CONTENTS
SMITH, CHARLES PIPER. Studies in the genus ah cpio The Mic-
rocarpi, exclusive of Lupinus densiflorus I
Harper, RoLAND M. The — ee of newbies: iste Michigan :
and its environment - 23
TENopYR, LILLIAN A. On the constancy of a shape i in ea of varying
shape 51
HARPER, Shikinn T. Two seacieaihe Discomycetes hie ae - 77
Stet, W. N. Studies of some new cases of apogamy in ferns (plates 4,5) - 93
Harris, J. ARTHUR, and Avery, B. T. Correlation of morphological varia-
tions in the seedling of Phesestus vulgaris - 109
Rock, JosEpH F. New species of Hawaiian slate (plate 6) - - - 133
ARTHUR, JosEPpH CHARLES. New species of Uredineae—X = - - - 141
Humpert, E. P. A striking variation in Silene satiate - - ~ 157
SMITH, aoe Preer. Studies in the eae ciple al eat den-
ifloru - 167
PETRY, pene C. Studies i in the eogeritian of New York pO ues - 203
ON Makoto. A carrier of the mosaic disease (plate sl - - 219
ANS, ALEXANDER W. The air chambers of Grimaldia fragran - 235
Macleans, VAUGHAN. The strand flora of the Hawaiian. Archipelago
—I. Geographical relations, origin, and composition- - 2
Berry, EpwarpD W. Notes on the fern genus Clathropteris - - - 279
Donce, B. C. Studies in the genus Gymnospora — Bo 2 on
cultures made in 1915 and 1916 (plate 8) - - - 287
WEATHERWAX, PauL. The evolution of maize - 309
Boas, HELENE M. The relationship between the number of sporophylls
and the number of stamens and pistils—a criticis
CouTANT, Mary WoTHERSPOON. Wound oan in ge cacti (plate
9 =. g00
ici! Py os P. The ions nad flowering ‘elite of Nantucket XIX 365
Brown, ELIZABETH Wuist. Regeneration in Phegopter. a - 391
FERRIS, ROXANA STINCHFIELD. Sharan: and a of Adenos
(plates 10- O-I2) -
CoKxer, DorortHy. Revision of the North Aan gras a iucilerie
(plates 13, 14) - se eae
GrarF, Paut W. Philippine Basidiomyeetes—II (plate rs) 51
PENNELL, Francis W. Notes on plants of the southern United States—IV 477
nesgeenertgge VAUGHAN. The sors flora of the Hawaiian Archipelago
—II. Ecological relations - 483
INDEX TO AMERICAN BOTANICAL LITERATU - 43,
87, 121, 159, pet 253, 301, pes 385, 425, 471, 503
INDEX TO VOLUME 45 - - - - - - - - - - SIL
1V DATES OF PUBLICATION
Dates of Publication
No. 1, for January. Pages I-50. Issued February 8, 1918°
No. 2, for February. 51-92. March 7, 1918.
No. 3, for March. 93-132. April 1, 1918.
No. 4, for April. 133-166, May 15, 1918.
No. 5, for May. 167-218. May 23, 1918.
No. 6, for June. 219-258. June 20, 1918.
No. 7, for July. 259-308. August 9, 1918.
No. 8, for August. 309-352. September 9, 1918.
No. 9, for September. 353-390. September 20, 1918.
No. 10, for October. 391-432. October 19, 1918.
No. 11, for November. 433-476. November 15, 1918.
No. 12, for December. 477-519. December 23, 1918.
Errata s
Page 241, line 10, for “‘created”’ read “ — es
Page 271, line 8, for ‘‘ Cyas”’ read ** C-
Page 383, line 3, for ‘‘ Law or’’ read “‘ eae on.’
TORREY BOTANICAL
aaa
FouNDED BY WILLIAM HENRY LEGGETT 1870
EDITOR
ALEXANDER WILLIAM EVANS
ba a
Jean Broapuurst
Vol. 45 No. 1
BULLETIN
OF THE
TORREY BOTANICAL CLUB
ee ee
JANUARY, 1918
Studies in the genus Lupinus—lI. The Microcarpi, exclusive of
Lupinus densiflorus
CHARLES PIPER SMITH
(WITH SIXTEEN TEXT FIGURES)
INTRODUCTION
‘The subgenus Platycarpos of Watson readily permits separation
into two groups, the Pusilli of Heller (Muhlenbergia 8: 87.
1912), or the loosely-flowered small species of the interior of west-
ern North America; and the Microcarpi (Bull. Torrey Club 44:
405. 1917), or the verticillate and commonly larger plants of the
Pacific Slope of both North America and central Chile.
The published names applied to members of the boc cd i
are as follows:
microcarpus Sims, Bot. Mag. 50: Pi 2413. 1823.
densiflorus Benth. Trans. Hort. Soc. II. 1: 409. 1835.
Menzies Agardh, Syn. Gen. Lup. 2. 1835.
palustris Kell. Proc. Cal. Acad. Sci. 5:16. 1873.
Menzies aurea Kell. ibid. 5:16. 1873.
lacteus Kell. ibid. 5: 37. 1873.
luteolus Kell. 1bid. 5: 38. 1873.
Bridges Gray; Watson, Proc. Am. Acad. 8: 538. 1873.
malacophyllus Greene, Pittonia 1: 215. 1888
ruber Heller, Muhlenbergia 2:75. 1905.
arenicola Heller, ibid. 2:75. 1905.
horizontalis Heller, ibid. 2: 74. 1905.
glareosus Elmer, Bot. Gaz. 39: 53. 1905.
. subvexus C. P. Smith, Bull. j Ses Club 44: 405. 1917.
{The BULLETIN for December (44: 535-579. pl 25, 26) was issued December 27, 1917.]
Pe orhrerrr ren
2 SMITH: STUDIES IN THE GENUS LUPINUS
L. Bridgesui Gray is evidently the same as L. luteolus Kell. and
is so acknowledged by Watson (Bib. Index 238). L. arenicola has
been referred by Heller (Muhlenbergia 6: 70) to L. lacteus Kell.,
which, as noted by Heller, was referred by Watson first to L.
densiflorus and later to L. microcarpus. L. malacophyllus Greene
might easily be considered as a connecting link between the
Pusilli and the Microcarpi. Since so many variations occur, it
is really surprising that no more names have been published.
Several specimens have been marked “var.” by the collector or
student; but evidently no one has cared to attempt a critical study
of the group. Except for the material easily determined as L.
luteolus Kell., most specimens have been labeled either L. micro-
carpus or L. densiflorus, or left unnamed. -
During four years of field observations in middle western
California, my efforts to determine the variations I collected led
to unsatisfactory results, and a survey of the material found in the
herbaria visited in 1908 brought me to the determination to some
day seek a classification that might possibly eliminate some of the
evident confusion as to proper identifications. With this object
in mind, I recently undertook to look into the status of the forms
already described, and then to work out the relationships and char-
acters of the seemingly undescribed forms known to me.
MATERIAL EXAMINED
The National, Stanford University, and University of Cali-
fornia herbaria kindly placed many sheets in my hands for study.
Two weeks were spent in Cambridge, New York, and Philadelphia,
where the material in the herbaria in those cities was given due
attention. Thus some 225 sheets have been considered in forming
the opinions put forth in this paper.
GEOGRAPHICAL DISTRIBUTION
This group is peculiar to the west coast of America, the range
generally ascribed to the initial species, L. microcarpus Sims, being
the range of the entire group. In South America, the plants seem
to be confined to central Chile, having an altitudinal distribution
from the coast far up into the Andes. In North America the
longitudinal range is from Vancouver Island to Lower California;
SMITH: STUDIES IN THE GENUS LUPINUS 3
Vancouver and the adjacent islands have one isolated form, the
Yakima Valley in Washington another. This latter, one local
form of central Oregon, and one of western Nevada are the only
ones peculiar to the country east of the Cascade-Sierra ranges.
The more widely distributed form northward is apparently com-
mon both to the waterways in Oregon east of the Cascades and
also to southwestern Oregon and the northern counties of Cali-
fornia. Central California is evidently the center of distribution
of the group, and there many specialized and localized forms occur,
from the coast well up into the Sierras.
DIAGNOSTIC CHARACTERS
Most of the characters ascribed by Sims and Bentham to their
species are common to all members of the group, and cannot be
used to distinguish between the two species. Agardh’s treatment
of the species was mainly based upon the specimens seen by him
in Lindley’s herbarium. A small-flowered plant collected by
Douglas in California, as well as all the Chilean material there, he
referred to Sims’s species. A more robust and longer-haired plant
he accepted as Bentham’'s L. densiflorus, and this he acknowledged
as having the aspect and nearly all the characters of L. microcarpus,
sed robustior et bina Supe tits oabitie in Saher floribus obsitus. Ceterum
petioli longiores, videntur. Unica autem dif-
ferentia, quae ad speciem distinguendam adele videatur, e bracteis sumenda est.
The yellow-flowered, short-pubescent form with long racemes
of distinct verticels was set apart as his L. Menziesii. Three
certain forms collected by myself in California were very readily
determined as these three species of Agardh; but when I con-
sidered other forms found by me, the original plates of L. micro-
carpus and L. densiflorus, and also the conflicting determinations
and opinions of various botanists, it seemed logical to conclude
that the group needed special attention.
Upon taking up the study, I soon found that reliable diagnostic
characters are not as readily selected as I expected them to be.
Many which at first recognition appeared to be all-sufficient for
clear-cut distinctions finally suffered complete or partial abandon-
ment after being carefully weighed. As a rule, the general aspect
will appeal to the eye first and the general structure of the vege-
4 SMITH: STUDIES IN THE GENUS LUPINUS
tative parts of the plant merits reasonably close attention. Evi-
dently the environment, and especially competition with plants of
other genera, has much to do with whether the plant is low or tall.
Presumably the amount of available plant food may determine
- whether the plant is acaulescent and unbranched, or more or less’
caulescent and branched. The axial terminal bud, unless injured
by some external cause, may be depended upon to produce a
fruitful peduncle. Branching is, of course, merely the result
of the development of axillary buds, and it is common, in large
well-branched plants, to find matured or ripened seeds in the pods
of this axial peduncle when the racemes of at least some of the
branches are still in bloom. A low unbranched acaulescent plant
is a record of its own environment, and a dry barren hillside or
desert area the habitat in which it is able to both exist and persist.
On the other hand, a tall plant, branched more or less above the
base perhaps, almost always comes from an environment also
favorable to plants of other genera; and abundance of nourishment,
assisted by the stimulation of competition, is fundamentally re-
sponsible for the extensive vegetative growth developed. The
“‘fistulous”’ character, especially, is certainly the evidence of a
rapid growth during a warm moist season. I see no “‘specific’”’
characters in the stem structures of these plants, excepting, to
some extent, in L. luteolus Kell.; but I have admitted into my
treatment of varieties more than one definite reference to such
stem characters, especially when coupled with other characters or
supported by geographical distribution. The length of the pet-
ioles and the size of the leaflets and stipules vary with the stem
portions and seem to me to be of no value for specific distinctions.
Even the pubescence failed to give satisfaction in my effort to
draw specific lines.
Most reliable of all, I have concluded, is the position of the
flowers in anthesis and soon after anthesis. This may not always
be strictly constant in all the flowers of a particular raceme, nor
in all the racemes of a certain plant; but I am convinced that, with
occasional allowance here for some little variation, this character
will prove to be as positive a one as can be found in the Micro-
carpi, and certainly is more usable than the very indefinite dis-
tinctions as yet proposed by those who have treated any of the
SMITH: STUDIES IN THE GENUS LUPINUS 5
group. The verticels are regularly crowded in L. luteolus and
L. horizontalis, usually well separated in L. microcarpus and
L. subvexus; but in L. densiflorus is found a full range of variations
from densely crowded to quite remote. The floral bracts vary
much in length, usually becoming reflex-withering as the flower-
buds open; but in one variety of L. subvexus they are rather con-
spicuous for their tardiness in reflexing. Their length seems to be
too inconstant to be of much diagnostic value.
With few exceptions, I have not been able to accept size and
color of petals for specific distinctions, though such are abundantly
used in the varietal characterizations. At the free edges above,
the keel petals are usually ciliate, often densely so, and a tendency
to be ciliate at the free edges below is more or less pronounced in a
few of the forms herein designated as varieties; but in L. luteolus
this ciliation below is as dense and prominent as that above,
usually extends backward onto the claws, and seems to be worthy
of special notice. Variation in size, shape, apex and base of
banner; ciliation, length, and width, but not detailed form, of
wings; size, shape, and curvature of keel—are all worthy of con-
sideration in the delineation of varieties. ‘The pubescence of the
calyx usually agrees rather closely with that of the peduncles,
petioles, etc. The lower lip, I am persuaded, yields good varietal
characters as to length, dentation, and inflation; but the upper
lip, at least in many cases, varies too much within the flowers of a
given raceme to be trusted for any distinctions. The presence or
absence of bracteoles is usually a decided uncertainty; but their
presence may be diagnostic in L. horizontalis and in Elmer’s L.
glareosus.
The size, relative thitkness, surface, color, and marking of the
seeds—when more of same have been collected and properly identi-
fied—will probably be found of value in the varietal, if not in the
specific, classification; but in L. luteolus again, I find a seed char-
acter seemingly distinctive and peculiar to that species. The
mature pods show much variation in size, but otherwise give little
evidence of deserving much consideration.
PATHOLOGICAL MATERIAL
Aboud specimens are occasional, and care should be exer-
cised to avoid them in the designation of varieties. Field work
6 SMITH: STUDIES IN THE GENUS LUPINUS
may sometimes be necessary in order to establish that a certain
variation is really the result of a diseased condition. Usually,
however, an abnormal development is indicated by pronounced
elongation or multiplication of some of the floral parts. One
very interesting case is that represented by sheet 366288 in the
U.S. National Herbarium (K. Whited 536, Ellensburg, Washington).
In both specimens the flowers dissected have three wing-petals
each, and in one all, the wing- and keel-petals are very abnormal
in shape (Fic. 1). In the case of sheet 620193 (V. Bailey gz,
NC ye
es
1 cm.
Fic. 1. Lupinus SUBVEXUS FLUVIATILIS C. P. Smith. K. Whited 536 (US
366288). 1. From specimen on right, 6, banner; w, wing ale like this); k, keel; c,
calyx as seen from above; side view of entire blossom above. . From specimen on
left: w, wing (two like this); ew, extra wing petal; k, keel; no fee present.
Toppenish, Washington) the flowers are of the same narrow form
and seemingly fruitful.
SPECIAL ABBREVIATIONS
The location of each specimen cited is indicated by an abbrevia-
tion. The abbreviations used are as follows:
DS, Dudley Herbarium of Stanford University;
UC, herbarium of the University of California;
ColU, herbarium of Columbia University;
T, Torrey Herbarium (at New York Botanical Garden);
NY, herbarium of the New York Botanical Garden;
G, Gray Herbarium of Harvard University;
US, United States National Herbarium
PA, Philadelphia Academy of Arts and ‘Sciences:
_ CPS, private herbarium of the writer.
SMITH: STUDIES IN THE GENUS LUPINUS r
ILLUSTRATIONS AND MEASUREMENTS
The figures are mere outline representations of the dissected
floral parts, mounted in water under cover-glass slips. The calyx
is represented as seen from above, the upper lip having been
pressed down to show its proportional relations to the lower lip.
By this method, the interstitial bracteoles, if present, are not apt
to be overlooked. One wing-petal is often added merely to show
the extent or absence of the ciliation. The banner is flattened out
sufficiently to get the measurements and show the shape of the
apex and claw. The measurements, with necessary exception in
a very few cases, were made from these dissected floral parts after
same had been softened in boiling water and mounted for drawing.
Key to the species of the Microcarpi
Bracts subulate throughout, mostly stiff and not reflex-wither-
ing, about 5-6 mm. long, little more than twice the length
pedicels; flowers 9-11 mm. long. 1. L. malacophyllus.
Bracts thant ladcoolake with a dilated thin base, mostly reflex-
withering as the flower-buds open, at least the lower much
longer than the pedicels; flowers 11-19 mm. long.
Flowers ascending to suberect in anthesis.
Banner 4-6 mm. wide; wings naked below and often
above; calyx ebracteolate.
Banner 7-11 mm. wide; wings and keel ciliate near the
claw both below and above; calyx bracteolate. 3. L. horizontalis.
Flowers spreading in anthesis.
lowers soon becoming suberect after anthesis, not
to
. L. microcar pus.
secund. 4. L. subvexus.
Flowers spreading or —— secund after anthesis. i
Plants 30-90 cm. tall, pend fibrous, often widely
branched above; banner ovate; keel ensely ciliate
at base both below vie shoes . aes crowded;
seeds uniformly blackish brown, regularly tuber-
] 5. L. luteolus.
rarely ovate; keel usually naked near base below;
but sometimes sparsely ciliate; verticels crowded
or remote; seeds smooth or irregularly roughened. 6. L. densiflorus. nai
1. LupINUS MALACOPHYLLUS Greene, Pittonia 1: 215. 1888.
I quote from the original description as follows:
al, erect, a span high, with a few ascending branches from the base; soft
throughout, with a long white villous pubescence .. . . racemes verticillate, at
di tai ONE NEN a RET
* L. densiflorus and its varieties will be di 1 in the third paper of this series.
8 SMITH: STUDIES IN THE GENUS LUPINUS
least below, .’. calyx-lips very unequal; the upper short, scarious, slightly notched;
lower . " dotinetle 3-toothed: corolla 1% inch long, _ blue and dark purple;
keel widikiegiaty falcate, naked . . . seeds orbicular, whit
Dry hills, near Verdi, wipvacie: 2 May, TB85; RENAE yo Mr. C.F. Sonne :s2.-.
related to the nen small-flowered L. pusillus Lane L. brevicaulis, but with .
showy verticillate racemes of large flowers, . .. .
This species might well be viewed as a connecting link between
the Pusilli and the Microcarpi. Its possession of verticillate
flowers is my only excuse for referring to it in this paper. Forms
of L. subvexus transmontanus, of northern California and Oregon,
are the only specimens of the Microcarpi that have already been,
and may yet be expected to be, confused with L. malacophyllus.
Seeds in my collection, collected by Heller, from about Reno, are
from pale flesh-color to dark reddish.
NeEvADA. Without definite locality, C. L. Anderson (UC
170306). Ormsby County: Carson City, 1865, C. L. Anderson
(UC). Washoe County: along track below Verdi, 16 May, 1897,
C. F. Sonne (UC); same locality, June, 1889 (UC); Reno, 13 May,
1901, 7. W. Cowgill (UC); seed only, road between Laughten
Springs and Reno, 26 July, 1911, K. Brandegee (UC).
2a. LUPINUS MICROCARPUS Sims, Bot. sia 50: pl. 2473. ‘1823.
[Fics. 2, 3.]
The original description is as follows:
Lupinus microcarpus; foliis digitatis, calycibus verticillatis in ane Pap AS
labio rate emarginato inferiore bifido ter breviore, leguminibus rhombeis hir-
sutis disperm
Descr ea branched. Leaves aaa cuiated 9-10, lanceolate, hairy on
the under surface, smooth on the petioles twice the length of the
leaflets, pubescent. Stipules ae ney go ncle terminal. Flowers blue, in a
verticillate ane six-flow red. Bracts small, hairy. Calyx Wace late
( which are asta from the calyxes, are to be called the appendices)
bilabiate; Mie pat much the shortest, emarginate; lower lip bifid. aun oblong.
g the vexillum arina monopetalous, sharp-pointed. Anthers 10,
five Pasi pe five hac. Style the length of the stamens. Stigma capitate.
Legume small, rhomb-shaped, mucronate by the persistent style, hairy; seeds two,
variegated with black lines and dots.
This species of Lupin is a native of Chili. It has not, we believe, been heretofore
described; and differs from all the known species by its small two-seeded We
regret however that we did not receive these in time enough to be added to me en-
graving.
- Raised from seeds Pe John Walker, Esq., of Arnos Grove. It flowered in April,
. and appears to be ann
SMITH: STUDIES IN THE GENUS LUPINUS 9
Authors generally, following Agardh, seem to have assumed
that any and all Chilean specimens are necessarily L. microcarpus.
Sims. One of my early hypotheses was that the Chilean plant
ought to be specifically distinct from all the
North American forms referred to this species
of Sims. As I now see it, however, neither
of these assumptions is altogether in agree-
ment with the facts. A new viewpoint is
evidently necessary and the conclusions finally
accepted by me are: (1) that the strongest
specific character possessed by L. microcarpus
has never been properly recognized; (2) that
most of the North American forms regularly,
or occasionally, referred here belong rather to
two other species; and (3) that both of these
“two other species’’ are likewise represented
in Chile. This viewpoint, as I see it, permits
a much more satisfactory classification of the
forms under consideration.
The original description serves little more
than to indicate the group, of which this spe-
cies was the first named and described. The Fic. 2. Luprnus
illustration (from which Fic. 2 was copied) M'CROC*? US Sims.
f Copied from _ original
shows quite plainly, however, that the flow- plate.
ers must have been ascending to suberect in
anthesis, a character that, as stated above, my studies have led
me to accept as of real diagnostic value. This character, I am
persuaded, will be found to persist under cultivation, at least
for a few generations. I find, moreover, amongst the very limited
array of Chilean material available for my study, specimens which
show this character and which certainly, it seems to me, represent
much better the form used for the original illustration than the
other Chilean specimens referred elsewhere by myself, or the
North American specimens referred here by others. It is very
probable that the species, as here limited, is a composite of more
than one recognizable form; but I await the opportunity to study
new and additional material before attempting to do more with
the South American varieties.
10 SMITH: STUDIES IN THE GENUS LUPINUS
CHILE. Vicinity of Santiago, springy hillsides, 4500 ft., near
Rio Colorado, 21 Jan., 1902, G. T. Hastings 468; without
definite locality, C. Gay, ex Herb. Mus. Paris (G); Coquimbo,
July—Aug., 1856, W. H. Harvey (G).
Fic. 3 represents the gan char-
acters of Hastings’ No. 468, as re-
corded above. The specimen may
-be briefly described as follows: about
15cm. tall, branches and foliage con-
gested; villous, branched near the
base and rebranched above; leaflets
about 10-15 mm. long, petioles 50-70
Lom mm. long; flowers ascending to sub-
QU
erect in anthesis; verticels 2-3; leg-
Se a umes 13-15 mm. long, seeds about
Fic. 3. Lupinus microcarpus 3 * 4 ™m., straw-colored, unmarked,
Sims. G. T. Hastings 468 (UC. perhaps immature.
65806). The species is ciate in
North America by the following variety:
2b. Lupinus microcarpus ruber (Heller) comb. nov. [Fic. 4.]
Lupinus ruber Heller, Muhlenbergia 2:73. 1905.
From Heller I quote:
Branched from the base, the branches diffuse . . . leaflets blue-green ...
smooth above . . . : flowers erect in from 2 to 4 whorls, . . . dull red, 1 cm. long,
about 4 mm. across lower calyx lobe 8 mm. long,
barely 3 mm. wide at the base, the apex cleft for 2 mm.,
with a short cusp in the sinus; upper lip practically ob- Sv
i ai
solete, represented by two short lanceolate teeth of 1 si
ban —
mm.:-banner almost plane, ovate-lanceolate, 4 mm.
wide at base, slightly keeled; wi ngs narrow, 2 mm. q
wide, the lower edges not meeting until near the apex; pees :
keel not strongly curved, barely 2 mm. wide at the
mee bearded only near the slightly narrowed base. . I
type is no 7827, collected May 5, 1905, at
deans Kern County, oo along the railroad Bias.
a short distance west of t town. is a species
remarkable for its small narrow red flowers. Some of Fic LUPINUS MICR
the specimens show a short central flowering CARPUS RUBER (Heller) C.
branch much shorter than the leaves, while others have P. Brandegee
in addition lateral branches about equalling the leaves. (UC 149863).
hat
blue”’ flowers, the parts all but the keel differing in shape, and that is glabrous, while
in ours it is bearded near the
SMITH: STUDIES IN THE GENUS LUPINUS 11
The calyx is mostly 7-9 mm. long, the teeth of the lower lip
are about 1.5 mm. long, commonly slender and curved-diverging,
and the interstitial ‘‘cusp’’ is conspicuous. The flowers vary
from dark red to merely pinkish. The seeds are yellowish gray
to flesh-color, about 3.5 mm.x 3 mm., flattish, coarsely rugose,
unmarked or nearly so. JL. brevicaulis, with which Heller com-
pares the plant, is a member of the Pusillt.
CALIFORNIA. Kern County: Tehachapi, 5 May, 1905 A. A.
Heller 7827 (US, PA, NY, G); Tehachapi, June, 1911, K. Bran-
degee (UC). San Diego County: Jacumba Hot Springs, near
Monument No. 233, 26 May, 1894, L. Schoemfledt, U. S. & Mex.
Int. Bound. Comm. No. 3307 (US); same locality, 29 May, 1894,
E. A. Mearns, U. S. & Mex. Int. Bound. Comm. No. 3336 (US);
Banner, 25 Mar., 1901, J. S. Brandegee (UC); Jacumba, 2500 ft.,
9 July, 1916, E. A. McGregor 88 (CPS). San Bernardino County:
Mesas, Argus Mountains, Apr.—Sept., 1897, C. A. Purpus 5437
(US, G, UC). ?San Luis Obispo County: La Panza, 26 May,
1888, Mrs. Mathews (UC
3a. LUPINUS HORIZONTALIS Heller, Muhlenbergia 2: 74. 1905.
[Fic. 5.]
I quote from the original description, as follows:
Branches several, all floriferous ... the lowest ones hor-
izontal, the ones above them more ascending, the middle
ones erect . .. lower lip of calyx green, pubescent with
somewhat ae, hairs, ovate-lanceolate, 6 mm. long, over 3
mm. wide at base, barely acutish, the apex minutely 2-tooth-
,
ed; upper os ovate, membranous, barely 2 mm. long, cleft
nearly to the base, the lobes lanceolate with rather wide () Sie
sinus. villous and ciliate: flowers pale violet-blue, whorled roe ne
and crowded . erect, I cm, long, 7 mm. across; banner
plane or only slightly turned back, with a prominent mid-
vein; wings . 2 mm. wide keel not much curved,
ciliate on the ‘ie third . . . seeds nearly 3 mm. across
marked with small dark spots.
I accept this as a well-marked species. It is oo.
probably peculiar to the region of the type local- oxiontaris Heller-
ity, though represented southeastward by adis- A. ‘ean 7725 (G).
tinct variety.
CaLIFoRNIA. Kern County: Sunset, 20 Apr., 1905, A. A.
Heller 7725 (US, PA, G, NY).
We
12 SMITH: STUDIES IN THE GENUS LUPINUS
3b. Lupinus horizontalis platypetalus var. nov. [Fic. 6.]
A L. horizontali differt partibus omnibus florum majoribus
latioribusque; vexillo 15 x 11 mm., alis 15 x 7 mm., carina 12 x 4
prope 3.5 X 3 mm. crassis.
All the floral parts are
much larger and broader
than in the typical form of
the species, and the outer
branches are apparently
not prostrate. The seeds
are of a buff ground, mar-
bled with purplish brown.
CALIFORNIA. San Ber-
Fic. 6. LUpiINUS HORIZONTALIS PLATYPET- :
Hee €. built. Boll & Chosdier ds60. coc, NaTdina. County: cabterd
127206). base of Fremont’s Peak,
Mojave Desert, 6 May,
1906, Hall & Chandler 6860 (TypE, UC 127296); Mojave Desert,
May, 1882, S. B. & W. F.Parish 1271 (G). Kern County: near
Randsburg, May, 1913, K. Brandegee (UC).
Platypetalus is Greek for ‘‘ broad petal.”
4a. LUPINUS SUBVEXUS C. P.
Smith, Bull. Torrey Club
44: 405. 1917. [Fic. 7.] U
The following is a part of
the original description: Serer
lous, the hairs 2-4 mm. long; whorls
3-7, well- separated; flowers spread- ee
ing in anthesis, evidently ascending
to suberect later; calyx ebracteolate,
quite villous below; pods not secund.
The typical plant has the lower lip of
the calyx evidently inflated (subsac-
cate) near the base, large flowers with
much of the banner and wings dark
purple, and the banner rounded apic-
ally. Most of the varieties have the
lower calyx-lip scarcely or not at all inflated and smaller flowers with the banner
gradually narrowed to an acute apex.
i cm.
Leen ene TEED
Fic. 7. Lrprnus susvexus C. P. Smith.
Heller & Brown 5415 (DS 9586).
SMITH: STUDIES IN THE GENUS LUPINUS 13
CALIFORNIA. Yolo County: Madison, Heller & Brown 5415
(DS, US, PA, NY, G). Contra Costa County: Antioch, 3 May,
1894, A. Eastwood (G, UC); Antioch, 8 Apr., 1895, A. Eastwood
(US); Clayton, 17 Apr., 1889, Chesnut & Drew (UC). Alameda
County: Oakland Hills, 1866, H. N. Bolander 100 (G); Alameda,
T. S. Brandegee (DS 9589); Livermore Pass, May, 1898, J. B. Davy
(UC); Cedar Mountain, May, 1903, A. D. E. Elmer 4371 (US,
NY). San Francisco County: San Francisco, Apr., 1893, I. Tide-
strom (UC).
Most, if not all, of these specimens are labeled L. microcarpus
Sims, though the type sheet bears a correction, by A. A. H.,
changing the determination to L. densiflorus Benth.
Subvexus, ‘‘sloping upward,” refers to the position assumed by
the flowers upon withering.
Key to the varieties of Lupinus subvexus
Calyx 9-11 mm. long.
Banner truncate, rounded, or obscurely angled at apex,
usually 7-8 mm. wide
Bracts normally reflex-withering as the flower-buds
open.
Flowers 14-17 mm. long; calyx 10-11 mm. long;
leaves not blackening in drying. 4a. typical subvexus.
Flowers about 14 mm. long; calyx 9 mm. long;
leaves blackening in drying. 4b. var. nigrescens.
hering 4c. var. fluviatilis.
Bracts tar dily or with
Banner evidently angled at apex, 5—7 mm. wide.
Wings usually non-ciliate; keel naked below.
Verticels approximate; plants 8-18 cm. tall, bushy. 4d. var. transmontanus.
Verticels remote; plants 20—40 cm. tall, with stems
it hes el ted and fistulous bel 4e. var. Leibergit.
Wings strongly ciliate near base above; keel ciliate
4f. var. insularis.
Calyx 7-8 mm. long; banner 5-6 mm. wide, angled at apex.
Leaflets 20-40 mm. long, 7-10 mm. wide, green; banner
Verticels distinct; keel straight. 4g. var. phoeniceus.
Verticels crowded; keel evidently arcuate. 4i. var. Wilkesit.
Leaflets 10-20 mm. long, 4-6 mm. wide, conspicuously
white woolly-villous; banner more slender, 12-14 mm.
ts 4h. var. albilanatus.
4b. Lupinus subvexus nigrescens var. nov. [Fic. 8.]
A L. subvexo typico differt foliolis nigrescentibus cum siccantes;
pilis brevioribus inconspicuuis; floribus paulo minoribus; labio
inferiore calycis vix inflato; alis superne ad basin ciliatis.
14 SMITH: STUDIES IN THE GENUS LUPINUS °
Differs from the typical form in that the leaflets turn black in
drying, hair-covering less conspicuous, flowers smaller, lower lip
of the calyx but slightly subsaccate, and the
wings ciliate above near the base.
CALIFORNIA. Ventura County: Griffins,
Mt. Pinos, July, 1902, A. D. E. Elmer 4006
(Type, US 466205; type-duplicates, DS, NY).
Kern County: hills, near the summit of Tejon
Pass, on desert slope, 27 May, 1914, S. B.
Parish 9256 (UC).
This variety merits special study in the
field, with collection of considerably more
material. It is very different from Elmer’s
L. glareosus, taken at the same locality, but
lem shares with that variety the peculiar char-
Fic. 8. Luprnus acter of the leaflets turning black in drying.
SUBVEXUS NIGRESCENS The specimens are labeled L. microcarpus.
C. P. Smith. A. D.
Whew joes eG 4c. Lupinus subvexus fluviatilis var. nov.
[Fig. 9.]
Eramosus vel ramosus ad basin ramis suberectis vel panden-
tibus, 10-25 cm. altus, conspicue villosus; foliis paulo congestis
(NOE
Fic. 9. Lupinus SUBVEXUS FLUVIATILISC. P, Smith. Fritillaria Club (US 205559).
cum planta humilis eramosaque est, plus patentbus cum planta
ramosa est; pilis 1-1.5 mm. longis; petiolis 4-10 cm. longis, vil-
losis, pilis saepe paulo plus 2 mm. longis; foliolis 5-10, 15-25 mm.
longis, saepe obovatis vel spatulatis nonnumquam oblanceolatis,
3-5, plus minusve appositis; floris ad anthesin pandentibus, postea
-ascendentibus vel suberectis, 14-16 mm. longis; pedicellis paulo
plus 1 mm. longis, plurimum gracilibus; bracteis verticilli humil-
limi calyce prope aequantibus longitudine, tarde vel nequaquam
SMITH: STUDIES IN THE GENUS LUPINUS 15
reflectentibus et marescentibus, laxe villosis vel sublaevibus:
calyce ebracteolato 9-10 mm. longo, subter dense villoso_pilis
prope 2 mm. longis, labio superiore diverso 2-dentato, prope 2 mm.
longo, inferiore subrecto vel ad basin paulo inflato 3 mm. lato
2-dentato, sinu plerumque sine vestigio; eee pallido-rosea vel-
purpurea; vexillo 12-14 mm. longo, 7-8 mm. lato, apice paulo
contracto, rotundato vel truncato, plus Scere ungue 2—4 mm.
lato; alis 11-14 mm. longis, 5-7 mm. latis, ad basin non ciliatis;
carina 9-11 mm. longa, subrecta: leguminibus usitatibus prope 14
x 8 mm.; seminibus pallidis maculatis, 4.5 x 4 mm.
WASHINGTON. Without definite locality, 1889, G. R. Vasey 250
(Typr, US 296664; type-duplicates, G, NY). Yakima County:
Yakima region, 1882, T. S. Brandegee 42 (UC); North Yakima,
Oct., 1885, T. S. Brandegee (UC); Fritillaria Club, North Yakima,
1890, (US); Toppenish, 17 July, 1897, V. Bailey 91 (US); Yakima,
3 June, 1898, A. B. Leckenby (US); North Yakima, 29 May, 1899,
J. B. Flett (US); Wenas, 1 June, 1902, Griffith & Cotton 85 (US,
NY). Kittitas County: Ellensburg, 25 June, 1897, K. Whited
536 (US); Ellensburg, June, 1897, A. D. E. Elmer 371 (US, NY).
Seemingly peculiar to the valley of the Yakima River in central
Washington. The flowers are relatively large with broad petals.
Perhaps the strongest individual character is that the floral bracts
are tardily or not at all reflex-withering as the flower-buds open.
The seeds are flesh-color, mottled with a pale reddish brown, the
hilum of the darker color. Fic. 1 illustrates abnormal specimens
of this variety.
Fluviatilis is Latin for
4d. Lupinus subvexus transmontanus var. nov. [Fic. 10.|
A var. fluviatili differt foliolis oblanceolatis plerumque acutis;
calyce 9-11 mm. longo, sinu labii inferioris plerumque dentis
vestigio instructo; vexillo apice angulato, 11-12 mm. longo, prope
6 mm. lato ad basin paullatim contracto.
Similar to var. fluviatilis, but the leaflets are usually acute at
the apex, the floral bracts usually reflex and wither as the flower-
buds open, the lower lip of the calyx has the vestigial median tooth,
and the banner is evidently acute at the apex. .
OREGON. Wasco County: Antelope, May, 1885, 7. Howell
(Type, US 20844; type-duplicate, PA). County not given: Currant
Creek, 11 May, 1885, 7. Howell 361 (G); John Day River, Crown
Rock, 19 June, 1896, V. Bailey 65 (US). Malheur County:
tay
of a river,’’ and refers to the habitat.
16 SMITH: STUDIES IN THE GENUS LUPINUS
Malheur River, June, 1883, Cusick 1113 (G). Jackson County:
Antelope Creek, near Eagle, 4 June, 1898, E. I. Applegate 2388 (US).
CALIFORNIA. Modoc County: meadow
bank along stream, 28 July, 1893, Milo S.
Baker (UC); shore of Goose Lake, Aug., 1895,
Mrs. R. M. Austin (US); sagebrush at Tule
Lake, 31 May, 1897, E. I. Applegate 879 (US);
Goose Lake, July, 1898, Mrs. R. M. Austin &
_ Bruce 154 (UC). Lassen County: Craigs,
1893, M. S. Baker (UC); Madeline Plains,
June, 1898, Mrs. R. M. Austin & Bruce 2145
: (DS, NY, UC). Siskiyou County: Klama-
ae thon, 2 July, 1903, E. B. Copeland, Baker
Distribution 3537 (US, NY, G); Klamath
Hills, 13 May, 1909, G. D. Butler 704 (UC);
Fic. 10. Lupmus Klamath Hills, 21 May, 1910, G. D. Butler
cisnipapen ee 1373 (US, UC).
‘Gowell (IS. 208aa), Transmontanus is Latin for ‘‘across the
mountains’’ and refers to the fact that the
form is found on both sides of the Cascade ranges. It has been
determined as L. densiflorus, L. microcarpus, and L. malacophyllus.
4e. Lupinus subvexus Leibergii var. nov. [Fic. 11.]
A var. transmontano differt
altitudine 25-40 cm.; caule fis- —
tuloso ramoso aliquantum super
gatis: verticillis 4—5, distantibus;
floribus suberectis mox post an-~
ri: Epp oie 2s SUBVEXUS LEI-
mith. J. B. Leiberg 317
vel prope acuminato, ungue 4 mm. lato basi; alis 11 mm. longis;
carina 8 mm. longa, reeta.
Unlike the other northern races of L. subvexus, this variety is
branched from well above the base and has the lower parts dis-
tinctly fistulous. The verticils are remote, the banner is almost
acuminate at the apex and its claw is poorly defined, and the
SMITH: STUDIES IN THE GENUS LUPINUS 17.
flowers are evidently far from being showy. It is probably very
local and rare and should be carefully studied in the field.
OREGON. Crook County: near Prineville, 955 m. alt., 1894,
J. B. Letberg 317 (Type, US 291134; type-duplicates, G, US
291133).
4f. . Lupinus subvexus insularis var. nov. [Fic. 12.]
Ad basin ramosus; foliis aliquantum congestis, foliolis gracil-
' ibus 2—3 cm. longis; racemis pluribus, floribus ad anthesin pandent-
ibus ascendentibus postea, bracteis reflectentibus, bracteis petiolis
pedicellisque laxe villosis; calyce dense villoso; carina curva.
Fic. 12. LupINUSSUBVEXUSINSULARISC. P.Smith. T. S. Brandegee (UC82003),.
Much branched near the base, foliage congested, leaflets slender
and 2—3 cm. long; racemes several with the flowers spreading in
anthesis but ascending soon afterwards, bracts reflexing as the
flower-buds open, the bracts, petioles and pedicels loosely villous;
calyx densely villous; keel distinctly curved.
CALIFORNIA. “Santa Cruz Island: Apr., 1888, T. S. Brandegee
(Type, UC 82003).
Mounted on the sheet with the above specimen are some small
plants (UC 82002) collected by Brandegee at San Telmo, Lower
California, 28 Apr., 1893. These may represent the variety
insularis, but should prove to be different. They are so small and
in such condition I prefer to attempt no definite classification of
them at this time.
4g. Lupinus subvexus phoeniceus var. nov. [FIG. 13.]
Eramosus vel ad basin laxe ramosus, ramis ramulosis laxe
breve-villosis: verticillis 2-5 vel plus, appositis vel distantibus;
floribus ad anthesin pandentibus, postea ascendentibus vel sube-
rectis,12 mm. longis; calyce ebracteolato, 7 mm. longo, subter paulo
18 SmitH: STUDIES IN THE GENUS LUPINUS
laxe villoso, pilis I-1.5 mm.:longis, paulo retrorsis, labio superiore
minus 2 mm. longo, inferiore lanceolato-oblongo, vix 3 mm. lato,
2-dentato dentibus 1 mm. longis parallelis plerumque dentis
vestigio instructo; corolla phoenicea vel pallidiora; vexillo 10-11
mm. longo, 6 mm. lato, apice angulato; alis 9 mm. longis ad basin
superne paulo ciliatis; carina 8 mm. longa, plerumque recta: legu-
OSA
13. LuprinuS SUBVEXUS PHOENICEUS C. P. Smith. A. A. Heller 8632
(US ee a
minibus 13 x7 mm.; seminibus planis angulatis, 5 mm. longis, 4
mm. latis, atro-cineraceis, minute nigromaculatis circum cicatricula
pallidioribus, superficie minute inaequaliterque rugoso simile
quartzo.
Simple or loosely branched from near the base, loosely short
villous; whorls usually two to five, crowded or distinct; flowers
spreading in anthesis, soon becoming suberect, about 12 mm. long;
calyx retrorse-villous below with hairs 1-1.5 mm. long, the lower
lip with a vestigial median tooth; petals reddish purple or paler,
the banner about 10 x 6 mm., angled at the apex; seeds 5 x 4 mm.,
somewhat flattened and angled, dark-gray, minutely dotted with
black, paler about the hilum, the entire surface minutely roughened,
resembling quartz.
CALIFORNIA. Santa Clara County: Mt. Hamilton road,
2300 ft. alt., ‘‘on a roadside bank near oak trees,”’ 31 May, 1907,
A. A. Heller 8652 (Typr, US 612641; type-duplicates, PA, NY,
G); seed only, type locality proper, 17 June, 1908, C. P. Smith
(CPS); tilled soil below Smith Creek bridge, 17 June, 1908, C. P.
Smith 1479; Calaveras Valley, May, 1914, K. Brandegee (UC
178195, except as to raceme in upper left hand corner).
This seemingly well-marked variety may be peculiar to the
Mt. Hamilton Range. The plants are low and mainly simple in
poor soil, but in richer and looser soil may be 2-3 dm. tall with a
spread of 3-4 dm. The Smith Creek specimens were in seed as
to the primary branches, while secondary branches, in the axils
of the primary branches, were in flower.
SMITH: STUDIES IN THE GENUS LUPINUS 19
Probably related to this, but certainly not typical, are the
following: Atascadero, San Luis Obispo County, 30 Apr., 1861,
W. H. Brewer 493 (US); Ojai and vicinity, Ventura County, 24
May, 1866, Peckham (US). Additional material from these lo-
calities would be welcome and worthy of careful study.
The type collection has been recorded as L. microcarpus by
Heller (Muhlenbergia 2: 294. 1907).
Phoeniceus is Latin for ‘‘ purple-red.”’
4h. Lupinus subvexus albilanatus var. nov. [Fic. 14.]
A var. phoeniceo differt foltis gracillimis acutis plurimum solum
10-20 mm. longis; foliis petiolis pedunculisque conspicue brevi-
lanato-villosis; vexillo 12-14 mm. longo, graciliore; seminibus
plurimum minoribus pallidioribus, immaculatis vel maculatis.
ee
ANS
wb eemee
Fic. 14. Lupinus SUBVEXUS ALBILANATUS C. P. Smith. K. Brandegee (UC 149886).
CALIFORNIA. San Luis Obispo County: Paso Robles, July,
1911, K. Brandegee (Typr, UC 149886; type-duplicates, UC
149884, 149885, and 149947). Monterey County: near Plas-
kett’s ranch, on road to Jolon, about seven miles from Kings City,
K. Brandegee (UC); Manfield’s ranch, Santa Lucia Mountains,
ten miles from Kings City, 1-12 May, 1897, A. Eastwood (G).
This variety is conspicuous for its whitish cast and the narrow-
ness of its banner. The seeds are variable in size and color, as
_ shown by Mrs. Brandegee’s collections. Those with sheet 149884
are about 5 x 4 mm., some milky- or bluish-white unmarked with
the hilum area flesh-color, while others are drab with an occa-
sional dark speck and a conspicuous bluish-white or pale area
about the scar. Those with sheet 149885 are 4x3 mm. with a
drab ground much speckled with black, the hilum area little or
conspicuously paler and unspotted. Those with sheet 125917
20 SMITH: STUDIES IN THE GENUS LUPINUS
(Monterey County) are 4x3 mm., heavily marbled with dark-
brown, the hilum area unmarked and pale. The variety is evi-
dently confined to the upper Salinas Valley.
Albilanatus is Latin for ‘“‘white-woolly,
of the general pubescence in this plant.
”
which is descriptive
41. See subvexus Wilkesii var. nov. [Fic. 15.]
A var. phoentceo differt verticillis con-
fertis et carina curva: caulo plus minusve
erecto, 20-30 cm. alto, gracili aliquantum
super basin ramoso, laxe villoso pilis 1
mm. longis; petiolis gracillimis 5-8 cm.
— longis; foliolis 7-9, oblanceolato- -spatu-
latis prope 15 mm. longis; pedunculis
5mm - folia aequantibus, laxe villosis; verticillis
Fic. 15. Lupinus susvexus Prope 8, inferioribus distantibus, super-
Wikes C.P. Smith. Capt. toribus confertis; floribus 9-10 mm.
Wilkes (US 20841). longis ad inttionin pandentibus mox
postea suberectis; pedicellis gracillimis;
bracteis prope 5 mm. longis mox reflectentibus; calyce 7-8 mm.
longo, labio inferiore 2-dentato, sinu dentis vestigio instructo:
legumina et semina non vidi.
CHILE. Valparaiso, Capt. Wilkes, U. S. Explor. Exped.
(Type, US 20841).
The specimen is labelled ‘“‘Z. microcarpus, var.,’’ but is so dif-
ferent from all other Chilean material seen by me that I think it
best to set it apart as a variety of L. subvexus. The flowers be-
come suberect soon after anthesis and have several characters in
common with var. phoeniceus, but this differs in the greater number
of whorls, more crowded above, in the very slender pedicels, and
in the curved keel.
5. Lupinus LutTeotus Kell. Proc. Cal. Acad. Sci. S: 36. 1874.
[Fic. 16.] |
Lupinus Bridgesii Gray; Watson, Proc. Am. Acad. 8: 538. 1873.
From Kellogg’s description I quote:
Stem 1-2 feet high, suffruticose, glabrous below, bark light peered sei satiny
y E g neei-
unio cone. ... Leaflets .. . silky above and ulone: . . . Flowers light yellow
- - . ina dense crowded spike 6 to 12 inches in length . a. tube scarious
dest and widely ibid 2-bracteolate . . . upper lip contend. acute, entire;
lower lip herbaceous, toned, Learenyd defiexed and eepeeaietig at the junction of
the scarious portion . . . We ted, glabrous, with scarcely
SMITH: STUDIES IN THE GENUS LUPINUS 21
a few hairs on the margin at the base; keel acute, villous on the margins above at the
lower third. .
Found on the Coast range of mountains, near Senal, Mendocino County, Cal.
1872.
From Watson I quote:
Stem 1-2 feet high . . . leaflets . . . pubescent both sides or somewhat smooth
above ... petals pale eliowe:
Sacramento Valley. Pistestan by Bridges (55), Bolander (6512), and Dr.
Kellogg
Jepson (Fl. West. Mid. Cal. 2d ed. 216. 1911) emphasizes
the ‘“‘herbaceous”’ character of the upper calyx-lip.
As shown by my drawings, I have made studies of dissected
flowers from specimens taken at. five different stations, repre-
senting the geographical range of the species. I also studied with
( : eras
Fic. 16. Lupinus LUTEOLUS Kell. By AX ees ye oo County (US
468711); 2. H. N. Bolander 6512 (US 321126); 3. F. P. Nutting, Contra Costa
County (UC 15918); 4. J. P. Tracy 3496, Humboldt Ried ie 542773); 5. M.W.
Gorman 411, Oregon (US 280475).
a lens several others of the specimens listed below. Only one
specimen evidently supports the claim for an herbaceous upper
lip, and just three sheets plainly show bracteoles present. The
banner is usually ovate, acute, with the claw distinct and stiffened.
The wings are sometimes strongly ciliate at the base below, as
well as above, and the keel seems to be generally as densely ciliate
below as above, the ciliation extending well out onto the slender
claw. The upper lip of the calyx is sometimes emarginate and
22 SMITH: STUDIES IN THE GENUS LUPINUS
the lower lip is sometimes. two-toothed. The seeds are about
4X 3.5 mm., uniformly dark brown, regularly and roughly tuber-
culate as to the specimens seen.
CALIFORNIA. County not given: Bridges 55 (T); 1866, H. N.
Bolander 6512 (US); head of Penn Creek, 5100 ft., Aug., 1875, J.
T. Rothrock 224 (US); near Cheswick, 30 July, 1899, J. B. Leiberg
4318 (US). Contra Costa County: Fish Ranch, 29 June, 1893,
F. P. Nutting (UC); Walnut Creek, 28 July, 1897, Mrs. T. J.
Maynard (UC). Lake County: Lakeport, 16 Aug., 1882, C. G.
Pringle (US); Allen’s Spring, 6 July, 1882, D. Cleveland (UC); be-
tween Potter Valley and Hullville, 21 July, 1902, A. A. Heller 5938
(US, CPS) ; Scott Valley, 21 Aug., 1905, J. P. Tracy 2372 (UC); Kel-
seyville, 2 July, 1911, K. Brandegee (UC); Kelseyville, July—Aug.,
1892, W. L. Jepson (UC); between Houghs and Bartlett Springs,
11 Aug., 1910, K. Brandegee (UC); Sanhedrin, June, 1894, Purpus
1132 (UC); Mt. Sanhedrin, 18 July, 1913, H. M. Hall 9514 (UC).
Mendocino County: Covelo, 20 July—3 Aug., 1897, V. K. Chesnut
543 (US); Cloverdale, Russian River, 9 July, 1902, A. A. Heller
5830 (US); Calito, 5 Aug., 1902, A. Eastwood (US). Humboldt
County: valley of Trinity River, near mouth of Willow Creek, 10
July, 1911, J. P. Tracy 3496 (US); valley of Van Duzen River,
opposite Buck Mountain, 27 June—30 July, 1908, J. P. Tracy 2606 ©
(US, UC); Kneeland Prairie, 26 July, 1912, J. P. Tracy 3871 (UC);
Eel River, July, 1894, Purpus 1206 (UC). Shasta County: near
Middle Creek station, 3 June, 1905, A. A. Heller 7951 (US).
Siskiyou County: Klamathon, 2 July, 1903, E. B. Copeland, Baker
Distribution 3535 (US); Yreka, 8 June, 1905, A. A. Heller 7990
(US); Hornbrook, July, 1887, K. Brandegee (UC); dry land near
Yreka, June, 1908, 1909, 1910, Butler 375, 910, 1447 (UC).
OREGON. Jackson County: Rogue River Valley, 14 July, 1887,
I. Howell (UC); Ash Creek, near Ashland, July, 1893, Mrs. R.
M. Austin (UC); Ash Creek, Rogue River Valley, July, 1893, Mrs.
Austin (UC); Medford, 25 Aug., 1897, Mrs. Austin 1651 (US);
Spouting Springs, thirteen miles south of Ashiand, 13 Aug., 1896,
M. W. Gorman 411 (US). County not given: Dead Indian road,
western slope Cascades, 12 Aug., 1902, W. C. Cusick 2945 (US).
COLLEGE Park,
MARYLAND
The plant population of northern lower Michigan and its environment
ROLAND M. HARPER
(WITH THREE TEXT FIGURES)
Introduction.—The northern third of the Lower Peninsula of
Michigan, an area of about 11,000 square miles, seems to differ
from all other parts of the United States sufficiently to be treated
as a distinct geographical region. Its most striking characteristic
is probably the prevailing sandiness of the soils. (In this particu-
lar, as well as in the abundance of lakes, swamps and bogs, it
reminds one strongly of Florida.) It differs further from the
Upper Peninsula, the nearest land tothe northward, in being warmer
and therefore in having less of the boreal conifer element in its
forests, and from the territory adjoining it on the south in being
colder and in having a somewhat different seasonal distribution of
rainfall (as will be pointed out farther on) and more swamps.
The southern boundary is very indefinite, but may be located
arbitrarily for statistical purposes at the parallel of latitude 44° 15’,
which crosses the state from near the mouth of the Manistee River
to the mouth of Saginaw Bay, and passes through or near Manistee,
Cadillac, and Tawas City. Only about one sixth of the area is
under cultivation, so that there is no lack of vegetation to study.
But as the lumbermen spoiled the looks of the country before
there were many botanists in Michigan, and nearly all the plants
happen to belong to widely distributed species, comparatively few
botanical explorers have investigated this region. Aside from
incidental references in Cowles’s well-known monograph on the
dunes of Lake Michigan (1899), Beal’s Michigan Flora (1904),
and a few local plant lists, the following seem to be about the only
easily accessible papers on this region that a phytogeographer
would need to consult. Some of them contain references to earlier
literature of some importance.. The arrangement is chronological.
V. M. Spalding. ‘‘The Plains” of Michigan. Am. Nat. 17: 249-259. 1883.
C.S. Sargent. (Forests of) Michigan. Tenth Census U.S.9:550-554. 1884.
23
i
24 HARPER: PLANT POPULATION OF MICHIGAN
C. Kedzie. The jack pine plains. Mich. Exp. Sta. Bull.37. 8 pp. 188
. M. Coulter. in Ge. al comparison of some typical swamp areas. Rep. Mo.
Bot. Gard. 15: 38-71. pl. 1-24. 1904. (See pp. 41-49 for Michigan
B. E. Livingston. gle: relation of soils to natural vegetation in EEA and
Crawford Counties, Michigan. Bot. Gaz. a 22-41, with map. Jan. 1905.
Also in Rep. Mich. Geol. Surv. 1903: 9-30. pl.
H.H. Rusby. Observations in economic seiciins at Oscoda, Michigan. Jour. N. Y.
ot sees Sept. 1906.
W. J. Geib. Soil survey of Wexford County, Michigan. Field Operations U. S.
Bur. Soils 1908: 10 51-10 066. 1911. (Advance copies, separately paged, dis-
tributed in October, 1909.)
S. Whitaker. A vacation in northern Michigan. Forest & Stream 77: 806-807.
Dec: 2,
Hu Max a ad. -using industries of Michigan. (State publication, no series or
number.) 101 pp.,2folded tables. Lansing, 1912. (The notes on this region are
mostly on pp. 7-10. See criticism in Proc. Soc. Am. Foresters 11: 350. 1916.
Frank Leverett. Surface geology and agricultural conditions of 8 southern penin-
sula of Michigan. (With a chapter on climate by C. F. Schneider.) Mich. Geol.
& Biol. Surv. Publ. 9 (Geol. Series 7). 144 pp., 15 plates saad 3 folded maps),
16 figs. 1912
F.C. Gates. The vegetation of the region in the ig tate of Douglas Lake, Cheboy-
gan County, Michigan, 1911. Rep. Mich. Acad. Sci. 14: 46-106, with 24 half-
tones on 17 plates. 1913.
H. A. Gleason & F. T. McFarland. The introduced vegetation in the vicinity of
Douglas Lake, Michigan. Bull. Torrey Club 41: 511-521. Oct. 1914.
az. Se Gates. The relation between evaporation and plant succession ina given area.
Am. Jour. Bot. 4: 161-178. f. 1-9 (including 5 half-tones). March, 1917. (Area
igan.)
H. A. Gleason. Some effects of excessive heat in northern Michigan. Torreya 17;
176-178. Oct. 1977.
The plant population statistics given below are based on notes
taken by the writer while occupying the post of research assistant
in botany at the Biological Station of the University of Michigan
in the summer of 1912. Most of the observations were made while
walking out in all directions from the station (which is on Douglas
Lake), as far as Cheboygan and Topinabee on the east and Pellston
on the west. Copious notes were also taken from trains on the
Michigan Central R. R. from Cheboygan to Mackinaw City, on
the Grand Rapids & Indiana Ry. from Cadillac and Mackinaw
City to Pellston, and on the Pere Marquette R. R. from Petoskey
southward to the limits of the region and beyond.
Geology and soils.—The whole of lower Michigan is underlaid
by nearly horizontal Paleozoic strata, largely limestones and
shales of Devonian and Carboniferous age, but in the region under
HARPER: PLANT POPULATION OF MICHIGAN 25
consideration these are exposed only in limited areas near the
Great Lakes, and have little influence on soil or topography. The
interior is covered by glacial drift, averaging several hundred feet
thick and composed of sand, clay, pebbles and small boulders in
various proportions. The pebbles and boulders are quite diverse
lithologically, but most of them are more or less calcareous. Few
of the boulders are more than a foot or two in diameter, or large
enough to support any characteristic vegetation.
There is only one government soil survey for this region as yet,
and that is for one of the counties at its southern edge. But
Leverett’s bulletin, above cited, divides the soils of the Lower
Peninsula into about half a dozen classes, and gives the approxi-
mate acreage of each in every township and county. From the
returns from the twenty-one northernmost counties the following
percentages have been computed; and the corresponding figures
for the remaining forty-seven counties are given in an adjacent
column for purposes of comparison.
Northern Central and
Soil classes counties southern counties
OGY CU oo as Pal ke va eee a 26.9 21.0
oo ee ers rater cep aes a ee 25.9 24.7
CAVey Ss oer ess SC a a eek 18.6 38.2
Swarine and lakeee os eee et 17.9 8.8
CHavely IOAN oe. os ee ee ea 9.7 6.7
CHAVGE Se oo ori eae Ct i ee ee 1.0 0.5
The clayey till includes both moraines and glacial-lake de-
posits, which could have been separated if it had seemed worth
while. (The latter type in Michigan is chiefly confined to the
vicinity of Saginaw Bay and Lake Erie, and is very sparingly
represented in the region under consideration). Some of the larger
interior lakes are excluded from the estimate for swamps and lakes.
The sand on many of the uplands is so deep and loose as to
make walking and hauling on unimproved roads somewhat diffi-
cult in dry weather, just as in many parts of Florida. No satis-
factory chemical analyses of the soils seem to be available, but
they are doubtless below the average in fertility.
Topography and hydrography.—The topography is that com-
mon to many glaciated regions, undulating to hilly, with numerous
depressions containing lakes, ponds, swamps, bogs, marshes, etc.
The largest lakes cover about thirty-five square miles. The highest
26 HARPER: PLANT POPULATION OF MICHIGAN
elevations are only about nine hundred feet above the Great Lakes,
and are near the southern edge of the region, so that there are prob-
ably no differences in vegetation that can be ascribed to altitude
alone. Several series of supposed ancient beaches, formed in glacial
times when the Great Lakes were considerably higher than at pres-
ent, have been traced by Leverett and others, but as in the case of
the supposed marine Pleistocene terraces of Maryland and other
Atlantic states,* these episodes of geological history seem to bear
no obvious relation to the present vegetation, except as they may
have locally influenced soil or topography. (In other words, the
vegetation of an area that has been above the water say a hundred
thousand years does not differ noticeably from that of one only
ten thousand years old, if the soil is the same. And if the soil is
not the same, that is a matter for the geologist to explain, not for
the botanist.)
Being on a peninsula not exceeding a hundred and fifty miles
in width, this region has no large rivers. The drainage area of
the largest, the Cheboygan, covers about sixteen hundred square
miles. On account of the absence of rock ledges there are no
waterfalls, but in descending a few hundred feet from their sources
to their mouths the rivers necessarily traverse some gravelly
rapids, which are being utilized more and more for water-power.
The presence of many lakes and swamps and the coincidence of the
hottest season with the season of greatest precipitation makes the
flow of all the streams pretty steady, and on account of the pre-
vailing sandy soil and the scarcity of cultivated fields they carry
very little sediment. (In these respects also this region resembles
Florida more than it does most of the intervening states.)
Climate.—The average temperature is 41°-45° F., the January
mean 18°-23°, and the July mean 66°-69°. The proximity of two
of the Great Lakes presumably makes the difference between sum-
mer and winter climate a little less than it would be otherwise.
The average growing season or period free from killing frosts is
125 to 145 days, the average annual snowfall about seventy inches,
and the absolute minimum temperature about ——40° F.
This is one of the driest parts of the eastern United States,
having only about thirty inches of rain and melted snow annually.
* See Geog. Review 4: 224-225. Sept. 1917. :
HARPER: PLANT POPULATION OF MICHIGAN a7
An interesting feature of the precipitation is that late summer is the
wettest season, by a small margin. In this particular our region
resembles the pine-barren portions of the coastal plain, and differs
notably from southern Michigan and the whole Mississippi valley,
where spring and early summer are the rainy seasons. Coming as
it does just when the evaporating power of the sun tends to be
greatest, the late summer rain helps keep the ground-water level
constant and favors the formation of peat. It may bea mere coin-
cidence that two such sandy and peaty and “piney” and thinly
settled regions as northern Michigan and peninsular Florida should
both have a late summer rainy season, but it is more than likely
that the rainfall has influenced the soil somewhat, or even vice
versa through the vegetation, for there is no apparent physiographic
reason why the seasonal distribution of rain should be any different
in Michigan from what it is in Ohio and Indiana, for example.
Vegetation.—The aspects of the vegetation or the composition
of the plant associations have been described at considerable
length in some of the papers cited, particularly those by Livingston
and Gates (and Gates’s are illustrated), so that it is not necessary
to say much more on the subject here, except for pointing out
some fundamental principles often overlooked, and giving quanti-
tative data, which have not been supplied before except for very
small areas or for only a few species.
The dry uplands have vegetation of three principal types,
correlated with soil differences. On the more clayey soils the
original forests evidently were mainly of hardwoods and hemlock,
making a dense shade and considerable humus. On the most
sterile sands forests of jack pine prevail,* while in intermediate
habitats, covering most of the upland area, white and red pines
seem to have been the dominant trees before the lumbermen ap-
peared on the scene.
The streams have more or less meadow and river-bank vege-
tation along them, but few species seem to be confined to such
situations in the region under consideration. Flat areas adjacent
* See Spalding’s paper on “The Plains’’ previously referred to, and Beal’s Michi-
gan Flora, pp. 16-18. The jack pine, Pinus Banksiana, resembles the Florida spruce
pine, P. clausa, very much in gen appearance, habitat, and relations to fire.
(See Ann. Rep. Fla. Geol. Surv. 7: 142-144, 155- 1915.)
28 HARPER: PLANT POPULATION OF MICHIGAN
to streams and lakes, with a slow circulation of water through the
muck or peat, are generally densely wooded with spindle-shaped,
short-leaved conifers of the type characteristic of snowy climates.*
Fic. 1. Peat bog in a small sandy depression close to south shore of Douglas
Lake, but apparently quite independent of the lake. Water deep and cool and not
subject to much fluctuation. The floating leaves are Nymphaea variegaia.
vegetation of the common slow-growing ‘‘high moor” or muskeg type, or the
Chamaedaphne association of Gates (1913, p. 57); mostly evergreen. The trees are
Picea mariana and Larix, and the shrubs mostly Chamaedaphne, Nemopanthes,
Andromeda, and Kalmia polifolia. Herbs are relatively inconspicuous, but there
is an abundance of sphagnum. (Thisis one of the few known localities in Michigan
for Razoumofskya pusilla, which is parasitic on the spruces.)
The basin-like depressions, which may or may not have visible
outlets, have quite a variety of vegetation, depending on their
size, depth, etc. The larger ones contain lakes, with little vege-
tation in their deeper parts, many characteristic aquatics in shel-
tered shallow bays, and still other species, mainly of rush-like
aspect, on wave-washed sandy beaches. Around the lakes are
also numerous lagoons cut off by barrier beaches, and these com-
monly contain marsh vegetation, composed largely of grasses,
sedges and rushes. The isolated depressions which are too small
for wave action are usually occupied, at least around their edges,
* See Pop. Sci. Monthly 85: 340-341. I9I4.
HARPER: PLANT POPULATION OF MICHIGAN 29
by either marsh or bog vegetation (corresponding to Warming’s
“low moor” and “‘high moor”’ respectively.*)
The most significant difference between marsh and bog vege-
tation (and one apparently overlooked by Warming in contrasting
his low and high moors) is in the rate of growth; and just why
rank grasses should occupy one pond and the slow-growing sphag-
num, evergreen shrubs, and stunted conifers another has never
been fully explained. The depth of the water undoubtedly has a
great deal to do with it, for a shallow pond is quickly warmed by
¢ A
Fic. 2. Looking north across a oe —— or sedgy pond, with stagnant
water quite warm in summer, a sort of “low moor’’, 20-25 acres in extent, about 2
miles southeast of Douglas Lake. Vegetation coms Carex filiformis, with a few
wand Lams of Chamaedaphne and Andromeda. Partly burned conifer swamp
n background. = little to the left of this view, near a moderately rich hillside,
e ra therefore faster-growing vegetation, including Calamagrostts
me ran
canadensis, ne peg Sparganium, and Iris versicolor.)
the sun, and one in Michigan, where the sun shines nearly sixteen
hours a day in midsummer, may be nearly as warm during the grow-
ing season as one in Florida, where there is not more than fourteen
hours of sun ina day. Furthermore, a deep pond, besides being
colder, may be too deep in the middle for aquatic plants rooted in
the soil (such as the Nymphaeaceae) to reach the surface, so that it
* See his Oecology of Plants (1909), pp- 196-199, 200-205.
30 HARPER: PLANT POPULATION OF MICHIGAN
can be filled only by means of a floating mat of vegetation growing
out from the edges.* And plants not connected with the soil,
whether epiphytic or floating, cannot grow very fast on account of
the dearth of mineral plant food. The peculiarities of bog vegeta-
tion have been commonly attributed to the acidity of the water, but
the acids in bog water are not a fundamental part of the environ-
ment, but are derived from the vegetation, so that offering such
an explanation is only reasoning in a circle.
_ The size of the basin in which the pond or bog is located also
influences the vegetation through the amount of seasonal fluc-
tuation of water (as was observed in Florida a few years agof).
For considerable fluctuation of water in or above the ground hastens
the decay of dead vegetation and the liberation of the food in it,
and peat is formed best in places where the water-level is nearly
constant.§
The occurrence of typical slow-growing (sometimes erroneously
called xerophytic) bog vegetation in flat, slowly drained areas as
well as in deep stagnant basins is probably to be explained largely
by the fact that seeping water, having just emerged from the
ground, is considerably cooler in summer than shallow standing
water.|| The dense growth of conifers in such places still further
protects the water from the heat of the sun, after they are once
established. Another important factor is that in a perpetually
saturated soil the lack of aération restricts the availability of the
mineral plant food in the soil, particularly the potassium com-
pounds. Of course it may be said that the soil of a marsh is also
perpetually saturated, which is true enough; but in a marsh,
whether stagnant or estuarine, the water has been exposed to
evaporation much longer than that in a seepage or spring-fed
swamp, and thus the soluble salts in it are more concentrated.
* Probably the best description of this process is that by C. os Davia: in Rep.
Mich. Geol. Surv. 1906: 125- 907.
{ For a summary of bog theories, in which however the question of mineral nu-
trients is hardly considered at all, see G. B. Rigg, Plant World 19: 310-325. ‘Oct.
[Nov.], 1916
t See Tocris TI; 225-234. 1911; Ann. Rep. Fla. Geol. Surv. 6: 202, 203. figs.
46-48. 1914.
§ See Ann. Rep. Fla. Geol. Surv. 3: 211. IOI,
|| The temperature of springs, other than thermal springs i is — very close
to the average annual temperature of the locality where they
HARPER: PLANT POPULATION OF MICHIGAN dl
Furthermore, most marsh plants have hollow or spongy stems or
petioles, or aerenchyma, or pneumatophores, all of which doubt-
less serve to conduct air to the roots.
It might be observed here, parenthetically, that many plants
which in the region under consideration are chiefly confined to
bogs grow equally well on uplands in colder climates.* This is
probably because the low temperature and short growing season
farther north so limit the availability of the nutrients in the soil
that none but slow-growing plants can thrive.
An elaborate system of hypotheses of succession has been
postulated by Gates and others who have worked in this or some-
what similar regions, and some have even gone so far as to try to
connect all the plant associations in a limited area by successional
relations. But some of the imagined successions can never take
place without profound topographic changes, which may or may
not come to pass, and with which the botanist is not particularly
concerned. There are, however, two genuine types of succession
(biotic succession as defined by Cowles,} and distinguished from
his regional and topographic successions) which can be studied to
advantage in this region. The first is that connected with the
filling of lakes, etc., with vegetation and the gradual accumulation
of peat and humus. In a coniferous swamp the falling leaves,
twigs, trunks, etc., gradually pile up high enough above the ground-
water level to be subject to ordinary decay, and thus form humus
or duff instead of true peat. In such humus grow many plants
which are equally characteristic of the upland hardwood forests,
and this has led some to believe that the swamps, barring human
interference and unforeseen complications, will ultimately be
replaced by beech-maple-hemlock forests. But there are quite a
number of plants in this region which seem to demand both humus
and access to mineral soil or alkaline peat, such as Acer saccharum,
Tsuga, Fagus, Tilia, Ulmus, Quercus alba, Viburnum acerifolium,
Vagnera racemosa, Carex arctata, C. laxiflora, Washingtonia, Actaea
alba, Circaea, Adiantum pedatum, Geranium Robertianum, Caulo-
phyllum, and several others less common here, and we have no
* See Livingston, Bot. Gaz. 39: 40. 1905; Harper, Pop. Sci. Monthly 85: 340.
I9r4.
+ Bot. Gaz. 51: 161-183. I9gII.
32 HARPER: PLANT POPULATION OF MICHIGAN
evidence that these will ever grow on top of deep sour peat.
Furthermore, the swamps are colder at night than the adjoining
slopes, and lack some of the characteristic soil fauna of clayey
uplands, and these fundamental differences can hardly be oblit-
erated by succession.
It has been assumed also that the pine forests on the sandy
uplands would likewise be succeeded by hardwood forests, if fire
and lumbering did not prevent the accumulation of humus.
Although the sand is undoubtedly poorer than the clay, one might
suppose that in the course of centuries the pines could bring up
enough mineral matter from a depth of several feet, and deposit it
on the surface in decaying leaves, to make a rich soil that would
support trees that make a complete new crop of leaves every year,
and that such a forest when once established would be self-per-
petuating, as it would return to the soil every fall what it took up
in the spring and summer. But the difficulty is that in sandy soils
leaching probably goes on fast enough to prevent the accumula-
tion of any considerable amount of plant food on the surface of
the ground. (On clayey slopes there is less leaching, and erosion
is constantly exposing fresh layers of soil, and thus maintaining
its fertility indefinitely.) It seems that in many if not most cases
the proportion of deciduous trees in a forest does not increase with
succession, but depends on fundamental soil characters.*
Another type of succession is a periodic one resulting from fire.
In a state of nature the hardwood forests were rarely visited by
fire, and one might say figuratively that the plants in such habi-
tats made no provision for such an occurrence. The pine forests
have been so nearly destroyed that it is difficult to determine what
the normal frequency of fire in them may have been. As Pinus
resinosa is not very sensitive, it may have been subject to ground-
fires every few years, like some of the southeastern pines; while
in the white-pine forests fire may have been almost as rare as in
the hardwoods. Be that asit may, the “‘ pernicious activities” of
the lumbermen a few decades ago removed the greater part of
these two valuable pines,t and the ground formerly occupied by
* See H
. W. Wiley, Science II. 17: 794-795. May ts, 1903; Harper, Bull.
. Torrey Club 41: 209-220. 1914; B. Moore, Bot. Gaz - 61: 59-66. Jan. 1916.
T See Pop. Sci. Monthly 85: 343. IQI4.
7 Th Spe £* 4 . eee 4 roe
L * 1 Ee BS
7a
ibed by several
HARPER: PLANT POPULATION OF MICHIGAN 33
them has now a low scrubby growth of birch and aspen, which is
burned too often for the white pine to reéstablish itself, though the
red pine is making some headway.
On steep bluffs and small islands, and in ravines, where fire is
necessarily rare, we find some plants that are sensitive to fire but
do not require as rich a soil as that of the hardwood forests, such
as the few woody vines of the region. (Most of the existing white
pines are found in such places, too.) The low sandy “‘ice ramparts”’
around the larger lakes are protected from fire on one side by the
water and at the same time are too sterile to support vegetation
dense enough to carry fire readily, so that certain fire-sensitive
(or pyrophobic, if one may coin a new term) plants, such as A melan-
chier sp., Prunus pumila,* Rosa sp., Rhus Toxicodendron, Arcto-
staphylos Uva-ursi, Equisetum hyemale, Elymus sp., and Potentilla
Anserina, are characteristic of such places.
The normal frequency of fire in the jack-pine and spruce types
of forest seems to be about once in the average lifetime of a tree.
Pinus Banksiana is one of several pines whose cones remain closed
and attached to the tree for many years, but open soon after a
fire and discharge their seeds, thus re-stocking the forest. In the
spruce bogs, as in the white-pine forests, one of the first effects of a.
fire sweeping through the crowns of the trees is to liberate the
potash and other mineral substances stored up in several years’
growth of leaves and twigs, which falls to the ground and acts as a
high-grade fertilizer. Several quick-growing and short-lived trees
and shrubs, such as Betula, Populus tremuloides, P. grandidentata,
Prunus pennsylvanica, and a few characteristic herbs, known
collectively as fire-weeds, soon invade the burned areas by means
of seeds carried by wind or birds, and flourish until the surplus
potash, etc., is exhausted. In the swamps one of the conifers
itself, namely the deciduous one, Larix, acts as a sort of fireweed,
being especially common in the first few decades after a fire; and
its career is commonly terminated by saw-flies instead of fire.
As it renews its whole crop of leaves every year it needs a larger
food supply than the spruces, and it also extends farther into the
of the writers cited in the bibliography, particularly Dr. Rusby. Dr. H. A. Gleason
told me in 1912 that he did not know of a single acre of virgin white-pine forest in
— Michigan
bes Rises 18: 202. Sept. 1916.
34 HARPER: PLANT POPULATION OF MICHIGAN
richer regions of southern Michigan, Ohio, etc., than they do.
While the fireweeds are flourishing, seeds of the original conifers
are being brought in in the regular manner, and by the time the
birches and aspens have run their course the conifers that have
Fic. 3. Scene in large tamarack swamp with a few feet of peat, about four
_miles north of Douglas Lake, bordering a sluggish stream flowing into Carp Lake.
Trees in background all or nearly all Larix laricina. (See Gates, 1913, pp. 64-66.)
Open space in foreground evidently severely burned several years before. Woody
plants in it mostly Larix seedlings, Salix pedicellaris, Betula pumila, Rhamnus alni-
folia, Chamaedaphne, Alnus, Lonicera canadensis and Ledum. Evergreens in the
minority; little or no sphagnum. (This view was taken a mile or more from the
stream, and near the south edge of the swamp, where it is bordered by clayey hills
mostly cultivated. Toward the stream the percentage of evergreens increases con-
siderably.)
been growing up in their shade may be ready to dominate the
situation again.*
Since the best pines have disappeared the lumbermen have
attacked the hardwood forests also, and much hemlock has been
destroyed for tan-bark. The slash left from these operations is
subject to fire too, but the fireweeds on hardwood land are not
quite the the same as on pine land. Sambucus pubens and Rubus spp:
* So ine many papers have been written about the effects of fire in ‘boreal coniferous
forests that it would not be worth while to attempt to cite them here, but references
t© several of them can be found in Bull. Torrey Club 35: 349. 1908, and in Pop. Sci.
Monthly 85: 341. 1914.
HARPER: PLANT POPULATION OF MICHIGAN 35
are quite characteristic, and the vacciniums are rare or absent.
A hardwood area after logging, and even after the slash is burned,
is very disagreeable to traverse, on account of the numerous logs
and tops cumbering the ground, many of the logs being held up by
stiff branches at such a height that it is just as hard to climb over
them as to crawl under them, and only the smaller branches are
consumed by fire, the larger ones making veritable chevaux-de-
frise that last for many years. (Ten or twelve degrees farther
south fallen trees decay much more rapidly, and do not materially
impede the explorer after two or three years.*)
Farmers have damaged the vegetation still further by totally
eradicating much of it to make room for crops. This influence has
been chiefly concentrated on the hardwood land, on account of its
richer soil, but only about 17 per cent. of the area was classed as
“improved land” in 1910. The extension of farms ought to have
one indirect beneficial effect on the pine land, however, by multi-
plying the barriers to fire and thus diminishing its frequency at
any one point.
Plant census.—The following plant list is based on the writer’s
observations in northern lower Michigan between June 28 and
August 24,1912. Although only nine of the twenty-one counties
were visited in that time, the results are probably representative
enough, except for the dunes and cliffs along the Great Lakes, which
were not examined. The relative abundance of the species has
been determined in the manner explained in several previous papers
which are easily accessible.{ Although some may question the
accuracy of my rapid reconnoissance methods, there i is probably
no one at the present time in a position to assert that the results
*In this connection see W. — Long, Investigations of the rotting of slash in
Arkansas. U.S. Dept. Agr. Bull. 496. Feb. 1917.
naomminmas’ —— however, aisaws degrees farther south the tropical hard-
subject to fire at long intervals, followed
Mah a few characteristic fireweeds (Trema, Carica, etc.), and are just as disagreeable
to traverse for some time afterward as these Michigan forests, on account of the
hardness and durability of the wood of many of the trees
} For an a account of the effects of pare on fire frequency in the
Ozarks, see Marbut, Field Operations U. S. Bur. Soils 1911: 1740. 1914. (Or page
20 of the separates.)
¢ Ann. Rep. Fla. Geol. Surv. 6: 177-180. 1914; Bull. Torrey Club 41: 557-559-
1914; 44: 47-50. .1917; Torreya 17: 1-2, 5-6. 1917.
36 HARPER: PLANT POPULATION OF MICHIGAN
are incorrect. Future explorations will doubtless necessitate
changing the sequence of many of the species remote from the
head of the list; but as it is, the present sequence corresponds very
well with what one might obtain by counting the number of times
each species is mentioned in previous descriptions of the vegetation
of the same region.
The species are first divided into large trees, small trees, vines
and large shrubs, small shrubs, and herbs. Large trees are large
enough to be sawn into lumber, small trees large enough for posts,
large shrubs for canes or bean-poles, and so on down. Woody
vines are combined with large shrubs, because otherwise in the
absence of numbers there would be nothing to indicate how rela-
tively scarce they are, but their names are italicized to distinguish
them. Some small evergreen plants which have perennial stems
above the ground, such as Eguisetum spp., Lycopodium, Chimaphila,
Gaultheria, Epigaea, Chiogenes, Oxycoccus, Mitchella, and Linnaea
( “chamaephyte” class of Raunkiaer, in part), are classed
with the herbs on account of their small size and lack of true woody
tissue.
Evergreens are indicated by heavier type, as usual, and the
names of weeds enclosed in parentheses, whether they are treated
as exotics in the manuals or not.* The three commonest modes of
dissemination are indicated by somewhat arbitrary but suggestive
symbols, as follows: wind-borne seeds, Y; fleshy fruits, O; barbed
fruits, X. It would have been interesting to indicate annuals,
biennials, perennials, etc.} (or better still perhaps the Raunkiaerian
growth-formst!}, as well as the blooming periods and color of flowers,
but there are too many cases in which these data are not yet
known accurately enough. The usual habitats of the different
species are given in a few words.
In each major group the species are arranged as nearly as
possible in order of present abundance. Very likely a similar
census taken fifty years ago would have placed Pinus Strobus at
the head of the list, and arranged some of the other trees dif-
ferently, and excluded some of the weeds entirely. On account of
* See Bull. Torrey Club 35: 347-360. 1908; 37: II7-120. 1910.
-T Asin Ann. N. Y. Acad. Sci. 17: 36-37. 10906, for example,
t For an easily accessible reference to Raunkiaer’s system, and an illustration of
_ use, see Taylor, Am. Jour. Bot. 2:32. 1915.
HARPER: PLANT POPULATION OF MICHIGAN 37
the incompleteness of the data, and the great changes that have
been made by lumbermen and others, percentage numbers are
not attempted. But the numbers when finally determined prob-
ably will not depart much from a geometrical progression.* (The
commonest herb, however, seems to be about four times as abun-
dant as the next one.) The rarer species, and all cellular crypto-
gams, are omitted, because they make up such an insignificant
part of the total vegetation.
TREES ;
is Lane IBTICIN Re ie Cy eh ee eal Cues Swamps and bogs
< MG GrtA i os sk ee es a ee
Ps Sent eo Sandy uplands, etc.
Te RoC ERR ERETE OG as or ee giao Rich uplands
‘Y; SOWew COneieneis, 3c ee ae ay
pg Way tate i Prt bs Wineaat, goa ee rice a eae eg ER Pri ie
1 a MT AS OUR he Pee es oe ies ews Swamps and bogs
VARIG DAR IBR yn es ek
CGR RN ek Pe ae ee Various habitats
I 2 idee OCIS 2 es cea oe he Rich uplands
AY Pewee WiNTe 2 a Se ee ee i es mps
Pe RR SOOGI so se ok eo ne Sandy uplands
re CA SUI Oe ee a Richer soils
Quercus borealis —— FEE ME NCU eee IR Sandy uplands
Y PiChe CANAOONEE oe Swamps and bogs
VY Betula lntes ooo i Se os ate Rich woods
Y Pinas Bauksiane 62 os a es Poorest soils
Quercus alas ie ra a a Uplands
Y Braxinuy americana. 6 525..5 sae ae eee Richer soils e
SMALL TREES
V Betula panyritera ci a ees ak ys Various habitats
Y Populus enenliides ras Une ee Oe ice eens Burned areas mostly
Y Populus cers ate OE Oates a a - ie
© Prunus peunsyivanica: ... 205.6. b os ee ss ey a
Y Populus ia ees ie. Ce pisos Rae ies ae Lake shores, etc.
Oatrea ViTeiAata 66 e655 ee Rich uplands
Y Salix Qeviatiliel 6 es ees Lake shores, etc.
LARGE SHRUBS AND V.
Amys Weenie ya ee ea > Swamps, etc.
W Sable lita ee ee we ee Wet places
Y SAue St os we sees ee ee Swamps, etc.
VY Acer pennsyivanicnm. 2). 0.25. 2.6 ee Rich woods
* See Torreya 13: 244. Oct. 1913.
~ Long known as Q. rubra. See Torreya 17: 135. hae IQI7.
t This includes several unidentified species. If y had been separated they
would of course stand lower in the list.
38 HARPER: PLANT POPULATION OF MICHIGAN
©: Sambuens pubens cv Sos oa a ities Pay aie Burned or cleared hardwood land
OO FURGIS BO ool Se ee ee Oe oe Caen es pees aan
Oo ic aa eg ee ly C2 SES gia SERRE Cee SS
Y Acer spi Ra ne ee eee ee Rich woods
O Cornus staloniferacs.) ise ea, Low grounds
() Nemopnanthes mucronata 0.8 oo ed sd caida: Swam
O Ilex tee eee es be ek eT ee baie Swamps
OA eM CEE Bl Oe acc oh ces Poe eis Various habitats
Vie Wal ae i ee ee eee Lake shores and bluffs
aioatieod BOOMERS Feo Pa ot Ait ka as < stes Bluffs and islands
Oo m cassinoides Decale acal ei pla Su iw aeeile ce Weace oes Swamps
Y Salix mee Ea ON ae Al Br i in, Serr ay ome Low grounds
SMALLER SHRUBS
D MA DOMME ey hs oo as coe oes Sandy uplan
Cr OO Be CLIERONUS 6 sree sa ore eek urned or cleared hardwood land
Chamaedaphne calyculata................... bog
OMIPtONIa Peregritg i804 oS a oc ee ok ales Sandy uplands
Mubus dileghenietiaia? un io cigs oo Burns, clearings, etc
mhus Poxitodentron.. 630 ee ct OO Ice-
Ledum groe WE ieee osc bre ee Bogs and swamps
w Dele POstratAr. | pb Da Vey eo a oe Swamps
O Vaccinium pennsylvanicum.................. Sandy uplands
O Sacus canadeusits.3 55 4 ich woods and swamps
© Hhanis atnifoliaig. 5 aoe eS Bogs and swamps
Myrica a chiens Cie) Ces e soll whe oon Ne ig
O Vaccinium cunadenat 6:0 568 Soo Ss tc 23 Sandy uplands
* Betula ppiiiia. eto) oo eC Ml ee ee Bogs and swamps
Lositers Caleieneit, ygiew vce). ss "
O Arctostaphylos Uva-ursi..................... B ns
RG BD rns ody oe et Cr das og Lake shores, etc
wOIreen TARO Fie ee Swamps and meadows
O Viburnum: aceroliamt . 665.0007 A Rich woods and bluffs
odon M pederspesia CP eri rint Cees AN Swamps and marshes
Oo ches nigra
ANS Rachie epenes COE Vi oe dee oe Sandy uplands, etc
Type ON ee aa Marshes, ete
Y Chamaenerion angustifolium................. Burns and clearings
(Poa piatensig ce oer ee Roadsides, etc
Calamagrostis canadensis Petts ieee eee arshes
O Aralia nudicaulis HEE Clank feel iy s Bene u ns Rich woods, shady bogs, etc.
O Unifolium canadense.................. cee: - a “ ecu
yv (Solidago COMAIPHMIEY cs as ee aes Fields and roadsides
* The sumacs of this region present a bewildering variety of ints of feecin rasta £ from
te petecny: ono R. ninbis to —— almost pubescent enough to be called R. ty-
R ty phi:
HARPER: PLANT POPULATION OF MICHIGAN
Y. Uigclentas eprigra) ii i Fields and roadsides
(Achillea Millefolium). .;....: 0... 0.65.00. c05 i ih .
Scirpus vali ie say sate REPRE N v oe hie Via et oie oe Lakes, e
pater Wienriniets ye rl Os he ee ax noe and ‘marshes
Y Dryopteris tinihinis are Bas ee ae stp eal gt weak
O Gaultheria procumbens.....................Sour soils, dry or moist
OC ornis tinadenaisd or wen el Rich woods and bogs.
Dulichium arundinaceum.................... Marsh
(Verbesrium Phaneus) ec! ek oa. sk gc eke Beheuck: roads etc.
¥ Eupatoriun, purpareums. 3364. = us Swamps and me
OO: Clibtonia: borealis ete age co ee ich woods and ns bogs
Nymphaea variezata™ 23.4 6602000 ve Quiet water
Sci pus americanus UC a ra ei As Lake shores, etc
O Kubus traflotuge 0 3 eo ce ee Bogs and swa
O Vagneta racemosa ie Rich woods
Ivis versicolor oo vis eee ee ere. arshes
Y Osmmunda reeatis 5.7 ae ea ee Swamps and marshes
Y oe ehh — ON OM cht nig gabe as Marshes and meadows
PAWEL ee Marshes, e
VY eae as Puen cle eek caneees Meadows, etc.
Thiadenum ViTsinicuMn. fics eS Bogs and marshes
Trientali ROR isc rr ees St a ich age and shady bogs
© ppeeias incarnate or i cs es ok Marshes,
iN aemere thiolacdis cei, Seo ee eed. Ri fea wood and aeee —
O Chiogenes hispidnla sip 0
¥ Osociea sensthilis co5cos aes occ es et Marshes, etc
Lysimachia terrestrias..¢ ofc fo a
(Pastinaca tativa).i¢ 26s sv oe Roadsides
O Mitchelia repens. 5c es Rich woods mostly
Potentilla Anserina (070. 322. ae Lake shores
Carex arctata ccd eet ee ee Rich woods and bluffs
Coptis trifolia “3. oo. a er eS Rich woods and shady bogs
spartinn Michanxintiae a. 570 ST is Lake shores, etc.
athyrus palustris. co 255 ey . i as
Y¥ (Leptilon camadense) or kee set a Burns and waste places
(Hatisetiit afvenue) ¢c0 665i. sa ei Along railroads, ete.
Linnaea MPICONE 5s a ee eek es Rich woods and shady bogs
Y Lycopodium otingme ee ea as he atthe ae *
Lilium philadelphicum. <3. 6 oe aes eee Swamps and meadows
Thalictrum dasyCarpum <0)0 eo. es ee. o e 2
rysanthemum Leucanthemum)............ Roadsides, etc.
Danthonia A eee | og eo Ne dy d
Equisetum as Og Rs i vat are Nae bogs
oe ek eae es oi ee Lake shores, etc
M (Cynoetossum officinale). 22. . 626i. ei 5... Roadsides
A Washinetonia Clayton. oo 6 eis ee es Rich woods
) Actaeet aes 6 ee a eo ee a a
* For a discussion of the nomenclature of this plant, see Fernald & St. John,
Rhodora 16: 137-141. Aug. 1914.
40 HARPER: PLANT POPULATION OF MICHIGAN
Wittella niudaevieg ee er co aS Rich woods and bogs
MOMMGR TONENR hes ON ee Sou ils
Campanula uliginosa . LUAU ie NOM Oe PG ca ipa ae DE Marshes
Piguisetitn Muvintie in. eo a ee As
OC ArsWi taro Rich woods
(Convolvulus spithamaeus) 2... 2. cs Sandy uplands
Y Apocynum androsaemifolium................ es ie
CPhienii pratenseln tas Poe a Roadsides, etc.
Menyanthes ee eebbiaaaet c/w. dy gts ep seis Gogg Bogs and marshes
Pileochaign palistrise: acy ie. oe ee Shallow water
AP Erin Seaneimoriiy) ye es ee Rich
‘ MOIR eee yo Lake shores
Y Eriophorum viridicarinatum................. Bogs, etc
© Oxycoccus macrocerpus.. 2.6603 ek Bogs
Y (Apocynum cannabinum).................... Ice ramparts, etc.
Usyropems Seperiolia: 6.2.8 Sandy bluffs, etc.
mreden alia ye eee aiias ok a eee es Rich woods
2 Carlin DURE ee SE eS oe Waste places
me PREUARNA Spe) So oa aig BB A Lake shores, etc.
Pediisctum inevieatim. 2 sy As 3
Porechinw Wireman. of so Rich woods
Panicum depaupetatum 2.45. .... 522.0. 5 Sandy uplands
Eq BCIEDOMIOR os os
Y Aster Np aera i eet nies a, Rich woods, etc
MG GLIOVITene 2 3 ee oS Meadows, etc
(Ambrosia paeumnelods HED ea eel ESL real Meer ne aoe Waste places
mohelia cavdinalig po Along brooks
Callie pacts... ee
» Gallant titeram ci oe Rich woods
rani Robettiangne 4.5 <0;
Tittle otltiibehc oto Roadsides, etc
(Tron Sen a
0 Polygonatum biotic. 4. io. Rich woods
Naumburpia Hivisiicing 20 oe
Comandra iwaolbelinte: occa 3 oe. Sandy uplands
Y Dryopt RUINGIOBR oo a a Rich woods
Riymee 6). cite a Lake shores
Phalarin arupdinnced 22.2065. 0 2 '
Cypripedium Birsatunis 660.0 Shady bogs
Summary.—The species above listed probably constitute
something like nine tenths of the vegetation and one tenth of the
flora of the region under consideration. About 275 additional
species, nearly all herbs, were observed, but not often enough to
be worth mentioning.
Not one of the large trees has fleshy or barbed fruits, while few
of the smaller shrubs are wind-distributed. There are no barbed
fruits on any of the woody plants, but a few of the small trees and’
HARPER: PLANT POPULATION OF MICHIGAN 41
many of the shrubs, vines and herbs have fleshy fruits.* Herbs
with barbed fruits are in this region chiefly confined to rich woods
and roadsides, probably partly because of the exemption of such
places from fire,t though it is not at present apparent why that
should affect them more than it does the fleshy fruits. (Barbed
fruits seem to be more characteristic of warmer climates and more
calcareous soils, too.) Plants with erect capsules on stiff stems
which stand up through the winter (called “tonoboles” by
Clementst) are much rarer here than a few degrees farther south,
possibly because the snow interferes with their dissemination.
About half the large trees are evergreen, but none of the small
trees, vines and large shrubs are, strange to say. It would seem
from this that no evergreens except conifers (and one of those is
deciduous) can stand the Michigan winters without the protection
of snow.§ Vines are scarce, only two being listed, and those are
not found much farther north.
The Ericaceae and allied families are largely represented among
the small shrubs and evergreen herbs, as in many other places
with similar climate. Other families pretty well represented in —
proportion to the total number of species in them, or the total flora
of this region, or both, are Equisetaceae, Cyperaceae, Orchidaceae,
Salicaceae, Rosaceae, and Caprifoliaceae, while the opposite might
be said of the Fagaceae, Cruciferae (native), Caryophyllaceae
(native), Leguminosae, Polygalaceae, Violaceae, Hypericaceae,
Umbelliferae, Labiatae, Scrophulariaceae, and Lentibulariaceae,
The sedges seem to be more numerous and also more abundant
than the grasses.
Nearly all the species listed are widely distributed, extending
from Nova Scotia to Minnesota at least, and most of them are
represented in northern Europe by identical or closely related
forms. Those peculiarly American plants which are confined to
* See Torreya 14: 16. :
(In the text on page 138 Allium and Deringa
ional references
that should have been given there are: S. M. Coulter, d
1904; Harper, Torreya 10: 60-61. 1910.
t. Surv. Neb. 7: 47. 1904
§ In this connection see Gates,
Acad. Sci. 1§: 194. 1914.
Torreya 12: 257-262. 1912; Harper, Rep. Mich.
42 _ HARPER: PLANT POPULATION OF MICHIGAN
the glaciated region and coastal plain or nearly so* are scarcely
represented here, presumably because the climate is a little too cold.
A list of the commoner plants of the eastern part of{the Upper
Peninsula, based wholly on car-window notes, was published by
the writer a few yearsago.f In that a slightly different method of
computation, which did not do justice to the conifers, was used,
and the different sizes of trees and shrubs were not separated.
But it is probably safe to say that Abies, Picea canadensis, Betula
pumila, and Andromeda glaucophylla (to mention woody plants
only) are more abundant in the Upper Peninsula than here; while
the reverse is true of Acer saccharum, Tsuga, Fagus, Tilia, Ulmus,
Acer pennsylvanicum, Sambucus, Rhus glabra (etc.), Diervilla,
Rubus strigosus, Comptonia, Rubus allegheniensis (?), Rhus Toxico-
dendron, Taxus canadensis, and a few others, most of which are
not evergreen.
Comparing this region with that adjoining it on the south we
get a greater contrast, due to better soil and climate both. The
commonest trees in the central third of lower Michigan (not count-
ing the lake plains around Saginaw Bay, which are still more
fertile), as determined from a few hours of car-window observa-
tions, seem to be as follows:
Quercus velutina (?), Pinus Banksiana,t P. Strobus, Ulmus
americana, Larix laricina, Quercus borealis maxima (?), Acer sac-
charum, Fagus, Thuja, Tsuga, Abies, Tilia, Pinus resinosa, Quercus
alba, and Picea mariana.
Still farther south the change in composition of vegetation
continues in the same direction, and in extreme southern Michigan
nearly all the trees are deciduous.
COLLEGE Pornt,
New Yor«
a ee ee es
* See Rhodora 7: 69-80. 1905; 8: 27-30. 1906.
T Rep. Mich. Acad. Sci. 15: 193-198. 1914.
f If I had left northern Michigan by way of the Michigan Central R. R. instead
of the Pere Marquette, Pinus Banksiana would doubtless have stood higher in the
first list and lower in the one on this page; for along the former railroad it is said to be
abundant in Crawford and Roscommon Counties, while along the oe i saw it
mostly in Lake and Newaygo Counties, south of the limits assigned in
(See second map between pages 550 and 551 of the oth volume of the fab Census. )
INDEX TO AMERICAN BOTANICAL LITERATURE
1913-1917
The aim of this Index is to include all current botanical literature written by
Americans, published in America, or based upon American material ; the word Amer-
ica being used in the broadest sense.
Reviews, and papers that relate exclusively to forestry, agriculture, horticulture,
manufactured products of vegetable origin, or laboratory methods are not included, an
made in favor of some paper appearing in an American periodical which is devoted
wholly to botany. Reprints are not mentioned unless they differ from the original in
Some important particular. If users of the Index will call the attention of the editor
to errors or omissions, their kindness will be appreciated.
This Index is reprinted monthly on cards, and furnished in this form to subscribers
at the rate of one cent for each card, Selections of cards are not permitted ; each
risnioned must take all cards published during the term of his subscription, Corre- .
pondence relating to the card issue should be addressed to the Treasurer of the Torrey
9
Barre, H. W. Report of the botanist and plant physiologist, Rep.
S. Carolina Agr. Exp. Sta. 27: 20-25. 1914; 29: 16-20. 1916.
Bates, C. G. Forest succession in the central Rocky Mountains.
Jour. Forest. 15: 587-592. My 1917.
Berry, E. W. A middle Eocene Goniopteris. Bull. Torrey Club 44:
331-335. pl. a2. 14 Jl et
Goniopteris claiborniana sp. n
Bessey, E. A. Yellow peice: a dangerous weed. Michigan Agr.
Exp. Sta. Spec. Bull. 80: 1-4. My 1917
Bessey, E. A. The hormone theory of chromosome actions.
Rep. Michigan Acad. Sci. 18: 53-58. 1916
Bessey, E. A. The origin of the Anthophyta. Rep. Michigan Acad.
Sci. 17: 142-151. pl. 13-15. 1916.
Bessey, E. A. The sexual cycle in plants.
Sci. 18: 59-77. f. 1-12. 1916.
Bigelow, M. A. Popular names of plants. Science II. 46: 16, 17.
6 Jl 1917.
Blake, S. F. A new Rudbeckia from Indiana. Rhodora 19: 113-115.
2 Jl 1917.
Rudbeckia Deamii sp. n
Blake, S. F. Veraonisa altissima Nutt. var. taeniotricha, var. nov.
Rhodora 19: 167, 168. 10S 1917.
43
Ann.
Ann. Rep. Michigan Acad.
44 INDEX TO AMERICAN BOTANICAL LITERATURE
Boncquet, P. A. Bacillus morulans n. sp. A bacterial disease organ-
ism found associated with curly top of sugar beet. Phytopathology
7: 269-289. f. 1-7. .7 S 1917.
Borzi, A. Studi sulle nux oficie. Nuo. Gior. Bot. Ital. N. S. 24:
65-112. Ap IgI7.
Includes several American species of Stigonema.
Boynton, K. R. Aster amethystinus. Addisonia 2: 39. pl. 60. 30
Je 1917.
Boynton, K. R. Notes from the herbaceous collections—II. Jour.
N. Y. Bot. Gard. 18: 141-143. Je 1917.
Britton, E. G. The conservation of wild flowers. Am. Mus. Jour.
17: 350-352. My 1917. [Illust.]
Britton, N. L. Forests, hygiene and ethics. New York Forestry 4:
24, 25, 32, Jl 1917. [Illust.]
Brigham, R. O. Sterilization of pop corn. Rep. Michigan Acad. Sci.
17: 190-193. I916.
Britton, N. L. El genero Rynchospora Vahl, en Cuba. Mem. Soc.
Cubana Hist. Nat. ‘‘Felipe Poey’’ 2: 151-166. [Je 1917.] é
Translated from the English text by Brother Léon. Reprinted as Contr. N. Y.
Bot. Gard. 194: 1-16. Je 1917.
Includes Rhynchospora siguaneana, R. joveroensis, R. Gageri, R. Shaferi, R-
nipensis, and R. Randit, spp. nov.
rooks, S. C. Methods of studying permeability of protoplasm to .
salts. Bot. Gaz. 64: 230-249. 15S 1917.
Brown, P. E. Bacteriological studies of field soils—III. Iowa Agr.
Exp. Sta. Research Bull. 13: 421-448. S 1913.
Bryan, G. S. The archegonium of Catharinea angustata Brid. (Alt-
richum angustatum). Bot. Gaz. 64: 1-20. pl. 1-8 + f.1. 17 J11917-
Burlingham, G. S. Methods for satisfactory field work in the genus
Russula. Mycologia 9: 243-247. 30 Jl 1917.
Burnham, S. H., & Latham, R. A. The flora of the town of Southold,
Long Island and Gardiner’s Island. First Supplementary list.
Torréya *7: 411-122. 18 Jl 1917.
Hysteriographium Vaccinii n. comb.
Burns, G. P., & Peitersen, A.K. Agricultural seed: concerning weeds
and weed seeds. Vermont Agr. Exp. Sta. Bull. 200: 5-79. f. I-52.
S 1916.
Bush, A. D. Pathophytes and pharmocophytology. Rep. Michigan
Acad. Sci. 18: 43-46. 1916. -
Butler, O. The cuprammonium washes, their preparation, biological
properties and application. Phytopathology 7: 235-268. pl. 3-10.
7 S1917 .
INDEX TO AMERICAN BOTANICAL LITERATURE 45
Butters, F. K. Pellaea atropurpurea (L.) Link and Pellaea glabella
Mett. ex Kuhn. Am. Fern Jour. 7: 77-87. Jl 1917.
Butters, F. K., & St. John, H. Studies in certain North American
species of Lathyrus. Rhodora 19: 156-163. 108 1917.
Candolle, C. de. Meliaceae centrali-americanae et panamenses.
Smithsonian Misc. Coll. 688: 1-8. 23 Jl 1917.
Includes rr new species in Gaurea (9) and Trichilin (2).
Childs, L. New facts regarding the period of ascospore discharge of
the apple scab fungus. Oregon Agr. Exp. Sta. Bull. 143: I-11.
My 1917.
Chupp, C. Studies on clubroot of cruciferous plants. Cornell Agr.
Exp. Sta. Bull. 387: 421-452. f. 95-110. Mr 1917
Plasmodiophora Brassicae.
Clinton, G. P. Report of the botanist for 1913. Connecticut Agr.
Exp. Sta. Rep. 38: 2-42. pl. 1-8. 1915.
Includes: Notes on plant diseases of Connecticut; (2) So-called chestnut
blight poiso: :
Cockerell, ca D. A. Adult characters in sunflower seedlings. Jour.
Heredity 8: 361, 362. f. 13. Au 1917. ;
Cockerell, T. D. A., Lamium amplexicaule in Colorado. Torreya
27:.123. 48 jl 1917.
Coker, W. C. The amanitas of the eastern United States. Jour.
Elisha Mitchell Sci. Soc. 33: 1-88. pl. 1-69. Je 1917.
Includes Amanita hygroscopica sp. nov. and Amanita mappa var. lavendula, A.
spissa var. alba and A. rubescens var. alba, var.
Collins, F. S., & Hervey, A. B. The aces 6f Bermuda. Proc. Am.
Acad. Arts and Sci. 43: 1-195. pl. 1-6. [31] Au 1917.
Includes descriptions of new species in Dermocarpa (1), Hormothamnion (1),
Oedogonium (1), Endoderma (1), Chaetomorpha (1), Cladophora (3), Codium (1).
(1), Spermothamnion (1), Gymnothamnion (1), Ceramium (2), and Rhodochorton (1);
also various new combinations and form names.
i. Common diseases of apples, pears and quinces. New
Jersey Agr. Exp. soe 24e 80: 1-27. f. I-23. 10 My 1917.
A revision of Circ. 44:
Cook, M. T. paste: decane of beans and peas. New Jersey Agr.
Exp. Sta. Circ. 84: 1-8. f. 1-4. 9 Je 1917.
Cook, M. T. Common diseases of the peach, plum and cherry. New
Jersey Agr. Exp. Sta. Circ. 81: 1-19. f.I-1Ir. 19 My 1917.
A revisoin of Circ. 45: I915-
Cook, M.T. A Nectria parasitic on Norway maple. Phytopathology
7: ats, 314. 67 5 1917:
Cook, M. T., & Martin, W. H. Diseases of tomatoes. New Jersey
Agr. Exp. Sta. Circ. 71: 1-8. f. 1-6. Ap 1917.
46 INDEX TO AMERICAN BOTANICAL LITERATURE
Cook, O. F. Staircase farming of the ancients. Nat. Geog. Mag. 29:
474-534. My 1916. [Illust.]
Includes numerous references to Peruvian plants.
Coons, G. HH. An undescribed bark canker of apple and the associated
organisms. Rep. Michigan Acad. Sci. 17: 117-122. pl. 6. 1916.
Coons, G. S. The Michigan plant disease survey for 1914. Rep.
Michigan Acad. Sci. 17: 123-133. pl. 7-10. 1916,
Davie, R. C. Some Brazilian plants. Jour. Bot. 55: 215-223. Au
Includes Gaultheria Willisiana sp. nov.
Dearing, C. Muscadine grape breeding. Jour. Heredity 8: 409-424.
J. 40-18... S 10917.
Delavan, C. C. The relation of the storage of the seeds of some of the
oaks and hickories to their germination. Rep. Michigan Acad.
Sci. 17: 161-163. 1916.
Doolittle, S. P. Cucumber scab caused by Cladosporium cucumeri-
num. Rep. Michigan Acad. Sci. 17: 87-116. 10916.
Farwell,O.A. Fern Notes. Ann. Rep. Michigan Acad. Sci. 18: 78-94.
FoL7s 1916:
Fernald, M. L. Contributions from the Gray Herbarium of Harvard
University.— New Series, No. L. Rhodora 19: 133-155. I0S 1917.
I. Some polygonums new to North America. II. New or critical species or
varieties of Ranunculus. III. Some color forms of American anemones. IV. New
species, varieties and forms of Saxifraga. V. A new Vitis from New England. VI.
Gentiana clausa a valid species. VII. Some forms of American gentians. VIII.
Some new or critical plants of eastern North America.
Gleason, H. A. A prairie near Ann Arbor, Michigan. Rhodora 19:
163-165. 10S 1917. 3 ‘
Goss, R. W.; & Doolittle S. P. The effect of fungicide on the spore
germination of Longyear’s Alternaria. Rep. rile Acad. Sci.
¥7: 183-187. tors.
Gourley, J. H. Fruit bud formation—a criticism. Bull. Torrey Club
44:455-457-f. 2. 15S 1917.
Gruzit, O. M., & Hibbard, R. P. The influence of an incomplete cul-
ture solution on photosynthesis. Rep. Michigan Acad. Sci. 18:
50-52. I9I16.
[Abstract.]
Hahn, G. G., Hartley, C., & Pierce, R.G. A nursery blight of aaah
Jour. Agr. Research 10: 533-540. pl. 60, 61. 3S 1917
Hance, R. T. An attempt to modify the germ plasm of Oenothera
hrditaga the germinating seed. Am. Nat. 51: 567-572. S 1017.
O. E., & Hibbard, R. P. A simplification of the present
INDEX TO AMERICAN BOTANICAL LITERATURE 47
freezing-point method for the determination of the osmotic pressure
of plant sap. Rep. Michigan Acad. Sci. 18: 47, 48. 1916.
[Abstract.]
‘ Hedgcock, G. G., & Hunt, N.R. Notes on Razoumofskya campylopoda.
Phytopathology 7: 315, 316. 7S 1917.
Higgins, J. E. The litchi in Hawaii. Hawaii Agr. Exp. Sta. Bull. 44:
I-21... pl. 7-§. 27 Jl-4917.
Hitchcock, A. S., & Chase, A. Grasses of the West Indies. Contr.
Ve S. Nat. Herb. 18: i-xviii seal Meath 18 Au 1917.
ncludes descriptions of the new genus Saugetia and 11 new species in Saugetia
Ci); gona (1), Paspalum (5), Raddia (2), Spieigcs (1), and Thrasya (1), and
several new combinations.
. Hollinger, A. Does the movement of air affect the growth of plants?
Rep. Michigan Acad. Sci. 17: 159, 160. 1916.
[Abstract]
Hottes, A. C. Gladiolus studies—I. Botany, history and evolution
of the Gladiolus. Cornell Extension Bull. 9: 93-188. f. z-9. D
1916; II. Culture and hybridization of the Gladiolus. Cornell
Extension Bull. 10: 195-271. f. ro-38. D 1916; III. Varieties of
the garden Gladiolus. Cornell Extension Bull. 11: 277-451. f. 39-68.
D 1916.
House, H. D. The Peck testimonial exhibit of mushroom models.
Science II. 46: 204. 31 Au 1917.
House, H. D. Photographing wild flowers. Kodakery 4: 10-13. Jl
1917. [Illust.]
Howitt, J. E. Phytophthora infestans, causing damping-off of tomatoes.
Phytopathology 7: 319. 7S 1917.
Hurd, A. M. Winter condition of some Puget Sound algae. Puget
Sound Mar. Sta. Publ. 1: 341-348. 10 Au 1917.
Hurst, B. Peloria flowers on ivywort or ivy-leaved toadflax. Ottawa
Nat. 31: 45. Jl 1917
Kauffman, C. H. Uinveborind Michigan fungi for 1911, 1912, 1913, and
1914. Rep. Michigan Acad. Sci. 17: 194-223. 1916.
Kenoyer, L. A. Environmental influences of nectar secretion. Iowa
Agr. Exp. Sta. Research Bull. 37: 219-232. N 1916.
Levin, E. Light and pycnidia formation in the Sphaeropsidales. Rep.
Michigan Acad. Sci. 17: 134, 135- 1916.
MacDougal, D. T. To increase the yield of food. N. Am. Review
206: 62-69. “Jl” [Je] 1917.
Mains, E.B. Some factors concerned in the germination of rust spores.
Rep. Michigan Acad. Sci. 17: 136-140. 1916.
Markle, M. S. Root systems of certain desert plants. Bot. Gaz. 64:
177-205. f. I-33. 15 S 1917.
48 INDEX TO AMERICAN BOTANICAL LITERATURE
Martin, W. H. Sclerotium bataticola. The cause of a fruit-rot of
peppers. Phytopathology 7: 308-312. f. r-12. 7S 1917.
McIndoo, N. E., & Sievers A. F. Quassia extract as a contact insec-
ticide. Jour. Agr. Research 10: 497-532 3S 1917.
Contains some botanical information.
{Moore, G. T.] Native wild mushrooms for food. Missouri Bot.
Gard. Bull. 5: 119-129. pl. 17-23. Au 1917.
Muenscher, W. C. A key to the Phaeophyceae of Puget Sound.
Puget Sound Mar. Sta. Publ. 1: 249-284. pl. 47-67. 6 Je 1917.
Murrill, W. A. A giant puffball. Jour. N. Y. Bot. Gard. 18: 193. pl.
£05.20 3 1917.
Nash, G. V._ Hardy woody plants in the New York Botanical Garden.
Jour. N. Y. Bot. Gard. 18: 167-170. Jl 1917; 189-192. 20S 1917; *
203-207. 23 O 1917; 217-224. O 1917.
Piemeisel, F. J. Factors affecting the parasitism of Ustilago Zeae.
Phytopathology 7: 294-307. 7S 1917.
Popenoe, W. The pollination of mango. U.S. Dept. Agr. Bull. 542:
1-20. pl. r-4 +f.7. 11 Au 1917.
Povah, H. W. Helicostylum and Cunninghamella: two genera of
Mucorales new to the state. Rep. Michigan Acad. Sci. 17: 152-155.
pl. 76, 17. - 1916.
Praeger, W. E. Growing alien cacti in Michigan. Rep. Michigan
Acad. Sci. 17: 156-158. 1916.
Prince, F. S. The soy bean in New Hampshire. New Hampshire
Exp. Sta. Dept. Agron. Bull. 181: 1-20. Mr 1917. [Illust.]
Rands, R. D. The production of spores of Alternaria Solani in pure
cultures. Phytopathology 7: 316, 317.f.1. 7S 1917.
Rigg, G. B. Seasonal development of bladder kelp. Puget Sound
Mar. Sta. Publ. 1: 309-318. 10 Au 1917.
Roberts, J. W. Control of peach bacterial spot in southern orchards.
U.S. Dept. Agr. Bull. 543: 1-7. 8 Au 1917.
Disease caused by Bacterium pruni Smith.
Rumbold, C. Notes on effect of dyes on Endothia parasitica. Bot.
Gaz. 64: 250-252. 15S 1917.
Rydberg, P. A. Phytogeographical notes on the Rocky Mountain
region—VII. Formations in the Subalpine Zone. Bull. Torrey
Club 44: 431-454. 15 S 1917.
Sackett, W. G. A bacterial stem blight of field and garden peas.
Colorado Agr. Exp. Sta. Bull. 218: 3-43. pl. 1-2 + colored plate +
Jo i-3. Ap i916. ;
Sawyer, W. H., Jr. Development of some species of Pholiota. Bot.
_ Gaz. 64: 206-229. pl. 16-20. 15S 1917.
INDEX TO AMERICAN BOTANICAL LITERATURE 49
Seaver, F. J. Damage from soil fungi. Jour. N. Y. Bot. Gard. 18:
186-188. 20S 1917.
Also published in Sci. Ann. Suppl. 2184: 294. 10 N 1917.
Selby,A. D. Diseases of wheat. Methods of control possible by seed
treatment. Ohio Agr. Exp. Sta. Monthly Bull. 2: 219-222. Jl
1917.
Brief notes on Colletotrichum cereale, Fusarium roseum, Ustilago tritici and Til-
letia foetens.
Shear C. L., & Stevens, N. E. Studies of the Schweinitz collections of
fungi—I. Sketch of his mycological work. Mycologia 9: 191-204.
bl. 8, 9. 36 JL 1617.
Sherbakoff, C. D. Fusaria of potatoes. Cornell Agr. Exp. Sta.
Het 6: 97-270. pl. 1-7 +f. 1-51. My 1917.
veral new ipecies and varieties described.
pen C. D. Report of the associate plant pathologist. Florida
Agr. Exp. Sta. Report 1916: 80 R-98 R.f. 12-16. My 1917.
Sherbakoff, C. D. Some important diseases of truck crops in Florida.
Florida Agr. Exp. Sta. Bull. 139: 193-277. f. 76-112. Je 1917.
Sherff, E. E. Studies in the genus Bidens—IV. Bot. Gaz. 64: 21-41.
17 Jl 1917.
pines Bidens mollifolia, B. cornuta, B. leptocephala, B. Langlassei, B. a
lifolia, B. carpodonta, B. pseudalausensis, B. Brandegeet and B. Holwayi, spp.
Shimek, B. The plant geography of the Lake Okoboji region: me
Bull. Lab. Nat. Hist. Univ. Iowa 7‘: 3-5. O 1917.
tional notes.
Bull. Lab. Nat. Hist. Univ.
Shimek, B. The sand flora of Iowa.
Iowa 74: 6-24. pl. 1-5. O 1917.
Shufeldt, R. W. Forms of leaves. Am. Forest. 23: 412, 413. Jl 1917.
[Illust.]
Shufeldt, R. W. Midsummer flowers. Am. Forest. 23: 403-406. f.
r-5., Jl 1917.
Shufeldt, R. Ww. Remarkable buttonballs.
2 As A
Shufeldt, R. iv oe flowers that boys and girls should know. Am.
Forest. 23: 474-480. f. 1-20. Au 1917.
Shull, G. H. Pitcher-leaved ash trees. Jour. Heredity 8: 431. f. ro.
S 1917.
Siggers, P. V. Some cultural characteristics of Pestalozzia funera
Desm. Rep. Michigan Acad. Sci. 17: 141. pl. 12. 1916.
Sladen, F. W. L. Some investigations of the honey flora in Ontario.
Ann. Rep. Beekeepers Assoc. Ontario 37: 31, 32. 1917.
Small, J. K. The earciaale walnut, oe: N. Y. Bot. Gard. 18: 186.
pl. 203, 204. 20S 1917.
Jour. Heredity 8: 310.
50 INDEX TO AMERICAN BOTANICAL LITERATURE
Smith, E. F. A new disease of wheat. Jour. Agr. Research 10: 51-53
pl. 4-8. .2 Jl 1917.
Smith, R. E. A new apparatus for aseptic ultrafiltration. Phyto-
pathology 7: 290-293. f. 7, 2. 7S 1917
Spaulding, P., & Gravatt,G.F. Inoculations of Ribes with Cronartium
ribicola Fischer. Science Il. 46: 243, 244. 7 S 1917.
Spaulding, P., & Pierce,R. G. State and national quarantines against
the white pine blister rust. ‘Phytopathology #741G9,'3205 9 BS 1917:
Spegazzini, C. Espigando en el herbario. Ann. Soc. Cien. Argentina
82: 217-232. D nate [1917] {Illust.}
Includes new genera: Chiovendae, Cavaraea and Piroltantha and the new species’
Portulaca pha coe 7 eae uypolencs, Cavaraea elegans, Pirottantha modesta,
Jaborosa leptophylla and J. oxipetala.
St , E. C., & Piemeisel, F. J. Biologic forms of Puccinia gra-
minis on cereals and grasses. Jour. Agr. Research 10: 429-496.
pl. 53-59. 27 Au 1917.
Standley, P. C. The Chenopodiaceae of the North American Flora.
Bull. Torrey Club 44: 41 1-429. 155 1917.
Standley, P. C. New East African plants) Smithsonian Misc. Coll.
68: 1-20. 23 J 1917.
Includes numerous new species.
Stevens, H. E. Report of the plant pathologist. Florida Agr. Exp.
Sta. Report 1916: 66 R-79 R. f. 10, 11. My 1917
Contains mention of lightning injury to citrus trees as well as discussion of citrus
Ss.
Stevens, O. A. Plants of Manhattan and Blue Rapids, Kansas, with
dates of flowering—I]. Am. Mid. Nat. 5: 98-112. Jl 1917;— III.
Am. Mid. Nat. 5: 113-129. S$ 1917.
Stevenson, J. A. Lightning injury to sugar cane. Phytopathology 7:
ty, S16. ff. 20° 9S 0987.
St. John, H. Remarks on several North American species of Alope-
curus. Rhodora 19: 165-167. 10§ 1917.
Stout, A. B. Fertility in Cichorium Intybus: the sporadic occurrence
of self-fertile plants among the progeny of self-sterile plants. Am.
Jour. Bot. 4: 375-395. f. 12,2. Jl 1917.
Sudworth,G.B. The pine trees of the Rocky Mountain region. U.S.
Dept. Agr. Bull. 460: 1-47. pl. 1-28 + maps 1-13. 26 My 1917.
Swingle, W. T. The botanical name of the lime, tobe aurantifolia.
Jour. Washington Acad. Sci. 3: 463-465. 4 N 19
Swingle,W.T. Citrus and Poncirus. In Sargent, ‘“ S., Plantae Wil-
somianae 2: 141-151. 24 Mr 1914.
Swingle, W. T. eetvoes, clonetypes, and spermotypes, means for
botanical type specimens. Jour. Washington Acad.
Sci. oe Saiesl a Wt ,
BULLETIN
OF THE
TORREY BOTANICAL CLUB
FEBRUARY, 1918
On the constancy of cell shape in leaves of varying shape
LILLIAN A. TENOPYR
(WITH ONE TEXT FIGURE)
INTRODUCTION
Since the publication of Sanio’s observations on the size of the
wood cells of Pinus sylvestris, the subject has aroused considerable
interest, as is shown by the work of Amelung, Sachs, Strasburger,
Gates, Keeble, Neilson Jones, Jakushkine and Wawilow, and Sierp.
Among the zoologists who have investigated the same problem are
Gaule, Donaldson, Hardesty, Levi, Morgan, Driesch, Rabl, Cham-
bers, Popof, Berezewski, Jennings, and Conklin.
Sanio (1872) shows that in the Scotch pine, the wood cells
attain a definite (final) size, which is constant for the following
annual rings. He finds, however, that the size of the wood cells
does vary according to the height at which they are situated in the
stem. The size of the wood cells increases as the stem is ascended,
until a maximum is reached, then decreases toward the apex. The
branches always have smaller wood cells than the main stem at
the level at which the branch arose; but branches which arise from
that portion of the main stem which had the largest wood cells,
have also larger wood cells than the branches situated higher or
lower on the stem.
Amelung (1893) studied cell size in its relation to the size of
the organ. He raised the question whether differences in the size
of homologous organs within a species are accompanied by corres-
[The BuLLetin for January (45: I-50) was issued February 8, 1918]
51
52 TENOPYR: CONSTANCY OF CELL SHAPE
ponding variations in cell size, or whether the cell size of each
‘species is a constant and distinct character. That is, do environ-
mental conditions, which modify the size of plant organs, affect
the size of their cells? Are giants and dwarfs due to greater or
lesser development in size of the individual cells, or to more or less
numerous cell divisions?
Amelung estimated the size of the various cells of plants having
homologous organs differing in size. He made ten longitudinal
sections, or, in other plants, ten cross sections of each of the organs
to be compared. These he mounted in water or in glycerine. In
each section he counted the number of cells of a given tissue that
were intersected by a line on the stage micrometer, a line usually
I mm., sometimes 5 mm., in length. He thus obtained ten cell
counts on each organ. From these he determined the average
number of cells to 1 mm. and the average length or average
breadth of the cells of a given tissue both of the giant and of the
dwarf organs. He compared the palisade and epidermal cells of
large leaves with similar tissue cells of small leaves, the wood cells
of poorly developed and of well-developed shoots, and the paren-
chyma of large and small fruits. The organs compared were
taken from the same plant, or from different plants of the same
species, or from closely related species. He found that the cell
size of a given tissue of an organ is constant for the species, re-
gardless of the average size of the individual or of the organ.
Thus, the epidermal cells of a small leaf of Ficus macrocar pus
were as large as those of a leaf twice as long and broad; the
wood cells of a twig of Vitis vinifera, 6 mm. in thickness, measured
as much in cross section as those of another shoot, only half as
thick; a leaf of Victoria regia, measuring 900 X goo mm., and
one of Nymphea alba, 190 X 190 mm., had cells that were identical
in size.
Amelung concludes that not the cell size, but the cell number,
determines the size of an organ.
Sachs, with whom Amelung worked, confirms the statement
that the cell size of a given species is constant, and is not influenced
by body size.
Strasburger (1893), in a series of studies on the relation of
we nuclear and cell size in the embryonic cells of the growing points
TENOPYR: CONSTANCY OF CELL SHAPE 53
of stems and roots of many species of plants, reached the same
conclusion, regarding the size of these cells in large and small indi-
viduals of the same species, although varieties of the same species
- may differ greatly as to their cell size.
Conklin (1912) has made an elaborate and extended study of
cell size compared with body size in the genus Crepidula. He
found great variability in the body size of these gasteropods. In
Crepidula plana, there are dwarf and normal females. These
latter are larger than the normal-sized males. The average size
of individuals of the same sex differs greatly for the different
species. Yet, in spite of all these differences in body size, the tissue
cells of corresponding organs or parts of organs were in ‘general of
the same size in all the adult animals examined.
The muscle fibres and ganglion cells formed an exception to
this rule. In Crepidula plana, the size of these cells was greater
in the largest animals. This, he notes, agrees with the observations
of Gaule, Donaldson, Hardesty, and Levi, who measured these
cells in frogs and various mammals. In connection with the size
differences found in these cells, Conklin cites the observation made
by Levi, that these cells cease to divide early in life, whereas such
cells as the epithelial and the gland cells, where Levi found no
correlation with body size, continue to divide throughout life.
The size of the sex cells, also, varied in the different species.
In general, the smaller species produced larger eggs. These re-
sulted in larger embryos, having larger cells. Conklin says the
fact that the species with smaller eggs produce, in the end, larger
adults, and that the final cell size is almost the same for all the
crepidulas, is due to a longer duration and more rapid rate of cell
growth and division in the larger species.
A very notable case bearing on the question of the constancy
of cell size in a species, is that of Oenothera gigas, a so-called mutant
of Oenothera Lamarckiana, with double the number of chromo-
somes found in the parent species. Gates (1909) found that the
cells of Oenothera gigas, which have twenty-eight or twenty-nine
chromosomes, were conspicuously larger than the cells of Oeno-
thera Lamarckiana, whose chromosome number i is only fourteen.
Boveri (1905), on the basis of extended studies on sea-urchin
larvae, formulated the law that the cell size in sea-urchin larvae
\
54 ‘TENOPYR: CONSTANCY OF CELL SHAPE
is directly proportional to the number of chromosomes. Accord-
ing to this law, one would expect to find here a direct relation be-
tween the amount of chromatin, the nuclear size, and the size of
the cell. Gates (1909) does not find that this holds equally for
all the tissues. While the chromosome number of Oenothera gigas
is double that of Oenothera Lamarckiana, the cells of the former are
not uniformly twice as large as those of corresponding tissues of
Oenothera Lamarckiana.
Gates estimated the size of rectangular cells, as those of the
epidermis, by multiplying the three dimensions. The length and
breadth of the cells he estimated from measurements made on
camera-lucida drawings, magnified about 1,380 diameters, the
third dimension being considered as identical with the length of
the cell. By length of the cell he means the measurement along
the long axis of the organ, though this is not necessarily the longest
dimension of the cell; the width being the measurement at right
angles to the surface of the organ, that is in the direction of the
thickness of the organ in which it occurs. The epidermal cells of
the petals of O. gigas were 1.9 times as large in volume as those
of O. Lamarckiana, which is closely in accordance with Boveri's
law. However, other tissue cells depart widely from the expected
ratio of 2:1. Thestigmatic cells, the cells of the anther epidermis,
and the inner wall of the anther, were more than three times as
large in O. gigas as in O. Lamarckiana, while the pollen mother
cells of the former, both during synapsis and in the reduction divi-
sions, were only 1.5 times as large. :
Moreover, the increase in size of the O. gigas cells was not
equally great in all dimensions. The epidermal cells of the anther
increased 72.8 per cent. in length, and only 28.4 per cent. in width;
the stigma cells increased 51.9 per cent. in length, and 32.2 per
cent. in width. The anthers of O. gigas are approximately twice
as long as those of O. Lamarckiana. Gates believes that both have
approximately the same number of cells, ‘‘and that the greater
length of the O. gigas anthers is accounted for by the greater length
of the individual cells.” Gates concludes that there is evidently
some regulating factor determining that the increase in length o
the epidermal cells shall be greater than the increase in thelr
in the anther epidermis, but less in the petal epidermis.
TENOPYR: CONSTANCY OF CELL SHAPE 55
Similarly, Keeble (1912), ina comparative study of the cells of
the stem and leaf tissues of White Queen Star, a horticultural
variety of Primula sinensis, and those of its mutant, Giant White
Queen Star, showed that the latter is a giant because its cells are
larger and not because they are more numerous than those of the
parental type. Indeed, in the cortex of Giant White Queen Star,
Keeble found fewer cell layers than in the normal type. The meas-
urements he gives for the cortical cells of the flower peduncles, taken
for the layer immediately external to the endodermis, are:
Radial measurements..........:..5.... g:m:: 100: 48
Tangential measurements.............. £2 :- 100; BI
Longitudinal measurements............ £72 032: 1002 S97
The cells are larger in all three dimensions, radial, tangential
and longitudinal, and the gigantism is common to all tissues. He
does not state how many cells were measured nor does he describe
his methods. The chromosome number in both varieties is the -
same. The nuclei of the giant form are larger, as Keeble shows in
his drawings of pollen grains, but he does not give any measure-
ments of them. It is therefore impossible to tell, in the case of
White Queen Star and its giant mutant, Giant White Queen Star,
whether or not the amount of chromatin material is the factor de-
termining the relative cell size in these plants.
In Avena sativa, Jakushkine and Wawilow (1913) found that
different varieties may have different-sized cells, but they were
unable to discover any relation between the size of either the
stomata or the epidermal cells, and that of the leaf surface, when
varieties with different-sized organs were compared. For in-
stance, a small-celled variety had the largest leaves. Like Sanio
these authors found that the cell size of the leaves varied according
to their position on the stem, the cells of the highest leaf being
smaller than those of the third leaf from the topof the stem. They
- found that in any given variety, the cell size of any tissue at a
definite part of an organ was a constant character, and further
that each of the seventeen pure lines obtained through selection
from a mixed population of German and west Russian oats, fell
into one of two groups, namely:
1. A large-celled group, the average length of the stomata of
the highest leaf being 0.063 mm., that of the third leaf down being
0.0735 mm.
‘
56 TENOPYR: CONSTANCY OF CELL SHAPE
2. A small-celled group, the first leaf from the summit having
stomata whose average length is 0.054 mm., those of the third leaf
being 0.067 mm. long.
In his paper on species hybrids of Digitalis, Neilson Jones
(1912) does not give any measurements of the cell size of D. grandi-
flora and D. purpurea; but his general observations are most in-
teresting. He finds that D. grandiflora, which has smaller leaves
than D. purpurea, has larger leaf cells. Hybrids of these two
species are intermediate in both external and cytological charac-
ters, but in both respects there is a tendency to favor the seed
parent. Thus, in the hybrid resulting from the cross | hee ae Gas
¢ the leaves are somewhat larger and the cells smaller, than they
are in the progeny of the reciprocal cross. Evidently the size of
the leaves and cells of the hybrids cannot be due to heterozygosity,
since in that case we would expect the reciprocal crosses to have
the same effect on the leaf and cell size of the resulting hybrids.
One of the most careful studies of the relation between body
size and cell size is that of Sierp (1913). _He made measurements
of the cells of various tissues of dwarf and normal plants, the
dwarfness being in some cases a fluctuating characteristic, in other
cases hereditary. Under the first group come dwarfs of Panicum
sanguineum, Draba verna, Aethusa Cynapium and Urtica dioica.
This type of dwarfs always had cells more or less reduced in size.
Some of the differences in cell size were very slight. True, that is,
hereditary dwarfs, he found were of three kinds:
1. Those having smaller cells than the normal plants, as certain
dwarf varieties of Solanum tuberosum, Pisum sativum, Clarkia
pulchella, and Zea Mays.
2. Dwarfs with cells slightly smaller than, or almost the same
as, those of the normal type, as those of Lathyrus odoratus, Mira-
bilis Jalapa, Phaseolus vulgaris, and Lens esculenta (Rote Kleine
Winter).
3- Dwarfs with larger cells than found in the normal plants,
as Nigella damascena nana.
Sierp found great variability in the cell size of each tissue,
though the average value was quite constant. He lays great
stress on the importance of measuring exactly corresponding
_ places, when comparing cell size of any tissue of the stem or leaf,
~
TENOPYR: CONSTANCY OF CELL SHAPE 57
since it varies according to the height of the stem. Moreover, in
each leaf, the cells at the apex, at the base, and midway between
these two regions differ in size.
Sierp used the Schwendener-Ambronn method for finding the
average cell size. This method gives the average area of the cell
from a surface view, or of its cut surface in longitudinal or cross
section. A camera-lucida drawing is made of a group of the cells
of the epidermis, or of the cells of any other tissue, of each of the
organs compared. These drawings are made on a piece of Bristol
board of as uniform thickness as possible, the area and weight of
this piece of Bristol board being carefully noted. The drawing
of the group of cells is then cut out and weighed also. The product
of the weight, times the area of the original piece of Bristol board,
divided by the weight of the camera-lucida drawing, gives the area
of the group of cells drawn. This area, divided by the number of
cells in the group, gives the average area of the cell. The area of
the organs compared is ascertained in a similar way.
Possibility of error lies in the uneven thickness of the Bristol
board, in its absorption of moisture from the atmosphere, and in
the lack of precision in cutting inside or outside of, not through,
the outline of the group of cells. However, as many tissue cells,
especially those of the epidermis, are very irregular, this method of
determining cell size would seem preferable to that of basing one’s
comparisons on the average length or average width of the cell or
on the product of ‘these two dimensions, as if the cells were indeed
rectangular.
The work of Sanio, Amelung, Jakushkine and Wawilow, and
Conklin seems to warrant the conclusion that while the cells
of an organ may differ greatly in size for various and obscure
reasons, the average cell size of an organ is constant for the species
or variety. The work of Gates, Keeble, Neilson Jones and Sierp
shows that hereditary differences in body size of varieties or species
may be due to corresponding or reverse differences in cell size, in
cell number, or to both these factors.
THE PROBLEM OF CELL FORM
The further questions suggest themselves: Is there any correla-
tion between the shape of the plant organ and the shape of its con-
58 TENOPYR: CONSTANCY OF CELL SHAPE
stituent cells? Are the long narrow stem leaves of many plants
due to the length and narrowness of their cells as compared with
those found in the more rounded basal leaves, or to a greater num-
ber of cell divisions in the long axis of the leaf? Are the cells of
the lobes of incised leaves wider than those of the constricted por-
tions of the leaves, or are the constrictions the result of a slower
rate or shorter duration of cell division?
Gates (1909) seems to think such a direct relation between cell
shape and leaf shape is probable. “Increase [in cell size],’” he
says, “has been greater in one dimension than in another, resulting
in a change in the relative dimensions of the cells. This in all
probability accounts for the altered shape of some of the organs,
as leaves and capsules.” He holds that the greater cell size, to-
gether with the difference in cell shape of O. gigas, is sufficient to
have produced external differences between the two plants, with-
out the introduction of any new factors. Except for stating that
the anthers of O. gigas, which are about twice as long as those of
O. Lamarckiana, have cells which are not only larger but rela-
tively longer than those of the parent plant, Gates does not give
any comparative measurements on corresponding dimensions of
organs and their cells, to prove the relation of cell shape and body
shape.
According to both Familler (1900) and Goebel (1908) there is
a direct relation between the light intensity and the form of leaf
produced in Campanula rotundifolia. Plants placed where they
were well shaded, instead of developing typical linear stem leaves,
produced only round leaves on the stem, similar to the basal leaves
which appear earlier in spring.
In my cultures of Lobelia Erinus, plants grown in the green-
house during the dull winter months flowered less profusely than
during the summer. These winter plants had only spatulate
leaves along the whole length of the floral shoot and none of the
small linear leaves which are usually found on the upper part of the
stem.
Familler’s explanation for such phenomena is that the light,
food supply, and other optima are higher for the production of the
flowers and the long leaves than for the production of the round
leaves. Thus, it is not until the warm weather of early summer,
TENOPYR: CONSTANCY OF CELL SHAPE 59
when the illumination is stronger and the plant already possesses
a well developed root system and numerous basal leaves, that the
narrow stem leaves and flowers appear. Even then, a diminution
of the light intensity, or any great disturbance in the development
of the plant can occasion the production of the juvenile form of
leaf. Thus, if the Campanula plant is propagated by cuttings from
shoots bearing only long narrow leaves, the young leaves sent out
by these plants are round, like the basal leaves of the parent plant.
These propagated plants cannot produce long leaves until they
have better developed root systems, and the plants have become
well established in their new food relations.
I have studied the shape, i. e., the relation of length to breadth
of leaves and their constituent cells, in the following three types of |
plants.
I. Species with broad basal leaves, narrow stem leaves near the
inflorescence, and transitional leaves on the lower part of the stem:
Campanula rotundifolia, Lobelia Erinus.
2. Broad-leaved and narrow-leaved species belonging to the
same genus: Plantago major and P. lanceolata, Linum angustifolium
and L. usitatissimum.
3. Varieties of the same species having entire leaves, as com-
pared with others having lobed leaves: Cichorium Intybus.
METHODS
The Schwendener-Ambronn method used by Sierp, which gives
the average area of the cells, was not suited for my investigations
on the relation of the length to the breadth of the cells of variously
shaped leaves. The method used in finding the average length
and the average breadth of the cell resembles that of Amelung,
except that an ocular micrometer instead of a stage micrometer
was used. The number of cells to a unit of the scale in the ocular
micrometer were counted, and the average dimensions in milli-
meters were estimated from these numbers. Whenever the line
of measurement passed through the two opposite sides of a cell,
as it appeared in the section regardless of how small a fragment of
the entire cell was thus cut, its measurement was recorded as the
measurement of the length or of the breadth of the cell in that par-
ticular portion of the leaf. If, as sometimes happened in the
irregular cells of the epidermis, the same line ‘passed twice through
*
60 TENOPYR: CONSTANCY OF CELL SHAPE
any cell, the two fragments were recorded together as the measure-
ment of the cell in that region. Frequently the end of the line of
measurement did not reach the opposite wall of a cell. In such
a case I estimated as accurately as possible what fractional part -
of the length or of the breadth of that cell fell upon the line of
measurement, and this fraction was recorded. Thus, if the line
passed through nine cells and reached only one fifth of the way
across the tenth cell, the cell count was recorded as nine and one
fifth cells. This method cannot of course be regarded as giving
absolute length and breadth of the epidermal cells especially, but
is probably the best available. It seems on the whole preferable
to record the fractional parts of cells lying on the scale unit. The
_ alternative would be to make all the measurements on single cells,
“and wh this method the problem of selection and the errors in deter-
mining the cell axes would probably involve still greater inaccu-
racies.
The maximum length and width of each leaf examined was
recorded, and an outline tracing was made. In most cases one
hundred counts were made on each type of leaf as the basis for
comparison, with the exception of Linum, in which case I made
nineteen counts on the palisade cells and forty-five counts on the
epidermis, and of the palisade cells of Plantago, on which thirty
counts were made.
Except in the comparison of leaf shapes in lobes and constric-
tions, the cell counts were made in the middle region of the leaf,
and about 2 mm. away from the midrib. As the comparison was
between leaves differing from each other in length and breadth,
only these two dimensions of their cells were measured. I have
not concerned myself with the third dimension, as the difference
in thickness of the leaves is too small for easy study.
All the measurements were made from the surface of the leaf.
The epidermal cells of the under surface of the leaf were measured
from below. The palisade cells were measured from above.
Thus in any one portion of a given tissue I could make measure-
ments of the length and of the width of the cells at the same time.
The length of the cell I considered to be that dimension of the cell
which was parallel to the midrib of the leaf and the long axis of
the entire leaf, the width of the cell as taken was the dimension
to the width of the leaf.
TENOPYR: CONSTANCY OF CELL SHAPE 61
I found that, if the sections of the lower epidermis to be meas-
ured were prepared by stripping this tissue away from the leaf,
it was difficult to determine which axis of the cell had been parallel
to the midrib. The possibility of error in this respect was greatest
in cells having irregular outlines. I therefore prepared these sec-
tions by removing the upper epidermis and the green tissues from
that portion of the leaf which was to be examined, leaving the
lower epidermis intact. The entire leaf, or the exposed epidermis
with a portion of the midrib, was then mounted in water and
placed under the microscope. Before measuring the palisade
cells, both the upper and the lower epidermis of the region to be
examined were stripped off, care being taken to remove none of
the green tissue. When the leaf was placed flat on a slide the
remaining tissues were sufficiently transparent to enable me to
make measurements of the two dimensions of the upper ends of the
palisade cells. All the measurements were made on the living
cells, thus avoiding the possibility of shrinkage or distortion due
to fixation.
I found an ocular micrometer preferable to a stage micrometer,
as, by moving the slide or by rotating the micrometer, I could
more easily bring the specimen to be examined in the proper posi-
tion in relation to the scale, without handling the specimen itself.
