset with minute proliferations. Branching subdichotomous or sub-
palmate, with branches linear and often much prolonged, occasionally alter-
nate below, becoming expanded and cuneate above, frequently divided or
lobed. Stalk linear, flat, with definite midrib, very often becoming cylin-
drical through wearing away of margin and thickening of midrib, frequently
twisted by wave action, often persistent and freely proliferating. Midrib

Bot.—Vo.. II.] MO7T—CALIFORNIAN NITOPHYVLLA. 35

conspicuous, branching, sometimes divided into usually conspicuous flabel-
late nerves and veins, or unfrequently remaining undivided and evanescent;
frequently weathered, thickened, persistent, and proliferating freely from
sides and end. Nerves flabellate, free or anastomosing, often conspicuous,
or inconspicuous and evanescent, sometimes dividing into minute veins.
Proliferations produced very freely, on stalk, on margin of frond or on
reduced frond, minute and rounded or large, linear, cuneate, frequently lobed
or divided, with midrib and flabellate nerves, often bearing sori and cystocarps.

Sporangia in linear sori flabellately disposed about margin of frond, or in
linear or irregular submarginal patches, or upon proliferations abundantly
produced along margin of frond or upon its surface. Antheridia as yet
unobserved. Cystocarps large, infrequent, projecting, irregularly disposed
over both surfaces, or gathered together along the margin, or borne upon
marginal or surface proliferations.

Ltemarks on the Species. —Considerable variation in color
may be observed in JV. ruprechtianum. The plants, when
young, are often bright red, becoming dull red or carmine
with increasing age. The proliferating fronds often exhibit
this change in color. When dried, the plant becomes a
deep carmine to blackish red. A length of 20-30 cm. is
not uncommon, as the plant is vigorous in its growth. Fre-
quently a large number of abundantly branched fronds
develop from a single stalk. Good specimens often form
masses 30 cm. in diameter and 30 cm. or more in height.

The prostrate frond in JV. ruprechtianum is developed
rather more extensively than in any other species of the
coast. It is usually membranous in character, and may
either be undifferentiated to any extent or occasionally be
provided with midrib and nerves. Owing to the free devel-
opment and frequent branching of the prostrate frond, there
is formed on the substratum an extensive ramification whose
outer ends, by the decay or accidental rupture of the older
portions of the frond, become separated, and constitute the
starting point for a new frond.

The predominant characters of /V. ruprechtianum are
displayed in the robust habit, the abundant proliferations,
and the variety in the position of the cystocarps and sori,
and the shape of the latter.

Proliferation takes place more abundantly in JV. ruprecht-
zanum than in any other of the Californian forms. The
frond very commonly becomes worn away by the action of

36 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

the waves, and by friction upon other alge and upon rocks.
The portions thus reduced persist for a considerable time
and give rise to numerous proliferations which exhibit all
the characters of the original frond.

It seems strongly probable that the remarkable degree of
variation in the shape and position of the sori in JV.
ruprechtianum has been a fruitful source of error to those
who have been called upon to identify the forms belonging
to this species. It is difficult, perhaps, without extensive
examination of plants on the shore at all seasons, to realize
what a diversity of form may be found within the specific
limits. Especially is this diversity important in considering
the sori, which are usually much employed in establishing
specific distinctions.

The examination of abundant material of JV. ruprecht-
zanum has shown that the plants fall into three groups dis-
tinguished by the differences in the method of production
of the sori, without regard to other features. In one group
the sori are produced on the upper, flabellately expanded
segments of the frond, and are arranged in rather wide
linear sori, or lines, distinct or occasionally confluent, and ex-
tending in a more or less connected fashion from the median
portion of the segment to the margin. A second group
exhibits these linear sori usually confined, however, to the
apices of the segments, while along the margin are pro-
duced numerous minute proliferations upon which are borne
sori having the form of rounded patches. In a third group,
the sori are confined to the marginal proliferations or to the
similar proliferations appearing upon both the margin and
the surface. While, in general, these variations in the posi-
tion and shape of the sori are seen on different plants, yet
it is of great importance in employing them as specific char-
acters to keep in mind the fact that these three different
dispositions of the sori are likewise found on one and the
same plant.

A similar habit with respect to the production of sori is
seen in JV. violaceum, where, however, the linear sori are
much narrower and the marginal ones are often widely

a

tak DEVELOPMENT OF THE KARYOKINETIC
SPINDLE IN THE POLLEN-MOTHER-CELLS
OF LAVATERA.!

BY EDITH SUMNER BYXBEE.

PLATES X—XIII.

THE way in which the spindle is formed varies widely in
the different families of plants that have been studied. In
the generative cells of higher plants, the spindle seems to
be always multipolar at first, but the formation of the poles
may proceed in several different ways.. With a view to
shedding further light on this question, the pollen-mother-
cells of a species of Lavatera were selected for study.’

This plant blooms throughout the year, so that the mate-
rial is plentiful at all times, while the arrangement of the
flowers in dense racemes and the number of anthers in a
flower make it easy to obtain cells in all stages of division.
The pollen-mother-cells are large. They may be examined
before preserving, so that much timeis saved. The flowers
were usually gathered in the morning, as it was found that
then the cells were dividing more rapidly. The anthers
were examined by crushing them, either with or without the
addition of one per cent. glacial acetic acid. All heads in
a favorable condition were then immediately dropped into
the fixing fluid. It may be mentioned here that all the more
striking appearances observed in preserved material were
also seenin the fresh. ‘The granular zone, especially, stands
out with great distinctness by reason of the strong refrac-
tive properties of its granules. %

1 Contributions from the Botanical Laboratories of the University of California, No.
13. Presented for the degree of Master of Science, Prepared under the direction of Dr.
W. Jj. V. Osterhout.

2 Owing to the destruction of the plants, it has been impossible to determine the spe-
cies with exactness. Itis probably either Lavatera uaguiculata Desf. or L. micans \,.

[63] October 29, rg00.

64 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

The following fixing fluids were tried: Flemming’s
strong mixture; one per cent. chromic acid; two per cent.
iron trichloride; Wilson’s corrosive sublimate-acetic; Bove-
ri’s picro-acetic: Lindsay’s potassium bichromate-platinum
chloride-osmic-acetic; one per cent. palladium chloride and
one-half per cent. iridium chloride.

Flemming’s strong mixture was used undiluted and also
with the addition of one, two and three parts of water. The
quantities of chromic and acetic acids in the original mixture
were also varied. Dilute solutions were found to shrink the
cells much more than the strong ones. Varying the amount
of chromic acid did not improve the action of the fixing .
fluid. An increase in the amount of acetic acid, however,
gave the best results obtained. Flemming’s strong mixture
with an excess of acetic acid was therefore almost exclu-
sively used. Fair results were also obtained with palladium
chloride and iridium chloride to which a small amount of
glacial acetic acid had been added.

After remaining in the fixing fluid for twenty-four hours,
the anthers were washed in running water for six hours.
They were then placed in a dehydrator! for twenty-four
hours, with 95 per cent. alcohol below and distilled water
above.

Some alcohol was then removed from the material and
mixed with an equal volume of 95 per cent. alcohol. The
material was transferred to this stronger mixture for two
hours. By repeating this process three or four times the
material was brought into 95 per cent. alcohol without
shrinkage. It was left in 95 per cent. alcohol for twenty-
four hours. It was then placed for six hours in each of the
following, successively: absolute alcohol; absolute alco-
hol and bergamot oil (equal parts); bergamotoil; bergamot
oil and paraffin, 47° (equal parts); paraffin 47°; paraffin
47° and paraffin 54° (equal parts); paraffin 54°.

Sections 3 to 4 u in thickness were cut with the Minot
wheel microtome. Of the stains tried, Flemming’s triple
stain (safranin, gentian violet and orange ‘‘G’’) gave the
best results.

1 For a description of the dehydrator see Lawson, 1898, and Williams, 1899.

Bot.—VOL. II.] BYXBEE—LAVATERA. . 65

The pollen-mother-cells are large, and the-diameter of the
nucleus is equal to fully one-half that of the cell. The
chromatin thread is thick and stains blue with gentian
violet. The large nucleolus stains red with safranin. It
containsa single vacuole. The linin either forms a complete
network filling the whole nucleus, or is present as broken
threads attached to the chromatin and nucleolus. In neither
case does it stain.

The cytoplasm is made up of two constituents, one fibrous,
the other granular. The fibrous part forms a network com-
posed of delicate threads crossing each other in every
direction. These threads stain deep blue with gentian
violet. The other element is composed of small granules,
varying somewhat in size.

These granules are scattered throughout the network,
both between and upon the fibers, usually in sufficient quan-
tity to give the cytoplasm a cloudy appearance. They have
a tendency to collect in small, denser masses at the inter-
section of the fibers of the network. This tendency is
especially noticeable in the earlier stages. A little later the
granular matter is distributed more evenly. It stains a
brownish yellow with orange ‘‘G’’. A cell in this stage is
shown in fig. I.

The first change that occurs in the cytoplasm is the elon-
gation in a direction parallel to the nuclear wall of the row
of meshes immediately surrounding the nucleus (fig. 2).
This process goes on until three or four rows of meshes
outside the nucleus have become pulled out in this way (fig.
3). These meshes become so long and narrow that, often,
on casual inspection, there seem to be threads wound round
and round the nucleus. On close examination, however,
the meshes can always be seen. Within the nucleus the
linin network has meanwhile broken up somewhat; the
threads lose their smooth, transparent appearance and begin
to stain blue. One or more additional vacuoles appears in
the nucleolus, which begins to stain purple rather than red.

Soon after this, the granular substance, which hitherto has
been equally distributed throughout the cytoplasm, begins
to collect in a denser mass immediately about the nuclear

66 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

wall, covering the elongated meshes of the cytoplasmic
reticulum (fig. 4). It increases rapidly in quantity about
the nucleus without decreasing throughout the rest of the
cytoplasm. It is always densest close to the nucleus and
from there shades gradually out into the cloudy mass of the
cytoplasm.

As this granular substance begins to accumulate, the
meshes of the cytoplasmic reticulum, with the exception of
a narrow zone surrounding the nucleus, become radially
elongated (figs. 4 and 5). This arrangement, however,
does not seem to have any particular significance and soon
disappears. Ina very short time a zone of granular matter
has collected about the nucleus, occupying from one-half
to one-third of the space outside of it. It becomes so
dense that it entirely obscures the reticulum within it,
except that a few fibers may sometimes be visible close to
the nuclear wall (figs. 5 and 6). By this time, the elon-
gated meshes of the cytoplasm immediately surrounding
the nucleus have been transformed into free fibers, which
lie between the granular zone and the nuclear wall. Occa-
sionally there is a felt of fibers bounding the outer edge of
the granular zone, and, by reason of their deep blue color,
standing out conspicuously against the yellowish brown
granular matter (figs. 6 and 8). This is by no means
constant.

By the time the granular zone is completely formed, the
radial arrangement of the reticulum outside of it has entirely
disappeared (fig. 6). The granular zone is composed of a
dense mass of granules, most of which are larger than those
that gave the cytoplasm its cloudy appearance in the earli-
est stage. In this form it remains unchanged until the end
of the anaphase.

While the granular zone has been gathering, the linin in
the nucleus has increased somewhat in quantity and in
staining power. About this time, the cytoplasm loses its
regular structure, as shown in figs. 6, 7, etc. Soon the
fibers immediately outside of the nuclear wall come into
clearer view, as though the granular matter had withdrawn

Bot.—Vot. II.] BYXBEE—LAVATERA. 67

from the wall a little, or had been used up at its inner edge
(figs. 7 and 8). These fibers, as has already been stated,
are probably derived from the elongated meshes of the
cytoplasm. They are of much greater diameter than the
linin threads, are smooth and stain deep blue. The extent
to which these fibers are visible varies greatly in different
cells.

At some point the nuclear wall disappears and through
the gap thus formed the fibers immediately without the
nucleus begin to grow into the cavity (fig. 9). At first,
these fibers can be distinguished from the linin threads of
the nucleus by their greater diameter and smoother appear-
ance. As the nuclear wall continues to disappear, how-
ever, and the linin to thicken up, the fibers from within and
without the nucleus mingle in an interwoven mass in which
those of different origin cannot be distinguished (fig. 10).

By the time the nuclear wall has entirely disappeared,
the nuclear cavity is filled with a mass of interwoven fibers
which is usually densest about the circle of chromosomes
which marks the situation of the old nuclear wall (fig. 40).
In some part of the nuclear space the fibers crowd closer
together to form a denser mass and at the same time tend
to range themselves so that they lie more or less parallel to
each other (figs. 11 and 12). Soon a number of projec-
tions appear in this mass as though it were being drawn out
at a number of points. Figure 13 shows an earlier and fig.
14 a later stage in this process. This continues until a dis-
tinct multipolar spindle is formed (fig. 15).

Soon two principal groups of fibrous cones can be distin-
guished in the multipolar spindle and in each of these
groups one cone becomes the most prominent. Into these
two all others are soon absorbed by the continued straight-
ening out and converging of the fibers. Figure 16 shows a
spindle in which the cones have almost reached their final
position and in which there is only a trace of the third cone.
The completion of this process brings the chromosomes
into position at the equatorial plate just at the time that the
perfect bipolar spindle is formed.

68 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

The completed bipolar spindle lies with its poles close
against the inner edge of the granular zone, which has as-
sumed an oval shape to accommodate itself to the outline of
the spindle. The spindle has sharply pointed ends (fig. 17).
It is composed of two sets of fibers. One set runs from pole
to pole forming the central spindle. The other runs only
from the poles to the chromosomes to which the fibers are
attached in bundles. These fibers contract and pull the
daughter chromosomes toward the poles. When this process
is completed the fibers of the central spindle no longer appear
straight but have assumed a wavy appearance. As the
daughter chromosomes approach the poles, the mantle fibers
appear (fig. 18). About this time the granular zone loses
its definite outline. It begins to break up in the plane of the
equatorial plate of the spindle and gathers about the poles
in two masses in which the daughter chromosomes lie em-
bedded. These two masses remain connected by a shell of
granular matter which outlines the old spindle. Figure 19
shows the beginning of this process and fig. 20 a later stage.
The cytoplasm seems to contain a greater quantity of gran-
ular matter than at any previous time.

Within these two granular masses are formed the daughter
nuclei. These have at first a decided indentation on the
side toward the spindle (fig. 20). Later they become
spherical (fig. 21). The daughter nuclei are thus from the
first surrounded by a granular zone which, by the time they
are completely formed, has become relatively as wide and
as dense as that about the mother nucleus. The central
spindle fibers seem to disintegrate and when the nuclei are
ready for the second division remain simply as lines of
granules connecting the two granular zones (fig. 21).

The second division, as far as could be observed, exactly
repeats the process of the first. It was impossible to follow,
under the dense granular zone, the elongating of the meshes
about the walls of the nuclei, but the concentric lines of
fibers were visible in many cases in the narrow space between
the zone and the wall. The breaking down of the nuclear
wall, the growing in of the fibers, and the formation of the
multipolar and bipolar spindles occur as in the first division.

Bot.—Vot. II.] BYXBEE—LAVATERA. 69

The planes of division of the daughter nuclei do not seem
to be at all constant. Sometimes they divide in the same
plane, sometimes in planes at right angles to each other, and
there are all possible transitions between thesetwo. Figure
22 shows a cell in which the planes are at right angles to
each other and fig. 23 one in which they are nearly parallel.

The four daughter nuclei resulting from the second divi-
sion become surrounded by granular zones just as did the
nuclei resulting from the first division. These zones are
usually very broad and dense. Connecting them are the
mantle fibers across which the cell-plates are formed later
on (fig. 24).

The granular zone persists even in the pollen-grain, at
least while itis young (fig. 25). It usually occupies at least
one-third of the cell space outside of the nucleus. The
cytoplasm outside of the zone also contains a great deal of
granular matter.

The most important fact in the method of spindle forma-
tion above described is that the spindle is formed from free
fibers and not from a network. That part of the cytoplas-
mic reticulum which aids in the formation of the spindle is
converted into free fibers at an early stage, long before the
nuclear wall breaks down (figs. 5, 6, etc.). The linin net-
work of the nucleus breaks up at an even earlier time.
The fibers derived from these two sources become inter-
woven but never form a true network.

The granular zone, too, is more than usually prominent
in Lavatera. Its significance will be discussed later.

The higher plants hitherto most exhaustively studied, as
Equisetum (Osterhout, 1897), Lartx (Belajeff, 1894),
Cobea (Lawson, 1898), and Passzifora (Williams, 1899),
all show a certain general resemblance to each other and to
Lavatera in the method of forming the spindle in the repro-
ductive cells. No two of them, however, agree fully in
the details. In all of them the first changes in the cyto-
plasm are either a radial elongation of the meshes of the
reticulum or a parallel drawing out of the first two or three
rows adjacent to the nucleus.

7O CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

In Zarzx the radial precedes the parallel elongation, and
is of short duration, to judge from Belajeff’s figures. In
Equisetum, Passtflora, and Lavatera the parallel elongation
is the first change observed in the cytoplasm; in Cobea
alone it has not yet been observed. In Lartw and Lavatera
this condition persists. In the former case, the elongated ~
meshes become part of the central network; in the latter
they become transformed into free fibers. In Passzflora
and Eguzsetum this condition is transitory. In Aguzsetum
the parallel elongated meshes are drawn out into an indefi-
nite mass of radially arranged fibers which grow out into
the cytoplasm, become parallel to each other, and finally
bend together into groups, so that, before its breaking
down, the nuclear wall is surrounded by a number of cones.
When the nuclear wall disappears, the fibers of these cones
grow into the nuclear cavity, come into contact with the
linin threads and the chromosomes, and form a multipolar
spindle.

In Passiflora the radial elongation of the meshes of the
reticulum persists for some time. Some of the threads
stain more strongly and present an outline suggesting the
cones in Hguisetum, though they are not formed of free
fibers. This condition of the cytoplasm is transitory and
seems to have nothing to do with the formation of the spin-
dle. This is formed directly from the network resulting
from the union of the linin reticulum of the nucleus with
that portion of the cytoplasmic reticulum immediately out-
side of the nucleus. On the breaking down of the nuclear
wall these unite to form a continuous network which fills
the entire space within the granular zone. The network
becomes pulled out at a number of points, and is changed
into free fibers which form the multipolar spindle.

In Larix, Cobea and Lavatera, no cones are present
before the nuclear wall disappears. It would seem that the
development of these cones is correlated with the slight
development of the granular zone. It hardly seems that it
would be possible to have such cones in forms like Cobe@a
and Lavatera, where the granular zone is very dense. In

Bor.—VOL. II.] BYXBEE—LAVATERA. ya

Larix, where there is less granular matter, long fibers
extend out from the central network—which results, as in
Passifiora, from the union of cytoplasmic and linin threads—
to form by their contraction the cones of the multipolar
spindle.

In Cobea and Lavatera the spindle formation goes on
within the dense granular zone. In Codea a network is
formed as in Larzx and Passiflora. In Lavatera the fibers
seem to be always distinct and bunch themselves together
into a dense mass. The multipolar spindle is formed by a
pulling out of the network (Cobe@a), or mass of fibers (Lav-
atera), as in the other cases.

The method of formation of the bipolar from the multi-
polar spindle does not seem to vary in the various cases.

One of the principal differences, then, in the method of
the formation of the spindle in the various plants studied,
seems to be the time at which the free fibers are formed
from the reticulum of the resting cell. In some cases this
occurs very early, as in Hguzsetum and Lavatera, which,
however, differ widely in other respects. In other cases,
as Larix, the spindle itself seems to be a network much
stretched out.

The granular zone, whichis so conspicuous in Codea and
Lavaiera, has been figured in most of the papers on the
division of the generative cells of the higher plants. Oster-
hout (1897) figures it in the bipolar stage in Hguzsetum and
Mottier (1897 a) in Podophyllum and Helleborus. Mottier
(1897 4) also speaks of its presence at several stages in
the divisions in the embryo-sacs of the Lz/zacee, but does
not seem to regard it as constant in or characteristic of these
divisions. Juel (1897) figures it as a prominent ring in
Flemerocallis, but does not discuss it at any length. In
Passiflora, also, it forms a well marked zone.

While it is so commonly, perhaps invariably, present in
the reproductive cells at the time of their division, it has not
been observed in any of the dividing vegetative cells that
have been studied. This fact seems to indicate a connection
with the two rapid divisions of the reproductive cells for

72 CALIFORNIA ACADEMY OF SCIENCES. [PRroc. 3D SER.

which it apparently furnishes food material. The granules,
too, strongly suggest the yolk granules present in animal
eggs. They appear in small quantities in the pollen-mother-
cells at an early period, but their exceedingly rapid increase
at the time when the first steps in the actual formation of
the spindle takes place strongly impresses one with the idea
that the cytoplasm is busy producing nutritive material to
serve the cell through the period of its activity.

The manner in which the granular zone accumulates sug-
gests the gathering of the deutoplasm in animal eggs ( Wil-
son, 1896, p. 115 et seq.). In Passzfora it appears first in
patches scattered through the cytoplasm and later gathers
into a somewhat loose ring at some distance from the
nucleus. This is the way in which the yolk collects in the
egg of Diemyctylus and other Amphibians, as described by
Jordan (Wilson, 1896, p. 116).

In Cobea and Lavatera the granular substance appears
first close around the nucleus and spreads out from this as
in the trout (Henneguy) and cephalopods (Ussow) (See Wil-
son, 1896, p. 117). The exceedingly dense zone formed
by this substance in Lavatera exactly resembles that figured
by Van Bambeke for a fish (.Scorpena) (Wilson, 1896; p.
116, fig. 59, C). In some cases, as Cobea, the granular
matter is used up as the divisions are completed. In Lava-
tera, however, it constantly increases in quantity up to the
formation of the pollen-grains in which it is present, at
least in the younger stages. It seems probable that it per-
sists in them and serves them for food during the develop-
ment of the pollen-tube and the succeeding divisions.

Bot.—Vot. IL.] BYXBEE—LAVATERA. 73
SUMMARY.

1. The cytoplasm of the young pollen-mother-cell is
made up of two constituents—a fibrous network and a
granular substance.

2. The spindle is formed in the following manner :—

a. ‘The meshes of the network, close to the nuclear
wall, pull out in a direction parallel to the wall,
forming a felt of fibers about the nucleus.

6. The granular constituent of the cytoplasm col-
lects in a wide, dense zone about the nucleus.

c. The linin increases in quantity.

d. The nuclear wall breaks down and the fibers
outside begin to grow into the nuclear cavity.

e. The cytoplasmic and linin fibers form a mass in
which the chromosomes lie.

jf. The mass of fibers projects out at a number of
points, forming the multipolar spindle.

g. Two of the cones become more prominent than
the others, which they finally absorb, thereby
forming the bipolar spindle.

3. The process in the second division exactly repeats
that in the first.

4. The granular substance, forming the dense zone, is
comparable with the deutoplasm of animal eggs.

3. Finally, the spindle is formed directly from elements
—cytoplasmic and linin reticula—present in the cell from
the first, and not from any special spindle-forming sub-
stance, or by the aid of centrosomes.

BOTANICAL LABORATORY,

UNIVERSITY OF CALIFORNIA,
October, 1899.

BotT.—VOL. II.] BYXBEE—LAVATERA. 75

1894.

1897.

1898.

1897a.

18976.
1897.

1897.

1896.
1899.

BIBLIOGRAPHY.

BeLajerF, W. Zur Kenntniss der Karyokinese bei den Pflanzen.
Flora, Bd. LXXIX, p. 430.

JueL, H. O. Die Kerntheilung in den Pollenmutterzellen von Heme-
rocallis fulva, und die bei denselben auftretenden Unregelmassig-
keiten. Pringsh. Jahrb., Bd. XXX, p. 205.

Lawson, A. A. Some Observations on the Development of the
Karyokinetic Spindle in the Pollen-Mother-Cells of Cobzea scan-
dens Cav. Proc. Cal. Acad. Sci., 3d Ser., Bot., Vol. I, p. 169.

Mortirr, D. M. Beitrage zur Kenntniss der Kerntheilung in den
Pollenmutterzellen einiger Dikotylen und Monokotylen. Pringsh.
Jahrb., Bd. XXX, p. 169.

Ueber das Verhalten der Kerne bei der Entwickelung des
Embryo-sacks, etc. Pringsh. Jahrb., Bd. XXXI, p. 125.

OstEerHoutT, W. J. V. Ueber Entstehung der karyokinetischen Spin-
del bei Equisetum. Pringsh. Jahrb., Bd. XXX, p. 155.

STRASBURGER, E. Ueber Cytoplasmastructuren, Kern- und Zelltheil-
ung. Pringsh. Jahrb., Bd. XXX, p. 375.

Witson, E. B. The Cell. New York. 1896.

Witurams, C. L. The Origin of the Karyokinetic Spindle in Passi-
flora coerulea Linn. Proc. Cal. Acad, Sct., 3d Ser., Bot., Vol. I,
p. 189.

76 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

EXPLANATION OF PLATE X.

All figures were drawn with the Abbe camera lucida: objective, Zeiss oil
immersion 1/12, compensating ocular No. 6. ;

Fig. 1. A young pollen-mother-cell. The chromatin thread is beginning
to break up. The cytoplasm is composed of two elements, one
a fibrous network, the other a granular substance.

Fig. 2. A little laterstage. The first row of meshes adjacent to the nuclear
wall has begun to pull out.

Fig. 3. A number of rows of meshes adjacent to the nuclear wall have be-
come elongated. The chromatin thread has broken up.

Fig. 4. The granular substance has begun to collect about the nuclear wall.
The outer meshes of the cytoplasm have become radially elon-
gated. The nucleolus shows four vacuoles.

Fig. 5. The granular zone has increased in width. Fibers can be seen
between it and the nuclear wall. The linin is thickening up.

Fig. 6. The granular zone is completely formed. The radial arrangement
of the meshes outside of it has disappeared.

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CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

EXPLANATION OF PLATE XI.

The space between the granular zone and the nuclear wall has
widened and in this space numerous fibers are seen. The cyto-
plasm has a less regular structure.

About the same stage as that shown in fig. 7, but there are many
more fibers outside the nucleus. There are also strongly staining
fibers in the cytoplasm.

The nuclear wall has disappeared at one point and the fibers out-
side the nucleus are growing into its cavity. The linin has
increased in quantity.

The nuclear wall has completely disappeared and the space within
the granular zone is filled with a mass of interwoven fibers; those
near the center of the space are linin.

The fibers begin to straighten out and arrange themselves in groups,
in which they are parallel to each other.

A more advanced stage in this process.

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CALIFORNIA ACADEMY OF SCIENCES. [PRroc. 3D SER.

EXPLANATION OF PLATE XII.

The fibers begin to converge at a number of points to form cones.

The cones project out farther from the mass of fibers.

The fibers have straightened out. The cones are in two groups.

The cones have nearly fused to form the bipolar spindle.

A completed bipolar spindle.

The daughter chromosomes have nearly reached the poles. There
are a few mantle fibers. The fibers of the spindle are no longer
straight.

The granular matter is becoming thinner at the equator and collect-
ing more densely about the poles of the spindle. There are
traces of radiations which seem to be formed of granular matter
extending out from the poles.

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CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

EXPLANATION OF PLATE XIII.

The daughter nuclei are formed. They are at this time crescent
shaped as seen in section. There is still a shell of granular mat-
ter bounding the remains of the spindle. It is seen in section as
two lines.

The daughter nuclei have become spherical. The granular matter
has increased in quantity.

A spindle of the second division. The other spindle, whose posi-
tion is indicated by the circular mass of granular matter, lies at
right angles to the first.

Two spindles of the second division that lie nearly parallel to each
other. The granular mass between them is not of equal density
throughout.

The four Giughter nuclei, each surrounded by a dense granular
zone. They are connected by mantle fibers.

A young pollen-grain. The nucleus is surrounded bya dense gran-
ular zone and the cytoplasm outside of this contains much gran-
ular matter.

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[BYXBEE] PLATE XIII.

Proc.CALACAD. Sc1.32 SER Bor. VoL IL

STUDIES ON THE COAST REDWOOD,
SEQUOIA SEMPERVIRENS ENDL.

BY GEORGE JAMES PEIRCE.
Associate Professor of Plant Physiology in the Leland Stanford Juntor University.

CONTENTS.
PEATE OXCRVE
PAGE.
. THE VEGETATIVE MODE OF REPRODUCTION IN THE REDWOOD... 83
I]. PECULIARITIES OF SOME VEGETATIVELY PRODUCED YouNG
RE DWOO DS pepe e err ole are sae rete Eo eS es Sicicmuem aloe 88
Alte ELAS GTADILO NM oaie erste tty aeteermere eter aes PEN cisicicle cocte aoe cicsskeoweh cre 88
(BAe OU AIEBINIS Mtn cieerin aoa cetcieomtie Sent sina sl ara crear ania Ass wsrdewe cs 89
Ill. THe SIGNIFICANCE OF THE WHITE REDWOODS IN CONNECTION
WITH OUR CONCEPTIONS OF PARASITISM AND OF HEREDITY..... 100

I. THe VEGETATIVE MopE oF REPRODUCTION IN THE
REDWOOD.

THE coast redwood (Seguoza sempervirens Endl.) is one
of the comparatively few coniferous plants which reproduce
themselves vegetatively by suckers. The majority of the
Coniferz multiply only sexually, by seeds. Seeds offer the
means by which plants may be carried in the dormant or
resting condition for long distances, thus permitting ex-
tended distribution. If the seeds retain their vitality, they
offer also the means by which the species as well as the life
of the individual may be maintained for long periods. A
forest composed of other coniferous trees than the redwood
will usually come to an end when the trees are felled. The
coniferous forest may be succeeded by a wholly worthless
growth of ‘‘scrub’’ oaks and other deciduous leaved plants,
as for example on the Ossipee Plains in southeastern New
Hampshire, or by hardwood forest of diverse composition,
as in other parts of New England, or it may be followed by
desolation, bare sand or rock, as in the region around
Truckee and about Lake Tahoe in California and Nevada.
Not so with the redwood forest.

[83] March 14, Igor.

84 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3p SER.

In the National Geographical Magazine for May, 1899,
Gannett asserts that a region naturally forested with red-
wood will not become reforested with the same tree if the
standing timber is felled. Hesays(l.c., p. 151) ‘*Nowhere
is there any young growth. The youngest trees, which are
found only in the northern portion of the [redwood] belt,
are several hundred years of age. When the timber has
been cut, there is no sign of reproduction from seed. In
many localities sprouts are starting from stumps in the cut
areas, but even this form of reproduction is limited. In-
deed, everything seems to indicate that for some reason,
probably a progressive drying of the climate, the forest
environment is not favorable to the growth of redwood, and
that with the clearing away of the present forests, the end
of the species as a source of lumber will be at hand.’’
Gannett furnishes a photograph of sprouted redwoods in a
cut area.

It is true, as Gannett says, that the major part of the red-
wood forest is north of San Francisco, especially in Hum-
boldt County, California; butin the Santa Cruz Mountains,
south of San Francisco, there are more than merely ‘‘ scat-
tered groves’ of redwood trees. The amount of redwood
lumber here cut is evidence that the redwood has attained
profitable size and that it still occurs in profitable quantity.
Redwood forests must have been abundant in the mountains
between the southern arm of the Bay of San Francisco and
the ocean, even within comparatively recent years. Much
of what was once forested land is now tilled, but by the
roadsides and along the fences one sees great blackened
stumps which prove the recent presence of redwood forest.
In the cafions and on the steeper hillsides, where land can-
not under present conditions be profitably cultivated and
need not be used for pasture, some of the old redwoods
remain. In these same places young redwoods are coming
up, some from the stumps, more from the uninjured and
still living underground parts of trees which have been
felled, and some from seed. So far as this region is con-
cerned, the alarm which Mr. Gannett’s remarks arouse
seems not to be well founded.

BoT.—VOL. II.] PETRCE—SEQUOIA SEMPERVIRENS. 85

The redwood is a tree of fairly rapid growth. I can not
base my opinion on measurements, but I believe that the
sprouts from the stumps and underground parts of old red-
woods which have been felled grow faster than plants
from the seed. It appears, from what I shall presently
report, that in the living underground remnants of old trees
there are great quantities of reserve food which are avail-
able for the nutrition of sprouts. These sprouts or suckers
are not wholly dependent upon the food they themselves
elaborate. So long as their connection with the parent tree
continues unbroken, and the remnants of the old tree retain
their vitality, the young trees can use the food stored in the
parent. For this or for some other reason, the young trees
which begin as suckers and sprouts attain a considerable
height and diameter within a few years. Young trees of
this sort, in the cafions and on the mountain sides in various
places which I have visited, not far from the Stanford Uni-
versity, already have very considerable dimensions. Such
trees may be seen above Portola and above Los Gatos. In
comparison with the great redwoods of the virgin forest to
the north, these young trees are very small. They are still
absolutely as well as relatively of no value as timber, but
they have been growing only a few years. If they are
allowed to continue to grow, if they are reasonably pro-
tected against drought by having the watershed above and
about them as little disturbed as possible, I can see no reason
why valuable redwood timber should not continue to be
produced in these cajions at least.

Mr. Gannett’s account of the lumbering operations in the
northern redwood forest reveals one, and it seems to me an
adequate, reason why the redwood forest is succeeded
neither by a second growth of seedlings of its own sort nor
by a vigorous growth of sprouts from the stumps and under-
ground parts. Because the redwood is free from resin and
contains much water, the freshly felled trees do not burn
readily. For this reason, the lumbermen commonly clear
away the rubbish around the trunk of a felled tree simply
by setting fire to the brush. In this way the foliage and

86 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

smaller branches are consumed and the main trunk becomes
accessible, blackened by the fire, the bark more or less
burned through, but the wood uninjured. A fire hot enough
to burn up so much green rubbish, though not hot enough
to impair the value of the great felled trunks as lumber, is
surely hot enough to do great damage to the superficial
parts of the stump if not to kill it. Such a fire would
probably heat the ground enough and deep enough to injure
or kill the underground parts, and it would surely destroy
all seeds not deeply buried in the soil. Land cleared in any
such way as this usually has to be restocked by plants that
wander in, their seeds being blown or brought in by wind,
animals, man, etc. It seems to me, therefore, that the habit
not of the redwood but of the lumberman is responsible for
the failure of the northern redwood forest to renew itself.
In view of these facts, is it not unnecessary to imagine any
harmful change, if change at all, in the climate of the
Pacific Coast since the redwoods have lived here ?

Young redwood trees grown from the seed under some- —
what artificial conditions often send up suckers from the
trunk at or slightly below the level of the ground. A con-
siderable number of redwoods in the Arboretum of Stan-
ford University are doing so. In some instances this may
be due to injury to the upper parts of the tree by fungus or
animal enemies, but apparently not in all. It seems to me
much more likely that the more abundant branching at the
bases of these young trees is a reaction to the larger
amount of light which falls upon the surface of the soil and
upon the lower parts of the trees in the comparatively open
Arboretum than in the natural forest. But the Arboretum
is not only more brightly lighted than the forest; it is warmer
by day and colder by night during some if not all seasons
of the year; it receives much less moisture both as rain and
as fog than the naturally forest-clad hill and mountain sides;
and the soil is not able to retain so much moisture because
the ground is naked or nearly so. In the natural forest,
unharmed by sheep or man, the ground is covered bya
thicker or thinner layer composed of humus, decayed

Bot.—Vot. Il.] PEZRCE—SEQUOIA SEMPERVIRENS. 87

leaves, leaves only recently fallen, and a great variety of
small plants—mosses, lichens, etc.—which form a turf.
This covering of the forest floor, as every one knows, is
the most important natural means of holding back water,
restraining it from too rapid flow, and holding it against
evaporation. Some or all of these differences between the
natural and the artificial habitat of these trees may act upon
them as stimuli to which the production of suckers is the
visible reaction.

The Palo Alto, the only large redwood tree still standing
on the floor of the Santa Clara Valley, so far toward the
Bay of San Francisco, has been subjected to many disturb-
ing influences. Its crown has been seriously injured and
its underground parts have been subjected to changed
environment. The proximity of the railway embankment
and bridge have caused changes in the drainage, both sur-
face and subsoil, and other disturbances less evident must
also have occurred. Around the base of this old tree,
growing thickly and closely about it, is a brush or thicket
of suckers. No young trees have grown up around the.
parent, forming a little grove such as one sees around the
stump of a felled or fallen redwood of advanced age. Only
these suckers are formed, close around the trunk, and these
are not likely to attain any considerable height or size.

So far as I know, it is only when conditions are unlike
those prevailing in the natural forest, or when an old tree
has been felled or injured or at least considerably disturbed
above or below ground, that it sends up suckers from the
trunk or stump, or that young trees come up from the re-
moter underground parts. These last often make circular
groves of greater or less size, known as ‘‘redwood temples.”’
Even in the group of giant redwoods at Felton, near Santa
Cruz, one sees this arrangement clearly marked. The
suckers and sprouts may attain great size in the course of
time, as some of these giant redwoods show.

In the production of suckers or sprouts from the trunks
and underground parts of Seguota sempervirens, we see
the vegetative mode of reproduction engaged in bya species
of the Conifer; but this recourse to the vegetative mode of

88 CALIFORNIA ACADEMY OF SCIENCES. [PRroc. 3D SER.

reproduction is probably the result of some external stimu-
lus or stimuli acting upon the plant which, under unchanged
conditions, would continue to reproduce itself by seeds if
at all. This method of vegetative reproduction is probably
sufficient, even under present climatic and other conditions,
to secure the continuance of redwood forests in the regions
where they now occur, frovided lumbering operations are
so conducted that the production of suckers and sprouts is
not made impossible by destructive fires.

II. PECULIARITIES OF SOME VEGETATIVELY PRODUCED
YounGc REDWoOODs.

A. Fasctation.

Fasciation of the young suckers coming up around the
trunks of redwoods is not uncommon. The view advocated
by Frank (1896), that they are the consequences of an
excess of food substances, is strengthened by the time and
manner of their appearance. Frank says that fasciations
on other plants appear especially when the ordinary
branches have been removed or injured in any way. We
have seen that the redwoods produce suckers probably
only when stimulated to do so by external influences, espe-
cially by the removal or at least the injury of the parts above
ground. The wound, or other injury, which stimulates the
redwood to form suckers, may occur when there is such an
abundance of food in immediately usable form, that the
production and growth of suckers is so prolific as to insure
the fusion of the adjacent parenchymatous parts of the very
young branches. In the autumn and early winter, I have
had no difficulty in finding fasciated redwood suckers in the
Arboretum of Stanford University; they are very notice-
able. In the spring and summer months they are by no
means so common. I have found no new ones this spring,
though there are many young suckers on the redwoods in
the Arboretum. Summer, at least the earlier half of the
dry season, is the time of food manufacture and storage.
In the latter half of the dry season little food can be manu-
factured because only little water is obtainable. In autumn

BoT.—VOL. II.] PEITRCE—SEQUOIA SEMPERVIRENS. 89

the food manufactured and stored becomes available and is
used after the first rains have made it possible for growth to
be renewed. If, as was the case last winter, there is much
mild damp weather, growth will be luxuriant, the stored food
will be freely used, the conditions for sucker-formation and
for fasciation will coincide. In the spring, especially after
a mild winter, during which much growth and comparatively
little food manufacture have taken place, the stores of food
having been considerably reduced, growth will be less luxu-
riant and food manufacture will become more necessary and
more active. We see, then, some reasons for the formation
of the fasciations and for the time of their appearance.

B. Albinism.

The most remarkable and, I am surprised to find, not an
especially rare peculiarity of the suckers or sprouts which
come up from the stumps or from the old roots of felled or
fallen redwood trees, is that they are sometimes perfectly
white. My attention was first attracted to this peculiarity
when, in the fall of 1898, a student brought some redwood
twigs bearing white leaves into the Botanical Laboratory of
this University. On inquiry I learned where these white
redwoods were growing, and in the fall of 1899 I went to
the spot. These are the only white redwoods which I have
seen growing, but I have heard of others much larger and
one which must be several years old was brought to the
laboratory from the ‘‘ Redwood Retreat,’’ about twelve
miles from Gilroy. This last I planted in my garden, but it
lived only a short time, whether because it was injured in
the transplanting or because it could not bear transplanting
late in the spring I do not know. Its behavior before it died
I will speak of presently (p. 95).

The white redwoods which I have visited are on the sum-
mit not far from the stage road between La Honda and
Redwood City, and on the line to the left of the road (as
one goes toward La Honda), where the forest gives place
to open fields. The tallest redwood tree in view marks the
spot where the white ones grow, ‘This tall tree is one of a

90 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

number which are several decades old. They, and other
smaller young trees of various ages, have come from the
stump and roots of a much older tree which must have been
very large when it was felled. The old stump has been
repeatedly burned under and into, apparently by camp fires,
but the heat could not have been enough to do more than
local damage, and even this is not great. There is little
left of the old stump above the general ground level, but as
the hillside falls away abruptly at that point, a good deal of
the underground part of the old tree is more or less exposed.
Because of the irregularity of the surface of the hillside
just there, the old tree sent its roots out more irregularly
than is usually the case and the trees which have sprung up
from them are not symmetrically placed. There is a
thicket, but not a circle or ‘‘ temple,’ of redwoods. All of
these second growth trees are perfectly normal, as far as I
could see. One buttress of the old parent tree, instead of
sending up a few more or less scattered sprouts which grow
up fairly rapidly and, within a season or two take on the
characters of young trees in bark, foliage, and manner of
branching, produces branches or bunches of scrubby,
thickly set, short and slender sprouts or suckers. These
are perfectly white as to leaves. The youngest parts of the
stems are of the same ghostly color as the leaves. These
white suckers may attain a height of thirty (30) centimeters
in the course of one season. They began growing early in
April this spring (1900), and they go on growing till hard
frost comes. In the same length of time, and with a simi-
lar origin, a green sprout or sucker would make two or
three or more times this growth in length. The white
suckers increase in thickness proportionally to their growth
in length, that is, slowly, but the surface of the stem be-
comes brown and develops cork sooner than the correspond-
ing parts of the green suckers. This precocious cork-for-
mation is not accompanied by other means of protection or
by such vigor that the white suckers survive the hard frosts
of winter. Even this last winter, milder than usual, was
fatal to the white suckers; they were killed down to or just

BoT.— VOL. II.] PETRCE—SEQUOIA SEMPERVIRENS. oI

below the surface of the soil. The green suckers, on the
other hand, are enough tougher to survive the winter. In
this spot, therefore, white suckers with parts above ground
which are two years old are not tobe found; but near and just
under the ground are well-formed buds which, surviving the
winter cold, form the next year’s growth of white suckers.
In this difference in ability to resist cold, we have one of
the physiological differences between the dependent white
suckers and the independent green suckers. Whether this
is merely a coincidence or a fundamental difference in vigor
which forces the white suckers, unable to form chlorophyll,
to draw food from the parent if they are to survive, who
can tell ?

This difference between white and green suckers is not
everywhere visible. White redwoods of fair height and
age are reported, indefinitely to be sure, from various
places in the Santa Cruz Mountains. I have seen white
redwoods several years old. These, however, came from
places of lighter frosts, if any frost at all touches them
during the winter.

Turning now to the anatomy of the white and the green
suckers, we see certain peculiarities in the white which
demand remark. The leaves evidently present the most
marked differences. The leaves of the white suckers are
similar in size, form, kind and arrangement to those of the
green. here are two kinds of leaves on both green and
white suckers—the early or young form, large, long, few,
scattered along the stem or branch—and the later or mature
form, smaller, shorter, more numerous, regularly placed along
the branches, giving to these leafy branches the typical flat
and thin dorsi-ventral aspect as compared with the more
nearly radial arrangement in the young plant. These two
forms, found on the suckers, are also found on seedlings.
According to Goebel (1898), the young form is to be re-
garded as presenting the original leaf-form and leaf-arrange-
ment in the Gymnosperms. The rudimentary characters to
be discussed later, which are found in the leaves of the
white suckers, would lead one to believe that sooner or later

Q2 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

suckers would be found which retain this primitive charac-
ter for considerable periods, if not throughout their exis-
tence. If Goebel’s other supposition is true—that the basal
branches, showing the young form of leaves, disappear by
correlation as the mature form develops—one would infer
the greater physiological perfection and effectiveness of
the mature as compared with the young form. But in the
white suckers the main function of leaves, that of photosyn-
thetically manufacturing non-nitrogenous food, is entirely
suppressed—the mature form of white leaves being as im-
potent as the young form. ‘Two other important functions
of leaves—that of securing and controlling transpiration,
and that of securing and controlling the aération of the
plant-body,—would apparently be as perfectly prepared for
and accomplished by the one form of leaves as the other,
provided the texture and the surface-area of the two kinds
of leaves are equal. In these two respects the mature
leaves are superior to the young form. There is, therefore,
some reason, other than the conception of an ‘‘inherited
tendency,’ for the development of the mature form of
leaves in the white suckers.

Since the Seguozas are geologically such old plants, it is
interesting to have the young form of leaves so clearly

marked and so constantly recurring. There is a striking
resemblance between some of the green suckers in their
young condition and the great fronds of Cycas revoluta, the
leaves of the redwood resembling the leaflets of Cycas in
form, thickness and arrangement. Can this be a hint as to
the origin, perhaps the common origin, of the Coniferz and
the Cycadacez ?

Comparing the mature form of leaves of green and of
white suckers from the same localities, one finds that,
despite the superficial likenesses, there are decided struc-
tural differences. These are at once evident in cross-
sections of the leaves, as shown in figs. 1 and 2. Figure 1
is a diagram of a cross-section of a small green redwood
leaf, the single vascular bundle occupying the centre of the
leaf, one resin-tube lying under it, the other two resin-tubes

Bot.—VOL. II.] PEITRCE--SEQUOIA SEMPERVIRENS. 93

being located at the ends. The upper surface of the leaf is
convex, the lower concave, the upper surface and the edges
being greatly strengthened by the thickened and heavily
cutinized walls of the epidermal cells and by the underlying
single layer of sclerenchyma. Beneath this is the very per-
fectly developed palisade parenchyma, extending from edge
to edge of the leaf. The remainder of the mesophyll is com-
posed of simple, unbranched parenchyma, enclosing many
intercellular spaces, and bounded on the under side of the
leaf by the single layer of fairly thick-walled epidermal cells.
In contrast to this, fig. 2, a similar diagrammatic view of
the cross-section of a larger white leaf shows a less convex
upper, a less concave lower, surface, and the almost or quite
complete absence of sclerenchyma cells except at the edges
of the leaf (see fig. 9). The most striking difference in
the structure of the two leaves, however, consists in the
complete absence of palisade parenchyma from the white
redwood leaf. The remainder of the mesophyll is com-
posed of somewhat larger parenchyma cells than in the
green leaves, and the intercellular spaces are also slightly
larger.

Examination of these cross-sections under higher magni-
fication (figs. 7-10) reveals still more plainly the contrast
between the white and the green leaves. Figure 7 is a
detail from near the middle of the green leaf represented
in fig. 1. Figure 7 shows the thick-walled epidermal, the
thicker walled sclerenchyma, cells, and the regular palisade-
parenchyma cells. These contain many chloroplastids,
slightly larger than the starch-grains indicated in the figure.
There are many of these starch-grains imbedded in the
cytoplasm. Vacuoles are numerous, and evident. The
cytoplasm and nucleus are sharply differentiated. Figure
8 represents the corner of fig. 1 cut off by the dotted line.
In fig. 8 are shown the very strong sclerenchyma cells im-
mediately underlying the epidermis at the edge of the leaf,
and extending almost continuously along the under side to
and around the resin-tube. This last is large and bounded
by many suitably supported, thin-walled, glandular cells.

94 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

At the edges, as well as elsewhere throughout the palisade
and mesophyll tissues of the green leaves, starch-grains
occur in considerable numbers in all the cells. The amount
of starch present undoubtedly varies at different seasons of
the year; but since these green leaves were collected at the
same time as the white ones, and near them, comparison of
the starch content is justified.

Figure 9 represents under higher magnification the part
of fig. 2 cut off by the dotted line. As this shows, scleren-
chyma cells are found at the edges of the white leaves, but
they are not so firm as in the green leaves. The resin-tube
is decidedly smaller and bounded by cells evidently less
active than the glandular cells of the resin-tubes in the
green leaves. There is no starch in any mesophyll cells of
the white leaves. The cytoplasm and nucleus are not easily
distinguishable from one another in most of the mesophyll
cells. The cytoplasm presents a thoroughly disorganized
or, as one may more truthfully say, a by no means organ-
ized, appearance, neither vacuoles nor plastids being dis-
cernible in most cells. The vacuoles occur in few cells
only and are unlike those of green mesophyll cells.

The plastids are variable. In sections of white redwood
leaves from the summit near where the Redwood City—La
Honda stage road crosses the first ridge between here and
the sea, I have entirely failed to detect even rudiments of
plastids or chromatophores. There are granules and gran-
ular aggregations in the cells, as fig. 9 shows, but material
carefully fixed in Flemming’s weaker mixture of chrom-
osmic-acetic acids and stained by two-tenths per cent. acid
fuchsine in distilled water* failed to exhibit any structures
which I could positively identify as even rudimentary chro-
matophores. On the other hand, the material from near
Gilroy, treated in exactly the same way, contained chro-
matophores which ranged in size from those about half as
large as the average chloroplastids in the normal green
leaves down to indistinguishable rudiments.

* See Zimmermann-Huniphrey, Botanical Microtechnique, pp. 196, 202, etc.

Bot.—VOt. II.] PEITRCE—SEQUOIA SEMPERVIRENS. 95

In this connection I may state that the white redwood
taken from near Gilroy and planted in my garden had at
least one leaf which, after the little tree was transplanted
and before it died, became pale green over half its surface
on either side of the midrib. I have made no attempt as
yet either to transplant, or to disconnect from their parents
without transplanting, the white redwoods growing on the
summit near the La Honda road. This I shall do presently.
Obviously the white redwoods must turn green if they are
to survive after being severed from the parent. Some white
redwoods can do this more readily than others, the condi-
tion of the chromatophores being one of the factors con-
trolling this change. Whythere should be these differences
in the rudimentary condition of the chromatophores of white
redwoods we can understand only after determining the
reason for the production of any white leaves at all.

Comparing white redwoods with cedars, which, in culti-
vation and in nature, not infrequently produce white leaves
or green leaves striped or otherwise variegated with white,
we find the cell-structure as well as the general conditions
for the nutrition of the plants quite unlike. The mesophyll
cells in the white parts of green variegated leaves, and of
white leaves, contain less protoplasm (Frank, 1896) than do
normal green cells, the cytoplasm forming a comparatively
thin layer lining the cell-wall, the greater part of the cell-
cavity being filled with the more than usually abundant
cell-sap. Chromatophores, if visible at all, are colorless,
small, and scarcely denser than the cytoplasm, or otherwise
distinguishable from it, but they vary in this respect with
the degree of whiteness of the leaves. The white redwoods
are similar, the cells of the whitest containing no structures
recognizable as chromatophores, while those leaves which
contain visible chromatophores are not perfectly white.
But between the unorganized though abundant protoplasm
in the mesophyll cells of the white redwood, and the meagre
but very definitely situated protoplasm of the white meso-
phyll cells of cedar and similar plants, there is a great dif-
ference. This we may perhaps account for thus.

96 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

Cedars and other plants with white or white variegated
leaves are independent, manufacturing in the leaves and
other parts that are green the non-nitrogenous foods needed.
Any reduction in the number of green leaves, or in the
number of chlorophyll-containing-cells in the mesophyll, is
a reduction in the capacity of the plant to manufacture
food; and if all the leaves turn white the plant will die as
soon as the food is consumed which was elaborated and
stored while its leaves were green. The turning white, or
the failure to become green, of the leaves or any part of
the leaves of cedars, etc., is a variation neither useful nor
permanent; it is really a morbid condition.

White redwoods, such as I have seen and here describe,
are not independent. They absorb from the still living
underground parts of the parent tree the non-nitrogenous
foods (starch, sugar, etc.) manufactured in its own green
leaves and stored in its own underground parts. These
stores of food are very great in old redwoods. When for
any reason a sucker starts with none of its leaves green, it
is exactly as well off, so long as the store of food in its par-
ent lasts, as if its leaves were green and as if it could manu-
facture food for itself. The activities and possibilities of the
white sucker are not abruptly terminated by the exhaustion
of its own very limited store of food. It can and does draw
on its parent for much food. In the variegated cedar we
have some leaves shirking their function as food-manufac-
turing organs, either because they were defective from the
time of their origin at the growing-point, or because they
have developed this pathological condition subsequently.
The white redwoods, on the contrary, are the vegetatively
produced offspring of a wholly independent organism which
live as parasites. They take on some of the characters of
parasites, as is shown by the absence of palisade cells in the
leaves, and by the rudimentary condition of the chromato-
phores and other protoplasmic contents of the mesophyll
cells. They are also less vigorous and grow less rap-
idly than wholly independent though similarly produced
individuals.

Bot.—VOL. II.] PETRCE—SEQUOIA SEMPERVIRENS. 97

Other differences in structure between the white and the
green leaves of redwood may be mentioned. As shown by
figs. 11, 12 and 13, the walls of the epidermal cells of the
white are not as thick as those of the green leaves. Figure
tr shows the epidermal cells around and above the some-
what depressed stoma on a white leaf. Figure 13 is a similar
view of a part of the surface of a green leaf. Both figures
are of a stoma from the upper surface. Though in the green
and in the white the mouth of the stoma is about equal in
size, the adjacent epidermal cells are smaller as well as thin-
ner walled in the white redwood. Figure 12 represents the
guard-cells of the stoma, only the upper part of which is
shown in fig. rr. There are no chromatophores in the
guard-cells, but the nuclei are well differentiated.

Though the numbers of stomata on the under side of the
green and the white leaves are about equal, there are more
stomata on the upper surface of the white leaves than of the
green. Figures 3 and 4 represent very diagrammatically
the shape and size, but exactly the numbers, of the stomata
in equal areas of epidermis from the upper side near the
midrib of a green and of a white leaf from redwoods in the
Santa Cruz Mountains. Figure 5 indicates the number of
stomata in an equal length of epidermis similarly situated
but from a green leaf from one of the redwoods in the
Arboretum of Stanford University. Figure 6 is another
strip from a white redwood leaf from the mountains. Two
facts are demonstrated by these figures: first, that the white
leaves always have more stomata on the upper side than do
the green ones; second, thatin the green leaves the number
of stomata on the upper side of the leaf is proportioned to the
humidity of the region in which the tree grows. ‘The sec-
ond isa fact well known and understood; the first is new
and not easy to understand. In all probability it will be
found that the white redwoods occur where transpiration is
not so great as in many places in the mountains where green
redwoods occur. Ina thicket there would evidently be less
rapid transpiration than in the open at one side or above the
thicket. The white redwood suckers which I have seen are

98 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

certainly in a situation in which transpiration can not be
very rapid while the suckers are actively growing.

We come now to consider what causes the suckers of
certain redwoods to be white while others are green. It is
evident that the white redwoods are not white from etiola-
tion, for other suckers similarly situated, and other plants
all about, are as green as usual. Nor can lack of iron be
the cause, for the same reason. So far as I can see, the
only reason for these plants being white is that the leaves
form, and attain nearly or quite their full size, at a season
when there is insufficient warmth for the formation of
chromatophores and chlorophyll pigment, though enough
for growth. This is in perfect harmony with Sachs’s obser-
vation (1864), since extended by Frank (1895), that seed-
lings growing and buds unfolding at low temperatures
produce leaves yellowish or white, either wholly or in
patches. I had occasion to notice this phenomenon par-
ticularly during the past winter in the leaves of Bur Clover
(Medicago denticulata Willd). In December and January
there were warmth, moisture, and light enough for a lush
vegetation composed of the common annuals, but the nights
were chilly and the days not warm. ‘There was an unusual
amount of variegation in the leaves of the common weeds.
That there was light enough for chlorophyll formation is
evident from the fact that Bur Clovers growing in the labo-
ratory had no white or variegated leaves. As Sachs proved
by experiment, the plant must have a certain minimum
amount of warmth in order to form chlorophyll. This min-
imum, higher than the minimum for growth, of course
varies with the species.

The white redwoods growing on the crest near the La
Honda road are killed down to the ground each year by the
frost. This is evidence of considerable cold. In January
of this year (1900), when I visited these white redwoods,
they had been frosted down but, buried in the leaf-mould
covering the branch of the old stump from which the white
redwoods spring, were many buds, healthy, with well formed
but perfectly white leaves. On that summit no temperature

BoT.— VOL. II.] PEIRCE—SEQUOIA SEMPERVIRENS. 99

records are kept. I therefore cannot tell the temperatures
prevailing when these buds were forming in darkness under-
ground. The darkness is of no significance, for all suckers
begin in darkness and ordinarily their leaves are then green.
Insufficient warmth seems, then, to be the reason why the
chromatophores and the chlorophyll pigments do not form
in the cells of these growing leaves.

Frank (1895) says that it frequently happens that plants
with leaves white because of cold at the time of their form-
ation often retain these white leaves even into the summer,
subsequent warmth being insufficient to stimulate to chro-
matophore and chlorophyll formation. On all such plants,
however, the leaves developed later than the white ones,
and when the temperature is higher, are green. If this
were not the case, the plants would die.

In the white redwood we have a different state of affairs.
Although the first leaves borne on a shoot of one year’s
growth may all have been formed in the cold, late in the
previous year, and therefore may not be able to turn green,
the leaves later formed, and the internodes forming or at
least elongating later, when there is sufficient warmth, one
would expect to find green. On the contrary, once started
as white redwoods, the suckers continue white as to leaves
and young cortex for an indefinite time.

The differences in the color, and in the condition of the
chromatophores between the white redwood leaves from
near the La Honda road, and those from near Gilroy, may
be accounted for thus. It is probable that near Gilroy, at
least in the spot where the white redwoods grow, the tem-
perature is not so low as on the exposed summit crossed by
the La Honda road. While the temperature is low enough
to prevent chlorophyll formation, it is not low enough com-
pletely to suppress the formation of chromatophores, and
by no means low enough to interfere with growth. Slight
variations in low temperatures at the times when the buds,
from which suckers spring, are forming, might permit the
formation of green buds, of yellowish buds with rudimentary
chromatophores, and of white buds with no chromatophores

(2) March 18, root.

I0O CALIFORNIA ACADEMY OF SCIENCES. [PRroc. 3D SER.

at all. Certain it is that all the white redwoods I have seen
or heard of grow where the temperature is low in autumn
and winter.

Ill. Tue SIGNIFICANCE OF THE WHITE REDWOODS IN
CONNECTION WITH OUR CONCEPTIONS OF PARASITISM
AND OF HEREDITY.

White redwoods are wholly dependent, absolutely para-
sitic, plants which are in their first generation. They are
not the offspring of other white redwoods, they are not the
descendants of a long line of more and more dependent,
more and more degenerate, organisms. Their parasitic
characters have been acquired, or developed, during the
brief course of their own existence, but they possess some
parasitic characters not yet acquired by plants which have
been semiparasitic for no one knows how long. /Phora-
dendron, Viscum, and the other ‘‘green parasites’’ have
long lived at the expense of the other plants upon which
they grow; but though attached to their hosts, these para-
sites manufacture in their own green leaves their own non-
nitrogenous foods.

As I have shown elsewhere (1893), the ‘‘green parasites”’
which have been studied differ from completely parasitic
flowering plants (e. g. Cuscuta, Brugmansia, Rafflesia) in
the completeness of the connection between the tissues of
host and parasite effected by the haustoria. In the com-
plete parasites, xylem and phloém of the parasite are directly
connected with the xylem and phloém of the host by means
of xylem and phloém tissues which are continuous through-
out the haustoria. In the ‘‘green parasites,’’ on the other
hand, only the xylems of host and parasite are directly con-
nected. This anatomical difference may be considered the
reason for the difference in the degree of parasitism in
these two sets of plants, or we may conceive that, so long
as the parasite remains green, and therefore able to manu-
facture its own food, a complete connection with both
sets of conducting tissues in its host is unnecessary and
unformed. There is at present no means to decide which

Bor-—VOoL. Il. PEIRCE—SEQUOIA SEMPERVIRENS. IOI

is true. In the first place, we do not know the actual func-
tions, and cannot determine the entire significance, of the
phloém tissues in higher plants. In the second place, so
long as Trelease (1894), or any one else, can say without
proof of error, that Leztneria floridana, a tree, contains no
sieve-plates and inferably no sieve-tubes, the elements of
phloém commonly regarded as most essential, it is impossi-
ble to conclude that sieve-tubes are indispensable or that a
phloém as well as xylem connection between host and para-
site is essential for complete parasitism. Until the chemical
physiology, and not the anatomy only, of the relation exist-
ing between parasite and host in Viscum, Phoradendron,
Cuscuta is worked out, it cannot be known how significant
and important are the tissue connections effected by the
haustoria.

As to the significance of these tissue connections, the
conditions presented by the white redwood may furnish
some idea. The dependent white redwoods are branches
of independent parents and are therefore connected from
their beginnings, xylem with xylem, phloém with phloém,
and parenchyma with parenchyma, with their parents. By
means of these connections the adequate supply of foods,
as well as of food-materials and water, by the parent to its
offspring, from old redwood host to parasitic sucker, is
assured from the first. On the other hand, a parasite
attacking the tree from the outside must establish these
connections. It may establish them only imperfectly, as in
Viscum, or completely, as in Cuscuta. The white redwood,
with its perfect connection with the parent, offers the coun-
terpart of the condition which accompanies complete para-
sitism, and though the leaves persist as such, they are struc-
turally no longer perfect leaves, and physiologically only
partly so. Because of its perfect connection with the host,
the white redwood is able immediately to develop some feat-
ures of the characteristic structure of parasites.

This is especially interesting because the white redwoods
are the vegetatively produced offspring of independent
plants, themselves the descendants of generations of inde-
pendent plants. The suckers of redwoods inherit the

102 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

tendency, which one would expect to find firmly fixed, to
develop into independent plants, green and manufacturing
their own foods. But these young suckers, originating when
the weather is too cool for chlorophyll and chromatophore
formation, though warm enough for a certain amount of
growth, beginning with white leaves and growing well
enough during the season without manufacturing food for
themselves, form no chlorophyll even in the leaves and
internodes later developed. The effect of low temperatures
when the first leaves were forming was either upon the
protoplasm itself, preventing its forming chromatophores and
chlorophyll, or upon the chemical processes by which these
organs and substances are produced. One naturally as-
sumes the former—that the powers of the protoplasm are
lessened by low temperatures. In these first leaves, the
protoplasm is prevented by the cold from following its in-
herited tendency to produce chromatophores and chloro-
phyll. In the leaves later formed, the inherited tendency
to form chromatophores and chlorophyll is not interfered
with by cold, but it does not cause these leaves to become
green. They do not need to be green; the plant obtains
food enough without turning green and manufacturing its
own food. The inherited tendency is not aroused by hun-
ger into action. The stimulus needed to set it in operation
not being given, the inherited tendency remains dormant as
long as the white suckers remain connected with the parent.
The parasitic habit forced upon the young white sucker by
its inability to manufacture its own food, and the parasitic
characters assumed by the young white sucker, are continued
as the plant grows. Continued healthy existence in spite of
inability to manufacture food induces in new leaves and
cortex those characters found in the earlier and older ones.
Environment, the influence of certain stimuli, induce a re-
action ordinarily characteristic of species of plants which
have been parasitic for generations.

The white redwood serves as an index of the relative
powers of heredity and of environment, or, more definitely,
of heredity and of the influence of, and the power of

BoT.—VOL. II.] PEIRCE—SEQUOIA SEMPERVIRENS. 103

reaction to, certain stimuli. There being no need to man-
ufacture food, the food-manufacturing apparatus is not
formed, a parasitic habit being successful so far as the
individual is concerned, the inherited habit is not entered
upon.

That the need to manufacture food would have an effect
upon the development of the white suckers is indicated by
the behavior of the white sucker which grew near Gilroy
and which I planted in my garden where it could obtain
little if any organic matter as food. It died soon after
transplanting, but zot untzl one leaf had become pale green.
The effect of cutting white suckers away from the parent
stock and from their supply of manufactured food I shall
test presently by experiment on the redwoods near the La
Honda road; but this experience is not without significance.
At least it strengthens my contention that inherited tendency
is less strong than environment, and that in some cases, at
least, inherited tendency must be called into action by some
specific stimulus or combination of stimuli operating upon
the plant from outside itself. In our white redwoods, the
descendants of an exceedingly ancient race of trees in which
heredity should be proportionally strong, we have a certain
amount of evidence that the irritability and the power of
response of the organism to external influences are stronger
than its heredity. May not this always be the case? May
it not be that what we call heredity is really the response to
similar stimuli and combinations of stimuli occurring in
orderly succession in the course of nature?

STANFORD UNIVERSITY,
CALIFORNIA,

June, 1900,

tee

eh

; ‘ "ec 7 rey,
ea eae. See ee te a

Bot.—VOL. II.] PEIRCE—SEQUOIA SEMPERVIRENS. IOS

1895.
1896.
1899.

1898.
1893.

1864.

1894.

1893.

BIBLIOGRAPHY.

FRANK, A. B. Krankheiten der Pflanzen. Bd. I. Breslau.

The same, Bd. III.

Gannett, Henry. The Redwood Forests of the Pacific Coast.
Nat. Geograph. Magazine, Vol. X.

GOEBEL, Kart. Organographie der Pflanzen. Bd. I. Jena.

PEIRCE, GEORGE J. On the Structure of the Haustoria of Some
Phanerogamic Parasites. Aznals of Botany, Vol. VII.

Sacus, JuLtus von. Ueber den Einfluss der Temperatur auf das
Ergriinen der Blitter. /Vora, 1864. Reprinted 1892, in Gesamm.,
Abhandlungen, Bd. 1.

TRELEASE, WILLIAM. Leitneria floridana. Sixth Ann. Report
Missouri Bot. Garden.

ZIMMERMANN, A. Botanical Microtechnique. Translated by J. E.
Humphrey. New York.

106

Fig.

Fig.

on

Io.

CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

EXPLANATION OF: PLATE XIV.

All figures were drawn with an Abbé camera lucida.

Cross-section of green redwood leaf, showing position of vascular
bundle and resin-tubes, the well developed palisade parenchyma
and sclerenchyma. X 30. Diagrammatic.

Cross-section of a larger white redwood leaf, showing the absence
of palisade parenchyma and sclerenchyma. xX 30. Dia-
grammatic.

Strip of epidermis from upper side of green redwood leaf, showing
distribution of stomata near midrib. X 60.

Similar strip from white redwood leaf, showing distribution of
stomata in an equal area. X 60.

Figures 3 and 4 are from mountain redwoods.

Similar strip from green redwood leaf, showing number and dis-
tribution of stomata. X 60.
Similar strip from white redwood leaf, showing the same. X 60.

Figure 5 is from redwood growing in Arboretum of Stanford
University, fig. 6 from mountain redwood.

Part of green leaf shown in fig. 1, from above vascular bundle,
showing palisade parenchyma, sclerenchyma, starch-grains, and
other contents of palisade cells. X 300.

Part of figure cut off by dotted line, showing starch-grains and
other contents of mesophyll cells, thick-walled sclerenchyma and
epidermis at edge of leaf, strengthening cells around resin-tube,
etc. XX 300.