Throughout the investigations I used a Leitz ocular microm-
eter. The scale consists of a large square, divided into four
smaller squares, one of which is again divided into twenty-five
equal squares. The value of these divisions of course depends
upon the magnification, and it was ascertained with the aid of a
stage micrometer for each eye-piece and objective used. For
example, when a one-inch ocular and two-thirds objective were
used with a Bausch and Lomb microscope, each side of any one
of the smallest squares was equal to 0.116 millimeter. All cell
counts on all the plants examined are stated in terms of cell
diameters per millimeter.
The section to be measured was placed on the stage of the
microscope in such a position that the midrib of the leaf was paral-
lel to one side of the ocular micrometer scale.
In measuring the palisade cells, a favorable area of this tissue,
uncrushed and free from veins, was chosen, equal to one of the
*
62. TENOPYR: CONSTANCY OF CELL SHAPE
smallest squares of the ocular micrometer. The number of cells
cut by each side of the square was counted. The length of this
line was then divided by the cell number to obtain the average cell
diameter. In this way, with one placing of the micrometer, I
could make four measurements, two giving the length and two
giving the width of the palisade cells of the region.
To measure the epidermal cells, a suitable area, free from veins,
was selected. To avoid errors due to the presence of stomata,
care was taken in each case to so place the micrometer that the
stomata came between but not on the lines of measurement. In
plants having numerous stomata, this was often a difficult matter.
In this case I used twice as large a micrometer unit, the sides of
two squares, corresponding roughly to the larger size of the cells.
CAMPANULA ROTUNDIFOLIA
Campanula rotundifolia is well known as a plant whose radical
and cauline leaves differ widely in shape. It produces numerous
radical leaves, which are often wanting at the time of flowering.
These are petioled, almost orbicular in shape, with cordate base.
In May or June the plant sends up several shoots which bear
flowers at their summits throughout thesummer. The upper stem
leaves are sessile and linear, but the lower stem leaves are inter-
mediate in shape, being ovate and acute. These three types of
leaves are designated in the tables as basal, transitional, and linear.
Measurements were made of the length and the width of the
cells of the lower epidermis of typical leaves of the three forms
described. TABLE I gives the results of these measurements.
The average number of cells in each leaf that were cut by a line
I mm. in length, is given for each of six leaves of each shape. The
final averages are from the original data and differ fractionally
from the average of the averages in the following table.
A comparison of the average number of cells to 1 mm. found
in the long and the broad axis of each basal leaf shows that there
is some variation in the size of the cells, for example, in leaf C the
average number being 22.02 cells to 1 mm. in the long axis and 20.91
cells in the transverse axis, while in leaf D the cell counts were
26.95 cells in the long axis and 29.56 cells in the transverse axis of
the leaf. These figures also serve as an example of the variability,
TENOPYR: CONSTANCY OF CELL SHAPE 63
found in the relative dimensions of the cells, the cells in leaf C
being somewhat broader than long, while in leaf D this relation of
length to breadth of the cells is reversed. This variability in cell
TABLE I
CAMPANULA ROTUNDIFOLIA
Comparative length and width of the three types of leaves and the corresponding
dimensions of the cells of the lower epidermis of each type of leaf. Numerals in paren-
theses show number of counts. Line of measurement = 1 mm
Basal leaves —— Linear leaves
eaves
Length | Width | Length | Width | Length | Width
Leer An ooo. ee ae eee FY: FAVS 7123, 5.5 | §6: 3.
Pek ae Call Bite 626) 55.6.0) woes Ie 0.037) 0.036; 0.04 | 0.029) 0.033) 0.03
Average number of cells (15)......... 26.99 | 27.74 | 24.81 | 33.45 | 29.78 | 32.97
Leaf “~- be Shiv s eMeE pe ores Se Seles ti 5. ET: 49- 3.5. 1.32: a;
pata CRE GER CI eo a oie 0.046; 0.047) 0.038) 0.038) 0.036) 0.035
Average number of cells (15) ......... 21.54 | 21.38 | 26.02 . 92 | 27.75 | 28.
Leaf ~~ PEE ae ee ee eas eG iy Bs 18, 55. 39. 4.5
AVerage Cell Gige 125) on oa 0.045} 0.047} 0.035) 0. sap 0.033} 0.034
as. number of cells (15).......-- 22.02 | 20.91 | 28.52 _ 55 | 29.98 | 28.92
Cee EME ae ved s Vel Cates = ¥¥. 49. a
iets CEN Giz@ £96) 23.4% 5 ces ck 0.037| 0.033) 0.043) 0. ee 0.03 | 0.027
Average sees or cella (8a ys oo eis 26.95 | 29.56 | 25.95 | 26.56 | 32.28 | 35.77
ROM Be ee ees 10. 19:5 a7. 6. 50. -
Averaze cell size (15)... dis. Soe tc 0.042) 0.037) 0.035) 0.036) 0.032) 0.033
Average number of cells (15)......... 23.55 | 26.97 | 28.3 | 27.32 | 30.6 | 29.86
EPA oe Ee ale ee ee Gea 19. 19. ai. 3 57- 4.5
Soba COL OG 086) os se ve ee 0.037} 0.039) 0.037) 0.03 | 0.04 | 0,032
Aver: number of cells a Berea atar 26.74 | 25.62-| 26.61 | 32.36 | 24.7 | 30.31
‘hetiag e measurement of leaves....... £4.16 | 15.33. |32.5 a 47.16 | 3:33
Average rere ase of bake gee 0.04 | 0.039) 0.038) 0.034) 0.034 0.032
Average number of cells (100)........ ‘| 24.85 25-34 | 26.26 | 29.21 | 28.74 | 30-91_
Ratio dimensions of leaf. .-........:.. Of 2% 4.64:1 T4.15 $2
Ratio dimensions of cell. ..........-. toa: 2 ee Gee 4.07 71
size and cell shape is found in all three forms of leaves examined.
However, in most leaves as in the final averages, the numbers
indicate that the cells are nearly isodiametric or somewhat longer
than they are broad. The table also gives the average measure-
ments of the cells in fractional parts of a millimeter.
A glance at the ratio of the length to the width of the cells of
the lower epidermis of the three forms of leaves shows that the
percentage of difference between these two dimensions does not
approach that found between the leaf dimensions. The basal
leaves are on the average slightly shorter than broad, and their
epidermal cells are slightly longer than broad, almost isodiametric.
64 TENOPYR: CONSTANCY OF CELL SHAPE
The transitional leaf is more than four times as long as broad, yet
its cells are of almost the same shape as those of the basal leaf,
being slightly longer than wide. In the linear leaves, which are
about 14.15 times as long as they are broad, the cells are only .07
longer than wide. That is, the difference between the length and
the width is 287 times greater in the leaf than in the cell. More-
over this difference in cell dimensions is least in the basal leaves
and greatest in the transitional leaves. There seems to be no rela-
tion here between the shape of the cell and that of the leaf.
The slight difference between the ratios of length to breadth
of cells obtained in the different leaves is almost equaled by the
variability of the individual cells in any one leaf. In some leaves
the cell proportions, as compared with the leaf proportions, may
be found reversed in certain areas. Thus fifteen measurements of
cells in a basal leaf E, which was 10 mm. long by 12.5 mm. broad,
showed that the cells averaged 0.042 mm. long and 0.037 mm. wide
(a ratio of 1.13 : 1) while similar measurements of cells of a linear
leaf, 50 mm. by 3 mm., showed that they had an average length of
0.032 mm. and an average width of 0.033 mm. (a ratio of .97 : 1).
The forms of the three types of leaves found in Campanula rotundi-
folia are not therefore directly correlated with the shape of their
cells.
The cells of the stem leaves show a smaller average size than
those of the basal leaves, the cells of the linear leaves being most
reduced. In Taste II are given the relative length and relative
TABLE II
CAMPANULA ROTUNDIFOLIA
Relative size of the lower epidermal cells of the three types of leaves.
Length of Width of
cells cells
Transitional leaf : ei leaf . 04:1 86:1
Linear leaf : basal leaf... .......; 86:1 Ae Bede
width of the cells of the three types of leaves. The difference in
the dimensions of the three types of cells is very slight, and might
seem at first to fall within the limits of individual cell variability.
I therefore plotted frequency curves (Fic. 1) showing the number
of times that I found a given number of cells to a line of 0.232 mm.,
in the long and in the broad axis of the basal and the linear leaves.
A greater number of cells to a given line indicates smaller cell size.
TENOPYR: CONSTANCY OF CELL SHAPE 65
All cases where there were four and a fractional part of a cell
to 0.232 mm. were recorded as four cells—all cases of more than
five cells and less than six cells were recorded as five cells, etc.
These curves show that both in the long axis and in the transverse
Pt coe | es eS a ea a Ret Si ee
+4 ee AM Be 8 TT
A rT] |
A
in Ga Os Oe Ly
ITA
- |
+. |
1 |
1. |
|
i] |
i ‘ \
8 |
5 \
} i
avi \
9423456789 i0ili2 x q
A B
Fic. Curves denoting frequency of dimensions of lower epidermal cells of
ie seats rotundifolia , Transverse axis. B, Longitudinal axis. Abscissas
denote the number of bells to 0.232 sastitaien Ordinates record the n
instances in which a given number of cells per 0.232 mm. occu U cai lines
used for cells of basal leaf; lines broken by x’s, for linear stem leaf.
axis of the leaf, there are most frequently five cells to 0.232 mm. in
the basal leaf, while in the linear leaf there are six. The curves
prove conclusively that the decrease in both dimensions of the
lower epidermal cells of the linear leaves is regularly present in the
material studied.
This observation agrees with that made by Sierp, who found
that in Mirabilis Jalapa, Nigella damascena, and Pisum sativum
(Laxton’s Alpha) the higher the leaf was situated on the stem, the
smaller its cell size, as well as that of the node from which the leaf
sprang.
66 TENOPYR: CONSTANCY OF CELL SHAPE
Neither in the case of Campanula nor the other plants studied
did I find the size of the individual cells to be as constant as Ame-
lung’s measurements would indicate. Because of the wide range
of variation in cell size, the average cell size could be determined
only through numerous counts. Usually one hundred counts
were made on each form of leaf, so that the average cell size given
is the average size of from six hundred to seven hundred cells.
LoBELIA ERINUS
This plant, like Campanula rotundifolia, has three distinct
forms of leaves. The orbicular radical leaves are developed before
the flower shoots appear, and are rarely found at the time of
flowering. The slender, somewhat recumbent shoots bear at first
only spatulate or obovate leaves, but later, as the stem elongates,
it has small linear leaves near its summit. The flowers grow from
the axils of these linear leaves. Each type of leaf produced is not
only smaller than the one which preceded it but it is also propor-
tionately narrower.
A comparative study was made of the shape st the cells of the
lower epidermis of the three forms of leaves, in relation to the
shape of the whole leaf. Only those linear leaves were examined
in whose axils the flowers had already faded, as I thought these
leaves were probably fully grown. Measurements were made on
six such leaves after two intervals of two weeks each, to see if
they had ceased to grow by the time the flowers had bloomed.
There was some further growth, but scarcely enough to seriously
affect the proportions of the leaf, or noticeably alter its shape
(TABLE III).
TABLE III
LoBeELIA ERINUS
Record of further growth of linear leaves, after flowers in their axils had faded. Line
of measurement =
Plant A B c pcb pd se
| Length eit: a Length Length| Length
| width width width wk} eta | wih
First ca ag | 10 % 25 ¢ 36: FT te a ae 142 119 X S| 190 X4
End second week.....| 10 X 1.87) 11 X 1.5| 12 X 2.5/15 X2.5| — 20 X 4
End fourth week... ../11.5 X 1.87! 11 X 1.5 Died 16 X 2.5 121 X 9\20.5 X 4
TENOPYR: CONSTANCY OF CELL SHAPE 67
The results of the cell measurements made are given in TABLE
IV, which shows both the number of cells per millimeter and the
dimensions of the cell. The ratio te the length to the width of these
TABLE IV
LOBELIA ERINUS
Comparative length and width of the three types of leaves and the corresponding
dimensions of the lower epidermal cells of each type : leaf. Numerals in parentheses
show the number of counts. Line of measurement =
Linear leaves
Basal leaves Transitional |
} | leaves
"Length | Width | Length| Width Length | Width
COT AT Oe Se eV eae ee : 27. | 19. 9. 10.5 a5
Average cell size (15).............-.} 0.066] 0.053] 0.044) 0.048) 0.032) 0.033
Average number of cells (15)........-. 14.99 18.73 | 22.8 | 20.79 |30.9 | 30.11
Lea eRe Ss. . | at, &. 1%. 2.
oe cell size (15) veneees| 0.06 | 0.058 0.052] 0.043) 0.057) 0.04
Average number of Celis. (15). gers, aren 0 | 16.53 | 16.95 | 19.02 | 23.2 | 17.45 | 24.46
peg 8 ea eee | 34. St. t5. 9. ta: 3-5
Average cell ize (19) oe Se ei 0.063! 0.057 0.04 | 0.042) 0.045) 0.041
Average number of cells (15)......--. 15.76 | 17.45 | 24.63 #8 27 | 22.02 | 24.16
TRALEE ea ee eee 33: 30. 18. a 2.5
Averave céll size (5)... 5. as. 2 ee 0.094 0.068 | 0.054} 0. ae 0.042) 0.045
Average number of cells (5).........- 10:6 |15.48 | 18.18 | 21.33 | 23.49 | 22.17
Dal eae i eae ees oe 33. 29. 16. 5.5: 134: a
Avetaee Gert size (0) 0. oe os ee 0.067) 0.074 0.036) 0.035) 0.047) 0.033
Average number of cells (9). .-...---- 14.83 |13.37 | 27-42 | 28.07 | 21.27 | 29.98
Deel Boo Co igo ices ob es Ci 30. a5: 20. 7. 20. ne
‘Average Gell siz6(Q) 0a cee 0.085! 0.062) 0.046) 0.045) 0.051 0.046
Average a heuer of cells (9) 2024. s. 11.68 | 15.91 | 21.37 | 21-8 | 19.29 | 21.47
PGE ss ce Pees 5 ees 0. 31. 22. 10. 15. 3.5
— eel sige (12) 6. oa eae ee 0.06 | 0.079; 0.042} 0.038 0.04 0.033
Average number of cells (12).......-. 16.49 | 12.56 | 23-45 | 25-77 - 54 | 29.64
Feat yo 8 eh ig es ee es ee Va aay £5. 8. ce
pki td pel bine (PL) oe ee. cies are 0.056} 0.06 | 0.036) 0.039 0. ee 0.031
Ave number of cells (11). .-...... 17.84 |16.57 | 27.03 | 25-57 | 23-92 | 31-55
Le we * re ea Pe ou ee cae Cae any Al. 37: ans" ye 19. 5.5
Average ne. Fat) Cores Cee rae 0.064; 0.062) 0.043] 0.04 | 0.054 0.0
erage hes of cells ie) ey ete 15.48 |15.96 | 23-16 | 24.85 | 18.44 | he 08
Average measurements of leaves...... a ee 29. 18.88 | 7.94 | 14.25 |
Ave Ps a measur adie _ cl Oy 0.065} 0.062) 0.043) 0.042) 0. 044 >: 037
Average number of cells (100).....--- 15.15 ' 15.06 | 22.84 | 23.54 | 22.47 | 26.31
Ratio dimensions of leaf.........---- 2 2% ie a75 -%
Ratio dimensions of cell ee i ing ew Ray I.05: TOG ck (F372 F
cells by no means agrees with that of the corresponding dimensions
of the leaves. While the difference between the length and the
width of the linear leaves is almost four times greater than that of
the basal leaves, the cells of the linear leaves are relatively only .11
68 TENOPYR: CONSTANCY OF CELL SHAPE
longer than those of the round leaves. That is, the difference in
these two leaf shapes is almost forty times as great as the difference
in the shapes of the cells. Furthermore, the transitional leaves
have proportionately wider cells than the basal leaves.
There is a considerable range of variability in the shape of the
cells of any one type of leaf. For example, in a basal leaf, D, 33
mm. long by 30 mm. wide, the average length of the cells of the
lower epidermis was 0.094 mm., the average width 0.068 mm.,
while in another leaf, G, 50 mm. by 31 mm., the average cell
measurements were 0.06 mm. for the length and 0.079 mm. for the
width. The great range of variability of the cell shape of each
type of leaf may account for the slight difference between the
ratios of the cell dimensions of the round, the narrow and the tran-
sitional leaves.
The decrease in cell size, that is, in both length and width,
found in the linear leaves as compared with the basal leaves, is
even more marked than in Campanula rotundifolia. The ratios
between the corresponding dimensions of the cells of the three
forms of leaves are given in TABLE V. It will be seen that the
TABLE V
LOBELIA ERINUS
Relative size of the cells of the lower epidermis of the three types of leaves.
Length of Width of
cells cells
Transitional leaf : Basal leaf..... oat 07 21
Narrow leaf : Basal leaf......... 07.7 OO 2%
cells of the transitional and linear leaves, both of which grow
on the stem and are produced at almost the same stage of de-
velopment of the plants, have cells which are almost identical
in size, although differing considerably from those of the earlier,
radical leaves.
PLANTAGO MAJOR AND PLANTAGO LANCEOLATA
The leaves of the two species of plantain, Plantago major and
_ P. lanceolata, differ greatly in shape. The largest and best de-
veloped leaves of each species that could be obtained, were chosen
for comparison. P. lanceolata, when crowded, produces leaves
which are somewhat dwarfed, particularly in their breadth, being
TENOPYR: CONSTANCY OF CELL SHAPE 69
relatively only two-thirds to three-fourths as wide as is usual.
Such leaves also were examined.
Measurements were made of the cells of the lower epidermis of
such leaves of P. lanceolata, as well as those of the normal P. lan-
ceolata and P. major leaves. Also the palisade cells of the two
species of plantain leaves were compared. The results of these
measurements are given in cell size and:in cell number per milli-
meter in TABLE VI.
TABLE VI
PLANTAGO
Measurements of the length and the width of the leaves of two species of plantain,
and of the corres ponding dimensions of the cells of their lower epidermis and their palisade
tissue in transverse section, expressed in millimeters and in number of cells to one milli-
meter. B = average size of cells in mm. C = average number of cells to 1 mm.
Numerals in parentheses show the number of counts.
Leaf ell £7 pi A ‘mi:
Length Width Length Width
POI 6 i 176.09 110.06 B(100), 0.031 0.044
C(100), 30.09 22.63
MOG. Se ge po Bia hs I Ry A Srp WE
Ps hanceolatin 05 ee 310.3 32.63 B(100), 0.035 0.041
C(100), 27.97 23-97
Pt eG or a ee. D2 fe I ee Ee
P. lanceolata (narrow).. 153-5 10.5 B(100), 0.032 0.038
C(100), 30.71 25.81
Wate oe cs ee ay oe 14.6% I Ke
Leaf Palisade sae
Length Width Length Width
FM ee 48. 27. B(30), 0.022 0.021
C(30), 44.1 46.28
Mao... ele ey 3 eer I 1.04:
FP faneedlete.. eo i eae 70. 23. B(30), 0.021 0.021
C(30), = 92 46.79
Mat eee ot eee 3-04 I SE
In both species the palisade cells are quite round in transverse
section. The cells of the lower epidermis of P. major are slightly
shorter than those of P. lanceolata, but the disproportion does not
approach that between the length and width of the leaves of the
two species. The P. lanceolata cells are only 7 per cent. longer and
6 per cent. shorter than those of P. major.
I have plotted frequency curves for each species showing the
number of times that a given ratio of the length to the width of
the epidermal cells occurred. It would be difficult to determine,
70 TENOPYR: CONSTANCY OF CELL SHAPE
from curves as irregular as these, the mean ratio of length to width
in either species.
This does not mean that the shape of the lower epidermal cells
is the same in both cases. While the difference does not lie in the
length and width of the cells, their outlines are quite different,
the Cells of P. major being quite wavy in outline, while in both
types of leaves. of P. lancéolata the cells are only slightly wavy in
outline. The cells of the normally developed and the narrow leaves
of Plantago lanceolata are practically the same in shape and size.
LINUM ANGUSTIFOLIUM AND LINUM USITATISSIMUM
The leaves of Linum usitatissimum are lanceolate, those of
L. angustifolium are oblong or oblong-elliptical. The leaves of
both species are sessile, acute and entire. Those of L. usitatissi-
mum are about as broad, but at least twice as long as the leaves of
L. angustifolium. The cells of the lower epidermis and the pali-
sade cells were measured in each species. The measurements
show a striking similarity in the cell shape of these two species of
Linum. The cells of Linum usitatissimum are only .o4 longer
than those of L. angustifolium, and show no relation therefore to
the greater difference in their leaf dimensions.
CicHorium INTyBUS
The first leaves of the common type of Cichorium Intybus are
entire but these are soon succeeded by other radical leaves which
are distinctly lobed. The degree of lobing varies greatly in dif-
ferent plants, but the leaves of any one plant are quite similar in
shape. For this reason all the leaves of Cichorium Intybus whose
cell shape was measured were selected from one plant, which had
deeply constricted leaves. The Witloof type of Cichorium Intybus
is a cultivated variety of chicory, which has large and relatively
entire leaves. All the leaves of this type studied were taken from
the same plant. These, as well as the leaves of the common type
of Cichorium Intybus with which they were compared, were basal
leaves. In both, the cells of the lower epidermis near the midrib
and near the margin of the broadest part of the leaf, and midway
between these two regions were measured and their two dimen-
sions compared with the length and breadth of the leaves to which
they belonged (TaBLe VII).
TENOPYR: CONSTANCY OF CELL SHAPE 71
TABLE VII
CICHORIUM INTYBUS, COMMON TYPE
Maximum length and width of each leaf, the width of each leaf across ihe lobes, and
the constricted portions, and the len eh vate idth of the cells of the lower epidermis in
these two regions of the leaf. A = width of leaf from midrib lo margin. B = average
size of cells in mm.. C = average ais of cells to 1 millimeter. Numerals in paren-
theses show the number of counts.
| Lobe below this constricted portion
Vent Size Constricted ni eso
pera: Near Midrib | Midway out | Apex of lobe
. from midrib
Length | Width Length | Width Length | Width | Length| Width | Length | Width
A 2. | 28.
250 | 95 B (15) 0.36 | 0.29 | 0.03 | 0.028} 0.33 | 0.03
© GS) fav.s7 [aa a2 | 34.6 30.11 | 32.48
A 4 28.
220 61 B (15) 0.033) 0.027} 0.028) 0.029! 0.03 | 0.029
C (15) |29.48 | 36.8 (35.65 | 34.28 | 32.09 | 33.8
A 6. £5)
135 53 B (15) 0.025|} 0.029) 0.026) 0.027| 0.034) 0.023
C (15) |38-55 | 34-12 |37-81 | 35-05 | 29.22 | 42.28
A 5-5 34-
186 65 B (15) 0.037} 0.027| 0.03 | 0.03 | 0.032) 0.027
C (15) | 26.43 | 35-85 |33-21 | 32.94 | 31-03 | 35-95
A 4. 29.
229 vee B (15) 0.033} 0.028! 0.033} 0.028) 0.029) 0.027| 0.028) 0.026
C (15) |29.42 | 34-99 |29.35 | 35-47 | 33-5 | 36-1 | 33-56 | 37-45
A A. 32.
153 57 B (15) 0.033} 0.029| 0.029| 0.028) 0.034) 0.028) 0.027) 0.029
€ (15) |30- 34-11 (34-02 | 34.68 | 29.36 |34.6 | 36.52 33-69
A 4. at:
I51 55 B (10) 0.034, 0.025| 0.03 | 0.032} 0.026) 0.026
C (10) |29.24 | 39.46 |32.92 | 30.41 | 37-21 | 37-07
: Av. A 4.5 26.
Av. 189|/Av.65|Av. B(100)| 0.033) 0.028) 0.029) 0.020) 0.031 0.027
v. C(100)| 30.14 | 35-92 |33-7 | 34-23 | 31-62 | 35-99
Ratio dimensions of cell...... 5 ee Sy a Orc t TisttT
The lower epidermal cells of the eee ones of each leaf
4 -
were compared with those of the i ly adjacent
toit. The ratio of the average length to the average width of the
cells is .o4 greater in the constricted portion than in the middle.
region of the lobe, and .16 greater than in the lobe near the midrib.
This rather slight difference is apparently due to the fact that the
72 TENOPYR: CONSTANCY OF CELL SHAPE
cells in the constricted portions of the leaf are longer, rather
than narrower, than the cells of the lobes, whereas in the
leaf as a whole the difference is plainly one of greater breadth of the
leaf in the lobed portion than in the constricted portion, the length
TABLE VIII
CICHORIUM INTYBUS, WITLOOF TYPE
Maximum length and width of the leaves of a cultivated chicory and tageccan :
dimensions of the cells of the lower epidermis in different regions of the dth
of leaf from midrib to margin. B = average size of cells in mm. C = average Coke
of cells in 1 millimeter. N seat in parentheses show the number of counts.
Broadest part of blade
Leaf siz Basal portion
aes cates Near midrib Midway out Marginal
from midrib
Length | Width Length | Width | Length | Width | Length} Width Length | Width
A 6. 45-.
350 92| B (15) 0.062} 0.029) 0.032} 0.032] 0.038} 0.026] 0.027| 0.024
C (15) | 16.01 | 33-98 | 30.35 | 30.87 26.15 37.18 | 35-82 | 40.5
A ace 57.
386) 105 B (15) 0.059! 0.027; 0.03 | 0.039] 0.032} 0.034
C (15) | 20.04 | 36.59 | 32.43 | 25.5 | 30.98 | 20.34
A by SE.
352\ +02 .B (15) 0.055| 0.03 | 0.033) 0.033} 0.034) 0.033
C (15) | 17-80. | 32.43 | 290.52 | 29.93 | 29.13 | 30.05
A 6. 48.
412| 104 SB (15) 0.053} 0.029} 0.036) 0.04 | 0.037| 0.036] 0.037| 0.037
C (15) | 18.85 | 33.71 | 27-75 | 24-67 | 26.63 | 27.66 | 26.94 | 27-01
A 8. 50.
472 94 B (15) 0.066| 0.026) 0.032) 0.037} 0.031] 0.032
C (15) | 14-95 | 37-17 | 30.65 | 26.42 | 31.29 | 30.69
A 8. 49.
425| 107 B (15) 0.062} 0.03 | 0.041, 0.034 0.033| 0.033} 0.029) 0.026
C (15) | 15.92 | 32.42 | 24.06 | 28.58 | 30.04 | 29.43 | 33-5 | 37-68
A 9. Si.
350| 104 B (10) 0.041} 0.035 0.03 | 0.03 | 0.035! 0.036] 0.032! 0.031
C (10) | 24.01 | 28.28 | 24.02 | 32.05 | 28.49 | 27.26 | 31.07 | 31.84
Av.A 9.07 49-42
Av. 404/Av. 99) Av.C (100), 0.055! 0.029 0.034) 0.035) 0.034! 0.032
Av. 4 ti 17.95 | 33-76 | 28.62 | 28.19 a8 08 bee i
Ratio dimensions of OO orci 7.885% 08 >t t.04:t
of the two regions being the same, or even less in the constricted
portion. The cell shape in the wild chicory leaf is practically the
same both in the lobes and the constrictions of the leaf.
TENOPYR: CONSTANCY OF CELL SHAPE 73
The cells near the base of the Witloof chicory leaf are appar-
ently modified in their shape by their relation with the numerous
veins. ;
The lobing of the wild chicory leaf is not due to greater width
of the cells in the lobes as compared with the constricted areas,
but rather to the greater number of cells in the long axis of the
lobe, which is almost at right angles to the midrib. This greater
cell number is apparently due to a more rapid rate of division in
the lobe cells as compared with those in the constricted regions
(TABLE VIII).
The cells of the leaves of the Witloof variety are larger than
those of the wild chicory leaves, but the shape of the cells is almost
the same in both varieties. The ratio of the length to the width of
the Witloof cells is 1.01: 1, that of the wild chicory cells being 1.11: 1.
SUMMARY AND CONCLUSIONS
Cell size
The cells of the plants above described show a considerable
variability in size in the same tissue but that the average cell
size for any one tissue of a species or variety, however, is a fairly
constant and hereditary character, has been previously shown by
Sanio, Amelung, Sachs, Strasburger and Conklin.
The cell size of closely related species, as Linum usitatissimum
and Linum angustifolium, may be the same, which agrees with
the conclusions of Amelung and Conklin, or again, in closely re-
lated varieties of the same species, as the common and Witloof
types of Cichorium Intybus, the cell size may differ considerably,
as was also proved tobe true incertain plants investigated by Gates,
Keeble, Neilson Jones, Jakushkine and Wawilow, and Sierp.
As Sanio, Jakushkine and Wawilow, and Sierp have already
shown, the cell size in an organ depends in some degree on the
stage of development of the plant at the time the organ was pro-
duced. Thus the transitional leaves of Campanula rotundifolia
and Lobelia Erinus which are developed after the basal leaves:
have smaller cells, while the linear leaves, which appear latest,
have the smallest cells. In both these plants, the average cell
size of each type of leaf is a constant characteristic. Differences
in the size of any given organ of a species are due to differences in
74 TENOPYR: CONSTANCY OF CELL SHAPE
the number of its cells, and not to variations in cell size. The
extreme limits of the number of cells of which each type of leaf
may be made up, are obviously determined by heredity, and the
hereditary size of the organ is due to factors of periodicity in
growth, which determine the rate and duration of cell division.
Cell shape
Livingston (1901) describes the cells of Stigeoclonium as having
two characteristic shapes: a spherical form when growing in loose
masses on the trunks of trees, in highly concentrated media, or
when partially desiccated; and a cylindrical form, when growing in
filaments in extremely dilute media. Instances like the above
might lead one to believe that cell shape is largely a matter of
environmental conditions, such as mutual pressure, turgor as in-
fluenced by environment, etc. However, there are unicellular
algae, for example the desmids and diatoms, having a characteristic
cell shape which cannot be due to pressure nor any simple turgor
relations.
My studies show that the cells of the lower epidermis of the
leaves of any species of plant, in the regions between the veins,
have a characteristic length and breadth. The cell shape may, in
other regions, be modified to a considerable extent by various
factors, as by the presence of stomata, veins, etc. The relative .
length and breadth of the cells of any tissue may be the same in dif-
ferent species or varieties, as in Linum usitatissimum and Linum
angustifolium, or it may be somewhat different, as in Plantago
major and Plantago lanceolata.
Differences in the shapes of the leaves of the same plant, or of
related species, are not correlated with corresponding differences
in the shape of their cells. The linear leaves of Campanula
rotundifolia and Lobelia Erinus are not composed of longer, nar-
rower cells than those found in the round leaves, but have a
larger number of cells in the long axis of the leaf. The cells of
Cichorium Intybus are of the same size and shape in the lobed and
in the constricted portions of the leaf.
The shape of the leaf obviously cannot be the result of the dif-
ferences in cell shape but must rather be due to factors for period-
ically limiting the number and direction of the cell divisions in
TENOPYR: CONSTANCY OF CELL SHAPE 75
each type of leaf. The form of incised leaves like those of Cichor-
tum Intybus must be due to a factor or factors for differential
periodicity, which determines that the rate or duration or both, of
cell division, shall be greater in one part of the leaf than in another,
thus producing the lobes and constrictions.
In concluding, I wish to express my gratitude to Dr. R. A.
Harper, under whose direction these investigations were under-
taken, for the many helpful suggestions he gave me throughout
the course of this work. I also thank Dr. A. B. Stout, from whom
I obtained my chicory material, for his kind interest and help.
DEPARTMENT OF BOTANY,
OLUMBIA UNIVERSITY
LITERATURE CITED
Amelung, E. Ueber mittlere Zellengréssen. Flora 77: 176-207.
1893.
Conklin, E. G. Body size and cell size. Jour. Morph. 23: 159-188.
f. I-12. I
Familler, J. Die verschiedenen Blattformen von Campanula rotundi-
folia L. Flcra 87: 95-97. f. I oO.
Gates, R. R. The stature and eae of Oenothera gigas: De
Vries. Arch. Zellforsch. 3: 525-552. 1909.
Goebel, K. Einleitung in die experimentelle Morphologie der Latins
Leipzig und Berlin. 1908. ws
e, O. W., & Wawilow, N. Die anatomische Untersuchung
einiger Haferrassen. Jour. Opitnoi Agron. 13: 830-861. 1912.
[Russian.
Reviewed in Bot. Centralbl. 123: 481, 482. :
Jones, W. Neilson. Species hybrids of Digitalis. Jour. Genetics 2:
71-88. pl. 3-5 + f. I-45. 1912.
Keeble, F. Gigantism in Primula sinensis. Jour. Genetics 2: 163-188.
pl. rr +f.1-5. 1912.
Livingston, B. E. On the nature of the stimulus which causes the
change of form in polymorphic green algae. Bot. Gaz. 30: 289-317.
pl. 17, 18. 1900.
Further notes on the physiology of polymorphism in green
algae. Bot. Gaz. 32: 292-302. 1901.
_ Sachs, J. Physiologische Notizen. VI. Ueber einige Beziehungen der
specifischen Grésse der Pflanzen zu ihrer Organization. Flora 77:
49-81. 1893.
76 TENOPYR: CONSTANCY OF CELL SHAPE
Sanio, K. Ueber die Grésse der Holzzellen bei der gemeinen Kiefer
(Pinus silvestris). Jahrb. Wiss. Bot. 8: 401-420. 1872.
Sierp, H. Uber die Beziehungen zwischen Individuengrésse, Organ-
grosse und Zellengrésse, mit besonderer Berucksichtigung des
erblichen Zwergwuchses. Jahrb. Wiss. Bot. 53: 55-124. f. I-3.
1913.
Strasburger, E. Ueber die Wirkungssphaire der Kerne und die Zell-
groésse. Histol. Beitrage 5: 97-124. 1893.
Two remarkable Discomycetes
EpWarpD T. HARPER
(WITH PLATES I-3)
1. UNDERWOODIA COLUMNARIS Peck
I have collected this rare plant in several localities in Michigan
and Illinois. The first collection was on Neebish Island, Michigan,
in August, 1897. A cluster of small plants about two inches high
were found growing beside a path in balsam woods. The next
year some larger plants were found growing among dead leaves in
a ravine on Mackinac Island. In September three different col-
lections were made on Neebish Island, and the photographs on
PLATE 1 were taken. In May, 1908, the two plants shown on
PLATE 2 were found on a hillside in open woods at Bureau, Illinois.
A single plant was found at Neebish during the past summer.
The species was described and illustrated by Dr. Peck in the
43d Report of the New York State Museum, p. 78, p/. 4 (1890),
from plants found near Kirksville, New York, and sent to Dr-
Peck by Professor Underwood. A note in Underwood’s Moulds,
Mildews and Mushrooms, p. 65 (1899), says that six plants were
found in the same locality in three different years. Peck’s descrip-
tion is accurate as far as it goes, but his account of the base and
method of branching of the plant is incomplete. The plant is so
unique in structure and so remarkable in habit and size that I
have thought it worth while to discuss its structure and relation-
ships on the basis of my specimens.
Peck’s illustration shows the part above the ground only, and
he describes it as stemless and everywhere acigerous. This led
Schroeter to place the genus among the Rhizinaceae. The plant
does, however, have a short stem and is more or less bulbous at
the base. The lower margin of the hymenium is uneven, running
down in points on the stem while naked strips extend upward for
short distances into the hymenium as in some species of Geo-
glossum. There is a definite though inconspicuous margin to the
*
78 HARPER: TWO REMARKABLE DISCOMYCETES
hymenium and in dried plants the hymenium becomes reddish
brown where the asci are numerous, while the stem remains
whitish. The base of the plant is rounded like the base of species
of Morchella. No thick mycelial root or sclerotium of any kind
was found.
Sometimes the clubs are simple as shown on PLATE 2 and’ in
Peck’s illustration but they usually divide into fingers. The
plant on PLATE 1 was divided near the middle. The halves
have split and sprung apart as shown in B. The left branch
shown at A is forked near the apex.’ The right branch was nearly
destroyed by insects but appeared to have been more deeply
divided. Other plants in the collections were divided a little
below the middle. PLATE 2 shows two plants, one of which has
been sectioned, closely caespitose and apparently connected at
the base. In the cluster of small plants first discovered there were
at least three connected at the base.
The whole interior of the plant—bulbous base, stem and as-.
coma—is perforated by irregular longitudinal cavities, as shown in
the section on PLATE 2,A. The walls of the cavities are of nearly
uniform thickness. The interior of the walls is composed of a
mesh of septate hyphae 6—-8u in diameter. The cavities are lined
with a palisade layer of regular, septate hyphae about 6y in diam-
eter. The layer is about 50u thick and there are three or four
septa in each hypha. The appearance of a cross section of one
of the walls is like that shown in Hesse’s Hypogaeen Deutsch-
lands, pl. 18, f. 12 (1894), except that the interior is composed
entirely of septate hyphae. |
The outside walls are covered with the hymenium on the outside
and the palisade layer on the inside. The surface of the hymenium
is wrinkled and corrugated, PLATE 2, C, but there appears to be no
definite relation in position between the wrinkles of the hymenium
and the internal cavities. The interior of the walls is composed
of hyphae, and there appears to be no definite subhymenial layer.
A palisade layer like that on the walls of the interior cavities
covers the stem and base of the plant.
easci, PLATE 2, B, are club-shaped, 10-12 X 160-200n. They
open by a lid as far as I can make out but I have not been able to
find empty asci, which Boudier says must be examined to deter-
HARPER: TWO REMARKABLE DISCOMYCETES 79
mine the nature of the opening. The character of the plant and
the spores is that of Boudier’s division Opercules. The paraphyses
are septate, a little longer than the asci, slightly thickening up-
ward and somewhat curved at the apex, 4—6y in diameter.
The colors of the plant are whitish or cream colored outside
and pure white within.
I have rewritten Peck’s description to include the form of the
base and method of branching, as follows:
Ascocarps single or caespitose, simple or more or less deeply
and somewhat dichotomously divided, divisions columnar, straight
or slightly curved, sulcate-costate and uneven; ascigerous layer
definite but not separated from the short stem. Base even or
slightly bulbous, interior perforated by irregular longitudinal
cavities, separated by thin walls, whitish or brownish outside,
pure white within, cavities lined with a regular palisade layer of
septate hyphae; asci club-shaped, IO-I2 X seGaawed ee
slightly longer than the asci, rarely branched, septate, 4—6y in
diameter; spores tuberculate, hyaline or slightly aca, elliptical,
10-13 X 18-24.
Varying greatly in size, 5-35 cm. high, 2-5 cm. thick.
Growing on the ground among grass or dead leaves in mixed
woods.
Underwoodia columnaris is not closely related to any known
discomycete. It forms a monotypic genus, as the term is usually
understood. It appears to be the only representative of its group.
The group arrangement is especially applicable to the large
discomycetes, for the groups are few and well marked; and, while
the forms within them are closely related, the kinship of the groups
themselves appears remote. Apart from Underwoodia there are
only seven groups in the family Helvellaceae in our region: the
Morchella esculenta, Morchella hybrida and Verpa digitaliformis
groups in the Morchella-Verpa series and the Gyromitra esculenta,
Gyromitra infula, Helvella crispa and Helvella elastica groups in the
Gyromitra-Helvella series. In the Geoglossaceae there may be
about thirteen groups, nine of which are in the large Geoglossum
series and one each in Spathularia, Cudonia, Leotia and Vibrissea.
The groups are also few and well defined among the larger Peziza-
ceae. The order Helvellales is based on little more natural re-
lationship than that its members are the large conspicuously
stemmed discomycetes.
80 HARPER: TWO REMARKABLE DISCOMYCETES
Forms within the groups offer a most fruitful field for ecological
and comparative study. Most of the groups of large discomycetes,
such as the Verpa, Cudonia, Leotia, and Geoglossum glutinosum
groups, appear in a variety of forms. In some groups the forms
are very abundant and variable. Boudier has named and given
beautiful illustrations of twenty-three forms in the Morchella
esculenta group, offering a fine opportunity for students to com-
pare other Morchella esculenta floras with that of France. Durand
lists seven forms in the Geoglossum glabrum group. Underwoodia
is so rare that its variation is little known, though it may have
had a bloom period at some time in the past.
But while the kinship and variation of Underwoodia are un-
known it possesses a striking combination of characters found
singly in other groups. Most of these characters are more highly
developed than in other groups where they occur and this gives to
Underwoodia a unique position and a special interest. I can only
briefly compare the most striking characters and do it without
implying any close kinship between Underwoodia and the groups
in which similar characters are found.
The cancellated stem of Underwoodia is more highly developed
than in any other discomycete. It contrasts strongly with the
hollow stems of species of Morchella and the solid stems of the
Geoglossaceae. The character is found in the Acetabula vulgaris
and the Helvella crispa groups, but the stems in these groups are
deeply sulcate as well as cancellate, the cavities are much less
regular and the walls of the cavities as far as I have examined
them are not covered with a regular palisade layer. The stems
of species of Morchella often have a few cavities near the base.
They are very markedly developed in the specimen of Gyromitra
gigas illustrated by Boudier (Icon. Mycol. pl. 221. 1904). Such
cavities are not lined with a palisade layer of hyphae.
Many discomycetes have lacunae and cavities in the stem. An
interesting case for comparison with Underwoodia was identified for
me by Durand as Lachnea (or Macropodia) semitosta. The stem
bears a close resemblance to that of Underwoodia. There area few
sulcations on the outside, but the walls of the cavities are not
lined with such a regular palisade layer. Macropodia semitosta is
remarkable in the development of the cancellated stem. Note the
e
HARPER: TWO REMARKABLE DISCOMYCETES 81
illustration of Peziza semitosta in Cooke’s Mycographia, f. rog
(1879). The palisade layer lining the cavities in the body of the
fungus is most highly developed in species of Hypogaei, but in
them it is connected with the hymenium. Bucholtz derives the
Helvellaceae from Lachnea through the Hypogaei.
The cancellated stem is combined in Underwoodia with a club-
shaped ascoma, not cup-shaped or saddle-shaped as in the Acetabula
and Helvella groups. Such club-shaped ascomata are found in the
groups of the Geoglossum series. In size, however, in substance,
in the spores, and probably in the opening of the asci Underwoodia
is widely removed from any of the groups in that series. The
hymenium is also more sulcate than in species of Geoglossum and
resembles in this respect the hymenium of Verpa. Compare
Boudier’s illustrations on pi. 219 and 220. We may also compare
the ascoma of Gyromitra gigas referred to above. Gyromitra
gigas, as illustrated by Boudier in pl. 221, is nearer in general shape
and appearance to Underwoodia than any other discomycete with
which I am familiar.
The spores of Underwoodia have a tuberculate wall and in this
respect are more highly developed than is usual among the Hel-
vellaceae, the other species of which have smooth, hyaline spores.
The question whether Underwoodia is a gymnocarp or an
angiocarp cannot be settled without the young stages. In the
ascoma of the species of Helvella the young hymenium is over-
grown by surrounding tissue as in very many cup fungi and the
same is true of the elements of the compound hymenium of Mor-
chella. The hymenium of Underwoodia does not appear to have
been so overgrown at any time, but whether it was covered at any
stage with a weft of fibers, as is the case with many of the Geo-
glossaceae is not known. Nor is it known whether the hymenium
originated on or below the surface of the primitive ascocarp.
Questions as to the protection of the young hymenium in the dis-
comycetes are not yet settled.
The high development of the different characters in Under-
woodia gives it a most important position. It is one of the largest
and certainly the most highly developed of all the discomycetes.
82 HARPER: Two REMARKABLE DISCOMYCETES
2. PUSTULARIA GIGANTEA Rehm
In July, 1899, I found some large cup fungi on the ground
among dead leaves on Mackinac Island, Michigan. Later I sent
them to Dr. H. Rehm, who described them as a new species under
the name Pustularia gigantea. The description was published in
Annales Mycologici (3: 517. 1915). I have since found the plant
several times in coniferous woods on Neebish Island. In Septem-
ber, 1907, it was abundant in a piece of cedar and balsam woods,
and I secured the photograph of the opened plant, PLATE 3, A,
and preserved a number of unopened plants in alcohol, from
which the photograph of the section, B, was made. The specimens
sent to Dr. Rehm were dried and somewhat torn, so that they did
not show the peculiar folding in of the edges of the apothecium and
the way in which the mature plants burst open, which are two
of the most striking characters of the species.
The unopened apothecia are irregularly ellipsoid with a deep
groove across the top. The apothecia often occur in clusters in
which the orientation of the members is irregular. The surface of
the apothecia is nearly smooth, whitish in color, and much soiled
by the earth or mould in which the plants are buried.
The apothecia have a definite interior structure which is shown
in PLATE 3,B. This figure represents a vertical section cut from
the center of an unopened apothecium. There is no stem and the
flat under surface lies directly on the soil, with which it is connected
by strands of white mycelium. Quite frequently the connection
with the substratum is lost, as is the case with the Hypogaei.
The cup is roughly and partially divided into two chambers by
the infolding of the edges of the apothecium. The chambers can
be recognized in all the plants examined, though they are at times
more or less distorted. The infolded edges extend to the bottom
of thecup. In a section like this made through the center of the
apothecium the infolded walls are divided at the base. In sections
near the ends of the apothecium they are united. The walls have
been infolded till the usual apical opening or mouth of the cup in
other Pezizaceae lies on the bottom of the cup. Neither the lips
nor the walls are grown together, though they are pressed closely
against each other. In the section before us the walls have sprung
apart. The lips are like those of other cup fungi with the hypo-
HARPER: TWO REMARKABLE DISCOMYCETES 83
thecium extending a little beyond and embracing the hymenium.
The bottom of the cup is thickened and raised into a sort of
cushion covered like the rest of the interior by the hymenium.
Such a cushion is evident on the bottom of all the cups examined.
Around the cushion is a narrow zone which is destitute of hyme-
nium. It has apparently resulted from a rupture of the hymenium
during growth. In the larger chamber is an irregular outgrowth
covered by the hymenium such as may occur in almost any position
on the hymenium of these plants.
An end section of another specimen shows an apothecium in
which the basal cushion is very large. The chambers are irregular
and there are several minor cavities. The general structure is
however the same as in the plant figured on PLATE 3. The basal
cushion in this second specimen is hollow and it was deeply in-
dented on the under side in a zone around the central portion,
which was attached to the soil as a sort of stem.
When the apothecia are mature and the growth tension becomes
great; instead of opening at the mouth as in most species of Peziza,
the walls of the chambers burst irregularly as shown in PLATE 3, A.
The appearance of such opened plants is striking. The walls of
the internal cavities are pure bluish white and the thick reflexed
segments give the plant the appearance of a large white flower.
When the weather is unfavorable to rapid growth the apothecia
do. not open as far as my observation goes but dry up or decay
without exposing the hymenium. Sometimes irregular openings
are found in the walls of old plants.
In regular plants the walls of the apothecia are of nearly equal
thickness throughout. They are covered on the inside by the
hymenium. The hypothecium is made up of irregular septate
hyphae of different diameters with many large irregular cells and
cavities, especially in the center. The hyphae are much more
dense directly beneath the hymenium where they form an indefi-
nite subhymenial layer. On the outside there is a compact corti-
cal layer of hyphae which run somewhat parallel to the surface.
In the center of the walls the tissue is looser and the cells larger.
Often the tissue breaks down and leaves hollow spaces in the walls.
as can be seen in the sections.
The asci are long and narrow and closely packed together with
84 HARPER: TWO REMARKABLE DISCOMYCETES
relatively few paraphyses. They measure 10-12 X 200~-300un.
They are linear for most of their length, rounded-truncate at the
apex and narrowed near the base. The base is enlarged and irreg-
ular. The paraphyses are about 2u in diameter, slightly enlarged
at the upper end, rarely branched and with few septa. The spores
are oblong with rounded ends, smooth, usually with two oil drops
when mature, 5-8 X I0-I4up.
Rehm placed the plant in the genus Pustularia and compared
it with Pustularia vesiculosa, noting that it differed from that
species in the larger size and smaller spores. According to Rehm’s
key Pustularia contains species of fleshy Pezizaceae, which have
smooth and entire cups, asci turning blue with iodine, elliptical
spores and sessile apothecia. It includes species like Peziza
vesiculosa, P. Stevensoniana, and P. coronaria. Boudier disregards
the surface of the apothecia and places species like Peziza coronaria
and Sepultaria sepulta in a genus Sarcosphaera, on the ground that
they are semi-subterranean with the apothecia closed at first and
bursting irregularly at maturity. Pustularia gigantea is semi-sub-
terranean in its habit and bursts irregularly at maturity and might
be placed in this genus, but it has a definite mouth the walls of .
which have simply become infolded during growth.
' The natural arrangement of the species of Pezizaceae has not
yet been discovered. All that was said about the groups of the
Helvellaceae above applies equally to the groups of the large cup
fungi. As Dodge has emphasized, a knowledge of the early stages
is a necessary prerequisite of a natural classification.
It is not certain to which group Pustularia gigantea belongs.
It is not close to Pustularia vesiculosa, as Rehm suggests, for in
addition to the difference in the size of the spores the subterranean
habit and character of the mouth are very different from that
species. Our collections of Pustularia-vesiculosa have mouths of
the ordinary form and grow in the mulching about trees and on
manure heaps. They agree with Lloyd’s photograph in Hard’s
Mushroom, f. 432 (1908). Pustularia gigantea appears to be closer
to Peziza coronaria, for that species is subterranean. Boudier’s illus-
tration, pl. 302, resembles our expanded plant. Cooke's illustra-
tion in Mycographia, f. 238, agrees in size and method of opening,
and the same is true of Kalchbrenner’s illustration of Peziza coro-
HARPER: TWO REMARKABLE DISCOMYCETES 85
naria var. macrocalyx (Icon. Select. pl. 40, f. 2). The spores also are
nearer in size to those of Peziza coronaria than to those of P. vesicu-
losa. Saccardo’s Sylloge (8: 81. 1889) gives the spores of P.
coronaria 8-9 X 15-18u. The spores of Pustularia gigantea are
often 8 X 14, though Rehm gives 5-6 X 10-124. The descrip-
tions of Peziza coronaria do not, however, give the internal struc-
ture of the unopened apothecia. It is supposed that the young
plants have no evident mouths. The species of Sepultaria also
are said to be closed at first and to open irregularly. Yet what
is known of the structure of the cup fungi suggests that in all
these forms the hymenium has been overgrown by surrounding
tissue, and the location of the mouth should be determinable
even in young unopened apothecia. Pustularia gigantea is very
enlightening on this point and the unopened apothecia of species
like Peziza coronaria should be carefully examined to see if there is
not evidence of the location of the mouth. Itis uncertain whether
Pustularia gigantea should be associated with Peziza coronaria or
placed in a group by itself. Further observation may bring to
light other more closely related forms.
The most striking feature of Pustularia gigantea is the degree
in which the walls of the apothecia are infolded. It is very com-
mon among the discomycetes for the hymenium to be covered by
the surrounding tissue during the formative period. In small
species of Peziza the ascoma usually remains circular, but in large
plants the walls of the cup are flattened and folded together from
the sides. This is also true of species of Helvella which have a
cup-shaped or saddle-shaped ascoma. The saddle-shaped ascoma
in the Helvella elastica group is folded and flattened in the same
way when young and later opens at the ends instead of in the
middle giving it the characteristic saddle-shaped form.
Bucholtz has shown that in many of the Tuberaceae the hy-
menium is formed on the outside of the ascocarp and then over-
grown by surrounding tissue until it is entirely enclosed in cavities
of the fruit body. Pustularia gigantea has many characters in com-
mon with the Tuberaceae. The closed chambers, subterranean
habit and loose connection with the soil are all suggestive of the
Tuberaceae. We need only mention the Hysteriaceae to show
how common the inrolling of the disk and consequent covering of
the hymenium is among the discomycetes and their allies. The
86 HARPER: TWO REMARKABLE DISCOMYCETES
striking feature of Pustularia gigantea is the extent to which the
walls are infolded The mouth is completely inverted and lies on
_ the bottom of the cup. In this the plant appears to be unique.
Explanation of plates 1-3
PLATE 1. UNDERWOODIA COLUMNARIS Peck
A, upper half of the left branch of a large plant; B, lower half of the plant.
PLATE 2. UNDERWOODIA COLUMNARIS Peck
A, longitudinal and cross sections; B, ascus and paraphyses; C, one of a pair
of plants grown together at the base.
PLATE 3. PUSTULARIA GIGANTEA Rehm
A, expanded plant. 3B, vertical section through an unopened apothecium; C,
asci and paraphyses.
*
INDEX TO AMERICAN BOTANICAL LITERATURE
1916-1917
The aim of this Index is to include all current botanical literature written by
Americans, published in America, or based upon American material ; the word Amer-
ica being used in the broadest sense.
eviews, and papers that relate exclusively to forestry, ee onirag horticulture,
manufactured products of vegetable origin, or laboratory methods are not included, and
no attempt is made to index the literature of bacteriology. An cote exception is
made in favor of some paper appearing in an American periodical which is devoted
wholly to botany. Reprints are not mentioned unless they differ from the original in
some important particular. If users of the Index will call the attention of the editor
‘o errors or omissions, their kindness will be appreciated.
This Index is reprinted monthly on cards, and farnishea | in this form to subscribers
at the rate of one cent for each card, Selections of cards are not permitted ; each
subscriber must take all cards published during the term of his subscription, e-
spondence relating to the card issue should be addressed to the Treasurer of the Torrey
Botanical Club,
Abrams, L. Flora of Los Angeles and vicinity. i-x + 1-432. Stan-
ford University. 10 Ap 1917.
Allard, H. A. Further studies of the mosaic disease of tobacco. Jour.
Agr. Research 10: 615-632. pl. 63. 17 5 1917.
Allen, C. E. A chromosome difference correlated with sex differences
in Sphaerocarpos. Science II. 46: 466, 467. 9 N 1917.
Armstrong, M. Common plant names. Science II. 46: 362. 12 0
1917.
Arthur, J.C. Cultures of Uredineae in 1916 and 1917. Mycologia 9:
2904-312. 24S 1917.
Barnhart, J. H. The first hundred years of the New York Academy
of Sciences. Sci. Mo. 5: 463-475. N 1917. [lllust.]
Beckwith, F., Macauley, M. E., & Baxter, M. S. Plants of Monroe
County, New York, and adjacent territory. Proc. Rochester Acad.
Sci. 5: 59-99. My 1917. :
Berry, E. W. Pleistocene plants in the marine clays of Maine.
Torreya 17: 160-163. f. I-3. 2 0 1917.
Boughton, F.S. Hymenomyceteae of Rochester, N. Y., and vicinity
Proc. Rochester Acad. Sci. 5: 100-119. My 1917.
Boynton, K. R. Aster tataricus. Addisonia 2: 51. pl. 66. 295 1917.
ree
88 INDEX TO AMERICAN BOTANICAL LITERATURE
Brenckle, J. F. North Dakota Fungi—I. Mycologia 9: 275-293.
24.S- 1917.
Britton, N. L. Harrisia gracilis. Addisonia 2: 41, 42. pl. 61. 29S
IQI7.
Britton, N. L. Harrisia Martini. Addisonia 2: 55, 56. $1, 08. 205
1917.
Brooks, S.C. A new method of studying permeability. Bot. Gaz. 64:
S0eat7. f,-2,:22 160 10t7.
Brown, P. E., & Hitchcock, E. B. The effects of alkali salts on nitri-
fication. Soil Sci. 4: 207-229. f. 1-14. S 1917.
_ Brown, W. H., & Heise, G. W. The relation between light intensity
and carbon dioxide assimilation. Philip. Jour. Sci. 12: [Bot.] 85-97.
t,o. Mir T6177.
Bunyard, E. A. The history and development of the strawberry.
Jour. Internat. Gard. Club 1: 69-90. Au 1917. [Illust.]
Burnham, S. H., & Latham, R.A. Corrections of the flora of the town
of Southold. Torreya 17: 164. 20 1917.
Burrill, T. J., & Hansen, R. Is symbiosis possible between legume
bacteria and non-legume plants? Illinois Agr. Exp. Sta. Bull. 202:
TiS“*61.. OF 2-17. Jl 1917:
Burt, E.A. Odontia Sacchari and O. saccharicola, new species on sugar-
cane. Ann. Missouri Bot. Gard. 4: 233. f. r, 2. 20S 1917.
Burt, E. A. The Thelephoraceae of North America—VIII. Conto-
phora. Ann. Missouri Bot. Gard. 4: 237-269. 20S 1917.
Includes Coniophora inflata, C. vaga, C. alvellanea, C. Harperi, and C. flava,
Spp. nov.
Butters, F. K. Taxonomic and geographic studies in North American
ferns. Rhodora 19: 169-216. pl. 123 +f. 1-6. 11 O 1917.
Also published in Contr. Gray Herb. 51: 170-216. 11 O 1917.
Cary, M. Life zone investigations in Wyoming. North Am. Fauna
42: 7-95. pl 15 +f. 1-17... 3.01917.
Cockerell, T. D. A. Somatic mutations in sunflowers. Jour. Heredity
8: 467-470. f. 10-12. O 1917.
Coons, G. H., & Levin, E. The leaf-spot disease of tomato. Michigan
Agr. Exp. Sta. Spec. Bull. 81: 1-15. f. 1-7. Je 1917.
Cooper, W.S. Redwoods, rainfall and fog. Plant World 20: 179-189.
TY; a Je tor.
i, C. S. Seed production in apples. Illinois Agr. Exp. Sta.
Bull. 203: 185-213. 7. 1-8. Au 1917.
INDEX TO AMERICAN BOTANICAL LITERATURE 89
Dallimore, W. D. New species of Rosa. Jour. Internat. Gard. Club
I: 213-218. Au 1917. [Illust.]
Downing, M. B. Edward Lee Greene. Catholic World 106: 13-24.
O 1917.
Duggar, B. M., Severy, J. W., & Schmitz, H. Studies in the physiology
of the fungi—V. The growth of certain fungi in plant decoctions.
Ann. Missouri Bot. Gard. 4: 279-288. f. 1-5. 20S 1917.
Elliott, J. A. Taxonomic characters of the genera Alternaria and
Macrosporium. Am. Jour. Bot. 4: 439-476. pl. 19, 20 +f. 1-6.
20.1997:
Erwin, A.T. Bordeaux spray for tip burn and early blight of potatoes.
Towa State College Agr. Exp. Sta. Bull. 171: 63-75. f. 1,2. Jl 1917.
Early blight and tip burn are illustrated in color.
Fairchild, D. The annual catalogue of plant emigrants. Jour. Hered-
ity 8: 500-508. f. 6-13. N 1917.
Faulwetter, R.C. Wind-blown rain, a factor in disease dissemination.
Jour. Agr. Research 10: 639-648. f. 1. 17S I9I7.
Fernald, M.L. The tardy flowering of plants in eastern Massachusetts
in the spring of 1917. Rhodora 19: 219, 220. 11 O 1917.
Floyd, B. F. Dieback, or exanthema of citrus trees. Florida Agr. Exp.
Sta. Bull. 140: 1-31. f. 1-15. Au 1917.
Fowler, L. W., & Lipman, C. B. Optimum moisture conditions for
young lemon trees on a loam soil. Univ. Calif. Publ. Agr. 3: 9-11.
pl. 25-36. 29 S 1917.
Fraser, W. P. Overwintering of the apple scab fungus. Science II.
46: 280-282. 21S 1917.
Freeman, G. F. Linked quantitative characters in wheat crosses.
Am. Nat. 51: 683-689. N 1917.
Fulmer, H. L. The relation of green manures to nitrogen fixation.
Soil. Sci. 4: 1-17. f. 1-4. Jl 1917.
Gainey, P. L., & Metzler, L. F. Some factors affecting nitrate-nitrogen
accumulation in soil. Jour. Agr. Research 11: 43-64. 8 O 1917.
Gano, L., & McNeill, J. Evaporation records from the Gulf coast.
Bot. Gaz. 64: 318-329. f. 1-4. 16 O 1917.
Garner, W. W., Wolf, F. A., & Moss, E.G. The control of tobacco wilt
in the flue-cured district. U.S. Dept. Agr. Bull. 562: 1-20. f. 1-5.
15.5 1017.
Gates, F. C. The revegetation of Taal volcano, Philippine Islands.
Plant World 20: 195-207. f. 1-4. Jl 1917.
90 INDEX TO AMERICAN BOTANICAL LITERATURE
Gates,R.R. The mutation theory and the species concept. Am. Nat:
51: 577-595: O 1917.
Gleason, H. A. Some effects of excessive heat in northern Michigan.
Torreya 17: 176-178. 31 0.1917.
Gleason, H. A. The structure and development of the plant associa-
tion. Bull. Torrey Club 44: 463-481. 1 O 1917.
Grantham, A. E. The tillering of wheat. Sicence II. 46. 392, 393.
19 O 1917.
Gravatt, G. F., & Marshall, R. P. Arthropods and gasteropods as
carriers of Cronartium ribicola in greenhouses. Phytopathology 7:
368-373. 30 1917.
Gregory, W. K. Genetics versus paleontology. Am. Nat. 51: 624-
635s. O 19t7.
Grossenbacher, J. G. Crown-rot of fruit trees: histological studies.
m. Jour. Bot..4: 477-512. pl. 21-27. O 1917.
[Grosvenor, G. H.] Our state flowers. The floral emblems chosen by
the commonwealth. Nat. Geog. Mag. 31: 481-517. Je 191
917.
Each state flower is illustrated in color from the original paintings by Miss
M. E. Eaton, of the N. Y. Bot. Garden.
Guilliermond, A. Levaduras del pulque. Bol. Direc. Estud. Biol.
Mexico 2: 22-28. Ja 1917.
Illustrated with numerous figures.
Giissow, H. T. Plant diseases in Canada. Science II. 46: 362. 12
O I917.
A report on the presence of Dothichiza Populea, Colletotrichum cereale, and
Sporidesmium exitiosum.
Harper, R.M. A preliminary soi! census of Alabama and west Florida.
Soil Sci. 4: 91-107. Au 1917.
Harper, R. M. The native plant population of northern Queens
County, Long Island. Torreya 17: 131-142. 14 Au 1917.
Harris, J. A. The weight of seeds as related to their number and posi-
tion. Torreya 17: 180-182. 31 O 1917.
Harris, J. A., & Lawrence, J. V. Cryoscopic Wererinionticies on tissue
fluids of plants of Jamaica coastal deserts. Bot. Gaz. 64: 285-305.
16 O 1917.
Harshberger, J. W. Pennsylvania men commemorated in the names
of plants. Alumni register 1917: pp. 3. Ap 1917. [Illust.]
A reprint without pagination.
Harshberger, J. W. A text book of mycology and plant pathology.
i-xili + 1-779. f. 1-270. Philadelphia. 1917.
INDEX TO AMERICAN BOTANICAL LITERATURE 9]
Haskell, R. J. The spray method of applying concentrated formalde-
hyde solution in the control of oat smut. Phytopathology 7: 381-
484. 3 © 19%9.
Herre, A. C. Preliminary notes on the lichens of Whatcom County,
Washington. Bryologist 20: 76-84. f. 1. S 1917.
Herrera, A. L. Estudios de plasmogenia. Bol. Direc. Estud. Biol.
Mexico 2: 29-45. 1917. [Illust.]
Hines, C. W. A study of the root system of the sugar cane and its
application to the production of ratoon crops. Philip. Agr. Rev. 10:
ERT“ 161.. f..2-6.<: 4917.
{Hitchcock, A. S.] Botanical explorations in the Hawaiian Islands.
Smithsonian Misc. Coll. 66": 59-73. f. 61-77. 1917.
Hofmann, J. V. Natural reproduction from seed stored in the forest
floor. Jour. Agr. Research 11: 1-26. pl. 1-7 + f. 1-4. 1 O 1917.
Hooker, ‘H. D., Jr. Mechanics of movement in Drosera rotundifolia.
Bull. Torrey Club 44: 389-403. 10 Au 1917.
Hotson, J. W. Notes on bulbiferous fungi with a key to described
species. Bot. Gaz. 64: 265-284. pl. 21-23 +f. 1-6. 16 O 1917.
Papulospora pallidula, P. byssina, P. aurantiaca, P. nigra, and P. magnifica,
_ 8pp-. nov., are described. :
House, H. D. The Peck testimonial exhibit of mushroom models.
Mycologia 9: 313, 314. 24S 1917.
Also published in Torreya 17: 178-180. O 1917.
Jackson, H.S. Two new forest tree rusts from the northwest. Phyto-
pathology 7: 352-355. 3 O 1917.
Chrysomyxa Weirii and Melampsora occidentalis.
Jehle, R. A. Susceptibility of non-citrus plants to Bacterium Citri.
Phytopathology 7: 339-344. f. I-3. 3 O 1917.
Jones, D. F. Linkage in Lycopersicum. Am. Nat. 51: 607-621. O
1917.
Jones, L. R. Soil temperature as a factor in Phytopathology. Plant
World 20: 229-237. Au 1917.
Kraebel, C. J. Choosing the best tree seeds. Jour. Heredity 8: 483-
492. f. 1-5 + frontispiece. N 1917.
The influence of paternal character and environment upon progeny of Douglas
fir.
Levin, E. Control of lettuce rot. Phytopathology 7: 392, 393-
1917.
Lioyd, C. G. The Geoglossaceae (viz., the genus Geoglossum and
related genera). 1-24. f. 782-807. My 1916.
=O
92 INDEX TO AMERICAN BOTANICAL LITERATURE
Lloyd, C. G. Mycological notes 48: 670-684. f. 992-1025. Jl 1917:
49: 685-700. f. 1026-1048. Jl 1917.
Long, W. H. Notes on new or rare species of Gasteromycetes. Myco-
aint 271-274... 24 S 19017:
Includes Geasteroides and eens gen. nov. and Geasteroides texensts tats
Leodilne albicans, spp. n
MacDougal, D. T. The Pe and physical basis of parasitism.
Plant World 20: 238-244. f. 7. Au 1917.
McCubbin, W. A., & Posey, G. G. Development of blister rust aecia
on white pines after they had been cut down. Phytopathology 7.
301, 392: 3 O 19017.
McDonnell, C. C., & Roark, R.C. Occurrence of manganese in insect
flowers and insect flower stems. Jour. Agr. Research 11: 77-82
15 O 1917.
McNair, J. B. Fats from Rhus laurina and Rhus diversiloba. Bot.
Gaz. 64: 330-336. f. 7. 16 O 1917...
Miles, L. E. Some diseases of economic plants in Porto Rico. Phyto-
pathology 7: 345-351. f. 1-3. 30 1917.
Moore, G. T. Algological notes. I. Chlorochytrium gloeophilum
Bohlin. Ann. Missouri Bot. Gard..4: 271-278. pl. 18. 20S 1917.
Murrill, W. A. Collecting fungi at the Delaware Water Gap. Jour.
N. Y. Bot. Gard. 18: 207. S$ 1917.
Murrill, W. A. A disease of the hemlock tree. Jour. N. Y. Bot. Gard.
18: 208. S$ 1917.
- Murrill, W. A. An excursion to Delaware Water Gap. Torreya 17:
148-150. 14 Au 1917.
Murrill, W. A.
Mlustrations of fungi—X XVII. Mycologia 9: 257-
260.
nopus strictipes, Cortinellus rutilans, Gymnopus dryophilus, Prunulus purus,
Clitocybe virens, Chaniarel Chantarellus are illustrated in color.
Nash,G.V. Bomarea edulis. Addisonia 2: 49, 50. pl. 65. 29S 1917.
Nash, G. V. Epidendrum oblongatum. Addisonia 2: us pl. 62. 29
S 1917.
Nash, G. V. Oncidium pubes.
Addisonia 2: 57. pl. 69. 29 S 1917.
Nash, G. V. Raphiolepis ovata.
Addisonia 2: 59. pl. 70. 29S 1917.
Nelson, J. C. The introduction of foreign weeds in ballast as illus-
trated by ballast-plants at Linnton, Oregon.
Torreya 17: 151-160.
2 O-1927.
CLUI
ORREY
UNDERWOODIA OLUMNARIS
VOLt ME
TORREY CLUB
BULL.
UNDERWOODIA COLUMNARIS PEcK
PLATE 3
Sy
T
VOLUME
TORREY CLUB
BULL.
CSRS BOS Ges « = =
a BN > © 9 :
rer RST TET ae
St na
semanas Bei, inn eS
aaa cane
CoO OS o8O
GIGANTEA REHM
A
ARI
PUSTUL
Vol. 45 No. 3
BULLETIN
OF THE
TORREY BOTANICAL CLUB
MARCH, 1918
Studies of some new cases of apogamy in ferns
W. N. STEIL
(WITH PLATES 4 AND 5)
INTRODUCTION
During the past six years, the writer has made an attempt to
determine to what extent undernormal cultural conditions apogamy
occurs in the homosporous leptosporangiate ferns and especially
in the genera Pellaea, Pteris, and Aspidium. Since the nuclear
history in only a few apogamous ferns has been investigated, it was
believed that further studies in the cytology of such ferns would be
desirable. Hence an investigation of this nature was undertaken
in the species in which apogamy was discovered, and this part of
the work has been considered more interesting and important than
the discovery of new cases of apogamy. In some of the ferns
studied the nuclear history is wholly or partly known, but the dis-
cussion of this subject is reserved for another paper. At this time
new cases of apogamy will be reported and briefly considered.
METHODS AND MATERIALS
On account of the great difficulty experienced in securing spores
for cultural work, the investigation, so far as the discovery of new
cases of apogamy is concerned, cannot be regarded as wholly suc-
cessful. Some of the spores were collected in the field. A large
number of plants were grown in the university greenhouse, and
[The BULLETIN for February (45: 51-92. pl. 1-3) was issued March 7, 1918]
93
94 STEIL: SOME NEW CASES OF APOGAMY IN FERNS
spores were collected from these. Spores from a still larger
number of species were obtained through the kindness of the fol-
lowing persons: Dr. A. B. Stout, New York Botanical Garden;
Dr. R. C. Benedict, Brooklyn Botanic Garden; Dr. G. T. Moore,
Missouri Botanical Garden; Dr. E. B. Copeland, Los Banos,
Philippine Islands; Mr. F. C. Greene, Rollo, Missouri; and Rev.
George Moxley, Los Angeles, California. To these gentlemen the
writer wishes to express his sincerest thanks, for in no small
measure they contributed to the investigation.
The spores were generally sown on the surface of a nutrient
solution or on sphagnum. The latter was placed in a Stender
dish and saturated with a nutrient solution. Before sowing the
spores, the medium was thoroughly sterilized. Other media, such
as nutrient agar, peat clay and loam, were also used, but none
of these proved as satisfactory for the cultural work as the sphag-
num. The Stender dishes were placed under bell-jars in a Wardian
case in the university greenhouse. The jars were tilted on edge,
so that the prothallia were provided with a sufficient supply of
oxygen and carbon dioxid. The temperature of the Wardian case
varied from 65° F. in winter to about 110° F. in summer. The
prothallia were protected from too intense illumination by shading.
The light was very favorable for the growth of fern prothallia
since, when they were not crowded in the cultures, they became
heart-shaped. The moisture supply was always sufficient for
fertilization to occur in the non-apogamous species grown at the
same time under precisely the same cultural conditions.
In a large number of species in which fertilization is known to
occur, including Pteris aquilina L., P. serrulata L., Osmunda re-
galis L., O. Claytoniana L., O. cinnamomea L., and Adiantum
pedatum L., sex organs were produced under the conditions just
described, and embryos were formed only as a result of the union
of the gametes.
The prothallia of Nephrodium molle Desv. and Asplenium
nidus L., in which Yamanouchi (1908) and Nagai (1914) respec-
tively induced apogamy, were grown under the same conditions,
_ but while embryos were produced in large numbers, none were
formed apogamously.
The spores of some of the apogamous species were sown on soil
STEIL: SOME NEW CASES OF APOGAMY IN FERNS 95
in the university greenhouse, but the prothallia never showed any
peculiarities in their development, and embryos were never pro-
duced by fertilization. Hence it appears that cultural conditions
were not a factor in inducing apogamy in any case.
In the majority of cases, the prothallia were grown until em-
bryos were formed, either as a result of fertilization or apoga-
mously. Parthenogenesis was not excluded in the non-apoga-
mous species. To determine this point a cytological investigation
would have been necessary.
The conclusions are in no case based on a single culture but on
a large number of cultures. Cultures in which only a few pro-
‘thallia were obtained were always discarded. Great care was
exercised in handling the spores before sowing in order to avoid,
so far as possible, mixing those of different species.
Two sets of cultures were made of each of the ferns tested for
apogamy. In one set only a small number of spores were sown
to avoid crowding the prothallia. The prothallia under these con-.
ditions grew to a good size and became heart-shaped. It was be-
lieved that these cultures were favorable for the development of
archegonia. The other cultures were made by sowing a large
number of spores. The majority of the prothallia in these cul-
tures were irregular in form and usually produced numerous an-
theridia. If embryos were produced, they appeared in both types
of cultures. From the two sets of cultures it was possible to
determine whether sex organs developed.
While the prothallia of the different species were growing, they
were carefully examined from time to time with a microscope.
Since in a number of species investigated tracheids appeared
among the prothallial cells, it was not difficult to determine the
apogamous forms. In all the apogamous species a region com-
posed of small cells made its appearance posterior to the apical
notch. In every instance when such an area of small cells ap-
peared on the surface of the prothallium, the embryo proved to
be of apogamous origin. When archegonia were produced in any
of the apogamous species, they were absent on many of the pro-
thallia. These, however, always produced embryos apogamously.
In some species longitudinal sections of the prothallia were made
in order to determine with more certainty the origin of the embryo.
96 STEIL: SOME NEW CASES OF APOGAMY IN FERNS
APOGAMY IN PELLAEA AND NOTHOLAENA
The first case of apogamy in the genus Pellaea was discovered
by Goebel (1905) in P. nivea (Poir.) Prantl. Later Woronin
(1907, 1908) found apogamy in P. flavens (Sw.) C. Chr., and P.—
tenera (Gill.) Prantl., and also in Notholaena Eckloniana Kunze
and N. sinuata (Lag.) Kaulf., belonging to a genus closely related
to Pellaea. Berggren (1888), however, had already described
- apogamy in JN. distans R. Br. The writer has described apogamy
in Pellaea atropurpurea (L.) Link (Steil, 1910) and P. adiantoides
J.Sm. (Steil, 1915). New cases have since been found in P. atro-
purpurea var. cristata Trelease and P. viridis (Forksk.) Prantl.
APOGAMY IN PTERIS
Farlow (1874) discovered apogamy in P. cretica L. var. albo-
lineata Hort. This was the first reported case in plants. Up to
the present time, no other case of apogamy has been reported in
any other species of Péeris, although Stephens and Sykes (1910)
assumed that apogamy occurred in Pteris Droogmantiana L.
Linden, on account of the presence of binucleate cells in the pro-
thallia.
Wigand (1849) appeared to be convinced that the fern embryo
did not owe its origin to fertilization. He undoubtedly described
apogamous embryos, and from his descriptions and figures it is
probable that he studied the development of such embryos in
some species of Pteris. Although Wigand gives a good descrip-
tion of an apogamous embryo, the true nature of apogamy was
first recognized by Farlow.
Tracheids were observed in the prothallium of P. sulcata
Meyen by Leszezyc Suminski (1848). Later Mercklin (1850)
confirmed the observation. Neither, however, knew the signifi-
cance of the presence of such sporophytic tissue elements in the
cells of the gametophyte. DeBary (1878) grew the prothallia of
P. quadriaurita Retz. var. argyraea Moore but failed to find the
fern apogamous.
Several years ago, without a knowledge of the observations of -
these investigators, the writer discovered apogamy in P. sulcata.
Spores were obtained from the New York Botanical Garden, and
STEIL: SOME NEW CASES OF APOGAMY IN FERNS 97
from Dr. E. B. Copeland, Los Banos, Philippine Islands. A
plant obtained from Mr. Anderson, fern specialist, Short Hills,
New Jersey, was grown in the university greenhouse, and spores
were also collected from this plant. Archegonia were never found
on any of the prothallia. However, apogamous embryos in large
numbers were produced. Since numerous cultures of P. sulcata
were made and from spores obtained from different plants, there
can be no doubt that apogamy is of constant occurrence in the fern.
P. argyraea Moore has also been found by the writer to be
apogamous. In many respects the prothallia and apogamous
embryos are similar to those of P. sulcata.* In P. Parkeri Hort.
The writer has also discovered apogamy. The prothallia become
large as compared with those of the former but develop in a similar
manner. :
During the course of the investigation a large number of P.
cretica varieties have been tested for apogamy, and so far none
have been found which form embryos as a result of fertilization.
The following is a list of these apogamous horticultural varieties:
albo-lineata Alexander, maxima, magnifica, Mayii, major, Wim-
settiit, Wimsettit compacta, Wimsettit multiceps, Wimsettii grandis,
and Ouvrardi. For the identification of some of the above var-
ieties the writer is indebted to Mr. James C. Clark, of Philadelphia.
P. cretica var. albo-lineata Alexander Hort. resembles the P. cretica
albo-lineata in which Farlow discovered apogamy only in its second
set of leaves, which are linear but with a broader band of white
along the main veins of the pinnae. The first leaves are nearly all
crested, while the linear leaves show sometimes a slight tendency
to become crested. From the investigations which have already
been made, it may be predicted that all Pteris cretica forms are
apogamous. Apogamy has not so far been found in any of the
varieties of P. serrulata L. f.
APOGAMY IN ASPIDIUM
Apogamy was found by De Bary (1878) in Aspidium falcatum
(L. f.) Sw. and in a crested cultivated variety of A. Filix-mas (L.)
* According to Christensen (Ind. Fil. 593, 608. 1906) P. sulcata, P. quadriaurita
and P. argyraea are all synonyms of P. biaurita L. According to Underwood and
Benedict (Bailey’s Stand. Cycl. Hort. 2852. 1916) P. quadriaurita is distinct from
P. biaurita, P. argyraea being given as a var. argyraea Hort. under P. quadriaurita.
98 STEIL: SOME NEW CASES OF APOGAMY IN FERNS
Sw. (A. Filix-mas cristatum’’). In the former Miss Allen (1911)
described nuclear and cell fusions in the sporangia, previous to the
formation of the spores. Kny (1895) discovered apogamy in an un-
crested form of A. Filix-mas (‘‘A. Filix-mas genuinum”’). Lang
(1898) found apogamy in the aberrant varieties of A. Filix-mas:
known as ‘‘ Nephrodium pseudo-mas var. polydactylum Wills”’ and
N. pseudo-mas var. polydactylum Dadds.”’ In these same varieties
(discussed under the name ‘‘ Lastrea pseudo-mas var. polydactyla’’),
Farmer, Moore and Digby (1903) and Farmer and Digby (1907)
described remarkable nuclear fusions in the prothallium before
the formation of the apogamous embryos. In a preliminary note
on apospory Miss Digby (1905) had already reported apogamy in
“ Lastrea pseudo-mas var. cristata apospora Druery.”’
Heilbron (1901) found apogamy in Aspidium aculeatum (L.)
Sw. var. cruciato-polydactylum Jones and A. angulare Willd.
forma grandidens Moore. Five years later (Steil, 1915 a and d)
apogamy was reported in A. hirtipes Bl. (Nephrodium hirtipes
Hook.), A. Tsus-Simense Hook. and A. chrysolobum Kaulf.
(Lastrea chrysoloba Presl). Apogamy has also been discovered
as a result of the investigations herein described in A. varium
(L.) Sw., A. auriculatum (L.) Sw., A. caryotideum Wallich, Cyr-
tomium Fortunei J. Sm. and C. Rochfordianum Hort.*
THE DEVELOPMENT OF THE PROTHALLIA AND SEX ORGANS
In all the species so far studied, the prothallia become typically
heart-shaped. Between the prothallia of apogamous and non-
apogamous species no difference was noted excepting that in the
latter tracheids sometimes appear. The prothallia of all the
species of Aspidium in which apogamy has been discovered bear
glandular hairs on both surfaces and on the margins (PLATE 4,
FIGS. I, 2,and 5; PLATE 5, FIGS. 20 and 21), while in both Pellaea
(PLATE 5, FIGS. 13, 14, and 15) and Pteris (PLATE 4, FIGS. 3, 4,
and 6) such hairs are always absent. The prothallia in Aspidium
grow to a much larger size than in Pellaea and in most species
* Benedict (Bailey’s Stand. Cycl. Hort. 2852. 1016) gives C. ca
asasynonym of C. falcatum J. Sm. (= A. falcatum Sw.); Christensen (Ind. F 1. 460.
1916) includes both A. caryotideum and Cyrtomium Fortunei among the synonyms of
the same species.
STEIL: SOME NEW CASES OF APOGAMY IN FERNS 99
of Pieris. Those of Aspidium chrysolobum, A. varium, and A.
auriculatum become especially large.
Antheridia are produced on the prothallia of all of the apoga-
mous species. The antherozoids responded to the chemotactic
influence of the archegonia of non-apogamous species in all cases in
which tests were made. The mature antherozoid appears perfectly
normal and is probably capable of functioning. Ina former note
(Steil, 1910) it was reported that antheridia had not been observed
on the prothallia of Pellaea atropurpurea, but in many of my cul-
tures made since this time they have been formed in large numbers.
Archegonia have been found on the prothallia of Aspidium
chrysolobum, but in a large number of the prothallia they are never
produced (PLATE 4, FIG. 5). The embryo always appears at the
anterior portion of the cushion and can be readily observed to
begin its development as a vegetative outgrowth from the pro-
thallial cells. Over 50 per cent. of the prothallia of Pellaea viridis
bear archegonia. Whether embryos are produced as a result of
fertilization in either case has not been determined. In a few
instances two embryos were observed to develop from a single
prothallium of P. viridis. One of these was apogamously pro-
duced, but the other appeared to owe its origin to an egg. Arche-
gonia were never found.in the prothallia of P. adiantoides, and for
this reason especial care was exercised in making the cultures of
P. viridis. Spores were obtained from the New York and the
Missouri Botanical Gardens and from two plants grown in the
university greenhouse, but the embryos in all of the cultures made
from the spores thus obtained were produced apogamously. In
some of the Pteris cretica varieties archegonia were observed very
rarely, but an embryo was never found to be developed from an
ege.
THE DEVELOPMENT OF THE APOGAMOUS EMBRYO
The apogamous embryo usually appears as a compact region
of small cells on the ventral side of the prothallium and posterior
to the apical notch (PLATE 4, FIGS. 2-5, and PLATE 5, FIGs. 13
and 20). When the embryo begins its development, the prothal-
lium has not yet attained its maximum growth. In some species,
such as Aspidium chrysolobum (PLATE 4, FIGS. I and 2), A. hir-
100 STEIL: SOME NEW CASES OF APOGAMY IN FERNS
‘ tipes and A. auriculatum, the prothallia increase considerably in
size as the young embryo is developing. In most instances the
cushion has not been formed when the embryo makes its appear-
ance. The embryo proceeds in development with the growth of
the cushion. On account of the early appearance of the embryo.
it was easy to determine when apogamy occurred in any species,
While the embryo usually occurs back of the apical region
its position varies in the different species and even in the same
species. In some cases the embryo is developed directly in the
apical notch (PLATE 5, FIGS. 14. and 15). A cylindrical or conical
process produced as an outgrowth of the apical region may bear
on some portion of it the apogamous embryo. Sometimes the
embryo may be produced at a considerable distance posterior to the
notch (PLATE 5, FIGS. 13 and 20). In still other instances the
embryo is formed on the lobes of the prothallium (PLATE 5, FIG.
19). From the foregoing, it is seen that the apogamous embryo
can be produced on portions of the prothallia where archegonia
have never been observed to be formed in any non-apogamous
species.
Tracheids are visible among the prothallial cells of some of the
apogamous species long before the prothallium has reached its
maximum growth (PLATE 4, FIG. 3, and PLATE 5, FIG. 20).
It is usually in the portion of the prothallium where the
tracheids appear that the apogamous embryo begins its develop-
ment. The tracheids are readily observed in many cases, since
the prothallium frequently becomes pale in the region of the notch
and where these elements are produced. The cells in this portion
of the prothallium contain fewer chloroplasts than the neighboring
prothallial cells (PLATE 5, FIG. 20). In the species of Pteris the tra-
cheids are most frequently observed (PLATE 4, FIGS. 3, 4, and 6).
In many instances the light area extends forward as a cylindrical
or conical process, already mentioned (PLATE 4, FIG. 6).
The embryo is usually surrounded by hairs, each composed ’of
a single row of cells (PLATE 5, FIG. 13). In some species, as in
Aspidium chrysolobum and A. Tsus-Simense, scales are also pro-
duced.
In only a few species has the development of the apogamous
embryo been studied in sections. It seems that the apical cell of
STEIL: SOME NEW CASES OF APOGAMY IN FERNS 101
the leaf always appears first, then that of the root, and finally that
of the stem. A foot has never been observed to develop in any
of the species in which the embryos were studied from prepared
slides. Usually the leaf is much in advance of the root (PLATE 4,
FIG. 9). A leaf, however, may be just making its appearance after
the root has grown considerably in length (PLATE 4, FIG. 8).
Both root and leaf, in the large majority of cases, are produced on
the ventral side of the prothallium (PLATE 4, FIG. 7). PLATE 4,
FIG. 12, represents a prothallium with an apogamous embryo the
leaves and roots of which are ventral. Either root or leaf or both
may appear on the dorsal surface (PLATE 4, FIGS. 10 and 11, and
PLATE 5, FIG. 16). In the cultures of Aspidium chrysolobum and
A. hirtipes embryos of this nature have been observed. Such
anomalies may be produced by light conditions as Leitgeb (1885)
has shown.
As a result of studies thus far made, it has not been determined
whether the apogamous embryo owes its origin to a single super-
ficial cell or to inner and outer cells of the prothallium.
Frequently, in some of the cultures, more than one embryo was
observed to develop from a single prothallium. This was espe-
cially the case when the prothallia showed a tendency to become
lobed (PLATE 4, FIGS. 3 and 4). PLATE 4, FIG. 3, represents a
prothallium of Pteris cretica albo-lineata Alexander with a young
embryo posterior to the apical region, and tracheids in each of
the two secondary regions of growth. Another prothallium of
the same variety is shown in PLATE 4, FIG. 4. In this instance
tracheids are present in the apical region, and two embryos have
been produced on other portions of the prothallium. One of these
is produced in the apical region of one of the main lobes, and the
other is a vegetative growth on the inner margin of a lobe itself.
Each of these embryos has produced a small leaf, J, and a root, r.
The embryos in such cases, as is readily seen, are wholly independ-
ent of one another. In some instances it was observed that some
of the tracheids extended from the apical notch of the prothallium
to one or both of the inner margins of the lobes where the embryo
was produced (PLATE 5, FIG. 19). It may be stated in this con-
nection that embryos are not always produced when tracheids are —
formed, but in most cases these tissue-elements indicate the be-
ginning of an apogamous embryo.
102 STEIL: SOME NEW CASES OF APOGAMY IN FERNS
In one of the cultures of Pteris sulcata, conical and nearly
spherical projections were observed in the lobes of the prothallia.
Some of these prothallia were transferred to another culture, and
the growth of the projections was followed in both cultures. The
projections produced either secondary prothallia or apogamous
embryos. The former were often cylindrical at the point of origin,
but in other respects they resembled the ordinary prothallia of
Pieris sulcata. As many as six embryos were observed on the
lobes of a single prothallium. In most instances these were nor-
mal, producing both roots and leaves.
By cultural conditions secondary prothallia have been pro-
duced from the primary prothallia of many of the apogamous
species. These prothallia also form embryos apogamously.
DeBary (1878) reported that such prothallia of Pteris cretica
albo-lineata seldom produced embryos. However, in my cultures
of the same species, the secondary prothallia usually produced
embryos of apogamous origin. Secondary prothallia of Pellaea
atropurpurea, Aspidium hirtipes, Pteris sulcata, Pteris argyraea,
and Pteris cretica albo-lineata Alexander frequently produce apoga-
mous embryos.
THE INFLUENCE OF WEAK ILLUMINATION ON THE DEVELOPMENT
OF THE- PROTHALLIA AND OF THE APOGAMOUS EMBRYOS
- When the prothallia of the apogamous species were placed
under the influence of weak light, the same results were obtained
as with the non-apogamous ferns. Filaments were formed from
the margin and both surfaces of the prothallia (PLATE 5, FIG. 21),
and these under the normal conditions of illumination in the war-
dian case became, independent prothallia, which in nearly all
instances formed also apogamous embryos. When the prothallia
were maintained in weak light, they remained simple or branched
filaments, producing neither sex-organs nor embryos (PLATE 5,
FIG. 18). Under somewhat more favorable conditions of illumi-
nation, ribbon-like plates were produced, which frequently bore
numerous antheridia and _ occasionally apogamous embryos.
When the illumination was slightly less than that in the greenhouse,
the prothallia became lobed (PLATE 5, FIGS. 14 and 15). In the
new apical regions which were formed embryos also made their
appearance (PLATE 5, FIG. 17).
STEIL: SOME NEW CASES OF APOGAMY IN FERNS 103
The conical or cylindrical processes already described grew
considerably in length when the cultures were placed in weak
light. In such instances an apical cell could be readily distin-
guished. The embryo in these cultures was frequently formed on
the process, and usually it was produced as a direct outgrowth of
the apical notch. The ‘‘light’’ area, or pale portion of the pro-
thallium where the embryo begins its development, remains more
conspicuous under these conditions. The colorless plastids,
present in large numbers in the cells of the pale region, become
chloroplasts under favorable conditions of light, and hence the
nearly colorless region is not so clearly differentiated in the latter
case.
ATTEMPT TO INDUCE APOGAMY IN OSMUNDA REGALIS
In the latter part of July, 1912, a large number of prothallia of
Osmunda regalis were found by the writer in a swamp in the vicinity
of Madison. Most of the prothallia at this time were small but
had produced numerous antheridia. Some of the prothallia were
removed with a depth of about three inches of soil and placed under
bell jars in the university greenhouse. Several cultures were kept
in a Wardian case where the illumination was very favorable for
the normal development of the prothallia. Other prothallia
were placed under bell jars and in different parts of the greenhouse
where strong light was obtained for the greater part of the year.
The latter were watered only from below and great care was exer-
cised to prevent condensation of moisture on the prothallia. In
this manner fertilization was prevented for nearly a year and a
half. During this period, however, the prothallia grew to a large
size, and numerous antheridia and archegonia were produced.
Many of the prothallia reached a length of three centimeters.
On such prothallia most of the archegonia were formed in acropetal
succession, but frequently a number were produced among the
older archegonia. In one instance the archegonia on one side of
the “midrib” of a prothallium, measuring two centimeters in
length, were counted and approximately five hundred were found
to be present. Therefore this prothallium had produced about
one thousand archegonia.
When the prothallia were freely watered, embryos were pro-
.
104 STEIL: SOME NEW CASES OF APOGAMY IN FERNS
duced in large numbers as a result of fertilization, Hence it is
certain that the cultural conditions which were maintained ren-
dered fertilization impossible. The prothallia grown under favor-
able conditions were smaller, but produced numerous sex organs
and embryos only as a result of fertilization.
Since the prothallia for the cultural work were found growing
under plants of Osmunda regalis, and since no other osmundas
were growing in the immediate vicinity, it could hardly be assumed
that the prothallia could be referred to any other species. Never-
theless, spores of O. regalis were collected and sown in the Wardian
case, and in every respect the prothallia grown corresponded to
those brought from the field.
The experimental work with Osmunda regalis was of especial
interest, since Leitgeb (1885) reported in this species the occa-
sional occurrence of apogamy. No one, however, has confirmed
the observation. Although Lang (1898), Yamanouchi (1908),
Pace (1913), and Nagai (1914) believed that apogamy might be
brought about by cultural conditions, Miss Black (1909) and
Mottier (1915) were unable to produce a single apogamous embryo
by cultural conditions.
SUMMARY
1. The prothallia of a number of species of ferns in which
apogamy was discovered were grown under cultural conditions
favorable for the development of sex-organs and embryos in non-
apogamous species.
2. The prothallia of all the apogamous ferns become heart-
shaped before the formation of the embryo. Antheridia are pro-
duced on the prothallia of all apogamous forms, but archegonia
are formed on the prothallia of only a few forms.
3. The embryo usually appears as a compact region of cells
posterior to the apical notch and on the ventral side of the pro-
thallium. In a number of species tracheids are visible among the
prothallial cells in the pale portion of the gametophyte.
4. First to make its appearance is the apical cell of the leaf,
then that of the root, and later that of the stem. A foot has not,
so far, been observed to develop in connection with the apogamous
embryos.
STEIL: SOME NEW CASES OF APOGAMY IN FERNS 105
5. Either root or leaf or both of these organs may develop on
the dorsal side of the prothallium. As a rule, however, they are
produced on the ventral side.
6. While the embryo is produced as a rule posterior to the
apical notch, it may be formed on a cylindrical or conical “ pro-
cess’’ and in some instances on the lobes of the prothallium.
7. Several apogamous embryos may be formed on a single
prothallium.
8. As in non-apogamous species, secondary prothallia are
readily produced, and these form embryos like those of the ordi-
nary prothallia.
g. The “‘light’’ area present on the prothallium of some of the
apogamous species is rendered more conspicuous in cultures main-
tained in weak light. The conical or cylindrical “process’’ in-
" creases considerably in length when the prothallia are grown under
these conditions. As a result of weak illumination, the embryo is
frequently produced as a direct outgrowth of the apical region of
the prothallium.
10. By growing the prothallia of Osmunda regalis in strong
light and preventing fertilization for a year and a half, no embryos
were produced apogamously.
11. An investigation extending over a period of six years has
resulted in the discovery of apogamy in a large number of ferns.
The conclusion that apogamy is of frequent occurrence in the
genera Pellaea, Pteris, and Aspidium, is justified on the basis of
the many cases so far found in these genera.
I wish to thank Professor C. E. Allen for suggestions and criti-
cisms received during the progress of the foregoing investigation.
UNIVERSITY OF WISCONSIN,
MapIsoN, WISCONSIN
LITERATURE CITED
Allen, R. F. (1911). Studies in spermatogenesis and apogamy in
ferns. Trans. Wisconsin Acad. 17: I-56. pl. I-6.
Bary, A. de. (1878). Ueber apogame Farne und die Erscheinung
der Apogamie im Allgemeinen. Bot. Zeit. 36: 449-487. pl. 14.
Berggren, S. (1888). Ueber Apogamie des Prothalliums von Noto-
chlaena. Bot. Centralblatt 35: 183-184. f. 1-3.
106 STEIL: SOME NEW CASES OF APOGAMY IN FERNS
Black, Caroline A. (1909). The development of the imbedded an-
theridium in Dryopteris stipularis (Willd.) Maxon and “‘ Nephrodium
molle.” Bull. Torrey Club 36: 557-572. pl. 26-28.
Digby, L. (1905). On the cytology of apogamy and apospory. II.
Preliminary note on apospory. Proc. Roy. Soc. 76B: 463-467.
f. I-3.
Farlow, W. G. (1874). An asexual growth from the prothallus of
Pieris cretica. Quar. Jour. Micr. Sci. 14: 266-272. pl. 10, 11.
Farmer, J. B., Moore, J. E. B., & Digby, L. (1903). On the cytology
of apogamy and apospory. I. Preliminary note on apogamy.
Proc. Roy. Soc. 71: 453-457. f. I-4.
Farmer, J. B., & Digby, L. (1907). Studies in apospory and apogamy
in ferns. Ann. Bot. 21: 161-199. pl. 16-20.
Goebel, K. (1905). Kleinere Mitteilungen. 3. Aposporie bei Asple-
nium dimorphum. Flora, 95: 239-244. f. I-3.
Heilbron, A. (1910). Apogamie, Bastardierung und Erblichkeits-
verhaltnisse bei einigen Farnen. Flora ror: 1-42. f. I-43.
Kny, L. (1895). Entwickelung von Aspidium Filixmas Sw. Bot.
Wandtaf. 411-438. pl. 93-100.
Lang, W. H. (1898). On apogamy and the development of sporangia
upon fern prothallia. Phil. Trans. Roy. Soc. London 190B: 187-
239. pl. 7-11.
Leitgeb, H. (1885). Die Sprossbildung an apogamen Farnpro-
thallien. Ber. Deutsch. Bot. Gesells. 3: 169-176.
Leszczyc Suminski, J. (1848). Zur Entwicklungsgeschichte der Farrn-
kraiiter. Berlin.
Mercklin, K. E. (1850). Beobachtungen an dem Prothallium der
Farnkrauter. St. Petersburg.
Mottier, D. M. (1915). Boebachtungen iiber einige Farnprothallien
mit Bezug auf eingebettete Antheridien und Apogamie. Jahrb.
Nagai, Isuburo (1914). Physiologische Untersuchungen iiber Farn-
prothallien. Flora 106: 281-330. f. 2
Pace, Lula (1913). Some peculiar fern prothallia. Bot. Gaz. so:
49-58. f. I-81. :
Steil, W. N. (1911). Apogamy in Pellaea atropurpurea. Bot. Gaz.
52: 400, 401.
- (1915a). Apogamy in Nephrodium hertipes. Bot. Gaz.
59: 254, 255-
——————. (1915). Some new cases of apogamy in ferns. Pre-
liminary note. Science II. 41: 293-294.
a
STEIL: SOME NEW CASES OF APOGAMY IN FERNS 107
Stephens, E. L., & Sykes, M. G. (1910). Preliminary note on apog-
amy in Pteris Droogmantiana. Ann. Bot. 24
Wigand, A. (1849). Zur Entwicklungsgeschichte der Farrnkriuter,
Bot. Ztg. 7: 105-116.
Woronin, Helene (1907). Apogamie und Aposporie bei einigen
Farnen. Ber. Deutsch. Bot. Gesells. 25: 85, 86.
. (1908). Apogamie und Aposporie bei einigen Farnen.
Flora 98: 101-162. f. I-72.
Yamanouchi, S. (1908). Apogamy in Nephrodium. Bot. Gaz. 45:
289-318. pl. 9, 10 +f. I-
Description of plates 4and 5
The photomicrographs of the prothallia on PLATE 4, and FIGS. 13, 14, 15, and 18
on PLATE 5, represent a magnification of about twenty diameters. FIGs. 19, 20, and
I on PLATE 5 show a much higher magnification. The apogamously produced
sporophytes were magnified about two and one half times. All the figures were re-
duced one seventh in reproduction.
PLATE 4
Fic. 1. A ventral view of a prothallium of Aspidium chrysolobum just before the
nate of the apogamous embryo.
Fic. 2. similar view of a prothallium of the same species. The apogamous
atau: is represented as a black region posterior to a pale region in the apical portion
of 8 dee sien
A ventral view of a prothallium of Pteris cretica albo-lineata Alexander.
In - ae region a young embryo is developing. In the apical regions established
in the lobes tracheids are present. _
Fic. 4. A ventral view of a prothallium of Pteris cretica albo-lineata Alexander.
An embryo with root and leaf has been produced in the new apical phateg of one of
the lobes. On the inner margin of the oener lobe a young embryo has also appeared.
In the main yeas region tracheids are visible
Fic. 5. rsal view of a prothallium ot Aspidium chrysolobum. The prothal-
lium produced no archegonia. The apogamous embryo appears as a dark region
posterior to the apical notch.
Fic. 6. A prothallium of Pieris cretica albo-lineata. A tongue-like portion has
developed as an outgrowth of the apical region. In the larger secondary prothallium
he * 20 ’’ region has already appeare
. 7. A ventral view of a prothallium of Aspidium hirtipes with an embryo
i root and leaf are produced on the ventral surface
Fic. 8. An embryo of A. hirtipes witha ‘eell Rewtion ve root.
Fic. 9. An embryo of A. hirtipes with only the leaf well developed.
Fic. 10. An embryo of A. chrysolobum, one root of which is on the dorsal surface,
and the other on the ventral
Fic. 11. An — 0 of a chrysolobum whose root is on the dorsal surface and
leaf on the ventral s
Fic. 12. A young sont of A. hirtipes with ice primary leaves and a
onee ventral primary r
108 STEIL: SOME NEW CASES OF APOGAMY IN FERNS
PLATE 5
Fic. 13. A ventral view of a prothallium of Pellaea adiantoides. The young
embryo is surrounded by hairs
Fic. 14. A prothallium of Pélines adiantoides grown in slightly weaker illumina-
tion than that maintained in the Wardian case. The prt has become dis
tinctly lobed, and the embryo is developed in the apical reg
Fic. 15. A similar prothallium of the same species. he lobes are still more
d the embryo has grown to a larger si
16. An embryo of Aspidium se scents one Ae of which is on the dorsal
surface and the other on the ventral surface of the prothallium. The long root is
also on the ventral surface.
Fic. 17. A portion of a lobed prothallium of A. hirtipes bearing two embryos,
each of as has a well-developed leaf and root.
G. 18. Filaments of a single row of cells of prothallia of A. hirtipes grown in
weak oes ion.
FIG - A prothallium of Pteris sulcata, showing the beginning of an embryo in
the ue sna Tracheids pass upward to the young embryo, which has been pro-
duced on the inner margin of a |
Fic. 20. An embryo of A. ee aes beginning its development i in the apical region.
The ‘ hog region and tracheids are clearly differentiated.
FIG. ondary prothallia of A. hirtipes produced from the margins and sur-
faces of a poultice placed in weak light.
Correlation of morphological variations in the seedling of
Phaseolus vulgaris
J. ARTHUR HARRIS AND B. T. AVERY
INTRODUCTORY REMARKS
During the past several years one of us has had under way
extensive experiments on the differential death-rate of bean seed-
lings. Individuals differing in structure also differ in their capa-
city for survival under field conditions,* and in such physiological
characteristics as capacity for the development of the tissues of the
primordial} and of the subsequent leaves.t{
Some tens of thousands of seedlings of known morphologica]
characteristics have been exposed to risk, as the life insurance
statisticians express it, in an attempt to determine the selective
value of the various morphological variations. These seedlings
were, for technical reasons, necessarily planted in the field at a
time when the cotyledonary node and the primordial node only
could be studied. It is evident that the capacity of the plant for
survival may be in some degree dependent upon characters de-
veloped later in ontogenesis, but correlated with characters of the
first or second node of the seedling.
However this may be, it is certainly true that a full knowledge
of the morphology and physiology of the variant bean seedling
demands a thoroughgoing investigation of the correlation between
the structure of the first two leaf whorls and that of later whorls.
We have, therefore, been forced to consider the problem of the
‘morphological character of the leaf whorls produced at the third
* Harris, J. Arthur. A simple demonstration of the action of natural selection.
Science II. 36: 713-715. 1912.
t Harris, J. Arthur. Studies on the correlation of morphological and physiologi-
cal characters: the development of the primordial leaves in teratological bean seed-
lings. Genetics 1: 185-196. 1916.
¢ Harris, J. Arthur. Further studies on the interrelationship of morphological
and ee characters in seedlings of Phaseolus. Mem. Brooklyn Bot. Gard.
I, in press
109
110 HARRIS AND AVERY: MORPHOLOGICAL VARIATIONS
node in the case of plants showing various structural abnormalities
at the first two nodes.
Phaseolus is well suited for such investigations. The normal
seedling has two cotyledons, inserted at the same level, and two
opposite primordial leaves. A large number of structural varia-
tions, four types of which will be considered in this paper, may
occur. The chief disadvantage lies in the rarity of many of the
variations in the lines with which we have dealt. The securing of
adequate series is excessively laborious. The present paper is
based upon a careful study of the variations in the first three nodes
of 16,348 plants, which were selected from about 450,000 seedlings
examined for the characters of the first and second node.
When in the following paragraphs we refer to normal and
abnormal plants or seedlings, it must be understood that this
applies to the characteristics of the individual as determined ‘on
the basis of the first two nodes, the cotyledonary and the pri-
mordial only. In its later development the “abnormal” plant
may remain ‘“‘abnormal’’ or become “normal,’’ and the ‘‘normal’’
plant may either continue to be “normal” or become “abnormal.”
The nature and method of Classification of the abnormalities
dealt with will be discussed in the presentation of the data below.
MATERIALS AND METHODS
The materials upon which this study is based are a series of
lines of White Navy beans grown at the Station for Experimental
Evolution during the past several years. The seeds were har-
vested from field cultures in 1915 and germinated in sand in the
autumn of 1916.
Seedlings which were abnormal in the characters of the first
or second node, i. e., in the number or insertion of the cotyledons
or of the primordial leaves, were sorted out for potting in soil and
subsequent study of the third node, that normally giving rise to
the first compound leaf.
For each abnormal individual, a normal control seedling from
the same parent plant was taken at random to serve as a basis of
comparison. Both were potted in soil and grown to a stage when
the characteristics of the third node could be accurately deter-
mined.
HARRIS AND AVERY: MORPHOLOGICAL VARIATIONS 111
For the onerous preliminary examination of nearly a half a
million seedlings we are greatly indebted to Miss Edna K. Lock-
wood, Miss Margaret Gavin and especially to Miss Lillie Gavin.
PRESENTATION AND ANALYSIS OF DATA
The slightest abnormality which we have been able to discover
occurring in considerable numbers of bean seedlings is the vertical
separation of the two normally opposite cotyledons. So imper-
‘ceptible is the line of transition between normal and abnormal that
personal equation must play some part in classification. The
cotyledons may be much more widely separated. The variation
is a purely graduated one, with no sharp lines of demarcation be-
tween the different degrees of separation. Generally we have
recognized three grades, but because of the rarity of plants with
more widely separated cotyledons we have in this paper grouped
our data into two classes only. The first comprises plants with
cotyledons 2-3 mm. apart. The second includes all those in which
they are more distant.
The number as well as the position of the cotyledons may vary.
Plants with three instead of two cotyledons fall into two groups;
those with the normal pair of primordial leaves and those with a
whorl of three leaves. The latter are by far the more abundant.
Abnormality developed subsequently to the selection of the
seedlings in the preliminary sorting may affect either the inter-
node between the second and the third nodes, that is, between the
primordial leaf whorl and the point of insertion of the first com-
pound leaf or leaf whorl, or it may be confined to the number or
structure (or number and structure) of the leaves inserted at the
third node.
In the original selection of individuals abnormal in the char-
acters of the first or second node, only those with sensibly normal
axes (hypocotyl and epicotyl) were chosen for the purposes of the
present study.
Two types of abnormality in the axis beyond the second (the
primordial) node have been considered.
The first is a sensible broadening of the axis, identical with or
similar to fasciation. This is a graduated character. The line of
demarcation between normal and abnormal is not clearly marked.
112 Harris AND AVERY: MORPHOLOGICAL VARIATIONS
and personal equation may influence in some degree the classifica-
tion of the seedlings.
The second is a division of the axis into two coérdinate branches
each with a terminal bud.
The frequencies of the two types of axial variation are too small
to justify detailed discussion. The entries in TABLE I show that
TABLE I
FREQUENCIES OF ABNORMALITY OF SECOND INTERNODE IN NORMAL AND ABNORMAL
SEEDLINGS
| Actual frequencies Percentage frequencies
| ——___—____—
. Broad-| Di-
ee eee. ‘Norma ened | vided | Normal | Broadened| Divided
inter- | inter- | internode | internode | internode
‘node node node
Two cotyledons slightly siseiuia: |
two primordial leaves.......... 4,017 5 8| 090.6774 .1240 -1985
PROFIAL CONEIOD. 6 eva ee fies o 4,029 (a) El GO.OFGE| feu near .0248
* erenCe ee Fb ee oe cs —12| + 5| + 7} —-2077| +. .1240| +.1737
Two cotyledons sien As grasa
two prim ye 55 leavea eGo yes 878 2) I} 99.6594 .2270 +1135
Normalcontrics 25 ccs eee ss 881 oO | ROO 0600 Soe Te ye.
Diff bes be 2 AGE oh 2990) 4 35
ere: cotyledons; two primordial
WOR ee Ve haces see ee 813 12 GO} O8;5454| 5.45401... ..
Peak PORNO fore a 825 ° © IO 00CG Fo a eo
Difference aed —I2} +12 @} 1.4546) 4+-1.4546] ......
Three cotyledons; three primordial .
VOR eae a5 .| 2,410 14 14} 98.8515 -5742 -5742
Norinal control... 2,436 2 0| 99.9179 ORI vies.
Diflerénce. 525 3 | —26| +12} +14) —1.0664| + .4922] +.5742
in every instance in which any individuals at all are available the
seedlings which are abnormal in either the cotyledonary or the
primordial node show a higher percentage of abnormality in the
structure of the internode beyond the second node than do the
normal controls.
We now turn to a consideration of variation in the leaves in-
serted at the third node. The leaves of plants with abnormality
of the axis should not be combined with those having normal axes,
They are not sufficiently numerous for separate consideration.
Confining our attention to seedlings which have a normal axis
for at least the length of the second internode of the epicotyl, we
Harris AND AVERY: MORPHOLOGICAL VARIATIONS 113
have the frequencies shown in TABLEs II-V.*. The character of the
control plants is also given.
TABLE II
SEEDLINGS WITH TWO COTYLEDONS SLIGHTLY SEPARATED AND TWO PRIMORDIAL
iS ES
Naatet of, Actual frequencies Percentage frequencies ae
| ifference
siete hil wie ot Abnormal Control Abnormal Control
oe: 3,791 3,853 94.37 95.63 —1.26
2 225 176 5.60 4.37 +1.23
3 I oO -02 -00 + .02
‘Potals.s 50s; 4,017 4,029 99-99 100.00
TABLE III
SEEDLINGS WITH TWO COTYLEDONS WIDELY SEPARATED AND TWO PRIMORDIAL
LEAVES
ilibae of Actual frequencies Percentage frequencies CG
acter aoe _ Abnormal Control Abnormal Control
I 81t 840 92.37 95-35 —2.98
2 67 4I 7.03 4.65 +2.98
Totals. i 878 881 100.00 100,00
TABLE IV
SEEDLINGS WITH THREE COTYLEDONS AND TWO PRIMORDIAL LEAVES
Numbes of Actual frequencies Percentage frequencies io IOs
AveE per Boe Abnormal Control Abnormal Control
I 591 792 72.69 96.00 —23.31
2 221 a4 27.18 4.00 +23.18
ee £ fs) Ae -00 ee
Totals...... 813 825 99-99 100.00
TABLE V
SEEDLINGS WITH THREE COTYLEDONS AND THREE PRIMORDIAL LEAVES
i P tage fi i
vcomiees ot Actual frequencies ercentage frequencies Rona
leaves per node Abnormal Control Abnormal Control
I 1,632 2,200 67.72 90.31 22.59
2 771 236 31.99 9.69 +22.30
= 7 oO +29 -00 299
Fotale, ..i:4 2,410 ° 2,436 100.00 1OO00) Pyke
* In these tables the numbers of control plants are not exactly identical with the
some of those selected as normal in the i
stage
here, where we are discussing abnormalities of foliar characters onl
114 Harris AND AVERY: MORPHOLOGICAL VARIATIONS
In each of the types of abnormality dealt with the abnormal
series show a higher proportion of the individuals with two or three
leaves at the third node than do their normal controls.
Furthermore, seedlings showing different types of abnormality
~ at the first nodes also differ among themselves in the extent of
abnormality at the third node. Thus plants which are normal
except for slight separation of the cotyledons have two or three
leaves at the second node instead of the single leaf normally found
in 5.63 per cent. of the individuals. Plants with the cotyledons
more widely separated have 7.63 per cent. of their number with two
or three instead of a single leaf.
When one turns to the groups of plants which have three in-
stead of twocotyledons, a conspicuous difference is at once apparent.
Plants which have three cotyledons and a normal pair of primordial
leaves produce two or three instead of a single leaf at the third
node in 27.31 per cent. of the cases. Seedlings with three coty-
ledons and a whorl of three primordial leaves instead of the normal
pair at the third node have 32.29 per cent. of the individuals with
two or three leaves at the third node.
Heretofore the number of leaves inserted at the third node has
furnished the only measure of variation at this region of the axis.
We now propose to consider variation in the organization of the
leaves themselves. It will not be possible to do this in the detail
in which we hope to treat the problem ultimately. The range of
variation in the division of the bean leaf is rather great, and the
laws governing it are doubtless very complicated. Some progress
has already been made on the problem, but for the present we shall
limit our discussion to the number of leaflets, leaving the problem
of their arrangement for treatment when even larger series of data
are at our disposal.
The actual frequencies of number of leaflets per leaf produced
at the third node are shown in TABLE VI.
The most conspicuous feature of this table is the bimodal
nature of the distribution. The modes are on three and Six, as is
to be expected from the fact that the distribution of the whole
number of leaflets depends upon plants with from one to three
leaves at the third node.
Because of the wide range of variation in leaflet number it is
HARRIS AND AVERY: MORPHOLOGICAL VARIATIONS 115
not feasible to redtice these frequencies for the individual classes
to a percentage basis for comparisons. This has, however, been
done for larger groups secured by combining all the seedlings
TABLE VI
NUMBER OF LEAFLETS PRODUCED AT THE THIRD NODE BY SEEDLINGS OF VARIOUS
TYPES
Two cotyledons Twocotyledons | Three cotyledons Three cotyledons
slightly —o =~ deg separated and and two and three
bp int ed two primordial leaves | two primordial leaves primordial leaves ‘} primordial leaves
Abnormal} Control | Abnormal | Control | Abnormal Control | Abnormal | Control
I 2 | I eS I
2 16 e 6 4 iis 3 2 2 5
x 3,741 3,825 801 835 572 782 1,602 | 2,185
4 27 at 5 4 I4 5 28 8
5 10 2 I 3 2
6 215 173 65 40 203 33 751 236
4 Meee I +2 I wets
8 ae e's I sigh
9 I ras I 3
10 I ae ee 2
Ea & v's ae ee I
Totals | 4,017 4,029 | 878 881 813 | 825 2,410 | 2,436
showing merely separation of the cotyledons and all those showing
three cotyledons instead of the normal two. The results are shown
in the accompanying TABLE VII, which is self explanatory.
TABLE VII
COMPARISON OF THE NUMBER OF LEAFLETS IN DICOTYLEDONOUS AND TRICOTYLEDO-
NOUS SEEDLINGS WITH THAT IN THEIR NORMAL CONTROLS
Number of Seedlings with cotyledons separated Seedlings with three cotyledons
leaflets
Abnormal Control Abnormal Control
I 12 .04 03
2 12 -16 21
a 92.79 94.91 67.45 90.98
4 -5t T.3 -40
5 -08 -4 15
6 5.72 3-52 29.60 8.25
<a 81 -74
8 -03
9 -02 32
10 .02 06
II -03
A comparison may be made without the combination of dif-
ferent grades of abnormality by grouping the number of leaflets
116 Harris AND AVERY: MORPHOLOGICAL VARIATIONS
around the modal classes 3,6 and 9. The results in TaBLe VIII
show essentially the same relationships as those given in TABLES
VI-VII. First, the higher leaflet numbers are more extensively
represented in the abnormal plants of each of the four types than
they are in the controls. Second, the tricotyledonous plants show
a far greater increase in the number of leaflets inserted at the third
node than do those abnormal only in the position at which the
two cotyledons are inserted.
TABLE VIII
PERCENTAGE FREQUENCIES OF NUMBERS OF LEAFLETS IN SEEDLINGS OF VARIOUS
TYPES
a
Number of leaflets
Class of abnormality
I-4 5-7 8-11
T. ty a 1 gl y Dp 4 4 primord al
IOAVOR hs a a Si es wey Pee eh se pee cles 94.32 5.62 .05
AOE oe Se ae ees tes es os 95.66 4.34 ape
Dieter ie as i pe eee — 1.34] + 1.28| +.05
ia PE OT ree | wa & A 1
sinh sia Y t a
VEE are WN See ee oe bees See eee ee 92.37 4.63
MRTG ene te se Ce wee ee Hew Ca ne ee ale ba 95.23 77
Dilerette is ee a ee eS ae — 2.8 + 2.86
Th tyled dtwop Hal leaves... oe. oss 72.57 27.18 24
Comirat. Soe a ee ee 95-64 4.
EBiif cross pier Were Carma eae | eter er Venera —23.07 | +22.82 +.24
Th tyled d ti Pp dial leaves. 2:..... 67.72 32.03 25
COOUIOR Se ea ees ee ee a ees 90.23 9.77 slay
Differenee oso oo re Po oe i et ee —22.51 | +22.26 +.25
In substantiation of these conclusions the reader will note that
in the class with slightly separated cotyledons 5.68 per cent. of the
plants have from five to ten leaflets as compared with 4.34 per
cent. with five and six leaflets in the control series. In seedlings
with more widely separated cotyledons but no other abnormality
there are 7.63 per cent. of the plants with five to seven leaflets as
compared with 4.77 per cent. of the normal controls with five or
six leaflets. Seedlings with three cotyledons but the normal number
of primordial leaves have 27.43 per cent. of the individuals with
from five to nine leaflets as compared with 4.36 per cent. with five
or six leaflets in the normal controls.
Plants with a trimerous cotyledonary and primordial whorl
have 32.28 per cent. of the seedlings with from five to eleven leaflets
HARRIS AND AVERY: MORPHOLOGICAL VARIATIONS 117
as compared with 9.77 per cent. with five and six leaflets in the
normal controls.
Taking the average number of leaflets per plant as a basis of
comparison between the abnormal plants and their controls we
find the results in TABLE IX.
TABLE IX
MEAN NUMBER OF LEAFLETS IN SEEDLINGS OF VARIOUS TYPES
a Mea
Class of abnormality — er of number of Difference
abnormals | controls
|
|
ee ee a aT oa coe ees mao Vcc es 3-170 3-133 | +0.037
Ne ie ea sar SAE) es 6 woe IE Awe ba a 3.228 3-191 +0.037
Three dae ais two primordial leaves............. 3-847 | 3.131 | +0.716
Th tyled three pri dial leaves........... 3-900, 3.204 | +0.606
Note (a) that for each type of normality the average number
of leaflets is greater in the abnormal individuals than in the nor-
mal, and (0) that the difference between the abnormal class and its
control is far greater in the case of the plants with three cotyledons
than in those in which the abnormality in the cotyledonary whorl
consists merely in the separation of the two cotyledons.
Thus the results for number of leaflets substantiates the con-
clusion based upon number of leaves.
Evidently, however, the number of leaflets is to a great extent
determined by the number of leaves. The problem now arises:
Are there differences in the average number of leaflets per leaf in
the abnormal and normal individuals? :
Means and their differences have been diternined, but are
so slight that conclusions must be deferred until further series of
data are available.
Just one other method of dealing with the problem of the cor-
relation in structural variation may now be considered.
Number of leaflets is, in the materials dealt with, practically
an integral variate. In examining a large series of plants those
with partial division of a leaflet, representing transition stages
between a leaf with » and one with m + 1 leaflets are sometimes
found. Such cases are, however, relatively rare. The lobing of
the leaflet has therefore been disregarded in the foregoing treat-
118 Harris AND AvERY: MORPHOLOGICAL VARIATIONS
ment. A leaf with three leaflets, one of which has a lobe, has been
recorded as 3 in the tables, not as an intermediate between three
and four. This has simplified the tabling of the data, and the cal-
culation of the simple constants necessary to the interpretation
the data, without any material loss in accuracy.
One may, however, inquire whether there are differences in the
degree of lobing of the leaflets produced at the third node in plants
which are normal and in plants which are abnormal in the char-
acters of the first and second node. Because of the very low per-
centages of lobing in the leaflet no stress whatever is to be laid
upon the exact values found, even in samples containing several
hundreds or thousands of plants, because of the great difficulties
of determining the probable error of a small percentage.
The results are given in TABLE X
TABLE X
PERCENTAGE OF LOBING IN THE LEAVES OF SEEDLINGS OF VARIOUS TYPES
: I Sontrol ‘
Cla fah lity peat ype Difference
_ leaves See Deere teense Se ee 0.349 0.273 +0.076
WORN CES 0.456 0.112 +0.344
Three cotyledons; two primordial leaves ............. I.599 0.242 +1.357
three primordial leaves ........... 0.954 0.328 | +0.626
Here the percentage frequency of plants with one or two lobes
on the leaflets* are given for each type of abnormality dealt with
and compared with that found in the control series.
Two relationships seem clearly indicated by the constants in
this table.
First, the tendency to the production of lobes is greater in the
leaflets produced by abnormal plants of all four types than in
their normal controls.
Second, the tendency to the production of lobes is greater in
the leaflets of plants with a trimerous cotyledonary whorl, and
either a dimerous or trimerous primordial whorl, than it is in
plants in which the sole abnormality consists in the separation of
the two cotyledons in their insertion on the axis of the plant.
* In the case of two lobes both may occur on the same leaflet or they may be on
different leaflets.
HARRIS AND AVERY: MORPHOLOGICAL VARIATIONS 119
RECAPITULATION
This paper presents the results of a first attempt to determine
some of the correlations in the structural variations of the seedling
of Phaseolus vulgaris.
rhe materials are drawn from a series of lines of Navy beans
grown for the past several years at the Station for Experimental
Evolution. The seeds used were harvested from plants of selected
ancestry. Neither of these factors will, we believe, invalidate the
conclusions drawn in this paper. These conclusions will not neces-
sarily apply to certain entirely abnormal races.
Fasciation-like broadening of the axis and longitudinal division
of the axis distal to the insertion of the primordial leaves are both
more frequent in seedlings showing separation of the cotyledons
and in tricotyledonous seedlings than in those which are normal.
Seedlings which are normal except for the separation of the
cotyledons and those which have three cotyledons and a normal
pair of primordial leaves or three cotyledons and a whorl of three
primordial leaves produce a larger number of leaves, a larger num-
ber of leaflets and a higher percentage of leaves with lobes at the
third node than do those which are normal in their cotyledonary
and primordial leaf characters.
Seedlings which are tricotyledonous, with either a normal pair
or a whorl of three primordial leaves, show higher percentages
of variation in the axis, or the leaves produced by the axis, distal
to the primordial leaves than do those which are normal except
for the separation of the two cotyledons.
These studies will be continued.
STATION FOR EXPERIMENTAL EVOLUTION,
CoL_p Sprinc Harsor, NEW YorK
INDEX TO AMERICAN BOTANICAL LITERATURE
1913-1918
The aim of this Index is to include all current botanical literature written by
Americans, published in America, or based upon American material ; the word Amer-
ica being used in the broadest sense
Reviews, and papers that ulate ee to forestry, agriculture, horticulture,
manufartared products: of vegetable origin, or laboratory methods are not included, and
Re n
Some important particular. If users of the Index will call the attention of the editor
to errors or omissions, their kindness will be appreciated.
This Index is reprinted monthly on cards, and furnished in this form to subscribers
at the rate of one cent for each card, Selections of cards are not permitted ; each
subscriber must take all cards published during the term of his subscription. Corre-
spondence arta to the card issue should be addressed to the Treasurer of the Torrey
Botanical Clu
Allen, C. E. The spermatogenesis of Polytrichum juniperinum.
Ann. Bot. 31: 269-291. pl. 15, 16. Ap 1917.
Allen, C. E., & Gilbert, E. M. Textbook of Botany. i-x + 1-459.
f. 1-223. Chicago. 1917. [Illust.]
Andrews, E. F. Agency of fire in propagation of longleaf pines. Bot.
Gaz. 64: 497-508. f. 1-5. 18 D 1917.
Arthur, J. C. Relationship of the genus Kuehneola. Bull. Torrey
Club 44: 501-511. 20 N 1917.
Includes Frommea gen. nov. and F. Polylepidis sp. nov so
Babcock, D. C. Diseases of ornamental plants. “Ohio Agr. Exp. Sta.
Month. Bull. 2: 323-328. O 1917. [Illust.]
Barnhart, J. H. John Eatton Le Conte. Am. Mid. Nat. 5: 135-138.
Beadle, C. Hedychium coronarium in Brazil. Kew Bull. Misc. Inf.
1917: 104, 105. Au I917.
Blake, S. F. Descriptions of new spermatophytes, chiefly from the
collections of Prof. M. E. Peck in British Honduras. Contr. Gray
Herb. Harvard Uniy. 52: 59-106. 5S 1917.
Includes one new genus and 50 new species.
Blake, S. F. New and noteworthy Compositae, chiefly Mexican.
Contr. Gray Herb. Harvard Univ. 52: 16-59. S 1917.
One new genus and nineteen new species are described.
122 INDEX TO AMERICAN BOTANICAL LITERATURE
Blake, S. F. Notes on the systematic position of Clibadium with de-
melas of some new species. Contr. Gray Herb. Harvard Univ.
1-8. S 1917.
Incl udes Clibadium See ice C. strigillosum, C. Sprucei, C. polygynum and .
C. divaricatum, spp. n
Blake, S. F. A revision of the genus Dimerostemma Cass. Contr.
Gray Herb. Harvard Univ. 52: 8-16. S 1917.
Dimerostemma asperatum sp. nov. is described.
Blake, S. F. Two new polygonums from New England. Rhodora
19: 232-235. 21 N 1917.
Polygonum achoreum and P. allocarpum, spp. nov.
Blake, S. F. The varieties of Chimaphila umbellata. Rhodora 1g.
237-244. 5 D 1917. ;
Bourquin, H. Starch formation in Zygnema. Bot. Gaz. 64: 426-434.
p27. 16 N ro17.
Brooks, C., & Cooley, J. S. Effect of temperature aération and hu-
-midity of Jonathan-spot and scald of apples in storage. Jour. Agr.
Research 11: 287-318. pl. 32, 33 +f. 1-23. 12 .N 1917
Brooks, S.C. Permeability of the cell walls of Allium. Bot. Gaz. 64:
509-512. 18 D 1917
Brown, M.M. The development of the embryo-sac.and of the embryo
in Phaseolus vulgaris. Bull. Torrey Club 44: 535-544. pl. 25, 206.
27 D 1617;
Brown, P. E. The importance of mold action in soils. Science II.
46: 171-175. 24 Au 1917.
Brown, W. H., & Heise, G. W. The application of photochemical
temperature coéfficients to the velocity of carbon dioxide assimila-
tion. Philip. Jour. Sci. 12: (Bot.) 1-25. f. 1-3. Ja 1917.
Brush, W. D. Distinguishing characters of North American sycamore
woods. Bot. Gaz. 64: 480-496. pl. 32-38 + f. 1-3. 18 D 1917.
Buchanan, R. E.. Studies on the nomenclature and classification of the
bacteria. Jour. Bact. 2: 603-617. N 1917.
Burt, E. A. Merulius in North America. Ann. Missouri Bot. Gard.
4: 305-362. pl. 20-22 + f. 1-39. N 1917. *
Includes descriptions of sixteen new species.
Choate, H. A. The earliest glossary of botanical terms. Torreya 17:
186-201. 30 N 1917.
Clute, W. N. Botany and common names of plants. Science II. 46:
483, 484. 16 N 1917.
Colley, R. H. Diagnosing white-pine blister-rust from its mycelium.
Jour. Agr. Research 11: 281-286. pl. 31 +f. 1. 5N 1917.
INDEX TO AMERICAN BOTANICAL LITERATURE 123
Copeland, E. B. New species and a new genus of Borneo ferns, chiefly
from the Kinabalu collections of Mrs. Clemens and Mr. Topping,
cae Jour.Sci. 32: te! moa Ja 1917.
Forty-four new species are descr
Cfoulter,], J. M. Ellsworth Fane Hill. Bot. Gaz. 64
15 Au 1917. [Portrait.]
Curjar, A. M. The adaptation of Truog’s method for the determina-
tion of carbon dioxide to plant respiration studies. Plant World 20:
gan-ag%..7,.15. 5 19tf.
Davis, W. E. Resistance of seed coats of Abutilon laa cee to intake
of water. Bot. Gaz. 64: 166, 167. 15 Au
i. 165, 106.
Dearness, J. New or noteworthy North eee fungi. Myco-
ceed 9: usobiae 24 N antl
Detjen, L. :. Poliacton of the rotundifolia grapes. ie Elisha
Mitchell Sci. Soc. 33: 120-127. N11
Dorsey, M. J. The Duchess apple ee Jour. Heredity 8:
565-567. 7.9. .-D 1917.
Douglass, [H.] B. Mushroom poisoning. Torreya 17: 171-175.
31 O 1917; 207-221. 24 Ja 1918.
Drennan, G. T. Volunteer plants. Am. Bot. 23: 86-88. Au 1917.
Dunham, E. N. Unusual habitats. Bryologist 20: 98, 99. N 1917.
Dupler, A. W. The gametophytes of Taxus canadensis Marsh. Bot.
Gaz. 64: 115-136. pl. 11-14. 15 Au 1917.
Durfee, T. Lichens of the Mt. Monadnock oem, N. H.—No. 9.
Bryologist 20: 99. N 1917.
Elliott, J. A. The conduction of potassium cyanide in plants.
pathology 7: 443-448. f. 1, 2. 3 D 1917.
Evans, A. W. A new Lejeunea from Bermuda and the West Indies.
Bull. Torrey Club 44: 525-528. pl. 24. 20 N 1917.
Lejeunea minutiloba sp
Evans, A. W. Pech ainacy list of Arizona Hepaticae.
60-62. 15 Au I917.
Faust, E. C. Resin secretion in Balsamorrhiza sagittata. Bot. Gaz.
64: 441-479. pl. 28-31 +f. 1, 2. 18 D 191
Fernald, M. L. A new alpine willow from the White Mountains.
Rhodora 19: 221-223. 21 N 1917.
Salix Peasei sp. nov
Fernald, M. L. The boreal and subalpine variety of Spiraea latifolia.
Rhodora 19: 254, 255. 5 D 1917.
Fromme, F. D. Tylenchus tritici on wheat in Virginia.
ogy 7: 452, 453, f. 1. 3 D 1917.
Phyto-
Bryologist 20:
Phytopathol-
124 INDEX TO AMERICAN BOTANICAL LITERATURE
Frye, T. C. Illustrated key to the western Ditrichaceae. Bryologist
20: 49-60. pl. I-19. 15 Au I917.
Gaines, E. F. Inheritance in wheat, barley and oat hybrids. Wash-
ington Agr, Exp. Sta. Bull. 135: 1-61. f. 1-9. Mr 1917.
Garman, H. A bean disease introduced in diseased seeds. Kentucky
Agr. Exp. Sta. Circ. 16: 91-95. f. zr. Jl 1917.
Anthracnose.
Gilbert, A. H., & Bennett, C. W. Sclerotinia trifoliorum, the cause of
stem rot of clovers and alfalfa. Phytopathology 7: 432-442. f. 1-5.
4 i) 19%7.
Gourley, J. H. Some observations on the growth of apple trees. New
Hampshire Agr. Exp. Sta. Bull. 12: 1-38. charts 1-9. Jl 1917.
Greenman, J. M. Two exotic Compositae in North America. Ann.
Missouri Bot. Gard. 4: 289-292. pl. 19. N 1917.
Grier, N. M. New forms of Calamites. Am. Mid. Nat. 5: 147-150.
pt. 2-3, N 1987,
Includes Calamites Fettermanni, C. multifoliotus, and C. cruciatus Harrisoni,
spp. nov.
Grier, N. M. Note on fruit of mountain magnolia. Rhodora 19:
anG, 5 LD igt?.
Giissow, H. T. The occurrence of Colletotrichum cereale, Dothichiza
populea and Leptosphaeria napi in Canada. Phytopathology 7:
450," 3 D:t017;
Halsted, B. D., & Owen, E. J- Environment of seeds and crop pro-
duction. Plant World 20: 294-297. S$ 1917.
Hanson, H.C. Leaf-structure as related to environment.. Am. Jour.
Bot. 4: 533-560. f. I-20. 24 N 1917.
Harger, E. B., and others. Additions to the flora of Connecticut.
Rhodora 19: 224-232. 21 N; 245-253. 5 D 1917.
Harper, R. M. Some North Carolina soil statistics and their signific-
ance. Jour. Elisha Mitchell Sci. Soc. 33: 106-1 19. N 1917.
Harris, J. A. On the distribution of abnormalities in the inflorescence
of Spiraea Vanhouttei. Am. Jour. Bot. 4: 624-636. pl. 30, 31 +
J. 4. 12'D 10917.
Harris, J. A. Sunspots, climatic factors and plant activities. Am.
Nat. 51: 761-764. D 1917.
Hendershot, L. B. The white or American elm. Nature Study Rev.
13: 298-301. O 1917.
Henry, J. K. Alberni notes. Ottawa Nat. 31: Bese. 0E N 1Gi7,
H W. An avocado monstrosity. Jour. Heredity 8: 557.
Jf. 'D 1917.
INDEX TO AMERICAN BOTANICAL LITERATURE 125
\
Hodgson, R. W. Some abnormal water relations in Citrus trees of the
arid southwest and their possible significance. Univ. Calif. Publ.
Agr. 3: 37-54. pl. 12. 29S 1917.
Hopkins, L. S. A new species of fern (Polystichum Jenningsi). Ann.
Carnegie Mus, 11: 362, 363. pl. 37. 5 N 1917.
Jeffrey, E.C. The anatomy of plants i-x + 1-478. f. 1-306. Chicago.
O 1917.
Jennings, 0. E. Pierygophyllum acuminatum at Ohio Pyle, Pennsyl-
vania. Bryologist 20: 100. N 1917.
Korstaim, C. F. The indicator significance of native vegetation in the
determination of forest sites. Plant World 20: 267-287. S 1917.
Latham, R. Habitat of Cephalosia Francisci on Long Island, N. Y.
Bryologist 20: 63, 64. 15 Au 1917.
Levy, D. J. Some experiments on the germination of moss spores on
agar. Bryologist 20: 62, 63. 15 Au 1917.
Lloyd, C. G. Mycological notes 50: 701-716. f. 1049-1074 + por-
trait. O 1917.
Léveillé, H. Catalogue des plantes du Yun-Nan, avec renvoi aux
diagnoses originales observations et descriptions d’espéces nou-
velles. 1-299. f. 1-68. 1917.
Contains reference to, and figure of, an American plant, Epilobium Arechavaletae
Lownes, A. E. Further notes on the orchids of the Asquam Region,
Rhodora 19: 235, 236. 21 N 1917.
Luckan, L. Ecological morphology of Abutilon Theophrasti. Kansas
Univ. Sci. Bull. 20: 219-228. pl. 1-3. Ja 1917.
MacCaughey, V. The guavas of the Hawaiian Islands. Bull. Torrey
Club 44: 513-524. 20 N 1917.
MacCaughey, V. The phytogeography of Manoa Valley, Hawaiian
Islands. Am. Jour. Bot. 4: 561-603. f. I-14. 12 D 1917.
MacCaughey, V. A survey of the Hawaiian land flora. Bot. Gaz. 64:
89-114. f. 1-5. 15 Au 1917.
MacCaughey, V. Vegetation of Hawaiian lava flow. Bot. Gaz. 64:
386-420. f. 1-22. 16 N 1917.
Macdonald, G. B. Evergreen trees for lowa. Yearbook lowa Dept.
Agr. 17: 437-464. pl. 1, 2 +f. 1-18. 1917.
Also published in Iowa Agr. Exp. Sta. Bull. 170: 1-16. Mr 1917.
MacDougal, D. T., & Spoehr,H. A. The measurement of light in some
of its more important physiological aspects. Science II. 45: 616-
618. 15 Je 1917.
, L. M. The crown canker disease of rose. Phytopathology
7: 408-417. f. 3-3. 3 D-19%7.
Cylin
ium scoparium.
126 INDEX TO AMERICAN BOTANICAL LITERATURE
Mortinez, M. Apuntes para una monografia del pochote o arbol del
Aegodon. Bol. Direc. Estud. Biol. Mexico 2: 150-158. f. I-4.
1917.
Murphy, L. S. The red spruce: its growth and management. U. S.
Dept. Agr. Bull. 544: 1-100. pl. 1-7 + f. 1-3. 31 O 191
Nanz, R.S. Note on Buxbaumia indusiata Bridel. Bryologist 20: 64.
15 Au 1917.
Nieuwland, J. A. Teratological notes. Am. Mid. Nat. 5: 156.
N 1017
Olive, E. W. Nea colors. Brooklyn Bot. Gard. Leaflets 5: [1-3].
17 O 1917.
, Olive, E. W. A trip to Texas to investigate cotton rust. Brooklyn
Bot. Gard. Rec. 6: 154-158. O 1917.
O’Neil, G. H. Messages of flowers or their floral code and dictionary.
1-142. frontispiece. Brooklyn. 1917.
“Embracing mythological stories and some floral facts.’’
Osterhout, G. E. A new Mertensia. Torreya 17: 175, 176. 31 O
1917.
Mertensia media sp. nov. :
Osterhout, W. J. V. The réle of the nucleus in oxidation. Science II.
46: 367-369. 12 O 1917.
Overholts, L.O. The structure of Polyporus glomeratus Peck. Torreya
17: 202-206. pl. 1. 30 N 1917.
Overholts, L.R. An undescribed timber decay of pitch pine. Myco-
logia 9: 261-270. pl. 12, 13. 24S 1917.
A rot caused by Polyporus amor phus.
Parish, S. B. An enumeration of the Pteridophytes and Spermato-
phytes of the San Bernardino Mountains, California. Plant World
20: 163-178. f. I-3. Je 1917; 208-223.. Jl 1917; 245-259. Au
1917.
Pearson, C. H. The almacigo or gumbo-limbo. Cuba Rev. 14°: I0-
14. F 1916. [Illust.]
Piper, C. V. Notes on Canavalia with the descriptions of new species.
Proc. Biol. Soc. Washington 30: 175-178. 23 O 1917.
Canavalia campylocarpa and C. luzonica, spp. nov.
Posey, G. B., Gravatt, G. F., & Colley, R. H. Uredinia of Cronartium
ribicola on Ribes stems, Science II. 46: 314, 315. 28S 1917
Pratt, D. J. An anatomical study of Cycloloma atriblicifolium.
Kansas Univ. Sci. Bull. 20: 87-120. pl. 1-19. Ja 1917.
Rands, R. D. Aliernaria on Datura and potato. Phytopathology 7:
327-338. f. 1-4. 30 1917.
Record, S. J. Ray tracheids in Quercus alba. Bot. Gaz. 64: 437. f. 1.
16 N 1917.
INDEX TO AMERICAN BOTANICAL LITERATURE 127
Reddick, D. Effect of soil temperature on the growth of bean plants
and on their susceptibility to a root parasite. Am. Jour. Bot. 4:
513-519. 24 N 1917.
Rehder, A. The genus Fraxinus in New Mexico and Arizona. Proc.
Am. Acad. Arts & Sci. 53: 199-212. O a
Includes Fraxinus Standleyi and F. Lowellii, spp. n
Reynolds, E. S. Internal telia of rusts. kes II. 46: 140, 141.
1o Au 1917.
Riddle, L. W. Pyrenothrix nigra gen. et sp. nov. Bot. Gaz. 64: 513-
15. f. Ig. 18 Dagre:
Riddle, L. W. The genus Parmeliopsis of Nylander. Bryologist 20:
69-76, pl. 20 +f. 1, 2. 13 01917.
Ridgway, C. S. Method of photographing culture plates. Phyto-
pathology 7: 388-391. f. 1. 30 1917
Rietz, H. L., & Smith, L. H. A statistical study of some indirect
effects of certain selections in breeding Indian corn. Jour. Agr.
Research 11: 105-146. f. 1-24. 22 O 1917.
Ritter, N. Histology of Astragalus mollissimus. Kansas Univ. Sci.
Bull. 20: 197-208. pl. 1-4. Ja 1917.
Rock, J. F. Hawaiian trees—a criticism. Bull. Torrey Club 44: 545,
> SG. a7 1) 1017,
Rock, J. F. The Ohia Lehua trees of Hawaii. Hawaii Board Agr. &
Forest. Bot. Bull. 4: 1-76. pl. 1-31. Au 1917.
Rock, J. F. Revision of the Hawaiian species of the genus Cyrtandra,
section Cylindrocalyces Hillebr. Am. Jour. Bot. 4: 604-623.
Re oe Pigs FE Gd
Cyrtandra Waianuensis sp. nov. and several new varieties are described.
Rogers, J. M., & Earle, F.S. A simple and effective method of pro-
tecting citrus fruits against stem-end rot. Phytopathology 7: 361—
367, 3 UF igi.
' Rolfe, R. A. Ondontoglossum chiriquense. Curt. Bot. Mag. IV. 13:
pl, 6725. 3S 4917.
A plant from Central Ameri
Rolfe, R. A. Odsatisbission platycheilum. Curt. Bot. Mag. IV. 13:
Pl. 8778. S$ 1917.
A plant from Guatemala.
Rolfe, R. A. New orchids: Decade XLV. Kew Bull. Misc. Inf.
1917: 80-84. Jl 1917.
Includes Pleurothallis mennlsege Epidendrum tricarinatum, Mazxillaria
Shepheardii, and Dichaea ciliolata from America.
Rorer, J. B. Algal renee eF cacao. Proc. Agr. Soc. Trinidad &
Tobago 17: 345-348. -S 1917.
128 INDEX TO AMERICAN BOTANICAL LITERATURE
Rose, J. N. Pachyphytum bracteosum. Addisonia 2: 53. pl. 67. 29
S 1917.
Russel, P.G. Uses of cacti. Am. Bot. 23: 90-97. Au 1917.
Sampson, A. W. Important range plants; their life history and
forage value. U.S. Dept. Agr. Bull. 545: 1-63. pl. 1-56. 8 O
1917.
Sawyer, M. L. Pollen tubes and spermatogenesis in Iris. Bot. Gaz.
64: 159-164. f. 1-18. 15 Au 1917. :
Sawyer, W. H., Jr.. The development of Cortinarius pholideus. Am.
Jour. Bot. 4: 520-532. pl. 28, 29. 24 N 1917.
Seaver, F. J. Sclerotinia and Botrytes. Torreya 17: 163, 164. 20
1917.
Also published in Science II. 46: 163. 17 Au 1017.
Shamel, A.D. Origin of the striped cane. Jour. Heredity 8: 471, 472.
f. 13. O 1917. |
Shantz, H. L., & Piemeisel, R. L. Fungus fairy rings of eastern Col-
orado and their effect on vegetation. Jour. Agr. Research 11: 19I-
246. pl. 10-30 + f. I-15. 29 O 1917.
Shear, C. L. Endrot of cranberries. Jour. Agr. Research 11: 35-42.
pl. A + f. 1-3. -8 O i917:
Fusicoccum putrefaciens sp. nov. is described,
Shear, C. L., & Stevens, N. E. The botanical work of Ezra Michener.
Bull. Torrey Club 44: 547-558. 27 D 1917.
Shear, C. L., & Stevens, N. E. Studies of the Schweinitz collections of
fungi-II. Distribution and previous studies of authentic speci-
mens. Mycologia 9: 333-344. 24 N 1917.
Sherwood, N. P. A study of several strains of pleomorphic Strepto-
cocci. Kansas Univ. Sci. Bull. 20: 245-257. Ja 1917.
Shufeldt, R. W. Marsh land and other aquatic plants. Am. Forest.
23: 611-618. O 1917. [Illust.]
Shufeldt, R.W. Necessity of greater accuracy in describing American
trees. Nature Study Rev. 13: 288-294. f. 1,3. O 1917.
Shufeldt, R. W. Queen Ann’s lace: the papaw tree, and self-heal.
Am. Forest. 23: 543-548. f. 1-6 + f. 98-124. S$ 1917,
Sinnott, E. W. A botanical criterion of the antiquity of the Angio-
sperms. Jour. Geol. 24: 777-782. D 1916.
Sinnott, E. W. The ‘‘age and area” hypothesis and the problem of
endemism. Am, Bot! 31: 209-216. Ap 1917.
Skan, S. A. Pyrola bracteata. Curt. Bot. Mag. IV. 13: pl. 8710 B.
Je 1917. :
A North American plant.
INDEX TO AMERICAN BOTANICAL LITERATURE 129
Skan, S.A. Pyrola uliginosa. Curt. Bot. Mag. IV. 13: pl. 8770 A.
Je 1917.
A plant from North America
Small, J. K. Aesculus parviflora. Addisonia 2: 45, 46. pl. 63. 29S
1917.
Small, J. K. The tree cacti of the Florida Keys. Jour. N. Y. Bot.
Gard. 18: 199-203. pl. 206. S$ 1917.
Includes description of Cephalocereus Deeringii sp. nov.
Smith, C.P. Studies in the genus Lupinus—I. A new species of the
subgenus Platycarpos. Bull. Torrey Club 44: 405, 406. 10 Au
1917.
Lupinus subvexus sp
Soth, B. H. Plant life o on the peaks. Am. Bot. 23: 77-85. Au 1917.
[Illust.] ‘
[Standley, P. C.] Botanical explorations in Florida and New Mexico.
Smithsonian Misc. Coll. 66'7: 53-59. f. 54-6o. IQI7.
Stapf, O. Pinus tuberculata. Curt. Bot. Mag. IV. 13; pl. 8717. S
1917.
A plant from western North America.
Stevens, F.L. Spegazzinian Meliola types. Bot. Gaz. 64: 421-425.
pl. 24-26. 16N 1917.
Stevens, O. A. Plants of Manhattan and Blue Rapids, Kansas, with
dates of fowecing—III. Am. Mid. Nat. 5: ii3-1290- -S ror.
Stevenson, J. A. Wood rot of citrus trees. Porto Rico Dept. Agr.
and Lab. Exp. Sta. Rio Piedras Circ. 10: I-10. 1917.
Stevenson, J. A. An epiphytotic of cane disease in Porto Rico. Phy-
topathology 7: 418-425. f. 7,2. 3 D 1917.
Stewart, V.B. The perennation of Cronartium ribicola Fisch. on cur-
rant. Phytopathology 7: 448, 449. 3 D 1917.
Stewart, V. B. A twig and leaf disease of Kerria japonica. Phyto-
pathology 7: 399-407. f. 1-6. 3 D 1917.
Coccomyces Kerriae sp. nov.
Stout, A.B. Notes regarding variability of the rose mallows. Torreya
27: 142-148. 14 Au 1917.
Stout, A. B. Observations on tulips. Jour. Hort. Soc. New York 2:
201~206. pl. 37, 38. At 1917.
Sturgis, W. C. Notes on new or rare Myxomycetes. Mycologia 9:
323-332. DF t4, 16.. 24 N 1917.
Includes Physarum _melanospermum P. lilacinum, Didymium fulvum and En-
teridium minutum, spp.
Swingle, L. A boighi- bole lemon. Jour. Heredity 8: 559. f. 8.
D 1917.
130 INDEX TO AMERICAN BOTANICAL LITERATURE
Swingle,W. T. The early European history and the botanical name of
the Tree of Heaven, Ailanthus altissima. Jour. Washington Acad.
Sci. 6: 490-498. 19 Au 1916.
Swingle, W. T. Le fruit mfir et les jeunes semis de /’Aeglopsis Cheva-
liert. Bull. Soc. Bot. France 60: 406-409. f. A. 1913.
Swingle, W. T. Merotypes as a means of multiplying botanical types.
Jour. Washington Acad. Sci. 2: 220-222. 4 My 1912.
Swingle, W. T. Microcitrus, a new genus of Australian citrous fruits.
Jour. Washington Acad. Sci. 5: 569-578. f. 1-4. 40 1915.
Swingle, W. T. - The name of the wood-apple, Feronia Limonia. Jour.
Washington Acad. Sci. 4: 325-329. 19 Je 1914.
Swingle, W. T. A new genus, Fortunella, comprising four species of
kumquat oranges. Jour. Washington Acad. Sci. 5: 165-176. f. 1-5.
4 Mr 1015.
' Swingle, W. T. Pamburus, a new genus related to Citrus, from India.
Jour. Washington Acad. Sci. 6: 335-338. 4 Je 1916.
Swingle, W. T. Pleiospermium, a new genus related to Citrus, from
India, Ceylon and Java. Jour. Washington Acad. Sci. 6: 426-431.
19 Jl 1916.
Swingle, W. T. Severinia buxifolia, a citrus relative, native to south-
ern China. Jour. Washington Acad. Sci. 6: 651-657. f. 1,2. 19 N
Tanaka, T. New Japanese Fungi. Notes and_ translations—II.
Mycological 9: 249-253. Jl ‘1917;—III. Mycologia 9: 365-368.
Taubenhaus, J. J. On a sudden outbreak of cotton rust in Texas.
Science IT. 46: 267-269. 14S 1917.
Taylor, M. W. Preliminary report on the vertical distribution of
Fusarium in soil. Phytopathology 7: 374-378. 3 0 1917.
[Taylor, N.] The Semi-centennial of the Club. Torreya 17: 183-186.
Taylor, N. Informal and wild gardening. Jour. Internat. Gard. Club
I: 91-104. Auso917. [Illust.]
Taylor, N. Plants of the Catskill aqueduct. Brooklyn Bot. Gard.
Leaflets 5: [3-8]. 100 1917.
Terao, H. On reversible transformability of allelomorphs. Am. Nat.
51: 690-698. N 1917.
Thériot, I. Contribution a la flore bryologique du Chili. Rev. Chilena
Hist. re 20: aed pl. 1-7. 1917.
New i ortula (2), Webera - Bryum (2), Pogonatum (1), Porotrichum
(2), and Costesia pei gen. et sp. no
INDEX TO AMERICAN BOTANICAL LITERATURE 131
Thom, C. C., & Holtz, H. F.. Factors influencing the water require-
ments of plants. Washington Agr. Exp. Sta. Soil Physics Bull.
146: 1-64. f. 1-18. Je 1917.
Tisdale, W. H. Relation of temperature to the growth and infecting
power of Fusarium Lini. Phytopathology 7: 356-360. pl. rz +
Fs lad O 1017:
Traver, J. The American larch or tamarck, a tree of the swamps.
Nature Study Rev. 13: 341-345. N 1917. [Illust.]
Trotter, A. Osservazionie ricerche istologiche sopra alcune morfosi
vegetali determinate da funghi. Marcellia 15: 58-111. pl. 1-3 +
f. toe 41 Au sot.
Trumbull, H. L., & Hotson, J. W. The effect of Roentgen and ultra-
violet rays upon fungi. Phytopathology 7: 426-431. f. 1, 2.
3 2) 1677.
Van Eseltine, G. P. Selaginella fusiformis, a new species in the S.
rupestris group. Proc. Biol. Soc. Washington 30: 161, 162. 10 O
1917.
Vestal, A.G. Foothills vegetation in the Colorado front range. Bot.
Gaz. 64: 353-385. f. 1-8. 16 N 1917.
Victorin, M. Découverte du lycopode petit-cypres dans les Lauren-
tides. Le Naturaliste Canadien 39: 166-170. My 1913.
Victorin, M. Notes botaniques. Le Naturaliste Canadien 40: 78, 79.
N 1913.
Victorin, M. Notes sur deux cas d’hybridisme natural. Le Natur-
aliste Canadien 39: 177-189. f. 1-4. Je 1913.
Victorin, M. Plusieurs ‘‘Cyperacies”’ nouvelles pour la flore de Québec.
Le Naturaliste Canadien 39: 117-123. f.1, 2. F 1913.
Waterman, W. G. Plant ecology and its relation to agriculture.
Science II. 46: 224-228. 7S 1917.
Weatherby, C. A. Color forms of Impatiens biflora. Rhodora 19:
195-418... 2 JL 1917.
Weatherby, C. A. Concerning Polystichum acrostichoides forma lan-
ceolatum. Am. Fern Jour. 7:90, 91. 2 Jl 1917.
Weatherwax, P. The development of -the spikelets of Zea Mays.
Bull. Torrey Club 44: 483-496. pl. 23 +f. I-33. 10 1917.
Weimer, J. L. Three cedar rust fungi, their life histories and the
diseases they produce. Cornell Agr. Exp. Sta. Bull. 390: 507-549.
jf. 130-157. My 1917.
Weir, J. R. A needle blight of Douglas fir. Jour. Agr. Research 1o:
99-103. pl. 12 +f. 1-3. 9 Jl 1917.
132 INDEX TO AMERICAN BOTANICAL LITERATURE
Weir, J.R. Notes on wood-destroying fungi which grow on both coni-
ferous and deciduous trees—II. Phytopathology 7: 379, 380.
O 1917.
Weir, J. R., & Hubert, E.E. Cvonartium cerebrum on Pinus resinosa.
Phytopathology 7: 450,451. 3 D 1917.
Weir, J. R., & Hubert, E. E. Observations on forest tree rusts. Am,
Jour. Bot. 4: 327-335. f. 1,2. Je 1917.
Uredinopsis pleridis on Abies grandis and Pteridium aquilinum pubescens.
Wernham, H. F. Tropical American Rubiaceae—VIII. Jour.. Bot.
85:169-177. Jl 1917:—IX. Jour. Bot. 55: 251-254. $1917.
Includes Neobertiera gracilis, Blandibractea brasiliensis, gen. et spp. nov.; also
six new species in Sipanea, three in Cephalanthus, and seven in Psychotris.
Wester, P. J. Additional observations on the citrus fruits of the Philip-
pines. Philip. Agr. Rev. 10: 104-115. pl. 2-8. 1917.
Wester, P. J. New or noteworthy tropical fruits in the Philippines.
Philippine Agr. Rev. 10: 8-23. pl. 2-9. 1917.
Wester, P. J. Plant names. Philippine Agr. Rev. 10: 1-9. 1917
10: 55-63. 1917.
Weston, W. H. Observations on an Achlya lacking sexual reproduction.
Am. Jour. Bot. 4: 354-366. Je 1917.
White, O. E. Inheritance of endosperm color in maize. Am. Jour.
White, O. E. Inheritance studies in Pisum. IV. Interrelation of
the genetic fectors of Pisum. Jour. Agr. Research 11: 167-190
22 O 1917
Williamson, J. A golden variety of rye. Jour. Heredity 8: 568.
Williams, R. S. Philippine mosses. Bull. N. Y. Bot. Gard. 8: 331-
378. pl. 171-174. 23 Jl 1914.
Wilson, J. K. Physiological studies of Bacillus radicicola of soy-bean
(Soja Max Piper) and of factors influencing nodule production.
Cornell Agr. Exp, Sta. Bull. 386: 369-413. f. 80-94. J 1917.
Wilson, P. The vegetation of Vieques. Bull. N. Y. Bot. Gard. 8: 379-
426. 9 Jl 1917.
Wilson, P. Micrampelis lobata. Addisonia 2: 47, 48. pl. 64. 29 5
1917.
Wilson, S.F. A giant cactus. Am. Forest. 23:427. Jl1917. [Illust.]
Winslow, C. E. A., Broadhurst, J., and others. The families and genera
of the bacteria. Jour. Bact. 2: 505-556. S 1917.
Winslow, E. J. The male fern in Vermont. Am. Fern Jour. 7: 87-90.
ji 1917.
BULL. TORREY CLU
RR CLUB VOLUME 45, PLATE 4
STEIL: APOGAMY IN FERNS
VOLUME 45, PLATE 5
IB
CLu
TORREY
BULL.
egae
SOF
eo
cA iby,
ie Oth
{
"re
sg th,
Pnee
Cie t
a
So ecte
Seep eS
a
AP gore 58
We f
»
yak
c
et
&
fool
js
G
shale
Y cies
rs
moth
4
0
APOGAMY IN FERN
STEIL:
Voi. 45 No. 4
BULLETIN
OF THE
TORREY BOTANICAL CLUB
New species of Hawaiian plants
JosEpH F. Rock
(WITH PLATE 6)
1, Cyanea Giffardii sp. nov.
A tree 5-10 m. tall with a single trunk 15 cm. in diameter near
the base, gradually tapering towards the apex, bark smooth,
grayish green, with scattered leaf-scars, woody zone thin near the
apex of the trunk, the medullary cavity septate by chartaceous
diaphragms; leaves obovate-oblong, about 50 cm. long and 12-15
cm. wide, broadest portion in the upper third, margins strongly
undulate and minutely denticulate, appearing sinuate, due to
strong undulation, subentire with exception of the base, which is
unevenly lobed, acuminate at the apex, mucronate, gradually
tapering at the base into a stout petiole 3.5-6.5 cn. long, thin,
subchartaceous, dark green and shining above, pale and dull
~ underneath, midrib stout and prominent as are the lateral veins,
the latter arcuate and united with other arcuate veins near the
margin, the whole surface of the leaf covered with a pellucid reticu-
late net work, glabrous above, pubescent underneath, especially
on the midrib and veins; peduncles axillary, drooping, stout, terete,
glabrous, 10-16 cm. long or slightly longer, about 1 cm. thick,
distantly bracteate the entire length, the upper bracts linear,
15-30 mm. long, 3-5 mm. wide, rounded and mucronate at the
apex, bracts less distant towards the apex and also smaller;
flowers on pedicels 2-3 mm. long with three small, acute bractlets
at the base and dorsal side of the flower; calyx glabrous, dark
purplish black, tube obconical, usually seven-ribbed, with two
[The BULLETIN for March (45: 93-132, pls. 4, 5) was issued April 1, 1918.]
134 Rock: NEw HAWAIIAN PLANTS
tubercles at the base, 18 mm. long, 15 mm. wide; calycine lobes
broadly triangular, acute, 5 mm. each way, with broad sinuses
intervening; corolla strongly arcuate, 7-8 cm. long, glossy, glab-
rous, dark purplish outside, slit at the back three fourths its
length only when fully mature, the five lobes of the corolla en-
tirely connate; staminal column protruding, perfectly glabrous,
pale, anthers glaucous, glabrous, the two lower only penicillate;
style black, the stigmatic hairs encircling the stigma, the latter
yellowish tinged with purple; flowers usually ten on a peduncle,
crowded at the apex; fruits globose, nearly 25 mm. each way, dark
purplish black, locules small, each containing from six to twelve
rather large whitish seeds; milky juice of the plant yellowish.
[PLATE 6.]
Hawall: in the forest on the windward slope of Mauna Loa,
near Glenwood at 22 miles, along the Homestead Road at an ele-
vation of 2,200 feet, August 27, 1917,W. M. Giffard 12802, (flower
buds) in the herbarium of the College of Hawaii; along the Vol-
cano Road at 23 miles, in wet forest, September 1, 1917, Rock &
Holm 128026, (flowering and fruiting specimens), TYPE, in the
herbarium of the College of Hawaii.
This very remarkable plant, which is closely related to Cyanea
superba (Cham.) Gray, differs from it in the pubescent, deeply
undulate leaves, which are lobed at the base instead of being entire;
in the glabrous, much shorter peduncle, and in the glabrous deep
purplish black flowers and fruits. ‘Cyanea superba occurs on the
island of Oahu in the gulches of Makaleha and Mt. Kaala, while
Cyanea Giffardii occurs on the southernmost island of the group.
It is also much statelier than Cyanea superba, as it reaches a height
of thirty feet, with a single crown of leaves at the apex. It grows
in company with Antidesma platyphyllum Mann, Labordia,
Cyrtandra, Straussia hawatiensis Gray, Strongylodon lucidum
Seem., Clermontia parviflora Gaud., Cibotium Menziesii Hook. and
C. Chamissoi Kaulf., the last two being the common tree ferns of
the region.
_ When the species was first discovered by Mr. W. M. Giffard,
in whose honor it is named, only two plants were observed, one with
large flower buds and the other without flowers, the latter plant
divided into three branches at the apex, on account of an injury.
The mature type specimens, with flowers and fruit, were col-
lected on the road to Glenwood and the Volcano, at an elevation
Rock: New HAWAIIAN PLANTS 135
of 2,400 feet. The largest plant seen was thirty feet in height.
The plants are difficult to see in the forest as the trunk, which is
usually covered partly with moss, does not branch and the crown
of leaves is hidden amongst the foliage of other trees.
Cyanea Giffardii may be Hillebrand’s 6 var. of his Cyanea
arborea from the woods of Hilo, Hawaii. The writer is well ac-
quainted with Cyanea arborea and can only state that the new
species is exceedingly different from the latter, and that it comes
much closer to Cyanea superba. Cyanea arborea has a much
larger and denser crown of leaves, which are sessile, and linear-
oblong; the peduncles are much longer and the flowers are very
thin, narrow and slender, suberect and whitish to gray.
2. Cyanea rollandioides sp. nov.
Plant I-1.5 m. high, stem simple, fleshy towards the apex,
woody towards the base, stem muricate to spinose in the upper
portion; leaves obovate-oblong, acute, fleshy when fresh, papery
when dry, dark green above, paler underneath, but with dark
purple midrib and veins and a prominent dark purple reticulate
network, puberulous or glabrous on both surfaces, but more or less
covered with spines on both sides, those of the upper surface yel-
low, those of the lower surface deep purple, margins eroso-dentate
to irregularly notched, and somewhat uneven-sided at the base,
30-50 cm. long, 8-15 cm. wide in the widest portion, which is in
the upper third, on fleshy stout spinose or muricate petioles, 8—1 5
cm. long; racemes glabrous, peduncle 3-6 cm. long, naked three
fourths its lower length, but distantly covered with scars of fallen
flowers, bearing in its upper fourth about fifteen flowers; bracts
subulate, 3 mm. long, supporting each pedicel, the latter filiform,
10-25 mm. long, bibracteolate, the bracteoles alternate, one at
about the middle of the pedicel the other near the apex, 0.25 mm.
long; calyx tube turbinate to obovate-oblong, 7-10 mm. high, the
linear calycine lobes as long as the tube; corolla deep purplish red
or purple to pale yellowish white with dark purplish streaks,
moderately arcuate, broadest at the middle, 5-8 mm., about 4-5
cm. long, thin and glabrous, dorsal slit very shallow, extending
only one fourth of the length of the tube or a little beyond the two
upper linear subulate lobes, the three lower lobes a little shorter;
staminal column glabrous, as are the pale greenish anthers, the
lower ones only penicillate; fruit unknown.
Hawatt: Forest of Puna in dense woods along the Kalapana
Road, not far from Pahoa, September 3, 1917, Rock & Newell!
136 Rock: NEw HAWAIIAN PLANTS
12831 (flowering specimens), Type, in the herbarium of the Col-
lege of Hawaii.
This rather variable species is remarkably like a Rollandia.
The plant varies considerably in the spinosity of the leaves,
petioles and stems; some of the specimens almost approach Cyanea
noli-me-tangere Rock in spinosity, while others are only muricate.
The color of flowers is also variable, ranging from pale yellowish
white with deep purplish streaks to entire dark purple. It is diffi-
cult to state to what known species the plant is closest related; in
habit it approaches Rollandia lanceolata Gaud., but actually seems
to come close to Cyanea noli-me-tangere; from the latter it differs in
the long-petioled leaves, which are much larger, and in the longer
naked peduncles which are not spinose. The plant as a whole is
much larger in every way, the flowers are purple as are the veins
and midrib of the leaves; the whole inflorescence is glabrous. C.
noli-me-tangere is very loosely foliate, while C. rollandioides is
simple-stemmed and has at its apex a dense crown of leaves, this
feature bringing it under the section Palmaeformes.
The plants grow in the wet forests of Puna on Hawaii, a little-
explored district and one of the most primitive regions on the
island of Hawaii. A stalwart Hawaiian gave the writer the native
name Aku-aku for the species in question, of which he said that
the leaves were cooked with meat and eaten like cabbage. The
name Aku alone is applied to Cyanea tritomantha Gray, to which our
plant has no resemblance.’ The species was collected in company
with Brother Matthias Newell, of Hilo, an ardent naturalist.
3. Rollandia angustifolia (Hillbrd.) sp. nov.
Rollandia longiflora 8 var. angustifolia Hillbrd. Fl. Hawaiian Isl.
246. 1888.
Stem smooth, 1-1.5 m. high, leaves linear-lanceolate, 18-35
cm. long, 2.5-3.75 cm. wide, thick, fleshy, dark green, glossy above,
pale whitish underneath, with dark purplish midrib and veins,
acuminate, mucronate at the apex, gradually narrowing at the
base into a fleshy petiole 2-4.5 cm. long, glabrous on both sides;
racemes slender, 3-4 cm. long, four- to five-flowered ; pedicels thin,
about 14 mm. long, bracteate at the base, bi-bracteolate about the
middle; calyx turbinate, 1 cm. long, the apex truncate-dentate, or
oftener lobed, the calycine lobes acute, of irregular length, usually
Rock: NEw HAWAIIAN PLANTS 137
3 mm. long, with a median nerve; corolla deep purplish red, 7-8.5
cm. long, I cm. wide, the lobes 1.5 cm., the dorsal slit extending
one fifth the length of the tube; staminal column glabrous adherent
up to about the middle of the corolla, the anthers glabrous, the
lower only bearded; fruits globose, crowned by the tubular limb
of the calyx, which disappears at the maturity of the fruit.
Oanu: Mt. Konahuanui trail, Palolo Valley, Mt. Olympus,
and Manoa Valley, Kalihi Valley, January, 1870, W. F. Hillebrand,
without number, in Berlin Herbarium; June 14, 1908, H. L. Lyon
8816 in the herbarium of the College of Hawaii; September, 1912
and 1914, J. F. Rock 10250a, 10250b, in the herbarium of the
College of Hawaii.
The plant in question is certainly worthy of specific rank.
Young plants which the writer observed were of the same habit as
mature ones, both having linear-lanceolate leaves, while the true
Rollandia longiflora Wawra has sinuate leaves when young and
also when in a mature state. R. angustifolia differs mainly in the
linear, entire, minutely denticulate leaves, which give the plant
an entirely different appearance from that of R. longiflora.
4. Lobelia oahuensis sp. nov.
Plant rather stout, stem short and thick, solid and not hollow;
rosette of leaves very dense and about 1 m., in diameter; leaves
densely packed around the apex of the stem, linear-oblong, acumi-
nate at both ends, merging at the base into a winged fleshy petiole
about 2.5 cm. in length, 50 cm. long, 4.5-5 cm. wide, thick, coria-
ceous, dark green, glabrous above and covered with a strongly
impressed, very close, reticular net work, young leaves densely
hirsute underneath, especially along the very prominent pro-
jecting midrib and veins, of a dirty grey or fawn color on the older
leaves, the margins revolute, denticulate with thick callous teeth;
flowers not seen, a single dead terminal flower stalk was seen on one
of the plants, which was about 1 m. long.
Oauu: at the very top of the main crest of the island, over-
looking the cliffs of Waimanalo at an elevation approaching 3,000
feet, September 14, 1917, J. F. Rock 12836, Type, in the herbarium
pS th
of the Collegeof Hawaii. Several pl g g tog ;
the lower ones of which could not be reached owing to the vertical
cliffs on which they grew, immediately below the knife-edge crest
of the backbone of the island of Oahu. The plant forms a large
138 Rock: NEW HAWAIIAN PLANTS
rosette with the leaves densely packed at the apex in an almost
horizontal position, that is, at right angles to the stem. It grows
in company with Trematolobelia macrostachys (Hook. & Arn.),
Zahlbr., Dubautia laxa Hook. & Arn., Metrosideros rugosa Gray, and
other species. Notwithstanding that the flowers of this new
species are unknown, the plant is so distinct from all of our other
lobelias that it can well be described at present.
The plant is evidently related to Lobelia hypoleuca Hillbrd.,
from which it differs in the solid stem, thick, coriaceous, closely
reticulate leaves, not silvery underneath but hirsute, being covered
with fawn-colored or dirty gray hair. In Lobelia hypoleuca, which
is a branching species, the leaves are few and more or less scattered,
thin and chartaceous; it does not ascend to such high elevations,
but remains more at the lower levels, from 1,000 to 1,500 feet, in
very sheltered situations, especially deep ravines.
5. Straussia glomerata sp. nov.
A medium-sized tree, 10 m. or more in height, trunk straight,
crown round and of rather small dimension; stipules cup-shaped,
not triangular, sheathing and of even height, 3 mm. high, soon
deciduous; leaves obovate, coriaceous, glossy and smooth above,
dull and pubescent beneath, especially along the midrib and veins,
pubescent glands present in the axils of veins and midrib, 5-10
cm. long, 4.5-6.5 cm. wide, on petioles 15 mm. long, rounded at
the apex, cuneate at the base; panicle erect, not drooping save
when in full fruit, 2-4 cm. long, distinctly angular, hirsute with
dirty yellowish hair, whorls two or three, of very short rays 2-6
mm. long; flowers sessile; calyx minute, 0.5-0.75 mm. long, densely
hirsute, truncate; corolla greenish white, minute, tube 0.5 mm.
long, lobes 1.5 mm. long, glabrous, stamens minute, filaments 0.2 5
mm. long; the convex disk densely hirsute; fruits numerous, densely
agglomerate, forming a densely packed globose head, hiding all
whorls and rays, fruits yellow, pubescent to hirsute, obovate, 9
mm. long, 6-6.5 mm. wide, crowned by a rounded, dome-shaped,
projecting disk.
Hawai: North Kona, in the forest of Waihou and on the lava
flows of Puuwaawaa, at an elevation of 3,700 feet, August 24,
1917, J. F. Rock 12829 (flowering and fruiting specimens), Tyre,
in the herbarium of the College of Hawaii. .
This interesting species is very distinct from the other species
Rock: NEw HawallAN PLANTS 139
in the genus, and is remarkable for the minute flowers, hirsute
panicles and fruits which are densely glomerate, forming a densely
packed, globose head, nearly as long as broad. It is related to
Straussia hawatiensis which differs however in the loose and open-
whorled drooping, glabrous panicle.
COLLEGE OF Haw
HONOLULU, kes ene oF Hawalrr
Explanation of plate 6
Cyanea Giffardii Rock. see pyeee eS a type specimen, showing the crown
of a flowering tree, twenty-four feet in hei
New species of Uredineae—X *
JOSEPH CHARLES ARTHUR
The preceding number in this series was issued in November,
1915. Since that time some collections of Uredinales have come
to hand, or have been awaiting study, which appear to be unde-
scribed. The discovery of additional spore forms for previously
named species admits of their transfer to more suitable genera,
thus necessitating new combinations. Both new names and new
combinations are required in order to make the material more
available in future studies. All the species are North American.
Although the localities range from Wyoming and Vermont to
Panama and the West Indies, yet the majority are in Mexico
and Central America, where the rust flora is abundant, but much
less known than northward.
Uromyces Atriplicis (Shear) comb. nov.
Aecidium Atriplicis Shear, Bull. Torrey Club 29: 453. 1902.
The aecia of this rust have been known for a number of years.
They are abundant in Colorado on Atriplex confertifolia (Torr.)
S. Wats., and were distributed in Griffiths, West Amer. Fungi
321 (type), Ellis & Ev., Fungi Columb. 1294, and Barth., North
Amer. Ured. gor. On August 29, 1911, telia were gathered by
Mr. E. Bethel at Delta, Colorado, from the same plants that had
yielded a large crop of aecia earlier in the season. This collection
shows the same plentiful sprinkling of telia over the under side of
the leaves as marks the presence of the aecia, a few sori in both
cases usually appearing on the upper side of the leaves. The col-
lection from Delta also includes aecia, both on the same leaves
* Reprints may be obtained by application to the Botanical Department, Purdue
University Agricultural Experiment Station, Lafayette, Ind., under whose auspices
the work was carried on.
New Species of Uredineae I-IX: Bull. Torrey Club (I) 28: 661-666. Igor.
(II) 29: 227-231. 1902; (III) 32: 1-8. 1904; (IV) 33: 27-32. 1906; (V) 33: 513-
522. 1906; (VI) 34: 583-592. 1907; (VII) 37: 569-580. 1910 (VIII) 38: 369-378;
ro11; (IX) 42: 585-593- 1915
141
142 ARTHUR: NEW SPECIES OF UREDINEAE
with the telia, and intermixed with them, and also separately.
The telia were also found at Pueblo, Colorado, by Mr. Bethel, and
on September 13, 1911, Dr. F. D. Kern and the writer, following
Mr. Bethel’s directions, visited the spot, but were only rewarded
with aecia. On August 13, 1912, Dr. Kern repeated the visit and
found plenty of telia. Both the position of the telia on the leaves,
and their succession to the aecia on the same plants, indicate their
genetic relationship. The telia may be described as follows:
III. Telia chiefly hypophyllous, scattered, round or irregular,
0.2-0.4 mm. across, early naked, pulverulent, cinnamon-brown, .
ruptured epidermis noticeable; teliospores broadly ellipsoid or
ovoid, 18-23 by 23-30 u, rounded at both ends, or somewhat nar-
rowed below; wall cinnamon-brown, 1.5-2 thick, 4-5» above,
including a low semi-hyaline umbo, smooth; pedicel colorless,
fragile.
The type collection, made by Shear & Bessey at Montrose,
Colorado, July, 1897, showing only aecia, was distributed as on
Atriplex Nuttallii, an error for A. confertifolia.
Uromyces fuscatus sp. nov.
I. Aecia amphigenous, in dense groups 3-5 mm. across;
peridia short-cylindric, the margin erect, coarsely and deeply
lacerate; peridial cells rhomboidal, 13-19 by 28-37 u, considerably
overlapping, the outer wall 7-9 u thick, finely striate transversely,
the inner wall 2.5-3.5 u thick, moderately verrucose; aeciospores
globoid, 16-22 » in diameter; wall colorless, 1-1 .5 thick, finely
and closely verrucose, often appearing smooth when wet.
II. Uredinia amphigenous, scattered, round, 0.3-0.8 mm.
across, elevated, early naked, puberulent, dark cinnamon-brown,
ruptured epidermis pi ; urediniosp globoid or broadly
ellipsoid, 18-24 by 22-29 yw; wall cinnamon-brown, 2-3 thick,
closely and finely verrucose, the pores 3 or 4, approximately
equatorial.
III. Telia amphigenous, scattered, round or irregular, 0.5—2 mm.
across, often confluent, early naked, prominent, dark chestnut-
brown, ruptured epidermis evident; teliospores broadly ellipsoid or
On Polygonum alpinum All., House Creek, Idaho, June 29,
ARTHUR: NEW SPECIES OF UREDINEAE 143
1912, Nelson & Macbride 1794 (type); near Gogorza, Summit
County, Utah, June 29, 1915, A. O. Garrett 2286. Among autoe-
cious species this appears to be most like Uromyces Acetosae
Schrét., a European species on Rumex, but the urediniospores are
verrucose and not echinulate and have a different number of pores,
while the teliospores are contrasted in the coarseness and arrange-
ment of the markings. There appears to be no correlated form
among the many species of Puccinia on either Polygonum or
Rumex. The rust covers the leaves heavily with the conspicuous
sori, inducing a more or less mottled, reddish appearance of the
tissues. The specimen from Utah was labelled Pentstemon pro-
cerus, but both host and rust appear identical with the one from
Idaho.
Uromyces Krameriae Long, sp. nov.
III. Telia amphigenous and caulicolous, soon opening by a
longitudinal slit, chestnut-brown, loosely and prominently pulvi-
nate, oblong, 0.1-2 mm. long, ruptured epidermis conspicuous;
teliospores broadly ellipsoid or globoid, 21-23 by 23-294; wall
cinnamon-brown, 2.5-3 » thick, thicker above, 6-9 », including a
broad paler umbo, smooth; pedicel hyaline, once to twice or more
the length of the spore.
On Krameria glandulosa Rose & Painter, Denton, Texas,
October 21, 1901, W. H. Long 1to7z. The host has usually
passed under the name of K. parvifolia, a species of Lower
California. This rust was received a long time since with
the name as given above. I do not find that the species has
yet had the name established, and so take this opportunity to
place it in use. It is one of the numerous finds made by Mr.
W. H. Long, while living in northern Texas, and affords one more
evidence of Mr. Long’s extended and efficient study of the rusts.
No urediniospores were detected, but the appearance of the rust
is not that of a short-cycle form; it is more likely to possess cupulate
aecia. Only the one collection is known.
Puccinia wyomensis sp. nov.
O and I. Pycnia and aecia unknown
II. Uredinia chiefly bet sca tose naied: biline to linear,
0.6-2 mm. in length, 0.2-0.4 mm. wide, tardily naked, finally
opening by a longitudinal slit, pulverulent, golden-brown, rup-
144 ARTHUR: NEW SPECIES OF UREDINEAE
tured epidermis conspicuous; urediniospores broadly ellipsoid or
- obovoid, 18-21 by 26-29 y; wall pale yellow or quite colorless,
1-1.5 uw thick, closely and rather inconspicuously echinulate, the
pores 6-8, scattered.
Ill. Telia amphigenous, scattered, oblong to linear, 0.2-3
mm. in length, compact, dark gray, long covered by the epidermis,
surrounded by a thin dark-brown stroma; teliospores oblong or
clavate, 15-21 by 50-60, truncate or somewhat rounded above,
narrowed below, slightly or not constricted at septum; wall cinna-
mon-brown, darker above, thin, I-1.5 uw, not or slightly thickened
above, 2—5 u, smooth; pedicel very short, tinted.
On Scirpus americanus Pers., Arcola, Wyoming, August 23,
1916, E. Bartholomew 6089. A subepidermal form of Scirpus
rust abundantly distinct from all others.
Puccinia Rosenii sp. nov.
Oand I. Pycnia and aecia unknown.
II. Uredinia hypophyllous, scattered or in series, oval or
linear, 0.5-2 mm. long, rather tardily naked, ruptured epidermis
conspicuous; urediniospores 30-35 u long, when seen with pores in ~
optical section oblong, 10-20 » wide, when seen with pores central
globoid or broadly ellipsoid, 28-35 » wide; wall cinnamon-brown,
1.5-2.5 » thick, moderately echinulate, the pores 2, superequatorial.
Ill. Telia few, disposed like the uredinia, tardily naked,
chocolate-brown; teliospores clavate, or elongated oblong, 16-23
by 43-67 p, rounded or truncate above, usually narrowed below,
slightly constricted at septum; wall chestnut-brown above, some-
what lighter below, 1-24 thick, 5-10» above, smooth; pedicel
colorless, nearly as long as spore.
On Schoenus nigricans L. (?), Daytona, Florida, December 27,
1915, R. A. Harper 5. This rust, found in scanty amount on a
somewhat uncertain host without inflorescence, is apparently
nearest like P. Eriophori Thiim. This species is named for H. R.
Rosen, who has made the careful study of this material on which
the diagnosis and conclusions are founded.
Puccinia Heliconiae (Diet.) comb. nov.
Uredo Heliconiae Diet. Hedwigia 36: 35. 1897.
The uredinial stage of this rust was described from a collection
made at Rio de Janeiro, Brazil, in December, 1891, on an unde-
ARTHUR: NEW SPECIES OF UREDINEAE 145
termined species of Heliconia (Bihai). In North America it was
collected in Martinique, West Indies, on Bihai borinquena Griggs,
August, 1913, and again by Whetzel and Olive in Porto Rico, West
Indies, on the same host, April, 1916, both collections showing
only uredinia. In a collection on Béihai latispatha (Benth.)
Griggs, made by E. Bethel, at Montelinio, Canal Zone, Panama,
March 4, 1913, an abundance of telia accompany the uredinia.
The characters of the telia are as follows:
III. Telia hypophyllous, scattered, soon naked, round, o. 3-0.
mm. across, cinnamon-brown, ruptured epidermis evident; telio-
spores elongated clavate, 15-19 by 60-724, rounded above, not
or slightly constricted at septum; wall pale cinnamon-brown, thin,
I w, much thickened above, 9-13 4, smooth, pedicel short, colorless.
: The urediniospores of this species appear to be quite variable.
Those of the West Indian collections are thin and dark, as com-
pared with most of those of the Panama collection, in which, how-
ever, all gradations occur. The sori of the present material are
much parasitized, and both uredinia and telia are consequently
unnaturally conspicuous.
Puccinia Viornae sp. nov.
O. Pycnia epiphyllous, crowded upon swollen areas 2.5-5 mm
in diameter, dark brown, noticeable, subepidermal, globoid,
98-110 uw in diameter, with ostiolar filaments
I. Aecia hypophyllous, crowded upon swollen areas opposite
the pycnia, cupulate, short, 0.2-0.5 mm. in diameter; peridium
recurved, the peridial cells rectangular, abutted, 19-23 by 22-24 u,
the outer wall 7-9 u thick, the inner wall 3-5» thick, finely ver-
rucose; aeciospores irregularly globoid or ellipsoid, 13-16 by 14-
18 »; wall colorless, thin, 1.5 u, finely and closely verrucose.
II. Urediniospores in the telia, ellipsoid or elliptic-obovoid,
16-19 by 24-324; wall pale golden-brown, thin, 1-1.5 4, thicker
above, 3-5 , closely echinulate, the pores indistinct, 2 or possibly
more, equatorial; paraphyses in the telia, intermixed with the
spores, or surrounding groups of spores, cylindric, clavate or
clavate-capitate, 7-23 by 47-644u, the wall nearly colorless to
golden-brown, darker above, thin, I-1.5 4, somewhat thickened
above, 2—7 u
Il. Telia caulicolous, scattered, oblong or linear, 1-3.5 m
long, somewhat tardily naked, inclined to be pulverulent, a
nut-brown, ruptured epidermis conspicuous; paraphyses sparingly
146 ARTHUR: NEW SPECIES OF UREDINEAE
intermixed, doubtless of uredinial origin, often wanting; telio-
spores apparently in groups, ellipsoid or oblong, 16-21 by 28-35 n,
rounded at both ends, deeply constricted at the septum; wall dark
chestnut-brown, uniformly 1.5-2 4 thick, moderately and closely
verrucose with conical warts, at times almost echinulate; pedicel
short, colorless, fragile.
On Viorna sp., Abilene, Texas, May 17, 1900, T. A. Williams.
The specimen, which was communicated by the National Museum,
Washington, D. C., has been in hand for a considerable time,
hoping that additional material would come to light. The ap-
pearance of the rust is much like that of species belonging to the
genus Tvranzschelia, but the uredinial stage is yet imperfectly
known; moreover, the pycnia are unequivocally subepidermal.
It finds its nearest representative seemingly in Tranzschelia cohaesa
(Long) Arth. (Puccinia cohaesa Long), a species from Texas on
Anemone. But not only are the pycnia subepidermal, both pycnia
and aecia are grouped instead of systematically scattered. It
might be assumed that the aecia here described, which occur upon
the leaves and are entirely dissociated from the caulicular telia, do
not belong to this species, but are heteroecious and the aecia of
Puccinia Agropyri E. & E., which occur upon species of Viorna
further north, and are common in Texas on Clematis Drummondii.
However, the aeciospores measure much smaller than those of any
collection known for that species. An interesting feature is the
grouping of the teliospores, which can be fairly well seen in sections
of sori, which have not yet opened. It is a character exactly in
accord with the telial characters selected for the genus Tranz-
schelia. The paraphyses are also essentially like those of that
genus, both in form and position in the sorus, and the uredinjo-
spores as well. The characters of the latter can not, however, be
fully made out until the uredinial sorus is studied. Altogether it
would seem that in this new and highly distinctive ranuncula-
ceous species of rust we have another candidate for the genus
Tranzschelia, and one which will necessitate some modification in
the generic characters.
Puccinia missouriensis sp. nov.
O. Pycnia epiphyllous, few in small groups, honey-yellow, in-
conspicuous, subepidermal, flask-shaped, 60-75 4 in transverse
diameter, with ostiolar filaments.
ARTHUR: NEW SPECIES OF UREDINEAE 147
I. Aecia hypophyllous, usually few in somewhat indefinite
groups opposite the pycnia, cylindric, 0.3-0.5 mm. long, 0.1-0.2
mm. in diameter; peridium erect, erose; peridial cells rhombic or
rectangular, abutted or somewhat overlapping, 15-23 by 18-27 n,
the outer wall thick, 10-15 n, transversely striate, smooth, the
inner wall much thinner, 2-3, coarsely verrucose; aeciospores
globoid or ellipsoid, 13-16 by 16-20 u: wall colorless, thin, I-1.5 u,
finely and closely verrucose.
III. Telia hypophyllous, scattered among the groups of aecia,
round, early naked, pulvinate, chestnut-brown, ruptured epider-
mis inconspicuous; teliospores oblong, 10-14 by 40-48 yu, obtuse or
rounded at both ends, slightly constricted at septum; wall dark
cinnamon-brown, 1.5-2 4 thick, much thicker above, 5-9u, smooth;
pedicel short, more or less tinted.
On Ranunculus recurvatus Poir., Creve Coeur Lake, Missouri,
May, 1887, L. H. Pammel. The collection came to the writer’s
attention ten years ago, when consulting the cryptogamic her-
barium of the New York Botanical Garden. It was there labelled
Aecidium ranunculacearum, and was for a long time considered one
of the common heteroecious aecia belonging to grass rusts. When
critically examined, however, it was found that the abundant
aecia differed somewhat from those belonging to grasses, and were
accompanied by telia, which had before been overlooked. It is
remarkable that no other similar collection has turned up in the
thirty years since it was gathered.
Puccinia obesispora sp. nov.
II. Uredinia chiefly hypophyllous, confluent on discolored
spots, often in rings about a central area that becomes dry and
dead, or scattered, early naked, cinnamon-brown, ruptured epi-
dermis prominent; urediniospores broadly ellipsoid or globoid,
19-26 by 24-30 u; wall variable in thickness, 1.5-3 u, dark cinna-
mon-brown, sparsely and strongly echinulate, the pores 2, equa-
torial.
__ If. Telia chiefly hypophyllous, scattered, usually following
the uredinia, irregular in outline, 0.I-0.5 mm. across, early naked,
dark chocolate-brown; teliospores ellipsoid or oblong, large, 26-35
by 55-70, rounded at both ends, or obtuse or even acute above,
slightly or not constricted at septum; wall chocolate-brown, thick;
3-4, much thicker above, 6-12 4, coarsely and rather sparsely
verrucose; pedicel tinted near the spore, and twice length of spore,
or less.
148 ARTHUR: NEW SPECIES OF UREDINEAE
On Achyranthes obovata. (Mart. & Gal.) Standley, Oaxaca,
Mexico, October 24, 1899, E. W. D. Holway 3732. The species
is remarkable for its large, coarsely verrucose teliospores, which
greatly resemble those of the euphorbiaceous species, P. Euphor-
biae P. Henn. In that species the uredinia are sometimes accom-
panied by pycnia. In the present species the annular groups of
uredinia appear like a primary stage, but no pycnia could be found
by sectioning or otherwise.
Puccinia incondita sp. nov.
III. Telia hypophyllous and caulicolous, densely pressed or
confluent into more or less rounded pulvinate masses, I-3 mm.
across, dark cinnamon-brown, early naked, ruptured epidermis
inconspicuous; teliospores irregularly ellipsoid or oblong, 16-20
by 23-29 un, rounded at both ends or somewhat narrowed below,
slightly or not constricted at septum: wall cinnamon-brown,
1.5-2.5 uw thick, not or somewhat thicker above up to 6 », smooth;
pedicel colorless, sometimes longer than spore, usually breaking
near the spore.
On Solanum triquetrum Cav.., Austin, Texas, January 5, 1916,
B. C. Tharp. A short-cycle rust which appears to be different
from any species on Solanum heretofore described.
Puccinia adducta sp. nov.
O. Pycnia epiphyllous, in small orbicular groups, dark brown,
noticeable, subepidermal, globoid, 80-105 » broad.
II. Uredinia hypophyllous, at first grouped opposite the pycnia
on larger yellowish spots 2-5 mm. across, afterward scattered, small,
round, rather tardily naked, cinnamon-brown, surrounding epi-
dermis noticeable; urediniospores broadly obovoid, 19-23 by 27-
35m; wall pale cinnamon-brown, 1-2 y thick, moderately and
strongly echinulate, the pores indistinct, possibly 3 and equatorial.
III. Telia hypophyllous, following the uredinia in the same
sori; teliospores ellipsoid or oblong, 19-26 by 35-40 uw, obtuse or
rounded at both ends, slightly or not constricted at septum, wall
cinnamon-brown, I-24 thick, usually with a slight papilla-like
thickening at apex, smooth; pedicel short, rather thick, often
deciduous.
On Solanum racemosum Jacq., Antigua, West Indies, Feb-
ruary 4-16, 1913, Rose, Fitch & Russell 3473. A seemingly dis-
tinctive species.
ARTHUR: NEW SPECIES OF UREDINEAE 149
Puccinia Notopterae sp. nov.
, O. Pycnia amphigenous, crowded on brownish areas 2-3 mm.
across, dark brown, conspicuous, subepidermal, globoid, 85-140 u
in diameter, with ostiolar filaments.
III. Telia epiphyllous, becoming amphigenous, circinating
about the pycnia at first, then becoming confluent and causing
hypertrophy, early naked, very pulverulent, chestnut-brown,
ruptured epidermis noticeable; teliospores ellipsoid, 19-24 by
26-34 wu, rounded at both ends, slightly constricted at septum; wall
dark cinnamon-brown, evenly 1.54 thick; coarsely and rather
sparingly verrucose; pedicel colorless, short, fragile.
On Notoptera hirsuta (Sw.) Urban, Mandeville, Jamaica,
February 23, 1915, E. W. D. Holway 226. A distinctive, short-
cycle rust, on a host genus not before represented among those
bearing rusts.
Aecidium anthericicola sp. nov.
O. Pycnia amphigenous, in small orbicular groups, small,
honey-yellow, inconspicuous, subepidermal.
I. Aecia hypophyllous, in annular groups 2-7 mm. across about
the pycnia, at first bullate then erect, the margin erose; peridial
cells quadrate, 16-20 by 20-26 un, abutted, the outer wall slightly
or no thicker than the inner wall, 3-6 uw; aeciospores globoid, 16-20
by 19-23; wall colorless, thin, 1.54, very finely verrucose, ap-
pearing smooth when wet.
On Anthericum nanum Baker, Dedregal near Tlalpam, Valley
of Mexico, Mexico, June 30, 1905, Rose, Painter & Rose 8246.
The species is similar to the aecia of Uromyces Hordei on Notho-
scordium, but both the peridial cells and aeciospores are smaller
than in that form.
Aecidium plenum sp. nov.
O. Pycnia amphigenous, crowded in groups 0.2-0.3 mm. across,
cinnamon-brown, small, inconspicuous, subepidermal.
I. Aecia hypophyllous, crowded in circular groups about the
pycnia, I-2 mm. across, without conspicuous spots, cupulate or
short cylindric; peridium erect, or somewhat recurved, irregularly
and deeply lacerate; peridial cells angularly oblong in face view,
about 20 by 30, narrowly oblong in radial section, somewhat
overlapping, about 8-10 by 304, the outer wall 5-9 u thick, smooth,
the inner wall 3-6 yu, coarsely verrucose; aeciospores globoid or
ellipsoid, 16-21 by 18-244; wall colorless, thin, Iu, finely and
closely verrucose.
150 ARTHUR: NEW SPECIES OF UREDINEAE
On Argemone intermedia Sweet, Burkburnett, Texas, May 29,
1917, B. O. Dodge. The collection shows a remarkably heavy
infection, the leaves being wholly covered with groups of aecia on
the lower side, with scarcely an aecium showing above. The
groups rarely become confluent and indistinguishable, although
often somewhat intermingling. The general appearance of the
rust is that of a heteroecious form, and is not unlike that on
Agoseris, Lactuca and Crepis, belonging to the Carex species,
Puccinia patruelis. No aecia have heretofore been reported on
any papaveraceous host, and no safe prediction can be made re-
garding its relationship.
Aecidium Thenardiae sp. nov.
I. Aecia hypophyllous, in circular groups 8-10 mm. across,
often circinating, on somewhat larger, yellowish spots, short
cylindric; peridium white, the margin erect, erose, or torn; peridial
cells in face view oblong, 16-22 by 24-30 », somewhat eee
moderately verrucose; aeciospores quadrately globoid, 14-16 p
diameter; wall volatiles: thin, I-I.5y in diameter, Selseghat
thicker above, 2-5 u, finely and closely verrucose.
On Thenardia Galeottiana Baill., near Oaxaca, Mexico, August
20, 1894, C. G. Pringle, communicated by E. W. D. Holway.
The form is undoubtedly heteroecious.
Aecidium Cyrillae sp. nov.
I. Aecia hypophyllous, in peg of 2-8, rarely more, on
reddish-brown spots, 2-4 mm. across, short-cylindric, white;
peridium erect, erose; peridial cells quadrangular i in radial section,
with a strongly overlapping projection, the outer wall A, a
smooth, the inner wall 7-9 thick, verrucose; aeciospores irregu-
larly ellipsoid or globoid, 19-23 by 23-26 nu; wall pene ates thin,
I-I.5u, greatly thickened above, 7-9 y, closely and finely verrucose.
On Cyrilla racemiflora L., Ocean Springs, Mississippi, June 15,
1896, Underwood & Earle. A sparse amount of the rust was
found on the same host in the phanerogamic herbarium of the
New York Botanical Garden, collected at New Orleans, Louisiana,
1832, Drummond 202. A well-marked species, which is probably
heteroecious.
ARTHUR: NEW SPECIES OF UREDINEAE 151
Aecidium Tithymali sp. nov.
O. Pycnia hypophyllous, scattered sparsely over the surface
of the leaf, preceding and accompanying the aecia, punctiform,
honey-yellow or brownish, noticeable, subepidermal, globoid or
flask-shaped, 110-150 broad; ostiolar filaments 60-100 u long.
I. Aecia hypophyllous, evenly and loosely scattered, at first
bullate and opening by a pore or irregular break of the epidermis;
peridia erect or recurved, torn, fragile; peridial cells rhomboidal,
10-15 by 23-274, somewhat overlapping, the outer wall 5-74
thick, striate, the inner wall 2-3, thick, verrucose; aeciospores
globoid or broadly ellipsoid, 18-22 by 19-24 u; wall nearly or quite
colorless, thin, I-1.5 , finely verrucose.
On Tithymalus commutata (Engelm.) Kl. & Garcke (Euphorbia
commutata Engelm.), Decorah, Iowa, June, 1883, E. W. D. Hol-
way; same, June 20, 1885, E. W. D. Holway (Barth. N. Am. Ured.
703); Beloit, Wisconsin, May 30, 1910, J. J. Davis; Lafayette,
Indiana, June 7, 1901, H. B. Dorner (type); same, May 13, 1910,
Kern & Billings; Plummer Island, Maryland, May 30, 1903,
P. L. Ricker 1055; Crawfordsville, Indiana, May 17, 1913, F. D.
Kern.
On Tithymalus leiococcus (Engelm.) Small (Euphorbia texana
Boiss.), San Antonio, Texas, March 16, 1900, Wm. Trelease;
same, February 16, 1914, Arthur & Fromme 5300.
On Tithymalus missouriensis (Norton) Small (Euphorbia
dictyosperma Auct.), Lincoln, Nebraska, April 28, 1902, Geo. G.
Hedgcock.
On Tithymalus robusta (Engelm.) Small (Euphorbia robusta
Small), Colorado, July, 1888, C. H. Demetrio (Ellis & Ev. N. Am.
Fungi 2215); Flagstaff, Arizona, June, 1891, D. T. MacDougal; -
Larimer County, Colorado, July 2, 1894, C. F. Baker 261; Colorado
Springs, Colorado, August 2, 1900, E£. T. Harper 368; Wasatch
County, Utah, August 3, 1905, A. O. Garrett (Fungi Utah. 96);
La Veta, Colorado, 2,100 m. alt., June 21, 1907, F. E. & E. S.
Clements (Crypt. Form. Colo., 594); Boulder, Colorado, May
13, 1910, E. Bethel; same, August 23, 1911, Bethel, Arthur &
Kern 509; Denver, Colorado, May 24 and June 2, 1915, E.
Bethel; Ft. Collins, Colorado, May 23, 1916, J. C. Arthur; Golden,
Colorado, May 27, 1916, J. C. Arthur; Rimrock Station, Montana,
August 2, 1917, E. Bartholomew 6213.
152 ARTHUR: NEW SPECIES OF UREDINEAE
This is a common and conspicuous species in the Rocky
Mountains, and extends sparingly eastward across the plains and
over the Allegheny Mountains. For a long time it has been con-
sidered heteroecious and probably the early stage of the common
Uromyces on Astragalus. Both its distribution and morphological
characters favored this view, and many attempts at cultures were
undertaken to substantiate the assumed connection but without
success. Observations in the field have yielded no strong evidence
that the aecia were followed by telia on Astragalus, but rather
that they were not. At various times it has been suggested that
the telial form on the same hosts as the aecia, Uromyces Tranz-
schelii, an apparently short-cycle species having pycnia associated
with the telia, might be derived at times from the aecia, and all
be one autoecious species. If this were true, we should have the
anomalous condition of a short-cycle and a long-cycle form asso-
ciated at times so intimately that only culture demonstration
could separate them. In the meantime while actual experimental
knowledge is awaited, it seems best to designate the aecia by a
name that will distinguish them from other aecia on similar hosts.
Some of the collections above cited are said to be on Euphorbia
montana, which may be true, although the preponderance of
probability points to E. robusta, a far more common species in the
region covered, and under this name they have been listed.
Aecidium Mozinnae sp. nov.
O. Pycnia chiefly epiphyllous, few in orbicular groups, honey-
yellow, inconspicuous, subepidermal, globoid, 90-125 across;
ostiolar filaments present, abundant.
I. Aecia hypophyllous, encircling the pycnial area, on yellowish
spots 2-5 mm. across; peridium cylindric, the margin erect and
erose; peridial cells quadrate or rectangular, 19-24 by 29-34 b,
abutted or slightly overlapping, the outer wall 10-12 u thick,
transversely striate, smooth, the inner wall 3-5 4 thick, strongly
verrucose; aeciospores angularly ellipsoid or globoid, 19-23 by
24-29 u; wall nearly or quite colorless, 22.5 » thick, closely and
noticeably verrucose.
On Mozinna spathulata (Miill.-Arg.) Ortega (Jatropha spathu-
lata Miill.-Arg.), State of Guanajuato, Mexico, July 11, 1899,
J. N. Rose & Walter Hough 8999.
ARTHUR: NEW SPECIES OF UREDINEAE 153
The species appears to be well defined, and is quite likely
heteroecious.
Aecidium conspicuum sp. nov.
O. Pycnia amphigenous, numerous, crowded in round groups,
0.3-I mm. across, small, honey-yellow, subepidermal, flask-
shaped, 100-125 uw in transverse diameter
I. Aecia amphigenous, in large, dense groups 3-8 mm. across,
on slightly larger discolored spots; peridium erect or somewhat
recurved, finely lacerate; peridial cells angularly oval in face view,
26 by 32-38, oblong in radial section, 16 by 32-38, strongly
overlapping, the inner wall 10-12 uw thick, verrucose, the inner wall
about I 4, smooth; aeciospores globoid, 19-23 4 in diameter; wall
colorless, thin, I wu, finely verrucose.
On Dugaldea Hoopesit (A. Gray) Greene (Helenium Hoopesii
A. Gray), La Plata River, Colorado, 9,500 ft. alt., July 16, 1898,
Baker, Earle & Tracy 1075; mountains near Pagosa Peak, Col-
orado, 9,000 ft. alt., August 10, 1899, C. F. Baker 113; Cloudcroft,
New Mexico, July 19, 1899, E. O. Wooton; Winsor Creek, Pecos
National Forest, New Mexico, July 28, 1908, Paul C. Standley
4581; Rio Pueblo, New Mexico, August 11, 1910, E. O. Wooton;
Little Colorado River, White Mts., Arizona, July 20, 1910, L. N.
Goodding; Trout Lake, Colorado, 10,000 ft. alt., August 2, 1912,
Arthur & Kern 5108; Snowball Creek near Pagosa Springs,
Colorado, 7,200 ft. alt., August 6, 1912, Arthur & Kern 5521
(type); Ute Park, Colfax County, New Mexico, August 25, 1916,
Paul C. Standley 13756.
This rust is quite common in the Rocky Mountains from 7,000
to 10,000 feet altitude. The plants grow from two to three feet
tall, and the long lanceolate leaves are often conspicuously yel-
lowed by the abundant aecia, which show strongly against the
cinereous surface of the host. It is undoubtedly a heteroecious
species. In 1912 Dr. F. D. Kern and the writer searched the two
localities where the rust was found by them for a possible alternate
form, but in vain. Similar efforts have been made by Mr. E.
Bethel, but with no better success.
Aecidium Pereziae sp. nov.
O. Pycnia chiefly epiphyllous, closely grouped, honey-yellow
becoming brown, punctiform, noticeable, subepiderma
154 ARTHUR: NEW SPECIES OF UREDINEAE
‘I. Aecia hypophyllous, numerous, opposite the pycnia, in well-
defined groups 0.3-0.6 mm. across, on roseate or yellowish spots
I0-I5 mm. across, cupulate; peridium usually erect, the margin
erose or lacerate; peridial cells easily separating, rhomboid, little
or no longer than broad, 19-26 by 26-37 u, the outer wall 8-10 p
thick, faintly striate, the inner wall 3-6 u thick, moderately ver-
rucose; aeciospores globoid, 13-18 by 16-19 y; wall nearly or quite
colorless, thin, I-1.5 4, very minutely verrucose, appearing smooth
when wet.
On Perezia sp., Barranca, Mexico, July 25, 1893, C. G. Pringle,
communicated by W. G. Farlow. A distinctive species, and
doubtless heteroecious.
Aecidium steviicola sp. nov.
O. Pycnia amphigenous, numerous, in loose groups, punctate,
honey-yellow becoming brown, noticeable, subepidermal.
I. Aecia hypophyllous, in loose groups surrounding the pycnia
on yellowish spots 10-15 mm. across, cupulate, the margin some-
what reverted, erose or lacerate; aeciospores globoid, large, 24-32
by 30-40; wall colorless, rather thin, about 1.5 uw, considerably
thickened above, 6-9 4, minutely and closely verrucose.
On Stevia sp., Popo Park, Federal District of Mexico, August
4-8, 1910, A. S. Hitchcock.
This is doubtless a heteroecious form. It much resembles
A. roseum Diet. & Holw., which occurs in Mexico on species of
Stevia, but more abundantly on Eupatorium, and which is be-
lieved to be the aecial stage of a grass rust, probably A egopogon,
but the spores are nearly half as much longer than in that form,
although in other respects there is close similarity.
Aecidium Keerliae sp. nov.
O. Pycnia amphigenous, in close groups I-2 mm. across,
honey-yellow becoming brown, punctiform, noticeable, subepi-
dermal.
I. Aecia hypophyllous, in groups 3-8 mm. across, on somewhat
larger, yellowish spots, cupulate, low; peridium slightly recurved,
erose or lacerate; aeciospores globoid or ellipsoid, small, 12-18
by 15-20; wall nearly or quite colorless, thin, 1 #, much thicker
above, 3-6 y, finely and closely verrucose.
On Keerlia mexicana A. Gray, Guadalajara, Mexico, July 14,
ARTHUR: NEW SPECIES OF UREDINEAE 156
1893, C. G. Pringle, communicated by W. G. Farlow. A species
of the general characteristics of A. roseum and A. steviicola, both
of which are also on carduaceous hosts, but with very much
smaller spores. It is likely to prove to be heteroecious.
Uredo egenula sp. nov.
II. Uredinia epiphyllous, Se oblong or linear, 0.1—-1.2
mm. long, early naked, pulverulent, cinnamon-brown, ruptured
epidermis evident; anes potas globoid to broadly ellipsoid,
21-25 by 26-344; wall cinnamon-brown, 1.5-2.5 thick, moder-
ately echinulate, the pores 2, equatorial, prominent.
On Sporobolus argutus (Nees) Kunth, alkali soil below Gregory
Park, Healthshire Hills, Halfmoon Bay, Jamaica, West Indies.
The specimen was removed from a phanerogamic collection in
the National Herbarium, made October 31, 1912, by A. S. Hitch-
cock 9760, and communicated by Mrs. Agnes Chase. The form
is one that seems to fit no recorded species on Sporobolus. It is
named and described in order to call attention to the need of
search for the telial stage, and of observations that may eventually
lead to finding the aecia.
Uredo panamensis sp. nov.
II. Uredinia hypophyllous, crowded in Sn ——
2-3 mm. across on slightly discolored spots of nearly same
size, round or oblong, 0.3-0.5 mm. in diameter, the Bet one
usually much larger, subepidermal, early naked, applanate,
cinnamon-brown, ruptured epidermis evident; urediniospores
globoid or broadly ellipsoid, 20-27 by 24-31 4; wall cinnamon-
. brown, rather thick, 2-3 u, very closely echinulate, often appearing
verrucose, the pores distinct, 3, approximately equatorial.
On Phytolacca decandra L., Panama, December 7, 1915,
E. W. D. Holway 234. The sorus is without special inclosing
structures, and resembles the form common for the genera Puc-
cinia and Uromyces. The species is quite unlike Puccinia
Rivinae (B. & C.) Speg., the only other rust known on this family
of hosts.
Uredo unilateralis sp. nov.
II. Uredinia hypophyllous, in groups 2-4 mm. across, more or
less circinate, or scattered, round, 0.2-0.8 mm. in diameter, soon
156 ARTHUR: NEW SPECIES OF UREDINEAE
naked, cinnamon-brown, somewhat pulverulent, ruptured epi-
dermis evident; urediniospores broadly obovate, or spatulate-
obovate, flattened or concave on one side, 20-26 by 26-32 u; wall
cinnamon-brown, uniformly thick, 1.5-2 4, sparsely and noticeably
echinulate except on concave side which is smooth, with one pore
on concave side, subequatorial, sometimes near hilum.
On Geranium mexicanum H.B.K., Amecameca, Mexico,
October 6, 1900, E. W. D. Holway. A unique form on account of
the spore having a smooth, concave side, bearing the single pore.
PuRDUE UNIVERSITY,
LAFAYETTE, INDIANA
A striking variation in Silene noctiflora
E. P. HUMBERT
'
(WITH TWO TEXT FIGURES)
From some Silene noctiflora seeds planted in the greenhouse
December 1, 1917, one very interesting plant has developed. The
normal Silene noctiflora seedling has two seed-leaves and suc-
ceeding leaves are in whorls of two, or opposite, each pair being
placed over the intervals between the preceding pair. The pairs
cross at right angles or decussate. This is illustrated in Fic. 1,
a reproduction of a normal seedling. The plant which is the
FIG. I. Fic. 2.
Fic. 1. Normal Silene noctiflora seedling with two seed-leaves and two leaves.
in each succeeding whorl, X 2.
Fic. 2. Silene noctiflora seedling with three seed-leaves and three leaves in each
of the two succeeding whorls, X 2.
occasion of this sketch produced three seed-leaves and the suc-
ceeding leaves were arranged in whorls of three, each leaf of the
157
158 HumBeErtT: A STRIKING VARIATION IN SILENE NOCTIFLORA
new whorl being placed above an interval between leaves of the
preceding whorl. Fic. 2 is reproduced from a photograph of this
plant.
It is no uncommon thing to find Silene noctiflora seedlings with
divided seed-leaves. All gradations have been noted from the
extreme where both seed-leaves are completely divided, giving the
appearance of four seed-leaves, to a partial division of one seed-
leaf. When one seed-leaf is divided the seedling has the appear-
ance of a plant with three seed-leaves. In all such monstrosities,
however, the leaves which follow the seed-leaves are opposite and
the plant is thereafter quite normal. The plant here pictured in
Fic. 2 is the first one observed to show a completely altered
phyllotaxy.
Silene noctiflora seeds were secured from the Department of
Plant Breeding, Cornell University, in the fall of 1916, and an
attempt was made to grow seedlings in the garden in 1917.
Only one (very much stunted) plant produced seed, due to un-
favorable environment. The seed from this plant produced the
seedlings pictured.
AGRICULTURAL EXPERIMENT STATION,
COLLEGE STATION, TEXaAs
INDEX TO AMERICAN BOTANICAL LITERATURE
1911-1918
The aim of this Index is to include all current botanical literature written by
Americans, published in scenes or based upon American material ; the word Amer-
ica being used in the broadest sen
Reviews, and papers that aie exclusively to forestry, agriculture, horticulture,
manufactured products of vegetable origin, or laboratory methods are not included, an
© attempt is made to index the literature of bacteriology. An occasional exception is
ag in favor of some paper appearing in an American peri riodical which is devoted
some important particular, If users of the Index will call the attention of the editor
to errors or omissions, their kindness will be appreciated.
This Index is reprinted monthly on cards, and furnished in this form to subscribers
at the rate of one cent for each card, Selections of cards are not permitted ; each
subscriber must take all cards published during the term of his subscription, Corre-
spondence relating to the card issue should be addressed to the Treasurer of the Torrey
Botanical Club
Alway, F. J., & McDole, G. Relation of movement of water in a soil
to its hygroscopicity and initial moistness. Jour. Agr. Research ro:
391-428. f. I, 2. 20 Au 1917.
Atkinson, G. F. Charles Horton Peck. Bot. Gaz. 65: 103-108. 17
Ja 1918. [Portraits.]
Bailey, L. H. The caoders systematist. Science II. 46: 623-629.
28 D 1917.
Bailey, W. W. Amphicarpaea. Am. Bot. 23: 120-122. N 1917.
Baker, C. F. The botanic garden of Para. Pomona Jour. Econ. Bot.
1: 64-72. f. 22-28. | F 1911.
Bakke, A. L. Studies on the transpiring power of plants as indicated
by the method of standardized hygrometric paper. Jour. Ecol. 2:
10-173. f. 22, 23. 5S 1914.
Barnhart, J. H. Philippe de Vilmorin. Jour. Hort. DOC I A: 2:
231-234. N 1917.
Barrett, J. T. Thomas Jonathan Burrill (1839-1916). Phytopathol-
ogy 8: 1-4. pl. zr. Ja 1918.
Bates, J. M. Anew oak. Am. Bot. 23: 119, 120. N 1917.
Species not named as yet.
Belling, J. Selection of plant-breeding. Jour. Heredity 9: 95. 28
1918.
— 159
160 INDEX TO AMERICAN BOTANICAL LITERATURE
Benoist, R. Descriptions d’espéces nouvelles de !égumineuses de
la Guyane Francaise. Not. Syst. 3: 271-274. 1916.
Parkia velutina, Dimorphandra polyandra and Eperua kourouensis, spp. nov.,
are described.
Berry, E. W. ‘The fossil plants from Vero, Florida. Rep. Florida
Geol. Surv. 9: 19-33. I917.
Bicknell, E. P. Aster cordifolius. Addisonia 2: 79, 80. pl. So. 31
D 1917.
Bowman, H. H. M. Ecology and physiology of the red mangrove.
Proc. Am. Philos. Soc. §6: 589-672. pl. 4-9 + f. 1-3. 8 D 1917.
Boynton, K.R. Centradenia floribunda. Addisonia 2: 65, 66, pl. 73.
2% D 101t7.
Boynton, K.R. Anneslia Tweediei. Addisonia 2: 75. pl. 78. 31 D
1917.
Bregger, T. Linkage in maize; the C aleurone factor and waxy
endosperm. Am. Nat. 52: 57-61. Ja 1918.
Britton, N. L. Piaropus azureus. Addisonia 2: 67, 68. pl. 74. 31
D 1917.
Brooks, C., & Fisher, D. F. Irrigation experiments on apple-spot
diseases. Jour. Agr. Research 12: 109-138. pl. 2-5 +f. I-10.
21 Ja 1918.
Brown, W. H., Merrill, E. D., & Yates, H. S. The revegetation of
Volcano Island, Luzon, Philippine Islands, since the eruption of
Taa Volcano in 1911. Philip. Jour. Sci. 12: (Bot.) 177-248.
bl. T“10 fe 1,72. TL AGF.
Claassen, E. Second alphabetical list of the lichens collected in several
counties of Northern Ohio. Ohio Jour. Sci. 18: 62,63. 31 D 1917.
Clute, W. N. Food from wild plants. Am. Bot. 23: 131, 132. N
1917; 24: 16-25. F 1918.
Clute, W. N. The fruit of the potato. Am. Bot. 23: 115, 116.
N 1917. [Illust.]
Coleman, D. A. Environmental factors influencing the activities of
soil fungi. Soil Sci. 2: 1-66. f. r-r10. Jl 1916.
Collins, G. N. & Kempton, J. H. Breeding sweet corn resistant to
the corn earworm. Jour. Agr. Research 11: 549-572. 10 D 1917.
Cook, M. T. The study of plant diseases in the high school. School
Sci. & Math. 16: 351-353. 1916
Crawford, D.L. A biological expedition to Southern Mexico. Pomona
Jour. Econ. Bot. 1: 57
Cribbs, J.E. A columella in Marchauias polymorpha. Bot. Gaz. 65:
Of-06. DI. y, 2. 17 Ja ors.
INDEX TO AMERICAN BOTANICAL LITERATURE 161
Cruz, D. da. A contribution to the life history of Lilium tenuifolium.
I-37. pl. 1-7. Washington. 1915.
Davis, A.M. A white-leaved hemlock in Vermont. Rhodora 19: 273.
27 D 1917.
Davis, B. M. A criticism of the evidence for the mutation theory of
De Vries from the behavior of species of Oenothera in crosses and
selfed lines. Proc. Nat. Acad. Sci. 3: 704-710. D 1917.
Dearing, C. Muscadine grape breeding. Jour. Heredity 8: 409-424.
f. ro-18 25 Au 1917.
East, E. M., & Park, J. B. Studies on self- peediee The behavior
of self-sterile plants. Genetics 2: 505-609.
Edgerton, C. W. A study of wilt resistance in Si seed-bed. Phyto-.
pathology 8: 5-14. f. 1-4. 24 Ja 1918. ;
Elmore, C. J. Thomas County diatomite. Nebraska Geol. Surv. 7°:
Bi, 52. dle re 35 Jota.
Evans, A. W. Notes on New England Hepaticae——XIV. Rhodora
19: 263-272. 27 D 1917.
Fairchild, D. The grafted jujube of China. Jour. Heredity 9: 3-7.
f. 1-4 + frontispiece. Ja 1918.
Fernald, M. L. A remarkable colony of Bidens in Connecticut.
Rhodora 19: 257-259. 27 D 1917.
Bidens heterodoxa var. monardaefolia and B. heterodoxa var. agnostica.
Gager, C.S. The near future of botany in America. Science II. 47:
IOI-115. 1 F 1918.
Giddings, N. J. Potato and tomato diseases. West Virginia Agr.
Exp. Sta. Bull. 165: 1-24. f. 1-20.. S 1917.
Grier, N. M. Sexual dimorphism and variation in Ginkgo biloba, L.
Torreya 17: 225. 24 Ja 1918.
Grisdale, J H. The black or stem rust of wheat. Canada Dept. Agr.
Exp. Farms. Div. Bot. II. Bull. 33: 1-15. 1917. [Illust.]
— illustration in the text is republished in color as Circ. No. 12 of the Central
Exp.
a a R.C. Rapid respiration after death. Proc. Nat. Acad. Sci.
3: 688-691. D 1917.
Hagstrém, J.O. Three species of Ruppia. Bot. Not. 1911: 137-144.
J. 28. 1911.
Halstead, B. D. Colors in vegetable fruits. Jour. Heredity 9: 18-
23. Jai9
Hansen, A. A. Petalization in the Japanese quince. Jour. Heredity
9: 15-17. f. 5, 6. Ja 1918
Hansen, A. A. A striking reproductive habit. Jour. Heredity 9: 85.
f. 19. 28 Ja 1918.
162 INDEX TO AMERICAN BOTANICAL LITERATURE
Hansen, A. A. Uses of the bay berry. Am. Bot. 23: 117-119. N
1917. [Illust.]
Harper, R. M. The plant population of northern lower Michigan and
its environment. Bull. Torrey Club 45: 23-42. f. 1-3. 8 F 1918.
Harter, L.L. Podblight of the lima bean caused by Diaporthe phaseo-
lorum. Jour. Agr. Research 11: 473-504. pl. 42, 43 +f. I-II.
3 D 1917. |
Hauman, L. Note préliminaire sur les Hordewm spontanés de la flore
Argentine. An. Mus. Nac. Hist. Nat. Buenos Aires 28: 263-316.
fl. 70-13 +f. 1. 223 1916.
Havemeyer, T. A. The foxtail lily. Jour. Internat. Gard. Club 1:
431-434. D 1917. [Illust.]
Hedrick, U. P. and others. The peaches of New York. Ann. Rep-
N. Y. Dept. Agr. 24-vol. 27. i-xiii + 1-541. 1917. [Illust.]
Hopkins, L. S. An interesting T rillium. Ain. Bot; 23:°126, 127.
N 1917. [Illust.]
Howitt, J. E., & Caesar, L. The more important fruit tree diseases
of Ontario. Ontario Dept. Agr. Bull. 257: 1-44. D 1917.
Ichimura, T. A new poisonous mushroom. Bot. Gaz. 65: 109—-IIT:
f. I-3. a Ja 1918.
iss, th. sp. nov
Jagger, I. C. Hoste of ‘e hite pickle mosaic disease of cucumber.
Phytopathology 8: 32, 33. 24 Ja 1918.
Jagger, I. C., tewart, V. B. Some Verticillium diseases. Phyto-
pathology 8: 15-19. 24 Ja 1918.
Jehle, R. A. Susceptibility of Zanthoxylum clava-hercules to Bac-
terium citri. Phytopathology 8: 34-35. 24 Ja 1918.
Jenkins, E. H., & Street, J. P. Manure from the sea. Connecticut
Agr. Exp. Sta. Bull. 194: 1-13. pl. 1-7. Jl 1917.
Contains considerable information of a botanical nature.
Johnston, J. R. Notas sobre micologia y pathologia vegetal en Cuba.
Mem. Soc. Cubana Hist. Nat. 2: 225-228. 1917.
Jones, L. R., Johnson, A. G., & Reddy, C. S. Bacterial-blight of
barley. Jour. Agr. Research 11: 625-644. pl. B, 47-49 +f. 1, 2
17 D 109t7.
Jones, M. E. Contributions to western botany. No. 14: 1-52. 29
Je 1912.
Kearney, T. H. A plant industry based upon mutation. Jour.
naa g: 51-61. f. 1-8 + frontispiece. 28 Ja 1918.
, & Anderson, E. M. Botanical specimens collected in
the — district, 1916. Rep. Provincial Mus. Nat. Hist.
British Columbia 1916. Q 25-Q 43. pl. 9, 10. 1916.
INDEX TO AMERICAN BOTANICAL LITERATURE 163
Knowlton, C. H. Preliminary lists of New England plants,;—XXIV.
Rhodora 18: 245-248. 1 D 1916;—XXV. Rhodora 19: 217-219.
11 O 1917.
Leonard, E. C. The Astereae of Ohio. Ohio Jour. Sci. 18: 33-58.
31 D 1917.
Lewis, F. J Vegetation distribution in the Rocky Mountains Park.
Canadian Alp. Jour. 8: 87-95. f. 1-5. 1917.
Lofgren, A. Especies, variedades, hybridos, seleccdo, natural, muta-
cors, hereditariedade, lei de Mendel, chromasomos. A Lavoura 21:
348-359. 1917. [Illust.]
Long, W. H., & Harsch, R. M. Pure cultures of wood-rotting fungi
on artificial media. Jour. Agr. Research 12: 33-82. 14 Ja 1918.
Love, H. H., & Craig, W. T. Small grain investigations. Jour.
Heredity 9: 67-76. f. 9-15. 28 Ja 1918.
Mackenzie, K. K. Solidago altissima. Addisonia 2: 69,.70. pl. 75.
i 2 1007.
Marsh, C. D., & Clawson, A. B. Eupatorium urticaefolium as a
Sere plant. Jour. Agr. Research 11: 699-716. pl. 52-55.
24 D1
Wovens x . Algae of the Hawaiian Archipelago. I. Bot. Gaz.
65: 42-57. 17 Ja 1918; II. Bot. Gaz. 65: 121-149. 15 F 1918.
MacCaughey, V. The Hawaiian taro. Am. Bot. 23: 122-126. N
1917 :
Colocasia antiquorum var. esculenta
McClintock, J. A. Further eine relative to the varietal resistance
of peanuts to Sclerotium Rolfsii. Science I] 47: 72,73. 18 Ja1g18.
Massey, L. M. Experiments for the control of blackspot and powdery
mildew of roses. Phytopathology 8: 20-23. 24 Ja 1918.
Melchers, L. E. Physoderma (Zeae Maydis?) in Kansas. Phyto-
pathology 8: 38, 39. 24 Ja 1918.
Melhus, I. E., & Durrell, L. W. The barberry bush and black stem
rust of small grains. Iowa Agr. Exp. Sta. Circ. 35: [1-4]. f. 1-6.
Ap 1917.
Merrill, E. D. New Philippine shrubs and trees. Philip. Jour. Sci.
42: (Bot.) 263-303. 5 1917.
Includes descriptions of 44 new species and the new genus Trifidacanthus.
Mitra, S. K. Toxic and antagonistic effects of salts of wine yeast
ACHE 6 ellipsoideus). Univ. Calif. Publ. Agr. 3: 63-102.
f. 1-12. 30N 19
Munn, ic T. Pathogenicity of Bacillus amylovorus (Burr.) Trev. for
blossoms of the strawberry (Fragaria sp.). Phytopathology 8: 33.
24 Ja 1918.
164 INDEX TO AMERICAN BOTANICAL LITERATURE
Murrill, W. A. The delicious fruits of Actinidia. Jour. N. Y. Bot.
Gard. 18: 257-259. D 1917.
Murrill, W. A. Western polypores. i-iv + 1-36. New York. 1915.
Nash, G. V. Crassula quadrifida. Addisonia 2: 77. pl. 79. 31 D
Nash, G. V. Dendrobium atroviolaceum. Addisonia 2: 63. pl. 72.
31 D 1917.
Nash, G. V. Freylinia lanceolata. Addisonia 2: 73. pl. 77. 31 D
Nash, G. V. Pentapterygium serpens. Addisonia 2: 71. pl. 76. 31
D 1917.
Nash, G. V. Rosa ‘Silver Moon.’ Addisonia 2: 61, 62. pl. 71.
31 D 1917.
A garden hybrid.
Nelson, A., & Macbride, J. F. Western plant studies. V. Bot. Gaz.
65: 58-70.
Includes Trifolium Leibergii, Gentiana Covillei and Pentstemon Albrightii, spp.
nov. :
Pammel, L. H., King, C. M., & Seal, J. L. Studies on a Fusarium
disease of corn and sorghum (preliminary). Iowa Agr. Exp. Sta.
Research Bull. 33: 115-136. f. 2-15. Mr 1916.
Piper, C. V., & Shull, J. M. Structure of the pod and the seed of the
Georgia velvet bean, Stizolobium deeringianum. Jour. Agr. Re-
search 11: 673-677. pl. 50, 51. 24 D 1917.
Popenoe, F. W. ‘The avacado in southern California. Pomona Jour.
Econ. Bot. 1: 3-24. f. 1-13. F 1911
Popenoe, P. Meanings of genetic terms. Jour. Heredity 9: 91-94
28 Ja 1918.
Porto, P. C. Contribuicgao para o conhecimento da flora orchidacea
da Serra do Itatiaya. Arch. Jard. Bot. Rio de Janeiro 1915: 1-22.
1915.
ing, G. Aquatic gardening. Jour. Internat. Gard. Club. 1:
435-454. D 1917.
Ramsey, G. B. Influence of moisture and temperature upon infection
of Spongospora subterranea. Phytopathology 8: 29-31. 24 Ja 1918.
Robbins, W. W. Native vegetation and climate of Colorado in their
relation to agriculture. Colorado Agr. Exp. Sta. Bull. 224: 1-56.
7.220; ¥F 1947.
Roig, J. T. Plantas nuevas o poco Conocidas de Cuba. Mem. Soc.
Cubana Hist. Nat. “Felipe Poey’’ 2: 109-123. Au 1916; 210-222.
1917.
INDEX TO AMERICAN BOTANICAL LITERATURE 165
Rolfe, R. A. Cryptophoranthus Dayanus. Curt. Bot. Mag. IV. 13:
pl. 8740. O 1917.
From Colombia, South America.
Russell, E. J. Recent investigations on the production of plant food
in the soil. Jour. Internat. Gard. Club. 1: 317-346. D 1917.
Rydberg, P. A. Flora of the Rocky Mountains and adjacent plains.
i-xii I-1110. . 31 D 1917.
Includes nas nia of 13 new genera and 52 new species, and many new
combinatio
Safford, E. Chelonocarpus, a new section of the genus Annona,
with descriptions of Annona scleroderma and Annona testudinea.
Jour. Wash. Acad. Sci. 3: 103-108. f. 1-3. 19 F 1913.
Sargent, C. S. Notes from the Arnold Arboretum. Jour. Internat.
Garden Club. 1: 361-380. D 1917. [Illust.]
Reprinted from various bulletins of the arboretum.
Schneider, C. A conspectus of Mexican, West Indian, Central and
South American species and varieties of Salix. Bot. Gaz. 65: 1-41.
17 Ja 1917.
Includes Salix Schaffneri, S. Rowleei spp. nov.
Seward, A.C. Fossil plants, 3: i-xviii + 1-656. f. 377-629. London.
1917.
Shamel, A. D. Chrysanthemum varieties. Jour. Heredity 9: 81-84.
28 Ja 1918. :
Sheepers, J. May-flowering tulips and how they may be advan-
tageously planted. Jour. Internat. Gard. Club. 1: 293-316. D
1917. [Illust.]
Shufeldt,R.W. Flowers, feathers, and fins. Am. Forest. 23: 669-674.
f, I-I12.. N' 1937.
Shufeldt, R. W. Multiple button-balls. Jour. Heredity 8: 550-552.
f.6. D 1917
Shufeldt, R. W. Plants and animals of the Atlantic and Gulf states.
Am. Forest. 23: 743-747. f. 1-11. D 1917.
Skan, S. A. Castilleja miniata. Curt. Bot. Mag. IV. 13: pl. 8730.
O 1917.
A North American plant.
Small, J. K. The Lo Anamomis in Florida. Torreya 17: 221-224.
J, 22°24: Ja-15
Anamomis PTE sp. nov. is described.
Smith, C. P. Studies in the genus Lupinus—II. The Microcarpi,
exclusive of Lupinus densiflorus. Bull. Torrey Club. 45: 1-22.
f. 7-16. 3 PF 193s.
166 INDEX TO AMERICAN BOTANICAL LITERATURE
Smith, J. J. The Amboina Orchidaceae collected by C. B. Robinson.
Philip. Jour. Sci. 12: (Bot.) sin age S 1917.
Includes descriptions of five new species
Standley, P.C. (Chenopodiales) Allioniaceae. N. Am, Fl. 23» 171-
254. 22 Ja1918.
Includes descriptions of nine new species.
Stapf,O. Sechium edule. Curt. Bot. Mag. IV. 13: pl. 8738. O 1917.
From tropical America.
Stevens, N. E. Temperatures of the cranberry regions of the United
States in relation to the. growth of certain fungi. Jour. Agr. Re-
search FE: 521-520. f. 1-3. 3 D 1917.
Stewart, F. C. A Phoma blight of red cedar. Phytopathology 8:
33, 34. 24 Ja 1918.
St. John, H. Arenaria laterifiora and its varieties in North America.
Rhodora 19: 259-262. 27 D 1917.
Stokey, A.G. Apogamy in the Cyatheaceae. Bot. Gaz. 65: 97-102. _
fe t-20.. 17 Ja 1918:
Stone, R. E. Orange rust of Rubus in Canada. Phytopathology 8:
27-29. f,.1. 24 Ja 1918.
Tisdale, W. H. Flaxwilt: a study of the nature and inheritance of
wilt resistance. Jour. Agr. Research 11: 573-606. pl. 44-46 +f.
T7400 P1917.
True, R. H. Notes on osmotic experiments with marine algae. Bot.
Gaz. 65: 71-82. 17 Ja 1918.
Tupper, W. W., & Bartlett, H.-H. The relation of mutational char-
acters to cell size. Genetics 3: 93-106. f. 1, 2. Ja 1918.
Uphoff, J. C. T. Cold resistance in spineless cacti. Arizona Agr.
Exp. Sta. Bull. 79: 1-144. pl. 1, 2+/f. 1-10. 1D 1916.
Van Fleet, W. Possibilities in the production of American garden
roses. Jour. Internat. Gard. Club 1: 485-496. D 1917. [Illust.]
Van Hermann, H. A. Marabo. Dichrostachis nutans. Mod. Cuba
Mag. 1: 7-9. N'1913. [Illust.]
Van Hermann, H. A. Citrus canker. Mod. Cuba Mag. 2: 78-84.
O 1914. [Hlust.]
Vries, a de. Croisements et mutations. Scientia 20: 1-12. S 1916.
Vsies, H. de. Mutations of Oenothera suaveolens Desf. Genetics 3:
Wakefield, E.M. Fungi exotici. XXIII. Kew Bull. Misc. Inf. 1917:
308-314. 1917.
Includes Tilletia Wilcoxiana from America.
BULL. ToRREY CLUB VOLUME 45, PLATE 6
CYANEA GIFFARDII Rock
AN ILLUSTRATED FLORA
OF THE
NORTHERN UNITED STATES, CANADA
AND THE BRITISH POSSESSIONS
From Newfoundland to the Parallel of the Southern Boundary of Virginia, and
from the Atlantic Ocean Westward to the 102d Meridian
By
NATHANIEL LORD BRITTON, Ph.D., Sc.D., LL.D.
Director-in-Chief of the N. Y, Botanical Garden; Professor in Columbia University
And M
HON. ADDISON BROWN, A.B., LL.D.
President of the New York Botanical Garden
Three Volumes. $13.50 Special Net; Expressage extra
SECOND EDITION—REVISED AND ENLARGED
CHARLES SCRIBNER’S SONS
FIFTH AVENUE AT 48th STREET NEW YORK
American fern Journal
An illustrated quarterly devoted to the general study of ferns, Subscription, with
membership in the AMERICAN FERN SOCIETY, $1.00 per annum; without mem
bership, 90 cents. Volume I, six numbers, $2.00; other volumes, $1.00 each, Matter
or publication should be sent to R. C. BENEDICT, 322 East 19th Street, Brooklyn,
N. Y,; subscriptions and other business communications to E, J. WINSLOW, Auburn-
Mass.
THE BRYOLOGIST
THE ONLY MAGAZINE IN THE WORLD RELATING
EXCLUSIVELY TO THE STUDY OF MOSSES,
HEPATICS, AND LICHENS
Illustrated with original drawings and half-tones. Twenty-first volume
begins with January, 1918. Send for sample copy; subscription price
$1.25 per year. Address
EDWARD B. CHAMBERLAIN,
18 West 89th Street, NEW YORK CITY.
Large Dissecting Stand
A new design which meets every requirement by reason of its
large size and wide range of usefulness. Accommodates inter-
changeably Binocular Microscope body (illustrated), Monocular
Erecting Body, using medium high power objectives, or simple lens
in jointed arm—all focused by rack and pinion. The microscope
bodies may be mounted on a sliding track, permitting the entire
width of the stage to be covered.
Stage, 8x7 inches, is provided with glass and metal plates,
4l4 inches in diameter, and with adjustable background stops be-
neath.
With 38 and 19mm. doublet lenses . . . . . . . $26.50
With Binocular Microscope body, 40 mm. yk a ies and
10X eyepieces, as illustrated above . . - $70.00
Write for illustrated circulars.
Rausch 6 lomb Optical ©.
NEW YORK FRANCISCO
LONDON ROCHESTER. NY. FRANKFORT
Vol. 45 No. 5
BULLETIN
OF THE
TORREY BOTANICAL CLUB
MAY, 1918
Studies in the genus Lupinus—Ill, Lupinus densiflorus
CHARLES PIPER SMITH
(WITH TEXT FIGURES 17-42)
Having, in my last paper, disposed of all the other species of
the Microcarpi, I will confine this paper to my discussion and
classification of L. densiflorus and its numerous varieties.
6. LUPINUS DENSIFLORUS Benth. Trans. Hort. Soc. II. 1: 410.
1835. [FiG. 17.]
The original description reads:
. annuus, caulibus adscendentibus basi foliosus foliisque subsericeo-pilosis,
foliolis oblongo-spathulatis, verticillis numerosis approximatis 6-10 floris, sh
ebracteatis, calycis labiis stapes ce superene membranaceo inferiore piloso dup
cigar leguminibus villosis disperm
he flowers, which grow in aes whorls, are om delicately stained with
oe they are also a little speckled at the base of the vexillu The leaves are closely
clustered atone are covered with fine soft hairs, and a has about nine narrow
divisions. The stem does not grow above six or seven inches high
is species nie hitherto produced its seeds, which are of an olive green, smooth,
and minutely dotted with black, in very small quantity. It is probable that it re-
quires shade
In the same year Lindley, in the Botanical Register (20: pl.
1689), published a very interesting plate of L. densiflorus (see Fic.
17), and, in addition to quoting Bentham’s description, gave a
description of his own. From this I quote:
Annuus; caulo erecto .. . villoso, in spontanea pedunculo communi multo™
breviore . . . foliolis in sis cickeinians pedancals adapts eulta eo: ree
erticilli villosissimi, 6-10 flori . -
Calyx villosus, in cultu tantum ca teegcetia? bracteolis setaceis labii superioris seniei-
[The BULLETIN for April (45: 133-166, pl. 6) was issued May 1, r918.]
167
168 SMITH: STUDIES IN THE GENUS LUPINUS
tudine; lab. sup. bipartito . . . inter
medio minimo, superiore breviore. (Obs.
partes in icone incuria pictoris false delineantur.) Vexillum lacteum, acutiusculum,
S e" ib)
wb dt
Soho
Fic. 17. LuPINus DENSIFLORUS
Benth. Copied from the original
plate.
asi viridi-punctatum; alae et carina acu-
minatae, roseae. . . . Semina olivacea,
laevia, nigro maculata.
Lindley thus records his obser-
vation as to the differences be-
tween the specimens collected by
Douglas in California and those
grown in England from seed also
collected by Douglas. While the
plate may not be altogether satis-
factory, it at least shows that the
flowers were spreading in anthesis,
and substantiates my conception
of the species sensu lato.
About the same time Agardh
saw the specimens in Lindley’s’
herbarium and decided that there
were at least two species. In his
monographic “ Synopsis Generis
Lupini,”’ also bearing the date of
1835, he described his L. Menziesii
and made his own disposition of
Bentham’s species, without mak-
ing mention of the specimens grown
from seed. His two descriptions,
in part, follow:
2. L. Menziesii nob. floribus in spica lon-
gissima verticillatis pedicellatis, pedicellis
bracteas setaceas persistens subaeque ntibus,
calycis ebracteolati labiis integris, superiore
scarioso, inferiore herbaceo duplo longiore.
Hab. A Douglas e California reportatum.
Vidi in Hb. Lindleyi.
Pedunculus aenstres longissimus (ultra
pedalis) . . . subglaber, inferne pilis sparsis.
- Foliola supra glabre, subtus piloga: . ...
Verticilli longe distantes, 5-6 flori,
Calyx ebracteolatus, ob bien , labio inkerts
ore viridi; maximo, 2 Regge supeérius scariosum, acuminatum plus duplo super-
ante. Corollae lu . . Carina valde curvata, apice ochracea.
SMITH: STUDIES IN THE GENUS LUPINUS 169.
ularis species primo aspectu corollis luteis verticillatis L. luteum referens;
notis ates tamen abunde diversa. Calyce singulari ad sequ. accedens, inter utram-
que collocanda videtu
4. L. densiflorus shia floribus in spica densa verticillat . bracteis per-
sistentibus reflexis, corollam aequa ric calycis ebracteolati Gea sup. emarginato,
ato
inf. duplo longiore oie las triden
; densiflorus Benth. Hort. “ane n. ser. v. I p. 409. “Edwards Bot. Register. t.
1689.
Hab. E. California reportavit Douglas. Vidi in Hb. Lindl.
Habitus et fere characteres omnes praecedentis, sed robustior et villo longiore
praecepue in partibus sndarseisies obsitus. . . n speciminibus, quae coram habeo,
a ia, insignis; . . chai etiam aliquantulum ats ita ut
es hic latere, rete aheiats
ae, treatment, ire because published in a mono-
graph, was followed by Torrey and Gray in their Flora of North
America; but Bentham (Pl. Hartweg. 303. 1848), some fifteen
years after publishing his species, recorded a Monterey specimen
thus:
1692 (53). Lupinus densiflorus, Ag. —Torr. & Gr. Fl. N. Amer. 1, p. 371, excl.
. Benth.—In pascuis spe Monterey.—Planta Douglasiana quam sub nomine L.
densiflori olim descripsi est L. Menziesii Ag. —Torr. et Gr. Fl. N. Amer. 1, p. 371,
quam etiam legit cl. Coulter inter San Miguel et Santa Barbara.
A few years later, Dr. Torrey (Pac. R. R. Report, 1853-4,
Botany 4: 81. 1856) gave us the following views:
Dood dennis DENSIFLORUS, Benth. 2 wars eoerenind hey PI. Ps pohi ig p. 303)
Agardh has founded his & t, which
i had described as L. densiflorus. All confusion ahint thee synony: suis may be
avoided, however, for the two species, L. densiflorus and L. Menziesii, Ag., cannot be
kept ee Both have white flowers (Agardh wrongly attributes yellow corollas
to his L. Menziesii, but = — eine the sppenrane' in dried specimens is cei? cor-
rect in thi
of Agardh’s L. densiflorus are weideuty not available pny a eens Metiastions. "De.
Bigelow’s specimens, however, correspond in this respect with L. Menziesii.
In Watson’s review (Proc. Amer. Acad. 8: 538. 1873) the
species is treated thus:
. L. pENsiFLorRUs, Benth. Much resembling the last [L. microcarpus], but more
sparingly villous with shorter hairs; bracts much shorter than the calyx, which is
smooth or short-pubescent, the upper lip often entire; petals yellow, or a
white or pink.—From the Sacramento Valley southward. It includes L.
ziesii, Agh.; and L. succulentus, Koch, is probably but a garden form.
None of the writers quoted above may be credited with having
done much field work in California, and it may be well to note here
a mere gleaning from the writings of a Californian, Dr. Kellogg
170 SMITH: STUDIES IN THE GENUS LUPINUS
of San Francisco, a man of extended field observations with a keen
appreciation for plant differences. In his comments upon the
plant named by him as L. lacteus (Proce. Cal. Acad. Sci. 5: 38.
1873), he says:
Admitting L. densiflorus to be the same as L. gems with variations, it
would then bring us toa “* dense subsessile spike’ . ith which to contend.
If these and many more varieties prove ie ately to run into one, it is not our ‘aie.
as the literature now stands, we are obliged, in self-defence, to set it apart, when
called upon for determinations.
Agardh’s descriptions are certainly more comprehensive than
Bentham’s, but the evidence is clear that the specimens selected
by him for his description of L. densiflorus were not typical of the
species as known to Bentham. The latter’s statement, however,
that L. Menziesti of Torrey & Gray is the same as his L. densiflorus
may be worthy of some careful weighing. I agree with both
Torrey and Watson that these two names do not represent two
separate species; but Torrey’s statement in respect to the “‘ yellow
corollas’’ in L. Menziesii is evidently erroneous, as attested by
many herbarium specimens and by my own and others’ field ob-
servations. I also appreciate Kellogg’s observation that ‘these
and many more varieties’’ exist, and feel, as he felt, the need of a
classification for their naming.
Very little can be gleaned from the published records of
Douglas’s sojourn in California. In his letter dated “ Nov. 23rd
1831,’’ written from ‘‘ Monterey, Upper California” and published
by Dr. Hooker (Comp. Bot. Mag. 2:149. 1836), he states that he
arrived at way opiate’ on iassscieicvesoe 22, 1830. About the end of
April, after vari of Monterey, he ‘ undertook
a journey southward, and reached Sisk Barbara, 34° 25’, in the
middle of May ... and returned late in June, by the same
route. . . . Shortly afterwards [he] started for San Francisco,
and proceeded to the North of that port.”’ His “last observation
was at 30° 45’; therefore in what is now Sonoma County, or
possibly, but not probably, in Solano or Yolo County.
Study of the collectors’ data with the numerous specimens
examined by me while preparing this paper permits of some specu-
lations that may be worthy of attention. One might thus con-
clude that Douglas’s specimens were secured before leaving
SMITH: STUDIES IN THE GENUS LUPINUS 171
Monterey for the north. Mature seed, however, might have been
taken by him almost anywhere from Santa Barbara to Sonoma
County. The ‘type’ of Agardh’s L. Menziesii probably came
from San Luis Obispo County, while the ‘“‘type’’ of L. densiflorus
Agardh almost certainly came from the vicinity of Monterey.
L. densiflorus Benth., however, was based upon specimens grown
from seed collected by Douglas. The seed characters given by
both Bentham and Lindley ought to be useful; but, unfortunately,
the seeds of most of the varieties herein recognized are unknown,
and none of those seen help much in the study from this angle.
Therefore, what may be accepted as reliable diagnostic seed char-
acters are yet to be worked out.
To sum up:—It seems to be an easy matter to establish the
identity of L. densiflorus Benth., when such is recognized as a
composite of numerous variable forms; but it is certainly a much
more difficult matter to determine which of the several forms is
properly to be designated as the typical plant. None of the vari-
eties herein considered are known to combine a low stem six or
seven inches high with pink-veined white petals and seeds smooth,
olive-green, and minutely dotted with black. For the present,
however, to serve as the typical form of the species, I am accepting
a Sonoma County plant whose inflorescence, flower-color, and
pubescence agree fairly well with Bentham’s description and the
plate in the Botanical Register, and whose period of blossoming
at Santa Rosa, June 2, as found by Heller, would have permitted
the collecting of seed by Douglas at the time of his visit to that
region. ;
The following key will probably be useful in determining most
of the material in the larger herbaria in terms of tHe varieties
recognized in this paper. Undoubtedly additional varieties are
yet to be added to the list.
Key to the varieties of Lupinus densifiorus
Calyx not bushy-villous below, the hairs 0.3-1.5 mm. long,
: bracteoles Sequenty. present.
Pub lly | less than I mm.
oe rather dense ‘and subappressed or
ppressed; lower calyx-lip aie sles
Beier and bent near base. DENSIFLORI.
Leaflets thin or thickish, not blackening in drying.
172 SMITH: STUDIES IN THE GENUS LUPINUS
Simple or branched well above the base, foliage
mostly ope
yx 7-8 mm. ge petals not yellow.
Banner 8 mm. wide; lower calyx-lip te
the two teeth slender and parallel. 6a. typical densiflorus.
Banner 6 mm. wide; lower calyx-lip with
broad sinus and a vestigial tooth. 6b. var. stenopetalus.
Flowers 16-19 mm. long; calyx 11-12 mm.
g; Danner 15-17 mm. long; petals
usually not yellow, often rose-tinted. 6c. var. perfistulosus.
Flowers 13-15 mm. long; calyx 9 (rarely
10) mm. long; banner 14 mm. long; petals
evidently yellow, often reddish- or pur-
plish-tinged at the edges. 6d. var. Menziesii.
Much branched at or near the base, the foliage
ngested.
alyx bracteolate; lower lip obtuse, the teeth
et and diverging; petals drying a dark
‘ose-purple. 6e. var. latilabrus.
ok spears lower lip -acutish, the teeth
small an ; petals drying a rich blue. 6/. var. Tracyi.
Leaflets almost ee blackening in dryi yx
bracteolate, lower lip acute and entire or palo
two- to three-toothed. 6g. var. glareosus.
Pubescence of stems and peduncles 1-1. 5 mm. long,
spreading or retrorsely spreading.
Lower calyx-lip nearly straight, inconspicuously or not
at all subsaccate and bent near base. LACTEI.
Acaulescent or nearly so, commonly simple with
contracted ra: a rare claw. 6h. var. lacteus.
Keel evidently curved; banner gradually con-
tracted into a poorly defined claw. 6l. var. vastiticola.
Stems and_ branches a the branches
usually all floriferou
Flowers 14-16 mm. das banner 15-16 mm
long; calyx conspicuously nities
spreading, about 1 mm. long. 67. var. sublanatus.
Flowers 12-14 mm. long; banner I1-14 mm. long.
Calyx 7 mm. long; banner rather abruptly
contracted into a well-defined claw; keel
nearly straight. 6j. var. McGregori.
Calyx 8 mm. long; banner gradually con-
tracted into a poorly defined claw; keel
evidently curved. : 6k. var. altus.
Lower calyx-lip usually distinctly subsaccate and bent
near base; flowers 13-19 mm. long, banner 13-16
mm. long, calyx 8-10 mm. long.
SMITH: STUDIES IN THE GENUS LUPINUS 173
Branches ascending to suberect; whorls five to sev-
“Fal,
Calyx commonly bracteolate, the hairs mostly
—0.5 mm. long.
Calyx 10 mm. long, lower lip acutish, the
lon
a not broader than long. 6m, var. versabilis.
alyx 8-9 mm. long, lower lip obtuse, the
gre feos than long. 6n. var. latidens.
Calyx rarely bracteolate, the hairs mostly tr. 5
mm. long. 60. var. Dudleyi.
— short and widely spreading; whorls two
0 five; flowers and fruits conspicuously secund. 6p. var. persecundus.
oe busyilous below, the hairs 1.5—-4 mm. long, ebrac-
PALUSTRES.
Core cates usually distinctly bent and inflated near
ase.
Flowers 13-14 mm. long; calyx 8 mm
Lower lip two-toothed; keel ne ade 30-60
6l. var. curvicarinus.
Lower lip three-toothed; keel stout; plants low. 6u. var. Reedii.
Flowers 15-17 mm. long; calyx 9-12 mm. long.
lants low, 8-15 cm. tall, the stems and branches
hort and usually widely spreading. 6s. var. crinitus.
0
Banner rounded or truncate at apex; lower lip
calyx two-toothed, the middle tooth ves-
ba or wanting. 6g. var. palustris.
Banner acute or distinctly contracted apically;
calyx three-toothed. 6r. var. stanfordianus.
Lower calyx-lip straight or nearly so, indistinctly or not at
bent and subsaccate near
Banner rounded or truncate at apex, 6 mm. wide;
calyx 8-9 mm. long, two-toothed; plant densely
villous. 6v. var. trichocalyx.
Banner mostly angled at apex.
Banner Ir mm. long, 5 mm. ; calyx 8 mm
long, lower lip three-toothed, upper lip less than
2mm. long. 6w. var. barbatissimus.
Banner 12-15 mm. long; calyx 10-13 mm. long,
upper lip usually 3-4 mm. long.
Banner gradually contracted into an undiffer-
entiated claw; wings naked or nearly so. 6y. var. scopulorum.
Banner con asia into a J cheaters broad
basal angle.
ein ts distinctly thoves weras four-)
toothed; aes conspicuously ciliate on
midrib below 6r. var. stanfordianus.
Lower lip ees. the Bese tooth
vestigial or wanting; leaflets not con-
spicuously ciliate. 6x. var. austrocollium.
174 SMITH: STUDIES IN THE GENUS LUPINUS
DENSIFLORI
6a. LUPINUS DENSIFLORUs Benth. (typical).
Pubescence less than 1 mm. long; verticils six to nine; calyx
about 8 mm. long, with short inconspicuous appressed pubescence,
lower lip barely 3 mm. wide, acute, the two
slender teeth with a very narrow sinus; petals
white, tinted or veined with violet. or rose,
banner 14 mm. long, 8 mm. wide, keel 12 mm.
long, relatively slender. Seed not seen.
CALIFORNIA. Sonoma County: Santa Rosa,
2 June, 1902, A. A. Heller 5630 (DS, US, PA,
NY, G). Napa County: Conn Valley, Napa
River Basin, 1 May, 1894, W. L. Jepson (UC).
Santa Clara County: Stanford University, 5
May, 1895, Cloud Rutter 173 (UC 308269, mid-
dle and right-hand specimens only). County
not given: Taylor Mountain, 20 Apr., 1898, M.
Fic. 18. Lupmsus - Baker (UC).
th. . :
DENSIFLORUS Bently 65, Lupinus densiflorus stenopetalus var. nov.
A. Ay Hed 5630 [
(US 416650). Fic. 19.]
Vexillo gracillimo, 13-14 mm. longo, 6 mm.
lato, ungue gracili prope 2 mm. lato, contracto; calyce 8 mm.
longo, labio inferiore bidentato, sinu lato dentis vestigio in-
structo, dentibus curvis; semina non vidi.
Petals white, more or less
rose-purple edged, the banner
unusually narrow, about 6 mm.
wide, with the claw only 2 mm.
wide; lower lip of the calyx ee / \
two-toothed, the teeth curved Se
and a vestigial median tooth tl
usually present. Seed not Fic. 19. Lupmvus pENSIFLORUS STENO-
seen. PETALUS C. P. Smith. A. A. Heller 7385
Cauirormia. Santa Clara ‘> 4%):
County: Los Gatos, 7 May, 1904, A. A. Heller 7385 (Typr, US
468403; type-duplicates, PA, NY, G). San Mateo County: San
Francisquito Canyon, 27 May, 1905, F. Grinnell (US).
Stenopetalus, Greek for “slender petal,” refers to the very
narrow banner.
I cm.
SMITH: STUDIES IN THE GENUS LUPINUS 175
6c. Lupinus densiflorus perfistulosus var. nov. [Fic. 20.]
Ramosus plerumque aliquantum super basin quamquam inter-
dum eramosus, 25-60 cm. altus; caule manifeste et ramis*pedun-
culisque plus minusve fistulosis, minute laxe pubescentibus vel
levibus; foliis patentibus; petiolis longissimis, 10-20 cm. longis;
foliolis 6-10, 20-40 mm. longis, oblanceolatis acutis arcuatis paulo
dense appresso-pubescentibus vel levibus subter: pedunculis
elongatis, plerumque folia excedentibus; verticillis plurimum 6-12
vel pluris, appositis vel distantibus subter; floribus pandentibus,
Fic. 20, LUPINUS DENSIFLORUS PERFISTULOSUS C. P. Smith. 1. Heller & Brown
5376 (US 413708); 2. W. H. Brewer 483 (US 321138); 3. Heller & Brown 5381 (US
413713).
I5-I9 mm. longis; pedicellis 3-4 mm. longis, prope I mm. latis;
bracteis humillimis 12-15 mm. longis, breve-pubescentibus vel
levibus; calyce 11-12 mm. longo, ebracteolato, appresso-pubescenti
vel levi, labio superiore 2-4 mm. longo, emarginato, inferiore mani-
feste inflato, prope 4 mm. lato, 2-dentato, sinu gracili vel latiore
dentis vestigio instructo; petalis diverse roseo- vel purpureo-
tinctis, interdum luteis; vexillo 15-17 mm. longo, 9 mm. lato, apice
rotundato paullatim contracto in unguem rigidum inflatum 5-6
m atum saepe retrorse dilatatum, labium superiore calycis
repellentem; alis 13-15 mm. longis, ad basin superne ciliatis, 7 mm.
latis, apicibus conspicue truncatis; carina 11-14 mm. longa, mani-
feste arcuata: legumina seminaque non vidi.
176 SMITH: STUDIES IN THE GENUS LUPINUS
The most robust and largest-flowered variety of the species,
unless equalled by var. palustris; often 60 cm. tall, stem usually
fistulose ; flowers 15-19 mm. long, banner 15-17 mm. long, rounded
at the apex, the claw inflated and stiff, sometimes pushing back
the upper calyx-lip, wings conspicuously truncate at the apex,
lower calyx-lip strongly subsaccate and bent downward; pods and
seeds, probably the largest of the. species.
CALIFORNIA. County not given: 1868-9, Kellogs & Harford
187 (Type, US 20645). Solano County: near Fairfield, 26 Apr.,
1902, Heller & Brown 5376 (US, G, NY, DS, PA); Gates Canyon,
mear Vacaville, 27 Apr., 1902, Heller & Brown 5381 (US, PA, G);
Vallejo, 11 May, 1874, E. L. Greene 171 (G); Wolfskill, 2-6 May,
1891, W. L. Jepson (UC). Contra Costa County: Muir Station,
July, 1904, Charlotta Case (UC). Yolo County: Rumsey, 7 May,
1903, C. F. Baker 3070 (US, G); Woodland, 16 May, 1893,
J. W. Blankinship (G). Butte County: “fields,” May, 1898,
Mrs. C. C. Bruce 2033 (NY). Lake County: Burns Valley, Apr.,
1902, Agnes M. Bowman 146 (DS). San Luis Obispo County:
San Luis Obispo, 27 Apr., 1861, W. H. Brewer 483, Geol. Sur.
Cal., (US); May, 1893, Mrs. Blochman (UC).
Two other specimens that perhaps should have been listed
. above are: Little Oak, Solano County, 2-6 May, 1891, W. L.
Jepson (UC), and Brighton, Sacramento County, Apr., 1884, K.
Brandegee (UC). These, and also the Contra Costa plant listed,
have a much denser pubescence than I have attributed to this
variety and would probably not be easily traced here by the use of
the key on pages 171-173. The last-mentioned plant is also a
pathological individual, having leafy branches and one raceme
developed from raceme-buds that normally should have produced
flowers. Seeds with Jepson’s “Little Oak’ specimen are dull
yellowish brown, finely and sparsely speckled with dark brown,
6x5 mm., decidedly flattened.
6d. Lupinus densiflorus Menziesii (Agardh) comb. nov. [FiG. 21.]
| Lupinus Menziesii Agardh, Syn. Gen. Lup. 2; 1836,
Lupinus Menziesii aurea Kell. Proc. Cal. Acad. Sci. $2. 16,
1873. | .
Agardh’s description will be found on page 168, included in my
SMITH: STUDIES IN THE GENUS LUPINUS 72
general discussion of L. densiflorus, so will not be repeated here.
The description of Kellogg’s variety is as follows:
Collected by Kellogg and Samuel Brannan, Jr., in Deer Valley, near Antioc
San Joaquin River, April 22d, 1869; chiefly differing from the accepted description of
the species—if we include also L. densiflorus—in the 2-toothed lower lip, relative
length of leaves, and the entire scarious tube of the calyx, etc
m fistulose, branching from near the axila]l summit, “leaflets about 10, one
third the length of the ada Scan above, pubescent beneath, stipules and bracts
ous, setaceously long acuminate, persistent; calyx tube scarious, upper lip 2-
toothed, deflexed, somewhat saccate; vexillum short, rounded outline, pubescent on
the back at the base, and along the claw above. Legumes hirsute, minute, 2-seeded.
Fic. 21. Lupmnus DENSIFLORUS MeENziEsiI (Agardh) C. P. Smith. 1. C. P.
Smith 1456; 2. A. A. Heller 7311 (US 468336); 3. A. A. Heller 7430 (US 468453); 4. C.
P. Smith 1484; 5. A. D. E. Elmer 4791 (US 665678); 6. H. A. Walker 2526 (US 669625).
As I have stated in the foregoing pages, I agree with others
that L. Menziesii is not specifically distinct from L. densiflorus;
but, on the other hand, I consider it a readily recognizable variety,
from which I do not feel justified in trying to separate Kellogg’s
var. aurea. The following description is supplementary to those
given pi others:
rs 14-15 mm. long; calyx rarely ( Heller'7439) bracteo-
late, pstbabann & 9 mm., rarely 10 mm. long, upper lip variable in
form, 2.5—3.5 mm. long, lower lip acute or Series as the tip and
entire or two- to three-toothed, the teeth mostly less than I mm.
long; petals bright yellow or paler, the apices and margins often
washed with some shade of reddish purple; banner 12-14 mm. long,
7-8 mm. ers rather abruptly contracted into a claw 4 mm. wide;
?
178 SMITH: STUDIES IN THE- GENUS LUPINUS
wings 11-12 mm. long, 6 mm. wide; pods conspicuously secund;
seeds about 4.5 x 4 mm. long, yellowish brown, speckled with darker
brown, the dark “lateral lines ’’ enclosing a paler area about the
hilum, plump or somewhat flattened.
CALIFORNIA. County not given: D. Douglas (T, G); T.
Bridges 59 (T, Col. U, NY, G, US). Sacramento County: Elk
Grove, May, 1882, E. R. Drew (UC). Contra Costa County:
Port Costa, 8 June, 1892, 7. S. Brandegee (DS, NY); Antioch,
3 May, 1893, A. Eastwood (G); Antioch, 5 May, 1907, K. Brande-
gee (UC 155195, as to specimen in center of sheet); near Point
Isabel, 8 July, 1911, H. A. Walker 2526 (US, G). Marin County:
Ross Landing, Aug., 1877, H. Edwards (NY). Alameda County:
between Mission San Jose and Livermiore, 13 Apr., 1904, A. A.
Heller 7311 (US, UC, PA, G); Mission San Jose, June, 1909, R. J.
Smith 7 (UC); Oakland, 1875, G. R. Vasey (US). Santa Clara
County: Evergreen, 11 Apr., 1893, J. B. Davy (UC); Gilroy, May,
1903, A. D. E. Elmer 4791 (US); Mt. Hamilton road, fourteen
miles from San Jose, 20 May, 1904, A. A. Heller 7439 (US);
Mt. Hamilton road, Halls Valley, 1950 ft. alt., 17 June, 1908,
C. P. Smith 1484 (CPS). Santa Cruz County: along Southern
Pacific Railway, east of Ellicott, 4 June, 1908, C. P. Smith 1456
(CPS). San Luis Obispo County: open hills near sea, 1 Mar.,
1883, Mrs. R. W. Summers (G); near boundary line of Santa
Barbara County, 8 May, 1896, A. Eastwood (G); Newhall Ranch,
8 May, 1900, J. H. Barber (G).
6e. Lupinus densiflorus latilabrus var. nov. [Fic. 22.]
Caule ad basin ramoso, ramulis foliisque congestis; foliolis 5-8,
10-25 mm. longis, 2-3 mm. latis; verticillis 3-5, floribus 13-15 mm.
seh
__Fic. 22. Lupinus DENSIFLORUS LATILABRUS C. P. Smith. E. Braunton 1081
(US 469834).
longis, pedicellis brevibus; calyce bracteolato 8 mm. longo, pilis
subappressis 0.5 mm. longis, labio inferiore 4 mm. lato apice obtuso
SMITH: STUDIES IN THE GENUS LUPINUS 179
2-dentato, dentibus latis brevibusque, sinu brevissimo; petalis ad
dimidia terminata roseo-purpureis, ad basin pallidis; vexillo prope
12 mm. longo, 8 mm. lato, apice rotundato; alis 11 mm. longis, vix
ciliatis ad basin superne; carina 9-10 mm. longa, paulo curvata;
leguminibus prope 14 mm. longis; semina matura non vidi.
Differs from var. Menziesii in the stem being branched at the
base, the branchlets and foliage congested; whorls few and pedicels
short and stout; calyx with bractlets, the lower lip relatively wide
at the blunt apex, the two teeth short and wide; keel not much
curved; terminal half of petals a dark rose purple when dried.
CALIFORNIA. Amador County: vicinity of Ione, 200-500 ft.
alt., June, 1904, E. Braunton 1081 (Tyre, US 469834; type-
duplicates, NY, UC).
Latilabrum is Latin for “ broad lip.”
6f. Lupinus densiflorus Tracyi var. nov. [Fic. 23.]
Caule ramosissimo ad basin foliis congestis; labio inferiore
calycis apice acuto bidentato, dentis sinuque gracilibus; petalis
pallido-carnosis, ad anthesin
atro-caeruleis cum siccatis; vex-
illo 12 mm. longo, 8 mm. lato,
ungue non claro; alis carinaque
latis: legumina seminaque non
vidi. |
Much branched at the base
with the foliage congested;
lower calyx-lip acute, the two : cm
teeth and their sinus narrow;
petals pale flesh-colored when
fresh (according to the label),
largely a rich blue in the pre- : ee
served specimen; banner 12 mm.
: Fic. 23. LUPINUS DENSIFLORUS TRACYI
long, 8 mm. wide, the claw not |p Smith. J. P. Tracy 3280 (UC
well defined; wings and keel 61546).
relatively broad.
CaLiForNIA. Humboldt County: gravel bar of Willow Creek,
Trinity River Valley, 4 July, 1911, J. P. Tracy 3280 (Type, UC
161546), labelled “ L. microcarpus.”
/
180 SMITH: STUDIES IN THE GENUS LUPINUS
6g. Lupinus densiflorus glareosus (Elmer) comb. nov. [F1G. 24.]
Lupinus glareosus Elmer, Bot. Gaz. 39:53. 1905.
I quote from Elmer’s description, as follows:
An almost acaulescent annual, 3s 5 dm. high, somewhat succulent though readily
outer ones curved upwards from the middle . . . leaflets 10 . . . sparsely pubescent
on both sides, soft and fleshy . . . the older ones with a very peculiar dead scarious
pany spikes cylindrical . . . 10-15 cm. long, erect and exceeding the leaves
wers. in whorls of 5, upon short pubescent pedicels . . . calyx . . . pubescent;
ae lip rather broad, 7 mm. long, obtuse, 3-nerved; upper one thinner, 3 mm. long,
notched; banner broadly elliptic, 12 mm. lon keel at least 12 mm. long, fal-
cate . . . with the margin of the aperture Seeks ciliate . . . seeds . . . smooth,
me ecu
Griffin’s Postoffice, Ventura nee California, July, 1902. Type specimen,
no. 3588, in Herb. Stanford University.
It is wholly confined to dry bteiels soil along water courses, hence its name.
Fic. LUPINUS DENSIFLORUS GLAREOSUS (Elmer) C. P. Smith. 1. A. D. E.
Elmer ar (US 465791); 2. Abrams & McGregor 262 (US 613146).
I consider this one of the most extreme variations of L. densi-
florus, and I would have upheld it as a species but for the evidence
presented by Abrams & McGregor’s collection listed below. The
following characters should also be mentioned.
Foliage blackening in drying; verticils crowded or distinct, six
to twelve or fewer; flowers spreading in anthesis, becoming secund
later, 13-14 mm. ‘beng: calyx bracteolate, 6.5-8 mm. long, very
shortly and almost sparsely appressed-pubescent, upper lip a little
an I mm. long and cleft, or 2-3 mm. long and emarginate,
lower lip entire and acute or minutely fer ba about 3 mm.
SMITH: STUDIES IN THE GENUS LUPINUS 181
wide; petals (according to Dr. Hall’s labels) light blue, the banner
with a white center; banner 12-14 mm. long, 8-9 mm. wide, ovate
to ‘‘ broadly elliptic,’’ obtusely angled, truncate, or rounded at the
apex, the basal inflation much thickened and stiff; seeds pale
yellowish brown, sparsely mottled with dark brown, the consider-
ably paler hilum area marked off by prominent V-shaped lines.
CALIFORNIA. Ventura County: Griffin’s Postoffice, June,
1902, A. D. E. Elmer 3588 (DS, US, NY); Griffin, Mt. Pinos, 20
June, 1905, H. M. Hall 6334 (G, UC); near Frazier borax mine,
Mt. Pinos, 12-14 June, 1908, Abrams & McGregor 262 (DS, US).
While with his description Elmer gives the date of collection
of his number 3588 as July, 1902, his labels give June as the month.
His number 4006, however, from the same locality, is labeled as
having been taken in July; but the latter is a very different plant,
and has been treated by me elsewhere as L. subvexus nigrescens
(Bull. Torrey Club 45:13. 1918).
LACTEI
6h. Lupinus densiflorus lacteus (Kell.) comb. nov. [Fic. 25,]
Lupinus lacteus Kell. Proc. Cal. Acad. Sci. 5: 37. 1873.
Lupinus arenicola Heller, Muhlenbergia 2: 75. 1905.
Kellogg’s description of L. lacteus is, in part, as follows:
Stem annual, fistulous, the elongated central peduncle from a mere depressed
crown, mostly solitary, spike 4 to 8 inches long, lateral radicle branches 2 to 6 inches
long, with secondary clusters of leaves and (when present) shorter spikes, soft pubes-
cent throughout, with white hairs. Leaves mostly clustered at the base . . . leaf-
lets . . . silky-pubescent beneath. . . . Flowers large, white, somewhat distant,
aber . . . calyx ebracteolate, hirsute . . . upper lip 2-cleft (rarely entire)
nc ¥ the length of the lower lip, lower lip straight, herbaceous, 2-toothed .. .
um .. . ciliate at the Ry prs juncture of the claw . . er, wings and
‘acs ora equal, wings oblon . . Margins ciliate at the ices or origin of the claw;
keel ciliate at the upper Ay margin toward the base.
In habit and general appearance this species resembles L. brevicaulis, but is
rather more robust, the flowers sic larger and not “‘deep blue,’ but quite white,
- Itisclosely allied to L. Menziesii. . . . Specimens collected by Mr. S. Brannan.
S on Oak Creek hillsides, Kern County, 14 miles from Tejon Pass
Extracts from Heller’s description of L. arenicola follow:
Branches several, ascending, commonly only the middle one floriferous .
flowers in 1 to 3 whorls, merely ascending, whitish or rose color, 1.5 cm. long . .
calyx with broad acutish lower lip 7 mm. long, 5 mm. wide at base, the apex sniectiake
2-toothed; upper lip ovate . . . the apex 2-toothed, the teeth 1 mm. long, slightly
spreading; banner turned back ... keel . . . little curved, ciliate on the lower
half, . eseile large, 4 mm. in Bisisiee. white or whitish.
182 SMITH: STUDIES IN THE GENUS LUPINUS
type is no. 7609, collected April 7, 1905, near the first crossing of the creek
west ei Caliente, Kern county, California, growing in sandy soil on steep banks near
the railroad. It also occurs on the bluffs of Kern River above Bakersfield.
Heller, writing some five years later (Muhlenbergia 6: 70.
1910), says:
It gives me great pleasure to reinstate this long unrecognized species [L. lacteus],
even at the expense of one of my own names. The species was barely published be-
fore it was suppressed by Watson in Proc. Am. Acad. 8: 542, November, 1873. He
there puts it down as a synonym of L. densiflorus, but evidently never saw the plant
itself, for he nae not mention it in the list of specimens
mined. During the course of five years he came
another conclusion, for in the Bibliographical In-
ce published in 1878, it is said to be the same as L.
microcarpus, a less satisfactory disposition than the
first, for it is clearly more related to L. densiflorus
than it is to the narrow-flowered L. microcar pus.
pe of L. iene shows a bractlet in the
sinus Pel the calyx lobes, whereas Kellogg says
it is not prese
pO as CALIFORNIA. Kern County: near Ca-
liente, A. A. Heller 7609 (US, G, NY,
PA); Bakersfield, 26 Apr—30 May, 1896,
J. B. Davy 1716 (G). Ventura County:
Ojai and vicinity, 25 Apr., 1866, S. F.
Fic. 25. Lupinus DEN- Peckham (US); Ojai Valley, 18 Apr., 1896, ©
weeds ages mage Aa 3 Hubby 34 (UC). Tulare County:
5301 (US 328676). North Tule River, Nisy, thob. C. A.
Purpus 1733 (UC); North Tule River,
May-Oct., 1896, C. A. Purpus 5694 (UC); “Southeastern Cal.”:
hillsides, Erskin Creek, Apr.Sept., 1897, C. A. Purpus 5301
(US, G, UC). Riverside County: near San Jacinto, 9 Mar.,
1898, J. B. Leiberg 3134 (US). San Diego County: Coyote Can-
yon (Colorado Desert), Apr., 1902, H. M. Hall 2853 (UC).
While not recognizing L. lacteus as of specific rank, I accept it
as a well-marked variety of L. densiflorus. As to the calyx brac-
teoles, the discrepancy between Kellogg’s and Heller’s descrip-
tions is not a critical point, for my studies have shown me that
these bractlets may or may not be present in different flowers of a
particular raceme. I find them to be usually absent in the speci-
mens listed above. The pubescence in this variety is quite var-
iable, some specimens being nearly smooth.
SMITH: STUDIES IN THE GENUS LUPINUS 183
Kellogg’s Napa specimen in the Gray Herbarium (Napa, dry
barren hillsides, 1 Apr., 1870), though distributed by Kellogg as
L. lacteus, cannot be referred here. It is possibly a diminutive
specimen of var. perfistulosus of this paper.
6i. Lupinus densiflorus sublanatus var. nov. [FiG. 26.|
Erectus, eramosus vel ramosus, 10-40 cm, altus, caule et cetera
dense conspicueque villoso, pilis plusculum 1 mm. longis: bracteis
calycibusque saepe sublanatis: verticillis 5—9; floribus pandentibus
ad anthesin et postea: calyce bracteolato vel ebracteolato, labio
superiore 3-4 mm. longo, gracile, acuto integroque vel bifido, in-
feriore recto vel ad basin paulo inflato 2-dentato, dentibus 1 mm.
longis parallelis, sinu cum vestigio vel sine eo: corolla alba vel
tincta; vexillo 13-15 mm. longo, 7-9 mm. lato, apice rotundato vel
interdum obtuse angulato; alis superne ad basin ciliatis, interdum
subter paulo; carina recta vel curva: fructus seminaque non vidi.
CALIFORNIA. Kern County: Water Canyon, Tehachapi Moun-
tains, 26 June, 1908, Abrams & McGregor 485 (Tyrer, DS 9584;
type-duplicates, US, NY, G); Tehachapi, June, 1911, K. Brande-
gee (UC); vicinity of Fort Tejon, 16-
1 cm.
T
17 June, 1908, Abrams & McGregor
jio (DS, US, NY, G); vicinity of Fort
Tejon, 1857-8, L. J. X. de Vasey 21
(US); Tehachapi, 1891, Coville & Fun-
ston 1119 (F. W. Koch) (US); Bakers-
field, 26 Apr.—30 May, 1806, J. B. 9
Fic. 26. LUPINUS DENSI-
Davy 1708 (UC); Keene Station, 1 May,
1905, A. A. Heller 7798 (DS, US, PA,
G, NY); vicinity of Havilah, 15 June, [°° i 6
1905, F. Grinnell 295 (US); Johnson 48, (ps 9584).
Canyon, Walker Basin, 3 June, 1905,
F. Grinnell 51 (US); Caliente Creek, 1 June, 1905, F. Grinnell
4 (US); near Caliente, May, 1909, K. Brandegee (UC).
Probably most closely related to var. lacteus, from which it
would seem to be easily distinguished by the caulescent habit and
the numerous racemes. It is noteworthy, however, that most of
the specimens cited above were collected in June, while the types
of both L. lacteus and L. arenicola were taken in April. It would
therefore be an interesting study for some student to locate, if
possible, the type station of Heller’s L. arenicola and Mrs. Brande-
a
184 SMITH: STUDIES IN THE GENUS LUPINUS
gee’s station at Tehachapi, say in April, and follow both through
until all the seed is matured and the season’s growth is evidently
completed. Cultural studies would also be of interest; but great
care would be very necessary in the selection and identification of
the seed to be used.
Sublanatus, “almost woolly,” refers to the general pubescence
of the typical phase of the variety, as represented by the type-
collection and Mrs. Brandegee’s Tehachapi plant. However, the
specimens referred here show quite a range of variation in density
of pubescence, the “least lanate’’ extreme being well represented
by Abrams & McGregor’s No. 310. This collection I had selected
as the type of an additional variety, but later decided that this
variation was not supported by other characters and, evidently,
was in itself too inconstant to be reliable. In the upper San Joa-
quin Valley, as at Bakersfield, where the two varieties overlap,
the nearly straight lower calyx-lip with retrorse-spreading hairs
should readily distinguish var. sublanatus from the related var.
versabilis, in which the lower lip is decidedly subsaccate and pubes-
cent with subappressed hairs about 0.5 mm. long.
Some other specimens, not placed elsewhere, which may have
some affinity to the above are: Nacimiento Canyon, San Luis
Obispo County, 8 May, 1900, J. H. Barber (UC); Knights Ferry,
Stanislaus County, 9 Apr., 1895, F. W. Bancroft (UC); Linden,
San Joaquin County, May, 1896, F. W. Gunnison (UC).
6j. Lupinus densiflorus McGregori var. nov. [Fic. 27.]
Erectus, super basi prope 10 cm. ramosus, ramis paucis suberec-
tis, caule ramisque gracilibus nequaquam fistulosis, laxe pubes-
centibus, pilis pandentibus 1-1.5 mm. longis: foliolis prope 20 mm.
longis, cuneatis, apice rotundatis vel emarginatis pallido-viridis,
pilis brevibus albescentibus subter: racemis brevibus, verticillis
2-3, distantibus; floribus parvis, 12-13 mm. longis, pandentibus ad
anthesin; pedicellis 1.5-2 mm. longis, gracilibus; bracteis 2-5 mm.
longis, i picui lyce ebracteolato, 7mm. longo, villoso inferne,
pilis prope 1 mm. longis, pandentibus vel retrorso-pandentibus;
labio superiore 2 mm. longo, inferiore obtuso, paulo plus 2 mm. lato,
2-dentato, dentibus prope 0.5 mm. longis: corolla alba vel pallida;
vexillo 11 mm. longo, 6 mm. lato, apice rotundato paulo abrupte
contracto in unguem gracilem 2 mm. latum; alis 10 mm. longis,
4 mm. latis, non ciliatis; carina 10 mm. longa, recta: leguminibus
15 mm. longis, pallido-stramineis; seminibus 4 mm. longis, 3.5 mm.
SMiTH: STUDIES IN THE GENUS LUPINUS 185
latis, pinguibus, paulo acute angulatis, dense atro-cinereis macu-
latis.
Erect, branched about 10 cm. above the base, the branches
nearly erect, slender, and somewhat fistulose; leaflets about 20
mm. long, cuneate, the apex rounded or
emarginate, pale green, whitened below with
conspicuous short subappressed hairs; whorls
two or three, flowers small, 12-13 mm. long,
spreading in anthesis; calyx only 7 mm.
long, the hairs spreading or retrorse-spread-
ing and about I mm. long, lower-lip two-
toothed; banner 11 mm. long, 6 mm. wide,
wings not ciliate, keel straight, all the petals er
white or pale: pods pale straw-colored; seeds
about 4 x 3.5 mm., plump, somewhat acutely ane
angled, densely spotted with dark gray.
CALIFORNIA. Los Angeles County: Rock Fic. 27. Lup
Creek, desert slope of the San Gabriel Cath ce agree
Mountains, 2-4 July, 1908, Abrams & Mc- gyyams oe MiGriiey Ga
Gregor 551 (Type, DS 9585; type-dupli- (Ds 9585).
cates, US, NY, G).
Apparently this form has a number of unique characters.
Specimens collected in late May or early June would be interesting.
Named si one of the collectors, my friend Ernest Alexander
McGreg
6k. Lupinus densiflorus altus var. nov. [Fic. 28.]
Ramosus, 30-60 cm. latus, ramis pedunculisque elongatis,
caulibus ramisque laxe villosis et pedunculis petiolisque dense
villosis, pilis 1 mm. longis: petiolis 10-20 cm. longis; verticillis 6-7,
distinctis; floribus 14 mm. longis, pandentibus; calyce ebracteolato,
8 mm. longo, subter dense pandento-pubescenti, labio geile
recto, paulo inflato, 3 mm. lato, 2-dentato, dentibus I mm
sinu dentis vestigio plus minusve instructo: petalis albis; vexillo
I3 mm. longo, 7 mm. lato, apice rotundato, paullatim contracto in
unguem; alis 12 mm. longis, dense ciliatis ad basin superne; carina
curvata: legumina seminaque non vidi.
Branched above the base, 30-40 cm. tall, the branches and
peduncles elongated, branches loosely villous, the peduncles and
petioles densely villous with hairs 1 mm. long; whorls six or seven,
186 SMITH: STUDIES {N THE GENUS LUPINUS
distinct; flowers spreading, 14 mm. long; calyx ebracteolate,
densely pubescent below with spreading hairs, the lower lip
straight, but slightly inflated, two-toothed,
with or without the interstitial vestigial
tooth; petals white, the banner rounded at
the apex and gradually contracted into a
poorly defined claw, wings densely ciliate
at the free edges above, keel evidently
curved; pods and seeds not seen.
CaLiForNIA. Los Angeles County:
Manzana, Antelope Valley, 9-24 May,
1896, J. B. Davy 2505 (Type, UC 130329).
The collector’s label gives generic de-
termination only; but an annotation label
ee calls the specimen an “undescribed sp.”
6/. Lupinus densiflorus vastiticola var.
Fic. 28. Lupinus pDEN-
SIFLORUS ALTUS C.P.S
J.B.Davy 2505(UC 130329). Acaulescens, eramosus vel saeaeuie
18 cm. altus: foliolis 10-20 mm. longis,
4-5 mm. latis, oblanceolatis, acutis, laxe villosis superne: pedunc-
ulis i apo dense villosis, subrufis rl
nov. [FIG. 29.]
colore corollae incerto; vexillo apice rotun-
dato, paullatim contracto in unguem, 11 ie
mm. longo, 5-6 mm. lato; alis prope 10 mm.
longis, 5mm. eis metey ad basin superne;
carina curvata, xima II mm. longa, 4
mm. lata, acumine ne brev latoque: legumina ‘Fic. 29. Lupinus pEN-
et semina non vi SIFLORUS VASTITICOLA C,
P. Smith. C.G. Pringle
Simple or eee at the base, 9-18 cm. (US 20638).
tall; peduncles short, reddish or purplish,
densely villous; whorls about four or five, distant; flowers spread-
~
SMITH: STUDIES IN THE GENUS LUPINUS 187
ing, about 13 mm. long; calyx ebracteolate, 8 mm. long, its hairs
spreading and about 1 mm. long, lower lip somewhat inflated
near the base, 3 mm. wide, two-toothed, with or without a ves-
tigial median tooth; color of the corolla uncertain; banner rounded
at the apex, gradually contracted into a poorly defined claw;
wings ciliate at the base above; keel curved, comparatively large,
the point short and wide; fruit and seeds not seed.