Corresponding part of fig. 2, a white redwood leaf, showing unor-
ganized contents of mesophyll cells, lighter strengthening tis-
sues, etC. X 300.

Showing structure of stoma of white redwood and absence of
chlorophyll and starch-grains. X 300.

Figs. 11 and 12. Two views from the surface of stoma from white redwood

leaf; fig. 11 from above, showing auxiliary cells (Vebenzellen);
‘fig. 12 from further down, showing guard cells. Stoma closed.
X 350.

Fig. 13. Surface view of epidermis and stoma from green redwood leaf,

showing larger and more vigorous epidermal cells, and decidedly
thicker walls of epidermal cells. X 350.

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[PEIRCE] PLATE XIV.

Proc. CALAcAD. Scr.3? Ser Bor. VonI!

A REVISION OF THE GENUS CALOCHORTUS.

BY CARL PURDY.

PLATES XV-XIX.

INTRODUCTION.

THE most widely diffused as well as the handsomest of
the liliaceous plants of the Pacific Coast are the Calochorti.
On the north they reach British America; one species is to
be found as far east as Nebraska; several are natives of
northern Mexico; and within these limits no considerable
section of country is destitute of some species.

While the range of the genus is so immense, that of sev-
eral of the species is also very extensive. What a diversity
of conditions C’. xuttal/zz meets in its range from the west-
ern side of the Sierra Nevada to Nebraska, and from the
Snake River to Arizona. C. nztidus is found from the
meadows of eastern Oregon to the shores of Yellowstone
Lake, C. albus from San Diego to Tehama, and many
others are scattered over hundreds of miles. A species
distributed over a region so varied in soil, climate, and alti-
tude cannot but be variable; and it hardly need be added
that the genus Cadlochortus is a very difficult one for botan-
ists to deal with.

I long since became convinced that it is only in the gar-
den, where plants from different localities can be grown
under identical conditions, that the relationship between
apparently different forms can be satisfactorily determined.
For some years I have grown a large variety of Calochorti
in my grounds, and have had nearly every known species
under cultivation, often in many forms. The culture of
Calochorti is most interesting, though not unattended with
cultural difficulties; but the beautiful flowers amply repay
all efforts, and the garden has proved the identity of forms

[ 107 ] November 27, 1go!.

108 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

apparently different: here, also, variations attributed to
environment are shown to be constant. In the garden, too,
strains, which from a botanist’s standpoint seem scarcely
distinguishable, show marked differences in vigor, flower-
ing time, or immunity from disease.

It is a peculiarity of our liliaceous plants, that as a rule in
a given locality there is little variation from a well marked
type, as little, indeed, as may be found between flowers
growing upon the same plant. Hundreds and thousands of
flowers may be picked, all conforming closely to this type.
In another locality, the same species will be found marked-
ly different. The difference between the forms in the two
localities may be slight, consisting merely of a marking or a
slightly varying leaf, habit, or gland; yet the variant, once
noted, is found to be constant. In Calochortus color forms
are frequent, the flowers from one bulb retaining the same
tints under any and all conditions. The difference between
forms from different localities is rather that which florists
designate by the word ‘‘strain’’ than what is usually under-
stood to constitute a botanical species or variety.

In cultivation it has frequently been found that a very
slight variability in strains is accompanied by a marked
constitutional difference. In two beds of Calochortus venus-
tus, planted in the same soil, and separated only by a thin
board, it would puzzle a botanist to state wherein the plants
vary. They come from widely separated localities, and the
difference is one more easily detected by the eye than con-
veyed by words. In one bed, two-thirds of the leaves are
already destroyed by mildew (otryézs), while in the other,
not one leaf is injured; and such is the case whenever and
wherever the two are planted. Many similar instances
occur in other species, but a single one is sufficient to show
that the slight variations which the eye detects are not the
only ones.

Such strains are present in nearly every species of Ca/lo-
chortus. The range of a strain may be very local—a few
miles square—or it may be found over half the length of a
state. In Calochortus venustus one strain runs through all

BotT.—VOL. II.] PURDY— CALOCHORTUS., 109g

the plants found for hundreds of miles along the Sierra;
another strain is found in the same species occurring in the
Coast Range and over an equal area. In some of the more
variable species there are several strains.

In many of the Calochorti the gradations from one species
to another are so slight that it is impossible to separate them.
The extreme types on which the species are founded are
easily distinguishable, but a perfect chain of variations links
them closely together. There is no doubt that C. weed7z,
C. plummere, and C. obispoénsis are variations of a greater
species.

While, as before stated, it is the rule that a given locality
produces specimens conforming closely to a type, yet this
is not always the case. In some localities the variations are
bewilderingly numerous. I have seen places where hun-
dreds of flowers of C. venustus could have been selected,
each differing in color and markings from the rest. Why
a species that remains so true to a type in some localities
should vary so remarkably in others is a subject that will
not be discussed at present. Hybridization will account for
it in some instances, while in others it is hardly a tenable
hypothesis.

I cannot say that I attribute any material share in the
origin of the many strains or varieties to hybridization,
although among the Calochorti it is not infrequent. Such
crosses as C’. albus and C. benthamz, C. maweanus and C.
pulchellus are frequently met with, but I have never yet seen
one that was fertile. Again, varieties of a species, e. g.,
C. luteus var. oculatus and C. luteus var. citrinus, readily
cross and produce fertile hybrids. Over a small area in-
numerable cross-breeds may be found, but a few miles away
the two varieties will be found separate and varying as little
as in any locality. Then again, hybridization often will not
take place between two apparently very closely related
species. I have often seen C. vesta in flower surrounded
by large numbers of C. luteus var. oculatus or var. cztrinus,
but not a plant could be found that in any way indicated
hybridization; while last summer, in the Sierra Nevada, I

IIo CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

visited a spot where C. /uteus var. oculatus and C. luteus
var. cztrtnus had hybridized to a remarkable degree. A
few yards away,indeed mingling with them, were thousands
of C. venustus. Within a quarter of a mile I do not doubt
there were fifty thousand plants in flower, yet close search
failed to reveal one that in any way suggested a cross of C.
venustus with any of the variations of C. luteus.

In the ‘‘ Botany of California,’’ Vol. II, published in
1880, Watson described twenty-six species of Ca/ochortus.
The work was carefully done, though the material at com-
mand was meager as compared with that obtainable now;
yet nearly every species recognized by Watson stands good
to-day. Many new species have been added, but by the
exploration of new territory rather than by the subdivision
of old species.

The work of preparing the present paper has been facili-
tated by the courtesy of the California Academy of Sciences
and the University of California, in allowing me to inspect
their herbarium specimens. Mr. J. W. Congdon of Mari-
posa very courteously permitted an examination of his
material, and to Mr. G. W. Hansen I am indebted for a set
of specimens from Amador County. My personal collec-
tion, including both herbarium and living specimens, covers
a wide range; still, with these facilities, probably as good as
can be obtained anywhere, the material is painfully unsatis-
factory in some species, several of which are represented
in the best herbariums by a single specimen, if at all.

With each year appear many new forms, even from Cali-
fornia. Last season brought three new species, and many
striking variations of old species were added to the already
large assortment. The field is immense and has never been
properly worked over. In view of these facts, it seems the
wisest course to disturb existing nomenclature as little as pos-
sible. As to whether a given degree of difference warrants
a specific or a varietal name seems to me to be very largely
a matter of personal opinion. While one can hardly agree
with the author who designates a color form by a specific
name, it will probably be consulting the convenience of

Bot.—VOL. II.] PURDY— CALOCHORTUS. DLL

botanists to allow sucha name to stand for the present,
especially as a more extensive knowledge of the subject
may result in still further changes. The pressing need is
for a work containing descriptions of all known species of
Calochorti, together with such grouping as will readily con-
vey to the student the relationship of the various species.

In the ‘‘ Botany of California,’’ the types of the species
known to Watson are usually very accurately described.
The only criticism to be made is that in many instances he
was acquainted with but afew representatives of the species.
Nineteen years have added much to our knowledge of the
range of the various forms of Calochorti, but it is still far
from complete. In the notes on distribution of species, the
range as I have accurate knowledge of it is given. A
species may, and in many cases doubtlessly does, extend
over a far wider range than that with which it is credited.

The measurements of any portion of a plant as given in
published descriptions of Calochorti are of little value, and
are apt to be misleading. Environment makes the greatest
difference in the size of the plants. Take, for instance,
those of the woods, such as C’. albus, C. pulchellus, and the
elegans group; the variations are almost limitless. Especially
after a forest fire is growth luxuriant. Plants which under
adverse conditions have leaves but a few inches in length,
and few-flowered, slender stems, will, under more favorable
circumstances, produce great leaves a foot or two long, stout
stems eight inches to two feetin height, and a dozenor more
fine flowers. Especially do the plants of the Mariposa sec-
tion respond prolifically under fertile conditions. If the
season be dry, the plants are sparsely scattered and but a
few inches above ground; but let the season be one of great
rainfall, they fairly hide the ground with tall, many flowered
stems upholding numerous large blossoms. But while
measurements based upon a series of specimens are almost
valueless, proportional measurements of the parts of the
same flower are often of great importance, the proportions
between the parts being usually the same whatever the size
of the flowers.

I1I2 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

In the present paper I have departed from the usual
method by describing a single type plant and indicating the
variations in the notes. Personal experience has proven
that the fuller a description is, the more value it has in the
determination of specimens; details seemingly of little im-
portance at the time when the specimen was described, are
often indispensable in the work of later students, making it
an absolute necessity for them to refer to the type specimen
in order to determine what was really described. This
brevity in some of the earlier descriptions makes them
entirely valueless for determination.

Having no personal knowledge of the Mexican species
outside of the herbarium, I have not included them in this
revision, being unable to add anything of value. There is
a large and almost unknown field in Arizona and western
New Mexico which will probably yield several new species.

Only the description and when possible the locality (the
original locality is quoted in nearly every instance) of the
species have been given in the following pages, the
synonymy having been omitted because of its often
doubtful nature.

Kry TO THE SPECIES OF CALOCHORTUS.
; Section |. Eucalochortus.

Flowers or fruit nodding; petals incurved or strongly arched; gland trans-
versely crested or hairy; capsule nodding, with thin acute or winged cells;
leaves long and glossy, not channeled.

Group 1. GLOBE TULIPS.
Type of Group C. albus.
Flowers subglobose, nodding. Woodland plants; California.

Flowers white; petals covered with scattered silky hairs within.
1. C. albus.
Flowers rose color; petals silky within, partly opening out. Foot-
hills of Fresno and Tulare counties, California..2. C. ameanus.
Flowers light yellow; petals silky within, gland bordered with stiff
hairs which cross each other. ...............: 3. C. pulchellus.
Petals very strongly inarched, not silky within, but margin thickly
set with short, stiff hairs; gland like last........ 4. C. amadbtlis.

BorT.—VOtz, II.] PURDY — CALOCHORTUS. LE Qs

Group 2. STAR TuLips.
Type of Group C. elegans.

Flowers campanulate, erect or ascending; capsule nodding (except in No.
13); stem low and flexuous (In 14, 15, 16, stout and erect), not bulbiferous or
very seldom so.

“Petals covered with hairs, and with a transverse scale covering upper part
of the gland. Woodland plants.

Flowers yellow. Foothills of the Sierra Nevada... 5: C. benthamt.

Flowers white or purplish blue, covered with long erect hairs; cap-

sule oblong-elliptical; stem branching........6. C. maweanus.

6a. C. maweanus var. major.

66. C. maweanus var. roseus.

Flowers blue, covered with silky hairs, longer and slenderer than

the last; capsule orbicular; inflorescence umbellate.

7. C. ceruleus.

Flowers greenish white; petals with very narrow scale and covered
with long hairs. Oregon and north..8. C elegans.

8a. C. elegans var. nanus.

Flowers yellow, green tinged; petals strongly inarched and pit

deeply set. Mt. Jefferson, Oregon............... 9. C. lobbi.

“Petals with a transverse scale closely appressed over upper portion of
gland, nude or nearly so. Woodland plants; in dry soil.

Flowers white with a single tuft of a few hairs at each end of scale
on petals; plants very low and slender. Sierra Nevada.

Io. C. nudus.

Flowers white with scant hairs on lower third; plants taller than the

last. Vicinity of San Francisco Bay........ 11. C. umbellatus.

“Petals nude or only lower portion hairy; flowers campanulate; plants
growing in open wet meadows.

Flowers lilac, hairy on lower third; one or several bulblets on stem
belowthe sroundl 4.00) ke ee ee, 12. C. unifiorus.
Flowers white, not bulbiferous; capsules erect... 13. C. shastensis.

"Petals covered with silky hairs; flowers and stems stout and erect.
Closely related to Group 1 of Mariposa. Plants growing in open
jields or hillsides. Mt. Shasta, California, and north.

Flowers blue; petals without scale, covered with long silky hairs.
Mt. Shasta, California (?), and Willamette Valley, Oregon.

14. C. folmtet.

Flowers white; petals with scale, otherwise the same as the last.

Willamette Valley, Oregon .............. 2.005. 15. C. purdyt.

Flowers straw color; petals without scale, otherwise like C

tolmiet. Lake Pend d. Oreille, Idaho ...... 16. C. apiculatus.

114 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

SecTION II. Mariposa.

Flowers open-campanulate; gland usually densely hairy; capsule and
pedicels erect.
Group I. OREGON MARIPOSAS.

Type of Group C. nitidus.

Capsule as in Section I, but erect; leaf (as in Section I) long and glossy,
not channeled. Oregon and northeast.

* Petals with an indigo blotch in the center.
Plowers larse, white'to lavender «2. ..0G.0% seed oe 17. C. nitidus.

**Petals not spotted in the middle; flowers lilac, smaller than in the

preceding.
Flowers strongly arched and barred with yellow..... 18. C. greene.
Flowers less arched; stem bracted midway...... 19. C. pavonaceus.
Similar to last; stem not often bracted midway. 20. C.longebarbatus.
Flowers white, densely hairy above gland.......... 21. C. howellit.

Group 2. Rocky MOUNTAIN MARIPOSA.
Type of Group C. gunnisoni.

Gland transverse and narrow; leaf usually as in Section I. East
of the Rocky Mountains... 2.035052 5-608 22. C. gunnisonti.

Group 3. WEED’s MARIPOSA.
Type of Group C. weediz.

Petals covered with slender hairs; capsule narrowly oblong, with thick,
obtusely angled cells; radical leaf as in Section I, solitary, long, shining, and
not channeled; bulb heavily coated with coarse black fiber.

Flowers orange or rarely pink or white.........-.... 23. C. weedit.
Flowers purple...23@. C. weedii var. purpurascens (C. plummere).

Petals brownish, short, truncate, not equaling sepals.
236. C. weedti var. vestus.

23c. C. weedit var. obispoénsis.

Group 4. GOLDEN BowL MArIposas.
Type of Group C. clavatus.

Petals yellow, lower half covered with clavate hairs; radical leaves linear
and deeply channeled.
Stem stifly ziezae.. f../s. sock Ae cies stream @ Soe suinen 24. C. clavatus.

Stem not zigzag. Doubtfully placed in this group, but has no cla-
vate hairs...25. C. concolor, sp. nov. (C. luteus var. concolor ).

Bot.—VOL. II.] PURDY— CALOCHORTUS. II5

Group 5.
Type of Group C. kennedy.

Petals nearly naked; gland round, small, and densely hairy with matted
hairs; leaves ashy blue, linear, deeply channeled. Desert plants.

Petalsivermilionton Oranees. esac cities) eee lee os 26. C. kennedy.
Flowers clear yellow ; petals densely hairy below; capsule narrowly
ODOM ee eal sie etsoterotinnerbaiatie cementite. aes 27. C. aureus.

Group 6. BUTTERFLY TULIPS.
Type of Group C. venustus.

Petals slightly hairy below, usually oculated and brilliantly colored; gland
prominent, round or lunate; leaves linear, channeled. California.

Flowers yellow; petals not oculated; gland lunate; capsule attenuate
from a broad base; plant dwarfed..............% 28. C. luteus.
Flowers yellow or lemon, otherwise same as in var. oculatus.
28a. C. luteus var. citrinus.
Flowers white, yellow or lilac; petals oculated, gland lunate.
286. C. luteus var. oculatus.
28c. C. luteus var. robusta.
Flowers lilac or white; gland narrow, doubly lunate...29. C. vesta.
Flowers white, cream, lilac, purple, red or pink; petals oculated,
in some varieties with a red blotch above eye; gland round;
capsule: linear.) .i: st) <s4) ois.cts 30. C. venustus.
30a. C. venustus var. roseus.
306. C. venustus var. eldorado.
30c. C. venustus var. purpurascens.
30d. C. venustus var. sulphureus.

Group 7. Litac MAriposas.

Type of Group C. splendens.

Petals white, lilac, or purplish, not oculated, more or less hairy; gland
small, round, and densely hairy; leaves linear, channeled.

Flowers lilac-purple; petals hairy on lower third. 31. C. splendens.
Flowers lilac to salmon, short yellow hairs on lower third of petal;

plants low and slender....... 31a. C. splendens var. montanus.
Flowers large; petals pale lilac with cobwebby hairs on middle
EES Vey, ies steal oie neces (etapa thao ee 316. C. splendens var. major.

Much larger and stronger than type; hairy on lower third of petal.
31c. C. splendens var. rubra.

Flowers white; gland ill defined; plants more slender than last.

MOlOLAMOCDESETED iin ance wc Uelve comets wlaa abies 32. C. palmer.
Flowers white or lilac with dark maroon spot at base of petal; cap-
sule oblong. Resembles C. splendens........ 33. C. cataline.

Flowers smoky white; stems stout and umbellate.
34. C. invenustus.

Flowers similar to last; petals deeply pitted...... 35. C. excavatus.

116 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

Flowers purple; sepals obtuse; stem flexuous and weak, almost
PLCC MINE Pate jenna pe te roles wee erie kos 2se we 36. C. flexuosus.
Flowers white. Resembles C. splendens........... 37. C. dunnit.

Group 8. GREEN BANDED MARIPOSA.
Type of Group C. macrocarpus.

Petals purplish lilac, with a greenish line down the back, obovate-
acuminate.

Stems stout and rigid; leaves linear and deeply channeled. North-
eastern California to eastern Washington and southern Idaho.
38. C. macrocarpus.

Group 9. SEGo LILIEs.

Type of Group C. xuttallit.

Petals white, lilac, yellow, or pink; gland round; stem prominently
bulbiferous at base, umbellate.

Flowers as described above. Stout desert plants of the Great Basin

ANG SaAStwaradly i. 235.0. %soc ee taken s seater eee 39. C. nuttallit.
Flowers smaller, usually white; anthers sagittate. Slender Alpine
plants; = SierrayNevadaceat 2.2 eee ere aac 40. C. leichthiniti.

DESCRIPTION OF SPECIES.

Calochortus.

Perianth deciduous, of six distinct, more or less concave segments, the
three outer (sepals) greenish and more or less sepaloid, the inner (petals)
mostly broadly cuneate-obovate, usually with a conspicuous glandular pit
near the base, and variously colored. Stamens six, on the base of the seg-
ments, included; anthers linear to oblong, basifixed, dehiscent laterally.
Ovary sessile, triquetrous and three-celled, many ovuled; stigmas sessile,
recurved, persistent; capsule elliptical to lanceolate, membranous, three-
angled or three-winged, mostly septicidally dehiscent; seeds numerous, two
rows in each cell, somewhat flattened, with a thin membranous white or
brownish, often loose, testa. Stems usually flexuous and branching from mem-
branous or but fibrous-coated corms; leaves few, linear-lanceolate, radical and
cauline, the latter alternate and clasping; all with many nerves and transverse
veinlets. Flowers one to twenty, showy, terminal, paniculate or umbellate.

The above generic description is in greater part that of
Sereno Watson as given in the ‘‘Botany of California.”’
The genus is confined to western America, from Nebraska
to the Pacific Ocean, and from northern Mexico to British
America.
Section I. ucalochortus.

Flowers or fruit more or less nodding; petals strongly incurved or arched,
with a broad, transversely crested or more or less hairy pit above the base;

BoT.—VOL. II.] PURDY — CALOCHORTUS. 7

sepals naked, rarely spotted; capsule elliptical or broadly oblong, deeply tri-
quetrous and septicidal, the thin compressed lobes acute or winged; seeds
ascending, close and pitted, the testa mostly brownish.

Group 1. G.iose TuLips.
Flowers subglobose, nodding. Woodland plants; California.
1. Calochortus albus Dougl.

Calochortus albus DouGLas 7 Litt.
Cyclobothra alba BENTHAM, Trans. Hort. Soc., N.S., Vol. I, p. 413, Pl. XIV,
fig. 3; Bot. Register, Vol. XX, 1835, Tab. 1661.

Stem stout, glaucous, usually branching, a foot or two high; radical leaves
a foot or two long, 8-12 lines wide, lanceolate-acuminate; bracts large and
foliaceous, I-5 inches long, 4-8 lines wide; sepals shorter than petals,
Ovate-acuminate, greenish white; petals pure white, purplish at base, ovate-
orbicular, acutish, 12-15 lines long, with scattering long silky hairs above
the gland; gland lunate, shallow, with four transverse imbricated scales,
fringed with close short yellow or white glandular hairs; anthers oblong-
obtuse, mucronate; ovary attenuate above; capsule 1 or 2 inches long, 6-12
lines broad, abruptly short-beaked; seeds brown, pitted.

The original specimens are in all probability from Mon-
terey, as Douglas visited there, where the species is plentiful.

C’. albus is found in the Coast Range of California, from
San Francisco Bay south to San Diego County, and in the
Sierra Nevada Mountains, from Butte County south to San
Diego County.

There is quite a difference between the plants of the
Coast Range and those of the Sierra Nevada. The flowers
of the former are darker in color, often tinged with rose,
and with petals thicker, more strongly inarched. The petals
never open out sufficiently to show the inside of the flower,
which after being in bloom a few days is half opened.
Calochortus amenus Greene is really a color form of this
Sierran form of Calochortus albus; but owing to the fact that
the original locality of C’. albus is unknown, the writer hesi-
tates to erect either the form from the Coast Range or that
from the Sierra Nevada into a new species.

Several variations of the form of the Coast Range have
been described, some as species and some as varieties; but
I fail to discover any characters by which they may readily
be recognized.

118 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.
2. Calochortus amenus Greene.

Calochortus ameenus GREENE, Pittonia, Vol. II, 1890, p. 71.

Similar to C. albus, but lower and more slender; petals a rosy pink with the
gland rose-purple, scarcely at all arched, and opening in full bloom.

Found in the higher foot-hill region of Fresno and Tulare
counties, California.
‘* Mountains east of Visalia, California.’’

3. Calochortus pulchellus Doug.

Calochortus pulchellus DouGuas in litt.
Cyclobothra pulchella BENTHAM, Trans. Hort. Soc., N. S., Vol. I, 1835, p.
415, Pl. XIV, fig. 1; Bot. Register, Vol. XX, 1835, Tab. 1662.

Stem stout, glaucous, usually branching, 8 to 16 inches high; radical leaves
a foot long, 6 to 12 lines wide, lanceolate-acuminate; bracts large and folia-
ceous, 2 to 3 inches long on the same plant, 4 to 6 lines wide; sepals shorter
than petals, ovate-acuminate, yellow tinged with brown on the back; petals
canary-yellow, ovate with the base cuneate, obtuse at apex, 9-12 lines long,
with scattering long silky hairs above the gland, and bordered with short stiff
hairs; gland deep, protruding outwardly, bordered with stiff hairs which
cross each other; anthers oblong-obtuse; ovary elliptical to elliptical-orbic-
ular, abruptly short-beaked.

The original specimens of this species were collected by
Douglas prior to 1835. The exact locality is not given;
but the only place in which the species has since been
found is in the Mount Diablo region, a section which was
easily accessible at the time of Douglas’ visit to California,
and often visited by his Mexican-Californian hosts.

Although in a region much visited by botanists since
then, no specimens were to be found in any of the herba-
riums of this State up to the year 1897, when C. pulchellus
was collected by Miss Alice Eastwood of the California
Academy of Sciences.

The very name Calochortus pulchellus had been appropri-
ated by another yellow-flowered species of the same group,
which is described and named below as C. amadzlis, and
which is clearly a distinct species.

Following are the original descriptions of the species
copied from the Botanical Register :—

BoT.— VOL. II.] PURDY— CALOCHORTUS. IIg

Cyclobothra pulchella.—‘‘Umbellis 2-3 floris, pedunculis bracteis breviori-
bus, floribus globosis, petalis ovatis obtusis serrulato-fimbriatis fovea valdé
excavatd extus callosd4, sepalis ovato-lanceolatis acuminatis vix brevioribus.”’

Calochortus pulchelius.—‘‘Caulis erectus, teres, glaber, subcorymbosus,
apice magis ramosus quam in precedente, et humilior. Folia plana, acumi-
nata, minus glauca; superioribus brevioribus. Pedunculi bracteis foliaceis
breviores, bini ternive. Flores globosi, minores quam in przecedente, lutei.
Sepala virescentia, viridi-striata, petalis paululum breviora, acutissima. _ Pet-
ala ovata, barbata, fimbriata, basi glabra: fovea nectarifera pilis abscondita.”’

4. Calochortus amabilis, sp. nov.

Stems stout, usually branching in pairs, 8 to 12 inches high, glaucous;
radical leaves to inches long, 4 to 6 lines wide, lanceolate-acuminate, tinged
with purple; bracts large and foliaceous, 2 to 3 inches long, 4 to 6 lines wide;
sepals shorter than petals, ovate, shortly cuneate at base, sharply acuminate
or even mucronate at apex, yellow tinged with brown on the back; petals clear
yellow, ovate, with a short claw, obtuse at apex, naked but margined with a
close row of short stiff hairs, very strongly inarched so that the tips of the
petals overlap each other much like a child’s pin-wheel; gland very deep,
projecting upwards and outwards like a knob, lined with short stiff hairs
which cross each other; anthers oblong-obtuse; ovary elliptical, short-beaked.

C. amabilés is found on the hills along the north side of
San Francisco Bay, from the redwood belt to the Sacra-
mento foot-hills, as far north as Burnt Ranch, Trinity
County, California.

The species has been distributed in large numbers among
the flower-growers of the world as Calochortus pulchellus,
which it resembles in habit and size. The latter is more
closely allied to C. albus, having the same globular flower
and petals silky-haired within. It is also of a much lighter
_ shade of yellow, and never could be confused with C. ama-
belts by anyone who had seen both.

Group-2:) STAR TuLies.

Flowers campanulate, erect or ascending; capsule nodding (except in No.
13); stem low and flexuous (in 14, 15, 16, stout and erect); not bulbiferous or
very seldom so.

“Petals covered with hairs, and with a transverse scale covering upper part
of the gland. Woodlard plants.

I20 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

5. Calochortus benthami Aaker.

Calochortus benthami BAKER, Journ. Linn. Soc. (Bot.), Vol. XIV, 1875, p.
304; also Bot. Mag., Pl. 6475.

Stems several-flowered, very flexuous, branching, dividing into pairs 7 inches
high; radical leaf linear-lanceolate, shining, 4 lines wide, exceeding stem;
bracts ovate-lanceolate, long, acuminate, 2 to 4 inches long, 3 to 5 lines wide;
sepals 8 to 1o lines long, narrowly ovate, mucronate; petals yellow, with a
yellow claw, naked, obovate, rounded above, a little longer than sepals, the
upper portion of the gland covered by a narrow crescentic scale which is
densely bordered above with short yellow hairs, some of which are clavate;
anthers lanceolate-acute, capsule nodding, nearly orbicular, 6 to 9 lines long.

Found in the lower Yellow-Pine belt of the Sierra
Nevada, from Mariposa to Butte counties.

‘¢ California.”’

The description is drawn from rather an extreme speci-
men. The plants are often very slender, simple, and but
three to four inches in height. The flowers vary but little.
In some sections flowers with the claw dark red or nearly
black are common (C.. wadlacez Hort.).

6. Calochortus maweanus Lezchilin.

Calochortus maweanus LEICHTL. ex BAKER, in Journ. Linn. Soc. (Bot.), Vol.
XIV, 1875, p. 305; also Bot. Mag., Pl. 5976.

Resembles the preceding; stem usually branching, very slender and flexu-
ous, 3-10 inches high, with from a few to ten flowers; leaves glaucous,
much exceeding stem, 2-6 lines wide; bracts lanceolate, narrow, Io lines
or more long; sepals ovate-lanceolate, acute or acuminate; petals a little
longer, broadly ovate, acute; gland covered above with a narrow transverse
scale, immediately above the scale densely hairy, the entire surface thickly
bearded with long, erect, white or bluish hairs; anthers lanceolate-acuminate,
2-3 lines long; capsule oblong-elliptic.

This is the type which is found in the Coast Range, from
San Francisco Bay, at least as far north as Trinity County,
California, and western Oregon.

‘¢California.”’

6a. C. maweanus var. major, var. nov. This variety is twice as large as
the type, from which it differs in its much stronger habit and lighter color.

BotT.— VOL. II.] PURDY— CALOCHORTUS. I2I

Grows in the Yellow-Pine belt, Butte County, California.

66. C. maweanus var. roseus, var. nov. The flowers of var. roseus are
tinged with rose; the bulb is distinctive, having a smooth, mahogany-colored
coat.

Its habitat is western Oregon.

7. Calochortus ceruleus Watson.

Calochortus ceruleus Watson, Proc. Amer. Acad., Vol. XIV, 1879, p. 263.

In general resembling a small specimen of C. maweanus. The plants are
very slender; leaves and bracts narrower, pedicels more slender; flowers
almost always in an umbel, petals more rhombic in outline, claw more slen-
der, scale broader and fringed, the remainder of petal densely covered with
long slender silky hairs; anthers oblong-obtuse; capsule orbicular, not
beaked, 6 lines long.

Specimens of this species show but little variation.
‘‘California (in the Sierra Nevada, Placer to Plumas
counties).”’

8. Calochortus elegans Pursh.

Calochortus elegans Pursu, Fl. Am. Sept., Vol. I., 1816, p. 240.

Scape very slender, 4-8 inches high; leaves lanceolate-acuminate, nar-
row, exceeding scape; flowers one to four in umbel, bracts one-half length of
pedicels, acuminate from a base 2 lines wide; sepals ovate-acute, greenish
white without, lighter within, purplish at base; sepals obovate-obtuse, whitish
or tinged slightly with green, with purple spot on claw, covered thickly with
rather short soft hairs, which are white on upper and purple on lower por-
tion, excepting that the margin and a band around upper portion of petal is
naked; scale narrow, ascending, and deeply fringed, covering about one-third
the width of claw; anthers long, acuminate; capsule elliptical, rounded at
each end.

The type specimens were collected by the Lewis and
Clark expedition on the headwaters of the Kooskoosky in
Idaho (?).

The writer was never able to obtain specimens of this
species until just as the present paper was going to press,
when flowers which are unquestionably the true C. elegans
were received from a collector in western Idaho, near
Spokane. As the original description of the species is very
brief, the fuller description, as given above, was drawn

122 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

from these specimens. There is much difference in general
appearance between the species and var. zanus, and in my
opinion the variety will eventually follow the other forms
which were originally included under C. elegans, and be
given specific rank. For the present, however, it is left
under the varietal name. No more definite locality than
western Idaho can be given for the type.

Some Californian specimens have been referred to C.
elegans but the writer has never seen any which come under
the species and does not believe that either the type or any
of its variations are found in California. The Californian
plants referred to as C. elegans are C’. nudus.

Sa. C. elegans var. nanus Woop. (Proc. Phil. Acad., 1868, p. 168.)
Dwarf and very slender, leaves very narrow; petals more hairy and ciliate,
often acute or even acuminate.—C. lyallii Baker, Journ. Linn. Soc. (Bot.),
Vol. XIV, 1875, p. 305.

‘‘High hills, Yreka [California]. Also on Mt. Hood
[Oregon].”’

The description is that of Watson in the Botany of
California.

The variety is known to the writer as it grows on Mt.
Hood and on Mt. Adams. Watson evidently had small
specimens, as under favorable conditions it is quite as
strong as any of the preceding. The color is a delicate
cream.

g. Calochortus lobbii, sp. nov.

Calochortus elegans var. lobbii BAKER, Journ. Linn. Soc. (Bot.), Vol. XIV,
1875, P. 305.

Stem 3 to 5 inches high, not so slender as the preceding; leaf a little exceed-
ing stem in length, 3 to 5 lines wide, lanceolate, abruptly acute; sepals ovate-
lanceolate, acute, greenish with a dark spot below, 6 to 8 lines long; petals a
little longer, white tinged with green, broadly rhombic-ovate, very deeply
pitted, the pit showing as a prominent knob on back of petal; scale very
narrow, deeply bordered with long, feathered, hairy fringes, and concealed in
the recess of pit; above the scale there is a nectar-producing gland covered
by a broad band of agglutinated feathered hairs, above this band lower half
of petal densely hairy with silky hairs, upper half less densely hairy; fila-
ments subulate; anthers oblong-acuminate, ending in a hook-like cusp; cap-
sule narrowly beaked.

Botr.—VOL. II.] PURDY— CALOCHORTUS. 123

So far found only on Mt. Jefferson, Oregon.

“Oreson:, 7

The above was identified for the writer by Baker. As
the other varieties which Baker mentions in his ‘‘Tulipex”’
have been erected into species, and this is more distinct in
character than any of them, it seems proper to raise it to
specific rank.

““Petals with a transverse scale closely appressed over upper portion of
land, nude or nearly so. Woodland plants; in dry soil.

10. Calochortus nudus Watson.

Calochortus nudus Watson, Proc. Amer. Acad., Vol. XIV, 1879, p. 263.
Calochortus elegans var. subclavatus BAKER, Journ. Linn. Soc. (Bot.), Vol.
XIV, 1875, p. 305.

Low and slender, scape 2 to 4 inches high, with a single leaf 3 to 6 inches
long, 3 to 6 lines wide, light green, of even width for most of length, abruptly
acute; flowers one or more, in all specimens examined in an umbel if more
than one; sepals narrowly oblong-ovate, acute, shorter than petals; petals
greenish white or lilac, greenish at base, obovate, somewhat acute, denticu-
late above, 5 to 7 lines long, the same in width, entirely nude except for a
tuft of two or three short stiff hairs at each extremity of the narrow, closely
appressed scale which covers the upper margin of gland; anthers blue,
oblong, two-thirds the length of the subulate filaments.

Probably C. elegans var. subclavatus of Baker.

On north sides of high mountains in the pine forests of
the Sierra Nevada, from Tulare to Plumas counties; in
loose dry soils.

‘‘California (in the Sierra Nevada, Yosemite Valley to
Plumas County).”’

The type as described from Tulare County is white, but
there seem to be variations tending to lilac, and in some
sections a nude petal. This is the smallest flowered of all
the Calochorti.

Ir. Calochortus umbellatus Wood.

Calochortus umbellatus Woop, Proc. Acad. Nat. Sci. Phila., 1868, p- 168.
Cyclobothra elegans var. Torr. ?, Pac. R. R. Rept., 1856, Vol. IV, p. 146.
Calochortus collinus LEMMON, Erythez, Vol. III, 1895, p- 49.
Stem low and branching, 3-15 inches high, flexuous; radical leaf exceed-
ing stem, narrow, 3-4 lines wide; bracts foliaceous, acuminate, 1-4 lines
(2) December 3, Igor.

124 CALIFORNIA ACADEMY OF SCIENCES. ‘[Proc. 3D SER.

long, 3-4 lines broad at base; flowers five to ten; sepals greenish white,
ovate-lanceolate, acute; petals white, broadly fan-shaped, denticulated,
obtuse; scale triangular, ascending, appressed on upper portion of gland,
which has many short white hairs just above it, remainder of petal nude; fil-
aments slender; anthers 1 line long, oblong-obtuse; capsule short, oblong,
6 lines wide, 8 lines long.

A native of California, found on the lower mountains
and hills back of Oakland, Berkeley, and Mills Seminary,
and on Mt. Tamalpais.

‘‘Oakland, California.’’

The species was long confused with C. unzflorus, and was
described as C. collinus by J.G. Lemmon. Professor E. L.
Greene called the writer’s attention to the earlier description
of Wood.

C. umbellatus can be confused only with C. unzflorus,
from which its lack of bulblets and the situation in which it
grows, as well as its color, easily distinguish it. It varies
but little.

*** Petals nude or only lower portion hairy; flowers campanulate; plants
growing in open wet meadows.

12. Calochortus uniflorus Hook. & Arn.

Calochortus uniflorus Hook. & ARN., Bot. Beech. Voy. Suppl., 1841, p. 398,
Tab. XCIV.

Stem low, flexuous, but often stout, usually branched, 4-8 inches high,
with one to four bulblets below the surface; radical leaves broad, 4-6 lines
wide, exceeding stem; bracts linear-lanceolate, long and conspicuous;
flowers four to ten, in one to three umbels, on long flexuous pedicels 3-10
inches long; sepals ovate, lanceolate-acuminate, greenish lilac; petals cuneate,
somewhat truncate, denticulate, 1o-12 lines long, color lilac, often with a
purple spot on each side of the scale, naked above, sparingly hairy imme-
diately above the gland; gland shallow, not pitted, a narrow triangular scale
appressed upward over upper center; filaments slender; anthers obovate-
obtuse, lilac, 2 lines long, one-half length of filaments; capsule elliptical.

Except in the spots on petal or sepal there are few color variations.

From Monterey, California, northward in the Coast
Range to Grant’s Pass, Oregon. Found in wet meadows.
C. uniflorus was originally described by Hooker and Arn-
ott from specimens collected on the Beechy Expedition.
The specimens were without doubt collected either in the

Bor.—VOL. II.] PURDY — CALOCHORTUS. 125

vicinity of San Francisco or at Monterey. A good cut
accompanies the description.

C. lilacinus was first described by Kellogg from specimens
collected near Calistoga, Napa County, California. Dr.
Kellogg’s original description, together with a water color
drawing, are in the Herbarium of the California Academy
of Sciences.

After carefully comparing all the data contained in the
descriptions of C’. unzflorus and C. dzlacinus with both fresh
and herbarium specimens from Monterey, Calistoga, and a
number of other localities, the writer is thoroughly convinced
that the two species are the same. The description of C.
unifiorus was published much earlier than that of C. /zla-
cinus; it is drawn from a fair specimen of the species.

It is obvious that Dr. Watson confused C.. /elacinus with
the quite different C’. wmdellatus.

The writer has seen C. wuzflorus in meadows high in the
mountains where the plants grew very low and slender and
were only one- to three-flowered. In rich ground several
bulblets are produced annually, and if left undisturbed large
and dense masses are formed, sometimes hundreds of them
to the square foot. In soft ground the stems are apt to run
along close to the surface a few inches to a foot before
coming through, and in these situations plants a foot high
with pedicels ten inches long are not uncommon.

13. Calochortus shastensis, sp. nov.

Scape low, slender, 4 to to inches high, but unusually erect, with a single
shining light green radical leaf 3 to 6 inches long, of almost uniform width
(3 to 6 lines), but abruptly acute at apex; bracts lanceolate, 6 lines long;
sepals long, ovate, acute and acuminate, greenish without, lighter within,
purple spotted near base; petals white or lilac, broadly fan-shaped, somewhat
truncated above, denticulate, naked except that some few specimens have a
few hairs above the narrow, fringed, ascending scale which divides the gland;
anthers linear, obtuse, slightly sagittate; capsule as in preceding but erect.

Found in open moist meadows in the vicinity of Sissons,
California, at the base of Mt. Shasta, and about springy
places on the western flank of the mountain.

C. shastensts has long been known and collected as C.
nudus, which it closely resembles in flower but from which

126 CALIFORNIA ACADEMY OF SCIENCES. [PRroc. 3D SER.

it is clearly distinguished by the erect capsule. Itis a curious
fact that a species linking the small Calochorti of the wet
lands with the C.. mztzdus section should be found at the very
point where the latter terminates its most southern extension.

The true C. xudus, it will be noted, grows only on dry
slopes in the Sierra Nevada, from Plumas County, Cali-
fornia, southward.

GIANT STAR TULIPS.

**** Petals covered with silky hairs; flowers and stem stout and erect.
Closely related to Group 1 of Mariposa. Plants growing in open
fields or hill-sides. Mt. Shasta, California, and north.

14. Calochortus tolmiei Hook. & Arn.

Calochortus tolmiet Hoox. & ArN., Bot. Beech. Voy. Suppl., 1841, p. 398.

Stem stout, erect, usually branched, 9 to 18 inches high; leaves 4 to 6 lines
broad, not greatly exceeding the stem, bracts foliaceous; petals 9 to 15 lines
Jong, very broadly obovate and scarcely acute, rather deeply pitted, covered
and fringed with long purple and white hairs; gland without scale, but the
upper circular edge with a dense fringe of reflexed hairs; anthers lanceolate
and acuminate, 2 to 3 lines long; capsule broadly elliptical, acutish at each
end, Io to 15 lines long.

C. tolmiez is frequently confused with C. purdyz. The
writer has seen specimens of the true C. tol/mzez from the
hills west of the Willamette River in Oregon. Dr. Watson
also refers specimens from Mt. Shasta, California, and Mt.
Adams, Washington, to this species. The original locality
is ‘‘ Banks of the Walamet River.”’

15. Calochortus purdyi Zastwood.
PLATE XV.

Calochortus purdyi EAstwoop, Proc. Cal. Acad. Sci., 3d Ser. (Bot.), Vol. I,
p. 137, Pl. XI, figs. 8a-8f.

Glabrous and glaucous; stem 2 to 3 dm. [18 to 16 inches] high, rather
stout, erect, branching, two to many-flowered, not bulbiferous at base; radical
leaf solitary, sheathing the stem, linear-lanceolate, acuminate, 2 dm. [8 inches]
long, 1 cm. [6 lines] wide, the upper surface bright green, the lower glaucous
and ribbed with the filiform nerves; bracts foliaceous, janceolate-acuminate,
amplexicaul, upper ones opposite; pedicels equalling or slightly surpassing

yh

Bot.—VOL. II.] PURDY— CALOCHORTUS. 127

the bracts, erect in flower, recurved in fruit; flowers broadly open-campanu-
late; sepals from elliptical to narrowly ovate, abruptly acuminate, tinged with
purple on the outer surface, purple-veined on the inner, two-thirds as long as
the petals; petals broadly obovate-cuneate, acute or rounded at apex, creamy
white or tinged with purple, bearded all over the inner surface with long hairs
which are white on the upper half of the petals, purple on the lower, some-
what arched by the narrow, transverse, semicircular, conspicuous gland, the
shallow pit of which is covered by a densely hairy narrow scale; anthers lan-
ceolate, abruptly acuminate, cream color or purplish, shorter than the fila-
ments, which broaden to the base; capsule 3 cm. [one-fourth inch] long,
2 cm. [ro lines] wide, broadly elliptical, with the three wing-like valves trans-
versely veined.

The above is the original description as given by Miss
Eastwood.

‘‘Willamette Valley [Oregon], in the foot-hills.”’

C. purdyt may be distinguished from C. ¢olmzez by its
color and the absence of the gland. The two are fre-
quently confused and many of the specimens in herbariums
labeled C. tolmzez are in reality C. purdyz.

Through C. howelliz, C. tolmiez and C. purdyz are closely
related to C. longebarbatus; while the large forms of C.

maweanus link them to the e/egans group.

16. Calochortus apiculatus Baker.

Calochortus apiculatus BAKER, Journ. Linn. Soc. (Bot.), Vol. XIV, 1875,
P- 305.

Stem stout, erect, a foot to 18 inches high; the single radical leaf 6-12
inches long, 3-9 lines wide; bracts linear, acuminate, I-3 inches long;
sepals lanceolate, greenish white, acute, 6-9 lines long; petals straw
colored, broadly obovate, one inch long, distinctly pitted, above with only
scattering hairs, pit densely hairy and without scale; anthers 4 lines long,
acuminate; filaments as long; capsule 12-15 lines long, narrowly oblong.

‘‘Columbia brittanica ad montes Pend Oreille et Koot-
enay.”’

The above is drawn from Baker’s original description
and from letters. The writer has no personal knowledge

of the species.

SecTion II. Mariposa.

Flowers and fruit erect on stout pedicels; flowers open-campanulate; gland
usually densely hairy; capsule (except in Group 1) narrow, with thick lobes,

128 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

septicidal; sepals often hairy or subglandular, or spotted within; seeds
ascending and somewhat turgid, with white, loose and spongy, minutely
tesselated testa.

Group 1. Orrcon Mariposas.

Capsule with thin, acute or winged cells (as in Section I, but erect), also
resembling Section I in the solitary, flat, shining leaf. Closely linked to the
preceding group of Eucalochorti (Giant Star Tulips). Oregon and to the
northeast.

* Petals with an indigo blotch in the center.

17. Calochortus nitidus Doug.

Calochortus nitidus DouG.as, Trans. Hort. Soc., Vol. VII, 1830, p. 277,
Tab. IX, fig. a.

Stem bulbiferous near the base, slender, stiffly erect, not bracted in the
middle, bearing an umbel of 2-4 flowers subtended by 2-4 linear bracts; ped-
icels often 3 inches long; sepals ovate-lanceolate, long-acuminate, exceeding
petals; petals white to lavender, a conspicuous indigo spot in the middle,
2 inches long, the same in width, broadly cuneate, rounded above, with a
small rounded gland densely matted with short hairs, and scattering long
hairs around and above; filaments filiform, winged below; anthers linear-
oblong, two-thirds as long as filaments; capsule elliptical-orbicular, strongly
winged and crested.

Described from specimens from Union, Oregon.

‘¢Columbia britannica’’ (Dougl.); ‘‘Oregon’’ (Spaulding).

This species is the largest flowered of the group; it is
found in wet fields, from eastern Oregon through Idaho to
Montana and northeastern Nevada. Specimens from Yel-
lowstone Lake are yellow. The purple blotch on the petal
seems to be distinctive.

** Petals not spotted in the middle; flowers lilac, smaller than in the preceding.

18. Calochortus greenei Watson.

Calochortus greenei Watson, Proc. Amer. Acad., Vol. XIV, 1879, p. 264.

Stem stout, branching, often a foot high or more, 2-5-flowered; leaf about
equalling the stem, an inch broad; bracts narrow, elongated; sepals greenish
with more or less of lilac within, and with a yellowish hairy spot above the
base; petals broadly fan-shaped and obtuse, 1% to 1% inches long, lilac,
somewhat barred with yellow below, strongly pitted and arched, the lower
part densely covered with very long yellow hairs, upper part of the blade
more thinly hairy, not ciliate; pit densely villous above a broad transverse,
laciniate scale; anthers broad, acute or obtuse, % inch long; capsule an inch
long, 4 to 6 lines broad, attenuate into a stout beak.

Bot.—VOL. II.] PURDY — CALOCHORTUS. 129

“Siskiyou County, California (Greene); Multnomah
County, Oregon (Howell).”’

The description and locality are copied from the original
of Watson, but the latter habitat is incorrect, as Howell’s
specimens are C’. longebarbatus.

While at work on this revision the writer had not access
to specimens of authentic C’. greenez, the examples from
Oregon and Washington referred to this species being in
reality C’. dongebarbatus Watson. Specimens from Modoc
County, California, a little east of the type locality, appear,
however, to be the true C. greenez.

Professor Greene, the discoverer of the species, is of the
opinion, which the writer also shares, that Dr. Watson con-
fused two distinct species in his description. C. greene? is
described as having the erect capsule characteristic of the
section, while Professor Greene reported that some of his
specimens had the nodding capsule of the Eucalochorti.
From the evidence, it would seem that the densely hairy,
strongly arched Calochorius described by Dr. Watson is
properly another one of the Giant Star Tulips; but this
cannot be definitely determined until there is fuller material
to work with.

19. Calochortus pavonaceus Fernald.

Calochortus pavonaceus FERNALD, Bot. Gaz., Vol. XIX, 1894, p. 335.

Stem erect, stiff, slender, a foot or so high, with a single linear bract near
the middle and two or more below the umbel; radical leaf flat, shining, lan-
ceolate, not channeled; flowers 1-4 in an umbel; pedicels 14-3 inches long,
exceeding bracts; sepals purplish, ovate-lanceolate, acuminate; petals 1%
inches long, cuneate-obovate, denticulate, with a broad claw, color lavender
to purple, with a circular band above the small round gland, which is covered
with densely matted yellow hairs; a few hairs on the margin and silky hairs
sparingly scattered over lower third; filaments slender, winged, exceeding
the obovate-obtuse anthers, which are 4-5 lines long; capsule elliptical,
acutely angled and strongly beaked.

The description is from specimens collected near the
type locality, ‘‘Pullman, Whitman Co., Washington.’’

130 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

20. Calochortus longebarbatus Watson.
PLaTE XVI, Fic. 1.

Calochortus longebarbatus WATSON, Proc. Amer. Acad., Vol. XVII, 1882,
p. 381.

Near C. pavonaceus. Stem slender, a foot or so high, bulbiferous at base,
stiffly erect; inflorescence umbellate if more than one flowered, short linear
bracts subtending the umbel or a single flower on a stem; true radical leaf
one, lance-linear, acute, 2 to 3 lines wide, nearly as long as or exceeding
stem, a linear bract-leaf on lower part of stem just above radical leaf; sepals
ovate-lanceolate, acuminate, a little shorter than petals, greenish lilac within;
petals lavender, lighter below, with a dark purple circular band above gland,
obovate-cuneate, denticulate, margin naked; gland small, roundish, covered
with densely matted brown hairs, with some long silky hairs above and
beside it; filaments slender, winged below, 2 to 3 times as long as anthers;
anthers narrowly ovate, obtuse; mature capsule 9g to to lines long, strongly
winged and beaked.

‘In low grassy grounds, Faicon Valley, Klickitat
County, Washington Territory.”’

The description given is of average specimens from the
type locality, and from Hood River, Oregon. From the
description it will be seen that C. longebarbatus closely
resembles C. pavonaceus except that the bracts are near the
base instead of the middle, there are no hairs on the mar-
gin of the petals, and the filaments are somewhat longer.
The two may run into each other, but to definitely ascer-
tain whether this is the case specimens from a much wider
range are needed.

21. Calochortus howellii Watson.

Calochortus howellit Watson, Proc. Amer. Acad., Vol. XXIII, 1888, p. 266.

Stem erect, a foot or more high; radical leaf solitary; cauline leaf narrow
and short; sepals ovate and shortly acuminate; petals white, an inch long,
denticulate, slightly ciliate near the base, covered within with short crisped
hairs, those above the gland denser and dark greenish; gland transversely
oblong, densely covered with short yellow hairs; anthers oblong-acute, and
apiculate, 3 lines long; capsule elliptical, acute, 9 lines long.

‘¢Found near Waldo, Oregon, in 1884, and at Roseburg
in 1607.

BoT.—VOL. II.] PURDY— CALOCHORTUS. 131

The above description is drawn from specimens collected
near the type locality.

Group 2. Rocky Mountain Mariposa.

Gland transverse and narrow, extending across petal from side to side;
leaf usually as in Section I.

22. Calochortus gunnisoni Watson.

Calochortus gunnisont Watson, Bot. King’s Rept. Ex. goth Par., p. 348.

The usually single radical leaf linear-lanceolate, acute, flat, and not chan-
neled; stem bulbiferous at base, a foot or so high, erect; inflorescence usually
umbellate, the umbels 1-4-flowered and subtended by two or more rather
broadly lanceolate, acuminate bracts; one or several cauline leaves below the
umbel; sepals narrowly ovate, acute, with scarious margins, yellowish within,
often marked with dark blue; petals creamy white, often banded with dark
blue above gland, cuneate, with a broad claw, usually rather truncated above,
sometimes slightly rounded, 12-15 lines long; gland narrow, extending trans-
versely nearly from side to side of petal, and covered with short, dense,
glandular clavate hairs; anthers equalling filaments, ovate, cuspidately acute;
capsule cylindric-triangular.

Described from an average sized specimen from Boulder,
Colorado. The species is found on the eastern side of the
Rocky Mountains from Nebraska to New Mexico.

‘‘Rocky Mountains of Colorado. Collected also in Utah
by Gunnison.”’

The color of the petals is variable, flowers from different
localities showing all the gradations from white, through
pink, lilac, purple, and blue, with considerable variety in
the color of the bands marking the lower portion of the
petal; but a greenish tinge and greenish hairs characterize
them all. Some of these color forms have been named.

Group 3. WeEED’s Mariposa.

Petals covered with slender hairs; capsule narrowly oblong, with thick
obtusely angled cells; radical leaf as in C. albus (Section 1), solitary, long,
shining, and not channeled; bulb heavily coated with coarse black fiber.

132 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

23. Calochortus weedii Wood.
PLATE XVII.

Calochortus weedii Woop, Proc. Acad. Nat. Sci., Phila., 1868, p. 169.

Corm deep seated, heavily coated with coarse dark brown fiber; stem 1-3
feet high, stout and flexuous, usually branched, leafy, one to many flowered,
not bulbiferous; radical leaf lanceolate-acuminate (the same as in C. albus),
solitary, broad, shining, flat and not channeled; cauline leaves broad, acumi-
nate; sepals often exceeding petals, narrowly ovate-lanceolate, acuminate,
yellowish within, often with a hairy spot at base, with scarious margins; pet-
als cuneate, denticulate, either rounded or more or less truncate above,
12-15 lines long, orange colored and covered with long silky yellow hairs,
each set in a dark brown spot, upper quarter usually naked; gland small,
circular to oblong, densely matted with short hairs; filaments filiform, exceed-
ing the oblong anthers; capsule narrow, attenuate upward, 1% inches long.

Described from specimens from San Marcos, San Diego
County.

“San Dieso:’

C. weediz in its various forms is found in the Coast
Range of California, from San Luis Obispo County south
to Lower California. It is easily distinguished from other
Mariposa Tulips by the single large radical leaf (one to two
feet long by five to eight lines wide), the very heavy,
coarse, stringy, fibrous coating of the bulb, and the long
silky hairs springing from the brown dots on the petals.

The species is almost as variable as C. venustus, each
variation having its own range, where it is found to the
exclusion of all others.

Beginning at San Diego, we have the typical C. weedzz,
a large orange-colored flower, covered with yellow hairs
and dotted with brown. In this form the petals are usually
full, although occasionally somewhat truncated. Nearly
all of the C. weedzz in San Diego County, whether in the
interior or on the coast, conform to the type.

23a. C. weedii var. purpurascens WATSON, Proc. Amer.
Acad., Vol. XIV, 1879, p. 265. (C. plummere GREENE,
Pittonia, Vol. II, 1890, p. 70). Going north to Los Ange-
les and San Bernardino, we find a broad belt of country
where the flower of C. weedzz is still full petaled, but the
color is lilac or lilac purple. While possessing hardly any

Bot.—VOL. II.] PURDY— CALOCHORTUS. 133

analytical points of difference, it may be said that in cultiva-
tion this form proves to be a stronger and hardier plant than
the type. A curious fact in connection with the bulb is also
worthy of mention. The coarse fibrous coating which the
variety has in common with the type is often found to cover
not one bulb, as is usual, but two large bulbs of about equal
size, laid so flatly face to face that outwardly the bulb
retains the round form of the species, and it is only by
breaking the coating that the presence of the second bulb
is discovered. This habit of the bulb can hardly be called
offsetting, as it is impossible to call either the parent bulb.
In the San Jacinto Mountains this form varies to pale pink
and white. It is the C. weedzz var. albus of horticulture.

A fine plate of this variety may be found in ‘‘The Gar-
den, 7) Heb. 2,795,

236. C. weed? var. vestus. Continuing still further
north to Santa Barbara, we again find the species, but the
petals are much more truncated and curiously fringed with
brown hairs, while the color is reddish brown.

23c. C. weedit var. obispoénsis, var. nov. (C. obispo-
énsis LEMMON, Bot. Gaz., Vol. XI, 1886, p. 180.) This is
an extreme form found in San Luis Obispo County. The
petals of the flowers are still more truncated than in the
preceding, and near the town of San Luis Obispo, which I
think is the northern extreme of its range, the flowers
assume a most fantastic form, the brownish petals being so
much truncated that the sepals far exceed them, and the hairs
which are scattered in typical specimens here seem to be
condensed upon the small remaining surface of the petals.

“‘On dry, stony hills near San Luis Obispo, Cal.”’

It is, of course, impossible to enumerate here all the
many gradations in this plant, which forms one of the most
interesting studies in plant variation.

Group 4. GoLpEN Bow. Mariposas.

Petals yellow, lower half covered with clavate hairs; radical leaves long-
linear and deeply channeled.

134 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

24. Calochortus clavatus Watson.

Calochortus clavatus WATSON, Proc. Amer. Acad., Vol. XIV, 1879, p. 265.

Stem stiff, very stout, strongly zigzag, branching, 1-3 feet high, if few flow-
ered umbellate, if many flowered with a terminal umbel and short laterals at
the axils of the rather large cauline leaves; the single radical leaf glaucous,
linear, deeply channeled, often 1-2 feet long; cauline leaves as broad and
prominent; pedicels stout, 2-5 inches long; sepals acuminate, broadly ovate-
lanceolate, nearly as long as petals, yellowish within, often greenish without,
spotted, with dry scarious margins; petals rather truncate, 18 lines wide, 21
lines long, broadly fan-shaped and strongly arched, with a broad claw, the
gland deeply pitted, with yellow hairs; the side of gland and the lower half
of petal densely hairy with long yellow hairs, each tipped with a knob-like
point, and purplish red at base; color a rich yellow, with claw often reddish
brown and a reddish brown band above the hairy zone; perianth bowl-
shaped (‘‘formed like a broad-based cup’’); filaments slender, a little exceed-
ing the ovate-oblong, obtuse, purplish-brown anthers; capsule narrow,
attenuate above and below, 3 inches long.

Described from medium sized plants.

Found on dry rocky points, usually in volcanic soils. The
species is widely but peculiarly distributed from Newhall
(Los Angeles County) to San Luis Obispo; it is also found
in the Sierra Nevada near Pleasant Valley (El Dorado
County), and on White Rock, an isolated quartz rock-
mass in the plains of Mariposa County. Doubtlessly found
also on lava formations at many intermediate points.

«‘California (San Luis Obispo; J. G. Lemmon, 1878).”’

This Calochortus is the stoutest stemmed, tallest, and
largest flowered of all the Calochorti. The heavy, strongly
zigzag stem, yellow bowl-shaped flowers, and clavate hairs,
are strongly marked characters shared in by no other mem-
ber of the genus. The knob-like tips of the hairs are trans-
lucent, having the appearance of little icicles. But although
the prominent characters remain constant, there is con-
siderable local variation.

In Ventura and Los Angeles counties (Piru City to New-
hall), for instance, the flowers are a very rich yellow and
the plant is rather low and stout. Near the city of San
Luis Obispo, the plant has the same habit, but the upper
half of each petal is backed with olive which showing
through gives a peculiar changeable color effect. In El

BoT.—VOor. II.) PURDY — CALOCHORTUS. 135

Dorado County the form is very tall and large, often three
feet in height, with flowers five inches across, and of a
lighter yellow. The hairs are longer and the knob-like tip
is smaller.

25. Calochortus concolor, sp. nov.

Calochortus luteus var. concolor BAKER, Plate in ‘‘The Garden,’’ Dec. ike
1895.

Bulb large, reddish; radical leaves narrow, glaucous, deeply channeled;
one or more cauline leaves below umbel; stem 2 feet in height, one to several
flowered, if more than one flowered, umbellate; pedicels stout, 1 to 3 inches
long; umbel subtended by linear bracts; all parts of plant very glaucous,
lower stem and sepals with a bluish bloom; sepals ovate-acute, with scarious
margins, yellowish within, brownish on back; petals a deep rich yellow,
tending toward orange, broadly fan-shaped, 15-18 lines long, and as broad
as long, slightly rounded above, the lower third densely hairy with long erect
yellow hairs; gland small, oblong; anthers yellow, linear, exceeding fila-
ments; capsule strongly triquetrous, lance-linear, attenuate above (imperfect
in type specimen).

Described from a large plant collected by Mr. D. Cleve-
land at Laguna, on the edge of the desert, San Diego
County.

This is one of the desert species found in rocky soil in
various places on the desert side of San Diego County and
in the Cuyumaca Mountains, also at Mill Creek, near San
Bernardino, and in Hermit Valley, Riverside County. The
writer does not agree with Mr. Baker in referring this spe-
cies to C’. duteus; its affinities, if any, are with C. nuttalliz
or C. clavatus. Following C. clavatus it is the largest flow-
ered, and the showiest of the yellow Mariposa lilies.

Group 5.

Petals nearly naked, gland round, small, and densely hairy with matted
hairs; leaves ashy blue, linear, deeply channeled. Desert plants.

26. Calochortus kennedyi Porter.

Calochorius kennedyi PorTER, in Coutt. Bot. Gaz., Vol. eS 77e8ps 70:
Plate in ‘‘ The Garden,”’ Feb. 11, 1893.

Stem very low, rather stout, often only 1-4 in. high, 2-4-flowered; radical
leaves linear, 2-10 in. long, channeled, very glaucous, as are the stem and
bracts; flowers produced in an umbel, which is subtended by short bracts;
pedicels 3-4% inches long; sepals ovate-oblong, about equalling petals, obtuse

136 CALIFORNIA ACADEMY OF SCIENCES. [PrRoc. 3D SER.

or acute, with white scarious margins, vermilion inside, often spotted brown
near base, brownish without; petals cuneate, rather truncate, 10-15 lines long,
not quite as broad as sepals, a dazzling vermilion, naked above, a few scat-
tering hairs below; gland small, round, densely matted with short hairs and
bordered nearly black; filaments one-half as long as the oblong-ovate, obtuse,
brownish purple anthers, which are 3-4 lines long; capsule 14-2 in. long, 4-5
lines wide, attenuate above.

Found in the arid regions of southeastern California from
near the Ojai Valley (Ventura County) and Tehachapi
Station, along the eastern flank of the southern continua-
tion of the Sierras into San Diego County, eastwardly to the
Argus Mountains in southwestern Nevada, and throughout
Arizona.

** Kern County, California.”’

In California the flowers are a vermilion color, in Arizona
and Nevada, orange. While the stem may reach eighteen
inches in height, it often hardly rises above the ground, and
the meager ashy foliage gives little promise of the dazzling
flower. It is probable that in eastern Arizona and in Nevada
C. kennedyz will be found merging into C. aureus.

27. Calochortus aureus Watson.

Calochortus aureus Watson, Amer. Nat., Vol. VII, 1873, p. 303.

Low, 4-6 inches high, with a single linear carinate radical leaf, 3-4 inches
long; scape short, 1-2-flowered, the single pair of bracts linear, 2 inches
long; sepals greenish-yellow, with a dark-purple spot near the base, oblong-
or ovate-lanceolate; petals broadly cuneate, 15 lines long, bright-yellow, with
a small well-defined, circular, densely hairy gland near the base, and a
lunate purplish spot above it; young capsule narrowly oblong, not winged.

‘* On sand-cliffs, Southern Utah.’’

The writer has no acquaintance with the species except
as scant herbarium specimens. The description and local-
ity are the original of Watson.

Group 6. BuTtTEerFLy TuLips.

Petals slightly hairy below, usually oculated and brilliantly colored; gland
prominent, round or lunate; stem bulbiferous at base, erect, slender,
branching; radical leaves, usually a pair, channeled, linear, slightly glaucous,
not ashy blue as in the desert species; cauline leaves and bracts narrow.

BoT.—VOL. II.] PURDY — CALOCHORTUS. 137

C’. luteus and C’.. venustus are the true Mariposa or Butter-
fly Tulips. It would be difficult to name any other group of
plants in which can be found such a wonderful diversity of
color and marking. The colors range from white through
lilac to purple, from delicate pink through light reds to deep
glowing reds; buffs, yellows and citrons are all present;
and the tintings, oculations, and blotchings are bewilder-
ingly numerous.

Watson bases his division of the group on the differences
in the gland: that of C. /utews being from transversely
oblong to narrowly lunate; that of C. venustus from round
to longitudinally oblong. While the casual observer would
place all together, Watson’s division is a true one. In C.
futeus the capsule is acuminate from a triangular base, in
C. venustus, linear.

28. Calochortus luteus Doxg7.
PLATE XVI, Fic. 2.

Calochortus luteus Dovuct., Bot. Register, Vol. XIX, 1833, Tab. 1567.

Plants usually dwarfish, 8-10 inches high, stem slender, stiffly erect, bul-
biferous at base, often branching; bulblets enclosed within sheath of stem;
radical leaves linear, channeled, 1-3 lines wide, bright green, slightly glau-
cous; cauline leaves or bracts linear, 1-3 inches long, peduncles 1-4 inches
long; sepals narrowly ovate-lanceolate, acute, 12 lines long, 2-3 lines wide,
yellowish within; petals cuneate, as long as broad, slightly rounding above,
yellow or tending toward orange, not oculated, but having penciled lines
radiating from gland to center of petal, claw broad; gland rather broad,
lunate, densely hairy with ascending matted yellow hairs, a very few scatter-
ing hairs above reaching to middle of petal; stamens about equalling style,
filaments slender, a little longer than the light yellow, oblong-linear, acute
anthers; mature capsule acuminate from a triangular base, 14-2 inches long.

The description is drawn from strong specimens collected
at Monterey.

‘* California.”’

The type described of C. /uteus follows the coast-line
from Anderson Valley (Mendocino County) to San Diego
County. It grows in adobe soil and is dwarfish. Speci-
mens from Mendocino, San Francisco, Monterey, and San
Diego counties are exactly alike.

138 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

Farther in the interior, from Sonoma County (Sebasto-
pol) to the Sacramento Valley, and south along the foot-
hills and higher plains of the east side of the San Joaquin
Valley, a second form grows, exactly like the type except
that it is tall and slender. This is the form commonest in
the Sacramento and San Joaquin plains. By slight grada-
tions through a multitude of forms, a dark eye develops,
and the habit becomes more luxuriant, until we have C.
luteus var. citrinus.

28a. C. luteus var. citrinus. This variety is merely a color variation of

that which follows, the color varying from yellow to deep lemon, and the
central spot being darker and not oculated.

This variety is found at its best in the Russian River
region from Hopland, Mendocino County, to the town of
Sonoma, in Sonoma County. In the latter place, the color
is a deep citron and the eye has disappeared. At various
other points, for example, eastern Lake County, Linden
(San Joaquin County), near Placerville (El Dorado
County), and in various localities in the foot-hills of the
Sierra this form is approached.

286. C. luteus var. oculatus. This variety is usually much taller, larger
flowered, and often many flowered; petals white to lilac or purple, often 2
inches long, the same in width, the center of each showing an oblong dark

brown spot oculated with yellow, and many radiating pencilings; sepals also
oculated.

The type of C. luteus var. oculatus as above described is
common in the Coast Range from Shasta County to San
Francisco Bay. It is to be expected that it would occur as
far south as Monterey County, but the writer has seen no
specimens from that region.

In the foot-hills of the Sierras, from Butte to Kern coun-
ties, there is a form slightly differing from the type. The
petals have less yellow in the middle third, the blotch is
smaller, the gland rather deltoid than lunate, and the pen-
cilings are lighter—all trifling differences which only close
study will detect; yet they are quite sufficient to separate
the oculatus of the Coast Range from that of the Sierras.