Vastiticola is Latin for ‘waste inhabiting.”
CALIFORNIA. ‘Mojave Desert,” 12 May, 1882, C. G. Pringle
(Type, US 20638; type-duplicate, PA).
VERSABILES
6m. Lupinus densiflorus versabilis var. nov. [F1G. 30.]
Eramosus vel ramosus, 20-40 cm. altus, plus minusve fistulosus
cum super basi ramosus, pubescens, pilis brevibus, paulo densis,
plurimum retrorso-pandentibus, vix I mm. long; petiolis laxe
mm. longis, subrectis vel arcuatis, oblanceolatis, acutis, subter
specie levibus, quamquam
ibus, medio-costis margini-
busque ciliatis; pedunculis
verticillis 4-10 appositis vel
distantibus; floribus ad an-
bus, 14-16 mm. longis; pedi- 7
mm. longis, marginibus cili-
atis; calyce frequenter brac-
teolato, 9-Io mm. longo, ali-
quantum dense pubescente,
pilis brevibus laxe appressis,
riore diverso, 2—3 mm. longo, C. P. Smith. 1. A.A. Heller 8174 (DS ere
inferiore inflato 2-3 mm.lato, 2. A. A. Heller 7638 (CPS).
mm. longis; corolla alba, diverse roseo- vel violaceo-tincta; vexillo
apice rotundato, 14-16 mm. longo, 7-8 mm. lato, paulatim con-
pubescentibus vel sublevibus, 6-18 mm. longis; foliolis 6-10, 20-35
minute appresso-pubescent-
paulo brevibus vel elongatis,
thesin et postea pandenti-
cellis prope 2 mm. longis,
paulo gracilibus; Dien
verticilli humillimi vix 10
0.5 mm. longis, labio supe- Fic. 30. Luprnus DENSIFLORUS VERSABIL.
2-dentato, dentibus 0.5-I
tracto ungue rigido, 4-5 mm. lato; alis 12-14 mm. longis, prope 6
188 SMITH: STUDIES IN THE GENUS LUPINUS
mm. latis, ad basin subter manifeste ciliatis; carina subrecta, vel
paulo curvata II-I2 mm
This variety of the San Saar in Valley and adjacent foothills
of the Sierras differs from var. Dudleyi in the shorter pubescence
of the calyx, the usually more richly tinted corollas, and the de-
cided prevalence of interstitial bracteoles. While seeds of var.
Dudleyi have not yet been seen by me, I have no doubt but that
such will be found to differ considerably from those of var. ver-
sabilis. Seeds with the San Joaquin County plant, cited below,
are milky-white, unmarked, rather smooth and plump, about
5x4 mm.
CALIFORNIA. Fresno County: Fresno, 13 Apr., 1906, A. A.
Heller 8174 (TyPE, DS 9593; type-duplicate, US). Kern County:
near Bakersfield, 10 Apr., 1905, A. A. Heller 7638 (DS, US,
CPS). Madera County: Madera, 22 Apr., 1897, W. A. Setchell
(UC); North Fork and vicinity, 30 May-8 June, 1903, D. Griffith
4622 (US). Amador County: Jackson Gate, May, 1892, G.
Hansen 1314 (DS, UC); Stony Creek, 26 May, 1896, G. Hansen
1674 (US). San Joaquin County: Tracy, to Apr., 1900, B. Cobb
(UC).
Versabilis is Latin for ‘‘ changeable” and refers to the variation
in the tinting of the petals.
6n. Lupinus densiflorus latidens var. nov. [Fic. 31.]
Habitu notisque var. versabilis, a ipso differt pilis pandentibus
plurimum laxe vix 0.3-0.5 mm. longis; labio calycis inferiore
interdum integro vel 2—3-dentato, dentibus vegans latioribus
quam longis; leguminibus secundis prope 15 mm. long
Differing from var. versabilis in the somewhat eet pubes-
cence, in the paler tinting of the petals, as a rule, and the lower
calyx-lip being entire or with very wide short teeth. It seems to
be local in the general vicinity of San Bernardino.
_ CALIFORNIA. San Bernardino County: vicinity of San Ber-
nardino, 20 May, 1896, S. B. Parish 4165 (Type, US 279032;
type-duplicates, NY, G, UC); Redlands, May, 1890, S. B. Parish
(US); vicinity of San Bernardino, 11 May, 1901, S. B. Parish
4783 (DS). Riverside County: Temicula, 1880, G. R. Vasey 96
(US). County not given: J. M. Biglow (Whipple’s Exploration
of the 35th — 1853 (US); foothills, May, 1887, S. B.
a AS.
SMITH: STUDIES IN THE GENUS LUPINUS 189
(OS0
Dara
UPINUS DENSIFLORUS LATIDENS C. P. Smith. 1. S. B. Parish 4165
(US yes: as a M. Bigelow (US 20644); 3. S. B. Parish (US 480801).
60. Lupinus densiflorus Dudleyi var. nov. [Fic. 32.]
Ramosus, prope basin manifeste pubescens, pilis retrorso-pan-
dentibus prope 0.5 mm. longis; racemis pluribus, verticillis 5-10
vel pluribus, plus minusve apposi-
tis; floribus 15-16 mm. longis;
bracteis humilioribus, calyces ex-
cedentibus longitudine; calyce in-
terdum -bracteolato, plerumque
ebracteolato, 10 mm. longo, subter
laxe villoso, pilis pandentibus, 0.5-
1.5 mm. longis, labio superiore 2.5-3
mm. longo, inferiore plerumque in-
p
sinu dentis vestigio instructo, den-
tibus 0.75-1.5 mm. longis; petalis
albis vel pallido-roseis; vexillo apice
rotundato, 14 mm. longo, prope 8
m. lato, paulo abrupte contracto
eusue 4mm. lato; alis 12-13 mm.
longis, apice prope 7 mm. latis, ad
basin oe ciliatis; carina 10-II
mm. longa, acumine subvexo; le-
Pf
See
Fic.32. LUPINUS DENSIFLORUS Dup-
Levi C.-P. Smith. 1. C. P.. Smith
1442, C. F. Baker 850 (US 440513);
zinta secundis, 15 mm. longis: , 4. p. £. Elmer 4847 (US 665734).
semina non vidi.
The rounded banner and the short ge spreading pubes-
cence distinguish this quite local variety of the San Francisco
190 SMITH: STUDIES IN THE GENUS LUPINUS
peninsula, etc. The flowers are white or tinted with rose-pink,
commonly large and showy. The pods are conspicuously secund.
CALIFORNIA. San Mateo County: serpentine rock west of
San Mateo, 26 Mar., 1894, W. R. Dudley (Typr, DS); Pilarcitos
Lake and Canyon, 21-23 June, 1893, J. B. Davy 1151 (UC);
Crystal Springs Lake, 1 May, 1902, C. F. Baker 850 (US); near
San Mateo on the Half Moon Bay road, 23 May, 1907, A. A.
Heller 8561 (DS, US); Lake San Andreas, June, 1903, A. D. E.
Elmer 4847 (US, NY); San Mateo Canyon, 31 May, 1908, C. P.
Smith 1442. Marin County: without definite locality, 4 June,
1895, I. Tidestrom (UC).
6p. Lupinus densiflorus persecundus var. nov. [FicG. 33.]
Humilis, 5-15 cm. altus, ramis brevis subpandentibus, pedun-
culis eorum plerumque plus pandentibus; foliis congestis, viridis,
peculiaribus; petiolis 3-8 cm. longis, pilis 1 mm. longis, laxe panden-
bcm.
Fic. 33. LUPINUS DENSIFLORUS PERSECUNDUs C. P. Smith. M.E. Jones 3320 (US).
tibus; foliolis6—8, prope 20 mm. longis, 4-6 mm, latis, acutis , superne
sublevibus, inferne laxe pubescentibus: pedunculis foliis aequan-
tibus vel ea excedentibus, laxe pubescentibus, racemis aliquantum
deflexis; verticillis 1-4, distinctissimis, floribus manifeste secundis,
prope 16 mm. longis; pedicellis 1.5-2 mm. longis; calyce ebrac-
teolato, 9 mm. longo, 4-5 mm. lato, laxe pandentibus pubescente,
pilis vix 1 mm. longis, labio superiore 3 mm. longo, emarginato,
inferiore 3-dentato, dento medio lateralibus minore interdum
vestigio; petalis atro-purpureis, praesertim ad dimidia externa;
vexillo 12 mm. longo, 10 mm. lato, apice rotundato, abrupte con-
SMITH: STUDIES IN THE GENUS LUPINUS 191
tracto ungue inflato 6 mm. lato superne, 4 mm. lato basi; alis 13
mm. longis, ciliatis ad basin superne; carina 12 mm. longa, curvatis-
sima, acumine longissimo gracile; fructibus manifeste secundis,
specie usitatibus: semina non vidi.
Low, 5-15 cm. high, the branches more or less spreading with
usually more widely spreading peduncles; whorls one to four,
remote, the flowers secund and largely dark purple in color, about
16 mm. long; calyx ebracteolate, 9 mm. long, its pubescence spread-
ing, about I mm. long, the lower lip mainly three-toothed; banner
12x10 mm., abruptly contracted into an inflated claw which is
6 mm. wide above; keel 12 mm. long, much curved and with a
long slender apex; fruits strongly secund, otherwise apparently
not distinctive; seed not seen.
CALIFORNIA. Sonoma County: Duncans Mills, 17 July, 1882,
M. E. Jones 3329 (Type, US; type-duplicates, NY, Utah Agr.
Col.); cliffs near Bodega Bay, 30 June, 1907, K. Brandegee (UC).
Marin County: Dillons, Dec., 1898, R. E. Gibbs (UC).
Persecundus is used in the sense of “strongly secund”’ and
refers to. the position of the flowers and fruits on the rachis.
The type collection is labelled L. microcarpus.
PALUSTRES
6g. Lupinus densiflorus palustris (Kell.) comb. nov. [Fic. 34.]
Lupinus densiflorus Agardh, Syn. Gen. Lup. 1835.
Lupinus palustris Kell. Proc. Cal. Acad. Sci. 5: 16. 1873.
Agardh’s description, in part, will be found on page 169, with
the general discussion of L. densiflorus Benth.
Kellogg’s description of L. palustris, somewhat abridged,
follows:
Stem stout, annual, fistu . «Ofte - branchin, . . beyond the main
axis and its elongated cess pars tae on. silky, pub, or subglabrous,
with Arca very minute villi... leaflets . glabrous above, subpubescent
12 inches; flowers are, viotetbie, eet —— or verging to
spik
ce mages Ssaslnicinia or verticellat ‘ u yx bracteolate
or ebracteolate hirsute . . . slightly saceate; upper lip 2-toothed, ceil herbaceous
oattine; petals equal. Legumes very appressed, (silvery?) hirsute, compressed, an
inch or more in length, about 8- ed.
Collected by Kellogg and Bloomer on the San Joaquin River, April 7th, -_
Differs from Mensziesii—a 2-seeded species,—whereas this has 8 or more; also on
var. (deep purple-blue flower) has very distinct bracteoles.
192 SMITH: STUDIES IN THE GENUS LUPINUS
This description has been a puzzle to me since I first
studied it in 1910, and no solution to the matter suggested itself
until recently, when it was my pleasure to examine, at the Gray
Herbarium, one of Kellogg’s specimens distributed by himself as
his L. palustris, and collected
5mm. in April, 1869. This is in flower
only, but it is the same as L.
densiflorus Agardh, and agrees
with Kellogg’s description, as
far as same applies at all to a
Platycarpos species. It may be
Fic. 34. Lupinus pENsIFLorus pal- that Kellogg’s type-collection
ustTrRis (Kell.) C. P. Smith. <A. Kellogg,
Apr., 1869 (G), not dissected.
included some very peculiar
pathological specimens; but I
am more inclined to believe that he actually confused flowering
specimens of this variety of L. densiflorus with fruiting specimens
of a quite different species, perhaps the robust annual now
known as L. affinis Agardh.
However, as attested by Bentham himself, Agardh’s L. den-
siflorus can not be accepted as the typical form of the species, and
it seems to me that this easily recognizable form, as a variety,
should bear Kellogg’s name.
Another specimen in the Gray Herbarium calls for special
attention, said specimen being the one labelled ‘‘L. Menziesii,
var. aurea, Kellogg and Brannan. Jr., San Joaquin River, 22 April
1870." As may be seen by referring to the original description
(see page 177), Kellogg’s type is said to have been taken at
“Deer Valley, near Antioch, San Joaquin River, April 22, 1869.”
One might easily assume that the discrepancy in date, 1869 vs.
1870, is a mere clerical error, overlooked by the person who wrote
out the label in question. The specimen, however, according to
the descriptions, is not Kellogg’s var. aurea, but is the same as his
L. palustris. As to the locality, Antioch and vicinity, my cita-
tions will show that Davy also collected var. palustris at Antioch
in 1895, and that Miss Eastwood and Mrs. Brandegee each col-
lected L. Menziesii (probably var. aurea) at Antioch in 1893 and
1907, respectively. Still a third variety of L. densiflorus and also
me _ the typical L. subvexus have been taken at Antioch. I accordingly
SMITH: STUDIES IN THE GENUS LUPINUS 193
wonder if this particular specimen was not labelled on the kasis of
the collectors and part of the date (22 April), rather than with
careful regard to the description of var. aurea.
Var. palustris shares with typical L. subvexus a pronounced
tendency toward a covering of long loose hairs, those of the calyx,
especially, being 2-4 mm. long. The “palustris varieties” of
L. densiflorus, however, may be distinguished from the varieties
of L. subvexus by the position of the flowers soon after anthesis,
the flowers soon turning up in L. subvexus, without becoming
secund, as is so frequent in many varieties of L. densiflorus as the
fruits develop. Some of the extreme ‘palustris variations’’ are
below designated as separate varieties, most of which have com-
monly been labelled as L. microcarpus. One of these, however,
has been named, but not described, by Miss Eastwood.
_To supplement Kellogg’s and Agardh’s descriptions, I would
mention the following diagnostic points:
Flowers spreading in anthesis and later; calyx usually ebracteo-
late, 12-13 mm. long, bushy-villous below, especially on the cup,
the hairs 2-4 mm. long, upper lip various but usually large, about
3 mm. long, lower lip inflated near base, 3-4 mm. wide, two-
toothed, with or without an interstitial vestige; petals white or
tinted with pink or purple; banner rounded apically, 15-17 mm.
long, 8 mm. wide, rather abruptly contracted at base, the basal
inflation more or less’saccate, pressing the upper lip backward;
wings 13-15 mm. long, sometimes ciliate at the free edges above;
keel 11-13 mm. long, slightly to considerably arcuate; pods 14-
16 mm. long; seeds probably dull yellow, sparsely speckled with
dark brown, about 5 x 4 mm. (DS 9588).
CALIFORNIA. County not given: San Joaquin River, Apr.,
1869, A. Kellogg (G); San Joaquin River, 22 Apr.,. 1870, A.
Kellogg & S. Brannan, Jr. (G); no data (DS 9588). Contra Costa
County: Antioch, Apr., 1895, J. B. Davy (UC). Alameda County:
vicinity of Berkeley, May-June, 1906, H. A. Walker 173 (UC);
Berkeley, June, 1894, W. S. Blasdale (US). Santa Cruz County:
Watsonville coast, 13 Apr., 1902, C. F. Baker 3009 (US, NY, G).
Monterey County: Monterey, T. Hartweg 1692 (G); Monterey,
J. Gi Cooper (US); dry ravine, Monterey, May, 1850, C. C. Parry
(T).
194 SMITH: STUDIES IN THE GENUS LUPINUS
6r. Lupinus densiflorus stanfordianus var. nov. [FIG. 35.]
A var. palustri differt labio inferiore calycis 3-dentato, dente
medio plerumque amplo; corolla pallido-luteolo-alba vel pallido-
rosea; vexillo apice acuto vel abrupte contracto ungue rigido;
seminibus 4 x 5 mm., asperibus, atro-fulvis maculis non claris.
Bes
oS
—— =
Fic. 35. LUPINUS DENSIFLORUS STANFORDIANUS C. P. Smith. 1. C. P. Smith
791; 2. A. D. E. Elmer 2190 (US 655041); 3. W. F. Wight 140 (US 467702).
Lower lip of the calyx three-toothed, the median tooth being
well-developed; petals creamy or rose-tinted, the banner acute or
abruptly contracted at the apex, the claw stiff; seeds rough, dark
brown with indistinct specks; leaflets conspicuously ciliate on the
midribs and margins.
CALIFORNIA. Santa Clara County: Stanford University,
base of foothills, 9 Apr., 1905, C. P. Smith’791 (Type, CPS);
16 Mar., 1900, W. F. Wight 140 (US); Stanford University, Apr.,
1900, W. A. Atkinson (DS); west of road near Rocy House, 6 Mar.,
1900, W. A. Atkinson (DS); near Stanford University Apr., 1900,
A.D. E. Elmer 2190 (US); foothills, Stanford University, 8 Apr.-
4 May, 1902, C. F. Baker 475 (US, DS, NY, G). San Mateo
County: Stanford University, by road from county bridge to
Basaltic Rocks, 10 May, 1897, W. R. Dudley (DS); Los Trancos
_ road, near Stanford University, 1 Apr., 1905, C. P. Smith 727
(CPS) ; Stockfarm bridge, near Stanford University, 13 Apr., 1908,
C. P. Smith t4o01 (CPS). .
SMiTH: STUDIES IN THE GENUS LUPINUS 195
6s. Lupinus densiflorus crinitus Eastwood, var. nov. [FiG. 36.]
Humilis, 8-15 cm. latus, villosissimus, pilis 3-5 mm. longis, caule
ramisque brevibus plerimum laxe pandentibus; foliis congestis;
pedunculis plus minusve a obentitnas vel deflexis, folia exceden-
tibus praesertim ad fructum; peas 2-4 distantibus, floribus
prope 14 mm. longis, pedice
pe 14
brevibus robustis; calyce ha
teolato, 10 mm. longo, 4 mm. lato,
dense villosissimo, pilis 2-3 mm.
longis, labio superiore 3 mm. longo,
emarginato, inferiore 2-dentato,
dentibus 1 mm. longis et I mm.
latis basi, sinu lato; Corellia pluri-
tundato abrupte contracto in un-
guem prope 4 mm. latum; alis I1 Icm.
mm. longis, 6 mm. latis, a! cili- Bh eae pie Anis
atis; carina 10 mm. longa, RODE: icin Rustwood: A. Rastwood (GC).
mm. longis; seminibus angulatis, luteofulvis, dense minuteque
maculatis maculis fulvis.
Low, 8-15 cm. tall, very villous with hairs 3-5 mm. long, stem
and branches short and usually loosely spreading with the peduncles
more or less decumbent or deflexed, exceeding the foliage at time
of fruiting; verticils two to four, distinct, flowers about 14 mm.
long; calyx ebracteolate, 10 mm. long, very densely villous with
hairs 2-3 mm. long; corolla mostly dark purple, the banner 13
mm. long, rounded at the apex and abruptly contracted into a
wide claw, wings ciliate, keel nearly straight; pods large; seeds
angular, yellowish brown, densely and minutely spotted with
brown.
CALIFORNIA. Sonoma County: Bodega Point, A. Eastwood
(Typr, G; type-duplicate, UC); Bodega, 25 May, 1900, H. P.
Chandler 686 (UC). :
The type sheet is labelled “ Lupinus crinitus n. sp.,”” but an
annotation, in pencil, calls it L. densiflorus, var. crinitus. Itseems
to be a local and well-marked variation of the “ palustris group”’
and peculiar to the coast line; while var. persecundus (page 190), of
similar general appearance, but very different pubescence, is
evidently nearer typical L. densiflorus and is found both near the
\
196 SMITH: STUDIES IN THE GENUS LUPINUS
coast and also up in the mountains of the same local region. This
region is not readily accessible to botanists; but I surmise that it
would prove to be an especially interesting locality to taxonomists
of ecological bent.
6t. Lupinus densiflorus curvicarinus var. nov. [Fic. 37.]
Eramosus vel super basi ramosus, 25-30 cm. altus, laxe vil-
losus; foliolis subter specie levibus sed vero pubescentibus, pilis
laxe appressis brevibus, medio-costis marginibusque ciliatis; ver-
ticillis 2-17, plurimum distantibus, floribus laxe pandentibus, 13-14
mm. longis, pedicellis 2 mm. longis, gracilibus, bracteis verticil-
Fic. 37. LUPINUS DENSIFLORUS CURVICARINUS C. P. Smith. 1. K. Brandegee
(UC 155195); 2. C. P. Smith 1460.
lorum humiliorum 15 mm. longis; calyce 8 mm. longo, subter laxe
villoso, pilis 1.5-2 mm. longis, labio superiore emarginato, prope
2 mm. longo, inferiore lanceolato-ovato, inflato, bidentato, dentibus
sinuque, gracilibus, 1 mm. longis; corolla pallido-rosea vel purpurea,
vexillo 12 mm. longo, 7 mm. lato, apice rotundato, alis 11 mm.
longis, carina curvatissima ad basin ciliata; lezuminibus pallido-
stramineis, 12 x 8-13x9 mm.; seminibus asperibus, atro-fulvis,
maculis non claris.
Loosely villous, 25-30 cm. tall, branched above the base or
simple; leaflets apparently glabrous below, but in reality pubescent
with loosely appressed short hairs, the margins and midribs often
ciliate; flowers loosely spreading, 13-14 mm. long; calyx 8 mm.
long, loosely villous below with hairs 1.5-2 mm. long, upper lip
_ emarginate, about 2 mm. long, lower lip lance-ovate, evidently
SMITH: STUDIES IN THE GENUS LUPINUS 197
bent and subsaccate, the two teeth and the sinus slender; een
pale rose or purple, banner 12 mm. long, 7 mm. wide, rounded a
the apex, keel decidedly curved, ciliate; seeds rough, penis poset
the markings indistinct. .
CALIFORNIA. Yolo County: Woodland, 20 May, 1893, J. W.
Blankinship (Type, G). Contra Costa County: Antioch, 5 May,
1907, K. Brandegee (UC 155195, except as to the specimen of var.
Menziesti in center of sheet). Alameda County: Livermore,
16-17 May, 1891, W. L. Jepson (UC). Santa Clara County:
Stanford University, 5 May, 1895, C. Rutter 173 (US 308269,
specimen on left only); Stanford University, 8 June, 1908,
Smith 1460 (CPS).
Curvicarinus is Latin for ‘‘curved keel.”
6u. Lupinus densiflorus Reedii var. nov. [Fic. 38.]
Eramosus, humilis, 4-10 cm. altus, villosus; petiolis I-4 cm.
longis, foliolis 1-2 cm. longis, villosis; pedunculis folia excedentibus,
‘verticillis 2-6, distantibus, floribus pandentibus, prope 13 mm.
longis, pedicellis gracilibus, prope 2
mm. longis; calyce ebracteolato, 8
mm. longo, pilis densis, prope 1.5
mm. longis, labio superiore emar-
ginato, vix 2 mm. longo, inferiore 3
manifeste inflato, 2 mm. lato 3-
_ dentato, dentibus 1 mm. longis, os ;
: a oe
medio graciliore; colore corollae
non claro, vexillo 12 mm. longo, 7
mm. lato, apice rotundato, carina
9 mm. longa, comparate brevi;
legumina seminaque non vidi.
Simple and low, 4-10 cm. tall,
villous; verticils two to six, distant,
flowers spreading, about 13 mm.
long; calyx ebracteolate, 8 mm. thks ae “ . anes Cee
long, upper lip emarginate, lower Comets (2).
lip manifestly bent and inflated
near base, three-toothed, the median tooth more slender; color of
corolla not evident, banner 12 mm. long, 7 mm. wide, rounded
at the apex, keel comparatively short, 9 mm. long; pods and
seeds not seen.
198 SMITH: STUDIES IN THE GENUS LUPINUS
CHILE: Concepcion, E. C. Reed (Type, G).
The sheet bears two plants. The flowers are well pressed, but
the color of the petals is not preserved. L. microcarpus is the
determination given, to which Miss Eastwood has added, ‘not
typical.”
6v. Lupinus densiflorus trichocalyx var. nov. [Fic. 39.]
Eramosus vel basi ramosus, 20-30 cm. altus, dense villosus;
petiolis 10-14 cm. longis, foliolis maximis, 30 mm. longis, subter
manifeste villosis; pedunculis prope folia aequantibus, verticillis
prope 7, distantibus, floribus ad anthesin et specie postea panden-
tibus, 12 mm. longis, pedicellis prope 1
mm. longis, robustis, bracteis brevibus;
calyce ebracteolato, 8 mm. longo, sub-
ter dense villoso pilis 2 mm. longis,
labio superiore bipartito, inferiore lan-
ceolato, bidentato, dentibus 1 mm.
longis, parallelis, gracilibus: colore
corollae non claro, specie purpureo,
vexillo 12 mm. longo, 6 mm. lato,
Elo Smm. #Pice rotundato ungue lato,alis 10 mm.
longis ad basin superne ciliatis, carina
9 mm. longa, curvata: legumina sem-
Fic. 39. LUPINUS DENSIFLO- inaque non vidi.
RUS TRICHOCALYX C. P. Smith.
D. Douglas (G). Densely villous, 20-30 cm. tall,
simple or branched at the base; ver-
ticils about seven, remote, flowers spreading in anthesis and
seemingly later, 12 mm. long, pedicels short and stout and bracts
short; calyx ebracteolate, bushy villous below with hairs 2 mm.
long, upper lip two-parted, lower lip lanceolate, the two teeth
about I mm. long, slender and parallel; color of the corolla not
certain but probably purplish, banner 12 x 6 mm., rounded at the
apex and with a broad claw, wings ciliate at the free edges above,
keel curved: pods and seeds not seen.
CALIFORNIA. County not given: 1829, D. Douglas, ex. Herb.
Lindley (Type, G); Saucelito, 21 May, 1874, J. G. Lemmon
46
.
Perhaps related to this, but not at all typical, are the following:
near Tulare Lake, July, 1878, J. G. Lemmon 124 (G); Cuyama
River, 27 May, 1896, A. Eastwood (G); Arroyo Grande, May,
1895, M. Alice King (UC).
SMITH: STUDIES IN THE GENUS LUPINUS 199
The Douglasian plant upon which this variety is primarily
based is probably one of the collection referred by Agardh to
L. microcarpus,—in his words, ‘‘ Etiam e California retulit Doug-
las.’’ An annotation on the type sheet states that this is the
L. microcarpus of Torrey & Gray’s Flora of North America. If,
as is very probable, the ‘‘Saucelito”’ of Lemmon is the same as the
town now called Sausalito, in Marin County, such might have
been the locality of Douglas’s collection of ‘‘ L. microcarpus.”
It seems to me more probable, however, that these plants came
from Santa Barbara or San Luis Obispo County.
6w. Lupinus densiflorus barbatissimus var. nov. [FiG. 40.]
Ramosus, prope 10 cm. super basi, 25—35 cm. altus, villosus, ad
basin speck petiolis longissimis, 12 cm. longis, foliolis 20-30
mm. longis, superne laxe villosis; verticillis prope 10 appositis,
floribus prope I2 mm. longis, pandenti us,
pedicellis prope 2 mm. longis, paulo graci-
libus, bracteis viliotiedtenda: calyce ebracte-
olato, 8 mm. longo, subter villosissimo, pilis
2 mm. vel plus longis, labiosuperiore prope
longo, emarginato, inferiore mani-
ae - SiBate, 2 mm. lato, 3-dentato, denti- ©
5mm.
bus 1 mm. longis, medio gracillimo; colore
corollae non claro, specie roseo vel purpureo,
vexillotI mm. longo, 5 mm. lato, apice
acuto, paullatim contracto in unguem non
clarum, carina prope 8-9 mm. longa, acu- «© p gph James
mine brevissimo: legumina seminaque non yyaerae (G).
vidi.
Branched above the base, 25-35 cm. tall, fistulose at the base,
villous; whorls about ten, approximate, flowers spreading, about
12 mm. long, bracts very villous; calyx ebracteolate, strongly
bearded below with hairs 2 mm. or more long, upper lip emarginate,
about 1 mm. long, lower lip manifestly subsaccate, tridentate,
the median tooth slender; corolla apparently rose or purple, banner
II X5 mm., acute at the apex, slightly contracted into an indis-
tinct claw, keel about 8-9 mm. long, with a short apical point:
pods and seeds not seen.
Cute. Bath of Collina, Andes, James Macrae (Tere, @).
The sheet bears the following annotations:
o. LUPINUS DEN-
200 SMITH: STUDIES IN THE GENUS LUPINUS
L. microcarpus ? B barbatus Gray in Hb. Kew
art fr. Hb. Hook. The plant which Agardh, from Hb. Lindl., under L. micro-
carpus, refers to as ‘huic proximum,’ and identifies with Dougl. Calif.
“Thus it is evident that neither Agardh nor Gray considered
this plant to be typical ZL. microcarpus. The name “barbatus”’
is not now available, as it has been applied by Henderson to an
Oregon lupine of the Polyphylli.
Barbatissimus, “very much bearded,” refers to the very shaggy
calyx, etc.
6x. Lupinus densiflorus austrocollium var. nov. [Fic. 41.]
A var. palustri differt labio inferiore calycis subrecto vix
inflato; floribus 13 mm. longis, vexillo plurimum apice acuto, 11-1
3
5
3
ongo, 6-7 mm. lato, alis 11 mm. longis, ad basin ciliatis;
leguminibus 13-16 mm. longis: semina matura non vidi.
i
eo,
Fic. 41. Lupmrnus DEN-
SIFLORUS aa ase
c
Abrams 3465 (US Sas
Differs from var. palustris in the lower
lip of the calyx being nearly straight and
hardly inflated; flowers 13 mm. long, banner
usually acute at the apex, 11-13 x 6-7 mm,
long, wings II mm. long; pods 13-16 mm.
long; matured seed not seen.
CALIFORNIA. San Diego County: near
St. Mary’s hospital, San Diego, 12 May,
1903, L. R. Abrams 3465 (Type, DSg 579;
type-duplicates, US, PA, NY, G); San
Diego, May, 1852, G. Thurber 579 (NY,G);
San Diego, 22 Apr., 1885, C. R. Orcutt
(Col U); San Diego, 1876, W. G. W. Har-
ford 185 (NY); Harbison Canyon, Sweet-
water Valley, 6 May, 1888, G. C. Deane
(G); San Ysabel, 12 May, 1893, H. W.
Henshaw 21 (US); El Cajon, 8 Apr., 1894,
T. S. Brandegee (UC); San Diego hills, June-July, 1895, S. G.
Stokes (DS); Canyon Road, San Diego Park, May,1905, K. Bran-
degee (UC); San Diego, May, 1906, Brandegee (NY).
Apparently confined to the foothills about San Diego. It is
probable that all the specimens listed above should not be in-
cluded under one and the same name. Considerable variation
exists and it is ae that none of them should be included in
SMITH: STUDIES IN THE GENUS LUPINUS 201
the L. densiflorus series. Some are labelled “ L. densiflorus,”
others “ L. microcarpus.” Careful field work is needed here.
Austrocollium, ‘of the southern foothills,” is from the genitive
plural of collis, ‘‘foothill,’’ and australis, “southern.”
6y. Lupinus densiflorus scopulorum var. nov. [FIG. 42.]
Ad basin ramosus et superne subramosus, caule 20-30 cm. alto,
villoso, pilis 2-3 mm. longis; petiolis 4-8 cm. longis, conspicue vil-
losis, foliolis 15-25 mm. longis, oblanceolatis, apice rotundatis vel
angulatis, subter laxe villosis; pedunculis folia excedentibus vel
brevioribus, verticillis 3-5, appositis, floribus ad anthesin et postea
pandentibus, prope 15 mm. longis, pedicellis 2!mm. longis, gracilibus,
Fic. 42. LUPINUS DENSIFLORUS SCOPULORUM C. P. Smith. J. Macoun (US
20837).
bracteis villosissimis, cito reflectentibus marescentibusque; calyce
ebracteolato, prope 11 mm. longo, labio superiore 2-3 mm. lo
diverso, interdum integro, snes villosissimo ad anthesi recto
ad fructum subinflato, 3-4 mm. lato, 2-dentato, sinu lato dentis
vestigio instructo; petalis pallido- juteolo-albis, vexillo 14mm. longo,
lato, medio maculato paullatim contracto, apice acuto et
ungue gracili non claroque, alis 12 mm. longis, 6 mm. latis, non
ciliatis, carina 10 mm. longa, subrecta; leguminibus prope 14 x10
mm.: semina matura non vidi.
Branched at the base and babbranched above, 20-30 cm. tall,
villous with hairs 2-3 mm. long; verticils three to five, approximate;
flowers spreading in anthesis and later, about 15 mm. long; bracts
soon reflecting and withering; calyx without bracteoles, about If
mm. long, upper lip 2-3 mm. long, entire or variously toothed,
lower lip very villous, straight in anthesis, slightly inflated in
fruit, two-toothed, the sinus wide with a vestigial tooth present;
petals pale yellowish white, banner 14x 6 mm., gradually con-
202 SMITH: STUDIES IN THE GENUS LUPINUS
tracted into an acute apex and an ill-defined slender claw, wings
12 x6mm., non-ciliate, keel nearly straight: mature seed not seen.
British CoLtumMBIA. Vancouver Island: sea cliffs, Beacon
Hill, 4 July, 1889, J. Macoun 21 (Tyre, US 20837; type-duplicates,
T, G); Aug., 1873, H. Edwards (NY); cliffs by sea, Beacon Hill
Park, Victoria, A. J. Pineo (UC).
WaAsHINGTON. Island County: Whidby Island, N. L. Gard-
ner (UC).
While this form and the var. austrocollium may not be properly
included in L. densiflorus, they certainly are less closely related to
typical L. microcarpus; and var. scopulorum is certainly specifically
distinct from its nearest neighbor, the Yakima Valley form I have
called var. fluviatilis of L. subvexus.
Scopulorum, ‘‘of the cliffs,” is the genitive plural of scopulus.
More extended and much more careful field study of these
forms should contribute both taxonomic and ecological data.
Interesting genetical data should also be obtainable from garden
cultures; but one would have to devise first a suitable method of
treating the seed in order to obtain proper germination, for the
“‘hard-seed problem’’ would probably be the first stumbling stone
to success in cultural studies—at least that has been my experience.
The collection and identification of seeds should be pursued
with especial care, as one may often carelessly, though uninten-
tionally, mix in collecting, at one time and locality, seed of more
than one species or variety.
To the many friends who have generously assisted me in many
different ways my full appreciation is herewith acknowledged.
Listing of all of them will not be attempted here; but I cannot
refrain from mentioning Drs. B. L. Robinson, H. M. Hall, and
P. A. Rydberg, Professors H. H. Bartlett and L. R. Abrams, and
Mr. Sidney F. Blake. Mr. Blake, while in England, looked up the
old material of L. microcarpus, L. densiflorus, etc., and gave me
certain valuable information.
COLLEGE PaRK,
MARYLAND
Studies on the vegetation of New York State—II. The vegetation
of a glacial plunge basin and its relation to temperature *
Loren C. PETRY
(WITH THREE TEXT FIGURES)
A striking feature of the topography of the region immediately
about Syracuse, New York, is the occurrence of numerous plunge
basins produced by waterfalls during the later stages of the glacial
period. The recently established Clarke State Reservation near
Jamesville contains several of these basins; Green Lake in the
Reservation occupies the bottom of a large typical one. Some of
these are known to botanists acquainted with this region as the
habitat of certain rare ferns, especially Botrychium Lunaria (L.)
Sw. (B. onondagense Underwood) and Scolopendrium vulgare Sm.
Maxon} and others have noted that these stations for Scolopen-
drium are always quite cool in summer. Some of these plunge
basins show such remarkably low summer temperatures and dis-’
play a vegetation so distinctly northern in character that a pre-
liminary study of one of them was made during the summers of
1916 and 1917.
The basin studied lies near White Lake, six miles southeast of
Syracuse. It is a natural amphitheater, elliptical in shape, with
sloping sides; at the top or rim it measures about six hundred feet
in length and four hundred fifty feet in width, and has a maximum
depth of about ninety-five feet. Along part of the rim there are
cliffs five to fifteen feet in height, and the slope begins at their
foot; elsewhere the slope begins at the rim and descends at an
angle of approximately thirty degrees until the opposite slope is
met. The longer axis of the basin lies on an east-west line, and
* The first of this series is a general discussion of the outstanding features of
the vegetation of New York state as a whole; the citation to this is
Bray, W. he development of ~ vegetation of New York State. N. Y.
State colt igure 4 Syracuse, N. Y. Tech. Pub. 3, 1915
TM R te. e occurrence a the Hart’s Tongue in America. Fern-
wort aan: aan: Dec. 1900. ‘
203
204 Petry: VEGETATION OF NEW YORK STATE
the bottom presents the appearance of a ravine running in that
direction. The outlet of the basin lies to the southeast, where
there is a notch about fifty-five feet in depth in the rim; this
lowest point of the rim is forty feet above the bottom of the basin,
which lies at an altitude of about six hundred ten feet above sea
level.
The basinlies in rather thick-bedded limestone for its entire
depth, and the slopes are formed of debris of this material.
iB
A
Fic. 1. Map of plunge basin, with isotherms of air temperatures (Fahr.) at
1 in. from the ground, at noon, September 12, 1916. A, B, C and D indicate location
of stations referred to in the text. Scale, 1 in. = 160 ft.
Throughout this region the rock is extensively fissured and it is
probable that the underlying bedrock here is traversed by crevices,
since water does not collect in the basin. Except on a considerable
portion of the north side, where there is a rock slide, the slopes are
covered by a thin layer of nearly pure humus. Under the condi-
tions which occur here this soil is always moist and plant growth
in the basin is not restricted by lack of available water.
The basin lies in a forest which has been somewhat disturbed
by cutting. The interior of the basin itself is heavily wooded
with Tilia, Ti huja, Betula lutea, etc. The outlet is a narrow notch
whic sitar only half the depth of the basin. As a result of
PETRY: VEGETATION OF NEW YorK STATE 205
these conditions, the bottom is completely free from air currents
due to winds; even on a day when the local weather bureau station
reported a wind velocity of thirty-five miles per hour, no move-
ment of the air was perceptible half way down the slope of the
basin. This absence of air currents is of prime significance in the
explanation of the striking conditions of temperature that occur
here.
Four stations for the securing of data on temperature and
humidity were established in the basin. The first of these (FIG. 1,
A) is six feet back from the rim, on the south side; it is in a forest
of Tila and Acer saccharum. The second station (B) is fifty feet
down the slope from A; Tilia, Scolopendrium and Impatiens are
the characteristic plants. Station C is one hundred feet further
down the slope, while D is at the bottom of the basin, about one
hundred feet below C. Temperature and humidity readings
were made at each of these stations several times during the sum-
mer of 1916 and once during 1917; the data obtained on August 22,
1916, the hottest day of the year, are given in TABLE I.
TABLE I
TEMPERATURE AND HUMIDITY DATA AT STATIONS IN PLUNGE BASIN. AUGUST 22,
IQ1I6, 2.30—4.30 P.M.
Temp (Fabr.) Humidity (4)
Sta- i } i |
tion | 3 ft. above Gin-above |... -; .. | 6in. above 4ft.above | 6in. above
ground ground be ee soil | ground | ground | ground
Aes 94.7 90.6 81.5 72.5 | 32 30
B.. 90.0 | 76.6 75.2 [se es F
Gi 90.0 78.0 | 73.0 68.5 3 43
oe 63-5 59-7 51.3 42.5 73 71
The remarkable character of the conditions in the plunge basin
is sufficiently shown by the data obtained at station D, where air
temperatures were thirty degrees below those of station A, and
where the soil temperature at a depth of six inches was only ten and
one half degrees above freezing point. Readings made on other
dates gave data similar to those of TaBLe I, but in no case were
any of the temperatures at station D higher than those given above.
Similarly, the humidity at D varied from 75 to 85 per cent. but was
at no time found to be below that recorded in TasLe I.
The cause of the low temperatures in the bottom of the basin
206 PETRY: VEGETATION OF NEW YorK STATE
is not clear, but is probably to be found in the accumulation of
ice in underlying fissures. In the basin itself ice persists in the
spring about a month after it has disappeared elsewhere. It is
_ possible that a sufficient amount of ice accumulates in large crev-
ices or fissures under the loose rock of the basin to maintain the
low temperatures through the summer season. Some such locali-
zation of the areas of lowest temperature is indicated by the data
on soil temperatures given in Fic. 2.
Fic. 2. Map of plunge basin with isotherms of soil temperatures (Fahr.) at depth
of 1 inch at noon, September 12, 191 Scale ft.
In order to determine with exactness the temperature condi-
tions over the entire bottom of the basin, a system of twenty
stations, at distances of thirty-five to fifty feet from each other,
was established; these covered an irregular area extending about
three hundred feet along the axis of the basin, some ninety feet
up the south slope and about seventy feet up the northern slope.
‘Two thermometers were placed at each station, one, completely
shaded, at one inch from the ground, the other with the center of
the bulb in the soil to a depth of one inch; after thirty minutes all
the thermometers were read simultaneously.
When the temperature data are plotted on a map of the basin,
isotherms can be drawn to represent the temperature conditions.
Fic. 1 shows air isotherms representing the conditions existing
PETRY: VEGETATION OF NEW YoRK STATE 207
between 11:30 A. M. and 12 M. on September 12, 1916, when the
temperature at three feet from the ground at station A (on the
rim) was 73° F.; Fic. 2 shows the soil isotherms of the same date.
These are typical of the results obtained; charts made from the
data obtained on other dates show that the isotherms vary but
little in shape, only moving up and down the slopes within narrow
limits with changes of temperature outside the basin. When the
outside temperature falls below that within the basin the system
of isotherms is of course destroyed.
As mentioned, Scolopendrium occurs abundantly in this basin,
which is a typical habitat for it, associated with Impatiens pallida
Nutt., Asarum canadense L., Rubus odoratus L., Aspidium mar-
ginale (L.) Sw., Cystopteris bulbifera (L.) Bernh., etc.; this asso-
ciation is restricted to the upper third of the south slope of the
basin and is therefore outside the region of very low temperatures.
The greater part of the basin is occupied by an association in which
Thuja occidentalis L., Betula lutea Michx. f., and Acer spicatum
Lam. are the dominant members; ground forms are few, mostly
Cystopteris bulbifera, Aspidium marginale (L.) Sw., A. spinulosum
(O. F. Miiller) Sw. var. intermedium (Muhl.) D. C. Eaton, and
Thuidium spp. The tree members of this association extend to
the bottom of the basin but are less abundant there than farther
up the slopes.
The forest floor association of the bottom area is the striking
feature of the vegetation of the basin; a list of the species follows.
DOMINANT SPECIES
Cornus canadensis L. Phegopteris Dryopteris (L.) Fée.
Lycopodium annotinum L. Pyrola asarifolia Michx.
Dalibarda repens L. Ribes lacustre (Pers.) Poir.
Coptis trifolia (L.) Salisb.
INCIDENTAL SPECIES
Lycopodium lucidulum Michx. Aralia nudicaulis L.
L. obscurum L. var.dendroideum Tiarella cordifolia L.
(Michx.) D. C. Eaton Streptopus roseus Michx.
Aspidium marginale Actaea rubra (Ait.) Willd.
A. spinulosum var. intermedium Diervilla Lonicera Mill.
208 PETRY: VEGETATION OF NEW YORK STATE
Of these species, Bray, in the first paper of this series, lists three
of the dominant ones as indicators of the Canadian zone—charac-
teristic of the Adirondacks above 3,500 ft.—and four others as
occurring abundantly in the Canadian Transition zone—character-
istic of the higher Catskills and of the Adirondacks up to 3,500 ft.;
and Cooper* lists four of the dominant species as characteristic of
the climax forest of Isle Royale, Lake Superior. In addition, the
manuals indicate a northern distribution for all the remaining
species. These citations are sufficient to indicate the distinctively
northern character of the plant association of the bottom of the
basin.
Without entering into a discussion of the general questions
of discontinuous distribution, it is evident that this may be con-
sidered to be a typical relict association, left behind at the final
recession of the ice sheet. In this interpretation, certain condi-
tions exist in this plunge basin which have caused or permitted
the maintainance here of a plant association characteristic of
more northerly regions or areas of higher altitude. That low
temperature is the controlling factor of these conditions is the
immediate and obvious conclusion from the data given above.
This conclusion is strongly supported by further analysis of con-
ditions in the basin.
During the work in the basin it was early noticed that the
characteristic species of the bottom association are restricted to a
common region of limited area and irregular shape; that is, the
conditions which caused or permitted the maintenance of the
association in the basin exist only in a small area of its bottom.
As already described, the conditions of soil moisture, humidity,
wind, etc., vary but slightly within the basin and cannot be con-
trolling factors. The growing season in the bottom area is con-
siderably shorter than in the upper part of the basin, due to the
late melting of accumulated snow; the frostless period is probably
about one hundred twenty-five days—an average period for the
Canadian-Transition zone of Bray. This short growing season
cannot however be effective in preventing the spread of the bottom
association up the slopes of the basin, for not only do most of the
* Cooper, W. S. The climax forest of Isle Royale, Lake Superior, and its -
Bot. Gaz. 55: Laas II5—140, 189-235. 1913.
PETRY: VEGETATION oF NEW York STATE 209
species of the association spread readily by vegetative propagation,
but ripe spores or seeds are also produced by most of them. On
the other hand, the short season cannot be effective in preventing
the invasion of plants of the higher parts of the slope, since these
species have a longer season in which to mature spores or seeds, and
every facility for their distribution into the bottom area.
With regard to the temperature conditions, however, the case
is different. Casual observation indicated that the characteristic
species of the bottom association do not occur outside the region
of noticeably low temperatures. To verify this observation the
Fic. 3. Map of plunge basin showing distribution of Ribes lacustre (Pers.) Poir.
tin. = 160 ft
exact distribution within the basin of five of the dominant species
was determined and plotted. The maps of the distribution areas
are remarkably alike; that for Ribes lacustre, given in FIG. 3, is
typical. By comparison with Fre. 1 it will be seen that the dis-
tribution area is only roughly that of low air temperatures, being
extended farther along the axis of the basin. A comparison of
Fics. 2 and 3, however, shows a remarkable similarity between the
outline of the distribution area of Ribes and the soil isotherm of
56° Fahr.; and the maps of the distribution areas of the other species
show an equally close resemblance. This coincidence of the areas
of low soil temperature and of distribution of five of the dominant
210 PETRY: VEGETATION OF NEW YorK STATE
species is held to indicate a direct causal relationship between the
two. The writer is of the opinion that we have here a case in
which the control of distribution of a plant association by the
single factor of temperature is demonstrable; and that the plunge
basin offers an exceptionally favorable opportunity to determine
experimentally the mechanism of this control. Work is now in
progress along this line.
SUMMARY
I. In the bottoms of certain plunge basins near Syracuse,
New York, low temperatures prevail throughout the year. Data
of air and soil temperatures are given for one such basin whose
bottom lies at an altitude of six hundred ten feet above sea level;
an area of some thousands of square yards in it remains throughout
the year at a temperature below 70° Fahr.
2. The bottom area of this basin is occupied by an association
of plants characteristically Canadian in distribution. Analysis
of the conditions in the basin demonstrates that temperature is
the factor controlling their distribution in this area.
SYRACUSE UNIVERSITY,
YRACUSE, NEw YorK
INDEX TO AMERICAN BOTANICAL LITERATURE
1912-1918
The aim of this Index is to include all current botanical literature written by
Americans, published in America, or based upon American material ; the word Amer-
ica being used in the broadest se
Reviews, and papers that res exclusively to forestry, agriculture, horticulture,
manufactured products of vegetable origin, or laboratory methods are not included, and
no attempt is made to index the cae of bacteriology. An occasional exception is
R a
Some important particular. If users of the Index will call the attention of the editor
to errors or omissions, their kindness will be appreciated.
This Index is reprinted monthly on cards, — furnished in this form to subscribers
at the rate of one cent for each card. Selections of cards are not permitted ; each
subscriber must take all cards published sata the term of his subscription, Corre-
spondence relating to the card issue should be addressed to the Treasurer of the Torrey
Botanical Club
Allard, H. A. Abnormalities in Nicotiana. Bot. Gaz. 65: 175~-185.
£.. I-10. 35-¥ 1018.
Allard, H. A. The mosaic disease of Phytolacca decandra. Phyto-
pathology 8: 51-54. f. Zz, 2. 25 F 1918.
Bachmann, F M. A bacteriological method useful for the study of
other micro-organisms. Am. Jour. Bot. 5: 32-35. f. 1, 2. Ja 1918.
Barrett, J. T. Some observations on wither-tip in 1914. Proc. Fruit
Growers Conv. Calif. 45: 242-244. 1915.
Benedict, R.C. Two serious fern pests. Am. Fern Jour. 7: 122-124.
23 F 1918.
Bergman, H. F. Comments on Malva rotundifolia L. and its allies.
Minnesota Bot. Stud. 4: 437-442. pl. 47, 48. 20S 191
Bergman, H. F., & Stallard, H. The development of climax ee
in northern Minnesota. Minnesota Bot. Stud. 4: 333-378. f. 1.
20 S 1916.
Blake, S. F. On the names of some species of Viburnum. Rhodora
20: II-15. 25 Ja 1918.
Boot, D. H. Plant studies in Lyon County, Iowa. Proc. lowa Acad
Sci. 24: 393-414. pl. 11, 12 +f. 80. 1917.
des, E. W. Report of the plant pathologist. Rep. Porto Rico
Agr. Exp. Sta. 1916: 28-31. 2 4; §. a F 1918.
Includes descriptions of Fusarium cubense S
211
A&A INDEX TO AMERICAN BOTANICAL LITERATURE
Braun, E.L. Regeneration of Bryophyllum calycinum. Bot. Gaz. 65:
1Ot- 19a. 7. fy ay. 15 F161,
Britton, N. L. The relatives of catalpa trees in the West Indies.
Jour. N. Y. Bot. Gard. 19: 6-9. pl. 209. Ja 1918.
Brown, J.G. Mistletoe vs. mistletoe. Bot. Gaz. 65: 193. *. 7. 155
1918.
Buchanan, R. E. Studies in the nomenclature and classification of
the bacteria—V. Subgroups and genera of the Bacteriaceae. Jour.
Bact. 3: 27-61. Ja 1918.
Burnham, S.H. Lake George flora stations for Botrychium lanceolatum.
Am. Fern Jour. 7: 124. 23 F 1918.
Butler, O. On the preservation of phytopathological specimens in
their natural colors. Phytopathology 8: 66-68.
Butters, F. K. Notes on the species of Liagora and Galaxaura of the
Central Pacific. Minnesota Bot. Stud. 4: 161-184. pl. 24. 5 3
IQII.
ae peers maxims, L. intricata, L. ca ev geet L. hawaiiana, L.
Tildenii axaur and G. mauiana, spp. ni
“SeMRings : K. Some bbcuilne cases of plant phe in the Selkirk
Mountains, British Columbia. Minnesota Bot. Stud. 4: 313-331.
J. is f5 Me 1014,
Butters, F. K., & Rosendahl, C. O. Some effects of severe frost upon
vegetation in a condition of active growth. Minnesota Bot. Stud.
4: 153-180. pl. 23, 35S 1011.
Clements, F.C. Nova fungorum coloradensium genera. Minnesota
Bot. Stud. 4: 185-188. pl. 25. 15S 1911.
Includes Comoclathris lanata, C. ipomoeae, Pezoloma griseum, Leucopezis excipu-
lata, Sirodothis populi and Sirocyphis nivea, spp.
Co , T. D. A. A new hybrid uted Torreya 18: 11-14.
13 oS i518.
Coit, J. E., & Hodgson, R. W. The June drop of Washington navel
oranges. California Agr. Exp. Sta. Bull. 290: 203-212. f. 1-3.
Ja 1918.
Conard, H. S. The white water lily of Clear Lake, Iowa. Proc.
Towa Acad. Sci. 24: 449-454. f. 88. 1917.
Corson, G. E., & Bakke, A.L. The use of iron in nutrient solutions for
plants. Proc. Iowa Acad. Sci. 24: 477-482. f. 95-98. 1917.
Davis, W.H. The aecial stage of alsike clover rust. Proc. Iowa Acad.
Sci. 24: 461-472. pl. 15, 16 + f. 89-04. 1917.
Davis, W. H. Chlorotic corn. Proc. Iowa Acad. Sci. 24: 459, 460.
1917.
INDEX TO AMERICAN BOTANICAL LITERATURE 213
Deam, C. C. Cheilanthes lanosa and Isoetes in Indiana. Am. Fern
Jour. 7: 112-114. 23 F 1918.
Douglas, G.E. The development of some exogenous species of agarics.
Am. Jour. Bot. 5: 36-54. pl. 1-7. Ja 1918.
Includes description of Mycena subaicalina Atkinson sp. nov.
Elmore, C. J. Changing diatoms of Devils Lake. Bot. Gaz. 65: 186—
190. 15 F 1918.
Elrod, M. J. Ophioglossum vulgatum L. in Montana. Am. Fern Jour.
wr.125. +23 F 19018
Fairchild, D. The seated papaya as an annual fruit tree. Mod.
Cuba Mag. 1: 14-20. N 1913. [Illust
Fernald, M.L. American variations of Epilobium, section Chamaener-
ton. Rhodora 20: 1-10. 25 Ja 1918.
Fernald, M.L. Some American epilobiums of the section Lysimachion.
Rhodora 20: 29-39. 12 F 1918
Includes Epilobium Steckerianum sp. nov.
Folsom, D. Studies in the morphology of Yucca glauca. Minnesota
Bot. Stud. 4: 427-435. pl. 43-46. 20S 1916.
Godfrey, G. H. Sclerotium rolfsii on wheat. Phytopathology 8: 64-
66. f. i.° 25 ¥ 16th:
Gravatt, G. F., & Posey, G. B. Gipsy-moth larvae as agents in the
dissemination of the white-pine blister rust. Jour. Agr. Research
12: 459-462. 18 F 1918.
Harlan, H. V. The identification of varieties of barley. U.S. Dept.
Agr. Bull. 622: 1-32. pl; I-4. 2 F 1918
Harper, R. M. An interesting peat bog in New York City. Jour. Am.
Peat Soc. 11: 8-11. Ja 1918.
Harter, L. L., & Jones, L. R. Cabbage diseases. U.S. Dept. Agr.
Farm. Bull. 925: 1-30. f. 1-13. Ja 1918.
Hartley, C. Rhizoctonia as a needle fungus. Phytopathology 8: 62.
25 F 1918.
Hastings, G. T. Some abnormal! poplar flowers. Torreya 17: 16-20.
j..Pad.. 13 F 1918;
Hills, T. L. Influence of nitrates on nitrogen-assimilating bacteria.
Jour. Agr. Research 12: 183-230. 28 Ja 1918.
Hodgson, R. W. An account of the mode of foliar eee in Citrus.
‘Univ. Calif. Publ. Bot. 6: 417-428. f. 1-3. IF 1
pals R. H. A further note on the lichens of ie
0: 40. 12 F 1918.
dea N. L. Response of micro-organisms to copper sulphate treat-
ment. Minnesota Bot. Stud. 4: 407-425. f. I-5. 20
Rhodora
214 ' INDEX TO AMERICAN BOTANICAL LITERATURE
Johnson, A. G., & Vaughn, R. E. Ergot in rye and how to remove it.
Wisconsin Agr. Sep. Sta. Ext. Serv. Circ. 94: [1-4]. f. 1-4. Ja 1918.
Johnson, J. R. The citrus canker. Mod. Cuba Mag. 2: 63-65. N
1914. [Illust.]
Johnson, J. R. An insect fungus. Mod. Cuba Mag. 2: 76, 77. D
1914. [Illust.]
Johnston, J. R., & Bruner, S.C. A Phyllachora of the royal palm.
Mycologia 10: 43, 44. pl. 2. 14 F 1918.
Johnston, J. R., & Stevenson, J. A. Sugar-cane fungi and diseases of
Porto Rico. Jour. Dept. Agr. Porto Rico. 1: 177-264. pl. 19-31.
O 1917.
cludes Himantia stellifera, Chromocreopsis striispora, Arthrinium saccharicola,
Hormiactella sacchari, Septonema sacchari, Periconia sacchari, Tetracoccosporis
sacchari, and Verticicladium graminicolum, spp.
Jones, M. E. The willow family of jhe yore plateau. 1-32. Salt
Lake City. 9 M 1908.
Kellogg, H. S. Preservation of native plants in Iowa. Proc. lowa
Forest & Conserv. Assoc. I9I4-1915: 80-99. I916.
Kinman, C. F. The mango in Porto Rico. Porto Rico Agr. Exp. Sta.
Bull. 24: 1-30. pl. 11. 4 F 1918.
Kirby, R. S. Influence of orchard soil management on fruit bud de-
velopment and formation as found in the apple. Proc. Iowa Acad.
Sci. 24: 447, 448. 1917. :
Loeb, J. Chemical basis of correlation. I. Production of equal
masses of shoots by equal masses of sister leaves in Bryophyllum
calycinum. Bot. Gaz. 65: 150-174. f. 1-18. 15 F 1918.
Ludwig, C. A. The influence of illuminating gas and its constituents
on certain bacteria and fungi. Am. Jour. Bot. 5: 1-31. Ja 1918.
MacCaughey, V. The California laurel. Hawaiian Forester 14: 194,
195. Jl 1917.
MacCaughey, V. The carambola in Hawaii. Hawaiian Forester 14:
304-306. O 1917.
M ee V. The Hawaiian Violaceae. Torreya 18: 1-11. 13F
ees V. Lichen flora of the Hawaiian Islands. Hawaiian
Forester 14: 303, 304. O 1917.
MacCaughey, V. The Mangosteen in Hawaii. Hawaiian Forester 14:
125,126. My 1917.
Mann, A. The economic importance of the diatoms. Smithsonian
Rep. 1916: 377-386. pl. 1-6. 1917.
axon, The Aisatks range of Botrychium lanceolatum.
Rhodora 20-50; 20. a Ja 1918.
INDEX TO AMERICAN BOTANICAL LITERATURE 215
Maxon, W. R. A new Notholaena from the southwest. Am. Fern
Jour..7: 106-109. 23 F 1918.
Notholaena Jonesti sp. nov.
Maxon, W. R. Notes on western species of Pellaea. Proc. Biol. Soc.
Washington 30: 179-184. 1 D 1917.
Includes Pellaea compacta sp. nov.
McBeth, I. G. Relation of the transformation and distribution of soil
nitrogen to the nutrition of citrus plants. Jour. Agr. Research 9:
183-252. f. r-r9. 14 My 1917.
McGregor, E. A. List of plants from Batesburg, S. C., and vicinity.
Jour. Elisha Mitchell Sci. Soc. 33: 133-145. N 1917.
Merrill, E. D. Koordersiochloa javanica Merrill, a new genus and
species of Gramineae from Java. Philip. Jour. Sci. 12: (Bot.)
67-69. pl. r. Ja ¥917.
Merrill, E. D. Contributions to our knowledge of the flora of Borneo.
Jour. Straits Branch R. A. Soc. 76: 75-117. 1917.
Includes descriptions of thirty-five new species.
Merrill, E. D. The dates of publication of the third edition of Blanco’ s
“Flora de Filipinas.’ Philip. Jour. Sci. 12: (Bot.) 113-116. Mr
1917.
Merrill, E. D. An interpretation of Rumphius’s Herbarium Amboi-
nese. Bur. Sci. Manila Publ. 9: 1-595. pl. 1,2 +f. 1
Merrill, E. D. New Philippine Lauraceae. Philip. Jour. So. fa:
(Bot.) 125-141. My 1917.
Includes descriptions of nineteen new species.
Merrill, E. D. New Philippine Myrsinaceae. Philip. Jour. Sci. 12:
(Bot.) 143-158. My 1917.
Includes descriptions of eighteen new species.
Merrill, E. D. Notes on the flora of Kwangtung Province, China.
Philip. Jour. Sci. 12: (Bot.) 99-IIT. Mr 1917.
Includes descriptions of five new species.
Merrill, E. D. Two new ‘genera and four new species of Philippine
Compositae. pce Jour. Sci. 12: (Bot.) 117-124. Ol. 2) 3: My
1917.
Metcalf, H. Summary of the white- -pine blister rust situation. Jour.
Forestry 16: 85-89. Ja 1918.
Moore, B. Reproduction in the coniferous forests of northern New
England. Bot. Gaz. 64: 149-158. 15 Au 1917.
Moore, B. Some factors influencing the reproduction of red spruce,
balsam fir, and white pine. Jour. Forest. 15: 827-853- f. I-4.
N 1917.
Murrill, W. a Growing walnuts for food. Jour. N. Y. Bot. Gard.
19: 9-11. Ja 1918.
216 INDEX TO AMERICAN BOTANICAL LITERATURE
Nash, G. V. Hardy woody plants in the New York Botanical Garden.
18: 246-250, 259-263. N 1917; 18 D 1917; Ig: 11-15. Ja1g918.
Osterhout, G. E. Concerning some species of Carduus in Colorado.
Torreya 18: 14-16. 13 F 1918.
Pammel, L. H. The flora of Lake Vermillion, Minnesota. Amer.
Forester 5: 40-44. 1917. [Illust.]
Pammel, L.H The preservation of Iowa’s lakes from the standpoint
of a botanist. Proc. Iowa Forest & Conserv. Assoc. 1914-1915:
173 1755. (IGEG:
Pammel, L. H. The preservation of natural history spots in Iowa.
Proc. Iowa Forest. & Conserv. Assoc. 1914-1915: 105-108. 1916.
[Iflust.}
Pammel, L. H. The vegetation of Iowa lakes. Rep. State Highway
Com. 163-189. f. 61-79. 1917.
Pammel, L. H., & King, C. M. The germination and juvenile forms of
some oaks. Proc. Iowa Acad. Sci. 24: 376-391. f. 58-79. 1917.
Pammel, L. H., & Kenoyer, L. A. Some notes on pollination of red
clover. Proc. Iowa Acad. Sci. 24: 357-366. 1917.
Parker, J. H. Greenhouse experiments on the rust resistance of oat
varieties. U. S. Dept. Agr. Bull. 629: 1-16. pl. 1-3 +f. 1, 2-
Pease, A.S. Nuttall and Pickering in the White Mountains. Rhodora
20: 39. 12F 1018.
Pool, R. J. A study of the vegetation of the sandhills of Nebraska.
Minnesota Bot. Stud. 4: 189-312. pl. 26-40 +f. 1-16. 15 Mr
1914.
Prizer, J. A. Some experiments in treating citrus trees for gummosis.
Proc. Fruit Growers. Com. Calif. 45: 218-229. f. 30-36. 1915.
Ransier, H. E. Fresh pleasures from old fields. Am. Fern Jour. 7:
99-104. pl. 5, 6. 23 F 1918.
Reed, G. M. Phytopathological survey of the trees and shrubs of
Prospect Park and the Botanical Garden (Brooklyn).—Il. Report
of the second season’s work. Brooklyn Bot. Gard. Record 7: 14-
24. ja 1988.
Rohret, M. B. The morphology of the thallus and cupules of Blasia
pusilla. Proc. lowa Acad. Sci. 24: 429-454. f. 84-87. 1917.
Rogers, W. E. Notes on Melilotus alba, white sweet clover. Proc.
Iowa Acad. Sci. 24: 415-428. pl. 13, 14 + f. 81-83. 1917.
Roig, J. T. The Cuban walnut. (Juglans insularis Griseb.) Mod.
Cuba Mag. 2: 40-42. Au 1914.
Safford, W. E. Narcotic plants and stimulants of the ancient Amer-
icans. Smithsonian Rept. 1916: 387-424. pl. 1-17 +f. 1-6. 1917.
INDEX TO AMERICAN BOTANICAL LITERATURE 217
Salmon, S. C. A preliminary note on soil moisture and temperature
factors in the winter-killing of grain crops. Science II. 47: 173,
F
Schaffner, J. H. Additions to the catalog of Ohio vascular plants for
1917. Ohio Jour: Sci. 18: 99, 100. 31 Ja 1918.
Seaver, F. J. Photographs and descriptions of cup-fungi—VII. The
genus, Underwoodia. Mycologia 10: 1-3. pl. 1. 14 F 1918.
Shufeldt, R. W. Studies of leaf and tree (Part 1). Am. Forest. 24:
29-35. f. 1-15. Ja1918. Part II. Am. Forest. 24: 95-102. f. 18~
3r. F. 1918; Part III. Am. Forest. 24; 151-159. f. 35-50. Mr.
1918
Small, I. K. Collecting prickly-pears at Apalachicola. Jour. N. Y.
Bot. Gard. 19: 1-6. pl. 208. Ja 1918
Spurrell, J. A. The disappearance of native plants in Iowa. Proc.
Iowa Forest. & Conserv. Assoc. 1914-1915: 100-104. I916.
Standley, P. C. Rusts and smuts collected in New Mexico in 1916.
Mycologia 10: 34-42. 14 F 1918.
Stevenson, J. A. Citrus scab in Porto Rico. Porto Rico Dept. Agr.
& Lab. Exp. Sta. Bull. 17: 1-16. 1917.
Stevenson, J. J. Interrelations of the fossil fuels. III. Proc. Am.
Philos. Soc. 57: 1-48. 30 Ja 1918.
St. John, H. Additional wool waste plants. Rhodora 20: 20. 25 Ja
1918.
St. John, H. Lycopodium Selago var. Miyoshianum in North America.
Am. Fern Jour. 7: 121, 122. 23 F 1918.
Terao, H. Maternal inheritance in the soy bean. Am. Nat. 52: 51~
Thomas, A.O. Ona supposed fruit or nut from the tertiary of Alaska.
Proc. Iowa Acad. Sci. 24: 113-118. pl. 5. 1917.
Thomas, W. A. A Picea from the glacial drift. Proc. Iowa
Sci. 24: 455,456. 1917.
Thompson, W. P. Independent evolution of vessels in Gnetales and
Angiosperms. Bot. Gaz. 65: 83-90. f. I-11. 17 Ja 1918.
Thone, F. E. A. Pioneer plants on a new levee. III. Proc. lowa
Acad. Sci. 24: 457, 458. 1917.
Waksman, S. A. Soil fungi and their activities. Soil Sci. 2: 103-156.
pl, 1-5. Au 1916.
Wauters, W. M. (Mrs.). A plea for the laurel: mountain laurel as our
national flower [1-3]. Published by Hotel McAlpin, New York.
Ja 1918.
Weir, J R. New hosts for Razoumofskya laricis. Phytopathology 8:
62, 63. 25 F 1918.
Sead.
218 INDEX TO -AMERICAN BOTANICAL LITERATURE
Weir, J.R. Notes on the altitudinal range of forest fungi. Mycologia
10: 4-14. 14 F 1918.
Weir, J.R. A short review of the general characteristics and cytological
phenomena of the Uredineae, with notes on a variation in the
promycelium of Coleosborium pulsatillae (Str.). New Phytol. 11:
129-139. Ap 1912.
Weir, J. R. Some problems in conservation with reference to forest
hygiene. The Timberman. So Sk SOT
Treats of wood-destroying fungi.
Weir, J.R., & Hubert, E.E. Cronartium coleosporioides on Pedicularis
groenlandica. Phytopathology 8: 63. 25 F 1918.
Weir, J. R., & Hubert, E.E. A note on Hyalopsorae. Phytopathology
&: 47,38. 24 Ja 1918.
Weir, J. R., & Hubert, E. E. Notes on the overwintering of forest
tree rusts. Phytopathology 8: 55-59. ' 25 F 1918.
Wherry, E. T. Observations on the habitat of certain ferns. Am.
Fern Jour. 7: 110-112. 23 F 1918.
White, O. E. Studies of inheritance in Piswm—II. The present
state of knowledge of heredity and variation in peas. Proc. Am.
Philos. Soc. 56: 487-588. 8 D 1917.
Wolf, F. A., & Foster, A. C. Bacterial leaf spot of tobacco. Science
II. 46: 361, 362. 12019
Wolf, F. A., & Stanford, E. E. A Macrophoma disease of figs. ,. Phyto-
pathology 8: 24-27. f. 1, 2. 24 Ja 1918.
Woodward, K. W. Tree growth and climate in the United States.
Jour. Forestry 15: 521-531. My 1917.
Wright, C. H. Orthrosanthus chimboracensis. Curt. Bot. Mag. IV.
Wuist, E. D. Apogamy in Phegopteris a cage a Fée, Osmunda
cinnamomea L., and O. Claytoniana L. . Gaz. 64: 435-437.
16 N 1917.
Wylie, R. B. Cleistogamy in Heteranthera dubia. Bull. Lab. Nat.
Hist. State Univ. Iowa 7!: 48-58. pl. rr. 1917. :
Zimmerman, H. E. How Brazil nuts grow. Am. Bot. 23: 88, 89.
Au 1917. [Illust.] :
Zimmerman, H.E. A novel Hiniper tree. Am. Bot. 23: 130. N 1917.
[Ilust.]
Zinn, J., & Surface, F. M. Studies on oat breeding —V. The F, and
_ F2 generations on a cross between a naked and a hulled oat. Jour.
on Research 10: 293-312. pl. 39-47. 6 Au 1917.
Vol. 45 No. 6
BULLETIN
OF THE
TORREY BOTANICAL CLUB
JUNE, 1910
A carrier of the mosaic disease
MakKOTO NISHIMURA
(WITH PLATE 7)
INTRODUCTION
The symptoms and general character of the mosaic disease
of tobacco and other solanaceous plants are becoming very well
known through the work of Mayer (1886), Sturgis (1900), Woods
(1899, 1902), Lowe (1900), Iwanowski (1903), Hunger (1905),
Clinton (1908), Westerdijk (1910), Peters (1912), Allard (1914,
1915, 1916, 1917), Chapman (1913, 1916), Jensen (1913), and
Freiberg (1917).
‘The disease has been observed on all varieties of Nicotiana
Tabacum, on several of the more distinct varieties of tomato, on
Petunia violacea, on Physalis (two distinct garden species), on
Datura Stramonium and D. Tatula, on Hyoscyamus niger, On
Solanum nigrum and S. carolinense and on several of the more
distinct varieties of Capsicum. It has been transferred to all
these from infected N. Tabacum. To this list should be added now
Solanum aculeatissimum, the apple of Sodom, an infected plant of
which was brought from Florida to the greenhouse of Columbia
University in the winter of 1915.
According to Allard (1914) the following symptoms are charac-
teristic of different phases of the disease at one or another stage in
the infection: (1) partial or complete chlorosis, (2) curling of the
leaves, (3) dwarfing and distortion of the leaves, (4) blistered or
[The BULLETIN for May (45: 167-218) was issued May 23, 1918.]
219
220 NISHIMURA: A CARRIER OF THE MOSAIC DISEASE
‘“‘Savoyed’’ appearance of the entire plant, (5) mottling of the
leaves with different shades of green, (6) dwarfing of the entire
plant, (7) dwarfing and distortion of the blossoms, (8) blotched
or bleached corollas (in N. Tabacum only), (9) mosaic sucker
growth, (10) death of tissue (sometimes very marked in N. rustica).
To these should be added the narrowing of the leaves or frenching
and their sometimes uniformly lighter or more yellowish green
color.
In attempting to infect the alkekengi or bladder cherry (Phy-
salis Alkekengi) I have found that although this plant when inocu-
lated does not show any of the above visible symptoms, it may,
nevertheless, become a carrier of the disease in the fullest sense,
for I have infected N. Tabacum and the apple of Sodom with the
juice of plants of the alkekengi which appeared to be entirely free
from mosaic.
Allard reports (1914) that he has readily transferred the disease
from tobacco to two distinct garden species of Physalis, which
then showed the general symptoms of the disease, but he does not
tell us what species of Physalis he used.
As grown by me the alkekengi certainly shows none of the ordi-
nary symptoms, the plants remaining healthy in appearance for
indefinite periods, after inoculation with the juice of diseased
tobacco or apple of Sodom.
Allard (1917) reports some very interesting experiments on
the behavior of the mosaic disease in N. glauca. He inoculated
this species from diseased plants of N. Tabacum and found that
the symptoms were confined to a sparse and indistinct mottling
along the veins of some of the leaves. This mottling in some in-
stances was too faint to be detected readily, except in transmitted
light. Eight of these plants which showed these symptoms more
clearly were tested by injecting the expressed sap of each into a
series of plants of N. . Tabacum. The sap of all proved exception-
ally virulent, producing in most instances 100 per cent. of infection
in each lot of ten plants. After the initial expression of the disease
in N. glauca, Allard reported that the symptoms gradually became
more attenuated, until they were barely distinguishable. In
July three of these plants were cut back severely and were im-
— transplanted to the field. Growth was resumed, but
NISHIMURA: A CARRIER OF THE MOSAIC DISEASE ype |
no symptoms of the mosaic disease whatever could be detected.
However, inoculation tests demonstrated that the sap of these
‘plants contained the infective principle of the disease. These
plants were again taken from the field and transplanted into the
greenhouse for the winter. Although growth appeared normal
and symptoms of the mosaic disease could not be detected with
certainty, experiments showed repeatedly that the infective prin-
ciple of the disease was still present in the expressed sap. Allard
further shows that when scions of the immune species, NV. glutinosa,
are grafted upon mosaic-diseased plants of N. Tabacum the infec-
tive principle of the disease may pass into N. glutinosa without
the subsequent development of symptoms in it.
These experiments bear an interesting relation to my own obser-
vations on the alkekengi as a carrier of the disease. A so-called
carrier of the disease is, first, an organism which through acquired
toleration continues to harbor the germ or virus after recovery
from the diseased condition; or second, one which through natural
toleration may acquire and transmit the germ or virus without
itself showing any symptoms or suffering in any way from the
disease. A third type of carrier is found in those organisms which
are immune, but may passively transmit a disease (as does A butilon
arboreum in Baur’s grafting experiments), without necessarily
becoming a seat for the multiplication of the germ or the increase
of the virus. See Baur (1906).
MATERIAL AND METHODS
I have used two species of plants as a source of the mosaic
disease, namely:
1. Apple of Sodom (Solanum aculeatissimum). The first
plant of this species showing the mosaic disease was brought from
Florida by Professor Harper in December, 1915. So far as I am
aware this plant has not hitherto been reported as showing mosaic,
though there is of course every reason to suppose that most of
the species of Solanum are susceptible. The symptoms in apple
of Sodom include all those enumerated above.
2. Tobacco (Nicotiana Tabacum). I have used the juice of
mosaiced plants of the tobacco, kept as stock cultures in the green-
house of Columbia University.
Sac NISHIMURA: A CARRIER OF THE MOSAIC DISEASE
The juice for inoculation was procured by crushing the leaves.
The inoculations were made in several different ways as follows:
No. 1. One stab in each node. In addition to the puncture
the four upper leaves were rubbed on both surfaces with the same
inoculum used for the needle stab.
No, 2. Five stabs on stem in the following places: one at the
tip, one at the base, and three at equal distances between tip and
base. Besides these punctures the five upper leaves were rubbed
on both surfaces with the same inoculum as used for the stabs.
No. 3. Four stabs: one in the growing tip of stem, and three
additional punctures on each in the base of the three upper young
leaves near the midrib.
No. 4. Four stabs: one in growing tip of stem, one in middle
of stem, a third in its base, and a fourth on the youngest leaf near
the midrib.
No. 5. Three stabs on the stem: one in growing tip, one in the
middle, and another one near the base of the stem.
No. 6. Four stabs: one in the growing tip of the stem, another
in the middle of the stem, the third and fourth on the base of the
upper two young leaves near the midrib.
No. 7. One stab in the growing tip of the stem.
No. 8. Two stabs: one in the middle of the stem and another
in the blade of the youngest leaf.
EXPERIMENTS
1. Inoculation of the alkekengi with juice of infected apple of
Sodom plants.
On May 21, forty-eight young plants (3-5 inches high) of
the alkekengi were transplanted into five and a half inch pots
from the garden and brought into the greenhouse. These were
entirely free from mosaic disease and produced fine new growths
in two weeks’ time.
On June 5, nine plants out of the forty-eight were inoculated
with the juice of S. aculeatissimum having the mosaic disease.
_I used method No. 1 in this experiment. No symptoms of the
disease appeared in any of these plants. They were entirely indis-
tinguishable from the controls. On June 27 these same nine
: is — were inoculated again by the same method as before. On
NISHIMURA: A CARRIER OF THE MOSAIC DISEASE 223
July 16 each plant was examined carefully, but there was no
visible evidence of the disease.
On June 25 another series of inoculations with juice of diseased
tobacco plants was made on fourteen healthy plants of the alke-
kengi. These plants were inoculated by method No. 2. After
four weeks, although growth continued normally, they showed no
symptoms of the disease.
The same experiment was repeated with another series of
mature plants of the alkekengi (10-12 inches high). On August:
4, twenty of these plants were inoculated with the juice from
diseased S. aculeatissimum, another series of twenty plants were
inoculated with the juice of diseased N. Tabacum. Method of
inoculation No. 2 was used. After a period of thirty days no
symptoms of the disease had appeared on the plants in these two
series of experiments. See FIG. 1. :
2. Transfer of mosaic from the apple of Sodom to the tava and
then to tobacco
It occurred to me that the plants might carry the eee
though not showing symptoms, and I inoculated plants of N.
Tabacum and apple of Sodom with the juice from the alkekengi
plants which had been inoculated from the apple of Sodom appar-
ently without results.
TABLE I
SUMMARY OF EXPERIMENT No. 2
No. thc wes First symptom | Bom ge 25 days after inoculation
1 | July 16 | July 23, mosaic on the first new leaf | 7 | Disease See
aor | No reacti
a oe Jul y 24, mosaic on the ees new talnars 8 Disease eicleoing
4 er July 25, mosaic on the first sh 9 bd
s ieee _ July 24, mosaic on the first n 8
6 ‘ec ‘ee nye
EO nes | | July 23, mosaic on the first new leaf | 7 | Di isease co oelayiy
oS es | | July 24, mosaic on the os t new — f | 8
2 tas elas | July 25, mosaic on the first 9 a
i eee ee | Nor
II ey | July 23, mosaic on the first new leat | 7 | Disease ae developing
12 aa ors | No rea:
I 3 i os ae | aa
On July 16 thirteen young healthy plants of N. T abacum
(six- to ten-leaf stage) were inoculated with the juice of the P.
224 NISHIMURA: A CARRIER OF THE MOSAIC DISEASE
Alkekengi which had been inoculated on June 5 and June 27 with
the juice of mosaiced leaves of apple of Sodom. In this experi-
ment inoculation method No. 3 was used. July 23, experimental
plants 1, 7 and 11 showed mosaic on the first new leaves which
were just appearing. July 24, plants 3, 5 and 8 also showed
mosaic on the first new leaves. Finally on July 25, plants 4 and 9
exhibited mosaic on the first new leaves and these plants con-
tinued diseased, but plants 2, 6, 10, 12 and 13 did not show any
symptoms whatever. Eight plants were left as controls and at
the end of twenty-five days they showed no ) symptoms of mosaic:
See TABLE I and Fic. 2.
3. Transfer of mosaic from the tobacco to the alkekengi and then back
to the tobacco
I next tested plants of the alkekengi of the series which had
been inoculated with mosaic from diseased tobacco plants with
respect to their power to infect tobacco.
July 22, six young healthy tobacco plants (six- to eight-leaf
stage) were inoculated with the juice of the alkekengi which showed
no symptoms of the disease, although they had been inoculated
with the juice of diseased tobacco on June 25. This inoculation
was made by method No. 3. July 28, plant 3 showed mosaic
on the first new leaf. July 29, plant 1 showed mosaic on the first
new leaf. July 30, plants 2 and 5 showed mosaic on the first new
leaf, but plants 4 and 6 did not show symptoms of the disease.
For this experiment six control plants were kept, which after
twenty-five days showed no mosaic.
The experiment was repeated in the period from August 8 to
September 2.
August 8, twelve young healthy tobacco plants (six- to ten-
leaf stage) were inoculated by method No. 4. August 15, plant
10 showed mosaic on the first new leaf. August 16, plants 7 and
14 showed mosaic on the first new leaves. August 17, plants 9,
II, 15 and 16 showed mosaic on the first new leaves. August 18
plants 12 and 18 showed mosaic on the first new leaves. August
19, plant 17 showed mosaic on the first new leaf.
All of the inoculated plants developed the disease except plants
8 and 13 which did not show any symptoms. In this experiment
NISHIMURA: A CARRIER OF THE MOSAIC DISEASE 225
twelve plants were kept as controls and after twenty-five days
they were found to be free from any symptoms of mosaic. See
TABLE II.
TABLE II
SUMMARY OF EXPERIMENT NO. 3
No. erent | First symptoms a 25 days =. inocula-
I | July 22 July 20, mosaic on the first new leaf 7 Disease developing
2] ** * July 30, mosaic on the first new leaf | 8 eS =
3 ‘“ “| July 28, mosaic on the first new leaf 6 as "
OS poe leas, No reaction
5 | ** | July 30, mosaic on the first new leaf | 8 Disease developing
is gah | No reaction
7 | Aug. 8) Aug. 16, mosaic on the first new leaf 8 Disease developing
_o ehehe eee No reaction
9 | «| Aug. 17, mosaic on the first new leaf 9 Disease developing
ro | ‘ ‘| Aug. 15, mosaic on the first new leaf 7 * =
ir | ‘“ ‘ Aug. 17, mosaic on the first new leaf 9 “5 SG
12) “ “| Aug. 18, mosaic on the first new leaf | 10 2 “
6 A eather No reaction
14 “| Aug. 16, mosaic on the first new leaf | 8 Disease developing
15 | “Aug. 17, mosaic on the first new leaf | 9 a S
16 | ae ae | ae ae ee ae oe “oe ee oe | 9 ae “
I7 | «| Aug. 19, mosaic on the first new leaf | II - vs
18 | ‘“ ‘ Aug. 18, mosaic on the first new leaf _ Bae) es i
4. Inoculation of the tobacco with juice of healthy alkekengi plants
To prove that the juice of uninoculated plants of the alkekengi
will not produce the disease in tobacco twelve healthy young
N. Tabacum plants (ten- to eleven-leaf stage) were inoculated on
September ro with the juice of the leaves of healthy P. Alkekengi
by method No. 3, the same which was used in experiment No. 2.
After thirty days two new leaves or more appeared on each plant
but did not show any symptoms of the disease.
5. Comparison of inculation periods
To compare the length of the incubation period in my transfers
through P. Alkekengi with the ordinary incubation period when
tobacco is inoculated with juice from diseased tobacco, a series of
inoculations was made using methods Nos. 4, 7, and 8; twenty .
plants of NV. Tabacum were used in each of the three series. The
first series were inoculated on March 28 by method No. 8, the plants
having eight to ten leaves. The second series were inoculated
on April 5 by method No. 7, the plants having five to seven leaves.
226 NISHIMURA: A CARRIER OF THE MOSAIC DISEASE
The third series were inoculated on August 10 by method No. 4, the
plants having seven to eight leaves. Forty-eight of these showed
the symptoms of disease. The incubation period was from six to
seven days in each of the series. The mosaic symptoms always
appeared on the first or second new leaves that were formed after
inoculation. Fifteen controls were kept in each experiment and
after twenty-five days they were found to be in a healthy condition.
The incubation period is seen to be much the same in all the ex-
periments:
Pee: experiment No. 2, from 7 to 9 days;
In experiment No. 3, from 7 to 11 days;
In experiment No. 5, from 6 to 10 days.
This result agrees with the observations of other authors on
the length of the incubation period. In each experiment the
mosaic was observed on the first or second new leaf which appeared
after inoculation. :
The appearance of the mosaic leaves of apple of Sodom and
tobacco, infected through the alkekengi, is the same in design and
coloring (i. e., distribution of the dark green with relation to the
vein, etc.) as in direct infection from tobacco to tobacco.
There can be no question that the virus induces the same symp-
toms after passing through the apparently immune plant as when
transferred directly from one susceptible plant to another. The
affected plants developed the disease on all new leaves which
appeared after inoculation.
6. Transfer of mosaic from the apple of Sodom to the alkekengi and
then back to the apple of Sodom
The transfer through P. Alkekengi from apple of Sodom to
apple of Sodom is equally easy.
June 23, ten young healthy plants of apple of Sodom (3-4
inches high) were inoculated with the juice from the leaves of the
- same plants of P. Alkekengi (inoculated from apple of Sodom)
which were used in experiment No. 2 as a source for the inoculation
of tobacco. I used method No. 5 for this inoculation. June 30,
plants 2 and 6 showed mosaic on the first new leaves. July 1,
plant 4 showed mosaic on the first new leaf. July 2, plants 7 and
NISHIMURA: A CARRIER OF THE MOSAIC DISEASE 227
10 showed mosaic on the first new leaves. July 3, plant 1 showed
mosaic on the first new leaf. These six plants developed the
disease, but plants 3, 5, 8 and 9 showed no symptoms.
On July 20, the same experiment was repeated with easter
series of ten young healthy plants of apple of Sodom 4-5 inches
high. July 29, plant 12 showed mosaic on the first new leaf.
July 30, plants 11 and 13 showed mosaic on the first new leaves.
July 31, plants 14 and 16 showed mosaic on the first new leaves.
August I, plants 19 and 20 showed mosaic on the second new
leaves. Plants 15, 17 and 18 showed no symptoms.
Ten plants were kept in each experiment as controls and after
twenty-five days they were found to be in healthy condition.
TABLE III
SUMMARY OF EXPERIMENT No. 6
No Date of First symptom pen | 25 days after inoculation
“| inoculation j Sa days, 5
1 | June 23. July 3, mosaic on the first new leaf 10 Disease developing
2 : «| June 30, mosaic on the first new leaf | . - :
3 oe ee No rea
4 “| July 1, mosaic on the first new leaf 8 Hissaae a
5 hr etaly No reaction
6 ** | June 30, mosaic on the first new leaf +4 Di isease developing
7 *« | July 2, mosaic on the first new leaf 9
8 i ee No reaction
- «« «| July 2, mosaic on the first new leaf 9 Disease developing
11 | July 20 | July 30, mosaic on the first new le bas) ge
I July 29, mosaic on first new leaf 9 ue *
13 «* | July 30, mosaic on the first new leaf 10 i .
14 «| July 31, mosaic on the first new leaf II - *
I5 ae es No react
16 «July 31, mosaic on the first new leaf xi Disease developing
¥ 7 “sé “ee o No react
1s ae ae
19 « | Aug. 1, mosaic on the second new
lea. 12 Disease developing
20 « « | Aug. 1, mosaic on the second new
leaf 12 ae oe
7. Inoculation of the apple of Sodom with the juice of healthy alke-
kengi plants
To prove that the juice of uninoculated plants of P. Alkekengi
will not produce the disease in the apple of Sodom, twelve healthy
young plants of apple of Sodom (5-6 inches high) were inoculated
on September 12 with the juice of the leaves of healthy P. Alke-
228 NISHIMURA: A CARRIER OF THE MOSAIC DISEASE
kengi by the method used in experiment No.6. After a period of
four weeks two or three new leaves had appeared on each plant but
no symptoms of the disease had appeared.
8. Transfer of mosaic from the tobacco to the alkekengi and then to
the apple of Sodom
To make the evidence still more clear that P. Alkekengi is a
carrier, I tried an inoculation of the apple of Sodom with the
juice of P. Alkekengi used in experiment No. 3, that is from plants
inoculated from tobacco. The experiment is the reverse of experi-
ment No. 2.
August 20, twelve young healthy plants of apple of Sodom
(4-5 inches high) were inoculated with the juice of the leaves of
P. Alkekengi which had been inoculated with the juice of mosaic
leaves of N. Tabacum. Four punctures were made in each plant
by method No. 6. August 30, plants 2, 3 and 10 showed mosaic
on the first new leaves. August 31, plants 1, 9 and 11 also showed
mosaic on the first new leaves. September 1, plant 5 showed
mosaic on the second new leaf. September 2, plant 7 showed
mosaic on the second new leaf. These eight plants developed the
disease, but plants 4, 6, 8 and 12 did not show any symptoms.
TABLE IV
SUMMARY OF EXPERIMENT No. 8
No. Plies Seal First symptoms oe ave 25 days after incubation
r | Aug. me Aug. 31, mosaic on the first new leaf II Disease developing
2 = mel Aug. 30, mosaic on the first new ew leaf 10
3 ee ‘ “ ro “a “
4 = - No reaction
Ce eat ** Sept. 1, mosaic on the second new leaf, 12 Disease developing
6 av se No reacti
q se ** Sept. 2, mosaic on the second new leaf 13 Disease developing
8 - ce No reaction
9 “| Aug. 31, mosaic on the first new wie II Disease developing
ro ie So Pood 3 mosaic on the first new le: 10 ae te
II “ “| Aug e first new haat It “
I2 eee No reaction
In this experiment twelve controls were kept and after twenty-
five days they were found to be in healthy condition. See TABLE
IV and Fic. 3.
NISHIMURA: A CARRIER OF THE MOSAIC DISEASE 229
9. Inoculation of the apple of Sodom with juice of infected tobacco
plants
Before making the above experiments with the alkeKengi I
had already satisfied myself as to the relation of the mosaic of
tobacco to that of the apple of Sodom.
Forty young healthy plants of apple of Sodom (4-6 inches
high) were used in each of the two series. The first series was
inoculated on March 27 with juice from diseased apple of Sodom.
The second series was inoculated on March 25 with juice from
diseased tobacco. These inoculations were made by method
No. 5. In the first series thirty-three plants showed mosaic.
The incubation period was from eight to ten days. In the second
series thirty-six plants showed mosaic. The. incubation period
was from seven to twelve days. In every case the mosaic symp-
toms appeared on the first or second new leaves that were formed
after inoculation. The incubation period was much the same in
all these experiments. Twenty-five controls were kept and after
thirty days they were found to be in healthy condition.
The appearance of the mosaic leaves of apple of Sodom, pro-
duced by infection from the carrier, P. Alkekengi (which had been
inoculated from apple of Sodom or from tobacco) is the same as in
the case of direct infection from tobacco or apple of Sodom in the
distribution of the dark green with relation to the veins, in the
blistering, etc. There can be no question that the virus induces
the same symptoms after passing through the apparently immune
plant to another as when transferred directly from one susceptible
to the disease.
10. Reinoculation of tobacco plants
The N. Tabacum plants inoculated in experiments 2, 3, and 4,
which did not take the disease, were later used for further inocu-
lations with virus from the diseased tobacco which had been
inoculated from P. Alkekengi. September 10 the fourteen plants
of N. Tabacum were inoculated by method No. 1. September 18,
plants 3 and 5 showed the mosaic on the first new leaves. Sep-
tember 19, plants 7 and 11 also showed the mosaic on the first new
leaves. September 20, plants 8, 10 and 12 showed the mosaic.
September 21, plants 2, 6 and 9 showed the mosaic. September
230 NISHIMURA: A CARRIER OF THE MOSAIC DISEASE
22, plant 13 showed the mosaic on the second new leaf. These
eleven inoculated plants developed the disease, but plants 1, 4
and 14 did not show any symptoms.
It is evident that the failure to infect these plants in the earlier
inoculations was due to the uncertainties of the method of inocu-
lation. Twelve of these same plants were kept as controls (four
plants which had been used in experiment No. 2, five plants from
experiment No. 3 and the other three plants from experiment
No. 4). All of the controls but one remained in a healthy condi-
tion. The results show that the chances of accidental inoculations
in the greenhouse where the work was carried on were very small.
These control plants with one exception had remained uninfected
for over two months.
11. Reinoculation of apple of Sodom plants
The same test was made on the plants of the apple of Sodom
which had remained healthy after inoculation in the earlier
experiments. }
September 13, the twelve plants of apple of Sodom of experi-
ments 6, 8 and 9, which had failed to develope the disease were
inoculated by method No. 6 with juice from diseased tobacco
which had been inoculated from P. Alkekengi. September 21,
plants 8 and 11 showed the mosaic on the first new leaves. Sep-
tember 22, plants 7 and 6 showed the mosaic as above. Septem-
ber 23, plants 2, 4 and 5 showed the mosaic as above. September
24, plant 9 showed the mosaic as above. September 25, plant
12 showed the mosaic on the second new leaf.
Nine plants developed the disease, but plants, 1, 3 and 10 did
not show any symptoms.
SUMMARY
Physalis Alkekengi is found to be a carrier of the mosaic disease
without itself showing any symptoms. The experimental results
were as follows:
1. Thirty-four healthy plants of Physalis Alkekengi were
inoculated with juice of mosaic-diseased plants of N. Tabacum
and showed nosymptoms. To test the method of inoculation and
the virulence of the disease sixty young healthy plants of N.
NISHIMURA: A CARRIER OF THE MOSAIC DISEASE 231
Tabacum were inoculated with juice of the mosaic-diseased tobacco
and forty-eight of these plants showed the disease.
2. The same test was made with the juice of mosaic-diseased
Solanum aculeatissimum on twenty-nine healthy plants of P.
Alkekengi, and none showed any symptoms of the disease. Of
forty healthy plants of S. aculeatissimum inoculated with juice of
mosaic-diseased S. aculeatissimum thirty-three plants showed the
disease.
3. Of thirty-one healthy plants of N. Tabacum inoculated with
juice from leaves of P. Alkekengi which showed no symptoms of
mosaic but had been inoculated from diseased tobacco or S. acu-
leatissimum, twenty-two plants showed typical mosaic disease.
Of twelve young healthy plants of N. Tabacum inoculated with
juice from leaves of P. Alkekengi which had not been inoculated
none showed the disease.
4. Of thirty-two healthy plants of S. aculeatissimum inocu-
lated with juice from leaves of P. Alkekengi which showed no
symptoms of mosaic but had been inoculated from diseased
tobacco or S. aculeatissimum, twenty-one plants showed’ mosaic
also. Of twelve young healthy plants of S. aculeatissimum
inoculated with juice from leaves of P. Alkekengi which had not
been inoculated, none showed the disease.
5. Experiments have shown that the mosaic diseases of S.
aculeatissimum and N. Tabacum are similar with respect to the
symptoms and incubation periods.
6. The juice from diseased N. Tabacum when passed through
P. Alkekengi to N. Tabacum has produced the disease,in 80 per
cent. of the inoculations on healthy plants of S. aculeatissimum or
N. Tabacum. It is evident that there is no marked weakening
of the virus by the carrier P. Alkekengt.
The author is especially indebted to Professor Harper of
Columbia University for his kindly interest and valued super-
vision throughout these experiments.
DEPARTMENT, OF BOTANY,
COLUMBIA UNIVERSITY
232 NISHIMURA: A CARRIER OF THE MOSAIC DISEASE
LITERATURE CITED
Allard, H. A. (1914). The mosaic disease of tobacco. U.S. Dept.
Agr. Bull. 40: 1-33.
(1914). A review of investigations of the mosaic disease of
tobacco, together with a bibliography of the more important con-
tributions. Bull. Torrey Club 41: 435-458.
(1915). Distribution of the virus of the mosaic disease in
capsules, filaments, anther, and pistils of affected tobacco plants.
(1916). The mosaic disease of tomatoes and petunias.
Phytopathology 6: 328-336.
(1917). The mosaic disease of tobacco. Jour. Agr. Res. 10:
zungsber. K. Preus. Akad. Wiss. Math.-Naturw. Kl. 32: 11-29.
Chapman, G. H. (1913). ‘ Mosaic’’ and allied diseases with especial
reference to tobacco and tomatoes. Rept. Mass. Agr. Exp. Sta.
of tobacco. Science II. 44: 537-538.
Clinton, G. P. (1915). Chlorosis of plants with special reference to
calico of tobacco. Conn. Agr. Exp. Sta. Rept. 1914: 537-424.
pl. 25-32.
Freiberg, G. W. (1917). Studies in the mosaic disease of plants. Ann.
Missouri Bot. Gard. 4: 175-232.
Hunger, F. W. T. (1905). Untersuchungen und Betrachtungen itiber
die Mosaik-Krankheit der Tabakspflanze. Zeitschr. Pflanzenkr. 15:
257-311.
Iwanomski, D. (1903). Ueber die Mosaikkrankheit der Tabakspflanze.
Ibid. 13: 2-41.
Jensen, H. (1913). Mosaick-Zcikta. Mededeelingen von ‘Het proef
station voor Vorstenlandsche Tabak”’ 5: 61-67.
Lowe, O. (1900). Remarks on the mosaic disease of the tobacco plant.
U.S. Dept. Agr. Rept. 65: 24-27.
Mayer, A. (1886). Ueber die Mosaik krankheit des Tabaks. Landw.
Versuchs Stat. 22: 450-467.
Peters, L. (1912). Krankheiten und Beschadigungen des Tabaks.
Mosaikkrankheit Mitteil. Kais. Biol. Anst. Land. Forstrw. 13: [58].
Sturgis, W.C. (1900). On the effects on tobacco of shading and applica-
tion of lime. Conn. Agr. Exp. Sta. Rept. 23: 259-261.
Westerdijk, J. (1910). Die Mosaikkrankheit der Tomaten. Phyto-
path. Lab. ‘“‘ Willie Commelin Scholten,’’ Mededeel. 1: 1-20.
NISHIMURA: A CARRIER OF THE MOSAIC DISEASE 233
Woods, A. F. (1899). The destruction of chlorophyll by oxidizing
enzyms. Centralbl. Bakt. Abth. II. 5: 745-754.
(1902). Observations on the mosaic disease of tobacco.
U. S. Dept. Agr. Bur. Pl. Ind. Bull. 18: 1-24. pl. 1-
Explanation of plate 7
Fic. 1. Typical leaf of Physalis Alkekengi, the carrier of the mosaic disease,
Ldaiictiaiee! with the juice of the leaves of eran aculeatissimum having mosaic
sen but not sea any symptoms of the
be cal mosaic leaf of Nicotiana kes inoculated with the juice
of Pe etionee - ian Alkekengi which had been inoculated with the juice of
mosaic ssa of 7 lanum aculeatissim
Fic mosaic leaf of Sa aculeatissimum, inoculated with the
juice of ee Alkekengi which had been inoculated with the juice of mosaic
leaves of Nicotiana Tabacum.
The air chambers of Grimaldia fragrans *
ALEXANDER W. EVANS
(WITH FOURTEEN TEXT FIGURES)
INTRODUCTION
In most of the Marchantiales the thallus shows a layer of
green tissue with air spaces or chambers below the dorsal epidermis.
These chambers exhibit many differences when the group as a
whole is considered, but it is possible to refer the majority to three
distinct types. To these the names of the representative genera
Riccia, Reboulia and Marchantia may be applied.
In the Riccia type the chambers occupy a single layer and are
in the form of canals with their long axes approximately vertical;
the canals are usually narrow and bounded by only four rows of
cells, but in certain cases they are broader and bounded by a
greater number of cells. In the Reboulia type the chambers are
in two or more layers (at least in the median portion of the thallus)
and are in the form of irregular polyhedrons, often tending to be
isodiametric; this type is sometimes complicated by cellular out-
growths into the chambers. In the Marchantia type the chambers
are again in a single layer (as in the Riccia type) but are in the
form of more or less flattened polygonal prisms with their longer
dimensions approximately horizontal; they are further distin-
guished by the presence of numerous simple or branched green
filaments, extending from the floors of the chambers nearly or
quite to the epidermis. In all three types the chambers communi-
cate with the outside air by means of openings in the epidermis.
In the Riccia type these may be nothing more than continuations
of the canalicular chambers, but in the two other types the open-
ings are usually surrounded by specialized epidermal cells and
form the characteristic air pores or epidermal pores of the group.
Among North American genera the Riccia type is restricted to
Riccia and Oxymitra; the Reboulia type is found in Ricciella,
* Contribution from the Osborn Botanical Laboratory.
235
236 Evans: ArR CHAMBERS OF GRIMALDIA FRAGRANS
Ricciocarpus, Peltolepis, Sauteria, Clevea, Plagiochasma, Reboulia,
Grimaldia, Neesiella, Cryptomitrium, Asterella and Bucegia; while
the Marchantia type occurs in Corsinia, Targionia, Conocephalum,
Lunularia, Preissia and Marchantia. The reduced air chambers
of Cyathodium conform best perhaps to the Marchantia type, in
spite of the absence of green filaments, while the adult thallus of
Dumortiera lacks air chambers altogether.
The genus Grimaldia Raddi, as understood by most recent
writers, contains about half a dozen species. The most widely
distributed of these is G. fragrans (Balb.) Corda, which is found in
Europe, Asia and North America. Other well-known species,
closely related to G. fragrans, are the Mediterranean G. dichotoma
Raddi and the Californian G. californica Gottsche. In the eastern
parts of the United States G. fragrans is sometimes locally abun-
dant, preferring sunny trap ridges and growing on earth among
rocks, rather than on the rocks themselves. It was in such a
locality as this, on West Rock Ridge, near New Haven, Connecti-
cut, that the material used in the present study was collected.
The narrow thallus is firm and compact and produces an abun-
dance of purple ventral scales with bleached-out appendages.
The upper surface is grayish green and shows no indications of the
boundaries of the air chambers beneath the epidermis. The
margins, as well as the ventral surface, are more or less pigmented
with purple. The species is markedly xerophytic, the margins
becoming involute when dry, thus covering over and protecting
the upper surface.
THE AIR CHAMBERS OF THE MATURE THALLUS
The green tissue of the thallus in Grimaldia has been repeatedly
described, most of the observations having been based on either
G. fragrans or G. dichotoma. Unfortunately the descriptions show
marked discrepancies. Stephani (11), for example, states that
the air chambers are densely filled with erect green filaments
composed of long cylindrical cells, and K. Miiller (6, p. 259)
notes the presence of vertical plates of cells in addition to the
filaments. Schiffner (9, p. 309) criticizes these descriptions.
According to his account the chambers of Grimaldia undergo a
secondary partitioning by means of irregular green lamellae which
Evans: AIR CHAMBERS OF GRIMALDIA FRAGRANS Paes
grow upward from the floors and lateral walls of the chambers.
A spongy tissue is thus formed in which narrow air spaces run,
scarcely broader than the thickness of the lamellae, and the original
partitions of the chambers soon become unrecognizable. He
admits that in section the plates of cells one cell thick look like
filaments and that marginal cells of the plates sometimes project
as teeth, but he maintains that actual filaments are never present
and that this fact is at once made evident by sections of the green
tissue cut parallel with the surface of the thallus. Massalongo
(5, p. 730), on the other hand, agrees with Stephani and states that
the chambers are filled with vertical uniseriate filaments, some of
Fic. 1. Transverse section through epidermis and green tissue, X 270 A
air chambers; e-g, apparent filaments; h, plate-like outgrowth; 7, k, 1, boundaries
between chambers.
them reaching the epidermis. His figures not only show filaments
clearly but indicate that the boundaries of the chambers are
distinct, in this respect also differing from Schiffner’s account.
The green tissue of G. fragrans is so compact that it is difficult
to make out its true structure from ordinary hand sections. Even
microtome sections are not always easy to interpret, but they give
a much clearer idea of the complex arrangement of the cells and
of the intricate system of aérating chambers and help to explain
some of the conflicting statements in the published descriptions.
In a transverse section, such as the one shown in Fie. 1, the cham-
238 Evans: AIR CHAMBERS OF GRIMALDIA FRAGRANS
bers are seen to be in three or four layers in the thickened median
portion of the thallus. As the margins are approached the thallus
becomes thinner, and the number of layers decreases until only
the uppermost layer is left. Except in this uppermost layer the
chambers are usually polygonal in outline and tend to be isodia-
metric. In the uppermost layer they tend to be elongated verti-
cally, as shown in the spaces a and 6. That the spaces communi-
cate with one another is also indicated in the figure. The space
c, for example, is connected with a space nearer the epidermis, and
the space d probably represents a passageway to a chamber in
another section. The figure seems, at first sight, to confirm the
statements made by Miiller, that both filaments and cell plates
are present. Immediately beneath the pore there are apparently
three filamentous outgrowths, e, f, and g, and a plate-like outgrowth
is clearly shown at h. Of course, as Schiffner intimates, apparent
filaments may be nothing more than sections of cell plates. In
the section drawn careful focusing does indeed show that e and f
are in close contact with another apparent filament in another
plane, and the same thing is true of other apparent filaments in
the section. Some of the cell plates, moreover, appear to have a’
fluted surface, so that a section cut parallel with the surface of the
plate might readily give the impression of a series of filaments.
At the same time there are many apparent filaments which seem
to be entirely free from one another, and it is impossible to deter-
mine their true status except by the study of other sections. It
will be noted that the more deeply situated chambers are free or
nearly so from outgrowths of any kind.
The figure is of further interest in showing that some of the
apparent filaments and plate-like outgrowths end freely in the
chamber without reaching the epidermis, this being especially
true in the vicinity of the pores; others, as shown by the one be-
tween the spaces a and b, extend to the very epidermis and seem
to be connected withit. It is doubtful, however, if the connection
is ever anything more than a close contact, such as the free fila-
ments in Marchantia and Conocephalum often exhibit. No
instance has been observed where an outgrowth extends downward
from the epidermis and ends freely in a chamber, and there is no
adequate evidence that the epidermal cells themselves ever give
Evans: AIR CHAMBERS OF GRIMALDIA FRAGRANS 239
rise to outgrowths, as Schiffner suggests may be the case. The
original boundaries of the dorsal air chambers are not absolutely
unrecognizable, but they are by no means as distinct as Massa-
_ longo’s figure represents them. In Fic. 1 the boundaries of the
chamber with the air-pore are shown at i and k, while another
boundary is situated at I.
The longitudinal section drawn (Fic. 2) brings out the fact
that many of the air chambers are more or less elongated. This
is strikingly true of those most deeply situated but is also well
. Fic. 2. Longitudinal section through epidermis and green tissue, X 270.
shown by the chamber with the air-pore, although the actual
boundaries of this chamber are not definitely indicated. It will
be noted that the upper margin of the cell plate represented, which
extends almost longitudinally beneath the pore, is distinctly
dentate, some of the teeth being over a cell in length. This
accords, on the whole, with Schiffner’s statement that the marginal
cells of the plates may project as teeth. Although some of the
teeth shown are more than projecting cells, it would be a stretch
of the term to describe them as filaments. The figure, therefore,
presents no evidence of the occurrence of true filaments. Other
sections, however, show apparent filaments, similar to those
represented in Fic. 1.
According to Schiffner, a section through the green tissue paral-
lel with the surface of the thallus will at once show that the cham-
bers are destitute of free filaments. Fic. 3 shows a part of sucha
240 Evans: AIR CHAMBERS OF GRIMALDIA FRAGRANS
section, cut immediately below the epidermis, and seems at first
to belie his statement. The figure shows the partitions, almost
complete, of an air chamber, the cells being distinguished by stip-
pling. Only one end of the chamber is represented; the other end
did not show because the section was slightly oblique in that
region and passed through the epidermis instead of the green
tissue beneath. That the stippled cells represent the boundaries
of a chamber is evident from their close union and also from the
b
Fic. 3. Section parallel with the surface, just below epidermis, X 270.
Fic. 4. Section a short distance below the one shown in Fic. 3, X 270.
fact that an epidermal pore was situated above the middle of the
space which they enclose. It will be seen that the chamber con-
tains a number of cells, circular in section and either entirely free
or else loosely connected with one another or with the cells of the
partitions. Similar cells are shown elsewhere in the figure, and a
superficial examination would interpret them as the sections of
filaments, especially if they were considered in connection with
Fic: 1.
The incorrectness of this interpretation is brought out by a
comparison with Fic. 4, which shows the same chamber at a
lower level, the cells of the partitions being again indicated by
stippling. In this figure the complete boundaries of the chamber
Evans: AIR CHAMBERS OF GRIMALDIA FRAGRANS 241
» left
diit
are shown, but the cells enclosed present a very different app
They are not only much more numerous but are, with a few excep-
tions, more or less firmly united, and the entire chamber is thus
divided up into smaller chambers, some of which seem entirely
cut off while others show their connections with other chambers.
The seven cells shown on the right of the left-hand partition in
FIG. 3 are represented in Fic. 4 by seven united cells, showing at
once that these seven cells are not the cross sections of filaments
but simply the cross sections of teeth, like those shown in Fic. 2.
Similar conclusions would be created by comparing other apparently
free cells in Fic. 3 with their representatives in Fic. 4. It thus
becomes established that there are no free filaments in the cham-
bers. It will be noted further that Fic. 4 presents a much more
complicated condition than Fic. 3 and that the boundaries of the
air chamber would be hardly
distinguishable except through
comparison with the simpler -fig-
ure. It is probable that a sec-
tion like the one shown in FIG. 4
was responsible for Schiffner’s
statements, which it certainly
strongly supports.
In Fic. 5 a section from
another thallus is shown, cut at
a still lower level. This section
shows a loose spongy tissue, two
of the chambers being connected
by a passageway. Cellular out-
growths are very infrequent, but
a single cell, apparently free, is ae ‘
shown in one of the chambers, ee a ae ling
and a single short outgrowth in pers, » 270.
another. When compared with
Fic. 4 the spaces are relatively larger and fewer and the tissue in
consequence much less compact. Sections cut farther down show
elongated spaces, similar to those represented in Fic. 2, while
sections beneath these show the ventral parenchymatous tissue
without spaces of any sort.
Bt ok
3
242 Evans: AIR CHAMBERS OF GRIMALDIA FRAGRANS
It is clear from a comparison of transverse, longitudinal and
horizontal sections that a distinction may be made between the
dorsal layer-of air chambers in Grimaldia and the more deeply
situated layers. In the dorsal layer the original chambers show a
secondary partitioning by a system of more or less vertical cell
plates, the free margins of which sometimes bear scattered teeth,
apparently always less than two cells long. Except for these
teeth the chambers lack filaments completely. In the more
deeply situated layers, the chambers are much simpler and rarely
show evidences of any kind of outgrowth. These conclusions
show the incorrectness of certain statements made by Stephani,
K. Miiller, and Massalongo and the essential correctness of Schiff-
ner’s account.
The complex conditions found in the green tissue of Grimaldia
are duplicated by Plagiochasma and by certain species of As-
terella. The other genera showing the Reboulia type of air cham-
ber have a much looser green tissue, the secondary partitioning
being-less highly developed or absent altogether.
ORIGIN AND ENLARGEMENT OF THE AIR CHAMBERS
The development of the air chambers in the Marchantiales
has aroused a good deal of interest among students of the Hepati-
cae, and the history of the subject is fully given by Barnes and
Land (1). The explanation which they advance to account for
the origin of the chambers differs in certain respects from the older
explanation advanced by Leitgeb and accepted by many of his
successors. Leitgeb’s explanation was based primarily on his
study of Riccia, but he extended its application to the more com-
plex genera. According to his ideas the air chambers do not
originate in compact tissue, and no splitting of cell walls is involved
in their formation. They arise, rather, on the surface of a young
thallus and are due to a cessation of upward growth in certain
jimited areas, the surrounding parts growing upward vigorously.
The areas where growth is supposed to cease are situated in most
cases where four of the surface cells come together; they mark the
lower ends of the chambers, the vertical extent of which depends
upon the degree of upward growth which the surrounding parts
exhibit.
Evans: AIR CHAMBERS OF GRIMALDIA FRAGRANS 243
According to Barnes and Land there is nothing to support
Leitgeb’s views. In their opinion the chambers always originate
in compact tissue below the surface of the young thallus by a
splitting of cell-walls and, in case the mature chamber has an
epidermal pore, the splitting extends upward until the surface is
reached. They based their conclusions on a study of Ricciella
(Riccia fluitans L.), Ricciocarpus (Riccia natans L.), Marchantia,
Lunularia, Conocephalum, Dumortiera, Asterella (Fimbriaria),
and Plagiochasma, and assumed that they would apply as well to
Riccia (in its restricted sense). After the chambers are once
established their increase in size need not involve any further
schizogenous processes. It is largely brought about by the growth
of the cells surrounding the chamber, and may be wholly brought
about in this way; in other words, by the surface extension of the
bounding cell-walls. According to Leitgeb’s ideas the increase in
the size of the chambers is brought about in much the same way,
except that a total absence of splitting is always assumed. It
will be seen, therefore, that the most important differences between -
the two explanations are concerned with the very beginning of the
developmental process: according to Leitgeb the chamber is
superficial in origin and no splitting occurs; according to Barnes
and Land the chamber is not superficial in origin and splitting does
occur.
Among recent papers dealing with air chambers, those by
Miss Hirsh (4), Pietsch (8), Deutsch (3), Miss O’Keeffe (7) and
Miss Black (2) may be briefly noted. Miss Hirsh’s work is based
largely on Ricciocarpus natans (L.) Corda and Riccia Frostii Aust.
She reaches the conclusion that the first of these species agrees
with Barnes and Land’s explanation, while the second agrees with
Leitgeb’s. Her figures of R. Frostii, however, by no means support
this conclusion fully. Although they show that the chambers
drawn may have been superficial in origin, they show as well that a
splitting must sometimes have occurred, because some of the cham-
bers extend below the original surface of the thallus. This is
brought out clearly by her f. 6, upon which she lays especial
emphasis. This figure, in fact, presents no convincing evidence
that the chamber may not have been initiated by a schizogenous
process.
244 Evans: AIR CHAMBERS OF GRIMALDIA FRAGRANS
Pietsch’s work is remarkable for its thoroughness and accuracy.
It deals with species of Riccia and Ricciella, and his account is
therefore based on the group of plants from which Leitgeb drew
his conclusions. Although he criticises the work done by Barnes
and Land, his observations lead to similar conclusions, so far as
the development of the air chambers is concerned. He finds that
even in Riccia the chambers originate from a splitting of cell walls,
the split beginning below the surface = then extending upward
until the surface is reached.
Deutsch’s paper, devoted to Targionia hypophylla L., includes
an interesting observation on the development of the air chambers.
He states that they arise by a splitting apart of cells close to the
apical cell but maintains that the split begins on the outside and
extends inward, instead of beginning below the surface and extend-
ing outward. The f. 3, which he cites as evidence, would be more
convincing if the youngest chamber shown did not extend into
the hypodermal tissues; as the figure stands it might equally well
bear the opposite interpretation from the one drawn. Deutsch
does not consider that his account differs in any essential respect
from the explanation of Barnes and Land, in spite of the super-
ficial origin which he assigns to the chambers. Miss O’Keeffe,
who also worked on Targionia, fully supports Deutsch in his
statements about the origin of the chambers. Fortunately, the
youngest chamber which she shows (f. z, A, a) seems to be con-
clusive; it appears in longitudinal section as a split between two
superficial cells and does not extend beyond them.
Miss Black’s paper deals with Riccia Frostii, one of the species
investigated by Miss Hirsh, and the same conclusions are drawn
as to the origin of the air chambers. Her f. 6, however, is open to
the same criticism as Miss Hirsh’s figures. It represents the
apical region of a thallus cut longitudinally and including five
young air chambers, but even the youngest of these projects below
the original surface, showing that a splitting of a cell wall must
have taken place. Miss Black emphasizes the fact that she ob-
served no cases in which an intercellular space appeared below the
surface and then broke through to the outside, so that her con-
clusion regarding the superficial origin of the chambers seems
justified. At the same time her figure presents no evidence that
Evans: AIR CHAMBERS OF GRIMALDIA FRAGRANS 245
the chambers may not have originated from splits between super-
ficial cells, as Deutsch and Miss O’Keeffe maintain is the case in
Targionia. ¥
The thallus of Grimaldia fragrans is so complex that it is
impracticable to trace the cell divisions which take place in the
segments cut off from the apical cell, as Pietsch has so ably done
in the case of Riccia glauca L. Fics. 6-8, however, give some idea
of the apical region and bring out the fact that a single apical cell
with four cutting faces is present. In Fic. 6, immediately above
the apical cell the meristematic tissue forms a compact mass
Fic. 6. Longitudinal section through a growing point, X 500.’ x, apical cell.
Fic. 7. Longitudinal section through another growing point, X 500. x, apical cell.
without intercellular spaces. Between the fourth and fifth cells
the first indication of a chamber appears in the form of a split a
short distance below the surface. Between the fifth and sixth cells
an older and longer chamber is visible, which has reached the
surface, apparently through the upward extension of a similar
split. The elongation and widening of the chamber have been
largely due, it would appear, to the growth of the bounding cells,
The still older chambers shown in the figure are not cut squarely
in the middle and need not be further considered.
246 Evans: AIR CHAMBERS OF GRIMALDIA FRAGRANS
In Fic. 7, which represents the apical region of another thallus,
a somewhat different condition is revealed. In this case the
first indication of an air chamber appears between the third and
fourth cells and is likewise in the form of a split, but this time the
split evidently began on the outside and extended inward. Al-
though the chamber is thus superficial in origin, there is no evidence
that a surface area has had its upward growth arrested, as Leit-
geb’s explanation demands. The split clearly extends inward
from the original surface. The chamber between the fourth and
fifth cells is considerably deeper and broader, and it is clear that
its increase in size has involved further schizogenous processes.
The next chamber shown gives evidence of a further horizontal
extension.
It would appear from these two figures that the air chambers
in Grimaldia fragrans owe their origin to a splitting of cell walls,
but that the place where the split first makes its appearance is not
always the same. It may be below the surface and extend out-
ward, in which case it agrees fully with the explanation advanced
by Barnes and Land; it may be at the surface and extend inward,
thus agreeing with Deutsch’s account of Targionia hypophylla.
In the writer’s opinion the figures published by Miss Hirsh and
Miss Black might be interpreted in the same way as FIG. 7, so
that there still seems to be no conclusive evidence that Leitgeb’s ©
explanation ever applies.
Fics. 8-10 yield further evidence as to the origin of the cham-
bers; they were all drawn from a single section, cut at right
angles to the long axis of the thallus, and show for the most
part superficial cells. In Fic. 8 the apical cell appears in the form
of a rectangle. Directly above it an air chamber reaching the
surface is shown between the fourth and fifth cells, corresponding
apparently with the chamber between the fourth and fifth cells
of Fic. 7. The schizogenous origin of this chamber seems clear,
but there is nothing to show whether the split began at or below
the surface. Between the third and fourth cells no signs of a
chamber can be discerned, although a superficial split may be
present like the one shown in Fic. 7. The figure at any rate gives
no evidence of a split beginning below the surface.
Fics. 9 and 10 are much more conclusive. They represent a
Evans: AIR CHAMBERS OF GRIMALDIA FRAGRANS 247
portion of the thallus to the right of the apical cell and derived
from lateral segments. In drawing Fic. 9 the microscope was
focused on the surface of the cells in a circumscribed area; in
drawing Fic. 10 it was focused a little below the surface of the
same area. FIG. 9 shows a series of cells in close union and two
air chambers which have reached the surface; F1c. 10 shows the
same two chambers and six additional ones. The latter clearly
represent schizogenous spaces below the surface and demonstrate
an origin like that of the youngest chamber in Fic. 6.
Fic, 8. Transverse section through a growing point, xX 500. x, apical cell.
Fic. 9. Superficial cells to the right of the apical cell shown in Fic. 8, X 500.
Fic. 10. The same region as that shown in Fic. 9, but at a slightly lower focus,
more very young dorsal pores bene anes x bh
FI Slightly older d h in Fic. 10, the section
G. II. oiightly older
parallel with the surface, X 500.
Section just below the one shown in FiG. 11, the numerous intercellular
00.
Fic. 12
spaces being the beginnings of more deeply situated chambers,
The rudimentary chambers shown in FiGs. 6-10 represent the
beginnings of the complex dorsal chambers shown in Fics. 1-4.
The later stages in the development of these chambers and the
origin and development of the more deeply situated chambers are
exceedingly difficult to follow. For a while the dorsal chambers
are distinct enough in sections cut immediately below the epider-
mis. Such a section is shown in Fic. 11, where two complete
chambers and parts of six others are represented. The increase
248 Evans: AIR CHAMBERS OF GRIMALDIA FRAGRANS
in size which these chambers show, when compared with the small
intercellular spaces in Fic. 10, is due to the vigorous growth of
the bounding cells, accompanied by rapid cell divisions. At this
stage the partitions show no evidence of outgrowths. Fic. 12
represents the section just below the one shown in Fic. 11, the
cells drawn, in part at least, forming the floors of the dorsal cham-
bers. The figure shows many intercellular spaces, -which are
clearly schizogenous in origin; these spaces represent the begin-
nings of the more deeply situated chambers or, in some cases, the
passageways leading from these chambers to the dorsal chambers.
A longitudinal section, repre-
senting about the same stages as
those shown in Fics. 11 and 12,
may be seen in Fic. 13, the left-
hand side of the figure being
toward the apical cell. The very
rapid development of the cham-
bers is clearly indicated, and light
is thrown on the way in which
the deeper chambers originate,
such chambers being indicated by
the letters a—e. It will be seen
that some of these chambers
seem to be completely enclosed,
showing that they may have originated by a splitting of cell walls
in compact tissue, and that others already communicate with
more dorsally situated chambers. Whether the connecting
passageways are always formed subsequently to the chambers,
or whether the formation of the passageways may sometimes
precede that of the chambers is not altogether clear. If the older,
right-hand side of the figure is compared with the younger, left-
hand side, it becomes evident that the tissue with intercellular
spaces has almost tripled in thickness and that the dorsal chambers
have become distinctly deeper. The rapid growth involved in
these changes has taken place in the original partitions of the
dorsal chambers, in the cells which formed their irregular floors
and in the cells immediately beneath. As the writer conceives
the process, the growth of the partitions is both horizontal and
Fic. 13. Longitudinal section through
young chambers, 500. a-ée, more
deeply situated chambers.
Evans: AIR CHAMBERS OF GRIMALDIA FRAGRANS 249
vertical, the growth in the latter direction being often equalled
by the upward growth (accompanied by cell division) of the cells
forming the floors of the chambers; these in turn remain more or
less united with one another and with the cells of the partitions
and in this way form the system of united cell-plates in the dorsal
chambers. At a later stage the margins of some of the plates
which end freely in the chambers give rise to teeth as shown in
FiG. 2.
It is difficult to secure direct evidence from the vegetative
thallus that the partitions form surface-outgrowths. Fic. 14,
however, which is drawn from a sec-
tion of the young female receptacle,
shows that such outgrowths are pos-
sible. The section was cut parallel
with the upper surface of the recep-
- tacle, and the figure shows two com-
plete chambers and parts of eight
others; two of the latter contain sec-
tions of the tubular epidermal pores
which hang down from the roofs of
the chambers. The partitions are one
cell thick but give the impression of
being thicker when cut obliquely.
The outgrowths originate as projec- Section parallel with
; ‘ the surface of a young female re-
tions of cells which become cut off by sindbis Yaak tebits the eplaeriaia
walls and then continue their growth x 270.
and cell-divisions. In the vegetative
thallus such outgrowths evidently play a very minor part in the
development of the green tissue.
The chambers below the dorsal layer make their appearance
very early, as seen in Fics. 12 and 13, although they always appear
later than the dorsal chambers. As the thallus becomes differen-
tiated, these chambers increase rapidly in size through the growth
of the bounding cells, but the appearance of new chambers, except
in the apical region, has not been demonstrated and seems im-
probable. If schizogenous processes play a part in the enlarge-
ment of these chambers, it is only to a very limited extent.
The green tissue in the thallus of Plagiochasma bears a strong
250 Evans: AIR CHAMBERS OF GRIMALDIA FRAGRANS
resemblance to that of Grimaldia. Its development has been
described by Miss Starr (10), her investigation having been based
on an undetermined species from Mexico. She confirms the
earlier observation of Barnes and Land that the air chambers of
Plagiochasma owe their origin to a splitting of cell walls below the
surface. She notes further that the chambers are at first deep
and narrow but that they soon become wide and irregular, and
she ascribes the changes in size and form which they show to a
“stretching and tearing of tissues between neighboring chambers.”
In other words she considers that schizogenous processes play a
leading part in the enlargement of the chambers as well as in their
origin. This conclusion is hardly supported by her f.. 11 or by
the earlier figures published by Barnes and Land (x, f. 17-22).
Although these figures indicate a schizogenous origin of the cham-
bers, they do not disprove that the enlargement is mainly due to
the growth of the surrounding cells.
SUMMARY
The air chambers of Grimaldia fragrans are in several layers in
the thickened median portion of the thallus.
The dorsal chambers communicate with the outside by means
of epidermal pores. They are subdivided by an irregular system
of more or less vertical, united cell plates, enclosing narrow spaces,
so that the boundaries of the chambers are difficult to distinguish.
The cell plates sometimes.reach the epidermis and sometimes do
not; in the latter case the free margins sometimes bear scattered
teeth, less than two cells in length, especially in the vicinity of the
pores. Except for these teeth the chambers lack filaments com-
pletely.
The more deeply situated chambers communicate with one
another and with the dorsal chambers by means of passageways;
they are scarcely or not at all subdivided by cell plates.
The chambers all owe their origin to a splitting of cell walls in
closely united tissue. In the case of the dorsal chambers the split
sometimes begins below the surface and extends outward; some-
times at the surface and extends inward.
The dorsal chambers appear first, very close to the apical cell,
but the more deeply situated chambers appear soon afterwards.
Evans: AIR CHAMBERS OF GRIMALDIA FRAGRANS 251
The increase in the size of the chambers is due largely to the
growth of the bounding cells and only slightly to further splittings
of cell walls. The system of united cell plates in the dorsal cham-
bers and the partitions between the chambers increase in vertical
height simultaneously. Direct outgrowths from the surfaces of
cell plates play a very small part in the process of subdivision.
The material upon which this investigation was based was col-
lected and prepared by Mr. John F. Logan, who expected to
utilize it in his own studies. Through the pressure of other work
his plans could not be realized, and his preparations were placed
at the disposal of the writer for examination. The writer would
therefore express his sincere thanks to Mr. Logan for his courtesy.
SHEFFIELD SCIENTIFIC SCHOOL,
YALE UNIVERSITY
LITERATURE CITED
_ Barnes, C. R., & Land, W. J. G. Bryological papers. I. The
origin of air chambers. Bot. Gaz. 44: 197-213. Ji 22. . 1907.
2. Black, C. A. The morphology of Riccia Frostii, Aust. Ann. Bot.
27: 511-532. pl. 37, 38. 1913.
Deutsch, H. A study of Targionia hypophylla. Bot. Gaz. 53:
492-503. f. I-13. 1912.
4. Hirsh, P. The development of the air chambers in the Ricciaceae.
Bull. Torrey Club 37: 73-77- f. 1-6. 1910.
Massalongo, C. Le Marchantiateas” della Flora Europea.
Atti R. Ist. Veneto 75: 669-816. pl. 1-27. I91
Miiller, K. Die Lebermoose Deutschlands, Oesterreichs u. d.
Schweiz. In L. Rabenhorst, Kryptogamen- -Flora, ed. 2, 6.
Leipzig. 1906-II.
O’Keeffe,L. Structure and ot of Targionia hypophylla.
New Phytol. 14: 105-116. f. I, 2 15.
Pietsch, W. Entwicklungsgeschichte eb vegetativen Thallus,
insbesondere der Luftkammern der Riccien. Flora 103: 347-
484. f. I-21. {Orr
g. Schiffner, V. Aaepholopteclie und biologische Untersuchungen
iiber die Gattungen Grimaldia und Neesiella. Hedwigia 47:
306-320. pl. 8. 1908.
Stur, A. M. A aoe Aytonia. Bot. Gaz. 61: 48-58. pl.
1-4 + f. 30-33-
. Stephani, F. ae an [In Species Hepaticarum 1: 89-
93.] Bull. Herb. Boissier 6: 792-796. 1898.
i
“
Py
ai
-
9
bn
_
INDEX TO AMERICAN BOTANICAL LITERATURE
1911-1918
The aim of this index is to include all current botanical literature written by
Americans, published in America, or based upon American material ; the word Amer.
ica being used in the broadest sense.
Reviews, and papers that relate epee to forestry, agriculture, horticulture,
manufactured products of vegetable oe r laboratory methods are not included, and
empt is made to index the literature of bacteriology. An occasional exception is
made in favor of some paper ‘shies in an American periodical which is devoted
wholly to botany. Reprints are not Sapaieatie unless they differ from the original in
oe ‘ a
to errors or omissions, their kindness will be appreciated.
This : ndex is reprinted monthly on cards, and furnished in this form to subscribers
at the rate of one cent for each card. Selections of cards are not permitted ; each
su Rider must nee all cards published during the term of his subscription, Corre.
spondence relating to the card issue should be addressed to the Treasurer of the nent
Botanical Club,
Andrews, E. F. A botanist’s suggestion for a national flower. Am.
Bot. 24: 8-10. F 1918.
Andrews, F. M. Closterium moniliferum. Proc. Indiana Acad. Sci.
TOIG! 42%, 324; . 1Ot7-
Anthony, R. D. Inheritance of sex in strawberries. New York Agr.
Exp. Sta. Tech. Bull. 63: 3-10. 5S 1917.
Atkinson, G. F. Selected cycles in Gymnoconia peckiana. Am. Jour.
Bot. 5: 79-83. 9 Mr 1918.
Beardslee, H. C. The russulas of North Carolina. Jour. Elisha
Mitchell Sci. Soc. 33: 147-198. pl. 7o-111. Ja 1918.
Blake, S. F. Further new or noteworthy Compositae. Contr. Gray
iar II. 53: 23-30. pl. r. 26 F 1918.
cludes Cirsium acanthodontum, Liabum pcg en Verbesina Arthurii,
Sseeolt denticulatum and A phanostephus Kidderi, spp. n
Blake, S. F. Lycopodium sabinaefolium Willd. var. sharonense (Blake)
comb. nov. Rhodora 20: 60. 2 Mr 1918.
Blake, S. F. New plants from Oaxaca. Contr. Gray Herb. II. 53:
55-65. 26 F 1918.
Specics it e=4
254 INDEX TO AMERICAN BOTANICAL LITERATURE
Blake, S.F. New Spermatophytes collected in Venezuela and Curacao
by Messrs. Curran and Haman. Contr. Gray Herb. II. 53: 30-55.
26F 1
Includes oe new genera Hecatostemon and Oxycarpha, and 24 new species in
various genera
biieen: G. S. New species of Russula from Massachusetts.
Mycologia 10: 93-96. Mr 1918.
Four new species are described.
Britton, N. L. An undescribed Scirpus from California. Torreya 18:
56. f. fo 9 Mr 1018.
Scirpus Congdoni sp. nov.
Buchanan, R. E. The evolution of the bacteria. Science II. 47: 320-
324. 29 Mr 1918.
A review of Dr. I. J. Kligler’s doce on the evolution and relationship of bacteria
containing additional informatio
Burkholder, W. H. The anthracnose disease of the raspberry and
related plants. Cornell Agr. Exp. Sta. Bull. 395: 155-183. f. 12-21.
N 1917.
Bush, B. F. The genus Euthamia in Missouri. Am. Mid. Nat. S:
157-177. Mr 1918.
Carpenter, C. W. Wilt diseases of okra and the Verticillium-wilt
problem. Jour. Agr. Research 12: 529-546. pl. A, 17-27. 4 Mr
1918.
Clute, W. N. Note and comment. Am. Bot. 24: 26-33. F 1918.
Includes notes on: Origin of coal, Sir Walter Raleigh’s potatoes, The bluet in
cultivation, The first frost, Erigenia bulbosa, Lycopsida and Pteropsida, Evolution
he larch, Thymol.
Conard, H. S. Prairie plants for the garden. Jour. Internat. Gard.
Club 2: 129-140. Mr 1918. [Illust.] '
Cooper, J. R. Methods of controlling blister canker. Nebraska Agr.
Exp. Sta. Bull. 161: 1-18. pl. 1-7. 15 D 1917.
Coulter, J. M. A century of botany in Indiana. Proc. Indiana Acad.
Sci. 1916: 236-260. 1917.
Crozier, W. J. Cell penetration by acids—IV. Note on the penetra-
tion of phosphoric acid. Jour. Biol. Chem. 33: 463-470. f. 1-3.
Mr 1918.
Deam, C. C. Plants new or rare to Indiana—VII. Proc. Indiana
_ Acad. Sci. 1916: 315-322. 1917.
INDEX TO AMERICAN BOTANICAL LITERATURE 255
Downer, H. E. Forcing native plants. Jour. Internat. Gard. Club.
2: 119-123. Mr 1918. [Illust.]
Dunbar, J. Notes on Cotoneasters. Jour. Internat. Gard. Club 2:
83-85. Mr 1918. [Illust.]
Fairchild, D. Gardens for plant breeders. Jour. Heredity 9: 112-116.
f. 5-8 Mr 1918.
Faulwetter, R. C. The Alternaria leaf-spot of cotton. Phytopa-
thology 8: 98-105. f. 7-3. Mr 1918.
Fink, B. The distribution of fungi in Porto Rico. Mycologia 10: 58—
61. Mr 1918.
Fitzpatrick, H. M. Sexuality in Rhizina undulata Fries. Bot. Gaz.
65: 201-226. pl. 3, 4. 15 Mr 1918.
Frye, T. C. The rhacomitriums of western North America. Bryolo-
gist 20: 91-98. pl. 20-23. 5 D 1917; 21: 1-16. pl. I-34... 35 Me
1918.
Fuller, G. D., & Bakke, A. L. Raunkiaer’s “life forms,” “leaf-size
classes,”” and statistical methods. Plant World 21: 25-37. f. r.
F 1918.
Galloway, B. T. Some of the broader phytopathological problems in
their relation to foreign seed and plant introduction. Phyto-
pathology 8: 87-97. Mr 1918.
Gruber, C.L. Herbs with fleshy fruits. Am. Bot. 24: 13,14. F 1918.
Hanson, H. C. The invasion of a Missouri river alluvial flood plain.
Am. Mid. Nat. 5: 196-201. pl. 1, 2. Mr 1918.
Harper, E. T. The Clavaria fistulosa group. Mycologia 10: 53-57.
pl. 3-5. Mr 1918.
Harper, E.T. Two remarkable Discomycetes. Bull. Torrey Club 45:
77-86. pl. 1-3. 7 Mr 1918.
Underwoodia columnaris and Pustularia gigantea.
Harper, R. M. Some dynamic studies of Long Island vegetation.
Plant World 21: 38-46. f. 1, 2. F 1918.
Harris, J. A., & Avery, B. T. Correlation of morphological variations
in the seedling of Phaseolus vulgaris. Bull. Torrey Club 45: 109-
119. 1 Ap 1918.
Hendrickson, A. H. The common honey bee as an agent in prune
pollination. Calif. Agr. Exp. Sta. Bull. 291: 215-236. f. 1-13.
Ja 1918.
256 INDEX TO AMERICAN BOTANICAL LITERATURE
Herrington, A. Lilies. Jour. Internat. Gard. Club 2:°5-29. Mr 1918.
[Illust.]
‘Hoffer, G. N. An aecium on red clover, Trifolium pratense L. Proc.
Indiana Acad. Sci. 1916: 325, 326. 1917.
Hoffer, G. N., & Holbert, J. R. Results of corn disease investigations.
Science II. 47: 246, 247. 8 Mr 1918.
Holmes, E. M. Medicinal herbs: their cultivation and preparation in
Great Britain. Jour. Internat. Gard. Club 2: 35-82. Mr 10918.
[Illust.]
Jackson, H. S. Apple diseases in Indiana, with spray schedule.
Indiana Agr. Exp. Sta. Circ. 7: 1-23. f. 1-14. $1917.
Jackson, T. F. The description and stratigraphic relationships of
fossil plants from the lower Pennsylvania rocks of Indiana. Proc.
Indiana Acad. Sci. 1916: 405-428. pl. 7-10 + f..r. 1917.
ourteen new species are described in Lepidodendron (1), Trigonocarpum (3),
and Cardiocar pon (10).
Jackson, H. S., & Osner, G. A. Potato diseases in Indiana. Indiana
Agr. Exp. Sta. Cire. 71: 1-16. f. 7-5.. S 1617,
Jones, L. R. Disease resistance in cabbage. Proc. Nat. Acad. Sci. 4:
4240.7. 9,92 15 F 918;
Kendall, J. N. Abscission of flowers and fruits in the Solonaceae, with
special reference to Nicotiana. Univ. Calif. Publ. Bot. 5: 347-428.
pl. 490-53 +f. I-10. 6 Mr 1918.
Knowlton, C. H., & Deane, W. Reports on the flora of the Boston
District,—XXVI. Rhodora 20: 15-18. 25 Ja 1918;—XXVII.
Rhodora 20: 55-59. 2 Mr 1918. :
Long, B. History of the American record of Scirpus mucronatus.
Rhodora 20: 41-48. 2 Mr 1918.
Ludwig, C. A., & Rees, C. C. The structure of the uredinium in
Pucciniastrum Agrimoniae. Am. Jour. Bot. 5: 55-60. pl. 8. 9
Mr 1918.
Lunell, J. The collecting, drying and mounting of plant specimens.
Am. Mid. Nat. 5: 191-195. Mr 1918.
Lyman, G. R. The need of organization of American botanists for
more effective prosecution of war work. Science II. 37: 279-285.
22 Mr 1918.
INDEX TO AMERICAN BOTANICAL LITERATURE 257
Macbride, J. F. New or otherwise interesting plants, mostly North
American Liliaceae and Chenopodiaceae. Contr. Gray Herb. II.
53: I-22. 26 F 1918.
Includes Cirsium praeteriens, Lycium Spencerae, Lomatium Nelsonianum, Lotus
Spencerae, dibs Paes and Tricyrtis clinata, spp. nov. and several new varieties
and new combinatio
MacCaughey, V. The native bananas of the Hawaiian Islands.
Plant World 21: 1-12. Ja 1918.
McGee, J. M. The imbibitional swelling of marine algae. Plant
World 21: 13-15. Ja 1918.
McKay, M. B., & Pool, V. W. Field studies of Cercospora beticola.
Pisoni halopy 8° 419-136. (2-3, 2. Mr igis.
McNair, J. B. Secretory canals of Rhus diversiloba. Bot. Gaz. 65:
268-273. 15 Mr 1918.