C. luteus var. oculatus of the Sierra foot-hills is very con-
stant. While in some localities it crosses with var. cztrznus,

Bot.—VOoL. II.] PURDY — CALOCHORTUS. 139

specimens from near Chico (Butte County), from Placer-
ville (El Dorado County), and from Dunlap (Fresno
County), are identical in every respect. As before stated,
C. luteus var. citrinus and var. oculatus hybridize readily,
and in many localities where both are found, there are
cross breeds in endless variety, running as is usual in such
cases to greater extremes than either parent. In Mendo-
cino County, from Ukiah to Hopland, and in El Dorado
County, near Camp Creek, these crosses are particularly
plentiful and beautiful. There are other strains of C.
luteus, each found in a more or less extensive territory.

28c. C. luteus var. robusta (C. venustus var. robusta (Hort.).) In por-
tions of E] Dorado County a form approaching C. ocudatus is found growing
in wet grounds (oftener in wire grass lands), which is there dwarfish and with
three to five almost spheroid bulblets on the stems. The colorings are very
rich; the forbidding surroundings seem to have developed unusual vitality in

this form, for in ordinary soil the bulbs produce unusually tall, stout stems
but retain the bulbiferous habit.

29. Calochortus vesta Purdy.’

Calochortus vesta PurDy, The Garden, Oct. 12, 1895, with colored plate;
Gardeners’ Chronicle, 1895, Part II, page 14.

Tall, large flowered, stem leafy, pedicels elongated, 8-12 inches long;
petals more narrowly cuneate than in C. luteus var. oculatus, white tinged
with lilac (with rare albinos), instead of an oculated spot having a broad red-
dish or dark brown band across the middle; gland narrow, doubly lunate,
extending across from side to side of petal; bulblets long and _ slender,
I to 4 to the stalk, not enclosed in sheath of stem, but set at an angle.

While C. vesta is grouped with C. duteus and C. venustus,
it is a strongly marked form which does not hybridize with
either. C. vesta is found in adobe soil (sticky black or
blue clay) from Sonoma and Napa counties to Humboldt

County, California.

1C. vesta and the varieties of C. venustus—var. roseus, vat. purpurascens, and var. eldo-
vrado—werte first mentioned in a catalogue of bulbs issued by the writer.

C. vesta was first described in an unsigned article in ‘‘ The Gardeners’ Chronicle”’
(August, 1896). It was so named in honor of the author’s wife. In the same article,
C. venustus var. roseus and var. purpurascens were also described under the names C. voseus
aud C. purpurascens, thus giving specific rank to the catalogue varieties.

A fine colored plate of C. vesta, C. venustus (the type, known as var. vosexs), and
C. venustus var. purpurascens, was published in ‘‘ The Garden” (Irondon); and a good plate
of C. venustus (type), C. luteus (type), and C. luteus var. citrinus, appeared in the same mag-
azine 1883 (?).

(3) December 5, 190%.

140 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

Calochortus bulbs move from year to year by means of a
spongy process growing from their base, at the end of
which the new bulb is formed. By this means the bulblets
of C. vesta, being set at an angle, travel some distance before
at maturity they take a vertical position. By digging into
a patch where C. vesta has been undisturbed for some
years, the ground will be found literally full of bulbs, the
immature ones placed at every angle. In most bulbiferous
Mariposas, the bulblets being placed inside the stem sheath,
and parallel to the stem, grow straight down and form a
close clump.

C. luteus var. robusta has a habit somewhat similar to
that of C. vesta, though not so marked; and in C. venustus
var. purpurascens the habit is exactly the same as in
C. vesta.

30. Calochortus venustus Doug/.

Calochortus venustus DOUGL. ex. BENTH. in Trans. Hort. Soc., Ser. II, Vol.
I, 1835, p. 412, Tab. XV, fig. 3.

Stem stiff, erect, usually branching, 4 inches to 3 feet in height, bulbiferous,
with a single bulblet at base; one or two linear, radical leaves, 1-3 lines
wide, quite glaucous; pedicels 2-5 inches long; sepals oblong-lanceolate,
4 lines wide, 18 lines long, acute, as long as petals; petals broadly obovate-
cuneate, slightly rounding above, wider than long, 16 lines long, 20 lines
wide, color white to lilac above, carmine below, oculated like C. Zuteus, with
reddish pencilings on each side of eye, and a transversely oblong, rose-
colored blotch near apex, claw reddish brown; gland roundish or oblong,
densely matted; filaments a little exceeding the narrowly oblong, obtuse
anthers; capsule linear, 2-3 inches long.

Described from a specimen collected in San Benito
County, California.

C’. venustus is even more variable than C. Juteus. While
C’. Juteus is almost always found in clay, sometimes heavy
clay, C. venustus usually grows in light open ground, often
in sandy soil.

“~s

Bot.—VOL. II.] PURDY— CALOCHORTUS. 14!

30a. C. venustus var. roseus, The Garden, Oct. 12, 1895.

The typical form described above is known as var. roseus,
Or C. s0seus.

306. C. venustus var. eldorado, var. nov. Differs from var. voseus in
having the more cuneate petals always narrower than long. Found in all
colors from white through lilac to purple, from pink through many shades to

deep claret, with or without rose-colored blotch near the top of petal. The
plants are stout, usually tall (1-2 feet), branching, and large flowered.

C. pictus (The Garden, Oct. 12, 1895) is merely a white
form of the above variety.
30c. C. venustus var. purpurascens', The Garden, Oct. 12, 1895.
This is a strong growing variety, like the type but taller; flowers lilac to
purple, petals oculated and more leafy, but without the rose-colored blotch.

The plants are very bulbiferous, producing 1-4 bulblets a year, which scatter
as do those of C. ves¢a,; they are not enclosed in a stem sheath.

Found growing in adobe (heavy clay) soils, from San
Luis Obispo County to Suisun Bay, California.

30d. C. venustus var. sulphureus, var. nov. As in the preceding, like
the type but taller; petals a light, warm yellow, with eye in center, and rose-
colored blotch at top.

Occurs at Newhall and Alcalde (Kern County), in the
lower end of the San Joaquin Valley.

C. venusius is as widely scattered as C. luteus, and has
many strains, some local, others widely spread. There are
two principal strains. One commences in the Coast Range
at Antioch (Contra Costa County) and extends southward
into the interior or dryer portion of the Coast Range to Los
Angeles County.

The type described, as found from Antioch to Paso
Robles, always has a rose-colored blotch at the top of the
petal. In this strain the petals are broader than long, giving
a very full flower. At Paso Robles there is a break, and
we find a wonderful variety of color forms, from white to
purple, and from pink to deep red, always with a rose

1 The original C. venustus purpurascens Watson was described from specimens from
Kern County,in the southern Sierras, and is without doubt only the purple extreme
of the El Dorado strain. The name has for years been used for a purple variety of the
Coast Range form, which is distinct in every way. As the latter is perfectly distinct,
with features by which it can be readily identified, and the former is a mere color
extreme, occurring usually with other colors, the rule of priority could, I think, be
shaded a little, and the name be used for the Coast Range type.

142 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

blotch at the top of the petal. From this point south to
Los Angeles County the forms seem to change frequently.

At Preston and Newsom’s Springs (San Luis Obispo
County) there is a pretty white form, in color exactly like
C. cataline (white, tinged lilac), destitute of both eye and
rose-colored blotch, but with the ovate dark brown spot on
the claw of the petal, as in C. cataline.

At Santa Barbara the type with the rose blotch reappears,
and it is found at intervals southward.

In the Ojai Valley and at Newhall there is a white-flow-
ered variety with a rose blotch and narrow petals, closely
resembling the white-flowered plants at Paso Robles. It is
also tall and slender.

At Newhall a new color strain comes in, quite novel in
C. venustus; this (var. sulphureus) is light yellow, narrow
petaled, and has the rose blotch. At Elizabeth Lake, a
point about midway between Newhall and Alcalde, the
species is again seen in the infinite variety of color found
about Paso Robles and in the southern Sierras, but with the
additional yellow form.

Newhall is the most southern point from which the writer
has specimens of C. venustus.

Mr. S. B. Parish has written a very interesting descrip-
tion of these color forms.

The Eldorado strain, so termed by the writer in a cata-
logue of bulbs, is found in the Sierra Nevada. It begins
to appear a little north of Camp Creek (El Dorado
County), high up in the foot-hills in the upper Yellow-Pine
belt (growing oftenest in open woods), and extends at a
nearly uniform altitude in the Sierras south to the Tehachapi
and Tejon mountains. The colors of the flowers are richer
than those of the Coast Range forms. They vary from
white to lilac and deep purple, from pink to deep red, and
sometimes light cream flowers are found. The eye is dark
brown, not much oculated, the blotch at the top of the petal
varies from rose to gold, and flowers can be found with rose
or gold rays across the entire top of the petal; but the rose
blotch at the apex, instead of being a constant feature as is

BotT.—VOL. II.] PURDY— CALOCHORTUS. 143

usual in the Coast Range forms, is rare. At all points from
which specimens of this form have been collected the variety
of color is shown to vary greatly in proportion; sometimes
nineteen-twentieths of the flowers are white, again red or
purple will predominate, and this in localities only a few
miles apart; but no single color form is ever found to the
exclusion of all others.

Group 7. Litac Mariposas.

Petals white, lilac, or purplish, not oculated, more or less hairy; gland
small, round, and densely hairy; leaves linear, channeled; capsules linear
except in C. cataline.

31. Calochortus splendens Dozg/.

Calochortus splendens Douci. ex BENTH. in Trans. Hort. Soc. oer lr
Vol. I, 1835, p. 411, Tab. XV, fig. 1.

Stem slender, often bulbiferous at base, usually a foot or two high; sepals
often spotted purple, ovate-acuminate, about equalling petals; petals broadly
fan-shaped, circular above, 1-5 lines long, 18 lines broad, upper half naked,
scattering short hairs around the gland and on lower third, the claw very
short; color from lilac to purple, lighter about gland, usually reddish purple
on claw; gland small and round with a mound of matted agglutinated hairs
which are scarcely distinguishable as hairs, and are frequently absent; anthers
purple, obtuse, one-third as long as filaments; capsule linear.

Described from strong plants from San Diego, California.

In horticulture the type is known as C. splendens var.
atroviolacea. It is found from San Diego to Santa Barbara
counties, and on the coast islands.

31a. C.splendens var. montanus, var. nov. Like the type but very slender,
8-12 inches high, smaller flowered, and more densely hairy about the gland
with short yellow hairs; color lilac to salmon pink; often bulbiferous.

Described from specimens from Raynetta, San Jacinto
Mountains.

316. C. splendens var. major, var. nov. This resembles var. montanus but
is not bulbiferous, is stouter and much larger flowered; petals pale lilac,
lighter below, with very long tangled hairs scattered on middle third.

Described from specimens from Monterey County, Cali-
fornia. Found also in San Luis Obispo County.

144 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

The figure shown in Douglas’ plate is that of the form
here described as the type. Baker, in his Tulipe, gives an
accurate description of this same form, calling it Calochortus
splendens; but Watson, in the ‘‘Botany of California,’’ has
described as the type the form here given as var. major.
The marked difference between these two forms is in the
hairs. In the type the hairs are short and found on the lower
third of the petal; in var. major they are on the middle
third of the petal, and are long and cobwebby. In addition
to this, the latter variety is of a stronger habit, has larger,
lighter colored flowers, and is without bulblets.

31c. C. splendens var. rubra,a form found in eastern Lake and Napa
counties, is singularly enough in gland and hairs almost exactly like the
type at the other extremity of the range. It is a very large plant, with a
deep-seated yellow bulb, tall (1-3 feet), stout stem, and a large reddish lilac,
pink or purplish flower, the petals quite hairy with short hairs on lower
third, and with a deep reddish purple claw.

C. splendens var. montanus has been mistaken for C. fal-
merz, but it is. closely related to the type; in fact, aside from
the color of the hairs above the gland, and the more slender
habit, it hardly differs from the type except in its habitat.
The writer has it from ‘‘cienagas’’ (wet springy spots) near
Tehachapi, from wet spots near Bear Valley (San Bernar-
dino County), and from wet meadows in the San Jacinto
Mountains. The other three forms always grow in dry,
rocky soil. .

32. Calochortus palmeri Watson.

Calochortus palmeri Watson, Proc. Amer. Acad., Vol. XIV, 1879, p. 266.

Stem very slender, lax and flexuous, a foot or two high, 1-7-flowered,
bulbiferous near the base; sepals with narrowly acuminate recurved tips,
spotted; petals 6-12 lines long, white (or yellowish below) with a brownish
claw, and with scattered hairs around the ill-defined, broad, densely hairy
gland; anthers obtuse, 3 lines long, capsule very narrow, an inch long or
more.

‘‘California, near the Mojave River.’’
The above description and locality are quoted from the
original of Watson.

BotT.—VOL. II.] PURDY— CALOCHORTUS. I 45

The writer has never seen a specimen which conformed
to the description; all of those seen from Southern Cali-
fornia collections were either C. zuvenustus or C. splendens
var. montanus.

33. Calochortus cataline Watson.

Calochortus cataline Watson, Proc. Amer. Acad., Vol. XIV, 1879, p. 268.
Calochortus lyoni Watson, Proc. Amer. Acad., Vol. XXI, 1886, Pp. 455.

A foot or two high, general habit as in C. sflendens; stem bulbiferous at
base; sepals broadly ovate-lanceolate, acute, a little shorter than petals,
greenish, spotted dark purple on claw; petals cuneate, longer than wide,
rounded above, narrowing abruptly to a short narrow claw; color from white
tinged with lilac, to lilac purple, with a large ovate purplish-maroon mark,
the apex covering claw and the rounded end above gland; gland oblong,
densely matted with short agglutinated hairs, a few scattering hairs around
and above; filaments slender, about four times as long as the yellowish,
oblong-obtuse anthers; capsule oblong, rounding at both ends.

Described from specimens taken near Los Angeles,
Calitornia.

‘‘Santa Catalina Island.’’ [ California. ]

C. cataline is found from Los Angeles and San Bernar-
dino along the coast to Santa Barbara, and on all the islands
along the coast. It is one of the least variable and best
marked of all the Mariposas. The oblong capsule is a
distinctive feature, while the peculiar ovate mark on the
petals is equally characteristic.

The species was first described by Watson from a plant
in the capsule, the specimens having been collected on
Santa Catalina Island. Later, he described the same spe-
cies, under the name of C. lyonz, from flowers collected at
Los Angeles and Newhall by Dr. Asa Gray and W. S.
Lyon. C. lyonz also grows on Santa Catalina Island, and
there is no doubt of its identity with C. cataline. In an
able article in ‘‘Erythea,’’ Dr. Davidson of Los Angeles
shows that the two species are the same.

34. Calochortus invenustus Greene.

Calochortus invenustus GREENE, Pittonia, Vol. II, 1890, p. 7I.

Stem very stiff, stout as compared with C. sp/endens, strongly bulbiferous
at base, the bulblets large, ovate-oblong as in C. zuttallii; radical leaf linear,

146 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

deeply channeled; plant very glaucous, as in most desert species; inflores-
cence umbellate or at least the larger divisions of stem so; umbels, or branch
divisions, subtended by three or four lanceolate bracts an inch or two long;
sepals ovate-lanceolate, acute, considerably shorter than petals, yellowish
green within; petals obovate-cuneate, short clawed, rounding above or
abruptly acute, smoky white, claw purplish; gland small, oblong, densely
hairy with matted hairs, yellowish short tangled hairs above gland; anthers
2-3 lines long, not equalling filament; capsule 3-4 lines broad in the middle,
tapering both ways.

The description given is drawn from specimens taken at
Bear Valley (San Bernardino County), California.

‘‘Higher mountains to the westward of the Mojave
Desert.’’

C. cnvenustus was discovered by the namer, Professor
Greene, near Tehachapi. It is also found in the Tejon
Mountains, and doubtlessly grows all along the line from
there to Bear Valley. Specimens from the former locality
are identical with those here described.

The characteristics of the species place it between C.
nuttallii and C. splendens. The prominent offset, stiff,
stout stem, and tendency to flower in umbels, are strongly
suggestive of C. nuttalliz, while the flowers resemble those
of C. splendens var. montanus.

35. Calochortus excavatus Greene.

Calochortus excavatus GREENE, Pittonia, Vol. II, 1890, p. 71.

Resembling the last [C. zzvenustus], but the bracts ovate-lanceolate, scar-
ious almost to the striate-veined middle portion, their acuminate tips re-
curved; petals white shaded with lurid purple above, but dark purple below
and about the broad obovate hairy gland, which is deeply impressed, appear-
ing like a yellow saccate body on the outside of the petal; stamens as in the
last, but anthers dark maroon.

‘‘From Bishop Creek, Inyo County, California, collected
by Mr. W. H. Shockley (No. 427).”’

Having little knowledge of this species, the writer has
quoted the original description and locality of Professor
Greene. From specimens seen, the species would seem to
come between C. znvenustus and C. nuttalliz.

Bot.—VOoL. II.] PURDY — CALOCHORTUS. 147

36. Calochortus flexuosus Watson.

Calochortus flexuosus Watson, Amer. Nat., Vol. VII, 1873, p. 303.

Stem slender, very flexuous or almost decumbent, a foot or so high,
branching; bracts linear or lanceolate, 6-15 lines long; sepals oblong-lanceo-
late, greenish with a deep purple spot; petals broadly obovate-cuneate, 12 to
15 lines long, purple, claw deep purple; gland obscure, purplish yellow, scat-
tering glandular hairs above gland; capsule broadly oblong.

The description given is substantially that of Watson.

‘¢ Southern Utah and Northern Arizona.”’

In 1897, the writer received a number of fresh flowers
from St. George, Utah, the type locality. They showed
the color to be a deep, rich purple (the flowers are some-
times white), the markings varying considerably, showing
bands or spots on either petal or sepal; scarcely any two
of the flowers were alike in this particular. The very flex-
uous habit is distinctive. The stems are often even decum-
bent (‘‘creeping’’ a correspondent has it). Found also in
southern Nevada.

This Calochortus grows on the hills in red granite soil.
Mr. S. B. Parish collected a species in the desert region
of Southern California, which he identified as C. flexuosus.
This, he writes, grows in tufts of grass in saline meadows.
It is, however, hardly probable that the same species grows
under such diverse conditions.

37. Calochortus dunnii, sp. nov.

Stem not bulbiferous at base, a foot or two high, slender; leaves linear,
deeply channeled; sepals ovate-acute, with white scarious margins, a little
over one-half the length of the petals, never recurving, light green without,
greenish white within, faintly spotted; petals broadly cuneate, as broad as
long, rounded above, white, with a reddish brown transverse band above the
gland; gland small and round, densely hairy with short matted hairs, short
scattering hairs on each side of the gland only; capsule linear as in C
venustus.

Described from specimens flowering in the writer’s
grounds; they were originally collected near Julian (San
Diego County), California, by the veteran naturalist,
George W. Dunn, in whose honor the name is given. So

148 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

far, this pretty new species is known only from the type
locality. It forms a connecting link midway between C.
splendens and C. venustus.

Group 8. GREEN BANDED Mariposa.

Petals purplish lilac, with a greenish line down the back, obovate-acuminate.

38. Calochortus macrocarpus Dozg/.
PiaTe XVIII.

Calochortus macrocarpus DouGt., Trans. Hort. Soc., Vol. VII, 1830, p. 276,
Tab. VIII.

Stem bulbiferous at base, stout, erect and rigid, 1-2 feet high, one or more
flowered; the single radical leaf, linear, deeply channeled; cauline leaves
3-5, narrow and convolute; sepals about equalling petals, acuminate, purple
inside or tinged with purple, darker at base, sometimes spotted and hairy,
with a broad scarious margin; petals obovate, narrowly acute or acuminate,
or often distinctly cuspidate, a deep purple lilac, lighter at base, with a broad
light-colored claw 1%4-2% inches long and half as wide, a greenish line down
the middle, the lower third above the gland with scant glandular hairs; gland
oblong, densely hairy; anthers purple or yellow, lanceolate, obtuse, 4-6 lines
long, about equalling filaments; capsule attenuate upward, 1%4-2% inches
long.

In sandy deserts from northeastern California (Modoc
and Lassen counties) to eastern Washington and into Idaho.

A very clearly marked species varying but little. There
is a white form near Pullman, Washington. The Indians
of the northwest are very fond of the bulbs.

Group 9. Serco LIiLigs.

Flowers lilac, white, yellow or pink; gland round; stem prominently bul-
biferous at base, umbellate.

39. Calochortus nuttallii Zorr. & Gray.

Calochortus nutiallii Torr. & Gray., Pacif. R. R. Rept. (Bot.), Vol. II,
1854, p. 124.

Stem erect, stiff, a foot or two high, with a large oblong bulblet at base;
infloresence simply umbellate, 1-5 flowered; radical leaf linear, deeply chan-
neled; cauline leaves 1 to 3, narrow, glaucous, revolute; sepals ovate-lanceo-
late, with scarious margins, yellowish within, with or without a dark spot at
base, which is sometimes hairy, much shorter than petals; petals broadly

Bot.—VOL. II.] PURDY — CALOCHORTUS. 149

obovate-cuneate, with a rather narrow claw, abruptly acute or rounding above,
white, yellowish below, with a small round or oblong gland which is covered
densely with agglutinated hairs, a few long hairs scattered about the gland;
anthers oblong-obtuse, more or less sagittate at base, shorter than filaments;
capsule lanceolate, 1-2 inches long.

Described from specimens from southern Utah.

C. nuttallit has the widest range of any Calochortus. It
is found from the western flank of the Sierras to Nebraska,
and from the Snake River to New Mexico. In color it
varies greatly and in other characters slightly. The region
of its habitat is too little known to enable any one to mark
the range of the variations definitely. In Nebraska and
eastern Colorado it is yellow; in many places in central
Colorado the color varies wonderfully from white and pink
to purple; from Salt Lake to Reno, Nevada, and southern
Utah, but one form seems to be common, that which is the
type here described; at Pine Valley, southern Utah, the
flowers are cream colored; and at Dimee, New Mexico,
they are orange.

40. Calochortus leichtlinii! Hooker.

Calochortus leichtlinit HOOKER, Bot. Mag. Tab. 5862.

A dwarfed or alpine form, quite different from any of the other forms,
often but an inch or two in height; stem more slender; anthers strongly sag-
ittate; petals smoky white with a dark spot.

Found in the Sierras at an altitude of from 6,000 to 9,000
feet.

There is another variety of this dwarfed alpine Calochor-
tus also found in the Sierras, but the description of this
form will be reserved until there is fuller material to draw
from.

1C. leichtlinit is included under C. uuttallii by Watson, in the Botany of California,
Vol. II, p. 177.

150

CALIFORNIA ACADEMY OF SCIENCES.

EXPLANATION OF PLATE XV.

Calochortus purdyt.

eRe er aS

MP QO

Plant, actual size.

Petal.

Stamen, enlarged.
Sepal.

Scale on petal, enlarged.
Ripe pod.

[PRoc. 3D SER.

Proc.CaLAcan. Sct.4? Ser. Bor. Vou II {Puroy]PLATE XV

PHOTO -LITH BRITTON & REY, SE

CALOCHORTUS PURDYT.

>

152 CALIFORNIA ACADEMY OF SCIENCES. [PRoC. 3D SER.

Fig. I.

Fig. 2.

EXPLANATION OF PLATE XVI.

Calochortus longebarbatus.

a, (Petal:

6, Petal, less enlarged.

Calochortus luteus.

a, 6, Petals.

c,d, Capsule at different stages of development.

PrRoc.CAL.ACAD. Sct.32 SER Bor. Vou Il PuRDY| PLATE XVI

er ae Ta |r "yal,

wi WL,
w \ 5 i} fi
‘ 4

PHOTO -HITH BRITTON & REY, SF.

Matrnrinomec TANPIDVADRATNTC
L LALOCHOURTHS LUNGIBARBATUS
UAT MAC 7 TE mMPETTe
(aie LALUCHORTUS LUTEUS

&

Eke Win Ae FP els
i oe = ne

154 CALIFORNIA ACADEMY OF SCIENCES. [PRroc. 3D SER.

EXPLANATION OF PLATE XVII.

Calochortus weedit.

a, 6, Capsule at different stages of development.
C, Sepal.
d,e, Petals.

PHOTO

O-WITH, BRITTON & REY, SF.

‘ g
- i
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ay ‘ . :
:
7
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ety '
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'
. i .
- . . ~ ;
’
tw
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i { “ me
“ ‘ vs eee ml fe - ae e .
i wry ‘ eS ay
ear ? i - 1? ‘
7 ad *
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| Ais; 3 :
i f
eu) ws
: ‘ 4 S y .
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Wai 54
Hee

156 CALIFORNIA ACADEMY OF SCIENCES.

EXPLANATION OF PLATE XVIII.

Calochortus macrocarpus.

)
;
°
} id
:
7
-
_ a - 7
en Py 7 hie
- a ; a Ng
—— > ain hb a nm a fa Y

PRoc.CAL. ACAD. 5c1.32 SER. Bot. Vou ll

}
>

158 CALIFORNIA ACADEMY OF SCIENCES. [PrRoc. 3D SER.

EXPLANATION OF PLATE XIX.

Petals of Calochortus species.

1. C. amabilis.

2. C. benthamt.

3. C. maweanus.

4. C. amenus.

5. C. albus.

6. C. vesta.

7. C. gunnisont.

8. C. longebarbatus.
g. C. nuttallit.

1o. C. venustus.

11. C. venustus roseus.
12. C. cataline.

13. C. howellit.

14. C. dunnit.

15. C. clavatus.

16. C. howellii, flower.
17. C. dunnit, flower.
18. C. clavatus, flower.

Proc CAL Aca, $c1.32 SER. Bor Vou I] | PURTIY | PLAT}

A GROUP OF WESTERN AMERICAN SOLANUMS.

BY S. B. PARISH.

THE first notice in botanical literature of any of the group
of Solanums which is the subject of the present paper was
in 1826, when in the Memoirs of the Academy of St.
Petersburg the name Solanum umbelliferum was bestowed
by Eschscholtz on a plant from ‘‘ Nova California.’’ As
the original description is not easily accessible, a transcrip-
tion of it, for which I am indebted to Dr. J. N. Rose, is
subjoined in the foot-note.'

The floral characters here assigned are common to a whole
series of plants that the student of Californian botany finds
it unsatisfactory to unite, and as difficult to separate on
stable lines; nor are the vegetative characters defined with
sufficient accuracy to enable one to determine which of these
plants the author had in hand.

'4. Solanum umbelliferum sch.

Solanum umbelliferum Escu., Mem. Acad. Petersb., Vol. X, 1826, p. 283.

‘*(Inerme, foliis integerrimis, calycibus quinquedivisis, staminibus zequalli-
bus, inflorescentia terminali. )

“Caule suffruticoso, erecto, foliis ovalibus, acutis, integerrimis, pubescenti-
bus; umbellis terminalibus.

“Tn fruticetis Nove Californiz.

‘“‘Caulis orgyalis, suffruticosus, fistulosus, angulatus, pubescens; ramis sub-
herbaceis, nutantibus, tomentoso pubescentibus.

‘Folia alterna, petiolata, ovalia, acuta, integerrima, utrinque pubescentia,
vix pollicaria, caulina interdum late ovata sesquipollicaria.

‘Flores terminales umbellati; umbella plerumque quadriflora, interdum bi-
vel-triflora; involucrum parvum urceolare, integrum, pubescens; pedunculi
zequales, elongati, pubescentes. Calyx urceolaris quinquefidus pubescens,
laciniis acutis. Corolla calyce triplo major, dilute violacea, quinquefida,
extus pubescens. Antherz flave.

“Bacca magna purpurea.”’

The ‘‘involucrum parvum’? is, of course, the organ better designated by
Dr. Gray in the Synoptical Flora as a ‘‘cupulate node’’ at the insertion of the

pedicels on the peduncle.
[159] October 23, Igol.

160 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

Dunal (1852), in De Candolle’s Prodromus, substantially
repeated the above diagnosis for the original species, and
added to it two others. The significant points in the
descriptions of these species are that the one, S. genzs-
tordes, is characterized as ‘‘ramudlosissimis pilosis,’’ and the
other, S. californicum, as ‘‘ tomentosis candicantibus.’’ ‘The
first of these was founded on a specimen collected in Cali-
fornia by Douglas, and the latter on another plant from the
same region and collector, and one of Pavon’s from ‘‘Nova
Hispania.”’

In 1876, Dr. Gray, in a contribution to the Proceedings
of the American Academy, reduced Dunal’s two species to
the original Eschscholtzian S. wmbelliferum, making the
branched hairs of the pubescence the essential character of
that species, and at the same time proposing a new species,
S. xantz, for the reception of certain plants which had since
come to hand, and in which the pubescence was of ‘‘simple
and few-jointed hairs, some of them glandular.’’ No type
is specified, but reference is made to specimens collected
by Xantus de Vesey, Bigelow, Anderson, and Lemmon.
A variety, S. xanti wallacet, was proposed for a plant
collected on Santa Catalina Island by Wallace.

There are preserved in the Gray Herbarium of Harvard
University authentic specimens of all the above species,
those representing Eschscholtz’s, and Dunal’s types coming
from the Herbarium of Trinity College, Dublin. By an
examination of these it is possible to ascertain exactly what
were the plants intended by the different authors, and what
more recently collected material may be included with
them.

A specimen collected by Hartweg is accompanied with a
note in Dr. Gray’s hand, certifying it to be an original of
Eschscholtz’s S. umbelliferum. It has a stem moderately
hirsute (but not canescent) with a mixture of unbranched
and few-branched hairs in about equal proportion. The
leaves are ovate, obtuse at base, about two centimeters
long, sparsely hirsute, the hairs short and mostly unbranched.
All the hairs are unilocular and not glanduliferous.

Bot.—Vot. II.] PARISH—WESTERN AMERICAN SOLANUMS. 161

An original Douglasian specimen represents .S. genis-
totdes. The stems are slender; the leaves few, somewhat
fascicled, minute (5-6 mm.), and ovate; the hairs on the
peduncles are mostly branched, on other parts of the plant
they are entire, or a few once-branched. As in the first
species, all are one-celled and glandless. The specimen has
the appearance of coming from a starved plant, as Dr. Gray
suggests. Coulter's No. 590, also from Trinity College
Herbarium, is a like form.

A specimen of the plant of Douglas, on which S. cali-
fornicum was based, has stout stems canescently tomentose,
with hairs all of which are branched. The leaves are broadly
ovate (2-4 cm.), cuneate at base, and sparsely hirsute with
mingled unbranched and few-branched hairs. The struc-
ture of all the hairs is as in the other species. A specimen
by Fremont, mounted on the same sheet, is a better repre-
sentative of this form, and has like characters.

The plant collected by Xantus de Vesey, No. 73, at or
near Fort Tejon, California, may be taken as the type of
the species which bears his name. It has oblong leaves
(circ. 3 cm.) mostly acute at summit and cuneate at base,
except a few leaves lobed and truncate at base. The whole
plant is moderately hirsute with rather short plurilocular
hairs. Anderson’s, Lemmon’s, and Bigelow’s plants are on
the same sheet. The first is quite like that of Xantus; the
other two are mere fragments with similar pubescence and
oblong leaves, which are obtuse or subcordate at base, and
probably were taken from plants whose lower leaves were
cordate. Wallace’s Santa Catalina Island specimen (type
of the variety wa//acez) is likewise fragmentary. It consists
of the summit of a branch with an ovate leaf (9 cm.) and a
few young leaves, and an umbel of flowers with large corollas
(4 cm.); the pubescence of the stem is of long tawny
hairs, multilocular and viscid-glandular; the leaf is nearly
glabrous above and tawny-villous below. All the hairs
appear to be unbranched, but other specimens commonly
show some few-branched hairs.

162 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

From the above notes on these type specimens it will be
seen that the character which has been relied upon as dis-
tinctive of S. umbelliferum, namely, ‘‘ hairs branched,”’ is
only partially applicable, not one of them being without a
considerable admixture of unbranched hairs. Indeed, I
have not been able to find a single specimen of this species,
among the many examined, in which more or less unbranched
hairs could not be detected. On the other hand, many
plants are found with a few, often a very few, branched
hairs intermixed with the prevalent simple hairs. More-
over, the character of the pubescence, at least in most
specimens of S. wmbelliferum, is different on the various
parts of the same plant. On the young stems, notably
towards the tips, it is exclusively, or nearly so, of many-
branched hairs, while on the leaves, notably on the older
ones, it is largely, sometimes exclusively, of unbranched
hairs.!

A more satisfactory character is found in the structure
of the hairs. In S. wmbelliferum these, whether branched
or unbranched, are without cell divisions and are not
glanduliferous. In the plants which have been referred
to .S. xantz the hairs consist of elongated cells, some of
the cells usually evacuate and collapsed, or atrophied, so
that the hairs have a peculiar unevenness; many of them
are tipped with black globular glands, causing the plant to
be more or less viscid. In glabrate forms the hairs are very
short and mostly reduced to a single cell, but they remain
glanduliferous. Unfortunately, this character is not entirely
constant, and it is possible to find specimens on which
there are branched hairs which are also plurilocular and
glanduliferous.

The leaves of the group exhibit a wide range of variation
in shape, passing from orbicular to oval, oblong, elliptical,
and even lanceolate; the apices are either acute or rounded,

1In a specimen collected by Kellogg and Harford near Bear Harbor, on the northern
coast of California, a third form of hair occurs on the oldest stems. These are densely
tomentose with the usual mixture of branched and unbranched hairs, from which stand
out scattered spinose branched hairs, 2-4 mm. long.

BoT.—VOL. II.] PARISH—WESTERN AMERICAN SOLANUMS. 163

the bases attenuate, cuneate, rounded, truncate, or cordate.
Often a part of this range of variability may be seen ona
single plant, and series of specimens can be arranged
readily passing by intergradations from one extreme to the
other. A tendency to segmentation is manifested by the
occasional occurrence on a specimen of a few leaves with
a pair, or even two pairs, of basal lobes. The leaves are
either smooth or papillose-roughened beneath the pubes-
cence, and are either entire or crenate margined, the latter
character appearing to be constant and of some diagnostic
value.

The floral characters are practically the same throughout
the group, except that .S. wal/aceZ usually has a narrower
and deeper lobed calyx than the others. The corollas are
from two to five centimeters in diameter, and in color vary
from light to very dark violet, the centers having green
markings; the long, bright yellow anthers are sagittate at
base, and are on very short, stout filaments; the style
exceeds them one-half to one-third its length, is usually
straight, but the included portion is sometimes bent. The
fruit is very little known; it may possibly afford some satis-
factory characters when better understood. It is a smooth,
globular, many-seeded berry, about two centimeters in
diameter, or perhaps sometimes smaller. In S. wadlacei
it is certainly dark purple when ripe, and this color has
usually been assigned to the fruit of all the members of the
group; but according to Professor Greene! the ripe fruit
of S. umbelliferum is ‘‘yellow;’’ that of S. xantz I have
never seen more than a light or whitish green when
apparently ripe.

Such being the generally inconstant characters of this
group of plants, it is evident that their satisfactory segrega-
tion is a matter of no little difficulty. The exercise of that
botanical industry which multiplies ‘‘ species’? by the min-
ute description of individuals might reap here an abundant
harvest. On the other hand, a rigorous insistence on sharp

1 Man. Bay Region, p. 268.

164 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

and absolute delimitations, which should exclude all inter-
grading forms, would reduce the whole group to a single
heterogeneous species, a disposition, it is safe to say, unac-
ceptable to any one who has studied these plants. Avoid-
ing these two extremes, I have attempted a classification
with reference to certain diverging lines of development
which are manifested, without insisting upon definite cleav-
ages, which do not exist. Such differences, as every
botanist is aware, are more easily perceived in the exam-
ination of copious material than characterized. In the
present instance the facts as they are in nature are repre-
sented by such a treatment; for we have here a series of
plants in which variation has outrun the processes of selec-
tion, and in which the connecting forms yet remain to unite
the diverging lines of evolution. It has also the advantage
of preserving the species recognized by Dr. Gray, who
with his accustomed discernment fixed upon the best diag-
nostic character, that of the pubescence, although it does
not possess the definiteness he seems to have supposed.

To the group I have prefixed an Arizona plant with a
very different corolla, but with further characters shared
by the other members of it.

KEY TO THE GROUP.

Corolla deeply 5-cleft; nodes of the peduncle obsolescent. 1. S. arizonicum.
Corolla angulately 5-lobed; nodes cupulate, prominent.
Leaves mostly linear-lobed at base. 2. S. tenutlobatum.
Leaves seldom lobed.
Plants viscidulous; hairs unbranched.
Leaves crenate; corollas large.

Stems long-hirsute. 3. S. wallacei.
Hispidulous or glabrescent. 4. S. wallacei viridis.
Leaves with entire margins.
Acute or merely obtuse at base. 5: 3. SOME
Cordate or subcordate at base. 6. S. ranti intermedium.
Attenuate at base, small. 7. S. xanti glabrescens.
Plants not viscid; hairs branched.
Stems villous. 8. S. umbeliiferum.

Stems canescently tomentose.
g. S. umbelliferum californicum.

Bot.—Vot. II.] PARJSH—WESTERN AMERICAN SOLANUMS. 165

* Suffrutescent or suffruticose plants; peduncles lateral, or by the suppres-
sion of the growing apex apparently terminal, styles clavate; fruit a many-
seeded berry.

+ Corollas small, 5-cleft; peduncles slightly thickened at the articu-
lation of the pedicels.

I. Solanum arizonicum.

Barely suffruticose or even herbaceous; stems 3 m. high, not striate or
angled, pubescent with unbranched hairs, the upper part canescent, as are
the lower surfaces of the leaves; leaves ovate-lanceolate, 2-3 cm. long, prom-
inently anastomose veined, the lower half of the margins coarsely toothed;
flowers in small corymbs (about 7-flowered); peduncles surpassing the leaves;
pedicels short, 2-5 mm.; calyx 3 mm. high, the lobes ovate; corolla light
purple, pubescent without, 5-6 mm. wide, 5-cleft nearly to the base into
ovate-acuminate lobes; anthers 3 mm. long, on filaments 1 mm. long; style
hirsute below; fruit not seen.

flabitat: Wot Springs, Arizona (397 Toumey, June 17,
1892 [N]).’

+ + Corollas rotate, angulately 5-lobed, violet, with green markings
at base; peduncles thickened into a cupulate node at the articulations of the
slender pedicels.

++ Pubescence of several-celled, unbranched hairs.

2. Solanum tenuilobatum.

Suffrutescent, stems slender, angled, glabrescent below, hirsutulous above
with short, one- to several-celled, non-glanduliferous hairs; leaves linear to
narrowly oblong, 2-3 cm. long, the midrib prominent, all but the uppermost
with a pair of hastate linear lobes at base; umbels 1-4-flowered; corolla 12-15
mm. wide; fruit not seen.

Fflabitat: Mexico,—Lower California (probably near

Ensenada, C. C. Parry, April, 1882, type [G]); (Carrizo
Creek, Brandegee, April 19, 1893 [A]).

1 The lettersin brackets denote the herbaria in which specimens are deposited. [A]
Herbarium California Academy of Sciences, San Francisco; [G] Gray Herbarium, Har-
vard University; [N] National Herbarium, Washington; [P] Herbarium of S. B. Parish,
San Bernardino, Calif.; [U] Herbarium University of California.

It isa pleasure to record my thanks to Miss Alice Eastwood, Dr. B. L. Robinson, Dr.
J. N. Rose, and Dr. Willis L. Jepson, for the opportunity of examining the collections of
which they are the custodians. Iam also under obligations to Mr. H. M. Hall for speci-
mens and other favors.

166 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

3. Solanum wallacei.

Solanum xanti wallacei Gray, Proc. Am. Acad., Vol. XI, 1876, p. 90; in
Brew. & Warts. Bot. Calif., Vol. I, 1876, p. 539; Syn. Fl., Vol. IT, Pt. 1,
1878, p. 229. GREENE, Bull. Cal. Acad. Sci., Vol. I, 1885, p. 226; id.,
Vol. II, 1887, p. 408. Lyon, Bot. Gaz., Vol. XI, 1886, pp. 204, 334, 336.
BRANDEGEE, Zoe, Vol. I, 1890, p. 143. FRANCESCHI, Zoe, Vol. IV,
1893, p- 137. Davipson, Pl. Los Angeles Co., p. 21 (1896). TRASK,”
Erythea, Vol. VII, 1899, p. r4o.

Solanum aanti WATSON (not Gray), Proc. Am. Acad., Vol. XI, 1876, p. 117.

Suffrutescent, ‘‘ often forming round masses;’’ stems about a meter long,
densely tawny-villous with long, multilocular, viscidly glanduliferous hairs
which are unbranched, or usually a few once-branched; leaves thickish,
sometimes pustulose, usually less densely villous than the stems, crenate
margined, the lower ample, cordate, the upper ovate, rounded, or subcordate
at base; calyx narrowly funnel-form, deeply cleft, or wider and less deeply
divided; corolla 2-4 cm. wide; style glabrate, or villous below; ripe fruit dark
purple.

Flabitat: Islands off the Coast of California and Lower
California, and near the seacoast in Central California.

California (Brandegee [G]); Central California (429
Palmer [A});

Los Angeles County (Santa Catalina Island, Wallace,
type [G]; 76 Lyon, 1885 [G]; McClatchie, Nov., 1893
[PiseMes> Trask, Dec. 2894 (A, Ni )s

Santa Barbara County (Santa Cruz Island, Greene,
1886 [A]; U.S. S. Albatross, Feb., 1889 [A]) (Santa
Rosa Island, Brandegee [A]) (Santa Barbara [no collect-
or’s name] Jan., 1892 [A]) (Santa Inez Mountains, Brande-
gee, 1885) [/A]);

San Louis Obispo County (Mrs. Blochman, May, 1893
[A]; 428 Palmer, June, 1876 [A]);

Marin County (Miss Eastwood, Oct., 1896 [A]).

Mexico (Guadaloupe Island, 62 Palmer, 1875 [G];
Greene, April, 1885 [P]; Franceschi, Jan., 1893 [A
(much reduced form), N, P, U]).

4. Solanum wallacei viridis.
Stout, erect, glabrate, or above hispidulous; hairs mostly reduced to a

single cell; leaves ovate, cordate, or rounded at base; calyx cup-shaped, with
short lobes.

Bot.—VOL. II.] PARISH—WESTERN AMERICAN SOLANUMS, I 67

FHlabitat: Central California, near the coast.

California (586 Coulter [G]);

Monterey County (Pacific Valley, Miss Eastwood, May,
1897, type [A, P]) (Santa Lucia Mountains, Willow
Creek, RK Ay Plaskett) Keb.,'1898 [A ]);

Marin County (Mt. Tamalpais, Miss Eastwood, May 30,
1896 [A]).

5. Solanum xanti Gray.

Solanum xantt Gray, Proc. Am. Acad., Vol. XI, 1876, p. 90; in Brew. &
Wats. Bot. Calif., Vol. I, 1876, p. 539; Syn. Fl., Vol. II, Pt. 1, 1878, p.
229. Davipson, Pl. Los Angeles Co., p. 21 (1896). McCrarcuie, FI.
Pasadena, p. 641 (1895). COvILLE, Death Valley Rept., pp. 167, 251
(1893).

Stems slender, 3-10 dm. long, woody, or at high altitudes herbaceous from
a lignescent base, the younger angled, moderately villous, with many-celled,
unbranched hairs, most of them gland-tipped; leaves ovate, ovate-oblong to
oblong-lanceolate, 1-4 cm. long, acute or obtuse at base; corolla 1-2 cm.
wide; mature fruit apparently light green in color.

Hairs with some of the cells atrophied, and sometimes a few are once-
branched.

Hlabitat: Throughout California, except in the desert
region, ascending to 6,500 feet altitude in the mountains,
and reaching the borders of Arizona and Lower California.

California (428 Palmer, 1876 [N]; 186 Thomas Bridger
[N], a canescently tomentose form); central California
(429 Palmer, 1876 [N]) Sierra Nevada Mountains
(Lemmon, 1875 [N]);

Alameda County (Piedmont, F. W. Koch, March, 1895
[U]);

Calaveras County (1358 Davy, May, 1898 [P, U].
*‘Viscid to the touch, herbage malodorous;’’ leaves thin,
neurose) ;

Fresno County (Fort Miller, Heermann, July, 1853
[N]) (Toll House, 2,050 feet altitude, 7 Hall. & Chandler,
June, 1900 [P]) (Pine Ridge, 5,000 feet altitude, 93
halle @handier,. \juney) 1900) (Pi) (Dinkey Creek;
5,300 feet altitude, 355 Hall & Chandler, June, 1900
[P]) (North Fork King’s River, 6,000 feet altitude, 448
Hall & Chandler, July, rg00 [P]);

168 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

Inyo County (Willow Creek, 789 Coville & Funston
[G, N, PJ);

Kern County (Fort Tejon, 73 Xantus, 1857-8 [G, N]);

Los Angeles County (San Gabriel, 108 Brewer [G],
the specimen in [U] under this number is of the var.
glabrescens) (Borders of Mojave Desert, Antelope Valley,
Pringle, May, 1882 [N]) (Elizabeth Lake, 1888 Parish,
June, 1887 [P]) (Saugus, Brandegee [A]) (Antelope
Valley, 2343 Davy [U]);

Mariposa County (Yosemite Valley, Mrs. Dodd [U]);

Mendocino County (Eel River, W. G. Wright, 1894 [ A]);

Placer County (Truckee River, July, 1886, 49 Sonne
[A]; 398 Sonne [P]);

Plumas County (Mrs. Austin [G]);

San Bernardino County (Bear Valley, altitude 6,500 feet,
H. M. Hall, July, 1899, [P]; 3382 Parish, June, 1894 [N])
(Bloomington, Parish, March, 1897 [P]) (Reche Cafion,
Parish, 1897 [P]);

Santa Barbara County (Bartlett Canon, 131 Rothrock,
1875 [G]);

Santa Clara County (Saratoga, 254 Davy, Sept., 1893
[U]);

Sierra County (Webber Lake, Lemmon [G]);

Sonoma County (Freestone, Miss Eastwood, March,
1899 [A]);

Tulare County (Mineral King, Brandegee [A}]).

Arizona (Palmer, 1869 [N]) (Central Arizona, 427
Palmer, 1876 [N]) (Fort Apache, 607 Palmer, June, 1890
[G, N]).

Mexico,—Lower California (Ensenada, 3711 Jones, April,
1882 [G], a transition to var. elabrescens) (San Pedro
Martir, Brandegee, May, 1893 [A]).

6. Solanum xanti intermedium.
Solanum 2aanti CoviLLe (not Gray), Death Valley Rept., p. 257 (1893).

Stems woody, lax, up to 2 m. long, viscid, leaves cordate to oblong, at least
obtuse at base, 3-15 cm. long; corollas 2-4 cm. wide.

Few-branched hairs are often present, indicating a transition to S. wmbel-
liferum; while in size and shape of leaf this form passes into S. wadllacet
through its variety vzridis.

Bor.—Vou. I.] PARISH-WESTERN AMERICAN SOLANUMS. 169

FHlabitat: California, from Sonoma County southward,
chiefly in the foot-hills, but ascending the mountains to
8,000 feet altitude in Southern California.

California (Bigelow [N]; Chas. Sayre, 1875 [N]);

Kern County (Havilah, 1064 Coville & Funston, June,
1891 [N]);

Los Angeles County (Cucamonga, Bigelow, 1853-4
[G]) (‘‘ Cahuenga Pass,’’ 189 Brewer [G]) (‘¢ San Fer-
nando Valley,’’ 189 Brewer [U]) (‘‘ San Fernando Plains,”’’
207 Brewer [U]) (‘‘ Santa Susana Mountains,”’ 207 Brewer
[G]) (Compton, McClatchie, 1897 [P]) (Pasadena, Mrs.
Brandegee [A]);

Marin County (Redwood Cajfion, Miss Eastwood, March,
1896 [U]);

Monterey County (Santa Lucia Mountains, 440 G. R.
Vasey, 1880 [N]; 20 R. A. Plaskett, Feb., 1898 [N]);

Napa County (Jepson, May, 1897 [P, U]);

Riverside County (Santa Ana River, 141 H. M. Hall,
May, 1895 [P]);

San Bernardino County (San Bernardino, 441 G. R.
Vasey, 1880 [N]); 4388 Parish, May, 1897, type [G,
N, P, UJ) (San Antonio Mountains,—Lytle Creek, altitude
e750) Leet, idl M), Halls June, 1s09; [N, (P50 Swarthout
Canin, altitude 6,500 feet, H. M. Hall, June, 1899 [G, N,
P, U));

Santa Barbara County (Santa Inez Mountains, G. W.
Dunn, May, 1891 [A];

Sonoma County (191 Samuels [N], a form with shorter
hairs) ;

Tulare County (Long Meadow, altitude 8,o00-9,000 feet,
206 Palmer, June, 1888 [N]).

7. Solanum xanti glabrescens.

Stems woody, slender, ro-15 dm. long, glabrate, or above hirsutulous with
short, mostly one-celled hairs; leaves smaller (2-6 cm.), oblong, elliptical or
lanceolate, mostly attenuate or acute at base; corolla 2 cm. wide.

Habitat: From southern Oregon, throughout California
(excepting the desert region), to northern Arizona and
Lower California. Also doubtfully reported from New

170 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

Mexico. This is the most widely distributed form, but
apparently confined to lower altitudes. Stems usually lax,
and seeking support from other shrubs, but in open ground,
notably near the coast, forming low, compact clumps.

Oregon (Josephine County, Howell, May, 1884 [G}).

California (294 Fremont, 1846 [G]; ‘‘R.N. A.’’ 1896
[N]) (Sierra Nevada Mountains, Lemmon, 1875 [N])
Southern California (285 Parry & Lemmon [G]);

Alameda County (Berkeley, April, 1900, H. M. Hall [P]);

Butte County (Clear Creek, 191 H. E. Brown, April,
1897 [A, N]) (Little Chico, Mrs. C. C. Brown, April, 1897
[A])

Calaveras County (Mokelumne Hill, 83 Blaisdell [A]);

Colusa County (Epperson’s, Mrs. Brandegee [A]);

Lake County (Mrs. Brandegee, July, 1884 [A]) (Snow
Mountain, Mrs. Brandegee [A]):

Los Angeles County (San Gabriel Canon, 108 Brewer
[U]) (Compton, McClatchie, 1896 [P]);

Modoc County (Goose Valley, M. J. Baker [U]) (Little
Hot Springs Valley, Baker & Nutting, July, 1894 [U]);

Monterey County (Santa Cruz, 2223 M. E. Jones, June,
1881, in part [A]);

Napa County (Zem Zem, Jepson, July, 1892 [U]) (Vaca
Mountains, R. H. Platt, March, 1898 [A]);

Placer County (Mrs. M. M. Hardy, 1893 [A]) (Apple-
gate, Mrs. H. Smith [A]);

Riverside County (San Jacinto River, 3115 Leiberg,
March, 1898 [N]);

San Bernardino County (San Bernardino, 4384 Parish,
May,'2897, type [G, 'N; P, U])3.

San Diego County (San Diego, Cleveland, 1874, 1875
[G]; April, 1881 [P]; Dec., 1883 [A]; Mrs. Brandeges
[A]; Greene, March, 1885 [A], avery leafy form; Miss
Cummings, April, 1896 [G]) (Alpine, Mrs. Brandegee [A])
(Temecula Cafion, Greene, 1885 [A]) (Fallbrook, Parish,
Nov., 1891 [P]) (Witch Creek, Alderson, May, 1894 [P])
(San Isabel, A. W. Henshaw, April, 1893 [N]; H. M. Hall,
May, 1899 [P]) (Oceanside, 4437 Parish, June, 1897 [A,
G, N, P, U], a compact, maritime form) (‘‘ Southwestern

BoT.—VoL. II.] PARISH—WESTERN AMERICAN SOLANUMS. I yi

part of Colorado Desert,’’ Orcutt, April, 1889 [N], but
an error in locality is probable) ;

Siskiyou County (Yreka, 877 Greene, June, £376). Gal):

Arizona (Fort Mojave, Cooper [G]).

New Mexico (‘‘Chiefly in the Valley of the Rio Grande,
below Dofia Ana,’’ torr Mexican Boundary Survey eae
perhaps an error of locality).

Mexico (Lower California, Pringle, April 6, 1882 [N}.
A transition to S. tenuilobatum).

++ ++ Pubescence of one-celled hairs, at least those of the stems
mostly many-branched, not gland-tipped.

8. Solanum umbelliferum Zsch.

Solanum umbelliferum Escu., Mem. Acad. Petersb., Vol. X, 1826, p. 283.
Duwat in DC. Prodr., Tome XIII, 1852, p.86. Gray, Proc. Am. Acads
Vol. XI, 1876, p. 90; in Brew. & Wats. Bot. Calif., Vol. I, 1876, p. 539;
Syn. FI., Vol. II, Pt. 1, 1878, p. 220.

Solanum gentstoides DuNAL, in DC. Prodr., Tome XIII, 1852, p. 86.

Suffrutescent or suffruticose, stems slender and erect, I m. or more in
length, moderately hirsute, the hairs glandless and without cell divisions,
mostly branched, but some simple, and these often predominating on the
leaves; leaves thin, ovate to oblong, obtuse or somewhat acute at base.

ffabitat: Coast counties of central California as far south
as Santa Barbara. A doubtful form from Mexico.

California (Hartweg, type [G]; 587 Coulter [Gul
Bloomer [G]; Kellogg [G]; Douglas, type of |S. genis-
tordes [G]; 590 Coulter, same form [G]);

Alameda County (Berkeley, Greene [A]; McLean [U])
(Sunol, Congdon, May, 1892 [P]; Jepson, March 9, 1900
[P]); .

Mendocino County (Bear Harbor, 717 Kellogg & Harford,
July, 1869 [N1);

Monterey County (Fremont, Jan. 31, 1846 [G]);

San Francisco County (Bigelow, 1853-4 [G]; Kellogg
[G]; Bolander, 1866 [N]);

San Mateo County (Bolander, 1892 [G]) (Crystal
Springs, Miss Eastwood, April, 1896 [A, U]);

Santa Clara County (Stanford University, 104 C. Rutter,
Bely..10, 1992 {IN ]);

172 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

Santa Cruz County (Santa Cruz, 2223 M. E. Jones, in
part [A]) (Glenbrook, Santa Cruz Mountains, H. Davis,
April, 1899 [A]).

Mexico (San Martin Island, 30 Anthony [G, N]. Nearly
leafless, leaves small, orbicular, trifoliately lobed; probably
distinct).

g. Solanum umbelliferum californicum.

Solanum californicum DUNAL, in DC. Prodr., Tome XIII, 1852, p. 86.
Solanum umbelliferum GREENE (not Escu.), Man. Bay Reg., p. 267 (1894).

Stems stout, erect, densely and canescently tomentose with many-branched
hairs, those of the leaves sometimes in part unbranched; fruit said to be
“‘vellow’? when mature.

Flabitat: Coast Mountains from San Francisco to Santa
Barbara, and possibly on the borders of Nevada.

California (Douglas, type [G]; Fremont, 3d Exped.
[G]; 589 Coulter [G]; Brandegee [G]) (Santa Maria
Mountains, Mrs. Watts [A]);

Contra Costa County (Oakly, March, 1900 [P]);

Fresno County (Alcalde, Mrs. Brandegee [A]) (Huron,
Miss Eastwood, May, 1893 [A]);

Monterey County (Monterey, Fremont, 3d Exped., Jan.
31, 1846 [N]; 633 Brewer [G]; 633 Guirardo [U]; Bran-
degee [A]) (Pacific Grove, Tidestrom, Jan., 1893 [U],
a stout form with ovate leaves, 3-4 cm. long, same as
Guirardo’s plant) (Nacimiento River, Miss Eastwood,
May, 1897 [A]) (Los Burros Trail, Miss Eastwood, May,
1897 [A]);

San Benito County (New Idria, Miss Eastwood, May,
1893 [A]):;

San Francisco County (Lone Mountain, 14 Kellogg &
Harford [A});

Santa Barbara County (San Rafael Mountains, H. C.
Ford, 1887 [G]) (Howard Cafion, Miss Eastwood, May,
1896 [A], albino) (Duford’s Ranch, Miss Eastwood, May,
1896 [A]) (Sespe, May, 1897, F. W. Hubby [P]).

Nevada (Carson City, Anderson, 1865 [G], but an error
in the locality label is probable).

AN ACCOUNT OF THE SPECIES OF PORPHYRA
POUND ON THE PACIFIC COAST OF
NORTH AMERICA.

BY HENRI T. Aw HUS:*

CONTENTS.

PLATES XX-XXII.

IE RPELISTOR Vier crsiatei near achieve stot aellvetekot ar teers ouniersaotiak [= fake ne c2-6 81 ol. csSsel 173
MS MORPHOLOGY e.. «is sissies stom eile miedo a orca site sel aoe siete ede sie cs 175
WM LD ISTREBUMION se: tape) veoetyel cate che rere eter vaes © ohm ei as ole oleh ee oi 192
Vee DESCRIPTION OFS SPECIES) eyec orer ieee raat reek teisie acer ie See wile 195
KEvy TO THE PACIFIC COAST eee sia gS Sea ee eae 195

Wats IS CONOMIGUUSESE s2 erty cnn le reer Eras oe ete a ie eae we eve tj 230
Walls, (ANY REA SGD Sine ate crac a pen ante be a ay heme ee 231
LITERATURE AND EXSICCAT Cae. By aafoTa feLecote Moan 5 a8 “op nya ehatists tanec Sanelsie too sie 234
PREPEANATION( OF ATES); 5st We Aa estes ohm OEE aes chee eal eine Soe aes 236

I. HisrTory.

FRom a systematic point the genus Porphyra has been
scantily dealt with. Created in 1824, by C. A. Agardh, to
contain those species of ‘‘ V//va’’ which possess a red color-
ing matter, it has since been fully treated by but two
authors—J. G. Agardh (1882) and J. B. de Toni (1897).
The former author was the first to distinguish between
monostromatic and distromatic species. This idea was car-
ried still farther by Kjellman, who in 1883, in his ‘‘Algz
of the Arctic Sea,’’ distinguished between the genus Por-
pryra and the subgenus Diploderma, the latter to include
all distromatic species.

The name Diploderma was changed by de Toni (1897)
to Wildemania.

L. Kolderup Rosenvinge (1893) used the name Por-
phyra for both monostromatic and distromatic species,
retaining Dip/oderma as a subgeneric name. This appears

*Contributions from the Botanical Laboratories of the University of Cal-
ifornia and presented in partial fulfillment of the requirements for the degree
of M.S., May, 1899. Prepared under the direction of Professor W. A. Setchell.

[173] December 14, 1901.

174 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

to be a step in the right direction, since, as Rosenvinge
points out, the distromatic species are frequently monostro-
matic in portions of the frond. The writer has often found
specimens of P. minzata, P. tenuissima and P. abyssicola
which were partly monostromatic, partly distromatic in the
purely vegetative portions of the fronds.

Even though the distromatic character is far more con-
stant than was originally supposed, it seems to the writer
there exists no sufficient reason to subdivide the genus,
since the plants agree so entirely in habit and external
characters as to be readily recognized by the collector as
belonging to the genus Porphyra.

Other contributors to our knowledge of the genus Por-
phyra are Foslie (1890) and Strémfelt (1886), both of
whom chiefly investigated the waters of Northern Europe.

The species of the genus Porphyra in Asiatic waters
have been little studied. Suringar (1870) mentions P. vu/-
garis as occurring in Japan. Afterwards Kjellman (1897)
studied the species of Porpfhyra of the coast of Japan, and
in his paper enumerates six new Japanese species.

One of the first to mention Porphyra in America was
Ruprecht (1852). In an account of a species of Phyllo-
spadix collected by Wosnessenski near the mouth of the
stream Slavjanka (Russian River!), he refers to a para-
sitic species of Porphyra, occurring on the blades; ‘‘gegen
die Blattenden zu, finden sich kleine parasitirende Exem-
plaren von Porphyra.’’ From the fact that P. nazadum
And. is the only species of Porphyra occurring with any
regularity on Phyllospadix, it is more than probable that
this is the species referred to.

Harvey (1858) in his account of American Algae, men-
tions but a single species, P. vulgaris, found on both the
east and west coast. He is inclined to unite P. vulgaris
Ag., P. lacintata Ag., P. purpurea Ag., P. linearis Grev.,
and P. amethystea Kiitz. under the name P. vulgaris. Later,
Farlow (1881) described P. dacinzata Ag. as a cosmopolitan
species, and mentions P. J/eucosticta Thur. as_ probably
occurring in New England, but not yet certainly observed.

Bot.—Vot. II.] HUS—PORPHYRA, 175

Collins (1882) reported P. /aciénéata from the east coast, and
later (1884) P. leucosticta and P. miniata.

J. Agardh (1882) reports P. coccinea (P. natadum?) and
P. perforata from the Pacific Coast. Since then Dr. Ander-
son (1892) has added two new species to the number,
P. natadum and P. nereocystts.

Miss Tilden in Century III of ‘‘Algz of North America’’
has distributed four species of Porpiyra from the Pacific
Coast under the names P. miniata, P. natadum, P. leuco-
sticta, and P. laciniata.

Up to 1898, however, there had been reported from the
west coast of North America but four distinct species.

In the winter of 1897 Professor Setchell suggested that
the writer investigate the peculiar base of P. natadum
And., but specimens of Porfhyra gathered on collecting
trips, and a consideration of those in the herbarium, showed
such a variety of morphological and anatomical characters
that the desirability of a collection and investigation of the
species of Porphyra occurring on this coast became appar-
ent; itis the results of these investigations which are set
forth in the following paper.

Il. Morpuouocy.

The shape of the fronds of the various species of Porphyra
is exceedingly variable, but that of most of them can be
reduced to the elongated type of frond. ‘The variation is
between linear and oblanceolate, and nearly all the species
mentioned in this paper exhibit both characters at various
periods of their existence. A striking exception to this rule
is Porphyra perforata {. lanceolata, which, as a rule, is
constantly linear; yet there are specimens in our herbarium
which are decidedly lanceolate. On the other hand, I
believe there are but few mature specimens of P. fenuzss7ma,
if any, which ever exhibit a linear form. The nearest
approach the plant makes to the linear form is when young;
it then-possesses an oblong outline.

Closely connected with the shape of the fronds are their
length and width. These three characters seem to be

176 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

closely related and are evidently determined by the same
conditions, viz., age, zone, and locality. It would be diffi-
cult to say which of these three agencies exerts the greatest
influence on the plants. All seem to be of equal importance.
A very good illustration is yielded by P. perforata. While
young specimens of P. perforata (three to four centimeters
long) are usually irregularly expanded, with a tendency
towards the orbicular, we notice that very soon a change in
the shape of the frond takes place. Asa rule, specimens
five centimeters or more in length already show the type,
lanceolate with undulate margin; but older specimens, and
this applies particularly to those found in the lower part of
the litoral zone, and in the upper part of the sublitoral
zone, possess a great width and are frequently much lobed
and laciniate. Especially those plants which grow on the flat
surfaces of rocks, for instance on reefs, show a marked iso-
diametric development. But if the plant grows pendant from
an overhanging rock, it develops the elongated type of frond.
Another condition, and dependent upon locality, is the
movement of the water. Plants growing where they are
continually exposed to the wash of the waves, back and
forth, and from side to side, show far less marked longi-
tudinal development than those which are exposed to the
movement of the water in but one direction. This is very
well illustrated by P. xazadum And. f. major. The author
had an opportunity to observe this plant growing on Zostera
in the lagoon at Bolinas, Marin County, California. The
lagoon, which is long and narrow, was, in the summer of
1899, protected by a high bar, so that at the rise of the tide
the water flowed in very regularly for a number of hours,
till the turn of the tide, when it flowed out as regularly.
The blades of Zostera and the fronds of P. nazadum were
bent in the direction taken by the water and the latter showed
a marked elongation, so great, indeed, that the writer felt
entitled to consider them a special form of the species,
since they were fully twice as long as the blades of P.
naiadum growing on Phyllospadix, and since there existed
some other minor differences as well. The latter form was
consequently designated as P. nacadum f. minor.

Bot.—VOL. II.] HUS—PORPHVRA. ae

An interesting instance of great length attained by
exposure to the motion of waves in one direction was found
in a specimen of P. perforata f. lanceolata, which grew on
a rock buried in the sand of the gently sloping shore of the
Presidio, San Francisco, California. This specimen attained
a length of 325 centimeters, which to the author’s knowledge
is the greatest length ever attained by any specimen of this
species of Porfhyra. This extraordinary longitudinal devel-
opment (the average length is but thirty to forty centime-
ters) was evidently due to the plants being stretched out at
full length every time a wave rolled in or went out.

P. nereocystis, growing in three to five fathoms of water
on the stipes of (Vereocystis liitkeana, often attains a great
length, specimens of over three meters in length having
been collected at Monterey. A specimen of P. variegata
collected at Santa Cruz by Dr. Anderson measured seventy-
nine centimeters. On the other hand, we have found a
fertile specimen of P. perforata but two centimeters long.
But the plant which in its adult stage is the smallest of all
Pacific Coast species of Porphyra is P. natadum, which
often bears fruit when but one centimeter long.

The width of the fronds also varies considerably. While
the writer has measured specimens of P. nereocyst7s which
were fully forty-nine centimeters in diameter, some mature
specimens of P. perforata f. lanceolata collected by Dr. W.
A. Setchell at Monterey, California, measured in their
widest part but twenty-five hundredths of a centimeter.

In regard to the part age plays in the determination of the
shape of the fronds, it must be said that while the younger
plants asa rule possess the elongated type of frond, the
older plants generally have a greater width. A microscop-
ical examination reveals the fact that the divisions of the
cells of the younger fronds are usually parallel to each other
and at right angles to the longer axis of the frond. In the
older plants, where the development is more isodiametric,
we can readily recognize more or less isodiametric groups
of cells, which evidently arose from a single cell. Of
course, environmental conditions have much to do with this.

178 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

The base in Porphyra varies from cuneate to cordate, or
even umbilicate, and is sometimes cucullate, as in P. nerveo-
cystts. These various forms of base depend just as much
as size, etc., upon age, zone, and locality. An umbilicate
base is found on the fronds of plants growing on flat sur-
faces (P. laciniata £. umbzlicalis); one is liable to meet
with a cordate base in the older fronds; while a cuneate
base is found on the fronds of plants which grow in exposed
places.

In the genus Porphyra, we may distinguish between two
kinds of attachment, the one cushion-shaped and paren-
chymatous, the other discoid and rhizoidal. The latter
form of base has been amply discussed and illustrated by
Bornet and Thuret (1878), and it will suffice to say here
that the cells in the immediate neighborhood of the base
produce rhizoid-like projections, in thickness from two-
tenths to one-tenth the diameter of the cell, which grow
down through the jelly between the cells and the cuticle.
That these hyphe actually grow, and that the older the
plant grows the more of these projections are produced, is
demonstrated by treating the base of a young specimen of
Porphyra perforata with Schultze’s macerating fluid. This
dissolves the jelly, and the weight of a cover-glass crushes
the preparation sufficiently to show the details. From each
of the thick-walled cells near the base, a hypha may be
seen growing out, which may be longer or shorter, some
being even but a few microns in length. Their course is
more or less direct. Most of them extend down to the
substratum. Consequently, though the frond about one
centimeter above the base is normal, the part lower down is
very much thickened by an ever increasing number of these
projections, which finally form a dense network, in which
it is impossible to trace the individual hyphe. According
to Agardh (1882), these hyphz possess no septa, at least,
he has been unable to see them: for he observes: ‘*‘ Hec
fila radicantia Porphyrz mihi semper inarticulata obvener-
unt, * * * .’’? They are long, slender, tapering threads,
averaging one to two microns in thickness. Of course, the
longer they become, the less evident their tapering nature

Bor:—VOtz. II.] HUS—PORPHVRA. 179

is. In fact, the diameter of the larger hyphe appears to be
the same for the whole length. While the majority agree
in this regard, if we follow them down to the base a differ-
ence soon becomes apparent. Some of the hyphz come to
an abrupt end, their diameter remaining constant, the con-
tents remaining hyaline and parietal, and no septa being
present; but others show a greater or lesser increase of
thickness at the tip fora greater or lesser length, and a few
even branch or at least show indications of branching
(Pl. XX, figs. 7-10). In some of these, septa have been dem-
onstrated. The swollen ends contain protoplasm. Whether
these ends are to be considered as haustoria, and whether
the hyphe enter the cells or intercellular spaces of the host-
plant, or whether they merely adhere to the substratum,
are questions to which the author can give no definite reply.
In sections of the base of a specimen of Porphyra perforata
which grew on Phyllospadix, it was impossible to deter-
mine the course of the hyphae. The same was true for
plants growing on wood. Young specimens of Porphyra
perforata growing on barnacles were treated with one per
cent. nitric acid; but after dissolving the calcium salts, it
was impossible, partly owing to the confusing mass of par-
asitic algz which flourished in large numbers on and in the
shells, but especially to the presence of chitin, to follow
the hyphe in their course.

The ever increasing number of hyphz adds considerably
to the thickness of the frond, the latter within one-half a
centimeter of the disc often measuring two hundred microns
or more, while the strength must be increased a hundred-
fold. The cells which give rise to these hyphe, especially
those situated more towards the disc, are but imperfectly
seen, even in a specimen not quite four centimeters long,
being obscured by the hypha-like projections which sur-
round them. It is easy to conceive that being thus partly
excluded from the light these cells should undergo some
change. They lose their purplish color, have yellow-
brown cell-contents, and their walls are considerably
thickened.

180 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

At first it seemed exceedingly improbable to the author
that the cells to which the function of the attachment of the
frond was delegated should finally produce fruit; but a
knowledge of the fact that the formation of spores in these
cells was observed by Bornet and Thuret (1878) led him
to make a more careful study of the basal cells. Up to
this time, however, the author has been unable to demon-
strate a single cell which both emitted a hyphal thread and
bore fruit, though the oldest obtainable specimens were
investigated. It must therefore be concluded that if ever
the contents of these cells are transformed into spores, this
must be but rarely the case.

The areolate, lighter colored portion of the frond, about
one centimeter in diameter, directly surrounding the attach-
ment found in all specimens of Porphyra as known to the
writer, with the exception of P. nazadum, probably finds its
reason in two causes. The first is the partial loss of color
of the cells near the base, caused by decreased activity
owing to the large number of rhizoid-like projections which
separate them from the surface of the frond. The other '
cause may be looked for in the large number of rhizoid-like
projections with hyaline walls, resulting in an increased
thickness and consequently increased density of the lowest
part of the frond.

Some of the species of Porphyra are slightly stipitate.
On this coast only P. /eucosticta shows this to any marked
extent. The stipes appear to possess the same structure as
the discs.

A cushion-shaped base is, as far as the writer is aware,
found in but a single species of Porphyra, viz., P. natadum
And., a species peculiar to the Pacific Coast. P. natadum
has been found growing on eel-grass, either on Phyllo-
spadix in exposed places in the sublitoral zone or on Zos-
tera, sheltered, in lagoons.

On examining the blades of Phyllospadix during the win-
ter months, we find here and there small reddish brown,
cushion-shaped growths, which to the superficial gaze
appear like colonies of diatoms. Continued observation
shows a gradual increase in the number of these wart-like,

Bot.—Vot. II.] HUS—PORPHYRA. ISI

more or less flattened structures. Finally, they cover the
blades of eel-grass in such large numbers that they grow
next to and over each other, and lose their natural hemi-
spherical shape, obscuring the normal color of the eel-grass,
and giving a rough appearance to the blade.

As the season advances, examination with a lens shows a
greater or smaller number of short, blunt protuberances
issuing from the wart-like growths. Under the microscope
they appear to be composed of a number of cells placed
end to end. Further observation demonstrates the fact
that these cells, by division in two planes, give rise to a
monostromatic frond. From this it is but a step to estab-
lish a genetic connection between the hemispherical struct-
ures on eel-grass and the fully grown fronds of P. nazadum
on the same host-plant. Evidently we have the prothalloid
form of P. naiadum before us. This was already suspected
by Dr. Setchell when he called my attention to the matter.

The prothallium, when young, consists of but a single
layer of cells, placed side by side on the blade of the eel-
grass. For a certain length of time these cells continue to
divide in a single plane. After that, division in the second
plane begins to take place, gradually giving rise to the
wart-like growths referred to above. In section they appear
to consist of layers of large, thin-walled, parenchymatous
eels (PIP XXII fis. 19). The célls of the central layers
possess ordinary cell-contents, but only a very small chro-
matophore. The two or three outer layers are made up of
slightly smaller cells, and possess a large chromatophore.
The cells of the layer adjacent to the surface of the blade
of the host-plant also contain a large chromatophore. Upon
these latter cells evidently devolves the function of attach-
ing the prothallium to the eel-grass. Each cell is extended
so as to form a short, sharp, unicellular rhizoid. The
writer has been unable to determine with any satisfaction
whether these rhizoids entered the cells of the host-plant or
not. Careful sectioning and staining has failed to reveal
anything of the kind. But in material which had been
shrunken by reagents, only the rhizoids at the periphery of
the cushion-shaped base were attached to the eel-grass, the

182 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

central part of the base having shrunk away, exposing the
rhizoids. From this it would appear as if the rhizoids
entered, to a slight extent only, the cuticula of the host-
plant.

The young fronds arise from the cells of the external
layer of the prothallium. Evidently any cell may give rise
to a frond by a division in one plane, in advance of the
surrounding cells which form part of the external layer of
the cushion-shaped base. When in this manner a filament
of some five or six cells has been formed, the cells of the
filament begin to divide in two planes, thus giving rise to a
membranous frond, the length of which when fully grown
seldom exceeds six centimeters.

The number of fronds a prothallium may give rise to
appears to be indefinite, every cell of the outer layer of
the base seemingly being capable of producing a frond. A
frond may be formed during the first stage of the existence
of the prothallium, cases having been observed where frond
formation evidently took place when the prothallium was
but two cells in thickness. The formation of a frond by a
cell of the outer layer does not mean cessation of growth
for the other cells of the outer layer, since frequently a
frond may be found, the base of which lies in a depression
of the prothallium several cells deep.

An attempt was made to ascertain, if possible, if when
the frond has reached a certain size, the cells of the frond,
in the neighborhood of the base, produced rhizoid-like pro-
jections such as are found in the corresponding cells of
other species of Porphyra. But an examination of the
bases of a large number of mature fronds of P. nazadum
failed to reveal these structures.

When Porphyra naiadum occurs on Zostera, it produces
the same wart-like growths, but only on the extreme mar-
gins, not on any part of the surface of the blade, and they
appear to be smaller.

Other species of Porphyra probably occur but seldom on
eel-grass. The only species found by the writer to occur
occasionally on the same host-plant were P. perforata,
P. laciniata, and P. abyssicola.

Bot.—Vot. II.] HUS—PORPHYRA. 183

The value of the prothalloid base lies evidently in the
power to form a large number of fronds rapidly, which in
the production of new fronds in case of accident is clearly
of great importance.

As far as the author is aware, P. nazadum has never
been found growing on any substratum other than eel-grass,
nor has a cushion-shaped base such as here described ever
been found in any other species of Porphyra. The nearest
approach to such a description is that of the base of P.
coccinea J. Ag., such as is found in Agardh’s ‘Till
Algernes Systematik’’ VI (1882). But judging from this
account, the base is hollow, being formed by the involution
of the edges of the young frond. When the frond grows
older the base finally flattens out.

It is worthy of note, that in connection with P. coccinea
Agardh mentions a Porfhyra occurring in large numbers
on the Pacific Coast; it grows on seaweeds (!), and judging
from the description given might possibly be P. xazadum.

The color of the fronds of the different species of Por-
pryra is such as to lead one to place the genus among the
Floridee. But the color is far from being constant. A
hundred different shades may be met with, for even the
color of the different fronds belonging to one species
varies; so that an attempt to describe a species by the
color would be futile. While the color of one species (P.
tenuissima ) is, as a rule, a delicate pink, others are a bright
red or even crimson, as P. abyss¢cola. P. laciniata exhibits
a decidedly purple color, while P. ferforata appears mostly
yellow-brown. The frond of P. variegata is crimson when
sterile, while when fruiting it acquires the beautiful varie-
gated appearance indicated by its name.

An important fact is that the color of herbarium specimens
generally changes. This was most notable in P. perforata.
Specimens which when collected had a yellow-brown tint
generally became a deep blue-purple. Some of the fronds
of P. nereocystis underwent a change in the herbarium,
while others retained the original dull brown-red color.

Especially did specimens which were rough-dried and
afterwards soaked in fresh or salt water for mounting

184 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

purposes seem to change color on drying. A similar change
was noted in sheets of Asakusa Nori (see Economic Uses.)
which were of a yellow-brown color when bought in San
Francisco, but having been kept for several months ina
closed paper box turned purple. A sheet of Asakusa Nori
which had accidently been left partially exposed to the air
and light for about a month, showed after that time a bril-
liant violet coloring in the exposed portion, while that part
of the sheet which was not exposed retained its original
yellow-brown tint.

As far as can be judged from the statements of various
authors, as well as from our own observations, it appears
that the color of certain species varies according to the
locality. This is well illustrated by P. /eucostzcta Thur. It
seems that the European specimens of P. Jeucosticta are of
a distinctly yellow color when fresh, and when dried a deli-
cate purple-pink tint. But the specimens of P. /eucosticta
found on the Pacific Coast, if gathered early in the season,
are deep-pink, becoming lighter as the season advances.

From the above, it will be seen that specimens of Porphyra
should, whenever possible, be mounted fresh; that even
then the color is of small value from a systematic point of
view; and that it is most undesirable, in fact, impracticable,
to use the color of a frond as the criterion for the species,
though it is often of great value in indicating its position.

It remained for J. G. Agardh to call attention to the
monostromatic and distromatic nature of the fronds of the
different species of Porphyra. These characters have
been found to be absolutely constant in all species, with the
exception of those belonging to what we may call the
‘‘miniata’’? group, which includes besides P. mznzata, P.
amplissima, P. tenuissima, and P. abyssicola. The first
three species are, as a rule, distromatic, though places may
be found which exhibit a monostromatic character, especially
towards the edges. Fronds of P. abysszcola, which species
was first described by Kjellman as monostromatic, have been
found by Rosenvinge and by the author to sometimes exhibit
a distromatic character, either through the whole frond or
in portions of it.

Bot.—Vot. II.] HUS—PORPHYRA. 185

Kjellman applies the name Dzploderma to all distromatic
species, but the above mentioned results lead the writer to
agree with Rosenvinge in applying the name Porphyra to
all members of the genus, while retaining D7floderma
(Wildemania de Toni) as a subgeneric name for the di-
stromatic fronds, the more so as in habit and external char-
acter the monostromatic and distromatic species agree in all
respects.

Two sources of confusion in determining the number of
layers in the frond exist. The first is the age of the frond,
the second, the formation of reproductive cells. Young
fronds of distromatic species are frequently monostromatic.
This monostromatic character persists in the vegetative por-
tion of the frond even after the fruit has been formed (P.
abyssicola) (Rosenvinge, 1893). The same results were
obtained by the author in his observations on P. adysszcola,
but he cannot confirm Rosenvinge’s statement, that in the
distromatic forms the inferior portion of the thallus is com-
posed of a single layer of cells. ‘* Dans les formes distro-
matiques, du reste, la partie inférieure du thalle est composée
d’une seule assise de cellules’’ (Rosenvinge, 1893, p. 84).
On the contrary, if we do not consider those of P. abysszco/a,
but few, if any, distromatic specimens were found which
were monostromatic at the base. As a rule, whenever a
frond of a distromatic species was partly monostromatic, the
monostromatic portion was found in the region of the tip at
the edge.

Monostromatic species practically become distromatic as
soon as the reproductive bodies begin toform. ‘The second
division of either the antheridium or sporocarp-mother-cells
takes place ina direction parallel to the surface of the
frond, and in this manner gives rise to two layers of cells
which are often difficult to distinguish from purely vegetative
cells; so that in deciding their nature, the cells of the sur-
rounding tissue must be taken into account.

The thickness of the fronds of the various species of
Porphyra is more or less variable, and though not an abso-
lutely specific character, taken in conjunction with other

186 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

characteristics it is of great value, especially in indicating
the position of a sterile frond.

As a rule, the thickness of the fertile part of the frond is
much greater than that of the sterile part. This seems to
be due to the swelling of the jelly surrounding the repro-
ductive bodies at the time of ripening. Also, wherever we
have a dicecious frond, or where the antheridia and sporo-
carps are born on separate portions of the plant, the thickness
of the antheridial portion is greater than that of the sporo-
carpic portion. This would bear out our hypothesis that
increased thickness is due to the swelling of the jelly, since
there exists a larger number of partitions consisting of jelly
between the antherozoids than between the carpospores.

P. variegata exhibits a very marked thickness and strati-
fication of the jelly-walls surrounding the vegetative cell.
The walls of the vegetative cells of all fronds of this species
examined were of this nature; consequently the author feels
entitled to consider this a diagnostic character, the more so
as the only other Porphyra which possesses much thickened
cell-walls differs widely from P. variegata in habit and
external characters.

The plant here referred to is P. perforata f. segregata.
The walls not only of the vegetative cells but also of the
reproductive cells are much thickened, especially those
produced by the first reproductive division of the antheri-
dium. In fact, the upper and lower groups of antherozoids
are noticeably separated, which lends the cross-section of
the antheridial portion of the frond a most characteristic
appearance.

The outer jelly-walls of nearly all the fronds examined
were infested with bacteria which formed narrow lines per-
pendicular to the surface of the frond, reminding one of the
canals formed by the ‘‘spermatium”’ at the time of the fer-
tilization of the ‘‘procarp’’, as described by Berthold (1882).
These ‘‘ canals’’ were found in the jelly surrounding the
vegetative and antheridial cells, as well as in that surround-
ing the sporocarps.

In regard to the shape of the vegetative cells, it may be
said that while in the monostromatic fronds the cells are

Bot.—Vo-. IL] HUS—PORPHYRA. 187

either cubical or more frequently higher than broad, the
vegetative cells of the fronds of the distromatic species
vary, as a rule, from cubical to broader than high. Excep-
tions to this rule are found in P. xereocysizs, a monostromatic
species which sometimes possesses cells which are broader
than high, and in P. varzegata. While the vegetative cells
of the younger sterile fronds of the latter species are usually
square, the vegetative cells of the older fertile fronds are
much higher than broad and often have a fusiform appear-
ance. Judging from the fact that the vegetative cells are
found between the reproductive cells, it is suggested that
the shape of the former is due to pressure exerted by the
reproductive cells, which before dividing gorge themselves
with protoplasm, and when fully ripe swell to an abnormal
size, owing to the partial dissolution of the jelly partitions
separating the individual spores.

During the study of the species of Porfhyra of the Paci-
fic Coast, the fact gradually made itself felt that the repro-
ductive bodies are of the greatest diagnostic value, and that
habitat, color, and thickness of frond can only be used to
determine species in connection with the number of divi-
sions of the antheridia and sporocarps.

Since the object of this paper is merely to give a syste-
matic account of the species of Porphyra of western North
America, and it is not designed to throw light on the sexu-
ality or nonsexuality of the genus Porfhyra, the author uses
the terms sporocarp and antheridium merely to indicate the
larger and smaller bodies, which by some are believed to
play a part in sexual reproduction, without necessarily
ascribing a sexual character to these bodies. The same is
true for the asexuality of the monospores. But it must be
said that in no case even the slightest indication of sexuality
has been observed, though many sections of sporocarps in
all stages of development were examined. Neither has the
author been able to observe an amceboid movement of the
liberated carpospores, nor flagelliform appendages to the
bodies contained in the antheridia, nor any movement on
the part of these bodies, though observations were made to

188 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

determine this point if possible. The results obtained by
the writer therefore agree rather with those obtained by
Reinke and Bornet than with those of Berthold and others.

While some species of Porfhyra are moneecious, others
are dicecious. In some cases fronds have been found
which evidently approached closely to those of a mone-
cious species, but which differed from it in being dicecious
and in exhibiting some slight differences of habit, etc.
Such fronds have been referred to the original species but
were separated from it under a form-name.

We may distinguish two forms of moncecious fronds. In
the one, antheridia and sporocarps are separated in patches.
These patches are usually sharply defined, especially at the
edges of the frond, owing to the lighter color of the ripe
antheridia and to the more intense color of the ripe sporo-
carps. This arrangement is usually met with in the fronds
of the monostromatic species. In the other form, the
antheridia and sporocarps occur side by side, so that the
frond has a uniform color. This occurs in the distromatic
species, and more particularly in what the writer has found
convenient to designate the ‘‘mznzata group,’’? which in-
cludes P. ampilissima, P. miniata, P. tenuissima and P.
abyssicola.

Vegetative cells are frequently found mixed with the spo-
rocarps, among both monostromatic and distromatic species,
and in larger or smaller patches. While the distromatic spe-
cies show this constantly, the monostromatic species often
fail to show these vegetative cells among the sporocarps.
Hardly ever has the author found any vegetative cells
mixed in with the antheridia of the monostromatic species.

Among the sporocarps there appear frequently bodies
which by various authors have been called monospores.
They seem to be formed by the arrest of division in one of
the segments of the sporocarp or by one of the vegetative
cells lying among the sporocarps. What differentiates them
from the vegetative cell proper is a greater thickness of the
cell-wall and a larger amount of protoplasm. The chromato-
phore may be seen lying in or near the center of the cell.

.
,

Bot.—Vot. II.] HUS—PORPHYRA. 189

These monospores can easily be distinguished from the dead
vegetative cells lying among the sporocarps. The dead
cells possess likewise a thick wall but apparently contain
yellowish, homogeneous, highly refractive cell-contents, in
which no chromatophore can be discerned.

Whether monospores are sexual or not, or whether they
possess any reproductive power, the author has, notwith-
standing a series of careful experiments, been unable to
determine.

The reproductive bodies are usually first formed at the
margins and gradually spread over the whole frond. Bornet
even observed spores in the basal cells of P. /aciniata.
Under the microscope we can trace the various stages of
division from the original vegetative cell to the fully ripe
sporocarp. This is especially easy in the species where
antheridia and sporocarps occur side by side in patches.
Observation shows that each vegetative cell gives rise to a
single sporocarp. The sporocarp by two more or less
simultaneous divisions at right angles to each other and to
the surface of the frond finally consists of four segments.
In some species division proceeds no farther, and four carpo-
spores are the result; but in other species, where the fully
ripe sporocarp contains more than four carpospores, the
cruciate division is followed by a division parallel to the
surface of the frond, giving rise to eight segments, which
by further cruciate division perpendicular to the surface of
the frond in each of the resulting cells may give rise to
thirty-two carpospores.!

In the formation of the antheridia, starting from the vege-
tative cell equivalent to the mother-cell of a sporocarp, there
first takes place a vegetative cruciate division perpendicular
to the surface of the frond, which gives rise to four anther-
idial cells. Consequently, owing to this additional vegetative

1The author understands under “cruciate” division, two divisions in different direc-
tions, at right angles to each other and to the surface of the frond, and which are simul-
taneous or nearly so. Under ‘‘transverse”’ or ‘‘parallel” division the writer understands
a division parallel to the surface of the frond. Though these divisions are, of course,
but seldom if ever strictly parallel or at right angles, the use of the terms “cruciate”
and “‘parallel’’ is a great convenience, doing away with a lengthy explanation.

(2) December 19, 1901.

190 CALIFORNIA ACADEMY OF SCIENCES. [PRroc. 3D SER.

division of the antheridium-mother-cell each sporocarp cor-
responds to four antheridia.

The author distinguishes between a vegetative division
and a reproductive division by the aid of the partition walls
laid down by these divisions. A jelly-wall formed after a
vegetative division is, as a rule, much thicker than one
formed after a reproductive division. Furthermore, a wall
of the latter kind dissolves when the frond is fully ripe, so
that the reproductive bodies become arranged more or less
irregularly; something which is very clearly shown in the
antheridia of P. /eucosticta.

The first reproductive division of the antheridium is
parallel to the surface of the frond, corresponding to the
first transverse division of the sporocarp, and is followed by
a cruciate division in both segments. In fact, parallel and
cruciate divisions alternate until the number of antherozoids
peculiar to the species has been formed. The only differ-
ence, therefore, between antheridia and sporocarps lies, if
we do not consider their origin, in the larger number of
divisions which the former undergo.

Though the manner of division of antheridia and sporo-
carps is fairly constant, yet a large number of variations
take place. The most frequent among these is the direc-
tion of the last division in either antheridia or sporocarps,
which is not necessarily parallel or perpendicular to the
surface of the frond, but is often oblique, and is occasion-
ally omitted altogether in some of the segments of the spo-
rocarp or antheridium. Rarely an additional division takes
place in some or all of the segments of the sporocarp or
antheridium.

Cases have been met with, where the vegetative division
of the antheridium-mother-cell is not as evident as usual,
and the whole vegetative cell apparently becomes an
antheridium, so that four times the usual number of anther-
ozoids are formed.

Occasionally the first cruciate division of the sporocarp-
mother-cell is vegetative instead of reproductive, and only
one-fourth the usual number of carpospores are found.

i dC ee” See a.

Bot.—VOL. IT.] HUS—PORPHYVRA. I9gI

With the material at the disposal of the author, he has
been able to distinguish between four types of division of
the reproductive bodies, the differentiation into types being
based upon the number of antherozoids and carpospores
produced. The first is the Porphyra perforata type. Here
thirty-two carpospores are produced, the sporocarp under-
going first a cruciate division, followed by a parallel division

in each of the segments, which is again followed by a cru-
ciate division of each segment. If we represent a vegeta-
tive cell by a cube, and indicate the two horizontal lines
respectively as a and 64, and the perpendicular as c, we

can, by the aid of the formula 32 (“ 7 =) readily form

a diagram such as is represented in fig. 25, which shows
the manner of division.

The antherozoids contained in each antheridium number,
in the P. perforata type, 128, and are formed by alternat-
ing parallel and cruciate divisions, the first division of the
antheridium being parallel to the surface of the frond.! The
manner of division may be represented by the formula
128 (4 ze, 5) (fig. 28). To this type belong P.
perforata, P. perforata £. lanceolata, P. perforata f. segre-
gata and P. nereocystis.

The second is the P. /eucosticta type. Here eight carpo-
spores arise from a cruciate division of the sporocarp-
mother-cell, followed by a parallel division of the four
segments. This may be represented by the formula

CE
8 (<, a ) (fig. 24).

The antherozoids contained in each antheridium of the
fronds of species belonging to the P. /eucosticta type num-
ber sixty-four. They are formed by first a parallel division
of the antheridium, followed by a cruciate division, after
which a second parallel and a second cruciate division take

1It must be remembered that an antheridium is but one-fourth as large as a sporo-
carp, the first division of the antheridium-mother-cell, which corresponds to both a
vegetative cell and a sporocarp-mother-cell, being vegetative and cruciate.

192 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

place in each segment, so that the arrangement of the anther-
wa %b Cc

AS =)
(fig. 27). To this type belong P. leucosticata, P. laciniata,
and it may be predicted, judging from the carpospores,
that P. natadum also belongs to this section, the antheridia
in this species being as yet undetected.

Among the distromatic forms the remainder of the types
are found. First we have the P. amplissima type, with
eight carpospores and sixteen antherozoids, and the form-

en a %a %b c
ule 8 (< ee =) (fig. 24) and 16 ( an cee >) (fig.
26); then the P. mznzata type, with four carpospores and

ozoids is represented by the formula 64 (

eight antherozoids, and the formule 4 (=, “ c) (fiz, 22)

;
2 2
tata, P. tenuzssima and P. abyssicola.
Of the two other distromatic species which occur on the
Pacific Coast, but one form of fruit has been found, so that
they can hardly be brought forward as types.

and 8 = ae = (fig. 25). To this belong P. mzn-

III. DuisTRIBUTION.

It is almost impossible to obtain a correct idea of the dis-
tribution of the older species of Porfhyra, as frequently the
name P. /aciniata was applied to various species which since
have been separated fromit. However, the author believes
that it may be said with some degree of certainty that P.
lactntata occurs on the western shores of Europe, from the
Norwegian Polar Sea (71° N. lat.) to the Mediterranean
(40° N. lat.) and on the Atlantic coast of North America
from Greenland (67° N. lat.) to New Jersey (40° N. lat.).
It has never been authoritatively reported from the eastern
shores of Asia; for though older authors have mentioned
it, yet P. daciniata was not included by Kjellman (1897)
among the Japanese species, and he even expresses some
doubt as to its occurrence. On the Pacific Coast of North

EE

BoT.—VOL. II.] HUS—PORPHVRA. 193

America P. laciniata has been reported from Orca, Alaska,
from Yakutat, and from Amaknak Island (between 61° and
54° N. lat.). P. laciniata f. umbilicalis was reported by
Professor Setchell (1899) from the Pribilof Islands.

Porphyra leucosticta Thur. does not appear to possess
such a wide range as the species just discussed. It is found
on the Atlantic coast of England, Germany, and France,
and appears to be abundant in the Mediterranean. Collins
(1884) and Holden (1897) have detected it on the eastern
shores of North America, and while it may have a wide
distribution on the Pacific Coast, it has as yet been reported
from buta single locality, Monterey Bay, California (36° 45°
IN: lat: ):.

Up to the present time the members of the ‘‘mzndata’’
group, under which the author includes P. amplissima, P.
miniata, P. tenutssima, and P. abyssicola, have been reported
by European collectors only from the more northern latitudes
(60°-80° N. lat.). On the west coast of North America
these species have not been reported from so far north,
ranging between 36° 45’ and 60° N. lat.

P. amplissima was first found by Kjellman (1883) in the
Norwegian Polar Sea, and it has since been reported from
both the east and west coast of Greenland. Kjellman did
not detect this species in any of the localities visited by the
Vega Expedition. Since then it has been collected at Orca,
Alaska (60° 30’ N. lat.), and near Coupeville, Washington
(48°"10' N.. lat.).

P. tenuissima occurs on the shores of Norway, Iceland,
and Greenland. It has never been found in the Bering
Sea, but it has been collected at Sitka, Alaska (57° N. lat.).

P. miniata is met with on the coasts of Norway, and
attains its greatest latitude on the northwest coast of Spitz-
bergen (79° 49’ N. lat.). It is also found on the east coast
of Norway and in Baffin Bay, whence it descends to New
Foundland.

There exists some doubt in the author’s mind as to the
occurrence of the typical P. mznzata on the Pacific Coast.
Only an extensive collection of specimens gathered on

194 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

numerous expeditions along the coast can lead to a definite
conclusion. But the occurrence of a variety designated by
us as P. miniata f. cunetformzs is an undoubted fact. It
has been collected in the Gulf of Alaska (60° N. lat.), at
Coupeville, Washington, and as far south as Monterey Bay,
California (36° 45’ N. lat.).

P. abyssicola is reported by Kjellman and others from the
north coast of Norway and Russia and from Greenland.
What the author believes to be P. abysstcola has been col-
lected on the Pacific Coast at Whidby Island, Washington
(48° 10 N. lat.).

Five species of Porphyra appear to be peculiar to the
Pacific Coast. In some cases it is possible to ascribe a
reason for this. P. nereocystts, which as far as can be
judged selects WVereocystzs liitkeana exclusively as its host-
plant, is necessarily limited to the region of distribution of
this species of /Vereocystis. It has been reported from St.
Paul, Kadiak Island (57° 30 N. lat.), from Coupeville,
Washington, and from the Californian shores (33°40 N.lat.).

P. naiadum, growing on Zostera and Phyllospadix, seems
limited to the Pacific Coast. Though other species of
Porphyraoccur on Zostera, both in Europe and on American
shores, yet no case is known to the author where P. nazadum
was found growing on eel-grass in waters other than those
of the Pacific, where it extends from Coupeville, Washing-
ton (48° 10 N. lat.) to San Diego, California (32° 20 N.
lat.).

P. perforata, so closely allied to P. /acinzata, attains
nearly the same northern latitude as the latter species, but
extends far lower down the Pacific Coast. Of the two
varieties of this species the author has been able to find
only one, P. perforata f. lanceolata, at San Francisco and
at Monterey, while the other occurs from Washington to
Mexico (San Roque) (47° 30-27° 8%’ N. lat.).

P. variegata was first found by Kjellman at Bering
Island (Vega Expedition). Since then it has been reported
by various collectors along the Pacific Coast, from Whidby
Island, Washington, to San Pedro, California (48° 10’—33° 40’
IN. at);

Bot

.—VOL. II.] AUS—PORPHVRA. 195

P. occidentalis has been found in but a single locality,
Monterey Bay, California (36° 45’ N. lat.).

DISTRIBUTION OF THE PaciFic Coast SPECIES OF PORPHYRA.

10)4) peel ae) Beas) las! es sl ast flee vail ee

S/8)2/2/8/5/ 8/8) 8) 8) =

SPS S/S] FI Si S/S) S/S) 8
x x Behring Sea. 5

alee sale » | Gulf of Alaska. o
Vancouver, Whidby Island, | 2
* * * * * * * °
Seattle. ae
Pee San Francisco. 3
* | % x x |x | x | x Monterey Bay. aa
x | Santa Barbara. 5
x Be || 02) lh) 36 San Pedro, San Diego. a
% x | x» | New England.
x | « | West Europe.
« | » | Mediterranean.
| eM tee flees « | Greenland.
«| * | * | x » | Arctic Ocean and Spitzbergen.
IV. DESCRIPTIONS OF SPECIES.
Key TO THE PaciFic Coast SPECIES OF PoRPHYRA.

REEL OMG SHIM OMOSEROMNAMES 25.5.0) eis ys eis etnies raat le cee ieieo ne heat ws 2
PIC OMC SACs tr OMMAtEC LAs) st), tiie eoce ahebe catia! eee trite a ete beg maces ai
Fronds monostromatic or distromatic............ Ease GeO DUES Rear II

2. Base cushion-shaped, consisting of parenchymatous cells... P. maiadum
Base discoid, consisting of agglutinated, rhizoid-like cells............ 3

Bane nirty-two spores in: each SpOrocanp .24.2\iccek. covet eae es os ave ces 4
Mim hiespares snl each) SpoLrocanp as at orise ade ek Eee ass Setcle sid oheae aalers 6

4. Fronds moncecious..... SHIGE G GOB OO.CD A Sh DASHES MORE ORO aE AEE eae 5
PP RONGS CHOCCIOUS ee. ane ses vera ys Sate eens tae P. perforata f. lanceolata

5a rondsybrown-purple; 45-150 thick... 51.66 sccss sees cee. P. perforata
Fronds brown-purple, 60 thick, with thick partition walls. ............

P. perforata f. segregata
Fronds red-purple, 25-60 thick, with thin partition walls. . P. zereocystis
SM PRONG MONCE IOUS. sae. casera ee aire NG tase bea enes P. leucosticta

IE OMG Ss GIG ClO US tan we arte rr tat arerr eee ren) a eases ea ey ee P. laciniaia

196 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.
%. - BEONGS GIGEGMIGHS 6 64 sc8 wise dae ds ors ue mae iseypae eee oe hee oes 8

Prends apparentiy GueenOns oe. aor asa es aeeee ie Dem eee eaeee Io

8. Sixteen antherozoids in each antheridium............... P. amplissima

Eight antherozoids in each antheridium................--. 22-20-00 9

9.. Fronds 254 thick, except ab DASE: «65.0203: «skeet ones eee P. tenuissima

Fronds 50-757 Mitek oe oo os Sb cu sicnss sacs uwes P. mintata f. cunetformis

LO? }SPLONGSEOO—220/faTHICKe: Scene Sue cciclease ec eee tee ees P. variegata
rOnGs m5 56 tulekKe ser wice aioe oe sco ole tals cielo oe eieiee ee P. occidentalis

Tis" xP ONES 25/0 tiniG kes siete Ss fA ceky cae eee ee oboe eee P. abyssicola

1. Porphyra laciniata (Lighi/.) Ag.
Systema Algarum, p. 190, 1824.

Porphyra laciniata (LicutF.) C. A. AGARDH, Systema Algarum, p. 190,
1824; Icones Algarum Europzarum. Tab. XXVII, 1828. Harvey,
British Marine Algz, p. 216, 1849. JANCZEWSKI, Etudes Anat. sur les
Porphyra, p- 352, 1873. BORNET et THuRET, Etudes Phycologiques,
p. 58, 1878. J. G. AGARDH, Till Algern. Systematik., VI, p. 67, 1882.
FarLtow, Marine Alge of New England, p. 111, 1881. Hus, Zoe,
Vol. V, p. 62, 1900.

Ulva laciniata AGARDH, Species Algarum, p. 404, 1822.

Porphyra vulgaris HARVEY, Phyc. Brit., Pl. CCXI. fig. 1, 185r.

Wildemania ? laciniata DE Ton, Sylloge Algarum, Vol. IV, p. 20, 1897.

Fronds membranous, 1o-80 cm. long, 5-30 cm. broad, linear when young,
becoming lanceolate or broadly expanded and much laciniate when older;
base obtuse to cordate; sessile on disc; gray-purple; monostromatic, vegeta-
tive part of frond 30-45 thick, cells square with rounded angles or higher
than broad, surface jelly 7-8 thick; dicecious, sometimes moncecious, sporo-
carps and antheridia forming a marginal zone, 8 (or 16) carpospores in each
sporocarp, 64 (or 128) antherozoids in each antheridium.

The author has, up to this time, been unable to find any
data in regard to the size this plant attains, but has received
the impression from the writings of others, as well as from
an examination of specimens, both American and European,
that 30 centimeters is the average length attained by the
fronds of this species, which suspicion was confirmed by
the study of a number of specimens of our western coast.
But recently there have come to our notice two specimens
from Orca, Alaska, one of which measured 60 centimeters,
while the other was 80 centimeters long, with a breadth of
30 centimeters. But such specimens do not indicate the
size the species normally attains; a length of 30 centimeters
and a breadth of 10-15 centimeters represent the average
measurement.

_—

:
e

BoT.—VOL. II.] HUS—PORPHYRA. 197

The shape of the fronds is far from being constant, and
appears, as in other species of Porphyra, to be strongly
influenced by environmental conditions. While the young
frond is liable to be more or less linear, even up to the time
when it attains a length of from 15 to 20 centimeters, most
specimens of that size show a considerable lateral develop-
ment. Older specimens are liable to be much laciniate.

Dhe form’) of the base, lke the shape of the frond,
depends on the surroundings; some specimens, probably
those which were exposed to the motion of the waves,
exhibiting an obtuse base when young, and a more or less
cordate base when older.

The color of the fronds is fairly constant, being as a rule
an even gray-purple, which increases in intensity with age.
Various shades may be met with.

The frond is proliferous and deeply folded and at times
beautifully crenulate.

Attachment is by a disc. The aureole around this disc,
so pronounced in P. perforata, is here but slightly marked.

The vegetative portion of the frond is uniform in thick-
ness, measuring from 30-45 microns. ‘The cells are square
or from two to three times as high as broad. The angles
are always rounded. There is but little jelly between the
cells, and the outer layer of jelly is not very thick, though
this varies more or less owing to different conditions of
exposure to air and heat, measuring about 7 or 8 microns.

The fronds are strictly monostromatic in the vegetative
part.

Porphyra lactniata is as a rule dicecious, though some
instances were found when the fronds were monececious.
The fruit occupies a marginal zone. When antheridia and
sporocarps are found on the same frond, they occur in
patches very much as in P. ferforata. The writer has
never met an instance where a few sporocarps were inter-
mixed with a large number of antheridia, such as are found
in the mznzata group; but a larger or smaller number of
vegetative cells and monospores may be found among the
sporocarps. Nor has he ever found vegetative cells among
the antheridia.

198 CALIFORNIA ACADEMY OF SCIENCES. [PRroc. 3D SER.

Sporocarps.—The reproductive bodies of P. Jaciniata
belong to the P. /eucosticta type. Each sporocarp contains
eight carpospores, which arise from the vegetative cell by a
cruciate division, followed by a parallel division, thus giving
rise to eight carpospores in two tiers of four each. This
appears to be the normal number, but frequently this mode
of division undergoes changes Either the vegetative division
proceeds one step further and only a parallel division takes
place, thus giving rise to but two spores, or in the latter case
an additional parallel division may occur, so that the sporo-
carp contains four carpospores in four tiers of one each.
Again it is possible that after the normal cruciate and par-
allel divisions have taken place, another more or less com-
plete division, usually parallel or oblique, forms sixteen or
less carpospores.

A ntheridia.—The antheridium-mother-cell first undergoes
a cruciate division, giving rise to four antheridia. Each
antheridium now undergoes its first reproductive division,
parallel to the surface of the frond. This division is fol-
lowed by a cruciate division and by a division parallel to the
surface of the frond, in all segments, thus giving rise to
sixteen bodies in four tiers of four each. These bodies
divide by a cruciate division, giving rise to 64 antherozoids.
This division is in many cases followed by a division parallel
to the surface of the frond, so that each antheridium now
contains 128 antherozoids arranged in eight tiers of 16 each.

The drawings of P. /actnzata of Bornet and Thuret
(1878) admirably illustrate these points, as well as those
cases where the vegetative division of the antheridium-
mother-cell is less apparent; so that we find 256 or 512
antherozoids in an antheridium, or where the vegetative
division of the antheridium-mother-cell goes one step farther
so that each antheridium contains but 32 antherozoids.

Economic Use.—According to the Rev. Albin Johnson,
the Indians of Yakutat, Alaska, collect, cook and eat this
plant.

Habitat.—On rocks or epiphytic on Fucus evanescens.
Throughout the litoral and lower litoral zones.

Bot.— VoL. II.] HUS—PORPHVRA., 199

Distribution.—P. laciniata appears to be limited on the
Pacific Coast to the shores of Alaska (61°-54° N. lat.).

Localities. —Orca, Alaska (W. A. Setchell, No. 5164!) ;
Yakutat, Alaska (Rev. Albin Johnson, No. 14!); Uyak
Bay, Kadiak Island, Alaska (W. A. Setchell, No. 5099!) ;
Sitka, Alaska (de A. Saunders, No. 136!); Annette Island,
Alaska (de A. Saunders, No. 26!); Amaknak Island,
Alaska (W. A. Setchell, No. 3269!, No. 3270!).

2. Porphyra laciniata f. umbilicalis A ¢.
Icones Algarum, Tab. XXVI, 1828.

Porphyra laciniata f. umbilicais C. A. AGARDH, Icones Algarum, Tab.
26, 1828. SETCHELL, Algz of the Pribilof Islands. Fur and Fur-seal
Islands of the North Pacific Ocean, 1899, p. 593. Hus, Zoe, Vol. V,
1900, p. 62.

Distribution.—Pribilof Islands, Bering Sea; U.S. S.
Albatross (according to Setchell, 1899) (57° N. lat.).

3. Porphyra leucosticta 7hur.
PLATE XX, Fics. 1a-30.
In le Jolis, Liste des Algues Marines de Cherbourg, 1864, p. too.

Porphyra leucosticta THURET in LE JoLis, Liste des Algues Marines de
Cherbourg, 1864, p. 100. DE JANCZEWSKI, Etudes Anat. sur les Por-
phyra, 1873, p. 241. CoLtiins, Bull. Torrey Bot. Club, Vol. XI, 1884,
p. 131. FarLow, Marine Algz of New England, 1881, p. 112. HOLDEN,
in CoLLins, HOLDEN & SETCHELL, Phyc. Bor.-Amer., Fascl. VIII, 1897,
No. 376. Hauck, in Hauck & RICHTER, Phyc. Universalis, Fascl. IX,
1891, No. 4o1. Hus, Zoe, Vol. V, 1900, p. 63.

Porphyra atropurpurea DE Tont, Syll. Alg., Vol. IV, 1897, p. 17.

Membranous, 7-70 cm. long, 2-25 cm. broad, oblong with slightly undulate
margin; base cordate, stipitate; color light pink; monostromatic, vegetative
part of frond 25-50 thick, cells once and a half to twice as high as broad,
surface jelly thin; fronds moncecious, antheridia forming small, elongated,
colorless patches among the dark-colored sporocarps; fruit marginal, gradu-
ally spreading over the whole frond, no vegetative cells intermixed with the
reproductive cells, sporocarpic part of frond 25-50 thick, eight carpospores
in each sporocarp; antheridial part of frond 30-50y thick, 64 antherozoids in
each antheridium.

Porphyra leucosticta Thuret is an annual, only to be
found in the spring months. It has never, to the writer’s

200 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

knowledge, been collected on the Pacific Coast north of
Monterey Bay. It appears to be exceedingly delicate, espe-
cially those parts of the frond which bear fruit. When
examining an herbarium specimen, the method of soaking
the desired portion in water, as pursued with P. perforata
and others, is entirely inadequate, water causing the almost
instantaneous dissolution of the jelly. It becomes there-
fore necessary to use a concentrated solution of corrosive
sublimate.

P. leucosticta is fairly constant in shape, irrespective of
size, varying from oval in the young plants to oblong in the
older ones. It is but seldom laciniate, and the margin is but
slightly undulate. The base is decidedly stipitate, some-
thing in which the western specimen differs from the east-
ern and European plants, which are at the most substipi-
tate. Another difference between eastern and European
specimens of P. /eucosticta on the one hand, and the west-
ern plants on the other, lies in the color, which in the latter
varies from cerise to dull brown, while the former appear
much lighter in color.

This plant is, in the vegetative portion of the frond, con-
stantly monostromatic. No indications of a distromatic
nature have ever been found.

The fronds are moneecious. At first the fruit is found
only at the tip and along the margins. In a ripe frond we
find a colorless margin, consisting of antheridia, together
with an empty network formed by the jelly-walls of those
sporocarps and antheridia which have discharged their con-
tents. Inside this margin, the sporocarps and antherida are
intermixed, the antheridia usually forming irregular, elon-
gated, colorless patches among the dark cerise sporocarps.

The thickness of the reproductive portion of the frond
does not differ materially from that of the vegetative por-
tion, measuring from 25-50. ‘This is an additional reason
for ascribing the increase in thickness in the reproductive
portions of the fronds of other species of Porphyra to the
swelling of the jelly, since in P. /eucostecta Thur. there is

Bot.—VOL. II.] HUS—PORPAVRA. 201

but a very small amount of jelly surrounding the cells, and
it could not swell enough to make an appreciable difference
in the thickness of the frond.

Vegetative cells are but rarely found intermixed with the
reproductive cells.

Sporocarps.—Each sporocarp contains eight carpospores.
They are formed from the vegetative cell by a cruciate
division, followed by a division parallel to the surface of the
frond. These divisions give rise to two tiers of four carpo-
spores each (Pl. XX) figs. 22-26; Pl. XXII, fig. 24).
Occasionally the vegetative division of the sporocarp
mother-cell goes one step farther; so that we find but two
carpospores in each sporocarp.

Antheridia.—The antheridium-mother-cell, by a cruciate
division perpendicular to the surface of the frond, gives
rise to four antheridia. Each antheridium now undergoes
a division parallel to the surface of the frond, then a cruciate
division perpendicular to the surface of the frond, followed
by another parallel division in all segments. The anther-
idium is now divided into sixteen parts, each of which, by a
cruciate division, gives rise to four antherozoids; so that
the whole antheridium now consists of sixty-four anthero-
zoids arranged in four tiers of sixteen each (Pl. XX, figs. 3a
and 36; Pl. XXII, fig. 27). The direction of the walls laid
down by the last cruciate division is often decidedly oblique,
so that, when the walls separating the individual antherozoids
dissolve, the groups of antherozoids are arranged in the form
of a hollow sphere. The wall laid down by the first, paral-
lel, reproductive division of the antheridium is thicker than
those laid down afterwards, and only dissolves when the
frond is fully ripe; so that for a long time two separate
spherical groups of antherozoids exist in each antheridium.

This spherical arrangement of the antherozoids seems to
be peculiar to the plants collected on the Californian shores,
since specimens of P. leucosticta collected on the coasts of
France and of Helgoland fail to show this.

Porphyra leucosticta was founded by Thuret (1864), and
has since been more fully described by Janczewski (1873).

202 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

According to de Toni (1897), it is identical with U/va atro-
purpurea Olivi.'

Flabetat.—On rocks and epiphytic on alge ( Gracilaria
confervoides); lower litoral and sublitoral zones; March
to May.

Distribution.—The author is aware of buta single region
of the Pacific Coast where this species occurs; i. e., Mon-
terey Bay, California (36° 45’ N. lat.).

Localities. — Pacific. Grove, California (M. A. Howe!,
Mrs. J. M. Weeks!, W. A. Setchell No. 5161!); Santa
Cruz, California (Mrs. J. M. Weeks!, de A. Saunders!).

4. Porphyra perforata 7. Ag.
PLATE XX, FIGs. 4a-10.
Till Alg. Syst. Afd. 3, VI, 1882, Ulvacez, p. 69.

Porphyra perforata J. AGARDH, Till Alg. Syst., Afd. 3, VI, 1882, Ulvacee,
p- 69. Hus, in Phyc. Bor.-Amer., Fascl. XIV, CoLitins, HOLDEN &
SETCHELL, No. 682, 1900. Hus, Zoe, Vol. V, 1900, p. 63.

Porphyra vulgaris ANDERSON, Zoe, Vol. II, 1891, p. 221. Howe, Erythea,
Vol. I, 1893, p. 67.

Wildemania perforata DE Tont, Syll. Alg., Vol. IV, 1897, p. 21.

Membranous, 7-70 cm. long, 3-20 cm. broad; linear-lanceolate, with undu-
late margin, often much expanded and laciniate; base cordate to umbilicate;
attached by a disc; gray to brown-purple, becoming blue-purple on drying;
monostromatic, vegetative part of frond 45-140 thick; cells once to two and
a half times as high as broad; surface jelly thick, often forming two-fifths of
the thickness of the frond; moncecious, sporocarps and antheridia in irregular
patches, the latter radiating towards the margin; vegetative cells often mixed
in with the sporocarps, never with the antheridia; each sporocarp containing
32 carpospores, each antheridium containing 128 antherozoids.

This species is known to the author from specimens col-
lected by Mrs. Snyder at San Diego, some of which were
communicated to the late Professor J. G. Agardh, and
referred by him to P. ferforata.

It is an annual, found throughout the year, the young
fronds making their appearance before the old ones have
altogether disappeared. Where the fronds are exposed to

1Saggi Accad. di Padova III, 1, (pres. die 18 Martii 1791), p .153, Tab. I-III (fide de
Toni).

Bor:—VOr. IT] HUS—PORPHYVRA. 203

the violence of the waves they disappear in the fall, some-
thing to which the dry north winds which frequently occur in
California during the latter part of the year contribute not
a little. The result of this is that in December and January
the fronds are found in sheltered places only.

P. perforata occurs throughout the litoral zone and in the
upper part of the sublitoral zone. There is some difference
in form between the plants of the upper part of the litoral
zone and those of the sublitoral zone. While the former,
which the writer considers typical, are lanceolate, with a
cordate base and a frond which is but slightly perforate, the
latter are far more irregular, being much lobed and laciniate,
with an umbilicate base, and are much perforate. They
also seem of finer texture and are as a rule much thinner.
This, however, is not always the case. The author has
collected specimens in the sublitoral zone which in the
sporocarpic portion of the frond measured r1rop,a thickness
which corresponds to the average measurement of the sporo-
carpic parts of the fronds of the litoral zone. The differ-
ence in thickness seems to lie chiefly in the jelly, the latter,
in the fronds of the litoral zone being as a rule more abun-
dant than in the fronds of the sublitoral zone.

The specimens of P. perforata found in the sublitoral
zone also possess a color different from that of the fronds
of the litoral zone. While the latter are a gray- to brown-
violet, the former are altogether devoid of a violet shade
and appear a gray- or yellow-brown. This difference in
color is very striking, and at first led to many conjectures.
But in the light of the fact that herbarium specimens after
having been preserved for six months or more begin to take
a violet hue, a color, the various shades of which are often
characteristic of the various species to which the genus
owes its name, and chiefly its inclusion among the Rhodo-
phycez, it seems but reasonable to suppose that the deeper
violet color of the specimens of Porphyra perforata found
in the litoral zone is due to the action of the air. What
the other causes for the difference may be has not yet been
solved. Perhaps the slug, which in all probability causes

204 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

the perforations, is more numerous in the sublitoral zone
than in the litoral zone,’ or perhaps the light has something
to do with it; but whatever the cause, both the anatomical
and morphological differences are too slight to allow us to
consider the sublitoral zone form a variety of the typical
Porphyra perforata J. Agardh.

The shape of the frond of Porphyra perforata is as a
rule linear-lanceolate, with an undulate margin and a cor-
date base. But frequently more or less expanded, broadly
laciniate fronds are met with, which possess an umbilicate
base. Between these two extreme forms which the frond
may assume, numerous transition stages are found.

The size of the frond is variable. Fruit has been found
on specimens but seven centimeters long. A well-devel-
oped specimen usually measures from 50-70 centimeters.
Both size and shape of the frond depend apparently largely
upon local conditions. For instance, those plants which
grow on the rocks in the upper part of the litoral zone
seem but seldom to attain a great development, and are
usually lanceolate. Those which grow in the elitoral zone
or in places where they are exposed to the tow of the tide
and are always, or for the greater part of the time, stretched
out by the water—such as those which grow on lagoons
and on long, low reefs,—are more linear-lanceolate, while,
as has been said before, those growing on rocks exposed
to the irregular wash of the waves are much laciniate and
rather broadly expanded.

The plants are attached by a disc, the structure of which
is of much interest. It agrees in all respects with that of
P. laciniata as described by Bornet (1878). P. perforata

1 Von Martens (1866), in a discussion on the use of seaweed as food by marine ani-
mals, describes some specimens of P. vulgaris Ag. (?) which were covered by a large
number of sea-slugs (Nasa corniculum Olivi), and which were much perforate. In their
immediate vicinity grew plants of Grateloupia filicina and of Chetomorpha linum, which
were not inhabited by the slug and had not been damaged. From this he concludes
that the perforate condition of the specimens of P. vulgaris was due to the action of
the slug, which used the plants for nourishment. The slug which infests the plants
Porphyra perforata on the Californian shores is a species of Lacuna. On this subject
Agardh (1882) says: ‘‘ * * * ; preterea frequenter foraminibus minutis rotundatis,
linem aut paucas diametro equantibus, precipai in disco perforatam vidi, et hoc
quidem aliquando jam in speciminibus minoribus et angustis.”’

BoT.— VOL. II.] HUS—PORPHVRA. 205

possesses a pronounced aureole around the point of attach-
ment, often one centimeter in diameter.

The writer has had an opportunity to examine a large
number of specimens of P. Zerforata collected at numerous
localities on the shores of the Pacific Ocean, but has up to
this time failed to encounter a single distromatic specimen.
A thorough examination of the fronds left no doubt as to
the purely monostromatic nature of the species, all consist-
ing in the purely vegetative part of but a single layer of
cells.

The thickness of the frond varies from 40 to 140m, the
difference in thickness being oftener due to a greater
amount of jelly than to a difference in the height of the
cells. The cells in the lowest part of the frond are usually
square or nearly so, but more towards the tip they are from
one and one-half to two and one-half times as high as
broad.

Porphyra perforata is monecious. Patches of sporo-
carps and antheridia occur here side by side, usually alter-
nating; but not a single instance has been met with, where
the antheridial and sporocarpic cells were intermixed, such
as we meet with in P. amplssima, or where one-half of the
cell formed’ antherozoids and the other half carpospores,
such as described by Janczewski (1873) for P. leucosticta.
The frond is often nearly entirely sporocarpic or antherid-
ial. Vegetative cells are sometimes encountered among
the sporocarps but never among the antheridia. There are
also cases where one or several of the divisions of the cell
in the formation of the sporocarp have not developed, so
that but one or more have formed spores, while the others
are to all appearances dead. The walls of such a cell are
thickened, the contents are hyaline and of a deep yellow
color, which sharply contrasts with the dark violet tinge of
the carpospores, and the unchanged, granular contents of
the vegetative cells or of the monospores. Some instances
have been noted where the sporocarps occurred in long
narrow patches, instead of in the broader patches usually
found.

(3) December 27, 1901.

206 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

The patches of sporocarps are easily distinguished from
the patches of antheridia. The former are of a uniform,
dark violet color, which gradually, towards the center of
the frond, fades into the gray- or brown-violet of the vege-
tative part of the frond. The patches of antheridia, on the
other hand, are nearly colorless when fully ripe. They are
usually very narrowly triangular and radiate from the center
of the frond. These patches contain the unripe antheridia
and are, towards the center of the frond, of a light violet
color, lighter than that of the surrounding sporocarps, but
deeper violet than that of the vegetative cells.

Sporocarps.—The first division of the sporocarp is cru-
ciate and perpendicular to the surface of the frond. This
is followed by a division parallel to the surface of the frond,
thus giving rise to eight segments in two tiers of four each,
every one of which undergoes a cruciate division, thus giving
rise to thirty-two spores in two tiers of sixteen each (Pl.
XX, figs. 52 and 54; Pl. XXII, fig. 25). Rarely there is
an additional parallel division in some or all the segments
formed by the last cruciate division.

Antheridia.—The antheridium-mother-cell undergoes a
cruciate division perpendicular to the surface of the frond,
giving rise to four antheridia. The first reproductive divi-
sion of the antheridium is parallel to the surface of the
frond. This is followed by a cruciate division, after which
another parallel division takes place; so that each anther-
idium now contains sixteen segments arranged in four tiers
of four each. Each of these segments now undergoes first
a cruciate division and then a parallel division; so that each
antheridium now contains 128 antherozoids, arranged in
eight tiers of sixteen antherozoids each (Pl. XX, fig. 6; PI.
XXII, fig. 28). In this species the last division is nearly
always regular and fully carried out.

Sometimes the vegetative division of the antheridium-
mother-cell does not take place; so that four times the
number of antherozoids are formed in an antheridium.

Economic Use.—Porphyra perforata is one of the edible
seaweeds, and is largely collected for food by Indians and
Chinese on the Pacific Coast.

Bor.—VOL. II.] HUS—PORPHYRA. 207

Habitat.—On rocks, wood and barnacles. Epiphytic on
Zostera, Phyllospadix, Nitophyllum, Fucus and Gigartina.
Throughout the litoral and in the upper part of the sublitoral
zones. Found throughout the year, but during the winter
months in sheltered places only.

Distribution.—At present known from the Pacific Coast
only, from Alaska to Southern California (58° 30’—32° 20’
N. lat.).

Localities. —Glacier Bay, Alaska (de A. Saunders, No.
100!); Baranoff Island, Alaska (de A. Saunders, No.
130!); Shumagin Island, Alaska (de A. Saunders, No.
394!); Whidby Island, San Juan County, Washington (N.
L. Gardner, No. 295!); Chehalis Bay, Washington (Ralph
Emerson, No. 1792!); Crescent City, Del Norte County,
California (H. Hus!); Trinidad, Humboldt County, Cali-
fornia(H. Hus!); Duxbury Reef, Marin County, California
CW. A: Setchell; No. 1055!; H. Hus, No. 81!); Farallon
Islands, California (H. Hus!); Lands’ End, San Francisco,
California (W. A. Setchell, No. 2034!, No. 2068!, H. Hus,
ING 26) .INo. 69! ,e¢ a/.);, Santa’ Cruz, Calitornia:(C, L.
Anderson!); Monterey, California (W. A. Setchell, No.
5159!, H. Hus!); San Simeon Bay, California(Dr. Palmer!) ;
Santa Barbara, California (Mrs. Cooper!); San Diego, Cali-
fornia (Miss Reed, No. 25!); Coronado, California (Mrs.
M.S. Snyder!).

5: Porphyra perforata f. segregata Setchell & Hus.
Zoe, Vol. V, 1900, p. 64.

Porphyra perforata f. segregata SETCHELL & Hus, Zoe, Vol. V, 1900,
p. 64. SNYDER, in Phyc. Bor.-Amer., Fascl. XIV, CoLitins, HOLDEN &
SETCHELL, No. 684, 1900.

Porphyra leucosticta TILDEN (not THuRET), Amer. Algze, Cent. III, No.
228, 1898.

This plant agrees in many respects with the species, but
there are some fundamental differences, which, though they
do not entitle it to a separate specific name, make it desir-
able to distinguish it as a distinct variety. As a rule the
plant is much smaller and has a rather more umbilicate base

208 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

than P. perforata. The frond is much thinner, never ex-
ceeding sixty microns. The vegetative cells are smaller
than those of the species, and the jelly walls separating the
different cells are thicker, giving the vegetative portion of
the frond when seen in surface view a most characteristic
appearance.

The difference between the reproductive cells of P. per-
forata and this variety can be best seen in cross-section.
It is especially noticeable in case of the antheridia. The
wall laid down by the first reproductive division of the
antheridium is very thick and widely separates the anther-
ozoids into an upper and a lower group.

Distributton.—F rom Washington southward to Lower
California (47° 30’—27° 8%’ N. lat.).

Localities.—Shillshole Bay, Seattle, Washington (Miss
J. E. Tilden!); San Pedro, California (de A. Saunders,
No. 1034!); San Diego, California (Mrs. M. S. Snyder!);
San Roque, Mexico (G. Eisen!).

6. Porphyra perforata f. lanceolata Setchell & Hus.
Zoe, Vol. V, 1900, p. 65.

Porphyra perforata f. lanceolata SETCHELL & Hus, Zoe, Vol. V, 1900, p. 65;
in Phyc. Bor.-Amer., Fascl. XIV, CoLtins, HOLDEN & SETCHELL, No.
683, Igoo.

Porphyra laciniata TILDEN, Amer. Algz, Cent. III, No. 229, 1898.

Membranous; 10-325 cm. long, I-to cm. broad; linear with undulate mar-
gin; base cuneate to cordate, attached by a disc; steel-gray to gray- or yellow-
brown, becoming purple on drying; monostromatic, vegetative part of frond
75-150 thick; cells one and one-half to four times as high as broad; jelly
very thick, forming two-fifths to one-half the thickness of the frond; dice-
cious; sporocarps containing 32 carpospores, each antheridium containing
128 antherozoids.

This variety of P. perforata agrees in many respects
with the species. The chief difference lies in the fact that
it is dicecious. Besides this there are numerous minor
differences.

P. perforata {. lanceolata can be distinguished at first
sight from the species by its form. This is either distinctly

BorT.—VOL. II.] HUS—PORPHVRA. 209

linear with undulate margin, or deeply laciniate, producing
a two-forked frond. Sometimes a specimen, and thisis true
for those of even 50 centimeters long, measures but one
centimeter across; but the larger specimens may be as wide
as 10 centimeters. The average width is 5 centimeters,
with a length of 50 centimeters. Mr. R. E. Gibbs, how-
ever, collected a specimen of this variety at the Presidio
which measured 325 centimeters in length, and with a width
of Io centimeters in the broadest part. The plant was
growing on a rock buried in the sand of a gently sloping
beach, where it was carried back and forth by the waves,
which perhaps brought about its extreme development.
Something similar is met with in the linear-lanceolate fronds
of P. perforata when growing in lagoons.

The base is cuneate to cordate, and is but rarely umbili-
cate. The plant is attached bya disc, the structure of which
is identical with that of the disc of P. perforata.

The color of the fronds varies considerably. Asa rule
the fronds are steel-gray to gray- or yellow-brown; some-
times they are in parta bright green. In mature fronds the
edges, if the frond is antheridial, are yellow and appear
much swollen. The latter is caused by the swelling of the
jelly preparatory to breaking down and setting the anther-
ozoids free. The sporocarpic frond is usually of a darker
color, the edges having a reddish bfown appearance.

The difference between antheridial and sporocarpic
frondsis brought out still better on drying, when the anther-
idial fronds become distinctly yellow at the edges, and the
sporocarpic fronds, in the region of the sporocarps, red-
violet. The latter fronds are also less shiny than the
antheridial fronds.

Occasionally forked fronds are met with; and in such
cases it is not unusual to find one fork bearing antheridia,
while the other is strictly sporocarpic. These subdicecious
fronds form a connecting link between the form and the
species proper.

The number of divisions in the sporocarps and antheridia
seems to be the same as in P. ferforata.

210 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

Flabitat.—P. perforata f£. lanceolata is an annual, occur-
ring throughout the year, but in winter found only in shel-
tered places. It usually grows on rocks in the highest part
of the litoral zone. In some cases it has been found on but
a single rock for a distance of overa mile. This was in
December, but the rocks in the same locality (Fort Point,
Presidio, San Francisco) were found to be thickly covered
with it in June of the next year.

Distribution.—Up to now this plant has, to the author’s
knowledge, been found in but two localities, both in Cali-
fornia (37° 47-36° 45’ N. lat.).

Localities.—Lands’ End, San Francisco, California (W.
A; Setchell!, H. Hus!, R. E. Gibbs, Miss J. EB. Gilden!)
Carmel Bay, Monterey County, California (W. A. Setcheli!,
R. E. Gibbs!).

7. Porphyra nereocystis Anderson.
PLATE XX, FIGs. 11a-12.
Zoe, Vol. III, 1892, p. 148.

Porphyra nereocystis ANDERSON, Zoe, Vol. II, 1891, p. 221, (name only);
Vol. III, 1892, p. 149 (descr.). Howe, Erythea, Vol. I, 1893, p. 67.
SETCHELL, in HOLDEN, COLLINS & SETCHELL, Phyc. Bor.-Amer., Fascl.
XII, No. 583, 1899. Hus, Zoe, Vol. V, rgoo, p. 65.

Pyropia californica J. G. AGARDH, Anal. Algol., Cont. V, 1899, p. 153-

Fronds 3-270 cm. long, 2-40 cm. broad, linear to oblong, with laciniate
margin and obtuse or cucullate base; attached by a disc; fronds red to purple,
monostromatic, vegetative part 25-6oy thick, little jelly between the cells;
fronds moncecious, antheridia forming light-colored, sharply defined spots
and streaks among the dark-colored sporocarps, 32 carpospores in each
sporocarp, 128 antherozoids in each antheridium.

This species, first mentioned by Dr. Anderson of Santa
Cruz, California, is one of the largest of known species of
Porphyra. It attains a great length, specimens go centi-
meters long being by no means rare. From notes appended
to a fragment of one specimen it was gathered that the
whole specimen measured 270 centimeters. The shape is
fairly constant. The younger specimens are lanceolate
and have an even margin. Older specimens are oval and

Botr.—VOL. II.] HUS—PORPHVRA. PAPAL

have a cucullate base. The aureole around the disc is very
pronounced in this species. The frond is not plicate, but
on drying several folds appear in some of the older speci-
mens, owing to the concave nature of the frond caused by
the cucullate base. The margin in the older fronds is
much laciniate.

The fronds are monostromatic in the vegetative part. No
deviation from this has been observed. The cells are
square with rounded angles, or may be twice as high as
broad or twice as broad as high.

Sporocarps.— The mature frond becomes gradually
changed into reproductive cells of which the sporocarps
form the majority. They can be readily recognized by
their deep red color, and are sharply outlined against the
lighter colored antheridia. The sporocarp first divides
cruciately, giving rise to four cells. Each of these now
undergoes a parallel division, followed by a cruciate divi-
sion; so that in each sporocarp we finally have thirty-two
carpospores arranged in two tiers of sixteen each (Pl. XX,
Meth; el RON, fig 25).

Antheridia.—They are found in larger or smaller, irreg-
ular, light-colored patches among the sporocarps. The
antheridium-mother-cell, by a cruciate division, gives rise
to four antheridia, each of which by alternating parallel
and cruciate divisions, as in P. perforata, give rise to 128
antherozoids, arranged in eight tiers of sixteen each.

Economic Use.—This plant is largely collected by China-
men on the Californian coast, along with /. perforata.
From what could be learned, it seemed that it is much more
esteemed than the latter.

Flabitat.—This species is found attached to the stipes of
Nereocystis liitkeana in from three to five fathoms of water,
and is also found on rocks. After a storm it is frequently
found floating.

Distribution.—Porphyra nereocystis has a very wide
range, having been collected at St. Paul, Kadiak Island,
Alaska, and as far south as San Pedro, California (57° 30’—
ga? 40) Neilat:);

212 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

Localities.—St. Paul, Kadiak Island, Alaska (W. A.
Setchell and A. A. Lawson, No. 5149!); Coupeville,
Washington (N. L. Gardner!); Bolinas Bay, Marin
County, California (W. A. Setchell, No. 1275!); Monte-
rey Bay, California (W. A. Setchell, No. 3065a!; C. L.
Anderson!); San Pedro, California (de A. Saunders!).

8. Porphyra naiadum Anderson.
PLATE XXI, Fics. 19-22.
Zoe, Vol. III, 1892, p. 148.

Porphyra naiadum ANDERSON, Zoe, Vol. III, 1892, p. 148. Howe, Erythea,
Vol. I, 1893, p. 67. McCratcuir, Proc. So. Cal. Acad., Vol. I, 1897,
p. 356. TILDEN, Amer. Algz, Cent. III, No. 231, 1898. SETCHELL in
Phyc. Bor.-Amer., Fascl. XIII, Cottins, HOLDEN & SETCHELL, No.
632, 1899. Hus, Zoe, Vol. V, rgoo, p. 66.

Porphyra coccinea AGARDH, J. G., Till. Alg. Syst., Afd. 3, VI, 1882, p. 58;
in part (as to the Californian specimens ?).

Porphyra sp. RUPRECHT, Neue oder unbek. Pflan. d. N. Th. d. St. Oceans,
1852, p. 65.

Fronds 2-10 cm. long, obovate when young, oblanceolate when older;
base cushion-shaped; fronds wine-red to blue-purple; monostromatic, vege-
tative part 25-30 thick, cells square or slightly higher than broad, 15-20p
high; surface jelly measuring about 5y, little jelly between the cells; fronds
dicecious ?; sporocarps with eight carpospores.