Middlebrook, C. A. Odorous flowers of Texas. Am. Bot. 24: 3-6.
F 1918.
Moore,G.T. Algological notes.—II. Preliminary list of algae in Devils
Lake, North Dakota. Ann. Missouri Bot. Gard. 4: 293-303.
N 1917
Moore, W., & Williams, J. J. Studies in greenhouse fumigation with
hydrocyanic acid: physiological effects on the plant. Jour. Agr.
Research 11: 319-338. pl. 34 +f. I-11. 12N 1917.
Mottier, D. M. Chondriosomes and the primordia of chloroplasts and
leucoplasts. Ann. Bot. 32: 91-114. pl. 7. Ja 1918.
Munn, M. T. Neck-rot disease of onions. New York Agr. Exp. Sta.
Bull. 437: 365-455. pl. r-1z. Jl 1917.
Murphy, P. A. The morphology and cytology of the sexual organs of
Phytophthora erythroseptica Pethyb. Ann. Bot. 32: 115-153. fl. 2,
3. Jasgi18.
Murrill, W. A. The Agaricaceae of tropical North America.—VII.
Mycologia 10: 15-33. 14 F 1918
Includes 18 new species in Atylospora (11), Psathyrella (5), Psilocybe (1), and
Campanularius (1).
Murrill, W. A. The Agaricaceae of tropical North America—VIII.
bork 10: ih Mr 1918.
Include cies in Drosophila (8), Hypholoma (1), Gomphidius (1),
Stropharia ais asia eek and Coprinus (4).
Murrill, W. A. Murrill’s and Saccardo’s names of polypores thahpered:
1-13. New York. 1918.
Dos +. INDEX TO AMERICAN BOTANICAL LITERATURE
Nelson, J. C. Additions to the flora of western Oregon. Torreya 18:
21-35. 8 Mr 1918.
Osner, G. A. Additions to the list of plant diseases of economic im-
portance in Indiana. Proc. Indiana Acad. Sci. 1916: 327-332.
1917.
Pammel, L. H., & Dox, A.W. The protein contents and microchemical
tests of the seeds of some common Iowa weeds. Proc. Iowa Acad.
Sci. 24: 527-532. 1917.
Pipal, F. J. Weed seeds in the soil. Proc. Indiana Acad. Sci. 1916:
368-377. 1917.
Popenoe, W. Avocados as food in Guatemala. Jour. Heredity 9:
99-107. f. I-4. Mr 1918.
Porter, A. A., & Coons, G. W. Differences between the species of
Tilletia on wheat. Phytopathology 8: 106-113. f. 1-4. Mr 1918.
Rathbun, R. The Columbian Institute for the promotion of arts and
sciences. U.S. Nat. Mus. Bull. ror: 1-85. 18 O 1917.
h +
chapters Botanical Gardens; Washington Botanical Society, etc.
Rice, T. B. A study of the relations between plant growth and com-
bined nitrogen in Winona Lake. Proc. Indiana Acad. Sci. 1916:
333-362. f. I-16. 1917.
- Rock, J. F. Pelea and Platydesma. Bot. Gaz. 65: 261-267. f. I.
15 Mr 1918.
Includes Pelea Gayana and P. recurvata, spp. nov.
Rosendahl, C. O. Observations on Betula in Minnesota with special
reference to some natural hybrids. Minnesota Bot. Stud. 4: 443-
459. dl. 40 + J. 1, 2. 20°35 1616.
Rosendahl, C. O., Observations on the morphology of the underground
stems of Symplocarpus and Lysichiton, together with some notes on
geographical distribution and relationship. Minnesota Bot. Stud.
4: 137-152. BW. 24,22 © 1-17, 2, 18 5 1081.
Rosendahl, C. O., & Butters, F. K Reputed Minnesota plants which
probably do not occur in the state. Minnesota Bot. Studies 4: 461—
473: 2058 1916.
Sax, H. J. Spore formation in Philocopra coeruleotecta. Am. Jour.
Bot. 5: 61-78. pl. 9-11. 9 Mr 1918.
Schaffner, J. H. The expression of sexual dimorphism in heterosporous
sporophytes. Ohio Jour. Sci. 18: 101-125. f. 7-28. F 1918.
BuLL. ToRREY CLUB VOLUME 45, PLATE 7
NISHIMURA: CARRIER OF THE MOSAIC DISEASE
Vol. 45 No. 7
BULLETIN
OF THE
TORREY BOTANICAL CLUB
JULY, 1918
The strand flora of the Hawaiian Archipelago—l. Geographical re-
lations, origin, and composition
VAUGHAN MACCAUGHEY
‘GEOGRAPHICAL RELATIONS
For a long time botanists have manifested particular interest
in the strand vegetation of various countries. The floras of many
continental and insular strands have alike yielded significant
material. There is, however, absolutely no comprehensive ac-
count of the Hawaiian littoral. Fragmentary and _ uncodrdi-
nated notes concerning various Hawaiian strand plants are given
by Hillebrand, Wawra, Gray, Chamisso, Heller, Mann, Schauins-
land, Forbes, and other botanists, who at various times during
the past century have studied the Hawaiian flora. In the works
of none of these investigators is the littoral flora given any special
prominence or consideration. This is somewhat surprising, as
much of the Hawaiian coast line is readily accessible by boat or
trail, whereas the montane districts present innumerable diffi-
culties to the explorer and collector.
Guppy (’06),* in connection with his suggestive studies of plant
dispersal in the Pacific, visited the Hawaiian Archipelago and
studied the strand flora with reference to the general problems of
evolution and distribution. Frequent references are made to
Guppy’s work in the present paper. His brilliant theories will
undoubtedly require more or less revision as data concerning the
Pacific flora become more comprehensive and standardized.
* The literature cited will be listed in the second paper of this series.
[The BULLETIN for June (45: 219-258. pl. 7) was issued June 20, 1918.]
259
260 MacCAuGHEY: THE STRAND FLORA
The unique position of the Hawaiian Islands, as the most
northern group of the great Polynesian island series, and as a
region of extreme isolation, gives particular significance to
its strand flora. The absences from this flora are as important
criteria as are the species actually present, *and in many ways
contribute as effectively to our knowledge of the origin and history
of Hawaii’s strand flora.
The attempt is here made to present a salient account of the
Hawaiian littoral flora, both from the standpoint of content and
dispersal, and also from the ecological viewpoint. In the latter
phases of the subject the author has been particularly interested.
During a residence of nine years in the islands he has made hun-
dreds of excursions along Hawaiian strands, including the prin-
cipal islands of the archipelago. The present papers incorporate
the important data of these field studies.
A noteworthy feature of the littoral floras of the tropical
Pacific islands is their remarkable similarity. As Hedley (’15)
expresses it: ‘The same species are repeated from atoll to
atoll over enormous distances across the Pacific Ocean. The
identity of the vegetation possessed by tiny islets separated by
thousands of miles of deepest ocean is very striking, since para-
doxically they present a greater continuity of life range than any
continent can show.’’ Many of the more common Hawaiian lit-
toral plants occur on practically all the islands of the archipelago,
along an axis of nearly two thousand miles, whereas the montane
species are highly localized.
Just as the interior mountainous districts of a high Pacific
island contain the majority of the endemic species, so the strand
regions are characterized by a majority of the cosmopolitan or
wide-ranging species. Tansley and Fritsch (’05) find two main
causes for ‘‘the striking uniformity of strand plants through the
tropics—first, the great similarity of life conditions prevailing on
tropical coasts, and secondly, tropical strand plants are mostly
adapted for distribution by ocean currents.’’ These factors will
be considered in detail in later sections of this paper.
As will be shown later, the Hawaiian littoral flora comprises
many species that occur in other parts of the Pacific, and in many
other parts of the world. The mountain flora, on the contrary,
OF THE HAWAIIAN ARCHIPELAGO 261
is highly endemic and precinctive, and each island possesses an
array of peculiar forms. The Hawaiian Islands therefore obey
the general law of the cosmopolitanism of littoral constituents.
The geographic situation of the Hawaiian Islands is unique,
and has a very important bearing upon the character of the strand
flora. There is no other land area of equal magnitude on the
earth that is so far removed from continental land areas. The
Hawaiian Islands are the most isolated islands, of their size, in
the world, and their flora strikingly and faithfully registers this
profound and prolonged isolation. An examination of a map
of the North Pacific Ocean will show the nature of this isolation.
Using the island of Oahu, which is situated in the eastern part of
the archipelago, as a base point, the distances to adjacent land-
masses, in terms of nautical miles, to ports specified, are as follows:
San Francis a 2,100 Pe oer i ee ee ee 2,700
Sl) DIEGO. Gs Sc. es ee 2,260 PC RIA CS ee 3,810
Patianie oy osha ee. 4,665 ide 2 ale ay Oa Re Re 4,980
COAG A ian sis Ga aes eee ee 5,147 natn er a eee dee. 4,920
Cane Hom Salvi sd ee: 6,488 Cama er 3-400
Vahiti ic ever ees Let 2,440 ‘cnet Poca v erin et 4,721
Ramoee Ss wes aero ey 2,290 OTRO ce esa ie icine. 2,360
ARIAS os ee sn a ics 2,106
The isolation is further emphasized by an examination of the
deep-sea soundings in the vicinity of the Hawaiian Islands, which
show that the islands rise from abyssal depths. The great deeps
of the Pacific Ocean, which lie between the Hawaiian Archipelago
and the continents, are:
Name of deep | — bine we tathoms. depth in A in sq +
May oe oe | Due north 3,540 1,033,000
DBUpAN oo ess eee | Due north
Sry Se as | Due north |
pn nn Mee, Fae er are Northea
FUERA, Gt ks Northeast
Tanner oe es an Northeast
GHOVErS io Cee! Northeast
Agassi ee). Northeast
Bache... es cer | Northeast
MOG. Fa | Southeast }
OCHAY i. ol eee Southeast
Belknap. . 06.2 ies | South | 3337 165,000
Canipbell 23.3 Gsaa: | South
ANNER pic ek Southwest |
BYOOKE 5 oN oc | West 3,429 282,000
Batley ie | ' West | 3432 241,000
Tuscarora 226s is 5: | Northwest 4,655 :
262 MacCauGHEY: THE STRAND FLORA
The isolation of the Hawaiian flora is reflected in the vegetation
of the littoral zone by the high proportion of endemic species,
32 out of a total of 110, or 30 per cent. Nine of these belong to
endemic genera. This is a remarkable showing, since littoral
floras, in all parts of the world, are usually comprised almost wholly
of cosmopolitan and non-endemic constituents.
SouTH PACIFIC EQUIVALENTS OF HAWAII
In the comparisons which are likely to be made of the Hawaiian
flora with those of the South Pacific islands it is important to
recognize the geologic and topographic status of the various
island groups. Guppy (’06, p. 14) makes the eee significant
observationt:
The Hawaiian Islands, standing alone in the North Pacific, form a floral region
in themselves, a region that is the equivalent not of one group in the South Pacific,
such as that of Fiji or of Tahiti, but of the whole area comprising all the groups ex-
tending from Fiji to the Paumotu Archipelago.
For the purposes of this paper the islands and continental
coasts of the Pacific region may be roughly divided into the fol-
lowing great phyto-geographic provinces, listed clockwise in and
around this greatest of marine basins. These provinces are not
presented as of equal biological value or range, but merely for
purposes of convenience in description.
f ct woes SIBERIA AND ALASKA: Kamchatka, raat Tschuktsche,
Al Islands, Alaska, British Columbia, and coastal islan
‘ ie aie oF NortH AMErica: Washington, Oregon, see and coastal
islands.
3- Paciric Coast oF MEXICO AND CENTRAL AMERICA: Lower California, Manion:
Guatamala, Honduras, Nicaragua, Costa Rica, Panama.
4. Paciric COAST OF NORTHERN SOUTH sega Colombia, Ecuador, Galapagos
Islands, Peru, Chile, Juan Fernandez Islan
vd
5. PAciric COAST OF SOUTHERN SOUTH AMERICA: eee and coastal islands; Tierra
del Fuego.
6. oe ipampey Tasmania, New Zealand, and adjacent islands.
7. MALAYSIA: matra, caine epone: and other Sunda Islands; Moluccas, New
uinea, Pi ine Isla
8. SOUTHEASTERN ASIA: leis Hai-nan, Indo-China, Cochin China, Cambodja,
Siam, vei Peninsula.
. PACIFIC Cons ST OF CENTRAL AsIA: Japan, Riu-Kiu Islands, Formosa, and adja-
‘0
9
x
&
Ge
n
:
7
marck Archi ee) Louisiade, Solomon, Santa Cruz, New
and Loyalty Islands, New Caledonia, Fiji, and intervening smaller
OF THE HAWAIIAN ARCHIPELAGO © 263
II. MIKRONESIA: Mariana, Pelew, Caroline, Marshall, and Gilbert Islands, and
intervening islands.
12. Potynesta—(a) Nuclear Polynesia: Samoa, Tonga, Fiji, Tokelau, Ellice Islands,
etc.
(b) Central Polynesia: Cook, Phoenix, Tubuai, Rokahanga, Ton-
garewa, Manahiki Islands.
(c) Southeast abaireae Society, Marquesas, Taumotu, Gambier,
Pitcairn, Easter, Ducie Islands, etc.
(d) Northern Feito: HAWAIIAN ISLANDS
(e) Southern Polynesia: New Zealand and Cthiahatn Islands.
f) Polynesian Verge: scattered islands between Polynesia and
Melanesia, such as Ticopia.
EXTENT OF THE HAWAIIAN LITTORAL
The great length of the archipelago gives the littoral zone a
much larger significance and extent than if the archipelago con-
sisted of but a few islands situated close together. The Hawaiian
littoral, ranging for nearly two thousand miles, contrasts sharply
with the compact littoral of such groups as Samoa, Tonga, New
Caledonia, Ellice and Phoenix. Other Pacific island groups which
are extended over long axes, similar to Hawaii, are the Aleutian,
Kurile, Paumotu, Marshall, Caroline, and Solomon Islands.
The Hawaiian strand occupies an island series extending from
18° 54’ to 22° 15’ north latitude, and between 154° 50’ and 160° 30’
of longitude west of Greenwich. This range should greatly in-
crease the mathematical probability of plant dispersal, and in
some measure tend to neutralize the powerful isolation-factor.
The east-and-west range of the littoral naturally results in a much
greater homogeneity of flora than would be the case in an archi-
pelago with a dominant north-and-south axis. In this respect
the Hawaiian Islands may be contrasted with such archipelagoes
as the Philippines, and the Mariana and Maldive groups.
The great variation in the size and elevation of the several
islands markedly influences the extent of the littoral. In general,
the low islands have strands that extend further back into the
interior than do those of the high islands; the small islands have a
larger proportion of strand, relative to their total area, than do
the large islands. The small, low coral islets that predominate
in the western end of the archipelago are littoral throughout
practically all their area; the large, high islands of Maui and Hawaii,
at the eastern extremity of the archipelago, have a narrow and
closely defined strand.
264 MacCaAuGHEY: THE STRAND FLORA
The following islands have relatively wide strands:
I. Ocean, paps or Cure Island: circular barrier atoll, 16 miles in circumference;
lagoon about 38 square miles; several low sand islets in the lagoon;
aids eneeneety and other low vegetation on the largest sand islet,
**Green Island
a) gerne or Brook's Island: circular barrier atoll, 18 miles in circumference; area
agoon about 40 square miles; several low sand islets in the lagoon; native
os introduced vegetation.
3. Pearl and Hermes Reef: irregular, oval barrier atoll, 42 miles in circumference;
area of lagoon about 80 square miles; numerous low sand islets in the lagoon,
with grasses and other low i
4. ieee Island: low, oval isle of coral sand two > miles by three miles; lagoon
empty of water; surrounded by reef whic seven miles from isle;
Vv
5. Laysan Island: elevat latoll; t iles b il da half; well-developed
fringing reef; briny lagoon; ébinrdent acct vegetation.
eS revch F ete s Shoal: crescentic atoll, with numerous low sand isles, and several
ocky volcanic isles; area of shoe! about 30 square miles; extensive reefs;
w vegetation on san es
7. Small isles ee the coasts of Oahu, most of them formerly connected with the
island; sparse vegetation: Kihewamoku, Mokuaia; Pulemoku; Kukuihoolua;
Mokualai; Mokolii; Kapapa; Ahuolaka; Kekepa; Mokuoloe; Mokolea
Cea Kaonikaipu; Mokauea; cs. Onini; Moku Umaume; Lau-
lau
8. Smali nei along the coasts of Molokai, most of them formerly connected with
the island; vegetation sparse or ete Namoku; Mokapu; Okala; Kuelo-
Lepau; Mokuhooniki; Kanaha; Pu
g. Small isles along the coasts of Hawaii: Casi Island, etc.
Narrow strands are characteristic of the following islands:
10. Nithan: area 97 square miles; highest point, 1,300 feet.
13- Molokai: area 261 square miles; highest point, 4,958 feet.
14. Maui: area 728 square miles; highest point, 10,032 feet.
15. Lanai: area 139 square miles; highest point, 3,400 feet.
16. Kahoolawe: area 69 square miles; highest point, 1,472 feet.
17. Hawaii: area 4015 square miles; highest point, 13,825 feet.
A number of the smaller islands of the Hawaiian Archipelago
rise very abruptly from the sea, and are characterized by steep or
precipitous coasts. The following are of this type:
18. Gardiner Island: conical rock, 700 fee t in diameter, 175 feet high; cliffs 60-70
eet high on all sides; a smaller, precipitous rock nearby.
1g. French Frigates Shoal: volcanic rocky islet in center of formerl agoon; 180 feet
long, 45 feet wide, 120 feet high; vertical walls; barr
20. Necker Island: remnant of volcanic crater, eeucd by reef; isle is three
q ts of a mile long, 500 feet wide, and 300 feet high; scanty vegetation
sea-cliffs on all sides.
OF THE HAWAIIAN ARCHIPELAGO 265
21. Nihoa, Bird Island or Moku (Modu) Manu: volcanic remnant; three quarters of a
mile long, one third of a mile wide, 600-900 feet high; sea-cliffs on all sides;
vegetation scanty.
22. Small isles along the coast of Niihau, probably at one time connected with the
island: Kaula and Lehua, small eroded cinder cones with sea-cliffs and steep
es
Small isl th t of Oahu: Moku M : volcanic remnants); Manana
(Rabbit fant: eroded crater, with sea-cliff:
24. Small isles and rocks along the coasts of Shaaonilt: Mokolea, Mokohola, Moko-
25. . pes along the coasts of Maui: Molokini,
26. Five Needles: a group of detached pinnacle a dbeat 120 feet high, situated
five and a half miles north of Cape Kaea, Lanai, and about the middle of the
bight on the west side of the island.
The largest strand areas on any single island occur on the
island of Hawaii; the smallest strands are those of the tiny islets
in the westward end of the archipelago. On the whole, the
Hawaiian strand, as a phytogeographic province, is poorly de-
veloped when compared with the Indo-Malayan or West Indian
strands, or with those of numerous other archipelagoes.
SUBSIDENCE AND ELEVATION
A factor of far-reaching importance in any biological studies in
the Hawaiian Archipelago is that of subsidence, i. e., the islands
are but the apices of lofty and slowly-sinking submarine moun-
tains. Physiographical evidence is accumulating to show that
during previous stages in the history of the central Pacific, these
islands undoubtedly stood thousands of feet higher than they do at
present. Many stages of subsidence and erosion may be found
today within the group, ranging from the large, actively volcanic
island of Hawaii (nearly 14,000 feet in elevation), at one end of
the chain, to the tiny coral atolls, but a few feet above sea-level,
which are scattered along the other extremity.
_ Considering the strand zone of any given island, it is evident
that through a long period of time this zone has been slowly
creeping up the slopes of the island, and the terrestrial vegetation
has been crowded into steadily diminishing areas. In other words,
the total mileage of strand was formerly much greater than at
present, other things being equal. Granting slow subsidence as
the prevalent condition of Pacific islands (see, in this connection, an
important contribution by Bryan, '16), the great strand mileage
266 MacCauGHEeEy: THE STRAND FLORA
of an island like Hawaii steadily decreases until ultimately the
condition exhibited by the tiny strand of Laysan or Midway is
reached. In this manner the subsidence-factor, although not of
especial force at any one time, has been through long periods of
time a powerful influence on the strand flora.
Although subsidence has been the dominant note in Hawaii’s
geological history, there have been minor elevations within recent
times. Raised coral reefs and beaches occur at various points
along the coasts, e. g., along the southern and western shores of
Oahu. The highest reef known is on the southwestern end of
Mailiilii, elevated 120 feet above the sea. Just as local elevation
at such points as Mokapu, Kalihi, and the Coral Plain has pushed
the littoral zone seaward, so at Hanalei, Kahana, Kaneohe, and
Pearl Harbor the drowning or submergence of valley-mouths has
developed deep embayments, and the littoral flora extends deep
into the lowland zone which surrounds it on three sides. It is
evident that sufficient study and emphasis has not been given to
the ecological effects of these gradual changes of land- and sea-
level.
In his illuminating studies of the New Zealand flora, Aston
(’12) concludes that the raised marine beaches at Cape Turakirae
show that there has occurred comparatively recently, and perhaps
within historic times, rapid elevation of the coast line. Violent
earthquakes have so altered the physiography of the littoral as to
result in some unusual ecological features.
ORIGIN '
RELATION OF PACIFIC OCEAN CURRENTS TO THE DISSEMINATION
OF PLANTS OF THE HAWAIIAN LITTORAL
The importance of ocean currents as agencies for the distri-
bution of plants, and more particularly for the wide dissemination
of beach species, has long been recognized. In a vast body of
water, like the Pacific, dotted with thousands of scattered islands,
the surface currents assume special significance in relation to the
migrations of plants, animals, and man.
The surface circulation of the Pacific is, on the whole, notably
less active than that of the Atlantic. The vertices of the rota-
tional movements are marked by “Sargasso Seas”’ in the north and
OF THE HAWAIIAN ARCHIPELAGO 267
south basins, but these quiet areas are of small extent when com-
pared with the well-known Sargasso Sea of the North Atlantic
Ocean.
Reference should here be made to the Northern Equatorial
Current, which receives important contributions from the great
stream that sweeps down the North Pacific coast of America.
Many of the largest and most famous of the Hawaiian double
canoes were hewn from Douglas spruce (Pseudotsuga taxifolia)
which had been carried to the shores of Niihau, Kauai, and other
islands by the currents. It is a well-known fact that the natives
of the Alaska islands obtain much of their fire-wood as drift
from the Asiatic coast. Japanese fishing-boats, at various times
in history, have drifted to the Hawaiian Islands and to the north-
west coast of America.
Between the two great equatorial currents flowing westward
on either side of the equator there is a narrow counter-equatorial
current flowing to the east. This stream is largely assisted during
the latter half of the year by the southwest monsoon, and from
July to October the southwest winds prevailing east of 150° E.
further strengthen the current, but later in the year the easterly
winds weaken or even destroy it. The currents of the South
Pacific are well shown in Schimper’s (’91) monograph of the Indo-
Malayan strand flora.
_ A feature of ocean currents as seed carriers that has not been
sufficiently emphasized is the definiteness of their courses. This —
fact is well illustrated by the large number of tree trunks and logs
from the North Pacific coast that are annually cast upon the
Hawaiian coasts. These trees occur in a relatively small and well-
defined region, and evidently follow a definite course across the
North Pacific. _Wood-Jones (’12) performed an interesting ex-
periment to determine the course of drift material in the Indian
Ocean. He cast adrift, in the Cocos-Keeling Islands, bottles
containing messages. One was picked up on the beach of Brava,
Italian Somaliland, after a journey of three thousand miles across
the Indian Ocean, and a second one, sent out nearly a year later,
was washed ashore at precisely the same place. This definiteness
of course gives to the ocean currents a high potential cumulative
effect as carriers, that merits more than passing mention.
268 MacCauGHEy: THE STRAND FLORA
CURRENTS IN THE VICINITY OF HAWAII
The United States Coast Pilot contains data concerning the
local island currents, which may be summarized as follows. The
strong northeast trades begin early in March, blowing well from
the northward until May, and from then until October they are
more easterly. During October the trades are light with frequent
calms, and occasionally a west southwest swell sets in. During
November and December the trades are strong and variable,
occasionally being interruped by light southerly winds. During
anuary and February southerly and southwesterly gales often
prevail. These are konas and are from a few hours to two or three
days duration, followed by rain.
Hawatl.—Generally, the currents follow the trades but occa-
sionally they set against the wind. A current follows the coast
north of Cape Kumukahi around Upolu Point; another one fol-
lows the trend of the coast offshore southwestward from Cape
Kumukahi around Kalae and northward as far as Upolu Point.
There is an inshore current that sets southward from Okoe Landing
along the west coast around Kalae, and thence northeastward
along the shore as far as Keauhou.
Maul.—Generally the currents set with the trades... A current
follows the north shore of Maui westward from Kauiki Head and
draws down through Pailolo Channel; the current is stronger on
the Molokai side of the channel. A strong current follows the
coast southward of Kauiki Head until past Kahoolawe. In the
vicinity of Lahaina the current generally sets northwestward.
Mo.oxkal.—The current sets westward along the entire north-
erly coast, and about half the length of ‘the southerly shore, where
an easterly current prevails.
OaHu.—The currents around Oahu are variable in strength
and direction, but the general movement of the water along the
coast is westward or northwestward, the direction being modified
by the trend of the coast.
Kaval.—Currents are very uncertain as to direction but they
generally follow the winds, though frequently setting in the oppo-
site direction during the first calms after strong trades.
A careful study of any good map which depicts the ocean cur-
OF THE HAWAIIAN ARCHIPELAGO 269
rents of the North Pacific Ocean will graphically show that the
Hawaiian Archipelago is practically outside the zone of influence
of the great currents that would naturally bring the seeds of
tropical plants to her shores. Guppy’s statements (’06, pp. 75, 64)
are pertinent in this connection:
The currents of the Pacific have failed to establish the numerous beach-trees
(possessing buoyant fruits) of the Pacific Islands, not only in the Hawaiian group,
but also on the coast of America; and it is therefore argued that we should expect the
Hawaiian group to have received through the currents its shore-plants with buoyant
seeds or fruits Beal the tropical west coasts of America.
In support of this contention it is pointed out that most of the Hawaiian strand-
plants that are dispersed by the currents are found in America, and some indeed in
America to the exclusion of the Old World.
The arrangement of the currents in the North Pacific also favours the view that
the Hawaiian Islands are more likely to receive plants by the agency of the currents
from America than from the Asiatic side of the Pacific.
ee
Speaking generally of the extension eastward of the Indo-Malayan strand-plants
over the Pacific, Professor Schimper (['91] page 195) remarks that they become fewer
and fewer in ie eons need -ghueig: from their original home, their number
shrinking toa f the Marquesas and the Hawaiian
Islands. The canes eo introduced through the currents into Hawaiiin
all likelihood, therefore, d
IMPORTANCE OF DRIFT MATERIAL
Drift material is much more abundant along the Hawaiian
windward shores than on the leeward shores. Nowhere does it
attain the proportions that characterize many other regions else-
where on the globe. Certain districts, for example, the south-
east coast of Hawaii, between Honuapo and Kalae, particularly
the Kamilo beach near Kaluwalu, seem to be much more favor-
ably situated for the reception and accumulation of drift than do
others.
Tansley and Fritsch ('05) describe the abundant drift on
portions of the Ceylon littoral, and note the great variety of plant
fragments, fruits, and seeds:
The thickest masses of drift were very moist and quite warm to the hand, and
intervals ost striking sight. Of these the most conspicuous were Cerbera
odollam, Saaiilins inophyllum, Bruguiera gymnorhiza, Crinum asiaticum, and
Colocasia antiquorum (from bits of old rhizome).
Moseley (’79, p. 367) reports from the Moluccas living epiphytic
270 MaAcCAUGHEY: THE STRAND FLORA
orchids and young palms as part of the drift, washed high up on
the beach and growing. He states:
We passed large quantities of leaves, fruits, and flowers, and branches of trees
floated off from the shores. . . .- I was astonished at the large quantities of fresh vege-
table matter thus seen floating on the sea. . . . Not only are large quantities of
fruits [containing seeds] capable of RIE FE thus transported from island to
island, but entire ier “ea even trees, are washed from island to island and
transplanted by the w
Hooker (’47, p. met states that the majority of the littoral species
of the Galapagos Islands have reached the islands through oceanic
and aerial currents. There are about twenty such plants, mostly
species common to warm latitudes. Some of these are: Cissam-
pelos Partera, Tribulus cistoides, Tephrosia littoralis, Verbena
littoralis, Avicennia tomentosa, Scaevola Plumieri, Ipomoea mari-
tima, Calystegia Soldanella, and Heliotropium curassavicum.
Hooker attributes the following plants of the Peruvian and Chilean
littoral, which occur on the Galapagos strand, to ocean currents:
Vigna oahuensis, Acacia Cavenia, Nicotiana glutinosa, Dictyocalyx
Miersit, Lycopersicum peruvianum, Verbena littoralis, V. polystacha,
and Plantago tomentosa.
FLOTATION ADAPTATIONS OF STRAND PLANT SEEDS
In his ‘monograph on the Indo-Malayan strand flora Schimper
(91, pp. 163-178) makes the following classification of strand
plants, based upon the flotation characters:
I. oe und Driftsamen mit grossen luftftthrenden annie Examples:
fru
maritima, Ipomoea pes caprae, Pangium edule (?), cities asiatica (?),
orinda citrifolia.
2. Driftsamen mit schwammigen Samenkern. Examples: many Leguminosae, such
as Sophora tomentosa a sth on of paisa: ‘out Canavalia; embryos of
Rhizophora and Avic
3. Driftfriichte und Driftsa wad deren Schwimmfahigkeit durch luftfiihrendes
halengewebe bedin
A. sates e saat a Examples: Clerodendron inerme, Car rapa
species, nahi subcordata, Wollastonia glabra, ene este argentea,
Pemphis aci
a. phair mit grossen Intercellularriumen. Examples: Cerbera
Odollam, Laguncularia racemosa, Nipa fruticans.
- Schwimmgewege ohne oder nur mit winzigen Intercellularriumen. Ex-
amples: fruits of Cocos nucifera, Barringtonia speciosa, B. excelsa, Ter-
OF THE HAWAIIAN ARCHIPELAGO Zit
minalia Katappa, Conocarpus erecta, Lumnitzera racemosa, L. coccinea,
Scyphiphora hydrophyllacea, Guettarda speciosa, Tournefortia argeniea,
Wollastonia {glabra, Scaevola Koenigii, Clerodendron inerme, Cynometra
cauliflora, Cordia subcordata; seeds of Carapa moluccensis, C. obovata,
cana; seeds of Cyas circinalis and Excoecaria A gallocha.
Guppy (06, p. 531) enumerates the following seeds or seed
vessels that remained afloat after a year’s flotation in sea-water:
Thespesia populnea, Mucuna gigantea, Dioclea sp., Strongylodon
lucidum, Sophora tomentosa, Caesalpina Bonducella, Entada scan-
dens, Morinda citrifolia, Scaevola Koenigti, Cordia subcordata,
Tournefortia argentea, Ipomoea grandiflora, and Tacca pinnatifida.
In Helmsley’s classification of the Bermudian flowering plants
(’85, p. 48) the following indigenous genera, chiefly littoral forms,
are listed as having probably been conveyed to the island by ocean
currents: Cakile, Hibiscus, Suriana, Elaeodendron, Sapindus,
Dodonaea, Cardiospermum, ‘Rhus, Sophora, Vigna, Canavalia,
Centrosema, Conocarpus, Rhizophora, Opuntia, Sesuvium, Rhachi-
callis, Chiococca, Morinda, Solidago, Borrichia, Scaevola, Tourne-
fortia, Heliotropium, Ipomoea, Convolvulus, Avicennia, Coccoloba,
Atriplex, Salicornia, Euphorbia, Croton, Ruppia, Zostera, Cenchrus,
- Spartina, Stenotaphrum, Sporobolus, Chloris.
Helmsley (’84, p- 304) has also recorded the actual germination
of various drifted seeds after being cast ashore. He lists Hibiscus
tiliaceus, Vitis vinifera, Sapindus Saponaria, Anacardium occidentale,
-Aleurites moluccana, Ricinus communis, Cocos nucifera, and Sagus
sp. Of Vitis vinifera he records the foundering of a vessel laden
with a cargo of white Lisbon grapes, off the south shore of Bermuda.
Many of the grapes were washed ashore, and the seeds germinated
at high-water mark. Numbers of plants were taken up, out of
curiosity, and transplanted, and bore fruit. Martins raised plants
from seeds of Ricinus communis that had been floating for ninety-
three days upon the surface of the sea.
Shull’s (’14) extensive experiments show that the seeds of
many species will germinate after four years of continuous sub-
mergence in fresh water, and that the seeds of three species were
viable after seven years of continuous submergence.
Zi MacCauGHEey: THE STRAND FLORA
Guppy (’06, p. 529) shows that of the littoral plants of Fiji and
Tahiti, 75-80 per cent. have seeds or fruits that will float un-
harmed for two months or more, and that about 30 per cent. of
this number are legumes. He says:
n the course of the ages the plants with buoyant seeds or seed vessels have been
gathered at the coast. This is indicated: (1) By the far greater proportion of species
with buoyant seeds and seed vessels amongst the shore plants than among the inland
plants. (2) By the circumstance that almost all the seeds or seed vessels that float
unharmed for long periods belong to shore plants. (3) By the fact that when a genus
has both inland and littoral species, the seeds or fruits of the coast species as a rule
float for a long time, while those of the inland species either sink at once or float only
for a short period. .
Guppy (’06, p. 563) makes the following list of ‘Hawaiian
plants with buoyant seeds and fruits known to be dispersed by the
currents either exclusively or, as in a few species, with the assistance
of frugivorous birds”: Colubrina astatica, Dioclea violacea,
Mucuna gigantea, M. urens, Strongylodon lucidum, Vigna lutea,
Caesalpinia Bonducella, Scaevola Koenigit, Ipomoea glaberrima,
I. Pes-caprae, Vitex trifolia and Cassytha filiformis. Although
Many strand plants possess seeds or fruits that can float for long
periods, other widely distributed species possess feeble or no flota-
tion power. It is necessary to recognize other agencies.
TREES AND LOGS AS DISSEMINATORS
Logs and tree-trunks of various coniferous species from the
Puget Sound region are commonly cast ashore upon the Hawaiian
windward coasts. It is a matter of common observation that on
all windy coasts, small seeds, like sand, are blown into every
available cranny. In this way many lodge in the holes and cracks
in drift-wood, which is floated off at high tide or during storm time,
and thus the seeds or fruits may be carried to new localities.
Strand seeds or fruits which do not possess special flotation devices
may be carried to new shores. Moreover, the seeds of inland
species may be carried by trees which have been uprooted by
inundations or storms, either in the soil around the roots, or in the
bark, etc.
Ernst (’08, p. 56) states that “tree stems and branches played
an important part in the colonization of Krakatau by plants
and animals.”’ Hedley (’15) records a log of Dammara dustralis
OF THE HAWAIIAN ARCHIPELAGO 273
the New Zealand Kauri, as stranding on the windward reef of
Funafuti. Wood-Jones (’05) gives an excellent account of
tree-trunks and ‘“‘floating islands’’ of storm-washed vegetation
as carriers of seeds, animals, etc., to the Cocos-Keeling group.
He emphasizes the importance of trees with buttressed bases as
disseminators:
These buttresses are in ine ee of large thin wings, which taper to the trunk
and below form like stalls in a circular stable. With-
in a stalls much earth is held ae, by the interlacing of smaller roots, and when
such a tree is uprooted, and set adrift to sea, it carries its earth withit. It may carry
it for very great distances, and I have seen a buttressed tree come ashore in the atoll,
from whose base a wheelbarrow-load of fine red earth might have been collected.
FLOATING ROCKS AS DISSEMINATORS
The idea of floating rocks as disseminators of littoral plants
might be met with incredulity, were it not for the testimony of
many reliable observers. Among the volcanic islands of the East
Indies large blocks of pumice float for many weeks, and are carried
many hundred miles from their points of origin. The salient
points—prolonged flotation of the blocks; presence of numerous
kinds of seeds in the crevices and pores of the pumice; and the
germination of these seeds when the block is cast upon a favorable
beach-situation—have all been corroborated by careful investi-
gators. Ernst (’08, p. 56) states that floating blocks of pumice con-
stitute an important dispersal agency in the Sunda-Straits region.
Although there is very little pumice to be found on the Ha-
waiian coasts at the present time, there is abundant evidence that
in earlier periods in the geologic history of the islands, repeated
volcanic explosions, resulting in pumice production, have taken
place. There are today extensive pumice beds aroundsthe vol-
cano Kilauea. Therefore, although pumice blocks play little or
no part in the dispersal of plants in the Hawaiian group at present,
it is entirely possible that they had a more important réle in
earlier times, at least in distributing seed from island to island.
Floating masses of dead coral may also be ranked as possible
seed-carriers. _Wood-Jones (15) found numerous instances of
this in the Cocos-Keeling group. The innumerable air-cavities
in certain kinds of coral render it buoyant. The block is cast
upon a beach at storm time; it lies there for an indefinite period;
274 MaAcCAUGHEY: THE STRAND FLORA
earth, sand, and seeds lodge in its many crevices; another storm
sets it again adrift; and it may be cast ashore upon a distant strand.
Coral blocks of this sort are infrequent on Hawaiian shores, owing
to the relative paucity of fringing reef, and have probably been of
minor significance in seed dispersal. They constitute, however,
a possible factor, soloaepietied on Oahu, Kauai, and the leeward
isles.
COMPOSITION—A CLASSIFIED LIST OF THE
HAWAIIAN STRAND PLANTS
I. True littorals
Species which occur only or chiefly within the strand zone.
ENDEMIC LITTORALS
Trees and shrubs
There are no endemic trees that are strictly littoral. This is a
significant feature of the Hawaiian flora. The shrubs are:
Lycium sandwicense Gray Scaevola coriacea Nutt.
Nototrichium humile Hillebd. Solanum Nelsoni Dun.
Santalum Freycinetianum Gaud. SS. laysanense Bitter
var. littorale Hillebd. Wikstroemia Uva-ursi Gray
Phyllostegia variabilis Bitter
Herbaceous plants
Achyranthus splendens Mart. Lipochaeta connata (Gaud.) DC.
A. splendens var. rotundata Hil- var. littoralis Hillebd.
lebd. L. integrifolia (Nutt.) Gray
Campylotheca molokaiensis Hil- L. succulenta (Hook. & Arn.) DC.
lebd. Schiedea globosa Mann
Fimbristylis pycnocephala Hil- S. Lydgatei Hillebd.
lebd. Sporobolus virginicus (L.) Kunth
Kadua littoralis Hillebd. var. phleoides Hillebd.
Lefidium owathense Cham. & Tetramolopium sp.
Schlecht. ;
2. INDIGENOUS LITTORALS
Trees and shrubs
There are no indigenous trees that are strictly littoral. Shrubs:
Colubrina asiatica (L.) Brongn. Gossypium tomentosum Nutt.
OF THE HAWAIIAN ARCHIPELAGO 215
Heliotropium anomalum Hook. Sesbania tomentosa Hook. & Arn.
& Arn Vitex trifolia L.
Scaevola iets Murr.
Herbaceous plants
Argyreia tiliaefolia (Desr.) Wight Ipomoea Pes-caprae (L.) Sweet
Boerhaavia diffusa L Lepturus repens R. Br.
Cressa cretica L. Lysimachia spathulata Benth. &
Cyperus laevigatus L. Hook.
Euphorbia cordata Meyen Ruppia maritima L.
Heliotropium curassavicum L. Scirpus maritimus L.
Herpestis Monnieria H. B. K. — Sesuvium Portulacastrum L.
Ipomoea acetosaefolia (Vahl) Sporobolus virginicus (L.) Kunth
Roem. & Schl. Tephrosia piscatoria (Soland.)
I. glaberrima Bojer ers
I. insularis Steud.
3. LITTORALS INTRODUCED BY THE PRIMITIVE HAWAIIANS*
Trees
Calophyllum pees i Cordia subcordata Lam.
Cocos nucifera L
4. LITTORALS INTRODUCED SINCE THE ADVENT OF EUROPEANS
(1555-1778 TO DATE)
Trees and shrubs
No true littoral trees and shrubs have been introduced.
Herbaceous plants .
Batis maritima L. Polypogon littoralis (With.) Sm.
II. Pseudo-Littorals
Species which chiefly inhabit the lowlands or other zones, and
which appear on the strands in the role of invaders from the interior,
I. ENDEMIC PSEUDO-LITTORALS
Trees and shrubs
Acacia koa Gray Erythrina monosperma Gaud.
Cassia Gaudichaudti Hook & precios sandwicense (A. DC.)
Arn. Gra
Chenopodium sandwicheum Moq.
* Thespesia populnea Soland. should be noted here.
276 MaAcCAUGHEY: THE STRAND FLORA
Herbaceous plants
Carex sandwicensis Boeckl. Peucedanum sandwicense Hillebd.
Jacquemontia sandwicensis Gray Sicyos hispidus Hillebd.
Nama sandwicensis Gray S. microcarpus Mann
2. INDIGENOUS PSEUDO-LITTORALS
Trees and shrubs
Caesalpinia Bonducella (L.) Pritchardia spp.
Flem Tribulus cistoides L.
M seaside polymorpha Gaud.
Herbaceous plants
Argemone mexicana L. Fleurya interrupta Gaud.
Cassytha filiformis L. Kyllingia monocephala Rottb.
Cenchrus calyculatus (Spreng.) Lythrum maritimum H. B.
Cav. K.
Chenopodium album L. Malvastrum tricuspidatum (Ait.)
Cladium leptostachyum Nees Gray
Cyperus pennatus Lam. Mucuna gigantea (Willd.) DC.
C. phleoides Nees ° Ophioglossum vulgatum L.
Eragrostis hawaiiensis Hillebd. Sida spp.
Erythraea sabaeoides (Griseb.) Vigna lutea (Sw.) Gray
Gray Waltheria americana L.
3. PSEUDO-LITTORALS INTRODUCED BY PRIMITIVE HAWAIIANS
Trees
Aleurites moluccana (L.) Willd. Morinda citrifolia L.
Aibiscus tiliaceus L. Pandanus odoratissimus L. f.
4. PSEUDO-LITTORALS INTRODUCED SINCE THE ADVENT OF
EUROPEANS
Trees and shrubs
Acacia farnesiana Willd. Leucaena glauca (Willd.) Benth.
Casuarina equisetifolia Stickman Prosopis juliflora (Sw.) DC.
Hibiscus Rosa-sinensis L.
OF THE HAWAIIAN ARCHIPELAGO Ze
Herbaceous plants
Abrus precatorius L.
Achyranthus aspera L.
Cardiospermum Halicacabum L.
Crotalaria spp.
Cynodon Dactylon (L.) Pers.
Cyperus umbellatus Vahl
Datura Stramonium L.
Euphorbia pilulifera L.
Euxolus viridis Moq.
Hydrocotyle verticillata Thunb.
Indigofera Anil L.
Mesembryanthemum spp.
Portulaca oleracea L.
Salvia occidentalis Sw.
Samanea Saman (Benth.) Merrill
Xanthium echinatum Murr.
There are numerous ruderals, in addition to those indicated
in the last section, that occur at random on the various beaches.
COLLEGE oF Hawaltl, HONOLULU
Notes on the fern genus Clathropteris
Epwarp W. BERRY
(WITH TWO TEXT-FIGURES)
Among the more fascinating objects of paleobotanical investi-
gation are the abundant and varied forms which have now come
to be rather generally recognized as constituting two distinct
families of ferns, the Matoniaceae and the Dipteriaceae, members
of which are such characteristic and striking objects in Mesozoic
fern floras. This interest associated with their far distant an-
cestry is heightened by the fact of the singular association of the
few surviving representatives of these two families at a limited
number of localities in the oriental tropics.
It is not my purpose, however, to attempt an elaboration of
this subject in the present connection, since it has already been
discussed by others* and there is, moreover, a rather extensive
literature dealing with the different extinct generic types that
seem to be referable to the one or the other of these families
All that will be attempted in the present brief contribution will
be the placing on record of certain observations on new material
belonging to the genus Clathropteris and a discussion of its bear-
ing on the probable habit of these ferns.
During a visit to the Richmond (Virginia) coal field during
1911 I collected for the United States National Museum a re-
markably fine specimen of the so-called Clathropteris platyphyiia,
which, in so far as I recall, was the most complete specimen of this
ubiquitous form that has ever been collected. This specimen
was about 40 X 55 cm. and showed several dichotomies of the
stipe. During its shipment to Washington the edges were broken
and the surface abraded so that only a very inferior specimen
remains. A counterpart of a portion of the face of this specimen
“* Seward, A. C. On the structure and affinities of Matonia pectinata, R. Br.,
with notes on the geological history of the Matonineaer Phil.
&
Trans. Roy. Soc.
Lond. B, 191: i hae — Seward, A. C., & Dale, E. On the structure and
affinities of Dipter the geological hist of the Dipteridineae. Idem.,
B, 194 : 487-513. I90r.
279
280 BERRY: FERN GENUS CLATHROPTERIS
was carefully brought back for the collections of the Johns Hopkins
University and it is upon this fragment that the following re-
marks are mainly based.
The latter has maximum dimensions of about 22 X 27 cm.
and shows a fragment of a large stipe 12 cm. long and 1 cm. in
diameter, part of a whorl of large pinnae, and a fragment of the
terminal part of a stipe with eight palmately arranged and mostly
attached pinnae. Some of the marginal dentations of the pinnae
are perfectly preserved, as is the peculiar netted venation. This
specimen is shown in FG. 1, one half natural size.
The genus Clathropteris was proposed by Brongniart* in 1828,
the type being his Filicites meniscioides} from the Rhaetic beds
of Hoer in Scania. In 1849 Brongniart{ transferred the Camp-
topteris platyphylla of Goeppert§$ to the genus Clathropteris, and
these two species have usually been maintained as distinct, al-
though Nathorst states|| that after an examination of Brongniart’s
type material of Clathropteris meniscioides he is convinced that it
is identical with Clathropteris platyphylla. If this is true then the
former name has priority.
The genus has been discussed recently by Nathorst (op. cit.),
Zeiller** and Seward7j{ so that it is unnecessary to attempt to rede-
fine it at the present time. Seward{t{ has advocated the merging of
Clathropteris and the allied genus Thaumatopteris Goeppert (op. cit.)
with Dictyophyllum, but this course has fortunately not been fol-
lowed, and Nathorst (op. cit.) has given excellent reasons why such
a consideration would be unwarranted. The last author proposes
that these fossil genera should be segregated from the existing
family Dipteriaceae under the family name of Camptopteriaceae
and on philosophical grounds it would seem that such a course
would come nearer to representing the true status of these forms
since the two groups are separated by the whole time interval of the
* Brongniart, A. Prodrome, 62. 1828.
+ Brongniart, A. Ann. Sci. Nat. Bot. 4: 200. pl. rr. 1825.
t Brongniart, A. Tableau 32. 1849.
§ Goeppert, H.R. Gen. pl. foss. 5-6 : 120. pl. 18, 19, f. I-3. 1846.
|| Nathorst, A.G. Kgl. eccuae Vetens. ~Akad. Handl. 41:4. 1906.
** Zeiller, R. Flore foss. gites charbon Tonkin. Ministére trav. publ. Etudes
gites minér. France, Atlas, 1902, texte, 1903. :
tt Seward, A. C. Fossil Plants 2: 386. 1910.
tt Seward, A.C. Phil. Trans. Roy. Soc. Lond. B, 194 : 503. 1901.
BERRY: FERN GENUS CLATHROPTERIS 281
Cenozoic, generally estimated at several millions of years, during
which it is almost inconceivable that family boundaries did not
shift. At the same time there seems to be a consensus of opinion
that the existing Dipteriaceae represent the last relics of this
adaptive radiation of the Camptopteriaceae, so that the question
of family nomenclature is really not of great importance.
Specimens referable to Clathropteris and probably representing
several botanical species, but not certainly distinguishable in the
present state of our knowledge, have a very wide geographic and a
very considerable geologic range. In this country they are found
in the rocks of the Newark formation, probably of Keuper age, in
Massachusetts, Connecticut, New Jersey and Virginia. They
occur in the Keuper of Prussian Saxony and Switzerland. In the
succeeding Rhaetic they are found in Sweden, Bornholm, Germany,
France, England, Persia, China and Tonkin. In rocks referred
to the lower Lias they are recorded from Hungary, Saxony, Silesia
and France. Owing to the peculiar habit of these ferns and the
often great length of the pinnae the specimens are usually much
broken, the best foreign material probably being that described
by Zeiller (op. cit.) from Tonkin.
As regards the habit it appears that the dichotomously forked
rhizomes described by Nathorst (op. cit.) as Rhizomopteris cruciata
represent the rhizomes of Clathropteris. The scars on these
rhizomes indicate that in the Swedish region the fronds were not
as crowded as they were in the allied Dictyophyllum growing at the
same locality. The Virginia material shows that the stipes were
stout and somewhat curved (in this respect suggesting Mertensia),
as much as a centimeter in diameter, and with a longitudinally
striated epidermis.
These stipes, rising for a considerable ditcare from the creeping
rhizome, divided dichotomously at a wide angle and bore on the
upper side of this fork from ten to thirty pinnae as in the genera
Dictyophyllum and Camptopteris, species of both of which genera
have been admirably restored by Nathorst. These pinnae are
said to be fused proximad but it may be considered certain that
the amount or absence of fusion was a variable feature as it is
demonstrated to have been in the allied genus Dictyophyllum.
It appears from Zeiller’s Tonkin material that these primary pinnae
BERRY: FERN GENUS CLATHROPTERIS
282
are more inclined to separation, are more slender, have simpler
margins and are more numerous than those pinnae which I pro-
pose to consider as ultimate or secondary pinnae. I regard the
tyse
lee,
Me,
MAS
TA
uN Z
aan f SOR Mr
NN, SGML
SS SUAS NN SST 7 NG 2a ane
ma LN BN) BI 0 ar 8 0 om
SAAN UE Ng MLZ Mo lly 2:
WRIGHT = Cee
ROSARY My od The peserce
SOREN nel J 2222745.
: : ey
if gj
LZ
|
SN
SS
SSS
LX
ty
;
vi
“|
oY,
I)
|
‘ |
as
We;
Dah
LU
9
ss
a
i“
2
es
Oe:
ale
Ls
a
ge
4
3
VB
4
3
ay
é
MS
<
\
Li
7]
TIL
g
e yf}
Al)
HTL
ae
=
>
SQ
ge)
SS
_ ay
LTR, 1G
RE cai
Ae
eB
Lo
tl //
E]
My,
age,
Mase
£7
ae
SF,
LL
U/7
wh
7)
Let
|
27.
EF |
Ae
wane
22%
Patt
sae
Be:
c/
Lt:
Li
<1
B
Ne
ATT
Fic. 1. Specimen of Clathropteris platyphylia from Triassic of Virginia, one half
natural size. :
palmately arranged pinnae on the right of the specimen figured
These are lan-
(Fic. 1) as representing such primary pinnae.
ceolate in form, show a maximum width of 5 cm. at about one
BERRY: FERN GENUS CLATHROPTERIS 283
third the distance above the base, and have an indicated length
of about 30 cm. They taper conspicuously proximad and are
still free at a point where their width has narrowed to less than
1.5 cm. Although their extreme bases are obliterated they were
evidently either entirely separate or only slightly united. The
margins are not entire but the marginal teeth are very much re-
duced, even at a distance of 10 cm. above the base, and proximad
they appear to have been entirely wanting, although the preser-
vation is not sufficiently good to be positive on this point.
It has been assumed by most students that the Clathropteris
frond consisted of a single dichotomy and therefore had a habit
somewhat like the modern Dipteris and more conspicuously like
the existing Matonia pectinata, or like that of the extinct genus
Dictyophyllum as restored by Nathorst. It would appear from the
Virginia material that, in addition to such a group of what I
have called primary pinnae, arranged en éventail, each branch of
the dichotomy continued for some distance and then expanded
into palmately arranged ultimate or secondary pinnae. These
were fewer in number than those that I have called primary pinnae,
being eight in the preserved material. They are wider and more
conspicuously toothed and are clearly united for a distance of 3-5
cm. above their bases in my material.
While this interpretation is based in the first instance on ma-
terial no longer extant and is therefore to be accepted with reserve,
I feel justified in calling attention to it and in offering the tenta-
tive restoration of this interesting species as shown in Fic. 2
That the fronds were sometimes much larger than I have in-
dicated is shown by fragments in my possession and by the
extremely large fragments collected by F ontaine, which according
to this author showed a width of pinnae of 20 cm. and an indicated
length of 60 cm.
I have seen no fertile specimens from Virginia, te certain
foreign material shows, on the lower surface of the lamina, an
abundance of crowded sori without indusia and made up of from
five to fifteen annulate sporangia, similar to those of Dictyophyllum
and Dipteris, although the annulus is said by Goeppert to have
been complete—a feature that it would seem would be impossible
to decide in the absence of structural material.
284 BERRY: FERN GENUS CLATHROPTERIS
The venation of Clathropteris has always excited great interest.
In the proximal region where the pinnae are fused it is somewhat
irregularly anastomosing, as is admirably shown in Zeiller’s
vi
6 es ay of
hs BONE by
it Ai, of Vi Vat
1 OM | hs ory AG
Sy AG ft OP Me iia
bids AD ki f A Lhd
" L] i iy
NS : ae Ne ie Ne ry We ded lig ee
SH =i sant ea Es Hama Sta la ie a vee
ANA \ he ai a a a a Wo Ma yp TS V2
i ys ae he OG
tata c I yA aa Ya CANIN ray HE
ARON a oy A a aa RE ls sy
AULA Re te EO CAM as Cine ey
oe sy ee ae, ry Wi LE p
QS aN ye Chee LGD pay CHE LZ tg ess
WX A QE ‘ ann ae ee KEE LEG ie
QS RY ey Be ay hi ie” Ss Le
. Mt gst 3e U cf a
SSN W ca
SSS 5 LE BES
x Ne
KY
ee = SS >
ee ee =
LES ae
SSS
eS
Ye
~
x
xy
AY
Saree ere Se,
——
H
HH 4
a I
1)
t /
y |
l
Vi
Sse
SELES
=— ——
A =e
Cg on ee _.
Si ey LEE: LOY = SN
4 7 Ly SN ? f => “SEES
Lh? MGA JEE cS, SS
ti P toe HU fe ea A ‘
j y ay BO
Lip LS LR, Lee Lyf Gite SINS
Lh GER We
1G. hi so NS Swng
iy ip by: wal a Wh
aap yRiey PK SS
Ny ye P oy a
Y TK) LUT)
yh i: x ay
Ly
hs
4
=
oa
=,
4
Fic. 2. Tentative
restoration of Clathropteris platyphylla, one eighth natural
restoration.* In the free portion of the pinnae is very stout
midribs give off, at regular intervals and at very wide angles ap-
* Zeiller, R. Eléments de Paléobotanique, 116, f. 89. 1900.
BERRY: FERN GENUS CLATHROPTERIS 285
proaching 90 degrees, stout parallel secondaries. These usually
alternate but not invariably. They proceed outward in almost
straight courses nearly to the margin where they curve conspic-
uously upward, becoming rapidly much reduced in size and ter-
minate at the tips of the marginal teeth. The secondaries are
connected at approximately regular intervals by relatively stout
straight percurrent tertiaries at nearly right angles to the second-
aries, and subordinate divisions result in an ultimately fine rec-
tangular areolation with blind endings.
This rectangular venation is rather consistently more regular
than in the allied fossil genera and rather different from that of
the existing Dipteriaceae. It is, however, approached very closely
by certain existing Polypodiaceae with the so-called Drynaria
composita type of venation. Among modern ferns that have a
comparable venation might be mentioned various oriental species
of Polypodiaceae, belonging to the genera Lomariopsis, Dryostach-
yum, Polybotrya and Drynaria. The latter genus is especially
like Clathropteris in the form of its pinnules, in their venation and
(in some of its species) in their toothed margins. Drynaria
comprises about a dozen species of epiphytes of the oriental tropics
and the species Drynaria quercifolia is particularly like Clathrop-
teris in respect to the characters just enumerated, although the
general habit is very different.
Although Schenk refers the fossil forms to the family Dictyop-
terideae, the venation characters, more readily ascertainable from
fragmentary specimens, which are the kind usually collected, have
resulted in the usual reference of the fossils to the Polypodiaceae
or Acrostichaceae, as is done by Ettingshausen, although it is
obvious that they constitute a unique and distinct line of forms
ancestral to the modern family Dipteriaceae.
JoHNs HopKINS UNIVERSITY,
BALTIMORE
Studies in the genus Gymnosporangium—ll, Report on cultures
made in 1915 and 1916
B. O. DopGE
(WITH PLATE 8)
The writer (2) reported the infection of Chamaecyparis with
Roestelia transformans, and later (3) gave an account of experi-
ments which resulted in the infection of the red cedar with G.
clavipes, G. macropus, G. globosum and G. nidus-avis. . Arthur’s
results (1) were confirmed in the case of G. clavipes, and it was
stated that the other species required nearly two years to mature.
G. globosum as well as G. macropus may develop strictly foliicolous
galls. In 1917 the writer reported the infection of Chamaecyparis
with G. Ellisiit (4). Weimer (12) states that G. clavipes matures
in two years, although it seems he was unable to make the infec-
tions. He also reports the development of one small gall of G.
macropus on cedars which he had previously inoculated. The
abstract of the writer’s paper read at a meeting of the Botanical
Society of America, 1916, has not appeared in print, and as further
work has resulted in the accumulation of considerable data, a
summary of the cultures of 1915 and 1916 is presented at this time.
GYMNOSPORANGIUM CLAVIPES
A red cedar eight inches high obtained from Cold Spring
Harbor, Long Island, in May, 1915, had at the time a few sori of
G. clavipes on the stem. This material was used to infect Crataegus
oxyacantha.
Six red cedars were sprayed with aecidiospores August I, 1915,
and left in the infection frame two days. Several other cedars
were growing in the same greenhouse, otherwise no controls were
provided. TasLe I includes all cedars that were growing in the
greenhouse at the time aecidiospores were being shed from the
rust on the Crataegus.
The results shown in the table indicate that spores may mature
the first spring after inoculation, although in some cases they do
287
288 DopGE: STUDIES IN GENUS GYMNOSPORANGIUM
not appear until the second year. All of the plants that were
inoculated became so heavily infected that by 1918 many of the
branches had been killed. Those plants that became infected
without inoculation through being exposed near hawthorns bearing
aecidia bore only a few sori in 1917 and had about twice as many
in 1918. The invasion of new regions of the host by the mycelium
TABLE I
INOCULATION OF Juniperus virginiana WITH AECIDIOSPORES OF G. clavipes
Plants inoculated August 1, 1915 Ced — d Sg Sahar — _—
No. Results, 1916 | Results, 1917 No. Results, 1916 | Results, 1917
401 | — | +, many sori | 405 - | +, 6 sori
i ad Blea See Bate eae 407 =
403 hae +, se “cc 408 wre cigs
4I2 +, II sori ee 4 411 _ -
415 +, 30 sori mk eel fe 938 — _
416 ‘+, I sorus “hy 2 413 - _
414 _ -
417 - _
418 — +, 2 sori
590 cm aap
591 ce -
594 as =<
609 - _
is not very rapid. It has, however, completely connected spaces
between most of the groups of sori that were evident on the plants
in 1917. In some cases small witches’ brooms have been formed,
and spindle-shaped swellings are beginning to appear on some of
the larger branches and main stems.
GYMNOSPORANGIUM MACROPUS
The life histories of the two ‘‘cedar apple” rusts have become
fairly well understood through field observations made by many
investigators of the gymnosporangial stages, and by cultures of the
rusts on their aecidial hosts.
The cedar plants used in this work were obtained at Cold
Spring Harbor, New York, February 1,1914. Some of them bore
a few galls of G. macropus and G. globosum. Spores from these
galls were used to infect seedling apples and hawthorns. The
galls were marked with tags for further observation. The cedars
were carefully inspected during the summer of 1914 and until
DopGE: STUDIES IN GENUS GYMNOSPORANGIUM 289
July 1, 1915, and any new galls that appeared were marked with
tags. The heights of the plants were taken in February, 1914, and
each spring since that date. These precautions enabled me to
obtain evidence relating to the length of life of the galls, especially
of G. globosum, which has been assumed to be perennial, although
records bearing on this point appear to be meager. Hawthorns
bearing aecidia of G. globosum were grown in the greenhouse at the
same time and this fact required that infections be made with
teleutospore material from galls that appeared on the cedars later,
since the smaller galls of G. globosum and G. macropus are not
always so characteristic as to be easily distinguished. The results
are given in TABLE II.
TABLE II
INFECTION OF THE RED CEDAR WITH G. macropus
Height of cedars, inches
N | Date of | Results, number Ss
o- Feb., Feb. Feb., Feb., | inoculation of galls } pores matured
| tOr4 1915 1916 19*7
O14 |
401 II 16 24 z July 13 fs)
403 7 12 206 35 June 13 2 | March 17, 1916
504 6 10 I5 23 June 7 3 1916
4II 14 24 39 53 June 22 ts)
414 16 24 39 gs June 20 2 Ma 18, 1916
415 9 13 rg 20 June 28 2 April 2, 1916
417 16 at 34 48 June 20 ra) :
403 7 12 26 35 I9I5 3 April 14, 1917
july . 2
407 I2 22 35 50 Aug. I 6 April 24, 1917
416 pas) 14 17 24 June 5 | 2 April 25, 1917
418 yg ? 20 42 June 10 I March I, 1917
The last date upon which the plants were exposed to natural
infection was in the summer of 1913. The point at which the
lowest gall, for example on plant No. 403, appeared was five inches
above the tip of the main stem of this plant February 1, 1914, as
shown by the records of measurements. The cultures, especially
those made in 1915, prove that Heald’s conclusions were correct
as regards the time required for the maturity of the rust. There
were fourteen galls on these four plants; they were for the most
part comparatively small, bearing from one to five sori.
290 DopGE: STUDIES IN GENUS GYMNOSPORANGIUM
GYMNOSPORANGIUM GLOBOSUM
The inoculations of cedars with G. globosum were made in
1914 and 1915 under the conditions described in the preceding
experiments. The results appear in TABLE III.
TABLE III
INFECTION OF THE RED CEDAR WITH G. globosum
Height of cedars in inches
No. SU meas a et er hte ot. Results, number d
°. Feb. Feb., Feb., Visi Viancchation at pals Spores mature
IgI4 1915 1916 1917
IOI
405 6 sae) I5 a3 June I5 March 7, 1916
407 12 22 35 50 June 25 5 Feb. 28, 1916
408 II 20 30 48 June 20 (0)
IQIS
401 II 16 24 38 Aug. I ts)
402 14 ? 26 a7 Aug. I oO
403 7 I2 26 35 Aug. I 2 March 10, 1917
405 6 10 15 23 Aug. I I April 4, 1917
416 Io 14 +7 24 Aug. I te)
The first indications of infection were discovered August 1,
1915; there were two small green galls on plant No. 407.
The writer is rather inclined to disregard these galls, especially
as they were on the lower branches and grew to considerable size
like those one finds in nature. There were no sorus-scars such as
we should expect to see if the plants had been infected in nature in
1913. The other galls were clearly the result of inoculation as
most of them developed at points on the plants represented by
new growth since they had been brought in. None of the fifteen
leaf galls on plant No. 405 developed sori a second time. The ones
on No. 407 matured spores two seasons. Most of the large galls
on naturally infected plants, so far as observed, bore sori three
years in succession. There is no record of one bearing four crops
of spores. It was impossible to inspect the cedars thoroughly
during July and August, so that it is uncertain at just what
time the galls made their first appearance. Most of them were
visible in September, although some of the galls of G. globosum
were so small that they might have been overlooked. In some
cases there is very little hypertrophy of the leaf tissue upon which
they appear. Weimer ieee has very adequately described these
smal] galls.
DopGE: STUDIES IN GENUS GYMNOSPORANGIUM 291
GYMNOSPORANGIUM ELLISII
Fromme (6) connected Gymnosporangium Ellisii with a oosatvai
myricatum. Dodge and Adams (5) found the aecidium on
Comptonia asplentfolia. Attempts were made to infect Comptonia
in 1915 and 1916 without success. The plants brought in March
25, 1917, leaved out in about two weeks and were inoculated April
12. The spermogonia appeared April 1 on several leaves. By
May 14 the peculiar ram’s horn twist of the leaves was very strik-
ing. Absence prevented further observation of the infected plants
but at least two of them matured aecidia.
Certain seedling plants of Chamaecyparis were sprayed with
aecidiospores from Myrica in 1915 and othersin 1916. The cedars
in the first lot showed no signs of infection in 1916, three years
after they had been exposed to natural infection. The second lot
consisted of small seedlings, one to three inches high. The results
of these experiments are given in TABLE IV.
TABLE IV
INFECTION OF Chamaecyparis thyoides wiTH G. Ellisit
A. Inoculation, 1915
fi Results
No. ys orton | Controls
1916 1917 1918
345 June 21 — + 5 Infec + Eighteen xepac A prions: sat
349 June 21 -_ ~ - exposed in the e greenhou
400 June 21 = oe + s greed rat
404 June 21 - - _ spores were being shed, but they
406 June 21 _ peg + | were n y
410 June 21 _ Se ee + Sear chambers. Only
420 June 21 - i401 , No. 431, bécame “Se 5
426 June 21 — - - On ne sorus sri d on young
433 June 21 a en nae | + | growth of a branc
436 June 21 - - | =-—
B. Inoculation, 1916
906 April 24 - | _ oo | sy eg similar plants
907 April 24 oe + + | were sed, but not pada
908 April 24 eee. _ + /in it infection ram me. No
909 April 28 ete + | fection has appeared on paces
914 | May 4 ee oe + | plants.
O15 May 4 eet rote - 4
916 May 4 —- |- + |
918 May 17 - | — —
953 June I ~ Plant died
954 June 1 | oe =
955 ex os +
956 June 1 - a oe
957 June 1 - aS a4
958 June I _ = ee
292 DopGE: STUDIES IN GENUS GYMNOSPORANGIUM
The table shows that eighteen of the twenty-four plants
inoculated became infected, one of them in fourteen different
places. Sori matured about twenty months after inoculation,
except in the case of plants Nos. 907, 908, and 909, where sori
developed the year following inoculation. It is possible that in
some cases this species may mature in one year especially where
inoculations are made as early as April 24.
In many cases there is but little hypertrophy or distortion of
the leaves or twigs when the sori are first formed. There is a
slight bending of the tip of the branch and a cushion-like swelling
is developed beneath the sorus. After another year a witches’-
broom of considerable size is formed, or if the main stem is infected
a slight spindle-shaped swelling occurs. Sori may be foliicolous
with the primordia in the leaf tissue, but in all cases the mycelium
penetrates into the wood of the branch.
GYMNOSPORANGIUM CLAVARIAEFORME
The telial stage of G. clavariaeforme was obtained from New
Haven, Connecticut, through the courtesy of Dr. Clinton and
Dr. Nichols. Several plants of Amelanchier and other pomaceous
hosts were inoculated May 9-13, 1916. The most abundant
aecidia were produced on Amelanchier canadensis and A. inter-
media. If spores are floated on water in damp chambers the per-
centage of germination is ordinarily about 95 per cent.
Five plants of Juniperus communis had been obtained from a
nursery in New Jersey in 1915. They were about one foot high.
Six smaller plants, three to six inches high, came from Glen, New
Hampshire. Nine of these junipers were sprayed with spores in
June, 1916. They were taken from the cold frame March 7, 1917.
Two plants from New Hampshire were used as controls and showed
no signs of being infected. Minute sori were discoveted on three
plants in April. No further inspection was made from May 14
until September 10. At this time slight swellings along the stems
of two of the infected plants could be distinguished. Three of the
nursery plants died before January 7, 1918. The results of these
inoculations are given in TABLE V.
DopcGE: STUDIES IN GENUS GYMNOSPORANGIUM 293
TABLE V
INFECTION OF Juniperus communis WITH G. clavariaeforme
No. Date of inoculation sei 7 cau w noted 1017 Results January 7, 1918
I916
925 June 10 Plant died gee
026 June 6 dl infectio f Plant died 1917.
931 June 6 e infec rosa Two infections.
932 June 6 No: signs of infection. Plant died 1917.
9033 une 6 Soaee Cong es “e One infection.
959 June 29 Two infections, lost Three infections.
many leaves.
June 29 No signs of infection. One infection.
961 June 29 One infection. Two infections.
962 June 29 No infection. No infection.
There were nine separate infections in all. One of the plants
from New Hampshire was not infected. The two control plants
remained uninfected. Sections of three of the swollen stems were
made January 17, 1918, ten days after the plants had been taken
from the cold frame. It was found that teleutospores were being
formed, although the sori had not yet broken through the bark.
In sections even at the lowermost points of the swellings there
appeared only two annual rings of wood. There were at first no
signs of infection but minute sori with a few spores each developed
on three of the plants during the last of April, 1917. The fact
that sori formed the first year may be so small as not to break
through the cork or epidermis, and therefore not be detected,
should always be considered. Sections made the next year will
show the cork callus at the point where the sorus was located. The
infection of these plants undoubtedly occurred on the new growth
of the spring of 1916. These cultures do not furnish much evi-
dence that, as Plowright maintains (9), sori are not produced the
spring following the year of infection. Tubeuf (10) gives a de-
tailed and quite convincing account of his cultures of the species
and there is no doubt that the incubation time may vary. This
period does not appear to be absolutely fixed in the case of the in-
fection of the red cedar with G. clavipes and G. Ellisii, which the
writer has previously noted, and it will be shown later that the in-
cubation time varies when cedars are infected with G. nidus-avis. .
294 DopGE: STUDIES IN GENUS GYMNOSPORANGIUM
GYMNOSPORANGINM JUVENESCENS
The telial stage of G. juvenescens was obtained from Dr. J. J.
Davis, Madison, Wisconsin, in April, 1916. Sori were present in
the axils of the acerose leaves, but the witches’-broom effect was
not pronounced. Three or four plants of Amelanchier intermedia
and A. spicata that were in bloom at this time were inoculated.
Infection was evident in five days. Aecidia matured in abundance
on fruit and leaves as early as May I.
Nine red cedars were inoculated in 1916. No infection ap-
pears to have followed these inoculations; there are as yet, April,
1918, no indications of swellings or the development of sori.
GYMNOSPORANGIUM NIDUS-AVIS
The red cedars on Long Island and in the vicinity of New
York are badly infected with a Gymnosporangium which, if the
infections are due to a single species, is certainly multiform in its
manifestations. Amelanchier and Malus have been infected with
spores taken from each of the following forms, although the inocu-
lations were not made in each case with a single sorus: (1) Sori in
the axils of densely crowded acerose leaves; (2) sori caulicolous,
large branches forming coarse witches’-brooms; (3) trunk infec-
tions, sori appearing in deep fissures in the thickened bark. There
is perhaps another form recognized by the presence of long parallel
cork ridges, about one centimeter in width, that mark the location
of sori of former years.
Several apple seedlings had been infected with spores from
material resembling the second type mentioned above. A con-
siderable number of aecidia matured. Spores from these aecidia
were sowed on a red cedar, No. 418, in June, 1914. No infection
was discovered on this cedar in 1915, but two sori developed on
the main stem in May, 1916. In 1917, even on this small plant,
the swollen sori coalesced in masses two inches long, The infec-
tion has spread over five inches vertically during the past three
years. Old and new branches growing from the infected portion
of the stem appear not to be infected at all. It is a typical trunk
. infection, and slightly erst: seen and Malus have
been infected wit two years in succession
and there can be no » dame that both the shad bush and the apple
are host plants for this form of G. nidus-avis.
DoDGE: STUDIES IN GENUS GYMNOSPORANGIUM 295
Twelve red cedars were sprayed with spores from G. nidus-avis
on Amelanchier in 1916. Only three of the plants have become
infected, one of these in five different places. Cedar No. 414 was
inoculated with spores from Amelanchier No. 878, June 11, 1916.
Three small tongue-shaped sori were found among the young leaves
of a side branch on April 5, 1917. This branch has since died.
Cedar No. 929 had not been exposed to infection since 1913.
The original teleutospore material with which the experiment was
begun was obtained at Fort Lee, New Jersey, May 10, 1916.
The infected branch, about two inches in diameter, had a very
rough appearance and was covered with corky mounds and
swellings. The sori when swollen were tremella-like and about an
inch long. Amelanchier “‘canadensis’’ No. 878 was sprayed
May 14. Spermogonia appeared on the leaves May 21, and aecidia
were fully matured on the fruits June 9. Only a few aecidia
developed on the leaves, although they had previously borne
a great many spermogonia. No. 929 cedar was then sprayed
with aecidiospores June 9; it was put in the cold frame October 22
and taken out March 7, 1917. A swelling and distortion of the
main stem at the tip was plainly visible about November 1, al-
though no sori had been noticed during the early months of spring.
This plant was brought back to the green house again January 6,
1918. Sections of the swollen region were made January 17,
1918 and showed two annual rings of wood and two developing
sori. No callus scars have been found on these sections, showing
that a sorus had developed in 1917. In this case the resulting
infection was just about of the type we should expect, and we can
imagine it might have in years come to look much like the original
infection with which we started. Photographs and specimens of
all stages have been preserved.
The history of the infection of cedar No. 609 is more interesting.
The teleutospore material was obtained from the largest infected
red cedar on the grounds of the New York Botanical Garden.
The trunk of this tree is heavily infected for a distance of several
feet and has developed a large spindle-shaped swelling three feet
long about ten feet above the ground. Amelanchier “ canadensis’
No. 886 was sprayed May 17, 1916. Spermogonia appeared May
26 and aecidia ripened on the fruit June 12. Red cedar No. 609
296 DopGE: STUDIES IN GENUS GYMNOSPORANGIUM
(six inches high June 6, 1915) was inoculated June 21, 1916.
On March 22, 1917, twenty-one days after it had been taken from
the cold frame, one very light-colored orange-yellow sorus de-
veloped in the axil of the leaf. This was supposed to be a sorus of
G. clavipes that resulted from accidental infection in 1917. The
little branch was marked with a tag. The plant was taken from ~
the cold frame again January 7, 1918. On February 15, there
were two light-colored sori in the axils of the leaves of the tagged
branch. February 19 two more sori were discovered in the axils
of leaves of another branch, and two strictly foliicolous sori on
opposite leaves of another branch. March 5 another foliicolous
sorus appeared on another branch, and at the base of a larger
branch a small brown sorus was found, upon removing a piece of
loose cork. All of the leaf sori were very light colored and looked
like G. clavipes. The mycelium has been traced from the infected
leaves down into the branches for about two centimeters. The
infected branches are all on parts of the plant that have grown
since it could have been exposed to natural infection. I have
proved by the examination of spores, by sections of infected leaves
and branches and by infection of the shad bush that this cedar
was infected in 1916 with G. nidus-avis. Some of the infections
appeared one year after inoculation, while it undoubtedly re-
- quired two*yearsefer other sori to mature. This is the fourth
species of Gymnosporangium which I have found requiring either
one or two years in which to mature the teleutospores. Under
favorable conditions the rust may develop in one year.
GYMNOSPORANGIUM TRANSFORMANS AND G. FRATERNUM
Kern (8) describes G. fraternum as an annual species. The
writer (3) called attention to the fact that there are two leaf forms
of this genus on Chamaecyparis, both of which are frequently
perennial, and in another paper (4) showed how the two forms
could be distinguished by the character of the buffer cells that
precede the teleutospores in the sori of each.
Blueprints were made of infected branches so that leaves
bearing sori could be located on the prints. In many cases it was
found that sori developed a second and third year on the same
leaves.
DopGE: STUDIES IN GENUS GYMNOSPORANGIUM 297
To prove that the species were distinct, spores from individual
sori were germinated and smeared on selected leaves of both
Amelanchier and Aronia. Slide mounts of the spores were then
preserved and photographed. A few of the photographs are
shown in PLATE 8. TABLE VI, A and B, is arranged to show the
effect of inoculating both trial hosts with spores from the same
sorus.
TABLE VI
PARALLEL INOCULATIONS OF Amelanchier AND Aronia WITH INDIVIDUAL SORI FROM
LEAVES OF Chamaecyparis, 1916
A. G. fraternum
Date Amelanchier Result 1 Aronia Result | Source of sorus
|
Feb. 22 491 = | 642 — 435-3"
Ap. 18 558 + 806 _ 721
Ap. 18 556 ~ 808 - 721.10
Ap. 18 651 - 809 — 9721.5
Ap. 18 807 + 344 _ 721.8
Ap. 18 810 _— 256 — 721.4
Ap. 18 485 + 250 _ 7AItt
Ap. 20 820 + 818 _ 721.3
Ap. 20 554 + 819 = 721.2
p- 20 821 Be 822 7at7
May 8 864 + 456 _ 740.1
ay 8 484 7 470 746.2
B. G. transformans
Feb. 22 441 aes 465 “ 435.2
Feb. 26 438c 452 + 423.1
Feb. 22 503 643 wt 435.2
Feb. 26 552 a 452 me 423.2
I 553 - 454 7 424.2
Mch. 7 649 an 518 a 400.2
Mch. 3 4304 oe 645 = 424.1
Ap. 27 653 ae 457 + 701.0
Ap 9 653 = 652 ri 437-1
7 651 - 650 - 437-2
My. 10 654 _ 808 + 710.1
My. 17 800 = 806 738.1
Referring to the table it can be seen under “A”’ that G. frater-
num infected nine of the twelve amelanchiers inoculated, while the
aronias were not infected. Under ‘“‘B” where the thicker-walled
teleutospores, G. transformans, were used seven Aronias were
infected, five gave no results, and none of the amelanchiers was
infected. In addition to the cultures shown in the table, 32
amelanchiers have been heavily infected by spraying with care-
- Means that the sorus used was sorus No. 3 from plant No. 435.
298 DopGE: STUDIES IN GENUS GYMNOSPORANGIUM
fully chosen sori of G. fraternum and 50 aronias with G. transfor-
mans without the infection of control plants used in either case.
So far as color, size and thickness of walls are concerned, one
finds sori made up of intermediate types of spores not so easily
identified. Compare Fics. 3 and 5 (G. transformans) with Fic. 7
(G. fraternum). On the contrary, such types as are shown in
Fic. 6 (G. transformans) and Fic. 8 (G. fraternum) are easily dis-
tinguished. Spores of G. biseptatum are shown in FIGs. 9 and 10
for comparison. G. fraternum does not become G. biseptatum
when grown in the greenhouse.
INFECTION OF CHAMAECYPARIS WITH G. TRANSFORMANS
The ninety-eight seedling plants of Chamaecyparis described
in connection with cultures of G. Ellisii (Table IV, B) were used
for this work. None of them developed rust in 1916. Most of
them had no branches and only subulate leaves when brought in
(1915). All new growth could be determined readily. Twenty-
four plants were inoculated and seventy-four were kept in another
greenhouse as controls. The results of these experiments are given
in TABLE VII.
TABLE VII
INFECTION OF Chamaecyparis witH G. transformans IN IQI6
ee Pe Fae esults, i
No. ¥ “var oe me ag Controls
615 May 2, Aug. 28 5 Plants numbered 610-
617 June 1, Aug. 28 Ir 614, 616, 620, 625, 629,
618 June 1, Aug. 28 2 790-798, 906, 908, 909,
619 July 16, Aug. 28 ‘3 914-921, 948-9 , 963-
621 June 1, Aug. 28 z 992, 1012, 1014 re-
622 Apr. 24, Aug. 28 I mained entirely free from
623 July 23, Aug. 28 5 this rust in 1917 and 1918
624 pr. 24, Aug. 28 3
626 Apr. 24, Aug. 28 2
627 Apr. 24, Aug. 28 I
ts Aug. 2
628 July 16, Aug. 28 I2
Jun g. 28 Plant died.
789 May 7, Aug. 28 °
900 Apr. 26, Aug. 28
Qor Apr. 24, Aug. 28
902 Apr. 24, Aug. 28
903 Apr. 24, Aug. 28
904 Apr. 24, Aug. 28
905 Apr. 24, Aug. 28
A
to
COR HF OMHO
g. 28
910 May 2, Aug. 28
OIL May 2, Aug. 28 Ir
912 May 2, Aug. 28 Plant died.
013 May 2, Aug. 28 oO
DopDGE: STUDIES IN GENUS GYMNOSPORANGIUM 299
Sixteen of the twenty-four plants inoculated gave positive
results, showing seventy-nine separate infections. Six plants
were not infected, two died. The time required for the full de-
velopment of this species is only nine or ten months.
The endeavor to infect Chamaecyparis with G. fraternum and
G. biseptatum has not as yet resulted in success.
LITERATURE CITED
1, Arthur, J.C. Cultures of Uredineae in 1910. Mycologia 4: 7-33.
Ja 1912.
2. Dodge, B. O. Effect of the host on the morphology of certain
species of Gymnosporangium. Bull. Torrey Club 42: 519-542.
pl. 28, 29. N 1915.
—. Report on further cultures of Gymnosporangium. Paper
read at the December meeting of the Botanical Society of
America, New York, 1916.
Studies in the genus Gymnosporangium. I. Notes on
the distribution of the mycelium, buffer ceils, and the germina-
tion of the aecidiospore. Brooklyn Bot. Gard. Mem. 1: 128-140.
pl. 1 +f.1-6. 6 Je 1918.
5. Dodge, B. O., & Adams, J. F. Notes relating to the Gymno-
sporangia on Myrica and Comptonia. Mycologia g: 23-29.
N29 FF Ja tory.
6. Fromme, F. D. A new gymnosporangial connection. Mycologia
6: 226-230. S$ 1914.
7- Heald, F. D. The life history of the cedar rust fungus (Gymno-
sporangium Juniperi-virginianae Schw.). Ann. Rep. Nebraska
Agr. Exp. Sta. 22: 105-113. pl. I-13. 1909.
8. Kern, F. D. Gymnosporangium. N. Am. Fl. 7: 188-211. Ap
9. Plowright, C. B. British Uredineae and Ustilagineae. London.
10. Tubeuf, C. Mitteilungen ueber einiger Pflanzenkrankheiten.
Zeits. Pflanzenkrankheiten 3: 201-205. 1893.
11. Weimer, J. L. The origin and development of the galls produced
by two cedar rust fungi. Am. Jour. Bot. 4: 241-251. pl. 12-16
+f.2. My 1917.
. Three cedar rust fungi, their life histories and diseases
they produce. Cornell Agr. Exp. Sta. Bull. 390: 507-547. f.
236-157. My 1917.
300 DopDGE: STUDIES IN GENUS GYMNOSPORANGIUM
Explanation of plate 8
_ The photographs of spores were made from sori used in making the inoculations
reported in TABLE VI. Magnification, about 350.
GYMNOSPORANGIUM TRANSFORMANS
Fic. 1. A typical spore of this species showing a germ pore at the apex of each
cell. He cell wall is comparatively thick.
Fic. 2. A three-celled spore, the lower cell showing two germ pores near the
septum.
Fic. 3. A bcuseae! long PPO of the type that is difficult to distinguish
from such spores of G. fra in Fic. 7. The germ pore at the apex
of the terminal cell, ‘ase own in Fic. 3, appears to be a very Scasislidkd feature.
Spores of G. Pesce frequently germinate at the apex but the germ pore is not
eect mar
eS rae spores comparatively short. Both cells of three of them have
ae germinated.
1G. 5. A group of very thin-walled spores shaped very much like spores of G.
fraternum. The three-celled nai: Kae is easily oT by its shape from
the baie spores of G. bi in Fic.
6. Large, broad, dark SREES spores, none of ‘ohh has germinated.
GYMNOSPORANGIUM FRATERNUM
. 7. Spores from a dark brown sorus. The spore wall of the upper cell is
acs aes at the a
Fic. 8. Spores from a fn Se sorus. The pore at the apex
of the upper cell is visible in one of t
thi
“
The spore walls are very
GYMNOSPORANGIUM BISEPTATUM
F om ah £. +h
FIG. 9. 1 mentioned in another paper
(Dodge, 4). This i is the youngest infection I have been able to find in nature.
spores have from four to seven cells. There are very few three-celled spores.
Fic. 10. Spores from a sorus on a large burl about eight inches in length and
two inches in diameter. Nearly every spore is three-celled.
INDEX TO AMERICAN BOTANICAL LITERATURE
1916-1918
The aim of this Index is to include all current botanical literature written by
Americans, published in aaa or based upon American material ; the word Amer-
ica being used in the broadest se
Reviews, and papers that sales ag to sd agriculture, horticulture,
manufactured products of vegetable origin, or laboratory m gre not included, an
no attempt is made to index the literature of bacteri laey en occasional exception is
made in favor of some paper appearing in an American periodical which is devoted
wholly to botany. Reprints are not mentioned unless they differ from the original in
some important particular. If users of the Index will call the attention of the editor
to errors or omissions, their kindness will be appreciated.
This Index is reprinted monthly on cards, and furnished in this form to subscribers
at the rate of one cent for each card, Selections of cards are not permitted ; each
subscriber must take all cards published during the term of his subscription, Corr
spondence relating to the card issue should be addressed to the Treasurer of the rey
Botanical Club,
Andrews, A. L. Bryological notes—IV. A new hybrid in Physcomit-
rium. Torreya 18: 52-54. 10 Ap 1918.
Anthony, S. A. An anomaly of wheat anthers. Jour. Heredity 9:
166-168. f. 6,7. Ap 1918.
Ashe, W. W. Notes on Betula. Rhodora 20: 63, 64. 9 Ap 1918.
Bailey, C. H., & Curjar, A. M. Respiration of stored wheat. Jour.
Agr. Research 12: 685-713. f. 1-7. 18 Mr 1918
Berger, A. Agave fourcroydes. Curt. Bot. Mag. IV. 14: pl. 8746.
Mr 1918
A plant from Yucatan.
Bessey, E. A.. & Makemson, W. K. Notes on the control of rye smut
(Urocystis occulta) Ann. Rep. Michigan State Board Agr. 56: 305-
307. 1917. [Illust.]
Bidwell, G. L. A physical and chemical study of the kafir kernel.
U. S. Dept. Agr. Bull. 634: 1-6. f. 1. 4 Ap 1918.
Blake, S. F. Notes on the Clayton Herbarium. Rhodora 20: 21-28.
f. 1-5. 12 F 1918; 48-54. 2 Mr 1918; 65-73. f.6-8. 9 Ap 1918.
Blake, S. F. A variety of Smilax glauca. Rhodora 20: 78-80. 9 Ap
1918
Smilax glauca genuina.
301
302 INDEX TO AMERICAN BOTANICAL LITERATURE
Borzi, A. Studi sulle Mixoficee. Nuo. Gior. Bot. Ital. N. S. 24: 65—
112. p I9I7
Includes several American species of Stigonema.
Corrected title.
Boyce, J. S. Perennial mycelium of Gymnosporangium blasdaleanum.
Phytopathology 8: 161, 162. 30 Ap 1918.
Brown, W. H. The rate of growth of Podocarpus imbricatus at the
top of Mount Banahao, Luzon, Philippine Islands. Philip. Jour.
Sci. 12: (Bot.) 317-329. pl. 17 +f. 1, 2. N 1917.
Carpenter, C. W. Bean spot disease. Hawaii Agr. Exp. Sta. Ext.
Bull. 8: [1-4]ef. 7, 2. 14 Mr 1918.
Coe, H.S. Origin of the Georgia and Alabama varieties of velvet bean.
Jour. Am. Soc. Agron. 10: 175-179. f. 25, 26. Ap 1918.
Collins, G. N. Tropical varieties of maize. Jour. Heredity 9: 147-
154. f. I-3 + frontispiece. Ap 1918
Coons, G. H. Notes on Michigan plant diseases in 1916. Ann. Rep.
Michigan State Board Agr. 56. 310-317. 1917.
Coons, G. H.. Oat smut. Ann. Rep. Michigan State Board Agr. 56:
308, 309. 1917.
Coons, G. H. A Phoma disease of celery. Ann. Rep. Michigan State
Board Agr. 56: 318. 1917.
Coons, G. H. The relation of weather to epidemics of late blight of
potato. Ann. Rep. Michigan State Board Agr. 56: S17, 318. 2017:
Coons, G. H., & Nelson, R. The plant diseases of importance in the
transportation of fruits and vegetables. 1-60. i. 2°08. (Chicago.
F 1918.
Copeland, E. B. The genus Christiopteris. Philip. Jour. Sci. 12:
(Bot.) 331-336. N 1917.
Day, M. A. Dates of Eaton’s Ferns of North America. Rhodora 20:
74,75. 9 Ap 1918.
Elliott, J. A. Nematode injury to sweet potatoes. Phytopathology 8:
169. f. 1. 30 Ap 1918.
Emig, W. H. Travertine deposits of Oklahoma. Oklahoma Geol.
Surv. Bull. 29: 1-76. pl. 1-15 +f. 1-5. Au 1917.
Contains lists of plants found in these deposits.
Evans, A. W. Noteworthy Lejeuneae from Florida. Am. Jour. Bot.
5: 131-150. f. 1-5. 26 Ap 19018.
Cololejeunea contractiloba, Lejeunea cladogyna, Euosmolejeunea parvula and
Ptychocoleus heterophyllus, spp. nov., are described.
INDEX TO AMERICAN BOTANICAL LITERATURE 303
Faull, J. H. Fomes officinalis (Vill.) a timber destroying fungus.
Trans. Royal Canadian Inst. Toronto 11: 185-209. pl. 18-25.
1916.
Fernald, M. L. The North American Littorella. Rhodora 20: 61, 62.
9 Ap 1918.
Fernald, M.L. The validity of Oxalis americana. Rhodora 20: 76-78.
9 Ap 1918.
Gericke, W. F. Effects of rest and no-rest periods upon growth of
Solanum. Bot. Gaz. 65: 344-353. 15 Ap 1918.
Green, M. L. Echeveria setosa. Curt. Bot. Mag. IV. 14: pl. 8748.
Mr ro18.
A plant from Mexico.
Greenman, J. M. Monograph of the North and Central American
species of the genus Senecio—Part II. Ann. Missouri Bot. Gard. 5:
37-108. pl. 2, s. I
Includes Senecio molinarius sp. nov.
Hall, F. H. Poor ventilation injures stored potatoes. N. Y. Agr.
Exp. Sta. Bull. 436: 1-11. pl. r- O17
Halsted, B. D. Reciprocal breeding in tomatoes. Jour. Heredity 9:
169-173. Ap 1918.
Hansen, A. A. A national floral emblem. Science II. 47: 365-367.
12 Ap 1918.
Henry, J.K. A new variety of Rubus parviflorus. Torreya 18: 54, 55.
f. it. 40 Ap 1918.
Hermessen, J. L. A journey on the Rio Zamora, Equador. Geog,
Rev. 4: 434-449. f. 1-7. D 1917.
Contains some information of botanical interest.
Hicken, C. M. Una Aracea curiosa Felipponia. An. Soc. Cien. Argen-
tina 84: 240-244. I917.
Hodgson, R.W. Black smut of figs. Month. Bull. State Comm. Hort.
Calif. 7: 188, 189. f. 28. Ap 1918.
Holway, E. W. D. Infected grass seeds and subsequent rust develop-
ment. Phytopathology 8: 169. 30 Ap 1918.
Horsford, M. Dr. Cyrus Guernsey Pringle. The Vermonter 23: 12-
14. 1918.
Howe, M. A. The marine algae and marine spermatophytes of the
Tomas Barrera Expedition to Cuba. Smithsonian Misc. Col. 68":
1-13. 9 Ap 1918. [Illust.]
Includes Phormidium Hendersonti sp. nov.
304 INDEX TO AMERICAN BOTANICAL LITERATURE
Hutcheson, T. B., & Wolfe, T. K. The effect of hybridization on
maturity and yield in corn. Virginia Agr. Exp. Sta. Tech. Bull.
18: 161-170. Je 1917.
Jackson, H. S. Carduaceous species of Puccinia—lI. Species occur-
ring on the tribe Vernoniae. Bot. Gaz. 65: 289-312. 15 Ap 1918.
Includes Puccinia fraterna, P. hyalina, P. rata, P. idonea, P. notha, P. praealta,
P. discreta, P. inaequata and P. Kuntzii, spp. nov.
Jones, D. F. Bearing of heterosis upon double fertilization. Bot.
Gaz. 65: 324-333. f. 1-3. 15 Ap 1918.
Keith, G. W. Control of cherry leaf spot in Wisconsin. Wisconsin
Agr. Exp. Sta. Bull. 286: 1-11. f. 1-7. F 1918
Kline, W. A. A rare hybrid oak in Pennsylvania. Forest Leaves 16:
120, 121. Ap1rgr18._ [Illust.]
Langdon, L. M. The ray system of Quercus alba. Bot. Gaz. 65:
313-323. f- I-22. 15 Ap 1918.
Lipman, C. B. A new method of extracting the soil solution. Univ.
Calif. Publ. Agr. Sci. 3: 131-134. 15 Mr to18.
Lloyd, C.G. Letter No. 66. 1-16. Cincinnati. O 1917.
Lloyd, C. G. Mycological notes. 51: 717-732. f. 1075-1100 + por-
trait. N 1917; 52: 733-748. f. ITI0I-1123 + portrait. D 1917;
53: 749-764. f. 1124-1148 + portrait. Ja 1918.
Lloyd, C. G. The myths of mycology. 1-16. Exhibits A-E. Cin-
cinnati. D 1917.
art 1
Loeb, Il The law controlling the quantity and rate of repeNOrIRIDA:
Proc. Nat. Acad. Sci. 4: 117-121. 15 Ap 1918.
Love, H. H., & Craig, W. T. Methods used and results obtained in
cereal investigations at the Cornell station. Jour. Am. Soc. Agron.
10: 145-157. pl. 4+/f. 24. Ap 1918.
Long, E.R. Further results in desiccation and respiration of Echino-
cactus. Bot. Gaz. 65: 354-358. f. 1. 15 Ap 1918.
MacCaughey, V. The genus Gleichenia (Dicranopteris) in the Hawaiian
Islands. Torreya 18: 41-52. 10 Ap 1918.
McClelland, C.K. The time at which cotton uses the most moisture.
Jour. Am. Soc. Agron. 10: 185-1809. Ap 1918.
McCubbin, W. A. Dispersal distance of urediniospores of Cronartium
ribicola as indicated by their rate of fall in still air. Phytopathology
Bt 48 36. f. ¥... 24 Ja 1638:
INDEX TO AMERICAN BOTANICAL LITERATURE 305
McKee, R. Glandular pubescence in various Medicago species. Jour.
Am. Soc. Agron. 10: 159-162. Ap 1918
Merrill, E. D. Alabastra Borneensia. Jour. Straits Branch Roy.
Assoc. 77: 189-247. 1917
Forty-eight new species are described.
Merrill, E. D. An interpretation of Rumphius’s Herbarium Am-
boinense. 1-595. pl. z, 2. Manila. 1 N 1917.
Merrill, E. D. New Philippine Melastomataceae. Philip. Jour. Sci.
12: (Bot.) 337-360. N 1917.
Twenty-eight new species are described.
Merrill, J. H., & Melchers, L. E. Insects and slant diseases attacking
garden crops. Kansas Agr. Exp. Sta. Circ. 65: I-12. Ap 1918.
Mitchell, J. A. Incense cedar. U.S. Dept. Agr. Bull: 604: 1-40.
pl. 1-5 +f. 1-4. 16 Mr 1918.
Morse, A. P. List of the water-color drawings of fungi by George E.
Morris in the Peabody Museum of Salem. 1-70. Salem. 1918.
Murphy, P. A., & Wortley, E. J. Determination of the factors inducing
leaf roll of potatoes particularly in northern climates. (First
progress report.) Phytopathology 8: 150-154. f.z. 30 Ap 1918.
Osterhout, W. J. V. A demonstration of photosynthesis. Am. Jour.
Bot. 5: 105-111. f. 7, 2. 26 Ap 1918.
Osterhout, W. J. V., & Haas, A. R. C. Dynamical aspects of photo.
synthesis. Proc. Nat. Acad. Sci. 4: 85-91. 15 Ap 1918.
Osterhout, W. J. V., & Haas, A.R.C. A simple method of measuring
photosynthesis. Science II. 47: 420-422. 26 Ap 1918.
Parish, S. B. Notes on some southern California plants. Bot. Gaz.
64: 334-343. 15 Ap 1918.
Pease, V. A. North Pacific coast species of Desmarestia. Puget
Sound Marine Sta. Publ. 1: 383-394. pl. 83, 84. 28 D 1917.
Piper, C. V. Cutthroat grass (Panicum Combsii). Jour. Am. Soc.
Agron. 10: 162-164. Ap 1918.
Pittier, H. New or noteworthy plants from Colombia and Central
America—6. Contr. U. S. Nat. Herb. 18: 225-259 + i-x. pl. 106.
I5S1
ee he 26 new species in Cowssapoa (2), Cecropia (4), Roupala (3), Schinolobinm
(2), Terminalia (2), Miconia (1), Mimusops (1), Cordia (7), Adenocalymna (4), an
several new or imperfectly known species in other genera.
Rhoads, A. S: Some new or little known hosts for wood-destroying
fungi—II. Phytopathology 8: 164-167. 30 Ap 1918.
306 INDEX TO AMERICAN BOTANICAL LITERATURE
Rigg, G. B. Growth of trees in Sphagnum. Bot. Gaz. 65: 359-362
15 Ap 1918.
Rollins, W. Letters from an old to a young gardener. Horticulture
27: 247-249. f. I-6. 16 Mr 1918.
Includes notes on corn breeding.
Sando, C. E., & Bartlett, H. H. The flavones of Rhus. Am. Jour.
Bot. 5: 112-119. 26 Ap 1918.
Shafer, J. A. Narrative of a trip to South America for collecting cacti.
Jour. N. Y. Bot. Gard. 19: 21-43. F 1918.
Shear, C. L. Pathological aspects of the federal fruit and vegetable
inspection service. Phytopathology 8: 155-160. 30 Ap 1918.
Sinnott, E.W. Conservatism and variability in the seedling of Dicoty-
ledons. Am. Jour. Bot. 5: 120-130. f. r-4. 26 Ap 1918.
Skan, S. A. Petunia integrifolia. Curt. Bot. Mag. IV. 14: pl. 8749.
Mr 1918.
A South American plant.
Small, J. K. A winter collecting trip in Florida. Jour. N. Y. Bot.
Gard. 19: 69-77. pl. 210, 211. Ap 1918.
Small, J. K. Cactus hunting on the coast of South Carolina. Jour.
N. Y. Bot. Gard. 18: 237-246. pl. 207. N 1917.
Soth, B. H. Pasque flowers. Am. Bot. 24: 1, 2. F 1918. [Illust.]
Spessard, E. A. Prothallia of Lycopodium in America. Bot. Gaz. 65:
362. 15 Ap 1918. :
Squires, W. A. An interesting case of seasonal inversion. Am. Bot.
24:.6, 7.8 20%8.
Stakman, E. C., & Hoerner,G.R. The occurrence of Puccinia graminis
tritici-compactt in the southern United States. Phytopathology 8:
141-149. f. ¥, 2. 30 Ap 1918.
Steil, W. N. Studies of some new cases of apogamy in ferns. Bull.
Torrey Club 45: 93-108. pl. 4,5. 1 Ap 1918.
Stevens, F. L. Some meliolicolous parasites and commensals from
Porto Rico. Bot. Gaz. 65: 227-249. pl. 5,6 +f. 1-5. 15 Mrig18.
Stevens, H. E. Melanose—II. Florida Agr. Exp. Sta. Bull. 145.
103-116. f. 26-31. F 1918.
Stevens, O. A. Plants of Blue Rapids and Mahnattan, Kansas. Cor-
rections and Index to common names. Am. Mid. Nat. 5: 201-204.
Mr 1918.
Stevenson, J. A. Report of the department of pathology and botany.
Ann. Rep. Ins. Exp. Sta. Porto Rico 1917: 37-83. 1917.
INDEX TO AMERICAN BOTANICAL LITERATURE 307
Stevenson, J. & Rose, R. C. Vegetable diseases. Ann. Rep. Ins.
Exp. Sta. Porto Rico 1917: 83-98. 1917.
Includes Cercospora acrocomiae and Tubercularia coccicola, spp. nov.
Stewart, F. C., & Mix, A. J. Blackheart and the aeration of potatoes
in storage. New York Agr. Exp. Sta. Bull. 436: 321-362. pl. 1-10.
Je 1917.
Stout, A.B. Observations on tulips—II. Jour. Hort. Soc. New York
2: 235-243. pl. go-g2. F 1918.
Stout, A. B. Fertility in Cichorium intybus: self-compatibility and
self-incompatibility among the offspring of self-fertile lines of de-
scent. Jour. Genetics 7: 71-103. pl. 4-6. 9 F 1918
Tanaka, T. New Japanese fungi. Notes and translations—IV. My-
cologia 10: 86-92. Mr 1918.
Tenopyr, L. A. On the constancy of cell shape in leaves of varying
shape. Bull. Torrey Club 45: 51-76. f. r. 7 Mr 1918.
Thom, C., & Church, M. B. Aspergillus fumigatus, A. nidulans, A.
terreus n. sp. and their allies. Am. Jour. Bot. 5: 84-104. f. I-3.
9g Mr 1018.
Thomas, H. E. Cultures of Aecidium tubulosum and A. passifloriicola:
Phytopathology 8: 163, 164. 30 Ap 1918.
Valleau, W. D. Sterility in the strawberry.. Jour. Agr. Research 12:
613-670. pl. BE + 35, 36. 11 Mr 1918.
[Vries, H. de.] Hugo de Vries opera e periodicis collata. Vol. 1: 1-
630. Utrecht. 1918.
Waller, A. E. Crop centers of the United States. Jour. Am. Soc.
Agron. 10: 49-83. f. 5-12. F 1918.
Walton, L. B. Entetramorus globosus. A new genus and species of
algae belonging to the Protococcoidea (Family Coelastridae). Ohio
Jour. Sci. 18: 126-128. f. 1. F 1918.
Weatherwax, P. A remarkable case of fasciation in Oenothera biennis.
Proc. Indiana Acad. Sci. 1916: 363, 364. 1917. [Illust.]
Weatherwax, P. A variation in Plantago lanceolata. Proc. Indiana
Acad. Sci. 1916: 365-367. f. I, 2. 1917.
Weaver, J. R. The effect of certain rusts upon the transpiration of
their hosts. Minnesota Bot. Stud. 4: 379-406. pl. 41, 42 + f. I-9.
20 S 1916.
Weir, J. R., & Hubert, E. E. Notes on forest tree rusts. Phyto-
pathology 8: 114-118. Mr 1918.
308 INDEX TO AMERICAN BOTANICAL LITERATURE
Weston, W. H. The development of Thraustotheca, a peculiar water-
mould. Ann. Bot. 32: 155-173. pl. 4,5 +f. 1, 2. Ja 1918.
Whitaker, E.S. Anatomy of certain goldenrods. Bot. Gaz. 65: 250-
260. pl. 7,8 +f. 1. 15 Mr 1918.
White, O. E. Environment, variation and the laws of heredity.
Brooklyn Bot. Gard. Leaflets 6: [1-16] f. I). 17. Ap 1988:
Wilson, G. W. Rusts of Hamilton and Marion Counties, Indiana.
Proc. Indiana Acad. Sci. 1916: 382, 383. IQI7.
Wilson, G. W. Studies in North American Peronosporales—VII.
New and noteworthy species. Mycologia 10: 168, 169. My 1918.
Rhysotheca Acalyphae sp. nov. is described
Wilson, O. T. Notes upon a market disease of limes. Phytopath-
ology 8: 45-50. f. 1-5. 14 F 1918.
Wolf, F. A. Tobacco wildfire. Jour. Agr. Research 12: 449-457.
J. 4, 2. -¥B:F :igi8
Wolf, F. A. SPE with a note on mechanical injury as a
cause of their development. Jour. Agr. Research 13: 253-260.
pl. 18, 19 +f. 1. 22 Ap 1918.
Woodward, R. W. Some Connecticut plants. Rhodora 20: 97, 98
My 1018.
Yates, H. S. Some recently collected Philippine fungi. Philip. Jour.
Sci. 12: (Bot.) 361-380. N 1917.
ncludes 38 new species in Meliola (17), Asterina (6), Asterinella (1), Morenoella
(1), Nectria (1), Trabutia (1), Melanopsamma (1), Mycosphaerella (1), Stigmatea (1),
Merrilliopeltis (1), Pleospora (1), Hypoxylon (1), Nummularia (1), Xylaria (1),
Pirostoma (1), Phyllosticta (1), and Melanconium (t).
Zeller, S. M. Correlation of the strength and durability of southern
pine. Ann. Missouri Bot. Gard. gs: 109-118. pl. 7, 8+f.7
Ap 1918.
Zeller, S. M., & Dodge, C. W. Rhizopogon in North America. Ann.
Missouri Bot. Gard. §: 1-36. pl. 3... F188.
Nine new species are described.
Zeller, S. M., & Dodge, C. W. Gautieria in North America. Ann.
Missouri Bot. Gard. 5: po ah hae 9. Ap 1918.
Gautieria plumbea sp. nov. is describe:
Zimmerman, H. E. The siascak fe: Am. Forestry 24: 231: Ap
1918.
BULL. ToRREY CLUB VOLUME 45, PLATE -8
DODGE: STUDIES IN THE GENUS GYMNOSPORANGIUM
Vol. 45 No. 8
BULLETIN
OF THE
TORREY BOTANICAL CLUB
The evolution of maize
PauL WEATHERWAX
(WITH THIRTY-SIX TEXT FIGURES)
The ancestry of Indian corn, the place of its origin, and its
relation to other plants of its family have been the subjects of
extensive study and discussion. As a New World contribution
to science and agriculture, the maize plant found, after a time,
its present systematic position, and it has made its way into a
place of prime importance in the economic life of the world.
When first seen by the civilized explorers of this country, maize
was a cultivated plant; and no wild form of it has ever been found.
Theories maintaining that it was of Old World origin have been
shown to lack sufficient proof, and it is now very generally be-
lieved to have originated somewhere in the plateau region of
Mexico or Central America.
Concerning the ancestry of maize a number of different theories
have been advanced. In 1892 Bailey (1) expressed the opinion
that Zea canina, which had been described a short time previously,
was the progenitor of maize. Harshberger’s work (17), published
in 1893, was also based upon the assumption that this was the
primitive maize plant. The latter authority and others have since
shown that Zea canina is a hybrid between teosinte and ordinary
maize. In 1911 Harshberger (20, pp. 51-52) described, as the
theoretical ancestor, a plant whose inflorescence would resemble
that of Tripsacum; and in 1912 he suggested (21, p. 399) that
[The Bulletin for July (45: 259-308. pl. 8) was issued August 9, 1918.]
309
310 WEATHERWAX: THE EVOLUTION OF MAIZE
“our maize is of hybrid origin, probably starting as a sport of
teosinte, which then crossed itself with the normal ancestor,
producing our cultivated corn.’’ Characterizing these earlier
theories is Hackel’s suggestion, made (16, p. 20) as early as 1889,
that the female spike of corn was evolved by monstrous fasciation,
from a number of parts like the lateral branches of the male
inflorescence. In 1906 Montgomery (24) described a theoretical
ancestor of maize and pointed out the fact that the ear is not the
fasciated homologue of the lateral branches but the equivalent of
the central spike of the tassel. Mrs. Kellerman had given a
similar interpretation of the case in 1895, but her article (22)
seems not to have been available to many botanists. In 1912
Collins made the point (9, p. 525) that the homology between the
ear and the central spike of the tassel is inconclusive, because the
central spike of the tassel is as anomalous and as hard to explain
as is the ear. He discussed two theories as to the structure and
origin of these organs and favored the idea that they are the results
of fasciation. According to his theory, ordinary maize originated
as a hybrid between teosinte and some plant possessing the char-
acters of pod corn. In 1912 East (12) discussed the origin of
maize and said that he agreed, in a way, with both Collins and
Montgomery.
These are the more comprehensive discussions of the subject,
but the discovery of an abnormal ear or tassel, a new variety, or
an unusual habitat for the plant has, from time to time, been
enough to provoke a new theory or a modification of an old one.
The present status of the question may be summarized as
follows: most of the evidence concerning the evolution of maize
has come from a comparison of its gross morphology with that of
its near relatives, and from a study of hybrids between maize and
-teosinte; the ear of corn is generally supposed to be a fasciated
structure; two theories are now held as to the origin of the plant—
(1) that maize and teosinte have descended along slightly different
lines from a common ancestor, and (2) that maize originated ina
cross between teosinte and some closely related grass.
But in a detailed examination of maize and its near relatives,
we meet with many inconspicuous but significant organs which
have not thus far been considered in any theory of the evolution
WEATHERWAX: THE EVOLUTION OF MAIZE 311
of the plant. The writer has made a thorough study of many
varieties of corn, probably representing the full range of varia-
bility of the species, and other species of the Maydeae have been
examined in detail by way of comparison. As a result of these
studies, a theory is here proposed, which is believed to explain,
in a manner more consistent with known facts, the evolution of
maize.
Acknowledgment is here made of valuable assistance received
in this study from a number of sources. The investigation is an
outgrowth of a problem suggested to me five years ago by Pro-
fessor D. M. Mottier, who has since then also aided me with many
suggestions and criticisms. American consuls in several cities of
Central and South America have assisted me in getting seeds of
many varieties of corn; and very material assistance of the same
kind has been given me by a large number of friends in this
country. The Bureau of Plant Industry has supplied me with
seeds of corn and teosinte. Rhizomes of Tripsacum were obtained
from the Missouri Botanical Garden. The late Mr. Juan J.
Rodriguez, of Guatemala, sent me rhizomes of a species. of Trip-
sacum and seeds of a number of varieties of corn; he also told me
of many interesting observations that he had made on the grasses
of Guatemala. Mr. George F. Will, of Bismarck, North Dakota,
supplied me with seeds of thirty varieties of corn collected from
Indian reservations. Mr. A. Heinisch, of Clarcona, Florida,
sent me seeds and morphological material of teosinte, aided me in
making observations in his teosinte fields in the summer of 1917,
and gave me the benefit of many observations that he had made
as a grower of the plant.
CLASSIFICATION
The tribe Maydeae includes a number of grasses characterized
by unisexual spikelets, the male being above the female, in the
same or in different inflorescences. The seven genera of the tribe
fall naturally into two classes.. Zea, Euchlaena, and Tripsacum
bear evidences of close relationship and are all native of America.
Having the same general characteristics as these, but differing
from them somewhat in appearance and detailed structure, are
four genera, Coix,, Chionachne, Sclerachne, and Polytoca, which
312 WEATHERWAX: THE EVOLUTION OF MAIZE
are all native of southeastern Asia and the neighboring islands.
An anomalous American genus, Pariana, has been included in
this tribe by some authorities, but its natural position seems to be
in another group.
The best evidences concerning the immediate ancestry of
maize come from a comparative study of the three American
genera of the Maydeae. The Asiatic species are probably much
_ t. A variety of corn from an
servation; this plant may be con-
ad ihe type of the species, which is to-
day essentially the same as when first seen
by the white man.
less closely related to those of
America than they are to each
other; and, for present pur-
poses, they will not be con-
sidered in detail.
Zea and Euchlaena are suf-
ficiently closely related to
cross readily, and there is a
record of at least one case of *
hybridization of Euchlaena
and Tripsacum; but I have
found no record of any hybrid
between Zea and Tripsacum.
MorpHoLocy OF ZEA
The genus Zea, represented
by the single species, Zea
Mays L., comprises all the
varieties of maize. Being the
Maydeae, it is taken as the
basis for comparison. It isa
very variable species, but its
variability is mostly quanti-
tative in character. Good
general descriptions of the
plant have been available for
a long time, but some sig-
sifrant details have not thus far been included in any of these.
A complete working description makes necessary here the repe-
tition of some well-known facts as a proper background for new
details.
WEATHERWAX: THE EVOLUTION OF MAIZE
The aerial part of the
typical maize plant (Fic. 1)
consists of a tall, jointed stem,
provided with distichous
leaves, and bearing in the
axil of each leaf, except pos-
sibly some of the upper ones,
a potential branch. Some of
these branches do not nor-
mally develop, and microtome
sections are sometimes neces-
sary to disclose their rudi-
ments. One or more near
the middle of the stem develop
into ear-bearing shoots. The
lowest branches, arising at or
below the surface of the
Fic, 2. A typical male inflorescence of corn.
’
The main stem terminates in a
Male «nflorescence.—The
terminal panicle of spikelets
(FIG. 2) is made up of a num-
ber of simple or branched
lateral organs attached to a
central axis which terminates
in a spike. The spikelets are
borne in pairs, one pedicelled
and the other sessile (Fics. 3,
5). Occasionally in all, and
very regularly in a few varie-
ties, groups of three or more
are found (Fic. 4), instead of
pairs of spikelets, and in such
Fic. 3. Portion of .the central spike
of a corn tassel. Fic. 4. A group of
spikelets, such as often occurs itistead of
a pair. Fic. 5. Spikelets on the lateral
rachis of a corn tassel.
a group one is usually pedi-
celled. Both of a pair or all
of a larger group are occasion-
ally sessile. In’ the typical
314 WEATHERWAX: THE EVOLUTION OF MAIZE
case, where the spikelets are paired, the pedicelled ones are ar-
ranged symmetrically in a horizontal plane along the sides of the
lateral rachis, and the sessile ones are ventral to these on the
rachis, giving the structure a definite dorsoventral aspect (Fic.
5). The central spike is symmetrical with respect to the pairs
of spikelets, which are arranged in several longitudinal rows upon
it (Fic. 3).
The spikelets have been described in detail elsewhere (25, 26).
Each has two flowers and a full complement of glumes, palets,
and lodicules. Only the male essentials are functional, but each
flower has a rudimentary pistil. A superabundance of pollen is
produced, and it depends upon gravity and the wind for distribu-
tion. The flowering period of an individual plant is limited to
two or three weeks, depending upon weather conditions and the
size of the inflorescence.
Female inflorescence-—The ear-
bearing branch has the same gen-
eral structure as the main stalk,
except that its inflorescence is fe-
male; and the shortness of its axis
enables the leaf sheaths to cover
the inflorescence and mature fruit.
In some cases the laminae and
ligules of these prophylla are pres-
ent (Fic. 6), but often they are
each prophyllum is a bud which
may develop into an ear-bearing
shoot (Fic. 8). This behavior is
common to some varieties, and it
seems possible to bring it about in
any variety by subjecting the plant
Fic. 6. An ear of corn whose
teak idee Milnes: to proper physiological conditions.
The clusters of ears formed in this
way and occurring in ordinary maize (6, f. 96, 97) or in teosinte-
maize hybrids (17, pl. 1) are not at all to be confused with branched
ears of maize (Fic. 9). They have much the same appearance to
the casual observer, but they have a different origin and a different
WEATHERWAX: THE EVOLUTION OF MAIZE 315
Fic, 7. An ear-bearing shoot of a high-grade dent corn. The prophylla have
lost their laminae and ligules. Fic. 8. An ear branch bearing small lateral ears.
meaning. When these secondary branches develop far enough,
buds are formed in the axils of their prophylla, indicating the
possibility of tertiary branches.
The female spikelets are borne in pairs,
sessile or almost so, on a thickened axis, the
whole structure being the “ear.” The “cob”
is the axis of the mature ear after the grains
have been removed. Each row of pairs of
spikelets is responsible for two rows of grains
on the ear, thus accounting for the fact that an
ear always has an even number of rows. The
spikelets are two-flowered and have the same
parts as those of the male inflorescence; but the
whole lower flower of each is aborted, the lodi-
cules are present only in early stages of develop-
ment, and the stamens are aborted. The pistil wie ‘Aiea:
of the upper flower is usually the only functional aie ear suggest-
organ of either flower, but occasionally in any ing the relationships
bet al
plant, and as a fixed characteristic of some Ge sad tek Guisas
varieties, the pistil of the lower flower may also spike of the tassel
316 WEATHERWAX: THE EVOLUTION OF MAIZE
function. The pistil is characterized by the greatly elongated
“silk,’’ which reaches beyond the husks of the ear. I have given
in other papers (25, pp. 131-137, and 26, pp. 488-493) a detailed
description of the parts of the female spikelet.
As has already been noted, the normal ear is doubtless the
homologue of the central spike of the tassel. Every part of either
can be identified in the other, and they bear identical relations to
the other parts of the shoots which they terminate. The apparent
difference between the two is one of develop-
ment. Teratological forms (Fics. 9, 10) and
the inflorescences of suckers (Fics. 11-13) sug-
gest connecting links between the two organs.
Suckers—The sucker branches may arise
singly or two or more from one node (FIG. 1),
an appearance of the latter condition often
being the result of secondary branching. Most
of the suckers develop root systems more or less
independent of that of the main plant. Some
suckers have all the appearance of ears growing
low on the stalk; others grow tall and become
structurally and functionally the same as the
main stalks; and still others have characteristics
intermediate between these two extremes. These
intermediate types, while they have axillary
buds, usually develop only terminal inflores-
10. Atera- cences, which may represent any possible grada-
tological ear having tion between an ear and a normal tassel (Fics.
11-13). The most ear-like of these have, no
silat aailiees doubt, been made the subject of many of our
popular articles on freak ears. The figure given
by Mrs. Kellerman (22) and some of Montgomery’s photographs
(24) look suspiciously like sucker inflorescences. Many freak ears
given to me by non-technical collectors have turned out, on close
inquiry, to have been taken from suckers. Their value as evi-
dences of evolution are none the less for this; but they must be
interpreted in a different way, for they are not in any way terato-
logical.
‘
)
- Variations—Anomalous inflorescences are common, however,
WEATHERWAX: THE EVOLUTION OF MAIZE 317
and there are several more or less definite varieties of maize
Fics. 11-13. Normal terminal inflorescences of suckers, having characteristics
of both the ear and the tassel.
which do not agree with the general description just given. A
well-known variation is found in pod corn, each mature grain of
which is covered by the enlarged
glumes of the female spikelet.
Many plants of pod corn have
perfect or female flowers in their
tassels and produce fruits there,
and some of these plants have a
tendency to become earless; but
no independent earless variety
has ever been isolated. Other
varieties of maize occasionally
produce female flowers in the
tassel, probably as a result of
physiological conditions. It is
significant that silks produced in
the tassel are usually fasciated
like those of the normal female
flower, but bipartite silks have
been seen in a few cases. Most
Fic. 14. A tassel of Gernert’s Branch
Co
-
female spikes of maize have sterile tips, and these are often re-
318 . WEATHERWAX: THE EVOLUTION OF MAIZE
placed by staminate spikes (Fic. 10). Emerson (13, p. 83)
has isolated a dwarf variety which
has perfect flowers in the ear.
Gernert (15) has;isolated from a yel-
low dent variety. a strain in which
the ear is a loose panicle like that of
one of the sorghums. The tassel is
also branched more than in the typi-
cal variety (Fic. 14), and many
groups of spikelets are borne on pe-
duncles instead of being in sessile
groups (Fics. 15-19). A common
type of variation consists of a nor-
: mal ear bearing a whorl of small,
Fics. 15-19. Peduncled groups
of spikelets from Branch Corn, four-rowed branches around the base
showing the probable steps inthe (FIG. 9). Ears variously divided or
evolution of the pair of spikelets. — branched at the tip are also of com-
mon occurrence.
MorpHoLocy oF EuCHLAENA
Without discussing the systematic treatment of Euchlaena,
we may take Euchlaena mexicana Schrad. (synonymous with
Euchlaena luxurians Dur. and Reana luxurians Dur.) as the type
of the genus. It is the well-known teosinte, an annual forage plant
of the warmer parts of America. The species is much less variable
than maize.
Teosinte has much the same general appearance as maize,
especially since the casual observer’s mental picture of the latter
is obtained from seeing two or three plants growing inaclump. It
has the same sort of leafy stem, and the same potential branches
in the axils of its leaves; but the branches arising from the base
of the plant—ten to sixty or more in number—grow as tall as
the main stem and resemble it in every way. These continue to
branch indefinitely, a branch being possible at practically every
node; generally speaking, a branch has about as many nodes
as the main stalk above the point of branching. The main stalks
and some of the higher branches are terminated by male panicles.
At the top of any stem, the branches tend to be male, while the
lower ones of the same order tend to be female.
WEATHERWAX: THE EVOLUTION OF MAIZE 319
The plant makes a very ready response to conditions of soil
and climate, and herein lies the main cause of its variability.
/
Fic. 20. Tassel of teosinte Fic. 21. Portion of a spike of a teosinte tassel.
Rich, wet soil and warm weather are most favorable for luxuriant
growth. Plants are not injured by standing for some time in
water a foot deep. Under favorable conditions the stems often
Fic. 22. Portion of a corn tassel. The lateral spikes, LL, are jointed to the
in axis, which continues into the central spike, C. Fic. 23. Part of a tassel of
‘teosinte;. all the spikes are jointed to the main axis. :
320 WEATHERWAX: THE EVOLUTION OF MAIZE
fall prostrate, take root at the nodes, and send up branches,
becoming almost if not quite perennial at times.
Male inflorescence——The male panicle (Fic. 20) resembles
that of maize in appearance. The lateral spikes are so much
alike in the two that a close examination is necessary to reveal
any difference; but the central spike, which is characteristic of
maize, is lacking in teosinte. All the branches (Fic. 21) of the
teosinte tassel are dorsoventral, and it is probable that they are
all morphologically lateral, the true end of the axis being not
in any one rachis but between the bases of the uppermost two.
This is further supported by the fact that all the tassel branches,
including that (or those) apparently occupying a
central position, are jointed at their bases (Fic. 23),
which is not true of the central spike of the tassel of
maize (Fic. 22). This organ seems merely to have
failed to develop in teosinte. Following anthesis,
the rachids become brittle, easily separating at the
nodes.
The male spikelet is much like that of maize.
The same parts are present, including the rudi-
mentary pistils, and, in so far as has been observed,
there is no significant difference in the development
or the final form of the two. The pollen and pol-
lination are as in maize. :
Female inflorescence—The female inflorescence
is usually a single spike (Fic. 24), enclosed in the
sheath of a single leaf, which often has a ligule and
a short lamina (Fic. 25). Many of these shoots,
each borne on a short stem in the axil of a leaf,
may be enclosed in the sheath of a larger leaf of
the main stem, and the silks, protruding from the
sheath, give the whole the appearance of a young
ear of maize (Fic. 26). This whole structure is
the equivalent of an ear-bearing shoot of maize,
having branches of the third, fourth, or higher orders, but with
the characteristic female spike of teosinte, instead of an ear,
borne on each branch. ;
The female spike has received the most unsatisfactory treat-
Fic. 24. A
female spike of
teosinte.
WEATHERWAX: THE EVOLUTION OF MAIZE 321
ment of any part of the plant. As a typical instance of mis-
interpretation of its structure may be cited a statement in a
recent textbook (2, p. 137), where it is spoken of as being made
up of kernels fastened end to
end, the whole structure hav-
ing nothing to correspond to
the cob in maize. Collins’s
description: (9, p. 525) of the
spike as ‘one-rowed”’ is
equally misleading.
This spike is made up of ¢----
two rows of functional spike-
lets borne alternately in alve-
structure (F1GS. 27, 28) which
represents the other. spikelet
of the pair. Structurally,
then, the spike has four rows
25
of spikelets It will be re- Fic. 25. An “ear” of teosinte, covered
h : h | by its single husk, which has a rudimentary
membered that in the male lamina, L. Fic. 26. Portion of a stem of
spike the pedicelled spikelets teosinte bearing a cluster of female spikes,
are symmetrically arranged
on the rachis, with the sessile ones below them; but, in the
female spike, the fully developed sessile spikelets occupy the
symmetrical positions, and the rudimentary pedicelled ones have
been crowded up to a dorsal position (Fics. 34, 35). In
teosinte somewhat mixed with maize the rudimentary spikelet
is sometimes replaced with a sterile or male, pedicelled one, and a
little further contamination with maize produces two functional
female spikelets side by side. Paired female spikelets are also
occasionally found in pure teosinte. These facts indicate that
the female spike is the homologue of one of the branches of the
tassel, and it probably does not morphologically terminate the
shoot of which it is a part.
The female spikelet consists of the same parts as that in maize. _
The lower flower is aborted, rudimentary stamens are present,
aon WEATHERWAX: THE EVOLUTION OF MAIZE
and the development of no part shows any significant difference
except for the indurated outer glume. The silk has much the
same structure as in maize, but it is shorter.
Because of the large number of inflorescences, the flowering
period of a teosinte plant is much longer than that of a maize
plant, and the chances for self-pollination are better. When the
cae §
Fic m of a longitudinal section of an internode and its spikelet in
the pits erat of Euchlaena or Tripsacum. = saving I, internode; GG, glumes;
LL, lemmae; PP, paleae; S, rudimentary s' F, rudimentary flower; ste
functional pistil; Sp, position of the ties ae which does not appea
Fi
in a median section. G, 28. The rudimentary spikelet shown in the racine
rectangle in Fic, 27.
seeds are mature, the female spike separates at the nodes, each
internode bearing a spikelet and its enclosed fruit.
Mixed inflorescences.—The homologies suggested between the
male and female inflorescences are further indicated by the occur-
rence of mixed inflorescences. In male tassels, functional female
spikelets have several times been observed near the base of one
or more of the branches; and the sterile tip of the female spike,
as in maize, is often replaced by a short staminate portion (FIG. 29).
Some single spikes are so much like those of Tripsacum, to be
described later, that it would be hard to determine whether it is
a case of male spikelets being in a female inflorescence, or vice
versa, In such a mixed spike, each male spikelet is usually
WEATHERWAX: THE EVOLUTION OF MAIZE 323
accompanied by another male or sterile one; and each female
spikelet is characteristically alone, has an indurated outer glume,
and is sunken into the rachis. No functionally bisexual spikelets
have been seen, but it must be emphasized
that all are structurally so.
MorpPHOLOGY OF TRIPSACUM
A number of species of Tripsacum are
native of various parts of America, but the
best known is Tripsacum dactyloides L., the
gama grass of swamps and stream banks of
the central and southern parts of the United
States. Being the only species available for
my work, it is the basis of my conclusions
for the genus.
Tripsacum dactyloides (FIG. 30) is peren-
nial by means of a branched rhizome, from
the nodes of which arise the aerial shoots.
Many of these are sterile, and their stems
are so short that only the leaves appear above
the ground. In early summer, however, some
of the shoots elongate for flower bearing and
show the adult structure. These stems attain
the height of four to six feet and resemble
somewhat those of maize or teosinte. The
flowering period continues for two or three
months. The main stem bears a branch at each node, and this
has secondary and even tertiary branches. The main stem is
terminated by a panicle, and each branch by a spike or a pan-
icle. Erect or curving until after flowering, the stems are, by
means of some growth-mechanism near the base, caused to lie
prostrate by the time the seeds are mature.
Inflorescence—The male spikelets occupy the terminal and
the female the basal part of the same spike (Fic. 31). The mode
of branching of the panicle is similar to that in the male inflores-
cence of Euchlaena, There is no evidence of any central spike,
all the branches being much alike.
There is a very evident tendency for the highest inflorescence,
Fic. 29. A mixed spike
of teosinte.
324. WEATHERWAX: THE EVOLTUION OF MAIZE
that is, the one terminating the main stem, to be male, and the
lower ones, terminating the branches, to be female. Counts of
male and female spikelets in a number of inflorescences showed in
those of the main stem a ratio
of 1 female to 18.5 male; in those
of the primary branches the ratio
was I to 7.5; and on secondary
branches the ratio was. still
higher. No unisexual inflores-
cence has been observed, how-
ever, on any plant.
Male spikelets.\— The male
spikelets are arranged on the
rachis in two rows of pairs; and
the male portion of the spike,
even where there is but one spike
in the inflorescence, has the same
dorsoventral appearance as in
maize or teosinte (FIG. 32); one
Fic. 30. Habit of Tripsacum dactyloides. spikelet of a pair sometimes has
usually sessile.
a short pedicel, but both are
The rachis of me male part of the spike becomes
brittle soon after the pollen is shed.
Fic, 31.
Terminal inflorescence of Tripsacum dactyloides.
The outer glume of the male spikelet is somewhat thickened
and hardened, but otherwise the bracts, lodicules, stamens, and
rudimentary pistils are essentially the same as those of maize or
teosinte.
WEATHERWAX: THE EVOLUTION OF MAIZE 325
Female spikelets—The female portion of the spike (Fic. 32)
is similar to that of teosinte, but there is usually more or less
external evidence of dorsoventrality. The
spikelets are alternately imbedded in alveoli in
the rachis, forming two rows arranged almost
symmetrically or along one side of the rachis.
But the real four-rowed structure is clearly
indicated, as in teosinte, by the presence of
a rudimentary spikelet (Fics. 27, 28) near the
base of each functional one. This rudiment
is sometimes replaced by a functional spikelet,
which may be either male or female (Fic.
33). As in teosinte, the female portion of
the spike easily separates at the nodes when
the seeds are mature.
The female spikelet has the same parts as
that of teosinte, and they are arranged in the
same way. The stigma is bipartite (Fics. 32,
33), in contrast with that of Zea or Euchlaena.
Seeds—The plants that I have grown
ordinarily set seeds in only a few flowers, and
these do not germinate well. Mr. Rodriguez
has informed me that Tripsacum laxum, a solitary female spikelets.
Central American species, has never been FS 33- Pair of female
known to produce seeds. The morphological messin hokek
basis of this peculiarity has not been deter-
mined, but it is doubtless correlated with the perennial habit.
SUMMARY OF MORPHOLOGY
As is evident from the foregoing descriptions, these genera are
identical in structural plan; for every significant organ in any one
of them, there is a homologue, fully developed, rudimentary, or
indicated, in each of the others.
Common to all three is the jointed vegetative stem, bearing a
leaf or a leaf rudiment at each node and a branch or a bud in the
axil of almost every leaf. Buds, representing undeveloped shoots,
bear no indication of giving rise to branches, because their nodes
have ordinarily not developed far enough; but practically every
fully developed node has a bud.
326 WEATHERWAX: THE EVOLUTION OF MAIZE
Some of the stems of Tvipsacum live underground, thus making
the plant perennial, a condition which, in some of the species at
least, is associated with the swamp habitat and limited seed
production. When Euchlaena grows in a swampy place, it has a
tendency to assume the perennial habit by means of prostrate
34. Diagram of a cross section of one internode of a male spike of maize or
teosinte, showing the rachis, R, with one sessile, S, and one pedicelled, P, spikelet,
IG. 35. Section of an internode of a female spike of Tripsacum or Euchlaena.
showing a functional sessile, S, and a rudimentary pedicelled, P, spikelet imbedded
in the rachis,
stems rooting at the nodes; its seed production, however, seems
in no way impaired by this habit. Maize produces a large number
of viable seeds and has no tendency to become perennial.
The general inflorescence of the group is a panicle with a
central spike and lateral branches, like the tassel in maize. In
most of the inflorescences some parts of this general structure are
lacking; in the maize ear it is the lateral branches, and in the
teosinte tassel and the inflorescence of Tripsacum it is the central
spike; in the female spike of Euchlaena all that is present is a
single lateral branch. The spikelets occur usually in pairs, but
sometimes in larger groups, in all the inflorescences.
The spikelet has two glumes and two florets. Structurally
the flowers are as in the typical grass, but the regular suppression
of some of the essentials makes all the flowers functionally uni-
sexual. Tripsacum has two feathery stigmas, but the homologue
of these in Zea and Euchlaena is the silk, a fasciated organ divided
at the tip.
THEORIES OF THE EVOLUTION OF MAIZE
As has been noted by way of introduction, several theories |
have been advanced to explain the origin of the maize plant or
some of its parts; and it is appropriate here to examine in a critical
way some of the more prominent of these.
WEATHERWAX: THE EVOLUTION OF MAIZE 327
Origin of the ear.—Most of the attempts to explain the struc-
ture and origin of the ear of corn have been based upon Hackel’s
idea (16, p. 20) that it has resulted from the union of several
organs like the lateral branches of the tassel. Harshberger at
first supported this theory (17, pp. 75-83); after describing the
ear and some of its teratological variations, he says: ‘‘These
structural and teratological arrangements point to the probable
union of several spikes into a thick, fleshy axis, with grains on the
circumference, each paired row limited at the sides by a long,
shallow furrow, a row corresponding to a single spike of Euchlaena
or Tripsacum.”’ Gernert (14, p. 37) agrees with Harshberger in
support of the fasciation theory and gives some additional evi-
dences, most of which, however, are irrelevant or obviously open
to objection. Mrs. Kellerman (22) and Montgomery (24) object
to this theory and point out that the ear is the homologue of the
central spike of the tassel; and later (20, p. 51) Harshberger
apparently agrees with the former. Collins (9, p. 525) removes
all doubt of this homology and shows that the central spike of
the tassel is as anomalous and as much in need of explanation as
is the ear. He thinks it probable that both have resulted from
fasciation. One of the latest discussions of the structure of the
ear is that given by Worsdell (28, p. 58). Due to a special defini-
tion of terms, he does not call this a case of fasciation; but, as
the following statement indicates, he is in accord with most of
the others in his interpretation of the structure and origin of the
ear: “It consists of the fusion of numerous spikes with flattened
rachis, each bearing two rows of female spikelets, to form the thick
female inflorescence usually termed the ‘cob.’” (The word
“cob” is a misnomer; he is evidently talking about an ear.) A
new term, “disruption,” is proposed to cover the abnormality,
which ‘‘consists in the appearance of a ‘cob’ as a copiously-
branched paniculate inflorescence, closely resembling, in its ex-
treme form, the male inflorescence; and is due to the dissolution
of the compound organ into its separate parts.”
In spite of the apparent consistency of this widely accepted
theory of the formation of the ear, it is untenable at least in its
present form. There can be-no reasonable doubt of the homology
between the ear and the central spike of the tassel. Ears termin-
328 WEATHERWAX: THE EVOLUTION OF MAIZE
ated by male spikes (Fic. 10), ears with small female or mixed
branches attached (Fic. 9), and perfect ears replacing the central
spikes of sucker tassels (F1G. 12) all give indication of this. More-
over, the early development of the two organs is the same except
that in the case of the normal ear no side branches are developed.
But the idea that either or both of these organs is fasciated is
inconsistent with some important morphological facts. Carefully
prepared microtome sections fail to show any evidence of fascia-
tion, the development in either case being essentially the same as
that of any ordinary floral or vegetative shoot. The only kind of
organ of which we have any knowledge, the like of which could
have united to form an ear, is the lateral branch of the tassel, and
this involves a mathematical difficulty. The rows of grains on an
ear are paired because the spikelets are borne in pairs. Ifa lateral
spike of the tassel should contribute to the formation of an ear
anything in terms of paired spikelets, it would be responsible for
four rows, and this would make impossible an ear of ten, fourteen,
or eighteen rows; but the fact remains that such ears are common.
Harshberger’s idea that each pair of rows of grains corresponds
to a spike of Euchlaena or Tripsacum is of no avail in getting around
this difficulty; they bear two rows of functional spikelets, but not
at all because of the presence of paired spikelets, for one spikelet
of each pair is rudimentary. It may be noted, too, that although
Worsdell has the ear formed by the union of spikes bearing two
rows of spikelets, he has it “disrupt” into a central spike bearing
eight rows and a number of lateral spikes each bearing four rows
(28, pl. 30, f. 13). Incidentally it may be said that the ‘‘dis-
rupted”’ ear figured is not abnormal at all but a normal tassel of
a sucker.
Collins (9, p. 526) advances another theory which probably
offers the best explanation of the origin of both of the organs in
question. He suggests that the male inflorescence might have
been developed from a loose panicle by the shortening of some of
the branches until pairs of spikelets were left on a central spike
and a few lateral branches; the ear could easily have been de-
veloped from this by the loss of the side branches. He abandons
this theory for want of sufficient evidence in the form of inter-
mediate steps, but Gernert’s Branch Corn (Fics. 14-19) supplies
the needed evidence and strengthens a very consistent theory.
WEATHERWAX: THE EVOLUTION OF MAIZE 329
Hybrid origin of maize.—A new variety of corn known as Zea
canina was reported from Mexico about 1890, and Bailey (1)
and Harshberger (17) concluded that it was the long-sought wild
ancestor of the species, the latter making it the basis of his mono-
graph on corn. But correspondence with Mexicans who knew
the plant showed that Zea canina could be produced at will by
hybridizing teosinte and ordinary maize. Accordingly, previous
conclusions had to be revised, and along with Harshberger’s
revision (18) came the suggestion of three possible explanations
of the botanical nature of maize: (1) that maize is a distinct
species; (2) that maize originated as a hybrid between teosinte and
some unknown grass; and (3) that maize is the result of a cross
between teosinte and some variety of teosinte which had been
changed by cultivation. He favors the last-named possibility,
and in his latest expression on the subject (21, pp. 398-399)
this is the only theory given.
His conclusions were reached as a result of hybridization experi-
ments with maize and teosinte, in which a graded series of inter-
mediates between the ear of the one and the female spike of the
other were produced. A series of this Kind always suggests
evolution, but this one does not possess the advantage of having
maize at one end and its two hypothetical ancestors at the other.
Moreover, the status of the hybridization problem at the time
at which this theory was proposed (1896) and the evolutionary
influences attributed to cultivation, especially to irrigation, do
not show up so well alongside the genetics of this later day.
Collins’s theory (9) is similar to the second possibility suggested
by Harshberger; that is, he thinks that maize originated in a
cross between teosinte and some unknown grass similar to pod
corn and belonging to the Andropogoneae. With respect to a
large number of characteristics, ordinary maize is shown to be
intermediate between the primitive pod corn and the highly
specialized teosinte; and this is considered evidence that Zea is a
hybrid between the two extremes.
Although this theory is the most elaborate and the most
widely accepted of all the attempts that have been made to
explain the origin of maize, yet it falls short of its aim in some
respects. Granting the accuracy of the observations upon which
330 WEATHERWAX: THE EVOLUTION OF MAIZE
‘it is based, we may still question the value of the facts as con-
structive evidence leading to the conclusion. Is it safe to assume
on this basis alone that an intermediate of this kind is necessarily
a hybrid between the two extremes? And is a hybrid usually
intermediate in character between its parents? But many of the
supposed facts upon which this theory is based will not stand the
light of a close examination; and the actual conditions are capable
of a more simple and more direct explanation in another way.