The first mention of this species of Porfphyra was prob-
ably made by Ruprecht (1852), who speaks of a parasitic
Porphyra occurring on Phyllospadix scoulert. It was first
recognized asa distinct species by Dr. Anderson in 1892;
but for reasons given below it is evident that it was known
before that time to J. G. Agardh. Formerly it had been
distributed along with P. nereocystis and P. perforata as
P. vulgaris, but, as Dr. Anderson says, ‘‘ without much
more reason than our early botanists had for placing all sea-
weed in the genus Fucus.’’ It is found growing on Phyllo-
spadix at extreme low-water mark, and in sheltered and
exposed places alike. The plants occurring on a single
blade of eel-grass are usually so numerous that the color of
the adult fronds literally hides the green color of the eel-
grass. It is interesting to note that most of the fronds

BoT.—VOL. II.] HUS—PORPHYVRA. 213

occurring on Zostera usually attach themselves to the edges
of the blade, forming a broad fringe.

In the spring of 1898, Dr. Setchell called the attention of
the author to some plants of Phyllospadix, the leaves of
which were densely covered by numerous small, reddish-
brown, cushion-shaped growths, which he suspected to
stand in some relation to P. natadum. On examination,
these cushions proved to consist of a number of angular cells
(Pl. XXI, fig. 19), the outer layers of which contained chro-
matophores, which were deeply lobed, somewhat after a stel-
late fashion, resembling those described by Schmitz (1882)
for Helminthocladia purpurea. Each cell of the layer
adjacent to the substratum possessed a short rhizoid, which
apparently attached itself to the cuticle of the host-plant.
In the beginning of February, these cushions begin to show
short hair-like projections. These projections are composed
of cells placed in a single narrow layer. The youngest of
them consist of but a few cells placed ina single row.
Older specimens show that these cells divide in two direc-
tions, giving rise to a monostromatic frond which proves to
be P. naiadum (Pl. XX], fig. 22).

The cells of the lower part of the frond, unlike those at
the base of fronds of other species of Porphyra, do not
produce hypha-like projections. Even in the oldest plants
no sign of this was to be found. This prothalloid base is
different from the bases of all known species of Porphyra,
as described by Berthold, Thuret, and others, with the
exception, perhaps, of one described by J. G. Agardh (1882)
under the name P. coccinea.

The size of the frond varies somewhat. The young
fronds are obovate, measuring about one centimeter across;
but the frond gradually broadens and lengthens, finally
assuming an oblanceolate shape, being from four to ten
centimeters long and two or three centimeters broad. There
is a considerable difference in size and color between speci-
mens growing in sheltered places and those growing in the
open. In fact, we may distinguish between two forms—a,
minor and 4, major. Under the first, we understand the

214 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

form of P. naiadum found on Phyllospadix, growing in
exposed places, with a smaller, rather orbicular frond and a
laciniate margin; while by P. xazadum f. major, we mean
the plant which grows on Zostera,in lagoons and other
sheltered places, with a larger, broader frond, and which
usually possesses an even margin.

Only one form of fruit was found, viz.—sporocarps. As
in the case of the reproductive bodies of the other species
of Porphyra, they are here first formed in the region of
the tip, gradually spreading over the whole frond.

Sporocarps.—In the formation of the sporocarp, the first
division is cruciate, followed by a division parallel to the
surface of the frond, thus giving rise to eight spores in two
tiers of four each (Pl. XXI, figs. 21a and 216; Pl. XE
fig. 24). Atthe time of the ripening of the sporocarp the
jelly around the spores swells, thus making the frond a few
microns thicker, but finally dissolves as does the outer jelly-
wall, setting the spores free.

Dr. Anderson suggests that small molluscs, in rasping for
food on eel-grass, cause abrasions, ‘‘in which the spores of
this Porphyra find a place to adhere;”’ but careful section-
ing has failed to show any injury to the epidermal layer of
the eel-grass at the places of attachment of very young
fronds.

Habitat.—The plant has been collected throughout the
year. It grows exclusively on Phyllospadix and Zostera.
In sheltered places we find it all through the winter, but
where the eel-grass is exposed to the direct influence of the
tides, the Porpfhyra disappears about August, to reappear
in the prothalloid form about the latter part of January.

Distribution.—This plant appears to be limited to the
Pacific Coast, occurring from Washington to Southern
California (48° 10—32° 20 N. lat.).

Localities.—Puget Sound (de A. Saunders!); Coupeville,
Washington (N. L. Gardner!); Ballard Beach, Seattle,
Washington (Miss J. E. Tilden!); Crescent City, Del
Norte County, California (H. Hus!); Fort Ross, Sonoma
County, California (W. A. Setchell, No. 1789!); Duxbury

Bor.—VOL. II.] HUS—PORPHVRA. 215

Reef, Marin County, California (W. A. Setchell, No.
1037!, No. 1276!, H. Hus, No. 80!); Farallon Islands,
California (T. W. Blankinship); Land’s End, San Fran-
cisco,’ California (W: A. Setchell, No. 1118!, H: Hus!);
Monterey, California (W. A. Setchell, No. 5158!, C. P.
Nott!,) Mrs. Bush, RE! Gibbs!; H. Hus!); Santa Cruz,
California (C. L. Anderson!); San Pedro, California (A.
J. McClatchie!); San Diego, California (Miss Reed, No.

107!).

g. Porphyra amplissima (A7e//man) Setchell & Hus.
PLATE XX, Fics. 13a-130.
Zoe, Vol. V, Ig00, p. 67.

Porphyra amplissima (KJELLMAN) SETCHELL & Hus. Hus, Zoe, Vol. V,
1g00, p. 67. GARDNER, in Phyc. Bor.-Amer., COLLINS, HOLDEN &
SETCHELL, Fascl. B, No. XLIX, Igor.

Diploderma amplissimum KjeELitmM., The Algze of the Arctic Sea, p. 188,
Pl. 17, figs. 1-3; Pl. 18, figs. 1-8, 1883.

Diploderma amplissimum f. typica Fostiz, Cont. to Knowledge of the
Marine Algze of Norway, I, 1890, p. 56.

Wildemania amplissima Fos.ir, Cont. to Knowledge of the Marine Algze of
Norway, JI, 1891, p. 14. DE Tont, Sylloge Algarum, Vol. IV, 1897, p. 24.

Fronds membranous, 20-60 cm. long, 1o-15 cm. broad, broadly elliptical
to ovate-lanceolate, with much undulate margins deeply folded, the folds usu-
ally reaching to the median line of the frond; color deep red-purple; base
slightly cuneate, sometimes cordate, sessile with small disc; fronds distro-
matic, 50-S0f thick; cells in cross-section square or slightly higher than
broad, with rounded angles; surface jelly 5-10 thick; fronds moncecious,
antheridia and sporocarps intermixed in a marginal zone, sometimes together
with vegetative cells; each sporocarp consisting of 4 or 8 carpospores, each
antheridium consisting of 16 antherozoids.

This alga is the Dizploderma amplissimum of Kjellman
(1883), agreeing in every respect with both drawings and
description. It varies considerably in size. Kyjellman sug-
gests that the greater size is due to the floating condition.
It is, however, doubtful if plants of this species continue to
grow after being torn away. The author, not having col-
lected this species, is unable to express an opinion, but
judging from the fact that plants of other species of

216 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

Porphyra when found floating are in a very poor condition
it seems very unlikely. One would rather be inclined to
ascribe the greater development of specimens of P. amplis-
stma to little exposure, since, judging from notes appended
to specimens examined, it appears to be due to the plants
growing in undisturbed water, while specimens collected in
exposed places always were small.

The shape of the fronds is to a certain extent variable.
The extreme forms are broadly elliptical and ovate-lanceo-
late. Between these two a large number of intermediate
forms may be noted. The diversity in form is perhaps due
to a great extent to mechanical influences. An important
factor is also the age of the specimen. The older, longer
plants seem to possess a greater lateral development, out of
proportion to the increase in length.

The color seems likewise to depend on the age of the
plant. The younger specimens vary from lake to deep red-
purple, but the color seems to fade out in the older speci-
mens, which are often a faint brownish red.

The fronds are densely folded, the folds extending to the
median line of the frond. ‘The margin is smooth or lacini-
ate, a matter which also probably depends on greater or
less exposure.

The base is asa rule more or less cuneate. In some
specimens the base is strongly cordate. Many intermediate
forms may be met with. Our plants from Alaska are not
stipitate and in this regard differ from Kjellman’s descrip-
tion. They are sessile upon asmall disc, much smaller than
the one we find in Porphyra perforata.

The thickness of the frond varies from fifty to eighty
microns. The majority of the specimens, however, meas-
ured about sixty microns, both in the vegetative and in the
reproductive parts. The shape of the cells in the middle
of the frond is, in cross-section, square or slightly higher
than broad.

The plant is as a rule moneecious, sporocarps and anther-
idia occurring side by side, as shown in figs. 13a and 130.
Frequently one cell develops into antheridia, while the

Bot.—VOL. II.] HUS—PORPHYVRA. 217

corresponding cell on the other side of the frond becomes
asporocarp. Sometimes vegetative cells are found among
the reproductive cells to a greater or less extent. Some-
times specimens are met with which are dicecious or nearly
so; but the majority of the fronds are monecious.

Sporocarps.—The number of spores in each sporocarp is
four or eight. The sporocarp divides into four parts by a
cruciate division. The division may take place in one
direction only (perpendicular to the surface of the frond),
or one line only may be perpendicular and the other paral-
lel to the surface. These divisions would give rise to four
spores in one tier or to four spores in two tiers of two
spores each. A cruciate division perpendicular to the sur-
face seems to occur most frequently. In this case a third
division, parallel to the surface of the frond, sometimes
takes place, thus giving rise to eight spores in two tiers of
font each (Pl. XX, digs. 13¢ and, 136; Pl. 2X11, fig. 24).
In making these observations care was always taken to
select an absolutely ripe portion of the frond.

Antheridia.—The division of the antheridium-mother-cell
goes one step further than that of the sporocarp-mother-
cell, a cruciate vegetative division perpendicular to the sur-
face of the frond taking place before the formation of the
antheridium proper is begun. A sporocarp therefore corre-
sponds to four antheridia. Each of these antheridia sur-
rounds itself with a wall of jelly. By subsequent divisions,
the first parallel to the surface of the frond, the second a
perpendicular cruciate division, and the third again parallel,
there arise sixteen antherozoids arranged in four tiers of
fone each (Pl. XX, figs. 13¢ and 136; Pl. XXII, fig. 26).
This number may, however, vary, and from two causes.
The division of the antheridium-mother-cell is sometimes
incomplete, so that each antheridium becomes four times as
large as the normal ones. Usually in such cases an addi-
tional reproductive division takes place, so that each anther-
idium possesses sixty-four antherozoids. Frequently also
the divisions in the antheridium are incomplete, so that the
number of antherozoids is less than sixteen.

218 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

flabttat.—On rocks in the sublitoral zone. Also found
floating. June, July, August.

Distribution.—From Alaska southward to Washington
(60°—48° 10’ N. lat.).

Localittes.—Orca, Alaska (W. A. Setchell and A. A.
Lawson, No. 5165!, de A. Saunders, No. 259a!); Unga,
Alaska (W. A. Setchell and A. A. Lawson, No. 5047!);
Amaknak Island, Alaska (W. A. Setchell, No. 3268!);
Coupeville, Washington (N. L. Gardner, No. 47! No. 280!
No. 1990!).

10. Porphyra miniata (Lyngb.) Ag.

Porphyra miniata (LYNGB.) AGARDH, C., Syst. Algarum, 1824, p. I9gI.
AGARDH, J., Till Alg. Syst., VI, 1882, p. 59.

Ulva miniata LYNGBYE, Hydrophyt. Dan., 1819, p. 29, Tab. 6, D.

Diploderma miniatum KyELLM., Algze of the Arctic Sea, 1883, p. 189.

Wildemania mtniata Fos.ie, Cont. to Knowledge of Marine Algz of Nor-
way, II, 1891, p. 14 (in part); DE Ton, Sylloge Algarum, Vol. IV, 1897,
p: 23.

11. Porphyra miniata f. cuneiformis Setchell & Hus.

Prats TD Bic. 74.

Porphyra miniata f. cunetformis SETCHELL & Hus. Hus, Zoe, Vol. V,
1900, p. 68.

Fronds membranous, 15-50 cm. long, 4-15 cm. broad, lanceolate with
undulate, often crenulate, margins; color red-purple; base strongly cuneate,
attached by a disc; fronds distromatic, often monostromatic near edge in
vegetative part, 30-75 thick; cells square to twice as long as broad; surface
jelly to-12.54 thick; fronds moncecious, antheridia and sporocarps inter-
mixed in a marginal zone, gradually spreading over the whole frond, each
sporocarp containing 4 carpospores, each antheridium consisting of 8
antherozoids.

Nearly two hundred specimens of this species, collected
at different times on the Pacific Coast, agreed essentially in
all respects among themselves as well as with specimens of
Porphyra miniata from the Atlantic coast; among others
with those distributed in the Phyc. Bor.-Amer. by Mr.
Isaac Holden (No. 377). Agardh’s notes on U/va purpurea
(P. miniata) are too meager to give a definite clue in the
determination of the species, especially since the number

Bor.—VOL. II.] HUS—PORPHYVRA. 219

of antherozoids in each antheridium, which the author pro-
poses as a criterion for the species of Porphyra, is not
mentioned. But, since the specimens do not agree with the
description of Diploderma amplissimum by Kjellman, and
since it is evident that they cannot belong to P. tenuzss*¢ma
(Stromf.) S. & H. or P. abyssicola Kjellm., the writer has
deemed it advisable to assign the name P. mznzata to these
specimens, until such time as others who have access to the
type-specimens may determine its rightful position.

This species differs from the preceding one in color, in
being less deeply folded, and in possessing a crenulate mar-
gin which gives the fronds a very characteristic appearance.

It differs from Kjellman’s description of P. mzndata in
being monececious, and in possessing a strongly cuneate base.
These two characters were deemed sufficient to differenti-
ate the plant under a form name.

Sporocarps.—The sporocarps contain four carpospores,
formed by a cruciate division of the mother-cell (Pl. XX,
HentAs Pl, XOX, fig. 23).

Antheridia.—The antherozoids appear here in groups of
eight arranged in two tiers of four each, thus showing in
cross-section four antherozoids (Pl. XX, fig. 14; Pl. XXII,
fig. 25). At first it was thought that this appearance was
due to an unripe condition of the part of the frond exam-
ined, but a large number of sections from carefully selected
portions of the frond showed the same character.

Sometimes sixteen antherozoids were found in one anther-
idium; but in that case they were arranged in two tiers of
eight each. This showed that the double number of anther-
idia was caused by the fact that the last vegetative division
in the antheridium-mother-cell did not take place.

ffabitat.—F ound floating.

Distribution.—From Alaska southward to middle Cali-
fornia (60°—36° 45’ N. lat.).

Localities. —Lowe Inlet, Gulf of Alaska (de A. Saun-
ders, No. 20!); Coupeville, Washington (N. L. Gardner,
No. 199@2!); Monterey Bay, California (Mrs. J. M.
Weeks!).

220 CAEIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

12. Porphyra tenuissima (Strémf.) Setchell & Hus.
Zoe, Vol. V, 1900, p. 68.

Porphyra tenuissima (STROMF.) SETCHELL & Hus. Hus, Zoe, Vol. V, 1900,
p- 68.

Diploderma tenutssimum STROMF., Bot. Centralbl., Bd. XX VI, 1886, p. 173;
Om Algvegetationen vid Islands Kiister, 1886, p. 33.

Diploderma amplissimum f. tenuissima Fos.ie, Contribution to Knowl-
edge of the Marine Algz of Norway, I, 1890, p. 56.

Porphyra miniata f. tenuissima ROSENVINGE, Grénlands Havalger, 1893,
p. 827; Les Algues Marines du Greenland, 1894, p. 83.

Wildemania tenuisstma DE ToNt!, Sylloge Algarum, Vol. IV, 1897, p. 23.

Fronds membranous, 25 cm. long, 14 cm. broad, round-ovate, not lobed,
with undulate margin, folded; base cordate, sessile, with small disc; color
delicate pink; fronds distromatic, 254 thick, except at the base, where they
measure 65-754; cells near base square to twice as broad as high, ro-15u
broad, cells near edge of frond three to four times as broad as high, 20n
broad, with rounded angles; surface jelly 2.5y, little jelly between the cells;
fronds moncecious, antheridia and sporocarps intermixed in a marginal zone,
4 carpospores to each sporocarp, 8 antherozoids to each antheridium.

The description is drawn from several specimens collected
by Mr. de Alton Saunders when with the Harriman Alaskan
expedition during the summer of 1899. The study of a
large number of specimens of other species of Porphyra has
led to the conviction that, to describe a species from but a
few specimens is, to say the least, a very unreliable method
of procedure; and were it not for the fact that the author
had access to a numberof European specimens of P. ¢enuzs-
stma, as well as to Strémfelt’s ample description (1886),
with which the specimen in question agreed in nearly every
respect, such a course would not have been undertaken. In
matters such as length and breadth of the frond, the evi-
dence submitted is of course not conclusive. It must be
said that among our specimens are the largest which have
as yet come to our notice. The European specimens are
less than half the size. In the collection of Porphyra
kindly placed at our disposal by Mr. Collins, a specimen
was found, collected at Nahant, Massachusetts, which was
originally determined as P. J/eucosticta' and afterwards
designated P. mznzata,? but which is evidently P. tenuzssima.

1 Bull. Torrey Bot. Club, Vol. IX, 1882, p. 7o.
2Bull. Torrey Bot. Club, Vol. XI, 1884, p. 131.

Bot.—VOL. II.] HUS—PORPHVRA. PIPER

It is about fourteen centimeters long and ten centimeters
broad. It is the writer’s opinion that the specimen in ques-
tion belongs to the set distributed in the Phycotheca Gener-
alis, Fascl. I, No. 8, and referred to by Foslie (1890) when
speaking of a specimen of P. mznzata communicated to him
by Collins, and collected at Nahant, Massachusetts.

The shape of the fronds of P. tenuzss¢ma seems eminently
constant. Most specimens are roundish ovate, and are not
lobed in the slightest. The fronds are folded, more so than
in P. miniata but less than in P. amplissima. ‘The color is
a delicate pink. This character together with the thinness
of the fronds aftords the means of readily recognizing P.
tenuissima from other species of Porphyra having the
miniata-type of antheridia.

The base seems to be cordate, and is attached by a small
disc.

The fronds are distromatic throughout. There is some
variation in the thickness of the frond and in the shape and
size of the cells. While toward the edges the frond is but
little more than twenty to thirty microns thick, and vegeta-
tive cells, where they occur, are from three to four times as
broad as high, the cells being fifteen to twenty microns high
and from four to six microns broad, in the lower part of the
frond, near the base, the thickness varies from sixty-five to
seventy-five microns, and the cells are sometimes square,
sometimes twice as broad as high, being about twenty
microns broad and from ten to twenty microns high. In
this regard our specimens from the west coast agree fully
with the description of Strémfelt.

But it is different with the moneecious character. Strém-
felt describes this species as dicecious, only having found
sporocarps. But what we know of the variability of the
different species of Porphyra in this respect does not warrant
us in referring these specimens other than to P. tenuzssima,
awaiting a more extended comparison between our forms
and those of northern Europe.

(4) January 2, 1902.

222 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

The antheridia and sporocarps are intermixed in a mar-
ginal zone. Occasionally one or more vegetative cells are
found; but this is rarely the case. The antheridia are
present in predominating numbers. The reproductive part
of the frond has very much the appearance of that of P.
miniata in regards arrangement of sporocarps and anther-
idia. Often a cell on one side of the frond will form
antheridia, while the corresponding cell on the other side
of the frond will form carpospores.

Sporocarps.—The first division of the sporocarp is per-
pendicular to the surface of the frond. This is followed by
a division at right angles to it and perpendicular to the sur-
face of the frond, or nearly so, thus giving rise to four car-
pospores in a single tier, showing in cross-section two
carpospores (Pl. XXII, fig. 23). The divisions are more
or less regular. ‘They may take place simultaneously, thus
forming a cruciate division, or they may be consecutive,
and the second division in one half of the carpospore is in-
dependent of that in the other half.

Antheridia.—The vegetative division in the antheridium-
mother-cell goes one step farther than in the sporocarp-
mother-cell, a cruciate division perpendicular to the surface
taking place. After this a division parallel to the surface
of the frond occurs in each antheridium, followed by a very
regular cruciate division perpendicular to the surface of the
frond. This last division, however, may precede the paral-
lel division. These divisions finally give rise to eight an-
therozoids in each antheridium, arranged in two tiers of
four each, so that the original antheridium-mother-cell now
contains four groups of eight antherozoids each, each group
corresponding to a carpospore (Pl. XXII, fig. 25).

From the above it will be seen that P. tenuzsstma possess-
es the mznzata type of antheridia. It is for this reason that
the writer doubts the wisdom of Foslie’s action in making
P. tenuissima a form of P. amplissima.

Hlabitat.—Epiphytic on alge; also on rocks?.

Distribution.—As yet only known on the west coast of
North America from Alaska (59° 40-55° N. lat.).

Bor.—VOL. II.] HUS—PORPHVRA. 223

Localities.—Yakutat Bay, Alaska (de A. Saunders, No.
214!); Baranoff Island, Alaska (de A. Saunders, No. 136!,
No. 137!, No. 148!); Shumagin Island, Alaska (de A.
Saunders, No. 384!).

13. Porphyra abyssicola A7je//m.
Algz of the Arctic Sea, 1883, p. 191.

Porphyra abyssicola KjELLMAN, Algz of the Arctic Sea, 1883, p. 191; Pl. 17,
fig. 4; Pl. 18, figs. ro-11. Fostre, Cont. to Knowledge of the Marine
Algee of Norway, I, 1890, p. 60. Hus, Zoe, Vol. V, 1900, p. 68. DE
Tonl, Sylloge Algarum, Vol. IV, 1897, p. 14.

Fronds membranous, 2-10 cm. long, 1-4 cm. broad, lanceolate-obovate,
with undulate margins; color livid cerise; base cuneate, sessile with small
disc; distromatic, often monostromatic, 25 thick; cells of the middle of the
frond square or nearly twice as long as broad, with rounded angles; fronds
moncecious, often dicecious; when moncecious antheridia and sporocarps
intermixed in a marginal zone, 2-4 carpospores to each sporocarp, 8 anther-
ozoids to each antheridium.

This species is known to the author by some twelve spec-
imens found among a number of specimens of P. xazadum
growing on Zostera. ‘They agree in every respect with the
description of P. adbysstcola of Kjellman. Only a few of
the specimens were fertile. On examination it was found
that some were monececious and others dicecious.

The specimens examined grew on Zostera, and were
easily distinguished by their more elongated frond and by
the absence of the characteristic cushion-shaped base of
P. naiadum. The plants of P. abyssicola were also deeply
though sparsely plicate.

In this species the difficulty of assigning it to the mono-
stromatic or distromatic group makes itself more felt than
in any other case. The specimens which were available
for examination were for the greater part distromatic, and
I hesitated considerably before assigning them to P. adyssz-
cola. But it must be remembered that plants of this group
vary considerably in this regard, and since all other char-
acters were those of Kjellman’s P. aéyssccola, they were
finally assigned to this species.

224 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

The possibility that a species might possess plants some
of which were monostromatic while others were distromatic,
or that one plant might be monostromatic in one part and
distromatic in the other, and this distromatic nature not due
to reproductive division, was evidently not considered by
Kjellman when he divided the genus Porfhyra into Por-
phyra proper and Dzploderma, Not only do we meet
monostromatic and distromatic forms in P. abysszcola, but
P. miniata and P. tenuzssima also exhibit this character in
a greater or less degree. It therefore seems to the writer
unwarranted to separate such closely allied species by a
criterion which does not hold true in all cases; and he fully
concurs in the action of Rosenvinge, who, after finding
many intermediary forms, reunited the two subgenera.
But the writer cannot agree with Rosenvinge in uniting P.
miniata, P. amplissima, P. tenutsstma, and P. abyssicola
under Porphyra minzata, since there exist too many char-
acteristic differences, of which the difference in the number
of antheridia is but one. The advantage of having but one
genus is not only exemplified in the mznzata group, but is
also shown in others. Not only are several distromatic spe-
cies monostromatic during some part of their existence, but
all monostromatic species become practically distromatic
when in fruit. Besides, the monostromatic nature of the
frond does not alter the external appearance by which
we can always readily recognize a member of the genus
Porphyra.

Sporocarps.—The formation of the sporocarps of P.
abyssicola is apparently identical with that of the carpospores
of P. miniata, so that, when ripe, each sporocarp contains
four carpospores (Pl. XXII, fig. 23).

Antheridia.—Like the sporocarps, the antheridia of P.
abyssicola agree entirely with those of P. mznzata, each an-
theridium containing when ripe eight antherozoids (PI.
PEI fig 2h)

Habitat.—On Zostera; also on rocks (fide Kjellman).

Distribution.—As yet known on the Pacific Coast from
but a single locality (48° 10’ N. lat.).

Bor.—Vot. II.] HUS—PORPHYRA. 225

Locality.—Whidby Island, San Juan County, Washing-
ton (N. L. Gardner, No. 273@!).

14. Porphyra variegata Aje//m. in litt.
PLATE XXI, Fic. 18.
Zoe, Vol. V, 1900, p. 69.

Porphyra variegata KJELLM. in litt.

Diploderma variegatum KJELLMAN, K. Sv. Vet. Akad. Handl., Bd. XXIII,
1889, p. 33, PI. Il, figs. 1-4.

Wildemania variegata DE Toni, Nuova Notarisia, Annata V, 1890, p. 148;
Sylloge Algarum, Vol. IV, 1897, p. 23.

Fronds membranous, 10-80 cm. long, 2-20 cm. broad, obovate to lanceo-
late, with slightly irregular margin, slightly undulate, base cuneate to obtuse,
or even cordate, sessile with small disc, areolate; color rich crimson, often
variegated; fronds distromatic, too-220y thick; cells in immediate vicinity of
attachment spherical or square with rounded angles, other vegetative cells
from 3-5 times as high as broad, 30-60 high, 3-304 broad, with rounded
angles; surface jelly thick, 20-45; cell-wall thick, composed of several layers
of jelly; fronds dicecious, sporocarps spreading over the whole frond, more
or less intermixed with vegetative cells; each sporocarp containing from 8 to
32 carpospores.

The average length of some hundred specimens of Por-
phyra variegata, collected at various localities on the Pacific
Coast is 20 to 25 centimeters. Several plants in our
herbarium fall below that mark, measuring from ten to
fifteen centimeters, while a few attain a length of from
forty to eighty centimeters. These measurements all refer
to dried specimens, and judging from the fact that plants of
this species, when soaked out, increase to about twice the
size of the dried specimen, it is supposed that the plants
contract considerably when drying; something which has
also been indicated by Kjellman (1889), who was able
to collect Porphyra variegata himself. The author was less
fortunate, and had to depend on dried specimens collected
by others. The impossibility of examining freshly gathered
specimens was keenly felt at first,—since specimens as soon
as soaked out disintegrated so rapidly as to make the prep-
aration of satisfactory sections out of the question. After
a number of experiments, it was found that if the dried

226 CALIFORNIA ACADEMY OF SCIENCES, [PRroc. 3D SER.

plants were treated with a boiling concentrated solution of
corrosive sublimate the cells retained their natural position
and size, at least sections from material treated in this man-
ner presented apparently a normal appearance. Asa mount-
ing medium twenty per cent. glycerine in water was found
to act very satisfactorily; but this was afterwards super-
seded by Farrant’s Medium, on account of the easier
manipulation.

The shape of the younger fronds of P. varzegata is ob-
ovate. The larger fronds are more or less lanceolate, and
are but seldom lobed, though a few cases were noted in
which the frond exhibited considerable lateral development,
together with a much laciniate appearance. The fronds are
slightly undulate and possess as a rule a cuneate to obtuse
base. A few specimens exhibited a cordate base. The
plants are attached by a small disc, surrounded by a slight
aureole. In this regard the plants differ from the descrip-
tion given by Kjellman, who calls them substipitate. How-
ever, a careful examination of our specimens has failed to
reveal a single instance of this.

The sterile frond has a red-purple color, but the fertile
frond presents a variegated appearance, especially towards
the center of the frond, while along the edges it is a rich,
uniform crimson, sharply contrasting with the yellowish
rim, some two to three millimeters broad, which surrounds
the fully ripe frond till near the base. This yellowish rim
consists of an empty network of gelatinous cell-walls, which
formerly contained the carpospores. The latter, when ripe,
left the frond because of the partial dissolution of the sur-
rounding cell-walls. This network is full of infusoria.

The crimson edges consist almost entirely of sporocarps,
with but few vegetative cells intermixed; while the varie-
gated appearance of the central part of the frond is due to
an intermixing of groups of deeply stained sporocarps with
a greater or lesser number of colorless vegetative cells. In
the sterile frond the vegetative cells are all colored, but not
intensely so. It has been suggested that the lack of color in

BoT.—VOL. II.] HUS—PORPHVRA. 227

the vegetative cells mixed with the sporocarps is due to the
absorption of the coloring matter by the reproductive cells;
but this hypothesis is not altogether satisfactory.

The fronds are distromatic throughout, and any one
frond is fairly constant in thickness. But among the
various fronds a great difference in thickness may be
noted, some measuring but 100 microns while others
measure as much as 220 microns. But if the thickness
of each frond is constant throughout, the size and shape
of the celis in the various parts of the frond are exceed-
ingly variable. While at the base, the cells, especially
those possessing the hypha-like projections which form
the disc, are spherical or pear-shaped, the cells a short
distance away from the place of attachment are square,
measuring in each direction from thirty to fifty microns.
But the vegetative cells among the fully ripe sporocarps
are from thirty to sixty microns high and but from three
to ten microns broad. They appear to have assumed this
shape owing to much lateral pressure, being much elongated
and ofted dumb-bell shaped (Pl. XXI, fig. 18).

The great thickness of the surface jelly and the peculiar
markings of the walls of the vegetative cells are character-
istic of this species. The cell-wall appears to consist of
layers of jelly of different density. These seem to denote
stages in the cells existence; but what these stages are
must at present be left to conjecture.

This species appears to be dicecious, no antheridia having
been found.

Sporocarps.—The first division of the cell destined to
become a sporocarp is cruciate, which is followed by a
division parallel to the surface of the frond in each of the
subdivisions of the sporocarp, thus giving rise to eight spo-
rocarps arranged in two tiers of four each. Often another
cruciate division takes place, giving rise to thirty-two carpo-
spores (Pl. X XI, fig. 18).

In Porphyra variegata the jelly, especially that which
forms the partitions between the individual carpospores,

228 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

seems to disintegrate more readily than in any other species
of Porphyra, with the exception perhaps of P. /eucosticta.
Consequently we find in the arrangement of the fully ripe
carpospores an irregularity such as is met with in the
arrangement of the antherozoids in P. leucosticta. The
carpospores arrange themselves along the lines of least
pressure, and consequently the whole group becomes spher-
ical and each carpospore more or less polygonal.

Habitat.—On rocks. Zone? August.

Distribution.—As far as the author is aware, this species
has been reported but once, and that by Kjellman (1889),
its author, from Bering Island. Judging from the table of
distribution in the paper above referred to, P. varzegata
seemed to be very local, and the impression was created
that it belonged to colder waters. Much surprise was there-
fore experienced when it was reported from San Pedro,
California. Later it was found at Coupeville, Washington
(55°-33° 40’ N. lat.). It was not observed in Alaska.

Localities. —Bering Island (Kjellman); Coupeville, Wash-
ington (N. L. Gardner, No. 177!, No. 179!); Monterey
Bay, California (C. L. Anderson!, C. P. Nott, No. 863],
Mrs. J. M. Weeks!); San Pedro, California (de A.
Saunders, No. 1206!, No. 1207!).

15. Porphyra occidentalis Setchell & Hus.
PLATE XXI, Fics. 150-170.

Porphyra occidentalis SETCHELL & Hus, Zoe, Vol. V, 1900, p. 69.

Fronds 15-30 cm. long, 1.5-5 cm. broad, linear, with slightly crenulate mar-
gin and a cuneate to orbicular base, sessile on small disc; color dull red;
fronds distromatic, vegetative part of frond, 45-75 thick, cells square or
once and a half to twice as broad as high, 12-15 high, 12-304 broad, surface
jelly 10-15 thick; apparently dicecious, antheridia forming a yellowish margin,
measuring 65-75 in cross-section, 64 antherozoids in 4 tiers of 16 antherozoids
in each antheridium.

This species is known to the author by but a few speci-
mens collected by Mrs. Weeks at Carmel Bay, Monterey
County, California. The plant has very much the external

Bot.—VOL. II.} HUS—PORPHYVRA., 229

appearance of a young frond of P. nereocystis, being linear
to slightly lanceolate in the more developed fronds. The
margins are slightly crenulate and more or less irregular,
but the frond is not lobed or even laciniate. The base is
cuneate to slightly orbicular, exhibiting a slight aureole
around the point of attachment. Attachment is by means
of a disc.

The distromatic nature of the frond readily distinguishes
the plant from P. nereocystis. The cells are square or from
once and a-half to twice as broad as high. The surface
jelly is thick (Pl. XXI, figs. 15@ and 150).

The specimens examined show only antheridia, and from
this it is supposed that the plant is dicecious. The anther-
idia are found towards the tip of the frond and along the
edges. They evidently spread gradually over the whole
frond.

The antheridia form the chief characteristic of the
species. The antherozoids are arranged in groups such as
are found in no other species, and exhibit a marked regularity
in the division. As in all species of Porphyra, the anther-
idium-mother-cell, to produce the antheridia, undergoes a
cruciate division perpendicular to the surface of the frond.
The first reproductive division (in the antheridium) is par-
allel to the surface of the frond. This is followed by a
cruciate division perpendicular to the surface of the frond,
after which another parallel and another cruciate division
take place, thus giving rise to sixty-four antherozoids in
four tiers of sixteen each (Pl. XXI, figs. 16, 17a, 176. PI.
XXII, fig. 27).

When the antheridia are fully ripe, the jelly-walls between
the antherozoids dissolve before the surface jelly goes into
dissolution, so that the antherozoids are arranged more or
less irregularly in rectangular groups.

flabitat.—On rocks. Zone?

Distribution.—As yet reported from but a single locality
on the Pacific Coast (36° 45’ N. lat.).

Locality Carmel Bay, Monterey County, California
(Mrs. J. M. Weeks!).

230 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

V. Economic USEs.

It is generally known that in Europe certain seaweeds are
used as food and even sold in the markets, under the name
of ‘‘laver.’’ One of these seaweeds is a species of Por-
phyra (P. laciniata). Kjellman (1897) devotes a part of his
paper to a highly interesting account of the use of Porphyra
species in Japan. From this it was learned that not only are
the plants collected, but they are even cultivated, and after
harvesting are carefully cleaned and prepared for market.

This led the writer to start on a voyage of discovery
through Chinatown, San Francisco, to find out, if possible,
whether the Chinese also use the species of Porphyra which
occur in such abundance on our coast. It was found that
formerly the Chinese used to depend altogether on Porphyra
imported from Japan, but that now P. ferforata and P.
nereocystis are gathered in large quantities, especially at
Monterey, and are partly consumed here and partly exported
to China. The Chinese understand that there is some dif-
ference between the two species, having different names for
them, and seemingly esteeming P. nereocyst7s more highly
than P. perforata, at least the latter costs only about one-
fifth the price of the former. Perhaps the distinction is
identical with the distinction made by the Chinese between
the plants collected from rocks and plants collected from
LNVereocystis.

Othes species of Porphyra, as yet undetermined, are sold
in Chinatown in flat, round, purple-colored cakes, about one
foot in diameter and from one-half to one-fourth of an inch
thick. They are exported from Japan for Chinese use.
This product forms a sharp contrast with ‘‘Asakusa Nori,”’
also made of Porfhyra sp. and exclusively used by Japan-
ese. This is sold in bundles of ten sheets each, each sheet
being about eight inches square, exceedingly thin, and light
brown in color. While the latter are scrupulously clean
and freed from all foreign alga and animal matter, the
product used by the Chinese contains other algz as well as
numerous molluscs.

Bot.—VOL. II.] HUS—PORPHYVRA. 231

The Indians of the slopes of the Pacific Coast also use
Porphyra as a food, the tribes making yearly trips to the
seashore to collect it, along with other economic seaweeds.

VI. METHODs.

On account of the fact that in most of the species of
Porphyra examined the jelly makes up a large part of the
frond, considerable difficulty was encountered before the
writer finally succeeded in preparing the necessary number
of mounts. The specimens collected fresh were either
dried or killed on the shore in Flemming’s mixture, strong
and dilute, one per cent. chromic acid, one per cent., two
per cent. and five per cent. chrome alum, Wilson’s corro-
sive sublimate, one per cent. formalin and alcohol. Of all
these, Wilson’s solution, formalin and alcohol, gave the
best results, the others either shrinking the specimens or
failing to fix them. As soon as this point was settled, all
except the latter three were discarded. In some cases, as
in P. natadum, formalin gave the best results; while in
others, Wilson’s solution was to be preferred.

The necessity of the salt of a heavy metal to fix the jelly
became apparent when rough dried and herbarium speci-
mens were examined. The first ones to be studied were
dried specimens of P. perforata which, when soaked in
water, resumed their original outward form. But when the
same process was tried with others, especially with P.
variegata, it was found that the jelly dissolved very readily,
so that some means had to be resorted to by which the
specimens might be preserved in good condition. At first,
glycerin, twenty per cent. glycerin in water, and twenty
per cent. alcoholic glycerin were tried, but without good
result. Finally Professor Setchell suggested the use of a
saturated aqueous solution of corrosive sublimate. When
used cold, this answered all requirements in every case ex-
cept that of P. varzegata, where it was necessary to use a
boiling saturated aqueous solution.

Of all species of which fresh material could be obtained
sections of from three to five microns in thickness were

232 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

made on the Minot microtome. To prepare the material
for this purpose, small parts of the frond of which a sec-
tion was desired were taken from the Wilson’s solution in
which they had been left for about six hours, or from the
formalin in which they had remained an indefinite time, and
placed in ten per cent. alcohol from which they were dia-
lyzed into 95 per cent. alcohol. The specimens remained in
95 per cent. alcohol for an indefinite time, but never less
than twenty-four hours. The parts of the frond were then
folded in small bundles of about one-fourth of an inch or
less square and one-eighth of an inch in thickness; after
which they were transferred for three hours to 100 per
cent. alcohol. From this they were placed in a mixture of
equal parts absolute alcohol and bergamot oil. In this they
remained two hours, as they did in the bergamot oil to
which they were transferred. From the bergamot oil they
were passed into 45° paraffin, a mixture of equal parts of
45° and 54° paraffin, and 54° paraffin, being during that
time placed in a water-bath which was kept at a constant
temperature of 56° C. In each of these they remained
from two to three hours.

After this they were imbedded in 54° paraffin and on
cooling were ready for cutting. In most cases good results
were obtained.

This method was extremely useful in determining the
number of divisions in the antheridia, which in compara-
tively thick razor sections with pith was very difficult to do.

Razor sections were usually stained with eosin, while
microtome sections were stained with acid fuchsin or with
safranin. Of the two, the latter gave the better results.

In a single case, that of P. nazadum, a double stain,
methyl-blue-safranin, was used with exceedingly gratifying
results. The slides were passed from xylol into absolute
alcohol, 95 per cent. alcohol, and a saturated alcoholic solu-
tion of safranin. Here they remained for about half an
hour, after which the slides were placed for five minutes in
95 per cent. alcohol to which some drops of methyl-blue,
enough to give the alcohol a deep blue tinge, had been

Bor.—Vot. II.] HUS—PORPHYRA. 233

added. After this they were transferred to absolute alcohol,
xylol, and finally mounted in Canada Balsam. On examin-
ation the slides showed that while the cushion-shaped base
of P. natadum stained a bright red, the tissue of the host-
plant, Phyllospadix, was stained blue, as were the chromat-
ophores and cell-contents of P. nazadum. In some cases,
where the slides had remained longer than five minutes in
the methyl-blue, the young fronds emerging from the
cushion-shaped bases were stained purple.

The ribbon was fixed on the slides with albumen, and
floated out in water and placed on the top of the water-bath
where the slides remained till dry.

As regards mounting media, dilute glycerin jelly and
Farrants’ Medium gave excellent results. While the
former can be used to great advantage in preparing
surface views, the latter is to be preferred where razor
sections are to be mounted. It appears to have a clearing
action on the tissues. Great care must be taken to wash out
all corrosive sublimate in alcohol to which some potassium
iodide is added, while hardening preparatory to cutting,
since even the slightest trace of mercuric chloride will
after a time attack the mounting medium and destroy the
preparation.

In conclusion, the author begs to acknowledge his in-
debtedness to Professor W. A. Setchell, at whose sugges-
tion the work was undertaken, and whose advice has been
of invaluable aid in compiling this paper. Thanks are also
due Professors Rosenvinge, Kjellman and Farlow, Messrs.
Bornet, Collins, Janczewski, de A. Saunders, Lawson,
Gibbs and Nott, and Mrs. Weeks and Mrs. Snyder, for
specimens received.

BOTANICAL LABORATORY,

UNIVERSITY OF CALIFORNIA,
BERKELEY, CALIFORNIA.

234

1821.
1824.

1828.
1882.

1899.

1891.

1892.

1882.

1882.
1884.
1890.

1897.
1881.

CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

LITERATURE AND EXSICCATZE CITED.

AGARDH, C. A. Species Algarum. Gryphiswaldiz.

Systema Algarum. Lund.

Icones Algarum Europzearum. Leipzig.

AGARDH, J. G._ Till Algernes Systematik, Afd. 3, VI, Ulvacez.
Lund’s Univ. Arsskrift, Tome XIX. Lund.

Analecta Algologica, Continuatio V. Acta Reg. Soc. Physiogr.
Lund., Tome X. Lund.

ANDERSON, C. L. List of California Marine Algz, with Notes. Zoe,
Vol. II. San Francisco.

-Algze of the Farallons. (In Blankinship and Keeler: ‘‘On the
Natural History of the Farallon Islands’’). Zoe, Vol. III. San
Francisco.

BERTHOLD, G. Die Bangiaceen des Golfes von Neapel und der
angrenzenden Meeres-Abschnitte. Fauna und Flora des Golfes von
Neapel und der angrenzenden Meeres-Abschnitte, herausgegeben
von der zoblogischen Station zu Neapel. Leipzig.

Coins, F.S. Notes on New England Alge. Aull. Torrey Bot.
Club, Vol. 1X. New York.

Notes on New England Alge, IV. Bull. Torrey Bot. Club,
Vol. XI. New York.

DE Tonl, J. B. Frammenti Algologici. La Nuova Nolarisia, Ano I,
No. 3. Padova.

Sylloge Algarum, Vol. 1V. Padova.

FarLow, W. G. Marine Alge of New England. Report U. S.
Comm. Fish and Fisheries, 1879. Washington.

1890-1891. Fosire, M. Contribution to Knowledge of the Marine Algz of

Norway, land Il. Zvromsé Museums Aarshefter, XIII. Tromso.

1846-51. Harvey, W. H. Phycologica Brittanica. London.

1849.
1858.
1893.
1900.
1873.
1883.
1889.

1897.

1864.

British Marine Algze. London.

Nereis Boreali-Americana, Part III. Washington.

Howe, M. A. A Month on the Shores of Monterey Bay. Avrythea,
Vol. I. Berkeley, California.

Hus, H. T. A. Preliminary Notes on West Coast Porphyras. Zoe,
Vol. V. San Francisco.

JANCzEwsk1, E. pe. Etudes Anatomiques sur les Porphyra. Amn.
Sct. nat. (Bot.), 5¢ Sér., Tome XVII. Paris.

KJELLMAN, F. R. The Algze of the Arctic Sea. XK. Svenska Vet.-
Akad. Handl., Band 20, No. 5. Stockholm.

Om Beringhafvets Algflora. A. Sv. Vet.-Akad. Handl., Band

23, No. 8. Stockholm.

Japanska Arter af Slagtet Porphyra. Azhang till K. Svenska
Vetenskaps-Akademiens Handlingar, Band 23, Afd. III, No. 4.
Stockholm.

LE Jouis. Liste des Algues Marines de Cherbourg. Mem. Soc. Imp.
Sct. nat. de Cherbourg., Tome X. Paris.

i,

Bot.—VOL. II.] HUS—PORPHYRA. 235

1819.
1866.

1897.

1893.

1878.

LyncsyeE, H.C. Tentamen Hydrophytologiz Danice. Copenhagen.

Martens, G. von. Die Preussische Expedition nach Ost-Asien.
Botanischer Theil. Die Tange. Berlin.

McCrarcuie, A. J. Seedless Plants of Southern California. Proc.
Southern Calif. Acad., Vol. 1. Los Angeles.

ROSENVINGE, L. K. Grgnlands Havalger. Meddelelser om Grgn-
land, I77. \Kjobenhavn.

Les Algues Marines du Grgnland. Ann. Sci. nat. (Bot.) 7e
Sér., Tome XIX. Paris.

Ruprecut. Neue oder unvol. bek. Pflanzen a. d. nérdlichen Theile
des Stillen Oceans. J7ém. Acad. de St. Petersb. sc. natur., Tome
VII. St. Petersburg.

Scumitz, F. Die Chromatophoren der Algen. Bonn.

SETCHELL, W. A. Algz of the Pribilof Islands. The Fur Seals and
Fur-Seal Islands of the North Pacific Ocean, Part II]. Washington.

STROMFELT, H. F.G._ Einige fiir die Wissenschaft neue Meeresalgen
aus Island. Sot. Centralblatt, Bd. XXVI. Cassel.

Om Algvegetationen vid Islands Kuster. Gé6tebore.

SuRINGAR, W. F. R. Algze Japonice Musei botanici Lugduno-
Batavi. Harlemi.

Tuuret ET Borner. Etudes Phycologiques. Paris.

1895-I901. CoLLIns, HOLDEN & SETCHELL, Phycotheca Boreali-Americana,

Fascl. I-X VII, and 4-2. Malden, Mass.

1885-1896. Hauck und RicuTer, Phycotheka Universalis, Fascl. I-XV.

Leipzig.

1894-1901. TILDEN, American Algz, Cent. I-V. Minneapolis.

236

CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

EXPLANATION OF PLATE XxX.

Figs. 1-6, and 11-14, drawn with Abbé’s camera lucida, obj. Leitz ,

ocular No. 3.

Figs. 7-10 drawn with Abbé’s camera lucida, obj. Leitz 4, ocular No. 1.

Fig. ra.
Fig. 10.
Fig. 2a.
Fig. 20.
Fig. 3a.
Fig. 30.

Fig. 4a.
Fig. 46.
Fig. 5a.
Fig. 50.

Fig. 6.

Porphyra leucosticta,

Surface view of vegetative part of frond.
Cross-section of vegetative part of frond.
Surface view of sporocarpic part of frond.
Cross-section of sporocarpic part of frond.
Surface view of antheridial part of frond.
Cross-section of antheridial part of frond.

Porphyra perforata.

Surface view of vegetative part of frond.
Cross-section of vegetative part of frond.
Surface view of sporocarpic part of frond.
Cross-section of sporocarpic part of frond.
Cross-section of antheridial part of frond.

Figs. 7-10. Cells of the base, showing hypha-like projections; after treatment

Fig. 11a.

with Schulze’s Macerating Fluid.

Porphyra nereocystis.

Surface view of cells of vegetative and sporocarpic part of frond.

Fig. 116. Cross-section of vegetative part of frond.

Fig. 1c.
Fig. 12.

Cross-section of sporocarpic part of frond.
Cross-section of antheridial part of frond.

Porphyra amplissima.

Fig. 13a. Surface view of antheridial and sporocarpic part of frond.
Fig. 136. Cross-section of antheridial and sporocarpic part of frond.

Fig. 14.

Porphyra miniata f. cuneiformts.

Cross-section of antheridial and sporocarpicspart of frond.

Proc.CALACAD. Sci.3? SER. Bor. VoL II

Janusog » —————
Sos TMT
o> ais 2)

ALOE M3 anyonanien
WESO : make
OSB (\ 38 2¢

BSG{Cwlos

Bi) US SS = es
<eslogy She
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5) CEN 30 com mop LOPSx aD

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5 Ay Se BS & ae SS So
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46

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RE
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[Hus] PLATE XX.

simian

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Ph Gee as

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“ woes, ' ; P . ; i j
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238 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

EXPLANATION OF PLATE XXI.

Figs. 15-18, and 20-22, drawn with Abbé’s camera lucida, obj. Leitz },
ocular No. 3.

Fig. 19 drawn with Abbé’s camera lucida, obj. Leitz 3, ocular No. 3.

Porphyra occidentalis.

Fig. 15a. Surface view of vegetative part of frond.

Fig. 156. Cross-section of vegetative part of frond.

Fig. 16. Cross-section of antheridial part of frond, showing stages in the
development of the antheridia.

Fig. 17a. Surface view of antheridial part of frond.

Fig. 174. Cross-section of antheridial part of frond.

Porphyra variegata.

Fig. 18. Cross-section of vegetative and sporocarpic part of frond.

Porphyra naiadum.

Fig. 19. Cross-section of blade of Zostera and of base of frond of P.
naiadum.

Fig. 20a. Surface view of vegetative part of frond.

Fig. 206. Cross-section of vegetative part of frond.

Fig. 21a. Surface view of sporocarpic part of frond.

Fig. 216. Cross-section of sporocarpic part of frond.

Fig. 22. Cross-section of young frond with base.

[Hus] PLATE XXI

nga
i

i
(

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anys
Sh at
i}

%
<4}
es

Si
SAG
aan
ESSA AY
LESS
AN

ie
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Sse PANS

99 Oa \R8 ER

PBBENYS Cd

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‘
?,
re oe Se ee

Vy

a eS. Sow Ae

240 CALIFORNIA ACADEMY OF SCIENCES. [PROC. 3D SER,

EXPLANATION OF PLATE XXII.

Diagrams illustrating Formation of Reproductive Bodies.

Fig. 23. Cube, representing cell of Porphyra sp., which has undergone a
cruciate division perpendicular to the surface of the frond. For-

mula for sporocarps, 4 (& Lx c ) ;
Derg

Fig. 24. The same, after undergoing a division parallel to the surface of the
frond in each segment. Formula for sporocarps, 8 (4 & =)

2 se
Fig. 25. The same after undergoing another cruciate division perpendicular

to the surface of the frond in all segments. Formula for sporo-

ae 4
carps, 32 ( m7 <) ; for antheridia, 8 (4%, as =),

Fig. 26. The same after undergoing another division parallel to the sur-
face of the frond in all segments. Formula for sporocarps,

64 eS is =e for antheridia, 16 Cae Ye 4B =a
A de A 3 2 Zui a

Fig. 27. The same after undergoing another cruciate division perpendicular
to the surface of the frond in all segments. Formula for anther-
dais epee eee SES

4 ft 4

Fig. 28. The same after undergoing another division parallel to the sur-
face of the frond in all segments. Formula for antheridia,
128 (Gee, ho —) :

4 4 §

eA
OA

SeGESERiars:
BEaMESaay)

S Ll Yeeeah
YS BSE,

PrRoc.CALACAD. Sc1.3? SER. Hor. Vou II

n

OF REPRODUCTIVE Rooles oF POoRPHYRA

PORMATION

e

OR ed

SOME NEW SPECIES OF PACIFIC COAST RIBES.

BY ALICE EASTWOOD.

Curator of the Department of Botany.

PLATES XXIII AND XXIV.

TueE genus /’zdes is represented on the Pacific Coast by
four subgenera, which include an indefinite number of
species and forms.

A recent attempt on the part of the author to identify the
unnamed specimens of this genus, which had for years been
accumulating in the Herbarium of the California Academy
of Sciences, has brought to light the following species
which seem to be undescribed.

The first six belong to the suborder Ribesia; of these,
numbers one and two fall into the group typified by /tzdes
sanguineum Pursh, number three into that typified by A.
malvaceum Smith; numbers four, five and six into that typi-
fied by #. nevadense Kellogg. The last three belong to
the suborder Grossularia.

Some botanists might include the first six under /’zbes
sanguineum, the last three under /t. menzzeszz, ‘This aggre-
gation may satisfy the amateur to whom generic differences
are sufficient, but the real student desiring to learn the truth
regarding a genus will find it a source of great confusion,
and altogether unsatisfactory.

While it is to be kept in mind that nature knows no
boundaries, and that orders, genera, and species are divi-
sions made by man for his own convenience, yet these
methods of classification have a scientific value beyond that
of pure utilitarianism, and ought to show as far as possible
the life-history of a group of related plants and of the entire
plant world, when the knowledge of man makes it possible.

[ 241 ] April 14, 1902.

242 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

1. Ribes brandegei, sp. nov.
PLATE XXIII, Fics. 1a AND 10.

Shrub with erect, branching stems unarmed; older bark dull brown; young
bark glossy, not shreddy. Leaves three-lobed, reniform, 3-4 mm. long, and
of about equal width; upper surface sparsely pubescent with fine, silky hairs,
mostly on the veins, and with scattered sessile or shortly stipitate glands;
lower surface pale green, with appressed, glandular hairs, veins at base vil-
lous; margin incised-dentate, glandular-ciliate; petioles generally shorter than
the blades, glabrous or clothed with a fine pubescence under the gland-tipped
hairs; stipular dilation 3 mm. broad, fringed with uneven, glandular hairs.
Inflorescence racemose, erect in flower, the peduncles equalling or longer
than the flowering portion, generally surpassing the leaves; pubescence as
on the petioles; bracts foliaceous, oblanceolate to obovate, acuminate,
incised, glandular-ciliate; flowers three to ten, on slender, erect pedicels
which later become as long as the flower. Calyx rose-color, 8 mm. long,
pubescent on the outside, glandular at base, puberulent within; divisions as
long as the tube, oblanceolate to obovate, cucullate at summit, 2 mm. wide.
Petals white, half as long as the calyx divisions, 1.5 mm. wide, orbicular-
spatulate, on short claws. Stamens with slender filaments, 1.5 mm. long;
anthers oblong, tipped with a blunt mucro. Style two-cleft at apex, with the
stigmas broad. Berry glabrous, globular.

This species is related to Azbes sanguineum Pursh from
which it differs in the pubescence, inflorescence, and shape
of floral organs, as can be seen by the figures.

Collected by Mr. T. S. Brandegee, in whose honor the
author takes pleasure in naming it, first at Sierra de Laguna,
Lower California, January 25, 1890, later in the mountains
of the Cape Region, March 26, 1892.

2. Ribes scuphami, sp. nov.

PLATE XXIII, Fics. 2a AnD 26.

Shrub with the upper bark reddish, shreddy, puberulent, unarmed. Leaves
orbicular, three- to five-lobed, truncate to reniform at base, 2-5 cm. wide,
about as long, unevenly dentate; upper surface pubescent with crisp, spread-
ing hairs; lower, canescent with matted hairs; stipular dilation of the petiole
broad, glandular, and tomentose, fringed with glandular hairs; petioles about
as long as the blades, with pubescence like the stipules. Racemes numerous
at the ends of the branches, 9 cm. long, slender, when flowering erect on
peduncles which are shorter than the leaves; bracts oblanceolate, red, gland-
ular, 8 mm. long, denticulate at apex; pedicels filiform, erect, a little longer
than the bracts. Flowers subtended by two small, red bracteoles which
are soon deciduous. Calyx rose-color, with tube 5 mm. long, divisions

a

Bot.—VOEz; I{f-] EASTWOOD—PACIFIC COAST RIBES. 243

linear-oblong, 7 mm. long. Petals white turning reddish, oblanceolate, cune-
ate, 4 mm. long. Stamens a little shorter than the petals; anthers globular.
Ovary sparingly pubescent, and with scattered, stipitate glands.

This is nearest to bes sanguineum Pursh. It differs
especially in having the racemes erect in flower, also in the
more slender flowers with narrowed divisions. This species
is the most beautiful of all belonging to the group of which
LF. sanguineum is the type.

It was collected on Smith River, Del Norte County,
California, by Major J. R. Scupham, May, 1808. It is
a pleasure to name this plant in honor of one who has
brought many interesting plants to the herbarium of the
California Academy of Sciences from little explored parts
of California.

3. Ribes indecorum, sp. nov.
PLATE XXIII, Fics. 3a AnD 36.

Shrub with erect stems, having dark brown, shreddy bark on the older
growth, the younger parts tomentose and glandular. Leaves three-lobed,
2-4 cm. long, 2-3 cm. wide, finely rugose on the upper surface, clothed with
stipitate glands, and a fine, sparse, silky pubescence; lower surface white
with a felt-like tomentum, and with a few gland-tipped hairs on the veins;
margins irregularly, doubly crenate; petioles stout, shorter than or equalling
the blades, glandular and tomentose, the stipular dilation (as wide on each
side as the petiole) fringed on the margin with uneven, gland-tipped hairs.
Inflorescence racemose, spreading or pendent, in fruit surpassing the leaves;
flowers sessile but erect; peduncles short; bracts foliaceous, almost equalling
the flowers, lanceolate, 6 mm. long, 2 mm. wide, with the margins fringed
with long, gland-tipped hairs. Peduncles stout, glandular, and tomentose.
Flowers at base subtended by two membranous, glandular, and tomentose
bracteoles; calyx-tube more than twice as long as the broad, rounded divi-
sions; these tomentose and glandular on both sides, almost 2 mm. wide;
petals orbicular, reniform, 1 mm. wide, crenulate, on very short and broad
claws. Stamens as long as the petals, on stout, short, deltoid filaments;
anthers .75 mm. long, longer than the filaments. Style stout, hairy at base,
two-cleft at apex, with broad, yellow stigmas; ovary tomentose and somewhat
glandular.

Collected by the author at Cajon Heights, near San
Diego, California, March 14, 1891. There is also a
specimen in the Herbarium of the California Academy of
Sciences collected by Dr. George Thurber at San Pasqual,
San Diego County. It is labeled Azbes sanguineum, No. 606.

244 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

Pibes indecorum is nearest to Azbes malvaceum but differs
most noticeably in the much smaller and sessile flowers.
The floral organs, too, are not the same.

4. Ribes ascendens, sp. nov.
PLATE XXIII, Fics. 4a AND 46.

Erect shrub unarmed, with gray-brown bark on older stems, younger stems
paler and shreddy. Leaves three- to five-lobed, orbicular, reniform, 3-6 cm.
wide, 2-5 cm. long, crenate-dentate, almost glabrous on the upper surface,
the lower clothed with fine, spreading pubescence; petioles equalling or
shorter than the blades, glandular; stipular dilation narrow, fringed with
long, gland-tipped hairs. Peduncles generally surpassing the leaves, at first
erect, later nodding, glandular-pubescent; flowers crowded at the summit of
the peduncle, which is naked for more than half its length; bracts oblanceo-
late, rounded at apex, 7 mm. long, 2 mm. wide, with gland-tipped hairs on
the surface and margin; pedicels half as long as the bracts, lengthening with
age, and recurving upwards, so that the berries are erect. Flowers subtended
by two membranous bracteoles which are soon deciduous. Calyx open-
campanulate, rose-color, the tube about half as long as the divisions; these
ovate, obtuse, 3.5 mm. long, 2.5 mm. wide, slightly pubescent. Petals white,

‘orbicular, narrowed to a short, broad claw, 2 mm. wide. Stamens not equal-

ling the petals, filaments linear, anthers oblong. Ovary clothed with gland-
tipped hairs. Berry veiny, sparingly glandular, becoming 7 mm. or more in
diameter.

This species is near 7’. xevadense Kellogg, but the racemes
are ascending when in flower. The floral organs also differ
in shape.

The type was collected by the author at Millwood,
(Sequoia Mills) Fresno County, California, in flower, May
4, 1895; in fruit, July 18,1893. There are specimens from
General Grant Grove in the same vicinity, and from Coburn’s
Mills in Fresno County, collected by T. S. Brandegee; the
former, July, 1892, the latter, May 29 (year not given).

4a. SLRibes ascendens jaspere, var. nov. (Plate XXIV,
fig. 5). This is similar to the species in general appear-
ance, but the sepals and petals are more orbicular, the
filaments are shorter, and the anthers broader and more
orbicular. It is named in honor of Mrs. William Jasper,
who sent the specimen from San Emidio Cafion, Kern
County, California, May, 1895.

Bot.—VOL. II.] EASTWOOD—PACIFIC COAST RIBES. 245

5. Ribes hittellianum, sp. nov.
PLATE XXIV, FIGs. 6a AND 60.

Erect shrub with spreading branches, 2-3 feet high; bark smooth, un-
armed, gray-brown, shreddy on the younger branches. Leaves three- to
five-lobed, orbicular, reniform or truncate at base, 3-4 cm. wide, 2-3 cm.
long, irregularly dentate and somewhat revolute, rugulose veiny, glabrous on
both sides but with some scattered glands on the lower; petioles about as
long as the blades, sparingly tomentose and glandular; stipular dilation
broad, truncate, membranous, as wide on each side as the petiole. Racemes
1-2 cm. long, at first erect, later nodding, but with the pedicels erect; flowers
three to eight, crowded, subtended by deciduous bracteoles. Calyx tubular-
campanulate, with the tube 1 mm. long, the divisions rose-color, oblong,
obtuse, 4 mm. long, 2 mm. wide. Petals white, narrowly oblong, three-
fourths as long as the sepals and one-half as wide. Stamens with subulate
filaments, half as long as the sepals; anthers orbicular. Stigmas two, capitate;
ovary and immature fruit clothed with stipitate glands.

This species belongs in the group of which A. nevadense
Kellogg is the type. It differs from that species in the
inflorescence and the shape of the floral organs.

Collected near the head-waters of Cafion Creek, Trinity
County, California, not far from Twin Lakes, July 9, 1901,
and named in honor of Mr. Carlos T. Hittell, one of the
party on a trip to these little known mountains.

6. Ribes glaucescens, sp. nov.
PLATE XXIV, Fics. 7a AND 76.

Unarmed shrub with older bark gray-brown, younger bark bright brown
glossy, shreddy. Leaves three-lobed, orbicular-reniform, about 3 cm. long,
3-5 cm. wide, irregularly dentate, glabrous except for some minute glands on
the lower surface, glaucescent, paler on the lower surface; petioles about as
long as the blade, minutely puberulent, with the stipular dilation on each side
narrower than the petiole, and sparingly fringed with glandular hairs. Inflor-
escence in fruit spreading or erect, generally shorter than the leaves, rather
loosely flowered with from five to ten flowers; peduncles as long as the raceme,
striate, puberulent; pedicels slender, becoming 5 mm. long, shorter than the
brown, membranous, gland-tipped bracts. Flowers subtended by two small,
deciduous, reddish bracteoles. Calyx open-campanulate, with very short
tube, and spreading divisions; these rose-color, oblanceolate, 4 mm. long,
I.5 mm. wide, glabrous. Petals white, spatulate two-thirds as long as the
sepals, denticulate near the apex. Stamens half as long as the sepals, with
suborbicular anthers and broad filaments. Ovary glabrous except for the
scattered stipitate glands.

246 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

This species is related to Azbes nevadense Kellogg from
which it differs in the glaucous color of its foliage, the
racemes erect in fruit, and the shape of the floral organs.
It was collected by the author on Mount Shasta, August 13,
1893. No note was taken of the exact locality, but it must
have been some place on the trail from Sisson.

7. Ribes oligacanthum, sp. nov.
PLATE XXIV, Fics. 8a AND 80.

Erect, branching shrub with the younger stems puberulent, older gray-
brown, not shreddy, somewhat tortuous, unarmed, except for the simple or
triple thorns at the leaf-axils; these often short and abortive. Leaves broadly
ovate or orbicular, three- to five-lobed, 2-3 cm. wide and about as long, with
margin laciniate-dentate, base truncate, but cuneate where it joins the petiole,
glabrous; petioles slender, as long as or twice as long as the blades, glabrous
or puberulent; stipular dilation twice as wide as the petiole, ciliate with white,
silky hairs. Peduncles filiform, one- to three-flowered, 1-2 cm. long; pedicels
almost as long, together becoming 4 cm. long, slightly puberulent; bracts
broadly ovate to orbicular, acuminate, three-angled, clasping, reflexed.
Calyx 12 mm. long, the linear, acute segments more than twice as long as
the tube, 2 mm. wide, apparently white, veined with parallel veins that
branch near the apex, puberulent or glabrous. Petals involute, obovate
when spread out, irregularly denticulate at apex, 3 mm. long, veins palmate.
Filaments almost twice as long as the petals, dilated at base; anthers oblong-
ovate, cordate at base, tipped with a recurved mucro. Style divided two-
thirds of its length, slightly exserted. Young fruit puberulent, clothed with
a few long spines (about ten) each about 5 mm. long.

Related to Arébes californicum Hook. & Arn. from which
it differs in the sparsely spinous fruit, the glabrous leaves,
the lax inflorescence, and the shape of the floral organs.

Collected by the author on the road between Jolon and
King City, in Monterey County, California, near Mans-
field’s Ranch, ten miles from King City, May, 1897.

8. Ribes sericeum, sp. nov.
PLATE XXIV, Fics. ga-9/f.

Erect, branching shrub, several feet high; stems clothed with numerous
fine, weak, short prickles, which are gland-tipped on the young shoots, also
with short, close, silky pubescence; axillary thorns three, orange-color, stout,
united, the middle one longest, more than 1 cm. long, broadening at the

BoT.—VOL. II.] EASTWOOD—PACIFIC COAST RIBES. 247

base, pubescent and glandular on the lower part, glabrous on the upper.
Leaves thin, three- to five-lobed, broadly ovate-orbicular, reniform or trun-
cate at base, 2-4 cm. long, not quite so wide, incised-crenate, clothed with
fine, white, silky hairs which are appressed or spreading, also with fine
gland-tipped hairs; petioles about as long as the blades, more glandular and
more spreading-pilose, dilated only at the very base, and without the appear-
ance of stipules. Peduncles one- to three-flowered, slender, erect, with
pubescence like the petioles; pedicels about half as long; bracts orbicular or
three-lobed, foliaceous; bractlets similar but smaller. Flowers 2 cm. long,
open-campanulate in the bud. Calyx with the divisions at length reflexed,
longer than the tube and the ovary, oblong, purplish red, greenish near the
apex, softly silky villous on both sides; tube campanulate, veined, slightly
glandular at base. Petals white, 5 mm. long, involute, erose along the almost
truncate apex. Stamens with filiform, purple filaments, exserted beyond the
sepals in the opening flower, and also beyond the pistil; anthers narrowly
linear-oblong, almost 2 mm. long, obtuse. Pistil two:cleft for 2 mm. Ovary
densely clothed with horizontally spreading fine, silky hairs mixed with some
longer, glandular hairs, the glands purple. Fruit purple, clothed with short,
weak bristles and scattered hairs. Some of the bristles retain the purple
glands on the fruit.

Collected in flower by Mr. R. A. Plaskett, at Spruce
Creek; also at Gorda, in flower and fruit. Collected by
the author at Pacific Valley, with immature fruit. . serz-
ceum flowers in December and January and fruits in June.
At Point Sur specimens were collected by the author in
June, 1893, with very large, pear-shaped fruit, almost 4 cm.
long, and specimens with globular fruit were collected at
about the same time at Slate’s Hot Springs. All these
localities are on the coast of Monterey County, California,
at the base of the Santa Lucia Mountains, and the range
extends from south of Point Gorda to north of Point Sur.

Libes sericeum is related to PR. subvestitum Hook & Arn.
but it has different leaves, different pubescence, and the
floral organs are not the same.