The theory depends upon some hypothetical pure variety of
pod corn, which can be approached in reality only by inbreeding
some one of the many available genetic complexes commonly
known as pod corn. But inbreeding for purity implies intelligent
selection among the pure lines reached or approached; and this
places a burden of responsibility upon him who makes such selec-
tion the basis of a theory of this kind. Collins has in mind, how-
ever, some strain of pod corn the like of which he considers one of
‘the ancestors of maize; and, in describing its simple, primitive
nature, he makes some statements which are not in accord with
the morphological facts that I have already set forth. The ex-
treme type of pod corn is described (g9, p. 527) as having no ear,
the absence of branches being a primitive characteristic. Dis-
‘regarding the difficulty occasioned by the fact that these earless
plants are incapable of self-propagation, we may cite the fact
that they have buds in the axils of their leaves, indicating the
suppression of ears. If a plant that never had any ears is primi-
tive, one that has vestiges of ears that it has lost must be highly
specialized. He says (p. 528): “In the branched forms of pod
corn staminate flowers have never been observed on any of the
branches.”” Contrary to this is the fact that sucker branches of
pod corn often bear mixed or staminate inflorescences, these having
often been mistaken, no doubt, for earless plants, especially when
more than one plant is grown in a hill; and in several cases ears
of pod corn have been found bearing well-developed stamens.
Applying the specialization test to the ears, he says (p. 528):
“In pod corn branches have never been observed in the axils
of prophylla;” but I have often seen such branches, especially
when the main ear had been injured and buds are to be found in the
axils of the husks of all kinds of corn. From the foregoing cita-
WEATHERWAX: THE EVOLUTION OF MAIZE 331
tions it may be seen that Collins does not necessarily hold to any
one variety of pod corn for his evidence, switching from earless to
branched types as he finds structures to fit his hypothesis; and,
even with this latitude of choice, he fails to find sufficient evidence
that pod corn is wholly primitive. It is readily granted by any-
one that the tunicate character and the tendency toward her-
maphroditism are primitive characters, but otherwise the plant
is modern, as the fasciated silk, the husks of the ear, and the many
degenerate organs go to show. The earless plants are the most
highly specialized of all, as indicated by the vestigial ear-buds in
the axils of their leaves.
Similar inconsistencies occur in the description of teosinte,
which is held to be highly specialized; the mention of a single
example will be sufficient. In support of a detailed argument
that the extreme differentiation between the male and female
inflorescences of teosinte points toward specialization higher than
that in maize, he says (p. 524) that female flowers have never
been observed in the male inflorescence of Euchlaena. I have
already described mixed inflorescences of this plant; but the im-
portance of this point is diminished by the fact that in this entire
group of plants we are sone with flowers which are structurally
bisexual.
Readily granting that monoecism is a less fixed characteristic
and that the separation of male and female flowers is not so sharp
in maize as in teosinte, yet I do not believe that pod corn has been
shown to be sufficiently different from ordinary corn to merit
the prominent position that it holds in this theory.
In a recent report on this theory (11) the intolerance of self-
pollination in maize is given as another evidence of its hybrid
‘nature, it being almost if not quite unique among the grasses in
this respect. But possibly another explanation for this may be
found in its having a very small number of monoecious inflores-
cences and, because of protandry or protogyny, a limited chance
for self-pollination under normal conditions, in which respect it
is also unique among the grasses. All the other members of the
Maydeae, teosinte, for example, produce a large number of in-
florescences and have a flowering period much longer than that
‘of maize, the chances for self-pollination being correspondingly
gas WEATHERWAX: THE EVOLUTION OF MAIZE
better. Tripsacum can be excluded from this consideration be-
cause it depends mostly upon vegetative propagation for increase
in number of individuals. In those grasses which have perfect
flowers self-pollination is the rule. Tolerance of self-pollination
may thus be interpreted as a matter of adjustment to the condi-
tions which have prevailed during the development of the plant
in question.
Traumatic evolution.—Maize was one of the principal plants
considered by Blaringhem (3, 4, 5) in arriving at his theory of
mutation by traumatic influences, and his conclusion as to its
evolution may best be stated in his own words (6, p. 228):
“L’étude des variations observées A la suite de mutilations permet
de reconstituer l’évolution du genre Zea, et d’établir que l’ancétre
sauvage du Mais cultivé est l’espéce Euchlaena mexicana. Le
genre Zea est une forme monstreuse du genre Euchlaena née et
propagée par les soins de l’homme.”’_ The chief interest in his work
in this connection lies not so much in his contribution to our
knowledge of the evolution of the plant as in the peculiar methods
employed and the peculiar interpretation of results.
By mutilating plants in various ways he causes them to send
up sprouts whose terminal inflorescences bear both male and
female flowers. Seeds from these sprouts provide the starting
points for numerous new varieties characterized by such peculiari-
ties as suckers with mixed inflorescences, ears with perfect flowers,
branched ears, tubular leaf sheaths, etc. Many of these breed
true and are given the rank of elementary species, a number of
new varietal names being contributed to the already overburdened
list associated with the variations of this monotypic genus.
Throughout the discussion, the sucker bearing a female or mixed
inflorescence is considered an abnormality; but if this is the
correct interpretation for the suckers of the whole genus, it con-
stitutes a case where the exception is quite as common as the rule.
In his description of the spikelets of the normal plant (6, p. 21)
he states that neither the male nor the female flower contains
organs of the opposite sex; and this fallacy becomes the basis
for his assumption (5, p. 1253) that the acquisition of perfect
flowers by some of his new varieties constitutes a progressive step.
‘No change that he has produced in the plant involves any charac-
WEATHERWAX: THE EVOLUTION OF MAIZE Ba]
teristic that is actually new, and, as Gernert suggests (14, p. 6),
most of the results may be explained on a basis in no way con-
nected with the original mutilation. As the theory stands, we
can justly ask that the experiments be repeated successfully with
more than one variety of maize, and the results interpreted on the
basis of a more sound understanding of the morphology of the
plant throughout its normal range of variation, before we consider
this a substantiated explanation of the past history and present
tendencies of the plant.
Montgomery’s theory —Along with his explanation of the ear
of corn as the homologue of the central spike of the tassel (24),
Montgomery proposes a theory of evolution, which, while not
worked out in full detail, is probably the most consistent solution
of the problem yet offered. He considers maize a distinct genus
codrdinate with teosinte, both having developed from a common
teosinte-like ancestor.
A THEORY BASED UPON MORFHOLOGY
Exact knowledge of the past conditions being out of the ques-
tion, the ancestry of the maize plant and the steps taken in its
evolution can only be built up in theory from a study of the
modern plant and its near relatives; and some attempts previously
made in this direction have already been discussed. Any theory
that is to receive serious consideration must be based upon, and
consistent with, recognized facts; and, of two or more explanations
of the same thing, the simplest and most direct is to be preferred.
Several kinds of evidence are available for use in the study of
maize; but morphology, the basic constructive factor of most of
our theories of evolution, has never been utilized to the full extent.
To be sure, the gross structure of this group of plants has several
times been considered, but a morphological study that does not
go to the bottom of the question is likely to mislead the investi-
gator. The study of abnormal structures is always instructive,
but it is more significant to find in the normal plant a morpho-
logical basis for its abnormalities. Other considerations, such as
may be occasioned by experiments in hybridization and selection
for a few generations, constitute instructive checks on other
methods; but they must not be over-emphasized as constructive
334 WEATHERWAX: THE EVOLUTION OF MAIZE
material, and they are never any better than the morphology upon
which they are based.
Ancestry of maize.—It is neither necessary nor desirable, I
believe, to look for the ancestor of maize and its near relatives
in any plant now living; its evolution along different lines to
the present forms would imply its own probable disappearance.
There are found in different members of the Andropogoneae,
however, all the characteristics necessary for a theoretical pro-
genitor of maize and its American relatives. In the evolution of
these plants from their common ancestor, many steps were prob-
ably taken, no suggestion of which has passed down to modern
times. It is almost certain that the Asiatic species of the Maydeae
arose from this same stock, and many lines of descent may have led
to other forms now long extinct. Geology and archeology are of
little value to us in solving these problems, since the oldest remains
of these plants found in the rocks or in human habitations are
practically modern.
Theoretically the ancestor of these plants was an herbaceous
Maydeae and some of the Andropogoneae as well, have been
evolved by the suppression of parts, whose rudiments are, in most
instances, still to be found in the modern plants.
The cause of this suppression and the mechanism of its accom-
plishment are unsolved problems, but the phenomenon is not
peculiar to this group of plantsalone. It is known that monoecism
among the angiosperms has been reached independently in many
groups as a result of the suppression of one set of organs in each
WEATHERWAX: THE EVOLUTION OF MAIZE 335
flower; and the suppression of whole inflorescences and of vegeta-
tive parts is also common. Whether this suppression is due to
environment, or to inherent tendencies, or to a selection by the
environment among the results of inherent tendencies, is unknown;
but the comparative structure of Zea, Euchlaena, and Tripsacum
indicates that, in their development from a common ancestor,
certain tendencies have found expression in different ways in the
different genera; and this expression has often come in the nature
of a response to external conditions. In other words, we assume
that these three genera are closely related, not because the char-
acteristics which they have were present in a common ancestor,
but because they have been evolved from an ancestor which had
tendencies to suppress pistils in some flowers and stamens in
others, whole flowers in some cases, and in some cases vegetative
organs. Unless this assumption is made, we cannot base any
close relationship between Euchlaena and Tripsacum upon monoe-
cism, for the separation of the sexes in these two genera has been
accomplished:in different ways.
The first division of the progeny of this ancestral form set off
a group in which the upper spikelet of each pair had lost its pistils,
this giving rise to a number of genera of the Andropogoneae. On
the other hand, in some of the plants whose inflorescences had
retained their perfect flowers, the peduncles bearing groups of
spikelets grew shorter (Fics. 15-19) until the spikelets were ar-
ranged as in the tassels of maize or teosinte (FIGS. 3, 4, 5, 21).
Pairs or larger groups of spikelets could have been left in this way.
The plants in which the central spike of the inflorescence survived
this process became the ancestors of Zea; while those in which
this spike was lost gave rise to Euchlaena and Tripsacum. In the
separation of these two genera, two lines of evolution were fol-
lowed, depending upon the manner in which monoecism came
about.
Origin of Euchlaena.—In the line tending toward Euchlaena, the
uppermost inflorescences—those terminating the main branches
—lost their female elements and assumed the form of the present
teosinte tassel (Fic. 20), while the lower inflorescences lost their
male elements; between these two extremes were formed some
mixed spikes, the upper portions being staminate and the lower
336 WEATHERWAX: THE EVOLUTION OF MAIZE
pistillate. In the pistillate spikes, or portions of spikes, the
pedicelled spikelet and the lower flower of the sessile one became
aborted (Fics. 27, 28). Each branch bearing a female inflores-
cence shortened enough that the inflorescence was covered by the
sheath of the leaf just below it; at the same time, all but one (or
rarely more) of the spikes of this inflorescence disappeared, and the
two stigmas of each pistil, lying side by side and elongating to
keep their tips exposed beyond the enveloping leaf sheath, united,
except at their exposed tips, to form the silk. The brittle rachis
came with the hardening of the outer glume of the spikelet and
the walls of the alveolus in which the spikelet was imbedded.
The leaf whose sheath enclosed the female spike has since then
almost lost its lamina (Fic. 25). By a continued shortening of
the internodes, the sheaths of lower and larger leaves were later
caused to envelop clusters of these simple female spikes and their
husks (Fic. 26). There is nothing to indicate when Euchlaena
became an annual, but this habit i is probably a acerag with a
reduced supply of moisture.
Origin of Tripsacum.—In those individuals a toward
Tripsacum, monoecism came about as a result of the loss of female
elements from the upper part and male elements from the lower
part of each spike of the inflorescence (Fic. 32). The point of
‘division between the male and female portions of a spike was
higher in the inflorescences of the branches than in that of the
main stem. In the male portion, the pedicellate spikelet became
almost sessile; the female portion became essentially as in teosinte.
There was no appreciable shortening of any internodes, and the
inflorescence was not covered by a leaf sheath; consequently
there was no reduction in the number of its branches, and the
two stigmas have persisted (Fic. 31). This plant usually grows
in rich, wet soil, and it has remained perennial; its ability to
produce viable seeds is restricted.
Origin of Zea.—In the evolution of Zea from its progenitor,
the inflorescence terminating the main stem lost its female ele-
ments and approached its present form (Fic. 2). Those terminat-
ing some of the primary branches near the middle of the main
stem lost their male elements, and these branches began to shorten
their internodes. As this shortening continued, the secondary
WEATHERWAX: THE EVOLUTION OF MAIZE 337
branches were reduced to buds; and, as the leaf sheaths began
to enclose the terminal inflorescence of the branch, this inflores-
cence lost its lateral branches, only the central spike remaining.
Up to this time the mature fruits were covered by the palets
and glumes, but, as a response to the covering of husks, these
bracts became shorter, leaving the mature fruits naked except
for the husks of the ear. The leaves whose sheaths formed the
husks, tended to lose their laminae and ligules (Fic. 7); these
remain, however, in some varieties (Fic. 6). The two stigmas
united, as in teosinte, to form the silk. At some time in the
development of the ear, the lower flower of each female spikelet
lost its function, but when and why this occurred is not clear at
present. Probably all the primary branches of the stem, except
those low enough to take root, went through this process, and
most of them became still further reduced; at present, one to
five or six usually remain as ears, and the rest have been reduced
to buds in the axils of the leaves of the main stem. Those
primary branches low enough to take root (Fic. 1) did so and
have met with the varied fate characteristic of the suckers of
the plant. Some are like the main stalk and others like ears,
depending probably upon their relative independence; but the
great majority share the characteristics of both (Fics. 11-13).
Meaning of variations in maize——In the light of this the-
oretical history of the species we are able to interpret many of the
ordinary variations of maize. The podded ear is a reversion to
the condition of the ear unprotected by husks, and very probably
not a primitive form that has come down to us unchanged.
Grains in the tassel mark a resumed function of the pistils in the
male flowers, and they are usually characterized by the fasciated
silk, showing that their flowers have not reverted in all char-
acteristics. Gernert’s Branch Corn (15) is a reversion to the
primitive paniculate inflorescence by a plant which is modern in
other respects. Emerson’s dwarf variety (13) and one of Blaring-
hem’s new varieties (5), both of which have perfect flowers in the
ear, are, in so far as the perfect flowers are concerned, analogous
to the varieties with perfect flowers in the tassel. Two-flowered
female spikelets, such as those regularly found in Country
Gentleman sweet corn (25, p. 135) and a few other varieties, are
338 WEATHERWAX: THE EVOLUTION OF MAIZE
reversions to the primitive two-flowered condition. We should
not expect any one plant to show reversion in all respects; char-
acteristics probably behave as units in reversion as much as in
heredity.
Some freaks, such as ears divided at the tip (17, p. 81; 14, p.
37) and fasciated kernels (27), are probably best interpreted as
anomalies of ontogeny, giving no more clue to the past than is
afforded by Siamese twins or lilies with two-parted flowers.
Indeed, an ancestral form that would be consistent with all the
teratological ears that have been used as evidence would be an
impossibility. It is a fact worth mentioning, also, that many of
the teratological conditions that do not fit into the foregoing
theory as reversions are not inherited, while those in accord with
this theory often form the starting points of new varieties tending
to breed true.
It is not deemed advisable here to attempt to trace the prob-
able origin of all the more or less fixed varieties of corn that have
come into existence after the generic characteristics ‘were reached;
but no constant characteristic is known in any variety, which is
inconsistent with the general theory here outlined. The develop-
ment of the annual habit was similar to that of teosinte, and no
further explanation is offered. The restricted period of flowering
has a morphological cause in the reduced number of inflorescences.
CONCLUSION
It is granted that the foregoing theory involves a measure of
speculation, but that is to be expected of any theory; it is as con-
servative as any that have been advanced to explain the origin
of maize, and much more so than some of them. It is believed
to be a logical deduction from the best and most recent evidences
available, and in it are embodied parts of some of its predecessors.
No point in it is known to be out of harmony with a rational
interpretation of any established fact of structure, history, or
genetic behavior concerning the plants with which it deals.
How much of this evolution of maize was due to natural
agencies, and how much to the influence of primitive agriculture,
we have no means of knowing; but, from the botanical point of
view, the changes wrought by four hundred years of civilization
WEATHERWAX: THE EVOLUTION OF MAIZE —- 839
Fic. 36. A corn plant very highly specialized by scientific breedin,
suckers aes all been lost, and the entire energy of the plant is directed el the
production of a single large ear.
340. WEATHERWAX: THE EVOLUTION OF MAIZE
are insignificant as compared with those preceding the advent
of the white man. The varieties of corn which Columbus first saw
in the West Indies in the fifteenth century were probably in no
essential way different from those now grown on many Indian
reservations (Fic. 1); and the highest attainment of corn breeding,
as represented by the dent varieties of the Mississippi valley
today (Fics. 7, 36), is merely one of a combination of these
varieties, with a few more organs dropped, a little more concen-
trated fructification, and a little greater vegetative vigor.
SUMMARY
Vestigial organs being considered, Zea, Euchlaena, and Trip-
sacum are identical in structural plan. The present aspect of
each is due to the suppression of some parts which were present
in a primitive ancestor having perfect flowers borne in one type
of inflorescence.
The ear of maize is the homologue of the central spike of the
tassel. There is no morphological evidence to support the view
that either of these organs originated in the fusion of more simple
parts; and there is in no one of the genera here considered any
organ the like of which could have united to form either the ear
or the central spike of the tassel.
The prevailing theory that maize is a species of hybrid origin
has little to suggest it when maize and its near relatives are
thoroughly understood, and it is not in harmony with the most
significant facts of morphology. It seems much more probable
that Zea, Euchlaena, and Tripsacum have descended independently
from a common ancestral form now extinct.
INDIANA UNIVERSITY.
REFERENCES
1. Bailey, L. H. A new maize and its behavior under cultivation.
Cornell Agr. Exp. Sta. Bull. 49: 332-338. 1892.
2. Bailey, L. H., & Gilbert, A. W. Plant Breeding. New York.
1916.
3. Blaringhem, L. Anomalies héréditaires provoquées par des trau-
matismes. Compt. Rend. 140: 378-380. 1905.
Production d’une espéce élémentaire nouvelle de mais par
traumatismes. Compt. Rend. 143: 245-247. 1906.
4.
coos]
WEATHERWAX: THE EVOLUTION OF MAIZE 341
. Blaringhem, L. Production par traumatisme et fixation d’une
variété nouvelle de mais, le Zea Mays var. pseudo-androgyna.
Compt. Rend. 143: 1252-1254. 1906.
Mutation et traumatismes. Paris. 1908.
. Candolle, A. de. Origin of cultivated plants. New York. 1885.
Collins, G. N. A new type of Indian corn from China, U. S.
Dept. Agr. Bur. Pl. Ind. Bull. 161. 1909.
The origin of maize. Jour. Washington Acad. Sci. 2: 520-
530.
Hybrids of Zea tunicata and Zea ramosa. Proc. Nat. Acad.
Sciences 3: 345-349. 1917. Also Jour. Agr. Research 9: 383-
396. pl. 13-21. 1917
—— Maize: Its origin and relationships. Jour. Washington
Acad. Sci. 8: 42-43. 1918.
. East, E. M. A chronicle of the tribe of corn. Pop. Sci. Mo. 82:
S35-296. J. 2-12. | 1913.
. Emerson, R. A. Genetic correlation_and spurious allelomorphism
in maize. Nebraska Agr. Exp. Sta. Rept. 1911: 59-90
Gernert, W. B. Analysis of characters in corn and their behavior
in transmission. Champaign, Illinois. 1912.
A new sub-species of Zea Mays. Am. Nat. 46: 616-622. f.
a—c. I912.
. Hackel, E. Gramineae. Engler & Prantl, Nat. Pflanzenfam. 2?:
I-97. 1889.
. Harshberger, J. W. Maize: a botanical and economic study.
Contr. Bot. Lab. Univ. Pennsylvania 1: 75-202. pl. 14-17. 1893.
Fertile crosses of teosinte and maize. Gard. & For. 9: 522-
523. 1896
. — Astudy of the fertile hybrids produced by crossing teosinté
and maize. Contr. Bot. Lab. Univ. Pennsylvania 2: 231-235.
1gol.
. — An unusual form of maize. Proc. Delaware Co. Inst. Sci.
6: 49-53. I9QII.
. — Maize, or Indian corn. Cycl. Amer. Agri. Ed. ed. 4. 2:
398-402. 1012.
. Kellerman, Mrs. W. A. The primitive corn. Meehan’s Monthly
5: 44. 1895.
. Kempton, J. H. Floral abnormalities in maize. U.S. Dept. Agri.
Bur, Pl. ind. . Bull. 278... Fors.
. Montgomery, E. G. What is an ear of corn? Pop. Sci. Mo. 68:
55-62. f. I-14. 1906.
342 WEATHERWAX: THE EVOLUTION OF MAIZE
25. Weatherwax, P. Morphology of the flowers of Zea Mays. Bull.
Torrey Club 43: 127-144. pl. 5, 6. +f. 1-4. 1916.
The development of the spikelets of Zea Mays. Bull. Tor-
rey Club 44: 483-496. pl. 23 +f. 1-33. 10917.
27. Wolfe, T.K. Fasciation in maize kernels. Am. Nat. 50: 306-309.
f. I-33 1916.
28. Worsdell, W. C. The principles of plant teratology. 2. London.
1916.
26.
The relationship between the number of sporophylls and the
numbers of stamens and pistils—a criticism
HELENE M. Boas
In a paper entitled, ‘The interrelationship of the number of
stamens and pistils in the flowers of Ficaria,’’ Harris* has at-
tempted, by statistical methods, to throw some light on the
biological factors which determine the sex of an organism. The
relative numbers of pistils and stamens present are assumed to
indicate which sex is more influented as the total number of
sporophylls increases. Without a knowledge of the number of
functional spores that are produced the ratio of the two kinds of
sporophylls can hardly be considered as a fundamental measure
of the sex of a flower. Nevertheless the quantitative relations
between stamens and pistils, if handled so as to be of biological
and not purely statistical significance, may suggest more precisely
the factors that influence the development of the two sexes.
Harris correlates the deviations of the total number of sporo-
phylls and those of the pistils and stamens from their “probable
values.”” He means by this the average frequency of pistils and
stamens to be expected according to the total number of sporo-
phylls. He does this rather than correlate directly with the
numbers of pistils and stamens in order to eliminate, as he
believes, the spurious correlation which would exist in the latter
case. He finds the correlation between the number of sporophylls
and the deviation of the pistils from their “probable number’ to
be of equal magnitude to that of the stamens, but positive,
while that of the stamens is negative. From this he concludes that
as the number of sporophylls increases, the pistils increase rela-
tively more rapidly than the stamens.
It can be shown that this result necessarily follows from the
fact that there are more stamens than pistils while their varia-
* Biol. Bull. 34: 7-17. 1918.
343
344 3 Boas: NUMBERS OF
bilities are almost equal, and consequently has only mathematical
and not biological significance.
According to Harris’s formula, if (n + s) represents the mean
number of sporophylls with their deviations and, let us say, p the
percentage of pistils and z, their deviations from their ‘probable
number” the pistils would be represented by the formula
(n + s)p + zp, and the stamens by the formula (m + s)(I — p) — 2s
where z, is the deviation of the stamens from their “probable
number” and zg, = 2». The sum of the pistils and stamens must,
of course, equal (7 + 5), the total number of sporophylls. He then
correlates the variables s and gz, assuming that z is not contained
in s.
We may analyze Harris’s formula for pistils to determine which
values are known and which are unknown. If » represents the
average total number of sporophylls, 7, that of pistils, 7, that of
stamens then» =”,+,. m,+ x are the pistils with their devia-
tions, », + y the stamens with their deviations. The deviation of
the total number of sporophylls from their. mean, s, equals x + 9;
so that Harris’s value for sporophylls (zn +s) =”, +n,+x+y.
Since p represents the percentage of pistils, it is equal to p/(m»+Ms),
and Harris’s value for pistils, given above, becomes
in +a + ¥)——— + &
ak
Z» is the only unknown value. We may find its value from the
following equation:
\gusesese”
Pag 2p = =p -_ x,
the number of pistils plus their deviation. Solving this
= = ek = Moy
. Na+ n,
S=x+y.
Therefore, correlating s, the total number of sporophylls, with 2,,
the deviation of the pistils from their “probable value,”
: i ae see NX — Nyy te
SPOROPHYLLS AND STAMENS AND PISTILS 345
and averaging
ah N32 raed tyty oe (ns, ep Ny) ¥Ox0y
Ny + Me :
[Zps]
According to Harris’s result [z,s] is always positive. The
formula shows that this must be the case when
NOx” apy tt? + (ns aes Ny) O20y >o
or when
n,(o22 + forty) > Np(e,? + roze,).
If o, and gy are equal this can happen only when 7, > mp,
or in other words when the stamens are more numerous than
the pistils.
From the original data from which Harris has drawn the
material for this study (see Harris for references) it is evident
that the standard deviations for pistils and stamens are nearly
equal and that the stamens are more numerous than the pistils.
For the average values for Europe given by Pearson the mean
number of pistils is 19.432 with a standard deviation of + 4.8508;
the mean number for stamens 26.498 with a standard deviation
of + 4.2562. The coefficient of correlation is + 0.5584. Sub-
stituting these values we find the results obtained by Harris
necessarily follow from the existing numerical relations.
From this consideration it seems evident that the results
obtained by Harris do not add anything to the observations on the
numbers of pistils and stamens and their variabilities. Just so
the higher coefficients of variation given by Harris in Table I of
his paper result from the fact that the same value (standard
deviation) is divided in the case of the pistils by a lower value
(mean for pistils) than in the case of the stamens (mean for
stamens). :
It might be said that the total number of sporophylls and the
per cent of stamens and pistils vary independently, in which
case the expression (7 + s)(p + v) would represent the number
of pistils. Disregarding the value sv, since it is small in com-
parison to the other values, the expression becomes p(n + s) + nv
and the same relations will hold as in the case discussed above.
New YorK BOTANICAL GARDEN
INDEX TO AMERICAN BOTANICAL LITERATURE
1918
The aim of this Index is to include all current botanical literature written by
cacti published in America, or based upon American material ; the word Amer-
ca bei eing u used in the broadest sen
Reviews, and papers that sis sere to forestry, agricalture, cate.
manufactured products of vegetable origin, or laboratory methods are not included, an
no attempt is made to index the literature of bacteriology. An seabed: soa is
made in favor of some paper appearing in an American periodical which is devoted
wholly to botany. eprints are not mentioned unless they differ from the original in
some important particular. If users of the Index will call the attention of the editor
to errors or omissions, their kindness will be appreciated.
This Index is reprinted monthly on cards, and furnished in this form to subscribers
at the rate of one cent for each card, Selections of cards are not permitted ; each
spondence Ny to the card issue should be addressed to the Treasurer of the Torrey
Botanical Cl
Appleman, C, O. Respiration and catalase activity in sweet corn.
Am. Jour. Bot. 5: 207-209. f. z. 16 My 1918
Arthur, J. C. New species of Uredineae—X. Bull. Torrey Club 45:
141-156. 1 My 1918.
New species are described in Uromyces (2), Puccinia (8), Aecidium (10), and
Uredo (3).
Arthur, J.C. Uredinales of Costa Rica a on wana i by E. W. D.
cess Mycologia 10: 111-154. My
species are described in Ravenelia (1), on dat ae Puccinia (13), Aecidium
(3), ia Uredo (2). :
Arthur, J. C. Uredinales of the Andes, based on collections by Dr.
and Mrs. Rose. iat, ote 65: iia 15 My1
ew species in me ie (1), Puccinia
(4), and Aecidium (1).
Ashe, W. W. Notes on southern woody plants. Torreya 17: 71-74.
Vaccinium Margarettae sp. nov. is described.
Atkinson, G. F. Some new species of Jnocybe. Am. Jour. Bot. 5:
210-218. 16 My 1918.
Twenty-four new species are described.
Blake, S.F. Note on the proper name for the sassafras. Rhodora 20:
98, 99. I My 1918.
347
348 INDEX TO AMERICAN BOTANICAL LITERATURE.
Britton, E.G. “The catkin-hypnum, with long hoses.” Bryologist
at: 32.. 40 Ap t9ts.
Britton, E. G. Mosses from Florida collected by Severin Rapp.
Bryologist 21: 27, 28. 30 Ap 1918.
Britton, N. L. John Adolph Shafer. Jour. N. Y. Bot. Gard. 19: 97-
99. My 1918.
Britton, N.L. Ared pine Are eat Jour. N. Y. Bot. Gard. 19: 105-
106.. My 1918.
Brandes, E. W. Anthracnose of lettuce caused by Marssonina panat-
toniana. Jour. Agr. Research 13: 261-280. pl. C, 20 +f. 1-4:
29 Ap 1918.
Brotherton, W. Jr., & Bartlett, H. H. Cell measurement as an aid in
the analysis of quantitative variation. Am. Jour. Bot. 5: 192-206.
f. 1,2. 16 My 1918.
Buchanan, R. E. Bacterial phylogeny as indicated by modern types.
Am. Nat. §2: 233-246. f. 1-3. F 1918.
Burnham, S. H. Lichens of the Berkshire Hills, Massachusetts.
Bryologist 21: 29-32. 30 Ap 1918.
Burt, E. A. Corticiums causing Pellicularia disease of the coffee plant,
Hypochnose of pomaceous fruits, and Rhizoctonia disease. Ann.
Missouri Bot. Gard. 5: 119-132. f. 1-3.. 1 Ap 1918.
Coville, F. V., & Blake,S.F. Notes on District of Columbia juncseee:
Proc. Biol. Soc. Washington 31: 45, 46. 16 My 1918.
Cowgill, H.B. Studies in inheritance in sugar cane. Jour. Dept. Agr.
Porto Rico 2: 33-41. Ja 1918.
Ducke, A. As especies de Massaranduba (genero Mimusops L.)
descriptas pelo botanico brasileiro Francisco Freire Allemao. Arch.
Jard. Bot. Rio de Janeiro 2: 11-16. pl. 2A.
Includes Mimusops huberi sp. nov
Duggar, B. M., & Bonns, W. W. The effect of Bordeaux mixture on
the rate of transpiration. Ann. Missouri Bot. Gard. 5: 153-174.
Ap 1918.
Emig, W. H. Mosses as rock builders. Bryologist 21: 25-27. pl. 15,
I6. 30 Ap 1918.
Enlows, E. M. A. A leafblight of Kalmia latifolia. Jour. Agr. Re-
search 13: 190-212. pl. 14-17 + f. 1, 2. 15 Ap 1918.
Fairman, C.E. Notes on new species of fungi from various localities—
II. Mycologia 10: 164-167. My 1918.
Includes new species in Phoma (1), Phomopsis (1), Sphaeropsis (2), Camaro-
sporium (1), Rhabdospora (1), Microdiplodia (1), Hendersonia (1), Dictyochora (1),
Platystomum (1).
INDEX TO AMERICAN BOTANICAL LITERATURE 349
Fernald, M. L. Rosa blanda ees its allies of northern Maine and ad-
jacent Canada. Rhodor : 90-96. 1 My 10918.
Rosa johannensis and _R. se ant spp. nov., are described.
Freeman, G. F. Producing bread making wheats for warm climates.
Jour. Heredity 9: 211-226. f. 12-16. 25 My 1018.
Gates, R.R. A systematic analytical study of certain North American
Convallariaceae, considered in regard to their origin through dis-
continuous variation. Ann. Bot. 32: 253-257. Ap 1918.
Gleason, H. A. The local distribution of introduced species near
Douglas Lake, Michigan. Torreya 18: 81-89. 4 Je 1918.
Hall, H. M. Walnut pollen as a cause of hay fever. Science II. 47:
516, 517. 24 My 1918.
Hansen, A. A. Preserving our wild flowers. Torreya 18: 65-69.
8 My 1918.
Harper, R. M. Changes in the forest area of New England in three
centuries. Jour. Forestry 16: 442-452. f. 7. Ap 1918.
House, H. D. Report of the state atone: 1916. N. Y. State Mus.
Bull. 197: 1-122. f. 1-12. 1 My
Includes new species’in Cercospora (1), Rae (1), Cryptospora (1), Crypto-
sels (1), Dendrodochium (1), Diplodia (2), Eutypella (1), Gloeosporium (1), Lepto-
Sphaeria (1), caincnaonoinents (1), Microdiplodia (1), Phoma (1), Ramularia (1), Septoria
(3); Spaleee is (3).
Humbert, E. P. A striking variation in Silene noctiflora. Bull. Torrey
Club 45: 157, 158. f. Z, 2. 1 My 1918.
Humbert, J. G. Tomato diseases in Ohio. Ohio Agr. Exp. Sta. Bull.
321: 159-196. f. r-r2. F. 1918... [Ilust.]
Humphrey, H. B., & Potter, A.A. Cereal smuts and the disinfection of
~ seed grain. U. S. Dept. Agr. Farm. Bull. 939: 1-28. f. 1-16.
Ap 1918.
Husmann, G. C. Girdling the Corinth grape to make it bear. Jour.
Heredity 9: 201-210. f. s-rz. 25 My ro18.
Ishikawa, M. Studies on the embryo sac and fertilization in Oenothera.
Ann. Bot. 32: 279-317. pl. 7 +f. 1-14. Ap 1918.
_ Jensen, G. H. Studies on morphology of wheat. Washington Agr.
Exp. Sta: Bull. 150: 1-21. pl. 1-5. Mr 1918.
Jensen, L. P. A plea for the preservation of our native plants. Gard.
_ Chrom. Am. 22: 147-149. My 10918.
Jones, F.R. Yellow-leafblotch of alfalfa caused by the fungus Pyreno-
peziza medicaginis. Jour. Agr. Research 13: 307-330. . DD, 25,
(_<o 6 My 1018.
350 INDEX TO AMERICAN BOTANICAL LITERATURE
Lee, H. N., & Hovey, R. W. The principal properties, structure and
identification of Canadian pulpwoods. Pulp & Paper Mag. 16:
419-422. f. I-6. 9 My 1918.
Lehman, 8. G. Conidial formation in Sphaeronema fimbriatum. My-
cologia 10: 155-163. pl. 7. My 1918.
Levy, D. J. Astation for Ephemerum near New York City. Bryolo-
gist 21: 33. 30 Ap 1918.
Lipman, C. B., & Gericke, W. F. Copper and zinc as antagonistic
agents to the ‘alkali’ salts in soils. “Am. Jour. Bot. 5: 151-170,
f. 1, 2. 16 My 1918.
Léfgren, A. Novas contribuicoes para as Cactaceas brasileiras sobre
os generos Zygocactus a Schlumbergera. Arch. Jard. Bot. Rio de
Janeiro 2: 19-32. pl. 3-6. 1918.
Zygocactus candidus sp. nov. is described.
Lofgren, A. Novas contribuigoes para o genero Rhipsalis. Arch.
Jard. Bot. Rio de Janeiro 2: 35-45. pl. 7-17. 1918.
Six new species are described.
Lofgren, A. Novas subsidios para a flora Orchidacea do Brasil. Arch.
Jard. Bot. Rio de Janeiro 2: 49-62. pl. 18-26. 1918.
Includes new species in Pleurothallus (8), Epidendrum (1), Leptotes (1), and
Moaxillaria (1).
Long, W. H. An undescribed canker of poplars and willows caused by
Cytospora chrysosperma. Jour. Agr. Research 13: 331-345. pl.
27, 28. 6 My 1918.
Long, W. H., & Harsch, R. M. Aecial stage of Puccinia oxalidis.
Bot. Gaz. 65: 475-478. 15 My 1918
Ludwig, C. A. The effect of tobacco smoke and of methyl] iodide vapor
on the growth of certain microérganisms. Am. Jour. Bot. §i-171~
177- 16 My 1918.
Maxon, W. R. A new hybrid Asplenium. Am. Fern. Jour. 8: 1-3
Mr 1918
Asplenium Gravesii hybr. nov.
McAtee, W. L. Note on the plants of Wallop’s Island, Virginia.
Torreya 18: 70, 71. 8 My 1918.
McColl, W. R. The male fern at Owen Sound, Ontario. Am. Fern
Jour. 8: 18,19. Mrig18.
Millspaugh, C. F., & Sherff, E.E. New species of Xanthium and Soli-
dago. Field Mus. Nat. Hist. Publ. 199: 1-7. pl. 1-6. 22 Ap 1918.
Six new species are described.
Mosher, E. , The grasses of Illinois. Illinois Agr. Exp. Sta. Bull. 205;
261-425. f. I-287. Mr 1918.
INDEX TO AMERICAN BOTANICAL LITERATURE 351
Murrill, W. A. [lustrations of fungi—XXVIII. Mycologia 10: 107-
110. pl. 6.
Pycnoporus cashed: Poronidulus conchifer, Polyporus Polyporus, Bjerkan-
dera adusta, Tyromyces amorphus and Cerrena unicolor, are illustrated in color.
Nichols, G. E. Additions to the list of bryophytes from Cape Breton.
Bryologist 21: 28, 29.. 30 Ap 1918.
Osner, G. A. Siemphylium leafspot of cucumbers. Jour. Agr. Re-
search 13: 295-306. pl. 21+24 + f. I-3. 29 Ap 1918.
Osterhout, G.E. A new Hymenopappus from Colorado. ‘Torreya 18:
go. 4 Je 1918.
Hymenopappus polycephalus sp. nov.
Parr, R. The response of Pilobolus to light. Ann. Bot. 32: 177-205.
f. 1-4. Ap 1918.
pin se E. B. Notes on certain Cruciferae. Ann. Missouri Bot.
ome -151. Ap 1918
Includes alee Osterhoutii, Dithyrea membranacea and D, clinata spp. nov.
Porto, P. C. Um caso de hybridacgdio natural. Arch. Jard. Bot. Rio
de Janeiro 2: 65, 66. pl. 27. 1918.
Pratt, O. A. Soil fungi in relation to diseases of the Irish potato in
southern Idaho. Jour. Agr. Research 13: 73-100. pl. A., B. +f.
I-q. 8 Ap 1918.
Prince, S. F. Fern notes. Am. Fern Jour. 8: 4-8. Mr 1918.
Rangel, E. Alguns fungos novas do Brasil. Arch. Jard. Bot. Rio de
Janeiro 2: 69-71. pl. 28-30.
New species are described in Puccinia (3), Septoria (1), Cercospora (1), and
Helmintosporium (1).
Ransier, H. E. More pleasures from old fields. Am. Fern Jour. 8:
8-12. pl. r, 2. Mr 1918.
Record, S. J. Intercellular canals in dicotyledonous woods. Jour.
Forestry 16: 429-441. f. 1-8. Ap 1918.
Roberts, J. W., & Pierce, L. Apple bitter rot and its control. U.S.
Dept. Agr. Farm. Bull. 938: 1-14. f. 1-3. Ap 1918.
Rock, J. F. New species of Hawaiian plants. Bull. Torrey Club 45:
133-139. pl. 6. 1 My 1918.
Includes new species in Cyanea (2), Lobelia (1), and Straussia (1).
Sargent, C. S. Notes on North American trees—I. Quercus. Bot.
Gaz. 65: 423-459. 15 My 1918.
One new species and many new varieties and new hybrids are described.
Seaver, F. J. The black locust tree and its insect enemies. Jour.
Y. Bot. Gard. 19: 100-104. pl. 212. My 1918.
352 INDEX TO AMERICAN BOTANICAL LITERATURE
Sherbakoff,C.D. Tomatodiseases. Florida Agr. Exp. Sta. Bull. 146:
119-132. f. 32-44. Mr 1918.
Shive, J. W., & Martin, W. H. A comparison of salt requirements for
young and for mature buckwheat plants in water cultures and sand
cultures. Am. Jour. Bot. 5: 186-191. 16 My 10918.
Shufeldt, R.W. The much-despised skunk cabbage—earliest of spring
flowers. Am. Forestry 24: 225-231. f. 3-12. Ap 1918.
Silveira, A. A. da. Contribuicio para as Eriocaulaceas brasileiras.
Arch. Jard. Bot. Rio de Janeiro 2: 7, 8. pl. 1, 2. 1918.
Includes descriptions of Paepalanthus densifolius and P. capanemae, spp. nov.-
Sinnott, E.W. Evidence from insular floras as to the method of evolu-
tion. Am. Nat. 52: 269-272. F 1918.
Small, J. K. Ferns of tropical Florida. i-ix + 1-80. pl. 1-5. New
York. 1918. [Illust.]
Pycnodoria pinetorum sp. nov. is described.
Smith, G. M. The vertical distribution of Volvox in the Plankton of
Lake Monona. Am. Jour. Bot. 5: 178-185. 16 My 10918.
Stevens, F. L., & Anderson, H.W. Protect the wheat crop. Eradicate
the common barberry from Illinois. Illinois Agr. Exp. Sta. Ext.
Cire. 22: 1~4. f. 1-3. Ap 1918.
Stewart, F. C. The velvet-stemmed Collybia—a wild winter mush-
room. N. Y. Agr. Exp. Sta. Bull. 448: 79-98. pl. 1-10. F 1918.
{Illust.]
Stevenson, J. A. The green muscardine fungus in Porto Rico. Jour.
Dept. Agr. Porto Rico 2: 19-32. pl. r. Ja 1918.
Sumstine, D. R. Fungi of Chautauqua County, N. Y. N. Y. State
Mus. Bull. 197: 111-118. 1 My 1918.
Truog, E. Soil acidity—I. Its relation to the growth of plants. Soil
Sci. 5: 169-195. Mr 1918.
Turner, W. F. Nezara viridula and kernel spot of pecan. Science II.
47: 490, 491. 17 My 1918.
Vinall, H. N., & Reed, H.R. Effect of temperature and other meteoro-
logical factors on the growth of sorghums. Jour. Agr. Research 13:
133-148. pl. 12, 12. 8 Ap 1918.
Vries, H.de. Mass mutations and twin hybrids of Oenothera grandiflora
Ait. Bot. Gaz. 65: 377-422. f. 1-6. 15 My 1918.
Vries, H. de. Mass mutation in Zea Mays. Science II. 47: 465-467.
10 My 1918.
Weir, J. R. Effects of mistletoe on young conifers. Jour. Agr. Re-
search 12: 715-718. pl. 37. 18 Mr 1918.
AN ILLUSTRATED FLORA
OF THE
NORTHERN UNITED STATES, CANADA
AND THE BRITISH POSSESSIONS
From Newfoundland to the Parallel of the Southern Boundary of Virginia, and
from the Atlantic Ocean Westward to the 102d Meridian
By
NATHANIEL LORD BRITTON, Ph.D., Sc.D., LL.D.
Director-in-Chief of the N. Y, Botanical Garden; Professor in Columbia University
And
HON. ADDISON BROWN, A.B., LL.D.
President of the New York Botanical Garden
Three Volumes. $13.50 Special Net; Expressage extra
SECOND EDITION—REVISED AND ENLARGED
CHARLES SCRIBNER’S SONS
FIFTH AVENUE AT 48th STREET NEW YORK
American fern Journal
An illustrated quarterly devoted to the general study of ferns, Subscription, with
membership in the AMERICAN FERN SOCIETY, $1.00 per annum; without mem-
bership, 90 cents. Volume I, six numbers, $2.00; other volumes, $1.00 each, Matter
or publication should be sent to R. C. BENEDICT, 322 East 19th Street, Brooklyn,
subscriptions and other business communications to E, J. WINSLOW, Auburn-
dale, Mass.
THE BRYOLOGIST
THE ONLY MAGAZINE IN THE WORLD RELATING
EXCLUSIVELY TO THE STUDY OF MOSSES,
HEPATICS, AND LICHENS
Illustrated with original drawings and half-tones. Twenty-first volume
begins with January, She Send for sample copy; subscription price
$1.25 per year.
EDWARD B. CHAMBERLAIN,
18 West 89th Street, NEW YORK CITY.
Large Dissecting Stand
A new design which meets every requirement by reason of its
large size and wide range of usefulness. Accommodates inter-
changeably Binocular Microscope body (illustrated), Monocular
Erecting Body, using medium high power objectives, or simple lens
in jointed arm—all focused by rack and pinion. The microscope
bodies may be mounted on a sliding track, permitting the entire
width of the stage to be covered.
Stage, 8x7 inches, is provided with glass and metal plates,
4}4 inches in diameter, and with adjustable background stops be-
With 38 and 19mm. doublet lenses . . . ae BO
With Binocular Microscope body, 40 mm. objec
tives and
10X eyepieces, as illustrated above . :
- - $70.00
Write for illustrated circulars.
Bausch €§ lomb Optical ©.
NEW YORK WASHINGTON CHICAGO FRANCISCO
LONDON ROCHESTER. N Y. FRANKFORT
Vol. 45 - No. 9
BULLETIN
OF THE
TORREY BOTANICAL CLUB
SEPTEMBER, 1918
Wound periderm in certain cacti
Mary WOoTHERSPOON COUTANT
(WITH PLATE 9 AND THREE TEXT FIGURES)
NORMAL TISSUE
The two cacti upon which the work of this paper is based are
Opuntia versicolor Engelmann and O. discata Griffiths. These
are jointed, branching forms, characteristic of the flora in the
vicinity of Tucson, Arizona. Morphologically they differ in that
the joints of the O. versicolor are composed of fluted cylinders,
while those of the O. discata are more or less disc-shaped.
Necessarily, a cross section of the former presents the appearance
of a circle with an undulating margin, while one of the latter is
more of an elongated ellipse. Anatomically considered, they are
identical in all essentials. There is (1) a distinct pith in the
center, bounded (2) by a ring of bundles equidistant from the
exterior; outward from the bundles (3) a ring of cortical tissue,
which extends to the (4) hypoderm, outside of which is the (5)
epidermis. Subepidermally, in plants several years old, normal
periderm occasionally, but not typically, forms in isolated areas.
Schleiden (3), in his early work on the cacti, describes fairly
accurately several forms, which, however, do not include either
O. versicolor or O. discata.
A further discussion of the tissues in their normal condition is
prerequisite to a complete understanding of their behavior as a
result of wounding.
[The BULLETIN for August (45: 309-352) was issued September 9, 1918.]
353
354 COUTANT: WoOOD PERIDERM IN CERTAIN CACTI
The pith is composed of thin-walled, isodiametric cells, loosely
packed together, and with consequent large intercellular spaces.
The cells themselves contain, aside from the usual protoplasmic
contents, a considerable quantity of stored starch, and small
crystals of calcium oxalate. These crystals are found in large
quantities in practically all of the tissues of both opuntias, some-
times causing, parallel with their own growth, an hypertrophy of
the cells containing them. In a concentrated solution of hydro-
chloric acid, the calcium oxalate of a medium-sized crystal does
not dissolve for over half an hour. Lauterbach (2) describes these
crystals as being star-shaped clusters composed of monoclinic
prisms, which have a short principal axis, and states that they
make up 85 per cent. of the weight of the ash.
The bundles vary somewhat in their structure with the age
of the plant and their position in the stem. Essentially, they are
of the open collateral type, with hadrome and leptome on the same
radius, separated by a cambial layer. The hadrome, much greater
in extent than the leptome, consists of annular and spiral ducts,
occasional parenchyma cells, and, in many cases, large interspersed
masses of stereome tissue. The ducts and stereome cells are
lignified, as shown by testing with phloroglucin and hydrochloric
acid, and in cross section the latter greatly resemble the median
optical view of the lignified-walled cells, to be described later in
the formation of the periderm. The cells of the cambial region,
as seen in cross section, present the typical brick-shaped appear-
ance, and form, at the most, a layer not more than three or four
cells deep. The leptome is composed of the characteristic sieve
tubes and their accompanying companion cells, interspersed with
parenchyma tissue. In cross section the peripheral portion of the
leptome mass appears dome-shaped, and is capped by a mass of
collenchymatic cells which resemble those of the hypoderm, re-
ferred to later on. The walls of these cells often become more or
less mucilaginous, an occurrence similar to that found by Lauter-
bach (2, p. 262) in Pereskia, also one of the Cactaceae. The sub-
cortical cells in the bundle region appear isodiametric, like those
of the pith. However, there is a gradual merging of these cells
with those in the outer cortical region, which are distinctly
cylindrical and arranged end on end, with their long axes at right
CouTANT: WooD PERIDERM IN CERTAIN CACTI 355
angles to the periphery. The outer five or six rows are normally
rich in chloroplasts, and practically all of the cortical tissues con-
tain a plentiful supply of stored starch. In seedlings a single-
layered epidermis covers the distinctly parenchymatic cortical
cells. However, as the plant matures, the outer cortex gives rise
to a clearly differentiated tissue, the hypoderm, socalled by
Solereder in his Systematic Anatomy of the Dicotyledons (4).
The cells of the hypoderm are, in both opuntias here studied, from
five to seven layers in depth, and decidedly collenchymatous in
character. Schleiden (3, p. 348) describes this condition as arising
through loss of water, from cells originally mucilaginous. PLATE
9, FIG. 1 will serve to give an idea of the thick cellulose walls,
traversed by pit canals which radiate out from the comparatively
small lumina, indicating protoplasmic continuity between the cell
contents. The outer layer, which lies just underneath the epi-
dermis, is characteristically so filled with oxalate crystals as to
almost obliterate the walls. Occasional crystals are found in the
other cells. This tissue has been described for different species
of Opuntia by Lauterbach (2, pp. 259-264), as well as by the two
authors mentioned above.
The epidermis is a single layer of cells in thickness and covered
thickly with cutin, as shown by staining with Sudan III and heat-
ing. The stomatal openings extend through the hypodermal
tissue down to the cortical cells. The guard cells are somewhat
sunken, and the whole cavity is lined with cutin continuous with
that on the outside. This fact was brought out by Mohl (see
Solereder, 4, p. 408) in his work on the cacti. :
The normal periderm, mentioned in the introduction as forming
in plants several years old, arises superficially, and gives the ap-
pearance of circular markings on the surface. When examined in
a transverse section of the stem, it is seen to be composed of
alternating zones of thin- and thick-walled tissue, each zone being
itself several layers of cells in thickness (PLATE 9, FIG. 6). The
thin-walled cells appear brick-shaped, and superficially greatly
resemble meristematic tissue, except that their walls are highly
suberized. The thick-walled cells are, on the contrary, lignified,
reacting with phloroglucin and hydrochloric acid as did the
stereome mentioned in discussing the structure of the bundles.
356 § COUTANT: WOOD PERIDERM IN CERTAIN CACTI
A single layer of this lignified tissue is never deeper than three cells,
arranged one on top of another; the outer tangential and the
radial walls are thicker than the inner tangential ones, and in
consequence the peripheral portions of the lumina are dome-
shaped, as is so common in thickened cutinized tissue of the
epidermis. In very thin sections stained with Bismarck brown,
distinct striations can be seen in the walls, coarser near the lumen,
and becoming finer toward the exterior; while running at right
angles to these striations are numerous fine pores. PLATE 9,
FIG. 5, gives an optical transverse view of this tissue. The middle
lamellae are shown, but the striations and pits above referred to
are not indicated. In describing the origin of this particular
tissue, Schleiden (3, p. 352) says that there arises a thick, yellow-
brown, granular, slimy mass, in which eventually cells are formed.
At present such an hypothesis is not to be accepted. However,
although omitting any reference to lignification, he brings out
clearly the fact that in the mature periderm there are alternating
zones of suberized thin- and thick-walled tissue.
EFFECT OF WOUNDING ON NORMAL TISSUE
Having studied the normal tissues, with especial reference to
the periderm, the next problem was to find out what reactions
would follow as a result of injury. Wounds were brought about
by making longitudinal slits down the plants with a razor, usually
cutting deeper than the bundle ring, but never beyond the center
of the pith. A contraction of the cells near the cut surface
resulted, thereby opening up the wound, and exposing the tissue
to the air. After the elapse of definite periods this material was
placed in alcohol. Some stems were wounded with glass, in
order to avoid a possible stimulus which might result from the
acids of the tissues acting on the steel of the razor, and the resultant
formation of soluble iron salts. In running a parallel series, how-
ever, there was found to be no difference in the rate of wound
reaction between the two, so the use of the razor was adhered to.
All of the material used came originally from Tucson, Arizona.
In the spring of 1913, cuttings were sent from Tucson, which were
immediately wounded and killed, but gave unsatisfatory results.
Plants from Tucson were wounded while growing in the Barnard
CouTANT: Woop PERIDERM IN CERTAIN CACTI SY f
College greenhouse, but, although giving fairly good results, they
were by no means as dependable as those obtained from material
wounded and killed directly in its natural habitat in Tucson.
Material of this character, collected by Dr. H. M. Richards in
1911 and by Dr. D. T. MacDougal in 1914, serves as the basis
for the observations which follow. :
Practically all of the studies have been made from transverse
sections through the stems, where, owing to the above mentioned
opening of the wound, the two exposed surfaces form a “V.”
Hand sections stained with Bismarck brown served to show clearly
all cell walls, and the unstained portions were easily tested for
cutin, cellulose, lignin, etc. Although most of the figures were
made from such hand sections, those cut with a microtome were
used for reference.
he only visible change during the first day after wounding
is the partial loss of starch in a region running parallel with the
exposed surface of the wound, but not bordering directly upon it.
By the second day, that region is entirely starchless, and the
reaction with iodine of the starch in the cells on the wound surface
does not give the normal purplish blue, but more of a reddish
color, suggesting the presence of a greater proportion of erythro-
dextrin.
In three-day-old wounds, some of the oxalate crystals in the
most exposed cells have increased in size. The starchless area in
the upper part of the “V”’ has become divided into two practically
equal parts, by the foreshadowing of a meristematic layer running
parallel to the surface of the wound. The cells in the outer region,
that is, between this future meristem and the cut surface, are some-
what discolored. The cells comprising this prospective wound-
phellogen are one layer in depth, and although no divisions have
as yet taken place, there seems to be an adjustment of the contain-
ing cytoplasm to form a flat plate, continuous from one cell to
another, and parallel to the wound surface. PLATE 9, FIG. 2, will
give an idea of how this appears in section.
A wound four days old shows the meristematic or phellogen
layer more clearly, especially near the periphery, where the cells
seem about to divide. Near the base of the cut, particularly if
the wound is deep and the circulation of air slight, it is un-
358 CoOuUTANT: WooD PERIDERM IN CERTAIN CACTI
developed, possibly due to lack of available oxygen or perhaps to
the lessening of the transpiration. See TExT FIG. 1. The latter
is suggested by a fact mentioned by Kiister (1, p. 187), that a
preliminary condition for the production of wound-cork is that
at least a small degree of transpiration must be possible for
the exposed tissue. Here, as pointed out, the cells on the cut
surface near the periphery must have at least a normal amount
of transpiration, while those deep in the cut can have much less.
Fic. Cross section of a four-day-old wound of an Opuntia discata plant,
showing os characteristic V-shaped cut and the formation of the wound-phellogen
only in the outer parts of the wound, X 28. a, phellogen layer.
Another change from the three-day-old wound is that the walls
of the discolored cells nearest to and outside of the developing
phellogen, have become lignified.
A study of a five-day-old wound can best be made by referring
to Text FIG. 2. The phellogen is clearly distinguishable and has
produced, except near the periphery, what may be spoken of
as a distinct phellem. This phellem is about three cells deep,
and the walls are highly suberized. The lignified cells just outside
COUTANT: WOOD PERIDERM IN CERTAIN CACTI 359
FIG, 2. ag ti tion through a five-day-old wound of an Opuntia
versicolor plant, X 24; the open white areas consist of normal cellulose-walled cells;
starch is found only in regions indicated. a, epidermal cells, with heavily cutinized
outer walls; 6, hypodermal tissue; c, cellulose-walled cells, containing stored starch;
d, vascular bundles; e, cells, the walls of whi ve me discolored and lignified;
f, cells, the walls of which have become discolored; g, suberized, thin-walled cells,
produced through action of the first wound-meristem.
360 CouTANT: WOOD PERIDERM IN CERTAIN CACTI
of them also give the suberin test, leaving, however, the large
number of discolored cells unlignified, or giving the lignin test and
no other. This phenomenon of cell walls giving the tests for
both suberin and lignin Kiister (1, p. 165) finds in the callus
tissue of poplar cuttings. He says he does not doubt that the
same substance, or a similar one, is the cause of these reactions.
However, this statement is perhaps open to question. There
seems no reason why both suberin and lignin might not be present
together.
In wounds six and seven days old the phellogen has produced
on the outside more suberized-walled phellem cells and on the
inside what may be called a distinct phelloderm, the walls of which
are pure cellulose (PLATE 9, FIG. 3). For the sake of convenience
these tissues will be spoken of as phellem, phellogen and phello-
derm, with the realization that, although analogous to the normal
periderm of plants, they are not necessarily homologous. Although
the phellem extends up through the hypoderm, few cells have
been produced there. At this stage, still more of the cells in the
discolored tissue have become lignified.
A wound ten days old shows several marked advances. The
periderm is in all about eighteen cells deep. However, the
phellogen, instead of producing only suberized phellem tissue,
has given rise to a few thick-walled lignified cells, five or six in a
cross section, appearing on one side, about half way down the
“V."" These cells were previously described in discussing the
normal periderm. A further lignification of the discolored tissue
is apparent. Another change is in the distribution of the stored
starch. With the formation of waterproofing cells covering the
wound, the normal activities of the tissues seem to be resumed,
and small grains of storage starch appear in the phelloderm cells,
and in the cortical cells below the phelloderm. These grains are,
however, very few and very scattered.
The next wound studied in detail was twenty-four days old.
Considering the various zones of tissues which extend from the
wounded surface into the interior of the plant, we have (1) on
the outside, the cells, in no way discolored, which retain the original
stored starch they contained at the time of wounding. Some
enclose oxalate crystals which have increased considerably in size.
COUTANT: WOOD PERIDERM IN CERTAIN CACTI 361
Within is (2) the layer of discolored cells whose protoplasmic
contents appears to have degenerated and the walls of which are
Qu
Fe iter me gag =" 5
APE OHO:
ci ee
'
'
4
1
AAO taney
Sie Site
Dy
“T4
Hh
hae
09
ae,
r
Fic. 3. Diagrammatic cross section through a thirty-one-day-old wound of an
Opuntia versicolor plant, X 24. a, old tissue, bordering upon wound surface; very
‘ol igni
m; ¢, first
fem; e, suberized thin-walled cells, produced by first
adios teaalencaes f, young vascular bundles produced by second wound-meristem;
g, “ph erm” tissue, produced by first and second wound-meristems, the cells
being cellulose-walled, and containing small amounts of stored starch and oxalate
crystals; &, old vascular bundles.
almost completely lignified. Bordering upon them is (3) the
suberized phellem tissue, averaging four or five cells deep, below
362 COUTANT: WOOD PERIDERM IN CERTAIN CACTI
which are (4) the thick-walled lignin cells, usually two, but
occasionally three deep. Inside is (5) more suberized tissue, which
arises directly from (6) the phellogen. The phelloderm (7) below
this, averages from ten to twelve cells deep, but is considerably
thicker near the base of the “‘V,’”’ where the appearance of some,
of the cells seems to indicate the formation of a second meristem.
The walls of the cells in this region are of cellulose, and the con-
tents include small, probably newly formed oxalate crystals, and
a considerable quantity of stored starch. The cortical cells (8)
underneath the phelloderm have now practically regained their
normal starch content.
TExtT Fic. 3 will aid in the study of a thirty-one-day-old
wound. The periderm is more developed, and now includes in
certain areas two more alternating layers of thick- and thin-walled
tissue. However, the big advance made here comes in the further
development of the second layer of meristematic tissue forming
within the old periderm, parallel to the exposed surface. The
function of this meristem seems to be to produce new vascular
bundles.
These bundles develop first near the innermost portion of
the wound, where the meristem was first seen to form. Kiister
(1, p. 164), in his discussion on the histology of callus, mentions the
formation of tracheids, especially in the inner layer. By the
union of many of these tracheids, he says, primitive vascular
bundles, and a wood-like tissue are produced. However, here in
the cacti, we have not just scattered rudiments of bundles, but
distinct masses arranged in a definite position.
Eventually the first wound-meristem extends peripherally into
the hypodermal tissue, but not until it is elsewhere very well
developed. Another point to be noticed is that where the bundles
were cut through, or where they are merely in a line with the
wound phellogen, their prosenchymatic as well as their parenchy-
matic cells seem capable of reverting to a meristematic condition.
Whether the stereome and hadrome tissues are able to revert is
possibly questionable, but in these wounds it does not look as if
the meristematic cells adjoining them had arisen from parenchyma
cells some distance away.
CouTANT: Woop PERIDERM IN CERTAIN CACTI 363
DIsCUSSION AND CONCLUSIONS
The series of changes which take place after injury, as described
in the different ages of the wounds, really gives in itself a summary
of what reactions took place.
The disappearance of starch in certain cells of the cacti prior
to normal cork formation, is noted by Soraurer (5, p. 428), so it
is not surprising to find it here, prior to the development of wound
periderm, which, as can be seen, greatly resembles that produced
normally.
This latter fact was noted by Schleiden (3, p. 353) who states
that the best analogy to the natural cork building process is seen
in the effect of wounding, and that this can be observed nowhere
so easily as among the cacti. He does not, however, go greatly
into detail, but describes the “drying up” of the cells on the
surface of the wound, due to the ‘‘evaporation of their liquid
contents” and says that this layer of ‘“‘dried out”’ cells forms a
protection to the cells underneath, out of which is formed the cork.
He also states that the wound formation does not take place
exclusively in cells exposed to air, but in behalf of this, gives no
convincing proof.
Kiister (1, p. 186) in his work on the histology of wound-cork,
says that the walls are always thin, and often folded, and that
differentiation of any kind whatsoever is entirely absent. This,
of course, is not the case in these cacti. The formation of the
thick-walled lignin cells is in itself enough to disprove the fact.
The essential new facts brought out by this paper are (1) the
lignification of the old outer cortical cells, (2) the production of
both thin- and thick-walled cells by the activity of the wound
phellogen, and (3) the formation of the second interior meriste-
matic layer, forming the new vascular bundles, parallel to the
wound surface.
I wish to thank both Dr. D. T. MacDougal, of the Carnegie
Desert Laboratory at Tucson, and Dr. H. M. Richards, of Barnard
College, for their kindness in providing me with material, and to
express my sincerest appreciation to Dr. Richards for the aid
he has given me in my work.
BARNARD COLLEGE BOTANICAL LABORATORY,
CotumsBiA UNIVERSITY
364 CouTANT: Woop PERIDERM IN CERTAIN: CACTI
LITERATURE CITED
_
. Kiister, E. Pathologische Pflanzenanatomie. Jena. 1903.
. Lauterbach, C. Untersuchungen iiber Bau und Entwicklung der
Sekrethehalter bei den Cacteen. Bot. Centralbl. 37: 257-264,
289-297, 329-336, 369-375, 409-412. pl. 2, 3. 1889.
. Schleiden, M. J. Beitrage zur Anatomie der Cacteen. Mém. Acad.
Sci. St. Pétersbourg (Sav. Etrang.) 4: 335-380. pl. I-ro. 1839.
. Solereder, H. Systematic Anatomy of the Dicotyledons. English
translation. Oxford. 1908.
Sorauer, P. Handbuch der Pflanzen-Krankheiten. 1. Berlin. 1909.
nN
io
-
ch
Explanation of plate 9
All figures are Zeiss camera lucida drawings from hand sections, which, with the
exception of that shown in Fic, 5, were taken transversely through the stem. In
some cases. as noted. th -e * Cee eat Men rie _
Fic. 1. Section through the hypoderm and epidermis of stem of an Opuntia
versicolor plant, stained with Bismark brown, X 730. a, epidermis; 6, outer layer
of hypodermal cells, filled with calcium oxalate crystals; c, stomatal opening, leading
down through hypodermal cells to cortical tissue; d, hypodermal cells; e,
cortical
tissue.
1G. 2. Section taken near the periphery of a three-day-old wound in an Opuntia
versicolor plant, X 245. a, cells ‘bordering on wound surface which have retained
their starch content; b, outermost Portion of starchless layer, which has become
t discolored; c¢, cells which will later divide, and so form the first wound-
meristem; d, inner portion of starchless layer.
Fic. Section from near the periphery of a seven-day-old wound in an Opuntia
versicolor plant, X 390, a, starchless cells, with lignified walls; b, starchless cells,
the walls of which give test for both lignin and suberin; c, suberized-walled cells
which have arisen through the activity of the first wound-meristem; d, first
meristem; e, “‘phelloderm" tissue, produced by first wound-meristem;
cortical cells,
Fic. 4. Section through a twenty-four-day-old wound of an Opuntia versicolor
plant, stained with Bismark brown, 7 4, starchless cells, with lignified walls;
wound-
f, starchless
cells, produced by first wound-meristem; d, inner mass
of suberized-walled cells, produced through activity of first wound-meristem; ¢,
first wound-meristem; f, “‘phelloderm” cells, produced by first wound-meristem.
Fic. 5. Transverse optical section of lignified-walled cells, produced through
activity of first wound-meristem, stained with phloroglucin and hydrochloric acid,
X 390, a, cell lumen; 6b, middle lamella.
1G. 6. Portion of wound-periderm produced in an Opuntia versicolor plant,
injured thirty-one days; all tissues shown hav
-walled cells; c, middle mass of suberized thin-walled
cells; d, inner mass of lignified thin-walled cells; ¢, inner mass of suberized thin-walled
cells, arising from first wound-meristem (not shown).
The ferns and flowering plants of Nantucket—XIX
EUGENE p. BICKNELL
SUPPLEMENTARY NOTES
Ophioglossum vulgatum L. Abundant in a meadow near the
Polpis schoolhouse, June, 1909, and in a Madequet meadow,
July, 1912. New stations of only a few plants are in Squam and
Shawkemo, in the shade of thickets, and by a pool on the Nan-
tucket golf links.
Osmunda spectabilis Willd. Of luxuriant growth among wet
thickets in Squam.
Osmunda Claytoniana L. When mentioning Clayton’s Fern
in Part I of this series no other evidence of its occurrence on Nan-
tucket could be added to Mrs. Owen’s record of a single specimen
long ago found by Mr. Dame. Later exploration has discovered
it in the present-day flora of theisland. A single tuft in a meadow
near the Creeks, observed first in 1908, was still there in 1912,
when, also, a thriving colony was met with in a thicket in Squam.
One tuft seen on Tuckernuck, June £7, 1911.
Asplenium Filix-foemina (L.) Bernh, Common locally, espe-
cially in Squam. The largest examples, in a Quaise thicket, were
over three feet in length and fourteen inches in maximum width.
Two different kinds of lady ferns grow near together at
Watt’s Run, one green throughout, and having small fronds with
close set, merely pinnate divisions, the other, characterized espe-
cially by a red-purple rachis, having broader fronds with less
crowded pinnae and more deeply cut pinnules. This red-stalked
plant, which I have long thought to be a distinct species, is com-
-mon enough among the forms combined under this fern, but I
have not seen it elsewhere on Nantucket.
It has recently been shown by Dr. F. K. Butlers (Taxonomic
and geographic studies in North American ferns. Rhodora 19:
179-216. pl, 123. 51917) that, “in the eastern parts of North
365
366 BICKNELL: FERNS AND
America there are two species of the lady fern group which appear
to be amply distinct from each other and from the true Athyrium
Filix-femina [of Europe]. These are A. asplenoides (Michx.) Desv.
and A. angustum (Willd.) Presl.”’ In accordance with the indica-
tions of this paper the common Nantucket plant should bear the
name Athyrium angustum. The large form that I have mentioned
answers to var. elatius (Link) Butters, and the red-stalked form
to var. rubellum (Gilbert) Butters. Athyrium asplenioides seems
not to have been found on Nantucket, although it is cited by Mr.
Butters from Sandwich, Massachusetts, and from Block Island.
Dennstaedtia punctilobula (Michx.) Moore. Abundant in dry-
ish open places in parts of Trot’s Swamp.
Equisetum arvense L. Proves to be well distributed, and is
abundant in many localities.
Lycopodium obscurum L. Additional stations are Taupawshas
swamp and south of the Fair Grounds, well-fruited at both places.
Lycopodium complanatum L. The Nantucket plant is the var.
flabelliforme Fernald. Localities on the eastern side of the island
are in Squam, where it fruits abundantly, near Sachacha Pond,
Pout Ponds, Shawkemo.
Lycopodium tristachyum Pursh. In great abundance in the
pine barrens towards the South Pasture, where it fruits pro-
lifically, the peduncles sometimes bearing seven spikes.
Pinus rigida Mill. Observed on the north side of Tuckernuck
Island, June 17, 1911, a few scattered trees, none of them over
five feet high.
Larix decidua Mill. In 1911 the tallest trees were estimated to
be perhaps twenty feet in height.
Juniperus virginiana L. The tallest red cedars are found
scattered through the oak woods in Coskaty, and probably exceed
twenty feet. Among the extensive growth of red cedars on Coatue
bright green and blue glaucous forms are everywhere grouped
together in agreeable contrast. On July 13, 1912, both were well
fruited, the blue form being the more prolific.
Typha angustifolia L. Under this name are two rather distinct
FLOWERING PLANTS OF NANTUCKET 367
appearing cat-tails that seem to call for critical study. One is
the abundant species of our coastwise marshes, the other a scarcer
and much more slender and narrower-leaved plant perhaps exclu-
sively of fresh water bogs. A comparison of living plants, in
July, 1912, showed a very obvious difference in their general color-
ing, the narrower-leaved plant quite wanting the bluish tinge of
leaf and stem so characteristic of the other; the leaves were also
more attenuate, and considerably longer at the time the plants
were beginning to bloom, and the very slender spikes were sepa-
rated by an unusually wide interval. Less obvious differences
were the more striate stem of the smaller plant and the absence
from its cortex of the numerous pale puncticulae that, under a
lens, appeared distinctly in the contrasted form. On July 1
the slender plant was just in flower, the broader-leaved form some-
what more advanced. Typha latifolia was not seen in flower until
July ro.
Sparganium eurycarpum Engelm. Much earlier flowering than
the other island species; fruiting heads are of full size sometimes
as early as the third week in June. S. americanum had no mature
staminate flowers up to July 12, 1912.
Potamogeton Oakesianus Robbins. Flowering and fruiting
abundantly in June.
Potamogeton pulcher Tuckerm. Flowering abundantly, some-
times before the end of May. As early as June 5, IQII, it was
blooming freely and bore well-formed fruit, while yet the leaves
were far from maturity, even the uppermost retaining their early
tenderness and pinkish-brown color.
Potamogeton pectinatus L. In full flower and with immature
fruit as early as June 10, 1908.
Ruppia maritima L. Professors Fernald and Wiegand, in
their paper, “The genus Ruppia in eastern North America”
(Rhodora 16: 119-127. pl. ro. 1914), have referred the Nantucket
plant to var. longipes Hagstrém, citing and figuring a specimen
collected in Sachacha Pond by Professor F.S. Collins. Specimens
of this form collected in Sachacha Pond, June 12, 1911, were in
full flower. It is quite possible that plants observed in a ditch on
368 BICKNELL: FERNS AND
Swain’s Neck, only just in flower September 17, 1907, were of a
different form.
Zanichellia palustris L. Fresh water ditches in Shawaukemmo
Spring meadow; abundant in a salt creek on Little Neck, where it
was flowering and fruiting freely June 22, 1910. At both stations
‘the plants appeared to be perfectly typical of var. pedunculata
J. Gay, the peduncles becoming 1-3 mm. long and bearing stipitate
nutlets; these were denticulate, and margined along the dorsal
ridge by a narrow semi-transparent membranous expansion.
Naias flexilis (Willd.) Rostk. & Schmidt. A second station is
Hummock Pond where it was collected by Miss Gardner, Septem-
ber 12, 1915.
Natas guadalupensis (Spreng.) Morong. In Hummock Pond,
the third Nantucket pond in which it is now known to occur.
Triglochin maritima L.- In flower as early as May 30, 19090;
some plants still in bloom July 10, 1912.
Alisma subcordatum Raf. Pool north of the town, 1908;
pool not far from the original station, 1912.
Sagittaria Engelmanniana J.G.Sm. In early flower July 10,
I9I2.
Sagittaria latifolia Willd. <A large form occurs at No-Bottom
Pond having the leaf blades and their basal lobes narrowly elon-
gated and tapering, sometimes becoming over 20 cm. in length
while only 10 mm, wide at the base.
Panicum linearifolium Scribn. True P. linearifolium has not
been collected on Nantucket. The grass somewhat intermediate
between it and P. depauperatum, previously referred, with reserva-
tions, to P. linearifolium, has been determined by Professor
Hitchcock to be the variety of P. depauperatum mentioned by
Hitchcock and Chase in North American species of Panicum
(Contr. U.S. Nat. Herb. 15. 1910), p. 152.
Panicum Owenae Bicknell. A fully developed mature tuft,
_ collected July 1, 1912, well attests the distinctness of this grass
from P. Addisonit Nash, to which less perfect examples were
referred by Hitchcock and Chase (North American species of
FLOWERING PLANTS OF NANTUCKET 369
Panicum, p. 243). It has now been collected at four stations
scattered within a distance of little more than a mile north and
west of the town. At each station only a single tuft was found,
or a few together, growing in association with P. depauperatum
and P. meridionale. In many of its characters P. Owenae is
somewhat intermediate between these abundant Nantucket grasses
and, although they belong to rather distinct groups in their genus,
the possibility seems to appear that P. Owenae may be a hybrid
between them. P. Owenae has been figured in Britton and
Brown’s Illustrated Flora from a specimen collected June 20, 1908.
Panicum Bicknellii Nash. Collected a second time on Nan-
tucket, June 17, 1911, on the downs east of Long Pond.
Panicum dichotomum L. Additional localities are in Siasconset
and Squam and near Sachacha Pond.
Panicum tennesseense Ashe. This proves to be rather a com-
mon species on Nantucket.
Chaetochloa versicolor Bicknell. A restudy of this grass has
not enabled me to view it as being the same as C. imberbis perennis
(Hall) Scrib. (C. occidentalis Nash) to which it has been referred.
Nor does its obviously closer relationship to true C. imberbis
establish identity with that species. Although it is said to inter-
grade with C. imberbis, I have yet to see specimens of mature
plants that cannot readily be separated.
Stipa avenacea L. Quite unexpectedly this grass has proved
to be a characteristic species of Nantucket occurring, however,
mainly on the eastern side of the island where it is well distributed
and locally abundant. It is found in scrub oak barrens along the
railroad and towards Siasconset and Sankaty Head, and in
Wauwinet, Pocomo and Quaise; it is also common on the Smooth
Hummocks on the western side of the island, and was observed on
Tuckernuck. It is locally common on Chappaquiddick Island.
Inflorescence appearing June 9, 1909; just in flower June 15, 1911,
June 24, 1910.
Alopecurus geniculatus L. Stations distant from the town are
Quidnet and Millbrook swamp.
Aira caryophyllea L. In full flower June 5, 1910, the larger
370 | BICKNELL: FERNS AND
plants 7 cm. high, but many not over 1 cm. and bearing only a
terminal spikelet. These diminutive plants were massed so thickly
over damp sandy tracts as to have the appearance of beds of
flowering moss.
Poa pratensis L. A variety that grows in sandy soil, often in
pure sand, is low and pale glaucous green, with firm culms and
contracted purple panicles, the leaves often slightly pubescent
on the upper surface and on the sheaths. In such forms the middle
and marginal veins of the palea were found to be more bearded
than in the ordinary pasture form.
Pantcularia grandis (S. Wats.) Nash. P. americana (Torr.)
MacM. Of luxuriant growth about Millbrook swamp and along
ditches west of the town.
Panicularia acutiflora (Torr.) Kuntze. Millbrook swamp;
head of Hummock Pond; Polpis; Quidnet.
Panicularia pallida (Torr.) Kuntze. A reduced form, abun-
dant in Rotten Pumpkin Pond, June 11, I9II, seemed to answer
quite exactly to descriptions of var. Fernaldii Hitche., but were
regarded by Professor Hitchcock, to whom specimens were sub-
mitted, as being scarcely typical. Ina recent paper on the “‘Status
of Glyceria Fernaldii”” (Rhodora 19: 75-76. 1917) Dr. Harold
St. John has placed this grass in specific rank under the generic
name Glyceria, marking it off sharply from P. pallida by certain
characters hitherto unnoticed, among which very small anthers
and reflexed lower branches of the panicle seem especially note-
worthy. By the test of these characters the Nantucket plant is
definitely excluded from G. Fernaldii, its anthers, even in the dried
state measuring as much as I-1.5 mm. long, and all the panicle
branches remaining flexuously ascending or erect. In other re-
spects, however, such as narrow leaves and small few-flowered
spikelets its agreement with G. Fernaldii seems to be quite perfect.
On Long Island the problem is even more confused. Some
Long Island specimens appear to have all the characters of G.
Fernaldu except reflexed panicle branches; other equally small-
anthered forms are as broad leaved as typical P. pallida; in yet
others the size of the anthers is half way between the two extremes.
It sometimes occurs that summer rains following a drought will
FLOWERING PLANTS OF NANTUCKET 3/1
revive a growth of P. pallida in drying pond holes where it had
earlier flowered as an aquatic, not infrequently bringing it to a
second flowering period, and it is to be noted that this interruption
of its course of growth has a repressive influence, sometimes very
obvious, both on the plant as a whole and in its parts. Neverthe-
less between P. pallida and authentic G. Fernaldii there is a certain
difference of aspect not readily definable and this, together with
the differing distribution of each as worked out by Dr. St. John,
may well point to something in their diversity more fundamental
than a mere response to casual seein in the conditions that
immediately affect growth.
Puccinellia distans (L.) Parl. At the only station for this
grass known to me on Nantucket, where it is in abundance, well-
developed plants possess all the characters of typical P. distans
as defined by Fernald (Rhodora 18: 12-13. | 1916), while reduced
examples, these the more numerous, correspond perfectly to the
description of var. tenuis (Uechtritz) Fernald; intermediate forms
grow with the extremes.
Festuca octoflora Walt. Not uncommon, and rather widely
distributed: near the Cliff; Island View farm; Tom Never’s
Swamp; Madequet.
Festuca myuros L. Widely scattered and growing in close
abundance at many places.
Festuca capillata Lam. Common and well distributed, even
far out on the plains and commons.
Festuca ovina L. Abundant and widespread in dry sandy soils.
Bromus tectorum L. Up to 1908 this grass had become estab-
lished only sparingly, although growing freely on the low dunes
near the bathing beach and observed at stations as far east as
Polpis and Pocomo and west towards Madequet. In succeeding
years it was found to be spreading freely and fast becoming com-
mon.
Bromus hordeaceus L. The prevailing grass in many lots and
fields.
Bromus secalinus L. Observed only at two stations in the
town and in waste ground near Surfside.
giz BICKNELL: FERNS AND
Hordeum jubatum L. Reported by Mrs. Flynn as having
been found by her as early as 1895. Collected at Siasconset,
September 14, 1915, by Miss Gardner.
Hordeum vulgare L. Awned and awnless forms are frequent in
old fields.
Eleocharis tricostata Torr. Almanac Pond remains the only
known station on Nantucket for this spike rush. It was in early
flower there June 11, 1912.
Scirpus validus Vahl. Less infrequent than at first appeared.
Additional stations are: head of hummock Pond; Squam Pond;
South Shore ponds; Coatue.
Carex hirta L. A second station is in a damp lot back of the
Springfield House, where it was well established June II, 1911.
Carex stricta Lam. Additional localities are near Long Pond
and in a bog on Little Neck.
Carex virescens Muhl. At Beechwood, in woodland shade,
occurs a taller form, dark green, with much reddened sheaths and
narrow spikes, liable to be mistaken for C. costellata Britton.
This is not an uncommon form in shaded woodland, and furnishes
examples, such as I referred to in Part III of this series, that
correspond accurately with Schkuhr’s original illustration of this
species. ; |
Carex leptalea Wahl. Additional localities are Shawankemmo
meadow, Sachacha Pond, Little Neck bog.
Carex muricata L. Found as far from the town as beyond the
water works, at Tristram Coffin’s monument around the base of
the shaft, south of the Thorn lot and east of Island Home.
Carex incomperta Bicknell. A second station is a boggy place
near Monomoy, about one and one half miles from the type
locality—June 28, 1912.
Carex Howei Mackenzie, Bull. Torrey Club 37: 245. .- ror.
Mr. Mackenzie has thus renamed this distinct sedge, the name C.
delicatula, given to it in Part III of this series, having been pre-
occupied by its employment earlier the same year by C. B. Clarke
in the Kew Bulletin. .
FLOWERING PLANTS OF NANTUCKET 373
Arisaema pusillum (Peck) Nash. On Nantucket extremes of
this plant are at marked contrast. At Watt’s Run and in Trot’s
Swamp the smallest and most delicate form, with narrow black-
purple spathes and slender linear spadix, may be seen growing near
by much larger examples having broader, striped, or even wholly
green spathes and with the spadix sometimes slightly clavate.
In some plants the spathe is fluted or ribbed on the inner as well
as the outer side. Observed in flower from June 1 to July 4.
Lemna minor L. Sachacha Pond; Shawkemo; Capaum Pond.
Lemna trisulca L. Choking the ditches in meadows north of
the town, and in great masses floating and submerged in No-
Bottom Pond.
Tradescantia virginiana L. Scattered through a sandy lot on
the north side of the town; also a large tuft in a grassy opening
in pine woods south of the Fair grounds, June, 1910.
Juncus balticus Willd. At several localities in Shawkemo
growing in abundance along the borders of fresh water bogs near
the shore as well as in brackish soil.
Lilium philadelphicum L. Leaves commonly very narrow and
often somewhat scattered, suggesting an approach to L. umbel-
latum Pursh; but the capsules, although often attenuate at base,
are very variable in size and form and are mostly obovoid or
obovoid-oblong.
Vagnera stellata (L.) Morong. Additional localities are Trot’s
swamp, where it is locally common, and Thorn lot, Shawkemo.
Small plants growing on the open plains near the South Shore
have the later leaves perfectly glabrous beneath.
Polygonatum biflorum (Walt.) Ell. Salomonia biflora Farwell.
To be regarded as not uncommon rather than scarce or local,
but not met with on the western side of the island.
Medeola virginiana L. Restricted to the same parts of the
island as the preceding and mainly to the same thickets, in some
of which, especially in Shawkemo, it is a frequent plant.
Smilax herbacea L. Not infrequent, extending from the locali-
ties previously reported into Pocomo.
374 BICKNELL: FERNS AND
Hypoxts hirsuta (L.) Coville. A second station is near Long
Pond where a solitary flowering plant was found June 1, 1909.
Gymnadentopsis clavellata (Michx.) Rydb. A second station,
for this orchid is ‘‘‘near the Cliff, 1888,’ Miss Mary Foster Coffin
and Leroy Schumacher,” F. G. Floyd.
Blephariglottis ciliaris (L.) Rydb. ‘‘‘Not far from the first
station, about twenty-five plants, 1897,’ Lorin L. Dame,” F.
Floyd.
Liparis Loeselii (L.) Richard. ‘‘Near the Cliff, one plant,
1888, and Monomoy, about a dozen plants, 1891,’ Miss Mary
Foster Coffin,” F. G. Floyd.
Salix tristts Ait. On the north side of the island observed
only near Capaum Pond, a single patch.
Salix petiolaris Sm. A second locality is Trot’s Swamp, I910.
SALIX BEBBIANA X CINEREA? A willow collected in Tom
Never’s swamp, June 3, 1908, growing with other willows, includ-
ing the tree of Salix cinerea that is established there, and also
Salix Bebbiana, quite unmistakably, I think, shows in combina-
tion leaf and branch characters of both these species as well as
other evidences of being a hybrid.
Another ambiguous willow collected near Gibbs’ Pond, Septem-
ber 19, 1899, is strongly suggestive of a cross between Salix erio-
cephala and S. Smithiana, both in the shape of the leaves and
their dense velvety pubescence.
Myrica Gale L. A second station is by a tidal pond near
Abraham’s Point where a thick cluster six feet across and three
feet in height was observed June 7, 1911, no specimens being taken.
In specimens collected at the Capaum Pond station, September 12,
1907, the leaves are either wholly glabrous beneath or with some
obscure pubescence on the midrib near the base. But specimens
in the herbarium of Miss Gardner collected at the same station,
August 2I, 1915, are more or less evidently pubescent on the
lower surface. I do not know, therefore, whether an exact deter-
mination of the Nantucket plant would place it with the typical
form of the species or with var. subglabra (Chevalier) Fernald
(Rhodora 16: 167. 1914).
FLOWERING PLANTS OF NANTUCKET 375
Myrica carolinensis Mill. At Coskaty twelve to fourteen feet
high, the stoutest trunk thirteen inches in circumference; a some-
what lower shrub near Abraham’s Point was fourteen inches in
girth of trunk.
Hicoria alba (L.) Britton. A wide-spreading tree in a Shaw-
kemo thicket, in 1909, was about twelve feet in height and thirty-
one inches around the trunk; the lower branches descended to the
ground and in their widest reach overspread a space paced at
forty feet.
Hicoria glabra (Mill.) Britton. The stump of a tree in Shaw-
kemo, recently felled in 1901, measured thirty-seven inches in
circumference.
Hicoria microcarpa (Nutt.) Britton. Not far from the tree
at Wauwinet previously reported is a group of three trees dis-
covered June 10, 1911, the largest of which, about fifteen feet in
height, had a maximum trunk circumference of twenty-six inches
and a spread of branches close to the ground of not less than thirty-
five feet.
» 3
Carpinus caroliniana Walt. ‘‘‘Rare and shrublike, 1901,
Lorin L. Dame,” F. G. Floyd.
Betula populifolia Marsh. The largest trees seen grew among
pines near the Wauwinet road and, in 1911, were estimated to be
eighteen feet in height.
Fagus grandifolia Ehrh. Undoubtedly the largest of our forest
trees growing on Nantucket today are beeches. The tallest are
in Beechwood and must be fully thirty-five feet in height. The
stoutest trunks seen there were, one fifty inches, another forty-four
inches in girth near the base. A much stouter tree in Squam,
difficult of access through its encompassment of dense thickets,
although not over twenty feet in height, measured seventy-three
inches in circumference one foot above the base.
Quercus coccinea Muench. Native trees occur at Coskaty, the
largest twenty-five to thirty feet in height, by estimate, the stoutest
trunk forty-one inches around one foot above the base.
Quercus velutina Lam. The stoutest black oak seen on Nan-
376 BICKNELL: FERNS AND
tucket, in Squam, measured forty-two inches in maximum girth.
But much stouter trees grow on Tuckernuck where trunks were
measured forty-seven and fifty inches and one sixty-eight inches
in circumference about one foot above the ground, decreasing to
forty-five inches a span higher up.
Quercus stellata Wang. New stations are Acquidness Point,
a close group of five trees, the largest eight feet high (1911) and
near Wigwam Ponds, four scattered trees from four to seven feet
high (1912).
Quercus pagodaefolia (Elliot) Ashe. On June 9, 1911, the oak
tree discussed in Part IV under this name was found to be dead,
but a second and perfectly healthy tree of about the same height,
and twenty-five inches in girth of trunk near the base, was dis-
covered not far off in the same thicket. Some old acorn cups
found beneath the tree are saucer-shaped to hemispheric, some of
then contracted to a short scaly base, 1.5-3 cm. wide, the slightly
tomentulose scales closely imbricated to a firm margin, the indi-
vidual scales contracted to a lanceolate obtuse termination. A
few partly decayed nuts were ovoid-globose, the exposed portion
apparently about one half their length.
Quercus ilicifolia Wang. Two forms of the bear oak differing
markedly in leaf pattern are common on Nantucket. In char-
acteristic form the leaf of this oak has short lobes of more or less
triangular general outline. In the variant the lobes of the leaves
are narrower and more tapering, becoming lanceolate or even
somewhat falcate, the sinuses cut much nearer to the midrib and
the terminal lobe often elongated after the manner of Q. falcata
Michx. Some leaves indeed are strikingly similar to those of that
species. The two forms of leaf present extremes of divergence
that are very noteworthy and suggest an Sree subject for
critical study.
Quercus rufescens (Rehder) sp. nov.
Quercus prinoides var. rufescens Rehder, Rhodora 9: 60. 1907.
This scrub oak proves to maintain so notable a constancy in its
characters and is so readily recognizable that, as a result of field
observations both on Nantucket and on Long Island I have come
FLOWERING PLANTS OF NANTUCKET 377
to accept it as essentially distinct from Q. prinoides; nor have I
yet encountered any trees so nearly intermediate between the two
as to have the appearance of being hybrids.
It may be noted that the anthers of Q. rufescens are perceptibly
smaller on relatively shorter filaments than those of Q. prinoides,
that is to say this proved to be true in numerous cases when I
was enabled to make satisfactory comparison in the field.
QUERCUS ALBA X RUFESCENS? An oak about five feet high of
somewhat straggling form, observed near the Wigwam Ponds,
July 6, 1912, conveyed a strong impression of being a hybrid
between Q. alba and Q. rufescens. The pubescence of the leaves
and branchlets is much as in Q. rufescens, while the size of the
leaves, some of them 14 X 8 cm., and their deep lobing seem to
point rather clearly to Q. alba as one of its parents.
QUERCUS ILICIFOLIA X VELUTINA Rehder. A thriving oak, a
hybrid, there seems little reason to doubt, grows in the dense
- thicket on the western side of Dyleave Swamp. It was about
twelve feet in height and nineteen inches in girth low on the trunk
June 26, 1910. In its foliage it is very distinct from any other
Nantucket oak that I have seen, and the forms of its leaves and
their pubescence suggest, rather convincingly, I think, a mixed
origin from the two oaks above indicated as probable parents.
Such a hybrid was described and figured by Mr. Rehder (Rhodora
3: 137: pl. 24. 1901). The leaves of the Nantucket tree show
remarkable variation in form, and have no close similarity to
those figured by Mr. Rehder, but the pattern characters by which
they differ appear to be such as might well result from a crossing
of Q. velutina with the form of Q. ilicifolia having narrow deeply
cut leaf lobes rather than with the more usual short-lobed form.
They are, however, considerably larger than those which Mr.
Rehder has described, the largest being 17 X 14cm. When young
the lower surface is whitened with a close tomentulose pubescence,
although in less degree than those of Q. ticifolia, but this is im-
permanent, the leaves becoming quite green beneath in age. The
cups of some old acorns found beneath the tree were turbinate,
their scales tomentulose to partly glabrate; nuts conic ovoid or
oblong, 10-14 mm. long, tomentulose and strigose towards the
apex.
378 BICKNELL: FERNS AND
QUERCUS COCCINEA X VELUTINA? Ambiguous trees at Coskaty
growing with Q. coccinea and (Q. velutina are difficult to under-
stand except as hybrids of those species.
Quercus palustris Muench. Besides the introduced pin oaks
still persisting in the Thorn lot, a few scattered trees, the largest
eight to ten feet in height, were observed among the Miacomet
pines in 1909, and a single tree among pines south of the fair
grounds in 1910. The latter had the appearance of being ad-
ventive.
Quercus bicolor Willd. Some small trees persist in the Thorn lot
among the other introduced oaks previously reported as growing
there.
Humulus Lupulus L. Localities far from cultivated ground
where the hop has every appearance of being indigenous are
Trot’s swamp; west side of Long Pond; Swamp in Madequet,
where there is much of it.
Cannabis sativa L. Prospect Hill, herbarium of Miss Gardner;
Pleasant Street, Mrs. Flynn.
Rumex crispus L. Plants with the grain short and rounded
seem to be less common than those with attenuate and acute
grain (Rumex elongatus Guss.); the character, however, seems to
be inconstant and even varies much in the same plant.
Rumex persicarioides L. Dr. Harold St. John, who has revised
our former understanding of this species (Rhodora 17: 73-83.
pl. 113, f. 34. 1915), considers that the Nantucket plant should
stand as Rumex maritimus var. fueginus (Phil.) Dusén.
Polygonum maritimum L. Professor Fernald has convincingly
shown that our plant is distinct from the European P. maritimum
and should bear the name Polygonum glaucum Nutt. (Rhodora
15: 68-73. 1913).
Persicaria pensylvanica (L.) Small. Two varieties of this com-
mon plant have recently been recognized by Professor Fernald
(Rhodora 19: 70-73. 1917). The prevailing Nantucket form,
described in Part V of this series, is Polygonum pensylvanicum
var. nesophilum Fernald. The closely related var. laevigatum Fer-
FLOWERING PLANTS OF NANTUCKET 379
nald also occurs on Nantucket as well as the more distinct type
form of the species, which is scarce.
Agrostemma Githago L. Occasional in fields and waste places.
Dianthus Armeria L. In abundance by the railroad near the
town; on the Cliff; near the waterworks.
Alsine graminea (L.) Britton. Old cartway over the commons
east of the town, June 14, 1911; north of the town, June, 1912.
Castalia odorata (Ait.) Woodville & Wood. In Wigwam Pond,
July 7, 1912, some leaves were as large as twelve and one half
inches long and wide.
Liriodendron Tulipifera L. Mr. Floyd has sent me a note that
young tulip trees were found by Mr. Dame in 1901 “orowing
freely in the Thornlot.” Had any of these trees long survived
it would seem that other collectors would have since observed
them.
Coptis trifolia (L.) Salisb. Bloomingdale Swamp, June 17,
1914, collected by Mrs. G. A. Spear; specimen examined by me
in Miss Gardner’s herbarium. It is a matter of much satisfaction
that the gold thread is thus definitely established as a Nantucket
plant. It had been admitted in its place in this catalogue because
included in Mrs. Owen’s list, and also reported to me as having
been found in the Thorn lot, but it has since appeared that both
of these records are open to serious doubt. In a letter to Mr.
Floyd under date of August 21, 1911, Mrs. Owen wrote, “*T find
no authority for Coptis on Nantucket,’’ adding that she could
not understand how the name got into the original Godfrey list.
‘When I was compiling my ‘Flora of Nantucket,’ she stated
further, ‘‘I probably used that printed list and took without
question what I found there.” The Thorn lot report I feel sure
should be regarded as an error involved in some way with the
trifoliolate-leaved Rubus hispidus, an abundant plant in that
locality.
Actaea rubra (Ait.) Willd. In dense thickets eastward from
the locality previously reported as far as Beechwood. Much
scarlet fruit June 26, 1912.
Ranunculus repens L. Abundant all over the island, mainly
in low grounds.
380 BICKNELL: FERNS AND
Thalictrum polygamum Muhl. Previously admitted because
of Mrs. Owen’s record on the authority of Mr. Dame. The dis-
covery that Thalictrum dasycarpum Fisch. & Lall. occurs on Nan-
tucket now makes necessary a substantiation of this record.
Sassafras Sassafras (L.) Karst. Trees as high as twelve to
fifteen feet were seen in Shawkemo and Quaise, and in Squam one
not less than eighteen feet, its trunk twenty-three and one quarter
inches in circumference one foot above the ground. |
Radicula palustris (L.) Moench. Waste ground north of the
‘ town, June 9, I91I, a group of plants just in flower; bog near
Oldest House, August 18, 1915, herb. Grace Brown Gardner.
Erysimum cheiranthoides L. Recently collected by Miss
Gardner on the Cliff road, a second station for Nantucket.
Conringia orientalis (L.) Dumort. Yard on North Water
Street, June 17, 1911, one well-fruited plant; waste ground at
' Surfside, July 9, 1912, several fruiting plants and one just in bloom.
Reseda lutea L. A few plants in a field near Island Home,
just in flower June 14, 1911; Fay farm, Squam, July 11, 1912, in
flower and fruit. The original station in Polpis mentioned by
Mrs. Owen was visited by Mr. Floyd in 1895; some plants were
still growing there ‘‘although subjected to occasional mowing.”
Drosera rotundifolia L. Racemes of buds but no open flowers,
_ July 6, 1912. On Martha’s Vineyard belated flowers of this
sundew, as of the next, persist into September.
Drosera intermedia Hayne. D. longifolia L. in part. First
flowers July 9, 1912.
Drosera filiformis Raf. Not yet in flower July 2, 1912.
Rosa cinnamomea L. Here and there by fieldsides and fence
borders in the suburbs.
Amelanchier nantucketensis Bicknell. Further observation of
this shadbush, later in the summer than I had previously studied
it, show that the fruit ripens early in July, when it is red or purple
in color, later becoming deep purple blue. By the middle of the
month the shrubs, sometimes even those of the smallest size, may
be seen thickly ornamented with the glaucous fruit in all stages of
FLOWERING PLANTS OF NANTUCKET 3881
maturity and varying in color from white through pink and red
to purple all in the same cluster, producing a variegated and
very pleasing color effect.
It has been suggested by Professor Wiegand in a paper entitled,
‘““The genus Amelanchier in eastern North America’”’ (Rhodora 14:
117-161. pl. 95, 96. 1912) that this Nantucket shrub is a hybrid
between A. oblongifolia and a proposed new species, named by him
A. stolonifera. This theory, I think, scarcely takes sufficiently
into account that A. nantucketensis is one of Nantucket’s char-
acteristic shrubs, more numerous indeed, and more generally
distributed than the other shadbushes of the island, and also
bearing fruit more abundantly and with greater regularity season
by season. In Professor Wiegand’s description of his A. stolonifera
there seems to be little to differentiate it from A. nantucketensis
beyond larger petals, a variable character, I find, in A. nantucket-
ensis, and densely woolly summit of the ovary. In A. nantucket-
ensis the exposed surface of the ovary is, as I have described it,
nearly or quite glabrous; this is its condition at maturity, in its
earlier stages it is clothed with a white woolly pubescence. Its
close relationship to Professor Wiegand’s new species is also to be
inferred from its having been attributed to Nantucket on the basis
of Miss Day’s collection, No. 95, June, 1900, a sheet of which in the
herbarium of the New York Botanical Garden formed part of the
material used in formulating my description of A. nantucketensis.
This specimen is the counterpart of numerous specimens of my
own collecting, and was not cited only for the reason that the
species was so abundant on Nantucket that no need appeared of
any Citation of specimens other than the type and co-type.
Crataegus chrysocarpa Ashe. A group of four shrubs, June 29,
1912, about two miles west of the original station.
Prunus maritima Wang. I do not know that the yellow or
amber fruited form of the beach plum has ever been found on
Nantucket. I was told that it occurred on Tuckernuck and it is
locally abundant on Chappaquiddick Island and in other parts of
Martha’s Vineyard. It is not generally disseminated and is evi-
dently scarce elsewhere than in the localities where it abounds.
I have myself seen little of it except as gathered by the islanders
382 BICKNELL: FERNS AND
who, in collecting it, are accustomed to keep it separate from the
purple form for use, I was told, in making a light-colored jelly.
On September 16, 1913, a group of plum gatherers returning across
Chappaquiddick Island were laden with baskets of these plums,
as many filled with the amber fruit as with the purple. I have
likewise seen large sacks of the amber plums at Edgartown which,
it was said, had been brought in from Chilmark. No constant
difference in size or form between the yellow and purple fruit could
be discovered, but on Chappaquiddick I observed a particular bush
of the yellow-fruited form on which the plums were of unusually
large size and ovoid rather than globose. The yellow fruit is
somewhat translucent, and varies in color from amber to light
yellow and pale orange.
Laburnum vulgare Gris. Several well-grown trees of the la-
burnum are scattered in partly open ground among the pines
near Miacomet Pond, where they were in full bloom June 11, 19rt.
Erodium cicutarium (L.) L’Hér, “A well-established colony of
about twenty plants in a field near the Thorn lot,” in flower and
fruit, September 15, 1915; specimens in the herbarium of Miss
Grace Brown Gardner. Not reported as growing in Nantucket
since 1851.
Linum striatum Walt. Collected by Miss Gardner at Hum-
mock Pond, September 12, 1915, the second known station on
Nantucket.
Cicuta bulbifera L. Collected by Miss Gardner, September,
1915, at the original station discovered in 1899 by Mrs. Flynn.
Arctostaphylos Uva-ursi (L.) Spreng. Mr. Floyd writes me
that he has occasionally observed on the commons a form with
crimson flowers, and that Mrs. Owen had told him that this
crimson-flowered bearberry had been familiar to her for many
years,
AMSINCKLA ARENARIA Suksd. <A recent paper by Dr. J.
Francis Macbride (A Revision of the North American Species of
Amsinckia, Contr. Gray Herb. 49. 1917) makes clear that the
plant reported in Part XIV of this series as A. intermedia F. & M.
is not that species. Specimens have been submitted to Dr.
FLOWERING PLANTS OF NANTUCKET 383
Macbride who considers them referable, perhaps not without
some uncertainty, to A. arenaria Suksd. of the northwest.
Thymus Serpyllum L. Law nor Sankaty Bluff, August 28,
1915, Miss Gardner.
Setiscapella subulata (L.) Barnhart. Peat bog between Polpis
and Quidnet, 1915, Miss Gardner.
Houstonia caerulea L. Miss Alice O. Albertson has sent me an
interesting specimen of this species collected on the Nantucket
golf course, May 13, 1916. It is matted and subspreading, the
stems and long peduncles relatively stout, and the flowers large,
spreading 1.5 cm. in the pressed plant; the calyx lobes are oblong
to obovate-oblong and more or less foliaceous, becoming 1.5 mm.
or more wide, with somewhat spreading tips, and equalling the
short and broad tube of the corolla in the single specimen seen.
Viburnum venosum Britton. The largest example seen, Rattle-
snake bank, June 8, 1911, was of the stature of a small tree, the
trunk eleven inches in girth near the base.
Lactuca virosa L. Collected by Miss Gardner in the town,
August 24, 1915, a second station on Nantucket for this intruding
weed, and too probably an indication that it is becoming estab-
lished there.
ADDENDUM
Under Sabbatia campanulata in Part XIV of this series read,
on page 32, S. stellaris for S. campanulata.
The following two plants, new to Nantucket, have recently
come to light in a collection of specimens kindly sent to me by
Mrs. Flynn:
*HELIANTHUS PETIOLARIS Nutt. Old field near Windmill,
August 5, 1911, collected by Mrs. Nellie F. Flynn.
*KOELLIA INCANA (L.) Kuntze. Collected by Miss Mina K.
Goddard, August 17, 1895. : Locality not mentioned. The species
is an interesting addition to the native flora of Nantucket. Mrs.
Flynn, who has recently made inquiries in regard to this specimen,
writes me that Miss Goddard believes it was probably found at
Wauwinet where, as disclosed by her records, she collected plants
on the very date that appeared on the specimen label.
INDEX TO AMERICAN BOTANICAL LITERATURE
1902-1918
The aim of this Index is to include all current botanical literature written by
stein published i in America, or based upon American material ; the word Amer-
ica being used in the broadest sense.
Batis and papers that relate exclusively to forestry, agriculture, horticulture,
eee products of vegetable origin, or laboratory methods are not included, a
o attempt is made to index the literature of bacteriology. An occasional exception is
mis in favor of some paper appearing in an American periodical which is devoted
wholly to botany. Reprints are not mentioned unless they differ from the original in
Some important particular. If users of the Index will call the attention of the editor
to errors or omissions, their kindness will be appreciated.
This Index is reprinted monthly on cards, and furnished in this form to subscribers
at the rate of one cent for each card, Selections of cards are not permitted ; each
— must take all cards published during the term of his subscription, Corre-
pondence relating to the card issue should be addressed to the Treasurer of the Torrey
etna ical Club,
Adams, J. F. _ Origin and development of the lamellae in Schizophyllum
commune. Mem. Torrey Club 17: 326-333. pl.g +f. a,b. 10 Je
1918. j
Abrams, L.R. A phytogeographic and taxonomic study of the southern
California trees and shrubs. Bull. N. Y. Bot. Gard. 6: 300-485.
pliaj. 27S1
Includes Lupinus Brittoni, Amorpha occidentalis, Ceanothus austro-montanus,
and Malacothamnus Nuttallii, spp. nov.
Arthur, J. C., & Johnston, J. R. Uredinales of Cuba. Mem. Torrey
Club 17: 97-175. 10 Je 1918. [Illust.]
Includes new species in Cronartium (1), Cionothrix (1), Ravenelia (1), Uromycla-
dium (1), Puccinia (3), Aecidium (2), and Uredo (3)
Ashe, W. W. Loblolly or North Carolina pine (Pinus taeda Linnaeus).
N. Carolina Geol. & Econ. Surv. Bull. 24: i-xvi + 1-176. pl. 1-27.
1915.
Atkinson, G. F. Six misunderstood species of Amanita. Mem.
Torrey Club 17: 246-252. 10 Je 1918.
Barnhart, J. H. Historical sketch of the Torrey Botanical Club.
Mem. Torrey Club 17: 12-21. 10 Je 1918.
Beal, A. C. Gladiolus Studies-I. Botany, history, and evolution of
the Gladiolus. Cornell Ext. Bull. 9: 93-188. f. r-9. D 1916.
386 INDEX TO AMERICAN BOTANICAL LITERATURE
Beauvard, G. Monographie du genre Melampyrum L. Mém. Soc.
Phys. et Hist. Nat. Geneva 38: 291-657. f. I-31. 1916.
Berry, E.W. The lower Eocene floras of southeastern North America.
U.S. Geol. Surv. Prof. Paper 91: 1-481. pl. 1-117. 1916.
Many new species are described. \
Blodgett, F.H. Weather conditions and ‘crop diseases in Texas. Mem.
Torrey Club 17: 74-78. 10 Je 1918.
Beach, S. A. Grape breeding: size, weight and specific gravity of the
seed as correlated with germination and vigor of the seedling.
, Proc. Soc. Hort. Sci. 1903-1904: 42-53. 1905.
Boas, H. M. The individuality of the bean pod as compared with
that of the bean plant. Mem. Torrey Club 17: 207-209. 10 Je
1918,
Bowman, H. H. M. Report on botanical investigation at Tortugas
laboratory, season 1916. Year Book Carnegie Inst. Wash. 1916:
188-192. 1916.
Braun, E. L. The vegetation of conglomerate rocks of the Cincinnati
formation. Plant World 20: 380-392. f. 7-5. D 1917.
Breazeale, J. F. Effect of sodium salts in water cultures on the ab-
sorption of plant food by wheat seedlings. Jour. Agr. Research
7: 407-416. f. I-8. 27 N 1916.
Britton, N. L. Botanical contributions. Contributions to the flora
of the Bahama Islands—IV. Bull. N. Y. Bot. Gard. 5: 311-318.
8 F 1909.
Includes descriptions of Zamia lucayana, Ibidium lucayanum, Badiera oblongata,
Passiflora bahamensis, Rochefortia bahamensis and Tetranthus bahamensis, spp. nov.
Britton, N. L. Torrey Botanical Club reminiscences. Mem. Torrey.
Club 17: 24-28. 10 Je 1918.
Brown, N. A. Some bacterial diseases of lettuce. Jour. Agr. Research
13: 367-388. pl. E., 29-41. 13 My 1918.
Burgess, E.S. A method of teaching economic botany. Mem. Torrey
Club 17: 52-55. 10 Je 1918.
Burlingham, G. S. A preliminary report on the Russulae of Long
Island. Mem. Torrey Club 17: 301-306. 10 Je 1918.
Buscalioni, L., & Muscatello, G. Studio anatomo-biologico sul gen.
“Saurauia” Willd., con speciale riguardo alle specie americane.
Malpighia 28: 49-81. 1917; 140-162. 1917; 239-270: 1917.
Buscalioni, L., & Muscatello, G. Studio monografico sulle specie
americane del gen. “Saurauia’’ Willd. Malpighia 28: 1-48.
1917: 107-138. 1917; 223-238. 1917.
INDEX TO AMERICAN BOTANICAL LITERATURE 387
Campbell, D. H. The origin of the Hawaiian flora. Mem. Torrey
Club 17: 90-96. 10 Je 1918.
Candolle, C. de. The Hawaiian peperomias. College Hawaii Publ.
Bull. 2: 5-38. 16 O 1973.
Cannon, W. A. Relation of the rate of root growth in seedlings of
Prosopis velutina to the temperature of the soil. Plant World
40. 320-333. J. 1-3.. O 1917.
Chavarria, A. P, Apuntes para el estudio de la flora nacional. Bol.
Soc. Cien. Nat. Inst. La Salle 6: 135-138. O 1917.
Cobb, F., & Bartlett, H. H. Purple bud sport on pale flowered lilac
(Syringa persica). Bot. Gaz. 65: 560-562. f. 1. 18 Je 1918.
Coblentz, W. W. The exudation of ice from stems of plants. Mo.
Weather Rev. 42: 490-499. f. 1-18, Au 1914.
Cooper, J. R. Studies of the etiology and control of blister canker on
apple trees. Nebraska Agr. Exp. Sta. Research Bull. 12: 1-117.
pl. 1-24. 15D 1917.
Cortes, S. Flora de Colombia. Rev. Médica de Bogota 34: 523-528.
oS aa
Includes a. tricolor and Dimorphylia iconongiana, gen. et spp. nov.
Demarest, S. A. A sketch of the life of Coe Finch Austin. Mem.
Torrey Club 17: 31-38. 10 Je 1918.
Dodge, B. O., & Adams, J. F. Some observations on the development
of Peridermium Cerebrum. Mem. Torrey Club 17: 253-261.
pl. 4-6 + f.I~3. to Je 1918.
Dodge, C. K. Catalog of plants. Michigan Geol. & Biol. Surv,
Publ. 4: (Biol. 2) 65-120. 1911.
Duggar, B. M. The physiological effects of shading plants. Proc.
Soc. Hort. Sci. 1903-1904: 15-26. 1905.
Evans, A. W. The air chambers of Grimaldia fragrans. Bull. Torrey
Club 45: 235-251. f. 1-14. 20 Je 1918.
Farwell,O.A. Sisyrinchium Bermudiana. Mem. Torrey Club 17: 82,
83. 10Jerg18. :
Fawcett, W., & Harris, W. Elementary notes on Jamaica plants—V.
Jamaica Bot. Dept. Bull. 9: 145-148. pl. 5-7. O 1902.
Flint, E. M. Structure of wood in blueberry and huckleberry. Bot.
Gaz. 6§: 556-559. pl. ro, rz. 18 Je 1918.
Graff, P. W. Philippine micromycetous fungi. Mem. Torrey Club
17: 56-73. 10 Je 1918.
Ascophanus verrucosporus, Meliola Litseae, Phyllosticta Brideliae and Actinotiy-
rium Hopeae, spp. nov., are described.
388 INDEX TO AMERICAN BOTANICAL LITERATURE
Graham, M. Centrosomes during early fertilization stages in Preissia
quadrata. Mem. Torrey Club 17: 323-325. pl. 8. 10.Je 1918.
Gruber, C. L. Experiences with a fern garden—IJ. Am. Fern Jour.
7: 114-121. Dirg17:-—III. Am. Fern Jour. 8: 12-16. Mr 1918.
Haasis, F. W. Comparative length of growing season of ring-porous
and diffuse-porous woods. Plant World 20: 354-356. N 1917.
Harper, R. A. The evolution of cell types and contact and pressure
responses in Pediastrum. Mem. Torrey Club 17: 210-240. f. I-27.
10 Je 1918.
Harper, R. M. The vegetation of the Hempstead Plains. Mem.
Torrey Club 17: 262-286. pl. 7 + f. 1-3. 10 Je 1918.
Harris, J. A. On the osmotic concentration of the tissue fluids of
desert Loranthaceae. Mem. Torrey Club 17: 307-315. 10 Je
1918.
Harris, J. A., Gortner, R. A., & Lawrence, J. V. Studies on the phy-
sico-chemical properties of vegetable saps. Biochem. Bull. 4:
52-78. Mr 1915.
Hauman-Merck, L. Observations d’éthologie florale sur quelques
espéces chiliennes. Rev. Chilena Hist. Nat. 17: 153-159. f. 17, 18.
Je 1913.
Hazen, T. E. The trimorphism and insect visitors of Pontederia.
Mem. Torrey Club 17: 459-484. pl. 14, 15 +f. 1-12. 10 Je 1918.
Hedrick, U. P. Effects of superheated soils on plants. Proc. Soc.
Hort. Sci. 1905: 51-55. Mr 1908.
Hicken, C. M. Sobre las Polipodiaceas Argentinas. Rev. Chilena
Hist. Nat. 14: 123-134. Ig10.
Hochreutiner, B. P. G. Critical notes on new or little known species
in the herbarium of the New York Botanical Garden. Bull. N. Y.
Bot. Gard. 6: 262-299. 27 S 1910.
Nineteen new species in various genera are described.
Holden, R. On the anatomy of two Palaeozoic stems from India.
Ann. Bot. 31: 315-326. pl. 17-20. O 1917.
Includes Dadoxylon indicum and D. bengalense, spp. nov.
Hollick, A. Torrey Botanical Club reminiscences. Mem. Torrey
Club 17: 29, 30. 10 Je 1918.
Horst, A. Observaciones sobre la biologia de la Tetilla hydrocotylifolia
DC. Rev. Chilena Hist. Nat. 19: 22-29. pl. 4-6. Ap 10915.
Léveillé, H. Les Rubus de l’Argentine et du Chili. Rev. Chilena
Hist. Nat. 24: 90-93. 30 Je 1917.
INDEX TO AMERICAN BOTANICAL LITERATURE 389
Levine, M. The physiological properties of two species of poisonous
mushrooms. Mem. Torrey Club 17: 176-201. pl. 1, 2+/f. 1, 2.
Io Je 1918.
Lewton, F.L. Kokia: a new genus of Hawaiian trees. Smithsonian
Misc. Coll. 605: 1-4. pl. 1-5. 220 1912.
Lewton, F. L. Rubelzul cotton: a new species of Gossypium from
Guatemala. Smithsonian Misc. Coll. 604: 1, 2. pl. 1, 2. 21 O 1912.
Lloyd, F. E. The effect of acids and alkalis on the growth of the pro-
toplasm in pollen tubes. Mem. Torrey Club 17: 84-89. 10 Je
1918.
Macdougal, D. T., & Cannon, W. A. The conditions of parasitism in
plants. 1-60. pl. 1-10 +f. 1, 2. IgIo.
Carnegie Inst. Wash. Publ. 129.
Ma4rquez, C.C. Tratado elemental de botanica. 1-533. Bogota. 1913,
McAtee, W. L. A sketch of the natural history of the District of
Columbia together with an indexed edition of the U. S. Geological
Survey’s 1917 map of Washington and vicinity. Bull. Biol. Soc.
Washington 1: 1-142. My 1918.
Contains lists of plants of the District, a chapter on ‘‘ Magnolia bogs near Wash-
ington, D. C., and their relation to the Pine Barrens,” etc.
McDougall, W. B. Some edible and poisonous mushrooms. Bull.
Illinois State Lab. Nat. Hist. 11: 413-555. pl. 86-143 +f. 1. N
I9I7.
Medsger, O. P. Two months in the southern Catskills. Mem. Torrey
Club 17: 294-300. 10 Je 1918.
Murrill, W. A. Collecting fungi at Delaware Water Gap. Mem.
Torrey Club 17: 48-51. 10 Je 1918.
Nichols, G. E. The interpretation and application of certain terms
and concepts in the ecological classification of plant communities.
Plant World 20: 305-319. O1917;—II. Plant World 20: 341-353.
N 1917.
Nishimura, M. A carrier of the mosaic disease. Bull. Torrey Club
45: 219-233. pl. 7. 20 Je 1918.
Olive, E. W. Potato diseases. Brooklyn Bot. Gard. Leaflets. 6:
[1-4]. 29 My 10918.
Rands, R. D. Early blight of potato and related plants. Wisconsin
Agr. Exp. Sta. Research Bull. 42: 1-48. f. 1-10. Ap 1917.
Renaudet, G. Notas sobre las adquisiciones recientes de la fitoqui-
mica y de la botanica médica. Rev. Chilena Hist. Nat. 14: 55-65.
I9I0.
390 INDEX TO AMERICAN BOTANICAL LITERATURE
Richards, H. M. Determination of acidity in plant tissues. Mem.
Torrey Club 17: 241-245. 10 Je 1918.
Robbins, W. J. Direct assimilation of organic carbon by Ceratodon
purpureus. Bot.,Gaz. 65: 543-551. f. 1-5. 18 Je 1918.
Robbins, W. W. Successions of vegetation in Boulder Park, Colorado. -
Bot. Gaz. 65: 493-525. f. 1-14. 18 Je 1918.
Rock, J. F. New species of Hawaiian plants. College Hawaii Publ.
Includes Cyrtandra cyaneoides, Clermontia montis- ae C. Waimeae, Cyanea
communis, Rollandia puirweerdisfolia; R. truncata, spp. ., and five new varieties.
Rusby, H.H. Recent botanical collecting in the Republic of Colombia.
Mem. Torrey Club 17: 39-47. 10 Je 1918.
Ruthhoven, A. G. A biological survey of the sand dune region of the
south shore of Saginaw Bay, Michigan. Michigan Geol. & Biol.
Surv. Publ. 4: (Biol. 2) 1-347. pl. 1-15 + map. 1911.
Scott, J. G. Early horticultural journalism in the United States.
Mem. Torrey Club 17: 79-81. 10 Je 1918.
Seaver, F. J., & Horne, W. T. Life-history studies in Sclerotinia.
Mem. Torrey Club 17: 202-206. pl. 3. 10 Je 1918.
Sclerotinia Geranii sp. nov. is described.
Shaw, J.K. A study of variation in apples. Massachusetts Agr. Exp.
Sta. Bull. 149: 21-36. Ap ro14.
Shufeldt, R. W. Flowers of late spring and early summer. Am.
Forestry 24: 289-294. f. 1-10. My 1918
Spegazzini, C. Sobre algunos Hongos Chilenos. Rev. Chilena Hist.
Nat. 21: 79-81. 30 Je 1917.
Spegazzini, C. Uredinaceas nuevas chilenas. Rev. Chilena Hist.
Nat. 14: 139, 140. I9gI0.
Spoehr, H. A. The pentose sugars in plant metabolism. Plant World
20: 365-379. D 1917.
Steil, W. N. Method for staining antherozoid of fern. Bot. Gaz. 68:
562, 563. f. r. 18 Je 1918
Steinberg, R. A. A study of some factors influencing the stimulative
action of zinc sulphate on the growth of Aspergillus niger. I.
effect of the presence of zinc in the cultural flasks. Mem. Torrey
Club 17: 287-293. 10 Je 1918.
Stiles, W., & Jérgensen,I. Quantitative measurement of permeability.
Bot. Gaz. 65: 526-534. 18 Je 1918.
PLATE 9
VOLUME 45,
TORREY CLUB
BULL,
COUTANT: WOUND PERIDERM IN CACTI
Vol. 45 i No. 10
BULLETIN
OF THE
TORREY BOTANICAL CLUB
OCTOBER, 1918
Regeneration in Phegopteris polypodioides*
ELIZABETH WUIST BROWN
(WITH THREE TEXT FIGURES)
INTRODUCTION
The fact that plants are able to reproduce lost parts was
known long before it was discovered that animals possessed this
same power. It was natural that the first experimental investi-
gation on regeneration in plants should have been carried on with
the higher plants as it was a common practise to propagate many
plants by means of cuttings. However, the study of regeneration
has since been extended to include not only the lower groups of
plants, such as the algae, fungi, liverworts, mosses and ferns, but
also many groups of animals. This has resulted in the accumu-
lation of a large amount of evidence regarding the possibilities of
regeneration by most groups of organisms.
Experimental evidence has also indicated that the regenerative
power of some plants is much greater in earlier than later life,
while in others this power is lost completely in later life.
Goebel (2, pp. 196-203), experimenting with ferns, found that
the primary leaves of the young sporophytes of some ferns, either
while attached to the sporophyte or cut off and placed under
moist conditions, were able to regenerate new fern plants or
prothallia or intermediate forms between leaves and prothallia.
Here the regenerative power seemed confined to the primary
* Contribution from the Osborn Botanical Laboratory.
{The BULLETIN for September (45: 353-360. #1. 9) was issued September 20, 1918.]
391
392 BRowN: REGENERATION IN PHEGOPTERIS POLYPODIOIDES
leaves of the young sporophytes, as in no case did regeneration
take place with leaves of the older plants. It was thought of
interest to see if the primary leaves of the young sporophytes of
Phegopteris polypodioides Fée could regenerate in the manner
indicated by Goebel, as apogamy had occurred so frequently in
cultures of this fern.
EXPERIMENTAL
Spores of Phegopteris polypodioides were obtained from Brook-
lin, Maine, through the kindness of Dr. A. H. Graves. Cultures
were started in the early part of October. The spores were sown
on Prantl’sand Knop’s full nutrient solutions and modifications of
these solutions. After the spores were sown the cultures were
placed before an east window. In an effort to induce apogamy the
prothallia were not transferred to fresh nutrient solutions from
time to time, but were allowed to develop upon the same nutrient
solutions upon which the spores had been sown. As a result
growth and development of the prothallia was slower and fewer
sporophytes formed, the majority of which were apogamous.*
The primary leaves of both normal and apogamous young sporo-
phytes were used in the regeneration experiments.
March 14, 1917, primary leaves 5-7 mm. in length were cut
from the young sporophytes and placed on sand in watch glasses.
In some cases the petiole of the leaf was inserted in the sand to
a depth of 1-2 mm., placing the blade of the leaf in an upright
position. In other cases the leaf was laid on the sand; while in
still others various parts of the blades were covered with the sand.
The sand in some of the watch glasses was moistened with Knop’s
and Prantl’s full nutrient solutions, while in the others it was
moistened with distilled water. All the cultures were placed in
large plates and covered with bell jars. The sand was never al-
lowed to dry and water was kept standing in the plates. In this
way the air under the bell jar was always moist.
No experiments were tried to see if the leaves would regenerate
while attached to the young sporophytes and no such cases were
observed among either the solution or soil cultures, although fre-
quently the leaves of this and other species of ferns, especially in
*Wuist, Elizabeth Dorothy. Apogamy in Phegopteris polypodioides Fée, Osmunda
cinnamomea L., and O. Claytoniana L. Bot. Gaz. 64: 435. 109
BROWN: REGENERATION IN PHEGOPTERIS POLYPODIOIDES 393
the soil cultures became appressed to the soil as a result of acci-
rents in watering.
Only one case of regeneration was obtained, although a large
number of leaves were experimented with. Regeneration began
in about six weeks after the leaf had been removed from the sporo-
phyte and laid on sand moistened with Knop’s full nutrient solu-
tion. One side of the petiole, near its base, was destroyed by
decay, and a short distance above this point on the opposite side
of the petiole a slight swelling occurred, from which a cellular mass
developed. At first it was slightly elongated, Fic. 1, afterwards
it became much thickened and broadened, Fic. 2, and finally as-
sumed the shape shown in Fic. 3. From this cellular mass there
developed two intermediate structures between leaves and pro-
thallia, then rhizoids and four normal leaves. Neither a true root
nor a stem “‘ Anlage’’ was formed.
The first of the intermediate structures resembled a very much
elongated prothallium, one cell in thickness, with an expanded
heart-shaped apex. Tracheids arranged in rows resembling a true
midrib extended up through the portion of the structure corres-
ponding to the petiole into the expanded or blade-like part. Here
the rows of tracheids branched dichotomously one branch going
to each lobe (Fics. 1 and 2). The margins of both the elongated
and expanded regions were for the most part smooth, with the
exception of two papillae, one of which developed on the elongated
and one on the expanded part.
The second intermediate structure which developed from the
cellular mass was even more thalloid in form and structure than
the first one. Both the elongated and broadened portions like
those of the first were only one cell in thickness. The expanded
portion differed somewhat in appearance as it was not so distinctly
heart-shaped and its margin bore many more papillae. There
were no indications of tracheids present in either of the portions
representing petiole or blade.
Growth was rapid and apparently normal in both of these in-
termediate structures, but the second one never attained the size
of the first. Neither was long-lived, death occurring soon after
the normal leaves began to develop. The rhizoids resembled in
every way those of a normal fern gametophyte. They formed from
394 BROWN: REGENERATION IN PHEGOPTERIS POLYPODIOIDES
the cells of both the upper and lower surfaces of the cellular mass,
although the majority came from the upper surface cells. Their
ag 0 oO,
Sereno
Men HID
3
Fics. 1-3. Stages in the regeneration of the young leaf of a sporophyte of
Phegopteris polypodioides Fée,x 550. R, rhizoids; LS, leaf of sporophyte; CM, cellular
mass , intermediate structure; MR, midrib; T, tracheids; S, stomata; rst L, first
leaf; 2d L, second leaf; 3d L, third leaf; 4th L, fourth leaf.
development began at about the same time as that of the inter-
mediate structures.
After a number of rhizoids had been formed from the surface
cells of the mass, normal leaves began to develop. The first and
second of the four leaves appeared almost simultaneously and
¥
BROWN: REGENERATION IN PHEGOPTERIS POLYPODIOIDES 395
their growth was very rapid. They resembled, in all respects, the
leaves of a normal young sporophyte of this species of fern (Fic.
3); although the blade of the second leaf was somewhat simpler
in form, having only two main divisions, each of which was lobed,
instead of three main lobed divisions. However, such irregulari-
ties are to be noted in leaves of normal young sporophytes of this
and other species of ferns. These two leaves attained a height of
6 mm. The third leaf, which was much slower in growth and
development reached a height of only 4 mm. and was much
simpler in form. The blade consisted of only two lobes, which
were entire. The fourth leaf showed still greater simplicity in
form and reached a height of only 3 mm. The blade was
almost entire, with a slight lobe on the one side. The blades of
these latter leaves, like those of the first and second ones, bore
stomata.
DISCUSSION AND CONCLUSION
The various theories which account for regeneration, as ad-
vanced by the many writers on the subject, consider the following
influences: (1) external influences to which the plant is subjected;
(2) tendencies inherent in the plant body.
The factors considered as the possible controlling or influencing
ones may be placed in the following classes: (1) nutrition dis-
turbances; (2) wound stimuli; (3) changes in the water content;
(4) the accumulation, at certain places, of definite formation
substances; (5) the presence of dormant or latent rudiments;
(6) correlation; (7) age and maturity of the parts; (8) form
disturbance; (9) growth tension; (10) interruption of the func-
tions of respiration, transpiration or photosynthesis; (11) iso-
lation from the influence of the whole; (12) presence of enzymes
which are responsible for the formation of the part regenerated.
The fact that regeneration was never observed in attached
leaves of Phegopteris polypodioides and did occur in a leaf which
had been separated from the plant would seem to indicate that the
separation from the repressing influence of the plant body played
an important part, as believed by Loeb (4, p. 153); although the
fact that cases have occurred in other species of ferns when the
leaves were still attached would not seem to refute this theory.
Experimental evidence has shown that many species of ferns,
396 BROWN: REGENERATION IN PHEGOPTERIS POLYPODIOIDES
either in the younger or older stages of their gametophytic or
sporophytic life history and especially in the younger stages, does
not always react in the same way to the same environmental con-
ditions; but each has a sort of individualism which enables it to
react in the way best fitted for its particular needs. It seems very
probable that in this particular case regeneration is closely con-
nected with nutrition for the reason that the severed leaf was not
able to regenerate immediately another sporophyte, but could
produce a cellular mass which appeared to be a reversion to a
prothallus-like structure. This structure after rhizoids developed
gave rise, in an apogamous manner, first to structures intermediate
between leaves and prothallia; then, as the number of rhizoids in-
creased, enlarging the absorptive surface, which in turn increased
the amount of nourishment, true leaves were produced. The
first cf these were the ordinary type of young sporophytic leaf but
later ones were much more primitive in character, due doubtless
to the lowered vitality of the prothallus-like structure. This
coincides with Goebel’s (1, vol. 2, Pp. 42) views. He considers a
reversion to a juvenile form as being the result of unfavorable con-
ditions to which the plant is subjected. It is also in keeping with
the results obtained by Miss Kupfer (3, p. 229) in her experi-
ments, which indicated that at the time of cutting under normal
conditions reserve food was present in sufficient quantities to
initiate the first stages of regeneration. If this food was absent
and its formation prevented regeneration was prohibited.
Such an explanation as this does not lose sight of the impor-
tance of the other factors so strongly emphasized by various
workers, for all these factors doubtless play an important part in
nutrition.
Experimental data in this particular case are not extensive
enough to permit drawing conclusions as to which of the above
factors or groups of factors acting separately or together, was the
determining one. However, since an intimate relation exists be-
tween growth and regeneration and since growth is dependent
upon nutrition, it would seem as if some phase of nutrition must
be an important factor in regeneration,
if not the most important
factor.
Brown: REGENERATION IN PHEGOPTERIS POLYPODIOIDES 397
SUMMARY
1. Regeneration took place near the base of the petiole of a
detached leaf of a young sporophyte of Phegopteris polypodioides,
placed upon sand moistened with Knop’s solution in moist air.
2. A cellular mass, resembling a prothallium, was formed, from
which rhizoids, intermediate structures between leaves and pro-
thallia, and true leaves developed.
3. At first true leaves resembling those of normal young sporo-
phytes were formed; then leaves of a much simpler type developed.
LITERATURE CITED
~_
. Goebel, K. Organography of plants. English translation. Oxford.
1900
2, —_———. Einleitung in die experimentelle Morphologie der
Pflanzen. Leipzig. 1908.
Kupfer, E. Studies in plant regeneration. Mem. Torrey Club 12:
195-241. f. 1-3, 1907.
4. Loeb, J. The organism as a whole. New York. 1916.
es
Taxonomy and distribution of Adenostegia
ROXANA STINCHFIELD FERRIS
(WITH PLATES 10-12)
The genus Adenostegia was first described by Bentham* with
the description of one species, A. rigida. Ten years later the same
author published in De Candolle’s Prodromus{ four Nuttallian
species, substituting Nuttall’s manuscript name, Cordylanthus.
This change was made on the ground that the meaning of the word
Cordylanthus (cordule, club; anthos, flower) was more character-
istic of the new species than was Adenostegia (aden, gland; stege,
covering). The proper generic name, according to present-day
rules of nomenclature, was revived by Greene in 1891, and later
accepted by Kuntze § and Wettstein. ||
The affinities of Adenostegia are with Castilleia and Orthocarpus,
and in Wettstein’s generic arrangement of the Scrophulariaceae
it is placed between them. The most noticeable likeness to
Castilleia is shown in the section A nisocheila in the elongated upper
lip of the corolla. A much more marked connection is found
between the section Chloropyron and Orthocarpus, for here there
are points of resemblance not only in the distinctly three-saccate
flower but in the spike-like inflorescence as well.
The sections Euadenostegia and Chloropyron of Adenostegia
exhibit the conspicuous degree of variability that is characteristic
of many of the Scrophulariaceae. In Euadenostegia there are,
besides the fixed species, two plastic groups, rigida and pilosa, in
which the range of intraspecific variation creates difficulties in
defining the species. The species of the rigida group are char-
acterized by their hispid or hirsute pubescence and their tri-
* In Lindley, J. A natural system of botany. Ed. 2. 445. 1836.
+ Prodr. Syst. Nat. 10: 507, 598. 1846.
t Some neglected priorities in generic nomenclature. Pittonia 2: 180-181.
§ Rev. Gen. Plant. 2: 456, 457- 1891.
|| In Engler & Prantl, Nat. Pflanzenfam. 4: 98. 1891.
39
400 FERRIS: TAXONOMY AND
partite, callous bracts. The species of the pilosa group have a
puberulent or pilose pubescence and their bracts are typically
linear, though they are parted in A. viscida and A. Hanseni.
The genus is confined to western America and is characteristic
of California and the Great Basin. Of the twenty-one species,
there are but five that are not recorded from California and nine
are known only from that state. According to the present records
the range of the genus extends from Washington and southwestern
Montana to Sonora, Chihuahua, and northern Lower California.
The members of the genus are found principally in open,
exposed places in the Upper and Lower Sonoran and the Arid
Transition Zones. Certain of the species may at times be met
with in the Humid Transition Zone, but they occur only on
exposed slopes that are truly “islands” of Upper Sonoran. All
the species of the section Chloropyron, which is in part coastal,
are found in salt marshes and on alkaline soils. This unusual
habitat may account for the fact that this group appears not to
conform to the ordinary zonal lines.
In preparing this paper I have had the opportunity of examin-
ing the material in the herbaria of the following institutions: the
National Herbarium, the University of California and Stanford
University.
I wish to express appreciation of the kindness shown me at the
University of California during my work in that herbarium and
to the National Herbarium for the loan of material. I also wish
to thank Dr. L. R. Abrams for his advice and assistance and Dr.
B. L. Robinson, of the Gray Herbarium, for fragments and
photographs of types.
ADENOSTEGIA Benth.
Adenostegia Benth. in Lindley, Nat. Syst. ed. 2. 445. 1836.
Cordylanthus Nutt.; Bentham in De Candolle, Prodr. ro: 597-
1846.
Chloropyron Behr, Proc. Calif. Acad. 1: 6r. 1855.
Rigid, summer-blooming annuals with divaricate or paniculate
branches and yellow roots. Leaves alternate, entire, parted or
dissected, those subtending the branches much longer than the
others, deciduous with age; flowers in spikes, heads, or scattered
along the branches; floral bracts entire, dissected, or parted;
DISTRIBUTION OF ADENOSTEGIA 401
calyx one- to two-leaved, lower leaf, when present, saccate at the
base, upper leaf erect, winged at the base except in the section
Chloropyron; corolla cylindrical, bilabiate, the lips nearly equal
except in A. laxiflora; upper lip enclosing stamens and pistil,
lower lip obscurely three-crenulate or entire, more or less saccate;
stamens two or four, in unequal pairs, the anthers one- or two-
celled, the lower cell, when perfect, subtending the upper, filaments
generally hairy; capsule flattened, lanceolate or slightly rounded;
seed irregular, more or less reticulate.
Key to sections
Lower lip of the corolla one-half the length of the upper lip. I. ANISOCHEILA. —
Lower lip of corolla equalling upper lip.
Tube of the corolla one fifth the length of the throat. If, PRINGLEA.
Tube of corolla more than one fifth the length of the throat.
Calyx diphyllous. III. EUADENOSTEGIA,
Calyx-leaf cxnalliie corolla.
Inflorescence capitate. IV. KINcIA,
Inflorescence spicate. VI. CHLOROPYRON
Calyx leaf one half the length of the corolla. V. Dicemnerik.
Section I. ANISOCHEILA
Cordylanthus § Adenostegia (in part) Gray, Proc. Am. Acad. 7:
381. 1868; Wats. Bot. King’s Exped. 459.
Cordylanthus § Anisocheila Gray, Syn. Fl. 2': 303. 1886.
Adenostegia § Anisocheila Wettstein in Engler & Prantl, Nat.
Pflanzenfam. 4: 98. 1891.
Flowers scattered along the stems; calyx diphyllous, upper lip
of corolla twice as long as lower; anthers one-celled or with vestige
of second cell.
1. ADENOSTEGIA LAXIFLORA (Gray) Greene
Cordylanthus laxiflorus Gray, Bot. Mex. Bound. 2: 120. 1859;
Gray, Proc. Am. Acad. 7: - Wats. Bot. King’s Exped.
232, 460; Gray, Syn. FI. 2!:
Adenostegia laxiflora Greene, ba 2: 181. 1891; Kuntze,
Rev. Gen. 2: 457; Wettstein in Engler & Prantl, Nat.
Pflanzenfam. 4%: 98.
Paniculately branching annual, 3-4 dm. high, pubescent
throughout with soft, glandular hairs; leaves 8-13 mm. long,
one- to three-parted; flowers many, solitary or in groups of two to
four; floral bract one half the length of calyx, deeply three-parted
402 FERRIS: TAXONOMY AND
into linear divisions rounded at the apex; upper calyx-leaf 11-16
mm. long, six- or seven-nerved, winged at the base, lanceolate or
bidentate, pubescence sparse; lower calyx-leaf 10-15 mm. long,
four-nerved, wider than the upper, rounded at the apex; corolla
16-18 mm. long, glabrous, the lower lip saccate; filaments gla-
brous; capsule 7-8 mm. long, slightly rounded; seeds reticulate.
[PLATE 10, FIG. I; PLATE I1, FIG. 1.]
Tyre LocaLity: ‘‘Rocky hills, Sonora, Mexico, Thurber.”’
DiIstRIBUTION: Hills and ravines in Arizona and northern
Sonora.
SPECIMENS EXAMINED :—ArIzoONA: “‘ Point of Mountain,” Roth-
rock 721; Pine Creek near Pine, MacDougal 693; hills, Beaver
Creek, Purpus 8208.
Section II. -PRINGLEA
_ Flowers in heads; calyx diphyllous; tube of corolla short and
dilated; stamens four, perfect, the filaments villous.
2. ADENOSTEGIA PRINGLEI (Gray) Greene
Cordylanthus Pringlei Gray, Proc. Am. Acad. 19: 94. 1883;
Gray, Syn. Fl. 2!: 453; Jepson, Fl. W. Mid. Calif. ed. 2. 387.
Adenostegia Pringlei Greene, Pittonia 2: 18t. 1891; Kuntze,
Rev. Gen. 2: 457; Jepson, Erythea 7: 112; Jepson, Fl. W.
Mid. Calif. 416.
Slender, glabrous annual, 4-6 dm. high; leaves 5—9 mm. long,
linear-filiform, early deciduous, the lower sparsely pubescent, the
upper glabrous; heads many, compact, three- to five-flowered;
floral bracts 5-7 mm. long, glabrous, with five to seven equal lobes;
calyx-leaves 9-11 mm. long, nearly equal, upper part covered
with large, tack-like glands; corolla 8-9 mm. long, lips densely
hairy; capsule rounded. [PLATE 10, FIG. 2; PLATE 40. 716.3.)
TYPE LocaLity: “California, on dry hills in Lake Co., August,
1882, Pringle.”
DISTRIBUTION: Exposed slopes in the mountains of Lake and
Napa Counties, California; Upper Sonoran Zone.
SPECIMENS EXAMINED:—CatirorNia: Lake County, August,
1882, Pringle; Snow Mountain, Lake County, August 25, 1892,
Katharine Brandegee; Cobb Mountain, Lake County, 1910, Kath-
arine Brandegee; near Bartlett Springs, Lake County, August,
1916, A. Stinchfield.
DISTRIBUTION OF ADENOSTEGIA 403
Section III. EUADENOSTEGIA
Cordylanthus § Adenostegia (in part) Gray, Proc. Am. Acad. 7: 381.
1868; Wats. Bot. King’s Exped. 459; Gray, Bot. Calif. 1: 580;
Gray, Syn. Fl. 2!: 303.
Adenostegia § Euadenostegia Wettstein in Engler & Prantl, Nat.
Pflanzenfam. 4%: 98. 1891.
Flowers scattered along the branches or in heads; calyx
Ape corolla lips equal; anthers one- or two-celled, filaments
illous
Key to species of § Euadenostegia
THAMENS CWO E.G i cn Sel ore fe ee ees 3. A. capitata.
Stamens four.
Antherpione-celled of 8/05. See a ae 9. A. Nevinit.
Anthers two-celled.
Bracts entire or with the callous tips notched.
e row cinereous pilose. ......... aijA.
Her puberulent with few pilose hairs. . SA
Bracts cin parted.
B s three-parted.
Ficlets scattered along the branches;
age pilose.
Tips of bracts enlarged or calloused.
Flowers 14 mm. or less long; middle
lobe of bract exceeding lateral
sees Cae Ree es EG 6. A. viscida,
Flowers mm. or more long;
lobes e pie nearly equal. 7. A. Hanseni.
Res of bracts not enlarged or eal:
8. A. parviflora.
ote ie eee at te de ge tt eee ae a eS eae
lou
F mies in ae. herbage hispid or hir-
ute. >
hese parted nearly to base; pubes-
10. A. filifolia.
Pubescence sparsely hispid ... 11. A. rigida.
t 11a, A poe brevibracteata.
12. A. litioralis.
Bracts five- to seven-
Corolla 1 s-1y- mmm. ones: 5665 ok eae ke 13. A. ramosa.
Corolla 39-306 mim: long. oi. 35 se ee 14. A. Wrightit.
404 FERRIS: TAXONOMY AND
3. ADENOSTEGIA CAPITATA (Nutt.) Greene
Cordylanthus capitatus Nutt.; Bentham in De Candolle, Prodr.
10: 597: 1846; Gray, Proc. Am. Acad. 7: 382; Wats. Bot.
King’s Exped. 231, 459; Gray, Bot. Calif. 1: 580; Gray, Syn.
FI.:2*: 304;
Adenostegia capitata Greene, Pittonia 2: 180. 1891; Kuntze,
Rev. Gen. 2: 457; Howell, Fl. N. W. Am. 537; Piper, Contr.
U.S. Nat. Herb. 11: 518; Rydberg, Bull. Torrey Club go:
484. 1913.
Adenostegia ciliosa Rydb. Bull. Torrey Club 34: 35. 1907.
Cordylanthus bicolor A. Nels. Bot. Gaz. 54: 416. 1912.
Paniculately branching annual, 4-6 dm. high, short-pilose
throughout with glandular hairs: leaves many, 2-5 cm. long,
linear or three-parted; heads two- to five-flowered; flowers spread-
ing, giving an open appearance to the head; floral bracts 8-10
mm. long, three-parted, the divisions linear-lanceolate, spreading,
the middle division twice as long as the lateral divisions; calyx-
leaves purplish, the upper 8-9 mm. long, thin, two-nerved, two-
toothed, the teeth 2-3 mm. long, the lower 10-13 mm. long,
five-nerved, broad at the apex and curved outward; corolla 10-12
mm. long, covered with reflexed hairs, the tube longer than the
throat, purple, tipped with yellow; stamens two, the anthers one-
celled with vestiges of a second cell, the filaments nearly glabrous,
with a U-shaped curve near the anther; capsule 5-6 mm. long,
slender, pointed; seeds few, reticulate. [PLATE 10, FIG. 3.]
Co-types of Cordylanthus bicolor examined by the author are
not distinct from A. capitata. Specimens of A. ciliosa have not
been examined, but Rydberg (Bull. Torrey Club 40: 484. 1913)
states that his species is identical with Nelson’s.
TYPE LocaLity: “In Nova California (Nuttall!).”
DISTRIBUTION: Mountain ranges of Washington and Idaho to
Lassen County, California; Arid Transition Zone.
SPECIMENS EXAMINED:—WASHINGTON: Yakima Region, 1882,
Brandegee; Falcon Valley, Suksdorf 201. Ipano: Redfish Lake,
Evermann 408; Blaine County, Nelson & MacBride 1239; Pine-
hurst, Boise County, Macbride 1671. OrEGon: Powder River,
Cusick 1784; same locality, Piper 2482; Hepburn Ridge, Wal-
lowa County, Sheldon 8643; same locality, Howell. CatirorNIA:
hills near Lassen Creek, Mrs. Austin 146. NeEvapa: .Coleman
Valley, Coville & Leiberg 95; Clover Mountains, Watson 816;
Gold Creek, Elko County, Kennedy 4282.
DISTRIBUTION OF ADENOSTEGIA 405
4. ADENOSTEGIA PILOSA (Gray) Greene
Cordylanthus pilosus Gray, Proc. Am. Acad. 7: 382. 1868; Wats.
Bot. King’s Exped. 459; Gray, Bot. Calif. 1: 581; Gray, Syn.
Fl. 21: 304; Jepson, Fl. W. Mid. Calif. ed. 2. 387.
Adenostegia pilosa Greene, Pittonia 2: 180. 1891; Kuntze, Rev.
Gen. 2: 456; Jepson, Fl. W. Mid. Calif. 416.
Stout, paniculately branching annual, 6-12 dm. high; stems
somewhat reddish, glandular with rather short, pilose hairs; leaves
10-20 mm. long, truncate and often callous-emarginate at apex;
flowers scattered along the branches; floral bracts linear, 15-25
long, cinereous-pilose, the callous tip dilated, sometimes
three-notched, three-nerved; upper calyx-leaf 20-22 mm. long,
shallowly bidentate, three-nerved, pubescence sparse, the lower
calyx-leaf 19-21 mm. long, broadly lanceolate, with pubescence
as on the bracts; corolla 15-18 mm. long, the tube shorter than
the throat, greenish white with reddish-purple markings at base
of throat and on lower lip; stamens four, perfect, the filaments
villous; capsule not sharply pointed; seeds few, slightly reticu-
late. [PLATE 10, FIG. 4.]
Type LocaLity: ‘‘Dry soil near San Jose.”’
DistRIBUTION: Interior valleys and foothills of California,
from Trinity and Mendocino Counties southward to Santa Clara
County; Upper Sonoran Zone. The specimens from the higher
elevations in Lake and Mendocino Counties are less pilose but do
not differ structurally from the typical form.
SPECIMENS EXAMINED:—CALIFORNIA: Biréka = Red: Bluff
Road, Trinity County, Abrams 6158; between Harris and Alder
Point, Trinity County, Abrams 5964; near summit of Mt.Sanhedrin,
Lake County, Heller 5997; Cobb Mountain, Lake County, August
13, 1910, Katharine Brandegee, near Bartlett Springs, Lake County,
August, 1911, A. Stinchfield; Princeton, Colusa County, October,
1905, Chandler; St. Helena Sanitarium, Napa County, Abrams
5752; foothills, Yolo County, Stinchfield 345; Petrified Forest,
Sonoma County, Bioletti & Michener 1731a; Weldon Cajon, Vaca
Mountains, Salona County, September 13, 1891, Jepson; near San
Jose, Santa Clara County, Brewer; foothills west of Los Gatos,
Santa Clara County, Heller 7532; same locality, Eastwood; Palo
Alto, Baker 1714; Stevens Creek Road, Santa Clara County,
Stinchfield 247; near Stanford University, Stinchfield 255; Ray-
mond Ranch, Santa Cruz Mountains, August 14, 1911, Blasdale;
406 FERRIS: TAXONOMY AND
near Saratoga, Pendleton 270; near Colley’s Landing, Palo Alto,
September 29, 1906, Abrams.
5. ADENOSTEGIA TENUIS (Gray) Greene
Cordylanthus tenuis Gray, Proc. Am. Acad. 7: 383. 1868; Wats.
Bot. King’s Exped. 232, 460; Gray, Bot. Calif. 1: 581; Gray,
Syn. Fl. 2': 304; Hall, Yosemite Flora 229.
Cordylanthus pilosus var. Bolanderi Gray, Proc. Am. Acad. 7: 382.
1868; Gray, Bot. Calif. 1: 581; Gray, Syn. FI. 2!: 304.
Adenostegia tenuis Greene, Pittonia 2: 180. 1891; Kuntze, Rev.
Gen. 2: 456; Wettstein in Engler & Prantl, Nat. Planzenfam.
a**: G8.
Adenostegia pilosa var. Bolanderi Greene, Pittonia 2: 180. 1891.
Adenostegia Bolanderi Kuntze, Rev. Gen. 2: 456. 1891.
Slender, paniculately branching annual, 2-6 dm. high; stems
puberulent with short, scattered, glandular, pilose hairs, sometimes
glabrous; leaves 1-3 cm. long linear, occasionally with callous tips;
flowers scattered along the branches; floral bracts 12-20 mm. long,
puberulent, ciliate with glandular, pilose hairs, linear-lanceolate,
entirely herbaceous or with callous tips; calyx-leaves with pubes-
cence as on bracts, the upper 15-18 mm. long, three-nerved,
lanceolate, sometimes bidentate, the lower 14-18 mm. long, five-
nerved, broadly lanceolate; corolla 12—1 5 mm. long, the tube
about equalling the throat, inconspicuously hairy; capsule 8 mm.
long; seeds few, somewhat reticulated. [PLATE 10, FIG. 5.]
The glabrous or puberulent type of A. tenuis is found in the
Lake Tahoe region and in western Nevada, while the form de-
scribed as A. pilosa var. Bolanderi is characteristic of the mountain
ranges north of this region. This last-named form was described
by Gray as being more nearly related to A. pilosa, but an examina-
tion of co-type material shows that the bracts are not notched as
they are in A. pilosa and that in the pubescence it is like A. tenuis
except for the presence of scattered, pilose hairs.
The specimens from Lake County are like the Nevada form,
_ while those from the Kings River region more closely resemble A.
pilosa. j
TYPE LocaLity: “Dry sandy soil near Lake Tahoe, Nevada,
Brewer, Dr. C. L. Anderson.”’
DIsTRIBUTION: Exposed slopes in Lake County in the Coast
DISTRIBUTION OF ADENOSTEGIA 407
Range and in the Sierra Nevada Mountains from Lassen County
to Fresno County, California, and in western N evada; Transition
and Canadian Zones.
SPECIMENS EXAMINED:—CALIFORNIA: Mt. Hannah, Lake
County, Tracy 3247; Susanvillé Summit, Lassen County, July 2,
1897, Jones; Grizzly Hill, Plumas County, Leiberg 5207; Clio,
Plumas County, Eggleston 6240; Sierra Valley Hot Springs, Sierra
County, August, 1909, Dudley; Soda Springs, Nevada County,
Jones 2580; Susie Lake Trail, Eldorado County, McGregor 125;
Lake Tahoe, Eldorado County, Leiberg 5 327; Agricultrual Sta-
tion, Amador County, Hansen 697; between Big Trees and Gard-
ner's, Calaveras County, August, 1906, Dudley; near Mariposa
Big Trees, Mariposa County, Abrams 5397; same locality, August
II, 1895, Congdon; Mariposa Big Tree Grove, Bolander 4903 (co-
type of A. pilosa var. Bolanderi); Yosemite National Park, Glacier
Point, Hall 9202; Tamarack Flat, Abrams 5474; near foot of
Yosemite Falls, Abrams 5 466; trail between Illillouette and Glacier
Point, Abrams 5430; Converse Basin, Fresno County, October,
1900, Dudley; Kings River Region, Fresno County, August, 1904,
Dudley; ‘west of Bearskin Meadow, August, 1904, Dudley; same
locality, Hall & Chandler 179. NEVADA: near Lake Tahoe,
Brewer 2150; Clear Lake Cafion, Ormsby County, Baker 1408;
mountains west of Bowers, Washoe County, Heller 10663; Galena
Creek, Washoe County, Heller 10671; Lake Tahoe, Washoe
County, Kennedy 1458.
6. ADENOSTEGIA VISCIDA Howell
Adenostegia viscida Howell, Fl. N. W. Am. 537. 1903.
Paniculately branching annual, 2-4 dm. high; short pilose
throughout with viscid-glandular hairs; leaves 18-25 mm. long,
linear-lanceolate, occasionally three-parted; flowers scattered
along the branches, solitary or in groups of three or four; bracts
0-14 mm. long, slender, three-parted, enlarged and slightly
calloused at the apex; calyx-leaves 15-16 mm. long, the upper the
longer, four-nerved, the lower five-nerved; corolla 12-14 mm. long;
stamens four, perfect, the filaments villous; capsule 6-7 mm. long.
[PLATE 10, FIG. 6.]
A. viscida in Shasta and Plumas Counties is inconspicuously
glandular-pilose as compared with the typical form and approaches
A. tenuis, to which this species is very closely related.
408 FERRIS: TAXONOMY AND
TYPE LocaLity: “On dry slopes, eastern base of the Coast
Mountains, near Waldo, Southern Oregon.”’
DISTRIBUTION: Siskiyou Mountains, Oregon, south to Plumas
County, California.
SPECIMENS EXAMINED :—OREGON: Illinois River in the Siskiyou
Mountains, Cusick 2937; same locality, July, 1887, Howell; near
Fort Klamath, Letberg 636; Klamath County, Mrs. Austin &
Bruce 1773; Rogue River, Brackenridge 1192; Upper Metolins
River, Crook County, Coville & Applegate 700; Wimer, Jack-
son County, Hammond 318. Catirornia: Mt. Eddy, Siskiyou
County, Heller 11744; Weed, Heller 11723; Burney Falls, Shasta
County, August, 1899, Dudley; Bear Valley Mountains, Shasta
County, June and August, 1893, Baker; Big Meadows, Plumas
County, Mrs. Bruce 411; Lassen Buttes, Plumas County, Brown
637; Grizzly Hill, Plumas County, Leiberg 5107.
7. Adenostegia Hanseni sp. nov.
Stout, paniculately branching annual, 5-12 dm. high; stem
reddish, glandular throughout with long, cinereous, pilose hairs
which are often 4-5 mm. long; leaves 12-30 mm. long, truncate
and often calloused at the apex; flowers scattered along the
branches; floral bracts 15-30 mm. long, 3-nerved, 3-parted, the
divisions 4-7 mm. long, nearly equal, enlarged and callous-
emarginate at apex; calyx leaves 15-22 mm. long, the upper
lanceolate, the lower rounded; corolla 16-18 mm. long, the tube
nearly equalling the throat; stamens 4, perfect, the filaments
villous; capsule 7-8 mm. long. [PLATE 10, FIG. 7: PLATE iT;
FIG. 3.] |
This species closely resembles A. pilosa in habit, but is readily
distinguishable by the tripartite bract and the long, pilose hairs.
TYPE Locatity: Agricultural Station, Amador County, Cali-
fornia; alt. 2,000 ft., September, 1893, Hansen 138. Tyrer, No.
21142 of the Dudley Herbarium.
DIsTRIBUTION: Foothills of the Sierra Nevada from Shasta
County to Tuolumne County, California; Upper Sonoran Zone.
SPECIMENS EXAMINED :—CaLIForRNIA: north of Redding, Shasta
County, Heller 12501; between Bellavista and Ingot, Shasta
County, July, 1914, McMurphy; four miles east of Placerville,
Tehama County, Heller 12551; east of Oroville, Butte County,
Heller 11569; Agricultural Station, Amador County, alt. 2,000 ft.,
DISTRIBUTION OF ADENOSTEGIA 409
Hansen 138; near Ione, Amador County, Braunton 121; Wards
Ferry, alt. 1,500 ft., Tuolumne County, Abrams 4717; French
Flat, near Rawhide, Tuolumne County, Stinchfield 64; plains of
the Sacramento, Pickering 1348.
8. Adenostegia parviflora sp. nov.
Divaricately branching annual, 3 dm. high, puberulent and
minutely viscid-pilose throughout; leaves 10-15 mm. long;
flowers many, scattered along the branches; floral bracts 5-6 mm.
long, three-parted into linear divisions with rounded, herbaceous
tips; calyx leaves 11-13 mm. long, the lower slightly exceeding
the upper; corolla 10-11 mm. long, tube nearly equalling throat;
stamens 4, perfect, the filaments villous; capsule lanceolate.
[PLATE 10, FIG. 8; PLATE II, FIG. 4.]
Type Locatity: Grand Canyon of the Colorado River, near
the San Francisco Mountains, Arizona, Knowlton 270. TYPE,
No. 48859 of the U. S. National Herbarium.
9. ADENOSTEGIA NEVINII (Gray) Greene
Cordylanthus Nevinit Gray, Proc. Am. Acad. 17: 229. 1882;
Gray, Syn. FI. 2!: 454.
Adenostegia Nevinii Greene, Pittonia 2: 181. 1891; Kuntze, Rev.
Gen. 2: 457; Hall, Univ. Calif. Pub. Bot. 1: 114.
Slender, paniculately branching annual, 3-4 dm. high; hirsute
when young, puberulent in age; lower leaves dense, 20-25 mm.
long, three-parted into linear divisions, the upper leaves scarcely
bracts scarcely one half the length of the calyx, short pubescent,
three-parted into linear divisions with emarginate, glandular tips;
calyx leaves hispidulous throughout and with soft ciliate edges,
the upper 12-13 mm. long, 6-nerved, winged at the base, the lower
equalling the upper, 5—6-nerved, broadly lanceolate; corolla 11-12
mm. long, glandular, the tube less than one half the length of the
throat; stamens 4, the anthers 1-celled, the filaments villous;
capsule not sharply pointed. [PLATE 10, FIG. 9.]
TypE Locauity: “California, in the San Bernardino Mountains,
at about 5000 feet, Rev. J. C. Nevin, 1880; S. B. & W. F. Parish,
1881.”’
DistripuTIon: Mountain ranges of Southern California, from
410 FERRIS: TAXONOMY AND
Mt. Pinos to the Cuyamaca Mountains, San Diego County;
Arid Transition Zone.
SPECIMENS EXAMINED:—CALIFORNIA: Upper San Antonio
Cafion, Los Angeles County, Johnston 1662; Swartout Cafion, San
Gabriel Mountains, San Bernardino County, September, 1904,
G. R. Hall; Bear Valley, San Bernardino County, Abrams 2070;
same locality, Davidson, 1894; same locality, June 29, 1897,
Chandler; San Bernardino Mountains, Parish Brothers 992; San
Jacinto Mountains, Riverside County, Hall 2619; Lockwood
Valley; Ventura County, Hall 6644; Cuyamaca, San Diego
County, 1884, Orcutt; same locality, October 15, 1894, Brandegee.
10. ADENOSTEGIA FILIFOLIA (Nutt.) Abrams
Cordylanthus filifolius Nutt.; Bentham in De Candolle, Prodr. 10:
597. 1846; Wats. Bot. King’s Exped. 459 (in part); Gray,
Bot. Calif. 1: 581 (in part); Gray, Syn. Fl. 2!: 303 (in part).
Adenostegia rigida Greene, Pittonia 2: 180 (in part). 1891; Hall,
Univ. Calif. Pub. Bot. 1: 114.
Adenostegia filifolia Abrams, Fl. Los Angeles 372. 1904.
Cordylanthus rigidus var. filifolius McBride, Contrib. Gray Herb.
N.S. 49: 58. 1917.
Slender, paniculately branching annual, 3-5 dm. high, the
stems hirsute, the lower part puberulent; leaves linear or three-
parted, puberulent with scattered, hirsute hairs; mature floral
fourth the length of the bract, densely covered with long, spread-
ing, hirsute hairs, tips of the divisions enlarged, calloused, emar-
ginate, with a large gland in the emargination; calyx-leaves
13-16 mm. long, short-hirsute, the upper 4—5-nerved, rounded at
apex; corolla 12-15 mm. long, the tube shorter than the throat;
stamens four, the anthers two-celled; capsule not sharply pointed;
seeds slightly reticulate. [PLATE 10, FIG. 10.]
TYPE LocaLity: “Ad San Diego (Nuttall!).’’
DIsTRIBUTION: Dry ridges and open places from Ventura
County in California to the northern part of Lower California;
Upper and Lower Sonoran Zones.
SPECIMENS EXAMINED:—CAaLirorniA: San Felipe, Los Angeles
County, Hayes 582; Glendora, Los Angeles County, 1892, Miss
Palmer; Ramona, Los Angeles County, July, 1903, Brandegee;
DISTRIBUTION OF ADENOSTEGIA 411
Hobart, Los Angeles County, Braunton 513; Newhall, October
14, 1882, Pringle; Victorville, San Bernardino County, Parish
10523; Lytle Creek Cafion, San Bernardino County, Abrams
2745; same locality, Hall 1423; San Jacinto Mountains, Riverside
County, Hall 2621; San Diego, San Diego County, August, 1916,
McMurphy; same locality, September, 1903, Brandegee; same lo-
cality, August, 1906, Katharine Brandegee; same locality, July,
1895, Stokes; Point Loma, San Diego County, Hall 8325; Viejas
Grade to Descanso, San Diego County, July, 1906, Katharine
Brandegee; between La Mesa and Jamul, San Diego County,
Abrams 5006; Del Mar, San Diego County, August 5, 1906,
Katharine Brandegee; Witch Creek, San Diego County, Anderson
248; Tecate River, San Diego County, Shoenfeldt 3741; same lo-
cality, Mearns 3797; Laguna Mountains, Imperial County, Mc-
Gregor 97; Southwestern Colorado Desert, Imperial County, June,
1889, Orcutt. ae
11. ADENOSTEGIA RIGIDA Benth.
Adenostegia rigida Benth. in Lindley, Nat. Syst. ed. 2. 445. 1836;
Bentham in De Candolle, Prodr. 10: 537; Greene, Pittonia 2:
180 (in part); Wettstein in Engler & Prantl, Nat. Pflan-
zenfam. 42°: 98; Jepson, Fl. W. Mid. Calif. 416.
Cordylanthus filifolius Gray, Bot. Mex. Bound. 2: 120. 1859;
Gray, Proc. Am. Acad. 7: 382. Not Nuttall.
Cordylanthus rigidus Jepson, Fl. W. Mid. Calif. ed. 2. 387. I9II.
Diffusely branching annual, 3-6 dm. high; stems often reddish,
puberulent with scattered, hirsute hairs, glabrous with age; leaves
12-15 mm. long, linear-filiform, occasionally three-parted, canes-
cent-puberulent; mature heads compact, six- to twelve- rarely
fifteen-flowered; floral bracts 12-20 mm. long, sparsely hispid,
the ciliate margins interspersed with soft hairs, three-parted into
linear divisions, the apex calloused, enlarged and truncate or
slightly emarginate, the veins prominent; calyx-leaves 18-21 mm.
long, sparsely hispid, five-nerved, the lower broad, I mm. longer
than the upper; corolla 16-19 mm. long, the tube equalling or
slightly shorter than the throat; stamens four, the anthers two-
celled; capsule not sharply pointed; seeds slightly reticulate.
[PLATE 10, FIG. IT.]
Type Locauity: “Herba Novo-californica.” The exact type
locality of this species is doubtful, but of various specimens sent
to the Kew Herbarium by Dr. Abrams in 1903, one from Naci-
412 FERRIS: TAXONOMY AND
miento River, Monterey County, was said to agree most closely
with Douglas’s specimen. It is known that Douglas visited San
Antonio Mission and collected in the Santa Lucia Mountains.
So we may safely assume that the type locality of this species is
in that region.
DIsTRIBUTION: Exposed slopes in the inner Coast Ranges of
California from the Santa Cruz Mountains to the Santa Lucia
Mountains; Upper Sonoran Zone.
SPECIMENS EXAMINED:—CALIFORNIA: Crystal Springs, Santa
Cruz Peninsula, 1896, Eastwood; Permanente Creek, July, 1903,
Dudley; Castle Rock Ridge, October, 1906, Abrams; near Congress
Springs, Stinchfield 248; near Saratoga, Pendleton 234; near
Wrights, Dudley, 1894; California Redwood Park, Stinchfield 253;
same locality, Abrams 6384; Glenwood Station, August, 1900,
Davis; head of Aptos Creek, Abrams 3028; Santa Lucia Moun-
tains, Condit 8; same locality, Vasey 483; same locality, May-—
July, 1892, Vortriede; Jolon, July 30, Brandegee; Tassajara Hot
Springs, July 18, 1908, Cox.
Ila. ADENOSTEGIA RIGIDA BREVIBRACTEATA (Gray) Greene
Cordylanthus filifolius var. brevibracteatus Gray, Bot. Calif. 1: 622.
1876; Gray, Syn. FI. 2!: 304.
Adenostegia rigida var. brevibracteata Greene, Pittonia 2: 180.
: 1891; Coville, Contr. U.S. Nat. Herb. 4: 173.
Habit as in typical A. rigida; bracts with distinctly calloused,
emarginate tips, often dark colored, with prominent veins; bracts
and calyx-leaves copiously hirsute-ciliate. [PLATE 10, FIG. 12.]
The short bracts upon which Gray based the name are char-
acteristic of the specimens from Fresno and Kern Counties.
These specimens, however, have the hirsute-ciliate pubescence of
all the Sierran forms. A form growing at Santa Barbara and near
Visalia, Tulare County, has less conspicuously calloused bracts,
but seems to grade into typical brevibracteata.
I have examined the following three collections from the
Kaweah region: meadows near Monarch Lake, Dudley 1230;
Bearskin Meadow, King’s River region, August, 1904, Dudley;
Grant Forest Reservation, August, 1910, Katharine Brandegee.
These undoubtedly belong to the A. rigida group but cannot be
DISTRIBUTION OF ADENOSTEGIA 413
correctly identified because of insufficient material. The form
represented in these collections differs from A. rigida brevibracteata
in being glabrous throughout, except for the slightly ciliated
bracts, and in the shape and herbaceous character of the bracts.
TYPE Locality: ‘‘Near Soda Spring on Kern River, at 8,500
feet, Rothrock, in Wheeler’s Exped., 1875.”
DIsTRIBUTION: Mariposa County to Tulare County, California,
and also on the eastern side of the Mt. Hamilton Range; Upper
Sonoran Zone. This variety is characteristic of the Digger Pine
Belt and ranges from Mariposa County along the foothills of the
Sierra Nevada to Tehachapi Pass, while a slightly different form
occurs in Santa Barbara and Tulare Counties.
SPECIMENS EXAMINED:—CALIFORNIA: Western slope of Pecheco
Pass, Santa Clara County, Abrams 5285; Mormon Bar, Mariposa
County, 1883, Congdon; Wawona, Mariposa County, October, 1895,
Ward; Yosemite National Park, Bolander 5012; Marble Fork
of Kaweah River, Tulare County, August, 1905, Katharine Bran-
degee; Soda Springs of Kern River, Tulare County, Hall & Bab-
cock 5572; Visalia, Tulare County, October, 1881, Congdon;
North Fork of Kern River, Kern County, Coville & Funston 1602;
Poso Creek, Kern County, August, 1853, Heerman; Tehachapi,
Kern County, August, 1894, Eastwood; Santa Barbara, Santa
Barbara County, Elmer 3741; La Cumbre Trail, Santa Inez
Mountains, Santa Barbara County, Abrams 4302; road to
Monteci, Santa Barbara County, Eastwood 199; Mountain Drive,
Santa Barbara, Abrams 4113.
12. Adenostegia littoralis sp. nov.
Diffusely branching annual, 3-6 dm. high; stems puberulent,
without hirsute hairs; leaves 13-15 mm. long, entire; flower-heads
compact, five- to ten-flowered; floral bracts 14-22 mm. lo
three-parted, the divisions slender, pubescence soft-puberulent with
an occasional hispidulous hair, tips truncate or with a tack-like
enlargement of the veins, not distinctly calloused; lower calyx-leaf
at least 2 mm. longer than the upper, pubescence as on the bracts;
corolla 16-19 mm. long, the tube equalling or slightly shorter
than the throat; stamens four, the anthers two-celled. (PLATE
10, FIG. 13; PLATE 12, FIG. 1.]
This species was recognized as a form distinct from 4. rigida
by Heller (Muhlenbergia 2: 251), but no name was applied to it.
*
414 FERRIS: TAXONOMY AND
It differs most noticeably from A. rigida in the divisions of the
bracts, which are narrow instead of broad, in the pubescence and
in the long lower calyx-leaf.
TYPE LOCALITY: Carmel, Monterey County, California, open
pine woods, August 17, 1909, Abrams 4254. TYPE, No. 21173
of the Dudley Herbarium.
DISTRIBUTION: Sand dunes and open woods of the Monterey
Peninsula, California; Transition Zone.
SPECIMENS EXAMINED:— CALIFORNIA: Carmel, Monterey
County, Abrams 4254; near Del Monte, Monterey County,
Heller 8247; Del Monte, Elmer 4o8o.
13. ADENOSTEGIA RAMOSA (Nutt.) Greene
Cordylanthus ramosus Nutt.; Bentham in De Candolle, Prodr. 10:
597. 1846; Wats. Bot. King’s Exped. 232, 459; Gray, Bot.
Calif. 1: 580; Gray, Syn. FI. 2!: 303; Coulter & Nelson, Man.
Bot. Rocky Mts. 462.
Adenostegia ramosa Greene, Pittonia 2: 180. 1891; Kuntze, Rev.
Gen. 2: 456; Howell, Fl. N. W. Am. 1: 537.
Paniculately much branched annual, 2-3 dm. high, cinereous
puberulent throughout; leaves 15-20 mm. long, linear-filiform,
one- to three-parted, flowers sometimes solitary, mostly in three-
to five-flowered heads; floral bracts often scantily hirsute-ciliate,
parted into five to seven linear divisions; upper calyx-leaf 18-22
mm. long, broadly lanceolate or iaconspicuously bidentate,
five-nerved, lower calyx-leaf 17-20 mm. long, lanceolate or two-
lobed, pubescence like that of bract; corolla 15-17 mm. long, the
tube longer than the throat; stamens four, the anthers perfect,
the filaments villous; capsule 10-11 mm. long, slender; seeds
many, small, reticulate. [PLATE I0, FIG. 14.]
TYPE LocaLity: ‘In mont. Scopulosis (Nuttall! Tolmie!).”’
DISTRIBUTION: Southwestern Montana to southwestern Colo-
rado and westward to Oregon and Lassen County, California;
Arid Transition Zone.
SPECIMENS EXAMINED:— OREGON: Grizzly Butte, Crook
County, Letberg 848; Lake County, Mrs. Austin 1772; Warner
Range, Lake County, Coville & Leiberg 55; White Horse Moun-
tains, Lake County, Griffiths & Morris 449; eastern Oregon,
Cusick 1750, 1197. CALIFORNIA: Plumas Junction, Lassen
DISTRIBUTION OF ADENOSTEGIA 415
County, Eggleston 6205; Goose Lake Valley, Lassen County, Mrs.
Austin 245. NEVADA: Kings Cafion, Ormsby County, Baker 1499;
Quinn River Crossing, Humboldt County, Griffiths & Morris 139;
Tuscarora, Elko County, Heller 9182; same locality, Kennedy 606;
Blaine, Elco County, Heller 11115; Humboldt Wells, Elco County,
Heller 9182; hills around Austin, Lander County, Kennedy 4038;
Toyabe Range, Lander County, Kennedy 4093; Wells, Lander
County, August, 1882, Jones. IpAHO: Pocatello, Snake Plains,
Palmer 408; near big Camas Prairie, Henderson 3160; Dry
Creek, Snake Plains, Palmer 336; Houston, Henderson 37090;
Flint Creek, Owyhee County, MacBride 496; Blackfoot, Snake
Plains, Palmer 292; Minidoka, Lincoln County, Nelson & Mac-
Bride 1313. Utan: Parley’s Park, Summit County, Smith 1887;
Echo Cajfion, Watson 817. Montana: Grasshopper Valley,
southwest Montana, Watson 323. WyominG: Fossil Station,
August 10, 1885, Letterman; Dubois, Nelson 711; Evanston, July
10, 1897, Williams; river bottoms, Carbon County, Tweedy
3409; Cokeville, Cary 704; Crook Creek, Fremont County,
Goodding 528; Fort Bridger, July 29, 1873, Porter; Slater, Colo-
rado-Wyoming line, Carbon County, Goodding 1727. COLORADO:
Mancos, July, 1890, Miss Eastwood. .
14. ADENOSTEGIA WricuTit (Gray) Greene
Cordylanthus Wrightii Gray, Bot. Mex. Bound. 2: 120. 1859;
Gray, Proc. Am. Acad. 7: 382; Wats. Bot. King’s Exped. 459;
Gray, Syn. Fl. 2!: 453; Coulter, Contr. U.S. Nat. Herb. 2:
316; Coulter & Nelson, Man. Bot. Rocky Mts. 462.
Adenostegia Wrightii Greene, Pittonia 2: 180. 1891; Kuntze,
Rev. Gen. 2: 457; Wooton & Standley, Contr. U. S. Nat.
Herb. 19: 590.
Paniculately branching annual, 3-5 dm. high; inconspicuously
puberulent or in age glabrous; leaves linear-filiform, often dis-
sected; heads five- to ten-flowered, spreading; floral bracts 18-20
mm. long, glabrous, deeply divided into dissected, filiform divi-
sions; upper calyx-leaf 23-28 mm. long, five-nerved, shallowly
bidentate, lower calyx-leaf 24-30 mm. long, three- to four-nerved,
tip lanceolate or two- to four-toothed, the teeth 1-4 mm. long;
corolla 22-30 mm. long, the tube longer than the throat; stamens
four, the anthers two-celled, the filaments villous; capsule 11-12
mm. long; seeds reticulate. [PLATE 10, FIG. 15.
416 FERRIS: TAXONOMY AND
A form with scattered flowers, corresponding to an herbarium
specimen named by Gray for Dr. Matthews, occurs within the range
of A. Wrightii. As it appears to differ from A. Wrightii only in
having the flowers in smaller clusters it is not here considered
distinct.
TyPE LocaLity: ‘‘Prairies from 6-30 miles east of El Paso,
western Texas; Wright (450). Sand Hills, Chihuahua; Thurber.”
DistRIBUTION: Utah and southern Colorado to Arizona and
south to Chihuahua, Mexico.
SPECIMENS EXAMINED:—UTAH: Willow Creek, southeastern
Utah, Miss Eastwood 98; La Salle Mountains, Purpus 7020.
CoLorapo: Mesa Verde, southwestern Colorado, August, 1892,
Miss Eastwood. Arizona: Chiricahua Mountains, Blumer 1744;
White Mountains, Griffiths 5379; Oracle, August 28, 1903, Jones;
Black River, Rothrock 795; Strawberry Valley, Toumey 360;
Keans Caiion, 1897, Hough; Navajo to Hawthorne, Griffiths 5801;
San Francisco Mountains, Knowlton 210; Cosmino, Jones 4026;
Flagstaff, Leiberg 5780. Nrw Mexico: Navajo Indian Reserva-
tion, in Tunitcha Mountains, Standley 7841; Cedar Hill, San
Juan County, Standley 7953; Fort Wingate, Matthews 1883;
San Lorenzo, July 26, 1896, Wooton; Mogollon Mountains, Rushby
319; Farmington, San Juan County, Standley 7121; Bear Moun-
tain, Grant County, Metcalf 695. Mexico: Chihuahua, Pringle
780.
Section IV. Kinota
Cordylanthus § Hemistegia (in part) Wats. Bot. King’s Exped. 460.
1871; Gray, Syn. Fl. 2!: 304.
Flowers in heads, calyx monophyllous; corolla as in Euadenos-
tegia; stamens four, perfect.
15. ADENOSTEGIA KINGII (Wats.) Greene
Cordylanthus Kingii Wats. Bot. King’s Exped. 233, 460. pl. 22, f.
3-6. 1871; Gray, Bot. Calif. 1: 581; Gray, Syn. Fl. 2': 304;
Parry, Am. Nat. 9: 346; Coulter & Nelson, Man. Bot. Rocky
Mts. 462.
Adenostegia Kingii Greene, Pittonia 2: 181. 1891; Kuntze, Rev.
Gen. 2: 457; Wettstein in Engler & Prantl, Nat. Pflanzenfam.
4°°: 98; Rydberg, Fl. Colo. 318.
DISTRIBUTION OF ADENOSTEGIA 417
Paniculately branching annual, 1-3 dm. high, glandular puber-
ulent throughout, sometimes short-villous; leaves 20-30 mm. long,
three-parted into linear divisions; heads few-flowered, the flowers
spreading, giving an open appearance to the head; floral bracts
15-18 mm. long, three-nerved, irregularly dissected into five to
seven linear divisions; calyx-leaf 22 mm. long, five-nerved, the
teeth at the apex 1-2 mm. long; corolla 21-22 mm. long, the tube
glabrous, somewhat longer than the throat, the latter covered
with soft, reflexed hairs; filaments hairy; capsule pointed.
[PLATE I0, FIG. 16.]
Type Loca.ity: ‘Rare; found only on a limestone ridge near
Roberts Station in Monitor Valley, Nevada; 6,000 feet altitude;
July.”
DistrIBUTION: White Pine County, Nevada; Emery, Garfield
and Iron Counties, Utah.
SPECIMENS EXAMINED:>—UTAH: two miles south of Ferron,
Jones 5454; Sink Valley, June 20, 1890, Jones; Panguitch Lake,
Jones 6015; head of Sevier River, Jones 6032; southern Utah,
1875, Siler.
16. Adenostegia Helleri sp. nov.
Paniculately branching annual, 2-4 dm. high; glandular-
villous throughout with short, spreading hairs; leaves 6-15 mm.
long, one-, sometimes three-, nerved; heads many, terminating
the branchlets, one- to four-flowered; floral bracts 8-12 mm. long,
five- to eight-parted; calyx-leaf lanceolate or shallowly two-
toothed, four-nerved, winged at base; corolla 1-2 mm. longer than
the calyx, throat soft-pubescent, equaling or slightly exceeding
the tube; filaments nearly glabrous; capsule sharply pointed.
{PLATE 10, FIG. 17; PLATE 12, FIG. 2.
This species has been confused with A. Kingii but it differs in
habit and in the bracts which in A. Helleri are smaller and regu-
larly instead of irregularly divided.
TYPE LOCALITY: Hills north of Reno, Nevada, September 20,
1910, Heller 10238. Type, No. 2122 of the Dudley Herbarium.
DISTRIBUTION: Western Nevada, in the vicinity of Reno.
SPECIMENS EXAMINED:—NEvADA: Reno, Brown 1564; hills
north of Reno, Heller 10238.
Section V. DICRANOSTEGIA
Cordylanthus § Dicranostegia Gray, Proc. Am. Acad. 19:95. 1883;
Gray, Syn. Fl. 2!: 454.
418 FERRIS: TAXONOMY AND
Adenostegia § Dicranostegia Wettstein in Engler & Prantl, Nat.
Pflanzenfam. 4°°: 98. 1891.
Inflorescence spicate; calyx monophyllous; calyx-leaf deeply
divided into two parts.
17. ADENOSTEGIA ORCUTTIANA (Gray) Greene
Cordylanthus Orcuttianus Gray, Proc. Am. Acad. 19: 95. 1883;
Gray, Syn. FI. 21: 454.
Adenostegia Orcuttiana Greene, Pittonia 2: 181. 1891; Kuntze,
Rev. Gen. 2: 457; Wettstein in Engler & Prantl, Nat. Pflan-
zenfam. 4°: 98.
Divaricately branched, often decumbent, annual, 15-35 cm.
high; stems stout, densely hirsute, sparsely so with age; leaves
25-30 mm. long, one-nerved, densely hirsute, sparsely so with age;
leaves 25-30 mm. long, one-nerved, hispidulous, irregularly dis-
sected into linear divisions; spike 2-6 cm. long; floral bracts
20-25 mm. long, setose-ciliate, three-nerved, tip broadly rounded;
calyx-leaf 6 mm. long, thin, two-nerved, deeply or completely
divided into acuminate divisions, the margins with soft pubescence
interspersed with hirsute hairs; corolla about equalling the bracts,
tube longer than the throat, curved outward, throat soft-pubes-
cent; stamens four, the anthers small, one-celled with vestiges
of a second cell, the upper anther sterile, the lower fertile, filaments
glabrous. [PLATE 10, FIG. 18; PLATE 12, FIG. 3.
Type Locatity: ‘“‘Lower California, about 70 miles below
the U.S. boundary, H. C. Orcutt and son.”’
DIstRIBUTION: Northwestern part of Lower California, from
the international boundary to Ensenada.
SPECIMENS EXAMINED:—LOWER CALIFORNIA: Tia Juana, June
30, 1884, Orcutt & Son; same locality, July, 1896, Stokes; Mexican
Boundary, Mearns 3927; Las Huevitas, 1893, Brandegee; En-
senada, October 5, 1892, Brandegee; San Pedro Martir, Robinson
45 (probably San Pedro Martir Island, not San Pedro Martir
Mountains).
Section VI. CHLOROPYRON
Chloropyron Behr, Proc. Calif. Acad. I. 1: 61. 1855; Heller,
Muhlenbergia 3: 133.
Cordylanthus § Hemistegia Gray, Proc. Am. Acad. 7: 383. 1868;
Wats. Bot. King’s Exped. 460 (in part); Gray, Syn. Fl. 2!: 304.
Flowers in spikes; calyx monophyllous; corolla slightly sac-
DISTRIBUTION OF ADENOSTEGIA 419
cate, pink with purplish tip; stamens two or four, the upper, when
present, imperfect, the lower with dilated filaments.
Key to species of § Chloropyron.
Stamens two.
Mracts densely’ villous-hiroute oy oes oe fs hh ee ae 18. A. mollis.
acts eoatsely hirsitte.. 0 Cl oie ee 19. A. palmata.
Stamens four.
Herbage canescent, bracts lanceolate...........000ee sees 20. A. canescens,
Herbage glaucous, bracts generally three-toothed........... 2%. A. marilima.
18. ADENOSTEGIA MOLLIS (Gray) Greene
Cordylanthus mollis Gray, Proc. Am. Acad. 7: 384. 1868; Wats.
Bot. King’s Exped. 460; Gray, Bot. Calif. 1: 582; Gray, Syn.
Fl. 2!: 304; Jepson, Fl. W. Mid. Calif. ed. 2. 287.
Adenostegia mollis Greene, Pittonia 2: 181. 1891; Kuntze, Rev.
Gen. 2: 457; Wettstein in Engler & Prantl, Nat. Pflanzenfam.
4°: 98; Jepson, Fl. W. Mid. Calif. 417.
Chloropyron molle Heller, Muhlenbergia 3: 133. 1907.
Simple or divaricately branching annual, 3-4 dm. high with
sparsely hirsute stems; leaves 5-7 mm. long, the lower entire, the
upper incised, pubescence dense villous-hirsute; spike 3-10 cm.
long, dense, and drooping; floral bracts 18-22 mm. long, villous-
hirsute with long, spreading hairs, shallowly five- to eight-parted
into finger-like divisions, the middle division exceeding the others,
three-nerved; calyx-leaf 17-18 mm. long, four- or five-nerved,
with two, sometimes three, erect teeth, completely hidden by the
wide floral bract, lower part nearly glabrous, upper villous-hirsute;
corolla 15~16 mm. long, the tube longer than the throat; stamens
two, the upper pair lacking; capsule 7-9 mm. long, rounded;
seeds 2-5 mm. long, deeply reticulate. [PLATE 10, FIG. 19.
Type Locatity: “Mare Island, Bay of San Francisco, Charles
Wright, in N. Pacif. Expl. Expedition, November, 1855.”
Distripution: A very local species, found only along the
northern side of San Francisco Bay from Suisun to San Rafael.
SPECIMENS EXAMINED:— CALIFORNIA: Vallejo, July, 1883,
Congdon; salt marshes near San Rafael and Petaluma, Davy 4063;
Suisun Marshes near Suisun, Heller 7551; Suisun Marshes, 1892,
Jepson; Mare Island, 1855, bees (co-type).
420 FERRIS: TAXONOMY AND
19. Adenostegia palmata sp. nov.
Low annual, 1-2 dm. high, branching divaricately from the
base, sparsely hirsute throughout with short hairs; leaves 8-18
mm. long, mostly incised, the lower sometimes entire; spike 5-10
cm. long, dense, erect; floral bracts 12-18 mm. long, deeply
parted into five to eight finger-like divisions, the middle division
exceeding the others, three- to five-nerved, often reddish; calyx-
leaf 11-15 mm. long, entire or bidentate; corolla 12-16 mm. long,
conspicuous, tube longer than throat; stamens two, the upper pair
lacking. [PLATE 10, FIG. 20; PLATE 12, FIG. 4.]
TYPE LocaLity: In alkaline soil, overflowed lands at Tule near
College City, Colusa County, California, June 17, 1916, Stinch-
field 284. Typr, No. 70613 of the Dudley Herbarium.
This species differs most conspicuously from A. mollis in its
sparse pubescence and its deeply parted, palmate bracts.
20. ADENOSTEGIA MARITIMA (Nutt.) Greene
Cordylanthus maritimus Nutt.; Bentham in De Candolle, Prodr.
To: 598. 1846; Gray, Proc. Am. Acad. 7: 383; Wats. Bot.
King’s Exped. 460; Gray, Bot. Calif. 1: 581; Gray, Syn. FI.
2': 304; Jepson, Fl. W. Mid. Calif. ed. 2. 387.
Chloropyron palustre Behr, Proc. Calif. Acad. 1: 61. 1855.
Adenostegia maritima Greene, Pittonia 2: 181. 1891; Kuntze, Rev.
Gen. 2: 457; Wettstein in Engler & Prantl, Nat. Pflanzenfam.
4°: 98; Jepson, Fl. W. Mid. Calif. 417; Abrams, FI. Los
Angeles 372. : :
Chloropyron maritima Heller, Muhlenbergia 3: 133. 1907.
Decumbent annual, 2—3 dm. high, branching diffusely from the
base or above, erect when young; stems puberulent, sometimes
sparsely pubescent; floral bract 2-3 cm. long, pubescence varying
from short-hairy to short-villous, I-nerved, broadly lanceolate or
shallowly three-toothed, the lateral teeth the smaller; calyx-leaf
12-15 mm. long, toothed at apex, the teeth scarcely 1 mm. long,
covering as on bract; corolla 10-18 mm. long, covered with soft,
short pubescence; stamens four, the upper pair with second cell
imperfect or absent, the filaments glabrous; seeds 1-2 mm. long,
deeply reticulate. [PLATE 10, Fic. 21.]
This variable species has two intergrading forms. Around
San Francisco Bay and to the northward the plant is, in general,
twice as large as the Southern California form. Both of these
DISTRIBUTION OF ADENOSTEGIA 421
forms, however, vary in the pubescence of the spike from nearly
glabrous to short-villous in their respective localities. The floral
bracts are either lanceolate or toothed, generally the latter. In
its extreme variations the San Francisco Bay form approaches
A. mollis. The species is very closely related to A. canescens,
but the lanceolate bracts, the soft villous pubescence and the
scantily hairy filaments of the latter furnish a basis for their
separation.
Type Locatity: ‘Ad San Diego Californiae (Nuttall!).”
DistriBuTION: Salt marshes along the Pacific Coast from
Humboldt Bay in California to San Quintan, Lower California.
SPECIMENS EXAMINED:—CALIFORNIA: Hookton, Humboldt
Bay, Humboldt County, Tracy 3697; Samoa, Humboldt Bay,
Tracy 1257; San Rafael, Marin County, Bolander 2403; Tiburon
Peninsula, Marin County, Heller 5722; near San ‘Francisco,
Vasey 1875; Shell Mound, Oakland, Alameda County, July, 1880,
Rattan; Belmont, San Mateo County, Davy 4063; Cooley’s Land-
ing, Palo Alto, September 14, 1901, Dudley; Ravenswood, Palo
Alto, Philips & Stinchfield 269; Palo Alto, Baker 3557; same
locality, 1901, Congdon; same locality, Elmer 3423; Alviso, Santa
Clara County, July 18, Dudley; Milpitas, Santa Clara County,
June, 1905, Smith; Playa del Rey, Los Angeles County, Abrams
1714; San Pedro, Los Angeles County, Grant 3124; Coronado Sand
Spit, San Diego County, Chandler 4003; South San Diego, Octo-
ber 3, 1903, Brandegee; mouth of the Tia Juana River, Mearns
3914; Mexican Boundary Monument 258, Mearns 3931. LOWER
CALIFORNIA: San Quintan, May, 1899, Brandegee.
21. ADENOSTEGIA CANESCENS (Gray) Greene
Cordylanthus canescens Gray, Proc. Am. Acad. 7: 383. 1868;
Wats. Bot. King’s Exped. 233, 460; Gray, Bot. Calif. 1 581;
Gray, Syn. FI. 2!: 304.
Cordylanthus Parryi Wats. in Parry, Am. Nat. 9: 346. 1875.
Cordylanthus canescens var. Parryi Gray, Syn. Fl. 2!: 304. 1886.
Adenostegia canescens Greene, Pittonia 2: 181. 1891; Kuntze,
Rev. Gen. 2: 457; Rydberg, Bull. Torrey Club 40: 484.
Adenostegia Parryi Greene, Pittonia 2: 181. 1891.
Chloropyron canescens Heller, Muhlenbergia 3: 134. 1907-
Chloropyron Parryi Heller, Muhlenbergia 3: 134- 1997-
422 FERRIS: TAXONOMY AND
Divaricately and sometimes corymbosely branching annual,
2-4 dm. high, more or less canescent throughout with spreading,
villous hairs; leaves 10-20 mm. long, lanceolate, prominently
one-nerved, rarely three-nerved; spike erect, 2-4 cm. long; floral
bracts 20-25 mm. long, more canescent than the leaves, lanceolate,
often purplish at the tip; calyx-leaf 1-3 mm. shorter than the
bract, two- or three-toothed at apex, the teeth erect or spreading
with age; corolla tube equaling or slightly shorter than the throat;
stamens four, the upper pair with second cell imperfect or absent,
the filaments with scattered hairs, pectage | glabrous; capsule
rounded; seeds reticulate. [PLATE 10, FIG.
Examination of a fragment and a Sis cask of the type of
Cordylanthus Parryi shows that this is an immature specimen
of A. canescens.
TYPE LocaLity: ‘Near Carson City, Nevada, Dr. C. L.
Anderson.”’
DISTRIBUTION: Saline lakes and springs in the Great Basin
from eastern California and Oregon to Utah.
SPECIMENS EXAMINED:—OREGON: Goose Lake Valley, Lake-
view, Cusick 2767; Denio, August, 1901, Griffiths & Morris.
CaLirornia: Lake Lessons, Modoc County, Mrs. Manning
328; Honey Lake Valley, Lassen County, Davy 3390; Alkaline
Meadows, San Bernardino County, 1888, Parish. NEVADA:
Steamboat Springs, Heller 10367; same locality, Kennedy 1488,
1499; Eagle Valley, Baker 1265; Black Rock Desert, Griffiths &
Hunter 535; Glendale, Kennedy 1953; near Carson City, Anderson
201. Utan: Smelter Beach, Tooele County, 1891, Smith; Gar-
field Beach, Salt Lake, Rydberg 6897; Salt Lake City, Jones 1403;
Ogden Hot Springs, August 16, 1893, Ries.
Explanation of plates 10-12
E 10
1. Adenostegia laxiflora (Gray) Greene; ras from specimen from Beaver Creek,
Arizona, X 4.
2. spares Pringlet (Gray) Greene; bract from specimen from near Bartlett
Springs, California, « 2
3. a anak wipltdie (Nutt.) Greene; bract from specimen from Gold Creek,
Elko County, Nevada, x 114.
4. mae Mass (Gray) Greene; — from topotyp i _ x1%.-
Pe i
(Gray) G co-type speci fi + Lake
Tahoe, Neva 14.
6. Adenostegi viscida Howell; bract from specimen from Siskiyou Mountains,
Oregon, X
DISTRIBUTION OF ADENOSTEGIA 423
7. Adenostegia Hanseni Ferris; bract from the type, X 1.
8. Adenostegia parviflora Ferris; bract from the type, X 34.
9. Adenostegia Nevinii (Gray) Greene; bract from specimen from Bear Valley,
San Bernardino County, California, X 4.
10. Adenostegia filifolia (Nutt.) Abrams; bract from specimen from San Diego,
rene x 1K.
. Adenostegia Me Benth.; bract from specimen from the Santa Lucia
ania California, X 414.
12. Adenostegia reid brevibracteata (Gray) Greene; bract from co-type speci-
men, X 3%.
13. Adenostegia littoralis Ferris; bract from type, X 14.
14. erase ramosa (Nutt.) Greene; bract from specimen from Grizzly
Butte, Crook Cou mented
14
: Adnstaie saeisee (Gray) Greene; bract from specimen from Chiracahua
ade, Arizon
16. Rasiuetdsia Kingii (Wats. ) Greene; bract from specimen from near Ferron,
Utah, pi 3
<7. ‘Adsanitvete Helleri Ferris; bract from type, X I
18. Adenostegia Orcuttiana (Gray) Greene; bract in ii specimen from Tia Juana,
Laie: California
19. ON imolés (Gray) Greene; bract f: i from Suisun Marshes
Solano County, California, x 114.
20. Adenostegia palmata Ferris; bract from type, X 1
. Adenostegia maritima (Nutt.) Greene; bract from specimen from Palo. Alto,
Or sa x 14.
22. Adenostegia canescens (Gray) Greene; bract from specimen from Smelter
Beach, Tooele County, Utah, K 14
PLATE I1
1. Adenostegia laxiflora (Gray) Greene; Beaver Creek, Arizona
2. Adenostegia Pringlet (Gray) Greene; co-type specimen His Lake County,
California.
3. Adeuiaeal Hlanseni Ferris; type.
4. Adenostegia parviflora Ferris; type.
PLATE 12
Pm littoralis — type.
—ihdenas @ Relleri F type.
fe neta Orcuttiana (Gray) Greene; Tia Juana, Lower California, Stokes.
. Adenostegia palmata Ferris;
See
INDEX TO AMERICAN BOTANICAL LITERATURE
1916-1918
The aim of this Index is to include all current botanical literature written by
Americans, published in America, or based upon American material ; the word Amer-
ica being used in the broadest sense.
eviews, and papers ine aie exclusively to forestry, agriculture, horticulture,
manufactured products of vegetable origin, or laboratory methods are not included, anc
no ettenspt is made to index the literature of bacteriology. An occasional exception is
made in favor of some paper appearing in an American periodical which is devoted
wholly to botany. Reprints are not mentioned unless they differ from the original in
some important particular. If users of the Index will call the attention of the editor
to errors or omissions, their kindness will*be appreciated.
This Index is os monthly on cards, furnished in this form to subscribers
at the rate of on t for each card, Selections of cards are not permitied ; each
subscriber must jake ‘aD cards published ae ihe term of his subscription, Corre-
mdence relating to the card issue should be addressed to the Treasurer of the Torrey
Botanical Club
Alderman, W. H. Experimental work on self-sterility of the apple.
Proc. Am. Soc. Hort. Sci. 1917: 94-101. Mr 1918.
Allard, H. A. Effects of various salts, acids, germicides, etc., upon the
infectivity of the virus causing the mosaic disease of tobacco..
Jour. Agr. Research 13: 619-637. 17 Je 1918.
Anthony, R. D. Some results in the breeding of small fruits. Proc.
Soc. Hort. Sci. 1915: 121-125. F 1916.
Anthony, R. D., & Wellington, J. W. Experiments in bud selection with
the apple and violet at Geneva. Proc. Am. Soc. Hort. Sci. 1916:
71-76. Mr 1917.
Arthur, J.C. Uredinales of Guatemala based on collections by E. W.
D. Holway. I. Introduction, Coleosporiaceae and Uredinaceae.
Am. Jour. Bot. 5: 325-336. 6 Jl 1918.
Atkinson, G. F. Twin hybrids from crosses of Oenothera Lamarckiana
and Franciscana with Oe. pycnocarpa, in the F; and F;. Proc. Am.
Philos. Soc. 57: 130-143. pl. 1-4. 14 Je 1918.
Babcock, E. B., & Clausen, R. E. Genetics in relation to agriculture.
i-xx + 1-675. pl. 1-4 +f. 1-239. New York. 1918.
Bates, J.M. Anew Kochia. Am. Bot. 24: 51,52. My 1918.
Kochia alata sp. nov.
425
426 INDEX TO AMERICAN BOTANICAL LITERATURE
Beal, A.C. Monographic studies with flowers. Proc. Am. Soc. Hort.
Sci. 1916: 17-22. Mr 1917.
Blake, S. F. Notes on the flora of New Brunswick. Rhodora 20:
IOI-107. I1 Je 1918.
Blake, S. F. A revision of the genus Viguiera. Contr. Gray Herb.
Il. §4: 1-205. pl. 1-3. Je 1918.
Includes twenty-eight new species and many new names and combinations.
Bliss, M. C. Interrelationships of the Taxineae. Bot. Gaz. 66: 54-60.
bled, 2., 15-31. 1918.
Boynton, K. R. Helianthus orgyalis. Addisonia 3: 25. pl. 93. 29
Je 1918. ;
Boynton, K. R. Stylophorum diphyllum. Addisonia 3: 31, 32. pl. 96.
29 Je 1918.
Britton, N. L. Aronia arbutifolia. “Addisonia 3: 33. pl. 97. 29 Je
1918.
Broadhurst, J. Botanical errors of some well-known writers. Torreya
18: 117-119. Je 1918.
Brown, F.B. H. Scalariform pitting a primitive : eatine i in angiosper-
mous secondary wood. Science II. 48: 16-18. 5 Jl 1918.
Carpenter, C. W. A new disease of the Irish potato. Phytopath-
Glogy:8: 286, 287. pl. rz. 15 Jl 1918.
Chase, A. Axillary cleistogenes in some American grasses. Am. Jour. :
Bot. §: 256-258. f. 1-5. 21 Je 1918.
Chodat, R. Un voyage botanique au Paraguay (1914). Actes Soc.
Helvétique Sci. Nat. 992: 68-86. 1918.
Clute, W. N. The bloodroot. Am. Bot. 24: 41, 42. My 1918
[Illust.]
[Clute, W. N.] Herbs with juicy fruits. Am. Bot. 24: 53, 54- My
1918.
(Clute, W. N.] Note and comment. Am. Bot. 24: 64-71. My 1918.
Includes notes on Syeenatte botany, Dandelion flowers, Effects of pollination,
Anemone nomenclature, etc
[Clute, W. N.] A Seg flower. Am. Bot. 24: 59, 60. My 1918.
{Illust.]
Conn, H. J. The microscopic study of bacteria and fungiinsoil. New
York Agr. Exp. Sta. Tech. Bull. 64: 1-20. Ja 1918
Cook, M.T. Common diseases of berries. New Vises Agr. Exp. Sta,
Circ. 88: 1-11. f. -6. 1D 1917.
INDEX TO AMERICAN BOTANICAL LITERATURE 427
Cook, M. T. Common diseases of garden vegetables and truck crops.
New Jersey Agr. Exp. Sta. Circ. 89: 1-22. f. 1-12. 1-D 1917.
Coons, G.H. Michigan potato diseases. Michigan Agr. Exp. Sta.
Spec. Bull. 85: 1-49. f. r-41. Mr 1918.
Davis, J. J. Tilletia on wheat in North Dakota. Phytopathology 8:
247. 11 My 1918.
Dearing, C. The production of self-sterile muscadine grapes. Proc,
Am. Soc. Hort. Sci. 1917: 30-34. Mr 1918.
Dickson, J.G. The value of certain nutritive elements in the develop-
ment of the oat plant. Am. Jour. Bot. 5: 301-324. f. 1-5. 6 Jl 1918.
Dodge, B.O. Studies in the genus Gymnosporangium—I. Notes on
the distribution of the mycelium, buffer cells, and the germination of
the aecidiospore. Brooklyn Bot. Gard. Mem. 1: 128-140. pl. 1
+ f.1-5. 6 Jl 1918.
Dorsey, M. J. The inheritance of some biological facts on bud varia-
tion. Proc. Am. Soc. Hort. Sci. 1916: 41-71. Mr 1917.
Duff, G. H. Some factors affecting viability of the urediniospores of
Cronartium ribicola. Phytopathology 8: 289-292. f.z. 15 Jl 1918.
Durst, C. E. Studies in lettuce breeding. Proc. Soc. Hort. Sci. 1915:
96-98. F 1916. :
East, E. M. The rdle of reproduction in evolution. Am. Nat. 52:
273-289. Jl 1918.
Evans, A.W. The American species of Marchantia. Trans. Connecti-
cut Acad. Arts & Sci. 21: 201-313. f. 1-20. Mr. 1917 °
Includes Marchantia breviloba sp. nov.
Fernald, M. L. The diagnostic character of Vallisneria americana.
Rhodora 20: 108-110. 11 Je 1918.
Fernald, M. L. The geographic affinities of the vascular floras of New
England, the maritime provinces and Newfoundland. Am. Jour.
Bot. §: 219-236. pl. 12-14. 21 Je 1918.
Fernald, M.L. The contrast in the floras of eastern and western New-
foundland. Am. Jour. Bot. 5: 237-247. pl. 15-17-21 Je 1918.
Fitzpatrick, H. M. The life history and parasitism of Eocronartium
muscicola. Phytopathology 8: 197-218. pl. 1+ f. 1-4. 11 My 1918.
Fletcher, S. W. Fragaria virginiana in the evolution of the garden
strawberry of North America. Proc. Soc. Hort. Sci. 1915: 125-137-
1916. Peas
Fracker, S. B. Effect of crown gall on apple nursery stock. Phyto-
pathology 8: 247. 11 My 1918.
428 INDEX TO AMERICAN BOTANICAL LITERATURE
Gaskill, A. A shade tree guide. Rep. Dept. Conserv. & Develop.
New Jersey 1-22. f. 1-13. Union Hill. My 1918.
Gates,R.R. A systematic study of the North American Melanthaceae
from the genetic standpoint. Jour. Linnean Soc. 44: 131-172.
pl.5. 22 My 1918.
Gillespie, L. J. The growth of the potato scab organism at various
hydrogen ion concentrations as related to the comparative freedom
of acid soils from the potato scab. Phytopathology 8: 257-269.
fide VS GE vei8.
Glaser, R. W. A new bacterial disease of gipsy-moth caterpillars.
Jour. Agr. Research 13: 515-522. i. 54. 3Je1918.
Gilbert, W. W., & Gardner, M.W. Seed treatment control and over-
wintering of cucumber angular leaf-spot. Phytopathology 8: 229-
233. My 1918.
Goodale, G. L. The development of botany as shown in this journal.
Am. Jour. Sci. 46: 399-416. Jl 1918. .
Hance, R. T. Variations in the number of somatic chromosomes in
Oenothera scintillans DeVries. Genetics 3: 225-275. ol. 1-7 +
f. I-35. My 1918.
Harlan, H. V. Cultivation and utilization of barley. U. S. Dept-
Agr. Farm. Bull. 968: 1-39. f. z-ro. Je 1918,
Heinicke, A. J. Factors influencing the abscission of fowers and par-
tially developed fruits of the apple. Proc. Am. Soc. Hort. Sci.
1916: 95-103. Mr 1917.
Hitchcock, A. S. Generic types with special reference to the grasses
of the United States. Am. Jour. Bot. 5: 248-253. 21 Je 1918.
Hood, G. W. Inheritance in tomatoes.
88-95. F 1916.
Howe, M. A. Further notes on the structural dimorphism of sexual
and tetrasporic plants in the genus Galaxaura. Brooklyn Bot.
Gard. Mem. 1: 191-197. pl. 3,4+f. 1-4. 6Jl 1918.
Jones, D. F. Segregation of susceptibility to parasitism in maize.
Am. Jour. Bot. 5: 295-300. 6 Jl 1918.
Jones, L. R., & Gilbert, W. W. Lightning injury to herbaceous plants.
Phytopathology 8: 270-282. f. #3. 25 Vi tors.
Judd, C.S. The true mahogany tree. Hawaiian Forest. & Agr. 15:
105. Ap 1918. [Illust.]
Keitt, G. W. Inoculation experiments with species of Coccomyces
from stone fruits. Jour. Agr. Research 13: 539-570. pl. 55-59 +
f. t3.. 10 Je 1018.
Proc. Soc. Hort. Sci. 1915:
INDEX TO AMERICAN BOTANICAL LITERATURE 429
Knight, L.I. Physiological aspects of self-sterility of the apple. Proc.
Am. Soc. Hort. Sci. 1917: 101-105. Mr 1918.
Knowlton, C. H. Plants from South Weymouth, Massachusetts.
Rhodora 20: 115. 11 Je 1918.
La Rue, C. L., & Bartlett, H. H. “An analysis of the changes involved
in a case of progressive mutation. Genetics 3: 207-224: f. TI:
My 1918.
Loeb, J. Healthy and sick specimens of Bryophyllum calycinum.
Bot. Gaz. 66:69. 15 Jl 1918.
Lumsden, D. Orchid breeding. Jour. Internat. Gard. Club 2: 203-
213. Je 1918. [I[llust.]
Lyman,G.R. The relation of phytopathologists to plant disease survey
work, Phytopathology 8: 219-228. 11 My 1918.
MacCaughey, V. The Hawaiian kamani. Hawaiian Forest. & Agr.
15: 69-73. Mr 1918.
MacCaughey, V. The paradise tree. Hawaiian Forest. & Agr. 15:
20-22. Ja 1918
MacCaughey, V. A rare fruit tree of Hawaii. Hawaiian Forest. &
Agr. 14: 97-98. Ap 1917.
Diospyros ebenaster.
MacDougal, D. T. Effect of bog and swamp waters on swelling in
plants and in biocolloids. Plant World 21: 88-99. f. zr. Ap 1918.
MacMillan, H. G. Sunscald of beans. Jour. Agr. Research 13: 647-
650. pl. 64-66. 17 Je 1918.
Martin, G. W. Brown blotch of the Kieffer pear. Phytopathology 8:
234-238. f. r-9. 11 My 1918.
McClintock, J. A., & Smith, L. B. True nature of spinach-blight and
relation of insects to its transmission. Jour. Agr. Research 14: 1-60.
pl. A, 1-11 +f..1. 1 Jl 1918.
McCubbin, W. A. Public school survey for currant rust. Phytopa-
thology 8: 294-297. 15 Jl 1918.
Mertill, E. D. New species of Bornean plants. Philip. Jour. Sci. 13:
(Bot.) 67-122. Mr 1918.
Sixty-one new species are described in various genera.
Miller, E. C., & Coffman, W. B. Comparative transpiration of corn
and the sorghums. Jour. Agr. Research 13: 579-604. #l. es 63 +
F.d-13. - 30 Je 1918.
Muncie, J. H. Experiments on the control of bean anthracnose and
bean blight. New York Agr. Exp. Sta. Tech. Bull. 38: 1-50. pl.
ergs. te 1G17-
430 INDEX TO AMERICAN BOTANICAL LITERATURE
Nash, G. V. Cotoneaster Simonsit. Addisonia 3: 21, 22. pl. 91.
29 Je 1918.
Nash, G. V. Hamamelis japonica. Addisonia 3: 35, 36. pl. 98. 29
Je 1918.
Nash, G. V. Sobralia sessilis. Addisonia 3: 39, 40. pl. 100. 29 Je
1918.
Nash, G. V. Symphoricarpos albus laevigatus. Addisonia 3: 27.
pl. 94. 29 Je 1918.
Nelson, J. C. Western malodorous plants. Am. Bot. 24: 42-45.
My 1918.
Newcombe, F. C., & Bowerman, E. A. Behavior of plants in unventi-
lated chambers. Am..Jour. Bot. 5: 284-294. 6 J! 1918.
Orton, W. A. Breeding for disease resistance in plants. Am. Jour.
Bot. 5: 279-283. 6 Jl 1918.
Overholser, E. L. Color development and maturity of a few fruits
as affected by light exclusion. Proc. Soc. Am. Hort. Sci. 1917:
73-85. Mr 1918.
Pammel, L.H. The extermination of the common barberry to prevent
crop leakage due to stem rust. Iowa Conservation 2: 4-8. Mr
1918. [Illust.]
Pierce, R.G. Notes on peridermiums from Ohio. Need of pathologi-
cal viewpoint in nursery inspection. Phytopathology 8: 292-294.
15 Jl 1918.
Piper, C. V. New plants of the Pacific northwest. Proc. Biol. Soc.
Washington 31: 75-78. 29 Je 1918.
Epilobium cinerascens, Vaccinium coccineum, Mertensia bella, Castilleja indecora,
Grindelia andersonii and Hoorebekia curvaia, spp. nov., are described.
Pittier, H. New or noteworthy plants from Colombia and Central
America. 7. Contr. U.S. Nat. Herb. 20: 95-132. pl. 7 + f. 44-62.
Ig
inenaltin descriptions of 32 new species in various genera.
Potter, A. A. The effect of disinfection on the germination of cereal
seed. Phytopathology 8: 248, 249. 11 My 1018..
Record, S. J. Significance of resinous tracheids. Bot. Gaz. 66: 61-67.
f. 1-5. 135 Jl 1918.
Reddick, D. Lightning injury to grape vines. Phytopathology 8:
298. Je 1918. | '
Rehder, A. The Bradley bibliography 5: i—xxxii + I-1008. 191 8.
Roberts, R. H. Winter injury to cherry blossom buds. Proc. Am.
Soc. Hort. Sci. 1917: 105-110. Mr 1918. [Illust.]
432 INDEX TO AMERICAN BOTANICAL LITERATURE
Stone, R. E. Common edible and poisonous mushrooms of Ontario,
Ontario Dept. Agr. Bull. 263: 1-24. Je 1918.
Stout, A. B., & Boas, H. M. Statistical studies of flower number per
head in Cichorium Intybus: kinds of variability, heredity, and
effects of selection. Mem. Torrey Club 17: 334-458. pl. 10-13 +
702.10 Je 191%
Stout, A. B. Duplication and cohesion in the main axis in Cichorium
Intybus. Brooklyn Bot. Gard. Mem. 1: 480-485. pl. 12 +f. 1.
6 Jl 1918.
Stout, A. B. Hibiscus Moscheutos. Addisonia 3: 37, 38- pl. 99. 29
Je 1918.
Stuart, W. Some correlations in potatoes. Proc. Ann. Soc. Hort.
Sci. 1917: 39-45. Mr 1918.
Stuckey, H. P. The two groups of varieties of the Hicoria pecan and
their relation to self-sterility. Proc. Soc. Hort. Sci. 1915: 41-44.
F 1916.
Taubenhaus, J. J. Pox, or pit (soil rot), of the sweet potato. Jour.
Agr. Research 13: 437-450. pl. 51,52. 27 My 1918.
Tehon, L. R. Systematic relationship of Clithris. Bot. Gaz. 65:
552-555- pl. 18 Je 1918.
Includes Clithris clusiae, C. minor, and C. pandant, spp. nov.
Weir, J. R. Experimental investigations on the genus Razoumofskya.
Bot. Gaz. 66: 1-31. f. r-r9. 15 Jl 1918. aoe
Weir, J.R. Forest disease surveys. U.S. Dept. Agr. Bull. 658: I-23.
J. 1-23. 32 Je 1918.
Weldon, G. P. Pear growing in California. Month. Bull. State
Comm. Hort. Calif. 7: 222-410. f. 7-186. My 1918. [Illust.]
Contains chapter on diseases of pears.
Wells, B. W. The Zoocecidia of northeastern United States and
eastern Canada. Bot. Gaz. 65: 535-542. .18 Je 1918.
Whipple, O. B. Methods in pure line selection work with potatoes.
Proc. Am. Soc. Hort. Sci. 1917: 34-38- Mr 1918.
White, O.E. Breeding new castor beans. Jour. Heredity 9: 195-200.
f. 1-4 + frontispiece. 25 My 1918.
White, O. E. Our common garden vegetables, their history and their
origin. Brooklyn Bot. Gard. Leaflets 6: [1-19]. f. 5. 1 My 1918.
White, O.E. Inheritance studies in Pisum. Ill. The inheritance of
height in peas. Mem. Torrey Club 17: 316-322. f. I- 10 Je 1918.
INDEX TO AMERICAN BOTANICAL LITERATURE 431
Rock, J. F. Cyctandrene Hawaiienses, sect. Crotonocalyces Hillebr.
Am. Jour. Bot. §: 259-277. pl. 18-23. 21 Je 1918
Includes 4 new species in Cyriandra. .
Rose, J. N. Echeveria nodulosa. Addisonia 3: 23. pl. 92. 29 Je
1918.
Rose, J. N. Sinningia speciosa. Addisonia 3: 29, 30. pl. 95. 29 Je
1918.
Rosenbaum, J., & Ramsey, G. B. Influence of temperature and pre-
cipitation on the blackleg of potato. Jour. Agr. Research 13: 507—
513. 3 Je 1918.
Salmon, S. C., & Fleming, F. L. Relation of the density of cell sap to
winter hardiness in small grains. Jour. Agr. Research 13: 497-506.
pl. 53. 3 Je 1918.
Sampson, H.C. Chemical changes accompanying abscission in Coleus
Blumei. Bot. Gaz. 66: 32-53. 15 Jl 1918.
Saunders, A. P. Peony culture in America. Jour. Internat. Gard.
Club 2: 157-180. Je 1918. [Illust.]
Sax, K. The inheritance of doubleness in Chelidonium majus Linn:
Genetics 3: 300-307. My 1918.
Seifriz, W. Observations on the structure of protoplasm by aid of
microdissection. Biol. Bull. 34: 307-324. f. 1-3. My _ 1918.
[Ilust
Setchell, W. A. Parasitism among the red algae. Proc. Am. Philos.
Soc. 57: 155-172. 14 Je 1918.
Shufeldt, R. W. Pitcher-plants—what are they? Am. Forestry 24:
347-352. f. 2-9. Je 1918.
Smith, E. F., & Godfrey, G. H. Brown rot of Solanaceae on Ricinus.
Science II. 48: 42, 43. 12 Jl 1918.
Stanford, E. E., & Viehoever, A. Chemistry and histology of the
glands of the cotton plant, with notes on the occurrence of similar
glands in related plants. Jour. Agr. Research 13: 419-436. pl.
42-50. 20 My 1918.
Steil, W. N. Bisporangiate cones of Pinus montana. Bot. Gaz. 66:
OS. 7: 7." 45 fF 16x68.
Stevens, H. E. Lightning injury to citrus trees in Florida. Phy-
topathology 8: 283-285. f. z. 15 Jl 1918.
Stevens, N. E., & Wilcox, R.B. Further studies of the rots of straw-
berry fruits. U.S. Dept. Agr. Bull. 686: I-14. 24 Je 1918.
St. John, H. Spiranthes in Dover, Massachusetts. Rhodora 20:
FIt—114. 11 Je 1018.
BuLL. TORREY CLUB VOLUME 45, PLATE 10
Lh eS fo
t
TN
FERRIS: ADENOSTEGIA
BULL. TORREY CLUB VOLUME 45, PLATE II
FERRIS: ADENOSTEGIA
BULL, TORREY CLUB VOLUME 45, PLATE I2
FERRIS: ADENOSTEGIA
Vol. 45 No. 11
BULLETIN
OF THE
TORREY BOTANICAL CLUB
NOVEMBER, 1918
Revision of the North American species of Encalypta
Dorotuy COKER
(WITH PLATES, 13 AND I4)
The family Encalyptaceae, of which there is only one genus,
Encalypta, is closely related to the Pottiaceae, because of the
twisted and crisped habit of its leaves when dry, the small, thick-
walled, very papillose cells of the upper portion of the leaves,
and the large hyaline cells at the base. The costa also is strong
and often excurrent. The genus is noted for the great diversity
in the structure of its peristome, ranging from double, with several
remarkable variations, to single or absent.
Hedwig (4, p. 88), in 1782, based the genus Leersia on two spe-
cies, Bryum pulvinatum and B. extinctorium, referring the latter
to its Linnean synonym (see p. 103 in the descriptions of figures
19 and 24). Of these two species Bryum pulvinatum, which is a
Grimmia, precedes B. extinctorium; and since Grimmia, according
to Ehrhart (6, p. 176), antedates Leersia by one year, the name
Leersia has been discarded. Hence, the adoption of the name
Encalypta Schreb., 1791 (9, p- 759), instead of Leersia Hedw.,
1782, is due to the fact that Leersia, which in reality antedates
Encalypta, was originally used by Hedwig to include Grimmia.
Subsequently Hedwig, in 1801 (12, pp. 60-63), accepted the name
ncalypta in place of Leersia.
Following the older authors, Lesquereux and James (32, pp.
180-184) placed Encalypta under the Orthotrichaceae, and Schim-
[The Butterin for October (45: 391-432. pl. 10-12) was issued October 19, 1918.]
433
434 CoKrer: NortH AMERICAN SPECIES OF ENCALYPTA
per (28, pp. 307-340) with the Grimmiaceae; Lindberg (30, p. 26)
and Braithwaite (33, pp. 279-287) adopted Leersia for Encalypta,
referring it, as a subfamily, to the Tortulaceae, because of the
leaf-characteristics. Limpricht (32, pp. 102-123) and Dixon
(40, pp. 227-231) agree in recognizing the Encalyptaceae as dis-
tinct from the Pottiaceae, but Dixon places it under the subgroup
Aplolepideae and Limpricht uses the terms Haplolepideae and
Diplolepideae only in the specific descriptions. Loeske (48, p. 100)
rejects Lindberg’s usage of Leersia under the Tortulaceae and also
the placing by Brotherus (44, pp. 436-439) of the Encalypteae as a
subfamily of the Pottiaceae, and agrees with Fleischer (46, p. xiii),
who places them under a new group, the Heterolepideae, because
the peristome varies from the Haplolepideae to the Diplolepideae,
stating that they should be treated as a separate family.
In 1904 Paris Index (45, pp. 119-126) listed 30 species, of
which 18 were recorded for North America. According to ‘Broth-
erus, in 1902 (44, pp. 436-439), there were 35 species, of which 21
were recorded for the whole of America, 14 being endemic and
18 occurring north of Mexico. We* have reduced this number
to 8 by careful comparison of original and authentic specimens and
by studying the types whenever possible, having seen type speci-
mens of E. longipes, E. Macounii, and E. Selwyni from the Austin
and Mitten Herbaria, and authentic material of E. alaskana, E.
leiocarpa, E. subspathulata, E. cucullata, E. subbrevicolla, and
E. labradorica from the Macoun collections. The accompanying
lists and synonymy show the changes that have been made.
Only one species remains doubtful, E. lacera Ren. & Card. (38,
p- 91); it was described in a footnote to their check list of North
American mosses from specimens collected in Oregon by L. F.
Henderson, and no specimens have been obtainable.
ae 7 z 2 W2c4 £4
1753. EE. extinctoria (L.) Sw. Disp. Musc. Suec. 24. 1799.
1782. . laciniata (Hedw.) Lindb. Acta Soc. Sci. Fenn. 10: 267.
1872.
1788. EE. contorta (Wulf.) Lindb. Oefv. K. Vet. Akad. Foerh.
20: 396. 1863.
* Th
ts largely with Mrs. Britton.
f Adapted from shisthetne eg and Paris Index (45).
1897.
CoKER: NorTH AMERICAN SPECIES OF ENCALYPTA 435 ©
E.
E.
EE.
EE.
by
E.
E.
E.
EE.
E.
alpina Smith; Sowerby, Engl. Bot. pl. 1479. 1805.
rhabdocarpa Schwaegr. Suppl. 1:56. pl. 16. 1811.
apophysata Nees & Hornsch. Bryol. Germ. 2: 49. pl. 15,
Je $s 2827:
procera Bruch, Abh. Akad. Miinch. 1: 283. pl. rz. 1832.
mexicana C. Mill. Syn. 1: 516. 1849.
longipes Mitt. Jour. Linn. Soc. 8: 29. pl. 5. 1865.
Macount Aust. Bot. Gaz. 2:97. 1877.
Selwynt Aust. Bot. Gaz. 2: 109. 1877.
leiocarpa Kindb. Bull. Torrey Club 17: 275. 1889.
subspathulata C. Miill. & Kindb.; Macoun, Cat. Can.
Pl. 6393:° 1892.
. leiomitra (Kindb.) Kindb.; Macoun, Cat. Can. Pl. 6:
cucullata C. Mill. & Kindb.; Macoun, Cat. Can. PI.
6: 96. 1892.
alaskana Kindb.; Macoun, Cat. Can. Pl. 6: 269. 1892.
lacera Ren. & Card. Rev. Bryol. 19: 91. 1892.
subbrevicolla Kindb. Eur. & N. Am. Bryin. 2: 295.
1897.
labradorica Kindb. Eur. & N. Am. Bryin. 2: 295. 1897.
Arrangement of the North American species of Encalypta®* -
§ 1. Pyromitrium (Wallr.) Kindb.
1. E. alpina Smith (E. commutata Nees & Hornsch.).
~
fy
TAH
§ 2. XanTHOPUS Kindb.
E. ciliata (Hedw.) Hoffm. = E. laciniata (Hedw.) Lindb.
E. Macounii Aust. = E. apophysata Nees & Hornsch.
. E. alaskana Kindb. = E. laciniata.
§ 3. RHABDOTHECA Kindb.
. E. vulgaris (Hedw.) Hoffm. = E. extinctoria (L.) Sw.
E. rhabdocarpa Schwaegr. (including £. levomitra Kindb.).
E. lacera Ren. & Card. = E. rhabdocarpa?
8. E. subspathulata C. Miill. & Kindb. = E. rhabdocarpa.
9. E. mexicana C. Miill. = E. laciniata.
* Adapted from Kindberg (42, pp. 292-297) and Brotherus (44, pp. 436-439).
436 CoKER: NortTH AMERICAN SPECIES OF ENCALYPTA
§ 4. DipLoLepis Kindb.
10. E. longipes Mitt. = E. procera.
11. E. leiocarpa Kindb. = E. apophysata.
12. E. subbrevicolla Kindb. = E. brevicolla Bruch.
13. E. labradorica Kindb. = E. extinctoria.
§ 5. STREPTOTHECA Kindb.
14. E. contorta (Wulf.) Lindb. (E. streptocarpa Hedw.).
15. E. cucullata C. Mill. & Kindb. = E. procera + E. extinctoria.
16. E. procera Bruch.
17. E. Selwyni Aust. = E. procera.
ENCALYPTA Schreb. Genera 2: 759. 1791
Leersia Hedw. Fundam. 2:88, in part. 1782
Plants growing in dense compact cushions or mixed with other
cespitose ; stems usually low and crowded, seldom more than
4 cm. high; branches usually simple and sub-apical, erect, the leaves
uniformly spaced; leaves usually curled and twisted when dry,
spreading above when moist, from an erect clasping oblong base,
seldom more than 6 mm. long by 1.5 mm. broad; costa stout,
either ending below the apex or excurrent into a subulate or mu-
cronate point, cross-section of costa usually papillose above on both
faces, with a large dorsal band of stereid cells and several upper
layers of large ducts in 3-4 rows, the upper epidermal cells with
thick walls and smaller papillae; cells of the upper part of the leaf-
blade deeper than wide with clusters of prominent, minute papillae,
those of the lower part of blade larger, oblong and usually without
papillae, except in one species (E. apophysata) which has them on
the short end walls; margins entire, flat or slightly recurved,
rarely undulate. Perichaetial leaves generally smaller, more
acuminate and often subulate.
Monoicous, or in one exception (E. contorta) dioicous; some- -
times sterile and frequently propagating by elongated septate
brood-bodies; the antheridia in small lateral axillary buds; arche-
gonia terminal, vaginule enlarged, cylindric and often ochreate;
seta usually elongate, smooth except in E. streptocarpa, seldom more
than I-2 cm., rarely 3 cm., long (E. longipes), usually twisted;
calyptra large, 0.5-1 cm., completely covering the capsule to the
, cylindric and glossy, smooth or papillose at apex, sometimes
slightly papillose over the entire surface, ragged or fringed at base;
lid (operculum) large, never equalling the urn (theca), long-
CoKEeR: NortH AMERICAN SPECIES OF ENCALYPTA 437
rostrate; peristome originating at or below the mouth; simple
(haplolepideous) or compound (diplolepideous), sometimes lacking;
preperistome sometimes present; annulus simple or compound;
teeth varying in length, usually papillose or striate, entire or
split along the median line, rarely more or less united and attached
to the endostome by a hyaline membrane, sometimes nodose or
with short intermediate cilia; walls of the urn (theca) either smooth
or striate, striae sometimes visible only after sporosis, sometimes
spirally twisted or deeply grooved; neck (column) short, stoma-
tose, or the stomata scattered along the wall; spores smooth or
rough, usually maturing in spring or summer, variable in size.
Type species, Bryum extinctorium L.
The genus divides itself naturally into two groups or sections:
§ 1. Haplolepideae, with the peristome single or absent, and § 2.
Diplolepideae, with the peristome double and variable.
In §1 are included 4 species that are found in Europe and
North America: E. alpina, E. extinctoria, E. laciniata, and E.
rhabdocarpa, the last two approaching the Diplolepideae by having
a preperistome.
In § 2 are included 4 species, also common to Europe and
North America: E. apophysata, E. brevicolla, E. procera, and
E. contorta; of these E. contorta has not yet been found fruiting
on this continent. :
Key to species
§1. HapLoLePmear. Peristome simple or lacking; teeth, when
present, short, lanceolate; capsule not twisted.
Capsule striate, or becoming so when old.
Calyptra lacerate at base, nearly smooth at apex.
Peristome only occasionally found. Walls of cells
at base of leaves thickened. 1. E. exiinctoria.
Peristome usually present, often with preperistome.
alls of cells at base of leaves not thickened. 2. E. rhabdocarpa.
Capsule smooth.
Calyptra lacerate at base but not fringed. Peristome
never present, spores papillose.
Calyptra with a persistent or fugacious fringe of larger
cells. Peristome usually present, deeply inserted. . E. laciniata.
§ 2. DreLoLeripear. Peristome double, teeth usually long and
slender; ostome more or less adherent to the teeth.
3. E. alpina.
of leaves with short walls papillose. 5. E. apophysata.
438 CoKkEeR: NortH AMERICAN SPECIES OF ENCALYPTA
Teeth more or less united in pairs, unequal in length.
Basal cells of leaves not papillose. 6. E. brevicolla.
Capsule striate and spirally twisted, calyptra lacerate.
Monoicous, seta smooth, spores 15-25 y; leaves’slightly
toothed at base. 4. E. procera.
Dioicous, seta slightly papillose, spores 8-12 yu, leaves
entire 8. E. contoria.
§ 1. Haplolepideae
1. ENCALYPTA EXTINCTORIA (L.) Sw. Disp. Musc. Suec. 24. 1799
Bryum extinctorium L. Sp. Pl. 1116. 1753.
Leersia extinctoria Hedw. Fundam. 2: 88. 1782.
Leersia marginata Hedw. Fundam. 2: 103. 1782.
Leersia vulgaris Hedw. Descr. 1: 46. 1787.
Encalypta vulgaris Hoftm. Deuts. Fl. 2:27. 1796.
Encalypta cucullata C. Miill. & Kindb.; Macoun, Cat. Can. Pl.
6: 96, in part. 1892.
Encalypta extinctoria subsp. tenella Kindb.; Roll, Hedwigia 35:
65. 1896
Plants small, about 0.5-1 cm. high; leaves up to 4 mm. long,
I mm. wide; apical blade lingulate, apex cucullate contracted to
an abrupt point; costa ending below the apex, smooth on both
faces except slightly toothed at tip on back; margins plane, erose
above; papillose cells 12-14 long; cells of hyaline base oblong, .
up to 55 long by 15” wide, walls brown, slightly thickened at
ends, with 7-8 rows of long narrow cells at margin; perichaetial
leaves shorter and blunt at apex, usually carinate when moist.
Monoicous; vaginule about 1.5 mm. long; seta 5~8 mm., red brown,
not twisted; calyptra entire or ragged at base, slightly papillose
at apex; lid about 1.5 mm. long; capsule 2-3 mm. long by I mm.
wide, cylindric, smooth when young, ribbed when old; annulus
simple, narrow; mouth marked by an irregular, broken row of 1-2
quadrate, small, thickened cells; urn with the stomata sparsely
scattered over the entire surface; neck short, red, without stomata;
peristome, when present, of simple fugacious teeth; spores rough
with large rounded papillae, 24~32u, maturing in early spring.
TYPE LOCALITY: European.
DIsTRIBUTION: On rocks and earth in the Rocky Mountains,
from British Columbia to Colorado, and South Dakota; western
states from Nevada to California. Also Eurasia and Australasia,
according to Paris Index (45).
ILLustRaTIONS: Dill. Hist. Musc. pl. 45. f. 8. 1741 (as
Bryuwm); Hedw. Descr. 1: pl. 18. 1787 (as Leersia); Bryol. Eur.
pl. 199. 1838 (as E. vulgaris).
CoKER: NORTH AMERICAN SPECIES OF ENCALYPTA 439
ExsiccaTAE: Macoun, Can. Musci gor (as E. cucullata);
Holz. Musci Acro. Bor. Am. 214. 1906.
Ia. ENCALYPTA EXTINCTORIA APICULATA Wahl. Fl. Lapp. 344. 1812
Costa usually excurrent into a short hair-point; capsule when
mature striate and somewhat ribbed.
DISTRIBUTION: Colorado, Montana, and Assiniboia. Also
Europe.
tb. ENCALYPTA EXTINCTORIA MuUTICA Brid. Musc. Recent. Suppl.
4: 28. 1819
Costa disappearing far below the blunt apex; mature capsule
ribbed.
DistriguTION: Colorado to British Columbia. Also Europe.
2. ENCALYPTA RHABDOCARPA Schwaegr. Suppl. 1: 56. 1811
Leersia rhabdocarpa Lindb. Musci Scand. 26. 1879.
Encalypta rhabdocarpa var. leiomtira Kindb. Ottawa Nat. 4: 61.
1890.
Encalypta subspathulata C. Mill. & Kindb.; Macoun, Cat.: Can.
Fi.6: 0%. <1802.
Encalypta leiomitra Kindb.; Macoun, Cat. Can. Pl. 6:94. 1892.
Plants 1.5-2 cm. high; leaves 3-4 mm. by 0.66-I mm. wide,
lingulate, flat and spreading when moist; costa extending beyond
the suddenly contracted apex of leaf into a long mucronate hair-
orange, not thickened, with a distinct marginal border of 6-8
rows of cells, 60n long by 6—10p wide; perichaetial leaves smaller
and tapering to a hair-point. Monoicous; seta 6-8 mm., orange,
not twisted; calyptra entire or ragged at base, papillose at apex
and sometimes to about the middle; lid about 2 mm.; capsule
2-3 mm. long by 1 mm. wide, cylindric, striate, each ridge of
about 5-6 rows of cells; annulus simple; rim of the mouth marked
red, deeply wrinkled with large loose cells; peristome usually pres-
ent, single, of 16 red, finely striate, papillose teeth, with 4-5 seg-
ments, and occasionally with a narrow lateral preperistome cov-
ering 1-2 segments at base of the teeth; spores 40-50u in diameter,
very rough with large granular warts, ripe in late spring.
440 CoKER: NortH AMERICAN SPECIES OF ENCALYPTA
According to Limpricht (39, p. 115. f. 245, 246) there is great
variation in the peristomes of this species. However, we have not
found any peristome to correspond with his f. 246 in American
specimens.
TYPE LOCALITY: European.
DistrIBUTION: Arctic America, Greenland, Labrador to
Quebec and northern New York, Rocky Mountains from Mon-
tana to New Mexico, Pacific Coast ranges from Washington to
California. Also Europe and Asia.
ILLUSTRATIONS: Schwaegr. Suppl. 1: pl. 16. 1811; Bryol.
Eur. pl. 205. 1838.
ExsiccaTaE: Drummond, Musci Am. 50, 57 in part; also
52. 1828. Sull. & Lesq. Musci Bor. Am. 112. 1856; ed. 2. 166.
1865; Macoun, Can. Musci 731, 427 (as E. subspathulaia).
E. leiomitra differs from E. rhabdocarpa only in that the apex
of the calyptra is nearly smooth.
E. subspathulata is undoubtedly E. rhabdocarpa, but all of the
older capsules are badly infected by fungi and filled with hyphae.
2a. ENCALYPTA RHABDOCARPA PILIFERA (Funck) Nees & Hornsch.
Bryol. Germ. 2: 41. 1827
Encalypta pilifera Funck, in Sturm, Deuts. Fl. 17: pl: 5. 1819.
Leersta extinctoria var. pilifera Lindb. Musc. Scand. 20. 1879.
Leaves somewhat broader and more ovate; costa excurrent
into a long toothed hair; peristome perfect.
DISTRIBUTION: Fraser River Valley, Canada. Also Europe.
ILLUSTRATION: Sturm, Deuts. Fl. 17: pl. 5.
2b. ENCALYPTA RHABDOCARPA MICROSTOMA Breidler; Limpr.
Laubm. 2: 115. 1895
Capsule narrowing to a small mouth. Lid small, extended
into a long point; peristome perfect or rudimentary. A parallel
form to E. laciniata microstoma.
DisTRIBUTION: Alpine regions of the Rocky Mountains. Also
Europe.
3. ENCALYPTA ALPINA Smith; Sowerby, Engl. Bot. pl. rg19. 1 F
: 1805
Encalypta afinis Hedw. f. Weber & Mohr’s Beitr.r: 121. Mr 1805.
CoKER: NORTH AMERICAN SPECIES OF ENCALYPTA 441
Encalypta commutata Nees & Hornsch. Bryol. Germ. 2: 46. 1827.
Leersia alpina Lindb. Musci Scand. 20. 1879.
Plants 4-6 cm. high; leaves 3-4 mm. by I mm. subspatulate,
carinate; apex cucullate when moist; costa excurrent into a long
hair-point or ending below the apex, smooth except for a few teeth
just below the apex of leaf; margin plane; upper cells hexagonal,
I2u-16u, not densely papillose; cells of hyaline base 32y-48z
long by 16u wide, walls not thickened, becoming narrower and
longer toward the margin; perichaetial leaves smaller and tapering
to a long hair point. Monoicous; vaginule about 1 mm. long;
seta 7-9 mm. long, orange, seldom twisted; calyptra laciniate
at base, very slightly papillose at apex; lid 1.5 mm. long; annulus
of 2-3 rows of cells; capsule 2-3 mm. long by 0.75 mm. wide,
cylindric, smooth when young, appearing striate when old; mouth
marked by 1-2 rows of small red-brown quadrate cells often
irregularly broken; neck short, wrinkled, stomatose; peristome
none; spores 25u—35u, warty, often flattened and irregular, ripe
in late summer.
TYPE LOCALITY: European.
DIsTRIBUTION: Alpine regions of the Rocky Mountains from
Colorado to Washington; Alaska and Greenland. Also Europe
and Asia.
ILLUSTRATIONS: Sowerby, Engl. Bot. pl. 1419. 1805; Weber
& Mohr’s Beitr. 1: pl. 4. 1805; Schwaegr. Suppl. 1: pl. 16. 1811
(as E. affinis); Bryol. Eur. pl. 198. 1838.
ExsiccaTaE: Drummond, Musci Am. 49. 1828 (as E. affinis).
4. ENCALYPTA LACINIATA (Hedw.) Lindb. Acta Soc. Sc. Fenn.
10: 267. 1872
Bryum extinctorium var. 8. L. Sp. Pl. 1116. 1753 (see Dillen, 1,
Pp. 349; also Druce & Vines, 47, p. 210).
Leersia laciniata Hedw. Fundam. 2: 103. pl. 5, f. 24a. 1782.
Leersia ciliata Hedw. Descr. 1: 49. 1787.
Encalypta ciliata Hoffm. Deuts. Fl. 2:27. 1796.
Encalypia mexicana C. Mill. Syn. 1: 516. 1849.
Encalypta alaskana Kindb.; Macoun, Cat. Can. Pi. 6: 269. 1892.
Plants growing on wet limestone rocks; about 1-3 cm. high;
leaves carinate when moist, up to 5 mm. long, to about 1.3 mm.
wide; apical blade elliptic or lingulate, suddenly contracted to
a short mucronate point; costa thick, tapering into the short
finely serrulate tip or ending below the apex, slightly toothed on
442 CoKER: NorTH AMERICAN SPECIES OF ENCALYPTA
the back; margins revolute below the middle, above entire or
erose with truncate, minutely papillose cells up to 13-24u in
diameter; cells of the hyaline base not papillose, oblong, up to
60u long by 13-214 wide, becoming narrower toward the margin,
the basal cells with brown walls; perichztial leaves slightly smaller.
Monoicous; seta yellow to brown, 5-10 mm. high; calyptra
broadening at base into a deep regular fringe of larger cells,
smooth or slightly roughened at apex; lid beaked, up to 2 mm.
long; annulus present, of one row of cuneate cells; capsule 3-5
mm. long by I mm. wide, cylindric, smooth; mouth narrow
bordered by 3-5 rows of smaller thickened cells; urn with numerous
large stomata from the middle to the neck; peristome deeply
inserted, single; teeth lanceolate with 5-7 joints, which are longi-
tudinally papillose inside and occasionally with a darker colored
preperistome partially covering the basal segments; spores up to
37u in diameter, with radiating stellate lines, maturing in summer.
TYPE LOCALITY: European.
DistRiBuTION: Alpine and mountain regions; Eastern States
from Maine to northern New York: North Central States from
Michigan to Wisconsin, Minnesota; Rocky Mountains from
Montana to New Mexico; west coast from Washington to Cali-
fornia; British America from Ontario to British Columbia. Also
Europe, Asia, Africa and Australia.
ILLustrations: Dill. Hist. Musc. pl. 45. f. 9. 1741; Hedw,
Descr. 1. pl. 19. 1787; Bryol. Eur. pl. 200. 1838.
ExsiccaTaE: Drummond. Musci Am. 50. 1828; Sull. & Lesq.
Musci Bor. Am. rz. 1856; ed. 2. 165. 1865; Austin, Musci Ap.
174. 1870; Macoun, Can. Musci 132 (as E. ciliata), and 133 (as
E. Macounii); Allen, Mosses of Cascade Mts. Wash. 45; Holz.
Musci Acro. Bor. Am. 213. 1906; Pringle, Plantae Mexicanae
10547 (as E. mexicana).
4a. Encalypta laciniata microstoma (Schimp.) comb. nov.
Encalypta ciliata var. microstoma Schimp. Coroll. Bryol. Eur. 38.
1855.
Seta only 3-6 mm. high; calyptra with brown fringe; capsule
narrowing to a small mouth. Neck somewhat longer and running
down into the seta; Peristome smaller, irregular, often lacking;
spores very finely papillose, less transparent, and the radiating lines
less distinct; ripe in August
DIsTRIBUTION: Northern New York and in the alpine regions
of the Rocky Mountains. Also Europe and Asia.
CoKER: NortH AMERICAN SPECIES OF ENCALYPTA 443
Diplolepideae
5. ENCALYPTA APOPHYSATA Nees & Hornsch. Bryol. Germ. 2:
49. 1827
Encalypta Macounii Aust. Bot. Gaz. 2:97. 1877.
Encalypta leiocarpa Kindb. Bull. Torrey Club 17: 273. 1890.
Plants 1.5-2 cm. high; leaves 3-4 mm. long by 1 mm. wide,
carinate, lingulate; costa ending in the blunt apex or rarely excur-
rent into a short mucronate point, densely papillose on both surfaces
with coarse spinose teeth on dorsal apex; margins revolute above;
cells of upper blade 8—10u, irregular, those of the hyaline base
50-60 long by 8-1ou wide with the end walls thickened, with
papillose projections; perichaetial leaves slightly broader. Monoi-
cous. Seta 10-12 mm. long, smooth; calyptra 6-7 mm. long by
Imm. wide, very scabrous at apex, papillose over the entire sur-
face, basal fringe sometimes fugacious, cells of fringe narrow; lid
about 2 mm. high; capsule 2-3 mm. long by 0.5 mm. wide, not
striate, neck apophysate when dry or when wet long and tapering;
annulus of 2-3 rows of cells, more or less persistent; mouth bor-
dered by 3-4 rows of small thick-walled hexagonal cells; peristome
obscurely double; teeth slender, very papillose, perforate, rarely
bifid, inner peristome white, papillose, adhering closely and almost
invisibly to the outer; spores 18-24, finely papillose, maturing
in summer.
TYPE LOCALITY: European.
DistripuTION: Rocky Mountains of British Columbia to Mon-
tana; and (according to Paris Index) Scoresby Straits, Arctic
America. Also Europe and Asia.
ILLustRaTIoNs: Bryol. Eur. pl. 201. 1838; Limpricht, Laubm.
a: fj. 287. 180.
ExsiccataE: None. Drummond, Musci Am. 50, is E. lact-
niata; so are many of the specimens cited in Macoun’s Catalogue
(37) for E. Macounii. The American specimens of this species
seem to have the leaves more often blunt than is usual in the
European ones, though Limpricht (39) describes them as obtuse
or short-pointed. The type specimens of E. Macounii in Austin’s
herbarium are immature and no spores were formed, but in all
other characters they agree with E. apophysata. The description
of E. leiocarpa is erroneous in two important characters, for the
calyptra is fringed and the peristome is double.
444 Coxer: NortH AMERICAN SPECIES OF ENCALYPTA
6. ENCALYPTA BREVICOLLA (B. S. G.) Bruch; C. Mill. Syn.
I: 519. 1849
Encalypta longicolla var. brevicolla B. S. G. Bryol. Eur. (4:)
Encalypta 12. 1838.
Encalypia labradorica Kindb. Eur. & N. Am. Bryin. 2: 295. 1897.
Encalypta subbrevicolla Kindb. Eur. & N. Am. Bryin 2: 295. 1897.
Plants 1-1.5 cm. high; leaves 4-5 mm. long by 1 mm. wide,
subacuminate, carinate; costa excurrent into long colorless hair-
point, sometimes toothed at the base of awn, papillose on both.
surfaces; margins plane; upper cells of blade 12—16y in diameter,
irregular; those at the hyaline base 40-48u long by about 16y wide,
becoming narrower and colorless at the smooth margins; walls
orange-colored, slightly thickened at ends; perichaetial leaves
slightly shorter and broader. Monoicous; seta 1 cm. long, smooth,
red; calyptra 5 mm. long by 1 mm. wide, very scabrous at the
apex, papillose over the entire surface, lacerate at base; lid 2-2.25
mm. high, with a red border; capsule about 3 mm. long by 1 mm.
wide, not striate; neck short, stomatose, with large basal cells;
annulus none; mouth bordered by 4-5 rows of small, thick-walled,
quadrate cells; peristome double, deeply inserted; teeth .8 mm.
high, irregularly broken and branched, usually united in pairs at
base and perforate above, papillose; inner peristome similar and
attached to the outer, median segments longer than the lateral
ones; spores 28-32yu, very rough.
TYPE LOCALITY: European.
DiIsTRIBUTION: Labrador and, according to Paris Index, the
eastern coast of Greenland. Also Europe.
ILLUSTRATIONS: Bryol. Eur. pl. 202, 8. 1838.
ExsIccATAE: None.
7- ENCALYPTA PROCERA Bruch, Abh. Akad. Miinch. 1: 283. 1832
Encalypta longipes Mitt. Jour. Linn. Soc. 8: 29. 1865.
Encalypta Selwyni Aust. Bot. Gaz. 2: 109. 1877.
Leersia procera Lindb. Musci Scand. 20. 1879.
Leersia Selwynt E. G. Britton, Bull. Torrey Club 18: 50. 1891.
Encalypta cucullata C. Mill. & Kindb.; Macoun, Cat. Can. Pl. 6:
96, in part. 1892.
Plants 2-4 cm. high; leaves more or less spreading when dry,
5-6 mm. long by I mm. wide; apical blade subspathulate, apex
blunt; costa ending below apex, papillose on. upper surface, sca-
brous on back; margin revolute above; upper cells 12—16y in diam-
eter, round; those of the hyaline base, 48-60 long by 12-16u wide;
COKER: NortH AMERICAN SPECIES OF ENCALYPTA 449
walls deep orange, thickened at ends, basal margin slightly serrate;
with 3~—4 rows of narrow cells, walls colorless and ends unthickened ;
perichaetial leaves acuminate, tapering, with the costa percurrent
into a long hair-point. Monoicous; seta about 1.5-2 cm., smooth,
purple shading to orange above; calyptra 6-7 mm. long by 1.5
mm. wide, papillose at apex, very slightly so over the entire surface,
lacerate at base but without differentiation of cells; lid 2 mm.,
marked by ragged broken cells at base; capsule 3-4 mm. long by
0.4 mm. wide, cylindric, slightly striate when young, marked when
old by 8 striae, spirally twisted once or twice around the capsule;
annulus large, compound; mouth bordered by 2 rows of small
thick-walled quadrate cells; neck short, stomatose; peristome
double, teeth about 0.5 mm. long, narrow, red, smooth or papillose,
basal segments of teeth united and perforated; endostome papil-
lose, orange, as long as the teeth, attached to a papillose basal
membrane, the segments alternating with short cilia; spores I5~25u,
smooth, granular inside.
TyPE LOCALITY: European.
DISTRIBUTION: On earth in crevices of rocks and on banks;
Ontario to British Columbia and Alaska; the Rocky Mountains
of Idaho and Montana; Greenland. Also northern Europe and
Asia.
ILLustRaTIoNs: Abh. Akad. Miinch. 1: pl. 2. 1832; Bryol.
Eur. pl. 205. 1838; Mitt. Jour. Linn. Soc. 8: pl. 5. 1865 (as E.
longipes).
ExsiccaTaE: Drummond, Musci Am. 48. 1828 (as E. strep-
tocarpa); Macoun, Can. Musci 134 (as £. Selwyni), 474 (as E.
longipes), 491 in part (as E. cucullata), and 565 (as E. procera).
8. ENCALYPTA CONTORTA (Wulf.) Lindb. Oefv. Sv. Vet.-Akad.
Forh. 20: 396. 1863
Bryum céntortum Wulf.; Jacq. Coll. 2: 236. 1788.
Encalypta streptocarpa Hedw. Sp. Musc. 62. 1801.
Leersia contorta Lindb. Musci Scand. 19. 1879.
Plants 2-4 cm. high; leaves spreading when dry, 5-6 mm. long
by 1.5 mm. wide; apical blade lingulate, carinate; apex tapering
to the blunt point; costa ending below the apex, very papillose
on both surfaces, scabrous on back at basal portion; margins
plane; upper cells 12-16 in diameter, round, those of the hyaline
base, 40-48 long by 16 wide, not papillose; walls deep orange,
slightly thickened at angles, basal margins bordered by 2-3 rows
446 CoKER: NortTH AMERICAN SPECIES CF ENCALYPTA
of long narrow cells; perichaetial leaves 2-3 mm., more acuminate.
Dioicous. Seta about 1.5-2 cm. long, slightly papillose, purple;
calyptra 8-10 mm. long by 1 mm. wide, brown, lacerate at base,
very rough at apex, entire surface slightly papillose; lid 1.5 mm.
long; capsule 4-5 mm. by 2-3 mm. wide, larger at base, deeply
grooved with 8 striae, which are spirally twisted 2—3 times around
the capsule; annulus large, compound, persistent; mouth bordered
by 2-3 rows of small thick-walled quadrate cells; neck short,
stomatose, red; peristome double; teeth long, narrow, orange-
colored, very papillose ; endostome with 32 paler papillose segments,
one half the length of teeth, united at base by a thin papillose
membrane; spores 8—12y, smooth, irregular, ripe in early summer.
TYPE LOCALITY: European.
DIsTRIBUTION: On limestone rock, sand and earth in tem-
perate and alpine regions of Canada and Ontario to the Rocky
Mountains; Eastern States from Vermont to Virginia; Central
States from Ohio to Minnesota; Colorado and California, accord-
’ ing to Paris Index. Also Europe and Asia. Fruit plentiful in
Europe but not yet found in North America.
ILLUSTRATIONS: Hedw. Sp. Musc. pl. ro. 1801 (as Encalypta
streptocarpa); Bryol. Eur. pl. 204. 1838.
' ExsiccataE: Sull. Musci Allegh. r52. 1845; Aust. Musci App.
175. 1870; Macoun, Can. Musci 135; Holz. Musci Acro. Bor. Am.
I4I. 1904.
I would like to acknowledge my appreciation of the assistance
and unfailing interest given by Mrs. N. L. Britton, without which
this work could not have been done. I am also indebted to the
officers of the New York Botanical Garden for help and the use of
the Library and the Herbarium, and to Professor J. M. Macoun,
of the Department of Mines, Geological Survey, Ottawa, Canada,
for the loan of the Macoun collections of Encalypta, containing
the types and co-types of Kindberg and Miller.
COLUMBIA UNIVERSITY.
CHRONOLOGICAL BIBLIOGRAPHY
1. Dillen, J. J. Historia muscorum. Oxford. 1741.
2. Linnaeus, C. Species plantarum. Stockholm. 1753.
3. Swartz, O. Methodus muscorum illustrata. Upsala. 1781.
4. Hedwig, J. Fundamentum historiae naturalis muscorum fron-
dosorum 2. Leipzig. 1782.
5.. Leysser, F. W. von. Flora halensis. Ed. 2. Halle. 1783.
F
co
o
CoKER: NorTH AMERICAN SPECIES OF ENCALYPTA 447
Ehrhart, F. Beitrage zur Naturkunde 1. Hannover. 1787.
(Grimmia and Hedwigia. Hannov. Mag.19: 1089-1098. 1781.)
Hedwig, J. Descriptio et adumbratio microscopico-analytica
muscorum frondosorum 1. Leipzig. 1787.
. Wulfen, F. X. von. In Jacquin, N. J. von, Collectanea ad botani-
cam, chemiam et historiam naturalem spectantia 2. Vienna.
1788.
. Schreber, J.C. D. von. Genera plantarum 2. 1791.
. Hoffmann, G. F. Deutschlands Flora, oder Botanisches Taschen-
buch 2. Erlangen. 1795.
. Swartz, O. Dispositio me dayse. los muscorum frondosorum Sue-
ciae. Erlangen. 1799.
. Hedwig, J. Species muscorum frondosorum. Leipzig. 1801.
. Hedwig, R. A. Observationes de plantis caly ptratis, adjectis
novarum specierum descriptionibus. Weber & Mohr’s Bei-
triage zur Naturkunde 1: 106-131. pl. 4-7. 1805.
. Sowerby, J. English Botany 20. London. 1805.
. Schwaegrichen, C. F. Supplementum 1. [To Hedwig’s Species
muscorum frondosorum.] Leipzig. 1811.
. Wahlenberg, G. Flora lapponica. Berlin. 1812.
. Funck, H.C. Encalypta pilifera. In Sturm, J., Deuts. Fl. Crypt.
17t Di. 5. 1BI9.
. Bridel, S. E. Muscologiae recentiorum supplementum 4. Gotha.
1819.
. Bridel, S. E. Bryologia universalis 1. Leipzig. 1826.
Nees von Esenbeck, C. G., Hornschuch, C. F., & Sturm, J. Bryo-
logia germanica 2'. Nuremberg. 1827.
h, P. Beschreibung einiger neuen Laubmoose. Abh. Akad.
Minch. 1: 277-286. pl. 10, 11. 1832.
. Bruch, P., Schimper, W. P., & Giimbel, W.T. Bryologia europea
(4) Encalypta. Stuttgart. 1838.
. Miiller, C. Synopsis muscorum frondosorum I. Berlin. 1849.
Schimper, W. P. Corollarium Bryologiae europaeae. Stuttgart.
1855.
. Lindberg, S. O. Bidrag till mossornas synonymi. Oefv. Sv.
Vet.-Akad. Foérh. 20: 1863.
Mitten, W. The bryologia of the survey of the forty-ninth parallel
of latitude. Jour. Linn. Soc. 8: 12-55. Pl. 5-8. 1865.
. Lindberg, S.O. Contributio ad floram cryptogamam Asiae boreali-
orientalis. Acta Soc. Sci. Fenn. 10: 223-280. 1872.
imper, W. P. Synopsis muscorum europacorum. Ed. 2.
Stuttgart. 1876.
448
w
ot
CoKER: NorTH AMERICAN SPECIES OF ENCALYPTA
. Austin, C.F. Bryological notes. Bot. Gaz. 2: 95-98. 1877.
. Lindberg, S. O. Musci scandinavici. Upsala. 1879.
. Kindberg, N.C. Die Arten der Laubmoose (Bryineae) Schwedens
und Norwegens. Bih. Sv. Vet.-Akad. Handl. 7°: 1-167. 1883.
. Lesquereux, L., & James, T. P. Manual of the mosses of North
America. Boston. 1884.
. Braithwaite, R. The British moss-flora 1. London. 1887.
. Philibert, H. Etudes sur le péristome. Huitiéme article. Rev.
Bryol. 16: 39-44. 188
9.
. Kindberg, N. C. Contributions to Canadian bryology.—No. 3.
Bull. Torrey Club 17: 271-280. 1890.
. Britton, E. G. Contributions to American bryology.—II. Bull.
Torrey Club 18: 49-56. pl. 114. 1891.
. Macoun, J. Catalogue of Canadian plants. Part 6—Musci.
Montreal. 1892.
. Renauld, F., & Cardot, J. Musci Americae Septentrionalis, ex
operibus novissimis recensiti et methodice dispositi. Rev.
Bryol. 19: 65-96. 1892; 20: I-32.
. Limpricht, K. G. Die Laubmoose HEN Oesterreichs und
der Schweiz. Rabenhorst’s Kryptogamen-Flora. Ed. 2. 4’.
1895.
. Dixon, H. N., & Jameson, H. G. The student’s handbook of
British mosses. London. 1896.
Roll, J. Nachtrag zu der in der Hedwigia (Bd. X X XII, 1893)
erschienenen Arbeit tiber die von mir im Jahre 1888 in Nord-
Amerika gesammelten Laubmoose. Hedwigia 35:58-72. 1896.
. Kindberg, N.C. Species of European and Northamerican Bryineae
(mosses). Linképing. 1
al
897.
. Barnes, C. R., & Heald, F. D. Analytic keys to the genera and
species of North American mosses. Bull. Univ. Wisconsin 1:
157-368. 1896.
Brotherus, V. F. Pottiaceae. Engler & Prantl, Nat. Pflanzen-
fam. 1°: 380-439. f. 239-292. 1902.
. Paris, E.G. Index bryologicus. Ed. 2. 2. Paris. 1904.
. Fleischer, M. Die Musci der Flora von Buitenzorg 3. Leiden.
1906.
. Druce, G. C., & Vines, S. H. The Dillenian Herbaria. Oxford.
1907.
. Loeske, L. Studien zur vergleichenden Morphologie und phylo-
genetischen Systematik der Laubmoose. Berlin. 1910.
CoKER: NortH AMERICAN SPECIES OF ENCALYPTA 449
Description of plates 13 and 14
PLATE 13
A. ENCALYPTA EXTINCTORIA (L.) Sw. 1. Plant, natural size. 2. Single oe
. RE Meters forms of leaves. 4. Base of leaf showing hyaline cells with
5. Ape leaf showing smaller papillose cells. 6. Capsule enlarged.
7. Capt with cone apex. 8. Roughs
. ENCALYPTA RHABDOCARPA Schwaegr. 1. Plants, about natural size. 2.
Forms of Saks es. 3. Smooth Mout cells. 4. aaa vieptibomt cells. 5. Ribbed
capsule. 6. Calyptra with rough apex. 7. Peristome showing basal preperistome.
8. Rough spores.
C. ENCALYPTA ALPINA Smith. 1. Plants, natural size. 2. Forms of leaves,
3. Basal cells. 4. Apex with subulate awn. 5. Capsule. 6. Calyptra. 7. Por-
ie
sule. 6. Fringed calyptra. 7. Basal portion enlarged. 8. Rough peristome, and
rim of capsule. 9. Smooth, stellate spores.
PLATE 14 ,
A. ENCALYPTA APOPHYSATA Nees & Hornsch. 1. Plants, natural size.
Leaves. 3. Basal cells, with papillose transverse walls. 4. Apical cells. 5. Cap-
sule. 6. Papillose calyptra. 7. Base of same, fringed. 8. Double peristome, the
es.
B. ENCALYPTA BREVICOLLA Bruch. 1. Plants. 2. Leaves. 3. Cells of base-
4. Cells of apex. 5. Capsule. 6. Papillose calyptra. 7. Double papillose peri-
stome, the inner more or less branched. 8. Rough spore.
C. ENCALYPTA PROCERA Bruch. 1. Plant. 2. Stem and perichaetial leaves.
3. Smooth basal cells. 4. Papillose apical cells. 5. Twisted capsule. 6. Papillose
ragged calyptra. 7. Double peristome, the inner perforate at base. 8. Fragment
of annulus. 9. Smooth spores.
D. ENCALYPTA CONTORTA (Wulf.) Lindb. 1. Plants. 2. Leaf. 3. Basal cells.
4. Apical cells. 5. Twisted capsule. 6. Papillose and ragged ianioans: 7. Double
peristome more or less united with intermediate cilia. 8. Smooth spores
‘
Philippine Basidiomycetes—II|
PAUL W. GRAFF*
(WITH PLATE 15)
A large number of basidiomycetous fungi have been added from
time to time to those known from the Philippine Islands and, as
opportunity has offered, numerous tropical species described by
different botanists have been checked. The result is that, with
the extension of our knowledge, many ranges have been extended,
and not a few supposedly distinct species have proven to be iden-
tical. The confusion of names has, to a considerable extent, been
due to a failure to compare authentically named specimens from
the larger museums and herbaria and to a slight and possibly un-
conscious tendency to limit species distribution with geographic
boundaries. A brief review of the synonymy in the following
pages, in company with a comparison of named specimens, will
show the value of a comparative study of species from various
localities, and the need for a continuation of such studies. Ex-
‘tensive studies should also be made of material from Japan,
Formosa, China, the Malay States, Borneo, Java, Celebes, New
Guinea and Australia. This would bring to light many interesting
facts regarding little-known species. Likewise, a comparative
study of material from these localities with fungi collected in other
more distant and less related tropical countries would probably
broaden many vietws regarding the localization of fungal distri-
bution. For example, Awricularia mesenterica (Dicks.) Fr. is
also of common occurrence in the American tropics and southern
Europe; Corticium caeruleum (Schrad.) Fr. is found in North
America, southern Europe and northern Africa; and Pleurotus
flabellatus Berk. & Br. has been collected in tropical America,
Ceylon, and South Africa, as well as in the Philippines.
The material in the following list of fungi was collected on the
islands of Luzon, Mindoro, Polillo, Leyte, Negros, Culion, and
* Formerly Mycologist in the Biological Laboratory, Bureau of Science, Manila.
4
452 GRAFF: PHILIPPINE BASIDIOMYCETES
Mindanao. All of the collections cited are preserved in the
herbarium of the Bureau of Science, Manila. The preceding
parts of this series have been published in the Philippine Journal
of Science (8: Bot. 299-307. 1913; 9: Bot. 235-254. 1914).
USTILAGINEAE
UsTILAGO Persoon
UstiLaGo ANDROPOGONIS-ACICULATI Petch, Ann. Roy. Bot. Gard.
Perad. 4: 303. 1909.
Luzon: Manila, May, 1910, Merrill 7070, on Andropogon
aciculatus Retz.
Originally described from material collected on the same host
at Peradeniya, Ceylon.
UsTILAGO EMODENSIS Berk. Jour. Bot. & Kew Gard. Misc.
Sg: 200: 1851.
Luzon: Subprovince of Benguet, December, 1906, Mearns, on
Polygonum chinense L.
Described from material in Hooker’s Sikkim-Himalayan col-
lection from Tonglo, at an elevation of 3,300 m. The species is
closely related to U. Candollei.
UstiLAGO KoorDErsIANA Bref. Unters. Mykol. 12: 132. pl. 8,
15,0) 1808,
Necros: Cabacalan, March, 1910, Merrill 6766,on Polygonum
barbatum L.
Originally described on the same host from Java.
UsTILAGO PANICI MILIACEI (Pers.) Wint.; Rabenhorst, Krypt.-
Fl. 11:89. 1884
Uredo (Ustilago) segetum var. Panici miliacei Pers. Syn. Fung.
224. 1801.
Uredo carbo var. Panici miliace: DC. Fl. Franc. 6:76. 1815.
Caeoma destruens Schlecht. Fl. Berol. 2: 130. 1824.
Uredo destruens Duby, Bot. Gall. ed. 2.2: 901. 1830.
Erysibe Panicorum Wallr. Fl. Crypt. Germ. 2: 216. 1833.
Ustilago carbo var. destruens Tul. Ann. Sci. Nat. Bot. III. 7:
81. 1847.
Tilletia destruens Lév. Ann. Sci. Nat. Bot. III. 8: 372. 1848.
GRAFF: PHILIPPINE BASIDIOMYCETES 453
Ustilago destruens Schlecht.; Rabenhorst, Herb. Myc. Nov.
400.
MinpANnao: District of Cotobato, Cotobato, May 8, 1904,
E. B. Copeland, Bur. Sci. 1349, on Panicum sp.
Previously reported from North and South America and
southern Europe. Probably of very general distribution.
UsTILAGO TONKINENSIS (P. Henn.) Sacc. Syll. Fung. 11: 232.
1895.
Uredo tonkinensis P. Henn. Hedwigia 34:11. 1895.
Luzon: Province of Rizal, Caloocan, November 20, 1909,
C. B. Robinson, Bur. Sci. 9554, on Ischaemum aristatum var. gibbum.
Collected previously in the vicinity of Handi, French Indo-
China, parasitic on Andropogon sp.
CINTRACTIA Cornu
CINTRACTIA LEUCODERMA (Berk.) P. Henn. Hedwigia 34: 335.
1895.
Ustilago leucoderma Berk. Ann. Mag. Nat. Hist. II. g: 200.
1852.
Cintractia Krugiana Magn. Bot. Jahrb. 17: 490. 1893.
Cintracthia affinis Peck, N. Y. State Mus. Bull. 67: 28. 1903.
MINDANAO: District of Davao, Lake Lanao, Camp Keithley,
September, 1907, Mary S. Clemens, on Rynchospora glauca Vahl;
District of Zamboanga, November—December, 1911, Merrill 8322,
on Rynchospora aurea Vahl.
Reported from the southern portion of the United States, the
West Indies, Mexico, Brazil, and Borneo.
UstTILAGINOIDEA Brefeld
USTILAGINOIDEA VIRENS (Cooke) Tak. Bot. Mag. Tokyo to: 19.
1896.
Ustilago virens Cooke, Grevillea 7:15. 1878.
Ustilaginoidea Oryzae Bref. Unters. Mykol. 12: 194. 1895.
Tilletia Oryzae Pat. Bull. Soc. Myc. Fr. 13: 124. 1897.
Luzon: vicinity of Manila, P. W. Graff; Province of Laguna,
Santa Rosa, December, 1912, P. W. Graff, on Oryza sativa L.
This fungus is a common parasite on rice and is found in most
_ 454 GRAFF: PHILIPPINE BASIDIOMYCETES
of the rice-growing districts of India, Japan, the Se aee and
Malaya.
PUCCINEAE
Hamaspora Koernicke
HAMASPORA ACUTISSIMA Syd. Monogr. Ured. 3: 80. 1912.
Luzon: Province of Benguet, Pauai, June, 1909, R. C. Mc-
Gregor, Bur. Sci. 8510a, at an altitude of about 2,100 m.; Province
of Tayabas, Mount Banjao, March, 1907, F. W. Foxworthy,
Bur. Sci. 2399a. NEGROs: Canlaon Volcano, April, 1910, Merrill
222a. The host plant of the three collections is Rubus Rolfet
Vidal.
The species has also been reported on Rubus moluccanus L.,
collected in Java and Australia.
PucciniA Persoon
PUCCINIA iis (M. A. Curt.) Tracy & Earle, Bull. Miss. Agr.
Exp. Sta. 34: 86. 1895.
Uredo Hyptidis M. A. Curt. Amer. Jour. Sci. 2:6. 1820.
Uredo Cibertti Speg. Anal. Soc. Cien. Arg.g: 15. 1880.
Puccinia Ciberti Speg. Anal. Mus. Nat. Buenos Aires Il. 6:
220. 1899.
Aecidium Hyptidis P. Henn. Hedwigia 34: 337. 1895.
Aecidium tucumanense Speg. Anal. Mus. Nat. Buenos Aires
III. 9: 35. 1908.
Mrinpanao: District of Davao, Davao, March 21, 1904, Cope-
land 595, on Hyptis spicigera Lam. (Uredo stage).
Previously collected in the southern portion of the United
States and the tropical and temperate portions of South America.
PHRAGMIDIUM Link
PHRAGMIDIUM SUBCORTICIUM (Schrank) Wint.; Rabenhorst,
Krypt.-Fl. 11: 228. 1881.
Lycoperdon subcorticium Schrank, os Tasch. Hoppe 68.
1793
Uredo miniata var. Eglanteriae Pers. Syn. Fung. 216. 1801.
Uredo elevata Schum. Enum. PI. Sael. 2: 229. 1803.
Aecidium Rosae Rohling, Deuts. FI. ed. 2. 3: 122.
1813.
Uredo Eglanteriae H. Mart. Fl. Mosq. ed. 2. 230.
1817.
GRAFF: PHILIPPINE BASIDIOMYCETES 455
Caeoma miniatum Schlecht. Fl. Berol. 2: 120. 1824.
Caeoma Rosae var. miniatum Link; Willd. Sp. Pl. 62:30. 1825.
Erystbe miniata var. Rosarum Wallr. Fl. Crypt. Germ. 2: 200.
1833.
Coleosporium miniatum Bon. Gatt. Coniom. 60, in part. 1860.
Phragmidium Rosarum f. R. pimpinellifoliae Rabenh. Fungi
Eur. 7677 (hyponym). 1873.
Phragmidium Rosae pimpinellifoliae Diet. Hedwigia 44: 339.
1905.
Caeoma exitiosum Syd. Ured. 1700 (hyponym). 1913.
Luzon: Subprovince of Benguet, Baguio, June 10, 1913, J. C.
Wagner, Bur. Sci. 21002.
This species is of wide distribution, having been reported from
North America, Europe, Africa and Australia. It is probable
that the fungus was introduced into the Philippines with either
European or American cultivated rose stock.
AURICULAREAE
AURICULARIA Bulliard .
AURICULARIA LOBATA Sommerf.; Fries, Elench. Fung. 2: 34. 1828.
Luzon: Province of Rizal, Antipolo, October, 1912, M. Ramos,
Bur. Sci. S76.
Previously collected on the River Niger, Africa.
AURICULARIA MESENTERICA (Dicks.) Fr. Epicr. Myc. 555. 1838.
Helvella mesenterica Dicks. Plant. Crypt: 1:20. 1785.
Auricularia tremelloides Bull. Hist. Champ. Fr. 278. pl. 290.
1791-1798.
Auricularia violacea Bull. Hist. Champ. Fr. 278. 1791-1798
Thelephora mesenterica Pers. Syn. Fung. 571. 1808.
Thelephora purpurea Pers. Syn. Fung. 571. 1808.
Auricularia corrugata Sowerb. Engl. Fungi pl. 290. 1797-1815.
Auricularia ornata Pers.; Gaudichaud, Bot. Voy. Uranie 177.
pl. 2,f. 4. 1826.
Phlebia mesenterica Fr. Elench. Fung. 1: 154. 1828.
Oncomyces mesentericus Klotz. Linnaea '7: 195. 1832.
Luzon: Province of Rizal, January 16, 1906, F. W. Foxworthy,
Bur. Sci. 121; same locality, February, 1911, M. Ramos, Bur. Sct.
456 GRAFF: PHILIPPINE BASIDIOMYCETES
12552. MiInDANAO: District of Davao, Davao, April, 1904, Cope-
land “ F.
Of common occurrence in the American and European tropics.
AURICULARIA RUGOSISSIMA (Lév.) Bres. Hedwigia 53: 78. 1912.
Phlebia rugosissima Lév. Ann. Sci. Nat. Bot. II].2:214. 1844,
Phlebia reflexa Berk. Jour. Bot. & Kew Gard. Misc. 3: 16.
1851.
Mrinporo: Mount Halcon, November, 1905, Merrill 6118.
NeGros: Gimagaan River, January, 1904, Copeland 25.
Previously collected in the West Indies and northern India.
HIRNEOLA Fries
HIRNEOLA AFFINIS (Jungh.) Bres. Annal. Myc. 8: 587. IgIo.
Merulius affinis Jungh. Fl. Crypt. Java 76. 1838.
Laschia velutina Lévy. Ann. Sci. Nat. Bot. III. 2: 217. 1844.
Luzon: Province of Laguna, Mount Maquiling, February,
1912, P. W. Graff, Bur. Sct. 15.934.
Originally described from material collected in Java. This
fungus has also been reported from Japan.
HIRNEOLA AMPLA (Pers.) Fr. Fungi Nat. 26. 1848.
Auricularia ampla Pers.; Gaudichaud, Bot. Voy. Uranie 77.
1826.
Exidia nobilis Lév. Ann. Sci. Nat. Bot. III. 2: 218. 1844.
Exidia ampla Lév. Ann. Sci. Nat. Bot. II]. 5: 159. 1846.
Hirneola nobilis Fr. Fungi Nat. 26. 1848.
Luzon: vicinity of Manila, February, 1904, Copeland 41, on
Gliricidia maculata H.B.K. NrGrRos: Gimagaan River, January,
1904, Copeland 30, on Dipterocarpus sp.
Reported previously from Japan and Natal.
HIRNEOLA AURICULA-JUDAE (Fr.) Berk. Outl. Brit. Fung. 289.
1860; Fr. Hym. Eur. 695. 1874.
Tremella auricula L. Sp. Pl. 1157. 1753.
Merulius auricula Roth, Tent. Flor. Germ. 1: 535. 1788.
Peziza auricula L. Syst. Veg. ed. 13. 1018. 1789.
Tremella auriformis Hoftm. Veg. Crypt. 2: 31. bl. 6, f. 4. 1790.
Tremella caragana Pers. Syn. Met. Fung. 625. 1808.
GRAFF: PHILIPPINE BASIDIOMYCETES 457
Auricularia sambucina Mart. Fl. Crypt. Erlang. 459. 1817.
Exidia auricula-judae Fr. Syst. Myc. 2: 221. 1822.
Auricularia auricula Underw. Mem. Torrey Club 12:15. 1902.
Luzon: Province of Rizal, Bosoboso, July, 1906, M. Ramos,
Bur. Sci. 1185, on Gliricidia maculata.
Reported from southern Europe, Guiana, Japan, and the East
Indies.
HIRNEOLA TENUIS (Lév.) Fr. Fungi Nat. 27. 1848.
Exidia tenuis Lév. Ann. Sci. Nat. Bot. III. 2: 219. 1844.
Luzon: Province of Isabela, Cordon, May 8, 1909, M. Ramos,
Bur. Sct. 8135.
Previously collected in Natal, Borneo and Brazil.
THELEPHOREAE
CortTic1iuM Persoon
CorTICIUM CAERULEUM (Schrad.) Fr. Epicr. Myc. 562. 1838.
Thelephora caerulea Schrad. Spicil. Fl. Germ. 187. 1794.
Thelephora fimbriata Roth, Catal. Bot. 2. pl. 9, f. 2.
Auricularia phosphorea Sowerb. Engl. Fungi pl. 350. 1797-
1800,
1815.
Thelephora indigo Schwein. Schr. Nat. Ges. Leipzig 1: 107.
1822.
Merisma fimbriata Schwein. Schr. Nat. Ges. Leipzig I: 110.
1822.
Thelephora atro-caerulea Trog, Flora 15: 560. 1832.
Minporo: Mount Halcon, November, 1906, Merrill 5545, on
decaying tree branches, host plant undetermined.
Previously collected in North America, southern Europe and
northern Africa.
HyMENOCHAETE Léveillé
HYMENOCHAETE ApusTA (Lév.) Bres. Hedwigia 51: 323. 1912.
Stereum adustum Lév. Ann. Sci. Nat. Bot. III. 2: 213. 1844.
Thelephora adusta Lév.; Gaudichaud, Bot. Voy. Bonite 192.
pl. 139, f. 2: 1846.
Mrinpanao: District of Davao, Mount Apo, May, 1909, Elmer
10607. ae
Originally described from material collected in the vicinity
458 GRAFF: PHILIPPINE BASIDIOMYCETES
of Manila by Gaudichaud-Beaupré, on the occasion of the visit
of the French vessel ‘‘Bonite’’ to that port, and deposited in the
herbarium of the Museum of Paris.
HyMENOCHAETE CACAO Berk. Jour. Linn. Soc. Bot. 10: 333. 1869.
Stereum cacao Berk. Jour. Bot. & Kew Gard. Misc. 6: 169.
1854.
Luzon: Subprovince of Ifugao, Payauan, January 26, 1913,
R. C. McGregor, Bur. Sci. 20038; Province of Nueva Ecija, Cabana-
tuan, September, 1908, R. C. McGregor, Bur. Sci. 5250; Province
of Bataan, November, 1909, H. M. Curran, For. Bur. 19209.
MINDANAO: District of Davao, March, 1904, Copeland 495,
523; District of Zamboanga, Zamboanga, January, 1908, Whit-
ford & Hutchinson, For. Bur. 9269, 9296.
This species is of very general tropical distribution.
HYMENOCHAETE NIGRICANS (Lév.) Pat. Bull. Soc. Myc. France
mo iD. A9ey
Thelephora nigricans Lév. Ann. Sci. Nat.'Bot.III.2: 212. 1844.
Stereum nigricans Sacc. Syll. Fung. 6: 561. 1888.
NeEGROs: Province of Negros Oriental, Dumaguete, April, 1908,
Elmer 0851.
Previously reported from Cochin China.
HYMENOCHAETE RHEICOLOR (Mont.) Lév. Ann. Sci. Nat. Bot. III.
“Bi21r. 1846.
Stereum rheicolor Mont. Ann. Sci. Nat. Bot. II. 17:23. 1842.
Luzon: Province of Laguna, Mount Maquiling, February,
1912, P. W. Graff, Bur. Sci. 15948.
Collected previously in the East Indies and French Guiana.
HYMENOCHAETE TENUISSIMA Berk. Jour. Linn. Soc. Bot. 10:
333: 1869.
Stereum tenuissimum Berk. Jour. Bot. 6: 510. 1847.
Luzon: Province of Tayabas, Lucban, May, 1907, Elmer 7548.
Previously reported from the island of Ceylon.
STEREUM Persoon
STEREUM BELLUM Kunze, Flora 13: 370. 1830.
Mrxpanao: District of Davao, Todaya, April 26, 1904, Cope-
land 1294, on a decaying log.
GRAFF: PHILIPPINE BASIDIOMYCETES 459
Previously collected in the Madeira Islands.
STEREUM CONCOLOR Berk. Hook. Flor. Tasm. 259. 1860. Not
Stereum concolor (Jungh) Sacc., 1888.
Luzon: Province of Tayabas, Lucban, May, 1907, Elmer 8166,
on dead and decaying branches.
Described from material collected in Tasmania.
STEREUM CRENATUM Lév.Ann. Sci. Nat. Bot. III. 2: 210. 1844.
MinDANAO: District of Zamboanga, San Ramon, May, 1904,
Copeland 763, on a root clump of living Cocos nucifera L.
Previously collected in the island of Java.
STEREUM INVOLUTUM Klotz. Linnaea 7: 499. 1832.
NeEGROs: Province of Negros Occidental, Mount Canlaon,
September, 1909, H. M. Curran,’ For. Bur. 13738. MINDANAO:
. District of Davao, Davao, April, 1904, Copeland 1204.
Previously collected in Mauritius.
STEREUM LUGUBRE Cooke, Grevillea 12: 85. 1884. ©
Luzon: Province of Rizal, Bosoboso, July, 1906, M. Ramos,
Bur. Sci. 1203. Leyte: Dagami Panda, September 5, 1913,
Wenzel 3. :
Collected previously in New Zealand.
STEREUM OSTREA (BI. & Nees) Fr. Epicr. Myc. 547. 1838.
_ Thelephora ostrea Bl. & Nees, Fung. Java 13. pl. 2, f.1. 1826.
Luzon: Province of Benguet, Pauai, June,’ 1909, R. C. Mc-
Gregor, Bur. Sci. 8709, at an altitude of 2,100 m.
Found in Malacca and Java, and probably distributed through-
out Malaya in general.
STEREUM OSTREA var. CONCOLOR (Jungh.) Bres. Hedwigia 53: 76.
1912.
Thelephora concolor Jungh. Fung. Java 38. 1826.
Thelephora lobata Kunze, Linnaea 5: 527. 1830.
Stereum lobatum Fr. Epicr. Myc. 547. 1838.
Stereum perlatum Berk. Lond. Jour. Bot. §: 153. 1842. _
Stereum concolor Sacc. Syll. Fung. 6: 561. 1888. Not Stereum
concolor Berk., 1860.
460 GRAFF: PHILIPPINE BASIDIOMYCETES
Luzon: Province of Rizal, Bosoboso,July, 1906, M. Ramos,
Bur. Sci 1210; Palawan, Mount Pulgar, May, 1911, Elmer 13057.
Mrinpanao: District of Davao, Mount Apo, May, 1909, Elmer
10550.
This appears to be a case where the variety is much more
widely known than the species. It is certainly much the com-
moner in the Philippines. While the species, so far as is at present
known, seems to be quite rare in the Islands, the variety is one of
the most common of this group of plants. The three collections
enumerated above indicate range rather than prevalence. Ber-
keley’s species, Stereum perlatum, was originally described from
Philippine material.
Reported from the West Indies, Guiana, Brazil, Cochin China,
Sumatra, and generally throughout the East Indies.
STEREUM PRINCEPS (Jungh.) Lév. Ann. Sci. Nat. Bot. III. 2: 210.
1844.
Thelephora princeps Jungh. FI. aes Java 39. 1838.
Stereum scytale Berk. Jour. Bot. & Kew Gard. Misc. 6: 170.
1854.
Stereum contrarium Berk. Jour. Linn. Soc. Bot. 16: 52. 1878.
POLILLO: October-November, 1909, R. C. McGregor, Bur. Sci.
10547. MINDANAO: District of Davao, Mount Apo, June, 1909,
Elmer 11006.
Collected previously in Cuba, Brazil, Japan, Java, and north-
ern India. .
STEREUM SPECTABILE Klotz.; Meyen, Nov. Act. Acad. Nat. Cur.
19: Suppl. 1. 238. pl. 5, f. 2. 1843.
Stereum radiato-fissum Berk. & Br. Trans. Linn. Soc. II. 2:
63. pl. 14, f. 8-11. 1883.