Sa. Libes sericeum viridescens, var. nov. The variety
is similar to the type, but the flowers are smaller and green-
ish, the leaves are more densely clothed with silky white
hairs, and are more orbicular-reniform. The peduncles in
the specimens examined all have single flowers.

This variety was collected by R. A. Plaskett at Gorda,
Monterey County, California, January, 1808.

248 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

g. Ribes hystrix, sp. nov.
PLATE XXIV, Fics. toa-tod.

Shrub several feet high, with light brown, tortuous branches, minutely
pubescent and thickly beset with stout, rigid, horizontal, yellow prickles,
some gland-tipped, generally small on the new growth, and increasing in size
with age; axillary thorns triple, stout, distinct at base, middle one longest,
becoming 15 mm. long, lower part pubescent, upper, glabrous. Leaves thin,
three-lobed or some five-lobed, with the basal lobes small, 2-4 cm. wide,
orbicular-reniform, incisely dentate, minutely pubescent and dotted with
sessile glands on the lower surface, almost glabrous on the upper; petioles
about as long as the blade, tomentose and slightly glandular. Peduncles
one- to three-flowered, 1-2 cm. long, ascending, slender, sparingly pilose
and clothed with gland-tipped hairs; pedicels less than half as long, occasion-
ally longer; bracts orbicular or lobed, clasping, acuminate to obtuse. Calyx
pubescent and glandular; tube a little longer than the ovary; divisions 1 cm.
long, surpassing the rest of the flower when reflexed, 3 mm. wide, lower
part purple, near the apex greenish, obtuse. Petals white, broadly obovate
when spread out, acute, narrowed to a short claw, involute, 4 mm. long.
Stamens with broad filaments dilated at base, as broad and long as the
anthers; these about 3 mm. long, sagittate at base, tipped with a blunt mucro.
Styles surpassing the sepals in the opening flower, divided about half; stigmas
small, capitate; ovary globular, tomentose, and densely clothed with purplish
bristles, some near the calyx gland-tipped. Fruit purple, more or less
densely clothed with stiff, spreading prickles, 2-5 mm. long.

This species is nearest to A. menzzeszz Pursh but differs
in the glandular pubescence, the shape and texture of the
leaves, the size of the flower, and shape of the parts.

Collected in flower by Mr. R. A. Plaskett, at Gorda,
Santa Lucia Mountains, California, December, 1897. The
fruiting specimens were collected by the author at Pacific
Valley, in the same vicinity, May, 1897, and June, 1893.

Bot.—VOL. II.] EASTWOOD—PACIFIC COAST RIBES. 249

KEY TO THE SPECIES OF PaciFic Coast RIBEs.

The following key, made by the author for convenience
in looking up the described species of Pacific Coast Azbes,
is appended, as it may be useful to others under the same

‘

circumstances.
Chrysobotrya SPACH. GOLDEN CURRANT.

Calyx with long tube; flowers yellow, berry smooth, leaves convolute in
bud; stems without thorns or prickles.

1. &. tenuiflorum LINDL., Bot. Reg., Vol. XV, Tab. 1274. Oregon
and California.

2. KR. aureum PursH, FI. Am. Sept., Vol. I, p. 163. Oregon and
Washington.

Ribesta BERLANDIER. WILD CURRANT.

Calyx-tube cylindrical to rotate; berries smooth; leaves plicate in bud;
stems without thorns or prickles; flowers in racemes.

Calyx-tube cylindrical.

3. &. bracteosum Dovctr., Hook. Fl. Bor. Am., Vol. I, p. 233.
Northern California to Alaska.

4. RR. cereum DovuGu., Bot. Reg., Vol. XV, Tab. 1263. California
to British Columbia.

5. LR. viscosisstimum Pursu, FI. Am. Sept., Vol. I, p. 163. Sierra
Nevada of California to British Columbia.

6. KR. sanguineum PursH, Fl. Am. Sept., Vol. I, p. 164. Northern
California to British Columbia.

7. R. scuphami, sp. nov. Del Norte County, California.

8. &. brandegei, sp. nov. Lower California.

9. . glutinosum BENTH., Trans. Hort. Soc., Ser. II, Vol. I, p. 476.
Coast Mountains of California.

10. AR. malvaceum Smitu, Rees. Cycl., Vol. XXX. Coast Mountains
of California.

11. R. palimert Vasey & RosE, Proc. U. S. Nat. Mus., Vol. XI,
p. 529. Lower California.

12. R. indecorum, sp.nov. San Diego County, California.

13. RR. nevadense KELLOGG, Proc. Cal. Acad. Sci., Vol. I, 1855, p. 65

= (R. sanguineum variegatum Wats., Bot. King’s Exped.
1ooth Par.) Sierra Nevada of Central California.

14. &. ascendens, sp. nov. Sierra Nevada of Central California.

15. &. Aittellianum, sp. nov. Trinity County, California.

16. A. glaucescens, sp. nov. Mount Shasta, California.

17. R. viburnifolium Gray, Proc. Am. Acad., Vol. XVII, p. 202.
Lower California, and islands off the coast of Santa Barbara,
California.

250

18.

19.

20.

21.

22.

23.

CALIFORNIA ACADEMY OF SCIENCES. [PrRoc. 3D SER.

Calyx rotate or saucer-shaped.

R. prostratum WV HErRIT, Strip. Nov. 3, Tab. 2. British Columbia
and northward.

R. hudsonianum RicHARDs., Frank. Journ. 2nd Ed., App., p. 6.
British Columbia and northward.

R. migratorium SuxsporF, Deutsche Bot. Monats., Bd. XVIII,
p- 86. Washington.

R. laxiflorum Pursu, Fl. Am. Sept., Vol. II, p. 731. Wash-
ington and northward.

R. ciliosum Howe 1, FI. N. W. Am.,-p. 208. Mount Hood,
Oregon.

R. erythrocarpum COviILLE & LEIBERG, Proc. Biol. Soc. Wash.,
Vol. X, p. 132. Crater Lake, Oregon.

Grossulavia A. RICHARD. GOOSEBERRY.

Stems thorny under the fascicles, generally prickly besides; leaves plicate
in bud; flower solitary, in corymbs, or racemes.

24.

Berry prickly or glandular.

R. menziesit PursH, FI. Am. Sept., Vol. II, p. 732 (= R. ferox
Smitu, Rees. Cycl., Vol. XXX. [Index Kewensis]). Coast
Mountains of California.

R. hystrix, sp.nov. Santa Lucia Mountains, California.

R. californicum H. & A., Bot. Beech. Voy., p. 346. Coast
Mountains of California.

R. occidentale H. & A., Bot. Beech. Voy., p. 346. Coast Moun-
tains of California. A doubtful species.

R. oligacanthum, sp. nov. Santa Lucia Mountains, California.

R. victoris GREENE, Pittonia, Vol. I, p. 224. Coast Mountains
of California.

R. mariposanum CONGDON, Erythea, Vol. VII, p. 183. Mariposa
County, California.

R. subvestitum H. & A., Bot. Beech. Voy., p. 346. Coast Moun-
tains of California.

R. sericeum, sp. nov. Santa Lucia Mountains, California.

R. lobbit Gray, Am. Nat., Vol. X, p. 274. Northern California
to Vancouver.

R. marshallii GREENE, Pittonia, Vol. I, p. 31. Northern Cali-
fornia to Oregon.

R. amictum GREENE, Pittonia, Vol. I, p. 69. Sierra Nevada of
California.

R. wilsonianum GREENE, Erythea, Vol. III, p. 70. Mountains of
Kern County, California.

R. cruentum GREENE, Pittonia, Vol. IV, p. 35. Coast Mountains
of California to Oregon.

R. aridum GREENE, Pittonia, Vol. IV, p. 35. Sierra Nevada in

Kern County, California.

Bot.—Vot. II.] EASTWOOD—PACIFIC COAST RIBES. 251

39. R. hesperium McCtrarcuir, Erythea, Vol. II, p. 79. Los Angeles
County, California.

40. R. amarum McCvatcuis, Erythea, Vol. II, p. 79. Los Angeles
County, California.

41. R. montigenum McC atcuis, Erythea, Vol. V, p. 38.

42. R. lacustre Porr, Encycl. Suppl. Il, p. 856 (= &. echinatum
Dovuct., Bot. Reg., sub. Tab. 1349 [Index Kewensis] ).
California, northward.

43. RR. lacustre var. molle Gray, Bot. Cal., Vol. I, p. 206. Sierra
Nevada Mountains, northward.

Berry smooth.

44. R. divaricatum Dovuct., Trans. Hort. Soc., Vol. VII, p. 515
(= R. villosum Nutt., T. & G. Fl. N. Am., Vol. I, p. 547).
California.

45. R. oxyacanthoides L., Sp. Pl., p. 201 (= R. saxosum Hook.,
Fl. Bor. Am., Vol. I, p. 231 [Index Kewensis]). Sierra
Nevada of California to British Columbia.

46. FR. gracile Micux., Fl. Am. Bor., Vol. I, p. 111. Oregon to
British Columbia.

47. R. niveum LINDL., Bot. Reg., Vol. XX, Tab. 1692. Washington.

48. &. leptanthum Gray, Mem. Am. Acad., N. Ser., Vol. IV, p. 53.
Sierra Nevada of California.

49. . cognatum GREENE, Pittonia, Vol. III, p. 115. Oregon.

50. A. lasianthum GREENE, Pittonia, Vol. III, p. 22. Sierra Nevada
of California.

51. &. velutinum GREENE, Bull. Cal. Acad. Sci., Vol. I, No. 3, p. 83.
California to northern Oregon.

52. R. quercetorum GREENE, Bull. Cal. Acad. Sci., Vol. I, No. 3,
p. 83. Coast Mountains of California. This is the same as
R. leptanthum brachyanthum Gray, according to Greene.

53. Rk. ambiguum Wats., Proc. Am. Acad., Vol. XVIII, p. 193.

Washington. This has been changed to 2. watsonianum
KOEHNE.

54. R. montanum Howett, Fl. N. W. Am., p. 210. Siskiyou
Mountains, Oregon. This has been changed to &. dinomt-
natum HELLER.

Robsonia BERLANDIER.

Stems thorny; parts of the flower commonly four; calyx with erect lobes.

55. . speciosum Pursu, Fl. Am. Sept., Vol. II, p. 731 (= R. stamt-
neum SMITH, Rees. Cycl., Vol. XXX = R. fuchsioides BERL.,
Mem. Soc. Geneva, Vol. III, Pt. 2, p. 58.) Monterey, Cali-
fornia, southward.

The references not being available, the following species could not be
placed :—
56. R. roezli REGEL, Gartenfl., p. 226. Am. Bor. occ. [Index
Kewensis].
57. RR. spethianum KOEHNE.

252 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

EXPLANATION OF PLATE XXIII.

Fig. 1. Ribes brandeget, sp. nov.
a. Diagrammatic representation of the flower; X 5.
&. Leaf, natural size.

Fig. 2. Ribes scuphami, sp. nov.
a. Diagrammatic representation of the flower; X 5.
6. Leaf, natural size.

Fig. 3. Ribes indecorum, sp. nov.
a. Diagrammatic representation of the flower; X 5.
6. Leaf, natural size.

Fig. 4. Rzibes ascendens, sp. nov.
a. Diagrammatic representation of the flower; X 5.
6. Leaf, natural size.

Fig. 5. Ribes ascendens jaspere, sp. et var. nov.

Pao. GALACAn, Stt.3” See Bor VoL,

TWOOD, DEL.

ALAS

‘dl ee

254

Fig.

Fig.

Io.

CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

EXPLANATION OF PLATE XXIV.

Ribes hittellianum, sp. nov.

a.

Diagrammatic representation of the flower; X 5.

6. Leaf, natural size.
Ribes glaucescens, sp. nov.

a.
b.

Diagrammatic representation of the flower; x 5.
Leaf, natural size.

Ribes oligacanthum, sp. nov.

a,

b.

Diagrammatic representation of the flower; X 5; dotted lines
indicate the shape of the ovary and calyx-tube.
Leaf, natural size.

Ribes sericeum, sp. nov.

Flower, natural size.

Buds, natural size.

Leaf, natural size.

Petal, spread out; X 2.

Petal as it appears in flower; X 2.
Stamen; X 2.

Ribes hystrix, sp. nov.

a.

5 GUS

Flower and bud, natural size.
Leaf, natural size.

Stamen; X 2.

Petal; X 2.

——_ a

A.EFASTWOOD, DEL. PHOTO -T9TH. BRITTON & REY, SF

Fig.6. RIBES HITTELLIANIIM, SE NOV. FIG.4. RIBES OLIGACANTHUM, SP NOV.
Fig.7. RIBES GLAUCESCENS.SE Nov. FIG 4. RIBES SERICEIUM, SP NOV.

Fig.10.RIRES HYSTRIX, SP NOV.

CELE STUDIES?

I. SPINDLE FORMATION IN AGAVE.

BY W. J. V. OSTERHOUT,

Assistant Professor of Botany in the University of California.

CONTENTS.

PLATES XXV-XXVIII.

I. INTRODUCTION AND TECHNIQUE.
II. SprinDLE FORMATION IN AGAVE.

First Mitosis: PAGE.
PL ONSEN ICDC 1 PAGE) 5:5 bie: oht.a are brand aot Sete ase ac eee Bee ate eet 261
PECEEIN BEC CS TAGGIE 18, 50) ts vents, e, shay eters oneha sie si eisai es ae eae Mae 262
PSHE EE EA WS ANG ES 6, ro bs tae) © Cina © eieheuer sete eae he Pe a et Ne 265
MEIVAS CICUEAR ODAGE ots sonia od stiesclacidslont ation eile were eae Lhd tcle tiers s cee 266
EE SIPOUARMSDA GID hss bir Ws 2S. a Ain oh cane aia SAE incite 266

Second Mitosis:
5, (LES WnY0s 8) 0678 10). | Repeat aa Eee OE rere acl US ce ee Se ee a ee 267
PE EROGENINDT Gg DA GBs stata, Ache al cit eras at att bar are etd renee ett se 268
Bo. CHES TBAB CAST ONC OPE arte nner A a wes ie ne pe eer a 268
Pe SOM EB DASee STAGE ore eo ti 3 ha) ests hvac. 2 aerate IES ete Mae a ore 269
SP A SCICUIEARY SAGE 92% ocala vias ses oe Sele oR ana ee OAs Heese eads 270
CMD IPOUAR MS DA GH pay poee ar yoy, Set cere cucicieroln sei eter ene Waclon oeren 271
PENPRAL AC ONSIEN RATIONS 41 ss ss 5s tee eas Gauls Lem eiee ae tees be aoe 271
SSPE RUC DBAS SSE eC U Pe eras tS ofall oe te CUA aie Pee Cie a Sis oes 274
LLIN EDINA UT con Co ol Oe pee i ge AU OS ee ao ea 276
EARP ONG OBE LGA TES 215/0;0) Seis Vaya eye hese abet ea lesone oe enene eet oe 278

I. INTRODUCTION AND TECHNIQUE.

UnpbER this title I propose to publish the results of a
series of investigations bearing on various cell questions.
The especial attention which has been devoted to technique
in these investigations makes it desirable to refer at the

1 Contributions from the Botanical Laboratories of the University of California.

[ 255] April 30, 1902.

256 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

outset to the methods employed. The particular point of
technique which demands the most care in dealing with plant
cells is fixation. The artifact caused by many fixing agents
cannot be too carefully guarded against; it was deemed ad-
visable therefore to test each fixative by observing its effect
on living cells. This was accomplished by teasing out the liv-
ing cells on the slide in the liquid expressed from surround-
ing tissues, and observing them while irrigation with the
fixative was going on. By employing a Zeiss 2 mm. apoch-
romat with comp. Oc. 6, 8 and 12, the effect of the fixative
on the finer details of structure could be carefully followed.
Some of the most highly recommended fixatives were seen
to produce profound disturbances in the structure of the
cytoplasm and were therefore modified in various ways in
hope of better results. The fact that in different stages of
development the cell reacts differently necessitated a sepa-
rate test for each stage. The principal fixatives used are
as follows :—
1. Flemming’s strong chromic-osmic-acetic.
2. Flemming’s strong chromic-osmic-acetic diluted with
from one to twenty parts of water.
3. Flemming’s strong chromic-osmic-acetic plus from
one-tenth to one part of glacial acetic.
4. Flemming’s strong chromic-osmic-acetic modified as
follows :—
Chromic: 7: per cent. \ 1s:
Osmic. > 2 pet cent.;2 ) “cox:
Glacial acetic OL5 1. 1C:
5. Flemming’s strong chromic-osmic-acetic with the
acetic omitted.
6. Hermann’s platinic chloride-acetic-osmic.
7. The same diluted with from one to twenty parts of
water.
8. Iridium chloride 1 per cent.
g. Iridium chloride plus 1 per cent. of glacial acetic.
10. Iridium chloride .5 per cent.
t1. Iridium chloride .5 per cent. plus I per cent. of
glacial acetic.

BotT.—VOL. II.] OSTERHOUT—AGAVE, 257

r2\.
pe

14.
TS)

16.
I7.

18.

19.
20.

Palladium chloride 1 per cent.

Palladium chloride 1 per cent. plus 1 per cent. of
glacial acetic.

Osmium chloride 1 per cent.

Osmium chloride 1 per cent. plus 1 per cent. of
glacial acetic.

Platinum chloride I per cent.

Platinum chloride 1 per cent. plus 1 per cent. of
glacial acetic.

Chromic 1 per cent.

Glacial acetic I per cent.

Potassium bichromate C. P. 3 gm.
Glacial acetic BGs C.
Distilled water Q5.e4 0:

Iodine, saturated solution in distilled water.

Potassium iodide 0.5 gm.

Iodine Py (om

Distilled water 1000 c. Cc.

Boveri’s picro-acetic.

Wilson’s sublimate-acetic.

Sublimate, saturated solution in alcohol.

Rawitz’s picro-nitric.

Guignard’s iron chloride-chromic-acetic.

Guignard’s iron chloride-picric.

Merkel’s platinic chloride-chromic.

Lindsay Johnson’s_ bichromate-platinic chloride-
osmic-acetic.

Von Rath’s sublimate-picric-osmic.

Tellyesniczky’s bichromate-acetic.

Zenker’s bichromate-sodium — sulphate-sublimate-
acetic.

Kleinenberg’s picro-sulphuric.

Carnoy’s alcohol-chloroform-acetic.

Mann’s sublimate-tannin-picric.

Mann’s alcohol-sublimate-tannin-picric.

Rabl’s sublimate-platinic chloride.

Keiser’s sublimate-acetic.

Mayer’s saltpetre-picric.

258 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

Flemming’s strong mixture (1) proved decidedly the
best in most cases; very fair results were obtained also
with iridium chloride (8), platinum chloride (16), palla-
dium chloride (11),and Flemming’s strong mixture with an
excess of chromic (4). Keiser’s sublimate-acetic (39)
gave good results in some Cases.

Some objects could not be satisfactorily fixed in any of
these. In such cases good results were usually obtained
by the use of Flemming’s strong mixture with the acetic
omitted (5); where this failed resort was had to fixing in
potassium iodide-iodine (22) and transferring after a longer
or shorter time to Flemming’s strong mixture.

In most cases fixatives were found which produced no
visible change in the structure of the living cell so far as
could be ascertained by careful observation with high
powers during the application of the fixative. It was
found, however, that material placed in some of these fixa-
tives undergoes structural alterations after a few hours; in
these cases the time of fixation was shortened accordingly.

During the formation of the spindle the cell is more
sensitive (i. e., more apt to shrink or produce artifact)
to the action of fixatives than at any other time; it is
not quite so sensitive after the spindle is fully formed;
during the anaphase it grows less and less so. Pollen-
mother-cells are usually quite sensitive just after division
into tetrads. Resting stages of the cell are the least sen-
sitive of all, and the degree of sensitiveness seems to be
in inverse ratio to the amount of protoplasm in the cell.
The nucleus is always far more resistant than the cytoplasm.

At every step in the processes subsequent to fixing, the
material was subjected to careful examination in order to
guard against artifact. These observations showed that
the points requiring most care are the washing out of the
fixative and the infiltration with oil. Either of these pro-
cesses if too prolonged will frequently affect the structure
of the cytoplasm; they should therefore be carefully
controlled.

The treatment of the material after fixing was as follows:

Bot.—VOL. II.] OSTERHOUT—AGAVE, 259

After being washed from two to eight hours in running
water it was transferred to a dehydrator (for description of
this see Lawson, 1898, Williams, 1899, or Osterhout, 1900).
Dehydration in no case produced shrinkage when allowed
to proceed with sufficient slowness. The material remained
in the dehydrator until the strength of the alcohol in the
upper part was equal to that in the lower. The relative
strength of the alcohols may be ascertained as follows: Dip
a pipette into the alcohol in the upper part and allow it to
take up a few drops by capillary attraction; now dip the
pipette into the other alcohol, if on diffusing out of the
pipette it falls, it is weaker (on holding it up to the light the
course of the diffusion current is easily seen). The mate-
rial was transferred from the dehydrator to a mixture con-
sisting of equal parts of 95 per cent. alcohol and alcohol
from the dehydrator. After standing two hours a part of
this alcohol was removed, mixed with an equal quantity of
95 per cent. alcohol, and the material transferred to this
mixture. It then passed in succession through 95 per cent.
alcohol (2-6 hours), absolute alcohol (2-6 hours), absolute
alcohol and bergamot oil, equal parts (6-12 hours), and
bergamot oil (3-6 hours). It was then placed on the top
of the paraffin oven and allowed to warm up slowly, and
was then transferred to bergamot oil and paraftin—equal
parts—and allowed to remain on the top of the bath.
After six to twelve hours it was placed within the oven,
and after an hour transferred to paraffin 43° (12 hours),
and finally to paraffin 52°, 60° or 72°.

Sections 1-5 mw thick were cut on the Minot wheel micro-
tome, and fastened to the slide by the water-albumen
method. For preliminary examination of ribbons Eisen’s
alcohol method (Eisen, 1897) proved exceedingly useful;
the drying on the oven was omitted, thus effecting a great
saving of time without any corresponding disadvantage (the
sections were examined in xylene only, in which they do
not tend to float off as they do when examined in water,
without having been carefully dried on the oven). In view
of the fact that the kinoplasmic fibres, and especially the

260 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

genetic fibres, are more difficult to stain than other constitu-
ents of the cell, search was made for a stain which should
differentiate these structures sharply. Out of the exten-
sive series of stains tested, gentian violet proved the best,
especially when combined with safranin and orange G in
the Flemming triple stain. Benda’s iron- hematoxylin,
used in combination with Bordeaux red, congo red, or
ruthenium red, gives good results in the study of non-kino-
plasmic elements. The use of gentian violet is attended
with some difficulties; in many cases sections can be left
but a short time in the violet, and it is then found that the
absolute alcohol washes out the violet too rapidly. This
may be avoided by dissolving enough gentian violet in the
absolute alcohol to make a saturated solution. The wash-
ing out of the violet can then be done entirely by means of
clove or anilin oil (either pure or diluted with xylol), and
this in many cases is of great advantage. Dipping the
slides from two to twenty seconds in dilute potassium-
iodide-iodine solution just before transferring to absolute
alcohol was found in many cases to give a sharper differ-
entiation of all the structures stained by the violet.

The intense illumination necessary for work with high
oculars was secured by using a Welsbach light, and inter-
posing a glass globe (about six inches in diameter) between
it and the mirror. The globe, being filled with a colored
solution, acted as a light filter and condenser; by changing
the solution the same preparation could be studied with
several different kinds of light. Many details were
observed in this way which are hardly visible with white
cloud illumination.

II. SprnpLE FORMATION IN AGAVE.

First Mitosis.

The species studied is Agave americana L. ‘The material
was fixed in the field; the best results were obtained with
Flemming’s strong mixture. Flemming’s triple stain was
used almost exclusively.

Bot.—VOL. II.] OSTE RHOUT—AGA VE. 261

The following account relates to the first division of the
pollen-mother-cells. The cytoplasm of the resting cell
presents the appearance of a network with fairly regular
meshes. The fibres of the metwork are covered to a
greater or lesser extent by a deposit of granular substance
which tends to stain somewhat more deeply with orange G
than the substance of the fibres. The network appears
practically alike in all parts of the cell.

1. PROGENETIC STAGE.

During the formation of the dense chromatin thread and
the separation of the chromosomes the meshes of the cyto-
plasm become radially elongated, until they present the
appearance seen in fig. 1. The meshes immediately sur-
rounding the nucleus are somewhat smaller and more
regular than the others. Just outside these is a layer
of somewhat denser and more granular cytoplasm, while
the peripheral layer consists of greatly elongated meshes,
which produce at first glance the impression of free fibres.
The fibres are not free, however, but are constituent parts
of a network; it is interesting to observe, nevertheless,
that a large proportion of these fibres run more or less in
a straight line directly to the nucleus. The layer of cyto-
plasm, one mesh in thickness, immediately in contact with
the nuclear wall, is of quite even width (see fig. 1) and
gives rise to the genetic or spindle-forming layer; it will
therefore be called the rogenetzc /ayer, and may be defined
as that layer of the cytoplasm from which the genetic layer
immediately arises. The stage in which this layer takes on
its characteristic condition will be called the progenetic
stage, and may be defined as the stage immediately preced-
ing the genetic stage, and in which preparation for the latter
is going on in the cytoplasm.

In fig. 2 the progenetic layer may be seen undergoing a
characteristic change: at one end of the nucleus (the
upper end in the figure) it retains its original form, on the
sides it is growing narrower, its radial fibres are becoming
less distinct, and granules are making their appearance

262 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

along its periphery: these begin to stain violet with the
Flemming triple stain. At the same time a few delicate
fibres are seen along its periphery, which resemble the
granules in their manner of staining; they seem, in fact, to
be composed largely of granules of the same sort as those
just described.

2. GENETIC STAGE.

The granules in the periphery of the layer increase in
number and size and stain deeper and deeper violet, as is
seen in figs. 3 and 4. Eventually they become densely
crowded, and apparently fuse to form an unbroken line
which stains deep violet. Careful focussing shows that
this line represents sooner or later a continuous membrane
which forms a complete and unbroken investment of the
spindle during its formation, and which will therefore be
called the spindle wall. The spindle wall is very often, at
least in the early stages, not in contact with the nuclear
wall, but in many cases it coalesces at one or more places
with the nuclear wall. This may happen even in the ear-
liest stages, as is shown in fig. 2.

During the formation of the spindle wall there are grad-
ually developed, between it and the nuclear wall, radial
fibres such as are seen in fig. 4. These fibres are at first
very delicate, stain faintly orange, and contain irregular
scattered granules, which tend to stain violet. As the
formation of the spindle wall progresses, the granules
increase in number and size, become crowded, and finally
seem to fuse to produce continuous violet fibres. There-
upon the fibres lose their granular appearance, become
thicker, straighter, and stain more and more violet, as is
shown in figs. 5-8. Inasmuch as these fibres subsequently
form the spindle, they will be called the genetic fibres; the
layer containing them, i. e., the space between the nuclear
and spindle wall, will be called the genetic layer, and it is
suggested that this term be used generally to designate the
layer containing the genetic fibres, whether that layer be
bounded by a definite wall or not; the stage which is

Bot.—VOL. II.] OSTERHOUT—AGAVE. 263

characterized by the presence of the genetic fibres will be
called the genetic stage, and may be defined as lasting from
the first appearance of the genetic fibres to the time when
the cones are fully developed, and the nuclear wall has dis-
appeared. As in many cases these two processes do not
occur simultaneously, it should be understood that the ge-
netic stage is not ended until both are completed.

The genetic fibres are attached to both the nuclear wall
and the spindle wall; the point of insertion on the latter is
marked by a granular enlargement (see figs. 4 to Io).
They may be derived in part from the original radial fibres
of the cytoplasmic layer abutting on the nucleus, but to a
great extent they appear to be new formations, since they
are much more numerous than the original radial fibres.
The fact that the nucleolus disappears about this time is in
accord with Strasburger’s assumption that its substance is
used to furnish material for the kinoplasmic fibres (cf.
Strasburger, 1900, page 124, f.; 1897, page 378, /7.; 1895).

During the development of the genetic fibres, the genetic
layer gradually widens, and the radial arrangement of the
cytoplasm becomes less pronounced (figs. 3 to 7). In the
meantime, the irregular fibres lying outside the spindle wall
have undergone a change similar to the genetic fibres inside
the wall. The former will be called the exterzor fibres to
distinguish them from the genetic fibres. The granules in
the exterior fibres become crowded, and fuse just as in the
genetic fibres; the fibres become smoother, straighter,
thicker, and less granular in appearance, staining deeper
and deeper violet. At the same time, they gradually take
on a more radial arrangement, and grow longer, until they
finally present the appearance seen in fig. 7. The stages in
this process are clearly seen in figs. 2 to 7. These fibres
appear to be the result of a transformation of the fibres of
the cytoplasmic network, the process of transformation
proceeding from the spindle wall outward.

It is in connection with certain of these exterior fibres
that the cones are formed. It is usually where the more
conspicuous exterior fibres (those which extend to the

264 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

limiting membrane of the cytoplasm) are attached to the
spindle wall that the latter begins to form conical projec-
tions such as are seen in figs. 7 and 8. Many very con-
spicuous fibres seem to be in no way connected with the
formation of cones; around these fibres, however, other
fibres frequently group themselves to form fibre cones; but
this is not followed by the development of conical projec-
tions of the spindle wall.

The width of the genetic layer at this time varies from
that shown in fig. 7 to a width not exceeding that in the
early stage, shown in fig. 3.

The subsequent development of the cones is shown in
figs. 8 to 10; the larger ones often extend completely to the
limiting membrane of the cytoplasm, and the smaller ones
are usually connected with it by a conspicuous fibre.

During this period, the exterior fibres diminish in number,
lose their power to stain violet, and begin to stain slightly
orange. ‘They are apparently retransformed into cytoplas-
mic reticulum. They have usually ceased to be recognizable
(or at least the majority have) at the beginning of the
disappearance of the nuclear wall.

During the development of the cones the genetic fibres
contained in them undergo a certain amount of rearrange-
ment, tending to make them converge more or less toward
the apex of the cone. At the same time, they lengthen to
keep pace with the elongation of the cone, and the gran-
ular enlargement at the point of their insertion on the
spindle wall gradually disappears as though its substance
were being drawn into the fibre to provide material for its
elongation.

The pollen-mother-cells of Agave are very favorable
material for the study of the disintegration and disappear-
ance of the nuclear wall, and the penetration of the genetic
fibres into the nuclear cavity. The nuclear wall first becomes
granular, and just before it begins to disappear consists of
numerous small granules between which a delicate mem-
brane is visible. Thereupon the membrane presents the
appearance of being slowly dissolved, growing thinner and

Bot.—Vot. II.] OSTERHOUT—AGAVE. 265

at the same time losing its capacity for staining violet.
After it has entirely disappeared, the granules remain intact
for some time, and then disappearin turn. The dissolution
of the wall begins beneath one of the cones; the wall then
breaks down under the others, and the process slowly
extends from these points until the wall has completely
disappeared (see figs. 10 and 11).

The development of linin fibres keeps pace with that of
the genetic fibres, and at the disappearance of the nuclear
wall these fibres are quite numerous (see fig. 10).

Usually the spindle wall begins about this time to fray
out more or less into free fibres and lose its membranous
character. This disintegration of the spindle wall occurs
sometimes earlier, sometimes later, and is decidedly variable
as regards the time of its commencement.

3. STELLAR STAGE.

With the disappearance of the nuclear wall, and the com-
plete development of the cones, the genetic stage may be
considered to end. In the next stage the spindle has a star-
shaped appearance, due to the irregular position of the
cones; this stage will therefore be called the s¢ed/ar stage;
it may be defined as lasting from the end of the genetic
stage to the time when the cones are separated into two
groups and the fibres begin to take on a parallel arrange-
ment. From this time on the spindles resemble fasces, and
this stage will accordingly be called the fascécular stage; it
may be considered to end with the fusion of the cones to
form the bipolar spindle, or, in cases where this does not
occur, with the arrival of the daughter chromosomes at the
poles.

In Agave the beginning of the stellar stage is marked (in
the pollen-mother-cells) by the penetration of the genetic
fibres into the nuclear cavity (figs. 10 and 11), where they
soon become mingled with the linin fibres. 'The number of
poles is usually, at this time, from three to eight.

The chromosomes retain their original arrangement for
some time after the disappearance of the nuclear wall (fig. 12),

266 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3p SER.

but gradually become displaced. The fibres increase in
number and soon begin to attach themselves here and
there to the chromosomes, at first singly, and later in bun-
dles; the latter condition is shown in fig. 13. The fibres
now begin to take on a parallel arrangement, and the cones
separate into two groups marking the beginning of the
fascicular stage. It may be remarked, in passing, that the
condition shown in fig. 12, where two of the cones seem to
dominate the rest, and the fibres stretched between them
tend to be parallel, is but temporary.

4. FascicuLAR STAGE.

The fascicular stage is characterized by the increase in
number of the cones, as is clearly seen in fig. 15. This is
the result of a rearrangement of the fibres, whereby each
cone becomes broken up into several. In the meantime
the arrangement of the fibres becomes more parallel, and
the chromosomes collect in the region of the equator (figs.
14 and 15); at the same time the mantle fibres begin to
develop.

5. Breotar STAGE.

The transition to the bipolar stage is easy to follow in the
preparations; the cones gradually fuse until the stage shown
in fig. 16 is reached. This, as will be seen in the figure, is
not strictly bipolar, but the ends of the spindle are drawn
out into two or more long points which often extend to the
limiting membrane. In many spindles the fusion is com-
plete at one or both of the poles,so as to forma sharp point;
but a large proportion of them remains even during the
anaphase, in the condition shown in fig. 16.

During the stellar and fascicular stages the cytoplasm
loses almost all trace of its radial arrangement; there is a
layer of less dense cytoplasm around the spindle (figs.
10-16).

The completed spindle presents in surface view the
appearance shown in fig. 16; there is little or no trace
of connecting fibres; in median optical section, however,

aS

Bot.—Vot. II.] OSTERHOUT—AGAVE. 264

these may be seen, and in cross sections they appear as a
sort of central core of the spindle (fig. 30). After the
chromosomes leave the equator this central core of con-
necting fibres stands out prominently.

As will be seen in fig. 30, these fibres are connected with
each other by small delicate fibres, so as to form a sort of
network. It will be remembered that Belajeff (1894)
described the spindle of Zarzx as a network, but his descrip-
tion refers to its appearance in longitudinal section. The
appearance of the spindle of Agave in longitudinal section
would not justify his description.

Second Mitosts.

1. INTRODUCTION.

The cell-plate formation following the first mitosis pre-
sents very interesting features which will be fully described
in another paper. The migration of the chromosomes to
the poles reveals the presence of numerous connecting
fibres forming a sort of central core of the spindle. These
fibres remain for some time closely compacted and then
begin to spread apart until they finally extend completely
across the cell. In the meantime the cell-plate has com-
menced to form, and as it nears completion, the connecting
fibres begin to disappear. Those in the center are the first
to go, leaving a peripheral shell of fibres of great regularity
and distinctness, which persists for some time. Finally, this
in turn disappears, leaving a completed cell-plate extending
from wall to wall.

Coincident with the disappearance of the fibres is the
reappearance of the cytoplasm, which begins in the center
and extends toward the periphery of the space previously
occupied by the spindle fibres. After the completion of
the cell-plate it increases rapidly in thickness, especially
where it joins the cell-wall, until the condition shown in
fig. 17 is reached.

The daughter nucleus has in the meantime grown to its
full size, and presents the appearance shown in fig. 17.

268 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

2. PROGENETIC STAGE.

The region occupied by the spindle remains distinctly
marked for some time owing to the fact that the cytoplasm
contained within it is less granular and less dense than the
rest (figs. 17-22). Throughout the entire cell the cyto-
plasm shows a marked tendency to a radial arrangement,
being denser in the neighborhood of the nucleus (fig. 17).

The first indication of spindle formation is the appear-
ance of granules (which stain violet) in the layer of
cytoplasm immediately adjoining the nucleus. They are
imbedded in the strands or fibres of the cytoplasmic net-
work and are first seen close to the nuclear wall, from
which they gradually extend outward into the cytoplasm.
_ They are at first irregularly scattered, but soon become
more numerous and more closely crowded together until
finally they appear to fuse together to form continuous
fibres which are at first granular and irregular in outline,
but gradually become regular and smooth in appearance
(fig. 18).

3. GENETIC STAGE.

The fibres so formed are attached to the nuclear wall
and their formation always begins at the wall and proceeds
outward towards the periphery of the cell. The fibres at
first are wavy and irregular and follow the outlines of the
cytoplasmic meshes, but gradually straighten out and come
to lie in a radial direction (fig. 19). Since these fibres go
to form the spindle, they will be called the genetic fibres.
The genetic fibres gradually increase (figs. 20 and 21) in
number and size—some being much larger and more prom-
inent than others—until the stage shown in fig. 22 (upper
half) is reached. They then begin to assemble in groups,
as shown in the lower half of the same figure, in which
various stages of the process are shown. So far as can be
judged from their appearance, the groups are formed simply
by a change in position of the fibres, whereby their free
ends gradually come together at certain places and finally
fuse to form cones. The location of the cones is variable;

Bor.—Vot. IT.] OSTERHOUT—AGAVE, 269

the number is usually not less than four or more than ten.
Not all the fibres are used in the formation of cones, since
some remain, like the exterior fibres of the first division,
radiating out into the cytoplasm after the cones are fully
formed (fig. 23).

The disappearance of the nuclear wall begins, as is
usually the case, under one of the cones, and gradually
continues under first one and then another of the remaining
cones until complete. The genetic fibres penetrate rapidly
into the interior of the nucleus and become attached to the
linin network, which increases rapidly during cone forma-
tion (fig. 23, lower half).

The details of the process just described are extremely
variable. Sometimes the formation of the genetic fibres is
greatly delayed, and scarcely anything can be seen of such
fibres until just before the nuclear wall breaks down; then,
however, they appear to be formed very rapidly. In such
cases the cones do not usually reach the size shown in
He. 23..

During the genetic stage the nucleolus disappears com-
pletely while the linin increases in amount (fig. 23).

From the figures it will be seen that while in general the
process of spindle formation goes on simultaneously in
both daughter cells, this does not quite hold true of such
stages as are shown in figs. 22 and 23. This is probably
to be accounted for by the fact that these stages are passed
through with extreme rapidity. These remarks apply par-
ticularly to the assemblage of the fibres into cones and the
breaking down of the nuclear wall. The stage shown in
fig. 23, upper half, is not passed through quite so rapidly,
and it is more common to find both daughter cells in this
stage at the same time.

4. STELLAR STAGE.

The stellar stage is variable in length and presents con-
siderable diversity as regards the form of the spindle and
the number of cones composing it. Frequently the cones

270 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

are prominent and well defined, and may reach nearly to
the limiting membrane (figs. 24 to 26), while in other cases
they are merely small, ill-defined elevations. Free fibres,
which radiate out into the cytoplasm, persist for a time, but
gradually disappear. The cones are generally unequal,
some having larger, longer, more numerous and more
prominent fibres than others.

The chromosomes preserve for a time their original
arrangement (fig. 25), but gradually become displaced and
irregularly scattered (figs. 25 and 26). The fibres in the
meantime combine to form more or less definite strands
which are at first irregular and loosely aggregated, but
which gradually become more compact and regular in
appearance. Gradually these strands or groups of fibres
become attached to the chromosomes: the attachment is at
first partial, involving only a few of the fibres of the strand,
but gradually extends to the other fibres of the group
(fig. 26).

5. FAscicuLAR STAGE.

While the attachment of the fibrous strands to the chro-
mosomes is taking place the arrangement of the strands
themselves begins to undergo a change whereby they tend
to place themselves parallel to one another (fig. 26); at
the same time, certain cones lying at the ends of the in-
cipient spindle become increasingly prominent, and seem
to serve as places of assemblage for the other cones which
appear to gather around them, thus finally forming two
distinct groups which lie at the opposite ends of the fascicle
(fig. 28). Inthe meantime, the strands of fibres become
more and more compact, and form well defined, thick fibres
(fig. 27).

The cones in each group gradually approach each other
more closely, and at the same time the number of cones
increases (fig. 28); the fusion of the cones is gradual, and
frequently takes place sooner at one end of the spindle
than at the other (fig. 28). The assemblage of chromo-
somes in the equatorial plate does not take place until the
fusion is well advanced.

_—

Bot.—Vot. II.] OSTERHOUT—AGAVE. 271
6. BrIrPoLAR STAGE.

Strictly speaking, there is not, as a rule, a bipolar stage,
since the fusion does not become complete but remains in
the stage shown in fig. 29. It is not uncommon to find the
fusion complete at one end but not at the other, and in
some cases the spindle may become truly bipolar.

GENERAL CONSIDERATIONS.

The spindle formation of the first mitosis of Agave, as
set forth above, differs markedly from anything hitherto
described. The formation of a spindle wall which com-
pletely encloses the spindle during the genetic stage is
unique; and the origin and development of the genetic
fibres is peculiar.

The spindle wall would seem to be a structure of great
physiological importance, comparable in its functions with
the nuclear wall and the limiting layer of the cytoplasm.
Its special function appears to be to segregate the important
process of spindle formation, thus carrying the division of
labor in the cell to a point otherwise impossible. Its pres-
ence therefore indicates a high degree of cell organization.

In appearance and mode of origin the spindle wall resem-
bles the nuclear wall; its close relation to the nucleus is
shown by its origin and development; it persists, however,
after the nuclear wall has disappeared, and may then per-
haps assume some of its functions. It is either in actual
contact with the nuclear wall during the genetic stage or is
constantly connected with it by the genetic fibres; on the
other hand, it early forms connections with the limiting
membrane by means of the exterior fibres. The relations
existing between the kinoplasm and the limiting membrane
have recently been emphasized by Strasburger (1900, p.
144 e¢ seg.), who calls attention to the fact that in many
forms the kinoplasmic fibres can be traced to this membrane
and that the spindle is in many cases inserted on it. Agave
offers direct confirmation of these statements.

It is possible that there may be some homology between

(2) May 2, 1902.

272 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

the spindle wall and the structures, more or less membran-
ous, which form at varying distances from the nuclear wall
in the pollen-mother-cells of Zz/zwm (Mottier, 1897), Cobea
(Lawson, 1898), and Lavatera (Byxbee, 1900). These
structures consist at first of a rather loose weft of fibres
(which stain violet with the Flemming stain); later the
fibrous weft becomes condensed and compressed so as to
form in many cases a thick continuous membrane which
stains violet; other cases may usually be found in the same
anther, in which the membrane is not continuous or frays
out in places into a fibrous weft. These structures stand in
no such intimate relation with the nucleus and the spindle
as the spindle wall of Agave, and seem in comparison with
it irregular and ill defined. That they may have some
phylogenetic connection with the spindle wall is not improb-
able; if so, they are probably to be regarded as rudimentary
structures.’

On the other hand, it seems quite certain that many of
the important resemblances between animal and plant cells
are to be regarded not as homologous but as analogous, and
the same is probably true of resemblances in cell structure
between different groups of plants. We must therefore
exercise considerable caution in assuming homologies
between cell structures.

The development of the genetic fibres is peculiar in that
there is no weft stage such as is characteristic of most
forms of higher plants hitherto studied; on the contrary, the
fibres are radial from the beginning. This may very likely
be connected with the presence of the spindle wall. Numer-
ous observations incline me to believe that the weft stage is
caused by the expansion of the nucleus, which at first
causes a tangential elongation of the meshes immediately
surrounding the nucleus; as the expansion continues the
meshes are broken and form free fibres. It is easy to see
how the turgidity of the genetic layer (contained within the

1 An interesting case is that described by Eisen (1900) for Batrachoseps, where a ‘‘false
nuclear wall” is formed around the daughter chromosomes: within this the true nuclear
wall is formed.

Bor.—Vot. II.] OSTERHOUT—AGAVE. 273

spindle wall) could hinder or altogether prevent the expan-
sion of the nucleus and so eliminate the weft stage. The
osmotic pressure of the genetic layer, if equal to or slightly
greater than that of the nucleus, would bring about this
result.

The exterior fibres do not seem to be essentially different
from the genetic fibres save that, being outside the spindle
wall, most of them take no direct part in the formation of
the spindle. Both kinds of fibres seem to be similar to the
radial kinoplasmic fibres of Aguzsetum. Some of the exte-
rior fibres form fibre cones in much the same way as in
Equisetum, but these structures are, in the case of the first
mitosis of Agave, merely temporary.

The formation of the spindle cones is peculiar on account
of the part played by the exterior fibres. According to
Belajeff (1894), the spindle cones of Zarzx are formed in
connection with certain fibres which extend from the gene-
tic layer to the cell wall. In this respect the process
resembles that which occurs in the first mitosis of Agave,
but the fibres do not group themselves to form the spindle
cones in the manner described by Belajeff.

While the method of spindle formation in the first mitosis
stands unique, that of the second mitosis recalls the origin
of the spindle in the spore-mother-cells of Aguzsetum
(Osterhout, 1897), with which it agrees in most of its prin-
cipal features, ¢. @., in the origin of the genetic fibres in
contact with the nuclear wall, their radial arrangement and
outward growth into the cytoplasm, their assemblage in.
groups accompanied by a fusion of their free ends to form
cones, and, finally, the fusion of these cones to form the
spindle. In Hguzsetum, however, there is at first a modi-
fied weft stage of which no trace is seen in Agave. This
agreement with Hguzsetwm is the more surprising since the
relationship between the two plants is so remote, while the
spindle formation of the immediately preceding mitosis of
A gave presents such profound differences.

Investigations on spindle formation, though as yet but
few in number, have revealed great diversity, especially in

274 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

plants, where almost every careful investigation has dis-
covered a new type. Widely divergent types have been
shown to occur in different tissues in the same individual
(Nemec, 1898; Strasburger, 1900, p. 119, also, p. 112, 7.)
or even in the same tissue (Davis, 1901; Hertwig, 1898).
In view of this we may conclude that the process of spindle
formation is a plastic one, easily modified in the course of
phylogeny and even of ontogeny, and that its study deserves
to be seriously taken up as a matter of vital importance to
an understanding of cell mechanics.

SUMMARY.

The first spindle formation of the pollen-mother-cells of
Agave americana L. presents the following features of
especial interest.

1. During the early stages of the spindle it is enclosed
in a special membrane (of cytoplasmic origin) which forms
a complete investment around it, and is called the spzndle
wall; the functions of this membrane appear to be compa-
rable to those of the nuclear wall and the limiting membrane
of the cytoplasm; it appears to be a unique structure.

2. Unlike most cases hitherto described, no weft stage is
present; this is probably due to the presence of the spindle
wall, which prevents the expansion of the nucleus, and con-
sequently the formation of the weft.

3. The genetic (7. e., spindle-forming) fibres are radial
from the beginning, and are attached to both the nuclear
and the spindle wall.

The second spindle formation differs zz toto from the
first and may be summarized as follows: The genetic
(spindle forming) fibres form in close contact with the
nuclear wall, take on a radial arrangement, extend outward
into the cytoplasm, assemble in groups, and form cones
which by their fusion give rise to the spindle. The process
resembles in general the spindle formation of the spore-
mother-cells of Aguzsetum.

Bot.—VOL. II.} OSTERHOUT—AGAVE, 275

It is proposed that the stages of spindle formation be
characterized as
(1) Progenetic
(2) Genetic
(3) Stellar
(4) Fascicular
(5) Bipolar.

The progenetic stage may be defined as the stage preced-
ing the genetic, and in which preparation for the genetic
stage goes on in the cytoplasm; the region of the cytoplasm
in which this takes place and from which the genetic layer
arises may be called the progenetzc layer.

The genetzc stage lasts from the first appearance of the
genetic fibres (2. e., the spindle-forming fibres) to the com-
plete development of the spindle cones and the disappear-
ance of the nuclear wall. When these processes do not
take place simultaneously, it is to be understood that the
genetic stage does not end until both are completed. The
layer containing the genetic fibres is called the genetzc layer.

The ste//ar stage lasts from the end of the genetic stage
to the separation of the cones into two (more or less oppo-
site) groups, and the commencement of the parallel
arrangement of the fibres.

The fascicular stage lasts from the end of the stellar
stage to the complete fusion of the cones (to form the
bipolar spindle), or, where this does not occur, to the
arrival of the daughter chromosomes at the poles.

The dzfolar stage lasts from the end of the fascicular
stage to the complete disappearance of the poles.

276

1894.

1900.

IgoOI.

1897.

1900.

1898.

1898.

1900.

1897.

1898.

1897.

1900.
1895.
1897.
1900.

1899.

CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

LITERATURE CITED.

BELAJEFF, W. Zur Kenntniss der Karyokinese bei den Pflanzen.
Flora, Bd. LXXIX, p. 430.

ByxBEE, E. S. The Development of the Karyokinetic Spindle in
Lavatera. Proc. Cal. Acad. Sct., 3d Ser. (Bot.), Vol. II, p. 63.

Davis, B. M. Nuclear Studies on Pellia. Azmnals of Botany, Vol.
XNVE p. 147.

Eisen, G. Notes on Fixation, Stains, the Alcohol Method, etc.
Zeitschr. f. wiss. Mikr., Bd. XIV, p. 195.

The Spermatogenesis of Batrachoseps. /ourn. Morph., Vol.
SQA roe, Le

Hertwic, R. Ueber Kerntheilung, Richtungskérperbildung und
Befruchtung von Actinosphzrium Eichornii. Adh. d.k. bayr. Akad.
ad: wess; Vi Cl. Bd. XTX, 'p: 636.

Lawson, A. A. Some Observations on the Development of the
Karyokinetic Spindle in the Pollen-Mother-Cells of Cobzea scan-
dens Cav. Proc. Cal. Acad. Sci., 3d Ser. (Bot.), Vol. I, p. 169.

Origin of the Cones of the Multipolar Spindle in Gladiolus.
Bot. Gazette, Vol. XXX, p. 145.

Mortier, D. M. Beitrage zur Kenntniss der Kerntheilung in den
Pollenmutterzellen einiger Dicotylen und Monocotylen. /ahré. /.
wiss. Bot., Bd. XXX, p. 169.

Nemec, B. Ueber die Ausbildung der achromatischen Kerntheilungs-
figur, etc. Bot. Centralblatt, Bd. LXXIV, p. t.

OsTERHOUT, W. J. V. Ueber Entstehung der karyokinetischen Spin-
del bei Equisetum. /ahrb. f. wiss. Bot., Bd. XXX, p. 159.

Befruchtung bei Batrachospermum. Flora, Bd. LXXXVII,
p. 109.

STRASBURGER, E. Karyokinetische Probleme. /ahrd. /. wiss. Bot.,
Bd. XXVIII, p. 151.

Ueber Cytoplasmastructuren. Kern- und Zelltheilung. /ahré.

f. wiss. Bot., Bd. XXX, p. 375.

Ueber Reductionstheilung, Spindelbildung, Centrosomen und
Cilienbildner im Pflanzenreich. Jena.

Wituiams, C. L. The Origin of the Karyokinetic Spindle in Passiflora
coerulea Linn. Proc. Cal. Acad. Sct., 3d Ser. (Bot.), Vol. 1, p. 189.

SS eee

248 CALIFORNIA ACADEMY OF SCIENCES. | [PRoc. 3D SER,

EXPLANATION OF PLATE XXV.

The material was fixed in the field with Flemming’s strong mixture; sec-
tions were cut 1-5 uw thick and stained with Flemming’s triple stain. Figures
were drawn with the aid of the Abbé camera lucida, Zeiss’ Apoc. Hom. Im.
Obj. 2.00 mm., Comp. Oc. 8. All pertain to the division of the pollen-
mother-cells of Agave americana L.

The lithographic plates reproduce the details of colour and form of the
preparations with great fidelity; the differentiation of the kinoplasmic
elements is in no way exaggerated.

Fig. 1. Pollen-mother-cell before the beginning of spindle formation. The
cytoplasm shows the characteristic radial arrangement, the inner
portion being denser.

Fig. 2. Progenetic layer in process of transition to the genetic (spindle
forming) stage. It shows violet granules along its periphery, and
outside this a few delicate fibres which later develop into the
exterior fibres. ;

Fig. 3. The periphery of the layer more violet; exterior fibres more
developed.

Fig. 4. Periphery of the layer shows violet granules more closely crowded
together; radial genetic fibres appear within the layer; exterior
fibres still more developed; beginning of genetic stage.

Fig. 5. Spindle wall definitely formed; genetic and exterior fibres still more
developed.

Fig. 6. Genetic layer widening; exterior fibres begin to form fibre cones.

Beginning of cone formation.

Further development of the spindle cones.

=
Gay
on

"TT =

[OsrerHour] PLATE XXV.

CaL AAD Sc1.32 Ser. Bor. VaLIl

RU

280

CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

EXPLANATION OF PLATE XXVI.

Fig. g. The spindle cones completely developed; the exterior fibres dis-

appearing.

Figs. toand 11. Disappearance of the nuclear wall; beginning of stellar

Fig. 12.

Fig.

Fig.

cA

oo

ER:

16.

stage.

The nuclear wall has completely disappeared, but the chromosomes
retain their original arrangement. Two of the cones seem to
dominate the rest, but this is only a temporary arrangement.

Beginning of fascicular stage; the fibres are assuming a parallel
arrangement; the cones are separated into two groups; fibres are
becoming attached to chromosomes in bundles.

The parallel arrangement of the fibres and their attachment to
chromosomes is more pronounced.

The cones have increased in number; the chromosomes are assem-
bled in the equatorial region.

Completed spindle; not strictly bipolar, since each end is drawn out
into two or more points.

OsTerRHouT] Phare YX

[

132 Ser Bor. Vou Il.

SCI

ACAD

Al
ALS

a Ay ire
- =i

282 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

EXPLANATION OF PLATE XXVII.

Fig. 17. Progenetic stage of second mitosis. The cytoplasm shows a radial
arrangement; in the region previously occupied by the spindle
fibres of the first mitosis the cytoplasm is less dense. Granules
which stain violet begin to make their appearance in strands
of cytoplasm nearest the nucleus.

Fig. 18. Genetic stage. The violet granules gradually increase in number,
become crowded, and fuse to form the genetic fibres.

Fig. 19. The genetic fibres increase in number and size.

Figs. 20 and 21. The genetic fibres increase and assume a more regular
arrangement. The lower half of the figure shows in both cases a
slightly more advanced stage than the upper.

Fig. 22. The upper half shows the genetic fibres fully formed; in the lower
half they are assembling in groups to form the spindle cones;
various stages in this process are shown.

Fig. 23. The upper half shows the cones fully formed: the lower half shows
the breaking down of the nuclear wall and the penetration of the
genetic fibres into the nucleus, marking the commencement of
the stellar stage.

Fig. 24. The nuclear wall has completely disappeared; the chromosomes
preserve their original arrangement.

i Kenn
cy a :

a a i - :

284

Fig.

Fig.

Fig.
Fig.
Fig.

Fig.

25.

26.

27.

28.

29-

30.

CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

EXPLANATION OF PLATE XXVIII.

In the upper half of the figure the chromosomes ‘still preserve
their original arrangement; in the lower half they have already
become somewhat disarranged. The genetic fibres tend to col-
lect into strands which have not yet attached themselves to the
chromosomes. 2

The fibrous strands now show a more or less complete attach-
ment to the chromosomes and a tendency to assume a parallel
arrangement.

The strands have become compact fibres and are more nearly
parallel.

Transition to bipolar stage. In the lower half of the figure the
chromosomes are assembling in the equatorial region.

Completed spindle; not strictly bipolar, since the fusion of cones is
not complete.

Cross-section of spindle of first mitosis near the equatorial region,
showing the central core of connecting fibres interconnected by
delicate threads.

[OsteRHout] PLATE XNVIII

Ft od
= Uh SSE at BY
at Ht at ay

=

NEW SPECIES FROM THE SIERRA NEVADA
MOUNTAINS OF CALIFORNIA.

BY ALICE EASTWOOD,

Curator of the Department of Botany.

Most of the plants described below were collected on a
trip to the south fork of Kings River from Millwood to the
Kings-Kern Divide (Harrison’s Pass), and to Kearsarge
Pass. The trail lies near the boundary of Fresno and
Tulare counties, and is one of the best known of the south-
ern Sierra Nevada.

It is a country of magnificent forests, beautiful mountain
meadows, rocky slopes down which dashing torrents rush,
and cafions through which the river flows serenely but
swiftly. The upper elevations are characterized by jagged
peaks and ridges which are clothed with everlasting snow
and enormous granite boulders, and are gemmed with little
lakes of great beauty.

The collection was made between July 2nd and 12th,
1899. To the care and assistance of Messrs. Pierson Dur-
brow, S. L. Berry, and Benjamin Brooks, members of the
party, the success of the trip is due.

The types are in the Herbarium of the California Acad-
emy of Sciences.

1. Streptanthus gracilis, sp. nov.

Annual or biennial from a slender tap-root, with delicate branches, chiefly
from the base, 1-2 cm. high, glabrous, glaucous. Basal leaves orbicular to
narrowly elliptical or spatulate, sinuate-dentate to obtusely lobed or lyrate,
tapering to long slender petioles as long as or twice as long as the blades,
together 1-3 cm. long; cauline leaves linear-oblong to ovate, entire to crenately
lobed, 5 mm. to 2 cm. long, auriculate at base, either sessile or on very short
petioles. Racemes few-flowered, those from the slender basal branches one-
to six-flowered; pedicels erect, 1-5 cm. long, generally shorter than the calyx;
bracts wanting except with the lowest flowers, which are in the axils of the

[ 285 ] June 2, 1902.

286 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

upper leaves. Calyx rose-purple, urceolate, narrowed under the spreading
lobes; the lobes obtuse, membranously margined, and undulate, becoming as
long as the tube formed by the united sepals; disk-like receptacle conspicu-
ous. Petals rose-pink, the slender claws exserted, blades spatulate to
obovate-orbicular, half as long as the claws (3 mm.), and more than twice
as broad (2 mm.), sinuate. Stamens in three pairs, with filaments all distinct,
one pair with filaments 1 mm. long, the next pair 2 mm., and the third
5 mm.; the anthers of the two shorter pairs 3 mm. long, those of the longest
pair not quite 2 mm. Pods (immature) erect, linear, glabrous, tipped with a
broad, sessile stigma; the largest 1.5 cm. long, 1 mm. wide. Seeds not known.

This delicate little plant was collected by the author
below timber line on the trail from East Lake to Harri-
son’s Pass (the type locality), and by Miss Catherine E.
Wilson, in 1898, on the trail to Bull-frog Lake. It seems
to be rare.

2. Polygonum exile, sp. nov.

Annual, with slender, wiry stems, erect, four-ribbed, about 3 dm. high,
glabrous or minutely puberulent, especially near the top, with few erect,
virgate branches from near the base. Leaves erect, linear-lanceolate, or
terete from the inrolling of the margins, jointed to the ochrez, cuspidate,
the upper surface minutely dotted; lowest leaves 2 cm. long, diminishing
upwards, shorter than the internodes; ochrez with three hyaline, long-acumi-
nate divisions. Flowers solitary or few at the nodes, erect, sessile, or on
short pedicels; divisions of the perianth extending to below the middle;
white, pinnately veined with green or rose-colored veins, elliptical, hooded at
apex, 2.5 mm. long, investing the ripe seed. Stamens three, with ovate,
long-acuminate filaments half as long as the perianth; anthers minute.
Akene three-angled, rhombic in outline, acute at each end, 2.5 mm. long,
brown, glossy, minutely papillate; styles very short, deflexed.

This is similar in general appearance to P. douglasiz
Greene, but differs in having but three stamens and in the
erect flowers.

Collected by the author in Kings River Caijion, July 4,

1899.
3. Eriogonum scapigerum, sp. nov.

Caudex branched, the divisions clothed with the dead brown bases of
former leaves. Leaves all radical, oblong to orbicular, 5-15 mm. long,
densely white-matted-tomentose on the lower surface, becoming somewhat
glabrate on the upper, with undulate margins, obtuse apex, and cuneate
base, tapering abruptly to the long petiole; petiole 3-5 cm. in length, flat,
with a central rib, and broadening at base for about half the length. Scapes
many, very slender, glabrous, 5-17 cm. high, terminated by a solitary head

Bot.—VOL. II.] EASTWOOD—NEW PLANTS FROM CALIFORNIA. 287

not more than 1 cm. in diameter. Bracts united, ternate, the lobes deltoid-
acuminate, dark red, glabrous, except for the long, white-woolly hairs on the
margin; bractlets at base of involucres similar but smaller, with the lobes
more deeply divided on one side. Involucres turbinate, glabrous, indistinctly
ribbed, woolly-ciliate along the entire to undulate margins; pedicels ex-
serted, distinctly jointed to the perianth at apex; divisions orbicular to
obovate, hooded at apex, glabrous on the outside, hairy within at the base ;
the outer ones broader, 2 mm. long. Stamens slightly exserted, anthers
two-lobed, suborbicular. Akenes three-sided ; styles spirally coiled.

This is near &. nudum Douglas, of which it may be
only an alpine variety. It looks quite different from the
common form as found in the Coast Mountains.

Collected by the author on Harrison’s Pass, above tim-
ber line, at an elevation of almost 14,000 feet, July 9, 1899.

4. Garrya pallida, sp. nov.

Branching shrub, several feet in height; older stems glabrous, dark brown;
younger ones cinereous, with densely appressed, silky pubescence. Leaves
oval, elliptical, ovate, or obovate, becoming stiff and thick with age, pale
green and glaucous, strongly veined, cinereous, with an appressed pubes-
cence of fine silky hairs on both surfaces but denser on the lower, and
becoming sparser with age; tapering somewhat at each end, the apex ab-
ruptly acuminate, with the point recurved, margin entire, thickened, rarely
slightly undulate; blade 3-7 cm. long, 2-4 cm. wide; petiole stout, .5-1.5 cm.
long. Pistillate spikes pendent, solitary or clustered, 4-6 cm. long; lowest
bracts deeply cleft, long-acuminate, upper ones cleft above the middle,
‘abruptly acuminate, silvery, silky canescent from the densely appressed
hairs; ovaries ovate, on short, thick pedicels, with pubescence similar to
that of the bracts; styles divaricate; berries becoming almost glabrous. The
staminate flowers have not been seen.

This species is nearest to G. fremont? Torrey, and is
found in the southern Sierra Nevada. The type was col-
lected in Kings River Cafion, July, 1899. Specimens from
San Emidio Cafion, Tejon Pass, and Tehachapi, all in Kern
County, also from the region of the Kaweah River, in Tulare
County, seem to be the same species.

5. Convolvulus berryi, sp. nov.

Stems perennial, trailing but scarcely twining, 5 dm. or more long, with
few branches from near the root, densely white-woolly throughout, with
fine, spreading hairs. Lower leaves on petioles longer than the blades;
upper shorter, 1-4 cm. long, broadly deltoid, with spreading sinus or
Sagittate with the sinus less open, apex mucronate, the two basal angles

288 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

lobed with mucronate or obtuse lobes, 2-6 cm. wide at base, 2-5 cm. long.
Flowers chiefly near the base, on long peduncles surpassing the leaves, often
upwardly curving, 5-8 cm. long; bracts similar to the upper leaves, close
under the flower, 1 cm. wide, 1.5 cm. long. Sepals ovate-oblong, unequal
in breadth but of the same length, hairy with appressed hairs except on the
membranous margins of the inner ones, mucronate, 14 mm. long, the broad-
est Io mm. wide. Corolla yellowish white, hairy on the angles and at their
tips, 4cm. long. Stamens with narrow sagittate anthers, 7 mm. long ; fila-
ments hairy below, attached to the corolla for half their length. Style as
long as the stamens (2.5 cm.), with ovate-lanceolate stigmas, 3 mm. long,
I mm. wide.

This beautiful species comes nearest to C. vz/losus Gray,
from which it differs in the bracts, peduncles, and stamens,
also in the broader leaves and longer peduncles. The
pubescence is less velvety. Itis also near C. tomentellus
Greene, from which it differs in almost the same organs.

Collected at Millwood by the author, July, 1893 and
1899, also near Converse Basin. The plant is named in
honor of Mr. S. L. Berry.

6. Castilleia brooksii, sp. nov.

Perennial, branching from the base, but with a few short branches above,
3 dm. high, glandular, viscid throughout and with a pubescence of uneven,
silvery, weak, jointed hairs. Leaves sessile, linear-oblong, entire to three-
lobed, about 2 cm. long, less than 5 mm. wide; lobed leaves generally
subtending the branches, the divided portion half the length. Branchlets
terminated by short, compact spikes, with subsessile flowers; bracts variously
and unevenly lobed, with the tips colored. Calyx a little longer than the
corolla tube, obliquely gibbous at base, equally cleft before and behind, with
divisions shorter than the tube, each two-cleft, with unequal, triangular-sub-
ulate, one-nerved, obtuse divisions, 3 mm. long. Corolla 2.5 cm. long; galea
longer than the tube, straight at first but later curving outwards, having three
blunt teeth at apex, the middle one smallest; lower lip truncate, 3 mm. long,
2 mm. wide, the sharply acute teeth incurved, folds noticeable. Stamens
exserted, with filaments smooth and anthers narrowly linear, with unequal
cells. Ovary glabrous, obliquely acuminate; stigma clavate, exserted from
the top of the galea.

This comes under the group to which belong C.. parviflora
Bongard, and C. mznzata Douglas, with neither of which it
agrees. The flowers are yellowish red but probably vari-
able in color, as in most species of this genus.

Collected by the author on the trail up Bubbs Creek,
early in July, 1899, and named in honor of Mr. Benjamin
Brooks.

es

——

Bot.—VoL. II.] EASTWOOD—NEW PLANTS FROM CALIFORNIA. 289

7. Castilleia disticha, sp. nov.

Perennial, erect, 6 dm. high, branching from the base and also above with
generally short, slender, spreading branches; somewhat viscid and with a
close, often somewhat scanty, cinereous pubescence mixed with longer,
jointed hairs. Leaves linear, acute or obtuse, the lower 4 cm. long, 3 mm.
wide, diminishing upwards, sessile by a truncate or subauriculate base,
distinctly three-veined, the middle vein most conspicuous; margin entire or
undulate-crisped. Inflorescence spicate, elongating in fruit, especially on
the main stem, 1-2 dm. long; flowers after anthesis distichous, becoming
more or less remote, sessile or almost so, with the capsule appressed to the
stem, the calyx and corolla persisting and spreading; bracts foliaceous, the
upper ones, only, colored, variously toothed, with the middle tooth longest,
equalling or shorter than the corolla. Calyx slightly surpassing the corolla
tube, about equally cleft before and behind for half the length; each division
tipped with red, two-cleft, with triangular, subulate, unequal lobes, 2-3 mm.
long, three-nerved, thin in texture, somewhat gibbous, but not broadest at
base. Corolla red, 3 cm. long, with galea as long as the tube, truncate or
emarginate at apex; lower lip three-toothed, the middle tooth much smaller
than the lateral, separated by a broad sinus, thin, not callous. Stamens
exserted, filaments glabrous; anthers narrow, with unequal cells, more than
2mm. long. Stigma exserted from the summit of the galea, capitate. Cap-
sule obliquely oblong-ovate, 8-ro mm. long, chartaceous; seeds elliptical to
orbicular, light brown, invested with a membranous, foveolate outer coat.

This is more closely allied to C. minor Gray and C. sten-
antha Gray than to any of the perennial species. Its ses-
sile or almost sessile flowers, more brightly colored and
differently shaped, together with the different habit of
growth, mark it as distinct.

The type was collected by the author at Converse Basin,
on the trail to the south fork of Kings River, July, 1899.
The species is also found at Millwood, where it was col-
lected the same year by the author, and by Mr. T. S. Bran-
degee, July 19, 1892.

8. Castilleia nana, sp. nov.

Low, 3-6 cm. high, with several stems from a woody caudex, which is
thickly clothed with the dead stems of former seasons ; somewhat cinereous
and viscid, the pubescence of the inflorescence of longer, jointed, arachnoid
hairs. Leaves simple and linear or three- to five-divided, with narrow, linear,
acuminate lobes which are shorter or longer than the undivided portion,
together 1-2 cm. long. Flowers in heads terminating the stems; bracts
broad, similar to the upper leaves, embracing the sessile flowers and more
than twice as long; the division terete from the involute margins, once and a
half to twice as long as the lower, undivided part, which is 4 mm. wide

290 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

and 5 mm. long. Calyx glandular, subequally cleft before and behind, with
the divisions twice as long as the tube, surpassing the corolla, each with two
narrow, linear lobes 5-6 mm. long, extending below the throat of the in-
conspicuous corolla. Corolla 7 mm. long, the lower lip longer than the
galea, the three lobes thin, ovate-orbicular, obtuse, 2 mm. wide, somewhat
saccate below; galea with the middle portion of thicker texture than the
membranous sides, three-toothed at apex. Stamens with the upper pair of
anthers exserted, the lower included, each distinctly two-celled, clothed
with a few long hairs at base. Pistil capitate, .5 mm. in diameter, exserted;
ovary lanceolate. Fruit and seeds unknown as the plants were too young.

This is an alpine species found only above timber line.
It is related to the group formerly included under Ortho-
carpus, section Castilleioides, but differs from all other
species in the peculiar corolla, it being the only species
described with the galea shorter than the lower lip.

Collected by the author on Harrison’s Pass, above East
Lake, July 9, 1899.

There are either one or two other low, alpine species in
the same region which seem to be related to C. pallida
Kunth, but the material is too scanty for satisfactory deter-
mination.

g. Mimulus bioletti, sp. nov.

Annual, branching diffusely, chiefly from the base, about 2 dm. high,
glandular-villous throughout, except the corolla. Leaves near the root
spatulate; cauline leaves rhombic obovate to lanceolate, thin, tapering to a
broad petiole, or sessile, sparingly serrulate or entire, 2-4 cm. long, 2-15 mm.
broad. Flowers axillary, on slender, upward-spreading peduncles, almost as
long as the internodes, generally shorter than the subtending leaves. Calyx
tubular, 8 mm. long when in flower, 12 mm. in fruit, often purplish-dotted
below the middle, plicately carinate-angled, the ribs rugose, rounded; divi-
sions deltoid, with the margins involute, the obtusely pointed apex spreading
outwardly. Corolla as long again as the calyx, with ampliate throat and
scarcely two-lipped border, crimson, the upper lip with a yellow blotch dotted
with crimson in the throat; limb 12-15 mm. across, with divisions rounded,
crenulate, or entire. Stamens and style included; the former four in two
sets, each united by the anthers, one set longer than the others; anthers
ciliate, explanate, one above the other; stigma bilammelar, cuneate in out-
line. Capsule included in the rigid calyx-tube, obtusely four-ribbed, opening
at the sides from the base up; placenta free, except at the top; seeds numer-
ous, minute.

This belongs to section Eumimulus Gray, and is most
closely related to 1. palmeri Gray. It differs chiefly in
the larger flowers and different calyx.

Eee

= _-

EEE

Ee

BoT.—VOL. II.] EASTWOOD—NEW PLANTS FROM CALIFORNIA. 291

Collected in Hetch-Hetchy Valley, Tuolumne County,
by Mr. F. T. Bioletti, in July, 1900.

10. Phacelia stimulans, sp. nov.

Stems tall, simple from a branched caudex, becoming 5-6 dm. high, erect,
sparsely leaved, generally flowering from the middle, viscid-pubescent, and
clothed besides with fine, long, stinging hairs. Radical leaves forming a
rosulate cluster, simple or with a few lobes at base, ribbed between the
hispid veins, elliptical, acuminate, 3.cm. long; petioles very hispid with
spreading hairs. Spikes of the panicle simple, the lowest and uppermost
geminate, horizontally spreading, somewhat distant, 5-6 cm. long; pedun-
cles very glandular, becoming shorter near the top; pedicels capillary, half
as long as the calyx. Divisions of the calyx oblong-spatulate, hispid, net-
veined, shorter than the corolla, surpassing the capsule. Corolla tubular,
the lobes conniving after anthesis and persistent, held to the calyx by the
tangling together of the long, persistent stamens and style. Filaments
exserted, conspicuously clothed with long white wool. Capsule ovate-
acuminate, hispid; seeds ovate, brown, not glossy, pitted.

This is allied to P. cercinata Jacq. f., but is entirely
unlike any of the described species which were formerly
included under that species. On account of the stinging
hairs of the stems and leaves it might be confused with
P. nemoralis Greene; but this has an altogether different
habit, pubescence, and inflorescence.

Collected by the author, July, 1899, in Kings River
Cafion, not far from the swampy meadow near which
campers stop on the way to Bubbs Creek.

11. Gilia sparsiflora, sp. nov.

Annual, a foot or so high, branching above, with slender, spreading stems,
minutely glandular-pubescent. Leaves few, terete from the infolding of the
margins, about an inch long, tipped witha short bristle. Flowers few, termi-
nating the branchlets, two to three in the clusters, sometimes solitary in the
upper axils; bracts keeled at base, three-lobed, the middle lobe much larger
than the lateral, all subulate-aristate, surpassing the flowers. Calyx mem-
branous between the ribs, clothed with dense, white, cottony wool, the un-
equal, aristate-subulate divisions as long as the corolla tube. Corolla salver-
form, 1 cm. long, white with some purple dots in the funnel-form throat; the
divisions elliptical-obtuse, half as long as the tube. Stamens equally in-
serted, with arrow-shaped anthers, obtuse at apex, exserted from the throat
of the corolla. Capsule oblong, 1 cm. long; seeds few, oblique at base, three-
sided, generally with rounded angles, developing mucilage and spiracles.

292 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

Gilia sparsifiora belongs to the same group as G. vzr-
gata Steud.

Collected by the author in Kings River Caijion, in July,
1899; and also along Bubbs Creek trail.

12. Cryptanthe vitrea,’ sp. nov.

Annual, with several stems from a tap-root, 1-2 dm. high, very hispid
throughout with white bristly hairs which are pustulate at base. Leaves
linear, 1-2 cm. long, 2 mm. wide, strongly nerved. Flowers sessile, in
numerous short spikes from almost the lowest axils, on slender peduncles.
Calyx 3 mm. long, the sepals conniving to form a tube around the nut-
lets, the tips free, densely clothed with long stiff bristles, 3 mm. long,
and also a fine, white, hispid pubescence. Corolla only about 2 mm. long,
with a smalllimb. Nutlets, only two maturing, ovate, obtuse, almost 2 mm.
long and 1 mm. wide at base, very glossy brown mottled, sharply tuberculate,
attached to the gynobase for the entire length, the groove closed except at
the forks.

The species comes nearest to C. muriculata Greene, but
the nutlets are broader, the flowers much smaller, and the
entire plant so floriferous that in the dried specimen it is
almost impossible to distinguish the peduncles.

Collected by the author on Bubbs Creek trail, July 5,

1899.
13. Aster durbrowi, sp. nov.

Stems erect, perennial from creeping root-stocks, disposed to grow in
tufts, 1-3 dm. high, sparingly leafy, erect, branching only at the inflores-
cence, glabrous and green except for some white woolly hairs which are
sparse on the lower part, but which make the upper part almost cinereous.
Radical leaves on long winged petioles which are dilated and clasping at
base, lanceolate, the blades about as long as the petiole, together 5-8 cm.
long, 5-10 cm. wide, entire or distantly serrate, ciliate on the margin, the
hairs becoming longer towards the base of the petiole and decurrent on
the stem; cauline leaves similar but sessile by an auriculate or cordate clasp-
ing base, the upper ones broadening and becoming shorter and more
pointed, the lower ones narrowing towards the insertion. Heads cymose,
the branchlets terminated by one to four middle-sized heads, 1-2 cm. across,
on short bracteate pedicels; involucre of six rows of imbricated bracts with
green, foliaceous tips, not spreading, the inner ones with purplish acuminate
tips, the outer linear, mucronate, glabrous except for the ciliate margins,
distinctly one-nerved and chartaceous at base; rays pistillate, reddish purple,
1 cm. long, I.5 mm. wide, dentate at apex, sparingly ciliate on the lower
part; disk flowers purplish; corolla shorter than the pappus, which extends

a

a

il

BoT.—Vor. II.] EASTWOOD—NEW PLANTS FROM CALIFORNIA. 293

about to the exserted style and stamens; the tube slightly pubescent.
Akenes (immature) hispid with dense, white, upwardly appressed hairs;
pappus simple, scabrous.

This is near A. yosemztanus Greene (A. ascendens yosem-
vtanus Gray), of which it may prove to be a variety. The
habit ef growth is quite unlike that of the above species,
while its larger heads, sparingly leafy stems terminated by
few heads, and the auriculate- or cordate-clasping cauline
leaves make it appear even more distinct. It is the com-
mon aster of the wet meadows at the upper altitudes in
this region.

Collected in Horse Corral Meadow, July 11, 1899, and
named in honor of Mr. Pierson Durbrow.

14. Madia villosa, sp. nov.

Stems simple, erect from an annual root, about 3 dm. high, slender, vil-
lous with long, white, soft, spreading hairs, also becoming glandular near the
top with black, stipitate glands. Lower leaves generally opposite, upper
alternate, linear or the lowest oblanceolate, sessile, entire, or glandular-
serrate with distant teeth, revolute, strongly ribbed, villous, with the hairs
finely pustulate on the older leaves, obtuse at apex, 4-7 cm. long, 2-5 mm.
wide. Heads few, terminating slender peduncles near the top of the stem,
the uppermost first in bloom, clothed with few small bractlets; outer bracts
of the involucre 8 mm. long, the foliaceous tips equalling that which encloses
the akene, linear acuminate, villous and glandular, half as long as the deeply
three-lobed rays, these often with a reddish-brown spot at base; inner bracts
scarious with short foliaceous tips; disk flowers all sterile, the corollas as long
as the abortive akenes, together 8 mm. long, the tube somewhat villous and
the lobes clothed at tip with spreading hairs; fertile akenes flattened laterally,
semilunate, black and brown mottled, minutely papillate in rows, 1.5 mm.
wide, 4 mm. long, glabrous; anthers purple and exserted, giving a purplish
color to the disk.

This is nearest to M/adia corymbosa DC. (Aladaria cor-
ymbosa Greene). It is also close to M7. hispida Greene,
and really seems to be intermediate between the two
species.

The type was collected at Converse Basin, July 12, 1899.
It was also found on Bubbs Creek trail.

THE ROOT-TUBERCLES OF BUR CLOVER (MED-
ICAGO DENTICULATA WILLD.) AND OF
SOME OTHER LEGUMINOUS PLANTS.

BY GEORGE JAMES PEIRCE,

Associate Professor of Plant Phystology in the Leland Stanford Junior University.

CONTENTS.

PLATE XXIX.
PAGE
EN TRODUCTION, AND. NETHOD e1).trtse ease eeieid sss He wieleve 6) + oe sé 295
II. ORIGIN AND MORPHOLOGY OF ROOT-TUBERCLES...........-.--. 298
III. THE Form AND DISTRIBUTION OF ROOT-TUBERCLES............ 312
Nie HE STRUCTURE OF, ROGT-TUBERCLES:. 055506 bs cies see os oss ole ee 316
SSRINGMMAUR Const te ccs ety | totes ve Sater ana tepcteree es Cara eeaT Oe Ie ME Reve tchey ich cits 324
PB UBEVOG ADEE. 2 Sicr2\'s)slei cts eae tiatetesajuie teneie ave ieleteteteions, ote aie mmpoteniare oie vena B27)
EXPLANATION OR! PUATE 0 )2 cicsija vain rtaa tse a lea a tieteli oa ea tiers 6 328

I. INTRODUCTION AND METHOD.

SOME time ago, on casually examining some hand-sections
of the root-tubercles of Bur Clover (Medicago denticulata
Willd.), I was struck by the great differences between the
cells containing bacteria or bacteroids and those in which
there were none. In these sections, the bacteria-containing
cells looked so unhealthy, as compared with the cells free
from bacteria, as plainly to suggest that the intimate rela-
tions of bacteria and leguminous cells were not mutually
advantageous, but that the bacteria were parasitic. There-
upon I began a careful microscopic study of the root-
tubercles of Bur Clover and other leguminous plants! in

1 Bur Clover was especially favorable for my work because, at most seasons, I could
get living plants from out of doors very near the laboratory, and I could grow such ma-
terial as I needed at other times from seed very quickly in the laboratory. Besides this,
however, I have studied Lupinus micranthus Doug. var. bicolor Watson, L. rivularis Doug].
var. latzfolius, Melilotus parviflora Desf., Medicago sativa Linn., Hosackia subpinnata Torr. and
Gray. The points which I wished to determine are essentially the same in all of these,
and hence my descriptions, though specifically of Bur Clover, are applicable to the
others.

[ 295 ] June 17, 1902.

296 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

order, if possible, to ascertain the real relations of the
tubercle organism to the cells in which it is found. This I
believe I have done. The bacteria are parasites, not bene-
fiting but injuring, if not finally killing, the cells in which
they occur. Whether the association of these bacteria with
a leguminous plant benefits the plant as a whole is another
question, answerable not from microscopic examination but
solely by experimental cultures. According to Frank (1890,
p- 109), the Legumznose are not all similarly affected by the
bacteria. Some are greatly benefited, stimulated to in-
creased growth and other activities, while in others the
bacteria are ordinary parasites, not benefiting the host in
any way in return for the food derived from it. Without
implying whether the results of infection benefit or injure
the plant as a whole, one may speak of the roots or cells of
leguminous plants being zzfected by tubercle bacteria. I
shall later, however, take occasion to discuss whether the
presence of the tubercle organism is really beneficial.

The material studied was either fresh, growing out of
doors wild or sown in boxes in the laboratory, or alcoholic.
The latter was fixed in Flemming’s chrom-osmic-acetic
mixture, dilute, and after washing for twelve to twenty-four
hours in running water was dehydrated and kept in go per
cent. alcohol. These tubercles, which were of different sizes,
ages and conditions, according to the season, were imbed-
ded in paraffin melting at 54° C., sectioned and mounted in
the usualway. The youngest tubercles scarcely turn brown
in the fixing fluid, but older ones become brown or almost
black. In any case I transferred the slides, after the paraf-
fin had been removed by turpentine, tora solution contain-
ing one part commercial peroxide of hydrogen in twenty
parts 80 per cent. alcohol. In this solution they remained a
half hour, or until the sections were no longer in the least
brown, and were then run down into water for staining.

The method of staining which I found most useful is a
combination of Flemming’s well known and now very popu-
lar triple stain—anilin safranin, anilin gentian violet, and
orange G.—with Ehrlick’s method of staining cover-glass

CC

Bot.—VOL. II.| PEITRCE—ROOT-TUBERCLES. 297

preparations of bacteria. The stains were made up
according to the directions given in Humphrey’s Zimmer-
mann’s Botanical Microtechnique (p. 186), and were used
as Hof (1898) directs, except that after the sections had
been for not more than two minutes in the gentian violet
solution they were rinsed with water and placed for a
half hour or longer in Gramm’s iodine solution to differ-
entiate the bacilli and the infection threads from the cyto-
plasm. Hof says that the sections may be left from two to
three minutes in the anilin gentian violet. I often found
this quite too long, and had difficulty in washing out enough
of the violet without taking out the safranin also. One
minute is usually long enough for these tissues. Washing
off the Gramm/’s iodine with water, the slides were then
allowed to remain for one to two minutes in staining bottles
containing orange G; they were then washed with absolute
alcohol so long as gentian violet came off abundantly or
needed to be removed (as shown by microscopic examina-
tion), were cleared in clove oil, and mounted in xylol
balsam. I decidedly prefer clove oil to xylol for clearing,
as it aids in the differentiation for which this staining
method is so highly prized.

So far as my experience goes, this method of fixing and
staining is perfectly certain to demonstrate the infection
threads and to differentiate the bacteria in the cytoplasm
and in the unstained matrix of the threads. I am, therefore,
somewhat at a loss to understand the difficulties reported
by some authors in staining tubercles and their contents.
Miss Maria Dawson (1899, p. 8) reports, for example:
‘¢For some time I made use of both hand-sections and
microtome sections of paraffin material. The latter method
I afterwards abandoned, however, since I found the tuber-
cle tissues very difficult objects to stain upon the slide, and
also ordinarily thin hand-sections serve better for the exam-
ination of the filaments within the cells—a point to which I
wished to devote special attention.’’ With the stains Miss
Dawson used on hand-sections, and of which she speaks

298 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

favorably, I, too, failed to get satisfactory results from
microtome material, butowing to the success of my own
method I did not try hers very long.

II. OricIn AND MorPHOLOGY OF ROOT-TUBERCLES.

Although most that I can say about the entrance of the
tubercle bacteria into the roots of leguminous plants has
already been reported by others, I wish to describe the
infection in Bur Clover (Medzcago denticulata Willd.) from
the beginning, and to discuss some of the stages in the
process. I must also frankly admit that I do not know all
that has been written on the subject, for the literature is
copious and scattered, and I have been able to see only the
papers herein referred to. I therefore bespeak lenient
criticism of my acquaintance with the literature, remote-
ness from the centres of scientific and other work making
it very difficult to secure the papers, and even references
to the papers, of my subject.

How the tubercle bacteria in the soil come into contact
with the root-hairs of the leguminous plants which they
attack is not known. The majority of authors consider
these bacteria (Bacillus radicicola Beyerinck, Ahizobium
leguminosarum Frank) only slowly motile if motile at
all. In artificial cultures they are usually quite motion-
less (Migula, 1900, p. 772). Zinsser (1897, p. 447) says
they are small and actively motile. Miss Dawson (1900, p.
59) reports that in drop-cultures, a week or more old, the
chains become motile, the shorter moving more rapidly than
the longer, but none actively, and the motion resembles the
pendulum movement of Oscz/latorza filaments. In younger
drop-cultures, containing 2.5 per cent. gelatine, I have seen
this movement. The movement of the chains formed in
artificial cultures may be only a feeble index of a much
more active movement of the separate bacilli when these
occur in natural conditions. Artificial cultures are unsatis-
factory at best, and it may very well be that the tubercle
bacilli are actively motile, for a time at least, in damp soil.

Bot.—Vor. II.] PEIRCE—ROOT-TUBERCLES. 299

It is well known that many bacteria are actively motile only
for a short time after division. When the conditions in the
soil—whatever these conditions may be—favor the repro-
duction of these bacilli, the young and separate rodlets may
be able to move with a fair degree of rapidity. They may
possibly be stimulated into motility by the proximity of the
root of some leguminous plant, and may be characteristically
attracted to it by the substances diffusing from it into the
soil-water (see Czapek, 1896). At first glance this seems
improbable because of the very small proportion of root-
hairs attacked to the total number of root-hairs formed;
but on this point the following observations throw some
light.

In the field of a Leitz objective III and ocular 3 I counted
one hundred root-hairs on the sides of a young lateral root.
Of these hairs one was infected. ‘There must have been an
equal number of root-hairs on the top and bottom of the
root as it lay on the slide. The zone of hairs was about
five times as long as the distance through which I counted
hairs. This would make the total number of hairs on this
small root at least one thousand. I searched carefully, but
found no other infected hair anywhere on this rootlet. The
proportion of infection in this case is as one to a thousand.
The rootlet examined was from a young Bur Clover seed-
ling growing in sandy soil in the laboratory. There must
have been a large number of Bur Clover bacteria in the
soil used, for it was taken from a spot where Bur Clover
throve last year and again this. Out of doors I fancy that
the number of root-hairs would be greater in the same sandy
soil than in the laboratory, for I watered my indoor mate-
rial, and that outside was watered only by the rains. The
number of root-hairs attacked is probably no greater, how-
ever. The proportion of one to a thousand is therefore
conservative. If, then, these bacteria are motile only
slowly, if at all, it is apparently the mere chance of a root-
hair’s growing to or very near where the bacteria are which
makes infection possible. We may infer either that the

1 This I estimated with considerable precision by means of a mechanical stage.

300 CALIFORNIA ACADEMY OF SCIENCES. [PrRoc. 3D SER.

bacteria are not motile or move only slowly, or that it is
chance which governs the infection of the root-hairs. This
latter inference is encouraged by a statement of Miss Daw-
son’s (1899, p. 21), that she saw ‘‘ on a very small piece of
a lateral root from one of the plants no less than twenty-
seven hairs, side by side, with well grown infection tubes
within them. This observation may serve to show how
successful the attacks of this organism may be, provided
suitable conditions can be arrived at.’’

To ascertain whether the number of infections in root-
hairs in nature is smaller than it might be under other
conditions, I tried the following experiment. Three layers
of filter paper, moistened with tap-water, were laid in each
of four small saucers and covered by tumblers. These
were sterilized in an Arnold steam sterilizer on three suc-
cessive days. A half dozen Bur Clover seeds were then
placed under each tumbler on the damp filter paper. The
seeds had just been removed from the little bur-like pods,
dipped in corrosive sublimate solution of one to one thou-
sand, and rinsed in distilled water which had been repeat-
edly boiled. The filter-papers were watered daily with
boiled distilled water. In a few days the seeds germinated.
When the roots had grown to an inch or so in length and
had developed many root-hairs, they were watered with
boiled distilled water in which had been ground healthy
growing tubercles. The water therefore held tubercle
bacteria in suspension. The next day nearly every hair
in the field on one side of a root was found to be enlarged
and twisted at the ends and showed the beginning of an
infection thread. Given the contact or close proximity of
the tubercle bacteria with the root-hairs, infections may
take place in great numbers simultaneously, at least when
the roots are very young. Whether the roots are always
susceptible, or whether older root-hairs or root-hairs on
older roots are susceptible, is another question. Any change
in the composition of the walls of the root-hairs may affect
their solubility or at least their permeability by the bacteria
(cutinization?).

Bot.—VOL. II.] PETRCE—ROOT-TUBERCLES. 301

It would appear from another experiment that not all con-
tacts and infections of root-hairs with tubercle bacteria lead
to the formation of tubercles. Among the seedlings grow-
ing in sterilized moist chambers, I infected some with
bacteria from a gelatine culture of Bur Clover tubercle
bacteria. The next day showed a great increase in the
number of bacteria, but the tips of the root-hairs, though
bent in many instances, were not coiled in the manner usual
in infections, but instead, were cut off into short sausage-
shaped, often non-nucleated segments. In this way the
bacteria which have entered a root-hair are excluded from
the more vital parts of the root, just as gonidial cells in
lichens are known to exclude the haustoria of the fungus
by so dividing that only one daughter-cell contains any part
of the haustorium which has penetrated the mother-cell
(Hedlund, 1892; Peirce, 1899). Once given the contact
with the bacteria, the root-hairs can become infected; but
these infections may be resisted by the leguminous plant by
cutting off the infected parts.

I am by no means ready to attach especial weight to the
result of this experiment for the following reasons: First,
I did not repeat the experiment, important though it would
be to prove that the root-hairs do cut off the infected por-
tions; second, this result followed the infection of the
sterile root of a seedling, not by bacteria suspended in
water but by stroking the root with a platinum needle which
had been dipped into a culture of the bacteria. By this
means not only bacteria but also their accumulated pro-
ducts in the culture-medium were put upon the root. It
might well be that these products, rather than the bacteria
themselves, so irritated the root-hairs that they segmented
as above described. It would be interesting to follow this
matter to a decisive conclusion, but it was not possible at
the time to do so, and this point was not directly connected
with the main object of this investigation. It may be that
by similar means the root-hairs, and thereby the roots, of
other than leguminous plants resist and escape infection
by the bacteria which so characteristically affect the

302 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

leguminous plants growing in the same soil and under the
same conditions. This possibility also deserves the test of
experiment.

Returning now to the question of the motility of the tu-
bercle bacteria, we see that the experiments just described
indicate that fewer infections take place in nature than when
‘many bacteria are brought directly into contact with young
and sterile roots, but the experiments leave the matter of
the motility of the bacteria still undecided. The behavior
of the bacteria in artificial culture is inconclusive, and ap-
parently we cannot now imitate the conditions which prevail
in the soil.

Figures 1-3 show root-hairs of Bur Clover plants infected
by tubercle bacteria. Figure 2 shows the lower and longer
of the two hairs in fig. 1 more highly magnified. Figure 3
is a hair on another plant drawn with the same magnification
as fig. 2 (300). In these figures, as in those of Frank,
Dawson, and many others, it is noticeable that on the con-
cave side of the curved tip of the root-hair there is a small
mass of bacteria, this mass being continuous with the line
or strand of bacteria extending through the hair. The
wall of the hair seems intact and uninjured except where
the small mass of bacteria is. At this point there is no ap-
parent rupture of the wall. The wall may be actually per-
forated, though to see this with the mass and the strand of
bacteria in place would be very difficult or impossible even
in very thin sections. It is much more probable that the
wall is merely softened, the cellulose digested at the point
where the bacteria are—a soft place large enough in area
to permit the bacteria to enter either by actual locomotion
or by the formation and growth of new cells in this direction.

The little group of bacteria on the surface and near the
tip of a root-hair is very often at the point of greatest curv-
ature of the hair. This curvature is due to the bacteria.
The bending is the evident response to irritation. The
irritation may consist in the softening and partial solution of
the cell-wall by enzyms formed by the bacteria—a mechan-
ical irritation—or in the stimulation of the cell by the same

Bor.—VoL. II.] PEIRCE—ROOT-TUBERCLES. 303

or other substances. In the one case we have a traumatropic
bending, in the lattera chemotropic. (See Spalding, 1894.)
This bending is entirely different in appearance and dis-
tinct in cause from that which carries the root-hairs closely
around particles of soil. This last is due to irritation by
contact (thigmotropism) and by water (hydrotropism).

Since the majority of infected root-hairs show the bend-
ing at or near the tip, as shown in figs. I-3, we may infer
that the bacteria enter uninjured hairs which are able by
growth curvatures to respond to mechanical or chemical
stimuli. If the hairs were broken, the ability to respond,
and the responses (curvatures) would be greatly lessened,
and instead of a short, close spiral (figs. 1-3), or a pro-
nounced bend, we should have little or no curvature. The
curvature of a broken hair is doubtful, and for mechanical
and physiological reasons certainly difficult to understand.

The roots of young Bur Clover grown in sandy soil in the
laboratory showed very few broken hairs when I dug up
the plants to search for infected hairs. The soil was very
friable, but even then I expected to find more hairs broken
as the result either of taking up the plants or of their growth
in the soil.

It would appear, then, that these bacteria are able to
soften or dissolve cell-wall, and when they come into con-
tact with a root-hair, enter it, whether it is broken or not.
The very slow movements, which are all that most observers
report having seen in these bacteria, their ability to soften
or dissolve cellulose, the small number of infected hairs,
and the small number of broken hairs, make Fischer’s
graphic description (1897, pp. 91-2) of how the infection
of leguminous plants takes place at least doubtful though no
less graphic and interesting. He says: ‘‘ Die feinen Wur-
zelharchen einer jungen, noch kndllchenfreien Legumino-
senpflanze schieben und drangen sich iiberall zwischen die
Bodentheilchen ein, um hier Wasser und mineralische Salze
aufzunehmen, ja sie scheiden sogar besondere Stoffe aus,
um die Erdteilchen, mit denen sie dicht verkleben, zu lésen.
So wird schon die unverletzte Oberflache der Wurzeln

304 CALIFORNIA ACADEMY OF SCIENCES. [PrRoc. 3D SER.

chemotaktisch wirkende Stoffe vielfach absondern. Dazu
kommen noch zahlreiche verletzte Wurzelhaare oder andere
leichte Wunden der Wurzel, die anlockend auf Knd6llchen-
bakterien wirken werden, wenn diese in den wasserer-
fiillten Raumchen zwischen den Bodentheilchen herum-
schwarmen. Wovon hier die Bakterien leben, bedarf noch
weiterer Untersuchung, denn sie miissten hier natiirlich mit
bescheideneren Kohlenstoff- und Stickstoff quellen vorlieb
nehmen als in der Reinkultur mit Asparagin und Zucker.
Gerade solche Stoffe, besonders das chemotaktisch sehr
wirksame Asparagin ist in den Keimpflanzen der Legumi-
nosen stets reichlich enthalten und wird bei jeder Verletzung
der Wurzel hervortreten. So konnte ihm wirklich die
Rolle des Anlockungsstoffes fur die Knéllchenbakterien
zufallen, die in ein aufgerissenes Wurzelhaar genau so
einschwarmen wiirden, wie in eine mit Asparagin gerfiillte
Kapillare.’’ Fischer’s conviction, expressed at length and
supported as well as possible by example and argument
(1. c., pp. 131-2), that bacteria do not enter uninjured
plant-cells and hence cannot produce disease by being par-
asitic on or in plants, is probably responsible for this state-
ment, which seems to me the opposite of correct. If it can
be shown that bacteria of any one species penetrate the
cell-wall of healthy uninjured plants, producing unusual
growths therein, Fischer’s contention that there are no
bacterial diseases of plants breaks down. On this question
Smith (1901) takes issue with Fischer, and seems to prove
that there are bacterial diseases of plants.

To help himself over the unavoidable difficulty of the
tubercle bacteria entering the roots of plants, Fischer says
(1. c., p. 92): ‘*Ja es scheint sogar, als ob die Legu-
minosen durch Auflockerung der Zellwande an manchen
Wurzelhaaren u. s. w. die Anlockung der bacterien vorbe-
reiten. In dicht gedrangten Ziigen dringen sie von der Ober-
flache der Wurzel in deren Inneres vor, wobei ihnen wie-
derum die Leguminose den Weg zu ebnen scheint dadurch,
dass sie die schwer durchdringbaren Zellwande etwas auf-
lockert.’? Such a loosening or softening (Auflockerung)

Bot.—VOL. II.] PEITRCE—ROOT-TUBERCLES. 305

of the cell-walls is well enough known in other cases; but
it is not the host which softens its own walls in order to
facilitate the entrance of a foreign organism, but rather the
foreign organism which, by enzyms secreted by itself,
softens or dissolves the walls of its host which lie across its
path of growth.’

Having entered the root-hair by softening or dissolving
a small portion of the cell-wall, and moving or growing
through this, the tubercle bacteria multiply rapidly, forming
a thread-like zooglcea from the infection spot along the hair
into the epidermal cell of which the hair is a branch (figs.
2and 3). From the epidermis the infecting zoog-cea grows
fairly straight into the underlying cortical parenchyma.
Figure 4, drawn from one of a series of thin microtom sec-
tions stained as previously described, indicates the course
of the infecting strand (purple). This course is nearly,
though not quite, straight toward the central cylinder of the
root, for within a series of five or six sections—a distance of
20-30 #—the infection thread was traced from the base of
the root-hair (7. 4. in fig. 4) to one cell (10) in the layer
next to the endodermis of the central cylinder. The cells
in this layer are distinguished from the cells of the cortical
parenchyma by somewhat larger and denser nuclei. This
layer is the one from which the lateral roots arise. The
direction of the infection thread—which is solid, and is
incorrectly termed infection ‘‘ tube’’—#is too regular not
to encourage one to suppose that the course of the growing
strand of bacteria is determined by attraction exerted by
the host-cells upon the bacteria. This then is chemotropic
growth of the strand or, if the bacteria are motile in the
cells, chemotactic movement of the bacteria. The course
of the thread is toward the conducting tissues of the host.
This is similar to the growth of the haustoria of Dodder,
Cassytha, Viscum, Phoradendron, and other phanerogamic
parasites (Peirce, 1894; Cannon, 1901). The growth does

1 See De Bary (‘‘Morphology and Biology of the Fungi, Mycetozoa, and Bacteria,”
Oxford, 1887) and many others as to this in fungi, and Peirce (Annals of Botany, 1894) as
to Dodder.

306 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

not extend into the central cylinder and the conducting tis-
sues, so far as I have seen. Instead, in the layer of cells
just outside the endodermis of the root, division takes place
in the cell into which the infection thread has penetrated
and in the cells adjacent to it. The daughter-cells grow,
repeated divisions and growth follow, and there arises a
conical mass of cells which are somewhat larger, and which
contain more protoplasm, than the adjacent cortical paren-
chyma cells. This conical mass is the young tubercle. At
first all of its cells are merismatic, but later the divisions
become more and more limited to the cells near the rounded
apex of the blunt cone. Thus aregular cambium is differ-
entiated in the tubercle. This cambium, as shown in fig. 7,
lies near the tip of the tubercle, and forms a bowl-shaped
or shallow thimble-shaped layer.

The growing tubercle pushes out the overlying cortical
parenchyma and epidermis, forming an increasing swelling
on the side of the root. Cortical parenchyma and epider-
mis, at least for a time, nearly keep pace with the growth
of the tubercle. Thus, although the cortical cells are com-
pressed somewhat, the epidermis is not ruptured, and the
tubercle does not burst out of the side of the root as a
lateral root does.

The layer of cells which, on infection, gives rise to the
tubercle, forms new cells not only centrifugally but also
centripetally, so that by these new and growing cells the
tubercle is pushed outward, away from the central cylinder.
In this way the cylindrical mass of the root itself is kept
fairly uniform. An older tubercle appears to be attached
to the root. Only by tracing its development can one see
that it originates internally. The course of development
was only very imperfectly traced by Schneider (1893);
hence his bold and erroneous statement that ‘‘ tubercles
seem always to develop exogenously.’’

Frank (1890, p. 70) states that the tubercles are new
organs formed and well nourished by the plant. He com-
pares them with galls which are formed by plants at the
points attacked by parasites (insects, worms, etc.), and

Bot.—VOL. II.] PEITRCE—ROOT-TUBERCLES. 307

adds, to strengthen his comparison, ‘‘ die Wurzelkndllchen
sind kein Organwelches der Leguminose urspriinglich eigen
ware, ebensowenig wie dies bei den anderen Pflanzen der
Fall ist, sondern eine erst von dem Rhizobium angeregte,
dann aber selbst aufgebaute Bildung.’’ Further on, he says
that he has repeatedly seen lupines, cultivated in sterilized
and uninfected soil, which formed swellings on the roots
closely resembling young tubercles, but showing neither
infection threads nor any traces of the cell-contents char-
acteristic of true root-tubercles. He accounts for this not
on the ground of slight infections producing only abortive
tubercles, but on the hypothesis that the lupines, accus-
tomed for thousands of years to symbiotic existence with
the tubercle bacteria, have so firmly acquired the habit of
forming tubercles that they begin to form them even before
and without infection. On these points I wish again to call
attention to the fact that the tissues of the tubercle orig-
inate from the same layer of cells as gives rise, by similar
divisions, to the lateral roots (see figs. 4, 5, 6). When one
compares a very young mass of tubercular tissue, still
enclosed in the cortex of the root, with a very young
lateral root also still enclosed in the cortex of the root, the
resemblance between the two structures is strong. Figures
5 and 6 show this. Figure 5 is a diagram of a section in
which a tubercle and a lateral root are growing side by side
and from the same layer. In the figure the tubercle is to
the left, the lateral root to the right. Figure 6 is a drawing
of tubercle and lateral root on a larger scale, the root to
the right, the tubercle to the left. In the tubercle some
infection threads show. The tubercle has the same form as
the root, but shows no differentiation among its cells. The
lateral root already shows a differentiation of dermatogen and
there is a foreshadowing of the vegetative point. Central
cylinder and periblem are not yet distinguishable. The
cells of the tubercle are larger than those of the lateral
root, but the nuclei of the tubercle cells are not proportion-
ally larger and most of them are actually no larger than
those of the lateral root.

308 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

As tubercle and lateral root grow, the resemblances be-
tween the two decrease and finally disappear altogether.
The tubercle has no cap! and no central strand of con-
ducting tissue. The tubercle cells differentiate into definite
tissues more slowly than do those of the lateral root, but
near the tip of the older tubercle there is a mass of meriste-
matic cells similar to the growing-point of a lateral root (see
fig. 7). This meristem forms cells forward and backward
as does the growing-point. The central mass of the tuber-
cle is proportionally much larger than the central cylinder
of the lateral root, but it is wholly undifferentiated. The
cortex of the tubercle contains vascular bundles, small and
separated from each other by considerable spaces of paren-
chyma (see fig. 7), and is enclosed by layers of cork-cells.
These may and usually do become powdery on the surface
and rub off as the tubercle forms, just as the cap cells do
from the tip of a root.

In point of origin and in their earliest growth, tubercle
and lateral root are similar. In subsequent growth they are
more and more dissimilar. Morphologically, then, the root-
tubercles are lateral roots. Though called into activity by
very different causes, the cells of the pericycle give rise by
division to masses of cells which, on the one hand, develop
into tubercles, on the other develop into lateral roots. In
the one case we know the stimulus which causes the tuber-
cle to form. It is the infection of the root-cells down to
the innermost layer of the periblem by bacteria. Do lateral
roots form as the result of external stimuli or are they the
effects of causes internal to the plant? The latter is the less
likely from the fact that the size, number, and position of
lateral roots varies in plants of the same species according
to the soil, to the number and kind of other plants living in
the same soil, to the distribution of moisture and other
matters in the soil, and to a great number of other factors
not now recognized. This subject merits investigation. As

1 According to Life (Botanical Gazette, April, 1901) the roots infected by Axabena and
certain other organisms in certain species of Cycas also have no caps, yet he unhesitat-
ingly describes them as roots.

BotT.—VOL. II.] PETRCE— ROOT-TUBERCLES. 309

plants are studied with a view to ascertaining the effects of
each factor in the environment, it will become more and
more evident that many of the effects which are now
attributed to internal causes, lumped together under the
name inheritance or distributed among the various func-
tions of the organs of the body, are the reactions of the
parts to stimuli exerted upon them from outside. If the
formation of lateral roots by the division of the cells of
the pericambium should prove to be the result of external
stimuli, it will be found that these stimuli operate upon
the cells immediately concerned.

The diverse development of tubercle and lateral root, the
result of the persistence of the different stimuli which called
them into existence, obscures the common morphology of
the two organs so that it is only natural that Frank should
have called the tubercles new organs. If the tubercles
were the result of hypertrophy indiscriminately among the
cells of the cortex of the root, as may be the case in several
of the species which Frank studied and described, this
would be no evidence that the tubercles are morphologically
roots. But in Bur Clover the case is clear. Doubtless the
same can be made out in other leguminous plants when
the origin of the tubercles is studied in properly stained
microtome sections of young roots. I have not examined
Frank’s Lupinus, bean, pea, etc., for it is a matter costing
much time and patience to find just the right stages, and I
preferred to study my own plants and to leave a review of
the origin and structure of the tubercles of the plants stud-
ied by Frank to a later time or to others interested.

In Bur Clover, at least, I think I have advanced strong
evidence against Frank’s contention that the tubercles are
new organs. As to his hypothesis, involving inheritance as
one of the factors in their origin and development, that
lupines grown in sterilized and uninoculated soil may form
small enlargements like young tubercles, I think my demon-
stration of the common origin of tubercles and lateral roots
also has some significance. Assuming that the soil remained
sterile, which is not wholly probable, the plant might begin

310 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

to form lateral roots which, for some unknown reason,
aborted while still in the cortex of the mother root. Lat-
eral roots are known to do this, and if the abortion took
place early enough, the root character of the new formation
might be lost if it had already developed. It seems to me
much more probable, however, that both causes were in
operation; that the soil did not remain sterile; that the
plants were infected by so few or so feeble tubercle-bacteria
that the tubercles stimulated to begin to form aborted because
the infection was not strong enough. If the leguminous
plant, or its separate cells, and the bacteria are parasitically
associated, the plant would resist the entrance and growth
of the bacteria, and would be much more likely to succeed
in this if the attacking bacteria were few or feeble. Over-
coming the bacteria, the stimulus to tubercle formation
ceases, the tubercle remains rudimentary. That infection
of sterilized soil by the tubercle-bacteria is possible, and
even difficult to avoid, is known to all who have worked on
the subject. This, then, rather than inheritance, accounts
for the rudimentary tubercles which Frank describes.

The bacteria in the infection thread, which grows through
the root-hair and the cortical parenchyma cells of the root
to the pericambium layer, multiply, but they multiply most
rapidly in the infected cells farthest from the surface of the
root. New threads form, which grow out into and infect
the cells of the mass of new cells composing the embryo-
tubercle. Thus a majority of the cells in the young tubercle
contain bacteria.

Though infected cells do divide (see pp. 322-323), they
probably divide less often than the uninfected cells. The
primary infection is in a nearly straight line from the root-
hair inwards. The infection of the daughter-cells com-
posing the embryo-tubercle is accomplished by branching
infection threads growing in fairly straight lines radiating
from the base of the tubercle. In this way the cells near
the base of the growing tubercle are most infected, those
near the tip least. It may be in consequence of this that
the cells at and near the tip of the tubercle retain their

BoT.—VOL. II.] PETRCE—ROOT-TUBERCLES. Zitat

merismatic quality, and that they form the bowl-shaped
layer or layers of meristem which continue the growth of
the tubercle (see fig. 7). The infection threads grow out
toward the tip of the tubercle (fig. 10), but the meristem
continues to form new cells between itself and the cells
containing bacteria and infection threads. By the layer or
layers of uninfected daughter-cells the meristem protects
its own cells from infection. Perhaps because they escape
infection, they retain their ability to divide. If they can be
prevented from forming a sufficient number of daughter-
cells to enable them to escape infection, what will be the
result?

To answer this, and some other questions, I tried the
following experiment. I imbedded young tubercles on
growing roots of potted plants of Bur Clover in plaster of
Paris, according to the method devised by Pfeffer (1892)
and used by his pupils (Newcombe, 1894, Richter, 1894,
etc.). The roots were disturbed as little as possible and
were put back carefully in the soil as soon as the plaster
had been applied to the tubercles. The plants were grow-
ing in coarse sandy gravel, so that it was not difficult or
dangerous to lift out one or more roots when the soil was
well loosened by being soaked with water. After the lapse
of twenty-nine days, I again took up the roots, cutting them
off for careful examination. T'wo tubercles had been firmly
held at the base by the plaster, but had broken the casts
sufficiently at the tips to grow fresh and pink out of their
investments. New tubercles seemed to have been formed
since the roots were operated upon. One tubercle, which
had stayed in its cast, was taken out and sectioned by hand.
It was evidently dying. The bacteria in it seemed active,
but were fewer and much smaller than in unconfined tuber-
cles. There was no meristem; the cells of the tubercle
were in their definitive condition; there were smaller quan-
tities of starch in this than in ordinary tubercles. What
was true of this tubercle was equally true of others which
had failed to break and grow out of their casts.

(2) June 18, 1902.

312 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

It would seem, then, from the results just described, that
the tubercle meristem is preserved from loss of its meris-
matic qualities by escaping infection, and that the presence
of bacteria in the cells ultimately costs them their power
of division (see pp. 322-323). The meristem near the tip
of the tubercle is a survival, as is the vegetative point at
the root-tip, of the merismatic cells which constitute the
tubercle and the root in their embryonic condition.

The result of imbedding an infected tissue, a root-tuber-
cle, in plaster, is different from that described by Newcombe
(1894), who worked on healthy plants. He says that the
external mechanical resistance causes developing cells to
attain their definitive condition more slowly, continues the
merismatic power and activity of the cambium cells, and
prolongs the life of such cells as ordinarily die early.
Newcombe worked on healthy plants, and his results exhibit
the effect of pressure only on growing cells and organs. In
my experiment the pressure which checked the growth of
the tubercle-cells may not have mechanically affected the
bacteria. Since the bacteria and the cells of the tubercle
are competitors, the plaster investments handicapped the
latter to such an extent that the ultimate results of bacterial
activity appeared earlier than in unconfined tubercles, the
bacteria gaining an advantage. To the action of the bac-
teria rather than of the plaster cast are due the early loss
of merismatic power and the early assumption of their
definitive condition by the cells of the tubercle.

Ill. THe Form anp DISTRIBUTION OF ROOT-TUBERCLES.

The form and distribution of root-tubercles merit some
discussion, and since those of Bur Clover are typical I will
continue to describe them. The tubercles grow both in
length and in thickness at the ends, not at their bases,
and thus become club-shaped. They may and often do
branch. The growth takes place solely at the tip, because
the only meristem is there. The tubercle tissue is supplied
with food through vascular bundles which are neither large

Bot.—VOL. II.] PEIRCE—ROOT-TUBERCLES. 313

nor numerous but adequate for a time. A lateral root
grows both at the tip and throughout its length, thickening
and elongating. In this way its cylindrical form is main-
tained. The lateral root contributes food-materials and
water in increasing amounts to the plant which forms it.
The tubercle receives food from the plant. Perhaps it
contributes also to the nutrition of the plant. Experiment
so far seems to justify this belief. But if the tubercle were
altogether beneficial and increasingly so, one would suppose
that it would grow at the base, by secondary thickening, as
well as at the tip, by primary growth, in order through
increasing conducting tissues to contribute more and more
to the nutrition of the plant. The absence of such secondary
growth and the ultimate fate of the tubercle—dying and being
cut off—suggest that the leguminous plant limits as far as
possible the supply of food to the tubercle, and finally stops
it. Herein we have another item of evidence against Frank’s
hypothesis that the leguminous plant encourages tubercle
formation. It does not cut off the tubercle immediately;
the irritation which results in tubercle formation is too
great and the osmotic demand for food is too strong to be
resisted at once by the plant. Only after a time is tubercle
growth checked—perhaps by the remoteness of the tubercle
meristem from the source of food-supply—and later still,
the tubercle is cut off.

The tubercles may be rosy pink at and near the tip,
creamy white elsewhere, nearly or quite the same shade
as the roots bearing them. Later, the oldest tubercles,
those nearest the surface of the soil, may branch, taking on
a flat, though thick, fan shape. By no means do all of the
tubercles branch. Those very near or almost on the surface
of the soil do not, and in the lower half of the infected
portion of the root-system I have seen almost no branched
tubercles. The difference in age between the branched and
unbranched tubercles in the same or the adjacent layers of
soil is not sufficient to account for the difference in form.
The branched tubercles are the first ones to lose the plump
and healthy appearance of active life; they grow thin and

314 CALIFORNIA ACADEMY OF SCIENCES, [PRoc. 3D SER.

shriveled. They have grown fast, attained maturity early,
and they die young. The reasons for this are probably
two: First, in the upper layers of soil, which are certainly
best aérated, the bacteria in the tubercles obtain the uncom-
bined nitrogen which they absorb and fix (Mazé, 1897)
more readily and more abundantly than those in tubercles
farther down; hence, second, they grow and multiply more
rapidly, the tubercle-cells are irritated proportionally. Be-
cause the rapid growth and multiplication of the bacteria,
and, probably as a consequence, of the tubercle-meristem
cells also, are not uniform, branching occurs as a result of
some parts of the tubercle growing faster than others. The
greater activity of the bacteria and of the host-cells in these
branched tubercles is not accompanied by adequate, much
less proportional, growth of the base, and of the conducting
tissues in the base, of the tubercle. These tubercles sooner
cease to receive as much food from the leguminous plant as
they need, and hence are the first to die. Since these
branched tubercles are the largest and contain most bac-
teria, one would suppose they would benefit the plant more
than the smaller ones (if any tubercles are beneficial), and
that they would be best supplied with afferent and efferent
conducting tissues, as indicated by the proportional size of
their bases. This is not the case.

The vertical distribution of tubercles on the roots has
been reported by Frank (1890, pp. 22-3). The greatest
number and the largest tubercles occur on the lupine within
seven centimeters of the surface of the soil, and there is
a rapid decrease in both number and size as the depth
increases, till below fifty-three centimeters none was found.
The distribution on Bur Clover roots corresponds. The
strictly aérobic character of the tubercle bacteria, as shown
by artificial cultures, accounts for this, but the relations of
these organisms to the uncombined nitrogen of the air as
well as to the oxygen should be borne in mind when these
bacteria are said to be aérobic. The distribution of air in
a soil varies with the nature of the soil—a well drained
gravelly soil being well aérated to a greater depth than a

Bot.—VOL. II.] PEITRCE—ROOT-TUBERCLES. 315

heavy, compact, clay soil. It also varies with the tillage.
I have found tubercles much lower on the roots of Bur
Clover growing on a heap of gravel than in an undisturbed
and compact clay. Mechanical reasons are insufficient to
account for this. The only inference to be drawn is that
the bacteria are limited in their natural distribution to those
soils and those layers of soil which contain considerable
volumes of air, for only there will they find enough oxygen
and nitrogen for their needs.

It is the general habit of leguminous plants to send their
roots fairly deep into the soil. In a natural field, or one
returned to a state of nature, where the soil is covered by
a mixed vegetation, it is found that different plants send
their roots to different depths. In this way the resources
of the soil are more perfectly exploited by the plants and
destructive competition is avoided. But it is to be noted
that of those plants which send their roots deeper, many are
members of the Leguminose. Is this merely to escape
competition with other seed-bearing plants, or to reach a
more abundant and constant supply of water, or to escape
the attacks of the bacteria which cause them to form tuber-
cles? It is mainly the Leguminose which are successfully
attacked by tubercle bacteria, and they, as a rule, send
their roots fairly deep into the soil. Furthermore, the
number of roots increases with the distance from the sur-
face. It would appear not inconsistent with the evidence
so far obtained, to suppose that the habit of the Leguminose
of sending their roots deep, and of causing them to branch
copiously only after they have reached some depth, is one
means which these plants have of avoiding the attack of
tubercle bacteria.

Alfalfa or Lucern (J/edicago sativa Linn.) has notori-
ously long roots. They are reported to go down to the
permanently water-bearing levels of the soil. The best
chance for examining these roots would be offered when a
well is being dug where Alfalfa is growing. I have not
been so fortunate as to have such a chance, but digging
around Alfalfa plants growing as weeds in a grass field

316 CALIFORNIA ACADEMY OF SCIENCES. [PRroc. 3D SER.

shows that below a depth of twenty centimeters from the
surface the number of tubercles decreases rapidly. The
roots of this plant are perennial, and the new roots each
season are most of them formed so far below the levels in
which tubercles ordinarily occur on the roots of leguminous
plants that this plant should form a good test object of the
vertical distribution of tubercle bacteria in the soil. Com-
pared with such annuals as Bur Clover, a member of the
same genus, there were far fewer tubercles on Alfalfa roots
than on Bur Clover at the time that I dug around Alfalfa
(December). ‘There were young roots near the surface as
well as further down, but the greater number of young
roots must have been formed far below where I reached by
digging, for there were not enough young roots above
to meet the needs of the plant. Alfalfa and many other
perennial Leguminose, may therefore form the majority of
their new roots each year so deep in the soil that they can-
not become infected. As the tubercles are not perennial,
whatever advantage may accrue to the perennial legumi-
nous plant by its association with bacteria would be limited
in time and quantity to the early life of the individual, when
its roots were all in the layers of soil containing active
tubercle bacteria. The question is well worth study.

TV. THe STRUCTURE OF ROOT-TUBERCLES.

The structure of a tubercle is shown somewhat diagram-
matically in fig. 7. This is a sketch, at a magnification of
thirty-five diameters, of a section of a young and still grow-
ing tubercle. The section is parallel with the long axis of
the tubercle and at right angles with the root. The cam-
bium of the tubercle lies between @ and 4, parallel with and
in the broken curved line. This meristem is composed of
two or three layers of cells. Those toward the periphery
of the tubercle as well as those toward the center divide,
the cells toward the periphery differentiating rapidly into
cells which round off from one another and form the pow-
dery, cap-like tissue which wears away but protects
the meristem within just as the root-cap protects the

Bot.—Vot. II.] PEIRCE — ROOT-TUBERCLES. 317

growing-point of the root. The cells toward the center
differentiate somewhat diversely according as they become
infected by bacteria or continue free from them. The un-
infected cells remain comparatively small, and present the
characters of ordinary parenchyma cells, the protoplasm
becoming vacuoled and containing numerous starch grains.
There may be several vacuoles in these cells or one tra-
versed by strands of cytoplasm.

The infected cells grow larger and in their definitive con-
dition are from half as large again to twice as large as the
uninfected cells. This increase in size may be attributed
to one of three causes: First, to the stimulation of the
protoplasm by the bacteria and the substances produced
by them in the cells; second, to the actual irritation (in-
flammation) of the protoplasm; and third, to the increased
pressure set up in the cell by the rapidly multiplying bacteria.
By the plaster of Paris method we can test the relative
value of two of these influences; the third must be de-
termined by ascertaining microscopically the actual con-
dition of the protoplasm of infected cells. On imbedding
the young tubercles of growing roots, as above described,
the pressure normally or abnormally developed in the cells
will be resisted by the plaster, the cells expanding against
the plaster will be subjected to compression. Nine days
after the tubercles were enclosed in plaster, I opened the
casts and sectioned the tubercles by hand. There were
many more starch-grains in the uninfected cells than in
ordinary tubercles; I saw no infection threads (I did not
fix and use the triple stain previously described and hence
infection threads might have been present which escaped
my notice), the bacteria and bacteroids were smaller, and
the general appearance suggested that the leguminous cells
were better able to bear confinement than the bacteria
were. Increase in size both of the tubercle and of its com-
ponent cells being prevented by the plaster investment, the
bacteria have no increasing space in which to grow, and
continuing to multiply, for a time at least, they remain

318 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

small, in consequence consuming less of the food provided
for them by the leguminous plant. Starch can therefore
accumulate in the uninfected cells from which the infected
cells are osmotically supplied with food. Depositing the
non-nitrogenous food in solid form (starch) of course re-
duces the turgor pressure in the cells of the tubercle and
thereby reduces the resistance to the plaster investment.
The component cells of the tubercle enclosed in plaster of
Paris remain smaller than in unconfined tubercles, the in-
fected and uninfected cells being more nearly equal in size.

Enclosing a tubercle in plaster may diminish the stimulus
or inflammation produced in the cells by the bacteria, but
as the bacteria survive and multiply there can be no great
diminution of their chemical effect on the cells in which
they occur. The physical result, pressure, is much more
affected by the plaster investment. Since the infected cells
remain more nearly the same in size as the uninfected ones,
the inference is plain that the excessive increase in size of
the infected cells is due to increased pressure in them.

The infected cells, as shown by fig. 8, are thin-walled
and contain only one large vacuole. This is not traversed
by cytoplasmic strands. The quantity of bacteria may vary
in infected cells, and with this there is a corresponding vari-
ation in the appearance of the cells. Thus, fig. 8 shows a
typical infected cell in which the bacteria have multiplied
enormously, while fig. 9, magnified about one-third larger,
represents a cell in which there are comparatively few bac-
teria, most of which are at the point indicated by the line
from &. In the cells containing relatively few bacteria
there may be some starch-grains, as indicated by the line
from S. In unstained hand-sections, the degree to which
the older parenchymatous cells in the central part of a
tubercle are infected is indicated at a glance by the amount
of starch in the cells, the cells with the average amount of
bacteria containing no starch-grains, the cells with no bac-
teria containing many starch-grains, the cells with few
bacteria containing starch grains in inverse proportion to

Bot.—VOoL. II.] PETRCE— ROOT-TUBERCLES. 319

the number of bacteria. The cells with few or no bacteria
receive more non-nitrogenous food than they consume. The
excess they deposit in solid form as starch. The cells with
many bacteria presumably receive at least as much non-ni-
trogenous food; but either they themselves or the bacteria
in them consume this so that there is no excess to deposit.

From the cells toward the center of the tubercle the new
cells formed by the tubercle meristem are infected by means
of infection threads running fairly straight toward and into
the daughter-cells of the meristem (see fig.10). This figure,
magnification two hundred, was drawn from a thin micro-
tome section of a young tubercle, and is colored as nearly
as possible like the cells of the preparation. The prepara-
tion was stained, as previously described, by Flemming’s
triple stain, Gramm’s iodine solution being used after the
anilin gentian violet in order to differentiate the strands of
bacteria. The cell walls are drawn in black, though they
were only very faintly stained and of course were not black.
The cytoplasm is brownish yellow from orange G., the
nuclei a somewhat deeper shade of the same color, the
nucleoli red from anilin safranin, the infection threads pur-
ple from anilin gentian violet. In a fresh preparation,
whatever starch grains are present in the section are stained
the usual color by the iodine, but this color is fugitive. The
arrow to the left in this figure (fig. 10) shows the direction
in which the tubercle meristem lies. It is evident from this
figure that the infection threads run very definitely toward
the new cells formed by the merismatic layer. Since all
the food of the tubercle cells comes from the opposite direc-
tion, from the root, there must be some other directive
influence than this food to cause the infection threads to
grow so definitely toward the tip of the tubercle. This
influence must come from the cambium or from its daugh-
ter-cells, and must consist in the substance or the products
of these cells rather than in the food supplied to them. If
we are to assume any chemotactic influence, it must be
exerted by some diffusing substance or substances and

320 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

hence must be by the products and not by the living sub-
stance of the meristem and its daughter-cells. The direction
of growth of these infection threads cannot be determined
by the oxygen or nitrogen (or both) of the air, for if this
were the case, we should find strands of bacteria running
from the central cells in all directions toward the periphery
of the tubercle. This is not the case. The strands run
toward the growing-point of the tubercle. In consequence,
the daughter-cells successively formed by the repeated divi-
sion of the cells of the meristem become infected.

Not only do the infection threads run definitely toward the
growing-point of the tubercle; they also grow toward the
nucleus of each cell which they enter. This statement has
been repeatedly made and denied in papers on the subject
of root-tubercles. In hand sections, especially if the mate-
rial were not carefully fixed and differentially stained, it
would be easy to find evidence in support of the affirmation
and of the denial. Microtome sections, differentially stained
as before described, of carefully fixed growing tubercles of
the species of leguminous plants which I have especially
studied, show that in most cases the infection threads run
definitely toward the nuclei of the tubercle cells. This is
evident in fig. ro. Figures 11 and 12 also show this. In
fig. 13 are shown two tubercle cells in which the main in-
fection thread is not directed toward the nuclei, but the
lower of these two cells shows that the infection thread
bends toward the nucleus. In the next section of the series
(not figured) a branch runs from the main infection thread
to the nucleus. In the cell shown in fig. 11 the infection
thread is divided, one part running beneath, the other
above the nucleus. For the sake of clearness this upper
part of the thread was omitted in drawing. The nucleus
of the upper cell in fig. 13 was not in the plane of the sec-
tion. An adjacent section (not figured) contains this and
has a branch of the main infection thread running to it.
There must be some reason for this definite growth of the
strands of bacteria toward the nuclei of the cells which

BotT.—Vot. II.] PEIRCE— ROOT-TUBERCLES. 321

they enter.’ The effect on the nuclei is marked, as will
be shown presently.

The changes which take place in infected cells as they
develop are indicated in figs. 14 a, 6, c,d. These repre-
sent successive cells from the meristem backward toward
the center of the tubercle, which is shown in fig. 7. A
part of the cambium layer at a—0 in fig. 7 is shown at a-y
in fig. 14 a. This series is stained by Flemming’s triple
stain, but Gramm ’s iodine was not used. For this reason
the bacteria and infection threads are not differentiated.
The magnification of figs. 14 6, c, dis 300, that of fig. 14 @
is 270, hence 14 6 and 14 a do not meet exactly. The fig-
ures in the series had to be drawn separately by reason of
the limited field of the objective which was used to give the
necessary magnification.

Two cells of the tubercle cambium are shown at «-y.
Recently formed daughter-cells le toward the outside as
well as toward the center of the tubercle. The cell z is
already beginning to show the effects of infection, vacuoles
of considerable size, which later become confluent, form-
ing in the cytoplasm, and a distinct vacuole, like a halo,
appearing around the nucleolus. As shown by the cells
further toward the center of the tubercle, the nucleolus is
the first part of the nucleus to be evidently affected by the
presence of tubercle bacteria in the cell. It is the first part
of the cell to decrease in size and to disappear. If the
nucleolus is in fact an accumulation of food in the nucleus,
one would expect it to disappear, to be consumed, when-
ever there arose a special need of food in the nucleus, or
even in the cell as a whole perhaps. Furthermore, the

1W. Magnus (1900) discusses the endotrophic mycorrhiza fungus of Neottia nidus
avis I,. in this connection, stating (pp. 7-9) that the hyphe of this fungus do not grow
toward the nucleus of the cell with any regularity, but that in many other parasitic
fungi the hyphe do grow toward and around the nucleus, in some cases, however, with
no greater regularity than toward and around starch-grains or other solid contents of
the cell. He adds (p. 61): ‘‘Dass sich parasitare Pilze mit ihren Haustorien oft an den
Zellkern legen und sich in seiner Nahe eigenthiimlich verzweigen, gestattet keinen
Riickschluss auf die Bedeutung des Kernes als Nahrungscentrum der Zelle.’’ To con-
clude, because fungus hyphe or strands of bacteria grow toward the nucleus ofa cell
which they have entered, that the nucleus is the center of nutrition of the cell is
illogical; but it may well be in these cases that the nucleus contains or produces sub-
stances which nourish and attract the parasites.

322 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

greater solubility of the substance of the nucleolus than of
the substance of the nucleus and cytoplasm would make it
the first to disappear under the attack of a parasite.

In fig. 14 6 two characters of the cells are especially
noticeable: the nuclei are becoming disorganized, cavities
taking the place of the nuclear substance; and the cyto-
plasm is no longer so dense as in the younger cells shown
in fig. 14 a, nor does it stain so deeply with the orange G.
The degeneration of the nucleus is especially marked in
the cells c, 7, A. These cells have lost their nuclei, except
the last traces of acromatic substance, before they have
themselves grown to their full size.

The two upper cells of fig. 14 c have reached the maxi-
mum size of infected cells. It is impossible to say whether
they have grown to this size after their nuclei were reduced
to the condition depicted or while their nuclei were being
destroyed. I cannot say whether the smaller cells c, ¢,h
of fig. 14 6 would ever have grown to the size of the upper
two in fig. 14 4, but the unusual growth of infected cells is
due to the bacteria, and it is the bacteria which also destroy
the nucleus or cause it to be destroyed. In the upper cells
of fig. 14 c the cytoplasm is no longer the clear color pro-
duced by orange G. This brownish yellow color is dulled
by the purple of the now extremely numerous bacteria.
The smudgy appearance of the cytoplasm of the cells in 14 d
is due to the mixture of the purple stain of the bacteria and
the brownish yellow of the cytoplasm, an effect which is
very striking in the preparations. The cells of fig. 14 d
also show the greatly reduced nuclei and the large central
vacuoles. Fig. 8 represents a cell from near the center of
the tubercle shown in fig. 7. The cytoplasm is crowded
with bacteria, the central vacuole is very large, the nucleus
is reduced to an elongated lumpy mass as seen in section,
or to a thin lumpy plate in the entire cell.

When infected cells contain any considerable number of
bacteria, they cease to be able to divide. Freshly infected
daughter-cells of the cambium layer do divide, as fig. 15
shows. One is inclined to say that the disturbing, if not

Bot.—VOL. II.] PETRCE—ROOT-TUBERCLES. 323

already destructive, effect of the bacteria upon the nucleus
and other parts of the cells causes them to lose their power of
division. As has been previously pointed out (p. 311), be-
tween the tubercle meristem and the infected cells there
normally lie two or more layers of uninfected daughter-
cells. These divide also and grow. By this means the
meristem cells are kept from infection. When, however,
the growth of these daughter-cells is prevented, and the
rate of division of the meristem cells is reduced by such
mechanical pressure as a plaster of Paris investment of the
whole tubercle, the infection progresses to and into the
merismatic cells and they presently lose their merismatic
character and take on their definitive form.

One result of the division of infected cells is the produc-
tion of new cells already infected and therefore not requiring
the entrance of infection threads. As shown by Hedlund
(1892) and myself (1899), the gonidia of lichens frequently
divide in such directions as to exclude the haustoria from
some of the daughter-cells. No such result can follow
when a cell contains a number of minute parasites distrib-
uted in its cytoplasm. The cell may divide under these
conditions, but the daughter-cells share the parasites as well
as the substance of the mother-cell.

The figures above referred to, especially figs. 14 a, 6, c,
d, and fig. 8, plainly show that the presence of tubercle
bacteria is not beneficial to the cells which contain the
bacteria. So far as the relations of the bacteria and their
host-cells are concerned, no one would hesitate to call the
association a clear case of parasitism of bacteria in the cells.
Whether it is a benefit to the leguminous plant to form
tubercles, to harbor bacteria in these, to have the cells of
the tubercles destroyed, and all the food supplied to the
tubercle consumed, is another question. The most careful
experimentation and the critical examination of the results
of experiment have so far led to the generally accepted
belief that the association of bacteria with the roots of
leguminous plants is beneficial to the leguminous plants. I
find it hard to understand how association with bacteria

324 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

which destroy the cells in which they occur can benefit the
larger member of the association. The bacteria are known
to fix free nitrogen from the air. By this means they form
in the cells of their host-plants nitrogenous compounds which
the host may use. Apparently the bacteria form more than
enough of these valuable nitrogen compounds in the cells
of their hosts to compensate for the extra material used in
forming and maintaining tubercles the cells of which are
ultimately destroyed by the bacteria. This implies a mar-
velous balance of profit and loss, the more remarkable since
the profit apparently exceeds the loss.

One point more needs to be made clear. Miss Dawson
(1899, p. 14) says that it is difficult to conceive how
such strictly aérobic bacteria as these can flourish in
the cells of such compact tissue as composes the tuber-
cle. This difficulty is of her own conceiving, for do not
the cells of the tubercles respire and are they not neces-
sarily supplied with oxygen for respiration? Again, inter-
cellular spaces in the infected tissue do occur, as figures
140 G STANC,' 135-124) 85;).0,7ane 9 plainly show. Even if
intercellular spaces did not occur, as asserted by Schneider
(1893, pp. 786 and 787), the existence of the living cells of
the tubercle tissue would prove the presence of sufficient
quantities of oxygen, if not of nitrogen, in the tissue and
therefore in the cells. Unless we are to imagine anaérobic
respiration for these cells, it is unnecessary to assume it for
the bacteria which infest them. Fischer (1897, note 63 to
p. 92), shows this clearly, and my sections reveal the pres-
ence of intercellular spaces through which a diffusion of
gases could take place even if the diosmosis of solutions of
the gases concerned were inadequate to supply the demand.

SUMMARY.

1. Though the bacteria which form root-tubercles on
leguminous plants are usually only slowly motile, if
motile at all, in artificial cultures, this proves nothing
as to their motility in the soil.

Bor.—VoL. II.] PEIRCE—ROOT-TUBERCLES. 325

2.

LO.

The proportion of root-hairs infected to the total num-
ber formed is small, in one case computed to '
be 1:1000.