Stereum luzoniense Ricker, Philip. Jour. Sci. 1: Suppl. 283.
1906.
Luzon: Province of Bataan, summit of Mount Mariveles,
on prostrate logs, October, 1903, Merrill 3531 (type of Stereum
luzoniense); Province of Laguna, Calauan, December 7, 1910, R.
C. McGregor, Bur. Sci. 12520.
Ricker’s species must be combined with Stehouths spectabile, as a
synonym, the two proving, on a careful comparison of the type
GRAFF: PHILIPPINE BASIDIOMYCETES 461
material of Stereum luzoniense with authentically named specimens
of the former, to be identical.
Besides being found in the Philippines, this species has been
reported from Australia and Mauritius.
CLADODERRIS Persoon
CLADODERRIS ELEGANS (Jungh.) Fr. Fung. Nat. 21. 1848.
Cymatoderma elegans Jungh. Ann. Sci. Nat. Bot. II. 16: 320.
1841.
Luzon: Province of Rizal, Bosoboso, July, 1906, M. Ramos,
Bur. Sci. 1189. MINDANAO: Subprovince of Butuan, March—July,
1911, Weber 128}.
This species shows a considerable tendency toward a transition
into Cladoderris infundibuliformis. So far, however, not enough
comparative material has been obtained to enable one to state
definitely whether this is a general condition or that the few speci-
mens showing the similarity are exceptional.
Collected previously in South Africa and the East Indies.
CLADODERRIS INFUNDIBULIFORMIS (Klotz.) Fr. Fung. Nat. 21.
1848.
Actinostroma infundibuliformis Klotz.; Meyen, Nov. Act.
Acad. Nat. Cur. 19: Suppl. 1.237. 1843.
Cladoderris Blumei Lév. Ann. Sci. Nat. Bot. III. 2: 213. 1844.
Cladoderris Decandolleana Lév. Ann. Sci. Nat. Bot. III. 2: 214.
1844.
Luzon: Province of Rizal, Bosoboso, July, 1906, M. Kamos,
Bur. Sci. 1184.
Cladoderris Blumei Lév., described from Javan material in
Blume’s herbarium and Philippine material in Delessert’s her-
barium, should unquestionably be placed as a synonym of C.
infundibuliformis.
Collected previously by Meyen, between 1830 and 1832, in the
vicinity of Manila. The fungus has been also collected in Java,
South Africa, and Brazil.
HYMENOLICHENES
Cora Fries
Cora GyroLoputa Fr. Epicr. Myc. 556. 1838.
Gyrolophium aeruginosum Kunze; Krombh. Consp. Fung. pl. 5,
Je 2G. 1821. .
462 GRAFF: PHILIPPINE BASIDIOMYCETES
LeyTE: Dagami Panda, September 8, 1913, Wenzel 5, growing
in the moist forest at an altitude of 60 m., on tree ferns.
Previously collected in Mauritius.
Cora PAVONIA (Web. & Mohr) Fr. Epicr. Myc. 556. 1838.
Thelephora pavonia Web. & Mohr, Beitr. Naturk. 1: 326. 1805-
Luzon: Province of Benguet, vicinity of Baguio, May, I9gII’
C. B. Robinson, Bur. Sci. 14107.
Collected previously in the West Indies and other, parts of the
American tropics.
CLAVARIEAE
LACHNOCLADIUM Léveillé
LACHNOCLADIUM FLAGELLIFORME (Berk.) Cooke, Austr. Fungi
179. 1892.
Clavaria flagelliformis Berk.; Hooker, Fl. New Zeal. 186. 1855.
MINDANAO: District of Davao, Mount Apo, May, 1909, Elmer
11301.
Described from material collected in New Zealand. As yet
reported only from these Pacific regions.
LACHNOCLADIUM USAMBARENSE P. Henn. Bot. Jahrb. 29: 44.
1904.
Minporo: Bulalacao, August-September, 1906, Bermejos, Bur.
Sct. 1553.
Previously known only from East Africa.
PHYSALACRIA Peck
PHYSALACRIA ORINOCENSIs Pat. Bull. Soc. Myc. Pr. i: 43. ol. 13,
f.2. 1884.
Luzon: vicinity of Manila, October, 1912, Merrill, on decaying
bamboo stalks.
This interesting little species of “Bubble Cap”’ has also been
collected by the writer in Manila, where it was growing in profusion
on bamboo joints which were being used for flower pots. Being
used for seedling plants and cuttings, these pots were kept con-
stantly moist and in a shady place so that they formed an ideal
substratum for a fungal growth.
Collected previously in tropical America.
GRAFF: PHILIPPINE BASIDIOMYCETES 463
HYDNEAE
IRPEX Fries
IRPEX FLAVUS Klotz. Linnaea 8: 488. 1833.
Polyporus flavus Jungh. Flor. Crypt. Java 46. 1838.
NEcRos: Gimagaan River, January, 1904, Copeland 26, on
decaying wood.
Coriolopsis melleoflavus Murr. (Bull. Torrey Club 35: 393.
1908) is not a synonym for this species as stated by Bresadola
(Hedwigia 53: 75. 1912) but belongs rather among the numerous
synonyms of Polystictus cervino-giluus (Jungh.) Fr. For this ar-
rangement see Graff (Philip. Jour. Sci. 9: Bot. 239. 1914).
Collected in North America, Java, and the East Indies. When
further known the species will, in all probability, be found to grow
in the American tropics as well.
POLY POREAE
Potyporus (Micheli) Fries
PoLyporus Apustus (Willd.) Fr. Syst. Myc. 1: 363. 1821.
Boletus adustus Willd. Fl. Berol. Prod. 392. 1787.
Polyporus carpineus Sowerb. Engl. Fungi pl. 231. 1799.
Polyporus crispus Fr. Obs. Myc. 1:127. 1815.
Polyporus pallescens Fr. Syst. Myc. 1: 369. 1821.
Boletus isabellinus Schw. Schr. Nat. Ges. Leipzig 1:96. 1822.
Polyporus subcinereus Berk. Ann. Mag. Nat. Hist. II. 3: 391. -
1839.
Polyporus Halesiae Berk. & Curt. Ann. Mag. Nat. Hist. II.
12: 434. 1853.
Polyporus Lindheimert Berk. & Curt. Grevillea 1: 50. 1872.
Bjerkandera adusta Karst. Medd. Soc. F. et Fl. Fenn. 5: 38.
1879.
Myriadoporus adustus Peck, Bull. Torrey Club 11: 27. 1884.
Polyporus Burtii Peck, Bull. Torrey Club 24: 146. 1897.
Mrnpanao: District of Davao, Mount Apo, April 20, 1904,
Copeland 1075; Lake Lanao, Camp Keithley, July, 1907, Mary S.
Clemens. :
A fungus of very general distribution.
Po.yporus atypus Lév. Ann. Sci. Nat. Bot. III. 2: 184. 1844.
Trametes aurora Ces. Myc. Borneo 5. 1879.
464 GRAFF: PHILIPPINE BASIDIOMYCETES
Coriolus atypus Pat. Tax. Hymén. 94. 1900.
Polystictus atypus Bres. & Syd. Philip. Jour. Sci. 9: Bot. 348.
1914.
Mrnporo: Puerto Galera, June, 1912, P. W. Graff, Bur. Sci.
530, growing on dead branches.
Previously collected in Java and Borneo.
POLYPORUS BENGUETENSIS (Murr.) Graff, Philip. Jour. Sci. 9: Bot.
236. 1914.” [PLATE 15, Fic. 1].
Coltricia benguetensis Murr. Bull. Torrey Club 35: 391. 1908.
Polystictus benguetensis Sacc. & Trott.; Saccardo, Syll. Fung.
222 32t.° 1912.
Luzon: Province of Benguet, Baguio, October-November,
1905, Merrill 5003 (type of Coltricia benguetensis Murr.); same
locality, March, 1904, Elmer 6047—both collections from prostrate
' logs of Pinus insularis Endl.
Upper surface and pore layer of the same shade of brown and
quite distinct in color from the flesh of the pileus which is a shining
golden yellow-brown when broken. The characteristics are so
distinct that its place in the genus Polyporus cannot be questioned.
As yet, only collected in the Philippines.
POLYPORUS LUZONENSIS (Murr.) Sacc. & Trott.; Saccardo, Syll.
Fung. 21: 266. 1912.
Spongipellis luzonensis Murr. Bull. Torrey Club 34:473. 1907.
Luzon: Province of Bataan, Mount Mariveles, November,
1904, Elmer 6944, on dead wood.
Described from Philippine material and, so far, not reported
from elsewhere.
| Fomes Fries
FOMES ALBOMARGINATUS (Lév.) Sacc. Syll. Fung. 6: 185. 1888.
Polyporus albomarginatus Lév. Ann. Sci. Nat. Bot. III. 2:
I9t. 1844.
Polyporus kermes Berk. & Br. Jour. Linn. Soc. Bot. 14: 49.
1875.
Polyporus laeticolor Berk. Jour. Linn. Soc. Bot. 16: 46. 1878.
Fomes pyrrhocreas Cooke, Grevillea 34:32. 1885.
Polyporus ochrocroceus P. Henn. Monsunia 1: 145. 1899.
GRAFF: PHILIPPINE BASIDIOMYCETES 465
Pyropolyporus albomarginatus Murr. Bull. Torrey Club 34:
478. 1907.
MINDANAO: District of Davao, Mount Apo, April 20, 1904,
Copeland 1077, growing on a fallen log in the forest.
Collected previously in Ceylon, Java, and New Guinea.
Fomes FuLLAGER! Berk. Jour. Linn. Soc. Bot. 16: 54. 1878.
Necros: Province of Negros Occidental, Farar, October 8,
1909, H. M. Curran, For. Bur. 17476, growing on living Shorea sp.
According to the collector’s note, the fungus sporophore was
growing from a wound, and the fungus had caused a rotting of the
heartwood of the tree. The rot is not accompanied by any stain-
ing of the wood. It is possible that this fungus may be found to be
one of the important timber destroyers of the Philippine Islands.
This fungus has been previously collected in Australia.
FoMES MELANOPORUS (Mont.) Sacc. Syll. Fung. 6: 196. 1888.
Polyporus melanoporus Mont.; Ramon de la Sagra. Hist.
Phys. Polit. Nat. Cuba 9: 422. 1841.
Polyporus cinereo-fuscus Curt. Trans. Linn. Soc. IT. t: 124.
$l. 10, {7 2. 1876.
Fomes melanoporoides Ces. Myc. Borneo 6. 1879.
Fomes cornu-bovis Cooke, Grevillea 13:2. 1884
Nigrofomes melanoporus Murr. Bull. Torrey Tank at: 425."
1904.
Luzon: Province of Bataan, Mount Mariveles, November,
1904, Elmer 6050.
Reported from Cuba, Borneo, Malacca and India.
Fomes MErritiit (Murr.) Sacc. & Trott.; Saccardo, Syll. Fung.
21:207. 3912,
Pyropolyporus Merrillit Murr. Bull. Torrey Club 34: 479.
1907.
CuLion: December, 1902, Merrill 3575, 00 decaying tree trunks
near the seashore.
As yet reported only from the Philippines.
Fomes rosgvs (Alb. & Schw.) Cooke, Grevillea 14: 21. 1885.
Boletus roseus Alb. & Schw. Consp. Fung. 251. 1805.
466 GRAFF: PHILIPPINE BASIDIOMYCETES
Polyporus roseus Fr. Syst. Myc. 1:372. 1821.
Polyporus carneus Nees, Nov. Act. Acad. Nat. Cur. 13: 14.
Ol. 3: £887.
Polyporus rufo-pallidus Trog, Flora 15: 556. 1832.
Fomitopsis rosea Karst. Rev. Myc. 3: 18. 1881.
Fomes carneus Cooke, Grevillea 14: 21. 1885.
Luzon: Province of Laguna, Mount Maquiling, April, 1913,
P.W. Graff, Bur. Sci. 21042, on a prostrate log near the trail at an
altitude of 150 m.
Previously collected in North, Central and South America,
Europe, Australia and Java. In the Americas its habitat is con-
fined to the tropical and warmer temperate portions. In all
probability Africa will be added to this list in the course of time
and the species will be found to be of universally tropical distribu-
tion.
Potystictus Fries
POLYSTICTUS ABIETINUS (Dicks.) Fr. Syst. Myc. 1: 370. 1821.
Boletus abietinus Dicks. Plant. Crypt. Brit. 3: 21. pl. 9, f. 9.
1793-
Polyporus incarnatus Schum. Enum. Plant. Saell.2: 391. 1803.
Sistotrema violaceum Pers. Syn. Fung. 551. 1808.
Polyporus dolosus Pers. Myc. Eur. 2:77. 1828.
Coriolus abietinus Quél. Enchir. Fung. 175. 1886.
Luzon: Bontoc Subprovince, February—March, 1911, Vanover-
bergh 1117, on the bark of Pinus insularis Endl.
Found in northern North America, Cuba and Europe.
POLYSTICTUS PERPUSILLUS (Murr.) Sacc. & Trott.; Saccardo,
Syll. Fung. 21: 319. 1912.
Coriolus perpusillus Murr. Bull. Torrey Club 35: 396. 1908.
Minpanao: District of Davao, Lake Lanao, Camp Keithley.
July, 1907, Mary S. Clemens.
Reported only from the Philippines.
POLYSTICTUS RUBRITINCTUS (Murr.) Sacc. & Trott.:
Syll. Fung. 21: 320. 1912.
Coriolus rubritinctus Murr. Bull. Torrey Club 35: 396. 1908.
Mrinporo: Mount Halcon, November, 1906, Merrill OIT7.
As yet collected only in the Philippines.
Saccardo,
GRAFF: PHILIPPINE BASIDIOMYCETES 467
POLYsTICTUS TABACINUS Mont.; Gay, Hist. Fisica Pol. Chili 7:
361. pl. 7, f. 6. 1845.
Polyporus tabacinus Mont. Ann. Sci. Nat. Bot. II. 3: 349.
1835.
Polyporus spadiceus Jungh. Flor. Crypt. Java 54. pl. 3o.
1838. Not P. spadiceus Berk.
Polystictus microcyclus Zipp.; Léveillé, Ann. Sci. Nat. Bot. III.
2: 188. 1844
Polystictus spadiceus Fr. Nov. Symb. Myc. 1851.
Polystictus xerampelinus Kalchbr. Grevillea 4:72. 1876.
Cycloporellus microcyclus Murr. Bull. Torrey Club 34: 468.
1907.
Polystictus substygius Bres. Hedwigia 53: 66. 1912.
Cyclomyces spadiceus Pat. Philip. Jour. Sci. 10: Bot. 95. 1915.
Cyclomyces tabacinus Pat. Philip. Jour. Sci. 10: Bot. 95. I9I5.
Luzon: Province of Laguna, Mount Maquiling, February,
1912, P.W. Graff, Bur. Sci. 15956, 15081.
Originally described from material collected in Juan Fernan-
dez, later from material collected in Chili, New Zealand and Java.
POLYSTICTUS TABACINUS var. barbatus (Murr.) comb. nov. [PLATE
15, FIG. 2.}
Cycloporellus barbatus Murr. Bull. Torrey Club 35: 397. 1908.
Polystictus barbatus Sacc. & Trott.; Saccardo, Syll. Fung. 21:
3%. tars
Polystictus sindieni var. barbatus Graff, Philip. Jour. Sci. 9:
Bot. 242. 1914.
Luzon: Province of Zambales, November—December, 1907,
H. M. Curran & M. L. Merrit, For. Bur. 8208 (type of Cyclo-
porellus barbatus Murr.).
As yet this variety has only been collected in the Philippine
Islands.
POLYSTICTUS TABACINUS var. substygius (Berk. & Br.) comb. nov.
Fomes substygius Berk. & Br.; Cooke, Praec. Polyp. 522.
Fomes spadiceus var. hal a Bres. Hedwigia 53: 59. 1912.
Luzon: Province of Rizal, February, 1911, M. Ramos, Bur.
Sci. 13467; Mindoro, Mount Halcon, November, 1906, Merrill
6114 (var. type).
468 GRAFF: PHILIPPINE BASIDIOMYCETES
There are two varieties of Polystictus tabacinus, which represent
two different sorts of development and variation from the type.
The first variety, barbatus, does not vary in pilear texture but in
the fact that the upper surface is covered with a long tomentum,
and has a somewhat darker shade of color. This tomentum varies
in length from short, at the outer margin, to about 3 mm., near
the place of attachment of the fungus.
In the case of the second variety, substygius, the variation is
one of which insufficient notice has been taken; namely, a tendency
toward the assumption of a perennial development in an occasional
species, which is typically annual, when growing under the warm
conditions of a tropical climate. This second variety has de-
veloped in this manner. The context of the plant develops much
thicker and harder than normally. The upper surface loses its
characteristic velvety tomentum, becoming hard and smooth.
Otherwise than this the plant retains its usual characters. To one
unfamiliar with the variations through which this plant may pass
the natural tendency would be to place it in the genus Fomes.
While pore layers have been occasionally observed in this variety
they are the exception rather than the rule.
Fomes spadiceus (Berk.) Cooke is not this species, but a quite
distinct plant.
Reported from Samoa, the Philippines, Malacca, and the East
Indies.
TRAMETES Fries
TRAMETES ELMERI (Murr.) Graff, Philip. Jour. Sci. 9: Bot. 243.
I9I4.
Tyromyces Elmeri Murr. Bull. Torrey Club 34: 475. 1907.
Polyporus Elmeri Sacc. & Trott.; Saccardo ,» oyll. Fung. 21: 279.
1912.
Luzon: Province of Bataan, Mount Mariveles, November,
1904, Elmer 6954, on dead wood.
The generic characters of this fungus are those of Trametes
rather than Polyporus. Saccardo evidently made the change on
general principles rather than after a critical study of the species.
As yet collected only in the Philippines.
TRAMETES INSULARIS Murr. Bull. Torrey Club 35: 405. 1908.
Luzon: Province of Benguet, Baguio, April, 1909, R. C. Me-
Gregor, Bur. Sct. 8311, on Pinus insularis Endl.
GRAFF: PHILIPPINE BASIDIOMYCETES 469
The limited distribution of pines makes this fungus very local
in the Islands. As this is but the second collection which has
been made to date, this form must be rare even in the pine regions.
Not reported outside the Philippine Islands.
UNIVERSITY OF MONTANA,
MissouLta, MONTANA
Explanation of plate 15
Fic. 1. Polyporus benguetensis (Murr.) Graff. Specimen from the herbarium
of the Bureau of Science, Manila, Merrill 5003, co-type material of Coltricia ben-
guelensis Murr
Fic. 2. Polystictus tabacinus var. barbatus (Murr.) Graff. Specimen from the
herbarium of the Bureau of Science, Manila, Curran & Merritt, For. Bur. 8208 co-type
of Cycloporellus barbatus Murr.
INDEX TO AMERICAN BOTANICAL LITERATURE
1914-1918
The aim of this Index is sto include all current botanical literature written by
Americans, published in Am or based upon American material ; the word Amer-
ica being used in the sicedtoat
eviews, and papers that ‘abel exclusively to forestry, agriculture, horticulture,
manufactured products of vegetable origin, or laboratory methods are not included, and
no attempt is made to index the literature of bacteriology. An occasional exception is
made in favor of some paper appearing in an American periodical which is devoted
wholly to botany. Reprints are not srinstioisd unless they differ from the original in
some important particular. If users of the Index will call the attention of the editor
to errors or omissions, their kindness will be appreciated.
This Index is reprinted monthly on cards, and furnished in this form to subscribers
at the rate of one cent for each card, Selections of cards are not permitted ; each
subscriber must take all cards published during the term of his subscription, Corre-
spondence relating to the card issue should be addressed to the Treasurer of the Torrey
Botanical Club,
Africa, E. M. The minimum Bordeaux application for the control of
Hemileia. Philip. Agr. Forest. 6: 251-271. My 1918.
Appleman, C. O. Physiological basis for the preparation of potatoes
for seed. cit hoses Agr. Exp. Sta. Bull. 212: 80-102. f. .I-T1.
F 1918.
Bailey, L. H. The indigen and cultigen. Science II. 47: 306-308.
29 Mr 1918.
Beal, W. J. Systematic botany. Ann. Rep. Michigan Acad. Sci.
1G: 237. 1077-
Beardslee, H. C. Michigan collections of Myxomycetes.
Michigan Acad. Sci. 19: 159-162. 1917.
Berry, E. W. Notes on the fern genus Clathropteris.
Club 45: 279-285. f. 1,2. 9 Au 1918.
Bessey, E. A. The phylogeny of the grasses.
Acad. Sci. 19: 239-245. 1917-
Britton, E.G. Musci. In Britton, N. L., Flora of Bermuda, 430-
448. 28 F 1918. [Illust.]
Britton, N. L. Flora of Bermuda.
[Illust.]
Includes contributions by Britton, E. G., Evans, A. W., Riddle,
F. J., and Howe, M. A., here indexed separately.
A471
Ann. Rep.
Bull. Torrey
Ann. Rep. Michigan
i—xi+1-585. 28 F 1918.
L. W., Seaver,
472 INDEX TO AMERICAN BOTANICAL LITERATURE
Brown, F. B. H. Flora of a Wayne County salt marsh. Ann. Rep.
Michigan Acad. Sci. 19: 219. 1917.
Brown, F.B. H. The forest associations of Wayne County, Michigan.
Ann. Rep. Michigan Acad. Sci. 19: 209-217. 1917.
Burkholder, W. H. The production of an anthracnose-resistant white
marrow bean. Phytopathology 8: 353-359. 17 Jl 1918.
Burnham, S. H. The flora of Indian Ladder and vicinity: together
with descriptive notes on the scenery. Torreya 18: 101-116.
f. 1-5. Je 1918; 127-149. f. 6-9. 15 Au 1918.
Coker, W. C. The lactarias of North Carolina. Jour. Elisha Mitchell
ci, Soc. 34: I-61. pl. -go. Je 1918.
Lactarius Allardii, L. subtorminosus, L. furcatus, L. coleopteris, L. Curtisii,
L. subplinihogalus, and L. lentus, spp. nov., are described.
Conzatti,C. Exploracion botanica por la Costa Meridional de Oaxaca.
Bol. Dir. Est. Biol. 2: 309-319. 1918. [Illust.]
Cook, M. T., & Martin, W. H. Leaf blight of the tomato. New
Jersey Agr. Sta. Circ. 96: [1-4]. 1 Ap 1918.
Dodge, B. O. Studies in the genus Gymnosporangium—II. Report
on cultures made in 1915 and 1916. Bull. Torrey Club 45: 287-300.
pl. 8. 9 Au 1918.
Dodge, C. K. Contributions to the botany of Michigan. Univ.
Michigan Mus. Zoo. Misc. Publ. 4: 1-14. 23 F 1918.
Edson, H. A., & Shapovalov, M. Potato-stem lesions. Jour. Agr.
Research 14: 213-220. pl. 24-26. 29 Jl 1918.
East, E. M., & Park, J. B. Studies on self-sterility—II. Pollen-tube
growth. Genetics 3: 353-366. f. 1-3. Jl 1918.
Evans, A. W. Hepaticae. In Britton, N. L., Flora of Bermuda.
448-469. 28 F 1918. [Illust.] :
Farwell, O. A. New species and varieties from Michigan. Ann. Rep.
Michigan Acad. Sci. 19: 247-249. 1917.
Farwell, O. A. Rare or interesting plants in Michigan. Ann. Rep.
Michigan Acad. Sci. 19: 251-261. 1917.
Fernald, H. T. The pine blister rust. Month. Bull. State Comm.
Hort. Calif. 7: 451-453. f. 60, 6r. Jl 1918.
Fernald, M. L. The American representatives of Equisetum sylvati-
cum. Rhodora 20: 129-131. 22 Jl 1918.
Fernald, M. L. Carex paupercula Michx., var. brevisquama, n. var.
Rhodora 20: 152. 1 Au 1918.
Fernald, M. L. Some allies of Rynchospora macrostachya. Rhodora
20: 138-140. 1 Au 1918.
- INDEX TO AMERICAN BOTANICAL LITERATURE 473
Fernald, M. L. The specific identity of Bidens hyperborea and B.
colpophila. Rhodora 20: 146-150. 1 Au 1918.
Fernald, M. L., & Wiegand, K. M. Some new species and varieties
of Poa from eastern North America. Rhodora 20: 122-127.
a2. 4b 3 ;
Includes Poa saltuensis and P. paludigena, spp. nov. _
Gardner, N. L. New Pacific coast marine Pepe Univ. Calif.
hue Bot. 6: 429-454. pl. 36, 37. 16 Jl 1918
Inc $ new species in Chlorogloea (1), Xenococcus (1), Dermocarpa (4), Hyella
(3), ce port (4)
Gladwin, F. E. A non-parasitic malady of the vine. New York Agr.
Exp. Sta. Bull. 449: 99-110. pl. 1-3. Mr 1918.
Goodspeed, T. H., & Davidson, P. Controlled pollination in Nico-
tiana. Univ. Calif. Publ. Bot. 5: 429-434. 10 Au 1918.
Harris, J. A. Further illustrations of the applicability of a coefficient
measuring the correlation between a variable and the deviation of a
dependent variable from its probable value. Genetics 3: 328-352.
Jl 1918.
Harter, L. L. A hitherto-unreported disease of okra. Jour. Agr.
Research 14: 207-212. pl. 23 +f. 1-3. 29 Jl 1918.
Harvey, L. H. Intra-microsporangial development of the tube in the
microspore of Pinus sylvestris. Ann. Rep. Michigan Acad. Sci.
19: 333-335. f. 16, 17- 1917.
Harvey, L. H. Polyembryony in Quercus alba. Ann. Rep. Michigan
Acad. Sci. 19: 329-331. 1917-
Hibbard, R. P. Physiological balance in the soil solution. Ann.
Rep. Michigan Acad. Sci. 19: 163-166. 1917.
Hillman, F. H. The agricultural species of bent grasses. Part II.
The seeds of redtop and other bent grasses. U. S. Dept. Agr. Bull.
692: 15-26. f. 3-11. 23 Jl 1918.
Howe, M.A. Algae. In Britton, N. L., Flora of Bermuda, 489-540.
28 F 1918.
pastas descriptions of Boodlea struveotdes,
, Seirospora purpurea, Ceramium leptozonum,
spp. nov., and new combinations, Zonaria zonalis,
encia Corallopsis, and Ptilothamnion bipinnatum.
- Jackson, H. S. The Uredinales of Oregon. Brooklyn Bot. Gard.
Mem. t: 198-297. 6 Jl 1918
described in catidopeetalh (1), Melampsora (x), Milesia (1),
Dasya Collinsiana, Callithamnion
and Nemastoma gelatinosum,
Gracilaria mammillaris, Laur-
ip
New
Puccinia @). and Uromyces (1).
474 INDEX TO AMERICAN BOTANICAL LITERATURE
Kauffman, C. H. Unreported Michigan fungi for 1915 and 1916, with
an index to the hosts and substrata of Basidiomycetes. Ann,
Rep. Michigan Acad. Sci. 19: 145-157. 1917.
Kirkwood, J. E. The conifers of the northern Rockies. Dep. Int.
Bur. Ed. Bull. 53: 1-61. f. 1-37. 1918.
Kirkwood, J. E. A Mexican hacienda. Nat. Geog. Mag. 536-584.
My 1914. [Illust.]
Contains some botanical information.
Kitchin, P.C. The effect upon the growth of some coniferous seedlings
of various conditions of shade and moisture. Ann. Rep. Michigan
Acad. Sci. 19: 337-356. pl. 17-23. 1917.
Krakover, L. J. The leaf-spot disease of red clover caused by Mac-
rosporium sarcinaeforme Cav. Ann. Rep. Michigan Acad. Sci.
19: 275-328... To-FO-- f2 947s: 1917.
Levine, M. N., & Stakman, E. C. A third biologic form of Puccinia
graminis on wheat. Jour. Agr. Research 13: 651-654. 17 Je 1918.
Macbride, J. F. A new Perezia adventive in Massachusetts. Rhodora
20: 150-152. 1 Au 1918.
Macbride, J. F. A new species of bladdernut. Rhodora 20: 127-129.
22 Jl 1918.
Staphylea Brighamii sp. nov.
MacCaughey, V. The Hawaiian Lehua. Am. Forestry 24: 409-418.
Jl 1918. [Illust.]
MacCaughey, V. The strand flora of the Hawaiian Archipelago—I.
Geographical relations, origin, and composition. Bull. Torrey
Club 45: 259-277. 9 Au 1918. :
Marsh, C. D. Stock-poisoning plants of the range. U. S. Dept. Agr.
Bull. 575: 1-24. pl. 1-30. 23 Jl 1918.
Martin, W. H. Dissemination of Septoria lycopersici Speg. by insects
and pickers. Phytopathology 8: 365-372. 17 Jl 1918.
Massey, L. M. The diseases of roses.
Trans. Massachusetts Hort.
Soc. 1918: 81-101. pl. z, 2. 1918.
Matz, J. Diseases and insect pests of the pecan. Florida Agr. Exp.
Sta. Bull. 147: 135-162. f. 45-93. My 10918.
Maxon, W. R. Polystichum Andersoni and related species. Am.
Fern Jour. 8: 33-37. 6 Au 1918,
Includes Polystichum alaskense sp. nov.
McKee, R. Purple vetch. U.S. Dept. Agr. Farm. Bull. 967: 1-12.
Ff. &-6,., Je 1918.
INDEX TO AMERICAN BOTANICAL LITERATURE 475
Merrill, E. D. Species Blancoanae. A critical revision of the Philip-
pine species of plants described by Blanco and by Llanos. Manila
Dept. Agr. Bur. Sci. Publ. 12: 1-423. pl. 1. 15 Je 1918.
Merriman, C. H. Two new manzanitas from the Sierra Nevada of
California. Proc. Biol. Soc. Washington 31: 101-104. pl. 2-5.
10 Jl 1918.
Arctostaphylos newukka and A. nissenana, spp. nov.
Nash, G. V. Injury to evergreens. Jour. N. Y. Bot. Gard. 19: 159-
164. Jl 1918.
Nichols, G. E. The vegetation of northern Cape Breton Island, Nova
Scotia. Trans. Connecticut Acad. Arts and Sci. 22: 249-467.
f. 770. Je 1918.
Noriega, J. M. EI Cuapinole. Bol. Dir. Est. Biol. 2: 357-363.
7 2a 26r8,
Ochoterena, I. Una nueva especie de Mamillaria. Bol. Dir. Est.
Biol. 2: 355, 356. f. 1-3. 1918.
Pennell, F. W. A botanical expedition to Colombia. Jour. N. Y-
Bot. Gard. 19: 117-138. pl. 213, 214. Je 1918.
Piper, C. V. The agricultural species of bent grasses. Part I. Rhode
Island bent and related grasses. U. S. Dept. Agr. Bull. 692: I-14.
fede mee gat 168. :
Rhoads, A. S., Hedgcock, G. G., Bethel, E., & Hartley, C. Host re-
lationships of the North American rusts, other than gymno-
sporangiums, which attack conifers. Phytopathology 8: 309-352.
17 Jl 1918.
Riddle, L. W. Lichens. In Britton, N. L. Flora of Bermuda,
470-479. 28 F 1918.
Rigg, G. B. Some energy relations of plants. Science II. 48: 125-132-
9 Au 1918.
Rugg, H.G. A Vermont fern garden 8: 50-52. 6 Au 1918.
Sargent, C. S. Charles Edward Faxon. Rhodora 20: 117-122. 22
Jl 1918.
Sax, K. The behavior of the chromosomes in fertilization. Genetics
3: 309-327. pl. r, 2. Jl 1918.
Seaver, F. J. Fungi. In Britton, N. L., Flora of Bermuda, 489-540.
28 F 1918.
Seeley, D. A. The relation between temperature and crops. Ann.
Rep. Michigan Acad. Sci. 19: 167-196. Pl. 5. +f.7,8. 1917.
476 INDEX TO AMERICAN BOTANICAL LITERATURE
Shamel, A. D., Scott, L. B., & Pomeroy, C. S. Citrus- fruit improve-
ment: a aiaiy of bud variation in the Valencia orange. GS
Dept. Agr. Bull. 624: 1-120. pl. 1-14 + f. 1-9. 25 Jl 1918.
Shive, J. W., & Martin, W. H. A comparative study of salt require-
ments for young and for mature buckwheat plants in solution cul-
tures. Jour. Agr. Research 14: 151-175. f. 1-3. 22 Jl 1918.
Shufeldt, R. W. Flower and other studies for the summer of 1918.
Am. Forestry 24: 433-438. f. r-zz. Jl 1918.
Stakman, E. C., Parker, J. H., & Piemeisel, F. J. Can biologic forms
of stemrust on wheat change rapidly enough to interfere with
breeding for rust resistance? Jour. Agr. Research 14: I1I—124.
Pi T7277. LOTS.
Stewart, V. B. Exclusion legislation and fruit tree production. Phy-
topathology 8: 360-364. 17 Jl 1918.
Sutton, J. M. Flora of the Detroit zoological tract. Ann. Rep.
Michigan Acad. Sci. 19: 263-271. f. 13. 1917.
Tanaka, T. Notes on some fungous diseases and a new coddling moth
attacking the persimmon in Japan. Month. Bull. State Comm.
Hort. Calif. 7: 461-463. Jl 1918.
Taylor, N. Effects of the severe winter on the woody plants in the
Garden. Brooklyn Bot. Gard. Rec. 7: 83-87. Jl 1918.
Teesdale, C. H., & MacLean, J.D. Tests of the absorption and pene-
tration of coal tar and creosote in longleaf pine. U.S. Dept. Agr.
Bull. 607: 1-42. pl. 1-11 + f. 1-14. 7 Je 1918.
Van Pelt, W. Some important clover diseases in Ohio. Month. Bull.
Ohio Agr. Exp. Sta. 3: 239-243. f. 1, 2. Au 1918.
Waterman, W. G. Ecology of northern Michigan dunes: Crystal
Lake Bar region. Ann. Rep. Michigan Acad. Sci. 19: 197-208.
pl. 6-11 +f. O-1I. 1917.
Waugh, F. W. Wild plants as food. Seevws Nat. 32: 2-6. f. 1-7:
Ap 1918.
Waynick, D. D. The chemical composition of the plant as further
proof of the close relation between antagonism and cell permeability.
Univ. Calif. Publ. Agr. Sci. 3: 135-240. pl. 13-24 +f. 1-26. 12Jl
1918.
Wiegand, K. M. Some species and varieties of Elymus in eastern
North America. Rhodora 20: 81-90. 1 My 1918.
Elymus riparius sp. nov. is described.
Wiegand, K.M. A new variety of Triosteum aurantiacum. Rhodora
20: 116. 11 Je 1918.
BuLL. TORREY CLU
r VOLUME 45, PLATE 13
A. ENCALYPTA EXTINCTORIA (L.) Sw.
B. ENCALYPTA RHABDOCARPA SCHWAEGR.
C. ENCALYPTA ALPINA SMITH :
D. ENCALYPTA LACINIATA (Hepw.) Lrivpe.
BuL_. TORREY J
CLUB VOLUME 45, PLATE 14
=
2S
—
'
ENCALYPTA APOPHYSATA Nees & HoRNSCH,
ENCALYPTA a ae
ENCALYPTA PROCERA B
ENCALYPTA CONTORTA poe L:NDB.
ad
Buti. TORREY CLUB
b VOLUME 45, PLATE I5
[ \Q < - baat .
1. POLYPORUS BENGUETENSIS (Murr.) GRAFF
2. POLYSTICTUS TABACINUS var. BARBATUS (Mcurr.) GRAFF
Vol. 45. No. 12
BULLETIN
OF THE
TORREY BOTANICAL CLUB
DECEMBER, 1918
Notes on plants of the southern United States—IV
FrRANciIS W. PENNELL
THE GENUS CROTONOPSIS
[In 1803 Michaux published in his “Flora Boreali-Americana”’
a new genus of plants to which, from its evident likeness to Croton,
he gave the name of Crotonopsis. The genus has been maintained
continuously from that time, although twice have attempts been
made to rechristen it. In 1826 Sprengel, without comment,
proposed to substitute Friesia, and twelve years later Rafinesque,
_ remarking that ‘‘the name previously given was absurd and incor-
rect,’ announced his Leptemon. According to current rules of
nomenclature Crotonopsis must be held.
Michaux collected Crotonopsis twice, and, in the plate which
accompanies the description of his species C. linearis, he fortunately
figured both the specimens obtained. The drawing to the left
hand shows a low plant with lanceolate or elliptic-lanceolate
leaves and with fruits few and axillary, that to the right is of a
side-branch of a taller plant with narrower longer leaves and with
conspicuously elongated spikes. Two localities are cited in the
text, Long-bay, Carolina, and the Illinois region. From evidence
to be presented it is clear that two species are illustrated, and,
from the form of its leaves, that to the right is the one entitled
to the name C. linearis. Fortunately it is also that from the
locality first cited. True C. linearis Michx. is a plant restricted
to the Coastal Plain of the southeastern states.
[The BULLETIN for November (45: ain ar pl. 13-15) was issued November 15,
478 PENNELL: PLANTS OF SOUTHERN UNITED STATES
In 1805, but two years later, Willdenow added a second species,
Crotonopsis elliptica. His plant is stated to differ from the linear-
leaved C. linearis Michx. in its leaves being elliptic, rounded to
each end, and in its spikes being shorter. Evidently C. elliptica
is our oldest name for the widely-ranging northern and inland
species. As this plant extends to the Gulf coast, incidentally
overlapping the range of C. linearis, the type-region, ‘‘Carolina,”
is well within its normal range.
The later history of the genus may be briefly sketched. Pursh
in 1814 combined both species, though as varieties, in one, his
C. argentea. Nearly to the close of the past century the genus
was uniformly considered monotypic. But in 1895 Nash, from a
single collection from Florida, added a second species, C. spinosa.
As a matter of fact he was actually recharacterizing Michaux’s
C. linearis, laying primary emphasis upon newly discovered
features of the fruit.
Spikes short, of but one or two fruits. Staminate flowers less than 1 mm. in diameter;
ents shorter than the sepals, and but little longer than the anthers. Frui
ovoid, with an evident median vein on each side; scale-like hairs on fruit with
broad brown disk, umbonate to tuberculate-raised, even ge ane into a gate
spine, and with its margin of are tively uniform closely appressed white
nh 1
—3 cm. long; stellate pin on upper sur-
cs with long rays which overlap those of adjoining hairs. Plant weer! 1-5 dm.
na Sie Nee
Spikes ares slender, of three to six fruits. Staminate flowers more ee Im
; filaments longer than the sepals, and much longer than the Bie:
Fruit pane: without evident vein on the side; scale-like
inane disk, which is usually raised
hairs on fruit with
into a decided tubercle or spine, and with
tellate not closely appressed — brownish rays. Leaves
Rip 2-4 cm. long; stellate hairs on upper surface with short rays
which do not overlap those of adjoining hairs. ma usually 4-8 dm. tall.
i Gs linearis.
I. CROTONOPSIS ELLIPTICA Willd.
Crotonopsis elliptica Willd. Sp. Pl. 4: 380. 1805. ‘‘Habitat in
Carolina.”
Crotonopsis argentea elliptica Pursh, Fl. Am. Sept. 1: 206. 1814
Dry sandy soil, Connecticut to northern Florida, west to
eastern Kansas and central Texas; northward in or near the
Coastal Plain, southward mostly inland, on granitic rocks of the
Piedmont and southern Appalachian regions. Numerous speci-
mens seen. The most southwestern studied are from sandy post-
PENNELL: PLANTS OF SOUTHERN UNITED STATES 479
oak woods, Sheridan, Colorado County, Texas, my number 5533,
and are unique in that the plants were uniformly 7-8 dm. tall,
and the fruit relatively large and somewhat brownish instead of
being nearly black.
2. CROTONOPSIS LINEARIS Michx.
Crotonopsis linearis Michx. Fl. Bor. Am. 2: 186. pl. 46 p.p.
1803. ‘Hab. in maritimis Carolinae, juxta Long-Bay, et in
regione Illinoensi.’”” Two plants figured. One is a plant with
lanceolate-linear leaves and slender spikes, the other with
lanceolate to ovate-lanceolate leaves and flowers one to two
together. As Illinois specimen certainly the latter, and as the
former is known in the maritime region of Carolina, and is the
plant to which the name /inearis better applies, this is selected
as the type.
Crotonopsis argentea Pursh, Fl. Am. Sept. 1: 206. 1814. Con-
sists of two varieties; name is here applied to first.
Crotonopsis argentea linearis Pursh, I. c.
Friesia argentea Spreng. Syst. 3: 850. 1826. Without citation
of Pursh.
Leptemon lineare Raf. Sylva Tellur. 67. 1838.
Crotonopsis spinosa Nash, Bull. Torrey Club 22: 157. 1895.
“Collected by Mr. W. T. Swingle [13974] at Dunellon [Florida],
July 24 [1894].”’ Type seen in the herbarium of Columbia
University at the New York Botanical Garden.
Dry sandy soil, in the Coastal Plain, South Carolina to central
Florida and eastern Texas, extending inland near the Mississippi
River to southern Illinois and southeastern Missouri, and in
Texas to Dallas.
SoutH Carona. Beaufort: Bluffton, huss (M).
Charleston: Mt. Pleasant, L. R. Gibbes (Y).
Grorcia. Lowndes: Olympia, R. M. Harper 1593 (M, U, Y).
Mitchell: R. M. Harper 1168 (M, U, Y).
FLoripa. Baker: Macclenay, L. H. Lighthipe 586 (Y).
Brevard: Melbourne, Curtiss 5715 (M, U, Y). Escambia: Pen-
sacola, J. M. Macfarlane (P). Hillsboro: Tampa, A. P. Garber
(U). Jefferson: Hitchcock (M). Lake: Eustis, Nash ro7z (M,
U, Y).. been: T Tallahassee, N. K. Berg (Y). Marion: Dunnellon,
480 PENNELL: PLANTS OF SOUTHERN UNITED STATES
Swingle 1397a (U, Y). Orange: A. Fredholm 5389 (Y). Pinellas:
Ozona, F. L. Lewton (Y). St. John: St. Augustine, M. C. Rey-
nolds (P). Sumter: Wildwood, H. J. Webber (M). Suwanee:
Live Oak, Curtiss 6897 (M, U, Y).
Mississippi. Oktibbeha: Starkville, S. M. Tracy (M).
IniinoIs. Mason: “sandy barrens,”’ E. Hall (M).
MissouriI. Dunklin: Campbell, Bush (M). Stoddard: Bush
Texas. Dallas: Dallas, Reverchon 869 (M, Y), 3177 (M),
4306 (M,U). Waller: Hempstead, E. Hall 575 pi tet, Y).
MISCELLANEOUS RECORDS
HyPERIcuM opacum T. & G.
Collected in flower August 15, 1912, in moist long-leaf pine-
land, Ozone Park, St. Tammany Parish, Louisiana, my number
4210.
LecHeaA LEGGEtTTI Britton & Hollick
Collected in fruit August 14, 1912, in dry open long-leaf pine-
land, Abita Springs, St. Tammany Parish, Louisiana, my number
- 4162. Wide-spread through the southeastern states, reaching
Florida and Louisiana.
RHEXIA LUTEA Walt.
Collected in fruit August 14, 1912, in moist long-leaf pineland,
Abita Springs, Louisiana, my number 4198.
Ru#eEx1A Nasu Small
Collected in flower August 16, 1912, moist sandy soil near
Mandeville, St. Tammany Parish, Louisiana, my number 4239.
Through the long-leaf pineland of the Coastal Plain, North
Carolina to Florida and Louisiana. When seen living, readily
distinguished from R. mariana L. by its flowers, the petals of
which are much larger, 18-25 mm. long, and deep purple-pink.
Rhexia interior Pennell, nom. nov.
Rhexia latifolia Bush, Rhodora 13: 167. I911. Not Rhexia
latifolia Aubl. Pl. Gui. 1: 336. 1775. Aublet’s plant is not
retained in the genus Rhevia as today understood, a fact which
PENNELL: PLANTS OF SOUTHERN UNITED STATES 481
under the Vienna Code permits the repetition of the same
specific name.
Collected in fruit September 8, 1913, moist shady soil, west of
Sapulpa, Creek County, Oklahoma, my number 5 380.
The following key to the species of Rhexia is offered:
Anthers relatively short, oblong, straight, not spurred at base.
Pet llow. Leaves lanceolate. 1. R. tulea Walt.
Petals pink-purple. Leaves ovate.
Hypanthium glabrous. Upper surface of the leaves hirsute,
. petiolata Walt.
Hypanthium glandular-hirsute. Upper surface of ee leaves glabrous.
3. R. serrulata Nutt.
Anthers longer, linear, curved, spurred at the base.
Anther-sacs very slightly spurred. Petals less than 10 mm. long, white.
4. R. parviflora Chapm.
Anther-sacs evidently spurred. Petals more than 10 mm. long, pale pink (or
white in R. lanceolata) to pink-purple.
Leaves membranous, green; lateral nerves not close to the margin; upper
surface of some or all leaves hirsute. Hypanthium glandular-hirsute
to glabrous
Apex of hypanthium not lanose. Calyx-lobes less than one-third
length of hypanthium
Stem aeeig if at all ie internodes conspicuously hirsute.
datb Neck of hypanthium equaling
or sey aida shorter than the body.
Petals sa to white, 10-15 mm.long. Buds with sepal
tips mostly s thine ed.
Leaves clliptic-ovat o lanceolate, all evidently three-
meee pert over 2 cm. long, the lower with a
more iolar
Leaves pe ae may on evident Aicoes 3-4 mm.
long. Petals about 10 mm. long
ee cae Small.
Leaves lanceolate to elliptic-lanceolate, on_ ill-
defined eeosiaeeg Petals 12-15 mm
6. R. mariana L.
Leaves narrowly-lanceolate to linear, only the main
stem-leaves if any three-veined, mostly less than 2
cm. long, sessile. Petals 10-12 mm. long
oR lucedinta Walt.
Petals Soni Se aa Buds with sepal-tips mostly
appressed.
eet 1825 mm. long. oe onatepma Io-14 mm. long,
ck equaling the body. Stem obscurely angled.
aoe ‘Wont si epee lateral nerves ob-
ensis Grise
482 PENNELL: PLANTS OF SOUTHERN UNITED STATES
Leaves lanceolate, conspicuously hirsute, lateral
nerves pe nent. 9. R. Nashii Small.
Petals 10-14 m g. Hypanthium 7-8 mm. long,
neck slightly oe than the body. Sica! eee
sharply angled. 10. R. interior Pennell.
Stem conspicuously wing-angled, the internodes sparsely hirsute
to glabrous aves rounded at base, the upper clasping.
Neck of hypanthium much shorter than the body. Petals
eep purple-pink. :
asics of stem sparsely hirsute. Leaves ovate. Hypan-
thium hirsute. Il. R. virginica L.
Internodes of stem glabro Leaves sh oo Reg
Hypanthium sparsely eae ar-pubescent to co)
. R. stricta Pursh.
Apex of hypanthium lanose with oe Sieh cs hairs. Calyx-lobes
more than one-half length of hypanthiu
13. R. aristosa Britton.
Leaves firm, glaucescent, lateral nerves following closely the margin of the
blade; upper surface glabrous. Hypanthium red ea
cent. R. Alifanus Wal
RAIMANNIA Drummonpt (Hook.) Rose
Collected in flower October 12, 1912; beachs and, Sullivan
Island, Charleston County, South Carolina, my number 4857.
Extensively spreading from Texas along the coastal dunes of the
southeastern states.
MYRIOPHYLLUM PROSERPINACOIDES Gill.
Pool in long-leaf pine-land, Mandeville, St. Tammany Parish,
Lousiana, my number 4205. Established also in ditches at
Houma, Terre Bonne Parish. Naturalized from Chile.
The strand flora of the Hawaiian Archipelazo—ll, Ecological relations
VAUGHAN MACCAUGHEY
TEMPERATURE
The Hawaiian littoral zone is characterized by relatively warm
and uniform thermal conditions throughout the year. Sudden
fluctuations are exceedingly rare and are never of great magnitude.
The lowest recorded littoral temperature is 47° F., the highest is
98° F.; the mean of all littoral temperatures is 74° F. The mean
monthly temperature at Honolulu, which may be taken as a repre-
sentative coastal station, varies from about 70.5° F. to 76.8° F.,
in January and July respectively.
The following table, arranged from data of the Hawaiian
Section, U. S. Weather Bureau, will show the temperature con-
ditions (for 1915) of a number of littoral stations on the various
islands. Data are not available from more strictly littoral sta-
tions—i. e., at the actual beach line itself. This is a problem that
awaits future field investigation.
; Temperature
‘ Elevation
goes aa feet Mean | Max. Min.
KMAUAE: Mane. o i ee a a 30 73.9" 93° x?
PAA eos rss eee es I5 73.3 88 47
OaHU Faheke coc 25 76.6 | 88 60
Honoltla: 65 <2 ws ee FII FER oP Oy 58
US. Mag. Sta oe 45 Yeg. | 92 50
Waianne. 2.5 6<-....-55: 6 76.2 93 52
Waialua Mill........-.. 30 74.3 93 52
Motoxar: Kalawao.........---+-: 70 75.2 57
Bganapalic. 20.06 eee 12 72.8 OI 55
PIQUA. cies 2 ease 145 75.6 89 58
Hawai: Mahukona...........-- II 75-5 98 58
i Ae ee as 200 73-5 87 58
Pepeekeo.......:-+---+> 100 93.3 86 57
BiG oe 100 72.6 87 54
TGDONO. - ss oie en ws IIo 72.8 89 58
Olsson-Seffer (’10) presents data to show that on tropical and
subtropical coasts the variations in the temperature of sea-water
are mainly due to changes in the direction of the winds and cur-
484 MAcCCAUGHEY: THE STRAND FLORA.
rents. Close to the shore, or where the water is shallow, the tem-
perature of the water is higher when the surface is calm, but low
when the sea is rough. This is the natural consequence of the
solar radiation in the former case, and of the mixing by the waves
of the surface water with the cooler water from below when the
sea is disturbed.
The annual thermal ranges of the oceanic waters of four repre-
sentative regions will make a significant contrast with the condi-
tions prevailing in Hawaiian waters: Sydney Harbor, Australia,
55-8°-72.4° F.; San Francisco Bay, California, 42°-69° F.;
Woods Hole, Massachusetts, below freezing—70° F.; Plymouth,
England, 44.1°-58.9° F.
The great oceanic current from the northeast, which travels
down the Pacific Coast and out past Hawaii, as part of the Equa-
torial Drift, has so profound an effect upon the Hawaiian climate
in general and the littoral zone in particular, that it merits special
consideration here. Dr. Sereno O. Bishop, who made Hawaii his
home for many years, writes (’04) of this current, as follows:
at remarkable stream of cold water, which flows ina vast stream southerly,
skirting southeast Alaska, Vancouver’s Island, the Pacific States of Washington,
Oregon, and California, and finally out westward to Hawaii, beyond whic
group it becomes merged into the great equatorial current running westward.
This stream is of very low temperature, of immense volume, and of great velocity.
It is unique in its powerful effects upon the climates of the coasts along which it
flows. . . . Finally turning westward like the trade winds under the impulse of the
earth’s rotation, this mighty stream broadens out into the open ocean, gradually
gaining warm
Dall ( ‘04) states:
As it moves down the coast it loses its heat and prod th d fogs of the
Oregonian region, cooling off so that when it peaches the latitude of the ‘eetien Gate
in late winter. This
imparts to that favored group a uniformly subtropical climate such as is unknown to
any other land in the same latitude.
Cowles, in describing the strand of the Lake Michigan dune
region (’99, p. 107), states that on the beach, due to the ‘‘absence
of vegetation and the general exposure . . . the temperature is
higher in summer and lower in winter than in most localities.
This great divergence between the temperature extremes is still
OF THE HAWAIIAN ARCHIPELAGO 485
further increased by the low specific heat of sand.”’ This is also
true of the Hawaiian sand strands. There isa greater temperature
range on the beach itself than in the protected zone lying immedi-
ately behind the beach. However, the temperature range of the
littoral is insignificant when contrasted with that of the mountains
that ascend directly from the lowlands, and in many places directly
from the littoral. The diurnal range in temperature increases as
one ascends the mountain slopes, and this range reaches its maxi- °
mum on the high summits of Kea and Loa (nearly 14,000 ft.).
Guppy (’03—’06) found the mean daily range of temperature on
the summit of Loa, August, 1897, to be 30.6° F.; the lowest reading
was 15°, the highest 61.2°.
Although sand has low specific heat, the upper dry layer becomes
excessively hot under a cloudless sky. Cockayne (’11) records
surface temperatures of 120°-127° F.on the New Zealand strand;
these figures are even higher than the generalization made by
Guppy in the table given below. It should be noted, however,
that the wet underlayers of sand absorb heat much more slowly.
At the depth of a few inches below the surface the sand is always
moist, so that the roots of sand-strand plants descend very quickly
into relatively cool soil.
Guppy (’03~’06) makes the following generalizations concern-
ing beach temperatures, the data applying to ordinary beaches
under an unclouded sky, in the hot season, during the early after-
noon:
Surface—half-inch deep Four inches deep
0°-105°
Temperate about 50°-55° lat. ae)
Subtropical 80 235 105 —I10 80
Tropical 270 =20 = =< II0 —120 85
Olsson-Seffer (’09, pp. 88, 89) gives an extensive series of
strand temperatures secured by him in various parts of the world.
With reference to temperature, Hawaiian strand regions may
be classified as follows:
I. Warmer strands—leeward, facing SE., S., or SW.
1. Typical leeward beaches: e. g., Mana, Barber’s Point, leeward
Molokai, Lahaina, and Kawaihae.
2. Slope approaches plane perpendicular to incident sunlight:
flat beaches, like those of Midway, Laysan, leeward Kauai
and Puna.
486 MacCauGHEY: THE STRAND FLORA
3. Not subjected to shadows: as for 2.
4. Heat reflected by neighboring objects: e. g., Mana-dune strand,
Koko Head tufa cliff coast, Mokapu.
5. Composed of dark materials which absorb and retain heat:
lava beaches of Hawaii, Maui, and parts of Oahu and
Kauai.
6. Texture unfavorable for rapid evaporation of moisture: mud
beaches and tufa cliff beaches, e. g., Pearl Harbor Inlet and
Mokapu.
7- Arid or semiarid: not receiving the cooling effects of rain,
waterfalls, etc., e. g., southwestern coasts of Maui and
Hawaii.
II. Cooler strands—windward, facing NE., N., or NW.
8. All of the windward beaches.
9. Slope more or less precipitous: high beaches, e. g., Hamakua
coast, windward East Molokai, Napali and Nihoa.
: ILLUMINATION
The brilliant illumination of the Hawaiian strand is one of its
most distinctive ecological features. The intense light of open
beaches as contrasted with other regions, has been commented
upon by ecologists in various parts of the world, but nowhere is
this better exemplified than in the Hawaiian Islands. On the low
islands the sky is cloudless, except during the infrequent rains.
On the high islands the clouds heap over the mountainous interior,
leaving the peripheral strand zone almost continuously exposed.
The total insolation, in diurnal or in annual terms, is therefore ©
exceedingly high. The Hawaiian coast, with its excessive inso-
lation, may be contrasted, for example, with the gray, foggy coast
of Washington and Oregon. On the coral and tufa beaches the
intensity of the direct illumination js greatly increased by reflec-
tion. The glare on a coral beach, during the middle part of the
day, as almost as intolerable to the eyes as that from a snow-field.
There are few data as to the direct and indirect effects of ex-
cessive insolation, save as an integral part of the xerophyte-pro-
ducing complex. In general, light retards growth, and too great
an intensity of light causes cessation of the growth of an organ.
Pfeffer (’03, p. 87) states:
OF THE HAWAIIAN ARCHIPELAGO 487
The internodes become shorter and the plant more condensed as the intensity of
the light increases, while the leaves attain their maximal size at a certain medium in-
tensity of illumination. This latter is owing to the fact that moderate light stimu-
lates the growth of the leaves, whereas intense light retards it
Jost (’07, p. 125) makes the following statements:
are not accurately acquainted with the precise way in which assimilation is
light; it is questionable, however, whether this is the rule with higher intensities . . .
may be easily imagined that a further increase in assimilation, following on in-
crease in light, is impossible owing to the deficiency in carbon-dioxide. Carbon-
dioxide may be present in sufficient quantity under ordinary circumstances to em-
ploy all the energy of sunlight, but when light is aNd increased the usual
amount of carbon dioxide would constitute a sub-optim
Schimper (’03, p. 58) states as a result of excess 2 aye
£ éhai
Terrestrial plants . . . frequently suffer f a Ee ae
chlorophyll. The veemasion of very sunny spots is never pure grea; but always
exhibits an admixture of yellow and brown tints due to the products of decompo-
intense tropical light may even completely bleach the
sition of chlorophyll...
foliage.
Many of the Hawaiian beach plants are grayish or yellowish
green. This is characteristic, not only of Hawaii, but also of other
strands. As Cockayne (’11) aes
The yellow colour of
is doubtless correlated
heredity of an acquired character.
The author cannot share the latter view, as beach plants which
have happened to grow in the shadedevelop normal green pigment,
instead of the bleached beach pigment. Such species as Santalum
Freycenetianum var. littorale, Lepidium owathense, Euphorbia
cordata, Batis maritima, Argyreia tiliaefolia, and Cressa cretica fur-
nish excellent examples of this pronounced difference between the
sun leaves and shade leaves of littoral plants.
A further comparison may be made which will illustrate the
intensity of the strand illumination. In the rain-forest and
summit-bog zones, and in the deep ravines of the lower and middle
forests, the ordinary illumination on cloudy days—and these are
regions of almost continuous cloudiness—necessitates a photo-
graphic exposure of say three minutes, whereas the same subject,
distance, and aperture on the strand would require but one
seventy-fifth of a second. The difference in illumination indi-
ey L 43 or es sae
with excess of ‘light, onde seems to me a possible example of
488 MacCauGHEY: THE STRAND FLORA
cated by these figures is thus about 13,500, that is, the beach
illumination is approximately 13,500 times as great as that of the
cloudy rain-forest. Of course, these figures do not include all of
the light-factors involved, but they are sufficient to indicate the
great differences in the illumination of regions not far removed
from one another.
PRECIPITATION
All precipitation on the Hawaiian littoral occurs in the form of
rain, dew, and rarely hail; snow is unknown. The strand is char-
acteristically xerophytic or semi-xerophytic, as contrasted with
the mesophytic lowland areas, and the montane rain-forests.
The following data from representative stations of the U. S.
Weather Bureau, 1915, will show these differences, the amount of
rainfall being given in inches:
Island Littoral station gsr a pag A Be 6
MRR See oN 16.98 38.02 75-52
Oahu Bim Sieh aly Vee cers Oe eae 18.99 42.29 II4.22
Maui (914) ee wh bie owes 4 20.91 42.70 397.26
MAMAN slates ebay we ae a 8.41 39.10 159-60
A few littoral stations are mesophytic or hygrophytic, but this is a
relatively unusual condition—e. g., Na Pali coast of Kauai, north
coast of East Molokai, and Hamakua coast of Hawaii.
The relationship between the annual precipitation on the strand
and the character of the vegetation is very close. The paucity of
the Hawaiian arborescent strand flora is undoubtedly due in part
to the xerophytic character of the littoral. In those few strand
regions which do possess sufficient rainfall, the forest extends down
to the beachline. The Puna, Hilo, and Hamakua regions, illus-
trate this condition. Schimper (’03, p. 407) makes the generali-
zation:
On Hee £.
ann te Aiat eine
rainfall they are almost the only ones. The close wood!
all?
AUR ad Siidis
In the case of the Hawaiian littoral vegetation, much emphasis
must be laid upon the exterminating and limiting agencies which
OF THE HAWAIIAN ARCHIPELAGO 489
have been introduced into the islands within historic times. The
ravages of wild live-stock, such as cattle, goats, sheep, and swine;
the clearing of the lowlands for agricultural and other purposes;
the building of roads; the large quantities of firewood which were
drawn from the nearest and most easily available sources; the
introduction of a great variety of pernicious foreign weeds—all
of these factors have contributed largely to the depletion of the
lowland and littoral floras, and have given them an aspect of
meagerness that they probably did not possess in their primitive
state. Man and his domestic animals have been much more potent
limiting factors than has been precipitation.
WIND ACTION
The wind is a powerful agency in its direct mechanical effects
upon beach vegetation. Many strand plants have forms that
are more or less protective, i. e., prostrate, creeping, rosette, or
hemispherical aerial bodies. ‘Plants of upright habit are per-
manently deflected and shaped by the wind; the windward branches
are stunted and warped, and growth takes place chiefly on the lee-
ward side of the plant. Seaside plants of Acacia farnensiana,
Prosopis juliflora, Santalum Freycinetianum var. littorale, Calophyl-
lum inophyllum, and Kadua littorale commonly illustrate this
condition. These wind-beaten plants are invariably dwarfed.
The mechanical effect of the wind is greatly augmented on
those coasts upon which it is able to pick up quantities of beach
sand. At storm times, in such regions, the wind becomes a veri-
table sand-blast. The evidences of this sand-blast action, upon
the trunks of both living and dead trees, and upon the local topog-
raphy itself, are familiar to all who have travelled along a windy
coast. The fantastic sculpturing of the seaward slopes of tufa in
the Koko Head region, and at Mokapu Peninsula, admirably
illustrate this sand-blast work. The herbaceous vegetation on
these slopes is either prostrate or rosette, ©. g., Sesuvium Portula-
castrum, Argyreia tiliaefolia, Boerhaavia diffusa; or tough and wiry,
e. g., Sporobolus virginicus, Fimbristylis pycnocephala.
On the Hawaiian littoral the wind is not as important an
ecologic agent as in such a region as the Lake Michigan sand dunes.
Here, according to Cowles (’99, p. 108), it ‘‘is the chief destroyer
490 MacCAuGHEY: THE STRAND FLORA
,
of plant societies,’ acting in two ways—either by undermining
plant individuals and groups, or by burying them with dune sand.
Neither of these processes is particularly conspicuous along Ha-
waiian coasts; the vegetation is not sufficiently luxuriant to em-
phasize the former, nor are the dunes of sufficient size or mobility
to give much importance to the latter. Here and there, however,
both of these processes may be witnessed.
On the whole, the Hawaiian beach winds are retarding rather
than destructive agencies. The Hawaiian strands may be divided
into the following classes, based upon the relative exposure to wind
action:
1. Shores exposed to prevailing winds.—The windward strands
are much more exposed than are those along the leeward sides of
the islands. Excellent contrasts are: the Hamakua and South
Kona shores of Hawaii; the northeast and southwest shores of
Haleakala; the north and south coasts of East Molokai; the Koo-
lau and Waianae shores of Oahu; the Kilauea and Kekaha shores
of Kauai.
2. High or promontory-like shores.—These are more exposed
than are low flat shores. Good examples are: Hamakua, Hawaii;
Hana and Kaanapali, Maui; the great pali of Molokai; Makapuu
and Kaena, Oahu; Napali, Kauai; and the cliffs of Nihoa.
3. Shores devoid of surface irregularities or vegetation sufficient
to break the force of the wind.—Low, flat shores, not protected by
mountains behind them nor covered with forest, are exposed to
the full force of the wind. In this class belong the coral atolls,
and such shores as Kahuku and Mokapu, Oahu, the western ex-
tremity of Molokai, the Maui isthmus, and the extreme south
point of Hawaii.
TRANSPIRATION
This is unquestionably the most important single physiological
factor in determining the character of the Hawaiian strand flora.
Only those plants which possess comparatively low transpiratory
rates are able to permanently establish themselves upon the strand.
Those species which invade the strand from the interior, and are
characteristically mesophytic, undergo marked structural changes
when they appear as members of the strand association.
OF THE HAWAIIAN ARCHIPELAGO ~ 491
The importance of the evaporation factor, particularly in the
early stages of an association, is admirably emphasized by Tran-
seau (’08, p. 230), who states:
The greatest decrease in the demands for transpiration on the part of seedlings
takes place during the first stages. This greatly aids in accounting for the well-known
called to the importance of pioneers as shade-producers, while their effectiveness in
reducing transpiration has been underestimated.
On the Hawaiian beaches a combination of factors—warmth,
brilliant insolation, and exposure to powerful and rarely inter-
mittent winds—tend to augment transpiration, and make of it an
influence of great potency in retarding certain plants and com-
pletely inhibiting many others.
HAWAIIAN TIDES AS RELATED TO THE LITTORAL
On all oceanic coasts and embayments the tides exert an in-
fluence of greater or less power in determining the seaward limits
of the land vegetation, and the landward extensions of the marine
flora. In regions where the tidal range is great the effect upon
the shore-line vegetation is proportionately augmented; in regions
where the range is slight, its influence is small or negligible. The
Hawaiian Archipelago comes under the latter class.
The greatest tidal contrasts in the Hawaiian Islands are due to
coastal topography, i. e., sea-cliffs contrasted with mud-flats that
lie only a few inches above low tide. At the foot of the sea-cliffs
which rise directly from the water is a tidal (and wave-splash)
zone of two or three feet. This zone is conspicuously marked by
the coralline algae, which form a reddish-purple or lavender band.
If conditions for land-plants are favorable, they may occur only a
few feet above this zone, within reach of the salt spray, and from
a distance appear to rise from. the sea itself. Plant-clad cliffs of
this character also occur in many of the South Pacific islands.
The mud-flats, however, present broad, rocky, mud-covered
platforms, a few rods to a half-mile in width, almost free from sea-
water at low tide, but covered at high tide with six to twenty-four
inches of water. The land vegetation is restricted to the shore-
ward limits of these flats. The absence of the land-building
halophytes—Rhizophora, Brugutera, Ceriops, Kandelia, etc.—
makes invasion very slow.
492 MacCauGHEy: THE STRAND FLORA
The mean range of the Hawaiian tides is very slight, that at
Honolulu being 1.3 feet, and that at Hilo, 1.8 ft, These ranges are
typical for all the islands, and contrast forcibly with the large
ranges of many other littorals. For example at Apia, Samoa,
the average rise is 3 ft. per tide; the tides in Sydney Harbor rise
6-7 ft.; Johnson and York (’15, p. 131) in their comprehensive
ecological study of the tide-levels at Cold Spring Harbor, New
York, found a mean tidal range of 7.63 ft., with a variation of
from 4.2 ft. to 10.8 ft.
The following data from the U. S. Coast and Geodetic Survey
Tide Tables show the tidal range through a single typical month,
January, at Honolulu, in feet:
Date Moon I 2 3 4
3 New; farthest south....... 2.2 0.1 0.6 —0.2
4 POTIRER SS. un eee oo NDS 2:3 0.1 0.6 —-O.1
9 Mavator. We. heen Le hs 0.5 I.4 0.0 1.0
Io Mitet duatter <6 i 0.7 1.2 0.0 2
16 Parthest south. 02.07. 22 Qt 0.2 0.6 0.1
18 Moon; Sporee.. 2... .. re 0.2 0.6 “*O.1
24 Hquatee.: oes ee 0.4 i 0.0 1
26 Abin auarter 0.7 0.9 0.0 5
30 Briheet South ooo Sc: 2: 0.2 0.7 —0.2
Contrasting sharply with the poorly defined tidal zones of the
Hawaiian littoral are those regions with large tidal fluctuations.
For example, Ganong (’06, p. 85) in his studies of the Miscou
Island littoral, in the extreme northeast corner of New Bruns-
wick, on the Gulf of St. Lawrence, found three well defined
beach zones:
First—‘‘a broad sloping inter-tidal beach of pure sand without
vegetation.” This corresponds to the mud-flats along the southern
shores of Oahu.
Second—a “narrow band between ordinary and extreme high
tides.” This zone was practically barren of vegetation. This
zone is negligible in Hawaii.
Third—a ‘‘broad shelf, . . . reached only by the very highest
tides.” This is an “upper beach,” and is characterized by scat-
tered drift-wood and dry, ever-shifting sand. Some of the typical
plants of this zone are Salsola Kali, Cakile edulenta, Atriplex
patula hastata, and Ammophila arenaria. Ganong states ‘that
OF THE HAWAIIAN ARCHIPELAGO 493
this vegetation is distinctly adjusted to the physical conditions,
for it is of great paucity, of small and slow growth, annually re-
newed, closely ground-appressed, and strongly xerophytic.”’
Brownlie (’02) makes much of the irregularity of the tidal
intervals in the Pacific, and states that at Honolulu the intervals
of time from one high water to the next vary from ten and three-
quarters to thirteen and three-quarters hours. A range so wide
apart shows great irregularity compared with the absolute regular-
ity of the movements of the moon. Although this tidal phenome-
non is of great significance from the standpoint of tide studies
themselves, it is not a factor of any special importance in the lit-
toral ecology of land-plants.
Although the Hawaiian marine flora is closely limited by tidal
intervals, these intervals are of little significance in determining
the zonation of land-plants, as compared with the importance of
other ecological factors. This contrasts with the findings of John-
son and York (’15, p. 149), at Cold Spring Harbor, who state:
ft tical distributi f the littoral plants about this harbor
careful study OL he v
shows that this depends primarily and very definitely on the relative time of their
b d ith the ri d fall of the tide. Moreover, the vertical
range of littoral species is strictly, sometimes very narrowly limited. There are no
algae, that are distributed “ between tidemarks”’
: fe rted
species here, except two or three
e nearest approach
(i. e., from low water up to high
this range found for any seed pla
It must be noted that the above statement uses the term littoral
in a relatively narrow sense, as does their further statement that
the vertical range of a littoral plant is exactly proportional to the
range of the tide. This does not apply to the customary usage of
the term littoral, which includes all vegetation along the coasts or
strands, the ranges of which are more or less closely maritime.
ter
nt is that of Spartina glabra.
NATURE OF THE SUBSTRATUM
The character of the littoral substratum obviously plays an
important rdle in determining the nature and distribution of the
littoral flora. Warming (’09, p. 223) groups halophytic plants
into four classes, according to the nature of the substratum upon
which they reach their optimum development: lithophilous,
psammophilous, pelophilous, and helophilous. This will serve asa
convenient basis for classifying the Hawaiian strand, with the
494 MacCauGHEY: THE STRAND FLORA
reservation that not all strand plants are halophytes. The
Hawaiian types to be considered may he referred to the litho-
philous and psammophilous classes, as follows:
A. LITHOPHILOUS
Sheet or flow lava.
Vertical rock shores or sea cliffs.
Littoral creviced rocks.
Lava boulder and pebble beaches.
Tufa beaches.
Coral limestone strands.
2 ee
B. PSAMMOPHILOUS
Coral sand.
Root molds.
~—
1. Sheet or flow lava.—This type of strand is of first impor-
tance in the Hawaiian group, both quantitatively and from the
standpoint of ecologic history. There are more miles of lava
beach than of any other type, or of all the other types combined.
The relatively large areas and recent formation of Maui and
Hawaii have caused this to be the dominant type. Historically
it is first to appear, and it eventually gives way to strands of other
types. The lava flows may be either relatively recent and un-
eroded, like many on the island of Hawaii, or they may be of ex-
treme antiquity and deeply carved, like those of the Na Pali
coast of Kauai, or Kaena and Makapuu, on Oahu. The beach
itself, in either case, is formed of exposed lava beds, very rocky,
with practically no sand or soil, and distinctly uncongenial to
plant life. Lava sheet beaches occur on all the larger islands, but
are best exhibited on the shores of Hawaii and East Maui. Every
gradation may be found from very low, flat lava strands, only a
few feet above sea-level, to bare sea precipices 600-700 feet in
height.
From the historic standpoint the lava beach is of twofold sig-
nificance. If of recent flow material—e. g., the Hawaii flows of
, and the exposure of a new littoral
OF THE HAWAIIAN ARCHIPELAGO : 495
logical history of the islands the Hawaiian strand was wholly of
sheet lava, and today remnants of this primitive beach condition
exist here and there along the coasts. Thus there is great diversity
in the ages of the various shores, and in the amount of plant in-
vasion, both from within and from without, to which they have
been exposed.
2. Vertical rock shores or sea cliffs—These are composed
either of sheet lava or laminated tufa. They are produced by sea
action and fracture. Some of the stupendous sea-clifis of Hawaii, —
Molokai, and Kauai, may have been produced by volcanic or
seismic activity. The sea-cliffs either rise sheer from the water,
or have a narrow strand at the base; this depends upon the depth.
of the off-shore water and both conditions are of frequent occur-
rence. None of the sea-cliffs, no matter how precipitious or ap-
parently uncongenial for plant life, are wholly devoid of vegetation.
Here and there on the surface of the cliff are crevices, ledges, and
little pockets where plants establish themselves.
On the arid precipices—e. g., Koko Head, Makapuu, Mokapu
and Kaena—occur such plants as Euphorbia cordaia, Lepidium
owathense, Schiedea globosa, Kadua littoralis, Tetramolopium spp.,
Lipochaeta integrifolia, Gossypium tomentosum, Sida spp., Jac-
quemontia sandwicensis, Boerhaavia diffusa, Cassia Gaudichaudit
and Capparis sandwichiana.
On the humid sea-cliffs—e. g., Waipio, Nahiku and East Mo-
lokai—are such forms as Campylotheca molokaiensis, Schiedea
Lydgatei, Lysimachia spathulata, Metrosideros polymorpha, Tri-
bulus cistoides and Nama sandwicensis.
3. Littoral creviced rocks.—Along the lava flow and tufa-cliff
coasts, and to a lesser degree along the uplifted coral limestone
shores, the rock crevices are the special habitats of many strand
plants. The crevices are of two kinds: those due to the lamina-
tion of the rock, that is of the sheets or layers of lava or tufa; and
those due to the vertical faulting of these layers. . The former con-
dition produces horizontal crevices, which on the sea-cliffs often
front on ledges of greater or less magnitude. The vertical check-
ing and faulting produces numberless irregular crevices upon the
surface of the exposed strata, and in these crannies soil, seeds, and
spores are lodged. Many of the crevices are less than an inch in
496 MacCauGHEY: THE STRAND FLORA
width, and the plants grow on top of the crevice, merely rooting
in it. Others are several inches wide, and the smaller plants, such
as Lipochaeta succulenta, Lepidium owaihense, and Cressa cretica,
grow down within the crevice, only the upper parts of the branches
showing above the rock. In the very large fissures,—one to three
feet wide, the entire plant body may be concealed within the
fissure.
4. Lava boulder and pebble beaches——Wherever the shore line
lava-sheets are subjected to the action of the sea, they are gradu-
ally broken into massive boulders, which in turn are slowly ground
into pebbles. These metamorphoses are abundantly illustrated
along the windward coasts of Hawaii, Maui, and Molokai, and
in such places as Kaena, Oahu, and Kilauea, Kauai.
The boulders are usually 2-3 ft. in diameter, more or less
oblate, smooth, black, very hard, heavy, and resonant. A beach
composed of these ponderous rocks is very impressive, particularly
during storm time, when the sea mills them with irresistible power.
In various places, especially on the coasts of Kauai and Oahu, the
lava boulders are consolidated in a calcareous matrix, formed of
re-deposited coral lime.
The seaward portion of a boulder beach is barren of terrestrial
vegetation, as is to be expected, but the upper or landward por-
tion, which is not disturbed by ordinary wave action, is the habitat
of such forms as Sesuvium Portulacastrum, Ipomaea glaberrima,
I. insularis, Euphorbia cordata, Tetramolopium sp., Kadua lit-
toralis, and Wikstroemia Uva-ursi.
The pebble beaches are relatively uncommon on Kauai and
Oahu, but are more common on Maui and Hawaii. The pebbles,
at the upper margin of the beach, are intermingled with soil, and
the line of demarcation between beach and lowland is not distinct.
5. Tufa beaches—Tufa craters occur here and there through-
out the islands, from sea-level up to the highest mountain summits.
In a few instances tufa cones stand so close to the shore line that
the sea has cut beach platforms in their slopes. In these cases the
strand is made of the solid wave-cut tufa rock. Leahi, Koko
Head, Koko Crater, Manana, and Mokapu, illustrate this condi-
tion. The tufa is soft and easily sculptured by the waves; it
usually does not form boulders or pebbles, but fractures easily and
OF THE HAWAIIAN ARCHIPELAGO 497
disintegrates into mud. The main part of the tufa platform is
wave-swept and barren of vegetation; the upper portion is the
habitat of such forms as Nama sandwicense, Sicyos hispidus, Jac-
quemontia sandwicensis; Tribulus cistoides, Cenchrus calyculatus,
Waltheria americana. te
Tufaceous sand or mud is brownish or yellowish green in color,
much finer in texture than the coral sand, and much more retentive
of moisture. It often contains large quantities of olivine; this
imparts the greenish hue. Some common plants of the muddy
beaches are: Batis maritima, Polypogon littoralis, Thespesia
populnea, Ruppia maritima, Cyperus laevigatus, Chenopodium
album.
A number of small islets, such as Moku Manu, Manana and
Molokini, are made up largely or wholly of tufaceous deposits, and
represent the eroded remains of former tufa cones.
6. Coral limestone strands.—Within recent geological times
there has been a slight uplift at various points in the Archipelago,
which has resulted in elevating above sea-level broad shelves of
coral limestone. Such areas are especially abundant on the island
of Oahu, and portions of the shore around Pearl Harbor, Kaneohe
Bay, Waianae, Kahuku, and Barber’s Point are formed of the
exposed coral limestone. These shores are often undercut by the
surf, and are sculptured from above by the action of rain-water.
The limestone along the beach may be actually wave-washed, or
may be more or less buried beneath coral sand. Further back
from the shore it is usually covered with a layer of soil.
7. Coral sand.—Beaches of pure coral sand are abundant on
the islands of Oahu and Kauai, and also occur on the islands of
Maui and Molokai. On the coral atolls to the leeward the coral
strand is, of course, dominant. The sand is washed ashore from
the coral reefs, and sometimes accumulates in sufficient quantities
to form dunes. Mana, Kauai, Makaha and Heleloa, Oahu, and
West Molokai, are representative dune regions. Fine stretches of
coral beach may be found at Waikiki, Makapuu, Waimanalo,
Mokapu, Kanuku, Waianae, and Ewa.
The Hawaiian coral strands correspond to the xerophytic
beaches of Cowles (’99, p- 112), who recognizes two classes of
beaches: hydrophytic and xerophytic, defining the latter as essen-
498 MacCauGHEY: THE STRAND FLORA
tially a product of wave action and comprising the zone which is or
has been worked over by the waves. Hence the beach may be
defined as the zone between the water level and the topographic
form produced by other agents. This definition is closely appli-
cable to the Hawaiian coral sand beaches, which are uniformly
xerophytic in their characteristics. Olsson-Seffer ('10) has
stated that the competition for food is more intense, the water
supply less, the light stronger, the temperature higher, the trans-
piration greater, the foothold more uncertain and difficult, the
conditions for plant life generally more adverse, than on any other
soil. |
Shaler (’94) has made some significant generalizations con-
cerning sandy beaches. He points out that dunes and beaches of
coral sand never march far inland, as do quartz sand dunes, for
the reason that the limestone grains speedily become consolidated
into a tolerably firm set rock. It is characteristic of coral beaches
that the materials of which they are composed, unlike those of
ordinary shores, are readily taken into solution, and in that state
may be borne away by the currents to any distance. Notwith-
standing the constant robbery of their materials, which is effected
by the solving process, the coral beaches often widen with great
_Yapidity. Shaler emphasizes that one of the most noticeable
features which is exhibited by beach sands is their extraordinary
endurance of the beating of the waves. He compares the rapid
abrasion of rocks and boulders to the very insignificant abrasion
of sand particles. Though subjected for ages to the beating of
the waves, with perhaps a hundred’ times as much energy applied
to the surface of which it forms a part as would suffice to reduce a
granite boulder containing a cubic foot of material to a granular or
powdery state, the beach sand remains unworn. This endurance
is due to the capillary water. So long as the beach is full of water
the particles do not touch each other. Thus the blow of the waves
is used up in compressing the interstitial water and is converted
into heat without wearing the mineral matter in an appreciable
degree. |
Sandy beaches have a relatively slight water capacity, as: the
percolation is very. rapid. The capillarity is not as pronounced as
in soils of finer texture, and the evaporation from a sandy surface
OF THE HAWAIIAN ARCHIPELAGO 499
is quite rapid. All of these conditions tend to greatly reduce the
available water supply of a beach, even though the latter be ex-
posed to normal precipitation. In other words, the physical char-
acteristics of the sandy beach, as has already been suggested, tend
strongly toward xerophytism. Olsson-Seffer (’10) shows that it
is the volume of water which a soil is capable of placing at the
disposal of the plant, which is the limiting factor in the production
of its vegetative covering and the controlling condition in the
distribution of this vegetation. Percolation in sand is so rapid
that were it not for the counteracting influence of surface tension
very little water would be retained by sand. Permeability in-
creases as the sand particles increase in size. Internal dew for-
mation in the sand is the direct cause of a portion of the permanent
moisture of the strand or dune. It is also to be noted, in this
connection, that extreme changes in the salinity of the soil water,
due to flooding by fresh water, are detrimental to the strand
flora. :
The two important constituents of the soil water of the sandy
beach are lime and salt. On coral beaches the percentage of lime
is very high. It is dissolved out by the rain-water, and ultimately
forms consolidated limestone. The soluble salt content is coastal
and is not of as great ecologic importance as was formerly sup-
posed. An excellent statement is given by Olsson-Seffer (’10).
Sandy soil yields its water to plants more freely than do other
soils, and below the superficial layer of dry sand there is always a
surprising amount of water. Fuller (’12) found this to be more
than double the wilting coefficient of dune soil.
Owing to the frequent inundations by waves and subsequent
rapid changes in evaporation, the soil temperatures of the lower
sand beach are more variable than on any other formations of the
sandy strand. On account of the low specific heat of sand, the
surface layers are rapidly heated in the daytime and quickly
cooled at night. Thus there is considerable fluctuation of diurnal
temperatures.
The list of plants enumerated by Schauinsland (’99) as oc-
curing on the Laysan atoll may be taken as representative of the
coral strand flora of the leeward isles. This list includes:
500 MAcCAUGHEY: THE STRAND FLORA
Lepidium owathense Phyllostegia variabilis
Capparis sandwichiana* Boerhaavia diffusa
Portulaca oleracea Achyranthus splendens
Tribulus cistoides Euxolus viridis
Sicyos hispidus Chenopodium sandwicheum
Sicyos microcarpus Santalum Freycinetianum
Sesuvium Portulacastrum Cyperus laevigatus
Lipochaeta integrifolia Cyperus pennatus
Scaevola Lobelia Cenchrus calyculatus
Solanum laysanense Sporobolus virginicus
Ipomoea Pes-caprae Eragrostis hawaiiensis
Ipomoea insularis Lepturus repens
Nama sandwicensis Pritchardia Gaudichaudii
Heliotropium curassavicum
8. Root-Molds.—An interesting formation along the Hawaiian
littoral, that has also been recorded from other parts of the world
(see Dolley, ’89, pp. 131, 132; and Darwin, ’60), is the root-
mold. This is well developed on sandy shores with persistent
winds, where there has been considerable vegetation. The west
end of Molokai, the Maui isthmus, Makapuu and Kaena on Oahu,
and the Mana region of Kauai all possess notable root-mold for-
mations. The molds are produced by the cementing together of
the sand particles which lie near the ramifying roots of beach
plants; the cementing process is undoubtedly due to specific root
excretions, as well as to the percolating rain-water which follows
the courses of the larger roots. In the course of time the vegeta-
tion dies, the winds Sweep away the loose sand from around the
more solid molds, and the latter are eventually exposed. They
appear either as isolated cylinders, rising here and there above the
sand, or as irregular masses of branching tubes. They rise to a
height of three to twenty inches above the present level of the
sand, and in color are white or yellowish brown.
The lumen varies from a fraction of an inch to nearly a foot
and is rarely open; it is more or less completely filled with lime-
stone. The smaller sizes are the most common, as the majority
* This abundant indigenous pseudolittoral was inadvertently omitted ito the
list on p. 276. It isa straggling shrub, inhabiting arid rocky lowlands and beaches;
a favorite habitat, for example, is a rocky talus slope near the sea.
OF THE HAWAIIAN ARCHIPELAGO 501
of beach plants are slender-rooted. Upon close examination the’
wall is found to be composed of sand, coral particles, and other
minutiae compactly cemented together. In cross section the
wall shows a very much hardened inner layer forming a distinct
zone. The outer layer is relatively soft and easily crumbled.
These molds are the fossils of the root-ramifications of a pre-
vious plant formation. Molds identical in mode of formation
with those of the coral strands are also plentiful in the tufa slopes,
and attain much larger sizes than the sand molds. Punchbowl,
Round Top, Diamond Head, and Koko Head are typical regions
where these molds occur in abundance and in all stages of de-
velopment.
COLLEGE OF Hawau, HONOLULU
LITERATURE CITED
Aston, B. C. The raised beaches of Cape Turakirae. Trans. & Proc.
New Zealand Inst. 44: 208-213. pl. 13, 14. 1911.
Bishop, E. The cold-current system of the Pacific. Science II. 20:
338-340. 1904.
Brownlie, A. The tides in the midst of the Pacific Ocean. Bull.
Geogr. Soc. 34: 17-25, 117-131. f. I-3. 1902.
Bryan, W. A. Deep subsidence of the Waianae Mountains. Thrum’s
Hawaiian Annual, 1916: 95-125. pl. 1, 2. Honolulu. 1916.
Cockayne, L. A report on the sand dunes of New Zealand. Pub-
lished by the Department of Lands, Wellington, New Zealand. 1911.
Cowles, H. C. The ecological relations of the vegetation of the sand
dunes of Lake Michigan. Bot. Gaz. 27: 95-117, 169-202, 281-308,
361-391. f. I-26. 1899
Dall, W. H. North Pacific currents. Science II, 20: 436. 1904.
Darwin, C. Journal of researches into the natural history and geology
of the countries visited during the voyage of H.M.S. Beagle round
the world. London. 1860.
Dolley, S. The botany of the Bahamas. Proc. Nat. Sci. Philadelphia
1889: 131-134. 1889.
Ernst, A. The new flora of the volcanic island of Krakatau. English
translation. Cambridge. 1908.
Fuller, G. D. Evaporation and plant succession. Bot. Gaz. 52: 193-
208. f. 7-0. © 1911.
Ganong, W. F. The nascent forest of the Miscou beach plain. Bot.
Gaz. 42: 81-106. f. I-14. 1
502 MaAcCAuGHEY: THE STRAND FLORA
Guppy, H. B. Observations of a naturalist in the Pacific between 1896
and 1899. 2. London. 1906.
Hedley, C. Ecological sketch of the Sydney beaches. Jour. and
Proc. Roy. Soc.tNew South Wales 49: 15-77. pl. 1-7. 1915.
Hemsley, W. B. Report on the Botany of Juan Fernandez, the south-
eastern Moluccas, and the Admiralty Islands. Challenger Rep.
Botany 1°. 1884.
Report on present state of knowledge of various insular
floras. Challenger Rep. Botany 1. 1885.
Hooker, J. D. On the vegetation of the Galapagos Archipelago, as
compared with that of some other tropical islands and of the con-
tinent of America. Trans. Linn. Soc. 20: 235-262. 1847.
Jost, L. Lectures on plant physiology. English translation. Ox-
ford.
Johnson, D. S., & York, H.H. The relation of plants to tide-levels,
Carnegie Inst. Wash. Publ. 206: 1-162. pl. 1-24 + f. I-§. 1915.
Olsson-Seffer, P. H. Relation of soil and vegetation on sandy sea-
_ _ shores. Bot. Gaz. 47: 85-126. f. I-12. 1909.
——_————._ Genesis and development of sand formations on
marine coasts. Augustana Libr. Publ. 7: 6-183. f. 1-16. 1910.
Pfeffer, W. The physiology of plants. 2d ed. English translation.
2. Oxford. 1903.
Schauinsland, H. an Drei Monate auf einer Koralleninsel. Bremen,
1899.
Schimper, A. F. W. Plant geography upon a physiological basis,
English translation. Oxford. 1903.
Die indo-malayische Strandflora. Bot. Mitt. aus den
Tropen 3. Jena. 1891.
Shull, G. H. The longevity of submerged seeds. Plant World I7:
329-337. J: 1, 2. 1914.
Shaler, N. S. Phenomena of beach and dune sands. Bull. Geogr.
Soc. Amer. 5: 207-13. 18
Tansley, A. G., & Fritsch, F. E. Flora of the Ceylon littoral. New
*Phytol. 4: 1-17. 1905.
Transeau, E. N. The relation of plant societies to evaporation. Bot.
Gaz. 45: 217-231. f. I-19. 1908.
Warming, E. Oecology of plants. English edition. Oxford. 1909.
Wood-Jones, F. Coral and atolls. A history and description of the
Keeling-Cocos Islands, with an account of their fauna and flora.
London. 1912.
INDEX TO AMERICAN BOTANICAL LITERATURE
1913-1918
The aim of this Index ts to include all current botanical literature written by
Americans, published in America, or based upon American material ; the word Amer-
ica be at used in the broadest s
ws, and papers that he exclusively to forestry, egriculture, horticulture,
made in favor of some paper appearing in an American periodical which is devoted
wholly to botany. Reprints are not mentioned unless they differ from the original in
some important particular. If users of the Index will call the attention of the editor
to errors or omissions, their sats will be appreciated.
This Index is reprinted monthly on cards, and furnished in this form to subscribers
at the rate of one cent for each card, Selections of cards are not permitied ; each
subscriber must take all cards published during the term of his subscription, Corre-
spondence relating to the card issue should be addressed to the Treasurer of the Torrey
Botanical Club,
Adams, J. F. Keithia on Chamaecyparis thyoides. Torreya 18: 157-
160, f. I-28. 9S 1618:
Keithia Chamaecy parissi sp. nov. is described.
Anderson, P. J. Rose canker and its control. Massachusetts Agr.
Exp. Sta. Bull. 183: 11-46. pl. 1-3+f. 1-11. My 1918.
Andrews, A. L. A collection of mosses from North Carolina. Bry
ologist 21: 61-67. Jl 1918.
Arny, A. C., & Garber, R. J. Variation and correlation in wheat, with
special reference to weight of seed planted. Jour. Agr. Research
14: 359-392. f. 1-8. 26 Au 1918.
Artschwager, E. F. Anatomy of the potato plant, with special refer-
ence to the ontogeny of the vascular system. Jour. Agr. Research
14: 221-252. pl. 27-47. 5 Au 1918.
Atkinson, G. F. Preliminary notes on some new species of agarics.
Proc. Am. Philos. Soc. 52: 354-356. 8 Au 1918
ew species are described in Amanita (2), Hypholoma (2), Lactarius (2), Lepiota
(1) and Pholiota (1).
Bakke, A. L. Determination of eines Bot. Gaz. 66: 81-116. f. 1-5.
15 Au 1918.
503
504 INDEX TO AMERICAN BOTANICAL LITERATURE
Bicknell, E. P. The ferns and [flowering plants of Nantucket—XVIII.
Bull. Torrey Club 44: 369-387. 10 Au 1917;— XIX. Bull. Torrey
Club 45: 365-383. 20S 1918.
Includes Quercus rufescens sp. nov.
Bisby, G. R., & Tolaas, A. G. Copper sulphate a disinfectant for
potatoes. Phytopathology 8: 240, 241. 11 My 10918.
Blakeslee, A. F. Unlike reaction of different individuals to fragrance
in Verbena flowers. Science II. 48: 298, 299. 20S 1918.
Boas, H. M. The relationship between the number of sporophylls
and the numbers of stamens and pistils—a criticism. Bull. Torrey
Club 45: 343-345. 9S 1918.
Boyce, J. S. Imbedding and staining of diseased wood. Phytopath-
ology 8: 432-436. Au 1918
Brenckle, J. F. North Dakota fungi—II. Mycologia 8: 199-221.
Jl 1918
Hendersonia Crataegi sp. nov. is described and 9 new species in various genera
are listed. :
Britton, E. G. Jagerinopsis squarrosa, n. sp. Bryologist 21: 48-50.
pl. 24. 30 Jl 1918.
Britton, N. L. The botany and plant products of northern South
America. Jour. N. Y. Bot. Gard. Ig: 182-185. Au 1918.
Published also in Science II. 48: 1 56. rg18.
Brown, J. G. Abnormal conjugation in Spirogyra. Bot. Gaz. 66.
269-271. f. I-3. 16S 1918.
Buchanan, R. E. Studies in the classification and nomenclature of the
bacteria—VII. The sub-groups and genera of the Chlamydo-
~ bacteriales. Jour. Bact. 3: 301-306. My 1918.
Buchholz, J. T. Suspensor and early embryo of Pinus. Bot. Gaz.
66: 185-228. pl. 6-10+f. I-3. 165 1918.
Campbell, D. H. Studies on some East Indian Hepaticae. Ann. Bot.
* 319-338. pl. 8, o+f. I-10. Jl 1918.
Dumortiera calcicola sp. nov. is described.
Clute, W. N. The barberry and the wheat rust. Am. Bot. 24: 85-87.
Au 1918.
Clute, W. N. Flowers of varying colors. Am. Bot. 24: 103-105.
Au 1918.
[Clute, W. N.] Noteandcomment. Am. Bot. 24: 106-113. Au1g18.
Includes notes on Sedum ternatum in Illinois, = flowers in autumn, Wilting
of cut flowers, Kochia alata, Odor of Amelanchier, et
[Clute, W. N.] Two forms of leaves on oe same plant. Am. Bot. 24.
IoI, 102. Au 1918. [Illust.]
INDEX TO AMERICAN BOTANICAL LITERATURE 505
Collins, F. S. Notes from the Woods Hole Laboratory—1917._ Rho-
dora 20: 141-145. pl. 124. 1 Au 1918
Includes Microchaete naushonensis, Bulbochaete F urberae, and Erythrotrichia rhi-~
zotdea, spp. nov.
Coutant, M. W. Wound periderm in certain cacti. Bull. Torrey Club
45: 353-364. pl. o+f. 1-3. 20S 1918.
Cunningham, B. Cross-conjugation in Spirogyra Weberi. Bot. Gaz.
OG. 274, 27%. f.-3. 16 8 1918:
Davidson, A. Additions to the local flora. Bull. So. Calif. Acad.
Sci. 17: 60, 61. Jl 1918.
Davidson, A. Lupinus mollisifolius spec. nov. Bull. So. Calif. Acad.
Soi. 29° 57. 4.10138.
Davidson, A. Lupinus Paynei spec. nov. Bull. So. Calif. Acad. Sci.
17: 58,59. Jli1918. [Illust.]
Detwiler, S. B. Battling the pine blister rust. Am. Forestry 24:
451-457. Au s918. _ [Illust.]
Dodge, B. O. Studies in the genus Gymnosporangium—lII.
origin of the teleutospore. Mycologia 8: 182-193. pl. g-iz. Jl
1918.
Drechsler, C. The taxonomic position of the genus SER
Proc. Nat. Acad. Sci. 4: 221-224. Au 1918.
East, E. M. The home of the sovereign weed. Sci. Mo. 7: 170~178.
Au 1918.
Emig, W. H. Octodiceras Julianum Brid. var. ohioense, new variety.
Bryologist 21: 60, 61. pl. 26. Jl 1918.
Farr, C. H. Cell division by furrowing in Magnolia.
5: 379-395- pl. 30-32. 3 Au 1918.
Felt, E. P. Gall insects and their relations to plants.
509-525. Je 1918. [I[llust.]
Foxworthy, F. W. Philippine Dipterocarpaceae, II.
13: (Bot.) 163-199. pl. r, 2. My 1918.
Seventeen new species are described.
Fred, E. B., & Davenport, A. Influence of reaction on nitrogen-
assimilating bacteria. Jour. Agr. Research 14: 317-336. 19 Au
1918.
Frye, T. C. The age of Pterygophora californica.
Biol. Sta. 2: 65-71. pl. 17. 1 Jl 1918.
Frye, T. C. Illustrated key to the western Sphagnaceae. Bryologist
21: 37-48. pl. 17-23. 30 Jl 1918.
Am. Jour. Bot.
Sci. Mo. 6:
Philip. Jour. Sci.
Publ. Puget Sound
506 INDEX TO AMERICAN BOTANICAL LITERATURE
Goodspeed, T. H. Modified safety-razor blade holder for temperature
control. Bot. Gaz. 66: 176,177. f. r. 15 Au 1918.
Graham, M. Centrosomes in fertilization stages of Preissia quadrata
(Scop.) Nees. Ann. Bot. 32: 415-420. pl. ro. Jl 1918.
Harper, R. M. A new seasonal precipitation factor of interest to
geographers and agriculturists. Science II. 48: 208-211. 30 Au
1918. :
Harris, J. A. Secondary parasitism fin Phoradendron. Bot. Gaz. 66:
275,°276. 16S 1918.
Harshberger, J. W. Ecological society of America. The preservation
of our native plants. Torreya 18: 162-165. 7S 1918.
Hasselbring, H. Effect of different oxygen pressures on the carbo-
hydrate metabolism of the sweet potato. Jour. Agr. Research 14%
273-284. 12 Au 1918.
Henderson, M. P. The black-leg disease of cabbage caused by Phoma
lingam (Tode) Desmaz. Phytopathology 8: 379-431. f. 1-10. Au
IQIS,.° ;
Hibbard, R. P. Physiological balance in the soil solution.
_ Agr. Exp. Sta. Tech. Bull. 40: 1-44. f. 1-8. S$ 1917.
Holm, T. Joan Baptista Porta. Am. Nat. 52: 455-461. S 1918.
Michigan
Hotson, J. W. Sphagnum as a surgical dressing. Science II. 48:
203-208. 30 Au 1918,
Johnston, I. M. A few notes on the botany of southern California,
Bull. So. Calif. Acad. Sci. 17: 64-66. Jl 1918.
Johnston, I. M. Some undescribed plants from southern California.
Bull. So. Calif. Acad. Sci. uf 63, 64. Jl 1918
Lupinus elatus sp. nov. is describe
Jones, D.F. The effect of REP and crossbreeding upon develop=-
ment. Proc. Nat. Acad. Sci. 4: 246-250. 7% 2. Au rors.
Keeler, H. L. The wayside flowers of summer. i-xiii+1-—288. New
York. 1917. [Illust.] :
Koessler, J. H. Studies on pollen and pollen disease.—I.
ical composition of ragweed pollen.
S 1918.
Lipman, C. B., & Waynick, D. D. A bacteriological survey of the soil
of Loggerhead Key, Tortugas, Florida. Proc. Nat. Acad, Sc: 4:
232-234. Au 1918.
Livingston, B. E., & Tottingham, W. E. A new three-salt nutrient
solution for plant cultures. Am. Jour. Bot. 5: 337-346. 3 Au1918.
The chem-
Jour. Biol. Chem. 35: 415-424.
INDEX TO AMERICAN BOTANICAL LITERATURE 507
Lorenz, A. Notes on Radula obconica Sull. Bryologist 21: 56-59.
pl. 25. Jl 1918.
Love, H. H., & Craig, W. T. The relation between color and other
characters in certain Avena crosses. Am. Nat. 52: 369-383. S
1918.
MacCaughey, V. An endemic Begonia of Hawaii. Bot. Gaz. 66: 273-
275. 165 1918.
MacCaughey, V. The olona, Hawaii's unexcelled fibre plant. Science
II. 48: 236-238. 6S 1918.
MacCaughey, V. A survey of the Hawaiian coral reefs. Am. Nat. 52:
409-438. f. 1-9. S 1918.
MacDaniels,*L. H. The histology of the phloem in certain woody
angiosperms. Am. Jour. Bot. §: 347-378. pl. 24-29. 3 Au 1918.
Matz, J. Some diseases of the fig. Florida Agr. Exp. Sta. Bull. 149:
1-10. f. 1-5. Au 1918.
McCubbin, W. A. Peach canker. Canada Dept. Agr. Dom. Exp.
Farms Bull. 37: 1-20. pl. 1-O+f. 1, 2. 1918.
McCulloch, L. A morphological and cultural note on the organism
causing Stewart’s disease of sweet corn. Phytopathology 8: 440,
441. pl. 1. Au 1918.
Merrill, E. D. Notes on the flora of Loh Fau Mountain, Kwangtung
Province, China. Philip. Jour. Sci. 13: (Bot.) 123-161. My 1918.
Twenty-three new species are described.
Moxley, G. L. Additions to the lichen flora of southern California.
Bull. So. Calif. Acad. Sci. 17: 61, 62. Jl 1918.
Murrill, W. A. Illustrations of fungi—X XIX. Mycologia 10: 177-
181. pl. 8. Jl 1918.
Prunulus viscidipes sp. NOV., Laccaria amethystea, L. striatula, Leptoniella conica
sp. nov., Prunulus galericulatus, Omphalopsis fibula, Clitocybe farinacea, Marasmius
dichrons and M. institius are illustrated in color. :
Nash, G. V. Hardy woody plants in the New York Botanical Garden.
Jour. N. Y. Bot. Gard. 19: 58-62. Mr 1918; 86-91. Ap 1918;
108-112. My 1918; 139-142. Je 1918; 167-171. Jl 1918; 192-196.
Au 1918.
Nelson, J. C. An addition to our food plants. Am. Bot. 24: 97-99-
Au 1918
Crambe maritima. .
Nelson, J.C. A new form of Prunella vulgaris. Am. Bot. 24: 82-85.
Ison, J
Au 1918. [Illust.]
/
508 INDEX TO AMERICAN BOTANICAL LITERATURE
Nichols, G. E. War work for bryologists. Bryologist 21: 53-56.
Jl 1918.
Nothnagel, M. Fecundation and formation of the primary endosperm
nucleus in certain Liliaceae. Bot. Gaz. 66: 143-161. pl. 3-5. 15
Au 1918.
O’Gara, P. J. The white-spot disease of alfalfa. Science II. 48: 299-
301. 20S 1918.
Paulsen, O. A new Cereus from the West Indies. Jour. Bot. 56: 235.
Au 1918.
Cereus venditus.
Peltier, G. L. Susceptibility and resistance to citrus-canker of the
wild relatives, citrus fruits, and hybrids of the genus Citrus. Jour.
Agr. Research 14: 337-358. pl. 50-53. 26 Au 1918.
Peltier, G. L., & Neal, D. C. A convenient heating and sterilizing
outfit for a field laboratory. Phytopathology 8: 436-438. f. 1, 2.
Au 1918.
Pennell, F. W. A plea for Aureolaria. Rhodora 20: 133-137. 1 Au
1918.
Potter, R. S., & Snyder, R. S. The production of carbon dioxide by
molds inoculated into sterile soil. Soil Sci. 5: 359-375. pl. r+f. 1-4.
My 10918.
Reichert, E.T. The differentiation and specificity of starches in rela-
tion to genera, species, etc. Part I. i-xvii+1-342+ [1-21]. pl. 1-
roz. Washington. 1913; Part II. i-xvii+343-900+[1-18]. Wash-
ington. 1913.
Carnegie Publ. No. 173.
Riddle, L. W. Some extensions of ranges. Bryologist 21: 50. 30 Jl
1918.
Roberts, J. W. The source of apple bitter-rot infections. U.S. Dept.
Agr. Bull. 684: 1-25. pl. 1-5. 19 Au 1918.
Robertson, C. Pollination of Asclepias cryptoceras. Bot. Gaz. 66:
177. 15 Au 1918.
Sargent, C. S. Notes on North American trees.—II. Carya. Bot.
Gaz. 66: 229-258. 16S 1918.
Includes Carya leiodermis sp. nov. and Many new varieties and combinations.
Schneider, C. Notes on American willows—I. The species related to
Salix arctica Pall. Bot. Gaz. 66: 117-142. 15 Au 1918.
Schwarze, C. A. The parasitic fungi of New Jersey. New Jersey
Agr. Exp. Sta. Bull. 313: 1-226. f. 1-1056. S$ 1917.
INDEX TO AMERICAN BOTANICAL LITERATURE 509
Shear, C. L., & Stevens, N. E. Plant pathology today. Sci. Mo. 7:
235-243. $1918.
Shear, C. L., Stevens, N. E., Wilcox, R. B., & Rudolph, B. A. Spoilage
of cranberries after harvest. U. S. Dept. Agr. Bull. 714: 1-20.
9g Au 1918.
Shufeldt, R. W. Midsummer flower-hunts. Am. Forestry 24: 489-
496. f. 1-10. Au 1918.
Sinnott, E. W. Factors determining character and distribution of food
reserve in woody plants. Bot. Gaz. 66: 162-175. Jody 22015 AU
1918.
Skottsberg, C. The islands of Juan Fernandez. Geog. Rev. 5: 362-
383. f. 1-20. My 1918.
Soth, B. H. Blossoms by the wayside in the Rockies. Am. Bot. 24:
88-91. Au 1918. [Illust.]
Spegazzini, A. Observaciones microbiolégicas. An. Soc. Cien. Argen-
tina 85: 311-323. Je 1918.
Includes the new genera Amphoropsis, Myriapodophila, Thaxteriola, Entomo-
cosma, and several new species in the Laboulbiniales, and two new species in Chan-
transiopsis.
Stevens, N. E. American botany and the great war. Science II. 48:
177-179. 23 Au 1918.
Stevenson, J. A. Citrus diseases of Porto Rico. Jour. Dept. Agr.
Porto Rico 2: 43-123. Ap 1918.
Stone, R. E. Incubation period of Cronartium ribicola on the white
pine. Phytopathology 8: 438-440. Au 1918.
Sudworth, G.B. Miscellaneous conifers of the Rocky Mountain region.
U.S. Dept. Agr. Bull. 680: I-45. pl. 1-13 +maps 1-9. 14 Au 1918.
Thériot, I. Note sur une mousse du Chili, Barbula flagellaris Schp.
Rec. Publ. Soc. Havraise d’Etudes Div. (1er Trimestre 1917), pp.
I-7. 1917.
Turner, C. E. Plant and animal life in the purification of a polluted
stream. Sci. Mo. 7: 34-45- Jl 1918. [Illust.]
Veitch, F. P., & Rogers, J. S. American sumac: a valuable tanning
material and dyestuff. U.S. Dept. Agr. Bull. 706: 1-12. pl. 1-5.
26 Jl 1918.
Weatherwax, P. The evolution of maize.
309-342. f. I-36. 9 S 1918. .
Weir, J. R., & Hubert, E.E. Cultures with Melampsorae on Populus.
Mycologia 8: 194-198. Jl 1918.
Bull. Torrey Club 45:
510 INDEX TO AMERICAN BOTANICAL LITERATURE
Weniger, W. Fertilization in Lilium. Bot. Gaz. 66: 259-268. pl. r1-
Fz. 16:5 1938,
Wieland, G. R. The origin of dicotyls. Science II. 48: 18-21. 5¥J.
1918.
Williamson, E. B. A collecting trip to Colombia, South America.
Univ. Michigan Mus. Zoo. Misc. Publ. 3: 1-24. 22 F 1918.
Wodehouse, R. P. Direct determination of permeability. Jour.
Biol. Chem. 29: 435-458. 1917.
Wolf, W. Quercus bernardiensis sp. nov. Torreya 18: 161, 162. 7S
1918.
Woodward, R. W. Some Connecticut plants. Rhodora 20: 97, 98.
1 My 1918.
Woodward, R. W. Some Rhode Island grasses. Rhodora 20: 116.
11 Je 1918.
Woodward, R. W. A Salix rostrata hybrid? Rhodora 20: 132. ° 39
1 1918.
Young, H. C., & Cooper, E. H. A method for determining the fungi-
cidal coefficient of lime sulphur and other common fungicides.
Ann. Rep. Michigan Acad. Sci. 19: 241-236. .10Ty.
Young, V. H. Some factors affecting inulase formation in Aspergillus
miger. Plant World 21: 75-87. Ap 1918.
Zeller,S.M. An interesting fungus from F riday Harbor, Washington.
Publ. Puget Sound Biol. Sta. 2: 95,96. 1 Jl 1918.
Rhizopogon diplophloeus.
Zimmerman, H. E. A tree that owns itself. Am. Bot. 24: 81, 82.
Au 1918. [IIlust.]
INDEX TO VOLUME 45
New names and the final members of new combinations are in bold face type.
Abies balsamea, 37
Abrus precatorius, 277
cia “caven nia, 270; farnesiana, 276,
489; 275
Acer pennsylvanicum, 37, rubru
Aa aon charum, 31, 37, 42, eer wane
Aca
Achillea Millefotium, 39
380; oblongifolia, 381; sp., 33, 38;
stolonifera,
Ammophila arenaria, 492
msinckia arenaria, 382; intermedia,
2
Anacardium occidentale, 271
pores pines cate 38, 42
Antheri
8; Paterna alatephetton. ¢
Achryanthus aspera, 277; 0 obovata, 14
poe ay ik 274, 500; splendens rotun- <, y andros seinttatuih: 40; canna-
ata,.274 binu
Actaea alba, 31, 39; rubra, stig Aralia ‘nudicaulis, 38, 207; racemosa, 40
Actinostroma pea orm ae Arctium minus,
Adiantum pedatum, Arioieashvies Ah Pia 33, 38, 382
Bolandert 406; Bolan- | Argemone intermedia, 150; mexicana,
Adenostegia, 400;
deri capitata, 403, 404 iosa, 404;
27
canescens, vag A421; fiitolia, 403, 410; | Argyreia
0, 403, 408; Helleri, 4173
ni,
laxiflorus, pet littoralis, Woecal 4133
Kingii, 416; mollis, 419 maritima,
420; Nevinii, 403, 409; " Occuttiann,
418; , 419; Parryi, 421; par-
viflora, 403, 409; pilosa, 403, 405;
Pringlei, 402; ramosa, 403, ida,
403, 411; rigida brevibracteata, 403,
412; aig 403, 400; ‘
3, 407; Wrightii, 403
40
ose et gi, Taxonomy and distribution
399
FE anthericicola, 149; Atriplicis,
ae onspicuum, I5
; 150; ¢ 33
Hyptidis, 454 Keerliae, 154; Mozinnae
152; myricatum, 291; Pereziae 153;
plenum, 149; ranu culacearum, 147;
Rosae, nee roseum, 154; steviicola,
1543 nardiae, 150; Tithymali, 151;
ee 4
ei 154
ethusa Cynapium, 56
joa stemma Githago, 379
Aira psi es Orbea? 369
cham of Grimaldia fragrans,
The, ont
eurites m —— 271, 276
\lisma rcs 368
\Inus incana, 3
\lopecurus ge eniclatus 369
\lsine graminea,
.mbrosia eh ondaita lia, 40 ‘
lanchier canadensis, 295; i
os. 2943 Trace
be i Be th ts es Se
«oe
292,
termedia, nantu
tiliaefolia, 275, 487, 489
Arisaema pusillum, 373
;| Aronia nigra, 3
ARTHUR, J. é New species of Uredi-
neae— X, I4I
Asclepias incarnata, Fol syriaca, 39
Asarum canaden
cppcreere eet; 98; angulare, 98;
uriculatum , 100; caryotideum,
obum, 98-100; falcatum
; Filix-mas cristatum,
inum, 98; hirtipes,
100-102; marginale, 207; spinulosum
intermedium, 207; Tsus-Simense, 98,
ium
Filix-foemina, 365; nidus, 94
te)
m, 366; wrcorning
atum rubellum
seat no
fe) fo) 457
sima, 456; sambucina, ee Se cactichden
455; vi — 455
Auriculareae, 455
Avena wat 55
cenn
Avi ia a paren 270, 2
Avery, B. T., HARR oe A. i. “sei
ng of morpho variations
he seedling of ka vulgaris, on
Barringtonia excelsa, 270; speciosa, 270
512
Batis maritima, 275, 487, 407
Berry, E. W. Notes on acid fern genus
4, 207; papyrifera,
37; populifolia, 375; pumila, 34, 38
BICKNELL, -
i » 365
so na, = ‘sag ona 145
erkandera adusta,
Blephariglotts wana 374
M..-T elationship between
sre number of 9 a Ear des d the num-
ber of stamens and pistils—a criticism,
3
Boerhaavia diffusa, 275, 489 495, 500
B preien B abietinus, 466; isabellinus, 463;
465
Borrichia a, 271
Botrychium Lunaria, 203; onondagense,
203; virginianum, 40
Bromus hordeaceus, 371
ne m,: 37%
E.
secalinus, 371;
corsage in Phe-
wens polypodioldes, 3
ryum conto on nctorium,
433, 437. 438, yn parieecu, 433
aeoma destruens, Sybat exitiosum, 455;
' miniatum, 455; Rosae miniatu um, 455
nero Bonducella, 270, 271,
275;
Cail, id i nee
canadensis
Calophyitum. inophy ae ch eos 489
Caltha palustris,
Calystegia Soldanella,
Campanula ‘essere 58, 59, 62, 66;
iveings. 40
Cs ana avalia, 270, 271
¢ +378
Ca moluccensis, 271; obovata, 271
€ aareyistane molokaiensis, ne 495
Capparis sandwichiana, 495,
Cardiospermum, 271; hadi acabael 277
Carex arctata, 31, 39; filiformis, 05.30}
hirta, 372, ei, 372; imcomperta,
372; laxiflora, 31; leptalea, 372; muri-
Carrier of the mosaic disease, A, 219
Cassia Gaudichaudii, 275, 4
Cassyt iformi 2
Castalia odorat
=
Cen ort stn acta 276, 497, 500
Centrose: 271
Corte | GasBat, 270
Chaetochl imberbis, 369; imberbis
perennis, 369; occidentalis, 369; ver-
sicolor, 3!
INDEX
Chamaecyparis thyoides, 291
7 38
497; sand-
Cc
Chiogenes, be hispidula, 39
Chloris,
Ch loropyron canescens, a maritima,
ustre, 420; Parryi, 421
1 vacations 39
s, 70
a, 382
ibotium Chamissoi, 134; Menziesii, 134
pp aeons affinis, 453; 4533
1 a
Krugiana,
eucode + 45
ircaea alpina, 4
Cissampelos Pariera,
aie oo 276; maris-
oides
Fe hiprvie nites Blumeri, 461; elegans, 461;
Decandolleana, 461; ntcadibatttocnis
4
Clarkia pulchella,
56
Clathropteris meniscioides, 280; platy-
phylla, 279, 280-28
Clavaria — 462
Clavarieae, 462
Clem cane Drummondii, a
Clermontia parviflora
SP eg ego oo 271
Clintonia —
Coccoloba
Cocos sad hess hs ‘ ie
Coker, D. Revi the North
American species veges tacaicnen, 433
Col i i 55
Colubrina asiatica, 270, 272, 274
palustre,
mptonia aiphitinis, 291; peregrina,
wer
Conocarpus
Conringia orientalis, 380
Convolvulus epithaimeeus, 40, 271
’
Coptis trifolia, 39, 207, 379
Cora gyrolophia, 461; pavonia, 462
Cordea subcordata, 270, 271, 275
Cordylanthus bicolor, 404; c us,
404; c ; ; filifolius, 410,
411; filifolius brevibracteatus, 412;
Kingii, 416; laxiflorus, 401; mollis,
41 maritimus, 0; ma 409;
Orcuttianus, 418; yl, 421; pilosus,
5, 406; Pringlei, 4o2; rathouie: 414;
rigidus, 411; tenuis, 406
— atypus, 464; melleofiavus,
ig abietinus, 466; perpusillus, “
rubritinctus, 466
INDEX
Cornus canadensis, 39, 207; stolonifera,
3
Correlation of morphological variations
in ag seedling of Phaseolus vulgaris,
Contkean caeruleum, 451, 457
COUTANT ot pis Wound periderm in
certain. cac
Cratacg us prin at 381; oxyacantha,
Crepidula plana, 53
ressa cretica, ag 487, 496
uae sp.,
Crotonopsis argentea, 478, 4793 pectin
478; linearis, 477, 478, 479; spinosa
478,
Cudonia
Cyanea pects 135; Giffardii, 133, 134;
age BN ngere, 136; 2
135; superba, 134; t ne sca ar
Cycas circinalis, 271
Cyclomyces spadiceus, 467; tabacinus,
pe die Sores omens 44
elegan
na, 5
ra. filiformis, 380; intermedia, 380;
Drose
ino ifolia, 3
Drynaria a composita, 285; quercifolia, =
— spinulosa, 49; Thelypteris,
Dryostachyum, 285
Dubautia laxa,
Dulichium nae 37 39
Elaeodendron, 271
3 haris “arog 40; tricostata, 372
Elymus sp., 33,
Encalypta tadle. 440; alaskana, 433,
435, 441; eee 435, 437, 440; apophy-
443; brevicolla, 437,
;
a 437s 8; extinctoria apiculata,
439; extinctoria mutica, cera
lac 435; laciniata, 434, » 441;
me : rica, 434, 435, 436,
444; ie poet 435, 436, 444;
Macounii, 433, 435, 443; mexicana,
435, 441; pilifera, 440; procera, 435,
438, 444; rhabdocarpa, 435, 437, 439,
440; rhabdocarpa micr Sie :
Cymatoder
* Cynodon oR neva oe 435, 436, 444; a arg SUE ie 4453
Cynoglossum officinale, subbrevicolla, 434, 436, 444; subspa-
Cynometra cauliflora, 271 Sager 434; 435, 439, 440; Sola
iainan bs laevigatus 275, 497; SF meck
natus, 276, 500 pheloides, ee Sacalyoea, cog ape of the North Ameri-
bellatu a can species of, 433
Covet hirsutum, 40 ea scandens, 271
Cyrilla racemiflora, 150 banc — 40
pndona tr Fortunei, 98; Rochfordia- Equise' arvense, 39, 366; fluvi atile,
40; cone 33, 393 laevigatum,
Cyaan bulbifera, 207 str ; ides, 40
Cyrtandra, 134 Eragrostis hawaiiensis, 276, 500
Eriophorum viridicarinatum, 40
Dalibar deaativntooe ts 07 Erodium cicutarium,
Dammara australis, 272 Erysibe a Rosarum, 455; pani-
ia spicata, cor
Danthonia <p - oni ae 219, 277; Tatula, Erysimum chelranthoides, 380
9 Erythraea sabaeoides,
on verticillatus, 3 Erythrina re aroanrecelg :
Dennstaedtia pu acne 306 Euchlaena luxurians, 318; mexicana,
Derris uliginosa, 2 318, 332 é
ctyocalyx Miersil, ager Eupatorium, 154; perfoliatum, 39; pur-
Diervilla Lonicera, 38, 2 pureum, 39, 154
Digitalis grandiflora ee ails PNK 56 | Euphorbia, 271; Atoto, 270; —
ioclea violacea, 272 275, 487, 495, 496; pilulifera, 277
DGE, . Studies in the gen = aren ~ 4
um—II. Report on cul- | Euxolus viridis, 277, 500
iy i . Evans, A. W. The air chambers of
ha, 271
Exidia ampla, 456; auricula-jude, AST:
nobilis, 456; tenuis, 457
ii a. 37-3
‘erns and flow: as late of Nantucket
Fe XIX, The, 3
514
FERR R. S. Taxonomy and distri-
Sain of ys Nia 399
Festuca capillata, 371; myuros, 371;
toflora, 371; ovina, 371
Ficus macrocarpus,
Filicites crags
Fimbristylis Potties. 274, 489
Fleurya rcracas: 276
Fomes albomarginatus, 4; carneus,
466; cornu-bovi 5; Fullageri, 465;
oe aps 3; Merrillii, 465; pyr-
oseus, 465; spadiceus,
4 . 468 cbr, 407
raxinus ameri a, 37; nigra, 37
Friesia, in Scones 479
Galium a.
Gaultheri um mbens, 39
pcoanaibag 79; glabrum, 80; glutinosum,
oye mexicanum, 156; Robertianum,
Sollee Fernaldii, 370
Gossypium to omientosum, 274, 495
RAFF, P. W. ar, ippine Basidiomy-
cetes—III, 4
Grimmia,
Grimaldia —— 236
Grimaldia grans, The air |
Seep 2963
2 ee Pity enescens, 94;
macr cropus, 287, 288; ‘iitue avis 287,
293, 204; transformans.
Gyromitra esculenta,
infula, 79
79; gigas, 80,
ce ae eg eA 454
HARPER T. Two remarkable Dis-
The plant eg npc
rthern lower Michigan a its
ERY, 6, Eo. Co
ological variations in
of Phaseolus vulgaris, 109
» 383
7
375; glabra, 375; micro-
Hi imeola mule, 456; ampla, 456; auri- |
INDEX
cula-judae, 456; nobilis, 456; tenuis,
5
Hordeum jubatum, get vulgare, 372
Houstonia caerulea,
sop - tie variation
€ noctiflora, 157
Hume ne upulus, 378
Hydneae, 463
evasive ei useage 257
Hymenochaete adusta, 457; cacao, 458;
nigricans, puis rheicolor, Bie tenuis-
sima, 45
oe 461
yoscyamus ni
pacum
Hypoxis hirsuta, 374
Ilex verticillata, 38
ag — pte pa
o Am
Index rican “botanical literature,
43, pod i 159, 211, 253, 301, 347,
385, 42 Pit 503
74
oea, 271; acetosaetoia, 275; glaber-
rim a, 272, 275, 4 randiflora, 271;
ari gin martina, 270;
aprae, 270, 272, 27
Iris aon 29, 39
Irpex flavus, 463
a a aca 276, 495, 497
Juncus balticu
Juniperus roel
288,
292; virginiana,
Kadua littoralis, hg 489, 495, 496
Kalmia poli a
Keerlia mexicana, -
Koellia i incana, pet
Krameria glandulosa
Kyllingia anneal pe
Labordia, 134
aburnum oe oa
Lachnea semitos
Li dehaadiees, becsitiierine. 462; usam-
rense, 462
Lactuca
vaguncularia siti
to)
rix decidua, 366; pasa 37,42
Laschia Dgariays 6
Lathyrus odoratus, "ig palustris, 39
ection) Leggettii, 4
Ledum groen ai 38
Leotia, 79
Leersia alpina, 441; ciliata, 441;
torta, 445; extinctoria
ell eal eal wall wall eal en!
; vulgaris, 43
Lemna minor, 373; trisulca, 373 .
Lens esculenta, 56
INDEX 515
Lepidium owaihense, 274, 487, 495, 496,
00
Leptemon lineare, 479
Leptilon canadense, 39
teal Se 275, 500
aena g
rie: piledelpbienl 39, 373
Linnaea americana
Linum angustifolium, 59, 70; striatum,
382; usitatissimum, 70
Liparis Loeselii, 374
Lipochaeta commana littoralis, 274; in
tegrifolia, 274, 495, 500; ciiinlenihy
Lycopodium annotinum, 39, 207; com-
partie um, 366; lucidulum, 207; 0
scu Pa se Eager dendroideum,
mindy ;
Ly sachin pss ame 275, 495; ter-
restris, 39
ythrum maritimum, 276
pgp oe tine V. The strand flora of
he Hawaiian Archi pela go—I. a
hical relations, origin, and
Sse 259;—II. Ecological a
t10:
, 483
274, 496 Varonat ld naira ge ai 276
Liviodendron Tulipifera, 379 Matonia pectin:
ardinalis, 40; ia me 59, 66;| Medeola virgini ee
bride 138; oahuensis, enyanthes trifoliata, 40
Lonicera canadensis, 34, 38 Merulius s, 456; auricula, 456
Lumnitzera c 271; ra a, 271| Mesembryanthemum spp., 2
Lupi a 192; arenicola, I, 2; Metrosideros polymorpha, 276, 495;
brevicaulis, 11; Bridgesii, 1 20;| rugosa, I
iflo: 1-7, 167, 172, 174; den- Michigan, The plant population of
siflorus altus, 172, 185; densiflor northern low 3
ustrocollium, 173, 200; de siflorus| Mirabilis Jalapa, 56, 65
imus, 173, 199; densiflorus itchella ot
crinitus, 173, 195; densiflorus cur- Mitella n
i 96; di orus Dud-) Mor ithe: as. 79, 80; giao 79
leyi, 173, 188, 189; densiflorus glareosus Morinda, 271; citrifolia, 270, 271,
; densiflorus la , 172, 181;| Mozinna spathulata, 152
densiflorus latilabrus, 172, 178; den- Mucu na gigantea, 271, 272, 276; urens, 272
ifio idens, 173, 188; densiflorus Myoporum sandwicense, em
enziesii, 172, 176; densi Mc- | Myriadoporus adustus, 4
Gregori, 172, 184; densiflorus palustris, | Myrica carolinensis, ase “Gate, 38, 374
173, 176, 101; pai ig perfistulosus, Myriophyilu um prosperinacoides, 482
172, 175; densiflor persecundus
173, 190; densiflorus Deed, £73) 1973
densiflorus scopulorum, 173, 201; den-
orus si
trichocalyx, 173; 198; densiflorus vas-
titicola, 172, 186; ravine sap sierra
Menziesii, I, 3, 170; Menzi re
I; microc s, I-9, 200; micr us
ruber, 10; palustris, 1; ruber, I, 10
subvexus, I, 5, 7, 12, 13, 1933 eubvexus
ee 13, 19; subvexu us fi uviatilis,
a or Leiber; 13, 16; subvexus
rgii,
nigrescens, 13; subvexus phoeniceus, sinu
sag 8, | Nol
13,235 hate xUus so eee Sa
13, 15:8 Wilke
vexus esii, I
Lupinus, ie in the pate) a; —
—IIlI, 16 Nympha'
Lyciu m sandwi 4
Lycoperdon subcordat 454
Nama sandwicensis, 276, 495, 496, 500
Naias s flexilis, 368; guadalupensis, 368
ew species of Uredineae— 41
Nicotiana _— 220; — 270;
Tabacum, 219, 2
Nigella da saan ai ee 65
oie fruticans, 27
SHIMURA, = A carrier of the mosaic
oaas
;| Notes on plants of the southern United
States—IV, 4
ayers on the a genus Clathropteris,
scion distans, 96; Eckloniana, 96;
74
ea alba, 52; ’ variegata, 28, 39
pene bee 53, 54, 58; Lamarckiana,
Lycopersicum peruvianum, 279
Sar 5
516
On the Scena of cell shape in leaves
eo ng shape, 51
on mesentericus, 455
Onloclea sensibilis, 39
Ophioglossum vulgatum, 276, 365
ie a 271° discata, 353; versicolor,
Ceyeoneie asperifolia, 40
Osmunda cinnamomea, 94; Claytoniana,
94, 3653 “i 94, 39, 103, 104; spec-
tabilis, 3
Ostrya stare Baas te 3
coccus eaicoen teh 40
hi tela nea fo rae rommgaaty 276
angium €, 270
Paideulacta: acutiflora, 370; american
370; arta 370; pallida, 370; saltidd
Fernal
Panicum Addieonit. 368; Bicknellii, dt
69; dic
depauperatum, 40, 368,
mum, 369; linearifolium, 368; ae
ionale, 369; Owenae, ro 369; san-
guineum, 56; tennesseense, 369
nel wie
Past tiv.
rae seein atanesae Tage 99; atropurpurea,
ae 102; atropu rpurea cristata, 96;
cre’ 99; flavens, 96; nivea, 96;
te sha "06; idis
Pemphis acidula, 270, 27t
PENNELL, F. W. Notes on plants of the
southern United States—IV, 477
378
Studies oh the vegetation
of New York State—II The ides
ion of a cial plun,
anum sandwicense, 276
Peziza auricula, 456; coronaria, | ~ cor-
onaria macrocalyx, 85; Stev niana,
phological variations in
ext I
Phegopteris Dryopteris, 207
Phesopteris polypodioides,, Regeneration
5!
vulgaris, Correlation of mor-
the seedling
Philippine ae 451
hlebia 455; reflexa, 456;
' Physalis Alkekendi, 2 220, pee!
Phytolacca decandra 155
INDEX
Picea ipa rs 37, 42;- mariana, 28,
Pinus antec. a2. 37, 43} insularis,
igida, 366;
464; resinosa, 37, 42
bus
Strobus, 37, 42; ee ae 51
Pisum sativum, 56, 65
Plant population of rthern lower
ichigan and its environment, The,
Plantago sors 59, 68; major, 59, 68;
tom 270
Plat
elite ily Oa A5I
a tensi
Polygonatum pe 0; 373
ie m a 142; barbatum,
452; chine ar ; maritimum, 378;
panuiealdsgns laevigatum, 378; pen-
sylvanicum n hilum, joe
Polypogon littoralis, 275,
Pol rus a us, 463; Bis omarginatus,
404; atypus, 463; benguetensis; 464;
Burtii, 463; carneus, . rpinus,
463; cinereo-fuscus, 465; crispus, 4633
dol 466; Elmeri, 468; fl , 463;
Halesiae, 463; incarnatus, 46 es,
464; laeticolor, 464; Lindheimeri, 463;
luzonensis, 464; melano , H
hrocroceus, hd pallescens, 463;
roseus, 466; rufo-pallidus, 466; spadi-
ceus, 467; ib cercas, 463; tabacinus,
Polyporea
Pelystiotne ee 466; atypus, 464;
barbatus, 467; be bactbnin 464;
cervino-gilvus, 463; microcyclus, 467;
perpusillus, 466; rubritinctus, 466;
ite Rt 467; Hoe tidy 467; tabaci-
nus, 467; tabacin barbatus, 467; .
tabacinus Sekutyssaa: 467; xeram-
pelinus, 46
Pongamia glabra, 270
opulus balsamifera, — grandidentata,
33, 37; tremuloides, 33, 37
Portulaca oleracea, rigs 5
otamoge ianus, 367; pecti-
natus, 367; pulcher, 36
as
Paar Gaudichaudi, 500; spp., >is
Proso os juliflora, 276, :
maritim: 381; pennsylvanica,
33, 395 “pumila, 33
Pteris aquilina, 38, 94; argyraea, 97, 102;
cretica se eata, 96, 97, 101, 102;
gmantiana, 96; Parkeri, 97; quad-
riaurita gbreed, 96; serrulata, 94, 97;
sulcata, 96, 97, 102 :
Puccineae,
adducta, 148; a 146;
Ciberti, 454; aie 146; Eriophori,
144; Heliconiae,
eon in 4545
INDEX
incondita, 148; missouriensis, 146;
nike iP 149; Rubles 147; pat-
pai set ses 144; as rae,
145; ap Rivinae, 15
Bacciaain: "aatine 391; ileus tenuis,
I
37
Pustularia gigantea, 82, 84; vesiculosa,
Pyrola cn 207
Pyropolyporus albomarginatus, 465; Mer-
rillii, 46 i
; alba x rufes-
Quercus alba, 31, 37,
cen
42;
bicolor, 378; borealis maxi- Sam:
517
Ruppia maritima, 271, 275, 367, 497
Rynchospora aurea, 453; glauca, 453
Sabbatia campanulata, 383; stellaris, 383
Sagus, 271
alix Bebbiana x cinerea, 374; discolor,
38; Bib ru lucida, 37; pedicel-
laris, 34; iolari id yo rostrata,
38; Smithi ane wae 37
Sagittaria Engelmanniana, 368; latifolia,
6
Salsola Ka li, 492
nap ath PIA 277
aman ,
8, 377;
’ ea : bacttnocs srr 4, 38
velutina, 378; ilicifolia, 376; ilicifolia Santalum Fre fea tlaoaee littorale, 274,
velutina, 377; pagodaefolia, 370; 487, 489, 500 ;
palustris, 378; prinoides rufescens, Sapindus Saponaria, 271
376; escens, ure stellata, 376;| Sassafras Sassafras, 380
velutina, 40, 375 Scaevola, 271; coriacea, 274; Koenigii,
271, 272; Lobelia, 275, 500; Plumieri,
oe pene 380
Raim ummondii
Randers ecinenten repens, 379
p oe 28
eana luxurians,
Beveti eration in peli polypodi-
oides
Relationship between the number
ea we lls and ake number of stamens
nai edo criticism, The, 343
Reseda core iw: 4
Revision of the North American species
270
sresph globosa, 274, 495; Lydgatei,
a nus ni s, T44
Scirpus americanus, 39; 144, atrovirens,
4 rinu:
0; cype pelius, 39; maritimus,
275; validus anh sina
ae Hae rium
Sesuvium, Der olecwatstai, 275,
496, 489, 500
Setcpeelie subulata, 383
S, Paes 495, 500; micro-
Rh s alnifolia, 34, 38 Sicyos ag ec oa
Rhe Alifanus, 482; ; carpus
cubensis, 481; delicatula, 481; interior, Sida eb. = pegs
480, 482; la ; latifolia, | Silene ie aa A striking variation in,
480; lutea, 480, 481; oa, A8I;| 157
Nashii, 480, 482; parviflo 481; | Sistotrema violaceum, 466
petiolata, 481; serrulata, pov acts. Smilax herbacea, 373
482; virginica, 482 SMITH be P. Studies in the —_
iata, 281 Lu upin s—lIl.
Rhlzomopters ¢ ag nr of Lupinus cauitiowos: I ear’
Rhus, 271; glabra, ee oh sp., 38: Toxi- Lupinus densiflorus, 16
codendron, 33, Solanum actleatisimum, 219, 221-231;
Ribes lacustre, 20 olinense ‘ ; laysanense, 274, 500;
ia fluit ae OFrostii, 243, 244;| Nelsoni, ot Sa 219; racemosum,
glauca, 245; natans, 243 148; triquetrum, 148; tuberosum, 56
Ricciocarpus natans, — Solidago, 271; sis
mneratia sp.,
Ricinus BE
om eget of Hawaiian
71
Psa Rigen 271
ricanum, 367; eurycar-
Rock,
sso ial 8 >
Roestelia : ormans, 287
Rollandia an, , 136; longiflora, 137 : “a Michausiana, 30, 271
Rosa cinnamomea, 380; sp-» 33+ 38 pathu
Rubus alleghenie 3 , 42; odoratus, | Spiraea geek a
207; strigosus, 38, 42; triflorus, 39 Spongipellis semen on
Rumex crispus, ; elongatus,. 378; pon oy 271; argutus, 155; Vir:
3783 persicarioides, tae 9, 500; virginicus oak
37
maritimus fueginus,
78
INDEX
518
Stem, W. N._ Studies of some new cases | Thymus serpyllum, 383
of apo y in ferns, 9. Tiarella satin ng “tes
Stereum adustu 457; bellu 458; | Tilia amer .
cacao, int concolor, 4593 contrarium, Tilletia i ccpomeshags 452; Oryzae, 453
460; um, 459; involutum, 459; Ti commutata, 151; leiococcus,
ibebafa, apenas aoa: luzoniense, I51 issouriensis, 151; —— I51
46: 61; nigri , 458; ostrea, 4593 Tournefortia argentea, am :
ostrea concolor, 459; perlatum, bab Tradescantia virginian
460; pri s, 460; radio-fissum, Trametes aurora, me "Bieri, 468; in-
Pheicolor, 458; re 460; acne. sularis, 468
460;
Stevia oo
Stipa a T, 3690
Strand flora of — veclnnary Archipelago,
,
uissi imum,
154
and c Bs Re ig 259;—II.
marth i 483
ao seg glomerata 138; hawaiiensis,
139
suring variation in Silene noctiflora, A,
Siecewell roseus, 207
Stro: aaplegee — m, oe. as 232
Studies in the genus Gymnosporangium
Re nae a catecnes made in
IQI5 ‘and x IQI6, 2
Studies in the pice Lupinus—II.
Mi saat exclusive of Lupinu
—-[k.
e
s den-
ical relations, origin, | Tri 39
. Ecological Tribulus cis PN “20, 276, 495, 500
Tranzschelia cohaesa, 146
| Trematolobelia macrostachys, 13
8
caragana, 456; auricula, 456;
enum vitginicu
talis amer.
Trifolium prate ense, 40; repens, 40
a
Tsuga canaden
A bo peserrsge tg a6: latifolia, 29, 38,
acti myces Elmeri, 468
Tw
wo remarkable Discomycetes, 77
| Ulmus americana, 37
» 42
siflorus, 1;—III. Lupinus Seniors. Un oodi Cees 7
16 be nape canadense, 38
cay ar of some new cases of apogamy in | {Jredo ca 452; Ciberti, 454; destruens,
452; i
S, 93
Seniics on the vegetation of New York
ate—II. The vegetation of a glacial
plunge basin and its relation to tem-
perature, ° 203
Suriana mari sg 270 271
Stenotaphrum, 2
Tacca pinnatifida, 27
Bagh erin rosaries 244, 246
onomy and distribution of Adeno-
stegia 399
AALS ay 38,
TENopyrR, L. A. On ae constancy of
shape in leaves of varying shape,
eadvcali littoralis, 270; piscatoria, 275
Terminalia Ka’ 277
Tetramolopium sp., 495, 496
Thalictrum dasycarpum, 39; polygamum,
380
Tictophics adusta, 457; atro-caerulea,
457; caerulea, 457; concolor, 450;
fimbriata, 457; indigo, 457; lobata,
459; rica, 455; nigricans, 458;
purpurea, ‘455; pavonia, 462; prince
Ecsta sa 457
Galeottiana, 150
airy populnea, 270, 271, 497
Thuja occidentalis, 37
teriae, 454; pana-
; tonkinensis, 453; unilat-
Scrape dow: 143; bo 141;
fusca 142, eria
Urtica dicken, a.
Ustilagineae,
Ustilago a tiene ecibulatt:
carbo wnaasoe 452;
modensis,
leucoderm , 453 '; Panici peat 452;
ached. 453; virens, 4
thesapionaes Oryzae, 4533 lens 453
Vaccinium canadense, 38; pennsylvani-
cum, 3
Vagnera racemosa, 31, 39; stellata, 373;
ifolia
Verbascum Thapsus
telat 2 littoralis, ae polystacha, 270
rpa digitaliformi
Vitroen >; 19
} venosum, 383
ven oria regia, 52
Vi igna, fide, lutea, 270, 272, 276; oahuen-
Vitex trifolia, 272
INDEX 519
Vitis vinifera, 52, 271; vulpina, 38 Wound periderm in certain cacti, 353
Wi ria americana, 276, 497 ear echinatum, 277
Mloarsiacintits Claytoni, 39 Ximemia americana, 271
poe wax, P. The evolution of Zanicheli palustris, 368
mg Zea ner: 09, 329; Mays, 56, 312
a Uva-ursi, 274, 496 isi age
boiaganttes. ag ek 270, 271
Live Music Archive
Librivox Free Audio
Metropolitan Museum
Cleveland Museum of Art
Internet Arcade
Console Living Room
Books to Borrow
Open Library
TV News
Understanding 9/11