Given the contact or close proximity of tubercle bac-
teria with the root-hairs, infections may take place
in great numbers simultaneously, at least when the
roots are very young.

Infection of the root may be resisted by cutting off the
infected ends of the root-hairs.

The tubercle bacteria enter and infect a root-hair by
softening or dissolving a small portion of the wall
and moving or growing through this. There is no
evidence that they usually enter through broken
root-hairs, and the curvatures of infected root-hairs
are evidence against these hairs having been broken
at any time.

The infection thread grows fairly straight, being
chemotropically attracted, through the cortical paren-
chyma, from the root-hair to the layer of cells next
outside of the central cylinder of the root.

The tubercles originate only endogenously and from
the same layer as gives rise to lateral roots. We
may therefore conclude that the tubercles are mor-
phologically lateral roots, though greatly modified by
the influence which caused them to be formed.

Tubercles form only as the result of stimulation by
bacteria. Do lateral roots form as the result of
internal causes or of external stimuli?

The growth of the tubercle is apical, the daughter-cells
of a bowl-shaped terminal meristem constituting the
growing part of the tubercle. There is little or no
secondary growth in thickness. Because of this,
the conducting tissues do not keep pace with the
growth of the tubercle. The growth of the tubercle
is correspondingly limited.

Tubercles are largest and most numerous near the
surface of the soil. It is possible that perennial
Leguminose form few if any tubercles after their
roots have grown deep into the soil.

326

II.

12.

i:

14.

TS:

16.

I7.

CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

The presence of bacteria in the cells of a tubercle pre-
vents the infected cells from forming starch granules.
Uninfected cells do not attain the size usually
reached by infected cells. The larger size of in-
fected cells is due to increased pressure, probably
also the greater irritation, in these cells.

The bacteria cause the degeneration and almost com-
plete destruction of the nuclei of the cells in which
they occur.

The infection strands grow definitely, chemotropically,
toward the daughter-cells formed by the tubercle
meristem, and seem also to grow definitely toward
the nuclei of the cells into which they penetrate.

Infected cells soon lose their power of division, though
not of growth.

The presence of bacteria in the cells of the tubercle is
injurious to these cells, and the relation of the bac-
teria to their host-cells is parasitism.

It is difficult to understand how the leguminous plant
as a whole can profit by an association which is
injurious and finally destructive to the cells in which
the bacteria occur.

Intercellular spaces occur in the tissues of root-
tubercles. Even if they did not, it would not be
necessary to assume that the bacteria live anaérobi-
cally therein, since the tubercle cells do not live
anaérobically.

LELAND STANFORD JR. UNIVERSITY,

PaLo ALTO, CALIFORNIA,
July 27, 1901.

BoT.—VOL. II.] PEIRCE— ROOT-TUBERCLES. 327

1890.
1893.
1898.
1899.
1900.
1897.
1900.
1896.

1892.

1899.
1894.
1893.

1897.
1goI.

1894.
Igor.
1893.
IgoI.
1892.
1894.
1894.
1897.

1900.

BIBLIOGRAPHY.

Frank, A. B. Ueber die Pilzsymbiose der Leguminosen. Sonder-
abdruck aus den Landwirthschaftlichen Jahrbiicher. Berlin.

ZIMMERMANN, A. Botanical Microtechnique. New York. Transl.
by Humphrey.

Hor, A. C. Histologische Studien an Vegetationspunkten. of.
Centralblatt, Bd. LX XVI, pp. 65-69.

Dawson, MariA. Nitragin and the Nodules of Leguminous Plants.
Phil. Trans. Roy. Soc.

MicuLa, W. System der Bakterien. Bd. I, 1897; II, 1900. Jena.

ZINSSER, O. Ueber das Verhalten von Bakterien, insbesondere von
Knollchenbakterien, in lebenden pflanzlichen Geweben. Jahrb. /.
wiss. Bot., Bd. XXX.

Dawson, Marta. Further Observations on the Nature and Functions
of the Nodules of Leguminous Plants. Phil. Trans. Roy. Soc.

CzaPEK, F. Zur Lehre von den Wurzelausscheidungen. /ahrd. f.
wiss. Botanitk, Bd. XXIX.

HEDLUND, T. Kritische Bemerkungen tiber einige Arten der Flech-
tengattungen Lecanora, Lecidea, und Micarea. Bihang till K.
Sven. Vet.-Akad. WHandl., Bd. XVIII.

PEIRCE, G. J. The Nature of the Association of Alga and Fungus in
Lichens. Proc. Cal. Acad. Sci., 3d Ser. (Bot.), Vol. I, No. 7.

SPALDING, V. M. The Traumatropic Curvature of Roots. <Azn.
Botany, Vol. VIII.

PerRcE, G. J. Structure of the Haustoria of Some Phanerogamic
Parasites. Ann. Botany, Vol. VII.

FiscHER, A. Vorlesungen tiber Bakterien. Jena

SmitTH, E. F. Entgegnung auf Alfred Fischer’s ‘‘ Antwort” in be-
treff der Existence von durch Bakterien verursachten Pflanzenkrank-
heiten. 2ter Theil. Centralbl. f. Bakteriol., Parasitenkunde u.
infektionskrankh., 11 Abtheilung. (The special literature is here
cited.)

Peirce, G. J. A Contribution to the Physiology of the Genus Cus-
cuta. Ann. Botany, Vol. VIII.

Cannon, W. A. The Anatomy of Phoradendron villosum Nutt.
Bull. Torr. Bot. Club, Vol. XXVILI.

SCHNEIDER, A. The Morphology of Root-tubercles of Leguminosz.
American Naturalist, Vol. XXVII.

Lire, A. C. The Tuber-like Rootlets of Cycas revoluta. Bot. Ga-
zette, Vol. 31.

PFEFFER, W. Ueber Anwendung des Gipsverbandes fiir pflanzen-
physiologische Studien. Ber. d. K. Sachs. Gess. d. Wiss.

NewcomeE, F.C. The Influence of Mechanical Resistance on the
Development and Life-period of Cells. ot. Gazette, Vol. XIX.

RICHTER, J. Ueber Reaktionen der Characeen auf iiussere Reize.
Flora, Bd. LXXVIII.

Maze. Fixation de l’azote libre par la bacille des nodosités Légumi-
neuses. Azz. de l’Inst. Pasteur, T. x 1.

Macnus, W. Studien an den endotrophen Mycorrhiza von Neottia
Nidus avis L. /ahrd. f. wiss. Bot., Bd. XXXV.

328 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

EXPLANATION OF PLATE XXIX.

Most of the figures were drawn with the aid of a Leitz drawing prism
ocular, the rest with an Abbé camera lucida.

Fig. 1. Two root-hairs of Bur Clover infected by tubercle bacteria,
showing the characteristic bending at the point of infection; x 50.

Fig. 2. The lower of the two root-hairs of fig. 1. The mass of bacteria
in the concavity of the coil and the infection thread running from this point
through the hair; x 300.

Fig. 3. Another infected and coiled root-hair with the infection thread
growing close to the nucleus of the hair; x 300.

Fig 4. A section showing the course of the infection thread from the
base of the hair (7. #.) to the layer of cells which gives rise to the tubercle,
the same layer which, under other conditions, would give rise to a lateral
root; x 360.

Fig. 5. A section showing a young tubercle (left) and a young lateral
root (right) developing from the same layer; x 20.

Fig. 6. The same section enlarged. In the lateral root (right) differen-
tiation of tissues is already taking place, but notin the tubercle (left): x 360.

Fig. 7. Longitudinal section of a tubercle, young and still growing.
The largest part of the tubercle is composed of bacteria-containing cells.
Outside of this mass is the comparatively thin layer in which are the small
and scattered vascular bundles. Beyond this is the protective tissue, pow-
dery on the outside, which is continuous with the cortex of the root. At
a-6 is the tubercle meristem, which forms daughter-cells both forward and
backward, as does the growing-point of a root; x 35.

Fig. 8. Atypical infected tubercle cell, thin-walled, with the cytoplasm
surrounding a large single vacuole, and the nucleus reduced to a small lumpy
plate. Cytoplasm stained with orange G., but the color obscured and dulled
by the purple of the bacteria stained by anilin gentian violet. The nucleus
is purplish from the mixture of anilin safranin and anilin gentian violet, a
wholly different color from that of a healthy nucleus stained and differentiated
by Flemming’s triple stain and Gramm’s iodine solution. Intercellular
spaces are shown; x 660.

Fig. 9. Tubercle cell containing only small number of bacteria (at 2) and
enclosing some starch-grains (.S). The nucleus has degenerated only some-
what and the vacuolization of the cytoplasm has not progressed far; x 820.

Fig. 10. Section of a tubercle, part of tubercle shown in fig. 7, near the
meristem. The direction in which the meristem lies is indicated by the
arrow at the side. The section was stained by Flemming’s triple stain and
differentiated, after the anilin gentian violet, by Gramm’s iodine. Thus the
infection threads are clearly brought out. Note the course of the infection
threads, definitely toward the tubercle meristem and generally toward the
nuclei of the cells entered; x 200.

Fig. 11. One cell from fig. to (nearly in the center of the figure) showing
the solid infection strand (zodglcea) in which the separate bacteria can be
distinguished; x 1000.

Fig. 12 and 13. Tubercle cells showing the infection threads growing in
definite direction, generally toward the nuclei of the cells; x 820.

Fig. 14, a-d. A series of cells from the meristem backward in a longitu-
dinal section of a tubercle. The section is stained by Flemming’s triple
stain, but not differentiated by Gramm’s iodine, hence infection threads do
not show. In fig. 14 a, two meristem cells show at 2-y, with daughter-
cells both forward and backward. Z=already infected and degenerating
cell. In 14 6, degeneration of the cell and especially of the nucleus is shown
by cells c, 7, %. Degeneration of the nucleus, obscuring of the color of
the cytoplasm by the many bacteria in it, and the formation of the char-
acteristic vacuole shows in 14 ¢c, especially the two upper cells. Fig. 14d
shows the same still more markedly, especially the uppermost cell. In this
series the effect of bacterial infection of the cells is clearly exhibited; 14 a, x
270; 14 b-d, x 300.

Fig. 15. Division of an already, though recently, infected daughter-cell
of the tubercle meristem; x 300.

[PetRce)] Flare XXIX.

c..3? Ser.Bor. Vou.

A ica hes
Apap. athe

SPINDLE FORMATION IN THE POLLEN-
MOTHER-CELLS OF CASSIA
TOMENTOSA L.

BY HENRI T. A. HUS.

CONTENTS.

PLATES XXX-XXXII.

PAGE

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INTRODUCTION.

Cassia tomentosa L. possesses a number of qualities
which particularly recommend this plant to the cytologist.
Chief among these are the large number of buds produced
and the protracted flowering period. The stages of cell-
development which are of greatest interest at the present
time are passed through very rapidly, and are also the most
difficult to fix; hence the collection of an immense amount
of material is necessary to ultimate success. At the same
time the protracted flowering period affords an opportunity
to determine at leisure which is the best fixing-medium and
what shall be the duration of the fixing-period.

The following fixing-fluids were tried:

[329] April 6, 1904.

330 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

Bs Strong Flemming:
I per cent. chromic acid 15 pts.
2 per cent. osmic acid 4 pts.

Glacial acetic acid I pt.
2-6. No. 1 diluted with 1-5 volumes of water.
fe I per cent. chromic acid 15 pts.
2 per cent. osmic acid 4 pts.

I per cent. glacial acetic acid I pt.
8-12. No. 7 diluted with 1-5 volumes of water.

13. I per cent. chromic acid 15 pts.
2 per cent. osmic acid 4 pts.
14-18. No. 13 diluted with 1-5 volumes of water.
19. ¥% per cent. chromic acid 15 pts.
2 per cent. osmic acid 4 pts.
Glacial acetic acid Type
20. No. 19 diluted with 1 volume of water.
Ae I per cent. chromic acid.
22. 2 percent. osmic acid.
23. I per cent. osmic acid.

24-26. I, 5 and 10 per cent. aqueous solution of bichro-
mate of potash.
27. Tellyesniczky’s fluid (Arch. mikr. Anat 52247.
1898) :
Bichromate of potash 3 gm.
Glacial acetic acid 5 cc.
Distilled water IOO cc.
28-32. No. 27 diluted with 1-5 volumes of water.
33- Miiller’s solution:
Bichromate of potash 2-2% pts.
Sulphate of soda To) ab
Distilled water 100 pts.
34. Zenker’s mixture:
Corrosive sublimate 5 per cent.
Glacial acetic acid 5 per cent.
Miiller’s solution 90 per cent.
35. 1Saturated aqueous solution of bromine I pt.

a

1 The material was immersed for two hours in this solution, and then removed to
Flemming’s strong mixture, where it remained eight hours.

Bot.—VOL. II.] HUS—CASSIA TOMENTOSA L. 331

Distilled water 9 pts.
36. Y per cent. iodine

1% per cent. potassium iodide I pt

Distilled water

Distilled water 9 pts.

I cannot here enter upon an extensive discussion of the
effects of these fixing-fluids. It is sufficient to say that
material fixed for ten to twelve hours in Flemming’s strong
mixture showed as a rule no shrinkage; while material
fixed in other solutions (especially in the very dilute ones)
or left longer than twelve hours in Flemming’s strong mix-
ture, showed considerable shrinkage, with the exception
perhaps of that fixed with Tellyesniczky’s fluid, which
gave fairly good results.

In fixing, two rules were observed: one, always to fix in
the field, since only in material fixed in this manner were
some of the earlier stages found in any number; the
other, to keep the material submerged, for only then could
the necessary penetration of the fixing-fluid take place.
Frequently air-bubbles adhered to the anthers, especially if
the latter were not separated, causing them to float. To
prevent this a very simple device was used: a disc was cut
off the lower end of the cork of the neckless fixing-bottle
and attached to a glass rod passing through the cork; this
kept the material submerged, while a notch in the edge of
the disc allowed the escape of any air-bubbles.

After fixing, the material was washed from six to eight
hours in running water; a longer stay in water apparently
injured the finer structure of the cytoplasm.

At this stage the material was sorted and then dehydrated.
It appeared here that a prolonged stay in weak alcohol
produced shrinkage. After reaching 95 per cent. alcohol,
the material remained for periods of about twelve hours
each, in absolute alcohol, a mixture of equal parts of abso-
lute alcohol and bergamot oil, bergamot oil, a mixture of
equal parts of bergamot oil and 43° C paraflin, 43° C
paraffin and 54° C paraffin. It was then imbedded in io C

332 CALIFORNIA ACADEMY OF SCIENCES. [PROC. 3D SER.

paraffin, the use of which was found necessary to obtain
good ribbons. Sections 3-5 ». thick were cut and fixed
on the slide by the water-albumen method.

Flemming’s triple stain was used, with some modifica-

tions. The length of time for each stain was ascertained
separately. It was finally found advisable not to use the
safranin solution usually employed, but to substitute for it
Babes’ safranin A, consisting of a mixture of equal parts
of a saturated alcoholic solution of safranin and a saturated
aqueous solution of safranin, because the former had a
tendency, when old, to give a muddy appearance to the
cytoplasm.
‘ In the case of orange G, even weak solutions took out too
much blue. By substituting for this a solution of iodine
and potassium iodide of the same strength and composition
as the fixing-fluid mentioned under number 36, much better
results were obtained. In fact the potassium iodide
iodine seemed to fix the blue in the fibers, so that absolute
alcohol and clove-oil could be relied upon to wash out the
superfluous blue.

The staining process was as follows: After dissolving the
paraffin in xylol and removing all traces of the latter by a
double washing in 95 per cent. alcohol, the slides were
placed in the safranin for five minutes and then decolorized
in absolute alcohol to which o.1 per cent. hydrochloric acid
had been added. After washing thoroughly in water to
remove all traces of the acid, the slides were placed for
exactly five minutes in a concentrated aqueous solution of
gentian violet, and then washed off in water; after which
they were immersed for twenty seconds in the solution of
iodine and potassium iodide above referred to. This
appeared to fix the blue in an admirable way, so that after
dehydrating in absolute alcohol for one second, the prepa-
rations, in clove-oil, could be watched under the microscope
until the cytoplasm was perfectly clear, the fibers a dark
blue, and the granular zone brown-violet. The nucleolus
was a bright red as well as the chromatin, except in the
earlier stages, when the chromatin stained blue.

ee eS es rr eC

Bot.—VOL. II.] HUS—CASSTA TOMENTOSA L. 333

DESCRIPTION.

The nuclei of the pollen-mother-cells of Cass¢a tomentosa,
in the stage indicated in figure 1, show a nucleolus, which is
sometimes vacuolate, and a broken up chromatin thread.
The former stains red, the latter blue or red. The cyto-
plasm is composed of two parts, one apparently fibrous,’
the other granular. The first is composed of thinner or
thicker fibers running in every direction, but more especially
from the nuclear wall to the cell-wall, so that the meshes
which they form are elongated ina radial direction (fig. 1).
The meshes are smallest in the neighborhood of the nucleus,
and gradually increase in size toward the periphery.

The other component of the cytoplasm appears in the
form of larger or smaller granules situated between and
upon the fibers, and showing a tendency to accumulate in
larger or smaller masses, especially at the intersection of
the fibers. The fibers can be stained blue with gentian
violet, while the granules stain brown-violet. The whole
presents a very uniform appearance, except here and there
where a thicker fiber or a denser accumulation of granules
shows more prominently.

Scattered irregularly through the cell may be seen small
dark round bodies, apparently oil globules, to judge by their
appearance in fresh material. It may be of interest to note
that while these bodies were very numerous in some cases,
especially in the early stages, in others they were rare and
were then found but seldom in the later stages.

Gradually the cytoplasm around the nucleus begins to
change in appearance: the change takes place in a single
layer of meshes immediately adjacent to the nuclear wall;
these meshes become smaller, due perhaps to the interpola-
tion of new fibers, and to the appearance immediately
around the nucleus of granular matter, situated upon and
between the fibers, often completely obscuring them. The
meshes now become elongated parallel to the nuclear wall;

1The description of the structure of the cytoplasm refers to microscopical appearance
only, without indicating whether it is a reticulum ora foam-structure.

334 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

they become very narrow, being usually from two to three
times as long as broad. The fibers are covered with gran-
ules; they begin to stain a deeper blue than the other fibers.
This appearance extends only partially around the nucleus,
usually about half way (fig. 2.)

Changes now begin to take place in the cytoplasm farther
away from the nucleus: the cytoplasm loses its uniform
appearance; here and there fibers have begun to thicken,
or granules have accumulated upon them. These fibers
stain a deep violet and show sharply against the lighter
background (fig. 3). They are arranged in more or less
conical groups with their bases directed toward the nucleus.

Very gradually, almost imperceptibly, the granules
around the nuclear wall increase in number, and a second
layer of elongated meshes is laid down next the one already
formed. Inthe meantime the granules situated upon and
between the fibers have disappeared, and the fibers have
become thick and smooth (fig. 3). We nowsee the nucleus
partly surrounded by a number of rectangular meshes
elongated in a direction parallel to the nuclear wall.

These processes seem to continue for some time, until
finally three or four layers of elongated meshes have been
laid down around the nucleus. Then the radial fibers of
these meshes begin to disappear, leaving a zone composed
of fibers running parallel to the nuclear wall. That this
zone is not attached closely to the latter can be easily
observed in later stages when one or more spaces exist
between the nuclear wall and the fibrous felt-like zone,
which stands out clearly as a dark blue band.

In the meantime granules have continued to accumulate
around the nucleus, and the meshes of the cytoplasm adjoin-
ing the felt-like zone have grown smaller. Together they
form a distinct zone (the granular zone) which possesses a
distinct outer boundary.

The cell now presents the appearance indicated in figure
4. The chromatin appears in the form of lumps, which
like the nucleolus, stain a brilliant red. Here and there
linin, in the form of violet-staining granular threads, may

Bot.—VOL. II.] HUS—CASSIA TOMENTOSA L. 335

be seen connected with the chromosomes. The nucleus
is more or less completely surrounded by a zone of smooth
parallel fibers which stain a dark blue. Next these lies
the granular zone, occupying about half the remaining
cell-space, and composed of a reticulum of small, more or
less isodiametric meshes, between which (especially at the
intersection of the threads) a great number of larger and
smaller granules may be seen. Surrounding this zone
(which stains brown-violet) is a dark blue line apparently
composed of fibers upon which granules, large and small, are
situated. These fibers are not continuous, and the number
of granules upon them is not the same in all places, so the
ring which they form is an irregular one. It stains a deep
blue, very effectually marking the boundary between the
brown-violet granular zone and the outer gray-yellow cyto-
plasm, which is composed of fibers forming large irregular
meshes. Upon and between these fibers are larger or smaller
masses of granules. The granular zone does not present
an absolutely uniform appearance; in places fibers upon
which granules have accumulated stand out clearer than
others. Usually they run parallel to, or at a small angle
with the nuclear wall. At first only one or two are seen,
but they rapidly increase innumber. Soon those nearest the
nuclear wall seem to establish connection with some of the
fibers of the felt-like zone (fig. 5); gradually other, deeply-
staining fibers are added, all running at a greater or lesser
angle to the nuclear wall, and finally arranging themselves
in conical groups with their bases resting on the fibrous
zone. Not only one cone like this may be observed, but
several (fig. 6). During this time the number of prominent
fibers in the granular zone continues to increase. Fre-
quently a space can be observed, at least in places, between
the nuclear wall and the felt-like zone.

At about this stage the nucleolus begins to disappear; in
the preceding one, however, the nucleolus was still present,
and the chromatin was in the form of curved, rod-shaped
chromosomes, or else in the form of rings, formerly con-
sidered so typical of the Liliacee.

336 CALIFORNIA ACADEMY OF SCIENCES. ([Proc. 3D SER.

The linin of the nucleus has gradually become more
prominent, staining a darker blue; but while in the pre-
ceding stage it showed as fine granular threads, it now
(fig. 6) appears as smooth fibers, running through the
nucleus, and usually more or less parallel to each other and
to the axis of the largest cone. The fibers stain blue.

The cones, which are composed of granular threads,
increase in number. Some are larger; some smaller.
Usually one has the ascendency over the others, and this,
as arule, is the first cone formed; it appears ordinarily at
one end of the ellipsoidal nucleus. As has been said, it is
usually parallel to the axis of this largest cone, and con-
sequently more or less parallel to the longer axis of the
nucleus, that the linin threads are directed. This seems to
indicate a polarity of the cell.

In figure 6 we find the cones composed of granular
threads; these threads are in connection with the fibers of
the felt-like zone. In the next stage we find cones, large
and small, composed of smooth fibers. These fibers appar-
ently originate from the felt-like zone. But do they push
out of their own accord? or are they pulled out by the
granular fibers ? or are the cones a result of a combination
of the granular fibers and those of the felt-like zone?
Judging from my preparations the last seems to be the
most probable.

As soon as the fibers of the cone have become smooth,
the nuclear wall just below the cone breaks down, and the
fibers penetrate into the nuclear cavity, where they anasto-
mose with the linin threads. The two kinds of fibers can
hardly be distinguished, for while it is true that the fibers
originating from the cytoplasm are thicker than those
formed from the linin, even this difference very soon dis-
appears.

Even at this early stage the fibers arrange themselves
in bundles (fig. 7), an arrangement which is far less promi-
nent in the fully developed multipolar stage, but which
reappears in the bipolar spindle.

The chromosomes retain their original position for a long

Ee lee

Bot.—VOL. II.] HUS—CASSIA TOMENTOSA L., 337

time after the nuclear wall has disappeared. Only when
the multipolar spindle has been fully formed, and in fact
not until a rearrangement of the poles (with a view to the
formation of a bipolar spindle) has begun, do they begin
their migration toward the spot where the equatorial plate is
to be formed. Throughout the formation of the cones and
the subsequent changes from multipolar to bipolar spindle,
no indication of a centrosome could be observed.

As soon as the fibers composing the cones have become
smooth, the nuclear wall breaks down and the blue-staining
fibers penetrate the nuclear cavity, where they lie in contact
with the bright red chromosomes. The multipolar spindle
thus formed has a variable number of cones; the largest
number observed in a single section was eight (fig. 10) ;
but as the cell was always cut into a number of sections,
the number of cones must be greater.

The granular zone now presents a uniform appearance,
and stains from yellow-brown to brown-violet; the granular
threads which were formerly so prominent have disappeared ;
nothing remains but small meshes which are fairly uniform
in size. Between and upon the threads composing the
meshes a large number of granules can be observed. The
poles protrude into the granular zone. Surrounding the
granular zone we find an irregular line still staining a deep
blue. The cytoplasm outside this line retains the same
structure as in the last stage.

In the formation of the multipolar spindle a highly inter-
esting phenomenon was observed. It has been stated
before that, as a rule, a cone situated at one end of the
nucleus had from the first an ascendency over the others;
and while it was not observed that another cone (situated
directly opposite this one) was always the second to be
formed, yet one could not fail to see that after a little while
two cones (one at each end of the nucleus) were more
prominent than-the others, or were even the only cones
present, in which case the nuclear wall disappeared beneath
these cones while persisting everywhere else. Such a case
is shown in figure 8. In nearly every instance this promi-

338 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

nence of two cones could be noted. If my observations
are correct, we have in Cassza tomentosa a most interesting
transition stage between the multipolar polyarchal and the
multipolar diarchal spindle.

The largest number of poles of the multipolar spindle
observed in cross section was eight; the most frequent
number noted in a section was from four to six. In what
manner the multipolar spindle finally became a bipolar one
could not be determined with certainty; the cones
approached each other in two groups (fig. 12), finally
forming the bipolar spindle.

The cell now presents the appearance indicated in figure
13. The chromosomes lie in the equatorial plate. In
polar view, twelve chromosomes can be observed. The
spindle has sharp-pointed poles. Apparently there are here
three kinds of fibers; some having the appearance of
strands are attached to the chromosomes; some run from
pole to pole; while from each pole mantle-fibers diverge
into the granular zone. The fibers stain a deep blue; the
granular zone is yellow-brown; but the irregular line around
the granular zone is as blue and sharp as it was before.
The poles of the spindle project into the granular zone.

The wandering of the chromosomes toward the poles
now begins in the usual manner; the continuous fibers still
run from pole to pole; while the mantle-fibers become
more prominent.

At this stage we find, sometimes inside the granular zone,
sometimes outside it, small round bodies which stain red,
sharply contrasting with other similar bodies, lying on the
outskirts of the granular zone, which stain a deep violet,
and look like the oil-globules previously observed.

Finally the chromosomes reach the poles, and the contin-
uous fibers as well as the mantle-fibers assume a wavy
appearance. The mantle-fibers during the last stage diverge
more than formerly.!. Why the continuous fibers should
assume this wavy appearance is not clear. Certainly the

1 Perhaps it would be well tostate here that these mantle-fibers were not equally
prominent in all preparations. In fact in some cases no trace of them could be found.

a

Bot.—VOL. II.] HUS—CASSIA TOMENTOSA L. 339

suggestion of Williams (1899, p. 194; wzde Went, 1887,
p. 258, figs. 8-10) that it is caused by the drawing together
of the poles, appears very acceptable. Inthe case of Cassza
tomentosa the spindle poles either reach to the outer edge
of the granular zone, or else pass through it (figs. 13, 14),
yet the daughter nuclei are always formed well within it.

The most interesting point at this stage is the behavior
of the granular zone, which shows a tendency to accumu-
late around the daughter nuclei as they are about to be
formed. Gradually it surrounds them on all sides except
where the connecting fibers remain. As soon as a wall is
formed around the daughter nuclei, they become entirely
surrounded by granular matter. |

It was the last stages of this process which were particu-
larly easy to follow in my preparations. To say that the
granular matter encroached upon that part of the cell which
contained the yet remaining continuous fibers would not be
quite true; the fibers seemed rather to thicken at various
places in the immediate neighborhood of the daughter
nuclei, and this apparently at the cost of the immediately
adjacent parts of the fibers. This process continued until
nothing was left of that part of the continuous fibers but a
granular thread, which sooner or later lost its continuity,
so that gradually the place where the threads were, was
entirely taken up by granules.

This process takes place very gradually, beginning at
the sides of the daughter nuclei and proceeding toward
the axis of the cell. At first the meshes formed by the
transformation of the connecting fibers into granular matter
are fairly large, but soon they can no longer be differen-
tiated from those of the rest of the granular zone. This
change, especially in its later stages, is very slow, and may
not be consummated until the daughter nuclei are already
forming spindles.

The daughter nuclei now lie in a granular mass in no
way different apparently from that which surrounded the
mother nucleus. The shape of the granular zone is not
quite circular, but more or less ellipsoidal, slightly con-
stricted about the middle.

340 CALIFORNIA ACADEMY OF SCIENCES. [PROC. 3D SER.

The shape of the daughter nuclei is decidedly ellipsoidal,
with the longest axis at right angles to the main axis of the
previous spindle. The daughter chromosomes are irregular
in shape: sometimes they take the form of lumps; at other
times they appear as chains of granules. They stain red
and are connected by bands of finely granular matter. A
body which seems to be a nucleolus appears in each
daughter nucleus.

Owing to the presence of a dense granular zone around
each daughter nucleus, it was impossible to follow in this
case the stages just preceding the formation of the multi-
polar spindle; yet what could be seen was sufficient to
indicate that these stages agreed in the main with the cor-
responding stages of the mother nucleus. The differences
observed are to be explained by the shape of the spindle,
which in all cases is of a pronounced multipolar diarchal
type.

The first indication of spindle formation that could be
observed was the appearance of a mass of clean-cut,
closely-woven fibers surrounding each daughter nucleus.
These fibers were particularly distinct where the nuclear
wall had shrunken somewhat. They stained a deep blue
(fig. 17).

The multipolar spindles of the daughter nuclei are formed
in a manner similar to that of the mother nucleus. Linin
fibers become more and more prominent within the daugh-
ter nucleus; they run more or less parallel to the longest
axis of the nucleus in which they are formed. Cones begin
to appear here and there on the nuclear wall, usually at the
ends; they increase innumber. In some places the nuclear
wall breaks down; the cytoplasmic fibers enter the nuclear
cavity, and become merged with the threads of nuclear
origin. Finally a multipolar spindle is formed, which, as
has been previously stated, is of a pronounced multipolar
diarchal type. The narrow, compressed appearance of the
entire multipolar figure is here very striking, far more so
than in the multipolar spindle of the mother nucleus. In
no case was as large a number of poles observed as in the

BoT.—VOL. II.] HUS—CASSTA TOMENTOSA L. 341

corresponding stage of the mother nucleus. The planes in
which the spindles of the daughter nuclei lie are sometimes
at right angles, sometimes parallel to each other.

Finally the daughter chromosomes reach the poles and
become surrounded with a membrane. The chromatin is
in the form of granules, arranged in some places to form a
thread, and interconnected by bands of the same finely
granular substance which we saw connecting the chromo-
somes of the daughter nuclei. The nucleolus, which dis-
appeared at the time of the dissolution of the membrane of
the daughter nuclei, now again makes its appearance.

The granular zone surrounds each of the four daughter
nuclei very much as in the stages just following the first
division of the mother nucleus; but instead of filling up the
entire space between the nuclei, it merely surrounds them,
leaving a clear space in the middle through which fibers
may be seen running from one nucleus to the other.

DISCUSSION.

In the formation, development, and subsequent behavior
of the spindle there are two points of special interest; the
nature of the multipolar spindle, and the origin of the
cones.

Tue NATURE OF THE MULTIPOLAR SPINDLE.

In the first as well as in the second divisions of the
pollen-mother-cells of Cassza tomentosa there is more or
less approach to the spindle formation described for vege-
tative cells. Instead of a multipolar polyarchal spindle
Anlage (Strasburger, 1900, p. 121) such as we usually
meet with in pollen-mother-cells, we have here a more or
less multipolar diarchal spindle. At the very beginning we
meet with an indication of this. The nucleus is usually
ellipsoidal. The first cone (afterwards the largest cone)
appears ordinarily at one end of the ellipsoidal nucleus.
The second cone formed lies in many instances directly
opposite the first one, or in other cases, the cone which

342 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

occupies this position becomes, as a rule, more prominent
than those surrounding it. These two cones remain the
most prominent ones. More or less parallel to the axis of
these cones, and consequently more or less parallel to the
longer axis of the nucleus, run the linin threads. The
multipolar diarchal nature of the spindle is even more
prominent in the second division.

Thanks to the numerous investigations of recent years,
we are now able to compare a number of modes of spindle
formation. Strasburger (1900, p. 118) distinguishes three
types of spindle formation; the multipolar polyarchal, the
multipolar diarchal, and, opposed to these, the bipolar
diarchal.

Multipolar polyarchal spindles are characteristic of the
first division of pollen-mother-cells (Lzlcum, Passzflora,
Lavatera, Cobea, Gladiolus) and of spore- and embryosac-
mother-cells (Osterhout, 1897; Strasburger, 1900, p. 121).
In Cassia tomentosa we have a multipolar diarchal spindle
Anlage in the first division, which thus forms a connecting
link between the multipolar polyarchal spindle Anlage of
reproductive cells and the multipolar diarchal spindle de-
scribed for vegetative cells, i. e., the root-tips of Vzcza
(Strasburger, 1900, p. 116), of Aldzum (Nemec, 18992),
and of ferns (Hof, 1898, p. 169). In Mymphea alba
(Strasburger, 1900, p. 122) the spindle Anlage of the
mother nucleus does not differ essentially from that in the
daughter nuclei described by Strasburger for /rzs squalens,
where it is multipolar diarchal.

It seems therefore that there exists no sharply marked
difference between spindle formation in reproductive and
in vegetative cells, and that the two processes are closely
related.

As suggested by Strasburger (1900, p. 122), the various
modes of spindle formation would readily lead one to
suppose them influenced by polarity: ‘‘ Eine Pollen-
und Sporenmutterzelle kénnte auch in ihren Protoplasten
multipolar sein und so die polyarche Anlage der Kern-
spindel bedingen. Auch liesse sich denken, das die

Bot.—Vot. II.] | HUS—CASSIA TOMENTOSA L. 343

diarche Anlage der Kernspindeln in Gewebezellen eine
Folge ihrer longitudinalen und radialen Polaritat sei.’’

Polarity is in many cases probably determined by causes
outside the cell; for though in the first division of the
pollen-mother-cells and in most divisions in vegetative cells,
the long axis of the spindle coincides with the long axis of
the cytoplasmic mass, yet there are numerous cases (both
in reproductive and in vegetative cells) where the long axis
of the spindle is placed at right angles to the long axis of
the cell.

Instances of these are the spindles of the daughter nuclei
in pollen-mother-cells, where the nuclei have probably a
reciprocal influence. Though the axes of the two spindles
are generally parallel and at right angles to the longest axis
of the cell, yet it sometimes occurs that the two spindles lie
in planes which are at right angles to each other. More
striking are those divisions in the cambium cells where the
axis of the spindle lies at right angles to the long axis of
the cell.

It seems probable that we must in many cases look out-
side the cell for the cause of the polarity, as is especially
indicated by the observations of Nemec (Vora, 18996,
p- 219) on cell-division in potato-tubers, where, in a cell
(the surrounding cells having died) the spindle formation
did not take place in ‘‘ monaxial’’ fashion, but cones were
formed on all sides.

THE ORIGIN OF THE CONES OF THE MULTIPOLAR
SPINDLE.

But few cytologists have investigated closely the origin
of the cones of the multipolar spindle. The first to devote
his special attention to this subject was Belajeff (1894), in
whose paper ‘‘Zur Kenntniss der Karyokinese bei den
Pflanzen’’ were published the results yielded by his obser-
vations on the division of the pollen-mother-cells of various
species of Larzx. Since then, in addition to Strasburger’s
observations, the spore-mother-cells of Eguzsetum (Oster-
hout, 1897), and the pollen-mother-cells of Coé@a (Lawson,

344 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

1898), Passzfora (Williams, 1898), Lzlium (Grégoire,
1899), Lavatera (Byxbee, 1900), Gladiolus ( Lawson,
1900), and now Cassza, have been studied more particu-
larly in this regard. The results of these studies are a
number of observations which agree in the main, though
in minor details they frequently differ.

It may therefore be of interest to compare the various
modes of formation and development of the cones of the
multipolar spindle in the spore- or pollen-mother-cells of
these plants.

About even the earliest stages the observations frequently
differ. In Larzx, Cobea, and Cassza the first stages ob-
served are described as showing a more or less radial
arrangement of the meshes of the cytoplasm (in Code@a in
the immediate neighborhood of the nucleus only). Similar
observations have been made on the pollen-mother-cells of
Lilium speciosum (Grégoire, 1899), and by myself on those
of Lilium Humboldit. In Hgutsetum and Gladiolus this
radial arrangement is not specially mentioned; but in regard
to Passifora and Lavatera it is distinctly stated that this
radial arrangement of the cytoplasmic meshes does not
appear until an elongation of the meshes parallel to the
nuclear wall has taken place. In all plants mentioned (with
the exception of Cobea) this parallel elongation of the cyto-
plasmic fibers has been observed and followed with great
care, especially in ZLavatera. Here the manner of the
formation of the meshes is identical with that observed in
Cassia, except perhaps, that in Cassza the fibrous zone in
its first stages is not so complete. But sooner or later the
number of rows of parallel elongated meshes increases to
such an extent that the felt-like mass is easily discernible.
At this stage it is figured by a number of authors. Stras-
burger (1888) figured and mentioned it; but Belajeff (1894)
was the first to describe this felt-like zone: ‘‘ Die erste
Veranderung, welche ich bei in Theilung begriffenen
Zellen habe beobachten kénnen, war die Bildung einer den
Kern umhiillenden, dichten, filzartigen Schicht, welche auf
den ersten Blick als eine concentrisch um den Kern gewun-

Bot.—VOot. II.] HUS—CASSTA TOMENTOSA L. 345

denener Fadenknauel erscheint. Eine nahere Untersuch-
ung feinster Schnitte zeigt jedoch, dass diese Filzschicht
aus der Kernwandung parallel in die Lange gezogenen
Schlingen (Maschen) besteht.”’

This description would fit the felt-like zone occurring in
the pollen-mother-cells of all the plants just mentioned; but
Belajeff’s words, ‘‘ Die obenbeschrieben Umgruppirung der
Plasmafaden ist, wie es scheint, die Folge ihrer Contraction
und des Zusammenziehens der Schlingen (Maschen) um
den Zellkern,’’ do not agree with the views of others in
regard to this matter. Both Byxbee and Williams are of
the opinion that the meshes become elongated because of a
drawing-out. This then would be caused, not by a con-
traction of cytoplasmic threads, but by their expansion.
What causes this expansion is not yet explained, but an
increase in size of the nucleus is not improbable, and how-
ever slight, would produce the appearance described.
Observations on Cassa tend to confirm this supposition.
But whatever the manner of its origin, a weft-like zone
seems to be formed around the nucleus in nearly all cases.

The manner in which the fibers of cytoplasmic origin
form cones seems to differ in the various plants observed.
In this connection it is necessary to call attention to the
fibers which may be seen in the cytoplasm at the time of the
formation of the fibrous zone. In Cassa these fibers appear
when the first changes in the cytoplasm immediately adjoining
the nucleus begin to take place. At various points the
presence of granular fibers, which stain a deep violet and
which are sometimes arranged in conical groups, be-
comes apparent. They remind one of the ‘irregular,
deeply staining strands’’ of Passzfora ( Williams, 1899, p.191)
or of the cones of Hguzsetum (Osterhout, 1897, p. 161, fig.
4). In Cassza these fibers apparently establish a connection
with the fibers of the fibrous zone, which latter gradually
become parallel, or nearly so, to the thicker granular fibers
running at a greater or lesser angle to the nuclear wall.
The origin of the cones is apparently determined by the
thicker granular fibers. No drawing-out of the cones takes

place.
(2) April 15, 1904.

346 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

In regard to the granular zone, which is so prominent in
Cassza, little remains to be said. This zone has been de-
scribed and figured for several plants such as Hemerocallis
(Juel,1897), Larix, Eguisetum, Lilium, Passiflora, Lavatera,
Cobea, Gladiolus, Podophyllum and Helleborus (the last two
by Mottier, 18972) but always in reproductive cells at the
time of division, and not in vegetative cells at the same
stage. The partial disappearance of the granules during
the formation of the fibrous zone seems to indicate that they
are an accumulation of material ready for immediate use.
The large quantity present in reproductive cells which
divide twice in rapid succession points to the same thing.
And though the granular zone is present even in the last
stages of the second division, though not always so promi-
nently (Lawson, 1898, p. 178), and in fact after the pollen-
grains have separated (Byxbee 1900, fig. 25), it must be
remembered that a large quantity of food material is needed
within a comparatively short time of activity, not only by
the pollen-mother-cell but also by the pollen-grain. As
Byxbee (1900, p. 72) points out, the manner in which the
granular zone accumulates, suggests the gathering of deuto-
plasm in animal eggs (Wilson, 1896, p. 115 e¢ seq.).

SUMMARY.

t. The cytoplasm of the young pollen-mother-cell is
made up of a network of more or less radially arranged
fibers, upon and between which larger and smaller granules
are found (fig. I).

2. The spindle is formed as follows:

a. The meshes adjacent to the nuclear wall be-
come smaller, and elongated parallel to the
nuclear wall (fig. 2).

b. A granular zone accumulates around the
nucleus. In the cytoplasm, especially in the
peripheral part, appear deeply-staining rough
fibers, frequently arranged in conical groups
(fig. 3).

Bot.—Vot. II.] HUS—CASSIA TOMENTOSA L. 347

c. The fibers forming elongated meshes around
the nucleus become smooth (fig. 3).

d. A felt-like zone is formed partially or com-
pletely surrounding the nucleus; granular
linin threads appear within the nucleus; the
granular zone now takes up about one-half the
remaining space (fig. 4.)

e. The deeply-staining rough fibers of the cyto-
plasm, united into cones, establish connection
with those of the felt-like zone (fig. 5).

f. The linin threads of the nucleus become more
prominent and finally smooth. They run
parallel to each other and to the axis of the
-cone which has the ascendency over the
others.

g. As soon as the threads of a cone become
entirely smooth, the nuclear wall breaks down
at the base and the linin and kinoplasmic fibers
anastomose. The fibers become grouped
into bundles (fig. 7).

h. A multipolar spindle is formed, two cones of
which, situated opposite each other, are more
prominent than the rest. Sometimes but two
cones are present (fig. 8). The two promi-
nent cones finally absorb the others, thus
forming a bipolar spindle.

3. The spindle formation for the second division takes
place in the manner described for the first, but is even more
pronouncedly multipolar diarchal.

4. The spindles of the daughter nuclei sometimes lie in
planes which are sometimes at right angles, sometimes par-
allel to each other.

s. The spindle formation in Casséa tomentosa L. forms
a connecting link between the multipolar polyarchal spindle
Anlage ordinarily met with in dividing pollen- , spore- , and
embryosac-mother-cells, and the multipolar diarchal spindle-
Anlage described for vegetative cells.

In conclusion I beg to acknowledge my great indebted-

348 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

ness to Dr. W. J. V. Osterhout, at whose suggestion the
work was undertaken, and under whose direction it was for
the greater part carried out.

Berkeley, California, and

Amsterdam, Holland, October, Igor.

Bor.—Vot. II.] HUS—CASSTA TOMENTOSA L. 349

1894.

1900.

1899.

18908.

1897-

1898.

1900.

1897a.

18970.

1899a.

1899).

1897.
1896.

1888.
1900.

1884.
1887.
1896.

1899.

BIBLIOGRAPHY.

BELAJEFF, W. Zur Kenntniss der Karyokinese bei den Pflanzen.
Flora, Bd. LXXIX, p. 430.

ByxBEE, E. S. The Development of the Karyokinetic Spindle in
the Pollen-Mother-Cells of Lavatera. Proc. Calif. Acad. Sci.,
3d Ser. (Bot.), Vol. II, No. 2.

Gr&GOIRE, V. Les cinéses polliniques chez les Liliacées. Za
Cellule, T. XVI, 2¢ fasc., p. 235.

Hor, A. C. Histologische Studien an Vegetationspunkten. of.
Centralblatt, Bd. LXXVI, p. 65.

Jue, H. O. Die Kerntheilungen in den Pollenmutterzellen von
Hemerocallis fulva, und die bei denselben auftretenden Unregel-
massigkeiten. /ahrb. f. wiss. Bot., Bd. XXX, p. 205.

Lawson, A. A. Some Observations on the Development of the
Karyokinetic Spindle in the Pollen-Mother-Cells of Cobza scan-
dens Cav. Proc. Calif. Acad. Sci., 3d Ser. (Bot.), Vol. I, p. 169.

Origin of the Cones of the Multipolar Spindle in Gladiolus.
Bot. Gazette, Vol. XXX, p. 145.

Mortiger, D. M. Beitrage zur Kenntniss der Kerntheilung in den
Pollenmutterzellen einiger Dicotylen und Monocotylen. /ahré.

f. wiss. Bot., Bd. XXX, p. 169.

Ueber das Verhalten der Kerne bei die Entwickelung des
Embryosacks und die Vorgange bei der Befruchtung. /ahkré. /.
wiss. Bot., Bd. XXXI.

Nemec, B. Ueber die karyokinetischen Kerntheilung in der Wur-
zelspitze von Allium Cepa. /ahrb. f. wiss. Bot., Bd. XXXIII,
Pp: 313-

Ueber Kern und Zelltheilung beiSolanum tuberosum. Fora,
Bd. LXXXVI, p. 214.

OsTERHOuT, W. J. V. Ueber Entstehung der karyokinetischen
Spindel bei Equisetum. Jahrb. f. wiss. Bot., Bd. XXX, p. 159.

Rosen, F. Beitrage zur Kenntniss der Pflanzenzelle. Cohn’s
Beitraége zur Biologie der Pflanzen, Bd. VII, p. 249.

STRASBURGER, E. Ueber Kern und Zelltheilung. Jena.

Ueber Reduktionstheilung, Spindelbildung, Centrosomen und
Cilienbildner im Pflanzenreich. Jena.

TANGL. Zur Lehre von der Kontinuitat des Protoplasma. Sztzber.
d.k. k. Akad. d. Wiss., Bd. XC.

WENT, F. A. F. C. Beobachtungen tiber Kern-und Zelltheilung.
Ber. d.d. Bot. Gesellsch., Bd. I], Heft 7, p. 247.

Witson, E. B. The cell in development and inheritance. New
York.

Wituiams, C. L. The Origin of the Karyokinetic Spindle in Passi-
flora coerulea Linn. roc. Calif. Acad. Sct., 3d Ser. (Bot.),
Voli aie ps 130:

Sees

CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER.

EXPLANATION OF PLATE XXX.

All figures were drawn with the Abbé camera lucida. Objective, Zeiss
Hom. Imm. ;3;; Comp. ocular, No. 6.

Fig.

Fig.
Fig.

Fig.
Fig.
Fig.

I.

nor

A young pollen-mother-cell. The cytoplasm consists of elon-
gated meshes.

Change in cytoplasm. The meshes of the layer immediately
adjacent to the nucleus become smaller. Beginning of accu-
mulation of granular matter around the nucleus.

Formation of violet-staining fibers in conical groups in the cyto-
plasm. The fibers parallel to the nuclear wall have become
smooth.

A dense granular zone has accumulated around the nucleus.

Granular fibers have appeared in the granular zone.

Formation of cones. More or less parallel linin threads are seen
in the nucleus.

_

> A

i

"E XAK.

LAT

ius] P

Ly
it

[

R Bor Voult.

ata

ord

je
pale

Roc. CALARAD

Pp

352 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

EXPLANATION OF PLATE XXxXI.

Fig. 7. The nuclear wall breaks down. The fibers are grouped in
bundles.

Fig. 8. Nucleus with two especially prominent cones.

Figs. 9-10. Multipolar spindles; these occur but rarely.

Fig. 11. Indication of bipolar spindle.

Fig. 12. Perfect bipolar spindle.

cS

ee -
LF pt he Diy OY n
~

-
7
’

of
on

te Oe gate tens

a

Pie, a)

~ 7 Ps A aay ; Mf
rire a hs)

a55

Fig.
Fig.

Fig.
Fig.
Fig.
Fig.

13.
14.

15.
16.
17.
18.

CALIFORNIA ACADEMY OF SCIENCES. [PROC. 3D SER.

EXPLANATION OF PLATE XXXII.

Daughter chromosomes proceeding toward the poles.

Daughter chromosomes at poles. Several small, red-staining
bodies present in granular zone and cytoplasm.

The daughter nuclei are surrounded by membranes.

Felt-like zone around daughter nuclei.

Multipolar diarchal spindle of daughter nuclei.

Bipolar spindle of daughter nuclei.

<a ry =

1. Scr. 3? Ser Bor Von It.

INDEX TO VOLUME II.

New names in heavy-faced type; synonyms in #talics.

AGAVE... . . 264, 265, 267, 271, 272,
@inericana . 302 46 =~ » « « 200;
Aglaophyllumruthenicum .....
EXISTE is Sweeter oe 6 “OUOROeCeObe Oin
Aster ascendens yosemitanus....
durbrowi......
YOSemitanus. 2.6 = - = =. =< «
BACILLUS RADICICOLA.......-.
BOERYVEIS!:;. <1 oes @ es 0 «6 85 =
BI StiIATISIA (ore sc) co) =) 6) = lel es

CALOCHORTUS . 107, 108, 110, 112, 116,
134, 147, 149

albus 107, 109, 111, 112, 117, 118, 119,
BMaAGMISN. |. 0-0. ci 112, 118,
TprpsiiGne 6 4.0 oe, 6 omIPA abe
DOH ENNG 455 86 oo Ono c 113,
RUG US raphe eed oe shreds of oss 1518 8 115,
eMmthattiece ce licks if) 109, 113,
GLETEHSY ) ajreniehs oF o) oh, 2) os is 113,
cataline.... ..- - 115, 142; 143,
CIAVALUSH = en. eeu lel ole) ors 114, 134,
RTHEP SS Olnn OE Ole CMD EOmEEIO 123,
RONCOLOL ER eee eae -s ess 14;
(obyvbrbebtilib, Sete ono on Goce cen 116,
elegans... .. 111, 113, 121, 122,

TGINEPEAS "Golan 5 GSR ae ORD

HEME Boho Go atame 6 113,

SHDCLAUALYS ac os jelel ise) es ete
CXCAVALCUS Ha Ni -i sites lemons) 7s). 115,

HEXHOSUS raiment aio) ns el;

greenei.. 2... = « . «114; 128,
gunnisoni 7 2 5. se. = 114,
iy galkins BiG oo oo oe 114, 127;
AAVETUISENSIc o8e) ile) Pe) LED, 1455,
eaves hats ons io Oe6 aes 115, 135,
PETC ELIIDAL sopra pie fr apees cu eterna rs 116,
TIIRSTIICS oe SP ene eceace ceicerc en

veya set by ighc Bea Be ducws ac
longebarbatus . . . 114, 127, 129,

luteus . 110, 115, 135, 137, 139, 140,
citrinus.. . 109, 110, 115, 138,
141
COMLOLOM Yo) eetrahake! iste faitieass 114,
oculatus . . 109, 110, 115, 138,
FODUSEA: o)<.<h @ Se eve 115, 139,
Iku te CG. Oe Ob Ga ceo eS 2

273
274

22
342
293
292
293

298
108
100

129

[355]

Calochortus macrocarpus. .. . 116,
maweanus.. . 109, 113, 120, 121,

major. -) .:-

roseus.....

rn eas 113,
crab ih tie’ 113,

MILICUSpomeedesuiis tots LOT 114k 26;
BITE S55 oka A 113, 122, 123, 125,
nuttallii 107, 116, 135, 145, 146, 148,
OpispOeHSis se) ees ee cee etlOOs
OVHMENATS 6) G bale eto mn toe
PANEL s) fle - 115,
pavonaceus... 114, 129,
Het 64 6 Fla olsvo co 6 oe
plunimerce 7 -) ieee 109, 114,
pulchellus . . . 109, 111, 112, 118,
Milo 4 6 Oo 4 Da Se 113, 126,
purpurascens sme emia: en cite diel ie
(AUG! 50 a0s Oy De oo la Cc - 139,
Shastensis:.< .)< 9s). 2... = 113,
splendens 115, 116, 143, 144, 145, 146,

AtEOVIOIACEAN =) «eee eie bos otis

PEN oa co & oho CUORH IES,

montanuts . 115, 143, 144, 145,

HMPA ele ecismieetomrem lilo.
Ose gang 6 omae 113, 126,
umbellatus..... 113, 123, 124,
LUTTILOTHS ses) =) ol ne 113, 124,

venustus.. 108, 109, 110, 115, 132,
139, 140, 141, 142, 147, 148

Eldorado 25) --) 2 115;1395
purpurascens. . 115, 139, 140,
TODUSE TA (Hi aes ost ee ant
TOSGCUSE-) iret 115, 139,
sulphuretus ... . .115, 141,
VeStal a ives «ssl « 109, 115, 139, 140,
wallaceitiir.. 5) 1 100<) =) «

WEEC Dia Pry Wiel eh oy clich cite
abi. go ao

obispoénsis. ..... .114,
purpurascens...... . 114,
VESEUSic oc) ele) cient is tic oma 114,
CASSIAM AN cobclp'clefpainic) 3! 1otte 344, 345,
tomentosa . 329, 333, 338, 339, 341,
347
CAGsitios Gian wo do eos os
Castilleia brooksil.)..)-) 2). s)-8 ene
Cistichaerc cree arene ances
sbi ee A 5s Gee Gg oon

spheres S. Rah oO eos orn OI ORC

nana,. -

356 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER.

Castilieia patlidal yo. sys 2 cites
PAT VINOLS ot ehe: eee
Steras Elia yoy ae Sion ey ail Se
CASIIIEIOIDES mee ise oan cee oe
Chetomorphalinum.........
Ghrysabotiy are cs) ole ia he teas
Cobzea 69, 70, 71, 72, 272, 342, 343, 344,
Contes fee eee on one 83,875
Convolvulus berryi.......
LORICHICUIIE Sc ecko eee Coes
villosus.....

Sel erim inl tet foo 7 o

WEELERN Se fer Fee ene oe eee
Cuscntals- is ers 2
Cyeadacere Fes oreo ere ats iti
PO ee OY BU of iL Te

revoluta..
Cyclobothraalba....

FYE atte 18 an aera cael Me be

PUICTPH A a) ic) = to 33

Diploderma. .
INDLISSEIEMIM ofa oo Ue fa) i= oe IS
RPSL SETI ais gs ah
EP PLCIE = FeV Woh SPE egal ee eer

PU EILALIEIG © 5, a > Were a vio. FS ws Hee
LENUISSIMMUM.. » ss
VAFIOLOTUIE cea ee a, aie ve

EQUISETUM . . 69, 70, 71, 278, 274, 343,
345, 346
Brioronum tindum).. > pe. 52. -
SCAPIS CEU 5 ce civ odsm te
Hucalochortus . +550 sce) Gs) x, 2 112,
POMS 87> oy Sota Fs SMa wie a he

GARRVA FREMONTYE ..355) 5 260.

pati. cares aero cae.
Gigartina..... el ckeWo Wojeem sees)
Gilia sparaiflora:.....°.°5 5 2: 291,

WAL SAA Kee wea rte ee, oh Ee
Gladiolus: 3 sews, (0: See ee 342, 344,
Gomimopitylnt cose. 25 obs ees

DUAN Swe 35 <3 eet aye ee
Gracilaria confervoides .......
Gratelonpia filicinay. = 2s 22
Grossulatiays/.) isisct ses cee es ee

TIRE ISBOR US! sioy. cslc, eo ute Neueiae ere 71;
Helminthocladia purpurea... ..
Premerocallisiiyo< ts. age ee orin mas
Hosackia subpinnata.........
Hymenenalatissima ....... 2, 16,

ERIS'*SOUALENS\. ove "s)s> otis. ee ose

290

LARIX . . 69, 70, 71, 267, 273, 343, 344,
Lavatera. . . . 63, 69, 70, 71, 72; 272,
344, 346
MICHNS eb e k ete. seers :
UNSWICMIATA, 5 lass uel eons nae
ES TIMIN OSA ors on ts ils note 2 « « 296,
Leitnera foridana =|; -). e600
Piliacew aoe sige ota se ca. wee eae
Ea laeisr Ff: Ac rretae tee Ue 272, 342, 344,
humboldsi. ...°.- 2. 2) cle aes
SPECiOSUIM <6. . pons hey cy sleet
ETI S oat ok oe see
micranthus bicolor... ....
tivularis latifolins). . 2 .< . sem
Madaria corymbosa. ...-+..2<-s
Madia corymbosa’. = ...)5 se = siee
hispidaic vo. si. 2 =) sas
villosa.. of ot noone perenne
Medicago denticulata.. - 98, 295,
SAbivalsiois Sty) eso eee 295,
Melilotus parviflora. ..... as
Minulus bioletti. <. . -). cise =
PaAlmetiycn.s - o-/- coil eae ee
NEOTTTA (NIDUS-A VIS» 2 15)'. us oe
Wereccystis).'<. << 66a <) se oode
lutkeana;..: 2.5 «=~ 177, 01945
Nenroplossatmt |. =). 4.) ane eee 4,
QNAEYSONTANUM .. 2 es we ew oe 4,
lobultferv um cs oe = 2 sw 4, 39,
Nitophyllum .. 2, 3, 4, 5, 7, 8, 12, 14,
25, 33, 207
andersonianum . 3, 4, 7, 8, 9, 10,
13, 15, 32, 33, 34, 43
ARDEFSORE foc) ors Oe eee
QPENALUIE 6 jac) Sai a eee 3,

corallinarum. .8, 9, 11, 12, 16,
25, 26, 43, 44

JAFLOTRARYIN Wo = oe) eA OA,
fissum) (2,2 2s) ee eee =a
flabelligerum...... 3, 4, 30, 34,
fryeanum . . 2, 3, 4, 9, 10, 11, 12,

14, 16, 21, 22, 23, 24, 43
harveyanum.. . 8, 12, 13, 15, 28,
30, 31, 37, 43
Hise Foy. ons aE Vay karen
TACEF LUI ee aes 2; 3; 145
latissimum . . 2, 3, 4, 7, 8, 9, 11,
13, 14, 16, 17, 18, 19, 20, 32, 43
macroglossum.......- 4, 16, 19,
marginatum. .
multilobum.. . 3, 4, 8, 9, 11, 12,
15, 23, 27, 28, 29, 30, 43
(Neuroglossum) andersontt.. . 3,

ruprechtianum . 3, 4, 8 9, 11, 12,
14, 16, 30, 31, 34, 35, 36, 37, 38,
41, 43

PUtHeMwiCw Mp. -.) <yake Ae eee rit

spectabile . . 3, 4, 9, 11, 12, 13, 14,
21, 22, 23, 43

346
342

23

BoT.—VOL. II.]

Nitophyllum stenoglossum.. . .4, 39,
uncinatum . . 3, 4, 11, 12, 15, 16,
27, 43
violaceum . . 2, 3, 4, 8, 10, 11, 12,
14, 16, 36, 37, 39, 40, 41, 43
Sormum crispulum.... 1...
Niyimipized albarisite es ie) «le oss
ORTHOCAREUSH am ehelmiciclne ero ios
Oscillatoriaung sue mew ment? ota i

PASSIFLORA .. 69, 70, 71, 72, 342, 344,
346
Phaceliaicircinatay. saci. +. e: « leh one
HEMOLALIS ee Ponca icici cel store
stimulans....
Phoradenaron. we) els) is! 100, 101,
Phyllospadix . . 15, 27, 174, 176, 179,
194, 207, 212, 213, 214, 233

es Se: ie es, 8 eo

Sonal otniduquaviohoatcss bec
Podopiiy lami <) ijee. ie ol ee ee ils
Polygonum douglasii........

xd e Rents iaited eels, vous encuee

Porphyra .173, 174, 175, 177, 178, 180,
183, 184, 185, 186, 187, 188, 192,
195, 197, 200, 210, 212, 213, 214,
219, 220, 221,. 224, 228, 229, 230,
abyssicola . 174, 182, 183, 184, 185,
192, 193, 194, 195, 196, 219, 223,
BUTLGUMY SECAR ie amin tsy nes eiitcleicl te
amplissima.. .
196, 205, 215, 216, 221, 222, 224
atropurpurea
RAEI PAS aca Seo NE
laciniata . . 174, 175, 182, 183, 189,
193, 194, 195, 196, 197, 198, 199,
208, 230
WH biliealis se
leucosticta . 174, 175, 180, 184, 190,
192, 193, 195, 198, 199, 200, 201,
207, 220, 228
WWMEATISM Ney icp ite teria eee)! fs
- 174, 175, 184, 188, 192,

= ewe Hele lta 6) he tela

miniata .
195, 197, 218, 219, 220, 221, 222,
cuneiformis! . 5.5 2: 194, 196,
LERUESSIINA TOM elle) all's) a¥ rel icles ie

naiadum . . 174, 175, 176, 177, 180,
182, 183, 192, 194, 195, 212, 213,
223, 231, 232, 233

nereocystis...
191, 194, 195, 210, 211, 212, 229,

occidentalis’.). <2. 2 5 195, 196,

perforata . . 175, 176, 177, 178, 179,
183, 191, 194, 195, 197, 200, 202,
204, 205, 206, 208, 209, 211, 212,
230, 231
lanceolata... 175, 177, 191, 194,
208, 210
segregata .-

INDEX. 357
41 Forphyra purpurea, oc si Ae 174
26 tenuissima . . .174, 175, 183, 184, 188
192, 193, 195, 196, 219, 220, 221, 222

13 224
variegata ..177, 183, 186, 187, 194, 195

39 196, 225, 226, 227, 228, 231

342 DUILATIS a Wale) te) 174, 196, 202, 204, 212
Prionitisy, ( oc:/c.taarecel Che one 8
290 Janceolata’’...; 4, Mise, ah ee 8
298 Pyropia california .......... 210
345 AERLESTAN. ha) sgtiask cue eu cine: Gian 100
Rhizobium leguminosarum. ... . 298
291 RIDES Hea se toes ts Slye vied hee 241, 249
291 AMALIE eypakiey i) ish ie ur cen aateptcees 251
291 GUD SUE IAG ta a SR tas he) BO 251
305 AULOUN TT Ai ei cace he, nae 250
180 cate rely aaah) Pearl Pa Opaes Come Se ne 2 ve 250
ascendens)..)5.0 2 2 ae 244, 249
212 Jaspers. aoe aN ieee 244
346 AMECH IN es ital oi)s ich toes Te acre 249
286- Pinominatitm::) is). ce) @\ eel 25H
286 Dracteosumin 20s Li eel te 249
182 DLrandeged asso ee 242, 249
194 CORSE ts fe joy, site fe) 6 te bathers! Wace DAS
216 ealifornieunis sie a. eth ae 246, 250
231 Ciliosim 4 geen etre en eee 250
188 cognatum... . 251
224 CHUSH LUM etch emsumet eve iioieais 250
174 Givaricatim Seles be eee
195 CORINALHI es es se sy rate ee
ery throcatpuiny..\ jo ls) =) eee 250
199 Suchstoides.. . ead
213 elancescens 1-202 anes 245, 249
192 SIUEMOSU ME 851) se ele enero ee 249
204 SLACUE 5) Ves. cline teas 251
HeESpertina cies sacs sects eineae 251
199 hittellianum .... 245, 249
191 hudsonianuittts<. st) ne) eile eee 250
205 Uh jeeb <5. asd OO pane a oer th bet)
indecorum) 29. -)-). . « 243, 244, 249
174 LACUSEKG yrs s yccel a) aie) a cet forte are 251
193 molle 251
224 haSiasubhirtmy-cs) tai 7., cas yieenat ier etc 251
218 Jeb oGitorn ibs: aMmine Peel ao. feo, ote Le 250
220 leptan thymic i- ts, lye ieh ence 251
181 BFACKYARTRUIO. se ode even te 251
214 TODDIlE eset oat Boba ch rant)
malvaceum’. =) 5. 3) sas 241, 244, 249
214 diatiposanuim ~ 2 6 2 «= seme 50
213 PMATSHAUT eioey ol sey es oy sees 250
187 menziesii. - 241, 248, 250
230 MIMSTACOLLUIM | 5 2a se) a) st oie 250
228 WORLAMUUL os (6.6 We, 1s ea, sy POLS 251
182 prQoyelrerhiiilonem Gg Oko oO clone 251
203 nevadense . . . 241, 244, 245, 246, 249
216 niveum.. ova Awol O55 251
OCCId en FALE Sa ses uie sls eess far as 250
195 oligacanthum........ 246, 250
oxyacanthoides..... eS
207 ceri brers alias a Seo e anal gtade ates 249

wv

\A)

2714

358

Ribes prostratum .-. <2). +. ss ss
quercetorim: 9. *) aie) 2h) =e
roezli.. .
sanguineum..

vartegatum .
SQ@XOSUIMN« » «© © © 2
Sctipuanii~ i. a bey 2 242,
SeTICCTHAN 2 <u +=. 31)-

VITIGESCENS i522, >, 3470 te) he
spethianum
speciosum
SIAZTNEREUIESS 5. « Aibeki es pas joe eke
subvestitum ...
tenuiflorum.....
velutinum
viburnifolium
WACOLLIS.. 5 fo. = = os
OINOSHME sonia = ks bee
VISCOSISSISLTT 5) =o). aa ce 5
WwatsOmianuiis: <<< 2 + 2% 2.
wilsonianum....

Rabesiieals eee e ee "|

ROUSONIAN te. = = 3 un

Te a EC ee BC

SEQUOIA SEMPERVIRENS ... . .- 83,
Solanum arizonicum.... . . 164,
COLSOTIACH ME > «le hid 160, 161,
PEMESTONMES Frm. i ieg ayo wa) a 160; 161,
tenuilobatum .. - - 164, 165;

ad

a

CALIFORNIA ACADEMY OF SCIENCES.

250
251
251
249
249
251
249
250
247
251
251
251
250
249
251
249
250
251
249
251
250
249
251

87
165
172
171
171

[Proc. 3D SER.

Solanum umbelliferum 159, 160, 162, 163,
164, 168, 171, 172

californicum.. - - 164, 172
wallace@o.) -. bos s.c 163, 164, 166, 168
viridis - 164, 166, 168
xanti. . 160, 162, 163, 164, 166, 167, 168
glabrescens.. . . . 164, 168, 169
intermedium...... 164, 168
t QUUIECEE co a. Se hae - 160,461, 166
Streptanthus gracilis..... 285
ULVA... ae ely és:
atropurphrea ..). . . . «sus 202
VACIHIAIEA so) = 3.6 Ba s: ORE te 196
IRPRELA «Renee ae oa 2 Se 218
purpureaig— -. =)<.--> sj See 218
VAISCTAS.-. 5S... het mes. 3, toy cee 342
WISCHIMN cee eke nee - 100, 101, 305
Wildemania......-. ss tlc eos
amplissIma .. «++. 215
PHS, oe a so heya) ce Royant a) Pa 196
TATRA BTS oe 68s, = Sota 218
Per foTasa ee sas == <=) ol oa 202
LONUESSIING Ma 2 we ew tw es 220
VOPLELAUL a tee tele 6 6) © oo mes 225

ZOSTERA. . . . 176, 180, 182, 194, 207, 213

214, 223, 224

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QK California Academy of
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C35 Proceedings. 3d ser.:

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